ITINERA GEOBOTANICA 12 1999

ITINERA GEOBOTANICA VOLUMEN 12 Fecha: 30 de Junio de 1999 ASOCIACION ESPAÑOLA DE FITOSOCIOLOGIA (AEFA) FEDERATION INTERNATIONALE DE PHYTOSOCIOLOGIE (FIP) ITINERA GEOBOTANICA es una publicación periódica de la Asociación Espola de Fitosociología (AEFA), adherida a la Federación Internacional de Fitosociología (FIP), en la que se darán a conocer monografías fitosociológicas itinerantes, pródromos biogeográficos y bioclimáticos y otros temas geobotánicos de interés regional o global Su difusión se asegurará a través del Servicio de Publicaciones de la Universidad de León ITINERA GEOBOTANICA is a journal of the Spanish Phytosociological Association (AEFA), which is adhered to the International Federation of Phytosociology (FIP), which will publish monographic works of phytosociological itineraries, biogeographic and bioclimatical prodromi, and other geobotanical works of regional or global interest Its difussion is guarateed by the University of León publishing service Editors (Editores) S Rivas-Martínez, Madrid, E T.E Díaz, Oviedo, E A Penas, León, E F Fernández, Madrid, E Editorial Board (Comisión editorial) F Alcaraz, Murcia, E A Asensi, Málaga, E J C Báscones, Pamplona, E C Blasi, Roma, I M Costa, Valencia, E J M Géhu, Bailleul, F J Loidi, Bilbao, E J Molero, Granada, E M Peinado, A Henares, E P Quézel, Marsella, F D Sánchez-Mata, Madrid, E F Valle, Granada, E J Amigo, Santiago de C., E M Barbero, Marsella, F A Benabid, Rabat, M O de Bolòs, Barcelona, E S Dafis, Tesalónica, GR J Izco, S Compostela, Es M Lousa, Lisboa, P G Navarro, Santa Cruz, Bo P.L Pérez de Paz, La Laguna, E F Roig, Mendoza, A J.P Theurillat, Ginebra, CH W Wildpret, La Laguna, E M del Arco, La Laguna, E M Barbourg, Davis, US E Biondi, Ancona, I E Box, Atlanta, US J A Fernández, Oviedo, E M Ladero, Salamanca, E L Llorens, P de Mallorca, E F Pedrotti, Camerino, I R Pott, Hannover, D P Sánchez, Murcia, E C Valle, Salamanca, E Technicals Editors (Responsables de la edición) L Herrero, León, E E Puente, Ln, E M.E García, Ln, E S del Río, Ln, E Subscription & Exchange information: Departamento de Biología Vegetal (Botánica) Facultad de Biología Campus Vegazana s/n Universidad de Ln 24071 - LEON (ESPA) Email: dbvlhc@unileon.es ISSN: 0213-8530 Depósito Legal: LE - 729 - 1987 Impreso en Gráficas CELARAYN S.A Editado por el Servicio de Publicaciones de la Universidad de León Publicado el 30 de Junio de 1999 NORTH AMERICAN BOREAL AND WESTERN TEMPERATE FOREST VEGETATION (Syntaxonomical synopsis of the potential natural plant communities of North America, II) Salvador Rivas-Martínez, Daniel Sánchez-Mata &Manuel Costa Itinera Geobotanica 12: 5-316 (1999) NORTH AMERICAN BOREAL AND WESTERN TEMPERATE FOREST VEGETATION (Syntaxonomical synopsis of the potential natural plant communities of North America, II) Salvador Rivas-Martínez (1, 2), Daniel Sánchez-Mata (2) & Manuel Costa (3) Abstract Natural potential vegetation broadly distributed throughout the Boreal and Temperate North American territories is globally studied, basically following the Braun-Blanquet phytosociological methodology Potential coniferous forests and their respective deciduous secondary woodlands are characterized and systematized according to bioclimatic features, biogeographic distribution, as well as floristic and dynamic patterns All the potential climactical plant communities are included in a broad, and newly-proposed phytosociological class: Linnaeo americanae-Piceetea marianae, which comprises eight orders, nineteen alliances and sixty-four associations This monograph also includes a comprehensive compilation of bioclimatic and biogeographic syntheses: The Worldwide Bioclimatic Classification System, the Worldwide Biogeographic Approach at regional level, the Synoptic Biogeographic Approach of America at provincial level and the Synoptic Biogeographic Approach of North America North of Mexico at subsectorial level Several thematic maps are given (including an attached and newly-proposed