TiliaceaeCLinden family

Tilia L.
linden or basswood

D. Bradley Rowe and Frank A. Blazich

Dr. Rowe is an assistant professor at Michigan State University=s

Department of Horticulture, East Lansing, Michigan; Dr. Blazich is professor of
plant propagation and tissue culture at North Carolina State University=s
Department of Horticultura; Science, Raleigh, North Carolina.

Growth habit, occurrence, and uses. The genus Tilia (L.)Clinden or

basswoodC
consists of about 40 species of large or medium-sized, deciduous trees that are
indigenous to the temperate Northern Hemisphere. Tilia is the only genus of its
family, the Tiliaceae. Species reach their maximum size in loamy, moist, fertile
soil, but they tolerate poor soils, pollution, windy conditions, and transplanting
and can be grown in full sun or partial shade (Dirr 1990; Haller 1995;
Kunneman and Albers 1991). Lindens possess a well-developed root system
and are long lived, with some species living between 500 to 1,000 years (Haller
1995; Kunneman and Albers 1991). Table 1 lists species native to North
America, as well as non-native species that are widely grown here.
Few shade trees vary so greatly in shape, leaf size, and growth rate as do
the lindens (Flemer 1980). They generally possess a uniform globular crown
and smooth, silver-gray bark that becomes fissured on old trees (table 2). The
winter form is striking, with stiff, erect branches growing upward at 30E angles
from a thick trunk (Burgess 1991). Considerable differences in growth habit
exist among cultivars of littleleaf linden, ranging from the very dense, formal
pyramidal habit of >Greenspire=, the dense upright oval shape of >Chancellor=, to
the more open, informal oval habit of >Fairview= (Pellett and others 1988).
There is much disagreement among taxonomists as to correct
identification of species and there are numerous names in the literature that
are no longer recognized by many botanists. For example, T. monticola Sarg.
and T. michauxii (Nutt.) Sarg. are sometimes seen in the literature or listed as
specimens in botanical gardens, but they are now considered to be varieties of
white basswoodCT. americana var. heterophylla (Venten.) Loud.Crecognized
previously as T. heterophylla Venten. (Ayers 1993; Rehder 1990).
Lindens are generally not suitable for lumber because the wood is soft
and rots easily. However, the soft, straight-grained and even-textured wood is
ideal for woodcarving and is utilized to make musical instruments, piano keys,
Venetian blinds, and veneer and can serve as a source of fiber (Haller 1995;
Kunneman and Albers 1991). The wood does not produce splinters, thus
making it ideal for tool handles. The inner bark (or Abast@) consists of long,
tough fibers that once were used in the production of cordage, mats, and
clothing. The common names for the speciesCbasswood, linden, and limeCare
derived from this characteristic: bast gives us the name bastwood or basswood;
linden and lime are thought to be derived from the Latin word for linen (Haller
1995). In addition, flowers of linden are quite fragrant and produce large
quantities of nectar that is very attractive to bees. The flowers of European,
bigleaf, and littleleaf lindens are brewed for tea (Bremness 1994). The nectar of
some species is so overpowering that bees can be found inebriated on the
ground beneath the tree (Haller 1995). The light-colored honey produced is
world famous.
Lindens are used primarily as ornamental shade and street trees (table
2), more so in Europe than in the United States. For example, Berlin=s most
famous boulevard is named AUnter den Linden@. They are well-adapted to a
broad range of soil and climatic conditions and are relatively free of major
disease problems that may threaten the survival or landscape value of
established trees (Pellett and others 1988). The European lindensClittleleaf,
European, bigleaf, and silver lindensChave greater importance in landscape
plantings in the United States because they are more tolerant and ornamental
than American species such as American linden (Dirr 1990; Heit 1977). In
addition, American linden becomes too large for the average home property and
is better left in the forest (Dirr 1990). However, silver linden possesses a
shallow root system and its canopy casts dense shade, making it unsuitable for
underplanting (Burgess 1991).
Geographic races and hybrids. As mentioned previously, there is
much disagreement among taxonomists as to correct identification of species.
