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Natural history and parasitism of Geocaulon lividum (santalaceae) Warrington, Patrick Douglas 1970

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The Natural History and Parasitism of Geocaulon lividum (Santalaceae) by Patrick Douglas Warrington B.Sc, University of British Columbia, 1964. A Thesis submitted in partial fulfillment of the requirements for the degree of Ph.D. in the Department of Plant Science We accept this thesis as conforming to the accepted standards. •The University of British Columbia, 1970. In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t he r e q u i r e m e n t s f o r an advanced degree a t t he U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and S t u d y . I f u r t h e r ag ree t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by the Head o f my Depar tment o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s thes. is f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Patrick D. Warrington Depar tment o f Plant Science The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada Date April 1970 Abstract The natural history, structure and parasitism of Geocaulon lividum (Richardson) Fern'ald of the Santalaceae are described. Both the present study and that of Piehl (1963) maintain the generic segregation of Geocaulon and Comandra. A l l available data on the distribution of Geocaulon are presented in a series of Maps, in the accompanying text and in an Appendix. The data are drawn from the literature, from herbarium material and from f i e l d work. The only vegetation zones of Canada in which Geocaulon i s not commonly found are . the prairie grasslands, the arctic and alpine tundra and the west coast Douglas Fir and Hemlock forests. The ecological conditions under which Geocaulon is usually found are noted. A l i s t of known hosts is given; many of them represented by voucher specimens in the herbarium of the University of Lethbridge (LEA). Anatomical and morphological data on the various portions of the Geocaulon plant are presented under a series of topics: rhizome, short-shoot, aerial shoot, leaf, inflorescence, flower, f r u i t , and root. These descriptions are accompanied by i l l u s t r a t i v e material in the form of figures, plates and photographs. Criteria for estimating the age of rhizomes and short-shoots are given along with a series of representative short-shoots with age determina-tions. Illustrations also indicate some of the diversity in the appearance of the inflorescence units. The haustorium is described in some detail. It is treated under several topics: general characteristics, development and structure. The structure is treated f i r s t under outer cortical region and second the,inner vascular.core. The core is further divided into the mother root, transitional and haustorial zones. The d i s t r i b u t i o n p a t t e r n o f v e s s e l elements w i t h i n the c o r e r e g i o n i s graphed. A b r i e f d i s c u s s i o n i s i n c l u d e d of as y e t u n r e s o l v e d problems a s s o c i a t e d w i t h t h e s t r u c t u r e and f u n c t i o n o f the h a u s t o r i u m of most S a n t a l a -ceous r o o t p a r a s i t e s . Some p r e v i o u s l y u n r e p o r t e d t y p e s of t h i c k e n i n g s on parenchyma w a l l s were found i n rhizomes and s h o r t - s h o o t s o f Geocaulon. These were found m o s t l y on c o r t i c a l c e l l w a l l s but were a l s o found on the w a l l s o f o t h e r t i s s u e s . A m i c r o c h e m i c a l i n v e s t i g a t i o n was undertaken t o c h a r a c t e r i z e t h e s e w a l l t h i c k e n i n g s . They were found t o be h o l o c e l l u l o s i c but were n o t a s s i g n e d t o e i t h e r t h e h e m i c e l l u l o s e o r c e l l u l o s e f r a c t i o n s . They were n o t composed o f a l p h a - c e l l u l o s e , t h e major s t r u c t u r a l m a t e r i a l o f p l a n t c e l l s . Table of Contents Ti t l e Page Abstract page i Table of Contents page i i i List of Figures . • page v List of Plates page v i List of Maps . . . • P aS e v i i List of Graphs page v i i i List of Photographs page ix Acknowledgements page x i i Introduction page 1 The Santalaceae page 2 Geocaulon vs. Comandra page 5 Geocaulon lividum page 6 Geocaulon - Distribution page 9 - Ecology page 33 - Parasitism page 48 Geocaulon - rhizome page 52 - short-shoot page 60 - aerial shoot page 76 - leaf page 80 - inflorescence page 88 - flower . page 98 - fruit. page 105 ; - root page 109 iv Geocaulon Haustorium page 111 - general characteristics page 111 - development page 111 - internal structure P a § e 116 - cortical region page 117 - mother root zone page 120 - transition zone page 121 - haustorial zone page 123 - vessel element distribution page 125 Geocaulon Cell Wall Thickenings page 129 Photographs page 136 Bibliography page 152 Appendix 1 - Techniques page 158 Appendix 2 - Geocaulon distribution data page 160 List of Figures Fig- 1. Examples of structures and their symbols as used in succeeding figures. page 40 Fig. 2. Old rhizome in i t s natural configuration. page 42 Fig. 3. Rhizome removed from a rotting log. page 45 Fig. 4. Rhizome removed from a rotting log. page 46 Fig- 5. Continuation of Fig. 4. page 47 Fig. 6 - Branching in the rhizome vascular system. page 58 Fig. 7. Representative short-shoots. page 69 Fig. 8. Representative short-shoots. page 70 Fig- 9. Representative short-shoots. page 71 Fig. 10. Representative inflorescences. page 92 Fig. 11. Representative inflorescences. page 93 Fig- 12. Representative inflorescences. page 94 Fig. 13. Vascular skeletons of 4 inflorescences. page 97 Fig- 14. Vascular skeleton of a young f r u i t . • page 107 Fig. 15. Sectional views of the haustorium. page 115 v i List of Plates Plate 1. Representative leafy shoot in f r u i t . page 35 Plate 2. Rhizome with short-shoots and aerial shoots. page 37 Plate 3. Two Geocaulon rhizome apices. page 54 Plate 4. One- and two-year-old Geocaulon short-shoots. • page 62 Plate 5. Details of one-year-old short-shoots. page 64 Plate 6. A complex old short-shoot. page 66 Plate 7. Old short-shoot indicating points at which sections were cut for anatomical comparisons of xylem structure. page 73 Plate 8. Details of the basal portion of a short-shoot, page 78 Plate 9. Venation pattern in a cleared leaf. " page 82 Plate 10. Upper epidermis of Geocaulon leaves. page 84 Plate 11. Lower epidermis of Geocaulon leaves. page 86 Plate 12. Three Geocaulon inflorescences. page 90 Plate 13. Bud, mature flower and young f r u i t of Geocaulon. page 100 Plate 14. Venation pattern in cleared tepals. page 10 3 Plate 15. Three views of a Geocaulon haustorium. page 113 V l l List of Maps Map 1. British Columbia and the southern Yukon; west of 120 degrees west longitude. page 11 Map 2. Alberta, southeastern British Columbia, western Saskatchewan arid the southern Northwest Territories;,east of 120 degrees west. page 13 Map 3. Manitoba, eastern Saskatchewan, western Ontario, and southwestern Hudson's Bay page 15 Map 4. Ontario, James Bay, west-central Quebec, and the northern Great Lakes region. page 17 Map 5. South and south-central Quebec, eastern Ontario, the northeastern United States, Gaspe and western New Brunswick. page 19 Map 6. Northern and eastern Quebec, Labrador and eastern Gaspe. page 21 Map 7. Southeastern Quebec, Gaspe, New Brunswick , Nova Scotia, Prince Edward Island, Maine, Newfoundland and southeastern Labrador. page 2 3 Map 8. Alaska, Yukon and northwestern British Columbia, page 25 Map 9. Eastern Yukon west and central Northwest Territories and northern Alberta. page 27 Map 10. , South shore of Hudson's Bay, northeastern Manitoba, northern Ontario and east-central Quebec. page 29 Map 11. Southeastern British Columbia, southern Alberta, southern Saskatchewan, and northern Idaho, Montana and Washington. page 31 vm List of Graphs Graph 1. Distribution of three kinds df vessel elements page 128 in the haustorium of Geocaulon. ix Photograph 1. Photograph 2. Photograph 3. Photograph 4. Photograph 5. List of Photographs Basal portion of a short-shoot in i t s f i r s t year of growth. Section A, Plate 5. Short-shoot/aerial shoot junction region, f i r s t year of growth. Section B, Plate 5. Base of aerial shoot dn one-year-old short-shoot. Section C, Plate 5. Vascular bundle of Geocaulon rhizome. Rhizome cross-section at base of a short-shoot. Section D, Plate 7. Photograph 6. Cross-section of base of short-shoot. Section E, Plate 7. Cross section of next oldest portion of short-shoot. Section F, Plate 7. Cross-section of oldest portion of the short-shoot. Section G, Plate 7. Cross-section of the base of the aerial shoot. Section H, Plate 7. Cross-section of aerial shoot. Median longitudinal section of Geocaulon rhizome apex. Photograph 12. Cross-section of a Geocaulon root near a large haustorium. Photograph 7. Photograph 8. Photograph 9. Photograph 10. Photograph 11. Photograph 13. Longitudinal section of f r u i t wall in an immature Geocaulon f r u i t . page 1.37 page 137 page 137. page 137 page 139 page 139 page 139 page 139 page 141 page 141 page 141 page 141 page 143 Photograph 14. Exocarp of mature Geocaulon f r u i t wall. page 14-3 Photograph 15. Longitudinal section of Geocaulon f r u i t showing endosperm and embryo. page 143 Photograph 16. Longitudinal section of mature Geocaulon f r u i t showing exocarp, mesocarp, endocarp and endosperm. page 143 Photograph 17. Mother root zone bf haustorial core in cross-section. Section A, Fig. 15. page 145 Photograph 18. Transition zone of haustorial core in cross-section. Just below level B, Fig. 15. page 145 Photograph 19. Haustorial zone of haustorium in cross-section, at level of the host cortex. Section C, Fig. 15. page 145 Photograph 20. Tip of the.haustorial wedge deep in the host. Cross-section of Geocaulon, longitudinal section of host. Section D, Fig. 15. page 145 Photograph 21. Longitudinal section of mother root zone of Geocaulon haustorium. Section E , above level B, Fig. 15. page 147 Photograph 22. Longitudinal section of a Geocaulon haustorium through the transition zone. Sections E, or F, between level A and a point below level B, Fig. 15. ' page 147 Photograph 23. Longitudinal section of Geocaulon haustorium. Sections E or F, between levels C and D, Fig. 15. page 147 xi Photograph 2 Ji. Longitudinal section of Geocaulon haustorium. Section G, below level B, Fig. 1 5 . page 1 4 7 Photograph 2 5 . Longitudinal section of a young Geocaulon haustorium and i t s mother root. page 1 4 9 Photograph 2 6 . Longitudinal section of a young Geocaulon haustorium. Section F, below level B , Fig. 1 5 . page 1 4 9 Photograph 2 7 . Cross section through a Geocaulon haustorium in the haustorial zone immediately below the transition zone. Sections E or F, between levels B and C, Fig. 1 5 . page 1 4 9 Photograph 2 8 . Cross-section through the transitional zone of a Geocaulon haustorium. Sections E or F, between levels B and C, Fig. 1 5 . page 1 4 9 Photograph 2 9 Representative c e l l wall thickenings in to 3 4 . Geocaulon rhizome cortex c e l l walls seen in cross section. page 1 5 1 X l l Acknowledgements I wish to thank. Dr. Job K u i j t , my research advisor, and Dr. Ian McTaggart Cowan, Dean of Graduate Studies, f o r t h e i r help in making possible the off-campus study arrangements. The Faculty and S t a f f of the Biology-Department of The University of Lethbridge, where most of the Work was c a r r i e d out, were most h e l p f u l at a l l times. Dr. John Powell of the Pathology Section of the Department of Forestry i n Calgary gave me a guided tour of known Geocaulon s i t e s i n the Kananaskis region of Alberta. Mr. John Nagy, a student i n the Biology Department of The.University of Lethbridge, dug out many hosts, helped with f i e l d work, and made independent c o l l e c t i o n s and observations of Geocaulon i n northern Alberta and the Northwest T e r r i t o r i e s . My wife Pamela helped dig out hosts and helped extend the d i s t r i b u t i o n maps. Many people, and i n s t i t u t i o n s throughout the world sent specimens of other members of the Santalaceae f o r my study. Data on the d i s t r i b u t i o n of Geocaulon i n northeastern Washington were supplied'by the Herbarium of Washington State Un i v e r s i t y , Pullman, Washington. The following Herbaria supplied sheet of Geocaulon f o r my study i n 1968. Department of Botany, University of B r i t i s h Columbia. Department of Botany, University of Manitoba. MacDonald College, University of M c G i l l . Herbier P. Louis-Marie, Laval University. Department of Botany, University of Saskatchewan. Department of Botany, Purdue University. Department of Botany, University of Washington. Osborne Botanical Laboratory, Yale University. Forest Research Laboratory, Fredericton, New Brunswick. X l l l The National Museum of Canada. , Forest Entomology Laboratory, Vernon, British Columbia. Petawawa Forest Experiment Station, Chalk River, Ontario. The New York Botanical Gardens. Department of Botany, University of Calgary. Department of Botany, University of Victoria. Department of Botany, University of Minnesota. Department of Botany, Queen's University, Kingston, Ontario. Department of Botany, University of Toronto. Ontario Agricultural College,. Guelph, Ontario Arnold Arboretum and Gray Herbarium, Harvard University. Plant Research Institute, Department of Agriculture, Ottawa. The Provincial Museum, Victoria, British Columbia. The Smithsonian Institution, Washington D.C. To the Curators of a l l these herbaria I would like to- extend my thanks for their cooperation in making specimens available to me. - 1 -Introduction Geocaulon, a root parasite, is one of two monotypic Canadian genera of the Santalaceae, a family composed mostly of root parasites. The other Canadian genus, Comandra, once contained Geocaulon lividum (as C_. livida) u n t i l Fernald segregated the two in 1928. this segregation is not followed by a l l workers and even today two major f l o r i s t i c works (Hitchcock, et al-1964; Gleason-1952) refer G_. lividum to Comandra. The comprehensive treatment of Comandra by Piehl (1963) excludes Geocaulon, thus following Fernald. Very l i t t l e published information is available for Geocaulon lividum per se and apart from the brief and partly erroneous description of the haustorium (Moss-1926) the information is entirely distributional. Fernald's work contains an important misconception, regarding the flowers, which needs cla r i f i c a t i o n . Piehl's comprehensive work on Comandra has pointed out the need for a parallel treatment on Geocaulon. This the present work attempts to f u l f i l l . This study does not contain any work on germination or on the seedling . simply because seed germination t r i a l s in the laboratory and the garden were unsuccessful and no seedlings were ever found in nature. . There is no reported physiological work on any aspects of parasitism in Geocaulon. Extensive literature research was carried out at the University of British Columbia in 1967-1968 on a l l the genera of Santalaceae. This work provided a broad background on the characteristics and inter-relationships of the members of the family. In the F a l l of 1968 study headquarters were moved to the University of Lethbridge which provided much better access to Geocaulon in the f i e l d . • ; ' - 2 -The Santalaceae T h e members of t h i s f a m i l y may be t r e e s , shrubs, herbs or woody climbers. A l l are presumed to be p a r a s i t i c on the roots or branches of other p l a n t s , although p a r a s i t i s m has not been documented i n a l l cases. Leaves may be o p p o s i t e , a l t e r n a t e or apparently s c a t t e r e d along the branch. Leaves are without s t i p u l e s and g e n e r a l l y e n t i r e . They may be l a r g e or reduced to minute s c a l e s ; t h i c k and l e a t h e r y or membranous; and l i n e a r to o r b i c u l a r . Terminal and a x i l l a r y i n f l o r e s c e n c e s are found and i n c l u d e cymes, s p i k e s , corymbs and racemes as w e l l as s o l i t a r y f l o w e r s . U s u a l l y there i s a s i n g l e greenish or p e t a l o i d p e r i a n t h whorl which i s g e n e r a l l y adnate t o the ovary. In Anthobolus and Exocarpos the p e r i a n t h i s f r e e t o i t s base. The p e r i a n t h i s 3 - t o 6-lobed above the tube p o r t i o n , v a l v a t e , i m b r i c a t e or f r e e . In many genera a t u f t of h a i r s a r i s e s on the a d a x i a l surface of the p e r i a n t h members at the base of the f i l a m e n t . The p l a n t s may be hermaphroditic, polygamous, monoecious or d i o e c i o u s . The number of stamens i s g e n e r a l l y the same as that of p e r i a n t h members. Stamens are u s u a l l y i n s e r t e d on the bases of the p e r i a n t h members and are thus opposite to the p e r i a n t h members. The mostly 2 - l o c u l a r anthers, with probable exceptions i n Choretrum and Leptomeria at l e a s t , commonly open by lengthwise s l i t s . Filaments take various forms. A f l o r a l d i s c i s present and may be lobed or e n t i r e . This d i s c may be b a s a l or i t may i n v e s t the p e r i a n t h tube. S t y l e s take v a r i o u s forms and stigmas range from e n t i r e t o 5 - l o b e d . I n f e r i o r through t o s u p e r i o r ovaries are found and though there may be 5 b a s a l p a r t i t i o n s there i s u s u a l l y only one l o c u l e , as the others g e n e r a l l y abort. These ovules are commonly pendulous from the apex of a f r e e - c e n t r a l p l a c e n t a l column. No seed coat e x i s t s i n the f a m i l y . The integument, when present, i s absorbed by the endosperm t i s s u e . Nuts and drupes are the u s u a l - 3 -fruits and they may be either dry or fleshy. Fruits are often crowned by the perianth and/or the disc. The family, of nearly 4-0 genera, has more or less world-wide distribution but i t is noticeably absent from Mexico, the West Indies and Central America. The currently accepted genera of the Santalaceae are: Acanthosyris, Amphorogyne , Anthobolus, Arjona, Austroamericium, Buckleya, Cervantesia, Choretrum, Cladomyza, Colpoon , Comandra, Daenikera, Dendromyza, Dendrotrophe, Dufrenoya, Eucarya, Exocarpos, Geocaulon, Hylomyza, Jodina, Leptomeria, Mida, Myoschilos, Nanodea, Nestronia, Okoubaka, Omphacomeria, Osyris, Osyridicarpos, Phaceliaria, Pyrularia, Quinchamalium, Rhoiacarpos, Santalum, Scleropyrum, Spirogardnera, Thesidium, Thesium. The description of the family and the l i s t i n g of i t s member'genera has had to be synthesised from many widely scattered sources as no really comprehensive, up-to-date treatment of the family exists. Some of the more important sources of information are the following works: Bentham and Hooker-1883 , Chatin-1860, Danser-1940, DeCandolle-1857, Engler and Prantl-1894, Fernald-1928, Gray-1846, Hendrych-1963, Hutchinson-1959, Kuijt-1969, Metcalf and Chalk-1965, Stauffer-1957, 1959, 1961, 1968, 1969, Stauffer and Hurlimann-1957. In the past several other genera have been included in the family but have now been excluded for various reasons. Champereia is now treated in the Olacacean complex (Kuijt-1969, Willis-1966). Calyptosepalum has been transferred to the Euphprbiaceae (Smith and Ayensu-1964, Steenis-1960). Grubbia and Myzodendron have been placed in their own families, Grubbiaceae • and Myzodendraceae respectively. Fusanus has been s p l i t into Colpoon, Santalum and Mida.and for lack of members the genus no longer exists. - 4 -Sarcopus is synonymous with Exocarpos (Stauffer-1959).• Henslowia has been s p l i t into Dendromyza,. Hylomyza, Dufrenoya and Dendrotrophe (Danser-1940). Austroamericium (Hendrych-1963) may well be united/reunited with Thesium in the future as more detailed comparitive work is carried out (personal opinion, Kuijt-1970). The five North American genera are Buckleya, Comandra, Geocaulon, Nestronia and Pyrularia.