The Project Gutenberg eBook of Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and Colorado

This ebook is for the use of anyone anywhere in the United States and most other parts of the world at no cost and with almost no restrictions whatsoever. You may copy it, give it away or re-use it under the terms of the Project Gutenberg License included with this ebook or online at www.gutenberg.org. If you are not located in the United States, you will have to check the laws of the country where you are located before using this eBook.

Title: Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and Colorado

Author: Sydney Anderson

Release date: March 22, 2010 [eBook #31730]
Most recently updated: January 25, 2021

Language: English

Credits: Produced by Chris Curnow, Simon Gardner, Joseph Cooper and
the Online Distributed Proofreading Team at
http://www.pgdp.net

*** START OF THE PROJECT GUTENBERG EBOOK SUBSPECIATION IN THE MEADOW MOUSE, MICROTUS MONTANUS, IN WYOMING AND COLORADO ***

University of Kansas Publications

Museum of Natural History

Volume 7, No. 7, pp. 489-506, 2 figures in text

July 23, 1954

Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and Colorado

BY

SYDNEY ANDERSON

University of Kansas
Lawrence
1954

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Robert W. Wilson

Volume 7, No. 7, pp. 489-506, 2 figures in text
Published July 23, 1954

University of Kansas
Lawrence, Kansas

PRINTED BY
FERD VOILAND, JR., STATE PRINTER
TOPEKA, KANSAS
1954

25-3560

Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and Colorado

BY

SYDNEY ANDERSON

Microtus montanus reaches the eastern limits of its geographic distribution in Wyoming and Colorado. There the mountains, but in general not the lowlands, are occupied by this species. A certain minimum of moisture may be of direct importance to the mouse and certainly is indirectly important, because certain hydrophytic or mesophytic grasses used by the mouse for food, for protection from enemies, and for shelter from the elements are dependent on the moisture. Areas suitable for Microtus montanus are separated by deserts that are dominated by sagebrush and other xerophytic plants or by forests or rocky exposures at higher altitudes. A relatively small percentage, probably less than ten per cent, of the total area even in the more favorable parts of the range of the species is suitable for occupancy. In these mice, as in other microtines (Elton, 1942; Piper, 1909), there are seasonal, and irregularly multiannual fluctuations in population density, which sometimes are extreme. Consequently the mice at some times seem to be absent from suitable habitats, and at some other times occur there in amazingly large numbers.

Because the species is broken up into partly isolated, or at times completely isolated, colonies or local populations it may be supposed that various evolutionary forces such as selection and random genetic drift operate to foster variation. The degree to which racial distinction is attained may depend upon these forces and the time available. In Microtus montanus in the eastern Rocky Mountains the degree of subspecific distinction is not great.

The study here reported upon is based on 1,187 specimens of Microtus montanus from Wyoming, Colorado, Idaho and Montana, and on work in the field. I spent approximately four months in the field in this area, in the summers of 1950, 1951, and 1952. The specimens studied were arranged according to localities and the larger series were compared statistically. Each of two series, totaling 136 specimens, was studied intensively to ascertain the kind and range of variation within single populations. Twenty-seven measurements, various proportions based on these measurements, and differences in color were analyzed. Fifteen characters, judged to be most significant, were selected for use in comparing all series. In addition, certain characters that can not be expressed easily by measurements, such as inflation of the auditory bullae and the curvature of the zygomatic arch, were observed. The studies by A. B. Howell (1924) of variation in Microtus montanus yosemite Grinnell in California and those by O. B. Goin (1943) of Microtus pennsylvanicus pennsylvanicus (Ord) were useful. The external measurements are from the collectors' field labels. The measurements of the skull all were taken with dial calipers reading to a tenth of a millimeter. The anteroposterior measurements of the skull all were taken along the shortest line between the points specified below and are not necessarily along a line parallel to the long axis of the skull. These measurements were taken on the left side of the skull whenever possible. Some of the skulls are damaged and therefore some measurements could not be taken and are omitted in the computations. Measurements are in millimeters.

The results of these studies were submitted to the Department of Zoology and the Graduate School of the University of Kansas in partial fulfillment of the requirements for the degree of Master of Arts (1952) and are available in manuscript form at the Museum of Natural History and the library of the University of Kansas.

Explanation of Measurements

Caudal index.—the length of the tail expressed as a percentage of the length of the head and body. The length of the head and body is the collector's measurement of total length less the length of the tail.

Cranial Measurements of Length.

Condylobasilar length.—from the exoccipital condyle to the most posterior point on the border of the alveolus of the upper incisor.

Alveolobasilar length.—from the posterior border of the alveolus of the third upper molar to the posterior border of the alveolus of the incisor.

Palatilar length.—from the anteriormost part of the posterior border of the bony shelf of the palate to the posteriormost part of the alveolus of the incisor.

Alveolar length of upper molar tooth-row.—from the most posterior point of the alveolus of the third upper molar to the most anterior point of the alveolus of the first upper molar.

Measurements of Breadth.

Zygomatic breadth.—greatest transverse width.

Interorbital breadth.—the breadth of the interorbital constriction.

Lambdoidal breadth.—between the lateralmost points of the lambdoidal crest.

Prelambdoidal breadth.—between the medialmost margins of the prominent fenestrae in the posterodorsal parts of the squamosal bones. To these fenestrae Howell (1924:995) applied the adjective "prelambdoidal," but other authors have used other names (see Hill, 1935:127).

Depth of braincase.—shortest distance from the ventral side of the cranium at the suture between the basioccipital and basisphenoid bones to the dorsal surface of the cranium (usually not perpendicular to the long axis of the skull).

The history of our knowledge of Microtus montanus in this area begins with the early work of the United States Bureau of Biological Survey directed by C. H. Merriam (1891), and participated in by Vernon Bailey (1900, 1917), Merritt Cary (1911, 1917), and others. The changes in nomenclature which grew out of increased understanding of these mice through additional collecting are expressed in the synonymies under the accounts of subspecies. As a result of my studies two of the three subspecific names previously proposed for mice from this area have been retained although changes are proposed in the ranges assigned to the two subspecies and two additional heretofore unrecognized subspecies are named and described. Furthermore the additional specimens and my studies of variation make modifications in the characterization of these subspecies necessary. Not all of the samples here assigned to a single subspecies are identical and I therefore list and discuss some of the local variants.

Numerous members of summer field parties from the Museum of Natural History at the University of Kansas collected most of the specimens studied and wrote field notes that have been helpful. I am grateful to these persons and to Professor E. R. Hall and Dr. R. H. Baker for their assistance and helpful suggestions. Specimens in the following museums were made available by their respective curators: Chicago Natural History Museum by Mr. Colin C. Sandborn, The Museum of Zoology at the University of Michigan by Dr. E. T. Hooper, The American Museum of Natural History by Mr. G. G. Goodwin, The United States National Museum by Dr. David H. Johnson and the Biological Surveys Collection by Miss Viola S. Schantz. A fellowship from the National Science Foundation made possible the studies at the museums other than at the University of Kansas.

Microtus montanus nanus (Merriam)

Arvicola (Mynomes) nanus Merriam, N. Amer. Fauna, 5:62, July 30, 1891.

Microtus montanus nanus, Hall, Proc. Biol. Soc. Washington, 51:131, August 23, 1938.

Microtus nanus, Bailey, N. Amer. Fauna, 17:30, June 6, 1900 (part).

Microtus montanus caryi Bailey, Proc. Biol. Soc. Washington, 30:29, February 21, 1917.

Microtus nanus nanus, Cary, N. Amer. Fauna, 42:43, October 3, 1917 (part).

Type.—Adult male No. 23853/31253, U. S. National Museum, Department of Agriculture collection, from Pahsimeroi Mountains, Custer County, Idaho; obtained by C. Hart Merriam and Vernon Bailey, September 16, 1890.

