Range Extensions of 35 Bryophyte

Range Extensions of 35 Bryophyte Species in the Black Spruce– Feather Moss Forest of Western Quebec, Canada MARIoN BARBé 1, 4 , L oUIS DUBoIS 2, J EAN FAUBERT 3, M ARTIN LAvoIE 2, Y vES BERGERoN 1, and NICoLE J. F ENToN 1

1Université du Québec en Abitibi-Témiscamingue, Institut de recherche sur les forêts, 445, boul. de l’Université, Rouyn-Noranda, Quebec J9X 4E5 Canada 2Université Laval, Département de géographie, 2405, rue de la Terrasse, Québec, Quebec G1v 0A6 Canada 3Société québécoise de bryologie, 47, 4 ème Rang Est, Saint-valérien-de-Rimouski, Quebec G0L 4E0 Canada 4Corresponding author: [email protected]

Barbé, Marion, Louis Dubois, Jean Faubert, Martin Lavoie, Yves Bergeron, and Nicole J. Fenton. 2017. Range extensions of 35 bryophyte species in the Black Spruce–feather moss forest of western Quebec, Canada. Canadian Field-Naturalist 131(3): 258 –269. https://doi.org/10.22621/cfn.v131i3.1901 Although the North American bryophyte flora are relatively well known, bryophytes of the Black Spruce–feather moss forest in the Nord-du-Québec administrative region, especially its southern portion (49–51°N, 74–79°W), remain under-sampled. Here, we report 169 bryophyte taxa for this region, of which 35 (14 true mosses, 20 liverworts, one sphagnum) represent note - worthy records, including 20 taxa new for the region. These new occurrences close several gaps in distribution in the study area and, more broadly, in the boreal Black Spruce ( Picea mariana ) forest of adjacent ontario. Microhabitat preferences of the species are also documented. This work represents a substantial contribution to knowledge of the bryophyte flora, which will help refine protection priority ranks of species of Quebec and Labrador. Key Words: Boreal forest; bryoflora; bryo-geography; liverwort; true moss; sphagna; northern Quebec Malgré une bonne connaissance globale de la bryoflore nord-américaine, la pessière noire à mousses de certaines régions telles que le Nord-du-Québec et notamment sa partie méridionale (49–51°N, 74–79°W) demeure sous-échantillonnée. Nous rapportons 169 taxons bryophytiques dans cette région, dont 35 (14 mousses, 20 hépatiques et une sphaigne) représentent des ajouts substan - tiels à la flore, incluant même 20 nouveaux taxons pour le territoire considéré. Ces récoltes permettent de relier les aires de répartition jusque-là disjointes de plusieurs taxons en pessière noire à mousses au Québec, mais aussi dans la province jouxtante de l’ontario. Les préférences des espèces en termes de microhabitats sont aussi décrites. Ce travail contribue à améliorer les connais - sances sur la bryoflore et permettra de redéfinir les rangs de priorité pour la conservation des espèces au Québec et Labrador. Mots-clés: aire de répartition; bryoflore; forêt boréale; hépatiques; mousses; sphaignes; Nord-du-Québec

