Preferred Citation: Warner, Richard E., and Kathleen M. Hendrix, editors California Riparian Systems: Ecology, Conservation, and Productive Management. Berkeley:  University of California Press,  c1984 1984. http://ark.cdlib.org/ark:/13030/ft1c6003wp/


 
Evolution and Riparian Systematics1

Evolution and Riparian Systematics[1]

David E. Brown[2]

Abstract.—Arizona's perennial streams and important marshlands have been mapped and a wetland classification system developed. To be effective and usable, a resource classification must be systematic, universal, and hierarchical, and must illustrate, or at least recognize, evolutionary relationships. Biogeography is therefore an important factor in the development of the taxonomy for any living (i.e., renewable) resource. Few renewable resources are as alive and dynamic as are our riparian ecosystems.

We have mapped Arizona's perennial streams and important wetlands at 1:1,000,000 (Brown etal . 1977, 1978, 1981). These maps show the potential for maximum riparian ecosystem development—various riparian communities are not illustrated perse . Riparian communities are too dynamic to present the same structure and composition for any length of time; similar communities may also occur along seasonal and even some ephemeral subterranean-fed waterways. This does not mean that riparian vegetation cannot be inventoried and mapped for study purposes and to document change.

We have developed a classification system that includes riparian and other wetland communities as well as upland ecosystems (see Appendix A) (Brown and Lowe 1974a, 1974b; Brown etal . 1977, 1979, 1980). Like the Linnean taxonomic system, this classification system is systematic in approach, universal in application, and hierarchical in arrangement. It is also digitized and therefore computer-compatible. Like other ecosystem classifications, this system uses vegetation, structure, climate, and vegetative components as criteria. However, an important distinction is that it is based on biogeography.

A classification based on this system for use in the Southwest has proven serviceable for classification, delineation, description, and data storage of that region's natural vegetation and biogeography. For examples of its application see Turner and Cochran (1975), Steenbergh and Warren (1977), Patton (1978), Martin (1979), Turner etal . (1980), and Volger (1980).

All classifications of vegetation consider structure (i.e., forests, woodlands, grasslands, etc.); the most successful employ phytogeographic descriptions (i.e., floodplain forest, montane forest, coastal scrub, etc.). Unfortunately, many of these same classifications rely on soil and/or chemical criteria which influence floristics only regionally. Soil-types or soil properties are of little use in describing vegetation on a worldwide or even continental basis. Some systems (e.g. Bailey 1976, 1978) use physiographic approaches that are wholly regional in scope and bear little relationship to biotic parameters. Few systems employ biogeography as it is used by biologists.

Biologists have long been developing systems of biographic realms, provinces, and districts (e.g., see Wallace 1876; Clements and Shelford 1939; Rasmussen 1941; Pitelka 1941; Dice 1943; Goldman and Moore 1945; Dansereau 1957; Darlington 1957; Lowe 1961; Shelford 1963; Walter 1973; Udvardy 1975; Cox etal . 1976; Dasmann 1976; Franklin 1977) to show the distribution of plants and animals. These distributions are the result of evolutionary origin and adaptation. The basic biogeographic unit is the biome (i.e., biotic community). The biome is also the primary component and mappable reality of any biotic classification system that attempts to illustrate evolutionarily significant plant and animal distribution. Distributions which are of evolutionary significance are of great importance to bird watchers, ornithologists, zoogeographers, mammalogists, herpetologists, phytogeographers, taxonomists, and wildlife managers. Biologists will generally not accept classifications and inventories that do not recognize the importance of biomes and biogeography. This is especially true of our riparian and other wetland resources, so valued for their biotic diversity.

[1] Paper presented at the California Riparian Systems Conference. [University of California, Davis, September 17–19, 1981].

[2] David E. Brown is Wildlife Biologist, Arizona Game and Fish Department, Phoenix; and Professor of Wildlife Management, Arizona State University, Tempe, Ariz.


289

Literature Cited

Bailey, R.G. 1976. Ecoregions of the United States (map, scale 1:7,500,000). USDA Forest Service, Intermountain Region, Ogden, Utah.

Bailey, R.G. 1978. Description of the ecoregions of the United States. USDA Forest Service, Intermountain Region, Ogden, Utah.

Brown, David E., Neil B. Carmony, and Raymond M. Turner. 1977. Inventory of riparian habitats. p. 10–13. In : R.R. Johnson and D.A. Jones (tech. coord.). Importance, preservation and management of riparian habitat: a symposium. [Tuscon, Ariz., July 9, 1977]. USDA Forest Service GTR-RM-43. 217 p. Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colo.

Brown, David E., Neil B. Carmony, and Raymond M. Turner. 1978. Drainage map of Arizona showing perennial streams and some important wetlands. Ariz. Game and Fish Department map.

Brown, David E., Neil B. Carmony, and Raymond M. Turner. 1981. Drainage map of Arizona showing perennial streams and some important wetlands. Ariz. Game and Fish Department map.

Brown, David E., and C.H. Lowe. 1974a. A digitized computer-compatible classification for natural and potential vegetation in the Southwest with particular reference to Arizona. J. Ariz. Acad. Sci. 9, Suppl. 2:1–11.

Brown, David E., and C.H. Lowe. 1974b. The Arizona system for natural and potential vegetation—illustrated summary through the fifth digit for the North American Southwest. J. Ariz. Acad. Sci. 9, Suppl. 3:1–56.

Brown, David E., C.H. Lowe, and C.P. Pase. 1977. A digitized classification system for the natural vegetation of North America with hierarchical summary for world ecosystems. In : A. Marmelstein (ed.). Proceedings of the national symposium on classification, inventory, and analysis of fish and wildlife habitat. [Phoenix, Ariz., Jan. 24–27, 1977]. USDI Fish and Wildlife Service, Office of Biological Science, Washington, D.C.

Brown, David E., C.H. Lowe, and C.P. Pase. 1979. A digitized classification system for the biotic communities of North America, with community (series) and association examples for the Southwest. J. Ariz.-Nev. Acad. of Sci. Suppl. 1:1–16.

Dansereau, P. 1957. Biogeography. Ronald Press. New York, New York.

Darlington, P.J., Jr. 1957. Zoogeography. John Wiley and Sons. New York, New York.

Dansmann, R.F. 1976. Biogeographical provinces. Co-Evolution Q. Fall:32–35.

Dice, L.R. 1943. The biotic provinces of North America. University of Michigan Press. Ann Arbor, Mich.

Franklin, J.F. 1977. The biosphere reserve program in the United States. Science 195: 262–267.

Goldman, E.A., and R.T. Moore. 1945. The biotic provinces of Mexico. J. Mammal. 26:347–360.

Lowe, C.H. 1961. Biotic communities in the sub-Mongollon region of the inland Southwest. J. Ariz. Acad. Sci. 2:40–49.

Martin, P.S. 1979. A survey of potential natural landmarks, biotic themes, of the Mojave-Sonoran Desert Region. Heritage Conservation and Recreation, U.S. Department of the Interior. 358 p.

Patton, D.R. 1978. Runwild-a storage and retrieval system for wildlife habitat information. USDA Forest Service GTR-RM-51:1–8, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colo.

Pitelka, F.A. 1941. Distribution of birds in relation to major biotic communities. Amer. Midl. Nat. 25:11–137.

Rasmussen, D.I. 1941. Biotic communities of Kaibab Plateau, Arizona. Ecol. Monog. 11:229–275.

Shelford, V.E. 1963. The ecology of North America. University of Illinois Press. Urbana, Ill.

Steenbergh, W.F., and P.L. Warren. 1977. Preliminary ecological investigation of natural community status at Organ Pipe Cactus National Monument. USDI Cooperative National Park Resources Studies Unit, University of Arizona Tech. Rep. No. 3:1–152.

Turner, D.M., and C.L. Cochran, Jr. 1975. Wildlife management unit-37B-pilot planning study. Arizona Game and Fish Department, Fed. Aid Prog. FW-11-R-8, J-1:1–128.

Turner, R.M., L.H. Applegate, P.M. Bergthold, S. Gallizioli, and S.C. Martin. 1980. Range reference areas in Arizona. USDA Forest Service GTR-RM-79:1–34. Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colo.


290

Udvardy, M.D.F. 1975. A classification of the biogeographical provinces of the world. Internat. Union Conserv. Nature and Natural Resources (IUCN, Morges, Switzerland). Occas. Pap. 18:1–48.

Vogler, L.E. 1980. The Arizona State Museum archaeological site survey system. Ariz. State Mus. Arch. Ser. 128:1–190.

Wallace, A.R. 1876. The geographical distribution of animals, with a study of the relations of living and extinct fauna and as elucidating the past changes of the earth's surface. MacMillan and Co. London, England.

Walter, H. 1973. Vegetation of the earth in relation to climate and the ecophysiological conditions. Translated from the Second German edition by Joy Wieser. English University Press, London; Springer-Verlag, New York, New York.


