THE EASTERN BELTED , MEGACERYLE ALCYON ALCYON (LINNAEUS), IN RELATION TO MANAGEMENT 1

J. CLARKSALYER, II U.S. Department of the Interior, Fish and Wildlife Service, Washington,D. C. AND KARL F. LACLER Department of Zoology, University of , Ann Arbor, Michigan

ABSTRACT

The kingfisher is the most common and universally distributed predator of fish in Michigan. Its principal migration routes are along the shores. Nesting territories are establishedalong streamsand lake shores;they are usually larger in the former than in the latter. The kingfisher is diurnal in its feeding with three peaks of activity--morning, afternoon, and early evening. First feeding of fledgelingsis on ;this food is followed by crayfish,then by fish. Fish eaten averageabout 2.3 inchesin length and at fish hatcheriesare mostly the speciesbeing propagated. On natural waters the food consistsmostly of non-food and non-game and crayfish. Predation pressure varies with season and within season according to weather, nest distribution, and en~ vironmental suitabliity of waters for feeding activity. Becauseof their feeding proclivities,it is undesirableto have kingfishersabout fish hatcheriesand rearing stations. At present,general control on natural waters is not biologicallyjustified.

INTRODUCTION This report is one of a seriesdealing with the relationshipsbetween fish predatorsand fish production.The purposeof this particularstudy has been to determine the relation between the eastern and the production,artificial or natural,of food,game, and foragefishes. It has thus been necessaryto determinethe nature and extent of fish predationby thisbird on naturalwaters and at outdoorfish hatcheries and rearingstations. As it becameincreasingly apparent that substantial lossesof highly valuedfish couldresult from the feedingactivities of this predatorat such stations,non-lethal controls were designedand tried (Laglet, 1939). Our data are of two kinds: field observationsof habits of and the natureof fish populations;laboratory analyses of stomachcon- tents. One or the other of the authors has devoted some time to this , Contribution from the Department of Zoology of the University of Michigan, and from the Institute for Fisheries Research of the Michigan Department of Conservation. Sub- stantial financial aid was given this study by the Wildlife Management Institute (Amer- ican Wildlife Institute) and by the Associated Fishing Tackle Manufacturers. 97 98 AmerJcanFisherJes Society study each year from 1930 to 1945 and field observationsand collections have been made in Michigan during all months. Most of the used for food analysesfrom fish-culturalestablish- ments were obtained by employees of the Michigan Department of Conservationwho followed instructionsissued by us for the care and preservationof this material. Many other individuals,including colleagues in the Institute for Fisheries Research,collected specimensfor us from natural waters. We extend our sinceregratitude to the many personsand agenciesthat have helped us. Special thanks are due to Carl L. Hubbs, Fred A. Westerman,A. S. Hazzard, J. W. Leonard, and Karl E. Goellner. Ralph Hile read the manuscript and offered many critical suggestionsfor its improvement. F•ELD O•S•:RV^T•ONS Since the emphasisof this paper is on the relationshipof the food habits of the kingfisherto practicalfishery management, we are present- ing no lengthy discussionof life history which has been summarizedby Bent (1940). Included only are apparently new materials based upon actual field observations and certain detailed information which will supply greater detail on activities of the bird which are generalizedin other, previous accounts. Salyer is responsiblefor the field observations and their interpretation in the sectionson distribution and migration, mating and nesting, and populations. Of all the observationsrecorded in the past on this species,those made by Gould2 appear to be the most accurate. In fact, this painstakingstudy well merits publicationeven at this late date (vide Lagler, 1943). Distribution and Migration.--Of all the winged fish predators of Michigan waters,the kingfisheris probablythe most widely distributed. The speciesis principallya summerresident in Michiganbut occasional individualsmay be seenall winterlong. The fact that this is a peninsular state with most of its boundaries on the shores of the Great Lakes has a profoundinfluence on the migration and distributionof the kingfisher in the region. The first thing that strikesthe observerwho worksdaily in the field throughoutthe spring and summer months is the fact that travel is not haphazardor "cross-country"as stated by many writers. In short, in both the Upper and Lower Peninsulas,the shoresof the Great Lakes are basic migration highways for the birds' movement. Kingfishers pass up the shoresof the Great Lakes in almost the same mannerand regularity that native hawksexhibit in their migrations.On April 12, 1931, Salyer observedindividual kingfishers moving along the west shore of Lake Huron north of the city of Port Huron at the rate of about 15 per hour during a 4-hour period in the morning. In the fall of the same year he found kingfishersmoving southwardon the .oGould, Victor Eugene. 1934. "A monograph of the belted kingfisher Megaceryle a•cyon (Linnaeus)", 311 pages plus illustrations, summary, and bibliography. Typewritten. Kingfisherin Relation to Fish Management 99 shoresof at the mouth of the Pentwater River at the rate of 12 per hour. This observationwas made on October 25; previous to this date Salyer had spent 3 days on the headwatersof the Pere Mar- quette River and had seen only one kingfisherthere. It seemsto the observer that on spring migration, kingfishersmove along the Great Lakes shoresand turn up major river valleys which they folow, dropping off as they locate, or return to, both new and old nestingsites. Mating and Nesting.--The female is slightly the larger, usually weigh- ing a little more than 6 ounces,whereas the male averagesa little less than 6 ounces. Observationsby Salyer indicate that the female is the more aggressiveof a nestingpair and takes the initiative in selectingthe nesting territory. On the Huron River in Washtenaw , the Platte River in Benzie County, and the Little Manistee River in Lake County, femaleshave been found activly defendingsites prior to getting a mate. A mated pair of birds not only has a definitenesting territory, which they defend against all other individuals with fury and persistence,but their territory can be marked almost to the exact yard by the observer becauseof the behavior of the birds, describedbelow. A great amount of time in early mating seasonis spent by the mated pair in traversing and defending their territory. During the daylight hours at least, most of the territory appearsto be patrolled at intervals of 10 to 15 minutes. All larger ,including man, cats, dogs,and larger birds are circledand scoldedand often followedthrough the area. Invadersof the samespecies are violentlyset upon forthwith. The sizeof the nestingterritories located on lakescontrasts sharply with that of siteson moststreams or rivers. On lakes the longestterritorial selec- tion found was 1• miles of shore, but the average comprisesabout • mile. This l•-mile territory was occupiedby a nesting pair of kingfisherson Black Lake, CheboyganCounty, in June and July 1931. On the other hand, two nestingpairs used about • mile each of shore- line on PleasantLake, WashtenawCounty, in May 1931. Another pair, during the same period, selectedabout ½ mile of shorelineof Portage Lake, Livingstonand WashtenawCounties, which includedan extensive cottage area. The stretch of shorelineselected invariably includesall or a portion of a small shelteredbay which providesa reasonableextent of unruffledfeeding surface regardlessof most directionsof the wind. Our studiesindicate that exposureto wind action, especiallyon the lake habitat of the species,is almostas importantas turbidity in determining the distributionof the populationand the locationand successof feeding. In this respect,it is significantto note that there are morekingfishers per unit of shorelineon the smaller Michigan lakes than upon such large, roughbodies of water as Houghton,Hubbard, and Mullet Lakes. Limitations of feedingarea on rivers by vegetativecover, deep and unfishablepools, and fast currentswhich reducevisibility, apparently requirethe birdsto establishmuch larger territories. Thus, the usualrange is mostlikely to be from 1• to 3 miles or more. In May and June lOO American FisheriesSociety

