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Cover: A Grizzly Bear (Ursus arctos) eating seeds from a Whitebark Pine (Pinus albicaulis Engelmann) cone ina 8
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THE CANADIAN FIELD-NATURALIST
Volume 129 MCZ LIBRARY 2015 MAY 18 2015 HARVARD UNIVERSITY Volume 131
The Ottawa Field-Naturalists’ Club Transactions
Promoting the study and conservation of northern biodiversity since 1880
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The Canadian Field-Naturalist
Volume 129, Number 1
January—March 2015
Experimental Evidence of Spatial Memory and Home Range Affinity in White-tailed Deer (Odocoileus virginianus)
MICHAEL E. NELSON
United States Geological Survey, Northern Prairie Wildlife Research Center, 8711 — 37th St. SE, Jamestown, North Dakota 58401-7317 USA; email: mekInelson2@gmail.com
Nelson, Michael E. 2015. Experimental evidence of spatial memory and home range affinity in White-tailed Deer (Odocoileus virginianus). Canadian Field-Naturalist 129(1): 1—7.
The role of spatial memory in the movement of animals through landscapes remains elusive. To examine spatial memory and home range affinity of White-tailed Deer (Odocoileus virginianus) in northeastern Minnesota during 1995—2007, I translocated 17 adult does with known home ranges to unfamiliar sites and radio-tracked them after their release. Twelve does wearing transmitting radio-collars returned to their home ranges. Death and collar expiration precluded determination of whether the remaining five does would have returned to home ranges. Three of five does wearing global positioning system collars traveled throughout hundreds of square kilometres, circling, backtracking, and returning to release sites, while two others exhibited directional movement for tens of kilometres. Four does that survived to parturition stopped traveling and moved at hourly rates similar to those of control does during the first three weeks of the typical fawn-rearing period, but continued traveling later. Their aberrant extensive travel before and after interruption by parturition suggests that they recognized they were in unfamiliar areas, demonstrating both their capacity and propensity to search for and occupy the familiar space of their individual home ranges. Their successful return to home ranges provided experimental evidence of spatial memory and further elucidated its pervasive role in White-tailed Deer spatial ecology.
Key Words: experimental translocation; home range; movements; Odocoileus virginianus; spatial affinity; spatial memory;
White-tailed Deer
Introduction
The role of spatial memory in how animals move through landscapes remains elusive and its quantifica- tion challenging (Moorcroft 2012; Spencer 2012; Fagan et al. 2013). Memory effects in the spatial ecology of animals were postulated in the earliest observations of the movements of individual animals and were eventu- ally confirmed with the development of radio-tracking technology (MJoorcroft 2012). With parallel advances in cognitive sciences and statistical analyses of animal movements, the influence of spatial memory is increas- ingly being recognized and incorporated into mathe- matical models of animal movement (Gautestad and Mysterud 2005; Borger et al. 2008; Gautestad 2011; Moorcroft 2012; Spencer 2012; Gautestad ef al. 2013).
Because spatial memory is an internal cognitive pro- cess, movements based on memory may be directed toward space beyond an animal’s current field of per- ception (Gautestad and Mysterud 2005; Moorcroft 2012; Fagan et al. 2013). Thus, to distinguish memory- based movements from movements elicited by the im- mediate external environment requires experimental manipulation (Moorcroft 2012). However, the spatial memory that animals possess before experimentation begins is unknown to researchers. Fagen et al. (2013)
suggest one solution to this problem is tracking the spatial dynamics of juveniles throughout ontogeny to obtain complete movement histories before experimen- tation. They further suggest translocating these animals to novel environments, which may help identify move- ments influenced by previous spatial memories.
The movements of adult female White-tailed Deer (Odocoileus virginianus) and their fawns in northeastern Minnesota indicated that spatial memories developed by fawns following their mothers was a primary influence on their adult home-range locations, seasonal migra- tions, and landscape distribution (Nelson and Mech 1981, 1987, 1999, 2006; Nelson 1994, 1998). Further evidence of spatial memory was inferred from a pilot study in which adult females and fawns were translo- cated to unfamiliar areas (Nelson 1994). Some deer returned to their home ranges and resumed their pre- vious movement patterns, while others did not return, but mimicked their pre-translocation movements near their release sites.
Traditional movements and home-range affinity not only suggest the capacity for spatial memory, but also imply a fitness advantage of the propensity to occupy familiar space. To collect experimental evidence of this capacity and propensity, | delineated the home ranges
2 THE CANADIAN FIELD-NATURALIST
of adult does for at least a year before translocating them outside their home ranges. I then examined their movements relative to their home ranges and to the movements of does not translocated. If spatial memory was a major factor responsible for home range affinity, then translocated does would be pre- dicted to return to their home ranges and not remain at their release sites or other sites they encountered that supported resident deer. A corollary of this pre- diction is that translocated does would employ differ- ent modes of movement than those of control does not translocated.
Study Area
I conducted this study in the Superior National For- est in northeastern Minnesota (48°N, 91°W). The top- ography was flat, and the area was dominated by lakes and mixed coniferous—deciduous forests (Heinselman 1996). Average monthly minimum temperatures ranged from 2°C to 18°C from May to October and —18°C to 7°C from November to April (Heinselman 1996). Snow cover generally occurred from November through April with weekly depths averaging 31—64 cm during Feb- ruary and March and 0-30 cm during April (Nelson and Mech 2006).
Most deer in the study area migrated from one of two winter concentration areas, roughly 30 km? each, and averaged 12-km and 25-km migrations to reach individual summer ranges, some up to 80 km distant (Nelson and Mech 1987). Deer migrated to summer ranges during late March and early April when summer range density was 1-3 deer/km? (Lenarz 2002). Partu- rition occurred primarily during the last week of May and first week of June (Kunkel and Mech 1994). Deer occupied 0.7—1.0 km? summer ranges (Nelson and Mech 1981) before returning to the winter concentra- tions during November to January, where densities were over 15 deer/km? (Nelson and Mech 1987). Gray Wolf (Canis lupus) predation and human hunting were the main causes of deer mortality (Nelson and Mech 1986). Wolves along with Black Bears (Ursus ameri- canus) were major predators of newborn fawns during their first weeks of life (Kunkel and Mech 1994).
Methods
I captured adult does during February-April when they occupied winter concentration areas (Nelson and Mech 1981). I anesthetized them (Kreeger 1996), ex- tracted an incisor for aging (Nelson 2001), and fitted them with very high frequency (VHF) radio collars or Global Positioning System (GPS) collars (Merrill e al. 1998). I radio-tracked VHF-collared does from the air two to four times weekly. GPS-collared does yielded locations every hour, which I downloaded to a spread- sheet after remotely releasing the collars (Mech and Gese 1992) or retrieving them after mortality occurred.
I subsequently recaptured, recollared, and translo- cated while sedated, those adult does radio-tracked a
Vol. 129
minimum of | year. I translocated them 10-25 km out- side their home ranges to unfamiliar (no previous radio- locations) and familiar (previous radio locations) re- lease sites. I separated the two groups when describing and analyzing their movements. I recorded the num- ber of days they used to return to their home ranges, or if not returning, the number of days radio-tracked until they died or their radio-collars expired.
I used data from both VHF- and GPS-collared does to measure propensity to return to home ranges after being translocated, but only data from GPS-collared does to describe and quantify movements. To provide experimental controls for comparing with translocat- ed GPS-collared does rearing fawns, I captured and released does at their capture sites wearing GPS col- lars programmed to record hourly locations starting 16 May, | week before parturition and continuing through the first 3 weeks of fawn rearing to 23 June.
I measured area used by GPS-collared does by cal- culating minimum convex polygons (MCP) of their locations (Mohr 1947). I differentiated two modes of movement: directional travel and all other movement based on rate (m/h) measured by distance between hourly locations. I identified and defined travel based on migrating GPS-collared deer traveling 1.5 km/h (SD = 0.6, n = 27, Nelson et al. 2004). Because 95% of their hourlymigration travel exceeded 300 m/h and was sustained for 3—6 h per travel periods (Nelson ef al. 2004), I classified as “travel” in this study, direc- tional movement of > 300 m during each of at least 3 sequential hours. These criteria separated directional travel from all other movements, including those that slowed, circled, or deviated from directional movement.
I contrasted movement behaviours and MCPs of translocated GPS-collared does with those of control GPS-collared does and further compared their hourly movement rates during each week starting 16-23 May and during the first 3 weeks of the fawn-rearing period. I assumed the timing of parturition based on a pattern of spatially constricted locations by parturient does (Kunkel and Mech 1994). This sampling corresponded to the following biological ontogeny: a period before fawns are born; the first week of fawn rearing, which requires maternal care and defensive behaviour by the doe and suckling and hiding behaviour of fawns; the second week of fawn rearing, when the transition from hiding to running begins as a response by fawns to dan- ger (Jackson et al. 1972); the third week of fawn rear- ing, when fawns generally run from danger.
I analyzed hourly movement rates by estimating means and 95% confidence limits (Cherry 1998; Ander- son et al. 2001; Johnson 2002). I further compared hourly movement rates of translocated does with those of control does during each week sampled in the pre- fawn and fawn-rearing periods, by using t-tests and accepting statistical significance at P < 0.05 and when 95% CLs on the mean differences did not include zero.
2015
I followed the American Society of Mammalogists’ guidelines (Sikes and Gannon 2011) and the Animal Care and Use Committee study plan 2700202, Patux- ent Wildlife Research Center, United States Fish and Wildlife Service.
Results
I captured and radio-collared 26, 1-13 year-old does (median = 6 years old) during February-April 1995— 2007. Of the 26 does, I radio-tracked 17 for 1-4 years (median = 2 years) before recapturing and translocat- ing them 10-25 km (median = 13 km) outside their current home ranges. | captured two of them a third time and translocated them to familiar sites that they previously occupied but located outside their current home ranges. I released the remaining nine does at their capture sites to serve as experimental controls for hourly movement comparisons to translocated does during the fawn rearing period.
Radio-tracking of translocated does yielded 2—111 (median = 46) locations from 12 VHF-collared does and 709-4150 (median = 2605) locations from seven GPS-collared does, acquiring 58-95% (median = 84%) of potential locations. Nine control GPS-collared does yielded 341-655 (median = 453) locations, 36-70% (median = 50%) of potential locations.
Return to home ranges
Twelve of 17 (71%) does that were translocated to unfamiliar sites returned to their home ranges. Ten re- turned within 1—89 days (median = 22 days) and the other two returned 1.3 and 3.2 years later. The remain- ing five does failed to return to home ranges as three died and the collars on two others expired (80-275 days, median 174 days). The two does translocated to familiar sites outside their current home ranges also returned to their home ranges.
GPS-collared does released at unfamiliar sites
From March through September 2004 through 2006, five translocated GPS-collared does released at unfa- miliar sites traveled directionally, circled, backtracked, and returned to or toward their release sites (Table 1, Figure 1). Does 8164, 8180, and 8252 roamed exten- sively and the other two (7958 and 8172) traveled directly. Doe 7958 moved directly to her adult summer range to which she had dispersed 3 years earlier as a 1 year old. The bearings and distances for her dispersal movements include the region of her release site, al- though she was never located there. Doe 8172 moved in a direction away from her home range before back- tracking 76% of the distance to her release site (Table 1). Hourly movement rates combined comprised 4% travel and 96% nontravel movement (Table 1). The does moved ten times faster when traveling than at other times (877 m/h vs. 83 m/h, respectively, Table 1).
GPS-collared does released at familiar sites In late March and early April, I translocated GPS- collared does 7904 and 7940 to familiar sites: 7904 to
TABLE |. Rates and patterns of travel and nontravel movement* of translocated GPS-collared adult female White-tailed Deer (Odocoileus virginianus) in northeastern Minnesota,
March-September, 2004-2007.
No. of
Direct travel, km
Nontravel rate Movement pattern
Travel rate
Travel
Doe
locations
a
Ja/olGle n Mean, m/h 9570 Cl n MCP, km
Mean, m/h
period
Released at unfamiliar site
7958
588
NA
NA NA 54
NA
1-30 Mar
NELSON: SPATIAL MEMORY AND Home RANGE AFFINITY IN WHITE-TAILED DEER
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tional movement.
Note: CL = confidence limits, MCP
4 THE CANADIAN FIELD-NATURALIST
5310000 —= =
Vol. 129
27 MAY | oy §305000 ae fe e PARTURITION ¢ 28 MAY-29 JUN % E 5300000 }— ——— = a =— sk | & = | 4 t | S$ : 3 = 5 5295000 - __-4+__—_ : ; oe? o¢ HOME RANGE 4 & 7 JUL-6 SEP ¢ caine 5290000 | — = At S — _ —— RELEASED 2 APR 5285000 — = = a 590000 595000 600000 605000 610000 615000 620000
UTM easting, m
Ficure |. The 3510 hourly locations recorded for GPS-collared White-tailed Deer (Odocoileus virginianus) doe 8252 from 2 April, when she was released at an unfamiliar site 10 km from her home range, to 6 September 2006 in northeastern Minnesota. Arrows depict her general direction of travel starting at the release site and arriving at her home range on
7 July.
a previous release site that she had used 7 years earlier and 7940 to her natal home range from which she had dispersed 5 years earlier as a | year old.
Doe 7904 traveled extensively, circling and back- tracking for 19 days before reaching her winter home range (Table 1). She remained there 15 h before migrat- ing directly to her summer range. Doe 7940 traveled directly for 22 km, including 4 km of backtracking to reach her winter range in 8 days (Figure 2). She re- mained there 8 days before migrating 11 km to her summer range. Their combined hourly movement rates comprised 7% travel and 93% nontravel movement (Table 1). They moved ten times faster when traveling than at other times (909 m/h vs. 85 m/h, Table 1).
GPS-collared does during fawn rearing
Translocated GPS-collared does 8164, 8172, 8180, and 8252 stopped their extensive travel in mid-May before parturition, which began for them and nine con- trol GPS-collared does between 25 May and 6 June (median = 31 May). During 12—28 (median = 23) se- quential days of fawn rearing, both groups occupied 0.2—1.3 km? (median = 0.5 km?).
Translocated and control does moved at the same mean hourly rates during 7 days in mid-May before fawn rearing and also during the first week of fawn rearing (Table 2). Control does moved faster than trans- located does during the second week of fawn rearing, but not in the third week (Table 2). Translocated does permanently departed their fawn rearing areas after
TABLE 2. Rate of movement of translocated and control GPS-collared adult female White-tailed Deer (Odocoileus virginianus) in northeastern Minnesota a week before parturition (16-23 May 2001-2006) and during three weeks after parturition while rear-
ing fawns. Period Translocated does Control does
Mean, m/h 95% CL n Mean, m/h 95% CL n Preparturition (1 week) 79 9 586 88 8 639 Fawn rearing First week 66 6 524 74 6 707 Second week 83* 9 475 109* 9 598 Third week 97 13 356 105 8 643
Note: CL = confidence limits.
*Significant difference between translocated and control does (p < 0.005, f test).
2 te ‘ 015 NELSON: SPATIAL MEMORY AND HOME RANGE AFFINITY IN WHITE-TAILED DEER 5 5300000 NATAL HOME RANGE 1999 5295000 ® RELEASED 1 APR 2004 y 5290000 + * ‘te 5285000 = “tes 8-9 APR = 18H ob @ s - gS 5280000 + 9-17 APR ADULT WINTER RANGE = | % 1999-2004 a | | | ay, 5275000 + & or 5270000 + —_——_— - 18 APR - 18 MAY S 5265000 +- ——————— : — a 590000 595000 600000 605000 610000 615000 620000
UTM easting, m
FiGURE 2. The 710 hourly locations, recorded between | April and 18 May 2004, for GPS-collared White-tailed Deer (Odocoileus virginianus) doe 7940 captured on her winter range and translocated 22 km to a familiar site (her natal home range) 5 years after natal dispersal. Arrows depict her general direction of travel starting at her release site, traveling 22 km, arriving on her adult winter home range on 9 April, subsequently traveling 11 km further to her adult summer home
range, and arriving on 18 April.
12-45 days (median = 26 days) and continued travel- ing, while control does remained on their home ranges.
Discussion
All translocated does left their release sites, which had resident deer present, suggesting that the resources for deer survival were present. Thus, lack of habitat appeared an unlikely factor influencing their departure. Aggression by resident deer toward the translocated does can also be excluded as a factor, as elsewhere in the study area, sympatric wintering deer moved inde- pendently of each other, suggesting that competition for space was not influencing their movements (Nel- son and Sargeant 2008). Similarly, Jones et al. (1997) observed no effect on movement of resident deer from the presence of translocated deer. Although conflict is observed at artificial feeding sites, which attract large numbers of deer, such disputes appear restricted to feed- ing behaviour (Ozoga 1972).
There was large variation in the amount of time tak- en to return to home ranges despite the fact that 87% of the does were translocated similar distances (13-15 km). I previously found that female yearlings made 7-22-km forays beyond their natal ranges (Nelson 1998), and some dispersed 18-168 km to new home ranges (Nelson 1993). Thus, spatial memories estab-
lished during exploratory or dispersal movements could have been one factor influencing variation in return time over similar distances. Conceivably, some does encountered areas they recognized from previous ex- ploratory or dispersal movements and then navigated accordingly to return to familiar space. Others not en- countering familiar areas would necessarily spend more time roaming if spatial memory was the primary mech- anism they used as they attempted to return to home ranges.
The movements of four of five GPS-collared does translocated to unfamiliar release sites suggest that they were looking for familiar space. Their extensive travel far exceeded that necessary to acquire the re- sources for daily survival evinced by adult does in the study area that were occupying home ranges < 1% the size of spaces traveled by translocated does (Nelson and Mech 1981). Their movements further suggest that translocated does simultaneously developed new spa- tial memories, evidenced by backtracking and return- ing to release sites.
The direct travel to her home range by the fifth GPS- collared doe suggests that the release site was part of the spatial memory she had developed during natal dispersal movements in the region of the release site as she established her adult home range. Direct travel
6 THE CANADIAN FIELD-NATURALIST
by the two GPS-collared does released on familiar sites after being absent from them 5 and 7 years also indi- cates that they recognized their surroundings, although only one traveled like deer migrating annually between summer and winter home ranges (Nelson et al. 2004). The additional roaming, circling, and backtracking of the other doe hints at variation in the longevity of spa- tial memory and highlights the challenge of understand- ing and including it in models of animal movement.
The movements of the one doe translocated to an unfamiliar release site, traveling directly and backtrack- ing toward her release site, contrasts sharply with her roaming cohorts. Given the small sample size and the fact that she was nonmigratory before being translo- cated, it is difficult to interpret her movements. She may have been behaving similar to three translocated deer that appeared to invoke memory of their previous migration pattern (Nelson 1994). Migrating Siberian roe deer (Capreolus pygarus), captured and translo- cated to unfamiliar areas while migrating, mimicked the same migration direction and distances as their cohorts (Danilkin et al. 1994). These examples sug- gest that at least two cognitive processes operate to achieve spatial orientation: one depending on memory of bearing and distance patterns to direct movement, and another involving roaming and searching for famil- iar space to determine direction of travel.
When translocated GPS-collared does stopped trav- eling just before parturition, they moved at the same hourly rates as control GPS-collared does and, subse- quently, constricted their movements, as did control does, indicating that both groups gave birth and cared for their fawns (Nelson and Mech 1981; Ozoga et al. 1982; Kunkel and Mech 1994). The similar movement rates of both groups during the first week of fawn rear- ing suggest that translocated does may not have experi- enced any negative behavioural or physiological effects from their extensive travel before parturition. The in- creased rate of movement by control does, compared with translocated does, in the second week of fawn rais- ing suggests possible differences as fawns matured. However, in the subsequent week, both groups moved at the same rate indicating an overall pattern of simi- lar movement rates. before and while rearing fawns.
The duration of restricted movement of does after giving birth further indicates that they nurtured their fawns beyond the period of fawn concealment and in- activity, well into the period when fawns regularly fol- low their mothers (Jackson et al. 1972). This further indicates adequacy of nutrition for maintenance as well as that needed to sustain lactation. I do not know if the fawns survived and followed their mothers when they continued roaming. Regardless, the relevant and over- riding result is that the biological imperative of par- turition and fawn rearing took temporary precedence over travel for the translocated does. This is a clear example of changes in movements determined by op- posing internal processes: one directing parturition and
Vol. 129
nurturing behaviour and the other emanating from the capacity for spatial cognition and the propensity to occupy familiar space.
All translocated does left their release sites, and those surviving with transmitting radio-collars returned to their home ranges. The translocated GPS-collared does did this despite having met their nutritional and physiological demands while traveling through hun- dreds of square kilometres, three orders of magnitude larger than home ranges of adult does. Thus they moved continually beyond daily field of perception to eventu- ally arrive at the exact site they occupied before being translocated. This clearly demonstrates not only the capacity for spatial memory, but also the propensity to return to familiar space. The extremely aberrant roam- ing of the GPS-collared does can only be understood as searching the landscape for space that was remembered and recognized when finally found.
The dominant paradigm of ungulate movement is based on the premise that movement is directed by in- nate optimal foraging in the field of perception. Math- ematical modeling has accepted this premise, ignor- ing behavioural mechanisms, such as the influence of spatial memory (Gautestad and Mysterud 2005). How- ever, it has become increasingly clear that spatial mem- ory plays an integral role in animal movement and must be included in models to achieve biological reality in predicting movements. Fagan et al. (2013) recognized this as part of an “emerging research interface” of be- havioural ecology, cognitive science, animal tracking, and quantitative ecology. The findings herein contri- bute to their call for experimental evidence of spatial memory and further elucidate its pervasive role in the spatial ecology of White-tailed Deer.
Acknowledgements
This research was supported by the United States Geological Survey, United States Department of Agri- culture North Central Research Station, and Special Projects Foundation funding to L. D. Mech. I thank L. D. Mech for his support of the study and review of the manuscript. I also thank associate editor Garth Mowat and two anonymous reviewers for comments and suggestions for improving the manuscript. I thank 160 volunteer wildlife technicians who assisted in live capture of deer and who retrieved collars. I also thank the pilots of the U.S. Forest Service for skillful and safe flying.
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Received 16 January 2014 Accepted 18 June 2014
Whitebark Pine (Pinus albicaulis) Seeds as Food for Bears (Ursus spp.) in Banff National Park, Alberta
Davip HAmer!:3 and IAN PENGELLY?
'300-379 Spring Creek Drive, Canmore, Alberta Tl W 0G8 Canada 2235 Lady MacDonald Drive, Canmore, Alberta T1W 1H2 Canada 3Corresponding author: j.david.hamer@gmail.com
Hamer, David, and Ian Pengelly. 2015. Whitebark Pine (Pinus albicaulis) seeds as food for bears (Ursus spp.) in Banff National Park, Alberta. Canadian Field-Naturalist 129(1): 8-14.
The large, nutrient-rich seeds of Whitebark Pine (Pinus albicaulis Engelmann) are important food for bears (Ursus spp.) in Yellowstone National Park. In Banff National Park, studies have shown that American Black Bears (Ursus americanus) eat these seeds, but little additional information is available. We studied Whitebark Pine in Banff National Park to address this information gap. Because bears obtain Whitebark Pine seeds from Red Squirrel (Jamiasciurus hudsonicus) middens, our objective was to measure the abundance, habitat characteristics, and use by bears of middens in Whitebark Pine forests. A second objective was to determine whether Grizzly Bears (U. arctos) in Banff National Park also eat Whitebark Pine seeds. In 2011-2012, we ran 29 ha of 20-50 m wide transects at 10 sites with accessible Whitebark Pine stands and found 0-3.7 middens/ha (mean 1.23, SD 1.17, n= 10). Midden density was weakly related to total basal area of all conifers but not to basal area of Whitebark Pine. Mid- dens were located in the upper subalpine at an average elevation of 2110 m (SD 90, 1 = 8) on 41-248° facing slopes with a mean steepness of 28° (SD 5, n = 8). Bears had excavated middens at all eight sites where we found middens; at the remaining two sites, middens did not occur within our transects. Overall, 24 (67%) of the 36 middens located in our transects had been dug by bears. In October 2013, we searched areas where three global positioning system (GPS)-collared Grizzly Bears had been located in late September 2013 and found five recently dug middens located less than 6 m from GPS fixes. These observations are, to our knowledge, the first conclusive evidence that Grizzly Bears in Banff National Park eat Whitebark Pine seeds. Because Whitebark Pine occurs at high elevations on steep slopes where human use is low, this resource may be important in keeping bears in habitat where risk of human-caused mortality is lower. Our results may assist managers responsible for conservation of bears in Banff National Park, where both American Black Bears and Grizzly Bears are subject to high levels of human-caused mortality.
