growth curves for calcium-tolerant lichens in the canadian ... · growth curves for...

Arctic and Alpine Research, Vol. 27, No. 3, 1995, pp. 290-297

Growth Curves for Calcium-tolerant Lichens in the Canadian Rocky Mountains

Daniel P. McCarthy Department of Geography, Brock University, St. Catharines, Ontario, L2S 3A1, Canada.

Daniel J. Smith

Department of Geography, University of Victoria, Box 3050, Victoria, British Columbia, V8W 3P5, Canada.

Introduction

The basic tenet of lichenometry is that thallus diameter can be used to provide a measure of the relative or absolute age of stable surfaces (Locke et al., 1979). While a wide variety of taxa have been used in dating studies using this principle, only a few

species provide reproducible and presumably accurate dates (In- nes, 1985). Prominent amongst this group are members of the

Rhizocarpon genus which are long lived and have a global distribution (e.g., Benedict, 1967; Denton and Karlen, 1973).

Proponents of lichenometry suggest that it is the best available technique for dating treeless Little Ice Age deposits (e.g., Innes, 1985). For instance, in the siliceous Main Ranges of the Canadian Rocky Mountains a number of researchers have suc-

cessfully used Rhizocarpon geographicum agg. to provide minimum dates for a variety of Little Ice Age moraine complexes (e.g., Luckman, 1977; McCarthy, 1985). Lichenometry has seen

comparatively little use in the calcareous front ranges of the Canadian Rockies, where the development of R. geographicum agg. thalli are inhibited (Osbom and Taylor, 1975). Attention in these areas has instead been directed to the calcium-tolerant lichen Xanthoria elegans (Link.) Th. Fr.

The growth curve for X. elegans produced by Osborn and

Taylor (1975) has long been the only published growth curve for a calcium-tolerant lichen in the Canadian Rockies. The curve is based on lichen size-age relationships on a variety of histor-

ically and tree-ring dated surfaces between Banff, Alberta, and Mount Robson, British Columbia (Fig. 1). Investigations of moraine ages at Bow Glacier by Leonard (1981) and at Athabasca Glacier by Luckman (1988), however, raise doubts concerning the accuracy of the three oldest control points used in the de-

velopment of this curve. The goal of our study was to use an independently cali-

brated data set to assess the validity of the X. elegans growth curve presented by Osbor and Taylor (1975). A fortuitous out- come of this work was the production of growth curves for both X. elegans and Aspicilia candida (Anzi.) Hue.

Abstract Lichen growth curves have been compiled from measurements of Xanthoria ele-

gans (Link) Th. Fr. and Aspicilia candida (Anzi.) on tree-ring dated moraines and

historically dated structures in the vicinity of Peter Lougheed and Elk Lakes Provincial Parks, Alberta and British Columbia. The newly calibrated growth curve for X. elegans and the previously published growth curve estimate radial

growth rates of 0.5 mm yr-~ for the first century. However, the new data show that the growth curve developed by Osborn and Taylor (1975) underestimates radial growth rates and overestimates substrate age beyond 150 yr. It should be

replaced by the newly constructed curve.

Aspicilia candida has lichenometric potential on carbonate-rich moraines in the front ranges of the Canadian Rockies. It has an ecesis of approximately 90

yr, followed by a maximum average growth of 0.9 mm yr-' for at least 60 yr. It is not known if the species has lichenometric potential beyond approximately 150

yr. More data are needed to better establish whether regional variations in growth and growth rates exist beyond the calibrated period.

XANTHORIA ELEGANS

Xanthoria elegans is a yellowish-orange through orange to reddish foliose to crustose lichen (Thomson, 1984; Clauzade and Roux, 1985). It has a circumpolar and alpine distribution, is found on all continents except Australia (Almborn, 1987) and was one of the first species to be used in lichenometry (Beschel, 1954). Despite this, we know of only three growth curves (Car- rara and Andrews, 1973; Osborn and Taylor, 1975; Chen, 1989) and a few short-term measurements that document the growth of this species (Table 1). The available data show X. elegans can have a short ecesis (10-30 yr), can grow quickly (0.2 to 1.75 mm yr-~) on a variety of substrates and may live for several centuries. Nevertheless, Osborn and Taylor (1975:111) report that X. elegans is less than ideal for lichenometry. They describe it as being: short-lived and relatively fast-growing, sensitive to natural fertilizers (i.e., orithocoprophilous), easily fragmented and weathered, rapidly out-competed by other plants and rare or absent for unknown reasons on some deposits.

