51' ERICKSON,Sarcocystis in Birds L[' AukOct.

SARCOCYSTIS IN BIRDS •

BY ARNOLD B. ERICKSON

Plate to THe.protozoan genus Sarcocystis, which is parasiticin the muscles of vertebrates,has been reported from eightorders, thirteen families, nineteengenera, and twenty species of birds. In Europeit hasbeen found in the DomesticFowl and Duck, the Wild Duck, and doubt- fully in the EuropeanBlackbird (Babudieri,1932, believesthat Sarcocystisturdi, reportedfor the EuropeanBlackbird, is an intestinal coccidian).In Africa it has been recoveredfrom one species--the Red-faced African Mouse-bird. All other references to its occurrence in birds relate to the Americas. In North America it has been re- ported as occurringin eleven speciesof birds; in South America, six species;in CentralAmerica, one species.As new hostrecords we may add Wilson'sSnipe (Capelladelicata), the AmericanPintail (Dafila acuta tzitzihoa),the Blue-wingedTeal (Querqueduladiscors), and the Gadwall (Chaulelasmusstreperus). The snipe was collectedsouth of Minneapolis,Minnesota, on Oc- tober 17, 1939, by R. M. Berthel and Arnold B. Erickson. When the bird was flushed,nothing abnormal was noted in its flight; but on skinning,it wasfound that the musclesof the breast,neck (Plate 10), legs,and the chin containedmany white cystsof Sarcocystis.They were especiallyprominent in the musclesof the neck and the tibiae, where they lay closeto the surface. In the pectoralmuscles, however, they were more deeply imbeddedand occurredall the way to the sternum. The spindle-shapedcysts averaged 2.69 mm. in length by 0.441 mm. in width at the middle. The smallest one was 2.18 mm. long by 0.469mm. at the middle;one end was0.486 mm. wide and the other 0.405 mm. wide. The largest one was 3.24 mm. long by 0.437 mm. wide at the middle; one end was 0.340 mm. wide and the other 0.324 min. wide. It is evident that there is a tendencyfor one end of the cystto be slightlywider than the other. The minute spores(Plate 10) measured8.15 micrain lengthby 1.63micra in width. Neither the sporesnor the cystsdiffer enoughfrom those of Sarcocystisrileyi to warrantthe descriptionof a new species. The Pintail was captured in a weakenedcondition along with several hundred Mallards at Heron Lake, Minnesota, on December

x Paper No. x775 Scientific Journal Series, Minnesota Agricultural Experiment Station, St. Paul. In cooperation with the Minnesota Conservation Department, Division of Game and Fish. ThE AuK, VOL. 57 PLATE 10

Sarcocystisrileyi •N g•Cl• OF •VlLSON'SSNIPE.

CRf).•S•$ECT1ONOF S•11•½o½y$•i$ rileyi FROM•ILSO1N'S SNIPE. SHOWINGMINUTE SPORES WITHIN THE TERETECYST. • 180. Vol.•94o•] a EVaCKSON,$arcocystis in Birds 515

