Check List 9(4): 883–893, 2013 © 2013 Check List and Authors Chec List ISSN 1809-127X (available at www.checklist.org.br) Journal of lists and distribution

n Distribution and natural history of carnivorous of istributio D Saskatchewan, Canada * raphic

g Fraser Baalim, Catherine Peters, and J. Hugo Cota-Sánchez

eo * Corresponding author. E-mail: [email protected] University of Saskatchewan, Department of Biology and W.P. Fraser Herbarium (SASK), 112 Science Place, Saskatoon, SK S7N 5E2, Canada. G n o Abstract: otes

N We provide distribution maps of carnivorous plants as well as an assessment of rarity status and potential ecoregionthreats to in diversity the northeast. of carnivorous Several areas taxa exhibiting in Saskatchewan high carnivorous using a biodiversity diversity informatics including rare approach. and endangered Saskatchewan species is home to ten carnivorous species ranging from the Mixed Grassland ecoregion in the southwest to the Selwyn Lake Upland as deserving conservation priority. are: the Nesbitt Provincial Forest, the Prince Albert National Park, and the Athabasca Sand Dunes. We propose these areas

Carnivorous plants, those that draw a portion of their their unusual ecological requirements. nutrients from animal prey rather than the soil, have long intrigued and confounded biologists. In 1796, American to different lineages of carnivorous plants: naturalist John Ellis described the insect trapping habit (),Saskatchewan Sarracenia is home L. (Sarraceniaceae), to four genera , belonging of Dionaea muscipula J. Ellis (Venus Fly Trap) to Carl and Linnaeus, who responded with disbelief, arguing that 2003). These three families are placed in three distinct a plant preying upon animals was “against the order of phylogenetic orders:() , (Harms Ericales, 1978, and nature as willed by God” (Barthlott et al. 2007). Nearly (Albert et al. 1992). The general distribution of carnivorous plants in Canada and is ofa centurycharacteristically later, Charles elegant Darwin experiments, offered theDarwin first showed formal e.g., Porsild and Cody (1980), Mellichamp thatscientific D. rotundifolia treatment along of carnivorous with other plants.species With of Drosera a series L. known based on distributional data in provincial and/or (Sundew), Utricularia L. (Bladderwort), and Pinguicula L. regional floras, (Butterwort) were indeed capable of capturing, dissolving, (2009). In Saskatchewan, the broad geographic range and absorbing insect prey (Darwin 1875). Despite the of these species has been presented (Harms 1978), but novelty implied by their unusual habit, carnivorous plants investigationlittle is known is regardinga case study the based extent on of digital their information distribution do not constitute a single branch of the evolutionary tree. fromand the herbarium specific ecological specimens areas stored these in plants the occupy. database This of Carnivory is a convergent lifestyle and has independently involving aspects of biodiversity informatics in the et al. 2007). the W.P. Fraser Herbarium (SASK) and is pioneer in evolvedCarnivorous at least plantsfive times are foundin the onplant every kingdom continent (Barthlott except the distribution patterns of carnivorous plants in Antarctica (Albert et al. 1992; Barthlott et al. 2007) and province. It provides the first approach in understanding exhibit a degree of morpho-physiological specialization levels stands for various reasons. Foremost, it will enhance theSaskatchewan. utility of herbarium Its relevance voucher at thespecimens regional and and contribute national the majority of carnivore plant diversity is found in the to the understanding of endangered and rare species by tropics,unrivalled various by most species species of inUtricularia the plant, kingdom.Pinguicula While and georeferencing and mapping updated occurrences of Drosera possess adaptations that allow their ranges threatened taxa. Also, it will facilitate targeting areas with to extend into higher latitudes of the Canadian arctic, high demographic indices and biodiversity of carnivorous Scandinavia, and Greenland (Barthlott et al. 2007). The areas deserving conservation priority. Maps integrating specimenplants in locality Saskatchewan, can help identify which areas can berequiring proposed further as withMalaysian, 133, 86, Australian and 65 species, and Brazilian respectively floristic (Barthlott regions et are al. study, sampling, protection and monitoring and ultimately the most significant centers of carnivore diversity globally, assist in predicting the future of ecosystems and the shifts the , especially the swamps and wetlands of common and rare species in view of human activities of2007). Florida, Within is North home America, to a high the diversity southeastern of carnivorous region of and climate change. taxa, including various endemics (Barthlott et al. 2007). Canada has 19 species of carnivorous plants (10 of which We used digitized label information from voucher These plants are widespread throughout Canada, mostly herbariaspecimens with at the the W.P. following Fraser objectives: Herbarium 1) (SASK) to generate of the occur in Saskatchewan) (Harms 1978; Canadensys 2012). aUniversity general provincialof Saskatchewan distribution and several map for other carnivorous Canadian occurring in small, well defined habitat patches due to 883 Baalim et al. | Distribution of carnivorous plants of Saskatchewan, Canada genera as well as create a searchable database of these paper) and species (not included in this paper but can be plants, 2) to re-assess the species’ rarity status on a provincial and national scale, and 3) to identify areas of data includes the geographic coordinates provided by diversity in carnivorous plants. The limited understanding produced at the above link) in Saskatchewan. The digitized

