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An Analysis of the Vascular Flora of Annapolis Heathlands, Nova Scotia

S. CARBYN1, P. M. CATLING2, S. P. VANDER KLOET3, and S. BASQUILL4

1Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, 32 Main Street, Kentville, Nova Scotia B4N 1J5 2 Agriculture and AgriFood Canada, Environmental Health, Biodiversity, Saunders Bldg., Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada; e-mail: [email protected] 3Department of Biology, Acadia University, Wolfville, Nova Scotia B4P 2R6 Canada 4Atlantic Canada Conservation Data Centre, PO Box 6416, Sackville, New Brunswick E4L 1G6 Canada; e-mail: sbasquill@ mta.ca

Carbyn, S., P. M. Catling, S. P. Vander Kloet, and S. Basquill. 2006. An analysis of the vascular flora of Annapolis Heath- lands, Nova Scotia. Canadian Field-Naturalist 120(3): 351–362. A description and analysis of the vascular composition of heathlands in the Annapolis valley were undertaken to pro- vide a basis for biodiversity preservation within a system of protected sites. Species presence and abundance were recorded at 23 remnant sites identified using topographic maps, air photos, and Nova Scotia Department of Natural Resources records. A total of 126 species was recorded, of which 94 were native and 31 introduced. The Annapolis heathland remnants are strongly dominated by with peregrina, Vaccinium angustifolium and Pteridium aquilinum var. latiusculum. A number of species, including Solidago bicolor, Carex tonsa var. rugosperma, Dichanthelium depauperatum, Lechea intermedia, Melampyrum lineare, and Rubus hispidus, were characteristic of heathland remnants, although they usu- ally contributed little to the total cover. The most frequent alien species were Hieracium pilosella and Festuca filiformis,but , present at 7 of 18 sites, appeared to have the greatest impact in displacing native species. Species listed as at risk and sensitive in Nova Scotia, including Helianthemun canadense, ericoides and Viola sagittata var. ovata, occur in open disturbed sand in the Corema heathlands. Distinctive patterns of variation occur in several species and varia- tion in crop relatives is noted with particular reference to the genera Rubus (blackberries), Amelanchier (Juneberries, Saska- toon) and Vaccinium (Blueberries). The available evidence suggests that the heathlands and sandy barrens in the Annapolis valley differ from those further west in Canada and from anthropogenic and coastal heathlands of Nova Scotia in their species composition including particularly the presence of Corema conradii, Hudsonia ericoides and Amelanchier lucida. The need to protect representative examples is supported. Key Words: Heathlands, Annapolis valley, barrens, vascular , flora, Corema, Nova Scotia, Canada.

In 1921, legendary Harvard botanist Merritt Lyndon accords. The only descriptions of Annapolis heathlands Fernald visited the Annapolis valley. He found exten- currently available (Fernald 1921; Roland 1946; Catling sive open heathlands. He noted: “near Berwick and et al. 2004) are brief, non-quantitative, and insufficient from there to Wilmot were vast uncultivated plains as a basis for protection of biodiversity. Here we pro- carpeted, wherever dry enough, with a close growth vide a description of remnants of natural heathland in of the barren Corema conradii, and, the Annapolis Valley along with an indication of dom- … remnants of them near Middleton” (Fernald 1921). inant species, rare and significant species, variation As recently as the 1960s open heathlands with scat- between sites and relationship to other eastern Cana- tered Red (Pinus resinosa) occurred for many dian sand barrens. This is designed to provide a basis miles along the Evangeline Trail (Figure 1). It has been for further study and for the establishment of a system estimated that in pre-settlement times the actual area of protected sites. of heathland encompassed approximately 200 km2. Today less than 3% of the original heathland vegeta- Methods tion remains in the Annapolis Valley (Catling et al. Sites and data 2004), and even that is threatened by loss of natural The study area consists of 23 sites in the Annapolis ecological processes, invasive species and conversion Valley (Table 1, Figure 2). Sites are defined as areas of the landscape (Catling et al. 2004; Catling and surveyed separated by at least 0.5 km. Information Carbyn 2004). Protection of this ecosystem is impor- from the Nova Scotia Department of Natural Resources tant for the protection of (1) insect pollinators of adja- and topographical maps were used to determine the cent crops; (2) protection of wild relatives of crops for most probable locations of heathland vegetation occur- crop improvement; (3) benchmark research examples; ring on sandy soil. (4) teaching examples; (5) nature-related recreational At sites 1–19 abundance of vascular plants was opportunities; and (6) protection of biodiversity gen- recorded. Areas surveyed varied from approximately erally in connection with national and international 0.5–61 hectares in extent. The majority of these 19 sites

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FIGURE 1. Open heathland dominated by Corema conradii (Broom Crowberry) with Pinus resinosa (Red Pine). Photo by P. M. Catling in 2004 at site 8.

were visited on at least two occasions in 2003, with mon, 4 = frequent and locally dominant, 5 = dominant) one visit in early summer and another in autumn. Ap- based on consensus of two or three observers. Although proximately two hours were spent at each site on each some larger sites were visited more often, almost all visit. Species lists were made at each site (Table 2). species recorded were recorded in the first hour of two An abundance value ranging from 0–5 was assigned visits. Time spent at sites was therefore considered to to each species (1 = rare, 2 = uncommon, 3 = com- be adequate and the lists are thought to be essentially

TABLE 1. Site number and location of heathland study sites in the Annapolis Valley, Nova Scotia. Site Number, Name Latitude ° Longitude ° 1 Evangeline Heathland E of 201 45.0059 –64.8882 2 Evangeline Heathland W of 201 45.0048 –64.8930 3 Trail Heathland 45.0005 –64.9004 4 Evangeline Main Heathland 45.0094 –64.8705 5 Brooklyn Street Wildlife Sanctuary 45.0872 –64.5245 6 Caribou Bog Heathland remnant 45.0377 –64.783 7 Trailer Park Heathland 45.0219 –64.8255 8 Exit 17 Heathland, NE corner 44.9992 –64.9411 9 Exit 17 Heathland, NW corner 44.9962 –64.9509 10 4 km W of Middleton 44.9204 –65.1048 11 W side of Middleton 44.9607 –64.9999 12 Evangeline Cemetary 44.9675 –64.9866 13 E side of Greenwood Military Base 44.9896 –64.8955 14 Aldershot Military Base behind shooting range 45.0966 –64.5333 15 Greenwood section 44.9833 –64.8873 16 Exit 17 far east 44.9974 –64.9470 17 Greenwood Military Base main site 44.9801 –64.9381 18 Greenwood Military Base satellite 44.9853 –64.9308 19 201 East near bog 44.9791 –64.9366 20 Plot – W of Whitman Road 45.0109 –64.8404 21 Plot – E of route 201 44.9992 –64.8854 22 Plot – N of route 201 44.9472 –65.0203 23 Plot – S of Ward Road 44.9681 –64.9367 12_05055_flora.qxd 11/1/07 11:11 AM Page 353

