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1981 Moss Sensitivity to Sulfur Dioxide John W. Wilcut Eastern Illinois University This research is a product of the graduate program in Botany at Eastern Illinois University. Find out more about the program.

Recommended Citation Wilcut, John W., "Moss Sensitivity to Sulfur Dioxide" (1981). Masters Theses. 2965. https://thekeep.eiu.edu/theses/2965

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Date Author

m Moss Sensitivity to

Sulfur Dioxide (TITLE) •

BY

John W. Wilcut �

THESIS

SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS

FOR THE DEGREE OF

Master of Science IN THE GRADUATE SCHOOL, EASTERN ILLINOIS UNIVERSITY

CHARLESTON, ILLINOIS

1981 YEAR

I HEREBY RECOMMEND THIS THESIS BE ACCEPTED AS FULFILLING

THIS PART OF THE GRADUATE DEGREE CITED ABOVE

\)y • i_YI �'fEl

.{)II<-:/

SULFUR DIOXIDE

BY

JOHN W. WILCUT

B.S. in Botany

Eastern Illinois Univers ity , 1978

ABSTRA CT OF A THESIS

Submitted in partia l fulfillment of the requireme nts for the degree of l\iasters of Science in Botany at the Gradua te School of Eastern Ill inois University

CHARIESTON , ILLINOIS

�

4·1.1.�29 ABSTRACT

Extensive research regarding the effects of sulfur

on vascular and nonvascular plants has been dioxide ( S02 )

reported . Sulfur dioxide has been shown to interrupt

normal metabolism , to interrupt normal reproduction, and

to alter the plant's morphology.

One hypothesis regarding the lack of bryophyte s in

urban areas is that the life cycle is be ing interrupted.

It has been shown that moss protonemata are sens itive to

so2 conc entrations in urban areas . Several authors noted

that sexual reproduction is blocked among many species which do occur in urban environments . Observations on

protonemal sensitivity are probably more important than the observations on the blockage of sexual reproduction

sinc e reproduction in mosses involves a protonemal stage .

Although moss es have not been studied as extens ively as lichens , published research has shown that mosses are at least as sens itive to air pol lutants as are liche ns .

i Unfortunately , most studies of air pol lutant effects on

mos ses have utilized unrealistic concentrations of as so2

compared to those to be encountered in urban are as . In

this study , some more realistic concentrations have so2

been used.

The focus of this research was the investigation of

the effects of on various age s and species of moss so2 protonemata. The mos ses use d in this investigation were

A ulacomnium heterostic hum (He dw . ) B.S.G., Bartramia

pomiformis Hedw., Leucobryum glaucum Agstr . ex Fr ., and

Polytrichum ohioense Ren. & Card . The protonemata of

these mosses were fumigated under varying conc entrations so2

(0.2 ppm , 1.0 ppm, 5.0 ppm ) in a flow-through chambe r for

eight hours . For each concentration, three different so2

flowrates liters pe r minute, 2 lpm, 4.0 lpm were (1.0 .0 )

used. The se flowra tes and concentrations resulted in nine

different fumigation treatments for each moss . In addition,

the protonemata and resulting young game tophytes fumigated,

were 2-rnonth, 4-month, and 8-month old cultures .

This research included using different flowrates

through the fumigation chamber to allow a comparison of

ii both flowrate effects and so2 conce ntrations on chlorophyll des truction.

Chlorophyll analysis, after S02 fumigation, revealed both a species and age different ial response to chlorophyll destruction from S02• Leucobryum glaucum was found to be the most so2- sensitive and Bartramia pomiformis somewhat less . Aulacomnium heterostichum and Polytrichum ohioense were found to be more so2 -resistant . Age of the moss

- game tophyte was shown to be an important factor in so2 resistance . As reported in the literature , game tophytes were more resistant as age increased.

I n addition to the relationship be tween so2 conc entra­ tions and chlorophyll des truction, there appe ars to be a corre lation be tween atmospheric turnover in the fumigation chambe rs and chlorophyll destruction. Data, obtained in this research, indicate that an increase in the amount of so2 brought in contact with the mos ses brings about addi­ tional chlorophyll destruction . That is , des truc tion of chlorophyll is re lated both to so2 concentration and to the arr�unt of so2 to which mosses are exposed. This is a variable that ne eds to be considere d in any future research

iii so on 2 effects on plants .

The four moss spec ies investigated exhibit sufficient

so 2 sensitivity in the young gametophyte stages so that

their life cyc le could be easily interrupted by moderate

so levels of 2 pollution. This could lead to an eve ntual

extin so . ction of these species in an area polluted by 2

iv ACKNOWIEDGE�::ENTS

I wish to express thanks to my advisor , Dr . Terry M.

We idner for his ideas , enc ouragement , guidance, and

ass istance in the writing of this thesis.

Thanks are also due to Dr. Charles B. Arzeni and Dr .

Wesley C. Whiteside for the ir helpful criticisms and

sugges tions in reviewing this manus cript .

Spec ial appreciation is extended to Dr . William A.

We iler for his assis tance in devising the experime ntal

des ign for this res earch. Thanks are due to Dr . Roger L.

Darding for help in the des ign of the fumigation chamber.

Most importantly, I would like to express appreciation

to my wife , Cathy , for her support and understanding only

to face a further degree .

Acknowle dement is made to the Botany Department and

Tiffany Graduate Research Fund for the fund ing of this study .

lastly, I wish to thank my parents and friends for

the ir encouragement and support .

v TABI.E OF CONTENTS

Page

ACKNOWI.EDGEMENTS ...... v

LIST OF TABI.ES ...... vii

LI ST OF FIGURES ...... viii

INTRODUCTION ...... 1

LITERATURE REVIEW ...... 4

WlATERIALS AND METHODS 24

RESULTS AND DISCUSSION 36

APPENDIX ...... 43

I.ITERATURE CITED ...... 110

vi LIST OF TABLES

Table Page

I. Chlorophyll content of so2 fumigated mosses 44

so II . Moles 2 exposure for three flow rates and

so concentrations during fumigation ... 82 three 2

vii LIST 0.2 FIGURES

?igure Page

I. Degradation of chlorophylla to phaeophytina . 84

II. Illinois distribution of mosses used in fumigation ...... 86

III. Fumigation apparatus for exposing mosses to sulfur dioxide ...... 88

IV. Response of gametophytes of 4- and 8-month old Aulacomnium heterostichum to so2 rates toncentrations at different flow 90

a V. Respons of gamstophytes of 4- nd 8-month old Bartramia pomif'ormis to so2 concentrations at different flow rates 92

VI. Response of gametophytes of 4- and 8-month

old Leucobryum glaucum to so2 concentrations at different flow rates ...... 94

viii Figure Page

ametophytes of VII . Response of g 4- and 8-month old Polytrichum ohioense to concentrations so2

at diffe rent flow rates . • . . • ...... • . . . • . . 96

VIII. Responses of 4-month old game tophytes of various mosses to differnt concentrations so2

at 1.0 liter per minute ...... •...... • 98

IX. Responses of 4-month old gametophytes of

various mosses to different so2 concentrations

at 2.0 liters per minute ...... •.... 100

X. Responses of 4-month old game tophytes of

various mosses to different so2 conc entrations

at 4.o liters per minute ...... •... 102

XI. Responses of 8-month old game tophytes of

various mosses to different so2 conc entrations at 1.0 liter per minute ...... 104

XII. Responses of 8-month old gametophytes of

various mosses to different so2 conc entrations

at 2.0 liters per minute ...... 106

ix Figure Page

XIII. Responses of 8-month old game tophytes of

various mosses to different so2 concentrations at 4.0 liters per minute ...... 108

x INTRODU�TION

Although mosses have not been studied as extensively as lichens, the available research suggests that mosses are at least as sens itive to air pollutants as lichens.

Both field data and experimental data support the hypothesis that mosses are sens itive to air pollutants .

Arnold (J ) noted that mosses were disappearing from the city of Munich. Later studies (24, 42 , 69) demonstrated that mosses are almost completely absent from city centers and that normal moss communities are only found in rural areas remote from the city ce ters. I n

Field studies (25, 2 6, 53, 64 ) have provided a great deal of information about mosses responding to air

pollutants . However, the critical question is whether air

pollutants are present in suffic ient concentrations to cause the observed effects in the field . Available experimental data are for the most part insufficient to answer this question. Coker (14 ) observed a differential

1 2 response among moss species to sulfur dioxide. The

) concentrations of so2 used by Coker (14 were considerably higher than those found in urban areas. Coker's lowest so2

5.0 concentration of ppm for 44 hours exposure is higher than the highest 24 hour concentration (0 .9 ppm) reported for New York City between 1957 and 1958 (9) and higher than the highest 24 hour concentration (1 .J ppm) of so2 during the infamous London smog of 1952 (81 ) .

Nash (5 5) ran a careful experiment using realistic concentrations of so2 on mature field collected gametophytes (unknown age) and cultured {gametophytes, known age) of the of the protonemal stages in mosses. Several authors

(24, 42) noted that sexual reproduction is blocked among many species which occur in urban environments plagued by so2 pollution . Nash's (5 5) observations on the sensitivity of the protonemata are probably more important because reproduction in mosses involves a protonemal stage in the life cycle.

The study reported herein is designed to examine the response of the protonemata of several more common species of moss to more realistic concentrations of ?his so2. 3

should provide further information on the relative

so . sensitivity of different ages of mosses to 2 Another aspect to be considered in this investigation is whether the rate of atmospheric turnover in a fumigation ch�mber has a bearing on S02-inflicted damage . In other words, is it solely the so2 concentration that determines an organism's

so ' so sensitivity to 2 or is it the amount of 2 brought into contact with organism, or is it a combination of both factors? LITERATURE REVIEW

Sulfur dioxide constitutes about of the (so2) 95% sulfur compounds resulting from the burning of fossil fuels that contain sulfur. Sulfur dioxide is a colorless, nonflammable gas which is very soluble in water (2 2% by weight at 0 C and standard pressure ). At concentrations of ppm, produces an acrid taste whereas, at 0.3-1.0 so2 levels of 3.0 ppm or greater it has a pungent odor which can cause pulmonary problems (11).

Sulfur dioxide is reduced to by bacterial action, H2S and in air it is further oxidized to The rate at so3. which this oxidation takes place depends on the presence of ultraviolet radiation and other reactions or catalysts

(37 ) . For an overview of the sulfur cycle refer to

Gatchel (21) and Kellog et al ( 37 ).

Sulfur is adsorbed by plants mainly in the form of sulfate ions and is reduced and incorporated into organic compounds (77) . It is a constituent of the amino acids

4 5

cysteine and methionine, and the proteins containing these amino acids. Thiamine, biotin, and coenzyme A are sulfur­ containing, low molecular weight coenzymes. These are essential in metabolism when attached to appropriate apoenzymes which require these enzymes for catalytic activity. The ferredoxins, non-heme iron proteins involved in photosynthesis and other electron transfers, contain sulfur in amounts equivalent to the iron present (29).

Volatile c·ompounds containing sulfur contribute to the characteristic odors given off by onions, mustards, and other plants. The functions of these compounds in the plant is not understood.

Reduced sulfur can be regarded as an oxygen substitute which extends the capacity of organic compounds to complex metals to enter into redox reactions and group transfers.

Protein structure is stabilized by disulfide linkages which are under redox rather than hydrolytic control (lJ).

Oxidized sulfur provides a stable acidic group to organic molecules, enhancing their water solubility (lJ) .

In healthy leaves sulfur content ranges from 500 to

14,ooo ppm by dry weight (0.5-14 mg/gr dry wt. ) depending 6

upon the species (44) . Concentrations below 250 ppm are considered critical , giving rise to deficiency symptoms

( and to the substitution of selenium when available ) for sulfur in amino acids and proteins (?8).

There are specific macroscopic symptoms in sulfur deficient plants . Deficiency symptoms include chlorosis of the youngest leaves and a spindly appearance of the plants .

Leaves can readily absorb so . 2 Part of all of the sulfur requirements of plants may be met by direct uptake

so so of 2 from the atmosphere if 2 is present at very low concentrations . In green_houses at the University of

�aryland, striking sulfur deficiency symptoms have often been produced but when_ atmospheric conditions cause smoke from a nearby power plant to settle in the greenhouse, symptoms disappear in 24 hours or less ( 22).

The sulfur requirements of plants may be met by the uptake of sulfate ions through the roots . If sulfur is present at very low concentrations the sulfur requirement may be completely fulfilled by the direct uptake of so 2 from the atmosphere . so If 2 concentrations exceed the 7

plant's biochemical threshold levels, the fundamental cellular processes such as respiration and photosynthesis

may be disrupted. Irreversible injury may occur if high concentrations persist. Tolerance varies with species and the various environmental factors.

Sulfur dioxide toxicity to plants can be related to its chemistry. may be a reducing or oxidizing agent so2 depending upon the pH of the medium in which it exists.

