NEW RECORDS OF THE ENDANGERED TEHUANTEPEC JACKRABBIT (LEPUS FLAVIGULARIS) FROM OAXACA, MEXICO

112 vol. 51, no. 1The Southwestern Naturalist

GEOGRAPHIC DISTRIBUTION, TAXONOMY, AND CONSERVATIONSTATUS OF NELSON’S WOODRAT (NEOTOMA NELSONI) IN MEXICO

NOE GONZALEZ-RUıZ, JOSE RAMıREZ-PULIDO,* AND HUGH H. GENOWAYS

Universidad Autonoma Metropolitana, Unidad Iztapalapa, Division de C.B.S., Departamento de Biologıa,A.P. 55-535, Mexico 09340, D.F., Mexico (NGR, JRP)

Laboratorio de Cordados Terrestres, Escuela Nacional de Ciencias Biologicas, Instituto Politecnico Nacional,Prolongacion de Carpio y Plan de Ayala, Colonia Santo Tomas, A.P. CON 256, 11340, D.F. Mexico (NGR)

University of Nebraska State Museum and School of Natural Resources, University of Nebraska–Lincoln,Lincoln, NE 68588-0514 (HHG)

*Correspondent: [email protected]

ABSTRACT We examined a specimen of Neotoma nelsoni collected in the Mexican state of Puebla,which represents the third known record for the species and the first one for the state. Thedistribution of the species is restricted to the eastern slopes of the volcanoes Pico de Orizaba andCofre de Perote, mainly within the cloud forest. The specimen of N. nelsoni has external andcranial characteristics that are different from other species of the genus. The distribution of thespecies is small, the sizes of populations are small, and the populations exist in geographic isola-tion. None are located within legally protected areas, which presents problems for the conservationof the species.

RESUMEN Examinamos un ejemplar de Neotoma nelsoni colectado en el estado de Puebla, Mex-ico, que representa el tercer registro de la especie y el primero para el estado. La distribucion dela especie se restringe a la ladera este de los volcanes Pico de Orizaba y Cofre de Perote, princi-palmente en el bosque mesofilo de montana. Neotoma nelsoni presenta caracterısticas externas ycraneales que la hacen distinguible de las otras especies del genero. La distribucion de la especiees pequena, los tamanos poblacionales son bajos, y las poblaciones existen en aislamiento geo-grafico. Ninguna esta localizada dentro de areas protegidas, lo que presenta problemas para laconservacion de la especie.

The woodrat, Neotoma nelsoni, is endemic toMexico, is a rare species, and is restricted tothe state of Veracruz (Hall and Dalquest, 1963;Hall, 1981; Prieto and Sanchez, 1993; Gonza-lez-Christen et al., 2002). This species was orig-inally described by Goldman (1905) based on11 specimens, the holotype, and 10 paratypes,from Perote, Veracruz; 58 years later, Hall andDalquest (1963) studied this material and con-firmed the taxonomic status of the species, butthey added no specimens. After 97 y, the rec-ord of Gonzalez-Christen et al. (2002) added aspecimen collected at Barranca Grande, Mu-nicipality of Cozautla (970 m elevation), Vera-cruz. This locality is 30 km southeast from Pe-rote, Veracruz, and represents the secondknown locality for the species.

During fieldwork conducted in the state ofPuebla, a new specimen of N. nelsoni was cap-tured at 5 km southwest of Huaxcaleca(198129080N, 97809450W) at 2,770 m elevation

in the Municipality of Chichiquila, Puebla.This locality does not greatly increase the geo-graphic range for the species because it is only16 km to the south-southwest of BarrancaGrande, Veracruz (Gonzalez-Christen et al.,2002; Fig. 1). However, this record representsthe first for the state of Puebla. Also, it consti-tutes only the third locality for the species, itis the southernmost locality in its known dis-tribution, and it is from the highest known el-evation. This specimen was trapped at night on27 January 1982 in the same trapline as 2 in-dividuals of Oryzomys chapmani and 2 of Reith-rodontomys sumichrasti. This specimen was cap-tured with a Sherman trap and using oatmealcombined with peanut butter as bait. The spec-imen was prepared in the conventional man-ner (Hall, 1981), and it is housed in the mam-mal collection of the Universidad AutonomaMetropolitana Unidad Iztapalapa (3865 UAM-I). Additionally, one of us (NGR) visited the

March 2006 113Notes

FIG. 1 Geographical distribution of Neotoma nelsoni. 1) Perote, Veracruz (type locality; Goldman, 1905);2) Barranca Grande, Municipio de Cozautla, Veracruz (Gonzalez-Christen et al., 2002); 3) 5 km southwestof Huaxcaleca, 2,770 m elevation, Puebla. The dots are cities and small towns. The contour lines are each400 m apart.

mammal collection of the Instituto de Investi-gaciones Biologicas de la Universidad Veracru-zana (IIBUV) to examine the specimen re-ported by Gonzalez-Christen et al. (2002).

Nelson’s woodrat has a combination of cra-nial and external characteristics that make itmorphologically distinctive from the other spe-cies of Neotoma: medium size for the genus; up-per parts cinnamon color, heavily overlaid withsmoky brown, this color predominating on topof head and along dorsally, becoming grayishbrown on cheeks and middle of face; seeming-ly clearer sides than the upper parts; underparts smoky gray; pectoral spot cinnamon;white-throat; tail monocolor, scaly, with little

hair; hind feet smoky gray to the middorsalmetatarsus, toes are dirty white. The skull of N.nelsoni is large; nasals with wedge-shaped pos-terior ending and expanded anterior; fronto-parietal region large and concave; supraorbitalborders remarkably beaded; palatal bone de-cidedly shorter than incisive foramina and end-ing with a short median spine; ascendingbranches of premaxilae long, the ends reach-ing posteriorly beyond nasal (2.5 mm), ap-proaching a line parallel to the interorbitalconstriction.

Neotoma nelsoni can be distinguished from N.leucodon and N. mexicana, 2 species with real orpotential sympatry, by its having upper parts


cinnamon color, under parts smoky gray, andcinnamon pectoral spot. The hair of the upperparts of N. nelsoni is shorter and silkier, tail isless haired and monocolor, tarsal and metacar-pal are smoky gray, and toes are white, nasalsare wedge-shaped posteriorly, and palatinebone has a short posterior spine.