biogeographic map of North America), as well as some original vegetation transections (catenas), various original keys to identify the diversity of potential plant communities, a checklist of the characteristic species and plant names used and finally, the bibliographic references This monograph on North America Boreal and Western Temperate potential natural vegetation follows that previously published (Itinera Geobot 10: 5-148 1997) on Mediterranean and Tropical vegetation in Western and South-Western United States North of Mexico (Heteromelo arbutifoliae-Quercetea agrifoliae, Calocedro decurrentis-Pinetea jeffreyi, Prosopido torreyanae-Fouquierietea splendentis and Artemisio tridentatae-Juniperetea osteospermae classes) Keywords: Bioclimatology, Biogeography, Phytosociology, North American Bioclimatology, NA Biogeography, NA Syntaxonomy, NA Boreal and Temperate Forest Vegetation Communities Introduction Bioclimatic, geobotanic and biogeographic notions Materials and methods 33 Discussion and results Description of plant communities 35 Vegetation transections and bioclimatic diagrams 215 Checklist of the characteristic species and plant names 249 Climatic and bioclimatic data 277 Resumen (in Spanish) 301 Acknowledgements 303 References 305 Index of syntaxa names .315 (1) PHYTOSOCIOLOGICAL RESEARCH CENTER J.M Usandizaga, 46 E-28409 Los Negrales, Madrid Tel.: (34) 91 851 1529 Fax: (34) 91 851 1873 Email: rivas-martinez.cif@tsai.es, srivas@eucmax.sim.ucm.es (2) DEPARTAMENTO DE BIOLOGÍA VEGETAL II (Botánica) Facultad de Farmacia, Universidad Complutense E28040 Madrid Tel.: (34) 91 394 1769 Fax: (34) 91 394 1774 Email: dsmata@eucmax.sim.ucm.es (3) JARDÍN BOTÁNICO, Beato Gaspar de Bono, Universidad de Valencia E-46008 Valencia Tel.: (34) 96 391 16 57 Fax: (34) 96 392 2823 Email: manuel.costa@uv.es Itinera Geobotanica 12 (1999) INTRODUCTION The considerable economic value of the forests as a natural resource in the United States of America and Canada and the centralization of their management in the respective Departments of Agriculture was the reason for the development of a scientific understanding of their vaste forests Within a short period of time, a close balance was achieved between explotation, regeneration and conservation of these resources In consequence, the development of the knowledge of dendroflora and forest management and the increase in the autoecological and ecophysiological studies has been an unexpected reality The vegetation typology approaches, mainly physiognomic-ecological and bioclimatic are a very important ensemble of studies and publications which have been synthesized in the valuable and veteran work of Küchler (1964) in the Conterminous of United States, in the recent Canadian monographs of Pojar & al (1987) and Klinka & al (1996), and mainly through the inestimable chapters compiled in the book ‘North American Terrestrial Vegetation’ by Barbour & Billings (1988) The studies and synthetic publications on the Boreal and Temperate vegetation of North America were initiated more than sixty years ago with Halliday (1937) and Weaver & Clements’ (1938) publications The influence of the Braun-Blanquet phytosociological approach was evident in the analysis and synthesis of numerous studies and publications: Braun-Blanquet, Sissingh & Vlieger (1939), Knapp (1957, 1965), Grandtner (1960, 1966), Krajina (1959, 1960, 1965), Damman (1964), etc However, for the last twenty years a global synopsis on North American potential natural vegetation following the BraunBlanquet phytosociological methodology has been in progress Within the publications involved in this project the works of Looman (1979, 1983/1987), Miyawaki, Iwatsuki & Grandtner (1994), Peinado & al (1994, 1997/1998) and Rivas-Martínez (1997) should be mentioned as important Our present work is a continuation of the previously published ‘Syntaxonomical synopsis of the potential natural plant communities of North America, I’ [Rivas-Martínez in Itinera Geobot 10: 5-148 (20.06.1997)] and a part of a ‘Global Bioclimatics’ project started in South America in 1977 In our first syntaxonomic compilation of climactical vegetation of North America we published four phytosociological classes, including the North American Mediterranean and Tropical potential climactical vegetation of South United States North of Mexico: I Heteromelo arbutifoliae-Quercetea agrifoliae (evergreen sclerophyllous and late-deciduous, oceanic, thermo- and mesomediterranean mesoforests, woodlands and thicket communities of the Californian Region), II Calocedro decurrentis-Pinetea jeffreyi (coniferous and mixed evergreen or broad-leaved deciduous, oceanic, supra-and oromediterranean forests of the Californian Region), III Prosopido torreyanaeFouquierietea splendentis (microphyllous-deciduous and succulent, xerophytic, thermoand supratropical open scrublands to closed microforests of the Mexican Xerophytic Region), IV Artemisio tridentatae-Juniperetea osteospermae (xerophytic, xeric and desertic continental, supra- and oromediterranean coniferous microforests, juniper-pinyon wood- Rivas-Martínez, Sánchez-Mata & Costa North American Forest Vegetation lands, and wooded or desertic steppe communities of the Great Basin Region) In this second part we focus on the potential natural Boreal and Temperate coniferous forests, broadly distributed throughout the territories of the North American Boreal and Rocky Mountain Regions Moreover, we also include their evergreen, riparian and deciduous secondary forests (V Linnaeo americanae-Piceetea marianae) BIOCLIMATIC, GEOBOTANIC AND BIOGEOGRAPHIC NOTIONS For an easier comprehension and use of this publication some bioclimatic, geobotanic and biogeographic simple syntheses and definitions are included [additional inedit and published information about these topics can be found in Rivas-Martínez (1996, 1997) and in ‘Global Bioclimatics’ (in progress)] BIOCLIMATIC PARAMETERS AND INDEXES In our Bioclimatic approach (Rivas-Martínez, in progress), which is almost a new generic climate worldwide classification, only easily available statistical and meteorological data have been used Those data have been treated as climatic parameters (single and summatory) or as bioclimatic indexes (combinations of parameters) both obtained through intentionally easy arithmetic formulas Both parameters and indexes have only been used when some thresholds have shown an accurate relation with the vegetation changes (boundaries of potential natural plant communities) and have a high predictive value The attached table summarizes the latest approach (16.12.98) to the Worldwide Bioclimatic Classification System (see pg 9) T: Yearly average temperature in centigrade degrees (Celsius) Ti: Average temperature any month of the year M: Average maximum temperature of the coldest month of the year (Tmin) m: Average minimum temperature of the coldest month of the year (Tmin) Tmin: Average temperature of the coldest month Tmax: Average temperature of the warmest month P: Yearly average precipitation in mm Pi: Average precipitation any month of the year Ps: Summer Precipitation Precipitation of the driest quarter of the year in mm (Pd) Total average precipitation of the three summer months measured in mm: In extratropical zones (north of 23ºN and south of 23ºS) precipitations of June + July + August for the Northern Hemisphere, and for December + January + February for the Southern Hemis- Itinera Geobotanica 12 (1999) phere Within tropical belts, this is understood as the ‘estival precipitation’, that is: the sum of precipitation occurring during the three driest consecutive months of the year, regardless of their average temperature Pw: Winter Precipitation (mm) In extratropical zones (north of 23ºN and south of 23ºS) precipitations of December + January + February for the Northern Hemisphere, and of June + July + August for the Southern Hemisphere Pp: Yearly Positive Precipitation In mm, total average precipitation of those months whose average