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by: Percy Wintheiser


Percy Wintheiser
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This 14 page Class Notes was uploaded by Percy Wintheiser on Wednesday September 9, 2015. The Class Notes belongs to FISH 311 at University of Washington taught by Staff in Fall. Since its upload, it has received 48 views. For similar materials see /class/192253/fish-311-university-of-washington in Aquatic And Fishery Sciences at University of Washington.

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Date Created: 09/09/15
BIOLOGY OF FISHES FISH 311 BIODIVERSITY TAXONOMY SYSTEMATICS AND CLASSIFICATION NUMBERS AND KINDS OF FISHES DIVERSITY THROUGH TIME MAJOR GROUPS OF FISHES General topics 1 quotTime to Revive Systematicsquot 2 What is taxonomy and how does it differ from systematics 3 Branching diagrams and classi cations 4 The importance of understanding phylogeny kl The numbers and kinds of living organisms vertebrates and shes 6 An ecological classi cation of shes 7 The groups of shes we Will study in FISH 311 1 quotTIME TO REVIVE SYSTEMATICSquot Start off with an editorial by E O Wilson the famous Harvard entomologist evolutionary biologist and behaviorist that appeared in SCIENCE back in December 1985 it may seem old to you but still as signi cant today as it was back then a plea to quotRevive Systematicsquot The important points are summarized as follows the full editorial makes up page 3 of these notes 1 Systematics is the study of biological diversity 2 At the present time we do not even know to the nearest order of magnitude how many species there are in the world 3 Approximately 17 million have been described but these numbers are far below the actual diversity 4 Recent studies in rain forests and other major habitats indicate the presence of as many as 30 million kinds of insects alone 5 Because of the largely unknown nature of diversity systematics remains a tremendous source of discoveries and new ideas in biology 6 Much of the research in systematics has economic and medical importance 1 The world supply of trained systematists is no where near the number required to research even a small part of unknown or poorly known aspects of biodiversity 8 Government support for systematic research in this country is so low that less than one percent of the known species of organisms are under active investigation Since 1985 with the help of strong advocates like Wilson some improvements have been made but there39s still a long way to go to even begin to tackle the enormous lack of basic information about the plants and animals around us The US National Science Foundation NSF is the only signi cant granting agency in this eld It now has a number of programs directed toward biodiversity 1 quotBiotic Surveys and Inventoriesquot BSI 2 quotPartnerships for Enhancing Expertise in Taxonomyquot PEET 3 quotBiocomplexityquot 4 quotBiological Research Collectionsquot 5 quotResearch Experiences for Undergraduatesquot REU But even with all this very little new monies have been directed toward basic systematic studies or toward the support of natural history collections The problem is of course much more crucial now than ever before because of global destruction of natural habitat and consequent Widespread extinction of species 13 December 1985 Volume 230 Number 4731 E 39Time to Revive Systematics Systematics the study of biological diversity is sometimes portrayed as the mere classi cation of organisms but in fact its range and challenge are among the greatest in biology At the present time we do not even know to the nearest order of magnitude how many species there are in the world Approximately 17 million have been described since Linnaeus including about 250000 owering plants 47000 vertebrates and according to one meticulous estimate published earlier this year 751012 insects But these numbers are far below the actual diversity Recent studies in rain forests and other major habitats indicate the presence of as many as 30 million kinds of insects alone The magnitude of biological diversity has many rami cations of general interest For example if there are really 30 or 40 million animal species why didn t just a thousand evolve or a billion It is not known to what extent diversity is controlled by physical properties of the planet as opposed to the mechanics of evolution itself Nor do we know to what degree species numbers can be raised or lowered arti cially without destabilizing local ecosystems In another dimension biologists have only begun to assess the complexity and potential of each species individually Every species is the terminus of an ancient lineage that has been hammered and shaped into its present form by a complex interplay of genetic recombination and natural selection In a purely technical sense the resulting genome is richer in con tent than a Caravaggio painting a Bach fugue or any other great work of art The billion bits of genetic information in the house mouse for example if transformed into an equivalence of printed English text would just about ll all editions