University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln

Faculty Publications from the Harold W. Manter Laboratory of Parasitology Parasitology, Harold W. Manter Laboratory of

4-2003

Survey for and Haemosporidia in the Lesser Prairie Chicken (Typanuchus pallidicinctus) from New Mexico with the Description of a New Species

B. H. Smith University of New Mexico, [email protected]

Donald Duszynski University of New Mexico, [email protected]

K. Johnson University of New Mexico

Follow this and additional works at: https://digitalcommons.unl.edu/parasitologyfacpubs

Part of the Parasitology Commons

Smith, B. H.; Duszynski, Donald; and Johnson, K., "Survey for Coccidia and Haemosporidia in the Lesser Prairie Chicken (Typanuchus pallidicinctus) from New Mexico with the Description of a New Eimeria Species" (2003). Faculty Publications from the Harold W. Manter Laboratory of Parasitology. 194. https://digitalcommons.unl.edu/parasitologyfacpubs/194

This Article is brought to you for free and open access by the Parasitology, Harold W. Manter Laboratory of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Faculty Publications from the Harold W. Manter Laboratory of Parasitology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. JOURNAL OF WILDLIFE DISEASES Thursday May 22 2003 03:14 PM jwdi 39_212 Mp_347 Allen Press • DTPro System File # 12em

Journal of Wildlife Diseases, 39(2), 2003, pp. 347–353 ᭧ Wildlife Disease Association 2003

SURVEY FOR COCCIDIA AND HAEMOSPORIDIA IN THE LESSER PRAIRIE-CHICKEN (TYMPANUCHUS PALLIDICINCTUS) FROM NEW MEXICO WITH DESCRIPTION OF A NEW EIMERIA SPECIES

B. H. Smith,1,2 D. W. Duszynski,1 and K. Johnson1 1 Department of Biology, The University of New Mexico, Albuquerque, New Mexico 87131, USA 2 Corresponding author (email: [email protected])

ABSTRACT: Blood films and fecal samples of the lesser prairie-chicken (Tympanuchus pallidi- cinctus) were examined for parasites when we surveyed specimens captured during a radio- tracking study conducted in Chaves County, New Mexico (USA). All were captured on the Caprock Wildlife Habitat Management Area, administered by the Bureau of Land Management. Samples were collected in late March, April, and early May 1998–2000. Oocysts were detected in five of 64 (8%) birds sampled and, upon sporulation, were determined to be an Eimeria species. This is the first eimerian reported from the lesser prairie-chicken and is described here as a new species. Sporulated oocysts are ellipsoidal, 27.1ϫ22.7 (22–32ϫ18–26) ␮m, with micropyle absent, but oocyst residuum and polar granule present. Sporocysts are ovoidal, 11.9ϫ7.8 (10–14ϫ6–10); a Stieda body, and sporocyst residuum are present, as is a small, indistinct substieda body. In- spection of blood smears revealed four cases of infection of 32 (13%) individuals sampled. The characteristics of this plasmodiid are consistent with the description of Plasmodium (Giovannolaia) pedioecetii, previously found in T. pallidicinctus (Stabler, 1978). Key words: Eimeria tympanuchi, lesser prairie-chicken, new species, Plasmodium (Giovan- nolaia) pedioecetii, Tympanuchus pallidicinctus.

