| A unit of Lasting Forests
evolving since March 30, 1999 |
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A Total Forest Management Plan
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| A class project on the quail was developed in 1976. The following unit used the results of that good work and revisions and additions have been made. It is important to evaluate the little change in knowledge now and then and the massive investments made in studies and habitat development that have produced little positive change in the quail populations or in public benefits from the remaining birds. |
The Meadowview Method is being designed to consist of computer aids for a total quail system, providing a profitable demonstration and visitation site, superior wildlife management, uses of GIS and Landsat, dynamic planning and management prescriptions, and research integration.
Context - Taxonomy
Quail-like birds have a long evolutionary history on this continent that dates back at least to the Oligocene. From the Oligocene, an indeterminate quail-like fossil is known adding to the fossil quail genus Nanortyx (Tordoff 1951). Johnsgard (1973) felt it is reasonable to assume quail are derived from cracid-like ancestors of the mid-Tertiary times. In the New World, quail had their center of evolutionary history and speciation in tropical America. The present array of quail indigenous to America north of Guatemala includes 15 species of quails. See the evolutionary tree in Johnsgard 1973.
The general taxonomy of the bobwhite (Colinus) quail can be summarized by the following table.
Table 1. Summary of suggested Galliformes classification.
The subspecies (ex A.O.U. Check-list and Aldrich, 1946) of bobwhite found in Virginia according to Johnsgard (1973:408-09) are two:
"C. v. virginianus: Eastern bobwhite. Resident of the southern Atlantic seaboard north to Virginia's southwest to north central Georgia, southeastern Alabama, and northern Florida," and
"C. v. marilandicus Linnaeus): New England bobwhite. Resident of New England north to southwestern Maine southwest to east central New York, Pennsylvania and central Virginia and south to southern Maryland and Delaware (not in A.o.U. Check-list; part of C. v. virginianus)" (Johnsgard 1973:409).
Bobwhites vary in length from 9.5-10.6 inches. "The sexes are very different in appearance, males having a white eye-stripe that extends from the bill through the eye back to the base of the neck, with brown to brownish black coloration above. The ear region is blackish to hazel brown in males makes this feathering extends backwards below the white eye-stripe and expands under the throat to form a blackish chest collar under the white chin and throat of most races. In the northern population the breast and abdomen in males are irregularly barred with black and white. Females of all races have buffy chins, upper throats, and eye-stripes, and buffy tones likewise replace the white underpart coloration of males. Females also lack black collars and in general are more heavily marked with brown and buff bearing or mottling both above and below" (Johnsgard 1973:410).
Callipepla, with little doubt, is the nearest living relative of Colinus. Holman (1971) indicated that on the basis of skeletal structure this species might be considered congeneric and Johnsgard (1973) has indicated that the same conclusion can be arrived at based on hybridization evidence. Were it not for the taxonomic problems at the species level existing within the bobwhite, this would probably be the best treatment (Johnsgard 1973).
It can be seen that the taxonomy of the bobwhite is far from being agreed upon. Depending upon the author, Virginia has one or possibly two resident subspecies. If Johnsgard's (1973) proposal of the subspecies is accepted, a program is needed to prevent these two subspecies from intermingling and breaking down their species differences which may occur as people turn more and more land to his uses. The physical barriers must be maintained between these two subspecies and human land use should not prevent this distinction being made.
Also, if these subspecies are to be maintained, infiltration of native stocks with "inferior blood" must be prevented. From 1910 to the present, Texas (Mexican) bobwhites have been scattered freely from north to south over the eastern states. In the first 15 years, records of the Biological Survey showed that more than 233,000 Mexican quail were brought into this country (Allen 1962).
"Investigations by the Bureau of Biological Survey and by others have shown that Mexican birds in Pennsylvania mate with, and undoubtedly will eventually change materially the character of the native stock." In 1928 Phillips reported he had "seen a large series of present-day Pennsylvania and Georgia specimens that show various gradation from eastern-looking.to pure texanus type. There is little doubt that the native northern stock will be swamped out" (Allen 1962:187).
Allen stated (1962:187) that "ordinarily, it seems possible that a small infiltration of inferior blood would quickly be eliminated from a population by the selective dying off of weaker birds. But it is possible that particular conditions have altered this."
