Midterm 1 (lectures 1-13) Flashcards
How diverse are the vertebrate species? (Facts to know)
- extant species
- discovered species
- extinct species
- living species percentage of total
Over 63,000 extant species.
100-200 species discovered every year.
More than five million extinct species.
Living species only 1 percent of total.
Extant vs extinct
extant: still living
extinct: no longer exist
Name the two major groups.
- non-amniotes
2. amniotes
Which are aquatic and terrestrial vertebrates, the non-amniotes or amniotes?
non-amniotes=largely aquatic
amniotes=largely terrestrial
What are the non-amniotes and give examples.
Embryo enclosed and protected by membranes produced by reproductive tract of female.
ex) jawless fishes (Agnathans), cartilaginous fishes (Chondrichtyes), bony fishes, amphibians (salamanders, frogs, caecilians).
Largely aquatic.
What are the amniotes and give examples.
Additional set of three membranes associated with embryo (fetal membranes). One of these is the amnion.
ex) turtles (Testudinia), Lepidosauria (lizards, snakes, tuatara), Crocodilia, Birds (Aves), Mammals.
Largely terrestrial.
Which three species are part of the “reptiles?”
turtles (Testudinia) Lepidosauria (lizards, snakes, tuatara)
Crocodilia
Why do we use classification?
- useful for organization
- reflects evolutionary relationships
Taxon and Taxa
- named taxonomic unit at any level
- plural=taxa
Binomial nomenclature
- scientific naming of species, standardized by Linnaeus
- Genus+species (always italicized)
How does hierarchy of classification work?
- higher levels of classification are more inclusive (Kingdom)
- lower levels are more exclusive (Species)
- Should reflect degree of relatedness or evolutionary relationship (when they last shared an ancestor, not necessarily how similar they appear)
Which are the levels of classification? List them in order of most inclusive to most exclusive.
- Kingdom (e.g., Animalia) includes many phyla *most inclusive
- Phylum (e.g., Chordata) includes many classes
- Class (e.g., Mammalia) includes many orders
- Order (e.g., Carnivora) includes many families
- Family (e.g., Felidae) includes many genera
- Genus (e.g., Felis) includes many species
- Species (e.g., concolor) *most exclusive
Cladistics
- hierarchical classification of species based on evolutionary ancestry
- recognizes only groups that are monophyletic (one tribe)
Clade
(“branch”) = an evolutionary lineage
Nodes
node= common ancestor of the tree
Phylogenetic systematics
Use of cladistics, a hierarchical classsification of species on evolutionary ancestry/relationships. Excellent way to visualize and summarize interrelationships among taxa.
Monophyletic
- “one tribe”
- where same ancestor gave rise to all species in that taxon and to no species in any other taxon
How do we identify monophyletic groups?
- On basis of derived (apomorphic) characters
- presence of synapomorphy (shared derived characters) helps identify common ancestry
Apomorphy= derived
character different from ancestral condition (novel trait)
Plesiomorphy
ancestral character
ex) feathers in birds, hair in mammals, jaws in vertebrates
Synapomorphy
shared derived characters
ex) Notochord of Chordates
Symplesiomorphy
- shared ancestral characters
- uninformative
- retention of ancestral characters does not necessarily indicate recent common ancestry
ex) mammals (with hair) more closely related to each other, but absence of hair doesn’t indicate close relationship
Homologous
- characters are the same due to common or shared ancestry, regardless of whether or not structures perform the same function in extant organisms
- ex) pectoral fins, forelegs, wings
- ex) swim bladder of fish=lungs of tetrapods
If a derived character (structure) were to evolve independently in different lineages, will it be useful for determining evolutionary relationships?
No.
ex) endothermy (“warm blooded”) in birds and mammals evolved independently in each lineage
Analogous characters or Homoplasy
- resemblance between species from different evolutionary branches
- similar ecological roles but arose independently
- plas=form
- refer to structures which perform a similar function but are not necessarily homologous
- ex) gills of fish and lungs of tetrapods
What are the various causes of homoplasy?
