-

ELSEVIER Forest Ecology and Management 78 (1995) 85-98

Fores;i;ology

Management

Potentials and limitations of ecological classification as a tool for forest management: a case study of disturbed deciduous

forests in Quebec

Philippe Nolet a7 * , GCrald Domon b, Yves Bergeron a a Groupe a’e Recherche en lkologie ForestiZre, Universitk du Qu&bec b Montrial, C.P. 8888, WCC. A, Montrial, Qut. H3C 3P8, Canada

b Facultt? d’Am&agement, Universitt? de Montrkal, C.P. 6128, succ. Centre-Ville, Montrt?a( Qub. H3C 357, Canada

Accepted 22 May 1995

Abstract

The forest ecosystems of the most southern part of Quebec have been influenced by activities of various intensities and spatial distribution (harvesting, agriculture, pastureland, etc.) since the beginning of settlement. These activities have changed the natural relationships between soil and vegetation characteristics. The aims of this study were (1) to identify, with a concrete example, the difficulties encountered by a forest ecological classification program in disturbed deciduous forests and (2) to evaluate the efficacy, for forest managers, of the products derived from that classification. We first show, by a complete linkage cluster analysis, that the forest type concept is not appropriate for the disturbed deciduous forests context because of the great variability in forest composition and the low relationships between soil and vegetation. By comparing information required to manage effectively the disturbed deciduous forest to the information provided by the Qu6be.c Forest Ekological Classification Program (FECP), we demonstrate, in the second part of the study, some shortcomings of this program at the site scale. Three main reasons are given to explain the weaknesses encountered. The first one is that the disturbed characteristics of these forests increase the complexity of forest and ecological mapping. The second main reason is that some vegetation and soil characteristics are so variable that it is almost impossible to predict them with ecological classification. Deficiencies in the methodology used in the Quebec FJXP constitute the third main reason explaining the weaknesses encountered. From these results, we propose some solutions to enhance the support that the Quebec FECP can bring to forest managers in disturbed deciduous forests. Finally, shortcomings of ecological classification at the site scale do not necessarily limit its potential at the regional scale.

Keywords: Deciduous forest; Disturbed ecosystem; Ecological classification; Ecosystem mapping

1. Introduction

Several forest management objectives justify the development of an ecological classification of forest

* Corresponding author.

habitats. Some examples include: evaluation of site productivity; knowledge of competitive vegetation after harvesting; protection of fragile habitats; identi- fication of site vulnerable to windthrows; species choice for reforestation; prediction of plant succes- sion (Jurdant et al., 1977; Barnes, 1986). To attain these objectives, different programs have been estab-

037%1127/95/$09.50 0 1995 Elsevier Science B.V. All rights reserved

SSDI 0378-l 127(95)03595-8

86 P. Nolet et al. /Forest Ecology and Management 18 (19%) X5-98

lished in most Canadian provinces (Corns, 1992; Wells, 1992; Bowling and Zelazny, 1992; Meades and Roberts, 1992; Sims and Uhlig, 1992; MacKin- non et al., 1992) and other countries (Christian, 1951; Pfister, 1977; Rogister and Galoux, 1982; Avers and Schlatterer, 1991; Brenner and Jordan, 1991). Thus, over the past few years, many papers have been published to describe forest ecological classification programs. Most of these papers have explained the usefulness of programs for large terri- tories (countries, provinces or states). However, pa- pers that have evaluated these programs for their efficacy in particular regions are limited (e.g. B6land et al., 1992).

Similarly to other provinces and countries, the Mini&e des Richesses Naturelles du Qu6bec (Ministry of Natural Resources) has established the Qu6bec forest ecological classification program (FECP) (BCland et al., 1992; Bergeron et al., 1992, 1993). In this paper, we are interested in the efficacy of the Qu&bec FECP (and consequently in ecological classification) in heavily disturbed deciduous forests. These forests are situated in southern Qu6bec and are usually on private lands. The sites on which we find these forests today have been used for activities of various intensities and spatial distribution (harvest- ing, agriculture, pastureland, etc.) since the begin- ning of settlement. This brings a great complexity to relationships between abiotic variables (soil, slope, exposure, etc.) and the distribution of species. Con- sequently the ecological classification of these forests becomes difficult (Bergeron et al., 1988; Archam- bault et al., 1989; Leduc et al., 1992; Meilleur et al., 1994). This description of the disturbed deciduous forests highly contrasts with the rest of Qu6bec forests which consist mainly of boreal and mixed conifer-hardwood forests. These more northern forests have not been touched by human activities to the same degree of complexity as deciduous forests and most of them are on crown lands.

Some authors have recently pointed out that sev- eral schools or methods of ecological classification have failed to attain their main objective, which is to produce effective management tools (Damon et al., 1989; BClanger et al., 1990). Low consideration given to particularities of management contexts may ex- plain this fact (Domon et al., 1989). It is then interesting to verify if the Qukbec FECP indeed

takes into account the particular case of disturbed deciduous forests in QuCbec to the degree that ii should.

In order to verify this. the main goals of this paper are: (1) to identify the difficulties encountered by ecological classification in disturbed deciduous forests; (2) to evaluate the efficacy of the products derived from that classification for forest manage, ment. More precisely, we will first evaluate the use of forest type concept in disturbed deciduous forests. The forest type (see next section) is a key componeui of ecological classification in the FECP. Because oi the great diversity observed in disturbed deciduous forests, the usefulness of forest type concept must be reviewed. Second, we will evaluate how the products derived from that classification provide the ecologi- cal information needed for managing disturbed de- ciduous forests. In the Qu6bec FECP, these products are the physiographic sere. the integrated forest map. and the field guide to the identification of fores!

types (see next section for description). Forest ecologists usually agree to distinguish two

types of information: factual information and inter- pretative information. The following example will make this distinction clearer: texture and soil drainage are necessary to choose the best species for reforesta- tion. Texture and drainage represent here the factual information. The relationships between species pro- ductivity and theses variables represent the interpre- tative information. Factual information is then very important because it is necessary to make valid interpretations. This paper limits its evaluation tu factual information because most of the interpreta- tive information has not yet been elaborated in the Qdbec FECP. This paper also limits its evaluation to site scale (as opposed to regional scale) because evaluation of information at the regional scale is part of a very different context and would consequently need an another study.