Range.—Idaho; southwestern Montana; most of the southwestern half of Wyoming; southward to central Colorado. See figure 1.

Comparisons.—Comparisons with subspecies named as new in this paper will be found in the accounts of those subspecies beyond. From Microtus montanus fusus Hall, the subspecies to the south, M. m. nanus from Idaho differs as follows: averages smaller; slightly darker and less reddish and less yellowish in color; slightly wider braincase (see measurement of prelambdoidal breadth); larger bullae.

Measurements.—Average (= arithmetical mean) measurements of 34 specimens, both male and female, from several localities in eastern Idaho are: total length, 151; length of tail, 39; hind foot, 19.2; condylobasilar length of the skull, 25.0; zygomatic breadth, 15.0; alveolar length of upper molar tooth-row, 6.4; prelambdoidal breadth, 8.9; and lambdoidal breadth, 11.7.

Average and extreme measurements of six adult males from near Pocatello, Bannock County, Idaho, and nine adult males from near Afton, Lincoln County, Wyoming, are, respectively, as follows: total length, 143(135-150), 163(143-179); length of tail, 35.1(33-38), 42.8(36-49); caudal index, 32.0(28.0-33.1), 35.7(30.6-41.9); hind foot, 18.9(18-20), 18.8(17-20); condylobasilar length of skull, 24.4(24.0-26.0), 25.6(24.5-26.2); alveolobasilar length, 14.1(13.7-14.5), 14.6(13.8-15.0); palatilar length, 13.2(12.9-13.6), 13.8(13.2-14.5); alveolar length of upper molar tooth-row, 6.3(6.1-6.5), 6.3(6.0-6.6); depth of braincase, 7.7(7.5-7.9), 8.0(7.7-8.3); lambdoidal breadth, 11.4(11.0-11.7), 12.0(11.3-12.7); prelambdoidal breadth, 9.1(8.6-9.4), 8.7(8.0-9.4); zygomatic breadth, 14.3(13.8-14.7), 15.3(14.4-16.3); interorbital breadth, 3.6(3.5-3.7), 3.5(3.3-3.7). The average length of the nasal bones in the series from Pocatello is 7.1 mm. The averages, which have not been included in Table 1, for three measurements of the series from Carbon County, Wyoming, are as follows (Encampment, males; Encampment, females; Savery, males; and Savery, females, respectively): alveolobasilar length, 14.4, 14.3, 14.5, 14.3; interorbital breadth, 3.5, 3.4, 3.5, 3.4; depth of braincase, 7.8, 7.6, 7.9, 7.6. Additional measurements are included in Table 1 for other series.

Discussion.—The name Microtus montanus caryi Bailey is here placed in synonymy under M. m. nanus (Merriam). Vernon Bailey (1917) in his description of caryi made four assumptions that have been found to be entirely or partly invalid. First, he assumed that this is an "extreme variant which gradually changes in characters across Nevada and Utah, and reaches its maximum variation in Wyoming." The differences pointed out in subsequent descriptions of subspecies found in the above area do not show a gradual change in any character, or in the number of characters, nor is caryi an extreme when compared with the other subspecies. Second, Microtus nanus was not, as Bailey assumed, a different species than Microtus montanus. Third, he assumed that the characteristics of adults of nanus were adequately ascertainable from the thirteen topotypes available to him. Subsequent sampling from Idaho shows that the series of specimens available to Bailey was made up mostly of young and subadult animals. Finally, caryi does not occupy as Bailey stated "the meadows along streams in the arid sagebrush country of the Bear River, Green River, and Wind River valleys" exclusively, or characteristically. When the localities from which the species actually is known are plotted, it seems that the arid basin serves as a barrier and that the species is more commonly and abundantly found in montane meadows in the Transition and Canadian life-zones.

Certain samples, here assigned to M. m. nanus, that vary from the average of the subspecies deserve comment. For example, mice from the area in Wyoming southwest of the Green River (in the Uinta Mountains) have relatively smaller feet, but are larger in both total length and size of skull. Specimens from near Afton, Lincoln County, Wyoming, are relatively large in both total length and size of skull. This series and specimens from Teton County, Wyoming, are intermediate between nanus from Idaho and the newly named subspecies from near Cody, Park County, Wyoming, described below, in terms of both darkness and the amount of reddish color. Mice from Laramie County are more richly reddish-brown. The specimens from near Savery, in Carbon County, Wyoming, are darker. The alveolobasilar length relative to the condylobasilar length is smaller in the series from along Deer Creek, 16 mi. S, 11 mi. W Waltman, Natrona County, Wyoming. The series from the southern tier of counties in Wyoming and some of the specimens from Colorado have relatively wider zygomatic arches. The specimens from southern Sweetwater County, Wyoming, are relatively paler, have a relatively longer tail and longer hindfeet, lesser condylobasilar length, and wider braincase. Most of these variations are of questionable significance; they may be chance variations owing to errors in sampling.

Much of the south-central part of the state is relatively low and relatively arid. This area includes the arid basin of the Green River and its major tributaries and the arid Red Desert along the continental divide in Sweetwater County. This area might have acted as a barrier to the mice; gene flow might have been prevented between the populations of the western part of the state and those farther east in the Medicine Bow Mountains and Laramie Mountains. Nevertheless geographic variations of subspecific worth have not taken place. The barrier has either not been of as long duration, or has not been so complete and effective, as the other barriers in the state, namely the Absaroka Range, the Big Horn Basin, the Shoshone Basin, and the valley of the North Platte River. These four barriers presumably have led to the differentiation of the two subspecies that are newly named beyond. Each of the two areas which is set apart by these barriers and in which one of the newly named subspecies has evolved is small; therefore there is a lesser amount of suitable habitat available for each of the newly named mice than there is for M. m. nanus. It is conceivable, therefore, that in periods of adverse conditions in each of the small areas the size of the effective breeding population may have been so small that random genetic drift could have operated effectively, or that selection was more critical than in a larger, more stable population. It is difficult to test these possibilities because the selective value of the taxonomic characters is unknown. The observed pattern of variation and facts of distribution are, however, not contradictory to the above possibilities.

Specimens examined.—Total, 993, distributed as follows: All specimens unless otherwise indicated are in the University of Kansas Museum of Natural History. Specimens in other museums are labeled as follows: Chicago Natural History Museum (Chi); University of Michigan, Museum of Zoology (Mich); American Museum of Natural History (AMNH); United States National Museum (USNM); Biological Surveys Collection (USBS). Localities that are not represented in Fig. 1 because overlapping or crowding of the symbols would result are Italicized. Localities are arranged from north to south by states, within a state from northwest to southeast by counties, and within a county from north to south.

