Sugar or a One-Eyed Horse

On the far western edge of Mongolia, in the Altai Mountains of Bayan Olgii aimag, a national park nudges the border with China. This place is called Chigertei (Чигэртэй), although it is sometimes spelled Chikhertei (Чихэртэй), which is how I read and wrote it for the few months between first hearing about it and actually setting foot there. The g-versus-kh debate is a reminder of the region’s unique cultural dynamics; the majority of Bayan Olgii’s population is ethnically Kazakh, descendants of a small number of refugees who settled in Mongolia during various 18th and 19th century conflicts with the Russians and Chinese. In 1940, the socialist government of Mongolia formed Bayan Olgii as a Kazakh aimag, creating a place where Kazakhs were allowed to maintain traditions that were more harshly curtailed in Kazakhstan itself, which was then part of the Soviet Union. Following the advent of democracy in 1990 and the opening of Mongolia to the wider world, some of these traditions became world renowned. Eagle hunting and the richly embroidered Kazakh ger hangings made for weddings are synonymous with the region. Although all official business in the aimag is conducted in Mongolian, school is taught in Kazakh, Kazakh remains the language of everyday interactions, and many families in more remote regions don’t speak much Mongolian at all. Bayan Olgii is a region where Mongolian and Kazakh run up against each other, culturally and linguistically, in interesting ways.

This is the source of the Chigertei/Chixertei mashup. Chixer means “sugar” in Mongolian, and Chixertei, roughly translated, means “sugared,” which, as a Mongolian speaker, I took to be a poetic reference to the snow that girds the Altai peaks for most of the year. Chigertei, on the other hand, means “a one-eyed two year old horse” in Kazakh. A long time ago, my Kazakh counterpart from the park administration told me as we jolted along the road to the park, there was a one-eyed two year old horse that hung out in the valley, and that’s how the place got its name. Why do Kazakhs have a specific word for a one-eyed two year old horse? Are there a lot of one-eyed horses running around Kazakh-populated areas? Is there a different word for a one-eyed three year old horse? Mongolian is like this in its precision – there are verbs to describe a calf running around with its tail in the air, or the act of tying ribbons in a horse’s mane to designate it sacred. There are dozens of words to describe the colors of a horse’s coat, and a whole herd of words to describe yak-cattle hybrids depending on the generation of hybridity. There are several words for “friend,” some of which indicate a greater degree of affection and closeness than others. There are a whole set of verb tenses that you employ to indicate how recently something happened, and how certain you are that it actually happened if you didn’t witness it with your own two eyes. But with Kazakh, I had no idea where things get specific, because the sum total of my Kazakh skills could be summed up in the two most essential phrases in any language: “Thank you” (Rakhmet) and “Are there any wolverines around here?” (Kunnu bar ma?) The exchange on the road to Chigertei served as a reminder that I was on strange ground. I’ve worked in Mongolia for 17 years and I’m used to being agile and fluent, culturally and linguistically. It felt both off-balance and also exciting to be in a place where I didn’t know what was going on 100% of the time.

Whatever direction you take the name, Chigertei appears to be full of wolverines, and that was why I was there. In May, a colleague who had set up cameras for snow leopards in Chigertei contacted me to tell me that he had gotten dozens of photos of wolverines, and would I be interested in collaborating on a project? I’d been to the Altai before as part of my wolverine quest, most recently in 2011, to interview and collect pelt samples. I’d wanted to work there on a more extensive and systematic program, but hadn’t had the time to do the most laborious preparatory task for any wildlife work in Mongolia – setting up relationships with the various authorities and finding local counterparts who were interested in collaborating. Barry Rosenbaum, the wildlife biologist who had put up the snow leopard cameras, had already done that work, so it was easy to agree. Easier still because snow leopards were my first wildlife love, the animal that had led me to the high mountainous landscapes that wolverines inhabit. Barry would be collaring in Chigertei’s sister park, Khokh Serkh, which meant that I might, if I were very lucky, have the opportunity to see an animal that held enormous personal significance.

Luck seemed to be in short supply for the first ten days after we set snow leopard snares in Khokh Serkh (the name, definitively Mongolian, means “dark blue billy goat” and refers to both the Special Protected Area and the mountain range encompassed by the SPA). The plan was to spend the first ten days of the trip collaring snow leopards, and then a week in Chigertei, 40 kilometers to the west, setting up baited wolverine camera stations. After a frigid first two days in the Khokh Serkh base camp, the weather warmed, and the Mongolians and Kazakhs insisted that the snow leopards would stay up high and away from our snares, snoozing in the sun, until the weather changed again.

Whatever the cause, the snares remained empty. The cameras we’d set up nearby yielded beautiful photos of ibex, quail, and ermine, but no sign of big cats. I was happy to be out on a snow leopard expedition, but it was mostly waiting, interspersed with the daily anticipation and letdown of heading up to the higher valleys to check the snares. The time between was spent rambling around the mountains, but there was something edgy and almost spooky about the steep, narrow, rocky valleys, dark in the scant light of the failing year. On one of our daily excursions to check the snares, my horse’s cinch strap slid back, the horse panicked, and I was thrown, dragged through a boulder field on a steep slope, and kicked twice. The damage was relatively superficial – bone bruises, a torn ligament or some other trauma in my right wrist, the most shockingly purple hoof-shaped bruise on my left buttock – but I couldn’t hike for a while afterwards, or sit on a horse, which meant several long days sitting in camp trying to figure out how to do anything useful when my only functional limb was my left arm. I was more than ready to get out of Khokh Serkh by the time we piled into the range rover and headed to Chigertei.

The land of sugar and one eyed horses was enchanting by comparison, a wide open swath of valley flanked by dramatic high peaks, the weather moving fitfully and spectacularly across the mountains and the river and the moraines left behind by recent glaciers, and wolverine tracks everywhere.

