Chordates and Vertebrates: Evolution

BK
Books for reference:
• K. V. Kardong: Vertebrate: Comparative, Anatomy, Function, Evolution, 4th Edition
• R. L. Kotpal: Modern Text Book of Zoology: Vertebrates
• E.L. Jordan, P.S, Verma: Vertebrate Zoology, S. Chand Publication

Note: All the images and diagrams used in this ppt are meant for educational purpose only. Copyright of these images/diagrams lies with the original authors and/or publishers of respective reference books/sources
Relative abundance of species within animal phyla
Chordate Phylogeny
Two distinct and independent lines of evolution in Bilateria
Generalized chordates
characteristics

1. Pharyngeal slits
2. Notochord
3. Dorsal hollow nerve cord
5. Edostyle
• A glandular groove in the floor of the pharynx that arises embryologically from the floor of the
pharynx.
• Involved in filter feeding.
• Both endostyle and thyroid are involved in iodine metabolism,
• suggesting a homology between the two,
• ‘the endostyle, being the phylogenetic predecessor of the thyroid’
• Evidence: jawless fish (lampreys) have a true endostyle when they are young larvae, that
becomes a true thyroid when they become adults.
• Thus, all chordates have endostyles (urochordates, cephalochordates, larval lamprey) or thyroids
(adult lamprey, all other vertebrates).
4. Postanal Tail

• chordates possess a postanal tail that represents a posterior elongation of the body extending
beyond the anus.

• The tail is primarily an extension of the chordate locomotor apparatus, the segmental
musculature and notochord.
Outline classification of phylum Chordata
Class Mammalia
Subclass I. Prototheria (Gr., protos, first + therios, beast)
• Order 1. Monotremata. (Gr., monos, single + trema, opening).
• Cloacal opening present.
• Confined to Australian region.
• E.g. Platypus or duckbill (Ornithorhynchus) Spiny anteater (Tachyglossus = Echidna).
• Subclass II. Theria (Gr., ther, animal)
• Modern or typical viviparous mammals that give birth to living young.
• Subdivided into 2 living infraclasses :
• Infraclass 1. Metatheria (Gr., meta, between or after)
• Pouched. viviparous mammals
• without or with a rudimentary yolksac placenta.
• Confined mostly to Australian region.
• Order 2. Marsupialia.
• E.g. Opossum (Didelphis), kangaroo (Macropus), koala (Phascolarctos).
• Infraclass 2. Eutheria (Gr., eu, true + therios + beast)
• Higher viviparous, placental mammals without marsupium.
• Order 3. Insectivora. (L., insectum, insect + vorare, to eat).
• Small primitive mammals with long pointed snout.
• E.g. Mole (Talpa), common shrew (Sorex), Solenodon (Solenodon), hedgehogs (Erinaceus,
Paraechinus).
• Order 4. Chiroptera. (Gr., Cheiros, hand + pteron, wing).
• Flying mammals or bats in which forelimbs are modified into wings (patagium).
• Order 5. Dermoptera. (Gr., derm, skin + pteron, wing).
• Four equal-sized limbs and tail included in a lateral furry skin fold, the patagium.
• Flying lemur, resembling a flying squirrel.
• E.g: One living genus Cynocephalus (= Galaeopithecus) with 2 species from Southeastern Asia
• Order 6. Edentata. (L„ edentatus, toothless)
• E.g. armadillo (Dasypus)
• Order 7. Pholidota. (Gr., pholis, a horny scale).
• E.g. scaly anteaters or pangolins (Manis).
• Tubulidentata. (L„ tubulus tube-like + dens, tooth).
• E.g. Cape anteater (Orycteropus) of South Africa.
• Order 9. Primates. (L., primus, of the first rank).
• 3 suborders : Lemuroidea (lemurs, Lemur & loris, Loris), Tarsioidea (Tarsius spectrum), and Anthropoidea
(Monkeys, apes and man).
• Order 10. Rodentia. (L„ rodo, gnaw).
• E.g. Rat (Rattus), mouse (Mus), squirrel (Funambulus)
• Order 11. Lagomorpha. (Gr., logos, hare + morphe, form).
• E.g. Rabbit (Oryctolagus), hare (Lepus)
• Order 12. Cetacea. (Gr., ketos or L., cetus, a whale).
• Two suborders : Odontoceti (toothed whales) and Mysticeti or Mystacoceti (whalebone whales).
• Order 13. Sirenia. (Gr., siren, sea nymph).
• Large, clumsy, herbivorous, aquatic mammals with paddle-like forelimbs, no hind limbs and a flattened tail
with horizontal lateral fleshy flukes with or without a notch.
• Manatee (Trichechus), dugong (Dugong = Halicore)
• Order 14. Carnivora. (L., caro, flesh + vorare, to eat).
• Dog (Canis familiaris), wolf (C. lupus), jackal (C. aureus), red fox (Vulpes),& Walrus (Odobenus), fur seal
• (Callorhinus), common seal (Phoca).
• Proboscidea. (Gr., pro, in front + boskein, to eat).
• Indian or Asiatic elephant (Elephas maximus), African elephant (Loxodonta ofricana), pigmy african elephant
(Elephas cyclotis).
• Order 17. Perissodactyla. (Gr., perissos, odd + dactylos, toes).
• Horse (Equus cabalus), wild ass (Equus asinus), zebra (Equus zebra)
• Order 18. Artiodactyla. (Gr., artios, even + dactylos, digit).
• Pig (Sus), common hippopotamus (Hippopotamus amphibius), camel (Camelus), deer (Carvus), sheep (Ovis),
goat (Capra), giraffe (Giraffa), blackbuck (Antilope), Ox (Bos indicus), water buffalo (Bubalus bubalis), yak
(Bos grunniens), bison (Bison).
 Hemichordates: ‘half chordates’

