Eastern Illinois University The Keep

Masters Theses Student Theses & Publications

1981 A Key to the Hepaticae of Coles and Clark Counties, Illinois Michael J. Klopmeyer 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 Klopmeyer, Michael J., "A Key to the Hepaticae of Coles and Clark Counties, Illinois" (1981). Masters Theses. 2975. https://thekeep.eiu.edu/theses/2975

This is brought to you for free and open access by the Student Theses & Publications at The Keep. It has been accepted for inclusion in Masters Theses by an authorized administrator of The Keep. For more information, please contact [email protected]. Tl I F:StS R F:PRODUCTCON CERTIFICATE

TO: Graduate Degree Candidates who have written formal theses.

SUBJECT: Permission to reproduce theses.

The University Library is receiving a number of requests from other institutions asking permission to reproduce dissertations for inclusion in their library holdings. Although no copyright laws are involved, we feel that professional courtesy demands that permission be obtained from the author before we allow theses �o be copied.

Please sign one of the following statements:

Booth Library of Eastern Illinois University has my permission to lend my thesis to a reputable college or university for the purpose of copying it for inclusion in that institution's library or research holdings.

Date

I respectfully request Booth Library of Eastern Illinois University not

allow my thesis be reproduced because  ·��-

Date Author

m A KEY TO THE HEPA TICAE OF

- COLES AND CLARK COUNTI ES , ILLINOIS (TITLE)

BY

Michael J. Klopmeyer

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

IfI 3;: JE �I tfI 30 / cYI DATE A KEY TO THE HEPATICAE OF COLES AND CLARK COUNTIES, ILLINOIS

BY

MICHAEL J. KLOPMEYER

B.S. in Botany and Zoology, Eastern Illinois University, 1979

ABSTRACT OF A THESIS

Submitted in partial fulfillment of the requirements for the degree of Master of Science at the Graduate School of Eastern Illinois University

CHARLESTON , ILLINOIS 1981

40386? This study represents a taxonomic survey of the hepati­ cae of east-central Illinois . The counties included in this study were Coles and Clark, located in the eas t- central por­ tion af the state , near the Indiana stateline . Of the areas included in this study, two are in Clark county, Rocky Branch and Rocky Hollow, and one in Coles county, Rock's Park .

The influence of the Wisconsin glacier, and its ter­ minal moraine located in southern Coles county, greatly in­ fluenced the topography and ultimately the specific habitats for the hepaticae . The majority of the liverworts encoun­ tered in these counties are found in the numerous sandstone outcrops along rivers and streams . These outcrops were ex­ posed due to the cutting action of the glacial melt water .

A key was composed from the combination of the keys present in current literature . Illustrations and descrip­ tions are provided for each of the genera and species lo­ cated in these two counties . In some instances , specific locations within these counties are included for any of the species considered rare or endangered .

A total of forty-seven liverworts have been collected in these counties. Four are considered to be rare or en­ dangered . They are pul cherrimum (G .Web.) Hampe ,

Blepharostoma trichophyl lum (L.) Dum., tomen- tella (Ehrh .) Durn. , and Marchantia po lymo rph a L. The sur­ vival of these , and liverworts in general , as well as other rare , is dependent upon the preservation of these areas . TABLE OF CONTENTS

ACKNOWLEDGEMENTS ...... iii

LIST OF ILLUSTRA.TIONS I e • t I e • e I • I t I I I I I I I I I I I I I I I I I I I I iv

INTRODUCTION ...... 1

DESCRIPTION OF STUDY AREAS • . . • • ...... • ...... • • . • • • • • J

CHECKLIST OF THE HEPATICAE OF EAST-CENTRAL ILLINOIS ... 11

A KEY TO THE HEPATICAE OF EAST-CENTRAL ILLINOIS .••••. 14

GLOSSARY •••••••.••••••••••••••, • • . • • . • • . • • • • • • • • • • • • • 68

...... 71 CONCLUSION .

LITERATURE CITED ...... • . . . • ...... • • . 72

ii ACKNOWLEDGEMENTS

I would like to thank my advisor and friend; Dr .

Charles B. Arzeni , for his encouragement and advice

throughout my graduate studies . I would also like to

thank Dr . Wesley Whi teside and Dr . Roger Darding for

their review and helpful criticism of the manus cript.

I am also indebted to my parents for their patience and

support throughout my graduate work .

iii LIST OF ILLUSTRATIONS

1. tenella (L. ) P.Beauv ••••••••••••••••••. 32

2. trilobata (L. )S.Gray •••••.••••••••••••• 43

3. Blasia pus illa L .••.••••••.••••••••••••••••••••• 34

4. trichophyl lum (L.) Dum • ••••••••••• 35

5. Calypoge ,ja muelleriana ( S chiftn . ) K.Mull . • .•••••• 44

6. connivens (Dicks .) Lindb . ••.••••••••• 57

7. C • 1 unul i fo 1 i a ( D um • ) D um • • • • • • • • • • • • • • • • • • • • • • . 58

8. Cololeje unea biddlecomiae (Aust.) Evans .•••••••• 38

9. Conocephalum conicum (L.) Lindb • ••.••••••••.•••. 28

10. Dipl ophyl lum api culatum (Evans ) Steph . .•••.•• ••• 54

11. Frullania bolanderi Aust • ••••••••••••.•••.•••••• 62

12. F . infla ta Gott ...... 64

13. F. riparia ( Reinw. et al .) Dum . • .••••• •••••••••• 66

14. F. squarrosa (Reinw. et al .) Dum • ••••••••••••••• 63

15. F. tamarisci (L.) Dum . var . asagrayana (Mont.) ••• 61

16. gr aveolans (Schrad.) Nees • •.••••••••••• 59

17. (Web . & Mohr .)Spruce •••••••. 67

18. autumnalis Aust • .•••••••••••••••••• 47

19. crenuliformis Aust • .•••••••••••.••• 50

20 . J. gr acillima Sm ...... 49

21 . J. hyalina Lyell ...... 51

22 . J. leiantha Grolle ...... 48

iv 23 . cavifolia (Ehrh .) Lindb . ••••••••••••••• 41 . 24 . Lepi dozia reptans ( L.) Durn. • ••••••••· • ••••••••••• 42

