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A GUIDE TO THESTUDYOFFISHES BY DAVID STARR JORDAN President cf Leland Stanford With Colored and Frontispieces IN TWO J uiilor University 42"] Illustrations Vni,UMES VOL I " I am the wiser in respect to all edge and the better qualified for knowing brfjok." NEW that there is — Thoreait ^T)RK HENRY HOLT AND COMPANY 1905 a knowl- for all fortunes minnow in that l/'" c-^ i G Copyrig-ht, 1905 EV HENRY HOLT AND COMPANY Published March, 1905 ROBERT DRUMMOND, PRINTER, NHW YORK To trbeoJ)ore Gill, Ichthyologist, Philosopher, Critic, Master in this volume is dedicated Taxonomy, PREFACE This work treats ofthe fish from all the varied points of view ofthe different branches ofthestudyof Ichthyology In general all traits ofthe fish are discussed, those which the fish shares with other animals most briefly, those which relate to the evolution ofthe group and the divergence of^ its various classes and orders most fully The extinct forms are restored to their place in the series and discussed along with those still extant In general, the writer has drawn on his ichthyologist, and with this on all own experience as an the literature ofthe science Special obligations are recognized in the text To Dr Charles indebted for a critical reading of most of his proof-sheets to Dr Bashford Dean, for criticism ofthe proofsheets ofthe chapters on the lower fishes to Dr William Emerson Ritter, for assistance in the chapters on Protochordata; to Dr George Clinton Price, for revision ofthe chapters on lancelets and lampreys, and to Mr George Clark, Secretary of Stanford University, for assistance of various kinds, notably in the prep- H Gilbert, he is ; ; aration ofthe index many To Dr Theodore Gill, he has been for years constantly indebted for illuminating suggestions, and Barton Warren Evermann, for a variety of favors To Dr Richard Rathbun, the writer owes the privilege of using illustrations from the "Fishes of Xorti: and Middle America" by Jordan and Evermann The remaining plates were drawn for this work by Mary H Wellman, Kako Morita, and Sekko Shimada Many ofthe plates are original Those copied from to Dr other authors are so indicated in the text No bibliography has been included in this work A writers so complete as to have value to the student woulcl list of make viii Preface a volume of itself The principal works and their autliors are discussed in the chapter on the History of Ichthyolog}', and with must be contented a book valuable to technical students, interesting to anglers and nature lovers, and instructive to all who open its pages tliis for the present the reader The writer has hoped to make David Starr Jordan Palo Alto, S.xnt.a Cl.-vr.a October, County, Cal., 1904 CONTENTS VOL I CHAPTER THE What a Fish? is Fish LIFE OF —The THE FISH Long-eared Sunfish I (Lepomis megalotis) — Form PAGE ofthe Fish — Face ofthe — How the Fish Breathes —Teeth ofthe Fish — How the Fish Sees Color ofthe Fish.— The Lateral Line.— The Fins ofthe Fish.— The Skele- — The Fish — The Fish's Nest ton ofthe Fish Fish Action.— The Air-bladder.— The Brain in CHAPTER II THE EXTERIOR OFTHE Form of Body —Measurement of Ctenoid and Cycloid Scales — Lateral Line — Function ofthe — The Scales or E.xoskeleton — Bony and Prickly Scales Lateral Line — The Fins ofFishesthe Fish — Placoid ofthe FISH Scales Muscles 16 CH.\PTER III THE DISSECTIOX OFTHE FISH Sunfish — The Viscera — Organs of Xutrition — The — The Spiral Valve — Length ofthe Intestine The Blue-green tary Canal CHAPTER Alimen26 IV THE SKELETON OFTHE FISH — Homologies of Bones ofFishes — Parts ofthe — Names of Bones ofFishes — Bones ofthe Cranium — Bones ofthe Jaws — The Suspensorium ofthe Mandible — Membrane Bones of Head — Branchial Bones — The Gill-arches — The Pharyngeals — The Vertebral Specialization ofthe Skeleton Skeleton Column —The Interneurals Shoulder-girdle — The Primitive Fishes —The and Interhsemals.- The Pectoral Limb.— The Posterior Limb — Degeneration — The Skeleton Skeleton of Sharks.— The ;\rchipterygium in 34 Contents X CHAPTER V, MORPHOLOGY OFTHE FINS OFFISHES PAGE ofthe Origin ofthe Fins of Fishes.^Origin ofthe Paired Fins.— Development TheCurrent Paired Fins in the Embryo.— Evidences of Palaeontology.— Fold.— Lateral theof Theory ories as to Origin of Paired Fin.— Balfour's Objections.— Objections to Gegenbaur's Theory.— Kerr's Theory of Modi- External Gills.— Uncertain Conclusions.— Forms ofthe Tail in Fishes Homologies ofthe Pectoral Limb.