2. The QuBec FECP

This description of the Qu&ec FECP is quite brief since recent papers have described it well (B& land et al., 1992; Bergeron et al., 1992, 1993). The conceptual framework of the Qu6bec FECP is largely based on the work of Jurdant et aI. (1977), though it

P. Nolet et al./Forest Ecology and Management 78 (1995) 85-98 87

integrates concepts of various schools of ecological classification (see Bergeron et al., 1993). In the approach developed by the FECP, territory is subdi- vided into levels of increasing resolution, from the ecological region to the ecological district and then to the ecological type. The ecological region is de- fined as an area characterized by a distinctive re- gional climate, as reflected by vegetation (Jurdant et al., 1977). The ecological regions are represented at the scale of 1:1250 000 (Thibault and Hotte, 1985). The ecological district is composed of units compris- ing distinctive patterns of relief; they are represented at the scale of 1:25000 (Robitaille, 1988). Finally, the ecological type is defined by a relatively uniform combination of soil and plant succession (Jurdant et al., 1977). In the FECP, soil is described by surface deposits and its thickness, the soil drainage class and the slope class; plant succession is expressed by the successional series, regrouping all chronosequences leading to the same stable forest type. The identifica- tion of ecological type is based on field surveys during which data on geology, geomorphology, to- pography, soils and the various forest strata are collected (Saucier et al., undated). The ecological types of a region are first represented on a physio-

graphic sere, which is a schematic representation (toposequence) of relationships between abiotic vari- ables and vegetation. Initially, ecological types were to be mapped at the scale of 1:20000, but, because of high costs, the idea was abandoned. Presently, surface deposits are mapped at a scale of 1:50000 (using 1:40000 aerial photographs) and then en- larged to a 1:20000 map. The latter map is trans- formed into a surface deposit-drainage map using the physiographic sere and superimposed by a tradi- tional forest cover map (1:20000). This new map is called the forest ecological map.

In addition to these subdivisions (ecological re- gion, district and type), the FECP uses the concept of forest type that is characterized by the homogeneity of its overstory and understory species composition and by the homogeneity of its structure and its physiognomy. A forest type occupies sites whose abiotic characteristics are situated within known lim- its. It possesses its own successional dynamics and a particular distribution (Bergeron et al., 1992, 1993). Forest types are described in field guides that ensure the link between ecological and forest surveys. The forest types are also used to elaborate the succes- sional patterns of a given region. Lastly, forest types

Fig. 1. The study area: the Cazaville sector of the Municipalitt Rkgionale de Comtt (MRC) of Haut-St-Laurent.

88 P. Nolet et al. /Forest Ecology and Management 78 (1995) 85-98

are at the base of identification of treatment units. A treatment unit is a relatively homogeneous land unit in terms of site conditions and forest composition for which outcome and productivity may be expected from a given silvicultural regime (Delpech et al., 1985). The exact content of the treatment unit field guides is not yet determined.

In summary, the products derived from FECP that may provide information at the site scale are the forest ecological map, the physiographic sere and the forest type field guide. The forest ecological map is the only product that provide spatial information. The physiographic sere increases information given by the forest ecological map by serving as a photoin- terpretation key and by providing information on texture, stoniness, vegetation chronosequence, etc. The forest type field guide gives relatively the same information than the physiographic sere, but this information is more detailed and not schematically presented.

3. Methodology

3.1. The study area

The study area, which is part of the disturbed deciduous forests, is the Cazaville sector of the Municipalite Regionale de ComtC (MRC) of Haut- St-Laurent. This is the southernmost region of Quebec (Fig. 1). The area has about 3250 degree-days per year (Rousseau, 1974) and an average of 140 frost-free days. The mean minimum temperature is - 10°C in January and the mean maximum tempera- ture is 20.8”C in July. Mean annual precipitation is 960 mm, including 250 mm in snow.

The territory under study is part of the Montreal et Haut-Richelieu ecological region (Thibault and Hotte, 1985) and the Great Lakes and the St Lawrence River forest region (Rowe, 1972). In the Haut-St- Laurent section, deciduous forest cover (Sugar Maple, Beech, Red Maple, Basswood) is most com- mon on deep calcareous soils, while conifers such as the Eastern Hemlock, Eastern White Pine, Balsam Fir, and White Spruce grow on thin acidic or eroded materials (Rowe, 1972). The region is characterized by a flat relief rising from 50 m to 100 m. The dominant surface deposits are tills deposited during

the last glaciation, and clays deposited by the Cham- plain Sea. In addition, polygenic forms are found, i.e. deposits arising from a combination of glacial and postglacial events. The surface deposits cover dolomite, a calcareous sedimentary rock (Bariteau, 1988).

The region is mainly agricultural, with 65-70s of its soils dedicated to agriculture. Forests have been generally limited to soils that are unsuitable far farming. Most of the forests have been harvested several times since colonization (Stellar, 1988; Bouchard et al., 19891. Today, these woodlots are principally used as a source of firewood and as pastureland (Domon et al., 1993). Furthermore, the effects of anthropic disturbances on forest communi- ties in the Haut-St-Laurent region have been reported by many authors. Bergeron et al. (1988) observed that forested ecosystems of the Haut-St-Laurent re- gion represent a gradual replacement of a natural system, controlled by species adaptation to environ- mental gradient, by a system controlled by anthropic disturbances. Without explaining their results di- rectly by the importance of anthropic disturbances, Leduc et al. (1992) found that spatial distribution of more than half of the tree species studied do not track the measured environmental variables. Meilleur et al. (1994) found that most of the forest communi- ties not statistically related to the most important environmental factor (in this case morphogenic fea- ture) showed important disturbed history.

3.2. Inventory data

The data used in this study were part of a multi- disciplinary framework elaborated by Bouchard et al. (1985). During the summers of 1983 and 1984, 191 plots of 0.02 ha (10 m X 20 m> were sampled in the eastern portion of the MRC. of Haut-St-Laurent by Meilleur (1986). A first photointerpretation (1:15 000) was done in order to obtain a preliminary map of geomorphalogical systems (Bariteau, 1988). Sampling plots were determined following a second photointerpretation (1:2OooO). Exploratory field trips were made to choose visually homogeneous forest area. A sampling plot was chosen by walking 50 steps in a random direction from the center of each homogeneous forest areas. At each plot, the diameter at breast height (DBH) of all the trees- was measured.