The high peaks on the western border of Chigertei Valley.

We stayed with one of the few families that winters in the valley – there are many more in the summer. The family were relatives of Yelik, our liaison at the park. His forebears had lived in Chigertei Valley for generations, he said, and pointed to a crumbling cemetery that we passed on the drive in, and said, “My family are all there. Two hundred years.” His cousin Yerlan, her husband Jaksalak, and their daughter Raya were our generous hosts, giving us space in their adobe house, renting horses to us, and regaling me with tales of wolverine sightings up and down the valley. For the first time ever in eight years of wolverine work in Mongolia, I heard stories of regular wolverine depredation on small livestock. Jaksalak told me, through translation into Mongolian by our ranger Aska, that he’d seen five wolverines together in the early summer, that he’d spooked them out of a thin line of forest that ran east-west along the north-facing slopes of the valley, and that they’d fled up into one of the big bowls above treeline. Five wolverines together could only be a family, and if it was a female and her kits-of-the-year, that meant she’d successfully nursed a litter of four. When I asked where they habitually saw wolverines, Jaksalak and two fellow herders waved their hands across the valley: everywhere.

Adobe and stonework home in Chigertei

It was pointless to speculate based on unverified stories, of course, but the herders of Mongolia know their wildlife well – far better than the Americans who report wolverines in their backyards and then send me photos of porcupines or foxes – so I was inclined to believe them. The photos from the snow leopard cameras showed at least two individuals visiting the same camera station within days of each other, and returning repeatedly.  It was all very tantalizing. But we’d learn more only by doing some scientific documentation.

The first day out, back in the saddle for the first time since being thrown off, I had a hard time focusing on where I was, too busy clutching the reins and trying hard not to think about the horse slipping on the ice, kicking me, dragging me across boulder fields, snapping my neck. Then we came up over a massive moraine and stopped at a boulder covered in petroglyphs, and all the crowding fearful thoughts disappeared as we dismounted and looked at the animal figures chipped from stone ages ago. Aska opined that the figures were little wolverines; I thought they were ibex. They were probably from the Bronze Age, which, in Mongolia, extended from about 2000-1000 BCE. There are petroglyphs like this all over the country – very ancient ones showing woolly rhinoceros and ostriches and mammoths, more recent scenes of men in chariots and reindeer with the beaks of birds. Out of all of this vast, Pleistocene world, the wolverine remains like a talisman of a lost past.

Petroglyphs. Depending on which end you see as the head, you can imagine these as running ibex with curved horns, or open-mouthed, stripe-tailed carnivores.

We rode further up the moraine and came out into a side valley, walled in at its head by another jagged range of peaks. From here we went on foot up the valley, and then up an east-facing slope, walking carefully among the snow-covered large-boulder talus. Before long we crossed wolf tracks, and then, inevitably, the prints of a wolverine. They led up through the talus field onto the ridge, where they and the wolf tracks joined a well-worn game trail running along the crest of the slope.

Dukhai and Aska on the wolf and wolverine highway

We set up four cameras in that valley, all of them along the east-facing slope. The west-facing slope was sheer scree and bare of snow, steeper and higher, far less appealing as a travel route. Aska and Dukhai, another of the winter herders, guided the effort, choosing spots where they’ve seen wolverine tracks in the past.

That night the wind rose until it howled around the outside of the house. Inside, Aska practiced his English by reading aloud from the back issues of the New Yorker that I bring with me on these trips; his pronunciation was flawless, and after a a sentence or two he would tell me, in Mongolian, what he understood, and then I would explain where he’d understood correctly and where he’d gone wrong. Since it was the New Yorker, this necessitated trying to explain phrases like “pathologically incompetent president” and “hipster locovore Brooklynites,” which was both headache-inducing and also a welcome challenge to improve my Mongolian skills. Neither Yerlan nor Jaksalak spoke Mongolian, although Yerlan liked to order me to eat more of the various dishes she cooked, thus improving my Kazakh vocabulary by about 1000% in the space of 24 hours. At some point, I explained to Aska in Mongolian that the wolverine’s Latin name meant “glutton” and that it had a reputation for eating enormous amounts of meat; Yelik, overhearing this, explained to Yerlan and Jaksalak in Kazakh; and everyone began saying “Gulo gulo!” before digging in to whatever dish Yerlan put on the table in front of us. Most of the time this was besbaramak, the signature Kazakh dish, which involves most of a sheep or goat, boiled for several hours, and served with enormous flat noodles, boiled carrots and potatoes, and sometimes a side of horse sausage. Invoking the wolverine came shortly after we washed our hands and bowed our heads and lifted our palms up for a quick blessing on the food from Allah. Sometimes I ended up translating whatever I understood back into English for Barry, although I’m terrible at remembering to do this, because translation is a task that requires focus. Later that evening, Aska and I digressed from New Yorker-level topics into our mutual enjoyment of the show Vikings, discussed which characters we most liked, and chatted about Norse history and Kazakh history. There was more comfortably cosmopolitan code-switching going on in the confines of that little adobe house, under the howling wind in the remote Altai, than in any place I’ve been in a while.

The wind went right on howling, and when we got up the next morning, a blizzard was beating down on Chigertei. Aska and Yelik swore it wasn’t as bad as the snowstorm that had hit the valley a few weeks before, so with that in mind, we all put on our warmest clothes and rode out.

Parking the horses in the blizzard.

