MAMMALIAN SPECIES no. 125, pp. 1-4. 5 fis. Brachylagus idahoensis. _ By Jeffrey $. Green and Jerran T. Flinders Published 15 April 1980 by The American Society of Mammalogists Brachylagus Miller, 1900 Brochylagus Miller, 1900:157. Type species Lepus idahoensis ‘Merfiam, by author's designation. Proposed as subgenus, see Remarks for history of taxonomic usage. CONTEXT AND CONTENT. Order Lagomorpha, Family Leporidae. Brachylogus is one of nine living genera inthe family Brachylagus idahoensis (Merriam, 1891) Pygmy Rabbit Lepus idahoensis Merriam, 1891:76. Type locality Pahsimersi Valley, Maho (near Goldburg, Custer County, Davis, 1939 Brachylagus idahoensis Lyon, 1904-411, First usage of current name combination, CONTEXT AND CONTENT. Content given above. Bra chylagus is 8 monotypic genus (Nelaon, 1909-275). There are no “apparent subspecies of B.idahoensis (Grinnell etal, 1980; Davis, 1989; Orr, 1540), DIAGNOSIS. Body size is smaller than in any other North American leporid (Fig 1}, Hind legs are very short; ind feet are ‘comparatively broad and heavily haired. Ears are shor, rounded land densely haired inside end out and are edged with ull Vie bbnissae are black and white. Tail ie small and inconspicuous, sunderside is buff rather than white as in cottotals Sylilagus), Upper pars aze bully gray, nape and anterior surfaces of legs are ‘cinnamon buf. Rostrum is short and pointed (Fig. 2); supracrbital pacesses ate long compared with those of members of the gents Syletlagus, Postorbital extensions of supracrbitals, instead of ta pering to blunt end asin species of Syuilagus, ae broadest d Tally and this end is ether truncate ot slightly notched and elon- re. The skull is small and braincase and auditory bullae ively large. Antevior palatine foremina are broad (or ra Constricted) posteriorly: palatal bridge is short, usually with & posteromedian spine. Molarfeem teeth are relatively smalls terior surface of fist upper moleiform tooth possesses but a tingle reentrant angle: posterior halves of the second to fourth lower moleiform teeth possess lateral diameters equal to about one-half ofthe lateral diameters ofthe anterior halves. The Fidge of enamel separating individual molarform teeth into an terior and posterior sections lacks any crenulation (movie from Davie, 1989: Orr, 194) GENERAL CHARACTERISTICS. General descriptions are given in Merriam (1891), Grinnell eta. (1930), Davis (1989), (Ore (1940, Janson (1946), and Halland Kelson (1959). New pelage in Tallis Tong, almost silky, and gray on upper parts and clear ‘white, often tinged with butt. on the abdomen (Fig. 3). By mid- Wrinter, the far becomes worn and appears silver gray (Davis, 1930) "The dental formula is i 21, © 00, 8/2, m 33, total 28. ‘Weights of six adult males from California averaged 409 g (375 to-455), end nine adult females averaged 396 g (246 to 458) (Orr, 1940), In Utah six adult males averaged 405 g (873 wo 428), and four adult females averaged 486g (315 to 456) Ganson, 1946), in Tdaho adult males averaged 418 g and adalt females averaged 462 g (Wilde, 1928). The means and ranges (in parentheses) for external measurements (in mm) of six adult males from California fare: total length, 272.0 252 to 285); lengoh of tal, 17.5 (15 to 205, Teng of hind foot 69.7 (67 to 76) and length of ear feom crown, 60.8 (69 to 68). Correcponding measurements for nine adu males are: 275.3 (280 to 295) 18.1 (15 to 2471.7 (67 to 79) and 59.0 36 o 61) (Orr, 1940), Measurements of six adult male from Utah are: total length, 275 (261 to 283); Tength of tall 17 (15 to 19: length of hind foot 70 (66 to 75); and length of ear from noteh ST (4810.56). Corresponding measurements for eight adult females fare: 291 (273 to 