LATRINE USE BY SAh' JOrZQUIN KIT FOXES (\'CTLPES JlL4CROTISJICTTICA) AND COYOTES (CANIS LATRANS)
Scent marking ~vitliglantlular secretions, Scent marking 1)). kit fi)ses (1: t~~nct-otis)has urine, or fecc3 i3 comnlon in sl~eciesof all car- not 11c.en well tlescril~ed.Egoscue (1962) re- nil ore Ltn~ilies.In general, scent inarks are ported that he found no c\iclt.nce that kit fi~ses not distril~uteclranclomlv But are laced care- regularly mark "'sign' stations such iis olcl fully on ~isuall!conspi~uous andLoften tradi- skeletons or natnral ol)jects." O'Fi~rrell (1987) tionally used o1)jects or locations, such as trail said that kit foxes defecate along trails and intersections (hlitctlollald 1985, Gorman anel near clcns and that they mal-k novel ol)jects 1)y Tro\\lhridge 1989). In some species repeated depositing untlcrsized scats 011 them. HE atldetl, defecation at the same site results in accuniu- "There is no c\.idence that the! systematicall!. lations of feces (scats) in small areas kno\vn as mark either their territorial 1)oundarics or latrines, niidtl~ns,or spriiint piles. Use of 'sin1<, stations' \\it11 uriile or feces." Rcccnt stud- latrines is particularly \\?ell know11 in Euro- ics of scent marking I)! co!.otcs (Cht~isIntt-crn.s) pean I~adgci-s(JlelPs rtleles; Stewart et nl. 2001, ill lre11o\\7stone National Park (Gesc allcl Ruff SOOZ), European otters (Llrtr-n lrrtr-cr; Kruuk 1997, rlllcn ct al. 1999) ~.onclutlcclthat urine l99.5), l>ro\\.n hyenas (H!lcretirr brunnetr; (;or- marking \\.as the most innportant form of ollkc- man and ljlills l984),and spotted hyenas (Cro- tory cunlmunication ant1 that coyotes ditl not clrtcr croclrtcr; llills and C;orman 1987). use scats to mark tcl-ritorics. Urine marking is grnelxll!~thought to l)c IIo\vrvcl; j1.e haw ol~scrvcdman!. accumu- the most ilnportant form of scent marking in lations of kit fos scats in latriiles \vhilc survc!.- callids (Klciman 1966), Ijut accumulations of ing for c.ntlangc.rrd Sa11 Joacpin kit foxes (1: 1r1. scats ill latl-ines ha\,e I)eeil descri1)ed in raccoon r~~zrticajat a \wiety of locations throughout dogs (Xyctet-ecrtesp,u)c!lot~oicles), golden jackals their railec<. (CSF\\'S, 1998). \\i. helie\.cl that (C(rr1i.s nrrl-ells), tllloles (Clron (rl/~itllrs),and foxes also urillatetl at the-sc latl-incs I,cc.ausc mancd \vol\-cs (Cl~t-!~soc!~onbt-crc1.1 y trnrs; \lac- \17c often fount1 accuni~~lationsof'\\rhitc salts don~tltl1985). Ethiopian \valves (Cmis sirnensis) from tlricd urine and \\.c, somctirnc~sti~untl tend to concentl-atct scats at latrines (Sillero- dump soil or sinall putldles smelling of urinc Zul)ii-i 1998) hund gra! \\~ol\!es (Cnnis 11rpra) near accumulations of' scats in the earl\, morn- sometimes tlef'cate near trail junctions (Pctc%rs ing. \\i. also founcl coyotc scats at some of ant1 \lecll 1975), which can lead to itccumnla- these latrines. C;oyotc scats are geilcrall!. mr~ch tions of scats that could Ilc consitlcrctl difi~sc larger than kit fibs scats and also lack thc stroiig, latrines (Vila et a]. 1994). Some fi).ues, such as iniisk~odor chart~ctcristicof kit fos scats (H. island tows (llroc!lolz littot-crlis;Laugllrin 1977) Cypher l~ersoii:il communication). ant1 gra!, foxes (I! ~itt~l-c.o(rrgetlt(!11~~:Trapp 1978) 1 iere, \ve l~rovirledt,tails on latrines disco\.- rlsc latrines. IIo\vcvcl; reaular nsc of latrines crctl while using trained clt,tc~ctiondogs to has not been tlescribcd for foscs in the genus locate salts of kit foses in the Lokern Satural \irll)es. although the I-ed fox (B ~.rrl~)rs)clcposits Area war Bakcrsfielcl, California. It has I)een itllitI glitlld secrctio~ls011 S~I~Cof its scats demonstrated I~cforethat docs<. trainctl to locate (\\:hitc ct al. 1989) ill i~tltlitionto frequent scats of targclt species can provitle a s~iccessful miirking with UI-inca(\lacclonaltl 1979). mcthocl of scat rcco\.c.r!. (U. Brt~itel~iiios~rant1 C. Bi-eitenmoser-\1~11rstcn,unpul~lislied data, 1 l~ottle,1 sheep carcass, 1 coyote skull, and 2 1984-1994; I? Pacluet, unpu1)lishecrl data, 1982- pieces of' bone). Ten of the 61 contained coy- 1989; Smith et al. 2001, 2003; \Vasser et al. ote scats in addition to fox scats. We also fi)uncl 2004). The principal habitat types in Lokenl 2 latrines containing only coyote scats. are nonnative annual grasslancl, saltl~ushscru11, \Ve fi~undkit fox scats at 98 locations in and upper Sonoran sul~sl~rul)scrub. The climate 2003. Forty-nille locations hael only 1 scat, 17 is