WFB 232 Ichthyology Ichthyology – Introduction to Fishes Vertebrate classes: Agnatha Chondrichthyes Osteichthyes Amphibia Reptilia Aves Mammalia # species 85 850 23,000 2,600 6,500 8,600 4,100 % of vertebrates <1% 2% > 50% of all vertebrates are fishes 50% 6% 14% 19% 9% physiological and sensory modalities in fish that are not present in other taxa osmoregulation buoyancy control respiration (air and water) mechanosensory – lateral line electroreception electric generation light production Characteristics and constraints of the aquatic environment salinity - must deal constantly with osmoregulation - into or out of body temperature - water has high thermal capacity, 4x that of air motion - from still water, with little oxygen exchange, to torrential currents gas saturation - water contains (relative to air) very little oxygen; methods for withdrawal must be specialized; too little, fish must achieve oxygen elsewhere; too much results in gas bubble diseases pressure - increases 1 atm for every 33’ of depth (= 10 m) consequence for buoyancy and gas bladders few fish are neutrally buoyant – use air or fat to compensate fish need much less skeletal support than terrestrial animals, due to support surrounding body viscosity - much higher effort required to swim in water than air fastest swimming speed (tuna) - 21m/s in short bursts; killer whale does 15m/s little effort required to prevent sinking light - attenuates rapidly with depth; thus deep-sea fish may create own light sound - propagates rapidly in water (400x as well as in air) provides medium of communication by vibration, but with little directionality Distribution of aquatic habitats Salt water Fresh water oceans 97.10% ice, snow salt lakes 0.01 groundwater lakes atmosphere rivers Total 97.11 2.2% 0.6 0.01 0.001 0.0001 2.81 Fish distribution in habitat: freshwater saltwater move between (anadromous, catadromous) 41% 58% 1% WFB 232 Ichthyology Taxonomy Objectives: Learn how species are defined and named; understand how taxonomic nomenclature is used Begin to know the classes, orders, and representative families and species in each order of fishes Learn to ‘read’ taxonomic descriptions Recognize primitive (older) versus advanced (recent) traits in fishes Classification = hierarchical grouping of organisms (i.e. a process, an operation) Systematics = the study of relationships Taxonomy = the science of biological nomenclature (formal rules for use) Taxonomic hierarchy: (note standardized endings) Phylum: Chordata Subphylum: Vertebrata Superclass: Gnathostomata Class: Actinopterygii (formerly Osteichthyes) Order: Cypriniformes Suborder: Cyprinoides Family: Cyprinidae Subfamily: Cyprininae Genus: Cyprinus Species: carpio standardized endings (well-established in ichthyology) Order: -formes Suborder: -oides Family: -idae Subfamily: -inae Tribe: -ini Only genus and species are underlined or italicized with only the generic term capitalized. phylogenetic systematics – objective is to make the classifications non-arbitrary, and informative about evolutionary relationships cladograms: each branch represents monophyletic group - all from common ancestor if polyphyletic, then subsequent categories based on false assumption all members share one or more derived characters problem is to determine whether a given character is homologous or convergent plesiomorphies – primitive character states apomorphies – advanced character states autoapomorphies – specialization unique to one taxon synapomorphies – shared specializations WFB 232 Ichthyology Taxonomic relationships defined on the basis of morphometrics - usually measurements as ratio against SL, to account for absolute size meristics - counts, with ranges - may vary with environment anatomical traits - shape, presence/absence of structures color patterns - highly variable, change with maturity, subjective, fade in preservative karyotypes - changes in number and shape of chromosomes biochemical methods - look directly at genetic material Authorship of scientific names In original genus: Amia calva Linnaeus – usually followed by date In new genus: Lepisosteus osseus (Linnaeus) right of priority – given to first full written description, usually with type specimen from a type locality the description makes it a nominal species original species name must not be changed, even if genus is changed - unless previously used for another animal (nomen praeoccupatum) - if previously described, later description becomes junior synonym hybrids denoted with an x - Rutilus rutilus x Abramis brama Nomenclature (International Code of Zoological Nomenclature) pronounciation: ch = k (e.g., ichthyology) – there is no ‘k’ in Latin initial c = s (e.g., Cyprinidae) genus and species always italicized; genus is always capitalized, species name is not capitalized Neogobius melanostomus genus and subspecies can be abbreviated after they have been mentioned once in a text N. melanostomus N. m. caspia subgenus is usually in parentheses Dreissena (Pontodreissena) polymorpha Characteristics of the classes of extant fishes Character skeleton paired fins gill arches gill arches nostril(s) jaws fins with spines pectoral fins pelvic fins tail scales mouth Primitive cartilaginous absent absent not joined to brain case 1, median absent absent (trout) horiz. base low on body (minnows) far back on belly (pikes) heterocercal cycloid (herring) front of head (trout) Advanced ossified (calcified bone) - may be lost present present firmly joined to brain case paired present present (perches) - but may be lost (killifish) vert. base high on body (basses) forward, attached to pectoral girdle (sculpins) symmetrical ctenoid (sunfish) - absence is specialization up- or down-turned (suckers, killifish) WFB 232 Ichthyology Taxonomy of Fishes Kingdom Animalia Phylum Chordata Subphylum Vertebrata SUPERCLASS AGNATHA Class Myxini Order Myxiniformes (hagfishes) Class Cephalaspidomorphi Order Petromyzontiformes (lamprey) SUPERCLASS GNATHOSTOMATA Class (Placodermi) Class (Acanthodii) Class Chondrichthyes Subclass Holocephali Order Chimaeriformes (chimaeras) Subclass Elasmobranchii 9 orders (sharks, rays, skates) Class Sarcopterygii Subclass Coelacanthimorpha Order Coelacanthiformes (coelacanths) Subclass Dipnoi Order Ceratodontiformes (Australian lungfish) Order Lepidosireniformes (S. American, African lungfish) Class Actinopterygii - (rayfins, higher bony fishes) Subclass Chondrostei Order Polypteriformes (birchirs, reedfish) Order Acipenseriformes (sturgeons, paddlefishes) Subclass Neopterygii Order Semionotoformes (Lepisosteiformes) (gars) Order Amiiformes (bowfin) Division Teleostei 35-38 orders (modern body fishes) WFB 232 Ichthyology Example of a taxonomic description (adapted from Berg, 1949) Round goby, Neogobius melanostomus (Pallas, 1811). D1 VI (V-VII); D2 I + 14-16 (13-16); A I + 11-13 (11-14); P 18-19 (17-20). [ = the anterior dorsal fin has 5-7 spines, usually 6, and the posterior dorsal fin has one spine and 13-16 soft rays. The anal fin has one spine and 11-14 soft rays, and the pectoral fins have 17-20 soft rays.] Scaled on the parietal region, nape, back (all), throat (all or most), abdomen, pectoral fin peduncles, and one quarter of the gill covers. Scales on the middle and anterior nape are cycloid. Head is as wide as or wider than deep; depth is 0.9-1.2 times the width. Head length 4.2-4.5 of total body length. Angle of the jaw below the anterior quarter of the eye. Lower jaw not prominent. Snout 1.1-1.4 times the orbit diameter. Upper lip narrows slightly to the rear. Usually 6, rarely 7, transverse suborbital series of pit organs. Ventral fins reach or almost reach the vent. Pelvic disk is 0.6-0.8 times the abdomen length. If present, the anterior membrane width is very shallow, with rounded, lateral lobes. Caudal peduncle depth is about two-thirds its length. Lacks a gas bladder and chemoreceptors. WFB 232 Ichthyology Fish Anatomy Objectives: become acquainted with the general external and internal structures of fishes, and how they vary functionally and taxonomically become familiar with parts of fish and associated terminology learn to