Copyright (c) 2008 University of Nebraska - Lincoln All rights reserved. http://digitalcommons.unl.edu/bioscigrouse Recent documents in Grouse and Quails of North America, by Paul A. Johnsgard en-us Sat, 10 May 2008 02:30:32 PDT 3600 http://digitalcommons.unl.edu/bioscigrouse/40 http://digitalcommons.unl.edu/bioscigrouse/40 Thu, 08 May 2008 11:32:39 PDT 1-4. Sage Grouse 5-12. Blue Grouse 13-17. Spruce Grouse 18-22. Willow Ptarmigan 23-26. Rock Ptarmigan 27-31. White-tailed Ptarmigan 62-65. Ruffed Grouse 66-69. Sharp-tailed Grouse 70-79. Pinnated Grouse 80. Buffy-crowned Tree Quail Habitat 81. Bearded Tree Quail 82-84. Barred Quail 85-87. Scaled Quail 88. Mountain Quail 111. Hybrid Barred × Scaled Quail 112. Hybrid Mountain × California Quail 113. Hybrid California × Scaled Quail 114. Hybrid Scaled × Gambel Quail 115. Hybrid Bobwhite × Gambel Quail 116. Hybrid Bobwhite × California Quail 117. Hybrid Bobwhite × Scaled Quail 118. Elegant Quail 119-20. Gambel Quail 121-22. California Quail 123-26. Bobwhite Quail 127-29. Black-throated Bobwhite 130. Spotted Wood Quail Habitat 131-32. Spotted Wood Quail 133. Singing Quail Habitat 134-35. Harlequin Quail 136. Ocellated Quail 137-38. Gray Partridge 139-40. Chukar Partridge Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/39 http://digitalcommons.unl.edu/bioscigrouse/39 Thu, 08 May 2008 11:29:40 PDT 32-35. Sage Grouse 36-39. Blue Grouse 40-43. Spruce Grouse 44-46. Willow Ptarmigan 47-49. Rock Ptarmigan 50-51. White-tailed Ptarmigan 52-54. Ruffed Grouse 55-58. Pinnated Grouse 59-60. Sharp-tailed Grouse 61. Downy Young of Grouse and Partridges 89. Long-tailed Tree Quail 90. Bearded Tree Quail 91. Mountain Quail 92. Barred Quail and Scaled Quail 93. Elegant Quail 94. Gambel Quail 95. Scaled Quail 96. Gambel Quail 97. Hybrid Gambel x Scaled Quail 98. California Quail 99. Bobwhite Quail 100. Spotted Wood Quail 101. Masked Bobwhite 102. Black-throated Bobwhite 103. Buffy-crowned Tree Quail and Singing Quail 104-6. Harlequin Quail 107. Chukar Partridge 108-9. Gray Partridge 110. Downy Young of Quails Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/38 http://digitalcommons.unl.edu/bioscigrouse/38 Thu, 08 May 2008 11:26:52 PDT Index of vernacular and scientific names English vernacular names indexed here are for the most part those used in this book for species or larger groupings. Vernacular names for subspecies as well as alternative vernacular names for species are included only if they are in general usage or have been referred to in the text discussions. Plates and figures are identified by number, and pages containing major discussions of each species are indicated by boldface. Scientific Names: Names indexed here are restricted to those of subspecies, species, or larger groupings of galIinaceous birds mentioned in the text. Technical names of other animal groups and plants are not indexed. Entries shown here are for the major page references; the index to vernacular names should be consulted for secondary references and references to illustrations. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/37 http://digitalcommons.unl.edu/bioscigrouse/37 Thu, 08 May 2008 11:26:50 PDT Bibliography; more than 600 works cited Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/36 http://digitalcommons.unl.edu/bioscigrouse/36 Thu, 08 May 2008 11:24:48 PDT From: Alectoris--from Latin alector (Greek alectryon): cock ... through: Tympanuchus--from Greek tympanon: a drum Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/35 http://digitalcommons.unl.edu/bioscigrouse/35 Thu, 08 May 2008 11:23:14 PDT The three following keys can be used to identify unfamiliar species of North American grouse or quails that may be examined in the hand. Unless one is certain that the bird represents either a grouse or a quail, he should begin with the first key. The procedure, as in the use of all such keys, is to choose which of the two initial alternative descriptive couplets (A and AA) best fits the unknown bird. Having chosen one of these, proceed to the choice of couplets (B and BB) occurring immediately below the couplet chosen, without further regard for descriptions listed below the rejected alternative. After making a varying number of such choices in the first key, the reader will have identified his bird as to its major taxonomic group (family, subfamily, or tribe). If it belongs to either of the subfamilies of grouse or quails, the following two keys may be used to identify the specimen as .to its species. These two keys operate in the same fashion as does the first, by the reader starting again with the choice of couplets A and AA and proceeding until the bird has been identified as to species. Measurements, where they are given, refer to adult birds, but in general the keys have been devised in a manner that will allow for identification regardless of the specimen's sex or, within limits, its age. