Ethnobryology: Usos Tradicionais de Briófitas

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Ethnobryology: Traditional Uses and Folk Classification of Bryophytes 
Author(s): Eric S. J. Harris 
Source: The Bryologist, Vol. 111, No. 2 (Summer, 2008), pp. 169-217
Published by: American Bryological and Lichenological Society
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Ethnobryology: traditional uses and folk classification 
of bryophytes 
Eric S. J. Harris 
University Herbarium, and Department of Integrative Biology, 3060 Valley Life 
Sciences Building, University of California at Berkeley, Berkeley, CA 94720-3140, 
U.S.A. 
e-mail: e_harris@berkeley.edu 
Abstract. The term 
"ethnobryology" was introduced about 50 years ago in a paper about 
the bryophytes used by the Gosiute people of Utah (Flowers 1957). Although there are 
fewer literature reports about human uses of bryophytes than those about vascular plants, 
a number of references about ethnobotanically important bryophytes do exist. These 
instances of ethnobryological use are all the more interesting both because of their relative 
rarity, and for the insights they can provide about the relation between people and small 
plants, like bryophytes, that lie on the cusp of human perception. This paper presents 
a summary of traditional uses and folk classifications of bryophytes around the world. 
Comparisons are made about the way that bryophytes are classified in different cultures. A 
list of about 150 ethnobotanical species of bryophytes is given. Many of these species have 
reported uses in Traditional Chinese Medicine (?27%) and by Native North Americans 
(?28%). The most common use of bryophytes is for medicinal purposes. Sphagnum, 
Marchant?a and Polytrichum are the most commonly reported genera to have 
ethnobotanical uses. This paper concludes by discussing how these peculiar instances of the 
human use of little plants can inform the study of ethnobotany in general. 
Keywords. Ethnobotany, Medicinal plants, Ethnobryology, Ethnotaxonomy, Bryophyta. 
The grass became a huge forest, a little creature ran about 
in it; he himself became so small that he imagined himself 
walking therein. But now instead of being, as hitherto, 
a filled space, it became an open space or extension, and 
his attitude changed as he portioned it out. (Kierkegaard 
1958: 107) 
The term 
"ethnobryology" was first introduced by the 
bryologist Seville Flowers (1957) in an article about the 
uses of bryophytes by the Gosiute people of Utah. 
Although it has been in existence for about 50 years, 
the term 
"ethnobryology" has not entered popular 
discourse in either the biological or the ethnobiological 
sciences. This is, in part, the result of two main 
factors?one is that there just are not that many 
ethnobotanical uses of bryophytes, and the other is that 
there may be a bias on the part of ethnobotanical 
researchers to focus on big plants. Consequently, 
although the ethnobotanical uses of bryophytes are 
interesting to most bryologists, it is both uncommon 
and somewhat difficult to find any ethnobryological 
reports in the literature. As an indication of this, in 
How to Know the Mosses and Liverworts Henry S. 
Conrad (1956) wrote: "Perhaps no great group of 
plants has so few uses, commercial or economic, as the 
mosses." Of course, this statement can be expanded to 
include bryophytes in general. 
The Bryologist 111(2), pp. 169-217 0007-2745/08/$5.05/0 
Copyright ?2008 by The American Bryological and Lichenological Society, Inc. 
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170 THE BRYOLOGIST 111(2): 2008 
Despite the fact that the uses of the bryophytes 
may not be as frequent as the uses of their larger, 
more 
conspicuous tracheophyte cousins, there are 
a number of interesting records of bryophyte use 
around the world. The purpose of this review is two 
fold. The first is to provide some generalizations 
about the use and knowledge of bryophytes by 
different people in different cultures. In doing so, I 
hope to emphasize the interesting aspects of people's 
use and knowledge of bryophytes that might inform 
the study of ethnobotany in general. The second 
purpose is to provide a comprehensive list of the uses 
of bryophytes, summarized from the ethnobotanical 
literature. This list was created to serve as a reference 
for those with interest in traditional uses of 
bryophytes and to provide a stimulus for further 
research in ethnobryology. Previous reviews of 
human uses of bryophytes have mainly relied on the 
bryological literature, rather than the ethnobotanical 
literature more broadly. Thus, an ancillary aim of this 
review is to provide an introduction and synopsis of 
pertinent ethnobotanical literature to bryologists. 
Two main aspects included in the field of 
ethnobotany are the study of how people classify 
plants and how people use plants. Each of these 
aspects will be explored below as they pertain to 
ethnobryology. Following this general discussion, I 
provide a list of bryophytes with ethnobotanical uses. 
This review does not include a discussion of 
bryophytes that are or have been used as indicators of 
pollution, as ornamentals in horticulture, or as fuel 
(e.g., peat). There have been comprehensive reviews 
of these topics elsewhere (e.g., Ah-Peng 8c De 
Traubenberg 2004; Ando 8c Matsuo 1984; Chapman 
et al. 2003; Glime & Saxena 1991; Iwatsuki & 
Kodama 1961; Onianwa 2001; Schenk 1997; Thieret 
1956; Turner 1993; Zechmeister et al. 2004). This 
review also does not contain a detailed discussion of 
bioactive substances in bryophytes. For that, I turn 
the readers' attention to other reviews of the topic 
(e.g., Asakawa 1990a, b; Banerjee 2001; Chopra 8c 
Kumra 1988; Schofield 1985). 