Allen (1962:188) stated that "Roger N. Latham and C. R. Studholme studied the situation in Pennsylvania and revealed facts that may have key significance. Pennsylvania quail were drastically reduced by the hard winter of 1935-36, and they have failed to recover since then. In the period following the reduction, there were heavy importations of Mexican quail. These, coming at a time when native stock was at its crucial low, would have -had their maximum effort in degrading the northern birds. Latham and Studholme had indications of physical changes which were strongly suggestive that this had happened. If it has, many years of annual turnover and natural culling may be necessary before the population regains its former thrift."
Any suggestion to restocking with non-native stock must be blocked for this reason. If stocking is allowed the bobwhites of Virginia will quickly lose their subspecies characteristics that set them apart from various other bobwhites.
The bobwhite quail is generally distributed over the eastern half of the United States. The range is mapped by Johnsgard (1973:413). The quail is distributed through-out the State of Virginia. The greatest densities of quail occur in areas that provide nesting habitat, protective cover and adequate food throughout the year. These are associated with active farm lands, abandoned farms, and open areas bordering woodlands.
In order to manage properly the bobwhite, it is important that the game manager view this problem multi-dimensionally, noting the environmental constraints and opportunities of time, space, and diversity. In the past, educational practices have been directed as a guide to the landowner to execute certain land practices to increase bobwhite populations on farmlands. Some techniques that can be employed for enhancing the importance of the bobwhite quail as a nongame species have been suggested. Special television and radio programs on the bobwhite's life history, population dynamics, and song could serve to promote the aesthetics of this popular bird. County maps indicating quail range and density as well as management area locations could be made available. Articles on artificial feeding, game bird recipes, field trials, and information on other social and economic values could be published in magazines, brochures, booklets, and special management publications. The public can be further educated to use land-use practices beneficial to quail. Other techniques for importing such knowledge include news releases, speeches, and public announcements. An objective function could be devised noting the major assets of the quail [in order of their importance. This technique could serve as a valuable aid in helping the wildlife manager know what factors of the bobwhite are of greatest significance to the public. Steps could then be taken either to ameliorate management conditions to obtain desired effects, or to explain to the public why certain procedures cannot be administered.
The Range of Objectives:
Wildlife management involves simultaneously maximizing a wide range of benefits, from metaphysical, esoteric and non-consumptive benefits to physical, exotic, consumptive, and experiential benefits. The scope, variety, amounts (amplitude), diversity and personal range of experience of the wildlife resource user benefits must be considered (Giles 1975). Region-wide management of a natural resource, such as quail, should be constructed around a system of owner- and citizen-weighted objectives. The objectives of the management should be to maximize the achievement of these objectives.
The management also should provide the opportunity for citizens to achieve their weighted set of quail resource objectives. Over the past 50 years there has been a perceived reduction in quail populations of -2.5% per year. The causes are under study. An important policy issue is that of access to private land for hunting and /or observing quail. If the public does not have access, there may be little incentive for management efforts to sustain or increase the bird population. Without birds, hunting experience and incentives decline. The results is a declining system, devoid of useful feedback.
The set of objectives for a quail-oriented system are suggested below:
A. Attainment of natural completeness of the system as it relates to rural and urban wildlife and people using the resource.
B. Aesthetic values of a natural resource.
C. Temporal Insight (the historical and evolutionary messages of wildlife).
D. Production of a food resource.
F. Pride of possession.
F. Pride in preservation of and tending action toward quail habitat.
C. Personal economic gains (e.g., profits from selling meat, eggs, and operation of private hunting preserves).
H. Corporate or financial gains (e.g. selling of firearms and ammunition, housing and food).
I. Community and government agency gains as from hunting seasons.
J. Environmental system support utility (minimizing ecosystem entropy).
K. Utility of resource opportunity on quail
These objectives must contain flexibility to achieve the changing desires of the consumers. Without this flexibility the system could collapse.
Bobwhite quail probably remain the most important game birds in the Southeastern United States and are the third-most popular species of small game. It ranks behind the squirrel and rabbit in this respect (Environmental Research Group 1974). In terms of recreation, quail are important Virginia songbirds, enjoyed by many throughout the state. Also, quail hunting provides thousands of hours of recreation for Virginia hunters. The sale of guns, ammunition and equipment for quail hunting is an important economic factor.