1931, Salyer kept under closeobservation a pair of birds known to dominatethe headwatercountry of the Little ManisteeRiver for a dis- tance of 6 miles. Althoughthese birds were not marked,they appeared to be the only pair of kingfishersalong this entirestretch of river. Their nestinghole wasfound and the feedinghabits both of the adultsand the young after leaving the nest were followeddaily. Here the extensive "brushing-over"of long sectionsof the stream and the fast current seemedto limit feedingopportunities. On the lower reachesof some of Michigan's larger rivers which are ordinarilyquite turbid, the bird is an infrequentnester; however,all adjacentsloughs and backwaterareas with relatively clear water have their nestingbirds so that the nestingpopulation of the entire stateis of a somewhatuniform density. In May 1931 Salyer made someinteresting observations on a female bird which had selecteda territory on a Huron River cutoff a few miles southeastof Ann Arbor, Michigan. This female actively defendedthe regionfor 2 days and took a mate on the afternoonof the secondday. By the morningof the third day the male bird seemedto becomefamiliar with the territory originallydefended by the female. This male (K-318) was collectedat 9:00 a.m. on the third day but by 3:00 p.m. on the same day there had appeareda new male bird which also learnedthe territoryin a shorttime. The territorywas composed of two smallponded areas estimated over-all at about 35 acres of water and marsh. During the early mating seasonit is commonto see and hear male kingfishersflying at altitudes of about 300 feet up this river valley. At regular intervals the migrant gives his characteristicrattling call. If a determinedand vigorouschallenge is returnedby a prior occupantfrom below, he frequentlypasses on without stopping. Failing to get a chal- lenge, the bird is often seen to go into a spectacularaerial dive and fly at a lower level. Sometimeshe meets a female; if she already has a mate, she drives him away but if not, she may accept him. In any given watershedthere alwaysseem to be one or two bachelorbirds which have not mated and which give the mated pairs considerableconcern• especiallyin the early morninghours, as evidencedby repeatedobserva- tions followed by collections. FeedingHabits.---As is generallyknown, the kingfisheris a very adept fisherman,but its fishingis confinedto thoseindividuals of its prey near the surfaceor in shoalwater. It apparentlyprefers to feed in the shallow watersof streamsor lakes. In lotic habitats,fishing success is not as good in deeppools as it is in shallowerreaches having a moderateto slowcur- rent. The kingfisheris strictly a diurnal bird; it speedilyretires with the adventof twilight in the evening.On the other hand,it is an early bird on the stream,becoming active in the early light before sunrise.The first hour of daylightis spentin feedingand repeatedlypatrolling the territory duringthe nestingperiod. Kingfisher in Relation to Fish Management 101

Peaksof feedingactivity occurin the morning,afternoon, and evening. Stomachsof birds taken very early in the morningprior to 7:00 a.m. are most often quite empty or contain a single recently taken item. Full stomachsare found more frequently in birds collectedfrom 7:00 to 10:30 a.m. From about 11:00 a.m. to 2:30 p.m. apparentlyis largely a non-feedingperiod, with stomachscollected between 2:30 and 4:30 p.m. again being relatively full. Stomachstaken from around 4:30 to 6:30 p.m. generallyshow bone remainsand seeminglythe birds make their final feedingof the day just prior to dusk. The first food which the young get when they leave the nest and start feedingwithout parental aid is insects,particularly mayflies. They are very fat whenthey first fly; this goodcondition is apparentlyneces- sary becausethe parentsgive the youngbirds little foodjust prior to the departure,perhaps to compelthem to beginseeking food for themselves. Severalbroods have been watchedby Salyer throughoutthe first and secondweeks of their livesafter they left their nests. In general,the familyremains together for 10 to 15 days. Duringthe first 3 or 4 days the youngcatch only flyinginsects from advantageousperches near the water,the parentsgenerally perch on deadtrees or exposedbranches and the youngon nearbydogwood, alder, or othershrubs where they are somewhathidden. From these protected vantage points they fly out and seizea passingmayfly or caddisflyin a characteristicfly-catcher manner. All of the youngof a broodtaken on July 4, 1941, on the Black River (MontmorencyCounty) on theirfirst day on wingwere feeding on insects. Severalother broodsand individualyoung showed the samehabit. The youngcan be readilyidentified as suchby their tarsalnesting callosities whichgradually disappear toward the endof the thirdweek on the wing (determinedby fieldcollections of birdsof knownage by Salyer). Duringthis periodthe youngcan be approachedand observedquite closelydespite the frantic warningsof the parents.The cries of the parentsseem to rally the flocktogether and to keepit concentratedgen- erally within a 100-yardstretch along the marginof the water area. It is mostamusing to seeboth cedar waxwings and young kingfishers feed- ing on flying insectsalong northern trout streamsin an almostidentical manner. Afterlearning to catchinsects, the young devote the fifth to tenthday on the wingalmost exclusively to learningto capturecrayfish. Broods in thisperiod show a strikinguniformity of behavior.For example,J. R. Greeleycollected practically all of a broodon the Big SouthBranch of the Pere MarquetteRiver (in Masonand NewaygoCounties) in the periodfrom July 23 to July 28, 1931. All of thesebirds were feeding whollyon Cambarus.Again, Greeley collected a broodof youngon August14, 1931,on the same stream, of whichall hadfed upon crayfish whilethe male parent, also collected, was feeding normally upon fish; its stomachcontained a 4-inchredhorse (Moxostoma sp.). This transitional crayfish-eatinghabit is substantiatedby the stomachcontents of individual 102 AmericanFisheries Society

