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North American Jumping Mice (Genus Zapus)

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A comprehensive taxonomic revision of the genus Zapus examines geographic distribution, paleontology, and species-level relationships across North America. Based on about 3,600 specimens and extensive measurements, it provides an annotated chronology of names, diagnostic characters, an artificial key, and systematic accounts for named species and subspecies (including treatment of trinotatus, princeps, and hudsonius). The work evaluates taxonomically useful morphological characters, documents nongeographic variation, compares related genera, and supplies tables, figures, and synonymies to facilitate identification and clarify subspecific limits.

Zapus hudsonius acadicus (Dawson)

Meriones acadicus Dawson, Edinburgh New Philos. Jour., new ser., 3:2, 1856.

Meriones labradorius, Dawson, Edinburgh New Philos. Jour., new ser., 3:2, 1856.

Jaculus hudsonius, Baird, Rept. Expl. and Surv…., 8 (pt. 1):433, July 14, 1858 (part—the part from Nova Scotia, Vermont, and New York).

Zapus hudsonius, Coues, Bull. U. S. Geol. and Geog. surv. of the territories, 2nd ser., No. 5:260, 1877 (part—the part from Nova Scotia, Vermont, and New York); Preble, N. Amer. Fauna, 15:17, August 8, 1899 (part—the part from New Brunswick, Nova Scotia, Maine, New Hampshire, Vermont, Massachusetts, and northeastern New York).

Zapus hudsonius canadensis, Batchelder, Proc. New England Zool. Club, 1:5, February 8, 1899 (part—the part from Keene Valley in Essex County of New York, and Orivell in Vermont); Anderson, Ann. Rept. Provancher Soc. Nat. Hist., Quebec, 1941:35-37, July 14, 1942 (part—the part from the tip of the Gaspé Peninsula in Quebec, New Brunswick, Maine, New Hampshire, Vermont, and New York).

Zapus hudsonius hardyi, Batchelder, Proc. New England Zool. Club, 1:6, February 8, 1899, type from Mt. Desert Island, Hancock County, Maine; Bole and Moulthrop, Sci. Publ. Cleveland Mus. Nat. Hist., 5:165, September 11, 1947 (part—but excluding Pennsylvania and Ohio).

Zapus hudsonius acadicus, Anderson, Ann. Rept. Provancher Soc. Nat. Hist., Quebec, 1941:38, July 14, 1942.

Type.—No type specimen designated. Subspecies characterized from specimens obtained in Nova Scotia.

Range.—Gaspé Peninsula of Quebec, New Brunswick, Nova Scotia, Prince Edward Island, Maine, New Hampshire, Vermont, Massachusetts, northern Connecticut and northeastern New York. See fig. 47. Zonal range: Transition and Canadian.

Description.—Size medium; back from near Ochraceous-Tawny to near Yellow-Ocher with heavy admixture of black-tipped hair, the dorsal band distinct against color of sides; sides lighter than back and from near Cinnamon-Buff to near Ochraceous-Buff lined with black-tipped hair; lateral line usually [433] faintly marked but sometimes distinct and clear Warm-Buff; underparts white, sometimes suffused with color of sides; tail distinctly bicolored, brownish-black above and yellowish-white to grayish-white below; ears dark, edged with color of sides; feet grayish-white above; pterygoid fossae relatively narrow; zygomata relatively long and broad; auditory bullae relatively narrow, usually with depression on anterior surface; mastoid region relatively narrow; inferior arm of zygomatic process of maxillary relatively narrow.

Comparisons.—From Zapus hudsonius canadensis, Z. h. acadicus differs in: Size averaging larger; upper parts usually less brownish and more ochraceous, sides and flanks being more ochraceous and less yellowish; zygomata relatively longer; pterygoid fossae relatively narrower; auditory bullae relatively narrower and usually with depression on anterior surface.

From Zapus hudsonius americanus, Z. h. acadicus differs as follows: Size larger; color darker on upper parts, flanks duller (less ochraceous); underparts white, much less frequently suffused with color of sides; ears dark, usually without flecks of ochraceous; general appearance of pelage not so brightly colored; zygomata longer; condylobasal length greater; mastoid region relatively broader; bullae larger, more inflated and usually with depression on anterior surface; maxillary tooth-row relatively longer.

For comparison with Zapus hudsonius ladas see account of that subspecies.

Remarks.—Specimens from various localities in Nova Scotia, Prince Edward Island and New Brunswick are essentially similar. Anderson (1942:38) revived the name Z. h. acadicus for jumping mice from these areas, correctly considering them to be distinct from Z. h. canadensis, the geographic race immediately to the west.

In the size and shape of the auditory bullae, length of the zygomata, breadth of the pterygoid fossae, and general color of the pelage the populations from Nova Scotia and New Brunswick are essentially indistinguishable from material of Zapus hudsonius hardyi from Maine. Thus, Z. h. hardyi must fall as a synonym of the earlier proposed name Z. h. acadicus.

Bole and Moulthrop (1942:165) applied the name Z. h. hardyi (= acadicus) to the mice inhabiting a large area from coastal Maine and central New Hampshire through southern New England, New York, northwestern Pennsylvania, and northeastern Ohio. I agree with Bole and Moulthrop (loc. cit.) that the population of Zapus hudsonius from Maine, New Hampshire, west-central and northern New England are different from neighboring subspecies and are referable to Z. h. acadicus, but find that material from extreme southern Massachusetts, Connecticut, southern New York, northwestern Pennsylvania, and northeastern Ohio is best referred to Zapus hudsonius americanus (see account of that subspecies).

Intergradation between Z. h. americanus and Z. h. acadicus is indicated by specimens from Berlin, Rensselaer County, New York. In color of ears, length of zygomata, and size and shape of the incisive foramina these specimens are more nearly like Z. h. americanus but in size and shape of the auditory bullae, breadth of the mastoid region, and general appearance of the pelage they are more nearly like Z. h. acadicus and are here referred to acadicus. Specimens from Glenville, Schenectady County, New York, are intermediate in cranial characters between Z. h. americanus and Z. h. acadicus but in color are best referred to the latter. Specimens from 1 mi. S Ayer, Worchester County, Massachusetts, are like Z. h. americanus in their short zygomata, narrow mastoid region and suffusion of the underparts; nevertheless, in the shape of the auditory bullae, breadth of the pterygoid fossae, and greater condylobasal length the specimens are more nearly like Z. h. acadicus which they are here considered to be. Animals from Essex and Wilmington, Essex County, Massachusetts, are like Z. h. americanus in external size and in the size and shape of the auditory bullae; but they are more nearly like Z. h. acadicus in most cranial characters and in the general color of the pelage and are here assigned to Z. h. acadicus.