estimatedthe original based collector on the in given the herbarium locality information labels. When using no of distributional patterns of carnivorous plants, the lack thecoordinates Natural were Resources specified, Canada the latitude on-line and Atlas longitude of Canada were – of a complete Saskatchewan flora, and the accelerated toporama (Natural Resources of Canada (2013) reference institutionloss of natural devoted habitats to the justifies documentation this biodiversity-based and study maps. Our mapping protocol follows that of Choi and Cota- research. Furthermore, SASK herbarium is the primary Allium L. of the Canadian collection of 180,000+ specimens, covering over 125 years Prairie Provinces. First, locality data for each species of botanicalSaskatchewan’s exploration flora withand is the home most to comprehensive a systematic wereSánchez imported (2010), into implemented a customized in map development tool chronological, historical, taxonomic, and geographic based on an open-source GoogleTM Maps API on-line development tool. Once generated, the GoogleTM map was 2009), thereby offering an ideal data mining opportunity imported into Adobe Photoshop as a JPEG image, and the coverage of provincial plants (Cota-Sánchez and Harms boundaries were eliminated, leaving only the province of basedfor this collectors, study. Additionally, spanning theSASK time Herbarium period of contains ca. 1910 the to overall distribution of all records of carnivorous plants main collections of almost all significant Saskatchewan- inSaskatchewan. the collections We obtained created fromprovincial all the maps above depicting herbaria the in three different layers, namely general topographic and the present. Coverage of species at risk is equally strong at hydrological information, ecoregions, and soil types. In SASK Herbarium. No other research institution, with the addition, we generated distributional provincial maps possible exception of the Saskatchewan Conservation Data for each carnivorous with the ecoregions layer. Centre (SK-CDC), has such comprehensive information on This study also includes a literature review to provide geography, , and rarity status of Saskatchewan general information dealing with taxonomic diversity, whileflora. Finally, maximizing SASK theis part use of of the systematic Canadensys collections group, a morphological adaptations and specializations as well multidisciplinary network aiming to preserve biodiversity line with the ongoing biodiversity informatics program species. The rarity status of the species was determined andacross digitization Canada. Within of systematic this scope, collections the present to study mobilize is in basedas ecological on previously requirements published of eachdata such of the as Saskatchewan the Canadian the wealth of information stored in labels of herbarium Endangered Species Conservation Council (CESCC 2011), Maher et al. tools relevant to conservation management, modelling systems,specimens andand endow environmental policy makers programs and scientists dealing with (1979), Harms (2003), the SK-CDC, and the plantsNature Conservancy lies in three Ranking plant of rare families: species (Table Droseraceae, 1). biodiversity in general. The taxonomic range of Saskatchewan carnivorous the flora of Saskatchewan, in particular, and Canadian 2003). These three phylogenetically distant carnivorous representing the range of taxonomic diversity and familiesLentibulariaceae, exhibit a and perplexing Sarraceniaceae array of (Harms morphological 1978, geographicWe surveyed distribution herbarium of carnivorous voucher plants specimens in the traits that characterize their traps. The trap design in from 759 voucher specimens available from herbaria of the traps, pitfall traps, and suction traps (Table 1). A total of followingprovince. institutions:We compiled 13 and from digitized the University label information of Alberta fourSaskatchewan genera and carnivorous ten species species of carnivorous is of three plantstypes: sticky occur (ALTA), 25 from the Canadian Museum of Nature (CAN), 91 from the Agriculture and Agri-Food Canada (DAO), and common in the province, these taxa are restricted to boggyin Saskatchewan areas, which (Table are characterized1), and though by they acidic, are relativelynutrient- poor soils, , and standing water (Schnell varied630 from depending the W.P. Fraser on rarity, Herbarium distributional of the University range, and of representationSaskatchewan (SASK). within Thethe numberbotanical of collections vouchers per surveyed. species a latitudinal band (50°N to 70°N) which contains most The specimens labels contain information useful to 2002). Globally, the province of Saskatchewan falls within understand the taxonomic diversity, phenology, geographic boggy habitats, the preferred environments of carnivorous distribution, ecological range, and other attributes of species,of the world’s are widespread peatlands throughout (Moore 2002). the province. Hence, islands of of provincial carnivorous plants in the Specify database Droseraceae: , D. linearis (Figure 1A-C), carnivorous taxa. With these data, we created a catalogue andSaskatchewan D. rotundifolia houses three species fromDrosera the database program which facilitates the digitization of museumof SASK Herbarium.and herbarium Specify collection software data is (Specify an open Software source the brightly coloured(Harms mucilage 1978, secreted 2003). The by genusglands on the 2012) and provides a data-computing interoperable tipstakes of its tentacles common covering name, “Sundew”, the adaxial from surface the appearance of the of platform that is responsive to evolving community needs and to new research opportunities enabled by the general, Drosera Internet. An electronic database (Baalim et al. published (Figure 1B), forming the characteristic sticky trap. In online) was developed for the specimens of carnivorous species are small perennial herbs, weakly plants investigated and maps created depicting the rooted, with a rosette of modified trap leaves borne on a distribution of each carnivorous genus (included in this short, unbranched shoot (Harms 1978). The small, perfect, actinomorphic (Figure 1C) are ephemeral, self- pollinated, and presented above the sticky trap leaves on884 a Baalim et al. | Distribution of carnivorous plants of Saskatchewan, Canada determinate raceme (Judd et al. 2008; Schnell 2002). Late Carnivorous plants have long been considered precious in the fall, Drosera form hardy winter buds (hibernacula), specimens by collectors, and it appears that the rarer the which allow the plant to survive through the winter and plant, the more desirable the specimen. As a result of over- reproduce both asexually and sexually during spring and exploitation and habitat loss, several carnivorous plants summer (Schnell 2002). are listed as rare species. The family Sarraceniaceae is represented by a single Sarracenia purpurea Following, In we Saskatchewan summarize ca.the 35% information of native 1978, 2003), a perennial, rhizomatous herb with leaves availablespecies are regarding listed inthe five current categories rarity status as provincially of carnivorous rare species in Saskatchewan, (Harms (Harms 2003). 1D-F). Insects are drawn to the lip (peristome) of a pitcher Assessing the general status of Canadian and world traphighly by modified nectar secreting to form the glands pitcher and or red pitfall pigment, traps while(Figure a speciesplants in is Saskatchewan. a challenging, but essential process. To date, 25 slippery, waxy cuticle and downward pointing hairs on species have been extirpated from Canada the inside of the pitcher walls (Figure 1E) prevent the (CESCC 2011). Even though most Canada’s carnivorous prey from escaping the pitfall trap (Barthlott et al. 2007). A long scape (up to 35 cm) bears a single, actinomorphic, ( Butterwort, Pinguicula macroceras Pall. ex plants are ranked in the Secure category, twoUtricularia species shaped style nearly enclosing the stamens (Figure 1F) ochroleuca (Judddownward-pointing et al. with a five-lobed, umbrella- andLink, one and species Yellowish-white (Thread-leaved Bladderwort,Sundew, visits to large, strong insects (usually queen bees) while R.W. Hartm.) have a CanadaCESCC rank 2011) of Sensitive,. Several 2008). The unique flower shape limits pollinator schemes have been proposed to assign rarity status to odds of successful cross pollination (Schnell 2002). This Raf.), has the At Risk rank in Canada ( 2010 speciesimposing reproduces specific entry sexually and by exit seed points, and asexually maximizing through the report (as per the CESCC 2011) recognizes the following fragmentation of the rhizomes (Barthlott et al. 2007). Canada’s vascular plant species. The Wild Species the Lentibulariaceae, includes the genera Pinguicula and Exotic,general and categories: Accidental Extinct, (the reader Extirpated, is referred At to Risk, CESSC May 2011 Be UtriculariaThe third (“Bladderwort”) carnivorous . SaskatchewanPinguicula, commonly plant family,called forAt Risk, detailed Sensitive, description Secure, of Undetermined,each type Not Assessed, the categories applied for the species in includes two species in the province, P. vulgaris (Figure ). Here wePinguicula focus on 1G-I)“Butterwort” and P. villosa due to their fleshy, greasy looking leaves, villosa, P. vulgaris, Utricularia cornuta, and U. minor are perennial herbs with compressed rosette leaves measuring province of Saskatchewan. At the national level, (Harms 1978, 2003). These are small, 1978; Barthlott et al. 2007). The small, zygomorphic, assessments),included in the followed May Be Atby RiskD. anglica category and (may D. linearis be at placedrisk of 1-4 cm long and a highlyP. villosa reduced and up to six system in P. vulgaris (Harms) extirpationin the Sensitive and arecategory candidates (Species for thatadditional are not detailed believed risk to bloom on an erect scape (Figure 1I). Nectar is produced in purpleelongated, flowers rearward-pointing (one in spurs (Barthlott et al. 2007). require special attention or protection to prevent them As with Drosera, the trap leaves of temperate Pinguicula be at risk of immediate extirpation or extinction but may species species considered Secure (CESCC 2011) (Table 1). In hibernacula, which have the potential to re-emerge the from becoming at risk), with the remaining provincial next growingdie back season in the to winter, restore leaving the population bulb-like, mainlyperennial by plant species into seven provincial categories: Common asexual reproduction (Schnell 2002). (C),turn, Fairly Harms’ Common (2003) (FC), framework Uncommon divides (UC), Saskatchewan’sVulnerable (V), The second carnivorous genus of the Lentibulariaceae, Threatened (T), Endangered (END) and Extirpated (EXT). Utricularia, is cosmopolitan and includes four terrestrial, Utricularia cornuta has zygomorphic, bilabiate 1),According i.e. to this ranking scheme, one carnivorous species, sub-aquatic and aquatic herbs species in Saskatchewan it occurs in only six, isto threatened 15 localities. in SaskatchewanThree other species, (Table (Harmsprovince, 1978, namely 2003). U. cornuta, The genus U. intermedia, U. macrorhiza Drosera, it is linearis, likely to Pinguicula become endangered villosa and in P.the vulgaris province, are as (Figureflowers 1J-L), (Figure and 1L)U. minor and includes four species in the listed in the vulnerable category (Table 1), warranting special concern because of their low or declining numbers (Harms 1978, 2003). The plants with plants locally sparse in 16-25 sites in the province form stolon-like structures bearing filiform photosynthetic sisterleaves taxon, and modified Pinguicula trap, the leaves sophisticated (Figure 1J, suction K). Compared traps of 1978, 2003). Utriculariato the relatively are very simple specialized adhesive in form or sticky and function. traps of The its but free from obvious, immediate endangerment (Harms small, hollow bladders (0.2-1.2 cm), each with a single, evaluates the rarity of species at the global and subnational The Nature Conservancy Element Rankings (NCER) inward-openingDespite the door, widespread are borne geographicon stalks underwater distribution or (provincial) scale (reviewed in Harms 2003). According to andbelow the the relative soil surface abundance (Figure of1K). numerous species, over- plants,the Saskatchewan Conservation and Pinguicula Data Centre villosa (2011),, are which given collection and other human pressures, such as agricultural uses the NCER system, two of Saskatchewan’s carnivorous expansion, continue to threaten plant and animal life. not rare, typically less than 100 occurrences and less Changes in landscape, including habitat fragmentation and thana global 10,000 rank individuals), of G4, (apparently while the secure, others uncommonare considered but the removal of entire plants, are major factors that deplete “Demonstrably Secure-Common” (G5) (Table 1). On the wild populations and lead to the loss of genetic diversity. provincial scale, D. linearis