28, 1939,and was sent to theDivision of EconomicZoology, University of Minnesota,where it died on January2, 1940. There wasno evi- dencethat it had succumbedto lead poisoning,as manyof the Mal- lardssubsequently did. The fact, however,that it was heavilyin- festedwith Sarcocystisrileyi in all of the skeletalmuscles and the heart may have been a factor contributingto its death. It wasmuch emaciated,weighing but 525 grams. The Blue-wingedTeal wasgiven to Dr. Alan Morrow of Fergus Falls,Minnesota, on October1, 1959,by a hunter,and on skinning, white streakswere noted throughoutthe muscles. The bird wassub- mittedto Dr. R. Fenstermacherof theDivision of VeterinaryMedicine, Universityof Minnesota,who diagnosedthe infectionas Sarcocystis rileyi. Somedoubt developed as to whetherthe bird may havebeen a Green-wingedTeal, but Dr. Fenstermacherstates that the carcass was the sizeof a Blue-wingedTeal. The Gadwall was found dead at Clayton Lake, Martin County, Minnesota,on April 6, 1940, by Mr. C. E. Carlson. The bird had been shot through the left breastby a rifle bullet. On skinning,it wasfound to havea light infectionof Sarcocystisrileyi in the pectoral muscles. Sections of infected muscle were made which substantiated the field diagnosis. The frequencyof occurrenceof Sarcocystisin birds is not known. The infection is commonenough, however, so that a few casescome to the attentionof parasitologistsand conservationdepartments each year. Of 279 ducksof eighteenspecies examined in the Division of EconomicZoology, University of Minnesota,from 1951 to November, 1959, eight or 2.86% were infectedwith Sarcocystisrileyi. Six of these were Mallards and two were Shovellets. One of the latter, constitutinga recent record, was taken on November 11, 1959, at Wheaton, Minnesota, and submitted by Mr. C. Gordon Fredine. The autopsyrecords of the MinnesotaWildlife DiseaseInvestiga- tion indicate that of 45 ducksof eight speciescollected from 1955 to March 27, 1959,three or 6.97% were infectedwith Sarcocystisrileyi. All three were Mallards taken in Minnesota. Since a large propor- tion of the ducks that come in for examination are sick, diseased, or crippled,it is probablethat the percentageof infectionof Sarcocystis heregiven is muchgreater than in the duckpopulation as a whole. Previous to 1951 there were four records of the occurrence of Sarcocystisin ducksin Minnesota. Two of thesewere collectedin 1919 and two in 1950 by Dr. W. A. Riley, Chief of the Division of Entomologyand EconomicZoology, University of Minnesota,who reportedthem in 1931. 5 16 ERICKSON,$arcocystis in Birds L[' AukOct. Duringthe 1935hunting season there were three records brought to theattention of Dr. GustavA. Swanson,then Biologist with theMin- nesotaConservation Department. One was from the Mississippi River belowSpring Lake, Minnesota,and a secondwas received from the Upper Mississippi'Wildlife Refuge near Winona,Minnesota. The third reportwas from GenevaLake in FreebornCounty, Min- nesota,where Warden Herman Baudletreported that six Mallards from a bagof duckstaken in oneday were all heavilyinfested. One of these ducks was submitted for examination. A week later three more infected Mallards were taken at the same lake. To dateall recordsof Sarcocystisin duckspertain to the puddleor dabbling ducks. A possibleexplanation for this may be that the ducksbecome infected either by eating fleshcontaining ripe cystsor by eating food contaminatedwith excrementof an infectedanimal; and sincethe pond duckscharacteristically feed in shallowwater, they would probablybe much more exposedto fecal contaminationthan the diving ducksthat feed in deeperwater. Also they are moreprone to feed in cornand stubblefields than are the diving ducksand would againbe moreexposed to fecalcontamination. In order to make the published information on avian hosts of Sarcocystismore availablethe followingoutline has been prepared. The bird names,as usedin earlier lists, have been changedto con- form with the A. O. U. 'Check-list,'1931, and 'Catalogueof Birds of the Americas'by CharlesB. Cory and CharlesE. Hellmayr,and the ordinal and family nameshave been includedas a further source of information. Authorities for hosts and localities have in each case been listed. AVlAN Hosxs oF x•E GENus Sarcocystis Order ANSERIFORMES Family Anatidae •lnas platyrhynchosplatyrhynchos (Mallard).--$arcocystis rileyi (Stiles,1893). C. V. Riley, 1869;Leidy, 1875;Stiles, 1895; Crawley, 1911; W. A. Riley, 1951; Greenand Shillinger,1935, 1955; Shillinger and Wetmore,1958; this paper. United States. "Wild Duck" Anas?platyrhynchos platyrhynchos (Mallard).--$arcocystis anatina Krauseand Goranoff,1935. Bulgaria. Anas platyrhynchosdomesticus (Domestic Mallard).--$arcocystis sp., v. Betegh and Dorcich, 1912. Germany. $arcocystisrileyi Hall, 1925. United States, Anas rubripes (Black Duck).--Sarcocystisrileyi. Mathews, 1950; Gower, 1938. United States. Chaulelasmusstreperus (Gadwall).--$arcocystisrileyi. This paper. United States. Dafila acuta tzitzihoa (American Pintail).--Sarcocystisrileyi. This paper. United States. Vol.57] ERICKSON,$arcocysti$ in Birds 5 17 194o a