young birds from severalother points throughoutthe state. Thus, the kingfisherdoes not ordinarilybegin to catchfish until the latter part of its secondweek on the wing. Two juvenile males and a juvenile female, collectedat 7:30 a.m. on June 25, 1931, all had caughttrout; they were then 18 days on the wing. During the latter portion of the crayfish-feedingperiod, the family group is not as closely knit as previouslyand gradually disperses;the parents finally disappearand the young expand their feeding areas. The young, however,continue for a time to feed in the samegeneral region in which their nest was located. They are usually found feeding first at intervalsof ¬ to • mile from the nestingsite and at greater distances in late August and September. The writers have not spent much time studyingthe life of the king- fisher in the nest as this has been well coveredby other authors. It was obvious, however, from the observationswe made, that we could not accuratelyassay the food of nestlingsby examiningremains in the nests since femaleswere repeatedlyseen during the day to carry off a bolus or pellet of waste or undigestedfood remains from the nest. Becauseof the great quantitiesof food required daily to sustainthe nestlings,the parents must, of necessity,remove the waste or the nest soon would become so congestedwith this material as to leave no room for either adults or young. We have not substantiatedthe observationsof White (1939) that the youngdigest bones totally whereasthe adultsregurgitate the heavierbone material. It appearsto us rather that in the initial period of feedingthe nestlings,the parentspresent them with smaller and more delicatefish which disintegrateeasily . Adults seemcapable of digesting bone for in the stomachswe have examinedboth partially dissolvedas well as fragmentedbone have appeared. Thus, we hold, there is evidence of erosionand absorptionin contradictionto the findingsof White (1939). It is our opinionthat digestionin a kingfishertakes place more rapidly than in any other of the bird predatorsof Michigan fishes. The stomachs of kingfishersshot by us while they were feedingin their natural habitat are almost invariably either full of largely undigestedfood or contain only the traces of the last meal. One does not often get intermediate stagesof digestionas in the caseof the heronsof which we have examined many specimens. The total length of all fish eaten by kingfishersin Michigan averages lessthan 3 inchesand rangesfrom 1 to 7 inches(Table 1). The minimum and maximumlengths were encounteredonly onceeach in the 729 speci- mensfor which actual measurementsor estimatesbased on comparative material were available. Most of the lengths determinedwere less than 5 inches; it is likely that fish of greater length are swallowedwith diffi- culty. Individuals have been seen carrying fish in the 6- to 7-inch size group and sometimeshave been observedto drop or abandonthem. Fish which are intentionally or accidentallydropped by kingfishersshow dis- tintive marks on each side from the pincer action of the bill (as also Kingfisher in Relation to Fish Management 103 noted by White, 1936). Often these marks are numerouson larger fish and bear evidenceof manipulation in the unsuccessfuleffort to hold or swallow; we have several trout about 6 inches long in our collection which have these marks. The typical habit of attack and subsequent ingestionis for the bird to grasp a prey fish, forceps-fashion,with a lightning-likethrust aimed a little ahead of the middle of the body, fly to a perch, gradually turn the fish, and swallow it head first. Crayfish are taken similarly but are turned to passdown the esophagustail first. Most of the whole or nearly whole fish and crayfish taken from stomach or esophagusare found headed in the directions indicated and are marked as described. The kingfisherdoes not spear or puncture larger prey with the closedbill as the great blue heron frequentlydoes. The kingfisherhas been accused of taking nestlingbirds; we have never seenthis nor do our food analysesgive evidenceof it. Populations.--Inthe opinionof Salyer, the abundanceof kingfishers on Michigantrout streamsappears to parallelthe strikingincrease of beaver populationof the statein the decadeprior to 1934. Beaverponds opened up long reachesof the streamspreviously sheltered from kingfisher activitiesby overhangingtrees and shrubs.There was a steadyincrement in the kingfisherpopulation per streamunit on the GladwinGame Refuge, which grew in proportionto the increasedarea floodedby the activities of the ever growingbeaver population. In the summerof 1931 there was apparentlyonly onekingfisher on 6 milesof the JordanRiver (Charlevoix and Antrim Counties). In 1934, when $alyer was working in the same regionagain on beaver-troutinvestigations, 15 kingfisherswere found in the same stretch of stream. The relative richness of individual water courses also has a direct bearing on the number of kingfishersand other fish predators. For example,the Big South Branch of the Pere Marquettehad by far a larger populationof predatorybirds, includingkingfishers, than did the main stream. J. R. Greeley, in making a 14-mile float of 7 hours' dura- tion downthe Big SouthBranch, saw at least 12 individualkingfishers. Suchrich streamsas the Platte River (BenzieCounty) have a relatively heavypopulation of kingfishersin comparisonwith a poorerstream such as the Boardman River (Grand Traverse County). Such contrastsare repeatedmany times over the state. For the suitable portionsof the NortheastMargaree and its tributariesin Nova Scotia,White (1936) estimatedone kingfishernest per mile with a minimum of sevenbirds ultimately feeding from each nest. Predation Pressure.--Predationby kingfisherson natural waters is differentin quality and quantity in the differentecological situations repre- sented in the state and it varies in all situations with the weather and seasons.The pressurereaches its low during winter months when only a few birds are in the region. During this time the lakes are usually ice-coveredand feedingroutes are along streamsthat do not freeze over. These are, for the most part, trout streamsin the Lower Peninsulabut 104 AmericanFisheries Society winter-residentkingfishers are so few that no concernneed be felt over them eventhough they are restrictedas to the watersthat they frequent. In the spring,summer, and fall, kingfishershave beenobserved feeding in almostall kinds of waters,both inland and alongthe marginsof the Great Lakes. They are strikinglyabsent from feederor nurseryportions of streamsystems, both trout and non-trout,where the marginalvegeta- tion moreor lesscovers the water,often forming a completecanopy from one to a few feet aboveit. Examplesof the latter situationare the alder- coveredsections of the Maple River near Pellston,Emmet County,and of the HurricaneRiver near AusablePoint, Alger County. Feedingbirds are not seenalong the turbid, lowerreaches of the largerrivers, especially in the southernpart of the Lower Peninsula,such as thoseof the Grand, Kalamazoo,Huron, and Raisin Rivers. Birds are characteristicallyabsent from placesthat they ordinarily frequent along waterwayswhen these are temporarilymuddied by run-off immediatelyfollowing heavy rains. They furthermore retreat from the Great Lakes shoreswhen surf makes the water there unfishable.It may be presumedthat during both of the latter conditions,recourse is had to tributaries or to other shelteredclear waterswhich then experiencea temporaryincrease in the predationpres- sure. Trautman (1940) has describedthe utilization of tributaries for feedinggrounds by migrantsat BuckeyeLake, Ohio, when the main body of water is iced-overor turbid. On most waters the predationpressure is greatestin late summerand early fall, when, as a result of the flight of the young, the total effect is increased;but most "buffer," forage fisheshave reproducedand temporarilyincreased their numbersby this time sothat the relativeeffective pressure on gamespecies by the predators may actually be less. The average increasein daily numbers of birds seenin a givenlocality from springto fall may be fourfoldas indicated by Trautman (1940). FOOD STUDIES Specimensfor food study are from a wide range over the state (Figs. 1 and 2) but only a sample randomly selectedfrom each locality is re- portedhere since experience has shown that the numbersused are adequate and that inclusionof additional onesdiscloses insignificantly few new facts. The methodsemployed in the laboratory for stomach-contentanalyses, which were as preciseas possibleboth qualitatively and quantitatively, follow thosedescribed by Salyer and Lagler (1940). We did not study regurgitatedpellets such as those which formed the basis of White's (1937) work. All quantitative results,except those for lengths of fishes at capture (Table 1), are thus basedon actual countsand measurements. Fragmentaryremains of fish were most often found; in order to obtain informationon sizesof fish eaten, it was thereforeusually necessaryto estimate the former whole lengthsfrom fragments. This estimationwas made by comparisonof remainingparts with entire specimensor with skeletonsof fish of the samekind and of known length. Kingfisherin Relation to Fish Management 105

