Ecology and Management of Larix : A Look Ahead Proceedings of an International Symposium

Whitefish, Montana, U.S.A. October 5-9, 1992

Compilers:

Wyman C. Schmidt Kathy J. McDonald Around the World with Larix: an Introduction Wyman C. Schmidt

Circling the globe at 60° N latitude, one is seldom out of . Their light green hues in the spring and sum­ sight of Larix on the extended landmasses of Eurasia and mer, the gold in the fall, and the absence offoliage in the North America. forests essentially encircle the North­ winter are but a part ofthe charm that this unique genus ern Hemisphere, stretching from eastern westward adds to its environs. across Eurasia (but presently absent in Scandinavia), re­ The 10 most commonly recognized larch species and their suming in eastern North America and westward across the general distribution are listed in table 1. In addition to United States and Canada to Alaska, where except for the these 10 species there are a large number of subspecies and Bering Sea, they essentially reach our starting point back hybrids where natural ranges of species overlap. Larch tax­ in Siberia (fig. 1). But along that approximate 20,000-km onomy has had little attention internationally. The last real (12,000-mile) path, larch splits into 10 species and numerous defmitive examination was over 60 years ago by Ostenfeld varieties and hybrids. These 10 species occupy a wide vari­ and Larsen (1930). This is reflected in the lack of total ety of ecological conditions and zones ranging from lowland agreement in the international literature about what consti­ boreal to upper montane to upper subalpine conditions and tutes a Larix species or a subspecies. It is the age-old tax­ extend south to 25° latitude at high elevations and north to onomy discussion between the "splitters and the lumpers." 75° latitude in the boreal lowlands. Splitting the species into subspecies often makes biological have been in the same general area for a long sense at the local or regional level, but in the larger context time. Larch fossils recovered from sediments laid down in it makes generalizations difficult. For the purposes of this the Oligocene to the Holocene eras have been described in introduction, generalization to 10 species is in order, but for North America, Europe, and Asia. More species oflarch papers within this proceedings, breakdowns into subspecies have already gone extinct than the 10 presently surviving and hybrids are described that certainly prove helpful in species. Fossil larch have been found in northern Canada, relating to individual research activities around the world. Poland, , Japan, and Alaska, U.S.A. Lepage and To more readily illustrate the magnitude of and differences Basinger (1991) list many of the fossil species described in in Larix species distribution, I have divided the northern the world literature and describe in detail excellent fossil temperate zone into four geographic areas: North America, remains of Larix altoborealis found in the Canadian Arctic. Europe, Northern Asia, and Southern Asia (figs. 2, 3, 4, 5). AlllO larches are in the genus Larix, a needle­ Larix distributions shown here are only approximate and gymnosperm in the family . Although similar are based on adaptations made from maps and narratives in appearance, shade tolerance, and deciduous character, from several sources. Most ofthe descriptions in the litera­ larch species do differ substantially in growth, ability to ture are in at least partial, but usually not total, agreement. establish on different substrates, and ability to compete Good defmitive information on exact ranges of the species successfully with associated species. Larch's deciduous is just not available in some cases. characteristic clearly distinguishes the genus from ever­ Species boundaries in North America are relatively well green conifers with which it is almost always associated. defmed with practically no overlap between L. laricina, L. Larches are the exception in the characteristically ever­ occidentalis, and L. lyallii (fig. 2). Although L. occidentalis green world of the northern boreal and mountain subalpine and L. lyallii occur in much the same geographic area, they forests of the northern hemisphere. They possess morpho­ are usually elevationally separated by 300 to 500 m. Excep­ logical and physiological characteristics that distinguish tions to this are noted in this proceedings. them from their evergreen or deciduous counterparts and Larix laricina forests are by far the most extensive of the likely provide them with unique establishment and survival three species in North America, stretching from the east advantages. But they do well in spite oftheir differences, to the west of Canada and into Alaska, U.S.A. Its largely especially in adding the diversity that is advantageous to boreal habitat contrasts sharply with that of the upper associated flora and fauna. Aesthetically, Larix species have montane/subalpine habitat of L. occidentalis. no real match in the evergreen world of temperate Only one species of Larix occurs naturally in Europe. most commonly occurs in the subalpine habi­ tat of the Alps, but it also occurs in other areas of central Europe (fig. 3). Different subspecies and varieties are com­ monly recognized there and are described in this proceedings. Larix russica (often referred to as ) and L. Paper presented at the Symposium on Ecology and Management of gmelinii dominate the Siberian forest landscape in North­ Larix Forests: A Look Ahead, Whitefish, MT, U.S.A., October 5-9, 1992. ern Asia, with L. russica to the west and L. gmelinii to the Wyman C. Schmidt, Forestry Sciences Laboratory, Intermountain Re­ east (fig. 4). Their major boundary is contiguous in a gen­ search Station, Forest Service, U.S. Department of Agriculture, located at Montana State University, Bozeman, MT 59717-0278, U.S.A. erally north-south direction for thousands of kilometers,

6 WORLD DISTRIBUTION OF LARIX

Figure 1-Natural range of the genus Larix throughout the World (adapted from Krlissman 1985).

