SAGUARO
Ecology of the Saguaro: II
NPS Scientific Monograph No. 8
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CHAPTER 5:
GROWTH (continued)
Geographic Variation in
Growth
The rate of growth can vary significantly and even
widely between saguaro populations separated by relatively short
distances, especially where elevation, topography, and/or soil structure
also vary greatly over short distances. For example, several different
effective environments characterize different desert areas and habitats
within Saguaro National Monument, and between the east and west
divisions of the monument (Table 32).
TABLE 32. Six-year on-site record of summer precipitation
(inches; June-October) at Saguaro National Monument, east (SEF) and west
flats (SWF).
|
Month |
1968
| 1969
|
1970
| 1971
|
1972
| 1973
|
SWF | SEF |
SWF | SEF |
SWF | SEF |
SWF | SEF |
SWF | SEF |
SWF | SEF |
|
| Precipitation
(inches)
|
June | 0a | 0b | 0 | 0 |
0.14a | 0.20 | 0a | 0 |
1.94 | 0.01 | 0.32 | 0.72 |
July | 2.43 | 1.92 | 1.70 | 0.89 |
1.22 | 3.07 | 0.90 | 2.22 |
1.26 | 0.03 | 1.77 | 2.20 |
Aug. | 2.76 | 2.77c | 1.57 | 2.96 |
1.67 | 2.13 | 5.96 | 5.83 |
1.57 | 0.03 | 0.55 | 0.37b |
Sept. | 0a | 0.12c | 0.81 | 0.80 |
3.14 | 2.91 | 1.47 | 1.70 |
0.37 | 1.49 | 0a | 0 |
Oct. | 0.63 | 1.58 | 0.12 | 0.18 |
0.25 | 0.25 | 2.54 | 2.32 |
3.87 | 5.49 | 0 | 0.64 |
Cum. total | 5.82 | 6.39 | 4.20 | 4.73 |
6.42 | 8.56 | 10.87 | 12.07 |
9.01 | 7.05 | 2.64 | 3.93 |
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Still greater differences in growth rate can be
predicted for populations separated by greater distances and growing
under more widely distinctive effective environments, for example, as
indicated by the climatic data for Tucson, Ajo, and Yuma, Arizona (Table
33). We predict large order growth-rate differences between Tucson
populations and populations in the Yuma desert (Gran Desierto) area.
There the saguaros on bajadas and valley fill are essentially a desert
riparian species population with low marginal density under 3.38 inches
(86 mm) annual rainfall and 1.57 inches (40 mm) summer (May-October)
rainfall (Green and Sellers 1964:493, Yuma; Fig. 9B).
TABLE 33. Summary of climatic parameters for Tucson (elev. 2430
ft; 740 m), Ajo (elev. 1763 ft; 537 m), and Yuma Citrus Station (elev.
191 ft; 58.2 m). Arizona (precipitation given in inches; temperatures
shown in °F).
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| Tucson |
Ajo | Yuma |
|
Mean Precipitation |
Summer (May-Oct.)a | 6.7 inches | 5.0 inches | 1.4 inches |
Winter (Nov.-April)a | 4.5 inches | 4.0 inches | 1.9 inches |
Annualb | 10.91 inches | 9.10 inches | 3.28 inches |
Mean Temperature |
Summer (May-Oct.)a | 79.0° | 83.4° | 83.7° |
Winter (Nov.-April)a | 55.9° | 59.3° | 60.0° |
Annualb | 67.3° | 71.2° | 71.9° |
Maximumb | 82.9° | 84.0° | 87.7° |
Minimumb | 51.6° | 58.3° | 56.1° |
Potential Evapotranspirationc | 41.68 inches/yr | 47.19 inches/yr | 47.8 inches/yr |
Extreme Temperatures |
Summer maximumb | 115° (1960) | 115° (1960)e | 119° (1957) |
Winter minimumb | 6° (1913) | 17° (1937)f | 19° (1937)g |
Jan. 1937d | 15° | 17° | 19° |
Jan. 1962d | 20° | 25° | 23° |
Freeze Occurenceh |
Frost-free days/yr(>32°) | 231 | 349 | 294 |
First freeze (P 32° = 0.50) | Nov. 14 | Jan. 3 | Dec. 10 |
Last freeze (P 32° = 0.50) | Mar. 28 | Jan. 19 | Feb. 19 |
aGreen 1964.
bGreen and Sellers 1964.
cBuol 1964.
dU.S. Weather Bureau Climatological Records.
eAlso recorded on earlier years.
fNo available record of temperatures prior to 1915.
gNo available record of temperatures prior to 1920.
hGreen and Kangieser 1967; minimum temperatures of 32°F or less.