• Comandra, Nestonia and Geocaulon are monotypic. Comandra has one subspecies in Europe (Piehl-1963) but Nestronia and Geocaulon are confined to North America. Buckleya and Pyrularia both have one species in eastern North America; the former also has one species in Japan and the latter a second species in the Himalayas (Kuijt-1969). Buckleya, Nestronia and Pyrularia are confined to the eastern United States h i l l s ; Comandra grows throughout the southern portion of North America and Geocaulon in the northern portion. - 5 -Geocaulon vs. Comandra Geocaulon and Comandra were once included in the same genus Comandra. While they are no longer considered congeneric by most workers they are s t i l l treated as such by some more conservative writers (Hitchcock et al-1964; Gleason-1952). Some of the differences between these genera w i l l be presented below. The interested-reader should read the descriptive portions of this thesis and that of Piehl-1963 in order to compare the two genera more f u l l y . In the aerial stem of Geocaulon the vascular system is in the form of separate bundles with bundle caps. The vascular system of Comandra's aerial stems is in the form of a continuous ring of xylem and phloem produced by a complete ring of cambium. Isolated fibre caps occur in the cortex (Piehl-1963). Almost a l l Geocaulon haustoria are simple whereas a great ,many Comandra haustoria appear to be compound (Kuijt-1969, plate 7-1-e; Moss-1926, Fig. 8). The flowering stem of Comandra branches regularly and profusely (Piehl-1963) but the flowering stem of Geocaulon does not branch except for very rare and very small branches in place of inflorescences. In Comandra there i s aerial stem dimorphism with some stems being reproductive and some being vegetative (Kuijt-1969; Piehl-1963) but there i s no dimorphism in Geocaulon. In Comandra rhizomes, roots arise directly above axillary buds and the rhizome apex is swollen (Kuijt-1969; Piehl-1963) whereas no such root and bud association nor swollen apex is found in Geocaulon. - 6 -Geocaulon l i v i d u m The taxon was d e s c r i b e d o r i g i n a l l y as Comandra l i v i d a by R i c h a r d s o n i n 1 8 2 3 . In 1 8 2 5 S p r e n g e l t r a n s f e r r e d i t t o the genus H a m i l t o n i a W i l l d . Bentham and Hooker put i t back i n Comandra i n the s e c t i o n Haplocomandra i n 1 8 8 0 . F i n a l l y , i n 1928, F e r n a l d d e s c r i b e d t h e new genus' Geocaulon based on _C. l i v i d a and the taxon i s now r e c o g n i z e d by most b o t a n i s t s as G. l i v i d u m ( R i c h a r d s o n ) F e r n a l d . The genus i s monotypic and c o n f i n e d t o N o r t h America. . I t i s found i n A l a s k a , t h e n o r t h e a s t e r n U n i t e d S t a t e s and t h r o u g h o u t the B o r e a l - S p r u c e - F i r zone o f Canada. The f o l l o w i n g i s a b r i e f d e s c r i p t i o n o f t h e s p e c i e s . P o i n t s mentioned below w i l l be e l a b o r a t e d on i n l a t e r c h a p t e r s . T h i s d i s c u s s i o n i s p r e s e n t e d i n o r d e r t o b r i n g t o g e t h e r i n r e l a t i v e l y b r i e f form the s a l i e n t c h a r a c t e r i s t i c s o f the p l a n t f o r purposes of i d e n t i f i c a t i o n and f o r comparison w i t h o t h e r t a x a o f t h e f a m i l y . A s m a l l herbaceous a e r i a l stem, up t o 3 dm i n h e i g h t , a r i s e s from a-y e l l o w i s h t o r e d d i s h - b r o w n , smooth, c r e e p i n g rhizome. The p l a n t i s a r o o t p a r a s i t e and has s m a l l w h i t i s h o r tan h a u s t o r i a on i t s r o o t s . These h a u s t o r i a . j o i n i t s r o o t s t o t h o s e o f o t h e r a d j a c e n t p l a n t s . The l e a v e s may be t h i n , f l a c c i d , p a l e t o g r e y i s h - g r e e n and r e l a t i v e l y l a r g e ; o r s m a l l e r , f i r m e r and r e d d i s h t i n t e d . The former c o n d i t i o n o c c u r s i n m o i s t , shaded a r e a s and the l a t t e r i n open, exposed s i t e s . The l e a v e s a r e a l t e r n a t e l y a r r a n g e d on t h e a e r i a l stem but no c o n s i s t e n t p h y l l o t a c t i c o r d e r i s d i s c e r n i b l e . Major v e i n s a r e prominent on the lower s u r f a c e of t h e l e a f . Leaves a r e o f t e n i n f e c t e d w i t h a v i r u s c a u s i n g y e l l o w o r o c c a s s i o n a l l y r e d d i s h v e i n a l a r e a s . In extreme cases t h e i n t e r v e i n a l a r e a s a r e d i s c o l o u r e d t o o and l e a f shape i s d i s t o r t e d from i t s n ormal o v a t e t o obovate shape t o become q u i t e i r r e g u l a r . The o t h e r w i s e - 7 -entire margins become sinuate and the blade area much reduced. The basic inflorescence unit is a dichasium. Considerable variation from this basic unit is found. The lateral flowers usually abort while s t i l l immature and thus in many portions of the range a single f r u i t is found on the central flower. The inflorescence-bearing nodes are those with the largest, most ful l y expanded leaves, about one half way up the aerial stem. In eastern Canada one inflorescence bearing one fru i t is the rule. Western Canadian plants have up to six inflorescences. In the southwest one f r u i t is s t i l l the rule but in the northwest two or three fruits become quite common for each inflorescence. Peduncles are almost always in the axils of 5 contiguous mature leaves and pedicels are usually in the axils of bracts. The inconspicuous flowers are perfect, small, and greenish. The base of the single perianth whorl is adnate to the inferior ovary; the top of the perianth is rotate. A lobed disc lines the perianth tube. Stamens are inserted on the bases of the perianth lobes which alternate with the lobes of the disc. Filaments are short and ligulate; anthers 2-lobed and ovate. The anther dehisces introrsely by longitudinal s l i t s . A tuft of hairs arises on the adaxial surface of the perianth members at the base of each filament. This bundle of hairs is closely appressed to the abaxial surface of the filament, passes up the f u l l length of the filament and between the anthers along the' connective. Some hairs project beyond the anthers towards the stigma. The one-celled inferior ovary has a long, basal, contorted placental column bearing several ovules. Embryo sacs are also long and twisted when they develop. There is a short, cylindrical and f a i r l y massive style with a variously lobed stigmatic surface. The f r u i t is juicy, red or orange, ovoid-globose and edible but not very tasty. Although i t has been considered a type of drupe or - 8 -nut i t does not f i t s a t i s f a c t o r i l y i n t o common f r u i t c l a s s i f i c a t i o n schemes. V e s t i g e s o f t h e d i s c and p e r i a n t h crown the f r u i t , p a r t i c u l a r l y when i t i s s t i l l immature. A mature f e r t i l e f r u i t c o n s i s t s o f an embryo embedded i n abundant endosperm which i s surrounded by a t h i n l a y e r o f c r u s h e d gametophyte and endocarp c e l l s . Next i s the h a r d s t o n y mesocarp o f s m a l l t h i c k - w a l l e d c e l l s . The v a s c u l a t u r e o f the f r u i t i s found on and w i t h i n t h i s l a y e r . The ' whole i s s u r r o u n d e d by a s o f t , f l e s h y , l a r g e - c e l l e d exocarp w i t h a t h i n e p i d e r m a l l a y e r . N e i t h e r seed c o a t n o r integuments e x i s t i n Geocaulon. - 9 -Distribution Geocaulon is apparently confined to North America north of 4 4 degrees in the east and north of 48 degrees in the west. The plant is generally associated with the Boreal-Spruce-Fir zone of North America. This forest community sets the southern limits of the range of Geocaulon f a i r l y well but in the northern part of North America the distribution limit is more nearly determined by stands of Populus, Salix or Betula. Conspicuous gaps occur in the distribution of Geocaulon in North America. It is absent in the northern Ungava peninsula, the southern Canadian prairies and the southern west coast of British Columbia. Dots on the accompanying maps (1 through 11) indicate the range of Geocaulon. The data were obtained from several sources: personal observations, herbarium sheets borrowed from the institutions l i s t e d in the acknowledgements, and from the literature. The literature sources are: Anderson-1946, Baker-1961, Bowman-1932, Eggleston-1902, Fernald-1928, Fraser and Russel-1953, Harris-1965, Hulten-1968, 1969, Hustich-1951, LaRoi-1967, Macoun-1886, Perry-1931, Porsild-1939, 1950, 1951, Potter-1932, Raup-1930, 1935, 1947, Rouleau-1944, Scamman-1940, Scoggan-1950, 1951, 1952, 1957. With regard to distribu-tion one important fact must be emphasized. The maps do not show the actual distribution of Geocaulon but rather the wanderings of i t s collectors. It is readily apparent that most collections in the north are on sea shores, lake shores and river banks; while in the south collections tend to be near roads, railways, s k i - l i f t s , u t i l i t y company access t r a i l s and along recreation t r a i l s in parks. Appendix 2 l i s t s the latitude and longitude of a l l spots on the maps to the nearest minute with an error, in most cases, of at most 3 minutes. These entries are arranged in ascending order by latitude. - 10 -Map 1 Geographic distribution of Geocaulon: British Columbia west of 120 degrees, portions of Alberta, Washington, Yukon, Northwest Territories and the Alaska panhandle where they border on B.C. The scale line indicates 100 miles. - 12 -Map 2 Geographic distribution of Geocaulon: British Columbia east of 120 degrees, Alberta, western Saskatchewan, the southern portion of the Northwest Territories bordering on Alberta and Saskatchewan, and northern Washington. The scale line is 100 miles long. - 11+ -Map 3 Geographic distribution of Geocaulon: eastern Saskatchewan, Manitoba, western Ontario, and the southeastern corner of Northwest Territories. The scale line represents 100 miles. 3 - 16 -Map 4 Ge o g r a p h i c d i s t r i b u t i o n o f Geocaulon: O n t a r i o e x c e p t t h e s o u t h e a s t e r n c o r n e r , e a s t e r n M a n i t o b a , James Bay, s o u t h e a s t e r n Quebec, n o r t h e r n M i n n e s o t a and M i c h i g a n . The s c a l e l i n e r e p r e s e n t s 100 m i l e s . - 18 -Map 5 Geographic distribution of Geocaulon: eastern Ontario, western Labrador, southwestern Quebec, western New Brunswick and northern portions of Maine, Vermont, New York and New Hampshire. The scale line represents 100 miles. - 2 0 -Map 6 Geographic distribution of Geocaulon: Labrador, the eastern tip of the Gaspe peninsula, northern New Brunswick and northeastern Quebec. The scale line represents 100 miles. - 2 2 -Map 7 Geographic distribution of Geocaulon; Newfoundland, southeastern Labrador, Nova Scotia, New Brunswick, Prince Edward Island, southeastern Quebec, and northern Maine and New Hampshire. The scale line, represents 100 miles. - 2 4 -Map 8 Geographic distribution of Geocaulon: southeastern Alaska, southern Yukon, northern Alaska panhandle, and northwestern British Columbia. The scale line represents 100 miles. - 26 -Map 9 Geographic distribution of Geocaulon: Yukon, the southwestern portion of the Northwest Territories, northern British Columbia, Saskatchewan and Alberta The scale line represents 100 miles. - 2 8 -Map iO Geographic distribution of Geocaulon: James Bay, the southern Hudson's Bay shore line, west-central Quebec, northern Ontario, northeastern Manitoba, and the southeastern t i p of the Northwest Territories. The scale line represents 100 miles. - 30 -Map 11 Geographic distribution of Geocaulon: southeastern British Columbia, southern Alberta and Saskatchewan, and northern Washington. The scale line represents 100 miles. - 3 2 -The a c t u a l distribution i s not nearly as spotty as the maps indicate. In areas where Dr. John Powell or I have looked for Geocaulon intensively we have found many individual sites in an area that would be covered by one dot on the map. There is however a curious distribution on a local level. The plants grow in discretely bounded patches suggestive in clonal growth and spread. A patch, at least in the southern part of the range, is seldom more than several hundred square feet in area. The margins of a patch can be very precisely determined; there are no scattered plants between patches. In the southern part of the range where the f i e l d work was carried out no seedlings were found, nor was germination in the laboratory successful. Perhaps the reason for the discretely bounded patches is that no germination occurs from randomly scattered fruits and thus a l l growth is clonal and so spatially restricted. One can walk through a forest where Geocaulon is known to occur and miss the plant completely by passing between patches. This knowledge makes i t even more evident that the maps are not complete records of Geocaulon distribution. - 3 3 -Ecology The following brief descriptions of the sites where Geocaulon has been collected have been gleaned,,for the most part, from herbarium sheet labels and partly from personal observation. They give an impression of the diversity of habitats in which Geocaulon may be found. Pine woods; open sandy terraces; sandy terraces in pine woods; mossy places in spruce woods; wooded river flats; rich poplar woods; open spruce woods; lowland pine; boreal-spruce-fir zone of Canada; dry or moist woods, rock outcrops and gravel ridges almost throughout Manitoba; in bogs; gravel ridges; sphagnum bogs; wet coniferous woods; wet rotting logs in dry forests; among Vaccinium and Arctostaphylos on gravel and sandy flood plains; with Vaccinium and Linnaea on open slopes and ridges in pine forests; with Cornus canadensis in mossy spruce forests; gravelly river banks among willows and birches; black spruce and lichen forests; white spruce forests with thick moss cover; Equisetum bogs and their slopes; mossy stream banks; moss covered rock outcrops. The major portion of a Geocaulon plant is the rhizome which grows in the AQ or A^ zones of the soil; between the soil or rock surface and the moss cover or deeply enough in sand to reach permanent water. The leafy, flowering, aerial shoot (Plate 1) dies at the first hard frost of winter, but the rhizome and attached short-shoots (Plate 2) overwinter. In the Fall one can often find blackened, withered stems lying on the surface of the moss with the bright red or orange fruits s t i l l attached. In regions where personal observations have been made i t appears that the rhizome is the major i f not the only means of propagation as no seedlings have ever been observed. In the southern part of the range at least the number of fruits is small compared to the number of aerial - 3 4 -P l a t e 1 The a p i c a l p o r t i o n o f a Geocaulon a e r i a l shoot i n f r u i t . Note the c o n t i g u o u s i n f l o r e s c e n c e - b e a r i n g nodes and the sudden d e c r e a s e i n l e a f s i z e above the i n f l o r e s c e n c e - b e a r i n g nodes. Leaves a r e a l t e r n a t e but no c o n s i s t e n t p h y l l o t a c t i c p a t t e r n i s o b v i o u s . The a p i c a l p o r t i o n o f the stem c o n t i n u e s t o grow u n t i l k i l l e d by w i n t e r f r o s t s . 7 cm - 36 -P l a t e 2 P o r t i o n of a Geocaulon rhizome w i t h i t s a t t a c h e d a e r i a l s h o o t s and r o o t s . The background l i n e r e p r e s e n t s s o i l l e v e l . The b l a c k s t r a n d s a r e t h e v a s c u l a r b undles and f i b r e s o f t h e a e r i a l s h o o t s of p r e v i o u s seasons. - 37 - 3 8 -s h o o t s . The rhizome would appear to be perennial since one can f i n d long sections a c t i v e l y growing at one end and r o t t i n g at the other. A l l the a e r i a l shoots in a patch are l i k e l y part of the same clone. Moisture appears to be a decisive f a c t o r i n Geocaulon d i s t r i b u t i o n on the l o c a l l e v e l , thus i f there i s no standing or running water nearby there w i l l c e r t a i n l y be seepage f o r part of the year at l e a s t . However Geocaulon does not grow submerged. Like many other 'semi-parasites' Geocaulon w i l t s quickly when excavated and f o r t h i s reason i s d i f f i c u l t to transplant s u c c e s s f u l l y . In general shaded areas are preferred but deeply shaded plants may be up to a month l a t e r flowering than nearby plants exposed to morning sun. Geocaulon i s very often found in conjunction •with Linnaea b o r e a l i s var. americana and various members of Vaccinium in p a r t l y open habitat s , often under pine trees. The Vaccinium have extensive woody root systems of tough roots as well as rhizomes i n the upper layers of the s o i l . Through t h i s three-dimensional network the b r i t t l e Geocaulon rhizome twists and turns randomly, branching at i n t e r v a l s and sending up a e r i a l shoots here and there. It s haustoria are attached to the roots and rhizomes of the Vaccinium, Linnaea and many other plants present. In more exposed s i t e s the association i s usually Vaccinium and Arctostaphylos but the same excavation problems remain. In shady spruce woods Geocaulon i s associated with Cornus canadensis in deep moss around the base of the trees. Under these conditions the moss can be peeled back to expose long lengths of the rhizome which can be removed r e l a t i v e l y e a s i l y with l i t t l e damage. Conditions are s i m i l a r on moss covered rocks. Generally Geocaulon a e r i a l shoots a r i s e from the rhizome at i r r e g u l a r i n t e r v a l s and the rhizome twists and turns ( F i g . 