Habitat Selection by the Siberian Musk Deer Moschus Moschiferus

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Anthropogenic influences on the distribution of a Vulnerable coniferous forest specialist: habitat selection by the Siberian musk deer Moschus moschiferus

J ONATHAN C. SLAGHT,BRIAN M ILAKOVSKY,DARIYA A. MAKSIMOVA I VAN V. SERYODKIN,VITALIY A. ZAITSEV A LEXANDER M. PANICHEV and D ALE G. MIQUELLE

Abstract The Siberian musk deer Moschus moschiferus, ca- Introduction tegorized as Vulnerable on the IUCN Red List, is a small ungulate associated with coniferous forests of East Asia. In ogging is an important component of the economy of Russia the species is hunted both legally and illegally for Lthe southern Russian Far East. If conducted properly, the commercially valuable musk gland in males. Steep some logging can improve habitat for ungulate species by population declines recorded in recent decades have been creating a structurally diverse forest, promoting growth of generally attributed to intensive illegal hunting, but the de- understorey vegetation and providing forage (Slaght et al., cline has coincided with increased logging activity and the in press). However, the logging roads built to facilitate timber concomitant expansion of logging roads. We conducted removal in the region also provide access for poachers, who an occupancy analysis in Primorskii Krai, Russia, to eluci- target ungulates, such as Siberian musk deer Moschus date the relative importance of environmental, ecological moschiferus,sikadeerCervus nippon, roe deer Capreolus py- and anthropogenic features associated with the presence gargus, red deer Cervus elaphus and wild boar Sus scrofa,as of musk deer. The top model contained covariates related well as Asiatic black bears Ursus thibetanus and Amur tigers  to the abundance of bearded lichen Usnea spp., the distance Panthera tigris altaica (Dunishenko et al., ; Slaght &  to a main road and the distance to logging sites, suggesting Surmach, ; Slaght et al., in press). Although most ungu- that both intensive hunting of musk deer (associated with lates are poached for their meat, musk deer, categorized as  greater accessibility via roads) and logging of habitat are in- Vulnerable on the IUCN Red List (Nyambayar et al., ), fluencing the occurrence of this species. We propose several are usually targeted for the musk glands found in males. management actions to limit the negative influence of log- Musk is used in eastern traditional medicine and the per-   ging and logging roads on musk deer in Russia, including fume industry (Ostrowski et al., ). Since the s encouraging logging companies to set aside high conserva- Moschus spp. numbers in Russia and elsewhere have de- tion value forests (to retain intact forests) and to close log- clined sharply, and the decline is generally attributed to in- ging roads post-harvest (to reduce access by poachers). tensive harvesting (both legally and illegally) of the glands (Homes, ; Nyambayar et al., ; Ostrowski et al., ). Keywords Habitat selection, logging, Moschus moschiferus, Musk deer are found throughout much of the higher ele- occupancy, poaching, roads, Russia, Siberian musk deer vation forests of East Asia. Unlike most ungulates in the Supplementary material for this article can be found at southern Russian Far East, which are found in deciduous https://doi.org/./S or mixed forest, musk deer are closely associated with mature and over-mature coniferous forests (Zaitsev, ). The deer prefer such forests because of the abundance of pendulant epiphytic lichens (Usnea, Evernia, Bryoria and Ramalina spp., hereafter forage lichens), which can com-  ’ JONATHAN C. SLAGHT (Corresponding author) and DALE G. MIQUELLE* Wildlife prise up to % of the musk deer s winter diet, and because Conservation Society, 2300 Southern Boulevard, Bronx, NY 10460, USA the patchy structure of these forests (with dense patches of E-mail [email protected] undergrowth and abundant felled trees) provides excellent BRIAN MILAKOVSKY WWF, Amur Branch, Vladivostok, Russian Federation cover for these secretive ungulates (Zaitsev, , ; DARIYA A. MAKSIMOVA,IVAN V. SERYODKIN* and ALEXANDER M. PANICHEV Pacific Prikhodko, ; Domanov, ). Geographical Institute, Russian