Part 2

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Howell (op. cit.:75) referred three specimens from Glacier Basin, on the northeastern part of Mount Rainier, to T. a. ludibundus as he did also one specimen (loc. cit.) from Reflection Lakes, on the southern flank of the mountain. Our comparisons indicate the correctness of Howell's identification of the specimens from Glacier Basin; they more closely resemble ludibundus than caurinus. The specimen from Reflection Lakes, however, is only one of five or six from the same place; the others were lumped by him among the 49 that he recorded from Mount Rainier under the name caurinus. The series from Reflection Lakes, so far as we can detect, is not unusually variable and the differences that are apparent are within the normal range of variation ascribable to season, age, and individualism. Also, the series from Reflection Lakes, to us, is not appreciably different from the other series, representing the following places on Mount Rainier: Indian Henrys, 5300 ft.; W slope Mt. Rainier, 6600 ft.; St. Andrews Park, 5500 ft.; Spray Park, 5500 ft.; Paradise Park; Muddy Fork of Cowlitz River; Sunset Park, 5000 ft.; ridge between St. Andrews Park and South Puyallup River, 6000 ft.; and Owyhigh Lakes, 5350 ft.

Collectively, or individually, where there are as many as six specimens from a place, the material from Mt. Rainier (Glacier Basin excepted) is intermediate in color between T. a. ludibundus and T. a. caurinus and no more closely resembles one subspecies than the other. As may be seen from the cranial measurements recorded above, specimens from Mt. Rainier, although intermediate between the two subspecies just mentioned, resemble ludibundus in lesser zygomatic breadth and lesser cranial breadth (and, it may be added, in lesser dorsolateral inflation of the braincase), but resemble caurinus in longer skull (occipitonasal length), longer nasals and greater breadth across the rows of upper molariform teeth.

In summary: The animals from Mount Rainier, in features of taxonomic import, are almost exactly intermediate between T. a. caurinus and T. a. ludibundus. Being influenced by considerations of geographic adjacency, we refer the animals on Mount Rainier to Tamias amoenus ludibundus (Hollister).

Dalquest's (op. cit.: 85) explanation of the probable origin of Tamias amoenus caurinus is pertinent here. He writes: "The chipmunks of the Olympic Mountains [caurinus] probably reached their present range from the Cascades. Their probable path of emigration was westward from Mt. Rainier, along the glacial outwash train of Nisqualli Glacier, to the moraine and outwash apron of the Vashon Glacier and thence to the Olympics. So similar are the chipmunks of Mt. Rainier and the Olympic Mountains that Howell (1929) included Mt. Rainier in the range of caurinus."

~Tamias townsendii cooperi~ Baird

Some uncertainty exists concerning the subspecific identity of the Townsend Chipmunk in southern Washington because Dalquest (Univ. Kansas Publ. Mus. Nat. Hist., 2:262, April 9, 1948) identified as Tamias townsendii cooperi specimens that he examined from Yocolt, a place well within the geographic range of T. t. townsendii as defined by A. H. Howell (N. Amer. Fauna, 52: fig. 7, p. 107, November 30, 1929). Dalquest (op. cit.) referred other specimens, that he did not examine, from Mt. St. Helens (90654, 231112 and 231114 BS) to T. t. cooperi although Howell (N. Amer. Fauna, 52:109, November 20, 1929) had previously identified them as E. t. townsendii. By implication, and on his map, Dalquest (op. cit., fig. 83, p. 261) assigned to T. t. cooperi still other specimens, that he had not examined, from: Government Springs, 15 mi. N Carson (230514, 230515, 230559, 230560, and 230563 BS); Stevenson (230513 and 230517 BS); and Skamania (230518 BS). Earlier, Howell (op. cit.) had listed the specimens from the three mentioned localities as Eutamias townsendii townsendii.

