ANALYSIS OF MORPHOLOGY, GROWTH RATE, AND FRAGMENTATION OF
THE ENDANGERED LICHEN SPECIES CLADONIA PERFORATA
by
David Warren Witmer
A Thesis Submitted to the Faculty of
The Charles E. Schmidt College of Science
In Partial Fulfillment of the Requirements for the Degree of
Master of Science
Florida Atlantic University
Boca Raton, Florida
May 2014 ANALYSIS OF MORPHOLOGY, GROWTH RATE, AND FRAGMENTATION OF
THE ENDANGERED LICHEN SPECIES CLADONIA PERFORATA
by
David Warren Witmer
This thesis was prepared under the direction of the candidate's thesis advisor, Dr. Jon Moore, Department of Biology, and has been approved by the members of his supervisory committee. It was submitted to the faculty of the Charles E. Schmidt College of Science and was accepted in partial fulfillment' of the requirements for the degree of Master of Science.
Jon Moore, P,.D. Thesis Adviso'r 41,', -1'~ "- ,~ ~'~'I'f~T~~~ -- '-'._ / Susan Richardson, Ph.D. ~ Xing-Hai Zhang, Ph.D.
Dale E. Gawlik, Ph.D. Sciences Program
Russell Ivy, Ph.D. Inte' Dean, Charles E. Schmidt College of Science J(~IJ;(;~~ eborah L. Floyd,Ed.D. Interim Dean, Graduate Studies
ii ACKNOWLEDGEMENTS
The author wishes to express his sincere thanks and love to his fiancée for her support and encouragement throughout the research and manuscript preparation processes. The author is also grateful to Palm Beach County’s Environmental Resources
Management Department for permission to conduct his research on the Jupiter Ridge
Natural Area.
iii ABSTRACT
Author: David Warren Witmer
Title: Analysis of Morphology, Growth Rate, and Fragmentation of the Endangered Lichen Species Cladonia perforata
Institute: Florida Atlantic University
Thesis Advisor: Dr. Jon Moore
Degree: Master of Science
Year: 2014
Cladonia perforata is an endangered lichen endemic to the Atlantic Coastal
Ridge, Lake Wales Ridge, Southwest Florida, and the North Gulf Coast of Florida. In all but a single locality, C. perforata relies entirely on asexual reproduction through fragmentation for reproduction, dispersal, and recruitment. This study suggests a positive correlation between fragment size and survivability of fragments after one year. The average thallus grew at a rate of 10.42% per year and younger branches of a thallus grew at a quicker rate than older branches. Additionally, a review of thalli morphology suggests C. perforata has a diverse form, and becomes more bifurcated as it increases in size.
iv ANALYSIS OF MORPHOLOGY, GROWTH RATE, AND FRAGMENTATION OF
THE ENDANGERED LICHEN SPECIES CLADONIA PERFORATA
LIST OF TABLES ...... vi LIST OF FIGURES ...... viii LIST OF EQUATIONS ...... ix 1: INTRODUCTION ...... 1 2: STUDY AREA ...... 4 3: MORPHOLOGY ...... 6 Methods and Materials ...... 6 Results ...... 7 Discussion ...... 10 4: GROWTH RATE ...... 12 Methods and Materials ...... 12 Results ...... 14 Discussion ...... 16 5: FRAGMENTATION ...... 19 Methods and Materials ...... 19 Results ...... 21 Discussion ...... 22 APPENDIX ...... 24 Morphology ...... 24 Growth Rate ...... 37 Fragmentation...... 53 BIBLIOGRAPHY ...... 77
v LIST OF TABLES
Table 3.1 Number of Branches in Each Branch Order ...... 30 Table 3.2 Number of Tributary Branches in Each Branch Order ...... 31 Table 3.3 Number of Source Branches in Each Branch Order ...... 32 Table 3.4 Wet Weight, Length, Base Diameter, Branch Orders, and Area (Thalli 1-25). 33 Table 3.4 Continued (Thalli 26-50) ...... 34 Table 3.5 Wet Weight to Dry Weight Ratio ...... 35 Table 3.6 Weight of Branch Orders and Total Thallus Weight ...... 36 Table 4.1 Length (Thalli 01-25)...... 43 Table 4.1 Continued (Thalli 26-50) ...... 44 Table 4.2 Weight (Thalli 01-25) ...... 45 Table 4.2 Continued (Thalli 26-50) ...... 46 Table 4.3 Length of 2° Branches (Thalli 1-25) ...... 47 Table 4.3 Continued (Thalli 26-50) ...... 48 Table 4.4 Length of 3° Branches (Thalli 1-25) ...... 49 Table 4.4 Continued (Thalli 26-50) ...... 50 Table 4.5 Length of 4° Branches (Thalli 2-32) ...... 51 Table 4.5 Continued (Thalli 33-49) ...... 52 Table 5.1 Fragment Growth (Fragments 1-25) ...... 56 Table 5.1 Continued (Fragments 26-50) ...... 57 Table 5.1 Continued (Fragments 51-75) ...... 58 Table 5.1 Continued (Fragments 76-100) ...... 59 Table 5.2 Fragment Growth (Fragments 101-125) ...... 60 Table 5.2 Continued (Fragments 126-150) ...... 61 Table 5.2 Continued (Fragments 156-175) ...... 62 Table 5.2 Continued (Fragments 176-200) ...... 63 Table 5.3 Fragment Growth (Fragments 201-225) ...... 64 Table 5.3 Continued (Fragments 226-250) ...... 65 Table 5.3 Continued (Fragments 256-275) ...... 66 Table 5.3 Continued (Fragments 276-300) ...... 67 Table 5.4 Fragment Growth (Fragments 301-325) ...... 68 Table 5.4 (Continued (Fragments 326-350)...... 69 Table 5.4 (Continued (Fragments 356-375)...... 70 vi Table 5.4 Continued (Fragments 376-400) ...... 71 Table 5.5 Fragment Growth (Fragments 401-425) ...... 72 Table 5.5 Continued (Fragments 426-450) ...... 73 Table 5.5 Continued (Fragments 456-475) ...... 74 Table 5.5 Continued (Fragments 476-500) ...... 75 Table 5.6 Chi-square Values ...... 76
vii LIST OF FIGURES
Figure 3.1 Bifurcation Ratio ...... 25 Figure 3.2 Average Tributary to Source Ratio...... 25 Figure 3.3 Length to Wet Weight ...... 26 Figure 3.4 Base Diameter to Wet Weight ...... 26 Figure 3.5 Length to Diameter ...... 27 Figure 3.6 Average Thallus Length by Branching Network...... 27 Figure 3.7 Average Thallus Wet Weight by Branching Network ...... 28 Figure 3.8 Dry Weight to Wet Weight...... 28 Figure 3.9 Branch Composition of Total Thallus ...... 29 Figure 3.10 Weight to Surface Area ...... 29 Figure 4.1 Tethered Thallus ...... 38 Figure 4.2 Average Linear Growth ...... 38 Figure 4.3 Average Weight Change ...... 39 Figure 4.4 Thallus With No Branch Division After Two Years ...... 40 Figure 4.5 Branches Grew at Independent Rate ...... 40 Figure 4.6 Percent Growth Rate By Branch Order ...... 41 Figure 4.7 Thallus With Fragmented Branch ...... 41 Figure 4.8 Thallus With Degenerating Base ...... 42 Figure 5.1 Fragment Container ...... 53 Figure 5.2 Artificial Scrub ...... 53 Figure 5.3 Fragment Survivability ...... 53 Figure 5.4 Fragment Mortality Per Quarter ...... 54 Figure 5.5 Average Fragment Lengths ...... 54 Figure 5.6 Average Quarterly Mortality ...... 55 Figure 5.7 Average Percent Weight Change...... 55
viii LIST OF EQUATIONS
Equation 3.1 Bifurcation Ratio ...... 24 Equation 3.2 Tributary to Source Ratio ...... 24 Equation 4.1Percent Change in Length ...... 37 Equation 4.2 Percent Change in Weight ...... 37
ix 1: INTRODUCTION
Lichens are symbiotic conglomerations of fungi and algae, or cyanobacteria.
On April 27th, 1993 Cladonia perforata became the first lichen species to be listed as endangered by the U.S. Fish and Wildlife Services (USFWS 1999). It grows only in xeric scrub, and is only known to occur on thirty eight sites distributed in
Southwestern Florida, the Atlantic Coastal Ridge, Lake Wales Ridge, and the North
Gulf Coast of Florida (USFWS 1999 and 2007; Richardson and Moore 2009).
Cladonia perforata is a fruticose lichen, a term which refers to the lichen’s shrubby growth. The name is derived from the Latin word fruticis meaning shrub.
The podetia grow in tufts, are pale yellowish grey, and have a surface that appears glossy. The primary thallus (usually in the form of flat squamules) is unknown.
Branches may be sub-equal and divergent caused by successive dichotomies and whorls. When this branching process is repeated several times, a sympodium is formed (Evans 1952). The apothecia and spermagonia of C. perforata have only been observed in a single population from the Lake Wales Ridge (USFWS 2007).
Individual podetia are mostly forty to sixty millimeters in height and their larger axis measure three to six millimeters in diameter (Evans 1952); although Richardson and
Moore (2009) found specimens as large as 110 millimeters across. Apical branchlets are shorter and divide into two to four still shorter ultimate branchlets. Wherever a
1 branch divides, a circular opening is formed in the axis. Under a magnifying lens the podetial surface appears uniform and no groups of algal cells can be distinguished
(Evans 1952).
George Llano an Airman serving at Eglin Air Force Base during World War
Two first collected C. perforata in 1945 from Santa Rosa Island, Florida. In 1952, lichenologist Alexander Evans described and named this species. By the late 1980s, a revitalized interest in C. perforata took hold. In 1988, the Florida Natural Areas
Inventory launched a search for C. perforata in the Florida Panhandle (Milius 2000); twenty-seven years after Evans named the species, thirteen new locations in Central
Florida were identified (FWS 2007; Milius 2000). Since then, periodic investigations have continued to find new C. perforata populations throughout the Atlantic Coastal
Ridge (Richardson and Moore 2009), Lake Wales Ridge, and Northern Gulf Coast
(Wilhelm and Burkhalter 1990).
Natural disasters such as hurricanes and anthropogenic perturbations like controlled burns have provided opportunities to learn more about the ecology of
C. perforata. In 1995, Hurricane Opal formed in the Gulf of Mexico. The category four storm made land fall in the Florida Panhandle and impacted a large portion of the C. perforata population in Eglin Air Force Base. The storm destroyed several local sites entirely. Accounts from J. Douglas Ripley, the Air Force’s natural resource manager suggest 70% of the population was washed or blown away (Milius 2000). Rebecca Yahr and Paula DePriest aided in the efforts of saving the remaining C. perforata (Yahr and
DePriest 2005). Yahr created lichen fragment powder by grinding thalli in a food mill. 2 This was not successful in recolonizing suitable habitat (Milius 2000). Subsequently,
Yahr was successful when using un-powdered lichen fragments for re-colonization
(Milius 2000).
In July of 1993, a prescribed fire at Archbold Biological Station impacted three populations of C. perforata. Yahr studied the post fire recovery from January 1997 to
August 1999. All of the other co-occurring terrestrial lichen species recovered from juvenile stages, while no juvenile forms of C. perforata were documented. The area inhabited by C. perforata increased by 200%. However, this was a result of dispersal of preexisting thalli to newly suitable areas. Population growth was too slow to be documented over that time period (Yahr 2000).
The 2004 hurricane season created an excess of vegetation debris at the Bureau of
Land Management’s Jupiter Inlet Lighthouse Outstanding Natural Area (JILONA).
Cladonia perforata beds were vulnerable to a proposed prescribed fire treatment in areas with extensive hurricane debris. Ann DeBolt demonstrated successful translocation of
C. perforata, from proposed prescribed fire areas to areas where prescribed fire treatments would not occur for at least fifteen years (DeBolt 2009).
The extreme rarity of C. perforata accentuates the need to better understand its asexual fragment reproduction, and growth rate. Growth rate estimates are essential to estimate the time required for a community to recover from a perturbation. Also a morphological analysis will illustrate the variation in which thallus development patterns occur.
3 2: STUDY AREA
The Jupiter Ridge Natural Area is located in the southeast section of Jupiter,
Florida. That natural area covers 109 hectares (270 acres) of diverse vegetative communities including scrub, scrubby flatwoods, mesic flatwoods, and tidal swamps located within an urban and suburban area. About 64 hectares (159 acres) of the 109 hectare site is comprised of scrub habitat (PBC ERM 2010). The natural area is in the southern portion of the Atlantic Coastal Ridge, and contains large sand ridges. The sand ridges are remnants of ancient shorelines and are characterized by well drained, quartz- dominated, nutrient poor soils. Large open areas of sand have been maintained by fire in this pyric scrub community.
Prior to acquisition for conservation in 1991 and 1992, off road vehicles were used heavily on the site which also created and maintained open sandy areas. Cladonia perforata populations are found throughout the scrubby communities with smaller populations in oak scrub than rosemary scrub. Additionally, C. perforata aggregations can be found along the windblown edges of old closed off road vehicle trails. Steve
Farnsworth of Palm Beach County’s Department of Environmental Resources
Management reported a 1997 survey revealed approximately five thousand lichen fragments on the site (USFWS 1999). A more recent survey also concluded it likely for
4 there to be several thousand individuals (Richardson and Moore 2009) within the natural area.
Palm Beach County purchased the site with funds from the County’s
Environmentally Sensitive Land Bonds and matching funds from the state’s Conservation and Recreation Land (CARL) Program. The high quality natural communities found on the Natural Area are home to fifteen plant species and nineteen animal species that have been listed with some degree of endangerment by a government agency or nonprofit environmental organization (PBC ERM 2010). The natural area is open daily to the public. Some public use facilities that have been installed for passive recreation include a parking lot, accessibility trail, hiking trails, observation deck, and educational kiosk (PBC
ERM 2010). The natural area provides opportunities for the public to experience and learn about some of the natural environments found in Palm Beach County.
5 3: MORPHOLOGY
Methods and Materials
Fifty C. perforata specimens were collected from Jupiter Ridge Natural Area for the morphological analysis. Prior to handling, thalli were saturated with distilled water to decrease possible damage as they shrank and became brittle when desiccated (Pegau
1968). All of the thalli were cleaned of adhering sand and debris, and all of the necrotic tissues were removed from the thalli. The fifty thalli were photographed and sketched so internodes could be labeled and classified. In order to classify branch internodes, a branch hierarchy was developed based on Horton’s (1932) stream network branching system and a similar system used by Brazeau and Lasker (1988) in quantifying branching patterns in corals. Branches were defined by thalli elongation thus meristem bundles that have not elongated were not considered branches, but parts of their corresponding branch.