2 ) through the s o i l at random - 39 -Figure 1 Symbols as used in other Figures. Fig. 1.1 Aerial shoot axis with inflorescences, j-shoot axis, k-leaf, 1-peduncle of inflorescence, m-floral bract, n-flower pedicel, p-rudimentary f l o r a l bract, r-aborted or shrivelled flower, s-normal flower. Fig. 1.2 Actively growing end of a rhizome with the 'plumular hook' represented by a. Fig. 1.3 Section of rhizome, g, with i t s roots, d, bearing haustoria, c. Fig. 1.4 Aerial shoot of the current growing season with leaves, Fig. 1.5 Rhizome, g, with i t s short-shoot, f, bearing the dead remnants of aerial shoots of previous growing seasons, e. Fig. 1.6 Rhizome portion, g, which has been broken at B and is rotting away at R. The extent of the short-shoot is represented by f. Fig. 1.7 The dotted lines with arrows indicate that a l l tissue beyond the line is dead but s t i l l recognizable. The portion above h is a short-shoot system which no longer bears liv i n g aerial shoots. - 40 -- 41 -F i g u r e 2 An o l d rhizome i n i t s o r i g i n a l c o n f i g u r a t i o n as i t t w i s t e d and t u r n e d t h r o u g h t h e s o i l . Most o f the ends are r o t t i n g away, R, and t h e o n l y l i v i n g p o r t i o n b r o k e , B, and was l o s t . The heavy b l a c k areas a r e s w o l l e n p o r t i o n s o f t h e r hizome, one w i t h r o o t s a t t a c h e d . - 4 3 -so that there i s no apparent pattern to the shoots as they appear above the ground. Two nearby shoots may be adjacent or on f a r removed portions of the same rhizome; or they may be on d i f f e r e n t rhizomes. There are however several instances where a recognizable pattern of shoots i s seen above ground. One i s in the above mentioned moss covered Spruce woods. The rhizomes pass from hummock to hummock but a e r i a l shoots a r i s e only on the hummocks and not in the low spots between. Thus one sees l i t t l e c l u s t e r s of a e r i a l shoots around the bases of the trees but none i n between. In d r i e r pine forests one occasionally sees a s t r a i g h t l i n e of Geocaulon a e r i a l shoots which gradually become shorter towards one end. Upon excavation these are seen to a r i s e from moist r o t t i n g logs p a r t i a l l y buried in the s o i l or covered in deep moss. If the log i s in an appropriate state of decay i t can be pulled apart by hand to expose the one or more Geocaulon rhizomes growing lengthwise through i t . Branches are given o f f at i n t e r v a l s passing out of the log and disappearing in the surrounding Vaccinium roots and rhizpmes. A e r i a l shoots a r i s e at i n t e r v a l s , pass up through the log and into the a i r . The rhizome apex i s found at the end where the shorter a e r i a l shoots a r i s e . Roots of other plants are found i n these logs and h a u s t o r i a l connections can be found in and around the l o g . In the occasional chamber within the l o g , Geocaulon roots can be found which are i n contact with only the moist a i r of the chamber. On these roots, root h a i r s can be r e a d i l y seen and are e a s i l y c o l l e c t e d . These e n t i r e rhizomes (Figures 3 , 4 and 5 ) can be removed unbroken. Such logs were the source of a l l rhizome material i l l u s t r a t e d . - 44 -Figures 3, 4 and 5 Sketches of .rhizomes removed from rotting logs. They, are intact in that only roots were broken off when the rhizomes were removed from the logs. In Fig. 3 the rhizome has been cut into 6 pieces of near equal length and stretched out for ease of il l u s t r a t i o n . Ends with the same number are the points where the cuts were made. Section 0-1 is current years growth s t i l l covered with a whitish epidermis; 1-2 is in i t s second year and bears 1 year old short-shoots. The rhizome and short-shoots are progressively older the further back from the rhizome apex one samples them. Figures 4 and 5 show much the same phenomena. Fig. 5 is a direct continuation of Fig. 4. In Fig. 4, section 0-1 two lateral rhizomes can be seen on the two year old portion of the main rhizome. The point 'a' indicates the end of last years growth of the rhizome. - 46 -- 4 8 -A p a r a s i t i c v a s c u l a r plant may be defined as one which, in nature and under natural conditions, forms haustorial connections to other plants. The haustorium is at least partly composed of. liv i n g tissues and forms a physiological bridge between the host and the parasite. The xylem of host and parasite may or may not be. in contact, but nutrients and water are presumed to pass through this bridge in one or both directions. By this definition Geocaulon is a parasitic vascular plant. It is of the group known as root parasites since i t s haustoria attack the roots and rhizomes of the host plant in the s o i l . Geocaulon does, as do many other root parasites, attack and form haustoria-like organs on pieces of bark, dead wood, pebbles and other non-nutritive debris found in the s o i l . There appears to be a minimum size for a host since very small roots are not attacked but conifer roots up to 6 inches in diameter may have Geocaulon haustoria on them. Self-parasitism is common and Geocaulon appears to be quite unselective with regard to host. Similar observations have been made for Comandra (Piehl-1963) and Santalum (Barber-1906, 1907). In areas where Geocaulon is common many members of the Liliaceae and the Orchidaceae are also abundant but neither of these families have yet been found to serve as hosts for Geocaulon. According to Moss-1926 the following plants are hosts of Geocaulon. Pin.us banksiana Lamb. ; Ledum groenlandicum Oeder; Pinus murrayana Balf. (Pinus contorta Dougl. ex Loudon, ssp. l a t i f o l i a (Engelm. ex Watson) Critch-f i e l d ) ; Picea; Salix; Alnus; Betula; Ribes; Lonicera. The l i s t has been extended by the present writer, who, together with J. Nagy, is responsible for the following voucher specimens (in the University of Lethbridge Herbarium) with Geocaulon haustoria attached to their roots or - 49 -rhizomes. Equisetaceae Equisetum scirpoides Michx. (PDW 6 JN, #587). Equisetum arvense L. (PDW £ JN, #589). Betulaceae Betula occidentalis Hook. (PDW £ JN, #601) Salicaceae Populus trichocarpa T. £ G. ex Hook. (PDW 6 JN, #573). Cornaceae Cornus canadensis L. (PDW £ JN,.#598, #594; PDW £ PJW, #604) Ranunculaceae Thalictrum sp. (PDW & JN, #579). Rubiaceae Galium boreale L. (PDW S JN, #587) Leguminosae Hedysarum boreale (Richards) var. mackensii G. L. Hitch. (PDW 6 JN, #574). Ericaceae Ledum groenlandicum Oede'r (PDW £ PJW, #603) Arctostaphylos uva-ursi (L) Spreng. (PDW £ JN, #585) Rosaceae Dryas drummondii Richards (PDW £ JN, #595). Potentilla fruticosa L. (PDW £ JN, #593) Fragaria vesca L. (PDW £ PJW, #606). Fragaria virginiana Duchesne var. platypetala (Rydb.) Hall. (PDW £ JN, #597) - 50 -Gaprifoliaceae Lonicera involucrata (Richards) Banks (PDW £ JN, #578). Linnaea borealis L. var. americana (Forbes) Rehd. (PDW 6 JN, #599). Saxifragaceae Mitella nuda L. (PDW £ JN, #590). Scrophulariaceae Pedicularis bracteosa Benth. (PDW £ JN, #592). Castilleja miniata; Dougl, or possibly C_. rhexifolia Rydb. (PDW £ JN, #591.) Compositae Aster conspicuus Lindl. (PDW 6 JN, #580, #583). ? Aster occidentalis (Nutt) T. 6 G. (PDW & JN, #582). 1 Senecio canus Hook. (PDW 6 JN, #581). ? Senecio sp. (PDW £ JN, #584 ) Cyperaceae Carex sp. (PDW £ JN, #571). Gramineae Undetermined genus (PDW £ JN, #575). Cupressaceae Juniperus horizontalis Moench. (PDW £ JN, #596). Pinaceae Picea sp. (seedling), (PDW £ JN, #576). Several Voucher Specimens of Geocaulon lividum, aerial portions only, are also deposited in the herbarium. Other plants known as hosts from observations, and in part preserved in alcohol, but for which no herbarium - 51 -sheet is available include the following. Salix sp,; Lupinus sp.; Picea sp Vaccinium sp.; Geocaulon lividum (Richards) Fern.; Pinus contorta Dougl. ex Loudon ssp. l a t i f o l i a (Engelm. ex Watson) Gritchfield. In addition many other plants are suspected of being hosts but no documentation has yet been obtained. Many haustoria are collected on roots which are not those of known hosts but i t is virtually impossible to attach a name to an isolated portion of root. - 52 -The Rhizome The rhizome (Plate 2) i s the major portion of a Geocaulon plant and makes up by f a r i t s greatest mass. It and i t s attached short-shoots over-winter whereas the a e r i a l shoots die in the l a t e F a l l . The'rhizome grows in the topmost organic layers of the s o i l and appears to branch at random and spread throughout the s o i l along the l i n e s of least r e s i s tance. The t i p s of the rhizome (Plate 3) and i t s branches look l i k e the 'plumular hook' of a bean sprout. The t i p i s r e f l e x e d and the bend leads the way through the s o i l with the t i p and leaves following. Thus the apex i s protected and the enclosing leaves (Photograph 11) are not sheared o f f by passage through the s o i l . The rhizome expands markedly in diameter just d i s t a l to the bend and then gradually tapers down to the mature rhizome diameter. The whole t i p and attached leaves back to the point where the diameter decreases i s e t i o l a t e d and whitish. It i s covered with a smooth epidermis which bears stomata. The r e l a t i v e l y large leaves of the t i p are quite close together, but i n the bend region, as internodes elongate, the leaves become widely spaced, r a p i d l y wither, are r e f l e x e d and shorn o f f as they die and the rhizome elongates. These leaves and those of the short-shoots and subterranean portions of the a e r i a l shoots are l e s s expanded, l e s s d i s t i n c t l y veined, more clasping and have les s prominent p e t i o l a r regions than a e r i a l leaves. In the zone of decreasing diameter the whitish epidermis i s replaced by the brownish or yellowish-tan cork and the subsequent mature portions of the rhizome are a l l covered by t h i s cork. The cork region has s l i g h t l o n g i -t u d i n a l furrows i n i t and i s s l i g h t l y roughened. It bears l e a f scars, a x i l l a r y buds, short-shoots and adventitious roots. Older portions become quite rough - 5 3 -Plate 3 Rhizome apices. B has been cut at the bend but A shows the complete 'plumular hook'. The leaves, are lost only a short distance behind the bend. - 54 -- 55 -and dark brown in color, while dying ends are marked by blackening and withering (Fig. 2). Occasionally portions of older rhizomes were found t o be swollen (Fig. 2) from the normal 2 to 4 mm in diameter to 7 or 8 mm. These swollen areas were of irregular extent and not uniform in size. The swollen regions are far more positive for starch in the I/KI test than normal . rhizomes. In the rhizome apex there is a starch sheath around the bundles in the position of the endodermis, but further back in the cork-covered zone a l l of the cortical region is I/KI positive for starch. There are heavier deposits of starch just beneath the cork, in interfascicular regions of rhizomes with separate bundles, and around and in meristematic regions such as axillary buds and interfascicular cambium. The pith, xylem and cork are starch negative by the I/KI test. The paired prophylls of lateral appendages on the rhizome are whitish with pink tips when young. Cross sections of rhizomes show a thin layer of cork, a cork cambium zone and a cortical region with l i t t l e intercellular space. In the swollen rhizomes the extra diameter is made up of larger thinner-walled cortical c e l l s . Starch grains are visible as clusters in the cortical c e l l s , which also contain druses, usually in cells just outside the vascular system. Metcalf and Chalk-1955 mention the occurrence of crystals in the stem of Osyris and in the leaves of Osyris, Cervantesia and Leptomeria. Comandra also has such crystals (Piehl-1963). The walls of cortical cells also have: some apparently unique thickenings. These are discussed further in the chapter on Cell Wall Thickenings. A small pith of large irregular cells with thin walls forms the centre of the rhizome. This pith is generally surrounded by a eustele of about 8 vascular bundles (Photograph 4) but in many rhizomes interfascicular cambium formation has occurred between at least some adjacent vascular bundles. Thus - 5 6 -the bundles may be joined by xylem into groups of 2. or 3 or occasionally i n t o a complete ring. There is a tissue which looks very much like cortical tissue, and stains in the same manner. It is found between the xylem and the pith. This tissue is generally in contact with the cortex via the rays between the vascular bundles. The tissue layer is generally only 1 or 2 cells deep opposite.the xylem of a vascular bundle and a l i t t l e deeper between bundles. The tissue is positive for starch in the I/KI test and also contains the above mentioned c e l l wall thickenings. This tissue appears to be unique as i t is not described in standard anatomy texts. In most rhizomes the bundles (Photograph 4) consist of a matrix of small xylem cells with a few metaxylem vessel elements embedded in them. The phloem region is external to the xylem (collateral bundle) and of about equal volume. Fibre caps external to the phloem are found in many rhizomes. There is a great deal of variation in the appearance of the vascular system from rhizome to rhizome. The endodermal layer is d i f f i c u l t to discern. It is evident as the starch sheath in rhizome apices but i t is a questionable entity in the mature rhizome where no casparian strips can be found. The xylem of the rhizome follows the standard pattern with protoxylem vessel elements spirally or annularly thickened and jnetaxylem vessel elements pitted or scalariform. Rhizomes are endarch and polyarch in their development. Figure 6 shows some of the ways in which the vascular system of the rhizome branches. 'A' is a forking of the rhizome into two. The side with the gradual curve is the basal or oldest portion. These branches generally arise near the apex of the rhizome in the region before cork formation occurs. In 'B' the rhizome is giving rise to root vascular traces. The vascular supply - 5 7 -F i g u r e 6 The v a r i o u s ways i n which t h e v a s c u l a r system o f t h e rhizome b r a n c h e s . A , rhizome b r a n c h i n g i n t o two, o l d e s t end a t t h e l e f t . B, v a s c u l a r s u p p l y t o two r o o t s and r o o t a b s c i s s i o n zone. C, f o l i a r and a x i l l a r y bud t r a c e s i n t h e mature p o r t i o n o f t h e rhizome. D, f o l i a r t r a c e s a t t h e rhizome apex p r i o r t o c o r k f o r m a t i o n . Drawn from c l e a r e d whole organs. - 5 8 -fi g . 6 - 59 -is much smaller in diameter, as are the roots, and the root xylem is perpendicular to that of the rhizome. If a root arise on a rhizome with separate vascular bundles the root vascular supply comes from only 1 or 2 bundles. The rest of the root tissues arise separately from the meristematic pericyclic cells and are hot continuous with the rhizome" tissues. There appears to''be an abscission zone of sorts at the root/rhizome junction where roots break off cleanly leaving a bump dh the rhizome; 'C,' is near the apex of the rhizome but in the cork-covered region. The vascular system is generally not in discrete strands in this region but closed into a more or less continuous ring. The bud trace goes almost directly to the vascular cylinder which bulges out to meet i t . The leaf trace runs parallel to the vascular cylinder for a short distance before merging. 'D' is well out towards the apex of the rhizome in the epidermal region. There are no macroscopically visible buds in this region. The vascular system is in discrete bundles here and leaf traces simply add 1 more bundle. These bundles join into a vascular cylinder at about the point where cork formation occurs. There is a sudden increase in diameter< at this point which is well marked by the change from smooth white epidermis to rough tan cork. - 60 -The Short-Shoot Short-shoots are perennial and are borne on the rhizome at intervals along i t s length. It is on these short-shoots (Plates 2 and 4) that the aerial or leafy, flowering shoots are produced each Spring. Each.year new short-shoots with aerial branches are produced at the tip of the rhizome and new aerial shoots grow from existing short-shoots further back on the rhizome. Early in the season one may see a line of progressively shorter shoots arising towards the apex of a rhizome while nearly uniform shoots arise further back on older portions of the rhizome. Even in the f i r s t year the demarkation between what is to become perennial short-shoot and what is to become annual aerial shoot is readily visible (Plate 5). The basal short-shoot portion is a light tan in color, has smallish unexpanded leaves and has slight longitudinal furrows in the cork. It is completely subterr-anean. The base of the aerial shoot is whitish, has larger more prominently veined leaves and has smooth white epidermis. In the 1 or 2 internode transition zone there is often a region of horizontal folding (Plate 5) of the outer tissues. In the winter the aerial shoot portion dies and most of i t disintegrates; however for some years a few strands of dark stringy vascular bundles and fibres (Plates 4 and 6) remain attached to the short-shoot and indicate the number and location of previous aerial shoots. In the second year a lateral bud on the short-shoot develops into the new aerial shoot. Up to three aerial shoots per year may develop on each short-shoot but one or two is more common. In the f i r s t year only one aerial shoot i s produced since i t is a direct continuation of the axis of the short-shoot. The short-shoot is long-lived and produces aerial shoots for many years (Plate 6). These aerial shoots - 61 -P l a t e 4 Three s h o r t - s h o o t s w i t h s o i l l e v e l i n d i c a t e d by t h e h o r i z o n t a l l i n e . A, f i r s t y e a r o f growth w i t h t e r m i n a l a e r i a l s h o o t . B and C, second y e a r o f growth. The f i r s t y e a r s t e r m i n a l a e r i a l s hoot has d i e d and l a t e r a l a e r i a l s h o o t s have grown t h i s s e a s o n . Note t h e d i f f e r e n c e between the a e r i a l shoot buds and th e rhizome a p i c e s i n P l a t e 3 . . - 62 - 63 -Plate 5 Details of the short-shoot/aerial shoot transition zone. D, f i r s t year short-shoot attached to a section of rhizome. E, detailed view of the transition region of D. The lower short-shoot portion is covered by a tan cork with vertical striations while the upper aerial shoot portion has a smooth whitish epidermis. Horizontal folds occur at the junction. F, another transition zone showing the change in f o l i a r appendages over the transition zone. G, an aerial shoot bud on a two year old short-shoot. Note the extension of the short-shoot into the base of the previous aerial shoot. A, B and C indicate where sections have been cut for microscopic examination of the vascular system. Photo-graphs .1, 2 and 3 are taken from these sections. - 64 -- 65 -P l a t e 6 One o f t h e o l d e s t and most complex s h o r t - s h o o t s found. The numbers r e p r e s e n t the r e l a t i v e ages o f t h e b r a n c h e s . Number 1 was t h e f i r s t t o a r i s e and number 9 i s th e c u r r e n t a e r i a l s h o o t . Depending upon how many a e r i a l s h o o t s a r o s e each growing season t h e whole system c o u l d be from 5 t o 9 y e a r s o l d , assuming a t l e a s t one b r a n c h a r o s e each y e a r . The most l i k e l y age e s t i m a t e i s 7 o r 8 y e a r s . - 67 -may not die a l l the way back to their point of insertion on the short-shoot. When this occurs the short-shoot axis may be extended into this aerial shoot base which may now produce aerial shoots in succeeding years. New aerial shoots are almost always produced at successively lower nodes on any portion of the short-shoot. The short-shoot has thus an int r i n s i c a l l y limited lifespan determined by the number of nodes below the cork/epidermis boundary in the f i r s t year. Extra time can be gained by extension into the base of aerial shoots. A sample of 80 short-shoots was examined and categorized as indicated in Figures 7, 8 and 9. The most common types were B and C with 14 specimens of each. This was followed by A, E and D with 9, 7 and 6 specimens respectively. G and I were represented by 3, and F , J , L and V by 2 specimens. The others were one-of-a-kind. From these and other short-shoots and the arrangement of these short-shoots on the rhizome, the following rules for determining the age of a short-shoot and the portion of rhizome bearing i t were determined. Criteria for determining the age of short-shoots - in i t s f i r s t year a rhizome tip bears' no short-shoots. - in it s second year a rhizome portion bears an unbranched short-shoot with 1 terminal aerial shoot. - in succeeding years a rhizome portion bears more complex short-shoots with black strands and lateral aerial shoots. - in it s f i r s t year a short-shoot bears only 1 terminal aerial shoot. - in it s second year a short-shoot bears one terminal bundle of black strands and 1 or more lateral aerial shoots. - in succeeding years a short-shoot bears a terminal and some lateral black strands and several lateral aerial shoots. - 68 -Figures 7, 8 and 9 Sketches of short sections of rhizomes'with attached short-shoots. These form a representative sample of the types of short-shoots which were met with in Geocaulon. . Leafless branches are the black strands of previous years aerial shoots; leafy branches are the aerial shoots of the current years growth. 