P. Nolet et al. / Forest Ecology and Management 78 (1995) 85-98 89

Furthermore, all saplings (l-10 cm DBH) were counted along a 20 m X 1 m strip. The following information was also collected for each plot: the geomorphological feature, landform, parent material, texture, the topographical situation, the microtopog- raphy, the position on the slope, exposure, submer- sion, stoniness, drainage and ground water depth. This physical description of plots allowed us to identify their geomorphological type (surface de- posit-drainage combination) according to the Qutbec FECP codes. Finally, we must mention that 19 plots were rejected from the analyses because of missing data.

3.3. Data analysis

The evaluation of forest type concept for dis- turbed deciduous forests and the evaluation of the products derived from ecological classification need two distinct methodologies.

3.3.1. Forest type concept

3.3.1.1. Identification of forest types. In order to define forest types, we first divided the vegetation data set into three classes: saplings (l-10 cm DBH), trees lo-20 cm DBH and trees 20 cm and more DBH. These subdivisions were made in a manner to integrate vegetation structure of plots in the elabora- tion of forest types. This gave us a matrix in which every plot is described in terms of the numbers of individuals by species and diameter class. A matrix of similarity was then computed using the Kulczyn- ski’s similarity index. This similarity index was cho- sen because it is adapted to raw abundance data and because it eliminates double zeros in the matrix (Legendre and Legendre, 1984). Afterwards, we per- formed a complete linkage cluster analysis. This analysis assures that every sample in the group is similar (degree of similarity depends of the connex- ity level) to every other plot in the group. This method was preferred to other less constraining clus- tering methods (e.g. single linkage or arithmetic average clustering) or to divisive methods (e.g. TWINSPAN; Hill, 1979) which can lead to high intra-group variability. Data computation (matrix of similarity and clustering) was done with the software package ‘R’ (Legendre and Vaudor, 1991).

Two reasons explain why we did not use shrub and herb species in our analysis. First, their utiliza- tion together with trees and regeneration would have rendered even more difficult the grouping of plots into forest types, by increasing the heterogeneity of the plots. Second, Meilleur et al. (1992) have already written an interesting and complete article on this topic using, in part, the same data set as in this paper.

3.3.1.2. Relationships between soil and vegetation. We do not intend to make a complete ecological classification of the territory under study in this paper. Other studies in the same territory have al- ready discussed different aspects of ecological classi- fication (Bergeron et al., 1988; Brisson et al., 1988; Leduc et al., 1992; Meilleur et al., 1992, 1994). Thus, to express the relationships between soil and vegetation variables, we will limit our work to a contingency table between forest types and geomor- phological types.

3.3.2. Evaluation of the products

3.3.2.1. General evaluation. The procedure followed consisted of three steps: (1) identification of the main silvicultural activities on private forest land; (2) identification of the corresponding ecological infor- mation required; (3) an evaluation of the usefulness of the various products derived from FECP in re- spect to the information required.

First, the main sylvicultural activities were pri- marily identified through the grant program for pri- vate forest improvement (Anonymous, 1992) of the Ministry of Natural Resources (MRN) of QuBbec. This program describes a list of activities funded by the provincial government. The document ‘R&ussir ma For& (Anonymous, 1988) by the Canadian For- est Service was also used to identify activities. Table 1 shows the list of activities we used. The second step was to identify the ecological information re- quired to accomplish the various activities. Complete and precise definitions of these activities have al- lowed us to infer the main ecological information required for each of the silvicultural activities. The third and final step was to evaluate how the products of the FECP could supply the ecological information required. The evaluated products were the physio-

90 P. Nolet et al. /Forest Ecology and Management 78 (1995) 85-98

Table 1

Most important forest activities in private deciduous forests in Quebec

General activities

A. Land preparation for reforestation

Specific activities

1. Slashing and cleaning of logging residue

2. Soil scarification

B. Reforestation 3. Species choice 4. Planting

C. Plantation maintenance 5. Competitive vegetation control 6. Fertilization 7. Plantation protection

D. Cuts 8. Selection cutting 9. Regeneration cutting

10. Shelterwood cutting

11. Release (liberation) cutting 12. Sanitation cutting

13. Precommercial thinning 14. Commercial thinning

E. Miscellaneous 15. Machinery choice for timber extraction

16. Road construction 17. Forest protection

graphic sere, the forest type field guide and the forest ecological map. The evaluation of a specific product depended on the presence and the quality of the information supplied. Three classes of information quality were used: (1) no information, if no informa- tion is provided by the product; (2) partial informa- tion, if the product supplied only part of the required information and/or if the supplied information was not sufficiently reliable; (3) complete information, if the product provided all necessary information in terms of quantity and quality.

3.3.2.2. A specific evaluation of the forest ecological map.

Because the forest ecological map is the most important FECP product for forest management, an additional evaluation of this product was made. This was done by comparing the description of vegetation in the Meilleur data set (for which we have the spatial coordinates) to the description of vegetation on the forest map (obtained by photointerpretation and used to produce the integrated forest map). This map was made in 1983. For every plot sampled by Meilleur, there were five possibilities of evaluation

for the forest map: (1) the two first dominant species are identified on the forest map; (2) the first domi- nant species is correctly identified; (3) only the second dominant species is correctly identified; (41 none of the two dominant species is correctly identl- fied; (5) the forest map gives a vague identification (e.g. tolerant hardwoods). Afterwards, the number of plots that fell into each of the four categories was added up. This calculation provided an objective analysis of the accuracy of the forest ecological map in terms of composition and thus of its usefulness for forest managers.

4. Results and discussion

Evaluation of the forest type concept and of the Quebec FECP products will be presented and dis- cussed first separately, then they will be put into relation in the general discussion.

4. I. Evaluation of usefulness of forest type

The clustering of the 172 plots lead to the results presented in Table 2. Forest types, by definition, need to be homogeneous. This is why we considered first the samples grouped at a connexity level of 75%. Most of the samples (139) were not grouped at this level and most of the groups formed were composed of only two samples (Table 2). This level of division was then of low interest. At a connexity level of 50%, there were 51 groups (two or more samples) formed but there were still 31 samples that could not be regrouped. This difficulty of forming groups is not surprising since other studies have demonstrated that a large number of species combi- nations is possible in this region (NoIet, 1993; Meilleur et al., 1994). A short description for each

Table 2 Number of groups by group height and connexity level

Connexity Groups of Groups of 2 Groups of 3 Groups of 4 level 1 sample samples samples samples or more

0.75 139 13 1 1

0.50 37 28 13 9

Results obtained from a complete linkage clustering method.