While these limitations should be of concern to potential users, the response of X. elegans to natural fertilizers is likely the most disconcerting and least understood characteristic of the

species. Unfortunately, we know of no study that provides a detailed account of how the growth or establishment of this species is influenced by the addition of animal dung.

It is our observation that dung-fertilized X. elegans thalli are almost always found near rodent burrows or bird perches. At such sites thalli are thick, have irregular outlines and are clustered along nutrient and moisture-rich cracks and micropits on the crest of rocks. We have also seen that X. elegans thalli that have overtopped dung develop rippled surfaces and are only loosely attached to the rock. These thalli are invariably small and are excluded from lichenometric measurements because they overgrow or have marginal contact with other thalli. This makes it easy to recognize and avoid well-fertilized thalli and ensure that such thalli are an unlikely source of error in lichenometric

dating.

290 / ARCTIC AND ALPINE RESEARCH ? 1995 Regents of the University of Colorado 0004-0851/95 $7.00

FIGURE 1. Map of the study area.

Despite the concerns of Osborn and Taylor (1975), more recent research gives reason to question whether the lichenometric limitations of X. elegans have not been overestimated. Hooker (1980b), for example, reports X. elegans thalli that have lost their centers by fragmentation (i.e., thallus rings) can regen- erate. This observation implies that even the thallus rings of this species (cf. Beschel, 1961:1046) may have lichenometric utility. Furthermore, Hill (1984) reports that the circularity of X. elegans thalli may actually increase with age. This ability to maintain thallus circularity suggests a reduced likelihood for measurement

errors, less confusion regarding single and compound thalli, and accurate determination of thallus diameter by extrapolation from arcs of severely weathered thalli.

ASPICILIA CANDIDA

In the Canadian Rockies, A. candida (formerly Lecanora candida [Anzi] Nyl.) is often the only crustose lichen found in areas otherwise colonized by X. elegans. Little is known about its ecology or long-term growth. We have found that A. candida

D. P. MCCARTHY AND D. J. SMITH / 291

TABLE 1

Growth rate and ecesis data for Xanthoria elegans

Mean growth Ecesis Oldest lichen Site Lithology (mm yr-') (yr) L.I.C./age

Glacier National Park, Montana (Oelfke and Butler, 1985: 8) Slide Lake Limestone 0.30 30 21/70

South Orkney Islands, Antarctica (Hooker, 1980a: Table III, opp. p. 4) Shingle Cove Quartz-mica-schist 0.21a+(10) - not stated -

Factory Cove Quartz-mica-schist 0.38a2(8 - not stated - Lenton Point Quartz-mica-schist 0.17a2('8 - not stated -

Pageant Point Quartz-mica-schist 0.32a2+(5) - not stated -

Pageant Point Quartz-mica-schist 0.27a2+0) - not stated -

Spindrift Rocks Quartz-schist 0.50a5+(14) - not stated - Canadian Rockies (Osborn and Taylor, 1975: 111-120)

Banff to

Mt. Robson Carbonates 0.68b 10-20 85/253 San Juan Mtns., Colorado (Carrara and Andrews, 1973: 376-377)

Telluride Cement/marble 1.33 6c 70/59 Telluride Granite/granodiorite 1.76 16' 80/65 Telluride All of above 1.40 4c 80/65

Ouray All of above 1.50 21C 54/55

Qingua kujatdleq, West Greenland (Beschel and Weidick, 1973: 316) Outlet glacier

from Pakitsup ilordlia Basalt and gneiss 0.32 not stated 35/111

Axel Heiberg Island, Arctic Canada (Beschel, 1963: 212) White Glacier Basalt and gneiss 0.30d - not stated -

Southern Disko Island, Greenland (Beschel, 1963: 49) First and second

outlet glaciers at Lyngmarksbrae Basalt and gneiss 0.75 not stated 82/110

Tianshan Mountains, China (Chen, 1989: 1492)

