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"; + counter-increment: item; +} + +dd.small_margin { + margin-left:0px; +} + + +/* Fraction Fixes */ +sup, +sub { + height: 0; + line-height: 1; + vertical-align: baseline; + vertical-align: bottom; + position: relative; + +} + +sup { + bottom: 1ex; +} + +sub { + top: .5ex; +} + +table.key { + margin:0px auto; +} + +td.text { + width:700px; + padding-right:10px; +} + +td.text_left { + width:700px; + padding-left:20px; + text-align:justify; +} + +td.number { + width:100px; +} + +td.litcit_num { + width:50px; + text-align:right; + vertical-align:top; + padding-left:15px; +} + +td.num_up { + width:100px; + vertical-align:top; + text-align:right; +} + +td.litcit_txt { + width:750px; + padding-left:10px; +} + + + </style> +</head> + + +<body> + + +<pre> + +Project Gutenberg's The Baculum in Microtine Rodents, by Sydney Anderson + +This eBook is for the use of anyone anywhere 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 + + +Title: The Baculum in Microtine Rodents + +Author: Sydney Anderson + +Release Date: November 15, 2011 [EBook #38021] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK THE BACULUM IN MICROTINE RODENTS *** + + + + +Produced by Chris Curnow, Alex Gam, Joseph Cooper and the +Online Distributed Proofreading Team at http://www.pgdp.net + + + + + + +</pre> + + +<p class="smcap center">University of Kansas Publications<br /> +Museum of Natural History</p> + +<p class="center">Volume 12, No. 3, pp. 181-216, 49 figs.<br /> +February 19, 1960</p> + +<h1>The Baculum in Microtine Rodents</h1> + +<p class="center" style="font-size:larger;">BY<br /> +SYDNEY ANDERSON</p> + +<p class="smcap center" style="margin-top:3em;">University of Kansas<br /> +Lawrence<br /> +1960</p> + +<p class="smcap center" style="margin-top:3em;">University of Kansas Publications, Museum of Natural History</p> + +<p class="center">Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Robert W. Wilson</p> + +<p class="center">Volume 12, No. 3, pp. 181-216, 49 figs.<br /> +Published February 19, 1960</p> + +<p class="center"><span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas</p> + +<p class="center" style="font-size:smaller;">PRINTED IN<br /> +THE STATE PRINTING PLANT<br /> +TOPEKA, KANSAS<br /> +1960</p> + +<div class="imgcenter" style="width: 100px;"> +<img src="images/001.jpg" width="100" height="39" alt="" title="" /> +</div> + +<p class="center">28-774</p> + + +<hr /> + +<p><span class="pagenum"><a name="Page_183" id="Page_183">[Pg 183]</a></span></p> + +<p class="center" style="font-size:2em;">The Baculum in Microtine Rodents</p> +<p class="center">BY<br />SYDNEY ANDERSON</p> + +<h2>INTRODUCTION</h2> + +<p>Didier (1943, 1954) has described the bacula of several Old World +microtines, and other rodents. Argyropulo studied (1933a, 1933b) five +species of Cricetinae and <i>Microtus socialis</i>. Ognev (1950) +illustrated numerous species of Eurasian microtines. Hamilton (1946) +figured and described the baculum of 11 species of North American +microtines. Hibbard and Rinker (1942, 1943) figured the baculum of +<i>Synaptomys cooperi paludis</i> and of <i>Microtus ochrogaster +taylori</i>. Dearden (1958) studied the baculum in two Asiatic species of +<i>Lagurus</i>, in six subspecies of <i>Lagurus curtatus</i> of North +America, and in six other species of microtines of other genera.</p> + +<p>The baculum can be preserved easily with standard study skins, and is +potentially useful in interpreting relationships on any taxonomic level, +and especially in determining the relationships of species within a +genus, if used together with other structures.</p> + +<p>The anatomical orientation of the baculum needs comment because some +confusion exists in the literature, especially concerning the use of the +terms ventral and dorsal. The urethra lies on the anatomically ventral +side of the penis, and of the baculum. In the center of the penis lies a +single corpus cavernosum penis, shown in cross section proximal to the +baculum in Figure 1c. Dorsally an artery, thinner walled than the ventral +urethra, ends in a somewhat reticulate sinus surrounding primarily the +middle part of the baculum within the bulbous glans penis. The corpus +cavernosum penis (the structure has no median septum, at least distally) +terminates with the baculum and is closely knit to it. The site of this +bond is evident in the tuberosities and sculpturing of the base of the +baculum.</p> + +<p>The part of the penis enclosing the baculum, when not erect, is folded +back as shown in Figures 1a and 1b. As a result the anatomically ventral +surface faces upwards, or at least posterodorsally. The use of the term +ventral in this account refers to the anatomically ventral side, that is +to say to the side of the baculum facing the urethra.</p> + +<p>The baculum in microtines consists of an elongate stalk, having a +laterally, and to a lesser extent dorsoventrally, expanded base and + +<span class="pagenum"><a name="Page_184" id="Page_184">[Pg +184]</a></span> + +an attenuate distal shaft. Usually, three digitate processes of +cartilaginous material in which additional ossifications may occur arise +from the terminus of the shaft. The proportions and curvature of the +stalk vary as do the proportions of the terminal ossifications to each +other and to the stalk. In some species one or more of the digital +processes are frequently completely unossified.</p> + +<div class="imgcenter" style="width: 500px;"> +<img src="images/184a.jpg" width="500" height="203" alt="Diagram a, b, c." title="" /> +</div> + +<div class="imgcenter" style="width: 500px;"> +<img src="images/184b.jpg" width="500" height="268" alt="Graph. X-axis: Length of Animal in Millimeters. Y-axis: Length of Stalk of Baculum in Millimeters." title="" /> +</div> + +<p><span class="smcap">Figure 1.</span> The +baculum in <i>Microtus ochrogaster</i>—orientation and variation +with age. <i>a.</i> Diagram of a sagittal section of the posterior half +of a vole, natural size. The penis, containing the baculum (in black), +extends ventrally from a point posterior to the pubic symphysis +(stippled), along the body wall, and bends posteriorly at the distal end. +<i>b.</i> Distal end of penis (× 2) showing baculum (in black), the +urethra (solid lines) adjacent to the baculum, and the corpus cavernosum +(broken lines) proximal to the baculum. <i>c.</i> Oblique view of the +cross section of penis (× 4) shown in Figure 1 <i>b</i>. The +thick-walled urethra lies ventral to the curved corpus cavernosum. A +thinner-walled blood-vessel lies dorsal to the corpus cavernosum. The +anatomically ventral side of the baculum, in the normal non-erect penis +shown, is seen to face dorsally. <i>d.</i> Graph showing the relationship +between size of baculum, size of animal, and development of digital +ossifications. Circles show presence of ossification in stalk only; +circles enclosing dots indicate presence of secondary ossification in +median process also; large dots indicate the addition of tertiary +ossification in one or both of the lateral digitate processes.</p> + +<p><span class="pagenum"><a name="Page_185" id="Page_185">[Pg +185]</a></span></p> + +<p>Preserved specimens of <i>Microtus arvalis</i>, <i>Microtus +agrestis</i>, <i>Microtus orcadensis</i>, <i>Microtus nivalis</i>, +<i>Microtus guentheri</i>, <i>Microtus subterraneus</i>, <i>Clethrionomys +glareolus</i>, and <i>Ellobius lutescens</i> were provided by Prof. +Robert Matthey of Lausanne, Switzerland. J. Knox Jones, Jr. carefully +saved the bacula with specimens of <i>Microtus fortis</i> and +<i>Clethrionomys rufocanus</i> from Korea. Dr. W. B. Quay, Department of +Zoology, University of California, supplied specimens of <i>Synaptomys +cooperi</i>, <i>Phenacomys intermedius</i>, and <i>Microtus oregoni</i>. +Dr. Franklin Sturges and Mr. John W. Goertz, Museum of Natural History, +Oregon State College, Corvallis, have provided specimens including bacula +of <i>Clethrionomys occidentalis</i>, <i>Microtus oregoni</i>, and +<i>Microtus townsendii</i>. Dr. Randolph L. Peterson and Mr. Bristol +Foster, Royal Ontario Museum of Zoology, Toronto, Canada, provided +specimens of <i>Phenacomys intermedius</i>. Dr. J. N. Layne, University +of Florida, Gainsville, Florida, presented me with a baculum of +<i>Microtus parvulus</i>.</p> + +<p>I am indebted to all of these persons for their aid, and to various +collectors for the Museum of Natural History, who preserved bacula with +specimens. Many of these specimens were obtained through the assistance +of the University of Kansas Endowment Association and the National +Science Foundation.</p> + + +<h2>METHODS</h2> + +<p>Bacula were obtained from fresh specimens, specimens preserved in +alcohol or formalin, and dried study skins. The processing of bacula has +been discussed by Hamilton (1946), Friley (1947), White (1951), and +Dearden (1958). The methods used to preserve bacula for my study differed +some from any of those reported. The terminal part of each penis +including the baculum imbedded in the glans penis was removed in its +entirety and placed in a vial. The catalogue number was kept with each +specimen at all times. A two per cent solution of potassium hydroxide was +added. All specimens were examined at least once a day. If tissues other +than the glans penis were present they were removed with forceps when +softened usually at the end of one day. Several drops of Alizarin red-S +stain in a saturated alcoholic solution were added to the 3 to 5 ccs. of +KOH solution in each vial. Solutions were replaced if they became turbid +enough to obstruct observation of the clearing penis. After one day the +solution containing stain was removed and replaced with two per cent KOH +solution without stain. When the glans became sufficiently cleared that +the stained baculum could be seen easily, the solution was replaced by +glycerin in which clearing was completed. The time required for the +entire process varied from one day to more than two weeks depending on +the size of the specimen and on its condition. Fresh specimens clear more +rapidly than dried specimens, and those that are dried more rapidly than +those that are preserved. A three or four per cent solution of hydroxide +will hasten the process, but more frequent observation is required to +prevent excessive maceration.</p> + +<p>Specimens were then examined in a shallow dish containing glycerin +under a binocular microscope. The baculum can be viewed from any desired +direction. The method described above leaves the baculum intact within +the glans penis; therefore its orientation can be determined relative to +the thick walled urethra and the thin walled dorsal artery that extends +onto the dorsal side of the baculum. The ventral curvature of the penis +proximal to the baculum, and the distal extension, characteristic of most +species, of the dorsal border of + +<span class="pagenum"><a name="Page_186" id="Page_186">[Pg +186]</a></span> + +the glans (both shown in Figure 1) are other features aiding in correctly +orienting cleared specimens. The digitate processes are not so often +injured, lost, or displaced when the method described above is used as +they are when the penis is dissected. Specimens were stored in glycerin +in glass shell vials having polyethylene stoppers. A small card bearing +the name, number, locality, and other data was placed in each vial. A +specimen thus enclosed can be kept indefinitely, or removed and mounted +in balsam as described by White (1951:631) or in plastic as described by +Dearden (1958:541) and thus stored in the vial containing the skull of +the specimen.</p> + +<p>Drawings were made on millimeter ruled paper while the baculum was +viewed under a binocular microscope with a square ruled eyepiece.</p> + +<p>Unless otherwise noted all specimens listed are in the University of +Kansas Museum of Natural History. Catalogue numbers are cited. +Measurements are accurate to within less than one-tenth of a millimeter. +Proportions as stated in the text are approximations, accurate to within +one-twelfth (8.33 per cent). The range of variation is unknown for some +species. Mention is made if maturity is known or suspected to differ in +specimens being compared.</p> + +<p>The development of the baculum has been studied by Callery (1951) in +<i>Mesocricetus auratus</i> and by Ruth (1934) in the laboratory rat. In +the rat (<i>Rattus norvegicus</i>) the bone is of endoblastemal origin +being laid down by a condensation of undifferentiated mesenchymal cells. +At the distal end of the bone dense fibrous tissue is then differentiated +and at the proximal end hyaline cartilage. Growth is by substitution at +the proximal end and by subperiosteal lamellation circumferentially. A +marrow cavity is formed by resorption. In the baculum of the hamster the +primary center of ossification is in the stalk, and is present at the age +of three days; the secondary centers are in lateral processes and are +present at 80 days and enlarge subsequently. A tertiary center, in each +median process, may or may not develop later. Maximum development of the +baculum is reached late in the reproductive life of the hamster.</p> + +<p>The early ossification of the baculum noted in the rat and the hamster +occurs in <i>Microtus</i> also. A specimen of <i>Microtus montanus +fusus</i> (76831, from 5 mi. N, 26 mi. W Saguache, 9600 ft., Saguache +County, Colorado) only 74 mm. in total length and weighing only 6.6 +grams, had a slender ossified baculum having enlarged ends. This vole was +one-half of the average length and less than one-fifth of the average +weight of an adult, and of approximately the size at which weaning takes +place.</p> + +<p>The development of the baculum in <i>Microtus ochrogaster</i> was +studied in 32 specimens of various ages. The specimens (between Nos. +74994 and 75074) were collected between August 15 and September 4, 1957, +at localities on the Great Plains. These specimens were from breeding +populations, as evidenced by pregnancy of females and by large size of +testes of males. The length and width of the stalk of the baculum, the +presence of digital ossifications, the + +<span class="pagenum"><a name="Page_187" id="Page_187">[Pg +187]</a></span> + +total length of the animal, and the size of the testes were noted. +Variability in length of testes is greatest when voles are from 140 to +150 mm. in total length. Sexual maturity is reached rather abruptly when +the total length of most individuals is 140 to 150 millimeters. If the +baculum likewise underwent more rapid growth at the onset of sexual +maturity, greater variability should be evident in the length of the +baculum of voles 140 to 150 mm. in total length than in bacula of voles +of other sizes. This was the case (see Figure 1d). The baculum does not, +however, suddenly reach its maximum maturity.</p> + +<p>The primary ossification is in the stalk. The secondary ossification +is in the median process except in <i>Lagurus</i> (Dearden, 1958:551) and +some individuals of <i>Neofiber</i> (see account on page 258). Tertiary +centers of ossification are in the lateral processes. The primary +ossification is present at an early age and subsequently increases in +size and solidity. The secondary and tertiary ossifications are +progressively more common in older voles. The increase in degree of +ossification of all parts continues after sexual maturity is reached. +Individual variation and variation with age in the baculum of <i>Microtus +pennsylvanicus</i> have been illustrated by Hamilton (1946:380). Figures +14, 15, and 17 illustrate variation with size, which is correlated with +age, and also illustrate individual variation. The three bacula are from +adult voles having testes that measured 15, 16 and 16 mm. in length, +respectively. Each vole was trapped in late June. The total lengths in +millimeters of the three voles are 172, 167, and 181; weights are 55, +52.4, and 65.5 grams. I judge that the greater size of the stalk and the +better developed base shown in Figure 17 than in Figure 15 are +illustrative of age variation; the difference in the size of the lateral +digitate processes is, in this case, attributable to individual +variation. Differences in the distal end of the baculum in Figures 42 and +43, show individual variation also. Figures 35 and 36 represent two +different subspecies; different individuals of <i>M. mexicanus +mogollonensis</i>, however, exhibit individual variation of the same +degree.