Mosses Are Small: Comparative Ethnotaxonomy 
of Bryophytes 
A natural instinct teaches [us] to learn about the nearest 
things first, and the most minute last, for example... the 
larger plants first, and the smallest mosses last. (Linn? 
2002: 112) 
Bryophytes are smaller and less conspicuous 
than vascular plants. This in no way detracts from the 
beauty of bryophytes, but the fact that they lie on the 
cusp of what is visible does mean that they are often 
ignored by humans. Humans perceive their 
environment at a certain spatial and temporal scale. 
Consequently, humans are biased to only seeing 
things of a certain size. For the most part, humans 
deal in sizes ranging in magnitude from centimeters 
to kilometers, and times ranging from seconds to 
decades. For the average person, a micrometer is not 
a 
meaningful measure of size. By contrast,most 
measurements of 
morphological features in 
bryophytes are in micrometers, centimeters at the 
largest. 
Arguably, the most important invention in the 
history of bryology was the microscope. The 
landmark work in the taxonomy of bryophytes, 
Hedwig's Species Muscorum (1801), was published in 
the decades following the invention of the 
microscope. As an indication of the importance of 
this new technology, the frontspiece to the Species 
Muscorum features a person using a microscope to 
examine a moss specimen. Though moss taxonomy 
did exist prior to Hedwig (e.g., Dillenius 1763), most 
works on moss taxonomy before the Species 
Muscorum was published recognized few types of 
mosses. For example, Dillenius only recognized six 
genera of moss, in contrast to Hedwig's 36, and 
Dillenius' classification also included lichens, 
lycophytes and algae as types of "moss." Most 
traditional classifications of the botanical world do 
not 
rely on the aid of the microscope. As a result, 
cultures around the world 
rarely recognize bryophyte 
species as distinct things. In a study on the relation 
between the size of organisms and their distinctness 
in folk taxonomies, Hunn (1999) noted that there is 
a general cross-cultural trend of lumping small 
organisms into more broadly inclusive categories 
than those 
recognized by scientific taxonomy. Berlin 
(1992) also noted that groupings of small organisms 
in folk classification systems which linguistically 
correspond to folk genera are more similar to higher 
level ranks, such as classes or orders within the 
classifications proposed by scientific taxonomists. 
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That is, scientific classifications discern many more 
little organisms than those classification systems 
made without the aid of the technological edifice 
underlying scientific taxonomy. 
In general, bryophytes are traditionally 
categorized along with other little plants in a waste 
basket group of things that lack distinctness to 
humans. In his book, Cognitive Foundations of 
Natural History, Atran (1990) noted: 
" 
...the non-flowering plants... may generally be construed 
as "residual" categories with no clearly defined morpho 
logical aspect. Ray's Musci...is not so much the artificial 
product of some a priori system... but the perceptual 
remainder of those small and often hidden plants that lack 
phenomenal resolution for human beings." (Atran 1990: 
32) 
For example, the English word "moss" has been used 
to refer to a number of what we now consider very 
different types of plants: "true" moss refers to 
bryophytes, cup-moss refers to the lichen Cladonia 
spp., Irish moss refers to the red algae Chondrus 
crispuSy Spanish moss refers to the angiosperm 
Tillandsia usneoides, club moss refers to the 
Lycopodiaceae, and so on (Welch 1948: 32). In his 
classification, Dillenius (1763) followed this 
historical folk classification by including lichens, 
algae, lycophytes and bryophytes in his treatment of 
"moss." The pattern of using one general term to 
refer to a category of many evolutionarily disparate, 
little plants is repeated in many languages around the 
world. For example, Pacific Islanders use the term 
"limu sa" (Uhe 1974), the Seri in the southwestern 
U.S. and Mexico use the term "Yamasa" (Felger 8c 
Moser 1985), the Navajo in the southwestern U.S. use 
the term "dlad" (Franciscans 1910), the Karok in the 
Pacific Northwest of the U.S. use the term 
"asaxxe'm" (Schenck 8c Gifford 1952), the Chinese 
use the term 
"qing tai" (ff) (Beijing Foreign 
Studies University 1981), the Shuar in Ecuador use 
the term 
"juu" (Bennet et al. 2002) and so on. The 
name "liverwort" has, to a lesser degree, also been 
applied to organisms other than bryophytes. For 
example, in Wren (1988) the name "liverwort" refers 
to both the angiosperm Hep?tica nobilis and the 
lichen Peltigera canina. 
It should be noted that there is a difference 
between recognizing different plants and naming 
Harris: Review of ethnobryology YJ1 
different plants. Different plants may be recognized 
as distinct without giving those different plants 
a distinct name. The possibility exists that people 
may recognize many different types of bryophytes, 
but they do not label those differences. It is difficult 
to prove or disprove this statement, but one could 
argue that any plant with economic or cultural 
importance would most likely have a distinct name, 
even if that distinct name was used only among the 
botanical specialists of a culture. In any case, it 
remains true that many cultures have only a single 
linguistic term that groups together a diverse host of 
little green things, including bryophytes. 