25 . heterophyl la (Schrad .) Durn. e e I • I I I I I I 60

26 . bicrenata (Schmid . ex Hoff m.) Durn• ••••• 53

27 . L. incisa (Schrad .) Durn• •••..••.•....•.••••.•••• 52

28. fragrans (Balbis) Frye & Clark ••••••••••. 31

29 . Marchantia polymorph a L. • ... ..••..•.•...... •. 27

JO. Notothylas orbicularis (Schwein.) Sull . •••.•••, • 23

31 . curvifolia (Dicks .) Mitt. . ••••••••••••• 56

32 . denudaturn (Sw.) Trev • ••.•••••••••. 45

•••••••••. . • ...•.•••• 33. epiphyl la (L.) Corda . 33

J4 . Phaeoceros laevis (L. ) Prosk. . ••••••••••• , •••••• 22

35° asplenioides ( L,) Dum. • ••••••••••••• 46

36 . Perella platyphylloidea (Schwein .) Lindb • ••••..• 40

37 . Preissia quadrata (Scop.) Nees . • • • • • • • • • • • • • • • • • 29

38. Ptilidium pulcherrimurn (G .Web.) Hampe .•••••••••• 37

39. Radula complanata (L.) Durn...... 39

40 . hemisphaerica (L.) Raddi ...... •, . . . . JO

41 . Riccia fluitans L . •..•...... •.•....•••••• 24

42 . R. frostii Austo ...... 25

43 . Riccio carpus na tans (L.) Corda ••..· ....••.•••.••. 26

44. Scapa nia nemorea(L. ) Grolle .•••••••••••••••••••. 55

45 . (Ehrh .) Durn• ••••••.•••••. J6

v 1

INTRODUCTION

The purpose of this paper is to present a reasonably simplified key to the Hepaticae of east-centra� Illinois.

The liverworts are usually an overlooked entity of the due to their small size, lack of common names, dif­ ficulty in determining their correct identification, and absence of any economic importance . Therefore , the keys in th is paper have been greatly simplified by the elimina­ tion of many technical terms and descriptions to help alle­ viate the problems of confusion and frustration encountered in most keys . The illus trations are of the basic morpho­ logical characteristics of each of the species , sufficient enough to separate them from each other . These illustrations were mainly reproduced from Schuster (1953, 1966) and Steere

(1964) by use of an Arto-Graph drawing machine . The keys , with the exception of the key to the Frullania, which is based on Evans (1897) , are derived from the combined sources of the keys present in current literature� The combination was prepared in order to simplify the keying by differentiating the morphological characteristics of the plants through the use of a hand lens or a dissecting stereo microscope . The nomenclature is based on the current class­ ification by Stotlar (1977) . Some liverworts , which have 2 been included in this paper, are extremely rare or, are apparently thought to be absent in east-central Illinois .

These liverworts are fairly common in the Smoky Mountains and the northern regions of the United States . Therefore, this key can be utilized for plants collected in those areas .

It is hoped that th is paper can be used by beginning bry­ ologists to aid them in identifying the liverworts to their respe ctive genera, then consulting more complete references for specific identification . 3

DESCRIPTION OF THE STUD Y AREAS

This study was conducted in Coles and Clark counties , in east- central Illinois (Fig. 1) . These counties are bor­ dered to the north by Douglas and Edgar counties ; west by

Moultrie and Cumberland counties ; south by Crawford county; and , east by the state of Indiana . Coles county covers 509 square miles ·and Clark county 504 square miles . Farmland makes up over 90% of the to tal area of these counties.

The most important geological period in the history of

Coles and Clark counties was the glacial . At least two gla­

cial advances , the Illinoian and early Wisconsin, influenced

the topography and ultimately the specific habitats suitable

for hepatic development. The glacial debris , known as till ,

deposited by the Illinoian and Wisconsin ice sheets, consti­

tute the soils of these counties .

The primary effect of the Illinoian ice sheet was in

acting as a leveling force , rubbing down the preglacial hills

and filling the preglacial valleys . The Wisconsin ice sheet

then followed the Illinoian, and filled up the valleys formed

by the melting of the Illinoian . The Wisconsin glacier reached

its southernmo st advancement in the Coles and Clark counties

of Illinois . This te rminal moraine varies from 5 to 9 miles

in width and extends across the southern part of Coles county 4

1. Coles 2. Clark

1 2

Figure 1. The location of Coles and Clark counties in Illi­ nois. 5

to th,e northwest portion of Clark county . The northern boundary of this moraine is indistinct, while the southern boundary is marked by an abrupt drop-off . The outwash plain

is an area which is composed of the material carried by the glacier and deposited in front of it by the melting water

of the glacier . This plain forms a belt some four or five miles in width and borders the moraine on the south.

The areas of the rolling hill terminal moraine and the

outwash plain are the most interesting, in regarding to fre­ quency and variety of hepatics in eas t-central Illinois.

The outw ash plain was significantly altered as time passed by the influx of water from the retreating Wisconsin glacier.

This continuously flowing water over the years cut through

the glacial till and exposed the bedrock surfaces which are

sands tone outcrops in th ese areas today .

The exposed bedrock , or sandstone , can be found in dif­

ferBnt areas of Coles and Clark counties . The mo st produc­

tive areas in the morainal belt are along large rivers and

streams , and in the outwash plains , in scattered locations

along small streams . Sandstone , and the glacial till asso­

ciated with it, are the best habitats for the hepatics .

An excellent location in Coles county for hepatics is

an area along the Embarrass River about five miles eas t of

Charleston. This area (Fig.2) , commonly known as Rock 's

Park (Sec 17 T12N R10E) , contains numerous sandstone out­

croppings along the river . It was once part of the terminal 6

•

• Rock 's Park

• Rocky Branch

.& Rocky Hollow

� Moraine

Figure 2. The study areas and extent of the Wisconsin ter­ minal moraine in Coles and Clark. counties , Illinois . 7 moraine , and the cutting effect of the Embarrass River, due to glacial melt waters , slowly exposed the rock outcrops .

One of the most unique and rare liverworts found in

Rock's Park is (Dicks .) Mitt . . This small, leafy liverwor ·t is usually found on decor ti cated coniferous logs in the northern regions of th� United States .

However, it occurs here on a shaded, south-facing sandstone ledge . It is, perhaps , the on ly location for this hepatic in east-central Illinois . Another leatyliverwort of north­ ern regions , Frullania bolanderi Aust., is found in the mixed mesophytic woods above the sandstone ledges, on the bark of mature white oaks (Quercus alba L.).

An area directly across the Embarrass River from Rock's

Park also contains numerous sandstone bluffs . Jungermannia leiantha Grolle , another rare leafy liverwort of east-central

Illinois, can be found in scattered locations on shaded soil at the base of these sandstone outcrops .

Clark county is . perhaps, the best county in east­ central Illinois for the liverworts . Rocky Branch Nature

Preserve (Sec 29 T12N R12W) , an excellent study area (Fig.2) , contains many sandstone cliffs and ledges . These outcrops were exposed by the erosive action of the glacial mel.t water .

However, Rocky Branch , as well as most of Clark county, is south of the terminal moraine and is mostly considered the outwash plain of the Wisconsin glacier . Therefore, there are dramatic sandstone outcrops here , due to the longer 8 effects of exposure to erosion than the terminal moraine area of Coles county .

There are four liverworts at Rocky Branch which are considered rare or endangered in east-central Illinois .

The moist sandstone inhabitors , Bleph arostoma trichophyl lum

(L. ) Dum., and Lophozia incisa (Schrad .) Dum . are found scattered on the south sandstone wall of Rocky Branch Creek .