Academy of Sciences Far Eastern Branch, ’ Vladivostok, Russian Federation The musk deer s coniferous habitat is currently a priority target for logging in the southern Russian Far East. VITALIY A. ZAITSEV A.N. Severtsov Institute of Ecology and Evolution, Moscow, Russian Federation Although the region still contains large expanses of unman- *Also at: Far Eastern Federal University, Vladivostok, Russian Federation aged, roadless coniferous forests, many of which meet the Received  April . Revision requested  July . definition of intact forest landscapes (Aksenov et al., Accepted  November . First published online  May . ), industrial timber harvesting is continually expanding

Oryx, 2019, 53(1), 174–180 © 2017 Fauna & Flora International doi:10.1017/S0030605316001617 Downloaded from https://www.cambridge.org/core. IP address: 170.106.35.76, on 27 Sep 2021 at 15:42:13, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605316001617 Habitat selection by Siberian musk deer 175

into previously remote areas (e.g. in Ternei County of nor- Primorskii Krai, Russia, in and adjacent to the Sikhote- thern Primorskii Krai (or Province) the cumulative length of Alin Biosphere Reserve (Fig. ). The Biosphere Reserve is forest roads increased an estimated -fold during – an IUCN Category Ia federally protected area, where re- , from  to , km; Slaght et al., in press). source extraction (e.g. hunting, logging, collection of non- The role of logging and its subsequent impact on human timber forest products) is strictly prohibited and other access and forest structure is a poorly understood factor in- human use (e.g. recreation) is largely restricted (although fluencing musk deer populations. Industrial forest manage- all these activities occur in adjacent lands). The study area ment may affect musk deer populations directly (e.g. is mountainous, at –, m altitude, with forests domi- through habitat transformation) or indirectly (e.g. through nated by Ayan spruce Picea ajanensis, Manchurian fir Abies increased hunting pressure as a result of expansion of the nephrolepis and Korean pine Pinus koraiensis, and with a logging roads network), or both. Although there is a general significant component of Dahurian larch Larix dahurica consensus that intensive snaring (the most effective means and yellow birch Betula costata. Korean spruce Picea kor- of obtaining musk deer glands) is the primary culprit, aiensis sometimes replaces Ayan spruce in stream and Tukhbatulin () argued for the importance of indirect river valleys. At lower elevations temperate hardwood spe- factors, having found that the musk deer population in cies begin to appear, including Amur linden Tilia amuren- the Samarga River basin (northern Primorskii Krai) in- sis, small-leaf maple Acer mono, Manchurian ash Fraxinus creased significantly in the first year after selective harvest- mandshurica, mountain elm Ulmus laciniata and ing in spruce–fir forests (probably because of the abundance Mongolian oak Quercus mongolica. of accessible forage lichens in the logging slash) but then The dominant forest leaseholder in the study area (in the dropped catastrophically over the next  years as hunting lands outside Sikhote-Alin Biosphere Reserve) is Joint Stock and poaching pressure increased. Extensive anecdotal evi- Company TerneyLes (hereafter TerneyLes), which cuts tim- dence from the region confirms the key importance of ac- ber primarily using shelterwood harvesting, a management cess: many hunters, game managers and rangers have technique that removes enough canopy trees to allow light stated that musk deer populations plummet in newly roaded into the understorey to stimulate regeneration but leaves en- forests, irrespective of logging intensity. ough to provide some protective shade (hence the term shel- However, much literature also points to a significant dir- ter). This method leaves a high proportion of standing trees ect impact of timber harvesting on musk deer; for example, post-harvest, including many that will never be harvested, in the late Soviet period (when hunting and poaching of the such as the Korean pine, logging of which is banned species for its glands was relatively limited), sharp reductions (CITES, ) because of its high ecological and