Methods and Mechanisms of Pinus sabiniana Colonization As Observed Within the Context of the Bella Vista Study Range

by

Greg Mushial Barbara Pozek

17 Dec 2013

Summary

Although gravity is important in the local dissemination and colonization of Digger Pines ( Pinus sabiniana ); the Western Scrub Jay ( Aphelocoma californica) in a mutualistic relationship is the vector for more important non-local dissemination and colonization. This is a process which works as long as water is available for the Jays to inhabit the Digger Pine's domain. But in the absence of water, this mechanism fails, propagation is constrained and wider-spread colonization is delayed.

1

Figure 1 – Pinus sabiniana in typical grassland setting

“The tree is remarkable for its airy, widespread tropical appearance, which suggest a region of palms rather than cool pine woods. The sunbeams sift through even the leafiest trees with scarcely any interruption, and the weary traveler finds little protection in their shade.” − John Muir (1894)

Introduction

Pinus sabiniana, commonly known as: Bull Pine, California Foothill Pine, Digger Pine, Foothills Pine,

Ghost Pine, Gray Pine, Grayleaf Pine, Grey Pine, Nut Pine, Pineapple Pine, Sabine Pine, Smoky Pine,

Squaw Pine - “Little is known of the ecological life history of this species, even though it may be the most rare of the North American pines” (Barbour, 2007).

2 If one finds a single Digger Pine, one will typically find more than one Digger Pine in that area.

The purpose of this study is to try to discern the method of migration/propagation/colonization of the species – how the seeds from existing trees move/are moved, to new locations for possible germination and possible growth. For some tree species, the seeds are quite small (millimeters in size, milligrams in mass 1), and one can easily imagine the seeds being distributed by the wind, or a bird of almost any minimal size. Conversely, the seeds of the Digger Pine are much larger 2, large enough that the carriage of which would be limited as to which species might provide such service. In this study we try to discern if propagation/dissemination/colonization is via gravity, wind blow, or animal carriage.

Pinus sabiniana – The Species

The Digger Pine is one of the small collection of dry climate pines; a gymnosperm, a member of the family Pinaceae and native to United States (though since transplanted to the Adelaide area of

Australia). With coarse bark, a candelabra form, with a sparse adornment of characteristic long (12-

20cm) pale gray-green needles, three per fascicle; with large subglobose pitch heavy down-hanging cones (up to 25cm and up to 700g), nominally in groups of 3, which may be retained for multiple years

(3-5), and containing large edible seeds - an important food for the native indian population

(Brockman, 2012; Farris, 1982). The wood is light, soft and brittle, which is no longer used commercially, but was heavily used as mine shoring during the gold mining era of the 1850-60's

(Powers, 2009). The resin can be distilled to produce the oil abietine: an insecticide and cleaning agent

(Brockman, 2012). The Digger is a slow growing tree, may live 200 years, and nominally will reach

20-25m. Record Diggers have been found to reach 57m (Callahan, 2009) - the tallest found for this study measured 42.6m (a near record height for the modern era). Given its dry habitat, it will grow both

1 Eg, Sitka Spruce ( Picea sitchensis ) 2mg per seed, Western Red Cedar ( Thuja Plicata ) 1mg per seed (Hak, 2009). 2 Just over 600mg on average from this study.

3 a deep tap root, and wide reaching surface roots. Its active growth period is Spring and Summer; blooms in early Spring (Sagebud, 2013). The tree is not cold hardy and temperatures below -12C will kill it (GardenGuides, 1997).

Pinus sabiniana – The Habitat

A drought tolerant, pioneer species, possibly a climax species (in the context of poor soils). It is native to California and Oregon. A member of the Foothill Woodland or Northern Oak Woodland communities, with an indifference for serpentine soils, no salt tolerance, though moderate CaCO 3 tolerance (GardenGuides, 1997). It can be found thriving in all 5 orders of soil. It is found in all

California counties but 11; found at elevations from 20m through 2200m (Calfloria, 2013) – which is primarily constrained by minimum winter temperatures; and is quite shade intolerant (Brockman,

2012). Its drought tolerance and adaptation to near-xeric environments allows it to out-compete most other trees. The Digger can survive on 35cm of precipitation per year, or less (some of the oldest in our study area started growing during a period (1920s) where 20cm had been the norm for two decades).

The Digger is tolerant of greater rainfalls – again, those of the study area have survived multiple years of more than 2m of precipitation. But their high mass to root ratio makes them prone to fall or being blown over when the soil becomes saturated – a hillside location facilitates the drainage of the root area mitigating this to varying degrees (the biggest and oldest Diggers in our study were situated such). Few tree species are viable over such large temperature or precipitation ranges (Powers, 2009). Given their small sail area to mass ratio, they quite easily survive 110-120kmhr winds (assuming non-compromised roots). The Digger is frequently found in association with Blue Oaks ( Quercus douglasii) and to a lesser degree Valley Oaks ( Quercus lobata) .

4 Pinus sabiniana, or Pinus sabineana

Davis Douglas (of Douglas Fir fame) spent the years 1831-32 cataloging the plants of coastal North

America (Bird, 2010). This expedition was sponsored by Sir Edward Sabine, Member of the Royal

Society, past president thereof, and honorary secretary of the Royal Horticultural Society. Douglas found, described and published descriptions of Pinus sabiniana in 1833, named in honor and appreciation of his sponsor Edward Sabine. From then until 2001 that was the name used. In 2001

Aljos Farjon, Oxford University, maintainer of World Checklist and Bibliography of Conifers , changed the listing for the Digger Pine, to “ Pinus sabineana, Pinus sabiniana,” arguing first that the Douglas' original notes had been transcribed in error, and then that -iana was an improper suffix given Sabine's name (ending in a vowel). Many online sites followed the change in naming. And until it was pointed out that Article 60C.2 of the International Code of Botanical Nomenclature (ICBN) disagreed with this view, this change was retained. Of note, Farjon's 2008 update had returned to the -iana spelling, and had dropped the -eana alternative (Callahan, 2009).

Respectfully, Gray Pine, not Digger Pine?

The most commonly used common name for Pinus sabiniana is Digger Pine. One finds this in both scientific literature and non-scientific (popular) literature. Gray Pine is the second most commonly used name, but Digger probably represents 70% plus of the citations. One even finds it in Australian literature (the pines now grow there also). But in two of the website in researching this paper – out of forty to fifty - comments are made that the name Digger Pine is a pejorative, derogatory name, insulting to the Native Americans which used parts of the tree for food or tools. Callahan includes the passage: “'Digger' is a pejorative term used by settlers to collectively describe the native people of central California, who dug roots and bulbs and used the seeds, sap, cones, branches, leaves and bark of the pine. The native people resented the European settlers' name for them... “ (Callahan, 2009).

5 Callahan also includes the passage: “because the term 'Digger' is comparable in disrespect for Native

Americans to the similar sounding term for African Americans, and dendrology is under no obligation to prolong historic insults...” (Callahan, 2009). Labiste in her website on Native American use of seeds of the tree, includes the comment: “and the ethnically offensive common name of Digger Pine”

(Labiste, 2013). One is left wondering if this is political correctness run rampant, or is actually a name that should be avoided. The present author amongst his friends counts one that claims lineage from the

Modocs, and when he was asked about this, said that “he didn't care what I called the tree” (personal communications). The above is written just as a heads up, and noting that this might be a point to consider, or might be one to ignore.

6 Hypotheses

We approached this study with the intent of understanding the Digger Pine propagation mechanisms and not with the approach of testing a single hypothesis. As such we started with a group of hypotheses, with the hope of eliminating those which didn't match the data, assigning importance to those which did, and as we progressed being quite willing to add additional hypotheses as indicated by the data. At the end of the study this was our list of hypotheses:

1. Digger seeds are dropped from the cones while still attached to the trees, germinate where they

fall, and that's where seedlings would be found. This implies a generational dispersion limited

to the radius of the canopy. This also means that cones found at the base of the trees should be

seed-bare.

2. Per above, the seeds drop, maybe the wind blown them some distance from a parent tree. The

strong winds of the study area blow from two directions (the North, and North-West) – in the

opposite directions there should be lopsided seedling pattern about the base of the parent.

3. The cones from the Diggers, fall still full of seeds, upon impact with the ground, will eject the

seeds, and seedlings should be found associated with decaying cones. Seedlings should also be

found in clusters around these cones.

4. The cones from the diggers fall, roll down-gradient, disgorge their seeds there. This should lead

to a lopsided growth of seedlings down-gradient and clustered around decaying cones.

5. The cones fall, naturally or are eaten off by squirrels, the seeds are removed and stored. One

should find cones which have chew marks at the attachment and likewise chewed cones.

6. The cones open while still attached, birds extract the exposed seeds, lose some while trying to

extract the meat, which germinate where they fall. This should lead to seedlings located under

safe bird perching locations, and potentially well away from the parent trees.

7 Study Area

The area for this study is the 320 acres of oak grasslands owned by Shasta Community College of

Redding California. The land thereof other than two preexisting one-lane unmaintained dirt roads, the construction of a half-acre pond, border fencing, and being lightly grazed until 2001, has been kept in its native state. The land has been subjected to fires over the decades and as such has gone through natural regenerative successions. During such fires the understory has burned, most of the larger trees have been scorched, some were killed.

The study area is oak savanna and part of the foothill oak woodlands which rings the Central

Valley of California. The predominate trees are: Blue Oak ( Quercus douglasii) and Digger Pine. Also found there to a lesser degree are: Valley Oak ( Quercus lobata) and Coulter Pine ( Pinus coulten ). The understory is predominately grasses; or where the grasslands turn to chaparral: Chamise ( Adenostoma fasciculatum) , Manzanita ( Arctostaphylos spp.), Yerba Santa ( Eriodictyon californicum) and Buckbrush

(Ceanothus cuneatur) . Figure 2 below is representative of the grassland biome of the study area; figure

3 of the chaparral biome. Figure 2 is representative of the proportioning of Oaks and Digger Pines.

Geographically, the study area is the northern half of section 27, T33N R3W. The northwest corner of the area is at 40° 41.669'N, 122° 12.149'W; the northeast corner is at 40° 41.676'N 122°

11.008'W; the southwest corner at 40° 41.221'N 122° 12.145'W; and the southeast corner 40° 41.224'N

122° 11.002'W. The area is roughly rectangular. An overview of the study area and the data points are given in figure 4 (giving a larger context view), and figure 5 (showing just the study area) – the four corners of the area are denoted by the yellow Google Earth “push pins”. Figure 6 below is a subset of the USGS 7.5' 1998 topographic map covering the northern half of section 27. Figure 7 shows the locations of the study sites for this research - “1” through “5” denoting Data Trees 1 through 5, ES the

East-Slope site.

