15_Protacanthopterygii

Phylum Chordata Subphylum Urochordata (tunicates & sea squirts Subphylum Cephalochordata (lancelets) Subphylum Craniata Superclass Myxinomorphi V E R TE B R A T A J A W L E S S J A W E D Superclass Petromyzontomorphi Superclass Conodonta Superclass Pteraspidomorphi Superclass Anaspida Superclass Thelodonti Superclass Osteostracomorphi Superclass Gnathostomata Class Placodermi Class Chondrichthyes Subclass Holocephali Subclass Elasmobranchii Class Acanthodii Class Actinopterygii Subclass Cladistia Subclass Chondrostei Subclass Neopterygii Class Sarcopterygii Subclass Coelacanthomorpha Subclass Dipnotetrapodomorpha

autapomorphies unique derived traits Osteoglossomorpha Elopomorpha Ostarioclupeomorpha synapomorphies 1 mobile premaxilla, uroneurals 3 two uroneural bones extend over second caudal centrum, abundant intramuscular bones in caudal and abdominal regions 5 neural arch of first caudal vertebra reduced or missing, upper pharyngeal jaws fused to gill arch elements, jaw joint with unique articulation and ossification 7 dorsal adipose fin and nuptial tubercles on head and body, first uroneural bones of tail have paired anterior membraneous outgrowth synapomorphies shared derived traits Euteleostei 28 Orders, 346 Families 17,419 spp* DIVISION TELEOSTEI *Nelson 2006

Subdivision Euteleostei Monophyly of the subdivision and its organization area of debate Johnson and Patterson (1996) proposed a scheme, that emphasizes three shared traits: 1 Similarities in the pattern of embryonic development of the supraneural bones among euteleosts, which are the small T-shaped or rodlike bony or cartilaginous elements that lie within the musculature between the cranium and the dorsal fin. 2 The presence of an outgrowth on the stegural bone, a structure associated with the neural arches of the vertebral centra of the tail base. 3 The presence of caudal median cartilages, cartilages that lie between the hypural bones of the caudal base page 275-276 Text

Division Teleostei Subdivision Euteleostei? Superorder Protacanthopterygii Nelson 2006: classification has been and continues to be unstable characters exhibit a mosaic distribution, show reduction, are otherwise highly modified, or are primitive for the euteleosts Monophyly and definition of some groups are a matter of debate and no synapomorphies are given (i.e., the Protacanthopterygii (osmeriform smelts, salmoniform salmons) remain a problematic group that lacks well-defined, unifying characteristics).

Superorder Protacanthopterygii DEFINING TRAITS: Maxillary included in gape Adipose fin Lack fin spines Physostomus Physoclistus Swimbladder absent (some - acellular bone, 1st vertebra articulates with 3 bones, uroneural plate) Ancestral or Derived?

Table 1.1 The diversity of living fishes. Below is a brief listing of higher taxonomic categories of living fishes, in phylogenetic order. This list is meant as an introduction to majo of living fishes as they will be discussed in the initial two sections of this book. Many intermediate taxonomic levels, such as infraclasses, subdivisions, and series, a presented here; they will be detailed when the actual groups are discussed in Part III. Only a few representatives of interesting or diverse groups are listed. Taxa and illustrations from Nelson (2006). Superorder Protacanthopterygii pickerels, smelts, salmons Subphylum Cephalochordata lancelets Subdivision Euteleostei advanced bony fishes Subphylum Craniata Superclass Myxinomorphi Class Myxini hagfishes Superclass Petromyzontomorphi Class Petromyzontida lampreys Superclass Gnathostomata jawed fishes Class Chondrichthyes cartilaginous fishes Subclass Elasmobranchii sharklike fishes Subclass Holocephali chimaeras Grade Teleostomi bony fishes Class Sarcopterygii lobe-finned fishes Subclass Coelacanthimorpha coelacanths Subclass Dipnoi lungfishes [Order Esociformes pikes, mudminnows] b Superorder Stenopterygii bristlemouths, marine hatchetfishes, dragonfishes Superorder Ateleopodomorpha jellynose fishes Superorder Cyclosquamata greeneyes, lizardfishes Superorder Scopelomorpha lanternfishes Superorder Lampriomorpha opahs, oarfishes Superorder Polymixiomorpha beardfishes Superorder Paracanthopterygii troutperches, cods, toadfishes, anglerfishes Superorder Acanthopterygii spiny rayed fishes: mullets, silversides, killifishes, squirrelfishes, sticklebacks, scorpionfishes, basses, perches, tunas, flatfishes, pufferfishes, and many others Class Actinopterygii ray-finned fishes Subclass Cladistia bichirs Subclass Chondrostei paddlefishes, sturgeons Subclass Neopterygii modern bony fishes, including gars and bowfin a Division Teleostei Subdivision Osteoglossomorpha bonytongues Subdivision Elopomorpha tarpons, bonefishes, eels Subdivision Otocephala Superorder Clupeomorpha herrings a Gars and Bowfin are sometimes separated out as holosteans, a sister group to the teleosts (see Chapter 13). b The esociform pikes and mudminnows are not as yet assigned to a superorder (see Chapter 14). Superorder Ostariophysi minnows, suckers, characins, loaches, catfishes Subdivision Euteleostei advanced bony fishes Superorder Protacanthopterygii pickerels, smelts, salmons [Order Esociformes pikes, mudminnows] b Superorder Stenopterygii bristlemouths, marine hatchetfishes, dragonfishes Superorder Ateleopodomorpha jellynose fishes Superorder Cyclosquamata greeneyes, lizardfishes Superorder Scopelomorpha lanternfishes Superorder Lampriomorpha opahs, oarfishes Superorder Polymixiomorpha beardfishes Superorder Paracanthopterygii troutperches, cods, toadfishes, anglerfishes Superorder Acanthopterygii spiny rayed fishes: mullets, silversides, killifishes, squirrelfishes, sticklebacks, scorpionfishes, basses, perches, tunas, flatfishes, pufferfishes, and many others a Gars and Bowfin are sometimes separated out as holosteans, a sister group to the teleosts (see Chapter 13). b The esociform pikes and mudminnows are not as yet assigned to a superorder (see Chapter 14). 7

