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Home , Acanthomorpha, Acanthopterygii, Acanthuroidei, Atherinomorpha, Beryciformes, Carangiformes

OEB 130: OF Lecture 16: Overview of acanthomorph diversity Outline

Lecture outline: Very brief review of fish phylogeny to put the in phylogenetic context The Acanthomorpha • Key characters of the Acanthomorpha • Major acanthomorph • A tour and very general overview of some of the key groups

Please note: 1. The diversity of Acanthomorph fishes is a bit overwhelming 2. The Diversity labs will help, as will the aquarium trip 3. Look closely at the phylogenies in this lecture and match the major groups up to the Encyclopedia of Fishes book; focus especially on clades with dots on slide 6 The

Gadiformes () Scopelomorpha + ()

Smelts viperfish

Acanthomorpha ~18,000 species

Otocephala Neoteleost fishes

Euteleostei Today! The Acanthomorph fishes Much recent progress in understanding the phylogenetic relationships of fishes – can be pretty confusing. The latest phylogeny is quite different from our understanding even 10 years ago, but things could still change!

Betancur, R. R., Broughton, R. E., Wiley, E. O., Carpenter, K., Lopez, J. A., Li, C., Holcroft, N. I., Arcila, D., Sanciangco, M., Cureton Ii, J. C. et al. (2013). The tree of life and a new classification of bony fishes. PLoS currents 5.

Near, T. J., Dornburg, A., Eytan, R. I., Keck, B. P., Smith, W. L., Kuhn, K. L., Moore, J. A., Price, S. A., Burbrink, F. T., Friedman, M. et al. (2013). Phylogeny and tempo of diversification in the superradiation of spiny-rayed fishes. Proc. Nat. Acad. Sci. USA. 110, 12738–12743.

Near, T. J., Eytan, R. I., Dornburg, A., Kuhn, K. L., Moore, J. A., Davis, M. P., Wainwright, P. C., Friedman, M. and Smith, W. L. (2012). Resolution of ray-finned fish phylogeny and timing of diversification. Proceedings of the National Academy of Sciences of the United States of America 109, 13698–13703.

Another useful resource: http://www.fishbase.org/home.htm

The Acanthomorpha– more detailed branching

Tunas, and relatives

(pp. 168-171) Toadfishes

(Batrachoidiformes) Many more (pp. 135-136) Beryciformes

(pp. 160-164) (Next phylogeny (Next phylogeny slide)

Gadiformes (cods)

(pp. 130-132)+ zeiformes Lampridiformes (pp. 157-159) Percomorpha

(pp. 165-167)

Page numbers in green refer to the Encyclopedia of Fishes book relevant Acanthomorpha pages ~18,000 species Don’t panic! Note the red dots and the major groups that we’ll be looking at. Another view

The Acanthomorpha– more detailed branching

Tunas, seahorses and relatives

Toadfishes

(Batrachoidiformes) Many more

Beryciformes (Next phylogeny (Next phylogeny slide)

Gadiformes (cods) + zeiformes Lampridiformes Percomorpha

Acanthopterygii Beardfishes

The branching pattern is what is important to understand as it reflects Acanthomorpha the evolutionary history ~18,000 species of these fish groups. The Acanthomorph fishes (These names can cause confusion!)

Outline summary of some major names to keep in mind:

Actinopterygii (ray-finned fishes) Euteleostei Acanthomorpha Acanthopterygii Percomorpha (-like fishes) Many “orders” of fishes in this group e.g., (an “” of perch-like fishes

Names ending in “morpha” usually indicate a “superorder” Names ending in “es” usually indicate an “order” of fishes Names ending in “ei” usually indicate a “suborder” of fishes Names ending in “ae” usually indicate a “” of fishes Acanthomorpha (spiny-finned fishes)

Key characters for the Acanthomorpha: (means "thorn-shaped") 1. Dorsal and anal spines 2. Rostral cartilage (under premax) 3. Pectoral girdle 4. skeleton Key characters for the Acanthopterygii: (means "spiny finned") 1. Pelvic spine present 2. Pelvic fin bone support anatomy 3. Ligament anatomy associated with upper jaw protrusion 4. Two distinct regions of : one spiny and the other flexible

(also: ctenoid scales and physoclistous bladder in most species) Acanthomorpha: Polymixiomorpha “beardfishes”

• Mostly open : 200 - 600 meters deep • 10 species • An odd mixture of and derived anatomical traits • Chin barbels Acanthomorpha Gadiformes (cods and relatives)

Female can produce up to 9 million eggs per spawning season

Rattail fish; (, benthic)

Burbot … freshwater (popular for ice- in Canada)

• Mostly salt water (only 20 species are freshwater) • Includes cods, trout-, , - Acanthopterygii: Zeiformes (dories)

• Some • 2 species; highly protrusible jaws; tasty too Acanthomorpha: Lampriomorpha “lamprid fishes”

Wegner, N. C., Snodgrass, O. E., Dewar, H. and Hyde, J. R. (2015). Whole-body endothermy in a mesopelagic fish, the , . Science 348, 786-789.