It could have been worse. It could have been a few degrees colder, the wind might have had an even sharper edge, the snow could have been thicker, the visibility even more curtailed. As it was, I could see the dim outlines of the near mountains, if I squinted into the wind and snow, and I could remind myself that I’d been far colder and more uncomfortable, once or twice, on the ski transect of the Darhad that I’d done back in 2013. It was tolerable, in other words, in the minimal sense of the word. We were riding further out that day than we had the day before, but over less tricky terrain. The conditions did not make the horse any less nerve wracking, and Yelik and I had a moment of mutual vocabulary-building when I explained that I was nervous about galloping the horse in icy and snowy conditions. “Nervous” was new to him, and I realized with shock that I’d never used that word in Mongolian before – how had I avoided having to learn it when I’d dealt with everything from ornery horses to psychotic and potentially rabid dogs to drunk herders breaking into my ger?

I saw almost nothing of the valley we reached, except for the sites where we set the cameras. Those sites emerged out of the blinding white as we stumbled towards them, knee-to-thigh deep in drifted snow, the whole landscape eerie and markerless. We left the horses behind near a stone wall enclosing a hay field or corral and waded upslope. These stations we baited with the flanks of a goat, roadkill not being an option in Mongolia as it was in the US.

Dukhai, Aska, and Yelik setting up a camera. They did 100% of the physical labor since I couldn’t use my wrist. They deserve all the credit in the world for the success of this trip.

The crew resting at the top of the slope after wading up through the snow.

Mongolian snow conditions – never fun, but somehow still entertaining.

For our pains, whatever powers exist out there granted us bluebird conditions the next day, which took us up the north-facing slopes and through the thin larch forest to the base of the massive north-facing bowls. Where these bowls narrowed, and along obvious wildlife highways running east-west, we set the last of the cameras, baited with the rest of the goat.

On the north-facing slopes at the edge of the forest, glassing for elk, ibex, and argali

Yelik and Dukhai setting up a bait station.

I stayed high up on the mountainside, walking back along the wildlife trails; Yerlan and Jaksalak’s home was perched beneath the slope several kilometers to the east, and, in a move that probably fooled no one, I pretended that wading through the deep snow at high elevations for all of that distance was super fun, because my horse – which was really Aska’s horse, he’d trained it from a colt – had been edgy all morning on the ride out. I was happy to turn him over to Aska and hike back.

Headed home.

I hit a set of wolverine tracks and followed it up into the trees, out of sight of the group of men riding their horses across the plain below. Suddenly there was yelling, a huge commotion from the men, ringing dimly off the trees. I’d followed the tracks far enough into the forest that by the time I came back out, all I saw was the group of them leading their horses back to the house. They were on foot, which was strange, but I thought that they’d gotten off for a smoke break. I arrived back maybe a half hour after they did, opening the door and ducking into the dim light of the interior, ready to prattle on about the exciting wolverine tracks, but everything inside was silent and heavy and the hair went up on the back of my neck.

Yelik shook his head and said, in English “Today was so scary.”

Aska was lying on the floor, on the rolled out thin mattresses on which they slept, and I remembered the shouting and knew immediately that something had happened with the horse. And it had; while I’d followed those tracks up into the forest, Aska had stopped to take a photo. He’d spurred the horse to catch up with the others, and the horse had hit a wind-scoured snow drift; his front legs had plunged into the snow, Aska had slid forward so that his boots jammed into the stirrups, and then slid off the right side of the horse. In Mongolia horses are freaked out by anything approaching on the right, including riders who slide down on that side. The horse bolted, dragging Aska at a full gallop for somewhere between 30 and 50 meters through a boulder field. He was kicked in the shins, just as I had been, before he got free of the stirrups.

“I thought he was going to die,” Yelik said. And then he added, “Now I am nervous.”

Aska was stoic – more so than I’d been, definitely. His shins didn’t look bruised, although that didn’t mean anything; mine hadn’t visibly bruised either, but they ached terribly, some deeper, invisible bruise in the bone. More than the physical, though, he seemed shaken up by the flashbacks; after an hour or so he asked if I had any sleeping pills, because every time he shut his eyes he kept reliving the fall. I didn’t, but I gave him some advil and told him it should help, and apparently it did, because he said that he slept fine.

That night we enjoyed a final round of “Gulo gulo” over besbaramek. The next morning we set up a final bait station just above Yerlan and Jaksalak’s home. Aska hiked up with me to set the cameras, to prove that he was okay, and then rode his horse 40 km back to the town of Deluun. Watching him ride off so confidently, I took a deep breath and promised myself to stop being such a wimp about the horses henceforth. The enormous bruise on my butt, which I surreptitiously checked the day before when I was up in the forest alone, was starting to look less black and was fading to a violent dark purple. My wrist was still a problem, but I took the improvised cardboard brace off and decided to deal with it later, even though it still hurt to, say, hold a pen, braid my hair, or scratch my back. Horse danger was a fact of life in Mongolia and you either dealt with it or you were incapacitated by your need for safety.

The rest of us rode back to Deluun in a land rover. Over the winter, Aska would rebait the camera stations as needed, and Yelik would switch out the cards in December to see which stations were receiving wolverine visits. I was pretty sure that most of the bait would be devoured by foxes, but there were enough wolverine tracks to make me hopeful. Ultimately, we wanted to identify the best places to set up a trap or two for collaring, and, if we were very lucky, find places where multiple wolverines were visiting the cameras. The only thing to do now was wait, and head back to Khokh Serkh in hopes that we would capture that even more elusive animal, the snow leopard.

 

 

 

 

 

 

 

 

 

 

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Return to the Altai

A couple of weeks ago I got a call from a colleague who suggested, politely but also pointedly, that I should probably start updating this blog again. I said yes, of course, I really had every intention of doing so, very soon.

Then, when I was on the phone with my parents yesterday, talking about my upcoming travel plans, my mother, politely but perhaps also pointedly, said, “Well, maybe this trip will provide some good fodder for your blog?” And I realized that I hadn’t even posted a single update about my upcoming plans, despite the fact that they are both wolverine-relevant and also pretty exciting.