is given a rating of S1, defined885 Baalim et al. | Distribution of carnivorous plants of Saskatchewan, Canada

Utricularia and founder of the Department of Biology, which he led cornutaas “Critically Imperilled”, having five or fewer occurrences, was the University of Saskatchewan’s third president “Imperilled”or less than 1,000 and “very individuals vulnerable in Saskatchewan. to extinction”, typically specimen. Six specimens are from the 1920s. Five of these with six tohas 20 aoccurrences provincial or status1,000 to of 3,000 S2, definedindividuals. as recordsfrom 1913-1949, are for Utricularia which adds macrorhiza further , significanceof which two to were this Pinguicula villosa, P. vulgaris, and Utricularia minor are each given the provincial rating S2S3 (Table 1), indicating that their status lies somewhere between “Imperilled” and collected by William P. Fraser in 1922 (SASK 86773 and “Vulnerable” (21 to 100 occurrences, or between 3,000 to 86774) around the city of Saskatoon (Appendix 1). These 10,000 individuals). historical specimens have added value because W.P. Fraser In the cases of D. linearis and U. cornuta ofwas this responsible facility after for him. the The creation remaining of SASK three Herbarium specimens inof and the Nature Conservancy’s systems disagree in their U.1925 macrorhiza (Cota-Sánchez date from and Harms1923, 1924,2009), and hence 1927 the collected naming , Harms’ (2003) D. linearis a higher conservation priority than U. cornuta relative rankings. The Nature Conservancy (NC) gives by W.C. McCalla, ALTA 65412 (Buttress area), R.C. Russell, lists U. cornuta as requiring higher conservation priority fromSASK the 86772 1920s (Duncairn is for Drosera area), rotundifolia and W.H. collected Cameron by SASK R.C. (G4than S1D. and linearis G5 S2(threatened respectively), vs. vulnerable) while Harms (Table (2003) 1). 21178 (Saskatoon area) (Appendix 1). The sixth record (Appendix 1). The remaining specimens in the list of devoted to the preservation of ecological diversity, the historicalRussell, SASK accessions 91998, are in 1926from thein the 1930s area and of 1940s,Speddington when Although both organizations, the NC and the SK-CDC, are These pre-1950 collections are mostly concentrated in conflicting designation between these authorities and further botanical exploration took place in Saskatchewan. those of Harms (2003), which were based on personal and (Appendix 1). re-evaluatefield observations the categories (V. Harms, assigned. pers. comm., University of the Thevicinities mapping of the protocolscities of Saskatoon used here and Prince were Albert useful Saskatchewan),Our carnivorous highlights plants the databaseneed for further contains inquiry plants to in elucidating distribution patterns of the species collected since the late 1800s, but we established 1950 investigated. Carnivorous plants are distributed as a cutoff to consider vouchers in the historical category. borders (Figure 2A-C). The southernmost occurrences are representedthroughout Saskatchewan by Utricularia from found the in southernthe Mixed to Grassland northern AppendixWe gathered 1). A 83 common historical issue records among dating historical prior specimens to 1950 ecoregion in the southwest corner of the province, at infor our the list carnivorous is the amount plant and group accuracy in Saskatchewan of the information (see the northernmost occurrences are represented by have complete and detailed label information, earlier Drosera49.1°N and and 108.1°W Utricularia (Figures. found in2A-C the and northeast 3D). Conversely, corner of provided by the collector. While recent voucher specimens number. In spite of this limited label information, data Drosera, fromcollections the past often were lackuseful accurate to locate the locality oldest and collection(s), collector thePinguicula, province, and within Utricularia the Selwyn Lake Upland ecoregion main collectors and localities of carnivorous species. Our at ca. 59.9°N and 102.1°W (Figure 3A, D). Also, database survey indicates that the oldest collection of occur north of Lake Athabasca in the Tazin Lake Upland ecoregion at ca. 59.5°N and This specimen of Sarracenia purpurea occurrence108°W (Figure of carnivorous3A, C, D). Ongoing plants digitization goes well of beyond Canadian the Saskatchewan carnivorous plants dates back to 1911. northernvoucher specimens provincial atboundaries SASK Herbarium to areas indicatesbeyond the that Arctic the (SASK 12946) was Circle in the Northwest Territories, approximately 69.4°N collected by Walter P. Thompson in the area of Prince TableAlbert 1. (Appendix 1). It is worth noting that W.P. Thompson merges subsp. Families gibbosa and species under subsp. of carnivorous purpurea. plants reported in Saskatchewan, their rarity status as assigned by CESCC (2011), Harms (2003), and the Nature Conservancy (as per the Saskatchewan Conservation Data Centre), and their trap types. *Mellichamp (2009) in Flora of North America now Canadian Carnivorous Plants Saskatchewan Rarity status Saskatchewan Rarity status Family & Species Trap type (CESCC 2011) (Harms 2003) (Nature Conservancy) Droseraceae Drosera anglica Sensitive Uncommon Not threatened D. linearis Goldie Sensitive Vulnerable G4, S1 Huds. Sticky D. rotundifolia L. var. rotundifolia Secure Common Not threatened Sticky Lentibulariaceae Sticky Pinguicula villosa L. Vulnerable G4, S2S3 P. vulgaris L. Vulnerable G5, S2 May Be At Risk Sticky Utricularia cornuta Michx. Threatened G5, S2 Suction May Be At Risk Sticky U. intermedia Secure Common Not threatened Suction May Be At Risk U. macrorhiza Le Conte Secure Common Not threatened Suction Hayne U. minor L. Uncommon Not threatened Suction Sarraceniaceae May Be At Risk Sarracenia purpurea L. subsp. Secure Not threatened Pitfall gibbosa (Raf.) Uncommon/Fairly common Wherry* 886 Baalim et al. | Distribution of carnivorous plants of Saskatchewan, Canada