Ethnotaxonomy is the branch of ethnobotany 
concerned with investigating folk classifications and 
forming generalizations about the way people classify 
the natural environment around them (Berlin et al. 
1973; Medin 8c Atran 1999). In many cases, 
ethnotaxonomy is concerned with forming 
generalizations about folk classification systems that 
could be construed as universals of the human mind 
(e.g., Berlin 1992). That is, some ethnotaxonomists 
have attempted to uncover psychological universals 
that explain how people around the world classify 
their environment. The occurrence of a miscellany 
category for little plants seems to be a general feature 
of folk classifications, irrespective of culture. More 
succinctly, humans, regardless of their culture, are 
naturally inclined to ignore bryophyte diversity. 
However, often in the cases when different 
bryophytes or little plants are lumped together in one 
category, they are categorized according to where 
they are growing. Across cultures, some of the most 
common types of moss and moss-like things are 
referred to by terms such as "tree moss," "rock 
moss" and 
"ground moss." Turner (2000) noted that 
this is a general trend among language groups in 
northwestern North America. Within these language 
groups, there is a general term for moss and moss 
like things. This general term is modified by habitat 
and growth form to distinguish among moss and 
moss-like things (Turner 2000: 107). In another 
example, among the Tzeltal of Mexico there are three 
names for bryophytes that refer to the places where 
the bryophytes grow: con te1 refers to bryophytes that 
live on trees, con lumilal refers to bryophytes that 
grow on the ground and cone en refers to bryophytes 
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172 THE BRYOLOGIST 111(2): 2008 
in caves (Berlin et al. 1974). Similarly, Turner et al. 
(1980) noted that the Okanogan-Colville First 
Nations in Canada distinguish four types of "moss." 
This category includes mosses, liverworts, Selaginella 
and lichens. Nkweskwesp?la7xw refers to "moss" on 
the ground, tkweskwespisxn refers to "moss" on 
rocks, tkweskwesp?lekw refers to "moss" that grows 
on wood and nkweskwespitkw refers to "moss" that 
grows in the water. The Thompson First Nation in 
British Columbia, Canada also distinguishes 
bryophytes according to their ecological conditions 
and general growth form. The Thompson First 
Nation use the term /qwz?m to refer to mosses, 
liverworts, lichens and other moss like plants (Turner 
et al. 1990: 85). This term is modified to describe 
"long moss" (Hylocomium splendens)> "short moss" 
(Eurhynchium oreganum, Rhytidiadelphus triquetrus), 
"rock moss" (Selaginella, Racomitrium canescens, 
etc.), "water moss" (Fontinalis) and"swamp moss" 
(Sphagnum) (Turner et al. 1990: 85). 
In addition to being distinguished by habitat, 
some "mosses" are thought to have medicinal 
properties that relate to their location of growth. Not 
only are those "mosses" labeled according to their 
ecology, but their essence is fundamentally linked to 
it. For example, some European herbal books report 
that the medicinal properties of tree-moss "...doth 
partake of the nature of the tree from whence it is 
taken..." (Culpeper 1840: 224). Although "tree 
moss" is probably the lichen Evernia, whose common 
name is "oak moss" or a species of the lichen Usnea 
(Bland 1971: 106; Wickens 2001: 365), it is relevant 
that not only is this "moss" defined by its ecological 
location, but its medicinal properties are dependent 
on the type of tree where it is growing (see also 
Gerard 1927: 281). This reasoning presumably also 
applies to bryophytes with medicinal uses. 
The phenomenon of defining bryophytes by 
a combination of their location of growth and 
morphological features is also exemplified by the 
liverwort Marchant?a polymorpha and its 
nomenclatural brethren in Traditional Chinese 
Medicine. Marchant?a polymorpha is used as 
a medicinal herb in China, where it is sometimes 
called DiFuPing (e.g., Jiangsu New Doctor College 
1974). This name literally means "earth-duckweed." 
Duckweed (Lemna) is usually referred to as FuPing. 
The fern Azolla is sometimes called HongFuPing, or 
red-duckweed. In this case, FuPing can refer to little 
plants that may be angiosperms, liverworts or water 
ferns (Jiangsu New Doctor College 1974; Wang et al. 
2001). In this example, a single term subsumes 
evolutionarily different types of plants that 
morphologically resemble one another and, in the 
case of Lemma and Azolla, occupy similar ecological 
niches. The different plants are distinguished both by 
ecological context, as in the "earth-duckweed" or by 
morphology, as in the "red-duckweed." 
In contrast to the general trend of lumping small 
plants together, a few bryophytes are recognized as 
distinct. These bryophytes are distinguished 
according to features other than ecology. 