291

Appendix A—
A Digitized Classification System for the Biotic Communities of North America, with Community (Series) and Association Examples for the Southwest[1]

David E. Brown, Arizona Game & Fish Department, Phoenix Charles H. Lowe, University of Arizona, Tucson Charles P. Pase, USDA Forest Service

Introduction

In previous publications on the North American Southwest System we have addressed primarily the North American Southwest region as outlined in Fig. 1 (Brown and Lowe 1973, 1974a,b). Responses to both the classification system and the classification have been favorable in both general interest and use: e.g., Lacey, Ogden, and Foster 1975; Turner and Cochran 1975; Carr 1977; Dick-Peddie and Hubbard 1977; Ellis etal . 1977; Glinski 1977; Hubbard 1977; Pase and Layser 1977; Steenbergh and Warren 1977; Patton 1978; BLM 1978a,b; Turner etal . 1979. In this report we expand the classification nomenclature at digit levels 1–4 to represent the North American continent

The Southwest System is evolutionary in basis and hierarchical in structure. It is a natural biological system rather than primarily a geography-based one in the sense of Dice 1943; Bailey 1978; and others. The resulting classifications are, therefore, natural hierarchies.

Because of the open-ended characteristic of a natural hierarchical system, resulting classification provides for orderly change. The inherent accordion-type flexibility provides for expansion and contraction at all levels. It permits accommodation of new information into the classification—addition, transference, and deletion of both (a) ecological taxa, and (b) quantitative data on ecological parameters concerning taxa, as our knowledge accumulates on either or both. Digit levels 7 to n accommodate the latter and digit levels 1–6 accommodate the former (ecological taxa) on a world-wide basis.

The system's potential is the provision of a truly representative picture for biotic environment. It permits but does not require inclusion of any and all biotic criteria in a given classification—animals as well as plants. Thereby included in the system's uses are the mapping of wildlife habitats and the determination and delineation of natural areas on a local to world-wide basis (Brown, Lowe, and Pase 1977). On a local basis, overlapping soil mapping units can provide "habitat-types" with their implied biotic potential for land use planning purposes.

The digitation of hierarchy makes the system computer-compatible; e.g., a system or subsystem for storing and retrieving biotic resource data within or parallel to an overall management system. The Southwest System is currently in use in the RUNWILD program developed for field unit use on remote terminals by Region 3 of the Rocky Mountain Forest and Range Experiment Station, U.S. Forest Service (Patton 1978). The system and classification is similarly incorporated in the State of Arizona Resources Inventory System  (ARIS). It is currently used by both industry and agencies for biological studies, resource inventories, and procedures for environmental analysis, for example as required by the National Environmental Policy Act.

The system is responsive to scale. The hierarchical sequence permits mapping at any scale, and various levels of the system have been mapped at 1:1,000,000 (1 inch represents ca. 16 miles). 1:500,000, 1:250,000, 1:62,500 (1 inch represents ca. 1 mile), and others. Moreover, the use of hierarchical sequence permits the needed flexibility for mapping those complex communities where more intensive levels are impractical or needlessly time consuming in a given investigation.

The classification has been expanded to include the major biotic communities of North America (Brown, Lowe, and Pase 1977, 1979). To facilitate communication with potential users, we provide, in addition to some structural modification of the original classification, a number of additional definitions and explanations. Our fourth level (biome) examples for North America are representative; they are not intended as either a definitive or final classification. Examples of the use of the system to the fifth (series = community) and sixth (association) levels are given here for those biomes located wholly or partially within the North American Southwest.

Incorporated in the present classification are contributions from approximately one hundred investigators, primarily biogeographers, wildlife biologists, and ecologists, all of which pertain to or are in general use in the Southwest today. Additional references are given in Brown and Lowe 1974a,b, 1977.

A Digitized Hierarchy of the World's Natural Ecosystems

Where:
1,000 = Biogeographic (Continental) Realm
   1,100 = Vegetation
      1,110 = Formation-type
         1,111 = Climatic (Thermal) Zone
            1,111.1 = Regional Formation (Biome)
               1,111.11 = Series (Community of generic dominants)
                  1,111.111 = Association (Community of specific dominants)
                     1,111.1111 = Composition-structure-phase

A number preceeding the comma (e.g., 1 ,000) refers to the world's biogeographic realms (see Table 1). Origin and evolutionary history are recognized as primary in importance in the determination and classification of natural ecosystems. The mapable reality of the world's biogeographic realms is interpretive in part and dependent on criteria used. In those regions where the components of one realm merge gradually with those of another and the assignment of biogeographic origin is difficult, we include such transitional areas (wide ecotones) in both realms. The following seven realms are adapted from Wallace 1876; see also Hesse etal . 1937; Dansereau 1957; Darlington 1957; Walter 1973; I.U.C.N. 1974; DeLaubenfels 1975; Cox et al . 1976:

[1] A contribution of the Arizona Game and Fish Department with (publication funded by Federal Aid Project W-53R). The University of Arizona Department of Ecology and Evolutionary Biology, and the United States Forest Service, Rocky Mountain Forest and Range Experiment Station.


292

figure

Figure 1.
The Southwest. In delineating a  natural  Southwest region, approximately one half of the area falls in the
Republic of Mexico and one half in the United States; the U.S. states of "Arizona and New Mexico"
constitute less than half of the "American Southwest." Parts or all of the following states are
included: Arizona, Baja California, California, Chihuahua, Coahuila, Colorado, Nevada, New
Mexico, Sonora, Texas, Utah. All of Baja California and its associated islands (not
completely shown) are included in our concept of a natural North American Southwest
region; extreme northern Durango and Sinaloa are also included at Lat. 26º N.

 

1000 Nearctic

Continental North America exclusive of the tropics and certain highland areas south of the Tropic of Cancer. We include those tropic-subtropic regions in and adjacent to the North American Southwest and the Caribbean.

2000 Palaearctic

Eurasia exclusive of the tropics. Africa north of the Sahel.

3000 Neotropical and Antarctican

Continental South America, Central America, and most of Mexico south of the Tropic of Cancer. Antarctica.

4000 Oriental

Southeast Asia, the Indian subcontinent; the Phillipines, Indonesia, etc.

5000 Ethiopian

Africa south of the Sahara, Malagasy, and parts of the Arabian peninsula.

6000 Australian

Australia and Tasmania.

7000 Oceanic

Oceanic islands processing a high degree of endemism.


293

First Level.—The first digit after the comma (e.g., 11 ,00) refers to vegetation, the structural and readily measurable reality of ecosystems. Included are all potential and/or existing plant communities that are presumed to be established naturally under existing climate and the cessation of artificially disruptive (man-caused) influences[2](Table 1).

 

Table 1. Summary for the Natural Vegetation of the World to the First Digit level.

Biogeographic Realm

1. Upland Vegetation

2. Wetland Vegetation

1,000. Nearctic

1,100.

1,200.

2,000. Palaearctic

2,100.

2,200.

3,000. Neotropical-Antarctican

3,100.

3,200.

4,000. Oriental

4,100.

4,200.

5,000. Ethiopian

5,100.

5,200.

6,000. Australian

6,100.

6,200.

7,000. Oceanic

7,100.

7,200.

All existing and potential natural vegetation (PNV) is classified as belonging to uplands (1,100) or wetlands (1,200) as in Table 1. Cultivated lands are designated 1,300 (cultivated uplands) and 1,400 (cultivated wetlands). The evolutionary distinctions between plants and animals of terrestrial (upland) ecosystems and those of aquatic or hydric (wetland) ecosystems is recognized by this dichotomy (see Ray 1975).

As discussed here, wetlands include those periodically, seasonally or continually submerged ecosystems populated by species and/or life forms different from the immediately adjacent (upland) climax vegetation, and which are dependent on conditions more mesic than provided by the immediate precipitation. Certain ecosystems having both upland and wetland characteristics and components (e.g., riparian forests) could be properly considered as belonging to both divisions. They are treated in this report as wetlands (1,2 00).

Second Level.—The second digit after the comma (e.g. 1,11 0) refers to one of the following recognized ecological formations, which on a worldwide basis are the formation-types (biome-types); see Tables 2 and 3. On continents these are referred to as formations, which are vegetative responses (functions) to integrated environmental factors, most importantly plant-available moisture.

Upland Formations

 

Tundra[3]

Communities existing in an environment so cold that moisture is unavailable during most of the year, precluding the establishment of trees, and in which maximum vegetation development is of herbaceous root perennials, shrubs, lichens and mosses, with grasses poorly represented or at least not dominant.

Forest and Woodland Forest

Forest

Communities comprised principally of trees potentially over 15 meters (50 ft) in height, and frequently characterized by closed and/or multilayered canopies.

Woodland

Communities dominated by trees with a mean potential height usually under 15 meters in height, the canopy of which is usually open—sometimes very open[4] —or interupted and singularly layered.

Scrubland

Communities dominated by sclerophyll or microphyll shrubs and/or multistemmed trees, generally not exceeding 10 meters (31 ft) in height and usually presenting a closed physiognomy, or, if open, interspersed with other perennial vegetation.

Grassland

Communities dominated actually or potentially by grasses and/or other herbaceous plants.

Desertland

Communities in an arid environment—usually less than 300 mm (12 in) precipitation per annum—in which plants are separated by significant areas devoid of perennial vegetation.

 

Table 2. Summary for the Natural UPLAND Vegetation of the World to the Second Level (Formation-Type).