Specimens from Keene Valley, Essex County, New York, considered by Batchelder (1899:4) to be Z. h. canadensis, are in color, length of the zygomata, and size and shape of the auditory bullae more nearly like Z. h. acadicus to which subspecies they are here assigned. A specimen from Orwell, Addison County, Vermont, that Batchelder (op. cit.:5) referred to Z. h. canadensis is more nearly like Z. h. acadicus in the shape of the auditory bullae, length of the zygomata, and color of the pelage, and is here referred to Z. h. acadicus. Specimens from western New Brunswick, referred to Z. h. canadensis by Anderson (1942:37), are more nearly like Z. h. acadicus. Specimens from Ste. Anne des Monts, Gaspé Peninsula, Quebec, are intermediate between Z. h. canadensis and Z. h. acadicus in color and size and also in the shape of the auditory bullae but are best referred to Z. h. acadicus.

Zapus hudsonius acadicus as here understood is a relatively wide-ranging subspecies. Populations at the southern periphery of its range are difficult to separate from populations at the northern periphery of the range of Z. h. americanus. These two geographic races represent opposite extremes of a clinal gradient and, as would be expected, geographic intermediates are morphologically similar.

Specimens examined.—Total, 156, distributed as follows:

Maine: Aroostock County: Madawaska, 6 (MCZ). Hancock County: Mount Desert Island, 9 (6 MCZ, 3 UM). Piscataquis County: Mount Katahdin, 1 (USNM); Sebec Lake, 4 (USBS); Katahdin Lake, 1 (USBS). Sagadahoc Co.: Small Point Beach, 1 (Clev. MNH). Somerset County: east [435] branch Penobscot River, 2 (USBS). Washington County: Columbia Falls, 1 (USBS).

Massachusetts: Essex County: Essex, 4 (Clev. MNH); Wilmington, 4 (3 USBS, 1 USNM). Worchester County: Lunenberg, 2 (USBS); 1 mi. S Ayer, 2 (MVZ); 2 mi. N Gilbertville, 1.

New Brunswick: Charlotte County: 6 mi. N St. Andrews, 2 (NMC); 5 mi. N St. Andrews, 4 (NMC). Carleton County: Debec, 1 (MVZ). Gloucester County: Dalhousie, 2 (MVZ); Miramichi Road, 15 mi. from Bathurst, 4 (NMC); Youghall, 3 (NMC). Madawaska County: Baker Lake, 2 (NMC); 9 mi. NE Edmundston, 4 (NMC); 5 mi. N St. Leonard, 5 (NMC). Victoria Co.: Tobique Point, 1 (AMNH). York County: Queensbury, 1 (USBS).

New Hampshire: Carroll County: Intervale, 1 (UM); Ossipee, 4 (3 USBS); 2 mi. S Ossipee, 12 (2 USNM). Coos County: Nathan Pond, 1 (UM); Fabyans-Bretton Woods, Dartmouth Brook, 2 (UM); Fabyans, 1 (USNM); 3 mi. W Base Station, 1; Mt. Washington, 1 (MVZ); Pinkham Notch, 1900 ft., 1 (USNM). Grafton County: Franconia Notch, Profile Lake, 1 (UM); Lebanon, 3 (UM). Strafford Co.: 1 mi. E Durham, 1 (UM).

New York: Essex Co.: Keene Valley, 5 (MCZ); Keene Heights, 5 (MCZ); Minerva, 1700 ft., 1 (AMNH). Herkimer County: Northwood, 7 (AMNH). Rensselaer Co.: Berlin, 8 (AMNH). Schenectady County: Glenville, 1 (USBS). Warren County: Lake George, 5 (USBS). Washington County: Patterns Mills, 1 (USBS).

Nova Scotia: Annapolis Co.: Bear River, 7 (NMC); Lake Kedgemakooge, 5 (UM); 2 mi. S Milford, 1 (AMNH). Kings Co.: Black River Dist., 1 (NMC); no exact locality, 1 (NMC). Shelburne County: Doctors Cove, N Barrington Passage, 1 (NMC); Barrington Passage, 4 (NMC).

Prince Edward Island: no exact locality, 1 (USBS).

Quebec: Ste. Anne des Monts, 1 (AMNH).

Vermont: Addison County: Orwell, 1 (MCZ); Lamville County: Mt. Mansfield, 2 (USBS). Windham County: Whitingham, 2 (AMNH).

Marginal records.—Quebec: Ste. Anne des Monts. New Brunswick: Dalhousie. Prince Edward Island. Nova Scotia: Black River District; Doctors Cove, N Barrington Passage. Maine: Columbia Falls; Small Point Beach. Massachusetts: Wilmington; 2 mi. N Gilbertville. New York: Berlin; North Wood; Keene Valley. Maine: E branch Penobscot River. New Brunswick: Baker Lake.

Zapus hudsonius alascensis Merriam

Zapus hudsonius alascensis Merriam, Proc. Biol. Soc. Washington, 2:223, July 15, 1897.

Zapus hudsonius hudsonius, Osgood, N. Amer. Fauna, 24:37, November 23, 1904.

Type.—Male, adult, skin and skull, No. 73584, U. S. Nat. Mus., Biol. Surv. Coll.; Yakutat Bay, Alaska; obtained on July 5, 1895, by Clark P. Streator, original No. 4660.

Range.—Alaska Peninsula, coastal section of mainland of southern and southeastern Alaska including Revillagigedo Island; also southwestern Yukon. See fig. 47. Zonal range: Canadian and Hudsonian.

