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And Ainpiificatior7 of Genes from Fecal Material Were Aimed at Devel n P984, Putman~ remarked in and ainpiificatior7 of genes from his review FM:& ikxn Faeres fecal material were aimed at devel- that ‘for any frustrated field opirng slew tools for clinicai diagno- mammalogist seeking observa- sisz7. Similar techniques allowed tional da’s on their elusive study parasitofogicai studies of domestic animals, dung may represent the and wild populations; for example, most readily-avaiiabie and easily- tbe amplification of DNA from tape- collected source of information worm (.Ecl~inococcm muMocul0ris) upon which they may fall back in in feces of red foxes (Vdpes udprs) despair’. Despair is frequent these showed this parasite had a high days because many mammals puevalence~x. are becoming increasingly elusive Feces contain cells shd from as their numbers decline. Rarity the intestinal lining; thus DNA fForn makes direct observation difficult the bust itself could potentially but the need for genetic, derno- be is&ted ad aanalyzea.l.The ap- graphic and life-history infor- p8iciltiOlI of this idea to an anilnal mation for species Sacimgextinc4ioul ~bopulatiore involved a saaalil popu- is more pressing than ever. Ad- lation of elusive, higbiy eoadan- ditionally, the great expense arPd gc~ctl brown bcnrs (Unua c~n’los) i~i extensive number of permits re- the Brenta mountains of northern quired to capture and transport 11alyx!‘-.“. The population size and samples makes studies of endan- Michael Kohn and Robert Wayne are at the sex composition were unknown, but gered species unpalatable to many Oept ot Glogy, University of California, anecdotal observations suggested 621 Circle Drive South. Los Angeles, biologists2-6. that only a few bears remained. A CR 90095-1606, tJSA. Even in abundant species, large- single mitochondrial DNA controi- scale survevs cf DorPufations are region sequence was found in bear logistically &ffic& ii they depend feces collected from the Brerlta on opportunistic sampting (e.g. Ref. 6). Moreover, “any study mountainassuggesting that ;he popu%ation was small and of a natural population has the unavoidable side effect of isolated. This sequence was present also at high frequencv altering the dynamics of the population in ways that may be in bear populations from Slovenia, Croatia and Bosnia, indk- difficult to predict” or that may change some characteris- eating that the Rrenta bears formerly were in contact with tics that a researcher is attempting to measure’.x. these populations. A 8(-cbromosomal nuclear marker SRY Must we lose hope if observation and biological sampling was used to show that at least one male and one female 1~11 are not possible? Perhaps blot: in this article we discuss a&- WWe [H?Wtl~ in r!::! alYil. TBWSe re’.4ts also implit ihal cations of recently introduced genetic techniques for the 0thcr WlCl?Z ~llilt%tTS SUCh AS InicroSatellil~s ElKQ tH> ClSOd analysis of feces (molecuPar scatology) when studying Iree- ----I._~_ - ranging mammal populations. These techniques allow for r the purification of DNA from feces, which is then followed by amplification of specific DNA sequences using the polymer- Mitochondrioi DNA’“(mtDNA) is a matemalty inhented. haplord.nun-recombmmg ase chain reaction (PCR). An array of genetic markers with and extra-nuclear genome. Each cell has multiple mtDNA copses crangc IC-250G different properties is available9-‘a and call be used to ad- copies) and thus mtDNA sequences are more eastly amplified than smgle copy. dress taxonomic questions, individual f(ler~tificat~on through nuclear genes. Because the mtDNA cytoclx~ne 0 (~~40) gene has only moderate mtraspeciflc variation. it is well suited for use In species identlficatlon. The mtDNA ‘genetic fingerprinting’, and sex determination with gender control Pegion IS commonly vanable on the rntraspecific bevel and is sultable for specific markers (Box I ), Moreover, specific sequences from studies oi genetic variability. phylogeography. ass@?ment to management units. other passengers through the gut such as food and patho- and forens,?,. Chloroplast DNA” (cpDNA) and mtDNA ran be used to amplify plant gens can be determined by using CR primers specific fcx and animal sequences from feces to study diet. Moreover, a vast number of their amplification. Consequently, molecular techniques ap- pathogen sequences (“(ruses. bactena. protists and macroparasItes) are avallable for ampltflcation from fecal material. plied to feces allow the identification of animal species, indi- Mlcrosatellites~.‘~ [also referred to as simple sequence length polymorphism viduals, their sex, pathogens and food habits. Furthermore, (SSLP). or slmpfe sequence repeats Jare nuclear. single-copy DNA conslstlng of tan- comprehensive sampling of an area for feces, or systematic demly repeated short sequence mohfs such as (CA),. They are scattered through fecal transects of populations. may allow estimates of home out :he genomo and are highly polymorphic. 