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Parasites of the Extinct Shasta Ground Sloth, Nothrotheriops Shastensis, in Rampart Cave, Arizona

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Parasites of the Extinct Shasta Ground Sloth, Nothrotheriops Shastensis, in Rampart Cave, Arizona 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 10-1992 Parasites of the Extinct Shasta Ground Sloth, Nothrotheriops shastensis, in Rampart Cave, Arizona Gerald D. Schmidt university of Northern Colorado Donald W. Duszynski University of New Mexico, [email protected] Paul S. Martin University of Arizona Follow this and additional works at: https://digitalcommons.unl.edu/parasitologyfacpubs Part of the Parasitology Commons Schmidt, Gerald D.; Duszynski, Donald W.; and Martin, Paul S., "Parasites of the Extinct Shasta Ground Sloth, Nothrotheriops shastensis, in Rampart Cave, Arizona" (1992). Faculty Publications from the Harold W. Manter Laboratory of Parasitology. 181. https://digitalcommons.unl.edu/parasitologyfacpubs/181 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. J. Parasitol., 78(5), 1992, p. 811-816 ? American Society of Parasitologists 1992 PARASITESOF THE EXTINCTSHASTA GROUNDSLOTH, NOTHROTHERIOPSSHASTENSIS, IN RAMPARTCAVE, ARIZONA Gerald D. Schmidt*, Donald W. Duszynski, and Paul S. Martint Departmentof Biology,The Universityof New Mexico,Albuquerque, New Mexico87131 ABSTRACT:Of 7 dissecteddung balls of the extinct Shastaground sloth (Edentata)from RampartCave, Arizona, 4 (57%)were found to contain nematodejuveniles, helminth eggs, and/or coccidian oocysts. One dung ball was radiocarbondated at 10,500 + 180 yr, about the time of groundsloth extinction.It is supposedthat the parasites also are extinct. Agamofilariaoxyura n. sp. is proposed for first-stagejuveniles of an oxyurid. These juveniles measured 13-20 x 126-198 (16.8 x 159) Aim.Strongyloides shastensis n. sp. is reportedas first-stagejuveniles, some of which clearlyare molting.These juvenile worms were 23-27 x 270-345 (24.4 x 305.3) ,em.Operculated schistosomelikeeggs, each with an abopercularpoint, were 33-50 x 63-90 (43.0 x 81.9) ,um.A second type of object that resembled a helminth egg had a thick wall and was 11-14 x 13-18 (13.5 x 15.5) ,m. A new collective genus,Archeococcidia, is proposedto include 2 new fossil oocysts. Archeococcidiaantiquus n. sp. was the most abundantform found. Its unsporulatedoocysts were 19-21 x 21-23 (19.6 x 22.1) Am, and its outer wall bore punctations.Unsporulated oocysts of Archeococcidianothrotheriopsae n. sp. were spheroidal,31-35 (33.8) um wide, and had a smooth outer surface. Parasites of prehistoric extinct mammals rare- at the top of a talus slope, more than 200 m above the ly are reported. Home (1985) and Wilke and Hall level of the ColoradoRiver. Passageways extend roughly 100 m into limestone Cambrian About (1975) provided annotated bibliographies on the (upper Period). 220 m2 of dung, reachinga maximum depth of 1.5 m, analyses of ancient feces, mainly of human ori- covers the floor of part of the cave. Two-thirdsof this gin, and Fry and Moore (1969) found Enterobius was destroyedby a smolderingfire betweenJuly 1976 sp. in dry human coprolites up to 10,000 yr in and March 1977. Exact provenienceof the dung balls in this is were radiocarbon age. The fossil dung of some mam- study unknown; they obtained from of in back dirt left the 1940 excavation rats is found in heaps dung by mals, especially pack (Neotoma), of Remington Kellogg(Martin, unpubl.). abundance in certain dry caves of the North Portionsof each dung ball were brokenup and rew- American desert southwest, where ancient feces etted for 48 hr in a 0.5% solution of trisodium phos- may endure for tens of thousands of years (Be- phate, then filtered through coarse cheesecloth, and later 40- and 60-mesh brass screens. tancourt et al., 1990). Occasionally, dung of cer- through Aliquots of the resulting filtrate then were examined either di- tain extinct mammals is also includ- preserved, rectly as wet mounts or they were concentrated by ing entire and fragmentary boluses of the Shasta flotation with a concentrated sucrose solution (specific ground sloth, Nothrotheriops shastensis. One of gravity 1.15). Parasitesand parasitestages were mea- sured with an ocular micrometer and several giant (extinct) edentates of the Western photographed with Panatomic-X 35-mm film within a Zeiss Uni- the late Pleistocene Hemisphere during Epoch, versalPhotomicroscope equipped with both brightfield its last remains can be traced to about 11,000 yr (Neofluar) and Nomarski-interference100 x objec- ago (Martin et al., 1985) when Shasta ground tives. All measurementsare in micrometers(,im) with sloths disappeared along with mammoths, sa- the means given in parenthesesfollowing the ranges. Phototypes(Bandoni and Duszynski, 1988) of the coc- bertooths, and other megafauna (Martin, 1987). cidian oocysts and both phototypes and syntypes of Examination of these dried boluses can offer in- the worms have been