temperature is higher than 0ºC Tp: Yearly Positive Temperature In tenths of degrees Celsius, sum of the monthly average temperature of those months whose average temperature is higher than 0ºC Tps: Positive Summer Temperature Positive temperature of summer quarter of the year Athermic: The average temperature of every month of the year is 0ºC or lower than 0ºC: consequently, Tp is null Ic: Continentality Index (yearly thermic interval) Ic = Tmax - Tmin In degrees Celsius, the number expressing the range between the average temperatures of the warmest (Tmax) and coldest (Tmin) months of the year The simple continentality index-types and subtypes are: hyperperoceanic (0-11) [extremely hyperperoceanic (0-3), euhyperperoceanic (3-7), barely hyperperoceanic (7-11)], oceanic (11-21) [euoceanic (11-18), semicontinental (18-21)], and continental [subcontinental (21-28), eucontinental (28-45) and hypercontinental (45-65)] Types HYPEROCEANIC (Ic 0-21) OCEANIC (Ic 11-21) CONTINENTAL (Ic 21-65) Subtypes Ic Extremely hyperoceanic Euhyperoceanic Barely hyperoceanic Euoceanic Semicontinental 0-3 3-7 7-11 11-18 18-21 Subcontinental Eucontinental Hypercontinental 21-28 28-46 46-65 Io: Ombrothermic Index Io = (Pp/Tp) 10 Ten times the quotient resulting value between the yearly positive precipitation in mm (Pp) and the yearly positive temperature (Tp) (see ‘Pp’ and ‘Tp’ above) Iosi: Ombrothermic index of any month of summer quarter (Tr3: June, July and August, in the Northern Hemisphere; and December, January and February, in Southern Hemisphere) Rivas-Martínez, Sánchez-Mata & Costa North American Forest Vegetation Ios2: Ombrothermic index of the warmest bimonth of the summer quarter (Tr3): [Ios2 = (Pps2/Tps2) 10] (Tr3: June, July and August, in the Northern Hemisphere; and December, January and February, in Southern Hemisphere) Iod2: Ombrothermic index of dryest bimonth within the dryest quarter of the year [Iod2 = (Ppd2/Tpd2) 10] It: Thermicity Index It = (T + m + M) 10 Ten times the sum of T (yearly average temperature), m (average minimum temperature of the coldest month of the year), M (average maximum temperature of the coldest month of the year) Coldest month of the year: the one which has the lowest monthly average temperature (Tmin) Itc: Compensated Thermicity Index (Itc) Itc = It ± C In the extratropical zones of the World (northern and southern 27ºN and 27ºS parallels, respectively), the Compensated Thermicity Index is designed to equilibrate the cold ‘excess’ that occurs during winter in the continental climates (average tenperature of the coldest month of the year minimum), or the excessively mild winter in the marked oceanic territories, so that these index values can be significantly compared If the Continentality Index (Ic) lies between and 18, the Itc value is considere equal to the It value, that means that there is no modification In the other hand, if the Continentality Index not reach, or surpass, the mentioned values, it is needed to compensate the Thermicity Index adding or subtracting a figure called Compensation Value (C) In the extratropical markedly hyperoceanic zones (Ic < 9.0), the Compensation Value (C) is calculated by multiplying by ten the result of the difference between 9.0 and the simple Ic: C = (9.0-IC) 10 This Compensation Value is then substracted from the Thermicity Index corresponding value: Itc = It – C In the extratropical continental climates (Ic > 18.0) the Compensation Value (C) is a summatory to the Thermicity Index corresponding value: Itc = It + C This Compensation Value (C) must be calculated in dependence of the Continentality Index figure (Ic) So, if the continentality is moderate (18.0 < Ic ≤ 21.0) the Compensation Value (Cl) is obtained by multiplying by fl (fl = 5) the result of the difference between the Ic of the station and 18 When the continentality is high (Ic > 21.0), the Compensation Value must be calculated by means of a total whose partial values (Cl, C2, C3, C4) are proportionally higher due