INTRODUCTION and Plasmodium species. Parasite load is not currently perceived as a significant Tympanu- The lesser prairie-chicken contributor to lesser prairie-chicken mor- chus pallidicinctus (Phasianidae: Tetraon- tality. However, if remaining lesser prairie- inae) is a lekking grouse species found in chicken populations become concentrated the dry short grass prairie of Colorado, into smaller areas of acceptable habitat, Kansas, New Mexico, Oklahoma, and Tex- parasite transmission rates could rise as (USA). In New Mexico, the shinnery (Dobson and May, 1986). oak (Quercus havardii) shrublands of the Staked Plain are the principal breeding MATERIALS AND METHODS grounds. A 4 yr trapping and banding Lesser prairie-chickens were trapped in the study of T. pallidicinctus was conducted in wild on the Bureau of Land Management ad- Chaves County, eastern New Mexico. In ministered Caprock Wildlife Habitat Manage- the last 3 yr of the study, blood and fecal ment Area, Chaves County, New Mexico samples were collected in the field. This (33Њ28Ј22ЉN, 103Њ47Ј44ЉW). Members of four population of prairie-chickens inhabits leks were sampled for blood and fecal parasites (Johnson and Smith, 1998, 1999). All birds rangeland managed for wildlife, grazing, screened for parasites were trapped within and oil and gas development. Recent de- 3,200 ha of contiguous habitat. This constitutes clines in T. pallidicinctus populations are a a small percentage of the extant New Mexico consequence of habitat alteration and population. We trapped birds in circular, weld- ed-wire, walk-in traps connected by chicken- drought (Giesen, 1998). Chemical control wire drift fences (Toepfer et al., 1987). Samples of shinnery oak has further reduced avail- were taken in late March, April, and early May, able nesting and brood-rearing habitat. from 1998–2000. Average capture time was The species is a candidate for federal pro- 6:30 AM (0500–08:15 AM, nϭ56), with only one tection as threatened by the United States captured after 8:00 AM. Each bird was temporarily detained in a small cardboard box. Fish and Wildlife Service. This study Fecal samples were collected from holding found the eastern New Mexico population boxes, or during the banding and release pro- of T. pallidicinctus positive for Eimeria cess, and transported to the University of New

347 JOURNAL OF WILDLIFE DISEASES Thursday May 22 2003 03:14 PM jwdi 39_212 Mp_348 Allen Press • DTPro System File # 12em

348 JOURNAL OF WILDLIFE DISEASES, VOL. 39, NO. 2, APRIL 2003

FIGURES 1, 2. Photomicrographs of sporulated oocysts of Eimeria tympanuchi.SBϭStieda body, ORϭoocyst residuum. Barϭ10 ␮m.

Mexico (UNM; Albuquerque, New Mexico). 20 min (Ն150 fields/film) for hematozoa, in- Morphologic data including weight, tarsus cluding Plasmodium, , Leucocy- length, wing chord, culmen depth, and length tozoon, trypanosomes, and microfilarie. Preva- of pinna were taken on each donor bird, as well lence of parasitemias was determined from four as age, sex, date, time, and location of capture. randomly selected counts of 500 erythrocytes The unique identity of each bird was kept via (nϭ2,000) per positive slide (Godfrey et al., a numbered aluminum leg band provided by 1987). Positive slides were read extensively in the New Mexico Department of Game and order to best describe life stage characteristics. Fish. All measurements of Eimeria and Plasmodium Fecal samples were placed in separate vials species are in micrometers with means fol- containing 2% aqueous (w/v) potassium dichro- lowed by ranges in parentheses. mate (K2Cr2O7). In the laboratory each sample was incubated at room temperature and pro- RESULTS cessed following the methods outlined in Du- szynski and Wilber (1997). All samples were ex- The birds we examined were infected amined and oocysts measured and photo- with one Eimeria species, described here graphed within 1 yr of collection. Photosyn- as new, and with one malarial species, types of sporulated oocysts are deposited in the United States National Parasite Collection Plasmodium (Giovannolaia) pedioecetii (USNPC), Beltsville, Maryland (USA) follow- first described by Shillinger (1942), ing Bandoni and Duszynski (1988) and Du- emended by Stabler et al. (1973). szynski (1999). We drew 10–50 ␮l of blood from captured Eimeria tympanuchi n. sp. (Figs. 1, 2, 5) birds using brachial venipuncture. Smears of Description whole blood were made in the field. Each slide was fixed in 100% methyl alcohol to preserve Sporulated oocysts (nϭ56) ellipsoidal, the cells, and later were stained with Giemsa’s blood stain. Smears were scanned under oil im- 27.1ϫ22.7 (22–32ϫ18–26); shape index mersion (1,000ϫ) to detect the presence of (SIϭlength/width)ϭ1.2 (1.0–1.5); wall blood parasites. Blood films were scanned for smooth in optical cross-section, 1.8 (1.0– JOURNAL OF WILDLIFE DISEASES Thursday May 22 2003 03:14 PM jwdi 39_212 Mp_349 Allen Press • DTPro System File # 12em

SMITH ET AL.ÐPARASITES IN LESSER PRAIRIE-CHICKEN 349

FIGURES 3, 4. Photomicrographs of Plasmodium (Giovannolaia) pedioecetii macrogametocyte (3, arrow) and ring stage (4). Barϭ10 ␮m.