3~ assaying several m!crosatellrte 100. 3 multllocus genotyDc:can he obtaIned: a .genetlc fingerprint’, which IS untque to Indk range, reproductive patterns, kinship structure and popu v~dual animals These markers are cudon~~na~tl~II?W:IIC.~ :alleks !rom both par l&ion size. Thk information may be augmented by tech- ents are traceable in the offsprl:d) and are useful for studying paternity and [tinship. niques for analysis of feces that do not use DNA1~‘5-25(Rox2). genetic varlatton. population genetrc structure and gene flow9.11. M~crosatellites Analysis of feces will potentially provide a means to study Identified in one species can usually be employed IO study a related species! ‘. PCR pnmers for the SHVgene on the Y chromosome can be used t0 dlstin genetics, life history and population dynamics of mammals. gu,ulsll male from female DNA”. Genetic maiicer sequences and WI? pnmers are published or accesslb:e throu$ GenBanlc or EMBL, which are electronic databases avallablr: on tile lW?rnet tha! PCR can produce large quantities of specific DNA se- can be searched for sequences by entering the taxon. genomlc lOCUS or SequenCC quences from small, degraded and impure samples”). Stimu- of Interest. lated by this property of RX, early attempts of isolation fecal studies showed that the source for reintroductlcn should be bears taken from the Slovenian p ation. Mitocbondrial DNA has been amplified from feces Mammal specres can often be tdentified based on their feces, allowtng stuu’ies of spec~cs iompos~t~on~. However, without addrtional information feces ara assigned of the Indian elephant (Elephas maximw), European bison correctiy :n only about 59-6606 cf the case2 ?j5, This uncertatnty IS owing to con- (Bison Bonasrrs), polar bear (Ursus nlak+itimus)3ci~3’and dugong sideraM? size-overlap of feces rn related tdxa In some cases, the pH value and (Dugong dugo.cJng)“z.Four studies Rave now confirmed that b~lc acids of feces have be‘sn use;~! to rdentl& specie\!‘. Relative abundance and individuals could be ~f~~ge~~~i~ted’ from their feces; these populatton SIZE have been estimated from fecal counts1 lE Hocever. the resdmultng studies successfully amplified microsatellites from feces of of Illdivrduals constrtutes a problem. Census data baser1 on teces can be supple menteci by techniques described elsewhere1,16,17. wild baboons (.Pnpio cynoce@~lus)~~~, bonobos (PM panis- Habitat use and range size may be investigated by examining spatial variation cus)34,brown bears (Ursus arclos)3” and pinnipeds?s. Although in feces abundance. Simrlarly, territory boundaries r- ‘y be marked by fecal aggre- some technical problems were encountered (Box 3), these gations:. These studies often require individuals to /I gest colored plastic markers studies F:esent convincing evidence that such problems or radioactive halt, which uniquely mark their feces. Analvsis of food habits and dietary quality have frequently used fece9.l;. In can be overcomexr-36 (see also Refs Z&37,38), and feces may carnIvore& undigested ploy Items can be recovered and Identified. Herbivore dung some day be as useful as blood or tissue samples for genetic often contains plant remams. In general, this approach lhas shortcomingsl,‘r,‘i,ls. analysis. In part, our intent in this article is to provide incen- tive for additional t&mica0 research. Previously, a variety of other potential sources for DIVA such as hair, bones, feathers, saliva, skin and nails have been collected for Pson-invasive genetic analy&P’. However, these SOtiCC(5 may be more difficult to obtain than fecal samples ant4 may provide less intormation. The g.enetic an fife history iaIformatiom that potentially ran be extracted from fecal samples is sunmiarizcd in Fig. 1, on feces to identify individuai beam?. Finally, a glimpse of Two essential parameters that need to be assessed in the feeding habits of Brenta bears during late summer be~~~~v~o~a~studies are the number of offspring produced by (when feces were collected) was provided by e relationship of indivi plant chloroplast sequences of the &CL gene group to each other. This information allows behaviors feces. Fecal plant sequences were identified as belonging to lead to reproductive success to be identified and prov the hawthorn genusHzotini;13, thought to be important in the an assessment of the kinship, component of cooperative be- diet of brown bears. Although animals were never seen dur- haviors (e.g. Ref. 11). M ing the study, feces provided answers to establish paternity and usually require direct observation and the handling of ani- popu&ionsX,“.l l.:i’).~llland mals. The results also had conservation
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