semiarid with hot, dry summers and cool, had 2, and 32 had 3 or more. Eight of the 32 wet winters. Approximate summer high and latrines marked a conspicuous ol~ject(1 metal winter low tempel-atures are 37OC and 1°C, ol~ject,1 tire, 1 can, 1 fencepost, and 4 power- respectively Average yearly precipitation is line poles). Four of the latrines also contained 14.5 cnl, occurring priinarily as winter rains. coyote scats. Three of these contained 23 fox We established a transect system on an area and 3 coyote scats, 6 fox and 6 coyote scats, approximately 7 kin2. Transects ran fi-om north and 20 fox and 3 coyote scats, respectively. We to south and were spaced at 400-111 intei-vals. also found 1 latrine containing only coyote The entire transect system covered 56 kill. To scats. Although only about one-third (32193) of locate scats, a detection doghandler team sys- the locations where scats were fi)und were tematically searched along each transect (see latrines, latrines contained over 75% (2421315) Smith et al. 2003). We searched the entire tran- of the 315 fox scats we found. The largest sect system in Ja11u;ii-y 2002 and again in Janu- nuinl~erof kit fox scats we found at the 26 ary 2003, completing the searches within 5 latrines where we counted the num1)er of scats and 4 days, respectively. \i7e recorded the per- present was 30 (Fig. 1). pendicular distance in meters from the tran- \Ve measured distance fi-om the transect sect line to each location where a dog found 1 line for all 11ut 2 of the locations where scats or more scats. were found in 2002 (159) and all locations in A latrine is a place where 1 or more indi- 2003 (98). As distance from the transect was viduals defecate repeatedly, 11ut there are no independent of year (2-tailed t test, t = 1.510, standards for how many scats it takes to con- df =255, P = 0.132), we pooled data for 110th stitute a latrine. For example, Goi~nanand Mills years, yielding a sample of 257 locations. Scats (1984) reported that 1)rown hyena latrines con- were found at a minimum distance of 0 n1 tained 5-50 feces, while Begg et al. (2003) from the transect and at a maximum of 38.40 decided that 2 feces were enough to constitute in. The mean distance was 4.8 k 6.7 111 (s). a latrine in honey 11adgers (Mellivorn cnpen- Accumulations of multiple scats might have sis). Although 2 fox scats near each other might a stronger smell than a single scat, making it inark the beginning of a new latrine site, we easier fbr dogs to detect latrines than single believe that 2 scats might also be deposited scats. \F7e reasoned that, if this hypothesis were near each other simply 11y chance or mark a true, latrines would, on average, be found at site where 1 fi)x had encountered a scat of greater distances from the transect line than another fox ancl deposited another in response single scats. The mean distance for single scats even though this site was not hal~ituallyvisited. was 4.3 f 6.3 in and that for latrines was 5.9 + Because of these considerations, we defined a 'Ir .3, but there was no significant difference in latrine as an accumulation of at least 3 kit fox the distances recorded for single scats and scats. We recorded whether fox scats were latrines (%tailed t test, t = 1.801, df = 238, P found alone 01- in a latrine and whether they = 0.07). were located on or near a conspicuous object. In ur11an Bakersfield, where coyotes are rare In 2003 we also recorcled how 111any fox and (B. Cypher personal communication), we 011- coyote scats were present at most latrines used seived that kit foxes use lati-ines even when no 11y 1~0thspecies. coyotes are present. Therefore, use of latrines \.tie found kit fox scats at 161 locations in appears to be a typical feature of kit fox l~iol- 2002. Eighty-nine locations had only 1 scat, 11 ogy. Although the functions of these kit fox had 2, and 61 had 3 or more, thus nleeting our latrines are not known, they likely play some definition of a latrine. Latrines were rarely role in chemical communication. I11 several located near fox [lens: only 2 of the 61 latrines carnivore species where latrine use has been were near dens. Seven ofthe 61 latrines marked well studiecl, such as 11adgei-s (Stewart et al. a conspicuous object (1 cement object, 1 tire, 2001, 2002) and hyenas (Gorman and Mills [Volume 64
# of scats found at each location
Fig. 1. Kumher of locations at which various numbers of kit fox scats were found in the Lokern Natural Area in 2003.
