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U. S. Department of ~~rickfture Service Research Paper SO-90

Flowering and Fruiting Of Southern Browse Species

L.K. Halls

Southern Forest Experiment Station Forest Service U. S. Department of Agriculture

Flowering and Fruiting of Southern Browse Species L. K. Halls '

Flowering and fruiting dates are re- old . Nine of each of the 14 ported for 14 browse species growing in the species were spaced equally within each of four open and beneath trees in an east Texas contiguous %-acre blocks. and - forest. Dates for individual were planted 5 feet apart; small trees, 10 feet species generally were not influenced by apart. tree cover. Usually bloomed earli- The same planting schedule and arrangement est when March temperatures were high- were implemented in a nearby abandoned field. est. In the open, plants gene~allyproduced Here the .land was disked before planting, and more consistently and abundantly weeds were controlled by disking until the and at an earlier age than beneath the study plants were definitely established and trees. American beaut ybe~ryand yaupon thereafter by mowing. yielded the most fruit. Honeysuckle and Twenty-four plants of each species ( three yaupon persisted longest through the from each block in the two planting sites) were winter. selected for flowering and fruiting observa- Additional keywords: Phenology, fruit tions. A few of these plants died during the yields, Pinus elliottii, P. taeda. study. The age when plants first bore mature fruits was noted. Thereafter, the seasonal de- Because fruits are a vital food source velopment of flowers and fruits was recorded for wild birds and mammals, wildlife managers from beginning date of growth until final fruit need to know when various plants bear fruits fall the following winter. Samples of mature and how much they yield. This paper describes fruits were collected, dried at 70°C., and the flowering and fruiting characteristics of weighed to compute an average ovendry weight 14 common species of woody vines, shrubs, and per fruit. This figure was multiplied by num- small trees in southern . The inforrna- ber of fruits per plant to obtain the total annual tion is based on observations made at the Ste- yield. Number of mature fruits per gram, mois- phen F. Austin Experimental Forest near Nac- ture content, and ratio of pulp to were ogdoches, Texas, from 1963 through 1972. Cer- calculated when a sufficient sample was avail- tain phases of the study were reported previ- able from 1967 through 1972. ously (Halls and Alcaniz, 1968 and 1972). Temperatures were cor.tinuously recorded by hygrothermographs at both study locations, SITE TREATMENTS and daily rainfall was measured by a standard A sawtimber-size stand of shortleaf (Pinus rain gage nearby. echinata Mill.) and loblolly (P. taeda L.) was thinned to a tree basal area of 70 sq. ft. WEATHER RECORDS per acre. vegetation was cut or killed with chemicals before planting of l-year- Rainfall.-Annual rainfall ranged from a low 1967 66 On the staff of the Wildlife Habitat and Lab- of 32 inches in to a high of inches in oratory, which is maintained at Nacogdoches, Texas, by 1968 (table 1). For 7 years out of 9 it was the Southern Forest Experiment Station in cooperation with the School of . Stephen F. Austin State Uni- below the longtime average of 46 inches. versity. Droughts of several weeks duration occurred in the summers of 1964 and 1965. Total rainfall 1967; June 1971; July 1969; August 1964,1969, in 1966 was very close to the yearly average. and 1970; and September 1971. The 1967 rainfall was 14 inches below average, The date of the last spring freeze varied from and soils were especially dry from June March 9 in 1967 to April 7 in 1971, and the through November. In contrast, rainfall was first freeze in fall ranged from October 19 in much above average in 1968 and through May 1967 to November 15 in 1972. The length of 1969. This wet period was in turn followed by the frost-free season was from 205 days in 1970 a prolonged dry period that extended through to 234 days in 1968. June 1972. Heavy rains occurred in July, and rainfall was above average for the remainder of the year. SEASONAL DEVELOPMENT OF FLOWERS Flowering dates for most species were not Table 1.-Seasonal rainfall (inches) at Stephen F. consistently influenced by tree cover. How- Austin Experimental Forest ever, American beautyberry flowered a few Year I Winter I Spring (SummerI Fall 1 Total days earlier in the open; and flowering dog- , , and common greenbrier flow- ered a few days earlier beneath trees (fig. 1). Flowering of almost all species was earliest in 1967 and 1972, when March temperatures were highest. Generally flowers bloomed late when March was cool; but this relationship was inconsistent, particularly for species blooming in May and June. January and February temp- eratures had no obvious effect on initia- tion, nor did amount of precipitation. In a wet spring some species would flower early and others late. Such diverse responses to the same Temperature.-From 1964 through 1972 the environmental factors have been noted else- average maximum monthly temperature was where ( Blaisdell, 1958). 