Trichocolea tomentella (Ehrh .) Dum., a common inhabitant of the northern United States, can still be found in one loca­ tion along the south side of Rocky Branch Creek at the base of a sandstone wall . Because of indiscriminate ·collecting , this showy liverwort may soon disappear from the Illinois flora. Another northern-inhabiting leafy liverwort, Ptili­ dium pulcherrimum (G.Web.) Hampe can be found on a dry, ex­ posed sandstone boulder next to Big Creek, near the mouth of Rocky Branch Cr eek . Near this site on dry exposed sand­ stone is another uncommon thallose liverwort of east-central

Illinois, Mannia fragrans (Balbis) Fry & Clark .

Another field area included in this study is located in the outwash plain of the southern portion of Clark county .

Rocky Hollow (Sec 5 T9N R12W ) (Fig .2) is privately owned and consists of a narrow, d·eep sandstone gorge dissecting rolling farmland . This area was also formed by glacial water erosion over a period of time , cutting through the glacial till and exposing the sands tone bedrock. The steep sands tone cliffs and narrow width of the gorge offer a unique habitat for 9 the .

At Rocky Hollow, as well as at Rocky Branch, there are northern-inhab iting liverworts in various substrata . Bleph ­ arostoma trichophyl lum (L.) Dum. is found scattered on the sands tone walls and Trichocolea tomentella (Ehrh .) Dum. can be found in several places on sandy soil along the small stream in the gorge . Unlike the sandstone substrate at Rocky

Branch, Ptilidium pulcherrimum (G.Web.) Hampe is seen occa­ sionally on the decorticated logs of the eastern red cedar

(Junipe rus virginiana L.). Marchantia polymo rph a L., a thal­ lose liverwort, is found in one location on �andy soil along a small stream . This is now the only known location for this liverwort in east-central Illinois .

The presence of northern-inhabiting liverworts in east­ central Illinois can be possibly linked to the effects of the last glaciation . The composition of the vascular plants of Rocky Branch and Rocky Hollow are similar to the liver­ worts in that they, too, contain several species of the hem­ lock-hardwood forests of the Great Lakes region . The Amer­ ican beech, Fagu s gr andifolia (L.) Bart .; partridge berry,

Mitchella repens L.; beech drops , Epi fagu s virginiana (L.)

Bart.; and strawberry bush , Euonymu s obovatus Nutt., are a few o� the higher plants in these areas which are beyond their normal distributional ranges.

There are several theories which attempt to explain the presence of these disjunc.t plants in east-central Illinois . 10

The presence of the glaciers here for hundreds of years could have produced a suitable northern climate conducive to the growth of these plants . After the glacier retreated, the climate changed and mo st of the plants did too . How­ ever with the glacial till acting as a direct influence on the soil and ultimately the substrata of these· plants , and the protected niches provided by sandstone outcrops , these disjunct plants survived and persist today. 11

A CHECKLIST OF THE HEPATICAE OF EAST-CENTRAL ILLINOIS

Family

Blepharostoma trichophyl lum (L.) Dum .

Family Ptilidiaceae

Ptilidium pulcherrimum (G.Web.) Hampe

Family

Trichocolea tomentella (Ehrh .) Dum.

Family

Bazzania trilobata (L.) S. Gray

Lepidozia reptans (L. ) Dum .

Family Calypogejaceae

Calypogeja muelleriana (Schiffn.)K. Mull . subsp .muelleriana

Family

Cephalozia connivens (Dicks .) Lindb .

C. lunulifolia (Dum.) Dum .

Nowellia curvifolia (Dicks .) Mitt.

Family

Odontoschisma denudatum (Sw.) Trev .

Family

Jamesoniella autumnalis (DC .) Steph .

Jungcrmannia crenul iformis Au8t .

J. gr acillima Sm .

J. hyalina Lyell 12

J. leiantha Grolle

J. purnila With.

Lophozia bicrenata (Schmid . ex Hoffm .) Durn.

L. incisa (Schrad.) Durn.

Family Geocalyaceae

Geocalyx gr aveolans (Schrad .) Nees .

Harpanthus scutatus (Web .& Mohr .) Spruce

Family

Lophocolea heterophyl la (Schrad .) Durn.

Family

Plagiochila asplenioides (L .) Durn.

Family

Dipl ophyl lurn api culatum (E vans) Steph .

Scapania nemorea (L .) Grolle

Family Porellaceae

Farella platyphylloidea (Schwein .) Lindb .

Family Radulaceae

Radula compl ana ta (L.) Durn.

Family Jubulaceae

Frullania bolanderi Aust.

F. eboracensis Gott.

F. inflata Gott.

F. squ arrosa (R einw . et al .) Durn.

F. tamarisci (L .) Durn. var . asagrayana (Mont.) Hatt .

Family Lejeuneaceae

Cololejeunea biddlecomiae (Aust.) Evans 13

Lejeunea cavifolia (Ehrh .) Lindb . emend . Buch .

Family

Pellia epiphyl la (L.) Corda

Family Blasiaceae

Blasia pus illa L.

Family

Riccardia pi ngu is (L.) Gray .

Family Aytoniaceae

Reboulia hemisph aerica (L. ) Raddi

Asterella tenella (L.) P. Beauv .

Mannia fragrans (Balbis) Frye& Clark

Family Conocephalaceae

Conoceph alum conicum ( L .) Lindb .

Family

Marchantia po lymorph a L.

Preissia quadrata (Scop.) Nees

Family Ricciaceae

Riccia fluitans L.

R. frostii Aust.

Ricciocarpu s natans (L.) Corda

Family Anthocerotaceae

Noto thylas orbiculatus (Schwein .) Sull .

Phaeoceros laevis (L.) Prosk. 14

A KEY TO THE HEPATICAE OF EAST-CENTRAL ILLINOIS

I. Plants scale-like or ribbon-like or in rosettes; prostrate; not differentiated into stems or .

II . Plants with stems and leaves (leaves may be thread­ like); erect or prostrate .

