Vegetatio 118: 131-137, 1995. © 1995 Kluwer Academic Publishers. Printed in Belgium. Canadian wetlands: Environmental gradients and classification S.c. Zoltai 1 & D.H. Vitt2 I Forestry Canada, Northern Forestry Centre, Edmonton, Alberta, Canada T6H 3S5 Botanic Garden, The University of Alberta, Edmonton, Alberta, Canada T6G 2El 2 Devonian Key words: Bogs, Ecological gradients, Fens, Marshes, Peatlands, Shallow lakes, Swamps, tion, Water chemistry Vegetation-Class!fi§f~'-: Abstract The Canadian Wetland Classification System is based on manifestations of ecological processes in natural wetland ecosystems. It is hierarchical in structure and designed to allow identification at the broadest levels (class, form, type) by non-experts in different disciplines. The various levels are based on broad physiognomy and hydrology (classes); surface morphology (forms); and vegetation physiognomy (types). For more detailed studies, appropriate characterization and subdivisions can be applied. For ecological studies the wetlands can be further characterized by their chemical environment, each with distinctive indicator species, acidity, alkalinity, and base cation content. For peatlands, both chemical and vegetational differences indicate that the primary division should be acidic, Sphagnum-dominated bogs and poor fens on one hand and circumneutral to alkaline, brown moss-dominated rich fens on the other. Non peat-forming wetlands (marshes, swamps) lack the well developed bryophyte ground layer of the fens and bogs, and are subject to severe seasonal water level fluctuations. The Canadian Wetland Classification System has been successfully used in Arctic, Subarctic, Boreal and Temperate regions of Canada. Wetland classification Classification is a process of organizing information by forming groups within which the individual elements are similar in some respects, but are distinctly different from the elements of the other groups. However, the purpose of classification must be clearly defined, as the same objects, in this case wetlands, can be grouped differently, depending on the needs of the intended user. The perceptions of a wildlife or waterfowl biologist, an agronomist, a peat producer, a hydrologist, or an engineer can be vastly different when considering the same wetland. It was natural, therefore, that a number of wetland classification systems have been developed in Canada by different interest groups. Some ecologists developed a phytosociological classification of peatland vegetation (cf. Dansereau & Segadas-Vianna 1952; Gauthier & Grandtner 1975). Meanwhile, the mire classification developed in Europe (Osvald 1925) has been used to include vegetation indicators and environmental gradients in Canada (cf. Sjors 1952, 1969; Vitt et at. 1975; Gauthier 1980; Damman 1986). The same principles were incorporated into a classification of Ontario wetlands (Jeglum et al. 1974). A classification of the wetlands of the Canadian prairies was developed (Millar 1976) primarily to fulfil the needs of waterfowl ecologists. An organic soil classification system was developed by the Canada Soil Survey Committee (1978), characterizing the peatlands by their soil properties. Yet another classification system was developed to serve the engineering community (Radforth 1969), based on various surface morphology and peat structure features. This variety of classification approaches resulted in restricted communication between the various user groups. The Canadian Wetland Classification System (CWCS) has been developed to facilitate communication between different disciplines concerned about this segment of the natural landscape (Tarnocai 1988). This classification system is based on categorizing important ecosystem processes, such as water budget (amount and seasonality), carbon budget (peat 132 accumulation), and environmental parameters, such as water quality (dissolved solids) and quantity, which interact to allow the development of distinctive ecosystems. Such a classification is synthetic and was chosen because this places the emphasis on environmental processes that determine the development of different types of wetlands. The genesis and dynamics of wetlands are ill...