MONTANA STATE '0- ^-:- - . >■' ■4 ' This «cover" page added by the Internet Archive for formatting purposes S I .„.,„!(f,?.^l',Tftf^(!!',ST,*IinilrmmMl?lTl 587.33 Nlldnit ISS4 3 68(54 0014 0421 2 STATE DOCUMENTS COLLECTIOM. m 2 f) 1998 MONTANA STATE UBRAR* 1515 E. 6th AVE. HELENA, MONTANA 59620 DEMOGRAPHIC MONITORING OF THREE SPECIES OF BOTRYCHIUM (OPHIOGLOSSACEAE) IN WATERTON LAKES PARK, ALBERTA 1993 Progress Report Peter Lesica and Kathy Ahlenslager Herbarivun Division of Biological Science University of Montana Missoula, Montana 59812 February 19 94 © 1994 Montana Natural Heritage Program This report is to be cited as follows: Lesica, P. and K. Ahlenslager. 1994. Demographic monitoring of three species of Botrychium (Ophioglossaceae) in Waterton Lakes Park, Alberta. Unpublished report to U.S. Fish and Wildlife Service. Montana Natural Heritage Program, Helena. 19 pp. INTRODUCTION Moonworts fBotrvchium subgenus Botrvchium) are a group of small, nonvascular plants in the Family Ophioglossaceae with a center of distribution in western North America (Wagner et al. 1984) . Plants consist of a single frond divided into a sterile, leaf-like segment below a fertile, spore-bearing segment. Many of the species are difficult to find and are considered rare (Lellinger 1985) . Among these is B^ paradoxum, a candidate for listing as a threatened or endangered species by the U.S. Fish and Wildlife Service. This plant is currently known from a handful of sites in Alberta, Montana and Utah (Wagner and Wagner 1981, Lellinger 1985, W.H. Wagner pers. com.). This species is unusual in that the sterile segment of the frond is replaced by a second fertile segment. In Montana and Alberta B^ paradoxum has often been found growing with other species of moonworts, and the question of whether B^ paradoxum is a good species or simply an unusual form of other more common species has been raised. In Waterton Park, Alberta, B^ paradoxum occurs with B^ hesperium and their putative hybrid B^ X watertonense. Analysis of flavonoid chemistry between B_^ paradoxum and B^ hesperium was performed and nearly half (9) of the 20 compounds examined were unique to one of the two species (Wagner et al. 1984) . These results support the contention that B,^. paradoxum is a valid species but are not entirely convincing because the distribution of secondary metabolites such as flavonoids in tissues may be partially under environmental or developmental rather than strictly genetic controls. More powerful molecular techniques such as isozyme electrophoresis and DNA analysis have not yet been performed because of the difficulty in obtaining material (W. Hauk pers. com.). Direct observation of many individuals over a period of years will help to determine whether B_^ paradoxum is a stable genetic species or merely an aberrant phenotype of another species. Due to their small size and rarity, relatively little is known about the life history and ecology of moonworts. Lellinger (1985) states that moderate disturbance such as grazing seems to favor their establishment and continued survival. It is not known how long-lived the plants are or whether or not they produce a frond each year. Wagner (pers. com.) suggests that some populations may be ephemeral while others may persist for centuries. Observation of individual plants through time will help to answer these questions. The purpose of our study is to track morphological characters of individual moonwort plants over a period of many years in order to: (1) determine basic life history and population parameters such as average life expectancy and mean age for B^ paradoxum and B^ hesperium and (2) gather evidence to