of the Encyclopaedia Britannica published since 1768 Because of the largely unknown nature of diversity systematics remains a fountainhead of discoveries and new ideas in biology If a biologist is well trained in the classi cation of the organisms encountered the known facts of natural history are an open book and new phenomena come more quickly into focus The irony of the situation is that successful research then gets labeled as ecology physiology or almost anything else but its true source the study of diversity Much of the research also has economic and medical importance The discovery of new sources of biomass energy lumber pharmaceuticals and pollination complexes depend ultimately on taxonomic exploration Also the design of natural reserves critical to the preservation of diversity in tropical countries cannot be performed reliably without a thorough knowl edge of local faunas and floras The problem is intensi ed by the accelerat ing destruction of natural habitats and extinction of species At present the community of systematists is sadly inadequate to the immense task before it In North America about 4000 specialists most part time work on 3900 collections Probably no more than 1500 trained professional systematists in the world are competent to deal with tropical organisms To cite one striking example there are exactly two such persons quali ed to deal with termites which are among the principal insect pests and soil movers of the world In scal 1985 the National Museum of Natural History our largest institution of basic research spent 128 million to support the activities of 85 scientists engaged partly or wholly in systematics The Program in Systematic Biology of the National Science Foundation the principal funder of independent projects granted 12 million Other programs in the NSF and Department of Interior provided 138 million for support of museum services and other activities related to systematics At this level which re ects a low priority worldwide less than 1 percent of the species of organisms are under active investigation Systematics deserves more cultivation and the attention of our brightest minds It is in a position to yield increasing returns to scale with a variety of bene ts for both science and society EDWARD 0 WILSON Museum of Comparative Zoology Harvard University Cambridge Massachusetts 02138 2 DEFINITIONS WHAT IS TAXONOMY AND HOW DOES IT DIFFER FROM SYSTEMATICS Taxonomy The discovery recognition de nition and naming of groups of organisms Systematics The study of biological diversity or more speci cally the ordering of the diversity of nature through construction of a classi cation that can serve as a quotgeneral reference system quot The task of systematics and the task of the creator of classi cations is to nd the best possible quotgeneral reference systemquot But nding the best possible quotreference systemquot is not so easy and it turns out that there are several approaches or ways to go about performing this task But whatever the methodology the nal products of systematic research are two fold l Branching diagrams 2 Classi cations Branching diagrams sometimes called evolutionary or phylogenetic trees or quotcladogramsquot from the Greek quotcladquot or quotkladosquot meaning to branch are simply graphic views of the sequence of evolutionary divergence of groups of organisms through time Along the horizontal axis they show the relative primitiveness of organisms along the vertical axis they show how groups of organisms have evolved from one another that is the pattern of branching through time BRANCHING DIAGRAM OF FISH AND TETRAPOD RELATIONSHIPS Hagfishes Lampreys Cartilaginous shes Bonyfishes Latimeria Lungfishes Tetrapods Classi cations on the other hand are not pictorial but rather concise lists of organisms grouped or ranked according to the pattern of branching seen in the branching diagram A CLASSIFICATION OF FISHES AND TERRESTRIAL VERTEBRATES lnfraphylum MYXINI Hagfishes lnfraphylum MYOPTERYGII Superclass CEPHALASPIDOMORPHI Lampreys Superclass GNATHOSTOMATA Class ELASMOBRANCHIOMORPHI Cartilaginous fishes Class TELEOSTOMI Subclass ACTINOPTERYGII Bony fishes Subclass SARCOPTERYGlI lnfraclass CROSSOPTERYGII Latimeria lnfraclass DIPNEUSTI Series DIPNOI Lungfishes Series TETRAPODA Terrestrial vertebrates You39ll notice in the examples given that the classi cation follows precisely the pattern of branching in the branching diagram and Vice versa the branching diagram can be constructed directly from the classi cation and the listing of organisms in the classi cation follows directly from the branching diagram You39ll notice also that the classi cation is strictly dichotomous groups branching in successive pairs Taxa ie species or other formal taxonomic groups of organisms within likepairs are termed sister groups for example we say that hag shes are the sister group of all other shes plus tetrapods sharks and their allies the cartilaginous shes are the sister group of a group containing the bony shes the coelacanths genus Latimeria the lung shes plus all tetrapods tetrapods are the