305} 17 {15 to 19) 70 (67 to 78) and 30 (48 to 5D) Ganson, 1946) ‘Some cranial measurements (in mm) for six adult males from California are: basilar length, 38,9 (86.9 to 40.9); 2yg0 breadth, 27.3 (26:6 to 28.2) postorbital constriction, 9.2 (8.5 to 5:5 length of nasals, 19,4 (18-2 to 19,6); width of nals, 9.2 (8.4 6 10,0); length of molar series, 9.2 0.0 to 9.4; diameter of ex ternal auditory meatus, 5.1 (48:0 3,4); breadth of besinease, 202 (09-7 to 20.8); and length of palatal bridge, 4.1 (8.3 to 4.6). Cor responding measurements for nine adult females are: 38.3 (37.5 to-4L. 0); 27.3 (26.4 to 28.3); 9.5 (89 to 10,9); 18.8 (17.6 to 204), 9:7 @.1 to 10-4), 9.1 (BB to 9.0); 4.9 (4.6 to 5.4); 20.0 (196 t0 20.6); and 4.0 (5 to 4.6) (Ore, 1940). Additional measurements fare in Nelson (1909), Durrant (1952), Kenner (1965), and Wilde as} DISTRIBUTION. The geographic range of Brachylagus ldahoensis inckades most of the Great Basin and some of the adjacent intermountain areas ofthe western United States (Fig ‘, One apparently isolated population occurs in southeastern Washington, weet of Lower Coulee (Johnson et al 1951). The species s not ubiquitous over the outlined range: they are found primarily on big aagebrosh (Arcemisia ¢ridensata) dominated plans, and alluvial fans where plants occur in tall. dense clumps (Orr, 1940), Range of vertical distribution in Nevada is from about 1,810 to over 2,135 m in elevation (Nelson, 1908) and from about 11520 to 1,615 a in California (Orr, 1940, Jensen (1965) found no ehidence of B. ahoensis in his survey ofthe mammals of Rich County. Utah: however, the pygmy rebbit was collected there in 1978, ad was alan observed in southwestern Wyoming, northeast of Rich County, Ltah in 1978 by J. N. Jensen (personal com- ‘munication Stephenson (1966) noted an castern extension of the wn range of pyginy rabbits that included the upper Sevier River Valley, near Panguitch, Uteh. The communities of big sage brush that dominate many alluvial fans and riparian sites in areas near the reported range of pygmy rabbits probably supper pop- lations of these rabbits. The dense stands of big sagebrush grow= Ing adjacent to permanent and intermittent streams, along fence Tines, and in botrow ditehes next to roadways may be avenues of dispersal for these rabbits FOSSIL RECORD. Fossil remains of B. idahoensis have been found in Jaguar Cave, Lembi County, Maho, dating to late Wisconsin or early Postglacial time (earbon-I8 dates 11,580. 50 and 10,370 = 350 B.P. Kurten and Anderson, 1972) Butler (1072) stated thatthe total population of B idahoensis was greater prior to 1,000 years ago than at any time since. Changes in the Sagebrushvgrass-forb component ofthe habitat likely account for these postulated changes in numbers of pysiny rabbits, Ficune 1. follows: Lepus rownsendii, Sylidagus nusalli, and Brachylagus dahoonsis, Photograph by J.T. Finders ‘Three adult Ieporids, listed from top to bottom asFicume 2, Dorsal, ventral, and lateral views of cranium, aid lateral and occlusal views of mandible of Brachylagus udahoensis (from top: BYU 5908 female, 3906 male, 5905 female, 5909 male, and 5905 female; all from 3 mi. N- Dubois, Clark Co., Idaho: Al approx, Us. Photographs hy M. Huff FORM AND FUNCTION. This species has one molt per year in mid to late summer. Pelage changes through fall and tint are due to wear and abrasion and were duplicated on study ‘kine by scraping a dull knife over the far (Grinnell el, 1980), MAMMALIAN SPECIES 125 ‘There are 10 mamma, five on each side; two pairs are pec- toral, two pairs are abdominal, and one peit is Inguival. ‘The prostate gland of the male is usually inconspicuous, but in Utah abbite measured: 27 min long and 16mm