support or reject the hypothesis that B^ paradoxum is not a true species but an aberrant phenotype of B^ hesperium and/or other species of Botrychium. This report summarizes results from the first five years of the study. METHODS Study Site We conducted our study in the Red Rock Canyon area of Waterton Lakes National Park ca. 12 km northwest of the town of Waterton. The study site is located on the north side of Bauerman Creek ca. 50 m northwest of the footbridge and ca. 2 00 m southwest of the Red Rock parking and picnic area. Elevation of the site is ca. 1555 m. The compass bearing from the site to the uppermost outhouse along the creek is 139 degrees. The compass bearing from the site to the top of Anderson Peak is 270 degrees. The vegetation of the area is lodgepole pine (Pinus contorta) and spruce (Picea engelmannii) forest interspersed with meadows dominated by rough fescue (Festuca scabrella) , Idaho fescue (|\. idahoensis) and shrubby cinquefoil (Potentilla fruticosa) on hilltops and warm exposures. Our study site is in one of these meadows. Field Procedures We established two parallel belt transects at the study site. The upper (north) consists of eight adjacent 1-m^ plots, and the lower (south) consists of five plots. The start and end points of the transects were permanently marked by driving 1/2 inch diameter reinforcing bar into the ground. These marker pins were painted orange to aid relocation. Individual Botrychium plants were mapped following methods outlined in Lesica (1987) . For each plant we recorded the following morphological characters: Species (P = paradoxum. W = watertonense. H = hesperium) Degree of pinnation of sporophore (PI = once pinnate, P2 = twice pinnate etc.) Above ground height (H12 = a plant 12 mm tall from ground level to tip of longest sporophore branch) Length of sporophore (LIO = a plant with a fertile segment 10 mm long) Thus a plant scored as H-P2-H20-L14 is morphologically identified as B^ hesperium 20 mm tall with a twice-pinnate sporophore 14 mm long. Species were identified using the criteria presented in Wagner et al. (1984). Beginning in 1990 we also recorded the length and degree of pinnation of the sterile frond segment. In 1991 plants were too dense to map in portions of plots 3 and 4 in the upper line. These areas have been eliminated from the analysis. Data Analysis We found that some plants would go undetected for one or more years but reappear in subsequent years. The presence of underground plants can be inferred by comparing transect maps from the full sequence of years. We recorded these plants as "U." The proportion of underground plants ranged from 3-20% in 1991-92 among the three species. In 1990-91 among the three species, 93% of the plants stayed underground for only one year. Thus, we have chosen to eliminate the first and last years of the study from demographic analysis. Montgomery (1990) reported that plants of Botrvchium dissectum would often be "missing" for one or two years before appearing again. Mortality is the proportion of the plants present in year t that died between years t and t+1. Recruitment is the proportion of plants present in year t that were new recruits. Statistical significance of mortality and recruitment proportions among species were compared with an overall chi-square goodness of fit test. If a 2 X 3 test showed a significant result, we used 2X2 tests to determine which pairs of sites were different. Probability values were not. corrected for multiple tests. We employed an a-level of 0.05. RESULTS No plants "changed species" during the four years of the study. The few plants that we previously reported as changing from one