1 year old-A; 2 years old-B, D and K; 3 years old-C, E, F, J and M; 3 or 4 years old-G, L, N and P; 4 years old-H, I and S; 4 or 5 years old-O, Q, T and V; 5 years old-W and X; 5 or.6 years old-U; 5 to 7 years old-R; 4 to 8 years old-Y; 6 to 9 years old-Z. Plates 1, 3, 4, 5 and 6 also i l l u s t r a t e short-shoots. - 7 0 -- 71 -- 7 2 -P l a t e 7 S h o r t - s h o o t from 6 t o 8 y e a r s o l d . S e c t i o n s were cu t a t t h e zones marked D through I f o r . a m i c r o s c o p i c e x a m i n a t i o n o f t h e v a s c u l a r system. Photographs 5 t h r o u g h 10 c o r r e s p o n d t o s e c t i o n s D through I. The v a s c u l a r system changes from e u s t e l a r i n t h e rhizome t h r o u g h a c o n t i n u o u s c y l i n d e r i n t h e s h o r t - s h o o t and back t o a e u s t e l e i n the a e r i a l s h o o t . - 74 -- when s e v e r a l a e r i a l . s h o o t s a r i s e i n one y e a r from the. same s h o r t - s h o o t , the f i r s t t o d evelop i s on a h i g h e r node than s u c c e e d i n g ones, which f o l l o w on s u c c e s s i v e l y lower nodes. The uppermost a v a i l a b l e node need not be the f i r s t t o d e v e l o p , n o r a r e s u c c e s s i v e a e r i a l s h o o t s on c o n t i g u o u s nodes; but a node which i s s k i p p e d i s not a v a i l a b l e f o r f u t u r e development.' - an a e r i a l s hoot need not d i e a l l the way back t o i t s p o i n t o f i n s e r t i o n on t h e s h o r t - s h o o t . When t h i s o c c u r s the s h o r t - s h o o t a x i s i s extended i n t o the base of the a e r i a l s h o o t . Now s u c c e s s i v e development o f lower nodes as a e r i a l s h o o t s b e g i n s a g a i n i n t h e new p o r t i o n o f the s h o r t - s h o o t . - i t i s presumed t h a t a t l e a s t one a e r i a l shoot d e v e l o p s , on an e s t a b l i s h e d s h o r t - s h o o t , every y e a r . T h i s has not been proven. I f n o t t r u e then the ages g i v e n r e p r e s e n t t h e number of growing seasons i n which t h e s h o r t - s h o o t has been a c t i v e r a t h e r than i t s c h r o n o l o g i c a l age. P l a t e s 6 and 7 a r e examples o f o l d and q u i t e complex s h o r t - s h o o t s whose age has been det e r m i n e d u s i n g t h e s e c r i t e r i a . F i g u r e s 7, 8 and 9 i l l u s t r a t e a sample o f s h o r t - s h o o t s w i t h ages i n d i c a t e d . The rhizome g e n e r a l l y has a e u s t e l e o r may have some i n t e r f a s c i c u l a r c a m b i a l development between some a d j a c e n t b u n d l e s ; i t seldom has a complete v a s c u l a r c y l i n d e r . The a e r i a l s h o o t s a r e a l l e u s t e l a r . S h o r t - s h o o t s g r a d u a l l y change from t h e c o n d i t i o n o f t h e rhizome t o a complete v a s c u l a r c y l i n d e r and then back t o t h e e u s t e l a r c o n d i t i o n o f t h e a e r i a l s h o o t s . Photographs o f t h e s e changes, numbers 5 through 10, c o r r e s p o n d t o t h e i n d i c a t e d s e c t i o n s D t h r o u g h I on P l a t e 7. A s i m i l a r but a b b r e v i a t e d s e r i e s i s i n d i c a t e d by s e c t i o n s A, B and C on P l a t e 5 and i s i l l u s t r a t e d by Photographs 1, 2 and 3 . - 7 5 -A c r o s s s e c t i o n o f a s h o r t - s h o o t r e v e a l s the t h i n p e r i p h e r a l l a y e r o f cork u n d e r l a i n by a cork cambium. W i t h i n t h e cork i s a zone of c l o s e l y packed rounded c o r t i c a l c e l l s . The c e n t r e o f t h e s h o r t - s h o o t i s o c c u p i e d by p i t h w i t h i r r e g u l a r l y shaped c e l l s . Around the p i t h i s t h e v a s c u l a r bundle zone. In young s h o r t - s h o o t s i t may be composed o f m o s t l y d i s c r e t e b u n d l e s but i t i s u s u a l l y , and c e r t a i n l y so i n o l d e r s h o r t - s h o o t s , i n the form o f a more o r l e s s complete v a s c u l a r c y l i n d e r . The p o s i t i o n s o f the o r i g i n a l bundles a r e marked by th e appearance of f i b r e caps at t h e c o r t e x / p h l o e m boundary. The same t i s s u e mentioned i n the. c h a p t e r on t h e rhizome o c c u r s between the p i t h and the xylem of t h e b u n d l e s . - 76 -The Aerial Shoot Aerial shoot buds are not at a l l like the 'plumular hooked' rhizome apices. Aerial shoot buds are erect and leafy and remain so as they grow. Buds removed from s t i l l frozen, snow-covered ground in early May have macroscopically visible inflorescences in the leaf axils. Aerial shoot buds arise on the short-shoot (Plates 2, 4 and 5) as lateral appendages. Only the f i r s t aerial shoot is terminal on the short-shoot. The basal portion of an aerial shoot (Plate 8) begins with several very short internodes such that the cataphylls form an involucre at the aerial shoot/short-shoot junction. This zone is followed by one where the nodes gradually elongate and the closely appressed bracts sheath the stem. Above, this, but s t i l l below ground lev e l , the appendages become more lea f - l i k e , : have axillary buds and stand out from the stem, though s t i l l pointing upwards. Above the ground the internodes become shorter, leaves stand out from the stem at increasing angles, the leaf blade increases in, area and the axis becomes green. In the zone of the largest leaves (Plate 1) the petiole to stem angle is nearly 90 degrees but the petiole to blade angle causes the blade to point upwards a l i t t l e . , This zone is about one-half way up the stem and is the zone where the inflorescences are produced. Above this region the leaves become progressively smaller and make smaller angles with the axis. In cross section the aerial shoot shows a thin cuticle covering the rounded epidermal cells which are barely distinguishable from the underlying narrow zone of cortex. The pith may have a cavity in the centre. Vascular bundles are about one half xylem and one half cambium and phloem. The bundles (Photograph 10) may have extensive fibre caps. The aerial shoot of Geocaulon - 7 7 -P l a t e 8 S u b t e r r a n e a n p o r t i o n o f a e r i a l s h o o t . A, p o i n t o f i n s e r t i o n o f t h e a e r i a l shoot on t h e s h o r t - s h o o t . The f i r s t few i n t e r n o d e s a r e v e r y s h o r t g i v i n g t h e i m p r e s s i o n of an i n v o l u c r e o f l e a f y b r a c t s a t t h e j u n c t i o n . The n e x t few i n t e r n o d e s a r e s h o r t w i t h b l u n t l e a v e s c l o s e l y a p p r e s s e d t o the stem a x i s . B, i s a l i t t l e h i g h e r w i t h l o n g e r l e a v e s and i n t e r n o d e s and d e c r e a s i n g stem di a m e t e r . C, i s a l i t t l e above B and has t h e l e a v e s removed t o show a x i l l a r y buds i n the l e a f a x i l s . - 79 -i s e u s t e l a r w h i l e i n Comandra ( P i e h l - 1 9 6 3 ; P l a t e 1-A) the xylem o f the a e r i a l shoot i s i n a c o n t i n u o u s c y l i n d e r . In t h e Geocaulon a e r i a l shoot t h e c o r t e x 'rays and t i s s u e between t h e p i t h and xylem o f t h e bundles a r e a l l s t a r c h p o s i t i v e i n t h e I/KI t e s t . The o t h e r t i s s u e s a r e n e g a t i v e f o r s t a r c h . - 80 -The Leaf The leaf (Plate 1) is ovate to obovate and reaches a maximum length of about 5 cm, though most leaves are about 2 to 3 cm long. In shaded areas the leaf is thin and flaccid and du l l - or olive-green. In exposed areas the leaves are much smaller and firmer and also have varying amounts of red coloration. Leaf margins are entire and there are no epidermal appendages. Veins are somewhat raised on the lower surface of the leaf. The leaf base is decumbent and the petiole very short. In cross section the petiole has a f l a t adaxial surface and an inverted-bell-shaped abaxial surface with flaring upturned wings near the insertion of the leaf blade. Stomata are completely lacking on the upper epidermis (Plate 10) whereas they are found on the lower epidermis (Plate 11) with a normal density of about 50/square mm. Leaf epidermal cells above vascular bundles are rounded as are the cortical cells between the bundle and the epidermis. Epidermal cells over the major veins are f l a t and elongate in the direction of the longitudinal axis of the vein. Other epidermal cells are f l a t and paving-stone-like. They have an irregular outline and have some tendency to inter-lock. On the lower epidermis the longitudinal axes of most stomata are roughly perpendicular to the longitudinal axis of the leaf. When a whole leaf i s treated with I/KI only the guard cells stain appreciably. The ends of the guard cells stain more intensely than the centres showing starch localization even within the guard cells. The inside of the leaf contains rounded cortical parenchyma. On the lower surface of the blade the sub-epidermal cells are wider and longer than deep, while the upper sub-epidermal cells are palisade-like. Particularly in the lower portions of the leaf there is a large proportion of sub-stomatal - 81 -P l a t e 9 V a s c u l a r p a t t e r n i n a c l e a r e d Geocaulon l e a f . Major v e i n s a r e connected i n a r e t i c u l a t e p a t t e r n and the s m a l l e s t v e i n l e t s end b l i n d l y . There i s a f r i n g e o f s m a l l v e i n l e t s around t h e l e a f margin. - 8 3 -P l a t e 10 P o r t i o n s o f t h e upper e p i d e r m i s o f Geocaulon l e a v e s . Only c e l l w a l l s and n u c l e i a r e i n d i c a t e d . There a r e no stomata on t h e upper e p i d e r m i s . E x c e p t f o r a tendency t o be i n p a i r s t h e r e does n o t seem t o be any o r d e r l y p a t t e r n t o t h e arrangement o f t h e e p i d e r m a l c e l l s . - 8 5 -P l a t e 11 P o r t i o n s o f t h e lower e p i d e r m i s o f Geocaulon l e a v e s . Only c e l l w a l l s and n u c l e i a r e shown. In a l l c a ses t h e p a i r o f guard c e l l s f o r m i n g a stoma have t h e i r n u c l e i a t t h e same ends o f t h e i r c e l l s . There i s a s l i g h t tendency f o r stomata t o be a l i g n e d i n more o r l e s s t h e same d i r e c t i o n i . e . , r o u g h l y p e r p e n d i c u l a r to. t h e l o n g i t u d i n a l a x i s o f t h e l e a f . - 8 7 -and intercellular space. There is only one vascular bundle in the petiole. In transverse section i t is in the form of an arc opening towards the adaxial surface. The vascular pattern in a typical leaf is indicated in Plate 9. As is usual in leaf bundles the xylem is on the adaxial surface. In the leaf blade larger veins are connected in a closed reticulate network. The smallest veinlets end blindly in a small group of parallel xylem elements. The vascular system is fringed by a row of small veinlets around the leaf margin. Throughout i t s range Geocaulon is occasionally infected with a virus (Moss-1926) causing yellow or sometimes red veinal areas. In extreme cases the leaf is ch'lorotic, small, distorted in shape, wavy margined and often s p l i t and cut. These plants are often only several inches high but s t i l l flower and set f r u i t . The leaves of Geocaulon also serve as an alternate host for Cronartium comandrae Peck, the comandra bli s t e r rust (Piehl-1963, Powell-1968) as well as other fungal and bacterial parasites which often alter the appearance of the leaf. Comandra is also a host for Cronartium comandrae Peck (Piehl-1963).. ' - 88 -The Inflorescence The inflorescence-bearing nodes (Plate 1) are those about one half way up the aerial shoot, in the region where maximum leaf area is attained. A l l the inflorescences of a plant are generally on contiguous nodes and flowering is centripetal. The peduncles in the axils of the leaves generally bear a 3-flowered cymule or dichasium though 2- and 4-flowered inflorescences are not rare. This basic inflorescence unit (Plate 12) may however be altered (Figures 10, 11 and 12) almost beyond recognition. In the southwestern part of the range the commonest condition is three flowering nodes per plant, while in the east there is only one flowering node per plant. In a Geocaulon inflorescence unit the three flowers are a l l in the same plane such that one of the flowers is adjacent to the stem and the one on the other side of, the central flower is adjacent to the leaf in whose a x i l the peduncle arises. Thus one can differentiate between inside and outside , flowers with respect to the stem axis or possibly of adaxial and abaxial flowers respectively. This leads to the possibility of mirror image inflorescence units. Several such pairs are illustrated in Figure 10, As indicated in Plate 12 the fourth flower in a four-flowered inflorescence is not in the same plane as the other three. Bracts may or may not be present at the base of the pedicels of the flowers and may also occur at other places on the pedicel or on the peduncle. Bracts vary from a,leafy organ to a mere vestigal flap of tissue visible only through the dissecting microscope. In place of many large normal flowers there is often a small, shrivelled, blackish flower which aborts early. Usually only the central flower matures into a f r u i t and the laterals, even i f normal, abort.- This is the common condition in the eastern and southwestern - 8 9 -P l a t e 12 T h r e e , i n f l o r e s c e n c e s which can be c o n s i d e r e d cymules o r m o d i f i e d d i c h a s i a . The c e n t r a l 3 - f l o w e r e d i n f l o r -e s c ence i s the 'normal' c o n d i t i o n w i t h a c e n t r a l p r e c o c i o u s f l o w e r and 2 l a t e r a l s i n t h e a x i l s o f b r a c t s . The 2- and 4 - f l o w e r e d i n f l o r e s c e n c e s a r e n o t as common but are n o t r a r e . I - 91 -Figures 10, 11 and 12 Variations on the basic inflorescence plan of Geocaulon. The basic unit i s a terminate cyme or dichasium. When a c e n t r a l flower i s present i t matures f i r s t and i s usually the only one to form a f r u i t , the l a t e r a l s generally aborting. The open c i r c l e s are normal flowers, the dots abortive or withered flowers. This c o l l e c t i o n was taken from one s i t e and few examples are represented by more than one specimen. There are many possible permutations of normal flowers, abortive flowers and bracts. - 94 -PD, W. - 95 -p a r t o f t h e range. In the n o r t h w e s t e r n p a r t o f t h e range one o r both l a t e r a l s commonly, mature. One p e d i c e l i n the a x i l o f a b r a c t was f o u n d i n s e r t e d d i r e c t l y i n t h e a x i l o f a l e a f and not on a p e d u n c l e . Geocaulon stems do not n o r m a l l y branch but s e v e r a l i n s t a n c e s were found where v e r y s h o r t l e a f y s h o o t s a r o s e i n l e a f a x i l s i n t h e i n f l o r e s c e n c e zone. S e v e r a l i n f l o r e s c e n c e s were c l e a r e d i n o r d e r t o t r a c e t h e v a s c u l a r s t r a n d s from the p e d u n c l e t h r o u g h t o the t e p a l s . Four o f t h e s e v a s c u l a r s k e l e t o n s ( F i g . 13) were drawn i n diagrammatic u n f o l d e d form. - 96 -F i g u r e 13 Diagrammatic ' u n f o l d e d ' r e p r e s e n t a t i o n o f t h e v a s c u l a r s k e l e t o n o f f o u r c l e a r e d i n f l o r e s c e n c e u n i t s . The drawings b e g i n i n t h e p e d u n c l e s h o r t l y below t h e p o i n t where the p e d i c e l s a r i s e and c o n t i n u e i n t o t h e base o f each p e r i a n t h member. The t e p a l s a r e numbered i n s e r i a l o r d e r around t h e f l o w e r . - 98 -The Flower Macroscopic flower buds are visible in Geocaulon aerial shoot buds before the latter have emerged from the ground. In the early spring when the ground is s t i l l covered with snow macroscopic flower buds can be found by digging up Geocaulon short-shoots with aerial shoot buds on them. In northern regions flowering may not occur u n t i l August but in much of i t s range Geocaulon flowers in June and early July. The flowers (Plates 12 and 13) are small and inconspicuous. When fu l l y open the flower may be 2 mm deep and 4 mm wide. The perianth is greenish and ranges from somewhat campanulate to quite rotate. Geocaulon tepals are nearly equillateral-triangular. There are generally 5, occasionally 4, tepals in a whorl. The base of the perianth tube is adnate to the ovary and the whole tube is lined inside by a disc whose lobes alternate with the tepals. There are as many stamens as there are tepals or perianth lobes; the filaments, which are short and ligulate, are inserted on the inside face of the perianth lobes, between the lobes of the disc. The anthers are ovate, 2-celled, and dehisce longitudinally and introrsely, at or before the opening of the flower. At the base of the filament, where i t is inserted on the perianth, a bundle of hairs arises, follows along the abaxial surface of the filament and between the anthers along the connective, to protrude from the stamen towards the stigma. These hairs are common to. many other Santalaceae. The style i s short, massive and cylindrical. It is topped by a slightly capitate and irregularly shaped stigma. There is a 1-celled inferior ovary, comprised of 5 carpels, with 2 to 4 ovules near the apex of a long coiled placental column as in Thesium and most other Santalaceae. The venation pattern in 3 cleared tepals (Plate 14) clearly shows the - 99 -P l a t e 13 A, view from above onto an unopened bud. B, view o f an open mature f l o w e r from above and t o one s i d e . The b u n d l e o f h a i r s on t h e a b a x i a l s u r f a c e o f t h e stamens and t h e i n v e s t i n g d i s c a r e v i s i b l e . C, l a t e r a l view o f a young Geocaulon f r u i t w i t h i t s crown o f t e p a l s . The s c a r where the l a t e r a l f l o w e r p e d i c e l was a t t a c h e d i s v i s i b l e . - 100 -- 101 -di ITer'once between . te'pal and leaf (Plate 9) venation. There is no fringe ol ymall veinlets around the margin of a tepal. The veins in a tepal do not form a reticulate network as do those of the leaf. In Geocaulon flowers from the southwestern part of the range there is no apparent difference between lateral and central flowers of a dichasium. The external morphological characteristics and the internal anatomical ones are the same. Apart from the slightly greater size of central flowers, which open f i r s t , one can not t e l l them apart. Both laterals and centrals have normal-appearing pollen and both have ovaries with well developed placental columns and embryo sacs. Embryos have been observed in both lateral and central flowers and except in the eastern part of the range lateral flowers do form f r u i t s . The original generic description of Geocaulon (Fernald-1928) states that "the perfect flower (is) central in the umbel with the (lateral flowers) staminate". Fernald also writes "drupes solitary (rarely 2)". If by this he means that only one drupe is produced per individual umbel then the phrase rarely 2 contradicts the statement that lateral flowers are staminate. If he means that only one f r u i t matures per aerial shoot this is refuted by the observation of multiple-fruited plants. I have not observed l i v i n g material in the f i e l d in the Gaspe and northeastern United States region which was Fernald's 'home region'.. I have examined fruiting plants In the f i e l d along the north shore of Lake Superior and here there were very few exceptions to the condition of one inflorescence per aerial shoot with only one f r u i t per inflorescence. These exceptions were several plants with two inflore-scences per plant but s t i l l only one f r u i t per inflorescence. Moving west to southeastern Manitoba one finds more of these exceptions,.and rarely, two - 102 -P l a t e 14 V e n a t i o n p a t t e r n i n t h r e e c l e a r e d t e p a l s above t h e p o i n t o f s e p a r a t i o n from the f l o r a l t u be. A d a x i a l views w i t h a p o r t i o n o f t h e a n t h e r s v i s i b l e a t t h e bottom o f each t e p a l . - 103 -- 104 -f r u i t s p e r i n f l o r e s c e n c e . In the southwestern p a r t o f t h e range t h e r e are up t o 6 i n f