Wyoming: Yellowstone Park: Canyon Camp, 1 (USBS); Lower Geyser Basin, 1 (USBS); Upper Yellowstone River, 2 (AMNH); North end of Lake, Yellowstone National Park, 2 (AMNH). Teton Co.: Pacific Creek, 1 (USBS); Big Game Ridge, 3 (USBS 1, Mich 2); Whetstone Creek, 7 (Mich); Moran and environs (6 localities within a 5 mile radius), 28 (USBS 2, Mich 5); S fork Buffalo River, 7 (AMNH); 2 mi. W pass, Black Rock Creek, 1 (USBS); Jenny Lake, 5 (Mich); Bar BC Ranch, 2½ mi. NE Moose, 6500 ft., 2; Teton Pass above Fish Creek, 1 (USBS); Jackson and environs, 142 (Mich 141); Sheep Creek, 2 (Mich). Lincoln Co.: 13 mi. N, 2 mi. W Afton, 2; 10 mi. N, 2 mi. W Afton, 4; 9½ mi. N, 2 mi. W Afton, 3; 9 mi. N, 2 mi. W Afton, 9; 7 mi. N, 1 mi. W Afton, 12; Afton, 1 (USBS); Labarge Creek, 1 (USBS); Border, 6 (USBS); Cokeville, 2 (USBS); 6 mi. N, 2 mi. E Sage, 1; Cumberland, 5 (USBS). Sublette Co.: 34 mi. N, 4 mi. W Pinedale, 1; 33 mi. N, 2 mi. W Pinedale, 6; 32 mi. N, 1 mi. W Pinedale, 1; 31 mi. N Pinedale, 4; Dell Creek, on Ferris Ranch, 7 (Mich); Horse Creek, 7800 ft., Merna, 4 (USBS); Big Piney, 1 (USBS). Fremont Co.: 17½ mi. W, 2½ mi. N Lander, 9500 ft., 3; 17 mi. W, 2 mi. N Lander, 9300 ft., 4; Milford and environs (5 localities within a 1 mile radius), 23 (USBS 4); 15½ mi. S, 7½ mi. W Lander, 9200 ft., 1; South Pass City, 8000 ft, 8 (USBS); 23½ mi. S, 5 mi. W Lander, 8600 ft., 7. Natrona Co.: Deer Creek, 16 mi. S, 11 mi. W Waltman, 6950 ft., 44; 6 mi. S, 2 mi. W Casper, 5900 ft., 4; 6-4/5 mi. S, 2 mi. W Casper, 6100 ft., 1; 7 mi. S, 2 mi. W Casper, 6370 ft., 3; 10 mi. S Casper, 7750 ft., 33; Sun, 2 (USBS); 5 mi. W Independence Rock, 6000 ft., 4; 5 mi. W, 1 mi. S Independence Rock, 2. Converse Co.: Beaver, 1 (USBS). Uinta Co.: 1½ mi. W, ½ mi. S Cumberland, 6; 16 mi. S, 2 mi. W Kemmerer, 6700 ft., 3; 10 mi. SW Granger, 3 (Mich); Fort Bridger, 6650 ft., 25 (USNM 6); 9 mi. S Robertson, 8000 ft., 9; 9½ mi. S, ½ mi. W Robertson, 8600 ft., 1; 10 mi. S, 1 mi. W Robertson, 8700 ft., 25; 14 mi. S, 2 mi. E Robertson, 9000 ft., 5; 4 mi. S Lonetree, 1 (USBS). Sweetwater Co.: Farson, 3; Bitter Creek, 3 (AMNH); Kinney Ranch, 21 mi. S Bitter Creek, 6800 ft., 9 (USNM 1, AMNH 2); 32 mi. S, 22 mi. E Rock Springs, 7025 ft., on Vermillion Creek, 15. Carbon Co.: 18 mi. NNE Sinclair, 6500 ft., 10; Bridgers Pass, 18 mi. SW Rawlins, 7500 ft., 7; Saratoga, 1 (USBS); 6 mi. S, 13 mi. E Saratoga, 8500 ft., 5; 6 mi. S, 14 mi. E Saratoga, 8800 ft., 1; Lake Marie, 10,440 ft., 2; 1 mi. S Lake Marie, 2; ½ mi. S, 2 mi. E Medicine Bow Peak, 10,800 ft., 1; Encampment (12 localities from 10 mi. N, 14 mi. E to 9 mi. N, 3 mi. E Encampment and from 6500 to 8400 ft.), 63; 1/4 mi. N Riverside, 7380 ft., 2; S base Bridger Peak, 8800 ft., Sierra Madre Mountains, 1; 2 mi. S Bridger Peak, 9300 ft., 2; Savery (10 localities from 8 mi. N, 21 mi. E to 4 mi. N, 8 mi. E Savery and from 7300 to 8800 ft.), 80. Albany Co.: 30 mi. N, 10 mi. E Laramie, 6760 ft., 6; 29¾ mi. N, 9½ mi. E Laramie, 6350 ft., 1; 26 mi. N, 4½ mi. E Laramie, 6960 ft., 8; 26¾ mi. N, 6½ mi. E Laramie, 6700 ft., 3; 3 mi. N, 13 mi. E Laramie, 7500 ft., 1; 7 mi. N, 2 mi. E Laramie, 1 (Chi); 5 mi. N Laramie, 7400 ft., 15; Laramie, 4 (AMNH); 1 mi. E Laramie, 7160 ft., 4; 7-7/10 mi. SSW Laramie, 7200 ft., 4; 6½ mi. S, 8¾ mi. E Laramie, 8200 ft., 1; Headquarters Park, 10,200 ft., Medicine Bow Mountains, 3 (USBS); Centennial, 8120 ft., 1; 2-1/4 mi. ESE Brown's Peak, 10,300 ft., 3; 3 mi. ESE Brown's Peak, 10,000 ft., 12; 2 mi. S Brown's Peak, 10,600 ft., 1; Pole Mountain, 15 mi. SE Laramie, 4 (USBS 3); 1 mi. SSE Pole Mountain, 8350 ft., 4; 2 mi. SW Pole Mountain, 8300 ft., 13; 3 mi. S Pole Mountain, 8100 ft., 1; Sherman, 2 (AMNH). Laramie Co.: 5 mi. N, 1 mi. W Horse Creek P. O., 7200 ft., 1; Meadow, 2 (USBS); 11 mi. N, 5½ mi. E Cheyenne, 5450 ft., 7; 7 mi. W Cheyenne, 6500 ft., 10; Cheyenne, 3 (USNM).

Colorado: Moffat Co.: Lay, 6160 ft., 1 (AMNH). Routt Co.: Wright's Ranch, Yampa, 7700 ft., 2; Gore Range, 8 mi. E Toponas, 8000 ft., 2 (USBS). Larimer Co.: 12½ mi. W, 1½ mi. S Rustic, 1; 11 mi. W, 1 mi. S Rustic, 1; Cache La Poudre River, 1 (Chi); Estes Park, 3 (USBS 1, AMNH 2); 19½ mi. W, 2½ mi. S Loveland, 7280 ft., 6; 16 mi. W Loveland, 6840 ft., 1; 6 mi. W, ½ mi. S Loveland, 5200 ft., 1. Rio Blanco Co.: Meeker, 1 (USBS); 9½ mi. SW Pagoda Peak, 7700 ft., 3; 5 mi. S Pagoda Peak, 9100 ft., 2. Eagle Co.: Eagle, 1 (USBS); Pando, 2 (USBS). Grand Co.: Mt. Whiteley, 2 (USBS); Arapahoe Pass, Rabbit Ear Mountains, 2 (USBS); Coulter (near Granby), 5 (USBS); Arrowhead (near Dale), 1 (USBS). Boulder Co.: ¾ mi. N, 2 mi. W Allenspark, 8400 ft., 4; 3 mi. S Ward, 9000 ft., 3; Nederland, 16 (Chi). Clear Creek Co.: Mt. McLellan, 2 (USBS); Berthoud Pass, 4. Park Co.: Trout Creek Ranch, 2 mi. N Garo, 1 (USBS).

