'310^ HARVARD UNIVERSITY Library of the Museum of Comparative Zoology The Great Basin Naturalist VOLUME 37, 1977 Editok: Stephen L. Wo Plhlished at BaiciiAM YoL'.Nc; U.\i\i:usitv, hv Bru;ham Yolnc; U.ntvehsity TABLE OF CONTEXTS Volume 37 Xuinbcr 1 - March 3L 1977 ireeding biologv of the Douhle-cresled Ooiniorant on Ltali Lake. Houald M. Mit- chell \:. 1 >attail decline at Faiinini^ton Bay Waterfowl Manau;enient Area. Christine .Mmton Anderson 24 0*' 1^ Osteology and myology of the head and neck regions of Callimunis, Cophosaurus, Holbwokia, and (7?no (Reptilia: Iguanidae). Douglas C. Cox and Wilmer W. Tanner 35 Hydrological features of a California coastal fen. Don C. Erman, Kenneth B. Roby, and Michael Eames 57 Introduced and exported American Scolytidae (Coleoptera). Stephen L. Wood 67 The sagebrvish vole flea, Megabothris ckintoni princei, in western Utah, with com- ments on the distribution of Megabothris in the Bonneville Basin. Harold J. Egoscue 75 Interpopulational variation of blood proteins in pika (Ochotona princeps saxatilis). John T. Bninson, Richard N. Seaman, and Donald J. Nash 77 Observations on the distribution of microorganisms in desert soil. A. T. Volmer, F. Au, and S. A. Bamberg 81 Home range of the western jumping mouse, Zapus princeps, in the Colorado Rocky Mountains. Nathaniel Stinson, Jr 87 Preliminary mammal survey of Fossil Butte National Monument, Wyoming. Tim W. Clark 91 An undescribed species of Thelypodiopsis (Brassicaceae) from the Uinta Basin, Utah. Stanley L. Welsh and N. Duane Atwood 95 A mountain cercocarpus population— revisited. E. S. Nixon 97 A new status for Penstenion cyananthus Hook. ssp. subglaher Pennell (Scrophula- riaceae). Stephen L. Clark 100 The influence of predator control on two adjacent wintering deer herds. Dennis D. Austin, Philip J. Urness, and Michael L. Wolfe 101 Plants of Arizona: a new species of Astragalus from the Kaibab Plateau. Stanley L. Welsh and Kaye H. Thome 103 Bionomics of two aspen bark beetles, Trypophloeus populi and Procryphahis mucro- nattis (Coleoptera: Scolytidae). Jerold L. Petty 105 Food of townsend ground squirrels on the Arid Land Ecology Reserve (Washington). MarkK. Johnson 128 Number 2 - June .30, 1977 Host-induced morphological variations in the strigeoid trematode Posthodiplostomiim minimum (Trematoda: Diplostomatidae). II. Body measurements and tegument modifications. James R. Palmieri 129 The subfamily Anomiopsyllinae (Hystrichopsyllidae: Siphonaptera). I. A revision of the genus AnomiopsyUus Baker. A. M. Barnes, V. J. Tipton, and J. A. Wildie 138 New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part iV. Stephen L. Wood 207 Supplementary note on Venezuelan bat flies (Diptera: Nycteriobiidae). Lindolpho R. Guimaraes 221 The systematics of Crotaphytus wislizeni, the leopard lizards. Part III. The leopard lizards of the Great Basin and adjoining areas, with a description of a new sub- species from the Lahontan Basin. Wilmer W. Tanner and Benjamin H. Banta .... 225 Syzygioxylon chhindwarense, a new fossil wood from the Deccan Intertrappean series of India. E. M. V. Nambudiri and William D. Tidwell 241 Birds of Gunnison and Cub islands, Great Salt Lake, Utah. F. L. Knopf and V. C. Bachman 247 Varying prevalence of Taenia taeniaeformis strobilocerci in Microtus pennsylvanicus of Montana. Richard H. McBee 252 An annotated list of the aquatic insects of southeastern Idaho, Part II: Trichoptera. Robert L. Newell and G. Wayne Minshall 253 A new status for Penstemon cyananthus Hook, ssp. subglaher Pennell (Scrophula- riaceae). Stephen L. Clark 258 Range extension of Trionyx spiniferus enioryi into Utah (Reptilia). Carlos Y. Roby and Earl C. Loveless 259 The life cycle of Hydroperla crosbyi (Plecoptera: Perlodidae). Reed Y. Oberndorfer and Kenneth W. Stewart 260 Spring waterfowl migration in the Uinta Basin of northeastern Utah. Mary E. Sang- ster 274 Number 3 - September 30, 1977 Utah flora: Brassicaceae (Cruciferae). Stanley L. Welsh and James L. Reveal 279 Typhlodromtis mcgregori Chant (Acarina: Phytoseiidae) and its control of phyto- phagous mites in Utah and southern California apple orchards. B. A. Croft and C. D. Jorgensen 366 Host-induced morphological variations in the strigeoid trematode Posthodiplostomum minimum (Trematoda: Diplostomatidae). III. Organs of attachment. James R. Palmieri 375 New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part V. Stephen L. Wood 383 Biomass of fifty conifer forests and nutrient exports associated with their harvest. T. Weaver and F. Forcella 395 The algal flora of the campus of Brigham Young University, Provo, Utah. Cheng Mou-Sheng and Samuel R. Rushforth 402 Occurrence of ticks on mule deer in central Utah. Jordan C. Pederson 407 Number 4 - December 31, 1977 Population dynamics of the Ferruginous Hawk during a prey decline. Neil D. Wof- finden and Joseph R. Murphy 411 A new species of Rhopalomyia (Diptera: Cecidomyiidae) from Artemisia ludoviciana Nuttall (Compositae) in Utah. Raymond J. Gagne 426 Bionomics of a subterranean gall midge (Diptera: Cecidomyiidae) from Artemisia lu- doviciana. M. A. S. K. Ranasinghe 429 The alpine butterflies of Castle Peak, Nevada County, California. Arthur M. Shapiro . 443 Geographic variation in Contopus sordidulus and C. virens north of Mexico. M. Ralph Browning 453 Distributional patterns of mammals in Utah. David M. Armstrong 457 A new fossil sponge from the Ordovician Garden City limestone of southeastern Idaho. J. Keith Rigby and James K. Gilland 475 Host-induced morphological variations in the strigeoid trematode Posthodiplostomum minimum (Trematoda: Diplostomatidae). IV. Organs of reproduction (ovary and testes), vitelline gland, and egg. James R. Palmieri 481 A survey of breeding and migratory birds southwest of Farmington, New Mexico. Duane A. ToUe 489 A new monotypic genus of chiggers and four new species of Quadraseta from Vene- zuela (Acari: Trombiculidae). M. L. Goff and J. M. Brennan 501 A new name for Galium scahriusculum (Rubiaceae) of Utah. Lauramay T. Dempster and Friedrich Ehrendorfer 510 New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part VI. Stephen L. Wood 511 Nectar-sugar concentrations and flower visitors in the western Great Basin. Larry J. Gut, Robert A. Schlising, and Carol E. Stopher 523 New grass distribution records for New Mexico and the United States. Stephan L. Hatch 530 Stilt Sandpiper and Hudsonian Godwit in Nevada. C. S. Lawson 532 Index to Volume 37 533 HE GREAT BASIN NATURALIST )lume 37 No. 1 March 31, 1977 Brigham Young University MUS. COMP. ZOOU. LIBRARY JAN20^b/o HARVARD UNIVERSITY GREAT BASIN NATURALIST Editor. Stephen L. Wood, Life Science Museum, Brigham Young University, Provo, Utah 84602. Assistant Editor. Stanley L. Welsh, Life Science Museum, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; William D. Tidwell, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agri- cultural Sciences; Ernest L. Olson, Director, Brigham Young University Press, Univer- sity Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been pub- lished from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only previously unpublished manuscripts of less than 100 printed pages in length and pertaining to the biological natural history of western North America are ac- cepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manu- scripts are subject to the approval of the editor. Subscriptions. The annual subscription rate of the Great Basin Naturalist is $12 (outside the United States the rate is $13). The price for single numbers is $4 each. All back num- bers are in print and are available for sale. All matters pertaining to the purchase of sub- scriptions and back numbers should be directed to Brigham Young University Press, Mar- keting Department, 204 UPB, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same office at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either jour- nal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts or other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back cover. © 1977 Brigham Young University Press. All rights reserved Printed in United States of America 9-77 650 24964 The Great Basin Naturalist PuBLisHi:i) AT Pro\o, Utah, by Brigham Young University Volume 37 March 31, 1977 No. 1 BREEDING BIOLOGY OF THE DOUBLE-CRESTED CORMORANT ON UTAH LAKE' Ronald M. Mitchell- Abstract.— Research on the nesting of the Donble-crested Cormorant (Phalacrocorax auritus auritus) was un- dertaken to determine the breeding biology of the bird on Utah Lake, Utah County, and the possible effects that diking Provo Bay would have on the cormorants breeding there. In 1973, two colonies in Provo Bay were visited weekly to determine laying dates, incubation period, clutch size, and hatching dates. Another colony, located on the dike of the Geneva Steel Works Reservoir near Oreni, was visited daily during the spring and summer of 1973. At this colony egg-laying occurred from 13 April to 17 May. The average clutch size was 3.8, and the egg length and width averaged 60.50 mm x 39.05 mm. The average period of incubation was 28 days, and 29.5 per- cent of the eggs hatched, beginning on 20 May and ending on 9 June. The young weighed an average of 36 gm at hatching with a beak length of 17 mm and tarsometatarsal length of 11 mm. These measurements increased to 1543 gm, 73 mm, and 58 mm, respectively, in 23 days. The cormorant population on Utah Lake may be severely reduced by the proposed diking of Provo Bay. There have been no detailed studie.s of the biology of the Double-crested Cormo- rant {Phalacrocorax auritus auritus) in Utah, other than a brief but noteworthy account by Behle (1958) of those birds that nested on Egg Island, Great Salt Lake. Two de- tailed studies of this subspecies have been conducted on the Atlantic coast (Lewis 1929, Mendall 1936), but no extensive ac- counts of breeding biology exist for Double- crested Cormorants found nesting inland. This lack of scientific data may be due in part to a lack of qualified and interested ob- .servers, but I feel it is principally due to the sentiment expressed in the following quotation:"God no doubt had his reasons for creating each living thing, but when he cre- ated the cormorant he did himself little credit" (Demille 1926). This attitude may account for the fact that the cormorant is likely the least understood water bird nest- ing in Utah. Although there are three other subspecies (P. a. albociliatus, P. a. floridanus, P. a. cin- cinatus) found in North America, P. a. au- ritus is considered the most important due to its wide distribution, ranging from central Canada to Texas and from the Atlantic coast to Utah. The P. a. auritus populations in Utah are the westernmost breeding colo- nies found in the United States and provide excellent contrast for compari.son with the previously studied birds nesting along the Atlantic coast. The few publi.shed references that exist concerning Utah's cormorants are little more than sight records or brief mentions of nesting sites. Behle (1958) summarized the history of the seven cormorant colonies of which he was aware. At the time of his This manuscripl was prepared in partial fulfillment of the requirements for the degree of Master of Science in Zoolojjy at Brighi Provo, L'tah. The research was supported in part by a grant from the National Audubon .ScK-iety. Young University. PO. Box 28147, San Ant( Great Basin Naturalist Vol. 37, No. 1 writing, all but two (Egg Island and Bear River) had been deserted or destroyed (Fig. 1). In addition I found a reference to an un- reported colony located on White Rock, a small island located a few hundred yards northwest of Antelope Island, Great Salt Lake. A set of cormorant eggs was collected from there on 10 May 1901 (Woodbury et al. 1949). I also discovered an unreported colony located west of Bountiful in 1973. Behle (1958) did not mention Utah Lake as a nesting area, and it is mentioned only once in the other literature as a "former" nesting site (Woodbury et al. 1949). How- ever, I was able to find unpublished refer- ences that included a record of four cormo- rant eggs collected in Provo Bay in 1920 (Bee 1920). Cottam (1927) stated that the Double-crested Cormorant nested on Utah Lake. In 1937 John Hutchings of Lehi (Bee 1937) reported that several sets of Double- crested Cormorant eggs were collected on Rock Island in Utah Lake (Fig. 2). James Bee (1946) found six active cormorant nests in a tree on the Geneva dike in May 1946 and stated that the birds had nested there for two or three years. Bee (1949) reported V/Be« r^ A Logan / r ^ p^ Brigham ^ Great \ 1 Salt a\ Y*"^^ Bou„t..uX UTAH ^-N,_y salt Lake ■Rjoele 9- vernal Nephi price Clear LaKe ij » Fi Cedar Ci St. Ge9rge ty Moab A Deserte • Present <. or destroyed coloni a trip to a large cormorant colony in Provo Bay, south of the Provo airport dike, that contained 150 pairs of nesting cormorants. The purposes of this study were to obtain information on the breeding biology of the unreported colonies on Utah Lake, obtain data on the nesting ecology of inland nest- ing cormorants for comparison with the data already published concerning coastal nesting birds, determine the current status of Utah's cormorant population, and gather information to determine the impact on the nesting sites of the two colonies located in Provo Bay by the proposed plan of the Bu- reau of Reclamation to dike and drain the bay as part of the Central Utah Project. The Study Area of There are three nesting colonies Double-crested Cormorants located near Utah Lake. Two of the colonies are located in Provo Bay (Fig. 2), a shallow arm on the eastern side of Utah Lake. These have been designated the Springville and Bay colonies. ■ Springville Colo ^ Bay Colony ^ Geneva colony Fig. 1. Location of the deserted and active nesting Fig. 2. Location of the three study sites on Utah colonies in Utah. Lake. March 1977 Mitchell: Cormorant Biology The water depth in this area varies from a few milhmeters to 2 m and averages about 1 m. The principle vegetative types are hardstem buHrush (Scirpus acutiis), tamarix (Tamarix petandra), cottonwood {Populus fremonti), and willow (Salix amygdaloides). The third colony is located 11 km north of Provo Bay at the United States Steel Com- pany Geneva Works and is designated Gen- eva Colony (Fig. 2). The Springville Colony is located 0.5 km due west of the mouth of Hobble Creek in a stand of cottonwood trees that range from 10-20 m in height and extend in a straight line west from the shore into the bay. The tree line, which is about 0.5 km long, was established in the late 1930's during low lake levels. The westernmost trees are dead, in 1 m of water, and in 1971 and 1972 the trees most distant from shore were used by cormorants for nesting. In 1973, apparently due to increased numbers, the birds moved eastward into some living cottonwoods that had previously been used exclusively by Great Blue Herons {Ardea herodias) for nesting. The Bay Colony is located at the mouth of Provo Bay (Fig. 2), 2 km due south of the Provo airport. This site consists of a stand of cottonwoods and willows growing in an east-west line in about 0.6 m of wa- ter. In 1971 the area consisted of 18 trees ranging from 10-20 m in height. In 1973 cormorants nested with Great Blue Herons in one cottonwood and in four willow trees at the east end of the tree line. The Geneva Colony is located on a diver- sion dike, built in 1940, that extends about 1.2 km within the reservoir. This artificial pond is located west of the steel plant proper and is about 0.2 km from Utah Lake. The water is contained by 3 m high earthen walls (Fig. 3). The water, which is over 6 m deep in most places, is used for cooling purposes in the steel-making pro- cess. The cormorants nested in one cotton- wood and three Siberian elms {Ulmiis pii- mila), all living, at the distal end of the dike. Great Blue Herons, Snowy Egrets {Leucophoyx thula), and Black-crowned Night Herons (Nycticorax nycticorax) nested in the same trees, while thousands of Cali- fornia Gulls {Larus californicus) nested on the dike itself. Methods and Procedures The study began in the spring of 1972 and continued until the spring of 1974. Two blinds were set up at the Geneva Colony to allow observation of nesting ac- tivity. Both blinds were set up on the east bank of the Geneva pond. One blind was placed opposite trees 1 and 2 about 12 m distant (Fig. 3). Six cormorant nests were visible from this blind. Tree 3, which con- tained three cormorant nests, was also ob- servable when binoculars were used. The second blind was placed about 300 m far- ther north near the northeast corner of the pond. Tree 4 was about 30 m away, and nine cormorant nests were observable when binoculars or a spottingscope were used. All trees were mapped, and each nest was assigned an identification number. In order to determine reuse of old nests and the number of new nests built, the nests that remained intact from the previous year Primary Dike (1.2 km) Fig. 3. Map of Geneva reservoir showing the loca- tion of the bhnds and the trees containing cormorant nests. Great Basin Naturalist Vol. 37, No. 1 were mapped before the birds returned. Ob- servations of the eggs and young in the nests were made using four ahiminum poles, 1.5 cm in diameter and 1.6 m long, which could be inserted into one another and se- cured by a wing nut. A side-view mirror at- tached to the end pole afforded a view of the nest. When held at chest level, this as- semblage allowed visual access to most nests. Tape was placed at 0.2 m intervals on the poles to measure nest height from ground or water level. One new and one rebuilt nest were col- lected from the Springville Colony for fau- nal examination in Berlese funnels and for comparative measurements of numbers, av- erage thickness, and length of each piece of nesting material. Invertebrates found in the nests were collected for identification. Egg length and width were measured us- ing dial calipers accurate to one-twentieth (.05) of a millimeter. Measurements were made to compare eggs of the same clutch, eggs within the colony, and eggs from dif- ferent colonies. Egg volume was determined using the formula V = 0.526 LB^, where L = longitudinal axis and B = transverse axis (Romanoff and Romanoff 1949). Average clutch size was determined for each colony by including abandoned nests, but not unused nests. A nest was considered abandoned if it contained at least one egg at any time, but showed no signs of in- cubation for one week. Nests that never contained an egg were considered unused. Hatching success for each breeding pair was determined by dividing the number of hatchlings that survived one day by the to- tal number of eggs laid in the nest, and for the colony by the total number of hatch- lings that survived one day by the total number of eggs laid in the colony. Fledgling success was determined for the Geneva Colony by dividing the total num- ber of fledglings that left the nest by the number of hatchlings that survived one day. Three measurements— weight, length of bill, and tarsometatarsus— were made daily of the Geneva hatchlings to determine growth rates. A number indicating nest and order of hatching was written on tape and attached to the leg of each bird for identi- fication. Weight was determined using three Sargent- Welch dial spring-balances: a 250 gm balance accurate to 2 gm, a 500 gm balance to 5 gm, and a 2000 gm balance to 10 gm. A small linen bag was attached to the balance hook to hold the bird for weight readings. The bill and tarsometa- tarsus lengths were measured with a clear plastic 150 mm rule. The bill was measured from the corner of the mouth to the tip of the upper mandible. The tarsometatarsus was measured from the point of articulation below the ankle to its articulation with the middle toe. In 1973 trips were made to all recorded nesting sites in the state to determine which colonies still existed and to conduct a cen- sus of Utah's cormorant population. The colony at Bear River Migratory Bird Refuge was visited in order to photograph the use of an alternative nesting habitat and to de- termine location and size. Letters were also sent to managers of selected state bird ref- uges in an attempt to discover new breed- ing colonies. Results and Discussion Breeding colonies of cormorants in Utah have been reported in Cache, Rich, Millard, Box Elder, and Davis counties. The only published reference to breeding in Utah County was a brief mention of the area as a "former" nesting site (Woodbury et al. 1949). This paper, then, is the first detailed report of the nesting of the cormorant on Utah Lake, Utah County. On 20 June 1973 I visited the Newstate Gun Club located 4 km west of Bountiful, Davis County, Utah. An unreported cormo- rant breeding colony was located with nests in Russian olive (Elaeagnus angustifolia) trees along the south side of the Burton Canal. About 15 trees, averaging 8 m high, were scattered for 1 km along the canal. They contained 42 active nests with 66 well-developed young. In addition, 23 im- mature birds were seen swimming in the canal. The trees containing cormorant nests also held a total of 61 Great Blue Heron nests with young. The caretaker of the March 1977 Mitchell: Cormorant Biology property reported that the cormorants had nested there since 1962. Arrival at Nesting Sites.— In 1972 the first birds to return to the Springville Colo- ny were seen on 4 March. Cormorants were first seen at the Bay Colony on 13 March and at the Geneva Colony on 22 March. In 1973 the first cormorants were seen at the Springville Colony on 7 March and on 14 March at the Bay Colony. Three birds, seen on 26 March, were the first to arrive at the Geneva Colony. Lockerbie (1942) stated that over a peri- od of years the earliest observation dates for cormorants in Utah was 29 March. The earliest that cormorants have been seen at Bear River Refuge is 5 March (Behle 1958). The arrival of cormorants on 4 March 1972 at Springville during this study is the earliest record for Utah. In 1972 and 1973 the first returning birds encountered inclement weather conditions, and some ice remained on Utah Lake. It is interesting to note that the first birds re- turned to the Bay Colony a week later than the Springville birds. Yet, all of the birds that nested at the Bay and Geneva colonies returned within four weeks, while birds were still arriving at the Springville Colony and building nests in late May. Since the first cormorants to return to Springville are often subjected to extremely poor weather conditions, their early arrival must provide them with some adaptive advantage. It may be that since it is the largest colony, nesting space is at a premium and the first birds to return are assured of the choicest areas. I observed that those birds that did not return to Springville until May in 1973 were forced to build their nests west of the main cormorant colony. The reason for the time difference in re- turn to each colony is puzzling. The Bay Colony is subjected to essentially the same type of weather conditions as Springville. Geneva may have somewhat different weather, but the pond is always ice-free so the birds could obtain food at any time. The Bay and Geneva colonies are consi- derably smaller in numbers of nesting birds than the Springville Colony, so the lack of competition for nesting space may not re- quire an early return. It may also be that the birds in each col- ony remain associated at their wintering grounds and return as a group, or that the individual birds are site specific. Color- banding fledglings to determine if they re- turn to the same colony each year may help answer this question. Nest Construction.— The Double-crested Cormorant utilizes two different nesting sit- uations determined by local conditions. Of the thirteen colonies reported in this paper, seven were ground-nesting colonies and of these only the Bear River Colony is still in existence. Vermeer (1973) stated that cor- morants prefer to nest on the ground and nested in trees only after their ground nest- ing habitat was disturbed. However, this is not always the case, since the birds at Ge- neva could have nested undisturbed on the dike in association with California gulls, but instead nested in trees. Most cormorants that nest along the At- lantic coast nest at ground level on islands bare of vegetation (Bent 1922; Lewis 1929; Mendall 1936). Townsend (Bent 1922) found that these island nests consisted of seaweed, kelp, grass, and a few sticks. The nest di- mensions averaged 52 cm across by 12 cm deep. Unlike the flimsy nests described by Townsend (Bent 1922), Lewis (1929), and Mendall (1936), Behle (1958) found that the ground-nesting cormorants at Egg Island built elaborate interwoven nests of sage and greasewood twigs. The twigs averaged 12 to 18 mm thick and 290 to 580 mm long. The nests were built on or near rocks and some of the nest cups were lined with gull feath- ers. Nests that had been used for more than one season were almost cemented solid with excrement. Cormorants did not nest on the islands at Bear River Refuge until three years after the islands were built (Marshall 1937). At that time rocks were placed at each end of the islands and cormorants then used them for nesting. Yet, Behle (1958), Lewis (1929), Mendall (1936), and Munro (1927) report nesting on bare ground in comparable island situations. Nest Utilization.— Of the 83 cormorant 6 Great Basin Naturalist Vol. 37, No. 1 nests located in the Springville Colony in 1973, 14 (17 percent) of the nests left from the previous year were utilized by the first returning birds. In the entire colony only five nests were destroyed and two aban- doned. The height of the nests averaged 5.2 m (range 2.4 to 7.6 m) above the surface of the water. The nests per tree varied from 1 to 11. There did not seem to be any corre- lation between the size of the tree and the number of nests the tree contained. Nests were still being constructed in the eastern part of the colony during late May. Of the seven cormorant nests studied at the Bay Colony during the nesting season, none were lost, although two nests were abandoned. The nest height averaged 5.3 m (range 4.3 to 6.4 m). Nests per tree varied from 1 to 3. All nest-building ceased by early April. In 1972 cormorants nested in four trees on the dike at the Geneva Colony (Table 1). Trees 2, 3, and 4 were all being utilized by 1 April, but tree 1 was not used for nesting until about 10 days later (Fig. 3). Eighteen nests from the previous year were reused, although some were in very poor condition. One old nest in tree 1 was not used, although it seemed in good condition. No nests were abandoned. One nest each in trees 2 and 4 were lost when they fell out of the trees during a storm. The same trees used for nesting at Ge- neva in 1972 were occupied in 1973. How- ever, tree 1 was utilized first in 1973, fol- lowed by trees 2 and 4, and finally by tree 3. Twenty-four nests remained of the 33 used in 1972. Some of these were in very poor condition and only 11 (33 percent) of Table 1. Summary of nest utilization at the Geneva Colony indicating numbers of new and rebuilt nests in each tree used for nesting. The number in parentheses indicates the percent of total colony nests each tree contained. Tree 1 Tree 2 Tree 3 Tree i4 Year new rebuilt i new rebuilt i new rebuilt new rebuilt 1972 1973 0 0 4(12) 3(18) 3 6(27) 2 0(12) 3 2 3(18) 1(18) 9 2 5(43) 7(52) them were reused for nesting. Nest height averaged 4.7 m (range from 3 to 7 m). Three unused nests were destroyed during bad weather and another was lost due to rob- bing of material by cormorants. Two nests that contained eggs were abandoned before hatching. During both years nest construc- tion was finished by early May. The reuse of old nests requires a com- ment. Palmer (1962) stated that rebuilding a nest took only two days, compared to four to six days to build a new one. Rebuilt nests were also reported to be much stronger than new nests. Rebuilding generally con- sisted of adding some material to the top of the old nest. He gave one record of a nest being reused for four years. I know of four nests at Geneva which were reused for at least three consecutive years. At all of the colonies there appeared to be a limit to the proximity of nests, since, even when there was a suitable site for an- other nest, none was built closer than about 1 m to another. The average distance be- tween nests at the Geneva Colony was 1.8 m. Lewis (1929) described inland nests in Ar- kansas, Minnesota, and Quebec that were located in various species of trees growing on islands or standing in water. The nests were located 0.3 to 22 m from the surface of the ground or water. He reported that the nest dimensions averaged 22 x 48 cm and that there were never more than six nests in one tree. However, Palmer (1962) observed that trees have contained as many as 36 nests and I observed a tree with 11 nests at Springville. The nests consisted of dead twigs, devoid of bark, and the nest cup was often lined with green leaves or grass and gull or cormorant feathers. I observed that dead twigs from various species of trees were the primary materials used in nest building. However, large pieces of hardstem bullrush were often used. I was not able to identify the tree species used, but I often saw cormorants pick up dead twigs from the ground beneath the same trees they nested in. Large twigs were se- cured in the crotch of branches and then other twigs were interwoven until the nest was approximately 30 cm high and 40 cm March 1977 Mitchell: Cormorant Biology wide. Reused nests tend to increase in size each year, due to the addition of material. The nest cup was hned with finer materials, including leaves and cormorant feathers. Occasionally, parts of live and dead bull- rushes were also used to line the nest cup. One new and one rebuilt nest were collect- ed for comparison of building materials. The new nest had materials averaging 312 mm in length (range 150 to 610 mm) and 7.8 mm thick, the thickest being 26.0 mm. The rebuilt nest had materials averaging 333 mm in length (range 170 to 690 mm) and 8.1 mm thick, the thickest being 25.0 mm. Prior to measuring, the nests were placed in Berlese funnels to collect invertebrates. A beetle {Denneste.s lardarius) and several flies {Rhegoclema sp.) were identified. These in- vertebrates should not be considered a de- finitive list of organisms that are found in company with cormorants. They are only examples of what may be found, as I have encountered no other list of this type in the literature. From my observations in 1972 I con- cluded that old nests, in good condition, were the preferred nesting habitat of re- turning cormorants. There was an order of preference from those highest in a particu- lar tree to the lowest. Only after these were occupied were new nests built. Despite this general rule, at Geneva in 1973 new nests were built while old nests were left unoccu- pied. There is evidence that the most mature birds arrive at a colony and begin nesting first (Lack 1968, Palmer 1962). If that were true, the higher nests in a colony may have a better hatching success, since the older birds should be more prolific and make few- er nesting errors (Lack 1968, Lewis 1929). To test this, I took the average of all nest heights for each colony and determined the hatching success for the nests situated above the average height as compared to those be- low the average height. In every instance, the lower nests had a higher hatching suc- cess (Fig. 4). This may indicate that the first returning birds are not necessarily the most mature birds, or possibly that the higher nests, which are more exposed to the ele- 7,5 -1 r 7.0 - 6.5 67* ' r 6.0 19% 39% 5.5 . I 5.0 - 62* 4.0 69« 36% 3.5 3.0 - 2.5 . - 2.0 Fig. 4. Distribution of nest height showing hatching success above and below the average nest height. ments when the adults are disturbed (not shielded by leaves or branches), have a higher incidence of pre-hatching mortality. However, more research is needed since neither of these ideas seems a completely adequate explanation. In Utah it seems to be the rule that cor- morants nest in the same trees with Great Blue Herons. Presently in Utah only the cormorants at Bear River do not nest in company with herons. Former nesting colo- nies in Utah where herons nested with cor- morants were located at Bass Pond (Pearson 1927), Cache Valley (Stanford 1937), and Egg Island (Behle 1958). However, this seems to be an uncommon occurrence throughout most of the cormorants' range, since there are few other references to them nesting with Great Blue Herons. Howell (1911), Stoddard (1922), and Vermeer (1973) reported them nesting with herons in Il- linois, Wisconsin, and Manitoba, respective- Iv. Ganier (1933) described cormorants shar- Great Basin Naturalist Vol. 37, No. 1 ing a colony with Great Blue Herons, Water Turkeys, and American Egrets in Tennessee. Cormorants have also been re- ported nesting with Great Blue Herons and Black-crowned Night Herons in Nebraska (Lewis 1931). I have observed that when cormorants nest in the same trees with herons, the cor- morant nests are always located below the level of the heron nests. Stanford (1937) noted the same thing at the colony in Cache Valley. At Springville and Geneva, where only one heron was nesting in a tree occupied by cormorants, the heron's nest was the topmost in that tree. When heron nests were located at the same level as a cormorant nest, the heron nests were never situated closer than 3 m to it. At Geneva in 1972 the highest nest in tree 1 was utilized by a Great Blue Heron. In 1973 the same nest was used by a cormorant, later desert- ed, and again used by a heron during the same season. Although there is no explanation in the literature for partitioning of the nesting habitat by these two species, I feel this con- dition may exist for two reasons. Cormo- rants are noted for the large amount of ex- crement they deposit outside the nest, to the point that the very trees they nest in are often killed (Lewis 1929; Mitchell pers. obser.). The feces could also be harmful if they were deposited on heron eggs or young located beneath a cormorant nest. However, the most probable reason is the difference in take-off and landing patterns of the two birds. The cormorant is not an agile bird and, in order to become air- borne, must launch itself precipitously out of the nest (Hall 1926). This may result in eggs or young being kicked out of the nest. Likewise, when they land at the nest, they will sometimes miss their footing and fall out of the tree or crash into their nest-mate or other cormorants nesting nearby. In com- parison, the Great Blue Heron is able to fly away from the nest with considerable grace by launching itself straight into the air, or land at the nest with amazing agility so as to not disturb any of the nesting birds lo- cated nearby. Therefore, I feel that the stratification of the nests of these two spe- cies is a behavioral adaptation for partition- ing the nesting habitat to avoid interspecific collisions, which could result in the loss of eggs and young. Further research may answer important questions related to nesting, such as whether cormorants return to the same colony each year or even the same tree and nest. It is not known whether tree-nesting cormorants could effectively become ground nesters. An attempt to induce the ground-nesting cor- morants at Bear River Refuge to nest in trees was successful when old nests were placed in nest boxes in an artificial tree made of 2 x 4's on the island in unit no. 4 (Gunther, pers. comm.). It also needs to be determined whether cormorants could be enticed to a new nesting area by taking old nests from a former colony and placing them in a new location. Egg Laying and Incubation.— In 1973 egg laying began at the Springville Colony on 26 March and continued into late May. Egg laying began at the Bay Colony about 6 April and continued into early May, and at the Geneva Colony began on 13 April and continued until 17 May. The difference in the beginning egg lay- ing dates at each colony is a reflection of the time differential in early arrivals at each colony. There is a three-week period be- tween the date of first arrivals and the date that eggs are first laid, so that eggs are laid first at Springville and last at Geneva. The eggs laid on 26 March 1973 at Springville comprise the earliest record for Utah (Behle 1958), although Lewis (1929) stated that Utah had the earliest laying dates for any state and theorized that laying began during the second half of March. Five records from Utah, given by Townsend (Bent 1922), provided dates of 9 April to 17 May for egg laying. The earliest reported date for eggs laid in Utah was 28 March 1937 at Egg Island, Great Salt Lake (Behle 1958). Marshall (1937) reported a very late egg laying date at Bear River Refuge, where nests contained viable eggs on 2 Sep- tember 1936. Eggs were laid over a 12-week period at Springville and over more than a 4-week period at Geneva (Fig. 5), although the bulk March 1977 Mitchell: Cormorant Biology 14 16 18 20 22 24 26 28 30 2 4 6 April DATE 10 12 14 16 18 20 22 May Fig. 5. Distribution of egg laying at Geneva Colony from 14 April to 20 May 1973. of laying at Geneva came within a 2- week period. Vermeer (1969) reported egg laying lasted for a period of 2.5 months in Alberta, Canada. At Geneva 90 percent of the females laid one egg per day until the clutch was complete. The other females took more than a day to lay one or more of their eggs. Others have noted that females laid one egg per day until the clutch was complete (Behle 1958; Lewis 1929; Mendall 1936), although Lack (1968) stated, that cor- morants lay eggs every two to three days. Lewis (1929) and Palmer (1962) reported that incubation usually began after depo- sition of the third egg. Mendall (1926) and Behle (1958) observed that incubation may begin with the first egg, or at any time thereafter, until the last egg is laid. I found that incubation began immediately after the first egg was laid, which may be due to egg laying beginning early in Utah while the weather is still very cool. The incubation periods given by Behle (1958), Lewis (1929), and Mendall (1936) were 25 days. Bergtold (1917) reported that most North American cormorants averaged 28 days for incubation, and Worth (1940) gave the incubation period for the Eu- ropean Cormorant (Phalacrocorax carbo), though somewhat larger in size than P. a. auritus, as 28.5 days. Van Tets (1959), working with the svibspecies P. a. cincinatiis in British Columbia, had 27 records for in- cubation, varying from 25 to 29 days, with most eggs hatching after 28 days. For the 16 Geneva hatchlings in 1973 incubation varied from 26 to 30 days with an average of 28 days for incubation. The difference between the dates reported by others and my results may again be due to early nest- ing and cooler weather. At Geneva in 1972, 11 nests lost one or more eggs before hatching. All of the eggs from four of the nests were lost before 10 May and a replacement clutch was laid for each by 29 May. The replacement clutch consisted of the same number of eggs as lost in three of the nests and one more egg than lost in the other. In 1973, 12 nests lost one or more eggs prior to hatching. However, of the seven nests that eventually lost all eggs, no clutch was completely lost before 24 May, and no new clutches were laid. Renesting occurred in 30 to 50 percent of 10 Great Basin Naturalist Vol. 37, No. 1 cormorants in Manitoba whose eggs or nests were destroyed (McLeod and Bondar 1953). Double-crested Cormorants have never been reported to raise two broods in a single year. It appears that as long as one of the original eggs remains in the clutch, there is no replacement of lost eggs. Also, there is apparently a period during which the fe- male can receive a stimulus that will enable her to lay a new set of eggs. After that time (10 May to 24 May in Utah), she is unable to replace a lost ckitch. Clutch Size.— The normal clutch size of the Double-crested Cormorant is three or four eggs, with four eggs most common (Bent 1922; Davie 1900; Lewis 1929; Men- dall 1936). Palmer (1962) reported a range of two to seven eggs with one record of nine eggs. Lewis (1929) found that most cormorants along the Gulf of St. Lawrence had a clutch of four eggs, "rarely" five eggs, and 5-10 percent contained three eggs. Mendall (1936) reported the following per- centages for Maine: two eggs, 8 percent; three eggs, 40 percent; four eggs, 50 per- cent; five eggs, 2 percent. In a census taken on Egg Island, Great Salt Lake, in 1941, Behle (1958) found that over half the nests contained four eggs. I censused eggs weekly in the Springville and Bay colonies and daily in the Geneva Colony; then the maximum number of eggs present was counted before hatching began. A total of 126 eggs were found in the Springville Colony (average 3.8 eggs per nest), and 28 eggs in the Bay Colony (aver- age 4.0 eggs per nest) in 1973. At the Gen- eva Colony in 1972 a total of 80 eggs were found (average 4.2 eggs per nest), and in 1973, 61 eggs (3.6 average) were found. The average for all the colonies in 1972 and 1973 was 3.8 eggs per nest. Table 2 in- dicates the combined totals and percentages of all the clutches from the Utah Lake colo- nies. These figures agree with other find- ings, although there is a greater percentage of clutches containing four and five eggs than previously reported. £gg Measurements.— Measurements of the longitudinal and transverse axes of the cor- morant eggs indicated that the Bay Colony eggs had the smallest average length and Table 2. Numbers of eggs laid per nest at the Utah Lake colonies. The percent of the totals are in paren- theses. Year Number of eggs /nest Colony 2 3 4 5 6 Geneva Springville Bay Geneva 1972 1973 1973 1973 0 2 0 1 3 6 2 5 10 21 3 11 5 4 2 0 1 0 0 0 Totals ■3(4) 16(21) 45 (59) 11(14) 1(1) greatest average width of 59.7 x 39.0 (longi- tudinal X transverse in mm), while the Springville eggs showed the greatest aver- age length and smallest average width of 61.35 X 38.5. The Geneva averages were in- termediate in size at 60.43 x 38.99. Small variations from these measurements can be found in the literature as indicated in Table 3, but they are remarkably slight when the large geographic distances separating the samples are considered. In a discussion on various means of meas- uring eggs, Preston (1968) stated that meas- uring all the eggs in a colony produced a bias toward the larger clutches. If measure- ments are being made in order to establish an average or norm for a species, it is the number of clutches or sets of eggs, not the number of eggs, that is important. He thought that a random selection of one egg from each of 20 or more clutches would be a better indicator for the species. In order to check that hypothesis, I took an average of all the eggs measured at Geneva in 1972 and in 1973. I then compared those aver- ages with three random samples of one egg measured from each of 12 clutches during each year (Table 4). T-tests (two-tailed) at 5 percent indicated that the random means were not significantly diff^erent from the overall mean. However, since Preston rec- ommends that the comparison be done with 20 or more clutches, significant differences may appear in larger samples. Egg Volumes.— Volumes for the eggs measured at Geneva in 1972 and 1973 were calculated in order to establish an average March 1977 Mitchell: Cormorant Biology 11 Table 3. Egg measurements from various sources compared with those of the Utah Lake colonies, showing range in length and width measurements with averages in parentheses. Date Location Sample Size Length (mm) Width (mm) Source - U.S. Nat. Museum" Canada Maine Egg Island, Utah Total Bay Springville Geneva Springville Geneva Total 40 50 15 20 125 7 7 48 10 52 124 56.00-65.42(61.60) 54.50-66.50 (59.90) 36.50-42.00 (38.80) 34.50-42.00 (37.70) -- (37.60) 34.80-39.40(37.10) Bent 1922 Lewis 1929 Mendall 1936 Behle 1958 - 53.00-64.00 (59.30) 4/16/71 4/27/71 5/18/72 5/2/73 5/8/73 Mean (60.33) 58.06-60.51 (59.70) 58.34-64.05 (61.52) 56.80-63.90 (60.34) 58.08-64.52(61.18) 56.05-65.50 (60.52) Mean (37.80) ,38.17-40.38(39.00) ,35.47-41.33 (38.20) .36.90-41.00 (38.93) ,37.17-40.08(38.86) 35.40-40.80 (39.05) White, pers. comm. White, pers. comm. This study White, pers. comm. This study Mean (60.65) Mean (38.80) 'These were probably collected from Minnesota, North Dakota, Manitoba, Saskatchewan, and Quebec for Phalacrocorax auritus, since no informa- tion on egg volumes for this species was re- ported in the hterature at that time. The vokimes were calculated by using the for- mula Volume = 0.526LB2, where L = lon- gitudinal axis and B = transverse axis (Romanoff and Romanoff 1949). The mean volume for the 48 eggs measured at Geneva in 1972 was 48.09 ± 2.80 cc, with a range of 41.55 to 52.69 cc. The mean volume for the 52 eggs measured at Geneva in 1973 was 48.60 ± 3.89 cc, with a range of 38.00 to 54.24 cc. Table 4. Average length and width measurements of all eggs measured at the Geneva Colony in 1972 and 1973, compared to random samples of ont egg from each of 12 clutches. Year Sample Length(mm) Width(mm) 1972 Overall mean 60..34 1972 1st random sample 56.67 1972 2nd random sample 60.,38 1972 3rd random sample 60.69 1973 Overall mean 60.52 1973 1st random sample 60.08 1973 2nd random sample 60.55 1973 3rd random sample 60.67 38.93 ,39.18 38.74 38.94 39.05 ,39.06 .39.26 38.92 Romanoff and Romanoff (1949) have stated that the volume (V) for an egg can also be calculated if the weight (y) is known by the equation V = 0.913 y for precocial birds and V = 0.959 y for altricial birds, such as the cormorant. This seems a very unreliable method, since cormorant eggs have a tendency to lose weight after they have been incubated for a time. Rahn and Ar (1974) reported that during incubation all bird eggs may lose approximately 18 percent of their initial weight. The average weight for 12 unincubated cormorant eggs was reported to be 46.9 gm, while 21 in- cubated eggs averaged 46.2 gm (Lewis 1929). As a further check on the reliability of this method, I calculated the volumes of 20 eggs, for which the weight was known, and listed the volumes computed for each egg by using both methods (Table 5). The aver- age weight of all the eggs was very close to the average of 46.5 gm for 50 eggs weighed by Lewis (1929). However, there is a differ- ence of almost 4 cc between the average volume calculated from length and width measurements and the average volume cal- culated by weight. A T-test (two-tailed) at 5 percent indicated that there was a signifi- cant variation between these means. Further 12 Great Basin Naturalist Vol. 37, No. 1 Table 5. A random selection of 20 eggs, taken at different times and from different Utah Lake colonies, show- ing the volume as figured by length and width measurement compared to volume computed from weight. (Ar- ranged according to increasing weight.) Width Length Weight Volume " Volume " No. (mm) (mm) (gm) (cc) (cc) 1 35.47 58.54 37.61 .38.59 36.07 2 38.33 60.10 42.03 46.44 40.30 3 38.72 58.08 42.45 45.80 40.71 4 38.17 58.06 42.58 44.60 40.84 5 37.62 58.34 42.62 43..35 40.87 6 38.99 58.87 43.39 47.12 41.61 7 .37.17 62.74 43.54 45.64 41.75 8 .39.11 59.63 44.78 47.93 42.95 9 37.43 59.43 44.84 43.70 43.00 10 38.85 59.37 44.93 47.28 43.09 11 38.01 64.05 45.40 48.69 43.54 12 38.53 61.96 45.79 48.34 43.91 1.3 39.21 60.08 47.50 48.50 45.55 14 39.86 59.88 48.16 50.16 46.18 15 39.73 64.52 49.58 .53.47 47.54 16 38.92 62.97 49.64 50.15 47.60 17 41.33 61.92 50.86 55.54 48.77 18 39.86 61.59 51.74 51.58 49.61 19 40.08 60.48 52.05 51.17 49.92 20 29.16 61.14 52.79 49..39 50.63 Means 38.73 60.59 46.11 ±4.05 47.87 ±3.81 44.22 ±3.87 ' Volume as computed by formula; Volume = 0.526 leugth x breadth^. ' Voliune as computed by formula; Volume = 0.959 x weij^ht. evidence of the unreliability of using weight to calculate volume is evidenced by com- paring samples #18 and #20 (Table 5). Al- though sample #20 is smaller in both di- mensions, it weighs over 1 gm more than sample #18. Therefore, the formula given by Romanoff and Romanoff (1949) for cal- culating volume from the weight of the egg (V = 0.959 y for altricial birds) may hold true for some species, but is not accurate for the Double-crested Cormorant. Recently, Pagnalli et al. (1974) reported a mean volume for eight Double-crested Cor- morant eggs from Maine as 45.87 ± 2.75 cc. Their method involves tracing a profile of the egg and dividing the tracing into seg- ments, then calculating the surface area of each segment, and from that the volume. However, they appear to be working with a somewhat larger bird, since they report a mean egg weight of 49.88 ± 3.37 gm, and I found a mean of 46.11 ± 4.05 gm. The length of the eggs was also different since they reported a mean length of 6.19 ± 0.21 cm and mine were smaller at 6.04 ± 0.19 cm. Additionally, Preston (1974) has recently stated that the volume of an egg cannot be accurately calculated on the basis of length •and width measurements alone without er- rors of 5 percent or more commonly occur- ring. Because the cormorant egg is pointed at both ends and describes the shape of a bicone, the coefficient given by Romanoff and Romanoff (1949) would be too large. Therefore, my values for volumes would be overestimated and may result in the differ- ence between my data and those of Pagnalli et al. (1974). Hatching — Hatching at the Springville Colony began on 22 April and continued until the second week in June. At the Bay Colony hatching began on 30 April and ended on 27 May. In 1972 hatching began March 1977 Mitchell: Cormorant Biology Table 6. Summary of the egg laying and hatching success for the Utah Lake colonies. 13 Colony Year No. Average Total No. Average Total No. Hatching Nests Height (m) Eggs Clutch Size Hatched Success Geneva 1972 19 unk. 80 4.2 44 55% Springville 1973 33 5.2 126 .3.8 83 66% Bav 1973 7 5.3 28 4.0 15 54% Geneva 1973 17 4.7 61 3.6 18 30% at Geneva on 19 May and continued until 10 June. In 1973 hatching began on 20 May and lasted until 9 June. Clutch size, total number of eggs laid, and hatching success are presented in Table 6. The hatching dates reported for each col- ony reflect an average of 28 days for in- cubation. The earliest hatching began at Springville on 22 April 1973 but, owing to the extended arrivals of new birds, lasted well into June. There was a remarkable syn- chrony of hatching periods at Geneva dur- ing 1972 and 1973. In 1972 hatching began on 19 May and continued for 21 days, while in 1973 hatching began one day later and lasted 20 days (Fig. 6). Two to four days are usually required for the complete hatch- ing of a single clutch since incubation starts soon after the first egg is laid (Mendall 1936). Hatching of the first and last bird may be separated by as much as a week or more (Palmer 1962). When this is the case, the youngest birds do not usually survive. Ricklefs (1969) stated that the hatching success for most colonial nesting birds .should be over 50 percent. There is no re- port of hatching success for P. a. auritus, but a record for P. a. cincinatus revealed a hatching success of 60.4 percent (Drent et al. 1964). The hatching success at Spring- ville Colony (66 percent) and Bay Colony in 1973 (54 percent), and Geneva Colony in 1972 (55 percent), provided an average of 58 percent (Table 6). That is probably a fair indicator for this subspecies, although the hatching success for grovmd-nesting cormo- rants may vary considerably from that val- ue, due to other external factors. The very low hatching success at Geneva in 1973 (30 percent) requires some explanation other than predation. I believe that the daily vis- its I made to Geneva, in combination with the cool weather during incubation, may principally account for the low success. In order to complete my daily counts, it was sometimes necessary to keep the adults 3 3 ^ 2 19 20 21 22 23 24 25 26 27 28 29 30 31 1 2 3 4 5 6 7 May June DATE Fig. 6. Distribution of egg hatching at Geneva Colony from 19 May to June 1973. 