l o r e s c e n c e s p e r p l a n t w i t h s p o r a d i c cases o f more than one f r u i t p e r i n f l o r e s c e n c e . In the n o r t h w e s t e r n p a r t o f t h e range m u l t i p l e i n f l o r e -s c e n c e s and m u l t i p l e f r u i t s p e r i n f l o r e s c e n c e a r e common. Thus i t appears t h a t F e r n a l d made h i s d e s c r i p t i o n on the b a s i s o f l o c a l m a t e r i a l o n l y a l t h o u g h he c i t e s specimens from o t h e r p a r t s o f t h e range. - 105 -The Fruit The fruit of Geocaulon (Plate 13) has been called a false drupe (Fernald-1950). However a false drupe (Gray-1967) is nut-like with a fleshy lower part. A true drupe has a stony endocarp (Gray-1967) but Geocaulon has a stony mesocarp (Photographs 13 and 16) much the same as Exocarpos (Ram-1959A), Leptomeria (Ram-1959B), Osyris (Joshi-1960) and Comandra (Ram-1957). These fruits with hard mesocarps in the Santalaceae• have been called huts (Ram-1959A, 1959B; Joshi-1960) but a nut has a dry pericarp (Cronquist-1961). The term spurious drupe (Gray-1967) is also offered for fleshy f r u i t s , not true drupes, but drupe-like in that they enclose a stone. In view of this confusion in nomenclature for which the arbitrary nature of f r u i t c l a s s i f i c a -tion is partly responsible, the f r u i t of Geocaulon w i l l be called ' f r u i t ' . The flowers and young fruits are erect on the pedicel but large mature frui t s are pendant. A ripe f r u i t is about 1 cm in diameter. The ripe exocarp is bright red or orange and edible but tasteless. The fru i t s are crowned by the remains of the f l o r a l disc and perianth members when young but this crown generally f a l l s in older f r u i t s . The layers of a mature f r u i t from the outside towards the centre are: epidermis, fleshy exocarp, stony mesocarp, crushed endocarp, endosperm and embryo. There is a firm epidermal layer around a f r u i t with one or two layers of small cells beneath. Next is a deep layer of large thin-walled c e l l s , the exocarp (Photograph 14) which is composed of spherical cells next to the epidermis and columnar cells next to the mesocarp. Only the exocarp cells of the f r u i t are starch positive in the I/KI test. The mesocarp layer (Photograph 16) is about 10 to 15 cells deep and is very hard and dense, with small, thick-walled c e l l s . The mesocarp has grooves in i t s outer surface in - 106 -Figure A young f r u i t figuratively cut between tepals 1 and 5 and then opened out f l a t . The lower ends of the vascular system in the drawing are in the base of the f r u i t where i t joins the peduncle. Strands labelled 's' supply the stamens; strands labelled 'x' nearly meet in the centre of the f r u i t just beneath the style. The other, strands are on the surface of the mesocarp or within the tepals. The 'x' traces compare to those labelled "X" by Smith and Smith-1943.' - 107 -- 108 -which vascular bundles l i e . There are also tubes within the mesocarp, f i l l e d with vascular tissue. Internal to the mesocarp is the remains of the crushed endocarp sandwiched between hard mesocarp and massive endosperm. The entire center of the f r u i t is f i l l e d with endosperm within which is the embryo. The embryo (Photograph 15) is embedded in the endosperm at a point just beneath the crown of perianth members. The embryo is displaced well off center and is only a few cells inside the endosperm/endocarp boundary. In the young f r u i t wall (Photograph 13) the exocarp cells have not yet elongated, the mesocarp cells are not yet thick-walled and the endocarp has not yet been crushed or absorbed by the endosperm. Several fruits were cleared to render them transparent, at least to the mesocarp surface. Using these the vascular strands on the mesocarp could be traced. For one of these cleared fruits a diagrammatic unfolded vascular pattern was drawn (Fig. 14). The numbers indicate the s e r i a l order around the f r u i t . Traces labelled 's' go to the stamens and 'x' traces meet in the center of the f r u i t beneath the style. These latter compare to the 'X' traces of Smith and Smith-1943. - 109 -The Root Roots are q u i t e s m a l l ; up t o 1 mm n e a r some l a r g e h a u s t o r i a but u s u a l l y o n l y up t o 0.5 mm i n d i a m e t e r . They a r i s e from the rhizome a t a p p a r e n t l y random l o c a t i o n s and have not been found more than about 1 dm l o n g . There i s a p o o r l y d e f i n e d r o o t cap and a w e l l d e v e l o p e d r o o t h a i r zone. The l a t t e r i s b e s t seen on r o o t s which have d e v e l o p e d w i t h i n m o i s t c a v i t i e s i n r o t t i n g l o g s where t h e r e i s no c o n t a c t between the r o o t and the s u b s t r a t e . These are th e o n l y c o n d i t i o n s which a l l o w one t o c o l l e c t i n t a c t r o o t s . Roots are w h i t i s h when young becoming brown w i t h age. They break r e a d i l y from the rhizome a t w e l l d e f i n e d a r e a s ( F i g . 6, B) l e a v i n g a c h a r a c t e r i s t i c s c a r . Root t r a c e s a r i s e p e r p e n d i c u l a r t o t h e rhizome v a s c u l a r system i n c o n t r a s t t o f o l i a r and rhizome t r a c e s . Root t i p s a r e o f t e n s w o l l e n , b ulbous and d a r k e r than the r e s t o f t h e r o o t . H a u s t o r i a a r e i n i t i a t e d a t i n t e r v a l s a l o n g t h e r o o t as l a t e r a l organs ... h a v i n g complete t i s s u e c o n t i n u i t y w i t h t h e r o o t . When branch r o o t s a r i s e ( endogenously) t h e y do so by t h e f o r m a t i o n o f a primordium o f m e r i s t e m a t i c c e l l s from the p e r i c y c l i c r e g i o n o f the r o o t . T h i s group o f c e l l s m u l t i p l i e s and e l o n g a t e s , p u s h i n g t h r o u g h t h e s u r r o u n d i n g c o r t e x and e p i d e r m i s , p e n e t r a t i n g and c r u s h i n g t h e s e t i s s u e s , as t h e new r o o t a x i s emerges. In c r o s s s e c t i o n a r o o t i s bounded e x t e r n a l l y by a t h i n and p o o r l y d e f i n e d e p i d e r m i s a t a p i c a l r e g i o n s . Between h a u s t o r i a and t h e rhizome most r o o t s have a l a y e r o f cork as t h e o u t e r l i m i t i n g t i s s u e . In the a p i c a l r e g i o n s most of a r o o t ' s d i a m e t e r i s t a k e n up by t h i n - w a l l e d c o r t e x almost d e v o i d o f i n t e r -c e l l u l a r s p a c e s , but i n r e g i o n s b e a r i n g h a u s t o r i a the c o r t e x i s o n l y a few c e l l ' s deep. These c o r t e x c e l l s a r e l a r g e and rounded and c o n t a i n m a t e r i a l which does n o t become e x t r a c t e d i n t h e normal embedding p r o c e d u r e s . Towards - 110 -the apex the v a s c u l a r system c o n s i s t s o n l y o f a few s t r a n d s o f a n n u l a r o r s p i r a l l y - t h i c k e n e d p r o t o x y l e m and a few s c a l a r i f o r m o r p i t t e d metaxylem v e s s e l elements. Between a l a r g e h a u s t o r i u m and the rhizome t h e r e may be a massive secondary development o f l a r g e v e s s e l elements (Photograph 12) i n a p r o t o s t e l e which o c c u p i e s most o f t h e r o o t s d i a m e t e r . Roots a r e e x a r c h and d i a r c h . The endodermis, i n a p i c a l p o r t i o n s o f the r o o t and r e g i o n s where h a u s t o r i a a r e s t i l l b e i n g formed, has t h e e n t i r e i n n e r t a n g e n t i a l w a l l s u b e r i z e d ( E s a u -1965, pages 374, 491) and no e v i d e n c e o f c a s p a r i a n s t r i p s can be found. The r a d i a l w a l l s o f the endodermis may break down t o l e a v e a gap between t h e c o r t e x and the i n n e r t a n g e n t i a l endodermal w a l l . T h i s i s most c l e a r l y seen a t t h e apex o f some h a u s t o r i a where t h e i n n e r v a s c u l a r c o r e o f the r o o t p a s s e s through the a p i c a l c o r t i c a l r e g i o n o f t h e hau s t o r i u m . Only t h e c o r t i c a l p o r t i o n s o f the r o o t s g i v e a s t a r c h p o s i t i v e r e a c t i o n t o t h e I/KI t e s t . There i s a heavy s t a r c h s h e a t h i n the r o o t c o r t e x n e a r t h e attachment o f l a r g e h a u s t o r i a . - I l l -The Haustorium General C h a r a c t e r i s t i c s The haustoria of the p a r a s i t i c plant Geocaulon lividum are outgrowths of the root. They are l a t e r a l organs and form the p h y s i o l o g i c a l bridge between Geocaulon and i t s host's roots or rhizomes. Haustoria (Plate 15) are whitish i n color when young becoming tan or brown with age as a:cork layer forms. Whether the active l i f e of the haustorium i s to be measured i n months or years i s not known in Geocaulon. In Buckleya the haustoria are known to be a c t i v e f o r many years (Kusano-1902) but where annual plants are attacked the haust- .' orium can not be a c t i v e f o r more than one growing season. Haustoria i n Geocaulon are up to about 7 mm high and about 5 mm in diameter. They are roughly pear shaped ( F i g . 15) above the host and have clasping f o l d s around the host which may nearly e n c i r c l e i t . Occasionally a l i n e of haustoria a r i s e very close together on a root. They may then have a common h a u s t o r i a l cortex region with separate vascular cores and penetrating wedges. This phenomenon probably also explains the occasional haustorium with two separate vascular supplies from the mother root which merge very shortly to form one common vascular h a u s t o r i a l core. Development Haustoria do not a r i s e i n the same manner as l a t e r a l roots but simply as l a t e r a l extensions of the root as a whole rather than being new organs. The epidermis, cortex and endodermis of the root bulge but together to form an elongate protuberance on the root. These ti s s u e s are thus a l l continuous (Photograph 25) from root to haustorium. Within the endodermis of the mother root the vascular t i s s u e s give r i s e to a branch which supplies the upper portion of the haustorium. The endodermis of the root can be traced i n the - 112 -Plate 15 A, B and C, top, lateral and bottom views respectively of a Geocaulon haustorium on a Vaccinium rhizome, a, root of Geocaulon; b, Vaccinium rhizome. - 113 -2 m m - 114 -Figure 15 Sections through Geocaulon haustoria in two longitudinal planes and cross sections at several levels. H, sketch of an intact haustorium on a host as a reference. E, section longitudinal to the haustorium and transverse to the host. F, section at 90 degrees to E and longitudinal to the host. G, same section as E but showing a haustorial core.which has failed to penetrate the host as a wedge but instead clasped the host with two flanges. A, B, C and D are cross sections at the levels indicated by the dotted lines in E, F and G. The scale lines are 1 mm long. A is represented by Photograph 17, B is just above Photograph 18, Photographs 19 and 20 are at levels C and D respectively. Photographs 21 and 25 are sections E and F respectively down to level B. Photograph 22 is between levels A and half way between C and D. Photograph 24 is section G below level B and Photograph 23 and 26 are section F below level B. Photograph 28 is above Photograph 27 by about 200 microns and both are at a level slightly below B. - 115 -- 116 -ha u s t o r i u m through the "mother r o o t zone" t o the boundary w i t h the " t r a n s i t i o n a l y.ouo.". The. v a s c u l a r s u p p l y o f the lower p o r t i o n o f the h a u s t o r i u m does not de v e l o p u n t i l l a t e r , i . e . , a f t e r the h a u s t o r i u m has made c o n t a c t and p e n e t r a t e d a h o s t . The "mother r o o t zone" and " t r a n s i t i o n a l zone" a r e d e f i n e d below. I n t e r n a l S t r u c t u r e The h a u s t o r i u m can be d i v i d e d i n t o two main r e g i o n s ; an i n n e r c o r e ( K u i j t -1969) o r a x i s (Barber-1907) and an o u t e r p e r i p h e r a l o r c o r t i c a l r e g i o n ( B a r b e r -1907). The c o r t e x has an e p i d e r m a l l a y e r which may l a t e r become c o v e r e d by cork and a c o l l a p s e d zone which s e p a r a t e s the c o r t e x i n t o i n n e r and o u t e r p o r t i o n s . The i n n e r o r c e n t r a l c o r e i s composed o f the v a s c u l a r t i s s u e s and the wedge o f t i s s u e which p e n e t r a t e s the h o s t . T h i s c o r e may be d i v i d e d i n t o t h r e e zones a t r i g h t a n g l e s t o i t s l o n g i t u d i n a l a x i s o r p a r a l l e l t o the s u r f a c e of t h e hos t (Photograph 22). The uppermost zone, o r the one n e a r e s t t o t h e mother r o o t , i s t h e t r a n s i t i o n a l r e g i o n (Barber-1907) and w i l l be r e f e r r e d t o as t h e mother r o o t zone (Photograph 21 and 25). Next f o l l o w s t h e t r a n s i t i o n zone (Photograph 22) which c o r r e s p o n d s t o t h e i n t e r r u p t e d zone o f Barber-1907. The term ' i n t e r r u p t ' i m p l i e s a break i n t h e c o n t i n u i t y o r u n i f o r m i t y o f t h e v a s c u l a r system o f the c o r e . However t h e v a s c u l a r system o f the c o r e i s not c o n t i n u o u s and u n i f o r m but d i f f e r s on e i t h e r s i d e o f the t r a n s i t i o n a l zone w i t h r e s p e c t t o the n a t u r e o f t h e v e s s e l elements and t h e i r time o f development. Thus t h e term t r a n s i t i o n a l zone i s p r e f e r r e d t o t h e term i n t e r r u p t e d zone. From t h e t r a n s i t i o n a l zone down t o t h e base o f t h e h a u s t o r i u m w i t h i n t h e h o s t w i l l be c a l l e d the h a u s t o r i a ! zone w i t h the p a r t a c t u a l l y p e n e t r a t i n g t h e ho s t c a l l e d t h e wedge. These terms c o r r e s p o n d t o the " n u c l e u s " and " s u c k e r " (Barber-1907) which a r e f e l t t o be u n f o r t u n a t e and c o n f u s i n g terms which s h o u l d be dropped. The h a u s t o r i a l zone i s i l l u s t r a t e d i n Photograph 23. - 117 -Each region and zone w i l l be treated in greater detail below with constant reference to Fig. 15 as an aid in visualizing the structure being described. The Cortical Region The f i r s t clasping fold or cortical fold (Barber-1907) which envelops the host arises before differentiation of the haustorium into core and cortex and thus this fold can not be said to arise from any particular region of the haustorium. Clasping folds (Plate 15, C) are presumed to offer firm anchor-age for the haustorium (Barber-1907) so that the wedge of the core can force its way into the host without pushing the haustorium away from the host. In any case these folds certainly give structural support to the vascular portion of the haustorium. In mature haustoria which have successfully penetrated a host one sees layers of host bark, cortex and xylem apparently 'peeled back' by the prying folds of the haustorium. These folds penetrate the host and grow intrusively, tangential to the hosts surface. Due possibly to the action of the prying , folds, the growth reaction of the host or a combination of the two, host tissues are separated (Photograph 24) and the xylem exposed. In Santalum (Barber-1906, 1907) the normal appearance of the intrusive organ is as illustrated (Fig. 15, G) where the intrusive organ spreads out over the surface of the xylem. This is unusual in Geocaulon where the intrusive organ is normally wedge-shaped (Fig. 15, E) and drives right into or through the host xylem. In Santalum (Barber-1906, Plate i i i - 1 5 and Plate iv 18) what is called a compound haustorium arises when the intrusive organ f a i l s to penetrate the host on the f i r s t attempt. A new mantle arises between the f i r s t and the host and forms a new clasping fold. The origin of this second and any succeeding mantles in a compound haustorium is in the core tissues of the intrusive organ in contrast to the origin of the f i r s t clasping fold (Kuijt-- 118 -1969, 1970). No such compound haustorium was noted or examined anatomically in Geocaulon though such a structure or what appears externally to be so has been illustrated (Kuijt-1969, Fig. 7-1-b). In Geocaulon the intrusive organ is almost invariably wedge-shaped and penetrates directly into the host. There are a series of concentric collapsed zones about the core of a Geocaulon haustorium. These are diagrammatically illustrated in Fig. 15 by' heavy black lines. One collapsed zone, the outermost one, arises from the original clasping fold formation and the inner ones from points where the host tissues are separated by intrusive haustorial 'fingers'. Whether these fingers arise from the inner cortex region of the haustorium inside the collapsed zone or from the core tissues of the wedge is not known. If these intrusive organs do arise from the cores tissues as a result of attempted penetration there is no development of extensive secondary, tertiary, etc. mantles as in Santalum. I prefer to explain these collapsed zones as resulting from intrusive growths originating in the inner cortex which remains active even after the wedge has penetrated the host. This extended activity appears to be rare in Santalum (Barber-1907) but common in Geocaulon. It is also possible that the wedge, while passing through the host, produces lateral processes which grow tangentially to the host at various boundary layers. Commonly these growth separate bark from cortex and cortex from xylem. Whether the process of penetration i s entirely mechanical or aided by chemical or enzymatic processes is not known. It is entirely possible that growth responses' by the host may contribute to the confusion. Whatever their origin, these intrusive growths do occur and separate layers of the host. The collapsed layer, separation layer (Barber-1907) or separation strip - 119 -(Moss-1926) forms w i t h i n one o r two c e l l s o f the t i p o f t h e s e p r y i n g f o l d s (Moss-1926) and can be t r a c e d back a l o n g t h e c e n t r e o f the p r y i n g f o l d up i n t o t h e c o r t e x as f a r as the t r a n s i t i o n zone. C o n c e n t r i c s h e l l s of compressed o r c o l l a p s e d zones may be seen i n c r o s s s e c t i o n s o f h a u s t o r i a . The main c o l l a p s e d zone i s a s s o c i a t e d w i t h the f i r s t c l a s p i n g f o l d and t h e o t h e r s w i t h p r y i n g f o l d s . One can o f t e n f i n d a zone o f l a r g e empty spaces i n a s h e l l o u t s i d e a c o l l a p s e d zone i n young h a u s t o r i a but not i n mature ones. The s i g n i f i c a n c e o f t h e s e spaces i s not known. There i s a g r e a t d e a l o f u n c e r t a i n t y as t o cause and e f f e c t , o r i g i n and f u n c t i o n o f t h e c o l l a p s e d zones o f c l a s p i n g and p r y i n g f o l d s . The v e r y name of the t i s s u e s u g g e s t s t h a t i t r e s u l t s from e x t e r n a l l y a p p l i e d f o r c e s which c r u s h a zone o f c e l l s . T h i s was t h e concept o f e a r l y workers who named t h e t i s s u e . Growth o f the c o r e i n d i a m e t e r ( K u i j t - 1 9 6 9 , Barber-1907) a f t e r t h e o u t e r c o r t e x i s f i x e d i n d i a m e t e r has been s u g g e s t e d as one p o s s i b i l i t y . T h i s i d e a may account f o r th e c o l l a p s e d zones o f the p r y i n g f o l d s but not f O r t h a t o f the i n i t i a l c l a s p i n g f o l d . T h i s does not seem a l i k e l y e x p l a n a t i o n when one sees c o l l a p s e d zones r i g h t a t t h e t i p (Moss-1926) o f p r y i n g f o l d s . A p o s s i b i l i t y i n t h i s case i s t h a t ( o s m o t i c , Moss-1926) p r e s s u r e s e x e r t e d by s u p e r f i c i a l m e r i s t e m a t i c c e l l s , when c o n f i n e d by the h o s t t i s