Figure 1. Drosera linearis

0.5 cm). D-F: Representative Sarracenia purpurea native . speciesD. Plant ofin natural carnivorous habitat plants (scale in = Saskatchewan.5.0 cm). E. Close A-C: up of the pitcher, trap a vulnerable showing the species operculum in Saskatchewan and downward- with nationallypointing hairs rare inside status. the A. Plantpitcher in naturalwall (scale habitat = 1.0 (scale cm). F.= 1.0Flower cm). detail B. Close showing up of the the leaf umbrella-shape sticky trap with style insects (St) and (scale = 1.0 (p) cm). as indicatedC. Flower bydetail the (scale arrows = (scale = 1.0 cm). G-I: Pinguicula vulgaris Utricularia macrorhiza. J. Plant removed , a vulnerable species in Saskatchewan with nationally rare status. G. Plant in natural habitat (scale = 1.0 cm). H. cm).Close up of the rosette leaves (sticky traps) with insects (scale = 1.0 cm). I. Flower detail (scale = 1.0 cm). J-L. from its aquatic habitat (scale = 1.0 cm). K. Close up of stem showing linear leaves and suction traps (arrow) (scale = 1.0). L. Flower detail (scale = 1.0

887 Baalim et al. | Distribution of carnivorous plants of Saskatchewan, Canada

P. villosa exist. Porsild and Cody (1980) mapped Pinquicula vulgaris, P. villosa, and P. villosa andmost 133.2°W, Utricularia where species records northward of and northwestward to 134706;has been 20/06/1992,and continues SASK to be 136189; found predominantly 08/04/1993, along SASK the Arctic Ocean. The wide geographic coverage of these 163973; 01/07/2004, SASK 170442). Similarly, species within the province and beyond is evidence of the province (e.g. degree of specialization and adaptability of carnivorous the Wollaston Lake road in the northeast portion of the plants to a wide diversity of climatic conditions. , 26/07/1973, SASK 52755; 07/09/1973, It is noteworthy that the distributions of Drosera, SASK 52756; 13/07/1998, SASK 147567; 07/08/1998, Sarracenia and Pinguicula SASK 147687; 27/06/2006, SASK 171128; 17/07/2008, common pattern. Most occurrences were recorded in SASK 179297), though a historical record exists from the the Boreal Transition ecoregionin Saskatchewan (Figure 2B), which share is a north shore of Lake Athabasca (59.6°N, 109.2°W, by H.M. dominated by Populus tremuloides Michx., Picea glauca Raup, 24/07/1935, CAN 99311 (Appendix 1) and a single (Moench) Voss, and Pinus banksiana Lamb. (Fung 1999), specimenU. cornuta has been, currently collected in in thethe Wildcat threatened Hill Provincial category, among other species. These three genera are totally Park (53.3°N, 102.5°W [07/10/1992, SASK 107572]). excluded from the Mixed and Moist Mixed Grasslands, and only two records each of Drosera and Pinguicula has been reported only along the Wollaston Lake road (21/07/1973, SASK 53180; 28/07/1973, SASK 52748), characterized by P. tremuloides, Symphoricarpos Duhamel, where it continues to be collected (19/07/1983, SASK andhave beenRosa collectedL. (Fung in 1999),the Aspen though Parkland, these an collectionsecoregion 94565; 09/07/1998, SASK 147682; 12/07/2005, SASK were made in boggy or calcareous microhabitats. 170918), on the south shore of Lake Athabasca (59.0°N, Interestingly, the two Drosera records from the Aspen 70090),109.0°W and [07/08/1975, on the site ALTA of the 83184; since 11/08/1975, decommissioned SASK 145991; 07/07/1979, SASK 70096; 11/08/1979, SASK U. cornuta have been occurrencesParkland correspond suggest that with Drosera small’s patches of gray soil Cluff Lake uranium mine (58.4°N, 109.6°W [26/08/1978, bywithin soil achemistry large band than of byblack temperature soil (Figure and 2C). precipitation These two SASKThe 174243]). wide distribution No occurrences of carnivorous of plants depicted in as its distribution is restricted to niche areas is of better grey defined soil or ourreported maps at demonstrates Lake Athabasca the or ample Cluff degreeLake since of adaptability 1978-79. Canadian Shield (Figure 2C). Utricularia, in turn, is the of these plants to altitudinal gradients while remaining e.g., soils with streamsgenus with and broadest shares distributiona sympatric in distribution Saskatchewan. with It its is restrictedrestrictions to ofspecific the carnivorous ecological conditions, habit have resulted in a congenerica truly aquatic terrestrial plant foundcarnivorous in lakes, species ponds, in theand northern sluggish low pH and low nitrogen content. The ecological and habitat two thirds of the province. Species of Utricularia are also and physiology among disparate angiosperm lineages. The found in aquatic environments throughout the Aspen sharedremarkable preference set of homoplasious for open, moist, characters low nutrient in morphology habitats means that unrelated carnivores often co-exist in , only carnivorous species with occurrences in the Cypress and similar ecosystems (Barthlott et al. 2007). In fact, UplandParkland, region Mixed, (Figure and Moist 3D). Mixed Grasslands and are the more than 13 carnivorous plant species exist in a single of vulnerable and threatened species with more recent diversity converges in the fen and boggy areas of three collectionsOur work and has records expanded from previous Canadian geographic herbaria. ranges For provincialbog (Folkerts areas, 1982). namely Our study the Athabasca indicates Sandthat carnivorous Dunes, the Nesbitt Provincial Forest, and Prince Albert National the vulnerable species D. linearis in the province: two near instance, Harms (1978) reported only five occurrences of as candidates for conservation priority not only for the abundancePark region. of Ascarnivorous indicated species next, we but suggest also because these these areas Prince Albert (53.6° N, 106.2°W, [12/07/1933, SASK 12947 areas host a number of endemic and narrowly distributed and 05/08/1933, SASK 12952]), two near McKague (52.6° N, 103.9° W [10/08/1933, SASK 12950 and SASK 12951]), (Maher et al. (1999)and one indicating near Garthland its occurrence (53.0° N, in 106.3° Greenbush W [07/07/1971, on the Red species that are provincially rare and/or threatened SASK 54271], with updated D.information linearis was by again Schwab-Moe reported northern prairie 1979; province Harms 2003). regions Further, and Athabascaearly vegetation Sand Dunes.studies emphasized the unique floristic elements of the 179631);Deer River however, east of McKague. no further collections have been made carnivorous species (including U. cornuta in the threatened in the Garthland area in 2008 (12/07/2008, SASK category)Our emphasize findings indicatingthe uniqueness the presence of the Athabasca of seven of D. linearis in these regions, subsequent collections have Sand Dunes regions in terms of the diversity, endemism revealednear Prince that Albert this orspecies’ McKague. range Despite extends the north recent of absence Prince Albert into the boreal forest (e.g. genera, except Sarracenia and rarity of plant species. All Saskatchewan carnivorous , 08/09/1984, SASK79389 , range well north of SK, most to and 21/08/1996, SASK 139967), including the Athabasca carnivorousthe Arctic Ocean plant (V. biodiversity Harms, pers. deserving comm.). special attention Sand Dunes (59.0° N, 109.0° W [08/05/1975,P. vulgaris SASK, listed 68579 in in termsOur mapsof conservation. have identified Among severalthese areas, areas the ofNesbitt high andthe 07/08/1979, vulnerable category,SASK 145934]). occurs in sporadic patches According to Harms (1978), carnivorous species, two of which (D. linearis and P. vulgarisProvincial) are Forest in the (52.9°Nvulnerable and category 106.0°W) and are contains restricted six distributionthroughout the pattern Boreal, persists Aspen ( Parkland,e.g. and Athabasca Plain ecoregions. Recent collections confirm that this , 16/06/1983, SASK to calcareous fens. The Prince Albert National Park region888 Baalim et al. | Distribution of carnivorous plants of Saskatchewan, Canada