recognize fishes by similarities and differences in structure External anatomy Head mouth mouth position – superior, terminal, sub-terminal, inferior teeth types and locations; mostly on mandible (lower jaw), premaxilla and maxilla (upper jaw) – Bond Fig. 2-20 generally protrusible in derived bony fishes frenum - bridge of skin binding lips to snout or chin in non-protrusible jaw additional (or seasonal) structures include barbels, tubercules, kype, cirri sensory pores nares Body shapes – which taxa have each, what are the advantages of each type? fusiform sagittaform anguilliform compressiform depressiform filiform Fins – what are each used for? how does their presence or location vary among taxa? paired fins: pelvic – some modified to disk (gobies) or claspers (sharks) placement may be abdominal, subabdominal, jugular, mental (under chin) pectoral single fins: dorsal - one or two; rarely three; may be modified to disk (medial) anal – may be modified to intromittent organ (gonopodium) caudal – rounded, lobed, truncate, emarginated, forked; homocercal or heterocercal adipose – in only a few orders peduncles (base of fin or tail where they attach to body) supporting structures: ceratotrichia – cartilaginous elements in elasmobranch fins lepidotrichia – rays (double, segmented elements) sometimes modified to spines (ossified, single) Skin and scales – which are more advanced, or primitive? presence, absence, or partial covering of scales functions of scales types of scales: placoid, ganoid, cycloid, ctenoid, scutes skin pigmentation: melanophores WFB 232 Ichthyology Internal anatomy Internal organs GI tract (stomach, intestine, pyloric caecae), reproductive organs, kidneys, liver, gall bladder, spleen gas bladder may be present or absent, connected or not to other organs physoclistous – closed gas bladder physostomus – gas bladder has open connected to esophagus Body musculature hypaxial muscles epaxial muscles myomeres Osteology Skull premaxilla, maxilla, dentary, palatine may all have teeth attached operculum: opercle, subopercle, preopercle, interopercle otoliths – ear bones branchial arches - usually 5 pairs - bear gill rakers on inner surface fifth arch may be modified to pharyngeal teeth breathing apparatus (see diagrams in Bond) Vertebral column Vertebrae composed of centrum, with neural spine, neural arch, neural canal zygopophysis, basapophysis – small locking projections add rigidity, connection with ribs hemal spine, hemal arch (only near tail), hemal canal dorsal ribs, or epipleurals, project from pleurals Caudal skeleton urostyle – last vertebra, modified into plate hypurals - modified from hemal arch, connected to end of vertebral column and caudal elements epurals – remnant from hemal spine, one or more free bones above hypurals Appendicular skeleton pterygiophores articulate with fin rays pectoral girdle – cleithrum, scapula, coracoid bones pelvic girdle – simple system to support fins WFB 232 Ichthyology Orders of fishes, with selected families Class/subclass Myxini Number of Order Families Myxiniformes 1 Cephalaspidomorphi Petromyzontiformes Chondrichythes Holocephali Elasmobranchii Chimaeriformes Heterodontiformes Orectolobiformes Carchiniformes Lamniformes Hexanchiformes Squaliformes Squantiniformes Pristiophoriormes Rajiiformes Sarcopterygii Coelocanthimorpha Coelacanthiformes Dipnoi Ceratodontiformes Lepidosireniformes Actinopterygii Chondrostei Polypteriformes Acipenseriformes Neopterygii Semionotoformes Amiiformes Div. Teleostei Representative families Myxinidae Common names hagfish 1 Petromyzontidae lamprey 3 1 7 7 7 2 3 1 1 9 Chimaeridae Heterodontidae Rhincodontidae Cetorhinidae Hexanchidae Squalidae Squantinidae Pristiophoridae Rajidae chimaeras bullhead sharks whale sharks ground sharks basking sharks cow sharks dogfish angel sharks saw sharks skates, rays 1 1 2 Latimeriidae Ceratodontidae Lepidosirenidae coelacanth 1 Australian lungfish 1 S. Am., African lungfish 5 1 2 1 1 Polypteridae Acipenseridae Lepisosteidae Amiidae birchirs, reedfish sturgeons, paddlefish gars bowfin