Tianger Peak II Schist, gneiss, granodiorite 1.10b 30 115/403

aGrowth rate determined photographically. b Growth rate for first century, subsequent growth is 0.22 mm yr-'. c Estimated by linear extrapolation of growth line. d Annual growth rate determined photographically in 1960-1962. ' Growth in one year. 2 Mean growth for 2 years (1972-1974). 5 Mean growth for 5 years (1972-1977). + Site is nutrient enriched; frequent bird perch. '10) Number of thalli measured.

thalli are highly resistant to abrasion and do not exhibit central

fragmentation or coprophily. The thalli are always closely ap- pressed to the rock and have remarkably circular outlines. Ma- ture thalli have black apothecia and a bumpy, chalky-white ap- pearance (Noble et al., 1987; DeBolt and McCune, 1993). The

species also has bright, distinctive pigmentation that makes it

highly visible on dark lithologies. Unfortunately, for unknown

reasons, the species is absent or the populations are small at some sites.

Description of the Study Area The study was carried out at several locations in the Ca-

nadian Rocky Mountains. The Canadian Rockies consist of two

high-relief structural provinces with northwest-southeast trends. The Main Ranges form the highest peaks along the border between Alberta and British Columbia and expose both calcareous and siliceous lithologies. Immediately to the east, but separated from the Main Ranges by a series of major thrust faults, are the front ranges. The bedrock within this mountain range consists

primarily of strongly folded and thrust-faulted Palaeozoic car-

bonates and clastics (North and Henderson, 1954; Monger and

Price, 1979). The majority of data reported in this paper are from dated

moraines and structures located in the front range setting of Peter

Lougheed and Elk Lakes Provincial Parks, Alberta and British Columbia (Fig. 1). Data are also presented for lichens found on Barrier Dam on the Kananaskis River, and on moraines at the Athabasca and Bow Glaciers in Jasper and Banff National Parks (Fig. 1).

Climatological data are not available for the individual

study sites, but are available for the main valley at Kananaskis, Alberta (Fig. 2). Temperature-elevation trends estimated from stations in the vicinity of Peter Lougheed and Elk Lakes Pro- vincial Parks correspond to a mean lapse rate of 7.6?C km-'

(Cote, 1984:26). The greatest precipitation (up to 900 mm yr-') is found at high elevations close to the interprovincial border and the least amount of precipitation (approximately 500 mm

yr-') occurs on the floors of the main valleys. The net result is a vertical climatic gradient and a multitude of microclimatic set-

tings controlled by aspect and slope. Climatological data and detailed descriptions of the Athabasca and Bow Glacier fore-

292 / ARCTIC AND ALPINE RESEARCH

20 - /* MINIMUM

10 -

-10

TOTAL PRECIPITATION / MEAN RAINFALL - 100 mm MEAN SNOWFALL

h x 2 2 | | |- 50

J F M A M J J A S 0 N D MONTH

FIGURE 2. Climate normals for Kananaskis, Alberta, 1951- 1980. The station is located at 512'N, 115?2'W, 1390 m. Source: Environment Canada (1982).

fields are available in Osborn and Taylor (1975), Leonard (1981) and Luckman (1988).

Methodology The approach used in this research was first to establish the

age of a series of control surfaces. Preference was given to sites occupied by X. elegans, where the substrate age could be established from either historically dated structures or from an as- sessment of tree-ring records (e.g., Luckman, 1977; Desloges and Ryder, 1990). The largest thallus found on each dated surface was then used to establish a growth curve fitted to the largest thallus on deposits of different age (e.g., Benedict, 1967; Porter, 1981). In all cases, a complete search of the landform was used to locate the largest circular to oval thalli (<80% fragmentation) on each control surface (e.g., Webber and Andrews, 1973). Unless otherwise noted, all critical thalli are from grey limestone clasts that are embedded in the crest of a moraine.

The thallus size data were collected in the summer of 1989 using a flexible, transparent plastic ruler. Individual lichens were registered by including a measure of the Largest Inscribed Circle (L.I.C.) and the longest axis (Innes, 1985). Lichen size will therefore be reported as: L.I.C. x longest axis. Maximum-average growth rates were calculated by dividing the L.I.C. measurement by substrate age (i.e., substrate age = the date that measurement was taken minus the date that the landform surface became stable and/or exposed). Voucher samples of the lichens studied were identified by P. Y. Wong, National Museum of Can- ada and J. Sheard, Department of Biology, University of Sas- katchewan.