Formation Type

Biogeographic Realm

1. Tundra

2. Forest

3. Scrubland

4. Grassland

5. Desertland

6. Nonvegetated

1,000 Nearctic

1,110

1,120

1,130

1,140

1,150

1,160

2,000 Palaearctic

2,110

2,120

2,130

2,140

2,150

2,160

3,000 Neotropical-Antarctican

3,110

3,120

3,130

3,140

3,150

3,160

4,000 Oriental

4,110

4,120

4,130

4,140

4,150

4,160

5,000 Ethiopian

5,110

5,120

5,130

5,140

5,150

5,160

6,000 Australian

6,110

6,120

6,130

6,140

6,150

6,160

7,000 Oceanic

7,110

7,120

7,130

7,140

7,150

7,160

[2] Our thinking on the complex question of determining climax, successional, and potential vegetation is to consider (and map) ecosystems on the basis of the existing or presumed vegetation of the foreseeable future.

[3] The holistic integrity of a "Tundra" formation is not without question. Treated here, tundra may also be composed of grasslands, scrublands, marshlands (wet tundra), and desertlands in an Arctic-Boreal climatic zone (Billings and Mooney 1968; Billings 1973; and others.

[4] The "savanna" formation (Dansereau 1957; Dyksterhuis 1957; and others) is here recognized (in North America) as an ecotone between woodland and grassland. Those homogeneous areas in which the crowns of trees normally cover less than approximately 15 percent of the ground space are classified as grasslands where grasses are actually or potentially dominant (=savanna grassland). Mosaics of grassland and smaller or larger stands of trees and shrubs are "parklands" and are composed of two or more ecologically distinct plant formations (Walter 1973).


294
 

Table 3. Summary for the Natural WETLAND Vegetation of the World to the Second Level (Formation-Type).

Formation Type

Biogeographic Realm

1. Wet Tundra

2. Forest1

3. Swamp-scrub, Riparian Scrub

4. Marshland

5. Strandland

6. Submergent Aquatic

1,000 Nearctic

1,210

1,220

1,230

1,240

1,250

1,260

2,000 Palaearctic

2,210

2,220

2,230

2,240

2,250

2,260

3,000 Neotropical-Antarctican

3,210

3,220

3,230

3,240

3,250

3,260

4,000 Oriental

4,210

4,220

4,230

4,240

4,250

4,260

5,000 Ethiopian

5,210

5,220

5,230

5,240

5,250

5,260

6,000 Australian

6,210

6,220

6,230

6,240

6,250

6,260

7,000 Oceanic

7,210

7,220

7,230

7,240

7,250

7,260

1 / Swampforests, bog-forests and riparian forests.

Wetland Formations

 

Wet Tundra[5]

Wetland communities existing in an environment so cold that available plant moisture is unavailable during most of the year, precluding the establishment of trees and all but a low herbaceous plant structure in a hydric matrix.

Swampforest; Riparian Forest

Wetland communities possessing an over-story of trees potentially over 10 meters (31 ft) in height, and frequently characterized by closed and/or multilayered canopies.

Swampscrub;

Wetland communities dominated by short

Riparian Scrub

trees and/or woody shrubs, generally under 10 meters (31 ft) in height and often presenting a closed physiognomy.

Marshland

Wetland communities in which the principal plant components are herbaceous emergents which normally have their basal portions annually, periodically, or continually submerged.

Strandland

Beach and river channel communities subject to infrequent but periodic submersion, wind driven waves and/or spray. Plants are separated by significant areas devoid of perennial vegetation.[6]

Submergent Aquatic

Aquatic communities comprised entirely or essentially of plants mostly submerged or lacking emergent structures.

Some localized upland and wetland areas are essentially without vegetation or are sparingly populated by simple organisms, e.g., on some dunes, lava flows, playas, sinks, etc. For purposes of classification certain of such areas could be considered as belonging to a nonvegetated formation-type (Tables 2 and 3).

Third Level.—The third digit beyond the comma (e.g., 1,111) refers to one of four world climatic zones (c.f. Walter 1973; Ray 1975; Cox etal . 1976), in which minimum temperature remains a major evolutionary control of and within the zonation and the formation-types (Tables 4 and 5). All four of these broad climatic zones are found in North America and in the "Southwest."

 

Arctic-Boreal (Antarctic-Austreal)

Characterized by lengthy periods of freezing temperatures, with growing season of short duration (generally 60–150 days), occasionally interrupted by nights of below freezing temperatures.

Cold Temperate

Freezing temperatures of short duration although of frequent occurrence during winter months. Potential growing season generally 100–200 days and confined to spring and summer when freezing temperatures are infrequent or absent.

Warm Temperate

Freezing temperatures of short duration but generally occurring every year during winter months. Potential growing season over 200 days with an average of less than 125–150 days being subject to temperatures lower than 0 °C or to chilling fogs.

Tropical-Subtropical

Infrequent or no 24-hour periods of freezing temperatures, chilling fogs or wind.

Fourth Level.—The fourth digit beyond the comma (e.g., 1,111.1 ) refers to a subcontinental unit that is a major biotic community (=biome). Biomes are natural communities characterized by a distinctive vegetation physiognamy within a formation; accordingly, the natural geography of biomes is commonly disjunctive. A single biome is not to be confused with a single biotic (biogeographic) province; in distribution, a province is always a continuous (non-disjunctive) biogeographic area that may include several (e.g., five or more) biomes.[7]

Our nomenclature at the biome (fourth) level incorporates useful geographic terms in the same sense of Weaver and Clements (1938). While such terms are also associated with biotic provinces (as in Fig. 2) we are classifying biomes, not biotic provinces. Biomes are characterized by a distinctive evolutionary history within a formation; thus they tend to be centered in, but are not restricted to particular biogeographic regions or provinces (e.g., see Weaver and Clements 1938; Clements and Shelford 1939; Pitelka 1941, 1943; Dice 1939, 1943; Odum 1945; Allee etal . 1949; Kendeigh 1954, 1961; Dansereau 1957; Shelford 1963; Daubenmire and Daubenmire 1968; Udvardy 1975; Dasmann 1976).

This fourth level and the fifth level (below) have provided the most successful and useful mapping of states, regions, and continents (e.g., in North America, Harshberger 1911; Shreve 1917, 1951; Shantz and Zon 1924; Bruner 1931; Morris 1935; Wieslander 1935; Brand 1936;

[5] Treated here, tundra may also be composed of grasslands, scrublands, marshlands (wet tundra), and desertlands in an Arctic-Boreal climatic zone; see footnote 3.

[6] Strand communities are situated in harsh physical environments that produce their characteristic physiognomy. Accordingly, strandland is treated as the wetland equivalent of desertland. While occurring in the usual sense on beaches and other seacoast habitats, freshwater (or interior) strands also occur in river channels, along lake margins, and below reservoir high water lines.

[7] Originally termed biotic provinces by Lee Dice (1943) who developed this biogeographic concept in North America between 1922 (biotic areas) and 1943 (biotic provinces), they have been referred to variously in recent literature as "biotic provinces" (Dasmann 1972, 1974; IUCN 1973), "biogeograpt' provinces" (Udvardy 1975; Dasmann 1976). "ecoregions" (Bailey 1976, 1978), and "biotic gions" (Franklin 1977).


295
 

Table 4. Summary for the Natural UPLAND Vegetation of Nearctic and Adjacent Neotropical North America to the Third Level.

Climatic (Thermal) Zone

Formation

1. Arctic-Boreal

2. Cold Temperate

3. Warm Temperate

4. Tropical-Subtropical

1,110 Tundra

1,111

     

1,120 Forest & Woodland

1,121

1,122

1,123

1,124

1,130 Scrubland

1,131

1,132

1,133

1,134

1,140 Grassland

1,141

1,142

1,143

1,144

1,150 Desertland

1,151

1,152

1,153

1,154

1,160 Nonvegetated

1,161

1,162

1,163

1,164

 

Table 5. Summary for the Natural WETLAND Vegetation of Nearctic and Adjacent Neotropical North America to the Third Level.

Climatic (Thermal) Zone

Formation

1. Arctic-Boreal

2. Cold Temperate

3. Warm Temperate

4. Tropical-Subtropical

1,210 Wet Tundra

1,211

     

1,220 Forest1

1,221

1,222

1,223

1,224

1,230 Swampscrub

1,231

1,232

1,233

1,234

1,240 Marshland

1,241

1,242

1,243

1,244

1,250 Strandland

1,251

1,252

1,253

1,254

1,260 Submergent Aquatic

1,261

1,262

1,263

1,264

1 Swampforests, bog-forests and reparian forests.

Nichol 1937; LeSueur 1945; Jensen 1947; Leopold 1950; Castetter 1956; Küchler 1964, 1977; Brown 1973; Franklin and Dyrness 1973; Brown and Lowe 1977). Biomes and biogeographic provinces are also the bases for the biosphere reserve program (MAB) in the United States and elsewhere (IUCN 1974; Franklin 1977).

A partial summary of the biotic communities (biomes) for Nearctic and adjacent Neotropical America is given in Tables 6 and 7.

Fifth Level.—The fifth digit beyond the comma (e.g., 1,111.11 ) refers to the principal plant-animal communities within the biomes, distinguished primarily on taxa that are distinctive climax plant dominants. Daubenmire and Daubenmire (1968) organized their data according to major dominants in climax communities referred to as climax series. "Series," or "cover-types" (sensu Society of American Foresters 1954), or "vegetation-types" (sensu Flores etal . 1971), are each composed of one or more biotic associations characterized by shared climax dominants within the same formation, zone, and biome (Oosting 1950; Lowe 1964; Franklin and Dyrness 1973; Pfister etal . 1977). For example, within Rocky Mountain montane conifer forest (122.3), the Pine Series (122.32) includes all of the Rocky Mountain forest associations in which Pinus ponderosa is a dominant.