— The Girdle in Fishes other than fied — 62 Dipnoans CHAPTER VI THE ORGANS OF RESPIRATION Fishes Breathe.— The Gill Structures.— The Air-bladder.— Origin ofthe Air-bladder.— The Origin of Lungs.— The Heart ofthe Fish.— The Flow How of 91 Blood CHAPTER Vn THE NERVOUS SYSTEM The Nervous System.— The Brain ofthe Fish.— The Pineal Organ.— The 109 Brain of Primitive Fishes.— The Spinal Cord.— The Ner\-es CHAPTER VHI THE ORGANS OFThe Organs of SENSE of Sight — The Organs of — The Sense of Taste — The Sense of Touch Smell.— The Organs Voices ofFishes CHAPTER Hearing 115 LX THE ORG.ANS OF REPRODUCTION The Germ-cells, —The Eggs ofFishes — Protection ofthe Eggs — Sexual Modi124 fication CHAPTER X THE EMBRYOLOGY AND GROWTH OF — General Laws of Development — The — The Development ofthe Bony Fishes Development Fishes — Peculiar Larval Fomis — The DevelFlounders — Hybridism — The Age ofFishes — Tenacity of Postembryonic Development Signifi- cance of Facts of Development The Larval opment ofFISHESof Contents xi PAGE Life.— Effect Temperature on Fishes.— Transportation of Fishes.— Reproduction of Lost Parts.— Monstrosities among Fishes 131 of CHAPTER XL INSTINCTS, HABITS, AND ADAPTATIONS The Habits of Fishes.— Irritability of Animals.— Nerve-cells and Fibers The Brain or Sensorium.— Reflex Action.— Instinct.— Classification of —Variability of Instincts.— Adaptations Environment.— Flight — Quiescent Fishes.—Migratory Fishes.—Anadromous Fishes.— Pugnacity ofFishes — Fear and Anger Fishes — Calhng theFishes Sounds Fishes — Lurking Fishes — The Unsymmetrical Eyes the Flounder — Carrying Eggs the Mouth Instincts to ofFishes in ofof in CHAPTER 152 XII ADAPTATIONS OFFISHES — Venomous Spines — The Lancet ofthe Surgeon-fish — Spines ofthe Sting-ray — Protection through Poisonous Flesh ofFishes Electric Fishes — Photophores or Luminous Organs — Photophores the Iniomous Fishes — Photophores of Porichthys — Globefishes — Remoras Sucking-disks of Clingfishes — Lampreys and Hogfishes — The Sword—The Paddle-fishes — The Sawfishes — Peculiarities of Jaws and TemperTeeth — The Angler-fishes — Relation of Number of Vertebra; ature, and the Struggle for Existence — Number of Vertebrje: Soft-rayed Fishes; Spiny-rayed Fishes; Fresh-water Fishes; Pelagic Fishes — Varia—DegeneraConditions of tions Fin-rays — Relation of Numbers tion of Structures — Conditions of Evolution among Fishes Spines ofthe Catfishes in fishes to Life to in CHAPTER COLORS OF 179 XIII FISHES — Protective Coloration, — Protective Markings — Sexual Colora— Nuptial Coloration — Coral-reef Fishes — Recognition Marks — InPatSpirits.— Variation tensity of Coloration — Fading of Pigments Pigmentation tion in in 226 tern CHAPTER XIV GEOGRAPHICAL DISTRIBUTION OF Zoogeography riers — General — Species Changed Laws of Distribution.- Species Absent through through Natural Failure to FISHES Absent through Bar- Maintain Foothold.— Species Selection.— Extinction of Species.— Barriers Contents xii Checking Movements in Distribution Distribution ofFishes toral — of — Agency Marine Marine Species Temperature the Central Fact Centers of Distribution of Ocean Currents Fishes — Distribution Faunal Areas of — Equatorial tion of Fresh-water Fishes — Pelagic — FishesFishes Littoral — Bassalian Fishes Lines by Coast Lit- — Minor — Realms of Distribu— Equatorial Zone — Southern Fishes most Specialized — Xorthern Zone — Origin ofthe New Zealand Fauna Zone CHAPTER 237 XV ISTHMUS BARRIERS SEPARATING FISH FAUNAS of Suez — The Fish Fauna of Japan — Fresh-water Faunas of — Faunal Areas of Marine Fishesof Japan — Resemblance of Japan- The Isthmus Japan and Mediterranean Fish Faunas ese — Significance of Resemblances — Source of — Effects of Direction of Shore Lines — Numbers of DilTerent Faunas — Significance of Rare Forms — Distribution of Genera Shore-fishes — Extension of Indian Fauna — The Isthmus of Suez as a BarDifferences between Japanese and Mediterranean Fish Faunas Faunal Resemblances in rier to Distribution Suez to the Evidences of Submergence of Isthmus of Mediterranean Explained by Present Conditions Panama of — Geological — The Cape of Good Hope as a Barrier to Fishes — Relations of Japan the as a Barrier to Distribution Jenkins — Views Isthmus of Panama Panama — Catalogue of — Unlikeness ofof — The Hill of Dr Giinther on the Isthmus of — Conclusions — Final Hypothesis as Fishesof Panama — Conclusions of Dr Isthmus Species on the Shores CHAPTER X\ Evermann & Panama of to 255 I DISPERSION OF FRESH \V.