2.5) thick, with two layers; outer, lightly Type locality: USA, New Mexico, striated, ϳ2/3 of total thickness; micropyle Chaves County, 33Њ28Ј22ЉN, 103Њ47Ј44ЉW. absent; usually one small, refractile polar Prevalence: 5/64 (8%). granule present; two to five oocyst residual Site of infection: Unknown, oocysts re- bodies present as round, opaque spheres. covered from feces. Sporocysts (nϭ56) ovoidal, 11.9ϫ7.8 (10– Material deposited: Photosyntypes of 14ϫ6–10) with L : W ratio 1.5 (1.2–2.0); sporulated oocysts in the USNPC, No. Stieda body present at pointed end of spo- 092386.00. rocyst, with evidence that a small, indis- Etymology: The specific name is de- tinct substieda body also may be present rived from the generic name of the host. immediately below it; sporocyst residuum Remarks: There are now 15 Eimeria present, comprised of small uniformly species described from grouse and ptar- shaped spheres in a cluster, approximately migan (Phasianidae:Tetraoninae) (Table two, partially obscuring sporozoites; spo- 1). The sporulated oocysts of E. tympan- rozoites with two refractile bodies, a larger uchi are distinguished from the others by one at the blunt end and a second, smaller the presence of a distinct oocyst residuum, body at pointed end with a nucleus visible lacking a micropyle, and the presence of between them. both Stieda and substieda bodies (see Ta- ble 1). Taxonomic summary Plasmodium (Giovannolaia) pedioecetii (Shillinger, Type host: Tympanuchus pallidicinc- 1942) Stabler et al., 1973 (Figs. 3, 4) tus (Ridgway, 1885), lesser prairie-chicken Plasmodium was detected in five of 34 (Phasianidae: Tetraoninae). (17%) samples. The samples come from 32 JOURNAL OF WILDLIFE DISEASES Thursday May 22 2003 03:14 PM jwdi 39_212 Mp_350 Allen Press • DTPro System File # 12em

350 JOURNAL OF WILDLIFE DISEASES, VOL. 39, NO. 2, APRIL 2003 mea- ϭ Author(s) 1970 1970 and Gousseff, 1936 1955 Allen, 1934; Todd and Lepp, Allen, 1934; Todd and Lepp, Levine, 1953 Levine, 1953 Galli-Valerio, 1929 Galli-Valerio, 1927; Yakimoff Yakimoff and Gousseff, 1936 Simon, 1939; Honess and Post, Honess and Post, 1955 1 M, PG, SB, SR PG, SB, SR PG, SB PG, SB M(?) PG, SR PG, SB(?) M(?), PG, SB, SR M(?) Qualitative wall structures m) Present ? ? ␮ 1.5 1.5 M,2.0 MC, PG, SB, SR PG, SB, Stabler SSB, et SR al., 1979 Stabler et al., 1979 NG NG NG NG ( 1.5–2.0 1.5–2.0 Thickness 1.9 1.0 2.7 1.1 2.4 1.8 1.6 2.3 1.6 NG NG NG NG NG NG NG NG NG L/W (range) (2.0–2.9) (1.6–2.0) width Sporocyst 5–8) 6–8) 7–8) 6–7) 8–10) 6.1 7.1 7.6 6.7 8.8 7 12 3.6 10.9 ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ NG NG NG NG NG NG NG NG 1.2 M, MC(?), SB, SR Haase, 1939 (range) 12 13 9.8 12.8 14.8 11.8 15.4 14.6 12.2 Length species known from the subfamily Tetraonimae as given in the original description (NG (12–17 (10–14 (11–13 (13–18 (13–16 Eimeria 1.05 1.5 1.5 1.6 1.9 1.3 1.5 1.5 1.2 1.9 1.2 2.1 NG NG NG NG NG NG NG NG L/W (range) (1.4–1.6) (1.4–1.6) (1.7–2.2) (1.0–1.2) Oocyst width 15–19) 18–23) 13–18) 12–15) 15–20) 20–26) 18–20) 18–20) 12–20) 17–24) 15–15) 17.1 20.6 17.1 13.5 17.7 22.6 18.8 18.8 15 15.3 21.3 15 ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ ϫ NG (range) 24 31 22.6 21.6 20.2 28.3 24.9 32.5 29.6 26.6 26.0 28.6 Length (28–37 (18–25 (17–25 (18–23 (23–29 (26–30 (27–29 (22–37 (21–29 (30–31 (22–31.5 spp. Eimeria 1. Mensural characters of sporulated oocysts of Host/ E. centrocerci E. pattersoni E. angusta E. bonasae E. lyruri E. nadsoni E. tetricis E. leucuri E. oreoecetes E. brinkmanni E. fanthami E. lagapodi ABLE Centrocercus urophasianus T Bonasa umbellus Tetrao tetrix surements, structures not given/mentioned) and/or in subsequent re-descriptions of sporulated oocysts. Lagopus leucurus Lagopus mutus JOURNAL OF WILDLIFE DISEASES Thursday May 22 2003 03:14 PM jwdi 39_212 Mp_351 Allen Press • DTPro System File # 12em