Community diversity of tropical and subtropical upland climax dominants is inherently more complex than in boreal and temperate communities. Moreover, some taxa may exhibit polymorphism to the extent that the same species may be dominant—and ecotypically differentiated—in more than a single formation. As an extreme case in southwestern North America, mesquite (Prosopisjuliflora ) may be a dominant life-form in certain desertland, disclimax grassland, scrubland, woodland, and riparian forest communities, and exhibit phenotypic and presumably genotypic population differentiation across the complex gradient. Facultative growth-form is exhibited by dominant plant taxa in both cold and warm climatic zones.

The distribution of some plant dominants also may span more than a single climatic zone, as in Larrea, Prosopis, and the introduced Tamarix. However, important plant and animal associates of these dominant species are usually encountered when passing from one formation or climatic zone to another. When specific and generic dominants are shared by more than one biome, closer investigation may reveal genetic geographic variation within the shared species, as in the chromosome races of creosotebush (Larrea divaricata, Yang and Lowe 1968; Yang 1970).

It is clear that the determination of fifth and sixth (below) level communities in particular will require modification and revision in the classification as field data accumulate. Some of the more widely distributed and commonly recognized series in the Southwest are given in Tables 6 and 7 under the appropriate biome.

Sixth Level.—The sixth digit beyond the comma (e.g., 1,111.111 ) refers to distinctive plant associations, and associes (successional associations), based on the occurrence of particular dominant species more or less local or regional in distribution and generally equivalent to habitat-types as outlined by the Daubenmires (1968), Layser (1974), Pfister et al . (1977), and others. While we give examples for certain communities within southwestern biomes, the enormous numbers of sets precludes presentation here for the treatments given in Tables 6 and 7. Associations may be added at length for regional studies by using a, b, c, sets as is also indicated in the tables in Brown and Lowe (1974a,b).

Seventh Level.—The seventh digit beyond the comma (e.g., 1,111.1111 ) accommodates detailed measurement and assessment of quantitative structure, composition, density and other attributes for dominants, understories, and other associated species. This level and additional ones in the system provide the flexibility required for encompassing data for ecological parameters measured in intensive studies on limited areas (see e.g., Dick-Peddie and Moir 1970).


296

figure

Figure 2.
Biogeographic provinces of Nearctic and adjacent Neotropical North America (modified
after Dice 1943, and Dasmann 1974), discussed in text under fourth (Biome) digit level.

 

1. Polar

  8. Plains

15. Mogollon (Interior)

2. Alaskan

  9. Rocky Mountain

16. Chihuahuan

3. Greenlandian

10. Great Basin

17. Tamaulipan

4. Canadian

11. Sierran-Cascade

18. Madrean

5. Northeastern

12. Sitkan-Oregonian

19. Sonoran

6. Southeastern

13. Californian

20. Sinaloan

7. Gulf Coastal

14. Mohavian

21. San Lucan

   

22. Carribean


297
 

Table 6. Nomenclature of UPLAND Biotic Communities of Nearctic and Adjacent Neotropical North America with Community (Series) and Association Examples for the North American Southwest.

1,100 Nearctic Upland Vegetation

1,110 Tundra Formation

1,111 Arctic Tundras

1,111.1 Polar (High Arctic) Tundra

1,111.2 Alaskan (Low Arctic) Coastal Tundra

1,111.3 Canadian (Barren Ground = Low Arctic) Tundra

1,111.4 Arctic Alpine Tundra

1,111.5 Rocky Mountain Alpine Tundra

1,111.511 Lichen-Moss Series*

111.511 Rhizocarpon geographicum Association*

111.52 Mixed Herb Series*

111.53 Avens-Sedge Series*

111.531 Geum turbinatum Association*

111.532 Geum turbinatum-Carex bella Association*

111.54 Woodrush Series*

111.541 Kobresia bellardi -grass-forb Association*

111.6 Sierran-Cascade Alpine Tundra

111.61 Lichen-Moss Series*

111.62 Mixed Herb Series*

111.621 Selaginella watsoni-Eriogonum umbellatum et al. Association*

111.7 Adirondack-Appalachian Alpine Tundra

120 Forest and Woodland Formation

121 Boreal Forests and Woodlands

121.1 Canadian Subarctic Conifer Forest and Woodland (North American Taiga)

121.2 Appalachian Subalpine Conifer Forest

121.3 Rocky Mountain Subalpine Conifer Forest and Woodland2

121.31 Engelmann Spruce-Alpine Fir Series*

121.311 Picea engelmanni-Abies lasiocarpa Association*

121.312 Picea engelmanni Association*

121.313 Abies lasiocarpa Association*

121.314 Abies lasiocarpa arizonica Association*

121.315 Picea pungens Association*

121.316 Populus tremuloides subclimax Association*

121.32 Bristlecone Pine-Limber Pine Series*

121.321 Pinus aristata-Pinus flexilis Association*

121.322 Pinus aristata Association*

121.323 Pinus flexilis Association*

121.4 Sierran-Cascade Subalpine Conifer Forest

121.41 Limber Pine-Lodgepole Pine Series*

121.411 Pinus flexilis-Pinus contorta murrayana Association*

121.412 Pinus flexilis Association*

121.5 Madrean Subalpine Conifer Forest

122 Cold Temperate Forests and Woodlands

122.1 Northeastern Deciduous Forest

122.2 Pacific Coastal (Oregonian) Conifer Forest

122.3 Rocky Mountain (=Petran) Montane Conifer Forest

122.31 Douglas-fir-White Fir (=Mixed Conifer) Series*

122.311 Pseudotsuga menziesi Association*

122.312 Pseudotsuga menziesi-Abies concolor Association*

122.313 Pseudotsuga menziesi -mixed conifer (Abies concolor, Pinus flexilis, Acer glabrum,
       Populus tremuloides, Pinus ponderosa
) Association*

122.314 Populus tremuloides subclimax Association*

122.32 Pine Series*

122.321 Pinus ponderosa Association*

122.322 Pinus ponderosa -mixed conifer Association*

122.323 Pinus ponderosa-Quercus gamnbeli Association*

122.324 Pinus ponderosa-Quercus arizonica Association*

122.325 Pinus ponderosa-Juniperus deppeana Association*

122.326 Populus tremuloides subclimax Association*

122.327 Pinus flexilis reflexa Association*

122.328 Pinus ponderosa-Pinus leiophylla Association*

122.33 Gambel Oak Series*

122.331 Quercus gambeli Association*

122.4 Great Basin Conifer Woodland

122.41 Pinyon-Juniper Series*

122.411 Pinus edulis-Juniperus scopulorum Association*

122.412 Pinus edulis Association*

122.413 Juniperus scopulorum Association*

122.414 Pinus edulis-Juniperus monosperma Association*

122.415 Juniperus monosperma Association*

122.416 Pinus monophylla-Juniperus osteosperma Association*

122.417 Pinus monophylla Association*

122.418 Juniperus osteosperma Association *

122.419 Pinus monophylla-Juniperus californica Association*

122.411a Juniperus-californica Association*

122.412a Pinus quadrifolia-Juniperus californica Association*

122.413a Pinus quadrifolia Association*

122.414a Pinus monophylla-Juniperus californica- chaparral Association*

122.5 Sierran-Cascade Montane Conifer Forest

122.51 Mixed Conifer Series*

122.511 Abies concolor -mixed conifer (Pinus contorta
      murrayana, Pinus jeffreyi
et al.) Association*

122.52 Pine Series*

122.521 Pinus ponderosa Association*

122.522 Pinus ponderosa-P. jeffreyi Association*

122.523 Pinus ponderosa-Quercus kelloggi Association*

122.524 Pinus jeffreyi Association*

122.53 Black Oak Series

122.531 Quercus kelloggi Association*

122.6 Madrean Montane Conifer Forest

122.61 Douglas-fir-Mixed Conifer Series*

122.611 Pseudotsuga menziesi Association*

122.612 Pseudotsuga menziesi-Pinus flexilis, Acer glabrum, Populus tremuloides, Pinus ponderosa et al. Association*

122.62 Pine Series*

122.621 Pinus flexilis ayacahuite Association*

122.622 Pinus ponderosa Association*

122.623 Pinus ponderosa -mixed conifer Association*

122.624 Pinus ponderosa-Quercus spp. Association*

122.625 Pinus ponderosa-Juniperus deppeana Association*

123 Warm Temperate Forests and Woodlands

123.1 Southeastern Mixed Deciduous and Evergreen Forest

123.2 Californian Mixed Evergreen Forest

123.21 Mixed Mesophytic Series*

* Examples only.

1 The first "I" (in front of comma and representing the Nearctic Realm) is understood, and cropped for tabular convenience only, from this point onward.