1) convergent evolution
2) parallel evolution
3) evolutionary reversal
Causes of homoplasy: Convergent evolution
similar characters evolve independently in separate lineages which diverged a very long time ago
ex) bats and birds do not have a common ancestor with wings
Causes of homoplasy: Parallel evolution
species that diverged relatively recently develop similar specializations
ex) elongate hind legs in arid adapted rodents in Africa and North America
Causes of homoplasy: Evolutionary reversal
independent evolution of trait (not indicative of ancestry)
ex) re-evolution of streamlined body & fins of aquatic whales & dolphins whereas sharks retained these traits from the ancestral condition
* Independent evolution of trait (not indicative of ancestry)
Cladistics doesn’t recognize which two types of taxa?
1) Polyphyletic taxa: members are derived from two or more unrelated lineages; also called artificial grouping or homoplasy (plas=form)
2) Paraphyletic taxa: don’t include all species derived from common ancestor
What are some traditional groups that are paraphyletic?
- Dinosauria (dinosaurs) should include birds
- Reptiles should include birds but still useful descriptive terms
- chimps (Pongidae) and humans (Hominidae) are more closely related but don’t like to include ourselves in that relationship
Sister groups
monophyletic lineage most closely related to monophyletic lineage being discussed
ex)Pterosaurs are sister to dinosaurs
Crown groups
taxa at ends of branches (extant species)
Stem groups
- extinct forms that preceded the point where first member of crown group branched off
- May lack all of derived characters of crown group
In the Phylum Chordata, which three subphylums exist? Which two subphylums are the “non-vertebrate Chordates?”
1) Subphylum Vertebrata
2) Subphylum Urochordata (tunicates, sea squirts)
3) Subphylum Cephalochordata (lancelets)
* 2 and 3 are “non-vertebrate chordates”
Which are the two ways in which Chordates can develop?
- Protostomata (first mouth, blastopore=mouth) *paraphyletic
- Deuterostomata (second mouth, blastopore=anus) *monphyletic therefore, deuterostome development synapomorphy indicating shared ancestry
Where do Chordates fit in relation to other animals?
- 30 other animal phyla
-Chordates superficially resemble other active animals (e.g., insects)
-But more closely related to phylum Echinodermata
and phylum Hemichordata
-Chordata, Echinodermata, Hemichordata are the only deuterostomes
Name the five synapomorphies (derived characteristics) that appear during the development of all Chordates and distinguish them as a monophyletic group (whether or not they persist in adulthood).
1) Notochord
2) Dorsal Hollow Nerve Cord
3) Postanal Tail
4) Pharyngeal Gill Slits
5) Endostyle/Thyroid Gland
Synapomorphies of all Chordates: Notochord
-gives Chordata its name
-Semi-rigid rod made of large cells with tough
connective tissue covering
-Incompressible, preventing shortening, but flexible
-Point of muscle attachment
-Replaced by vertebrae in most vertebrates
Synapomorphies of all Chordates: Dorsal Hollow Nerve Cord
- Ectodermal cells grow upward at dorsal midline, forming a hollow tube
- Forms spinal cord
- Ventral, solid and of mesodermal origin in Arthropoda, annelids, etc
- Appears to be induced by presence of notochord
Synapomorphies of all Chordates: Postanal Tail
- Segmented, muscular tail
- Extending beyond gut region
- Present in all chordates during embryonic development
Synapomorphies of all Chordates: Pharyngeal Gill Slits
- Paired openings in walls of anterior pharynx
- Likely evolved as filter feeding mechanism
- Modified for respiration in fish
- Present in tetrapod embryos but modified during development
Synapomorphies of all Chordates: Endostyle / Thyroid Gland
-Endostyle in adult non-vertebrate chordates and
larval lampreys
-Secretes mucus to trap food
-Homologous to thyroid gland of vertebrates
-Groove of ciliates glandular tissue on pharynx floor
-Larval lamprey endostyle metamorphoses into adult thyroid
Subphylum Urochordata: Facts
- tunicates and sea squirts
- Uro=tail, so notochord extends into tail
- Approx. 2000 species
- All marine
- Approx. 100 species with free-swimming adults