·!" --t at the highest levels of the classification by grouph.o similar wetland functions. This aspect is v:ery ~mportant for all wetlands, but especially for pea,tlands that literally create their own environment ': oveFalOiigperiod of time. The classification system has a hierarchical structure where the various categories represent different levels of complexity. The higher levels have been assigned multispectral (ecosystemic) definitive and descriptive criteria, the lower levels having more specific criteria (Zoltai et al. 1975). The three higher categories have readily recognizable definitive criteria which allow their use by non-specialists. The formal classification ends at a level which is fairly specific, but still too generalized for specialized purposes. Specialists are encouraged to develop their classification that is compatible with the ewcs, based on their interests: ecosystems, floristics, hydrology, soils, engineering qualities, etc. The ewcs provides a common platform that allows the exchange of data and results between different disciplines, using a common language. The CWCS differs in its basic philosophy from the classification system developed for the US Fish and Wildlife Service (Cowardin et at. 1979). The CWCS is based chiefly on wetland functions: interrelationships of the biotic and abiotic components of the wetland ecosystems. The resulting units have implications of the hydrology, water quality, climate, vegetation interactions and chronology (genesis) of the particUlar wetlands. The USFWS system, on the other hand, is an objective approach that relies largely on observable features which require few process oriented decisions. It is designed to accommodate a wide range of environments, from deep waters to bogs and from rocky substrates to peatlands. This system answers the questions of what and where, but does not provide a framework for understanding the rationale of wetland development. In the ewcs a wetland is defined as land that is saturated with water long enough to promote wetland or aquatic processes as indicated by poorly drained soils, hydrophytic vegetation, and various kinds of biological activity that are adapted to a wet environment (Tarnocai et al. 1988). Wetlands include organic MESOTROPHIC Fig. 1. The five classes of the Canadian Wetland Classification System in relation to the important chemical gradients. The vertical axis is a qualitative expression of the influence of water depth. wetlands (peatlands) and mineral wetlands (generally non-peaty). The broadest category of wetlands, wetland classes are recognized on the basis of overall genetic origin of the wetland ecosystem. Five wetland classes have been recognized: bogs, fens, swamps, marshes and shallow open waters (Fig. 1). They are defined and described on the basis of distinctive abiotic parameters such as hydrologic regime, water chemistry, or mineral material, which interact with the biota to form characteristic vegetation cover, and in some classes, peat. In the next lower category are the wetland forms, which are based on surface morphology and pattern, water quality, relationship to open water, and morphology of the underlying mineral soil, as expressed by the ecosystems that are established on the wetlands. This is an open-ended category, with new wetland forms being described and defined as needed. The third category, wetland types, is based on vegetation physiognomy. The wetland is then named to identify all three categories, for example: a wooded (type) raised (form) bog (class); or shrubby (type) patterned poor (form) fen (class). Wetland characteristics Quantitative and qualitative differences in ecological processes, operating within given climatic conditions, are primarily responsible for differences in wetlands. Whereas it is often difficult to measure ecological processes themselves, it is relatively easy to determine the manifestations of these processes in terms of ecosystem characteristics. These characteristics in turn can 133 be used to define a synthetic classification system.- The classification itself serves to reflect the fundamental ecological processes. The classification of wetlands should be based on the factors that control wetland processes. Most of these factors are present as gradients, and these are often non-linear in form. Although these gradients are often correlated with one another, they do not necessarily vary at similar rates or quantities. Despite these complications, wetlands can be grouped into fundamental types that reflect basic ecological processes (Figures I, 2). The Canadian System recognizes five classes of wetlands that form nodes along several complex hydrological, chemical, and biotic gradients. Shallow open waters are wetlands that exist whenever the water levels are sufficient to create habitats for aquatic and floating vegetation. Although seasonal fluctuations in water level may expose the bottom substrate, aquatic processes characterize this wetland. Chemistry