2 Separation of this biotic community into Rocky Mountain and Great Basin units may be warranted.


298
 

123.211 Mixed hardwood Association*

123.212 Quercus chrysolepis -mixed hardwood Association*

123.22 Big-cone Spruce Series*

123.221 Pseudotsuga macrocarpa Association*

123.3 Madrean Evergreen Forest and Woodland

123.31 Encinal (Oak) Series*

123.311 Mixed Quercus (=Quercus spp.) Association*

123.312 Quercus grisea Association*

123.313 Quercus emoryi Association*

123.314 Quercus chihuahuaensis Association*

123.315 Quercus arizonica Association*

123.316 Quercus spp.-Pinus cembroides-Juniperus spp. Association*

123.317 Pinus cembroides Association*

123.318 Juniperus deppeana Association*

123.32 Oak-Pine Series*

123.321 Quercus hypoleucoides-Quercus rugosa Association*

123.322 Quercus spp.-Pinus leiophylla Association*

123.323 Quercus spp.-Pinus engelmanni Association*

123.324 Quercus spp.-Pinus spp. Association*

123.325 Quercus spp.-Arbutus xalapensis-Pinus spp. Association*

123.4 Californian Evergreen Woodland

123.41 Encinal (Oak) Series*

123.411 Mixed Quercus Association*

123.412 Quercus agrifolia Association*

123.413 Quercus agrifolia-Juglans californica Association*

123.414 Quercus engelmanni Association*

123.42 Walnut Series*

123.421 Junglans californica Association*

123.5 Relict Conifer Forest and Woodland

123.51 Closed-cone Pine Series*

123.511 Pinus attenuata Association*

123.512 Pinus muricata Association*

123.513 Pinus torreyana Association*

123.52 Cypress Series*

123.521 Cupressus arizonica arizonica Association*

123.522 Cupressus arizonica glabra Association*

123.523 Cupressus arizonica stephensoni Association*

123.524 Cupressus arizonica montana Association*

123.525 Cupressus forbesi Association*

123.526 Cupressus forbesi-Pinus muricata Association*

124 Tropical-Subtropical Forests and Woodlands

124.1 Caribbean Montane Rain Forest

124.2 Caribbean Cloud Forest

124.3 Caribbean Evergreen Forest

124.4 Caribbean Deciduous Forest

124.5 Tamaulipan Deciduous Forest

124.6 Sinaloan Deciduous Forest

124.61 Mixed Short Tree Series*

124.611 Mixed Deciduous Association*

124.612 Lysiloma watsoni -mixed deciduous Association*

124.613 Conzattia sericea Association*

124.614 Ceiba acuminata -mixed deciduous Association*

124.615 Bursera inopinnata -mixed deciduous Association*

130 Scrubland Formation

131 Arctic-Boreal Scrublands

131.1 Alaskan (Low Arctic) Coastal Scrub

131.2 Canadian (Low Arctic, Barren Ground) Subpolar Scrub

131.3 Alaskan Alpine and Subalpine Scrub

131.4 Adirondack-Appalachian Alpine and Subalpine Scrub

131.5 Rocky Mountain Alpine and Subalpine Scrub

131.51 Willow Series*

131.511 Salix brachycarpa Association*

131.512 Salix planifolia Association*

131.513 Salix nivalis Association*

131.52 Spruce Elfinwood Series*

131.521 Picea engelmanni Association*

131.53 Bristlecone Pine Elfinwood Series*

131.531 Pinus aristata Association

131.6 Sierran-Cascade Alpine and Subalpine Scrub

131.61 Limber Pine-Lodgepole Pine Elfinwood Series*

131.611 Pinus flexilis Association*

132 Cold Temperate Scrublands

132.1 Great Basin Montane Scrub

132.11 Oak-scrub Series*

132.111 Quercus gambeli Association*

132.12 Mountain mahogany Series*

132.121 Cercocarpus montanus Association*

132.13 Maple-scrub Series*

132.131 Acer grandidentatum Association*

132.14 Serviceberry Series*

132.141 Amelanchier alnifolia Association*

132.15 Bitterbush Series*

132.151 Purshia tridentata Association*

132.16 Mixed Deciduous Series*

132.161 Mixed Scrub Association*

132.2 Sierran-Cascade Montane Scrub

132.21 Manzanita Series*

132.211 Arctostaphylos glauca Association*

132.212 Arctostaphylos glandulosa Association*

132.22 Mixed Scrub Series*

132.221 Mixed scrub Association*

132.3 Plains Deciduous Scrub

132.31 Oak-Scrub Series*

132.311 Quercus harvardi Association*

132.32 Sumac Series*

132.321 Rhus lanceolata Association*

132.33 Mixed Deciduous Series*

132.331 Prunus americana et al. Association*

133 Warm Temperate Scrublands

133.1 Californian Chaparral

133.11 Chamise Series*

133.111 Adenostoma fasciculatum Association*

133.112 Adenostoma fasciculatum -mixed sclerophyll Association*

133.113 Adenostoma sparsifolium Association*

133.12 Scrub Oak Series*

133.121 Quercus dumosa Association*

133.122 Quercus dumosa -mixed sclerophyll Association*

133.123 Quercus dumosa-Quercus wislizeni Association*

133.13 Manzanita Series*

133.131 Arctostaphylos glauca Association

133.132 Arctostaphylos glandulosa Association*

133.133 Arctostaphylos glandulosa-Pinus coulteri Association*

133.134 Arctostaphylos glandulosa -mixed sclerophyll Association*

133.14 Ceanothus Series*

133.141 Ceanothus cordulatus Association*

133.142 Ceanothus spp.-mixed sclerophyll Association*

133.2 Californian Coastalscrub

133.21 Sage Series*

133.211 Artemisia californica Association*

133.212 Artemisia californica-Salvia spp. Association

133.22 Mixed Shrub Series*

133.221 Eriogonum fasciculatum-Simmondsia chinensis et al. Association*

133.222 Encelia californica -Mixed shrub Association*

* Examples only.


