9, 11, 13-14: Chordata, Vertebrata, Tetrapoda, Amphibia Flashcards

1
Q

Name the main groups that are within Deuterostomia, but NOT Chordata

A

Ambulacrania - a group containing Echinodermata (e.g., starfish) and Hemichordata (e.g., Acorn Worms!)

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2
Q

Name the main groups that are within Chordata, but NOT Vertebrata

A

Cephalochordata (30 sp.) e.g., Amphioxus and Tunicata (formerly Urochordata, 2000 sp.)

Note: Vertebrata = 70,000 sp. (mostly fishes)

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3
Q

Describe the common anatomical features of chordates

A

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

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4
Q

What is Amphioxus?

A

A small, filter-feeding Cephalochordate with simple morphology - often interpreted as the Archetypal Chordate

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5
Q

Describe the Tunicates

A

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

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6
Q

What are the defining features of VERTEBRATES?

A
  1. Developed brain
  2. Paired sensory structures
  3. Branchial arches
  4. (MOST) some sort of vertebral column
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7
Q

What has changed fairly recently about our understanding of Chordate phylogeny?

A

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]

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8
Q

What are some key anatomical differences between the general anatomy of plesiomorphic Chordates and Vertebrates?

A

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.)

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9
Q

What is the interesting feature about Vertebrate Genomes?

A

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

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10
Q

Describe the Jawless Vertebrates in terms of morphology and lifestyle

A

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”

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11
Q

Describe the jawless vertebrates in terms of anatomy

A

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)

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12
Q

Describe the jawless vertebrates in terms of Development

A

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

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13
Q

What are the key differences between Cyclostomata and GNATHOSTOMATA

A

GNATHOSTOMATA (Jawed vertebrates - Cartilaginous Fishes, Boney Fishes, Lobe Finned Fishes, Tetrapods) have:

Paired Nostrils
Hinged Jaws
Vertebrae
Bone
Adaptive Immune System

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14
Q

Describe the debate/discussion surrounding the phylogeny of the jawless vertebrates

A

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!]

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15
Q

Describe the rate of evolution seen over the course of vertebrate history

A

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

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16
Q

What are the oldest (potentially) Chordate fossils?

A
  • Pikaia from Burgess Shale (Cambrian)
  • Metaspringgia (possible Cambrian Vertebrate) has muscle bands, paired eyes and gill arches
  • Cambrian Chordates of China - Haikouichtyhes, Yunnanozoan, Cathamyrus
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17
Q

Why is chordate phylogeny so hard to resolve for groups before the Gnathostomata?

A

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

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18
Q

What is taphonomy?

A

The study of the process of decay and preservation, to help interpret fossils more accurately and see which characters decay fast or slow

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19
Q

What is meant by the phrase “stem-ward slippage”?

A

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

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20
Q

Define the difference between Agnatha and Cyclostoma

A

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

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21
Q

Describe the main sub-groups of Cyclostoma and Gnathostoma

A

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)

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22
Q

Describe some of the key features of Gnathostomata

A
  • 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)
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23
Q