sister group of lung shes etc In theory each branching point in the tree indicates a sistergroup pair of shared categorical rank but in practice in is nearly impossible to present relationships this way in a large taxon containing numerous subtaxa for example an order containing many families or a family containing many genera A case in point is the branching diagram of angler sh relationships shown on the next page 4 THE IMPORTANCE OF UNDERSTANDING PHYLOGENY You might think that knowing the evolutionary relationships among groups of organisms is interesting but beyond that phylogenies are not too important Do they in fact have a greater signi cance Do they have a practical use Just what good are they Why should we care about them and why should money be spent to support their construction Example reproductive strategies among deepsea ceratioid angler shes A lac Diceratiidae eirodidae Thaumatichthyidae Centrophrynidae Ceratiidae Gigantactinidae Linophrynidae k i k k A phylogenetic tree cladogram showing the presumed evolutionary relationships of deepsea angler shes Stars indicate taxa in which sexual parasitism has been found 5 THE NUMBERS AND KINDS OF ORGANISMS VERTEBRATES AND FISHES Let39s go back to E O Wilson39s estimate of 17 million known species of organisms and take a look at how the number of living shes stacks up against the numbers found in other groups Remember this 17 million is an estimate of that small part of world biodiversity that has been formally described in the scienti c literature Let39s look at a simple breakdown of plant and animal taxa taken from an article that appeared in the New York Times a While ago The Dominant Species Of the World staccterla There are 751000 known 4800 insect species versus a Fungi mere 4000 or so 69000 mammals There are 112000 species of Lepidoptera butter ies and moths 103000 species of Hymenoptera ams bees and wasps and 290000 beetles Source quotThe Diversny of Life quot Edward 0 Wilson Bemap Press of Harvard University Protozoa 30800 You can see that insects far outnumber other organisms with some 751000 described species A category called quotAll other Animalsquot which excludes protozoans Viruses fungi and bacteria comes next in order of abundance with about 281000 described species Plants excluding algae amount to 248400 Of these 281000 species of animals excluding insects how do living shes compare PRIMARY VERTEBRATE TAXA AND APPROXIMATE NUMBER OF LIVING SPECIES Mammals 5000 Birds 8600 Reptiles 8000 Amphibians 4700 Fishes 28400 about 519 of all vertebrates TOTAL 54700 APPROXIMATE NUMBER OF LIVING SPECIES OF THE MAJOR GROUPS OF FISHES Hagfishes Myxini 70 Lampreys Cephalaspidomorphi 38 Cartilaginous fishes Chondrichthyes 970 about 34 of all shes Bony fishes Actinopterygii 27300 about 962 of all shes Coelacanths Crossopterygii 2 Lungfishes Dipnoi 6 TOTAL 28400 These 28400 living species of shes are distributed among approximately 515 families and 4494 genera Of the approximate 970 living species of Chondrichthyes sharks rays and chimaeras more than half 534 or about 55 are rays Actinopterygians or bony shes constitute about 96 of all living sh species and all but 51 species or about 018 belong to a group called the Teleostei Of the 515 sh families the nine largest contain about 9300 or 33 of all species These families are Cyprinidae about 220 genera and 2420 species Gobiidae about 210 genera more than 1950 species Cichlidae about 112 genera and 1350 species Characidae about 165 genera and 962 species Loricariidae about 92 genera and 684 species Balitoridae about 59 genera and 590 species Serranidae about 64 genera and 475 species Labridae about 68 genera and 453 species and Scorpaenidae about 56 genera and 418 species Sixtyfour families are monotypic that is they contain only a single species Sixty seven families contain 100 species or more of which three contain more than 1000 The average number of species per family is 54 the median number is only 12 OK so we think we have about 28400 described species of living shes Once again you should realize that this gure is restricted not just to described species but also to those species that are recognized by the scienti c community There are lots of named species that aren39t recognized by anyone because they turn out to be the same as earlier described species we call them junior synonyms of recognized species About 300 new species are described each year Most of these are freshwater forms coming from poorly collected highdiversity tropical habitats like the Amazon and Congo basins but a signi cant number are coming from the deepsea The number of new descriptions of shes can be monitored rather easily but we have no good way to estimate the number of species that fall into synonymy each year The eventual number of living sh species is projected to be close to 32500 PANORAMA OF THE DOCUMENTED HISTORY OF VERTEBRATE EVOLUTION We ve already seen that there are more shes than all other species of vertebrates put together but I want to emphasize that it39s not only true today it has always been true at every point in the geological