299
 

133.3 Interior Chaparral

133.31 Scrub Oak Series*

133.311 Quercus turbinella Association*

133.312 Quercus turbinella-Cerocarpus breviflorus Association*

133.313 Quercus turbinella-Cercocarpus betuloides Association*

133.314 Quercus turbinella -mixed sclerophyll Association*

133.315 Quercus intricata Association*

133.316 Quercus intricata-Cercocarpus spp. Association*

133.317 Quercus intricata-Quercus spp. Association*

133.318 Quercus intricata -mixed sclerophyll Association*

133.319 Quercus pungens Association*

133.311a Quercus pungens -mixed sclerophyll Association*

133.32 Manzanita Series*

133.321 Arctostaphylos pringlei Association*

133.322 Arctostaphylos pungens Association*

133.33 Ceanothus Series*

133.331 Ceanothus greggi Association*

133.332 Ceanothus greggi -mixed sclerophyll Association*

133.34 Mountain mahogany Series*

133.341 Cercocarpus breviflorus Association*

133.342 Cercocarpus montanus Association*

133.35 Silktassel Series*

133.351 Garrya wrighti Association*

133.352 Garrya ovata Association*

133.36 Mixed Evergreen Sclerophyll Series*

133.361 Mixed sclerophyll Association*

133.4 Southeastern Maritime Scrub

134 Tropical-Subtropical Scrublands

134.1 Caribbean Thornscrub

134.2 Tamaulipan Thornscrub

134.3 Sinaloan Thornscrub

134.31 Mixed Deciduous Series*

134.311 Mixed scrub-Fouquieria macdougali Association*

134.312 Mixed scrub-lpomoea arborescens Association*

134.313 Mixed scrub-Lysiloma divaricata Association*

134.314 Mixed scrub-Acacia cymbispina Association*

134.315 Mixed scrub-Ceiba acuminata Association*

134.316 Mixed scrub-Mixed tree Association*

134.32 Mesquite Disclimax Series*

134.321 Prosopis juliflora velutina -mixed scrub Association*

140 Grassland Formation

141 Arctic-Boreal Grasslands

141. 1 Alaskan (Low Arctic) Coastal Grassland

141.2 Canadian (Low Arctic) Grassland

141.3 Appalachian Subalpine (Balds) Grassland

141.4 Rocky Mountain Alpine and Subalpine Grassland

141.41 Bunchgrass Series*

141.411 Festuca thurberi Association*

141.412 Festuca arizonica Association*

141.413 Mixed grass-forb Association*

141.42 Sedge-Forb-Grass Series*

141.421 Carex spp.-mixed forb-grass Association*

141.5 Sierran Cascade Alpine and Subalpine Grassland

141.51 Bunchgrass Series*

141.511 Sitanion hystrix -mixed forb Association*

141.512 Mixed grass-forb Association*

141.52 Sedge-Forb-Grass Series*

141.521 Carex spp.-mixed forb-grass Association*

141.6 Madrean Alpine and Subalpine Grassland

142 Cold Temperate Grasslands

142.1 Plains Grassland

142.11 Bluestem "tall-grass" Series*

142.111 Andropogon scoparius Association*

142.112 Andropogon spp.-mixed tall-grass Association*

142.113 Andropogon spp.-Quercus harvardi Association*

142.114 Mixed tall-grass Association*

142.115 Artemisia filifolia -mixed scrub disclimax Association*

142.12 Grama "short-grass" Series*

142.121 Bouteloua gracilis Association*

142.122 Bouteloua spp. Association*

142.123 Bouteloua spp.-mixed grass Association*

142.124 Bouteloua spp.-mixed grass-mixed scrub Association*

142.13 Buffalo-grass Series*

142.131 Buchloe dactyloides -mixed grass Association*

142.14 Mixed "Short-grass" Series*

142.141 Aristida spp.-Bouteloua gracilis-Buchloe dactyloides Association*

142.15 Shrub-Grass Disclimax Series*

142.151 Gutierrezia sarothrae Association*

142.2 Great Basin Shrub-Grassland

142.21 Wheatgrass Series*

142.211 Agropyron smithi Association*

142.212 Agropyron smithi -mixed scrub Association*

142.213 Agropyron smithi-Artemisia tridentata Association*

142.22 Mixed Bunchgrass Series*

142.221 Mixed grass Association*

142.222 Mixed grass-Artemisia tridentata Association*

142.23 Ricegrass Series*

142.231 Oryzopsis hymenoides Association*

142.24 Sacaton Series*

142.241 Sporobolus airoides Association*

142.242 Sporobolus airoides-Atriplex canescens Association*

142.3 Pacific Coastal (Oregonian) Grassland

142.4 Rocky Mountain Montane Grassland

142.41 Mixed Meadow Series*

142.411 Mixed forb-grass Association*

142.42 Rush Series*

142.421 Juncus spp. Association*

142.43 Fern Series*

142.431 Pteridium aquilinum Association*

142.44 Iris Disclimax Series*

142.441 Iris missouriensis Association*

142.5 Sierran-Cascade Montane Grassland

142.51 Mixed Meadow Series*

142.511 Mixed forb-grass Association*

142.52 Rush Series*

142.521 Juncus spp. Association*

143 Warm Temperate Grasslands

143.1 Scrub-Grassland (Semidesert Grassland)

143.11 Grama Grass-Scrub Series*

143.111 Bouteloua eriopoda-Yucca elata Association*

143.112 Bouteloua eriopoda-Prosopis juliflora Association*

143.113 Bouteloua eriopoda -mixed grass-mixed scrub Association*

143.114 Bouteloua spp.-mixed grass-mixed scrub Association*

143.12 Tobosa Grass-Scrub Series*

143.121 Hilaria mutica Association*

143.122 Hilaria mutica-Prosopis juliflora Association*

* Examples only.


300
 

143.123 Hilaria mutica -mixed scrub Association*

143.13 Curleymesquite grass-scrub Series*

143.131 Hilaria belangeri -mixed scrub Association*

143.14 Sacaton-Scrub Series*

143.141 Sporobolus wrighti Association*

143.142 Sporobolus wrighti-Prosopis juliflora Association*

143.15 Mixed Grass-Scrub Series*

143.151 Mixed grass-Yucca elata Association*

143.152 Mixed grass-Prosopis juliflora Association

143.153 Mixed grass-Acacia greggi Association*

143.154 Mixed grass-Fouquieria splendens Association*

143.155 Mixed grass-mixed scrub Association*

143.16 Shrub-Scrub Disclimax Series*

143.161 Aplopappus tenuisectus Association*

143.162 Aplopappus tenuisectus-Yucca elata Association*

143.163 Aplopappus tenuisectus-Prosopis juliflora Association*

143.164 Aplopappus tenuisectus -mixed scrub Association*

143.165 Gutierrezia sarothrae-Prosopis juliflora Association*

143.2 Californian Valley Grassland

143.21 Annual Disclimax Series*

143.211 Mixed annual grass Association*

143.212 Avena fatua Association*

143.213 Bromus rubens Association*

143.214 Mixed forb Association*

144 Tropical-Subtropical Grasslands

144.1 Caribbean Savanna Grassland

144.2 Gulf Coastal (Tamaulipan) Grassland

144.3 Sonoran Savanna Grassland

144.31 Mixed Root-perennial Grass Series*

144.311 Heteropogon contortus-Bouteloua spp.-Aristida spp.-mixed scrub Association*

144.32 Grama Series*

144.321 Bouteloua rothrocki-Prosopis juliflora Association*

144.322 Bouteloua spp.-mixed scrub Association*

144.33 Three-awn Series*

144.331 Aristida spp.-Prosopis juliflora Association*

144.332 Aristida spp.-mixed scrub Association*

150 Desertland Formation

151 Arctic-Boreal Desertlands

151.1 Polar Desertscrub

152 Cold Temperate Desertlands

152.1 Great Basin Desertscrub

152.11 Sagebrush Series*

152.111 Artemisia tridentata Association*

152.112 Artemisia tridentata -mixed scrub-grass Association*

152.113 Artemisia nova Association*

152.12 Shadscale Series*

152.121 Atriplex confertifolia Association*

152.122 Atriplex confertifolia -mixed scrub Association*

152.13 Blackbrush Series*

152.131 Coleogyne ramosissima Association*

152.14 Rabbitbrush Series*

152.141 Chrysothamnus nauseosus Association*

152.15 Winterfat Series*

152.151 Eurotia lanata Association*

152.152 Eurotia lanata -mixed scrub Association*

152.16 Mixed scrub Series*

152.161 Ephedra viridis-Eriogonum spp.-mixed scrub Association*

152.17 Saltbush Series*

152.171 Sarcobatus vermiculatus Association*

152.172 Atriplex canescens Association*

153 Warm Temperate Desertlands

153.1 Mohave Desertscrub

153.11 Creosotebush Series*

153.111 Larrea divaricata Association*

153.112 Larrea divaricata-Ambrosia dumosa Association*

153.113 Larrea divaricata-Yucca spp. Association*

153.12 Blackbrush Series*

153.121 Coleogyne ramosissima Association*

153.122 Coleogyne ramosissima-Yucca spp. Association*

153.13 Mesquite Series*

153.131 Prosopis juliflora torreyana Association*

153.14 Bladdersage Series*

153.141 Salazaria mexicana Association*

153.15 Joshuatree Series*

153.151 Yucca brevifolio-Acamptopappus sphaerocephalus-Larrea divaricata -mixed scrub Association*

153.152 Yucca brevifolia-Coleogyne ramosissima Association*

153.153 Yucca brevifolia-Larrea divaricata Association*

153.16 Catclaw Series*

153.161 Acacia greggi -mixed scrub Association*

153.17 Saltbush Series*

153.171 Suaeda torreyana Association*

153.172 Atriplex spp. Association*

153.2 Chihuahuan Desertscrub

153.21 Creosotebush-Tarbush Series*

153.212 Larrea divaricata-Parthenium incanum -mixed scrub Association*

153.213 Larrea divaricata-Flourensia cernua Association*

153.214 Flourensia cernua Association*

153.22 Whitethorn Series*

153.221 Acacia neovernicosa Association*

153.222 Acacia neovernicosa-Larrea divaricata Association*

153.23 Sandpaperbush Series*

153.231 Mortonia scabrella Association*

153.232 Mortonia scabrella-Rhus microphylla Association*

153.24 Mesquite Series*

153.241 Prosopis juliflora glandulosa (shrub hummock) Association*

153.242 Prosopis juliflora glandulosa-Artemesia filifolia Association*

153.25 Succulent Series*

153.251 Agave lecheguilla Association*

153.252 Agave lecheguilla-Yucca spp. Association*

153.253 Opuntia spp.-Agave spp.-Larrea divaricata Association*

153.26 Mixed Scrub Series*

153.261 Fouquieria splendens -mixed scrub Association*

153.27 Saltbush Series*

153.271 Suaeda torreyana Association*

153.272 Atriplex canescens Association*

153.273 Atriplex spp.-Artemisia filifolia Association*

154 Tropical-Subtropical Desertlands

154.1 Sonoran Desertscrub

154.11 Creosotebush-Bursage ("Lower Colorado Valley") et al Series*

154.111 Larrea divaricata Association*

154.112 Larrea divaricata-Ambrosia dumosa Association*

154.113 Ambrosia dumosa Association*

154.114 Prosopis juliflora torreyana (shrub hummock) Association*

154.115 Cercidium floridum-Olneya tesota-Dalea spinosa riparian Association*

154.116 Fouquieria splendens-Agave deserti Association*

* Examples only.


301
 

154.117 Opuntia bigelovi Association*

154.12 Paloverde-Mixed Cacti ("Arizona Upland") Series*

154.121 Ambrosia deltoidea-Cercidium microphyllum- mixed scrub Association*

154.122 Ambrosia deltoidea-Carnegiea gigantea -mixed scrub Association*

154.123 Simmondsia chinensis -mixed scrub Association*

154.124 Larrea divaricata-Canotia holacantha Association*

154.125 Larrea divaricata -mixed scrub Association*

154.126 Encelia farinosa -mixed scrub Association*

154.127 Mixed shrub-Cercidium microphyllum-Olneya tesota -mixed scrub Association

154.13 Brittlebush-Ironwood ("Plains of Sonora") Series*

154.131 Encelia farinosa-Olneya tesota Association*

154.132 Encelia farinosa -mixed scrub Association*

154.133 Mixed shrub-mixed scrub Association*

154.134 Mixed shrub-Prosopis juliflora velutina Association*

154.135 Mixed shrub-Forchammeria watsoni Association*

154.14 Copal-Torote ("Central Gulf Coast") Series*

154.141 Jatropha cinerea-Bursera microphylla Association*

154.142 Jatropha spp.-Bursera microphylla-Pachycereus pringlei Association*

154.143 Jatropha spp.-Ldria columnaris -mixed scrub Association*

154.15 Agave-Bursage ("Vizcaino") Series*

154.151 Ambrosia chenopodifolia-Agave shawi Association*

154.152 Ambrosia spp.-Agave shawi-Pachycormus discolor-Ldria columnaris -mixed scrub Association*

154.153 Ambrosia spp.-Agave shawi-Pachycereus pringlei -mixed scrub Association*

154.154 Mixed shrub-Agave shawi Association*

154.155 Eriogonum fasiculatum -mixed scrub Association*

154.16 Paloblanco-Agria ("Magdalena") Series*

154.161 Machaerocereus gummosus -mixed scrub Association*

154.17 Saltbush Series

154.171 Suaeda torreyana Association*

154.172 Allenrolfea occidentalis Association*

154.173 Atriplex spp.-Prosopis juliflora torreyana Association*

154.174 Atriplex polycarpa-Lycium spp.-Prosopis juliflora velutina Association*

154.175 Frankenia palmeri-Atriplex julacea Association*

 

Table 7. Nomenclature of WETLAND Biotic Communities (Fourth Level) of Nearctic and Adjacent Neotropical North America with Some Community (Series) and Association Level Examples for the North American Southwest.