Description.—Size large; back from near Ochraceous-Tawny to near Dresden Brown, sometimes darkened with black tipped hair usually with darker mid-dorsal area forming a band; sides lighter than back and from near Ochraceous-Tawny to near Clay Color; lateral line usually distinct, of clear Ochraceous-Buff; belly white, frequently with a slight suffusion of Ochraceous-Buff; tail bicolored, brownish to brownish-black above, white to yellowish-white below; ears dark, edged and flecked on the inner surface with color of sides; feet grayish-white [436] above; auditory bullae broad and moderately inflated; pterygoid fossae relatively broad; incisive foramina relatively long, zygomata relatively long and broadly bowed; mastoid region relatively broad; distance from incisors to postpalatal notch relatively great; occipitonasal length relatively great.

Comparisons.—From Zapus hudsonius tenellus, Z. h. alascensis differs as follows: Size larger; upper parts darker, less ochraceous; sides duller, less ochraceous more tawny; incisive foramina averaging longer; mastoid region broader; occipitonasal length greater; zygomata wider-spreading and longer; condylobasal length averaging greater; auditory bullae less broadly rounded; and distance from incisors to postpalatal notch averaging greater.

For comparison with Zapus hudsonius hudsonius see account of that subspecies.

Remarks.Zapus hudsonius alascensis is a fairly well marked subspecies retaining most of its characters throughout its range. Variation is noted in specimens from the southwest end of Dezadeash Lake, 2400 ft., Yukon Territory, and seems to be the result of intergradation between Zapus hudsonius hudsonius and Z. h. alascensis. These animals are like Z. h. hudsonius in the shape of the auditory bullae but are otherwise more nearly like Z. h. alascensis to which they are here assigned. Alaskan specimens from 7 mi. SSE Haines, and from a point 9 mi. W and 4 mi. N Haines average slightly larger than Z. h. alascensis in most measurements taken; however, in coloration they more nearly agree with Z. h. alascensis than with Z. h. hudsonius or Z. h. tenellus the geographic ranges of which adjoin that of Z. h. alascensis.

Specimens examined.—Total, 56, distributed as follows:

Alaska: Cook Inlet, Tyonek, 1 (USBS); head Chalitna River, 2 (USBS); Lake Clark, 4 (USBS); east side Chilkat River, 100 ft., 9 mi. W and 4 mi. N Haines, 8; Yakutat, 3 (USBS); Lake Iliamma, 1 (USBS); Lake Aleknagik, 1 (USBS); Kokwok, 1 (USBS); Nushagak River, 3 (USBS); Chilkat Peninsula, 10 ft., 7 mi. SSE Haines, 18; Nushagak, 3 (USBS); Chignik Bay, 1 (USBS); Portage Cove, Revillagigedo, 1 (MVZ); Izembek Bay, 1 (USBS); Frosty Peak, 1 (USBS).

British Columbia: west end Kelsall Lake, 2900 ft., 1; Stonehouse Creek, 51/2 mi. W junction Stonehouse Creek and Kelsall River, 4.

Yukon: SW end Dezadeash Lake, 2400 ft., 2.

Marginal records.—Alaska: Lake Aleknagik; head Chalitna River. Yukon: SW end Dezadeash Lake, 2400 ft. Alaska: E side Chilkat River, 100 ft., 9 mi. W and 4 mi. N Haines; Portage Cove, Revillagigedo Island; Yakutat; Cook Inlet, Tyonek; Chignik Bay; Frosty Peak.

Zapus hudsonius americanus (Barton)

Dipus americanus Barton, Trans. Amer. Philos. Soc., 4:115, 1799.

Jaculus americanus Wagler, Nat. Syst. Amphibien, 23, 1830.

Meriones microcephalus Harlan, Proc. Zool. Soc. London, p. 1, 1839, based on two specimens from "the farm of Mr. Beck, in Philadelphia County, a few miles northeast of the city [= Philadelphia, Pennsylvania]."

Jaculus hudsonius, Baird, Repts. Expl. and Surv. 111, 8 (pt. 1): 433, July 14, 1858 (part—the part from Massachusetts, Connecticut, New York, New Jersey, and Pennsylvania).

Zapus hudsonius, Coues, Bull. U. S. Geol. and Geog. Surv. of the territories, 2nd ser. No. 5:260, 1877 (part—the part from Massachusetts, Connecticut, New York, and Pennsylvania); Preble, N. Amer. Fauna, 15:17, August 8, 1899 (part—the part from Peterboro and Waterville, New York, southeastern Massachusetts, Connecticut, New Jersey, Pennsylvania, West Virginia, Maryland, North Carolina, and Ohio).

Zapus hudsonius americanus, Batchelder, Proc. New England Zool. Club, 1:6, February 8, 1899; Preble, N. Amer. Fauna, 15:19, August 8, 1899.

Zapus hudsonius hardyi, Bole and Moulthrop, Sci. Publ. Cleveland Mus. Nat. Hist., 5:165, September 11, 1942 (part—the part from New York, Ohio, and Pennsylvania).

Zapus hudsonius brevipes Bole and Moulthrop, Sci. Publ. Cleveland Mus. Nat. Hist., 5:168, September 11, 1942, type from Bettsville, Seneca County, Ohio.

Zapus hudsonius rafinesquei Bole and Moulthrop, Sci. Publ. Cleveland Mus. Nat. Hist., 5:169, September 11, 1942 (part—the part from southeastern Ohio), type from Cat Run, extreme southeastern Belmont County, Ohio.

Type.—No type specimen designated. Dipus americanus was characterized from jumping mice obtained by Barton near the Schuylkill River, a few miles from Philadelphia, Pennsylvania.

Range.—Southeastern United States and lower peninsula of Michigan; east of central Indiana; from central New York and Massachusetts southward to northern Georgia. See fig. 47. Zonal range: Austroriparian (Lower Austral), Carolinian (Upper Austral), Alleghanian (Transition), and Canadian.

Description.—Size small; back from near Light Ochraceous-Buff to near Ochraceous-Buff with admixture of black-tipped hair forming distinct dorsal band; sides bright, lighter than back from near Light Ochraceous-Buff to near Ochraceous-Buff; lateral line usually distinct and of color of sides; underparts white, sometimes with slight suffusion of color of sides; tail bicolored, brown to brownish-black above, yellowish-white to grayish-white below; ears narrowly edged and heavily flecked with color of sides; feet white to grayish-white above; skull short; braincase relatively narrow; incisive foramina relatively broad; skull relatively narrow across zygomata; interorbital region relatively broad; distance from incisors to postpalatal notch relatively short; auditory bullae relatively small.