Hall and Genoways (1970) associated N. nel-soni with the albigula species group of the sub-genus Neotoma along with N. albigula, N. pala-tina, and N. varia. They noted the peripheraldistribution of the latter 2 species to N. albigulaand the distinctly isolated location of N. nelsoni,with N. a. leucodon coming no closer than thestate of Hidalgo. Subsequently, Edwards et al.(2001; see also Planz et al., 1996) demonstrat-ed that 2 cryptic species are present in the tax-on formerly known as N. albigula. They appliedthe species name N. leucodon to populations inMexico east and south of Rıo Conchos, NewMexico east of the Rio Grande, and Texas, re-stricting the name N. albigula to populations innorthwestern Mexico, New Mexico west of theRio Grande, Arizona, California, and Utah. Ed-wards et al. (2001) placed N. leucodon and N.micropus in the micropus species group and N.albigula, N. floridana, and N. goldmani in thefloridana species group. On geographicgrounds, N. nelsoni would seem to be associat-ed with N. leucodon of the micropus speciesgroup, but given the complex evolutionary his-tory of this group envisioned by Edwards et al.(2001), this placement must be consideredpreliminary.

The external and cranial measurements(mm) of the adult female (3865 UAM-I) andadult male (1510 IIB UV) are, respectively: to-tal length 390, 380; length of tail vertebrate180, 198; length of hind foot 40, 42; length ofear 30, 27; occipitonasal length 47.6, 45.6; na-sal length 18.8, 16.5; length of incisive foramen10.6, 10.6; depth of cranium 13.6, 13.6;breadth of auditory bulla 6.7, 7.5; length of au-ditory bulla 6.5, 6.6; breadth of braincase 17.6,17.6; zygomatic breadth 24.4, 23.2; mastoidbreadth 17.9, 17.4; interorbital constriction5.5, 5.8; breadth of rostrum 7.6, 7.1; length ofrostrum 20.1, 18.3; length of maxillary tooth-row 10.1, 9.6; length of dentary 22.3, 24.3;height of articular process 11.4, 11.3; length ofmandibular toothrow 9.0, 9.1.

The locality of 5 km southwest of Huaxca-leca (2,770 m elevation) is surrounded by hills

with steep slopes. The place is a small area ofcloud forest with coffee plantations. The pre-dominant climate is humid and warm, withrains during all months of the year (INEGI,2000). The locality of Barranca Grande is inthe same region, but the vegetation is a tropi-cal rain forest (Gonzalez-Christen et al., 2002);however, the most contrasting vegetation typeis that of the type locality of Perote, Veracruz,where the vegetation was xeric shrub. Accord-ing to Hall and Dalquest (1963), ‘‘Nelson andGoldman found this rat in the cactus at Perote.The cactus is now nearly gone, as a result ofclearing the desert for wheat, corn and ma-guey. We failed to take specimens.’’

Given the ecological setting where the 2most recent specimens have been found, theoriginal description (Goldman, 1905) shouldbe reexamined. It must be noted that aroundPerote it is possible to encounter several typesof vegetation, including xeric shrub, pine-oak(Pinus-Quercus) forest, cloud forest, and tropi-cal rain forest in the deep ravines. BecauseGoldman (1951) referred all specimens col-lected in this area to Perote, Veracruz, it is pos-sible that the type series was collected in aplace other than xeric shrub. To support thisidea, Hall and Dalquest (1963) collected morethan 250 rodent specimens, and we collectedover 450 in the Perote region, but no speci-mens of N. nelsoni were obtained. Another ex-planation for this discrepancy could be that N.nelsoni has a high environmental plasticity andcan live in a variety of vegetation types.

The geographical distribution of this speciesis restricted to a small region along the easternslopes of the volcanoes of Orizaba and of theCofre de Perote (Fig. 1), with an elevationalrange between 970 and 2,770 m. We estimatethe territory of this species is a relatively smallarea of 1,350 km2, almost the same as for Per-omyscus bullatus (1,200 km2 Gonzalez-Ruız etal., 2005) and less than that recorded for Sper-mophilus perotensis (2,427 km2; Valdes and Ce-ballos, 1997), 2 other endemic species in theeastern part of Mexico. The distance betweenthe 3 known populations of N. nelsoni rangesfrom 16.6 to 38.4 km. Current agricultural ac-tivities probably increased the isolation amongthese populations and might contribute to adecline in the population size of this rare spe-cies (known from only 13 specimens).

The combination of a small geographical

March 2006 115Notes

range, the isolation of the remaining suitablehabitat, and the low population size in the sam-pled areas suggest that gene flow among theremaining populations has been reduced andmight no longer exist. There is evidence thatisolated and small populations often have ahigher risk of extinction from loss of geneticdiversity, anthropogenic and environmentalprocesses, or disease (Wilcox and Murphy,1985; Ceballos and Rodrıguez, 1993; Arita etal., 1997). Neotoma nelsoni has been registeredas threatened and placed under special protec-tion by the Mexican government (SEMARNAT,2002), and it was placed on the endangeredspecies list by the International Union for Con-servation of Nature in 1996 (IUCN, 2003). Fi-nally, the low density, restricted geographicaldistribution, fragmentation of the habitat, andisolation of localities can be serious conserva-tion problems that confront this species, butan equally important problem is the absenceof biological information, because any conser-vation strategy must be directly linked with theavailable knowledge of biological data for thespecies.

The distribution of N. nelsoni is in the areaof the national parks of Cofre de Perote andPico de Orizaba, but no specimen was foundinside the limits of these protected areas. Thisrepresents a problem because the area ofcloud forest vegetation in this part of Mexicohas a high biodiversity with many endemic spe-cies, such as Sorex macrodon, Peromyscus furvus,P. beatae, Habromys simulatus, Megadontomys nel-soni, Microtus quasiater, and Orthogeomys lanius,and all of these species are in unprotected ar-eas. This fact enhances the value of the easternpart of the Cofre de Perote and Pico de Ori-zaba volcanoes for the maintenance of themammalian diversity of Mexico, indicating theneed for the conservation of this region.

We thank A. Gonzalez Christen from the Institutode Investigaciones Biologicas de la Universidad Ver-acruzana (IIBUV) for allowing us to examine thespecimen of Neotoma nelsoni under his care. For fi-nancial support we thanks to Consejo Nacional deCiencia y Tecnologıa (COI-39619Q) and ComisionNacional para el Conocimiento y Uso de la Biodiv-ersidad (BK022, CS005).

LITERATURE CITED

ARITA, H. T., F. FIGUEROA, A. FRISCH, P. RODRIGUEZ,AND K. SANTOS-DEL PRADO. 1997. Geographical

range size and the conservation of Mexicanmammals. Conservation Biology 11:92–100.

CEBALLOS, G., AND P. RODRıGUEZ. 1993. Diversidad yconservacion de los mamıferos de Mexico: II. Pa-trones de endemicidad. In: R. A. Medellın andG. Ceballos, editors. Avances en el estudio de losmamıferos de Mexico. Asociacion Mexicana deMastozoologıa, Publicaciones Especiales 1:1–464.Pages 87–108.