Early tomorrow morning, I leave for Mongolia to set up camera traps in the Altai for wolverines. The Altai form Mongolia’s western border with China and Russia, and contain Mongolia’s highest peaks. The glaciated landscapes of Altai Tavan Bogd National Park are breathtaking, as I was made aware when I first visited this region in 2001. That was before I was very aware of wolverines, but not before I was aware of the other compelling denizen of the higher elevations of Asia, the snow leopard. I returned to Tavan Bogd and neighboring Khovd Aimag in 2010 to conduct interviews on wolverines and other wildlife, and had some thrilling encounters and near-encounters while I was up in the mountains.

This trip is in cooperation with a snow leopard project affiliated with the Denver Zoo, which means that I’m not only camera trapping for wolverines, but may also have the chance to fulfill a 17-year-long dream of seeing a snow leopard in the wild and participating in snow leopard collaring activities. So it’s pretty darn exciting. I’m also exhausted, however, and have about a dozen other writing projects right now, so sometimes it’s hard to remember to keep this blog up to date. But my mother is probably right – as mothers usually are. This trip should be story-worthy.

More details will be forthcoming once I reach Ulaanbaatar – thanks for reading and stay tuned.

 

Territoriality in Female Wolverines

For reasons that I’ll get into in a future post, I feel suddenly compelled to write up a quick literature review on the topic of territoriality in female wolverines.

Let’s get this out of the way immediately: female wolverines are territorial. This is evident throughout the recent literature. Out of concern for the safety of resident adult wolverines in the US, researchers generally don’t publish maps of wolverine home ranges, but anyone who has had the chance to look at the minimum convex polygons of study animals can see a pretty clear pattern. Adult female wolverines don’t overlap with other adult female wolverines. The territorial boundaries are pretty strict, often down a drainage that both animals will approach and traverse, but that neither will cross. There are instances of incursions and small amounts of overlap, but this is generally less than 2% of any female’s home range. Adult males are also territorial, but with varying degrees of territorial incursion or excursion. The territoriality of wolverines is one of the most important factors for understanding wolverine distribution, their natural rarity, and, importantly, conservation concerns and strategies as we look at a warming world.

Strictly speaking, a territory is a patch of ground that an animal defends, and a home range is a patch of ground that an animal occupies and uses for subsistence. In the wolverine research community, including in published papers, these terms are used in a conflating fashion, although some papers do make a distinction. In presentations and talks and casual conversation, we often use the words “home range” to refer to the area that a wolverine (male or female) occupies, but we mean this as a defended home range, which is actually a territory.

Territoriality in the Literature

While the territoriality of wolverines is widely understood at this point, I was surprised to note that there are few American or Canadian papers or book chapters that deal explicitly with the question of how territoriality functions, or the ways in which it potentially limits population in areas of restricted habitat where the population fragments into a meta-population structure. The Scandinavians, on the other hand, do tend to deal with territoriality, probably because the management imperatives are more urgent in places where depredation on domestic reindeer is a major problem. A Norwegian master’s thesis from 2014 looks at genetic sampling as a method to determine territoriality, in comparison to GPS collar data. Jens Persson’s dissertation deals in some depth with possible explanations for territoriality, and how those explanations may differ between males and females. Proving that the Scandinavians more or less own this topic, a 2017 thesis by Malin Aronsson closely focuses on the dynamics of territoriality and dispersal among female wolverines (and lynx). But the territoriality of the species is taken for granted in most publications, especially the North American publications.

I was also reminded of the absence of peer-reviewed publications focusing specifically on several of the major American wolverine research projects. Doug Chadwick’s popular-science book The Wolverine Way remains the best published account of the Glacier wolverine project. The Rocky Mountain Research Station has put out a number of impressive papers that draw on the Glacier data to ask large-scale questions about habitat relationships, population genetics, dispersal, and climate change effects, but the annual project reports remain the best source of information for the park population itself. The same is true of the Absaroka-Beartooth project, the project with which I got my start in the wolverine world, working as a volunteer. Peer-reviewed papers for that project were somewhat limited by the low numbers of detected wolverines within the study area – sample sizes and statistics being the perennial priority for most journals these days – but the eerie emptiness of prime modeled wolverine habitat deserves some consideration, hopefully in future publications.

Back in 1981, Hornocker et al published a study asserting that wolverines were not territorial. This was one of the earliest studies of wolverines in the lower 48, lasting from 1972-1977, on the Flathead National Forest. It was a telemetry study in which the researchers observed overlap among many different wolverines, and concluded that wolverines were tolerant of fellow wolverines. They reported “no intraspecific strife” and discussed how a wounded female wolverine, whose injuries they first attributed to another wolverine, were likely caused by a mountain lion. (Despite assertions that wolverines are not territorial, the home range maps that are included in this paper do show a familiar pattern – two male wolverines who don’t overlap, one female wolverine who sticks to a fairly tight home range, and a second female, overlapping the first, who makes wider movements that include the home range of the male who overlaps the first female. If I were to guess, many years later, what was up with this scenario, I’d suggest a male-female pair and their juvenile daughter, preparing to disperse. The second male looks like an unrelated individual as his range does not overlap with any of the other animals.) Earlier observational studies of wolverines, and books about the species, also fail to make note of territorial behavior. The same is largely true for the Mongolian hunters and herders who I interviewed; they were well aware of the wolverine’s rarity and ability to travel over long distances, but only a few noted that a wolverine would reappear in a particular spot at intervals. None of those interviewees made a leap to territoriality as an explanation, but some did refer (maybe jokingly) to a wolverine’s nutag, which is a Mongolian concept denoting an individual’s homeland.