Periods of occurrence of major jawed (gnathostome) fish taxa based on the fossil record

Neognathi The remaining Euteleosts, all 336 families and 17,000+ species, are collectively placed in the unranked taxonomic category Neognathi, which consists of the order Esociformes (10 species) and the Neoteleostei, the latter also an unranked taxonomic level. The esociform pikes and mudminnows are viewed as the sister group of all higher euteleosts because they share a distinct type of tooth attachment and a skeleton made up of acellular bone (Johnson & Patterson 1996). Consensus on this arrangement is by no means universal, with some authors contending that esociforms are protacanthopterygians and a sister taxon to salmoniforms (see Nelson 2006 for a review). 9

Phylogeny of the major taxa of Actinopterygii (based on Nelson) Cellular/Acellular Bone Kranenbarg, S. et al. J Exp Biol 2005;208:3493-3502

Figure 14.19 The esocid Muskellunge, Esox masquinongy, one of the largest freshwater predators in North America. From Scott and Crossman (1973). Stenopterygii Ateleopodomorpha Cyclosquamata Scopelomorpha Lampridiomorpha Polymixiomorpha Paracanthopterygii Acanthopterygii Acanthomorpha Ctenosquamata Eurypterygii Neoteleostei Figure 14.20 Phylogenetic relationships among formally recognized, advanced teleostean superorders above the level of the Protacanthopterygii. Along the diagonal line of the cladogram are commonly used designations that do not have formal rank but that are generally distinguishable by shared derived traits as discussed in the following accounts. After Nelson (2006). 11

Esociformes Pikes (2 families, 10 spp) Redfin pickerel Esox americanus Gmelin 1788 Esociformes considered SISTER to all higher Euteleosts 12

Esociformes Esocidae (pikes, pickerels) Esox, 5 spp. Umbridae (mudminnows) Umbra, Novumbra, Dallia Maxillary is toothless (but in gape of mouth) Median fins posteriorly placed No adipose fin

Esociformes Freshwater Northern Hemisphere Esox lucius Northern Pike - most widespread natural distribution of any completely freshwater fish Esox lucius and Esox masquinongy Introduced into GA, not established sluggish, fluvial environments Lie in wait, sight-feeding predators; GA: 2 Esox species, Chain & Redfin pickerels Mudminnows Nocturnal, forage on inverts Holarctic GA: Umbra pygmaea, Eastern mudminnow

Amur pike Esox reicherti

Superorder Protacanthopterygii 3 Orders - Argentiniformes Marine smelts (ca. 6 families,57 genera 202 spp) Osmeriformes Freshwater smelts (2 super families) Salmoniformes Trouts, (1 family, 66 spp) Osmeriformes freshwater smelts 2 superfamilies Osmeroidea 11 genera 31 spp Galaxioidea 3 Families, 22 genera era and 88 spp. Binocular fish Winteria telescopa Bauer 1901 Capelin Mallotus villosus (Müller 1777) Chinook Oncorhynchus tshawytscha (Walbaum 1792)

Argentiniformes marine smelts 6 Families, 57 Genera, 202 spp. Two suborders Argentinoidei Argentines, herring smelts Opisthoproctus, Barreleye Greater Silver Smelt Argentina silus Alepecephaloidei Macropinna microstoma 500-1000m Tubeshoulders

Siphonophores Apolemia

Osmeriformes freshwater smelts 2 Superfamilies Osmeroidea 11 genera and 31 spp Galaxioidea 3 Families 22 genera 88 spp Osmeridae marine, anadromous, coastal freshwater Northern Hemisphere Arctic, Atlantic, Pacific TAVERNE AND FILLEUL: CRETACEOUS SALMONIFORM FISH 933 Many diadromous Maxillary is usually toothless (but in gape of mouth) TEXT-FIG. 4. Spaniodon latus (Agassiz, 1844); skull in lateral view. Restoration from specimens MNHN 1939-7-14, 69, 133, 164, 166, 174g, 186d, 188, 195g and 1946-18-130, 137, 153, 1244, 1259, 1271, 1352, 1378. Adipose fin, or not!