1,200 Nearctic Wetland Vegetation

1,210 Wet Tundra Formation

1,211 Arctic Wet Tundra

211.1 Polar (High Arctic) Wet Tundra1 /

211.2 Greenlandian Wet Tundra

211.3 Alaskan (Coastal) Wet Tundra

211.4 Canadian (Low Arctic) Wet Tundra

220 Forest Formation

221 Boreal Swamp and Riparian Forests

221.1 Canadian Swampforest

222 Cold Temperate Swamp and Riparian Forests

222.1 Northeastern Bog, Swamp and Riparian Forests

222.2 Plains and Great Basin Riparian Deciduous Forest

222.21 Cottonwood-Willow Series*

222.211 Populus sargenti Association*

222.212 Populus sargenti-Salix amygdaloides Association*

222.213 Populus wislizeni Association*

222.214 Populus spp.-Salix spp. Association*

222.215 Salix exigua Association*

222.3 Rocky Mountain Riparian Deciduous Forest

222.31 Cottonwood-Willow Series*

222.311 Populus angustifolia-Salix spp. Association*

222.32 Mixed Broadleaf Series*

222.321 Acer negundo-Populus angustifolia -mixed deciduous Association*

222.322 Acer grandidentatum Association*

222.4 Sierran-Cascade Riparian Deciduous Forest

222.41 Cottonwood-Willow Series*

222.411 Populus trichocarpa-Salix spp. Association*

222.42 Mixed Broadleaf Series*

222.412 Acer macrophyllum-Populus trichocarpa-Alnus rhombifolia -mixed deciduous Association*

223 Warm Temperate Swamp and Riparian Forests

223.1 Southeastern Swamp and Riparian Forest

223.2 Interior Southwestern Riparian Deciduous Forest and Woodland

223.21 Cottonwood-Willow Series*

223.221 Populus fremonti-Salix spp. Association*

223.212 Populus fremonti Association*

223.213 Populus wislizeni Association*

223.214 Populus acuminata Association*

223.22 Mixed Broadleaf Series*

223.221 Platanus wrighti-Fraxinus velutina-Populus fremonti -mixed deciduous Association*

223.222 Platanus wrighti Association*

223.223 Fraxinus velutina Association*

223.224 Alnus oblongifolia Association*

223.225 Juglans major Association*

223.3 Californian Riparian Deciduous Forest and Woodland

223.31 Cottonwood-Willow Series

223.311 Populus fremonti-Salix spp. Association*

223.32 Mixed Broadleaf Series

223.321 Platanus racemosa -mixed deciduous Association*

223.322 Alnus rhombifolia Association*

224 Tropical-Subtropical Swamp, Riparian and Oasis Forests.

224.1 Caribbean Interior Swamp and Riparian Forests

224.2 Caribbean Maritime Swampforest

224.3 Tamaulipan Interior Swamp and Riparian Forests

224.4 Sinaloan Interior Swamp and Riparian Forests

224.41 Mixed Evergreen Series*

224.411 Ficus spp.-mixed evergreen and deciduous Association*

224.412 Taxodium mucronatum Association*

224.413 Populus sp.-mixed evergreen and deciduous Association*

224.42 Palm Series*

224.421 Sabal uresana Association*

224.5 Sonoran Riparian and Oasis Forests

224.51 Palm Series*

224.511 Washingtonia filifera Association*

224.512 Washingtonia filifera-Populus fremonti Association*

224.513 Washingtonia filifera-Brahea armata Association*

*One or more examples only are given for these levels.

1 The first "I" (in front of comma and representing the Nearctic Realm) is understood, and cropped for tabular convenience only, from this point onward.


302
 

224.514 Brahea armata Association*

224.515 Phoenix dactylifera-Washingtonia filifera Association*

224.52 Mesquite Series*

224.521 Prosopis juliflora velutina Association*

224.522 Prosopis juliflora velutina -mixed short tree Association*

224.53 Cottonwood-Willow Series*

224.531 Populus fremonti-Salix gooddingi Association*

224.532 Populus fremonti Association*

224.533 Salix gooddingi Association*

230 Swampscrub Formation

231 Arctic-Boreal Swampscrubs

231.1 Polar (High Arctic) Swampscrub

231.2 Greenlandian Swampscrub

231.3 Alaskan Swampscrub

231.4 Canadian Swampscrub

231.5 Adirondack-Appalachian Alpine and Subalpine Swamp and Riparian Scrub

231.6 Rocky Mountain Alpine and Subalpine Swamp and Riparian Scrub

231.61 Willow Series*

231.611 Salix bebbiana Association*

231.7 Sierran-Cascade Alpine and Subalpine Swamp and Riparian Scrub

231.71 Willow Series*

231.711 Salix spp. Association*

232 Cold Temperate Swamp and Riparian Scrubs

232.1 Northeastern Deciduous Swampscrub

232.2 Plains and Great Basin Swamp and Riparian Scrub

232.21 Willow Series*

232.211 Salix spp.-mixed scrub Association*

232.22 Saltcedar Disclimax Series*

232.221 Tamarix chinensis Association*

232.3 Rocky Mountain Riparian Scrub

232.31 Willow-Dogwood Series*

232.311 Salix spp.-mixed deciduous Association*

232.4 Sierran-Cascade Riparian Scrub

232.41 Willow Series*

232.411 Salix spp. Association

232.5 Pacific Coastal (Oregonian) Swamp and Riparian Scrub

233 Warm Temperate Swamp and Riparian Scrubs

233.1 Southeastern Mixed Deciduous and Evergreen Swampscrub

233.2 Interior Southwestern Swamp and Riparian Scrub

233.21 Mixed Narrowleaf Series*

233.211 Cephalanthus occidentalis-Baccharis glutinosa- mixed scrub Association*

233.22 Saltcedar Disclimax Series*

233.221 Tamarix chinensis -mixed deciduous Association*

233.3 Californian Deciduous Swamp and Riparian Scrub

233.31 Mixed Narrowleaf Series*

233.311 Salix lasiolepis Association*

234 Tropical-Subtropical Swamp and Riparian Scrub

234.1 Caribbean Interior Swampscrub

234.2 Caribbean Maritime Swampscrub

234.3 Tamaulipan Interior Swampscrub

234.4 Tamaulipan Maritime Swampscrub

234.5 Sinaloan Interior Swamp and Riparian Scrub

234.51 Mixed Evergreen Series*

234.511 Vallesia glabra-Baccharis glutinosa-Salix bonplandiana Association*

234.6 Sinaloan Maritime Swampscrub

234.61 Mangrove Series*

234.611 Avicennia germinans Association*

234.612 Rhizophora mangle Association*

234.7 Sonoran Deciduous Swamp and Riparian Scrub

234.71 Mixed Scrub Series*

234.711 Prosopis pubescens-Prosopis juliflora torreyana-Pluchea sericea Association*

234.72 Saltcedar Disclimax Series*

234.721 Tamarix chinensis Association*

234.722 Tamarix chinensis -mixed scrub Association*

240 Marshland Formation

241 Arctic-Boreal Marshlands

241.1 Polar (High Arctic) Marshland

241.2 Greenlandian Marshland

241.3 Alaskan Maritime (Coastal) Marshland

241.4 Canadian Interior Marshland

241.5 Canadian Maritime (Coastal) Marshland

241.6 Adirondack-Appalachian Alpine and Subalpine Marshland

241.7 Rocky Mountain Alpine and Subalpine Marshland

241.71 Rush Series*

241.711 Juncus balticus Association*

241.72 Manna Grass Series*

241.721 Glyceria borealis Association*

241.8 Sierran-Cascade Alpine and Subalpine Marshland

241.81 Rush Series*

241.811 Juncus spp. Association*

242 Cold Temperate Marshlands

242.1 Northeastern Interior Marshland

242.2 Northeastern Maritime (Coastal) Marshland

242.3 Plains Interior Marshland

242.31 Rush Series*

252.311 Juncus tenuis Association*

242.32 Bur-reed Series*

242.321 Sparganium angustifolium Association*

242.33 Cattail Series*

242.331 Typha latifolia Association*

242.34 Bulrush Series*

242.341 Scirpus validus Association*

242.4 Rocky Mountain Montane Marshland

242.41 Rush Series*

242.411 Juncus saximontanus Association*

242.5 Great Basin Interior Marshland

242.51 Rush Series*

242.511 Juncus spp. Association*

242.52 Saltgrass Series*

242.521 Distichlis stricta Association*

242.6 Sierran-Cascade Montane Marshland

242.61 Rush Series*

242.611 Juncus spp. Association*

242.7 Pacific Coastal (Oregonian) Interior Marshland

242.8 Pacific Coastal (Oregonian) Maritime Marshland

243 Warm Temperate Marshlands

243.1 Southeastern Interior Marshland

243.2 Southeastern Maritime Marshland

243.3 Chihuahuan Interior Marshland

243.31 Saltgrass Series*

243.311 Distichlis stricta Association*

243.4 Mohavian Interior Marshland

243.41 Rush Series*

243.411 Juncus cooperi Association*

243.42 Saltgrass Series*

243.421 Distichlis stricta Association*

243.5 Madrean Marshland

243.51 Rush Series*

243.511 Juncus mexicanus Association*

* Examples only.