In a sample of households, 25 percent indicated small game hunting was an ongoing recreational endeavor. Fifteen percent of the households participated in big game hunting while only 4 percent participated in waterfowl hunting. Thus, quail hunting seems to be quite important) based on the number of small game hunters statewide (Environmental Research Group 1974). A Virginia Game Commission questionnaire taken in 1968-69 showed 132,510 quail hunters which was 43.2 percent of the total number of hunters in Virginia at that time (i.e., 306,247).
To pursue the pleasure of quail hunting 669,807 trips were taken, an average of 5.05 trips per hunter. The quail harvest for the season was 1,178,720. The average number of quail taken per trip was 1.75 with 8.9 quail taken per hunter.
It is estimated that 2 billion dollars are spent yearly for small game hunting with the average dollar value--per -day -of small-game hunting being $39.14. Most households have said they would have to receive $10 - $25 to give up a day of hunting, which shows one measure of the value hunters place on their sport.
To hunt, bird-watch, camp or participate in other wildlife recreation, a person typically foregoes a day of work, or takes a vacation day valued at least as much as the value of a day (or hour) of work. The value of hours is estimated as:
Value = Income/(working days) (16 working hours)
then expanded by computing and adding hunter-specific income and vacation time. Thus a minimum estimate can be obtained.
INPUTS
Energetics
Beyond the basic energy requirements for maintaining basal metabolic rate, bobwhite quail require energy (productive energy) for a variety of essential activities such as food-gathering, predator evasion, social behavior, molting and reproduction (Robel et al. 1974). In developing a management plan for a species such as the bobwhite quail consideration of the factors affecting the energy budget of the species is of utmost importance.
Although energy is acquired through one basic mode (ingestion of food) it is utilized or expended through diverse physiological and behavioral activities. Both the energy gain and drain of a species are affected by a variety of environmental factors. Additionally this gain and drain rate does not remain constant throughout the year but varies as the environmental components (particularly climatic components) vary.
In a laboratory study of the bioenergetics of bobwhite quail Case et al. (1974) found that energy requirements decreased in a linear manner as the temperature was increased from 0 to 400 degrees C. This would substantiate the idea that the energy requirements of bobwhite are greater in cold weather (e.g., winter) than in warm weather (e.g. summer). However, it was also found that increasing the photoperiod increased the energetic drain of bob-white quail. The highest efficiency (76 percent) was the 10-hour photoperiod.
Sex was not found to significantly affect the energy requirements except in the female during egglaying. Energy requirements of an egglaying female decreased quadratically with increased temperatures (Case et al. 1974)
Several researchers (Robel 1973; Case et al. 1974) noted behavioral modifications of quail for maintaining higher temperatures, most notably huddling, which, at 50-C., provided a metabolic advantage but was energetically disadvantageous at higher temperatures (Case et al. 1974).
Food Habits
The development of a strong, stable, bobwhite population is very dependent on a few, particular, limiting factors. Of these, food is one of primary importance and thus an adequate food supply must be available during all seasons, especially in winter and early spring. In winter, undernourished birds are not able to cope with harsh conditions and in spring, quail require a large supply of food to coincide with the reproductive season and moulting (Massey 1938).
Table 2 shows the occurrence of foods by month in 1659 adult bobwhite stomachs taken throughout the southeast (Trippensee 1948). This table conforms to the general food-habit pattern of the bobwhite, including other parts of its range. Using this analysis, Massey (1938) posed a mere graphical representation of food preferences throughout the year. (Fig 3).
To be re-entered
Quail foods can be divided into approximately six major categories: miscellaneous seeds, legumes, grasses and sedges, mast, fruits, and animal foods. These six comprise about 93 percent of the total food intake by a quail per year. The graphs represent the variation in uptake of these categories. They are organized to begin with winter season and they run through a 12 month period. The numbers on the left express the percentage by bulk of the material found in the stomachs of 1,659 birds.
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| Figure 3. Feeding habits of the quail in Virginia and the Southeast (from Massey 1938:785; graphs: top-to-bottom 1 to 7). |
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Graphs 2, 3, and 5 in Fig. 3 show significant increases in the uptake of fruits, grasses, and sedges, and masts concurrent with increases in the availability of these food sources. Of particular interest are graphs 4 and 6 which show increases in uptake which do not coincide with periods of increased availability. It would seem that immediately after moults, the bobwhite quail reacts to the demand for high protein food sources by consuming more insects than vegetation. The poults do in fact feed almost exclusively on insect life (Martin 1951). The peaks of high intake of legumes is a means for the bobwhite to fulfill the high protein needs of the approaching laying season. A more extensive breakdown of summer-food habits can be found in Table 3 (Allen et al. 1945), and of winter habits in Table 5 (Baldwin et al. 1946).