8 Water is scarce in the study area. Two creeks run though the study area: Bacon Creek and

Willow Creek – both are seasonal. From Google Earth historic imagery: both flow after the first sustained heavy rains of the season – as early as early November some years, others not until January; and both are typically dry by, as early as April and no later than June. For the birds there is water marginally available via the irrigation at the house to the northwest of the study area – meaning a quarter mile flight from the western part of the area. Water is available to the extreme eastern part of the study area via the two residences, one to the immediate east of the study area and one just south of the southeast corner – requiring a 400-500m flight. Water in the central and east-central part would require 1000-1500m flights.

9 Figure 2 – Typical grasslands with interspersed trees at study site

Figure 3 – Typical chaparral at study site

10 Figure 4 – Large area Google Earth overview of study area with area corners denoted

Figure 5 – Google Earth image of study area with corners denoted

11 Figure 6 – USGS, 1998, 1:24000 topographical map of study area

Figure 7 – Map denoting approximate study site locations

12

Methods / Materials

Digger Pine Site Selection – isolated Digger Pines, a significant distance from any other Diggers (50m at least, though because of the number and density of Diggers in the area, rarely is more than 150m possible); on grassland, with no immediate surrounding/interfering chaparral [at the base of the tree, or over the area where the cones might be expected to fall]; either on negligible slope, or steep slope

(depending what was sought), were initially identified via Google Earth. Google Earth being a color based system allows the user to readily distinguish Digger Pines from the rest given their blue-gray foliage color. Once a candidate tree/site was identified, its longitude/latitude were recorded and then with a hand-held GPS was field checked for viability. This study would have been much more difficult without Google Earth. Google Earth was also used to identify paths/routes/game trails that could be used to reach a site.

Hand-held GPS Unit – A Garmin 60CSx was used extensively to find sites previously identified via

Google Earth: using the Long/Lat coordinates from one, and walking/bushwhacking with the Garmin in hand until the matching coordinates were found in the field. The Garmin operates in degree/minute/fraction of minute mode – Google Earth was configured to also display locations in that format. Of note: both display locations to the thousandth of an arc-minute, and in no case was there ever even a 0.001 arc minute difference seen between the Google Earth coordinates and the Garmin display values. This means that trees only 6 feet apart will have unique coordinates. Standard Duracell batteries were used in the Garmin, were used for over 40 hours, and the battery indicator never showed less than full.

13 Measuring Distances – A Harbor Freight 100 meter fiberglass backed nylon reel tape was used to measure in-the-field distances. Distances of less then 30 cm were measured to plus/minus 1 cm.

Distances greater than 30 cm but less than approximately 1 meter were measured to plus/minus 2 cm.

Distances greater than 1m through 4 or 5 meters were measured to 5 cm; distances out to 50 m were measured to 10 cm; distances beyond that to the 20-50 cm level. All measured distances are generally accurate to 1%; but absolutely better than 2%. In the case of after-the-fact estimates of distances to nearest Digger, or distances between sites, Google Earth ruler tool was used – which reads to the 10 cm resolution; but given the problems of absolutely identifying locations, those distances should be taken as 50cm resolution values.

In the case of measuring the distances from a site Digger to a cone or seedling/sapling: those distances are measured from the central vertical axis of the Digger, to the center of mass of the cone or the center of the stem of the seedling.

Azimuth Angles - cone and seedling/sapling locations relative to the site subject Digger, were specified in terms of distance from the central axis of that tree, and the azimuth angle measured around the tree (measured in “hours” and “minutes”). By definition, the “12 o'clock” direction at any site is taken as “up gradient” as measured at the Digger. To facilitate the accuracy of the azimuth angles, either lines were scribed in the ground around the base of the tree; or in the case of a single person taking data, “duct tape” stripes were temporarily attached to the tree so that even at the extent of a measurement, the stripes could be seen and identified. Comparing in-the-field recorded azimuth angles to images in Google Earth, they appear to be accurate to at least plus/minus ten “clock” minutes (and in many case better than half that) [ten “clock” minutes is equivalent to 5 degrees of a 360 degree circle].

[Of note azimuth angles generally are measured in degrees, clockwise, from North; but historically these right-displacements (from the Arabic origins of the term) have also been measured in radians

14 (mathematics), mils, grads (military), or hours/minutes (astronomy) – we used the astronomy usage for ease of field use.]

In the case of a single person taking data, a wire was tied around the tree loosely, the end of the

100m tape attached to that wire, and by pulling it taunt it would rotate around the wire clearly identifying the azimuth angle. Since the end of the tape as not being held at the central axis of the tree, the radius of the tree plus the distance the tape was pulled beyond the bark were added to the measured distances. This correction has an accuracy of 5 cm or better.

Slope Gradients – are represented in “1 in n” terms, and for two reasons: 1) such values are more

“visualizable” by most readers [1 in 38 is simply 1 foot rise in 38 feet of run; 1.51°, actually the same value, is much less so]; and 2) so as to not to introduce apparent significant digits in the data not present in the actual measurements [to distinguish 1:50 and 1:51 requires 5 significant digits, where the actual measurements are only with 2 significant digits].

Google – was used to locate reference material for this paper, no specialized databases were used to find such. Google Earth was used extensively to locate possible study sites, and after the fact to make

“to nearest” Digger measurements.

Birds – were identified visually either by eye, with 8x binoculars, or by song. Distances if visual, were estimated from working knowledge acquired of the study site over time; if by song but with a visual sighting, were roughly estimated by volume, attenuation/muting of the high-frequency components, and by the lateral movement of the source (matching that to the likely bird flight velocity). Visual sighting distances are probably accurate to 5-10%; by sound, +/- 25%.

15 Mammals – other than two squirrels visually identified, the presence of such were deduced by cone damage, or droppings.

Cone Scale Counting – since Digger cones have nearly 100 scales up through 140 scales, while counting them, to ensure that all were counted, and none were counted twice, each as counted was marked with a red stripe from a felt pen.

Seed Collection – Over three data collecting trips the seeds were collected from approximately a total of 15 cones. The majority of the seeds (approx 80%) were collected from 8 to 10 cones – these were cones with 80% plus of the seeds still present. All seeds were collected from what we've defined as

“first-year” cones. The desire was to collect fresh, hopefully viable seeds. Occasionally older seeds were found, but they were not added to the collection. The total collection of seeds weighs 378 grams – or approximately 625 seeds.

Seed Volume – computed by water displacement method. A 25 mL graduated cylinder was filled with

15 mL of soft tap water, 10 randomly selected seeds from the above collection were added to the cylinder, were tamped down into the bottom of the cylinder so that they would stay submerged, and the level of the top of the water was then read. The volume of the seeds was taken as the difference between the resulting volume value, and the initial 15 mL value. The glass rod used to tamp the seeds into the cylinder, after such was done, was tapped multiple times to return any drops of water adhering to it to the measuring cylinder. The average volume of each seed was taken as one-tenth of the combined volume of the ten seed lots (fig. 31). Of note, the tap water used, unlike distilled water had an approximate density of 1.000030 g/cm 3, or approx 30 ppm greater than distilled water, meaning that all the measurements/calculations are low by that amount – but since that deviation is well beyond the

16 significant digits reported, this discrepancy will be ignored.

Seed Masses – Before the volume of each group of 10 seeds was measured, the 10 seeds were weighed on a Ohaus 0.01g resolution electronic scale. The average mass of each seed was taken as being one tenth the combined mass of the ten seeds. The seeds were weighed before their volume was measured, so that they were dry to the ambient air RH level. The scale was zeroed before each group was weighed.

Cones per Square Meter – a 1 by 1 meter square was laid out using the 100m tape used for other measurements. The cones within the area were counted. If a cone straddled the area boundary, if its center of mass was within the area, it was counted as being in; if the center of mass was outside of the area, then it was not counted. Only discernible cones were counted – this is to say cones up through 6,

7 or 8 years old. Cones the presence was only hinted at, were not counted. If what was left of a cone could still be picked up, it was counted.

Cone Age Estimates – after thirty some field hours collecting data, the Digger cones one found on the ground appeared to fall into distinct groups in terms of appearance and state of decay. We have interpreted this as being indicative of the number of years the cones have been on the ground. Since we have no “proof” that these estimates are correct, we are using/offering the following images as representatives of what we've referred to as one year through five year old ground cones. Cones beyond these ages are quite often found, but they are typically found in advanced stages of decay – where one finds parts of a cone, where they come apart merely by picking them up.

17

One year cones – are still brown in color, the sap is still sticky, there is no sign of decomposition and the claws/hooks at end of scales are still sharp (fig. 8).

Figure 8 - Example of “1 year old” cone

18

Two year cones – no longer overtly brown in color, sap possibly still present, but not sticky, developed a gray color, still structurally firm, sound in structure, and the scale ends have lost some of their sharpness (fig. 9).

Figure 9 - Example of “2 year old” cone

19

Three year cones – gray in color, sap missing, though color hints of where it might have been are present, structurally no longer tight as in a two year cone, decay is setting in (fig. 10).

Figure 10 - Example of “3 year old” cone

20

Four year cone – gray taking on a whitish over-color, scales loose, fungus beginning to colonize cone, will have been on the ground long enough that dirt will be adhering to it (fig. 11).

Figure 11 - Example of “4 year old” cone

21

Five year cone – shiny gray has been replaced by a dirt brown, scales are loose, some may be missing, has lost structural integrity (if one steps on one they disintegrate), likely much dirt adhering to them

(fig. 12).

Figure 12 - Example of “5 year old” cone

22 Seeds per Cone – This is a difficult number to establish with certainty. When a cone falls to the ground some are always lost upon impact – which means counting the seeds within a cone found on the ground always produces an under-count. Within a cone, some scales will hold a single seed, some will hold two (figs. 29, 30). This generally can be deduced by careful inspection of the scales and looking for a single or double depression – getting this number perfect is more an art than science. Probably the only foolproof way to count seeds per cone is to remove a cone from a tree while it is still attached, and before it has been attacked by any Blue Jays or Woodpeckers. Short of that one can count the seeds in full cones on the ground, and use those counts as lower approximations. Beyond that, two references mention 135 and 143 (Eberle, 1919) as the number of seeds per cone. Farris (1982) presents data from

26 cones with an average of 113 seeds per cone. These numbers do not conflict with what we saw, though we can not prove or disprove them.

Diagrammatic Representations of Data Sites – These diagrams were drawn by a program written specially for this project. The input file was a standard Excel .csv file. Canopy drip lines are drawn to scale. Height of seedlings/saplings are drawn to scale. The orientation of the drawings are always with the top of the image being in the direction of the up gradient, as observed at the site data tree. The azimuthal directions are always with “12 o'clock” being toward the top of the image, 3 o'clock toward the right edge. The scale and direction a geographic north are given at the bottom right of each image.

Diagrammatic site representations only include items for which data was taken, and not other objects within the context; as such, Google Earth imagery, with same scale and orientation is presented following the diagrammatic plots to provide a more accurate/complete context for the data items.

Scatter Plots – The scatter plots used in this paper (figs. 26-28) were drawn by a program specially written for this project. The input file format is based on the TopDrawer file format used by another

23 plotting program developed by one of the authors years ago at Stanford.