SMITH ET AL.ÐPARASITES IN LESSER PRAIRIE-CHICKEN 351 Stieda ϭ ys certain Author(s) Hardcastle, 1943 1 or more polar granules; SB ϭ Yakimoff and Gousseff, 1936; 1 oocyst residuum; PG ϭ M, PG, SB, SR Haase, 1939

FIGURE 5. Line drawing of a sporulated oocyst of

Qualitative wall structures Eimeria tympanuchi. Barϭ6 ␮m. 1.8 OR, PG, SR, SB, SSB This study 1.0 NG Thickness Present micropyle cap; OR

ϭ subjects, because two birds were captured and screened in successive years. One in- 2.1 1.5 NG NG L/W

(range) dividual was found positive in successive (1.2–2.0) years, the only infected bird either recap-

micropyle; MC tured or resighted between years. There- ϭ width Sporocyst fore, four of 32 possible hosts were infect- 6–10) 5.1 7.8 ϫ ϫ ϫ ϫ NG NG ed (13%). Intensity of each case (0.005– (range) 11.9 10.8 0.007, nϭ5) suggests hosts persisted with Length

(10–14 chronic infections. Trophozoites either were ameboid (78%) with one to five (x¯ϭ1.5) small pig- 1.2 1.4 NG NG L/W

(range) ment granules or were ring-shaped (22%), (1.0–1.5) with central vacuoles and one to four ag- gregated pigment granules (Fig. 4); 70% of all trophozoites seen were subpolar or Oocyst width 20–23) 18–26) 22 22.7

ϫ polar. ϫ ϫ ϫ ϫ 20 (range) 32 Gametocytes (nϭ29) appeared imma- (24–19.5) 27.1

Length ture and measured 7.5ϫ1.9 (6.25– (31–33 (22–32 10ϫ1.25–2.5); 25/29 (86%) were found on

sporocyst residuum. If not stated in the original description or the re-description, this structure isthe presumed to be absent although this is notlateral alwa edge of the nucleus without ap- ϭ parent alteration of nuclear position or cell wall. Pigment granules, 3.4 (1–11), were often on the edge of developing gameto- spp. cytes. Mature gametocytes (Fig. 3) were elongate and narrow, with irregular edges substieda body; SR Eimeria ϭ and with pigment granules throughout

1. Continued. their cytoplasm. The ends of the mature Host/ gametocyte usually curled around the po- E. tympanuchi E. ventriosa E. yakisevi ABLE Oocyst and sporocyst qualitative structures present in each species. Abbreviations: M in some of the older work. body; SSB Tympanuchus pallidicinctus T T. urogallus 1 lar ends of the nucleus without distortion JOURNAL OF WILDLIFE DISEASES Thursday May 22 2003 03:14 PM jwdi 39_212 Mp_352 Allen Press • DTPro System File # 12em