9 10 11 14 Great Basin Naturalist Vol. 37, No. 1 from the nest for as long as 45 minutes, al- though the interval was usually less. While awaiting my departure, the adults flew to the reservoir and when I left immediately flew back to the nest. Their moist feathers and water on the eggs may have resulted in an extended period before optimum in- cubation temperature was again reached (Lack 1968). Ricklefs (1969) has stated that high tem- perature in eggs, caused by exposure to the sun, is a serious mortality factor. This may have contributed to the mortality of some of the eggs in exposed nests, but may not be an important factor, since temperatures were not high until the last of May, when 90 percent of hatching was already com- pleted. Seven of the Geneva nests that were regularly attended by the adults in 1973 contained only addled eggs and made up 39 percent of all Geneva eggs. That is consid- erably higher than the 24 percent of addled eggs reported for a colony of ground-nest- ing cormorants {P. a. cincinatus) in British Columbia (Drent et al. 1964). All nests at Geneva that contained only addled eggs were still being incubated by the adults two weeks after the expected hatching date. Three of these nests were still being in- cubated 30 days after the estimated hatch- ing date. Incubation of addled or infertile eggs by cormorants beyond the normal in- cubation period is well documented (Snow 1960). Preckition of Eggs and Ytiing.— Predation on the eggs and young of cormorants is common. Ferry (1909) observed predation by Ring-billed Gulls {L(irits delawarensis) on the eggs of cormorants in Saskatchewan. Lewis (1929) reported predation on eggs and young by Great Black-backed Gulls {Lams tnarinus), crows {Corvus brachy- rhynchos), and ravens (Corvus corax). This only occurred when the adults were absent from the nests, since they are usually in- trepid defenders of their nests. Herring Gulls {Lams argentatus) have also been ob- served attacking eggs and young of un- guarded cormorant nests (Bourget 1973; Mendall 1936). At Mandarte Island in Bri- tish Columbia, crows and gulls removed eggs from nests that had been vacated for only a few minutes (Drent and Guignet 1961, Van Tets 1959), and 28 percent of the eggs in the colony were lost due to these predations in 1963 (Drent et al. 1964). Cali- fornia Gulls are known to attack and con- sume the eggs and young of unguarded nests (Behle 1958, Odin 1957, Vermeer 1970). The personnel at Bear River Refuge informed me that they had to employ meth- ods to discourage gulls from nesting because they were so destructive to the eggs and young of water birds, including the cormo- rant. There are no records of mammalian pre- dators on the eggs and young of cormo- rants. Lewis (1929) reported that, even when there were large numbers of coyotes {Canis latrans) in an area where a nesting colony was accessible, they were never known to attempt any attack. Contrary to these reports and the state- ment made by Armstrong (1965) that gulls always attack eggs in vacated cormorant nests, I never observed mammalian or avian predation at any of the Utah Lake colonies. In addition, I never saw any evidence of predation, such as broken eggs, ravaged young, etc., which would indicate attacks during my absence, although I anticipated serious problems from the California Gulls when I disturbed the nesting birds at Gen- eva. But even when I kept the cormorants from their nests for as long as 45 minutes, 1 never observed an aggressive act toward the eggs or young from a gull, although hun- dreds were flying nearby. It is not sugges- ted, however, that the gulls never attack cormorant eggs and young in this area. Food Items.— Most of the studies dealing with the food of the Double-crested Cormo- rant have been along the Atlantic coast (Lewis 1929; Mendall 1936). Therefore, they contain lists of marine species of fish and do not provide good information on the food habits of inland cormorants. The few re- cords that do exist include those of the Bio- logical Survey (Lewis 1929) that reported the stomachs of nine birds collected at the mouth of Bear River contained the follow- ing: carp {Cyprinus carpio), 73 percent; Utah Sucker {Catostomus ardens), 19 per- March 1977 Mitchell: Cormorant Biology 15 cent; Chub {Leuciscus litieatiis), 3 percent; Chub {Leuciscus sp.), 6 percent. Behle (1958) found the following species regur- gitated by young birds at Egg Island, Great Salt Lake: carp, catfish. Yellow Perch (Perca flavescens), Silverside Minnows {Richanl- sonius hydrophlox), and Webug Suckers {Ca- tostomus fecundus). Other records (Lewis 1929, Mendall 1936) have reported cormo- rants eating Bluegill Sunfish (Lepomis pal- lidus). Pumpkin-seed Sunfish {Lepomis gih- bosus), Northern Crappies {Pomoxis sparoides). Common Crappies {Pomoxis an- nularis), Northern Pike {Esox lucius). Wall- eye {Stizostedion vitreum), and a mudpuppy {Necturus maculosus). Palmer (1962) record- ed crayfish, frogs, Fathead Minnows {Pime- phales promelas), and dace {Rhinicthys sp.) as additional food items. Food items were obtained only from nest- lings that regurgitated when I approached or handled them. Some items that had been partially eaten were taken directly from the nest. Remains of Black Bullhead {Ictahirus melas). Carp, and Utah Chub {Gila atraria) were found at all three colonies; Goldfish {Carassius auratus) were found only at Gen- eva Colony, and White Bass {Roccus chry- sops) only at the Springville Colony. The fish found ranged in size from 3 cm (gold- fish) to 22 cm (bullhead). Of these five species of fish the nestlings were known to use for food, the Utah Chub, White Bass, and Goldfish are report- ed here for the first time. Development of the Young.— When the young birds hatched they were naked, blind, and barely able to move, and they had a taut black skin, although Lewis (1929) and Men- dall (1936) reported that they had a dark- brown skin that turned black within a day or two. From about the fourth day after hatching (Table 7), the oldest birds in the clutch were very aggressive. This sometimes resulted in the smaller or younger birds being deprived of food and ultimately dying of starvation (Behle 1958, Hall 1926). At Geneva three birds apparently starved when only a few days old while their siblings grew normally and continued in good health. Although starvation may be the major cause of death of fledglings, very young birds are also susceptible to death by expo- sure during hot weather. On hot sunny days it was necessary for the adults to shade the young, and I have observed the young die within minutes when the adults were fright- ened away. Of the 18 young that hatched at Geneva in 1973, 13 survived long enough to leave the nest. After they were a week old, the birds were better able to move into shade and employed gular flutter to cool themselves as indicated by Bartholomew et al. (1968) and Lasiewiski and Snyder (1969). Table 7. Reactions to human handling by Double- crested Cormorant nestlings during specific age peri- ods. Age (days) Reaction Can barely hold head up; no physical re- sistance or escape attempts; cower and shiver when held; usually make no sound; crawl toward shade when exposed 4-7 Head held erect to peer at intruder; make a whining sound; cower when hand reached out; attempt to hide be- hind nest-mates; sit quietly when head or eyes are covered. 8-13 Defecate or regurgitate food when han- dled; peck at intruders; squawk loudly; peck at nest-mate when placed back in the nest; quiet down when head is cov- ered. 14-22 Climb out of nest onto limbs away from intruder; defecate or regurgitate when approached; use toes and beak to hang onto any object within reach; will not remain quiet even when head covered; peck viciously when handled. 23- Climb frantically out of nest when ap- proached; often fall or jump out of nest into water; swim well at first attempt. The eyes opened when the birds were about four to five days old and the egg tooth dropped off at four to seven days, al- though some birds retained the egg tooth for as long as 12 days. A fine black down appeared about the sixth day. Eight days 16 Great Basin Naturalist Vol. 37, No. 1 later the birds were completely covered by down. When two weeks old, the young were very adroit in the use of beak and feet for grasping. When taken out of the nest, they would grab onto anything with their beak and toes, and it was difficult to dis- lodge them. I observed a four-week-old bird climb 5 m up a tree to the nest from which it had fallen. Until about two weeks old, the young were fed a semiliquid diet by regurgitation from the adults. Thereafter, whole or par- tially digested fish were brought to the nest. At i 7 to 19 days the flight feathers first ap- peared along the alar tracts, and the light- colored gular pouch became a deeper yel- low. The young were fully feathered at eight weeks and were easily recognizable on the water or in flight by their brownish backs and cream-colored breasts, in contrast to the glossy black of the adults. Measurements of the Young.— Three measurements of the nestlings in accessible nests at Geneva Colony in 1973 were taken in order to determine growth patterns. Due to the low hatching success, only 16 young were available for measurements and three of those died within four days. Each bird was weighed daily and linear measurements of the beak and tarsometatarsus taken. Fig- ures 7, 8, and 9 contain summaries of these measurements. The average weight at hatching was 36 gm (Fig. 7), and tarsometatarsal length was 11 mm (Fig. 8). This is close to Lewis's (1929) measurements of 37 gm and 12 mm, respectively. Mendall (1936) reported an av- erage weight at hatching of 32 gm. The av- erage beak length of the Geneva birds at hatching was 17 mm (Fig. 9), but Lewis (1929) reported an average length of 11 mm. The discrepancy is due to different measuring techniques. I measured along the side of the beak from the corner of the mouth to the tip of the upper mandible, and Lewis measured from base to the tip along the top of the beak. The average weight for Mendall's (1936) birds at five days was 158 gm, and mine averaged 156 gm. At 14 days Lewis (1929) reported an average weight of 778 gm, Mendall report- ed 785 gm, and mine averaged 736 gm. At 20 days the Geneva young averaged 1158 gm; Mendall (1936) recorded an average of 1232 gm. The average weight for the Gen- eva young after the 23-day period was 1543 gm (Fig. 7). Gonsiderable weight addition takes place thereafter, since Lewis (1929) gave the average weight for adult males as 2100 gm and adult females as 1670 gm. Those values differ from birds collected in Maine where 10 adult males averaged 2233 gm and 12 adult females averaged 1861 gm (Kury 1968). The Utah birds are even larger than those in Maine. Four specimens taken from Springville in 1974 averaged 2247 gm for two females, and 2553 gm for two males. Other than the averages recorded at hatching, there is no additional information for beak measurements and tarsometatarsal growth in the literature. When 23 days old, the young had an average beak length of 73 mm and an average tarsometatarsal length of 58 mm. Palmer (1962) records the aver- age tarsometatarsal length of adult birds as 61 mm. Measurements of an immature cor- morant (BYU Life Sciences Museum) in- dicated a beak length of 89 mm and tarso- metatarsal length of 58 mm. The relationship of these measurements to the growth patterns values are indicated in Figs. 8 and 9. The growth curve of the beak indicates continual increase through the 23rd day, but the tarsometatarsal length levels out at approximately 57 mm after the 19th day. Nest Departure.— There is little agree- ment among authors as to when young cor- morants first leave the nest and how they accomplish their departure. Some, but not all, of the variation can be explained by the difference in nest location, such as nesting on low islands, on cliffs, or in trees. Towns- end (Bent 1922) noted that the young re- mained in tree nests until fully feathered and flew at about eight weeks of age. How- ever, if they fell into water before they could fly, they would make no attempt to swim. Lewis (1929) reported that the young stayed in tree nests until about six weeks old, when they made their first flight. Men- dall (1936) observed that young birds flew feebly to the water from cliff nests at six March 1977 Mitchell: Cormorant Biology 17 weeks of age, but were not able to dive un- til seven weeks old. Lack (1968) stated that cormorants remained in the nest until they could fly and did not swim until after they were able to fly. Cormorants on Egg Island, Great Salt Lake, were observed to swim and dive long before they were able to fly, although no age was noted for either event (Behle 1958). In contrast. Burns (1921) found that the young usually fell, rather 16 16 13 13 13 12 11 1 1 n 10 u 8 1600 1500 ■ 1400 ■ 1300 1200 1100 1000 .. 900 800 t 700 600 500 + 400 300 200 + f f 100 -♦- + + DAY Fig. 7. Mean increase in weight (gm) from measurements on alternate days. The vertical line represents the range, the horizontal line the mean, and the blackened area is equal to two standard deviations. The sample size is given at the top of each day's measurements. 18 Great Basin Naturalist Vol. 37, No. 1 than flew, into the water from nests in trees. Some of these birds were only 17 days old when they fell from the nests. I observed that very few tree-nesting cor- morants in Utah remained in the nest until they were able to fly. After the young were approximately three weeks old, they would leave the nest and move onto tree limbs in order to increase the distance between them and me, which often resulted in young birds falling into the water. At first I suspected that they had drowned, but I later observed them swimming as far as 15 m underwater. After a large number had fallen out of the nest they could be seen swimming in groups some distance from the colony. I presumed Fig. 8. Mean increase in tarsometatarsal length (mm) from measurements on alternate days. The vertical line represents the range, the horizontal line the mean, and the blackened area is equal to two standard deviations. The sample size is given at the top of each day's measurements. March 1977 Mitchell: Cormorant Biology 19 that my presence caused early departures until I visited the colony at Newstate Gun Club in June 1973. At the time of my visit the birds had not been disturbed for three weeks, yet when I arrived at the colony there were 23 immature birds between three and six weeks of age swimming in the canal. I satisfied myself that they were not able to fly by chasing them up and down the canal. While I was there, other young fell into the water and two birds that fell onto the ground immediately walked over Fig. 9. Mean increase in beak length (mm) from measurements on alternate days. The vertical line represents the range, the horizontal line the mean, and the blackened area is equal to two standard deviations. The sample size is given at the top of each day's measurements. 20 Great Basin Naturalist Vol. 37, No. 1 to the canal and swam away. Early depar- ture from the nest would require the young to learn quickly to fish or to continue to be fed by the adults (Behle 1958, Lewis 1929, Mendall 1936). I have never observed young cormorants being fed on the water, but I assume the adults continue to feed them until they are able to fish effectively for themselves. The dates stated below pertain to the time when the birds were sufficiently devel- oped to be absent from the nest and prob- ably survive. This does not imply that the birds were able to fly when they left the nest. In all observed cases the birds fell or jumped from the nest into the water and swam expertly away. This made it impos- sible to estimate the date when the fledg- lings were able to fly. The earliest date at which birds were absent from the nest at Geneva in 1973 was 17 June, and the last nestling left on 5 July. Age at the time of leaving was between three and six weeks. The oldest bird to leave was 37 days old and the youngest was 21 days old. The av- erage age for leaving the nest was 29 days. Migration.— Most of the local cormorants leave the Utah Lake area in October. Al- though I have seen them on Utah Lake as late as 18 November, there is no way to tell if these are local birds or others migrating through. The latest date they have been seen at Bear River is 1 December (Behle 1958). Palmer (1962) stated that almost all of the Atlantic coast migrants have arrived at their wintering grounds in Texas and Louisiana by 1 December. An unusual sighting was made by Reed Ferris in 1959 (Scott 1959) when he saw a cormorant at Geneva reservoir on 1 Janu- ary. Since the reservoir is supplied with warm water, it does not freeze during the winter and the bird would be able to fish. However, there was no way to tell whether the bird was a late southern migrant, a bird that had overwintered, a very early arrival, or an injured bird. Discussion Status of Utah Lake Colonies.— Provo Bay has been a major nesting site for cor- morants for over 50 years (Bee 1920), al- though it has received little attention. The population there has at times exceeded 300 nesting birds (Bee 1949). Smaller numbers of cormorants have nested in the Geneva Colony but have probably done so contin- uously for almost 30 years (Bee 1946). The total number of cormorants breeding in the Utah Lake area for the past five years has probably remained stable at around 100 nesting pairs, although the numbers utilizing each colony fluctuate (Table 8). Table 8. Numbers of active nests in the Utah Lake colonies over a five-year period. Year Spriri .gvi ille Bay Geneva Totals 1970 Unk. 14 26 Unk. 1971 68 17 18 103 1972 48 12 31 91 1973 83 6 15 104 1974 96 7 0 103 Utah's Cormorant Population.— In 1958 seven cormorant breeding colonies were re- ported in Utah, but five had been deserted or destroyed (Behle 1958). Behle made no mention of the colonies that were located on White Rock, Great Salt Lake, and Rock Island, Utah Lake, or the existing Utah Lake colonies (Fig. 2). However, the former two sites were limited to a few birds and were probably used only briefly. In 1974 the only Utah cormorant colonies in use were located at New State, Utah Lake, and Bear River Migratory Bird Ref- uge. The latter consists of two groups of birds nesting on two small islands, one lo- cated in unit four and the other in unit five. Although there is no published reference to nesting since 1949 (Behle 1958), records kept by the refuge personnel indicate con- tinuous usage through 1974. The population of Double-crested Cormo- rants in Utah has been steadily decreasing for the past 50 years. Behle (1935) reported a decrease in cormorant numbers on Egg Is- land from 500 birds in 1915 to 114 in 1935. He felt that the breeding cormorants in the March 1977 Mitchell: Cormorant Biology 21 region were threatened with extinction. Sugden (1936) noted an alarming decrease in the numbers of herons and cormorants nesting on the Great Salt Lake and made a plea for their protection. Of the 13 colonies that have existed at one time or another within the state, only five were still in use in 1974 (Table 9). Dur- ing the 1930s and 1940s the larger colonies may have easily supported a combined total population of well over 1000 birds. How- ever, in 1973 the total known cormorant population of Utah consisted of only 386 birds nesting in five colonies (Table 10). Table 9. Summary of the populations of Utah's cor- morant colonies and their current status. Table 10. Summary of Utah's population as of 1973. existing cormorant Largest recorded no. nesting 1973 Colony birds Year status Bear River 600 1947 94 New State 84 1973 84 Geneva 62 1972 30 Springville 166 1973 166 Bay 300 1949 12 Bass Pond 80 1927 destroyed Bear Lake 400 1929 deserted Logan 400 1939 deserted Dolphin Island 150 1919 deserted Egg Island 500 1915 deserted Farmington Bay °2 1951 deserted Rock Island °8 1937 deserted White Rock °2 1901 deserted "Thesf are the a luthor'^ , estimates base don the hteratiire. Colony Percent of No. nesting birds state total Springville 166 43 Bear River 94 24 New State 84 22 Geneva 30 8 Bay 12 3 Totals 386 100 for years. Entire colonies have been wiped out and their numbers reduced by the thou- sands (Buchheister 1944, Demille 1926, Lewis 1929, McLeod and Bondar 1953, Smith 1911, Stanford 1937). The diking and subsequent draining of Provo Bay will only compound an already critical situation for Utah's steadily declining Double-crested Cormorant population unless adequate measures are taken to protect their present nesting sites. Acknowledgments.— I appreciate the en- couragement and suggestions offered by Dr. Herbert H. Frost during my research and for his critique of this manuscript. I thank Dr. Clayton M. White for his suggestions and critique of this manuscript and Dr. Stephen L. Wood for the identification of insects found in the nests. Appreciation is also extended to the U.S. Steel Company, Geneva Works, for allowing access to the Geneva Colony. Effects of Diking Provo Bay — Two of the five remaining cormorant colonies in Utah are located on Provo Bay, Utah Lake (Fig. 2). In 1973 the Springville Colony was the largest in the state and supported 43 per- cent of the total remaining birds in Utah and, combined with the small Bay Colony, comprised 178 birds or 46 percent of the state's total cormorant population (Table 10). The proposal to dike and drain Provo Bay as part of the Central Utah Project (Fig. 2) would eliminate the nesting area of these birds. Cormorants have been subjected to per- secution in the United States and Canada References Armstrong, E. \. 1965. Bird display and behavior. Dover Publications, Inc., New York. Bartholomew, C. A., D. C. Lasiewiski, and E. C. Crawford. 1968. Patterns of panting and gular flutter in cormorants, pelicans, owls, and doves. Condor 70:31-34. Bee, J. 1946. Unpublished field notes. University of Kansas, Lawrence, Kansas. Bee, R. G. 1920. Bee Oological Collection. Life Sci- ences Museum, Brigham Young University, Pro- vo, Utah. 1937. Unpublished field notes, 3:105. Life Sci- ences Museum, Brigham Young University, Pro- vo, Utah. 22 Great Basin Naturalist Vol. 37, No. 1 1949. Unpublished field notes, 6:93-106. Life Sciences Museum, Brigham Young University, Prove, Utah. Behle, W. H. 1935. A history of the bird colonies of the Great Salt Lake. Condor 37:24-35. 1958. The bird life of Great Salt Lake. Univ. Press, Salt Lake. Bent, A. C. 1922. Life histories of North American petrels and pelicans, and their allies. U.S. Nat. Mus. Bull. 121:236-261. Bergtold, W. H. 1917. A study of the incubation pe- riods of birds. Kendrick-Bellamy Co., Denver. BoiRGET, A. A. 1973. Relation of eiders and gulls nest- ing in mixed colonies in Penobscot Bay, Maine. Auk 90:809-820. BucHHEisTER, C. W. 1944. The comeback of the cor- morants. Audubon 46:14-29. Burns, F. L. 1921. Comparative period of nesting life of some North American Nidicolae. Wilson Bull. 33:90-99. CoTT.\M, C. 1927. Distributional list of the birds of Utah. Unpublished master's thesis, Dept. of Zo- ology, Brigham Young University, Provo, Utah. D.-vviE, O. 1900. Nests and eggs of North American birds. David McKay, Philadelphia. Demille, J. B. 1926. Birds of Gaspe County, Quebec. Auk 43:510-514. Dre.nt, R. H. and C. J. GuiGNET. 1961. A catalogue of British Columbia sea-bird colonies. Occas. Pa- pers Brit. Columbia Prov. Mus., No. 12. Drent, R. H., G. F. Van Tets, F. Tampa, and K. Vermeer. 1964. The breeding birds of Mandarte Island, British Columbia. Can. Field Nat. 78:208-263. Ferry, J. J. 1909. Birds observed in Saskatchewan dur- ing the summer of 1909. Auk 27:189-190. Ganier, a. F. 19.33. Water birds of Reelfoot Lake. J. Tenn. Acad. Sci. 8:65-83. Gu.nther, L. 1974. Manager of Bear River Migratory Bird Refuge. Personal communication. Hall, R. E. 1926. Notes on water birds nesting at Pyramid Lake, Nevada. Condor 28:87-91. Howell^ A. H. 1911. Birds of Arkansas. U.S. Biol. Surv. Bull. 38:15-16. KuRY, C. H. 1968. Differences in weight of male and female cormorants. Auk 85:113. Lack, D. 1968. Ecological adaptations for breeding birds. Methuen and Co. Ltd., London. Lasiewiski, R. C. and G. K. Snyder. 1969. Response to high temperature in nestling Double-crested Cormorants and Pelagic Cormorants. Auk 86:529-540. Lewis, H. F. 1929. The natural ni. .ory of the Double- crested Cormorant {Phaiacrocorax aurittii aur- itu.s Lesson). Ru-.Mi-Lou Books, Ottawa. 1931. .additional information concerning the Double-crested Cormorai t (Phaiacrocorax aur- itus auritus Lesson). Auk 1 .5:207-214. Lockerbie, C. W. 1942. Bird ,■•. about the Great Salt Lake. News Bull. Mi^ieralogical Soc. Utah, 3:fv4-66. Marshall, W. H. 1937. Double-crested Cormorant nesting on the Bear River Refuge in Utah. Con- dor 39:36. McLeod, J. A. and G. F. Bondar. 1953. A brief study of the Double-crested Cormorant on Lake Win- nepegosis. Can. Field Nat. 67:1-11. Mendall, H. L. 1936. The home life and economic status of the Double-crested Cormorant {Phaia- crocorax auritus auritus Lesson). Univ. Maine Studies, Second Ser., No. 38, Maine Bull. 39(3). Munro, J. A. 1927. Observations on the Double- crested Cormorant (Phaiacrocorax auritus) on Lake Manitoba. Can. Field Nat. 41:102-108. Odin, C. R. 1957. California gull predation on water- fowl. Auk 74:185-202. Pagnalli, C. v., a. Olzowka, and A. Ar. 1974. The avian egg: surface area, volume, and density. Condor 76:319-325. Palmer, R. S. 1962. Handbook of North American birds I. Yale Univ. Press, New Haven. Pearson, T. G. 1927. A-birding in Utah. Bird-Lore 29:379-383. Preston, F. W. 1968. The shape of birds' eggs: mathe- matical aspects. Auk 85:454-463. Preston, F. W. 1974. The volume of an egg. Auk 91:132-1,38. Rahn, H. and a. Ar. 1974. The avian egg: incubation time and water loss. Condor 76:147-152. Ricklefs, R. E. 1969. An analysis of nesting mortality in birds. Smithsonian Contr. Zoology, No. 9. Romanoff, A. L. and A. J. Romanoff. 1949. The avian egg. Wiley and Son, New York. Scott, O. K. 1959. Great Basin, Central Rocky Moun- tain Region. Audubon Field Notes 13 (3):311- 312. Smith, F. 1911. Double-crested Cormorants breeding in central Illinois. Auk 28:16. Snow, B. 1960. The breeding biology of the Shag (Phaiacrocorax aristotelis) on the Island of Lun- dy, Bristol Channel. Ibis 102:554-575. Stanford, J. S. 1937. Cormorant and heron colonies in Cache Valley, Utah. Proc. Utah .\cad. Sci., Arts, Letters 14:195. Stoddard, H. L. 1922. Bird notes from southern Wis- consin. Wilson Bull. 24:67-71. Sugden, J. W. 1936. Sanctuaries of the bird islands of Great Salt Lake. Auk 53:289-294. Van Tets, G. F. 1959. A comparative study of the re- productive behavior and natural history of three sympatric species of cormorants (Phaiacrocorax auritus. Ph. penicillatus. Ph. pelogicus) at Man- darte Island, B. C. Unpub'ished master's thesis, Univ. British Columbia Vermeer, K. 1969. Coloni ; oi ' . .hlc •.• ' Cormo- rants and White Pi .i. .i" e { n. Field Nat. 83:.36-39. 1970. Some aspe. - " h nes'; .. Double- crested Cormorant., a, ^^, Saskatche- wan, in 1969; a plea t. r ,)roteccion. Blue Jay 28:11-13. 1973. Great Blue Heron and Double-crested March 1977 Mitchell: Cormorant Biology 23 Cormorant colonies in the prairie provinces. 1949. Annotated checklist of the birds of Utah. Can. Field Nat. 87:427-4.32. Bull Univ. Utah 39(16) Bio. Ser, 11(2): 1-40. White, C. Professor of zoology, Brigham Young Uni- Worth, C. B. 1940. Egg volume and incubation peri- versity, Provo, Utah. Personal communication. od. Auk 57:44-60. Woodbury, A. M., C. Cottam, and J. W. Sugden. CATTAIL DECLINE AT FARMINGTON BAY WATERFOWL MANAGEMENT AREA' Christine Minton Anderson' Abstract.— For the last seven years, 1969 to 1976, a steady decline of cattails (Typha latifolia L.) has been ob- served at Farmington Bay Waterfowl Management Area, Davis County, Utah. Several parameters of Farmington Bay's environment that could cause or enhance a decline of the marshland vegetation were studied. These pa- rameters included temperature, pH, phenols, oil and grease, heavy metals, fungus pathogens, and salt water in- trusion from the Great Salt Lake. Elevated soluble salt concentrations were found to be responsible for the de- cline. Cattails are weakened or killed when they are exposed to soluble salt concentrations greater than 5.0 gm /liter, and there is a significant (1 percent level) negative correlation (-0.68) between soluble salt concentra- tion and cattail height. Symptoms of elevated salt concentrations include stunted growth, leaf tip necrosis (burn- ing), and occasional browning of an entire cattail clone. Furthermore, those salt concentrations that cause phys- iological stress in cattails also facilitate the growth of a decomposition fungus, Chaetophoma confluens. This fungus causes a rot consisting of irregularly scattered lesions on the surface of the rhizomes and was consistently isolated from rhizomes of declining plants. For the last seven years, 1969 to 1976, a steady decline of cattails has been observed at Farmington Bay Waterfowl Management Area, Davis County, Utah. The management area is 4,205 feet above sea level, located on the east shore of the Great Salt Lake, 15 miles north of Salt Lake City. The primary water source is the Jordan River. Water flows northwesterly, and Farmington Bay flows directly into the Great Salt Lake. At Farmington Bay, cattails in many areas were less than 1 m high, as compared to healthy cattails that frequently attained heights of 2 m. During the summer, cattails appeared to grow normally until July; growth then ceased. Yellowing and necrosis followed rapidly. Older parts of the cattail clones died first, with the younger shoots gradually following the same pattern. In- spection of the rhizomes revealed the pres- ence of mahogany-colored cankers, or le- sions, scattered irregularly over the surface. In some cases the lesions had extended deep into the tissues, and some rhizomes had completely rotted away. These symptoms are characteristic of those caused by .soil fungus pathogens, and it was suspected that the lesions might be the result of such a fungus and, in turn, that the lesions might cause the plant's decline. A fungus might act either as a primary pathogen or a weak, secondary pathogen, affecting tissues subjected first to some other stress. If this were the case, it would be necessary to establish what stresses might be involved and to define their relative contri- bution to the cattail decline. The objectives of this study were to: 1) determine the cause or causes of the cattail decline; 2) evaluate the seriousness of the situation; and 3) provide information so that corrective or control measures could be de- veloped to protect existing marshland vege- tation and reestablish vegetation that had been destroyed. A fourth objective was to better under- stand the marsh ecosystem. In this system, decomposition is an ecological factor that may be important in affecting marsh plant interactions. The declining cattails offered an opportunity to study how fresh-water marshlands might undergo decomposition and change. The broad-leaved cattail {Typha latifolia L.) is common to North American marsh- lands and dominant in Utah's waterfowl management areas. Other common marsh 'This article is based on research that was done by the author for a master's thesis. The research was partially funded by the .\ssociated Students i the University of Utah's Environmental Studies Program. 'Biology Department, University of Utah, Salt Lake City, Utah 84n2. 24 March 1977 Anderson: Cattail Decline 25 plants are alkali bulrush (Scirpus paludosus A. Nels.), hardstem bulrush (Scirf)us acutiis iVIuhl.), salt grass {Distichli.s stricta Torr. Rydb.), and salicornia (Salicornia rubra A. Nels. and Salicornia pacifica Standi.). Important as a successional species, the cattail is a plant that traps sediment and nutrients, allowing the subsequent estab- lishment of bulrushes. Bulrushes, the climax species of the marsh, provide abundant wa- terfowl food. Cattails, on the other hand, have little value as waterfowl food, and, when it is dominant in waterfowl manage- ment areas, the cattail is considered a nui- sance because its dense stands make it use- less as nesting habitat or escape cover. Cattail rhizomes are palatable to humans, muskrats, and geese; the plants themselves are used by waterfowl for nesting and es- cape cover when their growth has been controlled. The most significant publication on the subject of Utah marshlands contains the re- sults of 25 years of experimental work at Ogden Bay Refuge, 12 miles west of Ogden, Utah, near the east shore of the Great Salt Lake (Nelson 1954). The purpose of the study was to determine how marshland could be developed and waterfowl popu- lations increased; records were kept to as- certain how plant community changes in- fluenced animal populations. Several of the conclusions were pertinent to the present study at Farmington Bay. Major factors limiting plant growth and waterfowl use at Ogden Bay included water supply, seasonal water fluctuations, fluctua- tions of the Great Salt Lake, and over- grazing. Salt tolerance differed among plant species. Of the emergent species, alkali bul- rush was the most tolerant; cattail was sec- ond. All of the marsh plants were able to tolerate soil salinities exceeding the salinity ranges for agricultural crops. As plant cover was increased, waterfowl populations in- creased because new nesting areas were cre- ated and abundant food was made available. After vegetation had been developed on the Ogden Bay Refuge, it became evident that cattails, especially in the less saline areas of the marsh, could effectively out- compete other plants. Dense stands, useless for waterfowl nesting, were established. In 1947, cattail control studies were initiated by the Utah State Department of Fish and Game (Nelson and Dietz 1966). The control methods used to experimentally eliminate cattails included cutting, crushing, and mowing; chemical sprays; explosives; fire; and drought. It was concluded that by ad- justing the water levels cattail overgrowth could be controlled; when mud flats were not covered with water during the growing season, large-scale cattail invasions occurred. At Fish Springs, a salt marsh west of Salt Lake City, Typha angustifolia (narrow- leaved cattail) does not form large commu- nities (Bolen 1964). Bolen noted that Scirpus acutus and Typha latifolia, common species of Utah's fresh-water marshes, did not occur at Fish Springs. He concluded from his data that while it was probable that salinity lim- ited the development of these species, water depth or some other factor was probably re- sponsible for their absence at Fish Springs. In contrast to the Fish Springs study, data on physical and chemical soil constituents at Bear River Bird Refuge indicated that so- luble salts affected the yields of aquatic plants. Areas low in soluble salts were more productive than areas high in soluble salts (Jensen 1940). It was determined that high salinity, in addition to affecting vegetative growth, interferes with seed germination and seed production of Utah marsh plants (Kaushik 1963). Literature published on other localities stresses the importance of soil salinity in in- teractions between marsh plants (Haller 1974). Salinity is considered to be a primary factor in determining plant distributions in Louisiana marshes; cattails are limited to fresh-water areas of less than 0.5 percent salt (Penfound and Hathaway 1938). Water levels, in addition to salinity, are also thought to determine plant distribution (Bourn and Cottam 1939). Cattail populations growing in disturbed .salt flats have salt tolerances characteristic of each species. A concentration of 1 per- cent sodium chloride in hydroponic tanks caused stunting, drying, and leaf curl in Typha latifoha. Typha angustifoUa was able to withstand 2 percent sodium chloride con- 26 Great Basin Naturalist Vol. 37, No. centrations. A hybrid intermediate, Tijpha glauca, had an intermediate salt tolerance (McMillan 1959). Since many terrestrial plants also differ widely in their salt toler- ance (Levitt 1972), an effort was made to determine the relationships between cattail decline and salinity at Farmington Bay. Methods Five different features of the problem were studied: 1) the isolation of a possible fungus pathogen; 2) the investigation of en- vironmental stress conditions; 3) the eval- uation of marshland by aerial photography; 4) laboratory experimentation on the effects of salinity on cattail growth; and 5) labora- tory experimentation on the effects of sali- nitv on the growth of an isolated fungus. 1. Isolation of the fungus: Diseased cat- tails were collected, and plant tissue was cut from necrotic lesions with a sterile scal- pel. The infected tissue was surface ster- ilized and placed on Difco-Bacto Agar with nitrogen or Difco potato dextrose agar. Plant tissue was surface sterilized with a 10 percent aqueous solution of Purex bleach. Pure cultures were obtained by transfer- ring the developing fungus mycelium onto fresh agar. Cultures were allowed to grow at room temperature. Slant test tube cul- tures provided a method of storage, and they were refrigerated until used. The fungi were identified. Fungus mycelium was mounted on temporary slides with a drop of water for most identifications. To aid in the identification of one species {Chaetophoma confluens), fixing, sectioning, and staining were used. Tissue was fixed in formalin- aceto-alcohol and embedded in paraffin. Sections, 15 microns thick, were stained with safranin 0 according to the methods described by Johansen (1940). 2. Environmental parameters: Studies were conducted to determine the environ- ment's contribution to the vegetation de- cline. This included measuring salinity, pH, biological oxygen demand (BOD), dissolved oxygen (DO), and temperature. Samples of soil and water were taken weekly during the months of May through September, 1974. All routine sampling was done at the four sampling sites shown on Figure 1. Cat- tails appear to be healthy at Site 1 and Site 3. Cattails are in a state of decline at Site 2; Site 4 represents the outlet of Farm- ington Bay and has no cattails. Conductivity of both soil extracts and wa- ter samples was measured in the laboratory at 25 C with a conductivity bridge (Rich- ards 1954). Then pH was measured to the nearest tenth of a pH unit with a Beckman pH meter. BOD samples were taken once a month and analyzed by the Utah State Di- vision of Health. Water samples for dis- solved oxygen were measured weekly in the field according to the azide modification; DO was measured in milligrams per liter and expressed relative to 100 percent satu- rated water (American Public Health Asso- ciation 1971). In addition, samples were chemically analyzed monthly by the Utah State Division of Health to determine which industrial pollutants were present. Air and water temperature were measured in the field with a centigrade thermometer. The data obtained were used to calculate the percent of dissolved oxygen in the wa- ter. Weather conditions for the summer were recorded and compared with previous summer conditions. 3. Mapping of vegetation: Aerial photo- graphs and moving films of Farmington Bay and the surrounding area were taken. The Ogden Bay Waterfowl Management Area, the Bear River Bird Refuge, and Red Butte Canyon were also examined for indications of vegetation decline in order to determine if the decline was in some way uniquely as- sociated with Farmington Bay. A vegetation map of Farmington Bay was made so that the affected area could be properly illus- trated and the extent of damage could be determined. A U-2 infrared aerial photo- graph (scale 1:120,000) was projected onto a United States Geological Survey Map (Farmington Quadrangle, scale 1:24,000) with a Bausch and Lomb Zoom Transfer Scope. Areas covered by marsh vegetation were traced onto the geological survey map. The U-2 infrared photograph, obtain- ed from the Geography Department at the University of Utah, was taken 14 September 1972 by a high-flying aircraft at about March 1977 Anderson: Cattail Decline 27 5(),()()() feet. Standard infrared film was used in a high-resolution camera. Healthy vegeta- tion is bright red, vegetation under stress is salmon colored, and dead vegetation is white. A compensating polar planimeter was used to determine the area covered by healthy marsh vegetation and the extent of diseased vegetation. Color aerial photography for assessing the quality of marshlands has just recently come into use. Further information on the meth- ods used in aerial mapping of marshlands can be obtained from Seher and Tueller (1973); the source also explains how infrared photographs should be interpreted. In addi- tion, a bibliography on the use of aerial photography (Anson 1975) has been com- piled that provides more detailed informa- tion. 4. Laboratory experiments determining ef- fects of salinity on cattail growth: Cattails were grown from seeds gathered at Farm- Healthy vegetation Stressed vegetation 'ying vegetation The dashed line shows where vegetation was in 1969. Fig. 1. Farmington Bay Waterfowl Management Area. 28 Great Basin Naturalist Vol. 37, No. 1 ington Bay. Seedlings were transferred to six hydroponic tanks when they were abont six inches high; the tanks contained a four-salt nutrient solution made according to the methods described by Arnon and Hoagland (1940). Nutrients were replaced monthly during the experiment. Each of six hydro- ponic tanks contained 10 individual plants. Two tanks served as the control; the other four tanks were used to determine the ef- fects of different salt concentrations on the cattails. After the seedlings had acclimatized and had shown vigorous growth, sodium chloride was added in small amounts (0.01 Mole per day) over a period of 30 days. This was done to determine a realistic salt tolerance for the cattails without putting them under undue stress. Shoot growth of each plant was measured every two days. The final concentrations at the end of the period of additions were two tanks with 0.0 M sodium chloride, two tanks with 0.1 M sodium chloride, and two tanks with 0.3 M sodium chloride. The cattails in one tank of each salt concentration were inoculated with Chaetophoma confluens, the fungus that was most often isolated from the le- sions on the cattail rhizomes. Inoculation was performed by injecting a fungus-water suspension into the host with a hypodermic syringe. Cattail rhizomes were observed through- out the experiment for the development of lesions and other symptoms of stress. When the experiment was terminated, a wet weight for shoots and rhizomes of each plant was obtained. Tissue from all experi- mental plants was cultured for isolation of the fungi that were present. 5. Effect of salinity on the fimgus, Chaeto- phoma confluens: Chaetophoma confluens was grown on potato dextrose agar, to which sodium chloride had been added. The following sodium chloride concentrations were prepared: 0.0, 0.1, 0.2, 0.3, 0.4, 0.5, 0.6, 0.7, and 0.8 M. Cultures of each salt concentration were inoculated with 0.5 cmsquares of mycelium and were allowed to grow for two weeks in an oven held at 30 C. The area covered by fungus myce- lium was determined by tracing the area onto paper, cutting, and weighing. The av- erage growth area at each salt concentra- tion was graphed. Results Three fungus species were isolated from diseased cattails that had been collected at Farmington Bay. Fusarium tricinctum was isolated in 12 (16.2 percent of the isolated fungi) out of 74 fungus isolations. Botrytis cinerea was isolated 14 times (18.9 percent). Both of these fungi are saprophytic on decaying plant material. F. tricinctum oc- curs frequently on moldy corn and grain; B. cinerea causes "gray mold" of numerous or- namental and crop plants (Barron 1968). Neither of these fungi has been reported to occur on Tijpha. Chaetophoma confluens was isolated 48 times (64.9 percent). Chaetophoma might be an important decomposition fungus in the marsh. It has been isolated from Spartina (cordgrass), a common emergent in salt-wa- ter marshes (Barnett 1956). Another fungus, tentatively identified as a species of Olpidium, was discovered when the stained sections of cattail rhizomes were examined; the relationship of this fungus to cattail rhizome rot is unclear. Only a few species of Olpidium are parasitic on higher plants. When Olpidium infects clover or vetch, it causes distortions of the leaves; in one species of grass, Agrostis stolonifera, it causes root hair distortion. Chytridiomi/cetes (the fungus class that includes Olpidium) are characterized by the production of motile cells and are typically found in aquatic hab- itats (Sparrow 1943).' The most probable cause of the vegeta- tion decline was revealed when the results of the water and soil analyses were exam- ined. Figures 2 and 3 show the results of the conductivity measurements. The figures show that the salinity increases dramatically during the summer at Site 2, reaching a maximum during the month of July. This peak coincides with the period during which cattail deaths were first observed dur- ing the summer. The salinity peak also came just after the Great Salt Lake reached its high of 4,201.30 feet above sea level in June (United States Geological Survey March 1977 Anderson: Cattail Decline 29 1975). The 4,205-foot level is shown along the east side of Farmington Bay on a United States Geological Survey Map of the area; the west side of the management area is approximately 4,200 feet above sea level. Apparently, the management area is at the same elevation as are the Great Salt Lake brines for part of the summer. Chemical analyses of the water at Farmington Bay showed that sodium and chloride ions were present in the greatest amounts at this time. It was concluded that these ions made the largest contribution to the measured con- ductivity values. To determine the significance of the rela- tionship between salinity and cattail de- cline, another experiment was conducted. This experiment consisted of taking water samples every tenth of a mile (0.16 km) along the east side of Farmington Bay and along the dike that runs east and west across the refuge. As water samples were taken, the average height of the cattails was recorded in centimeters. Cattails were meas- ured in a circular area of 10 m in diameter. The point where the water sample was taken served as the center of the circle. Av- erage heights were based on measurements of 20 to 50 cattail plants. Only living cat- tails were measured. Figure 4 shows a graph of conductivity versus cattail height. There is a moderately high, negative corre- lation (r = -.68 with 41° of freedom) be- tween conductivity and cattail height. These field data also showed that cattails exhibit disease symptoms when the dissolved salt content exceeds 5.0 gm/1. The pH of Farmington Bay lies between the normal ranges of river water: 5.0 to 9.0. The data showed that there was no correla- tion between pH and cattail height. The BOD at Site 2 progressively in- creased during the summer, while the BOD at Site .3 and Site 4 increased slightly and May May < Site #1 ASite #2 I Days Osite #3 QSite #4 < Site #1 ^Site #2 July August Time in Days <>Site 13 j^Site #4 Fig. 2. Conductivity of the water. Fig. 3. Conductivity of the soil. 30 Great Basin Naturalist Vol. 37, No. 1 then leveled off (Fig. 5). The large increase indicated that oxidizable organic material was accumulating at Site 2. Results of the DO measurements are shown in Figure 6. Dissolved oxygen at Site 2 got as low as 46 percent saturated. Despite some periods of low DO and high BOD, the dissolved oxy- gen was high overall during the summer and no detrimental effects were observed that could be related to low oxygen avail- ability. The results of the chemical analyses showed that the chemical parameters of the water were normal except for the high sali- nity. Small amounts of phenols, oil, and grease were found in the water, and a dis- cussion of their significance to this study is included in the Discussion section of this article. The effect of sodium chloride on the growth of the fungus, Chaetophoma con- fluens, is shown in Figure 7. The addition of sodium chloride was found to increase growth. Optimum concentration for growth of Chaetophoma is 0.30 M sodium chloride. Cattails growing in hydroponic tanks showed symptoms of salt toxicity (leaf burn and wilting) at sodium chloride concentra- tions of 4.68 gm/1 (0.08 M). At sodium chloride concentrations of 17.6 gm/1 (0.30 M), growth was completely arrested and rhizomes began to rot. The results correlate well with the results of the field study. Table 1 shows how increased salt concentra- tions affected the wet weights of shoots and rhizomes. Attempts to isolate Chaetophoma from le- sions that developed on cattails following inoculation were unsuccessful. Because of the observed effects of salinity on the cat- tails, and because Chaetophoma was only isolated in two (out of 20) cultures in pre- vious inoculation experiments, it could not be concluded that the fungus was a primary Conductivity (micromhos/cm at 25° C) Fig. 4. Conductivity vs. cattail height. May June July August September Time in Days A Site #2 OSite #3 (^Site #4 Fig. 5. Biological oxygen demand. March 1977 Anderson: Cattail Decline 31 cause of the cattail decline. The com- bination of the appropriate soluble salt con- centration and the fungus appears to be critical in producing the symptoms observed at Farmington Bay. Figure 8 shows that the optimum sodium chloride concentration for Chaetophoma growth (0.30 M) corresponds to the salt concentration (0.30 M) that causes cattail death. Table 1. Wet weights of cattail shoots and rhizomes in relation to salinity.' Sodium Shoot : Chloride Rhizome Shoot Rhizome Concentration Weight Weight Ratio 0.0 M 73.0 gm 209.7 gm 2.90 : 1 0.1 M 53.4 gm 58.6 gm 1.10: 1 0.3 M 26.2 gm U.lgm 0.42 : 1 These are the average weiijhts of all the plants tha Hiven sodium chloride concentration. Discussion At present, the vegetation in Farmington Bay and the neighboring duck clubs appears to be under stress. The stress conditions oc- cur throughout the marsh, leaving very few places unaffected. Healthy vegetation is found only in the higher areas of the marsh and where fresh water is flowing in from the Jordan River. The portion of Farmington Bay illu.strated in Figure 1 has an area of 11.88 square kil- ometers (2,934 acres). Of this, 4.88 km^ (1,205 acres) is emergent vegetation. In 1969, Farmington Bay had 5.92 km^ (1,462 acres) of emergent vegetation; 1.04 km^ (257 acres) of this vegetation has dis- appeared, and 1.28 km^ (316 acres) of vege- tation is dying and in the process of dis- appearing. It is interesting to note that in 1959 there were only 1.40 km^ (346 acres) of vegetation in the portion of Farmington Bay that is illustrated. At the present time. <3 Site #1 A5ite #2 Fig. 6. Dissolved oxygen. • 1 i I ; 1 I / ' "^A^^ I . 1 0.2 0.3 O.i. 0.5 0. Sodium Chloride Concentrati 0.7 0.8 0.9 {moles/liter) Fig. 7. The effects of sodium chloride on the growth of Chaetophoma confluens. 