EDWARDS, C. W., C. F. FULHORST, AND R. D. BRADLEY.2001. Molecular phylogenetics of the Neotoma al-bigula species group: further evidence of a para-phyletic assemblage. Journal of Mammalogy 82:267–279.

GOLDMAN, E. A. 1905. Twelve new wood rats of thegenus Neotoma. Proceedings of the Biological So-ciety of Washington 18:27–34.

GOLDMAN, E. A. 1951. Biological investigations inMexico. Smithsonian Miscellaneous Collection115:1–476.

GONZALEZ-CHRISTEN, A., S. GAONA, AND G. LOPEZ-OR-TEGA. 2002. Registros adicionales de mamıferospara el Estado de Veracruz, Mexico. VertebrataMexicana 11:9–16.

GONZALEZ-RUıZ, N., S. T. ALVAREZ-CASTANEDA, AND T.ALVAREZ. 2005. Distribution, taxonomy, and con-servation status of the Perote mouse Peromyscusbullatus (Rodentia: Muridae) in Mexico. Biodi-versity and Conservation 14:3423–3436.

HALL, E. R. 1981. The mammals of North America,volume 2. John Wiley and Sons.

HALL, E. R., AND W. W. DALQUEST. 1963. The mam-mals of Veracruz. University of Kansas Publica-tions, Museum of Natural History 14:165–362.

HALL, E. R., AND H. H. GENOWAYS. 1970. Taxonomyof the Neotoma albigula-group of woodrats in cen-tral Mexico. Journal of Mammalogy 51:504–516.

INEGI. 2000. Sıntesis Geografica del Estado dePuebla. Instituto Nacional de Estadıstica, Geogra-fıa e Informatica.

IUCN. 2003. 2003 IUCN red list of threatened spe-cies. IUCN, Gland, Switzerland. http://www.redlist.org (Accessed: 16 August 2004).

PLANZ, J. V., E. G. ZIMMERMAN, T. A. SPRADLING, AND

D. R. AKINS. 1996. Molecular phylogeny of Neo-toma floridana species group. Journal of Mam-malogy 77:519–535.

PRIETO BOSCH, M., AND V. SANCHEZ-CORDERO. 1993.Sistemas de informacion geograficos: Un caso deestudio en Veracruz. In: R. A. Medellın and G.Ceballos, editors. Avances en el estudio de losmamıferos de Mexico. Asociacion Mexicana deMastozoologıa, Publicaciones Especiales 1:1–464.Pages 455–464.

SEMARNAT. 2002. Norma Oficial Mexicana NOM-059-ECOL-2001, Proteccion ambiental-Especiesnativas de Mexico de flora y fauna silvestres-Cate-gorıas de riesgo y especificaciones para su inclu-


sion, exclusion o cambio-Lista de especies enriesgo. Diario Oficial de la Nacion, Marzo 6,2002. Pages 56–85.

VALDEZ, M., AND G. CEBALLOS. 1997. Conservation ofendemic mammals of Mexico: the Perote groundsquirrel (Spermophilus perotensis). Journal of Mam-malogy 78:74–82.

WILCOX, B. A., AND D. D. MURPHY. 1985. Conserva-tion strategy: the effects of fragmentation on ex-tinction. American Naturalist 12:879–887.

Submitted 28 September 2004. Accepted 1 August 2005.Associate Editor was Cody W. Edwards.

NEW RECORDS OF THE ENDANGERED TEHUANTEPEC JACKRABBIT(LEPUS FLAVIGULARIS) FROM OAXACA, MEXICO

CONSUELO LORENZO,* FERNANDO A. CERVANTES, FELIPE BARRAGAN, AND JULIETA VARGAS

Departamento de Ecologıa y Sistematica Terrestres, El Colegio de la Frontera Sur, Carretera Panamericana y PerifericoSur s/n 29290, San Cristobal de Las Casas, Chiapas, Mexico (CL, FB)

Departamento de Zoologıa, Instituto de Biologıa, Universidad Nacional Autonoma de Mexico Coyoacan, 04510,Mexico, D.F. Mexico (FAC, JV)


ABSTRACT We obtained new records of the Tehuantepec jackrabbit, Lepus flavigularis, in thestate of Oaxaca, Mexico. The records represent 3 new populations of L. flavigularis in the southernpart of the Isthmus of Tehuantepec. The total area where the new records of L. flavigularis oc-curred is 67 km2. It is necessary to conduct additional studies to obtain details of the actualdistribution of the endangered L. flavigularis and recommend actions for its conservation.

RESUMEN Encontramos nuevos registros de la presencia de la liebre de Tehuantepec, Lepusflavigularis, en el estado de Oaxaca, Mexico. Los registros representan 3 nuevas poblaciones de L.flavigularis en la parte sur del Istmo de Tehuantepec. El area total de ocupacion de los nuevosregistros de L. flavigularis es de 67 km2. Es necesario realizar investigaciones adicionales paradefinir a detalle la distribucion actual de la especie en peligro L. flavigularis y recomendar accionespara su conservacion.

The endangered Tehuantepec jackrabbit, Le-pus flavigularis, is an endemic species of thestate of Oaxaca, Mexico. It is known to occurin only 6 localities: Santa Marıa del Mar, SanMateo del Mar, San Pedro Huilotepec, SantaEfigenia, Santo Domingo Tehuantepec, andReforma de Pineda (Anderson and Gaunt,1962; Dixon et al., 1983; Uribe-Alcocer et al.,1989; Cervantes, 1993; Cervantes and Yepez,1995). Therefore, the range of this species ishighly restricted and circumscribed to theSouthern Isthmus of Tehuantepec, resulting ina listing of Endangered in the IUCN Red DataBook (IUCN, 2004). Furthermore, Cervanteset al. (2002) reported that L. flavigularis hadthe lowest levels of genetic variation among Le-pus species they examined. These demograph-ic and genetic characteristics suggest a high po-tential for extinction. Accordingly, we wished

to determine whether there existed additionalpopulations of the species in addition to thosereported in the literature.