Key Words: American Black Bear; Banff National Park; Grizzly Bear; midden; Pinus albicaulis; Red Squirrel; Ursus americanus; Ursus arctos; Tamiasciurus hudsonicus; Whitebark Pine; seeds
Introduction
The large, nutrient-rich seeds of Whitebark Pine (Pinus albicaulis Engelmann) are a major food for Grizzly Bears (Ursus arctos) and American Black Bears (U. americanus) in the Greater Yellowstone Eco- system (GYE; Fortin et al. 2013). As stated by Mattson and Reinhart (1997:926): “When whitebark pine seeds are abundant, grizzly bears [in the GYE] eat virtually nothing else.” Mattson ef al. (1992) found that in years of high seed availability, GYE Grizzly Bears were half as likely to use areas within 5 km of roads or within 8 km of other developments because Whitebark Pine’s high elevation distribution typically is remote from human facilities. In contrast, in years of small White- bark Pine seed crops, mortality of adult female Grizzly Bears averaged 2.3 times higher, and mortality of sub- adult males averaged 3.3 times higher than in years of large seed crops, which the authors attributed to the ten- dency of bears to range closer to human facilities in years of pine seed scarcity.
Use of Whitebark Pine seeds by bears in Canada has not been clearly described. Whitebark Pine seeds were recorded in the diet of American Black Bears in Banff National Park (Kansas eft al. 1989; Raine and Kansas
1990), although information on habitat use was limited to the general observation that American Black Bears feeding on Common Juniper (Juniperus communis L.) cones, Common Bearberry (Arctostaphylos uva-ursi [L.] Sprengel) fruits, and Whitebark Pine seeds fre- quented higher elevations in moderate to steeply slop- ing, south-facing, sub-xeric pine forests. Seeds of White- bark Pine were also eaten by a radio-collared Grizzly Bear in Yoho National Park in | year of a 3-year study along the Continental Divide immediately west of Banff National Park (Raine and Riddell 1991). No habitat information was provided other than that this feeding occurred on high-elevation slopes.
McLellan and Hovey (1995) noted that Whitebark Pines were common in their southeast British Columbia study area, but they observed only one case of Grizzly Bears apparently feeding on seeds, and Whitebark Pine seeds did not occur in their sample of scats. In south- western Alberta, Hamer ef al. (1991) did not record Grizzly Bear use of Whitebark Pine seeds in Waterton Lakes National Park. In central Alberta, Whitebark Pine seeds were not recorded in Grizzly Bear food-habit studies conducted in Jasper National Park, Banff Na- tional Park, or the Jasper—Edson area (Russell et al.
2015
1979; Hamer and Herrero 1987; Munro et al. 2006), although the Banff study area was northeast of the zone of Whitebark Pine abundance in the park. A Grizzly Bear study (Wielgus 1986) and an American Black Bear study (Holeroft and Herrero 1991) in Kananaskis, immediately east of Banff National Park, also did not record bear use of Whitebark Pine seeds.
We studied Whitebark Pine in and near Banff Nation- al Park during 2011-2013 to address the lack of spe- cific research regarding use of pine seeds by bears in this area. Because Mattson and Reinhart (1997) found that all Whitebark Pine seeds eaten by bears in the GYE were obtained from Red Squirrel (Jamiasciurus hudsonicus) middens, our principal objectives were to record the abundance, habitat characteristics, and evi- dence of use by bears of Red Squirrel middens in White- bark Pine forests. Because field evidence of bear use of middens is not species specific, we also checked areas where radio-collared Grizzly Bears had been lo- cated to address the specific question: do Grizzly Bears in Banff National Park eat Whitebark Pine seeds?
Study Area
Banff National Park occupies 6641 km? in the cen- tral Rocky Mountains of Alberta, Canada. The park ex- tends eastward from the Continental Divide to encom- pass mountainous habitat of both the Main Ranges and the more easterly, more arid Front Ranges. Elevation in Banff National Park ranges from 1330 m to 3610 m with the tree line at roughly 2300 m. The subalpine zone is at approximately 1500-2350 m, with the upper subalpine (generally cooler and wetter, with deeper and longer lasting snow) beginning at about 2000 m (Achuff 1982). Our study sites were between 1900 m and 2300 m where Whitebark Pine occurs.
We worked in Whitebark Pine stands that exhibited little mortality from White-Pine Blister Rust (Cronar- tium ribicola) or Mountain Pine Beetle (Dendroctonus ponderosae). Banff National Park currently has a low rate of White-Pine Blister Rust infection compared with locations north and south along the Rocky Mountains (Smith et al. 2008, 2013). Because we located study sites in Whitebark Pine stands, Whitebark Pine was often co-dominant in our plots. Based on basal areas recorded during our analyses, the relative abundance of Whitebark Pine was 38%, Interior Spruce (Picea engel- mannii var. engelmannii * P. glauca) 32%, Subalpine Fir (Abies lasiocarpa [Hooker] Nuttall) 19%, Lodge- pole Pine (Pinus contorta Douglas ex Loudon) 11%, and Subalpine Larch (Larix lyallii Parlatore) 1%. The understory included submesic Soapberry (Shepherdia canadensis [L.] Nuttall)-Common Juniper-Common Bearberry communities; mesic Grouseberry (Vaccinium scoparium Leiberg ex Coville) and Grouseberry—Soap- berry communities; and subhygric Subalpine Fir (sap- lings)-False Azalea (Menziesia ferruginea J E. Smith syn. M. glabella)-feathermoss (e.g., Hvlocomium splen-
HAMER AND PENGELLY: WHITEBARK PINE SEEDS AS FOOD FOR BEARS 9)
dens, Pleurozium schreberi) communities (Corns and Achuff 1982).
Methods Transects
We established belt transects at 10 sites in Whitebark Pine forests to measure the density, habitat character- istics, and use by bears of Red Squirrel middens (i.e., locations where squirrels cache large numbers of con- ifer cones, shred these cones to obtain seeds, and thus create conspicuous deposits of organic material). We identified Whitebark Pine stands from aerial survey (1. Pengelly and A. Buckingham, Parks Canada, unpub- lished data), knowledge of Whitebark Pine stands in the park, and reconnaissance from roads and trails. At 71% of our surveyed areas, Whitebark Pine basal area was greater than 4 m’/ha. We established six transect sites in the main Bow Valley, two in the North Saskat- chewan watershed at the north end of the park, and two 0.3 and 2.4 km west of Banff National Park in Kootenay and Yoho national parks, respectively. The average distance between transect sites was 57 km (range 0.2-157 km). Transect sites were 0.4-3.8 km from road access. Fieldwork was conducted from 7 September to 2 October 2011, except at one partly sur- veyed site where we completed work in 2012.
We used a transect width that allowed the enclosed area to be accurately surveyed without excessive cours- ing up and down slope. Most transects (51% of hectares surveyed) were 30 m wide and conducted by two peo- ple. We also ran 20-, 40-, and 50-m wide transects with one, three, and four people, respectively. We ran tran- sects on the elevational contour of the start point, with one person maintaining this elevation so that middens near a transect edge could be accurately placed inside or outside of the transect.
Transects were 114-607 m long and ended either at a natural feature, such as an avalanche slope or rock talus, or after a preselected distance, commonly 200 or 400 m. Transect length was measured with a hand-held global positioning system (GPS) unit. At the end point, a new transect was typically run in the reverse direction, starting at a preselected distance up or down slope.
Only middens whose centres were inside a transect were recorded. A midden centre was defined as the cen- tre of the “midden tree” (for those formed around the base of a large-diameter tree) when this was unambigu- ous: otherwise, it was the intersection of the axes of midden length and width. Only middens with conifer- cone debris more than 20 cm deep and covering more than 10 m2 (> 6 m? if depth > 30 cm) were included. These criteria were used to exclude the numerous smaller deposits of cones and cone debris across the landscape that result from squirrel feeding activity. We also defined a secondary (diffuse) midden (Gurnell 1984) as a smaller midden (but meeting our criteria of minimum depth and area) whose centre was also with-
10 THE CANADIAN FIELD-NATURALIST Vol.
in the transect and that was less than 25 m from a larger midden (average distance 13.8 m). These secondary middens were assumed to be part of the resident squir- rel’s caching and feeding activity (Gurnell 1984) and, hence, were not analyzed separately to avoid pseu- doreplication.
At each midden, we measured midden length (the longest axis of the midden) and width (the greatest dimension at right angles to the long axis) and multi- plied these numbers to obtain midden area (Mattson and Reinhart 1997). We recorded location and eleva- tion using a hand-held GPS unit, slope aspect and steepness using a compass with built-in clinometer, and conifer basal area using a 2 m*/ha prism. Visible White- bark Pine cones were counted, but we did not disturb middens to tally buried cones. Excavated middens and middens with nearby bear fecal deposits (scats) con- taining pine seeds were recorded as used by bears.
We also recorded site characteristics at systematic points along transects (null plots). These null plots were placed 160 m from the last midden or transect starting point when no midden occurred within 200 m. We measured distances with a GPS unit and used these distances to locate null plot centres without bias.
GPS-collared Grizzly Bears
During late October 2013, we searched areas where three GPS-collared Grizzly Bears had been located. These bears were collared by Parks Canada for anoth- er study and generated GPS locational fixes every 20 minutes to 4 h. We selected a small subset of fixes from those obtained during 7-29 September 2013 in Banff National Park, 1.9—9.0 km from vehicle access, and in upper subalpine areas where Whitebark Pines are found. Google Maps (satellite view) was used as a layer
[29
in the geographic information system, QGIS (open- source software, version 2.0.1), to exclude fixes in non- forested habitat. Fixes were searched for signs of bear activity. Red Squirrel middens, if present near the fix, were examined in the same way as those located on our transects.
Data analysis
Sites were our sampling units. For each transect site, we assessed the relation between midden density and conifer basal area and between midden density and the proportion of Whitebark Pines using the linear model in R (open-source software, version 3.0.2). Because of our small sample size, we present differences in character- istics among middens obtained by transecting, middens located at GPS sites, and plots located at null sites visu- ally using box-and-whisker diagrams in R. Secondary middens were excluded from all analyses except for calculation of total middens per hectare.
Results Middens located by transecting
The mean density of Red Squirrel middens in our 10 transect sites was 1.23 middens/ha (SD 1.17, Table 1) and 1.81 middens/ha (SD 1.84) if secondary middens were included. Mean midden size was 97 m? (SD 64, range 30-218 m’, n = 8).
All middens contained Whitebark Pine cone scales, but we found few cached Whitebark Pine cones in 2011 compared with the hundreds we found in several mid- dens during a 1|-day pilot project in 2010. In 2011, the three largest caches of Whitebark Pine cones held 195, 67, and 7 cones. No cached Whitebark Pine cones were found in the three middens located at the site completed in 2012.
TABLE |. Results of transect survey of Red Squirrel (Tamiasciurus hudsonicus) middens conducted in Whitebark Pine (Pinus albicaulis) stands in and adjacent to Banff National Park, Alberta, 2011-2012.
RRC TTT TT
Mean Mean basal Mean Mean slope Mean area of basal area Mean Area Slope steepness elevation Whitebark ofother Midden midden Evidence Transect surveyed No. aspect* a Pine* conifers* density area of use site (ha) — middens @) G) (m) (m?/ha) (m*/ha) (no./ha) (m2) _ by bears A AalT, — 2 4] 20 2000 7.0 26.0 0.48 137 Dug, scat B 3.01 3 240 31 2150 6.0 32.0 1.00 96 Dug, scat Cc 2.52 0 265 31 2020 9.0 20.0 0 = D 2.06 4 189 33 2210 27.0 Teds 1.95 39 ~=Dug E 2.99 ul 176 30 2210 11.6 24.7 3.68 51 Dug, scat F 4.43 6 190 30 2100 10.3 S21) 1.36 68 Dug G 1.84 0 240 33 1990 6.7 8.0 0 Soe H 2.44 6 248 2D 1960 22.0 27.0 2.46 218 Dug, scat I 2205) | 230 34 2150 10.0 24.0 0.49 135 Dug J 3.61 3 185 23 2070 18. Total 29.12 36 il = = ae sill kG Ad lia: Mean — 200 29 2090 12.8 2 hs} ——— SD -— 64 5) 90 DP OF) Ihe 64
Oe SSS——SSSssSssssssSsSs>—gg.
At null plots if no middens occurred at that transect site.
2015
"| a
Midden Density (middens/ha)
HAMER AND PENGELLY: WHITEBARK PINE SEEDS AS FOOD FOR BEARS 1]
30 40 50
Basal area of all conifers (m*/ha)
FIGURE l. Relation between density of Red Squirrel (Zamiasciurus hudsonicus) middens and conifer basal area in 10 Whitebark Pine (Pinus albicaulis) belt transect sites in and adjacent to Banff National Park, Alberta, 2011— 2012.
We found a weak positive relation between midden density and forest basal area, with total basal area of conifers explaining 27% of the variation in the data (adjusted ° = 0.272; F = 4.36; 1, 8 df; P = 0.07; Figure 1). We did not find a relation between midden density and basal area of either Whitebark Pines (P = 0.2) or other species of conifer (P = 0.3) or between midden density and the proportion of total conifer basal area accounted for by Whitebark Pine (P = 0.7).
Middens had been excavated by bears at eight of the 10 transect sites; at the two remaining sites, we did not record any middens within the transects (Table 1). Overall, 24 (67%) of the 36 middens found in our transect sites had been dug by bears. Eight (33%) of the 24 dug middens had been excavated recently (i.e., late summer or early autumn 2011). We also found bear scats containing Whitebark Pine seeds at five of the 10 transect sites (Table 1).
Null plots had similar elevation, slope aspect, slope steepness, and basal area of Interior Spruce compared with midden plots (Figure 2a, b, c, and g). However, the basal areas of all conifers, Whitebark Pine, and Sub- alpine Fir were, respectively, about 1.3, 1.5, and 2.0 times greater at middens than at null plots (Figure Ref, and h). Middens tended to occur on less-steep slopes (mean 27.7° [SD 5.5, n = 8] than null plots (mean 31.7° [SD 3.4, n= 8)).
Use of middens by GPS-collared Grizzly Bears
We located recently dug middens at three GPS fix- es of an adult female Grizzly Bear and at two fixes of a subadult male Grizzly Bear. All five dug middens
were less than 6 m from GPS fixes and, thus, were linked to Grizzly Bear activity. Bear scats containing Whitebark Pine seeds occurred at three of these GPS sites. The central axes of Whitebark Pine cones, some with attached cone scales, numbered > 100, > 100, 30, 1, and 0 at the five sites. The midden with no identified cone axes contained Whitebark Pine cone scales.
We searched more than 15 satellite fixes from a third GPS-collared Grizzly Bear, at four locations occurring over 8 km linear distance in the park. We did not find Whitebark Pine feeding signs or Whitebark Pine stands at or near any of the fixes from this adult female.
The five middens found at fixes of GPS-collared Grizzly Bears had habitat characteristics notably sim- ilar to those of the middens we recorded from tran- sects, including mean elevation (2150 m vs. 2110 m), aspect (210° vs. 190°), and slope steepness (29° vs. 28°). Mean midden size (94 m2 vs. 97 m2) and total conifer basal area (35 m2/ha vs. 37 m*/ha) were also similar. However, mean Whitebark Pine basal area was 7 m?/ha at GPS-located middens but 14 m?/ha at transect mid- dens. Contributing to this difference was one GPS- located midden on a bench with a 16° slope, with no Whitebark Pines at the site (although Whitebark Pine trees were abundant on a steep, 37° slope 35 m away). These comparisons are displayed non-parametrically (i.e., using medians and quartiles) in box-and-whisker diagrams (Figure 2). Our GPS sample is small, but the notable overall similarity between the GPS data and the transect data supports the validity of our midden sampling by transect.
12 THE CANADIAN FIELD-NATURALIST Vol. 129
Elevation (m) Slope Aspect (degrees)
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FIGURE 2. Comparison of habitat characteristics at 36 Red Squirrel (Zamiasciurus hudsonicus) middens located in eight belt transect sites. five middens within 6 m of satellite fixes of two GPS-collared Grizzly Bears (Ursus arctos), and 27 null plots established systematically at 200-m intervals when no midden occurred within a transect for that 200-m distance (8 sites) in Whitebark Pine (Pinus albicaulis) stands in and adjacent to Banff National Park, Alberta, 2011-2013. (a) elevation: (b) slope aspect; (c) slope steepness; (d) ratio of White- bark Pine basal area to total conifer basal area; and basal area of; (e) all conifers; (f) Whitebark Pine: (g) Interior Spruce (Picea engel- mannii * P, glauca); and (h) Subalpine Fir (Abies lasiocarpa). Diagrams show median (band inside box), first and third quar-
tiles (top and bottom of box), 1.5 times the interquartile range (ends of whiskers): and outliers beyond the 1.5 interquartile range limits (circles).
2015
Discussion
Our observations of five recently dug Red Squirrel middens within 6 m of fixes obtained from two GPS- collared Grizzly Bears during September 2013, plus the associated scats containing Whitebark Pine seeds, are, to our knowledge, the first conclusive evidence that Grizzly Bears in Banff National Park eat Whitebark Pine seeds.
We found that 67% of the middens located by tran- secting had been excavated by bears. All eight transect sites where middens were recorded contained excavat- ed middens; however, we did not identify the species of bear involved in these excavations. In 2011, when we ran most transects, middens contained few cached Whitebark Pine cones, and we did not find recently deposited bear scats containing Whitebark Pine seeds as required for DNA sampling. Hence, we were unable to differentiate American Black Bear use from Grizzly Bear use in our transect sites. These results contrast with our |-day pilot project in 2010 when we found four fresh scats containing pine seeds within 2 ha at a site where we established transects in 2011.
At our transect sites, Whitebark Pine basal area ranged from 0 to 27 m’/ha, which is higher than the range of 0.2-7 m*/ha reported for a study area in the nearby Willmore Wilderness Park (McKay and Graham 2010). The greater abundance of Whitebark Pine at our transect sites may partly explain why our mean midden density was greater than the 0.46 middens/ha reported by McKay and Graham (2010). Results from the GYE (Mattson and Reinhart 1997) were more comparable to ours, with Whitebark Pine basal areas of 2-23 m*/ha and midden densities of 0.2 to 1.1/ha, although only active middens were tallied in that study. Our midden densities were 0—3.7/ha (Table 1). We did not differen- tiate between active and inactive middens because we judged that such categorization would be subjective and likely unreliable.
The mean basal areas of Whitebark Pine and Sub- alpine Fir were greater at midden plots than at null plots, but Interior Spruce basal area did not differ sub- stantially. Because squirrels often establish middens at the bases of large trees, our midden basal areas may be high compared with those for the stand where they were located.
Null plots tended to occur on steeper slopes (26-36°) than middens (20-34°). Flatter microsites, including small benches interrupting the main slope, were loca- tions for some middens. These microsites appeared to provide for the accumulation of midden material, allow- ing Red Squirrels to store cones in the organic debris. In contrast, on many steep slopes, it appeared that cones and conifer debris would readily disperse downhill from gravity and surface water flow. ;
Whitebark Pine seeds are a valued resource for bears in the GYE (Kendall 1983; Mattson e¢ al. 1991; Fortin et al. 2013). Whitebark Pine cone abundance was the highest-ranked habitat covariate (along with year, sea-
HAMER AND PENGELLY: WHITEBARK PINE SEEDS AS Foop FOR BEARS 13
son, sampling regime, and sex of Grizzly Bear) in six best models that explained Grizzly Bear survival in the GYE for 1983-2001 (Schwartz et a/. 2006). Raine and Kansas (1990) identified Whitebark Pine seeds as part of the diet of American Black Bears in Banff National Park, and we have shown that Grizzly Bears in Banff National Park also eat these seeds.
American Black Bears in Banff National Park ap- pear to be in decline because of high human-caused mortality (Hebblewhite e¢ a/. 2003). Grizzly Bears in Banff National Park are at the eastern limit of their range, inhabit one of the most intensively developed landscapes in the world where Grizzly Bears still occur, have the slowest reproductive rate of any Grizzly Bear population yet studied, and also experience high levels of human-caused mortality (Garshelis e¢ a/. 2005). Our study provides managers with information on a poten- tially important, nutrient-rich food that may give some bears the energy necessary for reproduction (Rogers 1976), and that, when abundant, can move bears into remote, steep habitat where risk of human-caused mor- tality is lower.
Acknowledgements
Financial support and GPS fixes were provided by Parks Canada. We are also grateful for the contributions of A. Buckingham, M. Didkowsky, A. Forshner, B. Fyten, S. Herrero, T. Jung, B. Low, J. Park, D. Verhulst, J. Whittington, L. Wiggins, N. Woode, and anonymous reviewers. Publication costs for this article were sup- ported by the Thomas H. Manning Fund of The Ottawa Field-Naturalists’ Club.
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McKay, T., and K. Graham. 2010. Whitebark pine seeds as a food source for grizzly bears in west central Alberta. 2008/2009 pilot study. Pages 30-47 in Foothills Research Institute Grizzly Bear Program, 2009 Annual Report. Edit- ed by G. B. Stenhouse and K. Graham. Hinton, Alberta, Canada. Accessed January 2012. https://foothillsri.ca/sites /default/files/null/GBP_2010 04 AnnRpt_2009.pdf
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McLellan, B. N., and F. W. Hovey. 1995. The diet of grizzly bears in the Flathead River drainage of southeastern British Columbia. Canadian Journal of Zoology 73: 704-712.
Munro, R. H. M., S. E. Nielsen, M. H. Price, G. B. Sten- house, and M. S. Boyce. 2006. Seasonal and diel patterns of grizzly bear diet and activity in west-central Alberta. Journal of Mammalogy 87: 1112-1121.
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Rogers, L. L. 1976. Effects of mast and berry crop failures on survival, growth, and reproductive success of black bears. Transactions of the North American Wildlife and Natural Resources Conference 41: 431438.
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Smith, C. M., B. Wilson, S. Rasheed, R. C. Walker, T. Car- olin, and B. Shepherd. 2008. Whitebark pine and white pine blister rust in the Rocky Mountains of Canada and northern Montana. Canadian Journal of Forestry Research 38: 982-995.
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Received 8 April 2014 Accepted 1 December 2014
Horse Ranching Increases Biodiversity in a Foothills Parkland Prairie in Northern Kananaskis Country, Western Alberta
PAUL M. CATLING!:3, BRENDA KostIUK!, and DON THOMPSON?
Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, William Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada
“Agriculture and Agri-Food Canada, Lethbridge Research Centre, 5403 | Avenue South, P.O. Box 3000, Lethbridge, Alberta T1J 4B1 Canada
‘Corresponding author: catlingp@agr.ge.ca
Catling, Paul M., Brenda Kostiuk, and Don Thompson. 2015. Horse ranching increases biodiversity in a foothills parkland prairie in northern Kananaskis Country, western Alberta. Canadian Field-Naturalist 129(1): 15-23.
Vascular plant biodiversity was evaluated in two adjacent sections of a continuous prairie glade. One section has been sub- ject to moderate grazing by feral horses (Equus ferus caballus) in late summer and fall for the past 25 years, while the other has been protected. From 28 June to 2 July 2009, we recorded cover for all vascular plants present in ten 1-m? quadrats along five transects in each section. We calculated biodiversity measures, including species richness, evenness, and Shannon- Wiener and Simpson’s diversity indexes. Horse grazing did not affect richness but significantly increased evenness. Grazing increased the Shannon-Wiener index, but did not affect the Simpson’s index. Cover and frequency values for most species differed significantly between the two sites. Mountain Rough Fescue (Festuca campestris Rydberg) dominated the non- grazed site but several shorter grasses and different forbs dominated the grazed site. The plant community in the grazed areas can be seen as an earlier seral stage of the fescue community with a different contingent of plant species. Light grazing in part of the prairie glade increased overall plant diversity so that it provided more diverse animal habitat.