Comparisons.—Compared with Zapus hudsonius canadensis, Z. h. americanus differs as follows: Smaller; paler (in a sense brighter because more ochraceous and less tawny); skull smaller; auditory bullae narrower, less inflated; incisive foramina relatively more bowed; condylobasal length averaging less.

From Zapus hudsonius intermedius, Z. h. americanus differs as follows: Smaller; color brighter, more ochraceous, less yellow; braincase relatively narrower; auditory bullae usually smaller; incisive foramina broader; inferior ramus of zygomatic process of maxillary usually with median projection; interorbital region averaging broader.

For comparison with Zapus hudsonius acadicus see account of that subspecies.

Remarks.—Intergradation with Zapus hudsonius acadicus occurs in southeastern New York as indicated by a series of 25 specimens from Peterboro. They resemble Z. h. acadicus in width of the mastoid region and relatively longer tooth-row, but in the size and shape of the auditory bullae, width of the pterygoid fossae, and lighter, brighter, color of the sides they are more nearly like Z. h. americanus to which they are here referred.

Intergradation between Z. h. americanus and Z. h. acadicus is indicated also by specimens from Lawyersville and Schoharie, New York. In animals from both localities the length of the zygomata and the breadth of the mastoid region are more nearly as in Z. h. acadicus, but in size and shape of the auditory bullae, over-all length of the skull, color of the ears, and general color of the pelage they are more nearly like Z. h. americanus to which they are here referred.

Specimens from western Pennsylvania, judged to be Z. h. hudsonius by Preble (1899:17), and those from northwestern Pennsylvania and northeastern Ohio, allocated to Z. h. hardyi (= acadicus) by Bole and Moulthrop (1942:165), are more nearly like Z. h. americanus in size and shape of the auditory bullae, short zygomata, relatively narrow mastoid region, and color of pelage.

Specimens from the lower peninsula of Michigan, northeastern Indiana, and northwestern Ohio, described by Bole and Moulthrop (op. cit.:168) as belonging to a new subspecies (Zapus hudsonius brevipes), are to me indistinguishable from most specimens of Z. h. americanus. The characters which Bole and Moulthrop (loc. cit.) ascribe to Z. h. brevipes—color bright Ochraceous-Buff, tail and hind feet short, and skull narrow—are also those of Z. h. americanus.

Specimens from various localities in southeastern Ohio, all within the range ascribed by Bole and Moulthrop (op. cit.:169) to Zapus hudsonius rafinesquei, are indistinguishable from specimens of Z. h. americanus from eastern Tennessee, West Virginia, North Carolina, and Maryland. Zapus hudsonius rafinesquei (at least that part from southeastern Ohio) is indistinguishable from Z. h. americanus and therefore is synonymized under Z. h. americanus.

Specimens from Lagrange County, Indiana, show intergradation between Zapus hudsonius intermedius and Z. h. americanus in the color of the pelage but are more nearly like Z. h. americanus to which they are here referred. One from Porter County, Indiana, is more nearly like Z. h. intermedius in size and shape of the bullae and in breadth of the pterygoid fossae but in color and degree of lateral bowing of the zygomata is better placed with Z. h. americanus.

Z. h. americanus is a wide ranging subspecies. Animals at the northern periphery of the range (lower peninsula of Michigan to the west and southeastern Massachusetts to the east) are largest and darkest; to the southward there is a progressive reduction in size and a change to a lighter, brighter color. Animals from Maryland, Virginia, and North Carolina are more nearly average representatives of the subspecies than are those from the region of the type locality.

A jumping mouse allegedly of this subspecies has been recorded by Coleman (1941:91) from Caesars Head, 300 ft., South Carolina. This specimen and one from Athens, Georgia, provide the southeasternmost record-stations of occurrence for the species Z. hudsonius.

Specimens examined.—Total, 318, distributed as follows:

Connecticut: Hartford County: Windsor, 1 (USBS); East Hartford, 2 (MCZ). Litchfield County: Sharon, 3 (AMNH); Macedonia Park, 2 (AMNH). Middlesex County: Clinton, 1 (AMNH). Windham County: South Woodstock, 10 (AMNH); Pomfret, near Hampton line, 1.

Georgia: Clarke Co.: Athens, 1 (USBS).

Indiana: Lagrange Co.: no exact locality, 2 (UM). Porter Co.: Mineral Springs, 1 (FM); no exact locality, 1 (FM).

Maryland: Anne Arundel County: Patuxent Research Refuge, 1 (USBS). Charles County: no exact locality, 1 (USBS). Garrett Co.: Finzel, 6 mi. N Frostburg, 1 (USBS). Montgomery County: Sandy Springs, 2 (USBS); Kensington, 1 (USNM); Cabin John Bridge, 2 (1 USBS; 1 USNM). Prince Georges County: Laurel, 8 (USNM); Branchville, 1 (USBS); College Park, 1. Worchester County: Assateague, 5 mi. S Ocean City, 1 (USBS).

Massachusetts: Barnstable County: West Falmouth, 1 (USBS). Bristol County: Raynham, 1 (Clev. MNH). Dukes County: Martha’s Vineyard, 1 (USBS); West Chop, Martha’s Vineyard, 1 (Clev. MNH). Nantucket County: Nantucket Island, 1 (USNM). Plymouth County: Middleboro, 1 (USNM); Plymouth, 1 (UM); Marshfield, 6 (USBS); Wareham, 3 (1 Clev. MNH; 2 UM).