We collected 4 individuals in the Municipal-ity of San Dionisio del Mar in open xeric veg-etation dominated by brush (Byrsonima crassi-folia) and grasses (Bouteloua and Paspalum pli-catulum; Table 1). Similarly, 4 individuals werecaught in the Municipality of San Francisco delMar in open xeric vegetation dominated bygrasses. Five additional jackrabbits were col-lected in dune habitat of the Municipality ofSan Francisco Ixhuatan (Table 1). Voucherspecimens were deposited in the Mammal Col-lection of El Colegio de la Frontera Sur (ECO-SC-M) at San Cristobal de Las Casas, Chiapas,Mexico, and in Coleccion Nacional de Mamı-feros (CNMA) of Instituto de Biologıa, Univer-

March 2006 117Notes

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FIG. 1 Three populations of Lepus flavigularis around the Laguna Inferior in the South Isthmus of Te-huantepec, Oaxaca, Mexico. Dots indicate locality records of L. flavigularis.

sidad Nacional Autonoma de Mexico, in Mex-ico City.

Reforma de Pineda and Santa Efigenia arethe 2 collection localities closest to our studyarea where Tehuantepec jackrabbit was record-ed previously. The distances between theseaforementioned localities and the closest newrecords we report are 30.5, 30.9, and 28.2 kmbetween Reforma de Pineda and San Dionisiodel Mar, San Francisco del Mar, and San Fran-cisco Ixhuatan, respectively; 48.2, 46.4, and37.3 km between Santa Efigenia and San Dion-isio del Mar, San Francisco del Mar, and SanFrancisco Ixhuatan, respectively.

We computed the area occupied by these newrecords or the area where this species is presentwithin the total area sampled during the col-lecting period (1998 to 2003). The study area

comprised 673 km2 and was divided into 1-km2

units. The location of each individual was ob-tained with a Geographical Positioning Systemunit and recorded in a spreadsheet. Data wereexported to ArcView 3.2 (Environmental Sys-tems Research Institute, Redlands, California)to map and quantify the distributional range oc-cupied by Tehuantepec jackrabbit. The result-ing occupation area was 67 km2 (Fig. 1).

Both adult and young Tehuantepec jackrab-bits were observed in the study area. Our ob-servational data, along with reproduction dataobtained from collected individuals (Table 1)indicated 3 well established populations of Te-huantepec jackrabbit; however, we have docu-mented low reproductive potential because fe-males apparently carry only one embryo.

The finding of additional populations of Te-

March 2006 119Notes

huantepec jackrabbit, a species whose prospectof survival is bleak at best, is an important de-velopment for its conservation. However, thesepopulations are located in areas of high hu-man population pressure. The needs to ade-quately circumscribe the range of the speciesand assess its basic population ecology remain.

We thank M. Aquino, J. Bolanos, and the Muni-cipalities of San Dionisio del Mar, San Francisco delMar, and San Francisco Ixhuatan, for their help dur-ing fieldwork. The families Gutierrez Lopez and Gu-tierrez Vazquez housed our crew. Chicago Zoologi-cal Society and Lincoln Park Zoo Neotropic Fundpartly funded this research. E. Valencia digitalizedmaps and assisted with software. Instituto Nacionalde Ecologia, SEMARNAT, issued the special researchpermit DOO.02.0374 to collect L. flavigularis.

LITERATURE CITED

ANDERSON, S., AND A. GAUNT. 1962. A classificationof the white-sided jackrabbits of Mexico. Ameri-can Museum Novitates 2088:1–16.

CERVANTES, F. A. 1993. Lepus flavigularis. MammalianSpecies 423:1–3.

CERVANTES, F. A., C. LORENZO, AND T. L. YATES. 2002.Genic variation in populations of Mexican lago-morphs. Journal of Mammalogy 83:1077–1086.

CERVANTES, F. A., AND L. YEPEZ. 1995. Species rich-ness of mammals from the vicinity of Salina Cruz,coastal Oaxaca, Mexico. Anales Instituto Biolo-gıa, Universidad Nacional Autonoma Mexico,Serie Zoologıa 66:113–122.

DIXON, K. R., J. A. CHAPMAN, G. R. WILLNER, D. E.WILSON, AND W. LOPEZ-FORMENT. 1983. The NewWorld jackrabbits and hares (genus Lepus). 2. Nu-merical taxonomic analysis. Acta Zoologica Fen-nica 174:53–56.

INTERNATIONAL UNION FOR CONSERVATION OF NATURE

AND NATURAL RESOURCES (IUCN). 2004. 2000IUCN red list of threatened animals. IUCN Spe-cies Survival Commission, Gland, Switzerland.http://www.redlist.org (Accessed: January 2004).

URIBE-ALCOCER, M., F. A. CERVANTES, C. LORENZO-MONTERRUBIO, AND L. GUERENA-GANDARA. 1989.Karyotype of the tropical hare. Southwestern Nat-uralist 34:304–306.

Submitted 14 September 2004. Accepted 28 June 2005.Associate Editor was Cody W. Edwards.

BOBCAT ATTACK ON A COTTONTAIL RABBIT

DEAN E. BIGGINS* AND DENISE M. BIGGINS

U.S. Geological Survey, Fort Collins Science Center, 2150 Centre Avenue, Fort Collins, CO 80526*Correspondent: [email protected]

ABSTRACT We observed an attack by a bobcat (Lynx rufus) on a cottontail rabbit (Sylvilagus)that involved stealthy approach by the cat for .1 h, followed by a 12.3-s chase covering 116.0 mfor the cat and 128.4 m for the rabbit. During the chase, the route of the cat from starting pointto kill site was more direct than the semi-circular route of the rabbit. Stride lengths for the catand total distance covered by the chase were longer than those previously reported for bobcats.

RESUMEN Observamos como ataco un lince (Lynx rufus) a un conejo cola de algodon (Silvila-gus). El lince se aproximo cautelosamente por un poco mas de una hora seguidamente persiguien-do al conejo por 12.3 segundos, cubriendo 116.0 m para el lince y 128.4 m para el conejo colade algodon. Durante el ataque, la ruta del lince desde el punto de comienzo hasta el sitio dondemato al conejo fue mas directo que la ruta semicircular del conejo. La distancia de los pasos dellince y la distancia total cubierta para la caza, fueron mas largas que aquellas reportadas previa-mente para los linces.

Bobcats (Lynx rufus) are distributed widelyin North America, but there are few publishedaccounts detailing direct observations of thehunting behaviors of free-ranging, wild bob-

cats. We observed an interaction between abobcat and a cottontail rabbit (Sylvilagus) thatlasted .1 h, beginning with the initial detec-tion of the rabbit, progressing through a multi-


FIG. 1 Movements of cottontail rabbit (Sylvilagus)and bobcat (Lynx rufus) prior to and during an at-tack by the bobcat. During the stalk, the bobcatmoved from points 1 through 4 as the cottontail rab-bit fed at sites A and B.

stage stalk, a high-speed chase, and capture ofthe rabbit.