species to another were based on misidentif ications. Sample population sizes in 1990-92 were relatively stable; all three species had slightly more plants in 1992 compared to 1990 (Fig. 1) . Survivorship of the 1989 sample population was highest for Botrychium hesperium. lowest for B^ paradoxum and intermediate for B^ watertonense (Fig. 2) . Mortality was higher in B^ paradoxum compared to B_j. hesperium for all three years measured, and this difference was statistically significant in two of those years (chi-square test, P<0.05; Fig. 3). Mortality in B^ watertonense was more variable than the other two species, as high as B^ paradoxum in one year and lower than B^ hesperium another (Fig. 3) . Recruitment among the three species showed a pattern very similar to mortality. Recruitment was higher in B. paradoxum . and this difference was statistically different in two of the three years (Fig. 4). Recruitment proportion for B_^ watertonense was higher than B^ paradoxum in one year but lower than B_^ hesperium in the two other years (Fig. 4) . DISCUSSION We have obtained no evidence that would indicate that Botrvchium paradoxum and B^ watertonense are anything but morphologically stable species. The shape of the depletion curves for all three species suggests that they are short-lived perennials. However, it should be kept in mind that plants may exist underground as gametophytes for a period of time before emeriging as a mature sporophyte. Nonetheless, compared to Botrychium dissectum, all three species are short lived (Montgomery 1990) . The high rates of mortality and recruitment demonstrated by these three species of Botrychium suggest that they are adapted to occupying early successional sites or ephemeral gaps in more stable vegetation (MacArthur and Wilson 1967, Harper 1977, Solbrig and Solbrig 1979) • Botrychium paradoxum has the highest rates of mortality and recruitment, indicating that it has the most volatile demographic regime and suggest that of all three species, it is the most adapted to ephemeral habitats and is the most prone to cycles of extinction and recolonization (Diamond 1984, Pimm et al. 1988) . Botrvchium hesperium is longer lived and had lower levels of recruitment and mortality for all years measured compared to B. paradoxum. These differences in life history must be at least partially genetically determined since the two populations are growing in the same environment. Hybridization may result in the breakup or restructuring of coadapted gene complexes, and breakup of integrated genetic systems often results in increased additive genetic variation (Carson 1975, Templeton 1980) . Morphological and chemical evidence indicates that B. watertonense is a hybrid between these two species (Wagner et al. 1984) . The life history characteristics of the hybrid plants are more variable and encompass more than the range of either of the putative parent species. Our results provide evidence that balanced genetic systems interacting with environment to produce life history traits in the parent Botrychium species have been disrupted in the hybrid. LITERATURE CITED Carson, H. L. 1975. The genetics of speciation at the diploid level. American Naturalist 109: 73-92. Diamond, J. M. 1984. "Normal" extinctions of isolated populations. Pages 194-246 in M. H. Nitecki, (ed.) Extinctions. University of Chicago Press, Chicago, Illinois. Harper, J.L. 1977. Population Biology of Plants. Academic Press, London. Lellinger, D. B. 1985. A field manual of the ferns and fern- allies of the United States and Canada. Smithsonian Institution Press, Washington D.C. Lesica, P. 1987. A technique for monitoring nonrhizomatous perennial plant species in permanent belt transects. Natural Areas Journal 7: 65-68. MacArthur, R. H. and E. 0. Wilson. 