Puckett et al. (62) have pointed out that at physiological

pH values, dissolved can participate potentially in so2 both oxidation and reduction processes. In water at a pH of 7.2 (normal for the cytoplasm of most plants), sulfur exists approximately in the form of and in the 50% so; 50% form of HSO (16). At pH 1.8, and exist in the j H2so3 Hso3 ratio of 1:1. In general, an acidic solution favors the formation of , while an alkaline solution favors the H2so3 formation of may also be oxidized to so;. so2 so4 (44).

is extremely soluble in water; 22 by weight at so2 %

• • and at The concentrations normally found in 0 C 9.4% 24 C.

polluted areas dissolve completely upon contact with surface

or tissue moisture of plants. In so establishes lution, so 2 8

the following equilibria, which have an important bearing on its effects.

so + 0 ...... 2 H2 H2so3 + = H so ...... H + Hso , pK 2 3 3 1.76 + Hso- � H + so , pK = 3 3 7.20

so2 in jury on plants has received much attention in the last thirty years and symptoms of acute injury are well known. Acute necrosis results from rapid absorption of so . 2 When lethal concentrations accumulate in the most susceptible areas of the leaf, a dark green, water soaked discoloration develops. The affected area soon becomes flaccid, and upon drying, becomes white to ivory in most plants . In some cases the dead tissue may turn red, brown, or almost black (46, 52).

Acute so2 injury on broad-leaved plants may develop marginal or intracostal necrotic areas. Tissues along the ma jor veins are not affected, making the veins stand out clearly on the ivory-colored, necrotic background.

Intercostal areas collapse and dry out, leaving regions 9 that are ivory-�olored or, in some plants, irregularly­

shaped, dark brown necrotic areas. Organic sulfates

migrate to the leaf margins and produce marginal necrosis

which may extend between the major veins toward the midrib.

Bleached or chlorotic tissue may develop between the necrotic lesion and healthy appearing tissue.

In pine needles, in jury usually starts at the tip so2 and extends toward the base as successive exposures produce more severe in jury. Young needles are more susceptible to in jury than older needles. Acute in jury appears as apical, medial, or basal bands of orange-red tissue on the needles of the current year. Older needles exhibit a yellow-green color which later turns reddish brown. Needles will abscise in 1-2 years instead of the normal 3-5 years.

Both sulfur dioxide and sulfite ions can be utilized by plants in normal sulfur metabolism. It is the amount of these ions in excess of "normal" quantities that is in jurious to plant metabolism. Plant cells possess a buffering capacity capable of neutralizing some of the acidity caused by disassociation of (75, 80). However, H2so3 if sulfur in excess of plant requirement enters, no matter 10

how slowly, chronic in jury characterized by general chlorosis develops. If sulfur enters too rapidly, the plants' metabolic systems for coping with sulfur are overwhelmed and acute in jury occurs.

Most biochemical reactions take place within membrane­ enclosed structures. ?or any pollutant to affect metabolism, it must be capable of entering these structures in its active form (s ). Cell membranes have selective permeability as a result of their orientation, composition, ionic nature, and other chemical properties. On the basis of ionic strength, the membranes will be more permeable to and so2 than to the more highly charged ions. Following H2so3 soj the entry of into the cell, it is assumed that so2 H2so3 and are disassociated to cause acidification . Hso3 . The interference with the structure and permeabil­ S02 ity of cellular membranes and enzyme activity will affect many biochemical processes (66, 82). Photosynthetic pigments and many enzymes are associated with the membranes of chloroplasts. These membranes are very fine structures and can be easily disrupted by Aqueous is S02 (82) . so2 capable of interfering with the electron flow in the 11 electron transport chains of chloroplast systems I and II.

It was demonstrated that bleaching of extracted pigments

by was identical to that induced by potassium so2 permanganate, a strong oxidizing agent . The same bleaching process has been observed for air-oxidation of chlorophyll in acetone ( 36). Sulfur dioxide is also capable of inactivating many enzyme systems by splitting their disulfide link.ages (6).

Asada et al . (5) found that bisulfite compounds reduced fixation in spinach chloroplats . It was C02 suggested that bisulfite compounds probably inhibit co2

fixation by interfering with chloroplast membranes or by affecting transport systems associated with chloroplast membranes in photosynthesis (51).

Photosynthesis is one of the first processes to be affected by Ziegler (84) found that the inhibition so2. of fixation by for active sites on the enzyme co2 so3 ribulose 1, 5-diphosphate carboxylase . High concentrations resulted in non-competitive inhibition of the enzyme .

Osmond and Avadhani (58) found that bisulfite compounds are effective inhibitors of the phosphoenol pyruvate 12

. carboxylation system in c4-dicarboxylic acid pathway of photosynthesis .

The effects of so on the distribution of lichens has 2 been well researched (J0,40). The photosynthetic algal component of the lichen seems to be the most susceptible

so . (1 4, 23 , 25 , JJ , 54, 62 , to damage from 2 Studies 59,

63) have shown that the exposure of lichens to so2 resulted ++ in the breakdown of chlorophyll into phaeophytin and Mg ions . Similar results using bryophytes were obtained by

Coker (14), and Nash and Nash (55) . Liverworts were also

used in one study ( 21) and yielded the same results.

When treated with a weak acid, the chlorophyll

++ molecules can be converted to phaeophytins. The Mg ion is replaced by 2 atoms of hydrogen thus changing the spectral properties . Arndt (2) found that this could be acheived by

so hydrofluoric and hydrochloric acids and thus is not 2 specific. Arndt ( 2) also found that the a-carotene content was affected by so . 2

Hill (JJ) proposed that the breakdown of chlorophyll was a secondary effect of so and had little relevance to 2 the disappearance of plants from a polluted area. lJ

Showman (68 ) supported Hill by observing a decrease in net

photosynthesis of fumigate lichens in the absence of any

detectable chlorophyll breakdown . Griessmeyer (28)

reported that so binds irreversibly with the "chloroplast 2 iron" found in ferredoxin and cytochromes. He suggested

that the inhibitory effects of on photosynthesis may so2

be due to inactivation of these electron carriers.

Wellburn et al. (82) reported a reversible swelling of the thylakoid within the chloroplasts when broad bean plants were fumigated with S0 (. 25 ppm, 2 hours . Thylakoids 2 ) contain some of the dark reaction enzymes in their lumen and have photophosphorylation particles on their inter- connections between membranes and any disruption to these structures will affect so assimilation (J4). �alhotra 2 and Docking (44) showed that at concentrations of 10-100 ppm, aqueous had no effect on the concentration of so2 chlorophyll or chlorophyll or phaeophytin. In contrast, a b concentrations of 250-500 ppm, phaeophytin increased in . a concentration, but not phaeophytinb, which suggests a conversion of chlorophylla only.

Malhotra (44) suggested that low concentrations of aqueous so2 , which normally do not produce visual symptoms on vegetation, will result in in jury at the molecular

level after prolonged exposure . The affects enzyme S02 systems such as chlorophyllase. Higher concentrations may

. . . ++ cause conversion of chlorophyll into phaeophytin and Mg , and the loss of photosynthetic activity through competition between and for the active site on ribulose 1,5- so2 co2 diphosphate carboxylase (84).

Effects on General Metabolism

The formation of a-hydroxysulfonates in plants exposed to inhibits glycolic acid oxidase which is necessary in so2 the glycolic acid pathway (4, Suppression this 32). of pathway will adversely effect glycine and serine synthesis .

Tanaka et al . (72) reported that photosynthetic formation

of serine was reduced by .50% in plants exposed to s • o2 and are both capable of converting disulfide so2 H2SOJ enzymes or p�oteins to thiosulfonates and thiols. The S -S bonds in the poly�eptide chains are cleaved by (6, 2 H2so3 1 ).

By disrupting S-S bonds in methionine and cysteine, the structural integrity of proteins is also disrupted .

Sulfite, being nucleophilic in nature, can combine with 15

4 NAD to form hydropyridine- -sulfonic acid, which is capable of forming an undissociable complex with dehydrogenases such as lactate and malate dehydrogenase. Sulfite thereby inhibits the hydrogen transfer system (6 0, 61).

Sulfite can react with free radicals that can (49) lead to splitting of phosphodiester linkages of DNA (31).

at some of the toxicity caused It seems possible, then, th by S0 may be due to inactivation of DNA or of messenger 2 S02 RNA (6 6).

(4 ) Mansfield and Ka jernik 5 reported that so2 is capable of increasing the stomatal aperture openings.

Studies (48, 83) have shown that bisulfite compounds also affect the stomatal openings in a way that is similar in appearance to the effect caused by The increased co2 (48). opening of the stomatal complex has two disadvantages to normal metabolism. First, the increased stomatal opening enables so to penetrate more readily into the more delicate 2 o:f inner structures the leaf and disrupt cellular metabolism.

Second, wider opening of stomata results in a rapid rate of water loss. In jury symptoms due to S02 closely resembles

o those caused by excessive water loss or a high rate f transpiration (J 5). Thus, at least part of the toxicity so�c:.

may be due to water stress.

so Effects on Epiphytic �ryptogams 2

In the early nineteenth century, botanists became aware

that lichens and bryophytes were scarce near large urban

centers and started to recognize that air pollution aris-

ing from these urban centers could adversely effect the ecesis and development of these plants. The declining lichen vegetation in the outskirts of the Jardins du

Luxemburg in Faris was noted by Nylander in 1866. He wrote

, ' � "Les lichens donnent a leur maniere la mesure de salulrite

, ' ' , . hygiometre tris sensible" de 1 air et constituent une sorte d

( 57) .

Later, the effects of air pollution on vegetation was

noted by many other botanists in Europe. N. J. Winch (2 5)

compiled county floras in 1807 and 1831 for lichens and

bryophytes grov.;ing near Newcastle ,England in the early

years of the industrial revolution. Winch ( 25) recorded

a large number of species growing within 5 km of the city

center and made no mention of damage to the vegetation.

Hebarium specimens are on file to testify to the luxuriant 17

growth of the lichens and bryophytes at that time. In 1879

Rev. W. Johnson (25 ) could find no trace of foliose or fruticose lichens in the woods 8 km west of the city.

Crombie in 1885 observed a reduction in lichens in the

Epping Forest near London and even observed their extinction elsewhere in England as a result of extensive urbanization and resulting increased air pollution (JO). In 1892,

Arnold (J) noted the same reduction in epiphytic cryptogams in the city of Munich.

The story regarding bryophytes is less clear due to the scarcity of records. Winch (25 ) does mention several communities of bryophytes that occurred near Newcastle,

England in the early 19th century and these appear to have been typical. Even though the habitats remain little altered at the present time, most of the species have disappeared. Though data is incomplete, county floras seem

to indicate that, in the :)rd !quarter ·Of the nineteenth century, bryophytes were fast disappearing around urban areas (23 ). Other studies noting this phenomena were published on New York (10 ), Stockholm (69 ) , Montreal (18 ), and Montana (67 ). 18

The cause or causes of epiphytic deserts have yet to be resolved . Is it dryness or air pollution which causes the impoverishment of lichens in cities-and industrial areas? Some (8, 38, 47) believe that it is the drought

( drought hypothesis) that contributes to the epiphytic deserts . Most workers believe that it is air pollution

( pollution hypothesis) and some (7, 10, 20) believe that neither air pollution nor aridity alone are responsible; rather both together contribute to the paucity of lichens and bryophytes in cities .

Other factors may also contribute to the declining populations of lichens and bryophytes in the cities.

Altered temperature and humidity may be responsible.

Microclimates in urban areas tend to have higher tempera­ tures and reduced relative humidity in comparison to rural microclimates (24). It may be due to the reduced light intensity in urban areas which is a result of particulate matter . The absence or decline of plant species may be due to pollutant influences upon the reproductive phases.

Both Gilbert (24) and Nash III (55) reported a greater sensitivity to pollution in the protonemal stage than in 19

mature gametophytes. Gilbert (24) believed that the deserts were from species sterilization in the city centers while Nash III (55) believed that it was blockage in moss reproduction, namely in the protonema stage. The actual cause may be a combination of· the above factors and influences from the microenvironment.

Gilbert (25) compared 16 aspects of the ecology of the cryptogams and concluded that, despite significant differences in their morphology, bryophytes and lichens are similar in their response to air pollution. This suggests that it is possible for them to colonize inhospitable habitats only by adaptations which carry with them an inherent susceptibility to so . It has been widely assumed 2 that the highly efficient mechanisms which lichens and bryophytes possess for accumulating a wide range of minerals from dilute solution (27, 70) are at least partly responsible for their acute sensitivity to air pollution. They are c:omparitively slow growing and relatively long-lived and can, therefore, accumulate large quantities of toxic materials over a long period of time, whereas other plant types usually shed large areas of toxic-laden parts each 20

season (1 0). Gilbert (23) did sulfur determinations in the lichens growing in polluted atmospheres. He found that sulfur concentrations in the lichens were thousands of times greater than those of the air in which they were growing but the concentrations were proportional to the amount of local so pollution. 2

71) Research (7, indicates that bryophytes have greater

sensitivity than do vascular plants because (1) they so2 lack an impermeable cuticle and regulatory stomata, (2 ) they absorb rain water directly over their entire surface, in contrast vascular plants obtain most of their water indir­ ectly after it is filtered through the soil and has under­ gone soil water interactions, and (3) bryophytes are most active in the seasons of higher humidity: spring, fall, and winter . Studies (9) have shown that pollution levels are the highest in New York C::ity between November and March.