to the increment of the multiplier value (fi) as a function of the increasing continentality Therefore: Itc= It+ (C1+C2+C3+C4) The compensation values of application, as a function of the Continentality Index values (Ic) and of the multiplication factor (fi), are obtained as follows: Ic fi 18 < Ic ≤ 21 f1 = 21 < Ic ≤ 28 f2 = 15 Ci Ci highest values Ci = C1; C1 = f1 (Ic - 18) Ci = C1 + C2; C1 = f1 (21 - 18) = 15; C2 = f2 (Ic - 21) C2 = 105 C1 = 15 Ci = C1 + C2 + C3; C1 = 15; C2 = f2 (28 - 21) = 105; C3 = f3 (Ic - 28) C3 = 450 28 < Ic ≤ 46 f3 = 25 46 < Ic ≤ 65 f4 = 30 Ci = C1 + C2 + C3 + C4; C1 = 15; C2 = 105; C3 = f3 (46 - 28) = 425; C4 = f4 (Ic - 46) C4 = 570 10 Itinera Geobotanica 12 (1999) Pcm1: Precipitation of warmest quarter of the year [Pcm1= ΣPi3Trh] (Trh: the hottest quarter of the year) Pcm2: Precipitation of the quarter following the warmest quarter of the year [Pcm2= ΣPi3Tr following h] Pcm3: Precipitation of the quarter previous to warmest quarter of the year [Pcm3= ΣPi3Tr previous h] Ioe: Ombro-Evapotranspiration Index The quotient resulting value between the yearly positive precipitation in mm and the value of Thornthwaite yearly evapotranspiration [Ioe=Pp/PE] Iosc: Summer compensated ombrothermic indexes (Iosc: Ios2, Ios3, Ios4) By definition, the Mediterranean macrobioclimate is an extratropical macrobioclimate characterized by, at least, two consecutive dry months during the summer (the warmest period in the year) A month is defined as dry if the precipitation (mm) is less than twice the temperature (centigrade degrees) (Pi < 2Ti) Obviously, if the ombrothermic bimonthly quotient of July + August (Ios2) or of January + February, depending on the hemisphere, (Ps July+August)/(Ts July+ August), is higher than two, the territory is not Mediterranean; but if that quotient is less than 2.0, the territory may or may not Mediterranean, as the bimonthly deficient hydrical balance may or may not compensated with the previous month’s precipitation (June or December: Ios3) A quarterly quotient Ios3 of less than 2.0 could even be compensated in some cases (see ‘Compensation Table’) with the previous month’s precipitation (May or November: Ios4) These summer compensated ombrothermic indexes are very useful as they discriminate the frontier between Mediterranean-Temperate and Mediterranean-Boreal territories In some extratropical places, the warmest months in the year are not those that follow the summer solstice, but those near the next autumnal equinox and, in these cases, those late summer hottest month must be used in calculations Io Ios2 Ios3 Ios4 2.0-3.6 3.6-4.8 4.8-6.0 6.0-8.0 8.0-10.0 10.0-12.0 > 12.0 > 1.9 > 1.8 > 1.7 > 1.5 > 1.2 > 0.7 - > 1.9 > 1.9 > 1.8 > 1.8 > 1.6 > 1.4 - > 2.0 > 2.0 > 2.0 > 2.0 > 2.0 > 2.0 > 2.0 Figure Summer ombrothermic compensation values (Ios2, Ios3) from Mediterranean to Temperate macrobioclimates (Submediterranean bioclimatic variant) It will be Temperate if Ios2 > 2.0 or Iosc3 and Iosc4 > 2.0, with Ios2 and Ios3 inside the threshold values expressed in the table 302 Itinera Geobotanica 12 (1999) órdenes admitidos, así como numerosas tablas sinópticas información bioclimática y biogeográfica Las tablas fitosociológicas correspondientes a las asociaciones reconocidas incluyen únicamente una cuidada selección de los inventarios fitosociológicos que hemos realizado personalmente en Norteamérica, indicándose en cada caso los datos precisos de la localidad y fecha de realización, según el registro existente en el banco de datos del CIF (“Centro de Investigaciones Fitosociológicas”) En el caso de tipos nomenclaturales (bien inventarios propios o bibliográficos), se transcribe en el texto el inventario correspondiente indicándose su procedencia y referencia precisa, los datos bioclimáticos, geográficos y biogeográficos de la localidad en cada caso, así como la fecha de su realización y los autores del mismo cuando se trata de inventarios propios Las unidades sintaxonómicas admitidas se relacionan en el