For example, there is debate whether white basswood is a southern race of
American linden or a separate species. Also, hybridization between species
occurs naturally and has given rise to variability among seedlings (Kunneman
and Albers 1991). Of the more common hybrids, Crimean and European
lindenw are not considered superior landscape trees relative to littleleaf linden
(Dirr 1990).
Flowering and fruiting. Perfect, fragrant, yellowish or whitish flowers
that bloom in June or July are borne in short, pendulous cymes with stalks
attached to a large thin-textured oblong bract. Trees and clonal groups of trees
flower almost simultaneously over the exposed parts of their crowns. In each
inflorescence, the terminal flower of the dichasium opens first and in warm
weather is followed at intervals of a day by flowers on the branches of
successive orders (Pigott and Huntley 1981). Trees usually flower within 5 to
15 years when grown from seed. Shortness of blooming period (several days to
2 weeks, depending on weather conditions) and lack of consistent flowering
from year to year are problems for beekeepers harvesting honey (Ayers 1993).
In particular, the American lindens have a reputation for not flowering every
year. Some of the introduced species are more consistent (Ayers 1993).
Following pollination, temperatures must be >15 EC for growth of the
pollen tube and for fertilization to occur so that fruits will be produced (Pigott
and Huntley 1981). Fruits are grayish, nut-like, round to egg-shaped capsules
that mature in autumn but may persist on the tree into the winter. Each
consists of a woody pericarp enclosing a single seed (but sometimes 2 to 4
seeds) (figures 1 and 2) (Brinkman 1974; Pigott and Huntley 1981). The
pericarp consists of an outer layer of loose fibers forming a mat (or tomentum)
and a broad region of thick-walled lignified fibers that are responsible for its
hard, tough, woody character (Spaeth 1934). Fruits of American linden are
tough and leathery, whereas those of littleleaf linden tend to be thinner and
rather brittle (Heit 1967). Seeds possess a crustaceous seedcoat; a fleshy,
yellowish endosperm; and a well-developed embryo (figures 2 and 3). Natural
dispersion is primarily by wind and animals (Brinkman 1974).
Collection of fruits; seed extraction and cleaning. The ideal time to
harvest fruits is early fall, when seed moisture content is approximately 16%
(Vanstone 1982). During fruit ripening, moisture is lost from the seeds at a rate
of 1 to 2% per day, so that seeds must be monitored closely. Pericarp color is a
reliable indicator of moisture content in relation to germination. Fruits should
be picked when the pericarp is turning from green to grayish-brown and before
the pericarp becomes tough and leathery. Otherwise, seeds will require greater
efforts during extraction and scarification. There is generally uniform ripening
on any individual tree, but the exact date of ripening may vary by several
weeks among trees (Vanstone 1978). Because fruits of linden persist on the
tree, fruit collection is often postponed until after maturity. After a heavy frost,
fruits can be shaken from branches onto a canvas tarp and spread out to dry
(Brinkman 1974).
Once fruits have been collected, bracts can be removed by flailing or
passing the fruits through a de-winging machine. The hard pericarp must then
be removed. Fruits of littleleaf and bigleaf lindens have prominent sutures that
are helpful in the extraction process by serving as a breaking point for both
mechanical and rubbing techniques (Heit 1977). Fruits of European and silver
lindens can be handled similarly, and a combination of sieving, screening, and
blowing can readily remove debris (Heit 1977). However, fruits of American
linden have a hard, tough, leathery pericarp and must be run through a coffee
grinder or a similar device or treated with acid to accomplish this removal (Heit
1977). Mechanical extraction of seeds is often difficult. Any crushing force
sufficient to fracture the tough pericarp is likely to exert a shattering pressure
on the brittle seedcoat (Spaeth 1934). Seed yields and size vary by species
(table 3)
Storage. Seeds of the lindens are orthodox in storage behavior and
should be stored in sealed containers at a moisture content of 8 to 12%. Seeds
of American linden have retained their viability for 2 years when stored under
dry conditions at room temperature and for 5 to 6 years when stored at 1 to 4
EC (Heit 1977).
Pregermination treatments. In addition to their tough pericarps,