303
 

243.6 Californian Interior Marshland

243.61 Cattail Series*

243.611 Typha latifolia Association*

243.612 Typha domingensis Association*

243.7 Californian Maritime Marshland

243.71 Cordgrass Series*

243.711 Spartina foliosa Association*

243.72 Glasswort Series*

243.721 Salicornia virginica Association*

244 Tropical-Subtropical Marshland

244.1 Caribbean Interior Marshland

244.2 Caribbean Maritime Marshland

244.3 Tamaulipan Interior Marshland

244.4 Gulf Coast Maritime Marshland

244.5 Sinaloan Interior Marshland

244.51 Cattail Series*

244.511 Typha domingensis Association*

244.6 Sinaloan Maritime Marshland

244.61 Glasswort Series*

244.611 Salicornia spp. Association*

244.7 Sonoran Interior Marshland

244.71 Cattail Series*

244.711 Typha domingensis Association*

244.72 Giant Reed Series*

244.721 Phragmites communis Association*

244.73 Bulrush Series*

244.731 Scirpus americanus Association*

244.74 Threesquare Series*

244.741 Scirpus olneyi Association*

244.8 Sonoran Maritime Marshland

244.81 Saltgrass Series*

244.811 Distichlis stricta Association*

244.82 Glasswort Series*

244.821 Salicornia spp. Association*

250 Strand Formation

251 Arctic-Boreal Strands

251.1 Polar Maritime Strand

251.2 Greenlandian Strand

251.3 Alaskan Maritime Strand

251.4 Canadian Interior (Stream and Lake) Strand

251.5 Canadian Maritime Strand

251.6 Adirondack-Appalachian Alpine and Subalpine Stream and Lake Strand

251.7 Rocky Mountain Alpine and Subalpine Stream and Lake Strand **

251.8 Sierran-Cascade Alpine and Subalpine Stream and Lake Strand **

252 Cold Temperate Strands

252.1 Northeastern Interior (Stream and Lake) Strand

252.2 Northeastern Maritime Strand

252.3 Plains Interior (Stream and Lake) Strand

252.31 Annual Series**

252.311 Xanthium saccharatum -mixed annual Association*

252.4 Rocky Mountain Montane Stream and Lake Strand*

252.41 Annual Series**

252.5 Great Basin Interior Strand*

252.51 Annual Series**

252.6 Sierran-Cascade Montane Stream and Lake Strand*

252.61 Annual Series**

252.7 Pacific Coastal (Oregonian) Interior Strand

252.8 Pacific Coastal (Oregonian) Maritime Strand

253 Warm Temperate Strands

253.1 Southeastern Interior Strand

253.2 Southeastern Maritime Strand

253.3 Chihuahuan Interior Strand

253.31 Annual Series*

253.311 Xanthium saccharatum Association*

253.4 Mohavian Interior Strand

253.41 Annual Series*

253.411 Xanthium saccharatum Association*

253.42 Mixed Scrub Series*

253.421 Tamarix chinensis -mixed shrub Association*

253.5 Madrean Stream and Lake Strand

253.51 Annual Series*

253.511 Mixed annual Association*

253.6 Californian Stream and Lake Strand

253.61 Annual Series*

253.611 Nicotiana attenuata Association*

253.7 Californian Maritime Strand

253.71 Mixed Scrub Series*

253.711 Abronia maritima-Atriplex leucophylla-Cakile maritima et al. Association*

253.72 Sea-grass Series*

253.721 Phyllospadix scouleri Association*

253.73 Green Algae Series*

253.731 Ulva californica Association*

253.74 Brown Algae Series*

253.741 Pelvetia fastigiata Association*

253.75 Red Algae Series*

253.751 Gigartina canaliculata Association*

254 Tropical-Subtropical Strands

254.1 Caribbean Interior Strand

254.2 Caribbean Maritime Strand

254.3 Tamaulipan Interior Strand

254.4 Gulf Coast (Tamaulipan) Maritime Strand

254.5 Sinaloan Interior Strand*

254.51 Annual Series**

254.6 Sinaloan Maritime Strand**

254.61 Mixed Scrub Series*

254.7 Sonoran Interior Strand

254.71 Mixed Scrub Series*

254.711 Baccharis glutinosa-Solanum nodiflorum-Nicotiana spp.-Rumex hymenosepalus et al. Association*

254.72 Annual Series*

254.721 Amaranthus palmeri Association*

254.8 Sonoran Maritime Strand

254.81 Mixed Scrub Series

254.811 Abronia maritima-Helianthus niveus-Jouvea pilosa et al. Association*

260 Submergent Aquatic Vegetation

261 Arctic-Boreal Submergent Aquatics

261.1 Polar Marine Submergents

261.2 Greenlandian Inland Submergents

261.3 Alaskan Marine Submergents

261.4 Canadian Inland Submergents

261.5 Canadian Marine Submergents

261.6 Adirondack-Appalachian Alpine and Subalpine Submergents

261.7 Rocky Mountain Alpine and Subalpine Submergents

261.71 Pondweed Series*

261.711 Potamogeton natans Association*

261.8 Sierran-Cascade Alpine and Subalpine Submergents

261.81 Pondweed Series*

261.811 Potamogeton alpinus Association*

262 Cold Temperate Submergent Aquatics

* Examples only.

* Our incomplete knowledge of these biotic communities precludes presentation of representative fifth (series) and sixth level (association) examples.


304
 

262.1 Northeastern Inland Submergents

262.2 Northeastern Marine Submergents

262.3 Plains Inland Submergents

262.31 Pondweed Series*

262.311 Potamogeton foliosus Association*

262.4 Rocky Mountain Montane Submergents

262.41 Pondweed Series*

262.411 Potamogeton foliosus Association*

262.5 Great Basin Inland Submergents

262.51 Pondweed Series*

262.511 Potamogeton foliosus Association*

262.6 Sierran-Cascade Montane Submergents

262.61 Pondweed Series*

262.611 Potamogeton pusillus Association*

262.7 Pacific Coastal (Oregonian) Inland Submergents

262.8 Pacific Coastal (Oregonian) Marine Submergents

263 Warm Temperate Submergent Aquatics

263.1 Southeastern Inland Submergents

263.2 Southeastern Marine Submergents

263.3 Chihuahuan Inland Submergents

263.31 Pondweed Series*

263.311 Potamogeton pectinatus Association*

263.4 Mohavian Inland Submergents

263.41 Pondweed Series*

263.411 Potamogeton pectinatus Association*

263.5 Madrean Inland Submergents

263.51 Pondweed Series*

263.511 Potamogeton pectinatus Association*

263.6 Californian Inland Submergents

263.61 Pondweed Series*

263.611 Potamogeton pectinatus Association*

263.62 Milfoil Series*

263.621 Myriophyllum exalbescens Association*

263.7 Californian Marine Submergents

263.71 Ruppia Series*

263.711 Ruppia maritima Association*

263.72 Eelgrass Series*

263.721 Zostera marina Association*

263.73 Giant Kelp Series*

263.731 Macrocystis pyrifera Association*

263.74 Feather-boa kelp Series*

263.741 Egregia laevigata Association*

263.75 Southern Sea Palm Series*

263.751 Eisenia arborea Association*

264 Tropical-Subtropical Submergent Aquatics

264.1 Caribbean Inland Submergents

264.2 Caribbean Marine Submergents

264.3 Tamaulipan Inland Submergents

264.4 Gulf Coastal Marine Submergents

264.5 Sinaloan Inland Submergents

264.51 Pondweed Series**

264.6 Sinaloan Marine Submergents*

264.61 Phytoplankton Series**

264.7 Sonoran Inland Submergents**

264.71 Pondweed Series*

264.711 Potamogeton pectinatus Association*

264.72 Milfoil Series*

264.721 Myriophyllum brasiliense Association*

264.8 Sonoran Marine Submergents

264.81 Ruppia Series*

264.811 Ruppia maritima Association*

264.82 Eelgrass Series*

264.821 Zostera marina Association*

* Examples only.

* Our incomplete knowledge of these biotic communities precludes presentation of sixth level (association examples.)

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Evolution and Riparian Systematics1
 

Preferred Citation: Warner, Richard E., and Kathleen M. Hendrix, editors California Riparian Systems: Ecology, Conservation, and Productive Management. Berkeley:  University of California Press,  c1984 1984. http://ark.cdlib.org/ark:/13030/ft1c6003wp/