\TER FISHESThe Dispersion to of Dispersion — Generalizations as — Conclusions of Cope Fishes.— The Problem of Oatka Creek — Questions Raised by Agassiz Questions Raised by Cope.— Views of Giinther —Fresh-water Fishesof North America.— Characters of Species.— Meaning of Species.— Special Creation Impossible.— Origin of American Species ofFishes CHAPTER 282 X\7I DISPERSION OF ERESH-W.\TER FISHES (Co„li„„cd.) Local — Fa\-orable — Water-sheds — How Fishes Cross Water-sheds —The Suletind —The Cas.siquiare —Two-Ocean Pass — Mountain Chains — U]>land Fishes — Lowland Fishes — Cuban Fishes — Swampy Water sheds — The Great Basin of Utah —Arctic Species Lakes — Causes of Barriers to Dispersion of Fresh-water Fishes: IJarriers Waters Have Most Species in DisTXTsion still in Operation ^ 297 Subclass Dipneusti, or Lung-fishes 6io Palfeozoic time, their origin with that or through that ofthe be traced to the Ichthyotomi or other primitive sharks These two groups are separated from all the more primitive latter to In its origin vertebrates by the presence of lungs diverticulum from the aliarises as a or air-bladder lung the earliest fishes as a breathing-sac, mentary canal, used by thethe respiratory functions lost in the progress of further difish-like Nothing ofthe nature of lung or air-bladder is vergence found in lancelet, lamprey, or shark In none ofthe remaining groups offishes is it wholly wanting at all stages of development, although often lost in the adult Among fishes it is most completely functional in the Dipneusti, and it passes through all stages of degeneration and atrophy in the more specialized bony fishes In the Dipneusti, or Dipnoans, as in the Crossopterygians and the higher vertebrates, the trachea, or air-duct, arises, as from the ventral side ofthe oesophagus In the more specialized fishes, yet to be considered, it is transferred to the dorsal side, thus avoiding a turn in passing around the oesophagus itself From the sharks these forms are further distinguished by the presence of membrane-bones about the head From the Actinopteri (Ganoids and Teleosts) Dipnoans and Crossopterygians are again distinguished by the presence ofthe fringe-fin, or archipterygium, as the form ofthe paired Umbs From the Crossopterygians the Dipnoans are most readily distinguished by the absence of maxillary and premaxillary, the characteristic structures ofthe jaw ofthe true fish The upper jaw m the Dipnoan is formed of palatal elements attached directly to the skull, and the lower jaw contains no true dentary bones The skull in the Dipnoans, as in the Chiuuvra, is autostylic, the mandible articulating directly with the palatal apparatus, the front of which forms the upper jaw and of which the pterygoid, hyomandibular and quadrate elements form an immovable part The shoulder-girdle, as in the shark, is a single cartilage, but it supports a pair of superficial membrane-bones In all the Dipnoans the trunk is covered with imbricated cvcloid scales and no bony plates, although sometimes the scales are firm and enameled The head has a roof of wellaboA'c stated, Subclass Dipneusti, or Lung-fishes i i developed bony plates made of ossified skin and not corresponding with the membrane-bones of higher fishesThe fish- membrane-bones, like yet dift'erentiated opercles, branchiostegals, The teeth have the form on the pterygoid areas are etc., not of grinding-plates ofthe palate, being distinctly shark-like The paired fins are developed as archipterygia, rays, and the pelvic arch consists of a single the two sides symmetrical and connected in front in structure often without cartilage, There is but one external gill-opening leading to the which, as in ordinary end fishes, are fringe-like, gill-arches, attached at one In the young, as with the embryo shark, there is a bushy gill, which looks not unlike the archipterygium pec- external toral fin itself, although its rays are of dift'erent texture In bony and enamsome recent forms deep sunken in the skin The early forms, as in the Ganoids, the scales were but in claspers have disappeared, the nostrils, as in the frog, open into the pharynx, the heart is three-chambered, the arterial bulb with many valves, and the cellular structure ofthe skin and of other tissues is essentially as in the Amphibian The developed lung, fitted for breathing air, which seems the most important of all these characters, can, of course, be traced only in the recent forms, although its existence in all Besides the