Hiodontidae Megalopidae Albulidae Anguillidae mooneye 217 tarpon 8 bonefish 29 eels 738 swallowers, gulpers 26 herrings 357 milkfish 35 carp, shiners 2,662 suckers hatchetfish 1,343 catfish 2,405 knifefish 62 pikes 5 mudminnows 5 smelt 236 salmon, trout, ciscoes 66 whitefish, chubs lightfish, dragonfish 321 jellynose fish 12 lizardfish 219 lanternfish 241 Osteoglossiformes 6 Elopiformes 2 Albuliformes 3 Anguilliformes 19 Saccopharyngiformes 4 Clupeiformes 4 Gonorynchiformes 4 Cypriniformes 6 Characiformes Siluriformes Gymnotiformes Esociformes 10 31 6 2 Osmeriformes Salmoniformes 13 1 Stomiiformes Ateleopodiformes Aulopiformes Myctophiformes 9 1 12 2 Clupeidae Cyprinidae Catostomidae Characidae Ictaluridae Esocidae Umbridae Osmeridae Salmonidae Ateleopodidae # species in order 43 41 31 8 31 208 16 5 74 12 5 456 10 26 5 1 WFB 232 Ichthyology Lampridiformes Polymixiiformes Percopsiformes Ophidiiformes Gadiformes Batrachoidiformes Lophiiformes 7 1 3 4 12 17 16 Mugiliformes 1 Atheriniformes 5 Beloniformes 5 Cyprinodontiformes 13 Stephanoberyciformes 9 Beryciformes 14 Zeiformes 6 Gasterosteiformes 11 Synbranchiformes Scorpaeniformes Perciformes Pleuronectiformes Tetraodontiformes Totals: 5 classes 57 orders 3 20 128 6 9 ribbonfish, oarfish 19 beardfish 5 trout-perch 9 cusk-eels 355 Gadidae cod, hake 482 Batrachoididae toadfish 69 Lophidae anglerfish 297 Ogvocephalidae batfish Mugilidae mullets 80 silversides, grunion 285 needlefish, flying fish 191 Cyprinodontidae livebearers 807 Poeciliidae guppies whalefish 86 squirrelfishes 123 John Dories 39 Gasterosteidae sticklebacks 257 Pegasidae seamoths Syngnathidae pipefish, seahorses Indostomidae I. paradoxus Synbranchidae swamp eels 87 Cottidae scorpionfish, sculpin1,271 Dactylopteridae flying gunards Percichthyidae temperate bass 9,293 Centrarchidae sunfish Percidae perch, bass Sciaenidae drum Mullidae goatfishes Cichlidae cichlids Mugilidae mullets Gobiidae gobies (also: bluefishes, remoras, blennies, mackerels, dolphins, snappers, tunas, swordfish) Pleuronectidae flounder, flatfishes 570 Balistidae triggerfishes 339 Ostraciidae cowfish, boxfish Tetraodontidae puffers Molidae molas (ocean sunfish) Polymixiidae Percopsidae 478 families ~26,000 species WFB 232 Ichthyology Swimming Objectives Understand the physical characteristics of water that affect the ability to swim Understand the various modes of fish propulsion, factors that affect their efficiency, and representative species that use them physical properties of water density – 830 x more than air viscosity – 70 x more than air boundary layer turbulence Reynolds number (Re) (dimensionless variable) Re = LVr/m L = length of object V = velocity of object r = density of fluid m = viscosity of fluid Examples of Re: animal whale tuna copepod sea urchin sperm speed 10 m/s 10 m/s 20 cm/s 0.2 mm/s Re 300,000,000 30,000,000 300 0.03 unavoidable issues when swimming: flow changes to turbulent at Re ~ 2,000 boundary layer changes to turbulent as Re goes from 5x105-5x106 turbulent flow is a consequence of decreasing viscosity increasing density of liquid increasing speed increasing length for efficient swimming avoid separation of boundary layer from surface maximize laminar flow in boundary layer minimize turbulent flow in wake solutions: streamline body (tapering) aspect ratio of about 0.25 max. thickness of body 1/3 back drag reduction - keep body rigid slime layer to reduce frictional drag rough surface (cteni) keeps boundary layer attached WFB 232 Ichthyology Swimming modes “kick and glide” active - sustained for hours or days burst - only for up to 30 secs large fishes have greater difference between burst and active than small fishes active swimming accomplished using red muscle along sides of fish - high myoglobin and mitochondrial enzymes burst swimming with white muscle - great contractile speeds, low endurance Body/caudal fin propulsion Medial/paired fin propulsion non-swimming locomotion burrowing wriggling ‘walking’ push-and-hold walking on bottom vics leaping gliding hitchhiking passive drift jet propulsion