78.7OF. in the open and 76.g°F. beneath the The interval between first flower emergence trees. Occasionally, however, the maximum and peak of bloom was shortest for rusty black- temperature was higher under the trees. The haw, common greenbrier, and saw greenbrier widest differences occurred in the falls of 1966 (3 to 5 days) and longest for flowering dog- and 1968 and in the summer and fall of 1969, wood plants in the open ( 14 to 16 days). Other when temperatures in the open were 4" to 8'F. species had average intervals of 6 to 10 days. above those in the shade. The highest maxi- In a typical year species bloomed in the fol- mum temperature, 95.0 OF., was reported in lowing : (1) flowering dogwood and August. sassafras, ( 2) rusty blackhaw, red mulberry, The average daily minimum temperature and smallflower pawpaw, (3) yaupon and com- was 52.8"F. in the open and 53.4"F. beneath mon greenbrier, (4) saw greenbrier, dwarf live trees. The lowest mean minimum temperature, , Japanese honeysuckle, and Alabama sup- 35.6OF., occurred in February. plejack, ( 5 ) sweetbay and muscadine In comparison to longtime Weather Bureau grape, and ( 6 ) American beautyberry. records at nearby Nacogdoches, the following Flowering dogwood and sassafras formed months were several degrees colder than nor- in early winter and reached their peak of mal: January in 1964, 1966, 1970; February bloom in late March or early April. Since frosts from 1964 through 1968; March in 1965 and often killed sassafras flowers, only the late 1969; and July and August in 1968. Tempera- blooms developed into mature fruits. Although tures were considerably warmer than average dogwood flowers were often discolored by a in January and February of 1969, 1971, and freeze, the fruits were rarely damaged. When 1972; March 1967 and 1972; April 1965 and fruits set, they generally grew to maturity. For rusty blackhaw, red mulberry, and from 5 to 9 years. Sweetbay magnolia, dwarf smallflower pawpaw: blooming usually peaked live oak, saw greenbrier, rusty blackhaw, and during the first or second week in April. Red muscadine grape plants beneath trees did not mulberry -blossoms were occasionally nipped bear fruit during the study. by late freezes, and only a small proportion of all three species' flowers developed into mature Table 2.-Age at which species first bore mature fruits. fruit Yaupon and common greenbrier flowered Beneath Species open late enough (mid-April) so that they were trees never damaged by a freeze. Yaupon produced - - Years - - abundant flowers; but common greenbrier, Smallflower pawpaw 4 7 only a few. Alabama supplej ack 3 8 Flowering peaked in late April for saw American beautyberry 2 2 Flowering dogwood 4 9 greenbrier, dwarf live oak, Japanese honey- Yaupon 4 7 suckle, and Alabama supplejack. Greenbrier flowers were sparse. Live oak continued to Japanese honeysuckle 3 5 Sweetbay magnolia 6 (I) form new flowers several weeks after the peak Red mulberry 4 '2 of bloom; many of the flowers aborted and Dwarf live oak 4 (I) were replaced by others. Honeysuckle also Sassafras 4 8 formed a few flowers throughout the summer Saw greenbrier 4 (I) and fall if rainfall was above average; these Common greenbrier 6 9 late blooms seldom developed into mature Rusty blackhaw 5 (9 fruits, however. Supplejack flowers were pres-- Muscadine grape 4 only briefly in the spring, and most of them ' Plants beneath trees did not bear fruit by 1972, developed into mature fruit. Sweetbay magnolia and muscadine grape The proportion of fruit-bearers was much flowered in mid-May. The magnolia's sparse higher for plants growing in the open than for flowers. developed into one or two full-size those beneath trees (table 3). Usually this cones and a few immature fruits. Grape flow- proportion increased as the plants grew older; ers were also sparse; and many of the develop- however, this trend was not consistent for sev- ing fruits aborted, especially in hot, dry wea- eral species. On the average, American beauty- ther. and Japanese honeysuckle had the high- Flowering of American beautyberry did not est proportion of plants bearing fruit. peak until June. Although flowers continued Secrson of fruiting.-Usually when a plant to form through summer and autumn when bloomed early, the fruit matured early; but for moisture was plentiful, late blooms seldom several species there was no relationship be- developed into mature fruits. tween dates when flowers appeared and dates when fruit matured. For example, in any FRUITING given year a species might flower earlier than average but bear mature fruit later than usual. Age at first fruiting.-Whether planted in Generally, the fruits of Alabama supplejack, the open or under trees, American beautyberry American beautyberry, and red mulberry ma- first produced fruit at age 2 years. All other tured at least 10 days later on plants beneath species except red mulberry produced mature trees than on those in the open. No consistent fruit at an earlier age in open than beneath difference between locations was observed for trees ( table 2 ) . In the open, sweetbay magno- other species. lia and common greenbrier bore fruit at age 6; and most other species, at 3 or 4 years. Beneath Red mulberry was the only species that bore trees, the beginning fruit-bearing age ranged mature fruit in the spring (May), and the interval from first appearance to end of drop In study the flowering and fruiting dates were earlier for smallflower pawpaw and red mulberry, and the fruiting was only about 6 weeks. Smallflower pawpaw, dates were later for Alabama supplelack and rusty blackhaw sweetbay magnolia, sassafras, and muscadine than the dates presented in taxonomic texts (Vines, 1960; Correll and Johnston, 1910). grape produced mature fruit in summer or JAN. FEB. MAR. APR. M AY JUN. JUL. AUG. SE PT. OCT. NOV. OEC. 5 IS 25 5 I5 25 5 15 25 5 15 25 5 15 25 5 15 25 5 IS 25 5 I5 25 5 IS 28 8 IS 2s 5 15 25 S I5 25 IIIII ~IIII(IIIIIIIIIIIJIIIII I IIII IIIII IIIII II(I1 III1I IIIII IIIII FLOWERING DOGWOOD ICORNUS FLORIDA] 1- 1- 51 ; 4 21 : I OPEN 31 : I 4 ------4+ : I ------lrn 5- 2% $1 4 3m 7- 4) : I