Describe the Ostracoderms and name some of their groups

A

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

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24
Q

Describe the Conodonts

A

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

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25
Name 4 Ostracoderm Groups and describe their features
1. Sacamambapsis - armoured head and tail; two nostrils; widespread in Ordovician 2. Heterostracans - diverse head shield shapes, composed of multiple plates; single gill opening, no paired fins 3. Galeaspids - heavily armoured headshield; unique sensory line system (2 sensory lines going towards 1 nostril - possible step towards Gnathostome trait) 4. Osteostracans - BONY SHELLS (but still no jaws); complex paired fins and advanced bone; advanced nervous system and sense organs
26
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
27
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
28
Name the two main groups of Chondrichthyes
Elasmobranches (Sharks and Rays which grow and shed teeth) Holocephalans (Rat Fishes with tooth plate)
29
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
30
What are "Osteichthyes"
ALL BONY VERTEBRATES (so Actinopterygii and Sarcopterygii)
31
Name the three groups of Actinopterygii
Chondrostei (Sturgeons, Paddlefish and Reedfish) Holostei (Bowfin and Gars) TELEOSTOMI (Hyperdiverse, Classical "Fishes")
32
Describe the key anatomical features of Actinopterygii
- Bony Endoskeleton, including Lepidotrichia (Fin Rays) - Swim Bladder to control buoyancy - Operculum - Bony scales
33
Name the main groups of Sarcopterygii
The Lobe-Finned Fish (Sarcopterygii) include: - Actinistia (e.g., Coelocanth) - Dipnoi (e.g., Lungfish) - TETRAPODA (ofc)
34
Describe the key features of non-tetrapod Sarcopterygii
Specialised fleshy lobed fins Stylopod (homologus to humerus) in a single connection between fin and supportive girdle (monobasal articulation)
35
What major shift in our understanding of bone evolution (may have) occurred in 2020?
Evidence of endochondral bone in Placoderms - meaning it first evolved far earlier than we thought, and was simply lost in Chondrichthyes, rather than being a synapomorphy of Osteichthyes
36
State some examples of the different ways one could possibly define the complexity of a taxonomic group
Genetic complexity (e.g., WGD events) Anatomical complexity Taxonomic diversity Taxonomic longevity
37
Describe some examples of the fish mating strategies discussed in the lecture
Sexual Dimorphism - Male and Female individuals have morphological differences, e.g. females usually larger [SEEN IN SALMON, PARROT FISH, MANY OTHERS] Hermaphroditism - Either Synchronous or Transitional, only seen in 8 orders [INCLUDING CLOWNFISH] Parthogenesis - Virgin Bonnethead sharks can give birth (rare but may be important for species that live far apart?) Sexual Parasitism (e.g., Angler Fish)
38
State and define some of the Fish Rearing Strategies discussed in the lecture
Oviparity - internal fertilisation, external egg development (e.g., majority of Actinopterygii) Ovuliparity - External fertilisation Viviparity - Internal development of nurtured embryos (e.g., White Surf Fish, Sand Tiger Shark) Ovoviviparity - Internal development, but using resources from egg only
39
What is meant by "Euryhaline" Fish?
Those that are exposed to a wide range of salinity due to migration
40
What is the difference between Iteroparity and Semelparity?
Iteroparity = continuous breeding Semelparity = single mass breeding event (e.g., Pacific Salmon)
41
How does each fish in a school behave, leading to overall complex swarming behaviour?
Rules: - Same direction as neighbour - Stable distance from neighbour - Avoid collisions
42
What are the Teleostei and what is believed to have contributed to their hyperdiversity?
An extremely diverse group of fishes within the Actinopterygii (essentially all bony fish excluding Chondrostei such as Sturgeons and Holostei such as Bowfins) Whole Genome Duplication event in their stem lineage
43
What are two key features of Teleost anatomy?
Mobile Premaxilla for protrusion feeding (remember video of expanding jaw and sucking in prey) - protrusion has gradually increased since the Late Cretaceous Homocercal Tail - spinal column ends at caudal peduncle, allowing fins beyond to take a wide range of forms
44
What are the Acanthomorphs, and name a couple of examples of both Acanthomorphs and Non-Acanthomorphs
Acanthomorpha are an extremely diverse group within the Teleosts, making up a third of extant vertebrate species (over 14000) Acanthomorpha include Tuna, Mackerel, Sunfish, Anglerfish, Seahorses, Clownfish, etc. etc. Non-Acanthomorpha include Eels, Herrings, Carp and Salmon
45
Describe the two mechanisms used by fish for Buoyancy
Swim Bladder - Oesteichthyes control their buoyancy via gaseous exchange in and out of the swim bladder Oily Liver - Chondrichthyes have a large oily liver that provides buoyancy (oil is more buoyant than air), BUT less control, so must control depth using orientation
46
Name the four broad swimming morphologies (and an example of each), from most head movement to least What are the advantages/disadvantages?
Anguilliform (Eels) -> Subcarangiform (e.g., Salmon) -> Carangiform (e.g., Mackerel) -> Thunniform (e.g., Tuna) Anguilliform allows the most rapid acceleration, while Thunniform allows the most efficient cruising ALSO NOTE: Ostraciiform allows highest manoeuvrability
47
Name some niches that have been filled almost identically by both extinct and extant groups