Introduction Bryophytes (liverworts, true mosses, and sphagna), ral wildfires and anthropogenic exploitation (forest along with lichens, dominate the coniferous boreal for - harvest, mining, hydroelectric development). These cu - est in terms of biomass, species richness (Turetsky et al. mulative disturbances of the landscape threaten spe cies 2012), and net primary productivity ( Bisbee et al. 2001; that are ill-adapted to anthropogenic environments, in - Proctor 2011) . They form a continuous carpet several cluding many bryophytes (Fenton and Frego 2005; centimetres thick and inhabit a variety of microhabitats Hylander et al. 2005; Caners et al. 2013). (Dynesius and Hylander 2007). Bryophytes represent Since the publication of the Catalogue des bryophytes 25% of the plant diversity of Quebec (Faubert et al. du Québec et du Labrador (Faubert 2007), the number 2010). In 2016, the database of the bryophytes of Que - of bryophyte species documented in Quebec and Lab - bec–Labrador listed 231 species of liverworts, 582 spe - rador has continued to grow (Gauthier 2011; Moisan cies of mosses, and 62 species of Sphagnum (Faubert et and Pellerin 2011; Faubert et al. 2012; Faubert and al. 2014+). However, the distributional ranges of some Gagnon 2013). New occurrences are continuously being species are only partly defined, and the bryophyte flora compiled in the online database of the bryophytes of is unknown in certain areas (Faubert and Gagnon 2013). Quebec–Labrador (Faubert et al. 2014+), contributing This is the case for the administrative region of Abi - to continuous updating of the bryophyte flora (Faubert tibi- Témiscamingue and the adjacent southern portion 2012–2014). The current study contributes to our under - of the Nord-du-Québec administrative region, which standing of bryophyte distributional ranges at the scale have been neglected in terms of bryophyte sampling of the boreal Black Spruce ( Picea mariana (Miller) com pared with other regions. Understanding the fre - Britton, Sterns & Poggenburgh ) forest of Quebec–Lab - quency and distribution of species is of primary im - rador. We describe the bryophyte community of this for - portance in establishing conservation plans and in im - est bioclimatic domain, including microhabitat prefer - plementing resource management practices in these ences, which may permit better forest development regions, where boreal forests are disturbed both by nat- prac tices and bryophyte conservation.

258 ©The Ottawa Field-Naturalists’ Club (2017) 2017 BARBé ET AL .: B RYoPHYTE RANGE EXTENSIoNS IN QUEBEC 259

Study Area The study area is located in the Black Spruce–feather The study area covers 73 197 km² (48°83'N to moss forest bioclimatic domain that extends over 154 50°71'N and 74°50'W to 79°69'W) in the southern por - 184 km² in Quebec (Grondin 1996). Forest stands are tion of the Nord-du-Québec administrative region of dominated by Black Spruce, Jack Pine ( Pinus banksiana western Quebec (Figure 1). After the retreat of the Lau - Lambert), Trembling Aspen ( Populus tremuloides Mi - rentide Ice Sheet, the area was covered by the proglacial chaux), Balsam Fir ( Abies balsamea (L.) Miller), and lakes Barlow and ojibway, which existed 11 500 and Paper Birch ( Betula papyrifera Marshall). The under - 7900 radio carbon years before present, respectively storey is dominated by ericaceous shrubs on a ground (vincent and Hardy 1977). Sedimentation in the lakes cover of bryophytes (Saucier et al. 2009). The natural generated a layer of clay 10–60 m thick that forms the dynamics of these forests are driven primarily by stand- soils of the “clay belt” of northeastern ontario and replacing wildfires. The fire cycle has been estimated at northwestern Quebec. There is little topographic varia - 398 years (Bergeron et al. 2004), and the average age of tion in the region, with elevations ranging from 200 to the forest is over 150 years. Average annual temperature 300 m above sea level. and precipitation (1981–2010) are 1°C and 928 mm,

FIGURE 1. a. Location of the study area (white square) in the Nord-du-Québec administrative region of western Quebec, Canada, within the boreal Black Spruce ( Picea mariana ) forest bioclimatic domain (dark grey). b. The six wildfires sampled (dark grey shapes) in Black Spruce forest bioclimatic domain (light gray zone; adapted from Payette and Bouchard 2001). Triangles represent main cities. c. Black rectangles represent sample plots of 50 m² located along a transect crossing the residual patch (light grey) from burned area (B) to edge (E) and core (C). 260 THE CANADIAN FIELD -N ATURALIST vol. 131