seeds of linden exhibit double dormancy and thus require both scarification
and stratification (moist-prechilling) because of their impermeable seedcoat
and dormant embryo, respectively. For seeds of littleleaf linden, Heit (1977)
recommended a sulfuric acid treatment of 10 to 50 minutes at a temperature
ranging from 23 to 27 EC. Colder temperatures required a longer duration of
acid treatment. Because all species of linden and individual seedlots within the
same species are variable in their percentage and degree of hardseededness, it
is advisable to soak some seeds in water for 1 or 2 days to determine the degree
of hardseededness before treating with acid. Ten to 20 minutes of acid
treatment may be ideal for some seedlots, but 20 to 50 minutes would produce
the best results for others. The degree of hardseededness depends on many
factorsCas seed source, time of collection, and storage conditions, including
temperature and relative humidity (Heit 1967, 1977). Other scarification
treatments include mechanical scarification and hot water treatments, but
neither are as good as acid scarification (Heit 1977). Freezing to !80 and !185
EC had little effect on the permeability of the seedcoat (Spaeth 1934). Surface
sterilization with sodium hypochlorite (NaOCl) and ethanol proved to control
seed pathogens but lowered germination percentages of littleleaf, bigleaf, and
silver lindens (Magherini and Nin 1993).
In addition to scarification, stratification is essential for maximum
germination and seedling production. Following scarification, seeds must be
either fall-sown immediately or stratified at 1 to 3 EC) for about 3 months
before spring-sowing. Vanstone (1978) recommends stratification in a 1:1
mixture (by volume) of peat and sand containing 30% moisture by weight.
Enzyme activity and levels of soluble proteins and amino acids in the seeds
increase gradually during stratification at 4 EC (Pitel and others 1989).
Nontreated seeds have been known to lie in the ground for over 5 years without
germinating while still maintaining viability (Heit 1967). Bigleaf linden requires
3 to 5 months of warm stratification followed by 3 months of cold, and even
this treatment does not guarantee high germination (Dirr and Heuser 1987).
Flemer (1980) recommends burying seeds in a wooden box filled with damp
sand and leaving the box outdoors during the winter. Boxes are then dug up
the following fall and the seeds are sown. Seed treatments that consistently
result in good germination for all species and seedlots have not been developed.
Much variability exists among species and seed lots in regards to permeability
of the pericarp and seedcoat, as well as stratification requirements.
In Europe, dormancy in littleleaf linden is overcome by the use of warm
incubation or acid scarification, followed by stratification (Suszka and others
1996). Fully imbibed seeds are first stored for 4 months at 20 to 25 EC (or
scarified with concentrated sulfuric acid for 12 minutes), then stratified at 3 EC
for 14 to 18 weeks. Stratification should be stopped when the first seeds start
to germinate.This complete process may take 8 or 9 months.
Germination tests. Germination is epigeal (figure 4). Optimum
germination occurs at temperatures above 20 EC (68 EF), but seeds will
germinate at temperatures as low as 2 EC once stratification requirements have
been satisfied (Spaeth 1934). Thus, seeds should be checked periodically for
radicle emergence during stratification. Light is not required for germination
(Heit 1967). The use of any stratification procedure requires far too much time
to be used in routine germination testing, however, so rapid estimates of
viability are recommended for this purpose. This can be done with tetrazolium
staining, indigo-carmine staining, or excised embryo tests (ISTA 1996; Suszka
and others 1996). However, these tests require removal of the pericarp and the
seedcoat without damaging the embryo. Tetrazolium staining is the most
common test. It requires soaking seeds in water for 18 to 24 hours, removing
all or a large part of the seedcoat, and soaking the seeds in a 1% tetrazolium
solution for 24 to 48 hours at 30 EC.
Pitel and Wang (1988) found that both the rate and percentage of
germination of seeds of American linden were increased by treating scarified
seeds with a solution of kinetin (1 mg/l) and gibberellic acid (GA3, 500 mg/l.
Over 90% germination was obtained after 60 to 80 days at 4 EC. However, GA