are rare compared to their sister species, the paucity of of carnivorous plants, featuring records of eight species, records may be exaggerated by the fact that, when not in including(53.9°N and two 106.2°W) vulnerable is anothertaxa: D. linearis area of and high P. incidence vulgaris. other, more common Utricularia to seven carnivorous species including the vulnerable D. bloom, these plants are easily overlooked or mistaken for Thelinearis Athabasca and the Sand threatened Dunes (59.1°N species and U. 109.1°W) cornuta are. Slightly home species (Harms 1978). southwest of the Athabasca Sand dunes is another Since much of Saskatchewan’s landscape is uninhabited thatand difficultthe diversity to access, found botanical in local “hotspots” collections indicated are lacking above for abundance of carnivorous plants, including records of six andlarge in portions our maps of the is alsoprovince. representative However, ofwe the hypothesize diversity species,distinct areaamong (58.4°N them theand vulnerable 109.7°W) P.with vulgaris a relatively. Because high of present in other provincial regions with similar ecological the relatively rich levels of taxonomic diversity, including conditions, even though the vast majority of the landscape the presence of rare plants, we propose these areas as remains botanically unexplored. priority sites for conservation. Notwithstanding the potential biases indicated The distributional patterns of carnivorous plants presented in this paper provide useful clues and potential exhibit intrinsic and interdependent relationships with theabove, environment carnivorous and plants other have organisms. specific Due habitats to complex and interactions of these plants with other community soleguides use to of establish herbarium conservation specimens priorities. in the generation However, we of members, their conservation may have unintended maps.would Foremost, like to note the thegeographic intrinsic range biases of imposed collection by sites the positive effects on ecosystems. Along with pollinator and (and hence distribution) is non-random; typically more prey interactions, carnivorous plants may support endemic specimens are collected close to major population centres, communities of commensal within their traps especially cities with universities or herbaria (Crawford et al. 1983). The traps of some pitcher plants provide refuge for amphibians (Jennings and Rohr 2011), and(Istock a species of Drosera has been shown to compete with and Hoagland 2009), main roads, and accessible areas. For spiders for prey (Jennings et al. 2010). Carnivorous plants instance, the area of high incidence we identified at 58.4°N are particularly sensitive to changes in environmental sampledand 109.7°W heavily corresponds compared to withthe surrounding the decommissioned areas due conditions, and many species are conspicuous and easy toCluff environmental Lake uranium impact mine, assessments the site ofof mining which hasactivities. been Similarly, the band of high Utricularia density extending candidates as “indicator species” for assessing the health andto identify. integrity These of thefeatures ecosystems make carnivorous they inhabit plants (Jennings good spurious pattern caused by heavy collection efforts along and Rohr 2011), which, in conjunction with the role of northeast from La Ronge (55.1°N,U. cornuta 105.3°W) and is U. likely minor a Sphagnum the Wollaston Lake road. While as a keystone species in restoration (Rochefort

Figure 2. Drosera, Sarracenia, Pinguicula, and Utricularia Distribution maps of carnivorous plants in Saskatchewan. A-C: Combined distribution map of the four carnivorous genera ( ) in Saskatchewan. A. Map with topographic layer. B. Map with ecoregions layer. AP=Aspen Parkland, AB=Athabasca Plain, BT=Boreal Transition, CH=Churchill River Upland, CY=Cypress Upland, MBL=Mid-Boreal Lowland, MBU=Mid-Boreal Upland, MG=Mixed Grassland, MMG=Moist Mixed Grassland, SU=Selwyn Lake Upland, TU=Tazin Lake Upland. C. Map with soil types layer. BR=Brown, DB=Dark Brown, BL=Black, DG=Dark Grey, GR=Grey, CS=Canadian Shield. 889 Baalim et al. | Distribution of carnivorous plants of Saskatchewan, Canada

Figure 3. Distribution maps of each carnivorous genus. A. Distribution map of Drosera species. B. Distribution map of Sarracenia purpurea. C. Distribution map of Pinguicula species. D. Distribution map of Utricularia

species. AP=Aspen Parkland, AB=Athabasca Plain, BT=Boreal Transition, CH=Churchill River Upland, CY=Cypress Upland, MBL=Mid-Boreal Lowland, MBU=Mid-Boreal Upland, MG=Mixed Grassland, MMG=Moist Mixed Grassland, SU=Selwyn Lake Upland, TU=Tazin Lake Upland. 890 Baalim et al. | Distribution of carnivorous plants of Saskatchewan, Canada