s u e s , cause c o l l a p s e o f the a x i a l c e l l s o f t h e p r y i n g f o l d . T h i s i s n o t an e n t i r e l y s a t i s f y i n g s o l u t i o n e i t h e r . I t has been s u g g e s t e d ( K u i j t - 1 9 6 9 , and o t h e r s ) t h a t i n s t e a d o f b e i n g p a s s i v e l y compressed t h e c o l l a p s e d zone a r i s e s t h r o u g h a c t i v e c o n t r a c t i o n o f t h e c e l l s i n t h i s zone and t h a t t h i s g i v e s added t h r u s t t o the a c t o f p e n e t r a t i o n . No c l e a r u n d e r s t a n d i n g o f th e c o l l a p s e d zones seems p o s s i b l e i n the l i g h t o f p r e s e n t knowledge. They remain e n i g m a t i c . - 120 -The Mother Root Zone The mother root zone (Photograph 21 and 25) is a direct continuation of parent root tissues (Barber-1907, Kuijt-1969) and forms the proximal part of the haustorium. This zone arises exogenously from the mother root and is also known as the transitional region (Barber-1907) by some authors. The term transitional has connotations which do not appear to be applicable to this zone and moreover the term is needed for the next zone where these connotations of change from one form to another are exactly what is desired. The mother root zone is the part of the haustorial core between the mother root and the transition zone. The mother root zone i s quite readily delimited by i t s sharp boundary with the transitional zone (Photograph 22) and f a i r l y sharp boundary with the surrounding cortex (Photograph 17). In some haustoria the vague outlines of the endodermis of the mother root can be seen as far down the mother root zone as the junction with the transitional zone. Cell size, amount of intercellular space and staining characteristics differentiate the mother root tissues from the cortex even when the endodermis can not be distinguished, though one hesitates to c a l l i t an endodermis, a line of darker staining c e l l walls (Photograph 25) can sometimes be seen around the mother root zone tissues. The center of the zone is composed of a solid core or a ring of vessel elements which are identical to those of the mother root with regard to size, shape and staining characteristics. These vessel elements are relatively the widest and longest in the haustorium and stain a brighter red in Safrahin-0 than the other vessel elements. The zone is circular in cross section at the mother root end and becomes oval shortly before i t grades into the transitional zone. The longitudinal axis of the oval is parallel to the - 121 -l o n g i t u d i n a l a x i s o f t h e h o s t . In t h e mother r o o t zone the v e s s e l members • may be i n a l o o s e l y a g g r e g a t e d p r o t o s t e l e o r i n a c l u s t e r o f bundles n e a r the mo Hi o r r o o t end o f the zone. The v e s s e l elements become more d i f f u s e as one p r o g r e s s e s down the zone u n t i l n e a r the t r a n s i t i o n zone boundary t h e v e s s e l elements a r e i s o l a t e d . The v e s s e l elements have s i m p l e p e r f o r a t i o n s on s l a n t e d end w a l l s and range from s c a l a r i f o r m l y t h i c k e n e d t o p i t t e d . The T r a n s i t i o n Zone T h i s i s t h e i n t e r r u p t e d zone o f Barber-1907 and was c o n s i d e r e d t o be absent i n Geocaulon by Moss-1926. I t i s v e r y c l e a r l y seen i n c l e a r e d whole h a u s t o r i a as d i s t i n c t gaps i n the v a s c u l a r c o n t i n u i t y o f the c o r e s . The t r a n s i t i o n zone i s sandwiched (Photograph 22) between the mother r o o t zone at t h e p r o x i m a l end o f t h e h a u s t o r i u m and the h a u s t o r i a l zone a t the d i s t a l end. The boundary w i t h the h a u s t o r i a l zone i s v e r y s harp but t h e boundary w i t h the mother r o o t zone more g r a d u a l . At the top o f the t r a n s i t i o n a l zone the mother r o o t v e s s e l elements g r a d u a l l y d i s a p p e a r , and a t the bottom o f the zone h a u s t o r i a l v e s s e l elements b e g i n t o appear. There may be an o v e r l a p of t h e two ty p e s i n some h a u s t o r i a but t h i s i s d i f f i c u l t t o d e t e r m i n e . In c l e a r e d whole h a u s t o r i a t h e r e appears t o be a complete s e p a r a t i o n but t h e r e s o l u t i o n i s n o t s u f f i c i e n t i n such a p r e p a r a t i o n t o determine whether o r not a few v e s s e l elements o f each t y p e o v e r l a p . The t r a n s i t i o n a l zone i s n o t n e c e s s a r i l y p e r p e n d i c u l a r t o t h e l o n g i t u d i n a l a x i s o f t h e h a u s t o r i u m and can not o f cou r s e be seen i n embedded m a t e r i a l . Thus when a h a u s t o r i u m i s p o s i t i o n e d i n t h e microtome f o r s e c t i o n i n g t h e r e i s no way o f knowing whether t h e c u t s w i l l be p a r a l l e l t o the b o u n d a r i e s o f the t r a n s i t i o n zone. A median s e c t i o n beyond a c e r t a i n maximum number o f degrees from t h e p l a n e o f t h e t r a n s i t i o n zone b o u n d a r i e s w i l l i n c l u d e mother r o o t zone v e s s e l members on - 122 -one s i d e and haustorial zone vessel members on the other.. Such a condition i s illustrated in Photograph. 28. The few large scattered vessel members in cross section at the top of the photograph are mother root zone vessel members and the group of small vessel members in radial f i l e s at the bottom of the photograph are from the haustorial zone. The vessel elements, i f they can be called such, in the transition zone diff e r from those found in the other two zones of the core. The cells are the shortest of the three vessel element types found, are very narrow and most seem to be occluded (Photographs 18, 22 and 28) with a substance staining very dark red with Safranin-O. There are very few of these xylem cells compared to the.numbers in the other zones of the core. The transition zone vessel elements are embedded in a matrix of small dense cells which stain darker than the parenchyma of the mother root zone or haustorial zone. Since this transitional zone appears in a l l the cleared haustoria examined regardless of age i t does not appear to be a function of the age of the haustorium. Vascular continuity is certainly broken (Photograph 22) in the transitional zone and this raises some interesting questions regarding the functioning of the haustorium. With regard to Santalum and other genera (Barber-1907) the point has been raised that this gap makes i t d i f f i c u l t to understand how the passage of fluids can occur from host to parasite. On the face of i t i t does seem quite unreasonable for a plant to have vascular continuity from the tip of the aerial shoot through the short-shoot, rhizome, root and top part of the haustorium, then to have a gap of several hundred microns, followed by renewed vessel element production in the lower part of the haustorium. The extent of vessel element production in the haustorial zone is greater than that in any other part of the plant and approached only by vessel element - 123 -formation in some roots near large haustoria (Photographs 12 and 27). Materials from the host passing through the haustorial zone vessel elements, i f they do so, must leave the xylem and pass through the small dense parenchymatous cells of the transition zone in order to reach the vessel elements of the mother root zone. Is this perhaps a deliberate f i l t e r i n g system? The problem of the transition zone is not as isolated as i t seems. At the bottom of the haustorial zone the number of haustorial zone vessel elements decreases markedly before the host xylem is reached and contact between host vessel elements or other conducting cells and the vessel elements of the.haustorium- are rare and perhaps accidental. Material from the host must pass through the small dense parenchy-matous cells of the haustorial t i p in order to reach' the vessel elements of the haustorial zone. There is certainly no free-flowing "pipeline" from host to.parasite along which large volumes of materials can be.passed quickly. This leads one to wonder i f the essential substance, i f there is such for Geocaulon, might be microgram amounts of a growth hormone or some such substance needed in very small amounts, which Geocaulon and other green parasites can not make for themselves, at least in sufficient quantities. Against this'notion is the observation that Geocaulon wilts very quickly when uprooted, more so than other plants growing nearby. The sparsity of roots, the fact that i t is restricted to moist habitats and the tremendous vessel element production in the lower part of haustoria, and in roots next to large haustoria, are also contrary evidence. It appears that Geocaulon depends, to some extent i ;upon its hosts for water. No physiological work has been reported for this species. The Haustorial Zone The haustorial zone of the core is the zone most dist a l from the mother root. It follows immediately below the transition zone. The haustorial zone - 124 -in the context used here includes the wedge of tissue which penetrates the host. Barber-1907 calls the upper part of the haustorial zone the "nucleus" and the lower part or wedge, the "sucker". The vessel elements are arranged in radial f i l e s (Photograph 27) and arise by secondary growth from a vascular cambium near the edge of the core. Very few cells are produced external to the cambium. The f i l e s may be peripheral around the circular or oval cross section of the haustorial core or in two broad nearly parallel plates of vessel elements aligned parallel to the longitudinal axis of the host. Such an arrangement and derivation also occurs in Santalum (Barber-1907). The vessel elements of the haustorial zone are narrower and shorter than those of the mother root zone (Photograph 22) and stain more purple than red with Safranin-O. This and their radial arrangement makes them readily distinguishable from mother root vessel elements. In a cross section in the region of maximum haustorial zone vessel element production, several hundred microns below the transition zone, there are many more haustorial zone vessel elements than mother root zone vessel elements in the part of the mother root zone where the maximum number of vessel elements is found. Vessel elements do not seem to arise in the haustorial zone u n t i l after host penetration occurs. As the level of the host is approached the number of vessel elements decreases markedly and those remaining become peripherally arranged and scattered (Photograph 19) as in Santalum (Barber-1907). At this level and below, right down to the surface of the host the nonvascular cells of the core are arranged in f i l e s parallel to the longitudinal axis of the haustorium. These cells are long and narrow, have dense cytoplasm and stain darker than cortical parenchyme cel l s . At the surface of the host xylem these central cells are small, rounded and have dense cytoplasm. The vessel elements remain peripheral on the wedge - 125 -and s c a t t e r e d (Photograph 23) u n t i l the t i p o f the wedge i s r e a c h e d where t h e r e i s t o o l i t t l e wedge m a t e r i a l l e f t t o make t h e term p e r i p h e r a l m e a n i n g f u l . Throughout the whole h a u s t o r i u m o n l y the n o n - v a s c u l a r c e l l s o f t h e cor e r e g i o n a r e p o s i t i v e f o r s t a r c h i n the I/KI t e s t . The p e n e t r a t i n g organ i s almost always wedge shaped i n Geocaulon and may p e n e t r a t e r i g h t t h r o u g h the hos t xylem and i n t o the c o r t e x on t h e f a r s i d e . In r a r e c a ses p e n e t r a t i o n i s not a c h i e v e d and t h e wedge becomes i n s t e a d a p a i r o f c l a s p i n g f o l d s on t h e s u r f a c e o f t h e h o s t xylem (Photograph 24; F i g . 15, G) as i s common i n Santalum (B a r b e r - 1 9 0 7 ) . V e s s e l element d i s t r i b u t i o n i n t h e h a u s t o r i u m P r e v i o u s s e c t i o n s have d e a l t w i t h each zone o f the h a u s t o r i a l c o r e and d e s c r i b e d the typ e o f v e s s e l element found i n t h a t zone. There has been mention o f d i f f e r e n c e s i n t h e numbers o f v e s s e l elements i n c r o s s s e c t i o n s o f each zone as w e l l as t h e d i f f e r e n c e s i n s t r u c t u r e . I f s e r i a l c r o s s - s e c t i o n s o f h a u s t o r i a are made and v e s s e l element counts made a t i n t e r v a l s , a v e r y i n t e r e s t i n g d i s t r i b u t i o n o f v e s s e l elements i n the h a u s t o r i u m i s o b s e r v e d . T h i s i n f o r m a t i o n i s p r e s e n t e d i n Graph 1. The number o f v e s s e l e l e m e n t s , i n the c r o s s s e c t i o n . of the h a u s t o r i a l zone which had t h e most v e s s e l e l e m e n t s , i s equated t o 100% and the counts a t o t h e r l e v e l s i n t h e h a u s t o r i u m c o n v e r t e d t o t h e a p p r o p r i a t e p e r c e n t a g e . In the. f o u r h a u s t o r i a p r e s e n t e d i n the graph t h e v a l u e o f 100% ranged from 230 i n A t o 585 i n B. The r e l a t i v e p r o p o r t i o n s o f v e s s e l elements at a g i v e n l e v e l i s q u i t e c o n s t a n t i n a l l t h e h a u s t o r i a examined. The number o f mother r o o t zone v e s s e l elements d e c r e a s e s q u i c k l y as t h e t r a n s i t i o n zone i s r e a c h e d . Towards the end of t h e t r a n s i t i o n zone t h e number o f h a u s t o r i a l zone v e s s e l elements r i s e s v e r y r a p i d l y t o t h e maximum, and t h e n d e c r e a s e s a g a i n . The i n t e r s e c t i o n o f o r d i n a t e and a b s c i s s a r e p r e s e n t s v e r y n e a r l y t h e - 1 2 6 -t i p o f t h e wedge and t h e t o p o f the . o r d i n a t e c o r r e s p o n d s t o t h e m o t h e r r o o t •/.oni.' .••.ivM'U.y a f t e r t h e s e p a r a t i o n f r o m t h e m o t h e r r o o t . One can r e a d i l y see on t h e g r a p h s t h e e f f e c t o f s e c t i o n i n g t h e h a u s t o r i u m a t an a n g l e t o t h e p l a n e o f t h e t r a n s i t i o n a l z o n e . A and D show no o v e r l a p o f m o t h e r r o o t zone and h a u s t o r i a l zone v e s s e l e l e m e n t s and were p r e s u m a b l y c u t a t a s m a l l a n g l e t o t h e p l a n e o f t h e t r a n s i t i o n a l z o n e . I n B and C t h e c u r v e s f o r t h e m o t h e r r o o t zone and h a u s t o r i a l zone v e s s e l e l e m e n t s i n t e r s e c t s h o w i n g t h a t t h e s e c t i o n s were c u t a t an a p p r e c i a b l e a n g l e t o t h e p l a n e o f t h e t r a n s i t i o n a l zone.- ' - 127 -Graph 1 D i s t r i b u t i o n o f v e s s e l elements i n f o u r Geocaulon h a u s t o r i a . The a b s c i s s a i s i n 5% u n i t s and the o r d i n a t e i n 100 micron u n i t s . 100% i s t h e number o f v e s s e l elements found i n the c r o s s s e c t i o n o f the h a u s t o r i a l zone w i t h the most v e s s e l elements. The h a u s t o r i a were s e r i a l l y s e c t i o n e d i n 10 m i c r o n s e c t i o n s , m, mother r o o t zone v e s s e l e l ements; t , t r a n s i t i o n zone v e s s e l e l ements; h, h a u s t o r i a l zone v e s s e l e l ements; b , l e v e l o f the h o s t s b a r k ; and x, t h e l e v e l o f t h e h o s t s xylem. - 12 8 -GRAPH 1 - 129 -The Cell Wall Thickenings In some rhizomes and short-shoots of Geocaulon a type of c e l l wall thickening is found which does not seem to have been reported in the anatomical literature (Cutler-1970). The most similar structure that could be found is described in Esau-1943. In her material, pear roots, the "wall thickenings" are found only on cortical cells immediately outside the endodermis. The wall thickenings in Geocaulon are definitely not in the form of a continuous sheath such as an exodermis (Russow-1875) nor as an inner cortical sheath (Guttenberg-1940) but are found throughout the cortex, in the outer phloem and in the tissue between the pith and the bundles. Esau-1943 writes of the "wall thickenings" as mostly radial with some adjacent to intercellular spaces, and her illustrations indicate that they are not always paired.. In Geocaulon the wall thickenings are always paired, are never next to intercellular spaces and though most may be on radial walls there are many on tangential and transverse walls. The Geocaulon wall thickenings are isolated units centered on the common wall between two ce l l s . They are usually in the form of a hemispherical or lens shaped bulge into each c e l l . The pear root "wall thick-enings" (Esau-1943) are bands of material in the longitudinal axis of the root. In Geocaulon transverse sections the occasional c e l l w i l l show a wall in face view. If the wall has a thickening on i t there w i l l be a circular portion in the center which stands out from the rest of the wall by it s lack of pit fields. This contrast is more marked i f the stain methyl violet is used. There is a great deal of variation in the appearance of the c e l l wall thickenings (Photographs 29 to 3 4 ) . Bands of darker staining material appear in some (Photograph 3 3 ) , some have internal holes (Photographs 3 0 , 33 and 3 4 ) , and some have darkly stained central bands with swollen tips (Photographs 31 - 130 -and 32). P a r t o f t h i s d i f f e r e n c e i n appearance may be due t o s e c t i o n s through d i f f e r e n t p a r t s o f t h e w a l l t h i c k e n i n g but p a r t may be due t o an a c t u a l d i f f e r e n c e i n s t r u c t u r e . These v a r i o u s views may r e p r e s e n t a d e v e l o p -ment s e r i e s . O t h e r shapes have been ob s e r v e d . In some cases t h e r e a r e two w a l l t h i c k e n i n g s s i d e by s i d e w i t h a common l e n s shaped o u t e r p o r t i o n e n c l o s i n g them b o t h . A m i c r o c h e m i c a l i n v e s t i g a t i o n i n v o l v i n g s t a i n i n g and s o l u b i l i t y was undertaken i n an attempt t o i d e n t i f y the s t r u c t u r a l component(s) o f t h e w a l l t h i c k e n i n g s . The p o t e n t i a l components under c o n s i d e r a t i o n were: s u b e r i n , c u t i n , c h i t i n , l i g n i n , c a l l o s e , ' p e c t i n s ' , ' h e m i c e l l u l o s e s ' and c e l l u l o s e . The s e c t i o n s c o n t a i n i n g the w a l l t h i c k e n i n g s were mounted s i n g l y on m i c r o s c o p e s l i d e s w i t h S a s s ' s a d h e s i v e which h e l d the s e c t i o n s on t h e s l i d e s i n a l l but s t r o n g and/or hot a l k a l i e s . In a l k a l i n e s o l u t i o n s t h e s l i d e s were p l a c e d i n p e t r i d i s h e s and the s o l v e n t added and removed, e x t r e m e l y c a r e f u l l y , by p i p e t t e . S e c t i o n s which s t i l l came o f f c o u l d u s u a l l y be r e t r e i v e d by f l o a t i n g them c a r e f u l l y o n t o a s l i d e . The f i b r e g l a s s sandwich t e c h n i q u e (Wilson-1961) was t r i e d and found h e l p f u l . i n some i n s t a n c e s but w i t h t h e t h i n s e c t i o n s b e i n g used p a r t u s u a l l y came o f f w i t h the f i b r e g l a s s . Extreme c a r e and p a t i e n c e was found t o be the b e s t p r o c e d u r e . The f o l l o w i n g a r e some o f t h e s t a i n i n g and s o l u b i l i t y c h a r a c t e r i s t i c s o f s e l e c t e d c e l l w a l l components. C h i t i n - r e d i n S a f r a n i n - O . (Johansen-1940). - i n s o l u b l e i n 50% chromic a c i d . (Johansen-1940) L i g n i n - r e d i n S a f r a n i n - O . (Johansen-1940) - r e d w i t h p h l o r o g l u c i n o l . (Johansen-1940, Shaw-1929, Jensen-1962) - r e d i n a l k a l i n e methyl r e d . (Johansen-1940). - 131 -- soluble in Shultze's reagent (1:1, cone. HNO^sat. KCl). (Shaw-1929, Johansen-1940). - soluble in 50% chromic acid. (Shaw-1929, Johansen-1940). - insoluble in KOH, NaOH. (Shaw-1929). Cut in and Suberin ,. - red with Safranin-O. (Johansen-1940). - red with Sudan IV. (Johansen-1940, Shaw-1929). - soluble in 3% KOH, 4% NaOH. (Shaw-1929) - insoluble in 50% chromic acid. (Shaw-1929, Johansen-1940) Callose - blue with resorcein blue and aniline blue; red with rosolic acid (Corallin). (Shaw-1929, Johansen-1940, Currier-19 57). - soluble in 1% KOH, 1% NaOH, cone. CaCl 2. (Johansen-1940, Currier-1957). 