Tree-ring counts were done using sanded increment cores and a dissecting microscope. In each case all mature trees on the landform were sampled below 30 cm on the mainstem. Tree germination dates were adjusted for sampling height using data developed at Elk Glacier (McCarthy et al., 1990) and a tree ecesis of 24 yr was applied based on estimates done at Elk and Haig Glacier following McCarthy and Luckman (1993). Cross-

dating of an overridden snag at Elk Glacier was performed by M. E. Colenutt, Dendro-geomorphology Laboratory, University of Western Ontario, using a regional network of 11 chronologies (Colenutt, 1992, Colenutt and Luckman, 1994, Smith et al., 1995).

Results and Evaluation XANTHORIA ELEGANS DATA

The X. elegans L.I.C. and substrate-age data developed in this work are summarized in Table 2 and plotted in Figure 3. Detailed descriptions of the sampling sites and dating controls are given in McCarthy (1993) and Smith et al. (1995).

With the exception of the oldest control point (point 12), all points in Figure 3 match a single largest thallus size (L.I.C.) with the minimum age of the geomorphic unit on which it was found. Point 12, matches the largest lichen on Moraine I with the germination date of a snag that was overridden by Moraine II. This allows the construction of a growth curve that indicates radial growth continues beyond 75 mm L.I.C. Unfortunately, in- clusion of point 12 in the growth curve may overestimate growth rates beyond the first two centuries.

In Figure 3, a line representing the growth trend estimated from the L.I.C. data passes through five points (including point 12) and is near (within 10-25 yr) three others. An ecesis-dependent point from P6tain Glacier and data from Barrier Dam fall above the line. Presumably, all points would fall on the line if the long-term average lichen growth rates and tree ecesis were uniform on all surfaces. However, data from Barrier Dam (Fig. 1) and previous studies (Table 1) show that X. elegans growth rates are high at moist, low-elevation sites. Similarly, modem tree-ecesis data (McCarthy and Luckman, 1993) show that uni- formity in ecesis is rare and suggest it would be unreasonable to expect all tree-ecesis dependent points to fall on the growth curve.

Comparison of the growth curve developed in this study with that developed by Osbor and Taylor (1975) shows the two curves are identical for the first century, but diverge in their oldest sections (Fig. 4). This discrepancy is due, in part, to fun- damental differences in the approaches used in the two studies. Our newly developed growth curves come from a limited number of sites representing a small range of elevations and micro- climates. In contrast, the Osborn and Taylor (1975) curve at- tempted to reflect average conditions over a much larger range of elevations and climates. The main differences between the two curves may, however, be due to ambiguities in the three oldest control points employed by Osborn and Taylor (1975).

The oldest control point presented by Osborn and Taylor (1975) came from the outermost terminal moraine (Moraine I) at Athabasca Glacier (Fig. 1, Table 3). Unfortunately, most of that control surface has been destroyed by human activity and by 1989 only six acceptable X. elegans thalli (largest 53 X 71 mm) could be found at the site. Osborn and Taylor (1975) assumed that Moraine I stabilized in 1721 (Fig. 4, Table 3, point 23) within a few years of the date estimated from a tilted tree on a contiguous lateral moraine (Fig. 4, Table 3, point 24). How- ever, Luckman (1988:44,48) reports the control surface has a minimum date of only 1778 (tree A81131) (Fig. 4, Table 3, point 22). Consequently, the oldest point used by Osborn and Taylor (1975) remains enigmatic.

The second oldest point used by Osborn and Taylor (1975) assumed that ice retreated from Moraine II at Bow Glacier (Fig. 1) in 1852 (Fig. 4, Table 3, point 20). However, Leonard (1981)


TABLE 2

Data used to construct the growth curve for X. elegans

Lichen size Substrate agea/ Point (L.I.C. X largest calendar equivalent no. axis) (mm) (yr before 1989/yr A.D.) Location Dating control

A) Historical Control Points

1 19/19 73/1916 Haig Glacier, Moraine III At or near 1916 ice front mapped by International

Boundary Commission 2 30/37 73/1916 Haig "cairn" Erected by International Boundary Commission and

assumed lichen free in 1916 4 38/46 73/1916 Three Isle Lake cairn Erected by International Boundary Commission and

assumed lichen free in 1916 7 56/61 42/1947 Barrier Dam Assumed lichen-free at time of completion in 1947

B) Dendrochronologic Control Points

Date of earliest Germination Substrate

growth ring date surface age yr A.D. yr A.D. yr A.D.