Table 1-The 10 commonly recognized species of larch with their general location and ecological situation.

Latin name Common name General location Ecological zones Larix occidentalis Western larch Rocky and Cascade Mountains of U.S. and Canada Upper montane to lower subalpine Larix Iyallii Alpine larch Rocky and Cascade Mountains of U.S. and Canada Upper subalpine to timberline ecotone Larix laricina Tamarack Northeastern and Lake States and Alaska in U.S. Mainly boreal and a wide belt completely across Canada Larix russica Siberian larch A wide belt in northern Russia and in Boreal to northern timberline Asian larch Eurasia east of the Siberian larch range Subalpine to (includes L. dahurica, northern olgensis, cajanderi, timberline other subspecies) Larix mastersiana Masters larch Mountain areas in south Upper montane to lower subalpine Larix griffithiana Sikkim larch Himalayas in Nepal, Bhutan, Tibet and south China High subalpine Larix potaninii Chinese larch Western China High subalpine Larix leptolepis Japanese larch Honshu,Japan Subalpine Larix decidua European larch Alps area in France, Switzerland, Austria, Italy, Subalpine Yugoslavia, Germany with scattered areas in Romania, Czechoslovakia, and Poland

7 NORTH AMERICA

_ Larix Iyallli

iii Larix occldentalis 200 400 600 100 1000 kilo.".., I I I I ! 111111111111 Larix larlclna Jo In Jx,maloI

Figure 2-Natural range of Larix species in North America (adapted from Johnston 1990; Arno 1990; Schmidt and Shearer 1990).

and where their boundaries overlap L. x czekanowski (=L. For at least 200 years people have carried from one russica xL. gmelinii) is often recognized. Within the wide continent to the other in hopes of fmding the perfect Larix range of L. gmelinii there are numerous regionally recog­ species for their area. As a result, plantations of introduced nized subspecies or varieties such as L. cajanderi in north­ Larix can be observed at many locations in the world, par­ eastern Siberia, L. olgensis on the east coast of Russia and ticularly in Europe and eastern North America (Kriissman down into Korea, L. principis-rupprechtii in northeast 1985). Genetics research, particularly with hybridization China, and L. kurilensis and L. kamtschatica on objectives, has been extensive. Some hybrids exhibit supe­ Island and Kamchatka. These are described in other pa­ rior growth and survival characteristics, and some of that pers within the proceedings. information is presented in this proceedings. Southern Asia accounts for a wide variety of Larix species, The value of Larix forests for wood products, animal ranging from the montane conditions of L. mastersiana in habitats, water production, aesthetics, and other resources southwestern China to the high elevation forests of L. grit­ is impressive, but the values vary tremendously by species fithiana in Nepal, Bhutan, and Tibet, and L. potaninii in and ecological zones. These forests harbor a wide comple­ southwestern China to the island environment of L. lep­ ment of fauna ranging from the moose to the mouse, the tolepis on Honshu in Japan (fig. 5). Larix mastersiana bear to the shrew, and the eagle to the hummingbird, not and L. leptolepis are unique, along with L. lyallii in North to mention the vast array of micro flora and fauna, as yet America, in having limited, but important, ranges. only generally comprehended. Many of these values and To the casual observer most Larix species look essentially ecological principles of this truly international genus are the same, but their cone and needle characteristics and par­ described in this proceedings. ticularly their ecological niches separate them. Some of these characteristics are illustrated in figure 6.

8 EUROPE

40" 50"

o 200 400 600 IlOO 1000 kilo I I!' I i I 200 A()() (:J:XJmllu

Larix decidua .0·

"\,.,\ ...... ~ ,.~ .~) Belorussia -;:-: ": :i,\ ······· .... :··,···.. ·t. Ukraine 50·

Figure 3-Natural range of Larix species 4"" 40· in Europe (adapted from Ostenfeld 1930; Gower and Richards 1990; Holtmeier, this proceedings).

Northern Asia

o "

_ larix gmelinil larix russlca 200 400 600 800 un} kilomokn iii::: ililllill larix leptolepis I I I I I I I I I russlcalgmelinii China o ax> 400 600 mil ..