of the water does not differentiate this wetland class from the remaining four (Fig. 1), but may be important in determining the type of shallow water vegetation that is present. These wetlands form a transition to truly aquatic ecosystems and are largely influenced by the adjoining aquatic system (Vitt & Slack 1975). Marshes are treeless wetlands that are subject to relatively large seasonal water level fluctuations. These wetlands are strongly influenced by either ground water or surface water and have relatively high amounts of water flow. As a result, available nutrients (here considered only as nitrogen and phosphorus) are abundant and vascular plant production is high, however decomposition rates are also high. Bryophytes are nearly lacking, owing to their inability to compete with vascular plants under eutrophic conditions and when fluctuating water levels are present. The influence of surface and ground waters generally serves to increase base cations and HCO:;- , thus creating a well buffered wetland. Trees are lacking due to either regional climatic conditions and/or the seasonally wet conditions. As a result, marshes are wetlands that reflect eutrophic, temporally variable conditions that prohibit a well developed bryophyte layer, largely due to high vascular plant production. Rapid decomposition, along with the poorly developed ground layer, inhibits significant peat accumulation. Chemical differences greatly influence the vegetation composition of marshes. Freshwater marshes are dominated by calcium and bicarbonate (and are alkaline systems), while saline marshes have sodium and sulphate as their dominant ions (and are thus saline and non-alkaline). Tidal marshes, with very different vegetation, are dominated by sodium and chloride ions (and thus are also saline and non-alkaline wetlands) (Fig. 2). Swamps have many similarities to marshes in that swamps have strong seasonal water level fluctuations and relatively strong water flow. Like marshes, the bryophyte ground layer is poorly developed or lacking in most swamps. Both production and decomposition is generally high, and the seasonal lowering of the water level permits the establishment of a welldeveloped tree or shrub layer. When peat is produced in some quantity, it is well decomposed; however, in general, peat accumulation is small. Tree development often inhibits water flow and some swamps can become acidic, especially in areas of acidic ground water. These acidic treed wetlands rarely develop a bryophyte dominated ground layer due to the fluctuating water levels, but stable water levels under dense conifer cover allow the formation of a moss carpet. Peat accumulation is small owing to rapid decomposition. Vegetationally, swamps can be quite diverse, from deciduous alderash, to evergreen white cedar, to densely forested black spruce, to cypress swamps farther south. All of these wetlands, however, are characterized by the presence of trees or tall shrubs, generally little peat accumulation and extreme seasonal water level fluctuation. Peatlands are distinct from non-peat forming wetlands by a combination of hydrological and biotic factors that function together to create conditions suitable for decreased plant production and decreased decomposition. The stability of seasonal water levels restricts total water flow through a wetland and allows a ground layer of bryophytes to develop. Available nutrient levels are low, as nutrients are accumulated in nonavailable forms by this ground layer. As a result, production of the vascular plant component is reduced, but production of the ground layer increases due to active nutrient sequestering by the bryophyte layer (Bayley et at. 1987). Stable water levels and decreased nutrient availability lead to a decrease in decomposition and peat accumulation increases. Fens are always bryophyte dominated wetlands that are influenced by the chemistry of the surrounding mineral soil deposits. Fens having bicarbonate (thUS they are alkaline) as their dominant anion and calcium as their dominant cation are rich fens. These wetlands are characterized by brown mosses largely of the 134 OLIGOTROPHIC MESOTROPHIC EUTROPHIC TOTAL NUTRIENT AVAILABILITY -+ PRODUCTION -+ DECOMPOSITION ~ DTreed c:?J Sphagnum Q Brown Mosses Fig. 2. The relationship of bog, fen, marsh, and swamp wetland classes to the major chemical, biotic, and hydrologic gradients. Fens and marshes are divided into several wetland forms in order to illustrate the pattern of variability of these classes. Saline wetlands may be either fens or marshes. family Amblystegiaceae, and