Specimens examined of M. m. nanus from eastern Idaho and Montana are as follows: Idaho: Custer Co.: Challis, 7 (USBS); Mill Creek, Challis Nat. Forest, 1 (USBS); Pahsimeroi Mts., 12 (USBS); Lost River Mts., 1 (USBS). Fremont Co.: N fork Snake River, 10 mi. SW Island Park, 6200 ft., 2 (AMNH); Black Springs Creek, 4 mi. W Ashton, 5200 ft., 1 (AMNH); 5 mi. W St. Anthony, 5000 ft., 1 (AMNH). Camas Co.: Camas Prairie, Corral, 5100 ft., 2 (USBS). Blaine Co.: Alturas Lake, 3 (USBS); Sawtooth Lake, 2 (USBS); Craters of the Moon, Laidlow Park, 2 (Mich); Ticura, 10 mi. S Picabo, 1 (USBS); 19 mi. NE Carey (Lava Lake), 8 (Mich). Butte Co.: 26 mi. SW Arco, 12 (Mich). Bingham Co.: Shelley, 6 (USBS). Bonneville Co.: 10 mi. SE Irwin, 4 (USBS). Owyhee Co.: Three Creeks, 3 (USBS). Twin Falls Co.: Castleford Fenced Plot, 11 mi. W, 9 mi. S Twin Falls, 1. Minidoka Co.: Heyburn, 2 (USBS). Cassia Co.: 2 mi. S, 2 mi. W Burley, 5. Bannock Co.: Pocatello, 23 (USBS 4); Swan Lake, 1 (USBS). Bear Lake Co.: Montpelier Creek, 6700 ft., 3 (USBS). Montana: Gallatin Co.: W. Fork of W. Fork, Gallatin River, 1 (USBS). Park Co.: Lamar River, 7000 ft., 1 (USBS); Gardiner, 1 (USBS). Sweet Grass Co.: 14 mi. S Big Timber, 1 (USBS); McLeod, 1 (USBS); West Boulder Creek, 18 mi. SE Livingston, 2 (USBS).

Microtus montanus codiensis, new subspecies

Type.—Female, adult, skin and skull; No. 27578, Museum of Natural History, University of Kansas, from 3⅕ mi. E and ⅗ mi. S Cody, 5020 ft., Park Co., Wyoming; obtained on August 11, 1948, by James W. Bee, original number 18-8-11-48.

Range.—In northwestern Wyoming eastward from the Absaroka and Wind River ranges into the western part of the Big Horn Basin.

Diagnosis.—A relatively large Microtus montanus; tail actually and relatively long; hind foot actually but not relatively large; skull large; zygomatic expanse actually and relatively large; alveolobasilar length relatively large; upper molar tooth-row relatively long; color relatively light, not reddish.

Fig. 1. Geographic range of Microtus montanus in Wyoming, Colorado, and adjacent areas. Fig. 1. Geographic range of Microtus montanus in Wyoming, Colorado, and adjacent areas. The solid circles represent localities from which specimens have been examined; the hollow circles represent type localities. The ranges of subspecies in Utah are after Durrant, 1952.

Guide to subspecies

M. m. nanus

M. m. codiensis

M. m. zygomaticus

M. m. fusus

M. m. micropus

M. m. nexus

M. m. amosus

M. m. rivularis

Comparisons.—As compared with the specimens of M. m. nanus from Idaho, the size is larger (see diagnosis and measurements). Certain proportions which differ from those of nanus and which are not in close agreement with the observed differences with age in specimens of nanus of a size comparable to codiensis are relatively large alveolobasilar length, relatively long alveolar length of upper molar tooth-row, relatively wide-spreading zygomatic arches, and relatively long tail. The color in codiensis is lighter than in nanus. As compared to the new subspecies named below from the Big Horn Mountains to the east, codiensis is of similar size in head-body length, but has a relatively as well as actually longer tail; the hind foot averages longer; the upper molar tooth-row is relatively longer; the color is slightly paler and less grizzled; the bullae are larger and less flattened; the angle formed at the suture between the basioccipital and basisphenoid bones is less acute; and the region of the suture is less prominently elevated between the bullae when viewed from the ventral aspect. The pterygoid plates mesial and posterodorsal to the posterior end of the last upper molar are less fenestrated, and the incisive foramina are less constricted posteriorly.

Fig. 2. Map showing the major barriers to Microtus montanus in Wyoming and Colorado; the barriers are the low areas named on the map (the name "Black Hills" is on the map for another reason; these hills are not a barrier). The major mountainous areas higher than approximately 8000 feet in elevation in Wyoming, Colorado and Utah are stippled. These mountainous areas include the habitat that is most suitable for the montane meadow mouse. The Black Hills are unoccupied by this species but these hills seem to be ecologically suitable for the species.

Measurements.—The average and the extremes for some measurements of 34 males and females, 27 from the type locality and 7 from other localities in the range assigned to this subspecies, are as follows: total length, 165 (146-186); length of tail, 44.2 (35-55); hind foot, 19.6 (17-21); condylobasilar length of the skull, 25.5 (24.0-27.5); zygomatic breadth, 15.6 (14.7-16.6); alveolar length of upper molar tooth-row, 6.6 (6.2-7.0); prelambdoidal breadth, 8.8 (8.1-9.5); lambdoidal breadth, 12.0 (11.2-12.8). As an indication of variability and for comparison with other series the coefficient of variability and two times the standard error of the mean for each measurement in this series are included in Table 1. The averages for some measurements of 27 topotypes are as follows: total length, 162; length of tail, 45.5; hind foot, 19.9; condylobasilar length, 25.6; palatilar length, 14.0; molar series, 6.6; alveolobasilar length, 14.9; zygomatic breadth, 15.6; interorbital breadth, 3.5; lambdoidal breadth, 12.1; prelambdoidal breadth, 8.9; depth of braincase, 7.8.

Discussion.—Three species of Microtus were collected by James W. Bee at the type locality. Microtus montanus codiensis, Microtus longicaudus mordax, and Microtus pennsylvanicus modestus were taken in the same runways in the same meadow, at the same time. Microtus ochrogaster haydeni, although not taken at this locality, occurs in the Big Horn Basin. These four species differ in their geographic ranges, being largely allopatric, except M. montanus and M. longicaudus which are sympatric. Although the different species have ecological preferences and habits which differ, several species of Microtus may occur together in local areas such as the above. Certain of the characteristics of M. m. codiensis are intermediate between those of the species M. montanus on one hand and those of the other three species on the other hand. Could interspecific hybridization between "good species" of Microtus take place in nature and possibly alter the characteristics of a local population?

Specimens examined.—Total, 50, distributed as follows (abbreviations for collections are given in the account of M. m. nanus; localities that are not represented in Fig. 1 because overlapping or crowding of the symbols would result are Italicized):

Montana: Carbon Co.: Beartooth Mountains, 2 (USBS); Beartooth Lake, 1 (USBS).

Wyoming: Park Co.: Black Mountain, head of Pat O'Hara Creek, 3 (USBS); 13 mi. N, 1 mi. E Cody, 5200 ft., 1; SW slope Whirlwind Peak, 9000 ft., 1; 5 mi. N Cody, 6300 ft., 1 (USBS); 3⅕ mi. E, ⅗ mi. S Cody, 31; Ishawooa Creek, 6300 ft., 2 (USBS); Valley, 1 (USBS); Needle Mountain, 10,500 ft., 4 (USBS). Hot Springs Co.: 3 mi. N, 10 mi. W Thermopolis, 4950 ft., 3.

Microtus montanus zygomaticus, new subspecies

Type.—Male, adult, skin and skull, No. 32761, Museum of Natural History, University of Kansas, from Medicine Wheel Ranch, 9000 ft., 28 mi. E Lovell, Big Horn County, Wyoming; obtained by R. Freiburg, original number 105.

Range.—The Big Horn Mountains of north-central Wyoming.

Diagnosis.—A large Microtus montanus with a relatively short tail; short molar series; broad zygomatic arches well rounded in lateral outline when viewed from above; small and flattened bullae; raised basioccipito-basisphenoid suture.

Comparisons.—For comparison with M. m. codiensis from the west, on the other side of the Big Horn Basin, see the account of that subspecies. In comparison with nanus this subspecies is slightly paler, in this respect showing more resemblance to codiensis although not so pale, and more grizzled or unevenly colored. This difference in color between zygomaticus and codiensis may not be of taxonomic significance. From both the topotypes of nanus, and the series of it from Wyoming, zygomaticus differs on the average in having a relatively shorter tail, a relatively shorter upper molar tooth-row, relatively more rounded and relatively more wide-spread zygomatic arches, and smaller more flattened bullae.