TABLe.1.--Total length(inches) o! fisheseaten by the easternbelted kingfisher

i•inimum Average Maximum Number of Kind of fish length length length individuals lrouts ...... 1.0 2.3 7.0 651 •uckers ...... 2.7 3.4 4.0 2 Minnows ...... 1.5 2.3 4.0 6 Bullhead ...... 3.5 3.5 3.5 Northern pike . 4.7 4.7 4.7 1 topminnows ... 1.5 1.8 2.0 2 gudmlnnows ... 1.5 3.0 5.0 6 •ellow perch ... 1.8 3.1 4.5 3 Black basses ... 1.8 3.0 5.0 17 Sunfishes ...... 1.8 2.6 4.0 15 Muddlers ...... 1.7 2.7 4.0 14 Sticklebacks ... 1.5 1.8 2.3 11

Kingfisherscollected by shooting,if they contain any bony fragments of food at all, usuallyhave in them certain remainswhich are identifiable as to kind of organism. The stomachsof birds taken in pole traps are practicallyworthless in food analyses.Trapped individualsmake frantic strugglesto escapewith the result that the food is regurgitatedand lost or, if the trap is not so situatedthat the bird is plungedinto water and drowned, food materials which remain in the stomachare very rapidly digested.Furthermore, a strategicallylocated trap will almostinvariably take a kingfisherbefore he has fed at the particular site. Kingfishersat Fish Hatcheries.--It has long been known that king- fishersfeed upon fishesin pondsand racewaysat fish-culturalestablish- ments. Someobservations on predation and many complaintsabout losses to predators at such stationshave been recorded. For example, in 14 kingfishersthat had eaten mostly crayfishesand aquatic insectsfrom an Ohio fish farm, Langlois (1936) found two smallmouthbass in only one bird. He suggestedthat this speciesmight be more destructivethan the wading birds later in the seasonwhen the bassare larger and faster and do not spendso much of their time in the shallows.De la Torre Bueno (1936) reportedtaking reared trout from the stomachsof hatchery king- fishers and added that this bird offers the most serious threat when the young first leave the nestsand fly to rearing pondswith the adults in family groupsof four to six birds. He concludedthat three fingerling trout constitutea day's ration for a kingfisher. The remedy suggested as bestwas screeningof the ponds. Hoover (1936) found 16 rangingfrom 1.5 to $.$ inchesin length in the stomachsof nine king- fisherstaken at hatcheriesin New Hampshireand concludedthat the exact statusof this bird is in need of study and that until suchinvestiga- tion is completed,reasonable control is necessary. The fish rearing stationsin Michigan, where predation by kingfishers occurs,are of two kinds; pondfishhatcheries, usually used for centrarchids, in the southernpart of the LowerPeninsula; trout rearingstations mostly in the northernpart of the Lower Peninsulaand in the Upper Peninsula (Fig. 1). 106 AmericanFisheries Society

L E GEND BIRDS HAVING FED ON SPECIES •AT TROUTHATCHERIES OAT PONDFISHHATCHER/ES BIRDS NOT HAVING FED ON SPECIES BEING PROPAGATED []FROM TROUTWATERS FROMNON-TROUT WATERS

1.--Distribution,numbers, and classificationof kingfisherscollected at fish- cultural stations in Michigan and studied for food.

In a generalway, thesetwo typesof fish-culturalestablishments repre- sentdistinct ecological entities in that the watersof pondfishstations are impoundments(often sizable:one of severalnear Almena,Van Buren County,is about38 acres)whereas those of trout rearingestablishments are small,stream-like raceways with moderatecurrent and eitherof artificialconstruction near springsor sectionsof naturalwaters set off Kingfisherin Relation to Fish Management 107

LEGEND

] TROUTSTREAM O NON-TROUTSTREAM • LAKEOR POND

FmURE 2.--Distribution, numbers, and ecological classification of habitat for king- fishers collected on natural waters in Michigan. No birds shown in Figure 1 are repeated here. by low dams or screens.Both types are alike, however,in containing greaterconcentrations of fishesof the size particularlydesirable to king- fishersthan exist in similar areas of natural waters. Both also appear to attract more kingfishersper unit area than do natural waters. Once an individualdiscovers this great and apparentlyinexhaustible food supply 108 AmericanFisheries Society of an ideal size, it will come back to feed again and again so that it is almost necessaryto kill the bird in order to stop its depredations-- especiallyif it has young nearby requiring repeatedfeedings and a large quantity of food. It is commonpractice to kill kingfishersas a meansof controllingtheir depredationsat fish hatcheries,rearing stations,or holding ponds. It would seemthat the huge take of kingfishersat the varioushatchery sites in Michigan and elsewherein the last 15 years could ultimately be re- sponsiblefor a reduction in their numbers. A serious downward trend resultingfrom this causecould be arrestedin part by a more sane and understandinghandling of the bird's predatory activities. The critical period for protecting fish hatcheriesand feeding stations is the month of May and the first half of June. At that time the birds are seeking territories and nesting. If they once locate the huge supply of food organismsin the hatchery ponds, of exactly the length they require, it is almost impossibleto drive them away. However, if the hatchery force is alert and has guns at strategicpoints about the hatchery groundsdur- ing this interval, and if the bird is fired at when it first apears,even if there is no hope of hitting it, usuallyit will take the hint and seeka more congenialhabitat. These early birds seekingterritory make themselves mostconspicuous by constantlyuttering their shrill, rattling,challenge call. As a meansof control,it is importantthat no polesor other supports 4 feet or more in height be permitted in the vicinity of the ponds; they greatly facilitate activity of kingfisherssince they provideexcellent fishing and observationperches and thus may evenattract the birds. The general use of pole traps is to be discouragedsince so many songbirds are taken by them. Scarecrows,flashing lights, or mirrors, and even automatic detonatorsare only temporarilyeffective since kingfishers appear to be- come accustomed to them and have been seen to fish from the structures supportingsuch devices. Screening is the most effectivemeans of control and is most desirableesthetically but it is limited in its applicationto smallerbodies of water and is by no meansadapted to maximumefficiency in hatcherypractice (see Lagler, 1939, for descriptionof various types of screening). Someforesight in locatingfish hatcheriesand rearingstations may be valuable in reducingdepredations by certain bird speciesincluding the kingfisher.Because of the great numbersof migratingkingfishers along the shoresof the Great Lakes, it would be sheerfolly in the future to locate any new hatcheryor feedingstation within 5 miles of lake shores since such a locationwould be directly athwart the flight lanes of the birds (and of heronsand mergansersas well). The Harrisville Hatchery (AlconaCounty) near the shoreof Lake Huron is a casein point. Preda- tion by kingfishersalways has been heavy there and is a constantproblem. This problemis even more accentuatedat a private hatcherynear the Lake Michigan shore. The ownersof this hatchery told us that they shot and trapped more than 400 kingfishersin one spring migration. Kingfisherin Relation to Fish Management 109

In the springof 1931 there was to be seenon the groundsof this hatchery a dump of more than severalbushels of kingfisherbodies. This deplorable situationcan be eliminatedby future planning. Conversionof sectionsof headwaterstreams into rearing stationsmay invite kingfisherdepredation. The openingproduced in the streamside shrubberyby the creationof extendedpools would tend to attract and concentrate the activities of fish-eating birds. Trout Rearing Waters.--Of the many kingfishersstudied by us from waters of this type, 428 containedfood and many additional ones were empty. The stationswhere thesespecimens were obtainedand the number with food from each are shown in Figure 1. The principal food of birds at theseplaces were the salmonidsbeing propagated (Table 2) sincethey comprised80.1 percent of the total volume of food and averagedtwo per bird. Other prominent items eaten in decreasingnumerical order were aquatic insects,forage fishes (mostly minnows, muddlers, and stickle- backs), and crayfishes.