correspondiente esquema tipológico Para toda la vegetación Boreal y Templada de Norteamérica proponemos una nueva clase fitosociológica de muy amplia jurisdicción: Linnaeo americanaePiceetea marianae, quizás algo heterogénea y posiblemente desmembrable en varias clases en un futuro En ella incluimos todos los bosques desarrollados en territorios boreales y templados, que pueden alcanzar algunas áreas mediterráneas así como la vegetación forestal secundaria que los reemplaza en estaciones alteradas bien por causas naturales o antrópicas (a veces carácter de comunidades permanentes edafo-higrófilas) y las formaciones de abedules arbustivos climácicas en ciertas áreas árticas, boreales u orotempladas) La nueva clase propuesta ahora (Linnaeo americanae-Piceetea marianae ) incluye ocho órdenes, diecinueve alianzas y sesenta y cuatro asociaciones Los ocho órdenes propuestos son: Gaultherio-Piceetalia (bosques climácicos de coníferas de la región Boreal Norteamericana), Tsugetalia mertensiano-heterophyllae (bosques climácicos de coníferas de la subregión Noroccidental Pacífica Norteamericana), Alno rubrae-Populetalia trichocarpae (bosques deciduos de sustitución o riparios de la subregión Noroccidental Pacífica Norteamericana), Pseudotsugo glaucae-Abietetalia bifoliae (bosques climácicos de coníferas de la subregión Rocosiana), Betulo papyriferae-Populetalia tremuloidis (bosques de sustitución deciduos de la región Boreal Norteamericana), Populetalia angustifoliodeltoidis (bosques riparios deciduos de Norteamérica: templados, mediterráneos y subtropicales), Populetalia fremontii (bosques riparios deciduos de Norteamérica: mediterráneos y tropicales xérico-desérticos) y, por último, Ledo decumbentis-Betuletalia glandulosae (tundra de abedules y sauces chaparros de Norteamérica: árticos, boreales y orotemplados) La distribución biogeográfica general de estos órdenes es: Gaultherio-Piceetalia (región Boreal Norteamericana y región Atlántica Norteamericana), Tsugetalia mertensianoheterophyllae (subregión Pacífico-Noroeste) y Pseudotsugo glaucae-Abietetalia bifoliae (subregión Rocosiana), Alno rubrae-Populetalia trichocarpae (subregión PacíficoNoroeste), Betulo papyriferae-Populetalia tremuloidis (región Boreal Norteamericana), Populetalia angustifolio-deltoidis (subregión Rocosiana y región Atlántica Norteamericana), Populetalia fremontii (región Californiana y región Gran Cuenca) y Ledo decumben- Rivas-Martínez, Sánchez-Mata & Costa North American Forest Vegetation 303 tis-Betuletalia glandulosae (región Artica Norteamericana, región Boreal Norteamericana y región Rocosiana) Las alianzas consideradas en cada orden son: Gaultherio-Piceetalia: Pinion banksianae, Gaultherio procumbentis-Piceion glaucae, Piceion rubentis y Ledo decumbentisPiceion marianae; Tsugetalia mertensiano-heterophyllae: Tsugion heterophyllae, Tsugion mertensianae y Pinion contortae; Alno rubrae-Populetalia trichocarpae: Alno rubraePopulion trichocarpae; Pseudotsugo glaucae-Abietetalia bifoliae: Piceo engelmanniiAbietion bifoliae y Pino scopulorum-Pseudotsugion glaucae; Betulo papyriferaePopuletalia tremuloidis: Betulo neoalaskanae-Populion tremuloidis, Betulo cordifoliaePopulion tremuloidis, Alnion rugosae; Populetalia angustifolio-deltoidis: Populion angustifolio-moniliferae y Platano occidentalis-Populion deltoidis (prov.); Populetalia fremontii: Populion fremontii y Populion mesetae (prov.); Ledo decumbentis-Betuletalia glandulosae: Salici pulchrae-Betulion glandulosae y Betulion minoris (prov.) ACKNOWLEDGEMENTS We would like to thank many people (colleagues, research assistants, techniques, reviewers etc.), as well as several private and public institutions for their help during our work in America First of all, to Spanish public institutions which have supported and funded our research in North America at different times: The Ministry of Science and Education (Madrid) through different research programs in US, and the Universidad Complutense (Madrid) through its ‘Gregorio del Amo’ research fund program at the University of California; the Spanish ‘Phytosociological Research Center’ (CIF, Madrid) also supported our research by making its considerable scientific resources available to us We would also like to thank all our colleagues at the Departamento de Biología Vegetal II (Universidad Complutense) in Madrid for their help and understanding Of the North American Institutions we are grateful to: The US Department of Interior (National Parks Service) and the US Department of Agriculture (USDA, Forest Service) for their assistance with our field works throughout the US territories and for their help with some technical publications; the Missouri Botanical Garden (Saint Louis) gave us technical and bibliographic assistance during our stays; University of California at Davis (Departments of Environmental Horticulture and Plant Biology) offered us their resources during our stays; the UC Davis Shields Library made all its great bibliographic collections, research assistance and resources available to us ; the Davis Herbaria Society and the University of California at Berkeley helped us by giving us access to their herbaria for our taxonomic studies in the Davis Herbaria and Jepson Herbarium collections We would also like to thank the following individuals: Those that shared with us the research on several field trips throughout North American Boreal and Temperate territories: A Asensi, M.G Barbour, F Fernández-González, M Grandtner, M Ladero, L Llorens, J 304 Itinera Geobotanica 12 (1999) Molero, J.A Molina, G Navarro, A Penas, C Reynal, P Rodríguez-Rojo, S Sardinero, J Thieret and F Valle; B Díez Garretas, A Asensi and F Fernández-González ordered the phytosociological data from Alaska; S Sardinero transferred some data on Mount Washington (New Hampshire) forest vegetation to us (Sorbo americanae-Betuletum cordifoliae); A Penas (North America), G Navarro (South America) and J Giménez de Azcárate (Mexico) have collaborated in the biogeographic proposals; E Dean (Curator of the UC Davis Herbaria), J Shepard (UC Davis Herbaria) and the Curator of the Jepson Herbarium (Berkeley) helped us with our taxonomic research; C Ansseau (Québec), M G Barbour (Davis), M Chýtrý (Pruhonice), C García Navarro (Davis), R.G Gavilán (Athens), M Grandtner (Québec), P Rodríguez-Rojo (Davis) and F.J Rueda (Davis) helped us with bibliographic research; A Penas, M.A Luengo and S Rivas Sáenz, supplied some bioclimatic data and made the adequate computer bioclimatic programs; A Penas, J Cortizo and I Prieto adapted and drew the biogeographic subsectorial map of North America; J.M Pizarro drawing the distribution maps of the phytosociological orders as well as the bioclimatic and biogeographic maps; M.G Barbour and M.L López Fernández made some comments, suggestions and gave us some constructive criticism about our proposals; C Hontana carried out the computer treatment and design of our handwritten texts; and, finally, M.L López Fernández, F Llamas and Pru Brooke-Turner assisted us in the English version Everybody, once again, thanks Rivas-Martínez, Sánchez-Mata & Costa North American Forest Vegetation 305 REFERENCES Achuff, P.L (1989) Old-growth forests of the Canadian Rocky Mountain national parks Nat Areas J 9: 12-26 Achuff, P.L & La Roi, G.H (1977) Picea-Abies forests in the highlands of Northern Alberta Vegetatio 33 (2/3): 127-146 Alcaraz, F (1996) Fitosociología integrada, paisaje y Biogeografía In: Loidi, J (ed.) 