social in musk deer populations were observed after clear-cutting of value as a source of pine nuts. Many large trees are also mature coniferous forests in Chuguyevka County (central left standing when, because of the mountainous relief, Primorskii Krai; Ribachuk, ). Similarly, Domanov skid trails veer apart and the space between them is unreach- () observed that musk deer nearly disappeared after log- able by harvesters from either side. This increases the level of ging in relict patches of mature spruce–larch forests in the retention and often leads to unharvested islands of pine and Tukurungra Mountain range in Amurskaya Province, surrounding spruce and fir trees amidst logging sites. where commercial hunting of musk deer was almost absent. The loss of cover and sources of forage lichens after logging and forest fires was identified as an important driver of musk Methods deer population declines in both the Sikhote-Alin Mountains of Primorskii Krai and on Sakhalin Island, where the local Survey design subspecies of musk deer is listed as Endangered in the Red Book of the Russian Federation (Zaitsev, ; Ministry of We conducted surveys in or near the northern border of Natural Resources, ). Our objective was to assess the in- Sikhote-Alin Biosphere Reserve, with surveys focused on fluence of logging activities (including road management) on three sites: Nechet, Taezhnoe and Venera regions (Fig. ). the relative abundance of musk deer and to develop guide- By including transects both inside (with no logging and lines for reducing anthropogenic impacts on the species. few roads) and outside the protected area (with logging We believed that musk deer would prefer areas with fewer and many roads) we were able to capture the variety of con- roads, less logging, and greater availability of lichens. ditions available for musk deer in the study area. We placed a network of transects at regular intervals (c.  km apart), starting from access points (roads) within our grid of prede- Study area termined management regimes (e.g. protected and selectively harvested sites). Surveyors walked along transects, guided The study was conducted in the central Sikhote-Alin by global positioning systems, and stopped at data points Mountains of Ternei and Krasnoarmeisk Counties, every  m to record habitat parameters, including harvest

FIG. 1 Probability of musk deer Moschus moschiferus occupancy within the (a) Venera and (b) Taezhnoe–Nechet regions of the study area, in and adjacent to the Sikhote-Alin Biosphere Reserve, Primorskii Krai, Russia.

regime, snow depth, tree stand composition, canopy cover, Any cell that contained four or more data points (or spatial topography, aspect, elevation, distance to roads, distance to replicates) was included in the analysis. We selected this harvest sites, and abundance/availability of lichens number of spatial replicates based on recommendations in (Supplementary Table S). Fresh tracks (i.e. ,  hours MacKenzie & Royle () for the number of data points old) of musk deer, other ungulates, predators or people needed per survey unit for an accurate estimate of the prob- were recorded whenever they bisected the transect, and ability of occupancy, by assuming a true occupancy value of the same environmental variables were collected. . (i.e. % site occupancy) and a detection probability of Lichen data were assessed in two ways: lichen load and . (i.e. an individual musk deer had a % chance of being lichen availability. To estimate lichen load, trees in a given detected). These assumptions were conservative given that  plot (determined as being in the plot using a relascope with a an adult male has a winter home range of c.  km (twice basal area factor of .) were assessed visually for musk deer the size of our survey unit; Zaitsev, ) and we conducted forage lichens as being –, – or . % lichen covered. our fieldwork in winter (when musk deer were easily detect- Although much of the lichen load is unavailable to musk able from tracks in snow). deer at any given time (being high up on the tree), some por- To model occupancy we developed a suite of a priori can- tion of it becomes available when branches and attached li- didate models (based on results of single parameter tests) to chens