2000), supports the importance of protecting these areas. Literature Cited In fact, there is evidence that boreal peatlands have a higher phylogeny and structural evolution. Science 257(5076): 1491-1495. proportion of characteristic species, both plant and animal, Albert, V.A., S.E. Williams and M.W. Chase. 1992. CarnivorousCarnivorous plants: plant distribution in Saskatchewan. Electronic Database accessible at he Canadian boreal forests are under Baalim, F., C. Peters and J.H. Cota-Sánchez, (Published online). html. Captured on April 2012. intensethan adjacent pressure, upland but despite habitats, the such dramatic as Aspen changes Parkland to the http://www.herbarium.usask.ca/MapDevelopment/mapsCarnSK.The curious (Locky 2010). T world of carnivorous plants. Portland, OR: Timber Press. 224 p. of this area to a wide diversity of wildlife, limited efforts Canadensys.Barthlott, W., 2012.S. Porembski, Database R. of Seine vascular and I. plants Theisen. of Canada. 2007. Electronic Database accessible at . orSaskatchewan programs haveboreal been transition prepared zone to and stop the deforestation importance Captured on 13 May 2012. et al. 2002). Canadian Endangered Specieshttp://data.canadensys.net/vascan/search/ Conservation Council (CESCC). 2011. Wild Species 2010: The General Status of Species in Canada. National Electronic Database and pdf informationand the vanishing stored of in native herbarium species voucher (Hobson specimens be usedWe propose proactively that in data biodiversity generated conservation from the wealth efforts toof cfm?lang=e.General Status Captured Working on Group:25 July 3022013. p. Chauvel,accessible B., F. Dessaint, at http://www.wildspecies.ca/wildspecies2010/home. C. Cardinal-Legrand and F. Bretagnolle. 2006. The historical spread of Ambrosia artemisiifolia L. in France from accelerated destruction of natural habitats and practice herbarium records. Journal of Biogeography 33(4): 665-673. moretarget sustainable specific areas methods for preservation of natural and resources. mitigate Our the Allium method is suitable for generating distributional data at the (Alliaceae) in the Canadian prairie provinces. Botany/Botanique Choi,88(9): H.J. and 787-809. J.H. Cota-Sánchez. 2010. A taxonomic revision of regional, provincial and national scale, and its accuracy can

Cota-Sánchez,Blue Jay 67(2): J.H. 97-104. and V.L. Harms. 2009. The W.P. Fraser herbarium and geographic information system technology. (SASK) of the University of Saskatchewan: past, present and future. be In refined conclusion, with more specimens powerful in georeferencing biological collections methods contain valuable but often hidden information, which Cota-Sánchez, J.H., Polibotánica R.W. Olson 17(1): and 131-138 J. Haraldson. 2004. The type aids the investigation of a wide array of biological and collection of the W.P. Fraser (SASK) herbarium of the University of useSaskatchewan. DNA extracted with a CTAB method adapted for herbarium Cota-Sánchez,specimens J.H., and K. mucilaginous Remarchuk plant and K.tissue. Ubayasena. Plant Molecular 2006. Ready Biology to determine distributional patterns, major collector(s), Reporter 24(2): 161-167. environmental processes. Herbarium holdings help used to map alien species invasion and native species expansion over group. Additionally, they aid in preserving records of past Crawford,the past P.H.C. 100 years? and B.W. Journal Hoagland. of Biogeography 2009. Can 36(4): herbarium 651-661. records be and centers of diversity of a particular flora or plant Darwin, C. 1875. Insectivorous plants. London: John Murray. 462 p. American Scientist et al. 70(3): 260-267. distribution of extinct/rare species, tracking the spread Folkerts, G.W. 1982. TheAtlas gulf of coastSaskatchewan. pitcher plant bogs. changesand/or naturalization in community of composition invasive species and (Chauvel phenology in University Extension Press. 336 p. 2006; Crawford and Hoagland 2009), and determining. 2004). In Fung, K. (Ed.). 1999. University of Saskatchewan,Blue et al JaySaskatoon: 36(2): 71-81. et al. Harms, V.L. 1978. TheChecklist native of carnivorous the vascular plants plants of of Saskatchewan. Saskatchewan and response2004), they to environmentalare useful as an change archival (Primack source of DNA (Cota- the provincially and nationally rare native plants in Saskatchewan. addition to document 2006), type in genetic specimens studies (Cota-Sánchez for conservation Harms, V.L. 2003. et al. 301 p. biology (Maunder et al. et al. University of Saskatchewan, Saskatoon: University Extension Press. Sánchez e.g., scale conversion of forest to agriculture in the Boreal Plains of Suarez and Tsutsui 2004; 1999), Rossman barcoding and (Kress Farr 2006; Hobson, K.A., E.M.Conservation Bayne and Biology S.L. Van16(6): Wilgenburg. 1530-1541. 2002. Large Lehtonen2005), and and for Christenhusz other scientific 2010. andDatabases social of studies, herbarium pitcher-plantSaskatchewan. mosquito. Wyeomyia smithii: behavioural and genetic label information are powerful in identifying regions Istock,aspects; C.A., p. K. 191-204 Tanner In and H. Zimmer. 1983. Habitat selectionPhytotelmata: by the deserving conservation priority based on high diversity terrestrial plants as hosts for aquatic insect communities. Medford, NJ: Plexus Pub. J.H. Frank and L.P. Lounibos (ed.). and high concentrations of rare or narrowly distributed Jennings, D.E. and J.R. Rohr. 2011. A review of the conservation threats to species. This botanical information facilitates the carnivorous plants. Biological Conservation 144(5): 1356-1363. development of management strategies for the protection competition between carnivorous plants and spiders. Proceedings of of unusual species assemblages, such as carnivorous plant Jennings,the Royal D.E., Society J.J. Krupa B 277(1696): T.R. Raffel 3001-3008. and J. R. Rohr. 2010. Evidence for anthropogenic factors. Future studies using information 2008. Plant systematics: a phylogenetic approach. Sunderland, MA: Judd,Sinauer W.S., C.S. Associates. Campbell, 611 E.A. p. Kellogg, P.F. Stephens and M.J. Donoghue. storedpopulations, in natural that history fluctuate collections in response will be to instrumental natural and Juniper, B.E., R.J. Robins and D.M. Joel. 1989. The carnivorous plants. contributions to the understanding of past and present species distribution as well as predicting future patterns London: Harcourt Brace, Jovanovich. 353 p. Proceedings of the in relation to environmental changes. Kress,National W.J., K.J. Academy Wurdack, of Sciences A.A. Zimmer, 102(23): L.A. 8369-8374. Weig and D.D. Janzen. 2005. Use of barcodes to identify flowering plants. in plant molecular systematics: an example from the genus Acknowledgments: Lehtonen,Lindsaea S. and (Lindsaeaceae). J.M. Christenhusz. Biologia 2010. 65(2): Historical 204-208. herbarium specimens Boreal peatlands and plant diversity: what’s there We are thankful to D. Falconer, V. Harms, D. Litwiller, and why it matters. SFMN Research Note Series No. 58. Edmonton, and X. Walker for critical comments on early drafts of the manuscript; Locky D.A. 2010. D. Falconer provided valuable assistance in populating the database. We The rare plants foralso their thank assistance the curators and offor ALTA, facilitating CAN, DAO, the loan and SASKof material. for lending Our gratitude material ofCanada: Saskatchewan Knowledge. Syllogeus Exchange No. and 20. Technology National Extension. Museums of78 Canada.p. goesfor study to Glen and/or and providing Maureen digitizedLee for providing information, photographic and to SASK material personnel for Maher,Ottawa, R.V., G.W. ON: NationalArgus, V.L. Museum Harms andof Natural J.H. Hudson. Sciences. 1979. 55 p. inclusion in this manuscript. This research was partially supported by the Department of Plant Sciences (UofS) and grants from the Canada 1999. Genetic diversity and pedigree for Sophora toromiro Maunder,(Leguminosae): M., A. Culham, a tree A. extinctBordeu, in J. theAllinguillaume wild. Molecular and M. Ecology Wilkinson. 8(5): 725-738. Foundation for Innovation, the Flora of Saskatchewan Association, and the Museums Assistance Program to JHCS.