Plagiomnium insigne is uniquely specified in the 
languages of the Oweekeno and Bella Coola of the 
central coast of British Columbia where it is used 
medicinally (Compton 1993; Turner 1973). The 
name in Oweekeno (OCPc?maimnixwsAaus) refers to 
the morphology of this moss and literally means 
"tiny, tiny little trees." In an example from southwest 
China, the genus Rhodobryum is distinguished from 
other mosses. In fact, many people in this area of 
China do not consider Rhodobryum to be a moss 
(QingTai = ff) at all, but consider it a type of 
herb or grass (cao 
= 
^) (pers. observ.). The species 
R. giganteum is perhaps most commonly used there, 
but other species in the genus, such as R. roseum and 
R. spathulatum, are also collected and used 
medicinally (pers. observ.) (see Fig. 1). The name for 
Rhodobryum in Yunnan is HuiXinCao (HPLVJf?), 
a name 
referring to the heart-healing properties of 
this plant. The name literally means "return-the 
heart-herb." Both of these mosses are relatively 
large?Plagiomnium insigne is usually 3-8 cm high 
(Lawton 1971), and Rhodobryum giganteum can grow 
to be as large as 6 cm high and 4 cm wide (Eddy 
1996). Presumably these mosses are more accessible 
to human perception and more likely to have 
a distinct name. 
There are also some cases where differences 
below the species level are recognized. In the example 
given above, the Oweekeno recognize two kinds of 
Plagiomnium insigne with varying medical efficacy. 
Compton (1993) reported that it was known that P. 
insigne grows under both Douglas-fir trees 
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Harris: Review of ethnobryology 173 
Figure 1. A vendor in Yunnan Province, China displaying locally collected herbs, including Rhodobryum giganteum (in the bag in 
her hands and in the bag to her left). This moss is used in Yunnan Province to treat nervous disorders and cardiovascular disease 
(Dai et al. 2006; Gao et al. 2004; Wang et al. 2005; Yan et al. 1998; Zhou et al. 2004). Photo by Eric Harris ?2006. 
(Pseduotsuga menziesii) and spruce trees (Picea), but 
was considered more medically effective if collected 
under spruce trees. In another example, Gottesfeld 
and Vitt (1996) noted the recognition of different 
types of Sphagnum moss used as diapers among the 
Wet'suwest'en and Gitksan people of British 
Columbia in Canada. In this example, people 
recognize different types of Sphagnum and 
differentiate them according to color and size. In one 
case, the authors noted the avoidance of red-colored 
variants of Sphagnum due to the belief that these 
variants would cause skin irritation (Gottesfeld 8c 
Vitt 1996). People of the Carrier First Nation of 
British Columbia also avoid using the red variants of 
Sphagnum (Carrier Linguistic Committee 1973: 86). 
In an example from Southwest China, different types 
of the herb HuiXinCao (Rhodobryum) are sometimes 
recognized by herbalists and herb collectors. However, 
in many cases these different types of HuiXinCao 
come from different species of plants that might be 
considered quite unrelated by western taxonomists. 
People recognize a big (da = ^) and a little (xiao = 
'JN) kind of HuiXinCao, In this folk classification, the 
small type refers to Rhodobryum whereas the big type 
refers to plants other than Rhodobryum. In one 
instance, the big type was identified as the large moss 
Pogonatum ?rratum, which can grow as tall as 10 cm 
or more (Wu 8c Wang 2005: 331). But in other cases 
the big type was indicated as a species in the genus 
Ledum (pers. observ.). This particular taxonomic 
grouping could arise because the name HuiXinCao 
refers to medical effect, rather than morphology. The 
different kinds of HuiXinCao are all used for the same 
medicinal purpose?namely, the treatment of minor 
heart problems. 
If bryophytes are rarely named as distinct things, 
it may seem strange that certain species of bryophytes 
have ethnobotanical uses at all. The reasons that 
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174 THE BRYOLOGIST 111(2): 2008 
specific bryophytes may come to have cultural 
importance may be as many as there are useful 
bryophytes. But there are two different explanations 
that might account for ethnobotanically important 
bryophytes, two of which I will briefly mention here. 
The first is necessity. The Oweekeno First Nation of 
British Columbia live in an isolated area along the 
central coast of British Columbia. To this day, the 
village is not accessible by road, and the easiest way 
to get to the village is either by plane or boat. A 
member of the village band council noted that 
because of their relative isolation, the Oweekeno have 
had to rely on everything available to them, whether 
it be fishing off the coast, or moss on the forest floor 
(pers. observ.). The second possible explanation for 
use of bryophtyes is comprehensiveness. The moss 
Rhodobryum giganteum is a popular herbal remedy in 
southwest China (Gao et al. 2004; Lei et al. 2001a, b; 
Yan et al. 1998; Yu 8c Ma 1993; Yu et al. 1994, 1995; 
Zhou et al. 2004). The reason that it is so popular is 
probably a result of the comprehensive nature of the 
system of Traditional Chinese Medicine (TCM). 
Thousands of different plants, animals and minerals 
are used as components of Chinese medicine. Several 
encyclopedias have been written to enumerate the 
various ingredientsused in TCM (e.g., Jiangsu New 
Doctor College 1974; ZhongHuaBenCao 1999). 
Among these ingredients, several bryophytes have 
been used for medicinal purposes in China 
(ZhongHuaBenCao 1999). In fact, mosses have a long 
history of use in TCM. For example, the 
Compendium of Materia Medica (Li 2004) written in 
the 16* century by the famous Chinese doctor, Li 
ShiZhen, notes that Polytrichum (TuMaZong = 
Horse-Mane of the Earth) can be used as a diuretic 
(see Fig. 2) (Fan et al. 2004: 1555). Wu and Jia (2003: 
52) provided a short history of the ways that 
bryophytes figure into ancient Chinese medicinal 
texts. In their discussion, these authors noted that 
bryophytes were probably used medicinally as early 
as the 6* century in China, and continue to be used 
in areas of southwest China such as Yunnan and 
Sichuan provinces (Wu 1982: 5; Wu 8c Jia 2003: 52). 