P. Nolet et al. /Forest Ecology and Management 78 (1995) 85-98 91

diameter class of the groups formed at a connexity level of 50% is presented in Table 3. We considered that groups formed at connexity levels lower than

Table 3 Description of groups, obtained at a 0.50 connexity level, by

diameter classes

Groups n DBH km)

l-10 10-20 > 20

I 5 asa, 2 2 asa, fgr

3 4 asa 4 6 asa, ( ) 5 2 asa

6 2 asa 1 2 asa 8 2 asa

9 2 tam 10 2 ovi 11 2 ovi

12 2 fgr 13 2 asc

14 3 uam 15 3 uam 16 3 uam, fni

17 2 aru, fpe 18 2 uam, aru 19 2 aru, uam

20 3 uam, tot 21 2 uam

22 3 uam, fpe

23 2 pba, uam 24 2 tot

2s 3 tot 26 3 tot 27 4 tot 28 2 tot 29 2 tot 30 2 fni 31 2 fni 32 3 ovi, fni, asa

33 3 asc, fpe

34 4 asc, fpe 35 4 fpe

36 2 fpe, tot 37 2 tea 38 3 tea

39 2 aru

40 3 aru 41 3 aru

42 3 am, fni

43 2 bal, aru 44 3 pst, aru 45 2 pba

tam, asa, cc0 ( ), cm, pgr, asa tea, pgr, tam, ( 1 asa, tam, aru ( ), aru asa pgr, asa, tam tam tam

cc0 ( ), cc0 fni, tot bal, pst bal, tea tea

tam, uam tam, tot ovi bpa, aru ovi asa

tam, fgr tam, fgr asc ax, ( ) fpe, uam fpe ( ), uam, ptr ptr, ( ) pba, ( ) pba, ( ) a*, foe 0 uam 0 uam, ( 1 jci, ( ) 0 0 tram tram

uam, ( 1, fpe pba, fpe, ( 1 pba, uam pba, ( 1 tot tot

tot, tram ptr, uam, ( ) tot, fni tot tot tram, tot, ( )

lla lla fpe, tot tea, ( ) bal, ptr bal, ptr

aru, uam fni tot, tram, tam tot, asa, ( )

fpe, asc, ( ) asc, ptr, ( 1 fpe, asc fpe, asc

asc, ( ), fpe asc, ( ), fpe pba, ( 1 pba, ( 1 tea tea, ovi

tea aru, bal aru asi, ( 1 aru aru, tram

bo, pst bpo, pst fpe, fni fam, ( 1 aru aru

Pst pst, ( ) @a @a

Table 3 (continued)

Groups n DBH km)

l-10 10-20 >20

46 5 pde, am, ptr, ptr ptr, ( ) asa

47 4 aru, bpo w, ptr, bpo uam, ( 1 48 4 bo bpo, pba, ovi ( ) 49 2 bpo bpo ptr, ( ) 50 2 uam bpo uam, t ) 51 2 fni, bal bal bal

A name of a species indicates that this species is the dominant one

in at least one sample of the group. Consequently, when there is only one species in a class, it indicates that samples of the group are quite similar for that class. Conversely, a group shows more

variability for a class when two or more species are indicated for that class. ( ) means that there were no trees in that specific class

for at least one sample of the group; aru, Acer rubrum; asa, Acer saccharum; asc, Acer saccharum; bal, Bet&a alleghaniensis; bpa, Betula papyrifera; bpo, Betula populifolia; cco, Carya cordi-

formis; fam, Fraxinus americana; fgr, Fagus grandifolia; fni, Fraxinus nigra; fpe, Fraxinuspennsyluanica; jci, Juglans cinerea; lla, Larix laricina; ovi, Osttya uirginiana; pba, Populus balsam-

ifera; pde, Populus deltoides; pgr, Populus grandidentata; pst, Pinus strobus; ptr, Populus tremuloides; tam, Tilia americana; tea, Tsuga canadensis; tot, Thuya occidental& uam, Ulmus amer-

icana

50% were of no interest in the identification of forest types because the latter have to be homogeneous.

Table 3 first shows that the disturbed deciduous forests are a complex multidimensional continuum and then explains the difficulty of forming groups. One may also observe that some of the groups (forest types) contain an appreciable degree of het- erogeneity for each of the diameter classes. We recall that one of the most important objective in defining forest types is to use them In the elaboration of treatment units. The heterogeneity of the forest types described in Table 3 would not allow one to elaborate such treatment units. For example, a forester could not necessarily prescribe the same sylvicultural treatments for a stand dominated by Largetooth Aspen (Populus grandidentata) as for another dominated by Sugar Maple (Acer saccha- rum) (Group 4 in Table 3).

We observe in Table 4 that various forest types may be found on more than one surface deposit type and/or drainage class. Since various studies (e.g. Carmean, 1975) have demonstrated the important effect of different abiotic variables on site productiv-

92 P. Nolet et al. /Forest Ecology and Management 78 (1995) 85-98

ity, it would be misleading to base productivity studies on forest types that are supported by different types of soils. We also observe that most of the geomorphological types (surface deposit-drainage combination) support various forest types. These re- sults coincide with many studies in disturbed decidu- ous forests that demonstrate the difficulties in under- standing the relationships between soil and vegeta- tion (Archambault et al., 1989; Lamontagne et al., 1991; Meilleur et al., 1992). One could argue that the forest types found on a geomorphological type represent the various stages of a single vegetation chronosequence. However, many authors, in the same study area, observed various successional pathways on most of the soil types (Bergeron et al., 1988; Leduc et al., 1992; Nolet, 1993). It remains possible that they arrived at this conclusion because of the difficulty to obtain reliable historic information (time since perturbations; nature and intensity of the per- turbations), but even with this information, relation- ships between soil variables and vegetation composi- tion would still be extremely complex.

The methodology used may have contributed to overestimate the plots heterogeneity (and conse- quently increasing the number of forest types and/ or the heterogeneity within forest types) in three ways. First, the complete linkage cluster analysis is very constraining. The reasons we used this analysis have already been explained. Second, we have incorpo- rated regeneration (l-10 cm DBH) in the characteri- zation of the plots. We felt that it was essential to include regeneration because, had we abandoned it, we would have omitted a part of forest structure that is important for forest management. Third, the rela- tive small size of the plots (10 m X 20 m) may also have contributed to the observation of a high number of forest types. However, other studies in Quebec deciduous forests (Gauvin and Bouchard, 1983), us- ing the point-centered quarter method in 1 ha forest tracks, also showed a high diversity of forest types.