</p> + +<p>Hall and Cockrum (1953) list 44 species of microtines in North +America. At least twelve of these are insular or local forms perhaps +derived from some other species; for example <i>Microtus coronarius</i>, +an insular form derived from <i>Microtus longicaudus</i>; <i>Microtus +provectus</i>, considered by Chamberlain (1954:587) and by Wheeler +(1956:176) as a subspecies of <i>Microtus pennsylvanicus</i>; and +<i>Microtus ludovicianus</i>, a close relative of <i>Microtus +ochrogaster</i>.</p> + +<p>All North American genera have been studied. Of the genus +<i>Microtus</i> in North America, all subgenera but <i>Orthriomys</i> and +all species but the following nine, have been studied: <i>M. (Orthriomys) +umbrosus</i>, the insular <i>M. (Stenocranius) abbreviatus</i>, <i>M. +(Microtus) breweri</i>, <i>M. (Microtus) nesophilus</i>, <i>M.</i></p> + +<p><span class="pagenum"><a name="Page_188" id="Page_188">[Pg +188]</a></span></p> + +<div class="imgcenter" style="width: 421px;"> +<img src="images/188.jpg" width="421" height="500" alt="" title="" /> +</div> + +<p><span class="smcap">Figures</span> 2-13. Bacula of microtines. Unless +indicated otherwise views are (<i>a</i>) of the dorsum, (<i>b</i>) the +right side, and (<i>c</i>) the proximal end with the dorsal surface +upward. Exact localities are given in accounts of species concerned.</p> + +<ol id="start_02"> + <li><i>Lemmus trimucronatus</i>, 50678, Point Barrow, Alaska.</li> + <li><i>Dicrostonyx groenlandicus</i>, 50539, Porcupine Lake, Brooks Range, + Alaska.</li> + <li><i>Dicrostonyx groenlandicus</i>, 52524, Point Barrow, Alaska.</li> + <li><i>Synaptomys cooperi saturatus</i>, WBQ 3-C-454, 3 mi. S Demotte, Indiana.</li> + <li><i>Synaptomys cooperi paludis</i>, 13716, Meade County State Park, Kansas.</li> + <li><i>Phenacomys intermedius celatus</i>, SA 2044, Quebec.</li> + <li><i>Phenacomys intermedius intermedius</i>, WBQ 3-C-309, 5.4 mi. S Moran, + Teton Co., Wyoming.</li> + <li><i>Clethrionomys rufocanus</i>, 60438, 1 mi. NW Oho-ri, Korea, (<i>d</i>) ventral + view.</li> + <li><i>Clethrionomys gapperi</i>, 42108, 31 mi. N Pinedale, Wyoming.</li> + <li><i>Clethrionomys rutilus</i>, 42865, 5 mi. NNE Gulkana, Alaska.</li> + <li><i>Clethrionomys occidentalis</i>, FWS 30, Mary's Peak, Benton Co., Oregon.</li> + <li><i>Clethrionomys glareolus</i>, 67100, Zermatt, Valais, Switzerland.</li> +</ol> + +<p><span class="pagenum"><a name="Page_189" id="Page_189">[Pg +189]</a></span></p> + +<div class="imgcenter" style="width: 431px;"> +<img src="images/189.jpg" width="431" height="500" alt="" title="" /> +</div> + +<p><span class="smcap">Figures</span> 14-25. Bacula of <i>Microtus</i>. +Unless indicated otherwise views are (<i>a</i>) of the dorsum, (<i>b</i>) +the right side, and (<i>c</i>) the proximal end with dorsal surface +upward.</p> + +<ol id="start_14"> + <li><i>M. pennsylvanicus</i>, 42439, 1 mi. S, 2 mi. E Eagle Nest, Colfax +Co., New Mexico; abnormality perhaps owing to injury; dorsal view.</li> + <li><i>M. pennsylvanicus</i>, 42306, 5 mi. N, 26 mi. W Saguache, Colorado; + dorsal view.</li> + <li><i>M. pennsylvanicus</i>, 43043, 20 mi. NE Anchorage, Alaska, ventral view.</li> + <li><i>M. pennsylvanicus</i>, 42430, 1 mi. S, 2 mi. E Eagle Nest, New Mexico.</li> + <li><i>M. agrestis</i>, 67102, Gryon, Switzerland.</li> + <li><i>M. montanus amosus</i>, 62241, 1/2 mi. E Soldier Summit, Wasatch Co., + Utah.</li> + <li><i>M. montanus nanus</i>, 57470, 2 mi. N, 2 mi. W Pocatello, Idaho.</li> + <li><i>M. montanus fusus</i>, 42307, 5 mi. N, 26 mi. W Saguache, Colorado.</li> + <li><i>M. arvalis</i>, 67101, Vidy, Switzerland, possibly not mature.</li> + <li><i>M. guentheri</i>, 67104, Palestine.</li> + <li><i>M. orcadensis</i>, 67106, Orkney Islands, orientation uncertain.</li> + <li><i>M. fortis</i>, 63841, Chipo-ri, Korea, (<i>d</i>) ventral view.</li> +</ol> + +<p><span class="pagenum"><a name="Page_190" id="Page_190">[Pg +190]</a></span></p> + +<div class="imgcenter" style="width: 432px;"> +<img src="images/190.jpg" width="432" height="500" alt="" title="" /> +</div> + +<p><span class="smcap">Figures</span> 26-39. Bacula of microtines. Unless indicated +otherwise views are (<i>a</i>) of the dorsum, (<i>b</i>) the right side, +and (<i>c</i>) the proximal end with the dorsal surface upward.</p> + +<ol id="start_25"> + <li><i>M. (Pitymys) fatioi</i>, 67103, Zermatt, Switzerland, immature.</li> + <li><i>M. (Pitymys) pinetorum</i>, 76834, 2 mi. N Baldwin, Douglas Co., Kansas.</li> + <li><i>M. (Pitymys) pinetorum</i>, 68545, 1 mi. NE Pleasant Grove, Kansas.</li> + <li><i>M. (Pitymys) quasiator</i>, 30709, Teocelo, Veracruz, (<i>d</i>) ventral view.</li> + <li><i>M. (Pitymys) quasiator</i>, 19878, 5 km. N Jalapa, Veracruz.</li> + <li><i>M. (Pedomys) ochrogaster</i>, 75036, 1 mi. N, 2 mi. E Oberlin, Kansas.</li> + <li><i>M. (Stenocranius) miurus</i>, 51152, Lake Schrader, Brooks Range, Alaska.</li> + <li><i>M. (Stenocranius) miurus</i>, 51169, Lake Schrader, Brooks Range, Alaska.</li> + <li><i>M. (Stenocranius) gregalis</i>, 8059, "Eastern Europe."</li> + <li><i>M. mexicanus mexicanus</i>, 63094, Valle de Bravo, Estado de México, México.</li> + <li><i>M. mexicanus mogollonensis</i>, 63298, Mt. Taylor, Valencia Co., New Mexico.</li> + <li><i>M. californicus</i>, 76828, 1 mi. NE Berkeley, California; (<i>d</i>) ventral view.</li> + <li><i>M. (Arvicola) richardsoni</i>, 42454, 31 mi. N Pinedale, Sublette Co., Wyoming.</li> + <li><i>M. richardsoni</i>, 37903, 23-1/2 mi. S, 5 mi. W Lander, Wyoming; distal end.</li> +</ol> + +<p><span class="pagenum"><a name="Page_191" id="Page_191">[Pg +191]</a></span></p> + +<div class="imgcenter" style="width: 420px;"> +<img src="images/191.jpg" width="420" height="500" alt="" title="" /> +</div> + +<p><span class="smcap">Figures</span> 40-49. Bacula of microtines. Unless +indicated otherwise views are (<i>a</i>) of the dorsum, (<i>b</i>) the +right side, and (<i>c</i>) the proximal end with the dorsal surface +upward.</p> + +<ol id="start_39"> + <li><i>Microtus (Pitymys) parvulus</i>, UF 1508, 1 mi. W Micanopy, Florida.</li> + <li><i>Microtus townsendii</i>, 79186, Sec. 33, T. 11S, R. 5W, Benton Co., Oregon.</li> + <li><i>Microtus (Herpetomys) guatemalensis</i>, 65895, 2 mi. S San Juan Ixcoy, Guatemala.</li> + <li><i>M. guatemalensis</i>, 65921, 10 mi. E, 4 mi. S Totonicapan, Guatemala, dorsal view + of tip.</li> + <li><i>Microtus oeconomus</i>, 43048, Kelsall Lake, British Columbia.</li> + <li><i>Microtus (Chilotus) oregoni</i>, WBQ 3-C-248, 5 mi. N Orick, California.</li> + <li><i>Lagurus (Lemmiscus) curtatus</i>, 26053, 9 mi. S Robertson, Uinta Co., Wyoming.</li> + <li><i>Microtus (Chionomys) nivalis</i>, 65127, Wetterstein, Germany, orientation uncertain.</li> + <li><i>Microtus (Chionomys) longicaudus</i>, 50253, Crane Flat, Mariposa Co., California.</li> + <li><i>Neofiber alleni</i>, 27268, 2 mi. S Gainesville, Florida, orientation uncertain.</li> +</ol> + +<p><span class="pagenum"><a name="Page_192" id="Page_192">[Pg +192]</a></span></p> + +<p style="margin-bottom:2em;"><i>(Microtus) provectus</i> (the last three are probably insular +derivatives of <i>M. pennsylvanicus</i>), <i>M. (Microtus) +fulviventer</i> (perhaps derived from the same stock as <i>Microtus +mexicanus</i>), <i>M. (Microtus) xanthognathus</i> (perhaps related to +<i>Microtus chrotorrhinus</i>), <i>M. (Microtus) coronarius</i>, and +<i>M. (Pedomys) ludovicianus</i>.</p> + +<table> + <tr> + <th class="smcap" colspan="2">Species of Which Bacula Were Examined</th> + </tr> + <tr> + <td class="left">Subfamily: Microtinae</td> + <td class="right">Number of Specimens</td> + </tr> + <tr> + <td class="bump1" colspan="2">Tribe: Lemmi</td> + </tr> + <tr> + <td class="bump2"><i>Dicrostonyx groenlandicus</i> (Traill)</td> + <td class="center">4</td> + </tr> + <tr> + <td class="bump2"><i>Lemmus trimucronatus</i> (Richardson)</td> + <td class="center">6</td> + </tr> + <tr> + <td class="bump2"><i>Synaptomys cooperi</i> Baird</td> + <td class="center">5</td> + </tr> + <tr> + <td class="bump1" colspan="2">Tribe: Microti</td> + </tr> + <tr> + <td class="bump2" colspan="2">Genus: <i>Clethrionomys</i> Tilesius, 1850</td> + </tr> + <tr> + <td class="bump3"><i>Clethrionomys rutilus</i> Pallas</td> + <td class="center">4</td> + </tr> + <tr> + <td class="bump3"><i>Clethrionomys gapperi</i> (Vigors)</td> + <td class="center">9</td> + </tr> + <tr> + <td class="bump3"><i>Clethrionomys occidentalis</i> (Merriam)</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump3"><i>Clethrionomys glareolus</i> Schreber</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump3"><i>Clethrionomys rufocanus</i> Sundevall</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump2" colspan="2">Genus: <i>Phenacomys</i> Merriam, 1897</td> + </tr> + <tr> + <td class="bump3"><i>Phenacomys intermedius</i> Merriam</td> + <td class="center">5</td> + </tr> + <tr> + <td class="bump2" colspan="2">Genus: <i>Ondatra</i> Link, 1795</td> + </tr> + <tr> + <td class="bump3"><i>Ondatra zibethicus</i> (Linnaeus)</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump2" colspan="2">Genus: <i>Microtus</i> Schrank, 1798</td> + </tr> + <tr> + <td class="bump3">(<i>Herpetomys</i>) <i>guatemalensis</i> Merriam</td> + <td class="center">3</td> + </tr> + <tr> + <td class="bump3">(<i>Arvicola</i>) <i>richardsoni</i> (DeKay)</td> + <td class="center">2</td> + </tr> + <tr> + <td class="bump3">(<i>Chilotus</i>) <i>oregoni</i> (Bachman)</td> + <td class="center">3</td> + </tr> + <tr> + <td class="bump3">(<i>Stenocranius</i>) <i>gregalis</i> (Pallas)</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump3">(<i>Stenocranius</i>) <i>miurus</i> Osgood</td> + <td class="center">9</td> + </tr> + <tr> + <td class="bump3">(<i>Chionomys</i>) <i>longicaudus</i> (Merriam)</td> + <td class="center">6</td> + </tr> + <tr> + <td class="bump3">(<i>Chionomys</i>) <i>nivalis</i> Martins</td> + <td class="center">2</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>arvalis</i> (Pallas)</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>orcadensis</i> Millais</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>guentheri</i> Danford and Alston</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>fortis</i> Büchner</td> + <td class="center">2</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>montanus</i> (Peale)</td> + <td class="center">15</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>townsendii</i> (Bachman)</td> + <td class="center">3</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>oeconomus</i> (Pallas)</td> + <td class="center">10</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>mexicanus</i> (Saussure)</td> + <td class="center">13</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>californicus</i> (Peale)</td> + <td class="center">2</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>pennsylvanicus</i> (Ord)</td> + <td class="center">13</td> + </tr> + <tr> + <td class="bump3">(<i>Microtus</i>) <i>agrestis</i> (Linnaeus)</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump3">(<i>Pedomys</i>) <i>ochrogaster</i> (Wagner)</td> + <td class="center">41</td> + </tr> + <tr> + <td class="bump3">(<i>Pitymys</i>) <i>pinetorum</i> (LeConte)</td> + <td class="center">2</td> + </tr> + <tr> + <td class="bump3">(<i>Pitymys</i>) <i>parvulus</i> (Howell)</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump3">(<i>Pitymys</i>) <i>quasiater</i> (Coues)</td> + <td class="center">5</td> + </tr> + <tr> + <td class="bump3">(<i>Pitymys</i>) <i>fatioi</i> Mottaz</td> + <td class="center">1</td> + </tr> + <tr> + <td class="bump2" colspan="2">Genus: <i>Neofiber</i> True, 1884</td> + </tr> + <tr> + <td class="bump3"><i>Neofiber alleni</i> True</td> + <td class="center">2</td> + </tr> + <tr> + <td class="bump2" colspan="2">Genus: <i>Lagurus</i> Gloger, 1841</td> + </tr> + <tr> + <td class="bump3"><i>Lagurus curtatus</i> (Cope)</td> + <td class="center">7</td> + </tr> + <tr> + <td class="bump4">Total number examined</td> + <td class="center">184</td> + </tr> +</table> + + +<p><span class="pagenum"><a name="Page_193" id="Page_193">[Pg +193]</a></span></p> + +<h2>ACCOUNTS OF SPECIES</h2> + +<h3>Dicrostonyx groenlandicus (Traill)</h3> +<p class="center">Figs. 3 and 4</p> + +<p>Baculum: stalk elongate, greatest length (3.1 mm.) 2 <sup>1</sup>/<sub>5</sub> to 2½ +times greatest breadth, and 4½ times greatest depth; digitate +processes usually cartilaginous, occasionally lateral processes partly +ossified; basal tuberosities weakly to moderately developed, medially +confluent; posterior profile in dorsal view rounded with rounded +posterior apex or shallow notch; dorsal concavity in end-view shallower +and not so wide as ventral concavity; median constriction approximately +<sup>2</sup>/<sub>3</sub> greatest depth; ventral part of base in end-view wider than dorsal +part; shaft straight or slightly curved; base of stalk placed dorsally +relative to axis of shaft; stalk spatulate, sometimes with distal +enlargement; at mid-point stalk wider than high; lateral profile in +dorsal view sloping gradually without abrupt curvature anterior to point +of greatest width.</p> + +<p>The baculum of <i>Dicrostonyx torquatus</i> figured by Ognev +(1948:476) agrees with that of <i>D. groenlandicus</i> in shape of stalk, +and in lateral digitate processes that are small relative to size of +median process; but differs in more elongate, terminally enlarged, bulbar +shape of median process. None of my specimens showed ossification in the +lateral processes, observed by Hamilton (1946:381) in <i>Dicrostonyx +rubricatus richardsoni</i> [ = <i>D. groenlandicus richardsoni</i>]. In +all of my specimens the cartilaginous median process was larger than that +figured by Hamilton, or by Dearden (1958:542).</p> + +<p><i>Specimens examined</i>: Four from; Point Barrow, Alaska, 52524 +(Barrow Village), 67264 (died in captivity); Brooks Range, Alaska, 50536 +(Wahoo Lake, 69°08', 146°58'), 50539 (Porcupine Lake, 68°51'57", +146°29'50", 3140 ft.).</p> + + +<h3>Lemmus trimucronatus (Richardson)</h3> +<p class="center">Fig. 2</p> + +<p>Baculum: Stalk heavy, broad, greatest length (2.8 mm.) in mature +individuals (Fig. 2) as little as 1<sup>1</sup>/<sub>3</sub> times +greatest breadth, greatest length no less than 2<sup>2</sup>/<sub>3</sub> times greatest depth +of base; three ossified processes, median one from as long as to ½ +longer than the lateral processes, and approximately +<sup>2</sup>/<sub>3</sub> wider and twice as deep as lateral processes; +length of median process almost 3½ times its breadth, +approximately ½ length of stalk; basal fossae broadly confluent; +posterior profile in dorsal view evenly rounded; in end-view ventral +concavity deeper than dorsal concavity, constriction as little as +½ greatest depth in mature specimens; shaft straight, bluntly +rounded, or slightly decurved and laterally inflated terminally; lateral +profile in dorsal view a gradual slope from widest point of stalk +anteriorly onto shaft; in younger individuals stalk slenderer, otherwise +as described above.</p> + +<p>Five specimens examined by me differ from one figured and described by +Hamilton (1946:379) in that stalk is better developed, larger relative to +size of processes, length of stalk in my specimen (Fig. 2) 2.8 as opposed +to 2.1 mm. in Hamilton's specimen; median process shorter, 1.5 as opposed +to 1.8 mm., proximal end rounded rather than concave, not partially +enclosing tip of shaft; proportion of and relative sizes of median and +lateral processes approximately same as in Hamilton's <i>Lemmus +helvolus</i> [= <i>Lemmus trimucronatus helvolus</i>]. + +<span class="pagenum"><a name="Page_194" id="Page_194">[Pg +194]</a></span> + +A specimen figured by Dearden (1958:542) has a basally trilobed median +process.