deposited in the U.S. National sight about the parasites that these mammals Museum Parasite Collection (USNMPC), Beltsville, maintained. Maryland. MATERIALSAND METHODS RESULTS We examined7 sloth dungballs collected from Ram- part Cave, Arizona, at the western end of the Grand The dung balls are large, up to 12 cm and Canyon. The cave is about 7 km from Pierce's Ferry occasionally more than 20 cm in diameter, lo- bate, with a reddish-brown varnish of dried mu- Received 30 July 1991; revised 17 October 1991; cus. Evidently the animal was totally herbivo- accepted 17 October 1991. to Hansen the identifiable * rous; according (1978), Department of Biological Sciences, University of plant tissues in the dung consist principally of Northern Colorado, Colorado 80631 (de- Greeley, Sphaeralcea; Mormon tea, Ephed- ceased). globemallow, t Departmentof Geosciences, University of Arizona, ra; and cactus, Opuntia. These xeric plants re- Tucson, Arizona 85721. main important in the regional vegetation of the 811 812 THE JOURNALOF PARASITOLOGY,VOL. 78, NO. 5, OCTOBER1992 area especially at higher elevation. Vegetation in ascarid egg (Fig. 11), but the outer layer is smooth the immediate vicinity of the cave during the and its size is much smaller (n = 8), 10.8-14.4 sloth's occupancy included juniper (Juniperus), x 12.6-18.0(13.5 x 15.5). ash (Fraxinus), and other woodland species that n. only occur now at higher elevations (Phillips, Agamofilaria oxyura sp. (Figs.1-3) 1984). Indeed, there is no indication that the Description: Robust worms, coiled, climate and vegetation were much different then usually tightly resemblingfirst-stage pinworms (Fig. 1). Mouth sur- now. One dung ball, containing unsporulated roundedby 3 bilobed lips (Fig. 2); tail sharplypointed coccidian oocysts and schistosomelike eggs, has (Fig. 3). Posteriorend of esophagusexpanded into a been '4C-dated at 10,500 + 180 yr; the other 3 bulb, characteristicof the order(Fig. 1, arrow).Length = 13-20 x 126-198 x infected samples were not dated. The weighted (n 7) (16.8 159). average of the youngest 10 samples of dung de- Taxonomic summary posited in the cave was within a few years of Type host: Shastaground sloth, N. shastensis. 11,000 BP (before present) (Martin et al., 1985). Type locality: Rampart Cave, ca. 7 km upstream All specimens are presumed to be of this age or from Pierce's Ferry, 36?06'N, 113?56'W,Grand Can- yon, Arizona. greater. Material deposited: Phototype of representative found in 4 of the 7 sam- Parasites were (57%) structuresfrom 1 juvenile worm and 10 syntypes in ples. There were juveniles of 2 nematode species, the USNMPC No. 82076. 2 of coccidian and of morphotypes oocysts, eggs Remarks 2 helminth species. spores and pollen were Fungal One ball contained nematode abundant and had to be differentiated from dung many juveniles pos- (about 10 worms/dropof sediment)that resemblefirst- sible oocysts. The condition of the nematodes stagepinworms. Adult nematodes were not found, sug- precluded detailed observations and measure- gesting that this does not representa coprophagous, ments. free-livingform. This species was found in only 1 of the 7 balls examined. The International Code of Zoological Nomen- dung clature (Art. 42 (b)(i)) (Ride et al., 1985: 83) makes shastensis n. provision for "... certain assemblages of taxo- Strongyloides sp. (Figs.4-8) nomic convenience known as 'collective groups'." Slender coiled Included are known from fossil trac- Description: worms,only slightly (Fig. species only 4). Head small, lacking conspicuous lips. Esophagus es (ichnotaxa) and those known only from im- usuallydetached from mouth and difficultto interpret mature specimens, adult forms of which cannot (Fig. 5);in some specimensin which not detachedfrom be determined. Any of those can be reallocated mouth, a pair of rodlikestructures are present(Fig. 6). Esophagusthicker at with intestine, to the correct genus if it is later discovered. Ex- junction repre- senting a pseudobulb.Tail (Fig. 7) long and pointed, 1892 amples given are Agamodistomum Stossich, resemblinga first-stagejuvenile of Strongyloides.Some (Trematoda), Agamofilaria Stiles, 1907 (Nema- worms in process of molting (Fig. 8). Length (n = 7) toda), and Stelloglyphus Vyalov, 1964 (trace fos- 23-27 x 270-349 (24.4 x 305.3). use sil), among others. Presumably, the frequent Taxonomic summary of as a to these collective "agamo" prefix group Type host: Shastaground sloth, N. shastensis. genera stems from its derivation: a- (Gr., not, Type locality: Rampart Cave, ca. 7 km upstream without) and gam- (Gr., marriage). In applying from Pierce'sFerry, 36?06'N, 113?56'W,Grand Can- names to the organisms we found, we sought to yon, Arizona. Material deposited: of follow this basic concept. Phototype representative structuresfrom 1juvenile wormand about 10 syntypes Numerous bodies, presumably helminth eggs in the USNMPC No. 82075. representing 2 species, were found in 3 dung balls. Remarks One kind (Fig.
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