In this process, the most apical branches were defined as first order (1°) branches. When two 1° branches join, a second order (2°) branch was formed. Higher order branches only arose when branches of the same order connected. The number of branches in each order of branches were recorded (Table 3.1) and used to determine the average bifurcation ratio
(Equation 3.1). This is the ratio of one branch order to the next higher order (Brazeau and
Lasker 1988). Branches of the same order that joined to create a higher order branch were considered source branches. When a branch joined a higher order branch, it did not
6 increase the order of the system and was considered a tributary branch (Mock 1971). The number of tributary (Table 3.2) and source (Table 3.3) branches were recorded, and then the tributary to source ratio was calculated to provide another measure of branching analysis (Equation 3.2)
Each saturated thallus was then weighed with an Ohaus Explorer digital lab scale, and measured with dial calipers. The total length of the thallus was measured in addition to the width of its base (highest order branch). Thalli were photographed on one millimeter scaled sheet of graph paper to estimate the area they occupy (Table 3.4). Ten thalli were vacuum dried for one hour at 40 C and 93.125 kPa (27.5 in. Hg) (Table 3.5).
Ten sacrificial thalli were also dissected at each node. Branches were divided into branch orders, and then weighed to calculate their portion of the total thallus mass (Table 3.6).
Results
The fifty thalli collected consisted of sixteen 4°, thirty 5°, and four 6° branching networks. The bifurcation ratio for each thallus was calculated, and then averaged amongst thalli of the same branching network complexity (Figure 3.1). The average bifurcation ratio of each consecutive branch order decreased for 5°and 6° branch networks, while the ratio remained fairly constant for 4° branch networks. Furthermore,
1°, 2°, 3° branches are present in all three of the branching networks. A comparison showed the bifurcation ratio for these branches decreased when branch networks increased. Additionally, the tributary to source ratio was calculated and averaged amongst thalli of the same branching complexity (Figure 3.2). The results show that most
7 of the consecutive branches in a network have a lower tributary to source ratio than the previous branch. However, the tributary to source ratio for 3° branches was higher than
2° branches in the 4° and 6° branching networks. No correlation was observed between the complexity of branching networks and their tributary to source ratio.
After thalli were saturated with distilled water to ensure they were fully extended, their lengths, base diameters, and wet weights were recorded. A comparison of length to weight demonstrated a positive correlation between the two parameters, but with wide variations. One thallus may weigh twice as much as a thallus of similar length (Figure
3.3). A comparison of the thallus base diameter to wet weight (Figure 3.4) and base diameter to length (Figure 3.5) also showed a positive correlation with great variation among thalli.
The length of each thallus was measured, and then sorted by branch network complexity. Thalli of the same branch order were then averaged (Figure 3.6). The average thallus length for the 4° branching networks was 37.25 mm with a standard deviation of ± 5.84 mm. The average thallus length for the 5° branching networks was
39.98 mm with a standard deviation of ± 5.68 mm. The average thallus length for the 6° branching networks was 54.75 mm with a standard deviation of ± 12.75 mm.
Additionally, the saturated weight of each thallus was recorded, then sorted by branch network complexity and averaged (Figure 3.7). The average saturated weight for the 4° branching networks was 0.72 g with a standard deviation of ± 0.29 g. The average saturated weight for the 5° branching networks was 1.01 g with a standard deviation of ±
8 0.29 g. The average saturated weight for the 6° branching networks was 1.64 g with a standard deviation of ± 0.36 g.
Ten thalli were vacuum dried for one hour at 40 C and 93.125 kPa (27.5 in. Hg) then weighed. Their wet weight was compared to their dry weight (Figure 3.8). The average dry weight of a thallus was 23.85% of its wet weight. The wet to dry weight was very consistent with only a ±0.02 g standard deviation between the actual dry weight and the average wet weight to dry weight ratio.
To determine the distribution of weight throughout a thallus, ten 5° thalli were dissected at each node. The percent of mass distributed in each branch order was calculated by dividing the weight of all branches in that order by the total weight of the thallus. The results of all ten thalli were then averaged (Figure 3.9). The 1ᵒ branches were the most abundant branches and accounted for 28.98% of the total weight. The 2ᵒ and 3ᵒ branches comprised 24.02% and 20.67% of the total weight respectively. Additionally, the 4ᵒ branches contained 16.49% of the total weight. Lastly, the 5ᵒ branches contributed the least amount to the total weight at 9.82%.
The photographs of the thalli placed on one millimeter scaled sheet of graph paper were printed, and then the graph lines were transposed across each thallus. Cells that were more than 50% covered by the thallus were then counted to determine the surface area of each thallus. There was a positive correlation between surface area and weight
(Figure 3.10). The standard deviation for the actual thallus area versus the slope intercept of the graph is ± 83.39 cm2.
9 Discussion
A branching network with perfect dichotomous branching has a bifurcation ratio of two. This value increases as branches which do not increase the order of the system are added (Brazeau and Lasker 1988). The results indicate that as thalli grow and branching networks increase, branch networks become more bifurcated. A tributary to source ratio of one means the tributary and source branches are equal in quantity. A tributary to source ratio above one indicates there are more tributary branches, and below one indicates there are more source branches. As branch order increased (branches are older) the number of source branches increased and the number of tributary branches decreased.
If the meristem bundle could develop from within internodes, the tributary ratio among higher order branches would be much higher as new younger branches would not increase the order of the system. Therefore, new branch development is limited to the apical meristems, and tributary branches decline as branches age.
While some other Cladonia species such as Cladonia alpestris have well defined forms that develop regularly (Karenlampi 1970), C. perforata has high variability in its form. As was expected there is a positive correlation between thallus length, base diameter, and wet weight. There are significant overlapping ranges amongst thalli of different complexity in their weights, lengths, and base diameters. Thus, there would be little confidence in any prediction of length, wet weight, branch orders, or base diameter if calculated from one of these known parameters. The surface area to weight comparison also had a positive correlation, but had a fair degree of variability. Whereas the dry weight to wet weight ratio was seen to be fairly consistent. To determine the distribution 10 of weight across a thallus, branches of the same order were weighed together and divided by the total thallus weight. While the lower order branches (younger branches) are smaller than higher order branches, the lower order branches are more abundant.
Therefore it is logical that the weight distribution showed the lower order branches provided more to the total thallus weight than the higher order branches. These results, suggest an inverse relationship between branch orders and their contribution to the total thallus weight.
11 4: GROWTH RATE
Methods and Materials
One hundred C. perforata thalli were collected from the Jupiter Ridge Natural
Area. The thalli were gathered from multiple locations as not to significantly reduce the population in any individual lichen bed within the site. Exposure to heat and dry air will quickly desiccate thalli causing them to shrink and become brittle (Pegau 1968).
Subsequently, thalli were always saturated with distilled water prior to handling to abate unintended damage. Once collected, all necrotic tissues were removed from each thallus so the decay of dead tissue would not deduct from the overall growth of the thallus.
After the one hundred thalli had their necrotic tissues removed, they were then split in to two groups of fifty. One group was used to measure total growth rates, while the other fifty were used to measure sympodium branch growth rates. To measure total growth rates, fifty thalli were saturated by spraying them with distilled water, which also aided in removing sand and other debris. Baseline data was collected in August of 2011.
Total lengths from their base to most apical tip were recorded with the use of a dial caliper and a millimeter ruler (Table 4.1). Also while still saturated and pliable, the thalli were weighed using an Ohaus Explorer digital lab scale (Table 4.2). To insure thalli could be correctly identified and aid in documenting branching events, each thallus was then photographed on one millimeter scaled sheet of graph paper. These photographs 12 were also marked to ensure that the same branch was measured over the course of the experiment. Thalli were then tethered with nylon string to a four inch plastic zip tie. Each tether had a unique color combination of size ten and eleven Czech seed beads to assist with identifying each thallus in the field. The fifty thalli were then returned to ten monitoring stations that were created within the Jupiter Ridge Natural Area. Monitoring stations were located in small sandy openings. Stations were located near existing C. perforata populations, to ensure they were in suitable habitat. At each monitoring station, the zip ties of five thalli were inserted into the sand to act as anchors tethering each thallus to the ground (Figure 4.1). Stations were located with a hand held Garmen Etrex
Legend GPS. As this GPS didn’t have sub-meter accuracy, to delineate the stations location while remaining inconspicuous a small pile of branches were stacked near each station. The fifty thalli were then collected from the field quarterly (every thirteen weeks) and photographed, weighed and measured for two years. Thalli were photographed with a
Fugi Fine Pix Z camera. The thalli were placed on a one millimeter scaled sheet of graph paper and the camera was set to micro. During each monitoring event, thalli were removed from their tether and the data collection followed the protocols established in the base line data collection. All of the data was collected and the thalli were returned to the field within one week of their removal.
The remaining fifty thalli were used to track intercalary growth by branch order.
The branch hierarchy classification followed the Horton (1932) stream network branching system and a similar system used by Brazeau and Lasker in 1988 to quantify branching patterns in corals. All thalli were saturated with distilled water prior to
13 handling. Each thallus was drawn and branches were labeled with their corresponding branch orders. The sympodium was established as the axis with the highest number of contiguous branch orders. The nodes of the sympodium were marked with a dab of silicone applied to the thallus with a needle point. These control points ensured that all subsequent measurements would occur at the same locations. No silicone was placed at the apical tip and measurement started from the first node. The base line data was collected in October of 2011, and consisted of the total length and length of the marked branches on each thallus (Tables 4.3 – 4.5). Following the protocols established with the first group of fifty, thalli were tethered to zip ties marked with colored seed beads and returned to an additional ten monitoring stations located within the natural area. Thalli were then collected from the field quarterly (every thirteen weeks) and measured for two years. All the data was collected and the thalli were returned to the field within one week of their removal.
Results
As a result of placing the thalli in a public and natural setting, some thalli were disturbed by wildlife, weather, or people. Consequently, roughly one third of the thalli from the total growth subset were not retrieved for measurements over the duration of the two years period. Unlike dicot plants whose linear growth is limited to their apical meristem, the entire thallus of a lichen can extend due to intercalary growth causing linear expansion throughout the thalli. For this reason growth rates were calculated as a percent increase in length. After two years, the average growth was 10.42% per year. The range of the thalli which were measured for the full two years varied in size from 14 eighteen to thirty one millimeters at the time zero recording. The range of the thalli at the end of the two-year period were twenty to thirty eight millimeters. Length was recorded on a quarterly basis and growth was calculated as a percent linear increase from the previous quarter (Equation 4.1). The percent linear growth was then averaged for all the thalli. While the major growth events didn’t match between each year, a temporal pattern emerged showing the lowest growth for both years in the March to May quarter (Figure
4.2). During this time weight changes were also recorded. The average weight change for the two year period was a 79.33% increase. Furthermore, weight change was recorded on a quarterly basis and change was calculated as a percent weight increase from the previous quarter (Equation 4.2). The percent weight change was then averaged for all of the thalli. Once again, the major growth events didn’t match from year to year, but for both years the March to May quarter had the lowest growth (Figure 4.3). Additionally, linear growth and weight gain correlated well for both years. In the first year the largest percent linear growth and weight gain occurred in the quarter ending in November followed by the August, February, and May quarters. The second year the largest percent linear growth and weight gain occurred in the quarter ending in February, followed by the
November, August, and May quarters. Branch division was not observed between August
2011 and August 2013 (Figure 4.4). Additionally, linear growth didn’t occur evenly across the thalli. Often one side of a thallus would grow at a quicker rate than the other
(Figure 4.5).
15
The remaining fifty thalli were used to track intercalary growth of branch orders.
The total thalli lengths ranged from seventeen to forty one millimeters at time zero. As with the previous experiment some thalli were lost over the course of the two year period.
All fifty thalli had a branch network of four branch orders. Length was recorded on a quarterly basis and growth was calculated as a percent linear increase from the previous quarter’s branch length throughout the year. The younger, lower order branches grew by a higher percentage rate in all quarterly measurements except July 2012 (Figure 4.6). The average annual growth rate for second order branches was 52.34%. The third order branches had 15.9%, and forth order branches were 7.76% annual growth.
Discussion
Cladonia perforata’s low annual growth rate is expected considering the harsh environment in which it lives. With no waxy cuticle to protect the thallus, lichens can be quickly desiccated in inhospitable environments; but this also allows the thalli to absorb moisture from precipitation, dew, and fog. Due to the harsh nature of the scrub environment and the lack of precipitation during the dry season, morning dew may be a significant source of water for this species (Richardson and Moore 2009). As water is essential for photosynthesis and gas exchange, it is likely that C. perforata may only photosynthesize for a few hours a day, thus explaining its slow growth rate.
Weight gain seemed to correlate well with linear growth. Weight gain in the last quarter may have been skewed, due to fragmentation and damage to thalli. During this quarter, several thalli were observed to have lost large branches (Figure 4.7). The loss of
16 these branches would have offset any weight gained during that quarter without impacting linear growth.
It was observed that a complete cycle of branch division was not evident while reviewing photos from the two year time period. Early in the branch splitting process, the meristem will divide into two bundles separated by a central perforation (Hammer 2000).
Elongation in these meristem bundles and a widening of the perforations where observed in all of the thalli, but not in all meristems. Most of the meristems that were closer to the base showed no development. Several thalli started the experiment with elongated meristems in which new perforations were observed indicating division had begun. These meristem bundles didn’t elongate, hence returning to the stage at which this cycle started.
This suggests that it may take more than two years for some thalli to complete the branch division process. Additionally, it was observed that thallus branches grew at independent rates. This may be a continuation of the independent growth rates observed in early branch division. Meristem bundles at roughly the same height can be in different developmental stages, causing different vertical growth in different parts of the thallus
(Hammer 2000).
The intercalary growth of branch orders was tracked for two years. Cladonia lichens are known to have three life stages, the growth accumulation period, renewal period, and degeneration period (Pegau 1968). The results clearly showed a progression from high growth rates to low growth rates as branches aged. The three periods can also be seen when examining an individual C. perforata thallus. While many of the thalli continually showed growth, some also showed degeneration at the bases of their oldest 17 branch (Figure 4.8). When the base decay reaches a branching point, the thallus will fragment. This fragmentation allows for dispersal of the thallus, and may reduce competition amongst thalli that would have otherwise over shaded themselves.