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| Fig. 4. Virginia showing county divisions and arbitrary delineation of the three natural regions related to quail foods. |
In Virginia, the major components of food supply must be divided into three geographical regions. The Coastal Plain, the Piedmont, and the Mountain Section (See Fig. 4). These three divisions are based on a compromise between forest type and soil type. The Coastal-Plain is largely devoted to growing peanuts, cotton and truck crops; the Mountain Section is used for livestock -and general farming. Vast differences in winter food uptake of native and naturalized legumes, fruits, mast, grasses and forage can be attributed to the variations occurring between these geographical regions (see Tables 4, 5, and 6)(Baldwin et al. 1946).
Contrary to some beliefs, bobwhite quail do not require gravel in their crop for proper digestion. Seeds such as wheat, millet, milo, soybeans, field peas, and vetch can be successfully macerated and digested without the aid of grit. When very hard seeds are ingested in the wild, apparently the seeds themselves serve in the capacity of grinding agents (Nesther 1946).
| Kind of Seed | Pounds of seed per acre on the cut and burned area |
Pounds of seed per acre on the untreated area |
| Legumes Partridge Pea (Cassia nictitans) Beggarweed (Desmodium spp.) Korean lespedeza (planted L. stipulacea) Native lespedeza (Lespedeza spp.) Milkpea (Galactia spp.) Grasses Ragweed (Ambrosia spp.) Other Total |
16.4 5.4 3.5 0.9 2.7 3.0 0.2 0.9 11.9 44.9 |
0.2 0.4 0.1 0.5 0.1 4.5 5.6 |
Cover Requirements of Bobwhite Quai1
The quail is primarily a bird of farmlands, thickets and open woodlands, rarely occurring in great numbers in heavily timbered lands devoted to the production of forest crops. In the Southeast, quail may thrive under a canopy of forest trees, if the forest is not too dense and the correct interspersion of cover types are maintained. They do occur in early stages of pine clearcuts and re-planting. This correct proportion, as reported by Stoddard and Komarek (1941) should be about 25% of the entire area distributed in small fields.
Leopold (1933) suggested that quail require four vegetative types: woodland, brushland, grassland, and cultivated land. For all to be maximally used they must meet at a common point which is the focal point of a covey's activities. Quail do well in primitive style farming situations where weedy fields, brushy fence rows, and odd corners supply an abundance and variety of suitable cover. In contrast, 'cleaner' farming practices which eliminate brushy fence rows are detrimental to bobwhite numbers.
The above set of "needs" must be managed, eventually over a landcape, as analogous to a timber rotation. As tracts age and go out of superior conditions, new tracts must be prepared and begin to produce for quail. Advanced vegetation (e.g., the pine plantation) is moved to a condition of low or no production of quail until is harvest date.
Early thicket or brush succession, some woodland, weed succession and grain provide good cover and food habitat for bobwhite quail. It is also beneficial to have a small acreage of marsh border, especially where these include smartweed (Polygonum .).
Cover, in general, is an important factor in expanding the range of bobwhite quail. The growing of hedges and windbreaks and the introduction of grain crops were sufficient to extend the range of the bobwhite westward into the treeless plain states. The cutting of woodlands and the development of agricultural crops also served to extend the bird's range.
Cover as a general term can be divided into more specific areas, these being: feeding cover, escape-cover, shelter or winter cover, nesting, and roosting cover. Each subdivision is an important and integral part of the whole, and these requirements must be satisfied if the population is to be maintained. All forms of cover must come together to be connected in series in order to be of use to bobwhite quail. Each subdivision need not be a separate entity unto itself.