did not improve germination percentage of lots of littleleaf, bigleaf, and silver
linden seeds (Magherini and Nin 1993). The conflicting results are likely due to
the level of gibberellin present. Natural levels of GA exist in dormant,
nonstratified seeds and a sudden increase in the quantity of gibberellin is
observed from the sixth week of stratification (Nagy 1980). It is likely that a
specific quantity, rather than just the presence of free gibberellins, is required
to break dormancy and stimulate germination.
Traditionally, for an accurate germination test, the outer pericarp must
be removed and the hard seeds must undergo scarification and stratification.
However, excised embryos of American linden that were separated from the
seedcoat and endosperm were able to germinate and grow when placed on an
agar medium without any pretreatment (Vanstone 1982). However, if any of the
endosperm was retained around the embryo, no growth took place. This
indicates an apparent lack of embryo dormancy, for the naked embryo will
grow when it is separated from other parts of the seed. Some factor that
restricts germination seems to be present in the endosperm and must be
overcome before an intact seed can germinate. That factor would normally be
overcome by stratification. The same result was obtained with bigleaf linden,
for germination was induced by removing the endosperm tissue around the
radicle (Nagy and Keri 1984).
Nursery practice and seedling care. Most trees in culture are of
seedling origin (Kunneman and Albers 1991). However, some are propagated by
grafting, chip budding, layering, rooting winter hardwood or leafy softwood
cuttings, or tissue culture (Flemer 1980; Howard 1995; Kunneman and Albers
1991). For grafting, seedling rootstocks are used, preferably of the same
species as the scion, as incompatibility is a common phenomenon (Kunneman
and Albers 1991). Named cultivars are grafted commonly in spring or budded
in summer (LHBH 1976). Plants of littleleaf linden have been propagated by
somatic embryogenesis initiated from immature zygotic embryos and then
established successfully in soil (Chalupa 1990). Except for hybrids such as
Crimean and European linden, all can be seed-propagated (Dirr and Heuser
1987).
Production by seed at a specified time is often relatively difficult (Dirr and
Heuser 1987; Heit 1967). As described previously, seeds show delayed
germination because of a tough pericarp, an impermeable seed coat, and a
dormant embryo. Seeds may remain in the ground for several years and never
produce a good stand of seedlings. The degree of seedcoat hardness and
embryo dormancy varies within and among seed lots for most species
(Hartmann and others 1997). Also, germination is irregular, and unknown seed
sources and hybridization between species have given rise to variability among
seedlings (Kunneman and Albers 1991). In addition, Heit (1977) found that
several lots of seeds of bigleaf and silver lindens from Europe contained high
percentages of empty seeds, from 20 to 72% (Heit 1977). This condition should
always be checked before sowing or treating seeds.
Mature fall-collected seeds may be sown in spring following scarification
and stratification (see Pregermination treatments). An alternate method is to
collect seeds early, before the pericarp turns brown and sow in the fall. Early
seed collections may result in seeds that have soft seedcoats that do not
require scarification (Heit 1977). However, some propagators have harvested
early and obtained inconsistent results, with the seeds sometimes decaying.
Late-harvested seeds may also be germinated the first season but require more
treatment than seeds harvested at the ideal stage of maturity (Vanstone 1978).
Seeds are sown in shallow rowsC6 to 13 mm (3 to 2 in)Cin beds and
covered with sand to aid in seedling emergence. The emerging seedlings are
very delicate and subject to sun scald, so lathe screens or shade netting over
the seedbeds greatly improves seedling stands (Flemer 1980). Fall-sown
seedbeds should be mulched, protected from rodent damage, and kept moist
until germination begins in the spring (Vanstone 1978). Good stands of
littleleaf, bigleaf, l and silver lindens are normal, but seeds of American linden
exhibit great variation in germination from year to year (Flemer 1980). When
poor stands result, seedlings should be removed carefully so as not to disturb
the bed, for additional germination often occurs the second year after planting.
Seedlings are usually outplanted as 1+0 or 2+0 stock.