This is due to lower cloud formation, frequent temperature inversions, fogs, and extensive residence heating (9) .

Fumigation and thalli transplant studies (14, 25, 41, 4J,

54, 69) have shown that mosses and lichens exhibit similar morphological and physiological responses to sulfur dioxide 21

pollution.

Turk and Wirth (79) noted a decrease in photosynthesis J in mosses after exposure to 5.25 ppm so2/m air, suggesting

that mosses are more sensitive to so2 pollution than lichens.

Rao and LeBlanc (6J) found that Xanthoria algal cells,

5 exposure for after ppm so2 24 hours, exhibited bleached chlorophyll, brown spots on the chloroplasts, a permanent ++ plasmolysis of the cells, and an abnormal Mg ion and

phaeophytin content.

Several authors (25, 69) and others have rnairtained that sulfur dioxide is the principle factor responsible

for the disappearance of lichens and bryophytes from

polluted areas. However, the crucial question in assessing the validity of this hypothesis is whether is present so2 in sufficient concentration in these polluted areas to account for the disappearance of these plants. Examples confirming this hypotheis are a study (59) demonstrating a reduction in photosynthesis in Parmelia sulcata after

concentrations of 100-10, 000 ppm for exposure to so2 24 hours . Rao and LeBlanc (6J) observed degradation of chlorophylls a and b in four lichen species after fumigation 22

5.0 so 24 of ppm 2 for hours.

so The probability of 2 occuring in polluted areas at concentrations as high as 5.0 ppm for as long as 24 hours is rather remote. ?or instance, over a 12 year period 1957-

1968, the highest hourly average was 2-J ppm and the highest 24 ho.l!.1.:r average was 0.9 ppm in New York c::ity (9),

The highest concentration of so recorded for a hour 2 24 period was 1.J ppm during the infamous London smog of

1952 (81).

so More relevant studies have shown 2 sensitivity at concentrations that are likely to be found in the polluted areas. Nash III and Nash (55) noted mature moss gameophytes

so 2-4 were 2 resistant at concentrations of ppm for 8 hours while moss protonema were susceptible to damage at concentrations of 0.2 ppm for 8 hours. Taoda (74) noted in jury to bryophytes at various concentrations and time periods. The greatest damage recorded at 0.8 ppm for

10-40 hours and poor growth at 0.2 ppm for greater than

100 hours exposure.

Nash (54) established a short term fumigation susceptibility threshold for lichens at about 0. 5 ppm. 23

Rao and LeBlanc (63) found lichen and bryophyte chlorophyll to be sensitive at concentrations in excess of .154 ppm, so2 with long term levels of less than .002 ppm causing no injury, 0.006-0.03 ppm causing chronic injury, and greater than 0. 03 ppm causing acute in jury.

Gilbert (25) analyzed 16 aspects of the biology of the cryptogams to try to establish the effects of the microenvironments in buffering or increasing susceptibility to damage. Gilbert (25) noted several controlling so2 influences of the habitat, i.e. , substrate pH, buffering capac ity, nutrient flushing, shelter, and humidity. He proposed that a high pH reduces the toxic ity of the pollutant directly by ionizing the sulfurous aciu.,• ,.::i or indirectly as an exclusion agent of the habitat.

Sensitivity to pollution varies from species to so2 species. Numerous factors figure prominently in the sensitivity levels of the plants. The complex biogeochemical reactions of plus variou environmental so2 � factors must be considered in assaying lichen and bryophyte tolerance to so2. MATERIALS AND Iv'.:ETHODS

The four bryophytes listed below were selected as test organism because of their morphology: they have a large prominent gametophyte generation, they are common inhabitants of deciduous forests, and are widely distributed

in eastern United States and in many other countries. In addit�on, Polytrichum ohioense and Leucobryum glaucum had been used in earlier experiments by Nash (55).

A. Aulacomnium heterostichum (Hedw. ) B. S. G.

Description according to Crum (17). Plants in rather loose, green or sometimes yellow-brown tufts 1.5-4 cm high.

Leaves rather soft, concave and slightly oblong or oblong­ ovate, not decurrent, subacute or obtuse and apiculate; margins plane or narrowly recurved below the middle, coarsely serrate in the upper 1/2 or more; costa not flexuose, ending somewhat below the apex; upper cells sm�ll, green, smooth, thick-wall, irregularly rounded-quadrate, rounded-hexagonal, and transversely short-oblong, 9-1 1, often 15f wide; basal

24 25

cells similar, those at the insertion somewhat longer, not colored or swollen. Autoicous; perigonia small, axillary.

Setae 8-17 mm long; capsules 2.5-J.5 mm long, =uberect or inclined, somewhat curved; operculum oblic;_uely ::..�ostrate; endostome segments not perforate. Spores very finely 9-15f, papillose.

Aulacomnium heterostichum is quite common in eastern

North America . It ranges from Ontario south to �lorida and

Texas and west to Minnesota and Kansas. It has been

(47) ) reported from JO counties in Illinois ( ?igure II .

This moss is typically found in moist to dry woods on rich soil in the shade at the bases of trees. =:rum (17) reports that this moss is often associated with Bartramia pomi:formis.

2. Bartramia pomiformis Hedw. Description according to =:rum (1 7) . Plants in rather dense, soft tufts 2-6 cm high, light- or yellow-green or yellow-brown, sometimes � glacous, tomentose below. Leaves � spreading and flexuose or crisped from an erect base when dry, loosely erect or erect -spreading when moist, 4-5.5 mm long, linear-lanceolate, acuminate; margins r2volute almost to the apex, bistratose and doubly serrate; costa shortly excurrent, serrate at 26

back toward the apex; upper cells subquadrate and short­ rectangillar, about 1-2:1, thick-walled, unipapillose on both surfaces; basal cells pale, long-rectangular, rather thin-walled. Synoicous and autoicous (the perigonia next to the perichaetia ). Setae 8-20 mm long, erect-flexuose; capsules 1.5-2.0 mm long, inclined to horizontal, globose to ovoid, deeply furrowed when dry; annulus none; operculum convex; peristome teeth inserted somewhat below the mouth, lanceolate, red-brown, very finely papillose, strongly barred at back; endostome pale-brown, shorter than the teeth, consisting of a well-developed basal membrane and broad segments. Spores spherical and � ellipsoidal, brown, coarsely papillose, 20-24)'; n=8, 8+1; 2n=16.

This moss is commonly known as the "Apple Moss" because the capsules resemble miniature apples. It is a fairly common moss in eastern North America and is also found from Alaska to Oregon and Montana. It is also found in Europe, Asia, New Zealand, Greenland, and Canada. In

2) . Illinois it has been reported in 25 counties (47) ( Figure

This moss is found in the moist, shaded soils typical of the deciduous forests of eastern North America. It is 27

occasionally found growing on rock in crevices or on ledges.

3. Leucobryum glaucum (Hedw.) Angtr. ex Fr.

Description according to Crum (17) as follows: fairly

robust plants in deep, hemispherical cushions usually 2-9

cm high (but often much higher and forming very large

hummocks), sometimes bearing clusters of reduced leaves

at the stem tips. Leaves erect to erect-spreading or

sometimes subsecund, 3-8 mm long, lanceolate and concave

or subtubulose from an erect, oblong-obovate base 1.5-3 mm

long, acute or apiculate, usually� serrulate at the tip; costa in cross-section near the insertion showing 5-7

irregular layers of leucocysts on either side of the middle, with 3-4 layers below and 2-3 above the chlorocysts; lamina

5-11 cells wide. Setas 9-17 mm long, red-brown; capsules

1. 5-2 mm long, curved and strongly inclined, strumose at base; annulus none; operculum 1. 5-2 mm long. Spores 13-18f ,

nearly smooth or very finely papillose. Calyptrae often

not split completely to the base which clasps the tip of

+ 14 . the seta until maturity of the capsule.-n= 11, 11,

This moss is commonly known as the "White Moss " or

"Pincushion Moss" in reference to its color or its grov-.'th 28

habit of forming dense tufts that resemble pincushions.

It is found in Europe, the Azores, the Canary Islands,

Japan, the Caucasus, �adeira, and in North America from

Newfoundland to Minnesota south to Louisiana and ?lorida.

In Illinois it is reported from 24 counties (47) ( ?igure 2).

It prefers soil or humus in moist to dry forests and is often found in association with decomposed wood humus of decaying logs or stumps. Leucobryum glaucum often forms large mats on the ridge tops in oak-hickory forests. This plant is unusual in that it rarely produces sporophytes but instead reproduces asexually by producing caduceus leaves which can drop off to give rise to new plants.

::::ard. Description 4. Polytrichum ohioense Ren. & according to Crum ( 1 7) as follows: moderately robust plants in loose, dark-green to brown tufts 1.5-6 cm high.

Leaves erect or erect-spreading when dry, wide-spreading and recurved when moist, 6-10. 5 mm long, long-lanceolate from a sheathing base about 2 mm long, acuminate, ending in a short, reddish, toothed awn; margins erect and coarsley 29

toothed nearly to the shoulders; lamellae 30-41, covering almost the width of the leaf above the shoulders, entire or nearly so in profile, 3-7 cells high, the marginal cells in section somewhat broader, flat-topped or somewhat retuse, with thick, brown, smooth walls; costa r�ther shortly excurrent, sparsely toothed at back near the apex.

Dioicous or rarely autoicous; perichaetial leaves not much differentiated. Setae 15-85 mm long; capsules 2-5 mm long, pale-brown, suberect to strongly inclined, not or only slightly constricted between urn and apophysis, 4- or rarely 5-angled; annulus none; exothecial cells not bulging, without a central thin spot; stomat numerous (but difficult to observe), with 2 guard cells. Spores 11-13f,

-n=14.

This moss is commonly known as the "Haircap Moss" or the "Pigeonwheat Moss" in reference to the hairy calyptra.

Polytrichum ohioense is circumpolar in distribution. In

North America, it is common in the eastern United States, but it has also been reported in New Mexico. In Illinois it is reported from 17 counties (47) (Figure 2). This moss is found in soil or humus in mesic woodlands, but is 30

also occasionally found in old fields.

Specimens of live mosses were collected at the following sites:

T9N, A. Rocky Branch Nature Preserve ( Clark ::;o. ,

R12W, sec. 29).

B. Rocky Hollow ( Clark Co. , T9N, R12W, sec. 5).

Specimens were surface-sterilized by immersion in

(5.25% ::;hlorox bleach sodium hypochlorite ) for 5 minutes.

Then the bleach was rinsed off with 3 rinses of sterilized, deionized water. Immediately after rinsing, sections of the stem and leaves were placed on the surface of a growth medium to obtain a generation of protonemata. Knop's medium (50), 1/3 strength, pH 6.91, was used to supply minerals to the mosses. The minerals were dissolved in a water solution to which 0.35% agar was added. The agar eatablished a semisolid surface from which the moss protonema could be later washed free. The young protonemata were culture in sterile plastic petri dishes and were later sub-cultured to sterile, polypropylene Tri-Pour graduated

250 ml beakers to which approximately 150 ml of medium had been added. 31

Saran Wrap was placed over the mouth of the Tri-Pour beakers and was secured by rubber bands. The Saran Wrap provides a barrier to contamination while allowing a gaseous exchange between the outside atmosphere and container atmosphere as indicated by the following d t a a: / / / Oxygen Transmission Rate: 1.5 cc 100 in2 24 hr

0 atm at 75 F.

Moisture Vapor Transmission Rate: .45 g/100 in2/24 ° hr/ atm at 100 F.

C0 Transmission Rate: 9.98 cc/100 sq / mil 2 in ° thickness/24 hr/atm at 25 c.

Information regarding Saran Wrap permeability to gases was provided for by Charles L. Mott (50 A).

Mosses were cultured at + 1� 8 with 28°8 a 16:8 photoperiod. Most of the time the temperature was a nearly constant 28°C with only occasional flunctuations.

Illumination was provided by banks of fluorescent lights.

4 Each bank contained a cool white fluorescent light ( 0 watts) and a Vita Lite Power Twist fluorescent light

(40 watts) suspended 43 cm over the mosses. Light intensity at the level of the mosses w s lux 10 lux measured a 330 � as 32

by a Li-Cor, LI-185A Quantum/Radiometer/Photometer. The

translucent containers were placed on Kadry, a plastic­ backed white bench paper, to achieve a uniform background.