2006 CARBYN,CATLING,VANDER KLOET,andBASQUILL:ANNAPOLIS HEALTHLANDS 353

FIGURE 2. Heathland sites surveyed in the Annapolis Valley in 2004 (solid triangles) and location of plots evaluated for cover in 2005 (dots). The sandy soils of the valley (shaded) suggest the maximum extent of heathland.

complete. Voucher specimens collected at various sites Although many voucher specimens were collected were deposited in the herbarium at (and deposited at DAO), a sufficient number of speci- Agriculture Canada in Ottawa (DAO). Status of plant mens of the genus Aronia were not collected to enable species was determined by using Maher et al. (1978), a determination of which species (or hybrid) was pres- the recent listing of species at risk (Nova Scotia De- ent at a particular site. Thus only the genus name ap- partment of Natural Resources 2001*) and informa- pears in Table 2. Although Rosa carolina and Rosa vir- tion and lists on file at the Atlantic Conservation Data giniana were noted, both may have been present at any Centre in Sackville. site where either was recorded, and some intermediates At sites 20–23, plant association sampling methods were noted. Rubus hispidus may be over-represented were employed following provincial plot standards in the survey and Rubus arenicola may be under-rep- (Quigley et al. 2005*) and Canadian vegetation clas- resented due to difficulties in distinguishing these, and sification conventions (CNVC Technical Committee hybrids between them may also have been present. 2004*). In May of 2005, at each of the four sites, a four hundred square meter plot was subjectively placed in a Results and Discussion. homogeneous tract of heathland vegetation, where can- Native species and limitations of the vegetation opy cover exceeded 10 percent. The sampled asso- description ciation was the open woodland stage of the Annapolis Although 104 native species are recorded from the Valley sand plain. Species and cover values, represent- 23 sites surveyed, and although these sites covered a ing percentage of plot area within a vegetation layer, rather extensive area, they were mostly drier examples were visually assessed for five life forms (Table 3). of dry, open sandy habitats (Figure 1). The relatively Lichen names follow Esslinger (1997*), bryophyte short-lived wetter examples and those associated with names follow Anderson et al. (1990), and vascular plant natural disturbances, such as fire, are now much less names are from Kartesz and Meachum (1999). common than the drier examples. Thus as a reconstruc- 12_05055_flora.qxd 11/1/07 11:11 AM Page 354