1984, Mills and Gorman 1987), latrines serve described with piles of feces left by coyotes on to mark the territories of the resident social a footpath to mark the entrances of descending group. trails through thick brush in coastal California. Coyotes are known to urinate over places where wolves have urinated (Paquet 1991). If We thank the Christensen Fund, the Alter- they tend to urinate and defecate on kit fox natives Research and Development Foundation, scats. this would lead to the accumulation of the Smithsonian Institution Scholarly Studies coyote scats at some kit fox latrines. However, Program, the Friends of the National Zoo, the we found 3 latrines that contained only coyote National Center for Environmental Research, scats, indicating that coyotes use latrines on the EPA STAR program, and the Abbott Fund our study area even if no kit fox scats are pre- for financial support. Alice Whitelaw, Aimee sent. Thus, it is not clear whether joint latrines Hurt, and Mike Smith provided exceptional result from coyotes responding to kit fox scats, field assistance. Brian Cypher and James Mur- kit foxes responding to coyote scats, or both. doch made helpful comments on the draft If we had trained dogs to locate covote scats manuscript. rather than kit fox stays and had searched a larger geographic area, we likely would have found more latrines that contained only coyote scats. Although<, several detailed studies of scent ALLEN,J.J., M. BEKOFF,AND R.L. CRABTREE.1999. An ob- marking in coyotes do not mention latrine use servational study of coyote (Canis latrans) scent- (Bowen and Cowan 1980, Wells and Bekoff marking and territoriality in Yellowstone National Park. Ethology 105:'289302. 1981, Bekoff and Wells 1986, Gese and Ruff BEEOFF, M., AZID M.C. WELLS.1986. Social ecology and 1997. Allen et al. 1999). Camenzing (1978) men- behavior of coyotes. Advances in the Study of tions2 coyote latrines'discovered ii~yorning. Behavior 16:2Fjl-238. One was in an empty hay shed and contained BEGT., C.M., J. BEGC,,T. DU TOIT, AND M.G.L. MILLS. over 500 scats. The second, under an old 2003. Scent-marking behaviour of the honey badger, Mellitiorrr capensis (Mustelidae), in the southern wooden bridge crossing a dry creek, also con- Kalahari. Animal Behaviour 66:917-929. tained several hundred scats. Both latrines were BOWEN,W.D., AND I.M. Co~v.4~.1980. Scent marking in in inconspicuous, covered areas and Camen- coyotes. Canadian Journal of Zoology 58:473480. zind (1978) did not think that they "served ter- BRA~TSTROSI,B.H. 1999. Trail marking by coyotes, Canis ritorial functions." However. it seems likelv that latrans. Southwestern Naturalist 44:405406. CASIEKZIND,EJ. 1978. Behavioral ecology of coyotes on covotes do use accumulations of scats for the National Elk Refnge, Jackson, Wyoming. Pages chemical communication under some circum- 267-294 in M. Bekoff, editor, Coyotes: biology, be- stances, as in the example Brattstrom (1999) havior, and management. Academic Press, New York. lic:osc:( 1:. t1.J. l!)(i2. Ecu)log!. ;illtl life I~ihto~?ol'tlrc. kit fin S\IITII,D.. K. I{\I.I.S,13. I),\\.I:YIJoI~T.13. AI)\\IS, \\I) 1. ill 'li)oc.lc (:aunt!. Ltah. I'coloq 4:3:481-4!)7. \I.\I,I)C)\.\I)O.2001. (;.III~II~;ishistii~~ts fhl- co))x.l-\~i- GI