24 Data

This study started with a thought: if one sees a Digger Pine on top of a hill – how did it get there?

Digger Pine seeds are quite large – too large to be carried on the wind; and the cones are presumed to not possess some magical forces which would allow them to roll uphill. So, how does a Digger get there? A related motivating question was: given a stand of Diggers and a surrounding large un- colonized area – how fast could that area be colonized – how fast might the leading edge of that Digger stand “migrate” into the area?

When we started collecting data, we were hoping that the data would lead us along, ie, it would show us something indicative of how this dissemination/migration process worked. Initially we collected data with regards to about everything/anything that we thought might lead us in the direction of an understanding, an answer: the physical characteristics of the parent trees; the locations of the fallen cones and the condition thereof; the locations, apparent ages and heights of the seedlings/saplings; the degree of shade the saplings would have germinated under; the slope of the surrounding land, etc. For the first two trees we collected a couple hundred data points – but the data wasn't indicating anything obvious. Though a couple of the hypotheses were already looking to be not supported by the data. It wasn't until the third tree that the data started to reveal possibly the underlying processes. The first three trees were all on the western side/edge of the study area. For the 4 th , 5 th and

6th sites, just to make sure that what we were seeing wasn't unique to the west-side, we took data on the east-side. That did reveal more. Between these two parts of the data taking we took data across the middle of the site to see if there was a transition effect. The data from the sites is presented below. For each of these sites, a diagrammatic representation is given, and then an actual GoogleEarth satellite view of the site is given (with the same scale and orientation).

25 Data tree #1

Data for this Digger Pine was taken on 8 Oct 2013 (figures 13, 14). Data taken beyond that of the

Digger Pine itself, included data on location of cones which could be attributed as coming from it, also seedlings/saplings attributable to it. The location, probable age and presence of seeds was recorded for the 13 cones found there. Locations, height, whorl/knot counts and likely degree of shading at germination data was also taken for 40 seedlings/saplings. Using Google Earth “ruler” function, the nearest mature Digger Pine is 97.4m away, making the data tree well isolated.

If we knew where this study would lead us, in fact the first hints of the Scrub Jay symbiotic relationship was buried in this data: the 3 seedlings under the oak canopy at the bottom of the diagrammatic drawing were a hint; likewise the 4 seedlings under the oak up and to the right of the

Digger were a hint.

About 12m to the North of the data tree is the remains of a 1.6m girth Digger – which from its state of decay appears to have been down for 20 or so years. Given its size and proximity, it appears possible, maybe likely, the it was the source of seed from which the data tree grew (it can be made out in the Google Earth image.)

All the cones found on the ground, had been cleanly broken off, ie, no evidence of squirrels having chewed any cones down; nor any of squirrel predation after they were on the ground.

On the tree were found hanging 3 open cones; additionally 1 full sized but not yet open; and 5 immature cones. The canopy extended down to 3m above the ground.

26 Figure 13 - Diagrammatic view of Data Tree #1 Site

27 Figure 14 - Google Earth View of Data Tree #1 Site

28 Data Tree #2

This data was taken on 10 Oct 2013 (figures 15, 16). This tree is a couple hundred meters north of site

#1. It was picked, like site #1, for its isolation from other Digger: the nearest Digger, though younger is

40m to the north; the nearest equal age or older is 70m to the east-south-east; and its suspected parent tree over 130m to the northeast. The gradient at tree 2 is flatter than the gradient at Data Site #1, with a gradient of approx 1:22 – something that one wouldn't expect cones to roll down.

Unlike tree #1, this tree provided significantly more data: 82 cones and 41 saplings. Although this tree was not that far from tree #1, and the soil and environment appeared the same, the difference

(the number of cones and seedlings/saplings) is significant. At the time we wondered if the increased girth, indicative of a few greater years in age was responsible (from later data such could be unrelated).

Of the 82 cones cataloged, all were found within the drip-line, or drip-line plus a single bounce one might expect after falling 15m. Out of the 82 cones, 2 were chewed (by squirrel), and they were what we called 3 and 4 year old cones, respectively (ie, this predation took place some time ago). All cones appeared to have cleanly broken from the tree – none appeared to have been chewed off, ie, these cones appear to have been chewed on the ground, after they had fallen.

Unlike tree #1, most of the seedlings/saplings were clustered near or under the parent tree – 13 of 41 were within the drip-line. The furthest was only 22m away. Of the seedlings/saplings found within the drip-line, none appeared to be correlated to any cone.

On the tree we found 11 mature open cones hanging. Of those, 10 were clustered 10m up; the last at 7.4m off the ground.

29 Figure 15 - Diagrammatic View of Data Tree #2 Site

30 Figure 16 - Google Earth View of Data Tree #2 Site

31 Data Tree #3

This data was taken on 11 Oct 2013 (figures 17-19). This data-taking was the turning point in beginning to understand the dissemination process. Data tree #3 is a huge Digger – over 30m high, with a 20m x 15m drip-line, and although not cored, quite likely 80 to 100 years old. It is also likely the source of the seed for data tree #2 130m away. It is also likely the seed source which generated the line of Diggers which extends for a couple hundred meters to its east – 60-70 20m class trees. The tree was chosen for its size and for its relative isolation.

The ground under the tree was littered with cones – 20-25 per the half-dozen square meter plots counted. The cones on the ground fade into “ancient” history – there were recognizable 1, 2, 3 etc year old cones; but there were increasingly decomposed cones which are likely 10 plus years old. Walking under the tree meant literally walking on cones. The drip area under the tree is approx 235m 2, and within that area are likely between 4000 and 5000 cones. This number really precluded the counting and logging of them individually. Hanging on the tree, although difficult to get a total count, were at least 88 cones.

This is a very productive tree, and has been for some time. This tree has survived multiple fires: the bark is charred and split. Interestingly, the tree seems to have gone though a couple periods of producing offspring – probably related to the land being cleared by fire, and its seeds providing the stock to repopulate the area. Interestingly, currently, only 8 of its 38 associated seedlings/saplings are more than a meter tall; and the ground to the east of its drip-line is littered with 20-30cm seedlings (2 or

3 years old at most) – a bit surprising for such an old tree. None of the seedlings are actually within its drip-line.

It was while taking this data, that to the southwest, in the Oaks shown in extended site diagram

Jays (3 or 4) were heard “arguing.” This proved to be a seminal event in reaching the conclusions we did in this study.

32 Figure 17 - Diagrammatic View of Data Tree #3 Site

33 Figure 18 - Diagrammatic View of Extended Data Tree #3 Site

34 Figure 19 - Google Earth View of Extended Data Tree #3 Site

35 Data Tree #4

This data was taken 24 Oct 2013 (figures 20, 21). After taking data on the east-side of the study area, and seeming to come to the conclusions that: the Blue Jays were the primary carrier of seeds, but that they could only do such if there was readily available water, this data taking was to see if that notion held up.

Data tree #4 is the second largest and probably second oldest Digger of the “data trees.” The tree was selected because it was reasonably isolated (closest adult Digger just over 50m to the SSE), was fertile (lots of cones both on the tree (64), and on the ground (approx 400)), and it was near water

(70m to a pond, measuring 66m x 18m, or approx 930 m 2 surface area). What we found matched the model that had been developing with time, observations and data.

This tree had such a large collection of cones on the ground that we mapped only one quadrant about the tree – noting that the 4 quadrants had visually similar cone densities and distributions. In that

1 quadrant we mapped the locations of 99 cones. Around the tree, we mapped the locations of 47 seedlings and saplings. As one can see from the diagrammatic view of the site, all the cones were found within the dripline for the tree; yet all the seedlings/saplings but 1 (found in the open) were found not under the data tree, but under nearby oaks, making an even stronger case of what was seen at Data

Digger #3.

36 Figure 20 - Diagrammatic View of Data Tree #4 Site

37 Figure 21 - Google Earth View of Data Tree #4 Site

38 Data Tree #5

This data was taken on 25 Oct 2013 (figures 22, 23). It was the last data taken for this study. The goal here was to seek further confirmation of the model that the data up to this point was calling for; to see if the data tree being on a significant slope was important; and see if being closer to residences that could provide sufficient water (like in the NW corner of the study area) to allow the birds, especially the Jays, to act as agents of dissemination.

This tree is full sized, but not a huge Digger, and on a 1 in 3 slope. From the diagrammatic site drawing, one can see in fact the slope was sufficient to have some of the cones roll down gradient (this is the first of the Data Tree sites where the slope was finally sufficient to see this) – 23 of the 48 cones were found outside and down gradient of the drip-line of the tree.

This tree was also a woodpecker seed granary (fig. 33), with approx 200-250 seeds/holes visible from the ground. Like the granary in dh21 (fig. 34), zero Digger Pine seeds were found stored there – all visible seeds were oak acorns. The fact that approx 25% of the acorns had started to decay, this had been used as a granary for some time.

The suspicion that by being closer to two residences (one 180m to the east, one 250m to the south), such would allow for the habitation by Jays proved correct – unlike the other east-side sites,

Jays were heard in the local area while taking the data. Likewise, upon first arriving at the data tree, a pair of woodpeckers which had been perching near the top of the tree flew away.

As had become the norm, no seedlings were found within the drip-line of the tree; and all the seedlings (only 4) were found under perching spots in nearby oaks.

This data might have been unnecessary, but it was comforting in that it further buttressed what the previous data had been pushing us towards conclusion-wise.

39 Figure 22 - Diagrammatic View of Data Tree #5 Site

40 Figure 23 - Google Earth View of Data Tree #5 Site

41 “East-slope” Data Site

This data was taken 15 Oct 2013 (figures 24, 25). The previous data was taken along the western edge of the study area. The physiography of the east-side is quite different from the west. This data was taken to see if there was a difference between the two sides, and/or, if the east-side data would mirror the west-side data.

This data proved crucial to arriving at the final understanding of the dispersion or dissemination of seeds and the colonization of new areas. The east-side is not like the west-side. On the west-side a cone on the ground rarely was found intact with seeds. The west-side was rife with bird calls. On the east-side, we found cone after cone full of seeds. Likewise, we found the east-side with an eerie silence

– it was only after listening quietly for more than 5 minutes was a single bird (Titmouse) heard.

Even though this data site – on a 1 in 3 hillside – is immediately above a creek bed (Wilson

Creek), there is no water to be had here – the closest water is 1200m to the WNW, or 700m to the east.

The slope was steep enough that cones when they fell would roll until they were caught against fallen branches or logs. These catch points could have 20-50 cones all clustered together (in a 1m by

40cm space). Because of the steepness of the site, the number of cones, and the clustering of the cones, their location data was not taken.

More than 50% of the cones found here had seeds in them. Typically 1 year old cones had a full or nearly full (85%+) complement of seeds. Even 3 and 4 year old cones still had some (2-10%) of their seeds. This was entirely unlike the west-side sites (where seeds were found in 1-2% of the cones, and then only 1 or 2 seeds (of possible 140-150)).