Home range maps showing seasonal movements for four wolverines in NW Montana, from Hornocker et al 1981

Home range maps for four wolverines in NW Montana, showing seasonal movements, from Hornocker et al 1981

Observation, track surveys, and radio telemetry, of course, are limited in important ways. These methods allow glimpses of an animal only at the moments when the observer or listener happens to have their eyes or telemetry antenna trained on the animal. Finding dens is more difficult, which – in the absence of DNA techniques – makes understanding relatedness more difficult as well. With the advent of GPS collars and DNA analysis, we were able to observe wolverines more closely and consistently. Hundreds of locations for multiple animals, taken over months, in combination with VHF locations over the course of years, made the territorial behavior of the species clear. Den locations and kit collaring showed that wolverines will tolerate their own offspring within their territories for up to two years after birth. This could account for Hornocker’s and others’ observations of multiple wolverines sharing the same home range, and photos like those that Igor Shpilenok took in Kamchatka, of up to six wolverines on a bear carcass at the same time.

The Wildlife Conservation Society’s Yellowstone Wolverine Project likewise relied on annual reports and white papers to convey results for many years. The project director, Bob Inman, was in the process of getting his PhD, so his dissertation eventually yielded several published papers. Again, though, these deal mostly with larger-scale questions about habitat relationships and conservation priorities. Several of his papers do discuss territoriality, but again, for obvious reasons, most of the published papers don’t include home range maps.

What all of these papers and sources do include, however, is an assumption that territoriality is important. Some imply territoriality; for example, in a summary of draft papers from 2007, the WCS project notes in the abstract for a chapter on wolverine space use:

Mean annual (1 Mar–28 Feb) 95% fixed kernel home range size was 453 km2 for adult females (n = 15 wolverine years) and 1,160 km2 for adult males (n = 13 wolverine-years). Mean percent area overlap of same-sex adults was < 1% (SE = 0.00, range = 0–2%, n = 12 pairs) using annual 100% minimum convex polygon home ranges.

The “<1% overlap” suggests territoriality, although it’s not explicitly stated here. In a 2012 review of wolverine reproductive chronology, however, Inman et al do note this:

Throughout its distribution, the wolverine displays extremely large home ranges, territoriality, low densities, and low reproductive rates (Copeland 1996; Inman et al. 2012; Krebs et al. 2007; Lofroth and Krebs 2007; Magoun 1985; Mattisson et al. 2011a; Persson et al. 2006, 2010). These adaptations are necessary for exploiting a cold, low-productivity niche where growing seasons are brief and food resources are limited (Inman et al. 2012).”  

Another 2012 Inman et al paper on spatial ecology does deal explicitly with the question of territoriality vs. undefended home ranges and is probably the most extensive discussion of this topic in the literature on wolverines in the lower 48. This paper does include some home range maps, with the locations stripped out.

Spatial distribution patterns of the Mustelidae are typically described as intra-sexual territoriality, where only home ranges of opposite sexes overlap (Powell 1979). Wolverine-specific reports exist for both intra-sexual territoriality (Magoun 1985, Copeland 1996, Hedmark et al. 2007, Persson et al. 2010) and for a high degree of spatial overlap but with temporal separation (Hornocker et al. 1983). Arguments against territoriality by wolverines include the lack of ability to defend such a large home range (Koehler et al. 1980). Our data on movement rates in relation to home range size, temporal development of the home range, minimal overlap of same-sex adults, and relatively immediate shifts upon a death suggest that wolverines are capable of patrolling a large territory and provide further support for intra-sexual territoriality. Reproductive success is closely correlated to the amount of energy that a female wolverine can obtain (Persson 2005), and for predators that are capable of individually acquiring prey, the presence of conspecifics reduces foraging efficiency (Sandell 1989). Since wolverines feed on individually obtainable prey and occupy relatively unproductive habitats, it follows that behaviors for maintaining exclusive access to resources would likely have selective advantage. Frequent marking behavior (Pulliainen and Ovaskainen 1975, Koehler et al. 1980) is likely part of an adaptive strategy that involves maintenance of exclusive territories within sexes so that feeding and breeding opportunities are monopolized by dominant individuals and their immediate offspring.

Female wolverine territories in Wyoming, with takeover of one home range by a female kit after the death of the resident adult female. From Inman et al 2012

The Scandinavian literature is also rife with references to territoriality among both male and female wolverines. Jens Persson, who works on wolverines in Sweden, reflects in his dissertation on the reasons for territorial behavior in female wolverines, and concludes that it’s related both to food, and also potentially to the need to protect kits against infanticide. Historically, it was widely believed that male wolverines would kill any kits they encountered, even their own; this has since been proven false, but the idea that males kill unrelated kits persists. Persson is the first researcher I’m aware of to suggest that female territoriality may actually be a defense against other females intent on infanticide.

Females could also gain from infanticide by eliminating non-related progeny to decrease future competition for territories or denning areas for her and her progeny. In addition, the death of an unrelated infant could also reduce the net reproductive success of a competitor (Hrdy & Hausfater, 1984). Competition for territories determine dispersal behaviour in female wolverines (Paper IV), suggesting that there is strong competition for territories among female wolverines.