1967. The theory of island biogeography. Princeton University Press, Princeton, NJ. Montgomery, J. D. 1990. Survivorship and predation changes in five populations of Botrvchium dissectum in eastern Pennsylvania. American Fern Journal 80: 173-182. Pimm, S. L. , H. L. Jones and J. Diamond. 1988. On the risk of extinction. American Naturalist 132: 757-785. Solbrig, T. and D. J. Solbrig. 1979. Introduction to population biology and evolution. Addison-Wesley Publishing, Reading, Massachusetts. Templeton, A.R. 1980. The theory of speciation via the founder principle. Genetics 94: 1011-1038. Wagner, W. H. and F. S. Wagner. 1981. New species of moonworts, Botrvchium subg. Botrvchium (Ophioglossaceae) , from North America. American Fern Journal 71: 20-30. Wagner, W. H. , F. S. Wagner, C. Hauffler and J. K. Emerson. 1984. A new nothospecies of moonwort Ophioglossaceae, Botrvchium^ . Canadian Journal of Botany 62: 629-634. Figure 1. Number of Botrychium hesperium, B. paradoxum and B^ watertonense plants in sample populations in 1990-92. 200 00 c _D CL CD E 13 150 100 50 0 • hesperium V paradoxum ▼ watertonense 90 91 Year 92 Figure 2. Depletion curve for 1989 sample populations of Botrvchium hesperium. B. paradoxum and B^ vatertonense. if) i_ O > > C O i_ 100 80 - 60 40 20 0 • hesperium V paradoxum ▼ watertonense 89 90 91 Year 92 Figure 3. Proportion of plants dying in three consecutive 1- years intervals for Botrychium hesperiuro, B. paradoxum and B. watertonense populations. For each interval, values annotated by different letters are significantly different by chi-square tests (P<0.05, see Methods). c o 0.7 0.6 0.5 §- 0.4 o ^ 0.3 o 0.2 0.1 - 0.0 1 1 • 1 hesperium a V paradoxum _ T, T Avatertonense a ■■■<: - ■.. a ■■■.. — -.-.^_^ a T " V b ah a • — , b ^,-V' • ~~ \ - 1 1 T 1 - 89-90 90-1 Yeor 91-2 Figure 4. Proportion of new recruits in three consecutive 1- years intervals for Botrychium hesperium. B. paradoxum and B. watertonense populations. For each interval, values annotated by different letters are significantly different by chi-square tests (P<0.05, see Methods). 0.7 0.6 5 0.5 o Q. E 0.4 D. E 0.3 I 0.2 a ' a V 0.1 0.0 b «... V '•"■■■ a T b • hesperium V paradoxum ▼ watertonense 90-1 91-2 Yeor 92-3 9 Appendix A. Morphological codes for individual Botrvchium plants in permanent monitoring transects in 1989-93. See text for explanation of codes. 1989 1990 1991 1992 1993 Lower Line Plot 1 a H-P2- -H30- -Lll H-P2-H39-L17 H-P2-H54-L20 — — b H-P3- -H71- -L38 H-P2-H85-L35 — — — c H-Pl- -H19- -LIO H-P1-H30-L17 H-U H-P1-H24 — d H-Pl- -H24- -LIO H-U H-P2-H38-L11 H-P1-H12 H-P1-H19 e H-Pl- -H26- -LIO H-P1-H34-L14 H-P1-H26-L11 — — f P-Pl- -H51- -L15 P-P1-H60-L12 P-P1-H61-L16 P H51 P H81 g — H-P2-H25-L11 H-U H-P1-H20 H-P1-H50 h — H-P1-H15-L18 — — — i -- P-P1-H13-L08 -- — — j — H-P2-H50-L24 H-P~H13-L— — — k — H-P2-H70-L31 H-P2-H56-L16 H-P1-H40 H-P2-H89 1 -- H-P1-H60-L15 -- -- — m — H-P2-H54-L17 H-P2-H45-L14 — — n -- H-P1-H37-L13 H-P2-H49-L21 H-P1-H24 H-P1-H40 o — H-P2-H32-L12 H-P1-H29-L11 H-U H-P1-H27 P — H-P1-H21-L03 H-P1-H15-L05 H-U H-P1-H38 q -- H-P1-H16-L08 — — — r — P-P1-H27-L06 P-P1-H18-L03 P-U P H26 s — P-P1-H28-L10 P-U P H45 P H62 t — P-P1-H31-L09 -- — -- u -- H-P1-H32-L11 H-P2-H38-L13 H-P1-H25 H-P1-H29 V — H-P2-H60-L22 -- — -- w — H-P2-H50-L20 H-P3-H81-L29 H-P2-H61 H-P2-H79 X — H-P1-H22-L07 H-P2-H41-L21 H-P2-H38 H-P2-H50 y -- P-P2-H100-L35 P-P1-H79-L19 — — z — P-P1-H55-L12 P-P1-H73-L28 P H54 P H95 aa — P-P1-H45-L11 P-U P H59 P H103 bb — P-P1-H52-L15 — — — cc — — H-P2-H45-L10 — — dd — — P-P1-H20-L0 — — ee -- . -— P-P1-H49-L11 P-U P H56 ff -- — P-P2-H81-L22 — -- gg — — H-P2-H51-L15 H-P1-H21 H-P1-H37 hh -— . —^ H-P2-H50-L16 — — ii -- — _ H-P2-H53-L20 H-U H-P1-H46 j j — — — H-P2-H50-L18 H-P1-H42 H-P1-H50 kk — — H-P2-H63-L15 H-P1-H32 H-P1-H40 11 — — H-P2-H38-L19 — — mm — — H-P1-H31-L08 — — nn — — — H-P1-H24-L08 H-P1-H16 H-P1-H32 oo — — H-P1-H25-L11 — — PP — -> — H-P1-H44 — qq — — . — H-P1-H23 H-P1-H49 rr — — — H-Pl-Hll — ss — — -- H-P2-H56 H-P1-H104 tt — — — H-P1-H09 — uu — — — — H-P1-H35 — vv — — — H-P1-H16 H-P1-H48 WW — — -- -- P H42 XX — — — — P H51 yy — — — — P H55 zz — — — — P H53 aaa — — — — H-P1-H46 bbb — — — H-P1-H37 ccc — -— -- — H-P2-H65 10 ddd eee fff ggg hhh iii jjj kkk mmm Plot 2 a P-P1-H63-L22 P-U P-P1-H22-L06 P H27 b H-P1-H35-L15 c P-P1-H55-L17 d P-P1-H62-L25 e P-P1-H29-L05 P-P1-H40-L70 f H-P1-H21-L08 g H-P1-H37-L10 H-P2-H40-L19 h — H-P1-H38-L09 i — P-P1-H67-L20 j — P-P1-H53-L16 k ~ H K40-L— I — H-P1-H25-L05 m — H-P1-H33-L04 n — H-P1-H24-L05 o — H-P1-K20-L08 p ~ H-P1-H35-L09 q — P-P1-H38-L07 r -- P-P1-H14-L03 s — H-P2-H45-L21 t — H-P1-H21-L08 u — P-P1-H45-L07 V — P-P2-H94-L22 w — H-P2-H41-L16 X — H-P2-H45-L13 y — P-P2-H110-L35 z — P-P1-H70-L19 aa — P-P1-H52-L12 bb — cc — — dd — ee — — ff ~ gg — hh ~ ii — — jj ~ kk — II ~ mm — — nn — — oo — — pp ~ gq ~ rr — — ss tt — uu — vv WW XX — 11 H-P1-H20 H-P1-H25 P H42 P H41 P H37 P H26 H-P1-H28 H-P1-H15 111 __ — — ~ H-P1-H36 H-P1-H40 H-U H-U H-P1-H21 H-P2-H46-L03 H-U H-P2-H80 P-P1-H67-L16 — — P-P1-H26-L10 -- — H-U H-U H-P2-H70 H-P1-H26-L05 — — H-P2-H41-L11 — — H-P1-H22-L04 H-U H-P1-H55 K-P1-H28-L11 — — — P-U P-U P H40 H-P1-H24-L06 H-U H-P1-H36 P-U P-U P H67 H-P2-H46-L13 — — H-U H-Pl- -H25 H-P2-H62 P-P2-H90-L25 P-U P H60 P-P1-H55-L12 — — P-P1-H25-L05 P-U P H50 H-P1-H21-L04 — — P-P1-H32-L05 — — H-P2-H46-L03 H-U H-P1-H60 P-P2-H50-L22 — — H-P2-H41-L20 H-U H-P2-H79 H-P1-H11-L09 H-U H-P1-H36 P-P2-H61-L18 P-U P H95 P-P1-H13-L10 — — P-P1-H14-L07 P -H22 -- H-Pl-HlO-LOl — — H-P1-H25-L04 — — H-P1-H28-L08 — — H-P1-H22-L08 H-U H-P1-H27 H-P1-H33-L18 H-Pl- -H25 — — W-Pl- -H40 — — H-Pl- -H38 H-P2-H73 — H-Pl- -H37 H-P1-H47 — — H-P1-H19 — — P H44 — — H-P1-H30 — — H-P1-H53 — — P H59 yy zz aaa bbb ccc ddd eee fff ggg hhh iii P H80 P H35 H-P1-H38 H-P1-H37 H-P1-H55 H-P2-H52 H-P1-H35 H-P1-H39 H-P1-H30 H-P1-H64 P H58 Plot 3 a b c d e f g h i — J k 1 m — n o P q r s — t u — V — w — X H-P1-H40-L10 P-P1-H65-L10 P-P2-H43-L16 P-P1-H41-L14 H-P1-H20-L05 P-P1-H73-L20 P-P1-H50-L12 P-P1-H33-L12 P-P1-H29-L09 P-P1-H24-L05 P-P1-H56-L10 P-P1-H32-L08 P-P1-H64-L09 P-P1-H31-L13 P-U P H29 P-U P H39 p — -H17 p — -H35 — p — -H45 — p — -H45 P H65 — P H81 — P H57 -- P H22 — P H75 — P H88 — P H70 — P H75 — P H66 — P H99 — P H19 — P H43 Plot 4 a P-P1-H63-L23 H-P1-H37-L10 b c d e f g h i J k 1 m n o P q r s t u V w P-P2-H76-L30 H-P2-H60-L23 H-P1-H36-L07 H-P1-H62-L21 H-P2-H61-L16 P-P1-H60-L14 P-P2-H40-L15 P-P1-H17-L06 H-P2-H32-L22 P-P1-H44-L16 H-P2-H69-L25 P-P1-H28-L05 H-P2-H35-L10 H-P2-H71-L31 H-U H-Pl- -H38 H-P1-H71 H-P2-H53-L15 H-Pl- -H43 H-P1-H44 P-U P -H45 — P-U P-U P HBO P-P1-H23-L08 — — H-P2-H75-L40 H-P2- -H57 H-P2-H66 H-P2-H68-L31 H-Pl- -H72 H-P2-H112 P-P1-H28-L08 P-U P H20 P-P1-H40-L06 P-U P H30 P-P1-H62-L18 P-U P H30 P-P1-H31-L09 P-U P H44 P-P1-H90-L19 P-U P H40 P-P1-H45-L06 P-U P H33 H-P2-H70-L28 — — P-P1-H70-L20 — — P-P1-H30-L10 p -H62 P— -H82 H-P1-H29-L10 — — 12 X — — H-P1-H40-L06 y P-P1-H65-L01 P H26 z — — P-P1-H41-L06 aa — — H-P1-H77-L18 bb — ~ P-P1-H41-L12 cc — — — H-P1-H40 H-P2-H59 dd — — — P H32 ee — — — H-PI-HIO H-P1-H20 ff — — — P H67 gg — — — P H22 P H22 hh — — — H-P1-H22 H-P1-H45 ii — — — H-P1-H27 jj — — — H-P1-H40 kk — — — — P H32 11 — — — -- P H82 mm — — — — H-P1-H30 nn — — — — P H30 00 — — — — P H75 pp — — — — P H61 qg — — — — p H40 rr — — — — P H45 ss — — — — P H41 tt — — ~ — P H18 uu — — — . — P H15 vv — — — — H-P1-H24 WW — — — — p H50 XX — — — — P H17 yy — — — — p H19 zz — — -- — p H25 Plot 5 a P-P1-H60-L17 P-U P-P1-H75-L16 b P-P1-H30-L08 c P-P1-H31-L05 P-U P-Pl-Hll-LOl d H-P2-H76-L40 H-P1-H37-L15 H-U H-P1-H40 e H-P2-H26-L05 H-P2-H40-L15 H-U H-P1-H31 H-P1-H31 f H-Juvenile H-P2-H38-L17 H-P2-H50-L09 H-U . H-P2-H57 g W-P1-H45-L12 W-U W-P1-H64-L18 W-P1-H45 W-P2-H72 h H-P2-H42-L14 i H-P1-H38-L15 H-P2-H42-L15 H-U H-P2-H48 H-P3-H91 j H-P2-H50-L20 H-P1-H26-L04 H-P1-H46-L05 k H-P2-H52-L21 H-P2-H54-L22 H-P1-H30-L11 1 H-P2-H47-L22 H-P2-H50-L25 H-U H-P1-H18 H-P1-H42 m H-P1-H11-L08 n — H-P1-H30-L08 H-P1-H34-P08 o — H-P1-H29-L04 H-P1-H32-L08 H-P1-H33 H-P1-H32 p ~ P-P2-H59-L16 P-P2-H61-L17 q — H-P2-H68-L36 H-U H-P1-H38 H-P1-H53 r — P-P1-H58-L07 P-U P-U P H59 s — P-P1-H40-L09 P-P1-H27-L03 P H17 P H52 t — P-P1-H20-L03 u — P-P1-H49-L10 V — P-P1-H31-L09 w — P-P2-H50-L16 X — H-P2-H77-L30 H-P2-H65-L18 H-P2-H41 y — H-P1-H30-L06 2 — stem 1 H-P1-H19-L05 H-P2-H49-L24 H-P1-H27 H-P1-H49 aa —stem 2 H-P2-H32-L08 H-P1-H11-L03 H-P2-H53 H-P1-H42 t>b — W-P1-H61-L18 W-U W-P1-H41 W-P1-H52 cc — — P-P1-H47-L11 — • dd — P-P1-H31-L06 ~ — ee — H-sterile leaf H-P1-H12-L03 H-P1-H21 H-P2-H52 13 p — -H62 P H70 P-U P H40 H-Pl- -H35 H-P1-H65 P-U P H67 p -H47 — H-Pl- -H40 — H-Pl- -H32 — H-Pl- -H27 H-P1-H33 H-Pl- -H21 H-P1-H43 p -H89 P H86 p -H47 — -- H-P1-H16 — P H104 — P H27 — P H56 -- H-P1-H29 -- W-P1-H40 — P H23 — W-P1-H43 — H-P2-H63 — H-P2-H65 — H-P1-H32 ff — H-P2-H56-L26 H-P2-H48-L28 gg — H-P3-H80-L31 H-P2-H72-L32 H-P1-H22 H-P2-H57 hh — P-P1-H35-L06 P-P1-H10-L02 ii — H-P1-H35-L08 jj — — P-P1-H29-L03 kk — — H-P1-H31-L06 11 — — P-P1-H18-L04 mm — — P-P1-H40-L07 nn — — H-P1-H30-L10 oo — — H-P1-H22-L10 pp — — P-P1-H53-L18 qq — — P-P1-H53-L18 rr — ~ ss — ~ tt — uu — — WW -- -- XX — yy — zz -- — aaa -- — bbb ~ ccc — — ddd — eee — fff — ggg — hhh ~ iii — — Upper Line Plot 1 a W-P2-H55-L17 W-P2-H68-L27 W-P1-H47-L14 b H-P2-H57-L35 C — H-P2-H65-L30 H-P2-H65-L24 d — W-P1-H68-L18 W-P1-H39-L13 W-P1-H58 W-P1-H80 e — H-P2-H80-L26 H-U H-P1-H35 H-P10-H65 f — H-P3-H89-L53 g — H-P1-H35-L08 H-P2-H67-L34 H-P2-H66 H-P2-H87 h — W-P2-H60-L21 W-P2-H76-L23 W-P1-H53 W-P2-H113 i — W-P3-H70-L29 W-P2-H51-L16 W-U W-P2-H80 j — P-P1-H42-L13 k — H-P2-H41-L14 H-P2-H46-L14 H-P2-H54 H-P2-H80 1 — H-P1-H65-L18 H-P2-H67-L19 m — H-P1-H17-L07 ~ H-P1-H18 H-P2-H25 n — — H-P1-H33-L29 H-P2-H40 H-P2-H71 O — — H-P1-H31-L09 p — — P-P1-H19-L05 q — ~ — H-P1-H40 r — — — P H66 s — — — P H45 t — — — W-P1-H77 W-P1-H57 u — — -- H-P1-H28 H-P1-H33 V — — -- W-P1-H92 W-P2-H128 w — — — W-P1-H47 W-P1-H70 X — — — H-P1-H39 H-P1-H57 y — — — p H36 z — — — P H34 aa — — ~ H-P1-H35 bb ~ ~ ~ H-P1-H51 14 cc dd H-P1-H44 H-P1-H30 H-P1-H18 H-P2-H54 ee — -• - ff — — H-P1-H26 gg hh ii jj kk 11 mm nn oo PP qq rr H-P2-H77 H-P2-H85 P H28 P H48 H-P2-H74 W-P1-H45 H-P2-H95 H-P1-H56 H-P2-H94 H-P1-H45 H-P1-H54 H-P2-H65 ss — — — H-P1-H38 tt — — ~ — H-P1-H42 Plot 2 a H-P2-H69-L30 H-P2-H75-L30 b H-P2-H66-L23 H-P2-H66-L19 C H-P2-H44-L14 H-P1-H40-L12 d H-P2-H57-L23 H-P2-H75-L31 e H-P2-H31-L07 f H-P3-H51-L30 H-P2-H67-L30 g H-P3-H66-L38 H-U h H-P2-H45-L20 H-P3-H81-L33 i H-P2-H42-L18 H-P2-K41-L17 j H-P2-H34-L13 H-P2-H30-L11 k H-P3-H75-L45 H-P3-H90-L53 I H-P3-H75-L49 H-P3-H75-L35 m — H-P2-H63-L41 n — H-P2-H69-L26 o — H-P2-H65-L20 p — H-P1-H34-L13 q — H-P1-H49-L20 r — H-P1-H41-L12 s — P-P1-H62-L14 t — P-P1-H68-L16 u — P-P1-H91-L25 V — H-P1-H45-L10 w — W-P1-H40-L11 X — H-P2-H90-L51 y — H-P2-H42-L11 z — H-P1-H26-L11 aa — H-P2-H61-L32 bb — H-P2-H60-L24 cc — dd ~ ee ff ~ gg — hh ~ ii jj __ kk ~ II — mm nn oo — PP ~ 15 H-P2-H54-L11 