Summary of Bryophyte Uses 
Bryophytes have many different uses around the 
world. However, with the exception of peat 
?? ? 
Figure 2. This image represents one of the first ethnobotanical 
records of a medicinal moss. It was originally published in the 
16th century in the Compedium for Materia Medica (Li 2004). 
This herb is called i-SjJft (= TuMaZong,) or "Horse-Mane of 
the Earth" and has been identified in the Traditional Chinese 
Medicine literature as a species of the Polytrichaceae, usually 
Polytrichum commune. 
(Sphagnum), there are relatively few bryophytes that 
are commercially significant (Frahm 2004). I have 
recorded about 150 different species of bryophytes 
with traditional ethnobotanical uses. Appendix 1 is 
a taxonomic list of these bryophytes and their uses. 
Following this list, Appendix 2 gives information 
about bryophyte uses that did not have any 
associated scientific name. These records of 
ethnobotanical use have mainly been culled from the 
primary literature, but some rely on my own 
observations on medicinal moss use in southwest 
China. Most of the records of Chinese medicinal 
bryophytes were translated from the Chinese medical 
literature specifically for this paper. Note that this 
review does not include uses of bryophytes in 
horticulture (see Ando 1980; Ando & Matsuo 1984; 
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Gallnut Production 
(?) 
Absorbent (9) 
Cleaning (9) 
Bedding (It) 
Packing (13) 
Misc. (16) 
Decoration (26) 
Medicinal [77) 
Chinking (22) 
Figure 3. Relative proportion of different uses of bryophytes. 
Total number of species in each use category is indicated in 
parentheses. Only bryophytes classified to genus or species were 
counted. Species with multiple uses were counted in each 
category of use. Consequently, although a total of 150 different 
species are noted in the text, the total number of useful 
bryophytes shown here is 189. Miscellaneous category includes 
such uses as dyes, perfumes and creation of lampwicks. 
Iwatsuki & Kodama 1961; Schenk 1997), as peat (see 
Ando 1980; Ando & Matsuo 1984; Chapman et al. 
2003; Glime & Saxena 1991, Turner 1993) or as 
bioindicators (see Ah-Peng & De Traubenberg 2004; 
Ando 1980; Ando & Matsuo 1984; Onianwa 2001; 
Zechmeister et al. 2004). The majority of bryophytes 
with recorded uses are mosses and liverworts, but 
there is one report of the potential use of the 
hornwort, Anthoceros (see Appendix 1). Appendix 3 
provides a list of all ethnobotanical bryophytes 
according to their family classification. 
The various different uses of bryophytes are 
summarized in Fig. 3. Almost half of these 
bryophytes are used medicinally, accounting for 
about 41% of the total bryophytes that have reported 
uses. The next most common use of bryophytes is for 
decoration, which accounts for about 14% of useful 
bryophytes. The third most common use of 
bryophytes is for use as chinking material in the 
construction of log cabins or for caulking boats 
(shown in the category "chinking"). This accounts 
for about 12% of useful bryophytes. Other uses 
include bedding, packing and cleaning, each of which 
accounts for about 5% of the total ethnobotanical 
species. Lastly, about 8% of ethnobotanical 
bryophytes have miscellaneous uses. This category 
Harris: Review of ethnobryology 175 
includes uses of bryophytes for such purposes as 
dyes, lamp-wicks, perfumes and others. Another use 
of bryophytes is for the production of gallnuts in 
China (Li & Longton 1993). In this case, mosses are 
grown to encourage the growth of gallnut-producing 
aphids, which over-winter in species such as 
Plagiomnium maximoviczii (see Fig. 4). Obviously, 
uses of bryophytes for chinking log cabins or as 
packing material do not rely on specific properties of 
the species used. Many kinds of mosses could be used 
to fill the chinks in a log house (Bland 1971). But this 
is not necessarily the case with medicinal bryophytes 
because medicinal uses of bryophytes are presumably 
specific to the species or genus involved. Lastly, note 
that there are no records of bryophytes that are used 
as food. The only reference that mentions bryophytes 
that are eaten is in Thieret (1956), who mentioned 
the use of Sphagnum as a famine food in China and 
a last resort in bread baking in Lapland. Some 
references indicate the use of bryophytes in 
preparation of drinks, such as the use of Rhodobryum 
as a medicinal tea (YunNanZhongCaoYaoXuan 1970: 
250), or Lunularia and Plagiochasma in the 
preparation of maize beer (Franquemont et al. 1990: 
35). However, the general paucity of references on 
the use of bryophytes for food can be taken as 
indication that bryophytes have little or no value as 
a nutritional source for humans. 
The distribution of ethnobotanically important 
bryophytes by geographic region is shown in Table 1. 