18
5: FRAGMENTATION
Methods and Materials
All fragment source materials were collected from the Jupiter Ridge Natural Area.
The thalli were gathered from multiple locations within three management units, as not to significantly reduce the population in any individual lichen bed within the site. Exposure to the sun quickly desiccates thalli causing them to shrink and become brittle (Pegau
1968). Subsequently, thalli were always saturated with distilled water prior to handling to increase pliability, ensure they are fully extended, and to abate unintended damage to the thalli.
After saturating the thalli with distilled water, 100 two-millimeter fragments were generated by cutting one apical branch from 100 thalli. The process was then repeated to create 100 four-millimeter, six-millimeter, eight-millimeter, and ten-millimeter fragments respectively. Each individual thallus fragment was assigned a unique number. For identification purposes and to prevent the loss of fragments, individual fragments were placed in cylinders similar to those used by Hooker (1980) and Lindsay (1975). The cylinders were made of three inch long transparent plastic tubing with a two inch diameter. A screen with one-millimeter mesh openings was attached to the bottom of each cylinder to allow precipitation to drain from the cylinder, while retaining the thalli fragments (Figure 5.1). The cylinders were partially buried in sand, and then filled to an 19 equal level with sand to help stabilize the cylinder and allow the thalli to rest on a natural medium. After length and weight measurements were collected the thalli were placed in cylinders and labeled with their corresponding identification numbers. A forty eight square foot artificial scrub habitat was created by lining a twelve foot by four foot planter box with a weed barrier. The planter box was then filled with four inches of sand (Figure
5.2). The artificial scrub was created to store the cylinders on private property in order to prevent vandalism of the project. Additionally, weeds were removed from the artificial habitat on a quarterly basis. The original five hundred parent thalli were returned to their source lichen beds within the Natural Area.
Thalli fragment mortality, lengths, and weights were recorded quarterly (every thirteen weeks) for one year. Fragments retaining a recognizable form were recorded as viable. Thalli that have completely decayed or have degenerated to a loose ball of filaments were recorded as deceased. Fragments were saturated with distilled water to ensure they were fully extended prior to recording their length with the use of dial calipers and a millimeter scaled ruler. While still saturated and pliable, thalli weights were also recorded using an Ohaus Explorer digital lab scale (Tables 5.1-5.5). Due to the nature of classifying the fragments by size, changes in weight were calculated as a percent change instead of comparing total weight changes. Fragment weight changes were compared to their weight in the previous quarter then averaged for each class size.
The deceased fragments were recorded as -100% changes in length and weight, but were not used to calculate the average change in the quarter in which they died. Finally, a Chi- square test was used to determine if there was a statistically significant difference in
20 survivability among the fragment size classes. The p (probability value) value for the
Chi-square tests was .05 and the df (degrees of freedom) value was 1.
Results
The thalli fragment survivability was calculated based on size class. Survival generally increased with fragment size (Figure 5.3). After one year, the two-, four-, six-, eight-, and ten-millimeter fragments had an 8 %, 25 %, 24 %, 79 %, and 74% survival rate respectively. While mortality rates per quarter (Figure 5.4) varied for size classes throughout the year, typically larger fragments fared better and appeared more stable than small fragments. Using a p value of .05, the Chi-square test showed no statistically significant difference between the two-, four-, and six-millimeter fragments survivability.
Additionally, there was no statistically significant difference between the eight-millimeter and the ten-millimeter survivability. However, there was a significant difference between these two groups (Table 5.6).
The initial quarter showed a positive average weight increase for all thalli class sizes. The three subsequent quarters had a negative average weight change for most fragment class sizes, excluding the ten-millimeter fragments in the second quarter, two- millimeter fragments in the third quarter, and four-millimeter fragments in the fourth quarter which had a positive change. Over the course of one year, the average length of the fragment classes changed little. The ten-, six-, and four-millimeter fragments had a small increase in their average length. The two-millimeter fragments had no growth while the average length for the eight-millimeter fragments decreased (Figure 5.5). Despite the
21 small changes in the average lengths, individual fragments ranged in size from one to twelve millimeters in length. The two- and eight-millimeter class sizes each had one fragment that increased in length by 25 % to 2.5 and 10 millimeters respectively. The four- and six-millimeter class sizes each had a fragment which increased by 50 % to 6 and 9 millimeters. Additionally, one ten-millimeter fragment grew by 20 % to 12 millimeters.
Discussion
As vagrant lichens, C. perforata doesn’t attach to the substrate. Instead it may be blown around freely like a tumble weed. While other lichens colonize isolated habitat patches with spores or specialized long distance dispersal units, C. perforata predominately reproduces by vegetative fragments (Yahr, 2000). The persistence and dispersal of these fragments, is key to recovering from perturbations and expanding the species’ range. Fragment survival generally had a positive correlation with size, but the survivability did not increase incrementally with size. Instead, the four- and six- millimeter fragment survivability plateaued between twenty-four and twenty-five percent.
An additional plateau occurred between the eight- and ten-millimeter fragments at seventy-four and seventy-nine percent.
These plateaus are represented with similar slopes when observing mortality per quarter (Figure 5.4), and are replicated again in the average mortality rate (Figure 5.6).
Average percent weight change and average fragment length change had no correlation with mortality rates or fragment size. While not in the scope of the protocols established
22 for this research, changes in fragment coloration were observed in many thalli months prior to their deaths. Loss of the photo-symbiont may have been a precursor or a cause for at least a portion of the thalli mortality observed. The two-, four-, and six-millimeter fragments, had low survivability. The Chi-square test showed no statistically significant difference between these fragments with low persistence. The eight- and ten-millimeter fragments had a high survivability, with no statistical difference between the two fragment sizes. The statistically significant difference recorded between the low survival and high survival fragment groups, illustrates that large fragments have a higher survivability than smaller fragments, and are better suited for asexual fragment reproduction. Additionally, the results suggest there is a key transitional class size in which survivability increases significantly. These findings may also explain why attempts by others to create lichen fragment powder by grinding thalli in a food mill were unsuccessful at colonizing in suitable lichen habitat.
23
APPENDIX
Morphology
(Rb=Ni/Ni+1)
Equation 3.1 Bifurcation Ratio
N denotes any given branch order.
Ti/Si
Equation 3.2 Tributary to Source Ratio
T denotes a tributary which occurs when a branch joins a higher order branch. This does
not increase the order of the system. S denotes a source branch. Source branches are of
the same order and join to create a higher order branch.
24
Average Bifurcation Ratio
4.25
4.5 4.15
4.13
3.75
4
3.36 3.34
3.5 3.28
3.00 2.93
Rb 1ᵒ
3
2.50 2.27 2.5 2.25 Rb 2ᵒ 2 Rb 3ᵒ 1.5 Rb 4ᵒ 1 Rb 5ᵒ Average Average BifurcaionRatio 0.5 0 4° Branch Network 5° Branch Network 6° Branch Network
Figure 3.1 Bifurcation Ratio
As thalli branching networks increase, they become more bifurcated.
Average Tributary to Source Ratio
1
0.87
0.8
0.70
0.68
0.67
0.64 T/S1ᵒ
0.6 0.52 0.50
0.50 T/S2ᵒ 0.46
T/S3ᵒ
0.4
0.25 T/S4ᵒ 0.13 0.2 0.13 T/S5ᵒ
0 Average Average TributarySource to Ratio 4° Branch Network 5° Branch Network 6° Branch Network
Figure 3.2 Average Tributary to Source Ratio
As branch order increases, the tributary to source ration decreases. Thus source branches
increase and tributary branches decrease.
25
Length to Wet Weight Slope Intercept: y = 10.568x + 29.335 60 50 40 30 20 10
Thallus Thallus Lengthin mm 0 0 0.5 1 1.5 2 2.5 Saturated Thallus Weight in g
Figure 3.3 Length to Wet Weight
A comparison of thalli weigh to thalli length shows high variability in the form of thalli.
Thalli of relatively similar lengths have a wide range of weights.
Base Diameter to Wet Weight Slope Intercept: y = 0.587x + 3.4326
8
6
4
2
0 0 0.5 1 1.5 2 2.5 Saturated Thallus Weight in g
Diameter Diameter Thallus of mm Base in Figure 1.4 Base Diameter to Wet Weight
The thallus wet weight has a positive correlation to the base diameter of a thallus. High
variability was indicated as thalli with the same base diameter had a wide range of
weights.
26
Length to Diameter Slope Intercept: y = 0.0409x + 2.3509
8 6 4 2 0 20 30 40 50 60 70 80
Length of Thallus in mm Diameter Diameter Thallus of mm in
Figure 3.5 Length to Diameter
A comparison of thalli base diameter to thalli length shows high variability in the thalli
form. Thalli of relatively similar base diameters have a wide range of lengths.
Figure 3.6 Average Thallus Length by Branching Network
Thalli length had a positive correlation with branch network order. The average lengths for 4° and 5° networks were within the standard variation of each other. While the 5° and
6° order networks only overlapped by 2.66 mm (between 42.0 mm and 44.66 mm).
27
Figure 3.7 Average Thallus Wet Weight by Branching Network
Thalli wet weight had a positive correlation with branch network order. The standard
variation between the 4° and 5° order thalli had a large amount of overlap while the
standard variation of 5° and 6° order networks had only a slight overlap.
Dry Weight to Wet Weight
2
1.643
1.5 1.2729
1.2344
1.1481
1.088
1.0158
0.9855
Wet weight
0.8015
1
0.7654
0.5584 Dry Weight
0.3463 0.3386
0.5 0.3214
0.2624
0.255
0.2543
0.2299
0.1758
0.1722
0.1425 Thallus Thallus weight g in 0 1 2 3 4 5 6 7 8 9 10
Figure 3.8 Dry Weight to Wet Weight
The average dry weight of a thallus is 23.85% of its saturated weight. The wet to dry weight was very consistent with only a ±0.02 g standard deviation between the actual dry
weight and the average wet weight to dry weight ratio.
28
Branch Composition of Total Thallus
9.82%
28.98% 1ᵒ Branchs 16.49% 2ᵒ Branchs 3ᵒ Branchs 4ᵒ Branchs
20.67% 5ᵒ Branches 24.04%
Figure 3.9 Branch Composition of Total Thallus
The smallest branch orders are the most abundant and contribute the most to the total
thallus weight.
Weight to Surface Area Slope Intercept: y = 523.99x + 85.157 1800 1600 1400 1200 1000 800 600
Area in square mm 400 200 0 0 0.5 1 1.5 2 2.5 Hydrated thallus weight in g
Figure 3.10 Weight to Surface Area
There is a positive relationship between thallus weight and surface area. 29
Table 3.1 Number of Branches in Each Branch Order
Number of Branches in Each Branch Order Thallus Thallus 1ᵒ 2ᵒ 3ᵒ 4ᵒ 5ᵒ 6ᵒ 1ᵒ 2ᵒ 3ᵒ 4ᵒ 5ᵒ 6ᵒ ID ID 1 234 43 10 4 1 0 26 61 17 6 2 1 0 2 71 22 6 1 0 0 27 67 22 5 1 0 0 3 80 23 5 2 1 0 28 110 35 12 5 2 1 4 66 10 5 2 1 0 29 130 34 11 2 1 0 5 47 12 2 1 0 0 30 93 35 10 2 1 0 6 73 14 3 1 0 0 31 58 12 2 1 0 0 7 93 23 5 2 1 0 32 100 27 7 2 1 0 8 80 21 5 1 0 0 33 92 24 7 1 0 0 9 45 10 3 1 0 0 34 67 18 7 2 1 0 10 62 19 6 2 1 0 35 113 28 9 2 1 0 11 69 22 8 2 1 0 36 100 27 7 2 1 0 12 80 18 6 1 0 0 37 116 33 11 3 1 0 13 135 38 9 1 0 0 38 123 32 8 3 1 0 14 55 13 3 1 0 0 39 35 6 2 1 0 0 15 63 20 6 2 1 0 40 69 18 5 2 1 0 16 54 16 5 2 1 0 41 58 16 5 2 1 0 17 66 19 5 2 1 0 42 125 37 11 3 1 0 18 80 22 6 2 1 0 43 56 17 5 2 1 0 19 69 21 7 2 1 0 44 64 16 6 2 1 0 20 75 24 6 2 1 0 45 85 23 4 1 0 0 21 39 10 3 1 0 0 46 117 31 14 4 2 1 22 43 12 4 1 0 0 47 157 39 10 4 1 0 23 71 15 4 1 0 0 48 167 45 15 3 1 0 24 67 18 6 2 1 0 49 243 75 26 9 3 1 25 93 22 9 2 1 0 50 189 59 16 5 2 1
The number of branches in each branch order was recorded for each thallus
30
Table 3.2 Number of Tributary Branches in Each Branch Order. When a branch joined a higher-order branch, it did not increase the order of the system and was
considered a tributary branch.
Number of Tributary Branches in Each Branch Order Thallus Thallus 1ᵒ 2ᵒ 3ᵒ 4ᵒ 5ᵒ 6ᵒ 1ᵒ 2ᵒ 3ᵒ 4ᵒ 5ᵒ 6ᵒ ID ID 1 148 23 2 2 1 0 26 27 5 2 0 1 0 2 27 10 4 1 0 0 27 23 12 3 1 0 0 3 34 13 1 0 1 0 28 40 11 2 1 0 1 4 46 0 1 0 1 0 29 62 12 7 0 1 0 5 23 8 0 1 0 0 30 23 15 6 0 1 0 6 45 8 1 1 0 0 31 34 8 0 1 0 0 7 47 13 1 0 1 0 32 46 13 3 0 1 0 8 38 11 3 1 0 0 33 44 10 5 1 0 0 9 25 4 1 1 0 0 34 31 4 3 0 1 0 10 24 7 2 0 1 0 35 57 10 5 0 1 0 11 25 6 4 0 1 0 36 46 13 3 0 1 0 12 44 6 4 1 0 0 37 50 11 5 1 1 0 13 59 20 7 1 0 0 38 59 16 2 1 1 0 14 29 7 1 1 0 0 39 23 2 0 1 0 0 15 23 8 2 0 1 0 40 33 8 1 0 1 0 16 22 6 1 0 1 0 41 26 6 1 0 1 0 17 28 9 1 0 1 0 42 51 15 5 1 1 0 18 36 10 2 0 1 0 43 22 7 1 0 1 0 19 27 7 3 0 1 0 44 32 4 2 0 1 0 20 27 12 2 0 1 0 45 39 15 2 1 0 0 21 19 4 1 1 0 0 46 55 3 6 0 0 1 22 19 4 2 1 0 0 47 79 19 2 2 1 0 23 41 7 2 1 0 0 48 77 15 9 1 1 0 24 31 6 2 0 1 0 49 93 23 8 3 1 1 25 49 4 5 0 1 0 50 71 27 6 1 0 1
31
Table 3.3 Number of Source Branches in Each Branch Order. Branches of the same order that joined to create a higher-order branch were considered source branches.