352 JOURNAL OF WILDLIFE DISEASES, VOL. 39, NO. 2, APRIL 2003

of erythrocyte shape. Only one mature In this study there was a low prevalence segmenter was seen with nine merozoites of schizonts and mature gamonts. In a visible in a tight cluster. It did not distort study of P. (Giovannolaia) pedioecetii from the erythrocyte’s shape or the position of galliformes, Stabler and Kitzmiller (1976) its nucleus. reported a daily peak in merozoite pro- The stage of the life cycle most evident duction from 8:00 AM to noon. The earlier was ameboid trophozoites in erythrocytes average capture time of birds sampled in (60%). The gamont life stages were often our study (0500–08:15 AM) may explain immature (66%), hence the observation of the lack of segmenters and large number mean length and width falls beneath the of underdeveloped gametocytes. stated means of Stabler (1978). Mature ga- metocyte dimensions and morphology are ACKNOWLEDGMENTS consistent with P. (Giovannolaia) pedioe- We thank the Bureau of Land Management, cetii. Length/width measurements of a Roswell, New Mexico Field Office for funding subset of nine mature gametocytes, the majority of this study. Grants from T&E, 9.5ϫ2.1, almost exactly reflects the means Inc. and the New Mexico Department of Game reported by Stabler (1978), 9.2ϫ2.0. Ga- and Fish Share With Wildlife program allowed us to complete the 2000 field season. We thank metocytes from the present study, howev- the following for trapping work: M. Berry, D. er, were more prone to lateral orientation Bilyeu, M. Kline, B. Long, J. Montgomery, M. (85%) than those reported by Stabler Radke, and C. Westwood and I. Asmundsson, (33.8%), in 1978. L. Couch, K. Decker, J. Hnida, A. Lynch, and M. Ryan for assistance in the laboratory. We DISCUSSION are especially grateful to L. Hertel for the line drawing. There are currently 15 Eimeria spp. de- scribed from the subfamily Tetraoninae, LITERATURE CITED including the new species described here. The systematics of grouse and ptarmigan ALLEN, E. A. 1934. Eimeria angusta sp. nov. and Eimeria bonasae sp. nov. from grouse, with a key were recently amended, and this paper fol- to the species of Eimeria in birds. Transactions lows the American Ornithologists’ Union of the American Microscopical Society 53: 1–5. (1998) classification of species within Te- AMERICAN ORNITHOLOGISTS’UNION. 1998. Check- traoninae. The Tetraoninae has six genera list of North American birds, 7th Edition. Amer- including Tympanuchus, which has three ican Ornithologists’ Union, Washington, D.C., 829 pp. species: T. cupido, T. pallidicinctus, and T. BANDONI, S., AND D. W. DUSZYNSKI. 1988. A plea for phasianellus. The lack of clear genetic dif- improved presentation of type material for coc- ferences within this species complex (Ells- cidia. Journal of Parasitology 74: 519–523. worth et al., 1996) and proximity of his- DOBSON,A.P.,AND R. M. MAY. 1986. Disease and toric range, suggested a comparison of ex- conservation. In Conservation biology: The sci- ence of scarcity and diversity. M. E. Soule (ed.). isting eimerians of closely related species. Sinauer Press, Sunderland, Massachusetts, pp. In a comparison of the characters of spor- 345–365. ulated oocysts from Tetraonid hosts, the DUSZYNSKI, D. W. 1999. Revisiting the Code: Clari- oocyst residuum of E. tympanuchi is nov- fying name-bearing types for photomicrographs. el. There is no report of an oocyst resid- Journal of Parasitology 85: 769–770. , AND P. G. WILBER. 1997. A guideline for uum present in any of the Eimeria species the preparation of species descriptions in the Ei- found in hosts of the subfamily Tetraoni- meriidae. Journal of Parasitology 83: 333–336. nae. ELLSWORTH, D. L., R. L. HONEYCUTT, AND N. J. SIL- Light infections of the hematozoan Plas- VY. 1996. Systematics of grouse and ptarmigan modium (Giovannolaia) pedioecetii were determined by nucleotide sequences of the mi- tochondrial cytochrome-b gene. The Auk 113: found. This species was previously de- 811–822. scribed from lesser prairie-chickens of GALLI-VALERIO, B. 1927. Notes de parasitologie et New Mexico and Texas by Stabler (1978). de technique parasitologique. Zentralblatt fu¨r JOURNAL OF WILDLIFE DISEASES Thursday May 22 2003 03:14 PM jwdi 39_212 Mp_353 Allen Press • DTPro System File # 12em