Michigan: Alcona Co.: 2 mi. S Harrisville, 2 (UM). Allegan Co.: near junction Swan Creek and Kalamazoo River, 3 (UM). Berrien Co.: Warren Woods, 2 (UM); Three Oaks, 1 (UM). Charlevoix Co.: Thumb Lake, 1 (UM); Section 1 Norwood Township, 1 (UM); Boyne Falls, 12 (UM); 2 mi. S Boyne Falls, 2 (UM). Cheboygan Co.: Douglas Lake, 2 (UM). Clinton Co.: 2 mi. SE DeWitt, 1 (UM). Emmet Co.: Maple River, near Douglas Lake, 1 (UM). Huron Co.: Rush Lake, 1 (UM). Kalamazoo Co.: no exact locality, 1 (UM). Lake Co.: 1 mi. NW Chase, 1 (UM). Livingston Co.: George Reserve, Pinckney, 2 (UM); Upper Whitewood Lake, 1 (UM); Whitmore Lake, 1 (UM); Portage Lake, 3 (UM). Mason Co.: 9 mi. N Ludington, 1 (UM). Midland Co.: Sanford, 1 (UM). Montmorency Co.: T. 32N, R. 1E, Sec. 30, 1 (UM). Muskegon Co.: 4 mi. NW North Muskegon, 2 (UM). Oakland Co.: Bloomfield, 1 (UM); no exact locality, 1 (UM). Otsego Co.: Pigeon River, 1 (UM); T. 32N, R. 1W, Sec. 25, 1 (UM); Waters, 1. Roscommon Co.: T. 24N, R. 2W, Sec. 2, 1 (UM). Shiawassee Co.: 1/2 mi. NE Byron, 5 (UM); 1/4 mi. S Byron, 2 (UM); 2 mi. SE Byron, 1 (UM); 3 mi. SW Byron, 1 (UM). Van Buren Co.: Van Auken Lake, 1 (UM). Washtenaw County: Whitmore Lake, 1 (UM); 2 mi. W Cherry Hill, 1 (UM); Ann Arbor, 7 (UM); 2 mi. E Ann Arbor, 2 (UM); Willow Run Village, 1 (UM).

[440]

New Jersey: Bergen County: Harrington Park, 1 (AMNH); Englewood, 1 (USNM). Cape May County: Mays Landing, 3 (Clev. MNH). Morris County: Mendham, 1 (AMNH). Ocean County: Tuckerton, 3 (USBS).

New York: Broome Co.: 5 mi. N Binghamton, 2 (USNM). Cayuga County: E Aurora, 1 (USBS). Greene County: Catskills, 4 (USNM); Kaaterskill Junction, 1 (USNM). Madison County: Peterboro, 25 (2 MCZ; 19 USNM; 4 Clev. MNH). Nassau County: Locust Grove, 3 (USNM). Orange Co.: Cranberry Pond, 840 ft., Highland, 2 (USNM). Otsego County: Lake Charlotte, 1 (AMNH). Queens County: Woodside, Long Island, 1 (USNM); near Forest Hills, Long Island, 1 (AMNH); Ray Nu Beach, Long Island, 1 (USNM). Rockland County: Tappan, 1 (AMNH). Schoharie County: Lawyersville, 1 (AMNH); Schoharie, 1 (AMNH). Suffolk County: Montauk Point, Long Island, 8 (USBS). Tioga County: Owego, 1 (USBS). Westchester Co.: Bedford, 1 (AMNH).

North Carolina: Buncombe County: Weaverville, 1 (AMNH). Cherokee Co.: Martin Creek, 2 (UM). Mitchell County: Roan Mountain, 2 (USBS). Wake County: Raleigh, 5 (3 USNM; 1 UM; 1 NCS).

Ohio: Carroll Co.: Carrollton, 2 (UM). Cuyahoga County: Big Creek, Brookside Park, 1 (Clev. MNH); Dover, 1 (Clev. MNH); Rocky River Metr. Park, 3 (Clev. MNH); North Olmstead, 1 (Clev. MNH). Erie Co.: Milan, 1 (Clev. MNH); Mill Hollow, Vermilion River, 1 (Clev. MNH). Lake Co.: Holden Arboretum, 3 (Clev. MNH). Meigs Co.: Portland Station, 1 (Clev. MNH). Seneca Co.: Bettsville, 4 (Clev. MNH); Old Fort Seneca, 4 (Clev. MNH); Corners, 1 (Clev. MNH). Wayne Co.: Wooster, 1 (UM); Craighton, 1 (UM).

Pennsylvania: Beaver Co.: 1 mi. NE Darlington, 1 (CM); 2 mi. E Industry, 1 (CM); 4 mi. E Frankfort, 2 (CM). Bedford Co.: 1 mi. NE Osterburg, 1 (CM). Berks Co.: 2 mi. W Strausstown, 1 (USNM). Bradford Co.: 21/2 mi. NNW Wyalusing, 2 (CM). Bucks Co.: 2 mi. N New Britain, 1 (CM). Butler Co.: Thorn Creek, 4 mi. S Butler, 4 (CM); 2 mi. E Middle Lancaster, 1 (CM); Orphans Home, 2 mi. E Mars, 2 (CM). Cambria Co.: 21/2 mi. S Patton, 1750 ft., 1 (CM); 51/2 mi. NE Ebensburg, 1 (CM). Centre Co.: 2, mi. E Snowshoe, 2 (CM). Chester Co.: 2 mi. S West Chester, 1 (CM). Clinton Co.: Tamarack, 9 mi. NNW Renovo, 1 (CM). Crawford Co.: Pymatuning Lake, 3 (Clev. MNH). Erie Co.: 41/2 mi. SW [town of] North East; 2 (CM); East Springfield, 1 (CM). Fulton Co.: 11/2 mi. NE Warfordsburg, 580 ft., 1 (CM). Huntington Co.: 61/2 mi. S Shade Gap, 2 (CM). Indiana Co.: 1/2 mi. E Indiana, 1320 ft., 2 (CM). Lebanon Co.: 11/2 mi. SE Cornwall, 800 ft., 1 (CM). Mercer Co.: 21/2 mi. W Mercer, 2 (CM); 5 mi. S Mercer, 1 (CM). Monroe Co.: Pocene Lake, 1(CM). Pike Co.: Bruce Lake, 1 (CM). Potter Co.: Woodcock Run, 71/2 mi. WSW Ulysses, 2 (CM). Sommerset County: 4 mi. SW Somerset, 2100 ft., 2 (CM); New Lexington, 1 (USBS). Susquehanna Co.: 10 mi. NNW Montrose, 1 (CM). Union Co.: Glen Iron, 2 (CM). Warren Co.: Bensons Swamp, 5 mi. E Columbus, 1 (USNM); Miles Run, 5 mi. NW Pittsfield, 1 (CM); 11/2 mi. N Pittsfield, 1 (CM); 21/2 mi. N Kinzua, 2 (CM); 2 mi. N Kinzua, 1 (CM).