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Organ Pipe Cactus National Monument
Measurement of saguaro growth over a period of 26
years in populations at Organ Pipe Cactus National Monument provides an
important sample for the study of the Species that is summarized in
Table 34 and graphed in Figs. 43 and 44. The curve in Fig. 44 is fitted
by multiple linear regression as in the case for the Saguaro National
Monument data (Fig. 40). As at Saguaro National Monument and elsewhere,
the asymptote of the growth curve for Organ Pipe Cactus National
Monument is at approximately 2 m (6.6 ft) stem height (Fig. 44). As
noted above, important changes in form and functionand important
consequencestake place in the saguaro in southern Arizona when the
plant reaches a stem height on the order of 2 m (6.6 ft).
TABLE 34. Saguaro stem height and apical growth (cm) at Organ
Pipe Cactus National Monument, Arizona. Plant height in 1941 (Lightle
1941) and height of the same plants in 1967 provide a 26-yr apical
growth increment. Data graphed in Figs. 43 and 44.
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No. | Height (1941) |
Height (1967) | 26-yr growth
increment |
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ALAMO CANYON |
1 | 15.2 | 48.2 | 33.0 |
2 | 15.2 | 81.3 | 66.1 |
3 | 25.4 | 114.3 | 88.9 |
4 | 30.5 | 132.1 | 101.6 |
5 | 30.5 | 157.5 | 127.0 |
6 | 30.5 | 165.1 | 134.6 |
7 | 30.5 | 129.5 | 99.1 |
8 | 30.5 | 101.6 | 71.1 |
9 | 30.5 | 144.8 | 114.3 |
10 | 30.5 | 172.7 | 142.2 |
11 | 30.5 | 210.6 | 180.1 |
12 | 30.5 | 157.5 | 127.0 |
13 | 30.5 | 104.1 | 73.6 |
14 | 30.5 | 127.0 | 96.5 |
15 | 30.5 | 116.8 | 86.3 |
16 | 30.5 | 162.5 | 132.0 |
17 | 30.5 | 165.1 | 134.6 |
18 | 43.2 | 302.0 | 258.8 |
19 | 45.7 | 312.4 | 266.7 |
20 | 45.7 | 213.4 | 167.6 |
21 | 45.7 | 109.2 | 63.5 |
22 | 45.7 | 142.3 | 96.5 |
23 | 45.7 | 119.1 | 73.6 |
24 | 45.7 | 259.1 | 213.4 |
25 | 45.7 | 287.0 | 241.3 |
26 | 61.0 | 223.5 | 162.5 |
27 | 61.0 | 203.2 | 142.3 |
28 | 61.0 | 172.7 | 111.8 |
29 | 61.0 | 182.9 | 121.9 |
30 | 79.2 | 254.0 | 177.8 |
31 | 91.4 | 345.4 | 254.0 |
32 | 106.7 | 309.9 | 203.2 |
33 | 106.7 | 416.6 | 309.9 |
34 | 182.0 | 510.5 | 358.0 |
35 | 195.6 | 528.3 | 332.5 |
36 | 243.8 | 538.5 | 294.7 |
37 | 335.3 | 538.5 | 203.2 |
38 | 365.8 | 711.2 | 345.4 |
39 | 366.0 | 718.7 | 353.0 |
40 | 426.7 | 853.4 | 426.7 |
41 | 487.7 | 670.6 | 182.9 |
42 | 548.6 | 716.3 | 167.7 |
43 | 579.1 | 797.6 | 188.0 |
DOWLING WELL |
44 | 30.5 | 101.6 | 71.1 |
45 | 30.5 | 78.7 | 48.2 |
46 | 30.5 | 134.6 | 104.1 |
47 | 30.5 | 101.6 | 71.1 |
48 | 45.7 | 106.7 | 61.0 |
49 | 61.0 | 182.8 | 121.8 |
50 | 61.0 | 160.0 | 99.0 |
51 | 121.9 | 248.9 | 127.0 |
52 | 182.9 | 340.4 | 157.5 |
53 | 182.9 | 388.6 | 205.7 |
54 | 182.9 | 348.0 | 165.1 |
55 | 182.9 | 335.3 | 152.4 |
56 | 195.6 | 373.4 | 177.8 |
57 | 213.4 | 373.4 | 160.0 |
58 | 213.4 | 462.3 | 248.9 |
59 | 213.4 | 378.4 | 165.0 |
60 | 213.4 | 408.9 | 195.5 |
61 | 213.4 | 388.6 | 175.2 |
62 | 213.4 | 388.6 | 175.2 |
63 | 274.3 | 439.4 | 165.1 |
64 | 274.3 | 469.9 | 195.6 |
65 | 304.8 | 421.6 | 116.8 |
66 | 365.8 | 513.1 | 147.3 |
67 | 365.8 | 586.7 | 220.9 |
68 | 365.8 | 551.2 | 185.4 |
69 | 365.8 | 538.5 | 172.7 |