'Pectins' - red with ruthenium red; violet with methylene blue. (Johansen-1940, Shaw-1929). - red with ruthenium red. (Davenport-1960, Jensen-1952), - soluble in 0.5% ammonium oxalate. (Shaw-1929, Jensen-1962). - soluble in the series:hot water, hot dilute acid, dilute alkalie. (Shaw-1929, Johansen-1940) 'Hemicelluloses' - red with ruthenium red. (Davenport-1960) - green with methylene blue. (Johansen-1940). - soluble in Shultze's reagent. (Mitchell-1930). - soluble in 4% NaOH. (Jensen-1962). - soluble in NaOH from 4% to 17.5% (Davenport-1960). - 132 -Cellulose - red with congo red, blue with methylene blue. (Shaw-1929). - red with congo red, tan with bismark brown. (Johansen-1940). - soluble in 50% chromic acid. (Johansen-19'rO, Shaw-1929). - insoluble in Shultze's reagent and concentrated KOH. (Shaw-1929). ' - insoluble in 17.5% NaOH. (Jensen-1962, Davenport-1960). The material used was k i l l e d and fixed in FAA (5% formalin, 5% gl a c i a l acetic acid, 27% water, and 63% ethanol), dehydrated in the tertiary butanol series, embedded in Paraplast, mounted with Sass 1s adhesive, and run through a series of xylene and ethanol down to d i s t i l l e d water. One would expect most fats and water soluble materials to be removed by this time as well as ethanol soluble substances. In the stain tests carried out, the wall thickenings and the cortex c e l l walls reacted the same while other parts of the rhizome reacted differently. In the interests of brevity and since i t is the wall thickenings that are of concern, only the reactions of the c e l l wall thickenings and the cortex cells walls w i l l be reported. The following stains did not stain the wall thickenings: resorcein blue, aniline blue, methyl red, Safranin-O, Sudan IV, phloroglucinol, rosolic acid. The wall thickenings were stained red by ruthenium red and congo red, blue by methylene blue, violet by crystal violet and orange/brown by bismark brown. These preliminary stain test results indicate that the wall thickenings do not have appreciable amounts of cutin, suberin, chitin, lignin or callose. The pectins and hemicellulose are positive by the ruthenium red test but negative by the methylene blue test. The cellulose tests are positive. A solvent series was. developed and sections passed through in order. - 133 -After each step a slide was removed and examined microscopically to see whether the wall thickenings were s t i l l present or had dissolved in part or entirely. The series was: Step 1 D i s t i l l e d water; 90 degrees C; 20 hours. Step 2 3% HC1; 60 degrees C; 30 minutes; rinse in d i s t i l l e d water. Step 3 2.5% NH^ OH; 60 degrees C; 30 minutes; rinse in d i s t i l l e d water. Step "4 0.5% ammonium oxalate; 60 degrees C; 2 hours; rinse in d i s t i l l e d water. Step 5 1% KOH; 40 degrees C; 30 minutes; rinse in d i s t i l l e d water. Step 6 Saturated CaCl^; 60 degress C; 4 hours; rinse in d i s t i l l e d water. Step 7 4% NaOH; 25 degrees C; 18 hours; rinse very carefully in d i s t i l l e d water. step 8 Shultze's reagent; 25 degrees C; 1 hour; rinse in d i s t i l l e d water. Step 9 18% NaOH; 25 degrees C; 3 hours; rinse in d i s t i l l e d water. Other procedures were tried on material which had not gone through the above series of solvents. These are: A. 10% NaOH; 25 degrees C; 3 hours; rinse in d i s t i l l e d water. B. Shultze's reagent; 25 degrees C; 20 hours; rinse in d i s t i l l e d water. C. 50% chromic acid; 25 degrees C; 30 minutes; rinse in d i s t i l l e d water. D. Slightly modified Jensen-1960 procedure. D-l ammonium oxalate at 0.5%; 30 degrees C for 8 hours followed by 90 degrees C for 15 hours; rinse in d i s t i l l e d water. D-2 NaOH at 4%; 30 degrees C; 16 hours; rinse in d i s t i l l e d water. Results The wall thickenings remained after Step 8 and after procedures A, B and D-2. The wall thickenings were gone but an outline of the c e l l walls remained - 134 -after procedure C. -An a n a l y s i s by w a l l components f o l l o w s . A n e g a t i v e r e s u l t i n d i c a t e s o n l y t h a t the m a t e r i a l i s p r e s e n t i n t o o low a c o n c e n t r a t i o n t o be the major s t r u c t u r a l . c o m p o n e n t o f t h e w a l l t h i c k e n i n g s . C h i t i n - The w a l l t h i c k e n i n g s d i s s o l v e d i n p r o c e d u r e C i n which c h i t i n s h o u l d be' i n s o l u b l e . A l l the s t a i n t e s t s were n e g a t i v e f o r c h i t i n . L i g n i n - L i g n i n s h o u l d d i s s o l v e i n Step 8 but t h e w a l l t h i c k e n i n g s were not a f f e c t e d . . The s t a i n t e s t s f o r l i g n i n were n e g a t i v e . C u t i n and S u b e r i n - These s h o u l d be s o l u b l e i n Step 7, p r o c e d u r e A and pro c e d u r e D-2. These s o l v e n t s d i d n ot d i s s o l v e the w a l l t h i c k e n i n g s . The w a l l t h i c k e n i n g s d i s s o l v e d i n p r o c e d u r e C whereas c u t i n and s u b e r i n s h o u l d n o t . The s t a i n t e s t s were n e g a t i v e f o r a l l f a t s . C a l l o s e C a l l o s e i s s o l u b l e i n Steps 5 and 6 which d i d n o t a f f e c t the w a l l t h i c k e n i n g s . The s t a i n t e s t s f o r c a l l o s e were a l l n e g a t i v e . P e c t i n s These a r e s o l u b l e i n Steps 1 thr o u g h 4 and i n p r o c e d u r e D - l . These s o l v e n t s do not a f f e c t the w a l l t h i c k e n i n g s . The p o s i t i v e r e a c t i o n t o ruthenium r e d i s not s p e c i f i c f o r p e c t i n s . H e m i c e l l u l o s e s These s h o u l d be s o l u b l e i n Steps 7 and 8 and i n p r o c e d u r e s A, B and D-2. The w a l l t h i c k e n i n g s d i d n ot d i s s o l v e i n t h e s e s o l v e n t s . Step 9 w i l l d i s s o l v e h e m i c e l l u l o s e s as w e l l as b e t a - and gamma-c e l l u l o s e but n o t a l p h a - c e l l u l o s e . The w a l l t h i c k e n i n g s d i d d i s s o l v e i n Step 9. The s t a i n t e s t s a r e not s u f f i c i e n t l y s p e c i f i c t o d i s t i n g u i s h t h e s e s u b s t a n c e s . There i s no sharp d i v i d i n g l i n e between low m o l e c u l a r weight c e l l u l o s e s and h i g h m o l e c u l a r weight h e m i c e l l u -l o s e s . The l a t t e r can not be r u l e d out as major components o f t h e w a l l t h i c k e n i n g s . - 135 -C e l l u l o s e A l p h a - c e l l u l o s e s h o u l d be i n s o l u b l e i n Step. 9 but b e t a - and gamma-forms may d i s s o l v e (Davenport-1960). The s t a i n t e s t s are p o s i t i v e f o r c e l l u l o s e but may not be s p e c i f i c f o r a l p h a - c e l l u l o s e C o n c l u s i o n On t h e b a s i s o f t h e p r o c e d u r e s c a r r i e d out h e r e one c o u l d say t h a t t h e w a l l t h i c k e n i n g s were h i g h l y p o l y m e r i z e d h o l o c e l l u l o s e (Davenport-1960) but not d i f f e r e n t i a b l e i n t o c e l l u l o s e o r h e m i c e l l u l o s e . The w a l l t h i c k e n i n g s do no t appear t o be a l p h a - c e l l u l o s e which i s the most h i g h l y p o l y m e r i z e d and t h e main s t r u c t u r a l component o f p l a n t s . - 136 -Photographs 1, 2 and 3 These p h o t o g r a p h s ' l a b e l l e d A, B and C a r e taken from s e c t i o n s A, B and C r e s p e c t i v e l y o f P l a t e 5. A, i s i n t h e b a s a l p o r t i o n o f a s h o r t - s h o o t i n i t s f i r s t y e a r o f growth. B, i s i n the t r a n s i t i o n a l r e g i o n where s h o r t - s h o o t and a e r i a l shoot meet. C, i s i n the base o f t h e a e r i a l s h o o t . The v a s c u l a r system changes from a more o r l e s s c o n t i n u o u s c y l i n d e r i n t h e s h o r t -shoot base t o d i s c r e t e b u ndles i n t h e a e r i a l s h o o t . F i b r e caps i n c r e a s e i n development from t h e base o f t h e s h o r t - s h o o t t o t h e a e r i a l s h o o t . The cork l a y e r g i v e s way i n t h e t r a n s i t i o n zone t o e p i d e r m i s . The s c a l e l i n e s a r e 0.2 5 mm l o n g . Photograph 4 V a s c u l a r bundle i n t h e rhizome o f Geocaulon. These bundles a r e about 1:1, xylem t o phloem, A few t h i n -w a l l e d f i b r e s cap the phloem end o f the b u n d l e . S e v e r a l rows o f l a r g e v e s s e l elements a r e embedded i n the remainder o f t h e xylem. The s c a l e l i n e i s 0.10 mm l o n g . - 137 -- 138 -Photographs 5, 6, 7, 8, 9 and 10 The labels D, E, F, G, H and I correspond to sections D, E, F, G, H and I r e s p e c t i v e l y as i l l u s t r a t e d i n Fig.. 7. D, i s i n the rhizome at the base of the'short-shoot. E, F and G are i n progressively younger portions of the short-shoot. H and I are at the base and shortly above, the base, r e s p e c t i v e l y , of the a e r i a l shoot. The vascular bundles are mostly separate in the rhizome, j o i n into a complete vascular c y l i n d e r in the short-shoot and then separate again into d i s c r e t e bundles i n the a e r i a l shoot. Fibre cap development increases from the rhizome to the a e r i a l shoot. C e l l wall thickenings are evident i n D, E and F. H and I have only an epidermis, the others a l a y e r of cork. The scale l i n e s are 0.25 mm long. - 139 -- 140 -Photograph 11 Median l o n g i t u d i n a l s e c t i o n through a rhizome apex. No p e r i c l i n a l d i v i s i o n s a r e o b s e r v e d i n the 1^ and few i n the T^. There t h e n f o l l o w , two l a y e r s o f p r e d o m i n a n t l y a n t i c l i n a l d i v i s i o n s which might be c o n s i d e r e d t h e o u t e r p o r t i o n of a p a r t i a l l y s t r a t i f i e d corpus.. The topmost c e l l o f t h e c e n t r a l l i n e a r column i s d i s c e r n a b l e i n the f o u r t h o r f i f t h l a y e r . The s c a l e l i n e i s 0.20 mm l o n g . Photograph 12 C r o s s s e c t i o n o f a r o o t c l o s e t o t h e attachment o f a l a r g e mature h a u s t o r i u m . The ends o f r o o t s and r o o t s w i t h no h a u s t o r i a , do not have such e x t e n s i v e v e s s e l element development o r c o r k f o r m a t i o n . The s c a l e l i n e i s 0.25 mm l o n g . - 141 -- 142 -Photograph 13 Fruit wall of an immature Geocaulon f r u i t . Ex, exocarp; M, mesocarp and E, endocarp. Contrast this with Photographs 14 and 16 of the mature f r u i t wall. The scale line is 0.25 mm long. Photographs 14 and 16 Mature f r u i t wall of Geocaulon. Ex, exocarp; M, mesocarp; E, endocarp and EN, endosperm. Photograph 14 is the outer portion of the pericarp, the exocarp or epicarp. Note in Photograph 16 the extension of the exocarp by c e l l expansion and the decrease in the endocarp by crushing and absorption. The mesocarp becomes very hard and stony and has vascular channels on i t s surface as well as within the tissue. The scale line is 0.20 mm long. Photograph 15 Endosperm and embedded embryo of a mature Geocaulon f r u i t . The scale line is 0.25 mm long. - 1 4 3 -- 144 -, Photographs 17, 18, 19 and 20 C r o s s s e c t i o n s o f Geocaulon h a u s t o r i a a t t h e l e v e l s A, j u s t below B, C and D r e s p e c t i v e l y as i l l u s t r a t e d i n F i g . 15. These l e v e l s a r e t h e mother r o o t zone, the t r a n s i t i o n a l zone, t h e h a u s t o r i a l zone a t t h e l e v e l o f t h e h o s t c o r t e x and t h e h a u s t o r i a l zone a t t h e t i p o f t h e wedge. S e v e r a l c o l l a p s e d zones can be seen i n Photograph 19. The s c a l e l i n e s f o r 17, 18 and 20 a r e 0.25 mm l o n g ; t h a t f o r 19 i s 0.50 mm l o n g . - 1 4 5 -- 146 -Photographs 21, 22 and 23 Longitudinal sections through a Geocaulon,haustorium. Photograph 21, the mother root zone, corresponds to Fig. 15-E, from the top down to level B. Photographs 22 and 23 correspond to Fig. 15-F. Photograph 22 is centered on the transition zone, T, with the mother root zone, M, above and the haustorial zone, H, below. Photograph 23 i s between levels C and D and shows the peripheral vessel elements in the haustorial wedge. The scale lines for Photographs 21 and 23 are 0.40 mm long and that for 22 is 0.2 5 mm long. Photograph 24 Longitudinal section through a Geocaulon haustorium corresponding to Fig. 15-G from level B down. Rare in Geocaulon, Fig. 15-E is most common. The intrusive wedge usually penetrates the host xylem and does not spread out on the surface as has occurred here. The scale line is 0.40 mm long. - 147 -- 148 -Photographs 25 and 26 Longitudinal sections of different portions of the same young Geocaulon haustorium. Photograph 25 shows the continuity of tissues,from mother root to haustorium and a line of darkly stained c e l l walls in the position of the endodermis. Photograph 26 shows a view corres-ponding to Fig. 15-F. In this immature state there is no differentiation of.cortex and core regions. The scale lines are 0.25 mm long. Photographs 27 and 28 Cross sections of Geocaulon haustoria in the upper haustorial zone and in the transitional zone respectively. There is only a few hundred microns between these sections. The scale lines are 0.25 mm long. - 149 -- 150 -Photographs 29-34 R e p r e s e n t a t i v e s e c t i o n s o f c e l l w a l l t h i c k e n i n g s found In some rhizomes and s h o r t - s h o o t s o f Geocaulon. These a r e a l l rhizome c r o s s s e c t i o n s . The l e n g t h s o f the r e f e r e n c e l i n e s a r e : 29-0.06 mm; 30, 31 and 32-0.04 mm; 33-0.02 mm and 34-0.015 mm. - 151 - 152 -Bibliography Anderson, J.P. 1946.. 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New Phytologist 25(4):264-276. ; - T - — :; . Payne, W.W. 1969. A quick method f o r c l e a r i n g leaves. Ward's Bulletin.: May. Vol. 8., No. 61. Ward's Natural Science Establishment, Inc. P.O. Box 1712, Rochester, N.Y. 14603. Perry, L.M. 1931. The vascular f l o r a of St-Paul Island, Nova Scotia. Rhodora 33:105-126. . P i e h l , M.A. 1963. The n a t u r a l h i s t o r y and taxonomy of the genus Comandra (Santalaceae). Ph.D. Thesis. University of Michigan. D i s s e r t a t i o n Abstracts. 23(9):3094. P o r s i l d , A.E. 1939. Contributions to the f l o r a of Alaska. Rhodora 41:141-183; 199-254; 262-301. - 155 -P o r s i l d , A.E. 1950. Vascular plants of Nueltin Lake, Northwest T e r r i t o r i e s . National Museum of Canada. B u l l e t i n 118:72-83. P o r s i l d , A.E. 1951. Botany of southeastern Yukon adjacent to the Canol Road. National Museum of Canada. B u l l e t i n 121. Potter, D. 1932. Plants c o l l e c t e d i n the southern region of James Bay. Rhodora 36:274-284. Powell, J.M. 1968. Personal Communication.' Ram, M. 1959a. Morphological and embryological studies i n the family Santalaceae. 2. Exocarpus, with a discussion on i t s systematic position';, Phytomorphology 19(1):4-19. Ram, M. 1959b. Morphological and embryological studies i n the family Santalaceae. 3. Leptomeria R.Br. Phytomorphology 19(1) :20-33. Raup, H.M. 1930. The d i s t r i b u t i o n and a f f i n i t i e s of the vegetation of the Athabasca-Great Slave Lake region. Rhodora 32:187-208. Raup, H.M. 1935. Botanical investigations i n Wood Buffalo Park. National Museum of Canada. B u l l e t i n 74. B i o l o g i c a l Series 20. Raup, H.M. 1947. The Botany of southwest Mackensie. Sargentia 6:1-275. Rodin, R.J. and R.E. Davis. 1967. The use of papain i n c l e a r i n g plant tissues f o r whole mounts. Stain Technology 42(4):203-206. Rouleau, E. 1944. 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C e l l w a l l development o f c a m b i a l d e r i v a t i v e s i n A b i e s c o n c o l o r . Ph.D. T h e s i s . U n i v e r s i t y o f C a l i f o r n i a a t B e r k e l e y . - 158 -Appendix'1 - Techniques Collection and Preservation of Material Fresh material was k i l l e d and fixed immediately in FAA. The FAA used consisted of the following: 5% - glacial acetic acid. 5% - formaldehyde (40%). 90% - ethanol (70%). The material was l e f t in the fixative for days or weeks, rinsed in 70% ethanol and then stored in 70% ethanol u n t i l needed. Herbarium material was soaked in hot water and detergent u n t i l soft, rinsed thoroughly in d i s t i l l e d water and stored in 70% ethanol. Slide Preparation Stored material was dehydrated in the tertiary butanol series and embedded in 56-57°C Paraplast. Sections were cut on a rotary microtome, mounted with Sass's adhesive on glass slides and l e f t to dry overnight. The slides were soaked in xylene to remove the Paraplast, rehydrated in an ethanol series and soaked in Stockwell's solution for several days to remove tannins and pigments. A tr i p l e staining procedure was used: tannic acid/ferric chloride, Safrannin-0 and fast green. Canada balsam diluted with xylene was used as a mounting medium. ' Photomicrographs and Anatomical Drawings Photomicrographs were made with a Leitz Ortholux Microscope, Aristophot Stand Assembly and a 4 x 5 bellows camera using Tri-X Pan professional sheet film A l l developing and printing was done personally. Several procedures were used for the drawings. Some were done free-hand, some were drawn from the - 159 -negatives using an e'rilarger to project the image onto the paper at the desired magnification and some were projected directly onto the paper with a micro-dia, apparatus and reflecting mirror. Whole Mount Preparation A great deal of d i f f i c u l t y was encountered in clearing and staining whole mounts of Geocaulon material. Portions could be cleared satisfactorily by the method of Arnott-1959, but washing and dehydration preparatory to staining led to the formation of a whitish opacity in rhizomes, f r u i t s , stems and haustoria in particular. Several other clearing methods were tried to find out i f they would remove the material responsible for opacity in ethanol but none was satisfactory. These methods were: Payne-1969, Rodin and Davis-1967, Simpson-no date, Stebbins-1938, Bates-1931, Varrelman-1938. It was f i n a l l y decided to take material as far as the 250 gm Chloral Hydrate/100 ml step of Arnott-1959 where i t was quite transparent and leave the material in this solution in an unstained condition for observation. Apart from the lack of differential staining the major obstacle to observation of material in this condition is breathing the Chloral Hydrate vapours. Starch Distribution The starch staining solution used was 0.3 grams Iodine and 1.5 grams Potassium Iodide dissolved in ^00 m i l l i l i t r e s of d i s t i l l e d water. Thin free-hand sections were cut with a razor blade, placed in a depression slide and covered with a few drops of the I/KI solution. After a few minutes the starch grains showed up plainly under the dissecting microscope as dark blue spots. These preparations while quick and easy and very useful were not permanent and - 160 -faded o v e r n i g h t . Permanent p r e p a r a t i o n s were made by f o l l o w i n g the p r e v i o u s l y o u t l i n e d s e c t i o n i n g and mounting p r o c e d u r e but s u b s t i t u t i n g I/KI, aqueous . b l u i s h s a f r a n i n , and f a s t green f o r the s t a i n s . T h i s s t a i n does n o t g i v e t h e s t a r c h as i n t e n s e a c o l o r ( p a r t l y due t o t h i n n e r s e c t i o n s ) , but i t i s permanent and g i v e s b e t t e r r e s o l u t i o n . Appendix 2 - Geocaulon D i s t r i b u t i o n The l o c a t i o n s are g i v e n as f i r s t l a t i t u d e and then l o n g i t u d e i n degrees and minutes. The l o c a t i o n s a r e o r d e r e d by l a t i t u d e b e g i n n i n g w i t h t h e most s o u t h e r l y . Most l o c a t i o n s a r e p l u s o r minus 3 minutes o r l e s s . 