development others can be safely predicated ofthe lung we may notice the gradual forward movement ofthe shoulder-girdle, which in most ofthe Teleostomous In bony fishes generally to the head fishes is attached there is no distinct neck, as the pogt-temporal, the highest bone ofthe shoulder-girdle, is articulated directly with the skull In some specialized forms { Batistes, Tctraodon) it is In a few groups (Apodes, fused with it immovably even eled, Opisthomi, Heteromi, etc.) this connection ancestrally possessed is lost through atrophy and the slipping baclavard shoulder-girdle leaves again a distinct neck ians tinct In theofthe Amphib- and ah higher vertebrates the shoulder-girdle is disfrom the skull, and the possession of a flexible neck is an important feature of their structure In all these higher forms the posterior limbs remain abdominal, as in the sharks and the primitive and soft -rayed fishes generally In these the pelvis or pelvic elements are attached toward the middle Subclass Dipneusti, or Lung-fishes In the giving a distinct back as well as neck disappears, neck the well as as spiny-rayed fishesthe "back" the pelvic elements being attached to the shoulder-girdle, and ofthe body, in a few extreme forms Ophidion) the pelvis (as the chin of Classification divided We two into follow Dean distinct class, to Dipnoans classes, — By Woodward is fastened at the Dipneusti are the Sirenoidei and the Arthrodira in regarding the latter as representative of a leaving the Sirenoidei, with the Ctenodipterini, constitute the subclass of Dipneusti The Sirenoidei are divided by Gill into two orders, the Monopneumona, with one To the lung, and the Diplopneunioiui, with the lung divided the Lcpidosirenidcv belong order latter To the former the and presumably the extinct families also belong, Zittel although nothing is known of their lung structures and Hay adopt the names of Ctenodipterini and Sirenoidei for these orders, the former being further characterized by the very Ccratodontidcc, fine fin-rays, more numerous than Ctenodipterini — In Order their supports this order the cranial roof-bones and numerous, and the rays ofthe median fins are very slender, much more numerous than their supports, which are small are inserted directly on the vertebral arches In the UronemidcB the upper dentition comprises a cluster conical denticles on the palatine bones; the lower dentition consists of similar denticles on the splenial bone The vertical fins are continuous and the tail diphycercal of small, blunt, There is a jugular plate, as in Amia in the Carboniferous, Uronemus The few species are found lobatus being the best-known species In Dipteridce there is a pair of dental plates on the palatines, and an opposing pair on the splenials below Jugular plates are present, and the tail is usually distinctly heterocercal In Phaneropleuron there the very long dorsal; a distinct anal fin shorter than Phaneropleuron andersoni is known from is Scotland, and Scaumenacia curta is found at Scaumenac Bay Canada In Dipterus there are no marginal teeth, and the tail is in the Upper Devonian of heterocercal, not diphycercal, as in the other ally Numerous species of Dipterus occur in Dipnoans generDevonian rocks Subclass Dipneusti, or Lung-fishes In these the jugular plate is present, as in Uronemus Dipterus valenciennesi is the best-known European species Dipterus nelsoni and numerous other species are found in the Chemung and other groups of Devonian rocks in America In the CtenodontidcB the tail is diphycercal, and no jugular known specimens In Ctenodus and no jugular plate and there are no marginal The numerous species of Ctenodus and Sagenodus belong plates are present in the Sagenodus there teeth Fig 383 is Phaneropleuron andersoni 'H.uxley; restored; Devonian chiefly to the Carboniferous age Ctenodus wagneri the Cleveland shale ofthe Ohio Devonian (After Dean.) found in Sagenodus occidenis talis, one ofthe many American species, belongs to the coalmeasures of Illinois As regards the succession ofthe Dipneusti, Dr Dollo regards Dipterus as the most primitive, Scaumenacia, Uronemus, Protopterus, and Lepidosiren following Ctenodus, Ceratodus, in order The last-named genus he thinks marks the terminus ofthe group, neither Ganoids nor Amphibians being derived from any Dipnoans Order Sirenoidei — The living families of Dipneusti differ from these extinct types in having the cranial roof-bones reduced in number There are no jugular plates and no marginal