Mosses were arbritarily chosen as being one day old when the cultures of protonemata were in colonies of 1 cm

diameter. Culturing times ranged from 2 weeks to 5 weeks to acheive this size in the various mosses. Polytrichum ohioense and Aulacomnium heterostichum were the fastest growers and Leucobryum glaucum was the slowest. Apparently, species differed in ability to recover from the surface sterilization. The moss gametophyte cultures were composed of bulbils, gametophytes, and protonemata in varying ratios.

The fumigation apparatus is illustrated in ?igure III.

All connections were made with tygon tubing. Air pumped

thru the system was filtered initially through a Drierite filter. The air then went thru a Series 10A1460-LK Low

Flow Flowrator to regulate and measure the air flowrate.

Next, the air was passed thru a water trap to humidify the air. Sulfur dioxide was provided from a cylinder of

Matheson Certified Standard, 50 ppm in air. The S02 JJ

flowrate was regulated and ad justed with a Series 10A1460-

LK Low Flow Flowrator to the desired flowrate. The air and so 2 were mixed and then introduced into the exposure

chamber. The exposure chamber was a 7-liter glass bell

jar. It was situated on a Mc�ord Rubberized g�sket which, in turn, was placed on J/4 inch plywood. To achieve an airtight seal, the bell jar was Vaselined on the bottom and clamped onto the gasket and plywood. The so2 gas mixture was vented into the atmosphere.

The gas flowrates used during the experiments were 1

2 liter per minute, liters per minute, and 4 liters per minute. This rate resulted in an atmospheric turnover in the exposure chamber of, respectively, once every 7 minutes, once every J. 5 minutes, and once every 1.75 minutes. For each flowrate, J different so concentrations were used; 2

0.2, 1.0, 4.o ppm. These flowrates and concentrations resulted in 9 different fumigation treatments.

For fumigation the mosses were transferred to open the plas�ic petri dishes and then placed into fumigation

.2109 chamber. Sample tissue weights varied from .0791 to for each moss fumigated. The mosses were fumigated with 34

so2 for 8 hours and then so2-free humidified air was run

thru the chamber at 1 liter per minute for 12 hours to

remove any residual so2. During the fumigation period,

+ 2 (8 hours 1 hours ) the light intensity was maintained

at 330 lux with a 250 watt incandescent light bulb.

+ 3 � Temperature within the exposure chamber was 27•:

Each fumigation was replicated 3 times.

Chlorophyll analysis was performed within 24 hours

after the end of the fumigation period. Mosses were covered

with plastic petri dish covers and placed back under normal

germination conditions if not analyzed immediately after

fumigation.

The protocol for chlorophyll analyses of the treated mosses was as follows. The moss protonema was washed from the growth medium by placing the moss-agar combination on a

fine wire mesh screen and by rinsing first with tapwater

and then deionized water. This washed away the agar and

left moss behind. Then the mosses were blotted dry on

paper towels and weighted immediately on a Sartious

analytical balance. Extraction of the chlorophyll involved

maceration of the weighed plant tissue in several ml of 80% J .5

acetone for 5 minutes with a mortar and pestle. A few drops of 1M Caco3 were added to prevent plant acids from destroying the chlorophyll. The brei was suction-filtered on Whatman #1 filter paper into a glass Buckner funnel.

The filtrate was made up to 50 ml with 80% acetone.

::hlorophyll analysis was performed using a Bausch &

Lomb Spectronic 20 spectrophotometer. Calculations of extracted chlorophyll were based on the equations of

Comar and Zscheile (1 5). RESULTS AND DISCUSSION

Chlor osis of the moss pro tenema was a general morphological response of most of the fumiga ted culture s of mosses. The disc o loration was espec ially prevalent in the young protonema • In the higher S02 conc entrations , even the older game tophytes were affected. Generally the chlorosis be came visible in the latter part of the 8 hour

S02 fumigation period . However, the discoloration be c ame more obvious during the 12 hour exposure to so2 -free , humidified air which immediately fol lowe d the so fumigation 2 period . Although not quantified in this exper ime nt ,

apparently there is some time lag for so - cau ed chlorosis 2 s to appear in mosses.

The data from fumigations of the game tophytes (Ta ble I) , indicates that there is a species and age differential susceptibility to S02 injury . Reference to Figure s IV-

XIII illustrates the differing rates of injury by so2.

Le uc obryum glaucum is the most susceptible and

36 37

Bartramia pomiformis sec ond in degree of so injury as 2 measured by chlorophyll destruction. Aulacomnium heterostichum and Polytrichum ohioense are more resistant to S02 damage than Bartramia �omiformis and Le uc obryum glaucum at low so2 concentrat ions . 1his is consistent with data reported by Nash (55) , who found that Polytrichum ohioense is a S02 -tolerant species while Le uc obryum glaucum is a so -sensitive species. No research has be en done on 2

Bartramia pomiformis and Aulacomnium heterstichum protonemata with regard to so2 injury .

Data from Table I indicates that there i.s a decreasing sensitivity to so damage with an incre ase in the age of 2 the moss gametophytes. With the exception of a few scattere d cultures, the two-month old game tophyte s were killed at every so concentrat ion. In the four-month old 2 game tophytes, total chlorophyll destruc tior. was considerably

the 0.2 concentrations . Aulacomnium less in and 1.0 ppm so2 heterostichum and Polytrichum ohioense showed a total chlorophyll de struction of approximately 2 0 to 40%. Those of Bar tramia pomiformis and Le uco bryum glaucum (values) varied from about 40 to 9 0% for chlorophyll destruc tion. JB

The 8-month mo ss game tophytes demons trated a gre ater toleranc e to S0 . The two S0 -toleran 2 2 t species (Aulacomnium heterostichum, Bartramia pomif ormis ) generally exhibited less than a 20 des truction of total chlo % rophylls in so 2 concentrations of 0.2 and 1.0 ppm . ;Le ucobryum glaucum , the most so -sensitive moss , expressed 2 a 50 to 60% reduction in total chlorophylls at 0.2 and 1.0 ppm . Bartramia pomiformis showed a reduction of total chlorophylls of

35 approximately to 50% at 0.2 and 1.0 ppm .

Generally , all mosses fumiga ted were quite susceptible to chlorophyll destruction regardless of age when so 2 conc e ntrations were 5.0 ppm . Nash (55 ) found 1�-month old cultures of Polytrichum ohioense to be sensitive to fumigations with 4.0 ppm . However, Nash (55) also found that mature , field-collected game tophytes of Polytrichum ohioense , Dicrane lla heteromalla , and Pottia truncata were insensitive to sulfur dioxide at 4.0 ppm . Data from

Table I plus that of Nash (55 ) imply that the age of a moss is very important when determining a moss's potential for susceptibility to so damage . 2

One que stion that needs to be asked is whether the 3 9

flowrate in fumigation chambe rs is an important variable to

be cons idered. Nash (55) used a flowrate that re sulted in

an atmosphel�ic turnover of onc e C'-.-:-ery 6 minutes in the

fumigation chamber. This re search inc luded using different

flow rates to allow a comparison of the ir effect on

chlorophyll destruction. ?low rates used resulted in a

turnover of atmospheres of onc e every 7 minutes, once every

J.5 minutes, and once every 1.75 minutes.

Figure s IV-VII indicate that there is a corre lat ion

be tween atmospheric turnover and chlorophyll destruction.

This suggests that the incre ase in the amount of brought so2 in contact with the mosses brings about additional

chlorophyll destruction. That is , des truction of

chlorophyll is re lated to so2 conce ntrat ion and to the

so total amount of 2 to which mosse s are exposed. Table II

illustrates how the flow rate incre ase also leads to an

so , therefore , increase in the amount of 2 exposure and could

result in an increase in chlorophyll destruction.

4-month old at 1 liter Aulac omnium heterostichum ( ) . 2 ppm ,

per minute flow rate , average s a 2 9% chlorophyll de struction;

while the same concentration at 4 liters per minute 4 0

increases the average chlorophyll destruction up to 33% .

Leuc o bryum glaucum (8-month old), at 1 liter pe r minute

flow rate , 1.0 ppm so2 , has a average chlorophyll destruction

53% 61% of contraste d to a chlorophyll destruction at 1.0

ppm S0 at 2 liters per minute . However, as indicated by 2

the data .in Figure s IV-VII, there is not always an increase

of chlorophyll des truction with an increase in flow rate .

None thele ss, there seems to be an overall trend towards an

increase of chlorophyll destruction with the incre ase in

flow rate . I nve stigators should therefore allow for the se

a two vari bles, that is the so2 concentration and f low rate .

Pre vious studies ( 14 , 55, 63 , 71 ) indicate that

chlorophyll ' when subjected to a weak ac id or a re duc ing a

pol lutant such as so , is degraded to phaeophytin and free 2 a ++ Mg ions (Figure I). Table I denotes the different rates

of des truction in total chlorophyll ana lysis compared to

destruction of chlorophyll only . Chlorophyll is more a a

susceptible to so -injury than chlorophyll a s indicated 2 b

by chlorophyll analysis data reported in Table I.

The results obtained using cultured game tophytes

clearly establishes a greater sensitivity of the protonema 41

and young game tophytes relative to that of older

game tophytes. These re sults , toge ther with earlier studies

(24, 55) imply that the absence of a moss species from the

environs of a so2 source may be produc ed by a blocking of

the protonemata into mature game tophytes rather than by a

direct effect on the mature game tophytes. I n areas with

so interme diate 2 levels, which are toxic to protonema but whi ch do not affect the mature game tophyte , the species will gradua lly disappear as the old game tophytes die and the

species is unable to reproduce .

I t has been reported ( 24, 42 ) that sexual reproduction

is blocked among many species of mos s es which do occur in an urban environment . However, research indicate s that the

so sensitivity of the pro tonema to 2 pollution is probably more important than the blockage of sexual reproduc tion be cause both sexual and asexual reproduction in mosses

involves a protonematal stage .

The data ( Table I) demonstrates that the 4 specie s of

so mosses investigated are sufficiently sensitive to 2 so

so that the pre sence of moderate leve ls of 2 in the ir environme nt will be sufficient to exclude the se species 42

from the area. APPENDIX

4 3 44

Table I. Chlorophyll content of fumigated mosses. so2 - Cone�- -i1ow - Age Chl . -A ---- Av . v .-% Chl.- % Genus (mo.) so Rate x T-1 T-2 T-J A-1 A-J ppm2 . (lpm) 1 A-2 Aulacomnium 1 0 10(%0) 0 (%) F 100 Aulacomnium 2 .2 1 0 100 100 0 100 F 100 2 .2 } Aulacomnium 2 .2 1 0 100 0 100 Aulacomnium 0 1 1'' 1.566 0 } 0 2 c .882 + Aulacomnium 2 1. 0 1 0 100 0 100 \..n Aulacomnium 2 1.0 1 F 0 100 100 0 100 100 F } } Aulacomnium 2 1. 0 1 0 100 0 100 Aulacomnium 0 1 F 1.566 0 0 2 0 1 c 0 100 .8820 100 Aulacomnium 5. F 2 .0 1 0 100 0 100 100 Aulacomnium 2 5 100 0 1 F 0 0 } 0 100 } Aulacomnium 5. F 10 2 0 1 0 6 0 1.311 9 Aulacomnium . 2 c 8 Age Cone. Plow --v ---�-- A . l . Chl . - C % h Av. % Genus A-1 (mo. ) s x T-1 T 2 A-2 A- J ppom2 Ratelpm) 1 - T.- J Aulacomnium ( (�) {%) 2 Aulacomnium 2 .2 F 0 100 0 100 100 Aulacomnium 2 .2 2 F 0 } 100 0 100 } 100 Aulacomnium 2 .2 2 F 0 100 0 100 2 0 2 c 1.884 0 1.019 0 Aulacomnium +c- F (.), Aulacomnium 2 1. 0 2 0 100 0 100 Aulacomnium 2 1.0 2 F 0 100 } 100 0 1 00 100 Aulacomnium 2 1.0 2 F 0 100 0 100 } Aulacomnium 2 0 2 c 1.884 0 1.0 1 9 0 . 0 F Aulacomnium 2 5 2 0 100 0 100 Aulacomnium 2 5 . 0 2 F 0 100 100 0 100 } 100 Aulacomnium 2 5 .0 2 F 0 100 } 0 100 2 0 2 c 1.117 0 . 5 91 0 ------v---. 1" low --Av .-% --chT.- A -% Age Cone . Chl . Genus (mo. ) s Rate x T-1 T-2 T - 3 A-1 A-2 A-3 o 2 1 ppm (lpm) (%) (%)

