Euteleostei
Transcription
Euteleostei
OEB 130: BIOLOGY OF FISHES Overview of teleost fish diversity Outline Lecture outline: Very brief review of fish phylogeny: sarcopterygians and early actinopterygians The teleost fish clade (broad overview) • What is a teleost fish? • Major trends in teleost fish evolution -- fin placement, scales, tail shape -- maxilla and premaxilla • Introduction to the major groups -- Elopomorpha and Osteoglossomorpha -- Otocephala – (much more in March 30 lecture) -- Euteleostei -- Salmons and relatives -- Neoteleostei, viperfish, Scopelomorpha -- Acanthomorpha: Beryciformes and Percomorpha Exam next Monday IMPT. exam notes: 1) First hour exam next Monday Feb. 29th. 2) The exam will last 50 minutes 3) Questions will be very short and short answer 4) No multiple choice questions 5) Exam covers both the lectures and readings 6) Pay special attention to the drawings done in class, with a focus on the phylogenies. Practice drawing these for yourself, and correlate the major groups with the readings that you did First exam in one week! Vertebrate tree – Sarcopterygii (lobe-finned fishes) ~35,000 ray-finned fishes ~ 30,000 lobe-finned “fishes” Major divergence in vertebrate evolution Ray-finned fish (Actinopterygii) phylogeny: broad overview Actinopterygii Holostei Chondrichthyes Polypterus and relatives Chondrostei Teleostei † Cheirolepis 1. Single dorsal fin 2. Fin ray structure: lepidotrichia 3. Scale structure Actinopterygii = ray-fin Teleost fish phylogeny: broad overview Otocephala Elopomorpha (tarpons and eels) Clupeiformes Ostariophysi Osteoglossomorpha (“bony tongues”) Euteleostei Teleostei 1. Mobile premaxilla bone in the skull 2. Specialized tail bones 3. Median basibranchial tooth plates What is a teleost fish? Selected key shared derived characters (synapomorphies) Teleostei: 1) Mobile premaxilla bone in head 2) Elongate bones in the tail skeleton – uroneurals (yellow in diagram below) 3) Median basibranchial tooth plates 2 1 Remember the premaxilla with the ascending process neurocranium Ascending process eye operculum Pectoral girdle suspensorium Premaxilla maxilla mandible Jaw joint hyoid Pectoral fin Teleost character (1) Mobile premaxilla bone in head Early actinopterygians have a fixed premaxilla and maxilla in the upper jaw. The maxilla is actually fixed in place too. Functional implications? This is the head of a paleoniscoid fish, an early fossil actinopterygian. Teleost character (1) Mobile premaxilla bone in head Early actinopts: no mobile bones in the upper jaw Amia (the bowfin) has a mobile maxilla (blue arrow) Black area and arrow shows joint where movement can occur. Teleost fishes like Elops (the ladyfish or ten-pounder) has a both mobile maxilla and premaxilla (blue arrows) Teleost character (1) Mobile premaxilla bone in head pmx maxilla Many changes in the maxilla-premaxilla relationship during teleost evolution. Important functional implications here too! Ascending process (allows significant jaw protrusion) Teleost character (2): Elongate bones in the tail skeleton: uroneurals Teleost character (2): Elongate bones in the tail skeleton: uroneurals Caudal fin rays uroneural Modified last caudal vertebra Vertebra Haemal spine Hypural plate (midline) Teleost character (2): Elongate bones in the tail skeleton: uroneurals Amia calva (bowfin) – pre-teleost Elongate uroneural bones Early (fossil) teleost fish tail skeleton Functional implications are unknown: stiffen the upper lobe? Teleost character (2): Elongate bones in the tail skeleton: uroneurals The teleost fish tail Externally symmetrical tail … But internally asymmetrical skeleton Teleost character (2): some diversity of tail shapes Non-teleost tails Teleost tail Teleost character (3): Median basibranchial tooth plates Primitive actinop. condition – small toothplates scattered among the ventral gill arch elements (as in bichirs, gar, etc) Looking down from above on the ventral gill arches, to show the basibranchial tooth plates. Teleost condition – midline toothplates on the ventral gill arch elements Teleost fish phylogeny: broad overview Otocephala Elopomorpha (tarpons and eels) Clupeiformes Ostariophysi Osteoglossomorpha (“bony tongues”) Euteleostei Teleostei 1. Mobile premaxilla bone in the skull 2. Specialized tail bones 3. Median basibranchial tooth plates Actinopterygii = ray-fin Some major trends in fish evolution: fin positions Pectoral fin Note change in the relative position of these fins. Pelvic fin Evolutionary patterns of fins – another view pectoral fins pelvic fins dorsal fin(s) anal fin caudal fin (spiny-finned fishes) (“true” bony fishes) (ray-finned fishes – 28,000 species) Many unanswered questions: •Effect of fin position? •Effect of fin shape? •Hydrodynamic and mechanical function? (spiny-finned fishes) (“true” bony fishes) (ray-finned fishes – 28,000 species) Some major trends in fish evolution: scales Thinner, lighter cycloid and ctenoid scales replace the heavy dermal armor of primitive ray-finned fishes. Some fishes (e.g. eels) lack scales. Diagram is missing the peg and socket Fish scales Teleost fish phylogeny: broad overview Otocephala Elopomorpha (tarpons and eels) Clupeiformes Ostariophysi Osteoglossomorpha (“bony tongues”) More detail in a later slide Euteleostei Teleostei 1. Mobile premaxilla bone in the skull 2. Specialized tail bones 3. Median basibranchial tooth plates Actinopterygii = ray-fin Elopomorpha (Eels, tarpons) Eels and tarpons and relatives have leptocephalus larvae (leaf-head) • Eels are the best known members with an amazing life history • Include moray eels, conger eels, deep sea “spiny eels” and gulper eels • Bonefishes, tarpon, tenpounders (ladyfishes) – good fishing! • 24 families, around 860 species • United by the presence of a leptocephalus larva Elopomorpha (Eels, tarpons) A moray eel: recall the raptorial pharyngeal jaws Elopomorpha (Eels, tarpons) tarpon/ladyfish Elopomorpha (Eels, tarpons) The leptocephalus larva, a synapomorphy for elopomorph fishes Leptocephalus = “leaf head” Elopomorpha (Eels, tarpons) Metamorphosis of an eel larva Elopomorpha (Eels, tarpons) Movie of locomotion in a leptocephalus Elopomorpha (Eels, tarpons) Movie of locomotion in a leptocephalus Swimming of a leptocephalus larva Elopomorpha (Eels, tarpons) Freshwater eels are catadromous, they enter the ocean as adults, migrate to a spawning area in the Sargasso sea, where they spawn & then die. Live most of their lives in fresh water. Sargasso sea Elopomorpha (Eels, tarpons) Freshwater eels are catadromous, they enter the ocean as adults, migrate to a spawning area where they spawn & then die. Teleostei: Osteoglossomorpha (bonytongues) • • • • 4 families, around 220 species. Often exhibit parental care of young Clown knifefish, mooneye, arowanas, butterfly fish Elephant fishes (Mormyridae): 201 species, freshwater, African, and these fishes can both generate and detect weak electric fields for communication and prey location Osteoglossomorpha (bonytongues) arowana Elephant fish African knifefish Osteoglossomorpha (bonytongues) Separate lecture on the Otocephala clade coming on March 30. Otocephala 2 major groups Clupeomorpha 364 species herrings, anchovies, menhaden, etc. Abundant ocean schooling fishes of great commercial importance. Ear-swimbladder anatomy Caudal skeleton anatomy Ostariophysi 8,000 species 28% of all fishes 68% of freshwater fishes; carps, milkfishes, characins, zebrafish, goldfish, many popular aquarium species, catfishes, and gymnotids – South American knifefishes Otocephala: Clupeomorph clade Otocephala: Ostariophysi clade Much more in the March 30 lecture. Teleost fish phylogeny: broad overview Otocephala Elopomorpha (tarpons and eels) Clupeiformes Ostariophysi Osteoglossomorpha (“bony tongues”) Euteleostei Teleostei More detail next slide 1. Mobile premaxilla bone in the skull 2. Specialized tail bones 3. Median basibranchial tooth plates Actinopterygii = ray-fin The Euteleostei – more detailed branching Scopelomorpha (Lanternfish) Gadiformes (cods) + zeiformes Beryciformes Smelts viperfish Percomorpha Protacanthopterygii Acanthomorpha (~17,000 species) Otocephala (~9,000 sp) Neoteleost fishes: 1 trait Euteleostei: ~ 3 traits Euteleost = “true” teleost Euteleost fishes (the Euteleostei) • • • • Tremendous species diversity Characters defining the clade less than solid Around 20,000 species 346 families Figure 24.15 Selected key shared derived characters for Euteleostei Euteleostei: 1) Dorsal adipose fin – lost in many later clades ** 2) Nuptial tubercles on head and body – lost in many clades ** 3) Unique anatomy of the uroneural bones and other 1 skeletal elements 2 1 ** traditional characters but pretty lame; molecular analyses support this grouping too One fairly good character for Euteleostei 3 Bone stained red, cartilage blue Euteleost character 3 – midline caudal skeleton cartilage The Euteleostei – more detailed branching Scopelomorpha (Lanternfish) Gadiformes (cods) + zeiformes Beryciformes Smelts viperfish Percomorpha Protacanthopterygii Acanthomorpha Otocephala Neoteleost fishes: 1 trait Euteleostei: ~ 3 traits Euteleost = “true” teleost Protacanthopterygii argentiniform (202 species) and salmoniform (154 species) fishes • Not a very well-defined group and some controversy over the included species and higher taxa. • Includes the esocids – pikes and mudminnows • Trout and salmon, and walleye • Argentinoid fishes, smelts, and some other odd deep-sea fishes • 12 families, 366 species Protacanthopterygii Freshwater pike mudminnow Trout and salmon The Euteleostei – more detailed branching Scopelomorpha (Lanternfish) Gadiformes (cods) + zeiformes Beryciformes Smelts viperfish Percomorpha Protacanthopterygii Acanthomorpha Otocephala Neoteleost fishes: 1 trait Euteleostei: ~ 3 traits Euteleost = “true” teleost The Neoteleostei major defining character is the retractor dorsalis muscle Remember the UPJ! 1 2 3 4 5 Premaxilla Lower jaw Basihyal (“tongue”) Hyoid Urohyal – part of the hyoid embedded in the sternohyoideus muscle Pectoral girdle The Neoteleostei major defining character is the retractor dorsalis muscle Remember the retractor dorsalis muscle The Euteleostei – more detailed branching Scopelomorpha (Lanternfish) Gadiformes (cods) + zeiformes Beryciformes Smelts viperfish Percomorpha Protacanthopterygii Acanthomorpha Otocephala Neoteleost fishes: 1 trait Euteleostei: ~ 3 traits Euteleost = “true” teleost Viperfish (Stenopterygii) • Mostly deep-sea fishes: meso- and bathypelagic • 391 species: hatchetfishes, bristlemouths • Families of note include the Stomiidae (stomiiform fishes -dragonfishes) • Photophores, barbels, big teeth!, dark color The Euteleostei – more detailed branching Scopelomorpha (Lanternfish) Gadiformes (cods) + zeiformes Beryciformes Smelts viperfish Percomorpha Protacanthopterygii Acanthomorpha Otocephala Neoteleost fishes: 1 trait Euteleostei: ~ 3 traits Euteleost = “true” teleost Scopelomorpha (lanternfishes / myctophiform fishes) • • • • • 2 families, around 250 species. Lanternfishes / myctophiform fishes Highly abundant group of mesopelagic fishes in the oceans Often diurnal migration from surface to 2000 m presence of non-bacterial luminescent organs (forming the lanterns of these so-called lanternfishes). • myctophid light organs are photophores and are species specific Scopelomorpha (lanternfishes) • • • • • 2 families, around 250 species. Lanternfishes / myctophiform fishes Highly abundant group of mesopelagic fishes in the oceans Often diurnal migration from surface to 2000 m presence of non-bacterial luminescent organs (forming the lanterns of these so-called lanternfishes). • myctophid light organs are photophores and are species specific The Euteleostei – more detailed branching Scopelomorpha (Lanternfish) Gadiformes (cods) + zeiformes Beryciformes Smelts viperfish Protacanthopterygii Percomorpha Acanthomorpha Future lectures on the Acanthomorpha Otocephala Neoteleost fishes: 1 trait Euteleostei: ~ 3 traits Euteleost = “true” teleost