SASSAFRAS (SASSAFRAS ALBIOUM) I- sm 2- 2- OPEN 81 : 4 3- 71 : I ------4) : 1 Im 2m * 5'+- sm WOODS 71 : I 4H.1

------~- - -- I I I - RUSTY BLACKHAW (VIBURNUM RUFlDULUM) 5w-* OPEN 6-

4- 4- LEGEND: ------& FLOWERING f RUITIU6 I 1st EMERGENCE 4 1st APPEARANCE 2 181 FULL BLOOM 5 MATURE 3 PEAK OF BLOOM 6 MAXIMUM DROP 7 END OF OROP RED MULBERRY ( RUBRA) MEAN LRANGE11 ------4- -__------It I 8- 21 : r 6- WOODS 3t : I 7 I 1 4- 4-

SMALLFLOWERPAW~AW ( PARVIFLORA)

OPEN ------:-:-::-::-: 8- * 81 : 1 3CT WOODS 71 : t IF&4m I YAUPON OLEX VOMITORIAI 1- sm I, OPEN C I -LL---1--- sm 7 6- I 3- WOODS 7 4- 4- t~mmIIIBI IIIIIIIIIII IIIII IISII 1111, IIIII la111 IIIIB I~III,IIII&

Figure 1.-Flowering and fruiting chronotogy of plants growing in the open and beneath trees. (* indicates that data are avaikble only for 2 years.)