Our examination of specimens in the Museum of Vertebrate Zoology from 1-1/2 mi. W Yocolt (94238 and 94239 MVZ) and from 3-1/2 mi. E and 5 mi. N Yocolt (94240-94244 MVZ) reveals that the "average" of the coloration is nearer to that of the paler T. t. cooperi than to that of the darker T. t. townsendii and indicates why Dalquest, we think correctly, identified specimens from Yocolt as T. t. cooperi. We have examined also the specimens in the Biological Surveys Collection of the United States National Museum (catalogue numbers given above) and have compared them with specimens (comparable in age and seasonal condition of pelage) of T. t. townsendii (notably a series from Lake Quinalt, Washington) and of T. t. cooperi (including specimens from Bumping Lake and Blewett Pass, Washington). In color, the specimens from Mt. St. Helens are almost exactly intermediate between T. t. cooperi and T. t. townsendii. We choose to use for them the name T. t. townsendii as did Howell (op. cit.:109). The specimens from 15 mi. N Carson, those from Stevenson and the one from Skamania agree in nearly all features of color with the relatively paler T. t. cooperi, as Dalquest (op. cit.) thought they would, and we, accordingly, use for them the name Tamias townsendii cooperi.

In view of the findings resulting from our study of the above mentioned specimens of the Townsend Chipmunk in Washington, it seemed worthwhile to examine the material of the same species from Hood River, Oregon. Howell (op. cit.:109) listed one specimen from there as E. t. townsendii, but (op. cit.: fig. 7, p. 107) mapped the locality as within the geographic range of E. t. cooperi. The specimen (89061 BS) is a juvenile having external measurements of only 175, 80 and 31. Although the color is intermediate between that of the two subspecies concerned, greater resemblance is shown to T. t. townsendii. We have not examined any other specimen of the species Tamias townsendii so young as No. 89061, but suspect that older specimens from the same place would be paler by a slight degree. This suspicion, and more especially the light color of an older specimen from nearby White Salmon, Washington, and the light color of two older specimens from Parkdale, Oregon, which seem to us to be referable to T. t. cooperi, influence us to refer the specimen from Hood River to Tamias townsendii cooperi Baird.

~Tamias townsendii townsendii~ Bachman

A. H. Howell (N. Amer. Fauna, 52:111, November 30, 1929) referred specimens of the Townsend Chipmunk from the lower elevations on the Olympic Peninsula to Eutamias townsendii townsendii but referred specimens from the central mountains on that peninsula to Eutamias townsendii cooperi. The subspecies T. t. cooperi thus is represented as having a geographic range of two separate parts: (1) The Cascade Mountains from southern British Columbia into southern Oregon, and (2) the area of the Olympic Mountains, the latter area being entirely surrounded by the geographic range of T. t. townsendii. Dalquest (Univ. Kansas Publ. Mus. Nat. Hist., 2:261 and 262, April 9, 1948) employed Howell's arrangement.

We have examined the specimens, in the Biological Surveys Collection of the United States National Museum, from the Olympic Peninsula and fail to find significant differences in external measurements or in size or shape of skulls between specimens from the mountains (alleged T. t. cooperi) and those from other parts of the Peninsula (assigned to T. t. townsendii). Nevertheless, the specimens from the higher parts of the Olympic Mountains resemble T. t. cooperi in being less ochraceous than are specimens of T. t. townsendii from elsewhere on the Olympic Peninsula, and in this one respect, in series, they more closely resemble T. t. cooperi. Even so, the upper parts of the specimens from the mountains are darker than in T. t. cooperi of the Cascades. In dark color of the superciliary stripe the specimens in question are referable to T. t. townsendii. The over-all gray tone, resembling that of T. t. cooperi, upon close inspection is found to be in considerable degree the result of wear, and the difference in grayness from T. t. townsendii, when specimens in comparable pelage are compared, is slight. This tendency to lighter color in specimens from higher elevations is seen in other places in Washington within the geographic range of Tamias townsendii. We feel, therefore, that the mentioned resemblance in color between specimens from the Olympic Mountains and those of T. t. cooperi from the Cascade Mountains is not significant taxonomically. To us, all of the animals of the species Tamias townsendii from the Olympic Peninsula seem best referred to the subspecies Tamias townsendii townsendii Bachman.

