Syst Parasitol (2008) 71:223–228 DOI 10.1007/s11230-008-9150-x Two new species of Eimeria Schneider, 1875 (Apicomplexa: Eimeriidae) from Gerbilliscus guineae Thomas (Rodentia: Gerbillinae) in the Niokolo Koba National Park, Senegal David Modrý Æ Klára Petrželková Æ Miloslav Jirků Æ Petr Koubek Received: 8 January 2008 / Accepted: 22 May 2008 Ó Springer Science+Business Media B.V. 2008 Abstract We describe two new species of Eimeria Schneider, 1875 from the gerbiline rodent Gerbilliscus guineae in the Niokolo Koba National Park, Senegal. Faecal examination of samples revealed the presence of sporulated oöcysts of two eimerian coccidia, both possessing an oöcyst residuum. Eimeria permira n. sp. is remarkable in terms of oöcyst size and oöcyst wall texture. Sporulated oöcysts are ellipsoidal, 45.8 (42–50) 9 32.5 (31–38) lm; the oöcyst wall is 3–4 lm thick, composed of three layers, with the outer layer sheathed by rough granular material; and the sporocysts are broadly ellipsoidal, 15.4 (15–16) 9 11 and with a Stieda body present. Oöcysts of Eimeria gerbillisci n. sp. are D. Modrý (&) Department of Parasitology, University of Veterinary and Pharmaceutical Sciences, Palackého 1–3, 612 42 Brno, Czech Republic e-mail: modryd@vfu.cz D. Modrý
M. Jirků Institute of Parasitology, Biology Centre, Academy of Sciences of the Czech Republic, Branišovská 31, 370 05 České Budějovice, Czech Republic K. Petrželková
P. Koubek Institute of Vertebrate Biology v.v.i., Academy of Sciences of the Czech Republic, Květná 8, 603 00 Brno, Czech Republic K. Petrželková Liberec Zoo, Masarykova 1347/31, 460 01 Liberec, Czech Republic subspherical, 22.5 (19.5–24) 9 18.8 (16.5–20) lm, with a colourless, faintly granulated oöcyst wall 1.5 thick; and the sporocysts are 10.1 (10–12) 9 6.7 (6– 8), broadly ellipsoidal and often somewhat pointed towards both ends. Introduction Compared to the diversity of African rodents, coccidian parasites of these hosts represent a neglected group of symbiotic organisms. Among the coccidia, Eimeria Schneider, 1875 undoubtedly represents the most diversified. Although, there are more than 30 species of Eimeria described from African rodents, none of them are reported from the gerbils of the genus Gerbilliscus Thomas. In this report, we describe two new species of Eimeria from Gerbilliscus guineae Thomas, a species distributed in the Sub-Saharan part of West Africa (Gambia, Senegal to Ghana and Burkina Faso). Materials and methods Within the study undertaken in Niokolo Koba National Park, Senegal, during February and December, 2005, two specimens of Gerbilliscus guineae were trapped at Lingué Kountou (13°020 N, 13°050 W; 22.ii.2005) and Dar Salam (13°150 N, 13°120 W; 15.xii.2005) by snap-traps. Fresh contents of the 123 224 caecum and colon were placed in plastic vials with 2.5% (w/v) potassium dichromate (K2Cr2O7), mixed thoroughly, transported to the laboratory and examined. The faecal samples were examined microscopically after concentration by flotation with modified Sheather’s sugar solution (s.g. 1.30). Oöcysts were measured and photographed using DIC optics on an Olympus AX 70 microscope. Measurements were made using a calibrated ocular micrometer and are reported in micrometres, as the mean followed by the range in parentheses. Results Faecal examination of samples revealed the presence of sporulated oöcysts of two eimerian coccidia in one of the samples. These are described below as two new species. Eimeria permira