respectively (Environment Canada 2017). The region is located using a handheld global positioning system re - characterized by long winters, with 313 cm of snowfall ceiver (Garmin GPSmap 62, olathe, Kansas, USA). an nually, and a short growing season of 140–160 days. In each 50-m 2 plot, the bryophyte community was sampled using a modified form of “floristic habitat Methods sampling” (Newmaster et al. 2005), which consists of Bryophyte Sampling sampling all the bryophytes present in all microhabitats Bryophytes were sampled within the footprints of six (e.g., coarse woody debris, tree bases, peat mounds, natural wildfires (Figure 1b) varying in age, size, and water holes) . This method was used to ensure that all origin and used for a study on post-fire residual forest small non-visible species were captured. vouchers of patches (Barbé et al. 2016, 2017). Within each wildfire all specimens are stored at the Université du Québec footprint, we identified five residual patches (unburned en Abitibi-Témiscamingue (Rouyn-Noranda, Canada). forest areas) and three burned areas for a total of 30 Nomenclature follows Faubert et al. (2014+) except residual patches aged 36–3400 years (time since last for Sphagnum subtile (Russ.) Warnst. (Flora of North wildfire) and 18 burned matrices aged 10–44 years America Editorial Committee 2007). (time since last wildfire). The age of the patches was All samples were dried and later identified to species estimated by coring 10 dominant trees; if the 10 domi - level using a stereomicroscope and a compound light nant trees were approaching their maximum lifespan microscope following the specimen preparation and (> 180 years old for Black Spruce; Simard et al. 2007) identification method described in Faubert (2012). the age of the patch was determined by 14 C dating of Dam aged, senescent, or immature specimens were charcoal particles extracted from the mineral soil. Age iden ti fied only to genus level. The microhabitat in which of burned areas corresponds to time since the last fire each species was found was qualitatively compared with determined from Société de protection des forêts con - data from Flore des Bryophytes du Québec–Labrador tre le feu digital maps (SoPFEU 2011). All residual (Faubert 2012–2014; herein shortened as “Flora”) to patches were chosen based on the following criteria: determine whether a species was specific to certain Black Spruce dominance, accessibility (< 600 m from microhabitats in the study area. the logging road), flat topography, and no complete sub- Distribution Maps mergence of the soil except in local depressions (water - Provincial distribution maps were generated for spe - holes). No bogs or fens were sampled, but post-fire cies found in this study whose ranges differed from res idual patches located in paludified (i.e., natural suc - those previously known in Quebec. New occurrences cession to peatland; Crawford et al. 2003) Black Spruce were compared with those detailed in the open-access forests were included. Consequently, we sampled sites BRYoQUEL participative online database of the bryo- encompassing the range of natural Black Spruce forest phytes of Quebec–Labrador (Faubert et al. 2014+). succession: from recently burned to paludified areas. New occurrences were also compared with document - Patches varied in size (0.05–11.1 ha) and forest struc - 3 ed occurrences from the neighbouring province of on - ture (e.g., 7.4–109 m /ha of coarse woody debris). More tario (Ireland and Ley 1992; Ley and Crowe 1999; details about the characteristics of the patches are pre - CNALH 2017). Maps were generated using the geo - sented in Barbé et al . (2017). graphic information system, ArcGis 10.3.1 (Environ - Rectangular plots (50 m²) were used to sample the mental Systems Research Institute [ESRI], Redlands, bry ophyte communities in each residual patch and California, USA). original map layers were from the burned area. At each location, a north–south linear tran - GéoIndex+ platform of the Geographic and Statistic sect was established that included the burned zone, edge, Information Centre (GéoStat Centre) of the Université and core positions (Figure 1c). In patches smaller than Laval created with data from Statistics Canada, geo - 1 ha, five plots were placed along the transect: two in graphic division, DMTI Spatial Inc. (Richmond Hill, the surrounding burned zone, two straddling each edge of the patch, and one in the core. In patches over 1 ha, ontario, Canada), and ESRI. The projection used for a second core plot was added. Plots were 10–200 m all maps was NAD83 CSRS MTM 10. apart. In each wildfire footprint, three additional 50-m² Data Analyses plots of burned area were placed 200–850 m from resi - Provincial occurrences, extracted from Faubert et d ual patches. Size, number, orientation, and placement al. (2014+), are “previously documented occurrences” of sampling plots were chosen to include all microhab - and were classified into four categories: rare (< 5 oc - itats at each site, from more humid and cold microhab - currences), infrequent (5–10), uncommon (11–30), and itats found at the northern edge and in residual patch common (> 30; Table 1). There was no minimum dis - cores, to warmer and drier microhabitats found at the tance between occurrences. The local occurrences from south ern edge and in burned areas. In total, the bry - this study were treated as “new occurrences”. Local ophyte community was sampled over 9300 m²: 108 occurrences refer to the record of one species in a resid - plots in 30 residual patches (48 cores and 60 edges) and ual patch or a burned area. Indeed, even though a spe - 78 plots in burned areas (2 × 30 residual patches plus cies was found several times in the same residual patch 3 × 6 wildfires). The four corners of each plot were geo - or burned area (i.e., several records in the same 50-m² 2017 BARBé ET AL .: B RYoPHYTE RANGE EXTENSIoNS IN QUEBEC 261 f - o o