32 Great Basin Naturalist Vol. 37, No. 1 there is more vegetation in Farmington Bay than there was in 1959. The same general pattern of vegetation decline is occurring in the southern portion of Farmington Bay and in the marshland to the south that is not shown in the illustration. The decline ap- pears to be occurring slowly, and it would take several years of similar conditions to eradicate the emergent marsh vegetation. Several parameters of Farmington Bay's environment could cause or enhance a de- cline of the marshland vegetation. Among these are adverse temperature ranges, pH, phenols, oil and grease, fungus pathogens, and salt water intrusion from the Great Salt Lake. All of these potential pathogens were examined in this study. The effects that each agent or factor might have on the veg- etation, alone or in combination, were con- sidered. Since no unusual temperature changes have been recorded at Farmington Bay, temperature can be immediately eliminated as the cause of vegetation decline. Marshes Sodium Chloride Concentration (moles/liter) Fig. 8. The effects of sodium chloride on the growth of cattails and Chaetophoma confluens. in Red Butte Canyon that experience a sim- ilar range of air and water temperatures have shown no signs of diseased or dying vegetation. However, the cattails at Farm- ington Bay are dying during the warmest part of the summer, and it is possible that high temperatures might be enhancing the effects of some other stress to which the cattails have already been subjected. Soil and water pH greatly affect the availability of plant nutrients. The ideal pH range for most crop plants is from 6.5 to 7.5 (Sprague 1964). Average values above 8.0 or below 5.0 would be necessary to cre- ate serious nutrient deficiencies and cause disease. A pH range of 5.0 to 9.0, with an average pH around 8.0, was measured at Farmington Bay. Chlorosis (yellowing of the leaves), which is indicative of many nutrient deficiencies, and other characteristic symp- toms that commonly occur in response to the lack of essential elements, were not ob- served (Treshow 1970). In addition, no sta- tistical correlation could be obtained be- tween pH and cattail growth as determined by height. In other studies conducted on Utah marsh plants, it has been shown that vegetation grows well in the pH ranges that occur at Farmington Bay (Jensen 1940, Nel- son 1954, Kaushik 1963). There is little pos- sibility, therefore, that pH is contributing to the vegetation decline. Analysis of the water at Farmington Bay for possible industrial pollutants revealed the presence of phenols, oil, and grease. Phenolic compounds in water resources are usually a result of pollution from oil refi- neries. Phenols also result from the break- down products of herbicides such as 2,4- dichlorophenoxyacetic acid (2,4-D) and 2, 4,5-trichlorophenoxyacetic acid (2,4,5-T) (Goerlitz and Brown 1972). The oil refi- neries upstream from Farmington Bay and the herbicide treatment of some grain crops with 2,4-D are the most probable sources of the phenols, oil, and grease in Farmington Bay. Phenol concentrations ranged from none to 0.008 mg/1. These concentrations are extremely dilute when compared to the concentrations of phenolic compounds (such as 2,4-D) that are known to be toxic to broad-leaved dicots (Crafts and Bobbins March 1977 Anderson: Cattail Decline 33 1962). Since the concentrations of phenols are dilute, there is little possibility that phe- nols are causing the vegetation decline. However, phenolic compounds might be contributing to the decline by inhibiting the reestablishment of cattail seedlings. McNaughton (1968) has shown that phenols produced by Typha latifolia have an auto- toxic effect on seed germination and seed- ling growth. Oil and grease measurements ranged from 0.6 mg/1 to 8.0 mg/1. These concentrations are also too dilute to cause any vegetation damage (Goerlitz and Brown 1972). Study of the total salt content, or salinity, of the soil and water of Farmington Bay re- vealed that the existing salinities were in a range that is known to be toxic to many plant species (Richards 1954). Field studies showed that cattails are weakened or killed when they are exposed to soluble salt con- centrations greater than 5.0 gm/1. Penfound and Hathaway (1938) have also found that Typha latifolia plants in Louisiana marshes do not occur where salinities are above 5.0 gm/1. McMillan (1959) found that T. lati- folia grown in hydroponic tanks could not tolerate 10 gm/1, and Kaushik (1963) found that cattails from Ogden Bay Waterfowl Management Area were killed when sub- jected to a soluble salt concentration of 0.9 gm/1. Symptoms of the elevated salt con- centrations on cattails include stunted growth, leaf tip necrosis (burning), and oc- casional browning of the entire plant. Fur- thermore, the salt concentrations that cause a physiological stress to cattails facilitate the growth of a decomposition fungus, Chaetophoma confluens. This fungus causes a rot consisting of irregularly scattered le- sions on the surface of the rhizomes and was consistently isolated from rhizomes of declining plants. At the beginning of this study, it was thought that the fungus might be a primary cause of the decline. How- ever, elevated soluble salt concentration ap- pears to be the primary cause. The source of the elevated salt concentra- tion at Farmington Bay is undoubtedly the Great Salt Lake. Salinities in Farmington Bay increase and decrease following the ris- ing and falling of the lake. Salinity measure- ments of the water entering Farmington Bay from the Jordan River show concentra- tions of salt between 0.1 gm/1 and 0.6 gm/1. These concentrations do not seem to be increasing the salinity of Farmington Bay. The Great Salt Lake, on the other hand, with a salt concentration of 224 gm/1, is a natural source of salt for bordering marshlands because the brines are known to diffuse through and under the dikes around the marshes. The periodic fluctuations of the Great Salt Lake allow for a natural con- trol of the density of cattail stands. This could be either beneficial or detrimental, depending on the frequency and extent of the fluctuations. In order to maintain a de- sirable stand density, it is suggested that the salinity be monitored and controlled. Literature Cited American Public Health Association. 1971. Stand- ard methods for the examination of water and wastewater. American Pubhc Health Associ- ation, Inc., New York. 874 pp. Anson, A. 1975. Photogranimetry as a science and as a tool (an index). Photogramm. Eng. 41(2):225- 236. Arnon, D. I. AND D. R. HoAGLAND. 1940. Crop pro- duction in artificial solutions and in soils with special reference to factors influencing yields and absorption of inorganic nutrients. Soil Sci. 50:4&3-484. Barnett, H. L. 1956. Illustrated genera of imperfect fungi. Burgess Publishing Co., Minneapolis, Minnesota. 218 pp. Barron, G. L. 1968. The genera of hyphomycetes from soil. Williams and Wilkins Co., Baltimore, Maryland. .364 pp. Bolen, E. G. 1964. Plant ecology of spring-fed salt marshes in western Utah. Ecol. Monogr. .34(2): 143-166. Bourn, W. S. and C. Cottam. 1939. The effect of lowering water levels on marsh wildlife. Trans. North Am. Wildl. Conf. 4:343-350. Crafts, A. S. and W. W. Robbi.ns. 1962. Weed con- trol. McGraw-Hill Book Co., New York. 660 pp. Goerlitz, D. F. and E. Brown. 1972. Methods for analysis of organic substances in water. Tech- niques of water-resources investigations of the U.S.G.S., Book 5, Chapter A3. U.S. Government Printing Office, Washington, D.C. 40 pp. Haller, W. T. 1974. Effects of salinity on growth of several aquatic macrophytes. Ecology 55(4):891- 894. Jensen, G. H. 1940. The relation of some physical and chemical factors of the soil to the productivity and distribution of certain waterfowl food 34 Great Basin Naturalist Vol. 37, No. 1 plants at the Bear River Migratory Waterfowl Refuge. Master's thesis, Utah State University- 30 pp. JoHANSEN, D. A. 1940. Plant microtechnique. McGraw- Hill Book Co., New York. 523 pp. Kaushik, D. K. 1963. The influence of salinity on the growth and reproduction of mansh plants. Ph.D. dissertation, Utah State University. 123 pp. Levitt, J. 1972. Responses of plants to environmental stresses. Academic Press, New York. 697 pp. McMillan, C. 1959. Salt tolerance within a Typha population. Am. J. Bot. 46(7):521-526. McNaughton, S. J. 1968. Autotoxic feedback in rela- tion to germination and seedling growth in Typha kitifolia. Ecology 49(2):367-369. Nelson, N. F. 1954. Factors in the development and restoration of waterfowl habitat at Ogden Bay Refuge. Utah State Department of Fish and Game, Salt Lake City, Utah. 87 pp. Nelson, N. F. and R. H. Dietz. 1966. Cattail control methods in Utah. Utah State Department of Fish and Game, Salt Lake City, Utah. 31 pp. Penfound, W. T. and E. S. Hathaway. 19,38. Plant communities in the marshlands of southeastern Louisiana. Ecol. Monogr. 8(l):l-56. Richards, L. A. 1954. Diagnosis and improvement of saline and alkali soils. U.S. Department of Agri- culture Handbook No. 60, U.S. Government Printing Office, Washington, D.C. 160 pp. Seher, S. J. and p. T. Tueller. 1973. Color aerial photos for marshland. Photogramm. Eng. 39(5):489-499. Snedecor, G. W. 1957. Statistical methods. Iowa State College Press, Ames, Iowa. 534 pp. Sparrow, F. K. 1943. Aquatic phycomycetes. Univer- sity of Michigan Press, Ann Arbor, Michigan. 785 pp. Sprague, H. B. 1964. Hunger signs in crops. David McKay Co., New York. 461 pp. Treshow, M. 1970. Environment and plant response. McGraw-Hill Book Co., New York. 422 pp. United States Geological Survey. 1975. Water re- sources data for Utah. Part I, Surface Water Records. U.S.G.S. Water Resources Division, Salt Lake City, Utah. 505 pp. OSTEOLOGY AND MYOLOGY OF THE HEAD AND NECK REGIONS OF CALLISAURUS, COPHOSAURUS, HOLBROOKIA, AND UMA (REPTILIA: IGUANIDAE) Douglas C. Cox' and Wilmer W. Tanner^ Abstract.- A detailed study of the anterior osteology and myology of Callisatmis, Copho.saurus, Holbrookia, and Vma reveals the phylogenetic relationships among the sand lizards. An SPSS discriminant analysis of os- teological characters combined with myological characters indicates that Callimiirtis is most primitive, Cop- hosaurus and Holbrookia are most closely related, and Uma is the most distinct of the sand lizard genera. Be- cause of close relationships between Cophosaurus and Holbrookia, it is postulated that earlessness evolved once, and Cophosaurus is returned to synonymy under Holbrookia. Blainville (1835) wrote the first descrip- tion of a sand lizard and named it Calli- saurus draconoides. Since then various au- thors have pubhshed articles concerning sand lizards. Girard (1851) described Hol- brookia maciilata, Trochel (1852) described Cophosaurus texanus, and that same year Baird and Girard synonymized Cophosaurus with Holbrookia, providing the name Hol- brookia texana, which then remained un- changed for over 100 years. Subsequently, Baird (1858) described Uma notata. By 1858 all genera represented in the sand lizard group had been described. Since then new species and subspecies, as well as new combinations, have been added by var- ious authors as follows: Bocourt (1874) Hol- brookia elegans- Cope (1880, 1883, 1894, 1895, 1896, 1900) Holbrookia lacerata, Hol- brookia maculata fkwilenta, Uma scoparia, Uma inornata, Uma rufopuncatata, Calli- saurus crinitus, Callisaurus rhodostictus, Holbrookia maculata maculata, Callisaurus draconoides ventralis, and Callisaurus ven- tralis gabbii; Stejneger (1890) Holbrookia maculata approximans and Holbrookia maculata lacerata; Richardson (1915) Calli- saurus ventralis myurus; Dikerson (1919) Callisaurus carmenensis; Schmidt (1921, 1922) Holbrookia maculata campi, Hol- brookia pulchra, Holbrookia dickersonae, and Callisaurus ventralis inusitatus; Schmidt and Bogert (1947) Uma exsul; Barbour (1921) Holbrookia thermophila; Harper 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 'Life Science Museum, Brigham Young University, Provo, Utah 84602. (1932) Holbrookia propinqua stonei; Smith (1935, 1943, 1946:137, 145) Holbrookia ele- gans thermophila, Holbrookia elegans ele- gans, Holbrookia bunkeri, Holbrookia macu- lata ruthveni, Holbrookia maculata dickersonae, Holbrookia maculata pulchra, and Holbrookia maculata thermophila; Lins- dale (1940) Callisaurus draconoides myurus and Callisaurus draconoides gabbii; Heifetz (1941) Uma notata notata; Bogert and Dor- som (1942) Callisaurus draconoides brevipes; Smith and Burger (1950) Holbrookia prop- inqua propinqua and Holbrookia propinqua piperata; Peters (1951) Holbrookia texana texana and Holbrookia texana scitula; Axtell (1956) Holbrookia lacerata, Holbrookia lacer- ata subcaudalis and Holbrookia maculata perspicua; Smith and Cochran (1956) Calli- saurus draconoides rhodostictus; and Wil- liams, et al. (1959) Uma paraphygus. Some summaries, reviews, checklists, and comparative studies have also been written. Cope (1896) synonymized Uina and Calli- saurus in a short paper di.scussing the genus Callisaurus. He recognized Uma again in his large work on the crocodilians, lizards, and snakes of North America (1900) and recognized one species and three subspecies of Callisaurus. Smith (1946:137, 145), in his "Handbook of Lizards," recognized in Callisaurus one species and ten subspecies and stated (p. 145): 'The whole group of Callisaurus of 35 36 Great Basin Naturalist Vol. 37, No. 1 western United States is in need of revision. The subspecies are not adequately charac- terized, nor are their ranges well worked out. There is very little information on the life history." Although we now know much more about the life history of CaUisaurus, it is still in need of a comprehensive tax- onomic review and remains a monotypic genus. (A careful revision for CaUisaurus has not yet been attempted.) The genus Holbrookia has had two revi- sions. Schmidt (1922) made the first; a sec- ond was by Axtell (1958). Schmidt (p. 709) stated: "The taxonomy of the North Ameri- can lizards of the genus Holbrookia Girard offers one of the most interesting and diffi- cult problems in North American herpeto- logy." He recognized several species and states that (p. 712) ^'Holbrookia is obviously di- rectly related to CaUisaurus, from which it differs only in the concealed tympanum and with which it agrees in general features of color pattern and scutellation." Smith (1946:137, 145) doubts that a prac- tical means of characterizing the species ex- ists and believes that "until such a means is found there will remain indefinitely a prob- lem in defining the ranges of the several subspecies, or in defending their actual validity. Accordingly the genus, par- ticularly the maculata group, merits a care- ful study perhaps more than any other in the United States." Axtell (1958) recognized only three spe- cies, HoUjrookia lacerata, H. propinqua, and H. maculata. He considered H. texana to be a species of the genus CaUisaurus and therefore did not discuss it. He did list two subspecies for lacerata, two for propinqua, and 11 for maculata. Uma has been reviewed by Heifetz (1941), Norris (1958), and Mayhew (1964b). Heifetz recognized U. notata notata, U. no- tata cowlesi, U. inornata, and U. scoparia. He also referred to the taxonomic confusion that exists because of erroneous type local- ities in this genus. In his checklist Schmidt (1953) recognized only one species (notata), with three subspecies {notata, inornata, and scoparia). Norris (1958) recognized Uma n. notata. U. n. rufopunctata, U. n. inornata, U. sco- paria, and U. exsul. He discussed the evolu- tion of Uma and its relationship to other sand lizards. The conflict relative to the classification of the notata-scoparia group was discussed by Mayhew (1964b). He recognized U. in- ornata, U. notata, and U. scoparia all as full species on the basis of temperature toler- ance and reproductive data. Peters (1951) reviewed Holbrookia texana {Cophosaurus texanus). He described two sub- species, but mentions little concerning rela- tionships with other sand lizards. Clarke (1965) revived Cophosaurus texan- us on the basis of behavioral data collected in a large comparative study of the sand fizard group. Ecological and behavioral studies con- cerning sand lizards have also been pub- lished by Burt (1931a, 1931b), Stebbins (1944, 1954, 1966), Ramsey (1948, 1949), Cagle (1950), Williams and Smith (1958), Axtell (1960), Lannom (1962), Carpenter (1963, 1967), Clarke (1965), Mayhew (1964a, 1964b, 1966), Pianka and Parker (1972), Tanner and Krogh (1975), and Judd (1974, 1975). The anatomy of these lizards has not been thoroughly studied. Earle, in a series of articles (1961a, 1961b, 1961c, 1962), de- scribed in detail the comparative anatomy of the middle ear of sand lizards. Stebbins (1943, 1944) described the nasal structures and some aspects of the ecology of Uma, then (1948) described the nasal structures of lizards in general, which included the sand lizards. Axtell (1958) described the osteology of Holbrookia and stated that it is essentially the same as found in all sand lizards. Eth- ridge (1964) studied the skeletal morphology of sceloporine lizards, which includes sand lizards, and compared their relationships. Savage (1958) studied Urosaurus and Uta and made remarks concerning related gen- era, which included sand lizards. A few ref- erences to sand lizards were made by Lar- sen and Tanner (1974) while studying Sceloporus, and Guttman (1970b) also refers to them in his electrophoretic study of the hemoglobins of sand lizards. He found that March 1977 Cox: Lizard Morphology 37 all genera possessed the same major and mi- nor protein components. Only portions of the osteology have been adequately treated; the myology is essen- tially untouched. The objectives of this study are to: (1) describe the skull osteology and branchiomeric myology of sand lizards; (2) identify osteological and myological characteristics that distinguish the sand liz- ard generic groups; and (3) determine more accurately the relationships between these groups. No attempt will be made to deal with the species and subspecies except as they relate to the generic phylogeny. Materials and Methods Specimens used were obtained from the Brigham Young University Life Sciences Museum (BYU), the California Academy of Sciences (CAS), and the Strecker Museum at Baylor University. At least four speci- mens from each of the four genera were used for osteological examinations. The myology of these 16 specimens, plus four additional individuals from each genus, were also used in developing the myological description. The following list indicates the material used: Callisaums draconoides gabhi from N.T.S., Nye Co., Nevada: (BYU) 2943, 2967, 3079, 40037; C. d. inusatatus from Tiburon Island, Sonora, Mexico: (BYU) 30175, 30176, 30178; C. d. splendidus from Isla Angel de la Guardia, Gulf of California, Mexico: (BYU) 41112; C. d. carmenesis from Baja California Sur, Mexico: (BYU) 41095, 41231; Cophosaunis texanus texanus from Chihuahua, Mexico: (BYU) 14339, 15712; C. t. scitula from Sierra Co., New Mexico: (BYU) 30512, 30513, 30515; C. t. scitula from Pima Co., Arizona: (BYU) 34331, 34336; Holbrookia lacerata from Giaraz Co., Texas: (CAS) 73979; H. maciilata approx- imans from Colonia Dublan, Mexico: (BYU) 11370, 17099; H. maculata hunkeri from Chihuahua, Mexico: (BYU) 15782, 15785, 15788, 15789; H. propinqua propinqua from Padre Island, Cameron Co., Texas: (CAS) 16187; Uma notata inornata from Riverside Co., California: (BYU) 3263, 3266, (CAS) 22824, 22826; U. n. cowlesi from Sonora, Mexico: (BYU) 30144, 30156; U. scoparia from San Bernardino Co., California: (BYU) 11389, (CAS) 42072. Skeletal material was prepared by careful dissection. Bones were cleaned with forceps and dissecting needles and soaked in Clorox bleach for several minutes to loosen soft tis- sues, after which hirther picking and clean- ing was done. Skulls were not allowed to dry, but were preserved in 70 percent EtOH to insure that cartilaginous skeletal elements could be examined. The Statistical Package for the Social Sci- ences (SPSS) discriminant analysis was uti- lized to aid in the identification of charac- ters where they were not already obvious. The statistical analysis will be described be- low. Myological examinations consisted of careful dissection in which each muscle was separated and its origin and insertion deter- mined. General morphology (shape, relative size, and position) of each muscle was also noted. Muscle comparisons are based on both origin-insertion and muscle morpho- logy. Only the branchiomeric muscles asso- ciated with the hyoid arch and the jaws are described. Osteology Sand lizard skulls have been examined in detail. The length and width of individual elements were measured with a Golgau ver- nier caliper and a five-millimeter mini-tool. Comparisons were made on the basis of ra- tios, as well as on the shape and position of each bone in relation to other articulating bones. The lower jaw and hyoid have been studied in the same manner. The skull is streptostylic, with a freely movable quadrate bone. In mature individ- uals the ethmoid region of the braincase proper is not ossified, but consists of carti- lage plates from which cartilaginous rods extend dorsad to add to and support the membranes that protect the brain. Eyes are large, and only a thin sheet of cartilage sep- arates them medially; there is no apparent area of ossification, even in mature individ- uals. The sand lizard skull follows the basic plan of Sceloporine lizards as described by 38 Great Basin Naturalist Vol. 37, No. 1 Ethridge (1964) and Larsen and Tanner (1974, 1975); there is a great deal of unifor- mity within the sand lizard group. We will not give detailed descriptions of each bone here; however, these data are available to those who may desire them. We will, how- ever, describe the variations from the Sce- loporine skull as well as the differences be- tween the sand lizard genera. The following is a list of the skeletal ele- ments measured and described (they are il- lustrated on Figs. 1, 2, 3, 4): Basioccipital Supraoccipital Vomer Maxilla Frontal Parietal wing Quadrate Surangular Coronoid Orbit Suprateinporal Basisphenoid Pterygoid Palatine Nasal Infraorbital fossa fossa Jugal Postorbital Dentary Splenial Hyoid Posttemporal fossa Exoccipital Ectopterygoid Premaxilla Prefrontal Parietal Squamosal Articular Angular Temporal fossa The sand lizard skulls differ from the gen- eral sceloporine skull in that they lack the lacrimal and the postfrontal bones. They differ among themselves in several ways, but most of the variance seems to be associ- ated with the posterolateral position of the skull. The discriminant analysis selected 116 ratios that could aid in discriminating the four genera. Six measurements were found to be important. Every ratio selected in- cluded at least one of these six. They are listed along with their frequency of use: squamosal width (26), quadrate length (26), mandible length (20), hyoid length (20), mandible width (19), and maxillarv length (15). Representative ratio comparisons are giv- en to illustrate the separation and relation- ships of genera. Figures (1-4) of the lateral, dorsal, and ventral aspects of the skull are also presented, including the lateral and me- dial views of the lower jaw and the dorsal view of the hyoid. Earl (1961a, 1961b, 1961c, and 1962) re- ported on the osteological variations in the ear and indicated that all four groups were distinct, with Cophusaurus midway between CalUsaunis and Holhrookia. One of the first variations noticed, as skulls were being prepared, was that Co- phosaurus and Holhrookia had a para-occipit- al process directed anterolaterally. That of CaUisaurus and Lhna is directed lateral or slightly posterior. As a result of this varia- tion, variations in the length and width of the quadrate, squamosal, jugal, and postor- bital were noted. In lhna and CaUisaurus the ventral articulating process is ventral to the basioccipital, whereas in Cophosaurus and Holhrookia the quadrate is much short- er. Uma and CaUisaurus also have a wider squamosal, and the squamosal and jugal just meet. In the earless genera the jugal pushes in between the squamosal and postorbital. In one Uma individual the squamosal and jugal failed to meet, and the postorbital had pushed in between them. Some other variations (such as the configuration of su- tures between the frontal and parietal, the frontal and nasals, the basisphenoid and ba- sioccipital, and the maxilla and premaxilla, as well as proportional variations in the maxilla, postorbital, and premaxilla) were also found, but they were slight and had no significance. These are attributed to individ- ual variation on the species or subspecies level. Myology The myology of the neck and throat re- gion has been carefully studied. Axtell (1958) pointed out that the inter- mandibularis muscle in the most anterior re- gion of the throat is degenerate. The gen- ioglossus is therefore the most ventral muscle in that region. This condition is con- stant throughout the sand lizard genera. The variations within the sand lizard genera are found in the intermandibularis muscles, depressor mandibularis, constrictor colli, and the omohyoideus-sternohyoideus complex. The intermandihularis anterior super- ficiaUs (Figs. 5, 6) is a small muscle. In lhna, it is distinctly fan shaped. In CaUi- saurus and Holhrookia it is not fan shaped, and Cophosaurus shows a slight amount of fanning. In Holhrookia the fibers of this March 1977 Cox, Tanner: Lizard Morphology 39 Fig. 1. The skull osteology of Callisaurus BYU 3079: A. lateral view of skull; B. lateral view of lower jaw; C. medial view of lower jaw; D. dorsal view of skull; E. ventral view of skull; F. dorsal view of hyoid. The symbols used are: an-angular ch-ceratohyal ju-jugal pr-parietal ar-articular co-columella mx-maxilla pt-pterygoid bh-basihyal cr-coronoid na-nasal qu-quadrate bo-basioccipital de-dentary pa-palatine sp-splenial bp-basipterygoid ec-ectoptorygoid pf-prefrontal sq-squamosal bs-basisphenoid fr-frontal pm-premaxilla sr-surangular cbl-ceratobronchlal I gh-glossohyal po-post orbital vo-vomer cbll-ceratobronchial II hh-hypohyal 40 Great Basin Naturalist Vol. 37, No. 1 Fig. 2. The skull osteology of Uma BYU 3266: A; B; C; D; E; F; same as Fig. 1. The symbols used are the same as Fig. 1. March 1977 Cox, Tanner: Lizard Morphology 41 Fig. 3. The skull osteology of Holbrookia BYU 15783: A; B; C; D; E; F; same as Fig. 1. The symbols used the same as Fig. 1 42 Great Basin Naturalist Vol. 37, No. 1 Fig. 4. The skull osteology of Cophosaurus BYU 30518: A; B; C; D; E; F; same as Fig. 1. The symbols used are the same as Fig. 1. March 1977 Cox, Tanner: Lizard Morphology 43 Fig 5 Lateral view of head and neck musculature; superficial depth. A. Lhna BYU 3263; B. Callisaurus BYU 2943- C. Holbwokia BYU 15782; D. Cophosaurus BYU 30512. The symbols used are: am-adductor mandibulans externus medius; as-adductor mandibularis externus superficialis; au-auditory meatus; cc-constrictor colli; dm- depressor mandibularis; gg-genioglossus; iap-intermandibularis anterior profundus; ias-mtermandibulans anterior superficialis; ip-intermandibularis posterior; la-levator angularis oris; mhl-mandibulohyoideus I. 44 Great Basin Naturalist Vol. 37, No. 1 las iap 99 mh II mh I pm d m .cc sh I sh II oh Fig. 6. Ventral view of throat musculature; superficial layer at left and first depth at right. A. Uma BYU 3263; B. Callisauru-s BYU 2943; C. Ilolbrookia BYU 15782; D. Cophosaurus BYU 30512. The symbols used are: cc-con- strictor colli; dm-depressor mandibularis; gg-genioglossus; iap-intermandibularis anterior profimdus; ias-mter- mandibularis anterior superficialis; ip-intermandibularis posterior; mhl-mandibulohyoideus I; mhll-mandibulo- hvoideus II; oh-omohyoideus; prn-pterygomandibularis; shi-sternohyoideus I; shil-stemohyoideus II. March 1977 Cox, Tanner: Lizard Morphology 45 muscle pass posteromedial, whereas in the other three genera they pass transversely or fan out. The intemnandihukiris anterior profundus (Figs. 5, 6) occupies a position posterior and dorsal to the intermandibularis anterior su- perficialis, and anterior and dorsal to the in- termandibularis posterior. In Uma the fibers extend medially, with only a few fibers at each end fanning out. In Cophosaurus it is distinctly fan shaped, and in Callisaurus it fans out to a lesser degree. In Holbrookia it is slightly fan shaped, with most fibers slant- ing posteriorly. There were two areas of variation seen in the neck musculature. The constrictor colli (Figs. 5, 6) is the su- perficial muscle of the neck, originating in the dorsolateral fascia of the neck and in- serting in the ventral raphe of the throat posterior to the intermandibularis posterior. It is one or two muscle fibers thick, rather narrow, and the origin is broader than the insertion. In all four genera the muscle fibers of the constrictor colli reach the mid-throat area, but they originate in a facia without reach- ing the dorsal skeletoginous septum. The constrictor colli is widest in Uma. It covers from the posterior edge of the tym- panum to the shoulder, nearly covering the depressor mandibularis completely. In the other three genera it is more straplike and only fills half the space between the posteri- or edge of the tympanum and shoulder, being centered in this area. The depressor mandibularis (Figs. 5, 6) originates in the mid-dorsal raphe, along the posterior borders of the parietal bone and parietal wings. The insertion has three slips; one inserts deep to the pterygomandibularis on the articular process of the lower jaw. Another passes superficially to the pterygo- mandibularis and the intermandibularis pos- terior; it inserts on the ventrolateral surface of the mandibular rami by interdigitating at right angles with these muscles. The third slip inserts on both sides of a tendon that extends dorsally from the articular process. The attachment of this slip to the articular is superficial to the deep slip mentioned above. In the earless lizards (Cophosaurus and Holbrookia) a portion of this muscle is expanded anteriorly to partially cover the tympanic cavity. The sternohyoideus I and omohyoideus are closely related, and there is considerable confusion in the literature concerning them. The position taken here is that those por- tions that originate on the scapula and cla- vicle are omohyoideus, and those portions that originate on the sternum are ster- nohyoideus. Both muscles are deep to the constrictor colli, and the episterno-cleidomastoideus. The muscle dorsal to them is the sterno- hyoideus II. The omohyoideus (Fig. 6) takes two forms; in Uma and most species of Hol- brookia it has a single head originating on the scapula and clavicle. In Callisaurus, Cophosaurus, and some Holbrookia the omo- hyoideus is divided for its entire length, connected only by a myocomma located midway between the origin and insertion, bisecting the muscle and binding the muscle fibers together. This myocomma also bisects the sternohyoideus I, and the two muscles are bound firmly together by it. The omohyoideus can be easily separated from the sternohyoideus posterior to the myocomma, but they cannot be distin- guished anterior to it except by position. The sternohyoideus I (Fig. 6) originates on the sternum and inserts on the proximal end of the ceratobranchial I and II. The myocomma is at its junction with the omo- hyoideus, and it unites the two muscles. The medial portion of the omohyoideus cannot be distinguished from the sternohyoideus an- terior to this myocomma in all genera. Statistical Analysis The osteology of the skulls was analyzed by taking 43 measurements from each skull and by calculating all possible ratios. The Statistical Package for the Social Sciences (SPSS) discriminant analysis was employed to determine which ratios were of value in distinguishing genera and if the lizards could be classified by using them. The theo- ry and use of discriminant analysis is de- scribed by Klecka (1975). From each skull 903 ratios were generated, and those suf- 46 Great Basin Naturalist Vol. 37, No. ficient to separate one genus from the other three were used. The discriminant analysis proceeds in a stepwise fashion by selecting the single best- discriminating variable, and then selects a second on the basis of its ability to improve the value of the discrimination criterion in combination with the first variable. Third and subsequent variables are similarly se- lected according to their ability to contrib- ute to further discrimination. A plot of the discriminant score in two dimensions, a ter- ritorial map, and a classification based on the preceding analysis are given. Range, mean, and standard deviation of ratios for each genus are also given, and examples are presented in Figs. 7 and 8 along with a plot of the ratios (Figs. 7-11) that illustrates rel- ative relationships between genera. These figures are only examples of the ratios, and were selected because they clearly demon- strate the results referred to below. A multivariate analysis of variance was performed to determine the significance of differences between the four genera. The comparisons were based on a non-orthogan- al set such that the differences between Uma and the other three genera were test- ed (comparison #1), the differences be- tween CaUisaurus and Cophosaums were tested (comparison #2), and, finally, the differences between Cophosaums and Hol- brookia were tested (comparison #3). Using 116 ratios selected in the discrimi- nant analysis, it is possible to classify the four genera. Of 21 tests, classification agreed with the present taxonomic system in 17 of them. The first classification error was in Test 7, where a CaUisaurus and a Cophosaums were both classified as Hol- brookia. Only three ratios were involved, and all three involved the squamosal width divided by the vomer width, palatine length, and palatine width respectively. These ratios were effective in separating Uma from the other three, but were not useful in distinguishing between the other three genera. The second classification error occurred in Test 8. Here a CaUisaurus and a Hol- hrookia were both classified as Cophosaums. Four ratios were involved, and each one used the squamosal width divided by the pterygoid width, the epipterygoid width, the hyoid length, and the hyoid width. Whereas Uma was well separated in Test 7, it was not well separated in Test 8. The most confused classification was in Test 15. Here one CaUisaurus and one Hol- brookia were both classified as Cophosaums, one CaUisaurus was classified as Uma, one Cophosaurus was classified as Holbrookia, and one Holbrookia was classified as CaUi- saurus. Two ratios were used, and they were derived from the mandible width di- vided by the frontal width and the nasal width. Uma is the most distinct in this test; however, the individual plot scores show CalUsa mrus 9.6875 to 12.3636 10.8422 1.1974 Callis aurus 4.5405 to 5.6667 5.0379 0.4923 Cophosa lurus 10.3333 to 13.8869 11.7222 1.5437 Cophos aurus 5.2963 to 5.6000 5.4190 0.1337 Holbroc Jcia 11.5000 5.4375 to 13.7500 to 7.1379 12.1583 6.1048 1.1751 0.7827 Holbro okia 4.7500 to 6.7647 4.0200 to 4.3500 5.5560 4.1624 0.9255 ^ 0.1373 Total 5.4375 to 13.8889 10.2069 2.7175 Total 4.0200 to 6.7647 5.0438 0.7361 Fig. 7. Ratio of the skull length /squamosal width used in Test #5 of the Discriminant Analysis. Range, mean, and standard deviation plotted for the ratio of skull length/squamosal width in the four current gen- era. Fig. 8. Ratio of the skull length /mandible width used in Test # 14 of the Discriminant Analysis. Range, mean, and standard deviation plotted for the ratio of skull length /mandible width in the four current gen- March 1977 Cox, Tanner: Lizard Morphology 47 Fig. 9. Range, mean, and standard deviation in the four current genera plotted for the ratios of: A. squa- mosal length /squamosal width; B. squamosal width /quadrate length; C. jugal length/squamosal width; D. squamosal width/mandible length. considerable variation, and there is no dis- crete grouping to distinguish the separate genera. The last confused classification was in Test 17. Here four ratios were used that in- volved the mandible width divided by the palatine length, palatine width, epiptery- goid length, and the mandible length. In this classification a Callisaurus and Co- phosaurus were both mistaken for Hol- brookia. The plot indicates a complete sepa- ration of Uma, but with an overlap of the other three genera. Over all 21 tests the centroids were sepa- rated on the average of the following dis- tances: Uma was separated from Callisaurus by 2.10 mm, from Cophosaurus by 2.44 mm, and from Holbrookia by 2.67 mm; Cal- lisaurus was separated from Cophosaurus by 1.19 mm and from Holbrookia by 1.58 mm; and Cophosaurus was separated from Hol- brookia by 0.92 mm. These centroids are derived from the discriminant functions, which in turn are derived from linear com- binations of the variables used in each test. The purpose for the discriminant analysis was to find ratios that would discriminate Holbrookia Fig. 10. Range, mean, and standard deviation in the four current genera plotted for the ratios of: A. quad- rate length /quadrate width; B. maxillary length /quadrate length; C. jugal length /quadrate length; D. skull length /quadrate length. Fig. 11. Range, mean, and standard deviation in the four current genera plotted for the ratios of: A. man- dible length/mandible width; B. jugal length /mandible length; C. skull length/jugal length. 48 Great Basin Naturalist Vol. 37, No. 1 between the groups; this goal was achieved. It is also desirable to determine how well each group is defined, and if it is signifi- cantly different from all other groups. The SPSS program does not provide this infor- mation, but it is possible, by studying the plots (Figs. 7-11), to see that the range of Callisaurus overlaps the range of Holhrookia and/or Cophosaurus in nearly all ratios plotted. The range for Uma is only occa- sionally overlapped by the other genera, and it is usually only Callisaurus that over- laps Uma. The multivariate analysis of variance was performed to determine the degree of sig- nificance between the four genera. The non- orthoganal set of comparisons was as fol- lows: Comp. Cal Coph. Hoi. Uma 1 1 2 -1 3 0 1 1 -3 1 0 0 1 -1 0 The degrees of freedom, F-ratios, and actual probability for each F-value are presented in Table 1. The differences between Uma and the other three genera are highly significant, but the differences between the remaining three genera are only slightly significant, and in many cases the differences are in- significant. Uma is therefore the best differentiated by these tests. The other three genera are close together, and although they can be distinguished on the computer, their degree of separation is slight. The probability of error in differentiating between Callisaurus and Cophosaurus is similar to the probability of error in differ- entiating between Cophosaurus and Hol- hrookia (average between 9 percent and 12 percent), verifying these statements. These data are based on 116 ratios that were found to be the best discriminators of the 903 ratios examined. It is noteworthy that by using the very best osteological dis- criminators, the genera are often not clearly separated. D ISCUSSION Sand lizards are a closely related group according to Smith (1946), Norris (1958), Axtell (1958), Etheridge (1964), and Clarke (1965). Smith referred to them as a closely knit group and listed their common charac- teristics as being oblique labials, granular dorsal scales, small head scales, a gular fold, a peculiar median triangular postmental, several prominent postlabials, much the same habits, similar habitat, scoop-shaped heads, flaring labial regions, and a counter- sunk lower jaw. Axtell (1958) listed 22 char- acters that he felt would describe ancestral sand lizards. With only a few exceptions, this list of characters might just as well ap- ply to other sceloporine lizards. Callisaurus, Cophosaurus, and Holhrookia exhibit many of the primitive characteristics mentioned by Axtell. Each genus, however, shows specializations that vary from those listed. Callisaurus is similar to Axtell's cri- Table 1. Summary of F-ratios from eight multivariate analyses of variance tests. Comparisons 1 2 3 Test df. F-ratio/Probability F-ratio/Probability F-ratio/Probability 1 6/7 7.797/0.79% 1.858/21.8% 4.034/4.51% 2 7/6 11.055/0.468% 4.623/4.05% 1.973/21.29% 3 7/6 25.922/0.044% 4.596/4.106% 31.230/0.026% 4 5/8 96.377/0.000062% 1.141/41.24% 2.827/9.25% 5 6/7 12..387/0.20% 16.8.30/0.077% 4.450/3.56% 6 5/8 40.174/0.00185% 2.100/16.76% 6.117/2.54% 7 6/7 10.251 /0.,35% 14.480/0.124% 2.147/17.02% 8 4/9 7.807/0.532% 4.670/2.572% 2.118/16.07% March 1977 Cox, Tanner: Lizard Morphology 49 teria for the primitive condition, with the most striking speciahzations being increased length of tail and limbs and a more slender body form. Holbrookia is also similar to an- cestral sand lizards except for the covered tympanum. Body form in Holbrookia is not as proportionately slender as it is in Calli- saurus, nor is the tail as long. The problem- atic Cophosaums has characteristics of both. Its body form, limbs, and tail approach those of Callisaurus, but it has a covered tympanum like Holbrookia. Uma has devel- oped, to a greater degree, the dorsovent- rally flattened body and the toe fringes. Callisaunis, Holbrookia, and Cophosaurus occupy generalized habitats, probably sim- ilar to that of the ancestral stock. Uma, on the other hand, is restricted to a sand dune habitat. Osteology Comparisons of skulls of Ctenosaura (Oelrich 1956), as well as general accounts of reptile osteology by Williston (1925), Ro- mer (1956), and Avery and Tanner (1964, 1971) indicated that osteological character- istics of iguanid lizard skulls are generally stable within a genus. Studies on Sauro- malus (Avery and Tanner 1964), Croto- phytus (Robison and Tanner 1962), and Ctenosaura (Oelrich 1956) portrayed the ap- parent general stability of osteological char- acters found in iguanid skulls; however, skulls of Callisaurus, Cophosaurus, Hol- brookia, and Uma observed in this study are peculiar to iguanid skulls as portrayed by the above authors because there is stability within the sand lizard genera as a group rather than within a single genus. The lac- rimals and postfrontals are also absent in all the sand lizards, a condition not generally found in iguanid skulls, except that Jenkins and Tanner (1968) found that two species groups of phrynosoma also lack these skull bones. Etheridge (1964) pointed out the ab- sence of the lacrimals and postfrontals in sand lizards. Deviation by sand lizards from the gener- al iguanid skull, particularly evidenced by the fusion or loss of the lacrimal and post- frontal bones, is evidence supporting the hy- pothesis that sand lizards are highly special- ized among the iguanids. In discussing the osteology of Holbrookia, Axtell (1958:24) stated: In general the osteology of the genera Calli- saurus and Vma corresponds closely with that of Holbrookia, so this discourse may apply just as well to the entire sand lizard section of the family iguanidae. Development of the covered tympanum appears to be related to the osteological variations observed in this study. The quad- rate, squamosal, paraoccipital process, and mandible are all adjacent to the ear, and all exhibit variations (Figs. 1, 2, .3, and 4). With the loss of the external ear, the quadrate is reduced in size, and the paraoccipital proc- ess is directed forward, the mandible is shortened, the squamosal is narrowed, and the hyoid is also shortened. These modifica- tions of the skull are apparent in earless sand lizards and are less modified in sand lizards having an external ear. Callisaurus and Uma are alike in that the paraoccipital process is directed caudad, the quadrate proportionately larger, and the mandible long and wide in comparison to the earless forms (Figs. 1, 2, 3, and 4). In Uma the de- gree of development of these characters is different because the squamosal is wide. A ratio of skull length divided by squamosal width shows: (a) Uma ranging from 5.4375 to 7.1379; (b) Callisaurus ranging from 9.6975 to 12.3636; (c) Cophosaurus ranging from 10.3333 to 13.8889; and (d) Holbrookia ranging from 11.0833 to 13.7500 (from Fig. 7). The quadrate and mandible length and width are similarly enlarged in Uma when compared to the other groups. These differ- ences are sufficient to permit the computer to distinguish between these measurements in Uma and those same characteristics found in Callisaurus and the earless group. There is some variation in the position and articulation between the squamosal, jugal, and postorbital. In all except some Uma individuals, the jugal reaches the squa- mosal. In Callisaurus it edges between the squamosal and postorbital. In Holbrookia and Cophosaurus the degree of encroach- ment is increased. This may be a result of the forward direction taken by the para- 50 Great Basin Naturalist Vol. 37, No. 1 occipital process and the shortening of the jaw. Uma is unique in that the jugal fails to reach the squamosal in some individuals, in which case the postorbital is found wedg- ing between them (Fig. 3). In other Uma individuals a mere contact is made without any overlap of the squamosal and jugal. That these variations exist is verified by the statistical analysis. The SPSS discrimi- nant analysis was able to identify character- istics that were capable of separating the genera into distinct groups. However, the analysis of variance points out that differen- ces between Callisaurtis, Copho.saurus, and Holhrookia (as identified by SPSS) are insig- nificant and that Ihna is indeed a distinct group. The F-ratios derived from the analy- sis of variance illustrate these facts. The tests comparing Uma with the other three genera have high F-values (96.377 to 7.797) and consequently low probability of making classification errors (from 0.000062 percent to 0.79 percent). Therefore the differences between Uma and the other sand lizards are highly significant. In comparing Callisaurus with Cophosaurus, the F-values were found to be lower (16.830 to 1.141), and con- sequently the probability of classification er- ror is higher (41.24 percent to 0.077 per- cent). Only two of the eight tests were significant (below 2.5 percent probability of error). The tests comparing Cophosaurus with Holhrookia are similar to those com- paring Callisaurus and Cophosaurus. The F- ratios range from 31.231 to 1.973. The probability of error is again higher (21.29 percent to 0.026 percent). Low F-ratios and high error probability indicate that the dif- ferences between the three genera are in- significant. Myology The literature is void of studies dealing directly with myology of sand lizards. Earle (1961a, 1961b, 1961c, 1962) dealt with the middle ear and also touched on the myol- ogy in the ear region. In this study, we will deal only with the myology of the head and neck region. A comparison of the anterior anatomy of sand lizards shows some obser- vable differences that are discussed below (Figs. 5 and 6). The intermandibularis muscles show some variations that are useful in distinguishing the genera. Uma is most distinct, with its intermandibularis anterior superficialis mus- cle being fan shaped (Fig. 6). This condition is contrasted with that found in Holhrookia, where the transverse mandibulae muscles are so nearly parallel that the borders of each muscle are difficult to discern. Co- phosaurus and Callisaurus exhibit an inter- mediate condition with some fanning seen, but not to the extent seen in Uma. Co- phosaurus can be distinguished from Calli- saurus in that fibers of the intermandicularis anterior superficialis extend posteromedially, and in Cophosaurus, a portion of the in- sertion of the intermandibularis anterior profundus is found anterior to the super- ficialis, a condition not seen in Callisaurus. The depressor mandibulae also shows some variations. In the earless lizards there is an expanded anterior edge of this muscle, which partially covers the enclosed tympa- num. This muscle emerges from beneath the constrictor colli to insert upon the man- dible, with its most ventral fibers inter- digitating at right angles with the inter- mandibularis posterior. Furthermore, the depressor mandibulae fibers extend much further along the mandibular ramus than in the eared genera. In Cophosaurus these fi- bers reach the mandibular ramus with only a few interdigitating with the inter- mandibularis posterior. The omohyoideus is also variable in these genera. In the unfolding of the evolutionary development of the omohyoideus, a branch of the rectus cervicus originated on the sca- pula and inserted on the hyoid. In sand liz- ards it further developed into a complex of muscles. In Uma the omohyoideus has a second head that originates on the sternum. This then can be called sternohyoideus I. Another muscle, the sternohyoideus II, arises on the sternum, passes deep to the sternohyoideus I, and inserts on the pos- terodorsal edge of the ceratobranchial I. Avery and Tanner (1964) designated this muscle thyrohyoideus in Sauromalus, but, since it clearly originated on the sternum in March 1977 Cox, Tanner: Lizard Morphology 51 sand lizards, it is designated sternohyoideus II in this study. A division of the oniohyoidens has oc- curred in CaUisaurus, Cophosaiirus, and two specimens of Holbrookia {Holbrookia lacerata CAS 73979 and Holbrookia m. approximans BYU 17099). Thus, in sand lizards the muscle may have one to three origins: the most lateral dorsal one is on the scapula, the second is on the clavicle, and the most me- dial is on the sternum and interclavicle. The omohyoideus also has two insertions: the most lateral division inserts on the distal two-thirds of the ceratobranchial I, and the second and medial heads unite and insert on the proximal third of the ceratobranchial II and the basihyal. In Uma and for the most part in Holbrookia, such division of the omohyoideus has not occurred, and the in- sertion is continuous along the ceratobran- chial I, the basihyal, and the proximal third of the ceratobranchial II. These myological variations, although observable, are not of great magnitude and may not be sufficient to support generic status for CaUisaurus, Cophosaurus, or Holbrookia. In all there is an overwhelming myological similarity within these three genera, and they are ob- viously closely related to Uma. Phylogeny Norris (1958) and Axtell (1958) agreed that sand lizards began their radiation in early or middle Miocene. This is the time when the Sierra Madre Occidental Range of Mexico was being built by volcanism along the western and southern borders of the Mesa of Central Mexico (Schuchert 1935, Miller 1942). Axtell indicated that this vol- canism split the sand lizards into two groups, a Uma-Callisaurus group and a Hol- brookia prototype. Norris also believed the sand lizards were divided at this time, but indicated that Uma was isolated from the CaUisaurus- Holbrookia stock. Norris further indicated that Uma was subsequently split by continued mountain-building processes in the mid-Pliocene, giving rise to the exsul group and notata-scoparia stocks. He also postulated that it was during this same time that the CaUisaurus- Holbrookia stock was divided. The data available to us from this study tend to support Norris' concepts con- cerning the radiation of sand lizards. Ac- cording to Maslin (1952), "The basic as- sumption upon which all taxonomic practices rest is that similar organisms are related." If we assume that sand lizards have occupied similar habitats and have been subject to similar environments throughout their recent history, then we can assume that the degree of similarity be- tween groups is an indication of the close- ness of their relationship. Maslin (1952) also states that because internal characters are less variable, they are of much greater val- ue in establishing relationships than are ex- ternal characters. The internal characters considered here indicate a closer relation- ship between Holbrookia and CaUisaurus than between Uma and CaUisaurus, in- dicating that Uma was probably the first of the sand lizards to break away from the an- cestral line. Clarke (1965) examined the behavior and external morphology from the standpoint of 20 characters; in 14 of them Cophosaurus was like CaUisaurus, in 5 of them Co- phosaurus was unique, and in only one was Cophosaurus like Holbrookia. In discussing these comparisons he states: The distinctness of Cophosaurus is evident. It is intermediate in many features between CaUi- saurus and Holbrookia, with the data indicating a closer affinity to CaUisaurus than to Hol- brookia. The uniqueness of Cophosaurus is most clearly shown in the distinctness of the push-up pattern. On the basis of five characters, Clarke would separate Cophosaurus as a separate genus. These five characters are: (1) place- ment of the lateral bar: anterior for CaUi- saurus, posterior for Cophosaurus, and cen- tral for Holbrookia; (2) body shape: slender for CaUisaurus, intermediate for Co- phosaurus, and stout for Holbrookia; (3) middle ear: distinct with an external open- ing for CaUisaurus, distinct without an ex- ternal opening for Holbrookia, and inter- mediate without an external opening for Cophosaurus; (4) preferred body temper- ature: 39.2 C for CaUisaurus, .38.3 C for 52 Great Basin Naturalist Vol. 37, No. 1 Cophosaurtis, and 35.7-38.1 C for Hol- hrookia; and (5) push-up pattern: distinct in all three groups. In four of the five charac- ters the differences are only comparative, and do not indicate a clear-cut distinction; only in the fifth does Cophosauriis show a real distinctness. An alternate interpretation of these data would have to conclude that there is not enough difference to warrant generic status for Cophosaurtis; indeed, there also may not be enough difference to warrant generic status for Holbrookia. The variations de- scribed are of the kind and magnitude used in the descriptions of species. Clarke (1965) stated that the push-up pattern is the most distinct feature of Cophosaurtis. Carpenter (1963, 1967) described the same behavior for Uma, indicating that a genus is capable of supporting greater variation than Clarke has allowed for in Callisaurus, Cophosaurus, or Holbrookia. Guttman (1970b) also commented on Clarke's study, stating: A comparison of Callisaurus, Cophosaurus, and Holbrookia (Clarke 1965) indicated the great similarity among these genera. According to Clarke, the uniqueness of Cophosaurus is most clearly shown by its distinctive push-up pattern. A comparison of the display-action pat- terns of two species of Urosaurus (Carpenter 1962) or three species of Uma (Carpenter 1963) indicated to this writer that sufficient in- trageneric variation exists to refrain from estab- lishing a genus based on this display pattern. Guttman was reporting his electrophoretic study of sand lizards, in which he analyzed the hemoglobin components and found that they were all identical. This is highly un- usual, especially for different genera. Elec- trophoretic techniques have been of value in confirming taxonomic relationships. This has been demonstrated by Dessaur et al. (1962), Dcssauer (1966), Gorman and Des- sauer (1965), Gorman (1966), Maldonado and Ortez (1966), and Guttman (1970a and 1970b). The conclusion Guttman (1970b) came to, and the one that supports our con- clusions, was that the .sand lizards are more closely related than their present taxonomic status indicates. Simpson (1945) states: What is deplorable in splitting is the tenden- cy to raise the ranks of groups without need, that is, without gaining any practical advan- tage. One of the more evident symptoms of this tendency is the appearance of many monotypic groups in classification. The proposal resulting from this study and data examined from other studies would eliminate one monotypic genus and would provide a better indication of the close rela- tionships that are so evident in the sand liz- ards. Axtell (1958) believed that the sand liz- ards evolved under subhumid conditions, not greatly different than the conditions ex- isting today in the sand lizard range. He postulates that it was during the mid-Plio- cene that Holbrookia developed the covered tympanum. He then states, "The species previously known as Holbrookia texana, but which now appears to belong in the Calli- saurus line of evolution, has probably devel- oped the covered tympanum independ- ently." Axtell's phylogenetic tree for the sand lizards is presented in Figure 12. Earl (1961a, 1961b, 1961c, and 1962) in- dicated that Cophosaurus was intermediate between Callisaurus and Holbrookia in ear anatomy, but agreed with Axtell that ear- lessness evolved twice, reporting that re- ^ DIRECTION OF SPECIALIZATION 3, TYMPANUM COVERED SAND LIZARD PROGENITOR Fig. 12. Phylogeny of the sand lizards according to Axtell (1958).' March 1977 Cox, Tanner: Lizard Morphology 53 lated groups have the potentiaHty to devel- op identical clines, and under similar environmental conditions these identical clines may develop at different times and places. This concept was discussed by Mas- lin (1952), who put forth the idea while dis- cussing morphological criteria of phyletic relationships. Norris (1958) and Clarke (1965) also agreed with Axtell about the idea of separate earless evolution. The concept of two evolutions for earless- ness may be a major barrier to the under- standing of sand lizard relationships. Since we lack a fossil record of sand lizards, there is no way of knowing when or how such a character came about. It has been suggested (Earl 1961a) that it came about in response to the burrowing habit, but this is purely speculative, as there are many burrowing species that do not have a covered tympa- num (including Uma). A covered tympanum is not unique to "sand lizards " because the agamid genus Tympanocryptis in Australia is earless, and so are some of the members of Phrynocephahis. Phrynosoma has both eared and earless members. Norris's (1958) idea is acceptable when he says that CalU- saurus and Holbrookia split in the early Pliocene, with Holbrookia occupying the table lands of the mesa of Central Mexico, and Callisaurus having been isolated from Holbrookia before their radiation to the more northern habitats. The habitats of the Chihuahuan desert and Sonoran desert are similar; the draconoides and texanus groups would have had an excellent opportunity to parallel each other sufficiently to account for the external morphologic similarities. In- ternal structures are not as accessible to ex- ternal selective pressures and may, then, in- dicate more accurately the true relationships: that the texanus group is more closely related to Holbrookia than to Callisaurus, which it resembles through par- allelism. That earlessness may have evolved twice is possible, and it is most probable that there were separate evolutions for this char- acter in the genera Tympanocryptis, Phry- nocephalus, Phrynosoma, and sand lizards. To theorize that it evolved twice in the sand lizards is, however, a questionable con- cept. Earless sand lizards are closely related, their geographic ranges overlap, their habit- ats are similar, their food requirements are similar, the ear anatomy is similar, and we lack any evidence from fossil records that they diverged before the earless character arose. Where evidence indicates a close tax- onomic relationship, as it does here, the idea of a single evolution for the earless character is most plausible. It is very un- likely that the same character would evolve twice in the same way in two groups that are as closely related as are earless sand liz- ards. (A proposed phylogenetic tree is given in Figure 13.) These data, when added to that of earlier workers, seem to clearly indicate that sand lizards may best be represented by three genera: Uma, Callisaurus (as at present con- stituted), and Holbrookia. The characteristics that separate sand liz- ards into genera are few and not well de- fined when compared to distinctions be- tween other Sceloporine genera. The relationship between Callisaurus and Hol- brookia is especially close, with earlessness (and its associated skull characters) and body proportions being the most striking variants. Uma, on the other hand, appears to be well defined. It is felt that Holbrookia is, therefore, a recent derivative of Calli- ^ DIRECTION OF SPECIAIIZATION ^ TYMPANUM COVERED SAND LIZARD PROGENITOR Fig. 13. Proposed phylogeny of the sand lizards as dictated by data developed from this study. 