Aulacomnium 2 . 2 4 F 0 100 0 100

Aulacomnium 4 F . 8 8 91 . 005 91 95 2 .2 1 6 5 } } Aulacomnium 2 .2 4 F .134 88 . 036 94 0 Aulacomnium 2 0 4 c 1.117 0 . 591 .{::- ---.J Aulacomnium 2 1. 0 4 F 0 100 0 100 Aulacomnium 2 1 .0 4 F 0 1 00 1 00 0 1 00 1 00 Aulac omnium 2 1. 0 4 F 0 100 } 0 100 } 0 Aulacomnium 2 0 4 c 2.015 0 1 . 3 15 Aulac omnium 2 5 . 0 4 F 0 100 0 100

Aulacomnium 2 5 . 0 4 F 0 1 00 1 00 0 100 1 00 Aulacomnium 2 5 .0 4 F 0 100 } 0 100 } Aulacomnium 2 0 4 c 1.842 0 1.241 0 - - - Age Cone. --- - - ChI . ---A - 7fv . % Genus (mo. ) -1"Ratelo w v : �;, Chl . s x T-1 T - 2 T - J A-1 A-2 A - J ppom2 (lpm) 1 Aulacomnium (%) (�'a) 4 .2 1 F 1 . 8 5 9 31 1.036 3 6 Aulacomnium 6 23 29 1 .133 3 Aulacomnium 4 .2 1 F 2 . 074 JO .0938 42 Aulacomnium 4 .2 1 F 1.805 JJ 4 0 1 c 2.694 0 1.6 18 0 } } +:- Aulacomnium co .874 39 Aulacomnium 4 1.0 1 F 1.647 3 6 88 5 44 Aulacomnium 4 1.0 1 F 1.7 5 0 3 2 3 0 .7 4 } 4 2 4 48 Aulacomnium 1.0 1 F 2.007 2 .7 5 4 0 0 1.4 2 0 Aulacomnium 1 c 2.5 73 } 3 0 0 Aulacomnium 4 5 . 0 1 F 0 100 10 4 5 . 0 1 F 0 1 00 1 00 0 1 00 1 00 Aulacomnium - Aulacomnium 4 5 .0 1 F 0 100 } 0 1 0 0 } 4 0 1 c 2.22 2 0 .909 0 e Cone. iTow Av . Av . - f..g % --Chl % Chl . - . Rate A-2 G nus mo. s A- e T A- ( ) x T-1 T-2 - 3 1 3 o2 l 1 ppm ( pm ) (�) (%)

Aulacomnium F Aulacomnium 4 .2 2 2.298 24 1.224 34 Aulacomnium 4 .2 2 F 2.238 26 27 1.26 1 3 2 } 33 4 .2 2 F 2.087 31 1.243 33 Aulacomnium 4 0 2 c 3.024 0 } 1.8 55 0 +- a \() Aulcomnium 47 4 1.0 2 F 2 .11 7 .95 0 Aul comnium 4 a 48 9 .93 2 Aulacomnium 4 1.0 2 F 1.84 5 J39 37 } Aulacomnium 4 1.0 2 F 1.7 54 42O ,861 5 2 3 0 1.7 93 0 4 0 2 c .024 } Aul comnium F a 0 4 5 .0 2 0 1 00 0 10 Aulacomnium F Aulacomnium 4 5 .0 2 .262 91 } 96 .22 3 86 } 92 Aulacomnium 4 5 . 0 2 F .088 97 . 160 9 0 4 0 2 c 2.915 0 1.596 0 J..ge Cone . 1� low Chl . Av . hl. Av . r,, C % mo . Rate x T-1 T - 2 T - J A-1 A-2 A-J Ge nus ( ) s 1 o 2 ppm ( lpm ) ( '�) ( % )

Aulacomnium 4 .2 4 F 1.454 28 . 801 Aulacomnium 4 . 2 4 F 1.3 53 33 } -1 .)·: .764 39 Aulacomnium •. 44 4 2 4 F 1.2 52 3 8 . 69 0 Aulacomnium 0 4 0 4 c 2.019 0 1.23 2 �: } \;\ a 6 0 Aul comnium 4 1.0 4 F 1.2 37 58 . 527 9 7 Aulacomnium 4 1.0 4 F 1.59 0 46 48 . 8 5 0 5 0 5

Aulacomnium 4 1. 0 4 F 1.766 40 . 8 1 6 52 Aulac omnium 4 2 .944 0 1.700 0 4 0 c } } a Aul comnium 4 5.0 4 F 0 100 0 100 Aulacomnium 4 5.0 4 F 0 100 1 00 0 1 0 0 } 100 Aulacomnium 0 0 0 1 00 4 5.0 4 F 10 } Aulacomnium 4 0 4 0 4 c 2 . 640 0 1. 3 2 Blow ge Cone. Av .-% J.. Chl . Av.-% -c11-i . Rate x T-1 A -1 mo. T A- ( ) T -2 - 3 A-2 3 Genus so2ppm lpm 1 Aulacomnium ( ) ( ·� ) (%) 8 .2 1 F 1.12 5 . 620 Aulacomnium 1 : .6 13 Aulacomnium 8 .2 F 1 .089 1 1 0 35 8 .2 1 F 1.089 11 } . 627 13 Aulacomnium 0 8 0 1 c 1.2 2 3 0 .721 .__,, ��} ..... Aulacomnium 0 27 8 1 . 0 1 F 1.0 08 .5 3 Aulacomnium 86 22 8 1 . 0 1 F 1.082 1 5 . 5 15 } Aulacomnium �: 4 24 8 1 . 0 1 F 1.04 5 15 ,52 Aulacomnium 0 8 0 1 c 1.2 2 9 0 } . 689 Aulcomnium 8 a 84 .127 5 8 5 , 0 1 F . 226 Aulacomnium 8 8 5.0 1 F . 071 95 8 9 .017 9 93 A 6 ulacomnium 9 8 5,0 1 F .170 88 } .034 } Aulacomnium 0 8 0 1 c 1.4 15 0 . 843 Cone. Flow Chl. Av.-% Chl. Av .-% Age A-2 Rate A- mo. s T-1 T-2 T - A-1 3 Genus ( ) x 3 o2 1 ppm (lpm) (%) (%) Aulacomnium 1.076 17 8 .2 2 F .598 Aulacomnium F .584 19 17 a 8 .2 2 1.06 5 5 Aul comnium F 8 .20 2 1.088 0 .613 1 50 Aulacomnium 2 8 2 1 . 133 .7 1 c \F\. :} } I\.) Aulacomnium 4 22 8 1. 0 2 F . 895 1 8 . 5 1 19 Aulacomnium F 68 1.0 18 8 2 .949 13 15 . 5 } Aulacomnium 0 F 8 1. 2 1..903918 14 } , 57 5 1 7 Aulacomnium 0 0 0 .693 Aulaco 8 2 c mnium F 82 8 5 .0 2 .287 71 . 085

91 SJ • 019 Aulacomnium 96 92 8 5 .0 2 F . 089 } Aulacomnium F • 0 5.0 98 A 8 2 . 129 87 1 0 0 0 4 ulacomnium 4 8 0 . 7 8 2 c .9 9 } -Av Cone� -r,low �-% Chl. Av .-% J..ge - - Ch l.

A-1 A - A - Rate 2 3 mo. s T-2 T-3 Genus ( ) x T-1 o2 1 ppm (lpm) (%) (%)

Aulacomnium 21 a 8 .2 4 F . 8 97 14 .476 Aul comnium 488 1 9 1 9 14 14 . 8 �2 4 F . 897 } a 0 17 Aul comnium . 5 1 8 .2 4 F . 87 14 a 60 0 Aul comnium 3 8 0 4 c 1.04 3 0 . } \,J\ \.;.) Aulacomnium . 398 4 5 8 1.0 4 F . 802 3 5 Aulacomnium . 93 1 8 33 8 1 . 0 4 F 1.023 17 } 28 5 Aulacomnium 46 3 6 8 1. 0 4 F . 83 8 3 2 . 3 } Aulacomnium 23 0 8 0 4 c 1 . 233 0 .7

a • Aul comnium 02 8 5 .0 4 F . 07 95 7

Aulacomnium • • 0 2 92 5 8 5 .0 4 F 084 94 93 7 9

} • Aulacomnium 0 6 96 8 5 .0 4 F . 14 90 3 7 9 Aulacomnium . 8 99 0 8 0 4 c 1.4 00 0 } - -- � -% h.ge Cone� !"low Chl. Chl. 'o Genus mo Rate Av . % Av . ( . s ) x T - 1 T-2 T - A- A- ppom2 (lpm) 1 3 1 A-2 3 Bartramia (%) (%) 2 Bartramia �2 1 F 0 100 2 0 Bartramia .2 1 F o F10oo0 100 0 1 00 1 00 2 Bartramia . 2 1 F 0 100 0 100 } 2 0 1 c 1 . 943 0 .991 0 \J\ Bartramia .{:'" 2 Bartramia 1.0 1 F 0 0 100 2 0 0 0 Bartramia 1 . 0 1 F r10oo0 100 0 10 } 10 2 0 0 Bartramia 1. 0 1 F 100 0 10 2 0 0 Bartramia 1 c 2.111 0 1.106 2 0 Bartramia 5 . 1 F 0 0 100 2 0 Bartramia 5 . 1 F 0 ro100o 100 0 1 00 1 00 2 0 Bartramia 5 . 1 F 0 100 0 100 } 2 0 1 c 2.4 54 0 1 .33 2 0 A e Cone. flow Av. Av . g Chl. - % �c1a . - %

Genus o A - A-2 A-3 ( m ) s Rate x T-1 T-2 T - 3 1 . o2 1 ppm ( lpm ) (%) (%)

Bartramia 2 �2 2 F 0 100 0 1 0 0 Bartramia 0 0 0 0 2 .2 2 F 0 100 } 10 10 } 10 Bartramia 2 .2 2 F 0 100 0 100

Bartramia 2 0 2 c 2.4 54 0 1 . 332 0

Ui. \.. .n Bartramia 2 1.0 2 F 0 100 0 100 Bartramia 2 1.0 2 F 0 100 1 00 0 100 } 100 Bartramia 2 1.0 2 F 0 100 } 0 1 0 0 Bartramia 2 0 2 c 2.111 0 1.1 06 0

Bartramia 2 5.0 2 F 0 100 0 100

Bartramia 2 5 .0 2 F 0 1 00 1 00 0 1 00 1 00 Bartramia 2 5.0 2 F 0 100 } 0 1 0 0 } Bartramia 2 0 2 c 2.454 0 1.3 32 0 - - -J:i' - - - lo Chl. • • A.ge Cone . w Av - % -----ctiI Av .- % Genus ( mo . ) s Rate x T-1 T -2 T - 3 A-1 A-2 A- J o 2 1 ppm ( lpm ) ( 1o) (%)

Bartramia 2 .2 4 F 0 100 0 100 Bartramia 2 .2 4 F 0 1 00 l- 100 0 1 00 1 00 Bartramia 2 .2 4 F 0 100 0 100 } Bartramia 2 0 4 c 1.874 0 .848 0

\..n Bartramia 2 1.0 4 F 0 100 0 100 °' Bartramia 2 1.0 4 F 0 1 00 1 00 0 100 } 100 B artramia 2 LO 4 F 0 100 } 0 100 Bartramia 2 5 0 4 c 1.874 0 .848 0 Bartramia 2 0 4 5.0. F 0 100 0 100 Bartramia 2 4 F 0 1 00 1 00 0 1 0 0 } 100 Bartramia 2 5 .0 4 F 0 100 } 0 100 Bartramia 2 0 4 c 2 .36 9 0 1.473 0 -�-- Chl. A Chl. v. Age Cone . Flow Av . % % Genus mo . ) s Rate x T-1 T -2 T - J A-1 A -2 A- ( o2 1 J ppm ( lpm) (%) (%)

Bartramia 4 . 2 1 F . 4 08 72 .17 5 7 5

Bartramia 4 .2 1 F . 889 39 53 .412 4 1 56 Bartramia 4 . 2 1 F , 7 58 48 , JJ5 52

Ba.rtramia 4 0 1 c 1.45 8 0 .698 0 } \..!"\ } " Bartramia 4 1.0 1 F . 598 5 9 .246 6 5

Bartramia 4 1 . 0 1 F .787 46 57 .JJ 8 52 62

Bartramia 4 1.0 1 F , 4 96 66 .218 6 9

4 0 • 04 0 Bartramia 4 0 1 c 1. 58 } 7 } Bartramia 4 5 .0 1 F 0 100 0 100 Bartramia 4 5,0 1 F 0 100 } 100 0 1 00 1 00 0 100 Bartramia 4 5 .0 1 F 0 100 } Bartramia 4 0 1 c 1.J Ol 0 ,74 9 0 -- Age Cone , Flow Chl . -Av . �% Chl. Av . % Genus ( mo. ) s Rate x T -1 T -2 T -J A-1 A-2 A- J o2 1 ppm ( lpm ) ( 1o) (%)