China has the most records of ethnobotanically 
important species (~63 spp.). This may, in part, be 
a result of my focus on the medicinal bryophytes of 
China, but this pattern is reflected in other papers 
that discuss ethnobryology (e.g., Asakawa 1990a). 
Most of these species are used in herbal medicine. 
However, it is unlikely that these are all in common 
use. Certainly some bryophytes are still used in 
Chinese medicine, such as the moss Rhodobryum 
giganteum (pers. observ.). This herb is still frequently 
sold in markets in southwest China, and has been 
recently studied by Traditional Chinese Medical 
doctors for its efficacy. These medical studies have 
collectively helped to show that R. giganteum is useful 
in treating cardiovascular problems, and have helped 
illustrate the physiological mechanism of the herb 
(Dai et al. 2006; Gao et al. 2004; Lei et al. 2001a, b; 
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176 THE BRYOLOGIST 111(2): 2008 
Figure 4. Growing moss for the production of gallnuts in southwest China. The scaffolding to the right is holding large plates of 
Plagiomnium maximoviczii that host gall-aphids in the winter. During the summer the plates were placed underneath Chinese 
sumac (Rhus chinensis) so the gall-aphids could produce the tannin rich gallnuts on sumac leaves (for more information see Li et al. 
1988; Li & Longton 1993). Photo by Yang Bilun ?2006. 
Wang et al. 2005; Yan et al. 1998; Yu & Ma 1993; Yu 
et al. 1994, 1995; Zhou et al. 2004). In addition to 
ethnobotanical bryophytes in Asia, there are also 
many species with ethnobotanical uses in the United 
States and Canada. Most of these reports refer to uses 
by Native Americans. As in the case of current uses of 
bryophytes in Chinese herbal medicine, very few 
species of bryophytes are still usedcommonly in 
North America, although it may be that Sphagnum is 
still used by some Native American groups (e.g., 
Gottesfeld & Vitt 1996). 
There are 22 bryophyte species with records of 
ethnobotanical use in India, including one 
ethnobotanical species used in the Nicobar Islands 
(Dagar & Dagar 1999). Many of these ethnobotanical 
reports come from an article about the Kumaun 
region in Uttar Pradesh written by Pant and Tewari 
(1989a) and reports about the use of Plagiochasma 
appendiculatum in Himal Pradesh (Kumar et al. 
2000; Singh et al. 2006). Collectively, these reports 
refer to the medicinal use of bryophytes only in 
a relatively limited area of India, in the Himalayan 
region. However, despite the large and 
comprehensive literature on plants used in Ayurvedic 
medicine, no references were found that describe the 
medicinal use of bryophytes in Ayurveda. To 
emphasize this, in an article on economic plants of 
India Watt (1889: 85) remarked that "...few 
(bryophytes) are of any economic value, and, so far 
as the writer is aware, none is used in any way in 
India." It is interesting that both the Ayurvedic and 
Traditional Chinese Medicine systems are very 
comprehensive, but whereas more than 50 
bryophytes have reported uses in Chinese herbal 
texts, there are few, if any, reported in Ayurvedic 
texts. Additionally, besides one report on the use of 
the genus Timmiella in Egypt (Brent Mishler, pers. 
comm., 2006), no citations could be found of the use 
of bryophytes in Africa. This does not necessarily 
mean that there are no uses of bryophytes in Africa, 
but it does suggest that ethnobotanical use of 
bryophytes is not common. For example, there may 
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Table 1. Distribution of useful bryophytes by geography. Note 
that the most species are used in China, followed by the U.S.A. 
and Canada. Almost half of all bryophyte uses are reported 
from Asia. Note the relative lack of references of bryophyte use 
in Africa. If the same species were used in different countries, 
they were counted separately for each country. Consequently, 
although a total of 150 different species are noted in the text, 
the total number of useful bryophytes shown here is 235. 
REGION # SPECIES % of TOTAL 
Africa 1 
Egypt 1 
Asia 90 
China 63 
India 22 
Japan 1 
Vietnam 4 
Europe 41 
Country not specified 8 
France 2 
Germany 4 
Iceland 1 
Spain 5 
Sweden 3 
UK 18 
North America 69 
Canada 29 
Mexico 4 
U.S.A. 36 
Oceania & Australia 9 
Australia 1 
Easter Island 1 
Fiji 1 
New Zealand 4 
Papua New Guinea 2 
South America 25 
Bolivia 17 
Chile 
Colombia 
Cuba 
Ecuador 
Guatemala 
Peru 
0.4 
0.4 
38.3 
26.8 
9.4 
0.4 
1.7 
17.4 
3.4 
0.9 
1.7 
0.4 
2.1 
1.3 
7.7 
29.4 
12.3 
1.7 
15.3 
3.8 
0.4 
0.4 
0.4 
1.7 
0.9 
10.6 
7.2 
0.4 
0.4 
0.4 
0.4 
0.4 
1.3 
be ethnobotanical uses of bryophytes in Madagascar 
(Kirsten Fisher, pers. comm., 2001), but there are no 
literature reports of these uses. Appendix 4 lists all 
ethnobotanical bryophytes according to the country 
where they are used. 