 Wolff and Peterson (1998) hypothesized that a primary function of female territoriality in solitary mammals could be to protect vulnerable young from infanticidal conspecific females. Four predictions can be deduced from their offspring-defence hypothesis: 1) Female territoriality should be associated with young that are vulnerable to infanticide. 2) Female territoriality should be associated with defence of offspring, and therefore most pronounced during the offspring-rearing season. 3) Defence will be greatest against the segment of thepopulation that commits infanticide and against those individuals that females can dominate. 4) Optimal territory size should be a function of intruder pressure, intruder detectability, female response distances and offspring vulnerability, and changes in food abundance and distribution should not affect territory size directly unless they are correlated with the other factors. In concordance with predictions 1-3, wolverines have altricial young that are vulnerable from late winter until late summer (March – August) and female territoriality seem to be strongest during this period (Magoun, 1985; Landa, Lindén & Kojola, 2000). We lack data to evaluate prediction 4. However, in contrast to prediction 4, I believe that food actually is an important determinant of territory size in wolverine females (see Banci, 1994).

Malin Aronsson’s 2017 thesis examines the territorial dynamics of female wolverines, using the vast dataset from Swedish studies dating back to the 1990s. She makes some interesting observations about the counter-intuitive conclusion that wolverines are territorial despite living in low-resource environments, which is the opposite of what studies on other carnivores would suggest:

Wolverines are highly territorial (Persson et al. 2010), and by comparing space use overlap between years for the same individual I found that wolverines show high territorial fidelity resulting in a stable distribution of resident individuals. Interestingly, territorial fidelity in general is predicted to be low in habitats where food resources are low, variable, unpredictable or deplete fast (Wauters et al. 1995; Kirk et al. 2008; Edwards et al. 2009), which corresponds to the characterization of wolverine habitat in general (Inman et al. 2012b), and particularly in this study area (Person 2005). However, scavenging and caching are integral parts of wolverine biology (Inman et al. 2012b; Mattisson et al. 2016), which increase resource predictability, decrease depletion rate and create a valuable resource (i.e. cache sites) to defend, promoting high territorial fidelity despite the unpredictable environment (Tye 1986; Eide et al. 2004). In addition, occurrence of more efficient predators, such as the Eurasian lynx (Lynx lynx), provide carcasses for direct consumption and caching (Mattisson et al. 2011b). Furthermore, both males and females showed higher between-year fidelity at the territory level (i.e. 90% isopleth) compared to the core areas (i.e. 50% isopleth). That fidelity was lower at the core area compared to territory level suggests that it is critical to maintain the outer territory boundary to secure long-term resources, while the most used area within the territory may vary between years due to spatial fluctuations in key resources, or, for females, location of den sites may vary between years.

Aronsson also documents a few cases of territorial adult females shifting territories after successfully reproducing. Why this happened, and how frequently such moves occur, would be interesting questions for further investigation.

Another Scandinavian paper, a master’s thesis from 2014 by Espen Gregersen, compares home range and territoriality derived from GPS data to those derived from scat analysis. The question in this thesis was not so much “are wolverines territorial?” as “can wolverine territoriality be detected using non-invasive methods like scat analysis?” It’s an interesting question and one of relevance to those of us who have limited resources for large-scale trapping efforts in places like, say, Mongolia. Gregersen concludes that yes, we can indeed determine home range size and observe territoriality using scat samples – but it takes a very large number of samples. This thesis also addressed the question of territorial turnover after the death of a resident adult, which is particularly interesting at the southern edge of distribution, where wide separation of habitat patches makes recolonization less certain.

Finally, a forthcoming book chapter from Copeland et al. proposes a slightly different take on territoriality among male and female wolverines. That chapter will be out soon and I’ll look at it in depth once it’s published, but it too reinforces the idea that female wolverines are highly territorial, and maybe even more strictly territorial than males.

There are many other papers out there that include brief mentions of territoriality and intrasexual exclusion in home ranges. These are just a sample, and this write-up fairly cursory, but I hope they’re adequate to illustrate that wolverines – both male and female – are territorial.

Territoriality and Conservation

At this point, territoriality in wolverines is accepted as an important feature of their life history and ecology. The question of why hasn’t yet been answered, but the fact that wolverines require such large territories, and the fact that their reproductive rates are so low, accounts for their natural scarcity on any landscape – let alone one in which suitable habitat is located only at certain elevations in widely scattered patches across a sea of non-habitat.

Female territories structure the wolverine population. Females must have adequate resources to meet their needs and, hopefully, to reproduce. They occupy and defend territories that allow them to do this. Males in turn seem to select for territories that overlap with resident females. Whether this represents a territorial strategy for sexual monopoly, or whether it’s defined by the male’s capacity for paternal investment, or some combination, is worth investigation (male wolverines seem not to always entirely encompass a female’s territory, leaving her open to potentially overlap with other males, which raises questions about the accepted narrative of males “controlling” access to females). Both males and females disperse over long distances, although the longest movements have been observed in males like M56.

Habitat availability for females is the limiting factor on wolverine population growth and range expansion in the US Rockies. I’d hypothesize that keeping a certain number of territories occupied is critical to the long-term persistence of wolverines in the lower 48, and that there is some distinction to be made between female population numbers, strictly speaking; the percentage of habitat that’s occupied; and where that habitat is located in relation to other habitat. There’s been an enormous focus on “connectivity”– concurrent with the fashion for corridors among conservationists –  but a surprising lack of attention paid to the population nodes themselves. For example, the question that Gregersen raises about recolonization of vacant territories is interesting and important, especially given the observed disappearance of wolverines from places like the Tetons. Presumably this disappearance represents some kind of natural cycle of die-off for a relatively isolated population node, but how long does it take before those territories are reoccupied? And how is time-to-recolonization related to population density and occupancy of the next-nearest population nodes? Questions about functional connectivity among wolverine population nodes are important, but connectivity as a single conservation strategy for wolverines seems like an odd allocation of resources; wolverines don’t migrate, they disperse, and their dispersal patterns are unique and erratic. They are likely to benefit from the broad and intense focus on connectivity and road-crossing structures for other species, but trying to preserve wolverine-specific corridors seems like a good recipe for driving oneself nuts. As one of my Mongolian interviewees once said when discussing wolverines, “One day it’s here, the next day it’s 50 kilometers away. It could turn up anywhere!” I hope to see a greater focus on what’s going on within habitat in the future, including investigation of questions about what drives territoriality and territory size, and how territorial turnover works in a meta-population.