Bartramia 4 .2 2 F .8J O 5 0 . J46 58

Bartramia 4 .2 2 F . 647 6 1 54 .288 6 5 59

Bartramia 4 .2 2 F .81J 5 1 . 3 7 9 54 Bartramia 4 0 2 c 1 . 6 59 0 } .82J 0 } \Jl 6 (X) Bartramia 4 1.0 2 F . 40J 6 9 .26 3 5 64 6 Bartramia 4 L O 2 F . 520 60 62 .27 0 3

Bartramia 4 1.0 2 F , 559 57 .JOO 60 Bartramia 4 0 2 c 1.3 01 0 } .75 0 0 } Bartramia 4 5 .0 2 F 0 100 0 100

Bartramia 4 5 .0 2 F 0 1 00 1 00 0 100 1 00 Bartramia 4 5 .0 2 F 0 100 } 0 100 } Bartramia 4 0 2 c 1.2 38 0 . 5 80 0 �---- Age Cone , flow Chi . -Av .-% C hl. Av . % Genus (mo. ) s Rate x T - 1 T -2 T - J A-1 A-2 A- J o2 1 ppm ( lpm ) (%) (%)

66 Bartramia 4 .2 4 F . 47 0 62 . 197 55 .209 64 64 Bartramia 4 .2 4 F .55 7 } 59 } Ba rtramia 4 .2 4 F .495 60 . 220 6 2 0 Bartramia 4 0 4 c 1.23 8 .580 0 I'< Bartramia 4 1.0 4 F .281 7 5 .136 79 'D

562 0 60 .298 54 66 Bartramia 4 1. 0 4 F . 5 } Bartramia 4 1.0 4 F .505 55 } .227 65 Bartramia 4 0 4 c 1.123 0 . 647 0

Bartramia 4 5 .0 4 F 0 100 0 100 Bartramia 4 5 .0 4 F 0 1 00 100 0 100 } 100 Bartramia 4 5 . 0 4 F 0 100 } 0 100 Bartramia 4 0 4 c 1.123 0 .647 0 --·--- Cone , f1 ow Av. Chl . -- Chl. Av Age . % %

s - A - T-1 A Genus mo .) Rate x T - 2 T- A-1 ( o2 1 J 2 J ppm ( lpm ) (%) (%)

• Bartramia 8 .2 1 F 54 0 J6 .27 8 40

Bartramia 8 .2 1 F . 48 1 43 40 .245 47 44 4 8 4 .255 4 Bartramia 8 .2 1 F . 9 1 } 5 Bartramia 8 0 1 c . 844 0 . 46J 0 } °' 0 Bartramia 8 1.0 1 F . 405 52 .204 5 6

I Bartramia 8 1.0 1 F . 625 26 4J J 06 34 50 F 4 4 .18 5 60 Bartramia 8 1.0 1 . 1 51 844 0 . 46 0 Bartramia 8 0 1 c . } 3 } 0 100 0 100 Bartramia 8 5 . 0 1 F a 8 5.0 0 1 00 1 00 0 1 00 100 Bartrami 1 F } 0 0 100 Bartramia 8 5.0 1 F 0 10 } Bartramia 8 0 1 c .929 0 .49 6 0 - Age Cone . Flow Chl. Av.-% Chl�- -- Av . %

Genus (mo . ) s Rate x T-1 T - 2 T - 3 A-1 A -2 A- 3 o2 1 ppm ( lpm) (%) (%)

Bartramia 8 .2 2 F . 580 33 . 24 0 47 Bartramia 8 . 2 2 F .476 45 } 37 .23 6 48 46 a 4 Bartrami 8 .2 2 F . 580 33 .258 3 } Bartramia 8 0 2 c . 865 0 . 4 53 0

°' Bartramia 8 1.0 2 F .33 8 68 . 1 54 74 ,__..

Bartramia 2 F . 686 35 47 .372 4 9 8 LO } J7 } Bartramia 8 1.0 2 F .654 3 8 . 378 J 6

Bartramia 8 0 2 c 1. 055 0 .591 0 F Bartramia 8 5 .o 2 0 1 0 0 0 100 Bartramia 8 5 . 0 2 F 0 1 00 1 00 0 100 100 Bartramia 8 5 .0 2 F 0 100 } 0 100 } Bartramia 8 0 2 c 1 .055 0 . 5 91 0 Chl. Chl. ----- ge Av � Av A Cone �-- RateiTow x T-1 T-2 T-3- % A-1 A- . % s A-2 3 Ge nus ( mo . ) o 1 2 (l m) m p (%) Bartramia 8 pp.2 4 .562 J5 .231 (%49) Bartramia 8 4 F .467 46 42 .227 50 49 Bartramia 8 .2 4 F .476 45 } .236 48 } Bartramia 8 .20 4 F .865 0 .453 0 c °' Bartramia 8 1.0 4 .436 45 .189 5J I\) Bartramia 8 1.0 4 F .397 50 49 .193 52 55 Bartrarnia 8 1.0 4 F .381 52 } .160 60 } Bartramia 8 0 4 F .793 0 .401 0 Bartramia 8 5.0 4 c 0 100 0 100 4 F 100 Bartramia 8 5.0 0 100 0 100 100 F } } Bartramia 8 5.0 4 F 0 100 0 100

a r ramia 8 0 4 .969 0 .502 0 B t c Age Cone . -Flow Chl . �-Av .-% Chl. Av. % Genus s x T-1 T - 2 T - J A- A -2 A- . (mo. ) o2 Rate 1 1 3 ppm (lpm ) (%) (%)

Leucobryum 2 .2 1 F .110 84 .022 93 Leucobryum 1 .0 87 6 2 .2 F 9 0 BJ .012 9 } 93 Leucobryu m 2 .2 1 F .152 7 8 . 03 1 9 0

• 0 Le uc obryum 2 0 1 c 69 0 0 . 312 } °' Le uc obryum 2 1 . 0 1 F .069 .005 \.;.)

• Leucobryum 2 1.0 1 F 07 5 93 .01 1 98 Le uc obryum 2 1. 0 1 F 0 100 0 100

Le uc obryum 0 68 0 .26 0 2 1 c . 5 :: } 5 :: } Le uc o bryum uc 2 5 .0 1 F 0 100 0 100 obr um 2 . 0 1 F 0 1 00 1 00 0 1 00 1 00 Le ucobryyum 5 0 0 Leu b 2 5 .0 1 F 0 100 } 10 } co ryum 2 0 1 c . 685 0 .265 -- �% ---- Age Cone . flow Chl . Av . o;o Chl. Av. % Genus ( mo . ) s Rate x T-1 T - 2 T - 3 A-1 A-2 A- 3 o2 1 ppm ( lpm ) (%) (%)

F Leuc obryum 2 .2 2 .052 :� .006 :: 0 1 98 Le ucobryum 2 .2 2 F 0 1 93

Leuc obryum 2 F .096 87 • 014 87 2 .2 } } 0 .281 0 Leucobryum 2 0 2 c . 740 °' 0 0 100 +:- Leucobryum 2 1.0 2 F 0 10 Leuc obryum 2 1. 0 2 F • 05 0 93 } 98 • 012 96 } 99 100 Leuc obryum 2 1. 0 2 F 0 100 0 9 0 Leucobryum 2 0 2 c .713 0 .2 3 0 0 Leuc o bryum 2 5 . 0 2 F 0 100 10 Le uc obryum 2 5 .0 2 F 0 100 1 00 0 100 } 100 Le uc obryum 2 5 . 0 2 F 0 100 } 0 100 • .302 0 Leucobryum 2 0 2 c 7 02 0 - f..ge Cone flow -cfi1 .� Av. % Chl. �Av--:-% Rate T-1 T - 2 T - J A-1 A-2 A-J Genus ( mo . ) s x o 2 1 ppm ( lpm ) ( <(o) ( �1a)

Le uc o bryum 2 .2 4 F 0 100 0 100

028 6 99 . 005 98 99 Le ucobryum 2 .2 4 F . 9 } } Le uc obryum 2 .2 4 F 0 100 0 1 0 0 1 0 Le ucobryum 2 0 4 c . 6 93 0 0 . 27 °' l...r\ Le uc o bryum 2 1.0 4 F 0 100 0 100

0 00 100 0 100 100 Leucobryum 2 1.0 4 F 1 } Le ucobryum 2 1.0 ·4 F 0 100 } 0 100 0 Le uco bryum 2 0 4 c . 6 53 0 .24 9

Leuc o bryum 2 5 .0 4 F 0 100 0 100 Leuc o bryum 2 5 .0 4 F 0 100 1 00 0 100 } 100 0 1 00 J.,euc o bryum 2 5 .0 4 F 0 100 } 6 0 0 .3 2 0 Leucobryum 2 0 4 c . 9 1 -- - -- ·--Av. . % % v Age Cone . 1"1.ow Chi. Chl. A T-1 T 2 T - A- A- A-2 - ) Genus ( mo . ) s Rate x ) 1 o 2 1 ppm ( lpm ) ( %) (%)

Le uco bryum 4 .2 1 F .137 . 020 84

Leuco bryum 4 .2 1 F . . 371 67 . 125 6) 73 - Leuco bryum 4 .2 1 F . 291 64 . 095 7 2 J.,e uc o bryum 4 0 1 c . 807 0 .JJB 0 } :� } O'\ °' Le ucobryum 4 1. 0 1 F .174 7 5 .074 77 Leuco bryum 4 1 . 0 1 . 1 05 8 5 8 1 .01 6 95 89 F } } J.,e ucQbryum 4 1. 0 1 F .119 8) .01 6 95 0 Leuc obryum 4 0 1 c . 697 0 .)23 . 0 Luecobryum 4 5 0 1 F 0 100 0 10

Leuco bryum 4 5 .0 1 F 0 1 00 0 1 00 100 Leuco bryum 4 5 .0 1 F 0 100 } 100 0 100 } :I:,euc o bryµm 4 0 1 c .76 5 0 . J 04 0 -- · h e - v � - - --% g Chl v Cone . ·1;i]6w x T-1 . AT-3. -% A-1Chl .- A . Genus ( mo . ) s Rate 1 T-2 A-2 A-J o 2 4 ppm ( lpm ) .35 ( % ) .178 ( �'o)5 3 60 Leucobryum .2 2 7 4 F .4 52 56 .182 56 54 F 3 Le uc obryum 4 .2 2 .3882 56 } 49 } uc o bryum 2 .21 1 I.e 4 . F .41J 0 Leuco bryum 0 2 c . 882 0 °' 4 2 . 79 91 " 0 100 Le uc o bryl.lm 4 1. 0 F .1041 84 89 94 98 02 Le uc obryum 1.1.0 2 F } . 5 } 2 0 100 Le uc o bryum 4 0 2 F . 88071 9 .413 0 2 0 Le uco bryum 4 0 2 c 0 100 Le uc obryum 4 5 . 0 2 F 0 100 100 100 Leucobryum 5 . 0 2 F 0 1 00 1 00 0 4 F 5 0 Leuc obryum . 0 2 .74 100 } 1 0 } 4 0 3 0 .288 Le ucobryum 0 2 c - J...ge Cone . Plow Chl . Av.-% Chl. -Av.-% x T-1 T -2 T - 3 A-1 A-2 · A - 3 Genus ( mo . ) s Rate 1 o2 4 ppm ( lp4m ) ( ·1o)4 (%) Leucobryum .2 .184 7 .115 F .129 9 4 4 . 9 3 6 33 2 6 Leucobryum .2 F 5 } 4 4 . 4 3 81 . 080 1 77 J:,e uc obryum 4 .2 4 F . . 48 :: 3 0 u(! o bryum 0 c 7 06 0 lie °' 96 l co 4 4 00 Leucobryum 1. 0 4 F . o:r n 0 1 83 4 6 Leuc obryum 0 . 180 7 6 88 . 112 5 1. F } 4 4 92 4 - 051 8 Leuco bryum 4 1.0 4 F . 06490 . Leucobryum 4 0 4 c 7 0 } .32 1 0 Leucobryum 5 .0 F 0 100 0 1 00 4 4 00 100 0 00 00 0 1 Leucobryum 4 5.0 4 F 1 1 0 1 00 Le ucobryum 5 . 0 F 8 0 100 } .} 4 4 . 19 0 Leucobryum 0 c 0 . 344 A e Cone . - - Flow Av.-% g Chl . -cn-1. Av . -% Rate x T-1 T 2 T - A- A- - 3 1 3 Genus ( mo . ) s A-2 o 2 1 ppm ( lpm ) (%) � (%) 8 .2 1 .288 56 .112 60 Leuc obryum 8 .2 1 F .321 51 54 .154 45 53 Le uco bryum 8 .2 1 F .294 55 .128 54 } 1e uc obryum 8 0 1 F .654 0 .279 0 Leuc o bry\,lm c } °' 8 1. 0 1 .319 55 .092 62 \0 Le ucobryum 8 1.0 1 F . 54 50 53 .104 59 3 57 Leuco bryum 8 1.0 1 F . 26 54 } .101 } 3 8 5 Le uc obryum 0 1 F . 08 0 .241 0 Leucobryum 8 5.0 1 c 7 0 100 0 100 Le ucobryum 5.0 1 F 0 100 100 0 100 100 Le ucobryum 8 5.0 1 F 0 100 } 0 100 Leucobryum 8 0 1 F 0 .29 0 } c 7 Leucobryum .633 ---··------h.ge Cone . ?low x T-1Chl . -ATv.- �% A-Chl1 . A-Av3. % s 3 A -2 Genus (mo . ) o Rate 1 T-2 2 8 ppm (lpm) .319 (%)1 (%6)5 Le ucobryum 8 .2 2 F 6 5 .2.02690 74 63 L eucobryum 8 .2 2 F .J.45ll1 46 25 } 6 .174 50 Le uc o bryum 8 .20 2 F .819 0 .347 0 Leucobryum 2 c } --J 8 1. 0 2 F .401 51 .160 54 0 Leucobryum

6 • 07 79 60 8 1. 0 F . 29 7 1 3 Leucobryum 8 1.0 2 . 2 28 602 .184 47 F 3 Le uc obryu.m 8 0 2 .819 0 .347 0 Le uc obryum 8 5.0 2 cF 0 100} 0 100 } Leuco bryum 8 5.0 2 F 0 100 100 0 100 100 Le uc o bryurn 8 5.0 2 F 0 100 } 0 100 } Leuco bryurn 8 0 2 .767 0 .302 0 Leucobryum c ------Av:- Av.-% % Chl. Age Cone . Plow Chl .