3 34/43 58/1931 Elk Glacier, Moraine II 1931 1907 1907 5 42/63 109/1880 Putnik Glacier, Moraine I 1910 1904 1880

6 48/48 92/1897 P6tain Glacier, Moraine II 1921 1921 1897

8 66/70 129/1860 Elk Glacier, Moraine II 1891 1884 1860

9 66/70 197/>1792 Elk Glacier, Moraine II >1792

10 74/92 117/1872 Petain Glacier, Moraine I 1897 1896 1872 11 75/75 159/1830 Haig Glacier, East Moraine I 1862 1854 1830

12 95/116 425/1564 Elk Glacier, Moraine I 1595 1588 1564

found growth depression in a tree at the site and argued that ecesis was likely delayed. He suggested Moraine II probably formed sometime before 1852 (Fig. 4, Table 3, point 21).

The third oldest point (Fig. 4, Table 3, point 17) developed by Osbor and Taylor (1975) is from the second oldest terminal moraine (Moraine II) at the Athabasca Glacier (see Luckman,

1988). Here, Heusser (1956) used a tree rooted on outwash distal to the moraine to infer that the moraine stabilized in 1841. It has since been dated to 1853 (A8149) (Luckman, 1988:43) (Fig. 4, Table 3 point 16). If lateral-terminal moraine synchroneity at the Athabasca Glacier is assumed, then the surface of the control

site predates 1885 (tree AD8114 of Luckman, 1988:44) (Fig. 4, Table 3, point 18). Consequently, the third oldest control point used by Osbor and Taylor (1975) is equivocal.

In this study, we developed control points from Elk and

LONG

. 12=

I- Ll.C.

A9

0 BARRIER DAM A ELK D HAIG * HAIG "CAIRN" v PETAIN v PUTNIK * 3 ISLE L. W. "CAIRN"

-I LANDFORM DATE

/1

I''.1 '' ''1 . '' ''1''I'' 1' I''

) 50 100 150 200 250 SUBSTRATE AGE

300 350 400 450 (years)

FIGURE 3. Growth curves for X. elegans developed in this

study. Error bars show adjustments for tree ecesis and sampling height compensation. Dating controls are summarized in Ta- ble 3.

Haig Glaciers (Fig. 3, Table 2, points 8, 11, 12) and plotted a curve that accepts the revised control points. These data indicate that the Osborn and Taylor curve overestimates growth rates

beyond the first century.

ASPICILIA CANDIDA DATA

The dendrochronological and historical data from five sites

(Table 4) were combined to produce the A. candida growth curve shown in Figure 5. A growth-trend line has been drawn to in-

tercept four of the control points and is intended as an envelope "curve" (Innes, 1988:83). Points that fall below/above the line indicate slower/faster growth than is estimated by the line.

Points representing the first and second largest A. candida

120 -

E E 100 -

cj

- 80-

LJ

cn

J 60-

=D 40-

I <: 20-

n

22 23 _.._..

2 223 4* T THIS STUDY

20.. 21 OSBORN & TAYLOR (1975)