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/34 http://digitalcommons.unl.edu/bioscigrouse/34 Thu, 08 May 2008 11:23:13 PDT Alectoris chukar (Gray) 1830 Alectoris graeca (Meisner) Other vernacular names: Chukor, Indian hill partridge, rock partridge (refers to graeca only). Range: Native to Eurasia, from France through Greece and Bulgaria (typical graeca) southeastward through Asia Minor and southern Asia (typical chukar). These two populations should probably be regarded as separate species (Watson, 1962a, b), and all of the introduced United States stock is apparently referable to A. chukar. The racial origin of the birds introduced into North America is varied and includes not only Indian stock (probably A. c. chukar, as recognized by Sushkin, 1927) but also some Turkish stock (cypriotes or kurdistani). These Turkish birds probably merged with Indian stock or have disappeared, except in New Mexico and California. The present range of the North American population is from southern interior British Columbia southward through eastern parts of Washington, Oregon, and California to the northern part of Baja California, and east in the Great Basin uplands through Nevada, Idaho, Utah, western Colorado, and Montana, with small populations of uncertain status in Arizona, New Mexico, western South Dakota, and southern Alberta. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/33 http://digitalcommons.unl.edu/bioscigrouse/33 Thu, 08 May 2008 11:21:44 PDT Perdix perdix (Linnaeus) 1758 Other vernacular names: Bohemian partridge, English partridge, European partridge, Hungarian partridge, Hun, Hunkie Range: Native to Europe and Asia but introduced into North America and now widely established in southern Canada and the northern United States (see distribution map). The North American population was probably derived from stock representing several different geographic races. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/32 http://digitalcommons.unl.edu/bioscigrouse/32 Thu, 08 May 2008 11:21:42 PDT Cyrtonyx montezumae (Vigors) 1830 Other vernacular names: Black quail, Codorniz Encinera, Codorniz Pinta, crazy quail, fool quail, massena quail, Mearns quail, Montezuma quail, painted quail, squat quail Range: Southwestern United States south to Oaxaca, Mexico. The doubtfully distinct species C. ocellatus (Gould) extends from southern Oaxaca to Nicaragua. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/31 http://digitalcommons.unl.edu/bioscigrouse/31 Thu, 08 May 2008 11:19:49 PDT Dactylortyx thoracicus ( Gambel) 1848 Other vernacular names: Chiviscoyo, Cinco Real, Codorniz, Chifladora, long-clawed quail, long-toed partridge, long-toed quail Range: Mountainous areas from Mexico to El Salvador and Hondu Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/30 http://digitalcommons.unl.edu/bioscigrouse/30 Thu, 08 May 2008 11:17:51 PDT Odontophorus guttatus (Gould) 1838 Other vernacular names: Bolonchaco, spotted partridge, thick-billed wood quail Range: Forested parts of the subtropical zone of southeastern Mexico south through Guatemala, British Honduras, Honduras, Nicaragua, and Costa Rica to extreme western Panama. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/29 http://digitalcommons.unl.edu/bioscigrouse/29 Thu, 08 May 2008 11:16:11 PDT Colinus nigrogularis (Gould) 1843 Other vernacular names: Black-throated quail, Codorniz Garganta Negra, Cuiche Yucateco, Yucatán bobwhite. Range: The Yucatán peninsula, the Lake Petén district of Guatemala, and coastal portions of British Honduras and Honduras, to extreme northeastern Nicaragua. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/28 http://digitalcommons.unl.edu/bioscigrouse/28 Thu, 08 May 2008 11:14:33 PDT Colinus virginianus (Linnaeus) 1758 Other vernacular names: American colin, Codorniz Común, Cuiche Común, partridge, quail Range: Virtually all of the eastern United States north to southern Maine, New York, southern Ontario, central Wisconsin, and central Minnesota, west to southeastern Wyoming, eastern Colorado, eastern New Mexico, and eastern Mexico south to Chiapas and adjacent Guatemala, but excluding the lowlands of Yucatán. Also existing as isolated populations in Sonora (largely extirpated) and as introduced populations in the Columbia and Snake river basins of Washington, Oregon, Idaho, and northwestern Wyoming (Bighorn and Shoshone River valleys). Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/27 http://digitalcommons.unl.edu/bioscigrouse/27 Thu, 08 May 2008 11:13:01 PDT Callipepla californica (Shaw) 1798 Other vernacular names: California partridge, Catalina quail, Codorniz Californiana, crested quail, San Lucas quail, San Quintin quail, topknot quail, valley quail Range: From southern Oregon and western Nevada south to the tip of Baja California. Introduced into southern British Columbia, Washington, Idaho, northern Oregon, and Utah Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/26 http://digitalcommons.unl.edu/bioscigrouse/26 Thu, 08 May 2008 11:11:35 PDT Callipepla gambelii (Gambel) 1843 Other vernacular names: Arizona quail, Codorniz de Gambel, desert quail, Olanthe quail. Range: From southern Nevada, southern Utah, and western Colorado south to northeastern Baja California, central Sonora, northwestern Chihuahua, and western Texas. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/25 http://digitalcommons.unl.edu/bioscigrouse/25 Thu, 08 May 2008 11:11:34 PDT Callipepla douglasii (Vigors) 1829 Other vernacular names: Benson quail, Codorniz Gris, crested quail, Douglas quail, Lesson quail, Yaqui quail. Range: Western Mexico from Sonora and Chihuahua to Nayarit and Jalisco Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/24 http://digitalcommons.unl.edu/bioscigrouse/24 Thu, 08 May 2008 11:10:09 PDT Callipepla squamata (Vigors) 1830 Other vernacular names: Blue racer quail, blue quail, Cordorniz Azul, Codorniz Escamosa, cottontop quail, Mexican quail, scaled partridge, top-knot quail, Zollin. Range: From southern Arizona, northern New Mexico, eastern Colorado, and southwestern Kansas south to central Mexico. Introduced into central Washington and eastern Nevada. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/23 http://digitalcommons.unl.edu/bioscigrouse/23 Thu, 08 May 2008 11:07:58 PDT Oreortyx pictus (Douglas) 1829 Other vernacular names: Cordoniz de Montana, mountain partridge, painted quail, plumed quail, San Pedro quail. Range: Resident in the western United States from southern Washington and southwestern Idaho east to Nevada and south to Baja California. Also introduced in western Washington and western British Columbia (Vancouver Island). Introduced but of uncertain status in western Colorado Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/22 http://digitalcommons.unl.edu/bioscigrouse/22 Thu, 08 May 2008 11:07:57 PDT Philortyx fasciatus ( Gould) 1844 Other vernacular names: Banded quail, Chorrunda, Codorniz listada. Range: Resident in semiarid tropical scrub of the Pacific slope from southwestern Jalisco to southeastern Guerrero, and inland to Morelos and Puebla, Mexico. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/21 http://digitalcommons.unl.edu/bioscigrouse/21 Thu, 08 May 2008 11:07:56 PDT Includes: Long-tailed Tree Quail, Dendrortyx macroura (Jardine & Selby) 1828 Bearded Tree Quail, Dendrortyx barbatus (Gould) 1846 Buffy-crowned Tree Quail, Dendrortyx leucophrys (Gould) 1844 Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/20 http://digitalcommons.unl.edu/bioscigrouse/20 Thu, 08 May 2008 11:06:31 PDT Tympanuchus phasianellus (Linnaeus) 1858 Other vernacular names: Brush grouse, pintail grouse, prairie grouse, prairie pheasant, sharptail, speckle-belly, spike-tail, spring-tail, whitebelly, white-breasted grouse. Range: Currently from north central Alaska, Yukon, northern Mackenzie, northern Manitoba, northern Ontario, and central Quebec south to eastern Washington, extreme eastern Oregon, Idaho, northeastern Utah, Wyoming, and Colorado, and in the Great Plains from eastern Colorado and eastern Wyoming across Nebraska, the Dakotas, northern Minnesota, northern Wisconsin, and northern Michigan. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/19 http://digitalcommons.unl.edu/bioscigrouse/19 Thu, 08 May 2008 10:18:50 PDT Tympanuchus cupido ( Linnaeus) 1758 Other vernacular names: prairie chicken, prairie cock, prairie grouse, prairie hen SUBSPECIES T. c. cupido (Linnaeus): Heath hen or eastern pinnated grouse. Extinct since 1932. Formerly along the East Coast from Massachusetts south to Maryland and north central Tennessee. T. c. pinnatus (Brewster): Greater prairie chicken. Currently limited to several small isolated populations in Michigan, Wisconsin, and Illinois and to the grasslands of extreme southern Manitoba, northwestern Minnesota, North Dakota, South Dakota, Nebraska, Kansas, and western Missouri. T. c. attwateri Bendire: Attwater prairie chicken. Currently limited to a few isolated populations along the coast of Texas from Arkansas and Refugio counties to Galveston County, and inland to Colorado and Austin counties. T. c. pallidicinctus (Ridgway): Lesser prairie chicken. Currently limited to arid grasslands of southeastern Colorado and southwestern Kansas southward through Oklahoma to extreme eastern New Mexico and northwestern Texas. Recognized by the A.O.U. Check-list (1957) as a separate species. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/18 http://digitalcommons.unl.edu/bioscigrouse/18 Thu, 08 May 2008 10:17:32 PDT Bonasa umbellus ( Linnaeus) 1776 Other vernacular names: birch partridge, drummer, drumming grouse, long-tailed grouse, mountain pheasant, partridge, pine hen, pheasant, tippet, white-flesher, willow grouse, wood grouse, woods pheasant. Range: Resident in the forested areas from central Alaska, central Yukon, southern Mackenzie, central Saskatchewan, central Manitoba, northern Ontario, southern Quebec, southern Labrador, New Brunswick, and Nova Scotia south to northern California, northeastern Oregon, central Idaho, central Utah, western Wyoming, western South Dakota, northern North Dakota, Minnesota, central Arkansas, Tennessee, northern Georgia, western South Carolina, western North Carolina, northeastern Virginia, and western Maryland. Recently introduced in Nevada and Newfoundland Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/17 http://digitalcommons.unl.edu/bioscigrouse/17 Thu, 08 May 2008 10:16:06 PDT Lagopus leucurus (Richardson) 1831 Other vernacular names: Snow grouse, snow partridge Range: From central Alaska, northern Yukon, and southwestern Mackenzie south to the Kenai Peninsula, Vancouver Island, the Cascade Mountains of Washington, and along the Rocky Mountains from British Columbia and Alberta south to northern New Mexico Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/16 http://digitalcommons.unl.edu/bioscigrouse/16 Thu, 08 May 2008 10:14:48 PDT Lagopus mutus (Montin) 1776 Other vernacular names: Arctic grouse, barren-ground bird, Chamberlain ptarmigan, Dixon ptarmigan, Nelson ptarmigan, Reinhardt ptarmigan, rocker (in Newfoundland), snow grouse, Townsend ptarmigan, white grouse. Range: Circumpolar. In North America from northern Alaska, northwestern Mackenzie, Melville Island, northern Ellesmere Island, and northern Greenland south to the Aleutian Islands, Kodiak Island, southwestern and central British Columbia, southern Mackenzie, Keewatin, northern Quebec, southern Labrador, and Newfoundland (A. 0 . U.) Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/15 http://digitalcommons.unl.edu/bioscigrouse/15 Thu, 08 May 2008 10:13:06 PDT Lagopus lagopus (Linnaeus) 1758 Other vernacular names: Alaska ptarmigan, Alexander ptarmigan, Allen ptarmigan, Arctic grouse, red grouse (Scotland form), Scottish grouse, white grouse, white-shafted ptarmigan, willow grouse, willow partridge. Range: Circumpolar. In North America from northern Alaska, Banks Island, Melville Island, Victoria Island, Boothia Peninsula, Southampton Island, Baffin Island, and central Greenland south to the Alaska Peninsula, southeastern Alaska, central British Columbia, Alberta, Saskatchewan, Manitoba, central Ontario, central Quebec, and Newfoundland Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/14 http://digitalcommons.unl.edu/bioscigrouse/14 Thu, 08 May 2008 10:11:49 PDT Dendragapus canadensis (Linnaeus) 1758 [also Canachites canadensis] Other vernacular names: Black partridge, Canada grouse, cedar partridge, fool-hen, Franklin grouse, heath hen, mountain grouse, spotted grouse, spruce partridge, swamp partridge, Tyee grouse, wood grouse. Range: From central Alaska, Yukon, Mackenzie, northern Alberta, Saskatchewan, Manitoba, Ontario, Quebec, Labrador, and Cape Breton Island south to northeastern Oregon, central Idaho, western Montana, northwestern Wyoming, Manitoba, northern Minnesota, northern Wisconsin, Michigan, southern Ontario, northern New York, northern Vermont, northern New Hampshire, Maine, New Brunswick, and Nova Scotia Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/13 http://digitalcommons.unl.edu/bioscigrouse/13 Thu, 08 May 2008 10:11:48 PDT Dendragapus obscurus (Say) 1823 Other vernacular names: dusky grouse, fool hen, gray grouse, hooter, mountain grouse, pine grouse, pine hen, Richardson grouse, sooty grouse. Range: From southeastern Alaska, southern Yukon, southwestern Mackenzie, and western Alberta southward along the offshore islands to Vancouver and along the coast to northern California, and in the mountains to southern California, northern and eastern Arizona, and west central New Mexico Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/12 http://digitalcommons.unl.edu/bioscigrouse/12 Thu, 08 May 2008 10:11:46 PDT Centrocercus urophasianus (Bonaparte) 1827 Other vernacular names: Sage hen, spiny-tailed pheasant, sage cock, sage chicken Range: From central Washington, southern Idaho, Montana, southeastern Alberta, southern Saskatchewan, and western North Dakota south to eastern California, Nevada, Utah, western