North American boreal and western temperate forest vegetation

Salvador Rivas-Martínez, Daniel Sánchez-Mata & Manuel Costa

Itinera Geobotanica 12:5-316 (1999)


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Description of Plant Communities: Pinion banksianae

1. Pinion banksianae Lavoie 1968 nom. mut.

[Pinion divaricatae Lavoie in Nat. Can. 95: 402. 1968 (lectotype: Pinetum divaricatae Lavoie in Nat. Can. 95: 403. 1968 (art. 20), Vaccinio-Pinion divaricatae Looman in Phytocoenologia 15(3) 298. 1987 (art. 8), Pinion banksianae Lavoie 1968 nom. mut. propos. (art. 45)]

Typus nominis: Pinetum divaricatae Lavoie in Nat. Can. 95: 403. 1968 (lectotype art. 20) [Pinetum banksianae nom. mut. propos. (art. 45)].

North American Continental Boreal; thermo- and mesoboreal, hemiboreal and occasionally upper supratemperate pine meso- and macroforests dominated by Jack pine [Pinus banksiana (syn.: P. divaricata)]. These pine forests are broadly distributed throughout the Western and Eastern Canadian Boreal Sectors, as well as Northwestern Appalachian and Great Lakes Sectors, but they are uncommon in Canadian Prairies Sector and absent in Newfoundlandian Sector areas. The Pinion banksianae associations constitute mainly an ensemble of pyrogenic secondary coniferous forests, but also in Xeric Boreal territories it could represents the wooded climactical and permanent forest plant communities, often with edapho-xerophytic character. They are growing as primary or secondary formations, on podzols and regosols mainly dystric or haplic with histic, ombric, gelic o stagnic character.

Bioclimatic diagnosis: Ic 23-46, Io 2.4-12.0, Tp 500-1000, Tps 400-550, Tmin < -4, m < -10. [Boco, Bosc, Teco-Tbo, Mbo, Sbo, Hbo, Ste, Ote-Dry, Shu, Hum].

We distinguish four associations with different ecological features and biogeographic distribution (figure 13): 1a. Pinetum banksianae (Continental Boreal -eucontinental-, humid and hyperhumid, mainly pyrogenic secondary forests of the Eastern Canadian Boreal Sector territories), 1b. Carici pensylvanicae-Pinetum banksianae (Continental Temperate, hemiboreal, humid and hyperhumid, mainly pyrogenic secondary macroforests, of the western district of Temperate Laurentian Subsector), 1c. Oryzopsio asperifoliae-Pinetum banksianae (Continental Boreal, occasionally Temperate hemiboreal, dry and subhumid, primary or pyrogenic secondary forests of the Western Canadian Boreal Sector and uncommon throughout the Canadian Prairies Sector areas), 1d. Pinetum banksiano-resinosae (Continental Temperate -eucontinental-, subhumid-humid, pyrogenic secondary macroforests of the Great Lakes Sector).

Characteristic species: Anemone quinquefolia var. bifolia, Comptonia peregrina, Geranium bicknellii, Leymus innovatus, Lilium philadelphicum var. andinum, Melampyrum lineare var. latifolium, Oryzopsis asperifolia, Pinus banksiana (Pinus divaricata), Pinus banksiana x P. contorta var. latifolia, Pinus resinosa, Prunus pensylvanica, Solidago nemoralis var. longipetiolata, Solidago nemoralis var. nemoralis, Solidago puberula var. puberula, Spiraea alba var. alba, Spiraea alba var. latifolia.

Remarks: Peinado, Aguirre & Cruz 1998 [in Plant Ecology 137: 190] published as lectotype of the alliance Pinion divaricatae Lavoie 1968 the association Comptonio (peregrinae)-Pinetum (divaricatae) Lavoie in Nat. Can. 95: 402. 1968. It was a wrong decission because the obliged lectotype following the current CPN (art. 20) is Pinetum divaricatae Lavoie in Nat. Can. 95: 403. 1968 [Pinetum banksianae nom. mut. propos. (art. 45)].