Key Words: Horses; Feral Horse; Eguus ferus caballus; Mountain Rough Fescue; Festuca campestris; grazing; rangeland; vascular
) Gee L'CES QuUS grazing, rang cu plants; biodiversity; richness; heterogeneity; eveness; Alberta; prairie foothills; Kananaskis; fescue grassland; shifting mosaic; patch dynamics; management
Introduction Stavely, Alberta (Willms ef al. 1985), but this site may A huge ecological cost is associated with livestock not represent the foothills grasslands well. Only 5% of grazing in western North America, and continuing inter- the grasslands remain in a pre-settlement condition est from conservation biologists is essential to ensure (Vujnovic 1998), and they are now recognized as an en- that management protects biodiversity (Fleischner dangered ecosystem (Trottier 2002). Increased precip- 1994). The extensive literature on maintenance of _ itation in foothills prairies may make them more sus- healthy rangeland for livestock production in western _ ceptible to grazing (Lauenroth ef al. 1994). Canada has led to an understanding that many range- There are currently over 350 000 feral horses (Equus land species may benefit from moderate grazing (e.g., _ ferus caballus) in Alberta, mostly used in recreation, Tannas 2003a.b, 2004). However, the effects of graz- and the number is increasing (Westar 2003*). Conse- ing on biodiversity overall have not been sufficiently quently, the demands for grazing land are also increas- studied in Canada (Ollf and Ritchie 1998; Bai ef a/. _ ing, particularly in the foothills region. Grazing by hors- 2001), although some important research is underway _ es has been shown to reduce plant species richness in (e.g., the long-term east block grazing experiment in some situations (Beever and Brussard 2000; Beever et Grasslands National Park, see http://www.pe.ge.ca/eng _ al. 2008), but not in others (Detling 1998). These differ- /progs/np-pn/re-er/ec-cs/ec-cs01.aspx). Research to date — ences are likely a result of different grazing pressures. has involved mostly cattle on the prairies and parklands With the effects of grazing generally requiring more and suggests that rangelands can be important in pro- study, especially in the foothills, and especially con- tecting biodiversity, although ecological integrity can- _ cerning feral horses, any situation providing data is an not be maintained if grazing pressure is too high (Trot- important study opportunity. We encountered such an tier 1993; West 1993; McLaughlin and Mineau 1995). opportunity in a foothills parkland prairie in northern The grazing ecology of the prairies of the Rocky Kananaskis Country of western Alberta. Here feral hors- Mountain foothills has been poorly studied, although — es had been excluded for 25 years from half ofa contin- they are the most diverse and complex of the fescue —_ uous fescue prairie glade, but rest-rotation grazing at a grasslands in Canada (Tannas 2003a, P.M.C. personal — specific carrying capacity was continued in the adjacent observation) and among the most productive grasslands half. Data was collected to elucidate the effects of graz- in North America (Willms er al. 1996). The response of ing by the horses on floristic diversity, as well as to ob- fescue grasslands to cattle stocking rates has been stud- _ tain information for conservation and management. ied intensively at one site in the Porcupine Hills near
15
16 THE CANADIAN FIELD-NATURALIST
Study Area
The area studied was within the Kananaskis Coun- try in Bow Valley Provincial Park, at 51.0789°N and 115.0384°W. It consisted of a 5-ha prairie glade sep- arated into two parts by a 10-m wide road allowance. The glade was surrounded by a semi-open woodland that included Trembling Aspen (Populus tremuloides Michaux), Lodgepole Pine (Pinus contorta var. latifolia Engelmann), Douglas Fir (Pseudotsuga menziesii var. glauca |Beissner] Franco), and White Spruce (Picea glauca [Moench] Voss). The park is a significant natu- ral area with a rich diversity of flora and fauna (Pinel 1985*; Wallis and Wershler 1972*; Williams 1988*).
Requirements for comparison
To be able to attribute differences in vascular plant composition between the two sections to differences in grazing by feral horses, all other characteristics should be nearly the same. There is good evidence that this was the case. The two sections, “grazed” and “non-grazed,” had similar gradual slopes, elevation, and very stony substrates, and the geology of the area was uniform (Rutherford 1927; Greenlee 1974; P.M.C. personal ob- servation). The sections were separated only by a road and roadside fences, which clearly split a formerly con- tinuous prairie glade, the edges of which remained well defined.
The entire glade was grazed by feral horses until 1984 (G. Cowley, Rafter Six guest ranch, personal com- munication) when grazing was discontinued on the south side (2.2 ha). On the north side (2.8 ha), grazing by horses was permitted at a moderate stocking level in the fall (15 August to 15 October) for 652 horse-days on approximately 22 ha of prairie which included the north half of the glade as well as open woods and other glades nearby. Over this period, both halves were sub- ject to very light grazing by free-ranging Rocky Moun- tain Elk (Cervus elaphus nelsoni) and grazing to an even lesser extent by both Mule Deer (Odocoileus he- mionus) and White-tailed Deer (Odocoileus virgini- anus) (R. Jaeger, park conservation officer, personal communication). The Rocky Mountain Elk (n = 100- 160) also used the area from December to February as one of a large series of openings preferred because characteristic high wind and chinooks reduce snow ac- cumulation and adjacent wooded areas provide cover (R. Jaeger, personal communication). Other grazing mammals, including various rodents, are scarce in this area of gravelly soils and are thought to have little impact on flora (P.M.C., personal observation).
Methods Data collection and identification
Between 28 June and 2 July 2009, in each section (grazed and non-grazed), data on the presence and cov- er of species were gathered from |-m? quadrats along five parallel transects at a 45° angle from the road. Ten quadrats, each 10 m apart, were placed on each transect, resulting in 50 quadrats on either side of the road (i.e.,
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in each section). Cover for each species in each quadrat was estimated as a percentage of the 1-m* ground sur- face covered by living material of that species. The results were tallied so that species in the two sections could be compared with respect to both frequency and cover. No new species were recorded after 35 quadrats in either section suggesting that 50 was an adequate sample to describe the vegetation.
As grazing may sometimes result in more-or-less stable, heavily grazed and non-grazed patches, and in- creased heterogeneity may exist on a broad scale but not on a small scale (Willms et al. 1988), a relatively extensive sampling procedure, such as that used here, is advantageous in biodiversity comparison.
Most species were flowering at the time of sampling, and this aided in identification; the few exceptions were Gentianella amarella, Solidago simplex ssp., and Sym- phyotrichum laeve var. geyeri. Plants were identified using Packer (1994), Kuijt (1982), Hallworth and Chi- nappa (1997), and Tannas (2003a,b, 2004) as well as the online Flora of North America series (1993-2009).
The names mostly follow the recent compilation of Kartesz and Meachum (1999*) with some more recent changes from Brouillet et al. (2010+*). All species recorded in the prairie glade are listed in Appendix Table 1 with authorities, scientific names and frequent- ly used common names. Voucher specimens are pre- served in the National Collection of Vascular Plants of Agriculture and Agri-Food Canada in Ottawa (acronym DAO). Although the identifications are considered ac- curate, there were limitations. Some species were too immature to identify with certainty to the infraspecific level (e.g., Solidago simplex).
The dominant grass in the non-grazed area appeared to be Festuca campestris Rydberg, but this could be confirmed in only a few quadrats where it was flower- ing. As we believed that only one species of the rough fescue complex was present, all non-flowering plants were assumed to be this species. The site is near the boundary of the ranges of Fesctuca campestris and F altaica. Kananaskis is located at 51° N, very close to the southern limit of F. a/taica in the Canadian Cordillera (Pavlick and Looman 1984). Bowden (1960) deter- mined that the chromosome number of rough fescue plants at other sites near Banff, Alberta, was 2n = 56, which is characteristic of F campestris (F. altaica is reported with 2n=28). However, we observed some plants with characteristics of F. altaica but neverthe- less best placed with F campestris.
Data analysis
Mean cover (including zeros) for each species was calculated for each of the five, 10-quadrat transects for each grazing treatment. Frequency was the percentage of the 10 quadrats in a transect that contained that species. Covers and frequencies of each species were compared with a Proc T-test using the five transects as replicates. This test was generated with SAS soft- ware (SAS 2003*). The summary function of PC-Ord
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CATLING ET AL.: HORSE RANCHING INCREASES BIODIVERSITY IN WESTERN ALBERTA
i
TABLE |. Mean cover and frequency of : ieee . Miser is and frequency of the more abundant (> 1% mean cover in either group) of 91 species in adjacent sec- prairie either grazed or non-grazed by Feral Horses (Equus ferus caballus), northern Kananaskis Country, Alberta ; ;
Cover (%)
Frequency (%)
Species Grazed Non-grazed Grazed Non-grazed Grasses and sedges
Carex obtusata 4.14* 0.32* Lo 12* Elymus lanceolatus 9.48* 3.06* 76* 308 Fe estuca campestris 0).42* 30.26* 48* gg Koeleria macrantha 8.24* Le 88* 48* Muhlenbergia richardsonis 0.26* 3.30" 6* 26* Poa secunda 5.32* 0.72* g4* 70% Forbs > Anemone multifida 0.16* 1.08* 8* 40* Antennaria parvifolia 2.64% 0.26* 40* 8* Artemisia frigida 3.34* 0.04* 50" ae Anticlea elegans PY ONY as 0.066* 56* 28% Campanula rotundifolia 2.28 1.14 She 40* Cerastium arvense Sil oe 1.90* 94* 60* Comandra umbellata 1.40* 0.30* 36* 14* Erigeron caespitosus 1.48* OMG 44* Ge Galium boreale 3.98 4.36 90 82 Geum triflorum 0.06* 2.24* ys 52° Hedysarum boreale 0.84 1.26 Ds 18* Oxytropis monticola 4.60* Ih Sor 68* 46* Oxytropis sericea 21627 0.06* Se 8* Potentilla concinna 1.02 0.46 28 22 Primula conjugens OnlGr 1.00* 10* 42* Pulsatilla patens 2.66* 0.98* 58 46 Toxicoscordion venenosus Eo 0.06* Sy 4* Vicia americana 0.08* 1.94* De 40* Shrubs
Arctostaphylos uva-ursi 0.80* 14.84* 10* 42* Dasiphora fruticosa 0.40* 1.82* 6 16 Juniperus horizontalis 15.46 14.46 46 Sy Other
Selaginella densa 154" 1n04 68* 18*
oo — — OOOO SS OC ———_—_——<<uomo
*Significant difference between grazed and non-grazed sections (P < 0.05, Student’s ¢ test).
(McCune and Grace 2002*) was used to determine the two components of diversity (species richness and evenness) as well as two indices of diversity (Shan- non-Wiener and Simpson’s indexes) for each transect using mean species covers. The diversity variables were compared between grazing treatments (Proc T-TEST of SAS) using the results for the five transects as replicates.
Results
A total of 91 species were recorded in the 100 qua- drats and 96 species (not including surrounding trees) were recorded in the entire prairie glade including the quadrats. The full species list 1s presented in Appendix I (along with authorities, common names and the acces- sion numbers of voucher specimens at DAO). There were only five introduced species, and these were pres- ent in trace amounts.
The plant communities in grazed and non-grazed areas appeared different (Figure 1). The mean cover and mean frequency of species with a cover of at least 1% are listed in Table 1 by major group. There was consid-
erable divergence in the species composition between the grazed and non-grazed sites. Festuca campestris and Arctostaphylos uva-ursi dominated the non-grazed site where covers were much greater than in the grazed site (30.26% versus 0.42% and 14.84% versus 0.8%, respectively). Other species with greater cover in the non-grazed areas included Mulhenbergia richardsonis, Geum triflorum, Vicia americana, and Dasiphora fruiti- cosa.
With grazing, the cover of many species increased. Among the graminoids, Carex obstusata, Elymus lance- olatus ssp. lanceolatus, Koeleria macrantha, and Poa secunda ssp. secunda increased. Among the forbs Ane- mone patens var. multifida, Antennaria parvifolia, Anticlea elegans, Artemisia frigida, Cerastium arvense, Oxytropis monticola, O. sericea, and Toxicoscordion venosum var. venenosum were more abundant in the grazed sites. Selaginella densa increased considerably with grazing. Several species including Astragalus lax- mannii var. robustior, Galium boreale, Potentilla con- cinna, and Juniperus horizontalis had similar covers
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FiGurE |. Area in northern Kananaskis Country, Alberta, non-grazed by Horses (Equus ferus caballus) (left) showing clumps of Festuca campestris Rydberg and grazed area (right) showing more abundant wildflowers, including, in particular, the yellow-flowered species of Oxytropis, O. monticola and O. sericea). The blue-green colour of the prairie on the right is a result of more open-ground and pale plants such as Artemisia frigida and Elymus lanceolatus. Although still present as scattered culms, clumps of Festuca campestris are absent from the photo on the right. Photo by Paul M. Catling.
in both grazing treatments.
The mean number of species (species richness) did not differ between the grazing treatments (Table 2). Evenness was significantly greater with horse grazing. The Shannon-Wiener index indicated that feral horse grazing increased the species diversity of the plant com- munity, whereas Simpson’s index showed no effect.
TABLE 2. Effect of Feral Horse (Equus ferus caballus) graz- ing on mean diversity components determined from species covers in northern Kananaskis Country, Alberta.
Grazed Not grazed Species number 44.6 48.8 Evenness 0.802* 0.685* Shannon—Wiener index 3.041* 2.661* Simpson’s index 0.805 0.849
*Significant difference between grazed and not-grazed section (P< 0.05, Student’s ¢ test).
Discussion
Our list (Appendix 1) contains many species that have been observed in the various seral stages of the F campestris grassland at the Stavely site in the Porcupine
hills of southern Alberta (Willms e¢ a/. 1985). On the other hand, some species common in that grassland were either not present or were present in only trace amounts, including Danthonia parryi Scribner and F. idahoensis Elmer. Our site is at the northern limit of the former and beyond the northern limit of the latter (Packer 1994).
The low abundance of introduced species (5 of 94 in Appendix |) suggests that, even with grazing, this is a relatively pristine plant community. Most important, only trace amounts of Poa pratensis L., one of the main species seen to increase with grazing at many F. cam- pestris sites, were found here. Greater evenness with grazing reflects the great reduction in F. campestris and Arctostaphylous uva-ursi cover that favours a diversity of other species. Rough Fescue is known to be grazing sensitive (Willms ef al. 1985), but wl vy A. uva-ursi de- clines is not clear, as it is seldom grazed by horses. Tram- pling may be a factor,
Many of the species that increased with grazing have a more prostrate growth form and, thus, may partly avoid grazing. With grazing, four graminoids increased at the expense of F’ campestris. The “increasers” did not include Danthonia parryi, which was the main increaser at the Stavely site (Willms ef al. 1985). The cover of many of the lower growing forbs was also greater at the grazed site, including Antennaria parv ifolia, Anticlea elegans,
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Artemisia frigida, Cerastium arvense, Oxytropis spp.. and Toxicoscordion venenosum var. venenosum. Two palatable legumes (Hedysarum boreale and Vicia amer- icana) increased with protection from grazing. Because of their high nutritive value, these are likely favoured by horses.
A short stature helps some species tolerate grazing, but makes them more susceptible to competition from robust grasses like Festuca campestris; thus, they would be expected to decline with protection from grazing. Competitive exclusion occurs when removal of distur- bances, such as livestock grazing, allows a highly com- petitive species such as F. campestris to restrict the growth of other species through rapid canopy and root development, which limit both light and moisture avail- ability, and litter accumulation, which reduces recruit- ment from tillers or seed (Grime 1973). The reduced evenness without horse grazing suggests that this had occurred to some extent. It might take longer for com- petitive exclusion to reduce the species number, or there may be sufficient grazing by wild ungulates in the “non-grazed” area to slow it. Studies of recovery of rough fescue grasslands (Willms ef al. 1985; McLean and Tisdale 1972) show that, after several decades of recovery, species number was not reduced. However, in the longer term, Festuca campestris dominates to the exclusion of most other species (McLean and Tisdale 1972).
We found an increase in the Shannon-Wiener index with grazing, as shown by Bai et al. (2001) in Saska- tchewan. Also similar to Bai et al., we found no change in species richness with moderate grazing. The Simp- son’s index is less sensitive to the contribution of less- abundant species (DeJong 1975) and, thus, did not in- crease with grazing.
The results of this study correspond to those of many others (Trottier 1993; West 1993; McLaughlin and Mineau 1995) in suggesting some biodiversity bene- fits of moderate grazing. Studies of grazing effects at many sites throughout Alberta would determine whe- ther grazing affects biodiversity in all grassland types. A study conducted in the mixed prairie (Willms et al. 2002) suggests that grazing does not always improve diversity.
General observations
The species of vascular plants that were much more abundant in grazed areas are known to increase with increased grazing pressure and are mostly not grasses. These species are likely those previously avoided by bison and currently avoided to some extent by feral horses. Both of these ungulates are grass specialists. Some of the plants that apparently increased with horse grazing are toxic to horses (e.g., Oxviropis sericea and Toxicoscordion venenosum var. venenosum, Majak ef al. 2008) and were possibly avoided for this reason. Allowing only moderate grazing by horses may also be beneficial to the horses as they may avoid toxic plants while alternatives are available.
CATLING ET AL.: HORSE RANCHING INCREASES BIODIVERSITY IN WESTERN ALBERTA 19
Both the area grazed by horses and the area free of horses were significant in terms of overall biodiversity, with different species present and different abundance values of shared species and high biodiversity values according to different indexes. The grazed area fa- voured wildflowers that serve pollinators including numerous bees and butterflies (most of the forbs listed in Table 1). The Greenish Blue butterfly (Plebejus sae- piolus amica |W. H. Edwards, 1863]; Figure 2) was abundant in the grazed area, with up to 10 in view at any one time, but entirely absent from the non-grazed area. A population of Speckle-winged Rangeland Grass- hoppers (Arphia conspersa Scudder, 1875), which pre- fer areas with bare soil (P. M. C. personal observation), was present in the grazed area but entirely absent from the adjacent non-grazed area. The dominance of clumped grasses in the horse-free area provided struc- tural cover for nesting birds such as Vesper Sparrows (Pooecetes gramineus) that were absent in the grazed area. Other studies have shown that below-ground arthropods, scavenging arthropods, and grasshoppers increase with grazing (Laycock 1994) and that birds are variously adapted to the extent of the grazing (Knopf 1996). The differences in biodiversity between the two sections may be much greater for other groups than for vascular plants, further supporting the high biodi- versity value of the presence of both grazing regimes.
Management implications
Based on studies of recovery from grazing in rough fescue communities (Willms et a/ 1985; McLean and Tisdale 1972), the non-grazed area is likely at a mid- seral stage. There may be a number of distinct biodiver- sity-rich intermediate stages of succession in foothills prairies based on different levels of grazing pressure and other factors. Biodiversity was not necessarily greatest at the apparent intermediate level of distur- bance, as might be expected (Bai ef a/. 2001; Vujnovic et al. 2002). However, succession and disturbance levels may not provide the most informative view of prairies.
In pre-settlement times, prairies were likely a shift- ing mosaic of heterogeneous patches where fire and grazing played major roles, along with periodic drought (Fuhlendorf and Engle 2004). Other important factors affecting diversity may have been spatial and tempo- ral variation in movements of wandering bison herds, and fluctuations in numbers of ungulate predators, as well as variation in their distributions. For example, bison traces would have been heavily grazed, but areas where predator risk was high would not have been grazed. Drought and fire would have influenced both of these factors. In the case of our study site, biodiversity was increased by the creation of two patches with dif- ferent grazing regimes. An improvement in manage- ment would involve changing the patches over space and time by applying different ecological situations, especially using fire as well as grazing, as suggested by Fuhlendorf and Engle (2004). With the loss of free-
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FIGURE 2. The Greenish Blue (Plebejus saepiolus amica (W.H. Edwards, 1863) was an abundant visitor to flowers of Oxytropis spp., which were much more abundant in the area of prairie subject to moderate grazing by feral Horses (Equus ferus caballus) than in an adjacent grazed area in northern Kananaskis Country, Alberta. Photo by Paul M. Catling.
ranging bison and the patch dynamics on which prairie
diversity is based, the use of an increasing number of
feral horses to achieve various levels of grazing is a potentially useful management tool that deserves wide- spread but well-planned application. Grazing by horses in foothills prairies at moderate to light levels can be beneficial in terms of biodiversity.
Acknowledgements
Bonnie Smith and C. C. Chinnappa of the University of Calgary Herbarium provided invaluable logistical support. Bow Valley Provincial Park warden, Rod Jaeger, and Chief Ecologist, Melanie Percy, provided information on the history of the study area and ar- ranged for a permit to collect plant specimens. Gloria Cowley of Rafter Six guest ranch provided informa- tion on the history of the area west of the Kananaskis River.
Documents Cited (marked * in the text) Brouillet, L., F. Coursol, S.J. Meades, M. Favreau, M. Anions, P. Bélisle, and P. Desmet. 2010+. VASCAN. the
Database of Vascular Plants of Canada. Accessed 28 August 2014. http://data.canadensys.net/vascan
Kartesz, J. T., and C. A. Meacham. 1999. Synthesis of the North American Flora, Version 1.0 North Carolina Botanical Garden, Chapel Hill, North Carolina, USA
SAS Institute, Inc. 2003. Statistical analysis system. Version 9.1, Copyright © 2003 [computer program]. SAS Institute Inc. SAS and all other SAS Institute Inc. product or service names are registered trademarks or trademarks of SAS Institute Inc., Cary, North Carolina, USA.).
McCune, B., and J. B. Grace. 2002. Analysis of Ecological Communities. MjM Software, Gleneden Beach. Oregon, USA.
Wallis, C., and C. Wershler. 1972. Ecological sury ey of Bow Valley Provincial Park, vol. 1 and 2. Provincial Parks Plan- ning Department, Alberta Parks Division, Edmonton. Alberta, Canada.
Westar Inc. 2003. Horse industry profile and economic impact survey for Horse Industry Association of Alberta. Weststar Inc., Edmonton, Alberta, Canada. Accessed 28 September 2014. http//www.albertahorseindustry.ca/econo micsurvey/complete_profile.pdf ,
Williams, J. 1988. Vegetation iny entory, Bow Valley Provin- cial Park, Alberta. Recreation and Parks, Operations Branch, Kananaskis Country, Canmore, Alberta.
2015
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Received 21 January 2014 Accepted 8 May 2014
Pp THE CANADIAN FIELD-NATURALIST
Vol. 129
APPENDIX I. Total list of species recorded (including quadrats) in a prairie glade in northern Kananaskis Country, Alberta, with scientific name, common name, some synonyms, and accession number of specimens preserved in the National Collection
of Vascular Plants, Agriculture and Agri-Food Canada.
Scientific name
Achillea millefolium L.
Achnatherum richardsonii (Link) Barkworth
Agoseris glauca (Pursh) Rafinesque
Allium cernuum Roth
Androsace chamaejasme Wulfén ex Host
Androsace septentrionalis L.