70 | 365.8 | 518.1 | 152.3 |
71 | 396.2 | 556.3 | 160.1 |
72 | 426.7 | 609.6 | 182.9 |
73 | 426.7 | 563.9 | 137.2 |
74 | 426.7 | 589.3 | 162.6 |
75 | 487.7 | 777.2 | 289.5 |
76 | 487.7 | 647.7 | 160.0 |
77 | 487.7 | 579.1 | 91.4 |
78 | 518.2 | 640.1 | 121.9 |
79 | 548.6 | 624.8 | 76.2 |
80 | 548.6 | 685.8 | 137.2 |
81 | 609.6 | 868.7 | 259.1 |
82 | 609.6 | 853.4 | 243.8 |
83 | 609.6 | 680.7 | 71.1 |
84 | 609.6 | 823.0 | 152.4 |
85 | 731.5 | 934.7 | 203.2 |
86 | 731.5 | 787.4 | 55.9 |
87 | 792.5 | 861.1 | 68.6 |
88 | 792.5 | 990.6 | 198.1 |
89 | 853.4 | 1000.7 | 147.3 |
90 | 853.4 | 1117.6 | 264.2 |
91 | 853.4 | 1046.5 | 193.1 |
92 | 853.4 | 929.6 | 76.2 |
93 | 875.4 | 1051.6 | 76.2 |
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The data on apical growth of saguaros at Organ Pipe
Cactus National Monument,2 shown in Table 34 and graphed in
Fig. 44, provide, for the first time, a basis for estimating mean growth
rates and age-height relationships of saguaro populations whose growth
rates are not under the dominant depressing influence of catastrophic
freezing. Growth rates for this population, situated close to the
ecological and geographic center of the species range, differ markedly
from those for saguaro populations at Saguaro National Monument and
elsewhere near the cold-limited boundaries of the species range.
2Lightle (1941) field data and re-measured
in 1967 to obtain a 26-year apical growth increment.
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Fig. 43. Apical growth on stem height
(1941) of 93 saguaros at Organ Pipe Cactus National Monument, Arizona,
on uniform (arithmetic) scale. Note the reversal in the rate of growth
that occurs at aproximately 200 cm (6.6 ft) height. Data in Table 34.
(click on image for an enlargement in a new window)
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Fig. 44. Regression on logarithmic
coordinates of 26-year apical growth on 1941 stem height (Lightle 1941)
for 93 saguaros at Organ Pipe Cactus National Monument, Arizona. Note
asymptote of curve at approximately 2 m (6.6 ft); see text. Data in
Table 34. (click on image for an enlargement in a new
window)
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A comparative view of a significant difference in
growth rate between two saguaro populations in southern Arizonaat
Saguaro National Monument (east) in southeastern Arizona and Organ Pipe
Cactus National Monument in south-central Arizonais provided in
Fig. 45. The differences in growth-rate characteristics for these two
populations, as shown in Fig. 45, are attributable to differences in
temperature and precipitation. Gross differences in parameters of the
climate at Tucson and Ajo (Table 33) relate importantly to observed
differences in saguaro growth rates and to height-age relationships for
saguaros in these two different climatic environments.
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Fig. 45. Regression on logarithmic
coordinates of 26-year apical growth on height for young saguaros at
Saguaro National Monument (east) and Organ Pipe Cactus National Monument
(see Tables 8, 33, and 34 and Figs. 38, 39, 42, and 43). See discussion
in text.