Subphylum Urochordata: Adult forms vs Larvae forms
-Most species have sedentary adults
-Adult with large ciliated pharynx for filtering, encased in fibrous ‘tunic’
-Adults only have 2/5 synapomorphies of Chordates: endostyle and pharynx
-Free-swimming larvae with all chordate characteristics
-Transformation of larval tunicate involves loss of
tail, notochord, and dorsal nerve cord
Subphylum Urochordata: Larvacea
- Group of urochordates which never transform
- Reach sexual maturity in larval form
- Paedomorphosis
- Adults retain notochord and tail and dorsal nerve cord
Paedomorphosis
retention of larval characteristics by sexually mature adult
Subphylum Cephalochordata: Facts
- Cephalo=head, notochord extends into head
- 22 species small, superficially fishlike marine animals (in salt water)
- lancelet or amphioxus, Branchiostoma lanceolatum
Subphylum Cephalochordata: Characteristics and derived traits
-Mostly burrowing, sedentary
-Anterior elongation of notochord aids in burrowing
-Pharyngeal gill slits for filter feeding, not gas exchange
-Derived traits:
○ Skin forming vertebrate-like tail fin (caudal fin)
○ And obvious muscle segmentation (myomeres)
Myomeres
blocks of striated muscle fibres separated by sheets of connective tissue
Subphylum Cephalochordata: What new discoveries have been made regarding the relationships between the three subphylums Cephalochordata, Urochordata and Vertebrata?
- Caudal fin and myomeres previously considered to be synapomorphies showing cephalochordates and craniates (vertebrates) to be sister taxa
- But new (molecular) evidence suggests Tunicates and not cephalochordates are the closest living relatives of vertebrates
- Shows urochordates and vertebrates as sister taxa
- Previous assumption that earliest chordates sessile and that cephalochordates and vertebrates evolved from tunicate-like ancestor
- Now suggestions that sessile adult stage in tunicates is derived
- Ancestral form? Larvacea
- Sessile tunicates specialized, not “primitive”
- Evolution sometimes messy
Why is the Subphylum Vertebrata also termed Craniata in older references?
-not all animals included in this subphylum have vertebrae
To be considered in the Subphylum Craniates, what characteristics are key?
1) brain in a cranium
2) paired sensory organs
3) pharyngeal arches respiratory
* excludes vertebrae and dorsal fin
What are the five synapomorphies of Subphylum Vertebrata?
1) brain in a cranium
2) paired sensory organs
3) pharyngeal arches respiratory (no longer for filter feeding)
4) vertebrae (secondarily lost in hagfishes)
5) dorsal fin (secondarily lost in hagfishes?)
* in some lineages, these will be secondarily lost or modified
What are agnathans?
- means ‘no jaws’
- a Superclass of jawless fish in phylum Chordata, subphylum Vertebrata
- a paraphyletic group (including extinct jawless fishes)
What are the two living agnathans?
1) hagfishes: lack vertebral elements altogether and lack of eyes = plesiomorphic traits have been lost secondarily, is a Craniate but not Vertebrate
2) Lamprey: have rudimentary cartilaginous vertebrae
* molecular evidence showing the two are monophyletic and sister taxas
* was once a more diverse group
Vertebrates are larger, more active by finding prey and evading predators. Therefore, they need which of the following to be able to do these things?
- Organ systems (instead of relying on diffusion or ciliary action)
- Higher metabolic rate (greater oxygen intake, circulatory system)
- Muscles and skeleton
- Flexible but protective outer covering
- Some tissues mineralized with unique type of mineral, hydroxyapatite (Ca, P)
- More resistant to acid (e.g., lactic acid) than calcium carbonate in mollusk shells
Cephalization
Concentrated, recognizable head end
Vertebrates display which important physical characteristics?
- Distinct cephalization with paired sensory organs (Nerve cord expanded into brain)
- Inner ears include at least one semicircular canal (Important for sensing orientation) and lateral line to detect movement and vibration)
- Pharyngeal gill slits become respiratory in adults. In non-vertebrate chordates, gas exchange across body surface; pharyngeal arches for filter feeding.