</p> + +<p>The baculum of the Asiatic <i>Lemmus lemmus</i> figured by Ognev +(1948:413) agrees with my specimens in the ossification of three +processes, the relative sizes of these processes to each other and to the +stalk, the well-developed base of the stalk and heavy bluntly rounded +shaft; the baculum of <i>Lemmus lemmus</i> differs in greater +anterolateral extent of basal tuberosities, in proximal notch seemingly +separating these tuberosities, and in median process being slenderer.</p> + +<p><i>Specimens examined</i>: Five, of two subspecies; <i>Lemmus +trimucronatus alascensis</i>, Point Barrow, Alaska, numbers 50591, 50678, +50731, 50758; <i>Lemmus trimucronatus subarcticus</i>, Wahoo Lake, +69°08', 146°58', 2350 ft., Brooks Range, Alaska, 50948.</p> + + +<h3>Synaptomys cooperi Baird</h3> +<p class="center">Figs. 5 and 6</p> + +<p>Baculum: Stalk elongate, greatest length (2.7 to 2.8 mm.) 2 +<sup>1</sup>/<sub>3</sub> to 2½ times greatest breadth, 4 to 5 +times greatest depth; three processes ossified or lateral processes +unossified, ossifications relatively small (in 78380, median ossification +less than ¼ as large as lateral ossifications although median +cartilaginous process is larger), length of median process +<sup>1</sup>/<sub>5</sub> to <sup>1</sup>/<sub>6</sub> of length of +stalk, cartilaginous part of median process larger; posterior profile in +dorsal view convex throughout or bilobate; tuberosities moderately +developed, deflected dorsal to axis of shaft; in end-view medial +construction <sup>3</sup>/<sub>5</sub> greatest depth of tuberosities; +shaft tapered from point of greatest width, slightly inflated +terminally.</p> + +<p>The specimen (KU 13716) figured by Hibbard and Rinker (1942:29) has +been restudied. It was first cleared and stained to soften the dry +cartilage binding the digital processes together and to differentiate +bone and cartilage. The lateral processes are small and cartilaginous +(Fig. 6) and seem intact. The differences between this specimen and +others examined by Hamilton (1946:381), Dearden (1958:542), and myself, +namely the relatively larger median ossification, the absence of +ossification in lateral processes, and the distinctly bilobate base and +larger size, may represent geographic differences, or individual +variation. The proportions of length, width, and depth of the stalk, and +the appearance in lateral view do not differ greatly from others examined +by Hamilton, by Dearden (1958:546), and by me.</p> + +<p><i>Specimens examined</i>: Five, representing four subspecies; <i>S. +cooperi gossii</i>, 6 mi. N Midway, Holt Co., Nebraska 78379, 78380; +<i>S. cooperi relictus</i>, 5 mi. N, 2 mi. W Parks, Dundy Co., Nebraska, +72601 (immature); <i>S. cooperi saturatus</i>, 3 mi. S Demotte, Jasper +Co., Indiana, 3-C-454, collection of W. B. Quay; <i>S. cooperi +paludis</i>, Meade County State Park, Kansas, 13716.</p> + + +<h3>Clethrionomys rutilus Pallas</h3> +<p class="center">Fig. 11</p> + +<p>Baculum: Stalk elongate, and proximally enlarged, greatest length (2.7 +mm.) 2 times greatest breadth; less than 4 times greatest depth; three +well-developed ossified processes; length of stalk 2<sup>1</sup>/<sub>3</sub> times length of median process; median process +with basal (and ventral) protuberence and lateral lobes, arched in +dorsoventral plane; lateral processes as large as median process, +flattened distally, having ventromedial vane on distal half; basal +tuberosities of stalk well developed, medially confluent; posterior +profile in dorsal view trilobate or convex + +<span class="pagenum"><a name="Page_195" id="Page_195">[Pg +195]</a></span> + +throughout with rounded posterior apex; dorsal concavity well developed, +ventral surface but slightly concave, medial constriction of base as +little as ½ greatest depth; shaft straight, slender, at mid-point +of stalk but slightly wider than high; basal tuberosities largely dorsal +to axis of shaft in lateral view; lateral profile in dorsal view with an +abrupt curvature separating the gently sloping sides of the shaft from +the basal part at its greatest breadth.</p> + +<p>The specimen of <i>Clethrionomys rutilus</i> figured by Ognev +(1950:120) is essentially like the North American specimens examined by +me in the relative sizes of the ossifications and the general shape of +the stalk.</p> + +<p><i>Specimens examined</i>: Four, of one subspecies; <i>C. r. +dawsoni</i>, west bank Gakona River, 1700 ft., 5 mi. NNE Gulkana, Alaska, +42865, 42866; SW end Dezadeash Lake, 2400 ft., Yukon Territory, 42910, +42921.</p> + + +<h3>Clethrionomys gapperi (Vigors)</h3> +<p class="center">Fig. 10</p> + +<p>Baculum: Stalk elongate, greatest length (2.8 mm.) 1¾ times +greatest breadth, and 3¾ times greatest depth; proximally +enlarged, greatest depth ½ greatest breadth; three well-developed +ossified processes; length of stalk 2 <sup>1</sup>/<sub>3</sub> times +length of median process; median process arched in dorsoventral plane, +with basiventral protuberence or spine and lateral lobes; lateral +processes as large as median process, flattened distally, arched; basal +tuberosities of stalk well developed, medially confluent; posterior +profile in dorsal view trilobate or convex throughout with a rounded +posterior apex; dorsal concavity well developed, ventral surface but +slightly concave, or in some cases slightly convex; medial constriction +of base <sup>3</sup>/<sub>5</sub> greatest depth; shaft straight, +slender, at mid-point of stalk twice as wide as high; basal tuberosities +dorsally placed relative to axis of shaft; lateral profile in dorsal view +abruptly curved anterior to point of greatest width; slender stalk +distinct from angular enlarged base.</p> + +<p>The most noticeable difference between the baculum of <i>C. +rutilus</i> and <i>C. gapperi</i> is size. The proportions of the four +ossifications are approximately the same. Ventral vanes on the lateral +processes are not developed in <i>C. gapperi</i>. <i>C. gapperi</i> and +<i>C. rutilus</i> are more nearly alike in their bacula than any other +two species of <i>Clethrionomys</i> examined. <i>Clethrionomys +occidentalis</i>, the other New World species, is also much like <i>C. +gapperi</i> and <i>C. rutilus</i>. The differences are of a magnitude +comparable to those between the bacula in subspecies of <i>Microtus +montanus</i> (Figs. 19-21) for example, or in subspecies of <i>Lagurus +curtatus</i> (Dearden, 1958:542).</p> + +<p><i>Specimens examined</i>: Nine, of two subspecies; <i>Clethrionomys +gapperi athabascae</i>, British Columbia, 42922 (Indian Creek, Mile Post +234 of Alaskan Highway), 64281 (West bank Racing River, 89 mi. W Muskwa), +64287 (North bank Tetsa River, 56 mi. W, 11 mi. S Muskwa), 64290 (44 mi. +W, 9 mi. S Muskwa), 64310 (32 mi. W, 2 mi. S Muskwa); <i>Clethrionomys +gapperi galei</i>, 31 mi. N Pinedale, Sublette Co., Wyoming, 42108; Grand +Mesa, Delta Co., Colorado, 60014 and 60015 (5½ mi. E, 12 mi. S +Collbran), 60022 (8 mi. E, 1/2 mi. S Skyway).</p> + + +<h3>Clethrionomys occidentalis (Merriam)</h3> +<p class="center">Fig. 12</p> + +<p>Baculum: Stalk elongate, greatest length (2.8 mm.) 2½ times +greatest breadth, 6 times greatest depth; three well-developed ossified +processes; median process larger than lateral processes, ½ the +length of stalk, curved, basally + +<span class="pagenum"><a name="Page_196" id="Page_196">[Pg +196]</a></span> + +broad, ventrally keeled, trilobate posteriorly; lateral ossifications +large, flattened distally, curved; posterior profile of stalk posteriorly +slightly emarginate, thus bilobate in outline; in end-view dorsal +concavity deeper than ventral, constriction less than ½ greatest +depth, tuberosities confluent, visible in dorsal view at each side; shaft +slender, especially in depth, straight; at mid-point of stalk almost +twice as wide as deep, slight terminal inflation.</p> + +<p>The general proportions of the stalk and the relatively large, +uniquely shaped processes, are characteristic of most specimens of the +genus <i>Clethrionomys</i> examined.</p> + +<p><i>Specimen examined</i>: <i>C. occidentalis californicus</i>, one +from Mary's Peak, Benton Co., Oregon, 30, F. W. Sturges' collection.</p> + + +<h3>Clethrionomys glareolus Schreber</h3> +<p class="center">Fig. 13</p> + +<p>Baculum: Stalk elongate, greatest length (2.9 mm.) twice the greatest +breadth in the specimen examined, flattened proximally, greatest length +almost 6 times greatest depth of base; three well-developed ossified +processes; median process arched in a dorsoventral plane, with basal +notch and lateral lobes; lateral processes as long as median process, +bowed in dorsal view, flattened distally, with ventromedial vane; basal +tuberosities of stalk weakly developed, medially confluent; posterior +profile in dorsal view evenly rounded; in end-view dorsal concavity +shallow in comparison to most species but deeper than ventral concavity, +constriction ¾ greatest depth; shaft straight, at mid-point +slightly wider than high, elongate, widest point of stalk less than +¼ of total length from proximal end, slight lateral inflation at +tip; lateral profile in dorsal view sloping at first abruptly and then +gradually from widest point of stalk anteriorly onto shaft.</p> + +<p>The specimen of <i>Clethrionomys glareolus</i> figured by Ognev +(1950:31) in dorsal view as I interpret it, resembles my specimen in the +rounded base; in the elongate, distally inflated shaft; in the initially +abrupt slope of the lateral profile in dorsal view from the greatest +width of stalk anteriorly; and in the presence of three well ossified +processes. Ognev's specimen differs from mine in the median process being +more elongate relative to its width, and rounded proximally, lacking +lateral lobes and basal notch; in lateral processes being less curved; in +the greater terminal inflation of the shaft; and in the closer +approximation of the terminal processes to the shaft. The baculum of +<i>Clethrionomys glareolus</i> as described and figured by Didier +(1954:243-244) resembles my specimen in general proportions, but is more +pointed proximally and more curved in dorsoventral plane. Didier states +that the baculum is rather variable in form in this species, in different +regions, but that a large number of specimens must be examined to assess +the geographic nature of this variation.</p> + +<p><i>Specimen examined</i>: One from Zermatt, Valais, Switzerland, +67100.</p> + + +<h3>Clethrionomys rufocanus Sundevall</h3> +<p class="center">Fig. 9</p> + +<p>Baculum: Base of stalk broad but relatively flattened dorsoventrally, +greatest length (3.2 mm.) less than 1½ greatest width, 4 times +greatest depth; three well-developed ossified processes; median process +arched in dorsoventral plane, having basal notch and lateral lobes; +lateral processes as long as median process, + +<span class="pagenum"><a name="Page_197" id="Page_197">[Pg +197]</a></span> + +flattened distally, with ventromedial vane; basal tuberosities of stalk +weakly developed, medially confluent; posterior profile in dorsal view +convex with rounded posterior apex; dorsal surface of base almost flat, +ventral concavity broad and shallow; constriction ¾ greatest depth +(not including an unusual irregularity on the ventral surface of the +base); shaft straight, at mid-point of stalk distinctly wider than high, +slender at distal end, widest point of stalk almost +<sup>1</sup>/<sub>3</sub> of total length from proximal end, tip of shaft +rounded; lateral profile in dorsal view gradually sloping from widest +point anteriorly onto shaft.</p> + +<p>The specimen of <i>Clethrionomys rufocanus</i> figured by Ognev +(1950:97) resembles my specimen in the presence of three well ossified +processes. Ognev's specimen differs however in the lack of a proximal +notch on the median process, the lesser proportion of the stalk included +in the basal enlargement, the more posterior position of the point of +greatest width, and the presence of a concavity in the posterior profile +of the stalk in dorsal view. These differences in the stalk may be owing +to a difference in age (my specimen perhaps being older).</p> + +<p><i>Specimen examined</i>: One from 1 mi. NW Oho-ri, 6 M., Korea, +60438.</p> + + +<h3>Phenacomys intermedius Merriam</h3> +<p class="center">Figs. 7 and 8</p> + +<p>Baculum: Stalk slender, greatest length (2.9 mm.) 2¼ to +2½ times greatest breadth, 4 times greatest depth; three +well-developed ossified processes, median one almost ½ length of +stalk, curved, broad basally and slightly larger in all dimensions than +either lateral process; lateral processes flattened distally, curved; +base of stalk well developed, basal tuberosities medially confluent or +separated by medial emargination, posterolateral faces flattened or +rough; emarginations in the four adults examined; posterior profile in +dorsal view bluntly pointed or flattened except for emargination +posterially, abruptly curved at point of greatest width; shaft arising +broadly from distal side of base of stalk; in end-view hour-glass shaped, +medial constriction pronounced, both dorsal and ventral concavities deep; +shaft having relatively straight but distally convergent sides; at +mid-point of stalk, 1 to 1½ times as wide as deep; tip bluntly +rounded, or slightly inflated.</p> + +<p>The specimens from Quebec differ from the one from Wyoming in smaller +size, relatively smaller lateral digital processes, larger more medial +basal emargination, and slender shafts. The baculum of <i>Phenacomys +intermedius</i> differs much from that of <i>Phenacomys longicaudus</i>, +described by Hamilton (1946:381) and by Dearden (1958:547). Dearden +states that the three bacula examined by him of <i>Phenacomys +longicaudus</i> differ markedly from the specimen described by Hamilton. +It seems to me that in major features the resemblance is greater between +the specimens of <i>Phenacomys longicaudus</i> examined by these two +authors than between their specimens and specimens of other microtines, +including <i>Phenacomys intermedius</i>. Neither Hamilton nor Dearden +record the exact localities of capture, the collections in which the +specimens are deposited, or the catalogue numbers of specimens. +Consequently verification of identifications and observations is +difficult.</p> + +<p><i>Specimens examined</i>: Five, of two subspecies; <i>P. intermedius +intermedius</i>, 5.4 mi. S Moran, Teton Co., Wyoming, 3-C-309, collection +of W. B. Quay; <i>P. intermedius celatus</i>, four (including one +immature specimen) from Authiernord, + +<span class="pagenum"><a name="Page_198" id="Page_198">[Pg +198]</a></span> + +Abitibi-ouest Co., Quebec, specimens in collection of Bristol Foster +designated by numbers 2041-2044 of S. Anderson's field catalogue. Smith +and Foster (1957:107) were of the view that <i>Phenacomys ungava</i> +(including the above specimens from Quebec) may be specifically distinct +from <i>Phenacomys intermedius</i>.</p> + + +<h3>Ondatra zibethicus (Linnaeus)</h3> +<p class="center">Not figured</p> + +<p>Baculum: In the single specimen examined, less mature than that +figured by Hamilton (1946:384), the digitate processes are cartilaginous, +the basal tuberosities are less well developed, and the shaft is +slenderer throughout. The cartilaginous processes are of the same +proportions as ossified processes in the figure mentioned. The shaft is +also convex ventrally in lateral profile. The view of the side here +considered to be anatomically the ventral side (adjacent to the urethra) +is labelled dorsal view in Hamilton's specimen.</p> + +<p><i>Specimen examined</i>: One, from Reserve, Brown Co., Kansas, 72405.</p> + + +<h3>Microtus (Herpetomys) guatemalensis Merriam</h3> +<p class="center">Figs. 42 and 43</p> + +<p>Baculum: Stalk moderately elongate, greatest length (3.5 mm.) +2<sup>1</sup>/<sub>3</sub> times greatest breadth, spatulate, flattened +throughout, greatest thickness <sup>1</sup>/<sub>3</sub> millimeter; +three ossified processes; median process having three cornered base, +curved dorsally, wider than high, ¼ to <sup>1</sup>/<sub>5</sub> +greatest length of stalk; each lateral process bent at middle, as long as +median process, compressed laterally; base of stalk curved dorsally, +tuberosities marginal, hence narrow, lateral excavations of tuberous +margin not confluent medially; in end-view ventral concavity broad, no +dorsal concavity, medial constriction but slightly less than greatest +thickness (not depth); shaft wider than high throughout, at mid-point +more than 3 times as wide as high; tip of shaft slightly inflated both +laterally and dorsoventrally; lateral profile gradually sloping +anteriorly from widest point of stalk.