• Mostly open water, oceanic, and a strange diversity of shapes • 21 species: 7 families • Species of note include the opah and • Unique method of upper jaw protrusion Acanthomorpha: Lampriomorpha “lamprid fishes” Acanthopterygii: Beryciformes (soldierfishes, squirrelfishes, and roughies)

• 144 species: many are popular aquarium fishes • Mostly inhabit coral reefs • Roughies are deep sea species: a popular food fish, sadly as they don’t reproduce until age 20

The Acanthomorpha – more detailed branching

Tunas, seahorses and relatives

Toadfishes

(Batrachoidiformes) Many more

Beryciformes (Next phylogeny (Next phylogeny slide)

Gadiformes (cods) + zeiformes Lampridiformes Percomorpha (~ 13,000 species. Greatest diversity is marine.)

Acanthopterygii Beardfishes

Acanthomorpha ~18,000 species Acanthopterygii: Percomorpha Batrachoidiformes (toadfishes; midshipmen)

Bottom dwellers; marine Lack pleural ribs Large mouths Able to produce sound Muscular vibrations of arranged on their belly like the buttons of a midshipmen’s uniform

Acanthopterygii: Percomorpha: Tunas, seahorses and relatives Suborder

• Upper jaw not protrusible; teeth fused to jaw bones • ~147 species: most of great commercial importance (; sushi!) • Groups of note include the tunas and • Active predators with elevated body temperatures • Some species have heated retinas for enhanced vision • Tunas, mackerel, cutlassfishes Acanthopterygii: Percomorpha: Suborder Scombroidei

tuna mackerel Acanthopterygii: Percomorpha: Suborder Scombroidei

This group is known for exceptionally fast locomotion, long distance migration, many species have warm red muscle that is internalized, and streamlining; fold into groves on the body

finlets Acanthopterygii: Percomorpha (, seahorses, trumpetfish) Tunas, seahorses and relatives

males brood eggs and young possess pouch (eggs from multiple females) long snouts feed via sucking food into mouths also: trumpetfishes and coronet fishes tropical reef dwellers Acanthopterygii: Percomorpha: (, pipefishes, seahorses)

The Percomorpha– more detailed branching

Xiphiformes Pleuronectiformes , () swordfishes (pp. 223-226) Carangids (jacks) Anabantoids Swampeels etc. (pp. 173-174) , Many more Cardinalfishes

(pp. 218-222) (Next phylogeny (Next phylogeny slide) (jacks and relatives)

Tunas, seahorses and relatives Acanthopterygii: Percomorpha: Gobioidei

• Gobies: • 2000 species: very diverse mostly marine group • Some freshwater too • Often extremely colorful • Includes the mudskippers which inhabit mudflats and mangroves • Pelvic fins often united into an adhesive sucking disc for attachment to substrates Acanthopterygii: Percomorpha: Suborder Gobioidei Acanthopterygii: Percomorpha: Suborder Gobioidei

Mudskippers Acanthopterygii: Percomorpha: Suborder

• “Labyrinth” fishes: have an accessory labyrinth-like air-breathing organ above the • 120 species • Species of note include the bettas and Acanthopterygii: Percomorpha:

319.6-pound halibut Pleuronectiformes (flatfishes, soles)

• primarily benthic; important food fishes • 680 species: very diverse group • Species of note include flounder, halibut • flattened, with both eyes migrating to one side of the head Acanthopterygii: Percomorpha: Pleuronectiformes (flatfishes) Acanthopterygii: Percomorpha: Pleuronectiformes (flatfishes) Acanthopterygii: Percomorpha: Carangiformes

• NOT closely related to tunas! • Atlantic, Pacific, and Indian Xiphiidae: Swordfish • Sword is dorsoventrally flattened • Lack teeth as adults and pelvic fin • Highly prized as food fish Acanthopterygii: Percomorpha: Carangiformes