Anemone multifida Poiret
Anemone patens L. var. multifida Pritzel (syn. Pulsatilla patens)
Antennaria anaphaloides Rydberg
Antennaria parvifolia Nuttall
Anticlea elegans (Pursh) Rydberg (syn. Zigadenus elegans)
Arabis hirsuta M. Hopkins var. pycnocarpa (M. Hopkins) Rollins Arctostaphylos uva-ursi (L.) Sprengel Artemisia campestris L. Artemisia frigida Willdenow Astragalus agrestis Douglas ex G. Don Astragalus australis (.) Lamarck Astragalus laxmanii Jacquin var. robustior (Hooker) Barneby and S.L. Welsh *Bromus inermis Leysser Campanula rotundifolia L. Carex duriuscula C. A. Meyer Carex filifolia Nuttall Carex obtusata Liljeblad Carex richardsonii R. Brown Carex rossti Boott Carex scirpoidea Michaux Castilleja rhexifolia Rydberg Cerastium arvense L. Comandra umbellata (L.) Nuttall ssp. pallida (A. de Candolle) Piehl Crepis runcinata (E. James) Torrey & A. Gray Dactylorhiza viridis (L.) R.M. Bateman, Pridgeon & M.W. Chase (syn. Coeloglossum viride var. virescens) Dasiphora fruticosa (L.) Rydberg (syn. Dasiphora fruticosa ssp. floribunda (Pursh) Kartesz) Delphinium bicolor Nuttall Draba cana Rydberg (syn. Draba breweri var. cana) Draba nemorosa L. Elymus glaucus Buckley Elymus lanceolatus (Scribner & J.G. Smith) Gould ssp. /anceolatus Erigeron caespitosus Nuttall Erigeron glabellus Nuttall var. glabellus (syn. Erigeron asper Nuttall) Eriogonum flavum Nuttall Erysimum inconspicuum (S. Watson) MacMillan Festuca campestris Rydberg *Festuca cf. ovina L. Festuca saximontana Rydberg var. purpusiana (Saint-Yves) Frederiksen & Pavlick Fragaria virginiana Miller Gaillardia aristata Pursh Galium boreale L. Geum triflorum Pursh Hedysarum alpinum L. (syn Hedysarum alpinum ssp. americanum (Michaux ex Pursh) B. Fedtschenko) Gentianella amarella (L.) Borner
Common name
Common Yarrow Richardson’s Needlegrass Pale Goat-chicory
Nodding Onion Sweet-flowered Rock Jasmine Pygmyflower Rock jasmine Red Windflower
Prairie Crocus
Tall Pussytoes Little-leaved Pussytoes Mountain Death Camas Hairy Rockcress
Red Bearberry
Pacific Wormwood
Prairie Sagebrush Cock’s-head
Indian Milk-vetch
Ascending Purple Milk-vetch
Smooth Brome
Round-leaf Harebell Spikerush Sedge Thread-leaved Sedge
Blunt Sedge
Richardson’s Sedge
Ross’ Sedge
Canadian Single-spike Sedge Rosy Indian-paintbrush Field Mouse-ear Chickweed Pale Bastard-toadflax
Fiddle-leaved Hawksbeard Frog Orchid
Shrubby Cinquefoil
Flathead Larkspur Cushion Whitlowgrass Woodland Whitlowgrass Blue Wildrye Streamside Wildrye
Tufted Fleabane Rough Fleabane
Alpine Golden Wild Buckwheat
Shy Wallflower Northern Rough Fescue Sheep Fescue
Rocky Mountain Fescue
Wild Strawberry Great Blanketflower Northern Bedstraw Prairie Smoke Alpine Sweet-vetch
Autumn Dwarf Gentian
Accession number
843223 843236, 843298 843264 843301 843271 843228 843221 843267
843296 843319 843320 843250
843315 843316 843278 843286, 843298
843252 843238, 843275 843277 843302 843240, 843260 843272 843297 843262 843279
843306 843314
843239
843219 843292, 843318 843253, 843323 843237 843243
843224 843220
843300
843258
843231, 843273
843291, 843295
843241, 843313, 843324
843288, 843310
843256
2015
APPENDIX 1. (continued)
CATLING ET AL.: HORSE RANCHING INCREASES BIODIVERSITY IN WESTERN ALBERTA 23
—_—_—=*"“mwvxrV<woOwTYW™“006BaN“_5eaaeeee
Scientific name
Hedysarum boreale Nuttall Hedysarum sulphurescens Rydberg Helictotrichon hookeri (Scribner) Holub
Heuchera cylindrica Douglas
Juniperus communis var. depressa Pursh
Juniperus horizontalis Moench
Koeleria macrantha (Ledebour) Schultes
*Lappula squarrosa (Retzius) Dumortier
Lilium philadelphicum L. (syn. Lilium philadelphicum var. andinum (Nuttall) Ker Gawler)
Lithospermum ruderale Douglas ex Lehmann
Maianthemum stellatum (L.) Link
Muhlenbergia richardsonis (Trinius) Rydberg
Oxytropis borealis de Candolle var. viscida (Nuttall) S.L. Welsh
Oxytropis deflexa (Pallas) de Candolle ssp. sericea (Torrey & A. Gray) Cody (syn. Oxytropis deflexa var. sericea Torrey & A. Gray)
Oxytropis monticola A. Gray
Oxytropis sericea Nuttall
Oxytropis splendens Douglas ex Hooker
Packera cana (Hooker) W.A Weber & A. Love
Penstemon confertus Douglas ex Lindley
*Phleum pratense L.
Poa cusickii Vasey
Poa pratensis L.
Poa secunda J. Presl ssp. secunda (syn. Poa sandbergii Vasey)
Poa sp.
Populus tremuloides Michaux
Potentilla concinna Richardson
Potentilla pensylvanica L.
Primula conjugens (Greene) A.R. Mast & Reveal var. conjugens (syn. Dodecatheon conjugens)
Primula pauciflora (Greene) A.R. Mast & Reveal var. pauciflora (syn. Dodecatheon pulchellum)
Rhinanthus minor L.
Sabulina rubella (Wahlenberg) Dillenberger & Kadereit (syn. Minuartia rubella)
Selaginella densa Rydberg
Silene parryi (S. Watson) C.L. Hitchcock & Maguire
Sisyrinchium montanum Greene
Sisyrinchium septentrionale E.P. Bicknell
Solidago simplex ssp.
Symphoricarpos sp.
Symphyotrichum laeve (L.) A. Love & D. Love var. geyeri (A. Gray) G.L. Nesom
* Taraxacum officinale F.W. Wiggers
Thalictrum venulosum Trelease
Toxicoscordion venenosum (S. Watson) Rydberg var. venensoum
Vicia americana Muhlenberg ex Willdenow
Viola canadensis L.,
Zizia aptera (A. Gray) Fernald
Common name
Northern Hedysarum Yellow Sweet-vetch Hooker’s Alpine Oatgrass
Poker Alumroot Common Juniper Creeping Juniper Junegrass
Bristly Sheepburr Wood Lily
Columbia Puccoon
Starry False Solomon’s Seal Mat Muhly
Boreal Locoweed
White Pendant-pod Locoweed
Yellow-flower Locoweed
Rocky Mountain Locoweed Whorled Locoweed
Silvery Groundsel
Lesser Yellow Beardtongue Common Timothy
Cusick’s Bluegrass Kentucky Bluegrass Sandberg’s Bluegrass
Bluegrass (unknown) Trembling Aspen
Red Cinquefoil Pennsylvania Cinquefoil Bonneville Shootingstar
Dark-throat Shootingstar
Little Yellow Rattle Boreal Stitchwort
Dense Spikemoss
Parry’s Catchfly
Strict Blue-eyed Grass Northern Blue-eyed Grass Mt. Albert Goldenrod Snowberry
Smooth Blue Aster
Common Dandelion Veiny-leaved Meadow-rue Meadow Death Camas
American Purple Vetch Canada Violet Heart-leaved Alexanders
Accession number
843227
843230, 843232, 843299, 843311, 843321
843274
843234, 84326] 843218 843293
843308
843354 843305
843222, 843247
843249, 843276, 843282, 843283 843248, 843280
843287 843294
843254, 843303
843269
843233, 843235, 843246, 843281
843322
843244 843245 843290
843298
843259 843242 843317 843251 843257 843309 843226 843263 843265, 843312 843229
843266
ee ee
*Non-native species.
The Flora of Cunningham Inlet, Somerset Island, Nunavut: History, Analysis, and New Collections of Vascular Plants, Mosses, Lichens, and Algae
PAUL C. SOKOLOFF
Centre for Arctic Knowledge and Exploration, Research and Collections, Canadian Museum of Nature, P.O. Box 3443, Sta- tion D, Ottawa Ontario K1P 6P4 Canada; email: psokoloff(@mus-nature.ca
Sokoloff, Paul C. 2015. The flora of Cunningham Inlet, Somerset Island, Nunavut: history, analysis, and new collections of vascular plants, mosses, lichens, and algae. Canadian Field-Naturalist 129(1): 24-37.
New collections of vascular plants, bryophytes, lichen, and algae are reported for Cunningham Inlet on the north coast of Somerset Island, Nunavut. This list of 48 species of vascular plants, 13 bryophytes, 10 lichens, and five algae includes 136 specimens collected in 2013 and 39 previously unreported specimens from the National Herbarium of Canada at the Canadian Museum of Nature (CAN), Agriculture and Agri-Food Canada’s Vascular Plant Herbarium (DAO), and University of Alberta (ALTA). Ten vascular plants from previous collecting in 1958 are re-reported here to give a comprehensive account of the vascular plant flora of the region. Two vascular plants are recorded for the first time for Somerset Island: Smooth Draba (Draba glabella Pursh) and Edlund’s Fescue (Festuca edlundiae S. G. Aiken, Consaul & Lefkovitch).
Key Words: Arctic; Nunavut; Somerset Island; Cunningham Inlet; vascular plants; Festuca edlundiae; Draba glabella
Introduction Flora of the Canadian Arctic Archipelago (Aiken et al.
Located on the northern coast of Somerset Island, 2007). These specimens may have also been mapped Nunavut, Cunningham Inlet is a focal point of marine by Porsild and Cody (1980), but they are difficult to mammal research as thousands of Beluga Whales (De/- _ distinguish among the individual dots that cover nearly phinapterus leucas) enter it each summer (Smith and __ the entirety of Somerset Island on their location maps. Sjare 1990; Smith and Martin 1994). Because of this Before Savile’s work, only one known specimen had remarkable natural phenomenon, Arctic Watch Lodge been collected from Cunningham Inlet: a single sheet was established on the inlet as a destination for wilder- of Arctic False Wallflower (Parrya arctica R. Brown)
ness tourism. Although this remote area receives nu- collected by B. Shindman during “Operation Magnetic” merous visitors and the inlet is close to Canada’s high jy 949.
Arctic research hub in Resolute (79 km away), only lim- ited botanical research has previously been carried out at Cunningham Inlet. As a result, relatively few species of vascular plants, mosses, lichens, and algae are report- ed from the inlet. This might be because of the relative lack of plant diversity on the northern shore of Somerset
Island; the Circumpolar Arctic Vegetation Map team : 5 ; : opto rs 6 ‘ P f Steven V. Zoltai and V. Woo conducted extensive soil classifies this site as cryptogam-—herb barren: sparse
barren landscapes with little vegetation cover (Walker NERS 0 Bee Sos cs a Paots fs et al. 2005). a proposed gas pipeline through the Canadian Arctic The first significant collecting activity on Somerset 1” TT Woo and Zoltai 1977). While they used field Island occurred in 1958, when D. B. O. Savile (Agri- identifications from plots and transects to characterize culture Canada) collected vascular plants, mosses, and the vegetation of habitats sampled, voucher specimens fungi from 12 sites around the perimeter of the island Wete taken and sent to Agriculture Canada (DAO), in- (Savile 1959). Before this, only sporadic collections had Cluding three species of Poaceae from the vicinity of been made, many of which are reported by Polunin Cunningham Inlet. (1940). Savile visited Cunningham Inlet very briefly on More recently, on 13 July 2004, L. Consaul and A. I] August 1958, and “selective collecting was done, | Archambault of the Canadian Museum of Nature made principally of parasitic fungi” (Savile 1959). He collect- a brief stop at Cunningham Inlet and collected 11 pre- ed 11 vascular plants (10 species) and three fungi and viously unreported specimens for the National Her- deposited them at Agriculture and Agri-Food Canada’s, _ barium of Canada (CAN), focusing primarily on False Vascular Plant Herbarium (DAO) and National Myco- Wallflowers (Parrya R. Brown) and Alkaligrasses (Puc- logical Herbarium (DAOM). These collections were — cinellia Parlatore). In total, they collected four vascular reported in Savile (1959), and a subset was mapped for __ plant species.
Over a decade passed until the next collector, L. C. Bliss of the University of Alberta, visited Cunningham Inlet to study the plant communities in polar desert habi- tats (Bliss et al. 1984). His 23 previously unreported specimens (19 vascular plant species) are deposited at the University of Alberta herbarium (ALTA).
24
2015
In July 2013, | embarked on a 6-day plant collecting trip to Cunningham Inlet as a scientist-in-residence for Arctic Watch Lodge’s 2013 Steve Amarualik Youth Leadership Expedition, a program designed to bring out leadership potential and teach outdoor skills to youth from the south and the north. The collections from this trip are described here alongside the collections made on the four previously mentioned trips. This provides a comprehensive overview of the vascular plant flora on the inlet, which consists of 48 species and 31 genera in I] plant families. I include two vascular plant species previously not known to occur on Somerset Island. In addition, | report here the first substantial inventory of cryptogams and marine algae from Cunningham Inlet.
Study Area
From 6 to 12 July 2013, I collected vascular plants, lichens, mosses, algae, and fungi in the vicinity of Cun- ningham Inlet (74°04'N, 93°48'W; Figure 1).
Methods
I collected specimens of all vascular plant and marine algae species encountered and opportunistically col- lected mosses, lichens, and fungi (Figure 2). Two stu-
ety » ay? 4} 6 , > > Nunavut din Oo
SOKOLOFF: FLORA OF CUNNINGHAM INLE'
Sokoloff Consaul & Archambault Savile
Woo & Zoltai Shindman
Bliss
© Arctic Watch Lodge
nN Nn
dents, Zachary Halem (New York, New York, USA) and Alicia Manik (Resolute, Nunavut, Canada), pro- vided extensive assistance with the collection and found additional specimens; thus they are named as collectors where appropriate. In all, 136 numbers were collected: 93 vascular plants (11 families, 31 genera, and 48 spe- cies), 12 bryophytes (eight families, 13 genera, and 13 species), 16 marine algae and terrestrial cyanobacteria (five families, five genera, and five species), 12 lichens (seven families, 10 genera, and 10 species), and two fungi. All specimens have been deposited in the rele- vant collection (CAN for vascular plants, CANM for bryophytes, CANL for lichens, and CANA for algae) in the National Herbarium of Canada at the Canadian Museum of Nature, except the fungal collections, which were deposited at DAOM. Herbaria with duplicate specimens are indicated in the species accounts by their herbarium acronym. Twelve unidentified vascu- lar plant, lichen, algae, and fungal specimens collected during the trip are not treated in this paper: Sokoloff PAO ASO, LOZ 177, 179,194, 195, 196, 205,222, 229, and 24/.
I also examined and verified the vascular plant col- lections made by Savile (11 numbers), Bliss (24 num-
of PAY
FIGURE |: Locations of vascular plant, algae fungi, and bryophyte collections made in the Cunningham Inlet area, Somerset cal lie d 5 ot! 5 ; A A pecenat St ‘ ; ae ; _ a: Ch: Island, Nunavut. Locations of earlier collections (Sokoloff, Consaul & Archambault, Savile, Woo & Zoltai, Shind-
man, and Bliss)
and Arctic Watch Lodge are denoted by symbols (right inset). There is a peninsula on the northeast a o =
corner of Cunningham Inlet not shown due to map resolution.
26 THE CANADIAN FIELD-NATURALIST
Vol. 129
FIGURE 2: Diverse specimens from the Cunningham Inlet area, Somerset Island, Nunavut: A) Flat-top Draba (Draba corymbosa R. Brown ex de Candolle), Sokoloff 245; B) Ptychostomum wrightii, Sokoloff 187, C) Purple Mountain Saxifrage (Saxifraga oppositifolia L.), Sokoloff 181, D) Fucus distichus, Sokoloff 140. Photos by P. Sokoloff.
bers), Woo and Zoltai (3 numbers), Shindman (1 num- ber), and Consaul and Archambault (11 numbers) and report all except Savile’s for the first time. Some mate- rial was examined and identified by specialists, who are listed in the acknowledgements.
The species accounts are organized alphabetically by family, genus, and species within each major group collected: algae, bryophytes, lichens, and vascular plants. Common names for vascular plant species in English and French are taken from Vascan (Brouillet et al. 2010*). Inuktitut names are taken from the Com- mon Plants of Nunavut (Mallory and Aiken 2004). Collections were plotted using SimpleMappr (open- source software, David P. Shorthouse, http://www.sim plemappr.net) (Figure 1). Species distributions are tak- en from the Annotated Checklist of the Panarctic Flora (Elven et al. 2011*).
Results
Diversity in the algae, bryophyte, and lichen collec- tions was relatively low, and the opportunistic sampling strategy makes it difficult to compare our collections with known checklists from the area. Five species of algae were documented, in five genera representing five families (not including unidentified samples). For bryophytes, 14 genera and species from eight families were collected, and 10 species and genera from seven families of lichens were documented.
For vascular plants, 48 species in 31 genera and 11 families were documented for Cunningham Inlet. This accounts for nearly half of the 98 species recorded for Somerset Island by Savile (1959) and the 75 species recorded by Woo and Zoltai (1959) and represents 40% of the 119 species documented for the Island in Flora of the Canadian Arctic Archipelago (Aiken et al. 2007, data obtained from species maps). At Cunningham Inlet, Poaceae, Brassicaceae, and Caryophyllaceae were the most species-rich families, accounting for 25%, 23%, and 15% of the species collected. The majority of vascular plants documented (65%) possess a cireumpo- lar distribution pattern; others are amphi-Beringian and North American species.
Description of specimens
ALGAE Alariacaeae
Alaria esculenta (L.) Greville — NUNAVUT: Somerset Island, in sea ice in centre of Cunningham Inlet, 3 km north of Arctic Watch Lodge, thick sea ice with algae embedded, 74°5'54"N, 93°47'15"W, 0 m. 10 July 2013, P. Sokoloff 192 (CANA 93578), Costariaceae
Agarum cribrosum Bory de Saint-Vincent — NUNA- vuT: Somerset Island, on tidal flats on west edge of Cunningham Inlet, 1.5 km north of Arctic Watch
2015
Lodge, in rocky tidal zone, mostly bare stones, with Fucus distichus, 74°4'48.7"N, 93°49'27.7"W, 53 m, 8 July 2013, P. Sokoloff 139 (CANA 93579): sea ice in centre of Cunningham Inlet, 3 km north of Arctic Watch Lodge, thick sea ice with algae embedded, 74°5'54"N, 93°47'15"W, 0m, 10 July 2013, P Sokoloff 193 (CANA 93580).
Fucaceae
Fucus distichus L. — NUNAVUT: Somerset Island, tidal flats on west edge of Cunningham Inlet, 1.5 km north of Arctic Watch Lodge, rocky tidal zone, mostly bare stones, 74°4'48.7"N, 93°49'27.7"W, 53 m, 8 July 2013, P. Sokoloff 140 (CANA 93581); tidal flats on west edge of Cunningham Inlet, 1.5 km north of Arctic Watch Lodge, rocky tidal zone, mostly bare stones, 7T4°4'48.7"N, 93°49'27.7"W, 53 m, 8 July 2013, P Sokoloff 141 (CANA 93582); tidal flats on west edge of Cunningham Inlet, 1.5 km north of Arctic Watch Lodge, rocky tidal zone, mostly bare stones, 74°4'48.7"N, 93°49'27.7"W, 53 m, 8 July 2013, P Sokoloff 142 (CANA 93583); tidal flats on west edge of Cunningham Inlet, 1.5 km north of Arctic Watch Lodge, rocky tidal zone, mostly bare stones, 74°4'48.7"N, 93°49'27.7"W, 53 m, 8 July 2013, P Sokoloff 143a (CANA 93584); sea ice in centre of Cunningham Inlet, 3 km north of Arctic Watch Lodge, thick sea ice with algae embed- ded, 74°5'54"N, 93°47'1S"W, 0 m, 10 July 2013, P. Sokoloff 191 (CANA 93585).
Laminariaceae
Saccharina latissima (L.) C.E. Lane, C. Mayes, Druehl & G.W. Saunders — NUNAVUT: Somerset Island, sea ice in centre of Cunningham Inlet, 3 km north of Arctic Watch Lodge, thick sea ice with algae embed- ded, 74°5'54"N, 93°47'15"W, 0 m, 10 July 2013, P Sokoloff 190 (CANA 93586).
Nostocaceae
Nostoc commune Vaucher ex Bornet & Flahault — Nunavut: Somerset Island, north-facing ridge west of Arctic Watch Lodge, Eriophorum—Calamagrostis mead- ow, 74°4'27.7"N, 93°50'39.4"W, 118 m, 9 July 2013, P. Sokoloff 175 (CANA 93588); north-facing ridge west of Arctic Watch Lodge, Eriophorum—Calamagrostis meadow, 74°4'27.7"N, 93°50'39.4"W, 118 m, 9 July 2013, P. Sokoloff 176 (CANA 93587).
BRYOPHYTES Amblystegiaceae
Campylium stellatum (Hedwig) Christian Erasmus Otterstram Jensen — NUNAVUT: Somerset Island, 2 km south of point at Cape Anne, muddy wet ground in marshy field alongside river, with Saxifraga oppositi- folia, Salix arctica, Dryas integrifolia, 74°6'23.3"N, "94°23'44.25"W, 26 m, 10 July 2013, P. Sokoloff, Z. Ha- lem 239 (associated species in same packet as Ditri- chum flexicaule) (CANM 332657).
Scorpidium revolvens (O.P. Swartz ex Anonymo) W.V. Rubers in A. Touw & W.V. Rubers — NUNAVUT: Somerset Island, wet snowmelt valley at base of large
SOKOLOFF: FLORA OF CUNNINGHAM INLET Zt
unnamed mountain northwest of Arctic Watch Lodge, wet mossy tundra, with Eriophorum angustifolium, HPA 1.3"N, 93°S 11.7" W, 122 m, 9 July 2013, P Sokoloff 184 (CANM 332651).
Drepanocladus sordidus (Miller Hal.) Hedends NUNAVUT: Somerset Island, south end of Sunday Lake, 7 km south of Arctic Watch Lodge, wet Eriophorum meadow, 74°0'24.9"N, 93°43'40.9"W, 26 m, 11 July 2013, P Sokoloff 220 (CANM 332655); slope above alluvial plain of Cunningham River, 4 km east of Arctic Watch Lodge, mossy bank in wet sedge mead- ow, 74°3'40.7"N, 93°41'32.2"W, 40 m, 7 July 2013, P. Sokoloff 124 (associated species in same packet as Brachythecium cirrosum) (CANM 332646).
Hygrohypnum luridum (Hedwig) Jennings — NUNA- vurT: Somerset Island, Flat Rock Falls, east coast of Cunningham Inlet, 4 km northeast of Arctic Watch Lodge, wet seepy rocks at edge of falls, 74°5'S6.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P Sokoloff 204 (CANM 332653).
Brachytheciaceae
Brachythecium cirrosum (Schwagrichen) Schim- per — NuNAvuT: Somerset Island, slope above alluvial plain of Cunningham River, 4 km east of Arctic Watch Lodge, mossy bank in wet sedge meadow, 74°3'40.7"N, 93°41'32.2"W, 40 m, 7 July 2013, P Sokoloff 124 (CANM 332646).
Bryaceae
Ptychostomum wrightii (Sullivant) J.R. Spence — Nunavut: Somerset Island, garden spot below sewage lagoon at Arctic Watch Lodge, lush green patch in rocky scree, with Salix arctica, Saxifraga oppositifolia, Papa- ver sp., Parrya arctica, 74°4'13.1"N, 93°48'55.8"W, 16 m, 9 July 2013, P. Sokoloff 187 (CANM 332652); south end of Sunday Lake, 7 km south of Arctic Watch Lodge, wet Eriophorum meadow, 74°0'24.9"N, 93°43'40.9"W, 26 m, 11 July 2013, P. Sokoloff 221 (CANM 332656).
Ditrichaceae
Distichium capillaceum (Hedwig) Bruch & Schim- per — NuNAvut: Somerset Island, confluence of Cun- ningham River and stream immediately south of Arctic Watch Lodge at Cunningham River crossing, rocky talus on south-facing slope, 74°4'2.9"N, 93°48'3 1.7" W, 58 m, 8 July 2013, P. Sokoloff 147 (associated species in same packet as Ditrichum flexicaule) (CANM 332649).
Ditrichum flexicaule (Schwagrichen) Hampe — NUNAVUT: Somerset Island, sloping west wall at Gull Canyon over dry creek bed, 4 km east of Arctic Watch Lodge, wet mossy slope, with Saxifraga oppositifolia, Cerastium arcticum, Draba glabella, 74°3'37.5"N, 93°40'17.7"W, 40 m, 7 July 2013, PR Sokoloff 132 (CANM 332647); confluence of Cunningham River and stream immediately south of Arctic Watch Lodge at Cunningham River crossing, rocky talus on south- facing slope, with Cerastium alpinum, 74°4'2.9"N, 93°48'31.7"W, 58 m, 8 July 2013, P Sokoloff 147
28 THE CANADIAN FIELD-NATURALIST
(CANM 332649); 2 km south of point at Cape Anne, muddy wet ground in marshy field alongside river, with Saxifraga oppositifolia, Salix arctica, Dryas integri- folia, 74°6'23.3"N, 94°23'44.25"W, 26 m, 10 July 2013, P. Sokoloff, Z. Halem 239 (associated species in same packet as Orthothecium chryseum) (CANM 332657).
