Loaina Gen. N. (Filarioidea: Onchocercidae) for the Filariae Parasitic in Rabbits in North America
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Proc. Helminthol. Soc. Wash. 51(1), 1984, pp. 49-53 Loaina gen. n. (Filarioidea: Onchocercidae) for the Filariae Parasitic in Rabbits in North America MARK L. EBERHARD AND THOMAS C. ORIHEL Department of Parasitology, Delta Regional Primate Research Center, Covington, Louisiana 70433 ABSTRACT: A new genus, Loaina, is erected to accommodate the species of Dirofilaria parasitic in rabbits in North America. The genus is comprised of two species, Loaina scapiceps (Leidy, 1886) comb. n. and Loaina uniformis (Price, 1957) comb, n.; L. uniformis is designated as the type species. The genus Loaina is distinguished morphologically from Dirofilaria and other Dirofilariinae by a combination of characters, including an extremely short tail in both sexes, an undivided esophagus, a post-esophageal vulva, short, simple spicules, a small number of caudal papillae grouped at the posterior extremity of the body in the male, and a sheathed microfilaria. Two other species, D. timidi and D. roemeri are not regarded to be valid species of Dirofilaria. The genus Dirofilaria should be restricted strictly to those species having unsheathed microfilariae. Morphologically, the genus Loaina most closely resembles the genus Loa. The two genera are distinct, however, on the basis of host preference, size and cuticular ornamentation. A review of the morphological features of the ments of anatomical features are from stained speci- dirofilarias that parasitize Louisiana mammals mens. revealed that there were marked morphological Description differences between those species of Dirofilaria Railliet and Henry, 1910 infecting rabbits in Loaina gen. n. North America, i.e., D. uniformis Price, 1957 Dirofilaria Railliet and Henry, and D. scapiceps (Leidy, 1886), and other mem- 1910, in part bers of the genus. These differences were ob- General served in both the adult worms and in the Onchocercidae (Leiper, 1911); Dirofilariinae microfilariae. The distinctiveness of these mor- Sandground, 1921. Anterior and posterior ends phological features warrants that a new genus, bluntly rounded in both sexes. Male one-half to Loaina, which will accommodate the species D. one-third length of female. Cuticle without boss- scapiceps and D. uniformis, be erected. es, lateral alae sometimes present. Tail of both The report that follows presents a description sexes extremely short, less than twice width of of the new genus, a discussion of its taxonomic body at anus. Male tail not spirally coiled. Esoph- relationships to other genera in the subfamily agus not divided, without long, sacculate, glan- Dirofilariinae Sandground, 1921, as well as re- dular part. Vulva postesophageal. Spicules short, marks on the above-mentioned species. simple. Caudal papillae large, paired, few in Materials and Methods number, grouped at end of body. Microfilaria Nematodes in the authors' collections identified as sheathed, nuclei to tip of tail; circulates in blood. D. uniformis were recovered at necropsy by skinning TYPE SPECIES: Loaina uniformis (Price, 1957) freshly killed rabbits and searching the subcutaneous comb. n. and muscle tissues for worms, or, more commonly, OTHER SPECIES: Loaina scapiceps (Leidy, 1886) worms were recovered after soaking the skin and car- comb. n. cass in physiological saline. Worms identified as D. scapiceps were recovered by blunt dissections of the tarsal bursae of the hind feet. Worms recovered in Diagnosis either manner were examined alive, then fixed in gla- The genus Loaina is separated on morpholog- cial acetic acid or hot (60°C) Bouin's solution, and stored in 70% alcohol to which 5% glycerine (by vol- ical bases from other genera in the subfamily ume) had been added. Worms were studied micro- Dirofilariinae as outlined by Anderson and Bain scopically after being cleared in glycerine. Specimens (1976). The new genus is distinguished from Bos- examined from other collections are listed under each trichodera Sandground, 1938 and Dirofilariae- species. Microfilariae, obtained in cardiac blood, were fixed formia Lubimov, 1935 on the basis of the short in 2% formalin. They were examined as wet prepara- male tail. Loaina lacks the long, sacculate, glan- tions and as hematoxylin-stained smears. Measure- dular esophagus, characteristic of Edesonfilaria 49 Copyright © 2011, The Helminthological Society of Washington 50 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY Figures 1-7. Loaina uniformis comb. n. and Loaina scapiceps comb. n. 1. L. uniformis male tail, lateral view. 2. L. uniformis male tail, ventral view. 3. L. uniformis, female tail, lateral view. 4. L. uniformis female, lateral view. 5. L. uniformis microfilaria from 2% formalin concentration stained with hematoxylin. 