"Teal"? Querquedula discors (Blue-wingedTeal).--Sarcocystis rileyi. This paper. United States. Spatulaclypeata (Shoveller).--$arcoc•tis rilcyi (Stiles,1893); W. A. Riley, 1931. This paper. United States. Order FALCONIFORMES Family Cathartidae Cathartesaura scptentrionalis(Turkey Vulture).--Sarcocystissp., Osterudand Bascorn, 1928. United States. Family Falconidae. Leucopternissp. (Ghiesbrecht'sHawk).--Sarcocystis sp., Darling, 1915. Panama. Order GALL•FORMF_S Family Phasianidae Gallusgallus (Domestic Fowl).-Sarcocystis sp.,KiJhn, 1865. Germany. Sarco- cystissp., Hames (cited in Stiles, 1893); Stiles, 1894. United States. Sarco- cystishorwathi v. Rfitz, 1908. Hungary. Sarcocystisgallinarura Krause and Goranoff, 1933. Bulgaria. Order GRUIFORME$ Family Rallidae Aramidessaracura (SaracuraWood-rail).--Sarcocystis aramidis Splendore, 1907. Brazil. Order CHARADRIIFO•MES Family Scolopacidae Capelladelicata (Wilson's Snipe).--Sarcocystis rileyi. This paper. United States. Order CUCV•IFO•MES Family Cuculidae Guira guira (Guira Cuckoo).--Sarcocystiscorderoi Vogelsang, 1929. Uruguay. Order COLIIFORMES Family Coliidae Coliusindicus [• erythromelon](Red-faced African Mouse-bird).--Sarcocystis colii Fantham, 1913. Africa. Order PASSERIFORMES Family Turdidae Turdusraerula (EuropeanBlackbird).--Sarcocystis turdi Brumpt. Rivolta, 1874 (citedby Biitschli);Brumpt, 1913. Europe. Family Compsothlypidae ½ompsothlypis[• Parula] pitiayumi (Olive-backedWarbler).--Sarcocystis sp., Barrows(cited by Stiles,1893). Argentina. Setophagaruticilla (American Redstart).--$arcocystissetophagae Crawley. Stiles, 1894; Crawley, 1914. United States. Family Ploceidae Passerdomesticus (English Sparrow).--Sarcocystiscorderoi Vogelsang,1929. Uruguay. Family Icteridae Molothrusbonariensis (Shiny Cowbird).--Sarcocystis debonel Vogelsang, 1929. Uruguay. Family Fringillidae Hedymelesludovicianus (Rose-breasted Orosbeak).--Sarcocystis falcatula Stiles, 189•. Canada. Myospizahuraeralis raeridiana [=ttmraodrarnus raanirabe] (Venezuelan Grass- 515 •IcK•ON,Sarcocystis in Birds œi' AukOct.

hopper Warbler).--Sarcocystis[•Miescheria] ammodramiSplendore, 1907. Brazil. Ornithologists,taxidermists, sportsmen, and otherswho skin or handlebirds can help increasethe knowledgeof the geographicand speciesdistribution of Sarcocystisby watchingfor the white tube- shapedcysts in the skeletalmuscles. I wishto thankDr. W. A. Riley for readingthe manuscriptand for many valuable suggestions;Dr. Gustav A. Swanson,Dr. R. Fenster- macher,and Dr. R. G. Green,all of the Universityof Minnesota,for permissionto includetheir unpublisheddata; Mr. C. GordonFredine, Biologist with the Minnesota ConservationDepartment, for sub- mitting severalducks infected with Sarcocystis;Mr. C. E. Carlson of the Universityof Minnesotafor submittingan infectedGadwall; and Dr. Alexander Wetmore and Dr. C. C. Gregg for assistancewith cer- tain bird names.