4.2. Evaluation of FECP products

Before presenting and discussing the results of this section, we will provide two examples to make clear how the results of Table 5 have been obtained. The first example involves the need to know soil texture. The forest ecological map identifies a sur-

face deposit for every stand on the map. The texture of that surface deposit is described in both the physiographic sere and the forest type field guide. The reliability of surface deposit (and then of tex- ture) appearing on the forest ecological map de- pends, of course, on the quality of the photointerpre- tation. This quality is reduced by the following two facts: (1) With 1:40000 aerial photographs, it is possible to observe more than one surface deposit in the same polygon (but only one is identified on the map). It is also possible that the surface deposit identification is simply not the good one. (21 The forest cover in disturbed deciduous forests cannot really help the photointerpreter to identify the sut- face deposit (see previous section); this consequently increases the possibility of a misidentification,

The prediction of abundance of advance regenera- tion will be our second example (last section of Table 5). Two reasons may explain why the Quebec FECP provides only a part of the information re- quired for the description of regeneration. First, re- generation is so variable that it makes its prediction from ecological variables very difficult @6land et al., 1992). Second, the only information about regen- eration encountered in the Quebec FECP products is found in the forest type field guide; this information cannot reliably be used by forest managers because the link between the forest cover described on the map and the forest type described in the field guide is not clearly made.

A more general look at Table 5 can now be taken. One may rapidly observe that, generally, FECP prod- ucts do not provide the information required to man- age effectively the disturbed deciduous forests. This observation becomes more interesting when we look at the reasons why, as outlined in the last column of Table 5. We may first observe that ‘impossible to predict’ (IPE) is often given as a reason to explain the missing information. This is because parameters such as advance regeneration (B&and et al., 1992). stem quality, presence of seed banks and presence of nutrients (Lechowicz and Bell, 1991) are. by their very nature, so variable that it is almost impossible to predict them, even if great improvements were made to the Quebec FECP. The fact that ecological classification cannot provide all the information re- quired for some specific variables demonstrates that ecological classification cannot resolve all the forest

93 P. Nolet et al. /Forest Ecology and Management 78 (1995) 85-98

Table 4 Contingency table between surface deposit-drainage combinations and the groups formed by the complete linkage clustering

Drainage 2-3 4-5 6

Surface la deposit

6s/la 2a 4p 6s 9s la 2a 3an 4ga 5a 4p 6s 7e,7t

1

2 3 4

5 6

7 8 9

10

11 12 13

14 15

16 17

18 19 20

21 22 23

24 25 26

27 28 29

30 31 32

33 34

35 36 37

38 39 40

41 42 43 44

45 46 47

48 49 50

51

*** ** **

* **

** *** * * *

** *

** *

*

* *

* * * *

* * ***

* * **

*

* *

* *

**

**

*

* * *

* **

*

** ***

* * * *

** *

* *

* * **

* * * * *

* *

* *

*

*

*

* * *

*

*

* **

* *

*

*

**

*

* ** * *

* * * *

* *

** * ***

****

**

Drainage classes: 2, well drained; 3, moderately drained; 4, imperfectly drained; 5, badly drained; 6, very badly drained. Surface deposits: la, undifferentiated till, 2a, fluvioglacial, 3an, ancient alluvial, 4ga, deep glaciolacustrine (clays); 4p, actual lacustrine littoral; 5a, deep glaciomarine (clays); 6s, ancient marine littoral; 6s/la, undifferentiated till coverd by a thin ancient marine littoral deposit; 7e, thick (> 1

m) organic; 7t, thin ( < 1 m) organic; 9s, stabilized dune.

94 P. Nolet et al. / Forest Ecology and Management 78 (1995) 85-98

management problems, as already pointed out by Sims et al. (1986), and that a good sylvicultural treatment will always need a sampling of every stand to be managed. This contrasts with the current ten- dency in Quebec, in which field work is limited to a minimum.

Another main reason explaining weak results ob- tained by products derived from ecological classifi- cation is the complexity of disturbed deciduous forests (CRD in last column of Table 5). We saw in the previous section that forest types are difficult to define in these forests and, consequently, this com- plicates the relationships between stand composition and physical variables. The mapping of ecological

(physical and vegetation elements) units, which de- pends in great part on these relationships, is then much more laborious. Table 6 presents an evaluation of the forest map (which is the vegetation part of the forest ecological map) of the territory under study. The results show that most of the descriptions of forest cover on the map and in samples do not match, demonstrating the difficulties of photointer- pretation in disturbed deciduous forests. The forest cover map analyzed was made before Quebec FECP was in place, therefore no physiographic sere was available to help for photointerpretation. However, it is likely that the physiographic sere would not have been very useful since physical variables may not

Table 5

Evaluation of how Quebec FECP provides factual information required for forest management in disturbed deciduous forests (see various activities described in Table 1)

Factual information required FECPs products Major reasons explaining .--._

1 2 3 the missing information

Soil characteristics

Thickness (2,3,15) a Texture (1 2 3 5 7 8 9 10,11,15,16,17) ,tv,,,, Drainage (1 7 3 5 7 8 9 10,11,15,16,17) ,-, I I 1 1 1 Stoniness (2,3,15,17) Hardness (compaction) (2,3,4)

Humus type and thickness (2,15,16)

pH(31 Nutrients (3,6) Presence of seed banks (2,9)

Other physical characteristics

Slope (15,161 Ground roughness (15,161

Stand characteristics

Stand structure Presence of even-aged or uneven-aged forest

(8,9,10,11,14)

Abundance of advance regeneration (8,10,111 Stand composition

Mature trees (poles) (8,9,10,11,12,13,14,)

Regeneration ((S,lO,llI Merchandable volume (8,9,10,11,14) Stem quality and health (8,9,10,11,12,13,14)

Presence of seed trees for regeneration cutting (9) Presence of seed trees for shelterwood cutting (10)

Stands affected by insects or disease (12)

n n

n

P

P P P

P P

P 11

n n

n

P

”

P

P n

P

CRD. 1NW CRD, [NW

CRD, INW CRD. INW IPE

CRD, INW INWJPE INW,IPE

IPE

INW

INWJPE

CRD, INW

INW. IPE

CRD, INW INW, IPE

CRD. INW IPE CRD, INW CRD, INW IPE

a Numbers in parentheses refer to activities of Table 1. Product No. 1: physiographic sere. Product No. 2: field guide for identification of forest types. Product No. 3: forest ecological map. n, the product provides no information; p, the product provides partial inforfnatiom CR-D,

complexity of soil-vegetation relationships and/or forest heterogeneity in disturbed deciduous forests (see text& INW, intrinsic weaknesses in Quebec FEC Program (see text); IPE, impossible to predict adequately from ecological classification (see text).