I. Thalloid Liverworts

1. Thallus green, with many chloroplasts per cell .

...... 3

1. Thallus blue-green, with one chloroplast per cell; horn-like , erect or prostrate.

...... 2

2. Thallus large; horn-like sporphyte erect and most obvious part of plant; mo ist sandstone .

• • .•••••.•• ••••..•• .•••••..•••• Phae oceros Page 22

2. Thallus small ; horn-like sporopnyte small and pro­

strate ; disturbed sandy to clayey soil; rare •

• • • • • • . • . • • • • , •••.•••.•••.••••. . Notothylas Page 23

3. Thallus with air pores present; dichotomously branched, air-filled thallus ; rhizoids pegged...... 4

3. Thallus with no air pores ; no internal differenti­ ation; rhizoids smooth ...... 11

4. Sporophyte imbedded in thallus; aquatic, or we t mud ; thallus from 1 to 8 mm wide .

I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I a a I I I I I I I I I I I I I I 5

4. Sporophyte elevated above thallus; thalli more than 8 mm wide; seldom aquatic.

...... 6 1.5

.5· Thallus thin, ribbon-like , dichotomously forked; no t more than 1 mm wide, floating or on wet mud • ••• •••••••... ..•. •••• ••••••• •••Ric cia Pages 24,2.5

.5· Thallus rosette-shaped ; pore on upper surface very small ; purple scales below ; floating or on wet mud . •••• ••••• ••••••••••••••••••••Ricciocarpus Page 26

6. Thallus dark green, midrib conspicuous ; gemma cups present; rare •

• • • • • • • • • • • . . • . . . • • • • • • • • • • • • • • Marchan tia Page 27

6. Thallus without gemma cups ; varying colors , usually bright green ...... 7

7. Thallus large , usually bright green ; aromatic when crushed ; conic-shaped archegonoiphores; common •

• • • • • • • . • • • • • • • • • • • • • • • • • • • • • Conoceph alum Page 28

Thallus not aromatic when crushed; conic-shaped arch­ egoniophores absent .

I • I I I I I I I I I I I I I • I I I I I I I I I I I I t t t t I t t I I I I I I I I • I I I I I 8

8. Air pores easily seen, circular, on scal­ loped umbrellas ; margin of thallus purplish to red­ dish; rare ...... Preissia Page 29

8. Air pores small , visible with hand lens ; moist sand­ stone or disturbed soil .

I I I I I t t I t t t t t t t t t I t t t t t t I t t t t t t t t t I I t t I t I I I I I I I I I 9

9. Air pores surrounded by 4 or 5 concentric rows of cells ; margin of thallus usually purple, especially below ; sandstone • ...... Reboulia Page 30

9. Air pores surrounded by less than 4 or 5 concentric rows of cells ; sandstone or disturbed soil ...... 10 16

10. Air pores surrounded by 2 or J concentric rows of cells ; margin of thallus purple, usually curled up­

wards ; rare on sandstone •

• • • • • ...... • . • • . • . . • • Mannia Page 31

10. Air pores surrounded by irregular , thin-walled cells ; thallus narrow with margins sometimes purple ; dis­

turbed soil in pastures and fields ; rare •

• • • • • . . . . • ...... • . • • • • • • • As terella Page 32

11. Thallus dark green but transluscent, turning dark umn purple in aut ; moist sandstone ; common • ...... , ...... Pellia Page 33

11 . Thallus pale green; transluscent; dark spots of Nostoc colonies visible imbedded in the thallus ;

moist sandstone • ...... Blasia Page 34 17

II. Leafy Liverworts

1. Leaves divided into filiform segments or threads .

...... 2

1. Leaves undivided , entire or lobed .

e I • I I • • I • • I e I e I • • • I e I • I I e I I • • • I e I • I I I I I I I I I I I I I I I 4

2. Leaves consisting of 3 to 4 filamentous threads , uniseriate ; minute , delicate plants resembling an

alga, sandstone •

•..•.•...•...... •••••Ble ph ar ostoma Page 35

2. L�aves consisting of 3 to 4 lobes of 4-20 cells wide near base ; lobes with marginal cilia; plant 2-3 mm wide .

e I I I I • I I • I I I I I I I I I I I I I I I I I • I I I I I I I I I I I I I I I I I I I I I I 3

3. Plants large , whitish-green, plumose ; leaves divided into many filiform divisions , base 1-4 cells wide ; mo ist stream banks .

•.•.••.•...... •. .••••••Trichocolea Page 36

3. Plants small , reddish-brown ; leaf base consisting of 2 major lobes , 6-20 cells wide ; rare on sandstone and rotten coniferous wood • ...... Ptilidium Page 37

4. Leaves in dorsal view , distinctly incubous .

• I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I f I I I I I I I I I I I I I I I 5

4. Leaves in dorsal view, distinctly succubous .

...... 12

5. Leaves with small ventral lobule , hidden under large dorsal lobe ( complicate-bilobed) .

I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I 6

5, Leaves lacking a ventral lobule , simple , plane ; not complicate-bilobed .

...... 10 18

6. Underleaves absent; plants yellow green .

• • e e e e • e e e I e e e I I I I I • I e t I e • e • I • e e e e e e I t t e t I t t I I a I t 7

6. Underleaves present, large and obvious ; plants dark green to brown or black .

...... 8

7. Plants minute , less than .05 mm wide ; leaf cells unipapillose ; rare on rotten wood .

• . . . . • ...... • . • . • • . • • • • • • • • • Colole,j eunea. Page 38

7. Plants larger, 1-2 mm wide ; light green leaves flattened upon each other ; perianth distinctly

flattened ; bark at base of trees • ...... Radula Page 39

8. Underleaves entire, similar to lobules , lobules tongue-shaped, parallel to stem; rare .

. . •• •. . •...... •. .. •. . .• ••••.•••.••Perella Page 40

8. Underleaves bilobed, different shape than lobules ; small plant.

I I I I I I t I t t t t t I I I I I I I I I I I I I t I t t I I t t I I I I I t t I I t I I I I I 9

9. Lobules not inflated; very rare on rotten wood .

• ...... • . . Le,jeunea Page 41

9. Lobules very concave , resembling an inverted cup ; small, dark green to brownish-blackish plants on bark of trees and sometimes sands tone .

• • . • • . . . • • • • . • ...... • • • • • • • • • • . Frullania Page 21

10 . Leaves entire or slightly notched .

...... 11

10 . Leaves divided halfway down into 3-5 lobes ; rare .

••...... •. •. •. .•••••••••••Lep idozia Page 42

11. Leaves divided into 3 teeth or notches ; rare on sandstone ...... Bazzania Page 43

11. Leaves entire or notched, pale , watery green .

• • . . • . . . . . • ...... • . . . • • . • • • • • • CalyPoge ,ja Page 44 19

12 . Leaves entire, not at all lobed or toothed.

...... 13

12. Leaves 2 or more lobed or toothed .

I t t t t t t t t t t t t t t t t t t t t t t I f I t t t t t t t t t t t t t t t t t t t t t t t 17

13. Leaf cells with large, bulging, knot-like trigones; lower leaves circular , rigid , tending to stand on

edge facing one another • ••••••..•....•.....•.••.••••Odontoschi sma .Page 45

13. Leaves with small to moderate trigones, ·or without .

...... 14

14, Leaf cells with trigones.

...... 15

14. Leaf cells lacking trigones; large , leaves erect­ ascending, decurrent; marginal teeth few or absent .

•••.• .....•.• •...••..: ..•..•••Pla giochila Page 46

15. Mouth of perianth contracted , fringed with hairs ; leaves longer than wide .

• • . . . • ...... • • . . . • • • • • • • • • Jamesoniella Page 47

15. Mouth of perianth pleated, perianth truncate .

t t t t t t I t t t t t t t t t t t t t t t t f t t t t t t t t t t t t t t t t t I t t t t t t f 16

16 . Leaves rectangular ; trigones large , bulging .