Non-vertebrate Chordate vs. Generalized Primitive Vertebrate: Brain and Head End
Non-vertebrate: no cranium, simple brain (cerebral vesicle), only photoreceptive frontal organ
Vertebrate: cranium around brain, tripartite brain, multicellular sense organs
Non-vertebrate Chordate vs. Generalized Primitive Vertebrate: Pharynx and Respiration
Non-vertebrate: Gas exchange across body surface; gill (pharyngeal) arches for filter feeding. Pharynx not muscularized. Water moved by cilia.
Vertebrate: Gill arches support gills used for respiration. Musculature for active pumping
Non-vertebrate Chordate vs. Generalized Primitive Vertebrate: Feeding and Digestion
Non-vertebrate: filter feeders, food passage by ciliary action, intracellular digestion
Vertebrate: most particulate feeders, larger gut volume, gut muscles to move food (peristalsis), digestion by enzymes in gut lumen
*exception=hagfish: products absorbed by cells lining gut
Non-vertebrate Chordate vs. Generalized Primitive Vertebrate: Circulatory System
Non-vertebrate: no true heart, single-chambered pumping structure, no neural control, open system with large blood sinuses, accessory pumping regions, blood not involved in gas transport, no respiratory pigment
Vertebrate: ventral pumping heart, with sinus venosus, atrium, ventricle, neural control regulates pumping (except hagfishes), closed with extensive capillary system (except hagfishes), red blood cells with hemoglobin, accessory pumping regions=*hagfish only
Non-vertebrate Chordate vs. Generalized Primitive Vertebrate: Osmoregulation
Non-vertebrate: body fluids same ionic composition as seawater, no specialized kidneys
Vertebrates: body fluids more dilute than seawater (*except hagfishes since it is lots of work to maintain osmoregulation), glomerular kidneys
Non-vertebrate Chordate vs. Generalized Primitive Vertebrate: Locomotion
Non-vertebrates: no median fins beside tail fin
Vertebrates: median fins, caudal fin with dermal rays, dorsal fins (*except in hagfishes)
True or False? There are more than 60,000 extant vertebrate species.
*on exam
True
True or False? Extinct vertebrate species likely outnumber extant vertebrate species by about 100:1.
*on exam
True
True or False? Amniotes include the fishes and amphibians.
*on exam
False. These are non-amniotes.
True or False? Biological classifications will change as our understanding of the interrelationships among taxa changes.
*on exam
True
True or False? Only monophyletic taxa are recognized in cladistics.
*on exam
True
What was observed in the early vertebrate fossils discovered in China 520 years ago?
- 3 cm long, fish-shaped
- Notochord, cranium, paired sensory structures, myomeres, gill pouches
- Dorsal fin (derived character lacking in hagfishes)
- No bone or mineralized scales
In the extant vertebrates, name some examples for the following:
A) no jaws, no bone
B) jaws, no bone
C) jaws, bone
A) hagfish and lampreys
B) cartilaginous fish
C) bony fish and tetrapods
Which evolved first, jaws or bones?
Bones
Conodonts (Conodonta): Facts
- First known from microfossils called conodont elements
- Widespread and abundant in marine deposits from Late Cambrian to Late Triassic (approx. 500 mya – 200 mya)
- Mineralized structures, kept well during fossilization (Mineralization is an anapomorphy)
- Once thought to be skeletal parts of marine algae or invertebrates, but made of apatite, similar to dentine
Apatite
- Mineralized calcium compound unique to vertebrates
- Likely toothlike elements of vertebrates
Conodonts (Conodonta): What helped determine that they were vertebrates?
Vertebrate status confirmed with complete fossils showing:
- Notochord
- Cranium
- Myomeres
- No evidence of gill slits (lost secondarily?)
- Pharynx (in filter feeding) adapted to operate teeth
- Also large eyes and fins with rays
- Likely more derived than soft-bodied, extant jawless fishes
True or false? Extinct means primitive.
False. Extinct does NOT mean more primitive.