The tail is diphycercal in all, ending in a and the body is covered with cycloid scales To forms the name Sirenoidei was applied by Johannes teeth in the jaws long point, these Muller — The CeratodontidcB have the teeth Family Ceratodontidae above and below developed as triangular plates, set obliquely each with several cusps on the outer margin Nearly all the species, representing the genera Ceratodus, Gosfordia, and Conchopoma, are now extinct, the single genus Neoceratodus still Numerous fragments of Ceraexisting in Australian rivers todus are fovmd in Mesozoic rocks in Europe, Colorado, and 14 Subclass Dipneusti, or Lung-fishes India, Ccratodus by Agassiz m latissimus, figured 1838, being the best- known species The abundance ofthe fossil teeth of Ceratodus renders the discovery of a hving representative ofthe same type a matter of great interest In 1870 the Barramunda ofthe rivers of Queensland was described rS — Teeth of Ceratodus runcinatus PlieFig 38.3 (After Zittel.) ninger Carboniferous by Krefft, who recognized tionship to Ceratodus name of Ceratodus and gave forsteri rela- its it the Later, and was separated as a distinct group by Castelnau in 1876, under the name o generic differences were noticed, it of Neoceratodus (later called Epicera- todus by Teller) Neoceratodus forsteri and a second species, Neoceratodus miolepis, have been since very fully discussed by Dr Giinther and Dr Krefft Pig 385.— Archlpterygium of Neoceratodus forsteri Gilntlier «, Subclass Dipneusti, or Lung-fishes 615 They are known in Queensland as Barrainimda They inhabit the rivers known as Burnett, Dawson, and Mary, reaching a length of six feet, and being locally much valued as food From the they are known to the settlers in QueensAccording to Dr Gunther, "the Barramunda is said to be in the habit of going on land, or at least on mud-flats and this assertion appears to be borne out by the fact that it is provided with a lung However, it is much more probable that it rises now and then to the surface ofthe water in order to fill its lung with air, and then descends again until the air is so much deoxygenized as to render a renewal of it necessary It is also said to make a grunting noise which may be heard at night for some distance This noise is probably produced by the passage ofthe air through the oesophagus when it is expelled for the purpose of renewal As the Barramunda has perfectly developed gills besides the lung, we can hardly doubt that, when it is in water of normal composition and sufficiently pure to yield the necessary supply of oxygen, these organs are sufficient for the purpose of breathing, and But that the respiratory function rests with them alone when the fish is compelled to sojourn in thick muddy water charged with gases, which are the salmon-colored flesh, land as "salmon." ; products of decomposing organic matter (and this must be the case very frequently during the droughts which annually exhaust the creeks of tropical Australia), to breathe air with way indicated above in which it commences it lung in the its If happens to be unfit for breathing, have any function thethe is medium perfectly gills cease to only in a less degree, the gills may still continue to assist in respiration The Barra; if „ can breathe by either ^ „^„ Fig 386 — Lpper jaw of Neoceraby both simul- todusforsteri Gimtber (After taneously It is not probable that it lives freely out of water, its limbs being much too flexible for supporting the heavy and unwieldy body and too feeble munda, in ' fact, ,, -' , gills or lung alone or '^ ' '' „ Subclass Dipneusti, or Lung-fishes 6i6 Howgenerally to be of much use in locomotion on land ever, it is quite possible that it is occasionally compelled to leave the water, although we cannot believe that it can exist any length of time, we know nothing except development propagation or in a lively condition for AMthout " Of its that deposits a great it it number of eggs ofthe size of those of a newt, and enveloped in a gelatinous case that the young are provided with external toptcrus and Polyptcrus "The discovery infer as Pro- in does not date farther back Ccratodiis of We may gills, than the year 1S70, and proved to be ofthe greatest interest, not only on account ofthe relation of this creature to the other living Dipneusti and Ganoidei, but also because it threw fresh light on those singular fossil teeth which are found in and Jurassic formations strata of Triassic _Lower iaw 3S7 Yu! ,\,acer„tndn.'