wide in March and April Janson, 1946), ONTOGENY AND REPRODUCTION. In Idaho, testes of males began to descend in mid-December, were fully scrotal by late January, and contained copious sperm by at least March (Wilde, 1978). Tn Utah, testes size increased progressively from ‘January and peaked in March (lanson, 1946). Testicular devel- ‘opment is thought to be regulated by photoperiod and regression af testes begins in June (Wilde, 1978). Time of reproduction fx ‘apparently dependent on female readiness, whichis influenced By photoperiod and vegetative condition of the habitat (Wilde, 1078}. Gravid females were collected in Utah from late February through late March Janson, 1946), and in dah from late March through late May (Wilde et al. i976). The gestation period is lunkncwn, but is between 27 and 30 days in various species of cottontails (Sylvilagns) (Dalke, 1982; Chapman and. Harmar 1972), ‘Wilde etal (1976) found a mean of 77 corpora lutea (range, 7.108) forte female examined in Liao and etinted ap Pregnant fomales had « mean of 5.9 embryos (a on, 1946). Davis (1989) reported two females from tmbryos cach; the embryos averaged. 72 mm in. rump-crowe length, were naked, and were apparently clone t htt. An {mum of tree Itters pr yearhave been reported in Ldaho (Wilde, 1978). iis unlikely tha litters are born In the fall (Orr, 1940, “Apparently all pygmy rabbits that survive to the neat Breeding season are capable of reproduction; juveniles donot breed (Wilde, 19) ‘Sex ratios may be ¥ nificantly diferent from pendent on date of bith; earlier cohort grow larger Tonger developmental period prior to winter. The order of com: tion of growih of pygmy rabbits is length of foot, length of Pod. and weight, respectively. Mortality of adults was highest in Tate winter and early spring, wth a maximum estimated annual mortality of 88%. Juvenile mortality was highest from birth to 5 weeks of age (Wilde, 1978). ‘Although pygmy rabbits use burrow whether on the surface of underground ing females were taken, but no young were found (Janson, 1946). ECOLOGY. Pygmy rabbits are closely associated with areaations of sagebrush throughout thelr range (Merriam, 1891; Grinnell eta. 1980; Borel and Ellis, 1998; Orr, 1940). They also ‘occupy areas in Maho supporting greasewood (Sarcobamus spp.) avis, 1939), They apparently prefer tall, dense clumps of big sgebrosh (Or, 1910; Green, 1978), which may account for the fact that they have been “so easily overlooked by collectors” (Geveraid, 1950), Often extensive, wellused runways interlace the saxe thickets and provide travel and escape routes for the rabbits (Davie, 1989). Members of this species ake extensive use of burrows, largely of theirown construction, a fact that may be unique among nnalive North American leporids (Walker eta, 1968). Burrows hhave several entrances (typially four o five, but up to 10 have been noted) and are usualy located at the base of sagebrush plants lanson, 1946). However, Wilde (1978) found just two en- frances toe nost common, Burrows are usually found on slopes, find are oriented in north to east direction (Wilde, 1978) Em trance diameter of burrows ts about 10 to 12 om, Tunnels widen below the surface, forming chambers, and extend to a maximum depth of about lm Holes among volcanic rocks, in stone walls, oud abandoned buildings and thoee made by badgers Taxidea {axus) Grinnell eta. 1980) and marmots (Marmota flaiventis) (personal observation) are also used. Use of bureows is variable, but is greatest for avenile, Early reproductive activities of adults may be concentrated at burrows (Wilde, 1978). Forms under ‘egetation are used during the day ina manner similar to that of ther lagomorphs Janson, 1946) ‘Weasels (usta spp.) realy enter burrows and traps com taining pyamy rabbits (Wilde, 1978), and are the principal pred tors of B idahoenss. Coyotes (Canis fatrans), red foxes (Valpes Ealpet), owls Bubo spp.) and marsh hawks (Circus eyeneus) also -adly prey on pygmy rabbits Janson, 1986; end personal ob-MAMMALIAN SPECIES 125 FicuRe 8. Clark Co., Idaho. Photograph by J. S. Green. Adult Brachylagus idahoensis in winter pelage from servations, Bobeats (Felis rufus) Gashwiler et al, 1960) and bad- tere, as well as other predator, likely take pygmy rabbits “Janson (1946) reported the following parasites on_B. ida- Aoensis: nematodes Dermatoxysveligera and Nematodiras sp: Dotty larvae (Cuterebra sp.) fleas (Cediopsylla inaequalis): mi larvae: and ticks Dermacentor parumapertus, Haemaphysalis le= ‘porispalustris and Ornithodoros sp). Wilde (1978) reported two ‘additional flea species (Odonsopsylins dentatus and Orchopeas {exdentazus) on pygmy rabbits in Idaho, Davis (1939) often found fleas to be so abundant on pygany rabbits as to cause a wavy ‘motion in the fur. Bacon etal (1959) collected an adult Derma’ Centor anderson from 8 pygmy rabbit in Warhington, Heininths found in western species of Splllagus may also infect B. ida oenss (see Erickson, 1947. "The primary food of B. idahoensis is big sagebrush, which may comprise up to 9% of the food taken in winter. Grastes Garopyron spp. and Poa spp.) assume a more important rle (80 to 40% of the diet) in mid to late summer (Green, 1978; Wilde, 1978). This preference for big sagebrush by pygmy rabbits isnot just « function of limited availabilty of other nutritious woody ‘vegetation. Green (1978) quantified woody vegetation at six sites eastern Idaho occupied by pyemy rabbits. Antelope brush Parshia tridentata) equaled big sagebrush in percent cover (09%), and yielded 2,125 kg/ha in mean dry biomast, which com pared closely withthe 2,581 kaha mean dry biomass for big sage- brush. In this study, antelope brush never represented more than 2% mean relative frequency of occurrence in the diets of pyEmY bbits during any season ofthe year. Deposition rates of fecal pellets have been used to anseas telative population densities of pygmy rabbite (Green, 1978). Green compiled deposition rates for nine adult pygmy rabbits {our males and five females) for a nine day period tn captivity. Mean rate of defecation was 36.1 (SE = 2) pellets per hour (mean total for 24 hours was 868 pellets. Nine individual defecations of pellet groups had a mean of 67.8 (SE ~ 5.4) pellets per group. On the average, nyemy rabbits defecated 12.8 pellet groups per day. ‘There is no indication of population cycles for B.idahoensis, although in some years they seem scarcer than in others. Several, eas in Utah containing pygmy rabbits in 1981 had none in 1946 Ganson, 194). Janson (1946) reported densities of 1.75 and 3.5 rabbits per acre (0.7 and 1:4 rabbiteha, respectively) in two fx: ‘orable areas in Utah. Green (1978) reported 45 rabbits per heet- in ideal habitat in Idaho. Whether this apparent isa result of the in summer and greatest in winter (Wilde, 1978). In addition to ‘tapping, Green (1978) described techniques for capturing pyamy abbite. A noote at the end of « nylon cord attached toa cole lapsible ishing rod proved effective in taking rabbits in winter a2 they emerged from their butrows in the snow. A spotlight used At ight during the summer rendered some rabbis incapable of Kilometers N Ficune 4. Distribution of Brackylagus idahoenss (adapted from Hall and Kelson, 1959, ‘escape and allowed capture by throwing a coat oF burlap sack ‘ver them, BEHAVIOR. Brachylagus idahoensis may be active at any time; however, they are generally crepuscular (Davi, 