T^BLE2.--Food of the eastern belted kingfisher on trout rearing waters [Based on the stomach contents (747.8 cubic centimeters of food) of 428 individuals •]

Number of Percentage of Percentage individuals of total volume frequency of Food item each food item of food occurrence

Trout ...... 981 80.1 100.0 Other game and pan fishes.. 11 2.1 1.0 Forage fishes ...... 163 3.7 21.0 Fish remains ...... 2.5 10.3 Amphibians...... ' ' 9 1.8 1.6 Other vertebrates ...... 2 0.5 0.5 Crayfishes ...... 62 7.3 14.3 Insects ...... 347 3.2 24.1

• Detailed identifications and enumerations of the food items in these birds follow. lEgOUT:•almo gairrlnerii irirleus, 2; •. tr•tta ratio, 2; Salmo sp., 54; Salve•inus•. •on- tinalis, 213; Salmonidae, 710, OTHER GAME AND PAN FISHES: Percidae, 2; Lepomis cyanellus, 2; L. gibbosus, 1; L. m. macroehirus, 1; Centrarchidae, 5. FOr,tGE FISHES: Catostomusc. commersonnii,2; Catostomidae,1; Rhiniehthys atratulus meleagris, 1; Semotilus a. atromaculatus, 1; Margariseus margarita nachtriebi, 1; Hyborhynchus notatus, 4; Notemigonus crysoleucasauratus, 1; Notropis cornut•ts, 2; Cyprinidae, 12; Umbra limi, 4 lia ineonstans,81. CRAYFISHES:Cambarus virilis, 1; C. immunis, 1; C. propinquus,12; Cambarussp., 48. INSECTS: Orthoptera, 1; Baetinae, 27; Ephemeridae, 10; Libellula, 1; Anax }unius, 1; Aeschnaumbrosa, 2; Zygoptera, 4; Anisoptera nymphs, 17; 0donata, 5; Plecoptera, 1; Lethoeerusamericanus, 1; Belostomatidae,2; Aphidae, 1; Helicopsyche,4; Trichoptera, 80 ;Dytiscus,1; Dytiscidae,2; Carabidae,1; Staphylinidae,8; Chrysomelidae,1; Col- coptera. 29; Chironomidae,99; 8tratiomyidae, 1; Rhagionidae, 1; Diptera, 4; Form- icidae, 1: Itymenoptera, 8; unidentified, 42.

Someof the foodappearing in the stomachsof thesebirds obviously hadbeen eaten prior to theirdescent upon the hatcherywaters. This view is substantiatedby the considerablenumber of individuals(131) taken at thesestations but not containingrepresentatives of the speciesbeing propagated(Fig. 1). Suchbirds, it seems,were killed beforethey had 110 AmericanFisheries Society opportunityto prey on fishesin the rearingenclosures and for this reason they are discussedwith kingfishersfrom natural trout streams(Table 4). It is likewise possiblethat some of the kingfisherscollected at the hatcheriesand containingtrout may have obtainedthose fish from the adjacent natural waters. Since we had no way of detectingthese trout from natural waters for most birds and in view of the much lower in- cidenceof trout in the food of this bird on natural trout waters (Table 4) we believe the inherent error to be insignificant. Furthermore, trout hatcheriesusing springwater, such as the one near Almena, Van Buren Cotmty, or that at Oden, Emmet County, have practically no trout streamsnearby and the birds enteringwould thereforenot be likely to contain such fishes. Pondfish Rearing Waters.--Twenty-sevenkingaSshers are availablefor recordingthe food habits of this bird on suchwaters (Table 3). Several more that were openedwere empty and still otherscontained none of the speciesbeing propagated. The latter birdstaken at pondfishhatcheries are consideredto have flown in from adjacent non-trout waters and are discussedwith kingfishersfrom non-troutstreams (Table 5).

T^BLE3.--Foo'd of the easternbelted kingfisheron pondfishrearing waters [Based on the stomach contents (59.1 cubic centimeters of food) of 27 individuals •]

Number of Percentage of Percentage individuals of total volume frequency of Food item each •ood item of food occurrence

Pan fishes ...... 52 85.8 100.0 Forage fishes ...... 44 8.0 40.7 Fish remahm ...... 0.8 25.9 Crayfishes...... '• 2.7 18.5 Insects ...... 14 2.7 29.6

a Detailed identifications and enumerations of the food items in these birds follow. GAME AND PAN FISHES: Micropterus salmoides, 1; Micropterus d. dolomie•t, 9; black basses, 26; Lepomis gibbosus, 1; L. m. raacrochirus, 7; Lepominae, 2; Centrarchidae, 6, FORAGE FISHES: Catostomidae, 2; Notemigonus crysoleucas auratus, 1; Notropis cor- nutus, 4; Cyprinidae, 11; Umbra limi, 1; Cottus sp., 8; Eucalia inconstans, 17. GRAY- rISHr:S: Cambaru.q virilis, 1; C. immunis, 2; Cambarus sp., 3. INSECTS: Odonata, 4; Ephemeridae, 1; Coleoptera, 1; Chironomidae, 5; unidentified, 4.