are brought down by wind. The availability of lichens determine which covariates had the greatest impact on # . m from the ground (the approximate height a musk musk deer occupancy (Supplementary Table S), and then deer can reach) was estimated on a scale of – (low to estimated the maximum likelihood of each model being the high availability). best fit for the data. We used  competing models with nine covariates to provide a quantitative assessment of the probability of musk deer occupancy of a given area. Data were Data analysis analysed following the framework for a single-species, single-season occupancy model in MacKenzie et al. We used a presence/absence approach (MacKenzie & Royle, (). We determined model fit using Akaike’s informa- ) to elucidate habitat availability vs use for musk deer, tion criterion (AIC; Burnham & Anderson, ), and iden- and to assess the relative importance of environmental, tified the best model by the lowest AIC score. However, any ecological and anthropogenic features associated with occu- models within two AIC units are considered to be equally pancy by musk deer. Occupancy modelling uses maximum viable, and any model within seven AIC units is considered likelihood to incorporate detectability of a target species into to be competitive. the estimation of site occupancy (Royle & Nichols, ). To describe the predictive accuracy of the top model we  Covariates for the occupancy analysis were selected based estimated r following methods similar to those of Skalski on an earlier pilot study (Slaght et al., ) after removing et al. (). Specifically, we assessed the total variation in  any correlated covariates (where r $ .). We created a the data by comparing the global and null models, and  grid of  km survey units and laid it over the study area. then evaluated the proportion of that variation explained

by the top-ranked model. We defined the global model as a of musk deer across the landscape, both inside and outside model using all of the covariates in the occupancy param- the protected area. eter. We analysed the data in PRESENCE v. . (Patuxent Similarly our analysis suggests logging has a negative in- Wildlife Research Center, Laurel, USA), standardizing cov- fluence on the probability of musk deer occurrence at a site. ariates using z-scores prior to analysis, and displayed the re- Distance to harvest site was the only single-covariate model sults using ArcGIS v. . (ESRI, Redlands, USA). considered to be competitive (models with ΔAIC , ), and all six of the top models contained this covariate, suggesting that it may be an even more important predictor of musk Results deer occurrence than road access. There appears to be an additive effect of roads and harvest, given that separately During  December – March  we walked c.  km the covariate distance to main road scored low of transects ( km in Nechet,  km in Taezhnoe and  km (ΔAIC = .) but when coupled with distance to harvest in Venera) and collected habitat parameters from , it appeared in four of the top six models. points. All surveys were conducted in potential musk deer It is possible that the negative effect of timber harvesting habitat in coniferous forest (mean forest cover was %), on musk deer use is, like the effect of distance to roads, con- at elevations of –, m (mean =  m), in unlogged nected primarily to access. The logging roads (and even skid areas as well as areas that had experienced logging. We re- trails) that fan through the forest from the main roads facili- corded wildlife sign on  occasions, including  records tate access by off-road vehicles as well as on foot. Setting of musk deer tracks, by far the most common ungulate in snares for musk deer around the perimeter of freshly logged the study area. sites is a common practice, as the ungulates are attracted to The results of model selection are in Table . The top the accessible lichens in the logging slash and may even be model suggested that musk deer selected sites with high li- drawn to the sound of falling trees (Zaitsev, ; chen load (β = . ± SE .), larger distance to main road Tukhbatulin, ). (β = . ± SE .) and larger distance to harvest site The importance of lichen loading in all three top models (β = . ± SE .). There were two other