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Mellichamp, T.L. 2009. Sarraceniaceae: p. 348-363 In Flora of North Taxa list for vascular America Editorial Committee (ed.). Flora of North America North of plants. Mexico Saskatchewan Conservation Data Center. 15 2011. January 2013. Editorial Committee. Schnell, D.E. Electronic 2002. Carnivorous Database plants accessible of the at United http://www.biodiversity. States and Canada. Moore, P.D., Vol. 2002. 8. 1993+. The future New York of cool and temperate Oxford: Flora bogs. of NorthEnvironmental America London:sk.ca/SppList-Inter.htm. Timber Press. 468 Captured p. on Conservation 29(1): 3-20. Schwab-Moe, B. 1999. The distribution and habitats of the linear-leaved Natural Resources of Canada. 2012. The Atlas of Canada – Toporama. Sundew, Drosera linearis Goldie. M.Sc. Thesis, Department of Plant

html. Captured on 1 June 2012 Specify Software. 2012. Accessible at www.specifysoftware.org. Captured Accessible at http://atlas.nrcan.gc.ca/site/english/toporama/index.Vascular plants of continental Northwest onSciences, 13 June University 2012. of Saskatchewan, Saskatoon, SK. 165 p. Territories. National Museums of Canada. Ottawa, ON: National Suarez, A. and N. D. Tsutsui. 2004. The value of museum collections for Porsild,Museum E.A. and of Natural W.J. Cody. Sciences. 1980. 653 p. research and society. BioScience 54(1): 66-74.

Primack,response D., C. toImbres, warming R.B. inPrimack, Boston. A.J. American Miller-Rushing Journal and of Botany P. Del Tredici. 91(8): 1260-1264.2004. Herbarium specimens demonstrate earlier flowering times in Rochefort, L. 2000. Sphagnum: The Bryologist 103(3): 503-508. : April 2013 a keystone genus in habitat restoration. : August 2013 In 8th International Mycological Congress Abstracts Received : September 2013 Rossman, A.Y. and D.F. Farr. 2006. The value of herbaria in the DNA age. Accepted : Angelo G. Manzatto [Abstract]: p. 415 Published online Book 2, Cairns, Australia. Editorial responsibility Appendix 1.

Chronological list of historical accessions of carnivorous plants by COLL.genus and species collectedLATITUDE in Saskatchewan LONGITUDE prior to 1950.ACCESSION TAXON LOCALITY COLLECTOR DATE NO. (°N) (°W) NO. Drosera anglica Prince Albert 53.2 -105.7 D. anglica Prince Albert 53.2 -105.5 Fraser, W.P. 23/06/1933 SASK 12955 D. anglica Prince Albert 53 -105 Fraser, W.P. 23/06/1933 SASK 91996 D. anglica Prince Albert 53.2 -105.7 Fraser, W.P. 12/07/1933 SASK 12954 D. anglica Prince Albert 53.2 -105.8 Fraser, W.P. 25/08/1933 SASK12957 D. anglica Breitung, A.J. 52.6 -103.917 Fraser, W.P. 31/08/1933 SASK12592 D. anglica 6854 59.125 -109.317 DAO 568305 McKague 10/08/1934 SASK 12956 D. anglica Breitung, A.J. 28 52.61667 -103.933 CAN 504618 Lake Athabasca Raup, H.M. 13/08/1935 D. anglica Prince Albert Furniss, O.C. 2.216667 -4.40694 McKague 12/07/1936 D. anglica Prince Albert Furniss, O.C. 860 53.2 -105.7 19/07/1941 SASK 144018 D. anglica Prince Albert 53.2 -105.767 ALTA 58102 29/06/1941 SASK 77299 D. linearis Prince Albert 53.2 -105.8 Fraser, W.P. 23/07/1936 D. linearis Prince Albert 53.2 -105.8 Fraser, W.P. 23/06/1933 SASK 91997 D. linearis Breitung, A.J. 52.6 -103.9 Fraser, W.P. 12/07/1933 SASK 12947 D. linearis Breitung, A.J. 52.6 -103.9 McKague 10/08/1934 SASK 12950 D. linearis Breitung, A.J. 29 52.61667 -103.933 CAN 504400 McKague 10/08/1934 SASK 12951 D. linearis Prince Albert Furniss, O.C. 53.2 -105.767 McKague 12/07/1936 D. linearis Prince Albert Furniss, O.C. 858 53.2 -105.8 19/07/1941 SASK 144019 D. linearis Breitung, A.J. 52.55 -104.05 ALTA 58103 19/07/1941 SASK 77298 D. rotundufolia Speddington Russell, R.C. 52.8 -103.7 Wallwort 22/07/1941 D. rotundifolia 54 -106.2 14/07/1926 SASK 91998 D. rotundifolia 53.9 -106.1 Waskesiu Lake Fraser, W.P. 00/00/1930 SASK 12971 D. rotundifolia 53.9 -106.1 Waskesiu Lake Fraser, W.P. 00/00/1930 SASK 92000 D. rotundifolia 54 -106.2 Waskesiu Lake Fraser, W.P. 00/00/1930 SASK 12966 D. rotundifolia 53.0 -106.1 Waskesiu Lake Fraser, W.P. 08/07/1930 SASK 12967 D. rotundifolia 53.9 -106.1 Waskesiu Lake Fraser, W.P. 07/08/1930 SASK 149965 D. rotundifolia 54 -106.2 Waskesiu Lake Fraser, W.P. 21/07/1932 SASK 91999 D. rotundifolia Prince Albert 53.2 -105.7 Waskesiu Lake Fraser, W.P. 26/07/1932 SASK 12969 D. rotundifolia Breitung, A.J. 52.6 -103.9 Fraser, W.P. 12/07/1933 SASK 12963 D. rotundifolia Prince Albert 53.2 -105.767 ALTA 58104 McKague 10/08/1934 SASK 12964 D. rotundifolia Prince Albert 53.2 -106 Fraser, W.P. 23/07/1936 D. rotundifolia Ledingham, G.F. 54 -106.1 Fraser, W.P. 23/07/1936 SASK 106447 D. rotundifolia Prince Albert Furniss, O.C. 855 53.2 -106 Waskesiu Lake 07/03/1937 SASK 106448 D. rotundifolia Prince Albert Furniss, O.C. 53.2 -105.8 19/07/1941 SASK 77297 D. rotundifolia Breitung, A.J. 54.00005 -106 ALTA 123407 19/07/1941 SASK 144078 Pinguicola villosa 6307 59.6 -109.217 CAN 99311 22/07/1941 P. vulgaris MacDowall 53 -106 Lake Athabasca Raup, H.M. 24/07/1935 P. vulgaris Prince Albert 53.2 -105.8 Fraser, W.P. 07/10/1932 SASK 67551 P. vulgaris Prince Albert 53.2 -105.8 Fraser, W.P. 01/07/1935 SASK 21146 P. vulgaris Prince Albert 53.2 -105.8 Fraser, W.P. 01/07/1935 SASK 21147 P. vulgaris Prince Albert Furniss, O.C. 53.2 -105.767 Fraser, W.P. 27/07/1937 SASK 116259 Sarracenia purpurea Prince Albert 53.2 -105.7 14/06/1940 SASK 144120 Thompson, W.P. 10/06/1911 SASK 12946