T-2 T-3 - 1 A-3 x T-1 A-2 A Genus ( mo . ) s Rate 1 o 2 8 ppm ( lpm ) . 85 (%) .151 ( %5 8) F 3 2 Le uc obryum .2 4 0 550 .10 1 59 . 1 51 6 8 4 Leuc obryum .2 4 F . 4 9 51 .15 58 8 1 F 3 3 Leuc obryum 8 .20 4 . 80 0 .359 0 Le ucobryum 4 c 1 } -.,,) } � 55 • 0 8 0 .281 062 7 1. F Leuco bryuro 4 0 9 . 62 8 0 .250 53 62 F 60 7 Leuc obryum 1. 4 4 } .096 5 . 9 4 8 F 1.0 3 44 _I,e ucobryum 8 0 4 0 .208 0 Le uc obryum 8 5.0 4 c . 6 240 100 0 100 } Le_ uc o bryum 8 .0 4 F 0 100 100 0 100 100 5 F Leuc obryum 8 5.0 4 0 100 0 100 Leuco bryum 4 F } } ' 0 0 • 208 0 8 4 Le_ug_o J:>ryum 4 c . 62 J...ge Cone . PTow Chl . Av . % Chl. Av .-% x T-1 T -2 T - 3 A-2 A- Genus ( mo . ) s Rate 1 A-1 3 o 2 2 pp.2m ( lpm)1 0 (%100) 0 (%10)0 Polytrichum 2 .2 F 47 6 8 .06 89 95 1 F .1 9 9 5 Polytrichum } } 2 .2 1 I 021 98 I 020 96 Polytrichum F

2 0 1 1.05 0 I 506 0 Polytrichum c 3 --..) 2 0 0 0 0 100 I\) 1. 1 F 10 Polytrichum 2 0 0 100 100 0 100 100 1. 1 F Polytrichum 2 0 0 0 100 } 1.0 1 F 10 Polytrichum 2 0 1 1.091 } 0 Polytrichum c 0 ,543

2 . 0 1 F 0 100 0 100 Polytr ichum 2 5 .0 0 100 100 0 100 100 5 1 F :Eolytrichum 2 .0 0 0 0 100 5 1 F 10 Polytrichum 2 0 1 1.122 0 } 0 } Polytrichum c ,549 ------% -�-- - - Age Cone . Flow ChI. Av . o;" Chl. Av .-% Genus ( mo . ) s Rate x T-1 T-2 T - o2 1 3 A-1 A -2 A- 3 ppm ( lpm ) F (%) (%) Po lytr ichum 2 . 2 2 0 100 0 1 00

F • Polytrichl.l_I!l 2 .2 2 F . 14 9 85 } 95 037 91 } 97 Polytrichum 2 .2 2 0 100 0 100

Polytrichum 2 0 2 c , 993 0 . 4 09' 0 ---J Polytrichum 2 1.0 2 F 0 100 0 100 'vJ Polytrichum 2 1.0 2 0 100 } 100 0 1 00 1 00 Polytrichum 0 2 1.0 2 F 0 100 0 10 } Polytrichum 0 0 6 0 2 2 Fc 1.24 3 . 11 Polytrichum 2 5 . 0 2 F 0 100 0 100 Polytrichum 2 5,0 2 F 0 1 00 100 0 1 00 1 00 Polytrichum 2 5 . 0 2 0 100 } 0 100 } Polytrichum 2 0 2 c 1.2 88 0 ,73 2 0 Age Cone . F1 ow Chl . Av . % Chl. Av.-% Genus ( mo . ) s Rate x T-1 T - 2 T - J A-1 A-2 A- J o2 1 ppm ( lpm ) (%) ( %)

Polytrich_lJ.m 2 . 2 4 F 0 100 0 100 Polytrichum 2 . 2 4 F 0 1 00 1 00 0 1 0 0 } 1 00 Polytrichum 2 . 2 4 F 0 100 } 0 100 Polytrichum 2 0 4 ·c . 879 0 . J83 0 --.J Polytrichum 2 1. 0 4 F 0 100 0 100 +:- Polytrichum 2 1.0 4 0 1 00 1 00 0 1 00 } 100 Polytrichum 2 1.0 4 F 0 100 } 0 1 00 Polytrichum 2 0 4 c 1 . 222 0 . 540 0 Polytrichum 2 5 .0 4 F 0 100 0 100

Polytrichum 2 5 .0 4 F 0 1 00 1 00 0 1 00 1 00 Polytrichum 2 5 . 0 4 F 0 100 } 0 100 } Polytrichum 2 0 4 c l . 200 0 . 67 2 0 FI -- Age Cone . ow x T-1Chl . AvT-J.- % Chl . Av-.-% s T-2 A-2 Ge nus ( mo . ) o Rate 1 A-1 A - J 2 ppm ( lpm ) (%6) .40J (%)4 4 .2 1 F .918 2 3 Polytrichum 4 1 F ,956 23 24 .446 27 JO Polytrichum 4 .2 1 ,956 23 } .4J4 29 } Polytrichum 4 0 1 F 1.241 0 .611 0 Polytrichum c -.._J .J42 44 l...n 4 1.0 1 F , 5 39 Polytrichum 4 1.0 1 ,79973 20 28 .4 1 2 JJ Polyt�ichum F .4407 287 } 4 1. 0 1 F ,931 25 Polytrichum 4 0 1 1.241 0 .661 0 Polytrichum 4 5,0 1 c 0 100 } 0 100 Polytrichum F 0 0 100 100 4 5.0 1 F 0 100 10 Polytrichum 4 5,0 1 F 0 100 } 0 100 Polytrichum 4 0 1 1.419 0 1.419 0 } Polytrichum c -� �- Age Cone .- Flow ChL- Av .-% Chl . Av .-% Genus ( mo . ) s Rate x T-1 T - 2 T - 3 A-1 A-2 A- 3 o2 1 ppm ( lpm ) (%) (%)

Polytrichum 4 .2 2 F . 8 19 28 .383 2 9 Polytrichum 4 .2 2 F .854 2 5 } 26 .41 0 24 28 Polytrichum 4 .2 2 F . 8 54 25 . 372 31

0 .539 0 Polytrichum 4 0 2 c 1.13 8 } --.J Polytr-_ichum 4 1.0 2 F , 6 53 JJ .25 9 JB O'- Polytrichum 4 1. 0 2 F .53 6 45 } 3 9 .188 55 45 Polytrichum 4 1. 0 2 F , 594 3 9 .242 42 Polytrichum 4 0 2 c , 974 0 . 4 1 7 0 } Polytrj.chum 4 5 , 0 2 F 0 100 0 100 Polytrichum 4 5 . 0 2 F 0 1 00 100 0 100 } 100 Polytrichum 4 5 , 0 2 0 100 } 0 100 Polytrichµm 4 0 2 c 1. 539 0 . 743 0 - - ... - - e- Coric-. - low Chl . Av .- Chl. --- A Ag .ti' % v -. % Ge nus (mo. ) s Rate x T-1 T-2 o2 1 T - J A-1 A-2 A- J ppm ( lpm) (%) (%)

Polytrichum 4 .2 4 F . 865 24 .J72 Jl

Polytrichum 4 . 2 4 F . 922 19 21 .377 3 0 31

67 J2 Polytrichum 4 .2 4 F . 910 20 . 3 Polytrichum 4 0 4 c 1.13 8 0 } . 539 0 } -.._) -.._) Polytricb.um 4 1. 0 4 F I 7 81 4 5 .497 44

6 2 4 .53 2 40 44 Polytrich1.JJT1 4 L O 4 F . 6 5 3 3 } Polytrichum 4 1. 0 4 F . 681 52 . 461 48 Polytrichum 4 0 4 c 1.4 1 9 0 } .887 0 Polyt�ichum 4 5 . 0 4 F 0 100 0 100 Polytrichum 4 5 .0 4 F 0 1 00 1 00 0 100 } 100 Polytrichum 4 5.0 4 0 100 } 0 1 00 Polytrichum 4 0 4 c 1.539 0 .743 0 - - -- C Age Cone-� -Flow - hI . Av. % Chl.- - Av .-% o. s Rate x T-1 T - 2 T - 3 A-1 A-2 A-3 Genus (m ) o2 . 1 ppm ( lpm) (%) (%)

. 4 94 1 6 �olytrichum 8 . 2 1 F . 9 25 1 � . 45 1 2 14 Polytrichum 8 . 2 1 F ,978 11 5 } Polytrichum 8 .2 1 F , 935 12 . 5 06 14

Polytrichum 8 0 1 c 1.063 0 . 5 88 0 } --.) Polytrichum 8 1.0 1 F ,788 22 .34 1 26 ())

.373 1 9 20 Polytrichum 8 1.0 1 li' .828 17 } 19 } Polytrichum 8 1.0 1 F . 8 1 8 1 8 . 392 15

Polytrichum 8 1 c .998 0 . 461 0

. 06 8 Polytrichum 8 5..o0 1 F . 289 71 9 5 Polytrichum 8 5.0 1 F 0 1 00 83 0 100 93 f'_olytrichum 8 5 , 0 1 F .220 78 • 0?8 94 Polytrichum 8 0 1 c . 998 0 } . 461 0 } - � A - Age - Cone . -PTow Chl . v. % C hl . Av .--% Genus mo . s Rate x T -1 T - 2 T - 3 A-1 A -2 A- 3 ( ) o2 1 ppm ( lpm ) (%) ( %)

Polytrichum 8 .2 2 F , 967 .52 3 11 Polytrichum 8 .2 2 F . 999 9 . 517 12 } 1 5 Polytrichum 8 . 2 2 . F . 935 1 . 4_59 22

Polytrichum 8 0 2 c 1.063 0 ,588 0 : } -..J '° Polytrichum 8 1. 0 2 F ,958 1 6 ,56 1 22 Polytrichum 8 1.0 2 F ,958 1 6 } 1 7 ,59 0 18 } 20 Polytrichum 8 1.0 2 F . 924 19 ,57 5 20

Polytrichum 8 0 2 c 1.14 1 0 .71 9 0

Polytrichum 8 5 . 0 2 F 0 100 0 100 Polytrichum 8 5 . 0 2 F • 320 75 } 88 .101 80 1 92 Polytrichum 8 5 . 0 2 F . 14 1 8 9 . 020 96 -

Polytrichum 8 0 2 c L27 9 0 .503 0 �- - �- - - - - "'% - --- - v Age - Coric-. - Flow - �C-hl. Av . Chl. A .-% Genus (mo. ) s Rate x T-1 T -2 T - 3 A-1 A -2 A - 3 o2 . 1 ppm ( lpm ) (%) (%)

Polytrichum 8 . 2 4 F .768 1 .299 23 5

Polytric:_hJ.l.m 8 . 2 4 li1 . 813 1 0 14 .33 0 1 5 2 1

Polytrichum 8 .2 4 F . 7 5 0 1 7 . 291 25

4 • 0 .388 0 Politrichum 8 0 c 9 03 } } co 0 PolytriQ..hum 8 1.0 4 F . 54 37 .213 38 5 4 8 . 1 99 42 39 Polytrichuro 8 1 . 0 4 F . 5 5 3 } 33 Polytrichum 8 1 .0 4 F . 668 24 .216 37

4 87 .343 0 Polytrichum 8 0 c . 9 0 } 0 Polytrich-ym 8 5 . 0 4 F 0 100 0 10 Polytrichum 8 5 . 0 4 F 0 100 1 00 0 1 00 1 00

0 0 0 1 00 Polytrich'Ll.rn 8 5 . 0 4 B 10 } } Polytrichum 8 0 4 c 1 .15 9 0 . 624 0 S t

= c control fumigated

:::hl. T-1 total chlorophyll in mg/gr. fr. wt.