H-P1-H35 H-P2- -H61 H-P1-H25-L14 H-P1-H32 -- H-P2-H64-L22 H-P1-H52 H-P2- -H73 H-P2-H49-L19 H-U H-P2- -H45 H-P2-H60-L27 H-P1-H46 H-P2- -H34 H-P2-H43-L15 H-P2-H55 H-P2- -H114 H-U H-U H-P2- -H55 H-P2-H38-L10 H-P1-H20 H-P3- -H93 H-P3-H57-L34 H-P2-H79 -- H-P3-H69-L27 — — H-P2-H53-L18 H-P1-H35 — H-P2-H64-L17 — -- H-P2-H89-L38 H-P2-H86 H-P3- -H105 H-P2-H51-L18 H-P1-H50 H-Pl- -H62 H-P2-H37-L18 — — P-P1-H82-L25 P H38 — — P-U P H83 — W-U W-P1-H32 — H-P3-H84-L35 H-P2-H46 H-P2- -H69 H-P1-H31-L08 H-P1-H29 H-Pl- -H41 H-U H-P2-H58 H-P2- -H51 H-P2-H38-L13 H-P1-H35 H-P2- -H85 H-P3-H81-L23 H-P2-H73 — H-P2-H75-L20 H-P2-H54 H-P2- -H80 H-P2-H50-L14 — — P-P1-H49-L09 P H54 p -H75 P-P1-H108-L20 P H59 — H-P2-H44-L11 H-P2-H52 H-P2- -H55 H-P1-H25-L04 H-P1-H19 — , H-P3-H80-L30 — — P-P1-H42-L02 P-U p -HBO W-P1-H63-L10 W-U W-Pl- -H60 P-P1-H49-L10 P H54 p -H66 P-P1-H38-L07 — — H-P1-H34-L09 — — P-P1-H31-L04 P H46 P -H40 gq — — — H-P2-H57- -L21 H-P1-H50 H-P2-H55 rr — — — — H-P1-H35 — 55 — — — — H-P1-H25 — tt — — -- H-P2-H52 — UU — -— -- W-P1-H32 — vv — — H-P1-H29 — WW — — -- H-P1-H44 — XX — -- P H55 — yy — — — P H62 — zz -— — H-P1-H46 — aaa _- — — H-P1-H40 bbb __ __ — — H-P2-H75 ccc — _ — — H-P2-H25 ddd ._ __ — — H-P1-H53 eee ._ __ — — H-P2-H76 fff _— — — P H86 999 __ — — H-P1-H42 hhh __ — — H-P2-H70 iii — — -- — H-P3-H134 J J J — — -- — P H93 kkk __ — -- H-P2-H98 111 __ — — H-P2-H72 mnun __ — -- H-P2-H46 nnn m.— — . -- W-P1-H21 ooo — — — -- H-P1-H55 PPP — — — H-P1-H36 999 — — -- -- H-P2-H50 rrr __ — — P H45 sss — — — W-P1-H51 Plot 3 a H-P2- -H26- -LIO H-Pl- -H54- -L14 H-P2-H41- -LIO H-P1-H23 — b H-P2- -H55- -L22 H-U H-U H-P1-H45 — c H-P2- -H86- -L31 H-P2- -H81- -L31 H-P3-H44- -L15 H-P2-H92 H-P2-H86 d H-P2- -H41- -L14 H-U H-P2-H53- -L18 H-U H-P1-H57 e H-Pl- -H50- -L14 H-U H-P3-H89- -L28 H-P2-H48 H-P2-H77 f — H-Pl- -H36- -L04 H-U H-P2-H87 — g — P-Pl- -H67- -L24 — — — h — - P-Pl- -H81- -L17 — -- — i — H-Pl- -H51- -L12 — — — j — — W-Pl- -H55- -L12 — — — k — — H-Pl- -H45- -L17 H-U H-Pl-H?? — 1 — — P-P2- -H55- -L32 -- — ~ m » H-P2- -H19- -L07 H-P1-H20- -L17 — — n _— H-Pl- -H52- -L14 H-U H-P2-H48 H-P2-H66 o — — H-P2- -H57- -L22 — — — p __ — — H-P1-H49- -L13 H-P1-H45 H-P2-H75 q — H-P1-H30- -L18 H-P1-H30 H-P1-H45 r — — — H-P2-H38- -L09 H-P1-H43 — s — — — H-P2-H54- -L16 H-P1-H44 — t __ — — H-P2-H39- -L04 H-P1-H70 H-P2-H105 u — — -- H-P1-H28- -LOS H-U H-P1-H12 V — — H-P2-H29- -L15 — — w — — -- H-P2-H28- -LOl — — X — _- H-P3-H55- -L31 H-U H-P2-H55 y __ — P H20 — z __ H-P1-H24 H-P1-H47 aa __ .» P H31 P H41 bb — — — H-P1-H22 P H52 cc -— — — H-P1-H23 — dd — — — P H40 ee — -— — P H37 — 16 ff — — ~ P H29 H-P1-H26 H-P2-H52 gg hh ii -- jj kk ^ ^ 11 — mm — nn — oo — PP — qq — rr — ss -- tt — uu — vv — WW — Plot 4 a H-P2' -H19' -L09 b H-P2' -H32' -Lll c H-P2' -H31- -L15 d — e -- f — g h ^_ i — J k ^*. 1 — m — n — o — P — Plot 5 a H-P3- -H45- -L28 b P-P2- -H37- -L18 d W-Pl- -H42- -L12 e W-P2- -H49- -L17 f H-Pl- -H30- -Lll g H-P2- -H43- -L15 h — i — J k ^^ 1 — m — n — o — P — q -- r — s — t — u — V — w — X — y — P H18 H-P2-H56 H-P2-H42 H-P2-H38 H-P1-H49 H-P1-H44 W-P2-H102 P H31 H-P1-H45 P H31 P H29 H leaf H-P1-H62 H-P1-H23 H-P2-H55 H-P1-H47 H-P1-H28-L13 H-P2-H24-L09 ' H-U H-P1-H19 H-P2-H36-L16 H-P2-H33-L15 H-P1-H29 H-P1-H49 H-P2-H65-L27 H-P2-H67-L18 H-P1-H28 H-P2-H60 H-P1-H25-L08 H-P1-H21-L03 H-U H-P2-H35 P-P1-H27-L08 P-U P H15 H-P3-H61-L18 H-P1-H40 H-P2-H63 H-P2-H59-L16 H-U H-P1-H45 P-P1-H25-L06 H-P1-H27-L09 H-P1-H15 H-P1-H32 H-P1-H25 H-P2-H67 p H22 H-P1-H31 H-P1-H50 p H24 H-P1-H69 H-P1-H50-L16 H-P2-H43-L10 H-P1-H48-L15 H-P1-H49-L19 H-P1-H42-L15 H-P1-H36-L11 H-P1-H35-L14 H-P1-H45 H-P1-H41 . H-sterile leaf — — — H-P2-H57-L18 H-P1-H20-L02 H-P3-H82-L36 H-P3-H140-L58 