</p> + +<p>Specimen number 65921 (Fig. 43) differs from number 65895 (Fig. 42) +described above. Terminus of shaft of number 65921 has lateral lobes from +which arise lateral cartilaginous processes; median terminal ossification +irregular in shape, smaller, imbedded in terminally bilobate cartilage. +In the spatulate flattened stalk these two specimens are much alike. An +immature specimen, number 65908, is smaller (length of stalk 2.6 mm.) +also flattened and spatulate, has the terminal processes cartilaginous, +the lateral processes bent medially, and proportions as in the adult.</p> + +<p>The baculum shows no noteworthy resemblance to that of any other +species of North American <i>Microtus</i>; on the other hand the +differences between <i>M. guatemalensis</i> and some other species are no +greater than the differences between certain species included in the +subgenus <i>Microtus</i>. The baculum neither strengthens nor weakens the +case for subgeneric rank for <i>M. (Herpetomys) guatemalensis</i>.</p> + +<p><i>Specimens examined</i>: Three from Guatemala; 65895 (2 mi. S San +Juan Ixcoy), 65908, (3-1/2 mi. SW San Juan Ixcoy), 65921 (10 mi. E, 4 mi. +S Totonicapán).</p> + +<p><span class="pagenum"><a name="Page_199" id="Page_199">[Pg +199]</a></span></p> + + +<h3>Microtus (Arvicola) richardsoni (DeKay)</h3> +<p class="center">Figs. 38 and 39</p> + +<p>Baculum: Stalk broad, greatest length (3.7 to 4.3 mm.) 1½ times +greatest breadth, relatively flattened, greatest depth +<sup>1</sup>/<sub>3</sub> greatest breadth; single median ossified +process, in smaller of two specimens this ossification incomplete and of +unusual shape (Fig. 39); length of stalk 4 times length of median +process; concavities of basal tuberosities medially confluent, +constriction less than ½ greatest depth; widest point of shaft +less than ¼ length of shaft from posteriormost point; shaft wider +than high except at distal end that is inflated dorsally and sometimes +laterally; both ventral and dorsal concavities of base of stalk broad and +moderately deep; posterior profile in dorsal view evenly rounded or +having marginal notch.</p> + +<p>In the absence of ossified lateral processes my two specimens differ +from bacula of <i>Microtus (Arvicola) terrestris</i> figured by Didier +(1943:79, 1954:245, 247, 248) and by Ognev (1950:591). The median process +relative to the size of the shaft is smaller, and the shaft relative to +its length is wider in <i>M. richardsoni</i> than in <i>M. +terrestris</i>. The stalk of <i>M. (Arvicola) amphibius</i> figured by +Didier is like that of <i>M. richardsoni</i> in its greater breadth and +median notch on posterior border.</p> + +<p>The relationship of the New World water rat, <i>M. richardsoni</i>, to +the Old World water rats (genus <i>Arvicola</i> of some European authors) +is uncertain. Miller (1896:66) placed all of them in the subgenus +<i>Arvicola</i>. Subsequent authors, stressing differences in the teeth, +have placed <i>M. richardsoni</i> in the subgenus <i>Aulacomys</i> of +Rhoads. Zimmerman (1955) has shown that teeth in some <i>Arvicola</i> +approach the more complex pattern of <i>M. richardsoni</i>. He argues +also that <i>Arvicola</i> is generically distinct from <i>Microtus</i> on +the grounds that the two groups have separate origins, <i>Arvicola</i> +having descended from the genus <i>Mimomys</i> and <i>Microtus</i> from +some other group of microtines. This argument also was advanced by Hinton +(1926:47-48). Pending further studies of the possible polyphyletic origin +of other subgenera of the genus <i>Microtus</i>, I refer both <i>M. +richardsoni</i> and <i>M. terrestris</i> to the subgenus <i>Arvicola</i>.</p> + +<p>The evidence afforded by the bacula available is not conclusive as to +relations of Old World and New World water rats. No general agreement on +the number of species in this Palaearctic group has been reached, and +bacula of only three or four of the numerous Old World subspecies have +been figured. I have examined none.</p> + +<p><i>Specimens examined</i>: Two, from Wyoming; 42454 (31 mi. N +Pinedale, 8025 ft., Sublette Co.), 37903 (23-1/2 mi. S, 5 mi. W Lander, +8600 ft., Fremont Co.).</p> + + +<h3>Microtus (Chilotus) oregoni (Bachman)</h3> +<p class="center">Fig. 45</p> + +<p>Baculum: Stalk broad, greatest length (2.2 mm.) 1¾ times +greatest breadth, 3½ times greatest depth; three well-developed +ossified processes; median process <sup>2</sup>/<sub>5</sub> length of +stalk, rounded or tapered terminally, proximal end opposed to tip of +stalk and flattened obliquely; lateral processes +<sup>2</sup>/<sub>3</sub> length of median process, deeper than wide, +curved; tuberosities of stalk well developed, confluent medially, visible +in dorsal view; in end-view dorsal concavity narrow, moderately + +<span class="pagenum"><a name="Page_200" id="Page_200">[Pg +200]</a></span> + +deep, rounded, ventral concavity wide, deep, flattened; base wider +ventrally than dorsally; shaft tapering more or less uniformly, +terminally inflated.</p> + +<p>In the relative sizes, to each other and to the stalk, of the three +digitate ossifications <i>M. oregoni</i> resembles closely the Old World +representative of the same subgenus, <i>M. (Chilotus) socialis</i>, as +figured by Argyropulo (1933b:181). In <i>M. oregoni</i> the greatest +width of the baculum is more proximal on the stalk than in the <i>M. +socialis</i> figured by Argyropulo but closely resembles the baculum of +the <i>M. socialis</i> figured by Didier (1954:242). In possessing a +shallow emargination in the base of the stalk and in possessing a median +process that is smaller than the lateral processes, <i>M. socialis</i>, +as figured by Didier, differs from <i>M. oregoni</i>. The baculum figured +by Argyropulo (<i>loc. cit.</i>) of <i>Sumeriomys colchicus +schidlovskii</i> [ = <i>Microtus (Chilotus) socialis schidlovskii</i> +according to Ognev, 1950:392] differs from other <i>Chilotus</i> that +have been studied in having an unusually elongate median process and a +more distal placement of the widest part of the stalk.</p> + +<p><i>Specimens examined</i>: Three, of the subspecies <i>M. oregoni +oregoni</i>, from 5 mi. N Orick, Humboldt Co., California, 3-C-248, +collection of W. B. Quay; from Mary's Peak, Benton Co., Oregon, 66, +collection of F. W. Sturges; and from Sec. 3, T. 11S, R. 5W, Benton Co., +Oregon, 79183.</p> + + +<h3>Microtus (Stenocranius) gregalis (Pallas)</h3> +<p class="center">Fig. 34</p> + +<p>Baculum: Length of stalk (2.4 mm.) 1¾, times greatest breadth, +4<sup>1</sup>/<sub>3</sub> times greatest depth; median ossified process +well developed, more than <sup>1</sup>/<sub>3</sub> length of stalk, +higher than wide, slightly bowed, closely appressed to terminus of shaft; +basal tuberosities of stalk moderately developed, confluent medially, +posterior profile of medial apex rounded in dorsal view, lateral +indentations present, hence trilobate outline; in proximal end-view base +wider ventrally, ventral concavity broader than dorsal concavity but of +equal depth, medial constriction <sup>2</sup>/<sub>3</sub> greatest +depth; shaft slender in distal part, inflated terminally, and wider than +high at mid-point of stalk; lateral profile a smooth slope of gradually +decreasing curvature from point of greatest width to near distal end.</p> + +<p>The baculum of this species figured by Ognev (1950:461) differs in +having lateral ossified processes, and a more rounded base of the stalk. +Resemblance to the New World <i>Stenocranius</i> is discussed below.</p> + +<p><i>Specimen examined</i>: One from "Eastern Europe," 8059.</p> + + +<h3>Microtus (Stenocranius) miurus Osgood</h3> +<p class="center">Figs. 32 and 33</p> + +<p>Baculum: Length of stalk (2.8 mm.) 1½ times greatest breadth, +3½ times greatest depth; median process ossified, +<sup>2</sup>/<sub>5</sub> to <sup>3</sup>/<sub>5</sub> length of stalk, +laterally compressed, sometimes arched in dorsoventral plane; lateral +processes cartilaginous, slender; basal tuberosities well developed, +averaging less enlarged than shown in Figure 32, but more angular in +lateral outline than shown in Figure 33; tuberosities confluent +posteriorly; posterior profile smoothly rounded to trilobate, curvature +at point of greatest breadth usually acute; in proximal end-view base +wider dorsally, deep dorsal concavity, shallow ventral concavity, medial +constriction <sup>3</sup>/<sub>5</sub> of greatest depth; shaft slender +anteriorly, at mid-point of stalk + +<span class="pagenum"><a name="Page_201" id="Page_201">[Pg +201]</a></span> + +twice as wide as high, at tip higher than wide, laterally inflated; +lateral profile in most specimens abruptly curved anterior to point of +greatest breadth.</p> + +<p>The single specimen of the Old World <i>M. (Stenocranius) gregalis</i> +examined resembles the New World <i>M. (Stenocranius) miurus</i> in the +angular lateral profile at the point of greatest breadth of the stalk, +slender shaft in comparison to broad base of stalk, and presence of a +single well-developed laterally compressed median process. The base of +the stalk in the baculum of <i>M. gregalis</i> is less well developed and +smaller than in the baculum of <i>M. miurus</i>.</p> + +<p><i>Specimens examined</i>: Nine, all of the subspecies <i>Microtus +miurus muriei</i>, from the Brooks Range, Alaska; 51077 (Lake Schrader, +145°09'50", 69°24'28", 2900 ft., Romanzof Mts.); 51151, 51152, +51154, 51164, 51166, 51169 (last 6 from Wahoo Lake, 69°08', +146°58', 2350 ft.); 51210, 51213 (last 2 from Porcupine Lake, +68°51'57", 146°29'50", 3140 ft.).</p> + + +<h3>Microtus (Chionomys) nivalis Martins</h3> +<p class="center">Fig. 47</p> + +<p>Baculum: Greatest length of stalk (2.7 mm.) 2¼ times greatest +breadth, 4½ times greatest depth; three digitate processes, +lateral processes mostly cartilaginous in single adult examined; median +process well ossified, approximately <sup>1</sup>/<sub>3</sub> length of +stalk, basally notched, not arched, laterally compressed distally; base +of stalk broad and flat, basal tuberosities well developed, separate; +posterior profile in dorsal view rounded, convex except for medial notch +separating tuberosities; dorsal and ventral concavities deep, broad, +equal; medial constriction less than ½ greatest depth; in dorsal +view shaft tapering gradually from widest point, terminally rounded; at +mid-point of stalk almost twice as wide as high.</p> + +<p>In the elongate, largely cartilaginous lateral processes of the +baculum, the specimen described above resembles <i>M. longicaudus</i>. +The size of the median process in comparison to the size of the stalk is +also the same. The lateral processes have larger ossifications and the +base of the stalk is more robust in <i>M. longicaudus</i> than in <i>M. +nivalis</i>.</p> + +<p>The well ossified lateral processes and enlarged base of Didier's +(1954:240) specimen suggest that it is of a more mature individual than +the one described above. These specimens of <i>M. nivalis</i>, as well as +the specimens of <i>M. longicaudus</i>, exhibit dorso-ventral flattening +of the mid-part of the base of the stalk.</p> + +<p>The baculum of a specimen from Switzerland is weakly developed, of +small size (shaft 2.0 mm. in length), slender, thin, spatulate, and +terminally inflated. Digital processes were not observed, perhaps owing +to excessive maceration in preparation. The general appearance of the +baculum is that of an immature individual, although the animal was not +small (165 mm. total length in preservative).</p> + +<p><i>Specimens examined</i>: Two <i>Microtus nivalis nivalis</i>; +Zermatt, Valais, Switzerland, 67105; Wetterstein, Germany, 65127.</p> + + +<h3>Microtus (Chionomys) longicaudus (Merriam)</h3> +<p class="center">Fig. 48</p> + +<p>Baculum: Base of stalk well developed, greatest length (3 mm.) +1¾ times greatest breadth, 3<sup>2</sup>/<sub>3</sub> times +greatest depth; three ossified processes; base of median process rounded; +median process slightly curved in dorsoventral plane, in length almost +<sup>1</sup>/<sub>3</sub> greatest length of stalk; ossifications in +lateral processes + +<span class="pagenum"><a name="Page_202" id="Page_202">[Pg +202]</a></span> + +variable in size, frequently widely separated from shaft by cartilage, +rarely as large as median ossification; basal tuberosities usually +well-developed, medially confluent; profile of base in dorsal view +trilobate or irregularly convex throughout; constriction ½ +greatest depth; shaft relatively straight or slightly bowed ventrally or +dorsally, shaft at mid-point of stalk wider than high; tip of shaft +laterally inflated; widest point of stalk approximately ¼ length +of stalk from proximal end; lateral profile in dorsal view tapers +gradually onto shaft anteriorly from point of greatest width of stalk; +shaft variable, from slender terminally and nearly parallel sided (Fig. +48), to broad distally and tapered.</p> + +<p>In many of the features that distinguish <i>M. longicaudus</i> (and +the closely related insular species <i>M. coronarius</i>) from other +North American <i>Microtus</i>, <i>longicaudus</i> resembles the Old +World species of the subgenus <i>Chionomys</i> (that is to say, <i>M. +nivalis</i>, <i>M. gud</i>, and <i>M. roberti</i>). These features are +medium size, long tail, grayish color, montane habitat, relatively short +molar tooth-row, moderate sized and unconstricted incisive foramen, +relatively decurved upper incisors, elongate nasals, relatively broad +interorbital region without well-developed median ridge, and similar +chromosomes (Matthey, 1955:178). For these reasons I am here referring +<i>Microtus longicaudus</i> to the subgenus <i>Chionomys</i>; previously +it has not been referred to that subgenus.</p> + +<p><i>Specimens examined</i>: Six, of three subspecies; <i>Microtus +longicaudus littoralis</i>, Sullivan Island, Alaska, 42972, 42969; <i>M. +l. mordax</i>, 3/4 mi. N, 2 mi. W Allenspark, 8400 ft., Boulder Co., +Colorado, 50335, 76829; <i>M. l. sierrae</i>, Crane Flat, Mariposa Co., +California, 50252, 50253.</p> + + +<h3>Microtus arvalis (Pallas)</h3> +<p class="center">Fig. 22</p> + +<p>Baculum: In the single specimen examined, stalk small, greatest length +(2.3 mm.) 2<sup>1</sup>/<sub>3</sub> times greatest width, almost 6 times +greatest depth, flattened proximally; three well-developed digitate +processes, the median one ossified, the lateral processes cartilaginous; +median ossification laterally compressed and decurved at tip, bilobate at +base; basal tuberosities of stalk weakly developed, medially confluent; +posterior profile in dorsal view evenly rounded; ventral concavity deeper +and narrower than dorsal concavity, but both comparatively shallow; +medial constriction <sup>2</sup>/<sub>3</sub> greatest depth; shaft +straight, at mid-point twice as wide as deep; lateral profile tapering +from greatest width gradually to parallel sides of distal third of +stalk.</p> + +<p>From the baculum of <i>Microtus arvalis</i> figured by Ognev +(1950:173), and from the baculum figured by Didier (1954:238) my specimen +differs in the absence of lateral ossifications in the digitate +processes, smaller and slenderer median ossification, and weaker base. +These differences in part may be owing to a difference in age, my +specimen being the less mature.</p> + +<p><i>Specimen examined</i>: One from Vidy, Switzerland, 67101.