54 Great Basin Naturalist Vol. 37, No. 1 saiirus, evolving earlessness and an adapta- tion for varied habitats in a relatively short time, perhaps since their separation in the late Pliocene. Sand lizards, particularly the species and subspecies, particularly in H. maculata, in- dicate that this group has recently under- gone adaptive radiation. If recent geological past has been correctly interpreted by re- cent paleontological findings (Etheridge 1961, Wells and Jorgensen 1964), the desert areas of today, extending from Texas to California, were very different as recently as 10,000 years ago, indicating that sand liz- ard adaptative radiation must be relatively recent. Evidence from internal morphology and geographical distribution indicates that ear- less sand lizards should remain as two close- ly related groups in the genus Holbrookia. Data from comparative skull and throat anatomy, if used alone, indicate a very close relationship between all genera in the sand lizard group, but it is felt that the dis- tinctions, however small, do indicate that Holbrookia has evolved from CaUisaums stock and has achieved sufficient distinctness to be given generic status. It is, therefore, proposed that sand lizards be classified as they were before Cophosaiirus was split off by Clarke (1965). This is as follows: Uma notata Baird Uma scoparia Cope Uma exsul Schmidt Uma pamphijgas Williams, Chrapliny, and Smith CaUisaums draconoides Blainville Holbrookia texana Troschell Holbrookia lacerata Cope Holbrookia maculata Girard Holbrookia propinqiia Baird and Girard Literature Cited Avery, D. F., and W. W. Ta.nner. 1964. The os- teology and myology of the head and thorax re- gions of the obesus group of the genus Sauro- maliis Dumeril (Iguanidae). Brigham Young Univ. Sci. Bull., Biol. Ser. 5(.3):l-30. 1971. Evolution of the iguanine lizards (Sauria Iguanidae) as determined by osteological and myological characters. BYU Sci. Bull., Biol. Ser. 12(3): 1-79. AxTELL, B. VV. 1956. A .solution to the long neglected Holbrookia lacerata problem and the description of two new subspecies of Holbrookia. Bull. Chi- cago Acad. Sci. 10(11):161-179. 1958. A monographic revision of the iguanid genus Holbrookia. Diss. Abstr. 19(6): 1476-1477. 1960. Orientation by Holbrookia maculata (Lacertilia, Iguanidae) to solar and reflected heat. Southwestern Naturalist 5(l):47-48. Baird, S. F. 1859. Description of new genera and spe- cies of North American lizards in the museum of the Smithsonian Institution (Uma). Proc. Acad. Nat. Sci. Philadelphia 10:253 (1858). Baird, S. F., and C. Girard. 1852. Holbrookia texana. Proc. Acad. Nat. Sci. Philadelphia 6:124. Barbour, T. 1921. A new lizard from Guaymas, Mexi- co. Proc. New England Zool. Club 7:79-80. Blainville, H. M. D. de. 1835. Description de quelques especes de reptiles de la Californie precedee de I'analyse dun systeme general d'herpetologie et d'amphibiologie. Nouv. Ann. Mus. Natn. Hist. Nat. Paris, (3)4:232-296. BocouRT, M. F. 1873-1897. Etudes sur les reptiles. Mission scientifique au Mexique et dans FAmerique Central-Becherches Zoologiques, (q.v.) part 3. Bogert, C. M., and E. E. Dorsom. 1942. .\ new lizard of the genus Callisaurus from Sonora. Copeia 1942(3): 17,3-175. Burt, C. E. 1931a. On the occurrence of a throat-fan in the sand lizard, Uma notata Baird, with notes on the adaptive specialization of the form. Copeia 1931(1): 1.5-16. 1931b. On the occurrence of a throat-fan in Callisaurus ventralis gabbii and two species of Crotaplnjtus. Copeia 1931(2):58. Cagle, F. R. 1950. Notes on Holbrookia texana in Texas. Copeia 1931(2):58. Carpenter, C. C. 1962. A comparison of the patterns of display of Urosaurus, Uta, and Streptosaurus. Herpetologica 18:145-152. 1963. Patterns of behavior in three forms of the fringe-toed lizards, Uma. Copeia 1963(2):406-412. 1967. Display patterns of the Mexican iguanid lizards of the genus Uma. Herpetologica 23(4): 285-293. Clarke, R. F. 1965. An ethological study of the igua- nid lizard genus Callisaurus, Cophosaurus, and Holbrookia. The Emporia State Research Studies 13(4): 1-66. Cope, E. D. 1880. On the zoological position of Texas. Bull. U.S. Nat. Mus., No. 17:1-51. 1883. Notes on the geograhical distribution of Batrachia and Reptilia in western North Amer- ica. Proc. Acad. Nat. Sci. Philadelphia 36:10-11. 1894. On the iguanian genus Uma Baird. Am. Nat. 28:4.34-435. 1895. On the species of Uma and Xantusia. Am. Nat. 29:938-939. 1896. On the genus Callisaurus. Am. Nat. .30:1049-1050. 1900. The crocodilians, lizards, and snakes of North America. Report U.S. Nat. Mus. for 1898, pp. 15.3-1270. March 1977 Cox, Tanner: Lizard Morphology 55 Dessauer, H. C. 1966. Taxonomic significance of elec- trophoretic patterns of animal sera. Rutgers Univ., Serai. Mus. Bull. 34:4-8. Dessauer, H. C, W. Fox, and F. H. Pauch. 1962. Starch-gel electrophoresis of transferrins, este- rases, and other plasma proteins of hybrids be- tween two subspecies of whiptail lizard (genus Cnemidaphoms). Copeia 1962:767-775. DicKERsoN, M. C. 1919. Diagnoses of twentv-three new species and a new genus of Lizards from lower California. Bull. Mus. Nat. Hist., 41(10):461-477. Earle, a. M. 1961a. The middle ear of Holbrookia maculata maciilata, the northern earless lizard. Copeia No. 1:68-74. 1961b. An additional note on the ear of Hol- brookia maculata. Copeia No. .3:355. 1961c. The middle ear of Holbrookia and Cal- lisaitrus. Copeia 1961(4):405-410. 1962. The middle ear of the genus Uma com- pared to those of other sand lizards. Copeia 1962(1): 185-188. Etheridge, R. 1961. Late Cenozoic glass lizards (Oph- isaurus) from the southern Great Plains. Herpe- tologica. 17(3): 179-186. 1964. The skeletal morphology and systematic relationships of sceloporine lizards. Copeia 1964(4):610. Girard, C. F. 1851. On a new American saurian rep- tile. Proc. Am. Assoc. Advmt. Sci., 4:200-202. Gorman, G. C. 1966. The relationships of Anolis of the Roquet species group (Sauria: Iguanidae), electrophoretic comparison of blood proteins. Comp. Biochem. Physiol. 19:845-853. Gorman, G. C, and H. C. Dessauer. 1965. Hemoglo- bin and transferrin electrophoresis and relation- ships of island populations of Anolis lizards. Sci- ence 150:1454-1455. GuTTMAN, S. I. 1970a. Hemoglobin electrophoresis and relationships within the lizard genus Sceloporus (Sauria: Iguanidae). Comp. Biochem. Physiol. 34:563-568. 1970b. An electrophoretic .study of the he- moglobins of the sand lizards, Callisaurus, Co- phosaurus, Holbrookia, and Uma. Comp. Bio- chem. Physiol. 34(3):569-574. Harper, F. 1932. A new Texas subspecies of the lizard genus Holbrookia. Proc. Biol. Soc. Washington (45): 15-18. Heifetz, W. 1941. A review of the lizards of the genus Ihna. Copeia 1941 (2):99-lll. Jenkins, R. L., and W. W. Tanner. 1968. Osteology and myology of Phrynosoma P. Platijrhinos Gi- rard and Phrynosoma d. hernondesi Girard. BYU Sci. Bull., Biol. Ser. 9(4): 1-34. JuDD, F. W. 1974. Intraspecific variation in blood properties of the keeled earless lizard, Hol- brookia propinqua. Herpetologica .30(1):99-102. Klecka, W. R. 1975. Discriminant analysis, pp. 434- 467. In. N. H. Nie, ed.. Statistical Package for the Social Sciences, second edition. McGraw- Hill Book Company, New York. Lannon, J. R. 1962. A different method of catching the desert lizards, Callisaurus and Vma. Copeia 1962(2):437-4.38. Larsen, K. R., and W. W. Tanner. 1974. Numeric analysis of the lizard genus Sceloparus with spe- cial reference to cranial osteology. Great Basin Nat. 34(1):1-41. 1975. Evolution of the sceloporine lizards (Iguanidae). Great Basin Nat. .3.5(1): 1-20. Linsdale, J. M. 1940. Amphibians and reptiles in Ne- vada. Proc. Am. Acad. Arts Sci. 73(8): 197-257. Maldonado, a., and E. Ortiz. 1966. Electrophoretic patterns of serum proteins of some West Indian Anolis (Sauria: Iguanidae). Copeia 1966(2): 179- 182. Maslin, p. T. 1952. Morphological criteria of phyletic relatioiLships. Systematic Zoology 1:49-70. Mayhew, W. W. 1964. Photoperiodic responses in three species of the lizard Uma. Herpetologica 20(2):95-113. 1964. Taxonomic status of California popu- lations of the lizard genus Uma. Herpetologica 20(3):170-ia3. 1966. Reproduction in the arenicolous lizard Uma notata. Ecology 47(1): 9- 18. NoRRis, K. S. 1958. The evolution and systematics of the iguanid genus Uma and its relation to the evolution of other North American desert rep- tiles. Bull. Am. Mus. Nat. Hist. 114(3):247-326, Figs. 1-17. 1975. Activity and thermal ecology of the keeled earless lizard, Holbrookia propinqua. Herpetologica 31(2): 137- 149. Oelrich, T. M. 1956. The anatomy of the head of Ctenosaura pectinata (Iguanidae). Misc. Pub. Mus. Zool. Univ. Michigan 94:1-122, 59 figs. Peters, J. A. 1951. Studies on the lizard Holbrookia texana (Troschel) with descriptions of two new subspecies. Occ. Papers Mus. Zool. Univ. Mich- igan. No. 537:1-20. PiANKA, E. R., AND W. S. Parker. 1972. Ecology of the iguanid lizard Callisaurus draconoides. Copeia 1972:493-508. Presch, W. 1969. Evolutionary osteology and relation- ships of the horned lizard genus Phrynosoma (Family Iguanidae). Copeia 1969(2):250-"275. Ramsey, L. W. 1948. Hibernation of Holbrookia tex- ana. Herpetologica 4(6):223. 1949. Hibernation and the effect of a flood on Holbrookia texana. Herpetologica 5(6): 125-126. Richardson, C. H. 1915. Reptiles of northwestern Ne- vada and adjacent territory. Proc. U.S. Nat. Mus. 48(2078): 403-435. Robison, W. G., and W. W. Tanner. 1962. A com- parative study of the species of the genus Crota- phytus Holbrook (Iguanidae), BYU Sci. Bull., Biol. Ser. 2(1): 1-31. Romer, a. S. 1956. Osteology of the reptiles. Univ. Chicago Press, Illinois, xxi, 772 pp., 248 figs. Savage, J. M. 1958. The iguanid lizard genera Uro- saurus and Uta with remarks on related genera. Zoologica 4,3(2):41-54. Schmidt, K. P. 1921. New species of North American lizards of the genera Holbrookia and Uta. Am. 56 Great Basin Naturalist Vol. 37, No. Mus. Novit. (22): 1-6. 1922. A review of the North American genus of Uzards Holbrookia. Bull. Am. Mus. Nat. Hist. I6(12):709-725. 1953. A checklist of North American amphibi- ans and reptiles. American Society of Icthyolog- ists and Herpetologists. University of Chicago Press, pp. 1-280. Schmidt, K. P., and C. M. Bogert. 1947. A new fringe-footed sand lizard from Coahuila, Mexico. Amer. Mus. Nov. No. 1339:1-9. ScHUCHERT, C. 1935. Historical geology of the Anti- Uean-Caribbean region, or lands bordering the Gulf of Mexico and the Caribbean Sea. John Wiley and Sons, New York, xxvi and 811 pp. Simpson, G. G. 1945. The principles of classification and a classification of mammals. Bull. Amer. Mus. Nat. Hist., vol. 85, 811 pp. Smith, H. M. 1935. Notes on some Mexican lizards of the genus Holbrookia with the description of a new species. Kansas Univ. Sci. Bull. 22(8): 185- 201. 1943. The White Sands earless lizard. Zool. Ser. Field Mus. Nat. Hist. 24(30):339-.344. 1946. Handbook of lizards: Lizards of the United States and Canada. Comstock Publ. Co., Ithaca, New York. 1960. Evolution of chordate structure. Holt, Rinehart, and Winston, Inc., New York. XIV, 529 pp., .357 figs. Smith, H. M., and D. M. Cochran. 1956. Callisatirus draconoides rhodostictiis Cope revived for the western fringe-footed lizards, Callisatirus draco- noides gabbii Cope. Herpetologica 12(2): 153- 154. Smith, P. W., and W. L. Burger. 1950. Herpetologic- al results of the University of Illinois field expe- dition, spring 1949. Trans. Kans. Acad. Sci. S3(2): 165-175. Stebbins, R. 1943. Adaptations in the nasal passages for sand burrowing in the saurian genus Uma. Am. Nat. 77:38-52. 1944. Some aspects of the ecology of the igua- nid genus Uma. Ecol. Monogr. 14(3):31 1-332, Figs. 1-22. 1948. Nasal structures in lizards with reference to olfaction and conditioning of inspired air. Amer. Jour. Anat. 8.3(2): 183-222. 1954. Amphibians and reptiles of western North America. McGraw-Hill Book Co., New York. 1966. A field guide to western reptiles and amphibians. Peterson Field Guide Series. Houghton Mifflin Company. Stejneger, L. H. 1890. Reptiles of San Francisco mountain region. North American Fauna. (3):109-111. Tanner, W. W., and J. Khogh. 1975. Ecology of the zebra-tailed lizard Callisaurus draconoides at the Nevada Test Site. Herpetologica 31(3):302- 316. Troschel, F. 1852. Cophosaurus texanus neve Eide- chesengattung aus Texas. Arch. Naturgesch. 16(1)185, pp. 388-394. Wells, P. V., and C. D. Jorgensen. 1964. Pleistocene wood rat middens and climatic changes in Mo- jave Desert: A record of juniper woodlands. Sci- ence 143(3611): 1171-1 174. Williams, K. L., and H. M. Smith. 1958. Range and status of a Mexican earless lizard. Herpetologica 13(4):265-267, pi. 1. Williams, K. L., P. S. Chrapliwy, and H. M. Smith. 1959. A new fringe-footed lizard, Uma from Mexico. Trans. Kansas Acad. Sci. 62(2): 166-172, pis. 1-2. WiLLisTON, S. W. 1925. Osteology of the reptiles. Har- vard Univ. Press, Cambridge, xiii, 30 pp., 191 figs. HYDROLOGICAL FEATURES OF A CALIFORNIA COASTAL FEN Don C. Erman', Kenneth B. Robv% and Michael Eames' Abstract.— A unique fen peatland, on the CaUfomia coast, depends on six streams for its supply of flowing, mineral-rich water. About 25 percent of the water is supplied by surface streams that had average concentrations of Ca"'""'" from 0.15 meq/1 to 0.32 meq/1 and of Mg+ + from 0.25 meq/1 to 0.47 meq/1. During the dry sum- mer period oxygen concentration and pH were lower in the central area than in the ten margins. Water flow near the margins maintains high 0-2, pH, and cation content. The fen ranges from 4 m to over 11 m in depth, but the thickest peat layer is made up of very liquid, unconsolidated peat. In one year, an estimated 19.3 metric tons of suspended sediment entered tne 38.6 ha fen, which was equivalent to a 42 kg/ha loss from the total wa- tershed. Inglenook Fen near the coast of Fort Bragg, California, may be the southernmost example of a fen-type peatland on the Pa- cific coast and is the only recognized ex- ample on the California coast (Baker 1972). A fen is a physical land type that has dis- tinctive flora and vegetation. It is often con- cave in cross section, has strong inflows of mineral-rich waters, has near-neutral pH, contains high amounts of Ca and Mg ions, and is productive (Heinselman 1970). To emphasize the source of minerals and water, this land type is referred to now as a miner- otrophic (Heinselman 1970) or rheotrophic (Moore and Bellamy 1973) peatland. Fens evolve naturally over time to be- come bogs. Bogs (ombrotrophic peatlands) are characterized as being isolated from mineral-rich water, often convex in cross section, acid in pH, and unproductive (Heinselman 1970). Numerous examples of bogs exist along the Pacific coast of North America (Rigg and Richardson 1938). Sever- al are within a few km of Inglenook Fen in the "pygmy forest" on the upper coastal terraces (Rigg 1933, Jenny et al. 1969). The peatland type and rate of change from fen to bog is strongly influenced by certain critical or controlling factors. Water sources are the key element in peatland ev- olution (Heinselman 1970, Moore and Bel- lamy 1973). Little information is available on the important hydrologic features of In- glenook Fen. Our objectives were to deter- mine the sources and amounts of inflowing water, the nutrient status of the water, and the physical-chemical conditions in the fen. Study Area Inglenook Fen was described by Baker (1972), who included a floral list and some chemical conditions. The fen was formed by sand dunes blocking the stream outlet of a small valley on the second coastal terrace. A small 1.11 ha pond (Sandhill Lake) exists at the western edge, and radiating outward, especially to the east, are typical zones of marsh-fen vegetation— floating aquatics, emergents, sedge fen, and woody plants. The fen is 38.6 ha, of which about 4.6 ha are dominated by Cyperaceae fen and Ca- lamogrostis-CypersLceae-Menyanthes fen (cf. Baker 1972). The remainder is pre- dominantly woody vegetation (fen carr). Six streams drain five watersheds (A-E) into the fen (Fig. 1). The watersheds range from forest land (67 percent) and pasture to .some residential development in Area C and part of Area D. 'Department of Forestry and Conservation, University of California at Berkeley, Berkeley, California 94720. 'U.S. Forest Service, San Francisco, California. 'Department of Fisheries. State of Wa.shington, Olympia, Washington 98501. 57 58 Great Basin Naturalist Vol. 37, No. 1 M ETHODS AND MATERIALS M> Approximately twice a month from No- vember 1972 to November 1973, we meas- ured discharge, water temperature, pH, dis- solved oxygen, calcium and magnesium ion content, total suspended sediments, ortho- phosphate, and nitrate in the fen streams. Silica was measured occasionally. Aerobic limit, peat depth, O2, pH, and temperature were measured irregularly on the fen. Flow was measured in two ways. The first involved catching stream flow in a con- tainer of predetermined volume. The time required to fill a container was measured with a stopwatch. All streams except A^^^, Aj,^, and the outlet were measured this way as they emptied from culverts that passed beneath Highway 1 (Fig. 1). Flows of other streams that did not empty from culverts were determined by measuring a section for width, average depth, length, and average velocity. Velocity was determined by timing a float over a measured distance. Water temperature was determined with a thermistor or mercury thermometer, and oxygen concentration was determined with a YSI Model 51A Og-temperature meter or by the Winkler method. Small samples were taken for water pH determinations. Shortly after collection of the sample, pH was measured with a La- Motte colorimetric comparator (wide range pH 3-10, narrow range pH 5-8). Other water samples were taken for Ca- Mg ions and PO4-NO3 analysis. Samples for Ca-Mg analysis were preserved by addition of perchloric acid, filtered qualitatively be- fore analysis, and brought to a concentra- tion of 0.5 meq/1 of strontium chloride to reduce interference. Analysis was performed on a Perkin-Elmer Model 303 atomic ab- sorption flame spectrophotometer. Separate glass bottles were used for PO4- NO3 samples; they were stored for one to three weeks in a refrigerator prior to analy- sis. Determinations were made on a Hach #640 Direct Reading Colorimeter. Ortho- phosphate was measured by the Hach- Stannaver Method and nitrate (after nitrite adjustment) by the cadmium reduction method. An additional water sample was vacuum- filtered through a preweighed glass fiber fil- ter (approximately 0.3 jum pore size). The filter was then oven dried and reweighed to determine total suspended sediments. Silica was measured with a LaMotte col- orimetric comparator. The depth to which oxygen was present in the fen (aerobic lim- it) was determined by measuring the stain that developed on implanted redwood stakes (Erman 1973). The distance from fen surface to where staining begins (absence of oxygen causes stain) is the aerobic limit. To determine peat depth, we made cores along two transects, A and B (Fig. 1), of the fen with a Hiller-type peat borer. Fig. 1. Map of Inglenook Fen showing inlet streams (dashed hnes) and their watersheds (A to E). Lines a and b are transects for peat depth cores. The dotted area in the central fen region is predominantly sedge vegetation. Results and Discussion Stream Flows Mean surface flows are given in Table 1. Large seasonal fluctuations occurred in dis- March 1977 Erman, et al.: California Coastal Fen 59 Table L Yearly average (and standard deviations) of environmental conditions of outlet and inlets to Inglenook Fen, Mendocino Co., California. Environmental Factor Aso Ano Stream B C D E Outlet Discharge (1/sec) (S.D.) 4.93 (3.97) 3.26 (4.93) 14.6 (14.1) 9.17 (17.7) 9.03 (14.8) 4.25 (11.5) 179.8 (278) 0-2 (mg/1) (S.D.) 10.1 (1.24) 10.0 (2.13) 10.5 (0.94) 10.1 (1.31) 11.3 (1.66) 10.3 (1.01) 7.43 (1.92) O2 saturation (%) (S.D.) 94.4 (9.47) 89.3 (16.8) 101.2 (5.56) 95.1 (6.3) 101.5 (10.7) 95.8 (7.64) 70.5 (24.4) Temperature (°C) (S.D.) 12.7 (3.3) 12.0 (3.4) 12.8 (3.7) 12.7 (4.4) 113 (4.0) 12.9 (4.9) 13.4 (5.3) Median pH 6.5 5.6 6.7 6.8 6.8 6.3 6.6 Ca+2 (me/1) (S.D.) .16 (.09) .14 (.08) .18 (.08) .28 (.14) .32 (.15) .15 (.07) .62 (.25) Mg+2 (me/1) (S.D.) .27 (.06) .26 (.06) .33 (.09) .47 (.14) .47 (.14) .25 (.06) .47 (.10) Sediments (mg/1) (S.D.) 19.9 (22.9) 11.4 (12.6) 16.0 (20.8) 12.5 (11.6) 16.9 (18.6) 4.24 (3.50) 4.41 (5.59) PO, -P (mg/1) (S.D.) .09 (.08) .09 (.07) .08 (.05) .09 (.08) .09 (.08) .07 (.07) .13 (.08) NO3 - N (mg/1) (S.D.) .60 (.29) .23 (.26) .28 (.07) .13 (.07) .28 (.16) .06 (.03) .12 (.10) Conductivity'' (uMhos @ 25°C) 109 216 154 222 203 92 .366 ^Data from November only. charge with minimum flows from May to October. Stream B consistently had the highest flow (avg. 14.6 1/sec). The total out- flow from May to October (the dry season) averaged 47.6 1/sec (28-68 1/sec) and was more than twice the combined inflow (15.9 1/sec). Mean monthly discharge of the com- bined inlets and of the outlet are shown in Fig. 2. One of the most interesting results of this work is the relationship between inlet and outlet flows. R. Jackson (pers. comm.), who surveyed the fen for the Nature Con- servancy, surmised that the predominantly sandy soils of the watershed would result in much subsurface flow. For the entire sam- pling period the measurable surface inflow averaged only 25.2 percent (13.8-66.3 per- cent) of the measurable surface outflow. With one exception, during a rainstorm on 21 September 1973, measurable inflow nev- er exceeded 50 percent of outflow. Thus subsurface flow appears more important than surface flow in this area. Water, of course, also enters the fen from surrounding sand dunes and direct precipitation on the fen, and it leaves via evaporation and evap- otranspiration. These sources were not measured. By expanding the mean discharge value we estimated the total annual volume of water carried by each stream (Table 2). The estimated yearly inflow of all surface water was 1.43 million m^, compared to about 60 Great Basin Naturalist Vol. 37, No. 1 1549 K4a / \ FEN OUTLET 500 • \ ro" O . \ '^5 400 ■ "^SURFACE V Hi \ STREAMS \ DISCHAR( :\^ \ \ ; 200 - \ \ , / Z o 2 100 : / \ ^ __^ ■^ . ^^"--^j_^_ ^______. ^^^' , 1 1 1 1 1 • NOV DEC JAN FEB MAR APR MAY JUN JUL AUG SEP OCT 1972 1973 DATE Fig. 2. Estimated total discharge by month of all inlet streams combined and of the outlet. 5.67 million m^ of outflow. In Table 2 the discharge of each inflowing stream is shown as a percentage of the total inflow, and this value is compared with the percentage of the total watershed area for each of the streams. These results indicate differences between size of watershed and amount of surface flow. For example, based on surface flow. Watershed A accounted for 18.1 per- cent of the water volume while it included 42.7 percent of the total watershed. Water- shed B contained 12.4 percent of the area, but accounted for .32.3 percent of the total flow. Differences shown in these figures may be due to differences in vegetative cover, soil type, and slope gradient in the five wa- tersheds. Preliminary soil maps of the fen watershed show streams A to C drain Em- pire Sandy Loam, while streams D to E drain Hugo Sandy Loam. We have no data on subsurface flow, but it may be that when subsurface flow is added to surface flow, the total contribution of each stream is closer to its percentage of the total watershed. Mean monthly discharge of the combined inlets and of the outlet are shown in Fig. 2. Table 2. Comparison of surface stream inflow and area of watershed of Inglenook Fen. Total Area of Annual Flow % of Total Watershed % of Total Stream (M3 X 103) Flow (Hectares) Watershed Area A (No -(- So) 258 18.1 196.3 42.7 B 460 32.3 86.7 12.4 c 289 20.2 33.8 18.3 D 285 20.0 77.7 16.9 E 134 9.4 44.9 1.7 Total 1430 459.4 March 1977 Erman, et al.: California Coastal Fen 61 Jackson (pers. comm.) has suggested that only Stream E is seasonal. Stream A^^, was the only intermittent stream during the pre- sent study, although Stream E carried less than 0.3 1/sec from June to September. Temperature and Oxygen It was impossible to sample the streams at the same time of day on the various dates. Daily fluctuations in temperature are unknown for these streams. As expected, the winter water temperature (5-12 C) was lower than the summer water temperature (15-20 C), although temperatures below 10 C are unusual. High daytime air temper- atures are uncommon in summer because of frequent fog along the coast, and thus even small streams are not particularly warm. In general, oxygen concentration in inlet streams was near air saturation (average 95- 101 percent except for A^^^, which had low O2 (5.7-7.3 mg/1) at low flows (Table 1). Warmer water temperatures during low flows resulted in lower oxygen concentra- tions, as shown for Stream B in Fig. 3, than at other times of the year, although oxygen was still near air saturation. For most of the year, the outlet had significantly lower Og content than the inlets. The outlet is com- pared to Stream B in Fig. 3. This result is expected because the fen has a tremendous amount of decaying material that would consume oxygen as water flowed through its length. Much of the inflow water also even- tually passes through the small lake near the outlet. Lake O2 levels were consistently below saturation, especially at lower depths. The outlet stream generally was below 60 percent saturation (less than 7 mg/1) during the wet months; but from June to Septem- ber, when outflows were less than 50 1/sec, the oxygen levels were at their highest (89- 111 percent saturation, 8.4-11.2 mg/1). Higher oxygen content in the outlet during low water periods probably reflects less contact of inflowing water with fen peats. During the winter months water is ob- viously moving across much of the fen sur- face. But when inflows diminish to some minimum level (probably at least 50 1/sec combined surface and subsurface), the water tends to move only through the marginal channels ("moats," e.g. Rigg 1940) of the 10- 6 2 UJ X ^ 0 — INLET STREAM B —OUTLET STREAM pH NOV DEC JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV 1972 1973 □ATE Fig. 3. Seasonal changes in oxygen concentration and pH in the outlet compared to the major inlet stream (B). 62 Great Basin Naturalist Vol. 37, No. 1 fen. We observed that the central sedge area was noticeably drier during the period from June to September, and this observa- tion supports the idea of the sedge area's reduced contact with flowing water and of lower water levels. As long as this oxygen- rich (and relatively mineral-rich) water flows through the central fen, the rate of peat accumulation will be slower than if water flow is confined to the margins. As peat and living plants in the center of a fen become isolated from flowing water, charac- teristic changes occur (Gorham 1957, Deevey 1958, Heinselman 1970, Moore and Bellamy 1973): the rate of peat accumula- tion accelerates with a consequent rising of the surface; and the peaty soils become pro- gressively less rich in nutrients, more acid in pH and more bog than fen. Recent work in Manitoba, Canada, showed that about 74 percent of the net primary production of a bog remained after one year, whereas 52 percent remained in a marginal fen (Reader and Stewart 1972). Changes in the biota ac- company such changes in physical-chemical conditions. There is, then, a very important influence of the quantity of water and its oxygen and mineral conditions on the rate of succession of Inglenook Fen. Acidity (pH) Median pH of the inlet and the outlet streams is given in Table 1. There was a general trend of streams with lower dis- charge to have lower pH. From May through August, when inflows were derived almost entirely from aquafer rather than run-off, pH was generally above neutral (7.0-7.7) or, in the case of Aj^^, and E, was higher than at other periods, although it was less than 7.0. In contrast, streams had lower pH in months when flows were high. Tests of oxygen, temperature, and pH in the fen were made generally along Transect A. Average values from these collections are given in Table 3. Samples were taken near the fen edge, where moving water occurred all year, as well as toward the center, where water was low in summer and had little flow. These average values showed seasonal changes in oxygen, temperature. Table 3. Average selected chemical conditions Inglenook Fen. Date Number of Oxygen Temp. pH Samples (mg/1) (°C) Dec. 2, 1972 7 4.9 9.2 6.0 Jan. 1, 1973 4 5.8 7.1 — Feb. 10 4 5.8 11.2 6.2 Feb. 24 4 8.7 10.0 6.2 Mar. 24 6 6.2 13.9 6.0 Apr. 7 6 3.5 16.8 6.0 Apr. 21 5 3.5 16.8 5.9 May 10 3 2.6 13.3 6.0 May 19 3 3.0 17.2 5.8 June 19 8 2.5 14.6 5.8 July 2 3 3.3 16.0 5.7 July 14 6 0.7 18.3 5.8 Aug. 1 6 .3.0 18.0 5.9 Aug. 13 2 0.6 18.7 5.6 Sept. 4 5 3.0 18.8 5.6 Sept. 21 3 3.4 20.0 6.6 Nov. 3 5 5.0 9.2 5.8 and pH that were the opposite of the in- coming streams [see Stream B in Fig. 3). In winter months high water flushes the entire fen and elevates oxygen and pH (especially in the central region). During drier periods, oxygen and pH decline in the fen except near the marginal channels. Oxygen, pH, and aerobic limit (to be discussed later) are increased relatively close to a region of moving water (Lahde 1969, Heinselman 1970, Erman 1973). Recent studies on a raised bog-system in Germany (Lotschert and Gies 1973, Gies and Lotschert 1973) showed that the marginal fen areas were consistently higher in pH and cation con- tent than the raised, central portion of the peatland. The pH of the outflow stream also in- dicates isolation of water from the central fen region during dry periods. Instead of pH decreasing— a condition that would oc- cur if the water were in contact with the more acid central area— the pH of the out- flow increases (Fig. 3). The studies by Lot- schert and Gies (1973) also showed a sum- mer increase in pH and cations in the marginal fen of Schwarze Moor. This change in pH in Inglenook Fen could result from additions of water that drain the alka- line dunes and that are less diluted by stream inflows at this time. March 1977 Erman, et al.: California Coastal Fen 63 Gorham (1975) suggests a pH of 4.2 as a lower limit of fen conditions, and Lotschert and Gies (1973) give ranges of 4.9 to 6.3 for typical fen conditions. Previous work (Baker 1972) and our present samples from the sedge-fen region indicate pH levels of 5.4 to 6.6, depending on season (more acidic dur- ing dry periods). Calcium-Magnesium ions Calcium ions averaged 0.15 meq/liter (3 mg/1) at Stream E to 0.32 meq/1 (6.4 mg/1) at Stream D (Table 1). These values are lower than observed on one occasion by Baker (1972), who found 0.40 meq/1 in Stream E. The outlet was consistently high- er in Ca ions (average 0.62 meq/1) than the inlets, and probably reflects the influence of dunes that were found to have higher Ca ion concentration than inflow water (Baker 1972). As a further check on the possibility of the dunes influencing the outlet water, a test for silica was made 7 October 1973. If dune water is important, then silica should be much higher in the outlet than in the in- let streams. The average for the six inlets was 3.5 mg/1, compared to 8.5 mg/1 for the outlet. These results further indicate the rel- ative importance of direct drainage from the dunes into the fen margins. Magnesium ions varied similarly to cal- cium in all inlet streams (Table 1), and ranged from 0.25 meq/1 (E) to 0.47 meq/1 (C and D) or 3.5 to 5.7 mg/1. The outlet, however, contained about the same as the inlets (0.47 meq/1), which is in contrast to the situation for calcium as noted previously by Baker (1972). Sea water spraying directly onto the fen may also contribute to the concentration of various ions in the outflow, as was suggested by Baker (1972). Heinselman (1970:245) stated that "ac- cumulating evidence indicates that the ionic balance and cation content of peatland wa- ters in relation to water sources and hydro- topography are key factors influencing flo- ristics, vegetation types, and ultimately peatland evolution." He used pH, Ca, and Mg and specific conductivity of peatland waters to relate peatland types and vegeta- tion. We used overall averages of the six in- let streams for these factors and obtained these figures: pH = 6.5, Ca = .205 meq/1 (4.1 mg/1), Mg = .342 meq/1 (4.16 mg/1), and specific conductivity (measured in No- vember only) = 166. These values indicate minerotrophic to weakly minerotrophic wa- ters in Heinselman's classification (1970), and they are typical also for European fen conditions (Lotschert and Gies 1973). Sediments Mean sediment concentrations in the six inlet streams and the outlet are given in Table 1. Stream A^.^ had the greatest aver- age sediment load (19.9 mg/liter), and Stream E had the lowest (42. mg/liter). The average from all inlets was considerably higher than the outlet (13.5 mg/1 inlets, 4.4 mg/1 outlet). On an annual basis this sedi- ment influx is approximately 19.3 metric tons entering the fen and 25 metric tons leaving. The greater amount of sediments leaving than entering results partly from the influx of dune material near the outlet, where dunes encroach directly on the stream. Natural dune shifting, in addition to disturbance from off-road vehicles on the dunes, causes sands to be mobile in the vi- cinity of the fen. The small existing housing development consisting of 13 houses in Wa- tershed C and part of Watershed D and oc- casional plowing of some pastures as yet show little impact on stream-suspended sediments. One of the major factors that influences the rate of fen-bog succession is the rate of basin filling. This filling occurs from peat accumulation, dune shifting, and sedimenta- tion from the watershed. Soil disturbance in the watershed can rapidly increase the amount of sediments reaching the inlet streams and, subsequently, the fen. Most of the incoming sediments no doubt accumulate in the fen, but our methods cannot distinguish the net change. The rate of sediment loss from the entire watershed (459.4 ha) amounts to 42 kg/ha per year. In a study of the Hubbard Brook Watershed in New Hampshire, Bormann et al. (1969) found that particulate losses from a forested 64 Great Basin Naturalist Vol. 37, No. 1 watershed were 25 kg/ha per year, and a summary of studies on small, imdisturbed forested watersheds gave a range from 25 to 53 kg/ha (Likens and Bormann 1974). Phosphate and Nitrate Soluble phosphate-phosphorus (Table 1) was very similar in most of the inlet streams (.07-.09 mg/1) and was slightly higher in the outlet streams (.13 mg/1). This result suggests that PO^-P is being released by decomposition in the fen. Occasionally, every stream contained three to five times the average PO^-P found in other streams, but these higher levels occurred in no con- sistent pattern. They may have resulted from agricultural activities, since pasture- land is occasionally fertilized, but the inter- mittent nature of the higher readings does not suggest sewage contamination. Nitrates seem to be retained in the fen (Table 1). The average amount of NO3-N in the outlet (0.12 mg/1) was considerably lower than in most of the inlets. Nitrates were also occasionally high in each stream, although A^„ was consistently higher in NO3, and this result suggests some basic difference in the immediate vicinity of A^^, because A^^ did not appear similarly in- fluenced. A yellow color in A^.^ water may have interfered with colorometric determi- nations. The few homes on Watershed C and Wa- tershed D apparently had no major effect on PO4 or NO3 in those streams. The levels of these nutrients in the six streams may serve as a baseline in case of later devel- opment. Peat Depth and Aerobic Limit of the Fen Two series of cores to determine peat depth were taken along the transects shown in Fig. 1. Transect A was taken across the fen at 30 m intervals (Fig. 4). Maximum peat depth was 8.2 m along this transect. The peat was composed of three distinct layers: a narrow surface layer of dense liv- ing plant roots and organic matter; a layer DEPTH CM) north edge CHI dense peat and rcxjts S black sandy peat ^ dense peot [S] gray- sandy [2D clay-sand AEROBIC LIMIT JUNE 2,1973 DISTANCE (MJ Fig. 4. Peat depth profile (A) along transect a at 30 m intervals and aerobic limits (B) along the same transect at 15 m intervals. Dotted lines near surface of A indicate approximate location of stream channels. March 1977 Erman, et al.: California Coastal Fen 65 underlying the surface made up of loose, soft brown peat that was very liquid; and a basal layer of dense peat in the central fen region. When the peat borer cut through the surface layer of dense fibrous peat, wa- ter squirted up through the hole, especially on the southern edge. The corer then passed freely through the next layer without effort until the bottom peat layer was reached. It appeared that the entire surface was float- ing over the liquid middle layer, and exerted a pressure on the watery fluid be- low. Beneath the peat was a dark clay layer, except on the southeastern edge (0-30 m), where black organic ooze overlaid the dark clay layer directly below the surface channel. Along this same transect, a series of measures of aerobic limit were taken (2 June 1973) every 15 m. The influence of moving water is apparent in the aerobic limit measures; the limits tend to be deeper where moving water maintains higher O2 levels, and hence maintains deep O2 pene- tration (Fig. 4). Mean aerobic limit for the 11 samples was 7.1 cm. A second transect to determine peat depth was taken along the east-west axis of the fen (Fig. 5) and ended at Sandhill Lake. The layers of peat are similar to those in Fig. 4. Fen depth increased toward the lake, where depths exceeded 11 m, the limit of our coring device. This transect halfway along its length also revealed a ridge covered with dark sand on the bot- tom. The higher sand-topped ridge may be the remnant of a former dune now sub- merged by the fen. Depth measures of Sandhill Lake showed that it had vertical sides that drop off rap- idly to about 6 m. The peat beneath the lake was of the same loose, watery texture as the similar layer in the fen and extended beyond the 11 m limit of our device. Peat profiles of bogs along the Pacific coast made by Rigg and Richardson (1938) showed some bogs with lakes that had similar steep sides and were relatively deep in com- parison to the total bog depth. Heinselman (1970) showed that even small lakes sur- rounded by peat may persist for long peri- ods. Early workers on bog lakes believed that the lakes were in terminal stages of succession and would soon fill. While this conclusion may still be valid over a very long time period, Heinselman (1970) showed that Myrtle Lake in Minnesota had main- tained its size and depth and had even ris- en, rather than filled, with the height in- crease of the surrounding peat. Under existing conditions Inglenook Fen DEPTH (M) Jlllilll^^ LIQUID PEAT ^%^^ CD DENSE PEAT AND ROOTS B DENSE PEAT EZ] CLAY ^^DARK SAND -^? 30 60 90 DISTANCE (M) Fig. 5. Peat depth profile along transect b. Note sandy ridge at 180 m that may be a remnant sand dune. Cor- ing device could not extend beyond 11m near the lake. 66 Great Basin Naturalist Vol. 37, No. 1 is slowly filling, and during low flows it shows signs of succession toward bog (om- brotrophic peatland). The process of change from minerotrophic to ombrotrophic would still require a very long time. But as recent studies of lake eutrophication have shown, a natural successional change in productive status can be tremendously increased by man's activities. The data in this report pro- vide baseline information on some of the key elements essential to maintenance of a minerotrophic peatland. Acknowledgments We thank Ken McDonald and Dick Sand- er for field assistance and Clarence Johnson for analysis of Ca and Mg. The Nature Conservancy and the state of California contributed financial support for the work. We thank Evert Schlinger and Herbert Bak- er for criticizing an earlier draft of the manuscript and Mr. Ralston Ross for his per- mission to visit the fen on his land. Literature Cited Baker, H. G. 1972. A fen on the northern California coast. Madrono 21:405-416. BoRMANN, F. H., G. E. Likens, and J. S. Eaton. 1969. Biotic regulation of particulate and solution losses from a forested ecosystem. BioScience 19(7):600-610. Deevey, E. S., Jr. 1958. Bogs. Scient. Amer. Reprint #840 (October 1958). 8 pp. Erman, D. C. 1973. Invertebrate movements and some diel and seasonal changes in a Sierra Nevada peatland. Oikos 24:85-93. GiES, T., and W. Lotschert. 1973. Untersuchungen uben den Kationengehalt im Hochmoor. II. Jah- reszeitliche Veranderungen und Einfluss der Sphagnen-Vegetation. Flora 162:244-268. GoRHAM, E. 1957. The development of peatlands. Quart. Rev. Biol. 32:145-166. Heinselman, M. L. 1970. Landscape evolution, peat- land types, and the environment in the Lake Agassiz Peatlands Natural Area, Minnesota. Ecol. Monogr. 40:235-261. Jenny, H., R. J. Arkley, and A. M. Schultz. 1969. The pygmy forest podsol ecosystem and its dune associates of the Mendocino Coast. Madr- ono 20:60-70. Lahde, E. 1969. Biological activity in some natural and drained peat soils with special reference to oxidation reduction conditions. Acta. For. Fenn. 94:1-68. Likens, G. E., and F. H. Bormann. 1974. Linkages be- tween terrestrial and aquatic ecosystems. Bio Science 24:447-456. Lotschert, W., and T. Gies. 1973. Untersuchungen uber den Kationengehalt im Hochmoor. I. Ab- stufengen in den Vegetationskomplexen. Flora 162:215-243. Moore, P. D., and D. J. Bellamy. 1973. Peatlands. Springer- Verlag, New York. 221 pp. Reader, R. J., and J. M. Stewart. 1972. The relation- ships between net primary production and ac- cumulation for a peatland in southeastern Mani- toba. Ecology 53(6): 1024-1037. RiCG, G. B. 1933. Notes on a sphagnum bog at Fort Bragg, California. Science 77:535-536. 1940. Comparisons of the development of some sphagnum bogs of the Atlantic coast, the interior, and the Pacific coast. Amer. J. Bot. 27:1-14. RicG, G. B., AND C. T. Richardson. 1938. Profiles of some sphagnum bogs of the Pacific coast of North America. Ecology 19:408-434. INTRODUCED AND EXPORTED AMERICAN SCOLYTIDAE (COLEOPTERA)' Stephen L. Wood- Abstract.— Fourteen North, Central, and South American Scolytidae have reached areas outside of America, namely Hawaii (5), Australia (3), Southeast Asia (2), Africa (7), Europe (1), and worldwide (1). The 44 species in- troduced into North and Central America have originated in South America (10), Southeast Asia (15), Africa (14), or Europe (7). Four North and Central American species have extended their ranges into new territory within this region. Most of the species that breed in the bole or roots of their host were imported more than a half cen- tury ago; those imported since 1950 breed mostly in seeds, pods, dry fruits, or small branches. Forty-five of the 62 species apparently can reproduce by means of facultative arrhenotokic parthenogenesis. A brief report on the extraterritorial dis- tribution of American Scolytidae to the Fourteenth International Congress of En- tomology (Wood 1972) aroused interest suf- ficient to prompt a more comprehensive re- view. This report omits those species that have moved into or out of North and Cen- tral America through natural migration. Those species intercepted in routine border inspection and not actually established in an extraterritorial locality are also omitted. This report includes only those species for which I have examined specimens taken from a breeding population in an extra- territorial locality. In most instances the in- troductions to new areas have occurred within the past century, and the spread of each species is part of the historical record. In other cases the entry and spread was not recorded, and the importation is based on undocumented circumstantial evidence. An account of the nomenclature, distribution, hosts, classification, etc., will be published in my monograph of North and Central American Scolvtidae (currently in prepara- tion). The documentation for the many items cited below will appear in the mon- ograph. Of the 62 species listed here (Table 1), six are native North or Central American spe- cies that were introduced to another extra- territorial part of North or Central Amer- ica. The 14 species exported from North, Central, and South America have reached the following areas (some have reached more than one area): Hawaii 5 Australia 3 S.E. Asia 2 Africa 7 Europe 1 Worldwide 1 Of the 44 species imported into North and Central America, probable areas of origin are (three Asian species also occur in Af- rica): South America 10 Africa 14 S.E. Asia 15 Europe 7 The 62 species treated here are sorted on the basis of the host material usually se- lected for the excavation of breeding tun- nels as follows: Roots 2 Bole or logs 11 Twigs and branches 28 Both seeds and branches 7 Seeds, pods, dry fruits 10 Misc. branches, weeds, herbs, etc. 2 Fruiting stalks of grass 1 Unknown 1 Almost all of the imported species that breed in roots, logs, or boles were estab- lished in North and Central America a half century or more ago. Those species im- ported since about 1940 breed almost ex- clusively in seeds, pods, or other dry fruit- Tart of the research on which this studv was based was sponsored l.y the N -Department of Zoology, Bri^ham VoiniK University, Provo, Utafi S46()2. Sec 67 68 Great Basin Naturalist Vol. 37, No. 1 Table 1. North and Central American Scolvtidae that have been introdnced from or exported to foreign lands. Species Aniptus politus Coccotrypes acictilatus aclverui carpophagus dactijUperihi (li.stincttis indicus rliizoplumie Corthijlus spinifer Cryphalomorphus jalapae Cryptocareniis heveae seriatus Cryphirgm pusillus Dendroctonus frontalis Gnatbotrich us materiarius Hyldstinus obscttrus Hypocrypludtts mimgiferae llypothcncmus aequahclavatus africanus (ireccue binnanus bntnneus californicus coltimbi CTudiae cylindricus erectus eniditus hampei javanus obscurtis parallehis pubescens seriatus setosus Ips grandicoUis Microborus boops Pugiocerus frontalis Phloeosinus cupressi Pityophthorus confusus juglanais Premnobius cavipennis ambitiosus Scolytus malt multistriatus rugulosus Xyleborinus aspericauda saxeseni Xyleborus ajfxnis aispar ferrugineus obliquus rubricollis spinulosus lalidus volvulus xylographus Xylosandrus romparlus rrassiusculus geniianiis tnorifurus zimmerrnanni Probable origin Introduced to Mexico, C. .