The attack occurred on 7 February 2004 inthe Rocky Mountain foothills 4 km south of Liv-ermore, Colorado (40.76808N, 105.19338W), ina shrub-grassland community (Fig. 1). Thegrasslands were a mixture of cheatgrass (Anisan-tha tectorum), western wheatgrass (Pascopyrumsmithii), needle-and-thread (Hesperostipa comata),green needlegrass (Nassella viridula), and bluegrama (Chondrosum gracilis). The site is aboutmidway between 2 small streams separated byabout 400 m, each with dense riparian vegeta-tion consisting of an overstory of cottonwoods(Populus deltoides and P. angustifolia) and shrubthickets of American plum (Prunus americana),hawthorn (Crataegus macracantha), and choke-cherry (Padus virginiana).

On the morning of the attack, there was nowind, there were scattered clouds, the temper-ature was 228C, and snow depths were 2 to 8

cm. We used a measuring wheel of 2-m circum-ference to measure distances along snow tracksand a global positioning receiver to record keypoints and boundaries of vegetative covertypes.

We first noticed the bobcat at 0735 h, sittingupright in a patch of grassland (Fig. 1, Point1). At 0739 h, the bobcat changed its behaviorto a more rigid posture, focusing its attentionon a point in dense grass cover. The bobcatassumed a low crouch at 0742 h and began tomove slowly westward, freezing in mid-strideon several occasions. We noticed a cottontailrabbit at 0743 h (Fig. 1, Point A). The stealthymovements of the bobcat continued intermit-tently until 0748 h, when it sat upright (Fig. 1,Point 2); the bobcat had shortened the dis-tance separating it from the rabbit from an ini-tial 25.1 m to 10.1 m. The rabbit began movingnorthward, although it seemed not to have de-tected the bobcat, feeding intermittently. A fewminutes after the rabbit had stopped at a newfeeding site (Fig. 1, Point B), the bobcat beganto move northerly (0804 h, estimated) in aslow, upright walk, and crawled under an over-hanging sandstone ledge at 0807 h (Fig. 1,Point 3). A western scrub jay (Aphelocoma cali-fornica) circled about 3 m above the bobcatand scolded it at about 0810 and again at 0830h, but neither the bobcat nor the cottontailseemed to respond. The bobcat remained in asitting position, partially under the ledge,watching the cottontail. At 0832 h, the bobcatbegan to move slowly to the north in a lowcrouch, reducing the distance to the cottontailby 5.8 m; the 2 animals were separated by 8 mat 0835 h, and the bobcat was sitting uprighton a snow-free slab of sandstone (Fig. 1, Point4). At 0840 h, the cottontail moved behind ashrub and might have detected the bobcat.The reaction of the bobcat was to rise to astanding position, from which it took 2 stepsand a short leap toward the cottontail (about3 m total), causing the cottontail to take flightto the north. After 2 additional strides by thebobcat, the cottontail circled to the west andthe cat abruptly changed its course by almost90 degrees to parallel the course of the rabbit.At the time their courses converged, the 2 an-imals were separated by about 3 m. The bobcatclosed that distance in the next several sec-onds. At the point of presumed contact (Fig.1, Point 5), the 2 animals became concealed

March 2006 121Notes

by shrubs, preventing us from seeing the cap-ture. After 2 h had elapsed, we visited the sitewhere the bobcat had caught the cottontail,finding interrupted skid marks in the snowover the final 6-m distance of the chase (Fig.1, Points 5–6) and bits of rabbit fur at the stop-ping point. A burrow surrounded by rabbittracks (perhaps the intended destination ofthe cottontail) was located 6.7 m from the stop-ping point. Along the 4.1-m route from theend of the skid marks to the feeding site (Fig.1, Point 7), there were intermittent drag marksleft in the snow alongside the tracks of the cat,perhaps made by one of the legs of the cotton-tail. The only remains at the feeding site werea 14-cm length of the gastrointestinal tract, in-cluding the grass-filled stomach, and a 6 cm by6 cm piece of skin.

The distance traveled by the bobcat duringthe chase (Fig. 1, Points 4–5) was 116.0 m. Thecottontail traveled 128.4 m. Pursuit of the cot-tontail along its course would have resulted ina path that was 20.4 m longer for cat, likelyresulting in a failed capture attempt given theproximity of the capture site to a burrow en-trance. Prior experience might have enabledthis bobcat to predict the path of the cotton-tail, a species that commonly circles when pur-sued (Madson, 1959). The cottontail did notengage in the abrupt, small-scale changes indirection commonly described for cottontailspursued by predators (‘‘zig-zags’’ of Cahalane,1954:612). Perhaps escape tactics vary accord-ing to predator, habitat, or species of Sylvilagus;S. floridanus, S. audubonii, and S. nuttallii aresympatric in our area (Fitzgerald et al., 1994),and we are uncertain which species we ob-served.

Along the route of the bobcat were 2 stridesof 5.2 m and a final leap of 5.5 m, longer thanextreme leaps previously reported (Cahalane,1954; Young, 1958). Except for a few shortstrides at the beginning of the chase, moststrides by the bobcat were .4.3 m. The bobcatjumped over clumps of mountain mahogany(Cercocarpus montanus, height about 1 m) at 2points along its path. The longest leaps by thecottontail were 3.4 m and 3.5 m, but most ofits strides were much shorter. The chase wasuphill, rising from an elevation of 1,828 m to1,837 m.

We estimated the elapsed time for the chaseseveral hours later while the details of the

event were easily recalled. We envisioned theprogress of the cottontail and the bobcat fromthe elevated vantage point of our original ob-servation (a window of our home), using astopwatch to time the chase as we visualizedthe animals traversing their respective routesto the point of contact. Our independently es-timated means (12.66 s and 11.93 s) from mul-tiple replicates (6 for DEB, 5 for DMB) did notdiffer significantly (t 5 0.756, P 5 0.469), sowe used the overall mean of 12.27 s (CI0.95,11.28 to 13.38 s) as a reasonable estimate ofelapsed time for the chase. Estimated averagespeeds were 9.45 m/s (21.1 miles per h) forthe bobcat and 10.46 m/s (23.4 miles per h)for the cottontail. Although our reliance onmental recall is not ideal, credibility of themethod is enhanced by the similar result from2 observers, who did not communicate untilsampling was completed. Speeds of 6.71 to13.41 m/s (15 to 30 miles per h) are claimedfor Sylvilagus in a wide variety of sources, butwe have found just one often-cited estimate (15miles per h) to be supported by a descriptionof the method used for estimation. A desertcottontail (S. audubonii) observed from a trainwas running ‘‘about three-fourths as fast’’ asthe speed of the train which ‘‘was estimated tobe 20 miles an hour’’ (Grinnell and Storer,1924:224). An accompanying statement aboutthe rarity of opportunities to judge runningspeeds is especially germane in the case we ob-served, where the cottontail was attempting toescape a natural predator.