</p> + + +<h3>Microtus orcadensis Millais</h3> +<p class="center">Fig. 24</p> + +<p>Baculum: In the one specimen examined, stalk broad, greatest length +(2.6 mm.) 1½ times greatest breadth, 3½ times greatest +depth; three digitate processes ossified; median process relatively +broad, in length more than ½ length of stalk, triangular in dorsal +view, with small spurs posterolaterally, + +<span class="pagenum"><a name="Page_203" id="Page_203">[Pg +203]</a></span> + +middorsal ridge posteriorly; lateral ossifications slightly curved, +slenderer, less than ½ depth and less than ½ transverse +thickness of median process; basal tuberosities well-developed, confluent +medially; in end-view base wider dorsally than ventrally, dorsal +concavity broader and more abruptly curved at mid-point than ventral +concavity; constriction ½ greatest depth; posterior profile in +dorsal view notched, setting off a posterior shelf; stalk including shaft +wider than deep throughout, at mid-point width twice depth; lateral +profile abruptly curved anterior to point of greatest width, sides of +shaft tapering gradually anteriorly to rounded uninflated tip.</p> + +<p>The baculum of this insular species, placed in the "<i>arvalis</i>" +group by Ellerman (1941:595), resembles the baculum of both <i>Microtus +agrestis</i> and <i>Microtus guentheri</i> more than it resembles the +baculum of <i>Microtus arvalis</i>. Similarities in the chromosomes of +<i>M. arvalis</i> and <i>M. orcadensis</i> were noted by Matthey +(1953:254, 279), who was of the opinion that <i>M. orcadensis</i> is an +insular derivative of the <i>arvalis</i>-group.</p> + +<p><i>Specimen examined</i>: One from the Orkney Islands, 67106.</p> + + +<h3>Microtus guentheri Danford and Alston</h3> +<p class="center">Fig. 23</p> + +<p>Baculum: In the one specimen examined, stalk broad, greatest length +(2.9 mm.) 1½ times greatest breadth, 3½ times greatest +depth; three digitate processes ossified; median process slightly less +than ½ length of stalk, broad, dorsally curved; curved lateral +ossifications shorter and more slender than median ossification; basal +tuberosities well developed, angular, confluent across posterior border +of projecting shelf; in end-view tuberosities projecting ventrolaterally +from central shelf; dorsal surface at medial constriction flat, ventral +surface broadly and deeply concave; posterior profile in dorsal view +trilobate, central lobe formed by posteriorly flattened shelf, surface of +attachment visible only on lateral lobes; at mid-point stalk almost twice +as wide as deep, depth of shaft greater than width proximal to inflated +terminus.</p> + +<p><i>Specimen examined</i>: One from Palestine, 67104.</p> + + +<h3>Microtus fortis Büchner</h3> +<p class="center">Fig. 25</p> + +<p>Baculum: Stalk large, greatest length (3.8 mm.) +1<sup>4</sup>/<sub>5</sub> times greatest breadth, 4½ times +greatest depth; three digitate processes ossified; median ossification +almost <sup>1</sup>/<sub>3</sub> length of stalk; lateral ossifications +slender, smaller than median ossification; posterior profile of stalk in +dorsal view trilobate, basal tuberosities well developed, confluent +medially; in end-view dorsal concavity broader and deeper than ventral +concavity; medial constriction pronounced (less than ½ greatest +depth); lateral profile at widest point of stalk convex, becoming +abruptly concave as the flange of the basal tuberosities grades into the +shaft, then gradually converging to narrowest point +<sup>1</sup>/<sub>3</sub> of length of stalk from the terminus; stalk +wider than deep in proximal <sup>2</sup>/<sub>3</sub>, circular in cross +section in terminal <sup>1</sup>/<sub>3</sub>, slight terminal +inflation.</p> + +<p>A specimen figured by Ognev (1950:297) has the same general +proportions, slender lateral processes, and proximal placement of the +point of greatest breadth.</p> + +<p><i>Specimens examined</i>: Two from Chipo-ri, Korea, 60443, 63841.</p> + +<p><span class="pagenum"><a name="Page_204" id="Page_204">[Pg +204]</a></span></p> + + +<h3>Microtus montanus (Peale)</h3> +<p class="center">Figs. 19, 20 and 21</p> + +<p>Baculum: Stalk broad, greatest length (varying with subspecies from +2.3 to 3.1 mm.) 1½ to 1¾ times greatest breadth, +3<sup>1</sup>/<sub>3</sub> to 4<sup>1</sup>/<sub>3</sub> times greatest +depth; three ossified processes, median one largest, more than twice as +wide and as deep as shorter, slenderer, lateral processes; median process +laterally compressed distally except in one specimen in which moderately +inflated distally, proximally enlarged in some specimens (Fig. 21) and +<sup>1</sup>/<sub>3</sub> to <sup>2</sup>/<sub>5</sub> length of stalk; +base broad, posterior profile in dorsal view evenly convex throughout, at +widest point of stalk abruptly incurved; basal tuberosities moderately to +strongly developed, medially confluent; in end-view base wider ventrally +than dorsally, dorsal concavity slightly to much deeper than the nearly +flattened ventral concavity; medial constriction +<sup>2</sup>/<sub>3</sub> to <sup>4</sup>/<sub>5</sub> of greatest depth; +shaft relatively slender, at mid-point of stalk slightly wider than high +and ¼ as wide as base of stalk, terminally rounded or slightly +inflated; lateral profile in dorsal view a gradual curve from point of +greatest width anteriorly onto shaft.</p> + +<p>The different subspecies figured show the essential characteristics of +the species, differing primarily in size.</p> + +<p><i>Specimens examined</i>: Fourteen, of three subspecies; <i>Microtus +montanus amosus</i>, ½ mi. E Soldier Summit, Wasatch Co., Utah, +62241; <i>M. montanus fusus</i>, La Manga Pass, Conejos Co., Colorado, +42164; 5 mi. N, 26 mi. W Saguache, 9500 ft., Saguache Co., Colorado, +42307, 42315; 5 mi. N, 27 mi. W Saguache, 9350 ft., Saguache Co., +Colorado, 42308; 5 mi. N, 28 mi. W Saguache, 9325 ft., Saguache Co., +Colorado, 42309; 5 mi. S, 24 mi. W Antonito, 9600 ft., Conejos Co., +Colorado, 42327, 42330; Prater Canyon, Mesa Verde National Park, +Montezuma Co., Colorado, 69456, 69457, 69463; <i>Microtus montanus +nanus</i>, 2 mi. N, 2 mi. W Pocatello, Bannock Co., Idaho, 57470, 57472; +¼ mi. N, 2 mi. W Allenspark, 8400 ft., Boulder Co., Colorado, +50330.</p> + + +<h3>Microtus townsendii (Bachman)</h3> +<p class="center">Fig. 41</p> + +<p>Baculum: Stalk broad, greatest length (3.0 mm.) 1½ times +greatest breadth, 4½ times greatest depth; three ossified +processes, median one largest, deeper and more than twice as wide as +curved, shorter, compressed lateral processes and more than +<sup>2</sup>/<sub>5</sub> as long as stalk; base broad, in dorsal view +posterior profile trilobate, basal tuberosities visible; basal +tuberosities well developed, medially confluent; in end-view base wider +ventrally than dorsally, dorsal concavity deeper than ventral concavity; +medial constriction <sup>3</sup>/<sub>5</sub> of greatest depth; shaft +broad, at mid-point more than twice as wide as high and +<sup>1</sup>/<sub>3</sub> as wide as base of stalk, terminally +rounded.</p> + +<p><i>Specimens examined</i>: Three, all <i>M. t. townsendii</i>; Fort +Lewis, Pierce Co., Washington, 57998, subadult; Sec. 33, T. 11S, R. 5W, +Benton Co., Oregon, 79186; Sec. 5, T. 12S, R. 4W, Benton Co., Oregon, +79188.</p> + + +<h3>Microtus oeconomus (Pallas)</h3> +<p class="center">Fig. 44</p> + +<p>Baculum: Stalk broad and flattened, greatest length (3.5 mm.) +1<sup>2</sup>/<sub>3</sub> to 2 times greatest width, 4 to 5½ +times greatest depth; three ossified processes, median one largest, +lateral processes slender, relatively small; length of median process +<sup>3</sup>/<sub>8</sub> length of stalk; median process decurved, +dorsoventrally flattened in + +<span class="pagenum"><a name="Page_205" id="Page_205">[Pg +205]</a></span> + +some specimens, widened at base; attachment of processes to shaft +displaced ventrally; base of stalk widened, posterior profile in dorsal +view usually trilobate, in a few cases rounded, median lobe forming +posterior shelf, lateral lobes dorsally raised and forming margins of +lateral tuberosities; in end-view thickness frequently more or less +uniform throughout central part, broad depression dorsally, ventral +concavity narrower and shallower (as figured); base, and occasionally +shaft, flattened, width at mid-point of stalk 2 to 3 times depth, +narrowest point posterior to terminal inflation of shaft in terminal +<sup>1</sup>/<sub>3</sub> of shaft.</p> + +<p>The baculum of <i>M. oeconomus</i> (Old World) figured by Ognev +(1950:257) resembles but exceeds that of <i>M. oeconomus</i> (New World) +in the relatively large median process and slender lateral processes, but +differs noticeably in the presence of a deep median notch in the base of +the stalk. A specimen from Hungary is intermediate between Ognev's +specimen and those from the New World in both size of median process and +size of lateral processes, and has an unnotched base resembling that in +Figure 44.</p> + +<p><i>Specimens examined</i>: Ten, of three subspecies; <i>M. oeconomus +gilmorei</i>, Umiat, Alaska, 51354, 51361, 51399, 51408; Lake Schrader, +Brooks Range, Alaska, 51422; <i>M. o. macfarlani</i>, 5 mi. NNE Gulkana, +Alaska, 43039, 43041; 20 mi. NE Anchorage, Alaska, 43044; Kelsall Lake, +British Columbia, 43048; <i>M. o. mehelyi</i>, Kisbalatan, Hungary, +75159.</p> + + +<h3>Microtus mexicanus (Saussure)</h3> +<p class="center">Figs. 35 and 36</p> + +<p>Baculum: Stalk attenuate, greatest breadth relatively near proximal +end; greatest length (3.1 to 3.4 mm.) more or less twice greatest +breadth, 4 to 5 times greatest depth; usually a single process ossified; +lateral processes relatively small, cartilaginous or (in three specimens, +63094, 69453, 68019) with small ossifications; median process relatively +small, sometimes appressed to tip of shaft, in length less than ¼ +length of stalk; posterior profile in dorsal view rounded, flattened +posteriorly, or in some specimens trilobate with angular edges; in +end-view relative depths of dorsal and ventral concavities variable, +dorsal usually deeper than ventral; distal end of stalk frequently bowed +dorsally; shaft slender distally, sometimes slightly inflated terminally, +or (in one specimen, 63085) near tip small lateral projections that are +perhaps fused lateral ossifications; lateral profile in dorsal view a +gradual slope anteriorly from point of greatest width to slender tip.</p> + +<p><i>Specimens examined</i>: Thirteen, of four subspecies; <i>Microtus +mexicanus mexicanus</i>, Las Vigas, Veracruz, 30692; Nevada de Toluca, +México, 63101; Valle de Bravo, México, 63094; <i>Microtus mexicanus +mogollonensis</i>, Mt. Taylor, Valencia Co., New Mexico, 63298, 76830; +Park Well, Mesa Verde National Park, Montezuma Co., Colorado, 69448, +69453; Upper Nutria, McKinley Co., New Mexico, 69997, 70000; <i>Microtus +mexicanus phaeus</i>, Sierra Patamba, 9000 ft., Michoacán, 63085; +<i>Microtus mexicanus subsimus</i>, 2 mi. E Mesa de Tablas, Coahuila, +58916; 13 mi. E San Antonio de las Alazanas, Coahuila, 68019, 68021.</p> + + +<h3>Microtus californicus (Peale)</h3> +<p class="center">Fig. 37</p> + +<p>Baculum: Stalk elongate, greatest length (3.0 mm.) +2<sup>1</sup>/<sub>3</sub> times greatest breadth, 4½ times +greatest depth; median process ossified, ¼ length of stalk, +basally broadened, flattened and shallowly grooved ventrally to fit tip +of shaft, to which the process is closely appressed; lateral processes +cartilaginous; ends + +<span class="pagenum"><a name="Page_206" id="Page_206">[Pg +206]</a></span> + +of stalk bowed upwardly; posterior profile of base of stalk rounded or +slightly trilobate if posterolateral concavities form in tuberosities; +moderate development of tuberosities, in end-view dorsal concavity +slightly deeper and narrower than ventral concavity, both comparatively +shallow, median constriction <sup>4</sup>/<sub>5</sub> greatest depth; +shaft curved, more or less terete at mid-point of stalk, terminally +inflated dorsally; lateral profile in dorsal view gradually curved from +point of greatest width anteriorly onto shaft.</p> + +<p><i>Specimens examined</i>: Two, of two subspecies; <i>Microtus +californicus californicus</i>, 1 mi. NE Berkeley, in Contra Costa Co., +California, 76828; <i>Microtus californicus mohavensis</i>, ½ mi. +SE Victorville, San Bernardino Co., California, 63745.</p> + + +<h3>Microtus pennsylvanicus (Ord)</h3> +<p class="center">Figs. 14, 15, 16 and 17</p> + +<p>Baculum: Stalk heavy, broad, greatest length (2.2 to 3.0 mm.) +1<sup>1</sup>/<sub>3</sub> to 1<sup>2</sup>/<sub>3</sub> times greatest +breadth, up to 3¾ times greatest depth; three ossified processes, +median one largest, usually not twice so deep as lateral ossifications; +median process usually distinctly widened basally, in length +approximately ½ length of stalk; base broad, frequently angular +laterally and basally, sometimes bilobate; basal tuberosities well +developed, medially confluent; in end-view more or less uniformly +biconvex or ventral surface more flattened than dorsal surface, medial +constriction ½ to <sup>2</sup>/<sub>3</sub> greatest depth; shaft +relatively heavy, at mid-point stalk almost twice as wide as deep and +<sup>1</sup>/<sub>3</sub> as wide as base of stalk; shaft terminally +rounded and sometimes slightly inflated; lateral profile in dorsal view +abruptly or gradually curved anterior to point of greatest width and then +gradually curved anteriorly.</p> + +<p>Specimens examined averaged slightly smaller and were more variable +than those described by Hamilton (1946:382). The greater variation may be +in part geographic, as five subspecies are represented. Lateral processes +are the last to ossify. One specimen (75082) with well-ossified median +process lacks any lateral ossification. Four bacula of <i>M. +pennsylvanicus</i> (locality not specified) studied by Dearden (1958:547) +agree in general with the description above.</p> + +<p>One specimen shows a break, perhaps resulting from injury, in the +shaft (Fig. 14). One specimen has a posteromedian spine on the median +digital ossification (Fig. 16). Comparison with <i>M. agrestis</i> is +included with the description of <i>M. agrestis</i>.</p> + +<p><i>Specimens examined</i>: Thirteen, of six subspecies; <i>Microtus +pennsylvanicus alcorni</i>, 20 mi. NE Anchorage, Alaska, 43043; +<i>Microtus pennsylvanicus finitus</i>, Laird, Yuma Co., Colorado, 68544; +<i>Microtus pennsylvanicus modestus</i>, 5 mi. N, 26 mi. W Saguache, 9500 +ft., Saguache Co., Colorado, 42306; 3 mi. N, 16 mi. W Saguache, 8500 ft., +Saguache Co., Colorado, 42416, 42417, 42418; 1 mi. S, 2 mi. E Eagle Nest, +8100 ft., Colfax Co., New Mexico, 42430, 42439; <i>Microtus +pennsylvanicus pennsylvanicus</i>, 2 mi. S, 3 mi. E Ft. Thompson, 1370 +ft., Buffalo Co., South Dakota, 42379; Vermillion, Clay Co., South +Dakota, 37070; <i>Microtus pennsylvanicus pullatus</i>, 12 mi. S, 5 mi. E +Butte, Silver Bow Co., Montana, 57501, 57503; <i>Microtus pennsylvanicus +uligocola</i>, Muir Springs, 2 mi. N, 2½mi. W Ft. Morgan, Morgan +Co., Colorado, 75082.</p> + + +<h3>Microtus agrestis (Linnaeus)</h3> +<p class="center">Fig. 18</p> + +<p>Baculum: Greatest length of stalk (2.9 mm.) twice greatest breadth, +4½ times greatest depth; stalk well developed, shaft not flattened +dorsoventrally; large median ossified process, minute lateral +ossifications in single specimen + +<span class="pagenum"><a name="Page_207" id="Page_207">[Pg +207]</a></span> + +examined; length of stalk 2½ times length of median ossification +which is higher than wide, slightly decurved, sagittate in dorsal view, +with three-cornered base; basal tuberosities of stalk moderately well +developed, medially joined; posterior profile in dorsal view evenly +rounded; ventral concavity broader than, but of comparable depth to, +dorsal concavity in end-view, base of stalk wider ventrally, constriction +¾ greatest depth; at mid-point of stalk shaft is but slightly +wider than high; pronounced terminal inflation of shaft; lateral profile +in dorsal view sloping abruptly from widest point of stalk anteriorly +onto stalk which then tapers more gradually to terminal inflation.</p> + +<p>From the baculum of its New World counterpart, namely <i>Microtus +pennsylvanicus</i>, my specimen of <i>Microtus agrestis</i> and the +specimen figured by Didier (1954:239) differ in their minute lateral +processes, relatively larger median processes, and more elongate, less +dorsoventrally flattened shafts.