n. sp. Type-host: Gerbilliscus guineae Thomas (Rodentia: Muridae: Gerbillinae). Type-locality: Lingué Kountou (13°020 N, 13°050 W), Niokolo Koba National Park, Senegal. Type-material: Photosyntypes are deposited in the protozoological collection of the Institute of Parasitology of the Czech Academy of Sciences, České Budějovice, Czech Republic under collection number IP CAS Prot. Coll.: P-4. Symbiotype: An alcohol-preserved voucher specimen of the host is deposited in the Institute of Vertebrate Biology v.v.i., AS CR, under the collection number SE 489. Site: Unknown; oöcysts were collected from the intestinal contents. Sporulation: Apparently exogenous; some oöcysts observed in the faeces were unsporulated. Etymology: The specific epithet permira (= weird) reflects the unusual oöcyst morphology of the species described. Description (Figs. 1–4, 9) Unsporulated oöcysts contain single granular sporont. Sporulated oöcysts ellipsoidal, 45.8 (42–50) 9 32.5 (31–38); shape index (SI; length/width ratio) 1.41 123 Syst Parasitol (2008) 71:223–228 (1.11–1.56). Oöcyst wall 3–4 thick, composed of 3 layers. Inner colourless layer 0.5; outer yellowbrownish layer 2–2.5, thickened at oöcyst poles, sheathed by rough granular material c.1 thick with numerous external projections (extending to 4 above outer layer surface). Micropyle absent. Polar granule present, usually single (sometimes 2), subspherical, up to 3 in diameter. Apart from polar granule, 1–3 fine granules are present in some oöcysts. Distinct oöcyst residuum present, variable both in size and morphology, consisting either of several large globules accompanied by fine granular material, or, less frequently, of small granules. In many oöcysts, membranous structure surrounding oöcyst residuum present. Sporocysts dizoic, broadly ellipsoidal, single-layered, colourless, 15.4 (15–16) 9 11, SI = 1.4 (1.36–1.45). Stieda body (SB) present, lentil-like; substieda body large (although sometimes not discernible from SB using light microscopy), subglobular, c.2 in diameter. Sporocyst residuum present, composed of granules (0.5–2 wide) usually scattered among sporozoites. Finely granulated sporozoites possess 1 distinct elongate (c.7–9 9 4–5) refractile body each. Eimeria gerbillisci n. sp. Type-host: Gerbilliscus guineae Thomas (Rodentia: Muridae: Gerbillinae). Type-locality: Lingué Kountou (13°020 N, 13°050 W), Niokolo Koba National Park, Senegal. Type-material: Photosyntypes are deposited in the protozoological collection of the Institute of Parasitology of the Czech Academy of Sciences, České Budějovice, Czech Republic under collection number IP CAS Prot. Coll.: P-5. Symbiotype: An alcohol-preserved voucher specimen of the host is deposited in the Institute of Vertebrate Biology v.v.i., AS CR, under the collection number SE 489. Site: Unknown; oöcysts collected from the contents of the caecum and colon. Sporulation: Probably exogenous; oöcysts observed in the faeces were sporulated at the time of examination. Etymology: The specific epithet gerbillisci is the genitive of the generic name of the type-host. Syst Parasitol (2008) 71:223–228 225 Figs. 1–8 DIC photomicrographs of the two species of Eimeria described from Gerbilliscus guineae. 1–2. Sporulated oöcyst of E. permira n. sp., note the prominent Stieda body (arrow) and polar granule (arrowhead). 3–4. Sporulated oöcyst of E. permira with evident oöcyst wall micro-ornamentation, note the distinct oöcyst residuum (arrowhead). 