P. Nolet et al. /Forest Ecology and Management 78 (1995) 85-98 95

Table 6 Comparison of stand composition descriptions from forest map

with descriptions from field plots

Comparison class Number of samples a

The two first domianat species of field sample

are identified on forest map

0

First dominant species of field sample is identified on forest map

19

Second dominant species of field sample is 9

identified on forest map

None of the two first dominant species of

field sample are identified on forest map

60

Forest map just gives a vague description of forest composition (e.g., tolerant hardwoods)

76

a There are eight samples for which spatial coordinates are

missing.

help much in stand composition identification be- cause of the weak relationships between physical and vegetation variables. Thus, this interpretation ex- plains why the disturbed aspect of these forests decreases the quality of information provided by the FECP products, especially the forest ecological map.

The third main reason explaining the unsatisfac- tory results obtained by the Quebec FECP is its intrinsic weaknesses (INW in last column of Table 5). This reason is of less interest for readers outside Quebec. Briefly, however, we can mention: (1) the number of field plots used for ecological and forest cover mapping is insufficient; (2) the methodology that consists of enlarging a 1:50000 surface deposit map to a 1:20000 map is not legitimate in terms of cartographic principles; (3) the link between vegeta- tion description in forest type field guide and vegeta- tion description on forest maps is not clear.

We must mention that it is not the first time that some of the limitations of ecological classification are outlined for detailed forest management. Many authors have reported some problems with ecological classification or soil classification, especially for the prediction of site productivity (Schlenker and Kreutzer, 1976; ShSnau, 1988; Monserud et al., 1990; Rayner, 1992). Our study has the originality to show that the Quebec FECP does not provide reliable factual information, which is needed for various management activities at the site scale.

5. General discussion

The results expressed in this paper tend to demon- strate that ecological classification is difficult in disturbed deciduous forests. These difficulties may decrease the potential of the products derived from that classification. This begs the question: Is ecologi- cal classification nevertheless of interest for these forests? Before dismissing ecological classification as a tool for forest management, one must consider the various aspects of ecological classification. First, we must recall that our analyses were done without shrub and herb species which have been used for a long time in forest management (Cajander, 1926; Rowe, 1956; Becker, 1972). Using understory species only, Meilleur et al. (1992) showed, in the same area as this study, that sociological groups are related to many environmental variables. However, the groups could not be used to identify all the morphogenic features and drainage classes. Second, ecological classification is unfortunately often seen strictly as the understanding of the relationships between soil and vegetation distribution. Even though soils cannot explain tree species distribution, they can still be useful for forest interpretations. In Quebec, Bourque (1989) and Cartier et al. (1994) have elaborated very interesting forest interpretations largely based on soils and other physical variables. Thus, two other major roles of ecological classification programs are the description of soil types in terms of characteristics that are important for forest management, and the elaboration of research studies designed to seek the knowledge required for the development of forest interpretations. Third, limitations of ecological clas- sification as a tool for forest management at the site scale do not necessarily restrict its potential at the regional scale (Nolet, 1993). Furthermore, many au- thors have outlined that broadscale ecological classi- fication has an interesting potential for various val- ues such as wildlife management (Kikkawa and Webb, 1976; Davey, 19891, urban planning (Steiner, 1991) and ecosytem ‘health’ conservation (Kimmins, 1990). The reader is referred to Have1 (1980) for a complete literature review of the various uses of ecological classification.

Many authors have written about the importance of the choice of the mapping scale (Mueller-Dombois and Ellenberg, 1974; Bailey, 1985). In the Quebec

96 P. Nolet et al. /Forest Ecology and Management 78 (1995) 85-9X

FECP, the scale chosen was intended to serve both regional and site planning. This paper demonstrates that this choice has produced significant shortcom- ings in site scale mapping. In this context, we pro- pose important changes to the Quebec FECP map- ping. For regional planning, 1:20000 ecosystems maps are too detailed; maps at a scale of 1:50000 would be sufficient. For site planning, the optimal scale is hard to determine because it depends on the spatial variability of soils and vegetation. In this context, we propose a system in which a forest owner would ask for an ecological mapping adapted to his needs. This system would respond to the criticism often made that ecological mapping is not adapted to users’ needs (Domon et al., 1989; BClan- ger et al., 1990). It would also allow a good descrip- tion of vegetation characteristics that are variable in nature, such as advance regeneration and stem qual- ity, but very important for silvicultural treatments.

The analysis of ecological classification presented in this paper is not only applicable to the disturbed deciduous forests. First, in a context where pressure on the natural resources is ever increasing, we can assume that the effects of anthropic activities on the natural forested ecosystems will increase as well. In Quebec, it is possible that the relationships between soils and vegetation will be greatly affected by the repeated mechanical logging in the boreal and mixed conifer-hardwood forests. This, in turn, may render the ecological classification of these forests more difficult as has been observed for the deciduous forests. Thus, two of the solutions for ecological classification mentioned in this paper (emphasizing the description of soils in terms of characteristics important for forest interpretations rather than rela- tionships between soils and vegetation; conducting studies to refine these interpretations) are pertinent not only to disturbed deciduous forests but also to disturbed forests in general as well as to natural forested ecosystems that are planned to be harvested. Second, the methodology used in the second part of this study, i.e. the comparison of information re- quired for forest management with the information provided by the FECP, is a logical way to ensure that a forest ecological classification program gives the information desired. Consequently, it may constitute an interesting framework for other similar programs to be developed in other jurisdictions.

Ackmwledgn~ents

We are grateful to the research group ‘Haut-St- Laurent, Ecologic et Amenagement’, and especially to Alain Meilleur, for letting us use their data from the study area. We also want to thank Alain Leduc for his advice on data analysis, Deborah Payne for editing and three anonymous reviewers for their helpful comments. This research was mainly funded by the Quebec Ministry of Education (FCAR) and also received financial support from the Social Sci-, ences and Humanities Research Council of Canada (CR%).

References

Anonymous, 1988, R&&r ma For&t, Guide d’Aminagement des For&s Priv6es. Service canadien des for& Gouvernement du Canada, 133 pp.