Tennessee: Carter Co.: 3 mi. SSW Roan Mountain (town), 2900 ft., 1 (UM).

Virginia: Amelia Co.: Amelia, 1 (UM). Elizabeth City County: Near Hampton, 2 (UM). Fairfax County: Fall Church, 4 (2 USNM; 2 USBS); opposite Plummers Island, Maryland, 1 (USNM). Highland Co.: Laurel Park, 9 mi. NNW Monterey, 3100 ft., 4 (UM). Nelson Co.: no exact locality, 5 (USNM). Norfolk County: Deep Creek, 1 (USBS). Page Co.: no exact locality, 1 (USNM). Smyth Co.: Sugar Grove, 1 (UM); 1/2 mi. E Konnarock, 2800 ft., 1 (UM). Washington Co.: Konnarock, 2900 ft., 1 (UM).

Washington D. C.: Chevy Chase, 1 (USBS); no exact locality, 4 (3 USNM; 1 USBS).

West Virginia: Monongalia Co.: Morgantown, 6.

Marginal records.—Michigan: Douglas Lake; Bloomfield. New York: E Aurora; Peterboro; Catskills. Connecticut: Sharon; South Woodstock. Massachusetts: [441] Middleboro. New Jersey: Tuckerton. Maryland: Assateague, 5 mi. S Ocean City. North Carolina: Raleigh. Georgia: Athens. Indiana: Mineral Springs. Michigan: 9 mi. N Ludington.

Zapus hudsonius campestris Preble

Zapus hudsonius campestris Preble, N. Amer. Fauna, 15:20, August 8, 1899.

Type.—Male, adult, No. 65872 U. S. Nat. Mus., Biol. Surv. Coll.; Bear Lodge Mt’s [Crook County], Wyoming; obtained on June 21, 1894, by B. H. Dutcher, original No. 600.

Range.—Southeastern Montana, southwestern South Dakota, and northeastern Wyoming. See fig. 47. Zonal range: Transition.

Description.—Size large; back from near Ochraceous-Tawny to near Ochraceous-Buff with admixture of black tipped hair forming distinct dorsal band; sides lighter than back, from near Ochraceous-Buff to near Yellow Ocher with black hair interspersed; lateral line usually distinct, of clear Ochraceous-Buff; belly white, usually with moderate suffusion of Ochraceous-Buff; tail bicolored, brownish to brownish-black above, grayish-white to yellowish-white below; ears dark, edged with Ochraceous-Buff; feet grayish-white above; auditory bullae large, well inflated; incisive foramina long and usually truncate at posterior border; pterygoid fossae broad; zygomata relatively wide-spread and long; large medial projection on inferior ramus of zygomatic process of maxillary; condylobasal length and occipitonasal length relatively great; mastoid region and palatal region relatively broad; interparietal bone usually broad.

Comparisons.—From Zapus hudsonius pallidus, Z. h. campestris differs as follows: Coloration darker (more black and yellow but less orange); averaging larger in all measurements taken except in least interorbital constriction and distance from incisors to postpalatal notch which are slightly larger and breadth across zygomatic arches which is same; zygomatic arch heavier; incisive foramina larger; interparietal bone broader.

Compared with Zapus hudsonius intermedius, Z. h. campestris differs as follows: Coloration more tawny and ochraceous, less yellow; auditory bullae averaging larger, more inflated; condylobasal length averaging greater; zygomata averaging more wide-spread and longer; distance from incisors to postpalatal notch averaging longer; mastoid region broader; incisive foramina longer and more truncate posteriorly.

From Zapus hudsonius hudsonius, Z. h. campestris differs as follows: Size larger; color lighter, more ochraceous, less tawny; occipitonasal length averaging greater; mastoid region broader; zygomata averaging longer; zygomatic arch more widely bowed; distance from incisors to postpalatal notch averaging longer; incisive foramina longer; auditory bullae broader, more inflated.

For comparison with Zapus hudsonius preblei see account of that subspecies.

Remarks.—Animals from the Black Hills of South Dakota and Wyoming are thought of as most characteristic of this geographic race. Intergradation is noted with Zapus hudsonius pallidus and is discussed in the account of that subspecies.

Specimens examined.—Total, 66, distributed as follows:

Montana: Big Horn County: Rotten Grass Creek, north base Big Horn Mts., 2 (USBS); Little Big Horn River, 2 mi. from Wyoming line, 1 (USBS).

South Dakota: Custer County: Custer, 3 (USNM); Bull Springs, 6 (Clev. MNH); Beaver Creek, Wind Cave Nat’l Park, 1 (UM); Wind Cave Nat’l Park [442] Game Ranch, Cold Spring Creek, Wind Cave Nat’l Park, 2 (UM); Pennington County: Rapid Creek, 2 mi. W Pactola, 4800 ft., 3 (UM); Castle Creek, R. 2E, T. 1N, 6500 ft., 3 (UM); Nelsons Place, 3 mi. SE Hill City, 6 (UM); Palmer Gulch, 4 mi. SE Hill City, 3 (UM); Palmer Gulch, 9 (FM); no definite locality, 4 (UM).

Wyoming: Crook County: Devils Tower, flood plain Belle Fourche River, 3350 ft., 1 (USBS); Bear Lodge Mts., 4 (USBS); 15 mi. N Sundance, Black Hills Nat’l Forest, 5500 ft., 2; 3 mi. NW Sundance, 5900 ft., 17; Sundance, 2 (USBS). Weston Co.: 11/2 mi. E Buckhorn, 6150 ft., 5.

Marginal records.—Montana: Rotten Grass Creek, N base Big Horn Mts. South Dakota: Nelsons Place, 3 mi. SE Hill City; Wind Cave Nat’l Park Game Ranch, Cold Spring Creek. Wyoming: 11/2 mi. E Buckhorn, 6150 ft.

Zapus hudsonius canadensis (Davies)

Dipus canadensis Davies, Trans. Linn. Soc. London, 4:157, 1798.