models with that could be considered equally competitive (ΔAIC , )in ΔAIC , , and six models in total with ΔAIC , . After predicting use of a site by musk deer suggests that logging is eliminating top models with uninformative parameters also important as an instrument of habitat transformation. (those with confidence intervals that overlapped ), we The fact that high lichen load alone was uninformative were left with various iterations of models with the same (ΔAIC = .) seems to contradict this conclusion but three covariates. could be explained by the fact that mature coniferous forests We found that our top model explained % of the vari-  with relatively high lichen loads are common in the unman- ation in the data (r = .), suggesting very strong fit. aged portions of the landscape. In such areas the supply of However, we note that the global model used to estimate  lichens is likely to exceed the needs of musk deer, and so li- r was not a saturated model (i.e. one with occupancy beta  chens do not act as a limiting resource, but in managed for- coefficients for each site), and thus our r value is probably ests, where the supply of large, old trees with open crowns an overly optimistic assessment of the top model’s explana- (the best substrate for pendulant lichens) is greatly reduced, tory power. lichen load in combination with reduced human access drives musk deer occurrence. However, exactly how harvest Discussion is influencing lichen distribution and abundance remains unclear. Our analysis suggests that both environmental and an- Further research is necessary to clarify the influence of thropogenic parameters were important in delineating timber harvesting on lichen loads and musk deer occu- musk deer occurrence. Musk deer preferred areas with pancy. The type and intensity of logging vary across the high lichen loads that were further from the main road landscape, from low-impact selection of individual, large and further from areas that had been logged. The inclusion spruce trees to commercial clearcuts. We observed some of distance to main road as a covariate in four of the top six partially harvested logging sites where the lichen load re- models provides evidence that direct human harvest (e.g. mained relatively high, and in some cases extremely long hunting and poaching) is influencing musk deer distribu- ‘beards’ of pendulant lichens were most common on the tion. In other words, the more inaccessible an area, the sides of old trees that faced open logging trails. Harvest ex- greater the likelihood that musk deer will be present. periments in Finland have suggested the possibility that se- Clearly, it is easier and more efficient for hunters and poa- lection harvests may actually improve conditions for chers to establish snare lines for musk deer that are easily pendulant lichen growth (Storaunet et al., , ), accessible from a road network. Hence, our results provide where – mature spruce trees with Usnea longissima evidence that human harvest is influencing the distribution per ha were retained post-harvest, and the volume of the

TABLE 1 Models of musk deer Moschus moschiferus occupancy in and adjacent to the Sikhote-Alin Biosphere Reserve, Primorskii Krai, Russia (Fig. ), with ΔAIC (the difference between the model with the lowest AIC value and the top model), Akaike weight (the probability that a model is the best fit of all candidate models), λ (the likelihood that a given model is the top model), and K (the number of parameters used in each model). Competitive models (i.e. those within  ΔAIC units of the top model) are shaded grey.

Model ΔAIC Akaike weight λ K Distance to harvest site + distance to main road + high lichen load 0.0 0.43 1.00 5 Distance to harvest site + distance to main road + high lichen load + % dark conifers 0.9 0.28 0.64 6 High lichen load + distance to harvest site 1.9 0.17 0.38 4 Distance to harvest site + distance to main road 3.6 0.07 0.17 4 Distance to harvest site + distance to main road + lichen availability 5.5 0.03 0.06 5 Distance to harvest site 6.8 0.01 0.03 3 High lichen load + distance to main road 9.6 0.00 0.01 4 Distance to main road 10.1 0.00 0.01 3 Distance to harvest site + distance to main road 11.0 0.00 0.00 4 % Dark conifers 18.0 0.00 0.00 3 % Dark conifers + high lichen load 18.1 0.00 0.00 4 High lichen load 18.5 0.00 0.00 3 Forest harvested or not 20.5 0.00 0.00 3 Topographic position index + elevation + aspect + % dark conifers + high lichen load 22.3 0.00 0.00 7