•••.••..•..•..••••••Jun ge rmannia leiantha Page 48

16 . Leaves circular ; trigones small, rarely bulging •

• . • • . • . . • . . . . • . . • • • • • • • • •Jun germannia Pages 49-51

17. Leaves all or mostly 3-4 lobed, lettuce-like ; blue­

green or green ; rare on sandstone or disturbed soil •

• . ••.•••..• ...... •.. ..••••Lop hozia Pages 52 ,53

17. Leaves all bilobed; not 3-4 lobed .

...... 18

18. Leaves complicate-bilobed, dorsal lobe smaller than ventral lobe .

t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t t I I I t t 1 9 20

18. Leaves simple , plane; no t complicate-bilobed.

• • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 20

19. Ventral lobes lingulate; dorsal lobes similar in shape, entire margin; gemmae polygonal ; sandstone .

• • • . . . . . • . . . . . • • ...... • • . • • • • D ipl ophyl lum Page 54

19. Ventral lobes not lingulate, margins finely serrate; ovoid, rust-brown gemmae at apices; sandstone ...... Page 55

20 . Leaf lobes extending into long slender cilia, 4- 8 cells long; usually reddish; rare oh sandstone

or rotten coniferous logs •

...... · ...... N owe llia Page 56

20. Leaf lobes acute to acuminate , not ending in cilia­ like apices ...... 21

21. Underleaves absent or rudimentary; leaf cells and cortical cells large and pellucid, lacking distinct

oil bodies •

• • • . • • • . • ...... • • • • • • • . Cephalozia Pages 57, 58

21. Underleaves present, obvious .

...... 22

22 . Underleaves bilobed to t their length , margins entire ; leaves evenly bilobed; greasy yellow­

green in color • ...... Geocalyx Page 59

22. Underleaves bilobed to i their length , conspicuous

tooth on each side; common on decorticated logs •

• • • . • . . • . . • ...... • • . • . • . • • • • . • Lophocolea Page 60 21

III . A Key to the Genus Frullania

1. Dorsal lobe with a faint midrib of dark cells •

•••••••••••••F. tamarisci var . asagraya na Page 61

1. Dorsal lobe· without a midrib ...... 2

2. Lobes not cordate at base, leaf cells in middle of lobe with inconspicuous trigones ; branches often ending in erect leafless flagella .

• • • • • . • . • . • . • . • . . • • . . • • • • • • • • F. bolanderi Page 62

2. Lobes cordate at base; leaf cells in middle of lobe with conspicuous trigones.

...... 3

3. Leaves distinctly squarrose when moist •

• • • • . . • . • . • . • • . • . . . . . • • • • • • • • F. sq uarrosa Page 63

3. Leaves scarcely or no t at all squarrose when moist.

...... 4

4. Dorsal lobes of mature leaves truncate at base; intermediate thickenings between cells absent;

lobules often flat • ...... F . infla ta Page 64

4. Intermediate thickenings between cells present; lobules inflated or flat.

...... 5

5. Lobules inflated throughout; cells with numerous intermediate thickenings ; underleaves entire or 1-toothed on each side .

•••. .•...... ••. . .••.•••F. eboracensis Page 65

5. Lobules flattened ; green to red; on sandstone •

••••••..•.. .••••••••..•••••••.•F. riparia Page 66 22

Phaeoceros laevis (L.) Prosk., 5x

Phaeoceros laevis (L.) Prosk .

Thallus dark-green to blue-green , nearly circular, with shallowly divided lobes; sporophyte erect, long , cylindric, splitting longitudinally at maturity, into two equal parts , thus allowing for yellow to escape . Easily seen , when fruiting, on moist sandstone , but in sterile condition the dark blue-green , tightly appressed thallus is its · distin­ guishing characteris tic . 23

Notothylas orbicularis (Schwein.) Sull ., 10x

Notothylas orbicularis (Schwein .) Sull .

Thallus a small , green rosette , 1 cm in diame ter , sporo­ phytes prostrate , 1 mm long, difficult to see without hand

lens ; on disturbed , clayey to sandy soil. 24

Riccia fluitans L., Jx

Riccia flui tans L.

Thallus narrow , ribbon-like , 0.5-1 .0 mm wide ; dichotomous­ ly branched ; floating aquatic form or terrestrial form on moist soil of river banks ; not common .. 25

Riccia frostii Aust., 14x

Riccia frostii Aust.

Thallus dark-green to grayish-green or grayish-crystalline, in compact rosettes , 4-15 mm in diameter ; or, thalli narrow ,

0.5-1 .5 mm in diameter , linear , elongate , lying close and parallel to each other ; upper surface obscurely re ticulate when young, soon becoming spongiose when mature; when fruit­ ing in late summer or fall , the dorsal surface becomes pit­ ted with numerous black , coarsely granular , appearing masses of spores ; not common on clayey soil in fields or along riverbanks. 26

Riccio carpus na tans (L.) Corda ., 2x

Ricciocarpus natans (L.) Corda .

Thallus light green , rosette-shaped , with dichotomously branched furrowed lobes , 2-4 cm wide; ventral surface with

large, numerous sword-like purple scales ; rare on mo ist soil . 27

Marchantia po lymo rph a L., 2x

Marchantia po lymo rph a L.

Thallus dark green, dichotomously branched , with broad , dark midrib ; pores difficult to see without a hand lens ; disco id gemmae cups usually present; umbrella-shaped recep­ tacles; rare on moist shaded stream banks 28

Conoceph alum conicum (L. ) Dumort. , 2x

Conoceph alum conicum (L .) Dumort.

Thallus large , uniformly bright green, dichotomously branched ; dorsal surface with distinct diamond-shaped markings , each with a conspicuous air pore ; conic-shaped archegoniophores; crushed thallus strongly aromatic; common on moist sandstone . 29

Preissia quadrata (Scop.) Nees , Jx; pore ,225x

Preissia quadrata (Scop .) Nees

Thallus dull green, lateral margins somewhat purplish ; ventral scales always blackish-purple ; pores prominent, whitish, elevated and compound ; margins of older thalli are usually bleached and decolorated ; this characteristic, along with the elevated pores, distinguish th is plant from the very similar , but inconspicuously pored , Reboulia hemi­ sphaerica. JO

Reboulia hemisph aerica (L.) Raddi, 2.5x; pore, 240x

Reboulia hemisphaerica (L .) Raddi

Thallus yellow-green, smooth , margins of thallus purplish or brownish-purple , underside purpllsh-black ; upper cells of thallus with distinct, bulging trigones; pores small, in- conspicuous , surrounded by 4-5 concentric rows of cells ; rare on soil or sandstone. 31

Mannia fragrans (Balbis) Frye and Clark, 6x ; pore 295x

Mannia fragrans (Balbis) Frye and Clark

Thallus grayish-green; dorsal surface of rough texture, due to protuberant pores; thallus linear-lingulate , sparsely forked; margins purplish, during dry periods, thallus edges become upcurved and the blackish-purple underside becomes

exposed ; not common on exposed sandstone . 32

Asterella tenella (L.) P. Beauv., 5x; pore, 225x

Asterella tenella (L.) P. Beauv,

Thallus bright green, often purplish along margins and beneath, 0.5-1 .5 cm long and 1.5-3 mm wide and dichotomous­ ly branched ; cells around the pore irregular and thin-walled ; rare in pastures or along edges of old roads . 33

Pellia epiphyl la (L.) Corda, 6x

Pellia epiphyl la (L.) Corda

Thallus large , flat, thin, dull dark-green, often purplish or maroon in the fall ; fruits prof�sely in the spring, pro­ ducing long , pellucid archegoniophores ; common , forming large extensive mats on mo ist soil-covered sandstone . 34

Blasia pusilla L., 5x.; gemmae, 100x

Blasia pusilla L.