1a. Pinetum banksianae Lavoie 1968 nom. mut. et em. hoc loco

[Pinetum divaricatae Lavoie in Nat. Can. 95: 403, tb. I. 1968, Comptonio peregrinae-Pinetum divaricatae Lavoie in Nat. Can. 95: 402, tb. I. 1968 (syntax. syn.), Ledo-Pinetum divaricatae Lavoie in Nat. Can. 95: 407. 1968 (syntax. syn.), Gaylussacio-Pinetum divaricatae Lavoie in Nat. Can. 95: 407. 1968 (syntax. syn.), Kalmio angustifoliae-Pinetum banksianae comptonietosum peregrinae Grandtner, Camiré, Ansseau, Lamontagne & Villeneuve, Guide de l"excursion en Écologie Forestière: 118. 1991 (art. 1, 25), Pinetum banksianae Lavoie 1968 nom. mut. propos. (art. 45)]

Continental Boreal (eucontintental); thermo-, meso- and supraboreal; humid-hyperhumid; oligotrophic, primary and secondary pyrogenic Pinus banksiana meso-macroforests growing on sandy, iron or gley podzols in the Eastern Canadian Boreal Sector.

Bioclimatic diagnosis: Ic 28-38, Io 6.0-12.0, Tp 500-800, Tps 400-500, Tmin < -14º, m < -20º. [Boco-Tbo, Mbo-Hum].

North American associations of Gaultherio-Piceetalia North American biogeographic sectors
2.1a 2.1b 2.1c 3.1a 3.2a 3.3c 2.2a 2.2b
1. Pinion banksianae . + + + + + . .
1a. Pinetum banksianae . + . . . . . .
1c. Oryzopsio-Pinetum banksianae . . + . . + . .
1b. Carici-Pinetum banksianae . . . + . . . .
1d. Pinetum banksiano-resinosae . . . . + . . .
2. Gaultherio-Piceion glaucae + + + + + + . .
2a. Abietetum balsameae + + . . . . . .
2b. Kalmio-Piceetum marianae + + . . . . . .
2d. Mertensio-Abietetum balsameae . . + . . + . .
2c. Sphagno-Piceetum marianae + + + + + + . .
3. Piceion rubentis . . . + + . . .
3a. Abieti-Piceetum rubentis . . . + + . . .
4. Ledo-Piceion marianae . . . . . . + +
4a. Rubo-Piceetum marianae . . . . . . + +
4b. Piceetum glauco-marianae . . . . . . + +
4c. Hypno-Piceetum glaucae . . . . . . + +
4d. Pulsatillo-Pinetum latifoliae . . . . . . + .

Figure 12. Relationships between boreal and hemiboreal North American biogeographic sectors of North American Boreal Region and adjacent biogeographic territories and forest associations of Gaultherio-Piceetalia (1. Pinion banksianae, 2. Gaultherio-Piceion glaucae, 3. Piceion rubentis and 4. Ledo-Piceion marianae): 2.1. Canadian Boreal Province: 2.1a. Newfoundlandian Sector, 2.1b. Eastern Canadian Boreal Sector, 2.1c. Western Canadian Boreal Sector, 3.1. Appalachian Province: 3.1a. Appalachian Sector. 3.2. Great Lakes and Central Lowlands Province: 3.2a. Great Lakes Sector. 3.3. Prairies Province. 3.3a. Canadian Prairies Sector. 2.2. Yukonian-Alaskan Province: 2.2a. Cassiar and Mackenzie Mountains Sector, 2.2b. Continental Alaskan Sector.

In table 3 we assemble three relevés from some thermo- and mesoboreal localities of Québec official province, from Senneterre to Lac Aigremont in the Reserve Faunique de Ashwapmushuan, belonging to evolved phases with a pyrogenic origin from Abietetum balsameae and Kalmio-Piceetum marianae mature forests. The ombrophilous and acidophilous plants such as Vaccinium angustifolium and Kalmia angustifolia are frequent within Pinetum banksianae forests and could be used as differential species in front to their western subhumid geovicarious association Oryzopsio-Pinetum banksianae, common in the Western Canadian Boreal Sector but rare in the Canadian Prairies Sector territories.