Number of Source Branches in Each Branch Order Thallus Thallus 1ᵒ 2ᵒ 3ᵒ 4ᵒ 5ᵒ 6ᵒ 1ᵒ 2ᵒ 3ᵒ 4ᵒ 5ᵒ 6ᵒ ID ID 1 86 20 8 2 0 0 26 34 12 4 2 0 0 2 44 12 2 0 0 0 27 44 10 2 0 0 0 3 46 10 4 2 0 0 28 70 24 10 4 2 0 4 20 10 4 2 0 0 29 68 22 4 2 0 0 5 24 4 2 0 0 0 30 70 20 4 2 0 0 6 28 6 2 0 0 0 31 24 4 2 0 0 0 7 46 10 4 2 0 0 32 54 14 4 2 0 0 8 42 10 2 0 0 0 33 48 14 2 0 0 0 9 20 6 2 0 0 0 34 36 14 4 2 0 0 10 38 12 4 2 0 0 35 56 18 4 2 0 0 11 44 16 4 2 0 0 36 54 14 4 2 0 0 12 36 12 2 0 0 0 37 66 22 6 2 0 0 13 76 18 2 0 0 0 38 64 16 6 2 0 0 14 26 6 2 0 0 0 39 12 4 2 0 0 0 15 40 12 4 2 0 0 40 36 10 4 2 0 0 16 32 10 4 2 0 0 41 32 10 4 2 0 0 17 38 10 4 2 0 0 42 74 22 6 2 0 0 18 44 12 4 2 0 0 43 34 10 4 2 0 0 19 42 14 4 2 0 0 44 32 12 4 2 0 0 20 48 12 4 2 0 0 45 46 8 2 0 0 0 21 20 6 2 0 0 0 46 62 28 8 4 2 0 22 24 8 2 0 0 0 47 78 20 8 2 0 0 23 30 8 2 0 0 0 48 90 30 6 2 0 0 24 36 12 4 2 0 0 49 150 52 18 6 2 0 25 44 18 4 2 0 0 50 118 32 10 4 2 0
32
Table 3.4 Wet Weight, Length, Base Diameter, Branch Orders, and Area (Thalli 1-
25).
Wet Weight, Length, Base Diameter, Number of Branch Orders, and Area of Each Thallus Number Wet Thallus Length Diameter of Area sq weight ID mm mm branch mm g order 1 1.1774 43 4 5 856 2 0.7859 27 3 4 341 3 1.2083 33 5 5 598 4 0.5596 33 4 5 535 5 0.363 31 4 4 220 6 0.866 36 4 4 499 7 1.0273 42 6 5 647 8 1.4085 42 4.5 4 617 9 0.4631 45 5 4 335 10 0.774 35 4 5 489 11 0.9072 48 4 5 463 12 0.9998 39 3 4 454 13 1.2517 40 3 4 797 14 0.7948 31 3 4 540 15 0.9087 26.5 3 5 416 16 0.75 35 5 5 460 17 0.8784 36 3 5 497 18 0.4449 29 4 5 318 19 0.4659 28 3.5 5 347 20 0.9032 38 3 5 526 21 0.2695 26 3 4 216 22 0.2372 26 3 4 233 23 0.4867 40 6 4 310 24 0.977 41 4.5 5 455 25 1.693 48 5 5 651
33
Table 3.4 (Continued) Wet Weight, Length, Base Diameter, Branch Orders, and
Area (Thalli 26-50).
Wet Weight, Length, Base Diameter, Number of Branch Orders, and Area of Each Thallus Number Wet Thallus Length Diameter of Area sq weight ID mm mm branch mm g order 26 1.0158 49 5 5 518 27 0.7654 42 4 4 547 28 1.2344 45 4 6 749 29 1.643 50 5 5 953 30 0.8015 43 4 5 553 31 0.7187 41 3 4 417 32 1.1445 50 5 5 844 33 0.5584 41 3.5 4 461 34 0.4489 39 4 5 352 35 1.088 38 4 5 648 36 1.1481 44 5 5 683 37 1.2729 38 5 5 748 38 0.9855 31 4 5 598 39 0.2919 37 3 4 206 40 0.7093 39 4 5 475 41 0.8456 40 4 5 547 42 1.7572 54 4 5 1024 43 0.6495 36 3 5 766 44 0.7774 40 3 5 387 45 1.2335 52 4 4 708 46 1.3197 39 3 6 871 47 1.6286 44 3 5 1551 48 1.6869 49 4 5 789 49 2.2226 42 5 6 1257 50 1.7826 56 5 6 1111
34
Table 3.5 Dry Weight to Wet Weight Ratio
Dry Weight to Wet Weight Ratio
Thallus Wet Dry Wet/Dry ID weight weight
26 1.0158 0.2543 25.03% 27 0.7654 0.1722 22.50% 28 1.2344 0.3463 28.05% 29 1.643 0.3386 20.61% 30 0.8015 0.1758 21.93% 33 0.5584 0.1425 25.52% 35 1.088 0.255 23.44% 36 1.1481 0.2624 22.86% 37 1.2729 0.3214 25.25% 38 0.9855 0.2299 23.33%
Ten thalli were vacuum dried for one hour at 40 C and 27.5 in. Hg, and then weighed to
calculate the wet weight to dry weight ratio.
35
Table 3.6 Weight of Branch Orders and Total Thallus Weight. Ten sacrificial thalli
were dissected at each node. Branches were divided into branch orders, and then
weighed to calculate their portion of the total thallus weight.
Weight of Branch Orders and Total Weight 1ᵒ 2ᵒ 3ᵒ 4ᵒ 5ᵒ Total Thallus Branches Branches Branches Branches Branches thallus ID in g in g in g in g in g in g 3 0.1689 0.1357 0.0994 0.0383 0.0113 0.4536 4 0.038 0.033 0.0436 0.0158 0.0135 0.1439 10 0.025 0.0409 0.0432 0.041 0.0566 0.2067 11 0.0629 0.0441 0.0411 0.0223 0.0399 0.2103 15 0.0857 0.0476 0.0566 0.0552 0.0229 0.268 16 0.0615 0.0474 0.0492 0.0294 0.018 0.2055 17 0.1428 0.1427 0.1176 0.0775 0.0244 0.505 24 0.09 0.0859 0.0778 0.0732 0.0401 0.367 34 0.058 0.0402 0.0299 0.053 0.0469 0.228 40 0.122 0.0916 0.0513 0.0805 0.0159 0.3613
36
Growth Rate
(Li+1 –Li)/Li
Equation 4.1Percent Change in Length
Li = Initial length
Li+1 = One measuring event after initial length.
(Wti+1 –Wti)/Wti
Equation 4.2 Percent Change in Weight
Wti = Initial weight
Wti+1 = One measuring event after initial weight.
37
Figure 4.1 Tethered Thallus
To prevent the loss of thalli, they were tethered to a zip tie. A unique color sequence of
Czech seed beads were used for identification purposes.
Average Linear Growth
6.00% 5.07% 5.00% 4.47% 4.00% 3.00% 2.31% 1.75% 1.88% 2.00% 1.66% 1.45% 1.00% 0.53% 0.00% NOV 11 FEB 12 MAY 12 AUG 12 NOV 12 FEB 13 MAY 13 AUG 13 Week 13 Week 26 Week 39 Week 52 Week 65 Week 78 Week 91 Week 104
Figure 4.2 Average Linear Growth
Average linear growth varied across the two year period, but the March through May
quarter had the lowest growth rates for both years.
38
Average Weight Change 45.00% 42.59%
40.00%
35.00%
30.00%
25.00%
20.00% 16.68% 14.04% 15.00% 12.23% 9.50% 10.00% 7.85%
5.00% 2.15% 1.02% 0.00% Nov 2011 Feb 2012 May 2012 Aug 2012 Nov 2012 Feb 2013 May 2013 Aug 2013 Week 13 Week 39 Week 39 Week 52 Week 65 Week 78 Week 91 Week 104
Figure 4.3 Average Weight Change
Average weight change varied across the two year period, but the March through May quarter had the lowest growth rates for both years. Additionally weight change tracked
linear growth well, with the exception of the week 104.
39
5 mm 5 mm
Figure 4.4 Thallus With No Branch Division After Two Years
A portion of the thallus has been accentuated, to illustrate how meristems have extended
over the course of the experiment, but no additional branch division has occurred.
5 mm 5mm
Figure 4.5 Branches Grew at Independent Rate
Linear growth didn’t occur evenly across the thalli. As seen here, often one side of a
thallus would grow at a quicker rate than the other.
40
Percent Growth Rate By Branch Order 2nd order 25.00%
21.57% branch
20.00% 3rd order
branch
14.02%
15.00%
4th
orderbranch 7.92%
10.00% 7.52%
6.15%
5.98%
5.62%
5.53%
5.09%
5.03%
4.88%
4.16%
3.81%
3.71%
3.41% 3.08%
5.00% 2.98%
1.87%
1.85%
1.59%
1.57%
1.21%
0.35% 0.10% 0.00% JAN 2012 APR JUL 2012 OCT JAN 2013 APR JUL 2013 OCT Week 13 2012 Week 39 2012 Week 65 2013 Week 91 2013 Week 26 Week 52 Week 78 Week 104
Figure 4.6 Percent Growth Rate By Branch Order
The younger, lower order branches grew by a higher percentage rate in all quarterly
measurements except for in July of 2012.
5mm 5mm
Figure 4.7 Thallus With Fragmented Branch Thallus that lost branches, may have contributed to the disparities between the linear
growth and weight gained recorded in the last quarter of the experiment.
41
5 mm
Figure 4.8 Thallus With Degenerating Base
Some thalli showed degeneration at the bases of their base (oldest branch)
42
Table 4.1 Length (Thalli 01-25). The total length of each thallus from its base to its
most apical tip, was recorded in millimeters. (This table continues on to the next
page.)
Thallus Length in Millimeters AUG NOV FEB MAY AUG NOV FEB MAY AUG Thallus 2011 2011 2012 2012 2012 2012 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 1 21 21.5 22 22 22 22 22.5 21 22 2 22 22 23 23 23 22 23 23 24 3 20 21 22 22 22 4 25.5 27 27.5 28 28 30.5 30.5 31 32 5 24 26 26.5 27.5 27.5 26.5 27 6 24 26 27 27 27 28 28 29 30 7 22 23 25 25 26 26.5 25 26 26 8 22 23 24 25 26 27.5 27.5 28.5 29 9 27 27 27 27 28 27 28 27 29 10 19 19 19 20 20 21.5 23 23 23.5 11 22 22 22.5 22.5 22.5 22.5 24 24.5 24.5 12 20 21 22 22 23 24 24 24 25 13 17 18 18 18 18 18.5 19 19 20 14 22 23 23 23 23.5 23 25 24 26 15 22 23 23 23 23.5 24.5 24 26 27 16 23 24 24.5 24.5 25.5 26 27 26 17 20 20 20 21 21 22.5 23 23 18 19 20 21 21.5 22 22.5 24 24 19 25 25.5 25.5 26 27 27 29 30 22 20 23 23 23 23 24 24.5 27 23 24 21 22 23 23 24 25 25.5 25.5 27 27 22 26 28 28 28 23 21 22 22 24 23 24 24 24 24 22 23 23 23.5 25 22 24 24 25 26 27.5 28 29 30
43
Table 4.1 Continued Length (Thalli 26-50). The total length of each thallus from its
base to its most apical tip, was recorded in millimeters.
Thallus Length in Millimeters AUG NOV FEB MAY AUG NOV FEB MAY AUG Thallus 2011 2011 2012 2012 2012 2012 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 26 19 27 17 18 19 19 20 20.5 21 22 22 28 23 25 26 26 26 26 29 30 30 29 20 30 19.5 20 20 21 21 23 23 23 24 31 23 25 25 25 25 25 26 27 28 32 21 23 23 23.5 24 22.5 24 33 22 23 23 23 23 23 25 24 25 34 19 20.5 20.5 20.5 22 23 23 23.5 35 22 23 23.5 24 36 24 26 26 27 27 26.5 29 29 29 37 17 19 19 19 20 20 21 21.5 23 38 20 21 21 22 23 23 26 23 24 39 19.5 20.5 21 21 22 22 24 40 16 16 16 16 16.5 17 18 41 24 25.5 26.5 27 28 29 30.5 31 31 42 17 19.5 20 20 21 21.5 23 23 22 43 29 31 31 33 34 35 37 37 38 44 22 23 23 23 24.5 25 26 45 26 26 26.5 27.5 28 29 31 32 46 22 24 25 26 26.5 28 29 30 30 47 21 21 21 22 22.5 23 24 25 25 48 22 23.5 24 25 25 26 27 27.5 28 49 23 24.5 25.5 26 27 27.5 29.5 29.5 26 50 20 21 22 22 22 23 24.5 25 27
44
Table 4.2 Weight (Thalli 01-25). The weight of each thallus from the linear growth
subset, wase recorded each quarter in grams. (This table continues on to the next
page.)