The kingfisherat warm-waterfish rearing establishmentsshows a strikingresemblance (Table 3) in food habitsto thosefrom trout cul- tural stations(Table 2). The birds averageabout two per individual of the speciesbeing propagatedand thesefishes compose 85.8 percent of the total volume of food. The remainder of the organismseaten are againprincipally minnows, crayfishes, and insects. Kingfisherson Natural Waters.--Thefood habits of the kingfisheron natural watershave beensummarizd by Bent (1940) with the indication that juvenilesand adultsfeed on the mostavailable aquatic organisms of suitablesize. This generalconclusion is attestedby the followingdata from extensiveanalyses on record in the U.S. Fish and Wildlife Service (Howell, 1932). Perch, mainly Perca fiavescens,were found in 16 Kingfisherin Relation to Fish Management 111 stomachs;catfishes in 15; floundersin 13; sunfishesin 9; and carp in 3. Non-edible fishes were identified in 82 of 162 stomachs; small minnows were in 57, sculpinswere in 8, and sticklebacks,killifishes, and menhaden made up the remainder. Crayfisheswere presentin 39 individuals,com- prisingabout 16 percentof the food, frogsformed about 5 percent,and insects,4 percent. Fishesin all composedabout 75 percentof the food of the bird. In an analysisof the food of kingfishersin varioushabitats progres- sively further upstream in the Margaree and Apple River systemsof Nova Scotia, White (1937) found that in the lower reachesmany species of fish are taken, but further upriver the number of kinds becomesless until only salmon and brook trout or trout alone constitutethe food. In a more restrictedregion, though one possessinga diversefauna, namely in the vicinity of Ithaca, New York, breadth of food habit is again indi- cated (Gould, MS). a We have new data on the food of the kingfisheron trout streams,non- trout streams,and lakes (Fig. 2). Stomachsof this bird on natural watersare from two principalsources. One groupis from bona fide natural situations and was collected over most of the state in order to obtain a generalpicture of the kinds and amountsof food eaten. The other repre- sentsa more or lesschance accumulation of birds resultingfrom control operationsat fish hatcheriesand rearing stations(Fig. 1). Many birds shot or pole-trappedat fish-culturalstations contained none of the species being cultured and, as indicated,have not been previouslyreported upon in this paper.Data on thesebirds are presentedseparately in the tablesfor birds from trout streams (Table 4) and from non-trout streams (Table 5). Obviouslythese specimens taken at hatcheriesoffer significantinfor- mation on predationat fish-propagationestablishments but they appear to contribute more to an understandingof the food habits on natural waters. That suchkingfishers represent individuals taken prior to feeding in the hatcherywaters is substantiatedby their stomachcontents. The fact that these specimenswere selectedon the basis of the absenceof the speciesbeing reared from their stomach contents precludestheir being used in any way to indicate nonexistenceof predationin natural waters on the fish speciesprimarily concerned.They do afford, however, an amplification of the list of other organismspreyed upon by king- a A condensation of Gould's original records of analyses, made by the U.S. Fish and Wildlife Service, is reproduced here with the permission of Harrison F. Lewis, in whose custody the manuscript lies, since it is likely that it will remain unpublished for some time. The name of each food item and the number of times the item occurred in the 25 stomachs examined are given. For the fishes, the list is in decreasing order of number of individuals of each item. FisH: Pomolobus sp., 5; Salmo trutta fario, 9; Catostorages c. commersonnii, 14: Cyprinidae, 12: Cyprinus carpio, 1: Leucosomus cor- poralis, I; Semotilus a. atromaculat•ts, 15; Exoglossum maxillinglm, 2; Rhinichthys a. atratulus, 7; Hybopsis sp., 2; Hybognath•ts sp., I; Notropis sp., 4 • Notropls cornuttts, 9 Amei•trus sp., 4: Fund•tlus diaphan•t.q, 1; Perca )•lavesccns, 2: Bolcosoma nigr,rn, 4 Micropterus salmoides, 5; Lepomis sp., 2; Lepomis gibbosus, 4 • Cottus sp., 1. Rana sp., 6. REPTILES1 Snakes, 2. INSECTS1 Anisoptera. 1: Aeschnidae. 1: Hydrous triangularis, 1 • Berosus sp., 1: Hallplus sp., 1; Elateridae, 1: Chrysomelidae. 1; Cur- culionidae, 1; Camponotus herculeanus, 1; Hymenoptera, 1. • Cambarus sp., 19. 112 AmericanFisheries Society fishersand corroboratethe evidenceafforded by the bona fide samples from natural watersso well that additionalsacrifice of birds for foodstudy was unneccessary. Trout Streams.--Trout streamsof a wide variety of sizes and kinds are found in Michigan. As has been indicated,fishes in all except the smallestof them are exposedto predationby kingfishers.That trout are eatencannot be denied; our findings(Table 4) showan incidenceof ap- proximately one for every two birds. Almost two-thirds of the bulk of the food and most of the food organismsby number, however,are made up of forage fishesand crayfishes.It is suspectedthat the vulner- ability of foragefishes varies greatly from streamto stream. Furthermore, it is generallyconceded that such fishesare less wary than trout and that they are most abundant in the open, sunlit parts of the streams (Hankinson, 1923) which kingfishersfrequent whereasthe trout are in

T^• 4.--Food of the eastern belted kingfisher on trout streams [Based on the stomach contents (266.7 cubic centimeters of food) of 92 individuals collected on natural trout streams x and on the stomach contents (186.1 cubic centimeters of food) of 131 individuals taken at fish hatcheries but representing food habits on natural trout waters 2]

Number of individ- Percentage of total Percentage frequenc, uals of each food volume of food of occurrence item Food item From From From natural From natural From natural From trout hatch- trout hatch- trout hatch- streams eries streams eries streams eries Game and [Trout ... 40 . 29.8 35.9 panfishes [ Others .. 14 iJ• 13.0 • 13.0 • Forage fishes ...... 68 122 15.0 15.4 35.9 39.7 Fish remains ...... 0.9 0.9 14.1 9.2 Frogs ...... ' i ' i2 Trace 13.2 1.1 7.6 Crayfishes ...... 58 56 40.7 50.5 53.3 39.7 Insects ...... 8 91 0.6 15.5 8.7 34.4 x Detailed identifications and enumerations of the food items in birds collected on natur- al trout streams follow. GAME AND PAN FISHES: Salmo gairdnerii irideus, 6; S. trutta fario, 1; Salmo sp., 3; Salvelinus •. /ontinalis, 5; Salmonidac, 25; Ameiuridae, 1; Esox lucius, 1; Perca /lavescens, 5; l•epomis m. macrochirus, 2; Centrarchidae, 5. For•AoE FIs•{Es: Moxostoma sp., 1; Catostomidae, 2; Cyprinidae, 6; Umbra limi, 1; Etheosto- mattnae, 5; Cottus sp., 22; Eucalia inconstans, 29. FR00S: Rana sp., 1. CRAYFISHES: Cambarus virilis, 14; C. propinquus, 20; Cambarus sp., 24. INSECTS: Anisoptera, 3; Stratiomyidae, 1; unidentified, 4. 2 Detailed identification and enumerations of the food items in birds collected at hatch- eries but representing food habits on natural trout waters follow. GAME AND PAN FISItrS: Perca flarescerts, 2; Micropterus d. dolomieu, 1; Lepomis cyanellus, 4; Cen- trarchidae, 7. FORAGE FISHES: Catostomus c. commersonnii, 2; Catostomtdae, 3; Rhin- ichthys atratulus meleagris, 1; Semotilus a. atromaculatus, 2; Hyborhynchus notatus, 9; Notropis cornutus, 5: Cyprinidae, 24; Umbra limi, 1; Fundulus diaphanus menona, 7; Etheostomatinae, 5; Cottus cognatus, 1; Cottus sp., 34; Eucalia inconstans, 26. FROGS: Rana sp., 12. CRAYFISHES: Cambarus virilis, 3; C. propinquus, 24; C. immunis, 6; •. diogenes, 1; Cambarus sp., 22. INSECTS: Orthoptera, 1; Xnax }unius, 2; Epiaeschna sp., 1; Anisoptera nymphs, 19: 0donata, 1; Lethocerus americanus, 1; l•ethocerus sp., 2; Belostomatidae, 2; Cicadellidae, 1; Aphldae, 1; Ephemeridae nymph, 1 • Dy•iseus Har- risii, 2; Dyt•scus dauricus, 1; Dytiscus sp., 1; Dytiscidae, 15; Gyrinidae, 1; Haliplus sp., 1; Coleoptera, 1; Trichoptera larvae, 27; Diptera eggs, 6; unidentified, 2. One water mite, Acarina, was also found and is included in this category for convenience. Kingfisherin Relation to Fish Management 113