• marine “worms” with ‘apparent links to chordates on the one hand and to echinoderms on the other’
• chordates characteristics: pharyngeal slits present, a dorsal collar cord in the collar region (resembles the chordate
dorsal, hollow nerve tube), postanal appendage (in some)
• But, this appendage is not a derivative of the locomotor system, and hence a true postanal tail is absent.
• They also lack a notochord. Studies using gene expression fails to find homology between stomochord (hemichordate)
and notochord (chordate).
• Echinoderm affinities: similar ciliated bands, digestive system & planktonic lifestyle in both tornaria larvae and
auricularia larva (echinoderm)
 Origin of Chordates

• Chordates from Annelids and Arthropods

• ‘chordate body plan was derived from a flipped-over version of an
arthropod’s’, Geoffroy Saint-Hillaire, French zoologist (1822).
• Georges Cuvier led those opposing this theory, and in 1830 publicly
debated Saint-Hillaire.
• W. H. Gaskell and William Patten (20th century) supported (Segmented,
gross brain regionalisation)
• many of the supposed linking similarities between chordates and annelids
or arthropods result from homoplasy rather than homology
• Criticisms:
• Different pattern of myotomal segmentation
• Hollow nerve cord in chordates, embryological difefernt development
• Mouth and anus ventral in chordates
• Cleavage pattern of embryo different
• Proteostomes vs dueterostome
Chordates from Echinoderms
• ‘Auricularian Hypothesis’ (refers to auricularia larva found in holothurians, sea cucumbers)
• W. Garstang, a biologist of the late nineteenth and early twentieth centuries
• Echinoderms or a group very similar to echinoderms were the likely chordate ancestors.
• Echinoderms, like chordates, are deuterostomes,
• Share embryonic similarities of cleavage and mesodermal and coelomic formation.
• Echinoderm larvae, like chordates generally, are bilaterally symmetrical.
• Auricularian hypothesis begins with a dipleurula larva, an idealized version of this auricularian larva, thought
to represent the simplified ancestor of all echinoderm larvae.
• Garstang proposed that, in fact, chordate characteristics first debuted in this dipleurula larva
• Segmental swimming musculature, elongated body, and stiffened bar (notochord) are the supposed
solutions, first supplementing and then replacing the faltering ciliary system.
 But how might this echinoderm larva, now endowed with chordate characters,
achieve a separate evolutionary destiny from the adult it is fated to become at
metamorphosis?

Garstang’s answer
• ‘Paedomorphosis’
• Adult stage was eliminated and the larval stage enhanced.
• Larva became sexually mature, reproduce, and thus escape from a life cycle tied
to a benthic adult
• Early in the Cambrian period (nearly 530 mya) an immense variety of invertebrate animals inhabited Earth’s
oceans.
• Amidst this bustle, a slender, 3-cm long creatures (Myllokunmingia fengjiaoa) gliding through the water
• Although lacking armor and appendages, this ancient species was closely related to one of the most
successful groups of animals ever to swim, walk, slither, or fly: the vertebrates
Origin and evolution of vertebrates..
• For more than 150 million years, vertebrates were restricted to the oceans

• But about 365 mya they colonized land.

• Over time, as the descendants gave rise to the three groups of terrestrial vertebrates alive today:
amphibians, reptiles (including birds), and mammals.

• ˃ 57,000 species of vertebrates

• Disparity in mass prominent although lesser species diversity than invertebrates.

• the fish Schindleria brevipinguis, discovered in 2004, is just 8.4 mm long and has a mass roughly
100 billion times smaller than that of a blue whale.
Phylogeny of living chordates
Lancelets (Cephalochordata)
• The most basal (earliest diverging) group of living
chordates
• Bladelike shape, 6 cm in length
• The larvae feed on plankton in the water column
• Adult sinks and burrows with only anterior end exposed
to water
• Pharyngeal slits for respiration and filtering planktons (by
mucous network)
• Metameric somite muscles for lateral movement
 Tunicates (Urochordata)
• More closely related to other chordates than are lancelets.
• The chordate characters of tunicates are most apparent during their larval stage, which may be as brief as a few
minutes
• tunicates have 9 Hox genes, whereas all other chordates studied to date including the early-diverging lancelets
share a set of 13 Hox genes.
Lancelets and tunicates: clues to the evolutionary origin of
vertebrates!
 Advent of vertebrates: ‘chordates having a backbone’

• With a skeletal system and a more complex nervous system than that of their
ancestors, vertebrates became more efficient at two essential tasks:

‘capturing food and avoiding being eaten’

Derived Characters of Vertebrates
• As a result of gene duplication, vertebrates possess two or more sets of Hox genes (lancelets and
tunicates have only one).