\merica New Gninea? S.E. Asia, Indonesia Africa? Africa? Cevlon? S.E. Asia Indonesia C. and S. America Mexico C. and S. America S. .America Europe, .\sia USA E. North America Europe India Indonesia S. Africa S.E. Asia S.E. Asia Africa? Africa? Africa? S.E. Asia? Africa? Africa? Tropical .\merica? .\frica Africa? Tropical .\merica Mexico Tropical America? Tropical America? .\frica? N. .\m erica C. America S. America N. .America USA Arizona .\frica .\frica Europe Europe Europe C. and S. America Europe Tropical .America Europe Tropical .\merica S. .\merica? S.E. Asia Tropical .\merica S.E. .\sia Tropical America E. North America S.E. Asia? S.E. Asia Japan S.E. .\sia Tropical .America Florida, Antilles Panama and Brazil Cuba to S. .\merica Florida to S. .America USA to S. America Florida to S. .America Florida to S. America Florida, Calapagos Islands Florida Worldwide interceptions Africa, Florida Florida, .Antilles E. North America Honduras France N. America Florida to S. America, etc. Honduras, Jamaica USA to S. America Florida to S. America, etc. Florida, C. America, etc. USA to Trinidad USA to Mexico USA to S. America USA to S. America Mexico to S. America USA to S. .America Almost cosmopolitan C. and S. .America, etc. Florida to S. America, etc. Intercepted worldwide Hawaii Florida, Hawaii, etc. USA, Africa, etc. Antilles to S. .America .Australia .Africa, Jamaica USA, etc. Panama, .Australia, etc. Honduras California Florida to S. America S. America N. America N. America, etc. N. America, etc. Florida N. America, etc. Hawaii to Malaya, Africa N. America Hawaii to Australia, .Africa N. .America, Africa N. .America Hawaii New York Hawaii to Malaya, Africa California Florida, Cuba, Africa, etc. South Carolina, Hawaii USA C. and S. America Florida Host material Seeds Seeds or branches Seeds Seeds Palm fruits Seeds Mangrove "seeds" Branches or seedlings Jalapa root Branches, etc. Branches, etc. Picea boles Pinus boles Pinus boles Clover roots Mangifera indica Unknown Fruits, branches Seeds, branches, etc. Branches Branches Twigs, etc. Twigs, weeds, etc. Twigs, weeds, etc. Branches Branches Twigs, seeds, etc. Coffee berries Branches, pods Brazil nuts, etc. Twigs? Grass Twigs, seeds, etc. Fruits, pods, etc. Pinus limbs, etc. Branches, etc. Seeds Cupressus branches Pinus bole Juglans branches Limbs, etc. Limbs, etc. Fruit trees Ulmus limbs, etc. Fruit trees Limbs, logs Limbs, logs, etc. Logs, etc. Limbs, etc. Logs Branches Branches, boles, etc. Limbs Limbs, etc. Logs Qucrcus logs Twigs Branches Limbs, etc. Branches, etc. Branches March 1977 Wood: American Scolytidae ing structures or in small branches. It is of interest that 45 of the 62 species can repro- duce bv means of facultative arrhenotokic parthenogenesis, by which means one fe- male can establish an entire breeding popu- lation. Araptus politus Blandford, 1904, is native to southern Mexico and Central America, where it breeds in Muciina seeds. It has been taken in Haiti (Cola nuts), Jamaica and Cuba (host not recorded), and Miami, Florida (intercepted in "bird seed"; breed- ing in "seeds") since 1960. Coccotnjpes aciculatus Schedl, 1952, was named from New Guinea. It has been taken in Brazil (Cuma nuts), Costa Rica, and Pan- ama (at light) since 1938. Its hosts and habits are unknown. Coccotnjpes advena Blandford, 1894, is apparently a native of Indonesia. It has been taken in India, Java, Japan, Micro- nesia, Samoa, Hawaii, Surinam, and Cuba since 1915. It breeds in either bark or large seeds of a variety of tropical hosts. Coccotnjpes carpophagus (Hornung, 1842) was named from betel nuts from India, al- though it probably originated in Africa. It has spread to virtually all tropical and sub- tropical areas of the world, where it breeds in a wide variety of nuts and other large seeds. It is commonly intercepted through- out the world, but in North America it breeds permanently only in southern Florida and Mexico. Coccotnjpes dactijliperda (Fabricius, 1801) is probably of African origin. It breeds in virtually all tropical and subtropical areas of the world. It is most common in date pits, but may breed in a wide variety of nuts, seeds, and buttons or other objects made from vegetable ivory. It breeds in Florida and Arizona and probably in other southern states. Coccotnjpes distinctus (Motschulsky, 1866) was named from Ceylon, where it probably originated. It has also been reported from Micronesia, Hawaii, Honduras, Puerto Rico, British Guiana, and southern Florida, where it breeds in a wide variety of palm seeds. Coccotnjpes indicus (Eggers, 1936) was named from India. It is widely distributed in southeastern Asia and Indonesia, and has been reported from the Fiji Islands, Tahiti, Hawaii, Brazil, Surinam, Trinidad, Puerto Rico, Martinique, Guadeloupe, Costa Rica, Panama, and Coral Gables, Florida. It has been intercepted in various seeds at several American ports. It probably is capable of breeding in large seeds in most southern states of the U.S.A. Coccotnjpes rhizophorae (Hopkins, 1915) is apparently from Indonesia. It also breeds in the Galapagos Islands and southern Flor- ida. It infests the viviparous seeds of man- grove (Rhizophora mangle). Corthijlus spinifer Schwarz, 1891, was named from Florida, but it is very doubtful that it is a native species. In addition to Florida, its present distribution extends from Brazil to Veracruz, Mexico. Several related species occur in Central America and north- ern South America. In all probability, it has spread through commerce to Brazil and Florida in the past century. Cnjphalomorphus jtdapae (Letzner, 1848) is native to Mexico, where it breeds in ja- lapa root (presumably Exogonium jalapa). It is not known to be permanently established in any invaded territory. Cnjptocarenus heveae (Hagedorn, 1912) was named from Africa (Congo), where it damaged rubber trees. It is now known to be native to Mexico, Central America, and northern South America. It also breeds in Ghana, Congo, Brazil, Jamaica, Cuba, and southern Florida. It infests the pith of small, broken, or unthrifty stems of a wide variety of trees, shrubs, and woody vines. Cryptocarenus seriatus Eggers, 1933, is probably native to Bolivia and Peru, but now occurs throughout South America, Central America, Mexico, Jamaica, Haiti, the Virgin Islands, Cuba, Florida, and south- ern Texas. It appears to have entered the United States since 1930 and to have reached Central America and Mexico after 1915 but prior to 1953. It breeds in the small branches of a wide variety of trees, shrubs, and woody vines. Cnjpturgus pusUhts (Gyllenhal, 1813) is native to Europe, north Africa, and north- ern Asia. It was apparently introduced to eastern Canada and/or the northeastern United States in the colonial period. It is a 70 Great Basin Naturalist Vol. 37, No. 1 secondaiN' enemy of spruce (Picea), and breeds from Ontario and Newfoundland to New Jersey in the bark of the bole and larger branches of the host. Dejidroctonus frontalis Zimmerniann, 1868, is native to the southern United States, where it is a primary enemy of pine {Pinus). It was evidently introduced into Honduras between 1910 and 1964. It occurs in the same trees with D. mexicanus, with which it is easily confused. A simultaneous introduction of Pityophthorus confusus Blandford apparently occurred. Recent col- lections (not seen) report this species from southeastern Mexico. If correct, these re- cords indicate that this species reached Honduras by natural migration and not through commerce. Gnatlwtrichiis matcriarius (Fitch, 1858) is native to eastern North America, where it is a secondary enemy of pine (Pinus). Shortly before 1936 it was introduced into France. Since then it has spread into neighboring areas, and may eventually occupy the entire pine belt of Europe and Asia. It is a wood- iioring ambrosia beetle. Hylastiniis obsciiriis (Marsham, 1802) was introduced to North America (New York) from Europe sometime prior to 1878. By 1929 it had reached British Columbia and Oregon. It now occurs in all provinces and states where alfalfa or other forage clovers are grown. Ihjpocrijphalus mangiferae (Stebbing, 1914) is apparently native to India, but it has spread throughout the tropical regions of the world wherever its host, Mangifera indira, is cultivated. It was recorded from Guadeloupe Island in 1872 and was taken in Florida in about 1949. It is common throughout southern Mexico and Central .■\merica. Hypotlwnenuis acipialiclavatus Schedl, 19.39, was named from Indonesia. A speci- men was taken in Honduras in 1906 and an- other in Jamaica in 1967. Nothing more is known of this species. Hypoilwncmus africanus (Hopkins, 1915) was named from South Africa, where it probably originated. Since 1933 it has been reported from Indonesia, .Malaysia, Brazil, Venezuela,^ Jamaica, Bahama Islands, Do- minican Republic, Puerto Rico, Honduras, Alabama, and Louisiana. It breeds in the twigs and fruiting pods of various trees, shrubs, and vines. Hypothenemus areccae (Hornung, 1842) is apparently native to southeastern Asia. It now occurs in virtually all tropical and sub- tropical countries in the world. The first re- cord of its occurrence in America was in Brazil in 1928. It now occurs throughout South America, Hawaii, Martinique, the Virgin Islands, Puerto Rico, the Bahama Is- lands, and southern Florida. It breeds in seeds, dry fruits, twigs, and in a wide varie- ty of other materials. At times it becomes a serious pest of mature seeds and nuts and stored products. Hypothenemus hinnanus (Eichhoff, 1878) was named from Burma, where it probably is native. It has been reported from south- east Asia, Indonesia, the Philippine Islands, Australia, Micronesia, Hawaii, Jamaica, Central America, Mexico, and Florida. It breeds in the twigs and small branches of a wide variety of trees, shrubs, and woody vines. It was first taken in Jamaica in 1937 and in Florida in 1951. Hypothenemus brunneus (Hopkins, 1915) occurs from Alabama, Florida, and Texas to Panama, including the Bahama Islands, Cuba, Puerto Rico, the Virgin Islands, and Trinidad. It has no native American rela- tives, but several similar species are native to Africa. For this reason it is presumed to be of African origin. It was first reported from Texas in 1915. Hypothenemus californicus Hopkins, 1915, was regarded as a native North Amer- ican species until specimens were taken re- cently in Liberia and Israel. Since it is not allied to any American species, but has sev- eral near relatives in Africa, it is presumed to be of .-Xfrican origin. It occurs from southern California to New Jersey and southward to southern Mexico. Hypothenemus eohimbi Hopkins, 1915, was named from the southern United States and has since been reported from Mexico, Costa Rica, Colombia, Venezuela, Cuba, and the Bahama Islands. All representatives of this species group are of Old World ori- gin. It is presumed that this species also March 1977 Wood: American Scolytidae 71 will eventually be found in Africa. It breeds in a wide variety of materials, including twigs, weeds, straw, seeds (rare), etc. It is of some economic importance. Hi/potfienemus crudiae (Panzer, 1791) was named from India, but it has spread through commerce to virtually all tropical and subtropical areas of the world for so long that its true origin is unknown. It is common from the eastern United States to Argentina. Circumstantial evidence suggests a southern Asian origin. Although it breeds in small branches of trees, shrubs, vines, weeds, or other plants, its greatest popu- lations occur in seeds, pods, or other fruit- ing bodies, where it has caused much eco- nomic damage in mature seeds both in the field and in storage. Its importance in agri- culture or forestry is limited to its effect on seed production. Hypothenetniis cijUndricus (Hopkins, 1915) was named from Guatemala, but is now known from Mexico to Panama, Co- lombia, Venezuela, Guadeloupe, and Africa (Liberia). Since this species has no close rel- atives native to America, it is presumed that it is of African origin where several near relatives occur. It breeds in small branches of trees, shrubs, and vines. Hypothenemiis erectiis LeConte, 1876, was named from Texas, but it is now known from Texas to Honduras, Venezuela, Cuba, the Virgin Islands, and Africa. Several very closely related forms occur in Africa, but there are no known native American species that are closely related For this reason it is presumed to be of African origin. It breeds in small branches of a wide variety of trees and shrubs. Hypothenemiis eruditiis Westwood, 1836, is probably the most widely distributed and most frequently encountered scolytid species in the world. It is known from virtually all tropical and subtropical areas of the world and in temperate regions where sufficient moisture and vegetation are present. It breeds in the bark of the trunk or branches of numerous plants, in flowers, weeds, grass, fruiting bodies of fungi, and seeds; the origi- nal series was breeding in the cover of an old book. In America it is known from Michigan to Argentina as a common spe- cies. It probably originated in the American tropics. It is frequently of economic impor- tance in mature seeds or pods both in the field and in storage. Hypothenemiis hanipei (Ferrari, 1867) is the notorious coffee berry borer. It is appar- ently native to Africa, but it occurs throughout the coffee-growing regions of the Old World and in Brazil. More recently it has reached Colombia and Guatemala. It breeds only in coffee beans. Infested beans have been intercepted in most countries of the world. It is of great economic impor- tance. Hypothenemiis javoniis (Eggers, 1908) was named from Indonesia, although it may have originated in Africa. It is now known from Africa (Camerouns, Congo, Ghana, and Liberia), Indonesia, the Philippines, Mexico to Panama, Colombia, Venezuela, Guadeloupe, Puerto Rico, Cuba, and Flor- ida. It breeds in small branches and seed pods of a wide variety of plants. It has been of economic concern in some situations. Hypothenemiis obsciinis (Fabricius, 1801) originated in the American tropics. It breeds from Costa Rica and Puerto Rico to Brazil, but has been intercepted in seeds and nuts in most countries of the world. It is best known as a borer in Brazil nuts, al- though it breeds in a wide variety of seeds and dry fruits and, apparently, can repro- duce in bark (uncommon). It is an impor- tant pest of many tropical nuts and fruits. Breeding populations in Florida apparently hybridize with the native H. seriatiis (Eich- hoff) and are absorbed by that species. Hypothenemiis parallehts (Hopkins, 1915) was named from Mexico, where it may have originated. It is also known to occur in Hawaii. It is very similar to H. eruditus Westwood, and may be no more than a morphological variation of that species. Hypothenemiis piihescens Hopkins, 1915, was named from Florida, but it is now known to occur in Texas, Puerto Rico, Mex- ico, and Hawaii. Because of its very small size and its unusual habit of breeding in the fruiting stalks of grass {Andropogon, Cy no- don, and Paspahim) it probably has been overlooked. An American origin is sus- pected. 72 Great Basin Naturalist Vol. 37, No. 1 Hypothenemus seriatus (Eichhoff, 1871) was named from Louisiana. It occurs from West Virginia to Brazil, Hawaii to Austral- ia, Indonesia, Madagascar, and Africa (the Ivorv Coast). It is probably of American origin. It breeds most commonly in twigs and small branches, but may also occur in seeds, pods, weeds, or other plant material. It is of economic concern. Hifpothenemus setosus (Eichhoff, 1867) was named from Guadeloupe, although it is probably native to Africa. It occurs from Mexico to Brazil, Guadeloupe, Puerto Rico, and Africa (Camerouns, Congo). It breeds in small branches and fruiting bodies of a wide varietv of hosts. Ips g,r(indicolU.s (Eichhoff, 1868) occurs from southern Manitoba and Quebec to Florida and Honduras. It was introduced into pine plantations in Australia. It is of economic concern. Microhorus hoops Blandford, 1897, is a rare species that is known from Guatemala to Panama. Its occurrence in Jamaica prob- ably represents an introduction. It was also introduced into Africa (Camerouns, Gold Coast, Uganda) prior to 1919. Pagiocerus frontalis (Fabricius, 1801) probably originated in southern South America, where similar species occur. It has spread northward in America to North Carolina and Texas, and it is occasionally intercepted in corn (maize) in Europe and elsewhere. In the Andean region of South .\merica it is a significant pest of dry ma- ture corn in the field and in storage. Phloeosiniis cupressi Hopkins, 1903, is na- tive to the coastal region from Alaska to California, where it breeds in Cupressus and a few related tree genera. It has been in- troduced into Panama, Australia, and New Zealand. Pitijophtlwrus ronfustis Blandford, 1904, is apparently native to the southeastern United States, where it occurs in Pinus holes with Dcndroctomis frontalis Zimmer- mann. It appears to have reached Hondviras with D. frontalis either by natural migration or through commerce (see D. frontalis above). Pityophtlwrus jii}ilandis Blackman, 1928, infests native black waliuit {Julians) in southern Arizona and New Mexico. It was reported as new to California by Bright and Stark (1973). Since native potential hosts occur in the area, there is a question as to whether an oversight in collecting prior to 1959 occurred or an introduction actually did occur. Premnohius cavipennis Eichhoff, 1878, is native to Africa, where it is a common, widely distributed species. Prior to 1931 it was introduced into South America and has now spread northward to southern Mexico (1951) and Florida (1939). It is an ambrosia beetle that breeds in a wide variety of hosts in material about two to eight inches in di- ameter. It may be of some economic con- cern. Premnohius amhitiosus (Schaufuss, 1897) has not yet been reported from North or Central America, although it probably oc- curs there. It is native to Africa, but was in- troduced into Brazil and Colombia prior to 1958. It is now known from Bolivia and has probably reached most other South and Central American countries. Its habits and importance apparently are similar to those of P. cavipennis. Scohjtus mali (Bechstein, 1805) is native to Europe, where it is a pest of apples, pears, prunes, and other cultivated fruit trees. It was introduced into New York pri- or to 1868 but was virtually unknown here until about 1950. Since then it has spread into southern Ontario, Michigan, Ohio, and Maryland and should spread into most of the states in the U.S.A. within a few years. It is larger than S. nig^iilosus, but apparent- ly has similar habits. Scolytiis multistriatiis (Marsham, 1802) is native to temperate Europe and Asia. It was introduced into North America and now occurs in all of the 48 contiguous states of the U.S.A. and is apparently in all of the southern provinces of Canada. It is the well-known European elm bark beetle, the principal vector of Dutch Elm Disease. It has also been introduced into several oth- er areas of the world where elms grow. Scolytiis ru^ulosus (Miiller, 1818) is the shot-hole borer of cultivated fruit trees. It is native to Europe and Asia, but it was in- troduced into North America very earlv in March 1977 Wood: American Scolytidae 73 the settlement of this continent and now oc- curs in all states and provinces where ap- ples are grown. Xylehorinus aspericaiula (Eggers, 1941) is native to Central and South America. One specimen was taken in the field in southern Florida (Biscayne Bay). Whether or not it is established in Florida is not known. It is an ambrosia beetle that breeds in the limbs and logs of a wide variety of host plants. Xylehorinus saxeseni (Ratzeburg, 1837) is native to Europe, but has been transported to most temperate areas of the world. In North America it has been reported from 33 states of the U.S.A., from British Colum- bia, and from Baja California. It is probably more widely distributed, but has been over- looked due to the lack of collecting. It was also intercepted in Australia in logs from Brazil and it has been in Hawaii for many years. In desert areas it is of little or no economic concern, but may be a significant pest elsewhere. Xyleborus affinis Eichhoff, 1868, is prob- ably native to tropical America, but now occurs from Massachusetts to Argentina, from Hawaii to Malaya, and in Africa. It breeds in injured, dying, and cut trees and is a significant pest of logs. It does not nor- mally breed in sawed lumber. Xyleborus dispar (Fabricius, 1792) is na- tive to Europe, but was introduced into North America prior to 1817. It is now widely distributed in southern Canada (Nova Scotia to British Columbia) and in the northern United States (south tt) Vir- ginia and Utah). It is an ambrosia beetle that breeds in a wide variety of hosts in material two to ten inches in diameter. It is of economic concern in some areas. Xyleborus ferrugineus (Fabricius, 1801) probably originated in tropical America, but it has now spread to Arizona, Michigan, and Massachusetts in the continental United States, as well as Hawaii, Micronesia, Aus- tralia, and Africa, and may have reached other tropical areas where it has not yet been detected. Its habits are similar to those of A', affinis except that it is more aggres- sive and more abundant and, consequently, is of greater economic concern. Xyleborus obliquus (LeConte, 1878) was named from Florida, but it is probably of South American origin where it and similar species occur. It now occurs from Virginia to Brazil and in Africa (Congo and Zambia). It apparently breeds in unthrifty or cut branches of a wide variety of hosts. In the United States it has been reported from the District of Columbia, Florida, Georgia, Mis- sissippi, and Virginia; in Mexico from Mich- oacan; in Central America from Guatemala; and in South America from Colombia and Brazil. Xyleborus rubricollis Eichhoff, 1875, is na- tive to southeastern Asia from Malaya to Ja- pan. It was introduced into the United States prior to 1942, where it has been re- ported from Connecticut, Maryland, and Virginia. It is an ambrosia beetle that breeds in a wide variety of hosts. It may be of local economic concern in some areas. Xyleborus spinulosus Blandford, 1898, is native to tropical America from central Mexico to Brazil. It was introduced into Hawaii prior to 1934. It breeds in the branches of a wide variety of host trees, shrubs, and woody vines. Xyleborus validus Eichhoff, 1875, was named from Japan; it has also been report- ed from Taiwan and China. About 1975 it became established in New York. In recent years it has been intercepted from wooden packing crates of Japanese origin at numer- ous ports worldwide. It is anticipated that it will become established in additional areas in the near future. Xyleborus volvulus (Fabricius, 1775) oc- curs in two geographical races that prob- ably ranged from the coastal southeastern United States to Argentina. One or both races, including hybrids between the two, have spread to Hawaii, Micronesia, Austral- ia to Malaya, Africa, and Madagascar. Its habits are similar to X. affinis and X. ferru- gineus except that it is usually less abun- dant. It is of some conomic concern. Xyleborus xylographus (Say, 1926) is na- tive to eastern North America from Ontario and Quebec to the Gulf Coast and Cuba. A few very old specimens labeled "Brazil" and "Chile" have been examined, but they are assumed to be mislabeled. One speci- men from China Flat, Eldorado County, 74 Great Basin Naturalist Vol. 37, No. 1 C:alifornia (1948), apparently represents a new introduction. It breeds in Qiicrcus, rarelv in other trees. Xi/loscindrus compactus (Eichiioff, 1874) occurs from Hawaii to Sri Lanka (Ceylon) and Japan, Africa, Cuba (1958), Mississippi (1968). Georgia (1975?), and Florida (1941). It is probably of Asian origin. It is an ag- gressive primary borer in new, vigorous twig terminals in a wide variety of hosts. It has significant economic importance. When 1 reported the identity of this species per- sonally in 1952 to United States Depart- ment of Agriculture officials, I was told: "Suppress the information. We already have too many problems to worry about." At the time, its distribution was very local in Flor- ida and eradication would have been com- paratively simple. It is widely known as the black shot-hole borer. It may be of extreme economic importance in some areas. Xi/Umindrus crassiusculus (Motschulsky, 1866) is native to the Indo-Malayan region. It now occurs in Africa, Hawaii, Australia, and South Carolina (1974). It is an ambrosia beetle that breeds in branches and limbs of a wide variety of hosts. It has some eco- nomic importance. Xi/losandrus gennanus (Blandford, 1894) was named from Japan. It was introduced into the United States prior to 19.31 and it now occurs from Illinois and Connecticut to West Virginia. It breeds in a wide variety of hosts from the branches to the stump. It may be of local economic concern. Xijlosandrus morigerus (Blandford, 1894) is native to southeastern Asia and Indonesia. It was introduced into America prior to 1959 and now breeds from Mexico (Vera- cruz) to Colombia and Venezuela. It has been intercepted in ports of entry in Ari- zona, California, and New Jersey, but breeding populations are not yet known from the United States. It is an ambrosia beetle that breeds in small stems of a wide variety of trees, shrubs, vines, and her- baceous plants. It has some economic im- portance. It is widely known as the red shot-hole borer. Xijlosandrus zimmermonni (Hopkins, 1915) is probably native to northern South America and Central America. It apparently was introduced into southern Florida prior to 1915. It is rather uncommon in small branches of trees, shrubs, and vines of a wide variety of plants. It is of minor eco- nomic concern. ReferExNces Cited Bright, D. E., Jr., and R. W. Stark. 1973. The bark and ambrosia beetles of California, Coleoptera: Scolvtidae and Platvpodidae. Bull. California In- sect'Survey 16:1-169. Wood, S. L. Geologically recent and modern in- troductions of Scolytidae (Coleoptera) to North America. 14th International Congress of En- tomology Abstracts 1:112. THE SAGEBRUSH VOLE FLEA, MEGABOTHRIS CLANTONI PRINCEI, IN WESTERN UTAH, WITH COMMENTS ON THE DISTRIBUTION OF MEGABOTHRIS IN THE BONNEVILLE BASIN Harold J. Egoscue' Abstract.— The sagebrush vole flea {Megfibothris clantoni princei Hubbard) is reported from Utah for the first time, based on collections made in Tooele County. Distribution of the flea Megabothris ahantis in the Bonneville Basin is limited to the Oquirrh Mountains, where its presence appears to be determined by the occurrence of the western jumping mouse. Fleas previously recorded from the sage- brush vole, Lagunis curtatus intermedins (Taylor), in western Utah included Amphip- sijUa sihirica ssp. "near" washingtona (Ego- scue 1966), a primary sagebrush vole flea described from eastern Washington (Hub- bard 1954). This note reports the occur- rence of a second sagebrush vole flea in Utah, Megabothris elantoni princei Hubbard, with a range extension of about 604 km east-southeast from the type locality, 6.4 km west of Vya, Washoe County, Nevada. A male M. c. princei (H.J.E. No. 7) was among seven fleas collected 15 May 1963 from a spotted skunk (Spilogale ptitorius gracilis Merriam) that was trapped on John- son Pass between the Stansbury and Onaqui Mountains in Tooele County, Utah (elev. 1830 m). This association was no doubt ac- cidental. A second male (H.J.E. No. 6349) was the only flea from a sagebrush vole trapped 13 February 1969 about .4 km west of the previous locality. Both traplines were set among rock out- crops and ledges on dry hillsides where the dominant plants were sagebrush {Artemesia sp.) and juniper trees {Jiiniperus osteospertna (Torr.) Little). Other small mammals com- monly collected there were the deer mouse {Peromyscus maniculatiis sonoriensis (Le Conte)), canyon mouse {P. crinitus per- gracilis Goldman), long-tailed vole (Microtus longicaudus latus Hall), desert wood rat (Neotoma lepida lepida Thomas), bushy- tailed wood rat {N. cinerea acraia (Elliot)), cliff chipmunk {Etitamias dorsalis utahensis Merriam), and Great Basin pocket mouse {Perognathus parvus olivaceus Merriam). Despite several efforts to capture them in both livetraps and snaptraps, sagebrush voles were rarely trapped here or at numer- ous other places in the Bonneville Basin where conditions seemed more favorable. The two specimens of M. c. princei were originally misidentified by me as atypical Megabothris abantis Rothschild. At the time, this was the only species of Mega- bothris known from Utah (Stark 1958), and I was unfamiliar with M. clantoni. The dis- covery of my error made me realize that the range of M. abantis within the Bonne- ville Basin was limited to the Oquirrh Mountains. In the Oquirrh Mountains M. abantis was the most common flea on west- em jumping mice {Zapus princeps utahensis Hall), but was also found regularly in rippa- rian situations and on adjacent damp hill- sides above 1525 m on long-tailed voles, and less commonly in the same location on deer mice associated with Zapus. In years of collecting in suitable habitats, scientists have failed to trap jumping mice in other mountains within the Basin; M. abantis was not found among hundreds of fleas collected from long-tailed and montane voles, deer mice, and other small mammals in the West Tintics, Stansburys, Onaquis, Sheeprocks, and other western Utah mountains. Stark believed the preferred hosts of M. abantis in Utah were microtines. If this is so, I have National Zoological Park, Smithsonian Institution, Washington, DC. 20009. 75 Great Basin Naturalist Vol. 37, No. 1 no reasonable explanation for the restricted range of M. abantis in an area where Micro- tus are common and widely distributed. My evidence strongly suggests that the presence of this flea in the Bonneville Basin is de- pendent on the occurrence of jumping mice. Past climatic and ecological changes of the Pleistocene Epoch must explain the disjunct distribution of Z. princeps and oth- er essentially montane mammals in this part of Utah. Conditions in the Pleistocene Epoch apparently enabled certain small mammals to penetrate further west from the centrally located Wasatch cordillera than did other small mammals. The habitat that permitted this differential expansion no longer exists, and the distribution of some species was fragmented into scattered popu- lations confined to higher elevations. (See Durrant 1952 for further discussion of the influence of Pleistocene Lake Bonneville on the distribution and speciation of mammals.) I never trapped Lagurus in the Oquirrh Mountains, and I know of no fleas being saved from sagebrush voles collected there. Durrant (1952) reported collecting speci- mens of L. c. intermedius from a locality on the east side of the Oquirrh Mountains in Salt Lake County, and there is reason to be- lieve that M. c. princei and A. sibirica will eventually be found there. I collected M. abantis from Z. princeps and M. clantoni ssp. from L. curtatus; the specimens were trapped within 10 yards of each other in southeastern Oregon. Thrassis bacchi johnsoni Hubbard is the only known Lagurus flea that has not been found in Utah. Two specimens of Thrassis that I found on a sagebrush vole collected in Uinta County, Wyoming, a few miles from the Utah-Wyoming boundary were identified by Stark as Thrassis bacchi ca- ducus (Jordan). Because of their implication in plague elsewhere (Hubbard 1949 and others), all of Utah's Lagurus fleas should be added to Allred's (1952) Utah list of plague- important fleas. Assistance from two former associates, J. G. Bittmenn and J. A. Petrovich, is recog- nized with thanks. Suggestions for improv- ing the manuscript were made by D. M. Al- Ired, C. D. Jorgensen, H. E. Stark, and V. J. Tipton. Most mammal names are according to Durrant (1952). This work was accom- plished in part while I was a research mam- malogist at the University of Utah. Literature Cited Allred, D. M. 1952. Plague-important fleas and mam- mals in Utah and the western United States. Great Basin Nat. 12(l-4):67-75. DuRRAMT, S. D. 1952. Mammals of Utah. . . . University of Kansas, Mus. Nat. Hist. 6:1-549. Egoscue, H. J. 1966. New and additional host-flea as- sociations and distributional records of fleas from Utah. Great Basin Nat. 26(3-4):71-75. Hubbard, C. A. 1949. Fleas of the sagebrush meadow mouse. Entomological News. 60 (6): 141-144. 1954. My last new North American fleas. En- tomological News. 65 (7): 169-175. Stark, H. E. 1958. The Siphonaptera of Utah. U. S. Dept. of Health, Education, and Welfare, Com- municable Disease Center, Atlanta, Georgia. 239 pp. INTERPOPULATIONAL VARIATION OF BLOOD PROTEINS IN PIKA (OCHOTONA PRINCEPS SAXATILIS) John T. Brunson', Richard N. Seaman^ and Donald J. Nash' Abstract.— Studies were undertaken to examine the degree of divergence in four populations of pika in Colo- rado. Separation of blood proteins was accomplished utilizing acrylamide-gel disc electrophoresis. Separate analy- ses of variance were carried out for the relative mobilities of two of the transferrins (designated RM)9j and RM/82), the mean relative distance difference of the two tranferrins (DM), and the density of the most prominent albumin (oj) and transferrin {^2) bands. Although the four populations were characterized by a general similarity of the gel patterns, both interpopulational and sexual variations were obser>ed. Variations between the sexes had to do with the amounts of protein, not with differences in protein mobility. The only significant populational dif- ferences were found in reference to density and DM. The interpopulational differences may serve as an indicator of populational divergence that has not been indicated by standard morphological characters. The significant pat- terns of variation observed in the blood proteins of the four populations studied may reflect a lack of gene now between isolated populations. North American pika are boreal mammals that occur in "island" populations on moun- tains throughout much of western North America. Pika are restricted to a narrowly defined habitat (the talus zone), and move- ment between neighboring populations is likely to be quite limited. Smith (1974) has estimated, for example, that distances great- er than 300 m appeared to pose difficult barriers to dispersing juveniles. The extent of genetic isolation among dif- ferent populations is largely unknown, and studies aimed at determining geographic variation in pika are necessary. The sub- species of Ochotona princeps were revised by Howell (1924), and a synopsis is provid- ed by Hall (1951). A totaf of 35 subspecies are currently recognized. The subspecies do differ in their cranial dimensions, but pre- vious reports have indicated that there is little divergence in pika populations on the basis of size morphology. The present study was designed to exam- ine variations in blood proteins in four pop- ulations of Ochotona princeps saxatilis in Colorado. The interpretation of electrophoretic mo- bility of various blood constituents as a means of discovering differences between separate populations has received much dis- cussion and criticism over the past few years. However, the fact that most of these substances are partly protein, thus bearing a close translational relationship to the gene itself, points up at least hypothetically the value of such investigations. Methods Collecting Localities: A total of 41 pika were collected from four localities in Colo- rado. The areas were Evans and Goliath in Clear Creek County, Audubon in Boulder County, and Crown Point in Larimer Coun- ty. The ranges of elevations over which ani- mals were collected at each site were 3,902-4,085 m, 3,537-3,659 m, 3,598-3,841 m, and 3,354-3,506 m, respectively. Blood Samples: Animals were shot and blood was withdrawn from the heart usually within three minutes. Heparinized 1.4-1.6 mm micro-hematocrit capillary tubes were filled to within one-half inch of one end, and were subsequently made airtight at both ends with plastic Crito-caps. All sealed hematocrit tubes were placed in glass vials. 'Department of Zoology. State University of New York, Oswego, New York 13126. 'Sinclair Research Farm, University of Missouri, Columbia, Missouri 65201. 'Department of Zoology and Entomology, Colorado State University, Fort Collins, Colorado 80523. 77 78 Great Basin Naturalist Vol. 37, No. 1 packed in snow, and shielded from sunlight during transport to the laboratory. Immedi- ately upon arrival in the laboratory, he- matocrit tubes were centrifuged at 11,500 rpm using an lEC microcapillary centrifuge. The tubes were broken at the cell-plasma interface, and the plasma was frozen until electrophoretic determinations were made. Electrophoresis: Electrophoresis of plasma proteins from 41 animals was accomplished using the aery lam ide-gel disc technique (7 percent acrylamide) described by Smith (1968). The electrophoretic chamber (Buchler In- struments) held a total of 12 gel tubes. Cur- rent was applied initially for 12 minutes at 1.5 milli-amps/gel in a tris-glycine buffer at pH 9.5. Following this initial warm-up peri- od, gels were run at 4.0 milli-amps/gel for 30 minutes. A gelman six-volt power source was used for all separations. No spacer or sample gel was employed, and 20 jul of plasma was mixed with 40 fx\ of 50 percent sucrose and was placed directly on the nm- ning gel. Tubes containing purified bovine albumin were mn as controls. Following separation, gels were stained for 45 minutes in analine blue-black. Des- taining required approximately five days and was accomplished in a series of three 7 percent acetic acid baths. Whole gels were analyzed densitometrically using a Densi- cord recording electrophoresis densitometer (Photovolt Corporation). An integrater (In- tegraph Model 49) automatically measured the areas under the densitometric curves. Analysis of densitometric curves followed the procedures outlined by Dalby and Lil- levic (1969). Results and Discussion Table 1 summarizes mean values resulting from measurements of densitometric trac- ings of the stained acrylamide gels. Because no tests were run to confirm the chemical identity of the protein bands, RM^j and RM^.2 refer only to relative mobilities of what are presumed to be the same two bands in the transferrin range. Likewise, a^ and )8j are measurements of the height of the densitometric curve of the most promi- nent presumed albumin and transferrin band respectively. The parameter DM represents 2(RM)82-RM/3J for each of the eight sample groups in- vestigated. Analyses of variance were run on RM/S^ RM^2' albumin and transferrin heights, and the mean relative distance (DM). Patterns of statistically valid sexual variation were seen in regard to the height of both the a and P bands; sexual differences in RMySj, RM/Sg, and DM were not significant. Sta- tistically significant differences among geo- graphical collecting areas were found only in reference to ^8^ height and DM. None of Table 1. Electrophoretic data: Means ± Standard Deviations. Area Sex Sample Size RM^, RM)82 ^1 Pi DM Evans Evans Males Females 5 10 0.569 ±0.004 0.575 ±0.015 0.717 ±0.005 0.718±0.014 21.800 ±0.447 21.700 ±0.483 10.000±2.121 10.300 ±2.907 0.147 ±0.001 0.143 ±0.010 Audubon Audubon Males Females 6 3 0.,576± 0.006 0..575± 0.007 0.698 ± 0.006 0.716 ±0.003 2 1.166 ±0.408 21.666 ±0.,577 6. 166 ±1.602 7..3.33± 1.154 0.122 ±0.003 0.140 ±0.005 Goliath Goliath Males Females 3 0..569 ± 0.010 0..568± 0.007 0.717 ±0.008 0.721 ±0.011 21.666±0.,577 21.8.57 ±0..377 7.666 ±0.577 11. 285 ±.3.0.39 0.148 ±0.002 0.153 ±0.008 Crown Crown .Males Females 4 3 0.565 ±0.006 0..569± 0.006 0.704 ±0.024 0.702 ± 0.039 21.0(K)± 0.816 22.000 ±0.000 ,5. 7,50 ±1.500 8.666 ±1.. 527 0.1.39 ±0.023 0.1.32 ±0.0,36 March 1977 Brunson, et al.: Pika Blood Proteins 79 the Area X sex interactions were significant. The main objective in regard to exam- ination of patterns of electrophoretic plasma protein migration was to uncover possible interpopulational variations of a striking nature. If these variations were to exist, they perhaps might be an indication of a stable phenotypic characteristic, which, when viewed with phenotypic characters of a morphological type, might indicate a high or low degree of populational divergence. That this divergence might be expected is based on the relative altitudinal isolation of pika populations in Colorado. The fact that high talus regions are generally separated by non-talus areas and the general reluc- tance of the pika to leave the talus habitat seems to preclude the necessity of distance as a block to gene flow. This does not mean to suggest that gene flow is the salient fac- tor causing population divergence. It is pos- sible that even in the absence of continued gene exchange, these animals have evolved in parallel fashion due to the relative sim- ilarity of selective pressures. Crown Point and Mount Audubon are widely separated from each other as well as from Mount Evans and Mount Goliath. There would seem, in these instances, to be a reduced amount of reproductive activity between populations. Because they are sep- arated by a distance of only about five miles, Evans and Goliath might be treated as a single population. It is possible, how- ever, that gene flow even between these two areas has been minimal or non-existent. Although divergence of three of these populations might be expected on the basis of restricted gene exchange, the present classifications of North American pikas are based entirely on morphological characters which, with the exception of pelage color, seem to show little variation (Broadbooks 1965). It was deemed worthwhile, then, to examine plasma protein migration patterns as a more precise indication of possible populational differences. In interpreting any variation found in a study of this nature, one must resist the urge to attribute them all to stable gen- otypic characteristics. Indeed, more infor- mation is needed from breeding experiments and from expanded analysis of individual protein components before a more con- fident stand can be taken in regard to the genotypic stability of a difference. There is no question that the literature warns against over-zealous interpretation of any measured protein variations. It might be prudent, therefore, to approach interpretation of data of this type from the standpoint of similarities in migration patterns. Although patterns of both inter- populational and sexual variation are pres- ent, there is a general similarity of the gel patterns from the four areas. In addition to the obvious height differences in some curves, the presence of a dense band very close to the origin in some samples from both sexes from Mount Evans and from Mount Goliath are the most noticeable dif- ferences in an otherwise generally common pattern. The presence of significant sexual varia- tion is not surprising, as was pointed out by Moore (1945). Female animals do seem to have denser j3-^ and a^ bands than do males. It is interesting that thiere seem to be differ- ences only in the amount of a protein rather than the type of protein molecule, as there were no significant patterns of varia- tion in protein mobility between sexes. The significant pattern of variation between lo- cations in regard to the fi^ band is probably due to bias introduced by the larger number of female samples from some locations. The significant differences in DM be- tween geographic areas seem to indicate differential migration rates of the entire samples. The DM is, in fact, a measure of the overall linear variation in both the gels (and consequently the tracings), and it is precisely why the tracings do not super- impose exactly. These interpopulational dif- ferences seem to be real. That they are stable genetic characters cannot be deter- mined from this analysis; they may, how- ever, be an indicator of population diver- gence that cannot be measured in terms of standard morphological characters. Although interpretation of electrophoretic plasma protein migration patterns should be approached with caution, such inter- pretation has the potential of uncovering 80 Great Basin Naturalist Vol. 37, No. 1 more subtle differences that may not be mirrored in standard taxonomic characters. At this point, one can only say that the sig- nificant patterns of variation found in re- gard to plasma protein mobility may sup- port the possibility of a predicted lack of gene flow between isolated populations of pika. Literature Cited Broadbooks, H. E. 1965. Ecology and distribution of the pikas of Washington and Alaska. Amer. Midi. Nat. 73(2):299-335. Dalby, p. L., and H. a. Lillevik. 1969. Taxonomic analysis of electrophoretic blood serum patterns in tne cotton rat, Sigmodon. Mich. State Univ. Publ. Mus., Biol. Sec. 4(3):65-104. Hall, E. R. 1951. A synopsis of the North American Lagomorpha. Univ. Kansas Publ., Mus. Nat. Hist. 5:119-202. Howell, A. H. 1924. Revision of the American pikas (Genus Ochotona). N. Amer. Fauna 47:1-57. Moore, D. H. 1945. Species differences in serum pro- tein patterns. J. Biol. Chem. 161(l):21-32. Smith, A. T. 1974. The distribution and dispersal of pikas: consequences of insular population struc- ture. Ecology 55(5):1112-1119. Smith, I. (Ed.) 1968. Chromatographic and electro- phoretic techniques. Vol. 2. Zone electro- phoresis. Wiley, New York, 524 pp. OBSERVATIONS ON THE DISTRIBUTION OF MICROORGANISMS IN DESERT SOIL A. T. Vollmer', F. Au-, and S. A. Bamberg' Abstract.— Population estimates of fimgi, bacteria, and actinomycetes in desert soil were determined with re- spect to soil depth and distance from shrubs. In general the highest numbers of microbes were found at the shrub base; the lowest numbers were found in the interspaces. While the total number of organisms usually de- clined in deeper soil, the relative importance of the actinomycetes increased. These population trends are attri- buted to substrate availability and utilization and interspecific interactions. As the soils became drier and warmer the total number of microorganisms decreased. Mold populations re- mained at about the same level during the study. While the numbers of both bacteria and actinomycetes de- clined, the relative importance of the actinomycetes increased. The numbers and activities of soil micro- flora are important in desert soils in the processes of root and litter decomposition, for the timing and release of nutrients tied up in dead organic matter, and also for the physical binding of soil particles (Went and Stark 1968, Khudairi 1969, Fuller 1974). For this report we determined spatial and temporal differences in the populations of fimgi, bacteria, and actinomycetes in Mo- jave Desert soils. This work was conducted as part of a large study of primary productivity and nu- trient interrelationships sponsored by the U.S. International Biological Program (IBP). Soils were gathered from the IBP Desert Biome's Rock Valley validation site, Nye County, Nevada. This area is in the north- ern Mojave Desert about 100 km northwest of Las Vegas, and is part of the U.S. Energy Research and Development Administration's Nevada Test Site. In Rock Valley soils are well-drained and have moderate per- meability. Soil beneath shrubs is fine sand to depths of around 15 cm, gravelly loamy sand to 33 cm, and cemented gravel be- tween 33 and 57 cm. Soils in bare areas are gravelly sandy loam to around 43 cm, and cemented gravel below that (Romney et al. 1973). The vegetation is composed primarily of perennial shrubs, which cover about 20 percent of the surface. More de- tailed descriptions of edaphic and biotic features may be found in Turner (1974). Methods Soil was sampled from nine locations dur- ing the middle of the plant growing season, March-April 1973, around the shrub species, Lycium andersonii Gray. Samples were col- lected from three depths (0-10 cm, 10-20 cm, and 20-30 cm) at each of three dis- tances from the shrub: the shrub base (sam- ples 1, 2, and 3, starting at the surface), at the canopy edge or one shrub radius (sam- ples 4, 5, and 6), and in the shrub inter- space or three-canopy radii from the plant base (samples 7, 8, and 9). These soil sam- ples were aliquots of those used in other be- low-ground studies of root biomass and soil ATP activity (Bamberg et al. 1974). Dilution plate counts were made to quan- tify and differentiate the fungal and bacte- rial populations of the soil samples collected during the 12th, 13th, 15th, and 16th weeks of 1973. Twenty grams of oven-dried soil (ODS) from each soil .sample were added to 'Laboratory of Nuclear Medicine and Radiation Biology, University of C;alifornia, Los Angeles, California 90024. Environmental Monitoring and Support Laboratory, Environmental Protection Agency, Las Vegas, Nevada 8910 'Present address: Stone and Webster Engineering Corp., P.O. Box 5406, Denver, Colorado 80217. 81 82 Great Basin Naturalist Vol. 37, No. 1 100 ml of sterile, distilled water and were shaken for 10 minutes. Coarser particles were allowed to settle, and one-nil samples of the soil suspension were used to prepare dilutions for plate counts according to standard procedures (Clark 1965). The dilu- tions used were 5X10"^ and SXIQ-^ for fungi and SXIO"* and SXIO"'^ for bacteria- actinomycete counts. Five plates of each di- lution were poured. Cooke's Rose Bengal agar with 35 jUg Aureomycin/ml added was used for fimgal determinations, and sodium albuminate agar was used for bacteria and actinomvcetes. The plates were incubated at room temperatures. Mold counts were made after two days, and bacteria and actinomv- cetes were counted after 14 days. Penicillium, Aspergillus, and Streptomyces were differentiated and their percents of oc- currence were reported. Soil pH was measured using a 1:5 soil so- lution. Determinations of soil moisture were made after the soil was oven-dried at 105- 110 C for 24 hours. Total numbers of or- ganisms per gram of oven-dried soil were calculated according to the method of Clark (1965). Root biomass was determined by a com- bination of sieving and flotation in saturated MgSO^ solution (see Bamberg et al. 1974). Results Soil pH averaged about 9.0 and did not vary significantly with either location or time. Soil temperature and moisture are presented in Table 1. Temperatures, which were taken in the morning, showed little variation within a sampling period with the exception of Lo- cation 7. As one might expect, this surface sample in the interspace was slightly warm- er than soil from the other locations. Soil temperatures for April during the 15th and 16th weeks averaged 5-10 C higher than those from March. .