Thallus green or yellowish-green with scalloped margins, near the base of which occurs scattered dark swellings which are cavities filled with the blue-green alga, No stoc ; growth is usually in dis tinct rosettes , or sometimes branched , and the midrib is distinctly indicated by a whitish line; star­ shaped gemmae produced in bottle-shaped flasks often present on surface; found on clayey to sandy stream banks . 35

leaf

underleaf

Blepharos toma trichophyl lum (L.) Dum., 50x

Bleph arostoma trichophyl lum (L.) Dum.

Plants minute , resembling an alga ; leaves dissected to the base into 3-4 thread-like divisions , that are uniseriate throughout; rare on mo ist, shaded sandstone at Rocky Branch and Rocky Hollow. J6

leaf underleaf

Trichocolea tomentella (Ehrh .) Dum., 4ox

Trichocolea tomentella (Ehrh .) Dum .

Plants large , showy, pinnately frondose ; leaves densely ciliate nearly to the base ; on mo ist, shaded , sandy soil at edge of stream at Rocky Branch and Rocky Hollow . 37

Ptilidium pu lcherrimum (G.Web .) Vainio , 20x ; leaf ,40x

Ptilidium pulcherrimum (G. Web .) Vainio

Plants rather small to moderate in size; green to yellow­ ish or reddish-brown ; leaves ciliate at the margins , but clearly consisting of 2 or more lobes ; rare on exposed sand­ stone or decorticated Junipe rus logs at Rocky Branch and Rocky

Hollow . 38

Dorsal view , 4ox

Ventral view , 70x

leaf, 80x

Cololejeunea biddlecomiae (Aust.) K. Muell .

Plants minute , less than 0.5 mm wide ; leaf cells distinct­ ly unipapillose ; rare on moist, coniferous wood (Juniperus ) at Rocky Hollow . 39

Radula complanata (L.) Dum. , dorsal view , 25x; ventral view , with perianth , 25x

Radula complanata (L.) Dum .

Plants rather small, bright yellow-green ; leaves strongly flattened against the substrate ; oil bodies very large , brown,

1 ( rarely 2) per cell ; gemmae often produced at the leaf mar- gins ; perianths common, strongly flattened and spade-shaped ; not common at base of trees on bark . 40

Perella pl atyphylloidea (Schwein .) Lindb ., dorsal view, 4x; ventral view, 18x

Perella pl atyphylloidea (Schwein .) Lindb .

Plants dark green ; lobules broader than the stems , de- current, about as broad as the underleaves ; rare on shaded rock or at the base of trees on soil. 41

Lejeunea cavifolia (Ehrh .) Lindb ., dorsal view with per­ ianth , 65x; ventral view, 65x

Lejeunea cavifolia (Ehrh .) Lindb .

Plants pale to yellowish-green ; leaves deeply notched, and underleaves are nearly circular ; lobules small and inconspicuous ; rare on rotten wood . 42

Lepidozia reptans (L.) Dum., 25x

Lepidozia reptans (L.) Dum .

Plants small , dull-green, pinnately branched ; leaves small , deeply divided into 3-4 finger-like lobes ; under­ leaves similar , but smaller; rare on rotten wood . (L.) Gray , dorsal view , 10x; ventral view , 10x

Bazzania trilobata (L.) Gray

Plants large , rigid , dark-green, dichotomously branched, with leaves rectangular , 3-lobed across a broad apex; very rare on sandstone banks . 44

Calypoge ja muelleriana (Schiffn.) K. Mull . subsp. muelleriana dorsal view, 15x; ventral view , 15x

CalYPoge ja muelleriana (Schiffn.) K. Mull . subsp. muelleriana

Plants pale , watery in color ; leaves broadly ovate , bluntly obtuse, or rounded at the apex ; underleaves about as broad as long, bilobed to i their lengths ; on moist humus or sandstone (C. trichomani s, wi th blue oil bodies, has no t been collected in this area). Odontoschisma denudatum (Nees . in Mart.) Dum., 18x-

Odontoschisma denudatum (Nees in Mart,) Dum.

Plants green, stems ascending at tips and bearing terminal clusters of yellow gemmae ; leaves smaller at some distance below the tip, rounded , entire ; trigones bulging ; oil bodies very large , 1-4 per cell; rare on wet humus or logs . 46

Plagiochila asplenioides (L. ) Dum., 10x

Plagiochila asplenioides (L.) Dum .

One of the largest leafy liverworts in this area; plants erect-ascending; leaves rounded and dentate , but entire in this area, decurrent; common along streq.m banks . 47

Jamesoniella autumnalis (D .C.) Steph.,10x; perianth , 10x

Jame soniella autumnal is (D .C.) Steph.

Plants green; leaves rounded-quadrate to shortly oblong; cells thin-walled, with small to bulging trigones ; the distinctive characteristic of th is plant, is the perianth , which is tapere� and plicate to the apex and fringed at the mouth ; on soil-covered sandstone . 48

Jungermannia leiantha Grolle , 10x

Jungermannia leiantha Grolle (J. lanceolata L.)