That’s it for now. If any of you have any thoughts about the function of territoriality in female or male wolverines, if you want to point out an obvious resource on this question that I overlooked, or if you just want to say hi, please feel free to comment.

References (with apologies for lack of consistent style formatting and for referring to multiple authors as “et al” instead of writing them out. Time constraints!)

Aronsson, M. 2017. ‘O Neighbour, Where Art Thou?’ Spatial and social dynamics in wolverine and lynx from individual use to population distribution. Doctoral dissertation. Swedish University of Agricultural Sciences. Uppsala. ISBN (electronic version) 978-91-576-8822-4

Chadwick, D. 2010. The Wolverine Way. Patagonia press.

Copeland, J. P., Landa, A., Heinemeyer, K., Aubry, K. B., van Dijk, J., May, R., Persson, J., Squires, J., and Yates, R. 2017. Social ethology of the wolverine. In: Biology and Conservation of Musteloids. Edited by David W. Macdonald, Christopher Newman, and Lauren A. Harrington: Oxford University Press. DOI 10.1093/oso/9780198759805.003.0018

Gregersen, E. 2014. Assessing territoriality in wolverines (Gulo gulo) using non-invasive genetic sampling. Master’s thesis. Norwegian University of Life Sciences.

Hornocker, M and H Hash. 1981. Ecology of the wolverine in northwestern Montana. Canadian Journal of Zoology. V. 59. pp. 1286-1301.

Inman et al. 2007. Wolverine space use in greater Yellowstone. In Greater Yellowstone Wolverine Program Cumulative Report. Wildlife Conservation Society.

Inman R. et al. 2012. The wolverine’s niche: linking reproductive chronology, caching, competition, and climate. Journal of Mammalogy 93(3):634-644.

Inman, R. et al. 2012. Spatial Ecology of Wolverines at the Southern Periphery of Distribution. Journal of Wildlife Management. 76(4). 778–792. DOI: 10.1002/jwmg.289

Persson, J. 2003. Population Ecology of Scandinavian Wolverines. Doctoral dissertation. Swedish University of Agricultural Sciences. Uppsala.

 

Support the Wolverine Foundation through Give Big Gallatin Valley

Wolverine enthusiasts, please consider supporting the Wolverine Foundation during the 24-hour Give Big Gallatin Valley fundraising event. It starts tonight, May 4th, at 6 pm and ends tomorrow, May 5th, at 6 pm. All donations made through TWF’s Give Big donation page during the event will be partially matched by the Bozeman Area Community Foundation, increasing the impact of your support for wolverine research and conservation.

The Wolverine Foundation has been integral to the advance of wolverine science and management over the past 20 years. The organization supports research projects with small grants, conducts education and outreach through talks at community organizations and schools, and supports researchers and students through dialogue, mentorship, and organization of workshops and other events.

Thank you!

 

Wolverines in “Wild Hope”

Last year, author Colleen Morton Busch contacted me to report a possible wolverine sighting in the Sierra Nevada near Tahoe, California. She knew that her sighting, lacking the evidence of photographs or DNA, wouldn’t be conclusive, but her descriptions of the animal she’d briefly spotted sounded distinctly gulo, and we suspected that Buddy, the California wolverine, was still somewhere in the area. We’re always conservative in assessing these sorts of reports, so I had to tell her that I couldn’t consider it a definite sighting, but I felt that it was probable that she’d seen a wolverine.

As we continued to email, the conversation evolved into a meditation on broader themes in conservation, and how those themes tied to Buddhism, with which both Colleen and I have some background. She wanted to write an article about her wolverine encounter that dealt with some of these themes, which made an intriguing divergence from the usual reporter inquiries about species biology and the policy situation around listing. Our ongoing email conversation was a highlight of last spring, particularly as she asked questions about the toll that immersion in the climate change scene takes on researchers. These are questions that people don’t usually ask, and that touch on the weights that we all carry; depression is common among climate researchers and people in affiliated fields. So it was wonderful to talk with someone who was aware of the dynamic between loving what you do, and constantly searching for some small hope – or, failing that, at least the equanimity to continue to love, and to accept impermanence, in the absence of hope.

Appropriately, then, Colleen’s article appears in the most recent volume of Wild Hope, a magazine that celebrates biodiversity and relates well-written stories of species accompanied by lush photography. There is no digital link to this article, but I’d encourage people to buy a copy if you want to read a great reflection on what wolverines mean to the people who are lucky enough to catch even a quick glimpse of one. As scientists, the emotional or psychological meaning of nature and wildlife is a topic that we’re wary of engaging with, but if we’re being honest, most of us would have to admit that we’re in this field in part because of our own dependence on the wild for some form of sustenance, and that we believe that protecting that source of inspiration is important for humanity. So it’s nice to read an account of how much a single, fleeting encounter meant to one person. As Colleen writes, “One wolverine sighting is likely all I’ll get in this life, so I’m grateful to have crossed paths ever so briefly. But seeing the wolverine lit a fire in me. It led to my education. And now I’m telling you, who may or may not live in a state where wolverines can be seen, but who are likely concerned about the changes we humans have wrought on our planet, about any threat of extinction, because the loss of the wolverine is connected to our shared future. Because there’s a glimmer of hope in an encounter between two beings – one wild and the other, a lover of wild things – even if it’s undocumented and unverified.”