Anonymous, 1992. Programme d’Aide a la Mise enValeur des For&s Privtes, Cahier d’brstructions, Normes Techniques. Service de Mise en Valeur des Forgts Priv6es, Ministare des For&s du Qu&rec, 190 pp.

Archambault, L., Barnes B.V. and Witter J.A., 1989. EcoIogicat species groups of oak ecosystems of southeastern Michigan. For. Sci., 35: 1058-1074.

Avers, P.E. and Schlatterer, E.F., 1991. Ecosystem classification and mapping on National Forests. Presented at the Symposium on Systems Analysis in Forest Resources 3-7 March 1991, Charleston, SC.

Bailey, R.G., 1985. The factor of scale in ecosystem mapping. Environ. Manage., 9: 271-276.

Bariteau, L., 1988. La cartographic g&morphologi.que au 1:2Oooi) de model&s polyg&riques: un exemple des basses terres du Saint-Laurent. Memoire de Maitrise, Universite de Montreal, 185 pp.

Barnes, B.V., 1986. Varieties of experience in classifying and mapping forestland classification in relation to forest manage- ment. Symposium Proceedings, Sault-Ste.-Marie, Ontario, 27- 29 August 1985, pp. 5-23.

Becker, M., 1972. Etude dea liaisons station-production+ dans une for& sur sols hydromorphes. Rev. For. Fr., 24: 269-287.

B&nd, M., Bergeron, Y., Harvey, B. and Robert, D., 1992. Quebec’s ecological framework for forest management: a case study in the boreal forest of Abitibi. For. Ecol. Manage., 49: 247-266.

B&anger, L., Gag& R. and Pineau, M., 1990. L’utihsation des donnees ecologiques dans la pratique forest&e au Quebec. For. Chron., 8: 247-266.

Bergeron, Y., Bouchard, A. and Leduc, A., 1988. Les successions secondaires darts les for&s du Haut-St-Laurent, C&r&b&. Nat. Can., 115: 19-38.

P. Nolet et al. /Forest Ecology and Management 78 (I 99.5) 85-98 97

Bergeron, J.-F., Saucier J.-P., Robitaille, A. and Robert, D., 1992.

Qu6bec forest ecological classification program. For. Chron., 68: 53-63.

Bergeron, J.-F., Saucier, J.P., Robitaille, A., Grondin, P. and

Robert, D., 1993. The Qu6bec Forest Ecological Survey Pro- gram, a set of tools for sustainable forest management. Con- ference presented at the Symposium ‘Advancement in Forest

Inventory and Forest Management Sciences’, September 1993, Seoul, South Korea.

Bouchard, A., Bergeron, Y., Cam%, C., Gangloff, P. and Garitpry,

M., 1985. Proposition d’une m&hodologie d’inventaire et de cartographic tcologique: le cas de la MRC du Haut-Saint- Laurent. Cah. Gtogr. Q&b., 29: 79-95.

Bouchard, A., Dyrda, S., Bergeron, Y. and Meilleur, A., 1989. The use of notary deeds to estimate the changes in the composition of 19th century forests in Haut-St-Laurent,

Qutbec. Can. J. For. Res., 19: 1146-1150. Bourque, J.F., 1989. Cadre Ccologique de &f&em% du territoire

de Girardville: quelques interpr&ations pour l’amtnagement

forestier. Planification Fcologique, Contribution de la Car- tographie Ecologique, No. 39. Direction du Patrimoine Ecol- ogique, Division de la Cartographie Ecologique, Ministbre de

l’Enviromrement, 150 pp. Bowling, C. and Zelazny, V., 1992. Forest site classification in

New-Brunswick. For. Chron., 68: 34-41.

Brenner, R.N. and Jordan, J.K., 1991. The role of an ecological classification system in forest plan development and imple- mentation. Presented at the Symposium on Systems Analysis

in Forest Resources March 3-7 1991, Charleston, SC. Brisson, J., Bergeron, Y. and Bouchard, A., 1988. Les successions

secondaires sur sites mtsiques dans le Haut-St-Laurent,

Quebec, Canada. Can. J. Bot., 66: 1192-1203. Cajander, A.K., 1926. Thetheory of forest types. Acta For. Fenn.,

29: l-108.

Carmean, W.H., 1975. Forest site quality evaluation in the United States. Adv. Agron., 27: 209-269.

Cartier, P., Harvey, B. and Bergeron, Y., 1994. Classification

6cologique et amtnagement pour la region Ccologique des Basses-Terres-D’Amos. Unpublished Report, Service des In- ventaires Forestiers, Ministere des Ressources Naturelles, 138

PP. Christian, C.S., 1951. Regional land surveys. J. Aust. Agric. Sci.,

17: 140-146.

Corns, G.W., 1992. Forest site classification in Alberta: its evolu- tion and present status. For. Chron., 68: 85-93.

Davey, S.M., 1989. Thoughts towards a forest wildlife manage-

ment strategy. Aust. For., 52: 56-67. Delpech, R., Dumb, G., and Galmiche, P., 1985. Typologie des

Stations Forestieres-Vocabulaire. Ministere de l’Agricul-

ture/Direction des For&s, Institut pour le Developpement Forestier, Paris, 243 pp.

Domon, G., Garidpy, M. and Bouchard, A., 1989. Ecological

cartography and land use planning; trends and perspectives. Geofomm, 20: 69-82.

Domon, G., Bouchard, A. and GariCpy, M., 1993. The dynamics of the forest landscape of Haut-Saint-Laurent (Quebec,

Canada): interactions between biophysical factors, perceptions and policy. Landscape Urban Phum., 25: 53-74.

Gauvin, C. and Bouchard, A., 1983. La vCgCtation forest&e du Pam du Mont-Orford, Qu6bec. Can. J. Bot., 61: 1522-1547.

Havel, J.J., 1980. Application of fundamental synecological knowledge to practical problems in forest management. II. Application. For. Ecol. Manage., 3: 81-111.

Hill, M.O., 1979. TWINSPAN: A fortran program for arranging

multivariate data in an ordered two-way table by classification of the individuals and the attributes. Cornell’s Ecology Pro- gram Series, Ithaca, New York, 52 pp.

Jurdant, M., BClair, J.L., Ducruc J.-P. and Gerardin, V., 1977.

L’inventaire du capital-nature, mtthode de classification et de cartographic Ccologique du territoire (36me approximation).

Direction G6&rale des Terres, Enviromrement Canada, SCrie de la Classification Ecologique du Territoire No. 2, 202 pp.