18 0-1 o ATHABASCA

14 13 o ROBSON 1011 12 v STRUCTURES

8Y Y * PEYTO 65 o4 o SLIDE LAKE, MONTANA

2 * S.E. LYELL v BOW

~~~/ - ~-I ECESIS '1''1''1''1''1'1 . . I I

I

11111 I .. I - .. I I

0 50 100 150 200 250 300 350 400 450 SUBSTRATE AGE (years)

FIGURE 4. Comparison of growth curves for X. elegans devel-

oped in this study and that presented by Osborn and Taylor (1975). Both curves use L.I.C. measurements. Error bars show

adjustments for tree ecesis and sampling height. Dating controls

for both curves are summarized in Tables 2 and 3.


120 -

E E100 -

Lr - 80-

1 40- I

-o -

20 -

0

. . . . . . . . I . . . . . . . . . . . I11 1 1 1 1 1

. . . . . . . . . . . . . . . . . 1 1 ' . . . . ' ' '. . ' ' . 1 1 1 -I

u

(

TABLE 3

Growth rate data for X. elegans after Osborn and Taylor (1975), Leonard (1981), Oelfke and Butler (1985) and Luckman (1988). Unless otherwise indicated, the dating controls are those given by Osborn and Taylor (1975)

Substrate agea/ Lichen size calendar equivalent Ecesis interval

Point No. (mm) (yr/yr A.D.) (yr) Description of size and dating control point

1 15 40/1934 12 Robson Glacier. Moraine 7, earliest ring of oldest tree 2 17 54/1920 12 Peyto Glacier. Moraine 7, earliest ring of oldest tree 3 17 50/1924 - Teahouse near Lake Louise built in 1924 4 21 64/1910 - Landslide, Slide Lake, Montana, historical account (Oelfke and

Butler, 1985) 5 22 62/1912 - Wall at Banff Springs Hotel built in 1912 6 25 51/1923 - Bridge at Banff built in 1924 7 28 67/1907 12 Peyto Glacier. Moraines 4 and 5, earliest ring of oldest tree 8 28 38/1936 - Mausoleum at Banff cemetery, on quartzite 9 29 38/1936 38 Mausoleum at Banff cemetery, on limestone

10 30 59/1924 12 Robson Glacier. Moraine 5, earliest ring in oldest tree 11 30 55/1919 12 Robson Glacier. Moraine 6, earliest ring in oldest tree 12 35 74/1900 12 Peyto Glacier. Moraines 3 and 4, earliest ring of oldest tree 13 40 82/1892 12 Peyto Glacier. Moraine 2, earliest ring of oldest tree 14 45 71/1903 12 Robson Glacier. Moraine 4, earliest ring in oldest tree 15 52 121/1853 - S.E. Lyell Glacier, roche moutonnee, historical account (Hector,

1863) 16 54 121/1853 17 Athabasca Glacier, oldest tree at distal base of moraine resam-

pled by Luckman (1988) and use of Heusser's 17-yr ecesis 17 54 116/1858 17 Athabasca Glacier, earliest ring of tree in outwash distal to mo-

raine 18 54 89/1885 17 Athabasca Glacier, earliest ring of tree on 2nd up-valley lateral

correlated with 2nd down-valley terminal (Luckman, 1988) 19 55 98/1876 12 Robson Glacier. Moraine 3, earliest ring in oldest tree 20 62 108/1866 14 Bow Glacier. Moraine 2, earliest ring in oldest tree + 14-yr

ecesis (Heusser, 1956) 21 62 123/1851 25 Bow Glacier. Moraine 2, + 25-yr ecesis (Leonard, 1981) 22 85 196/1778 17 Athabasca Glacier. Earliest ring of oldest tree on this outermost

terminal moraine segment, (Luckman, 1988) and Heusser's 17-yr ecesis

23 85 236/1738 17 Athabasca Glacier. Earliest ring of tree rooted in outwash chan- nel distal to search area terminal moraine (Heusser, 1956)

24 85 260/1714 -Athabasca Glacier. Tilted tree on up-valley lateral moraine correlated with search area moraine (Luckman, 1988)

aAge in years A.D. 1974 as reported by Osborn and Taylor (1975).

thalli on the outer moraine at Foch Glacier are shown in Figure 5 as points 7 (proximal crest) and 10 (distal crest). The rapid growth rate suggested by point 10 seems inconsistent with the lack of standing water or lush vegetation (lichens or other plants) on this moraine. It is therefore concluded that the distal crest of the lateral moraine (point 10) predates the crest and proximal slope (point 7) of the terminal moraine. Accordingly, point 10 is rejected and point 7 is accepted.

Points 8 and 9 are based on the largest A. candida thallus

(46 x 52 mm) measured on the western lateral moraine at Ath- abasca Glacier in 1989. Since the exact age of this surface is not known, the line representing the growth trend is drawn to fit the other controls.

While specific attempts were made to determine ecesis rates for this species, extrapolation from the growth curve suggests that A. candida has an ecesis of approximately 90 yr at these sites. Once established, the thalli grow at a maximum average rate of 0.9 mm yr-I for at least 60 yr.

Conclusion The results of our investigations suggest that radial growth