Type relevé (lectotypus hoc loco): Lavoie in Nat. Can. 95: 403, tb. I, n. 11. 1968 sub Pinetum divaricatae pinetosum. Site: Ontario. Lac Abitibi (Ouest), 48°45'N-80°30'W. 200 m2. Estimated bioclimatic factors: Continental Boreal, eucontinental (Ic ~ 35.0), lower mesoboreal Table 3 (Tp ~ 670), lower humid (Io ~ 8.0), Positive Summer Temperature Tps ~ 480. Biogeographic location: North American Boreal Region, Canadian Boreal Province, Eastern Canadian Boreal Sector, Hudson Bay Basin Subsector. Floristic combination: Linnaeo-Piceetea characteristic species, trees: 4 Pinus divaricata [P. banksiana]. Others: + Alnus crispa [A. viridis subsp. crispa], + Gaultheria procumbens, 2 Kalmia angustifolia, + Melampyrum lineare [M. lineare var. lineare], 3 Vaccinium angustifolium, Vaccinium myrtilloides. Companion species: 3 Epigaea repens, + Epilobium angustifolium, + Salix discolor, 4 Cladonia sp. pl., 1 Bryophyta divers.

Table 3
1a. Pinetum banksianae

(Pinion banksianae, Gaultherio-Piceetalia)

Altitude (1=10m) 33 37 35 35
Number of species 17 17 17 17
Ordinal number 1 2 3 4
Trees:
Pinus banksiana 5 4 4 3
Picea mariana (S2) 2 3 2 3
Populus tremuloides 1 2 2 3
Betula papyrifera 1 . 1 2
Picea glauca (S2) . + 1 2
Characteristic species:
Kalmia angustifolia 3 3 4 3
Vaccinium angustifolium 2 3 4 3
Linnaea americana + 2 1 3
Vaccinium minus 1 + . 2
Gaultheria hispidula 4 . 3 2
Epigaea repens 1 . + 2
Coptis groenlandica + . 1 2
Maianthemum canadense . 1 + 2
Ledum groenlandicum 1 . . 1
Cornus canadensis . 1 . 1
Ribes glandulosum . 1 . 1
Companion species:
Pleurozium schreberi 5 5 5 3
Cladina sp. pl. 2 3 2 3
Ptilium crista-castrensis 3 . 2 2

Other species. Trees: Larix laricina (S2) 1 in 1. Abies balsamea (S2) + in 3. Characteristic species: Cypripedium pubescens 1 in 1. Alnus rugosa 1, Prunus pensylvanica 1, Amelanchier sanguinea + in 2. Monotropa hypopithys + in 3. Companion species: Epilobium angustifolium 1 in 2.

Localities: 1. Québec: 10 Km N from Senneterre. 48º15'N-77º30'W. N, 5%, 20 m, 30 cm, 500 m2, gleyic sand podzol (reg. 1995b: 27). 2. Québec: Reserve faunique Ashwapmushuan, Lac Aigremont. 48º40'N-72º43'W. S, 10%, 15 m, 20 cm, 400 m2, iron podzol (reg. 1995b: 46). 3. Québec: 15 Km N from Senneterre. 48º23'N-77º20'W. W, 10%, 18 m, 30 cm, 400 m2, sand podzol (reg. 1995b: 29). 4. Synthesized table.

1b. Carici pensylvanicae-Pinetum banksianae Lavoie 1968 nom. mut.

[Carici-Pinetum (divaricatae) Lavoie in Nat. Can. 95: 406. 1968, Carici pensylvanicae-Pinetum banksianae Lavoie 1968 nom. mut. propos. (art. 45)]

Continental Temperate (eucontinental and semicontinental); hemiboreal and upper supratemperate; humid-hyperhumid; oligotrophic secondary pyrogenic Jack pine meso- or macroforests (Pinus banksiana) with Populus grandidentata and Populus tremuloides in the undercanopy, growing on sandy podzols with the water table usually at less 1.5 m, in the Appalachian and Temperate Laurentian Subsectors.

Bioclimatic diagnosis: Ic 23-32, Io 6.0-10.0, Tp 800-1000, Tps 450-550, Tmin < -4, m < -10. [Teco-Hbo, Ste-Hum]