•\t the shrub base and the canopy edge the soil moisture decreased from about 15 percent during weeks 12 and 13 to about 5 percent during weeks 15 and 16. With the exception of week 13, little change in soil moisture was noticed over this period of time at the interspace locations. The high moisture levels for week 13 were the result of 25 mm of rain that fell during the period between the first and second sampling dates. Moisture content in the interspace was slightly lower at week 12 and higher at week 16 than in the other locations. There was considerable variation in root biomass under Lijcium andersonii during the study (Table 2). Most of the roots (64 per- cent) were found under the shrub and the least (less than 7 percent) were present in the interspaces. Root biomass at the shrub base usually decreased with depth, while vertical distribution was fairly uniform at the canopy edge and in the interspaces. Flowering and continued growth occurred during the sampling period; fruit formation had started by the end of this period. Estimates of the population sizes of fungi, bacteria, and actinomycetes are given in Table 3 for soil samples collected between the 12th and 16th weeks of 1973 from Rock Valley. Table 4 and Figure 1 show the per- centage of the total mold population made up of organisms of the genera Penicillium and Aspergillus. Over 95 percent of the ac- tinomycetes isolated belonged to the genus Streptomyces. Results are summarized according to loca- tion: 1. Shrub Base: At the plant base the mold numbers in the surface soil increased during the course of the study. As the spring prog- ressed the number of fimgi decreased by about half at the 20-30 cm level. Penicil- lium was the dominant fungal genus under- neath the shrub throughout the five-week period. The relative importance of Asper- gillus dropped off during weeks 15 and 16. Populations of bacteria and actinomycetes were much higher than those of the fungi. In April the numbers of bacteria decreased by 55-85 percent. During the study Strepto- myces increased its dominance under the shrub from 50 percent to 80 percent, even though actual numbers declined. Bacterial population size estimates did not appear to be related to sample depth. 2. Canopy Edge: Soil from the canopy edge taken at locations 4, 5, and 6 yielded mold population estimates that did not March 1977 Vollmer: Soil Microorganisms 83 change radically during the experiment. Soil from Location 6 usually yielded a lower number of fungi than did soil from shallow- er depths. PeniciUiwn dominance over As- pergillus increased sharply during weeks 15 and 16. Bacteria and actinomycete numbers were lower at the canopy edge than they were underneath the shrub. Population size and dominance trends of these two groups fol- lowed those found at the plant base. 3. Interspaces: The lowest total numbers of fungi were found in the shrub interspace. Mold counts in April were highest at the 0- 10 cm level. In the shrub interspaces, As- pergillus replaced Penicillium as the domi- nant genus. The populations of bacteria and acti- nomycetes at three canopy radii from the shrub base were slightly lower than those at the canopy edge. With the exception of week 13, there was little difference in pop- ulation size estimates with time or depth. Streptoniyces continued its strong dominant role. Discussion Generally both root biomass and total number of microbes were highest at the shrub base. This relationship indicates that the soil microflora utilize the roots or root exudates as a substrate (Starkey 1958, Fried- man and Galun 1974). It must be pointed out, however, that other types of organic matter, such as litter, had a distribution pat- Table 1. Soil temperature and moisture at nine sample locations during March and April 1973. Sample Location Week Shrub Base Canopy Edge Interspace 1 2 3 4 5 6 7 8 9 Temperature (C) 12 8.5 9.0 9.5 9.0 9.0 9.5 10.0 9.0 8.5 13 8.5 9.0 10.0 9.0 9.0 10.0 10.9 9.0 9.0 15 18.0 16.5 16.0 17.0 15.0 15.0 24.0 18.0 16.0 16 14.5 12.5 13.0 1.3.0 12.0 Moisture (%) 1,3.0 16.0 15.0 13.0 12 15 15 16 14 15 15 9 10 13 13 14 16 17 16 19 17 17 19 17 15 5 6 6 5 6 6 7 7 7 16 5 6 3 5 7 ' 10 13 Table 2. Root biomass (gODW/1 soil) in relation to sample location under Lyciiim andersonii in Rock Valley, 1973. Sample Locat on Week Shrub Base Canopy Edge Interspace 1 2 3 4 5 6 7 8 9 12 5.5 2.9 2.8 1.1 6.3 1.7 <0.4 <0.6 <0.4 13 1.4 8.9 1.5 6.0 1.6 2.6 <0.3 <0.2 <0.3 15 3.9 2.1 2.9 1.5 1.6 0.4 1.4 0.5 0.3 16 14.9 8.8 10.3 1.7 1.1 5.2 <0.3 0.3 1.9 84 Great Basin Naturalist Vol. 37, No. 1 tern similar to that of roots (Bamberg et al. 1974). Fungal populations remained relatively stable as the season progressed. Bacteria and actinomycetes, however, decreased with time. These changes in the microbial com- munitv are probably related to soil mois- ture. Soil water potentials reached about -25 bars in April in Rock Valley. Bacterial activity is known to decline rapidly when the soil water potential drops below -5 bars, while many fungi and actinomycetes can tolerate much lower water potentials (Griffin 1972). Increases in the relative im- portance of streptomycetes and fungi may be partially attributable to reduced com- petition by bacteria. The percent of Strepto- mijces increased in April, which may in- dicate the beginning of a more active decomposition role assumed by this group. This, increase is particularly evident at the shrub base and canopy edge. PeniciUium was the dominant mold genus close to the plant, but Aspergillus was also common in the surface soils and was domi- nant in the interspace zone. Since these two genera are spore-formers, plate counts may only indicate the potential rather than the actual activity during the sampling period. The sample locations at which Aspergillus numbers were highest are also those that usually become the warmest and driest dur- ing the year. These results agree with re- ports that have found Aspergillus to be a more xerothermic genus than PeniciUium (Griffin 1972). As mentioned previously, roots, root exu- dates, and other substrate matter influence the soil microflora. Mold populations were found to decline in deeper soil, while Streptotnyces dominance increased with soil depth. These differences may have resulted because the availability of preferred nutri- ent sources varied with soil depth. Most of the roots were located in the upper 20 cm of the soil. Siu (1951) reported that acti- nomycetes were generally poor cellulose utilizers, while fungi were highly cellulolyt- ic. Went and Stark (1968) postulated that fungi may play an important role in deserts not only in decomposition of soil organic matter Ijut also in the direct cycling of nu- trients to living roots from dead organic material. Table 3. Plate counts from soil gathered at nine locations during March and April 1973 (Organisms /gODS). Sample Location Week Shrub Base Canopy Edge Interspace 1 2 3 4 5 6 7 8 9 Fungi X103 12 25.5 24.9 .30.8 23.0 24.5 8.5 9.9 13.1 3.9 13 41.4 22.4 26.1 9.3 23.0 9.8 3.8 3.4 9.2 15 44.7 50.3 16.3 20.1 23.5 12.7 13.2 9.0 5.2 16 47.8 27.8 16.8 18.3 15.1 10.3 7.3 3.6 4.7 Bacteria XIO^ 12 .34.4 93.0 78.8 26.6 63.7 13.6 9.6 13.8 3.3 13 82.2 54.4 20.8 .34.3 22.3 22.3 23.1 38.0 16.6 15 5.8 16.9 4.6 11.8 9.4 11.5 18.4 10.7 13.3 16 15.6 19.9 10.4 7.0 Actinomycetes 9.9 6.8 17.3 4.8 5.9 12 51.6 73.0 67.2 45.4 66.3 54.4 19.4 29.2 33.7 13 72.9 92.6 78.2 5.3.7 70.7 74.7 53.9 72.0 66.4 15 .39.2 48.1 18.4 35.3 29.6 25.5 .35.6 26.3 24.7 16 .36.4 63.1 58.7 .30.0 33.1 38.3 16.7 25.2 21.1 March 1977 Vollmer: Soil Microorganisms Table 4. Percentage of total mold colonies composed of the genera Penicillium and Aspergillus. 85 Sample Location Week Shrub Base Canopy Edge Interspace 1 2 3 4 5 6 7 8 9 Penicillium 12 17 54 93 23 48 40 6 21 13 13 35 50 32 21 60 16 0 15 7 15 71 72 58 50 50 29 14 19 29 16 73 25 36 44 Aspergillus 75 56 3 6 4 12 13 15 4 10 42 12 62 61 10 13 27 13 8 27 17 16 47 71 1 15 2 3 3 4 4 2 78 57 64 16 1 6 21 8 9 22 71 39 19 A D 80 - - 60 - y\ - 4 0 - ^^ .--7^0 " 20 ~ '^ '^^--o--^ / B E 80 - - 60 - . - 4 0 20 - ^ ^ ,» ""^^ C F 80 - - 60 - ,,\ - 40 - •--,''' \ /-''^ ' 20 - 1 r'^ 1 1 1 1 1 1 WEEK Fig. 1. Percentage of all fungi represented by the genera Penicillium (solid line) and Aspergillus (dashed line) at: A. Shrub Base (locations 1, 2, and 3); B. Cano- py Edge (4, 5, 6); C. Interspace (7, 8, 9); D. 0-10 cm (1, 4, 7); E. 10-20 cm (2, 5, 8); F. 20-30 cm (3, 6, 9). Alexander (1971) stated that the extant conditions in a habitat will dictate that only one, or a few, of the indigenous species will actually exploit an energy source. The fact that seasonal variation in the soil environ- ment will cause a shift in the relative num- bers of various microbial populations is seen by the higher percentage of Streptomyces in week 16 over week 12 at the shrub base lo- cation. This response corresponds with a de- crease in moisture and an increase in tem- perature during that time interval. The numerical dominance of Streptomyces in this study may be attributable to the warm, dry nature of our desert soils. Throughout the sampling period acti- nomycetes outnumbered bacteria; fungi rep- resented only a small portion of the total number of microorganisms. These results agree with those reported for other desert soils (Fuller 1974). It has been pointed out, however, that enumeration techniques alone do not necessarily give an accurate in- dication of the importance of the various microbial groups. For example, while these methods usually give low population esti- mates for molds, fungal biomass often ex- ceeds that of other microorganisms (Went and Stark 1968, Fuller 1974). We feel, however, that differences in the number of observed colonies within the var- Great Basin Naturalist Vol. 37, No. 1 ions groups of microorganisms are good in- dicators of spatial and temporal fluctuations in population size. These population trends in turn give an indication of changes in the relative importance of the diff^erent micro- bial groups. Acknowledgments This work was supported by the U.S. /IBP Desert Biome Program and under contract E(04-1)GEN-12 between the U.S. Energy Research and Development Administration and the University of California. Literature Cited Alexander, M. 197L Microbial ecology. Wiley and Sons, New York. 511 pp. Bamberg, S. A., A. Wallace, G. E. Kleinkopf, A. VoLLMER, AND B. S. AusMus. 1974. Plant pro- ductivity and nutrient interrelationships of per- ennials in the Mojave Desert. US/IBP Desert Biome Res. Memo. 74-8. Utah State Univ., Lo- gan. 16 pp. Clark, F. E. 1965. Agar-plate method for total micro- bial counts, pp. 1460-1466. In C. A. Black (ed.). Methods of soil analysis. Part II. Chemical and microbiological properties. Amer. Soc. Agron., Madison. Friedman, E. I. and M. Galun. 1974. Desert algae, lichens, and fungi, pp. 165-212. In G. W. Brown, Jr. (ed.). Desert biology, volume II. Aca- demic Press, New York. Fuller, W. H. 1974. Desert soils, pp. .31-101. In G. W. Brown, Jr. (ed.). Desert biology, volume 11. Academic Press, New York. Griffin, D. M. 1972. Ecology of soil fimgi. Syracuse Univ. Press, Syracuse. 193 pp. Khudairi, a. K. 1969. Mycorrhiza in desert soils. Bio- science 19:598-599. RoMNEY, E. M., V. Q. Hale, A. Wallace, O. R. LuNT, J. D. Chidress, H. Kaaza, G. V. Alexander, J. E. Kinnear, and T. L. AcKERMAN. 1973. Some characteristics of soil and perennial vegetation in northern Mojave Desert areas of the Nevada Test Site. USAEC Report, TID-4500, 340 pp. Siu, R. G. H. 1951. Microbial decomposition of cellu- lose. Reinhold, New York. 531 pp. Starkey, R. L. 1958. Interrelationships between micro- organisms and plant roots in the rhizosphere. Bacteriol. Rev. 22:154-172. Turner, F. B. (ed.). 1974. Rock Valley validation site report. US/ IBP Desert Biome Res. Memo. 74-72. Utah State Univ., Logan 64 pp. Went, F. W. and N. Stark. 1968. The biological and mechanical role of soil fungi. Proc. Natl. Acad. Sci. 60:497-504. HOME RANGE OF THE WESTERN JUMPING MOUSE, ZAPUS PRINCEPS, IN THE COLORADO ROCKY MOUNTAINS Nathaniel Stinson, Jr.' Abstract.— Western jumping mice, Zapiis princeps, were live-trapped during the summer of 1975 in an aspen forest in the Front Range of the Colorado Rocky Mountains. Home range sizes were calculated using the exclu- sive boundary strip method, and it was discovered that males had larger home ranges (298-.3315 m^, x = 1743 m^) than females (680-1275 ni", x = 1041 m'^). Mean range length was 96.5 m for males and 78.5 m for females. Spatial relationships showed that females tended to be more territorial than males, based on the females' higher percentage of exclusive home ranges, greater distance between centers of activity, and more uniform spacing. Jumping mice (Zapodidae) are widely dis- tributed over North America. The two east- ern species {Zapus hudsoniiis and Napaeoz- opus insignis) have been the subject of considerable research (e.g. Snyder 1924; Sheldon 1934, 1938a, 1938b; Hamilton 1935; Blair 1940; Quimby 1951; and Whita- ker 1963a, 1963b). However, ecological lit- erature on the western jumping mouse {Zapus princeps) is sparse. Krutzsch (1954) briefly discussed the natural history of Z. princeps in the Rocky Mountains, and Brown (1967a, 1970) has provided informa- tion on its reproductive and .seasonal activi- ty patterns. Although information is avail- able on habitat selection by Z. princeps (Brown 1967b, Clark 1971, Armstrong 1972), little is known of their spatial pat- terns within a single habitat type. Analysis of these spatial patterns was the thrust of the present study. Materials and Methods The study was conducted in an aspen for- est adjacent to the University of Colorado Mountain Research Station, Nederland, Col- orado, at an elevation of approximately 2900 m. This forest was mesic and willow {Salix); monkshood (Aconitum columhia- mnn), cowparsnip {Heracleiim lanatimi), Ca- nadian reed grass {Calamagrostis cana- densis), and bluegrass {Poa spp.) dominated the understory. The study area consisted of an 8 x 10 grid of trapping stations spaced 10 m apart. Three small Sherman live traps were placed at each .station. The study area was sampled for four days twice a month during July, August, and September 1975. On capture, each individual was sexed, weighed, marked by toe-clipping, and then released. Location of capture for each individual was also re- corded. The exclusive boundary strip method was used to measure home range size. The ex- ternal points of capture are connected, and the resultant area is considered the home range of the individual. Centers of activity were calculated using Hayne's method (1949) and were used to assess nearest neighbor distance (Clark and Evans 1954). Results A total of 58 individuals was captured during the summer of 1975 in 5760 trap- nights, representing an average monthly density of 31.0/ha (range 28.0-35.0). This density was extremely high compared to values obtained by Brown (1970), whose population densities averaged 3.2/ha over a four-year period. Population numbers were stable on a monthly basis, probably due to Department of Environmental, Population, and Or(»anismic Biology. University of Colorado, Boulder, Colorado 80309. 87 88 Great Basin Naturalist Vol. 37, No. 1 the high longevity characteristic of Zapits (Quinibv 1951, Brown 1970). All animals captured five or more times were used in the calculation of home range size (N = 37). The average home range size of males was 1743 m^ (298-3315 m'-^), and that of females was 1041 m^ (680-1275 m2). The mean range length (greatest distance between captures) was 96.5 m for males and 78.5 m for females. These differences be- tween the sexes were highly significant (p < .01, t-test). Brown (1970) obtained mean range length values twice the present values (210.0 m and 155.0 m respectively), but this is because the population of Zapus princeps that he studied showed an extremely nar- row, elongated distribution along a moun- tain stream. Discussion Home Range Size Home range, as applied to mammals, has been defined by Burt (1943) as "the area transversed by the individual in its normal activities of food gathering, mating, and caring for young." Home range values for Zapus princeps in this study were consid- erably lower than those found by Myers (1969). Myers obtained home range values of 3075 m^ for males and 2350 m^ for fe- males. There are several possible reasons for the discrepancy: (1) track- versus trap-re- vealed home ranges, (2) different population densities, and (3) differences in vegetation structure of the habitat. Myers used the tracking method of Justice (1961) to mon- itor animal movements, and the results of track- versus trap-revealed home ranges do not seem to be directly comparable. Met- zgar (1973) found that track-revealed home ranges were larger than trap-revealed home ranges. Myers captured a total of 25 Zapus prin- ceps throughout the whole summer on a 1.1 hectare plot, whereas in the present study the average monthly density was 31.0/hec- tare. It has been well documented that home range size varies inversely with popu- lation density (Burt 1943, Getz 1961, Van Vleck 1969, ' Mazuriewicz 1971, and Maza et al. 1973). Also, Myers does not describe the habi- tat's vegetation structure, which may in- fluence home range size. In the present study, the aspen forest was very productive, and Kenagy (1973) proposed that in years of high resource levels animals forage over smaller areas since in the smaller areas food is readily available and less energy expendi- ture is required for foraging. And O'Farrell et al. (1975) point out that home ranges of Perognathus parvus were affected jointly by resource levels and population size; neither of these factors acted independently. The exact reason for the large difference in calculated home range sizes is not known, but home range size is quite labile and many factors, both intrinsic (e.g. popu- lation size) and environmental, influence its size. The home range size of Zapus princeps is similar to that of the meadow jumping mouse, Z. hudsoyiius, in Minnesota. Quimby (1951) used the inclusive boundary strip method, where the external points of cap- ture are considered centers of rectangles whose boundaries are half the distance to the next trapping station, to determine home range size in Z. hudsonius. To facil- itate comparison with the present study, Quimby's values were reduced 15 percent (see Stickel 1954); this reduction results in home range sizes of 1479 m^ for males and 1307 m- for females. The similarity in home range size is not surprising, since Quimby (1951) found that the preferred habitat of Z. hudsonius in Minnesota was willow thickets— and the understory of the aspen forest in the present study was dominated by willows. McNab (1963) proposed that "hunters" (carnivores, insectivores, and granivores) re- quire a larger home range than "croppers" (herbivores). The home range size of two other species of rodents {Microtus long- icaudus and M. montanus) found in the area were calculated using the same meth- od. Zapus princeps is a primary granivore, with seeds of the graminoids being most im- portant, while members of the genus Micro- tus are primary herbivores (Lechleitner 1969, Clark 1971). The average home range of Z. princeps was significantly larger (p < March 1977 Stinson: Western Jumping Mouse .001, t-test) than that of the microtines (Table 1). The rationale for the difference in home range size is that food resources of hunters show a more dispersed distribution than that of croppers, and therefore must travel farther on foraging trips. Table 1.— Home range size of three species of ro- dents. Home Range Size Range Length Species N Male Female Male Female Microtiis longicatidus 11 485 m^ 364 m^ 42.0 m .30.5 m Microtus montanm 4 283 m^ 25.0 m Zapiis princeps ,37 1741 m^ 1012 m^ 96.5 m 78.5 m Territoriality Getz (1961) suggested that territoriality can be inferred from five criteria: (1) per- centage of exclusive (non-overlapping) home ranges between members of the same sex, (2) percentage of at least partially exclusive home ranges between members of the same sex, (3) distance between centers of activity, (4) distribution of centers of activity, and (5) sex of individuals involved in multiple cap- tures. In this study 71 percent of the males had inclusive home ranges, while no females had inclusive home ranges. Both males and females had home ranges that were partially exclusive (29 percent and 84 percent re- spectively). Sixteen percent of the female home ranges were totally exclusive. Centers of activity were calculated ac- cording to the method of Hayne (1949). The average distance between centers of ac- tivity was 11.1 m for males and 15.3 m for females. This difference in mean distance between centers of activity was significant (p < .01, t-test). Getz (1961) hypothesized that if the .sex ratio of the population is equal and if terri- torial behavior is not displayed, then the nearest neighbor should be of the same sex at least half the time. In this study, the nearest neighbor was always the opposite sex. Hanson and Fleharty (1974) believed this indicated that territorial behavior was exhibited by both sexes, but there may be an alternative explanation. In some rodent species mated pairs often share the same nest or nest in close proximity, and this may be the case with Zapus princeps. The fact that the nearest neighbor is of the opposite sex may just be a by-product of this repro- ductive behavior, and not a by-product of territorial behavior. The distribution of centers of activity should approach uniformity if territorial be- havior is displayed (Getz 1961). Clark and Evans (1954) established "R" as the ratio of the observed mean distance to nearest neighbor to the expected mean distance to nearest neighbor in an infinitely large ran- dom distribution. R has a finite range, with values indicative of perfectly clumped (0.00), random (1.00), and uniform (2.1491). Distributions of Zapus princeps ba.sed on centers of activity showed that both sexes tended towards uniform spacing (R = 1.33 for males and 1.51 for females). This differ- ence between the sexes was significant (F = 3.455, p < .01). There were no multiple captures of Zapus princeps in the present study. Quimby (1951) indicated that Z. hudsonius was a solitary feeder, and multiple captures were rare. The same may be true with Z. princeps. The analysis of spatial relationships .sug- gests that males are quite tolerant to each other (71 percent of home ranges inclusive), and that females tend to be more territorial than males, based on their higher percent- age of exclusive home ranges, greater dis- tance between centers of activity, and more uniform spacing. Acknowledgments The financial support of a National Fel- lowship Fund Predoctoral Fellow.ship is ac- knowledged with gratitude. Thanks is ex- tended to Dr. Michael Grant, director, and the staff of the University of Colorado Mountain Research Station for use of their facilities. Special thanks to Dr. Larry N. 90 Great Basin Naturalist Vol. 37, No. 1 Brown for comments on an earlier draft of the manuscript. Literature Cited Armstrong, D. M. 1972. Distribution of mammals in Colorado. Monogr., Univ. Kansas Mus. Nat. Hist. 3:1-415. Blair, W. F. 1940. Home ranges and populations of the jumping mouse. Amer. Midi. Nat. 23:244- 250. Brow.n, L. N. 1967a. Seasonal activity patterns and breeding of the western jumping mouse {Zapus princeps) in Wyoming. Amer. Midi. Nat. 78:460-470. 1967b. Ecological distribution of mice in the Medicine Bow mountains of Wyoming. Ecology 48:677-680. 1970. Population dynamics of the western jumping mouse {Zapus princeps) during a four- year study. J. Mammal. 51:651-658. Burt, W. H. 1943. Territoriality and home range con- cepts as applied to mammals. Misc. Publ. Mus. Zool., Univ. Michigan 45:1-58. Clark, P. J., and F. C. Evans. 1954. Distance to near- est neighbor as a measure of spatial relation- ships in populations. Ecology 35:445-453. Clark, T. W. 1971. Ecology of the western jumping mouse in Grand Teton National Park, Wyom- ing. Northwest Sci. 45:229-238. Getz, L. L. 1961. Home ranges, territoriality, and movement of the meadow vole. J. Mammal. 42:24-36. Hamilton, W. J. 1935. Habits of jumping mice. Amer. Midi. Nat. 16:187-200. Hanson, C. M., and E. D. Fleharty. 1974. Structural ecological parameters of a population of Per- oniyscus maniculatus in west-central Kansas. Southwest. Nat. 19:293-303. Hayne, D. W. 1949. Calculation of size of home range. J. Mammal. .30:1-18. Justice, K. E. 1961. A new method for measuring home ranges of small mammals. J. Mammal. 42:462-470. Kenagy, G. J. 1973. Daily and seasonal patterns of ac- tivity and energetics in a heteromyid rodent community. Ecology 54:1201-1219. Krutzsch, p. H. 1954. North American jumping mice (genus Zapus). Univ. Kansas Publ., Mus. Nat. Hist. 7:349-472. Lechleitner, R. R. 1969. Wild mammals of Colorado. Pruett Publ. Co., Boulder. 254 pp. McNab, B. K. 1963. Bioenergetics and the determina- tion of home range size. Amer. Nat. 97:133-139. Maza, G. B., N. R. French, and A. P. Aschwanden. 1973. Home range dynamics in a population of heteromyid rodents. J. Mammal. 54:405-425. Mazurkiewicz, M. 1971. Shape, size, and distribution of home ranges of Clethrionomys glareolus (Schreber, 1780). Acta Theriol. 16:23-60. Metzgar, L. H. 1973. A comparison of trap- and track-revealed home ranges in Peromyscus. J. Mammal. 54:513-515. Myers, L. G. 1969. Home range and longevity in Zapus princeps in Colorado. Amer. Midi. Nat. 82:628-629. O'Farrell, T. p., R. J. Olson, R. O. Gilbert, and J. D. Helund. 1975. A population of Great Ba- sin pocket mice, Perognathus parvus, in the shrub-steppe of south-central Washington. Ecol. Monogr. 45:1-28. QuiMBY, D. C. 1951. The life history and ecology of the meadow jumping mouse, Zapus hudsonius. Ecol. Monogr. 21:61-95. Sheldon, C. 1934. Studies of the life histories of Zapus and Napaeozapus in Nova Scotia. J. Mammal. 15:290-300. 1938a. Vermont jumping mice of the genus Zapus. J. Mammal. 19:.324-.3.32. 19.38b. Vermont jumping mice of the genus Napaeozapus. J. Mammal. 19:444-453. Snyder, L. L. 1924. Some details on the life history and behavior of Napaeozapus insignis abieto- rum. J. Mammal. 5:233-237. Stickel, L. F. 1954. A comparison of certain methods of measuring ranges of small mammals. J. Mam- mal. 35:1-15. Van Vleck, D. B. 1969. Standardization of Microtus home range calculation. J. Mammal. 50:69-80. Whitaker, J. O., Jr. 196.3a. Food, habitat, and para- sites of the woodland jumping mouse in central New York. J. Mammal. 44:316-321. 1963b. A study of the meadow jumping mouse, Zapus hudsonius (Zimmerman), in cen- tral New York. Ecol. Monogr. 33:215-254. PRELIMINARY MAMMAL SURVEY OF FOSSIL BUTTE NATIONAL MONUMENT, WYOMING Tim W. Clark' Abstract.— The mammalian community of Fossil Butte National Monument, Wyoming, consisted of at least 45 species. Snap trap sampling of the smaller species combined with sight and sign observations of the larger species formed the basis of this preliminary survey in the summers of 1974 and 1975. Snap-trapping samples (2880 trap days) yielded 44 least chipmunks and 89 deer mice. The distribution of all species was characterized by plant community occupancy. Heavy domestic livestock use of the communities nearest water has resulted in severe alteration in community structure; this has probably affected the distribution and abundance of at least some of the smaller mammals. Fossil Butte National Monument, world famous for its ichthyological fossils, is a re- cent addition (1973) to the National Park System; it is about 18 km west of Kemmer- er, Lincoln County, Wyoming. A prelimi- nary survey of the mammals of the mon- ument was made in the summers of 1974 and 1975 and may serve to develop public understanding and appreciation of the fauna and to promote more comprehensive ecological investigations. Study Area Fossil Butte (3313 m above sea level) is a ruggedly impressive topographic feature ris- ing from 2272 m at base. The climate is semiarid and cool-temperate. Annual pre- cipitation, most of which falls as snow, av- erages about 23 cm. Winters are cold; sum- mer days are warm, but summer nights are cool. The monument lies in the Rocky Moun- tain Faunal Area (Long 1965). However, the Idahoan and Upper Green River divi- sions of the faunal area meet in the general area of the monument, bringing several spe- cies groups into contact. Six major plant communities, indentified on the basis of physiognomy and taxonomy of the pre- dominant overstory vegetation, are de- scribed in Table 1. The vegetation of Fossil Butte was described previously by Beetle and Marlow (1974); they divided the area into 13 plant communities based on floral distribution. The correspondence of their communities to mine is listed in Table 1. Many of the communities listed by Beetle and Marlow (1974) are only a few meters wide and, as such, their boundaries were not meaningful to many mammalian species, especially the larger forms. Therefore, a lumping of their community categories seems necessary. Beetle and Marlow (1974) listed the species in each community. Vege- tation of the monument appears character- istic of the surrounding area. Methods Methods depended on the species in- vestigated. Ecological distribution of larger kinds (e.g., beaver, carnivores, ungulates) was determined by direct observation of an- imals or of their signs. Smaller mammals were sampled using snap traps. Snap-trap transects of 20 stations 15 m apart were placed in each plant community. Three traps were set at each station. Traps were baited with rolled oats and peanut butter and were checked in the mornings and eve- nings of four consecutive days. Results Twenty-three mammalian species were Department of Biology, Idaho State University, Pocatello, Idaho 8.32()9. 91 92 Great Basin Naturalist Vol. 37, No. 1 found on the monument, representing five orders, 11 families, and 22 genera. In addi- tion, other unverified species no doubt oc- cur on the monument. Snap-trapping,.- In 2880 trap days (TD) equally spaced among the six plant commu- nities (Table 1), 134 small mammals were caught. The catch included only two spe- cies, the least chipmunk {Eutamias min- imus) (N = 45) and the deer mouse (Per- omyscus maniculattis) (N = 89). Both species were caught at relatively low rates (1.5 cap- tures per 100 TD for chipmunks, and 3.1 captures per 100 TD for deer mice) when Table 1. Comparison of plant communities occurring on Fossil Butte National Monument, Lincoln County, Wyoming. Corresponding plant communities Plant community identified by Beetle and Marlow Prominent Life-form (1974) species structure Characteristics Crass-Forb (not listed separately) Gramineae Grasses, forbs Occurs on more exposed areas, especially at higl er elevations (Xeric) Sagebrush-Grass Alkali sagebrush and grass, Mt. big sagebrush and shrub complex, Mt. big sagebrush and grass complex, basin big sagebrush complex, shadscale saltbrush and shrub complex, shadscale salt- brush and alkalibrush complex, serviceberry complex Artemisia tridentata Gramineae Grasses, forbs, shrubs Occurs on drier lowland sites (Xeric) Mt. Mahogany Mt. mahogany and serviceberry complex, Mt. mahogany complex, mixed shrub complex, choke- cherry and antelope bitterbrush complex Cercocarpus movtanus Grasses, forbs, shrubs Occurs at higher elevations on flats and in protected areas on slopes (Xeric) Mixed timber and shrub complex Pintis flexilis Grasses, forbs, shrubs, trees Occurs on pro- tected slopes, frequently on north- facing areas at higher elevations (Xeric-Mesic) Meadow (not listed separately) Aspen-willow Aspen complex Sedges {Carex sp.) Grasses, forbs. Popuhis tremiiloides; Grasses, forbs, Salix sp. shrubs, trees Occurs in areas adjacent to willows and aspen on wet sites (Mesic-Hydric) Occurs on wetter sites, usually on protected slopes, associated with springs and inter- mittent streams (Mesic-Hydric) March 1977 Clark: Wyoming Mammals 93 compared with capture rates for these spe- cies in studies 150 miles to the north (Clark 1975). Capture rates of chipmunks increased from the Sagebrush-Grass (1.7/100 TD) and Aspen- Willow (1.7) communities to Moun- tain Mahogany (2.3) and Pine (3.7) commu- nities. No chipmunks were caught in the two herbaceous communities of Meadow or Grass-Forb. Deer mice were present in all but the two wettest communities, Meadow and Aspen-Willow. They were least abun- dant in the Grass-Forb Community (1.8/100 TD), followed by Mountain Mahogany Community (2.8) and Sage-Grass Commu- nity (4.2), and were most abundant in the Pine Community (9.7). Observations and signs.— Twenty-one oth- er species or their signs were observed. Monument management philosophy and the author's time limitations precluded quan- titative sampling of many larger forms. Several species were distributed pre- dominantly in the Grass-Forb and Sagebrush communities: desert cottontail {Sylvilagus audubonii), white-tailed jackrabbit {Lepiis townsendii), Richardson ground squirrel {Spermophilus richardsonii), Uinta ground squirrel (S. armatus), and northern pocket gopher (Thomomys talpoides). Even though ground squirrels were abundant, none were captured. Three yellow-bellied marmots {Mannota flaviventris) were seen on the up- per slopes. A single white-tailed prairie dog colony (Cynomys leucurus) of about 12 ha existed in Sec. 2, T21N, R118W; one sage- brush vole {Lagurus curtatus) was seen. Six- teen pronghorns {Antilocarpa americana), two badgers (Taxidea taxus), and a striped skimk {Mephitis mephitis) were sighted. No species seemed to be restricted to the Mountain Mahogany Community, although nine mule deer {Odocoileus hemionus) and three elk {Cervus canadensis) were observed there. A moose {Alces alces) sign was evi- dent in this community and two moose were seen in the Willow-Aspen Community. Red squirrels {Tamiasciurus hiidsoniiis), bushy-tailed woodrats {Neotoma cinerea), and porcupines {Erethizon dorsatiim) were associated mostly with the Pine Community. Two species were associated commonly with the Aspen-Willow Community; they were moose and beaver {Castor canadensis). Several Microtus species (i.e., M. pennsylvanicus, M. montanus, and M. long- icaudus) expected in the Meadow Commu- nity were not found, nor were any runways or nests located (cf. Clark 1973). Long (1965) listed specimens of all three species collected within 25 km of the monument, which shows their general occupancy of southwestern Wyoming. A few ubiquitous species were present, including coyotes {Canis latrans), long-tailed weasels {Mustela frenata), and wild horses {Equiis cabalis) (N = 12). Species of unverified presence.— Long (1965) listed 39 other species known to oc- cur in the region. No evidence of their presence was found in this investigation. Domestic livestock.— Fossil Butte has a history of cattle, horse, and sheep grazing. Before the establishment of the monument, the area was managed by the Bureau of Land Management. The area is still utilized by domestic livestock under grazing permits that will expire in 1983; grazing by cattle and sheep has influenced the native plant communities (Beetle and Marlow 1974). Heaviest grazing is in the wetter commu- nities. Meadow and Willow-Aspen, and in the drier communities adjacent to these areas. Current management further concen- trates animals by placing salt blocks in these areas. In addition, sheep are wintered on the monument. Quantitative data on livestock numbers and ranging patterns were not taken in this study. Beetle and Marlow (1974) noted that the combined grazing by both cattle and sheep during the spring rapidly inhibits annual vegetation production. Discussion Currently no data exist on the relative abundance of each species present in mam- malian communities in the area with which to compare the results of this preliminary survey. The heavy, concentrated livestock use of the Meadow and Willow-Aspen com- munities and adjacent areas, which has re- sulted in an alteration of vegetation (close 94 Great Basin Naturalist Vol. 37, No. 1 cropped vegetation, trampling, possible rel- atively high nitrogen content of soil, etc.), probably has in turn affected the distribu- tion and abundance of at least some of the smaller mammals (e.g., Microtus complex). Since the area adjacent to the monument receives similar heavy livestock use, it would be difficult to find a control area to test this hypothesis. However, with the re- moval of livestock in the future, it may be expected that several mammals that are cur- rently rare or non-existent on the mon- ument but that are characteristic of the area will be found in greater numbers (e.g., Sorex cinreus, S. vagrans, S. nanus, and S. paUistris; Microtus pennsylvanicus, M. mon- tanus, M. longicciudus, and Zapus princeps). Acknowledgments I thank Wavne E. Welch, then director of Fossil Butte National Monument, and the field staff for their interest and encour- agement. David Armstrong and Denise Casey critically reviewed the manuscript. The Department of Biology, Idaho State University, aided in typing. Literature Cited Beetle, A. A. and C. B. Marlow. 1974. A vegetative survey of Fossil Butte National Monument, Kemmerer, Wyoming. Univ. Wyoming Coop. Res. Rept. to Yellowstone Envir. Stud. Center. Vol. 11-40. Clark, T. W. 1973. Local distribution and interspecies interactions in microtines. Grand Teton Nation- al Park, Wyoming. Great Basin Nat. 33:205- 217. 1975. Ecological notes on deer mice in Grand Teton National Park, Wyoming. Northwest Sci. 49:12-13. Long, C. A. 1965. Mammals of Wyoming. Univ. Kan- sas Publ., Mus. Nat. Hist. 14:483-758. AN UNDESCRIBED SPECIES OF THELYPODIOPSIS (BRASSICACEAE) FROM THE UINTA BASIN, UTAH Stanley L. Welsh' and N. Duane Atwood' Abstr,\ct.— Thehjpodiopsis argiUacea Welsh and Atwood sp. nov. is described from specimens collected on the Green River Shale Formation in Uintah Connty, Utah. The impact of the Endangered Species Act of 1973 is being felt throughout the federal agencies that manage Utah lands. Intensified field investigations have yielded not only known taxa that are rare and po- tentially threatened or endangered, but also those that have not been previously named or described (i.e., those new to science). One of those new species is the one de- scribed below. The search that yielded the type material was undertaken to rediscover, if possible, living plants of the rare and ob- scure monotypic and endemic genus Glaucocarpum. Although Glaucocarpum was not discov- ered until later in the summer of 1976, sev- eral other plants of interest were collected from the type locality of Glaucocarpum. In- cluded among the collections were speci- mens of a crucifer, which is distinctive among the numerous Utah species. The presence of two narrowly endemic species, Glaucocarpum suffrutescens (Rollins) Rollins and Thehjpodiopsis argiUacea Welsh and At- wood, in the same general area is not alto- gether surprising when one considers the role that the common substrate (Green Riv- er Shale) has played as a spawning ground for narrow endemics. The obscure Astra- galus lutosus Jones, for example, is also known from that formation, as are a num- ber of other species. In appearance and duration, T. .argiUacea is unlike any of the species of Thelypo- diopsis in Utah. Those previously known from the state are all biennials, with well- developed basal leaves; the stems of T. ar- giUacea arise from a branching subligneous caudex. The lowermost leaves are reduced in size; the others are merely sessile and not sagittate or auriculate as in the biennial species. In the diagnosis, T. argiUacea is compared to T. linearifolia, a perennial spe- cies known from Colorado and Arizona to Mexico. T. argiUacea differs from T. line- arifolia in the salient features noted in the diagnosis inter alia. The type material lacks mature siliques of the current season, and measurements of pods are taken from shattered fragments persisting on stems of the previous year. Thelypodiopsis argiUacea Welsh and Atwood, sp. nov. Thelypodiopsis linearifolia (Gray) Al-She- baz aemulans, differt plantis parviores (13- 30, nee 50-150 cm), foliis brevioris, petalis parvioris unguibus laminis nee distinguendis et siliquis et stylis brevioris et lobis stig- matis obtusis. Plantae perennes omnino glabrae glaucae; caules 13-30 cm altus simplices vel ramifi- cantes enascentes caudex sublignei; folia 9- 35 mm longa 0.8-2 mm lata sessilia omnino caulina nee auriculata linearia leviter car- nosa acuta vel rotundata; racemi (5) 8-22- floribus; pedicelli 7-18 mm longi adscen- dentes; sepala 4.2-6.5 mm longa violacea marginibus hyalinis; petala 7.8-10.9 mm longa 2.5-3.2 mm lata alba vel lilacina, venis purpuris conspicuis, ungue lamina nee 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. Bureau of Land Management, Cedar City, Utah 84720. 95 96 Great Basin Naturalist Vol. 37, No. 1 distinguenda; antherae 1.7-2.5 mm longae; siliquae 18-25 mm longae 1-1.2 mm latae subsessiles teretes adscendentes vel erectae; styli 0.5-1 mm longi obconici; stigmata bilobata. Holotype: Utah, Uintah County, hills west of Willow Creek, on the east slope of Big Pack Mountain, TIOS, R20E, Sec. 33, at 5000 feet elevation, on Green River Shale, N. D. Atwood 6627, 11 May 1976 (BRY, isotypes to be distributed). Fig. 1. Tfielypodiopsis argillacea Welsh and Atwood sp. nov.: A. Habit sketch (x 1); B. Flower (x 10); C. Flower (dissected) (x 10). A MOUNTAIN CERCOCARPUS POPULATION-REVISITED' E. S. Nixon' Abstract.— a mountain cercocarpus (Cercocarpus montanus Raf.) population was analyzed first in 1965 and again in 1975 to gain insight into the growth and reproduction of this species. The basis for comparison was plant height and number of basal stems. Growth (measured by height) was slow during the 10-year period, with plants on the northeast-facing slope having an average increase of 37 cm and those on the southwest-facing slope maintaining relatively the same average height. There was an overall decrease in average number of base stems for plants positioned on both slopes. Located about six miles south of Chadron, Nebraska, near its eastern distributional boundary, is a rather small population of mountain cercocarpus {Cercocarpus mon- tanus Raf.). During the summer of 1965, this entire population was analyzed by measuring the height and counting the number of basal stems on each shrub. This analysis was to provide a basis for future comparative studies. The objective of this study, therefore, was to reanalyze the popu- lation 10 years after the initial study to bet- ter understand the growth and reproduction of this shrub. Although mountain cercocarpus provides shelter and browse for big game and is ef- fective in erosion control (Richens 1967, U.S. Department of Agriculture 1974), little research in regard to its ecology had been done prior to 1960. Medin (1960) accom- plished a rather detailed study of this spe- cies in relation to physical site factors and found that soil depth and moisture were the most significant factors affecting mountain cercocarpus production. Additional informa- tion is needed, however, to better under- stand the population dynamics of this spe- cies. The mountain cercocarpus population in northwestern Nebraska was located on a hill that extended into a valley in a southeast to northwest direction. The vegetation of the valley was mainly mixed-grass prairie. The geographic position of the hill resulted in northeast- and southwest-facing slopes that ranged between 25 and 35 degrees. Moun- tain cercocarpus plants occupied both slopes. The northeast-facing slope, which ap- peared to be more mesic, contained a fairly uniform canopy of ponderosa pine {Finns ponderosa Dougl.) and a shrub layer con- sisting chiefly of wax currant {Ribes ceretim Dougl.), prickly rose {Rosa acicularis Lindl.), western snowberry {Symphoricarpus occidentalis Hook.), poison ivy {Rhus radi- cans L.), common chokecherry {Prunus virginiana L.), and skunkbush {Rhus trilo- bata Nutt.). The dry southwest-facing slope exhibited a more open ponderosa pine cano- py and a greater exposure of bare ground. The shrub layer consisted mostly of skunk- bush and Great Plains yucca {Yucca glauca Nutt.). Grasses, sedges, and forbs were vari- ously distributed on both slopes. Methods and Procedures Soils were analyzed at the Soil Testing Laboratory at Stephen F. Austin State Uni- versity, Nacogdoches, Texas. Analyses in- cluded pH, Ca, P, K, Mg, soluble salts, and particle size. Surface samples were taken from the northeast- and southwest-facing slopes and from the ridge top. Using the ridge top as the dividing line, mountain cercocarpus plants were divided into two groups, one associated with the Research was partially supported by a grant from the Stephen F. Austin State University Sigma Xi Club. Sii Nixon, Jane Nixon, and Doris Gates for their aid to the project. ■Department of Biology, Stephen F. Austin State University, Nacogdoches, Texas 75961, appreciation is extended to Suzanne 97 Great Basin Naturalist Vol. 37, No. 1 northeast-facing slope and the other with the southwest-facing slope. Height and number of stems were recorded for each plant. Height measurements were taken from the soil surface to the tip of the tallest stem. Stem counts included those stems originating at the base of the plant. Results Soils Soils of the study site, regardless of slope exposure, were generally similar. Texturally, they ranged from sandy-loam to loam. Nu- trient concentrations, with the exception of P, appeared adequate for sustaining plant life (to 2275 ppm of Ca, to 31 ppm of P, to 300 ppm of K, and to 150 ppm of Mg). Soil pH ranged from 6.5 to 7.5, and soluble salt content was generally low. Height Comparisons Sortheast- facing slope: Interestingly, the number of plants on the northeast-facing slope after 10 years was almost identical (425 plants in 1965 and 426 in 1975), result- ing in a general shift in height of already existing plants (Fig. 1). In 1965 the greatest number of plants was in the 51-75 cm size class, whereas in 1975 the paramount size class was 176-200 cm. This is further sub- stantiated by the occurrence of 98 fewer plants with heights between 1 and 150 cm as compared to 99 additional plants with heights between 150 and >300 cm. The av- erage stem height in 1975 was 154 cm, a noticeable increase over the 1965 average of 117 cm. Southicest-facing dope: Height growth trends were not as discernable on the south- west-facing slope (Fig. 1). There was a shift- ing of plants from the 51-75 cm size class to the 126-150 cm size class, indicating in- creased growth. A shift associated with the 201-225 cm size class peak in 1965 is bare- ly observable, although, with the exception of the >.30O cm size class, plants were gen- erally taller. The average height of moun- tain cercocarpus plants in 1965 was 149 cm; in 1975 it was 144 cm. The number of mountain cercocarpus plants on the southwest-facing slope in- creased from 360 to 397, an increase re- sulting from the establi.shment of a group of seedlings (Fig. 1). There were 36 more plants in the 1-150 cm category in 1975. The number in the 151- > 300 cm range, on the other hand, remained essentially the same (170 in 1965 and 171 in 1975). Stem Comparisons Northeast-facing slope: There was a re- duction in the number of stems per plant during the 10-year period on this slope (an average of 10.9 in 1965 and 8.8 in 1975). Size class information indicated that in 1975 there were an additional 28 plants with 20 stems or less, and 27 fewer plants with more than 20 stems (Fig. 2). In 1965, 83 percent of the plants had less than 20 stems; in 1975, 89 percent had less than 20 stems. The reduction in number of stems for all mountain cercocarpus on this slope was from 4648 (1965) to 3755 (1975). Results were accentuated on the northeast-facing HEIGHT SIZE CLASSES (C* Fi^. 1. Height comparisons of a mountain cerco- :arpus population sampled in 1965 and 1975. March 1977 Nixon: Mountain Cercocarpus 99 slope because it appears that the same plants were present in 1965 and 1975. Southwest-facing slope: The 10-year trend on this slope also indicated a general reduc- tion in number of stems (11.5 stems per plant in 1965 as compared to 8.7 in 1975). Plants with 15 stems or more decreased in number by 32 plants. Those with less than 15 stems increased by 69 plants, but it should be remembered that much of this in- crease is attributed to reproduction. There were at least 37 new seedlings recorded in 1975. The overall reduction in number of stems on this slope was from 4139 to 3471. Discussion Reproduction of mountain cercocarpus during the 10-year period from 1965 to 1975 was essentially non-existent on the northeast-facing slope, but had occurred in one particular location on the southwest- facing slope. If dependent on natural repro- Fig. 2. Stem (number) comparisons of a mountain cercocarpus population sampled in 196.5 and 1975. duction, seedling establishment appears to be very sporadic. This general lack of re- production appears to be characteristic of this species, and seems to be due in part to seedling susceptibility to drought and frost (Plummer et al. 1968). Consequently, one- or two-year-old nursery-grown transplants with proper management are reported to have better chances of survival, and thus have been used in some instances to maintain or extend populations (Springfield 1972). Population dynamics over 10 years showed a general increa.se in height accom- panied by a decrease in number of stems. Because of this reciprocal effect, it's doubt- ful that overall production was increased. The population appeared to be only lightly grazed by deer, so the consequence of graz- ing was minimal. Growth on the north- east-facing slope was more pronounced than that on the southwest-facing slope, due in part to seedling establishment on the south- west-facing slope and the fact that the seed- ling establishment lowered the average height. Medin (1960) found that soil depth, clay content of the A horizon, soil moisture, and surface stoniness (negatively) were site factors that contributed significantly to mountain cercocarpus production. This cor- related very closely with results concerning the Nebraska population. The northeast-fac- ing slope contained deeper soils, had a high- er clay content in the A horizon, appeared more mesic, and was less stony than the southwest-facing slope. Literature Cited Medin, D. E. 1960. Physical site factors influencing an- nual production of true mountain mahogany, Cercocarpus montaniis. Ecology 41:454-460. Plummer, A. P., D. R. Christensen, and S. B. MoNSEN. 1968. Restoring big-game range in Utah. Utah Div. Fish and Game Publ. 68-3. 183 pp. Richens, V. B. 1967. Characteristics of mule deer herds and their range in northeastern Utah. J. Wildlife Mgmt. 31:651-666. Springfield, H. W. 1972. Mulching improves survival and growth of cercocarpus transplants. USDA Forest Service Research Note RM-220. 4 pp. U.S. Department of Agriculture, Forest Service. 1974. Seeds of woody plants in the United States. U. S. Dept. Agric, Agric. Handb. 450. 883 pp. A NEW STATUS FOR PENSTEMON CYANANTHUS HOOK. SSP. SUBGLABER PENNELL (SCROPHULARIACEAE) Stephen L. Clark' Absthact.- Penstemon cyammthus Hook. ssp. subglaber Pennell, a plant found frequently in northwestern Utah, southeastern Idaho, and southeastern Wyoming, is elevated to species status. A new name, Penstemon holm- grenii Clark is proposed because P. subglaber is preoccupied. Penstemon cijananthus Hook. ssp. sub- glaber Pennell is a tall, densely puberulent to pubescent, blue-flowered plant found commonly within sage and pinyon-juniper areas of northwestern Utah, southeastern Idaho, and southwestern Wyoming. Pennell described it in 1920 and .separated it from P. cyananthus ssp. cijananthus on the basis that the leaves of the subspecies are more lanceolate and more densely puberulent. In- vestigations by this writer suggest, however, that other differences exist. Penstemon cya- nanthus ssp. cyananthus is never pube.scent. The staminode of P. cyananthus ssp. sub- ' Department of Botany. Weber State College. Ogden, Utah 84408. glaber is densely covered with long yellow hairs, and the two taxa are not sympatric. These differences suggest to the writer that the subspecies should be elevated to species status. The following new name is proposed to replace the epithet Penstemon cyananthus Hook. ssp. subglaber Pennell: Penstemon Iwlmgrenii Clark nom. nov., based on P. cyananthus Hook. ssp. subglaber Pennell in Scrophulariaceae of the Rocky Mountain States. Contributions from the United States National Herbarium 20(9):313-381. Not P. subglaber Rydb. Named in honor of Arthur H. Holmgren, a friend who first guided me into studies in Penstemon. 