early fall; but almost all their fruit dropped Seven other species produced fruit both in by October. Fruit persisted through November the 'open and beneath- trees, but yields. from or December on all other species and through woods-grown plants were insignificant. In the most of the winter on yaupon and honeysuckle. open, smallflower pawpaw and flowering dog- In these two species, the end of drop was later wood plants yielded only small quantities of for plants in the open, probably because they fruit in their early years; but in 1971 and 1972 produced much more fruit than those in the their yields were .much higher. Apparently the woods. plants were just approaching their optimum Fruit yields.-In any year the yields were potential then.+.In. contrast, the fruit yields for apt to be high for one species and low for an- Japanese honeysuckle were highest when the other. The period of high production also va- plants were.4 to 6 years old and considerably ried considerably between species. Wright less thereafter. Yields of Alabairia supplej ack (1953) previously noted that there was little were substantial and relatively consistent be- correlation among woody species in the occur- tween years. Although sassafras yields were rence of good seed years. typically small, substantial quantities were pro- duced occasionally. The fruit yields of red American beautyberry yielded more fruit mulberry and common greenbrier were always - .than any other species during this study (table small. Red mulberry. plants beneath trees be- 3 ) ., In the open, plants grew rapidly the first came heavily invested with white scale few years and produced abundant fruit. Yields in 1968 and 1969, and by 1971 all plants had peaked at 1,722 grams per plant in 1966, when died. plants were 5 years old. The following year, Five species produced fruit in the open but however, the plants deteriorated; and many not beneath trees. Yields were never very stems died back. This general decadence con- large for sweetbay magnolia and common tinued through 1969, and fruit yields declined greenbrier. Rusty blackhaw produced small sharply to 107 grams. In 1970 many of the amounts of fruit through 1971, but a large old stems sprouted at the base, and the plants increase in 1972 indicated that the species was again produced a substantial fruit crop. Dur- just beginning to reach its potential. Dwarf ing the next 2 years, severe dieback of stems live oak plants produced moderate amounts diminished fruit yields. In the woods, the of acorns each year, with relatively high yields young plants developed more slowly; and they in 1971; during the last 3 years of the study, produced less fruit. Beginning in 1965 the each plant bore fruit. In muscadine grape, yields tended to be relatively high and low on yields per fruiting plant were fairly consistent alternate years. between years, but only a small proportion of Yaupon had the second highest yield per the bore fruit. fruiting plant; but because this species is dioe- Fruit and seed characteristics.-Table 4 lists cious-i.e,, has separate staminate (male) and the species observed and describes their fruits pistillate (female) plants-the proportion of and . Information on the kind, color, and total plants bearing fruit was relatively small. size of fruits and seeds was taken from taxo- The ratio of fruit yields between open- and nomic texts; data on the number of fruit per woods-grown plants was 19 :1. After plants gram, moisture content, and pulp to seed ratios in the open began to bear fruit (age 4), their were derived in this study. , drupes, yields tended to be high or low on alternate aggregates, cones, and acorns were included. years. In 1972 the yield per fruiting plant was The fleshy berries, aggregates, and berry-like the highest recorded for any species during the fruits had high moisture' contents; for example, study. moisture in American beautyberry, red mul- The alternate high and low yields of Ameri- berry, and muscadine grape averaged about 80 can beautyberry growing beneath trees and percent of total weight. Sassafras had the least of yaupon in the open were apparently unre- moisture content (47 percent); and Alabama lated to weather conditions. Rather, as noted supplejack, the highest (60 percent) for the in other studies (Kozlowski, 1971), the vari- drupes. Only seeds were sampled for dwarf ance seemed to be a physiological characteristic live oak and sweetbay magnolia; their mean of individual plants. moisture content was 45 percent or less. Table 3.-Fruit yields of browse plants growing in the open and beneath pine trees . . Open Beneath trees Species Year Yield per Plants Yield per Plants fruiting plant bearing fruit fruiting plant bearing fruit Grams Percent Grams Percent Smallflower pawpaw

Alabama supplejack

American beautyberry

Flowering d0gwoo.d

Yaupon

Japanese 1965 4 100 0 0

honeysuckle 1966 138 100 0 '0 '

6967 127 100 ' 1 17 1968 222 100 3 42 1969 43 100 5 92 1970 68 100 11 92 ,1971 23 100 6 92 1972 26 100 . 2 92 . Sweetbay 1968 20 42 0 0 magnolia 1969 3 42 0 0 1970 5 27 0 0 1971 16 64 0 0 1972 6 33 0 0

.7 Table 3.-Fruit yields of browse plants growing in the open and beneath pine trees (Continued) Open Beneath trees Species Year Yield per Plants Yield per Plants fruiting plant bearing fruit fruiting plant bearing fruit Grams Percent Gram Percent

' Red mulberry

Dwarf live oak

Sassafras

Saw greenbrier

Common greenbrier

Rusty blackhaw

Muscadine 1967 56 17 0 0 grape 1968 107 18 0 0 1969 388 18 0 0 1970 239 27 0 0 1971 300 36 0 0 1972 149 36 0 0 'All plants beneath trees had died by 1971. Table 4.-Fruit and seed characteristics of 14 woody species

Fruit Pulp /seed Species Moisture Seed Kind Number Content ratio Color Size per gram ( percent ) Mean Range Mean Range Mean Range