The relatively long chase we witnessed wasunlike sequences of bobcat predatory behaviordescribed by others (e.g., Marshall and Jen-kins, 1967; Foldesh, 1982). Typical huntingstrategy for bobcats often involves stealth, sur-prise, and ambush (Cahalane, 1954; Jones etal., 1983; Fitzgerald et al., 1994; Lariviere andWalton, 1997; Anderson and Lovallo, 2003),with abandonment of the pursuit if a shortrush or pounce is unsuccessful (Cahalane,1954; Sunquist and Sunquist, 2002).

We appreciate the abstract translation by C. Millerand helpful suggestions from J. Truett, J. Hasler, andM. Hasler.

LITERATURE CITED

ANDERSON, E. M., AND M. J. LOVALLO. 2003. Bobcatand lynx. In: G. A. Feldhamer, B. C. Thompson,


and J. A. Chapman, editors. Wild mammals ofNorth America: biology, management, and con-servation, second edition. Johns Hopkins Univer-sity Press, Baltimore, Maryland. Pages 758–786.

CAHALANE, V. H. 1954. Mammals of North America.Macmillan Company, New York.

FITZGERALD, J. P., C. A. MEANEY, AND D. M. ARMS-TRONG. 1994. Mammals of Colorado. UniversityPress of Colorado, Niwot.

FOLDESH, W. E. 1982. An attack by bobcats on anadult javelina. Southwestern Naturalist 27:457.

GRINNEL, J., AND T. I. STORER. 1924. Animal life inthe Yosemite: an account of the mammals, birds,reptiles, and amphibians in a cross-section of theSierra Nevada. University of California Press,Berkeley.

JONES, J. K., JR., D. M. ARMSTRONG, R. S. HOFFMAN,AND C. JONES. 1983. Mammals of the northernGreat Plains. University of Nebraska Press, Lin-coln.

LARIVIERE, S., AND L. R. WALTON. 1997. Lynx rufus.Mammalian Species 563:1–8.

MADSON, J. 1959. The cottontail rabbit. Olin Mathie-son Chemical Corporation, East Alton, Illinois.

MARSHALL, A. D., AND J. H. JENKINS. 1967. Movementsand home ranges of bobcats as determined byradio-tracking in the upper coastal plan of west-central South Carolina. In: J. W. Webb, editor.Proceedings of the Twentieth Annual Confer-ence of the Southeast Game and Fish Commis-sioners, Columbia, South Carolina. Pages 206–214.

SUNQUIST, M., AND F. SUNQUIST. 2002. Wild cats ofthe world. University of Chicago Press, Chicago,Illinois.

YOUNG, S. P. 1958. The bobcat of North America.Wildlife Management Institute, Stackpole, Har-risburg, Pennsylvania.

Submitted 13 May 2004. Accepted 8 June 2005.Associate Editor was Cheri A. Jones.

MOLECULAR IDENTIFICATION OF COYOTE FECESIN AN URBAN ENVIRONMENT

PAUL R. KRAUSMAN,* MARTHA I. GRINDER, PHILIP S. GIPSON, GERALD L. ZUERCHER,AND GEORGE C. STEWART

Wildlife Conservation and Management Program, School of Natural Resources, University of Arizona,Tucson, AZ 85721 (PRK, MIG; MIG deceased 16 July 1999)

Kansas Cooperative Fish and Wildlife Research Unit, U.S. Geological Survey, Division of Biology,Kansas State University, Manhattan, KS 66506 (PSG, GLZ)

Department of Pathology and Diagnostic Medicine, College of Veterinary Medicine, Kansas State University,Manhattan, KS 66506 (GCS)


ABSTRACT Coyotes (Canis latrans) are common in urban areas throughout North America. How-ever, their urban diets have been studied rarely, possibly because tools to discriminate coyote feces(i.e., scats) from those of domestic dogs have not been available. We used molecular techniquesto discriminate between scats from coyotes and domestic dogs. Our objective was to assess fieldaccuracy in identifying coyote scat from dog scat. We collected and analyzed 34 scats (classified ascoyote scat) via DNA analysis to determine whether they were deposited by coyotes. Ten scatsclassified in the field as coyote were from domestic dogs. We could not distinguish between coyoteand dog scats in Tucson simply by observing scats. The role that available food items play infeeding strategies of coyotes needs to be examined thoroughly to fully understand their ecologyand relationships to dogs and humans in urban areas. DNA analysis used for scat identificationprovides a mechanism to distinguish canid scats.

RESUMEN Los coyotes (Canis latrans) son comunes en las areas urbanas a lo largo de todaNorteamerica. Sin embargo, sus dietas urbanas han sido estudiadas raramente, posiblementeporque las herramientas para distinguir entre las heces coyotes y las de los perros domesticos noestaban disponibles. Usamos tecnicas de biologıa molecular para distinguir entre las heces de

March 2006 123Notes

coyotes y las de perros domesticos. Nuestro objetivo fue de determinar la exactitud en identificarlas heces de coyotes y de perros en el campo. Recogimos y analizamos 34 heces (clasificadas comoheces de coyotes) usando analisis de ADN para determinar si las heces fueron depositadas porcoyotes. Diez heces clasificadas en el campo como heces de coyote provinieron de perros domes-ticos. Simplemente observando heces en el campo, no fue posible distinguir entre las heces decoyotes y las de perros en Tucson. El papel que desempenan los artıculos alimenticios disponiblesen las estrategias de alimentacion de los coyotes necesita ser examinada a fondo para entendercompletamente la ecologıa de los coyotes y su relacion con los perros y los seres humanos en lasareas urbanas. El analisis de ADN usado para la identificacion de las heces proporciona un me-canismo para distinguir heces en la familia Canidae.

Coyotes (Canis latrans) are adaptable carni-vores that can change their diet in response tofood availability (Todd et al., 1981; Andelt etal., 1987; Dibello et al., 1990). Recently, coy-otes have become established in urban areaswith dense human populations (Gill and Bon-nett, 1973; Shargo, 1988; Quinn, 1995; Grinderand Krausman, 2001a), where they have accessto human-related food sources (MacCracken,1982; Quinn, 1997).