plot), only one occurrence was drawn on the map (one p a s r

T d / o d e l cross) to avoid overloading maps with superimposed y W W P e l f ‡ v

s d E T D D r w W W W W t e / / / / r crosses. Local occurrences followed the same categor - e e a h o s C C T D D C D C C D t t

i c

b N

b ies as provincial ones but were based on the number of e o o

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were used to refine species occurrences in Quebec. For P

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f example, a species represented by 28 provincial occur -

i o

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n o o o o rences plus 10 new local occurrences was updated from n

i i y i i n

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uncommon to common in Quebec. Each occurrence i

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p (provincial as well as local) was presented regardless o r r i

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of plot and site location. Species were grouped by their n D D

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provincial and local occurrences: group 1: locally un - c

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common/infrequent species already recorded near the e

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study area; group 2: provincially common to rare spe - 2

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cies only recorded sporadically near the study area;

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d d d d d d Microhabitats of species in the study area were com - d

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pared with those documented in Flora (Faubert 2012–

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2014), which were compiled from an exhaustive list of x x x x x x x

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preferential microhabitats found in the literature ( Schus -

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ter 1966–1992; Crum and Anderson 1981; Ireland 1982)

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plus additional microhabitats from specimens collected n g r

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in Quebec. Thus, we compared species’ microhabitat

h a c

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m preferences between boreal Black Spruce forest and

a c →

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e r

L c l

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habitats in the rest of the province and the species’ →

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t p n n n n n

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Canadian distribution to identify differences in micro -

e Q

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habitat preferences in the Black Spruce forest of west - q

c c c c

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n n n n n n o o o o o a

d ern Quebec. Microhabitats were classified as humus (T:

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terricolous species living on soil and litter), peat (B:

- s species living among sphagna or on exposed peat in

r h

w 2 4 2 3 9 1 2 3 3 9 2 4

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1 9 paludified areas), dead wood (DW: facultative or obli -

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o y species on living trees and shrubs), and rock (R: saxi -

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l colous species). e

t c s s s h

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Q e 5 8 0 8 2 9 3 5 r e

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Bryophyte Community and Species Distribution

e n

Bryophytes were sampled in 11 036 microhabitats, x

i s

e e

o i

each containing on average four species (range 0–20) e

e i p l

i t i

e f

n r

f i o 2 3 3 3 2

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for a total of 169 species encountered (61 liverworts, e

r l

a r a

p r

g f

f p S

90 mosses, and 18 sphagna; Appendix S1). Locally, h

a c

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118 species (70% of all species) were common (found w

s e

in more than 30 plots); the remaining species occurred l

n t

i a

, y

more sporadically over the sampling area.

n i

p d

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Post-fire residual patches were dominated by the fea -

r d .

e ;

n d a

t ther mosses, Hylocomium splendens (Hedw.) Schimp., a

e a

a l

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r n l b

a Pleurozium schreberi (Willd. ex Brid.) Mitt., and Ptil - a

a o .