Istiphoridae: and billfishes

• Bills are more rounded in cross-section • Possess teeth and pelvic fins • Highly prized sport fish • heat exchanger to keep brain and eye warm during forays into cold deep water Acanthopterygii: Percomorpha: Carangiformes

• “jacks” and • Marine Atlantic, Indian, Pacific oceans • About 140 species • Deep and laterally compressed body The Percomorpha – more detailed branching

Mullets,

Surfperches Blennies (pp. 214-217) (Damselfishes) Cichlidae (pp. 205-208) () (silversides, Clingfishes (pp. 200-204) , (pp. 142-143) live-bearing fishes

(pp. 144-156)

Many Many more (Next phylogeny (Next phylogeny slide) Eupercaria Jacks and relatives, , labyrinth fishes, and others

Ovalentaria: a very important Acanthopterygii: Percomorpha: – a major clade

Ovalentaria: this group has demersal sticky eggs (well, most species)

Damselfish eggs just before hatching Percomorpha: Ovalentaria: Atherinomorpha (silversides, , needlefishes, flying fishes)

• Mostly surface-dwellers and shallow water marine • 312 species; have long snouts • Unique jaw protrusion mechanism • Livebearers have considerable sexual dimorphism Percomorpha: Ovalentaria: Atherinomorpha (silversides, livebearers, needlefishes, flying fishes) Percomorpha: Ovalentaria: Atherinomorpha (silversides, livebearers, needlefishes, flying fishes) Percomorpha: Ovalentaria: Atherinomorpha (silversides, livebearers, needlefishes, flying fishes)

killifishes, topminnows, and livebearers • no spiny rays • small in size (some aquarium fishes) • livebearers : • in estuaries () • livebearers () • internal fertilization (intromittent organ) mosquitofishes: • eat mosquito larvae

Percomorpha: Ovalentaria: African Family Cichlidae

• Biologically and economically important • Africa and South/Central America • Diverse group with ~1500 species • Tilapia impt. for aquaculture: invasive • Rapid speciation has resulted in much evolutionary study Percomorpha: Ovalentaria: South Family Cichlidae American Acanthopterygii: Percomorpha: Ovalentaria – (mullets)

• Abundant in warm climates • 72 species: used for food throughout world • coastal shallow-water marine species • Filter feeders Acanthopterygii: Percomorpha: Ovalentaria: Pomacentridae (damselfishes)

• Pomacentridae: damselfishes, clownfishes, anemonefishes (350 species) • Tropical and primarily indo-pacific, very colorful Acanthopterygii: Percomorpha: Ovalentaria: Pomacentridae (damselfishes)

• Pomacentridae: damselfishes, anemonefishes (350 species) • Tropical and primarily indo-pacific; very colorful Acanthopterygii: Percomorpha: Ovalentaria: Suborder Blennioidei

• Mostly small benthic fishes; mostly marine; very diverse • ~400 species; arch and hypural bone characters Acanthopterygii: Percomorpha: Ovalentaria: Blenniidae

• Blennies: • ~ 400 species: quite diverse mostly marine group • Most have lost scales • Most have lost swimbladder • Often mimic other species The Percomorpha – more detailed branching Snailfish, Sticklebacks, Groupers Sculpins (pp. 171-172) Icefishes (pp. 175-180) (pp. 195-199) Zoarcids Kyphosidae

Sunfishes Hawkfishes (Sea chubs) Labridae , ,

and relatives Many more

(pp. 209-213) (Next phylogeny (Next phylogeny slide)

Some Scorpaeniform perch-like fishes fishes

Ovalentaria

Eupercaria Acanthopterygii: Percomorpha: Labridae and Scaridae (wrasses; parrotfishes)

• Very colorful and common reef fish • Excellent pectoral fin swimmers • Specialized pharyngeal jaws Acanthopterygii: Percomorpha: Gasterosteiformes sticklebacks

Sticklebacks: -- mainly (some marine species or morphs) with large spines assoc. w/ dorsal fins -- Important for genetic and evolutionary studies -- Some have armored scutes along side of body -- Males build nests and become red in belly -- Many studies of behavior

Acanthopterygii: Percomorpha: (scorpionfishes)

• Many lack scales; primarily benthic; many are poisonous • Large bony skulls with spines – many species poisonous • 1500 species: very diverse group • Species of note include the lionfish, lumpfish, sea robin Acanthopterygii: Percomorpha: Scorpaeniformes (scorpionfishes) Acanthopterygii: Percomorpha: Notothenioidei: icefishes