The seedlings/saplings were clustered around their parent trees, but interestingly, up-gradient from them. We interpret this as down-wind scattering during extreme wind events (more on this in the discussion section) – probably ejected when the tree tops were being whipped back and forth by the winds.

42

Figure 24 - Diagrammatic View of “East-slope” Data Site

43

Figure 25 - Google Earth View of “East-slope” Site

44 Digger Seed Metrics

This data was taken to allow for the computation of a “ballistics coefficient” of the digger seeds (table

1). This value is important in determining the terminal velocity of such a falling seed; which is important in determining the time required for a Digger seed to fall to the ground from a particular height. Which all leads to answering the question: how far can a Digger seed be displaced laterally (the point of impact on the ground) relative to its point of release within the canopy). The second hypothesis

(from above) requires this value/data – and in fact in the Data Tree #5 site data, it is likely, but not proven, that some of the dispersion/shifting of the drip-line cone pattern is a result of wind-blow (but in cones in this case, but not seeds).

Measurement Group Mass – 10 Seed Group Volume – 10 Seed Group Specific Density Group 1 6.00g 6.6mL 0.9090 g/cm 3 2 6.57g 6.5mL 1.010 g/cm 3 3 5.97g 6.2mL 0.9629 g/cm 3 4 5.75g 6.0mL 0.9583 g/cm 3 5 5.78g 6.4mL 0.9031 g/cm 3 50 Seed Averages 0.6014g / seed 0.634ml / seed 0.9488 g/cm 3 Table 1 – Digger Pine Seed Metrics

Digger Pine Tree Height, Girth and Cone Counts

This data was taken as input to the tree fertility estimation: how a tree of age x, might have y many cones, each cone will on average have n seeds (table 2). If one sees z seedlings/saplings: what is the expected progeny rate per tree per year. If the Digger Pine method of dispersion was gravity or wind, then these values might be computable. Since the primary dissemination method turned out to be Scrub

Jays, this data proved unimportant in our conclusion, but we only learned that after the fact. The data was left here for others to make use of in generating girth, height, age generating functions. In taking

45 this data we yet again saw the transition from empty cones to full cones as we worked from the west- side toward the eastern-central part of the study area. A Google Earth image of the DH locations within the study area is given in Figure 34.

Tree ID Tree Tree Latitude Tree Height Tree Girth Cone Count Longitude dh1 40 41.628N 122 12.072W 19.9m 1.45m 8 dh2 40 41.629N 122 12.065W 26.0m 3.62m 7 dh3 40 41.632N 122 12.055W 16.7m 1.28m 2 dh4 40 41.631N 122 12.052W 14.9m 1.52m 13 dh5 40 41.622N 122 12.055W 18.45m 1.79m 19 dh6 40 41.625N 122 12.023W 24.3m 2.98m 31 dh7 40 41.583N 122 11.984W 11.8m 0.53m 1 dh8 40 41.559N 122 11.912W 21.1m 1.69m 33 dh9 40 41.562N 122 11.922W 18.6m 1.86m 32 dh10 40 41.540N 122 11.879W 19.9m 1.78m 29 dh11 40 41.545N 122 11.88W 17.5m 1.42m 10 dh12 40 41.528N 122 11.871W 22.2m 1.91m 67 dh13 40 41.540N 122 11.822W 8.9m 0.37m 2 dh14 40 41.541N 122 11.822W 7.1m 0.24m 0 dh16 40 41.514N 122 11.701W 13.6m 0.85m 0 dh17 40 41.482N 122 11.582W 26.3m 2.64m 21 dg18 40 41.455N 122 11.529W 26.2m 1.89m 22 dh19 40 41.458N 122 11.539W 24.8m 1.39m 0 dh20 40 41.437N 122 11.514W 21.8m 1.84m 58 dh21 40 41.333N 122 11.380W 25.9m 1.94m 17 dh22 40 41.344N 122 11.317W 26.0m 2.38m 23 dh23 40 41.352N 122 11.325W 42.6m 2.34m 23 dh24 40 41.363N 122 11.328W 21.95m 1.92m 27 dh25 40 41.370N 122 11.318W 33.1m 2.66m 11 dh26 40 41.372N 122 11.321W 38.9m 2.81m 53 Table 2 – Height / Girth / Cone Counts for 26 Sample Diggers

46

Cone Count – Height – Girth Data Represented as Scatter Plots

This data was taken hoping to find a correlation between tree “size” and (number of) cones (a surrogate for seed counts) – but no such correlation was found. In the following three plots (figures 26-28), each data point from Table 2 is plotted: cones as a function of stem girth; cones as a function of tree height; and (somewhat unrelated to this project, but interesting anyway), as tree height as a function of stem girth [which does show a noticeable correlation]. Each data point, instead of being plotted as a point, an

“x”, or some other symbol, is plotted as the data-tree number producing that point. The actual location on the graph for that data-point is the exact center of each data-tree number.

Figure 26 - Number of Open Cones on Diggers as Function of Stem Girth

47 Figure 27 - Number of Open Cones on Diggers as Function of Height

Figure 28 - Digger Heights as Function of Stem Girth

48 Discussion / Results

In seven trips to the study site, amounting to thirty plus hours of observations and data taking: we did not see a digger cone fall, we did not see a seed germinate, we did not see a seedling sprout, but given the exact location of 242 cones relative to their parent trees and the exact locations of 205 seedlings/saplings, not to mention the observations of another several thousand cone positions, we believe that the data argues pretty clearly what the methods and mechanisms of Digger Pine seed/seedling/sapling dispersion, dissemination and colonization are.

Hypothesis #1 – that cones fall, seeds fall out and the next generation grows there: Yes, this is one of the ways which Diggers propagate, but a low incidence pathway. Of the 205 seedlings/saplings we counted, only 18 were within the drip-lines of their parent trees: 2 in Site #1 (Fig. 13), 12 in Site #2

(Fig. 15), 2 in Site #3 (Fig. 17) and 2 at the East Slope site (Fig. 24). If one accepts all these seedlings/saplings were started this way, this would represent an 8.7% incidence, or about 1 in 12.

Though if one looks at the 2 for Site #3 - both are right at the edge of the drip-line, where one would expect a Jay to be trying to crack a seed (hidden in the needles at the branch ends). Likewise, if one looks at the 2 at the East Side site, they appear to simply be a part of a larger scattered pattern of seedlings/saplings about the tree bases – something that looks more like a North-wind driven scatter pattern. The ones at site #1 and #2 do look like what one might expect for gravity drop plantings. If one accepts that analysis, then that's 14 of 205 seedlings, or 6.8% incidence. Even if one accepts the larger number, this is still clearly not the primary method of propagation. Beyond that it's probably a suboptimal method: yes it means that seeds are (safely) located where the parent tree found viable and sufficient resources, but it also places the offspring in direct competition with the parent for those resources (and given that Diggers tend to grow in locations of constrained resources, such makes for potentially less than a win-win situation).

Hypothesis #2 - the seeds drop as the result of strong winds and drop in the direction of the

49 blowing wind. This looks like the dispersion seen at the East Side site (Fig. 24). But before one says the dispersion looks like such or not, one needs to compute ballistics coefficients of a nominal falling

Digger seed (as if it were a rifle bullet) and determine what a maximum likely range of dispersion would look like. By using the “bullet designer” function in RCBS.Load (written by one of the authors), and modeling a seed as a bullet with a diameter of 8mm, a length of 17mm, with both a 3 radius rounded “nose” and tail, a mass of 0.60g and a density of 0.95g/cm 3, one finds that the seed has a forward BC of 0.07 (which is basically unimportant in this case in that gravity is continually accelerating it during its fall), but crucially, with a specific lateral cross section. Given the above dimensions, and a 100km/h wind (a 1 sigma maximum likely value), and given a 30m drop (hence a

2.50 sec drop time), the maximum lateral displacement is 11.89m (which would represent a 11.3% coupling, which is a quite reasonable value given the low density of the “bullet”). If one looks at the seedling/sapling dispersion at the East Side site, and assume the 4 Diggers are at least 30m high (they are likely somewhat taller, but such was not explicitly measured), then all the seedlings/saplings fall within the expected dispersion given a NW wind. The other telling point here is: all the seedlings/saplings are uphill from their parent trees, ie, cones did not roll uphill to get then there.

Of note, although one does not see this skewing of the seedling/sapling pattern in the other sites, one does see a skewing of the cone locations to the south on sites #1 and #2.

Hypothesis #3 – the cones fall full of seeds, upon impact eject the seeds, engendering seedlings/saplings around the cones. On the East Side site, where one has agglomerations of cones, where they rolled downhill, but were stopped by fallen branches and stems, although one has gatherings 40, 50, 60 cones, one does not have a single seedling/sapling growing from such a cluster.

On Site #1 where one has 2 seedlings/saplings growing under the canopy and amongst the cones, one does not have a cone close to either. One might argue that for the larger sapling that the cone has already decayed, leaving no trace – the sapling only 30cm away from the parent tree and with a

50 height of 3m+, such might be possible in that it's a 8 to 10 year old tree (though we did find cones that were probably that old and not fully decayed). But the 26cm seedling, which at that height is only 2 or

3 years old, it is very unlikely that it came from a cone, a cone which has already decayed already, and that's why is no cone to be found.

On Site #2 one sees this same effect: small saplings (20-40cm tall), but no cones associated with them. At Site #3 there are only two saplings barely under the drip line, and yes since they are quite a bit taller and hence older (ca. 10 yrs), maybe they came from cones – though above, the argument is made that given their locations, they look more to be the product of Jays dropping seeds while trying to open them. Additionally at Site #3, there is a collection (22) of small saplings (20-50cm) to the east of the tree, none associated with a cone.

The other argument against this hypothesis might simply be: when cones were found with their seeds still present, they were exactly that, ie, the seeds were (still) in the cones, and not scattered on the ground around them. (This fact actually made it easy to collect seeds, in that one could simply pick up a cone and shake it into a bag and end up with the seeds).

Hypothesis #4 – the cones fall, full of seeds, roll downhill, drop their seeds there, resulting in clusters of seedlings about these locations. We didn't see such. We saw empty cones on the ground, but no associated seedlings; we saw full cones on the ground, again, with no associated seedlings/saplings.

The east-slope site is probably the best counter-example in that there were large collections of cones, and no seedlings; and conversely, where there were seedlings, and no cones – they were elsewhere.

Hypothesis #5 – “the squirrels did it.” The fact that we found no cones with bases that showed any indication of being chewed off; the fact that we saw only 2 squirrels in our 30 hours of taking data

(well away from any of our data sites): makes this seem unlikely. Likewise, the fact that we only found

2 cones in the west side data that had been squirrel chewed, and only 4 on from the east side data – and this is out of 400 plus cones cataloged and another couple thousand cones seen – the squirrel is not an

51 important factor in the dissemination.