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currences of the acrocarp species, Dicranum fus cescens l a y

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pulcherrimum (Weber) Hampe. The wettest sites also E

a e e

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o supported Au la comnium palustre (Hedw.) Schwägr.,

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a s s

P P P I I H H H D C B M B Q Sanionia uncinata (Hedw.) Loeske, and Warnstorfia T r 262 THE CANADIAN FIELD -N ATURALIST vol. 131

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N § † * ‡ T S T C U C L C C C C F F K S L S S S L S O M M L L 2017 BARBé ET AL .: B RYoPHYTE RANGE EXTENSIoNS IN QUEBEC 263 . a s i e e fluitans (Hedw.) Loeske, whereas Ceratodon purpureus c g n o e

p (Hedw.) Brid., Polytrichum juniperinum Hedw., and r r y l u

a Pohlia nutans (Hedw.) Lindb. were found mainly in c c C

o .

burned areas, which were also the driest sites. In addi - a

w : e

a tion, deep mats of sphagna ( Sphagnum capillifolium n e

r e

a (Ehrh.) Hedw., Sphagnum fallax H. Klinggr., Sphag - r

a y

s d num magellanicum Brid.) were found, as many of the n u t g s i sites sampled were undergoing paludification ( Fenton

e s h

t et al. 2005) . Some of the species des cribed as common u

l p m in the coniferous boreal forests of Quebec in Faubert ; o s r f e

(2012–2014) were under-represented (≤ 15 occurrences) c n m e k in our samples (e.g., Barbilophozia hatcheri (A. Evans) r

r 2 u 3

c Loeske, Bryum capillare Hedw., Tomenthypnum nitens – c 6 o

(Hedw.) Loeske). 2

d d e t

e In addition to these locally common species, 35 spe - n d e r cies (14 true mosses, 20 liverworts, and one sphagnum) o m c u e c r represent noteworthy records (Table 1). of these, four

o y d l

species (group 1) were locally uncommon or infrequent, s y u l s o but were expected to be found because they have already i u v o e i been recorded 25 km from the study area in Quebec r v p e

s and 50–150 km from the study area in on tario: Caly - p e

i c

w pogeia sphagnicola (Arnell & J. Perss.) Warnst. & e o p

h s .

s Loeske, Fuscocephaloziopsis pleniceps (Austin) váňa

t s u n t c

e & L. Söderstr., Lophozia guttulata (Lindb. & Arnell) A. o c u o D q

Evans, and Sphenolobus hellerianus (Nees ex Lindenb.) . e y s r l f u e

n Steph. (Figure 2). Ten other species (group 2), includ - n k i i

a l r i ing four true mosses ( Brachythecium erythrorrhizon r o s

e e l n i l

o Schimp., B. starkei , (Brid.) Schimp., Isopterygiopsis c e e h m

s muelleriana (Schimp.) Z. Iwats., Polytrichum commune s m

u o e b

c Hedw. var. perigoniale (Michx.) Hampe and six liv - h o t n l

u o e

erworts ( Cepha loziella hampeana (Nees) Schiffn. ex r n y e e l l h h Loeske, Kurzia pauciflora (Dicks.) Grolle, Lophozia a p i w

c S t ascendens (Warnst.) R.M. Schust., L. bicrenata (Schmi - n . s i d e v

r ,

o del) Dumort., Odontoschisma francisci (Hook.) L. o a r f t

p a

s Söder str. & váňa , Schisto chylopsis laxa (Lindb.) Kon - l s g u o n t i t stant.) were provincially common to rare, but had been s m u i

r g r

e recorded only sporadically near the study area (one to a h i m t z a o three occurrences 20–115 km; Figure 3). Some of these o c e h

f , p

– moss species are found west of the study area, in on - 1 o

e p c L

tario (i.e., Brachythecium erythrorrhizon , B. starkei , u . u r c o

r p , Poly trichum commune var. perigoniale ), where they g s S

p f k e

o have already been recorded 50–200 km from the prov - c

c i s a l e n

i incial border (Ireland and Ley 1992; Ley and Crowe e B c l

l e p

a 1999; CNALH 2017). p s e s i

r s r

o Finally, 20 species (group 3) are new occurrences for p u b o o

i f e

z the study area, with range extensions from 75 km to

h e o t l

h t

a more than 670 km in Quebec–Labrador. These include s

t h f n p o

nine true mosses ( Campylium protensum (Brid.) Kindb., e e s s c e e o

r Dicranum fulvum Hook., Helodium blandowii (F. g c p n s e a u r

r Weber & D. Mohr) Warnst. var. blandowii , Hypnum

F l a

. a e curvifolium Hedw., Isopterygiopsis pulchella (Hedw.) r b n

a , o

i a

t Z. Iwats., Pohlia elongata Hedw. var. elongata , P. d l u e o b d c

i sphagnicola (Bruch & Schimp.) Broth., Thuidium rec - i a r t n h s g s i ognitum (Hedw.) Lindb., Ulota crispa (Hedw.) Brid.),

a e D h h .