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Ordination and classification of communities: 617-626 Handbook of Vegetation Science, Dr Junk Publishers The Hague Whittaker, R.H (1975) Communities and Ecosystems 2nd ed MacMillan Publ 387 p New York Rivas-Martínez, Sánchez-Mata & Costa North American Forest Vegetation 315 INDEX OF SYNTAXA NAMES Abietetum balsameae 56 Abieti amabilis-Piceetum sitchensis 88 Abieti amabilis-Tsugetum heterophyllae 99 Abieti amabilis-Tsugetum mertensianae 106 Abieti balsameae-Piceetum rubentis 66 Abieti grandis-Pseudotsugetum menziesii 96 Abieti lasiocarpae-Pinetum albicaulis 115 Abieti lasiocarpae-Tsugetum mertensianae 108 Aceri macrophylli-Alnetum rubrae 127 Alnetum rugosae .192 Alnion rugosae 191 Alno rubrae-Populetalia trichocarpae 123 Alno rubrae-Populion trichocarpae 126 Alno sinuatae-Populetum trichocarpae 132 Alno tenuifoliae-Populetum trichocarpae 133 Aralio hispidae-Populetum tremuloidis 190 Arctostaphylo patulae-Pinetum flexilis .146 Betulo cordifoliae-Populion tremuloidis 187 Betulo neoalaskanae-Populion tremuloidis 175 Betulo occidentalis-Populetum angustifoliae 200 Betulo papyriferae-Populetalia tremuloidis .172 Carici pensylvanicae-Pinetum banksianae 52 Carici-Alnetum rugosae 193 Empetro hermaphroditi-Abietetum bifoliae 150 Gaultherio procumbentis-Piceion glaucae 56 Gaultherio shallonis-Pseudotsugetum menziesii 93 Gaultherio-Piceetalia 46 Hypno procerrimi-Piceetum glaucae 73 Junipero depressae-Pinetum contortae 122 Kalmio angustifoliae-Piceetum marianae 60 Larici occidentalis-Pseudotsugetum glaucae 164 Ledo decumbentis-Betuletalia glandulosae 205 Ledo decumbentis-Piceion marianae 68 Linnaeo americanae-Piceetea marianae 36 Lonicero ledebourii-Pinetum contortae 120 Menziesio ferrugineae-Abietetum bifoliae 148 Mertensio paniculatae-Abietetum balsameae .64 Oplopanaco horridi-Alnetum rubrae 130 Oryzopsio asperifoliae-Pinetum banksianae 53 Penstemono whippleani-Pinetum aristatae .157 Phyllodoco glanduliflorae-Laricetum lyallii .116 Piceetum engelmannio-albertianae 152 Piceetum glauco-marianae 72 Piceetum pungentis 155 Piceion rubentis 65 Piceo engelmannii-Abietion bifoliae 141 316 Itinera Geobotanica 12 (1999) Piceo engelmannii-Tsugetum heterophyllae .102 Piceo glaucae-Betuletum kenaicae 181 Piceo glaucae-Betuletum neoalaskanae 180 Pinetum banksianae 50 Pinetum banksiano-resinosae .55 Pinion banksianae 49 Pinion contortae .119 Pino scopulorum-Pseudotsugion glaucae 163 Platano occidentalis-Populion deltoidis 201 Platano racemosae-Populetum fremontii 203 Populetalia angustifolio-deltoidis .194 Populetalia fremontii 201 Populetum angustifolio-moniliferae 197 Populetum tremuloidis 189 Populion angustifolio-moniliferae 196 Populion fremontii 202 Populion mesetae 205 Pseudotsugo glaucae-Abietetalia bifoliae 138 Pseudotsugo glaucae-Pinetum flexilis 170 Pulsatillo multifidae-Pinetum latifoliae 77 Rhoo trilobatae-Pinetum scopulorum .169 Ribesio acerifolii-Chamaecyparetum nootkatensis 118 Roso sayi-Populetum tremuloidis .178 Rubo chamaemori-Piceetum marianae .69 Rubo stellati-Piceetum sitchensis .89 Salici barclayi-Betuletum glandulosae .209 Salici longistylis-Alnetum sinuatae 137 Salici pulchrae-Betuletum exilis .212 Salici pulchrae-Betulion glandulosae .207 Shepherdio canadensis-Pseudotsugetum glaucae .167 Sorbo americanae-Betuletum cordifoliae 187 Sphagno-Piceetum marianae 62 Spiraeo lucidae-Populetum balsamiferae 183 Symphoricarpo utahensis-Populetum tremuloidis 185 Tsugetalia mertensiano-heterophyllae 78 Tsugion heterophyllae 83 Tsugion mertensianae .143 Tsugo heterophyllae-Sequoietum sempervirentis 91 Tsugo mertensianae-Abietetum procerae 110 Tsugo mertensianae-Piceetum sitchensis 113 Vaccinio membranacei-Alnetum sinuatae 135 Vaccinio microphylli-Betuletum glandulosae 211 Vaccinio ovati-Piceetum sitchensis .85 Vaccinio scoparii-Abietetum bifoliae .144 Vaccinio scoparii-Pinetum albicaulis .161 Vaccinio scoparii-Pinetum latifoliae 159 ... SUPERREGION 12 Brazilian-Paranean Region 12. 1 Cerrado Province 12. 2 Tocantins Province 12. 3 Beni Province 12. 4 Pantanal Province 12. 5 Atlantic Brazilian Province 12. 6 Paranean Province 12. 7 Caatinga... formations, biocoenosis and plant communities Within almost every bioclimate, a number of 12 Itinera Geobotanica 12 (1999) variations in the seasonal rainfall patterns of rain allows us to recognize... Northern Hemisphere, and for December + January + February for the Southern Hemis- Itinera Geobotanica 12 (1999) phere Within tropical belts, this is understood as the ‘estival precipitation’,