Grimmiaceae
Schistidium rivulare (Bridel) Podpera — NUNAVUT: Somerset Island, Flat Rock Falls, east coast of Cun- ningham Inlet, 4 km northeast of Arctic Watch Lodge, dry rocks at edge of falls, 74°5'5S6.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P Sokoloff 205 (CANM 332654).
Hypnaceae
Orthothecium chryseum (Schwagrichen in Schultes) Schimper in Bruch & Schimper — NUNAVUT: Somerset Island, 2 km south of point at Cape Anne, muddy wet ground in marshy field alongside river, with Saxifra- ga oppositifolia, Salix arctica, Dryas integrifolia, 74°6'23.3"N, 94°23'44.25"W, 26 m, 10 July 2013, P Sokoloff, Z. Halem 239 (CANM 332657); slope above alluvial plain of Cunningham River, 4 km east of Arc- tic Watch Lodge, mossy bank in wet sedge meadow, 74°3'40.7"N, 93°41'32.2"W, 40 m, 7 July 2013, P Sokoloff 124 (associated species in same packet as Brachythecium cirrosum) (CANM 332646).
Mniaceae
Cinclidium cf. arcticum (Bruch & Schimper) Schim- per — NUNAVUT: Somerset Island, 2 km south of point at Cape Anne, muddy wet ground in marshy field along- side river, with Saxifraga oppositifolia, Salix arctica, Dryas integrifolia, 74°6'23.3"N, 94°23'44.25"W, 26 m, 10 July 2013, P. Sokoloff; Z. Halem 239 (associated species in same packet as Orthothecium chryseum) (CANM 332657).
Mnium blyttii Bruch & Schimper — NUNAvuT: Som- erset Island, eastern edge of Gull Canyon, 4 km east of Arctic Watch Lodge, extremely deep moss directly above snow line, 74°3'48.4"N, 93°40'57.2"W, 40 m, 7 July 2013, P. Sokoloff 137 (CANM 332648).
Scorpidiaceae
Sanionia uncinata (Hedwig) Loeske — NUNAVUT: Somerset Island, western edge of Gull Canyon, 4 km east of Arctic Watch Lodge, extremely deep moss di- rectly above snow line, 74°3'48.4"N, 93°40'57.2"W, 40 m, 7 July 2013, PR. Sokoloff 137 (associated species in same packet as Mnium blyttii) (CANM 332648).
LICHENS Icmadophilaceae
Thamnolia subuliformis (Ehrhart) W.L. Culberson — Nunavut: Somerset Island, mossy wet area outside whale biologist’s cabin, 1.5 km north of Arctic Watch Lodge, wet mossy rocks, 74°4'58.8"N, 93°50'2"W, 94 m, 8 July 2013, P. Sokoloff 157 (CANL 125971).
Lecanoraceae Lecidella patavina (A. Massalongo) Knoph & Leuckert — NUNAVUT: Somerset Island, rocky beach on
Vol. 129
west coast of Cunningham Inlet, directly across from Flat Rock Falls 4 km north of Arctic Watch Lodge, barren rocks, 74°5'40"N, 93°50'16"W, 5 m, 10 July 2013, P. Sokoloff 211 (CANL 125978).
Megasporaceae
Aspicilia candida (Anzi) Hue — NUNAVUT: Somerset Island, Flat Rock Falls, east coast of Cunningham Inlet, 4 km northeast of Arctic Watch Lodge, dry gravel scree above falls, 74°5'56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, PR. Sokoloff, R. Weber 206 (CANL 125980).
Megaspora verrucosa (Acharius) Hafellner & V. Wirth — NUNAVUT: Somerset Island, mossy wet area outside whale biologist’s cabin, 1.5 km north of Arctic Watch Lodge, wet mossy rocks, with Saxifraga cernua, Luzula confusa, Draba sp., Salix arctica, 74°4'58.8"N, 93°50'2"W, 94 m, 8 July 2013, P Sokoloff 160 (CANL 125974); mossy wet area outside whale biologist’s cab- in, 1.5 km north of Arctic Watch Lodge, wet mossy rocks, with Saxifraga cernua, Luzula confusa, Draba sp., Salix arctica, 74°4'58.8"N, 93°50'2"W, 94 m, 8 July 2013, P. Sokoloff 163 (CANL 125976).
Parmeliaceae
Allocetraria madreporiformis (Withering) Kaérne- felt & A. Thell — NuNAvutT: Somerset Island, mossy wet area outside whale biologist’s cabin, 1.5 km north of Arctic Watch Lodge, wet mossy rocks, with Sax- ifraga cernua, Luzula confusa, Draba sp., Salix arc- tica, 74°4'58.8"N, 93°50'2"W, 94 m, 8 July 2013, P Sokoloff 158 (CANL 125972).
Evernia divaricata (L.) Acharius — NUNAVUT: Som- erset Island, windswept dry rocks on ridge off north- west coast of Cunningham Inlet, dry rocky talus, with Saxifraga oppositifolia, 74°4'35.3"N, 93°50'7.9"W, 98 m, 9 July 2013, P Sokoloff 186 (CANL 125977).
Vulpicida tilesii (Acharius) J.-E. Mattsson & M.J. Lai — NuNAvuT: Somerset Island, west ridge overlook- ing Gull Canyon, 4 km east of Arctic Watch Lodge, dry rocky scree, with Festuca sp., Saxifraga oppositifolia, 74°3'42.4"N, 93°40'47.8"W, 50 m, 7 July 2013, P Sokoloff 129 (CANL 125969).
Peltigeraceae
Peltigera ponojensis Gyelnik — NUNAVUT: Somerset Island, south end of Sunday Lake, 6 km south of Arctic Watch Lodge, mossy knoll in wet tundra, 74°1'7.8"N. 93°45'41.5"W, 66 m, 11 July 2013, P Sokoloff 214 (CANL 125979).
Physciaceae
Physcia dubia (Hoffmann) Lettau — NUNAVUT: Som- erset Island, west ridge overlooking Gull Canyon, 4 km east of Arctic Watch Lodge, dry rocky scree, with Festuca sp., Saxifraga oppositifolia, 74°3'42.4"N. 2840)47,8 W. 50 mi, 7 iily 2018, P Sokoloff 130 (CANL 125970).
Teloschistaceae Xanthoria elegans (Link) Th. Fries — NUNAVUT: Somerset Island, mossy wet area outside whale biol-
2015
ogist’s cabin, 1.5 km north of Arctic Watch Lodge, wet mossy rocks, with Saxifraga cernua, Luzula confusa, Draba sp., Salix arctica, 74°4'58.8"N, 93°50'2"W, 94 m, 8 July 2013, P. Sokoloff 161 (CANL 125975). Vascular Plants
Brassicaceae
Braya glabella Richardson ssp. purpurascens (R. Brown) W.J. Cody (Purple Braya, braya purpurine, Airaujuit) [circumpolar-cordilleran] — NUNAVUT: Som- erset Island, on a mild slope, with Dryas integrifolia, Salix, 74°6'00"N, 93°5L'00"W, 63 m, July 13, 2004, A. Archambault & L. Consaul aa53 (CAN 603400); west ridge overlooking Gull Canyon, 4 km east of Arctic Watch Lodge, tundra in dry mud, with Dryas integri- folia, 74°3'41.1"N, 93°41'18.8"W, 50 m, 7 July 2013, P Sokoloff 125 (CAN 603292).
Cardamine bellidifolia L. (Alpine Bittercress, car- damine a feuilles de paquerette) [circumpolar-alpine] — Nunavut: Somerset Island, Flat Rock Falls, east coast of Cunningham Inlet, 4 km northeast of Arctic Watch Lodge, wet snow-patch community at foot of packed snowbank, near foot of falls, 74°5'56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P Sokoloff 197 (CAN 603293); south end of Sunday Lake, north of Cunningham River, 7 km south of Arctic Watch Lodge, mud flats, 74°0'23.3"N, 93°42'43.5"W, 41 m, 11 July 2013, P. Sokoloff 230 (CAN 603294).
Cochlearia groenlandica L. (Greenland Scurvygrass, cranson du Groenland, Tipitsiariktut nunarait) [cir- cumpolar] — NUNAvuT: Somerset Island, south end of Sunday Lake, north of Cunningham River, 7 km south of Arctic Watch Lodge, mud flats, 74°0'23.3"N, 93°42'43.5"W, 41 m, 11 July 2013, P Sokoloff 233 (CAN 603295).
Draba corymbosa R. Brown ex de Candolle (Flat- top Draba, drave en corymbe) [circumpolar] — Cun- ningham Inlet, 8 km from inlet, uplands, polar desert, Somerset plateau, 74°06'N, 93°55'W, 250 m, 20 July 1976, L.C. Bliss s.n. (ALTA 56724); Cunningham Inlet, 8 km from inlet, uplands, polar desert, 74°06'N, 93°55'W, 225 m, 20 July 1976, L.C. Bliss s.n. (ALTA 56723); dry stone ridge immediately west of Arctic Watch Lodge, in gravel scree, dry gravel, with Sax- ifraga oppositifolia, Salix arctica, Papaver sp. 74°4'20.2"N, 93°49'22.6"W, 20 m, 6 July 2013, P Sokoloff 112 (CAN 603296), western cliff wall of Gull Canyon, below gull nesting area, wet rocks directly under water seeps and waterfall, dense lush vegetation, with Saxifraga cespitosa, Bistorta vivipara, Saxifraga cernua, bryophytes, 74°3'48.4"N, 93°40'57.2"W, 40 m, 7 July 2013, P. Sokoloff 135 (CAN 603297); west ridge overlooking Gull Canyon, 4 km east of Arctic Watch Lodge, dry rocky scree, with Xanthoria elegans, Festuca sp., Saxifraga oppositifolia, 74°3'42.4"N, 93°40'47.8"W, 50 m, 7 July 2013, P Sokoloff 136 (CAN 603298); gravelly scree ledge above Cunning- ham River, directly adjacent to Arctic Watch Lodge (north side), rocky talus irrigated by water pipe, with
SOKOLOFF: FLORA OF CUNNINGHAM INLE1 29
Draba corymbosa, 74°4'10.8"N, 93°48'37.4"W, 58 m, 8 July 2013, P. Sokoloff 151 (CAN 603299); north- facing slope west of Arctic Watch Lodge, wet rocky seep in Dryas—Eriophorum tundra, with Dryas integri-
folia, Salix arctica, Draba corymbosa, Cerastium arc-
ticum, 74°4'17.5"N, 93°49'17.2"W, 16 m, 9 July 2013, P. Sokoloff 165 (CAN 603300); Flat Rock Falls, east coast of Cunningham Inlet, 4 km northeast of Arctic Watch Lodge, wet Dryas tundra with snow-bed com- munity, with Saxifraga oppositifolia, Salix arctica, 74°5'S56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P. Sokoloff 210 (CAN 603301); gravelly scree ledge above Cunningham River, 9 km southeast of Arctic Watch Lodge, dry gravel and barren rocks, 73°59'27.3"N, 93°40'57.5"W, 69 m, 11 July 2013, P. Sokoloff 234 (CAN 603302); garden spot below sewage lagoon at Arctic Watch Lodge, lush green patch in rocky scree, with Salix arctica, Saxifraga oppositifolia, Papaver sp., Parrya arctica, bryophytes, 74°4'13.1"N, 93°48'S5.8"W, 16 m, 12 July 2013, P Sokoloff, A. Manik 245 (CAN 603303).
Draba glabella Pursh (Smooth Draba, drave glabre) [circumboreal-polar] — NUNAvuUT: Somerset Island, western cliff wall of Gull Canyon, below gull nesting area, wet rocks directly under water seeps and waterfall, dense lush vegetation, with Saxifraga cespitosa, Bistor- ta vivipara, Saxifraga cernua, bryophytes, 74°3'48.4"N, 93°40'57.2"W, 40 m, 7 July 2013, P Sokoloff 134 (CAN 603304).
Draba lactea Adams (Milky Draba, drave laiteuse) [circumpolar] — NUNAvUT: Somerset Island, Cunning- ham Inlet, 1.6 km from inlet, uplands, polar desert, sedge meadow, 74°06'N, 93°55'W, 20 m, 22 July 1976, L.C. Bliss sn. (ALTA 56712);
Draba nivalis Liljebkad (Snow Draba, drave des neiges) [circumpolar-alpine] — NUNAvUT: Somerset Is- land, south end of Sunday Lake, north of Cunning- ham River, 7 km south of Arctic Watch Lodge, lem- ming mound in middle of mud flats, 74°0'23.3"N, 93°42'43.5"W, 41 m, 11 July 2013, P Sokoloff 228 (CAN 603305).
Draba simmonsii Elven & Al-Shebaz (Simmons’ Draba, drave de Simmons) [North American] — NUNA- vutT: Somerset Island, gravelly scree ledge above Cun- ningham River, directly adjacent to Arctic Watch Lodge (north side), rocky talus irrigated by water pipe, with Draba corymbosa, 74°4'10.8"N, 93°48'37.4"W, 58 m, 8 July 2013, P. Sokoloff 152 (CAN 603306).
Draba subcapitata Simmons (Ellesmere Island Draba, drave subcapitée) [circumpolar] — NUNAVUT: Somerset Island, scarce on dry gravel slope, 74°6'N, 93°5I'W, 11 August 1958, D.B.O. Savile 3786 (DAO S2S91S).
Eutrema edwardsii R. Brown (Edwards’ Mock Wall- flower, eutréma d’Edwards) [circumpolar-alpine] — NUNAVUT: Somerset Island, Cunningham Inlet, 1.6 km from inlet, Stand 15, uplands, polar desert, sedge mead- ow, 74°06'N, 93°55'W, 20 m, 22 July 1976, L.C. Bliss s.n. (ALTA 56711).
30 THE CANADIAN FIELD-NATURALIST
Parrya arctica R. Brown (Arctic False Wallflower, parrya arctique) [North American] — NUNAVUT: Som- erset Island, Cunningham Inlet, 0.16 km from inlet, uplands, polar desert, coastal beach ridge, 74°06'N, 93°55'W, 20m, 23 July 1976, 2.C. Bliss sm (ALTA 56729); Cunningham Inlet, on a mild slope, with Dryas integrifolia, 74°6'00"N, 93°51’00"W, 63 m, 13 July 2004, A. Archambault & L. Consaul aa48 (CAN 603404), aa49 (CAN 603403), aa52 (CAN 603402), aa53 (CAN 603401), aa55 (CAN 603399), aa56 (CAN 603397), aa57 (CAN 603398), aa59 (CAN 603405); scattered on dry calcareous gravel slope, 74°6'N, 93°S1'W, 11 August 1958, D.B.O. Savile 3783 (DAO 567232); Cunningham Inlet, 74°N, 94°W, 7 August 1949, B. Shindman s.n. (DAO 567235); west ridge overlooking Gull Canyon, 4 km east of Arctic Watch Lodge, tundra in dry mud, with Dryas integrifolia, 74°3'41.1"N, 93°41'18.8"W, 50 m, 7 July 2013, P Sokoloff 126 (CAN 603307); wet snowmelt valley at base of large unnamed mountain northwest of Arctic Watch Lodge, dry mud and clay mound in wet turfy tundra, with Festuca sp., 74°4'31.3"N, 93°51'1.7"W, 122 m, 9 July 2013, P. Sokoloff 180 (CAN 603308); wet snowmelt valley at base of large unnamed moun- tain northwest of Arctic Watch Lodge, wet mossy tun- dra, with Eriophorum angustifolium, 74°4'31.3"N, 93°S V1.7" W, 122 m, 9 July 2013, P Sokoloff 185 (CAN 603309); south end of Sunday Lake, 7 km south of Arctic Watch Lodge, wet Eriophorum meadow, 74°0'24.9"N, 93°43'40.9"W, 26 m, 11 July 2013, P Sokoloff 224 (CAN 603310); shoreline on north coast of Somerset Island, 7 km west of entrance to Cunning- ham Inlet, rocky, snow-pack mountain, with Parry arc- tica, Saxifraga oppositifolia, Salix arctica, 74°7'56.9"N, OF WE2ISeW, 23 mo Wl July 2013) 2 Sokolo#e Z. Halem 242 (CAN 603311).
Caryophyllaceae
Cerastium arcticum Lange (Arctic Chickweed, céraiste arctique, Nunarait qakuqtat) [North American— amphi-Atlantic-European] — NUNAVUT: Somerset Is- land, Cunningham Inlet, 8 km from inlet, uplands, polar desert, 74°06'N, 93°55'W, 200 m, 23 July 1976, L.C. Bliss s.n. (ALTA 56720); west ridge overlooking Gull Canyon, 4 km east of Arctic Watch Lodge, dry rocky scree, with Xanthoria elegans, Festuca sp., Saxifraga oppositifolia, 74°3'42.4"N, 93°40'47.8"W, 50 m, 7 July 2013, P. Sokoloff 131 (CAN 603312); confluence of Cunningham River and stream immediately south of Arctic Watch Lodge at Cunningham River crossing, rocky talus on south-facing slope, 74°4'2.9"N, 93°48'31.7"W, 58 m, 8 July 2013, PR Sokoloff 148 (CAN 603313); gravelly scree ledge above Cunning- ham River, directly adjacent to Arctic Watch Lodge (north side), rocky talus irrigated by water pipe, with Draba corymbosa, 74°4'10.8"N, 93°48'37.4"W, 58 m, 8 July 2013, P. Sokoloff 155 (CAN 603314).
Cerastium beeringianum Chamussi & Schlechtendal (Bering Sea Chickweed, céraiste du détroit de Béring)
Vol. 129
[Asian—amphi-Beringian—North American] — NUNAVUT: Somerset Island, occasional on slightly moist calcare- ous gravel slopes, 74°6'N, 93°51'W, 11 August 1958, D.B.O. Savile 3784 (DAO 562713).
Cerastium regelii Ostenfeld (Regel’s Chickweed, céraiste de Regel) [circumpolar] — NUNAVUT: Somer- set Island, dry stone ridge immediately west of Arctic Watch Lodge, east-facing wet muddy snow bed set in shale rocks, with Luzula confusa, Sabulina rubella, TACAD |. IN, 93°50'30,7 W122 mo, yo, 201s. ie Sokoloff 119 (CAN 603315).
Sabulina rossii (R. Brown ex Richardson) Dillen- berger & Kadereit (Ross’ Stitchwort, sabline de Ross) [amphi-Beringian—North American—amphi-Atlantic] — NUNAVUT: Somerset Island, Cunningham Inlet, 8 km from inlet, uplands, polar desert, 74°06'N, 93°55'W, 200 m, 23 July 1976, L.C. Bliss s.n. (ALTA 72149); north-facing ridge west of Arctic Watch Lodge, Eriophorum-Arctagrostis meadow, 74°4'17.5"N, 93°49'17.2"W, 16 m, 9 July 2013, P Sokoloff 169 (CAN 603316); Flat Rock Falls, east coast of Cunning- ham Inlet, 4 km northeast of Arctic Watch Lodge, wet snow-patch community at foot of packed snowbank, near foot of falls, 74°5'56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P. Sokoloff 201 (CAN 603317).
Sabulina rubella (Wahlenberg) Dillenberger & Kadereit (Reddish Stitchwort, sabline rougeatre, Kakil- larnait) [circumpolar-alpine] — NUNAVUT: Somerset Island, scarce on calcareous gravel slope, 74°6'N, 93°51'W, 11 August 1958, D.B.O. Savile 3787 (DAO 527824); Cunningham Inlet, 8 km from inlet, uplands, polar desert, 74°06'N, 93°55'W, 200 m, 20 July 1976, L.C. Bliss s.n. (ALTA 56728); dry stone ridge immedi- ately west of Arctic Watch Lodge, east-facing wet mud- dy snow bed set in shale rocks, with Luzula confusa, 74°4'21.3"N, 93°50'30.7"W, 122 m, July 6, 2013, P Sokoloff 118 (CAN 603318); Flat Rock Falls, east coast of Cunningham Inlet, 4 km northeast of Arctic Watch Lodge, wet snow-patch community at foot of packed snowbank, near foot of falls, 74°5'56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P Sokoloff 202 (CAN 603319); south end of Sunday Lake, 6 km south of Arctic Watch Lodge, mossy knoll in wet tundra, 74°1'7.8"N, 93°45'41.5"W, 66 m, 11 July 2013, P Sokoloff 213 (CAN 603320); lemming mound at top of hill at south end of Sunday Lake, 7 km south of Arctic Watch Lodge, with Potentilla sp., Alopecurus magel- lanicus, 74°0'38.8"N, 93°44'30.4"W, 49 m, 11 July 2013, P. Sokoloff 219 (CAN 603321). ;
Silene uralensis (Ruprecht) Bocquet ssp. uralensis (Nodding Catchfly, siléne de l’Oural, Pulluliujuit) [European—Asian—amphi-Beringian—North American] — NuNAvuT: Somerset Island, scattered on wet grav- elly slope, 74°6'N, 93°51'W, 11 August 1958. D.B.O. Savile 3781 (DAO 537745); gravelly scree ledge above Cunningham River, directly adjacent of Arctic Watch Lodge (north side), rocky talus irrigated by water pipe, with Draba corymbosa, 74°4'10.8"N. 93°48'37.4"W. 58 m, 8 July 2013, P Sokoloff 153 (CAN 603322).
2015
Stellaria longipes Goldie (Long-stalked Starwort, stellaire 4 longs pédicelles, Miqqaviat) [circumboreal- polar] — NUNAVUT: Somerset Island, confluence of Cunningham River and stream immediately south of Arctic Watch Lodge at Cunningham River crossing, rocky talus on south-facing slope, with Cerastium arc- ticum, 74°4'2.9"N, 93°48'31.7"W, 58 m, 8 July 2013, P. Sokoloff 145 (CAN 603323); wet snowmelt valley at base of large unnamed mountain northwest of Arctic Watch Lodge, dry mud and clay mound in wet turfy tundra, with Festuca sp., 74°4'31.3"N, 93°51'1.7"W, 122 m, 9 July 2013, P Sokoloff 182 (CAN 603324, NFM); gravelly scree ledge above Cunningham River, 9.6 km southeast of Arctic Watch Lodge, muddy bank, 73°59'27.3"N, 93°40'57.5"W, 69 m, 11 July 2013, P. Sokoloff 235 (CAN 603325, US); gravelly scree ledge above Cunningham River, 9.6 km southeast of Arctic Watch Lodge, muddy bank, 73°59'27.3"N, 93°40'57.5"W, 69 m, 11 July 2013, P Sokoloff 236 (CAN 603326).
Cyperaceae
Carex aquatilis var. minor Boott (Arctic Water Sedge, carex mineur, Kilirnait) [circumboreal-polar] — NUNA- vutT: Somerset Island, south end of Sunday Lake, 7 km south of Arctic Watch Lodge, wet Eriophorum meadow, 74°0'24.9"N, 93°43'40.9"W, 26 m, 11 July 2013, P Sokoloff 244 (CAN 603328).
Carex capillaris ssp. fuscidula (V.1. Kreczetovicz ex T.V. Egorova) A. Léve & D. Love (Dusky-spike Sedge, carex a epis sombres) [circumpolar-alpine] — NUNAVUT: Somerset Island, Cunningham Inlet, 1.6 km from inlet, Stand 15, uplands, polar desert, sedge meadow, 74°06'N, 93°55'W, 20 m, 22 July 1976, L.C. Bliss s.n. (ALTA 56726); north-facing slope west of Arctic Watch Lodge, wet rocky seep in Dryas—Eriophorum tundra, with Dryas integrifolia, Salix arctica, Draba corymbosa, Cerastium arcticum, 74°4'17.5"N, 93°49'17.2"W, 16 m, 9 July 2013, P. Sokoloff 168 (CAN 603329), Flat Rock Falls, east coast of Cunningham Inlet, 4 km northeast of Arctic Watch Lodge, wet Dryas tundra with snow- bed community, with Saxifraga oppositifolia, Salix arc- tica, 74°5'56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P. Sokoloff 207 (CAN 603330).
Carex membranacea Hooker (Fragile Sedge, carex membraneux, Kilirnait ajjikasangit iviit) [amphi- Beringian—North American] — NUNAvuT: Somerset Is- land, scattered in small sedge meadow below limestone hill, 74°6'N, 93°51'W, 11 August 1958, D.B.O. Savile 3777 (DAO 363708).
