LECTURE 6 - OUTLINE. Evolution & Classification - Part II. Agnatha (cont.) Gnathostomata

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LECTURE 6 - OUTLINE Evolution & Classification - Part II Agnatha (cont.) 6. Myxini 7. Cephalaspidomorphi Gnathostomata 1. Phylogenetic relationships 2. Placodermi 3. Acanthodii BIOL 4340 Lecture 6-1 Class Myxini (hagfishes) - cranium poorly developed and vertebrae absent - mouth four rows of keratinized teeth - feed on soft-bodied invertebrates & dead or dying organisms - water taken in through a single nostril connected to pharynx BIOL 4340 Lecture 6-2 1

Class Myxini (hagfishes) - 75 to 200 slime glands (each side) along ventrolateral body wall - each ovoid gland (1.5 to 3 mm diameter) with numerous cells - slime exudate makes up to 3 4% of hagfish body mass 60g hagfish = 24L slime!? Slime composition 1. 99.996% seawater 2. 0.0015% mucin 3. 0.002% slime thread Hagfish slime = 0.015 mg/ml mucin Salivary mucus = 14 mg/ml mucin BIOL 4340 Lecture 6-3 Class Myxini (hagfishes) - 75 to 200 slime glands (each side) along ventrolateral body wall - each ovoid gland (1.5 to 3 mm diameter) with numerous cells - slime exudate makes up to 3 4% of hagfish body mass 1. mucin vesicles (arrowhead) - holocrine release of cells 2. Slime threads (arrow) BIOL 4340 Lecture 6-4 2

Class Myxini (hagfishes) - rapid - forceful - local slime formation requires mixing! http://www.youtube.com/watch?v=bb2eop3ohne BIOL 4340 Lecture 6-5 Class Myxini (hagfishes) - slime functions include: - aid in burrowing - cover scavenged meal (mask/repellant/suffocate) - defense 1. predation 2. digestive enzymes Lim et al 2006. J Exp Biol 209: 702-710 BIOL 4340 Lecture 6-6 3

Class Myxini (hagfishes) Myxinid Reproduction - a single unpaired gonad - immature individuals anterior = ovarian tissue posterior = testicular tissue - fertilization external (male has no intromittent organ) - eggs produced in batches of 20-30 Hagfish eggs, ~ 3cm long BIOL 4340 Lecture 6-7 Class Myxini (hagfishes) Myxinid Reproduction - what we don t know - spawning times - spawning location - spawning frequency - spawning behaviour - embryological details (not observed since 1890s) - reproductive lifespan Cash prize for information on reproductive habits issued in 1854 (Copenhagen Academy of Sciences) still unclaimed BIOL 4340 Lecture 6-8 4

Cambrian (544-505 mbp) - to date - varied body shapes, suggesting varied lifestyles - fossil cephalaspidomorphs heavily armoured - united by the presence of a single nasal opening on top of the head - nasal opening leads to olfactory sac and hypophyseal sac to form a nasohypophysis BIOL 4340 Lecture 6-9 brain olfactory sac nasal opening hypophyseal sac branchial skeleton vertebral elements BIOL 4340 Lecture 6-10 5

Order Petromyzontiformes - contains extant forms (lampreys) - caudal fin and 1-2 dorsal fins present (no paired fins) - 7 gill openings along side of body - eyes and lateral line system both well developed - prominent oral disk with numerous tooth-like plates of keratin Ammocoete larva - body form, appearance & life habit similar to the lancelet BIOL 4340 Lecture 6-11 - oral disc attaches primarily by suction but also with the teeth - lingual cartilage and tongue muscles protrude tongue - rasping erodes tissue which is ingested with body fluids - for respiration, water is pumped in and out of gill openings BIOL 4340 Lecture 6-12 6

Order Petromyzontiformes - contains extant forms (lampreys) ~ 31 recognized species : 16 freshwater nonparasitic : 9 anadromous parasitic : 6 freshwater parasitic BIOL 4340 Lecture 6-13 : starvation Lamprey Life Cycles - parasitic European river lamprey (Lampetra fluviatilis) - anadromous Time (years) I II III IV V VI VII ~ 10-13 cm ~ 50 cm freshwater marine : larval phase : transformation : migration : parasitic phase : spawning ammocoete larva macrophthalmia transformation (4-5 weeks) BIOL 4340 Lecture 6-14 7

Lamprey Life Cycles - nonparasitic European brook lamprey (Lampetra planeri) non-anadromous Time (years) I II III IV V VI VII Max 20 cm freshwater : larval phase : transformation : migration/maturation : parasitic phase : spawning BIOL 4340 Lecture 6-15 Direction of Water Flow x.s. 20 cm BIOL 4340 Lecture 6-16 8

Female - leading edge of 2nd dorsal fin - region in front of the cloaca Male - spur or penis BIOL 4340 Lecture 6-17 Alterations in body form - reduced length/weight - changes in dentition - degeneration of eyes - enlarged oral disk Alterations in physiology - atrophy of intestine - closure of foregut - depletion of: (i) lipids ( in H2O) (ii) liver glycogen BIOL 4340 Lecture 6-18 9