Terms for Migratory Fishes Greek dromos: running, dia: through Migrate between the sea and freshwater - Anadromous: running up (ana = up) - Catadromous: running down (kata = down) - Amphidromous: running around - Potamodromous: running within rivers Many diadromous

Osmeriformes Freshwater smelts Osmeridae - Northern Hemisphere about 31 spp Distinct Population Segment (DPS): A DPS, or a distinct population segment, is a vertebrate or endangered under the population This evening or group we were of populations visited by Comowool that is discrete the from other populations of the species and significant Clatsop Chief in relation and 12 to men the women entire species. and children The ESA of his provides for listing species, that the petition subspecies, presents substantial or distinct nation population... The Chief segments and his of party vertebrate had brought species. for indicating that the petitioned action sail a Sea Otter skin, some hats, stergeon and a species of small fish which now begin to run, and are taken in great quantities in the Columbia R. about 40 ensure that the status review is miles above us by means of skimming or scooping complete and based upon the best available scientific and commercial nets... I find them best when cooked in Indian stile, information, we solicit information which is by roasting a number of them together on a wooden spit without any previous preparation whatever. They are so fat they require no additional sauce, and I think them superior to any fish I ever taste, even more delicate and luscious than the white EULACHON (Columbia River Smelt) Thaleichthys pacificus fish of the lakes which hae heretofore formed my standaart of excellence among the fishes. From the Journals of Captain Meriwether Lewis and William Clark (February 25, 1806) Ascends rivers in Northern Pacific to spawn in spring Larvae swept downstream to estuary, ocean DID YOU KNOW: When dried and fitted with a wick a eulachon can be burned like a candle Commercial fishery Published in 2008 NMFS, received a petition to list populations of Pacific eulachon (Thaleichthys pacificus) in Washington, Oregon, and California as a threatened Endangered Species Act (ESA). We find scientific and commercial information may be warranted. Accordingly, we will initiate a status review of the species. To March 2010, Southern DPS listed as Threatened under ESA regarding the population structure and status of Pacific eulachon throughout their range in Alaska, British Columbia, Washington, Oregon, and California

Osmeriformes Freshwater smelts Osmeridae - Northern Hemisphere about 31 spp Capelin Mallotus villosus 16-18cm

Osmeriformes Freshwater smelts Osmeridae - Northern Hemisphere about 31 spp over 300 pyloric caeca Plecoglossus altivelis Ayu

Osmeriformes Superfamily Galaxioidea usually no pyloric caeca, no supramaxillae Southern Hemisphere (S AM, Aus, NZ, S Africa) Two Families Retropinnidae (New Zealand Smelts and Graylings ) 3 genera 5 species and Galaxiidae (galaxiids) 8 genera 52 species Cool temperate - freshwater, brackish, diadromous Dominant component of freshwater fish fauna many species displaced/imperilled New Zealand galaxiid, Prototroctes oxyrhinchus t EXTINCT

Osmeriformes Freshwater smelts Galaxioidea Galaxiidae about 52 spp giant kokopu, Galaxias argenteus (Gmelin 1789) [Systema Naturae Linné] to 58 cm, quiet pools, cover, predaceous, first Galaxiid to be described IUCN Threatened

A galaxiid in a different habitat altogether Lepidogalaxiidae, Lepidogalaxias salamandroides Osmeriformes Freshwater smelts Galaxioidea Galaxiidae about 52 spp Australian Salamanderfish small (50 mm), invertivore posterior dorsal & anal fins flexible neck! Tim M. Berra (Tree of Life Web Project) ephemeral, low-ph ponds aestivates in damp sand, 2-60 cm deep

Salmoniformes trouts, 1 Family Salmonidae 11 genera, 66 spp. 3 subfamilies Coregoninae 3 genera and 32 spp Whitefish and Ciscoes Thymallinae 1 genus 4-5 spp Graylings Salmoninae 7 genera, about 30 spp. Salmon, trouts, charrs Coregoninae < 16 Dorsal Rays, < 110 scales in lateral line no teeth on maxilla Thymallinae > 17 Dorsal Rays, teeth on maxilla Salmoninae < 16 Dorsal Rays, > 110 scales in lateral line, teeth on maxilla