Eriophorum triste (Th. Fries) Hadac & A. Love (Tall Cottongrass, linaigrette triste) {amphi-Beringian—North American—amphi-Atlantic] — NUNAVUT: Somerset Is- land, Cunningham Inlet, 1.6 km from inlet, wet sedge tundra, sedge meadow, 74°06'N, 93°55'W, 20 m, 22 July 1976, L.C. Bliss s.n. (ALTA 56715); North-facing ridge west of Arctic Watch Lodge, Eriophorum—Arcta-
grostis meadow, 74°4'17.5"N, 93°49'17.2"W, 16 m, 9 July 2013, P. Sokoloff 170 (CAN 603335, US); south
SOKOLOFF: FLORA OF CUNNINGHAM INLET 3]
end of Sunday Lake, 7 km south of Arctic Watch Lodge, wet Eriophorum meadow, 74°0'24.9"N, 93°43'40.9"W, 26 m, 11 July 2013, P. Sokoloff 222 (CAN 603336); south end of Sunday Lake, 7 km south of Arctic Watch Lodge, wet Eriophorum meadow, 74°0'24.9"N, 93°43'40.9"W, 26 m, 11 July 2013, P Sokoloff 223 (CAN 603337).
Juncaceae
Juncus biglumis L. (Two-flowered Rush, jonce a deux glumes, Iviit) [circumpolar-alpine] — NUNAVUT: Som- erset Island, occasional on wet calcareous slopes, 74°6'N, 93°51'W, 11 August 1958, D.B.O. Savile 3778 (DAO 781829); Cunningham Inlet, 1.6 km from inlet, wet sedge tundra, coastal lowland beach ridges, 74°06'N, 93°55'W, 20 m, 23 July 1976, L.C. Bliss s.n. (ALTA 56721); north-facing slope west of Arctic Watch Lodge, wet rocky seep in Dryas—Eriophorum tundra, with Dryas integrifolia, Salix arctica, Draba corymbosa, Cerastium arcticum, 74°4'17.5"N, 93°49'17.2"W, 16 m, 9 July 2013, P. Sokoloff 166 (CAN 603341, US); wet snowmelt valley at base of large unnamed moun- tain northwest of Arctic Watch Lodge, wet mossy tundra, with Eriophorum angustifolium, 74°4'31.3"N, 93°51'1.7"W, 122 m, 9 July 2013, P Sokoloff 183 (CAN 603342).
Luzula nivalis (Laestadius) Sprengel (Arctic Wood- rush, luzule arctique) [circumpolar-alpine] — NUNAVUT: Somerset Island, scattered on moist calcareous slopes, 74°6'N, 93°51'W, 11 August 1958, D.B.O. Savile 3782 (DAO 780607); dry stone ridge immediately west of Arctic Watch Lodge, east-facing wet muddy snow bed set in shale rocks, with Sabulina rubella, 74°4'21.3"N, 93°50'30.7"W, 122 m, July 6, 2013, P. Sokoloff 117 (CAN 603343); north-facing slope west of Arctic Watch Lodge, wet rocky seep in Dryas—Eriophorum tundra, with Dryas integrifolia, Salix arctica, Draba corym- bosa, Cerastium arcticum, 74°4'17.5"N, 93°49'17.2"W, 16 m, 9 July 2013, P Sokoloff 167 (CAN 603344); Flat Rock Falls, east coast of Cunningham Inlet, 4 km northeast of Arctic Watch Lodge, wet snow-patch com- munity at foot of packed snowbank, near foot of falls, 74°5'56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P Sokoloff 199 (CAN 603345, US).
Orobanchaceae
Pedicularis lanata Willdenow ex Chamisso & Schlechtendal (Woolly Lousewort, pédiculaire laineuse, Ugjungnaq) [amphi-Beringian—North American] — NuNAvuT: Somerset Island, south end of Sunday Lake, 6 km south of Arctic Watch Lodge, wet Salix— Dryas meadow, 74°0'S1.7"N, 93°44'57.6"W, 64 m, 11 July 2013, PR Sokoloff 216 (CAN 603347); south end of Sunday Lake, 7 km south of Arctic Watch Lodge, wet Eriophorum meadow, 74°0'24.9"N, 93°43'40.9"W, 26 m, 11 July 2013, P. Sokoloff 225 (CAN 603348). Papaveraceae
Papaver cornwallisense D. Love (Cornwallis Island Poppy, pavot de Cornwallis) [North American—amphi-
32 THE CANADIAN FIELD-NATURALIST
Atlantic] — NUNAvuT: Somerset Island, Cunningham Inlet, 0.16 km from inlet, uplands, polar desert, 74°06'N, 93°Sa'W, 20im, 23° July 1976, 0. C. Bliss. (AICTA 56725); Cunningham Inlet, 0.16 km from inlet, up- lands, polar desert, 74°06'N, 93°55'W, 20 m, 23 July 1976, L.C. Bliss s.n. (ALTA 56710); dry stone ridge immediately west of Arctic Watch Lodge, in gravel scree, dry slaty gravel, with Saxifraga oppositifolia, Salix arctica, Papaver sp., 74°4'20.2"N, 93°49'22.6"W, 20 m, July 6, 2013, P Sokoloff 115 (CAN 603349); western cliff wall of Gull Canyon, below Gull nesting area, wet rocks directly under water seeps and waterfall, dense lush vegetation, with Saxifraga cespitosa, Bistor- ta vivipara, Saxifraga cernua, bryophytes, 74°3'48.4"N, 93°40'57.2"W, 40 m, 7 July 2013, P Sokoloff 136 (CAN 603350); gravelly scree ledge above Cunning- ham River, directly adjacent to Arctic Watch Lodge (north side), rocky talus irrigated by water pipe, with Draba corymbosa, 74°4'10.8"N, 93°48'37.4"W, 58 m, 8 July 2013, P. Sokoloff 150 (CAN 603351); south end of Sunday Lake, north of Cunningham River, 7 km south of Arctic Watch Lodge, mud flats, 74°0'23.3"N, 93°42'43.5"W, 41 m, 11 July 2013, P Sokoloff 231 (CAN 603352).
Poaceae
Alopecurus magellanicus Lamarck (Alpine Foxtail, vulpin boréal, Ivi) [circumpolar-alpine and South American] — NUNAVUT: Somerset Island, site no. Z-25, level lacustrine well- to imperfectly drained silt plain, grass-saxifrage foxhole mound, 74°2'N, 93°30'W, 53 m, 260 m, July 1, 1975, S.C. Zoltai 751152 (DAO 137589); mound at top of hill at south end of Sunday Lake, 7 km south of Arctic Watch Lodge, lush lemming mound, with Potentilla sp., Sabulina rubella, 74°0'38.8"N, 93°44'30.4"W, 49 m, 11 July 2013, P Sokoloff 218 (CAN 603353); south end of Sunday Lake, north of Cunningham River, 7 km south of Arctic Watch Lodge, lemming mound in middle of mud flats, 74°0'23.3"N, 93°42'43.5"W, 41 m, 11 July 2013, P Sokoloff 226 (CAN 603354); confluence of Cunningham River and stream immediately south of Arctic Watch Lodge at Cunningham River crossing, rocky talus on south- facing slope, with bryophytes, Cerastium arcticum, THAD ING S348 37 We Som, muly 20s Sokoloff 143b (CAN 603355).
Arctagrostis latifolia (R. Brown) Grisebach ssp. latifolia (Polargrass, arctagrostide a larges feuilles) [circumpolar-alpine] — NUNAvUT: Somerset Island, Cunningham Inlet, 1.6 km from inlet, wet sedge tundra on coastal lowlands, 74°06'N, 93°55'W, 20 m, 23 July 1976, L.C. Bliss s.n., (ALTA 56714); alluvial plain of Cunningham River on Cunningham Inlet, 3 km east of Arctic Watch Lodge, wet sedge meadow emerging from melting snowbank, with Deschampsia sp., Poa sp., 74°3'40.1"N, 93°42'17.6"W, 40 m, 7 July 2013, P Sokoloff 121 (CAN 603356); north-facing ridge west of Arctic Watch Lodge, Eriophorum—Arctagrostis
Vol. 129
meadow, 74°4'17.5"N, 93°49'17.2"W, 16 m, 9 July 2013, P. Sokoloff 172 (CAN 603357).
Arctophila fulva (Trinius) Andersson (Pendant Grass, arctophile fauve) [circumpolar] - NUNAVUT: Somerset Island, site no. Z-44, level fen with poorly drained or- ganic soil, with Drepanocladus sp., 74°1'N, 93°27'W, 53 m, 14 July 1975, S. C. Zoltai 751114 (DAO 137672).
Deschampsia brevifolia R. Brown (Short-leaved Hairgrass, deschampsie a feuilles courtes) [Asian— amphi-Beringian—North American] — NUNAVUT: Som- erset Island, alluvial plain of Cunningham River on Cunningham Inlet, 3 km east of Arctic Watch Lodge, wet sedge meadow emerging from melting snowbank, with Poa _ sp., Arctagrostis sp., 74°3'40.1"N, 93°42'17.6"W, 40 m, 7 July 2013, P Sokoloff 122 (CAN 603358); alluvial plain of Cunningham River on Cunningham Inlet, 3 km east of Arctic Watch Lodge, wet sedge meadow emerging from melting snowbank, with Poa sp., Arctagrostis — sp., 74°3'40.1"N, 93°42'17.6"W, 40 m, 7 July 2013, P Sokoloff 123 (CAN 603359); north-facing ridge west of Arctic Watch Lodge, Eriophorum—Arctagrostis meadow, 74°4'17.5"N, 93°49'17.2"W, 16 m, 9 July 2013, P. Sokoloff 173 (CAN 603360, US); south end of Sunday Lake, 6 km south of Arctic Watch Lodge, mossy knoll in wet tundra, 74°1'7.8"N, 93°45'41.5"W, 66 m, 11 July 2013, PR Sokoloff 215 (CAN 603361).
Festuca baffinensis Polunin (Baffin Island Fescue, fétuque de Baffin) [Asian—amphi-Beringian—North American—amphi-Atlantic] — NUNAVUT: Somerset Island, site no. Z-25, level lacustrine well- to imper- fectly drained silt plain, grass-saxifrage foxhole mound, 74°2'N, 93°30'W, 53 m, 1 July 1975, S. C. Zoltai 751153 (DAO 137687).
Festuca brachyphylla Schultes & Schultes f. (Short- leaved Fescue, fétuque a feuilles courtes, Ivilsugait) [circumpolar-alpine] — NUNAVUT: Somerset Island, con- fluence of Cunningham River and stream immediately south of Arctic Watch Lodge at Cunningham River crossing, rocky talus on south-facing slope, with bryo- phytes, Cerastium arcticum, 74°4'2.9"N, 93°48'31.7"W, 58 m, 8 July 2013, P. Sokoloff 146 (CAN 603362).
Festuca edlundiae S.G. Aiken, Consaul & Lefko- vitch (Edlund’s Fescue, fétuque d’Edlund) [amphi- Beringian—North American—amphi-Atlantic] ~ Nunavut: Somerset Island, west ridge overlooking Gull Canyon, 4 km east of Arctic Watch Lodge, dry rocky scree, with Xanthoria elegans, Saxifraga opposi- tifolia, 74°3'42.4"N, 93°40'47.8"W, 50 m, 7 July 2013, P. Sokoloff 128 (CAN 603363).
Poa abbreviata R. Brown ssp. abbreviata (Dwarf Bluegrass, paturin court) [nearly circumpolar] — NUNAVUT: Somerset Island, scarce on moist calcare- ous slope, 74°6'N, 93°51'W, 11 August 1958, D.B.O. Savile 3780 (DAO 57589): Cunningham Inlet, 8 km from inlet, uplands, polar desert, 74°06'N. 93°5S'W, 225 m, 20 July 1976, L.C. Bliss s.n., (ALTA S6727): Cunningham Inlet, 8 km from inlet, uplands, polar
2015
desert, coastal lowlands, 74°06'N, 93°55'W, 30 m, 23 July 1976, L.C. Bliss s.n., (ALTA 56716).
Poa arctica R. Brown (Arctic Bluegrass, paturin arctique) [circumpolar-alpine] — NUNAVUT: Somerset Island, Flat Rock Falls, east coast of Cunningham Inlet, 4 km northeast of Arctic Watch Lodge, wet snow-patch community at foot of packed snowbank, near foot of falls, 74°5'56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P. Sokoloff 198 (CAN 603364).
Puccinellia bruggemannii T.J..Serensen (Prince Patrick Alkaligrass, puccinellie de Bruggemann) [North American] — NUNAVUT: Somerset Island, Cunningham Inlet, 1.6 km from inlet, uplands, polar desert, sedge meadow, 74°06'N, 93°55'W, 20 m, 22 July 1976, L.C. Bliss s.n., (ALTA 56719); on wet gravel and clay soil, 74°6'00"N, 93°51'00"W, 63 m, July 13, 2004, L. Consaul & A. Archambault 3083 (CAN 603396).
Puccinellia vahliana (Liebmann) Scribner & Mer- rill (Vahl’s Alkaligrass, puccinellie de Vahl) [North American—amphi-Atlantic] — NuNAvuT: Somerset Island, on wet gravel and clay soil, 74°6'00"N, 93°51'00"W, 63 m, July 13, 2004, L. Consaul & A. Archambault 3082 (CAN 603395); north-facing slope west of Arctic Watch Lodge, wet rocky seep in Dryas— Eriophorum tundra, with Dryas integrifolia, Salix arctica, Draba corymbosa, Cerastium arcticum, 74°4'17.5"N, 93°49'17.2"W, 16 m, 9 July 2013, P Sokoloff 189 (CAN 603366).
Trisetum spicatum (L.) K. Richter (Narrow False Oats, triséte a epi, Iviit iviksugait) [circumpolar- alpine] — NuNAvuT: Somerset Island, confluence of Cunningham River and stream immediately south of Arctic Watch Lodge at Cunningham River crossing, rocky talus on south-facing slope, with bryophytes, Cerastium arcticum, 74°4'2.9"N, 93°48'31.7"W, 58 m, 8 July 2013, P. Sokoloff 144 (CAN 603370).
Polygonaceae
Bistorta vivipara (L.) Delarbre (Alpine Bistort, renouée vivipare, Sapangaralannguat) [circumboreal- polar] — NUNAVUT: Somerset Island, Cunningham Inlet, 1.6 km from inlet, uplands, polar desert, sedge meadow, 74°06'N, 93°55'W, 20 m, 22 July 1976, L.C. Bliss s.n. (ALTA 56713); Flat Rock Falls, east coast of Cunning- ham Inlet, 4 km northeast of Arctic Watch Lodge, wet Dryas tundra with snow-bed community, with Sax- ifraga oppositifolia, Salix arctica, 74°5'56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P. Sokoloff 209 (CAN 603371).
Oxyria digyna (L.) Hill (Mountain Sorrel, oxyrie de montagne, Qunguliit) [circumpolar-alpine] — NUNAVUT: Somerset Island, south end of Sunday Lake, 6 km south of Arctic Watch Lodge, mossy knoll in wet tundra, 74°1'7.8"N, 93°45'41.5"W, 66 m, 11 July 2013, P Sokoloff 212 (CAN 603372); mud flats on alluvial slope to Cunningham River, south of Sunday lake, 2 km south of Arctic Watch Lodge, mud flats alongside river, 73°59'16.5"N, 93°41'38.7"W, 40 m, 11 July 2013, 2
Sokoloff 237 (CAN 603373).
SOKOLOFF: FLORA OF CUNNINGHAM INLET 33
Rosaceae
Dryas integrifolia Vahl (Entire-leaved Mountain Avens, dryade a feuilles enti¢res, Malikkaat) [amphi- Beringian—North American] — NUNAvUT: Somerset Is- land, Cunningham Inlet, 0.16 km from inlet, uplands, polar desert, coastal lowland beach ridges, 74°06'N, 93°55'W, 30 m, 23 July 1976, L.C. Bliss s.n. (ALTA 56730); north-facing ridge west of Arctic Watch Lodge, Eriophorum—Arctagrostis meadow, 74°4'27.7"N, 93°50'39.4"W, 118 m, 9 July 2013, PR Sokoloff 174 (CAN 603374); 2 km south of point at Cape Anne, muddy wet ground in marshy field alongside river, with Saxifraga oppositifolia, Salix arctica, Dryas integri-
folia, bryophytes, 74°6'23.3"N, 94°23'44.25"W, 26 m,
10 July 2013, P. Sokoloff, Z. Halem 240 (CAN 603375).
Potentilla pulchella R. Brown (Pretty Cinquefoil, potentille jolie) [circumpolar] — NUNAVUT: Somerset Island, lemming mound at top of hill at south end of Sunday Lake, 7 km south of Arctic Watch Lodge, lush lemming mound, with Arctagrostis sp., Sabulina rubel- la, 74°0'38.8"N, 93°44'30.4"W, 49 m, 11 July 2013, P. Sokoloff 217 (CAN 603376); south end of Sunday Lake, north of Cunningham River, 7 km south of Arc- tic Watch Lodge, lemming mound in middle of mud flats, 74°0'23.3"N, 93°42'43.5"W, 41 m, 11 July 2013, P. Sokoloff 227 (CAN 603377).
Salicaceae
Salix arctica Pallas (Arctic Willow, saule arctique, Suputiit, Suputiksaliit, Uqaujait) [circumpolar-alpine] — NUNAVUT: Somerset Island, Cunningham Inlet, 1.6 km from inlet, uplands, polar desert, coastal lowland beach ridges, 74°06'N, 93°55'W, 30 m, 23 July 1976, L. C. Bliss s.n. (ALTA 56718); dry stone ridge immediately west of Arctic Watch Lodge, in gravel scree, dry slaty gravel, with Saxifraga oppositifolia, Salix arctica, Papaver sp., 74°4'20.2"N, 93°49'22.6"W, 20 m, July 6, 2013, P. Sokoloff 113 (CAN 603379); dry stone ridge immediately west of Arctic Watch Lodge, in gravel scree, dry slaty gravel, with Saxifraga oppositifolia, Salix arctica, Papaver sp., 74°4'20.2"N, 93°49'22.6"W, 20 m, July 6, 2013, PR Sokoloff 114 (CAN 603380); gravelly scree ledge above Cunningham River, direct- ly adjacent to Arctic Watch Lodge (north side), rocky talus irrigated by water pipe, with Draba corymbosa, 74°4'10.8"N, 93°48'37.4"W, 58 m, 9 July 2013, P Sokoloff 164 (CAN 603381); wet snowmelt valley at base of large unnamed mountain northwest of Arctic Watch Lodge, wet sedge meadow emerging from melt- ing snowbank, with Ste/laria longipes, Cerastium arc- ticum, bryophytes, 74°4'31.3"N, 93°S1'1.7"W, 122 m, 9 July 2013, PR Sokoloff 178 (CAN 603382); garden spot below sewage lagoon at Arctic Watch Lodge, lush green patch in rocky scree, with Salix arctica, Saxifra- ga oppositifolia, Papaver sp., Parrya arctica, bryo- phytes, 74°4'13.1"N, 93°48'55.8"W, 16 m, 9 July 2013, P. Sokoloff 188 (CAN 603383); 2 km south of point at Cape Anne, muddy wet ground in marshy field along- side river, with Saxifraga oppositifolia, Salix arctica,
34 THE CANADIAN FIELD-NATURALIST
Dryas integrifolia, bryophytes, 74°6'23.3"N, 94°23'44.25"W, 26 m, 10 July 2013, PR Sokoloff, Z. Halem 238 (CAN 603384); garden spot below sewage lagoon at Arctic Watch Lodge, lush green patch in rocky scree, with Salix arctica, Saxifraga oppositifolia, Papaver sp., Parrya arctica, bryophytes, 74°4'13.1"N, 93°48'55.8"W, 16 m, July 12, 2013, P Sokoloff, A. Manik 243 (CAN 603385).
Saxifragaceae
Micranthes nivalis (L.) Small (Snow Saxifrage, sax- ifrage des neiges) [circumpolar-alpine] — NUNAVUT: Somerset Island, Flat Rock Falls, east coast of Cun- ningham Inlet, 4 km northeast of Arctic Watch Lodge, wet snow-patch community at foot of packed snow- bank, near foot of falls, 74°5'56.1"N, 93°44'18.5"W, 55 m, 10 July 2013, P Sokoloff 203 (CAN 603386).
Saxifraga cernua L. (Nodding Saxifrage, saxifrage penchee, Nunaraq qupanuap niqinga) [circumpolar- alpine] — NUNAVUT: Somerset Island, western cliff wall of Gull Canyon, below Gull nesting area, wet rocks directly under water seeps and waterfall, dense lush vegetation, with Saxifraga cespitosa, Bistorta vivipara, bryophytes, 74°3'48.4"N, 93°40'57.2"W, 40 m, 7 July 2013, PR. Sokoloff 133 (CAN 603387); gravelly scree ledge above Cunningham River, directly adjacent to Arctic Watch Lodge (north side), rocky talus irrigated by water pipe, with Draba corymbosa, 74°4'10.8"N, 93°48'37.4"W, 58 m, 8 July 2013, P Sokoloff 154 (CAN 603388).
Saxifraga cespitosa L. (Tufted Saxifrage, saxifrage cespiteuse) [circumpolar-alpine] — NUNAVUT: Somerset Island, Cunningham Inlet, 0.16 km from inlet, uplands, polar desert, coastal lowland beach ridges, 74°06'N, 93°55 W, 30im, 23 July 1976, L.C Bliss sn. (ALTA 56722); dry stone ridge immediately west of Arctic Watch Lodge, in gravel scree, wet seep under boulder in shale and muddy rock, with Cerastium arcticum, Papaver sp., Saxifraga oppositifolia, 74°4'21.3"N, 93°50'30.7"W, 122 m, July 6, 2013, P Sokoloff 116 (CAN 603389, NFM); Flat Rock Falls, east coast of Cunningham Inlet, 4 km northeast of Arctic Watch Lodge, wet snow-patch community at foot of packed snowbank, near foot of falls, 74°5'56.1"N, 93°44'18.5"W, 55m, 10 July 2013, P. Sokoloff 200 (CAN 603390); garden spot below sewage lagoon at Arctic Watch Lodge, lush green patch in rocky scree, with Salix arc- tica, Saxifraga oppositifolia, Papaver sp., Parrya arctica, bryophytes, 74°4'13.1"N, 93°48'55.8"W, 16 m, July 12, 2013, P. Sokoloff 246 (CAN 603391).
Saxifraga oppositifolia L. (Purple Mountain Sax- ifrage, saxifrage a feuilles opposées, Aupilattunnguat) [circumpolar-alpine] — NUNAvuT: Somerset Island, com- mon especially on moist calcareous gravel slopes., T4°6'N, 93°51'W, 11 August 1958, D.B.O. Savile 3785 (DAO 886692); Cunningham Inlet, 0.16 km from inlet, uplands, polar desert, 74°06'N, 93°55'W, 20 m, 23 July 1976, L.C. Bliss s.n. (ALTA 56732); Cunning- ham Inlet, 8 km from inlet, uplands, polar desert, Som-
Vol. 129
erset plateau, 74°06'N, 93°55'W, 200 m, 20 July 1976, L.C. Bliss s.n. (ALTA 56731); west ridge overlooking Gull Canyon, 4 km east of Arctic Watch Lodge, dry rocky scree, with Xanthoria elegans, Festuca sp., 74°3'42.4"N, 93°40'47.8"W, 50 m, 7 July 2013, P. Sokoloff 127 (CAN 603392); wet snowmelt valley at base of large unnamed mountain northwest of Arctic Watch Lodge, dry mud and clay mound in wet turfy tundra, with Festuca sp., 74°4'31.3"N, 93°51'1.7"W, 122 m, 9 July 2013, P Sokoloff 181 (CAN 603393, NFM).
Saxifraga flagellaris ssp. platysepala (Trautvetter) A.E. Porsild (Spider Saxifrage, saxifrage a sépales larges, Kakillarnaliit) [circumpolar] —- NUNAVUT: Som- erset Island, mossy wet area outside whale biologist’s cabin, 1.5 km north of Arctic Watch Lodge, wet mossy rocks, with Saxifraga cernua, Luzula confusa, Draba sp., Salix arctica, 74°4'58.8"N, 93°50'2"W, 94 m, 8 July 2013, P. Sokoloff 156 (CAN 603394).