• “icefishes” or “cod icefishes” ; coastal Antarctic • 125 species: many lack hemoglobin; have antifreeze proteins • Weakly ossified Acanthopterygii: Percomorpha: Serranidae: groupers and relatives

• Very diverse; includes groupers • 475 species • Marine tropical; spines on Acanthopterygii: Percomorpha: Centrarchidae: North American sunfishes and

• Restricted to N. America (but introduced on other continents too) • Micropterus – “basses” • Lepomis – sunfishes – and • Pomoxis – black and white crappies • Important recreational species

• Sea chub: Kyphosidae • Very friendly and curious when diving! • ~55 species • Marine Atlantic, Indian and Pacific • Both herbivorous and carnivorous species Acanthopterygii: Percomorpha:

• Perches and darters • 10 genera with 201 species, most (187) in North America • Darters most speciose with 184 species: mostly benthic living in streams The Percomorpha – more detailed branching

Lophiiformes () (pp. 137-141) Tetraodontiforms (puffer fishes and relatives) (Angelfishes) (pp. 227-231) Haemulidae Surgeonfish

Sunfishes and some other perch-like fishes Scorpaeniform fishes Acanthopterygii: Percomorpha: (surgeonfishes and relatives)

Acanthuroidei: a suborder of the Percomorpha Includes 6 families and 130 species.

Most species are herbivorous, have restricted gill openings, and have a planktonic larval stage with larvae that are nearly transparent.

Includes the surgeonfishes, tangs, spadefishes, and moorish idols among others.

Acanthopterygii: Percomorpha: Acanthuroidei (surgeonfishes and relatives)

Blue tang = “Dory”

• : surgeonfishes, unicornfishes, tangs (80 species) • Worldwide marine waters (tropical and subtropical) • Spine on caudal peduncle (base of tail) Acanthopterygii: Percomorpha: Chaetodontidae (butterflyfishes)

• Chaetodontidae: butterflyfishes (122 species) • Atlantic, Indian, and pacific waters; body laterally compressed • Bright coloration, eyespot often present, usually live in shallow waters • Have a specialized -swimbladder connection Acanthopterygii: Percomorpha: Pomacanthidae

• Pomacanthidae: angelfishes (85 species) • Marine, shallow tropical waters; colorful • Color may change dramatically from young to adult • Not the freshwater angelfish, which is a ! Acanthopterygii: Percomorpha: Lutjanidae

• “snappers” • Marine tropical • 105 species • Important food fishes Acanthomorpha: Percomorpha Lophiiformes (goosefish, anglerfishes)

Goosefish, Lophius

“Handfishes”, Pediculati (old name) ~ 300+ species

Bat fish Acanthomorpha: Percomorpha Lophiiformes (goosefish, anglerfishes)

“”

Deep-sea anglerfishes can have extreme sexual dimorphism Dorsal fin ray modified into a lure Acanthopterygii: Percomorpha: (, pufferfishes, and porcupine fishes)

• 360 species; high degree of skull bone fusion • Scales modified into bony plates • Groups of note also include the boxfishes, pufferfishes • Includes the bizarre ocean “sunfish” Mola mola • Strong jaws; defense mechanisms to avoid • Fugu – toxic internal organs • Some species have the ability to inflate themselves with water Acanthopterygii: Percomorpha: Tetraodontiformes (triggerfishes, pufferfishes, and porcupine fishes) Acanthopterygii: Percomorpha: Tetraodontiformes (triggerfishes, pufferfishes, and porcupine fishes)

Scales form tight-fitting bony plates Acanthopterygii: Percomorpha: Tetraodontiformes (triggerfishes, pufferfishes, and porcupine fishes) Acanthopterygii: Percomorpha: Tetraodontiformes (triggerfishes, pufferfishes, and porcupine fishes)

Movie of inflation From Prof. Beth Brainerd, Brown Univ. Acanthopterygii: Percomorpha: Tetraodontiformes (triggerfishes, pufferfishes, and porcupine fishes)

• Strong dorsal spines • Wedge themselves into corals to avoid being eaten Acanthopterygii: Percomorpha: Tetraodontiformes (triggerfishes, pufferfishes, and porcupine fishes)

clown Acanthopterygii: Percomorpha: Tetraodontiformes (triggerfishes, pufferfishes, and porcupine fishes)

Very bizarre! (Mola mola): no tail; dorsal and anal fins meet Locomotion is by the dorsal and anal fins Acanthopterygii: Percomorpha: Tetraodontiformes (triggerfishes, pufferfishes, and porcupine fishes)

• Ocean “sunfishes” • Large • Thick leathery skin • Eat jellyfish