Thallus Weight in Grams AUG NOV FEB MAY AUG NOV FEB MAY AUG Thallus 2011 2011 2012 2012 2012 2012 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 1 0.124 0.193 0.16 0.149 0.172 0.184 0.224 0.204 0.247 2 0.098 0.159 0.143 0.123 0.149 0.114 0.111 0.116 0.13 3 0.105 0.137 0.141 0.135 0.112 4 0.15 0.23 0.208 0.198 0.161 0.203 0.263 0.214 0.189 5 0.18 0.239 0.217 0.171 0.2 0.222 0.253 6 0.213 0.257 0.307 0.276 0.29 0.371 0.383 0.387 0.422 7 0.176 0.236 0.288 0.314 0.28 0.348 0.362 0.389 0.448 8 0.123 0.165 0.179 0.163 0.189 0.258 0.249 0.286 0.344 9 0.139 0.207 0.206 0.2 0.273 0.283 0.297 0.367 0.372 10 0.064 0.092 0.091 0.089 0.099 0.123 0.116 0.134 0.146 11 0.229 0.341 0.349 0.347 0.389 0.448 0.458 0.493 0.575 12 0.11 0.143 0.164 0.162 0.168 0.209 0.19 0.228 0.258 13 0.083 0.132 0.141 0.125 0.145 0.166 0.182 0.208 0.216 14 0.126 0.169 0.191 0.173 0.188 0.211 0.248 0.241 0.251 15 0.1 0.177 0.179 0.161 0.214 0.199 0.214 0.228 0.251 16 0.169 0.239 0.173 0.237 0.344 0.374 0.431 0.333 17 0.101 0.142 0.145 0.143 0.19 0.203 0.204 0.211 18 0.146 0.207 0.232 0.225 0.238 0.28 0.295 0.285 19 0.118 0.158 0.192 0.179 0.203 0.264 0.252 0.246 0.144 20 0.217 0.336 0.357 0.361 0.402 0.486 0.57 0.664 0.703 21 0.133 0.199 0.241 0.217 0.355 0.266 0.31 0.305 0.319 22 0.175 0.233 0.3 0.252 23 0.112 0.156 0.167 24 0.151 0.213 0.262 0.239 0.285 0.273 0.298 0.315 0.337 25 0.116 0.181 0.219 0.223 0.277 0.341 0.405 0.421 0.505
45
Table 4.2 Weight Continued (Thalli 26-50). The weight of each thallus from the
linear growth subset, was recorded each quarter in grams.
Thallus Weight in Grams AUG NOV FEB MAY AUG NOV FEB MAY AUG Thallus 2011 2011 2012 2012 2012 2012 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 26 0.172 27 0.077 0.109 0.121 0.122 0.169 0.162 0.2 0.215 0.233 28 0.18 0.257 0.273 0.335 0.402 0.486 0.57 0.664 0.703 29 0.084 30 0.134 0.192 0.198 0.2 0.25 0.269 0.323 0.329 0.352 31 0.121 0.154 0.185 0.177 0.203 0.203 0.254 0.255 0.301 32 0.113 0.156 0.173 0.178 0.199 0.234 0.267 33 0.076 0.1 0.106 0.112 0.134 0.143 0.156 0.171 0.195 34 0.067 0.085 0.104 0.105 0.115 0.133 0.167 0.163 35 0.083 0.122 0.107 0.149 36 0.151 0.206 0.216 0.211 0.224 0.253 0.274 0.301 0.322 37 0.101 0.134 0.161 0.153 0.187 0.214 0.246 0.275 0.26 38 0.147 0.221 0.247 0.28 0.32 0.388 0.451 0.2 0.334 39 0.055 0.077 0.082 0.087 0.106 0.114 0.14 40 0.098 0.141 0.155 0.182 0.206 0.244 0.311 41 0.155 0.232 0.304 0.303 0.351 0.439 0.521 0.51 0.561 42 0.043 0.054 0.075 0.08 0.091 0.118 0.135 0.134 0.143 43 0.216 0.328 0.361 0.406 0.474 0.532 0.664 0.689 0.749 44 0.111 0.182 0.204 0.224 0.232 0.283 0.369 45 0.159 0.197 0.18 0.213 0.244 0.266 0.371 0.327 46 0.15 0.197 0.264 0.287 0.314 0.348 0.467 0.441 0.525 47 0.054 0.079 0.085 0.098 0.108 0.123 0.18 0.162 0.19 48 0.091 0.138 0.176 0.175 0.191 0.192 0.168 0.221 0.22 49 0.172 0.256 0.284 0.299 0.394 0.424 0.475 0.52 0.395 50 0.089 0.136 0.148 0.15 0.212 0.204 0.231 0.196 0.173
46
Table 4.3 Length of 2° Branches (Thalli 1-25). The length of the 2° branches was
recorded on a quarterly basis for two years.(This table continues on to the next
page.)
Length of 2⁰ branches in millimeters OCT JAN APR JUL OCT JAN APR JUL OCT Thallus 2011 2012 2012 2012 2012 2013 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 1 5 5 5 5 5 6 6 6 6 2 3 4 4 4 6 6 6 6 6 3 3 4 4 4.5 5 8 8 8 8 4 3 3 3 3.5 3.5 4 5 5 3.5 3.5 3.5 3.5 6 6 6 2.5 2.5 3 3 7 1.5 2 8 5 5 9 6 6 6 6.5 7 7 7 8 8 10 3 11 6 6 6 6 7 7 7 7 7 12 2 2 2.5 2.5 4 4 5 13 6.5 7 14 3 6 6.5 7 8 10 10 10 10 15 3 5 5 5 6 7 16 2 2 3 3 5 5 5 6 17 6 6 6 6 18 2.5 2.5 2.5 2.5 3 3 3 19 7 8 8 9 9 9 9 6 7 20 4 4 4 4 21 6.5 6 6 6 6 22 2 23 7 7 8 8 10 11 11 11 11 24 4 4 5 5 25 3.5 4 4 4 4
47
Table 4.3 Continued (Thalli 26-50). The length of the 2° branches was recorded on a
quarterly basis for two years.
Length of 2⁰ branches in millimeters OCT JAN APR JUL OCT JAN APR JUL OCT Thallus 2011 2012 2012 2012 2012 2013 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 26 2 2 2 2 27 6 6 6 6 6 6 6 10 11 28 5 5 5 5 29 4 4 4 4 5.5 30 2 2 2 3 3 3 4 31 4 4 4.5 5 5 6 6 8 11 32 10 10 10 10 10 11 11 12 12 33 1 1 1.5 34 9 9 9 9 9 10 10 10 10.5 35 4 4 4 4 4 4 5 6 10 36 7 7 7 7 7 7 7.5 8 8 37 6 6 6 6 7 7 7 8 8 38 1 1 2 2 2 2 2 5 5 39 6 6 6 6 6 40 5 5 5 5 5 5 5 41 6 6 6 7 7 7 7 7 8 42 3 3 3 3 3 43 3.5 3.5 3.5 3.5 3.5 3.5 4 6 6 44 3 3 3 3 3 3 3 6.5 6.5 45 4 4 4 4 4.5 5 5 5 5 46 4 4 4 4 4 47 3 3 3 3 3 3 4 6 9 48 3 3 4 4.5 7.5 11 11 11 12 49 6 6 6 6 7 7 50 6 6 6 5 6 13 13 13 14.5
48
Table 4.4 Length of 3° Branches (Thalli 1-25). The length of the 3° branches was
recorded on a quarterly basis for two years. (This table continues on to the next
page.)
Length of 3⁰ branches in millimeters OCT JAN APR JUL OCT JAN APR JUL OCT Thallus 2011 2012 2012 2012 2012 2013 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 1 3 3 4 4 4 4.5 7 7 7 2 9 9 9 10 10 10 10 11 13 3 3 4 4 4 4 4 4 4.5 4.5 4 6.5 7 7 7 7 8 8 5 3 3 3 3 3 4 6 3 4 4 4 7 2 3 8 15 15 9 3 3.5 3.5 4 4 4 4 6 6 10 10 11 5 5 5 5 5 5 5 5 5 12 2.5 2.5 2.5 2.5 2.5 2.5 2.5 13 10.5 11 14 15 16 16 16 16 16 16 16 16 15 9.5 10 10 10 10 10 16 2 2 3 3 5 5 5 5 17 13.5 14 14 14 18 7 7 7 7 7.5 7.5 7.5 19 12 12 12 13 13 13 13 13.5 13.5 20 11 11 11 11 21 4.5 5 5 5 5 22 3 23 9 10 10 10 10 10 10 10 10 24 7 7 7 8 25 9 9 9 9 9
49
Table 4.4 Length of 3° Branches Continued (Thalli 26-50). The length of the 3°
branches was recorded on a quarterly basis for two years.
Length of 3⁰ branches in millimeters OCT JAN APR JUL OCT JAN APR JUL OCT Thallus 2011 2012 2012 2012 2012 2013 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 26 4 4.5 4.5 4.5 27 11 11.5 11.5 12 12 12 12 12 13 28 9 9 9 11 29 7 7 7 8 8 30 4 4 4 4 4 5 5 31 6 6 6 6 7 7 7 7 8 32 6 6 6 6 6 6 8 8 9 33 15 15 15 34 8 8 8 8 8 8.5 8.5 8.5 9 35 11 11 11 11.5 12 12 12 12 13 36 14 14 14 14 14 14 14 15 15 37 8 8 8 8 8 8 8.5 8.5 9 38 5 5 5 5 5 5 6 6 6 39 4 4 5 5 6 40 3 4 4 4 4 4 4 41 4 4 5 5 5 5 5 5 6 42 3 3 3 3.5 3.5 43 11 11 10 10 10 11 11 11 11 44 7 7 7 7 7 8 8 8.5 8.5 45 9 9 9 9 9 9 9 9 9 46 4 4 4.5 5 10 47 5 5 5 5 5 5 5 5.5 6 48 3 3 3.5 4 4.5 4.5 6 6 6 49 10 10 10 10 10 10 50 16 16 16 16 16 17 19 19 19
50
Table 4.5 Length of 4° Branches (Thalli 2-32). The length of the 4° branches was
recorded on a quarterly basis for two years. (This table continues on to the next
page.)
Length of 4⁰ branches in millimeters OCT JAN APR JUL OCT JAN APR JUL OCT Thallus 2011 2012 2012 2012 2012 2013 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 2 6 6.5 7 7 7 7 7 8 8 3 8 9 9 9 10 10 10 10 10 4 6 6 6 6 7 8.5 10 5 12 12 12 12 12 12 6 9.5 10 10 10 7 16 16 9 14 14 14 14 14 14 14 10 4 11 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 5.5 12 4.5 4.5 4.5 4.5 5 5 6 13 2 2 16 10 10 10 10 10 10 10 10.5 18 7 8 8 8 8 8 8 20 7 7 7 7 21 10 11 11 11 11 22 7 23 5 5 6 7 7 7 9 9 9 24 7 7 7 8 25 14.5 15 15.5 16 16 26 8 8 8 8 28 23 23 23 23 29 7 7 7 8 8 30 22 23 23 24 24 24 24 31 6 6 6 6 6 7 7 7 7 32 16 16 16 16 16.5 16.5 16.5 16.5 16.5
51
Table 4.5 Length of 4° Branches Continued (Thalli 33-49). The length of the 4°
branches was recorded on a quarterly basis for two years.
Length of 4⁰ branches in millimeters OCT JAN APR JUL OCT JAN APR JUL OCT Thallus 2011 2012 2012 2012 2012 2013 2013 2013 2013 ID Week Week Week Week Week Week Week Week Week 0 13 26 39 52 65 78 91 104 33 5 5 5 36 6.5 6.5 6.5 6.5 6.5 6.5 7 7 7 37 6.5 6.5 6.5 6.5 6.5 6.5 7 7 7 38 16.5 16.5 17 18 18 18 18 18 18 39 9 9 9 9 9.5 40 18 19 19 20 20 20 20 41 14 15 15 15 15 15 15 15 15 42 12 12 12 12 12 43 21 21 21 21 21.5 21.5 21.5 21.5 21.5 44 6 6 6 6 6 6 6 6 6 45 8 8 9 10 10 10 10 10 10 46 15 15 15.5 16 16 47 15 15 16 16 16 16 16 16 16 48 18 18 18 18 19 19 19 19 20 49 2 2 2 4 4 4
52
Fragmentation
Figure 5.1 Fragment Figure 5.2 Artificial Scrub Container
For identification purposes and to prevent the loss of fragments, individual fragments
were placed in cylinders on artificial scrub at a private residence.
Thalli Fragment Survival
120 100 100 99 9397 95 86 8387 80 78 77 79 2 mm live 70 74 60 57 4 mm live 48 40 42 6 mm live 20 22 2425 8 mm live 8
Surviving Surviving Thalli Fragments 0 10 mm live time 0 13 weeks 26 weeks 39 weeks 52 weeks Time (3months per quarter)
Figure 5.3 Fragment Survivability
Fragment survival was recorded in each size class every quarter (13 weeks) for one year.
53
Percent Fragment Mortality Per Quarter 70.00%
63.64% 60.00% 57.89% 50.00% 46.15% 47.62% 47.92% 2 mm 40.00% 37.66% 4 mm 30.00% 29.29% 22.00% 22.22% 6 mm 20.00% 18.57% 10.84% Fragment Fragment Mortality 10.00% 8 mm 7.00% 7.53% 8.42% 9.20% 0.00% 1.00%3.00% 2.06% 3.49% 10 mm 1st quarter 2nd quarter 3rd quarter 4th quarter Time (3 months per quarter)
Figure 5.4 Fragment Mortality Per Quarter
The above figure illustrates the percent of fragments that died over the course of each 13
week quarter.
Average Fragment Lengths
15.00
10.00 10.00 10.00 9.76 9.77 10.14 8.00 8.00 7.92 8.11 7.96 6.58 6.62 5.00 6.00 5.96 5.96 4.00 3.99 3.96 3.90 4.52 2.00 1.99 2.02 2.00 2.00
Size Size millimeters in 0.00 time 0 13 weeks 26 weeks 39 weeks 52 weeks Time (3 months per quarter)
Figure 5.5 Average Fragment Lengths
Fragment length was recorded every 13 weeks and averaged to track growth for the one
year period.
54
Average Quarterly Mortality 50.00% 44.85% 40.00% 27.20% 26.96% 30.00% 20.00% 7.21% Average quarterly 10.00% 5.67% mortality rate
0.00%
2 mm 2 mm 4 mm 6 mm 8 10 mm10 Fragment class size
Figure 5.6 Average Quarterly Mortality
The percent mortality was recorded for each fragment size each quarter, and then
averaged to determine average quarterly mortality.
Average Percent Weight Change
120.00% 102.14%
100.00%
80.00% 64.16%
60.00% 45.94% 2 mm
37.85% 36.55% 40.00% 33.79%
4 mm 4.24% 20.00% 2.07%
0.00% 6 mm
-20.00%
-40.00% 8 mm
4.34%
5.44%
6.08%
6.20%
-
- -
-60.00% -
10.28%
11.13% Average Average Change Weight
13.15% 10 mm
-
15.87%
-
-
- 21.58%
-80.00% 24.53%
-
- 31.18%
-100.00% - 71.45% First Quarter Second Quarter- Third Quarter Fourth Quarter
Figure 5.7 Average Percent Weight Change
Percent weight change was recorded for each thallus, and then averaged for each quarter.