Discussion
Although five botanical collecting trips have taken place on Cunningham Inlet, the four trips previous to this study were focused primarily on selective collect- ing or ecological assessment. Combining these data with the 93 vascular plant specimens collected in 2013 pro- vides a relatively complete inventory of the vascular plants of Cunningham Inlet.
Although the 48 vascular plant species documented is a relatively low number in terms of the species diver- sity over the entire island (40% of those reported in Aiken et al. 2007, 48% of those in Savile 1959, and 64% of those in Woo and Zoltai 1959), it is important to consider that Cunningham Inlet is well within the “cryptogam-—herb barren” vegetation unit described in the Circumpolar Arctic Vegetation Map, which covers only roughly half of the island (Walker et al. 2005). The dominant growth forms described for this vegeta- tion unit include “Cushion forbs: Papaver dahlianum ssp. polare; Draba; Potentilla hyparctica; Saxifraga oppositifolia” and “Graminoid: Alopecurus alpinus; Deschampsia borealis/brevifolia; Poa abbreviata: Puc- cinellia angustata; Phippsia; Luzula nivalis: Luzula confusa”’ (CAVM Team 2003*), nearly all of which are documented in this paper. Cunningham Inlet is also classified by the Circumpolar Arctic Vegetation Map Team as belonging to Arctic Bioclimate Subzone B. where prostrate dwarf shrubs (such as Arctic Willow) are the dominant growth form, and the number of species is estimated to run from 50 to 100 depending on the site (CAVM Team 2003*). Considering the ecol- ogy of Cunningham Inlet’s polar desert and the five collecting trips to this site, it seems highly likely that we have documented all the vascular plants at this inlet and that the remaining species known for Somerset Island occur within the other ecosystems found on the island.
2015
SOKOLOFF: FLORA OF CUNNINGHAM INLE1
ws) WN
FiGURE 3: Edlund’s Fescue (Festuca edlundiae S.G. Aiken, Consaul & Lefkovitch) in habitat at the top of cliffs overlooking Gull Canyon, Somerset Island, Nunavut (Sokoloff 128). Photo by P. Sokoloff.
The summer of 2013 was unusually cold and late in the Canadian high Arctic (NASA 2013*), which de- layed the flowering time of many species we encoun- tered. Many species found on the open tundra had just begun their yearly growth, and the Purple Mountain Saxifrage, a benchmark spring ephemeral species in the Arctic, was still in full bloom when I left Cunning- ham Inlet. Although this resulted in taxonomically use- ful plant specimens rarely collected in flower (1.¢., Sax- ifraga oppositifolia and Salix arctica), care should be taken to look for late-flowering specimens and speci- mens with fruits on subsequent visits to Cunningham Inlet. Thus, although comparisons between past and current vascular plant communities at this site are im- possible given the paucity of earlier collections, in the future Cunningham Inlet could be used to monitor floristic change in the high Arctic using this inventory as a baseline, while keeping an eye out for any additions
to the flora that would have been missed because of
their later flowering time.
Two vascular plant species reported here, Festuca edlundiae (Figure 3) and Draba glabella, have not been reported before for Somerset Island (Savile 1959; Aiken et al. 2007). Festuca edlundiae is a widespread Cana- dian high Arctic endemic, found throughout the north- em part of the archipelago, including Cornwallis and Prince of Wales Islands, adjacent to Somerset Island (Aiken et al. 2007). This species has only recently
been recognized as a distinct taxon within the Cana- dian high Arctic Festuca brachyphylla complex (Aiken et al. 1995). Dwarf plants with a heavily marcescent habit, Festuca edlundiae, were first differentiated from the phenotypically variable Festuca hyperborea Hol- men ex Frederiksen (High Arctic Fescue) based on isozymes (Aiken ef al. 1995). Hybridization and intro- gression between Festuca edlundiae and the other high Arctic Festuca species have been documented (Saarela et al. 2013), but taxonomic boundaries between the various species are well understood and various keys exist separating the species using consistent morpho- logic characters (Fjellheim ef a/. 2001; Guldahl et al. 2001). Using these keys, it may yet be found that Fes- tuca edlundiae has been collected on Somerset Island before its recognition as a distinct taxon; nonetheless Sokoloff 128 is the first known report of this grass species on the island.
Draba glabella (Sokoloff 134) is a first collection for both Somerset Island and the central Canadian Arctic archipelago. This species is common and widespread within the archipelago, but although its distribution extends from Banks to Baffin Island (east to west) and from the mainland to Ellesmere Island (south to north), it is absent from the central Arctic islands, including Bathurst, Prince William, and Cornwallis (Aiken et al. 2007), and has been reported only once on the west coast of Devon Island by Polunin (1940;
36 THE CANADIAN FIELD-NATURALIST
Vol. 129
FiGURE 4: Habitat of Smooth Draba (Draba glabella Pursh) at Gull Canyon, Somerset Island, Nunavut. Draba glabella (Sokoloff 134) was collected at the base of the cliffs on the right side of the canyon. Photo by P. Sokoloff.
a second report in this volume is apparently a typo, a repetition of a collection made on Sugluk Island just off the coast of Quebec). These central islands are pri- marily polar desert (including the study site at Cun- ningham Inlet) and consist of shattered limestone with minimal tundra cover (Savile 1959; Bliss et al. 1984).
Accordingly, we found Draba glabella growing at only a single site: in wet moss and rich soil at a gull colony (Figure 4). Burt (2000) and Polunin (1940) in- dicate that such damp, nutrient- and soil-rich cliffs are ideal habitat for Draba glabella. Our discovery of this species at a bird colony in the middle of a gap in its range (Aiken ef al. 2007) either points to bird-borne dispersal of the plant or indicates that this colony may serve as a refuge for this species in otherwise inhos- pitable and nutrient-poor habitat (Odasz 1994). In either case, other bird colonies within the central Canadian polar desert could harbour this widespread species as well.
The four identified species of marine algae, Fucus distichus, Alaria esculenta, Agarum cribrosum, and Sacharrina latissima, are all previously known to occur in the Barrow Straight, and both Alaria esculenta and Fucus distichus have been previously collected in Cun- nigham Inlet (Lee 1980). Nostoc commune, known to be common in the Canadian high Arctic (Polunin 1947; Lennihan et al. 1994; Sheath et al. 1996), is poorly
represented in Canadian algal collections (CANA, data available through Canadensys). Thus, Sokoloff 175 and 176 are likely the first known specimens of Nostoc commune from Somerset Island.
Although our 20 lichen and 13 bryophyte specimens greatly expand on known cryptogamic species from Somerset Island (Savile 1959), they were collected opportunistically and there are almost certainly gaps in the collection that a trained lichenologist or bryol- ogist could fill.
Acknowledgements
I am immensely grateful to the following people who provided identification for many specialized tax- onomic groups: Dr. Jeffery Saarela (Poacaeae, Cyper- aceae, Juncaceae), Paul Hamilton (algae), Colin Free- bury (lichens), Jennifer Doubt (bryophytes), Dr. Lynn Gillespie (Papaver, Potentilla), Etienne Léveillé- Bourret (Potentilla), and Gerry Mulligan (Draba). I thank Micheline Beaulieu-Bouchard, Jennifer Doubt. Amaina Boloron, Laura Smyk, and the Canadian Mu- seum of Nature’s botany volunteers for processing this collection. I thank Dr. Jeff Saarela for thoughtful com- ments on this manuscript. I also thank Gisele Mitrow and Amanda Ward for providing helpful access to the Agriculture and Agri-Food Canada collection of vas- cular plants. Comments by Dr. Jacques Cayouette and
2015
Dr. Paul Catling greatly enhanced earlier drafts of this work. This fieldwork was carried out as part of Arctic Watch Lodge’s 2013 Steve Amarualik Youth Leader- ship Expedition and would not have been possible without the generous support of Arctic Watch Lodge and the Canadian Museum of Nature. In particular, | thank Richard Weber, Josée Auclair, Tessum Weber, Nansen Weber, and Ewan Affleck for organizing an exceptional expedition for the youth participants, for hosting me at Arctic Watch Lodge, and for going out of their way to contribute to the success of the collect- ing portion of the trip. Zach Halem and Alicia Malik both contributed as able collectors, and I commend all the expedition’s youth — Alicia, Anika, Belinda, Louis, Savannah, Simeonie, Ryan, Sky, and Zach — for their enthusiasm, curiosity, and assistance. This research was conducted under Nunavut Department of Environment Wildlife research permit WL 2013-026.
Documents Cited (marked * in text)
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Elven, R., D. F. Murray, V. Y. Razzhivin, and B. A. Yurtsey. 2011. Annotated checklist of the panarctic flora (PAF): vas- cular plants. Panarctic Flora Project Steering Committee. Accessed 30 May 2014. http://www.nhm.uio.no/english /research/infrastructure/paf/
NASA (National Aeronautics and Space Administration). 2013. Arctic sea ice minimum in 2013 is sixth lowest on record. National Aeronautics and Space Administration, Washington, D.C., USA. Accessed 14 March 2014. http: /www.nasa.gov/content/goddard/arctic-sea-ice-minimum- in-2013-is-sixth-lowest-on-record/#. UyNMTj9dV8E
Woo, V. B., and S. Zoltai. 1977. Reconnaissance of the soils and vegetation of Somerset and Prince of Wales Islands, N.W.T. Information Report NOR-X-186. Northern Forest Research Centre, Edmonton, Alberta, Canada. 127 pages.
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Aiken, S. G., L. L. Consaul, and L. P. Lefkovitch. 1995. Festuca edlundiae (Poaceae), a high arctic, new species compared enzymatically and morphologically with similar Festuca species. Systematic Botany 20: 374-392.
Aiken, S. G., M. J. Dallwitz, L. L. Consaul, C. L. McJan- net, R. L. Boles, G. W. Argus, J. M. Gillett, P. J. Scott, R. Elven, M. C. LeBlanc, L. J. Gillespie, A. K. Brysting, H. Solstad, and J. G. Harris, 2007. Flora of the Canadian Arctic Archipelago. NRC Research Press, National Research Council of Canada, Ottawa, Ontario, Canada. Accessed 18 February 2014. http://nature.ca/aaflora/data.
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Bliss, L. C., J. Svoboda, and D. I. Bliss. 1984. Polar deserts, their plant cover and plant production in the Canadian high Arctic. Holarctic Ecology 7: 305-324.
Burt, P. 2000. Barrenland Beauties: Showy Plants of the Canadian Arctic. Outcrop Ltd. Yellowknife, Northwest Ter- ritories, Canada. 238 pages.
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Received 25 March 2014 Accepted 17 June 2014
Osmia species (Megachilidae) Pollinate Cypripedium parviflorum (Orchidaceae) and Packera paupercula (Asteraceae): A Localized Case of Batesian Mimicry?
PAUL M. CATLING 170 Sanford Ave., Ottawa, Ontario K2C 0E9 Canada; email: catlingp@agr.ge.ca
Catling, Paul M. 2015. Osmia species (Megachilidae) pollinate Cypripedium parviflorum (Orchidaceae) and Packera paupercula (Asteraceae): a localized case of Batesian mimicry? Canadian Field-Naturalist 129(1): 38-44.
The pollinator-non-rewarding Large Yellow Lady’s-slipper (Cypripedium parviflorum var. pubescens [Willdenow] Knight) and the rewarding Balsam Ragwort (Packera paupercula [Michaux] A. Léve & D. Léve) exist together in some eastern Ontario alvars where they share a group of medium-sized, dark blue metallic pollinators in the bee genus Osmia (Mason Bees, Megachil- idae). I provide evidence of floral mimicry of the ragwort by the orchid based on several observations: (1) Osmia visiting P. paupercula also visit C. parviflorum var. pubescens; (2) Osmia are more frequent visitors to P paupercula than to other co- blooming plants; (3) Osmia are the primary pollinators of C. parviflorum var. pubescens; (4) the behaviour of Osmia on the flower of C. parviflorum var. pubescens involving landing on the staminode suggests mimicry; and (5) the largest populations of C. parviflorum var. pubescens in Ontario are in alvar landscapes where P. paupercula is abundant. Pollination in Large Yellow Lady’s-slipper may vary geographically from non-model to species-specific Batesian mimicry, which is rare in orchids. This latter pollination mechanism may have evolved in ice-front environments during the Pleistocene, but then became isolated to unusual partial analogues of ice-front habitat during the Holocene with pollination in most of the current range appearing to be a generalist strategy.
Key Words: Cypripedium parviflorum var. pubescens; Packera paupercula, Osmia; Large Yellow Lady’s-slipper; Balsam
Ragwort; pollination; mason bee; floral mimicry; Ontario; alvar
Introduction
The Large Yellow Lady’s-slipper (Cypripedium parv- iflorum var. pubescens [Willdenow] Knight, previously recognized as C. calceolus L. var. pubescens {Willde- now] Correll) is reported to be pollinated by a variety of small bees (Argue 2012). Observations of only Mason Bees (Osmia spp.) pollinating the yellow, pollinator- rewarding Balsam Ragwort (Packera paupercula |Mi- chaux] A. Léve and D. Léve) and the yellow, non- rewarding (i.e., deceptive) Large Yellow Lady’s-slipper in the Ottawa Valley led to consideration of the extent to which the orchid may be an example of localized Ba- tesian floral mimicry, where a species with non-reward- ing flowers has evolved floral characteristics of a re- warding species that confer an advantage of increased pollinator visitation (Dafni 1984).
Floral mimicry has been reported in a number of orchids (e.g., Dafni and Irvi 1981; Gigord et al. 2002; Pellegrino et al. 2008; Edens-Meier et al. 2014) and in some Asian species of Cypripedium (Sugiura et al. 2002; Li et al. 2006) and has been recently inferred in the North American Ram’s-head Lady’s-slipper (Cypri- pedium arietinum R. Brown; Catling and Kostiuk 20S):
For Large Yellow Lady’s-slipper and Balsam Rag- wort to be an example of Batesian floral mimicry, five criteria discernible in field study should be considered, apart from the fact that the Balsam Ragwort model and the orchid mimic occur locally together: (1) Mason Bees visiting the ragwort also visit the orchid; (2) Mason
Bees are more frequent visitors to Balsam Ragwort than to other co-blooming plants so that a specific model is suggested; (3) Mason Bees are primary pollinators of the orchid, i.e., the primary insect group transferring orchid pollen from one flower to another; (4) the behav- iour of Mason Bees on the orchid flowers suggests mimicry; and (5) the orchid achieves higher abundance in the presence of Balsam Ragwort than in its absence, due in part to higher fecundity.
The present work focuses on the first four criteria listed above and considers support for the fifth. The work is centred on alvar landscapes in the Ottawa Val- ley that are diverse, rich in restricted species, and con- tain globally imperiled ecosystems (Catling 1995).
Study Area
The three study areas, all in eastern Ontario, included Marlborough Forest (45.0818°N, 75.8099°W), Brae- side Alvar (45.4864°N, 76.4544°W), and Burnt Lands Alvar (Burnt Lands Provincial Park: 45.2530°N. 76.1486°W). The latitudes and longitudes represent ap- proximate locations, and all observations and collections were made within | km of these sites. The specific study areas at each site were approximately 2 ha in extent and included 200-300 plants of Large Yellow Lady’s-slipper and 200-700 plants of Balsam Ragwort. Large popula- tions of Large Yellow Lady’s-slipper occurred in the surrounding area, including up to 1000 plants within an area of | km’. All three sites were dominated by more or less open woodlands of Eastern White Cedar (7) huja
2015
occidentalis L.) with some Balsam Fir (Abies balsamea [L.] Miller) and White Pine (Pinus strobus L.).
Methods Gathering pollination data and voucher specimens
I gathered data by direct observation and collection of bees on orchid and ragwort flowers. On all days when observations of pollinators were made it was sunny and 20—24°C. Areas where pollinators were ob- served or collected were less than 0.4 ha in extent.
Examining bees reveals pollination because the ad- hesive pollen of the orchid flower is smeared onto the dorsal surface of the thorax as the bee leaves one of the basal side openings of the flower and can be seen on the insect’s thorax for several days afterward (personal ob- servation) indicating that it was at least a visitor to a Large Yellow Lady’s-slipper flower and a potential pol- linator.
Bees with Cypripedium pollen smears in the study areas were assumed to be pollinators of Large Yellow Lady’s-slipper. Although Ram’s-head Lady’s-slipper is present and blooms at the same time (or starts slightly earlier), robust bees (such as Mason Bees) that are over 8 mm long are likely too large to enter and exit the flowers of that species and have not been implicated in its pollination (Catling and Kostiuk 2013). The pollen of these two orchids cannot be differentiated by simple light microscopy using a double-staining technique with phloxine and methyl green (personal observation).
Specimens of bees collected as part of this study are in the collection of Dr. Laurence Packer at York Univer- sity, Toronto, Ontario, Canada, and that of Dr. Cory Sheffield at the Royal Saskatchewan Museum, Regina, Saskatchewan, Canada. Dr. Sheffield identified Osmia subaustralis Cockerell, 1900 and O. albiventris Cres- son, 1864 and Dr. Anna Taylor identified Osmia prox- ima Cresson, 1864.
Mason Bees visiting the orchid as well as the ragwort
On both | and 2 June 2011, I spent approximately an hour observing pollination of Balsam Ragwort in a patch of 400 plants at Braeside. On 3 and 4 June ZOU, | observed a group of approximately 600 plants of Bal- sam Ragwort for approximately 1.5 h on the Burnt Lands. In all cases, ragwort flowers were within 100 m of Large Yellow Lady’s-slipper plants. Observations were made between 11 a.m. and 12:30 p.m. I collected and examined bees visiting Balsam Ragwort for orchid pollen smears on the dorsal thorax and released them at the end of the observation period.
Mason Bees visiting other co-blooming plants
On 9 May and | and 2 June 2011, at each of the three study sites, I spent an hour between 10 a.m. and 2 p.m. determining whether Mason Bees were visiting the only other species blooming in the area: the introduced Com- mon Dandelion (Zaraxacum officinale F. H. Wiggers), Swamp Dandelion (7° palustre [Lyons] Symons , and Wild Strawberry (Fragaria virginiana Miller).
CATLING: OSMIA SPECIES POLLINATE CYPRIPEDIUM PARVIFLORUM AND PACKERA 39
Mason Bees, the primary pollinator of the orchid
On 29 May 2011, | attempted to observe pollinators at Braeside. I spent 3 h watching a group of 30 flowers of Large Yellow Lady’s-slipper during sunny, mild weather between |] a.m. and 2 p.m. On 2 and 11 June 2008, | spent 2 h each at Marlborough Forest and the Burnt Lands observing pollination in a group of 25 and 50 flowers, respectively.
Behaviour of Mason Bees suggesting mimicry
On 4 June 2011, between 9 a.m. and noon, I conduct- ed eight experiments with three Osmia subaustralis (one male, two females) and five unidentified Osmia. The bees had been isolated and maintained in captivity, resting at low light and low temperature (20°C) for 20 h. As a result of their inactivity, they were each eas- ily transferred into a fine mesh cage containing two fresh flowers of Large Yellow Lady’s-slipper. The 16 orchid flowers selected for the observations had lips 31-40 mm long with more or less circular distal open- ings with a maximum width of 8-11 mm (Figure 1) and showed no signs of visitation in the form of scales, hairs or displaced pollen. The maximum width of basal open-
FIGURE 1. Osmia proxima Cresson, 1864 exiting one of the
basal lateral openings of a flower of Large Yellow Lady’s-slipper (Cypripedium parviflorum var. pubes- cens [Willdenow] Knight). The anther is on top of the thorax of the bee. Photo by P. M. Catling, Burnt Lands Provincial Park, 11 June 2008.
40 THE CANADIAN FIELD-NATURALIST
ings was 2-3 mm becoming 3—5 mm when the lip was pressed downward. The flower stems were placed up- right in water in a coffee cup, through a small opening in the lid; this prevented bees from falling into open water. Observations were made through the glass front after strong light and heat rising to 24°C was focused on the cage and led to activity. An experiment was con- sidered to be completed when a bee became inactive or when it appeared intent on escape and showed little interest in the flowers.
Results
Of 76 bees captured while they were visiting Balsam Ragwort, 25 carried pollen likely originating from Large Yellow Lady’s-slipper (Table 1).
Both co-blooming dandelion species were past peak flowering and infrequent. Wild Strawberry was infre- quent but in peak bloom locally. Although many bees were seen on the flowers of these plants only one Mason Bee was seen visiting them — a flower of Wild Straw- berry.
All observations of pollination are summarized in Table 2, which includes seven pollinator and three vis- itor records, all of which involved Mason Bees. In most cases, the bees were captured after they left one flower
VoL 129
and entered another; they were removed from the sec- ond flower. Apart from the fact that flowers were en- tered within a few seconds, there were no detailed ob- servations of behaviour, except in the following two cases.
At Burnt Lands Alvar on 11 June 2008 on a sunny morning with air temperature 18°C, a bee was found inside the lip of a Large Yellow Lady’s-slipper flower. The plant was covered with a net and within five min- utes the bee exited the flower from a lateral basal open- ing with pollen on the thorax (Figure 1). It was later identified as Osmia proxima.
At Marlborough Forest on 2 June 2008 on a sunny late afternoon with air temperature 22°C, after a cloudy morning, a bee landed on the staminodium of a Large Yellow Lady’s-slipper flower, then fell into the lip cav- ity. It emerged seven minutes later, with much pushing downward of the lip, from a lateral opening with pollen on its back. It was later identified as Osmia proxima.
During experiments to investigate mimicry, the bees generally became active within 10 minutes and flew around inside the cage. Within 1-18 minutes of flight, nine of ten bees landed on the staminodium of a cen- trally located Large Yellow Lady’s-slipper flower. They then turned back and forth over the staminodium
TABLE I. Observations of Mason Bees (Osmia spp.) carrying pollen of Large Yellow Lady’s-slipper (Cypripedium parviflorum
var. pubescens [Willdenow] Knight).
OO TTT S55 ——————————————————__————_
No. of bees with pollen
(total bees observed) Location Collected on/in Date
5 (20) Braeside Balsam Ragwort (Packera paupercula [Michaux] 1 June 2011 A. Love and D. Léve)
9 (20) Braeside Balsam Ragwort 2 June 2011
5 (20) Burnt Lands Balsam Ragwort 3 June 2011
6 (16) Burnt Lands Balsam Ragwort 4 June 2011
EEE,
TABLE 2. Observations of Mason Bees (Osmia spp.) pollinating or visiting Large Yellow Lady’s-slipper (Cypripedium parv-
iflorum var. pubescens [Willdenow] Knight).
Bee species Location/origin
1 Osmia proxima Cresson, 1864 Burnt Lands
1 Osmia sp. Burnt Lands
Notes
Date
Pollinator. Found inside lip, exited with pollen.
Pollinator. Leaving flower with pollen and
11 June 2008
11 June 2008
entering another.
1 Osmia sp. with pollen.
Marlborough 3 June 2008
Visitor, leaving flower
| Osmia proxima Cresson, 1864 Marlborough Pollinator. Landed on staminodium with pollen 3 June 2008 then exited flower with pollen.
2 cf. Osmia albiventris
Cresson, 1864 (male) Braeside Pollinators. Both visiting two consecutive 29 May 2011 flowers with pollen taken to both flowers. ;
3 Osmia subaustralis Cockerell,
1900 (1 female, 2 males) Braeside Pollinators. All three visiting two consecutive 29 May 2011 flowers with pollen taken to both flowers.
| Osmia sp. Braeside Visitor. Visiting a single flower and emerging 29 May 2011
with pollinia.
rE eEE£-290XY-- =
2015
probing it actively with the tongue. After 15—120 see- onds of this, all nine bees fell off the staminodium into the lip. Time to exit was usually 5 minutes, but was 35 minutes in one case. In a few instances, a bee was inactive inside the lip for 5-10 minutes. Time spent struggling in the exit area below the stigma was 3-10 minutes and, during this time, the bee often pushed the lip downward, evidently with its back against the col- umn to increase the space. Once the bee’s head had reached the side opening, exit from the flower, as it passed under the stamen, required only 2—7 seconds. This was true for seven of nine trapped bees. Of the oth- er two, one exited the side opening upside down with pollen attaching to the underside of the thorax and abdomen. Another bit a hole through one of the win- dows at the base of the lip and exited that way within 9 minutes after entering a flower. Most bees spent a few minutes grooming after leaving the flower and one had the wings stuck together, with pollen, over the back. Two bees went to, and through, the second flower with- in 5 minutes of visiting the first.