~Tamias striatus ohionensis~ Bole and Moulthrop

A. H. Howell (Jour. Mamm., 13:166, May 14, 1932) referred a specimen (252979 USNM) from Athens, Ohio, to Tamias striatus fisheri. Subsequently, Bole and Moulthrop (Sci. Publs. Cleveland Mus. Nat. Hist., 5:83-181, September 11, 1942) named Tamias striatus ohionensis and Tamias striatus rufescens, both of which occur in Ohio. They (op. cit.: 137) also excluded T. s. fisheri from the state list of mammals of Ohio. The locality of Athens lies between the ranges of T. s. ohionensis and T. s. rufescens, as outlined by referred specimens, and thus the identity of the specimen from that place was left in doubt. We have examined the specimen and among named kinds find that it most closely resembles T. s. ohionensis in its less widely spreading zygomata, slender incisors and dull-colored pelage. We prefer the specimen to T. s. ohionensis.

The subspecific identity of specimen No. 174762 USNM, a skin only, from Nobleville, Hamilton Co., Indiana, assigned by Howell (N. Amer. Fauna, 52:21, November 30, 1929) to T. s. griseus and by Lyon (Amer. Mid. Nat., 17(1):191, January, 1936) to T. s. fisheri, was left in doubt by Bole and Moulthrop's (op. cit.) assignment of specimens to T. s. ohionensis. Although the specimen lacks a skull and tail, on the basis of its dull-colored pelage and dark brown (anteriorly) median dorsal stripe, we identify No. 174762 as T. s. ohionensis. For the same reason, specimen No. 125445 USNM, from Bascom, Indiana, referred by Howell (op. cit.:16) to T. s. striatus, and by Lyon (op. cit.:191) to T. s. fisheri, required re-examination. The specimen appears to be an intergrade between T. s. striatus and T. s. ohionensis; it is probably best referred to the latter subspecies which it resembles in having short nasals. In color it is intermediate, but it does not possess the narrowly spreading zygomata of T. s. ohionensis and, in this respect, more nearly approaches T. s. striatus.

Specimen No. 13815 USNM, an alcoholic, from Wheatland, Knox Co., Indiana, was assigned by Howell (op. cit., 1929:21) to T. s. griseus and by Lyon (loc. cit.) to T. s. fisheri. Although the specimen is much faded and cannot be identified with certainty, we assign it to T. s. ohionensis. Allowing for fading, it seems to resemble ohionensis more in the lighter color of the anterior part of the median dorsal stripe, than it does either griseus or fisheri. We are also influenced in making this allocation by Bole and Moulthrop's (op. cit.:137) finding intergradation between T. s. ohionensis and T. s. striatus in a specimen obtained at New Harmony, Posey Co., Indiana.

Howell (Jour. Mamm., 13:166, August 9, 1932) referred two specimens from Boone County, Indiana, to T. s. fisheri. We have examined a specimen (5675 AMNH) from that place and think it is one of the two seen by Howell. The specimen is a poorly made skin in worn winter pelage with the skull inside. Because it differs from T. s. fisheri and agrees with T. s. ohionensis in the color of both upper parts and underparts (comparisons made with material of comparable stage of molt), we assign it to the latter subspecies. Howell (loc. cit.) referred specimens from Overton (57394), Wooster (57398, 57399, and 57442), and Loudonville (57391-57393), all from Ohio, in the Museum of Zoology of the University of Michigan, to Tamias striatus fisheri. We have examined these specimens and find them to be readily separable from T. s. rufescens on the basis of darker coloration. The affinities of the specimens in question are with T. s. fisheri and T. s. ohionensis. As a standard for comparison we have used specimens in the Museum of Zoology, University of Michigan, in comparable pelage of T. s. ohionensis from Dearborn County, Indiana, taken in August and specimens of T. s. fisheri from "near" summit Butt Mtn. and Little Meadows, both places in Giles County, Virginia, as well as two specimens from Allair, Monmouth County, New Jersey. On the basis of buffy (instead of white) edging of the tail, buffy (not white) light dorsal stripes, and buffy (not black) anterior third of the median dark stripe, the specimens from Overton, Wooster, and Loudonville are referred to Tamias striatus ohionensis.

~Tamias striatus pipilans~ Lowery

A. H. Howell (N. Amer. Fauna, 29:16, November 30, 1929) recorded six specimens of Tamias striatus striatus from Greensboro, Alabama. Subsequently, Lowery (Occas. Papers Mus. Zool., Louisiana State Univ., 13:235, November 22, 1943) named T. s. pipilans and assigned to it specimens from northeastern Alabama. Lowery did not, however, mention the specimens from Greensboro and, thus, their subspecific identity was placed in doubt. We have examined five of the six specimens mentioned by Howell (loc. cit.) (57034-57036, 57588, and 77037 BS) and because of their brilliant color and large size, refer them to Tamias striatus pipilans Lowery.