354 THE CANADIAN FIELD-NATURALIST Vol. 120 1 5 1 1 2 3 4 5 4 3 1 1 4 6 1 1 8 3 3 1 6 4 9 1 9 2 9 2 6 5 3 1 1 1 8 4 1 6 C 1 1 4 9 2 4 5 1 1 5 3 1 4 3 1 2 1 2 T 6 1 3 1 1 9 2 3 2 5 7 4 2 1 1 4 4 1 8 2 2 1 4 9 0 9 9 8 9 8 1 6 6 4 8 4 3 0 S . 1 1 1 1 1 1 1 1 1 1 1 1 1 1 t T n a 9 n 3 – 1 3 1 5 – – 1 1 2 1 – 1 3 – – 3 – 1 1 3 – – – – 1 – – 1 1 – – 2 – – – 1 i 1 m o d 8 – – – 4 – 5 – – 1 3 3 – – – – – – – – 1 1 2 1 – – – – – – 1 1 – – – – 2 – 2 1 = 5 , t 7 n 3 – 2 4 3 5 – 1 1 3 4 – 1 – 2 – 1 1 – 3 1 4 3 – – – – – – 3 1 1 – – – 2 1 3 1 a n i m o 6 – – 1 2 – 5 – – 2 2 3 – – – – – – 2 1 – – – 2 – – – – – – – – 2 – – – – 2 4 d 1 y l l a 5 c – – 4 – – 5 – – 2 2 3 2 2 1 – – 4 2 – – 1 2 2 – – – – – – – 1 – – 2 – – 2 2 1 o l d 4 n 1 – 3 – 1 5 – – – 2 3 1 3 – – 3 2 – – – – – 2 2 – – – – – – – – – – – – 1 – 1 a t n e 3 – – 2 – 1 5 – – 2 2 3 – – – – – 4 1 – – – 2 1 – – – 1 – – 1 – – – 1 – – – – u 1 q e r f 2 2 – 3 – 1 5 – – 1 3 4 – – – 1 – 2 2 – – – 2 1 1 – – – 3 – 1 – – – – – – – – 1 = 4 , 1 – – 2 – 2 5 – – 2 3 4 – – – – – 4 2 – – – 1 1 – – – – – – 1 1 – – 1 – – – – n 1 o m 0 – – 1 1 – 5 – – 1 1 3 2 – – 1 – 4 1 – – – – 1 – – – – 1 1 1 – – – – – – – – 1 m o c = – – 4 – 2 5 – – 1 3 4 1 1 – – – 4 2 1 – – 4 2 – – – 1 1 – 1 1 – – 2 9 – – 1 5 3 , n – 1 3 – 2 5 – 1 2 4 4 – 1 – – – 1 2 1 2 – 3 2 – – – – – – 1 1 – – 1 8 – – 1 2 o m – – 2 – – 5 – – 2 1 2 1 1 – 1 – 4 1 – 2 – 3 2 – 1 1 – – – 1 – – – 2 7 – – 1 – m o c n u – – 1 – – 1 – – 1 1 – 1 – – – – 4 1 – – – – 2 – – – – – – 1 – – – 2 6 – – 1 – = 2 – – 2 – – 5 – – 2 – 4 – 3 – – – – – – – – – 1 – – – – – – – – – – 2 5 – – – – , e r a r – – 1 – 1 5 – – 2 2 2 – 1 – – – 4 2 1 – – 3 1 – – – – 1 – – 1 – 2 2 4 – – – – = 1 . y – – 3 – – 5 – – 2 3 4 2 1 – – – 4 3 – 1 – 4 3 – – – 1 – – 3 – – – 2 3 – – 1 1 e l l a V 1 – 3 – – 5 1 – 2 3 4 – 1 – – – 5 4 – – – 4 3 – – – – – – 1 – – – 3 2 – 1 – 4 s i l o p 1 – 2 – – 5 – – – 2 3 3 – – 1 1 – 4 4 1 1 – 4 1 – – – – 1 – – 1 2 1 – 1 1 – 2 a n n A e h t n n a i s m e k s t c i n s i e r r d F C n d ) l ) a . . l u r r . k r h o e t . c o C r y a a G T e a e D ( ) v M h B h ( o d m m s ) 8 c l a u a e u 1 u l t n n a N t m o a e t a l r a e c x G o e u . r t s e n c ) p a d i . i a u . s G t = l . t a u . c x ( . n o a r a s h g s e o l t h b a g C l a n u n t a i fl s L f v u e c r e o T e n . i r t r . . . c t d M b e x , e . d r l o ( . . p n n s e S L i . h k . r r a i m u B ( r a e S s v . r c d a a v t u o m r ) a i a u m ) o v v . e T r A . . u a m v s C a r e u c . . t m o ) L e d i u a M L e l a . ) n . ( n n n a i . l d n i u T ( n . a l p t . v r r h ) a . g c L B k k n o n r t t . ) y i r a s a f L ( o a e r e i l y . d p c c u . a h o i s ) a f r a ( * i e l r a i i l e h n p s . e t t i n . * r i r a u a L i A l c c . s c m r m u H F a a i B B u L . s u i o × L k W a . . * = l o h e e * i e a ( - m a a M n t t b l t M o e ) ) . i l s m T L h a a . i s s i n s e . . a a l w u p a i e ( n b i i u S . i y c B r a u u r r l d a L R s a n i e n n v L e c t a i ( e s d b r a i v v x i p i r i s o e a g S r r a T i l o i ) u l r d a l S a e e c e e a s n h e e . D e V . c r e r e e . g a m o d w e i r m H c o c e r a u a a fl c t s f s e r a f d i s f i m d L F F L i u l l l i c i a o a g u m m m e i m i i i . d e a o ( ( ( a r n r e a i p t r a l l e r f m n r l s l u h a d u u u r r r p a a i e l s c x s m a a i n i i i r m n c u y y p o i m o a a n u t i a e e e n s l l l n e t N l u o u a i i i p p a r a u s s r o . p p h e o n a S s i i e e e p a i s r a c fl m s m t r i d i m e h h h o c g n n i p o a p c r i s a : e u d h h h c n e g e i l m u c c c b c o n g u l m s t t t o o s r t p p a a a h e u fi a 2 g l a r o a d n t t e c s i a t s n n n u o n i n a n n n r t t u e t n e p h e s a i t s r a a h l u a a a y E a h i t h m i i s e m l x x x x x x x a a a n p a l l n v a l l l l i t n o l p t c r L s I n c t i e e e e e e e e n r l p o n a r h h h t e u u e e e e n e m l r r r r r r r r r s a t n r o p B o t t e a h e i a c c c e a c o = a o i i i i i e r e o a y a a a a a a a o o r r e e n c g n m m m c p r r r c A T * S A A A A A A A A A A A A B B C C C C C C C C C C C C C C C D D D D D D D D D 12_05055_flora.qxd 11/1/07 11:11 AM Page 355