Measurements.—Average and extreme measurements of 24 adult males and females from several localities here referred to M. m. zygomaticus are as follows: total length, 159(150-175); length of tail, 37.6(31-46); hind foot, 18.6(17-20); condylobasilar length of the skull, 25.3(24.2-26.7); zygomatic breadth, 15.3(14.1-16.7); alveolar length of upper molar tooth-row, 6.2 (5.7-6.8); prelambdoidal breadth, 8.7(8.3-9.4); lambdoidal breadth, 11.9(11.0-12.5). Average and extreme measurements of a series of 12 adult male topotypes are as follows: total length, 159(144-174); length of tail, 36.4 (30-41); hind foot, 18.2(16-20); condylobasilar length of skull, 25.8(24.7-26.7); alveolobasilar length, 14.8(13.8-15.3); palatilar length, 13.8 (12.7-14.2); alveolar length of upper molar tooth-row, 6.4(5.9-6.6); zygomatic breadth, 15.9 (15.0-16.7); interorbital breadth, 3.6(3.4-3.7); lambdoidal breadth, 12.1 (11.5-12.5); prelambdoidal breadth, 8.6(8.3-8.9); depth of braincase, 8.0 (7.6-8.3).

Discussion.—This subspecies is separated from M. m. codiensis to the west by the Big Horn Basin. A series from along Buffalo Creek, 27 mi. N, 1 mi. E Powder River, 6075 ft., in Natrona County, Wyoming, is intermediate between the topotypes of zygomaticus and nanus in the characters cited above as distinguishing the two, but shows greater resemblance to zygomaticus in the shape of the zygomatic arch, in color which is paler than in topotypes of zygomaticus, and in the short hind foot. On these and on geographic grounds this population is referred to zygomaticus. Unfortunately we cannot be certain in many cases that an intermediate condition in a certain character indicates a genetically intermediate population and therefore true intergradation between the two subspecies to which the population is geographically intermediate. The topotypes of this subspecies are the most distinct of all the series which I have studied from the eastern Rocky Mountains, in terms of the degree of morphological departure from the norm for the species. After zygomaticus the following populations are arranged according to their degree of deviation from this norm (codiensis deviates most): topotypes of codiensis, fusus and a population from southern Sweetwater County, Wyoming, and lastly the nanus-caryi complex. Within the latter group, as I have mentioned, there are a number of local variants most of which do not differ significantly and do not conform to any geographic pattern.

Specimens examined.—Total, 55, distributed as follows (abbreviations for collections are given in the account of M. m. nanus; localities that are not represented in Fig. 1 because overlapping or crowding of the symbols would result are Italicized): Wyoming: Big Horn Co.: Medicine Wheel Ranch, 9000 ft., 28 mi. E Lovell, 30; W slope, head of Trappers Creek, 9500 ft., 2 (USBS). Washakie Co.: 9 mi. E, 5 mi. N Tensleep, 7400 ft., 1. Johnson Co.: 7½ mi. W, 1 mi. S Buffalo, 6500 ft., 3; Big Horn Mountains, 3 (USBS). Natrona Co.: Buffalo Creek, 27 mi. N, 1 mi. E Powder River, 6075 ft., 16.

Microtus montanus fusus Hall

Microtus nanus, Bailey, N. Amer. Fauna, 17:30, June 6, 1900 (part); Cary, N. Amer. Fauna 33:123, August 17, 1911.

Microtus montanus fusus Hall, Proc. Biol. Soc. Washington, 51:131-134, August 23, 1938; Warren, The Mammals of Colorado, Univ. of Okla. Press, p. 229, 1942.

Type.—Male, adult, skin and skull; No. 61281, Museum of Vertebrate Zoology; 2½ miles east of summit of Cochetopa Pass, Saguache County, Colorado; Sept. 21, 1933; collected by Annie M. Alexander; original number 2568. Type not seen by me.

Range.—Southern Colorado and northern New Mexico.

Comparisons.—For comparison with M. m. nanus, the subspecies to the northward, see the preceding account of that subspecies. For comparison with M. m. amosus the subspecies to the west see Hall (1938) and Durrant (1952). I have not examined specimens of amosus.

Measurements.—Average and extreme measurements for 17 adults including both males and females from several localities in southern Colorado are as follows: total length, 160 (136-179); length of tail, 42 (35-55); hind foot, 19.2 (17-23); condylobasilar length of the skull, 25.2 (24.0-26.0); zygomatic breadth, 15.0 (14.1-15.5); alveolar length of upper molar tooth-row, 6.4 (6.0-6.7); prelambdoidal breadth, 8.7 (8.3-9.2); lambdoidal breadth, 11.7 (11.1-12.6).

Average and extreme measurements of 4 adults (2 males and 2 females) from the type locality and 11 adults (4 males and 7 females) from other localities in southern Colorado are as follows: total length, 162 (157-168), 157 (137-169); length of tail (means only), 44.5, 40.5; hind foot, 18.8 (18-19), 18.6 (18-23); condylobasilar length of skull, 24.5 (24.0-24.7), 25.2 (24.3-26.1); alveolobasilar length, 14.2 (13.9-14.5), 14.6 (14.1-15.1); palatilar length, 13.2 (13.0-13.4), 13.5 (13.1-14.2); alveolar length of upper molar tooth-row, 6.3 (6.0-6.6), 6.4 (6.3-6.7); zygomatic breadth, 15.0 (14.3-15.5), 14.9 (14.1-15.5); interorbital breadth, 3.5 (3.3-3.6), 3.5 (3.3-3.7); lambdoidal breadth, 11.8 (11.1-12.6), 11.7 (11.2-12.3); prelambdoidal breadth, 8.6 (8.3-9.2), 8.8 (8.3-9.0); depth of braincase, 7.5 (7.2-7.8), 7.6 (7.1-7.9).

Discussion.—There is no sharp boundary between M. m. fusus of southern Colorado and the subspecies to the north, M. m. nanus. Although the line separating these two subspecies is drawn somewhat arbitrarily, on the whole the samples from north of this line more closely resemble nanus. All of the means for total length given above are larger than the maximum given in Hall's description of fusus. The caudal index (38 and 35% in two series) is slightly larger than that cited by Hall (33.3%) and is not significantly different from that in nanus (35.2%). The color in both young and old mice is variable, but in general is more yellowish, and less grayish, than in any other series studied.

There is a large area in western Colorado and eastern Utah, between the known ranges of M. m. fusus and M. m. amosus from which there are no specimens. Probably the species occurs only at certain places in this arid region which seems to be a partial barrier to the species.

Specimens of M. montanus from northern New Mexico have been referred previously to M. m. arizonensis. When he named M. m. fusus, Hall mentioned its resemblance to arizonensis in reddish coloration, but pointed out that fusus is less reddish. Of six specimens from Valle Santa Rosa, Jemez Mountains (USBS), 8500 ft., Rio Arriba County, New Mexico, three are immature, and the skulls of the remaining specimens are damaged. In reddish color and relatively large size these few specimens resemble arizonensis more than fusus although the locality of occurrence is closer to the geographic range of the northern fusus than to that of arizonensis. The identification of these specimens as arizonensis is provisional; additional specimens are needed from the area, 200 miles wide, which separates the ranges as now known of arizonensis in Arizona from the occurrence in New Mexico. There is a single specimen from this area, the damaged skull of which prevents conclusive identification. The specimen is either M. montanus or M. mexicanus, and is from Nutria, on the southern edge of the Zuni Mountains (USBS). Detailed comparison of fusus and arizonensis is not attempted here although it may be stated that in several characters fusus is intermediate between arizonensis to the south and nanus to the north.