• Other important families of genes that produce transcription factors and signaling molecules are
also duplicated in vertebrates.

• This resulting additional genetic complexity may be associated with:

• innovations in the vertebrate nervous system and skeleton,
• Such as development of a skull and a backbone composed of vertebrae.
Innovations of vertebrates
• Vertebral Column

• Head: cranium, or skull

Origin of vertebrates
• Took place in marine waters; however, some, fossil and physiological evidence seemed to point to a
freshwater origin.
• Earliest fish fossils consisted of fragments of bony armor worn smooth
• Physiologist Homer Smith (1930s) argued that the vertebrate kidney worked well to rid the body of any
osmotic influx of excess water, a problem among freshwater but not among marine animals.
• However, the discovery of still older fish fossils (Cambrian) confirmed the earliest vertebrates in marine
waters
• ‘Vertebrate kidney, while good at maintaining water balance, need not be interpreted as an innovation
of freshwater forms’
• Invertebrates (like lobsters & squid) and their ancestors have always been marine.
• Further, the Ordovician sediments first thought to be from fresh water, instead proved to be from
shallow, near-shore parts of the sea.
• Today, few scientists insist that the very first vertebrates were products of freshwater environments.
Evolution of early vertebrates was characterized
by
increasingly active lifestyles hypothesized
to proceed in
three major steps
 Evolution of vertebrates
Step 1
Comprised a suspension-feeding prevertebrate resembling Amphioxus.
It deployed only cilia to produce the food-bearing current that entered the pharynx.

Step 2
Comprised an agnathan, an early vertebrate lacking jaws
but possessing a muscular pump to produce a food-bearing water current.

Step 3
Comprised a gnathostome, a vertebrate with jaws, random food collection.
fed on larger food items with a muscularized mouth and jaws that rapidly snatched and selected prey
from the water.
Step 1: Prevertebrate
• This arose within the protochordates
• Two contrasting views: it was an incipient predator or suspension feeder.
• Suspension feeding based on ciliary pumps is common to hemichordates, urochordates, and
cephalochordates.
• Most likely a marine form, very similar to amphioxus, but a nonburrowing, free-swimmer better
able to tolerate estuary environments.
• Shift from such a prevertebrate to a vertebrate condition involved two mechanical changes in the
pharynx that together produced a muscular pump.
• First, the pharynx developed an encircling band of muscles.
• Second, strong and springy cartilage replaced the collagen in the pharyngeal bars.
• These adaptations resulted in capacity to increase body mass, evolutionary development of gills
 Origin of vertebrates.
• A more active, predaceous lifestyle characterized vertebrate evolution,
leading away from the suspension feeding that typified vertebrate
ancestors.

• Prevertebrates are envisioned as suspension feeders, perhaps something

like amphioxus, but they changed and came to depend on a muscularized
pharynx to produce feeding currents of water.

• Following prevertebrates, an agnathan stage developed in which adults

might have been benthic feeders, but larvae continued the trend toward a
more active lifestyle.

• Selection and capture of specific prey may have next led to the jawed
gnathostomes.

• Thus, the early trend in vertebrate evolution was from ciliary to muscular
mechanisms of moving feeding currents, and then to jaws that directly
snatched prey from water.
Step 2: Agnathan
• Appearance of a muscular pharyngeal pump brought early vertebrate evolution to the agnathan
stage.
• Extensive diversification by exploiting an expanded pharyngeal pump.
• Suspension feeders on an unusually thick soup of particles, or deposit feeders, mud grubbers
• Pushed their mouths into loose organic or silty mud and drew in sediment rich in organic particles
and microorganisms.
• The new muscularized pharynx, not cilia, forced the stream of rich organic material through the
mouth.
• Some fossil agnathans (ostracoderms) possibly created thick suspensions of food by using
roughened structures around their mouth to scrape growths of algae from rock surfaces.
 Step 3: Gnathostome
• The transition from agnathan to gnathostome involved a switch in feeding method taking advantage of
larger particles with more food mass.
• Transitional species became raptorial feeders that plucked individual food particles selectively from
suspension or off surfaces.
• Zooplankton, small worms that required forceful suction effort to be ingested.
• Raptorial and suction feeding favored a sudden and forceful expansion of the pharyngeal pump followed
by firm mouth closure to prevent escape of captured food.
• Early jawless vertebrates’ system was too weak to allow forceful capture and ingestion.
• With the advent of jaws powered by quick muscle action, pharyngeal expansion and suction became
strong and active.
• Muscles serving the anterior pharyngeal bar (near the mouth) became especially large, to open and close
the mouth quickly with a strong bite, securing the “inhaled” prey.
• The anterior pharyngeal bar enlarged, becoming the jaws, so that limits to prey size were also removed.