lichen increased (probably as a result of heightened light our results. We believe that both of these factors contribute and moisture levels compared to the dense canopy of the to reducing the quality of managed spruce–fir forests with unlogged control). roads in comparison to inaccessible, intact forests. As such, a strategy for conserving populations of this Vulnerable, commercially valuable species should include Management recommendations protection of prime habitats from both road building and timber harvesting and, where harvesting occurs, eliminating A simple comparison of logged vs unlogged forests does not access along forest roads after harvesting is discontinued. tell the full story of how timber harvest is affecting musk Protected patches should be large enough to support a deer. Our data appear to show there are at least three factors musk deer social unit (–, ha; Zaitsev, ), al- driving musk deer distribution in a selectively harvested though for long-term conservation of genetic diversity lar- landscape: distance from roads, distance from harvest ger habitat blocks (,–, ha) may be necessary sites, and amount of lichen. In Canada, a strategy has (Zaitsev et al., ). been developed for conservation of the caribou Rangifer tar- The protected area network in the central Sikhote-Alin andus, another Vulnerable ungulate associated with lichens Mountains is already well developed, and the prospect of es- and old-growth forests, which includes both exclusion of tablishing a large number of additional protected areas is habitats from logging and the use of practices on logging unlikely. Consequently, other mechanisms for securing sites that maintain lichen-rich trees (Armleder & habitat should be considered. One possibility would be to Stevenson, ; Miège et al., ). Musk deer clearly re- set aside high conservation value forests from logging, in spond to high lichen loads and, given our observations the framework of Forest Stewardship Council voluntary for- that forage lichens thrive in some partially harvested stands est certification; for example, TerneyLes has agreed to set in the study area, it is possible that similar guidelines could aside several massifs of intact Korean pine–spruce–fir for- be developed in Russia. However, our results did not iden- ests to the north of Sikhote-Alin Biosphere Reserve. This tify an interpretable mechanism by which logging influences practice should be extended to forests leased by other log- high lichen loads (besides the crude and obvious observa- ging companies in the region and, where possible, adjacent tion that high intensity harvests such as clearcutting reduce massifs should be set aside to increase the effective size of lichen loads to near zero). In the absence of clear, negative high conservation value forest massifs. The conservation impacts of harvest on lichen abundance it is difficult to of intact, mature and overmature coniferous forests would make specific management recommendations to reduce li- benefit the conservation of other key game species, such chen loss and improve habitat for musk deer without further as the sable Martes zibellina and Eurasian lynx Lynx lynx, research on the role of timber harvest on lichen load. as well as rare species such as the Near Threatened We were unable to tease apart the direct (habitat trans- Siberian spruce grouse Falcipennis falcipennis. formation) and indirect (increased human access) impacts Roads could be closed post-harvest by removing bridges of timber harvesting on musk deer populations, based on or using barricades such as earthen berms at key access