Table 2 summarizes the most commonly used 
bryophyte genera. As a proxy, genera are listed 
according to the number of countries where they are 
Harris: Review of ethnobryology YJJ 
Table 2. Genera of bryophytes used in more than two 
countries. As previous reviews of ethnobotanical bryophytes 
have indicated (e.g., Thieret 1956), the bryophytes with the 
most widespread usage are in the genera Sphagnum^ March 
ant?a and Polytrichum. 
GENUS # of Countries Where Used 
Sphagnum 13 
Marchant?a 12 
Polytrichum 9 
Conocephalum 5 
Climacium 4 
Hylocomium 4 
Hypnum 4 
Rhytidiadelphus 4 
Thuidium 4 
Antitrichia 3 
Bryum 3 
Dicranum 3 
Fontinalis 3 
Funaria 3 
Philonotis 3 
Pleurozium 3 
Rhizomnium 3 
used. The top three genera that are used in different 
countries are Sphagnum, Marchant?a and 
Polytrichum. Perhaps for this reason, these three 
genera are most commonly noted in discussions 
about useful bryophytes (e.g., Ando & Matsuo 1984; 
Glime & Saxena 1991; Thieret 1956; Welch 1948). 
Widespread use of one plant for similar purposes, 
where the uses arise independently in different 
cultures, is suggestive evidence for the efficacy of 
a certain plant. This is best exemplified by the various 
uses of Sphagnum that relate to its absorbent 
properties, such as a surgical dressing, diaper or 
sanitary napkin. Sphagnum has been used as a diaper 
and sanitary napkin by many different cultures such 
as Native Americans in the U.S.A. and Canada 
(Adelson 2002; Kimmerer 2003) and the Maori in 
New Zealand (Macdonald 1974). And Sphagnum is 
one of the most well known mosses due to its use as 
a surgical dressing during World War I (Ando 1980; 
Hotson 1918, 1921a, b; Pinheiro et al. 1989). These 
apparently separate and similar uses suggest that 
Sphagnum is undoubtedly a useful absorbent and 
bandaging material, a fact corroborated by recent 
pharmacological studies (e.g., Painter 2003). In 
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178 THE BRYOLOGIST 111(2): 2008 
another example, Marchant?a is used in the same way 
in some countries, but has different uses in others. It 
is used for liver ailments in Europe (Thieret 1956) 
and South America (Garc?a Barriga 1992; Roig y 
Mesa 1945) but is used for skin disorders in India 
(Pant 8c Tewari 1989a). Interestingly, the uses of 
Marchant?a in these areas were determined by the 
doctrine of signatures (Pant 8c Tewari 1989a; Thieret 
1956; Watt 1889). However, in Europe it was 
determined that Marchant?a resembled a liver and 
was used accordingly, but in India it was determined 
to look like a small boil (Pant 8c Tewari 1989a). 
Marchant?a is also used primarily in China for 
external ailments, such as burns or cuts (Yan et al. 
1999). Polytrichum, too, has some similar uses in 
different areas. In China it is used primarily as 
a diuretic or to stop bleeding (Ando 8c Matsuo 1984; 
Ding 1982), but early reports of this moss also note 
that it can also be used to make hair long and black 
(Li 2004). In Europe it has been used to strengthen 
and beautify women's hair. This latter use is derived 
from the doctrine of signatures due to the 
propinquity between the hairs on the calyptra of 
Polytrichum and human hair (Ando 8c Matsuo 1984: 
148; Bland 1971: 108; Crum 1983: 377). It is not clear 
if the use of Polytrichum in China for hair was 
derived from the doctrine of signatures. Thus the 
efficacy of Sphagnum seems well supported by its 
widespread use, but further study of Polytrichum and 
Marchant?a would be required to make any 
conclusions about the medicinal effect of these two 
bryophytes. 
Conclusions 
Despite the relative rarity of scientific reports of 
human uses of bryophytes, it is clear that bryophytes 
can and do play a direct role in people's lives. 
Although bryophytes lie at the cusp of what is visible 
to humans, different cultures may develop traditions 
that incorporate bryophytes in different ways. These 
different instances are interesting because they can 
illustrate some of the specific circumstances that 
bring people into contact with the smaller things in 
their biological environment. For example, 
a 
comprehensive system of herbal medicine, such as 
that employed in China, can allow access to 
potentially useful plants likebryophytes, whereas 
other cultural traditions may not be as engaged with 
the world of little plants. Thus, examinations of the 
cultural reasons for the use of a specific plant may 
gain insight from studying those institutions and 
contingencies at play in the ethnobotanical use of 
bryophytes. It may be expected that a more particular 
edifice of cultural technology might need to be in 
place for people to become engaged with the smaller 
plants in their environment. Although there are 
relatively few uses of bryophytes as compared to 
tracheophytes, they can act as a fruitful system in 
which to study the cultural institutions and 
contingencies that bring people to plants. In addition 
to cultural studies, further investigations on the 
pharmacological properties of popular medicinal 
bryophytes such as Marchant?a, Polytrichum and 
Sphagnum are also warranted to explore the 
medicinal potential of these and other useful 
bryophytes, such as Rhodobryum (e.g., Dai et al. 2006; 
Gao et al. 2004; Wang et al. 2005; Zhou et al. 2004). 