Kikkawa, J. and Webb, L.J., 1976. Identification and classification

of wildlife habitats. In: Proceedings XVIth IUFRO World Congress, Norway 1976. Proc. Div. 1, pp. 744-762.

Kimmins, J.P., 1990. Monitoring the condition of the Canadian

forest environment: the relevance of the concept of ‘ecological indicators’. Environ. Monit. Assess., 15: 231-240.

Lamontagne, L., Cami&, C. and Ansseau, C.. 1991. La vhgttation

forest&e du delta de Lanoraie, Qutbec. Can. J. Bot., 69: 1839-1852.

Lechowicz, M. and Bell, G., 1991. The ecology and genetics of fitness in forest plants. II. Microspatial heterogeneity of the edaphic environment. J. Ecol., 79: 687-696.

Leduc, A., Drapeau, P., Bergeron, Y. and Legendre, P., 1992. Study of spatial components of forest cover using partial Mantel tests and path analysis. J. Veg. Sci., 3: 69-78.

Legendre, L. and Legendre, P., 1984. Ecologic NumCrique. 2. La Structure des Don&es; Verne Bdn. Presses de 1’UniversitC du QuCbec, Quebec, 335 pp.

Legendre, P. and Vaudor, A., 1991. Le Progiciel R. Analyse Multidimensionnelle, Analyse Spatiale. UniversitC de Mont&l, 144 pp,

Mackinnon, A., Meidinger, D. and Klinka, K., 1992. Use of the biogeoclimatic ecosystem classification system in British Columbia. For. Chron., 68: 100-120.

Meades, W.J. and Roberts, B.A., 1992. A review of forest site

classification activities in Newfoundland and Labrador. For. Chron., 68: 25-33.

Meilleur, A., 1986. Etude des communaut6s v&g&es du secteur Cazaville de la M.R.C. du Haut-Saint-Laurent, Qu6bec. MCmoire de Maitrise, Universid de Montreal, 116 pp.

Meilleur, A., Bouchard, A. and Bergeron, Y., 1992. The use of understory species as indicators of landform ecosystem type in heavily disturbed forest: an evaluation in the Haut-St-Laurent,

QuCbec. Vegetatio, 102: 13-32. Meilleur, A., Bouchard, A. and Bergeron, Y., 1994. The relation

between geomorphology and forest community types of the

Haut-St-Laurent, Qu6bec. Vegetatio, 111: 173-192. Monserud, R.A., Moody, U. and Breuer, D., 1990. A soil-site

study for inland Douglas-fir. Can. J. For. Res., 20: 686-695.

98 P. Nolet et al. /Forest Ecology and Management 78 (1995) 85-98

Mueller-Dombois, D. and Ellenberg, H., 1974. Aims and Methods of Vegetation Ecology. Wiley, New York, 547 pp.

Nolet, P., 1993. Potentiel et limites d’utilisation du cadre icol- ogique forestier quebfcois en for& privee. Memoire de Maitrise Present6 comme Exigence Partielle de la Maitrise en Sciences de l’Environnement, Universiti du Quebec ic Montreal, 126

PP. Pfister, R.D., 1977. Ecological classification of forest land in

Idaho and Montana. Reprinted from 1977 Proceedings Ecolog-

ical Classification of Forest Land in Canada and Northwestern USA, University of British Columbia, Vancouver, pp. 329-

358. Rayner, M.E., 1992. Evaluation of six site classification for

modelling timber yield of regrowth karri (Eucalyptus diuersi- color F. MueIl.1. For. Ecol. Manage., 54: 315-336.

Robitaille, A., 1988. Cartographie des Districts Ecologiques:

Normes et Techniques. Ministbre de 1’Energie et des Ressources, Service de 1’Inventaire Forestier, Division Ecolo- gie, 109 pp.

Rogister, J. and Galoux, A., 1982. Forestry and ecological map- ping in Belgium. In: G. Jahn (Editor), Handbook of Vegetation Science, Part 12. pp. 117-146.

Rousseau, C., 1974. Gtographie Floristique du Quebec/Labrador. Distribution des Principales Esp&es Vasculaires. Les Presses de L’Universite Laval, Quebec.

Rowe, J.S., 1956. Uses of undergrowth plant species in forestry. Ecology, 37: 461-473.

Rowe, J.S., 1972. Les Regions Forest&es du Canada. Ministbre

de 1’Environnement. Service Canadien des For&, Publication No. 13OOF, 172 pp.

Saucier, J.-P., Berger, J.-P. and Robitaille, A., Undated. Le Point d’Observation Ecologique. Service des Inventaires Forestiers, Ministere des For&s du Quebec, Quebec, 104 pp.

Schlenker, G. and Kreutxer, K., 1976. Vergleich von Klassifka-

tionsystemen fur forstliche Standortskartierungen. In: Proceed- ings XVlth IUFRO World Congress, Norway 1976. Proc. Div. 1, pp. 219-238.

Shiinau, A.P.G., 1988. Problems in using vegetation or soil classi-

fication in determining forest site quality. In: D.W. Cole and S.P. Gessel (Editors), Forest Site Evaluation and Long-term Productivity, University of Washington Press, Seattle, pp. 3-11.

Sims, R.A. and Uhlig, P., 1992. The current status of forest site classification in Ontario. For. C&on., 68: 64-77.

Sims, R.A., Towill, W.D. and Wickware, G.M., 1986. A status report on a forest ecosystem classification @EC) program for Ontario’s North Central Region. In: G.M. Wickware and W.C.

Stevens (Editors), Site Classification in Relation to Forest Management. COJFRC Symposium Proceedings, Canadian

Forestry Service, Great Lakes Forestry Centre, Canadian Forestry Service, Government of Canada, Sault-Ste-Marie, Ontario, O-P-14, pp. 83-87.

Steiner, F., 1991. Landscape planning: a method applied to growth management plan. Environ. Manage., 15: ~519-S29.

Stellar, R., 1988. The History of Huntingdon and the Seignioties

of Beauhamois and Chateauguay. Huntingdon Gleaner Inc.. Huntingdon, Quebec, Canada.

Thibault, M. and Hotte, D., 1985. Les Regions Ecologiques du

Quebec Mtridionnal (map) (2nd approximation). Ministere de I’Energie et des Ressources du Quebec, Quebec. Direction de la Recherche et du Diveloppement.

Wells, E., 1992. User information needs for forest management site classification in Manitoba. For. Chron.. 68: 78-84.