100 THE INFLUENCE OF PREDATOR CONTROL ON TWO ADJACENT WINTERING DEER HERDS' Dennis D. Austin,' Philip J. Urness,' and Michael L. Wolfe' Abstract.— Two mule deer herds were studied on comparable, adjacent winter ranges in Utah. Significant dif ferences in overwinter fawn survival were attributed to intensive predator control. An unknown number of deer are contin- ually killed on summer and winter ranges by predators, and in at least some locations these losses are significant. In Oregon, Trainer (1975), using radio-collared mule deer, found 14 percent fawn loss to pre- dation in the first 45 days following birth, and 40 percent loss between January and April. Knowlton (1968) found that predation on fawns significantly limited deer popu- lation growth on the Welder Wildlife Ref- uge. Beasom (1974) showed a three-fold in- crease in the number of surviving fawns in a location of intensive predator control when compared to adjacent ranges in Texas. Smith (1976) compared fawn survival through January by comparing a herd en- closed by a predator-proof fence to an adja- cent free-ranging herd; the enclosed herd had about twice the number of fawns/ 100 does. Conversely, other investigators (Ozoga 1966, Hancock 1974, Gipson 1974, Korsch- gen 1957) have indicated that predators have little influence on deer herds. From 1973 to 1976 mule deer were stud- ied on two adjacent pinyon-juniper winter ranges in northeastern Utah. Returns and observations from a deer tagging program indicated that both herds shared the Blue Mountain Plateau summer range. During late autumn one herd migrated into Miners Draw on the south side of the plateau, and the second herd migrated into the Cub Creek drainage on the west slope. Winter- ing herds were discrete, separated by about 11 km. Winter ranges were comparable in elevation, climate, and major browse spe- cies—big sagebrush {Artemisia tridentata). mountain mahogany {Cercocarpus mon- tanus), and Utah juniper {Juniperus os- teospemia). Browse utilization transects and field observations indicated that throughout the study deer numbers were well below carrying capacity on both ranges. Predator control in the two areas greatly differed. Miners Draw, which received only limited control, was accessible by an unim- proved road, and during the winter deep snow conditions often prevented any travel except via snowmobile. Consequently, this area received little use, and no predator hunters were observed in the area. In comparison, the Cub Creek drainage was accessible via a paved two-lane road, becoming unimproved about halfway through the winter range. Both sections of this road were kept open by snowplows for two-wheel drive vehicles during most of the winter. A ranch was located in the lower portion of the area. Sheep, cattle, and horses were allowed to graze on the drain- age throughout the winter. To minimize livestock losses, predators were intensively controlled by the owners and by a hired professional hunter-trapper. In addition, other predator hunters were occasionally contacted. The USFWS took approximately 80 coyotes in the Cub Creek drainage and 45 in Miners Draw during the winters from 1973 to 1976, and reported predator control work was less extensive in Miners Draw (personal communication. Bob Dickson, USFWS). The number of visual predator observa- tions, deer carcasses located, and deer den- 'This study was partially supported by funds from the Pittman-Robertson Act under Project W105R. 'Department of Range Science, Utah State University, Logan, Utah 84322. "^poartment of Wildlife Science, Utah State University, Logan, Utah 84322. 101 102 Great Basin Naturalist Vol. 37, No. 1 sities between areas also differed. With ap- proximately the same amount of effort expended in observation in both areas, only one covote was observed in the Cub Creek drainage compared to nine in Miners Draw. Fifteen deer carcasses were found in Miners Draw; three of them were directly attri- buted to predation. Most of the others showed evidences of predation, but carcass conditions were too poor for accurate veri- fication. In the Cub Creek drainage only two carcasses were found; one was evi- dently a road kill, the other was likely a poaching case. The mean deer densities for the two winters were six deer/km^ in Min- ers Draw and 19 deer/km^ in the Cub Creek drainage. The winter range in Miners Draw contained about 39.7 ki the Cub Creek drainage contained 29.8 km^. Deer classification counts were made dur- ing post-hunt and post-winter periods with 20x spotting scopes and binoculars. Data used included only those observations where all individuals within the group were posi- tivelv classified (Table 1). Using an adjusted chi-square test, fawn and adult classification counts were not statistically significant be- tween areas for the post-hunt periods 1973- 74. However, counts were significantly dif- ferent for the post-winter periods (1974 X^.l.9 >-80; 1975 X2 J jg >.80; 1976 X^ 1.7.5 >-99) as well as for all years combined (X^ 3.11.3 >-98). Thus a significantly higher proportion of the fawns entering the winter period survived in the Cub Creek drainage, which had more intensive predator control, than did in Miners Draw. Table 1. Fawn : adult ratios during post-hunt and post-winter periods (Fawns : 100 Adults) 1973-74 1974-75 1975-76 Cub Miners Cub Miners Cub Miners Creek Draw Creek Draw Creek Draw Post-hunt 46 41 Post-winter 49 29 74 65 50 31 79 27 Literature Cited Beasom, S. 1974. Relationships between predator re- moval and whitetailed deer net productivity. J. Wildl. Manage. 38(4): 854-859. GiPSON, p. S. 1974. Food habits of Coyotes in Ar- kansas. J. Wildl. Manage. 38(4): 848-853. Hancock, N. V. 1974. Coyote predation in relation to deer. Western Fish and Game Departments special meeting. Sparks, Nevada. February 20- 21. Knowlton, F. F. 1968. Coyote predation as a factor in management of antelope in fenced pastures. Proc. Biennial Antelope Workshop. 3:65-74. KoRSCHGEN, L. J. 1957. Food habits of the coyote in Missouri, J. Wildl. Manage. 21(4):424-435'. Ozoga, J. AND E. M. Harger. 1966. Winter activities and feeding habits of Northern Michigan coy- otes. J. Range Manage. 30(4):809-820. Smith, R. H. 1976. Factors affecting survival of mule deer fawns. Federal Aid Final Report Project W-78-R, Arizona Fish and Game Department. 15 pp. Trainer, C. 1975. Direct causes of mortality in mule deer fawns during summer and winter periods on Steens Mountain, Oregon— A progress report. Western Proc. Fish and Game Commissioners, p. 163-170. PLANTS OF ARIZONA: A NEW SPECIES OF ASTRAGALUS FROM THE KAIBAB PLATEAU Stanley L. Welsh and Kaye H. Thome' Abstract.— Astragalus atwoodii Welsh and Thome is named and described on the basis of materials collected on the Kaibab Plateau in Coconino County, Arizona. In his discussion of Astragalus pinonis Jones, Barneby (1964, Atlas of North Ameri- can Astragalus, Mem. N.Y. Bot. Card. 13:259-261) discussed the existence of two collections tentatively ascribed to that spe- cies. Both collections are from Coconino County, Arizona. The first one cited (Goodd- ing 92-49) is from South Canyon, south of House Rock Valley, and has not been exam- ined by us. The other (Swapp 37) is filed in the collection of BRY. Dr. N. D. Atwood's recent collections in both flower and in fruit have been compared to the Swapp specimen. The specimens are a close match and demonstrate that the differences in ovule number as noted by Barneby (1964) hold constant for the Arizona materials. Further, the pods are consistently more slender than noted for A. pinonis sens. str. The terminal leaflet, always jointed to the rachis in A. pinonis, is occasionally con- fluent in the leaves of specimens from Ari- zona. Thus, the features of the Kaibab plants vary in approximately the same de- gree of magnitude as in other species of the section Lonchocarpi as defined by Barneby (1964). It is proposed, therefore, that the plants from Arizona be designated as a new spe- cies, named to honor Dr. N. D. Atwood, tireless collector and student of the Hydro- phyllaceae. Astragalus atwoodii Welsh and Thorne, sp. no v. Astragalo pinoni Jones aemulans, differt legumine angustiore (2.8-5.0, nee. 5.5-8.5 mm) et seminibus pauciore (18-22, nee 32- 42). Plantae perennes; caules erecti enascentes caudice subterraneis 1.9-5.7 din altus, pilis basifixis adpressis; stipulae omnes distinctae; folia 0.6-9 cm longa; foliola (7) 9-17 (ple- rumque 13-17) linearia ad anguste oblonga obtusa ad rotundata 1.5-12.5 mm longa, 0.7-2.5 mm lata, foliolum terminale saepe confluens; pedunculi 4-7.5 cm longi; racemi 2-15-floribus 2-11 cm longi in fructum; bracteae tringulari-subulatae 1-2.2 mm longae; pedicelli 1-2.5 mm longi; calyces 5- 6.5 mm longi strigulosi, tuba campanulato 3.5-4 mm longo, dentibus triangulari-sub- ulatis 1.1-2.6 mm longis; flores ex purpureo ochroleuci 8-11 mm longi; legumina effusi- descendentes, subsessilia 21-34 mm longa 2.8-5 mm in diametros teretia vel dorsi-ven- trali compressa strigosa rigide chartacei-co- riacea brunnea grosse reticulata; ovula 18- 22. Holotype: Arizona, Coconino County, 17 miles southeast of Fredonia Sawmill, along logging road to Ryan, pinyon-juniper-cow- ania-big sagebrush community, N. D. At- wood 6794, 15 June 1976 (BRY, isotypes to be distributed). Additional specimens: Ari- zona, Coconino County; Kaibab Forest, T39 N, Rl E, Sec. 29, B. Swapp 37-S, 7 June 1926; do, 17 miles southeast of Fredonia Sawmill, N. D. Atwood 5053, 20 May 1973; do, 23 miles southeast of Fredonia Sawmill, N. D. Atwood 5055, 20 May 1973 (all BRY). 'Life Science Museum and Department of Botany and Range Science. BriKham Young University. Provo, Utah 84602. 103 104 Great Basin Naturalist Vol. 37, No. 1 Fig. I. Astragalus atwoodii Welsh and Thome (x 1), drawn from Atwood 6794 and 5055 (BRY). BIONOMICS OF TWO ASPEN BARK BEETLES, TRYPOPHLOEUS POPULI AND PROCRYPHALUS MUCRONATUS (COLEOPTERA: SCOLYTIDAE) Jerold L. Petty' Abstract.— Beetles of Tnjpophloeus popuH attacked green bark of unhealthy aspen (Populus tremuloides) and hastened the death of the tree. Beetles of Pwcryphahis mitcromitus favored dead bark and were of little signifi- cance in the death of the aspen. These two beetle species were distinguished from each other by their primary galleries, eggs, larvae, larval mines, and mating behavior. In T. populi, there were three larval instars and one to one and one-half generations per year; only larvae overwintered, and the eggs of the first generation were present by mid-July. In P. mucronatus, there were two larval instars and one and one-half to two generations per year; both larvae and adults overwintered, and the first eggs of the new season were present by late May. In North x\merica, quaking aspen {Popu- lus tremuloides Michaux) occurs coast to coast from the 49th to the 69th parallel and sporadically at the higher elevations in the southwestern United States and in Mexico (Little 1971). These trees average from 12 to 15 m in height and from 30 to 60 cm in diameter (Preston 1969). Aspen are important in several ways. Their role is well known as an intermediate (subclimax) cover species in plant succes- sion, leading to the climax coniferous com- munity (Oosting 1956, Krebill 1972). They recover quickly after extreme disturbances such as fire and clearcutting (Preston 1969). Directly, they provide browse for deer, elk, and livestock (Reynolds 1969, Wallmo 1972). Indirectly, they provide for a rich understory that in turn provides habitat for a diverse fauna and forage for livestock (Harper and Warner 1972). In the Rocky Mountains, the poplars (especially aspen) are the most important broadleaf trees and the only ones that could be considered us- able as timber (Davidson and Hinds 1968). Aspen can produce more wood over a shorter period of time than conifers at com- parable elevations (Davidson, Hinds, and Hawksworth 1959). Recently, they have be- come important as a timber resource (Lamb 1967). Aesthetically, anyone who appre- ciates nature cannot deny the unequaled beauty of a mountainside of aspen, espe- cially in their cadmium hues of autumn. The only American Scolytidae known to infest the bark of quaking aspen are Pro- cryphalus mucronatus (LeConte), Tnj- pophloeus populi Hopkins, and T. thatcheri Wood. Another scolytid, Trypodendron retu- sum (LeConte), attacks aspen, but it is a wood-boring ambrosia beetle. The published research on P. mucronatus, T. populi, and T. thatcheri has been mainly taxonomic. In fact, virtually nothing of a nontaxonomic nature has been written on the genera Pro- cryphalus and Trypophloeus. Exceptions are a one-page article on the habits of the Eu- ropean Trypophloeus binodulus (Hagedorn 1904) and a half-page article on four Eu- ropean and Asian Trypophloeus spp. (Palm 1959). There are only four known species of Pro- cryphalus in the world, two in northern North America and two in northeastern Asia. In North America, P. utahensis Hop- kins breeds in Salix scouleriana (Scouler Willow) and P. mucronatus in Populus tremuloides. There are 15 recorded localities for P. mucronatus, all of which are in the United States: Colorado 7, Idaho 2, New Mexico 2, and Utah 4 (Wood in press). About a dozen species of Trypophloeus occur in Europe and Asia, but only four in North America. Of these four, T. populi and 'Department of Zoology. Brinhain Young University, Provo. Utah. Cnrrent address; 576 Eastland North. Twin Falls. Idaho 83301. 105 106 Great Basin Naturalist Vol. 37, No. 1 /'. thatrlwri are the only ones that have been collected from P. tremuloides. Try- pophloeus papitU also breeds in P. acuminata iLanceleaf Cottonwood), P. angustifoUa (Narrow-leaf Cottonwood), and P. tricho- carpa (Black Cottonwood). There are 13 re- corded localities for T. popiili: Arizona 1, Colorado 1, Nevada (eastern) 1, Utah 6, Idaho 1, Manitoba 1, New Brunswick 1, and Saskatchewan 1 (Wood in press). Tlie objectives of this paper were: (1) to report investigations on the life history and habits of T. popuH and P. mucronatus, and (2) to determine the nature of the damage thev cause in the host. Emphasis was placed on the differences between the two beetles in habits and behavior. It is hoped this re- search will establish a better understanding of the ecological significance of these spe- cies and provide a foundation for further studies of these and related scolytid species. Materials and Methods Four sites were selected for this study, all of which were in Utah County, Utah. More tlian 80 percent of the total number of trees on each site were Populus tremuloides. The sites covered approximately 3,000 m- each and ranged from 2,256 to 2,377 m in eleva- tion. The sites were initially selected by lo- cating scolytid-infested aspen. The periods of study for each site and further details on the elevation, number of trees studied, and exact location of sites are given in Table 1. Throughout the study, periodic collec- tions and observations were made at the field research sites and in the laboratory at Brigham Young University from material brought from the sites. To facilitate obser- vations, scolytid galleries were exposed by cutting away the bark with a razor-blade scalpel. To rear the beetles, infested bolts, limbs, and bark sections were placed in 20-gallon, galvanized steel garbage cans, each of which had a pint glass bottle attached to collect the emerging scolytids. All rearing was conducted in the laboratory at room temperature. The number of larval instars was determined by measuring the width of the head capsules (Lekander 1968). Head capsules were measured from undamaged larvae and with an occular micrometer scale in a microscope at 80X magnification. The drawings of the larval head capsules (Figs. 22, 23) were composed from viewing 25 specimens of each beetle species under the light microscope and from 18 scanning electron micrographs of four specimens of each species. Photographs were taken with a Yaschica 35 mm SLR camera equipped with extension tubes and a flash. The mar- gin of a millimeter ruler was included in many of the photographs for measurements. Details of the territorial behavior and mat- ing ritual were obtained from extensive ob- servations in the field and from movies ob- tained in the field with a Vivitar 8 mm movie camera equipped with a close-up lens. A fungus was cultured from adult bark beetles by macerating them on the top of sterile pieces of aspen wood (four beetles per piece), and then placing each piece of wood upright on agar-agar medium inside petri dishes. The petri dishes were main- tained at room temperature in the labora- Table 1. Descriptions of research sites, Utah County, Utah. Site LotaUtv No. Elevation (m) Pm" Tp° DBH Study period (cm) I 1.6 km N Aspen Grove 2,256-2,259 II 1.7 km N .\spen Grove 2,274-2,286 ill 2.4 km \ Aspen Grove 2,365-2,377 l\ 1.9 km \E Squaw Peak 2,.32.3-2..332 8 2 2 30-50 VII-1972 to VI-1976 14 18 8 10-48 VII- 1972 to VI-1976 0 12 2 31-49 VII- 1975 to VI-1976 8 0 0 .30-40 V-1973 to XI- 1974 •Th4##v^.,», nf- ,'- ^^' V T / l"ig. 1. Tiijpophluius pi)puli, piiiiiaiy ^allciic-s: 10 of the 13 galleries pictured were exposed by cutting away the epidermal bark covering them: e, white pointer marks the gallery entrance; black pointer, the parental adult exit hole; ft, the food tunnel; the cir- cled arrow points upward. 108 Great Basin Naturalist Vol. 37, No. 1 Table 2. Details of 40 primary galleries from 4 trees (10 galleries each) 9 March 1976 (Trypophloeus populi). No. of Listing Category Galleries Bark surface at entrance Slightly rough or ir- regular (Fig. 20) .31 Black rough spot, en- trance at edge (Fig. 4) 5 Black rough spot 4 Smooth 0 Total 40 Blockage of entrance Partially filled w/frass 18 No blockage 18 Plugged w/dead male 4 Total 40 .\ngle of entrance above or below hori- zontal (degrees) 1-45 above 23 46-90 above 9 0-45 below 9 46-90 below 0 Total 4P Bark covering gallery Medium split (Fig. 6) 20 Slight split (Fig. 1, unexposed galleries) 16 Extreme split (Fig. 20) 4 Total 40 Parents (dead) in the 14 galleries w/sec- ondary chambers.'' Male and female'' 7 Female onM 4 Male only ' 0 -Neither parent 3 Total 14 Parents (dead) in the 26 galleries without secondary chambers.'" Unoccupied 17 Male and female 5 Male only 2 Female only 2 Total 26 ^DIle gallery had two entrance holes. "Exit holes were not present in any of the setondary chambers, and the epidermal bark covering them was not split. •In all seven galleries, the female was blocking the food Hmnel en- trance to the sec-ondary chamber. ■"In all four galleries, the female was blocking the fcxxi tunnel entrance to the secondary- chamber. There were 26 eiit holes (18 in the food tunnel and 12 in the primary chamber!. or she was continuing her excavations. This interrupted the male's attempts to copulate with her. Because of the angle of the en- trance into the bark, it was necessary for the female to back out of the entrance at least one- fourth of the way to be available for copulation. -^fter copulation, the male entered the gallery behind the female. If the gallery was not deep enough to accommodate both, the male entered as far as possible while the female continued to lengthen the tun- nel. As long as the male was protruding above the surface of the bark, another male could challenge him for his claim. This territorial competition was similar to that between fe- males (see Territorial Behavior). The female devoted her time to the exca- vation of the gallery, to oviposition, and probably to repeated copulation inside the gallery (observed on two occasions). She ex- cavated the cavelike primary egg chamber at the inner end of the entrance tunnel. The design of this chamber was varied (Figs. 1, 2, 3) and sometimes consisted of two or even three distinct areas for the deposition of eggs. The female deposited a cluster of approximately 14 eggs in the primary egg chamber. They were deposited near to or against the paper-thin epidermal bark cov- ering the gallery. Here, heat from the sun may have facilitated incubation, and aera- tion through the bark surface may have in- hibited or prevented bacterial or fungal growth. Sometimes two or three clusters of eggs were laid in one gallery. Additional clusters contained fewer eggs (see Eggs). Very often a lone egg or two were depos- ited elsewhere in the gallery (Fig. 3). If eggs were located below subsequent excava- tions, fra.ss accumulated on them (Fig. 2). From the primary chamber, the female u.sually excavated a food tunnel (Figs. 1, 5, 6, f.t.) that was usually directed upward and extended approximately 10 mm. Fra.ss col- lected in the tunnel behind her. Sometimes, she excavated a side pocket and deposited a few eggs therein. At the end of the food tunnel, .she either died, exited by boring out of the tree, or excavated a secondary egg chamber (Figs. 5, 6, s.c). March 1977 Petty: Aspen Bark Beetles Fig. 2. TrypopJihntis popuh, part q)(ist'(i iifw primary t;allfr\': pc, primary chamber. Fit;. 3. TrijpopliUnus popuh. rxp^scd ncu pniiiary gallery; pc, primary chamber; circled arrow points up- ward; parents (male blocking entrance), one and one- half egg clusters, and one lone egg. During the period of excavation and ovi- position, the male blocked the entrance to the gallery with his body (Figs. 2, 3). He died in this position, abandoned the gallery (bv way of the entrance or by boring out through the bark covering the primary chamber), or followed behind the female as she excavated the food tunnel. Either one or both died or exited at or near the end of the food tunnel (Table 2). When the female excavated a secondary egg chamber, the male (if present) usually blocked the food tunnel at the point where it entered the secondary egg chamber (Fig. 5, s.e.) and helped remove boring dust by packing it behind himself in the food tunnel. Once an egg hatched, the larva began its larval mine by boring into the wall of the primary gallery. As the mine in previously unexcavated bark was lengthened, it was also widened according to the growth of the larva. The dark brown to black frass de- posited by the larva accumulated behind it and filled the mine. Here, it stuck together and dried, forming a firm blockage. The 110 Great Basin Naturalist Vol. 37, No. 1 walls of the frass-filled mines were stained a cLuk brown color (Fig. 7). The larva exca- vated the first portion of its mine just under the hark epidermis (Figs. 6, 7). The length of this part of the gallery was approx- imately 20 mm, but this varied depending on the condition of the host, time of year, density of galleries, etc. This portion fol- lowed either a straight course, meandered, or reversed direction abruptly (Figs. 6, 7). At the end of this shallow excavation, the larva mined deeper into the bark (Fig. 6). When the larva was mature, it excavated a frass-ftee pupal cell. In this cell it pu- pated and transformed into an adult. The new adult bored out of the bark and flew to find another tree to repeat the life cycle. Primary gallery.— Forty primary gal- leries, which had been excavated by bark beetles (T. populi) the previous season, were studied 9 March 1976 (Tables 2 and 3). Fourteen of the 40 had a secondary cham- ber. The bark lining the secondary cham- l)ers was darker in color (apparently stained) than in the primary chambers (Figs. 5, 6). In all 14, the bark covering the pri- marv egg chamber and food tunnel leading to the secondary chamber was split, but this split did not extend into the secondary egg chamber. Normally, each gallery was shared by one parental male and female. Occasionally one Table 3. Measurements and details of 40 primary galleries from 4 trees (10 galleries each) 9 March 1976 {Trypophloeus populi). Category 5 m ± SD Range n Entire length 21±7.2(10-14)mm 40 Maximum depth 2±.86(l-5)mm 40 Maximum width 4.6±1.2(3-8)mm 40 Length of primary egg chamber 10.6 ± 2.8(7-20)mm 40 Length of food timnel 10 ± 4.2(3-20)mm 20 Gallery orienta- tion expressed in degrees to the vertical 72 ± 25.9(0-90)° 40 Percent of gal- lery filled with frass 60.5 ± 25.9(10-90)% 40 Percent of food tunnel filled with frass 67 ± 30.7(0.90)% 20 4-6. Trypophloeus populi. details of three primary galleries entrance (e, white pointer), parental adult exit link- lilack pointer*, primary chamber (pc), secondary chamber (sc), secondary entrance (se), food tunnel (ft), the circled arrow points upward; 4, split bark covering primary gallery; 5, bark covering primary gallery removed; 6, liark covering primary gallery and larval mines removed. March 1977 Petty: Aspen Bark Beetles 111 gallery was connected to a neighboring one. Some galleries contained from one to three entrances. Excess parental adults appeared to be permanent residents of the gallery. The frequency of these exceptions was high- ly variable and increased with increased density of the beetles. Of the 10 exposed galleries in Fig. 7, one had two entrances, one contained a lone female, and one con- tained three females. The angle of the entrance into the bark enabled the male to effectively block the entrance. In his blocking position the poste- rior surface of his elytra completely covered the entrance opening (Fig. 3, e.). If the en- trance had been more nearly perpendicular to the bark surface, as in P. tnucronatus, the ventral side of the posterior end of his ab- domen would have been visible from the bark surface, and his elytra would not have completely covered the opening. This would have been to the advantage of pre- dators, and rain could have entered the gal- lery more easily. The angle of the entrance into the bark, and the fact that it was usu- ally directed upward from the horizontal, facilitated removal of frass and boring dust. In P. mucronatus, the entrance was often plugged with frass instead of the male. The food tunnels studied were approx- imately 10 mm long. The reasons for the relatively great length of the food tunnel were not clearly understood. The fact that this tunnel was practically full of boring dust indicated that much more bark was ex- cavated than was ingested by the female parent. Much of the boring dust found in the old primary galleries may have come from the excavation of the first part of this timnel. The food tunnel may hasten the death of the bark in the surrounding area to the advantage of the larvae. The rate of mortality of the immatures was high in bark that remained alive. The greater the dis- tance between a secondary egg chamber and the primary chamber, the less chance there will be of competition for space among the larvae of the gallery. The fact that in some instances the fe- male excavated a secondary egg chamber Fig. 7. Trypophloetis populi, exposed larval mines: Im, larval mines; 1, larva; db, expansion of dying bark; pa, parental adult; E, edge of epidermal bark not cut away; G, primary gallery; white pointer (e), entrance to pri- mary gallery; black pointer, parental adult exit hole; the circled arrow points upward. 112 Great Basin Naturalist Vol. 37, No. 1 and oviposited therein and in other in- stances she exited, indicated that if she did exit she was capable of excavating another priinarv gallery and ovipositing there also. \\'hether or not the female exits at the end of the food tunnel may be a function of the condition of the bark or the proximity of other galleries. If gallery density is high, it is expected that the incidence of secondary chambers would be low. The male's habit of following the female while in the food tunnel suggests that he mav copulate again with the female. His position at the food tunnel entrance may also prevent predators from entering. EcGS.— The eggs were oblong, white, translucent, and had a tacky surface (Figs. 2, 3). Of 52 eggs measured, the mean width and length were .33 ± .030 x .65 ±.097 mm (Range .27-.42 x .55-. 77 mm). To estimate the number of eggs laid per gallery, the eggs and larvae in five primary galleries found 14 July 1973 were counted. The quantities per gallery were: 18 eggs and 3 larvae, 6 and 4, 11 and 6, 22 and 2, and 11 and 4. The parental adults were present and oviposition was probably not completed. .\ primary gallery initiated 13 July 1974 and examined 3 August 1974 (21 days later) contained 15 eggs and 4 larvae (11 eggs and 3 larvae in one cluster, and 4 eggs and one larva in another). The gallery was 1.5-3.5 x 13 mm. Both parents were present and alive, the male blocking the entrance. The tacky surface of the eggs normally held them together in a cluster in one area of the gallery. When frass fell on them, it adhered loosely. Apparently the parents did not remove this debris from the eggs be- cause the eggs might be removed also. Apparently the primary gallery protected the brood from predators and desiccation long enough for the eggs to hatch and for the larvae to mine out of it. After this, the bark covering the gallery usually split open (Fig. 4). The drying and shrinking of the gallery that followed appeared to make the entrances of the larval mines less accessible to entry by predators. Predators may gain entry into the pri- mary gallery when the male leaves the en- trance to feed or copulate inside the gal- lery, and when the bark covering the gallery splits. The male may not need to leave the entrance to feed. Some of the bor- ings pushed to him by the female might serve this purpose. There are some problems created by lay- ing the eggs in clusters. Egg mortality fac- tors such as desiccation, predators, bacteria, and fungi would likely destroy whole clus- ters of eggs at a time compared to one egg at a time if they were oviposited as in P. miicronatiis. Larvae.— The larvae were white, legless, and grublike (Figs. 2, 3). They appeared dark when their intestines were full of the brown bark. The relative length, distribu- tion, and number of head capsule setae (Fig. 22) were characteristic of the species. The pattern of sclerotization on the frontal shield (Fig. 22, f.s.) may have been con- sistent in the last instar. Widths of the head capsules of 232 larvae (Table 4) and numbers per size class (Fig. 24) indicate that there are three larval in- stars. Those .21-. 28 mm, .30-. 38 mm, and .40-. 50 mm were categorized as Instar I, II, and III, respectively. These categorizations resulted in four size classes for Instar I, five for Instar II, and six for Instar III (Fig. 24). In 24 scolytid species, Lekander (1968) found the average increase in larval head capsule widths from one instar to the next was 1.32 times (limits 1.20 and 1.48 times). By applying his method to T. populi, the average head capsule width increase (index) from Instar I to II and II to III was 1.31 and 1.35 times, respectively, compared to 1.33 from Instar I to II in P. mucronatus. The width of the head capsule in the last larval instar of T. populi was considerably wider (.46 ±.022 mm) than that of P. mucronatus (.36 ±.021 mm). Larval mines.— The larval mines were distinct and clearly separate from the pri- mary gallery and usually from each other. The expansion of the dying bark surround- ing the larval mines (Fig. 7, d.b.) indicated that the excavation accelerated the decline of the tree. The dark stain observed lining the mines may have been from oxidation of the bark or frass or both, or it may have March 1977 Petty: Aspen Bark Beetles 113 been from fungi or bacteria associated with the beetle. A dead tree in which emergence had taken place was examined 9 March 1976. The length of 16 larval mines (randomly se- lected) was from 26 to 42 mm (33 ±4.5 mm). For additional information, larval depths and distances away from the primary gallery were measured from four trees con- taining live larvae 9 March 1976 (Table 5). Adults.— Beetles were reared in the labo- ratory on various dates to determine the sex ratio and body lengths of the male and fe- male (Table 6). The total reared (438 males and 587 females) gave a sex ratio of 1:1.34. A ratio of 1:1 was expected because of the importance of the parental male in blocking the gallery entrance. From the 100 adults reared 7-13 April 1976 (Table 6), the first 50 to emerge (7-8 April) were longer (1.86 ± .106 mm, males, and 2.0 ±.097 mm, females) than the second 50 (1.8 ±.124 mm, males, and 1.9 ±.130 mm, females). The same was found in P. mucronatus. In T. popidi, the males were about 5 per- cent shorter than the females (widths were not determined). This was not expected be- cause a male that is smaller than a female cannot block the gallery entrance as effec- tively as one the same size. These measure- ments were taken from emergent adults. The measurements of paired beetles in the primary gallery at the new tree may reveal less difference because of mortality of the smaller beetle during the flight to the new tree, and/or because of the replacement of Table 4. Width of 232 larval head capsules (Try- pophloeus popiili). Ill ± SD Range (mm) Index Obtained 131 .26±.016(.21-.28) 7 .46/.26=1.71 50 .43±.016(.30-.38) 8 .467.34 = 1.35 51 .46±.022(.40-.50) 6 .347.26 = 1.31 "Sample size of larvae. ° "Sample size of trees. the smaller beetle by a larger one in territo- rial behavior. Overwintering.— The larvae constituted the overwintering life stage (Tables 7, 8). Instars II and III survived at a higher rate than did Instar I. The contrast in the total number of each instar from one collection date to the next during winter was appar- ently due to the condition of the bark sam- pled and the differences in the dates when the trees were attacked. The differences between the time mate- rial was brought into the laboratory for rearing and when the new adults began to emerge (Table 6) and the percentages of feeding larvae on different dates (Table 7), indicated that a dormant period occurred in the overwintering larvae and that it was controlled by something other than daylight and continuous warm temperature, because the rearing was conducted at room temper- ature and in the dark. Generations per year.— The presence (Table 8), relative abundance of over- wintering life stages (Table 7), approximate Table 5. Depth of larvae in bark and proximity to primary gallery 9 March 1976 (Trypophloeus populi). Instar Distance from primary gallery m±SD Range (mm) Depth from sur- face of bark m±SD Range (mm) No. of trees sampled and larvae7tree 7.5± 4.85(1-18) not determined 6.6 ±2.3(1-22) .75±.75(.25-4) 1.25±1.01(.24-.4) not determined 40 2(20) 4(20) 3(?) Total "Sample size of larvae. 114 Great Basin Naturalist Vol. 37, No. 1 11-nionth duration of the life cycle in one tree in the field, and approximate 6-month difference between collection date (15 Au- gust 1972) and emergence date (February 1973) under laboratory conditions (Table 6), indicated that there were one to one and one-half generations per year in the field. The overwintered Instar 11 larvae began to pupate and transform in late June. By earlv July thev were emerging and attack- ing new trees, and by mid-July they were ovipositing in the new primary galleries (Table 8). This was a late start compared to P. mucronatiis for the first eggs of the sea- son. Territorial behavior.— The territorial behavior observed was basically similar to that described for P. mucronatiis. However, the attempt of one beetle to dislodge the other by extension of the prothoracic legs (Fig. 26) was not observed. The male that can do the best job of blocking the gallery is apparently selected for both strength and sufficient size to com- pletely plug the entrance. A male that is smaller than the female that excavated the gallery entrance might be more easily dis- placed than one the same size. Locating infested trees.— Infestations were found in stands of aspen where there Table 6. Number and length of males and females reared on various dates, 1976 (Trypophloeiis populi). Collec- Emergence S ? Lengths S (mm) Lengths ? (mm) tion date dates (n) (n) m ± SD Range m ± SD Range 8Mar» 7-8 Apr 18 32 1.9±. 106(1.7-2.1) 2.0±. 097(1.7-2.2) 9-13 Apr 22 28 1.8±. 124(1.5-1.9) 1.9±. 130(1.6-2.1) 14-18 Apr 10 13 Not determined Not determined Totals 50 73 15 Aug'' Feb" 59 41 1.9±. 153(1.5-2.2) 2.0±. 142(1.7-2.3) 2.3 Apr & Mav& 24 May^ Jun 329 471 Not determined Not determined Grand total 438 587 •Collected 15 August 1972 and emerged during Februa •The most advanced life stage present was Instar III. The most advanced life stage present was Instar II. 'The most advanced life stage present was Instar II(?). itial attack of this tree was estimated to be six Table 7. Relative abundance of overwintering life stages of Trypophloeus populi. No. of trees sampled Eggs No. of immatures Instars Pupae No. of adults Collec- Trans- formed" Paren- tal °° Feeding larvae (%)••" I II III 1 1 Nov 75 51 195'' 39 8 0 0 5'- <10(?) 9 Mar 76 0'' .34 194 122 0 0 0" 0 23 Apr 76 0 3 42 155 0 0 0 89 24 .VI a v 76 0 0 13 67 0 0 0 (?) 25 Jun 76 0 0 9 139 48 4 0 (?) •|n pupal cell or brood tree. "In pnmary gallery. •••Percent of larvae thai appeared dark because of bark in their intestines. All the •Of lhe« larvae. 55 were dead. Of these 5.5, .35 were in the primary gallery. Though 6 eggs were collected, they were probably nonviable. Of 54 parental adults collected. 49 were dead. ■"Of 60 parental adults collected, all were dead. were white— their gut devoid of food March 1977 Petty: Aspen Bark Beetles 115 were many trees in the 30 to 50 cm DBH size. Unhealthy or diseased trees, especially those bordering washes, trails, roadways, and seepage areas, appeared to be particu- larly vulnerable to attack. In summer, the leaves of infested trees were usually stunted and lighter in color than normal. Susceptibi- hty to attack seemed correlated with pres- ence of the artist conk Fomes appkincittis (Pers. ex Wallr.) Gill. Trees from which brood had already emerged typically had split bark above the primary galleries and exit holes where the transformed adults had emerged (Fig. 20). Host damage.— The beetles attacked liv- ing bark of unhealthy trees (Figs. 8, 9). At site II, 5 of the 18 (27.8 percent) infested trees studied harbored artist conks (Fomes applanatus). Typically, a single tree was at- tacked en masse, and the tree was covered with the galleries from the lower bole to the branches. Immature beetles died at a high rate in galleries that were in the small- er branches and in bark that remained alive after attack. The dving bark turned some- what orange immediately around each gal- lery (Fig. 7, d.b.); the orange areas enlarged until they coalesced. The leaves died, the bark turned brown, and, within 3 to 10 weeks after the initial attack, the tree was virtually dead. Three healthy trees were inoculated June 1973 with a fungus cultured from adult bee- tles. By July 1974 one of the trees had died; the other two remained healthy. The decline of infested trees, expansion of dying bark tissue around individual gal- leries, and the death of one of the three trees inoculated with fungus cultured from adults indicate that this beetle has a definite Table 8. Number of trees bv calendar date containing the various life stages of Tnjpoplilociis poptili and Pro- :ryph(ihis mucronatus. Life stage Month Mar Apr May Jun Jul Aug Sep Oct °A few egf;.s were found, but the 'This hfe stage was expected. Dec-Feb T. poptili Eggs Larval Instars 0° 0° 0 0 2 5 2 1 3 0° I 4 1 0 0 4 1 2 1 3 o*- II 4 2 4 3 0 3 0" 0- 6 1 III 4 2 5 5 2 0 O'- 1 2 (r Pupae Adults in 0 0 0 0 0 0 5 3 1 1 1 3 0^ 0 0 0 0 0 0 pupal cells Adults starting 0 0 ^ 0 0 5 9 0^ 0" 0 0 primary gallery Adults in 0 0 0 0 2 3 2 1 4 0 primary gallery P. mucronatus Eggs Larval Instars 0° 0 2 '' " 4 (r 0 0 0 I 2 4 2 3 5 1 O" (r 2 o-- II 2 4 3 0 4 5 ()<" o*- 4 1 Pupae Adults in 0 2 0 2 I 1 0 3 8 6 6 0 0 4 0 1 pupal cells Adults starting 0 2 8 4 3 6 O'- o'- 0 0 primary gallery Adults in 1 4 5 6 6 2 er 0" 0'- o*- primary gallery probably nonviable. 16 Great Basin Naturalist Vol. 37, No. 1 role in hastening the dechne and death of unthrifty trees. A pathogenic fungus may be associated with the lieetle. The higli niortaUty of iniinatures in the galleries that were located in hark that re- mained green after attack indicates that the rapid decline of the tree is essential for sur- vival of the brood. Unhealthy trees that might otherwise live on for several years evidently die within weeks if attacked en masse by the beetle. This appears to be beneficial to the survival of the aspen stand, since unhealthy trees iiarbor disease spores and rob understory and new aspen saplings of sunlight and moisture. Procn/pluilus nuicronatus Behavioral sequence.— The behavioral sequence was basically the same as de- scribed for T. populi. The observed differen- ces and other details are mentioned here and under .subsequent sections. Adults walking on the bark of the host held their antennae at right angles to the side of the head. In T. populi the antennae were extended forward at an angle of about 45 degrees from the side of the head. After the male joined the female in the tunnel, the entrance was often blocked with a firm plug of frass. Some of the plugs ob- .served in summer did not completely cover the entrance. Plugged galleries were found that contained only the female parent. Ap- parently, once the entrance was plugged be- hind an unpaired female, a male did not join her. Of 37 galleries studied 1 Novem- ber 1975, the entrances of 35 were plugged completely. Of 90 studied 9 March 1976, all were implugged. On a few occasions, a dead male was found in the entrance. Both the male and female excavated the primary gallery. The female excavated the part of the tunnel where the eggs were de- posited and the male apparently enlarged the gallery near the entrance (Figs. 16, 17). The female deposited each egg in a niche that she excavated and then packed it in with boring dust. The male apparently fed. Figs. 8, 9. A.spen tree recently attacked by Trypophloetis populi: spots); 9. leaf condition (green and dying). 8, distribution of gallery entrances (dark March 1977 Petty: Aspen Bark Beetles 117 and remained available for repeated cop- ulation. After the eggs were laid, the parents died in the gallery. There was one exception to this. Parent exit holes were found in about 30 percent of the new galleries of two trees 25 June 1976. The trees were quite green. This may have been the reason that some of the parents abandoned the galleries. After the egg hatched, the larva began to feed, possibly first on the boring dust that surrounded the egg and then on the walls of the gallery. There appeared to be no rela- tionship between the position of the egg and the direction in which the larva fed. In- star I and II larvae were commonly col- lected in the primary galleries that they had enlarged (Fig. 18). Instar II larvae and pupae were collected at the end of short larval tunnels (Fig. 19). Unlike T. populi. Figs. 10-13. Procnjphalus mucronatus, mating behavior: 10, male mating with female who is searching for a site to begin the primary gallery; 11, female starting the primary gallery; 12, male copulating with female who is starting the primary gallery; 13, male copulating with female who is in the gallery entrance. Figs. 14, 15. Trypo phloem populi, mating behavior: male copulating with female who is starting the primary gallery. 118 Great Basin Naturalist Vol. 37, No. tliese tunnels were only partially filled with low the bark surface, where it pupated (Fig. frass and the walls were not stained black. 19) and transformed. After transformation, it \fter the larva was fully grown, it exca- fed, emerged, and attacked a new tree or vated and cleared a cell about .25 mm be- reattacked the brood tree to repeat the life Figs. 16, 17. PTOtriii '!i / ;/, n>natus. exposed primary galleries: eggs (white arrows), and parents. Figs. 18, 19. Procnjphnlus miicwnatui>, larval excavations and pupae; pa, parental adult; 1, larvae (Instar II); c, pupal cell; p, pupa; ex, exit hole of transformed adult; GI, primary gallery that has been enlarged by larvae; circled arrow points upward. March 1977 Petty: Aspen Bark Beetles 119 cycle. If it was late in the season when the pupa transformed, it overwintered in the pupal cell. Primary gallery.— The primary gallery was: (1) approximately 14 mm long with tunnels from 1 to 3 mm wide that were straight or crooked and branched or un- branched (Figs. 16, 17), (2) usually directed upward (5 to 50 degrees from the horizon- tal) from the entrance, and (3) kept relative- ly free from frass. Ten galleries on 23 April 1976 were filled to approximately 35 per- cent with this debris. Other galleries exam- ined in the summer (Figs. 16, 17) were free of frass. The question arises, how was frass removed when the gallery was plugged? It appeared that bacteria played an important role in reducing the size of the frass par- ticles. The following information was obtained from 25 galleries located July 1973: All contained eggs and parental adults, but no larvae. Galleries having their central axis approximately horizontal outnumbered two to one those that were approximately verti- cal (Fig. 17). Four entrances were located at the side, six at the top, and 15 at the bottom of the galleries. Nine of the galleries were fully branched (Fig. 16), 10 slightly branched (Fig. 17), 4 unb ranched, and 2 had just been initiated. Major differences in the primary gallery compared to T. populi were: (1) the main gallery was composed of comparatively nar- row tunnels instead of broad chambers, and it was usually excavated in dead rather than live bark; (2) the bark of the bole or larger limbs, not the branches, was attacked; (3) the bark was usually soft and fermenting in- stead of firm; (4) the entrance tunnel was more nearly perpendicular to the bark sur- face, penetrated deeper into the bark (Fig. 27), and was often plugged with frass in- stead of being blocked by the male; and (5) the bark above the gallery did not usually split open (Fig. 21) as in T. populi (Fig. 20)'. The excavation of the primary chamber and food tunnel in T. populi resulted in the splitting of the epidermal bark covering them, whereas the excavation of the second- ary chamber in T. populi and the primary gallery in P. mucronatus did not cause the bark to split. The excavations that resulted in splitting were in dying bark that was green, whereas the excavations that did not result in splitting were in dead bark. The angle at which the entrance tunnel penetrated the bark (Fig. 27) usually al- lowed copulation to take place when the male was on the bark surface (Fig. 13) without requiring that the female back out of the entrance. Compared to T. populi, the female was more protected in this position, and the male was closer to the bark surface; therefore, presumably both were less con- spicuous to predators. The angle of the en- trance into the bark may have also facil- itated its being plugged because of the slight elbow created where the entrance tunnel curved to become parallel to the sur- face of the bark. The angle of the entrance, plugging of the entrance of the gallery, and gallery design need further study in relation to the condition of the bark and the time of year when the gallery is excavated. Eggs.— The eggs were oval, white, trans- lucent, and had a tacky surface (Fig. 16). Of 20 eggs measured, the widths and lengths were .38-.44 mm (.41 ±.016 mm) x .62-.76 mm (.68 ±.032 mm). Eggs were counted from 20 primary gal- leries 25 June 1976. Because oviposition was apparently not completed in some of these galleries, only the 10 with the greatest num- ber of eggs were used to compute the mean number of eggs laid per gallery. Galleries with more than one adult female were not counted. The mean was 16.8 eggs (Range 14-20). Of 20 galleries studied during July 1973, the mean number of eggs per gallery was 8, with limits of 3 and 14; however, oviposi- tion was not complete. Of 23 galleries stud- ied that contained a total of 193 eggs, 95 were on the top, 60 on the bottom, 25 on the right side, and 13 on the left side of the primary gallery. This suggests an order of preference in egg placement. The eggs usually lined the tunnel walls (Fig. 16). They were deposited in rows or scattered throughout the margins of the tun- nels of the gallery. The area of the gallery near the entrance where the male was lo- cated was usually free of eggs. 120 Great Basin Naturalist Vol. 37, No. 1 The incubation period for P. mitcronatiis was longer than for T. populi. The larvae were seldom collected with eggs, and one gallery contained 20 eggs with no larvae. Compared to T. populi, fewer eggs were laid and the female took greater care in de- positing them, apparently to insure their survival. The larger size of the egg may be the rea.son whv fewer were laid than in T. populi. The egg niche appeared to support, protect, and hold the egg in place. The bor- ing dust packed around the egg (Figs. 16, 17) apparently holds it in place, insulates it, may serve for food when the egg hatches, and may protect it from predators. L.\RVAE.— The superficial appearance of the larva was similar to T. populi. The length, distribution, and number of setae on tlie larval head capsule were characteristic of the species (Fig. 23). Widths of the head capsules of 91 larvae from 5 trees were measured as follows: 31 from 4 trees were .24-.29 mm (.27 ±.013 mm), and 60 from 5 trees were .33-.40 mm (.36 ±.021 mm). The index obtained from the means was: .367.27 = 1.33. Those .24-.29 mm were cat- egorized as Instar I and those .33-.40 mm as Instar II. Head capsules were wider than comparable stages of T. populi. The feeding patterns of the larvae in- dicated that more bark per given area was utilized in P. mucronatus (Fig. 18) than in T. populi (Figs. 6, 7), possibly resulting in more larvae per unit area. Adults.— The adults that emerged in the laboratory 5 February- 1 March 1975 (163 and 237) and 30 March-5 April 1976 (75 and 170) all totaled gave the sex ratio (male:female) of 1:1.7 (Table 9). Lengths were determined for 100 of these emergents (35 males and 65 females). The males were 1.92-2.41 mm (2.20 ±.117 mm) and the fe- Figs. 20, 21. Bark of dead aspen after brood has eimruc d pnniarv uiili i\ (. . u.l i i m . , white point- er I, parental adnit exit hole (black pointer), and exit holes of brood adults {unlabeled holes): 20, Trypophheus popi//i, bark split above primary galleries; 21, Procryphalus mucronatm, bark not split above primary galleries and entrances plugged with frass. March 1977 Petty: Aspen Bark Beetles 121 males were 1.96-2.53 mm (2.29 ±.109 mm). These measurements were taken from emer- gents of 30 April, 2 May, and 3 May. The length means for the emergents of each of these days were 2.32, 2.29, and 2.26 mm for the females and 2.27, 2.18, and 2.21 mm for the males. The emergents of the first day were longer than the subsequent ones. Overwintering.— All life stages over- wintered except the eggs and pupae (Tables 8, 10). There was mortality in the parental adults that overwintered in the new pri- mary chambers, but some also survived. The empty gut appearance of larvae collected November 1975 and March and April 1976 (Table 10) indicated that the larvae were Figs. 22, 23. Larval head capsules: fs, frontal shield: 22, Trypophloeus populi (Instar III); 23, Pwcn/phahis mucronatus (Instar II). Table 9. Number of reared males and females in the first hundred to emerge per week (5 February 1975 to 1 March 1975) and per day (30 April 1976 to 5 May 1976). {Procryphalus mucronatus). Collection date Emergence dates (n) (n) 29 Jan 1975 23 Apr 1976° 5-8 Feb 8-19 Feb 19-26 Feb 26 Feb 1 Mar Totals 30 Apr 1 May 2 May 3 May 4 May 5 May Totals 45 27 37 54_ 163 7 13 16 17 12 10 75 55 73 63 46 237 42 36 27 23 23 19 170 "Life stages present: adults and Instar 122 Great Basin Naturalist Vol. 37, No. 1 dormant during the winter. Overwintering larvae were found as deep as 7 mm from the surface of the bark. Because parental and newly transformed adults were found successfully overwintering, the adults may have some mechanism that enabled them to survive the cold winter months, a mecha- nism which was not present in T. papuli. The first eggs of the new season were laid fom one to two months earlier than in T. populi. The overwintered adults were ready to start new galleries and lay eggs as soon as the weather warmed up. Seasonal occurrence.