6. L. scapiceps male tail, ventral view. 7. L. scapiceps male tail, lateral view. Yeh, 1960 and Macacanema Schad and Ander- tinguish Loaina from Skrjabinodera Gnedina and son, 1963. The short, simple spicules and small Ysevolodov, 1947, Tawila Khalil, 1932, and Di- number of large, pedunculated caudal papillae rofilaria Railliet and Henry, 1910. Also, the grouped near the end of the body serve to dis- sheathed microfilaria with a blunt tail in which Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 51, NUMBER 1, JANUARY 1984 51 the nuclei extend to the tip contrasts markedly with those of the genus Dirofilaria in which the microfilariae characteristically are unsheathed and possess thin, attenuated tails devoid of nu- clei. The postesophageal position of the vulva separates Loaina from Foleyella Seraut, 1917 and Pelicitus Railliet and Henry, 1910. Loaina can be distinguished from Loa Stiles, 1905 by its smaller size, the absence of cuticular bosses, and spicule morphology. The genus Loaina, on the basis of present knowledge, appears to be restricted to the order Lagomorpha in North America. Loaina uniformis (Price, 1957) comb. n. Dirofilaria uniformis Price, 1957 (Figs. 1-5, 8, 10) HOSTS: Sylvilagus floridanus, S. palustris, S. aquaticus. SPECIMENS EXAMINED: USNM Helm. Coll. No. 38160, 73652, numerous specimens in authors' collections from Cameron, Orleans, and St. Tammany Parishes, Louisiana, and Dade Coun- ty, Florida. Diagnosis Price (1957) provided an accurate and com- plete description of this species. However, he Figure 8. Formalin fixed, hematoxylin-stained mi- failed to recognize, at the generic level, significant crofilaria of Loaina uniformis comb, n., note sheath (arrows) ( x 425). morphological differences between the genus Di- rofilaria and the specimens he described. Most there is a more complete and accurate descrip- notable among these are the short tail in both tion, including illustrations of L. uniformis. sexes (Figs. 1-4), the small number of large cau- dal papillae grouped at the end of the body and Loaina scapiceps (Leidy, 1886) comb. n. the short, simple spicules in the male (Figs. 1, Dirofilaria scapiceps (Leidy, 1886) 2), and the sheathed microfilaria. The illustration (Figs. 6-7) of the microfilaria furnished in the original species description is one of an unstained, wet-mount HOSTS: Sylvilagus floridanus, S. palustris, S. preparation. Consequently, the internal mor- aquaticus, Lepus americanus, L. campestris, L. phology and nuclear arrangement were not clear- washingtonii. ly evident. Because of the taxonomic signifi- SPECIMENS EXAMINED: USNM Helm. Coll. Nos. cance, the microfilaria of L. uniformis has been 28647, 32016, 32621, 38066, 45326, 73651, and illustrated by both a line drawing and a photomi- specimens in authors' collections from St. Tam- crograph (Figs. 5, 8). many Parish, Lousiana. Loaina uniformis has been reported from sub- cutaneous tissues of rabbits in Maryland, South Diagnosis Carolina, Georgia, Florida, and Louisiana. The Leidy (1886) described in very cursory fashion distribution may be more extensive than indi- Filaria scapiceps, noting the features of size, gen- cated by these reports. eral body shape, and nature of the caudal papillae Loaina uniformis is designated as the type in the male. In 1916, Hall provided a more de- species for several reasons, the most important tailed and accurate description of the adult par- being the availability of type specimens. Also, asite, including measurements. However, the il- Copyright © 2011, The Helminthological Society of Washington 52 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY «8&m^ *•> > : Figures 9-10. Cross sections of female Loa loa and Loaina uniformis comb. n. at comparable levels. 9. L. loa, note various levels of uterus in same section and cuticular bosses (xllO). 10. L. uniformis, note ovary and various levels of uterus in same section of worm (xllO). lustration of the male tail is misleading as it bursa of the hind feet, whereas adult L. uniformis depicts the caudal papillae as being small and are found in subcutaneous tissues. slender rather than large and bulbous (Figs. 6- Loaina scapiceps apparently has widespread 7). Neither Leidy (1886) nor Hall (1916) provid- geographical distribution, which includes: New ed a description of the microfilaria. Only Highby Brunswick, Ontario, and British Columbia, Can- (1943), working on the life history of the parasite, ada, and the northwestern, northeastern, eastern has provided a substantial description of the mi- seaboard, and gulf coast regions of the United crofilaria. He noted the size, internal anatomy, States. and the presence of a sheath. We have been able, Specimens deposited as USNM Helm. Coll. in the present study, to examine microfilariae of No. 32016 (three female worms), and identified L. scapiceps. We confirmed