Discussion
The number of bees visiting Balsam Ragwort that carried pollen of Large Yellow Lady’s-slipper seemed high, based on low levels of capsule development in many areas suggesting low levels of insect visitation. However, the orchid was common in both areas (Brae- side and Burnt Lands) and generally flowered a little earlier than Balsam Ragwort; thus, it may have attract- ed attention when resources were limited.
Although the survey of Mason Bees on co-blooming plants included over 2 h of observation time, this is not considered an extensive survey. Nevertheless, it pro- vides evidence that Mason Bees were very much asso- ciated with Balsam Ragwort and less so with other flowering plants.
The 11 observations of Mason Bees and no other species of bees on Large Yellow Lady’s-slipper is eVi- dence that they were the primary pollinators at these study locations (Braeside and Burnt Lands alvars and Marlborough Forest). Although Mason Bees were the only pollinators in the present study, studies of the similar, although not closely related (Li et al. 2011), European Lady’s-slipper Orchid (C. ypripedium calceo- lus L.) have revealed a number of different bee polli- nators (Nilsson 1978, Antonelli e¢ al. 2009). For Large Yellow Lady’s-slipper in eastern North America, the only pollinators reported are halictid bees and small Carpenter Bees (Ceratina spp.), and these were based on few records (Argue 2012). I saw bees from both of these groups in the study areas and a significant bee fauna was present at least at the Burnt Lands site ( Tay- lor and Catling 2011), although not all of these species would be active at the time of blooming of the orchid. Many species of bees, including Mason Bees, have been reported as visitors to Large Yellow Lady’s-slipper (Argue 2012). Although this suggests that many bee
CATLING: OSMIA SPECIES POLLINATE CYPRIPEDIUM PARVIFLORUM AND PACKERA 4]
species may be involved in its pollination and some regional variation may be anticipated, Mason Bees may be the primary pollinators in some of the larger orchid populations in the study areas.
The pollination of slipper orchids is based on control of the path of the insect through the flower by morpho- logic attributes of the flower lip, where the insect enters the large frontal opening, deposits pollen, picks up a new pollen load, and then exits by one of the two small openings on either side of the lip base (Figure |). This well-known and accepted phenomenon was first elab- orated by Darwin (1862), but more correctly and in more detail in Darwin (1877) following help from Asa Gray, and it was later discussed by many others (e.g., Stoutamire 1967; Catling and Catling 1991; Argue 2012). What has been controversial is the attractant. Ideas have varied: the food value of hairs on the inner lip, the production of fragrances by the pollinating bees, the resemblance of the lip to a cavity nest site, and gen- eral food deception involving colour and nectar guides (Catling and Catling 1991; van der Cingel 2001). The latter of these has been the most agreed upon, but an interesting addition is that a flower may be more likely to be visited again if it has already been visited, as a result of accumulation of bee odours on specialized hairs (Nilsson 1978). The present work supports food deception, but further suggests that Mason Bees may be specifically attracted to the staminode due to its resem- blance to the centre of a Packera flower. The darker and orange areas in open disc flowers within a mass of yellow disc flowers in Balsam Ragwort are similar to orange spots on a yellow staminodium of similar size in the orchid. The only other case in Cypripedium where bees contacted the staminodium first before falling into the labellum involved C. guttatwm Swartz and mimicry was not implicated (Banziger er al. 2005).
A distribution map for Balsam Ragwort in Ontario (Catling 1995, Figure 7) indicates a concentration in alvar landscapes along the edges of the Canadian Shield. Similarly, a map for Large Yellow Lady’s-slip- per (Whiting and Catling 1986, map 3c) shows con- centrations in the same regions and absences from the Canadian Shield and parts of southwestern Ontario. Although the orchid is more widespread than the rag- wort, it is similarly associated with limestone rock (Whiting and Catling 1986). The areas of high abun- dance of the orchid in southern Ontario are well known and include Manitoulin Island, the Bruce Peninsula, and limestone landscapes near Kingston and Ottawa. For example, with regard to the Bruce Peninsula, the Bruce-Grey Plant Committee ( 1997) notes that “it may be more common [here] than in any other part of On- tario.” These same landscapes are the areas of abun- dance of Balsam Ragwort.
Conclusions
The following observations support the case for Batesian floral mimicry. A large proportion of Mason Bees visiting Balsam Ragwort also visited the orchid,
42 THE CANADIAN FIELD-NATURALIST
but not other co-blooming flowers. Mason Bees were the primary pollinators of the orchid and behaved on the orchid flowers as they did on the flowers of Balsam Ragwort by landing on the staminodium, which resem- bles the centre of the ragwort flower. Orchids were more abundant where the ragwort was abundant. Although it may be appropriate to consider ragwort species as important models in a generalized magnet species effect, pollination may vary geographically from non-model to species-specific Batesian mimicry where a single model can be readily identified. This lat- ter possibility is of interest for three reasons: the rarity of floral mimicry in orchids, the existence of specialist and generalist strategies within one taxon, and the pos- sibility of early evolution and mimicry in the past.
Floral mimicry is rare in orchids
Members of the genus Cypripedium have generally been regarded as generalist (non-model) food mimics (Catling and Kostiuk 2006; Pelligrino et al. 2008) like most food-deceptive orchids, which include a third of all orchids (Cozzolino and Widmer 2005), the largest family of vascular plants. Most deceptive orchids have not evolved species-specific Batesian floral mimicry (Johnson et al. 2003; Li et al. 2006), although it has been attributed to various species around the world, including Red Helleborine (Cephalanthera rubra [L.] Richard) from Europe (Nilsson 1983), Leopard Orchid (Diuris maculata Smith) from Australia (Beardsell et al. 1986), and Cluster Disa (Disa ferruginea [Thunb.] Swartz) from South Africa (Johnson 1994). Mimicry has been reported only recently in two of approximate- ly 50 species of Cypripedium, a genus of the northern hemisphere (Li e¢ al. 2011): a Japanese Lady’s-slipper (C. macranthos Sw. var. rebunense (Kudo) Miyabe & Kudo (Sugiura ef a/. 2002) and Ram’s-head Lady’s- slipper (C. arietinum R. Brown) (Catling and Kostiuk 2013).
A specialist and a generalist within one taxon Geographically based differences in food deception within taxa, ranging from specialization to generalist strategies, may be more frequent than is realized. The idea of such micro-ecological isolating mechanisms is not new and was discussed by Heslop-Harrison (1958) with regard to orchids and by Stoutamire (1967) with regard to lady’s-slippers. This within-taxon variation may not always be as obvious as the anomalous white- flowered var. rebunense of Cypripedium macranthos on Rebun Island, Japan, which visually mimics the white-flowered Pedicularis schistostegia Vvedensky (Sugiura 2001, 2002). The pollinator-mediated mating restriction that has been shown between the varieties of C. parviflorum (Case and Bradford 2009; Case and Bierbaum 2013) may also occur between geographic races. There is also evidence for different pollination races in European C. calceolus (Antonelli et al. 2009). That different pollination races exist in Large Yellow Lady’s-slipper is suggested by the 36 records of visita- tion and pollination by Mason Bees on alvar landscapes
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and no records of other bees, as reported here, com- pared with reports of many other kinds of visiting bees in other locations (van der Cingel 2001). The possibil- ity that some lady’s-slipper species, and other deceptive orchids, are both specialists and generalists requires more study.
Early evolution and mimicry in the past
Mimicry might have originated as a general re- semblance to co-occurring floral food sources, such as yellow blossoms. Although similar colours may not al- ways indicate mimicry or pollinator sharing (Bierzy- chudek 1981), they might. Supporting this is the obser- vation of the unusual abundance of Large Yellow Lady’s-slipper (thousands of plants in a few hectares) along with several co-blooming, yellow-flowered species in remnant prairies along the railway lines in southeastern Saskatchewan (Catling and Kostiuk 2006). Corolla colour is sufficient to achieve floral mimicry, and pollinators can select for it (Gigord et al. 2002). Yellow-flowered species prominent in the Saskat- chewan prairies are Hoary Puccoon (Lithospermum canescens [Michaux] Lehmann) and Heart-leaved Alexanders (Zizia aptera [A. Gray] Fernald), but spe- cies of ragwort are also present (personal observation). Following development of colour resemblance, the orchid may have evolved a more specific resemblance to Balsam Ragwort by developing reddish spots on the staminode and possibly in other ways. This may have occurred in the open, ever-changing ice-front environ- ments that lasted for many thousands of years during the Pleistocene.
More recently, during the Holocene, the mimetic pollination system may have become isolated to un- usual partial analogues of ice-front habitat with polli- nation in most of the current range appearing to be a generalist strategy. Thus we have a Batesian mimic that to a large extent became a generalist with only localized situations, as in the alvars studied here, that make the former pollination mechanism clear. Certainly, within the general range, there are areas of high abundance of Large Yellow Lady’s-slipper, and many of these are in places that resemble ice-front and early postglacial habitats, such as alvars (Catling and Brownell 1995), Regardless of the likelihood of this hypothesis, it does draw attention to the possibility that pollination in temperate plants may sometimes be better understood through reference to past conditions.
Future research
Although this articles presents evidence to suggest Batesian floral mimicry, the suggestion would be strengthened if the bees were found to carry only rag- wort pollen, rather than observing bees visiting only ragwort flowers, as sampling based on a longer period would be achieved in this way.
Two more protocols for data collection that can be considered in future studies are: (1) flowers of the orchid and ragwort can be compared under ultraviolet (UV) light to determine whether similar patterns exist,
2015
although floral mimicry is not entirely dependent on UV reflectance (Gigord et al. 2002); and (2) a bio- chemical analysis of scent would be helpful to deter- mine the extent to which that factor plays a role, al- though studies have suggested that visual attributes may be more important than scent chemistry in deceptive orchids (Jersakova et al. 2012). The scent components may be general or contain those of the ragwort or even those of the pollinating bees (Volterova er al. 2007). These protocols have been effectively applied to stud- ies of Batesian floral mimicry in other species (Edens- Meier ef al. 2014).
Acknowledgements
Dr. Cory Sheffield and Dr. Anna Taylor assisted with identification of bees. Dr. Peter Bernhardt provided helpful advice.
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Received 21 March 2014 Accepted 18 May 2014
Gastony’s Cliffbrake (Pellaea gastonyi) in Manitoba: New Records and Assessment of Conservation Status
CHRIS FRIESEN!:? and COLIN MuRRAY!
Manitoba Conservation Data Centre, Box 24, 200 Saulteaux Crescent, Winnipeg, Manitoba R3J 3W3 Canada
“Corresponding author: chris. friesen@gov.mb.ca
Friesen, Chris, and Colin Murray. 2015. Gastony’s Cliffbrake (Pellaea gastonyi) in Manitoba: new records and assessment of conservation status. Canadian Field-Naturalist 129(1); 45-52.
Gastony’s Clifforake (Pellaea gastonyi Windham) is a globally rare fern (Pteridaceae) that grows on limestone cliffs and ledges, including those associated with alvars. Until now, the only record in Manitoba was from a location just north of Fish- er Branch. We report additional records and locations, one of which is over 250 km north of the initial collection. We also provide a conservation status assessment of this species in Manitoba that indicates that this species is rare in the province and is threatened in a least a portion of its range by habitat loss and degradation.
Key Words: Gastony’s Cliffbrake; Pe/laea gastonyi; Manitoba; limestone; alvar; rare species; fern; habitat loss
Gastony’s Cliffbrake (Pellaea gastonyi Windham) is a fern (Pteridaceae) of calcareous outcrops and cliffs (Windham 1993a). Its distribution consists of scat- tered occurrences in central and western North America (Windham 1993b), including South Dakota, Wyoming, Missouri, and Washington in the United States (Wind- ham 1993b; Rocky Mountain Herbarium 2008). Aside from Manitoba, its Canadian range includes the cor- dilleran regions of British Columbia and Alberta and a disjunct population in northern Saskatchewan (Rigby and Britton 1970; Windham 1993a). The global con- servation status of Gastony’s Cliffbrake is G2G3 — Im- perilled—Vulnerable (NatureServe 2013), although it may be locally abundant (Brunton 1979).
The species arose through hybridization of apoga- mous triploid Purple-stemmed Cliffbrake (Pellaea atro- purpurea [L.] Link) and diploid Western Dwarf Cliff- brake (Pellaea glabella ssp. occidentalis [E. E. Nelson] Windham), except for Missouri material, which has the diploid Pellaea glabella ssp. missouriensis (Gastony) Windham as a parent (Gastony 1988; Windham 1993a). Because the hybrid is apogamous and, thus, able to reproduce autonomously from the parent taxa, it was described as a new species by Windham (1993a). Of the parent taxa, only Western Dwarf Cliffbrake occurs in Manitoba, where scattered populations are known from the southern half of the province, and plants are often locally abundant where suitable habitat, i.e., calcareous outcrops, exists (Manitoba Conservation Data Centre, unpublished data). The sparsely villous rachis and pur- ple-brown petiole of Gastony’s Cliffbrake reliably dis- tinguish it in the field from Western Dwarf Cliffbrake which has a hairless rachis and a light-brown to straw- coloured petiole (Windham 1993a; Harms and Leighton 2011). It is distinguished from Purple-stemmed Cliff- brake by the presence of long, divergent (versus more abundant short, curly) hairs along the rachis, smaller
45
ultimate leaf segments, and large spores (Windham 1993a).
The first Manitoba collection of Gastony’s Cliff- brake (MANITOBA: South side of Marble Ridge Road, 1.6 km west of its junction with Highway 17, about 12 km north of Fisher Branch, 51.18361°N, 97.62556°W, dolomite cliff face, 26 July 2001, B. A. Ford 0140, M. Piercey-Normore & D. Punter, WIN 67479; D. F. Brunton herbarium) has not been previ- ously reported in the literature. It was initially identi- fied as Smooth Cliffbrake (Pellaea glabella Mettenius ex Kuhn), then annotated to Gastony’s Cliffbrake by D. F. Brunton in 2005. Western Dwarf Cliffbrake is also present at this location (Manitoba Conservation Data Centre, unpublished data). The specimen was collected at the north end of a limestone outcrop, locally known as Marble Ridge, which extends approximately 20 km to the southeast. The north end of Marble Ridge has been of ecological interest for some time because of its near-surface limestone bedrock and limestone cliffs, and it is now recognized as an alvar ecosystem (Neufeld et al. 2012). Until the surveys reported here, the popula- tion at this location was the only one known in Mani- toba.
Study Area and Methods
Near-surface limestone bedrock occurs in large areas of Manitoba’s Interlake region, and alvars are known to occur in several of these areas (Manitoba Conser- vation Data Centre, unpublished data). In 2012, the Manitoba Conservation Data Centre (MBCDC) part- nered with the Nature Conservancy of Canada to con- duct surveys in the southern portion of the Interlake to determine the extent and quality of alvar ecosystems and limestone cliffs in this region and record observa- tions of rare plant taxa, including Pel/aea Link species (Neufeld et al. 2012). Additional surveys of limestone-
46 THE CANADIAN FIELD-NATURALIST
dominated areas near Grand Rapids in the northern por- tion of the Interlake were conducted in 2012 and 2013 by MBCDC staff. Because Scoggan (1957) reported Western Dwarf Cliffbrake at Grand Rapids, the MBCDC surveys focused on limestone cliffs and ledges in an effort to refine the known distribution of this species in the area and search for Gastony’s Cliff brake.
For the Manitoba Alvar Initiative, 67 candidate sur- vey sites were identified using aerial imagery and GIS shapefiles of geological data available through the Man- itoba Land Initiative (http://mli2.gov.mb.ca/) and 61 of these sites were surveyed in 2012 (Neufeld et a/. 2012). For surveys at Grand Rapids, reports from the Mani- toba Geological Survey (Bezys and Kobylecki 2003;
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Kobylecki and Bogdan 2004) were also used to identify potential survey sites. Additional sites were identified in the field while conducting surveys. At Grand Rapids, seven sites were surveyed in 2012. In 2013, five new sites were surveyed and additional surveys were carried out at one 2012 site. The locations of Pellaea species were recorded with a Global Positioning System (GPS) unit and specimens were collected when necessary for confirmation of identification or to document new loca- tions. All specimens were deposited at the University of Manitoba Herbarium (WIN). Habitat type and con- dition were noted, and a coarse visual estimate of the number of plants was made. All known Manitoba oc- currences of Gastony’s Cliffbrake are shown in Fig- ure |.
FiGURE |. Pellaea gastonyi Windham) locations (black triangles) in Manitoba. Note: i i / : ¢ dAngies da. 6 MRENEC IS = a go Centre, and South, respectively; SGR = Sturgeon Gill Road. i ae
Results Marble Ridge North
This site corresponds to the location of the first Manitoba collection of Gastony’s Cliffbrake (see Intro- duction); thus, no additional specimens were collect- ed during the surveys reported here. In 2012, surveys
expanded the areal extent of the known population of
Gastony’s Cliffbrake at the north end of Marble Ridge from approximately 1.3 ha to 148 ha. The population size was estimated to be 300-500 plants. The actual ex- tent of this occurrence and number of plants may be somewhat larger as additional suitable habitat in the area remains to be surveyed. Most of the northern por- tion of Marble Ridge is alvar, with limestone bedrock at or within several centimetres of the surface. There are numerous limestone outcrops and plateaus up to 2 m high on the alvar, and it was at the edges of these out- crops, in the cracks in the limestone, that Gastony’s Cliffbrake was most often observed (Figure 2). The limestone at the edges of the outcrops tended to be more fractured and less consolidated than limestone else- where in the alvar. Gastony’s Cliffbrake was also ob- served growing at the base of limestone boulders sit-
FRIESEN AND MURRAY: GASTONY’S CLIFFBRAKE IN MANITOBA 47
ting on the alvar (Figure 3). It was only observed in areas of full or part sun. Western Dwarf Cliffbrake was also frequently observed in the area, although it was most often found growing in horizontal cracks in verti- cal faces of limestone cliffs and boulders, sometimes in very shaded areas. Marble Ridge Centre
In 2012, a new location of Gastony’s Cliffbrake was discovered in the central portion of Marble Ridge (MANITOBA: NW25-24-01W1, 51°06'30"N, 97°28'53"W, small cliff limestone ledge, 6 July 2012, Chris Friesen MBCDC9, WIN 76794), approximate- ly 12 km southeast of Marble Ridge North. The areal extent of occurrence was approximately 3.3 ha with an estimated population size of fewer than 100 plants. This area is also alvar and, as at Marble Ridge North, Gastony’s Cliffbrake was found growing in open areas at the edges of limestone outcrops in cracks in the rock. This portion of Marble Ridge contains additional un- surveyed habitat that likely supports Gastony’s Cliff- brake. Western Dwarf Cliffbrake was also found in the area, but only on shaded cliff faces.
o ~ x . oe. . > ; os imes ridge arble Ridge alvar s ar to those that support Gastony’s Cliffbrake (Pellaea gastonyi FIGURE 2. Limestone ridge on Marble Ridge alvat Soe oe Bis \ Windham). Photo courtesy of the Nature Conservancy of Canada, 2012.
48 THE CANADIAN FIELD-NATURALIST
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FIGURE 3. Limestone boulders on Marble Ridge alvar, some of which support Gastony’s Cliffbrake (Pellaea gastonyi Windham). Photo courtesy of the Manitoba Conservation Data Centre, 2012.
Marble Ridge South
Approximately 4.5 km southeast of Marble Ridge Centre, another population of Gastony’s Cliffbrake was discovered in 2012 (no specimen collected). This pop- ulation was divided between two limestone outcrops separated by a 200-m wide wooded gully. The popula- tion consisted of fewer than 100 plants in a very small area (approximately 0.04 ha) on the west side of the gully, with one additional plant found on the opposite side of the gully. The outcrops on either side of the gul- ly were open alvars. The plants on the west side were growing in habitat similar to those found at Marble Ridge North and Marble Ridge Centre. The one plant found on the east side of the gully was growing in deep shade on the edge of the outcrop. This was the only location where Gastony’s Cliffbrake was found grow- ing in shade.
Vegetative cover at the Marble Ridge sites was dom- inated by lichens and mosses growing on limestone bedrock and low-growing vascular plants including Creeping Juniper (Juniperus horizontalis Moench), Common Bearberry (Arctostaphylos uva-ursi [L.] Sprengel), and Poverty Oatgrass (Danthonia spicata [L.] P. Beauvois ex Roemer & Schultes).
Grand Rapids, Sturgeon Gill Road
Many of the 12 Grand Rapids sites surveyed had apparently suitable habitat for Gastony’s Cliffbrake, but it was found at just one (MANITOBA: 53°28'49"N, 99°13'15"W, upper edge of limestone cliff, 21 Septem- ber 2012, C. Friesen MBCDC72, WIN 76797). This site, which is just north of Sturgeon Gill Road, approxi- mately 32 km north of Grand Rapids, was first sur- veyed in 2012 with additional surveys in 2013. Over the 2 years, up to 100 plants were found growing at the top of a limestone cliff in cracks in the limestone bed- rock (Figure 4). Near-surface limestone bedrock was commonly observed near Grand Rapids, and such areas were usually dominated by Jack Pine (Pinus banksiana Lambert) and Paper Birch (Betula papyrifera Mar- shall). The Gastony’s Cliffbrake site was a Jack Pine— Paper Birch stand with sparse to discontinuous tree cover (Figure 5). The limited tree cover was likely be- cause of the lack of soil and the fire history of the site: the most recent burn was in 1979 (Manitoba 2013). Western Dwarf Cliffbrake was observed at this and a number of other sites near Grand Rapids, typically growing in the horizontal cracks of limestone cliffs.
2015 FRIESEN AND MuRRAY: GASTONY’S CLIFFBRAKE IN MANITOBA 49
FIGURE 4. Limestone ledge habitat of Gastony’s Cliffbrake (Pellaea gastonyi Windham) north of Grand Rapids, Manitoba. Photo courtesy of the Manitoba Conservation Data Centre, 2012.
Photo courtesy
Jack Pine—Paper Birch habitat on near-surface limestone bedrock near Grand Rapids, Manitoba. Jd
FIGURE 5. ae of the Manitoba Conservation Data Centre, 2012.
50 THE CANADIAN FIELD-NATURALIST
Discussion
Brunton (1979) noted that in Alberta there was a clear association between site orientation and the Pel- laea taxa present. Sites supporting Gastony’s Cliff- brake in Manitoba are level or have only slight direc- tional orientation and are in full to part sun; only one plant (at Marble Ridge South) was found growing in the shade of adjacent vegetation. Plants in Missouri are also found in full sun (G. Yatskievych, curator, Mis- sour! Botanical Garden, 10 January 2014, personal communication). Western Dwarf Cliffbrake was found in both open areas and those shaded by adjacent veg- etation (e.g., trees growing immediately adjacent to cliff face at Marble Ridge North).
Many sites surveyed included apparently suitable habitat (exposed fractured limestone), yet supported few, if any, Gastony’s Cliffbrake. The paucity of Gas- tony’s Cliffbrake at such sites may indicate that micro- habitat or microclimatic characteristics are not suitable (Richard ef al. 2000). Alternatively, factors related to dispersal and establishment could be limiting the small- to medium-scale distribution (Wild and Gagnon 2005).
At a broader scale, Gastony’s Cliffbrake occurs in three ecozones in Canada: montane—cordillera (British Columbia and Alberta sites), boreal shield (northern Saskatchewan site), and boreal plains (Manitoba sites) (Ecological Stratification Working Group 1995). Its North American distribution ranges from the Great Plains (Missouri, South Dakota) in the south to the boreal forest near Lake Athabasca in northern Sas- katchewan. These data suggest that, as a species, it is tolerant of considerable climatic variation at a broad scale.
The typical habitat for this species throughout its range 1s calcareous rock, often limestone or dolomite (Brunton 1979; Windham 1993a), and the discontin- uous distribution of this substrate in North America likely accounts for the similarly discontinuous distri- bution of Gastony’s Cliffbrake. Hastings (2002) linked the distribution of Grimmia Dry Rock Moss (Grimmia teretinervis Limpricht), a moss of calcareous rock that occurs at Marble Ridge and near Grand Rapids (Caners 2011; Manitoba Conservation Data Centre, unpublished data), to the margins of ancient epicontinental seaways, with occurrences in central and western North America associated with the margin of the Cretaceous epicon- tinental seaway. The known Canadian