LITERATVP• C1TZD AMERICAN ORNITHOLOGISTS' UNION 1931. Check-list of North American birds. BABUDIERI, BRENNO 1932. I sarcosporidie le sarcosporidiosi.Archiv f. Protistenkunde,76: 421-580. V. BETEGH,L., AND DORCICH, P. 1912. Studicniibcr Sarcosporidicn.Ccntralbl. f. Backtcriol.u. Parasitcnk.,1 Abt. Orig., 63: 387-390. BRUMPT, F,MILIE 1913.Pr•cis dc Parasitologic.2d cdition,p. 110. BOTSCHLI, O. 1880--82. Protozoa. In H. G. Bronn's Klassenund Ordnungen des Tier-Reichs, vol. 1. CORY, CHARLESB., AND HELLMAYR, CHARLESF.. 1918--38. Catalogueof birds of the Americas. Field Museumof Natural History, zool. set., vol. CRAWLEY, HOWARD 1911. Observationsou Sarcocystisrileyi. Proc. Acad. Nat. Sci., Philadelphia,63: 457-468. 1914. Two new Sarcosporidia.Proc. Acad. Nat. Sci., Philadelphia, 66: 214-218. DARLI'NO, S. T. 1915. Sarco•poridiaencountered in Panama. Journ. Parasitol., 1: 113-120. FANTHAM, H. B. 1913. Sarcocystiscorn n. sp. a sarcosporidiauoccurring in the Red-facedAfrican Mouse Bird, Colius ½rytl•romdon. Proc. Cambridge Philos. $oc., 17: 221-224. GOWER,GARL 1938. A new ho•t and localityrecord for Sarcocystisril½yi (Stiles•1893). Journ. Parasitol., 24: 378. Vol.194oo?'l a EglCKSON,Sarcocystis in Birds 519

GREEN,R. G., ANDSHILLINGER, j. •.. 1933. Pathologicalreport on miscellaneousspecimens examined. MinnesotaWild- life DiseaseInvestigation, October, 1933, p. 7 (mimeographed). 1935. Pathologicalreport on miscellaneousspecimens. Minnesota Wildlife DiseaseInvestigation, October, 1935, pp. 7•-75 (mimeographed). HALL, R. P. 1925. Sarcocystisrileyi from the domesticduck. Journ. Parasitol.,lh 217. KRAUSE,C., ANO GORANOFF,S. 1933. Ueber Sarkosporidiosisbei Huhn und Wilderite. Zeitschr.Infektionskr. und Hyg. Haustiere,43: 261-279. KOHN,J. 1865. Untersuchungenfiber die Trichinenkrankheitder Schweine.Inst. d. Univ. Halle, 68. LEIDY,JOSEPH 1875. On Psorospermsin a Mallard Duck. Proc. Acad. Nat. Sci., Philadelphia, 27: 125. MATHEWS,F. P. 1930. Sarcosporidlosisin a duck. Journ. Amer. Vet. Med. Assoc.,76: 705-707. OSTERUD,H. L., AND BASGOM,K. F. 1928. Noteson the occurrenceof Sarcocystis.Science, 67: 531-532. •TZ, I. 1908. Az izmokban 61osk6d6v6g16nyekr61 es a Magyar fauna ban e16fordu16 fajaikr6d. Allattani k6zlem•nyek,7: 177. RxLEv, C. V. 1869. A measlywild duck. Amer. Entomologist,1: 89. RILEY, WILLIAM A. 1931. Sarcosporidiosisin ducks. Parasitology,23: 282-285. 1931a.Sarcocystis a parasitethat dines on duck flesh. Fins, Feathersand Fur, no. 95, Mar. 1931, p. 22. SHILLINGER,J. F,., AND WETMORE, P. W. 1938. Sarcosporidiosisa protozoandisease of wildlife. Trans. Third North American Wildlife Conference, 898-901. SPLENDORE,g. 1907. Breva nota sopraalcuni sarcosporididi uccellibrasiliani. Rev. da Soc. Scient. de S5o Paulo, 8:115. STILF•, C. W. 1893. On the presenceof Sarcosporidiain birds. U.S. Dept. Agric.,Bureau Animal Industry, bull. no. 3, pp. 79-88. 1894. New Americanfinds of Sarcosporidia.Vet. Mag., Philadelphia,1: 728. VOGELSANG,]•. G. 1929. Beitr•igezur Kenntnisder ParasitenfaunaUruguays. Sarkosporidien bei V6geln.Zentralbl. f. Bakt. und Parasitenkunde,1 Orig., 113:206-208. Division of EconomicZoology Universityof Minnesota St. Paul, Minnesota