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Reproductive Potential and Minimum Reproductive Size of Ferocactus Wislizeni (Cactaceae)

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Reproductive Potential and Minimum Reproductive Size of Ferocactus Wislizeni (Cactaceae) Reproductive Potential and Minimum Reproductive Size of Ferocactus wislizeni (Cactaceae) Item Type Article Authors Bowers, Janice E. Publisher University of Arizona (Tucson, AZ) Journal Desert Plants Rights Copyright © Arizona Board of Regents. The University of Arizona. Download date 04/10/2021 01:51:49 Link to Item http://hdl.handle.net/10150/554307 Ferocactus Bowers 3 most cacti are hermaphroditic, some are monoecious, dio- Reproductive Potential and ecious, or even trioecious (Gibson and Nobel 1986, Fleming Minimum Reproductive Size et al. 1994), and although fruit set is generally low in her- maphroditic, outcrossing species in other plant families of Ferocactus wislizeni (Sutherland and Delph 1984, Sutherland 1986), high fruit set is apparently not unusual in the Cactaceae (McGregor (Cactaceae) and Alcorn 1959, McGregor et al. 1962, Johnson 1992, Suzán et al. 1994, Roller 1996, Bowers 1996); these and Janice E. Bowers other traits suggest that cacti could be instructively em- ployed in a variety of comparative studies. U. S. Geological Survey This paper reports a four -year investigation of flower, fruit, 1675 W. Anklam Rd. and seed production of Ferocactus wislizeni ( Engelm.) Britt. Tucson, Arizona 85745 & Rose, a short -columnar, unbranched cactus that is widely distributed in the northern Sonoran Desert. Ultimately, these Abstract data will be used in studies of the reproductive biology and Flower, fruit, and seed production of the short -columnardemography of this species. Ferocactus wislizeni can live cactus Ferocactus wislizeni were studied in the northernas long as 45 years (Goldberg and Turner 1986). The plants Sonoran Desert from 1993 -1996. Most plants of Ferocactusbloom once a year from July- September. Medium -sized wislizeni start to reproduce when they attain a diameter of solitary bees are the main pollinators (McGregor and Alcorn about 19 cm (7.5 in). Number of flowers initiated and fruits 1959, Grant and Grant 1979); the flowers are self- incom- set increased linearly with plant diameter. Plants produced patible or largely so (McGregor and Alcorn 1959). Flow- 25 flowers /year on average, a very low value for a woody,ers are borne on subapical tubercules, nipplelike structures relatively long -lived plant. On average, 93% of initiatedthat contain vegetative and floral meristems (Gibson and flowers set fruit, and fruits contained about 600 -1000 seeds.Nobel 1986). As the plants mature, adjacent tubercules fuse Fruit set seemed unaffected by drought, but seed set wasinto vertical ribs. Annual height growth is largely a func- apparently diminished by unusually dry conditions. Al- tion of tubercule formation (Gibson and Nobel 1986, Nobel though the small number of flowers produced by Ferocactus1986), as is annual diametric growth, at least for part of wislizeni could be a substantial competitive disadvantage, the lifespan (Benson 1982). Mature plants have a large fruit set and fecundity are evidently high enough for popu-volume:surface area ratio. Specific questions addressed lations to maintain themselves. during this study were: 1) what is the minimum size for reproduction; 2) what is the relation between plant size Investigations into reproductive, floral, and pollination bi-and annual flower and fruit production; 3) what propor- ology often require quantitative measures of flower, fruit,tion of flowers set fruit; 4) how many seeds does a fruit and seed production. Such investigations might include al-contain and what proportion of fresh fruit weight do they location of plant resources to reproductive effort (Harperrepresent; and 5) does reproductive effort vary annually? 1977, Silvertown 1982), pollinator behavior on flowers and inflorescences (Gentry 1974a, 1974b; Thomson 1983), the Study Site and Methods trade -off between flower size and flower number (PrimackThe Tumamoc Hill study site (32°13'N, 111°05'W) is at 1985), the connection between breeding system and the 818 m (2684 ft) above sea level on a gently sloping bench relative sizes of ovary and mature fruit (Primack 1987),with a northeast aspect. Tumamoc Hill, an outlier of the the adaptive significance of excess flower production Tucson Mountains, Pima County, Arizona, has a maximum (Stephenson 1981), and the effect of drought on fruit pro-elevation of 948 m (3110 ft) and a basal elevation of 703 m duction and therefore fitness (Parker 1987). Precise repro-(2306 ft). The rocky, basaltic -andesitic slopes are domi- ductive data also contribute to demographic studies, as whennated by Cercidium microphyllum (Torr.) Rose & Johnst., age -specific seed production is used to calculate lifetimeCarnegiea gigantea ( Engelm.) Britt. & Rose, Larrea fecundity or to determine the maximum and actual rates oftridentata (DC.) Cov., Fouquieria splendens Engelm., increase of a population (Harper 1977, Silvertown 1982). Ambrosia deltoidea (A. Gray) Payne, Opuntia engelmannii