p 10 liverworts ( Calypogeia suecica (Arnell & J. Perss.) 2 s T

E Müll. Frib., Cephaloziella elachista (J.B. Jack) Schiffn., R

U C. spinigera (Lindb.) Jörg., Chiloscyphus coadunatus G I

F (Sw.) R.M. Schust. & J.J. Engel var. rivularis (Raddi) 264 THE CANADIAN FIELD -N ATURALIST vol. 131 . t e a s e m i h

e z u t t i c

r o c e n e h r e e b p e r h u t r o h a y u L

w F h c .

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a o e t ,

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c n t . o

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s a T

m m . e m u o a i h l c h x

T c c a

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r h . v r y d o u h w

r ,

c c p e a c o

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d h s e

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r .

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. o . l e s r 3 a e n a D E R U G I F 2017 BARBé ET AL .: B RYoPHYTE RANGE EXTENSIoNS IN QUEBEC 265

Frisvoll, Elvebakk, Flatberg & okland, Fuscocepha - for these species. These occurrences, together with those loziopsis loitlesbergeri (Schiffn.) váňa & L. Söderstr., already documented from eastern Quebec, suggest their Lophozia silvicola H. Buch, Mesoptychia heterocolpos continuous distributional ranges extend throughout the (Thed. ex Hartm.) L. Söderstr. & váňa var. heterocolpos , Black Spruce forest of Quebec–Labrador. Furthermore, Mesoptychia rutheana (Limpr.) L. Söderstr. & váňa, the dispersed and numerous locations of these species Scapania apiculata Spruce, S. uliginosa (Lindenb.) indicate that they are common but under-sampled in the Dumort.), and one sphagnum ( Sphagnum tenerum Sull. province. Further extension of their continuous distri - & Lesq. ex Sull.; Figure 4). Considering occurrences butional ranges to all of Quebec–Labrador may be pos - from ontario, we report a 50–200 km eastward exten - sible, but more sampling is needed to determine their sion of the distributional range of Helodium blandowii true distributions. var. blandowii , Isopterigiopsis pulchella , Pohlia sphag - our results also extend to the entire Black Spruce nicola , and Thuidium recognitum (Ireland and Ley forest domain the known distributions of the liverworts 1992; CNALH 2017). Cephaloziella elachista, C. hampeana , C. spinigera, Hypnum fauriei Cardot was sampled once in the Chiloscyphus coadnatus var. rivularis, Fuscocepha - study area. No map was produced for this species be - loziopsis loitlesbergeri, Kurzia pauciflora, and Odon - cause its distribution in the province is unknown as a toschisma francisci, and of the true mosses Brachythe - result of its recent separation from H. fertile Sendtn. cium starkei and Pohlia sphagnicola . However, some of (Faubert 2014). them ( Cephaloziella elachista, C. spinigera, Chiloscy - phus coadnatus var. rivularis, F. loitlesbergeri, Lopho - Bryophyte Microhabitat Preferences zia silvicola, and O. francisci ) have only rarely been The microhabitat preferences of the 35 species dis - recorded in eastern Quebec; further sampling is need - cussed above were more diverse than reported previous - ed to confirm their presence across the province. In ly in Flora (Faubert 2012–2014; Table 1). For ex ample, light of the 92 and 56 new occurrences of B. starkei and two mosses (Pohlia sphagnicola and Caly pogeia sphag - C. hampeana , respectively, as well as the 634 new oc - nicola ) and several liverworts were found in a greater currences of