5–8. Sporulated oöcyst of E. gerbillisci n. sp., note the oöcyst residuum (6, arrowhead), polar granule (7, arrowhead) and very fine granulation of the oöcyst wall (8). Scale-bar: 20 lm Description (Figs. 5–8, 10) Sporulated oöcysts subspherical, 22.5 (19.5–24) 9 18.8 (16.5–20), SI = 1.20 (1.05–1.35). Oöcyst wall 1.5 thick, bi-layered, colourless, faintly granulated externally. Micropyle absent. One to 3 subspherical polar granule(s) usually present, up to 2 in diameter. Oöcyst residuum distinct, consists of large globules accompanied by fine granular material. Sporocysts dizoic, 10.1 (10–12) 9 6.7 (6–8), SI = 1.51 (1.29– 1.71), broadly ellipsoidal, often somewhat pointed towards both ends, with single-layered colourless wall. Sporocyst residuum present, consists of several granules (0.5–2 in diameter) scattered among sporozoites. Stieda body present (2–2.5 9 1–1.5). Sporozoites vermiform, with single relatively large (4–5 long) refractile body each. Discussion Eimerian coccidia represent by far the most diversified group of apicomplexans (e.g. Upton, 2000). Also in rodents, they represent the most common, and, thanks to the easy detection of oöcysts, also most frequently described protozoans. However, studies on coccidian parasites of African rodents are still in their infancy. Slightly more than 30 123 226 Syst Parasitol (2008) 71:223–228 Figs. 9–10 Composite line drawings of the oöcysts of the two described species of Eimeria. 9. E. permira n. sp. 10. E. gerbillisci n. sp. Scale-bar: 20 lm species have been described from these hosts so far, sharply contrasting with the diversity of rodents inhabiting the African continent (Levine & Ivens, 1965; Koudela et al., 2000; Šlapeta et al., 2001a,b; Modrý et al., 2005). No species have yet been reported from rodents of the Gerbillinae in SubSaharan Africa. Oöcysts of Eimeria permira n. sp. are so distinct that they are easily distinguishable from other rodent Eimeria spp. The oöcyst dimensions, rough surface and characteristic oöcyst residuum represent key diagnostic features. The presence of an oöcyst residuum was found to correlate with eimerian phylogeny (Zhao & Duszynski, 2001; Kvičerová et al., 2008) and it is essential for distinguishing E. gerbillisci n. sp. from most of Eimeria spp. parasitising African rodents. Based on the presence of this morphological feature, only seven species can be compared with E. gerbillisci (Table 1). Of these, E. livingstonei Šlapeta, Modrý & Koudela, 2001 and E. shoutedeni Van Den Berge & Chardome, 1957 are much smaller, E. arvicanthis van den Berge & Chardome, 1957 possesses much more elongate oöcysts, E. tachyoryctis van den Berge & Chardome, 1957 differs in the absence of a polar granule, and E. heliophobii Modrý, Jirků & Šumbera, 123 2005 can be readily distinguished by its heavily pitted oöcyst wall (Šlapeta et al., 2001a; Modrý et al., 2005). In relation to oöcyst morphology, the most similar species are E. pretoriensis de Vos, 1970 from Rhabdomys pumilio Sparrman in South Africa and E. yamikamiae Modrý, Jirků & Šumbera, 2005 from Heliophobius argenteocinereus Peters in Malawi. The oöcysts of E. pretoriensis differ in having a thicker, more ornamented and brownish wall (de Vos & Dobson, 1970), and E. yamikamiae can be differentiated by the presence of numerous small granules within the oöcyst (Modrý et al., 2005). In contrast to those from African species, coccidia from gerbiline rodents in the Near and Middle East are much better understood, mainly