892 Baalim et al. | Distribution of carnivorous plants of Saskatchewan, Canada

Appendix 1. Continued.

COLL. LATITUDE LONGITUDE ACCESSION TAXON LOCALITY COLLECTOR DATE NO. (°N) (°W) NO. S. purpurea Prince Albert 53.2 -105.7 S. purpurea Prince Albert 53.2 -103.7 Fraser, W.P. & Russell, R.C. 10/06/1932 SASK 91991 S. purpurea Prince Albert 53.2 -105.7 Fraser, W.P. & Russell, R.C. 06/12/1932 SASK 91992 S. purpurea Prince Albert Russell, R. C. 53.2 -105.767 ALTA 58100 Fraser, W.P. & Russell, R.C. 12/07/1932 SASK 12936 S. purpurea Breitung, A.J. 52.6 -103.9 12/07/1932 S. purpurea 59.05 -109.567 ALTA 9968 McKague 06/09/1934 SASK 12943 S. purpurea Prince Albert 53.2 -105.7 Lake Athabasca Raup, H.M. 24/08/1935 S. purpurea Prince Albert 53.2 -105.7 Fraser, W.P. 07/05/1939 SASK 12940 S. purpurea Prince Albert Furniss, O.C. 870 53.2 -105.7 Fraser, W.P. 07/05/1939 SASK 12945 S. purpurea Prince Albert Furniss, O.C. 53.2 -105.7 21/06/1940 SASK 77258 S. purpurea Prince Albert Furniss, O.C. 53.2 -105.7 22/06/1940 SASK 145051 S. purpurea Chelan Robinson, D.R. 52.6 -103.4 17/08/1941 SASK 145049 Utricularia intermedia Prince Albert 53.2 -105.8 15/06/1944 SASK 91993 U. intermedia Prince Albert 53.2 -105.8 Fraser, W.P. 23/06/1933 SASK 21168 U. intermedia Dahlton Breitung, A.J. 52.5 -104.1 Fraser, W.P. 23/06/1933 SASK 86764 U. intermedia Russell, R.C. 51.9 -106.8 07/05/1936 SASK 21166 U. intermedia 51.9 -106.8 Pike Lake 07/07/1936 SASK 116266 U. intermedia Prince Albert 53.2 -105.8 Pike Lake Mead, H.W. 28/08/1936 SASK 86765 U. intermedia Prince Albert Furniss, O.C. 1659 53.2 -105.8 Fraser, W.P. 24/06/1940 SASK 21169 U. macrorhiza 52.1 -106.6 29/06/1941 SASK 144769 U. macrorhiza 52.1 -106.6 Saskatoon Fraser, W.P. 00/08/1922 SASK 86773 U. macrorhiza Buttress 50.20003 -105.634 ALTA 65412 Saskatoon Fraser, W.P. 00/08/1922 SASK 86774 U. macrorhiza Duncairn Russell, R.C. 51.8 -106.5 McCalla, W.C. 18/07/1923 U. macrorhiza 52.1 -106.6 02/08/1924 SASK 86772 U. macrorhiza Sutherland Arnason, T.J. 52.1 -106.6 Saskatoon Cameron, W.H. 27/07/1927 SASK 21178 U. macrorhiza 52.1 -106.6 17/07/1930 SASK 21195 U. macrorhiza 53.9 -106.2 Saskatoon Fraser, W.P. 17/07/1930 SASK 21196 U. macrorhiza 52.6 -105.6 Waskesiu Lake Fraser, W.P. 00/08/1930 SASK 21180 U. macrorhiza Somme Russell, R.C. 52.1 -104.8 Wakaw Fraser, W.P. 30/06/1933 SASK 21179 U. macrorhiza St. Gregor Russell, R.C. 52.2 -104.8 10/07/1934 SASK 21197 U. macrorhiza Bolton, J.L. 109 49.6 -109.8 10/07/1934 SASK 86771 U. macrorhiza Bolton, J.L. 49.6 -109.8 Cypress Hills 15/08/1936 SASK 21198 U. macrorhiza 54.1 -105.8 Cypress Hills 15/08/1936 SASK 116280 U. macrorhiza Russell, R.C. 53.6 -106 Montreal Lake Fraser, W.P. 15/07/1938 SASK 21182 U. macrorhiza Russell, R.C. 53.6 -105.9 Emma Lake 18/07/1940 SASK 86770 U. macrorhiza Campbell, J.A. 49.1 -108.1 Emma Lake 26/07/1940 SASK 21193 U. macrorhiza Russell, R.C. 54.0 -106.2 Besnard Lake 20/08/1941 SASK 116278 U. macrorhiza Russell, R.C. 50.3 -102.3 Heart Lakes 17/06/1944 SASK 86769 U. minor Dahlton Breitung, A.J. 52.5 -104.1 Whitewood 08/03/1945 SASK 86768 U. minor Dahlton Breitung, A.J. 52.5 -104.1 07/05/1936 SASK 21174 U. minor Dahlton Breitung, A.J. 52.5 -104.1 07/05/1936 SASK 21176 00/08/1936 SASK 21175

893