</p> + +<p>The specimen of <i>M. agrestis</i> figured by Ognev (1950:320), in +dorsal view has lateral concavities producing a somewhat trilobate +outline in the base of the stalk, and the lateral processes are well +developed; the median process is larger and bulbous, wider distally than +proximally. Without larger numbers of bacula of <i>M. agrestis</i> I am +unable to reconcile these differences. The differences between <i>M. +agrestis</i> and <i>M. pennsylvanicus</i> seem comparable to the +differences between some other species of <i>Microtus</i>.</p> + +<p><i>Specimen examined</i>: One, from Gryon, Switzerland, 67102.</p> + + +<h3>Microtus (Pedomys) ochrogaster (Wagner)</h3> +<p class="center">Fig. 31</p> + +<p>Baculum: Stalk broad, greatest length (3.2-4.0 mm.) +1<sup>2</sup>/<sub>3</sub> to 2 times greatest breadth, 2½ to 4 +times greatest depth; median process ossified, relatively small, less +than <sup>3</sup>/<sub>10</sub> length of stalk; lateral processes +arising from subterminal part of stalk, cartilaginous or with small +ossifications; posterior profile in dorsal view broadly rounded or +slightly angular, widest point of stalk <sup>1</sup>/<sub>6</sub> to +¼ the length of stalk from base; basal tuberosities well developed +and medially confluent, in end-view dorsally convex, or at least less +deeply concave than ventrally; shaft straight, base bent ventrally or +more commonly dorsally; at mid-point of stalk wider than high, often +twice as wide as high; viewed from above, lateral profile from point of +greatest breadth to middle of shaft a gradual sigmoid curve; slight +terminal inflation of shaft.</p> + +<p><i>Specimens examined</i>: Forty-one, of three subspecies; <i>Microtus +ochrogaster haydeni</i>, Muir Springs, 2 mi. N, 2½ mi. W Ft. +Morgan, Morgan Co., Colorado, 74995, 74998, 74999, 75002; 1 mi. W Laird, +Yuma Co., Colorado, 57304, 76833; 2 mi. N, 2 mi. W Haigler, Dundy Co., +Nebraska, 75016; 2 mi. S Franklin, Franklin Co., Nebraska, 75043, 75044; +Atwood, Rawlins Co., Kansas, 75020, 75023, 75025, 75027, 75028; 1 mi. N, +2 mi. E Oberlin, Decatur Co., Kansas, 75030, 75032, 75034, 75035, 75036; +1½ mi. N, ¼ mi. E Norton, Norton Co., Kansas, 68327; 1 mi. +SW Norton, Norton Co., Kansas, 75037; 2 mi. S, 1 mi. W Norton, Norton +Co., Kansas, 75038; <i>M. ochrogaster ochrogaster</i>, Rydal, Republic +Co., Kansas, 75047-75053, 75060, 75062, 75063, 75066, 75070, 75071, +75073; 1 mi. N, 1 mi. W Holton, Jackson Co., Kansas, 75077; 2 mi. W Court +House, Lawrence, Douglas Co., Kansas, 76832; Univ. Kansas Natural History +Reservation, Douglas Co., Kansas, 68536; <i>M. ochrogaster taylori</i>, +Meade County State Park, Kansas, 68539, 68542.</p> + +<p><span class="pagenum"><a name="Page_208" id="Page_208">[Pg +208]</a></span></p> + + +<h3>Microtus (Pitymys) pinetorum (LeConte)</h3> +<p class="center">Figs. 27 and 28</p> + +<p>Baculum: Stalk broad, greatest length (2.5 to 2.7 mm.) +1<sup>2</sup>/<sub>3</sub> times greatest breadth, 4 times greatest +depth; median process ossified, size small, <sup>1</sup>/<sub>5</sub> +length of stalk, higher than wide, having small anterodorsal prominence +in both specimens examined; lateral processes cartilaginous, relatively +small, displaced posteriorly, attenuate; posterior margin in dorsal view +broadly rounded, or having blunt median apex, convex throughout; basal +tuberosities moderately well developed, medially confluent, barely +visible in dorsal view when mature; in end-view median constriction +<sup>4</sup>/<sub>5</sub> greatest depth, ventral concavity deeper than +dorsal concavity, both comparatively shallow; stalk at mid-point +1½ times as wide as deep; shaft relatively slender, bowed dorsally +at tip, relatively straight otherwise; lateral profile in dorsal view a +gradual concave slope from point of greatest width anteriorly to distal +part of shaft.</p> + +<p><i>Specimens examined</i>: Two, from Douglas Co., Kansas, 76834 (2 mi. +N Baldwin), 68545 (1 mi. NE Pleasant Grove).</p> + + +<h3>Microtus (Pitymys) parvulus (Howell)</h3> +<p class="center">Fig. 40</p> + +<p>Baculum: Stalk broad, greatest length (2.4 mm. in specimen examined) +1¾ times greatest breadth, 4 times greatest depth; median process +ossified, size small, less than ¼ length of stalk, wider than +high, terminally flattened; lateral processes cartilaginous, relatively +small, attenuate; posterior margin in dorsal view flattened, irregularly +curved with concavities medially and laterally; basal tuberosities well +developed, medially confluent; visible in dorsal view; in end-view median +constriction <sup>2</sup>/<sub>3</sub> greatest depth, ventral concavity +well-formed, no dorsal concavity; stalk at mid-point twice as wide as +deep; shaft relatively slender, bowed dorsally toward tip; in dorsal view +lateral profile a gradual concave slope from point of greatest width +anteriorly to distal part of shaft; tip of shaft enlarged.</p> + +<p>The baculum of <i>M. parvulus</i> resembles that of <i>M. +pinetorum</i> more than it resembles the baculum of any other microtine +studied, differing primarily in smaller size.</p> + +<p><i>Specimen examined</i>: One, from 1 mi. W Micanopy, Alachua Co., +Florida, Univ. Florida No. 1508.</p> + + +<h3>Microtus (Pitymys) quasiater (Coues)</h3> +<p class="center">Figs. 29 and 30</p> + +<p>Baculum: Stalk broad, greatest length (2.6-3.2 mm.) +1<sup>1</sup>/<sub>3</sub> to 1<sup>2</sup>/<sub>3</sub> times greatest +breadth, 3<sup>1</sup>/<sub>3</sub> to 3<sup>2</sup>/<sub>3</sub> times +greatest depth; median process ossified, with ventral depression, process +¼ to <sup>1</sup>/<sub>3</sub> length of stalk, appressed to tip +of shaft, wider than high proximally, relatively broad terminally; +lateral processes cartilaginous, small, attenuate; posterior profile of +stalk in dorsal view broadly rounded, bilobate, or trilobate, median lobe +formed by posterior projection of dorsal shelf between enlarged lateral +tuberosities that form outer lobes, posterolateral faces of these +tuberosities visible in dorsal view of stalk; in end-view dorsal surface +slightly concave, ventral concavity broad and deep, median constriction + +<span class="pagenum"><a name="Page_209" id="Page_209">[Pg +209]</a></span> + +½ greatest depth; shaft flattened except tip that is more terete, +and bowed dorsally; at mid-point, stalk twice as wide as high; shaft +relatively slender terminally, narrower than median ossification.</p> + +<p>The baculum of <i>M. quasiater</i> is the largest and has the best +developed base and median process of the three American species of the +subgenus <i>Pitymys</i>. The three species closely resemble each other in +basic form.</p> + +<p><i>Specimens examined</i>: Five, all from Veracruz; Teocelo, 4500 ft., +30709, 30711; 4 km. N Tlapacoyán, 1700 ft., 24466; 5 km. N Jalapa, 4500 +ft., 19869, 19878.</p> + + +<h3>Microtus (Pitymys) fatioi (Mottaz)</h3> +<p class="center">Fig. 26</p> + +<p>The baculum of a single specimen (KU 67103) of <i>M. fatioi</i> from +Zermatt, Valais, Switzerland, was examined. The baculum is immature, as +evidenced by its small size, slender stalk and absence of ossified +processes, therefore no characterization is included.</p> + +<p>The baculum of another Old World species of the subgenus +<i>Pitymys</i>, <i>M. pyrenaicus</i> from France, figured and described +by Didier (1954:242-243), differs from all New World <i>Pitymys</i> +examined in processing ossified lateral processes.</p> + +<p>The status of <i>Pitymys</i>, as a genus or as a subgenus, is +uncertain. Hall and Cockrum (1953:448) considered the North American +<i>Pitymys</i> and <i>Pedomys</i> as subgenera of <i>Microtus</i>. They +did not state specifically the basis for this point of view, but mention +the fact that these two subgenera (<i>Pitymys</i> and <i>Pedomys</i>) +closely resemble each other cranially. These authors did not study nor +comment upon the status of the Old World <i>Pitymys</i>. It may be asked +whether the Old World and New World <i>Pitymys</i> have developed as +fossorial <i>Microtus</i> independently, or from an ancestor common to +both groups and not common to any other <i>Microtus</i>. Matthey +(1955:202) found 62 chromosomes (2N) in both the New World <i>Pitymys +pinetorum</i> and the Old World <i>Pitymys duodecimcostatus</i>. This +suggests, but does not prove, common ancestry.</p> + + +<h3>Neofiber alleni True</h3> +<p class="center">Fig. 49</p> + +<p>Baculum: Stalk massive, greatest length (4.7 mm.) 1¾ times +greatest breadth, 4 times greatest depth; ossification in digitate +processes variable; in one (KU 27123) of two specimens examined lateral +processes ossified and median process unossified, as in two specimens +examined by Hamilton (1946:379) from "southern Florida"; in my other +specimen (KU 27268) that is possibly more mature, median process ossified +although less deeply stained than lateral ossifications or stalk; +posterior profile in probable dorsal view roughly rounded; in end-view +probable dorsal concavity deep, ventral concavity broad but shallow, and +with center convex; median constriction <sup>3</sup>/<sub>5</sub> +greatest depth; shaft heavy, least depth <sup>2</sup>/<sub>3</sub> +greatest depth of base; stalk, at mid-point, slightly wider than deep and +more than <sup>1</sup>/<sub>3</sub> width of base; lateral profile in +dorsal view sharply incurved distal to point of greatest breadth, shaft +therefore relatively distinct from basal part of stalk; slight +subterminal constriction; tip less reduced in the two specimens examined +than in two figured by Hamilton. In preparation, the tissues that make it +possible to distinguish + +<span class="pagenum"><a name="Page_210" id="Page_210">[Pg +210]</a></span> + +with certainty the dorsal and ventral surfaces of the baculum were +removed in both specimens.</p> + +<p><i>Specimens examined</i>: Two, of the subspecies <i>Neofiber alleni +alleni</i>, 2 mi. S Gainesville, Alachua Co., Florida, 27268; 1 mi. E +Courtenay, Merritt Island, Brevard Co., Florida, 27123.</p> + + +<h3>Lagurus curtatus (Cope)</h3> +<p class="center">Fig. 46</p> + +<p>Baculum: Stalk slender, greatest length (2.5 mm.) 2 to +2<sup>2</sup>/<sub>3</sub> times greatest breadth, 4 to 5 times greatest +depth; three ossified processes; median one more than +<sup>1</sup>/<sub>3</sub> length of stalk, curved dorsally toward tip, +proximally flattened and having acute lateral angles in dorsal view, +wider than deep except in distal half; lateral processes smaller than +median one, slenderer, shorter, of approximately same depth, also curved +dorsally; base of stalk well developed, basal tuberosities medially +confluent, in part visible in dorsal view, in end-view wider ventrally +than dorsally, dorsal and ventral concavities of equal depth and both +wide, medial constriction ½ greatest depth; posterior profile in +dorsal view broadly bilobate; lateral profile with abrupt transition from +basal tuberosities to gradually converging, slightly curved sides of +shaft; shaft terminally inflated.</p> + +<p>Dearden (1958:543) described and figured the bacula of six subspecies +of <i>Lagurus curtatus</i> and two Asiatic species, <i>Lagurus +lagurus</i> and <i>Lagurus luteus</i>. He examined at least 34 specimens +of <i>L. curtatus</i> and found geographic variation in size, breadth of +shaft distally, and proportions of digital ossifications to each other +and to the stalk. The description that I have given above pertains to +<i>L. c. levidensis</i>.</p> + +<p>The baculum of the Asiatic <i>Lagurus (Lagurus) lagurus</i> figured by +Ognev (1950:554) agrees with that of <i>Lagurus (Lemmiscus) curtatus</i>, +described here, in the relatively elongate shaft and slender stalk, the +proportions of the processes, and the well-formed and moderately enlarged +base of the stalk. The bacula of three <i>Lagurus lagurus</i> examined by +Dearden (1958:545) were of older individuals than the specimen that Ognev +figures and differ from it and from bacula of <i>Lagurus curtatus</i> +(all subspecies) in the unusual, almost heart shaped, median process, and +in larger size. <i>Lagurus luteus</i> examined by Dearden (1958:545) +differs from both <i>Lagurus lagurus</i> and <i>Lagurus curtatus</i> in +lacking lateral digital ossifications and in having shorter median +digital ossifications and wider base of stalk.</p> + +<p><i>Specimens examined</i>: Seven <i>Lagurus curtatus levidensis</i> +from Wyoming; 9 mi. S Robertson, Uinta Co., 26045, 26053; 8 mi. S, +2½ mi. E Robertson, Uinta Co., 26049; Farson, Sweetwater Co., +37906; 16 mi. S, 11 mi. W Waltman, Natrona Co., 42457; 32 mi. S, 22 mi. E +Rock Springs, 42465, 42466.</p> + +<p style="margin-bottom:2em;">The following key to the bacula in some adult North American +Microtinae is intended to help point out some of the most important +differences. It should be noted that not all species can be keyed out on +the basis of the baculum. The most difficult group in this respect +includes the species of <i>Microtus</i> that have small or no ossified +lateral processes, especially species of the subgenera <i>Pedomys</i> and +<i>Pitymys</i>, and the two species <i>Microtus californicus</i> and +<i>Microtus mexicanus</i> of the subgenus <i>Microtus</i>. Another +complicating factor is the variability of bacula evident in some species +even in the small samples available. + +<span class="pagenum"><a name="Page_211" id="Page_211">[Pg +211]</a></span> + +It is to be expected that additional specimens will show variations not +yet observed.</p> + +<h2 class="smcap">Key to the Bacula of Some North American Microtines</h2> + + +<table class="key"> + <tr> + <td class="text">1. Length of lateral digital ossifications more than <sup>1</sup>/<sub>3</sub> breadth of stalk</td> + <td class="number">2</td> + </tr> + <tr> + <td class="text">1´. Length of lateral digital ossifications less than <sup>1</sup>/<sub>3</sub> breadth of stalk or absent</td> + <td class="number">15</td> + </tr> + <tr> + <td class="text">2. Size small (total length of baculum less than 5.5 mm.)</td> + <td class="number">3</td> + </tr> + <tr> + <td class="text">2´. Size large (total length of baculum more than 5.5 mm.)</td> + <td class="number">14</td> + </tr> + <tr> + <td class="text">3. Width at mid-point of stalk more than <sup>1</sup>/<sub>3</sub> greatest breadth of stalk</td> + <td class="number">4</td> + </tr> + <tr> + <td class="text">2´. Size large (total length of baculum more than 5.5 mm.)</td> + <td class="number">14</td> + </tr> + <tr> + <td class="text">3´. Width at mid-point of stalk less than <sup>1</sup>/<sub>3</sub> greatest breadth of stalk,</td> + <td class="number">8</td> + </tr> + <tr> + <td class="text">4. Stalk, viewed from proximal end hour-glass shaped, and width + of stalk less than ½ length of stalk.... <i>Phenacomys + intermedius</i>,</td> + <td class="number"><a href="#Page_197">p. 197</a></td> + </tr> + <tr> + <td class="text">4´. Stalk not both hour-glass shaped when viewed from proximal + end, and with width less than ½ length of stalk</td> + <td class="number">5</td> + </tr> + <tr> + <td class="text">5. Shaft thin basally, thickness less than <sup>1</sup>/<sub>3</sub> of greatest breadth</td> + <td class="number">6</td> + </tr> + <tr> + <td class="text">5´. Shaft thick basally, thickness <sup>1</sup>/<sub>3</sub> or more of greatest breadth</td> + <td class="number">7</td> + </tr> + <tr> + <td class="text">6. Stalk more or less straight, base not deflected. <i>Microtus + oeconomus</i>,</td> + <td class="number"><a href="#Page_204">p. 204</a></td> + </tr> + <tr> + <td class="text">6´. Stalk spatulate, and base deflected from axis of shaft.... + <i>Microtus guatemalensis</i>,</td> + <td class="number"><a href="#Page_198">p. 198</a></td> + </tr> + <tr> + <td class="text">7. Base enlarged, depth nearly ½ of breadth.... <i>Lemmus + trimucronatus</i>,</td> + <td class="number"><a href="#Page_193">p. 193</a></td> + </tr> + <tr> + <td class="text">7´. Base moderately enlarged, depth near <sup>1</sup>/<sub>3</sub> of breadth.... + <i>Microtus pennsylvanicus</i>, <a href="#Page_206">p. 206</a>, or <i>Microtus townsendii</i>,</td> + <td class="number"><a href="#Page_204">p. 204</a></td> + </tr> + <tr> + <td class="text">8. Base hour-glass shaped as viewed from proximal end.... + <i>Phenacomys intermedius</i>,</td> + <td class="number"><a href="#Page_197">p. 197</a></td> + </tr> + <tr> + <td class="text">8´. Not so</td> + <td class="number">9</td> + </tr> + <tr> + <td class="text">9. Lateral processes separated from tip of shaft by more than the + thickness of the lateral process</td> + <td class="number">10</td> + </tr> + <tr> + <td class="text">9´. Lateral processes separated from tip of shaft by less than the + thickness of the lateral process</td> + <td class="number">11</td> + </tr> + <tr> + <td class="text">10. Lateral processes more than ½ the width of median + process.... <i>Microtus longicaudus</i>,</td> + <td class="number"><a href="#Page_201">p. 201</a></td> + </tr> + <tr> + <td class="text">10´. Lateral processes slender, less than ½ the width of median + process.... <i>Microtus montanus</i>,</td> + <td class="number"><a href="#Page_204">p. 204</a></td> + </tr> + <tr> + <td class="text">11. Lateral ossifications equal in length to median + ossification.... <i>Clethrionomys</i>,</td> + <td class="number"><a href="#Page_194">p. 194</a></td> + </tr> + <tr> + <td class="text">11´. Lateral ossifications shorter than median ossification</td> + <td class="number">12</td> + </tr> + <tr> + <td class="text">12. Size small, less than 3.4 mm. in total length.... + <i>Microtus oregoni</i>,</td> + <td class="number"><a href="#Page_199">p. 199</a></td> + </tr> + <tr> + <td class="text">12´. Size medium, more than 3.4 mm. in total length</td> + <td class="number">13</td> + </tr> + <tr> + <td class="text">13. Greatest width of stalk at a point about <sup>1</sup>/<sub>3</sub> of length of + stalk from base.... <i>Microtus chrotorrhinus</i> (Hamilton, 1946:382).