Smallflower pawpaw Fleshy berry Black at 5-12 cm. long 62-66 Several, 1.5-3 cm. 1:1.23 I1:1.23-1.24 (Asimina parvif lora) maturity long, bony, turgid, flattened, dark brown Alabama supplejack Juicy drupe Bluish-black About 8 mm. 53-66 A 2-celled stone 1:1.23 1:0.90-1.71 (Berchemia long scandens) American beautyberry Berry-like Reddish-purple 3-6 mm. long 75-84 4, about 1.5 mm. long 1:0,68 1:0.38-0.98 ( drupe wide americana) Flowering dogwood Drupe Bright red 1-1.5 cm. long 52-56 1-2, about 9 mm. long 1:0.96 1:0.85-1.05 f f lorida) to yellow pale brown Yaupon Drupe Shiny red About 6 mm. 48-60 Usually 4 1-seeded 1:0.69 1:0.65-0.76 (Ilex vomitoria) long stones, up to 4 mm. long Japanese Berry Black About 7 mm. in 61-72 Several, irregularly 1:0.35 1:0.27-0.45 honeysuckle diameter ridged ( ) Sweetbay magnolia A cone with Red 4-5 cm. long ' 37-45 Each follicle with ...... (Magnolia drupe-like 1-2 seeds, about 7 mm. virginiana) seeds long, red

Red mulberry An aggregate Dark purple 2-3 cm. long 72-81 1.-2 mm. long 1:0.31 I1:0.26-0.35 f ) of small drupes Dwarf live oak Acorn Glossy brown Cup 10-15 mm. 97-54 15-20 mm. long, 8-12 . . ... (Quercus minima) broad, 8-16 mm. mm. thick, ?4 to H high included in cup Sassafras Drupe, borne Lustrous blue About 1 cm. long 43-50 Solitary stone, light 1:0.73 1:0.60-0.81 f Sassafras albtdum) on red brown, about 6 mm. long Saw greenbrier Berry Glaucous, About 6 mm. 59-69 Usually solitary, 4-5 1:1.59 131.00-3.91 f Smilax bona-nox) black thick mm. long, reddish brown with black basal disk Common greenbrier Berry Blue-black 6-8 mm. thick 58-69 Usually 2-3, about 4-6 1:1.23 1:0.93-1.76 f Smilax mm. thick rotundifolia) Rusty blackhaw Drupe Blue-black 8-15 mm. long 44-58 Solitary, flattened 1:0.50 1:0.41-0.56 () Muscadine grape Berry Purple-black 12-25 mm. in 78-84 2-3, 7-8 mm. long, 4-5 1:0.28 1:0.18-0.36 f Vitis rotunclifolia) diameter mm. thick ' Samples taken only 2 years. ' Includes only the seed portion of fruit. --

The proportion of pulp to seed was highest openings. J. Wild. Manage. 32: for muscadine grape, red mulberry, and Jap- 185-186. anese honeysuckle, and lowest for smallflower Halls, L. K., and Alcaniz, R. pawpaw, Alabama supplejack, and greenbriers. 1972. Growth patterns of -browse These ratios are significant in wildlife manage- plants in southern forests. USDA ment since the nutrient content of seeds differs For. Serv. Res. Pap. SO-75, '14 p. considerably from that of the pulpy and fleshy South. For. Exp. Stn., New Orleans, portions of fruit. Usually the seeds contain La. more crude protein than the pulp. The value of a fruit with a large proportion of seeds de- Kozlowski, T. T. pends mainly on how well the animal digests 1971. Growth and development of trees. the seeds ( Wainio and Forbes, 1941). . Vol. 2, 514 p. N.Y.: Academic Press. LITERATURE CITED Vines, R. A. Blaisdell, J. P. 1960. Trees, shrubs and woody vines of the southwest. 1104 p. Austin: 1958. Seasonal development and yield of native plants on the upper Snake Univ. Tex. Press. River plains and their relation to Wainio, W. W., and Forbes, E. B. certain climatic factors. USDA 1941. The chemical composition of forest Tech. Bull. 1190, 68 p. fruits and nuts f roin Pennsylvania. Correll, D. S., and Johnston, M. C. J. Agric. Res. 62: 627-635. 1970. Manual of the vascular plants of Wright, J. W. Texas. 1881 p. Renner, Tex.: Tex. 1953. Notes on flowering and fruiting of Res. Found. northeastern trees. USDA For. Halls, L. K., and Alcaniz, R. Serv. Northeast. For. Exp. Stn., Stn. 1968. Browse plants yield best in forest Pap. 60, 38 p.