points to reduce or eliminate vehicular access (Slaght & ARMLEDER, H.M. & STEVENSON, S.K. () Using alternative Surmach, ). Such actions have had significant impacts silvicultural systems to integrate mountain caribou and timber management in British Columbia. Rangifer, Special Issue No. , in protecting wildlife species and restoring habitat elsewhere –   . (Forman et al., ; Switalski & Nelson, ; Kleinschroth BURNHAM, K.P. & ANDERSON, D.R. () Model Selection and et al., ). Road closures have already been undertaken on Multimodel Inference. Springer, New York, USA. a limited scale by TerneyLes to reduce human access to eco- CITES () The CITES Appendices. Http://www.cites.org/eng/app/ logically sensitive riparian zones, and could be extended to index.shtml [accessed  February ].  coniferous forests as well. If human access could be re- DOMANOV, T.A. ( ) Ecology of musk deer (Moschus moschiferus, Linnaeus ) on the Tukurungra Range. PhD thesis. Irkutskaya stricted via road management, it is not only musk deer po- Agricultural Academy, Irkutsk, Russia. [In Russian] pulations that would benefit: in the southern Russian Far DUNISHENKO, Y.M., ERMOLIN, A.B., DARENSKY, A.A., DOLININ, V.V., East, roadless areas are typically bypassed by people seeking SOLOVEY, A.A., GOLUB, A.M. & ZHUKOV, A.Y. () Hunting areas that are accessible by vehicle, and thus closures are a Resources of Khabarovsky Krai. Khabarovsk Provincial Typography, simple and inexpensive means to convert a forest from high Khabarovsk, Russia. FORMAN, R.T.T., SPERLING, D., BISSONETTE, J.A., CLEVENGER, A.P., to low human use (Slaght & Surmach, ). Road closures CUTSHALL, C.D., DALE, V.H. et al. () Road Ecology: Science and are likely to reduce poaching pressures on a wide variety of Solutions. Island Press, Washington, D.C., USA. species, including ungulates, tigers, bears and salmon HOMES, V. (ed.) () No License to Kill: The Population and Harvest Oncorhynchus sp., and benefit logging leaseholders by redu- of Musk Deer and Trade in Musk in the Russian Federation and cing the risk of anthropogenic fire and illegal timber extrac- Mongolia. TRAFFIC Europe, Cambridge, UK. KLEINSCHROTH, F., GOURLET-FLEURY, S., SIST, P., MORTIER,F.& tion (Slaght et al., in press). As such, closures facilitate HEALEY, J.R. () Legacy of logging roads in the Congo Basin: how resource extraction in a way that meets the needs of the persistent are the scars in forest cover? Ecosphere, , –. local economy and global timber demand, and also mini- MACKENZIE, D.I. & ROYLE, J.A. () Designing occupancy studies: mizes the negative impact of this activity on natural general advice and allocating survey effort. Journal of Applied communities. Ecology, , –. MACKENZIE, D.I., NICHOLS, J.D., ROYLE, J.A., POLLOCK, K.H., BAILEY, L.L. & HINES, J.E. () Occupancy Estimation and Modeling. Academic Press, New York, USA. Acknowledgements MIÈGE, D.J., ARMLEDER, H.M., WATERHOUSE, M.J. & GOWARD,T. () A Pilot Study of Silvicultural Systems for Northern We thank field assistants and volunteers G. Bannikov, Caribou Winter Range: Lichen Response. Working Paper /. A. Gabrielson, E. Tabalykin, M. Pozdeev, E. Petrunenko, Research Branch, British Columbia Ministry of Forests, Victoria, M. Borisov, A. Kosolapova and D. Maksimov. F. Maisels Canada. MINISTRY OF NATURAL RESOURCES OF THE RUSSIAN FEDERATION and T. Rayden assisted with a literature review. D. Tempel () Strategy for conservation of the Sakhalin musk deer in provided advice and support regarding occupancy Russia. Official order №-R,  March . Moscow, Russia. [In modelling. This project was made possible by grants Russian] awarded by the WWF–IKEA Partnership and the Wildlife NYAMBAYAR, B., MIX,H.&TSYTSULINA,K.() Moschus Conservation Society Research Fellowship Program (for moschiferus. The IUCN Red List of Threatened Species :e.       DAM). T A . Http://dx.doi.org/ . /IUCN.UK. - .RLTS. TA.en. OSTROWSKI, S., RAHMANI, H., ALI, J.M., ALI,R.&ZAHLER,P.() Musk deer Moschus cupreus persist in the eastern forests of Author contributions Afghanistan. Oryx, , –. PRIKHODKO, V.I. () Musk Deer: Origin, Systematics, Ecology, JCS provided primary study design, primary statistical ana- Behaviour and Communication. GEOS Books, Moscow, Russia. [In lysis, and played a lead role in article preparation. BM and Russian] RIBACHUK, V.N. () Influence of Logging on Winter Feed of the DGM contributed significantly to editing the article, se- Musk Deer. Moscow University Herald (Vestnik Mosckovskovo cured funding for the project, collected field data, and Universiteta). Geografiya, Moscow, Russia. [In Russian] played an advisory role in study design and data analysis. ROYLE, J.A. & NICHOLS, J.D. () Estimating abundance DAM, IVS, VAZ and AMP provided primary data collec- from repeated presence–absence data or point counts. Ecology, , – tion, secured permits to work in the protected area, and .  