</td> + <td class="number"> </td> + </tr> + <tr> + <td class="text">13´. Greatest width of stalk at a point less than <sup>1</sup>/<sub>3</sub> of length of + stalk from base.... <i>Lagurus curtatus</i>,</td> + <td class="number"><a href="#Page_210">p. 210</a></td> + </tr> + <tr> + <td class="text">14. Size of baculum larger, base more than 3 mm. wide, processes + all well developed.... <i>Ondatra zibethicus</i>,</td> + <td class="number"><a href="#Page_198">p. 198</a></td> + </tr> + <tr> + <td class="text">14´. Size of baculum smaller, base less than 3 mm. wide, processes + poorly developed in some animals.... <i>Neofiber alleni</i>,</td> + <td class="number"><a href="#Page_209">p. 209</a></td> + </tr> + <tr> + <td class="text">15. At least one digital ossification present</td> + <td class="number">16</td> + </tr> + <tr> + <td class="text"> + <span class="pagenum"><a name="Page_212" id="Page_212">[Pg + 212]</a></span> + 15´. Digital ossifications not present.... <i>Dicrostonyx groenlandicus</i>,</td> + <td class="number"><a href="#Page_193">p. 193</a></td> + </tr> + <tr> + <td class="text">16. Breadth of stalk at least ½ length of stalk</td> + <td class="number">17</td> + </tr> + <tr> + <td class="text">16´. Breadth of stalk less than ½ length of stalk</td> + <td class="number">19</td> + </tr> + <tr> + <td class="text">17. Length of stalk greater than 3.6 mm. and less than 1½ + times its greatest breadth.... <i>Microtus richardsoni</i>,</td> + <td class="number"><a href="#Page_199">p. 199</a></td> + </tr> + <tr> + <td class="text">17´. Length of stalk usually less than 3.6 mm., or if more than + 3.6 mm. (up to 4.0 mm.) then length 1½ times or more its + greatest breadth</td> + <td class="number">18</td> + </tr> + <tr> + <td class="text">18. Median process attenuate distally in dorsal view, and + relatively long (more than twice its own breadth), <sup>1</sup>/<sub>5</sub> to <sup>3</sup>/<sub>5</sub> the + length of stalk; breadth of stalk usually <sup>2</sup>/<sub>3</sub> or more length of + stalk.... <i>Microtus miurus</i>,</td> + <td class="number"><a href="#Page_200">p. 200</a></td> + </tr> + <tr> + <td class="text">18´. Median process relatively blunt distally in dorsal view, + relatively short (usually less than ¼ length of stalk), breadth + of stalk usually less than <sup>2</sup>/<sub>3</sub> length of stalk.... + <i>Pitymys</i>, <a href="#Page_208">p. 208</a>, <i>Pedomys</i>, <a href="#Page_207">p. 207</a>, or <i>Microtus mexicanus</i>,</td> + <td class="number"><a href="#Page_205">p. 205</a></td> + </tr> + <tr> + <td class="text">19. Distal processes small and firmly ankylosed to distal end of + shaft.... <i>Phenacomys longicaudus</i>,</td> + <td class="number"><a href="#Page_197">p. 197</a></td> + </tr> + <tr> + <td class="text">19´. Distal processes if present not firmly ankylosed to distal + end of shaft</td> + <td class="number">20</td> + </tr> + <tr> + <td class="text">20. Dorsal concavity of base as viewed from proximal end usually + deeper than ventral concavity.... <i>Microtus mexicanus</i>,</td> + <td class="number"><a href="#Page_205">p. 205</a></td> + </tr> + <tr> + <td class="text">20´. Dorsal and ventral concavities of base equal in depth or + ventral one the deeper</td> + <td class="number">21</td> + </tr> + <tr> + <td class="text">21. Total length of baculum more than 3.6 mm.... <i>Microtus + californicus</i>,</td> + <td class="number"><a href="#Page_205">p. 205</a></td> + </tr> + <tr> + <td class="text">21´. Total length of baculum less than 3.6 mm.... <i>Synaptomys + cooperi</i>,</td> + <td class="number"><a href="#Page_194">p. 194</a></td> + </tr> +</table> + + +<h2>DISCUSSION</h2> + +<p>Owing to shortness of lower incisors and present geographic +distribution of the species, Hinton (1926:35) considered the Tribe Lemmi +(lemmings) to be more primitive than the Tribe Microti (voles). The +surviving lemmings are specialized in many features and therefore are +considered as advanced end-products of an evolutionary radiation of a +primitive microtine stock, of which all earlier stages are extinct.</p> + +<p>Hinton regarded <i>Dicrostonyx</i> as the most primitive of the genera +of lemmings on account of its more complex molar teeth (complexity was +considered to be primitive), and on account of the presence of three +primitive longitudinal rows of tubercles in unworn molars. The other +three genera were arranged in order of increasing specialization as +follows: <i>Synaptomys</i>, <i>Myopus</i>, <i>Lemmus</i>.</p> + +<p>If the baculum tended to retain its primitive character while +specializations in the external anatomy developed, and if the above +arrangement is correct the most primitive bacula would be found +in <i>Dicrostonyx</i> and in <i>Synaptomys</i>. The baculum in these two +genera in comparison to that in <i>Myopus</i> (as figured by Ognev, 1948:512) + +<span class="pagenum"><a name="Page_213" id="Page_213">[Pg 213]</a></span> + +and <i>Lemmus</i> has a slenderer stalk and smaller digital ossifications +or none at all. The baculum in the genera of lemmings increases in +robustness and the development of processes from <i>Dicrostonyx</i>, to +<i>Synaptomys</i>, to <i>Myopus</i>, to <i>Lemmus</i>—the same +order outlined above for total of specialization. The two extremes in +this series are near the extremes of variation in bacula to be found in +all microtines. The baculum in lemmings as a group cannot then be +considered more primitive than in voles as a group, although the voles +are usually considered to be more advanced. The situation in the voles, +as we shall see, casts a different light on the matter.</p> + +<p>The voles, Tribe Microti, were considered by Hinton (1926:40) to be +more advanced than the lemmings because the incisors of the voles are +longer and the root of their last lower molar is lingual to the root of +the incisor. Hinton thought also that the murine ancestors of microtines +had shorter incisors and that the backward extension of the incisors in +the voles is a more ancient feature than the hypsodonty of the molars. A +trend in the molar teeth has been toward greater hypsodonty. The voles in +which the molars are least hypsodont are thus considered primitive. These +include the living genera <i>Clethrionomys</i>, <i>Phenacomys</i>, +<i>Ondatra</i>, <i>Dolomys</i>, <i>Ellobius</i>, and <i>Prometheomys</i>. +Therefore, the baculum, in these assumedly primitive genera, would be +expected to resemble the baculum in the lemmings or at least the most +primitive lemmings. This is not the case.</p> + +<p>The bacula that I have examined of <i>Clethrionomys</i> and +<i>Phenacomys</i> have well-developed digital ossifications. In this they +resemble the baculum of the genus <i>Lemmus</i>, the most advanced genus +of lemmings according to Hinton. The baculum of <i>Dolomys</i> has not +been studied. The baculum in <i>Ondatra</i>, and in <i>Prometheomys</i> +as illustrated by Ognev (1948:552), also possesses well-developed +processes. The baculum of <i>Ellobius</i> is small and lacks processes +(as figured by Ognev, 1950:662). No ossification was found in a single +specimen of <i>Ellobius</i> examined by me although the entire glans +penis was removed and cleared without dissection. So far as known then, +with the exception of <i>Ellobius</i> and <i>Phenacomys longicaudus</i> +(Dearden, 1958:547), the primitive microtines having rooted molars +possess bacula having three well-developed ossified processes.</p> + +<p>Voles of the genus <i>Microtus</i> vary in the structure of the +baculum almost as much as do the lemmings. Within the single subgenus +<i>Microtus</i> some individuals of <i>Microtus mexicanus</i>, for +example, have minute ossified lateral processes and other individuals +lack these + +<span class="pagenum"><a name="Page_214" id="Page_214">[Pg 214]</a></span> + +processes; <i>Microtus pennsylvanicus</i> and some other species have +proportionately large lateral ossifications. If the well-developed +condition of the baculum in the microtines having rooted molars is +primitive, then within the genus <i>Microtus</i> those species having +well-developed bacula may be considered primitive.</p> + +<p>The genera <i>Lagurus</i> and <i>Neofiber</i> have moderately +developed or well-developed lateral processes. <i>Neofiber</i> exhibits a +tendency, not prominent elsewhere, to have a proportionately smaller +median process rather than reduced lateral processes.</p> + +<p>American species of <i>Microtus</i> (genus and subgenus) that have +moderately- to well-developed ossified lateral processes are <i>M. +townsendii</i>, <i>M. oeconomus</i>, <i>M. pennsylvanicus</i>, <i>M. +montanus</i>, and <i>M. chrotorrhinus</i>. <i>Microtus</i> of other +subgenera having this type of baculum include <i>M. (Herpetomys) +guatemalensis</i>, <i>M. (Chilotus) oregoni</i>, and <i>M. (Chionomys) +longicaudus</i>.</p> + +<p>American species of <i>Microtus</i> (genus and subgenus) in which the +lateral ossifications are weakly developed or absent (although +cartilaginous lateral processes are present) include <i>M. mexicanus</i> +and <i>M. californicus</i>. In other subgenera, species of +<i>Microtus</i> having reduced lateral ossifications are <i>M. (Pedomys) +ochrogaster</i>, <i>M. (Pitymys) pinetorum</i>, <i>M. (Pitymys) +parvulus</i>, <i>M. (Pitymys) quasiater</i>, <i>M. (Arvicola) +richardsoni</i>, and <i>M. (Stenocranius) miurus</i>.</p> + +<p>The microtines are essentially holarctic in distribution. Both of the +tribes, the lemmings and the voles, as well as primitive representatives +of each tribe (not considering <i>Ellobius</i>) occur in both the Old +World and New World. It is not certain on which continent (or continents) +the Microtinae first differentiated. It is certain, however, that at +various times, both early and late in the evolution of the subfamily, +representatives have crossed from Eurasia to North America or <i>vice +versa</i>. Each of 10 or more microtines in the New World is more closely +related to some microtine in the Old World than to any other microtine in +the New World.</p> + +<p>The similarities or differences in the baculum in Old World and New +World representatives placed in the same genus or subgenus, or thought to +be "companion species" have been commented upon in accounts of +<i>Lemmus</i>, <i>Dicrostonyx</i>, <i>Clethrionomys</i>, <i>Lagurus</i>, +<i>Arvicola</i>, <i>Stenocranius</i>, <i>Chilotus</i>, <i>Chionomys</i>, +<i>Pitymys</i>, and in accounts of <i>Microtus agrestis</i> as compared +with <i>M. pennsylvanicus</i>, and <i>Microtus oeconomus</i> (both Old +World and New World).</p> + +<p>The baculum in the Microtinae more closely resembles the baculum in +the Cricetinae of the Old World than in the Murinae, or than in any other +rodents known to me. This resemblance suggests relationship between +Microtinae and Cricetinae.</p> + + +<p><span class="pagenum"><a name="Page_215" id="Page_215">[Pg 215]</a></span></p> + +<h2>LITERATURE CITED</h2> + +<table> + <tr> + <td colspan="2" class="smcap">Argyropulo, A. I.</td> + </tr> + <tr> + <td class="litcit_num">1933a.</td> + <td class="litcit_txt">Die Gattungen und Arten der Hamster (<i>Cricetinae</i> Murray, 1866) der Paläarktik. Zeitschr. f. +Säugetierkunde, 8:129-149, 27 figs. in text.</td> + </tr> + <tr> + <td class="litcit_num">1933b.</td> + <td class="litcit_txt">über zwei neue paläarktische Wühlmäuse. Zeitschr. f. Säugetierkunde, 8:180-183, 3 figs. in text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Callery, R.</td> + </tr> + <tr> + <td class="litcit_num">1951.</td> + <td class="litcit_txt">Development of the os genitale in the golden hamster, <i>Mesocricetus (Cricetus) auratus</i>. Jour. Mamm., +32:204-207, 1 fig. in text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Chamberlain, J. L.</td> + </tr> + <tr> + <td class="litcit_num">1954.</td> + <td class="litcit_txt">The Block Island meadow mouse, <i>Microtus provectus</i>. Jour. Mamm., 35:587-589, 2 tables in text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Dearden, L. C.</td> + </tr> + <tr> + <td class="litcit_num">1958.</td> + <td class="litcit_txt">The baculum in <i>Lagurus</i> and related microtines. Jour. Mamm., 39:541-553, 1 fig. in text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Didier, R.</td> + </tr> + <tr> + <td class="litcit_num">1943.</td> + <td class="litcit_txt">L'os pénien des Campagnols de France du Genre <i>Arvicola</i>. Mammalia, 7:74-79, 10 figs. in text.</td> + </tr> + <tr> + <td class="litcit_num">1954.</td> + <td class="litcit_txt">Etude systématique de l'os pénien des Mammifères (suite), Rongeurs: Muridés. Mammalia, 18:237-256, 14 figs. in +text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Ellerman, J. R.</td> + </tr> + <tr> + <td class="litcit_num">1941.</td> + <td class="litcit_txt">The families and genera of living rodents. Vol. II. Family Muridae. The British Museum (Natural History), London, +pp. xii + 690, 50 figs.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Friley, Charles E.</td> + </tr> + <tr> + <td class="litcit_num">1947.</td> + <td class="litcit_txt">Preparation and preservation of the baculum of mammals. Jour. Mamm., 28:395-397, 1 fig. in text.</td> + </tr> + <tr> + <td colspan="2"><span class="smcap">Hall, E. R.</span>, and <span class="smcap">E. L. Cockrum.</span></td> + </tr> + <tr> + <td class="litcit_num">1953.</td> + <td class="litcit_txt">A synopsis of the North American microtine rodents. Univ. Kansas Publ., Mus. Nat. Hist., 5:373-498, 149 figs. in +text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Hamilton, W. J., Jr.</td> + </tr> + <tr> + <td class="litcit_num">1946.</td> + <td class="litcit_txt">A study of the baculum in some North American Microtinae. Jour. Mamm., 27:378-387, 3 figs. in text.</td> + </tr> + <tr> + <td colspan="2"><span class="smcap">Hibbard, C. W.</span>, and <span class="smcap">G. C. Rinker.</span></td> + </tr> + <tr> + <td class="litcit_num">1942.</td> + <td class="litcit_txt">A new bog-lemming (Synaptomys) from Meade County, Kansas. Univ. Kansas Sci. Bull., 28:25-35, 3 figs. in text.</td> + </tr> + <tr> + <td class="litcit_num">1943.</td> + <td class="litcit_txt">A new meadow mouse (<i>Microtus ochrogaster taylori</i>) from Meade County, Kansas. Univ. Kansas Sci. +Bull., 29:255-268, 5 figs. in text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Hinton, M. A. C.</td> + </tr> + <tr> + <td class="litcit_num">1926.</td> + <td class="litcit_txt">Monograph of the voles and lemmings (Microtinae), living and extinct, Vol. I. British Museum (Natural +History), London, pp. xvi + 488, plus 15 plates, 110 figs. in text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Matthey, R.</td> + </tr> + <tr> + <td class="litcit_num">1953.