Zapus hudsonius hudsonius, Preble, N. Amer. Fauna, 15:17, August 8, 1899 (part—the part from Ontario).

Zapus hudsonius canadensis, Batchelder, Proc. New England Zool. Club, 1:5, February 8, 1899 (part—the part from Quebec); Anderson, Rept. Provancher Soc. Nat. Hist., Quebec, 1941:35-37, July 14, 1942 (part—the part from Quebec excepting the Gaspé Peninsula).

Zapus hudsonius ontarioensis Anderson, Ann. Rept. Provancher Soc. Nat. Hist., Quebec, 1942:59, September 7, 1943, type from Pancake Bay (Batchawana Bay) southeast end of Lake Superior, Algoma District, about 40 miles northeast of Sault Ste-Marie, Ontario.

Type.—No type specimen designated, subspecies characterized on the basis of two specimens obtained by Major General Thomas Davies within a few miles of the city of Quebec.

Range.—Eastern Ontario and western Quebec from Hudson Bay southward to the Great Lakes and into northwestern New York. See fig. 47. Zonal range: Transition and Canadian.

Description.—Size medium; back from near Clay Color to near Cinnamon-Buff with admixture of black hair usually forming a dorsal band; sides from near Clay Color to near Cinnamon-Buff and lighter than back; lateral line usually distinct, and clear Cinnamon-Buff; belly white, sometimes with slight suffusion of Cinnamon-Buff mid-ventrally; tail bicolored, brownish to brownish-black above, grayish-white to yellowish-white below; ears dark, sometimes flecked with color of the sides, edged with Cinnamon-Buff; feet grayish-white above; auditory bullae large, relatively broad and flat; incisive foramina relatively short and narrow, widest posteriorly; zygomata not widely bowed outward; mastoid region relatively wide; frontal region well inflated; nasals relatively narrow, short, and parallel sided.

Comparisons.—From Zapus hudsonius hudsonius, Z. h. canadensis differs as follows: Upper parts generally dull averaging lighter, less black tipped hair; sides also lighter with less suffusion of dark hair; frontal region more inflated; mastoid region averaging broader; auditory bullae broader; distance from incisors to postpalatal notch averaging slightly longer.

For comparison with Zapus hudsonius acadicus, Zapus hudsonius ladas, and Zapus hudsonius americanus see accounts of those subspecies.

Remarks.—Bole and Moulthrop (1942:165) refer 2 specimens from Elba, New York, to Z. h. hardyi (= acadicus); they are more nearly like Z. h. canadensis in size and shape of the auditory bullae and general color of the pelage. A specimen from Spectacle Pond, New York, has the narrow pterygoid fossae and relatively narrow auditory bullae of Z. h. acadicus and the relatively short, narrow incisive foramina, inflated frontal region, and color of Z. h. canadensis to which the specimen is here referred. Intergradation is noted also in animals from Schreiber, Ontario. They resemble Zapus hudsonius hudsonius in their darker coloration and shape of auditory bullae but in the remainder of the characters studied resemble Z. h. canadensis to which they are referred. Specimens from Notre Dame de la Dore and 1/2 mi. N Mistassini Post, Quebec, in size and shape of the auditory bullae and in width of the pterygoid fossae, closely approach Z. h. ladas but in color, distinct dorsal band, and in narrower zygomata are all nearest Z. h. canadensis to which subspecies they are here referred.

Zapus hudsonius ontarioensis Anderson (1942:59) from eastern Ontario was based chiefly, in comparison with Z. h. canadensis, upon, “dorsal stripe less distinct and sides somewhat duller yellowish with more admixture of blackish hairs.” Examination of 68 of the 69 specimens from the type locality shows that 58 are subadult and in subadult pelage. Individuals which are adult are indistinguishable in color of pelage and in cranial features from comparable material from southern Quebec. Z. h. ontarioensis is, therefore, considered to be a synonym of Z. h. canadensis.

Specimens examined.—Total, 123, distributed as follows:

New York: Franklin Co.: Spectacle Pond, Brighton Township, 2 (AMNH). Genesee Co.: Elba, 2 (Clev. MNH).

Ontario: Schreiber, 2 (NMC); Franz, 5 (MVZ); Pancake Bay, Algoma District, 68 (NMC); Maclennan, Algoma District, 3 (ROM); Cache Lake, Algonquin Park, 1 (MVZ); Experimental Farm, Ottawa, 1 (NMC); Dows Swamp, Ottawa, 1 (NMC); Apple Hill, 1 (NMC); Clear Lake, Arden, 1 (NMC); Athens, 1 (NMC); Aurora, 4 (Clev. MNH); Pattageville, Toronto, 1; Lorne Park, Toronto, 1 (NMC); Credit, 2 (NMC); Pickering, 1 (MVZ); Preston, 1 (NMC); St. Thomas, 1 (NMC).

Quebec: Notre Dame de la Dore, 3 (NMC); 1/2 mi. N Mistassini Post, 1 (NMC); Lake Albanel, 1 (NMC); St. Felicien, 3 (NMC); Valcartier, 8 (NMC); Kiamika Lake, 4 (NMC); Ste. Veronique, 2 (NMC); Val Jalbert, 2 (NMC); St. Methode, 1 (NMC).

Marginal records.—Quebec: 1/2 mi. N Mistassini Post; Valcartier. New York: Spectacle Pond, Brighton Township; Elba. Ontario: St. Thomas; Pancake Bay, Algoma Dist.; Franz; Schreiber. Quebec: Kiamika Lake.

Zapus hudsonius hudsonius (Zimmerman)

Dipus hudsonius Zimmerman, Geog. Geschichte d Menschen u. vierfussigen Thiere, 2:358, 1780.

Dipus labradorius Kerr, Animal Kingdom:276 (based on the Labrador Jerboid Rat of Pennant, 1781—but Preble, N. Amer. Fauna, 15:11, August 8, 1899, states that the specimen came from Hudson Bay), 1792.

Gerbillus canadensis, Desmarest, Mammalogie, 2:321, 1822.

Gerbillus labradorius, Harlan, Fauna Amer., p. 157, 1825.

Meriones labradorius, Richardson, Fauna Boreali-Americana, 1:144, 1829.