Feeding cover is furnished by standing corn, cane, sweet clover in fields, shrubbery, wild grape, greenbrier, and other tangles along ditch banks, fence rows, ponds and streams. Significant feeding cover must noted for the chicks. The broods need vegetation that produces abundant insects ... every year. The no-till crop practices seem to produce abundant insects and to significant reported direct poisoning of birds. Chicks (10 day old) eat 5 grams of mass per day to gain 1 gram of mass per day. They forage for 8 hours. Poor correlation has been found between sweep-net samples of insects and insects eaten per unit time by chicks. Using biomass per sweep (with red and inedible arthropods excluded), the condition of areas can be standardized for comparisons. There have to be insects! Debating the number, species, nutrition per insect, or actual number consumed is an exercise that denies the between area, between year, between site and sampling variability that exists. Those doing sweeps add to the variability. A gross approximation of insect biomass probably available for the chicks is needed to compare areas and treatments with observed birds.
Escape cover is necessary for the survival of the bobwhite against house-cats, sharp-shinned hawks and other predators. In order for a range to be highly productive of bobwhites escape cover is essential. A denser cover than can be normally frequented is preferred for purposes of predator escape. Brush piles, tangled thickets, and fallen trees with untrimmed branches meet these requirements (Trippensee l948). Wild plum, rose blackberry, raspberry and other briars form excellent tangles and thickets which are effective predator repellants (Terrill, no date). Mowed walkways are all that are needed to allow the birds in these areas produced at high costs to be observed.
Winter shelter is necessary for protection against snow, sleet and wind. This may be furnished by heavy timber, brush piles, closely set thickets, windbreaks, hedgegrows and low-branching evergreen trees (Terrill, no date). Examination of flush data by Yoko (1972) illustrated that honeysuckle is important to quail during the winter.
Nesting cover is essential for establishing nesting sites. Johnsgard (1973:420) found these nesting requirements:
Roosting cover is required by bobwhites to maintain body heat during the winter for survival. Johnsgard (1973:419) noted these requirements and site differences for bobwhite roosting:
Casey (1965) noted that there appeared to be a headquarter concept for bobwhites that related to the amount of light present and activity. His observations are as follows:
It has been found that the number of coveys a given area could be expected to support was determined by the number of covey "headquarters" that occur there. To be habitable, a headquarters area had to have protective vegetation that was dense enough at the critical season (after the leaves of deciduous vegetation had fallen) to reduce the amount of incidental light at the birds' level to an amount less than 1000 ft. candles at midday.
He found daily movements are a function of light intensity. On sunny days the birds would loaf under cover, intolerant of strong light.
Management Inputs
In dealing with the management of the bobwhite quail, two approaches are often discussed: (1) Increasing numbers of resident populations by introducing artificially reared birds and (2) manipulating the habitat through food plantings, sharecropping-burning, and burning. (Note that benefits are assumed to be directly related to the number of birds. It may be that rarity of birds seen increases the value to people. Managing for bird-related total benefits (or opportunities) per unit cost may be the objective under continual review.)
To evaluate more efficiently the use of state funds for these procedures, each of the above methods will be examined on a cursory economic basis.
"Stocking" of game birds is often a rapid means of increasing natural populations. This is often thought by hunters to be the best means of providing game. However, the economic profile of this procedure reveals it to be expensive. One estimate of production of pen-raised Colinus ranges between $0.68 and $3.00 per bird released. However, the return rate for these birds in the hunter's bag was only 1.21%. This makes the cost of each bird shot around $57.85 each (Murray and Frye 1957). In addition, Phelps (1948) reported poor success of stocking programs of bobwhite quail in Virginia. Because of the expense of restocking quail, the current trend is for eliminating this procedure in the east and southeast (Frye 1961).
Much discussion has been devoted to habitat improvement as a means to increase resident populations of quail. Three specific methods of habitat management will be examined: (1) wildlife food plantings, (2) sharecropping-burning, and (3) burning.
The costs of food planting as a means of habitat improvement for quail include many factors. The cost of clearing the land is a major expense (costs reported here are from 1976).
Clearing land for food plantings by mowing or by a circular brush cutter averaged $5 an hour in one study (Ellis et al. 1969). Clearing land with a bulldozer costs around $15-20 an hour or about $90 an acre (Ellis et al. 1969; Trumbo and Chappell 1969). The most expensive method of clearing was the use of herbicides which range from $36.54 (2,4,5-T) to $106.50 (Fenuron) an acre (Trumbo and Chappell 1969). Additional costs of the food patch arise from the cost of the seed. Average cost (including planting) for multiflora rose, Sericea lespedeza, and grass for an acre is around $30 (Burger and Lindusko 1967). Another study showed that plantings may cost from $12 to $45 depending on the seed used (See Table 7) (Smart et al. 1972). Often fertilizer must be used to boost production of food plantings. Ellis et al. (1969) recommend a mixture of lime and P204 which costs an additional $32 per acre for soils of average fertility.