Specimens examined.—Total, 89, distributed as follows (abbreviations for collections are given in the account of M. m. nanus; localities that are not represented in Fig. 1 because overlapping or crowding of the symbols would result are Italicized):

Colorado: Pitkin Co.: 5 mi. W Independence Pass, 11,000 ft., 1 (Chi). Lake Co.: Independence Pass, 12,095 ft., 2 (Chi). Gunnison Co.: Gothic, 2 (USBS); Decker's Ranch, Crested Butte, 2 (AMNH); Almont, 3 (USBS). Montrose Co.: Coventry, 5 (USBS 4, AMNH 1). Saguache Co.: Cochetopa Pass and environs, 44 (USBS 22). Hinsdale Co.: Ruby Lake, 1 (USBS). Mineral Co.: 3 mi. E Creede, 1; 23 mi. S, 11 mi. E Creede, 9300 ft., 7. La Plata Co.: Florida, 6800 ft., 1. Conejos Co.: 1 mi. S, 19 mi. W Antonito, 10,200 ft., 3; 4 mi. S, 23 mi. W Antonito, 1; 5 mi. S, 24 mi. W Antonito, 9600 ft., 9.

New Mexico: Rio Arriba Co.: 6 mi. W Hopewell, 9900 ft., 6 (USBS); Tusas River, 8700 ft., 1 (USBS).

Some measurements not given above are included in Table 1, together with the number of specimens and the sex if restricted to one sex. So that the variability can be evaluated more adequately, the coefficient of variability and 2 times the standard error of the mean are included for the measurements in two series. The series consist of all the adult specimens (with a condylobasilar length of 24.0 mm. or more) of both sexes from the areas specified. Various barriers are shown in Fig. 2 for comparison with the distributions of the subspecies and the localities of known occurrence shown in Fig. 1. Microtus montanus has not been taken in the Black Hills area of extreme northeastern Wyoming. Suitable montane habitat is present and both Microtus pennsylvanicus insperatus and Microtus longicaudus longicaudus occur there. The arid basin of the Powder River presumably is a barrier that has prevented M. montanus from reaching this area.

Table 1. Average Measurements, in Millimeters, of Adults of Microtus montanus.

Key to column headings:

A: No. of individuals averaged
B: Total length
C: Length of tail
D: Length of hind foot
E: Condylobasilar length
F: Alveolar length of upper molar tooth-row
G: Zygomatic breadth
H: Lambdoidal breadth
I: Prelambdoidal breadth

Locality | A | B | C | D | E | F | G | H | I
M. m. codiensis, all
Average | 34 | 165.3 | 44.2 | 19.6 | 25.47 | 6.56 | 15.55 | 12.05 | 8.76
2 × stand. error |  | 3.56 | 1.84 | .395 | .308 | .067 | .198 | .144 | .129
Coeff. variab |  | 6.0 | 11.6 | 5.6 | 3.5 | 3.0 | 3.65 | 1.20 | 1.47
M. m. nanus, Eastern Idaho
Average | 21[1] | 151.1 | 39.4 | 19.2 | 25.00 | 6.44 | 14.99 | 11.74 | 8.94
2 × stand. error |  | 3.20 | 2.89 | .293 | .286 | 1.15 | .295 | .210 | .182
Coeff. variab. |  | 6.1 | 21.1 | 4.38 | 2.49 | 3.99 | 4.10 | 3.79 | 4.31
M. m. nanus, Wyoming
Teton Co. | 35 | 160.5 | 40.7 | 18.6 | 25.16 | 6.51 | 15.17 | 11.86 | 8.77
Fremont Co. | 26 | 157.0 | 41.4 | 19.6 | 25.23 | 6.25 | 15.05 | 11.88 | 8.91
Lincoln Co. | 24 | 159.9 | 41.8 | 18.9 | 25.08 | 6.26 | 15.10 | 11.82 | 8.75
Uinta Co. | 26 | 162.4 | 41.3 | 19.0 | 25.33 | 6.42 | 15.31 | 12.16 | 8.89
Sweetwater Co. | 12 | 159.8 | 43.7 | 20.1 | 24.98 | 6.31 | 15.00 | 11.84 | 9.02
Natrona Co. | 40 | 159.6 | 41.0 | 19.6 | 25.04 | 6.40 | 15.00 | 11.84 | 8.93
Carbon Co. | 108 | 158.7 | 40.0 | 19.1 | 24.96 | 6.27 | 15.05 | 11.83 | 8.72
   Encampment ♂ | 27 | 161 | 41.7 | 18.9 | 25.1 | 6.16 | 15.2 | 11.9 | 8.7
   Encampment ♀ | 11 | 159 | 41.1 | 19.4 | 24.9 | 6.18 | 14.9 | 11.6 | 8.6
   Savery ♂ | 23 | 159 | 41.0 | 19.2 | 25.3 | 6.3 | 15.2 | 12.2 | 8.8
   Savery ♀ | 25 | 155 | 37.1 | 18.8 | 24.7 | 6.3 | 14.8 | 11.6 | 8.6
M. m. nanus
Northern Colo. | 8 | 163.1 | 42.4 | 19.6 | 25.20 | 6.44 | 14.86 | 11.70 | 8.56
M. m. fusus
Southern Colo. | 17[2] | 159.8 | 42.4 | 19.2 | 24.97 | 6.43 | 14.98 | 11.73 | 8.69

[1] For external parts, 34 individuals were used.

[2] For external parts, 29 individuals were used.

REFERENCES CITED

Bailey, V.

1900. Revision of American voles of the genus Microtus. N. Amer. Fauna, 17:1-88, June 6.

1917. A new subspecies of meadow mouse from Wyoming. Proc. Biol. Soc. Washington, 30:29-30, February 21.

Cary, M.

1911. A biological survey of Colorado. N. Amer. Fauna, 33:1-256, August 17.

1917. Life zone investigations in Wyoming. N. Amer. Fauna, 42:1-95, October 3.

Davis, W. B.

1939. The Recent mammals of Idaho. 400 p., front., illus., maps, diagrs., Caldwell, Id., The Caxton Printers Ltd., April 5.

Durrant, S. D.

1952. Mammals of Utah, taxonomy and distribution. Univ. Kans. Publ., Mus. Nat. Hist., 6:1-549, 91 figs, in text, 30 tables, August 10.

Elton, C.

1942. Voles, mice and lemmings; problems in population dynamics. 496 p., illus. (maps), tables, Oxford, The Clarendon Press, London.

Goin, O. B.

1943. A study of individual variation in Microtus pennsylvanicus pennsylvanicus. Jour. Mamm., 24:212-224, June 7.

Hall, E. R.

1938. Notes on the meadow mice Microtus montanus and Microtus nanus with descriptions of a new subspecies from Colorado. Proc. Biol. Soc. Washington, 51:131-134, August 23.

Hill, E. A.

1935. Cranial foramina in rodents. Jour. Mamm. 16(2):121-129.

Howell, A. B.

1924. Individual and age variation in Microtus montanus yosemite. Jour. Agr. Res., 28(10):977-1015, June 7.

Kellogg, R.

1922. A study of the California forms of the Microtus montanus group of meadow mice. Univ. California Publ. Zool., 21:245-274, April 18.

Merriam, C. H.

1891. Results of a biological reconnaissance of south-central Idaho. N. Amer. Fauna, 5:1-113, July 30.

Piper, S. E.

1909. The Nevada mouse plague of 1907-8. U. S. Dept. Agr., Farmers' Bul. 352, pp. 1-23, 9 figs.

Warren, E. R.

1942. The mammals of Colorado, their habits and distribution. Second (revised) edition, Univ. Oklahoma Press, Norman, xviii-330 p., front., 50 plates.