2006 CARBYN,CATLING,VANDER KLOET,andBASQUILL:ANNAPOLIS HEALTHLANDS 355 . 5 9 1 1 1 1 1 6 1 1 5 3 2 4 1 7 2 5 5 0 0 1 2 2 5 9 3 6 8 5 2 1 0 5 0 1 0 6 4 t C 1 1 1 5 1 1 3 2 1 2 1 3 5 4 3 3 5 1 1 n T a n i m 6 4 8 1 1 1 1 1 4 1 1 3 3 8 7 2 3 1 6 2 2 1 2 1 0 7 0 4 4 3 3 7 0 8 9 6 6 2 5 S o 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 T d = 9 – 1 – 1 – 2 4 – – – 2 1 – – 1 – 3 – – 2 1 1 2 – 1 – – – – – 1 2 2 3 – – 2 – – 1 5 , t n a 8 3 – – – – – 3 – – – 1 2 – 3 – – – – 2 – 1 – 2 – – 2 – – – – 1 3 2 2 – – – – – n 1 i m o d 7 3 1 2 2 1 – 4 1 – 1 2 1 – 1 – – – – 2 1 2 – 2 – 1 2 – – – 1 1 3 2 2 – – – – – 1 y l l a c o 6 l – – – – – – – – – – – 2 – – – – – 2 – 1 – – – – – – 1 – – – 2 1 – 3 – 2 3 – – 1 d n a t 5 n 1 2 – – – – 3 – – – 1 – – – – – – 2 – 1 1 2 2 – – 3 – 2 – – 3 3 3 1 – 4 – – – 1 e u q 4 e – 1 – 1 1 – 3 – – – 1 – 1 – – 2 1 – – 1 1 – 2 – – 3 – – – 2 3 – 3 – – 4 – – – r 1 f = 4 3 – 1 – – 1 – 2 – – – 1 – – – – – 4 3 – 1 2 1 2 – – – – 1 – – 4 1 1 2 – 4 – – – , 1 n o 2 m – – – – 1 – 3 – – – 1 – – – – – 1 – – – 3 – 1 – 1 – – – – – 4 4 – – – – – – – 1 m o c 1 – – – 1 1 – 3 – – – 1 – – – – – 2 2 – – 1 1 2 – – – – – – – 3 4 1 – – 3 – – – 1 = 3 , 0 – 1 – – 1 – – – – – – 1 – – – – 3 4 – – 1 1 1 – 1 2 – – – – 1 – 4 – – – – – – n 1 o m – 1 1 – – – 4 – – – 2 – – – – 1 2 1 – 1 3 1 2 1 – – – – – – 4 3 1 2 9 – 4 – – – m o c n 1 – 1 – – – 4 – – – 2 – – – – 1 – 3 1 – 2 – 1 1 – 1 – – 1 – 5 2 – 2 8 – 3 – – – u = 2 – – – – 1 – 3 – – – 1 – – – – 1 4 2 – 2 1 1 2 1 – – 1 – – – 2 2 2 2 7 – 3 3 – – , e r a r 3 – – – 1 1 – 2 – – – – – – – 1 – 3 – – 1 2 2 – 2 – – – – – 2 – 3 – 6 – 2 – – 1 ` = 1 . – – – – – – 4 – – – – – – – – – 1 – – 3 1 1 1 – – 1 – – – – 1 5 1 – 5 – 4 – – – y e l l a V – 2 1 1 1 – 1 – – – – – – – – – 3 1 – 1 – – 1 – 3 – – – – 1 3 2 1 4 4 1 5 – – – s i l o – 2 – – – – 3 – – – – – – – – 3 1 1 – 1 1 1 2 – – – – 1 – 1 3 3 1 – 3 – 1 – – – p a n n 1 1 – 1 1 – 4 – – – 2 – 1 – – 1 3 1 – 2 2 – 2 – 2 – – – – 1 4 4 2 4 2 – 4 – – – A e h t 1 2 – 1 – – 2 – 1 – 1 – – – – 1 3 1 – 5 – 1 1 – 2 – – – – 1 3 3 1 4 1 1 4 – 1 1 n i s e t i s d n a l h t a e h 9 1 . t w a r s a . r u F t e a ) v t . o s x c h d h l c n s i a r a t * u o M e t ( P . c a h x n = n a a c i i a h s j l o c * s d C a o i . ? r e K f n T x i r . t u , . M K h h g r p . b f s * h K c * c ) e ) n P o e s a e . i s l . . . o . t d o ) e n j r s . e l i s l m L S l s . e l g s m u . M K s ( e k * D r i . t u * e e * . * i n a . o r t L . P c l a . B s r e . h c t d . . V a e a e L D i m a a V L v l s ) l s c e L e ) K s * t . * L v g L o . s u . m ) e ) s r . u h n H * p W u e n . l . o z . ) . * . r g e e b . s u ( l ( t h L t n s e r o u i e L m . D S i e . i L L u d d L D L y y l L c e e a d o L l d i u i ( m l a a a G ) s = o e L h h L t M n b i * l g t a i a a s a R a a a a o o M t s c . R r d . l s l c ) t i e P i l l a i ( i i t . r a n . n s n i . S e n n a . m l e l l L u y s a i g u r a a M o i g p s c n . d a s u u L P r t d l a a L e e u u o e L a T h n v m o i . . c i l i d n D ( o f s n a c a A s n ( s s a r c f L c e n e i n . p i s ( M i m s u P e e e a o v n i e L p o n i r t i o o m o t ( S o c r s y d a l i t d m a s l n k l l c y s d b i m r e r m d t g a r m a m fl u a s i i m s i y e n h a n t t g n u e u i i s e i l a a h u a p o u p r o n r s r u a r s u p p c c o e s t r i s a k o n a f m p n i c i g l n t m m e i m c l e i i e e n b k N u a n p c u a m e a e t a u c l v m s u n m r c e m i f n u a u u ( v a v e r i i r e m m o n s d e r c p f b i i n e a l fi t c s a o c u a a h r u a r r r . p i i a n y u u a l u o h s e m o t y t u d d h o e s a i d e fi c i i p a t i a 2 r r t n r a a o p s b s s r g r m a i i a s n l r l o o t g e a a s t c c o c h i a n t r e i c c n s o r n u E t t l u a a c e a p p n u a s s s s u r o a a u a a a e x l s a n p l e i u u l a u I a s L n t c m i u i r r h g o o c i g u u u u t t a e l i l e e n m e g l r y u d l l p i y B c z r o u i i c e e s s c c a n n c n n n n c c y n e o a a o y = e a u y i i a a p i i i i i l h i i r e a e c r u i e a e e A y y u u T * S E E E F F F G G H H H H H J J K L L L L L L L L L M M M M M M P P P P P P P P 12_05055_flora.qxd 11/1/07 11:11 AM Page 356