One might add a corollary to this: the woodpeckers are responsible for the dissemination of the seeds. Again, the evidence runs the opposite. Yes we saw 2 woodpeckers at Data Tree #5 upon arrival, but the two granary trees that were observed (DH21 and Data Tree #5) – representing maybe 500 stored seeds, no (zero) Digger Pine seeds were observed, only oak acorns (figs. 32, 33). Maybe one can argue that the woodpeckers treat Digger seeds differently from acorns – they crack and eat Digger seeds, but store acorns – but one would have to explain why, especially when Digger seeds are so much tougher and more difficult to open. We can not rule this behavior out, but it seems unlikely.

Nature has likewise made the woodpecker an unlikely candidate to try to pierce seeds while perching on a limb: they are of the order piciformes – the climbing birds, and not passeriformes – the perching birds. The feet of a Jay are like those of all passeriformes : three toes forward, one rearward, and when they land on a branch, the toes automatically grasp the perch (Ehrlich, 1988; Pedone, 2001).

This gives them great stability in that position. The woodpecker is unstable and uncomfortable on a branch – it is actually more comfortable hanging upside down from a branch, or trying to hang from the side of one – neither place suitable for trying to crack or pierce a Digger seed. So, when one finds

Digger seeds beneath branch perches, and not near the tree stem, those seeds were much more likely left/lost by a bird of the order passeriformes , which the Scrub Jay is, than one from piciformes , like a woodpecker.

A variation on this is unfortunately common in the published literature, where digger pine seeds are left on the ground, and then based on which animals take the seeds away, they are attributed as being the disseminators of the Diggers, failing to acknowledge the reality of the cones being retained on the trees for 3, 4, possibly 7 years. Johnson (2003) is an example of this seemingly flawed analysis.

They also conclude that the wings attached to the seeds are important in dispersal of the seeds, without computing or appreciating the aero-characteristics of the seeds, or realizing that cones are commonly

52 found without seeds but with wings still in place. This is also the type of “research” which has made commonplace in the literature that the woodpecker is a disseminator of Digger seeds.

Hypothesis #6 – the Scrub Jays find Digger seeds in open cones still on the trees, carry away the seeds with the hope of cracking them and eating the meat, but lose them in the process: such might be the most important mechanism in dissemination of Digger seeds and the colonization of new areas.

Although we saw some seedlings/saplings around the bases of Diggers, these for two reasons are not a

“winning” strategy for the trees: 1) if it takes 20 years before a Digger becomes fertile, and the furthest it can project itself in that time is to the edge of its drip-line – that doesn't represent a significant extension of its domain nor a terribly efficient one; and 2) doing such, given that the Digger nominally is found in resource constrained locations (this is where and how it out-competes the other tree species), having the next generation competing for resources which the parent tree has pretty well monopolized/grown to the limit thereof, helps neither the parent, nor makes for the likely success of the next generation. For a Digger, to project its next generation maybe nearby, but far enough away from the utilized resource area of the parent tree, has to be a more successful/desirable strategy. And maybe over time for this reason/goal, a symbiotic relationship with Scrub (or maybe other) Jays has evolved: the Jay gets high quality food (in an area nominally with little), and the Digger gets its seeds disseminated nearby but still well enough away so as to not compete for what are limited resources.

The first point in making this argument is: that we did in fact see and hear Jays, where we did see this projection of seeds away from the parent trees; and likewise, did not see nor hear them where we didn't see this effect.

The second point of this argument: the Digger retains its cones after opening for several years before dropping them – Farris (1982) says 3 to 4 years, up to an extreme of 7 years. Again, where we saw the projection of the seeds, we found very few cones on the ground with any seeds in them; likewise and conversely, were we didn't see this effect, we found cones full of seeds on the ground.

53 (Tying this to point one: were we heard and saw Jays, we found empty cones on the ground, and projection of seeds; and were we didn't see or hear them, we didn't find the projection and empty cones

(we found full ones).)

A Digger has a very open canopy – an unsafe place for a Jay to try to crack freshly obtained seeds (even though the source might be close at hand): a Jay focused on the difficult task of extraction the meat, is a meal waiting to be had by the many Redtail Hawks ( Buteo jamaicensis ) we heard and saw. It would be much safer for the Jay to take the seed to a safer more sheltered/obscured/hidden place, one with a strong “anvil” against which to hammer the seed and try to open it there (also from where it might be dropped out of frustration in not being able to open it, or from where it might be

“launched' by a powerful but off axis hit). We saw these venues from Data Tree #1 site, but didn't recognize them until we'd taken data on Tree #3.

Two “events” which lead to proposing this hypothesis were: while taking the data for Tree #3, several Jays (4 or 5) were squawking back and forth in the Oaks at the “top left” of the extended Tree

#3 site diagram (Fig. 18) – which lead one of the authors after the data taking was complete to wonder over there to see if there was a reason for this noise; and 2) finding under the oak trees a collection of nearly 20 Digger seedlings/saplings – not what one would to first approximation expect to find under

Oaks. These events/observations suggested a new direction for this study, one which we did pursue productively.

An additional important observation is: when one looks at the locations of the seedlings under the oaks at site #3, one finds that at least half, are at the drip-lines of these trees; and equally importantly: with prominent branches overhead – branches which could easily be flown to (from outside of the tree), sturdy enough to be used as an anvil to try to break the shell of the Digger seeds against, and at the same time sufficiently sheltered that a Hawk wouldn't be able to easily surprise a Jay and make a meal of it.

54 These Oaks at Tree #3 were a starting point. But it turns out that along the road which leads down the west side of the study area, under at least half, maybe 75% of the Oakes, one also finds

Digger seedlings – something that we had walked by multiple times, but only noticed after finding the

Site #3 out-of-place seedlings.

What we believe is happening is: the Diggers open their cones, and display their seeds (over a period of 2, 3, 4, maybe 7 years), over which time they are found by Jays; which quickly fly in, extract a single seed, and then fly (out of harm's way) to a safe place whey they can try to harvest it. For anyone that has tried to crack a Digger seed, it is likely that the Jays fail most of the time at this, but given the size of the “prize” (the lure), they will continue trying with seed after seed, especially if they succeed at 1 in a dozen (meaning that 11 of the 12 seeds are dropped under their safe perches, positioned to potentially germinate and to the parent Diggers advantage, away from its resources; and maybe even competitively, under a “competitor's” canopy, possibly challenging its resources... and if such is true, a pretty clever strategy).

And this is why on the west side where we heard and saw the Jays, and where we found this projection effect, we rarely found cones with any seeds in them on the ground. But this raises the question: why did this work on the west side of the study area, and on the extreme east side, but not in the middle? We believe the answer is simply: water. Not water for the trees, since the entire study area gets approximately the same amount of precipitation over a winter, but water for the Jays. As noted above: on the west side a Jay can find water for its use (drinking water), within a 400m flight. Since it's over 100m from Digger Tree #3 to the surrogate parent Oaks, and Jays were heard all along the road to the northwest corner of the study area, the site of the water, it should be a safe assumption that a single

400m flight, or a 400m flight in multiple hops is reasonable. The bottom line being: that the west side was a hospitable domain for the Jays to live.

Which leads to the point of why this effect isn't seen in the east side: simply, as the west side

55 has water, the eastern central part (excluding the extreme east side), has no year round water. There are

2 creeks though that area, but they are seasonal, and for at least 6 months a year, they are dry, and quite possibly, for 8 or 9 months a year they are dry. Jays have territories. If a Jay can only make use of a site

4 months (plus/minus) a year, then they are disinclined to make use of it those 4 months. This why we believe, even though the Diggers there “presented” their seeds to/for the Jays, in this symbiotic relationship, the seeds weren't taken and disseminated; and why we found many 1 year old cones on the ground full of seeds, found a few 2 year cones on the ground with significant seeds, and even found 3 year old cones still with some seeds – simply, the Jays weren't there to harvest them.

The follow on to this is: in the extreme east side, where water becomes a 500-600m flight, the birds appear again, the woodpeckers, and the Jays. And why for Data Tree #5, even though 48 cones were found on the ground – representing a seed pool of 5000 or so seeds (over 5 years), the only 4 seedlings/saplings found, were all under Oaks – one nearby the Digger, but in quite a thicket, ie, safe from Hawk attacks; and the other three some 30 plus meters away, and under significant canopies.

The other extreme demonstration of this effect is Data Tree #4 – a site specifically selected to see if this effect was what we thought we were seeing; a short distance to a full year water source – a pond, and with a large number of nearby Oaks. Of the 47 seedlings/saplings at the site, only exactly 1, was not under one of these oaks, or more specifically at its drip-line. Data Tree #4 drives the point home: we cataloged 99 cones under 1 quadrant of the tree, and suggest that under the full drip-line of the tree, there were approximately 400 cones, yet not a single Digger seedling or sapling was found there; but under the adjacent 13 Oaks were found 46 (of the 47) seedlings/saplings.

Some may wish to argue that these seedlings/saplings were planted by squirrels. But we again would note: none of the cones there showed any signs of squirrel destruction; none showed any sign of being chewed off the trees; and why would a squirrel preferentially select beneath the oaks and specifically at the drip lines of the Oaks to plant/store the seeds? We believe that the data argues much

56 more strongly for the Jay model, and that the onus is upon those suggesting the squirrel model to make their case.

The other aspect of this study, was an attempt to calculate a likely projection rate – the rate that a Digger could colonize/project its type forward into unoccupied land, or even into contested land.

When we thought we were going to be able to do such, we hadn't encountered the Jay data / model yet.

We would argue now that to actually compute such a value, would prove to be difficult, or at least difficult to do with reasonable error bars on the computed values. We after the fact believe the variable over which one has no control and would be hard to determine, would be the distance a Jay would choose/be willing to fly, after collecting a seed to “attack” it, and actually doing such – a distance which has a lot to do with the local “safe place” topology, something which is going to vary from site to site greatly.

The final aspect of this study was the desire to suggest “effective fertility” rates for Diggers – this is to say, how many offspring Diggers might be found per unit time nearby the parent tree. The other possible way to express this would be; how many seedlings/saplings per unit area per year. We believe a lower limit on this would be illustrated by the East Slope site – where one does not have the help of the Jays, one sees projection distances of 10's of meters, and few offspring. There one sees rate/values on the order of 1 offspring per 1000 m 2 yr -1 [16 seedlings/saplings, over 5 years, over 3100 m2].

On Data Site #4, one would see a value like: 42.1 offspring per 1000 m 2 yr -1 [a 90m x 62m area

– 5580 m 2 , 5 years, and 46 seedlings/saplings (1 excluded for being clearly > 5 years old)]. On Data

Site #3 this value would look like: 5.35 offspring per 1000 m 2 yr -1 [140m x 140m area – 19600 m 2 , 5 years, and 21 seedlings/saplings] – but in this case, one must understand given the ambiguous boundaries of the site that this value comes with significant error bars.