T-2 ) % chlorophyll destruction (gr. fr. wt.

average chlorophyll destruction T - J % A- 1 chlorophylla in mg/gr. fr. wt. A-2 chlorophyll destruction (gr. fr. wt. ) % a A- average chlorophyll destruction J % a 82

a Table II. Moles S02 exposure for three flow r tes and three so2 concentrations during fumigation. . 83

2lowrate Concentration Moles so2 S02

(Liters per minute ) ( ppm)

1 .2 . 0015

1 1. 0 . 007 5

5 .0 . 0375 1

2 . 2 • 0030

2 1. 0 .01 5 0

2 5 .0 .0750

0060 4 .2 .

1.0 . 0300 4

.0 .1 00 4 5 5 ?igure I. Degradation of chlorophyll to phaeophytin . a a H C-CH H H2 C-CH 2

tt3c tt3c C 2H 5 C H I " I 2 5 "\. N N N / < / low so2 ""' ,// "' ·?- g H H 2H /Nl / cone . ' N / N/ \ N \o;i ;'\ / I H c CHJ HJC CHJ 3

c CH �2 I 2 c CH 2 12 I p H39c2 0ooc c H ,., o oc' 2 0 OCHJ OCH 3 9\J J

Ch lor ophyll Phaeophytin a a 86

?igure II, Illino is distribution of mosses used in

fumigation. 87

Aulacomnium heterostichum Bartramia pomiformis

leucobryum glaucum Polytrichum ohioense 88

Figure III . ?um igat ion apparatus for exposing

mosses to sulfur dioxide. i A . E.

i

B. i'. c:®:J i ./. .. .j. � .., v.

r •J .

D .

�ylind er of A. Air pump E. S02 Drierite filter llowme ter B. ? .

.., v • liter exposure chambe r .:.-: lowme ter G. 7

D. Wa tel"' trap watt incandescent lamp H. 250 9 0

Figure . Res ponse of game tophytes of and IV 4- 8-

month old Aulacomnium heterostichum �o so2

concentrations at different flow rates . 9 1

...... '\ s \ A A \ ' \ .._.. 't:I Cll r--i \ s:: ' 0 \ 0 \ 0 ...... Cll \� \ . +> \ l ..c: ' C\J ro +> ' S-4 ' s:: " +> 0 s:: s Q) 0 () co s:: .-I 0 0

C\J 0 fJl C\J

Q) Q) (j) 0 +> +> +' ::s ::s :::s 0 0 0 0 0 0 -- s:: ...... c 0 co ..:::t C\J ...... -I '-{): s s E

S-4 � � • M • uo1+on.:r+sap TTAt{d .I Tt.£0 Q) Q) Q) ( '.l- ·�J .I� ) O O % A A A

Cll Cll er. � � � Q) Q) Q) +> +' +' ...... • -! r-i r-1 r-1

0 0 0 ...... -I C\l _, [ A ..._..

"O Cll r-i \ s:: 0 \ 0 \ \ 0 ...... Cll \ r. +> ..c: \ C\J ro +' \ � s:: \ \ +:> 0 \ s:: s \ Q) \ \ 0 () ..:::t \ s:: .-I 0 ' .. \ () ' ' C\J ' \ 0 ' ' fJl ...... C\J 0

0 0 0 0 0 0 0 co '-C ..:::t C\J .-I

( • +M • .I.J • .I�) uo1 + ;:m.r1sap TTAtidO .IO P..[2 " 92

Figure Response of garne tophytes of and V. 4- 8-

rnonth old Ba.rtrarnia porniforrnis to so2 conc entrations at different flow rates. 1.0 liters per minute --- liters er minute 2.0 p

...... e 4 .o li tero p r minute 4 months old 8 months old ·- ,...... 100 . 100 . +> +> � ;: � . . / S--4 / �l � I r1 -1 . 80 . 80 S-t f..t b.0 l:lD - ...... :;...... '° � . � ..-:: ... � - \.....) 0 - .. 0 ·.-i ...... � .... ·d 60 ,_ -- � -- 6_ o I +> I ...... +> 0 I (.) ::s ::s S--4 S--4 p +> Cl) Ul Ql Ql 'O 40 'O 40 rl rl rl rl » » � ..c: Pt Pt 0 0 H 20 H 0 0 20 rl rl � ..c: r) 0

\;� *- o r ' I 0 I � 0.2 1 .0 2.0 5.0 0.2 1.0 2.0 (p m) so2 c oncentrat ions p - ' so2 concentrations (ppm) 94

Figure Res ponse of game tophytes of 4- and VI . 8-

month old Leuc o bryum glaucum to so2

concentrations at different flow rates. liters er minute 1.0 p liters er minute --- 2.0 p

-·--- --· L� . 0 li ter�; pe r minute 4 months .old ] 8 months old . --... 100 ...... 100 +>. 1 � +>. 3: :;: - S... �I. r, _, / Ci-1 r- -=-=� 80 I 80 H. H. QI) QI)

- ...... � .,.,..., h h · '° 0 0 ,,,,...... � \...r\ ·ri 6 •rl 6 0 --- +> 0 +-> - () 0 -- ::l ::l S... H r-:-:-..- p +> (/) I Cl) (j) (j) 'd 40 'd 40 1 r-l r-l rl r-l � � ..c'. ..c: P! P! 0 0 H 20 H 0 0 20 rl r-l � ..c: r) 0

\" *- ...... 0 l f I 0 I I � 0.2 1. 0 2.0 5 .0 0.2 1.0 2.0 concentrations conc entrations so ( ppm ) so (ppm) 2 2 96

Figure Respons e of game tophytes of and VII. 4- 8-

month old Polytrichum ohioense to so2 concentrations at different flow rates. liters per minu te 1.0 --- liters er minute 2.0 p

---·--- · nu l� . 0 l i t.e ru pe r mi Le months 4 old 8 months old . .-... ,...... 100 . 100 , .. .. .p .p , .. , � � , .. . / . , . / S-i � �� .p , .,,...,·/ , UJ UJ , Q) Q) // / 'd 4 0 '/ 'd 40 "' "' rl rl "' rl // rl / » , » 4 / ;:: .. P. -a , , 0 0 . , S-i 20 S-i 20 .. '7 0 0 � , rl rl ..r...: ..c: ! ,,,� ( ) (.) I '/" I \'' � 0 0 . r---s.o 0.2 1 .0 2 0 5.0 0.2 1.0 2.0 concentrations concentrations so ( ppm ) so2 ( ppm ) 2 98

Figure VIII. Responses of 4-month old gametophytes

of various mosses to different so2

conc entrations at 1. 0 liter per

minute . ;:;:;:;:;:;:;:; Aulacomni um he te rostichum

� Ba rtamia pomi formis

Lc uc o bryum gJauc urn

f-H Po lytrichum ohi oense - . 100 +> � : � ; ; :•··· :·· �� . :::::· H ::.: c.-.. ::::::: � Bo . - ;1;��;::::::: �� �, S-t r-i ' 0 �- ti.() '° - s.. ������ � '° +> � � ��= 0 0 1l���ll.. . •rl 0 60 � +> . � :;:�:�: . 0 0 � � :::s +> �=1=�=� S-t +> "Cl � ::::::: � Ul Q) :·. ::::: � Q) s... �:i:i� "Cl 4 ····••· � a ...... � rl E :::::� � r-i 0 � » 0 .c - � P. � ::: 0 � J�J::: � S-t 20 0 � r-i � : � -� � : :::: I) t ii: �i�� ' ., 1�t � "' -.; 0 � :::. ·-::�· � 0.2 1.0 5 . 0 � (ppm ) so2 conc entrati ons 100

Figure IX. Responses of 4-month old game tophytes of

various mosses to different so 2 conc entrati ons at liters per 2.0

minute . ;:;:;:::;:;:;:; Aulacomnium heterostichum

am � Ba rt i a pomiformis

uc o urn a uc IA) twy L�J urn

I I I Po lytrichum oh ioense - . 100 +> � � � �.m� � . �. 't-1 � - 80 ...... f.-t r-i0 ...... !':ID 0 f.-t �· ...... +> 0s:: s::0 •rl () 60 � CJ 0 ::s +> � f.-t � 'd � (/) ()) � ()) f.-t 4 � 'O a m� �� r-i � �� rl s0 » 0 m ��� .c: ...... � ™ P. =�=== m � 0 !�� � m � f.-t 20 0 :1:1:1 rl � . � .J...� � �=:=:: I ) m::::: � � I � ' : .. "' � 0 i�!ti!�! � 8 j)l�jl � � 0.2 f 1. 0 5 .0 so2 concentrations (ppm ) 102

?igure X. Responses of 4-month old gametophytes of

various mosses to different so2 conc entrations at 4.0 liters per minute . ;:;:;:;:;:;:;:; Aulacomnium heterostichum

Bartamia formis � pomi Le ucobryum t�Juucum

Polytrichum I I I ohioense ...... 100 +>. w � � . � t ....I �� H. "'"" '+-i . � Bo . � H...... rl tU) 0 * ,._. 0 - H I....) +> � 0s:: s::0 @ro � •r-i 0 60 +> 0 0 � � � :::s +> � w* · � H �­� +> 'O . Ul Q) \I) $...t * 'O 4 �� a � � rl i:: rl 0 � 0 � :>, .£:; - m� � 0. ili �- 0 � H 20 � � . � 0 � rl M -� I ) � � @ ',, I. �. ; � .,, 0 � � � � 0 .2 1 .0 5 . 0 so2 concentr� ations ( ppm ) 104

?igure XI. Res ponses of 8-month old game tophytes of

various mosses to different so2

conc entrations at 1.0 liter per

minute . ·:·:·:·:·:·:·=· :.;.;.;.;.;.;.: Aulac omnium heterostichum

Ba rtamia pomiformis �

Lc uc ol>ry um g1auc um

+-f-f Polytrichum ohi oense ...... 100 fol � ;c . � l-1 �I 0 - s... .p •:·:·: Vt

0s:: s::0 . .,..; () 60 � .p () 0 �m :::3 .p � � s... �il .p "Cl Ul (),)

?igure Responses of 8-month old game tophytes XII .

of various mosses to different so2 concentration at 2.0 liters per minute . ;:;:;:;:;:;:;:; Aulacomnium heterostichum

� Bartamia pomiformis Lc uco bryum glauc um

� Polytrichum ohioense - • 100 +' � �. "'• ft..i 80 •it':· � · . - ··· · S... rl ·: · ��� � � 0 :� :�: �� - ::::::: � f--> S... 0 +' ...1�;�;�1.... � -..J 0� �0 •ri () 6 i;;;;�� +' � 0 0 · ····· � ••···· � :;::I +' · · S-i ::::::: � �� +' "O ::::::::; �· Ul Q) ::�: ::: � Q) S.. : :·:·:::: � ·:· : : "O · · ·: � a rl ::::::: � rl s0 ::::::: � � () :: ::: � ..c:: - : : At � 0 : S... 20 :: � � : : � 0 !��1�11: rl ..c:: ,_.,· ··· I) ·· � f.f.f.:• : : : : : � ��;���· · ::: � •······ � '"' : : :: � : =: "' �: I :: :: •- � :=�=�==: ��� 0 •.•. ::: : :::::: � � 0.2� 1. 0 5 . O· so2 concentrations ( ppm ) 108

Figure XIII . Responses of 8-month old game tophytes

of various mosses to differ9nt so2

concentrations at 4.0 liters per

minute . : . . .;.;.;· ; ;·.;·:·:·:·:·:;; ;·. AuJacomnium heterostichum

pomiformis � Bartam ia

-- kucobryum L': laucum

I I I Po lytrichum ohi oense ...... 100 +> � .... � . � H � *''� 80 ...... i11mi � H ,...; t ::::::: � � � 0 � @ 0 - H +> \,() i:: i:: I ± 0 0 •rl CJ 6 0 tl � +> m...i�. () 0 ::s +> :::::: . �·� +>H 'O Ul Q) �lll�l � Cl! H � ::;:::: � 4 :ll�::;��� � 'O a •!:;:::::: ,...; 6 � � rl 0 � �l�l�ll· · � » CJ ·•·.·.· •·· · · .c: - :::::: P. : � 0 · @m•·.•.• � H 20 ·• • · � ·�m�·· ·l· · · � ::·:::: · 0 · � : r-1 .•::: .• : .c: : •· � · �!l , ) � ...... : � ' •> "' �i1���i · :�:l:�:·• 0 I��:: :::: · •· : � � · � 0 . 2 1.0 5 . 0 � so2 concentrations (ppm ) LITE RA'rURE CITED

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