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As compared to the climatic environment of Saguaro
National Monument and the Tucson locality, the climatic environment of
Organ Pipe Cactus National Monument and Ajo locality is a much warmer
and drier one that results in the slower growth of juvenile
saguaros. In its overall effects on the growth of larger plants,
however, the warmer environment at Organ Pipe Cactus National Monument
offers conditions for saguaro growth more favorable than those at
Saguaro National Monument (see Hastings and Alcorn 1961). There, the
effects of catastrophic freezing offset the potentially greater benefits
of substantially higher plant-available moisture and lower transpiration
losses.
Tempe, Arizona
Investigation of saguaro growth in a natural
population of saguaros in the Phoenix, Arizona, area was conducted by
Marshall and Earle (Marshall 1955) on South Hill in the Desert Botanical
Garden, Tempe. Their data for 6 years (1949-54) are given in Table 35.
The data appear to be in general accord with our observations and those
reported by others on growth rates for young saguaros in this portion of
the extreme range of the species.
TABLE 35. Height measurements (cm) and subsequent one-year
apical growth increments (cm) for 14 young saguaros at Tempe, Arizona
from annual measurements by W. Hubert Earle, reported by Marshali
(1955:57). Original data in inches.
|
No. | Original height (cm) |
One-yr apical growth (cm)
|
1949 | 1950 |
1951 | 1952 |
1953 | 1954 |
Mean |
|
1 | 5.4 |
1.6 |
1.3 |
1.9 |
1.3 |
3.8 |
0.6 |
1.8 |
2 |
14.0 |
3.2 |
0.6 |
3.8 |
1.3 |
1.3 |
0.6 |
1.8 |
3 |
22.2 |
6.4 |
1.9 |
6.4 |
5.1 |
3.8 |
5.1 |
4.8 |
4 |
28.6 |
5.1 |
0.6 |
8.3 |
5.7 |
3.8 |
3.2 |
4.4 |
5 |
56.5 |
10.2 |
3.2 |
14.0 |
10.8 |
7.6 |
12.7 |
9.8 |
6 |
61.3 |
3.8 |
1.3 |
7.0 |
5.7 |
5.1 |
7.0 |
5.0 |
7 |
64.4 |
6.0 |
2.5 |
7.6 |
0.6 |
11.4 |
9.5 |
6.3 |
8 |
85.1 |
15.2 |
2.5 |
12.1 |
13.3 |
9.5 |
12.1 |
10.8 |
9 |
88.9 |
14.0 |
3.8 |
15.2 |
10.8 |
11.4 |
14.6 |
11.6 |
10 |
114.3 |
8.9 |
4.4 |
5.1 |
8.9 |
3.2 |
5.7 |
6.0 |
11 |
116.8 |
14.0 |
2.5 |
14.6 |
13.3 |
10.2 |
15.2 |
11.6 |
12 |
129.5 |
14.0 |
3.2 |
12.7 |
14.0 |
8.3 |
12.7 |
10.8 |
13 |
165.7 |
20.3 |
7.0 |
23.5 |
22.9 |
9.5 |
17.8 |
16.8 |
14 |
256.5 |
22.9 |
6.4 |
21.6 |
26.0 |
20.3 |
24.8 |
20.3 |
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Tucson, Arizona
In the Tucson area, other investigations of saguaro
growth have been reported by MacDougal and Spalding (1910), Shreve
(1910), Hastings (1961), and Hastings and Alcorn (1961).
MacDougal and Spalding (1910) measured and reported
yearly apical growth increments for individuals in a saguaro population
on Tumamoc Hill in the Tucson Mountains. The data are for years 1905-8
and, though long overlooked, they are the first growth data obtained and
reported for the species (see Table 36). It is a peculiarity that Shreve
(1910), reporting in the same year from the same laboratory, included no
reference to these important data.
TABLE 36. Original height measurements (cm) and subsequent
one-yr apical growth increments (cm) for 8 saguaros on Tumamoc Hill,
Tucson, Arizona, from periodic stem height measurements reported by
MacDougal and Spalding (1910: Table 8).