Acknowledgments 
This paper was prepared as part of a dissertation submitted at 
the University of California, Berkeley, in partial fulfillment of 
the requirements for a Ph.D. in the Department of Integrative 
Biology, conducted under the supervision of Prof. Brent 
Mishler and Dr. Tom Carlson. I would like to thank Jim 
Griesemer and Isao Kubo for comments on early drafts of the 
manuscript, Momei Chen for comments on some of the 
Chinese translations and Hieu Pham for his translations of 
Vietnamese. I would like to thank Yang Bilun, Kirsten Fisher, 
Brent Mishler and Jim Shevock for sharing information about 
ethnobryology with me. I would also like to thank the 
reviewers, Michael Balick (NYBG) and Janice Glime (MTU), 
for many helpful comments and insights. And I would like to 
thank the editor, Bill Buck, for his patience and helpful 
suggestions. Work for this paper was partially supported by 
a Department of Integrative Biology Franklin Henry award, 
a University of California Pacific Rim Research Program 
(UC-PRRP) minigrant and a Natural Science and Engineering 
Research Council of Canada (NSERCC) postgraduate 
fellowship. 
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Appendix 1. 
An Ethnobryological List. 
Below is a list of bryophytes (hornworts, liverworts 
and mosses) that have reported uses. The list is 
a 
compilation of reports from ethnobotanical texts, 
compendia of herbal medicines and some texts that 
focus specifically on ethnobotanically important 
bryophytes. Other references that review the uses of 
bryophytes are: Ando (1980), Ando and Matsuo 
(1984), Asakawa (1990a), Glime and Saxena (1992), 
Pant (1998), Saxena and Harinder (2004), Thieret 
(1956), Welch (1948) and Wu and Jia (2003). 
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Additionally, the reader is encouraged to look at the 
only article that uses the word "ethnobryology" 
(Flowers 1957). 
A caveat should be made about relying on 
literature reports of plant use. As with many 
ethnobotanical uses of plants, some bryophytes were 
used mainly in the past. It appears that fewer and 
fewer bryophytes are used today as a result of the 
general decline in ethnobotanical knowledge around 
the world. So, although a plant may have a reported 
use in the literature, it may not be as popular now. 
Many of the bryophytes used by Native Americans 
have followed this fate. Another problem with relying 
solely on literature reports is that certain bryophytes 
may be over-reported. For example, Vo (1999) 
included a list of four medicinal bryophytes in 
Vietnam. It seems that these four medicinal 
bryophytes were cross-referenced from Chinese 
herbal literature, as V? (1997, 1999) noted, and may 
be used very rarely, if at all, in Vietnam. None of 
these bryophytes was known or found in herbal 
shops or hospitals in Hanoi, or by herbal doctors and 
herb sellers in northern Vietnam along the Chinese 
border (pers. observ.). 
The following list is grouped according to the 
major lineages of bryophytes: hornworts, liverworts 
and mosses. Within each grouping, the bryophytes 
are arranged alphabetically by genus, with entries for 
some families also included. In cases where the 
bryophyte was only cited to genus, it is listed as such. 
Scientific names were sometimes changed to reflect 
current taxonomy (Crosby et al. 1992), but the 
original scientific name listed in the ethnobotanical 
report is also noted. The countries where the 
bryophyte is/was reported to have uses are written in 
small capital letters. In the cases that the plant was 
used in different countries, the countries are listed 
alphabetically. Following the country name, the 
ethnic group or specific geographic region relevant to 
the ethnobryological report is provided. If known, 
the local name of the bryophyte is given and 
translated. Words that have been transcribed from 
other languages have followed the orthography from 
the cited reference. For this reason, different 
orthographic systems appear in this paper. For 
example, 
" 
7 ", 
" 
? 
" 
and 
" 7 " can all to refer to the 
presence of a glottal stop, and are employed by 
Harris: Review of ethnobryology 185 
different references used for this paper. The reader is 
encouraged to check the original reference for more 
information about the orthographic system 
employed in each case. 
Where multiple references existed for the use of 
any particular bryophyte, attention was made to 
preserve as much information as possible, rather than 
summarizing. One of the goals of this list is to give 
the reader as much detail as possible since many of 
the references used in this list, especially those in 
other languages, can be difficult to obtain. As 
a consequence, there may be some redundancies in 
information. Information was only combined in 
cases where the information in different references 
was identical; usually this is an indication that those 
references got their information from the same 
primary source. Following the taxonomic list, 
Appendix 2 provides a list of general entries for 
references that did not indicate any specific type of 
bryophyte. These latter references are arranged 
alphabetically by country. Appendix 3 provides a list 
of all ethnobotanical bryophytes according to their 
family classification. Appendix 4 lists all 
ethnobotanical bryophytes according to the country 
where they are used. 
Lastly, many of the Chinese herbal texts used in 
the following list were translated specifically for this 
paper. The use of bryophytes in Chinese medicine 
should be interpreted within that cultural context. 
Descriptions of disease, symptoms and preparation 
of Chinese herbs need to be understood in terms of 
the Chinese medical system as a whole. Chinese 
medical terms are 
notoriously difficult to translate 
(Xie 2003). For example, phrases such as "nourishing 
Yin" and 
"clearing heat" need to be understood in 
the context of the system of Traditional Chinese 
medicine,