Coyotes might play an important role in theecology of human-altered landscapes. Soule etal. (1988) suggested that when coyotes reducethe abundance of house cats and other smallmammals that prey on songbirds, they indi-rectly contribute to the maintenance of nativeavifauna. The extent to which coyotes in Tuc-son, Arizona, prey upon domestic pets cur-rently is unknown. The Arizona Game and FishDepartment (AGFD) receives numerous com-plaints about coyotes in urban Tucson (S. Rich-ardson, Urban Wildlife Specialist, AGFD, pers.comm.). The complaints are frequently aboutpets believed to have been killed by coyotes (G.Frederick, AGFD, unpublished report). Re-searchers in other urban areas have found thatdomestic pets comprise 2 to 14% of the diet ofcoyotes (MacCracken, 1982; Shargo, 1988;Quinn, 1997). With information on coyote di-ets, managers might be able to better under-stand the ecological role of coyotes and do-mestic pets.

Although the diets of coyotes in suburbs ofLos Angeles and San Diego, California, andSeattle, Washington, have been described(MacCracken, 1982; Shargo, 1988; Quinn,1997), the diets of coyotes in urban areas havereceived little attention compared to those ofcoyotes in rural areas (MacCracken, 1984),possibly because researchers have not had a re-liable method for distinguishing the scat of do-mestic dogs from that of coyotes (Halfpenny,

1986; Quinn and Jackman, 1994). Advances inamplifying small amounts of DNA through apolymerase chain reaction (PCR) process, cou-pled with the ability to isolate DNA directlyfrom scat, provides such a technique (Foran etal., 1997; Paxinos et al., 1997; Kohn et al.,1999). Our objective was to assess accuracy ofidentifying coyote scat in areas of sympatrywith domestic dogs in southwestern Arizona.

We collected scats in Tucson, Pima County,Arizona, and in the desert directly outside thecity limits (328099 to 328229N, 1108449 to1118019W). The composition of the study areain Tucson was 65% natural (residential areaswith ,1 house/0.4 ha, state and federal parks,and privately-owned open space), 12% residen-tial (neighborhoods with .1 house/0.4 ha),8% dry river beds, 5% commercial (industrialareas, shopping centers, public buildings, of-fice buildings), 4% parks (schools, cemeteries,zoos, golf courses, neighborhood parks), 3%vacant areas, and 3% roads $4 lanes wide. Tuc-son encompasses about 500 km2 with an esti-mated human population of 475,000 (TucsonUpdate Demography 1999, www.ci.tucson.az.us/planning/update.htm.)

We opportunistically collected intact scats weidentified as from coyotes (Murie, 1975; Dan-ner and Dodd, 1982) along roads, in dry riv-erbeds, and in other areas frequented by coy-otes from June 1998 through May 1999 (Grind-er and Krausman, 2001a). Unless we saw coy-otes or coyote tracks, we did not collect scats,20 mm in diameter to reduce the possibilityof collecting scat of gray fox (Urocyon cinereoar-genteus) (Murie, 1975; Danner and Dodd,1982). We wore plastic gloves during collectionto eliminate the possibility of leaving humanepithelial cells on scats. We placed scats intoindividual plastic collection bags (Whirlpak,NASCO Plastics, New Hamburg, Ontario, Can-


FIG. 1 Gel electrophoresis results from presumedcoyote fecal samples from Tucson, Arizona. Bandsrepresent cytochrome-b fragments from mitochon-drial DNA (mtDNA) after treatment with the Sau 3Arestriction enzyme. M 5 a molecular referencemarker; B 5 a known coyote blood sample; 1, 3, 7–10, 12, 13, 15, 17 5 fecal sample numbers. All fecalsamples showing a coyote triple-band pattern (in thiscase, the known coyote blood sample is B with 3bands below the dark dye mark that extends acrossthe gel) represent coyote mtDNA samples (1, 12, 13)cut by the Sau 3A restriction enzyme. Samples withsingle bands (3, 7, 8, 10) represent domestic dogmtDNA that is not cut by the Sau 3A restriction en-zyme. Samples showing no bands represent inade-quate DNA extraction and were reanalyzed. Note:samples 15 and 17 were later confirmed as coyote;sample 9 was confirmed as domestic dog. The wide,light bands at the bottom of the figure representRNA, probably from plant material in the scats.

ada) labeled with the date and location of col-lection, and we froze them until analysis.

We randomly selected and analyzed 34 scatsvia molecular techniques to assess our accuracyat identifying coyote scats. The DNA was ex-tracted from approximately 60 mg of feces us-ing the IsoQuick extraction kit (ORCA Re-search, Bothell, Washington; Kohn et al.,1999). Species identification was determinedthrough the extraction of the D-loop region(Pilgrim et al., 1998) from the mitochondrialDNA found in sloughed colonic epithelial cells(Albaugh et al., 1992). We compared mito-chondrial D-loop sequences for coyote and do-mestic dog and designed 2 unique primers(canisbcl: 59-TGATCATGGGCTGATTAG-39 andcanisxho: 59-ATGCCTCGAGAAACCATC-39) toamplify a 200 base-pair fragment from both.

We amplified a fragment of the D-loop gene(500 to 1,000 base pairs; Foran et al., 1997)using the following conditions for a 25-mL sam-ple: 2-mL DNA, 14.875-mL sterile water, 2.5-mLof 0.1% bovine serum albumin (BSA), 2.5-mL10X PCR reaction buffer 1-mL of 25-mMolMgCl2, 0.75-mL each of 25-ng/mL canisbcl andcanisxho, 0.5-mL of 10-mMol dNTP’s, and 0.125-mL Taq DNA polymerase. Reactions took placein a programmable thermal cycler (Perkin El-mer Cetus Model 3600, Perkin Elmer Corpo-ration, Norwalk, Connecticut) with a 5-min de-naturation step at 948C; 30 cycles of 948C for60 s, 558C for 45 s, and 728C for 45 s; and a5-min final extension at 728C. The PCR prod-ucts were verified through electrophoresis ona 0.7% agarose gel and examination on an ul-traviolet transilluminator.

The PCR products were digested by the Sau3A restriction enzyme to differentiate betweencoyotes and domestic dogs, by the following1-h, 20-mL reaction: 15-mL PCR product, 2-mL10X restriction enzyme buffer, 1-mL Sau 3A,and 2-mL sterile water. Reactions were subject-ed to electrophoresis on a 2% small fragment-size agarose gel, and enzyme digestion prod-ucts were evaluated on an ultraviolet transillu-minator.

We randomly selected 34 scats from 231 scatscollected from 15 June 1998 though 10 May1999. We extracted DNA from all 34 scats andsubjected them to PCR and restriction enzymeanalyses. All fecal samples (n 5 34) yieldedDNA after extraction with the ORCA kit andPCR product with the designed Canis-specific

(coyote or dog) primers. The Sau 3A restric-tion enzyme consistently discriminated be-tween fecal samples of coyote and dog sources(Fig. 1). Restriction enzymes applied to DNAextracted from feces validated coyote identifi-cations for 24 of 34 samples with the remaining10 samples confirmed as from domestic dogs.