Figure 4-Natural range of Larix species in Northern Asia (adapted from Ostenfeld and Larsen 1930; Gower and Richards 1990; Milyutin and Vishnevetskaia, this proceedings). The area shown as russicalgmelinii is often referred to as L. x czekanowski, and the area in northeast China shown as L. gmelinii is often referred to as L. principis-rupprechtii.

9 SOUTHERN ASIA

...... _ Larix gmellnii Ililllilll Larix leptolepls Ili ~~lliliii Larix potaninil _ Larix mastersiana 11111111111Larix griffithiana

.). -.,.,

.l; • • '

o 200 400 600 800 1000 kilometer• . I iii i o 200 400 600 milea

Figure 5-Natural range of Larix species in Southern Asia (adapted from Ostenfeld and Larsen 1930; Gower and Richards 1990; Wang, this proceedings). The area shown as russicalgmelinii is often re­ ferred to as L x czekanowski, and the area in northeast China shown as L gmelinii is often referred to as L principis-rupprechtii.

REFERENCES Lepage, Ben A; Basinger, James F. 1991. A new species of Larix (Pinaceae) from the early tertiary of Axel Heiberg Arno, Stephen F. 1990. Larix lyallii. In: Agriculture Hand­ Island, Arctic Canada. Review of Palaeobotany and Pa­ book 654. Washington, DC: U.S. Department of Agricul­ lynology, Elsevier Science Publisher B.V., Amsterdam. ture, Forest Service: 152-159. 70: 89-111. Gower, Stith T.; Richards, James H. 1990. Larches: Decidu­ Ostenfeld, C.H.; Larsen, C.S. 1930. The species of the ge­ ous conifers in an evergreen world. Bioscience. 40(11): nus Larix and their geographic distribution. Kongelige 818-826. Danske Videnskabemes Selskab Biologiske Meddelelser. Johnston, William F. 1990. Larix laricina. In: Agriculture 9: 1-106. Handbook 654. Washington, DC: U.S. Department of Schmidt, Wyman C.; Shearer, Raymond C. 1990. Larix Agriculture, Forest Service: 141-151. occidentalis. In: Agriculture Handbook 654. Washington, Krtissman, G. 1985. Manual of cultivated conifers. Timber DC: U.S. Department of Agriculture, Forest Service: Press, Portland, OR. pp. 157-163. 160-172.

10 Figure 6-This series of photos depicts some of the characteristics of the world's Larix species.

Larix occidentalis

L. occidentalis in a fall landscape scene in west­ em Montana, U.S.A. A 15-year old L. occidentalis in the spacing study on Coram Experimental Forest, Montana, U.S.A.

A 300-year old stand of L. occidentalis on Coram Experimental Forest, Montana, U.S.A. Large are about 40 m tall and nearly 1 m An ovulate cone of L. occidentalis in the in- diameter. early spring. 11 Larix /yallii

Ovulate cones and emerging foliage of L. Iyallii A late September view of L. Iyallii, Carlton Ridge, in early spring. Bitterroot Mountains, Montana, U.S.A.

A robust stand of L. Iyallii, Carlton Ridge, Bitterroot Mountains, Montana, U.S.A.

A planted L. Iya/lii seedling.

12 -Larix laricina

A stand of L. laricina during the October needle fall period on a lowland in Alberta, Canada.

Larix laricina provenance trial in Alberta, Canada.

13 Larix russica

A mixed species forest of larch, birch, and pine in the fall season near Lake Baikal in Russia.

A mature forest in the steppe area of Mongolia.

14 Larix gmelinii -

A fall scene in an intermediate-age stand of the sub­ species L. cajanderi, Yakutia, Russia.

A vigorous young forest of L. gmelinii in Korea.

A landscape showing an extensive forest of L. gmelinii in Korea.

15 A young vigorous stand of L. gmelinii in northeast China.

A stand of L. gmelinii adjoining an agricultural area in northeast China.

Establishing weather instruments in a forest of the A plantation of L. gmelinii, subspecies o/gensis in subspecies L. o/gensis in northeast China. northeast China.

16 Larix mastersiana -

A branchlet and cone of L. mastersiana in south­ A I~c~-~ike crown showing the drooping charac­ west China. teristiC In a southwest China forest.

Larix potaninii

Mature cones of L. potaninii in southwest China.

17 Larix leptolepis

A vigorous intermediate age stand in Japan. A thinned intermediate age stand with a heavy understory in Japan.

Larix decidua

Intermediate age L. decidua in a mixed­ species stand of spruce and pine near st. Moritz. Switzerland.

18