thanks to researchers from countries of the former USSR (e.g. Levine & Ivens, 1965; Musaev & Veisov, 1965; Pellérdy, 1974). However, the hosts are only distantly related and the conspecificity of these species with coccidia from Sub-Saharan rodents is unlikely, due to the presence of several biogeographical barriers and the geographical distance itself. We believe, that observed differences substantiate the description of both Eimeria species as new to science. Species of Eimeria Type-host E. permira Tatera guineae E. taterae Site Oöcyst Sporocyst Distribution Reference PG/ OR Oöcyst wall Size (lm) Shape SI Size (lm) Sb/ssb nd 1.41 (1.11–1.56) +/+ 3–4 lm, 3 layers 45.8 (42–50) 9 32.5 (31–38) 1.4 (1.36– 1.45) 15.4 (15–16) 9 11 +/+ Senegal This study Tatera guineae nd 1.20 (1.05–1.35) +/+ 1.5 lm, bilayered, colourless 22.5 (19.5–24) 9 18.8 (16.5–20) 1.51 (1.29– 1.71) 10.1 (10–12) 9 6.7 (6–8) +/? Senegal This study E. arvicantis Arvicanthis abyssinicus rubescens intestine ovoidal -/± thin, rosecoloured 23–24 9 10–14 ellipsoidal 10–1 9 7 - Belgian Congo van den Berghe & Chardome (1956a) E. heliophobii Heliophobius argenteocinereus unknown broadly ellipsoidal, 1.25 (1.18–1.33) +/+ 2.5–3 lm, bilayered, brownish, heavily pitted 27.9 9 22.3 (22–31 9 18–24.5) oval 1.55 (1.5–1.62) 12.8 9 8.4 (12–14 9 8–9) + Malawi Modrý et al. (2005) E. livingstonei Lophuromys flavopunctatus nd spindleshaped to ellipsoidal, 1.4 (1.5–1.3) -/+ 1.3 lm, bi-layered, 19.6 9 13.6 (17–22 9 moderately 12.5–15) rough, colourless ellipsoidal 12 9 5.5 (10–15 9 5–6) +/+ Ruwenzori Mts, Šlapeta et al. Uganda (2001a) E. pretoriensis Rhabdomys pumilio small broadly intestine ellipsoidal, 1.23 (1.1–1.5) +/+ 1.5 lm, bi-layered, 24 9 19 (18–27 9 thick, outer layer 16–22) thicker, light brown with roughened surface broadly ellipsoidal 10 9 7 (8–12 9 5–8) +/+ South Africa de Vos (1970) E. shoutedeni Cricetomys dissimilis ovoidal -/± 0.5 lm, singlelayered, smooth 14–15 9 11–13 7 9 5.6 nd Kivu, Kongo van den Berghe & Chardome (1957) E. tachyoryctis Tachyoryctes ruandae intestine ovoidal -/+ single-layered, quite thick, colourless 23 9 17 ovoidal 12 9 8 nd Belgian Congo van den Berghe & Chardome (1956b) E. yamikamiae Heliophobius argenteocinereus unknown +/+ broadly ellipsoidal to sunspherical, 1.19 (1.13–1.23) 1.5 lm, bi-layered, faintly pitted 20.8 9 17.5 (19–22 9 15.5–19) oval 1.56 (1.53– 1.57) 10.7 9 6.8 (10–11 9 6–7) + Malawi Modrý et al. (2005) Abbreviations: Sb, Stieda body; ssb, substieda body; PG, polar granule, OR, oöcyst residuum, SI, shape index; nd, no data 227 123 Shape SI Syst Parasitol (2008) 71:223–228 Table 1 Basic features of the oöcyst residuum-possessing Eimeria spp. reported from rodents in Sub-Saharan Africa 228 Acknowledgements The study was funded by the Grant Agency of the Czech Republic (grants 524/03/H133 and 524/ 03/1548) and the Grant Agency of the Czech Academy of Sciences (grant no. IAA 6093403). References de Vos, A. J., & Dobson, L. D. (1970). Eimeria chinchillae de Vos & Van der Westhuizen, 1968 and other Eimeria spp. from three South African rodent species. Onderstepoort Journal of Veterinary Research, 37, 185–190. Koudela, B., Šumbera, R., & Sedláček, F. (2000). Eimeria burdai sp. n. (Apicomplexa: Eimeriidae) a new parasite species from subterranean African silvery mole-rat, Heliophobius argenteocinereus. Folia Parasitologica, 47, 97–99. Kvičerová, J., Pakandl, M., & Hypša, V. (2008). 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