past since the origin of vertebrates some 500 million years ago In contrast to amphibians reptiles birds and mammals the known biodiversity of living shes exceeds that of known fossil taxa Y I a omens i PERIODS be lining 1o Quaternary g Q 65 Tertiary 135 Cretaceous of 190 Jurassic it 225 Triassic 7 x m 250 Permian 9 345 Carboniferous 395 Devonian 430 Silurian 500 Ordovician 570 Cambrian Precambrian 7O Cambrian 7o Ordovician 35 Silurian 50 Devonian 65 Carboniferous 55 Permian 35 Triassic 0quot 55 Jurassic 70 Cretaceous 64 Tertiary C E p l A 10 Quaternary PERIODS 6 AN ECOLOGICAL CLASSIFICATION OF FISHES It39s important to recognize that shes are highly diverse in terms of number of species but also important to realize that they occupy diverse habitats PRIMARY 331 s H W4 Q 412 r flt DEEP BENTHIC 64 DEEPPELAGIC 50 SHALLOWAWARM 582 399 MARINE SHALLOW7COL 56 EPIPELAGICJ 13 NUMBER OF SPECIES OF BONY FISHES IN THE PRINCIPAL MARINE AND FRESHWATER HABITATS OF THE WORLD The number of bony shes is conservatively estimated at 27300 species more numerous than all other vertebrates put together When we analyze bony shes ecologically that is if we arrange them by major habitat we nd that about 412 are more or less con ned to freshwaters while 582 are marine Additionally we get the following estimates 1 Primary freshwater shes those unable to tolerate salt water about 9036 species 331 of the total number of species of these approximately 8494 94 belong to the Ostariophysi 2 Secondary freshwater shes those living primarily in freshwater but able to tolerate salt water at some point in their lifecycle about 2210 species 81 mostly Cichlidae Cyprinodontidae and Poeciliidae 3 Diadromous fishes those with a life history partly in fresh water partly in salt water about 2 about 160 species 06 includes primarily catadromous eels and anadromous salmonids and galaxiids Warm water littoral and shelf fishes to depths of 200 meters about 10890 species 399 Perciforms the largest group of acanthopterygian shes are the major component Percoids blennioids and gobioids are particularly numerous The major nonperciform component is probably anguilliforms eels Cold water littoral and shelf shes to depths of 200 meters about 1530 species 56 Important components are gadids cods hakes etc zoarcids coldwater blennioids and scorpaeniforms rock shes sculpins poachers etc PRIMARY 331quot DlADROMOUS 06 5 H W Q 4 7 Q 412 DEEP BENTHIC 64 DEEPvPELAGIC 50 HALLOWWARM 5 a 2 399w SHALLOWCOL 56 EPIPELAGICJ 13 Continental slope and deepsea benthic fishes below 200 meters about 1750 species 64 Important components are macrourids rattails grenadiers brotulids zoarcids anguilliforms and scorpaeniforms Epipelagic fishes high seas above 200 meters about 355 species 13 Important components are scombroids tunas and mackerels and exocoetids ying shes Deep pelagic fishes below 200 meters including mesopelagic and bathypelagic realms 1365 species 50 Chief components are stomiiforrns myctophiforms and deepsea angler shes 11 7 THE GROUPS OF FISHES WE WILL STUDY IN FISH 311 By now I imagine that some of you are a bit worried about all this diversity There39s no question that things are extremely complex The branching diagram shown on page 13 contains a lot of information particularly lots of names that are in some cases long hard to pronounce and dif cult to memorize There are a number of extinct groups indicated as well But let me assure you that this diagram is incredibly simpli ed compared to reality particularly in the diversity of fossil material that is available But what you see here simpli ed as it is is not what you will be responsible for in this class 12 EVOLUTION OF MAJOR GROUPS OF VERTEBRATES The width of the branches indicates the relative number of recognized genera for a given time level on the vertical axis Time in millions of years indicates the beginning of geological periods c b 0quot 39 4 59 v crl39bo e 9 gt0 0Q lt60 PERIOD OF TIME Cenozoic 65 Cretaceous 1 3 5 Jurassic 180 Triassic 93 6 lt9 6 6 gt5 quot C3939OI 230 clocln39b39l39ln Permian 280 Halo hall mum I Carboniferous 345 H C I1 I sPFAOMqvai I Devonian Paeoaw 400 39I39er ll 391 wquot a r d fcMh 5 Aconh Chi SHu an 425 ri Ordovician 500 39 P raspiaa39o Cambrian 570 Instead we will recognize only those major groups of shes that have survived to the present day that is we ll forgot about all the extinct groups and concentrate only on the ve major groups diagrammed below The Myxini containing the hag shes about 70 species The Cephalaspidomorphi the lampreys about 38 species The Chondrichthyes the sharks rays and Chimaeras about 970 species The Actinopterygii the bony or ray nned shes at least 27300 perhaps as many as 32000 The Sarcopterygii the lobe nned shes including two living species of coelacanth the crossopterygian genus Latimeria and six species of lung shes Evolution of the Major Groups of Living Fishes CHONDRICHTHYES Sharks CEPHALASPIDOMORPHI Rays v SARCOPTERYGII Chimaeras Lampreys Lobe finned shes MYXINI ACTI NOPTERYGII Hagfishes Bony or ray finned shes 14


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