played a secondary role in editing. SKALSKI, J.R., HOFFMAN,A.&SMITH, S.G. ( ) Testing the significance of individual- and cohort-level covariates in animal survival studies. In Marked Individuals in the Study of Bird Populations (eds J.-D. Lebreton & P.M. North), pp. –. Birkhauser References Verlag, Basel, Switzerland. SLAGHT, J.C., MAKSIMOVA, D.A., MIQUELLE, D.G., SERYODKIN, I.V.,  AKSENOV, D.E., DOBRYNIN, D.V. & DUBININ, M.Y. () Atlas of MILAKOVSKY, B., ZAYTSEV, V.A. & PIMENOVA, E.A. ( ) The Intact Forest Landscapes of Russia. World Resources Institute Press, influence of logging on musk deer resource selection: preliminary Washington, D.C., USA. results from the central Sikhote-Alin. In Innovative Technologies in

Veterinary Medicine, Animal Husbandry, and Nature Conservation ZAITSEV, V.A. () Musk Deer: Ecology, Population Dynamics, in the Russian Far East. Conference Proceedings, – November  Perspective for Conservation. Biodiversity Conservation Center (ed. L.V. Zhilyakov), pp. –. Primorskaya State Academy of Press, Moscow, Russia. Agriculture, Ussuriisk, Russia. [In Russian] ZAITSEV, V.A., SERYODKIN, I.V., PIMENOVA, E.A. & MAKSIMOVA,D. SLAGHT, J.C., MIQUELLE, D.G. & TUKHBATULIN, G.A. (in press) A. () Necessary measures for the conservation of the musk deer Logging roads and Amur tigers in Russia: demonstrating the threat (Moschus moschiferus), their theoretical basis and the condition of and proposing solutions. Proceedings of the International the protected areas network in Primorsky Province. In Proceedings Conference on the Amur Tiger: Population Status, Problems, and of the Far-Eastern Conference on Nature Conservation Problems, – Conservation Prospects. – December . Institute of Biology  September . BSPU-Press, Blagoveshchensk, Russia. and Soil Science, Vladivostok, Russia. [In Russian] SLAGHT, J.C. & SURMACH, S.G. () Blakiston’s fish owls Bubo blakistoni and logging: applying resource selection information to Biographical sketches endangered species conservation in Russia. Bird Conservation International, , –. J ONATHAN S LAGHT is interested in wildlife distributions (primarily STORAUNET, K.O., ROLSTAD,J.&ROLSTAD,E.() Effects of birds) in the Russian Far East, and in identifying the resources and logging on the threatened epiphytic lichen Usnea longissima:an conservation issues that influence those distributions. B RIAN experimental approach. Silva Fennica, , –. M ILAKOVSKY works on developing practical methods for maintaining STORAUNET, K.O., ROLSTAD, J., TOENEIET,M.&ROLSTAD,E.() populations of rare and threatened species during industrial forest Effect of logging on the threatened epiphytic lichen Usnea management in the Russian Federation. D ARIYA M AKSIMOVA uses longissima: a comparative and retrospective approach. Silva Fennica, radio tracking, camera traps and direct observation to study musk , –. deer ecology at the Sikhote-Alin Biosphere Reserve in Russia. I VAN SWITALSKI, T.A. & NELSON, C.R. () Efficacy of road S ERYODKIN is a specialist on large predators and ungulates, with a removal for restoring wildlife habitat: black bear in the focus on their conservation in the Russian Far East. V ITALIY Northern Rocky Mountains, USA. Biological Conservation, , Z AITSEV studies the ecology, behaviour and population dynamics –. of ungulates and predators, the structure of bird and mammal TUKHBATULIN, G.A. () Influence of Timber Harvesting and communities, and wildlife management and research methodologies. Hunting on Musk Deer Population Density. Unpublished report. A LEXANDER P ANICHEV studies ungulate ecology, with a focus on ex- Biological Soil Institute, Far Eastern Branch of Russian Academy of plaining the causes of their geographical distribution. D ALE Sciences, Vladivostok, Russia. [In Russian] M IQUELLE coordinates the Wildlife Conservation Society’s conserva- ZAITSEV, V.A. () Musk Deer of the Sikhote-Alin. Ecology and tion efforts in Russia, with a focus on developing strategies to conserve Behavior. Nauka, Moscow, Russia. [In Russian] and restore Amur tiger habitat.