an abundance of sedges. Extreme-rich fens have pH above 7.0 (and are basic) and often have deposits of marl in their wetter areas, while moderate-rich fens have pH above 5.5 and below 7.0 (and are acid). These rich fens are strongly influenced by water from surrounding uplands and from the larger watershed, and generally have sufficient flow to allow the wetlands to be mesotrophic. Although data are scarce, it appears that production of the ground layer in rich fens is similar to that in poor fens and bogs (Vitt 1990), while production of the vascular plant component may be greater than in these latter wetlands due to somewhat greater decomposition allowing more nutrients to be made available. Poor fens (and bogs) are acidic, non-alkaline wetlands that are dominated by the genus Sphagnum. Sphagnum, through its abilities to acidify and hold large quantities of water, restricts both water flow and nutrient availability, which in turn decrease vascular plant production and decomposition. Poor fens, as rich fens, are influenced by geogenous water. The presence of a well developed ground layer 'of Sphagnum serves to stabilize the wetland and increase the acidity. From both vegetational and chemical viewpoints, poor fens are more similar to bogs than they are to rich fens (MaImer et al. 1992); however, the hydrology of poor fens has greater similarities to that of rich fens. Bogs are Sphagnum-dominated, ombrotrophic wetlands. They are acidic, largely due to humic acid production during decomposition processes (Hemond 1980) and to cation exchange due to uronic acid production from Sphagnum, and non-alkaline. The base cation content is limited and ombrotrophy greatly restricts water flow. Nutrients are largely tied up in Sphagnum and accumulated peat and mineralization processes are reduced. Thus the rate of production in the ground layer remains relatively similar to that of fens seemingly due to internal cycling of nutrients within the Sphagnum plants, while vascular plant production is limited. Wetlands in Canada form distinct types along complex gradients. Especially important are: (1) Hydrology (in particular, seasonal water level fluctuation and amount of water flow through wetlands); (2) Chemistry (in particular nutrient [N, P] availability, acidity [hydrogen ion content], alkalinity [bicarbonate content], and base cation content rCa, Mg, Na, K]). Nutrient availability leads to classification of wetlands as oligotrophic, mesotrophic or eutrophic; and (3) Biotic 135 (in particular the development of a ground layer dominated by either brown mosses or Sphagnum and the presence of a tree layer). Proper classification of wetlands will lead to proper management and any classification system must recognize the complex interrelationships between the gradients formed from these controlling hydrological, chemical, and biotic factors. The ewcs has been used as the framework for evaluating wetlands for conservation or use in Canada (Bond et at. 1992). The needs of particular disciplines can be accommodated by combining the CWCS with parameters that are important to that discipline. It is evident from wetlands of west-central Canada that data sets of these characteristics can be used to formulate wetland sites into an ecologically relevant classification (Tables 1 and 2). Inventorying wetlands It must be clearly understood that wetland classification and the inventorying of wetlands are two distinct and separate operations. Although the wetland classification should be used as an inventory and mapping criterion, the purpose of the inventory, and the desired detail and scale of the inventory map will dictate the kinds of wetlands that will be recognized. Therefore, a universal, global inventory of wetlands is possible only if these parameters are determined in advance. This is not an easy task, as circumstances vary greatly around globe. In some countries with limited wetland resources, the few wetlands may be very well documented, whilst in other countries, such as Canada, vast tracts of wetlands are virtually untouched and never studied. The wetlands of Canada have not been accurately inventoried. A small scale map was generated (National Wetlands Working Group (1986) from available published and unpublished information. In recent years, however, a number of wetland inventories were initiated by various provinces that provide more accurate information. The mapping units are generally compatible with or were inspired by the CWCS. The wetland resources of New Brunswick were inventoried at a scale of 1:50 000 (Airphoto Analysis Associates