Jaculus labradorius Wagner, Suppl. Schreber’s Saugthiere, 3:294, 1843.

Zapus hudsonius hudsonius, Preble, N. Amer. Fauna, 15:15, August 8, 1899 (part—the part from Northwest Territory, Ontario, Michigan, northern Wisconsin and northern Minnesota).

Zapus hudsonius alascensis, Osgood, N. Amer. Fauna, 19:38, October 6, 1900.

Type.—Type specimen not known to be in existence; from Hudson Bay, locality now considered to be Fort Severn, Ontario (see Anderson, 1942:37).

Range.—Central Alaska southeastward to central Ontario, northern Minnesota, northern Wisconsin, and upper peninsula of Michigan. See fig. 47. Zonal range: Hudsonian, Canadian, and into Transition.

Description.—Size medium; back dark, from near Tawny-Olive to near Cinnamon with heavy admixture of black hair forming dorsal band; sides lighter than back and from near Tawny-Olive to near Cinnamon, sometimes with admixture of black hair giving sides streaked appearance; lateral line usually distinct, clear Ochraceous-Buff; underparts white, sometimes with slight suffusion of Ochraceous-Buff; tail bicolored, brown to brownish-black above, grayish-white to yellowish-white below; ears dark, usually edged with ochraceous; feet grayish-white above; incisive foramina relatively short and broadly rounded; zygomata relatively short; braincase relatively broad; auditory bullae flat, long, and relatively broad; pterygoid fossae relatively narrow; nasals relatively broad and short.

Comparisons.—From Zapus hudsonius alascensis, Z. h. hudsonius differs as follows: upper parts generally darker, more black tipped hair; sides darker with greater suffusion of dark hair; lateral line brighter, more distinct; size averaging smaller; zygomatic arches less bowed outward; distance from incisors to postpalatal notch shorter; zygomata shorter; occipitonasal length less; mastoid region narrower.

From Zapus hudsonius intermedius, Z. h. hudsonius differs in: color darker, more tawny dorsally; sides averaging darker, more black-tipped hairs; size averaging larger; braincase averaging broader; distance from incisors to postpalatal notch averaging slightly shorter; zygomata averaging longer; mastoid region averaging broader; incisive foramina averaging shorter.

From Zapus hudsonius tenellus, Z. h. hudsonius differs as follows: upper parts averaging darker; tail averaging shorter; condylobasal length averaging more; braincase averaging broader; auditory bullae broader and less inflated; interparietal averaging broader; incisive foramina more broadly rounded and averaging longer.

For comparison with Zapus hudsonius canadensis and Zapus hudsonius campestris see accounts of those subspecies.

Remarks.—Preble (1899:16) had available for study five specimens of Zapus hudsonius hudsonius from Hudson Bay. Four were preserved in alcohol and one as an incomplete skin (prepared from an alcoholic specimen). All were unreliable for comparative purposes owing to the effects of the preservative. Preble, therefore, (loc. cit.) selected as a fairly typical sample a series of specimens from Tower, St. Louis County, Minnesota; these formed the basis of comparison between Z. h. hudsonius and other subspecies of Zapus hudsonius. Now that additional material (well prepared skins and skulls) is available from the Hudson Bay region and from other localities in northern and western Canada it is evident that the specimens from Tower, although here considered to be Z. h. hudsonius, are not typical Z. h. hudsonius but are intergrades between hudsonius and specimens of Zapus hudsonius intermedius. Comparisons made in the present account are based on specimens from the vicinity of Hudson Bay (Fort Severen, Ontario, York Factory, Shamatawa River, and Robinson Portage, Manitoba). These individuals are considered typical of this subspecies. With these new data available the range of Z. h. hudsonius is now understood to include all of the region from eastern Alaska to the northern parts of Minnesota, Wisconsin, and Michigan.

Intergradation between Zapus hudsonius canadensis and Z. h. hudsonius is noted in specimens from 30 mi. NE Port Arthur and also in those from Silver Islet, Thunder Cape, Ontario. These individuals resemble Z. h. canadensis in size and shape of the auditory bullae and in the shape of the nasals, but in their darker coloration, broadly rounded incisive foramina, and relatively narrow pterygoid fossae they are more nearly like Z. h. hudsonius to which they are here referred.

Specimens from Minaki, Ontario, are tending toward Zapus hudsonius intermedius in lighter coloration but in the size and shape of the auditory bulla, size and shape of the incisive foramina, and in the width of the pterygoid fossae they are more nearly like Z. h. hudsonius to which they are here referred. Specimens from various localities in Menominee County, Michigan, are like Z. h. intermedius in shape of the incisive foramina and shape of the postpalatal notch, but in color of pelage, size and shape of the auditory bullae, and breadth of the pterygoid fossae they closely resemble Z. h. hudsonius.

In Wisconsin, intergradation occurs in color and in cranial characters in specimens from Mercer, Solon Spring, and in a single individual from Basswood Lake. All these specimens, however, are best referable to Z. h. hudsonius.

Specimens from one mile southwest of Fairbanks and from Fairbanks, Alaska, show intergradation with Zapus hudsonius alascensis in coloration (more brown, less black), but in small size, short, broadly rounded incisive foramina, and in size and shape of the auditory bullae are nearest to Z. h. hudsonius to which they are here assigned.

Intergradation with Zapus hudsonius alascensis is noted also in specimens from McIntyre Creek, Yukon. They are like Z. h. alascensis in the size and shape of the auditory bullae and in the more elongate incisive foramina, but in the coloration, size of the pterygoid fossae, and breadth of the braincase are more nearly like Z. h. hudsonius and are here referred to this geographic race.

In British Columbia, in specimens from 1 mi. NW junction of Irons Creek and Laird River as well as in those from Hot Springs, 3 mi. WNW junction of Trout River and Laird River, and in those from 1/4 mi. S of the junction of the same rivers, three way intergradation occurs. These animals are like Zapus hudsonius alascensis in color and in length of tail. They agree with Zapus hudsonius tenellus in shape of nasals. In degree of inflation of auditory bullae, in length and width of incisive foramina, and in shape of pterygoid fossae they are as in Z. h. hudsonius to which they are here assigned.