55
Table 5.1 Fragment Growth (Fragments 1-25). Length and weight of the 2 mm
fragment class for 1 year (continues on to next page).
2 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 1 2.00 1.73 2 2.00 2.00 2.50 2.00 1.90 2.10 4.50 8.70 3 2.00 1.97 4 2.00 1.30 5 2.00 2.00 2.00 2.00 2.80 3.80 2.00 0.80 6 2.00 2.00 1.27 2.50 7 2.00 2.00 1.27 3.40 8 2.00 1.20 0.00 9 2.00 2.00 2.00 1.45 4.50 3.00 10 2.00 2.00 2.00 1.00 1.90 1.50 11 2.00 2.00 2.00 2.00 2.00 1.30 4.40 2.10 3.60 2.50 12 2.00 2.00 2.00 2.00 2.27 5.30 3.60 2.50 13 2.00 1.40 14 2.00 2.00 2.00 2.00 2.33 5.30 2.50 7.20 15 2.00 2.00 2.13 4.60 16 2.00 2.00 2.03 2.80 17 2.00 1.40 0.00 18 2.00 2.00 2.00 2.00 2.00 1.27 2.90 4.10 8.50 3.30 19 2.00 1.30 20 2.00 2.00 2.03 4.30 21 2.00 2.00 2.50 2.00 1.90 4.40 3.40 11.30 22 2.00 2.00 2.00 1.40 4.90 2.50 23 2.00 2.00 2.00 1.73 2.20 5.00 24 2.00 2.00 2.90 5.90 25 2.00 2.00 2.00 2.17 3.00 1.10
56
Table 5.1 Continued (Fragments 26-50). Length and weight of the 2 mm fragment
class for 1 year (continues on to next page).
2 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 26 2.00 2.00 2.00 2.20 2.00 2.50 27 2.00 2.00 2.00 2.00 2.00 2.37 6.20 3.30 7.00 6.20 28 2.00 2.00 1.73 3.50 29 2.00 2.00 2.00 2.00 1.50 2.60 3.70 5.80 30 2.00 0.03 31 2.00 1.17 32 2.00 2.00 2.00 2.30 4.50 4.10 33 2.00 2.00 2.00 2.33 4.80 5.00 34 2.00 2.00 2.00 2.00 3.60 8.20 6.50 6.20 35 2.00 2.00 2.00 3.17 4.80 4.00 36 2.00 2.00 2.00 2.00 2.30 3.60 2.40 7.70 37 2.00 2.00 2.00 2.00 2.57 4.80 6.20 6.10 38 2.00 2.00 2.00 2.07 4.50 2.50 0.00 39 2.00 2.00 2.00 2.00 2.07 4.30 2.60 5.60 40 2.00 2.00 1.17 2.10 41 2.00 2.00 1.87 3.70 42 2.00 1.90 0.00 43 2.00 2.00 1.50 3.00 44 2.00 1.25 45 2.00 1.10 46 2.00 2.00 2.00 2.90 13.60 6.60 47 2.00 2.00 2.00 1.30 3.20 3.20 48 2.00 1.20 49 2.00 2.00 2.20 3.50 50 2.00 2.00 1.90 2.10
57
Table 5.1 Continued (Fragments 51-75). Length and weight of the 2 mm fragment
class for 1 year (continues on to next page).
2 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 51 2.00 1.10 52 2.00 1.00 53 2.00 1.00 1.90 1.00 54 2.00 1.10 55 2.00 2.00 2.00 2.00 2.00 1.80 2.80 2.90 5.70 6.00 56 2.00 1.20 57 2.00 0.90 58 2.00 2.00 1.60 3.70 59 2.00 2.00 2.00 2.00 2.00 3.20 5.60 4.00 7.60 5.50 60 2.00 2.00 1.10 2.30 61 2.00 2.00 2.00 2.00 3.20 7.40 3.70 -3.00 62 2.00 2.00 2.00 3.90 7.40 4.10 63 2.00 1.20 64 2.00 2.00 1.50 4.50 65 2.00 2.00 2.00 2.00 2.00 3.00 6.10 4.10 6.90 3.70 66 2.00 1.50 67 2.00 2.00 2.00 2.00 2.30 5.70 2.90 8.00 68 2.00 2.00 2.00 2.00 2.00 2.00 3.30 4.20 8.00 4.40 69 2.00 2.00 2.00 2.00 2.00 3.20 6.50 7.60 2.70 7.40 70 2.00 2.00 2.00 2.20 7.90 3.00 71 2.00 2.00 2.00 2.00 2.70 5.20 4.90 7.10 72 2.00 2.00 2.00 2.20 4.00 2.10 73 2.00 1.60 74 2.00 2.00 3.00 5.90 75 2.00 2.00 1.80 6.20
58
Table 5.1 Continued (Fragments 76-100). Length and weight of the 2 mm fragment
class for 1 year.
2 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 76 2.00 1.20 77 2.00 2.00 1.60 5.70 78 2.00 2.00 2.00 2.20 5.20 2.10 79 2.00 2.00 2.10 7.70 80 2.00 2.00 2.00 2.00 2.20 3.40 3.10 2.80 81 2.00 2.00 2.00 3.60 4.20 4.10 82 2.00 2.00 2.00 2.00 5.10 3.60 83 2.00 2.00 2.00 1.60 1.80 4.30 84 2.00 2.00 1.70 2.30 85 2.00 2.00 3.20 5.90 86 2.00 2.00 4.00 3.90 87 2.00 2.00 1.70 2.80 88 2.00 2.00 2.20 4.10 89 2.00 2.00 2.40 3.90 90 2.00 2.00 4.50 7.30 91 2.00 2.00 4.00 4.50 92 2.00 2.00 2.00 2.40 6.80 2.30 93 2.00 2.00 1.80 3.00 94 2.00 2.00 1.90 3.20 95 2.00 2.00 2.10 3.50 96 2.00 2.00 3.00 3.50 97 2.00 2.00 2.90 5.30 98 2.00 2.00 3.00 6.20 99 2.00 2.00 3.30 5.20 100 2.00 2.00 2.80 5.00
59
Table 5.2 Fragment Growth (Fragments 101-125). Length and weight of the 4 mm
fragment class for 1 year (continues on to next page).
4 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 101 4.00 4.00 4.00 4.00 4.00 4.18 6.80 6.50 6.20 3.50 102 4.00 4.00 7.60 13.60 0.00 0.00 0.00 103 4.00 4.00 4.00 4.00 4.00 14.10 24.30 22.40 13.80 5.70 104 4.00 4.00 4.00 4.00 7.67 15.60 13.60 9.40 0.00 105 4.00 4.00 3.00 4.00 4.00 9.85 19.60 9.90 6.80 3.40 106 4.00 4.00 4.47 7.50 0.00 0.00 0.00 107 4.00 4.00 6.27 9.50 0.00 0.00 0.00 108 4.00 4.00 4.00 8.80 14.10 8.60 0.00 0.00 109 4.00 4.00 4.00 4.00 8.23 11.80 12.90 6.20 0.00 110 4.00 4.00 8.47 12.20 0.00 0.00 0.00 111 4.00 4.00 4.00 4.00 5.00 12.27 15.80 14.30 23.90 20.50 112 4.00 4.00 4.00 14.27 29.00 11.00 0.00 0.00 113 4.00 4.00 4.00 4.00 4.00 12.90 24.50 20.70 8.10 7.00 114 4.00 4.00 4.00 4.00 4.00 9.07 11.90 17.40 11.60 4.10 115 4.00 4.00 4.00 5.70 7.80 1.90 0.00 0.00 116 4.00 4.00 4.00 11.77 17.50 17.50 0.00 0.00 117 4.00 4.00 4.00 4.00 8.47 14.90 10.70 2.70 0.00 118 4.00 4.00 4.00 8.87 15.90 7.20 0.00 0.00 119 4.00 4.00 4.00 5.37 8.50 11.50 0.00 0.00 120 4.00 4.00 6.63 9.90 0.00 0.00 0.00 121 4.00 4.00 6.70 8.90 0.00 0.00 0.00 122 4.00 4.00 4.00 4.00 5.00 12.10 28.00 19.00 5.60 11.10 123 4.00 4.00 4.00 12.83 20.90 14.50 0.00 0.00 124 4.00 4.00 4.00 12.73 16.00 10.40 0.00 0.00 125 4.00 4.00 4.00 15.53 17.70 23.50 0.00 0.00
60
Table 5.2 Continued (Fragments 126-150). Length and weight of the 4 mm fragment
class for 1 year (continues on to next page).
4 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 126 4.00 4.00 7.95 17.20 0.00 0.00 0.00 127 4.00 4.00 4.00 5.20 6.20 10.40 0.00 0.00 128 4.00 4.00 4.00 4.00 5.00 11.30 13.70 14.00 14.00 12.00 129 4.00 4.00 4.00 8.00 9.50 7.10 0.00 0.00 130 4.00 4.00 4.00 12.10 9.90 10.30 0.00 0.00 131 4.00 4.00 7.30 11.40 0.00 0.00 0.00 132 4.00 4.00 4.50 6.00 0.00 0.00 0.00 133 4.00 4.00 4.00 7.90 10.60 3.50 0.00 0.00 134 4.00 4.00 10.70 20.20 0.00 0.00 0.00 135 4.00 4.00 11.40 19.30 0.00 0.00 0.00 136 4.00 4.00 4.00 3.00 7.25 8.20 7.50 2.40 0.00 137 4.00 4.00 4.00 12.80 19.10 14.60 0.00 0.00 138 4.00 4.00 3.00 4.00 10.50 12.60 12.60 10.70 0.00 139 4.00 4.00 4.00 4.00 7.65 11.80 15.40 2.20 0.00 140 4.00 4.00 4.00 9.40 9.60 11.90 0.00 0.00 141 4.00 4.00 10.75 12.90 0.00 0.00 0.00 142 4.00 4.00 4.00 8.05 11.50 13.20 0.00 0.00 143 4.00 4.00 2.00 2.60 4.60 4.50 0.00 0.00 144 4.00 4.00 4.00 4.00 9.75 11.00 16.80 8.80 0.00 145 4.00 4.00 4.00 18.45 21.40 13.70 0.00 0.00 146 4.00 4.00 4.00 8.27 11.50 4.50 0.00 0.00 147 4.00 4.00 4.00 4.00 5.85 10.00 7.10 4.60 0.00 148 4.00 4.00 4.00 9.35 13.50 13.20 0.00 0.00 149 4.00 4.00 4.00 4.00 4.00 15.35 13.90 25.00 15.50 10.70 150 4.00 4.00 4.00 4.00 8.55 16.30 7.00 16.10 0.00
61
Table 5.2 Continued (Fragments 151-175). Length and weight of the 4 mm fragment
class for 1 year (continues on to next page).
4 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 151 4.00 4.00 4.00 4.00 11.35 14.00 13.80 8.80 0.00 152 4.00 3.00 7.10 9.90 0.00 0.00 0.00 153 4.00 4.00 4.00 9.90 11.10 11.60 0.00 0.00 154 4.00 4.00 4.00 4.00 4.00 19.45 25.80 27.80 21.50 24.60 155 4.00 4.00 4.00 4.00 19.05 22.00 18.90 11.40 0.00 156 4.00 4.00 4.00 4.00 11.00 11.90 13.00 15.10 0.00 157 4.00 4.00 4.00 4.00 14.80 18.20 10.80 11.00 0.00 158 4.00 4.00 4.00 4.00 4.00 12.25 13.90 13.30 13.30 20.20 159 4.00 4.00 4.00 4.00 5.00 7.70 9.20 11.80 7.70 7.10 160 4.00 4.00 4.00 4.00 7.55 15.50 13.70 4.80 0.00 161 4.00 4.00 4.00 4.00 5.00 10.45 19.70 24.00 8.80 13.30 162 4.00 4.00 4.00 4.00 4.00 13.45 19.70 22.40 20.70 14.50 163 4.00 4.00 11.30 12.80 0.00 0.00 0.00 164 4.00 4.00 4.00 4.00 14.65 17.30 18.60 17.20 0.00 165 4.00 4.00 4.00 4.00 12.43 13.30 13.20 3.30 0.00 166 4.00 4.00 4.00 4.00 5.00 16.60 19.00 17.70 16.40 20.00 167 4.00 4.00 4.00 8.90 13.50 17.00 0.00 0.00 168 4.00 4.00 4.00 12.35 20.80 14.30 0.00 0.00 169 4.00 4.00 4.00 4.00 5.00 7.05 9.00 11.20 10.20 12.10 170 4.00 4.00 4.00 4.00 4.00 9.75 13.60 13.20 7.60 12.20 171 4.00 4.00 4.00 4.00 14.05 21.30 24.40 18.20 0.00 172 4.00 4.00 4.00 4.00 5.00 27.10 42.80 30.00 11.80 28.40 173 4.00 4.00 4.00 15.95 20.20 11.60 0.00 0.00 174 4.00 4.00 4.00 4.00 12.40 21.70 23.00 15.00 0.00 175 4.00 4.00 4.00 4.00 4.00 5.85 7.20 7.00 11.60 6.90
62
Table 5.2 Continued (Fragments 176-200). Length and weight of the 4 mm fragment
class for 1 year.
4 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 176 4.00 4.00 4.00 4.00 4.00 9.70 11.10 12.90 14.10 10.60 177 4.00 4.00 4.00 4.00 5.00 17.60 16.80 18.50 16.60 18.10 178 4.00 4.00 4.00 4.00 6.00 9.00 6.20 2.20 0.00 179 4.00 4.00 4.45 8.90 0.00 0.00 0.00 180 4.00 4.00 8.13 11.00 0.00 0.00 0.00 181 4.00 4.00 8.27 11.50 0.00 0.00 0.00 182 4.00 4.00 6.70 8.10 0.00 0.00 0.00 183 4.00 4.00 10.57 14.80 0.00 0.00 0.00 184 4.00 4.00 4.00 4.00 5.00 6.73 9.40 7.40 8.20 6.80 185 4.00 4.00 4.00 4.00 6.00 11.23 15.20 12.80 12.90 12.40 186 4.00 4.00 4.00 12.23 23.80 8.90 0.00 0.00 187 4.00 4.00 4.00 8.55 15.20 7.10 0.00 0.00 188 4.00 4.00 4.55 6.10 0.00 0.00 0.00 189 4.00 4.00 6.30 12.90 0.00 0.00 0.00 190 4.00 4.00 4.00 16.67 18.70 7.50 0.00 0.00 191 4.00 4.00 4.00 14.90 20.30 13.10 0.00 0.00 192 4.00 4.00 4.00 4.00 4.00 13.05 16.80 16.40 9.90 11.60 193 4.00 4.00 4.00 4.00 11.20 17.70 12.80 10.20 0.00 194 4.00 9.15 0.00 0.00 0.00 0.00 195 4.00 4.00 4.00 8.70 10.40 8.90 0.00 0.00 196 4.00 4.00 4.00 4.00 5.00 10.75 12.30 11.90 12.80 10.50 197 4.00 4.00 4.00 3.00 7.65 9.30 5.60 0.80 0.00 198 4.00 4.00 4.00 4.00 15.47 27.20 5.40 7.50 0.00 199 4.00 4.00 5.00 1.00 8.33 19.70 11.50 1.40 0.00 200 4.00 4.00 6.55 8.40 0.00 0.00 0.00
63
Table 5.3 Fragment Growth (Fragments 201-225). Length and weight of the 6 mm
fragment class for 1 year (continues on to next page).