356 THE CANADIAN FIELD-NATURALIST Vol. 120 . 2 6 3 6 8 4 1 1 1 2 2 1 2 2 7 2 4 2 6 4 1 2 1 5 3 2 4 1 7 9 2 5 8 6 1 4 6 9 7 t C 5 1 3 1 1 1 3 3 4 4 1 5 n T a n i m 2 6 1 6 5 3 1 1 1 2 7 2 1 2 2 5 2 4 2 9 4 4 1 2 1 3 3 2 3 8 0 7 0 6 7 7 0 4 8 S o 1 1 1 1 1 1 1 2 1 1 T d = 9 1 1 – 1 – – – – 2 – – – – 2 – – – – – 1 – – – 2 1 – – – – 2 1 3 – 2 – – 1 3 – 1 5 , t n a 8 – 1 – 1 – – 2 – 1 – – – – 1 1 – – 1 – – – 1 – – 1 – 1 – – 1 4 3 – 1 – – – 2 – n 1 i m o d 7 1 1 1 1 – – 2 – 1 – 1 1 – 2 – – – 1 – 1 1 1 1 – 2 1 1 1 1 2 3 3 – 2 – – 1 2 2 1 y l l a c o 6 l – – – 3 – 3 – – 1 – – – – – 2 – – – 1 2 – 1 – – – – – – – 1 1 2 1 2 – – – 1 – 1 d n a t 5 n – – – 3 – 2 – – 3 – – – – – – – – – – – – – – 2 3 – – – – 2 3 3 – 2 – – – 3 – 1 e u q 4 e – – – 4 – 1 – – 4 – – – – – 1 – – – – – – – – – – – – – – – 4 3 – 2 – – – 5 – r 1 f = 4 3 – – – 3 – 1 – – 3 – – – – – – – – – – – – – – 3 1 – – – – 2 3 2 – 1 – – – 2 1 , 1 n o 2 m – 1 – 2 – 1 – – 2 – – – – – 1 – 1 – – – – – – – 2 3 1 – – 2 3 2 – 1 – – – 4 – 1 m o c 1 – – – 2 – 1 – – – – – – – – 1 – – – – – – – – – 3 – – – – 2 3 2 – 1 – – – 4 – 1 = 3 , 0 – – 2 – 1 – 1 2 – – – – – – – – – – – – – – – 1 1 – – – – 4 3 2 – – – – – 3 1 n 1 o m – – – 4 1 2 – – 3 – – – – 2 2 – – – – 1 – – – – 3 – – – – 2 2 2 – 1 – – – 9 1 – m o c n – – – 4 – 1 1 – 2 – – – – 1 3 – – – – 2 – – 1 – 3 – 1 – – 1 2 3 – 1 – – – 8 – – u = 2 – – – 3 – – – – 3 – – – – – – – – – – – 1 – – 2 2 – – – – 2 1 2 – – – – – 7 4 – , e r a r – – – 3 1 – – – 3 – – – – – – – – – – – – – – 3 – – – – – – – 2 – – – – – 6 4 – = 1 . – – – 3 – – – – 3 – – – – 1 – – – – – – – – – – 5 – – – – 3 – 2 – – – – – 5 3 – y e l l a V – 1 – 2 1 – – – 2 – – – – – – 1 – – – – – – – 2 2 – – – – 1 2 3 – 1 – 1 – 4 4 – s i l o – – – 3 – – – – 2 1 – – – – 1 – – – 1 – – – – – 2 – – – – 1 2 1 – – 2 – – 3 5 – p a n n – – – 3 1 4 – 1 2 – – – 1 3 2 – – – – – – – – 1 3 1 – – 1 2 4 4 – 1 2 1 – 2 4 – A e h t – 1 – 3 1 1 2 1 2 – – – 1 – – – 1 1 – – – 1 – 2 1 2 – – – 1 3 2 1 1 – 1 – 1 – 3 n i s e t i s . d n w r a l e h d t n a e U h ) . 9 v 1 s t e a D s ( . r u t e m e a v t u v o s l ö c u d L l c n s a . a t u D i o e t t c a & l n = . . a . r r d C r A a . n o T v ) b . u , T s i b n s . L ) l a ( e . h x a t u i s . . h L * S r s m ( e c x h K e . i i ) u l . t c . h . e c ) f r L a o . c s n x . . e . t L M . i . o t fl h . a k n ( t . h p L o t * i l L i . m f c P L t a e c . n ( L r f S M u . . t i a i a u s n B . * * d l h A s e i . a i . a s r ) a l = * i L * i s t r l a t L N a m R i t . t i L s s h * l i c M s * s d R e . o . L a h o s s i a i s n u n a u n . S l f s L . a e a i . n l d r B a . A x n e i L i L t n M l E e R a n e e a i n a . u e n t L l T a a L i r . e L n e l r n c a a i u i L d u n d l a o a l m e a r l n a n a . i a a a L i o i a * i l l r d e t v L s i n n a h t . a a e l . e p i u o e M e n l n t d n D n a i t d e c u o u a n r m e u m i s p s p a i r t c m n s o n r h y g e i c n L o a n t s c b a m i q a s n m o s i h d a c h a a o r i n o s n c a a r c i i m l c i b e h o i s t a i o e r i u e i r l r a a g e e r g b a c r s n b n e u d l N n u m u c i i r r r p e r n s r r l i j g b p c n e m r a p g t ( e r s t r i e n o e l s d c g t p s e e i c a a a a a c g m u m l i r n r . e i t p v s u e o l l l l o o o o o o o k r s a a a h a l l l l s l s fi u n u a h v a m a i a 2 r i y a t d n a c g g g g g g i i i i s s s i i u n t a u u r r x t t t t o h v c o t i h e n t e o a s s s s a a a a a a l l c E e u u u d m n t e p c a n n n n g n i p r a a n b d h u u u u I d d d d d d x L u u n l a a n n n r u r e e e e y i i i i i i i e r r l e e d b b b b o p s s m B e l t t t p t a p a l l l l l l l i i i a u u u d m i e l e = u u u u u u a h o o t u r r r c A o o o o o o o o o r o o o o o o i a t e y p p p T * S P P P P P P P P P P P P P Q R R R R R R R R R R S S S S S S S S S S S S S S T 12_05055_flora.qxd 11/1/07 11:11 AM Page 357

2006 CARBYN,CATLING,VANDER KLOET,andBASQUILL:ANNAPOLIS HEALTHLANDS 357 . 3 5 3 1 8 2 2 4 3 6 t C 1 5 tion of the Annapolis sand barren flora, the mesic and n T a n

i successional stages of dry barrens are to a large ex- m 3 4 2 1 8 1 2 2 1 8 S

o tent omitted. This may explain the lack of some species 1 1 T d in the survey, such as Agalinis neoscotica (Greene) = 9 1 – – 3 – – – – – – 1 5

, . (Middleton False Foxglove), which was des- t n

a cribed from Annapolis heathlands near Middleton, 8 – – – 2 – – 1 – – – n 1 i and also Bartonia virginica (L.) BSP., Carex atlantica, m o

d Polygala sanguinea L. and Sisyrinchium fuscatum 7 – – – 2 2 – 1 – – 2 1 y l

l Bickn. All of these species occur on heathlands near a c Middleton (personal observation), but were not seen o 6 l – – 1 1 – – – – – – 1

d during the present survey. Their abundance in the area n a

t in the past (Fernald 1921, page 138) suggests that the 5 n 1 – – 4 – – 1 2 – – 1 e more mesic and naturally disturbed areas were much u q 4 e – – 1 3 – – – – – – more prevalent only a short time ago. Fernald (1921) r 1 f

= referred to these as the “the damper Polytrichim-car- 4 3 1 – – 3 – – – – – –

, Bartonia virginica 1 peted areas.” He noted that “ was n

o everywhere,” but it was not recorded in any of the bar- 2 m – 1 – 3 – – – – – – 1

m rens in our survey and only our site 10 approached this o c 1 damper sand barrens habitat. 1 – – 4 – – – – – – 1 =