Rapid accelerators, High Manoeuvrability, Sustained Swimming [Examples for most of these among Palaeozoic Chondrostei, Mesozoic Actinops AND Modern Teleosts] Also Mega-Planktivores (Giant Filter Feeders) currently filled by Chondricthyes, but has previously been filled by Actinops (in Mesozoic), Placoderms (in Palaeozoic) and Anomalocaridids (in Cambrian)
48
What is meant by "Morphometrics"?
It is a way of quantifying morphological disparity by plotting "landmarks" to give geometry to fish fossils
49
When did the Acanthomorpha radiate most rapidly?
After the Cretaceous mass extinction event - they evolved into many niches that were previously occupied before the extinction
50
Which group within Osteichthyes did Tetrapods arise from, and in what period did they make the transition to land?
From Sarcopterygians (Lobe-finned fish) and they are thought to have made the transition in the Devonian (around 390 mya)
51
Why (fundamentally) did tetrapods make the transition to land, and what are some key evolutionary events that made this possible?
There was available space and a niche to occupy - they needed terrestrial locomotion, air breathing, structural support and reproductive strategies
52
Describe the main anatomical evidence for the evolution of amphibians from lobe-finned fish
1. Arrangement and development of bones in the skull 2. The amphibian limb appears to be derived from fin structure 3. Labyrinthodont tooth structure (ridges on outside and folded structure) 4. Lobe-finned fish have an opening to the nostrils inside the mouth - this feature is found in ALL LAND VERTEBRATES
53
Name some of the "key player" lobe finned fish and describe the features that make them notable in piecing together the evolution of amphibians
Glyptolepis (385 mya) - distinct loose scales, perhaps early progenitor Eusthenopteron (385 mya) - internal nostrils and distinct fin endoskeleton, first evidence of body support in fins Panderichthys (385 mya) - lacks anal and dorsal fins, tail and shoulder more closely resemble tetrapod features, but still fish-like front fins Acanthostega (365 mya) - early tetrapod, longer snout with fewer bones, lacks opercular bones, interlocking vertebrae, larger pelvis, separation of skull from shoulder Icthyostega, Balanerpeton = Fully tetrapods
54
What are the major general differences in characteristics between fish and amphibians?
Most relate to locomotion (moving in water vs on land requires skeletal changes, especially limbs, shoulder, vertebrae and pelvis) Others relate to skull structure (breathing and feeding) - general trend is that tetrapods have FEWER skull bones (consolidation) and lost the opercular bones, which are completely gone in Tiktaalik and Acathostega
55
Describe the specific skeletal changes between Fish and Amphibians (and discuss the reasons why)
Pelvis and Shoulder Girdle: Attached to Vertebral Column in Amph, but NOT in fish Brain Case and Notochord: Notochord replaced by bone in modern adult tetrapods, and no longer runs into brain case Gill Arches: Reduced in number and size Bone that supports gill arches was redeployed in formation of tongue support + musculature Hip is attached to vertebrae in tetrapods, but NOT in fish Also, amphibians GAINED A NECK which fish lack All generally because neck flexibility important for feeding and breathing on land
56
Describe specifically the change in shoulder structure between fish and amphibians and the importance of this
In fish, the shoulder (pectoral) girdle is attached to the skull (for hydrodynamics) In tetrapods, the shoulder girdle is attached firmly to the backbone, and is separated from the skull (it consists of an enlarged scapulocoracoid and clavicle) This allows force transduction and more flexible head movement when walking on land
57
What factors in the Late Silurian and Devonian may have contributed to the eventual appearance of lungs and terrestrial life?
Fluctuations in rainfall, drying of bodies of fresh water, water temperature increases, O2 levels decrease -> All of this makes gills less useful as O2 falls, so animals with the strongest "limbs" are selected for
58
Describe the problems with evolving air breathing from gills
Breathing in air via external gills would result in massive water loss through evaporation from moist respiratory surfaces -> therefore, need INVAGINATED LUNGS and FOLDED vascularised tissue bathed in fluid HOWEVER, all lungs must use a tidal flow (bidirectional) system, which is not as efficient as unidirectional through-flow due to dead space -> therefore, needs an EFFICIENT PUMP
59
Describe some of the novel adaptations that helped tetrapods to overcome the problems with air breathing
More efficient pumps: - Amphibians use a MODIFIED form of the BUCCAL PUMP (the system used by some air-breathing fishes, in which the mouth makes the pressure changes necessary to move water), in which the AXIAL muscles are used for expiration - Amniotes use an ASPIRATION PUMP, in which the axial muscles are used for both inspiration and expiration Within the amniotes, various specialisations include uncinate processes, pelvic aspiration, hepatic pistons and gular pumping -> possible BR ;)
60
Explain the significance of the fin structure seen in Eusthenopteron
Its forefins contain bones that appear to be homologous with the humerus, ulna and radius Its pelvic fins contain bones that appear to be homologous with the femur, tibia and fibula
61
Describe the changes in the backbone between fish and amphibians
The vertebrae in amphibians are INTERLOCKING (go from just neural arch to NEURAL SPINE + ARCH), fuse with Centrum Elements(?) This makes the backbone less flexible, but stronger, allowing the organism to support its own weight on land
