9, 11, 13-14: Chordata, Vertebrata, Tetrapoda, Amphibia Flashcards
Name the main groups that are within Deuterostomia, but NOT Chordata
Ambulacrania - a group containing Echinodermata (e.g., starfish) and Hemichordata (e.g., Acorn Worms!)
Name the main groups that are within Chordata, but NOT Vertebrata
Cephalochordata (30 sp.) e.g., Amphioxus and Tunicata (formerly Urochordata, 2000 sp.)
Note: Vertebrata = 70,000 sp. (mostly fishes)
Describe the common anatomical features of chordates
Notochord (of course) - a stiff rod along the dorsal axis of the body, mechanically important
Pharyngeal Gill Slits
Endostyle
V-shaped Myomeres
Dorsal Nerve Cord (i.e. more dorsal than the gut)
Post-anal tail
What is Amphioxus?
A small, filter-feeding Cephalochordate with simple morphology - often interpreted as the Archetypal Chordate
Describe the Tunicates
All juveniles (and adult larvaceans) have notochord, nerve cord, brain, muscular post-anal tail
Adult forms can be larvaceans, salps or Sea Squirts
Sea Squirts undergo metamorphosis from a mobile stage to a sessile stage - pedomorphosis
What are the defining features of VERTEBRATES?
- Developed brain
- Paired sensory structures
- Branchial arches
- (MOST) some sort of vertebral column
What has changed fairly recently about our understanding of Chordate phylogeny?
Previously, it was assumed that Tunicates were sister taxon to other chordates, and that Cephalochordates were the closest relatives of Vertebrata (based on Neuroanatomy, Metameric Segmentation and Molecular Data)
Now, taking into account LBA of the Tunicates, it is thought that Cephalochordates are sister taxon and Tunicates are closest relatives of the Vertebrata [Grouped together as Olfactores]
What are some key anatomical differences between the general anatomy of plesiomorphic Chordates and Vertebrates?
Vertebrates have W-SHAPED Myomeres (not V)
Vertebrates have a muscular pharynx with fewer gill slits
Vertebrates have a brain and paired sense organs
Vertebrates have a NEURAL CREST - a migratory population of multipotent cells that form around the developing Neural Tube, and give rise to many important cell types, allowing the formation of important vertebrate characters (e.g., teeth, heart, nerves, etc.)
What is the interesting feature about Vertebrate Genomes?
There appears to have been TWO Whole Genome Duplication events in early vertebrate evolution - one between amphioxus and the jawless vertebrate ancestor, and another one since
Describe the Jawless Vertebrates in terms of morphology and lifestyle
Main 2 groups - Lampreys and Hagfish
Lampreys: Eel-like morphology with an oral sucker and keratinous teeth, no bones or fins; parasitic blood-sucking lifestyle
Hagfish: Soft, blind, worm-like morphology, no bones or fins; slime as defense mechanisms and “tie themselves in knots”
Describe the jawless vertebrates in terms of anatomy
Both have a CAARTILAGINOUS SKULL (very different from out own) and just one nostril
Lampreys: Single median nostril, Cranial Cartilages, Branchial Cartilages, Arcualia (which MAY be homologous with our backbone despite not being bone)
Hagfish: Single terminal nostril, Slime Glands, MANY gill openings, Cranial Cartilages, NO SORT OF BACKBONE (unusual for vertebrates)
Describe the jawless vertebrates in terms of Development
Lampreys undergo Metamorphosis: juvenile stage (ammocoete) lacks eyes or oral sucker (similar to Amphioxus)
Hagfish: development very difficult to study but eventually neural crest was confirmed, as well as similarity to lamprey cranio-facial development
What are the key differences between Cyclostomata and GNATHOSTOMATA
GNATHOSTOMATA (Jawed vertebrates - Cartilaginous Fishes, Boney Fishes, Lobe Finned Fishes, Tetrapods) have:
Paired Nostrils
Hinged Jaws
Vertebrae
Bone
Adaptive Immune System
Describe the debate/discussion surrounding the phylogeny of the jawless vertebrates
The Traditional View: Jawless Vertebrates as their own clade, Cyclostomata (Based on feeding apparatus and nasohypophysial opening)
Challenger Hypothesis: Jawless Vertebrate Paraphyly - Lampreys may be more closely related to Gnathostomata based on Vertebrae, Dorsal Fins, Camera Eyes, Closed Circulation
Conclusion based on MicroRNA Genetic Data - Traditional View was actually correct. Cyclostomata are monophyletic [Also, character losses in Hagfish were confirmed, because a stem hagfish fossil was found to have eyes with melasomes and melanin!]
Describe the rate of evolution seen over the course of vertebrate history
Extremely rapid early vertebrate evolution with lots of innovation, and increasing complexity of development and anatomy (based on phylogenetic data, but limited fossils); slower evolution later on
What are the oldest (potentially) Chordate fossils?
- Pikaia from Burgess Shale (Cambrian)
- Metaspringgia (possible Cambrian Vertebrate) has muscle bands, paired eyes and gill arches
- Cambrian Chordates of China - Haikouichtyhes, Yunnanozoan, Cathamyrus
Why is chordate phylogeny so hard to resolve for groups before the Gnathostomata?