References

Ayers GS. 1993. Reconsidering the basswoods: 11. The native American
basswoods. American Bee Journal 133(5): 337B340.
Bremness L. 1994. Herbs. London: Dorling Kindersley: 88.
Brinkman KA. 1974. Tilia, basswood, linden. In: Schopmeyer CS, tech. coord.
Seeds of woody plants in the United States. Agric. Handbk 450.
Washington, DC: USDA Forest Service: 810B812.
Burgess KS. 1991. Tilia tomentosa. Public garden: The Journal of the
American Association of Botanical Gardens and Arboreta 6(1): 39.
Chalupa V. 1990. Plant regeneration by somatic embryogenesis from
cultured immature embryos of oak (Quercus robur L.) and linden (Tilia
 cordata Mill.). Plant Cell Reports 9(7): 398B401.
Dirr MA. 1990. Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation and uses. 4th ed.
Champaign, IL: Stipes Publishing Co. 1007 p.
Dirr MA, Heuser CW Jr. 1987. The reference manual of woody plant
propagation: from seed to tissue culture. Athens, GA: Varsity Press. 239
p.
Flemer W III. 1980. Linden propagation: a review. Combined Proceedings of
the International Plant Propagators= Society 30: 333B336.
Haller JM. 1995. Tilia americana, linden: a neglected jewel. Arbor Age 15(7):
32B33.
Hartman RT, Kester DE, Davies FT Jr, Geneve RL. 2002. Hartman and
Kester=s plant propagation: principles and practices. 7th ed. Upper
Saddle River, NJ: Prentice-Hall. 880 p.
Heit CE. 1967. Propagation from seed: 7. Successful propagation of six
hardseeded group species. American Nurseryman 125(12): 10B12, 37B41,
44B45.
Heit CE. 1977. Propagation from seed: 27. Collecting, testing and growing
tilia linden species. American Nurseryman 146(7): 10B11, 100B110.
Howard BH. 1995. Opportunities for developing clonal rootstocks from
natural seedlings of Tilia spp. Journal of Horticultural Science 70(5):
775B786.
ISTA [International Seed Testing Association]. 1996. International rules for
seed testing. Seed Science and Technology 24(suppl.): 335.
Kunneman BPAM, Albers MRJ. 1991. Linden trees (Tilia spp.). In: Bajaj YPS,
ed. Biotechnology in agriculture and forestry. Trees 3(16):152B163.
LHBH [Liberty Hyde Bailey Hortorium]. 1976. Hortus third: a concise
dictionary of plants cultivated in the United States and Canada. 3rd ed.
New York: Macmillan. 1290 p.
Magherini R., Nin S. 1993. Experiments on seed germination of some Tilia
spp. Acta Horticulturae 331: 251B258.
Nagy M. 1980. Dormancy in fruits of Tilia platyphyllos Scop.: 4. Changes in
the endogenous gibberellin content during stratification. Acta
Agronomica 29: 1B11.
Nagy M. Keri A. 1984. Role of the embryo in the cytolysis of the endosperm
cells during the germination of the seeds of Tilia platyphyllos Scop.
Biochemie und Physiologie der Pflanzen 179: 145B148.
Pellett H, Vogel K, McNamara S., Zuzek K. 1988. Relative growth rate and
plant habit of linden taxa. Journal of Environmental Horticulture 6(2):
48B52.
Pigott CD, Huntley JP. 1981. Factors controlling the distribution of Tilia
cordara at the northern limits of its geographical range: 3. Nature and
causes of seed sterility. New Phytologist 87(4): 817B839.
Pitel JA, Cheliak WM, Wang BSP. 1989. Some biochemical changes
associated with stratification and germination of basswood seeds. Seed
 Science and Technology 17(1): 57B71.
Pitel JA, Wang BSP. 1988. Improving germination of basswood (Tilia
americana L.) seeds with gibberellic acid. Seed Science and Technology
16(1): 273B280.
Plotnik A. 2000. The urban tree book: an uncommon field guide for city and
town. New York: Three Rivers Press 211B215.
Rehder A. 1990. Manual of cultivated trees and shrubs hardy in North
America. 2nd ed. Portland, OR: Dioscorides Press. 996 p.
RHS [Royal Horticultural Society]. 1994. The new Royal Horticultural Society
dictionary index of garden plants. Griffiths M, ed. London: Macmillan.
1234 p.
Spaeth JN. 1934. A physiological study of dormancy in Tilia seed. Cornell
University Agricultural Experiment Station Memoir 169: 1B71.
Suszka B, Muller C, Bonnet-Masimbert M. 1996. Seeds of forest broadleaves,
from harvest to sowing. Gordon A, trans. Paris: Institut National de la
Recherche Agronomique. 294 p.
Vanstone DE. 1978. Basswood (Tilia americans L.) seed germination.
Combined Proceedings of the International Plant Propagators= Society 28:
566B570.
Vanstone DE. 1982. Seed germination of American basswood in relation to
seed maturity. Canadian Journal of Plant Science 62(3): 709B713.
Figure 1CTilia americana, American linden; T. cordata, littleleaf linden: fruits,
actual size.