P. sphagnicola, the status of these species variety of microhabitats than the exclusive peat micro - in the province should be changed from uncommon to habitat mentioned in Flora (Faubert 2012–2014). Eight - common. Liverworts, especially Cephaloziellaceae and een species were recorded on tree or shrub bases in K. pauciflora , are minute (< 1 mm wide shoots) and Black Spruce forest, especially Picea mariana and Rho - especially difficult to detect, which has probably con - dodendron groenlandicum (oeder) Kron & Judd (Table tributed to the underestimation of their frequency and 1), although they were not described as corticolous or distribution. Directed sampling efforts focusing on these epiphytic at the provincial scale. Half were associated taxa might help distinguish between a lack of sam - with only one tree or shrub species, but multiple hosts pling versus true rarity in other regions. The distribu - were also identified for many bryophyte species (e.g., tional ranges of the true mosses B. starkei and P. sphag - Pohlia sphagnicola , Brachythecium starkei , Cepha - nicola are unclear because of past misidentifications loziella elachista ; data not shown). Similarly, numer - (Faubert 2012–2014). Indeed, in the past, the few and ous species not previously identified as epixylics in confusing diagnostic characters discriminating these Flora (Faubert 2012–2014) were found on dead wood species from others of the same genera have led to con - (Table 1; e.g., Brachythecium erythrorrhizon, Dicranum fusion between B. starkei and B. curtum (Lindb.) Limpr., fulvum ). P. sphagnicola and P. nutans , and between Sphagnum tenerum and S. capillifolium. Discussion Lophozia bicrenata also needs attention. This un - Among the 169 bryophytes species identified, we common and minute species, typical of disturbed land - have documented 20 new species for the study area and scapes, is present in dispersed locations across the en - increased the understanding of the distributional range tire province. It is possible that this species is found for 15 others. our results suggest that these species may province-wide, but this possibility can only be addressed be more common in Quebec–Labrador and especially by additional sampling to discriminate between true rar - in the bioclimatic domain of the Black Spruce-feather ity and under-collection. moss forest, than previous occurrences indicate. We Distributional Ranges Extended to the North and West examine here the noteworthy occurrences within groups The few new reported occurrences of Dicranum ful - 1, 2, and 3 as identified above. vum and Thuidium recognitum (three and five, respec - Distributional Ranges Extended to the Entire Boreal tively) extend the provincial distribution of these spe cies Black Spruce Forest Bioclimatic Domain 135 km to the north and 75 km to the west, respective - Even though already documented in the study area, ly. These species are common in the province, but have we extended the known distribution of four species of not been previously recorded in the study area. Helodi - liverwort ( Calypogeia sphagnicola , Fuscocephaloziop - um blandowii var. blandowii, Hypnum curvifolium , and sis pleniceps , Lophozia guttulata, and Sphenolobus Ulota crispa are also common. However, we do not hellerianus ), with 82 to more than 200 new occurrences suggest extension of their continuous distributions be - 266 THE CANADIAN FIELD -N ATURALIST vol. 131 e e e e a y l r r i h h l s t a a t