57 Conclusions

Although Digger Pines do disseminate some of their seeds and colonize new but local areas via gravity and wind-blow (less than 10%), the most important mechanism to colonize more removed areas, is a symbiotic relationship with the Scrub Jay: the Jays carry the seeds from the still hanging cones, to a place where they can attempt to crack them (a strong branch as an anvil), but in the process of trying, given the toughness of the hulls, either give up and let them drop, or misstrike the exact ellipsoid center, converting the kinetic energy of the blow into transverse energy, launching the seed away from their perch and onto the ground below. Possibly some of these launched seeds are retrieved and further attempts are made to crack and eat them, but at least some are left, and over time a percentage germinate and become members of new colonies some distance removed from the parent tree.

Beyond such, the two animals presumed to be important in the dissemination of seeds and the projection of the species – squirrels and woodpeckers – is simply not supported by the data. Of the several hundred cones cataloged for this study, and more then again that many observed: only a half- dozen (well less than 1%) showed any evidence of squirrel seed predation – implying even smaller planting rates. The woodpeckers, although they use Diggers as granary trees, zero evidence was found of their interest in Digger seeds, while absolute evidence of their interest in and use of acorns was.

As such, the first conclusion is: Jays and the Diggers work together in a symbiotic relationship: to feed the Jays and to disseminate Digger seeds thus projecting the species.

The larger conclusion reached is: the Jays, like all living organisms, need water. The water does not need to be immediately present, but it does need to be nearby. For the data trees on the west side of the study area, in the northwest corner is a house with irrigation; and in the southwest corner a pond – collectively making water on average available within 400m or less. In this area we found compelling evidence of the Digger-Jay symbiosis working. Conversely, within the east-central area, where Bacon and Miller Creeks are dry (approximately 8 months a year), water is 1500m away or more, making that

58 area inviable for Jay habitation; as such, an area where the Jays are unable to fulfill their needed – from the Digger's perspective – role, of seed dissemination. Although gravity as a means of dissemination is at work in this area – given the incidences of seedlings under the canopies of existing trees, down- gradient from where cones might be expected to fall and roll to, or where they might be projected to via wind-blow - the further reaching Jay assisted colonization of the west-side is quite absent.

Conclusions: water supports Jay-Digger symbiosis, thus projection; conversely, no water, no

Jays, no symbiosis and thus no projection.

Work still to be done

− A common theme throughout the Digger literature is that the cones on the trees are

grown/presented periodically/cyclically, and with an interval/periodicity of 3 to 4 years. With

the data included but not fully utilized in Appendix A, it would be interesting if this cyclic

presentation can be seen, likewise a suggested periodicity.

− The DH data from 25 Diggers gives girth/DBH, height and cone counts. The trees should be

cored and actual age be determined. The Data from such might reveal some patterns in terms of

cone counts as a function of age. If not that, at least it would provide a reference for converting

DBH to age – and with 25 data points the error bars should be useable small.

− One could test the lack of water / lack of Jays observation and suggested causal relationship, by

making water available on the east side of the study area, over a decade or more. And see if one

didn't start to see seedlings start to appear well removed from the drip-lines of the existing trees.

− One could do it with the existing DH data, would be more robust if more DH data was added,

but there appears to be a strong correlation between stem girth and stem height – this needs to

be regression fit.

59 − On the west-side it was highly unusual to see cones with seeds; in the east-central area is was

the opposite. While we were taking the DH data (fig. 34) moving from the west side toward the

east side we noticed an effect: as we got closer to where we should expect cones with seeds, but

not there yet: if we found a cone heavily covered with sap, it would quite often have seeds. Is it

possible that the Jays that normally would have harvested the seeds: were afraid of heavy sap

coatings, afraid of getting stuck, or their feathers stuck? Or, maybe there was a (warning) odor

effect? This might be studied further.

60 Cited Material

Barbour, Michael G., Todd Keeler-Wolf, Alan A. Schoenhem. Terrestrial Vegetaion of California. Univ Cal Press: Berkeley. 2007. Print.

Bird, Jean. “Friends of the Waite Arboretum.” Waite Arboretum Newsletter 64, Winter 2010. 12Nov2013. http://waite.adelaide.edu.au/arboretum/pdf/newsletter_64_winter.pdf . Web

Brockman Tree Tour. “45. Digger Pine.” Brockman Memorial Tree Tour. Univ Washington. http://www.sefs.washington.edu/BrockmanTreeTour/45_dpine.html . 28 Nov 2013. Web.

Calflora. “Taxon Report 6524.” Calflora: Information on California Plants for Education, Research, Conservation. http://www.calflora.org/cgi-bin/species_query.cgi?where-calrecnum=6524 . 28 Nov 2013. Web

Callahan, Frank. “Discovering Gray Pine ( Pinus sabiniana) in Oregon”. Native Plant Society of Oregon, Vol 16, 2009. 20 Oct 2013. http://www.npsoregon.org/kalmiopsis/kalmiopsis16/callahan.pdf . Web.

Eberle, E. G., ed. “American Pharmaceutical Association, Yearbook, 1916, Volume 6.” American Pharmaceutical Association, Chicago IL, 1916. Print.

Erhlich, Paul R., David S. Dobkin, Darryl Wheye. “Feet.” Stanford University, 1988. http://www.stanford.edu/group/stanfordbirds/text/essays/Feet.html . 12 Dec 2013. Web.

Farris, Glenn J. “Pine Nuts as Aboriginal Food Source in California and Nevada: Some Contrasts.” Journal of Ethnobiology Vol 2, #2, pg 114:122. Dec 1982. Print.

GardenGuides.Com. “California Foothill Pine (Sabiniana).” Demand Media, 2009. http://www.gardenguides.com/taxonomy/california-foothill-pine-pinus-sabiniana/ . 28 Nov 2013. Web.

Hak, O., J. H. Russell. “Increasing Quality Seed Production in Western Redcedar Orchards.” Jun 2007. FGC Extension Note 09. Forest Genetics Council for British Columbia. http://www.fs.fed.us/pnw/olympia/kids/WASeeds.pdf . 5 Dec 2013. Web.

Hogan, Michael C. “Pinus Sabinians, Gray Pine.” EOL – Encyclopedia of Life, 2010. http://eol.org/pages/1033632/details . 28 Nov 2013. Web.

Johnson, Matthew et al. “A Comparative Analysis of Seed and Cone Characteristics and Seed-dispersal Strategies of Three Pines in the subsection Sabinianae. ” Plant Ecology, 168: 69-84, 2003. Web. 10 Oct 2013.

Labiste, Susan. “CALIFORNIA INDIANS, The Ohlone Peoples: Botanical, Animal and Mineral Resources.” 2013. http://www.primitiveways.com/Ohlone%20Peoples2.html . 22 Oct 2013. Web.

61 Pedone, Sam. “The Biogeography of the Acorn Woodpecker ( Melanerpas formicivorus ). San Francisco State Dept of Geography, 2001. http://online.sfsu.edu/bholzman/courses/Fall01%20projects/AcornW.htm . 12 Dec 2013. Web.

Powers, Robert F. “Digger Pine, Pinus Sabiniana.” AV High Desert Forum. 28 Nov 2013. http://www.avhidesert.com/forum/showthread.php?tid=889 , also http://na.fs.fed.us/spfo/pubs/silvics_manual/Volume_1/pinus/sabiniana.htm . Web.

Sabine, Sir Edward. http://en.wikipedia.org/wiki/Edward_Sabine . 20 Oct 2013. Web.

Sagebud. “California Foothill Pine (Pinus Sabiniana).” Sagebud – A Directory of Plants. Modscape LLC, 2008. http://sagebud.com/california-foothill-pine-pinus-sabiniana/ . Web. 28 Nov 2013.

62 Appendix A

Raw Cone, Sapling, Tree Data

The data for this project was gathered and kept in a spreadsheet, but for the purposes of this paper that proved counterproductive – the spreadsheet programs seemed to have a mind of their own (unfortunately exhibiting a very low IQ), and instead of helping, added to the difficulty of processing the data. The data was exported to a .csv format file, and hand edited from there. Agnr65Grf (a PC based program) was developed to generate diagrammatic representations of each study site and written to process the hand maintained .csv as its input. That .csv file is displayed below.

Copy of “ProjectData.CSV” file

site title, Digger Pine Site 1 lat, 40 41.298N long, 122 12.041W girth(m), 1.20 ConvertGirth height(m), 14.5 dripline3(m), 2.9 dripline6(m), 5.1 dripline9(m), 4.8 dripline12(m), 2.3 gradient, 6/55 north, 21deg w of 3oclock date, 8 Oct 2013 Cones * n, dist(m), angle, age(yr), seeds 1,2.7,2:30,2,0 2,1,11:30,1,0 3,1.1,9:30,2-3,0 4,1.5,9:00,3,0 5,2.5,9:45,2,0 6,3.1,8:45,2,0 7,3.9,8:45,2,0 8,3.1,8:00,3,0 9,2.6,7:45,3,0 10,3.3,7:45,3,0 11,3.8,7:00,2,0 12,4.3,7:10,3,0 13,4.2,8:10,3,0 Saplings * n, dist(m), angle, height(m), shade, whorls 1,2.4,5:15,.5,0/8,0 2,3.4,8:00,.26,1/3,0 3,3.9,8:15,.15,2/3,0

63 4,3.5,8:45,.16,7/8,0 5,6.7,11:00,1,1/4,2 6,8.3,11:00,1.6,7/8,4 7,12.5,10:45,3.3,6/8,7 8,12.8,12,1.9,1/4,6 9,9.2,9,1.7,3/4,5 10,12.9,8:50,2.2,3/4,5 11,16.0,12:45,1.9,3/4,5 12,15.8,12:45,.4,1/2,0 13,7.7,2:30,1.6,1/2,6 14,9.4,2,1,7/8,7 15,7.9,1:30,.56,8/8,2 16,8.8,1:30,1.05,8/8,3 17,8.4,1:30,.40,8/8,0 18,8.1,1:10,.7,8/8,1 19,17.4,2:30,3.5,7/8,7 20,18.2,2:50,1.9,7/8,7 21,17.1,3:10,9ft,3/4,8 22,14.3,4:10,.9,3/4,2 23,14.7,4:20,11ft,2/8,7 24,15.7,4:30,8ft,8/8,8 25,18.5,4:48,20ft,3/4,8 26,17.5,4:50,18ft,0,9 27,26.4,4:50,14ft,0 28,0.3,4:50,13ft,0,5 29,14.3,6,16.5ft,0,9 30,46,5:30,11ft,0,6 31,48.2,5:25,1.4,8/8,3 32,46.7,5:35,8ft,0,8 33,37.4,6,9ft,0,6 34,42.7,6:10,8ft,0,5 35,22.7,7:30,8ft,3/8,6 36,18.6,7,14ft,1/8,6 37,17.9,7:30,15.5ft,2/8,6 38,17.9,7:30,8ft,8/8,5 39,16.6,7:45,12ft,8/8,5 40,14.1,6,.26,0,0 othertree dist(m),10.5 angle,1:30 NSdrip(m), 8 EWdrip(m), 10 othertree dist(m), 51.2 angle, 5:30 NSdrip(m), 11 EWdrip(m), 12