rates of X. elegans at various sites in the Canadian Rockies are

120 -

E

E100 -

- 80 -

LX w 60-

0 40-

D

< 20 - i-

I 1' 011

3/ 1' /?

/01

0 0 50 100 150 200 250

SUBSTRATE AGE 300 350 400 450 (years)

FIGURE 5. Growth curve and control points for A. candida (solid line) and growth curve for X. elegans (dashed line) developed in this study.


I I I t I l l I i, I I I I I I . .\ . i I I Il

TABLE 4

Data used to construct the growth curve for A. candida

Substrate age/ calendar

equivalent Lichen size (yr before 1989/

Point No. (mm) yr A.D.) Location Dating control

A) Lichenometric Control Pointsa

2 22 94/1895 Petain Glacier, Moraine III 45 mm X. elegans = 1895, youngest date pos- sible 1897 given by Petain Glacier Moraine II

6 45 121/1868 N.W. Northover Glacier 60 mm X. elegans = 1868 7 45 121/1868 Foch Glacier, Proximal crest of lateral 60 mm X. elegans on Moraine I terminal

moraine 8 46 114/1875 Athabasca Glacier, Moraine II lateral 56 mm X. elegans = 1875

10 60 121/1868 Foch Glacier, Distal lateral crest 60 mm X. elegans on Moraine I terminal 11 67 152/1837 Petain Glacier, Moraine I, upper fore- 74 mm X. elegans on Moraine I in lower fore-

field field to date Moraine I in upper forefield

B) Dendrochronologic Control Points

Date of earliest Germination Substrate

growth ring date surface age yr A.D. yr A.D. yr A.D.

1 13 92/1897 P6tain Glacier, Moraine II 1921 1897 3 27 109/1880 Putnik Glacier, Moraine I 1910 1904 1880 4 40 129/1860 Elk Glacier, Moraine II 1891 1884 1860 5 40 194/>1795 Elk Glacier, Moraine II >1795 9 46 146/1843 Athabasca Glacier, Moraine II Tilted tree at distal base of moraine

= 1843-1844, A84-1, Luckman (1988)

Ages based on X. elegans are estimated from the growth curve developed in this study.

remarkably similar, especially for the first century of growth. The data indicate that the regional growth curve for X. elegans presented by Osborn and Taylor (1975) is questionable beyond the first century and should be replaced by the newly constructed curve. The A. candida growth curve developed as a component of this research provides additional short-term dating control for carbonate surfaces in this region.

Potential users should recall, however, that the oldest portion of our X. elegans growth curve is a provisional estimate of

growth rates beyond the first two centuries. More and closer

dating controls are needed before this oldest portion of the

growth curve can be defined. Workers must also understand, that the sporadic and unpredictable distribution of these species in this and perhaps other regions limits their lichenometric potential. We hope that our findings will encourage others to inves-

tigate the use of lichenometry in alpine carbonate terrain. How-

ever, we caution that these environments are not well suited to the use of dating methods that use percentage cover, multiple species or quadrat sampling in lichenometry (e.g., Winchester, 1984, Matthews, 1974, 1975). Despite this, we believe that the

approaches demonstrated herein have the potential to provide accurate and reproducible ages for carbonate-rich surfaces.

Acknowledgments The research presented in this paper was supported in part

by an NSERC Operating Grant (A1930) awarded to D. Smith. We wish to thank J. Sheard and P. Y. Wong for identifying the lichen voucher samples and thank the various authorities at Jas- per National Park, Peter Lougheed and Elk Lakes Provincial Parks for allowing us to work at the study sites. Kelly Skuse

and Glen Blahut provided comic relief and much needed assis- tance in the field. Brian Luckman provided detailed maps of the Athabasca Glacier forefield. J. T. Andrews, J. L. Innes, E. M. Leonard, and K. A. Salzberg provide helpful comments on an earlier version of this paper. Ole Heggen, Cartographic Resource Center, Department of Geography, University of Victoria produced Figure 1.

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the Colorado Front Range, U.S.A. I. Establishing a lichen-

growth curve. Journal of Glaciology, 6(48): 817-832. Beschel, R. E., 1954: Eine Flechte als Niederschlagsmesser. Wet-

ter und Leben, 6-7: 56-60. Beschel, R. E., 1961: Dating rock surfaces by lichen growth and

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Ms submitted June 1994


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