Gnathostomata is the only clade with Skeletal Hard Tissues - before this, animals were entirely soft bodied -> significant decay
The “Cambrian Chordates of China” (H, Y, C) and Haikouella could be seen as stem deuterostomes, chordates or even vertebrates
What is taphonomy?
The study of the process of decay and preservation, to help interpret fossils more accurately and see which characters decay fast or slow
What is meant by the phrase “stem-ward slippage”?
Organisms tend to lose synapomorphies as they decay, and so appear more primitive, or stem-like; therefore, they risk being classified as more “stemward” than they actually were
E.g., Cathamyrus only has decay resistant characters, so may be more advanced than it appears -> Potentially Any Chordate
Haikouichthys has SOME decay-prone vertebrate characters, but NO SKULL, so it probably really did lack a skull and was a true stem-vertebrate
Define the difference between Agnatha and Cyclostoma
Cyclostoma refers to the EXTANT jawless fishes (Lampreys and Hagfish)
Agnatha refers to ALL jawless vertebrates, including extinct groups such as the Anaspids, Conodonts and Ostracoderms
Describe the main sub-groups of Cyclostoma and Gnathostoma
Cyclostoma:
- Lampreys (Blood-sucking “vampire-eel-like fish” with keratinous teeth and oral sucker)
- Hagfish (Slimy, blind “eel-like fish” with rasping keratinous teeth
Gnathostoma:
- Chondrichthyes (Cartilaginous fishes, e.g. sharks and rays)
- Actinopterygii (Bony fishes - most diverse group)
- Sarcopterygii (lobe-finned fish and tetrapods)
Describe some of the key features of Gnathostomata
- Opposing Jaws with True Teeth
- Most have paired appendages (e.g., pelvic and pectoral fins, arms, legs, wings, etc)
- Fossil record shows stepwise acquisition of Gnathostome characters among stem-Gnathostomes (e.g., Placoderms)
Describe the Ostracoderms and name some of their groups
The Ostracoderms are an extinct, paraphyletic group of jawless fossil fishes, ranging from the Ordovician to the Devonian, including the Anaspids, Thelodonts, Galeaspids, Osteostracans, etc.
They may show the first evidence of Biomineralisation:
Some Anaspids have a mineralised dermoskeleton
Describe the Conodonts
Early jawless fish, more stemward than the Ostracoderms
Appear to have “mineralised teeth” that may have been homologous with ours - now thought to be convergent
Name 4 Ostracoderm Groups and describe their features
- Sacamambapsis - armoured head and tail; two nostrils; widespread in Ordovician
- Heterostracans - diverse head shield shapes, composed of multiple plates; single gill opening, no paired fins
- Galeaspids - heavily armoured headshield; unique sensory line system (2 sensory lines going towards 1 nostril - possible step towards Gnathostome trait)
- Osteostracans - BONY SHELLS (but still no jaws); complex paired fins and advanced bone; advanced nervous system and sense organs
Describe the Placoderms
The Placoderms were Jawed Stem Gnathostomes - i.e. more closely related to extant Gnathostomes than Ostracoderms
They were a diverse group, with simple jaws, pelvic fins and possibly teeth
Include the first “mega-predators” - e.g., Dunkleostus, whose bladed jaws had 5000N biting strength, stronger than a Great White
Summarise the stepwise acquisition of Gnathostome Characters along the Gnathostome stem
BONE TRANSITION:
- Origin of unmineralised skeleton in Cyclostomes
- Mineralised dermal skeleton in Anaspids
- Armoured Head + Tail in Sacabambapsis
- Multiple Plates in Heterostracans
- Heavily armoured head shield in Galeaspids
- Axial skeleton and Appendicular Skeleton in Crown-Gnathostomes
TEETH TRANSITION:
- Inside-out hypothesis -> Teeth derived from dermoskeleton migrating into the oral cavity
- Stem-Gnathostomes have dermoskeletal bone, and some pharyngeal denticles
- Helicoprion (a Chondrichthyan) had a buzzsaw-like mouth
PAIRED APPENDAGES:
- Strange paired anal fin in Devonian Euphanerops (FIRST PAIRED APPENDAGES??)
- Osteostracans have paired pectoral fins with skeleton
BRAIN EVOLUTION:
- Development of cerebellum
- Transitional brains seen in Placoderms
Name the two main groups of Chondrichthyes
Elasmobranches (Sharks and Rays which grow and shed teeth)
Holocephalans (Rat Fishes with tooth plate)
Describe the key features of Chondrichthyes
The Cartilaginous Fish have:
- Cartilaginous Skeletons with placoid scales composed of dentine and enamel (HOMOLOGOUS WITH TEETH)
- Bony Teeth (and mineralised vertebral centra)
- Gill openings
- Closed Blood System
- More Complex Brain than Placoderms
- Sensory systems: Lateral Line System for water pressure changes, and Ampullae of Lorenzini for Electroreception
- Sharks have claspers
What are “Osteichthyes”
ALL BONY VERTEBRATES (so Actinopterygii and Sarcopterygii)