Ostracoderms: Facts
- “Ostracoderms” (= Shell Skins)
- Bone fragments from around the world (480–500 mya)
- Later complete fossils (400 mya)
- Approx. 10–50 cm long
Ostracoderms: Physical characteristics
- bony armour, not an actual shell
- Still no hinged jaws
- Also more derived (more novel traits) than extant agnathans (hagfishes and lampreys)
- ex) cerebellum, olfactory tract = additional anapomorphies
- physiological capacity to form mineralized tissues in dermis (dermal bone)
Dermal bone
- Not external to skin like a shell
- Little toothlike elements (odontodes) formed in the skin
- Base of acellular bone overlain by layer of epidermis
- Virtually unmodified in placoid scales of sharks
- Aggregations form larger scales, plates, and shield on heads of ostracoderms and early bony fishes for protection
- Human teeth similar and likely homologous
- But origin of this ability a puzzle
- Earliest-known mineralized structures no less complex than that of living vertebrates
- *No (known) transitional stages
Name the advantages for selection of mineralized tissues for organisms.
- Protection: Defence against predators (e.g., eurypterids)
- Sensory: Insulating coat around electroreceptors that enhanced prey detection
- Mineral storage and regulation: Ca and P (relatively rare element, especially in fresh water)
Which evolved first? Freshwater or saltwater organisms?
Saltwater then freshwater
How do Ostracoderms fit into evolution as a group?
- Diverse group
- Paraphyletic group: more derived groups (e.g., osteostracans) more closely related to jawed vertebrates (gnathostomes) than to other ostracoderms (e.g., heterostracans)
Ostracoderm: Heterostraci: Pteraspida
- Pteraspida (“Wing Shield”)
- Pter=wing or fin
- No paired fins or jaws
- Have Hypocercal tail
Clade Myopterygii
Myopterygii = “muscularized fins”
- monophyletic
- Paired lateral fin folds, dorsal and anal fin
Ostracoderm: Osteostraci
- derived Cephalaspid
- Appearance of perichondral bone (at least in skull)
- Calcified cartilage
- Cellular dermal bone
- Pectoral fins with narrow base
- Heterocercal tail
Identify the correct progression through evolution of:
A) Dermal bone
B) Perichondral bone
C) Mineralized teeth
1) mineralized teeth
2) dermal bone
3) perichondral bone
How did extinction affect the early agnathans, gnathostomes, and ostracoderms?
- Early agnathans coexisted with early gnathostomes for approx. 50 mya
- Decline in ostracoderm diversity at end of Early Devonian (416–400 mya) may have been due to lowering of global sea levels
- Extinction of ostracoderms in Late Devonian (approx. 365 mya) occurred at same time as mass extinctions among many marine invertebrates
- Only ones that survived are lampreys and hagfishes
Cyclostomes
- Extant agnathans
- round mouth
Describe what we mean by agnathans are a paraphyletic group, including the extinct jawless fishes.
- They are each other’s closest living relatives, but not necessarily closely related
- Lots of differences between them
- Branched off from each other a long time ago
A shared derived characters or Synapomorphy helps to identify what?
Presence of synapomorphy (shared derived characters) helps identify common ancestry
e.g., jaws show gnathostomes to be monophyletic group
Shared ancestral characters help to identify what?
- They don’t help
- Shared ancestral characters are uninformative
- Retention of jawlessness does not necessarily indicate monophyly
Which are more “primitive”? Lampreys, Ostracoderms or Hagfishes?
- Hagfishes and lampreys more “primitive” than ostracoderms
- Fossil record sparse
- Two fossil genera of each from approx. 360–300 mya
- Longer body today than in the past for Lampreys
How many mass extinctions have lampreys and hagfishes survived?
-Hagfishes and lampreys have survived through 4 mass extinctions
Hagfishes (Class Myxini; Order Myxiniformes): Facts
- Myx=slime
- Up to 75 species
- all marine, deep sea (why eyes have been lost)
- Also called “slime eels”
- Population depleted by fisheries use to demand for eel wallets
Hagfishes (Class Myxini; Order Myxiniformes): How are they in many respects “primitive?”
- Single semi-circular canal per side
- Rudimentary eyes (but secondarily lost) due to living in deep sea
- No vertebrae (but may have been secondarily lost)
- Body fluid isosmotic with sea water (does not osmoregulate, keeps salt and water concentrations the same as the sea water they live in)
- No heart innervation
- Open circulatory system and accessory hearts
Hagfishes (Class Myxini; Order Myxiniformes): Do their shared derived characters represent the ancestral vertebrate condition or are these derived specializations?