62
Name some of the key traits in the complex web of interconnected characteristics that had to evolve together in the transition to land
Grasping Feeding, Skull, Jaw, Olfaction Substrate Locomotion, Robust Limbs, Shoulder Girdle, Flexible Neck, Intervertebral Articulations, Ribs, Pulmonary Ventilation Many more, but these are some of the key connections
63
When were the most dominant land animals amphibians, and when did they split from the group that would become the Amniotes?
Dominant in the late Carboniferous and early Permian periods (around 310-250mya?) Amniote branch split off around 300 mya
64
Describe the key ecological features of Amphibians
They lay eggs in water, and newborn amphibians live underwater, before metamorphosing into adult forms, which often live on land. Their moist skin functions in gas exchange
65
What were the advantages of moving onto land for the first terrestrial vertebrates?
It opens up new niches and terrestrial prey options It allows for more diverse locomotive ability It reduces competition or predation from other aquatic species It facilitates selection of an increase in body temperature, which increases rates of digestion, development and other physiological processes
66
What is meant by the "amphibian gap" and what are some possible reasons for it?
The distinct lack of fossils for a clear ancestor of any of the modern amphibian groups, prior to the early Jurassic - we only have a good record of the archaic amphibians and the stereospondyls Taphonomy can help explain this: Amphibians have a reduced skeleton, lots of soft, delicate body parts, and many are burrowing species in a fertile environment -> all of these factors reduce their chances of being preserved Fortunately, shared features at least provide confidence in monophyly of the three crown groups, although very little fossil evidence of their ancestors until Cretaceous (one or two Salamander ancestors from Jurassic)
67
What were the stereospondyli?
A group of extinct amphibians that appeared in the Late Permian - two families survived into the Triassic They had a distinct anatomy (simple backbones and weak vertebral structure, suggesting an aquatic lifestyle) -> none are a likely ancestor for any modern amphibians
68
What are the three main theories for the origin of amphibians, and what two features provide reasonably strong evidence for one of these hypotheses?
1. Lepospondyl hypothesis (Lepospondyli are sister to all Lissamphibia) 2. Temnospondyl hypothesis (all modern amphibians are closely related to dissorophoid temnospondyls) 3. Different groups of Lissamphibians evolved from different groups of early tetrapods or lobe-finned fish A phylogenetic analysis from Sigurdsen and Greene (2011) supports the second hypothesis, as does the shared rare feature of Pedicellate Teeth among all modern amphibians, and the Lissamphibian Elbow Joint
69
Name the three groups of modern amphibians, and the period in which they likely underwent radiation
Anurans (Frogs), Caecilians and Salamanders (Early-Mid) Carboniferous
70
Describe some of the key anatomical features of Amphibians
Circulatory System: They have a 2-chambered heart (unlike the partially- or fully separated hearts seen in reptiles and mammals) Limbs: 4 limbs and 4 digits (except Caecilians) - limbs can support body weight unlike lobe-finned fish, but humerus much less developed than in reptiles Skull: Consolidation of skull parts compared to fish, reduction of opercular bone and loss of posterior skull bones
71
Describe the ecology of Salamanders
- Many show a classic amphibious lifestyle (aquatic larvae and terrestrial adults, others more neotonic) - Carnivorous - worms, arthropods and molluscs - Paedomorphosis (e.g., Axolotls) - Capture prey using prehension of sticky tongue (rapid projection and retraction)
72
Describe the anatomy of Salamanders and their different groups
- Limbs at right angle to gait (SPRAWLING) - Anatomy varies with ecology - aquatic and burrowing groups have reduced limbs, while terrestrial and metamorphosing genera have short bodies and stout limbs
73
Describe the features of Caecilians (and briefly describe their fossil record)
- Unique among modern amphibians in their (secondary) lack of girdles and limbs -> have a "kink" where pelvis used to be - Elongated body with small scales, good burrowers using INTERNAL CONCERTINA locomotion - Eat worms and small invertebrates - Young have special dentition to eat layers of mother's modified skin (possible intermediate step in evolution of viviparity) Fossil record ranges from Lower Jurassic to Upper Paleocene (though only 3 are well-preserved enough to have been named)
74
Describe the fossil record of Anurans (Frogs)
Frog fossil record dates back to Jurassic (Prosalirus bitis), but most (e.g., Gerobetrachus) are quite morphologically similar to modern Anurans -> this is useful for showing how long ago the modern form appeared, but less useful for determining what transitional forms were like Prosalirus bitis has uncinate processes on ribs and was capable of jumping
75
Describe the characteristics of modern Anurans
- REPRODUCTION: Aquatic reproduction, tailed larval form, with gills - Adults lack tails and have lungs, and are specialised for jumping - Extreme skeletal changes, including lots of fusion of bones (e.g., post-sacral vertebrae into urostyle, tibiofibula, radioulna) - Evidence of gliding ability (likely did not evolve flight because they are ectothermic, and that niche was already filled by pterosaurs and then birds)