~Tamias striatus rufescens~ Bole and Moulthrop

A. H. Howell (Jour. Mamm., 13:166, August 9, 1932) also referred a specimen (13154), from La Porte, Indiana, in the Chicago Nat. History Museum to T. s. fisheri. We find the specimen to be distinguishable from T. s. fisheri in darker, richer pelage, brown instead of blackish anterior third of the median dorsal stripe, more buffy light dorsal stripes, and more heavily constructed skull. The specimen most closely resembles T. s. rufescens in having, as compared to T. s. ohionensis, brighter, more rufescent color, wider incisors, proportionately narrower interorbital region, and more widely spreading zygomatic arches. We refer it to that subspecies.

~Sciurus carolinensis pennsylvanicus~ Ord

When J. A. Allen considered what name to apply to the gray squirrel of northeastern United States and adjacent parts of Canada, (Monogr. N. Amer. Rodentia, p. 709, 1877) he selected the name leucotis of Gapper (Zool. Jour., 5:206, 1830) as applicable. Allen rejected Ord's (Guthrie's Geog., 2nd Amer. Ed., Zool. App., 2:292, 1815) earlier name, Sciurus Pennsylvanica, because (loc. cit.) "it was given to specimens from the Middle Atlantic States, and hence from a locality bordering upon the habitat of the southern form, and consequently the name is not strictly applicable to the northern type as developed in the Northern and Northeastern States and the Canadas." It must be recalled that Allen had not at that time seen a copy of Ord's exceedingly rare work and was basing his comments on Baird's statements on Ord's treatment of the squirrels.

Subsequently, Rhoads obtained a copy of the second edition of Guthrie's Geography and had Ord's zoological appendix thereto reprinted. The reprinted version (now known generally as Ord's Zoology by Rhoads, 1894) contains (Appendix, p. 19) Rhoads' review of the pennsylvanicus vs. leucotis controversy. Rhoads concluded that pennsylvanicus must apply because it has priority and is available. The habitat was given by Ord as "those parts of Pennsylvania which lie to the westward of the Allegany ridge," not the "Middle Atlantic States" as Allen thought.

Notwithstanding Rhoads' comments, Bangs (Proc. Biol. Soc. Washington, 10:156, December 28, 1896), in his "Review of the Squirrels of Eastern North America," employed leucotis Gapper and rejected Ord's name because it "is a nomen nudum" and of uncertain application. There seems to have been no attempt subsequently to review the pertinent names.

We are of the opinion that Rhoads' (loc. cit.) analysis and conclusions are correct and as cogent today as then. We do not agree with Bangs that pennsylvanicus is a nomen nudum for the following reasons. The name was based on melanistic individuals and could conceivably be applied to three species of squirrels, the red squirrel, the fox squirrel, and the gray squirrel. Melanistic red squirrels, Tamiasciurus hudsonicus, are everywhere rare and in any case appear as individuals and not populations. Ord (loc. cit.) reported that his Sciurus Pennsylvanica was abundant. Ord, we think, was not referring to the fox squirrel, Sciurus niger, because he wrote that S. Pennsylvania "has always been confounded with... [Sciurus niger], but it is a different species," and (loc. cit.) described S. niger as a "Large Black Squirrel" and Sciurus Pennsylvanica as a "Small Black Squirrel." Therefore, pennsylvanicus Ord can refer only to Sciurus carolinensis. Further, melanistic gray squirrels then, as now, were common in western Pennsylvania and exceedingly rare in eastern Pennsylvania. Additionally, Ord described his animal, although admittedly inadequately (small, black, not S. niger). The name Sciurus Pennsylvanica Ord is clearly not a nomen nudum and must replace leucotis Gapper.

Allen's (loc. cit.) argument that the specimens were not representative of "leucotis" because they were from the Middle Atlantic States is based on an initial misunderstanding of the locality. Further, whether or not "topotypes" are representative of a subspecies has no bearing on the availability of the name appended to them. The name and synonomy of the northern gray squirrel are as follows:

~Sciurus carolinensis pennsylvanicus~ Ord