1989; Jan ton, 1946), They have been observed feeding at midday, some times inthe tops of sagebrush plant, but they usualy rest near tor inside their burrows throughout the day Uanson, 1946). Pygmy rabbits heve alow scampering gat, "not leaping as most rabbite do,” and they easily maneuver through the brush when pursued (Merriam, 1891). They may travel approximately 1S miles per hour 24 kmh) Janson, 191). Movements generally confined toa smal area (30 m ralus) around the burrow in wie ter, with longer movements in spring and summer Janson, 1946). Homing from #25 km displacement was reported for one juvenile {female (Green and Flinders, 1979). When approached, pygmy ab- Seconds FicUnE 5. Sonagraph of a postulated alarm yocaization from Brachylagus idahoensis: A, singe syllable, and B, double sylable ‘ending with a squeak.‘ bits may move leisurely through the brush rather than retreating immediately toa burrow (Davi, 1989), Pygmy rabbits have several vocalizations, They emit a loud squeal when captured, which i typical of atherlagomorphs Ct Son, 1016: Green, 1978; Wilde, 1978). A\“pikadlike” call ‘was noted by Janson (1946) and Wide (1978), and was described by Green (1978) as an alarm call Fig. 5). Two additional vocal- Feations, a squeak end a chuckle, were identified by Green (1970 ‘Brackylogus idahoensie has heen seen inthe same areas at young cottontails and black-tailedjackrabbits (Anthony. 1913). In Idaho, they often occur in habitat slso occupied by Sylelague utallis, Lepue townsendis (personal observation), and L. cali- Jornieus (Wilde, 1978). No inerspecific encounters have been Mdeteribed, but S.nattaili and B. Ldahoensis have been observed Simultaneously inthe same burrow system (personal observation) Several pygmy rabbits may occupy the saute burrow, particulary ‘when frightened, but one adult per burrow is usual Ganson, 1946) In the breeding season, male and female adults have heen cap. tured from the same burrow (Janson, 1946; Wile, 1978; and Dersonal abservation). Dusting areas have been found in pygmy rabbit habitat, al- though no rabbits have been observed using them (Green, 1978). Some pygmy rabbit habitats in Idaho are eumpletely covered with wera feet of snow for up to two of more months during winter. Pygmy rabbits apparently make networks of subnivean trail, which are at the bases of big sagebrush shrubs, and which provide larger feeding areas. Burrows extending to the surface of the snow may or may not be constructed th these areas (personal observation). REMARKS. There has been some controversy concerning the classification of the pygmy rabbit. Merriam (1891) assigned itto the genus Lepus, based on the assumption that the pyginy bit had two molts per year, as do hares. Miller (1900) made the pygmy rabbit the type of @ new subgenus, Brachylagus. On the bass of is unique eraial characters, Lyon (1904) established Brachylagus as a monotypic genus, Nelson (1909) conearred with Lyon, Anthony (1913) and, later, Grinnell etal. (1930) confirmed that only one annual molt occurred in pygmy rabbite, Grinnell et (1930) felt that the unique cranial characters of the pygmy Tabbit merely indicated a "well marked species within the genus Shlvitagus, Spvilagus was the genus in which the pygmy rabbit ‘was placed by Orr (1940) and Hall and Kelson (1959), Based upon his analysis of dental patterns among the Le- porinae, Hibbard (1963) regarded the pygmy rabbit as bei Closely related only to Nesolagus, and stated that “itis incorrect td st leading to conndr Brackyagu aoa aubges of Sylitagus.” Following a detailed ie of cranial and dental characters, Kenner (98) an placed toe paz rab Bit in the monotypic genus Brachylagus” Johnson and. Wicks (1964) and Johnson (1968) determined that electrophoretic serum tnd hemoglobin pattern ofthe pygmy rabbit were quite diferent from the otherwise similar species of Syuilagus (S-udubont, 8. achmani, S. floridanas, and S. nutali), Based on these nd- ings. as well as out own unpublished research which shows that the pygmy 3'Syletlagus LITERATURE CITED Anthony, H.E. 1913. Mammals of northern Malheur Ci ‘Oregon, Bull. Amer, Mos. Nat. Hit, 321-27 Bacon, MC. H. Drake, and N. G, Milicr, 1959, Ticks (Ac- ‘tina: Txodoidea) on rabbits and rodents of eastern and cen- tral Washington. J. Parast, 85:281-236, Borel, A. Band R-Blis. 1998, - Mammals of the Ruby Moun- tain Region of northeastern Nevada. J. Mamm., 15:12 44, radfcld, T. 1975. On the behavior and ecology of the pyemy rabbit Syloilagae tdahoensis. MLS. Thesis, Idaho. State Univ, Pocatello 43 pp. Butler, B.'R. 1972. The Holocene and Postglacial ecological ‘erisis on the eastern Snake River plain, Tebiwa, 15:49-63. Chapman, J. 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California Publ. Zoo, 3 a Homing by a pyemy 5 Hall, E-R., and K.R. Keleon. 1959. The mammals of North ‘America, Ronald Press, New York, I:xxx + 1-546 + 79. Hibbard, C. W. 1963." The origin of the P, pattern of Si gus, Caprolagus, Oryctolanas, and Lap. J Many, 4:1~ Janson, R. G. 1946, A survey of the native rabbits of Utah ‘ith reference to their classification, distebution, life his Tories and ecology. MS. Thesis, Utsh State Univ, Logan, 108 pp. Jensen, JN, 1965, The mammals of Rich County, Utah. MS. ‘Thesis, Univ. of Utah, Salt Lake City. 131 pp. Jobnson, M. L.”1968." Application of blood protein electro- ‘Dhoretic studies to problems Zool 17.2820. Jobnsoa, MLL.,P.W. Cheney.endT-H Scheffer. 1961, Mammals ‘of Grand Coulee, Washington, Mureelet, 31:39-42 Jobngos, M: Land MJ. Wicks, 1968, Secum-protein eleo- trophoresis _in mammals: Significance in higher taxonomic categories, Pp, 681-094, in Taxonomic biochemistry and se- ry. A. Leone (ed), The Ronald Press Co., New York, 728 pp. Kenner, © H. 1965. Comparative osteology of rabbits of the genera Brachylagus Miler and Sylelagus Gray. MLS. Thee Sis, Univ. of Utah, Salt Lake City, 125 pp. Kurten, B.-and E. Anderson. 1972,”"The sediments and f of Jaguar Cave: I—the fauna. Tebiwa, 15:21—45. Lyon, MLW. Jr. 1908, “Classification of the hares and their ‘ilies. Siithsonian Mise. Coll, 45:321-447 Merriam, C. H. 1891, Resuls ofa biological reconnaissance of south-central Idaho, N. Amer. Fauna, 5:1-416. Miller, .'S. jr 1900." A new subgenus for Lepus idahoensis Poe. Bil. Soe. Washington, 13157. Nelson, E. W. 1908. “The rabbits of North America. North “Amer. Fauna, 29:1-314. Or, RT, IMO. The rabbits of California, Oceas. Papers Cal- fornia Acad, Sei, 19:1-227, Severaid, 1.11. "1950. The pyamy rabbit Spbilagus idahoen- “is ia Mono County, Californie. J. Mamm., 31:1~b Stephenson, 8. N, 1966, Mammals of the Paunsagont Plates ‘Region, Utah. Great Basin Nat, 26:43-4, Walker, B.P.s et al, 1964, The atammals of the world, Jobns Hopkins Breas, Baltimore, 2:687-1500 Wilde, D. B. 1978, "A population analysis of the pygmy rabbit (Gylelagus idakoenss) on the INEL site. Ph.D. Dissertation, Idaho State Univ, Pocatello, 172 pp Wilde, D. B., JS. Fisher, and B. L.'Keller. 1976. A demo- graphic analysis ofthe pygmy rabbit Sylilagus idahoensis, Bp. 88-105, in 1975 Progress Report Idaho National Engi- neering Laboratory Site Radinecology-Eeology Programs (0. D. Markham, ed.) U.S. Energy Res. Devel. Admin., Idaho Fall, 205 pp. Balitors for this aecouat were Dante F. WILLIAMS and SYDNEY ANDERSON Jerrney S. Gace axp Jennan , FLuNDERS, BaiotaM YouNc Univensity, Provo, Uran 34002 TTADDRESS OF GREEN, US. Sheer ExremMent Savion, Dupots, IpaHo 83425.