Plants dark green ; leaves rounded-oblong, longer than broad ; leaf cells irregularly roughened , distinctly tri­ gonous ; perianths smooth cylindric, abruptly truncate , shortly beaked from a central depression ; uncommon on moist, shaded banks . Jungermannia gr acillima Sm . (= Plectocolea crenulata (Smith) Evans) , 17x

... �. . Plants pale , pellucid , whitish-green, upper portions of leaves dull red ; prostrate , closely appressed to substrate , irregularly but freely branching ; leaves orbicular , narrowed at base, marginal cells of leaves 2 to 3 times the inner cell size, forming an obvious border ; rhizoids colorless; in diff- erentiating this species from J. crenuliformis, the border of thick-walled cells is more ·sharply developed than J. cren- uliformis , also , J. crenuliformis has purplish rhizoids , is greener, no t whitish-green, and also tends to develop a pur- plish pigmentation at the leaf base rather than a reddish tint in the leaf apices of J. gr acillima; J. crenuliformis also possesses larger and more conspicuous oil bodies in the leaf cells and marginal cells ; not common on the moist soil of sandstone banks . 50

Jungermannia crenuliformis Aust. , 10x

Jungermannia crenuliformis Aust. (= Plectocolea crenuliformis (Aust.) Mitt.) Plants pellucid, pale to yellowish-green; prostrate with ascending tips , simple or sparingly branched ; leaves rotund , distinc tly bordered ; margins appear swollen, that is, the marginal cells are larger than the cells immediately within

. and forming a sJ.�gh tly elevated border; rhizoids purplish ; in differentiating this species from � gr acillima see the descrip tion for J. gr acillima . 51

Jungermannia hyalina Lyell, 18x

Jungermannia hyalina Lyell (= Plectocolea hyalina (Lyell) Mitt.

Plants pellucid , pale to pure green, reddish-pigmented in

exposed areas ; prostrate , sparingly branched ; leaves rotund ;

oil bodies large , papillose, 2-4 per cell ; perianth emergent beyond bracts ; rhizoids purplish ; not common on soil-cover­

ed sandstone . 52

Lophozia incisa (Schrad .) Durn ., 28x ; apex of shoot, JOx

Lophozia incisa (Schrad .) Durn.

Plants small, bright-green to blue-green ; leaves crowded ,

erect, crisped-concave , with 3-5 unequal , sharp lobes ; cells

large , thin-walled , without trigones, many oil bodies ; very rare on moist sandstone at Rocky Branch . 53

Lophozia bicrenata (Hoffm.) Oum., 23x

Lophozia bicrenata (Hoffm.) Oum .

Plants very small, brownish, with shortly curved-ascend­

ing tips ; leaves crowded, 2-lobed , entire or ragged ; leaf

cells thick-walled ; on dry, exposed soil along paths . 54

Dipl ophyl lum api culatum (Evans) Steph., 25x ; gernmae , 570x

Dipl ophyl lum api culatum (Evans ) Steph .

Plants small , dark green ; many times mistaken for Scapan ia, however easily separated by having the leaf lobes extremely elongate and pointing nearly straight outward from the stem ; dorsal and ventral lobes directed in different directions ; gemmae irregular (polygonal ); no t common on moist sandstone . 55

Scapa nia nemorea (L.) Grolle , 4x ; gemmae , 65ox

Scapa nia nemorea (L.) Grolle ( = S. nemorosa (L.) Durn. )

Plants of moderate size ; leaves normally finely serrate ; at apices of plant, cinnamon-colored gemmae are usually pre- sent; common on sandstone . Nowellia curvifolia (Dicks .) Mitt., 6ox

Nowellia curvifolia (Dicks .) Mitt .

Plants small, slender , green but becoming red in sunlight; leaves deeply concave , deeply divided into 2 very slender , incurved lobes ; rare on coniferous logs or moist sandstone. 57

Cephalozia connivens (Dicks .) Lindb ., 20x ; leaf, 65x

Cephalozia connivens (Dicks .) Lindb .

Plants minute ; leaves strongly decurrent, lobes 1/3 to

1/2- bilobed and sharply acute (ending in one cell) and connivent; common on mo ist rotten logs . 58

Cephalozia lunulifolia (Dum.) Dum. , 100x

Cephalozia lunulifolia (Dum.) Dum. ( = C. media Lindb. .) . Plants minute , pale green ; leaves strongly decurrent,

1/3- bilobed , the lobes acute , connivent; cells thin-walled ; common on shaded sandstone . 59

dorsal view , 17x ventral view; 17x

Geocalyx gr aveolans (Schrad .) Nees

Plants bright-green with a slightly darker , often bluish or grecsy-appearing stem when mo ist; leaves evenly lobed ; underleaves bilobed without accessory tooth-like lobes ; sometimes mistaken for the common Lophocolea heterophyl la, but the absence of the lateral tooth-like lobes on the un­ derleaves should distinguish them ; no t common on rotten logs or shaded , moist humus or sandstone . 60

dorsal view, 21x ventral view, 25x

Lophocolea heterophyl la (Schrad .) Dum.

Plants shiny light-green to dark-green; sterile plants sharply 2-lobed ; underleaves bilobed with lateral tooth- like accessory lobes ; common on decorticated logs and stumps . 61

dorsal view, 50x ventral view , 50x

Frullania tamarisci (L. ) Dum. var . asagrayana (Mont.) Hatt. : dorsal view ; ventral view

Frullania tamarisci (L.) Dum. var . asagr aya na (Mont .) Hatt.

Plants usually uniformly reddish-purple or copper red, becoming dark and blackish-purple when dry ; lobes ovate , lobule ovoid , contracted toward base ; leaf cells of lobe thick-walled, trigones and intermediate thickenings be- coming more conspicuous toward the base ; large discolored cells forming a median line in most of the larger lobes is its· distinguishing charac teristic ; rare on bark of white oak (Quercus alba) . 62

Frullania bolanderi Aust. , 25x, ventral view ; upright, leaf­ less flagella

Frullania bolanderi Aust.

Plants closely appressed to substrate , dark-green to red- dish; stems irregularly pinnate , branches often at right angles to the substrate as upright, leafless flagella, leaves and underleaves distant on stem, lobules helmet-shaped; not

common on bark of mature white oaks (Quercus alba) . 63

Frullania squarrosa ( Reinw . et al . ) Dum. , ventral view, 56x ; median leaf cells , 225x

Frullania sg uarrosa ( Reinw . et al .) Dum.

Plants closely appressed to substrate , green to reddish- brown ; leaves strongly squarrose when mo ist, broadly ovate , and cordate at base; lobule helmet-shaped , inflated ; under-

leaves slightly bifid ; leaf cells from middle of lobe with

distinct trigones and intermediate thickenings ; common on

shaded banks , preferring moist sandstone . 64

Frullania inflata Gott., ventral view, 45x

Frullania inflata Gott .

Plants closely appressed to substrate , brownish-green to reddish ; stems irregularly pinnate ; lobes orbicular , trun­ cate at base; lobule helme t-shaped, truncate at base; leaf cells of lobe thick-walled with inconspicuous trigones,and no intermediate th ickenings ; not common on bark of mature trees . 65

Frullania eboracensis Gott., ventral view, 35x ; median leaf cells , 275x

Frullania eboracensis Gott.

Plants closely appressed to substrate , green often tinged with brown or red ; stems irregularly pinnate ; lobes sub- orbicular , cordate at base, lobules much smaller than lobes , helmet-shaped and truncate at base ; underleaves distant, ovate and bifid about 1/3 its length ; leaf cells of lobe thick-walled with trigones and intermediate thickenings ; not common on bark of mature trees . 66

Frullania riparia Hampe ex Lehm., ventral view, 35x

Frullania riparia Hampe ex Lehm.