6 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 201 6.00 6.00 6.00 14.25 23.10 6.40 202 6.00 6.00 6.00 6.00 57.50 68.80 23.00 26.40 203 6.00 6.00 6.00 9.00 7.00 50.85 55.70 52.20 62.10 58.30 204 6.00 6.00 6.00 5.00 30.70 37.30 8.80 7.70 205 6.00 6.00 6.00 7.00 7.00 15.10 16.80 21.00 22.00 19.40 206 6.00 6.00 6.00 8.00 8.00 31.35 34.00 42.70 39.90 45.50 207 6.00 6.00 6.00 8.00 24.60 26.30 29.60 27.80 208 6.00 6.00 17.40 22.30 209 6.00 6.00 6.00 6.00 14.70 19.00 15.80 19.30 210 6.00 6.00 6.00 5.00 18.40 24.70 5.10 7.00 211 6.00 6.00 17.55 16.30 212 6.00 6.00 6.00 7.00 7.45 11.10 10.70 8.00 213 6.00 6.00 18.65 36.20 214 6.00 6.00 6.00 6.00 32.30 31.10 4.40 6.10 215 6.00 6.00 16.50 21.10 216 6.00 6.00 19.40 23.60 217 6.00 6.00 15.75 22.59 218 6.00 6.00 6.00 5.00 12.65 15.00 6.20 5.00 219 6.00 6.00 6.00 20.70 23.50 16.40 220 6.00 6.00 6.00 8.00 7.00 24.90 31.00 45.00 51.60 29.50 221 6.00 6.00 6.00 5.00 18.85 24.90 14.90 12.60 222 6.00 6.00 6.00 6.00 22.45 28.90 12.90 9.60 223 6.00 6.00 6.00 7.00 11.10 12.40 16.50 11.70 224 6.00 6.00 6.00 6.00 7.00 13.40 18.90 27.20 33.10 30.40 225 6.00 6.00 6.00 8.00 12.80 15.70 13.50 12.90
64
Table 5.3 Continued (Fragments 226-250). Length and weight of the 6 mm fragment
class for 1 year (continues on to next page).
6 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 226 6.00 6.00 6.00 15.10 20.60 11.30 227 6.00 6.00 6.00 8.95 10.50 9.40 228 6.00 6.00 6.00 18.35 23.30 29.10 229 6.00 14.20 230 6.00 6.00 11.45 20.60 231 6.00 6.00 5.00 5.00 23.20 32.00 23.60 15.20 232 6.00 6.00 6.00 5.00 16.75 32.20 23.10 16.30 233 6.00 6.00 6.00 47.65 66.00 20.20 234 6.00 6.00 11.80 14.60 235 6.00 6.00 6.00 7.00 7.00 29.45 37.30 43.90 36.20 55.70 236 6.00 6.00 6.00 6.00 16.40 18.90 19.30 16.60 237 6.00 6.00 34.45 32.60 238 6.00 6.00 7.55 12.30 239 6.00 6.00 10.65 14.80 240 6.00 6.00 6.00 6.00 23.35 29.20 21.00 21.30 241 6.00 6.00 6.00 6.00 28.50 45.20 43.60 0.60 242 6.00 6.00 6.00 4.00 31.40 31.10 29.50 16.40 243 6.00 6.00 17.30 28.60 244 6.00 6.00 23.70 32.10 245 6.00 6.00 29.40 34.60 246 6.00 2.00 31.90 7.00 247 6.00 6.00 21.90 34.50 248 6.00 6.00 6.00 7.65 11.80 6.10 249 6.00 6.00 6.00 28.40 29.40 10.40 250 6.00 6.00 4.00 4.00 17.30 21.70 25.75 29.80
65
Table 5.3 Continued (Fragments 251-275). Length and weight of the 6 mm fragment
class for 1 year (continues on to next page).
6 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 251 6.00 6.00 12.30 15.90 252 6.00 6.00 35.00 42.80 253 6.00 6.00 6.00 22.90 36.90 12.10 254 6.00 6.00 6.00 8.00 6.00 36.20 62.30 41.80 16.70 12.50 255 6.00 6.00 6.00 5.00 16.30 23.70 7.30 9.70 256 6.00 6.00 6.00 7.00 7.00 27.15 42.00 31.70 43.20 21.00 257 6.00 6.00 6.00 6.00 6.00 33.00 33.00 23.70 19.20 13.60 258 6.00 6.00 6.00 8.00 8.00 39.55 47.40 49.00 52.00 48.40 259 6.00 6.00 6.00 7.00 4.00 18.95 25.90 33.30 25.90 12.50 260 6.00 6.00 6.00 19.55 32.40 25.50 261 6.00 6.00 14.10 17.80 262 6.00 6.00 10.90 21.10 263 6.00 6.00 11.20 27.40 264 6.00 6.00 15.70 21.70 265 6.00 6.00 22.55 38.80 266 6.00 6.00 6.00 7.00 23.75 33.00 12.90 11.50 267 6.00 6.00 6.00 8.00 31.90 54.20 33.90 21.60 268 6.00 6.00 6.00 6.00 28.90 34.50 14.90 9.70 269 6.00 6.00 6.00 7.00 19.60 29.00 14.70 13.70 270 6.00 6.00 6.00 7.00 6.00 22.70 24.40 30.60 29.30 25.50 271 6.00 6.00 6.00 7.00 6.00 17.55 29.70 17.30 26.50 21.40 272 6.00 6.00 6.00 7.00 5.00 16.45 24.90 19.20 13.20 14.00 273 6.00 6.00 6.00 6.00 16.60 24.50 16.20 8.10 274 6.00 6.00 6.00 6.00 16.85 25.50 17.90 7.30 275 6.00 6.00 6.00 19.35 32.50 10.30 0.00
66
Table 5.3 Continued (Fragments 276-300). Length and weight of the 6 mm fragment
class for 1 year.
6 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 276 6.00 6.00 6.00 6.00 12.55 17.60 11.90 7.10 277 6.00 6.00 6.00 7.00 31.15 40.70 28.10 31.30 278 6.00 6.00 8.80 18.50 279 6.00 6.00 6.00 8.00 29.35 43.50 12.30 13.90 280 6.00 6.00 6.00 7.00 6.00 24.50 24.40 29.50 23.80 25.40 281 6.00 6.00 6.00 6.00 10.45 14.30 11.90 9.70 0.00 282 6.00 6.00 29.30 57.80 283 6.00 6.00 18.25 30.40 284 6.00 6.00 6.00 22.60 26.00 28.70 285 6.00 6.00 13.85 21.90 286 6.00 6.00 26.20 42.50 287 6.00 6.00 6.00 5.00 22.90 24.20 12.80 11.20 288 6.00 6.00 6.00 8.00 7.00 22.70 26.80 27.20 29.80 28.40 289 6.00 6.00 6.00 6.00 9.00 33.20 46.40 21.10 14.80 10.80 290 6.00 6.00 6.00 6.00 6.00 29.60 37.60 43.50 37.70 26.70 291 6.00 6.00 6.00 6.00 7.00 26.90 30.00 40.00 9.10 34.60 292 6.00 6.00 6.00 7.00 6.00 16.35 24.50 19.50 17.90 16.10 293 6.00 6.00 6.00 8.00 3.00 20.50 33.10 31.40 24.40 11.40 294 6.00 6.00 6.00 6.00 20.35 19.60 24.80 21.20 295 6.00 6.00 6.00 8.00 8.00 16.60 21.80 26.00 24.80 32.30 296 6.00 6.00 6.00 8.00 8.00 17.90 23.60 23.40 26.40 19.30 297 6.00 6.00 17.70 25.80 298 6.00 6.00 6.00 8.00 15.25 19.60 18.40 21.30 299 6.00 6.00 6.00 8.00 8.00 19.60 22.90 25.60 28.70 20.20 300 6.00 6.00 6.00 15.30 20.20 1.90
67
Table 5.4 Fragment Growth (Fragments 301-325). Length and weight of the 8 mm
fragment class for 1 year (continues on to next page).
8 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 301 8.00 8.00 6.00 7.00 20.20 38.30 26.90 13.90 302 8.00 8.00 8.00 7.00 7.00 20.10 31.80 31.40 29.68 21.50 303 8.00 8.00 8.00 8.00 8.00 13.90 22.80 26.90 23.80 22.80 304 8.00 8.00 8.00 9.00 8.00 37.70 75.10 78.30 61.60 45.10 305 8.00 8.00 8.00 9.00 8.00 40.80 52.10 58.10 47.10 35.00 306 8.00 8.00 8.00 31.50 49.50 36.40 307 8.00 8.00 8.00 9.00 9.00 18.00 26.90 27.40 29.50 33.70 308 8.00 8.00 8.00 7.00 5.00 27.10 44.50 41.00 13.60 10.80 309 8.00 8.00 8.00 9.00 9.00 38.10 44.50 53.30 54.30 58.50 310 8.00 8.00 8.00 8.00 8.00 35.40 60.20 51.90 71.60 69.50 311 8.00 8.00 8.00 9.00 9.00 38.00 58.40 56.70 56.00 61.50 312 8.00 8.00 8.00 9.00 9.00 29.00 45.30 52.60 42.70 34.70 313 8.00 8.00 8.00 9.00 9.00 29.50 49.60 61.20 42.00 46.20 314 8.00 8.00 8.00 9.00 9.00 42.30 75.30 45.20 58.70 65.60 315 8.00 8.00 6.00 23.30 45.30 14.30 316 8.00 8.00 8.00 9.00 9.00 42.20 55.60 54.60 49.30 64.70 317 8.00 8.00 8.00 8.00 9.00 46.90 58.90 66.20 55.70 72.70 318 8.00 8.00 8.00 8.00 8.00 28.00 42.20 32.50 35.10 22.90 319 8.00 8.00 8.00 8.00 8.00 22.30 35.00 33.80 14.10 11.90 320 8.00 8.00 8.00 8.00 8.50 44.50 50.00 52.40 46.50 52.50 321 8.00 8.00 8.00 8.00 8.00 51.90 70.60 58.20 66.90 24.50 322 8.00 8.00 8.00 8.50 9.00 20.70 37.80 35.90 38.90 30.60 323 8.00 8.00 8.00 6.00 7.00 41.60 71.50 46.50 42.50 28.50 324 8.00 8.00 8.00 8.00 8.00 35.60 43.90 30.10 22.90 18.20 325 8.00 8.00 8.00 8.50 9.00 54.40 80.10 54.00 62.90 74.90
68
Table 5.4 Continued (Fragments 326-350). Length and weight of the 8 mm fragment
class for 1 year (continues on to next page).
8 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 326 8.00 8.00 8.00 9.00 8.00 19.50 35.50 30.60 29.00 25.90 327 8.00 8.00 8.00 26.20 38.80 46.00 328 8.00 8.00 8.00 6.00 21.40 38.60 18.20 14.80 0.00 329 8.00 8.00 8.00 8.00 5.00 25.80 49.40 24.30 24.10 8.80 330 8.00 8.00 8.00 8.00 8.00 52.80 76.80 75.80 74.80 53.00 331 8.00 8.00 8.00 6.00 5.00 21.10 33.10 16.60 16.30 9.60 332 8.00 8.00 8.00 7.00 7.00 30.50 36.60 17.60 18.90 9.40 333 8.00 8.00 8.00 10.00 7.50 40.90 41.90 58.90 56.90 61.20 334 8.00 8.00 8.00 8.50 8.50 34.20 43.90 30.80 34.50 27.60 335 8.00 8.00 8.00 9.00 9.00 34.90 33.00 37.70 37.10 30.20 336 8.00 8.00 8.00 9.50 9.50 29.60 56.60 49.80 46.80 39.40 337 8.00 8.00 8.00 8.00 10.00 32.60 52.00 45.20 49.70 37.40 338 8.00 8.00 8.00 7.00 8.00 10.20 44.30 48.90 35.30 33.50 339 8.00 8.00 8.00 8.00 8.00 18.20 16.00 16.70 22.20 28.60 340 8.00 8.00 8.00 8.50 8.50 18.20 28.40 38.10 32.90 36.90 341 8.00 8.00 8.00 10.50 8.00 35.30 35.50 32.20 65.20 34.80 342 8.00 8.00 8.00 9.00 8.00 32.60 36.00 27.50 41.10 39.30 343 8.00 8.00 8.00 8.00 9.00 54.40 62.00 66.30 67.90 66.40 344 8.00 8.00 8.00 8.00 7.00 24.70 41.70 37.10 29.70 28.00 345 8.00 8.00 8.00 8.00 7.00 12.90 21.20 17.40 20.90 20.00 346 8.00 8.00 8.00 7.00 7.00 21.10 32.10 25.10 22.90 17.60 347 8.00 8.00 8.00 10.00 7.00 21.80 35.80 58.40 42.80 50.80 348 8.00 8.00 22.10 27.20 349 8.00 8.00 8.00 10.00 9.00 41.10 60.80 56.60 64.90 65.70 350 8.00 8.00 8.00 32.90 47.70 51.70
69
Table 5.4 Continued (Fragments 351-375). Length and weight of the 8 mm fragment
class for 1 year (continues on to next page).