Transmitted March 22, 1954.

25-3560

Transcriber's Notes

One typographical error was corrected: "Castlford" was corrected to "Castleford" in "Twin Falls Co.: Castleford Fenced Plot".

*** END OF THE PROJECT GUTENBERG EBOOK SUBSPECIATION IN THE MEADOW MOUSE, MICROTUS MONTANUS, IN WYOMING AND COLORADO ***

Updated editions will replace the previous one—the old editions will be renamed.

Creating the works from print editions not protected by U.S. copyright law means that no one owns a United States copyright in these works, so the Foundation (and you!) can copy and distribute it in the United States without permission and without paying copyright royalties. Special rules, set forth in the General Terms of Use part of this license, apply to copying and distributing Project Gutenberg™ electronic works to protect the PROJECT GUTENBERG™ concept and trademark. Project Gutenberg is a registered trademark, and may not be used if you charge for an eBook, except by following the terms of the trademark license, including paying royalties for use of the Project Gutenberg trademark. If you do not charge anything for copies of this eBook, complying with the trademark license is very easy. You may use this eBook for nearly any purpose such as creation of derivative works, reports, performances and research. Project Gutenberg eBooks may be modified and printed and given away—you may do practically ANYTHING in the United States with eBooks not protected by U.S. copyright law. Redistribution is subject to the trademark license, especially commercial redistribution.

START: FULL LICENSE

THE FULL PROJECT GUTENBERG LICENSE

PLEASE READ THIS BEFORE YOU DISTRIBUTE OR USE THIS WORK

To protect the Project Gutenberg™ mission of promoting the free distribution of electronic works, by using or distributing this work (or any other work associated in any way with the phrase “Project Gutenberg”), you agree to comply with all the terms of the Full Project Gutenberg™ License available with this file or online at www.gutenberg.org/license.

Section 1. General Terms of Use and Redistributing Project Gutenberg™ electronic works

1.A. By reading or using any part of this Project Gutenberg™ electronic work, you indicate that you have read, understand, agree to and accept all the terms of this license and intellectual property (trademark/copyright) agreement. If you do not agree to abide by all the terms of this agreement, you must cease using and return or destroy all copies of Project Gutenberg™ electronic works in your possession. If you paid a fee for obtaining a copy of or access to a Project Gutenberg™ electronic work and you do not agree to be bound by the terms of this agreement, you may obtain a refund from the person or entity to whom you paid the fee as set forth in paragraph 1.E.8.

1.B. “Project Gutenberg” is a registered trademark. It may only be used on or associated in any way with an electronic work by people who agree to be bound by the terms of this agreement. There are a few things that you can do with most Project Gutenberg™ electronic works even without complying with the full terms of this agreement. See paragraph 1.C below. There are a lot of things you can do with Project Gutenberg™ electronic works if you follow the terms of this agreement and help preserve free future access to Project Gutenberg™ electronic works. See paragraph 1.E below.

1.C. The Project Gutenberg Literary Archive Foundation (“the Foundation” or PGLAF), owns a compilation copyright in the collection of Project Gutenberg™ electronic works. Nearly all the individual works in the collection are in the public domain in the United States. If an individual work is unprotected by copyright law in the United States and you are located in the United States, we do not claim a right to prevent you from copying, distributing, performing, displaying or creating derivative works based on the work as long as all references to Project Gutenberg are removed. Of course, we hope that you will support the Project Gutenberg™ mission of promoting free access to electronic works by freely sharing Project Gutenberg™ works in compliance with the terms of this agreement for keeping the Project Gutenberg™ name associated with the work. You can easily comply with the terms of this agreement by keeping this work in the same format with its attached full Project Gutenberg™ License when you share it without charge with others.

1.D. The copyright laws of the place where you are located also govern what you can do with this work. Copyright laws in most countries are in a constant state of change. If you are outside the United States, check the laws of your country in addition to the terms of this agreement before downloading, copying, displaying, performing, distributing or creating derivative works based on this work or any other Project Gutenberg™ work. The Foundation makes no representations concerning the copyright status of any work in any country other than the United States.

1.E. Unless you have removed all references to Project Gutenberg:

1.E.1. The following sentence, with active links to, or other immediate access to, the full Project Gutenberg™ License must appear prominently whenever any copy of a Project Gutenberg™ work (any work on which the phrase “Project Gutenberg” appears, or with which the phrase “Project Gutenberg” is associated) is accessed, displayed, performed, viewed, copied or distributed:

This eBook is for the use of anyone anywhere in the United States and most other parts of the world at no cost and with almost no restrictions whatsoever. You may copy it, give it away or re-use it under the terms of the Project Gutenberg License included with this eBook or online at www.gutenberg.org. If you are not located in the United States, you will have to check the laws of the country where you are located before using this eBook.

1.E.2. If an individual Project Gutenberg™ electronic work is derived from texts not protected by U.S. copyright law (does not contain a notice indicating that it is posted with permission of the copyright holder), the work can be copied and distributed to anyone in the United States without paying any fees or charges. If you are redistributing or providing access to a work with the phrase “Project Gutenberg” associated with or appearing on the work, you must comply either with the requirements of paragraphs 1.E.1 through 1.E.7 or obtain permission for the use of the work and the Project Gutenberg™ trademark as set forth in paragraphs 1.E.8 or 1.E.9.

1.E.3. If an individual Project Gutenberg™ electronic work is posted with the permission of the copyright holder, your use and distribution must comply with both paragraphs 1.E.1 through 1.E.7 and any additional terms imposed by the copyright holder. Additional terms will be linked to the Project Gutenberg™ License for all works posted with the permission of the copyright holder found at the beginning of this work.

1.E.4. Do not unlink or detach or remove the full Project Gutenberg™ License terms from this work, or any files containing a part of this work or any other work associated with Project Gutenberg™.

1.E.5. Do not copy, display, perform, distribute or redistribute this electronic work, or any part of this electronic work, without prominently displaying the sentence set forth in paragraph 1.E.1 with active links or immediate access to the full terms of the Project Gutenberg™ License.

1.E.6. You may convert to and distribute this work in any binary, compressed, marked up, nonproprietary or proprietary form, including any word processing or hypertext form. However, if you provide access to or distribute copies of a Project Gutenberg™ work in a format other than “Plain Vanilla ASCII” or other format used in the official version posted on the official Project Gutenberg™ website (www.gutenberg.org), you must, at no additional cost, fee or expense to the user, provide a copy, a means of exporting a copy, or a means of obtaining a copy upon request, of the work in its original “Plain Vanilla ASCII” or other form. Any alternate format must include the full Project Gutenberg™ License as specified in paragraph 1.E.1.

1.E.7. Do not charge a fee for access to, viewing, displaying, performing, copying or distributing any Project Gutenberg™ works unless you comply with paragraph 1.E.8 or 1.E.9.

1.E.8. You may charge a reasonable fee for copies of or providing access to or distributing Project Gutenberg™ electronic works provided that:

• You pay a royalty fee of 20% of the gross profits you derive from the use of Project Gutenberg™ works calculated using the method you already use to calculate your applicable taxes. The fee is owed to the owner of the Project Gutenberg™ trademark, but he has agreed to donate royalties under this paragraph to the Project Gutenberg Literary Archive Foundation. Royalty payments must be paid within 60 days following each date on which you prepare (or are legally required to prepare) your periodic tax returns. Royalty payments should be clearly marked as such and sent to the Project Gutenberg Literary Archive Foundation at the address specified in Section 4, “Information about donations to the Project Gutenberg Literary Archive Foundation.”