r h u

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D u o n a . c w w c e o A e o e r c l 4 n a a n e d E R U G I F 2017 BARBé ET AL .: B RYoPHYTE RANGE EXTENSIoNS IN QUEBEC 267 . d . p e

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cause only one occurrence of each species was record - species had singular preferences in terms of habitat in ed in the study area, and the distance from the rest of the Black Spruce forest compared with other ecosys - their known Quebec range is substantial (325, 270, and tems across the province. Species found in our study 165 km, respectively). occupied, in general, more diverse microhabitats and Finally, the few occurrences (generally ˂ 10) of the particularly more woody ones. remaining 14 species do not suggest a continuous dis - The southern portion of the Nord-du-Québec admin - tribution of those species, but rather their sporadic istrative region is considered by Quebec bryologists presence over the study area. The new occurrences of to be a “black hole” in terms of knowledge about the Brachythecium erythrorrhizon, Isopterygiopsis muel - distribution of the bryophyte flora, reflecting the pauci - leriana, Polytrichum commune var. perigoniale, Lo - ty of sampling efforts in the region rather than species. phozia ascendens, and L. bicrenata close gaps in their Substantial work is still needed to map bryophyte dis - provincial distribution between southern and northern tributions accurately in the province. Furthermore, in - Quebec but also, in the case of B. erythrorrhizon and creased sampling and the consultation of herbarium P. commune var. perigoniale , between eastern Quebec specimens will be required to fully understand the dis - and ontario. Furthermore, we document the presence of tribution and microhabitat preferences of bryophytes Campylium protensum , Isopterygiopsis pulchella , Poh - throughout North America. Efforts should be made in lia elongata var. elongata, Calypogeia suecica, Mesop - boreal feather moss forests, which may be defined as tychia rutheana, M. heterocolpos var. heterocolpos, “bryo-diversity hotspots” in view of the occurrences Scapania apiculata, and S. uliginosa in the southern of uncommon, infrequent, and rare bryophyte species portion of Nord-du-Québec, 260–670 km west of their and, therefore, require particular conservation attention. previously documented Quebec occurrences. our discoveries of the rare true mosses Campylium Acknowledgements protensum and Hypnum fauriei and the rare liverworts Funding for this study was provided by fellowships Chiloscyphus coadnatus var. rivularis, Odontoschisma from the Fonds de Recherche du Québec –Nature et francisci , and Schistochylopsis laxa , confirm the impor - Tech nologies, the Natural Sciences and Engineering tance of bryophyte sampling efforts in the neglected Research Council of Canada, and the University du regions of Quebec–Labrador. However, some of the Qué bec en Abitibi-Témiscamingue. We thank Philippe spe cies described here may be misunderstood as a result Heine and Flora Joubier for their help in collecting of under-sampling or taxonomic confusion. Recent bryophyte samples. We also sincerely thank the Editor- changes in species taxonomy have made herbarium in-Chief Dwayne Lepitzki, the Associate Editor Jeff specimens unreliable describers of species abundance Saarela, and the reviewers of our paper, including Jen - until they can be re-examined and their identifications nifer Doubt, for the constructive comments that led to confirmed. this improved version of our paper. Finally, considering a larger geographic perspective, even though our findings represent significant distri - Literature Cited butional extensions within the province of Quebec, for Barbé, M., N. J. Fenton, and Y. Bergeron. 2016. So close some species the distance from the adjoining ontario and yet so far away: long-distance dispersal events govern populations is less substantial (50–200 km, on average, bryophyte metacommunity reassembly. Journal of Ecology from our study area). However, bryophyte distributions 104: 1707 –1719. https://doi.org/10.1111/1365-2745.12637 Barbé, M., N. J. Fenton, and Y. Bergeron. 2017. Are post- in ontario are not as well or as recently documented as fire residual forest patches refugia for boreal bryophyte spe - in Quebec (Faubert 2012–2014). Therefore, it is diffi - cies? Implications for ecosystem based management and cult to evaluate the exact distances between species oc - conservation. Biodiversity and Conservation 26: 943 –965 . currences. These species were found in eastern ontario, https://doi.org/10.1007/s10531-016-1281-9 in some localities of the Hudson Bay lowlands, and the Bergeron, Y., S. Gauthier, M. Flannigan, and V. Kafka. Clay Belt region. The Clay Belt straddles Quebec and 2004. Fire regimes at the transition between mixedwood ontario and its specific soil conditions may explain why and coniferous boreal forest in northwestern Quebec. Ecol - the two provinces shared bryophyte assemblages. ogy 85: 1916–1932. https://doi.org/10.1890/02-0716 Bisbee, K. E., S. T. Gower, J. M. Norman, and E. V. Nord - Conclusions and Implications for Management and heim. 2001. Environmental controls on ground cover spe - Conservation cies composition and productivity in a boreal black spruce This study suggests that numerous bryophytes may forest. oecologia 129: 261–270. https://doi.org/ 10.1007/ be more common than expected in Quebec–Labrador. s004420100719 We present a substantially revised and updated list of Caners, R. T, S. E. Macdonald, and R. J. Belland. 2013. the bryoflora of the southern Nord-du-Québec admin - Linking the biological traits of boreal bryophytes to forest habitat change after partial harvesting. 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SUPPLEMENTARY MATERIAL : APPENDIX S1. Number of plots where bryophyte taxa were found in residual forest patches and burned areas in boreal Black Spruce–feather moss forest in the southern portion of the Nord-du-Québec administrative region, western Quebec, Canada.