Type relevé (lectotypus hoc loco): Lavoie in Nat. Can. 95: 406, tb. I, n. 24. 1968 sub Carici-Pinetum betuletosum. Site: Québec. Cap-de-la-Madeleine, 46°22N-72°32W. 120 m2. Estimated bioclimatic factors: Continental Boreal, eucontinental (Ic ~ 32.0), hemiboreal (Tp ~ 870), lower humid (Io ~ 7.0), Positive Summer Temperature Ts ~ 540. Biogeographic location: North American Atlantic Region, Appalachian Province, Appalachian Sector, Temperate Laurentian Subsector. Floristic combination: Linnaeo-Piceetea characteristic species, trees: 3 Pinus divaricata [P. banksiana], 1 Populus grandidentata, + Populus tremuloides. Others: + Apocynum androsaemifolium [A. androsaemifolium var. androsaemifolium], 2 Aralia nudicaulis, + Cypripedium acaule, + Fragaria virginiana [F. virginiana subsp. virginiana], + Kalmia angustifolia, + Melampyrum lineare [M. lineare var. lineare], + Oryzopsis asperifolia, + Solidago nemoralis var. nemoralis, 2 Vaccinium angustifolium. Companion species: + Aster lineariifolius, + "Botrychium multiflorum", + Cladonia rangiferina [Cladina rangiferina], + Cladonia sp. pl., 1 Danthonia spicata, + lichens divers, + Lycopodium trystachyum [Diphasiastrum tristachyum], + Lycopodium flabelliforme [Diphasiastrum digitatum], 1 Polytrichum commune, + Solidago juncea.

Map Figure 13. Distribution of the forest plant associations belonging to alliance 1. Pinion banksianae: 1a. Pinetum banksianae (Eastern Canadian Boreal Sector), 1b. Carici pensylvanicae-Pinetum banksianae (Temperate Laurentian Subsector), 1c. Oryzopsio asperifoliae-Pinetum banksianae (Western Canadian Boreal Sector), 1d. Pinetum banksiano-resinosae (Great Lakes Sector: Western Hemiboreal and Eastern Great Lakes Subsectors).

1c. Oryzopsio asperifoliae-Pinetum banksianae Looman associatio nova hoc loco

[Oryzopsio-Pinetum banksianae Looman in Phytocoenologia 15(3) 300, tb. 1 (synthesized table) 1987 (art. 5)]

Continental Boreal (eucontinental), thermo- and mesoboreal, and Temperate hemiboreal; upper dry and subhumid, oligotrophic permanent and secondary pyrogenic Jack pine meso- and macroforests (Pinus banksiana) with Populus tremuloides in the undercanopy, growing in podzols throughout the Western Canadian Boreal and Canadian Prairies Sectors.

Bioclimatic diagnosis: Ic 28-46, Io 2.4-5.8, Tp 500-850, Tps 400-500, Tmin < -13º, m < -20º. [Boco, Teco-Tbo, Mbo, Hbo-Dry, Shu].

Table 4 gives two relevés from Alberta (Lesser Slave Lake) and Saskatchewan (Prince Albert) belonging to thermoboreal territories of the Western Canadian Subhumid Subsector in which are absent the most western and southern species such as Vaccinium angustifolium, Kalmia angustifolia and Pinus resinosa, which are frequent in other associations of the alliance.

Table 4
1c. Oryzopsio asperifoliae-Pinetum banksianae

(Pinion banksianae, Gaultherio-Piceetalia)

Altitude (1=10m) 51 40 45
Number of species 20 22 21
Ordinal number 1* 2 3
Trees:
Pinus banksiana 5 5 2
Populus tremuloides 2 1 2
Characteristic species:
Arctostaphylos adenotricha 5 3 2
Calamagrostis purpurascens 2 1 2
Galium boreale 1 3 2
Prunus pensylvanica 1 2 2
Linnaea americana 1 1 2
Cornus canadensis 1 1 2
Oryzopsis asperifolia 1 1 2
Solidago longipetiolata 1 + 2
Rosa sayi + 3 2
Geranium bicknellii + 1 2
Pyrola asarifolia + + 2
Melampyrum latifolium 2 . 1
Aster ciliolatus . 2 1
Companion species:
Campanula rotundifolia 2 1 2
Cladina sp. 2 1 2

Other species. Trees: Picea glauca (S2) 1 in 1. Characteristic species: Hedysarum americanum 1, Mertensia paniculata 1, Aralia nudicaulis + in 1. Prunus virginiana 2, Pulsatilla multifida 2, Maianthemum canadense 1, Alnus crispa +, Lilium andinum + in 2. Companion species: Pleurozium schreberi 2 in 2.

Localities: 1. Alberta: Slave Lake. Lesser Slave Lake Provincial Park. 55º28'N-114º51'W. W, 10%, 18 m, 40 cm, 200 m2, holotypus ass. (reg. 1995d: 156). 2. Saskatchewan: Between Prince Albert and Henribourg near North Saskatchewan River. 53º13'N-105º46'W. SW, 15%, 20 m, 40 cm, 400 m2 (reg. 1995d: 163). 3. Synthesized table.