|
Plant No. |
Original measurement
|
Observed one-yr apical growth
|
Year | Height |
1906 | 1907 |
1908a | 1908b |
Mean |
|
24 | 1906 | 4.5 | |
2.9 | 2.2 | | 2.55 |
25 | 1906 | 37.4 | |
6.1 | 8.3 | 15.0 | 9.80 |
21 | 1905 | 70.4 | 13.5 |
5.9 | 7.1 | | 8.83 |
10 | 1905 | 86.8 | 5.4 |
0.1 | 5.9 | | 6.47 |
4 | 1905 | 162.1 | 7.4 |
13.7 | 10.8c | 10.8c | 10.70 |
12 | 1905 | 163.8 | 8.6 |
2.0 | 16.2c | 16.2c | 10.75 |
19 | 1905 | 223.6 | 15.6 |
5.2 | 28.6c | 28.6c | 19.50 |
22 | 1905 | 230.2 | 10.9 |
13.6 | 14.2c | 14.2c | 13.25 |
a23 Mar. 1908.
b22 Oct. 1908.
cMeans of two seasons' growth from 23 Jan. 1907 to 22 Oct. 1908. |
The growth record provided by MacDougal and Spalding
(1910:30, Table 8) contains the highest natural rate of growth ever
reported for the saguaro. During the 9-month period from 23 January 1907
to 22 October 1908, one individual (No. 19) in that sample grew from a
height of 244.4 cm (96.2 inches) to 301.6 cm (118.7 inches), an average
of 28.6 cm (11.25 inches) per year. It is noteworthy in relation to that
unusually high rate of growth that the July to September precipitation
during the 1907 and 1908 summer growth periods was 8.53 inches (217 mm)
(Green and Sellers 1964: Tucson) and 12.05 inches (306 mm) (MacDougal
and Spalding 1910, on site), respectively. Both values are substantially
above the 5.01-inch (127 mm) Tucson mean for these 3 months (Green and
Sellers 1964).
Shreve (1910) provided none of the data used to
obtain the age-height relationships he reported for saguaros growing on
the northern slope of Tumamoc Hill in Tucson (Table 37). Using his data,
a plot of height on age, however, shows an exponentially linear
age-height relationship, with a distinct increase in the growth rate
occurring at a height of approximately 24 cm (9.4 inches). Our results,
however, do not suggest the occurrence of such a growth-rate change at
that height (Tables 28, 30; Figs. 40-42).
TABLE 37. Height, estimated age and one-yr apical growth for
saguaros on Tumamoc Hill, Tucson, Arizona; apical growth is subsequent
one-yr apical growth determined by graphic analysis of height-age
relationships given in Shreve (1910:237).
|
Height (cm) | Age (yrs) |
Apical growth (cm/yr) |
|
10 | 8.0 | 2.0 |
20 | 12.5 | 2.5 |
40 | 19.1 | 4.0 |
80 | 27.3 | 7.0 |
100 | 30.3 | 8.0 |
200 | 40.5 | 12.0 |
300 | 47.5 | 15.0 |
400 | 54.0 | 20.0 |
500 | 60.0 | 40.0 |
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The growth rates reported by Hastings and Alcorn
(1961: Figs. 5 and 6) also showed the reversal in the annual apical
growth rate occurring at a height of approximately 2 m (6.6 ft), typical
of saguaro growth-form ontogeny thus far known (see Tables 34 and 38;
Figs. 43 and 44).
Table 38. Mean annual apical growth of saguaros in USDA sample
at Saguaro National Monument 1951-59. Adjusted data in Hastings and
Alcorn (1961:Fig. 5).
|
Height-class (ft) |
Mid-point
|
Apical growth (cm/yr) |
(ft) | (cm) |
|
0-1.99 | 1 | 30.48 | 8.68 |
2-3.99 | 3 | 91.44 | 9.33 |
4-5.99 | 5 | 152.40 | 11.11 |
6-7.99 | 7 | 213.36 | 11.25 |
8-9.99 | 9 | 274.32 | 10.20 |
10-11.99 | 11 | 335.28 | 7.29 |
12-13.99 | 13 | 396.24 | 7.51 |
14-15.99 | 15 | 457.20 | 7.07 |
16-17.99 | 17 | 518.16 | 5.21 |
18-19.99 | 19 | 579.10 | 5.30 |
20-21.99 | 21 | 640.08 | 6.80 |
22-23.99 | 23 | 701.04 | 6.71 |
24-25.99 | 25 | 762.00 | 6.25 |
26-27.99 | 27 | 822.96 | 6.49 |
28-29.99 | 29 | 883.92 | 6.20 |
30-31.99 | 31 | 944.88 | 8.00 |
32-33.99 | 33 | 1005.84 | 9.25 |
34-35.99 | 35 | 1066.80 | 6.80 |
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Last Updated: 21-Oct-2005
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