SMITH ET AL.ÐPARASITES IN LESSER PRAIRIE-CHICKEN 353

Bakteriologie. I. Abtellung Originale 103: 177– Charadriiformes, with descriptions of three new 182. species. American Midland Naturalist 49: 696– . 1929. Notes de parasitologie. Zentralblatt 719. fu¨ r Bakteriologie. I. Abtellung Originale 112: 54– SCHILLINGER, J. E. 1942. Diseases of wildlife and 59. their relationship to domestic livestock. Yearbook GIESEN, K. M. 1998. Lesser prairie-chicken (Tym- of agriculture, USDA, US Government Printing panuchus pallidicinctus) In The birds of North Office, pp. 1217–1225. America, Number 364. A. Poole and F. Gill SIMON, F. 1939. Eimeria centrocerci n. sp. du Cen- (eds.). The Birds of North America, Inc., Phila- trocercus urophasianus (coq du Bruye`re). An- delphia, Pennsylvania, 17 pp. nales de Parasitologie Humaine et Compare´e, GODFREY,JR., R. D., A. M. FEDYNICH, AND D. B. Paris 17: 137–138. PENCE. 1987. Quantification of hematozoa in STABLER, R. M. 1978. Plasmodium (Giovannolaia) blood smears. Journal of Wildlife Diseases 23: pedioecetii from the lesser prairie-chicken, Tym- 558–565. panuchus pallidicinctus. Journal of Parasitology HAASE, A. 1939. Untersuchungen u¨ ber die bei 64: 1125–1126. deutschen Wildhu¨ hnern vorkommenden Eime- , AND N. J. KITZMILLER. 1976. Plasmodium ria-Arten. Archiv fu¨ r Protistenkunde 92: 329– (Giovannolaia) pedioecettii from gallinaceous 383. birds of Colorado. Journal of Parasitology 62: HARDCASTLE, A. B. 1943. A check-list and host index 539–544. of the species of the protozoan genus Eimeria. , , AND C. E. BRAUN. 1973. Plasmo- Proceedings of the Helminthological Society of dium in a Darwin’s tinamou from Colorado. Jour- Washington 10: 35–69. nal of Parasitology 59: 395. ,A.G.HASKINS,N.J.KITZMILLER,O.W.OL- HONESS,R.F.,AND G. POST. 1955. Eimeria of grouse (Family: Tetraonidae), with a description of Ei- SEN, AND C. E. BRAUN. 1979. Two new species meria pattersoni n. sp. from the sage grouse. Wy- of coccidia, Eimeria leucuri and E. oreocetes oming Game & Fish Commission, Bulletin 8, Ar- (Protozoa: ), in grouse from Colorado. Journal of Parasitology 65: 272–274. ticle 2: 5–11. TODD,JR., K. S., AND D. L. LEPP. 1970. Redescrip- JOHNSON, K., AND B. H. SMITH. 1998. Radio telem- tion of Eimeria angusta Allen, 1934 and E. bon- etry study of lesser prairie-chicken habitat use in asae Allen, 1934 (Protozoa: Eimeriidae) from the the Caprock Wildlife Habitat Management Area. ruffed grouse Bonasa umbellus. Journal of Pro- New Mexico Natural Heritage Program technical tozoology 17: 427–429. report to the Bureau of Land Management, TOEPFER, J. E., J. A. NEWELL, AND J. MONARCH. 17 pp. 1987. A method for trapping prairie grouse hens , AND . 1999. Lesser prairie-chicken on display grounds. Montana Agriculture Exper- habitat use on the Sand Ranch and population iment Station, Contribution #2144: 21–23. status in the Caprock Wildlife Habitat Manage- YAKIMOFF, W. L., AND W. F. G OUSSEFF. 1936. A pro- ment Area, 1999. New Mexico Natural Heritage pos des coccidies des oiseaux sauvages. Annales Program technical report to the Bureau of Land de Parasitologie Humaine et Compare´e 14: 449– Management, 19 pp. and maps. 456. LEVINE, N. D. 1953. A review of the coccidia from the avian orders Galliformes, Anseriformes and Received for publication 13 May 2002.