64 site title, Digger Pine Site 2 lat, 40 41.410N long, 122 12.092W girth(m), 1.85 ConvertGirth height(m), 17.0 dripline3(m), 5.2 dripline6(m), 6.3 dripline9(m), 6.0 dripline12(m), 5.8 gradient, 1.6/35 north, 11deg w of 3oclock date, 10 Oct 2013 Cones * n dist angle yr seeds 1,2.6,12:00,2,few 2,3.1,12:00,2,few 3,4.7,12:50,3,few 4,1.9,11:50,3,no 5,1.1,11:50,4,no 6,1.1,11:45,1,no 7,3.5,11:00,2,no 8,3.2,11:00,2,few 9,2.1,11:00,3,no 10,2.0,11:00,1,no 11,0.7,11:00,5,no 12,2.1,10:00,1,no 13,2.8,10:30,2,no 14,2.9,10:20,1,no 15,3.3,10:20,4,no 16,4.2,10:00,2,no 17,5.6,10:10,3,no 18,5.0,10:00,5,no 19,6.8,10:00,2,no 20,3.9,9:30,3,no 21,3.9,9:25,3,1 22,.5,9:20,4,no 23,.5,9:20,4,no 24,.5,9:10,5,no 25,.7,9:00,5,no 26,6.3,9:30,2,no 27,6,9:10,3,no 28,5,9:00,2,no 29,4.7,9:00,2,no 30,3.6,8:30,5,no

65 31,3.6,8:20,2,no 32,3.3,8:30,2,no 33,3.3,8:20,3,no 34,3.8,8:15,4,no 35,3.8,8:15,4,no 36,4.5,8:10,2,no 37,1.1,8:00,4,chewed 38,1.3,8:00,3,chewed 39,2.2,7:50,3,no 40,5.7,7:50,3,1 41,6.6,7:50,2,no 42,6.4,7:40,2,no 43,7.0,7:30,3,no 44,5.5,7:30,2,1 45,5.5,7:30,2,no 46,6.1,7:20,4,no 47,5.0,7:20,4,no 48,7.3,0:00,3,no 49,6.7,7:00,2,1 50,5.9,6:50,2,no 51,5.1,6:55,2,1 52,4.7,6:50,2,no 53,4.9,7:00,2,1 54,1.6,6:45,2,no 55,2.3,7:30,2,no 56,1.9,7:15,5,no 57,6.8,6:15,2,no 58,5.1,6:00,2,no 59,3.4,6:00,2,no 60,3.3,6:00,4,no 61,3.7,5:45,1,no 62,5.0,5:30,3,1 63,3.7,5:30,2,1 64,4.7,5:00,4,15+ 65,2.8,5:00,2,no 66,2.6,5:05,5+,no 67,2.1,4:40,5+,no 68,2.1,4:30,5+,no 69,2.2,4:00,5++,no 70,4.0,4:00,2,no 71,4.0,3:40,2,1 72,3.3,2:20,1,no 73,3.0,3:00,2,no 74,3.4,3:00,2,no 75,3.6,1:45,4,no 76,4.7,1:00,3,2 77,4.8,12:45,1,no

66 78,5.8,12:20,3,no 79,4.6,12:20,2,1 80,2.1,12:20,3,no 81,2.1,12:15,3,no 82,2.1,12:10,2,no Saplings * n dist(m) angle height shade whorls 1,12,1:30,0.7,.125-1,2 2,18,3:00,0.85,0.25,2 3,4.1,3:50,0.65,0.7,1 4,4,4:30,0.8,1,2 5,3.5,4:35,0.3,1,0 6,2.8,4:30,0.3,0.875,0 7,22.3,4:30,0.55,0,2 8,9.9,5:10,0.2,0.3,0 9,10.4,6:10,2.1,0.3,5 10,5.4,6:00,0.27,0.5,0 11,8.4,6:45,1.2,0.8,4 12,12.5,6:50,1.4,0.6,4 13,10.3,7:00,0.6,0.9,1 14,10.2,6:55,0.1,0.9,0 15,2,6:45,0.21,0.3,0 16,2.7,7:30,1.25,0.3,3 17,4.1,7:30,1.25,0.4,3 18,5.4,7:30,0.6,0.5,1 19,7.6,7:30,0.6,0.9,1 20,8.6,7:30,0.9,1,2 21,5.6,8:00,0.6,0.4,0 22,3.6,8:00,1,0.3,2 23,6.7,8:00,0.15,0.7,0 24,8.8,7:45,1.3,1,2 25,6,9:30,1.9,0.3,4 26,6.4,10:00,0.75,0.35,2 27,6.8,9:00,0.12,0.4,0 28,6.7,8:45,0.15,0.8,0 29,14,8:30,2.1,0,4 30,14.4,9:00,0.4,0,0 31,17,9:00,0.3,0.15,1 32,9.7,10:30,0.27,0,0 33,13.5,9:00,0.26,0,0 34,14.5,8:55,0.35,0.1,0 35,17,8:45,0.75,0.1,1 36,21.1,8:00,0.29,0.6,0 37,22.1,7:55,0.21,0.8,0 38,8,9:30,0.5,0.1,1 39,8.9,10:00,2,0,4 40,9.3,10:00,0.15,0.8,0

67 41,5.5,10:30,1.6,0.3,3 othertree dist(m),9.5 angle,8:00 NSdrip(m), 5.5 EWdrip(m), 7.1 height(m), 10.1 othertree dist(m),8.9 angle,7:30 NSdrip(m), 4.5 EWdrip(m), 3.9 height(m), 10.1 site title, Digger Pine Site 3 lat, 40 41.464N long, 122 12.026W girth(m), 3.20 ConvertGirth height(m), 31 dripline3(m), 8.5 dripline6(m), 5.0 dripline9(m), 9.9 dripline12(m), 12.5 gradient, 1.6/15.85 north, 0deg w of 3oclock date, 11 Oct 2013 Cones saplings, * dist(m) angle height(m) shade whorls 1,9.81,9:00,2.13,5 2,10.71,8:55,0.7,2 3,13.01,9:00,1.9,4 4,14.91,9:10,1.1,1 5,8.71,9:30,3.04,7 6,10.01,9:35,3.657,6 7,22.41,12:30,0.9,2 8,24.81,2:00,0.4,0 9,20.01,1:55,0.38,0 10,7.31,4:15,0.35,0 11,10.01,4:30,1.2,5 12,10.01,4:31,2.05,6 13,10.91,4:35,2.2,7 14,12.91,5:00,0.28,0 15,12.71,5:05,0.26,0 16,12.21,5:10,0.18,0

68 17,10.61,5:15,0.25,0 18,8.81,5:20,0.22,0 19,11.51,5:45,0.4,1 20,13.31,5:45,0.4,1 21,15.81,5:46,0.35,0 22,13.91,5:40,0.2,0 23,13.3,5:50,0.5,1 24,13.91,5:55,0.55,1 25,13.71,6:10,0.6,1 26,11.71,6:30,0.55,1 27,11.91,6:35,0.45,1 28,12.41,7:00,0.6,2 29,15.01,6:55,0.5,1 30,8.91,8:00,0.25,0 31,8.96,8:01,0.28,0 32,9.71,7:59,0.2,0 33,16.41,4:30,2.133,5 34,18.61,4:35,3.04,3 35,16.41,5:15,0.26,0 36,8.71,5:15,0.24,0 site title, Digger Pine Site 3 extended lat, 40 41.464N long, 122 12.026W girth(m), 3.20 ConvertGirth height(m), 31 dripline3(m), 8.5 dripline6(m), 5.0 dripline9(m), 9.9 dripline12(m), 12.5 gradient, 1.6/15.85 north, 0deg w of 3oclock date, 11 Oct 2013 Cones saplings, * dist(m) angle height(m) shade whorls 1,9.8,9:00,2.13,5 2,10.7,8:55,0.7,2 3,13.0,9:00,1.9,4 4,14.9,9:10,1.1,1 5,8.7,9:30,3.04,7 6,10.0,9:35,3.657,6 7,22.4,12:30,0.9,2 8,24.8,2:00,0.4,0 9,20.0,1:55,0.38,0

69 10,7.3,4:15,0.35,0 11,10.0,4:30,1.2,5 12,10.0,4:31,2.05,6 13,10.9,4:35,2.2,7 14,12.9,5:00,0.28,0 15,12.7,5:05,0.26,0 16,12.2,5:10,0.18,0 17,10.6,5:15,0.25,0 18,8.8,5:20,0.22,0 19,11.5,5:45,0.4,1 20,13.3,5:45,0.4,1 21,15.8,5:46,0.35,0 22,13.9,5:40,0.2,0 23,13.3,5:50,0.5,1 24,13.9,5:55,0.55,1 25,13.7,6:10,0.6,1 26,11.7,6:30,0.55,1 27,11.9,6:35,0.45,1 28,12.4,7:00,0.6,2 29,15.0,6:55,0.5,1 30,8.9,8:00,0.25,0 31,8.96,8:01,0.28,0 32,9.717:59,0.2,0 33,16.4,4:30,2.133,5 34,18.6,4:35,3.04,3 35,16.4,5:15,0.26,0 36,8.7,5:15,0.24,0 * o1 37,106,11:15,6ft 38,105,11:12,4ft * o2 39,101,11:09,1ft 40,101.1,11:09.4,1ft 41,98,11:07,2ft 42,97,11:09,1.5ft * o3 * o4 43,82,10:49,3ft * o5 * o6 44,92,10:40,3ft 45,90,10:32,1.5ft * o7 46,89,10:21,2ft 47,90,10:18,.40 48,89.5,10:18,2ft 49,85,10:18,5ft

70 50,87,10:16,4ft * o8 51,77,9:59,1.5ft * o9 52,79,9:50,1ft 53,78.5,9:45,3.5ft 54,80,9:44,1.5ft 55,87,9:50,6ft 56,88,9:49,2ft 57,85,9:45,1ft * othertree, O1 dist(m), 107.8 angle,11:12 NSdrip(m),7.2 EWdrip(m),6.6 othertree, O2 dist(m),101.2 angle,11:08 NSdrip(m),8.6 EWdrip(m),8.5 othertree, O3 dist(m),102.6 angle,10:55 NSdrip(m), 5.6 EWdrip(m),7.6 othertree, O4 dist(m),83.9 angle,10:46 NSdrip(m),5.1 EWdrip(m),5.2 othertree, O5 dist(m),97.7 angle,10:30.5 NSdrip(m),14.0 EWdrip(m),13.2 othertree, O6 dist(m),89.9 angle,10:34.5 NSdrip(m),9.5 EWdrip(m),10.5 othertree, O7 dist(m),88.2 angle,10:19 NSdrip(m),5.2 EWdrip(m),9.2 othertree, O8