TABLE 5.--Food of the eastern belted kingfisher on non-trout streams [Based on the stomach contents (67.7 cubic centimeters of food) of 22 individuals col- lected on natural waters • and on the stomach contents (157.8 cubic centimeters of food) of 95 individuals taken at fish hatcheries but representing food habits on natural waters •] Number of individ- Percentage of total Percentage frequenc.• uals of each food volume of food of occurrence item Food item From From From natural From natural From natural From non-trout hatch- non-trout hatch- non-trout hatch- streams eries streams eries streams eries

Game and pan fishes 5 7 10.5 3.4 22.7 6.3 Forage fishes ...... 12 89 31.3 17.1 40.9 35.8 Other fishes ...... 1 .. 16.2 4.5 Fish remains ...... 0.1 "•.• 13.6 '•'.3 Frogs ...... ' • ... 9.7 . .. 2.1 Other vertebrates ... 2 1.3 2.1 •rayfishes ...... i• 32 3'9'.6 18.8 5'9'.i 27.4 Insects ...... 4 57 2.2 49.0 18.2 42.1

•Detailed identifications and enumerations of the food items in these birds follow. GAME ANDPAN FISHES: Perca flavescens,2; Centrarchidae, 3. FORAGEFISHES: Catos- tomidac, 1; Hyborhynchus notatus, 3; Notemigonus crysoleucas auratus, 1; Notropis atherinoides, 2; iV. hudsonius, 1: Cyprinidae, 2; Umbra lim{, 1; Cottus sp., 1. OTHER FISHES: ESOp vermiculatus, 1. CRAYFISHES;Cambarns propinqu•ts, 14; Cambarns sp., 4. INSECTS:Belostomatidae, 2; Ephemeridae, 2. 2 Detailed identifications and enumerations of the food items in birds collected at hatch- eries but representing food habits on natural non-trout waters follow. GAME AND PAN FISHES: Perca flavescens, 3; Lepomis cyanellus, 2; L. m. macrochirus, 2. FORAGE FISHES: CatostomusC. Commersonnig,2; Catostomidae,i; Hyborhynchusnotatus, 12 l•o•ropis corn•tus, 50; Cyprinidae, 10: Umbra limi, 4; Fungulus giaphan•s menohs, i; Etheostomatinae,5; Coitus b. bairgii, 2; Cottus sp., 2. FROGS:Rana sp., 6. OTHER Tl•BRATES;Turtle egg, 1. CRAYFSHES: Cambarns virilis, 6; C. pro•inquus, 5; C. immunis, 7; Cambarussp., 15. INSECTS:Anax •unius, 4 • Aeschnldae,2; Anispotera nymphs, 11 Lethocerus americanus, 8; Lethocerus sp., 3; Benacus sp., 1: Dytiscus harrisii, 1; Dy•iscus sp., 13; Ac•lius sp., 2; Dytiscidae, 2; Coleoptera, 1; Chironomldaelarvae, 2; unidentified, 1. the darker, deeper, and more shelteredportions. Experimentson the effectsof predationby White (1937) led to the conclusionthat competi- tion and cannibalismwere a greater menaceto young trout than attacks by birds including kingfishersand membersof the heron family. A part of the food is composedof game and pan fishesother than trout, which have no recognizablevalue for sport in these situations. At least someof these fishesare strays from lakes situated in the drainage systemsof the streamsand are not regularinhabitants of Michigan trout waters. Some that are eaten, for examplethe predaciousnorthern pike (Esox lucius), are liabilities in trout streamsas indicatedby Hankinson (1923). Crayfishes,forage fishes, and insectsare utilized in commonby trout (Metzelaar, 1929; Needham, 1938; Lagler and Lagler, 1944) and by kingfishers.It has further been suggested(Hankinson, 1923) that two of the forage species (Semotilus a. atromaculatusand Notropis cornutus) eaten by kingfishersmay be incompatiblewith native trout sincethey offer competitionfor food. The interrelationshipsof predator and prey are here, as always, very 114 AmericanFisheries Society

complicated,and the exact statusof the kingfisherin relationto trout productionin natural waters is thereforeundecipherable on the basis of the data thus far obtained. Non-troutStreams.--The considerable importance of crayfishesas seen in the food of kingfisherson trout streams(Table 4) is also somewhat evidentin that of specimensfrom runningwaters that are not inhabited by trout (Table 5). To someextent the largenumbers of crayfishmay be the resultof the numericaldominance of youngbirds in our collections. Insectsare more importantin birds collectedat hatcheries,but not con- taininghatchery fish, than in thosefrom naturalwaters. Forage fishes againare the numericallydominant food organismsand, amongthe fish, they exceedthe othersby far in both numbersand bulk. This preponder- ance of forage fish, of course,reflects the leading numericalabundance which is usual for forage speciesin fish populationsin streamsof this kind. Vertebratesas a whole appear to be of greater significanceto the kingfisheras food on thesewaters than are the arthropods. Lakes.--A differencein the foodhabits of kingfishersin standingwaters from thosein runningwaters is suggestedby our data (Table 6). Cray- fishesare not a food item of major importanceto this bird in lacustrine situations. This fact would seem to indicate a lesser abundance of these decapodsper unit area of fishablelentic watersthan lotic ones,or it might suggesta greater proportionateabundance of forage fishesto crayfishes in the former sincemuch of the numericaland volumetricloss for cray- fisheshas been gainedby forage fisheswhich are by far the most im- portant item of sustenance.Insects are secondby volume in apparent importancebut game and pan fishesare probably of greater actual sig- nificance for their volume would exceed that of the insects if both were

T^BLE 6.--Food of the eastern belted kingfisher on lakes [Based on the stomach contents (92.1 cubic eentilneters of food) of 45 individualsX] Number of Percentage of Percentage individuals of total volume frequency of Food item each food item of food occurrence

•ame and pan fishes ...... 29 17.5 20.0 Forage fishes ...... 78 49.1 51.1 Other fishes ...... 2 2.0 4.4 Fish remains ...... 0.9 22.2 Frogs ...... ' 6 2.3 13.3 Crayfishes ...... 18 7.4 26.7 Insects ...... 15 21.0 37.8