Plants green to brownish ; stems irregularly pinnate ; leaves and underleaves distant along stem; lobes ovate , slightly squarrose when mo ist; lobules usually flattened into a small lanceolate leaf ; leaf cells in middle of lobe with slightly thickened walls and dis tinct trigones ; common on shaded sand­ stone . Harpanthus sc utatus (Web . et Mohr) Spruce , 20x ; leaf and underleaf, 20x

Harpanthus scutatus (Web . et Mohr) Spruce

Plants small , yellowish-green, irregularly branched ; leaves notched at the tip or shortly 2-lobed ; underleaves

triangular and undivided , united at its' base to the leaf

just below it; not common, growing intermingled with Ceph - alozia lunulifolia and Blepharostoma trichophyl lum on sand-

stone . 68

GLOSSARY

acuminate- gradually and longly riarrowed to a point. acute- ending in a sharp angle .. archegoniophore- the elevated female receptacle of some mem­ bers of the Order . bifid- deeply split into two leaves . complicate-bilobed- a leaf divided into two lobes, with the lobes folded over each other . connivent- with the distal parts approaching each other . cordate- somewhat heart-shaped , or expanded at the base . cortical- pertaining to the peripheral layer of cells of the stem. decurrent- extending dow� from the main part of insertion . dentate- bearing teeth.

dicho tomous- branching by forking, e.g. • , with the two forks about equal . entire- unlobed or smooth. flage lla- whip-like slender branches or stems , leafless or without rudimentary leaves . ge mma (e)- small , asexually produced reproductive bodies, formed of 1-several cells . incubous- leaves so inserted that the dorsal margin lies above the posterior margin of the leaf in front of it. lanceolate- narrower than ovate , rounded at base and grad­ ually narrowed to a point distally . lobule- the small ventral lobe of an incubously complicate leaf. median leaf cell- a cell fron the middle of a leaf . ° obtuse- with the apex more than 90 . oil body- a small cell organelle consisting of oil-like res­ idue , found mainly in the Order orbicular- circular or nearly so . ovate- egg-shaped , with the broader end downward . papi lla( e ) - small protuberances usually on cell walls . pa racys t- an enlarged or brightly colored cell , very differ­ ent from the surrounding cells . pe llucid- translucent. pe rianth- a tubular sheath, usually narrowed toward the apex , formed of 2-3 connate leaves, that surrounds the developing sporphyte . pinnate- branching like a feather . pr ostrate- lying flat on· the substrate . rhizoids- unicellular hair-like tubes , fastening the gameto­ phyte to its' subs trate . spo rophyte- the -producing generation, consisting of an unbranched structure terminating in a capsule . squarrose- of leaves with midrib bent back at right angles

to the stem . 70

succubous- the type of oblique leaf insertion where the dor­

sal (uppermost) leaf margin lies over the leaf immediately behind it.

thallus- a plant bQdy not differentiated into stems and leaves .

trigone- the thickenings , at the angles , where three or more

cells adjoin.

truncate- abruptly and squarely terminated .

underleaf- amphigastrum , the reduced lower row of leaves,

often hidden among the rhizoids .

uniseriate- composed of a single cell row . 71

CONCLUSION

This study represents a taxonomic survey of the hep­ aticae of eas t- central Illinois . The counties included in this study were Coles and Clark, located in the east-cen­ tral portion of the state , near the Indiana stateline .

These counties are composed primarily of 90% farmland and the remainder devoted to small woods along streams and riv­ ers . The terrain of these counties was greatly influenced by the Wisconsin glacier as evident by its· terminal moraine extending across southern Coles county . The erosive action of the glacial melt water exposed the sandstone bedrock in

th ese areas . The majority of the liverworts are encountered

in these outcrop areas . Of the areas included in this study,

two are in Clark county, Rocky Branch and Rocky Hollow, and

one in Coles county, Rock's Park .

A total of 47 liverworts have been collected in th is region . Four are considered to be rare or endangered . They are Ptilidium pu lcherrimum: (G.Web.) Hampe , Blepharos toma tri­

chophyl lum (L.) Dum., Trichocolea tomentella (Ehrh.) Dum.,

and Marchantia po lymo rph a L. The survival of th ese, and

liverworts in general, as well as other rare plants , is de­

pendent upon the preservation of th ese areas . 72

LI TERATURE CITED

Arzeni , C.B. 1946 . Some bryophytes of Coles and Clark counties , Illinois Acad . Sci . 40 :44-49 .

1950 . The Hepaticae of the Douglas Lake region, Michigan . Edwards Brothers Inc . Ann Arbor , Michi­ gan . 30pp .

---- . 1980 . Mosses and liverworts . Eastern Illinois University. 253PP •

Awal t, F.L. 1979 . Soil survey of Clark county, Illinois . United States Dept. of Agriculture, Soil Conserva­ tion Service . 176pp .

Conard , H.S. and Paul Redfearn . 1979 . How to know the mosses and liverworts . li\lm . C. Brown Co . Publishers Dubuque , Iowa . 302pp .

Crum, Howard . 1980 . Leafy liverworts of the Straits area. unpublished manus cript. 11pp .

Evans, A.W. 1897 . A revision of the North American species of Frullania, a genus of Hepaticae . Trans . Conn . Acad . Arts and Sci. 10 (1) :1-39, pls . 1-15.

Frye , T.C. and Lois Clark . 1937-1947 . Hepaticae of . Univ . of Washington Publications . Seattle , Washington . Vol . 1-5.

Gleason, H.A. and A. Cronquist. 1963 . Manual of vascular plants of Northeastern United States and adjacent Canada . D. VanNorstrand Co . 810pp .

Kurz , Herman and Thomas Little . 1933 . Liverworts of north and central Florida . Bulletin Florida State College for Women . Vol XXVI No . 3. 40pp .

Macvicar , Symers M. 1926 . The student 's handbook of Bri­ tish Hepatics . V.V. Sumfield , Station Street, Lon­ don . 464pp .

Mohlenbrock, Robert H. 1975 . Guide to the vascular flora of Illinois . Southern Ill. Univ. Press . 684pp . 73

Schuster , R.M. 1953 . Boreal Hepaticae- a manual of the liv­ erworts of Minnesota and adjacent regions . Am . Mid . Nat . Vol . 49 , No . 2. 684pp .

1966 . The Hepaticae and Anthocerotae of North America. Columbia Univ . Press . New York and Lon­ don. Vol . I-III .

Smith , R.S. and E.E.· DeTurk . 1929 . Coles county soils . Univ . of Illinois Agriculture Experiment Station . Soil Report No . 44 . 62pp .

Steere , William C. 1960. Liverworts of southern Michigan . Cranbook Institute of Science . Bloomfield Hills , Michigan . No . 17. 97pp .

Stotlar, R. and B. Stotlar . 1977 , A checklist of the liv­ worts and hornworts of North America . Bryologist 80 : 405-428.