• You provide a full refund of any money paid by a user who notifies you in writing (or by e-mail) within 30 days of receipt that s/he does not agree to the terms of the full Project Gutenberg™ License. You must require such a user to return or destroy all copies of the works possessed in a physical medium and discontinue all use of and all access to other copies of Project Gutenberg™ works.

• You provide, in accordance with paragraph 1.F.3, a full refund of any money paid for a work or a replacement copy, if a defect in the electronic work is discovered and reported to you within 90 days of receipt of the work.

• You comply with all other terms of this agreement for free distribution of Project Gutenberg™ works.

1.E.9. If you wish to charge a fee or distribute a Project Gutenberg™ electronic work or group of works on different terms than are set forth in this agreement, you must obtain permission in writing from the Project Gutenberg Literary Archive Foundation, the manager of the Project Gutenberg™ trademark. Contact the Foundation as set forth in Section 3 below.

1.F.

1.F.1. Project Gutenberg volunteers and employees expend considerable effort to identify, do copyright research on, transcribe and proofread works not protected by U.S. copyright law in creating the Project Gutenberg™ collection. Despite these efforts, Project Gutenberg™ electronic works, and the medium on which they may be stored, may contain “Defects,” such as, but not limited to, incomplete, inaccurate or corrupt data, transcription errors, a copyright or other intellectual property infringement, a defective or damaged disk or other medium, a computer virus, or computer codes that damage or cannot be read by your equipment.

1.F.2. LIMITED WARRANTY, DISCLAIMER OF DAMAGES - Except for the “Right of Replacement or Refund” described in paragraph 1.F.3, the Project Gutenberg Literary Archive Foundation, the owner of the Project Gutenberg™ trademark, and any other party distributing a Project Gutenberg™ electronic work under this agreement, disclaim all liability to you for damages, costs and expenses, including legal fees. YOU AGREE THAT YOU HAVE NO REMEDIES FOR NEGLIGENCE, STRICT LIABILITY, BREACH OF WARRANTY OR BREACH OF CONTRACT EXCEPT THOSE PROVIDED IN PARAGRAPH 1.F.3. YOU AGREE THAT THE FOUNDATION, THE TRADEMARK OWNER, AND ANY DISTRIBUTOR UNDER THIS AGREEMENT WILL NOT BE LIABLE TO YOU FOR ACTUAL, DIRECT, INDIRECT, CONSEQUENTIAL, PUNITIVE OR INCIDENTAL DAMAGES EVEN IF YOU GIVE NOTICE OF THE POSSIBILITY OF SUCH DAMAGE.

1.F.3. LIMITED RIGHT OF REPLACEMENT OR REFUND - If you discover a defect in this electronic work within 90 days of receiving it, you can receive a refund of the money (if any) you paid for it by sending a written explanation to the person you received the work from. If you received the work on a physical medium, you must return the medium with your written explanation. The person or entity that provided you with the defective work may elect to provide a replacement copy in lieu of a refund. If you received the work electronically, the person or entity providing it to you may choose to give you a second opportunity to receive the work electronically in lieu of a refund. If the second copy is also defective, you may demand a refund in writing without further opportunities to fix the problem.

1.F.4. Except for the limited right of replacement or refund set forth in paragraph 1.F.3, this work is provided to you ‘AS-IS’, WITH NO OTHER WARRANTIES OF ANY KIND, EXPRESS OR IMPLIED, INCLUDING BUT NOT LIMITED TO WARRANTIES OF MERCHANTABILITY OR FITNESS FOR ANY PURPOSE.

1.F.5. Some states do not allow disclaimers of certain implied warranties or the exclusion or limitation of certain types of damages. If any disclaimer or limitation set forth in this agreement violates the law of the state applicable to this agreement, the agreement shall be interpreted to make the maximum disclaimer or limitation permitted by the applicable state law. The invalidity or unenforceability of any provision of this agreement shall not void the remaining provisions.

1.F.6. INDEMNITY - You agree to indemnify and hold the Foundation, the trademark owner, any agent or employee of the Foundation, anyone providing copies of Project Gutenberg™ electronic works in accordance with this agreement, and any volunteers associated with the production, promotion and distribution of Project Gutenberg™ electronic works, harmless from all liability, costs and expenses, including legal fees, that arise directly or indirectly from any of the following which you do or cause to occur: (a) distribution of this or any Project Gutenberg™ work, (b) alteration, modification, or additions or deletions to any Project Gutenberg™ work, and (c) any Defect you cause.

Section 2. Information about the Mission of Project Gutenberg™

Project Gutenberg™ is synonymous with the free distribution of electronic works in formats readable by the widest variety of computers including obsolete, old, middle-aged and new computers. It exists because of the efforts of hundreds of volunteers and donations from people in all walks of life.

Volunteers and financial support to provide volunteers with the assistance they need are critical to reaching Project Gutenberg™’s goals and ensuring that the Project Gutenberg™ collection will remain freely available for generations to come. In 2001, the Project Gutenberg Literary Archive Foundation was created to provide a secure and permanent future for Project Gutenberg™ and future generations. To learn more about the Project Gutenberg Literary Archive Foundation and how your efforts and donations can help, see Sections 3 and 4 and the Foundation information page at www.gutenberg.org.

Section 3. Information about the Project Gutenberg Literary Archive Foundation

The Project Gutenberg Literary Archive Foundation is a non-profit 501(c)(3) educational corporation organized under the laws of the state of Mississippi and granted tax exempt status by the Internal Revenue Service. The Foundation’s EIN or federal tax identification number is 64-6221541. Contributions to the Project Gutenberg Literary Archive Foundation are tax deductible to the full extent permitted by U.S. federal laws and your state’s laws.

The Foundation’s business office is located at 809 North 1500 West, Salt Lake City, UT 84116, (801) 596-1887. Email contact links and up to date contact information can be found at the Foundation’s website and official page at www.gutenberg.org/contact

Section 4. Information about Donations to the Project Gutenberg Literary Archive Foundation

Project Gutenberg™ depends upon and cannot survive without widespread public support and donations to carry out its mission of increasing the number of public domain and licensed works that can be freely distributed in machine-readable form accessible by the widest array of equipment including outdated equipment. Many small donations ($1 to $5,000) are particularly important to maintaining tax exempt status with the IRS.

The Foundation is committed to complying with the laws regulating charities and charitable donations in all 50 states of the United States. Compliance requirements are not uniform and it takes a considerable effort, much paperwork and many fees to meet and keep up with these requirements. We do not solicit donations in locations where we have not received written confirmation of compliance. To SEND DONATIONS or determine the status of compliance for any particular state visit www.gutenberg.org/donate.

While we cannot and do not solicit contributions from states where we have not met the solicitation requirements, we know of no prohibition against accepting unsolicited donations from donors in such states who approach us with offers to donate.

International donations are gratefully accepted, but we cannot make any statements concerning tax treatment of donations received from outside the United States. U.S. laws alone swamp our small staff.

Please check the Project Gutenberg web pages for current donation methods and addresses. Donations are accepted in a number of other ways including checks, online payments and credit card donations. To donate, please visit: www.gutenberg.org/donate.

Section 5. General Information About Project Gutenberg™ electronic works

Professor Michael S. Hart was the originator of the Project Gutenberg™ concept of a library of electronic works that could be freely shared with anyone. For forty years, he produced and distributed Project Gutenberg™ eBooks with only a loose network of volunteer support.

Project Gutenberg™ eBooks are often created from several printed editions, all of which are confirmed as not protected by copyright in the U.S. unless a copyright notice is included. Thus, we do not necessarily keep eBooks in compliance with any particular paper edition.

Most people start at our website which has the main PG search facility: www.gutenberg.org.

This website includes information about Project Gutenberg™, including how to make donations to the Project Gutenberg Literary Archive Foundation, how to help produce our new eBooks, and how to subscribe to our email newsletter to hear about new eBooks.

back

back