Our molecular analysis indicated that it isnot practical to differentiate coyote scats con-sistently from those of dogs by observation inthe field. Thus, analyses of diets from scatsidentified only by field-observation do not ac-curately reflect the diet of coyotes. About 30%of the 231 scats collected in Tucson were fromdogs.

Most dietary studies of coyotes have beenbased on scats identified in the field. Coyotes

March 2006 125Notes

throughout North America consume a varietyof foods year-round (Voigt and Berg, 1987),and their diet varies regionally (Gipson, 1974;Bekoff, 1982). If our results are consistent inother areas, it is likely that #30% of selectedscats could have been from canids other thancoyotes, which likely would alter results if dietsamong canids differed. DNA analysis used forscat identification provides a technique thatcan be used to distinguish coyote scats accu-rately. Future studies should ensure that col-lected scats are from the target species to pro-vide accurate dietary information.

Understanding the diet of coyotes is impor-tant for better management. For example, stra-tegic plantings of preferred vegetation mightlure coyotes to areas where their presence ismore acceptable and away from areas wherethey are not wanted. Managers also need tobetter understand dietary interactions betweencoyotes and dogs in the urban environmentand the potential for disease and parasitetransmission (Grinder and Krausman, 2001b).

This study was funded by the University of Arizo-na, Tucson, Arizona, and Kansas State University,Manhattan, Kansas.

LITERATURE CITED

ALBAUGH, G. P., V. IYENGAR, A. LOHANI, M. MALAYERI,S. BALA, AND P. P. NAIR. 1992. Isolation of exfoli-ated colonic epithelial cells, a novel, non-invasiveapproach to the study of cellular markers. Inter-national Journal of Cancer 52:347–350.

ANDELT, W. F., J. G. KIE, F. F. KNOWLTON, AND K.CAMPBELL. 1987. Variation in coyote diets associ-ated with season and successional changes in veg-etation. Journal of Wildlife Management 51:273–277.

BEKOFF, M. 1982. Coyote (Canis latrans). In: J. E.Chapman and G. A. Feldhamer, editors. Wildmammals of North America. Johns Hopkins Uni-versity Press, Baltimore, Maryland. Pages 447–459.

DANNER, D. A., AND N. DODD. 1982. Comparison ofcoyote and gray fox scat diameters. Journal ofWildlife Management 46:240–241.

DIBELLO, F. J., S. M. ARTHUR, AND W. B. KROHN. 1990.Food habits of sympatric coyotes, Canis latrans,and bobcats, Lynx rufus, in Maine. CanadianField-Naturalist 104:403–408.

FORAN, D. R., K. R. CROOKS, AND S. C. MINTA. 1997.Species identification from scat: an unambiguousgenetic method. Wildlife Society Bulletin 25:835–839.

GILL, D. A., AND P. BONNETT. 1973. Nature in theurban landscape: a study of city ecosystems. York,Baltimore, Maryland.

GIPSON, P. S. 1974. Food habits of coyotes in Arkan-sas. Journal of Wildlife Management 38:848–853.

GRINDER, M. I., AND P. R. KRAUSMAN. 2001a. Homerange, habitat use, and nocturnal activity of coy-otes in an urban environment. Journal of Wild-life Management 65:887–898.

GRINDER, M. I., AND P. R. KRAUSMAN. 2001b. Morbi-dity mortality factors and survival of an urbancoyote population in Arizona. Journal of WildlifeDiseases 37:312–317.

HALFPENNY, J. 1986. A field guide to mammal track-ing in western America. Johnson Brooks, Boul-der, Colorado.

KOHN, M. H., E. C. YORK, D. A KAMRADT, G. HAUGHT,R. M. SAUVAJOT, AND R. K. WAYNE. 1999. Estimat-ing population size by genotyping faeces. Pro-ceedings of the Royal Society of London B 266:657–663.

MACCRACKEN, J. G. 1982. Coyote food in a southernCalifornia suburb. Wildlife Society Bulletin 10:280–281.

MACCRACKEN, J. G. 1984. Coyote foods in the BlackHills, South Dakota. Journal of Wildlife Manage-ment 48:1420–1423.

MURIE, O. J. 1975. A field guide to animal tracks,second edition. Houghton-Mifflin, Boston, Mas-sachusetts.

PAXINOS, E., C. MCINTOSH, K. RALLS, AND R. FLEISCH-ER. 1997. A noninvasive method for distinguish-ing among canid species: amplification and en-zyme restriction of DNA from dung. MolecularEcology 6:483–486.

PILGRIM, K. L., D. K. BOYD, AND S. H. FORBES. 1998.Testing for wolf-coyote hybridization in the RockyMountains using mitochondrial DNA. Journal ofWildlife Management 62:683–689.

QUINN, T. 1995. Using public sightings to investigatecoyotes use of urban habitat. Journal of WildlifeManagement 59:238–245.

QUINN, T. 1997. Coyote (Canis latrans) food habitsin three urban habitat types of western Washing-ton. Northwest Science 71:1–5.

QUINN, T., AND W. JACKMAN. 1994. Influence of dieton detection of fecal bile acids by thin-layer chro-matography. Journal of Wildlife Management 58:295–299.

SHARGO, E. S. 1988. Home range, movement and ac-tivity patterns of coyotes (Canis latrans) in LosAngeles suburbs. Unpublished Ph.D. dissertation,University of California, Los Angeles.

SOULE, M. E., D. T. BOLGER, A. C. ALBERTS, J. WRIGHT,M. SORICE, AND S. HILL. 1988. Reconstructed dy-namics of rapid extinctions of chaparral-requir-ing birds in urban habitat islands. ConservationBiology 2:75–92.


TODD, A. W., L. B. KEITH, AND C. A. FISCHER. 1981.Population ecology of coyotes during a fluctua-tion of snowshoe hares. Journal of Wildlife Man-agement 45:629–640.

VOIGHT, D. R., AND W. E. BERG. 1987. Coyote. In: M.Novak, J. A. Baker, M. E. Obbard, and B. Malloch,editors. Wild furbearer management and conser-

vation in North America. Ministry of Natural Re-sources, Toronto, Ontario, Canada. Pages 345–357.

Submitted 24 February 2004. Accepted 1 August 2005.Associate Editor was Cheri A. Jones.

NEW RECORDS OF THE ENDANGERED TEHUANTEPEC JACKRABBIT (LEPUS FLAVIGULARIS) FROM OAXACA, MEXICO

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