3 In addition to the lack of mesic sites it is of interest , 0 1 – – 3 – – – – – 1 n 1 that in two days in 1920 Fernald (1921) recorded two o

m species from the drier heathlands near Middleton that 1 – – 4 – – 1 – – – 9 m

o were not seen by us. These were Potentilla tridentata c n 1 1 1 – 1 1 1 – – 1 8 u and Pyrola rotundifolia. Although our data indicate the = general and dominant composition of the Annapolis 2 – – – 3 – – – – – – 7 ,

e heathlands, the differences with Fernald’s brief survey r a r 3 – 2 5 – – – – 1 – 6 suggest that it may never be possible to have a com- =

1 plete knowledge of their former floristc composition. . 2 – – 3 – – – – – – 5 y e

l Dominant vascular plants l a V – – – 4 – – 1 – – – 4 Relatively few of the 126 species (94 were native s i l and 31 introduced – Table 2) present in the 19 com- o 1 – – 4 – – – – 1 – 3 p

a pletely surveyed sites were dominant in the vegetation. n n Species present at all sites included Corema conradii, 1 – – 4 – – 1 – – – 2 A

e Danthonia spicata, Pinus strobus and Solidago bicolor. h t – 1 – 4 – – 1 – – 1 – n Species present at 18 of the 19 sites were Amelanchier i s

e laevis, Carex tonsa var. rugosperma, Comptonia peregi- t i s na, Dichanthelium depauperatum, Pteridium aquilinum d n

a var. latiusculum, Vaccinium angustifolium and Populus l h

t tremuloides. Species with the highest overall cover a e

h values included Corema conradii with 91 followed by 9 y 1 Comptonia peregrina with 59, Pteridium acquilinum a t r a

G var. latiusculum with 57, and Vaccinium angustifolium s . r u t & e with 56. A number of species, including Solidago bi- a . v t r . o s r

c color, Carex tonsa var. rugosperma, Dichanthelium o G d l T n a a t ) & depauperatum, Lechea intermedia, Melampyrum lin- . o . e m t L T c u ( eare and Rubus hispidus, were characteristic of heath- s n = ) s . a o t e t d C b land remnants, although they were rarely dominant. d u n i T m u o , * N y The most frequent alien species were Hieracium pilo- b . s ( n r h i a e x c . o t a s t h i s t c n sella and Festuca filiformis, both at 17 and 16 (respec- i s s c e e a a i i * × A l v o 4 6 1 c c . a tively) of 19 sites. The most serious invasive of the o 9 2 3 M . e t m m r M L . 1 p o s * r a u u t ) . * s i i

S heathlands in terms of displacing native species was e a . v l l . L n v d ) = . L o o i e . f f ( d L s Pinus sylvestris, present at 8 sites (Catling and Car- L t i i o i S b e i t t l l e a l u s s T m s i a m d * A e t n u u

. byn 2004). m . u n n e i u r r i g g t d a c e L c m d y t e c e n n n v o a u

u The more wooded plot sites (20–23) gave a very sim- a p fi c a r r e o a a m t f d s n c N u t c v a p o i o i a ( c d t m m m ilar picture of the Corema-dominated heathland, but r c r g m . t a a t o a m m u u u fi a r 2 u r i i i n c e u u i t s i t c N I i i t

n Pinus resinosa n n n with scattered , mostly (Table 3, E l l n i i i r n l l l n a I L a a o o c c c l a a a u o e i i f f B t t t c c c i

r Figure 3). Three species more often associated with i i b c c o = c i i i i A o o o r r a a a e T T T * S T T V V V V V V V V T woodland than with barrens (Clintonia borealis, Lyco- 12_05055_flora.qxd 11/1/07 11:11 AM Page 358

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TABLE 3. Cover values for vegetation layers in single 400 square meter plots at each of 4 sites in the Annapolis valley, based on data collected by Sean Basquill in May 2005. * = introduced, + = not listed in Table 2. Species Site 20 Site 21 Site 22 Site 23 Trees Amelanchier laevis Wieg. 2 – – – Betula populifolia Marsh 3 2 5 5 Picea mariana (P. Mill.) B.S.P. – – 5 – Pinus resinosa Ait. 20 25 20 10 Pinus strobus L. – – 5 8 Pinus sylvestris L. *–––10 Prunus pensylvanica L. f. – 1 – – Quercus rubra L. – 4 0.1 – and small trees Amelanchier laevis Wieg. – – 0.1 0.2 Betula populifolia Marsh 3 2 5.1 3 Comptonia peregrina (L.) Coult. 1 1 – – Juniperus communis L. var. depressa Pursh –––0.1 Picea mariana (P. Mill.) B.S.P. – 0.1 – – Pinus resinosa Ait. 4 4 1 1 Pinus strobus L. 0.5 – 1 5 Pinus sylvestris L. * – – – 5 Quercus rubra L. – 1 – – Vaccinium angustifolium Ait. 0.1 0.5 3 6 Herbs and dwarf shrubs Arctostaphylos uva-ursi (L.) Spreng. 30 5 10 8 Betula populifolia Marsh – – 0.2 – Carex tonsa (Fern.) Bickn. (sensu lato) 0.01 0.1 1 1 Clintonia borealis (Ait.) Raf. + – – 0.01 – Corema conradii (Torr.) Torr. ex Loud. 58 40 60 60 Deschampsia flexuosa (L.) Trin. – – 1 1 Dichanthelium depauperatum (Muhl.) Gould – – 0.1 – Epigaea repens L. – – 0.01 0.1 Festuca filiformis Pourret * 0.1 – 0.01 0.5 Festuca trachyphylla (Hackel) Krajina * – 0.001 – – Gaultheria procumbens L. 0.5 0.01 – – Hieracium piloselloides Vill. * – – 0.01 – Hudsonia ericoides L. 0.01 0.1 – 0.3 Kalmia angustifolia L. 0.1 0.01 – – Lechea intermedia Leggett 0.001 – – – Lycopodium obscurum L. + – – 0.1 – Lycopodium tristachyum Pursh 0.1 0.01 – – Maianthemum canadense Desf. ––0.01 – Oryzopsis asperifolia Michx. + – 0.1 0.2 – Poa pratensis L. – 0.1 0.1 0.1 Potentilla simplex Michx. –––0.2 Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underw. 10 1 0.02 – Rubus hispidus L. – 0.001 – 0.01 Solidago bicolor L. – 0.2 –– Solidago nemoralis Ait. – 0.001 0.2 0.1 Taraxacum officinale G. H. Weber ex Wiggers ––0.002 – Trientalis borealis Raf. 0.01 – 0.01 – Vaccinium angustifolium Ait. –––0.01 Mosses Ceratodon purpureus (Hedw.) Brid.+ – 0.001 – 2 scoparium Hedw.+ – 1 –– Pleurozium schreberi (Brid.) Mitt.+ 0.5 0.1 0.5 – Polytrichum commune Hedw.+ ––0.1 1 Polytrichum juniperinum Hedw.+ – 0.01 0.3 3 Lichens Cladina rangiferina (L.) Nyl.+ 18 40 7 10 Cladina stellaris (Opiz) Brodo + 2 10 3 5 12_05055_flora.qxd 11/1/07 11:11 AM Page 359