Salm- Dyck., Ferocactus wislizeni and other woody plants Cacti are admirably suited to an examination of these and characteristic of the Arizona Upland division of the Sonoran other problems. Plant dimensions (stem height, stem di-Desert. Annual rainfall averages 250 mm (10 in), almost ameter, stem or cladode number) can be readily determined, half of which falls during July, August, and September. making it possible to correlate reproductive effort and plantMaximum temperatures in summer frequently exceed 40° size with unusual precision (e. g., Schmidt and BuchmannC (104° F). Minimum temperatures rarely drop below -6° 1986, Bowers 1996). Flower and fruit production of many C (21° F) in winter. Although freezing nights can be com- cacti are not so large as to hinder the monitoring of totalmon in winter, daytime temperatures always rise above 0° plant reproduction on a weekly or seasonal basis. Although C (32° F). 4 Desert Plants 1998 Flower and fruit production. In September 1993, a total ofthat Ferocactus plants invariably flower by the time they 24 Ferocactus wislizeni plants growing on a gentle north -exceed 30 cm (11.7 in) in height; the sample was therefore facing slope were randomly selected within four qualita-limited to plants less than or equal to 30 cm. All plants of tive size classes (cylinders, parabaloids, large spheres, small this size on the survey site were examined for flower buds spheres). The selected plants were tagged, and their heightsand flowers, and the height and diameter of each plant were and diameters were determined. For every plant, the num- measured. ber of ribs, flower buds, flowers, fruits, fruit scars, and aborted buds was recorded. Flower buds and flowers wereFruit and seed characteristics. Mean fruit weight, length, considered to be aborted when they shriveled before anthe- and width were determined from 17 ripe fruits collected in sis or when they abscised without having been fertilized. October 1995. The seeds from each fruit were removed, air The reproductive census was repeated for the same plantsdried, and weighed as a group to the nearest 0.1 gram, then in August 1994, September 1995, and September 1996.mean seed length and width were determined using five Sampling in the latter half of the flowering season made itrandomly selected seeds from each fruit. The average weight possible to count the entire reproductive effort of each plantof an individual seed was calculated from the collective at a single visit except in 1994, when the census was tooweight of 500 randomly selected seeds, then used to esti- early for an accurate assessment of fruit production andmate the average number of seeds per fruit. Total seed flower abortion. weight per fruit, average weight of a single seed, and esti- mated number of seeds per fruit were determined in the Numbers of flowers initiated, fruits set, and flowers aborted same way for 17 fruits collected in November 1995. A were plotted as a function of plant height or diameter, and Pearson correlation matrix was used to analyze the strength curves were fitted to the data points. Percent fruit set, cal-of the associations among the various fruit and seed traits, culated as the ratio of fruits set to flowers initiated, was and independent t -tests were used to compare between months plotted in the same way. Mean values for the four yearsthe weight of a single seed and total seed weight per fruit. were used in all regressions except for those involving fruit data, which were based on three -year means. One -wayResults ANOVA was used to compare flower and fruit productionFlower and fruit production. Of the original 24 plants, 19 among years. All sample plants, whether reproductivelywere reproductively mature in 1993 and 1994. In 1995, mature or not, were included in these analyses. A regres-one mature plant died and a young plant bloomed for the sion equation was used to examine the effect of drought on first time; in 1996, two more plants reached reproductive fruit production. Number of fruits expected on the basis ofmaturity. Over three years, 93% of initiated flowers set fruit plant diameter was determined from the equation, thenon average (range = 91% to 96 %). The sample plants compared in a paired samples t -test with the actual numbershowed no significant difference among years in number observed in 1996, when precipitation on Tumamoc Hill from of flowers initiated (F = 0.09, P > 0.80) or fruits set (F = January to July was 43% of the long -term average. 0.11, P > 0.80). Plant heights and diameters were highly correlated (R2 =0.78, P < 0.001). In linear regressions, plant Minimum Size at Reproductive Maturity. In July 1996, aheight explained a high proportion of the variance in num- survey was conducted to determine plant size at first flow-ber of flowers and fruits initiated (73% and 72 %, respec- ering. The 0.4 ha survey area, located on the north slope oftively, P < 0.001). Plant diameter accounted for an even Tumamoc Hill, included the plot used to monitor flowergreater proportion: 80% for flowers, 79% for fruits (P < and fruit production. Preliminary measurements suggested0.001) (Figure 1).
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