In general, the use of wildlife food plots is an expensive investment for two reasons. First of all, the initial investment in clearing, seed, and fertilizer is considerable. Secondly, these areas have a high cost of maintenance because they are often small, scattered tracts. Ellis et al. (1969) estimated that the cost of management per acre is $4.57 over a large area. The cost of this procedure is $22.85 per bird harvested (See Table 8). The actual cost of the land, even the annual taxes per acre are rarely included in valuation of land used for the quail.
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| Changes in quail populations over time can be confusing. A - quail/hectare of potential range; B Quail sightings. At one point in time there may be two obeservations |
Another method of habitat improvement for increasing natural quail populations involves the use of sharecropping and burning. Sharecropping is the voluntary gift of the rental of agricultural lands to a game agency for the purpose of food planting. Burning was restricted to wood margins and scrub areas adjacent to the sharecropped land (Ellis et al. 1969). Ellis et al. (1969) found this combination of land use to be the cheapest method of habitat improvement at a cost of $l6 per acre over a large area (See Table 7).
The cost of management for each bird harvested with the sharecropping-burning method (assuming 50% harvest) was $0.32 (See Table 8) (Ellis et al. 1969).
Table 8. Approximate costs of producing harvested quail under three systems of management employed on the Forbes and Dale areas. See page 20 for table
The third method of habitat management for the quail which is often used is simple burning of wooded and scrub areas. Burning was considered to be the most economical means of habitat improvement in 23 quail producing states (Frye 1961). This study also estimated that prescribed burning costs about $1 per harvested bird. Other studies arrive at a cheaper figure. Burning In large areas of the southeast (cost $0.10 an acre, but a more reliable~ figure of $0.50 an acre was reported in most areas (Ellis et al. 1969). The cost of quail harvested under this program of management averaged $0.66 per bird, assuming 50 percent harvest (Ellis et al. 1969).
Habitat improvement by any of the above means has a longer effect on the size of the resident population than stocking programs. The cost per quail harvested of habitat improvement methods are considerably below the costs associated with stocking programs. However, there are underlying costs to habitat management which have no uniform value from year to year. Some of these costs are for maintenance of access roads, parking lots, drainage systems, hunter control and public relations. These factors should be added to costs of the procedures.
Frye (1961) stated that 23 state agencies responsible for quail management found that it would cost between $l.00-$5.00 per bird to double the quail in any available region. Comparison of the three land management procedures (Table 9) showed that quail are most economically produced with a sharecropping-burning procedure. The least economical of the three procedures is the food patch method. Oddly enough this is the most frequently recommended method (Ellis et al. 1969).
Table 9. Estimated costs of managing a 250-acre tract on the Forbes area for bobwhites under three systems of habitat management (Ellis et al. 1969). Page 21
There is considerable difficulty in evaluating the cost benefit of the land management for the quail. The difficulty lies not in the production of quail, but other "benefits" resulting from habitat management for a single species.
A list of possible management activities for selection, site by site:
PROCESSES
Hunting Phenomena
Sport hunting means legitimate harvesting of surplus wildlife following accepted sporting and safe practices. To regulate that harvest all birds and mammals receive some degree of legal protection.
Mangold (1951) showed that fall and winter mortality ranged as high as 88 percent of a population of bobwhite-quail, but hunting accounted for only 14 percent of the loss. Kozicky (1952) concluded that where mortality ranged from 20.3 to 87.8 percent, hunting was not a limiting factor.
For maximum harvest by the hunter, the quail season should be open as soon as a large proportion of the birds (80 percent) are of "unsatisfactory" weight (acceptable to the hunter). Thus fewer birds are lost for reasons other than hunter's harvest. Opening the season earlier, however, would result in a larger proportion of underweight birds in the hunter's bag (Haugen 1957).
Ripley (1957) demonstrated that about one-third of the total bobwhite harvest is realized during the first few days of the hunting season. Hunting pressure, and resultant harvest, are higher on Saturday. Weather conditions also obviously effect the daily kill.
Hunting success can be used as a parameter of population size if seasons are set in accordance with population excess and hunting pressure is constant (Kabat 1963).