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FIGURE 3. Open wooded heathland dominated by Corema conradii and Cladina rangiferina with scattered Pinus resinosa. Although there is much open area, the site exceeds the threshold 10% tree cover required for classification as a treed association under Canadian vegetation classification conventions (CNVC Technical Committee 2004*). Photo by S. Basquill on 25 May 2005 at site 20.

podium obscurum and Oryzopsis asperifolia) were cleared for agriculture as suggested by Roland. How- present a these more wooded sites. Although vascular ever there is little doubt that they also existed in pre- plants were the focus of the descriptive work, the plot settlement times. data included lichens and bryophytes, suggesting the Rare and significant species former (Cladina rangiferina and C. stellaris) to be a The only species considered to be at risk in Nova significant component. This suggestion is supported by Scotia that occurs in the Annapolis heathlands is general observations at the 19 other sites where quan- Helianthemum canadense (Long-Branch Frostweed, titative data on bryophytes and lichens was not ob- Figure 4). It was found at 6 of the sites, always in dis- tained. turbed habitats with some bare sand. Both Viola sagit- Although Roland (1946) described the Annapolis tata var. ovata and Hudsonia ericoides are listed as sand barrens and heathlands 60 years ago, his descrip- sensitive (Nova Scotia Department of Natural Re- tion, which recorded 35 species, corresponds very sources 2001*), and these also occurred in disturbed closely to the composition seen in the relicts that re- sandy areas (Figure 5). The provincially rare (Maher main today. The successional processes are possibly et al. 1978) Sisyrinchium fuscatum Bickn. (Coastal- also the same as described generally by Roland, but Plain Blue-Eyed-Grass) was reported from “sandy areas succession may have been much more rapid in some near Middleton” (sub Sisyrinchium arenicola Bickn., areas than in others where barrens and heathlands may Roland and Smith 1969), but was not seen during the have existed as a subclimax lasting for centuries, even present survey. without fire. Roland’s article was written at a time when In addition to the rare and threatened species there sand barrens, heathlands and savanna were regarded is a suggestion of some distinctive patterns of varia- as wasteland rather than as special places for native tion due to taxonomic recognition of several variants. biodiversity, but his strong theme is accompa- Among these is a sand barren ecotype of Amelanchier nied by many astute observations regarding the native lucida (personal observation). Taxa of blackberries flora. At the time of early settlement the barrens and (Rubus particeps, R. arenicola), although not recog- heathlands may have expanded due to cutting and burn- nized as discrete in some recent literature, have been ing of pine forests and abandonment of sandy land reported from the Annapolis heathlands suggesting at 12_05055_flora.qxd 11/1/07 11:11 AM Page 360

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least the presence of distinctive genetic variants with- in broadly defined species. Further study may indi- cate both of these to be worthy of taxonomic recog- nition. A restricted glabrous variant of Viola sagittata var. ovata (f. glabrata) occurs on the dry open sand with the more typical form. Three taxa have been described from Annapolis heathlands: Amelanchier lucida Fer- nald, Dichanthelium (sub Panicum) depauperatum var. psilophyllum f. cryptostachys Fernald, and Rubus par- ticeps Bailey. The heathland ecosystem is particularly valuable as a reservoir of genetic variation in crops and crop rel- atives. Fernald (1921) commented on the remarkable variation in wild blueberries (Vaccinium angustifolium and V. myrtilloides) in Annapolis heathland remnants near Middleton. In addition to blueberries (Table 2) there are potential genotypes of Aronia (Aronia sp.), of blackberries (Rubus hispidus and others), juneber- ries (Amelanchier lucida and A. laevis), huckleberries (Gaylussacia baccata and G. dumosa), cherries (Pru- nus pensylvanica, P. virginiana, and P. serotina) and a strawberry (Fragaria virginiana). FIGURE 4. Helianthemum canadense (Long-Branch Frost- Variation between sites weed), a native plant at risk in Nova Scotia, with its Although all sites shared dominant species (Table main Nova Scotian occurrence on the Annapolis valley 2) they varied in diversity from 26 to 57 native species. heathlands. Photo by P. M. Catling in 2004 at site 8. Much of the variation between sites appeared to be attributable to disturbance and soil moisture. Those St. Lawrence region south to New Jersey. It does not sites with lower lying and periodic moist areas and occur in sand barrens further to the west in Canada, for open sand had the highest vascular plant diversity. example in the Ottawa valley, where Vaccinium angus- Site 10 for example was the most unlike the other tifolium is the dominant heath accompanied by sites in native species composition, possibly a result other shrubs such as Prunus susquehanae and Comp- of parts of it having a relatively high table and tonia peregina (Carbyn and Catling 1995). Other spe- disturbed areas of periodically moist sand where dry cies present in the Annapolis heathlands but absent in ground species such as Carex tonsa var. rugosperma Ottawa valley sites were Amelanchier lucida, Des- and Danthonia spicata occurred with wetland species champsia flexuosa, Rubus hispidus and Solidago bi- color such as Drosera intermedia. . Among the prevalent species in Ottawa valley Characteristic native species and “at risk” or “sen- sand barrens but absent in the Annapolis sites were Carex siccata, Carex lucorum, Dichanthelium sabu- sitive” species were present at some sites but absent lorum var. thinium, Polygonella articulata and Prunus from others (Table 2). Sites also varied in the extent to susquehanae (Carbyn and Catling 1995). which alien species were present and dominant (Table Within Nova Scotia, “barrens” with heath vegeta- 2). For example parts of sites 1 and 4 had extensive tion occupied a large part of the western portion of the and spreading stands of Pinus sylvestris, but displace- province (Strang 1972). Some of these barrens are a ment of native vegetation by this introduced tree was short-lived successional stage following fire, whereas . either less extensive or not observed at other sites in other cases they are long persisting. Those of short Festuca filiformis was a co-dominant at site 5 but not duration have in some cases been produced by cut- elsewhere. ting and then maintained by fires. Two such barrens Species presence, diversity and extent of impacts were described by Hall and Aalders (1968). Such bar- all require consideration in selecting sites for protec- rens, produced by human activities, have been consid- tion. The variation between sites in composition and ered a degraded landscape of little economic value. The abundance, including that of rare and/or characteristic existence and importance of apparently natural bar- species, suggests that protection of a number of sites rens have only recently become apparent (Catling et al. will be necessary to protect representative ecosystem 2004). The sites described by Hall and Aalders (1968) components. differ from those in the Annapolis Valley in lacking Relationship to other heathlands and barrens Corema conradii, the dominant of the Annapolis bar- In Canada, Corema-dominated barrens are charac- rens. They were also without several other species, teristic of the maritime region. Corema conradii has including Hudsonia ericoides. On this basis the dif- a restricted distribution extending from the Gulf of ferences between man-made and natural barrens seem 12_05055_flora.qxd 11/1/07 11:11 AM Page 361