Colorado, and southeastern Wyoming Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/11 http://digitalcommons.unl.edu/bioscigrouse/11 Thu, 08 May 2008 10:09:46 PDT There can be little doubt that the grouse and quail provide the most important and most popular targets for more than ten million small-game hunters every year in North America (National survey, 1965). In much of the southeast, to go "bird" hunting simply means a day in pursuit of bobwhites, and likewise in New England "pa'tridge" hunting is regarded as the premier sport of all upland game hunting. These two species, the bobwhite and ruffed grouse, in 1970 were hunted in forty-seven states and eight provinces and are without question the most important of all North American upland game species (table 27). Although neither species was hunted during 1970 in Arizona or South Dakota, both have been legal game in South Dakota in recent years, and masked bobwhites originally occurred in southern Arizona, where they are now being restocked. In addition, the bobwhite occurs over much of Mexico and is an important game species in that country. In table 27 is presented a list of the grouse and quail occurring north of Mexico, as well as the states and provinces in which they could legally be hunted during the 1970-71 hunting season, based on information available to the author. Of course, the length of the season and the daily limits varied greatly in different areas and in a few instances the total season lasted only a day or two. However, the list does provide a method of estimating the relative importance of the species as game. On this basis alone, the ruffed grouse might be judged most important, while the bobwhite is almost as widely hunted. Other species that are currently hunted in ten or more states and provinces are the sage grouse, blue grouse, spruce grouse, sharp-tailed grouse, chukar partridge, and gray partridge. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/10 http://digitalcommons.unl.edu/bioscigrouse/10 Thu, 08 May 2008 10:08:14 PDT The rearing of grouse and quail for enjoyment, profit, or stocking in the wild has been an important aspect of grouse and quail biology. The very presence of chukar and gray partridges in North America, the occurrence of ruffed grouse in Newfoundland and Nevada, the presence of bobwhites, scaled quail, and California quail in Washington, and many other examples are ample testimony to the potential value of careful propagation and release programs. Between 1938 and 1968 a total of 110,663 bobwhites, 18,136 other native quails, 7,977 grouse, and 50,568 chukar partridges were released under Pittman-Robertson programs in the United States (based on a recent summary provided by the Bureau of Sports Fisheries and Wildlife). An additional but unspecified number of gray partridges was also part of the release program. Yeatter (1935) estimated that more than 260,000 of these birds had been released in North America by the 1930s. The problems of keeping and breeding grouse in captivity are distinctly different from and much greater than those of propagating quails and partridges, and as a result relatively few persons have succeeded in keeping and breeding grouse in large numbers or with consistent success. This is largely a reflection of the greater sensitivity of grouse to various poultry diseases and parasites that are transmitted by ground contact, forcing the game breeder to keep the birds on wire-bottom cages where they can have no direct contact with the ground or their own droppings. A summary of the diseases and parasites of grouse and quails has been provided by Bump et al. (1947) and Stoddard (1931), respectively, although the treatments recommended have been greatly modified in more recent years. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/9 http://digitalcommons.unl.edu/bioscigrouse/9 Thu, 08 May 2008 10:07:46 PDT One of the most complex and fascinating aspects of grouse and quail biology is their social behavior, particularly that related to reproduction. Natural selection in the quail group has seemingly favored the retention of a monogamous mating system with the associated advantages of maintaining the pair bond through the breeding season. This system allows the male to participate in the protection of the nest, possibly participate in incubation, and later care for the brood. It also provides the possibility, if not the frequent actuality, that the male might undertake the entire incubation or rearing of the first brood, while the female is freed to lay a second clutch and rear a second brood in a single breeding season. In addition, within the quails may be seen a breakdown of typical avian territorial behavior patterns, probably resulting