• Detailed identifications and enumerations of the food items in these birds follow. GAME AND PAN FISHES i Perca flarescerts, 17; Micropterus d. dolomieu, 1; Lepomis cyanellas, 1; L. gibbosus, 1: L. m. •acrochirus, 3; Centrarchidae, 6. •q•ORAGEFISHES: Catostomidae, 1: Cyprinus carpio, 1; tlyborhynchus notatus, 49; Notemigomts cryso- leucas auratus, I; Notropis hudsonius, I; Cyprinidae, 7; Umbra limi, I: Percina cap- rodes semifasciata, 1; Etheostomatinae, 7: Labidesthes s. sicc•tlus, 7; Cottus sp., 11 Eitcalia inconstans. 1. OTH•]R FISItES: Esocidae. 1; Lota lota mac•tlosa, 1. FROaS: Rana Sp., 6. CRAYEISHES; Cambarus virilis, 8; C. immunis. 1: C. propinquu8, 3:Cambaru8 sp., 11. INSECTS: Anax ]unius nymph, I; Anisoptera, 3; Belostomatidae, 1; Dytiscidae, 2; unidentified, 8. Kingfisherin Relation to Fish Management 115 restoredto their originalbulk on ingestion.The numberof specimensof kingfisherson which theseobservations are basedis, however,small. PRESENT STATUS OI2'THE KINGFISHER PROBLEM Several commentshave been publishedon the relationshipsbetween fish-eatingbirds, includingthe kingfisher,and fish. Vladykov (1943) alone has attemptedto presentin one paper an unbiasedsummary of the views of the severalauthors and has listed the three categoriesof inter- pretersof the role of thesebirds. (1) ultra-preservationists;(2) exponents of stringentcontrol; (3) thosewho take the middle course. In general one would find over-enthusiasticlaymen or thosedirectly dependentupon them for incomein the first two categories,and, thosewho try to apply a more scientificmethod in arriving at their conclusionsin the third. Obviously,then, our data constituteammunition for any of thesegroups. Ultra-preservationistswill emphasizethe broad diet of the kingfisher,as here attested,and will continueagainst control. Fish-culturistsand ardent anglerswill regard as of primary importancethe feedingof the bird on populationsof their selectedspecies. Neither group will, however,be fully able to justify its caseon the basisof what is now known. Stagger- ing and damning estimatesof the total numbersof fish consumedmay be made (e.g., Sinker, 1883; Sullivan,1912; White, 1936), but unlessthese estimatesare interpretedin light of existingpopulations of both predators and prey in specificareas, they are obviouslyinvalid and worsethan mis- leading. The conclusionsthat we draw from our study are very limited and not at all trenchant. Becauseof its predatorypotential of more than 50 small fingerlingsper day (determinedexperimentally by White, 1936) and the establishedfact that on fish rearing waters it consumesquantities of the fishesbeing propogated,the kingfisheris incompatiblewith fish- cultural practicesof the presenttime. In recognitionof this difficulty, may not organizationsfostering the ultra-preservationistattitude be ex- pectedto sharein the extra costof protectingfishes at hatcheriesby non- lethal meansrather than by shotgunor pole-trap? On natural waters, with the possibleexception of spawningareas or of nurserygrounds such as headwater trout streams,we concludethat the kingfisher is relatively innocuousas a fish predator. Such prey in the form of preferredfish speciesor their foodorganisms which kingfishers consume on mostnatural waters can easily be consideredas a small tax, gladly given, for the esthetic virtues of this bird. That there are possiblebenefits as well as undesirableeffects of preda- tion has often been stated (e.g., Lagler, 1944). A detailed discussionof this matter doesnot appearto be desirablehere. LITERATURE CITED BE•T, A. C. 1940. Life histories of North American cuckoos, goatsuckers, hummingbirds and their allies. Bull. U.S. Nat. Mus., No. 176, viii q- 506 pp. 116 American Fisherles Society

DE L^ TORREB•;ENO, J. R., JR. 1936.Some observations on the kingfisher.Fish Culture(N.Y. Cons.Dept.), Vol. 2, No. 9, pp. 11-14. HANKINSON,T. L. 1923.The habitatof the brooktrout in Michigan.Pap. Mich. Acad.Sci., Arts, and Lett., Vol. 2, 1922, pp. 197-205. HOOVER,EARL E. 1936. Fish-eating birds. Pros. Fish-Cult., No. 21, pp. 21-22. HOWELL,A. H. 1932.Florida bird life. Dept. Game and Fresh Water Fish, Tallahassee,Fla., xxiv q- 579 pp. LACLER,KARL F. 1939. The controlof fish predatorsat hacheriesand rearingstations. Jour. Wild- life Man., Vol. 3, No. 3, pp. 169-179. 1943. Neglectedsources of data. Wilson Bull., Vol. 55, No. 2, p. 132. 1944.Problems of competitionand predation.Trans. Ninth N. Am. WildlifeConf., pp. 212-219. LAGLER,KARL F., and MARY JANE LAGLER 1944. Natural enemiesof crayfishesin Michigan. Pap. Mich. Acad. Sci.,Arts, and Lett., Vol. 29, (1943), pp. 293-303. LANGLOI$,T. H. 1936.A studyof the small-mouthbass, Micropterus dolo.mieu (Lac•p•de) in rear- ing pondsin Ohio. Ohio Biol. Surv. Bull. No. 33, Obio State Univ. Stud.,Vol. 6, No. 4, pp. 189-225.

METZELAAR,JAN 1929. The food of the trout in Michigan. Trans. Am. Fish. Soc.,Vol. 59, pp. 146- 152.

NEED•^•, P^•L R. 1938. Trout streams. ComstockPublishing Company, Inc., Jlthaca,N.Y., x q- 233 pp.

S^LYER,J. CL•, II, and KaRL F. L^CLER 1940. The food and habits of the American merganserduring winter in Michigan, consideredin relation to fish management. Jour. Wildlife Man., Vol. 4, No. 2, pp. 186-219.

SINKER,WOODEN 1883. About kingfishers.Forest and Stream,Vol. 21, No. 10, pp. 188-189.

SULLIVAN, R. H. 1912. Economicvalue of bird life. Agri. Educ. (KansasState Agri. Coll.), Vol 7, pp. 1-47.

TRAUTMAN,MILTON B. 1940.The birdsof BuckeyeLake, Ohio. Univ. Mich. Mus. Zool.,Misc. Publ. No. 44, 466 pp.

Vr^DVKOV,V^DI• D. 1943. Relation between fish and fish-eating birds. Can. Field Nat., VoL 57, pp. 124-132. Kingfisherin Relation to Fish Management 117

WX•XTE,H. C. 1936. The food of kingfishersand merganserson the Margaree River, Nova Scotia. Jour. Biol. Bd. Can., Vol. 2, No. 3, pp. 299-309. 1937. Local feeding of kingfishers and mergansers.Jour. Biol. Bd. Can., Vol. 3• No. 4, pp. 323-338. 1939. Change in gastric digestionof kingfisherswith development. Am. Nat., Vol. 73, No. 745, pp. 188-190.