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FIGURE 5. Open sandy ground in a disturbed area of heathland with Comptonia peregrina (Sweet Fern) and Viola sagittata var. ovata (Arrow-leaved Violet). Photo by P. M. Catling in 2004 at site 8.

pronounced, but with only two anthropogenic sites, a with current information on status ranks. Lawrence more detailed comparison is unnecessary. Benjamin and Randy Milton, also of the Nova Scotia The apparently longer persisting natural barrens in Department of Natural Resources, provided informa- Nova Scotia are readily divided into two major vegeta- tion on site locations for rare plant species in the An- tion types based on either granitic or sandy substrates napolis Valley. Steve Javorek and M. Grant assisted (personal observation). Sandy heathlands like those with field studies (site 1). Geoff Mercer assisted with of the Annapolis Valley were also well developed in the the inventory of sites on military bases at Aldershot interior of Yarmouth County and near Debert, Spring- (site 14) and Greenwood (sites 17 and 18). hill, Parrsboro, and on coastal dunes in Guysborough and Kings counties and on Sable Island. The coastal Documents Cited (marked * in text) heathlands on sand differ from the interior sites in both CNVC Technical Committee. 2004. Plant association con- species composition and species abundance, and the cept for the Canadian National Vegetation Classification. differences suggest adaptation to different conditions. NatureServe Canada. 5 pages. http://www.cnvc-cnvc.ca. Esslinger, T. L. Differences also exist at the infraspecific level. For 1997. A cumulative checklist for the lichen- forming, lichenicolous and allied fungi of the continental example Lechea intermedia var. intermedia occurs in United States and Canada. North Dakota State University: interior heathlands while L. intermedia var. juniperina http://www.ndsu.nodak.edu/instruct/esslinge/chcklst/ occurs on the coastal sandy heathlands. The heathlands chcklst7.htm on Sable Island appear distinct (Catling et al. 1985). Nova Scotia Department of Natural Resources. 2001. The interior sandy heathlands also appear to vary re- General status ranks of wild species. Vascular plants. http:// gionally, presumably due to differences in climate. The www.gov.ns.ca/natr/wildlife/genstatus/specieslist.asp climate in southern Yarmouth and Shelburne differs Quigley, E., P. Neily, K. Keys, and B. Stewart. 2005. Nova from that of the Annapolis Valley (personal observa- Scotia Department of Natural Resources Forest Ecosys- tion). The data suggest that the Annapolis heathlands tem Classification; Methods for Plot Establishment and Vegetation Inventory. Unpublished Internal Document. are distinctive in their floristic composition. Literature Cited Acknowledgments Anderson, L. E., H. A. Crum, and W. R. Buck. 1990. List Mark F. Elderkin, Species at Risk biologist with the of the mosses of north of Mexico. Bryol- Nova Scotia Department of Natural Resources, assisted ogist 93: 448-499. 12_05055_flora.qxd 11/1/07 11:11 AM Page 362

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Carbyn, S. E., and P. M. Catling. 1995. Vascular flora of Kartesz, J. T., and C. A. Meachum. 1999. Synthesis of the sand barrens in the middle Ottawa valley. Canadian Field- North American Flora, version 1.0. North Carolina Botan- Naturalist 109: 242-250. ical Garden, Chapel Hill, North Carolina. Catling, P. M., and S. Carbyn. 2004. Invasive Scots Pine Maher, R. V., D. J. White, G. W. Argus, and P. Keddy. (Pinus sylvestris) replacing Corema heathland in the Anna- 1978. The rare vascular plants of Nova Scotia. Syllogeus polis valley. Canadian Field-Naturalist 119(2): 237-244. (National Museums of Canada, Ottawa) 18. 37 pages. Catling, P. M., S. Carbyn, S. P. Vander Kloet, K. MacKen- Roland. A. 1946. The vegetation of the Annapolis valley I. zie, S. Javorek, and M. Grant. 2004. Saving Annapolis – well drained sand areas. Acadian Naturalist 2(7): 1-20. Heathlands. Canadian Botanical Association Bulletin 37(1): Roland, A. E., and E. C. Smith. 1969. The flora of Nova 12-14. Scotia. Nova Scotia Museum. Reprinted from Proceedings Catling, P. M., B. Freedman, and Z. Lucas. 1985. Vegeta- of the Nova Scotian Institute of Science 26: 1-238, 277- tion and phytogeography of Sable Island, Nova Scotia. 743. Proceedings of the Nova Scotia Institute of Science 24: Strang, R. M. 1972. Ecology and land use of the barrens of 181-248. western Nova Scotia. Canadian Journal of Forest Research Fernald, M. L. 1921. The Gray Herbarium expedition to 2: 276-290. Nova Scotia. Rhodora 23: 130-171. Hall, I. V., and L. E. Aalders. 1968. The botanical compo- Received 25 September 2005 sition of two barrens in Nova Scotia. Le Naturaliste cana- Accepted 17 February 2007 dien 95: 393-396.