Consultants Ltd. 1975). The Ontario portion of the Hudson Bay Lowlands were mapped at a scale of 1:250 000 (Pala & Boissonneau 1982). The peatlands of southern Quebec have been mapped and an atlas published (Buteau 1989) at a scale of 1:250 000; the inventorying of the wetlands of northern Quebec is under way. The wetlands of Alberta (with the exclusion of southern parts of the province) have been mapped at 1:250 000 scale, and a summary map was published at a scale of 1:1000000 (Vitt 1992). The mapping of the wetlands of Manitoba is currently under way. In addition to wetland inventory maps, the surficial geology maps of the northern Mackenzie River valley show the extent of fens and bogs at a scale of 1:250000 (Duk-Rodkin & Hughes 1992). In our experience, technology must be combined with field work in order to obtain reliable data on the extent and kind of wetlands. Remotely sensed data in the form of aerial photographs or spectral data (passive or active systems) are necessary to cover areas of poor access. Although computer-enhanced remotely sensed information shows promise (Pala & Boissonneau 1982; Anon 1989), the evaluation and validation of this information is necessary before confidence can be established. This rapidly developing technology, however, dictates that this option must be monitored for possible future use. The use of aerial photos is a well-established and tried method of wetland inventory (Buteau 1989; Vitt 1992). Wetland types can be readily identified to a detail limited only by the scale of the photos. The thickness of the peat cannot be determined from air photos with confidence, although relationships between peat depth and topography can be established locally within climatic regions. Ground truthing is necessary to determine the vegetation types, cation and nutrient levels, in addition to peat thickness and stratigraphy measurements. Ideally, a multi-tiered approach should be applied: (1) field work limited to a small portion of the wetlands; (2) expansion of this knowledge to large areas through air photo interpretation; (3) extension of the results of air photo study to a regional scale through interpretation of remotely sensed information. We feel confident that the CWCS can be applied to Boreal, Subarctic, and Arctic regions of the Northern Hemisphere. The definition of the wetland classes is compatible with that of northern European wetland ecologists. The wetland forms are especially useful for air photo interpretation, and new forms can be defined, as needed. The broad vegetation physiognomy, used to identify wetland types, should not pose any diffiCUlty. However, we lack the necessary experience to judge the relevance of CWCS to wetlands of lower latitudes. 136 Table 1. Variability in surface water chemistry in the wetland classes from a small region (Elk Island National Park) of central Alberta, Canada (Nicholson 1992; n = 331). Bog Fen Swamp Marsh 3.5-3.6 5.2-6.4 pH Reduced Conductivity* (J.LS cm- I ) Ca (mg I-I) 4.0-6.2 5.9-6.1 16-27 40-160 230-330 160-530 (900)** 4-7 2-33 Na (mg I-I) 2-3 2-5 26-43 5-22 27-65 3-125 (800)** Organic N (J.Lg I-I) NO;- (J.LgI- I ) 2900-3000 1350-2850 2000-3000 200-2500(6400)** 13-20 8-23 7-10 9-175 160-250 23-80 28-146 73-130 (6400)** 350-480 135-400 220-650 250-520 (1100)** NHt (J.LgI- I ) P (total)(J.Lg I-I) * Conductivity minus the effect due to hydrogen ions (Sjors 1952) * * ( ) Data from saline marshes. Table 2. Average chemical compOSItion of surface water in various peatlands in west-central Canada (Zoltai & Johnson 1987). All data are from point measurements at mid-summer. Standard error of the mean in brackets. Bogs Poor fens Moderate- Extreme- rich fens rich fens No. sites 71 33 147 18 Depth to water table 37 12 11 5 (1.6) below surface (cm) (1.8) (4.0) (1.2) pH 4.5 (.02) 4.8 (.04) 5.8 (.05) 6.5 Reduced conductivity* (J.LS cm- I ) 62 (3) 53 (7) 212 (12) 374 Ca(mgl- I ) 2.04 (.18) 2.90 (.63) 24.98 (1.59) 53.60 (5.29) Mg(mgl- I ) 0.87 (.12) 1.19 (.32) 10.16 (.69) 14.20 (1.24) Na(mgl- 1) 2.59 3.89 (.28) (.93) 4.75 (.36) 6.54 (.87) 0.17 (.018) 0.14 (.024) 0.13 (.046) 0.12 (.020) 1.42 1.26 (.19) 1.42 (.11) 0.97 (.20) P(mgl- 1) K(mgl- 1) (.10) (.12) (33) * Conductivity minus the effect due to hydrogen ions (Sjors 1952). References Airphoto Analysis Associates Consultants Limited. 1975. Wetlands - peatlands resources, New Brunswick. Prepared for New Brunswick Department of Natural Resources, Fredericton, New Brunswick. Unpublished Report, 188 pp. Anon. 1989. Inventory of peatlands in northern Quebec. 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