8 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 351 8.00 26.80 352 8.00 8.00 8.00 9.00 8.00 27.60 44.40 31.00 21.00 22.10 353 8.00 8.00 8.00 7.50 8.00 18.10 28.50 24.20 31.00 35.20 354 8.00 8.00 8.00 7.50 8.00 20.70 25.90 20.30 35.30 21.10 355 8.00 8.00 8.00 9.00 9.00 28.20 31.90 41.70 48.20 53.50 - 356 8.00 8.00 8.00 9.00 9.00 62.40 47.60 39.30 39.00 43.40 357 8.00 26.20 358 8.00 8.00 8.00 9.00 16.10 31.50 17.40 22.90 0.00 359 8.00 8.00 8.00 9.00 9.00 31.20 37.50 46.30 42.40 38.60 360 8.00 8.00 8.00 6.00 5.00 24.80 38.10 25.70 32.70 23.40 361 8.00 8.00 8.00 6.00 8.00 23.80 23.20 28.20 24.30 24.50 362 8.00 8.00 8.00 8.00 8.00 18.60 29.30 14.10 41.60 31.00 363 8.00 8.00 8.00 9.00 8.00 17.80 30.90 24.60 26.90 17.70 364 8.00 8.00 8.00 8.00 8.00 34.00 41.30 34.50 42.90 27.40 365 8.00 8.00 8.00 8.00 8.00 27.80 35.90 28.70 33.90 25.10 366 8.00 8.00 8.00 6.00 7.00 19.80 35.30 32.80 29.80 17.70 367 8.00 8.00 8.00 9.00 9.00 20.90 27.60 21.80 22.50 20.80 368 8.00 8.00 8.00 8.00 4.00 6.90 12.80 7.60 12.50 5.40 369 8.00 8.00 8.00 9.00 9.50 42.70 59.90 76.60 67.70 66.20 370 8.00 8.00 8.00 8.00 9.00 11.30 16.50 12.50 14.30 14.90 371 8.00 8.00 8.00 8.00 8.00 25.90 48.30 35.70 41.10 33.10 372 8.00 8.00 22.20 41.90 373 8.00 8.00 8.00 7.00 8.00 16.20 25.90 20.80 17.80 14.80 374 8.00 8.00 6.00 6.00 18.30 28.80 14.10 11.30 375 8.00 8.00 6.00 35.60 58.40 8.50
70
Table 5.4 Continued (Fragments 376-400). Length and weight of the 8 mm fragment
class for 1 year.
8 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 376 8.00 31.80 377 8.00 8.00 8.00 6.00 9.00 35.10 320.23 43.70 44.70 33.60 378 8.00 8.00 8.00 9.00 8.00 33.80 44.20 57.40 57.00 48.00 379 8.00 8.00 8.00 8.00 7.00 15.50 21.00 17.00 21.20 19.70 380 8.00 8.00 8.00 9.00 24.70 33.70 29.30 35.10 381 8.00 8.00 8.00 20.60 34.00 23.90 382 8.00 8.00 8.00 8.00 32.60 44.50 50.30 35.40 383 8.00 8.00 8.00 8.00 8.00 30.70 39.70 33.80 37.90 20.80 384 8.00 8.00 8.00 6.00 5.00 25.10 34.80 39.60 37.10 11.50 385 8.00 8.00 8.00 9.00 10.00 40.30 49.80 41.90 45.30 46.60 386 8.00 8.00 8.00 8.00 8.00 43.00 67.30 18.00 35.80 19.50 387 8.00 8.00 8.00 8.00 33.30 56.10 45.40 23.40 388 8.00 8.00 8.00 7.00 35.70 39.40 16.00 22.00 389 8.00 8.00 8.00 8.00 9.00 30.80 47.30 31.00 29.30 24.50 390 8.00 8.00 8.00 29.10 320.60 18.40 391 8.00 8.00 8.00 8.00 8.50 19.70 31.60 23.70 19.60 23.30 392 8.00 8.00 8.00 8.00 8.00 38.80 55.90 32.00 33.90 33.70 393 8.00 8.00 8.00 7.00 6.00 20.60 25.30 16.30 15.60 8.30 394 8.00 8.00 8.00 10.00 10.00 48.20 61.70 49.10 40.00 56.10 395 8.00 8.00 8.00 9.00 8.00 29.40 41.10 37.10 38.90 31.20 396 8.00 8.00 8.00 8.00 8.00 33.30 34.00 37.30 41.50 34.40 397 8.00 8.00 8.00 9.50 9.00 42.10 46.80 40.30 38.10 23.50 398 8.00 8.00 8.00 36.20 58.40 3.70 399 8.00 8.00 8.00 8.00 7.00 33.10 46.80 48.40 42.50 22.10 400 8.00 8.00 8.00 6.00 4.00 30.40 56.00 33.80 29.40 7.80
71
Table 5.5 Fragment Growth (401-425). Length and weight of the 10 mm fragment
class for 1 year (continues on to next page).
10 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 401 10.00 10.00 10.00 10.00 11.00 66.10 90.50 106.70 85.00 66.10 402 10.00 10.00 10.00 10.00 12.00 57.50 73.90 79.60 61.10 50.00 403 10.00 10.00 10.00 10.00 10.00 50.20 77.00 100.80 76.40 76.90 404 10.00 10.00 10.00 10.00 11.00 45.40 66.50 92.00 71.20 39.80 405 10.00 10.00 10.00 10.00 11.00 69.40 79.70 96.20 84.70 89.70 406 10.00 10.00 10.00 50.15 82.90 22.80 407 10.00 10.00 10.00 10.00 12.00 55.70 92.40 85.20 67.80 81.10 408 10.00 10.00 10.00 10.00 12.00 37.80 48.40 45.80 70.00 55.00 409 10.00 10.00 10.00 10.00 10.00 46.60 56.10 54.10 50.80 56.60 410 10.00 10.00 10.00 10.00 11.00 56.40 72.70 52.80 92.60 81.10 411 10.00 10.00 10.00 10.00 10.00 51.00 68.80 66.50 47.90 66.00 412 10.00 10.00 10.00 10.00 11.50 55.30 74.50 79.00 95.20 93.80 413 10.00 10.00 10.00 10.00 12.00 35.50 51.40 55.80 45.20 61.80 414 10.00 10.00 10.00 10.00 11.50 33.30 56.10 65.00 75.00 69.80 415 10.00 10.00 10.00 10.00 11.00 62.00 90.90 94.90 77.70 67.60 416 10.00 10.00 10.00 10.00 12.00 65.10 106.90 91.20 104.40 108.30 417 10.00 10.00 10.00 10.00 11.00 62.00 89.00 93.20 73.80 82.20 418 10.00 10.00 10.00 10.00 11.00 47.50 66.30 93.80 71.00 54.80 419 10.00 10.00 10.00 10.00 11.00 87.70 108.80 109.70 74.00 70.50 420 10.00 10.00 10.00 10.00 56.90 83.00 117.50 99.70 421 10.00 10.00 10.00 10.00 11.00 27.70 30.90 34.70 32.90 35.70 422 10.00 10.00 10.00 10.00 35.30 43.80 48.70 50.30 423 10.00 10.00 10.00 10.00 10.00 40.00 47.90 70.00 48.90 63.00 424 10.00 10.00 10.00 10.00 10.00 67.70 57.70 93.50 101.60 98.50 425 10.00 10.00 10.00 10.00 10.00 60.70 84.60 85.30 72.40 59.20
72
Table 5.5 Continued (Fragments 426-450). Length and weight of the 10 mm
fragment class for 1 year (continues on to next page).
10 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 426 10.00 10.00 10.00 10.00 10.00 58.00 59.10 65.80 75.10 71.40 427 10.00 10.00 10.00 10.00 10.00 23.50 31.10 33.40 37.40 31.20 428 10.00 10.00 10.00 10.00 11.00 51.20 56.10 52.30 75.20 51.70 429 10.00 10.00 10.00 8.00 72.30 99.30 15.10 25.00 430 10.00 10.00 10.00 10.00 11.00 47.60 77.70 81.10 66.70 36.40 431 10.00 24.80 432 10.00 10.00 10.00 10.00 10.00 50.90 56.30 88.30 51.10 42.90 433 10.00 10.00 10.00 10.00 10.50 49.30 59.10 65.30 56.30 62.00 434 10.00 10.00 10.00 10.00 12.00 55.40 88.40 73.70 83.90 64.00 435 10.00 10.00 10.00 10.00 8.00 30.30 44.00 44.50 46.40 16.10 436 10.00 10.00 47.60 47.60 437 10.00 10.00 10.00 10.00 11.00 53.60 47.70 65.10 76.50 65.90 438 10.00 10.00 10.00 10.00 12.00 52.30 67.20 79.80 54.50 30.30 439 10.00 10.00 10.00 10.00 10.00 59.50 74.20 64.40 56.80 27.80 440 10.00 10.00 10.00 10.00 10.00 59.50 72.90 87.80 63.70 68.30 441 10.00 10.00 10.00 10.00 9.00 31.80 44.00 23.40 24.40 11.30 442 10.00 10.00 10.00 10.00 11.00 47.40 54.10 58.40 61.00 66.00 443 10.00 10.00 10.00 10.00 10.00 45.40 52.40 54.80 62.40 54.40 444 10.00 10.00 8.00 8.00 57.70 102.10 30.80 30.90 445 10.00 10.00 3.00 5.00 23.20 39.20 4.80 6.60 446 10.00 10.00 10.00 10.00 2.00 42.20 45.20 82.80 54.30 24.30 447 10.00 10.00 10.00 10.00 10.00 31.30 41.90 28.20 38.50 35.20 448 10.00 10.00 10.00 8.00 7.00 47.40 53.60 72.40 51.60 15.80 449 10.00 10.00 10.00 10.00 12.00 107.30 131.30 150.30 141.00 208.40 450 10.00 10.00 10.00 6.00 7.00 41.10 48.30 45.90 33.40 20.30
73
Table 5.5 Continued (Fragments 456-475). Length and weight of the 10 mm
fragment class for 1 year (continues on to next page).
10 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 451 10.00 10.00 10.00 10.00 10.00 45.80 56.50 51.20 48.30 22.30 452 10.00 10.00 10.00 10.00 12.00 39.20 60.60 64.00 58.30 72.60 453 10.00 10.00 10.00 10.00 10.00 55.20 62.40 85.00 66.90 91.20 454 10.00 44.90 455 10.00 10.00 10.00 10.00 11.00 66.50 87.30 90.50 72.10 96.40 456 10.00 10.00 10.00 10.00 12.00 52.70 57.60 68.80 76.20 74.10 457 10.00 10.00 43.50 61.50 458 10.00 10.00 10.00 10.00 11.00 61.00 84.10 113.30 104.20 101.40 459 10.00 10.00 10.00 10.00 10.00 22.20 12.70 37.30 23.50 26.10 460 10.00 10.00 8.00 10.00 73.70 66.00 34.20 36.50 461 10.00 10.00 10.00 10.00 11.00 47.70 46.60 63.60 74.30 65.10 462 10.00 10.00 10.00 10.00 11.00 45.60 56.90 87.00 35.60 108.80 463 10.00 10.00 10.00 10.00 10.00 28.90 57.70 45.00 46.60 42.70 464 10.00 10.00 31.70 35.10 465 10.00 10.00 10.00 10.00 38.60 43.20 32.80 28.10 466 10.00 10.00 3.00 32.80 41.10 2.20 467 10.00 10.00 10.00 4.00 42.90 36.90 58.20 0.10 468 10.00 10.00 10.00 10.00 5.00 56.60 71.20 83.30 52.60 17.00 469 10.00 10.00 10.00 10.00 5.00 57.50 63.60 67.30 67.30 92.80 470 10.00 10.00 10.00 10.00 11.00 54.60 42.50 37.60 60.90 38.30 471 10.00 10.00 10.00 10.00 8.00 70.10 68.50 71.41 50.50 33.80 472 10.00 10.00 10.00 10.00 7.00 31.00 58.10 62.50 71.00 56.40 473 10.00 10.00 8.00 10.00 11.00 66.70 60.10 24.30 28.50 70.80 474 10.00 10.00 10.00 10.00 71.20 127.90 99.60 104.50 475 10.00 10.00 10.00 10.00 11.00 79.10 125.00 104.60 102.00 91.70
74
Table 5.5 Continued (Fragments 476-500). Length and weight of the 10 mm
fragment class for 1 year.
10 mm Fragment Growth Thalli Length in mm Thalli Weight in mg ID Time Week Week Week Week Time Week Week Week Week # 0 13 26 39 52 0 13 26 39 52 476 10.00 10.00 10.00 10.00 11.00 61.60 76.50 102.90 124.30 105.50 477 10.00 10.00 10.00 10.00 11.00 47.60 65.20 62.80 72.30 134.30 478 10.00 10.00 10.00 10.00 10.00 44.40 77.80 59.90 74.20 50.90 479 10.00 10.00 10.00 10.00 10.00 30.00 36.50 40.70 38.10 69.80 480 10.00 10.00 42.60 321.50 481 10.00 10.00 10.00 10.00 10.00 50.60 77.30 82.00 53.40 32.10 482 10.00 10.00 10.00 10.00 11.00 44.30 47.00 42.70 42.70 41.90 483 10.00 10.00 10.00 10.00 6.00 72.00 94.20 67.20 49.70 17.30 484 10.00 33.20 485 10.00 10.00 10.00 10.00 6.00 38.00 42.80 49.50 40.40 11.30 486 10.00 10.00 10.00 10.00 11.00 46.60 65.80 61.90 58.00 56.60 487 10.00 10.00 10.00 10.00 11.00 36.10 54.10 55.15 56.20 19.20 488 10.00 10.00 10.00 10.00 11.00 57.60 80.90 77.20 57.30 45.60 489 10.00 10.00 10.00 10.00 11.00 60.70 77.20 93.50 89.60 87.20 490 10.00 10.00 41.40 47.10 491 10.00 10.00 10.00 10.00 10.00 31.70 40.70 47.70 50.90 67.00 492 10.00 10.00 10.00 10.00 9.00 35.00 58.40 65.70 92.20 37.00 493 10.00 10.00 29.00 49.10 494 10.00 10.00 10.00 12.00 11.00 64.80 98.80 151.40 121.60 96.30 495 10.00 10.00 55.60 97.40 496 10.00 67.00 497 10.00 10.00 9.00 38.20 50.50 6.50 498 10.00 55.30 499 10.00 49.40 500 10.00 47.90
75
Table 5.6 Chi-square Values. Using a probability value of .05 and a degree of freedom value of 1 requires a Chi square value of 3.841 or greater to be statistically
significant.
Chi-square Size Size value 10 mm 8 mm 0.174 8 mm 6 mm 15.14 6 mm 4 mm 0.022 4 mm 2 mm 2.62
76
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