Salmoniformes trouts, 1 Family Salmonidae 11 genera, 66 spp. Coregoninae 3 genera and 32 spp Whitefish and Ciscoes Thymallinae 1 genus 4-5 spp Graylings Salmoninae 7 genera, about 30 spp. Salmon, trouts, charrs A 30 cm long specimen of Eosalmo driftwoodensis from the Eocene of Smithers, B.C. This is the earliest known salmon. University of Alberta Collections. Smilodonichthyes rastrosus Oncorhynchus rastrosus Miocene to Pleistocene, large to 8, 100+ gill rakers, probably planktivorous

Salmoninae 7 genera, about 30 spp. Salmon, trouts, charrs Brachymystax lenok Acantholingua ohridana one lake in Albania Salmothymus 3 spp S. obtusirostris Adriatic trout Hucho 3 spp Salveninus (charrs) 11 spp Salmo (Atlantic trouts) 5 spp Oncorhynchus (Pacific salmon and Pacific trouts) 11 spp

Hucho taimen Taimen Largest salmonid, record size of 210 cm, 231 lbs. Caspian and Artic drainages in Eurasia, some Pacific drainages in Mongolia and Russia (Amur)

charrs, Salvelinus brook, lake, bull trout; Arctic charr, Dolly Varden,

Salmo salar, Atlantic salmon Salmo spp., Atlanto-European salmons Salmo trutta, brown trout

Oncorhynchus (Pacific salmon and Pacific trouts) 11 spp The scientific names of Pacific salmon and rainbow trout were derived from common names used by native peoples of Kamchatka. 1740-German naturalist George Steller penned manuscript describing 5 salmon and rainbow trout, recording phonetic translations into German, but he died in 1746 before it could be published. It was translated into Russian in 1755, then into English, then into French in 1768. Walbaum in 1792 published the the names originally recorded by Steller, as latinized scientific names, tshawytscha (Chinook), gorbuscha (pink), kisutch (coho), keta (chum) nerka (sockeye) and mykiss (rainbow trout) from Trout and Salmon of North America Behnke and Tomelerri.

Oncorhynchus tshawytscha Chinook Salmon Oncorhynchus nerka Sockeye salmon Oncorhynchus spp, Pacific salmons

Rainbow Cutthroats Oncorhynchus spp, Pacific trouts, from Behnke and Tomelleri

Oncorhynchus chrysogaster Mexican golden trout Steelheads Oncorhynchus gilae apache Apache trout Oncorhynchus spp, Pacific trouts, from Behnke and Tomelleri

Oncorhynchus spp, Cutthroat tree, from Behnke and Tomelleri

Pacific salmon are anadromous, migrating upstream to spawn in natal streams (they were hatched in). They may spend 2 to 5 years feeding in the ocean, after out-migrating from freshwater.

kype elongated jaws in spawning male salmon Pink Salmon Sockeye Salmon Redband Steelhead Oncorhynchus spp, Pacific trouts, from Behnke and Tomelleri

NATAL STREAM

THE OLFACTORY HYPOTHESIS Coho in Lake Michigan: Hatchery reared, imprinted as smolts w/ 2 alcohols; Released into lake. 18 months later, dripped alcohol into 2 streams, not into 17 others. 95% of experimental fish returned to experimental streams; Others entered streams at random.

SALMON IN THE ECOSYSTEM

Large runs of Salmon provide pulsed inputs of prey and nutrients to coastal and inland streams. Carcasses? * transfers nutrients from Ocean to the forests riparian trees grow faster, perhaps positive feedback? Time for Sitka Spruce to reach a size to form a pool (falling in) drops from >300 to <100 yrs in se AK *Nutrients subsidize aquatic foodwebs. > 90% N in AK brown bears comes from salmon, up to 1/3 of N in valley bottom forests swam up the river and streams as a fish!

It is now widely recognized that Pacific salmon in the Pacific Northwest (except Alaska) are faring poorly Nehlson 1994 What is a stock? 16 runs protected as Endangered Species

Table 1 shows the peak salmon cannery packs in major production areas south of the Fraser River. Area Peak Year Cases Kg of Salmon (Millions) Sacramento River 1882 200,000 4.30 Columbia River 1895 634,696 13.70 Fraser River 1901 998-913 21.50 Willapa Harbor 1902 39,492.81 Grays Harbor 1911 75,941 1.62 Coastal Oregon Rivers 1911 138,146 2.30 Klamath River 1912 18,000.39 Puget Sound 1913 2,583,463 558.00 Coastal Washington Rivers 1915 31,735 0.67

THE 4 H S CAUSING SALMON DECLINE 1. HYDROPOWER 2. HARVEST 3. HABITAT DESTRUCTION 4. HATCHERIES 5. (HISTORY)