</td> + <td class="litcit_txt">Les Chromosomes des Muridae. Revue Suisse de Zoologie, 60:225-283, avec les planches 1 à 4 groupant 84 +photomicrographies, 98 figures et 5 schemas dans le texte.</td> + </tr> + <tr> + <td class="litcit_num">1955. + <span class="pagenum"><a name="Page_216" id="Page_216">[Pg 216]</a></span></td> + <td class="litcit_txt"> Nouveaux documents sur les chromosomes des Muridae. Problèmes de cytologie comparée et de taxonomie chez les +Microtinae. Revue Suisse de Zoologie, 62:163-206, avec 114 figures.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Miller, G. S.</td> + </tr> + <tr> + <td class="litcit_num">1896.</td> + <td class="litcit_txt">Genera and subgenera of voles and lemmings. North American Fauna No. 12, pp. 1-85, 40 figs. and 3 plates in +text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Ognev, S. I.</td> + </tr> + <tr> + <td class="litcit_num">1948.</td> + <td class="litcit_txt">The mammals of Russia (USSR) and adjacent countries (The mammals of Eastern Europe and Northern Asia), Vol. 6. +Publ. Acad. Sci. USSR, pp. 1-587, 260 figs., 12 maps, and 11 color plates +in text (in Russian).</td> + </tr> + <tr> + <td class="litcit_num">1950.</td> + <td class="litcit_txt">The mammals of Russia (USSR) and adjacent countries (The mammals of Eastern Europe and Northern Asia), Vol. 7. +Publ. Acad. Sci. USSR, pp. 1-736, 347 figs., 15 maps, and 10 color plates +in text (in Russian).</td> + </tr> + <tr> + <td colspan="2" class="smcap">Ruth, E. B.</td> + </tr> + <tr> + <td class="litcit_num">1934.</td> + <td class="litcit_txt">The os priapi: A study in bone development. Anat. Rec., 60:231-249, 16 figs. in 3 plates.</td> + </tr> + <tr> + <td colspan="2"><span class="smcap">Smith, D. A.</span>, and <span class="smcap">J. B. Foster.</span></td> + </tr> + <tr> + <td class="litcit_num">1957.</td> + <td class="litcit_txt">Notes on the small mammals of Churchill, Manitoba. Jour. Mamm., 38:98-115, 3 figs. and 3 tables in text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Wheeler, B.</td> + </tr> + <tr> + <td class="litcit_num">1956.</td> + <td class="litcit_txt">Comparison of the Block Island "species" of <i>Microtus</i> with <i>M. pennsylvanicus</i>. Evolution, 10:176-186, +4 figs. and 2 tables in text.</td> + </tr> + <tr> + <td colspan="2" class="smcap">White, J. A.</td> + </tr> + <tr> + <td class="litcit_num">1951.</td> + <td class="litcit_txt">A practical method for mounting the bacula of small mammals. Jour. Mamm., 32:125.</td> + </tr> + <tr> + <td colspan="2" class="smcap">Zimmerman, K.</td> + </tr> + <tr> + <td class="litcit_num">1955.</td> + <td class="litcit_txt">Die Gattung <i>Arvicola</i> Lac. im System der Microtinae. Säugetierkundliche Mitteilungen, 3:110-112, 2 figs. in +text.</td> + </tr> + <tr> + <td colspan="2"><i>Transmitted August 14, 1959.</i></td> + </tr> +</table> + +<p class="center">28-774</p> + +<hr /> + + +<p><span class="pagenum"><a name="front_flap" id="front_flap">[Front Cover Inside]</a></span></p> + +<h2>UNIVERSITY OF KANSAS PUBLICATIONS<br /> +MUSEUM OF NATURAL HISTORY</h2> + +<p>Institutional libraries interested in publications exchange may obtain +this series by addressing the Exchange Librarian, University of Kansas +Library, Lawrence, Kansas. Copies for individuals, persons working in a +particular field of study, may be obtained by addressing instead the +Museum of Natural History, University of Kansas, Lawrence, Kansas. There +is no provision for sale of this series by the University Library which +meets institutional requests, or by the Museum of Natural History which +meets the requests of individuals. However, when individuals request +copies from the Museum, 25 cents should be included, for each separate +number that is 100 pages or more in length, for the purpose of defraying +the costs of wrapping and mailing.</p> + +<p>* An asterisk designates those numbers of which the Museum's supply +(not the Library's supply) is exhausted. Numbers published to date, in +this series, are as follows:</p> + + <dl> + <dt>Vol. 1.</dt> + <dd>Nos. 1-26 and index. Pp. 1-638, 1946-1950.</dd> + + <dt>*Vol. 2.</dt> + <dd>(Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948.</dd> + + <dt>Vol. 3.</dt> + <dd class="small_margin"><ol class="start_1"> + <li>*The avifauna of Micronesia, its origin, evolution, and distribution. By Rollin + H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.</li> + <li>*A quantitative study of the nocturnal migration of birds. By George H. + Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951.</li> + <li>Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-530, + 49 figures in text, 13 tables. October 10, 1951.</li> + <li>Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and + Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, + 1951.</li> + </ol></dd> + <dd>Index. Pp. 651-681.</dd> + <dt>*Vol. 4.</dt> + <dd>(Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 + figures in text. December 27, 1951.</dd> + <dt>Vol. 5.</dt> + <dd>Nos. 1-37 and index. Pp. 1-676, 1951-1953.</dd> + <dt>*Vol. 6.</dt> + <dd>(Complete) Mammals of Utah, <i>taxonomy</i> and <i>distribution</i>. By Stephen D. + Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.</dd> + <dt>Vol. 7.</dt> + <dd class="small_margin"> + <ol class="start_1"> + <li>*Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, 73 figures in text, + 37 tables. August 25, 1952.</li> + + <li>Ecology of the opossum on a natural area in northeastern Kansas. By Henry + S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. August + 24, 1953.</li> + <li>The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker. + Pp. 339-347, 1 figure in text. February 15, 1954.</li> + <li>North American jumping mice (Genus Zapus). By Phillip H. Krutzsch. Pp. + 349-472, 47 figures in text, 4 tables. April 21, 1954.</li> + <li>Mammals from Southeastern Alaska. By Rollin H. Baker and James S. + Findley. Pp. 473-477. April 21, 1954.</li> + <li>Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479-487. + April 21, 1954.</li> + <li>Subspeciation in the montane meadow mouse, Microtus montanus, in Wyoming + and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text. + July 23, 1954.</li> + <li>A new subspecies of bat (Myotis velifer) from southeastern California and + Arizona. By Terry A. Vaughan. Pp. 507-512. July 23, 1954.</li> + <li>Mammals of the San Gabriel mountains of California. By Terry A. Vaughan. + Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954.</li> + <li>A new bat (Genus Pipistrellus) from northeastern Mexico. By Rollin H. + Baker. Pp. 583-586. November 15, 1954.</li> + <li>A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp. + 587-590. November 15, 1954.</li> + <li>Geographic variation in the pocket gopher, Cratogeomys castanops, in Coahuila, + Mexico. By Robert J. Russell and Rollin H. Baker. Pp. 591-608. + March 15, 1955.</li> + <li>A new cottontail (Sylvilagus floridanus) from northeastern Mexico. By Rollin + H. Baker. Pp. 609-612. April 8, 1955.</li> + <li>Taxonomy and distribution of some American shrews. By James S. Findley. + Pp. 613-618. June 10, 1955.</li> + <li>The pigmy woodrat, Neotoma goldmani, its distribution and systematic position. + By Dennis G. Rainey and Rollin H. Baker. Pp. 619-624, 2 figures in + text. June 10, 1955.</li> + </ol></dd> + <dd>Index. Pp. 625-651.</dd> + <dt><span class="pagenum"><a name="back_flap" id="back_flap">[Back Cover Inside]</a></span> + Vol. 8.</dt> + <dd class="small_margin"> + <ol class="start_1"> + <li>Life history and ecology of the five-lined skink, Eumeces fasciatus. +By Henry S. Fitch. Pp. 1-156, 26 figures in text. September 1, 1954.</li> +<li>Myology and serology of the Avian Family Fringillidae, a taxonomic +study. By William B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables. +November 15, 1954.</li> + + <li>An ecological study of the collared lizard (Crotaphytus collaris). +By Henry S. Fitch. Pp. 213-274, 10 figures in text. February 10, +1956.</li> + + <li>A field study of the Kansas ant-eating frog, Gastrophryne olivacea. +By Henry S. Fitch. Pp. 275-306, 9 figures in text. February 10, +1956.</li> + + <li>Check-list of the birds of Kansas. By Harrison B. Tordoff. Pp. +307-359, 1 figure in text. March 10, 1956.</li> + + <li>A population study of the prairie vole (Microtus ochrogaster) in +northeastern Kansas. By Edwin P. Martin. Pp. 361-416, 19 figures in text. +April 2, 1956.</li> + + <li>Temperature responses in free-living amphibians and reptiles of +northeastern Kansas. By Henry S. Fitch. Pp. 417-476, 10 figures in text, +6 tables. June 1, 1956.</li> + + <li>Food of the crow, Corvus brachyrhynchos Brehm, in south-central +Kansas. By Dwight Platt. Pp. 477-498, 4 tables. June 8, 1956.</li> + + <li>Ecological observations on the woodrat Neotoma floridana. By Henry +S. Fitch and Dennis G. Rainey. Pp. 499-533, 3 figures in text. June 12, +1956.</li> + + <li>Eastern woodrat, Neotoma floridana; Life history and ecology. By +Dennis G. Rainey. Pp. 535-646, 12 plates, 13 figures in text. August 15, +1956.</li> + </ol></dd> + <dd>Index. Pp. 647-675.</dd> + <dt>Vol. 9.</dt> + <dd class="small_margin"> + <ol class="start_1"> + <li>Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 +figures in text. December 10, 1955.</li> + + <li>Additional records and extension of ranges of mammals from Utah. By +Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80. +December 10, 1955.</li> + + <li>A new long-eared myotis (Myotis evotis) from northeastern Mexico. By +Rollin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.</li> + + <li>Subspeciation in the meadow mouse, Microtus pennsylvanicus, in +Wyoming. By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, +1956.</li> + + <li>The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6 +figures in text. May 19, 1956.</li> + + <li>Additional remains of the multituberculate genus Eucosmodon. By +Robert W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.</li> + + <li>Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 +figures in text. June 15, 1956.</li> + + <li>Comments on the taxonomic status of Apodemus peninsulae, with +description of a new subspecies from North China. By J. Knox Jones, Jr. +Pp. 337-346, 1 figure in text, 1 table. August 15, 1956.</li> + + <li>Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp. +347-351. August 15, 1956.</li> + + <li>A new bat (Genus Leptonycteris) from Coahuila. By Howard J. Stains. +Pp. 353-356. January 21, 1957.</li> + + <li>A new species of pocket gopher (Genus Pappogeomys) from Jalisco, +Mexico. By Robert J. Russell. Pp. 357-361. January 21, 1957.</li> + + <li>Geographic variation in the pocket gopher, Thomomys bottae, in +Colorado. By Phillip M. Youngman. Pp. 363-387, 7 figures in text. +February 21, 1958.</li> + + <li>New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones, +Jr. Pp. 385-388. May 12, 1958.</li> + + <li>Pleistocene bats from San Josecito Cave, Nuevo León, México. By J. +Knox Jones, Jr. Pp. 389-396. December 19, 1958.</li> + + <li>New subspecies of the rodent Baiomys from Central America. By Robert +L. Packard. Pp. 397-404. December 19, 1958.</li> + + <li>Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. +405-414, 1 figure in text, May 20, 1959.</li> + + <li>Distribution, variation, and relationships of the montane vole, +Microtus montanus. By Sydney Anderson. Pp. 415-511, 12 figures in text, 2 +tables. August 1, 1959.</li> + + <li>Conspecificity of two pocket mice, Perognathus goldmani and P. +artus. By E. Raymond Hall and Marilyn Bailey Ogilvie. Pp. 513-518, 1 map +in text. January 14, 1960.</li> + + <li>Records of harvest mice, Reithrodontomys, from Central America, with +description of a new subspecies from Nicaragua. By Sydney Anderson and J. +Knox Jones, Jr. Pp. 519-529. January 14, 1960.</li> + + <li>Small carnivores from San Josecito Cave (Pleistocene), Nuevo León, +México. By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, +1960.</li> + + <li>Pleistocene pocket gophers from San Josecito Cave, Nuevo León, +México. By Robert J. Russell. Pp. 539-548, 1 figure in text, January 14, +1960.</li> + </ol></dd> + <dd>More numbers will appear in volume 9.</dd> + <dt><span class="pagenum"><a name="back_flap_out" id="back_flap_out">[Back Cover Outside]</a></span> + Vol. 10.</dt> + <dd class="small_margin"> + <ol class="start_1"> + <li>Studies of birds killed in nocturnal migration. By Harrison B. +Tordoff and Robert M. Mengel. Pp. 1-44, 6 figures in text, 2 tables. +September 12, 1956.</li> + + <li>Comparative breeding behavior of Ammospiza caudacuta and A. +maritima. By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December +20, 1956.</li> + + <li>The forest habitat of the University of Kansas Natural History +Reservation. By Henry S. Fitch and Ronald R. McGregor. Pp. 77-127, 2 +plates, 7 figures in text, 4 tables. December 31, 1956.</li> + + <li>Aspects of reproduction and development in the prairie vole +(Microtus ochrogaster). By Henry S. Fitch. Pp. 129-161, 8 figures in +text, 4 tables. December 19, 1957.</li> + + <li>Birds found on the Arctic slope of northern Alaska. By James W. +Bee. Pp. 163-211, plates 9-10, 1 figure in text. March 12, 1958.</li> + + <li>The wood rats of Colorado: distribution and ecology. By Robert B. +Finley, Jr. Pp. 213-552, 34 plates, 8 figures in text, 35 tables. +November 7, 1958.</li> + + <li>Home ranges and movements of the eastern cottontail in Kansas. By +Donald W. Janes. Pp. 553-572, 4 plates, 3 figures in text. May 4, +1959.</li> + + <li>Natural history of the salamander Aneides hardyi. By Richard F. +Johnston and Gerhard A. Schad. Pp. 573-585. October 8, 1959.</li> + </ol></dd> + <dd>More numbers will appear in volume 10.</dd> + <dt>Vol. 11.</dt> + <dd class="small_margin"> + <ol class="start_1"> + <li>The systematic status of the colubrid snake, Leptodeira discolor +Günther. By William E. Duellman. Pp. 1-9, 4 figures. July 14, 1958.</li> + + <li>Natural history of the six-lined racerunner, Cnemidophorus +sexlineatus. By Henry S. Fitch. Pp. 11-62, 9 figures, 9 tables. September +19, 1958.</li> + + <li>Home ranges, territories, and seasonal movements of vertebrates +of the Natural History Reservation. By Henry S. Fitch. Pp. 63-326, 6 +plates, 24 figures in text, 3 tables. December 12, 1958.</li> + + <li>A new snake of the genus Geophis from Chihuahua, Mexico. By John +M. Legler. Pp. 327-334, 2 figures in text. January 28, 1959.</li> + + <li>A new tortoise, genus Gopherus, from north-central Mexico. By +John M. Legler. Pp. 335-343. April 24, 1959.</li> + + <li>Fishes of Chautauqua, Cowley and Elk counties, Kansas. By Artie +L. Metcalf. Pp. 345-400, 2 plates, 2 figures in text, 10 tables. May 6, +1959.</li> + + <li>Fishes of the Big Blue River Basin, Kansas. By W. L. Minckley. +Pp. 401-442, 2 plates, 4 figures in text, 5 tables. May 8, 1959.</li> + + <li>Birds from Coahuila, México. By Emil K. Urban. Pp. 443-516. +August 1, 1959.</li> + + <li>Description of a new softshell turtle from the southeastern +United States. By Robert G. Webb. Pp. 517-525, 2 plates, 1 figure in +text. August 14, 1959.</li> + </ol></dd> + <dd>Another number will appear in volume 11.</dd> + <dt>Vol. 12.</dt> + <dd class="small_margin"> + <ol class="start_1"> + <li>Functional morphology of three bats: Eumops, Myotis, Macrotus. By +Terry A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, +1959.</li> + + <li>The ancestry of modern Amphibia: a review of the evidence. By +Theodore H. Eaton, Jr. Pp. 155-180, 10 figures in text. July 10, +1959.</li> + + <li>The baculum in microtine rodents. By Sydney Anderson. Pp. +181-216, 49 figures in text. February 19, 1960.</li> + </ol></dd> + <dd>More numbers will appear in volume 12.</dd> + </dl> + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of The Baculum in Microtine Rodents, by +Sydney Anderson + +*** END OF THIS PROJECT GUTENBERG EBOOK THE BACULUM IN MICROTINE RODENTS *** + +***** This file should be named 38021-h.htm or 38021-h.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/8/0/2/38021/ + +Produced by Chris Curnow, Alex Gam, Joseph Cooper and the +Online Distributed Proofreading Team at http://www.pgdp.net + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions 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. 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