Hunting results can also be used to estimate mortality. When birds are shot throughout the period from September to March adult to juvenile proportions can be set on a monthly and yearly basis. Proportion of adults is greater in spring than in fall and reflects mortality trends (Robel et al. 1970).
Habitat Manipulation
There are several ways in which the habitat may be manipulated to sustain quail. Among these are planning, plot planting, burning, timber cutting, and restocking. The most important seems to be plot planting and plowing. Fenced plots of one quarter to 2 acres in size, located 300 yards apart along trams or roads, in the edges of fields, or small clearings, are very successful if they contain a clump of blackberry, rattan, myrtle, yaupon or other year-round-cover. Shallow plowing or disking every two years is an effective means of providing food if brush and trees do not shade the plowed area.
Timber cutting, although not as successful, may be implemented. Extensive management can be exercised by lumber companies to rotate cutting of strips or blocks so that no more than 1/4 of the total area is logged in 10-12 years. Such cutting allows quail access to developmental or early intermediate states of succession. This is important since the objective of quail management is to offer well-distributed spots or strips of early succession vegetation, so that an adequate food supply (especially doveweed) and cover is available. Rotational cutting can provide acorns, pine mast, and various fruits, seasonally (Lay 1940). A questionnaire given to 23 states revealed that (Frye 1961) more than 90 percent of the bird harvest resulted in quail that occur with no effort from management. A persistent question is: Is management really ineffective or does a relatively small proportion of quail represent the harvest?
Population Analysis
The bobwhite quail is strictly a monogamous species. Since the sex ratio of other monogamous gallinaceous birds at hatching is 1:1 it is assumed that the same holds true for the bobwhite. However, in the adult population the sex ratio slightly favors the 113:100 males. This slight excess of males is beneficial since the bobwhite will re-nest. These excess, active males in-sure that all renesting females will be fertilized. Additionally, these excess males can be selectively harvested without significantly reducing the reproductive potential of the population (Johnsgrad 1973).
The maximum bobwhite density is generally agreed to be 1 per acre (Leopold 1933 and Stoddard 1932). Undeveloped preserves may reach one bird per 2 acres and under less favorable conditions, one to 4 or 5 acres. Lighter concentrations are the rule, however (Stoddard 1932). Some speak of high populations as being 10 flushes per day.
Age ratio data are valuable in estimating the nesting success of the past breeding season and the breeding potential for the following year (Johnsgrad 1973). Hendrickson (1945), Leopold (1937), and Emlen (1936) found age ratios of approximately 1:2 to 1:4 (adults:juveniles).
Factors Affecting Populations
Fluctuation of quail populations from spring to the following spring is well documented. Various factors contribute to the reduction of quail numbers.
The greatest mortality factor is natural enemies (Stoddard 1950). Cooper's and sharp-shinned hawks are examples (Trippensee 1948).
Nesting success has been reported to be as high at 91.8 percent but there is a nest loss of 30.3 percent due to abandonment and 57 percent nest lost related to predators (Klimstra 1950). Of those nests that are successful, 13 to 16 eggs hatch but this is reduced to 8.5 chicks by October-November (Johnsgard 1973). Dickson (1974) indicated that after initial brood mortality there was a consistent 3 percent per week loss and the average loss from September 1 to November 1 was 2 percent per week.
The major shift in the population structure occurred between the months of November and January. With an initial population of 88 percent juveniles and 12 percent adults the following spring population was 74.7 percent juveniles and 25.3 percent adults.
Quail have an average life span of one year with few birds surviving to breed the second year (Johnsgard 1973). Healthy, well fed birds with-stand the rigors of winter given almost any conditions although winter losses can be as high as 80 percent (Errington 1939). Winter losses are due to persistent adverse conditions or brief severe conditions (Errington 1939). Once the quails' weight has dropped to about 65 percent of normal, any extended or intermittently severe weather prevents the birds from finding adequate food to increase their weight (Errington 1939).
Losses are also associated with disease and parasites although other factors must contribute to the degrading of the birds' health before these factors are a problem since a well-fed bird can survive attack such as this (Stoddard 1950).
Law Enforcement
An optimum habitat cannot yield good fishing, hunting, or trapping with-out adequate regulation and enforcement (Siegler 1972). Damage to property by hunters and abusing landowner rights can cause problems for conservation officers. Twenty-four out of 42 states charged game wardens with some enforcement of trespassing laws (Siegler 1972).