414 17 71 19 46 03 64 05 47 29 75 26 48 10 69 44 44 25 81 22 .46 03 71 21 47 32 71 15 . 48 11 64 54 44 33 72 47 46 15 60 09 47 35 52 55 48 13 90- 10 44 55 70 30 46 21 68' 06 47 36 91 15 . 48 14 90 11 44 58 66 42 46 24 60 38 47 40 65 38 48 18 88 50 44 58 81 23 46 33 72 00 47 41 69 37 48 20 85 50 44 59 64 57 46 34 87- 22 47 41 69 48 48 20 88 50 46 43 60 23 47 48 54 48 48 20 116 50 45 11 81 35 46 47 60 50 47 48 61 26 48 22 68 49 45 16 66 03 46 51 71 04 47 50 68 55 48 28 119 15 45 18 81 38 46 53 71 38 47 57 84 54 48 29 71 39 45 20 78 27 46 55 84 37 47 58 66 11 48 30 64 10 45 23 67 20 46 57 84 44 47 58 89 42 48 32 54 43 45 26 64 54 47 59 80 41 48 40 87 00 45 30 80 32 47 00 84 45 48 42 • 79 45 45 47 70 14 47 12 60 09 48 00 88 55 48 43 79 31 45 49 82 56 47 18 87 56 48 01 65 19 48 43 87 58 45 50 82 20 47 20 84 38 48 01 89 35 48 45 86 15 45 50 84 35 47 21 84 37 48 05 65 34 48 45 86 25 45 54 68 54 47 25 66 04 48 05 71 32 48 45 87 15 45 55 74 00 47 26 70 30 48 05 89 26 • 48 45 117 25 45 55 85 55 47 26 84 44 48 06 69 45 48 46 55 38 45 56 81 46 47 27 70 32 48 07 69 11 48 46 86 32 47 28 87 54 • 48 07 75 22 48 46 116 54 1 6 1 -48 4 7 66 0 5 5 0 0 0 8 8 3 5 48 47 66. 5 3 5 0 0 6 1 1 9 0 4 4 8 47 . 8 7 ' 1 0 5 0 0 9 1 2 0 5 2 4 8 4 8 ' 8 6 3 7 5 0 ' 1 1 6 6 1 2 4 8 4 8 8 6 5 8 5 0 1 3 6 1 4 6 4 8 4 9 8 7 0 5 5 0 1 3 6 3 4 0 4 8 5 0 6 6 0 0 5 0 1 3 6 3 4 9 4 8 5 0 9 4 2 5 5 0 .14 6 3 3 7 4 8 5 1 6 4 2 4 5 0 1 4 6 6 2 4 4 8 5 1 8 7 0 2 5 0 1 7 . 64 . . 0 1 4 8 5 2 7 2 0 5 5 0 1 9 1 1 5 5 2 4 8 5 2 9 4 2 3 5 0 2 0 1 1 5 5 0 4 8 5 3 6 5 57 5 0 2 2 1 1 5 3 6 4 8 5 3 6 6 3 8 .-.'... 5 0 2 3 1 1 4 2 9 4 8 5 3 6 6 4 0 5 0 2 9 1 2 0 2 9 4 8 5 3 8 9 5 5 5 0 3 0 7 3 4 1 4 8 5 5 6 6 1 1 5 0 3 2 6 8 4 7 4 8 5 6 5 6 0 2 5 0 3 3 5 9 2 0 4 8 5 7 9 0 1 5 5 0 3 4 1 1 6 2 3 4 8 5 9 6 5 5 7 5 0 3 8 5 7 1 5 5 0 3 8 1 1 7 3 3 4 9 0 0 5 4 3 0 5 0 4 3 5 7 2 2 4 9 0 2 8 8 0 3 5 0 4 8 1 2 2 5 6 4 9 0 3 8 6 0 2 5 0 4 9 5 7 1 4 4 9 0 4 5 5 5 5 "50 5 0 5 8 5 5 4 9 0 4 1 2 0 4 7 5 0 5 1 1 2 0 2 9 4 9 0 5 1 2 0 4 2 5 0 5 4 1 1 5 0 9 4 9 0 7 1 2 0 3 9 5 0 5 5 1 1 4 0 8 4 9 0 4 6 5 0 2 5 0 5 6 1 1 5 3 5 4 9 1 0 5 7 5 9 5 0 5 7 1 1 5 0 8 4 9 1 1 7 7 5 7 5 0 5 9 1 1 4 5 8 4 9 1 1 1 2 0 3 7 5 0 5 9 1 1 5 0 5 4 9 1 3 5 8 2 0 5 0 5 9 1 1 8 1 4 4 9 1 8 5 6 2 4 4 9 1 8 5 7 0 0 5 1 0 1 9 5 2 8 4 9 2 0 5 6 3 4 5 1 0 1 1 1 5 0 8 4 9 2 1 7 6 3 6 5 1 0 2 1 1 5 0 2 4 9 2 5 6 8 0 5 5 1 0 2 1 1 5 0 5 4 9 2 5 1 1 5 2 7 5 1 0 2 1 2 1 4 3 4 9 2 8 5 7 4 2 5 1 0 3 1 1 6 4 8 4 9 2 9 1 2 0 3 5 5 1 0 3 1 1 8 0 5 4 9 3 2 8 8 0 4 5 1 0 4 5 6 5 5 4 9 3 8 9 5 3 6 51 0 4 1 1 4 4 8 4 9 3 8 1 1 4 3 6 5 1 0 4 1 1 5 0 8 4 9 3.9 9 6 1 5 5 1 0 5 1 1 5 2 1 4 9 4 1 1 1 5 5 9 5 1 0 5 1 1 8 0 1 4 9 4 6 8 5 2.7 5 1 0 6 1 2 1 3 5 4 9 4 7 8 6 2 7 5 1 0 7 7 3 1 9 4 9 4 8 6 4 2 0 5 1 0 8 1 1 5 3 4 4 9 4 8 8 6 3 1 5 1 0 9 1 1 8 1 1 4 9 5 2 5 5 4 0 • .. 5 1 1 1 8 0 3 1 4 9 5 2 6 4 3 1 5 1 1 2 1 1 6 1 9 4 9 5 2 7 7 1 5 5 1 1 5 8 0 4 5 4 9 5 3 6 4 2 9 5 1 1 6 8 0 3 6 4 9 5 3 1 1 5 5 3 5 1 1 6 1 1 6 5 5 4 9 5 5 9 3 0 0 5 1 1 8 5 6 4 4 5 1 1 8 9 8 2 3 . 5 3 1 7 1 1 7 5 3 5 1 1 9 1 2 1 4 7 5 3 1 7 1 2 2 3 0 5 1 2 0 9 8 1 2 5 3 1 8 6 0 2 5 5 1 .23 1 1 6 3 0 5 3 2 0 6 0 3 0 5 1 2 5 5 5 3 7 5 3 2 0 9 4 3 0 5 1 2 5 1 1 6 3 0 5 3 2 0 1 0 4 4 6 5 1 2 7 1 1 6 3 2 5 3 2 2 6 0 2 4 5 1 2 8 5 6 2 0 5 3 2 7 5 5 4 7 5 1 2 8 1 1 7 0 4 5 3 3 0 1 0 6 0 6 5 1 2 9 7 8 4 6 5 3 3 0 1 1 3 4 1 5 1 2 9 1 1 7 1 1 5 3 3 0 1 2 2 4 2 5 1 3 0 1 2 0 3 7 5 3 3 4 6 0 1 4 5 1 3 6 5 5 4 4 5 3 3 5 6 4 2 0 5 1 3 7 1 1 7 2 4 5 3 3 6 1 0 5 5 4 5 1 4 1 1 1 7 4 3 5 3 3 6 1 0 8 3 8 5 1 5 0 6 2 5 3 5 3 4 2 5 6 5 8 5 1 5 7 1 1 8 0 2 5 3 4 3 1 0 5 • 1 6 5 1 5 8 1 1 8 1 6 5 3 4 3 1 1 3 1 3 5 3 4 5 1 0 5 1 5 5 2 0 0 7 9 3 0 . 5 3 4 9 5 8 2 5 5 2 0 5 1 1 3 2 8 5 3 5 0 8 9 3 0 5 2 0 7 1 1 9 4 8 5 3 5 3 1 2 4 3 8 5 2 1 2 1 0 4 4 6 5 3 . 5 5 1 0 5 1 8 5 2 1 4 1 2 2 0 6 5 3 5 5 1 0 6 0 5 5 2 1 5 1 1 3 4 9 5 3 5 5 1 0 6 3 7 5 2 1 6 5 5 3 0 5 3 5 5 1 2 2 4 8 5 2 1 6 5 5 4 1 5 3 5 7 1 0 1 1 5 5 2 1 6 1 1 3 4 7 5 3 5 7 1 0 5 4 6 5 2 1 7 5 5 3 6 5 3 5 7 1 0 6 1 5 5 2 1 7 8 1 3 1 5 3 5 7 1 0 6 2 9 5 2 2 0 7 0 3 5 5 3 5 8 1 1 3 1 3 5 2 2 3 1 1 4 5 6 5 3 5 8 1 1 3 5 9 5 2 2 7 1 0 2 4 0 5 3 5 9 9 7 5 0 5 2 3 3 6 6 0 7 5 3 5 9 1 2 4 5 8 5 2 3 7 1 0 3 5 6 5 2 4 3 1 0 3 3 9 5 4 0 0 6 4 0 0 5 2 5 0 1 0 6 3 0 5 4 0 1 1 2 8 3 3 5 2 5 2 1 0 3 0 0 5 4 0 6 1 2 2 0 3 5 2 5 2 1 1 8 3 3 5 4 0 7 1 2 5 1 2 5 2 5 3 1 1 8 0 3 5 4 0 8 1 0 8 2 6 5 2 5 5 1 1 8 0 3 5 4 0 9 5 8 3 0 5 2 5 8 1 2 2 4 2 5 4 1 0 1 2 5 2 0 5 2 5 9 1 1 8 0 6 5 4 1 7 1 2 2 3 9 5 2 5 9 1 2 2 2 9 5 4 2 0 9 8 5 3 5 4 2 4 1 1 1 2 4 5 3 0 0 5 6 0 0 5 4 2 4 1 1 6 4 7 5 3 0 0 1 0 1 2 8 5 4 2 5 1 2 5 4 1 5 3 0 2 1 1 8 0 7 5 4 2 7 1 2 4 2 3 5 3 0 3 9 3 0 9 5 4 3 1 9 7 3 8 5 3 0 3 9 3 1 5 5 4 3 2 1 2 2 4 3 5 3 0 4 1 0 0 0 2 5 4 3 5 1 0 0 1 7 5 3 0 6 1 0 4 2 8 5 4 3 8 1 0 3 1 3 5 3 0 8 8 3 1 8 5 4 4 0 . 6 6 4 5 5 3 1 5 1 1 6 5 6 5 4 4 0 8 7 5 5 5 3 1 6 5 9 5 5 5 4 4 2 6 6 5 1 5 3 1 6 6 0 3 0 5 4 4 3 . 1 0 7 0 2 - 162 54 46 66 .49 54 48 66 49 54 50 79 25 54 57 118 57 54 57 122 50 54 58 122 58 55 00 117 21 55 02 59 02 55 02 67 09 55 02 129 36 55 03 94 40 55 03 129 35 55 04 118 48 55 05 59 11 55 07 114 02 55 13 119 16 55 13 119 26 55 17 77 47 55 17 112 28 55 20 104 58 55 20 105 05 55 21 77 30 55 23 119 36 55 25 118 05 55 25 122 35 55 27 107 53 55 29 125 58 55 30 122 34 55 30 127 46 55 35 93 40 55 39 131 41 55 40 124 22 55 43 120 26 55 45 60 20 55 45 126 17 55 49 100 23 55 55 95 50 55 57 87 22 55 57 126 37 55 59 87 38 56 00 87 38 56 02 104 00 56 02 121 54 56 02 129 09 56 04 123 39 56 05 76 30 56 07 116 50 56 07 121 46 56 10 76 25 56 15 64 45 56 31 94 06 56 38 125 25 56 46 126 25 56 ' 47 124 51 56 48 89 03 56 50 122 23 56 56 113 27 57 00 92 18 57 05 122 35 57 14 • 111 23 57 22 106 50 57 31 106 21 57 36 117 31 57 37 101 44 57 43 106 17 57 45 94 05 57 49 101 28 57 57 131 07 57 58 111 40 58 21 116 00 58 21 116 08 58 21 134 20 58 22 111 35 58 24 116 08 58 28 111 30 58 36 94 10 58 38 117 29 58 39 124 38 58 40 111 20 58 41 94 14 58 42 111 08 58 42 123 40 58 .46 94 10 58 47 125 37 58 49 66 10 58 50 110 50 58 51 113 57 58 51 114 09 58 57 96 47 58 57 113 55 58 58 125 45 58 59 125 47 59 01 109 00 59 14 135 27 59 16 105 50 59 17 129 16 59 17 129 48 59 19 129 51 59 20 107 30 59 20 117 10 59 21 106 00 59 22 97 46 59 24 126 02 .59 24 133 35 59 26 126 06 59 28 108 28 59 28 117 11 59 28 132 35 59 28 133 34 59 28 135 16 59 30 107 41 59 31 111 28 59 34 112 15 59 36 • 109 13 59 36 113 07 59 38 126 50 59 39 126 57 59 43 111 31 59 44 105 08 59 48 99 39 59 50 104 10 59 52 111 37 59 55 102 05 59 55 131 41 59 56 111 43 59 56 127 27 59 56 128 30 59 56 133 48 60 00 112 47 60 00 115 51 60 00 127 43 60 01 111 53 60 02 129 02 60 05 123 47 60 06 106 23 60 06 112 16 60 06 134 51 60 07 128 48 60 07 130 44 60 12 120 16 60 14 104 14 60 14 132 55 60 15 123 28 60 17 109 02 60 19 112 37 60 19 141 00 60 24 145 18 60 25 149 22 60 28 133 11 60 29 133 10 60 29 150 05 60 30 116 13 60 30 137 00 60 30 149 20 60 30 149 45 60 33 134 25 60 35 116 07 60 37 99 55 60 37 133 04 60 38 117 39 60 39 151 30 60 40 111 52 60 40 115 43 60 40 116 00 60 40 . 135 12 60 42 123 37 60 43 112 50 60 43 135 03 60 45 115 37 60' 46 137 23 60 47 107 38 60 47 136 28 60 50 117 25 60 50 137 2.0 60 51 115 42 60 51 133 00 60 .52 110 34 60 56 121 37 60 56 149 39 60 57 117 20 60 57 133 00 60 58 117 13 60 58 117 16 61 00 105 00 61 00 149 42 61 03 117 22 61 03 138 31 61 06 94 04 61 10 111 53 61 11 135 12 61 13 149 49 61 15 113 20 61 15 117 36 61 20 138 45 61 20 149 43 61 22 128 15 61 24 135 31 61 30 141 00 61 30 149 .20 61 31 108 06 61 35 133 05 61 36 115 38 61 40 114 00 61 43 133 05 61 45 114 38 61 50 109 10 61 50 121 21 61 52 121 22 61 55 113 15 61 57 127 13 61 59 113 32 62 00 140 34 163 -6 2 05 127 3 5 63 02 156 02 64 00 121 30 65 15 147 47 6 2 1 3 ' 123 2 0 63 07 126 30 64 00 125 05 65 17 126 52 6 2 16 1 2 3 37 63 08 117 16 . 64 00 145 45 65 20 143 47 62 18 117 39 63 12 145 30 64 02 145 45 65 23 127 27 62 19 136 23 63 13 109 10 64 04 139 25 65 25 126 15 62 20 106 25 63 13 114 15 64 12 145 50 65 30 •' 12 8 40 62 24 110 45 63 16 123 37 64 17 115 12 65 30 144 40 62 27 114 22 63 20 143 04 64 19 146 20 65 36 148 39 62 22 150 02 63 23 136 42 64 23 146 55 65 43 118 47 62 28 123 37 . 63 30 141 00 64 35 139 30 62 30 114 18 63 30 143 24 64 44 112 12 66 05 117 58 62 32 108 18 63. 31 128 40 64 45 116 00 66 05 118 02 62 34 111 31 63 33 115 55 64 45 147 30 66 08 117 40 62 35 109 38 63 35 110 02 64 50 147 25 66 •32 117 45 62 40 114 15 63 35 135 54 64 51 147 41 66 40 122 00 62 40 121 06 63 38 143 58 64 53 147 51 66 57 151 47 62 41 137 10 63 40 144 56 64 54 125 34 62 42 12 3 08 .63 41 136 41 64 56 147 45 67 02 119 50 62 43 109 10 63 42 144 23 67 25 140 59 62 45 116 05 .'• 63 42 148 49 65 00 150 47 67 30 150 00 62 51 121 23 63 46 137 17 65 01 127 20 62 59 155 27 63 48 13.5 48 65 03 127 17 68 14 133 28 63 51 136 17 65 04 124 37 68 20 133 30 63 02 110 57 63 56 117 15 65 11 12 3 28 68 39 133 42 In a d d i t i o n t o the above a r e a s which a r e p l o t t e d on the maps t h e f o l l o w i n g s i t e s are r e c o r d e d on h e r b a r i u m l a b e l s as l o c a t i o n s f o r Geocaulon. Due t o nebulous d e s c r i p t i o n s o r ambiguous names o f p l a c e s t h e s e s i t e s can n o t be a c c u r a t e l y p l o t t e d . Saskatchewan - Meadow Lake F o r e s t Reserve. O n t a r i o - A n v i l Lake; Bruce P e n i n s u l a . Quebec - Loup R i v e r (46° 30' N, 73° W). Newfoundland - s o u t h s h o r e , N o t r e Dame Bay; Bay o f I s l a n d s (49° N, 5 8° W).. A l a s k a - C h i t i n a R i v e r Head; Headwaters o f Sheep and Mascot C r e e k s . Northwest T e r r i t o r i e s - Gr e a t S l a v e Lake (61° N, 115° W); S l a v e Lake F o r e s t Reserve; McKenzie Mountains (63° N, 127° W); Nahanni Mountains, McKenzie Region; Yukon R i v e r between Tanana and Rampart; - 164 -Yukon F l a t s (65° t o 65° 30' N, 141° t o 142° W). A l b e r t a - Whitemud Creek, n e a r Edmonton (53° 30' N, 113° 30' W); C a r i b o o Mountains (59° N, 116° W); n e a r Edmonton (53° 30* N, 113° 30' W); n e a r J a s p e r (53° N, 118° W); Bow R i v e r n e a r B a n f f , 4500', (51° N, 115° 30' W); r o a d t o Cave S p r i n g s , B a n f f , (51° N, 115° 30' W); J a s p e r N a t i o n a l P a r k ; Athabasca V a l l e y ; Saskatoon Mountains, n e a r B e a v e r l o d g e . B r i t i s h Columbia - between T o p l e y and Burns Lake, 2300', (54° 30' N, 126° W); C l e a r w a t e r F o r e s t Reserve (52° N, 120° W); L a i r d Range, 2400', (58° N, 127° W); K i t s u m g a l l u m River.(54° N, 128° W); Headwaters o f t h e F r a s e r R i v e r (53° N, 120° W); K i c k i n g Horse R i v e r , 4000', (51° 30' N, 116° 30« W); Salmon R i v e r (49° N, 117° W); T e l e g r a p h T r a i l (54° N, 129° W); T r o u t Lake; F i s h Lake; F r a s e r Lake; ( t h e r e are a t l e a s t 15 T r o u t L a kes, 8 F i s h Lakes and 4 F r a s e r Lakes i n B r i t i s h C o l u m b i a ) ; Peace R i v e r a r e a ; the t r a i l t o Qualcho Lake i n Tweedsmuir Park (53° N, 125° W); south of T e r r a c e A i r f i e l d (54° N, 128° W); Kimpton Creek T r a i l , Kootenay N a t i o n a l P a r k , 4410'; H o r s e t h i e f Creek (50° 30' N, 116° 05' t o 20' W); L a i r d R i v e r V a l l e y , McKenzie Lowlands (59° t o 60° N, about 123° W); Other - Green Mountain, 7000'; h i g h p o i n t between the McKenzie and L a i r d R i v e r s ; Wood B u f f a l o P a rk; Upper Matanuska V a l l e y . I have p e r s o n a l l y examined and c o l l e c t e d Geocaulon i n t h e f i e l d a t t h e f o l l o w i n g s i t e s . B a n f f , Western A l b e r t a 115° 34' N, 51° 10' W In t h e B a n f f t o w n s i t e , a c r o s s the r o a d from t h e B a n f f S p r i n g s H o t e l t e n n i s c o u r t s . Open woods. The p l a n t s have up t o 6 - 165 -inflorescences but almost invariably only one ripe f r u i t on each inflorescence. Kananaskis Road, Southwestern Alberta 115° 05' N, 51° 02' W Over the dam and south along the west side of Barrier Lake, on the banks of a creek upstream from an.old wooden bridge. This is a damp mossy site at the edge of the forest. 115° 02' N, 51° 02' W A short distance in along a dirt road going southeast from the Kananaskis road, a short distance north of the Research Stations. Several small patches on the edge of a small bog and on the slopes behind and to the west of the bog. Across the road to the east, north of the point where the power line crosses the road, there are several small patches. Fairly dense mixed woods. 115° OS' N, 50° 59' W About a mile north of the intersection of Wasootch Creek and the Kananaskis road, another wide, f l a t , gravelly, creek passes under the road in culverts. On the open gravelly northwest bank of this creek there are three small patches of Geocaulon among Arctostaphylos and Vaccinium. The plants are short, s t i f f , reddish specimens typical of exposed sites. Across the road along the base of and on the slope of the h i l l above the creek, there is an extensive patch. About one half mile east a small creek runs north under the road. Along the banks of this creek in deep moss and dense shady woods there are several patches of Geocaulon. Across the road from this creek site there is an open Pine woods with several patches of Geocaulon. 115° 08' N, 50° 57' W At the base of the south-facing slope of a h i l l , on the west side of the road just before a sharp turn west, there are - 166 -s e v e r a l v e r y s m a l l p a t c h e s which are v e r y d i f f i c u l t t o f i n d as they are h i d d e n by o v e r h a n g i n g bush. Mixed woods and heavy under-brush . 115° 09' N, 50° 5i+' W An o l d e r p o r t i o n o f t h e r o a d branches t o t h e west and runs down t o the c r e e k . Behind a clump o f Elaeagnus commutata at r o a d s i d e and r i g h t down t o the creek bank t h e r e i s a p a t c h o f Geocaulon i n mixed woods. F r u i t i n g was heavy i n t h i s p a t c h i n 1968. These Kananaskis p l a n t s have a number o f i n f l o r e s c e n c e p e d u n c l e s but r a r e l y more than one f r u i t p e r p e d u n c l e and seldom more than one f r u i t p e r p l a n t . F r u i t i n g i s g e n e r a l l y q u i t e poor w i t h t h e e x c e p t i o n o f t h e l a s t s i t e mentioned. Manning Pa r k , Southern B r i t i s h Columbia 120° 37' N, 49° 29' W Behind t h e Manning Park Lodge, between the Amphitheatre and t h e r i v e r , t h e r e i s a men's outhouse. Geocaulon can be found i n c l e a r e d woods i n t h i s a r e a . 120° 35' N, 49° 29' W E a s t of t h e e a s t gate t o Manning Park i s t h e Towers M o t e l . A; d i r t r o a d w i t h a gate r u n s n o r t h and t h e n west b e h i n d t h e m o t e l and ends e v e n t u a l l y n e a r a swamp. A t r a i l l e a d s t o an o l d wooden b r i d g e a c r o s s a creek west o f th e swamp. 'Geocaulon grows on the west bank o f the creek n o r t h o f t h e b r i d g e , on t h e t o p and s l o p e s o f t h i s bank, west o f t h i s c r e e k a c r o s s t h e t r a i l and around a s m a l l swamp and n o r t h from t h e creek bank up t h e h i l l s i d e i n s e v e r a l p a t c h e s . The Manning Park p l a n t s a l l have from 1 t o 6 i n f l o r e s c e n c e s p e r p l a n t b u t almost i n v a r i a b l y o n l y 1 f r u i t p e r i n f l o r e s c e n c e . There a r e u s u a l l y no more than 1 o r 2 f r u i t s p e r p l a n t and t h e s e f r u i t s a r e q u i t e s m a l l . - 167 -W e s t - c e n t r a l A l b e r t a 117° 46' N, 53° 17' W- E a s t o f t h e J a s p e r Park gate on t h e highway t o Edmonton a r o a d l e a d s n o r t h t o Kinky Lake and W i l d Horse Lake which are a few m i l e s n o r t h o f t h e highway. South o f the Kinky Lake-shore campsite t h e r e i s a Spruce-Equisetum bog w i t h Geocaulon growing on i t s e a s t e r n s l o p e s . A few cases o f 2 f r u i t s p e r i n f l o r e s c e n c e were found here but f r u i t s were s c a r c e . 115° 17' N, 51° 58' W At m i l e 256.2 n o r t h o f Coleman on t h e F o r e s t r y Trunk Road t h e r e i s a s m a l l Spruce bog on.the west s i d e o f t h e r o a d and, j u s t s o u t h o f a summit on the r o a d . Geocaulon grows around t h e edges o f t h e bog. N o r t h of Lake S u p e r i o r , western O n t a r i o 86° 37' N, 48° 48' W N o r t h e a s t o f t h e b r i d g e , over t h e L i t t l e P i c R i v e r , on highway #17 an abandoned s e c t i o n o f highway runs n o r t h . On both s i d e s o f t h i s o l d r o a d and o n l y a few hundred f e e t i n from the highway t h e r e a r e p a t c h e s o f Geocaulon. On t h e e a s t s i d e i t i s growing on an exposed sandy bench among A r c t o s t a p h y l o s and V a c c i n i u m as w e l l as a l o n e on t h e sandy s l o p e s below. No f r u i t s were f o u n d h e r e . On t h e west s i d e o f t h e r o a d i t grows i n f a i r l y dense mixed secondary growth but i s q u i t e r a r e . Only 1 f r u i t c o u l d be found here b u t i t was 1 1/2 cm i n d i a m e t e r , t h e l a r g e s t f o u n d . 87° 02' N, 48° 51' W On t h e n o r t h s i d e o f #17 highway, 6.7 m i l e s e a s t o f t h e T e r r a c e Bay Motor H o t e l chimney t h e r e i s a l a r g e s i g n a d v e r t i s i n g Wiebens R e s o r t , 37 m i l e s t o t h e west. Geocaulon grows beneath and around t h i s s i g n a l o n g w i t h Cornus c a n a d e n s i s under an open Spruce - 168 -f o r e s t . There i s only one i n f l o r e s c e n c e b e a r i n g one f r u i t p e r p l a n t . These are the t y p i c a l p a l e green f l a c c i d p l a n t s o f shaded a r e a s . 87° 05' N, 48° 49' W A l a r g e rock o u t c r o p s p a r s e l y d o t t e d w i t h t r e e s and d e e p l y c o v e r e d i n moss i s on the s o u t h s i d e o f t h e #17 highway 2.3 m i l e s e a s t o f t h e T e r r a c e Bay Motor H o t e l chimney. The Geocauloh rhizomes are found between the moss and the r o c k o r s o i l s u r f a c e . The p l a n t s were the t y p i c a l r e d d i s h c o l o r and s h o r t f i r m h a b i t o f exposed specimens. Only 2 p l a n t s were found w i t h more than 1 f r u i t p e r p l a n t and 1 o f t h e s e had 2 f r u i t s p e r i n f l o r e s c e n c e . 88° 03' N, 49° 02' W On the n o r t h s i d e o f #17 highway and .10.3 m i l e s e a s t o f the j u n c t i o n w i t h highway #11 Geocaulon was found i n deep moss under a t h i c k Spruce s t a n d . No f r u i t s were found. 90° 15' N, 48° 57' W No f r u i t s were found i n t h i s s i t e which i s a wet Spruce bog 12.9 m i l e s e a s t of t h e Up.sala P r o v i n c i a l P o l i c e S t a t i o n . Geocaulon grows on mossy hummocks here i n r e l a t i v e l y d r y c o n d i t i o n s . S o u t h e a s t e r n Manitoba 95° 36' N, 49° 38' W The s i t e i s on t h e s o u t h s i d e o f #1 highway, 14.7.miles e a s t o f t h e j u n c t i o n w i t h highway #11 and j u s t e a s t o f t h e p i c n i c grounds. Geocaulon grows on mossy hummocks a t the base o f t h e Spruce t r e e s . Cornus c a n a d e n s i s and s e v e r a l s p e c i e s o f Equisetum were common h e r e . There were many p l a n t s w i t h 2 i n f l o r e s c e n c e s and 1 w i t h 3 p e r p l a n t . There was o n l y one f r u i t p e r i n f l o r e s c e n c e and f r u i t s i z e v a r i e d w i d e l y . 95° 15' N, 49° 39' W On t h e s o u t h s i d e of #1 highway and 8.7 m i l e s e a s t of t h e j u n c t i o n w i t h highway #302 t h e r e i s a dry Spruce woods. Geocaulon i s r a r e here and no f r u i t s were found. 

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