Figure 2CTilia cordata, littleleaf linden: seed , H 4.

Figure 3CTilia americana, American linden: longitudinal section through a

seed, H 12.

Figure 4CTilia americana, American linden: seedling development at 1 day (A)

and 3 (B), 16 (C), and 19 days (D) after germination.
Table 1CTilia, linden: nomenclature, and occurrences

Scientific name(s) & synonym(s) Common name(s) Occurrence

T. americana L. American linden, basswood, New Brunswick S to Virginia

T. glabra Venten. whitewood, American lime, bee-tree & Texas

T. anericana var. caroliniana Carolina basswood SE US

(P. Mill.) Castigl.

T. americana var. heterophylla white basswood West Virginia to Florida, W to

(Vent.) Loud. Indiana & Alabama

T. cordata P. Mill. littleleaf linden, small-leaved Europe

(T. parviflora J. F. Ehrh. ex Hoffm.) lime, European linden

T. H euchlora K. Koch Crimean linden, Caucasian SE Europe & Sw Asia

(T. cordata H T. dasystyla) lime

T. H europaea L. European linden, common Europe

(T. cordata H T. platyphyllos) linden, lime
T. H intermedia DC.; T. H vulgaris Hayne

T. mexicana Schldl. Mexican basswood Mexico

T. petiolaris DC. pendent silver linden, SE Europe & W Asia

pendent white lime, weeping lime

T. platyphyllos Scop. bigleaf linden, large-leaved Europe to SW Asia

T. H europaea var. grandiflora Hort. lime, largeleaf linden

T. tomentosa Moench silver linden, European SW Europe & Asia

T. argentea DC. white linden

Sources: Dirr (1990), LHBH (1976), Plotnik (2000), Rehder (1990), RHS (1994).
Table 2CTilia, linden: growth habit and general comments

Species Growth habit & maximum height General comments

T. americana Tree to 40 m with numerous, slender, Flowers pale yellow in summer; bee plant;
low-hung spreading branches; pyramidal wood used for making expensive furniture
when young, crown somewhat rounded & excelsior, inner bark used for fabric
at maturity

T. americana var. Tree to 20 m; close to habit of C

caroliniana T. americana

T. americana var. Tree to 30 m; crown conical C

heterophylla

T. cordata Tree to 30 m; pyramidal when young; Widely planted as a street tree; pollution-
upright-oval to pyramidal-rounded and tolerant; excellent shade tree
densely branched in old age;
crown outspread

T. H euchlora Tree to 20 m Similar to T. cordata

T. H europaea Tree to 37 m C

T. mexicana Tree to 20 m C

T . petiolaris Tree to 23 m Sometimes considered as a pendulous

selection of T. tomentosa

T. platyphyllos Tree to 40 m; crown conical to broadly Not widely planted in the US

conical

T. tomentosa Tree to 27 m; pyramidal when young; Can be grown effectively as a multi-stemmed

upright-oval to pyramidal-oval in specimen to highlight light gray, smooth
later years; crown dense bark; good street tree, tolerating heat &
drought better than other lindens

Sources: Dirr (1990), LHBH (1976), Plotnik (2000), Rehder (1990), RHS (1994).
Table 3CTilia, linden: seed yield data

Seeds/wt (H 1,000)
Seed wt/fruit wt Range Average
Species kg/45.4 kg lb/100 lb /kg /lb /kg /lb Samples

T. americana C C C C 6.6 3 2
34.1 75 6.6B17.6 3.0B8.0 11 5 15+
C C 20B32.1 9.1B14.6 C C C
T. cordata 36.3 80 24.9B38.3 11.3B17.4 30.4 13.8 57+
C C
48.8B65
.1
22.2B29
.6 C
CC
T. H europaea C C 23.3B29.7 10.6B13.5 C C C
T. platyphyllos C C 25.1B30.6 11.4B13.9 C C C
T. tomentosa C C 20.0B25.1 9.1B11.4 C C C

Sources: Brinkman (1974), Heit (1977).