Type relevé (holotypus): Table 4, relevé n. 1. Rivas-Martínez reg. 1995d: 156. 07.09.1995, made with Dr. M. Costa. Site: Alberta. Slave Lake. Lesser Slave Provincial Park. 55º28'N-114º41'W. 510 m, W, 10%, 18 m, 40 cm, 200 m2. Physiognomy and habitat: Jack pine (Pinus banksiana) mesoforest with scattered aspen trees (Populus tremuloides) in the understory and a dense herbaceous cover growing on sandy soils. Estimated bioclimatic factors: Continental Boreal, eucontinental (Ic ~ 38), upper thermoboreal (Tp ~ 740) lower subhumid (Io ~ 4.0), Positive Summer Temperature Ts ~ 480. Biogeographic location: North American Boreal Region, Canadian Boreal Province, Western Canadian Boreal Sector, Western Canadian Boreal Subhumid Subsector. Floristic combination: Linnaeo-Piceetea characteristic species, trees: 5 Pinus banksiana, 2 Populus tremuloides. Others: + Aralia nudicaulis, 5 Arctostaphylos uva-ursi var. adenotricha, 2 Calamagrostis purpurascens, 1 Cornus canadensis, 1 Galium boreale, + Geranium bicknellii, 1 Linnaea americana, 2 Melampyrum latifolium, 1 Mertensia paniculata, 1 Oryzopsis asperifolia, 1 Picea glauca var. glauca (S2), 1 Prunus pensylvanica, + Pyrola asarifolia, + Rosa acicularis subsp. sayi, 1 Solidago nemoralis var. longipetiolata. Companion species: 2 Campanula rotundifolia, 2 Cladina sp., 1 Hedysarum alpinum var. americanum.

1d. Pinetum banksiano-resinosae associatio prov.

Continental Temperate (eucontinental); hemiboreal and upper supratemperate; upper subhumid and lower humid; primary and secondary pyrogenic red pine (Pinus resinosa) and Jack pine (Pinus banksiana) macroforests, growing on iron or gleyic podzols throughout the Great Lakes Sector territories (Western Hemiboreal and Eastern Great Lakes Subsector). The optimum of this provisional forest association organized by many temperate and boreotemperate trees (Pinus banksiana, Pinus resinosa, Pinus strobus, Abies balsamea, Larix laricina, Picea mariana, Populus balsamifera subsp. balsamifera, Populus tremuloides, etc.) seems to be in the eucontinental humid hemiboreal plains of the Great Lakes areas western of Carici-Pinetum banksianae association territories.

Bioclimatic diagnosis: Ic 25-35, Io 5.0-10.0, Tp 800-1000, Tps 450-550, Tmin < -4, m < -10. [Teco-Hbo, Ste, Ushu, Lhum].

Revealing relevé: Rivas-Martínez reg. 1992d: 20. 28.05.92, made with Drs. A. Asensi, J. Molero & F. Valle. Site: Minnesota: St. Louis Co. Superior National Forest. Mesabi Iron Range, between Mountain Iron and Virginia. 47º31'N-92º24'W. 390 m, 25 m, 40 cm, 400 m2. Physiognomy and habitat: Red pine (Pinus resinosa) and Jack pine (Pinus banksiana) mesoforest growing in sandy soils. Estimated bioclimatic factors: Continental Temperate, eucontinental (Ic ~ 33.0), hemiboreal (Tp ~ 890), lower humid (Io ~ 6.5), Positive Summer Temperature Tps ~ 430. Biogeographic location: North American Atlantic Region, Great Lakes and Central Lowlands Province, Great Lakes Sector, Western Hemiboreal Great Lakes Subsector. Floristic combination: Linnaeo-Piceetea characteristic species, trees: 2 Abies balsamea, 2 Acer spicatum, 2 Betula papyrifera var. papyrifera, 1 Larix laricina, 2 Pinus banksiana, 4 Pinus resinosa, 1 Pinus strobus, 1 Populus balsamifera. Others: 2 Anemone quinquefolia var. bifolia, 2 Aralia nudicaulis, 3 Clintonia borealis, 2 Gaultheria procumbens. Companion species: 2 Luzula acuminata var. carolinae, 1 Osmunda claytoniana, 2 Pteridium aquilinum var. latiusculum, + Salix pyrifolia.