Despite it being costly and time-consuming, undercover work is often effective against illegal activities (Ritter 1975).
A 14-month investigation revealed that men were arrested on 54 separate charges (Ballew 1971). The public had begun to cooperate in reporting violations to wardens (Ritter 1975).
Eisner (1972) has suggested that the computer programming could be utilized to calculate optimum warden and enforcement levels for areas through time.
Quail hunting violations are relatively few except for the above trespass cases. Quail in good cover cannot be over-harvested.
The primary law enforcement needs are to assure license purchases, to assure equitable distribution of the harvest, and to assure property protection. Agents' educational activities are likely to enhance quail values for other users.
Land Use Effects
There are many factors relating to the carrying capacity of the environment for the bobwhite quail. The use of land for farming or for woodland are two extremes of land use. Farmlands, for example, support different numbers of quail (N. C. Wildlife. Res. Commission) than woodlands. Controlled, periodic burning can increase food plants for birds (Kabat and Thompson 1963). The numbers of quail are affected by the soil fertility as well as by the amount of cover provided by the land. Quail tend to be distributed away from more fertile soil where there is less human activity in farming (Bennitt 1951). In areas where humans and birds interact most closely, the population can be changed by providing cover in the form of hedgerows (Kabat and Thompson 1963), rough field edges, and managing fields for high insect populations for chicks..
Mortality
It is important to remember that despite hunting pressures quail have the ability to spring back during the breeding season (often 3 clutches per year). Yet, annual fluctuations indicate that mortality is regulated by several interacting factors. In controlling mortality the manager is faced with the environment. For example, work done by Raitt and Genelly (1964) showed that the lowest mortality rates (40 percent) occurred during winters with the lightest rainfall. The highest mortality of quail was evident in the winters with the highest rainfall (81 percent mortality). Stoddard (1950) noted that quail losses, especially young, are due to excessive rains which tend to chill the young who never really recover. Mosby and Overton (1950) collected data at the VPI farms which indicated that the fall population which survived to the next spring was inversely proportional to the number of days in which snow of one inch or more covered the ground.
Reproduction in the quail does not generally increase in great numbers after a harsh winter. Instead, the habitat quality regulates the rate of quail increase. This fact has been attributed to vitamin A or carotene, a precursor of vitamin A, found in green foliage. Hungerford (1964) found that as the habitat quality improved, the development rate of the reproductive glands of both sexes of quail increased. This increase is correlated with an increase in breeding rate. Yet, when habitat quality was down, the reproductive rate would be almost zero. Analysis showed that vitamin A or carotene was responsible for increases in gland weights.
A possible means of population control seems feasible through the use of vitamin A pellets. Naturally, climatic events cannot be regulated but introduction of vitamin A to the diet of quail would increase the clutch size (Hungerford 1964). However, vitamin A as an inducer for breeding must not be utilized unless habitat improvements accompany application.
In order to manage the bobwhite properly, it is important that the game manager view this problem multi-dimensionally, noting the environmental constraints and opportunities of time, space, and diversity. In the past, educational practices have been directed as a guide to the landowner to execute certain land practices to increase bobwhite populations on farmlands. Some techniques that can be employed for enhancing the importance of the bobwhite quail as a non-game species have been suggested. Special television and radio programs on the bobwhite's life history, population dynamics, and song could serve to promote the aesthetics of this popular bird. County maps indicating quail range and density as well as management area locations could be made available. Articles on artificial feeding, game bird recipes, field trials, and information on other social and economic values could be published in magazines, brochures, booklets, and special management publications. The public can be further educated to use land use practices beneficial to quail. Other techniques for imparting such knowledge includes news releases, speeches, and public announcements. An objective function could be devised noting the major assets of the quail in order of their importance. This technique could serve as a valuable aid in helping the wildlife manager know what factors of the bobwhite are of greatest significance to the public. On private lands, the objectives of the landowner are important, but ownership, farm size, and agricultural and pricing policy can change quickly. Retaining some uses of the population and a little land use flexibility may not be a bad idea. Steps could then be taken either to ameliorate management conditions to obtain desired effects. Often objectives can be achieved by explaining to the public or landowner why certain procedures cannot be administered and desired results not achieved.
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A rabbit and quail production modifier.
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