from the greater survival value of ecological adaptations favoring sociality in these birds. Not only do these fairly vocal species benefit from their mutual alarm signals by remaining together but also their small size and catholic feeding behavior reduce the likelihood that the optimal breeding densities will exceed the carrying capacities of the habitat. By contrast, in the grouse there is a clear indication that selective pressures have favored the retention of strong territorial behavior, and there is a direct relationship between a male's capacities to establish and maintain a favorable territory and his ability to reproduce successfully. This territoriality perhaps results mainly from the wide variation among males in their aggressiveness and reproductive vigor but also from the possibility that in these species the carrying capacity of the habitat in relation to the population density may be more significant for the species' survival than are the advantages of sociality. Thus, territorial behavior among males is conspicuous in all the grouse species. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/8 http://digitalcommons.unl.edu/bioscigrouse/8 Thu, 08 May 2008 10:06:45 PDT Like other animals, grouse and quail exist as natural populations dependent upon particular habitats and vary in population density between the absolute minimum populations that have permitted past survival to fairly dense populations that may approach or even temporarily exceed the carrying capacity of the habitat. Each species may also have an upper limit on the density of the population, or a saturation point, which is independent of the carrying capacity of the habitat but is determined by social adaptations. Within the population as a whole, individual birds or coveys may have home ranges, geographical areas to which their movements are limited and within which they spend their entire lives. Part of the home range may be defended by individuals so that conspecifics of the same sex are excluded for part or all of the year; such areas of localized social dominance and conspecific exclusion are called territories. Among species lacking discrete territories and in which the social unit is the covey or flock rather than the pair or family, dominance hierarchies, or peck orders, may serve to integrate activities in the flock. These behavioral adaptations and habitat relationships play important roles in population ecology, and will be considered in detail in the individual species accounts. However, a preliminary survey may help to provide generalizations that will be useful to keep in mind when considering individual species. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/7 http://digitalcommons.unl.edu/bioscigrouse/7 Thu, 08 May 2008 10:05:40 PDT The reproductive potentiaI of animal species is a compound result of numerous behavioral and physiological characteristics, most of which can be considered species-typical. These include such things as the time required to attain reproductive maturity, the number of nesting or renesting attempts per year once maturity is attained, the number of eggs laid per breeding attempt, and the number of years adults may remain reproductively active. These traits place an upper limit on the reproductive potential of a species, which is never actually attained. Rather, the actual rate of increase will only approach the reproductive potential, being limited by such things as the incidence of nonbreeding; the mortality rates of adults; decreased hatching success resulting from infertility, predation, or nest abandonment; relative rearing success; incidence of renesting and clutch sizes of renests; and similar factors that affect the reproductive efficiency. The relative involvement of the male in protecting the nest or the young may also influence hatching or rearing success. Among those species in which the male does not participate in nesting behavior, the relative degree of monogamy, polygamy, or promiscuity may strongly influence the reproductive ecology and population genetics of the species. Although many of these considerations will be treated under the accounts of the individual species, a general comparison of the grouse and quail groups as a whole are worth considering here, to see if any general trends can be detected. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/6 http://digitalcommons.unl.edu/bioscigrouse/6 Thu, 08 May 2008 10:04:54 PDT The study of hybridization between species, under either natural or artificial conditions, provides information of value for a variety of reasons. In general, it may be expected that the incidence of crossbreeding between populations existing under natural conditions will be related to their nearness of relationship, and information of taxonomic interest may be obtained from such study. Furthermore, the relative survival and fertility of the resulting hybrids should provide an indication of the degree of genetic difference between the parental types, and thus genetic information may be available through experimental hybridization studies. Hybrids provide favorable material for studying the chromosomal numbers and configurations among related species, and when they are fertile the degree of phenotypic variation in second or backcross generations may be used to estimate genetic differences controlling specific traits. Finally, the presence or absence of natural hybridization between closely related forms occurring in the same habitats may provide a clue as to the degree of niche overlap and interspecies competition for habitat resources. Therefore, if the basis for periodic or local hybridization between two forms that normally do not hybridize can be established, the ecological differences that normally prevent hybridization may possibly be deduced. For various reasons, the grouse and quails of North America exhibit a rather surprisingly high tendency to hybridize, even among species belonging to seemingly different genera. Peterle (1951) reviewed the cases of intergeneric hybrids reported in gallinaceous birds, and Cockrum (1952) provided a more complete survey of hybridization in North American birds. Sibley (1957) commented on the taxonomic significance of hybridization in grouse, and a similar review of the significance of hybridization in the New World quails is available (Johnsgard, 1970). For a complete listing of all known hybrids of gallinaceous birds, including those reported from Europe and Asia, the summary by Gray (1958) may be consulted. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/5 http://digitalcommons.unl.edu/bioscigrouse/5 Thu, 08 May 2008 10:03:45 PDT An understanding of the molts and plumages of the quails and grouse is of great importance to the applied biologist, for they provide clues that are valuable for determining age and sex of individual birds without resorting to internal examination. They thus offer a means of analyzing wild populations as to sex and age composition, which are basic indices to past and potential reproductive performances and probable mortality rates. Additionally, molts and plumages are generally speciesspecific traits, which have resulted from pressures of natural selection over a long period of time in a particular habitat and climate. The ecology of the species is of major importance in this regard; species occurring in more northerly regions may undergo their molts more rapidly than those in southerly ones or, as in the case of the willow ptarmigan, certain races may even lack particular plumages that occur in populations existing in other areas having different climates. From the time they hatch, all grouse and quails exhibit a series of specific plumages, separated by equally definite molts, that are comparable in nearly all species. The only known exception to this occurs in the genus Lagopus, which is unique in having an extra molt, and thus a supplementary plumage, intercalated between its summer and winter plumages. This special case will be dealt with as required; the following summary will thus serve to provide the basic sequences and terminology that describe the molts and plumages found in the North American grouse and quails. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/4 http://digitalcommons.unl.edu/bioscigrouse/4 Thu, 08 May 2008 10:01:36 PDT All of the grouse, quails, and introduced partridges of North America share a number of anatomical traits which provide the basis for their common classification within the order Galliformes. Among these are the facts that they all have fowl-like beaks and four toes. In all the North American species the hind toe is elevated and quite short, thus is ill-adapted for perching. There are always ten primaries, thirteen to twenty-one secondaries, and twelve to twenty-two tail feathers (rectrices). Aftershafts on the contour feathers are well developed, especially in the grouse, and true down feathers are infrequent. A large crop is present, and is associated with the largely granivorous (seed-eating) behavior of most quails, and the more generally herbivorous (leaf-eating) diets of grouse. The egg colors range from pastel or earth tones (buff, cream, olive, etc.) to white, with darker spotting prevalent among those species having nonwhite eggs. The nest is built on the ground, and incubation is by the females alone or occasionally by both sexes (some quails and partridges). The young are down-covered and precocial and are usually able to fly short distances in less than two weeks. They are cared for by the female (most grouse) or by both parents (some ptarmigans, all quails). A number of external structural characteristics typical of grouse, quails and partridges are shown in figures 3 and 4. Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/3 http://digitalcommons.unl.edu/bioscigrouse/3 Thu, 08 May 2008 09:59:29 PDT The modern array of grouse-, quail-, and partridge-like species occurring in North America is the result of three processes: evolution and speciation within this continent, range expansion or immigration from Central America and Eurasia, and recent introductions by man. The last category accounts for the presence in North America of the chukar and gray partridges, which are both natives of Europe or southern Asia and typical representatives of the quail-like and partridge-like forms that have extensively colonized those land masses. It is still necessary to account for the presence of the nine or so species of grouse-like forms that are native to this continent, as well as the fourteen or fifteen species of New World quails that occur north of the Guatemala-Mexico border. In general, the evidence clearly indicates that the New World quails had their center of evolutionary history and speciation in tropical America, whereas the grouse are a strictly Northern Hemisphere group that perhaps originated in North America but which now occur throughout both this continent and Eurasia and at present represent about an equal number of species in each of the two hemispheres. North America therefore has provided the common ecological conditions to which two distinctly different groups of gallinaceous birds have become independently adapted and have undergone somewhat convergent evolutionary trends. The evolutionary history of grouse- and quail-like birds on this continent is a long one, going back to at least Oligocene times, from which an indeterminate quail-like fossil is known, an addition to a unique fossil quail genus (Nanortyx) (Tordoff, 1951). Perhaps Paleophasianus from the Eocene represents the earliest grouse-like fossil (Holman, 1961), although it is more probably a species of limpkin (Cracraft, 1968). Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/2 http://digitalcommons.unl.edu/bioscigrouse/2 Thu, 08 May 2008 09:57:25 PDT Table of Contents: List of Illustrations Preface Introduction Paul A. Johnsgard http://digitalcommons.unl.edu/bioscigrouse/1 http://digitalcommons.unl.edu/bioscigrouse/1 Wed, 07 May 2008 11:49:11 PDT Nearly all of the gallinaceous birds that are native to North America are included in two taxonomic groups, the grouse-like species of the subfamily Tetraoninae, and the quail-like species of the subfamily Odontophorinae. The former represent a temperate and subarctic group of about sixteen species which collectively have a widespread distribution in the Northern Hemisphere, and over half of which are found in North America. The latter group is a strictly Western Hemisphere assemblage that collectively includes about thirty species, almost half of which occur north of the Mexico-Guatemala border. Most of the remaining quails are tropical forest birds of northern and western South America about which very little is known. Thus, evidence suggests that North America was originally doubly colonized by early gallinaceous stock; from the south by basically tropical-forest-adapted birds that have evolved into the present array of quail species, and from the north by relatively arctic-adapted forms that have given rise to the present species of ptarmigans and grouse. Convergent evolution of these two separate but related stocks has since allowed much of North America to become inhabited by birds having similar ecological adaptations and in some cases overlapping distributions. Within each of the two ancestral groups, evolutionary radiation has developed an interesting spectrum of anatomical variations, ecological adaptations, and behavioral specializations. These latter two aspects--adaptational niche variations associated with habitat differences, and behavioral variations associated with maximal reproductive efficiencies under varied climates, habitats and contacts with associated species--are the primary subjects of this book. Anatomical and physiological considerations will be given some attention in the early chapters, but the primary focus will be on the living bird in its natural environment. This complete work is 654 pages, including 140 color and black-and-white plates. The PDF file of the whole book is 48 megabytes. All chapters and species descriptions are also posted here separately, for faster downloading of specific portions. Paul A. Johnsgard