A single wolverine encounter changed my life, so I understand this sentiment. There’s something uniquely compelling about this species, something that causes the mind to open in particular ways. Colleen’s captured that in her article, and that’s a great thing. Check it out.

 

 

Weal and Wolverines

Weal is an old word. It goes way back, to the days when a bunch of European tribes were squabbling over Great Britain: Roman invaders building walls to keep wild Celts and Picts from the civilized environs of their empire, Germanic and Scandinavian and Norman French invaders crashing onto the shores of an island where immigrants weren’t necessarily welcome, mixing up their genes and their languages and their culture. Whatever the disagreements of these groups as they contested over the British Isles, words akin to weal are found across large swaths of northern Europe, all related to a single concept: collective well-being. This is the word that gives us “well,” “wealth,” and “welfare,” as well as “Commonwealth,” a political entity that foregrounds the idea of shared responsibility and shared prosperity. Words with similar meanings are found throughout the world, so this is hardly an exclusively European concept; something in the human psyche acknowledges that we are, at our core, a social species responsible for our fellow creatures.

Today we understand wealth as relating to capital, material goods, and private property, but at one time wealth inhered in this shared investment in one’s community. When Puritans overthrew the corrupt monarchy of England in 1649 and decapitated Charles I – a man whose inflated self-opinion and dictatorial policies eventually became untenable to the British public – the nation’s briefly monarch-free government was referred to as a Commonwealth (This is about the time that the first use of the word “wolverine” is recorded in the English language, in customs forms related to fur imports). When those same English Puritans landed on the shores of North America in the 17th century, bringing with them all the good of their anti-monarchial ideals and all the ills of their colonial and religiously intolerant agenda, the land that they colonized eventually became the Commonwealth of Massachusetts, the birthplace of the American Revolution. And if there has always been a struggle between oligarchy and democracy in this country, if there has always been a tension between Enlightenment ideals and terrible colonial realities, the country’s values are nevertheless rooted in an understanding that we come to our community, and to its governance, in a spirit of both individualism and shared investment in the well-being of our fellow citizens.

What does this have to do with wolverines?

Wolverines, like all wildlife, are a public good. They don’t belong to any single person or interest group in the US, which means that we are collectively responsible for them. They are part of the weal of this country, part of the wealth we share. When we discuss what we know about wolverines, when we discuss management of wolverines, we’re talking about how to negotiate within a common-interest space for common-interest outcomes. This process is essential to democracy as well as to conservation of wildlife and the preservation of other environmental values that involve shared resources.

This reality rests uneasily next to a narrative that has gained increasing power over the past few decades, however: the narrative of unfettered neoliberal capitalism. The neoliberal narrative is a study in the privileging of special interests over the well-being of the collective of citizens known as society. It’s a deliberate turning of our backs on the idea that there is a reciprocal relationship between the individual and the world in which that individual lives and makes a living. Neolilberalism privatizes all profit, while socializing the risks and costs of profit-making. It consolidates capital among those who already have it, and denies the idea of the common weal by allowing those with capital to horde it without giving back and without taking responsibility for the ways in which  they impose on the public good. It puts a price tag on everything, and dismisses things that cannot be assigned monetary value. How much is a wild wolverine population worth to the public? This is hard to quantify. Easier to quantify is the value of profit made by oil industry executives and shareholders. The oil industry, in a world that extolls the neoliberal narrative, wins out over public interest. Neoliberalism is a powerfully destructive ideology, insidious and pervasive, and it is this basic set of values that will end up driving wolverines – and other climate sensitive wildlife – to extinction. Unless, of course, we have a massive society-wide awakening, immediately.

To that end, for those of you who are wolverine fans, I’d like to ask you to take a moment to reflect on your values. What are your priorities? How do these priorities potentially have an impact on society? What do you expect or feel entitled to in life, and what do you consider to be your obligations to the wider world? How do you define that wider world, and do you think are there limits to your obligations? If so, where are these limits? Likewise, if you feel like you are entitled to certain things, where are the limits on those entitlements, and do you extend the same set of entitlements to others? If so, which others?

These may seem like large and abstract questions, but I suspect that most people who are concerned with the persistence of wildlife fall within certain parameters when these questions are considered on a spectrum. There’s a lot of work out there in the social sciences on values orientation, the psychology of concern for in-groups vs. out-groups, and where environmental folks tend to fall out on these tests. For now, I’ll provide a couple of graphics and a set of questions to help you situate yourself within this conversation, because this issue of common interest vs. special interest will feature in upcoming posts and you’ll probably find yourself having gut-level reactions to some of what I write. You should know where those reactions are coming from.

Wolverine Quiz: What’s your in-group? 

First, let’s look at in-groups and out-groups. There’s a theory that people subconsciously define other people and entities as being relatively closer or relatively farther from themselves, and care about those people or entities to a degree that is dictated by that distance.

circleofconcern

Beings that are closer to the self get more care. Beings that are further get less. There are entities that are seen as being definitively outside of the scope of care; those beings are the out-group.

In a very restricted or self-centered worldview, the in-group and out-group might look something like this:

circleofconcern_conservative

In a very generous worldview, everything might be part of the in-group:

circleofconcern_generous

Note that these graphics should actually contain another dimension, which is time – for example, I consider living and non-living future generations, up to a certain point, as part of my ‘in-group,’ but I don’t really think about non-living past generations as part of my responsibility since – unless we really take it to a science fiction level – my actions can’t affect them. These are very basic graphics and very general explanations, but you probably get the idea.

Most people will fall somewhere in between the restricted and generous versions of the in-group/out-group dynamics above. Where do you fall? Who or what is in your in-group? If wolverines are in your in-group, why? (If they aren’t, what brings you to this blog?)

That question of why certain entities are included or excluded from our circle of concern leads us back to values. And that will be the topic of an upcoming post.