71 dist(m),79.6 angle,10:00 NSdrip(m), 8.6 EWdrip(m),10.4 othertree, O9 dist(m),83.3 angle, 9:49 NSdrip(m),9.0 EWdrip(m),12.2 site title,Digger Pine Site 4 lat,40 41.271N long,122 11.843W girth(m),2.41 ConvertGirth height(m),26.5 dripline3(m),7 dripline6(m),9.3 dripline9(m),9.3 dripline12(m),8.2 gradient,1.8/14.0 north,68deg w of 3oclock date, 24 Oct 2013 cones * n,dist(m),angle,age(yr),seeds 1,2.4,11:55,3 2,2.6,11:55,3 3,3.55,11:50,4 4,4.45,11:50,3 5,4.6,11:51,4 6,4.85,11:53,2 7,5.3,11:53,4 8,2,11:30,5 9,3.335,11:40,3 10,3.2,11:39,5 11,3.75,11:40,1 12,4.15,11:38,4 13,4.7,11:37,5 14,4.9,11:30,3 15,4.25,11:31,5 16,3.5,11:32,3 17,3.1,11:30,3 18,6.5,11:30,5 19,7.15,11:32,3 20,7.45,11:25,2 21,9,11:30,3

72 22,7.75,11:20,2 23,4.9,11:22,3 24,2.75,11:19,5 25,3.1,11:19,3 26,2.85,11:15,4 27,3.65,11:15,5 28,3.1,11:13,3 29,4.65,11:15,3 30,5,11:13,5 31,5.2,11:12,5 32,6.1,11:12,4 33,6.25,11:11,5 34,6.35,11:13,1 35,7.2,11:13,1 36,6.5,11:10,4 37,6.7,11:10,5 38,7.55,11:05,3 39,7.7,11:03,3 40,8.7,11:00,3 41,9.75,11:01,1 42,2.8,10:59,3 43,3.1,10:58,5 44,4.1,10:57,1 45,3.95,10:57,4 46,3.85,10:56,5 47,4.8,10:57,5 48,5.8,10:56,2 49,7.4,10:55,3 50,2,11:15,5+ 51,2.95,10:45,5 52,2.75,10:43,5 53,4.9,10:43,4 54,4.6,10:40,3 55,4.75,10:39,2 56,5.3,10:43,4 57,6.6,10:40,3 58,7.35,10:38,1 59,7.85,10:38,3 60,8,10:39,3 61,8.15,10:37,3 62,1.8,10:00,4 63,3.1,10:00,5 64,3.2,10:00,2 65,4.3,10:00,5 66,5.5,10:05,4 67,5.9,10:10,2 68,6.7,9:55,3

73 69,8.1,9:53,4 70,6.75,9:50,5 71,6.95,9:59,1 72,6.7,9:58,4 73,7.45,9:50,2 74,1.8,9:51,5 75,2.1,9:50,2 76,3.3,9:45,5 77,3.55,9:43,4 78,3.8,9:42,4 79,4.75,9:40,3 80,5.3,9:42,3 81,5.85,9:42,3 82,6.1,9:42,5 83,4.8,9:40,3 84,2.1,9:35,5 85,3.5,9:35,5 86,3.6,9:34,5 87,3.55,9:34,4 88,3.75,9:34,5 89,3.75,9:32,5+ 90,3.2,9:20,1 91,5.2,9:10,2 92,5.2,9:15,4 93,5.5,9:20,3 94,5.65,9:20,5 95,5.4,9:20,5 96,5.2,9:21,3 97,5.3,9:25,3 98,6.7,9:01,2 99,6.5,9:02,3 saplings * n,dist(m),angle,height(m),shade,whorls, 1,13.8,9:20,0.6 2,14.1,9:20,0.8 3,13.8,9:21,0.8 4,14.5,9:20,1 5,16.1,9:20,0.7 6,16,9:20,0.5 7,9.6,9:30,2 8,14.2,9:30,1.5 9,20.7,10:00,0.8 10,26.7,11:50,0.22 11,26.7,11:48,1.6 12,30.4,11:45,0.75 13,36.6,10:16,1.3 14,36.1,10:15,1.1

74 15,38.2,10:15,0.8 16,40,10:17,1 17,41.3,10:18,1.2 18,40.6,10:18,1.5 19,36.9,10:17,1.2 20,36.92,10:16,1.3 21,34.6,10:18,1.2 22,36.3,10:17,1.2 23,36.9,10:19,1.4 24,40.4,11:05,1.6 25,25.6,11:20,0.4 26,26.5,11:23,1.2 27,21.3,1:00,1.1 28,21.8,1:00,2 29,24.4,1:15,1.9 30,19.6,5:30,0.9 31,20.8,5:32,0.75 32,20.2,5:33,1 33,24.2,5:34,0.8 34,23,5:35,1.1 35,24.7,5:36,1.1 36,24.7,3:59,0.9 37,23.8,4:00,0.7 38,25.1,4:01,1 39,34.5,2:29,1.2 40,34.9,2:30,1.1 41,35.3,2:30,2.1 42,36.5,2:31,1.7 43,42,2:01,1.6 44,42.9,2:00,1 45,5.9,11:54,0.15 46,6.5,12:05,0.1 47,30.2,12:00,8 othertree dist(m),11.6 angle,9:00 NSdrip(m),9.6 EWdrip(m),7 othertree dist(m),28.2 angle,11:50 NSdrip(m),11 EWdrip(m),10 othertree dist(m),34.6 angle,10:15 NSdrip(m),10

75 EWdrip(m),7.4 othertree dist(m),39 angle,10:14 NSdrip(m),5.8 EWdrip(m),7 othertree dist(m),43.5 angle,10:18 NSdrip(m),7 EWdrip(m),8.6 othertree dist(m),39 angle,10:19 NSdrip(m),4.8 EWdrip(m),4.3 othertree dist(m),37.6 angle,11:05 NSdrip(m),10.6 EWdrip(m),8.5 othertree dist(m),26.5 angle,11:10 NSdrip(m),8.5 EWdrip(m),9 othertree dist(m),22 angle,1:00 NSdrip(m),11 EWdrip(m),12 othertree dist(m),25.5 angle,5:30 NSdrip(m),8 EWdrip(m),9 othertree dist(m),24.5 angle,4:00 NSdrip(m),8 EWdrip(m),10 othertree dist(m),32 angle,2:30 NSdrip(m),9 EWdrip(m),9 othertree

76 dist(m),42 angle,2:00 NSdrip(m),2 EWdrip(m),2 othertree dist(m),5.9 angle,11:55 NSdrip(m),1.8 EWdrip(m),2 othertree dist(m),6.6 angle,12:05 NSdrip(m),1.2 EWdrip(m),1 site title, Digger Pine Site 5 lat, 40 41.357N long, 122 11.116W girth(m), 1.99 ConvertGirth height(m), 24.4 dripline3(m), 2.4 dripline6(m), -2.0 dripline9(m), 5.1 dripline12(m), 14.0 gradient, 1.8/6.4 north, 36deg E of 12oclock treecones, 14 date, 25 Oct 2013 Cones * n, dist(m), angle, age(yr), seeds 1,2.6,3:15,3,5% 2,6.1,1:00,3,0 3,8.4,12:10,1,90% 4,8.3,12:00,4,0 5,8.5,12:00,5,chewed 6,8.7,11:55,4,chewed 7,11.1,11:50,4,0 8,12.1,11:30,1,50% 9,12.7,11:25,5,0 10,10.3,11:20,4,0 11,12.6,11:15,3,0 12,6.7,11:30,5,0 13,6.9,11:10,4,0 14,6.95,11:10,1,15% 15,8.3,11:10,4,0

77 16,9.8,11:00,3,0 17,11.7,10:45,1,40% 18,7.4,11:00,4,0 19,6.4,11:00,5,0 20,6.2,10:45,3,0 21,6.4,10:40,2,0 22,3.6,11:15,5,0 23,3.8,11:10,5,0 24,3.9,11:00,5,0 25,4.4,10:45,4,0 26,6.3,11:00,3,0 27,6.5,11:05,3,5% 28,6.1,10:30,1,60% 29,6.4,10:20,3,0 30,11.6,10:45,1,30% 31,9.4,10:40,3,0 32,7.9,10:30,1,90% 33,7.8,10:40,5,0 34,7.9,10:20,5,0 35,7.2,10:10,3,0 36,6.6,10:00,4,0 37,7.3,9:50,5,0 38,7.3,9:45,5,0 39,6.4,9:50,3,0 40,7.1,9:55,5,0 41,7.3,9:57,5,0 42,7.5,9:55,4,0 43,6.1,9:50,4,0 44,7.1,9:46,5,0 45,5.4,9:30,5,0 46,6.4,9:10,4,0 47,6.0,8:55,4,chewed 48,4.0,7:00,3,0 saplings * n dist(m) angle height(m) knots 1,9.8,10:05,1.8,3 2,32.2,5:55,1.2,2 3,31.6,5:40,.65,0 4,35.2,5:10,1.3,1 othertree dist(m),9.0 angle,10:00 nsdrip(m),5 ewdrip(m),4 othertree dist(m),30.1 angle,5:45

78 nsdrip(m),8 ewdrip(m),9 othertree dist(m),35.1 angle,5:15 nsdrip(m),8 ewdrip(m),7 site title, Eastside Slope Diggers lat, 40 41.301 long, 122 11.304 girth(m), 2 ConvertGirth height(m), 30 dripline3(m), 4 dripline6(m), 4 dripline9(m), 4.5 dripline12(m), 4.5 gradient, 1/3 north, 5deg e of 9oclock date, 15 Oct 2013 saplings 1, 13, 12:25, 2.5, 0 2, 14, 12:41, 2.5, 0 3, 9, 12:55, 1ft, 0 4, 8, 1:20, 3, 0 5, 18.4, 2:06, 2, 0 6, 26.7, 12:35, 9ft, 0 7, 29.5, 12:19, 2.5, 0 8, 29.45, 12:20, 2.5, 0 9, 31, 12:25, 1.5, 0 10, 32.5, 12:36, 1ft, 0 11, 32.5, 12:37, 1ft, 0 12, 33, 12:55, 1.5, 0 13, 34.5, 1:03, 2.5, 0 14, 10.5, 11:00, 1.5ft, 0 15, 21, 10:00, 2.5, 0 16, 25, 11:12, 2.5, 0 alttree altdist(m), 25.5 altangle, 12:32 altnsdrip(m), 7 altewdrip(m), 8 alttree altdist(m), 28 altangle, 12:39

79 altnsdrip(m), 7 altewdrip(m), 8 alttree altdist(m), 26 altangle, 11:05 altnsdrip(m), 7 altewdrip(m), 8

80 Appendix B

Digger Seeds

Figure 29 – Example of Double Seeds per Scale

81

Figure 30 – Double Seeds Can Even Occur on the Top Scale

82 Figure 31 – Digger Seeds

83 Figure 32 – Granary Tree DH21, Close-Up of Content

84 Figure 33 – Data Tree #5 as Granary Tree Close-Up of Content

85 Figure 34 – DH Data Sites Within Study Area (North is to Right)

86