H-P2-H124 H-P2-H142 H-P2-H78-L25 H-P3-H65-L25 H-P2-H90-L50 H-P1-H29-L14 H-P1-H21 H-P2-H60 W-P1-H39-L10 W-P1-H41-L11 W-P1-H34-L09 H-P2-H68-L40 H-P1-H32-L13 H-P3-H51-L24 H-U H-P1-H31 H-P2-H40-L15 H-P2-H38-L14 H-P2-H42-L27 H-P3-H74-L50 H-P1-H44-L16 H-P2-H76 H-P2-H51 H-P2-H50-L30 H-U H-P2-H55 17 z — aa — bb — cc — dd — ee — ff — gg — hh — ii — jj — kk — 11 — mm — nn — oo — PP — qq — rr — ss — tt — uu -- vv — WW — XX — yy — zz ■"■" Plot 6 a H-P2' -H49- -L15 b H-P2' -H30- -L17 c H-P2' -H83- -L27 d W-P2- -H89- -L36 e W-P2' -H44- -L17 f W-P3- -H83- -L30 g W-P2- -H44- -L22 h W-P2- -H42- -L37 i W-P2- -H52- -L36 J — k — 1 — m — n — o — P — q — — r — - s — t — u ~ V — w — X ~ y — z — aa — Plot 7 a H-P2- ■H34- ■L12 b H-P2- ■H31- ■LOS c H-P2- ■H20- ■L06 d H-P3- •H40- ■LIO H-U W-P2-H97-L41 W-P2-H55-L23 H-P2-H95-L42 H-P2-H65-L38 H-P3-H80-L52 H-sterile leaf W-P2-H74-L26 H-P3-H110-L41 H-P2-H66-L21 H-Pl- -H35- -Lll H-Pl- -H18 H-P1-H46 H-Pl- -H19- -L03 H-U H-P1-H31 H-P3- -H76- -L29 H-Pl- -H48 — H-Pl- -H18- -L03 H-Pl- -H21 H-P1-H12 P-Pl- -H39- -L08 P-U P H27 H-Pl- -H34- -L08 — — H-P3- -H68- -L37 H-U H-P2-H93 H-P2- -H36- -L20 H-U H-P1-H21 H-P3- -H56- -L22 H-P2- -H50 H-P2-H68 H-Pl- -H13- -L02 H-Pl- -H16 — H-Pl- -H28- -LOS H-Pl- -H20 -- P-Pl- -H09- -L03 — — — H-P2- -HS2 — -- p -H17 — — H-Pl- -H45 — — H-Pl- -H21 — -- H-P2- -H46 -- — H-Pl- -H4S -- -- H-P2- -H75 -- — — P HIS -- — W-P1-H30 -- — H-P1-H21 -- H-P2-H8S — H-P2-H58 -- — H-P1-H20 — — H-P2-H62 — — P H34 H-U W-P2-H71-L29 W-P2-H56-L21 H-P3-H105-L39 H-P2-H72-L35 H-P3-H78-L46 H-P3-H10S-L34 H-P2-H41-L21 H-P1-H63-L16 P-P1-H62-L26 H-U W H41 W H44 H-P2-H88 H-P1-H66 H-P2-H76 H-P2-H57 H-P1-H40 H-P2-H64 H-P1-H3S H-P1-H41 H-P1-H52 W-P1-H18 W-P1-H22 H-P2-H80 H-P2-H98 H-P2-H96 H-P2-H91 H-P1-H29 H-P1-H40 H-P1-H48 H-P2-H115 H-P1-H64 H-P1-H52 H-P2-H71 H-P3-H38-L20 H-P2-H28-L11 H-P2-H40-L14 H-P~H21-L — H-P2-H30-L10 H-P1-H22 H-P1-H35 18 H-P2-H42-L18 -- — — - H-P2-H26-L16 H-P2-H34-L10 H-Pl- -H28 -- H-P2-H64-L26 H-P2-H68-L27 H-Pl- -H56 H-P2- -H56 W-P2-H42-L15 W-P1-H45-L14 W-U W-Pl- -H39 H-P2-H56-L20 -- -- — H-P1-H22-L05 H-P1-H30-L09 H-Pl- -H15 H-Pl- -H25 P-P1-H50-L17 -- — — H-P3-H106-L38 H-P3-H68-L19 H-P2- -H62 H-P2- -H72 H-P2-H52-L14 — — — P-P1-H17-L06 P-P1-H32-L08 P-U p -H31 H-P1-H23-L07 H-P1-H31-L05 — — W-P2-H41-L20 — — — H-P3-H56-L31 — — — H-P1-H30-L10 — — — H-P2-H43-L26 -- — — H-P2-H65-L31 H-P2-H41-L18 H-U H-Pl- -H36 H-P1-H26-L06 H-P2-H51-L21 — — e H-P3-H45-L22 f H-P2-H33-L11 g H-P2-H40-L16 h H-P2-H34-L13 i H-P2-H59-L29 j W-P2-H35-L14 k H-P2-H64-L28 1 H-P2-H43-L17 m P-P1-H52-L15 n o P q r s — t u — V — w — X — — P-P1-H34-L07 y — — H-P1-H16-L02 2 ~ — H-P2-H57-L23 H-P1-H40 H-P1-H33 aa — — — W-P1-H36 bb — — -- H-P1-H40 cc — — -- W-P1-H52 dd — — ~ — H-P1-H13 ee — — — — H-P1-H19 ff — — — — P H46 Plot 8 a W-P4-H126-L73 — b H-P3-H96-L34 H-P2-H79-L33 c H-P3-H78-L30 H-P2-H42-L11 H-P2-H72-L20 H-P1-H25 d H-P3-H85-L31 e H-P2-H49-L17 H-P2-H38-L12 H-P1-H17-L04 f H-P2-H31-L10 g P-P1-H10-L03 h H-P1-H29-L16 i H-P3-H54-L26 H-P3-H75-L32 j H-P2-H39-L15 k — H-P3-H65-L21 1 — P-P1-H92-L23 m — W-P2-H54-L16 n — W-P2-H140-L67 o — W-P1-H49-L05 p — P-P1-H35-L05 g — W-P1-H51-L13 r — W-P1-H32-L04 s — — y aa — -- bb — - — . cc — — dd ~ 19 H-P3- -H90-L34 H-U H-P1-H46 P-U P-U P H48 W-P3- -H114-L53 W-P1-H120 W-P2-H53 P-Pl- -H44-L16 — W-Pl- -H32-L12 W H34 W-P1-H38 W-U W-P1-H56 W-P2-H95 H-Pl- -H48-L18 H-P1-H46 — H-P3- -H107-L30 H-P2-H80 H-P3-H57 H-P2- -H40-L21 H-P1-H45 — W-P3- -H80-L54 W H84 W-P3-H118 H-P2- -H41-L15 — — — W H58 W-P2-H80 — W H59 W-P2-H81 — W-P1-H55 W-P2-H79 -- W H31 W-P2-H76 — H-P1-H60 H-P2-H92 — H-P2-H65 H-P2-H80 — H-P1-H19 H-P1-H27 MONTANA STATE 'x This «cover" page added by the Internet Archive for formatting purposes