Today, studies are showing that their shared derived characters are actually specializations and not the case of ancestral vertebrate conditions.
Hagfishes (Class Myxini; Order Myxiniformes): Feeding
- Scavengers (earthworms of the deep, feed on dead fish in the sea)
- Greatly reduced eyes (no lens)
- Good sense of smell and touch
- Large folded tongue with ‘horny’ tooth plates (keratin=same structure as our fingernails are made)
- Single nostril for smell
- Able to tear off pieces of prey using knotted body for leverage (using tongue)
- Food engulfed in mucous membrane for digestion
Hagfishes (Class Myxini; Order Myxiniformes): Gills
- 1 to 15 external gill openings to let water out
- External opening displaced posteriorly
- Pouched gills with water intake through nostril
- Also capable of anaerobic and cutaneous respiration (while feeding)
Hagfishes (Class Myxini; Order Myxiniformes): Slime glands
- Also known as “slime eels”
- Mucus+protein threads
- Function of the slime? Probably for protection from predators and also protection for the skin
- Applications of the slime? Use of protein threads for surgical thread (doesn’t initiate immune response), used in cooking
- 70-200 pairs of slime pores
Hagfishes (Class Myxini; Order Myxiniformes): Reproduction
- Little known about reproduction
- Single gonad
- Likely external fertilization
- No obvious seasonality, so year round
- But highly female-biased sex ratios
- Some species hermaphroditic
- Produce few large eggs which appear to hatch directly into fully formed hagfishes, rather than larvae = Direct Development
Lampreys (Class Petromyzontida; Order Petromyzontiformes): Facts
- petro=stone, myzon=sucker
- Approx. 40 species in 10 genera
- Temperate regions of N and S hemispheres
- 18 species are parasitic on bony fishes
Lampreys (Class Petromyzontida; Order Petromyzontiformes): Characteristics
- Attach with large oral disk
- Horny teeth (keratin) on disk and tongue rasp hole in side of fish
- Oral gland secretes anticoagulants
- Blood rich and easily digested
- Generally do not kill host
Lampreys (Class Petromyzontida; Order Petromyzontiformes): Which characteristics of the lamprey are currently being studied for medical purposes?
- Spinal cord regeneration (only vertebrate who can do this)
- Iron loading
Arrange the following characters in order of their appearance in the chordates:
- dorsal fin
- jaws
- mineralized tissue
- notochord
- pectoral fins
- pharyngeal gill slits for respiration not filter feeding
1) The first: Notochord
2) Pharyngeal gill slits for respiration, not filter feeding
3) Dorsal fin
4) Mineralized tissue
5) Pectoral fins
6) Jaws
How do lampreys appear more “advanced” than hagfishes?
-have cartilaginous vertebral structures homologous with neural arches in gnathostomes
Which group were lampreys previously thought to be a sister taxon to? But recent evidence shows that they are actually a monophyletic group with which species?
- Were generally considered sister taxon to gnathostomes
- But, as mentioned, recent evidence that hagfishes with “long overlooked vertebral elements” and that hagfishes and lampreys do form monophyletic group
How are lamprey’s characteristics more derived than hagfishes?
- Two semicircular canals per side
- Heart innervated by parasympathetic nervous system (PNS)
- Well-developed kidneys and ability to osmoregulate
- One of the first vertebrate groups to invade fresh water
- Anadromous: species that reproduce in both fresh and salt water
- Well-developed eyes (in adults)
- Well-developed pineal gland (Important in seasonality for reproduction)
- Always seven pairs of gills
- Tidal ventilation (in adults, inefficient water comes out one opening and coming out through same opening, effective for eating and breathing separately) rather than flow-through (in larvae, other fish, water in mouth and out the gills)
- Velum prevents water from flowing out of respiratory tube into mouth
- Single nasal opening on top of head
Anadromous
Anadromous: species that reproduce in both fresh and salt water
What are lampreys called in German and why?
In German they are called Neunaugen=nine eyes (nine hole on one side)