— The eggs of the first generation were found as early as 24 May. One tree was under initial attack 17 June 1972. By August 1972 (58 days later), pupae and newly transformed adults were present. In the laboratory rearing, the dura- tion of time from the initial attack to the next generation was 60 days in one gallery. The seasonal occurrence of the various life stages (Table 8), the abundance and presence of overwintering stages (Table 10), and the two-month life cycle in one tree in the field and in one bolt in the laboratory indicated that there were one and one-half to two generations per year and that the overwintered adults laid the first eggs of the next season. These were followed by the eggs laid by the adults that developed from the overwintered Instar II larvae. Adults that developed from the overwintered Instar I larvae may have a later egg-laying period. 60 — n ^^ SS- 50- U5- ho- 35- 30 — 25- ~ 20 — [-1 15- ~i f— 10 — 5-_ -r Iff 30 — 25 25 — r— 1 20 — 15 — 10 — - 5 — 1r — --tvpoOOC\jjvOQOOc\j_SvOOOO r^ cr\ r^ .a ,j _3_:n_cjxr\ (\i C\J CM CVl "LAt^ON (HrAlTNt^ONr-t C\J CM CM r^r^r^r^o-\_3 O O CM (M tMCOCMr-> r-\ fr\ cr\ frs j:] o 1 8 1 8 C\J 1 O -3 s gR C\J OJ r^ r^ ro r^ _3 Figs. 24, 25. Numbers of larvae per size class of head capsule widths: 24, Trypophloeu.s populi, 232 larvae measured from 8 trees; 25, Procryphalus mucronatus, 91 larvae measured from 5 trees. March 1977 Petty: Aspen Bark Beetles 123 Territorial behavior.— The intraspecific territorial behavior consisted of the at- tempts of one beetle to take over another's place at the entrance to the primary gal- lery. Conflict for territory occurred between males and between females: between two females when one female tried to replace another female who had started her primary gallery, and between two males when one male tried to replace another male who had joined a female. The intruding beetle re- moved the resident by butting him (her) un- til he (she) backed out or was pushed out of his (her) entrance. This territorial behavior did not take place when the resident beetle was below the surface of the bark. At least half of the body of the resident needed to be protruding above the surface of the bark for an intruder to be successful. Combat between females was observed as follows: (1) The intruding female butted the defending female until she backed out of her gallery. (2) Both beetles repeatedly met head on (prothorax to prothorax). In this position they butted each other back and forth (Fig. 26, a). (3) The beetle with the apparent advantage repeatedly extended her front legs (Fig. 26, b) in an effort to dis- lodge the other from the tree (Fig. 26, c). The asperities and marginal teeth on the prothorax appeared to be well adapted for butting and dislodging. Dislodgment of one of the beetles from the tree also resulted from butting without extension of the legs. Butting occurred head on, from the side, and from behind. Very often the less aggressive of the two contenders was driven away be- fore dislodgment occurred. When an intrud- er located a defender who was too far into the gallery to be butted out, he either moved on or, apparently, tried to pull the defender out by clasping onto the posterior end of his elytra with his mandibles. In both beetle species, the territorial be- havior display was by the adult males, and females apparently selected for the more vigorous males at the new gallery. Mating behavior.— When a male located a female who was starting a gallery (Fig. 11), he either moved on or copulated with her (Fig. 12) and then continued to search for other females. For example, three males copulated in a 25-minute period with one female who was starting the primary gal- lery. The duration of each copulation was 5, 3, and 3 minutes. Copulations of one oth- er female with three males lasted for 3, 8, and 10 minutes. Repeated copulations were commonly observed. On three occasions a male was observed copulating with a female that had not yet located a site to initiate the gallery (Fig. 10). This promiscuity ap- parently insured fertilization even though the females outnumbered the males nearly two to one. Lone females were frequently found in plugged galleries with eggs and larvae, but no parental male. When a male located a female that was below the surface of the bark, he performed a mating behavioral sequence. The sequence Table 10. Relative abundance of overwintering life stages of Procrypliahis mucronatus. Collec- tion date No. of trees sampled Eggs No. of immatures Instars Pupae No. of adults Trans- Paren- formed^ tal" Feeding larvae I II (%)^ 11 Nov 75 9 Mar 76 23 Apr 76 23 Apr 76 3 1 1 4 0 0 0 0 27 63 0 4 76 0 20 24 0 pf pf 0 86" 20 0 0 0 0 pf < 10(?) 0 84 (?) ''In primary gallery. 'Percent of larvae that appeared dark because of bark in their intestines. .\ll the i 'There were 60 females and 26 males. "Female. 'Present, but numbers were not determined. white— their gut devoid of food material. 124 Great Basin Naturalist Vol. 37, No. 1 consisted of three distinct antics: nudging, stroking, and positioning. .Vf«^/gjng; The male repeatedly nudged the female bv brushing across the posterior end of her abdomen with his mandibles (Fig. 27). This motion was repeated from 6 to 12 times at a rate of approximately 4 per second (sample size: n=18). Stroking: The male vigorously (approx- imately 7 strokes per second, sample size: n= 1) stroked the female's abdomen with his prothoracic legs. It appeared as though he stroked both legs at the same time. He was positioned at the edge of the entrance (Fig. 27). Positioning: The male put his mandibles Tabl£ 11. Comparison of characteristics of Tnjpophloeus populi and Procryphalus mucronatus. Category T. populi P. mucronatus Attack 1. Live bark that is green— unhealthy trees 2. Mass attack— entire tree 3. Kills unhealthy trees Primary gal- lery 1. Cave-type plus food tunnel and secondary egg chamber 2. Parents often exit 3. Male blocks entrance 4. Bark covering gal- lery usually splits Eggs 1. Egg cluster Larvae 1. Three instars 2. Four prominent setae on frontal shield; 10 other prominent head capsule setae Larval mines 1. Distinct larval mines; fully plugged with frass Overwintering life stages 1. Larvae only Generations per year 1. 1-1'-^ Territorial behavior 1. Butting Male mating behavior 1. Position directly over entrance 2. Nondescript strok- ing, all legs in motion Antennal position when walking 1. Extended forward at 45- degree angle from side of head 1. Dying bark that is orange or brown, fermenting, and soft 2. Scattered or mass attack- bole of the tree 3. Little effect on decline of host 1. Linear, tunnel-type 2. Parents seldom exit 3. Entrance often plugged with frass; male inside gallery 4. Bark seldom splits 1. Egg niches 1. Two instars 2. Six prominent setae on frontal shield; 14 other prominent head capsule setae 1. Larvae enlarge primary gallery; mines short, partially plugged with frass 1. Larvae and adults 1. 11^-2 1. Butting and extension of prothoracic legs 1. Position at edge of en- trance 2. Nudging, stroking (prolegs only), positioning jerks 1. Extended at right angle to side of head March 1977 Petty: Aspen Bark Beetles 125 in contact with the abdomen of the female. He apparently clasped the posterior margin of her elytra with his mandibles and also grasped her with his prothoracic legs. In this position, he often jerked the posterior end of his body up and down. He jerked from 5 to 11 times at a rate of approx- imately 2 per second (sample size: n=ll). It appeared as though he was trying to po- sition her or stimulate her to position her- self for copulation. She appeared to be very imcooperative. She repeatedly backed part way out of the entrance to remove boring dust and then went back in to continue ex- cavation. The sequence of nudging, stroking, and positioning took approximately 2, 3, and 10 seconds, respectively (sample sizes: n=18, 18, and 8). The jerking antic (which was part of the positioning antic) took approx- imately two seconds (sample size: n = ll). The entire sequence was usually repeated several times before copulation was success- ful. From 3 to 30 minutes elapsed from the time the male located the female until cop- ulation. The apparent reasons for the mating an- tics were to insure intraspecific mating and prevent wasted time and energy in attempts to mate interspecifically. It was not unusual to find both T. popiili and P. mucronatus at- tacking the same tree. The antics of P. mucronatus were more complex and appar- ently more advanced phylogenetically than in T. populi. Locating infested trees.— Infestations of P. mucronatus were found in the same aspen groves as T. populi infestations and sometimes in the same tree. Diseased trees with fermenting bark were commonly at- tacked. Unhealthy trees typically died faster on the side that received the greater expo- sure to the sun. Sometimes P. mucronatus attacked this side while T. populi attacked the healthier, shaded side. Sometimes both species were in the same part of the tree, but P. mucronatus attacked several weeks after T. populi. Trees from which brood had emerged {P. mucronatus) did not have split bark covering the galleries, and many of the gallery entrances were still plugged with frass (Fig. 21). Host damage.— Infested trees were usual- ly diseased and infested with cerambycids and buprestids. The symptoms of the fungi Cenangium singular (Rehm.) Davidsen and Fig. 26. Prucrypluihts asic legs; c, dislodgement. uicronatus, territorial dislodgement behavior: a, butting; b, extension of prothor- 126 Great Basin Naturalist Vol. 37, No. 1 Figs. 27, 28. Mating ritual positions: 27, Procryphahis mttcronatits; 28, Trypophlocus popidi. Cash, and Cytospora spp. appeared on many of the infested trees. The following beetles were reared from P. miicronatus-iniested holts: Trypodendron rettisiim (LeConte) (Scolytidae); Saperda calcamta Say, Paral- lelina filicomis Casey, and Xylotrechus an- nosus Say (Cerambicidae); and Agrilus anx- iiis Gory and Dicerca tenebrica (Kirby) (Buprestidae). Two trees were studied that were half dead. The bark of the dead half was heavily infested with P. mucronatiis; nevertheless, the leaves that were present remained healthy and green from June to the end of the sea.son. These beetles {P. mucronatiis) were sel- dom important in hastening the death of the host. They accelerated the decomposi- tion process of dead parts of the tree. They tunneled and mined in the outer bark while the other borers attacked the cambium and xvlem of the host. Summary There are behavioral aspects and mor- phological characters that clearly distinguish T. populi from P. mucronatus in all stages with the exception of the pupae, which were not studied in detail in this work (Table 11). Both species play an important role in the decomposition process of dis- ea.sed and weakened aspen. Probably the most unusual discovery of this research was the territorial and mating behavior. An un- expected outcome was the success in locat- ing T. populi in sufficient numbers to in- clude it in the study. These beetles were associated with many other arthropods. Over 30 species of mites were collected from the beetles and their galleries. Most were phoretic; some were predacious. In addition, over 50 species of parasites and predators were collected and reared from the aspen infested with these beetles. A list of these species is in preparation. Acknowledgments This study was part of a Ma.ster of Sci- ence research project. The author thanks Dr. Stephen L. Wood for his direction and suggestions in the research and writing. Thanks are also extended to Dr. M. M. Fur- niss for his critical review, Dr. Samuel R. Rushforth for assistance with photography. Dr. Gary M. Booth for his suggestions, and to the late Dr. W. D wight Buchanan for his help in rearing the bark beetles. Special thanks are extended to my father. Dr. Wendell Pettv, for his support and en- couragement during this work. March 1977 Petty: Aspen Bark Beetles 127 Literature Cited Davidson, R. W., T. E. Hinds, and F. G. Hawkswobth. 1959. Decay of aspen in Colorado. Rocky Mt. Forest and Range Exp. Sta., Fort Col- lins, Colorado. U.S. Dept. Agr. Davidson, R. W., and T. E. Hinds. 1968. Disease of aspen in central and southern Rocky Mountains of U.S. Unpublished manuscript. Hagedorn, J. M. 1904. Biologischer nachtrag zur revi- sion unserer papelborkenkafer. Miinchener Koleopt. Zeitschr. 2:372-373. Harper, K. T., and J. H. Warner. 1972. Understory characteristics related to site quality for aspen in Utah. Brigham Young University Sci. Bull., Biol Ser. 16(2): 1-20. Krebill, R. G. 1972. Mortality of aspen on the Gros Ventre elk winter range. U.S. For. Ser. Res. Pap Int. 129. Intermountain Forest and Range Exp Sta., For. Ser., Ogden, Utah. Lamb, F. M. 1967. Aspen wood characteristics, proper ties, and uses. U.S. Forest Service Res. Pap. NC 13. 15 p. Lekander, a. B. 1968. The number of larval instars in some bark beetle species. Entomol. Tidsk. Arg. 89 (Hafte l-2):25^4. Little, E. L. 1971. Atlas of U.S. trees. Vol. 1. Conifers and important hardwoods. U.S. Dept. Agr. For. Ser. Misc. Publ. 1146. Costing, H. J. 1956. The study of plant communities. W. H. Freeman Co., San Francisco, California, 440 p. Palm, T. 1959. Die Holz- und Rinden-kafer der sud- und mittelschwedischen laubbaume. Opusc. Ent. Suppl. 16:350-351. Preston, R. J. 1969. North American trees. Iowa State University Press, Ames, Iowa. .395 p. Reynolds, H. G. 1969. Aspen grove use by deer, elk, and cattle in southwestern coniferous forest. U.S. Dept. Agr. For. Ser. Res. Note RM-1.38. 4 p. Wallmo, O. C. 1972. Forage use by mule deer rela- tive to logging in Colorado. J. Wildlife Mgt. .36:1025-10.33. Wood, S. L. A taxonomic monograph of the Scolytidae (Coleoptera) of North and Central America, ms. in preparation. FOOD OF TOWNSEND GROUND SQUIRRELS ON THE ARID LAND ECOLOGY RESERVE (WASHINGTON) Mark K. Johnson' Abstract.— Diets of townsend ground squirrels were quantified ft-om the Arid Land Ecology (ALE) Reserve eastern Washington. Little has been reported on townsend ground squirrel {Spennophilus townsendi) diets. Alcorn (J. Mamm. 21:160-170, 1940) stated that the species fed mainly on green vegetation. Davis (J. Mamm. 20:182-190, 1939) reported several species of plants in their diets. Scheffer (J. Mamm. 22:270-279, 1941) related the seasonal activity of the species to the seasonal abundance of green vegetation. Twelve ground squirrels were live- trapped on the Arid Land Ecology (ALE) Reserve during March-May 1974 in an Ar- temisia tridentata (big sage) /Agropj/ron spicatum (bluebunch wheatgrass) association (Daubenmire, Wash. State Agric. Exp. Sta. Tech. Bull. 62:131, 1970). The ALE Reserve is 15 miles northwest of Richland, Washing- ton, on the Energy Research and Devel- opment Administration's Hanford Works Reservation. This ground squirrel is not ac- tive above ground during the remaining months of the year at the elevation (1200 ft:366 m) of the study site. The diet analysis technique employed was the same as that reported by Hansen and Flinders (Science Series No. 1, Range Sci- ence Dept., Colorado State Univ., 1969). Five microscope slides were made from the stomach contents of each ground squirrel, and 100 fields were quantified for each stomach sample. There were 14 foods identified in the analysis: Poa sandbergii (49 percent), Ach- illea lanulosa (11 percent), Liipinus laxif- lorus (10 percent). Astragalus purshii (11 percent), Descurania pinnata seed (8 per- cent), Erigeron filifolius (3 percent), Anten- naria dimorpha (2 percent), Salsola kali (2 percent), Artemisia tridentata (2 percent), Sisymbrium altissimum (2 percent), unclassi- fied forb (3 percent), Balsamorhiza careyana (<1 percent), Lactuca serriola (<1 per- cent), lichen (<1 percent) and arthropod (Ol percent). Staple foods of townsend ground squirrels were almost entirely green vegetation. The above-ground activity of the species seemed to be highly correlated with the seasonal abundance of the major food, Poa sand- bergii. 'Range Science Department, Colorado State University, Fort Collins, Colorado 80523. 128 NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be di- rected to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be sim- ilarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. They should be typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. Use a recent issue of either journal as a format, and the Council of Biology Edi- tors Style Manual, Third Edition (AIBS 1972) in preparing the manuscript. An abstract, about 3 percent as long as the text, but not exceeding 200 words, written in accordance with Biological Abstracts guidelines, should precede the introductory paragraph of each ar- ticle. Authors may recommend one or two reviewers for their article. All manuscripts re- ceive a critical peer review by specialists in the subject area of the manuscript under con- sideration. Manuscripts that are accepted and that are less than 100 printed pages in length will automatically be assigned to the Great Basin Naturalist. Those manuscripts larger than 100 printed pages in length will be considered for the Memoirs series. Illustrations and Tables. All illustrations and tables should be made with a view toward having them appear within the limits of the printed page. Illustrations that form part of an article should accompany the manuscript. Illustrations should be prepared for reduction by the printer to either a single-column (2 5/8 inches) or double-column (5 1/2 inches) width, with the length not exceeding 7 1/2 inches. Costs Borne by Contributor. Contributors to the Great Basin Naturalist should be pre- pared to donate from $10 to $30 per printed page toward publication of their article (in addition to reprint costs outlined in the schedule below). Authors publishing in the Great Basin Naturalist Memoirs may be expected to contribute $35 per printed page in addition to the cost of printed copies they purchase. No printed copies are furnished free of charge. A price list for reprints and an order form are sent with the galley proof to contributors. Reprint Schedule of the Great Basin Naturalist 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each 100 copies $20 $24 $28 $32 $36 $40 additional 200 copies 28 32 36 40 44 48 2 pp. 300 copies 36 40 44 48 52 56 $4 Great Basin Naturalist Memoirs No. 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodburry, H. H. Frost. $10. No. 2 Intermountain biogeography: A symposium. By K. T. Harper, J. L. Reveal, et al. $15. GREAT BASIN NATURALIST Editor. Stephen L. Wood, Life Science Museum, Brigham Young University, Provo, Utah 84602. Assistant Editor. Stanley L. Welsh, Life Science Museum, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; William D. Tidwell, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agri- cultural Sciences; Ernest L. Olson, Director, Brigham Young University Press, Univer- sity Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been pub- lished from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only previously unpublished manuscripts of less than 100 printed pages in length and pertaining to the biological natural history of western North America are ac- cepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manu- scripts are subject to the approval of the editor. Subscriptions. The annual subscription rate of the Great Basin Naturalist is $12 (outside the United States the rate is $13). The price for single numbers is $4 each. All back num- bers are in print and are available for sale. All matters pertaining to the purchase of sub- scriptions and back numbers should be directed to Brigham Young University Press, Mar- keting Department, 204 UPB, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same office at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either jour- nal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts or other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back © 1977 Brigham Young University Press. All rights reserved Printed in United States of America 650 12-77 27561 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University Volume 37 June 30, 1977 No. 2 HOST-INDUCED MORPHOLOGICAL VARIATIONS IN THE STRIGEOID TREMATODE POSTHODIPLOSTOMUM MINIMUM (TREMATODA: DIPLOSTOMATIDAE). II. BODY MEASUREMENTS AND TEGUMENT MODIFICATIONS James R. Palmieri' Abstract.— Extensive feeding experiments were undertaken to determine if physiological host specificity was a major characteristic of Posthodiplostomum minimum. This involved the feeding of experimentally infected sunfish livers containing metacercariae of P. minimum to amphibian, reptilian, avian and mammalian hosts. Host-induced morphological variations of adult P. minimum were shown to be associated with the genus and class of host em- ployed as well as with the inherent variability of P. minimum exclusive of host factors. Morphological characters such as body size and shape and tegumental surface and spines are indicated as undergoing morphological varia- tion. Of all the experimental definitive hosts used, avian and mammalian are the most suitable for normal devel- opment of P. minimum. Amphibian and reptilian hosts demonstrate marked variation in the adult worm devel- opment usually manifested by pronounced size decrease. Posthodiplostomum minimum (MacCal- lum, 1921) is a strigeoid trematode of the family Diplostomatidae Poirier, 1886. Adults of this species (Fig. 30) parasitize the intestine of piscivorous birds and the meta- cercarial stage is found in various fresh- water fishes. Two subspecies of P. minimum have been reported, based upon the ability of cer- cariae to penetrate and develop either in centrarchid or cyprinid fish hosts (Hoffman, 1958). The subspecies used in this in- vestigation is the centrarchid strain (Pal- mieri 1975). Since Stunkard's report on intraspecific variation in 1957, several more recent ex- perimental studies have shown that size, shape, and position of various organs and structures in helminths may be considerably modified by the host. For many years, in- vestigators such as Dubois (1944, 1955, 1968, and 1970) have delineated species of strigeoids largely on the basis of host specif- icity. Recently, however, several in- vestigators have shown that parasites can indeed develop within hosts that normally would be ecologically isolated from in- volvement in the normal life cycle of the parasite (Blankespoor 1971, Campbell 1972, Palmieri 1973, Ulmer 1961, Watertor 1967). The lakes region of northwestern Iowa is an area rich in conditions requisite for the production of both snail and fish inter- mediate hosts of Posthodiplostomum min- imum. This area also serves as both a feed- ing and nesting area for piscivorous avian hosts needed in maintaining the life cycle of P. minimum. Experimental infections of a variety of amphibian, reptilian, avian, and mammalian hosts with experimentally developed meta- cercariae of P. minimum were carried out 'Department of Zoology, Iowa State University, Ames, Iowa 50010. Present address. Institute for Medical Research, Pahang Road, Kuala Lumpur, 12- 14-Malaysia. UCICMR. 129 130 Great Basin Naturalist Vol. 37, No. 2 from 1971 to 1974 at Iowa State University and the Iowa Lakeside Laboratory. Adult P. minimum recovered from these ecologically abnormal hosts were examined for host-in- duced morphological variation. Materials and Methods Three eggs of Posthodiplostomum min- imum, obtained from a single gravid worm from an experimentally infected chicken 48 hours postexposure, were placed in an em- bryological watch glass with filtered lake water. Hatching of the miracidia occurred 20 to 21 days later. A single miracidium was exposed to a laboratory-reared Physa gyrina and pentra- tion was observed. This snail was isolated in a one-gallon aquarium and maintained in the laboratory until shedding of cercariae took place (48 days postpenetration). Twice daily for 10 days, contents (shed cercariae) of the one-gallon aquarium were poured into an aquarium containing parasite-free, laboratory-maintained simfish. Once in- fected, sunfish were then maintained at room temperature for 45 days. These sun- fish livers served as the source of metacer- cariae for subsequent experimental feedings to definitive hosts. All definitive hosts which had been ex- posed to laboratory-developed metacer- cariae were autopsied from 49 to 96 hours postinfection. Adult worms so obtained were washed in the appropriate Ringer's so- lution and were prepared for light micro- scopy or scanning electron microscopy. x\ficroscopi/.— Specimens to be examined by scanning electron microscopy were fixed in a modified Parducz (1967) fixative (6.0 ml of 2 percent O^O^ and 1.0 ml of saturated mercuric chloride) for one minute at 0 C. All specimens were then washed in distilled water three times at 15-minute intervals. Entire specimens were rapidly dehydrated in ethanol using critical point drying tech- niques as described by Hearle, Sparrow, and Cross (1972), Cohen and Shaykh (1973), Pol- liack, Lampen, and de Harven (1973), and Lewis and Nemanic (1973). Dried specimens were then affixed by electrically conductive aluminum paint to cleaned brass plates and secondarily affixed to brass specimen holders. Specimens were initially coated with carbon and were sub- sequently given a double coat of gold-palla- dium. All specimen coating was done with the aid of an Edwards vacuum evaporator. Coated specimens were viewed and photo- graphed on a Jeolco JSM-Sl scanning elec- tron microscope at an accelerating voltage of 10 KV. All micrographs were recorded on Kodak Ektapan 4162 negative film and developed in a mixture of six parts Kodak D-76 and one part Kodak D-19 for max- imum resolution and negative contrast. Experimental /n/ecf tons.— Definitive hosts were force-fed simfish livers experimentally infected with over 100 metacercariae of P. minimum. Once fed, all hosts were main- tained in appropriate cages or aquaria and fed only water. After a suitable devel- opmental period of 49 to 96 hours, these hosts (Table 1) were examined for the pres- ence of P. minimum adults, using standard routine laboratory methods. Details of all hosts exposed to cercariae and those fed metacercariae of P. minimum have been re- ported by Palmieri (1976). Results and Discussion Body Measurements.— Five variables and relationships were analyzed for the body measurements of P. minimum (Table 1). These include (1) body length (BL) (from the anterior margin of the forebody to the posterior of the hindbody, exclusive of the extended bursa, when present); (2) body width (BW) (at the widest portion of the forebody); (3) distance from the anterior Table 1. Total number of vertebrate hosts exposed to fish hvers experimentally infected with metacercariae of P. tninimum. Total No. No. Class Positive Negative Species Hosts Amphibians 17 4 4 21 Reptiles 8 2 3 10 Birds 18 9 8 27 Mammals 13 9 6 22 TOTAL 56 24 21 80 June 1977 Palmieri: Trematode Variations 131 margin of the forebody to the anterior mar- gin of the acetabuhim (DAA); (4) from the posterior margin of the acetabuhim to the posterior margin of the hindbody (DPP); and (5) the ratio of DAA /DPP. This rela- tionship (DAA/DPP) was chosen rather than forebody and hindbody lengths because of the lack of uniformity of a distinct de- marcation between these regions in some specimens. It was also decided that using the ratio DAA/DPP would minimize the ef- fect of size differences of adults of P. min- irnum because the position of the acetabu- lum is relatively constant and serves as a more uniform reference point. A complete tabulation of the above data follows in Table 2. Body size (length and width) of adult P. minimum is a significant characteristic in identifying the class of definitive host used for experimental development of adult worms. At the class level, both body size and position of demarcation between the forebody and hindbody regions of P. min- imum vary significantly. In worms recov- ered from amphibian and reptilian hosts, very little demarcation can be noted (Figs. 1-3, 9-13). Furthermore, in many worms developed within these poikilothermic hosts, considerable invaginations of the anterior margin of the forebody occur (Figs. 11-12). In many worms the hindbody is not present (Fig. 1-2) or is poorly developed (Fig. 3). In some experimental avian hosts fed metacer- cariae of P. minimum, however, adult worms appear normal and well developed with a well-demarcated forebody and an elongate, cylindrical hindbody (Figs. 6, 8, 14, 16-19). In the most common definitive host for P. minimum, the great blue heron, worms recovered from this naturally in- fected host demonstrate the most character- istic body form (Fig. 19). Those worms re- covered from mammalian hosts appear to be more normal in their development than those recovered from amphibian and repti- lian hosts but vary somewhat from those re- covered from avian hosts in possessing hind- bodies whose lengths are reduced in proportion to the forebody (Figs. 4, 5, 7, 15, 17, 18, 20-21). Tegumental modifications.— Seversd host- induced modifications of the tegument were noted during the course of this study. Scanning electron microscopy of the tegu- ment of P. minimum specimens recovered from a variety of experimentally fed hosts revealed that tegumental spines underwent morphological modification. Two regions of the adult worm were selected for observa- Table 2. Body measurements of P. minimum from experimental definitive hosts.' Variable Mean of Combined Standard '^^^"^ Host Class Deviation Amphibian Reptilian Aves Mammalia Body Length (BL) 01 Body Width (BW) 0.2 Distance from Anterior Margin of Body to Anterior Margin of Acetabulum (DAA) Distance from Posterior Margin of Acetabulum to Posterior Margin of Body (DPP) Ratio of DAA/DPP 0.188 0.260 0.735 0.083 0.517 0.435 0.614 0.043 0.269 0.230 0.288 0.056 0.589 0.296 0.184 0.067 0.243 0.198 0.,321 0.274 0.238 0.867 0.554 0.664 0.634 'AH measurements in mm 132 Great Basin Naturalist Vol. 37, No. 2 Figs. 1-8. Morphological variation of body .shape of adult P. minimum recovered from vertebrate hosts. Note variations in body shape and hindbody demarcation. All specimens 72 hours old (Scale: 1 inch = .120 mm). Hosts are: 1, Bufo americanus Note lack of hindbody; 2, Amblystonui tigrinum Note lack of hindbody; 3, Chry- June 1977 Palmieri: Trematode Variations 133 semys picta Note bulblike hindbody and extended; 4, Didelphis marsupialis Note slightly reduced forebody; 5, Felis catus Note large forebody; 6, Larus argentatus Note well-developed forebody and hindbody; 7, Meriones unguiculatus Note reduced forebody; 8, Gallus domesticus Note reduced forebody. 134 Great Basin Naturalist Vol. 37, No. 2 Figs. 9-21. Outline drawings of adult P. minimum from amphibian (Figs. 9-11), reptilian (Figs. 12-13), avian (Figs. 14-16, 19), and mammalian (Figs. 17-18, 20-21). All specimens 72 hours old (Scale: 1 inch = 0.1 mm). Hosts are: 9, Amhystoma tigrinum; 10, Bufo americanus; 11, Rana pipiens; 12, Thamnophis radix, 13, Chrysemys picta; 14, Lartis argentatus; 15, Columha livia; 16, Zenaidura macroura; 17, Meriones unguiculatus; 18, Didelphis marsupialis; 19, Ardea heriodias; 20, Cants familiaris; 21, Felis catus. Figs. 22-29. Morphological modification of the tegument of P. minimum due to influences of various verte- brate hosts. Figs. 22-25 from an area lateral to and between the acetabulum and holdfast organ. Figs. 26-29 from an area of the dorsal hindbody (Scale: X = 30,000). Hosts are: 22, Ambystoma tigrinum (Note the complex na- ture of the tegumental spines and surrounding tegumental surface); 23, Meriones unguiculatus (Note similarity to Fig. 22 with some loss of complexity of tegumental spines and surface); 24, Chrysemys picta (Note that tegumen- tal spines have been greatly reduced and are joined to the tegument by a netlike or weblike process); 25, Iguana iguana (Note the reduction of tegumental spine serration and surface tegument); 26, Chrysemys picta; 27, Bufo americanus; 28, Rana pipiens; 29, Amhystonui tigrinum. June 1977 Palmieri: Trematode Variations 135 #v-^'«^-.^fr, ""^''^Vfv:.]!?, 136 Great Basin Naturalist Vol. 37, No. 2 tion: (1) an area lateral to and equidistant between the acetabulum and holdfast (Figs. 22-25) and (2) a middorsal area on the hindbody (Figs. 26-29). Although no phy- logenetic relationships or trends could be discerned, tegumental spination of the ven- tral forebody surface was reduced from the normal complex structure (Fig. 22). Normal spines are large with serrated margins and are surrounded by tegument containing many surface modifications (Figs. 22-23). In specimens collected from Chrysemys picta, spines were so reduced that only a netlike or weblike surface area remained. Such greatly reduced spines are connected to one another as well as to the underlying tegu- ment by filamentous strands (Fig. 24). Tegu- mental spines examined from most speci- mens recovered from vertebrate hosts were reduced when compared to others which were more highly developed. There is no apparent relationship between the class of host and complexity of tegumental spine structure. A typical example of a reduced tegumental spine can be found in Figs. 23 and 25, taken from Iguana iguana and Me- riones unguiculatiis, respectively. On the middorsal hindbody of P. min- imum, the tegument shows some surface modification ranging from a folded appear- ance (Figs. 26 and 27) {Chrysemys picta and Biifo americamis) to one in which bleblike modifications of the tegumental surface pre- dominate (Fig. 28) {Rana pipiens). A tegu- ment consisting of irregular ridges (Fig. 29) is also common among worms developed within amphibian hosts. Morphological modification of the tegument and associated surface structures are independent of the class of host used for experimental devel- opment of the adult P. minimum. Acknowledgments The author thanks Dr. Martin J. Ulmer for his time and guidance throughout this research project. Dr. D. Cox and Paul Du- bose for data analysis. Dr. Darwin Wittrock for aid in fieldwork and Mr. James Amrine for specimen preparation for S.E.M. Appre- ciation is also extended to the Iowa State Conservation Commission, and Federal Bu- reau of Sport Fisheries for animal collec- tion; the Iowa State University Departments of Zoology, Genetics, and Statistical Com- putations. This study was supported in part by the National Science Foundation Grant Fig. .3(). Diagram of adult P. minimum from the gull {Lams argentatus) depicting major organs undergoing morphological variation: A— acetabulum; E— egg; EO— esophagus; F— forebody; G— genital bursa; H— holdfast organ; HB— Hindbody; I— intestinal cecum; O— oral sucker; OV— ovary; P— pharynx; TA— anterior testis; TP— posterior testis; V— vitellaria. V 1977 Palmieri: Trematode Variations 137 No. GB-23057; the Thomas H. MacBride Scholarship from the Iowa Lakeside Labora- tory; Brigham Young University Depart- ment of Zoology; the Department of Inter- national Health, School of Medicine, University of California, San Francisco; and the National Institute of Allergy and In- fectious Diseases, National Institutes of Health, U.S. Public Health Service Grant AI 10051 (UC ICMR). Literature cited Blankespoor, H. D. 1971. Host-parasite relationships of an avian trematode, Plagiorchis noblei Park, 1936. Ph.D. Dissertation, Iowa State University, Ames, Iowa. (Diss. Abstr. 31:12). Campbell, R. A. 1972. New experimental hosts of Posthodiplostomum minimum (Trematoda: Diplos- tomatidae). J. Parasitol. 58:1051. Cohen, A. L., and M. Shaykh. 1973. Fixation and de- hydration in the preservation of surface structures in critical point drying of plant material. Proc. Workshop Scan. Electr. Microscop. Path. 3:371- 378. Dubois, G. 1944. A propos de la specificity parasitaire des Strigeida. Bull. See. Neuchat. Sci. Nat. 69. 103 pp. 1955. Nature de la specificity chez Strigeides (Trematoda). Rev. Iber. Parasitol. Tomo Extraor- dinario: 133-155. Synopsis des Strigeidae et des diplos- Hearle, J. W. S., J. T. Sparrow, and P. M. Corss. 1972. The use of the scanning electron micro- scope. Pergamon Press, New York, 278 pp. Hoffman, G. L. 1958. Experimental studies on the cercaria and metacercaria of a strigeoid trema- tode, Posthodiplostomum minimum. Lewis, E. R., and M. K. Nemanic. 1973. Critical point drying techniques. Proc. Workshop Scan. Electr. Microsc. Path. 3:767-774. Palmieri, J. R. 1973. Additional natural and experi- mental hosts and intraspecific variation in Post- hodiplostomum minimum (Trematoda: Diplos- tomatidae). J. Parasitol. 59:744-746. 1975. Host parasite relationships and in- traspecific variation in the strigeoid trematode Posthodiplostomum minimum (Trematoda: Diplos- tomatidae). Utah Acad. Sci., Arts and Letters. 52:68-69. 1976. Host parasite relationships and tomatidae (Trematoda). Soc. Neuchdt. Sci. Nat. 1-258 pp. — 1970. Synopsis des strigeidae et des diplo- stomatidae (Trematoda). Soc. Neuchdt. Sci. Nat. 259-727. traspecific variation in the strigeoid trematode Posthodiplostomum minimum (Trematoda: Diplos- tomatidae). Controlled experimental development in ecologically abnormal hosts. Great Basin Nat. 36(3)334-346. Parducz, B. 1967. Ciliary movement and coordination in ciliates. Int. Rev. Cytol. 21:63-67. PoLLiACK, A., N. Lampen AND E. DE Harven. 1973. Comparison of air drying and critical point pro- cedures for the study of human blood cells by scanning electron microscopy. Proc. Workshop Scan. Electr. Microscop. Path. 3:529-534. Stunkard, H. W. 1957. Intraspecific variation in para- sitic flatworms. Syst. Zool. 6:7-18. Ulmer, M. J. 1961. Passerine birds as experimental hosts for Posthodiplostomum minimum (Trema- toda: Diplostomidae). J. Parasitol. 47:608-610. Watertor, J. L. 1967. Intraspecific variation of adult Telorchis bonnerensis (Trematoda: Telorchiidae) in amphibian and reptilian hosts. J. Parasitol. 53:962-968. THE SUBFAMILY ANOMIOPSYLLINAE (HYSTRICHOPSYLLIDAE: SIPHONAPTERA). I. A REVISION OF THE GENUS ANOMIOPSYLLUS BAKER^^ A. M. Barnes, ' V. J. Tipton, ' and J. A. Wildie' Abstract.— In this taxonomic revision of the Nearctic flea genus AnomiopsyUus Baker morphological charac- ters, phylogeny, affinities with other genera in the subfamily Anomiopsyllinae, evolutionary adaptations, geogra- phic distribution, ecological parameters, and host preferences are discussed. Characters primarily associated with the male genitalia, in particular the number and placement of spiniforms on the movable process (finger) of the clasper, are used for species discrimination. A key to the males of the 15 species and subspecies is provided, in- cluding illustrations of both males and females. The Nearctic genus AnomiopsyUus Baker is a group of small, eyeless fleas associated primarily with wood rats of the genus Neo- torna and usually found in their nests. These fleas are characterized by the extreme re- duction in the number of setae and spiny structures common to most Siphonaptera, and by the reduction and displacement of certain skeletal structures which presumably strengthen the thorax for jumping. These anatomical modifications are carried further in the genus AnomiopsyUus than in any oth- er flea genus and reflect a high degree of adaptation to the nest environment. The genus was proposed by Baker (1904) to include Typhlopsylla nudatus Baker, 1898. The subfamily Anomiopsyllinae was established by Baker (1905) to include Ano- miopsyUus, which was then thought to be phylogenetically isolated and without near relatives. Since Baker's description of nu- datus, 11 additional AnomiopsyUus species have been described, of which caUfornicus Baker has been synonomized (Jordan and Rothschild 1915) and congruens Stewart has been reduced to a subspecific rank (Hopkins 1952). Traub and Tipton (1951) gave an ac- count of the relationships of the genera within the subfamily Anomiopsyllinae, using characters fotmd in females. Hopkins and Rothschild (1962) reviewed the seven known species and presented keys classifying all males and some females (three species and one subspecies). While the genus has been treated ade- quately so far as its placement among the higher taxa is concerned, very little is known of the distribution, geographical var- iation, biology, or ecology of any of the species. No taxonomic evaluation using all available specimens has been made for any of the species, nor have the intrageneric relationships been analyzed. The primary purpose of this paper is to provide a tax- onomic and systematic evaluation of the genus and the member species on the basis of morphological, geographical, and ecolog- ical data available from slide-mounted spec- imens. Distribution, host-parasite relation- ships, and phylogenetic relationships are also discussed. Acknowledgments Gratitude and deep appreciation are due Dr. Deane P. Furman, under whose direc- tion the original Ph.D. studies of the senior author were conducted, and to Dr. E. Gor- This paper is based on part of an original dissertation completed by the senior £ modified and updated to include all known species of AnomiopsyUus. •This work was supported in part by grant number 2 ROl AI 11292-04 from the National Institutes of Health. 'CDC Laboratory, Box 2087, Fort Collins, Colorado 80522. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. the University of California, Berkeley, in 1963. It has been 138 June 1977 Barnes et al.: Anomiopsyllus Fleas 139 ton Linsley and Dr. Frank E. Pitelka for their advice and criticism. Special thanks are expressed to Dr. Frank M. Prince and Dr. Harold E. Stark (who were formerly with the U.S. Public Health Service) as well as to Mr. John R. Walker and Mr. Keith F. Murray of the State of California Department of Public Health for their kindness and cooperation in making the studies possible. Appreciation is also expressed to the fol- lowing persons for providing advice, infor- mation, or material loaned from personal or institutional collections: Dr. D. Elden Beck, Dr. Cluff E. Hopla, Dr. G. F. Augustson, Mr. F. G. A. M. Smit, Dr. John Garth, Dr. William L. Jellison, Dr. Luis Vargas, Dr. Alfredo Barrera, Dr. Robert Traub, and Dr. G. P. Holland. Host-Parasite Relationships Species of Anomiopsyllus are most fre- quently associated with wood rats of the genus Neotoma; however, they have been collected occasionally from many ground- dwelling rodents, including species of Per- omyscus, Dipodomys, and Spermophilus. There are currently three species of Ano- miopsyllus that have not yet been associ- ated with Neotoma: A. oaxacae Barnes, in which the single specimen was taken from Baiomys musculus; A. traubi Barrera, which is known only from a "rodent's nest"; and A. martini Holland, which was described from Sciurus aberti, a squirrel. The rodent genus Neotoma Say and Ord is widely distributed in North America. The species are normally restricted to a given type of habitat varying from low, dry deserts or humid jungles to rocky slopes above the timberline (Hall and Kelson 1959). Geographical distribution of this genus ranges from Honduras and Nicaragua in Central America to British Columbia in Canada and from California east to New York and Florida in the United States. There are 22 known species of Neotoma comprising 114 taxa on the species and sub- species level. Most flea species of the genus Anomiopsyllus are associated with wood rats, and most occur in the arid south- western United States and Mexico where the average rainfall does not exceed 76 cm. The temperatures in these areas undergo great daily fluctuations and are generally quite high. Nest fleas are somewhat pro- tected from ambient fluctuations because the temperature and humidity in the nest remain at a fairly constant level. Under these circumstances a nest environment may provide some selective advantages over a host environment inasmuch as a host may be away from the nest for long periods of time foraging for food and, thus, the fleas on the host would be exposed to environ- mental fluctuations. Distribution of the Genus Anomiopsyllus is exclusively Nearctic, and species have been collected in a geo- graphical area bounded by Banff, Alberta, Canada, on the north (A. montanus Collins) and the state of Oaxaca, Mexico, on the south (A. oaxacae Barnes). Anomiopsyllus is restricted in the United States between the coastal range of California on the west (A. falsicalifornicus Fox) and the 101st meridian on the east (A. nudatus hiernalis Eads and Menzies). No species have been collected east of the Mississippi River, even though species of Neotoma do occur in the eastern United States. Generic Relationships Eopsylla Argyropulo is the only genus in the subfamily which occurs outside the Nearctic realm. Eopsylla was included in the tribe Anomiopsyllini by Hopkins and Rothschild (1962) because it is more closely related to the North American genus Callis- topsyllus Jordan and Rothschild than it is to other Asian genera. Careful collecting in northwestern North America may reveal the presence of taxa which will further sub- stantiate this relationship. Genera of the Anomiopsyllinae which are exclusively North American include: Ano- miopsyllus, Callistopsyllus, Megarthroglossus Jordan and Rothschild, Stenistomera Rothschild, and Conorhinopsylla Stewart. Callistopsyllus has the most northern distri- 140 Great Basin Naturalist Vol. 37, No. 2 bution of these genera and is well repre- sented in western Canada by three species (Holland 1949), two of which enter the United States by way of the Sierra-Cascade mountain chain and the Rocky Mountains. Megarthroglossus is also represented in Can- ada and extends as far south as Texas where M. divisus (Baker) and M. bisetis Jordan and Rothschild have been collected (Mendez 1956). Conorhinopsylla is an eastern genus and is not found further west than Kansas. Mendez (1956) has stated that Megarth- roglossus is more closely related to Ano- miopsyllus than any other genus, and affi- nities have been shown between these genera based on comparative female charac- ters (Traub and Tipton 1951). However, structures of the male genitalia of Callistop- syllus are strongly reminiscent of Anomiop- sylhis, especially with regard to the clasper and its movable process. The articulation of the movable process in CallistopsyUus deiit- erus Jordan is very similar to that of A. am- phibolus Wagner and A. montanus; the process resembles that of A. amphibolus, both in shape and in the presence of two spiniforms. The aedeagi of all three genera have much in common. However, the ae- deagus of Anomiopsyllus is reduced and has no prominent dorsal or subterminal lobes. The affinities of Conorhinopsylla and Ste- nistomera with the rest of the Anomiopsyl- linae are also borne out by similarities in male genitalia. Both distributional and morphological evidence indicate that Anomiopsyllus is a member of a monophyletic group, and that the genus has evolved in North America from an ancestor common to other members of the Anomiopsyllinae. Anomiopsyllus is more closely related to both CallistopsyUus and Megarthroglossus than to either Ste- nistomera or Conorhinopsylla. In all proba- bility, Anomiopsyllus has evolved exclusi- vely in North America in conjunction with the Nearctic genus Neotoma, its wood rat host. Skeletal Anatomy of Adults The remarkably reduced nature of some structures in Anomiopsyllus has evoked comment from every author discussing the genus, and in two instances (Ewing and Fox 1943, Hubbard 1947) the genus has been in- cluded in the Pulicoidea rather than in the Ceratophylloidea where it was rightfully placed by Jordan and Rothschild (1915). The past taxonomic confusion has undoubt- edly resulted from loss of the more obvious distinctive characters typical of Ceratophyl- loidea, which has brought about a super- ficial resemblance of Anomiopsyllus and the Pulicoidea. The outstanding features of Anomiop- syllus anatomy include complete eye- lessness, an extreme reduction in number of setae and spines, extensive loss and fusion of structures in the metathorax, and loss of structures in both meso- and metacoxae. In contrast to the highly modified nature of the thorax, the abdominal segments are only moderately specialized, though marked by loss of some structures in the male genitalia. Of these characters, eyelessness has arisen in a number of widely separated genera and seems to be associated with nest dwellers (Traub 1953). Reduction in setation in Siphonaptera has been carried to its greatest extreme in Ano- miopsyllus. Loss of setae and of important spines imdoubtedly has great adaptive as well as taxonomic significance. The adap- tive significance of such losses is not clear because several highly successful nest fleas which are in some respects as specialized as Anomiopsyllus (e.g. Stenistomera) have gone in the opposite direction by developing nu- merous large setae. In the following section, the more fimda- mental aspects of Anomiopsyllus anatomy are emphasized, especially those in which the genus differs from its relatives and those which would seem to have clear adaptive significance. Snodgrass (1946) has dealt ex- tensively with the skeletal anatomy of fleas; his terminology and interpretations are used here as a basis of comparison. Head Fig. 1 The head capsule is oval in shape, but its outline is broken anteriorly by a well-de- June 1977 Barnes et al.: Anomiopsyllus Fleas 141 fined clypeal tubercle. There is no inter- antennal suture. The eyes are completely lacking, as in Callistopsyllus; the allied Me- garthroglossus has vestigial eyes. The degree of sexual dimorphism in an- tennal size and shape is striking. The small, oval antennae of the female do not reach the prostemum. The male antennae are greatly lengthened with their bases higher up on the head capsule and their apices ex- tending back upon the prosternosome on which the antennal fossae are continued. Though deep, the antennal fossae as seen in a posterior view of the head capsule do not meet in the interior of the head capsule, nor are they connected by a trabeculum. The mouthparts bear no structures not duplicated elsewhere in Siphonaptera. The extended members are long in proportion to the remainder of the flea and reach the tro- chanter of the procoxae. The labial palpi are 4-segmented, and the stylets are armed their full length with minute, laterally placed nodules. The epipharyngeal stylet is completely unarmed, as illustrated by Snod- grass (1946) for Opisodasys Jordan, and is unlike those of Hystrichopsylla Taschenberg and others which bear teeth. .A. /ti ^ •^'■•'■, .,----— K- L_ yi^'^j .--^ ^ '- AT. 1 1 sy **^ ■ — V-v \. / Fig. 1 Anomiopsyllus amphibolus: Head and Thorax. 142 Great Basin Naturalist Vol. 37, No. 2 The anterior tentorial arms remain com- plete and extend from a point near the ten- torial bridge to connect anteriorly near the genae. Snodgrass suggested that these arms are in reality the dorsal tentorial arms since true anterior arms arise in the clypeal area. The completeness of these arms and lack of a trabeculum centralis are considered by Johnson (1957) to be primitive characters associated with the Hystrichopsyllidae. In the Ceratophyllidae, the anterior arms are in the process of being lost and replaced by the trabeculum. In its remarkable antennal sexual di- morphism, Anomiopsyllus is highly ad- vanced within the Hystrichopsyllidae, in which the more primitive members show little difference in this regard between the male and female. Within the Anomiopsyl- linae all genera show some degree of sexual dimorphism, but of the genera examined Anomiopsyllus and Callistopsyllus show the greatest antennal enlargement. Thorax Fig. 1 Structurally, the thorax is characterized by the total lack of the pleural arch and an extreme degree of fusion in the metathorac- ic segments. The pronotal comb is absent, and setae are reduced both in number and size. The prothorax, aside from the absence of the pronotal comb, is not remarkable. The fused pleurosternal plate is not divided into the proepisternum and proepimere by a ridge as in Hystrichopsylla. The ventral margin of this plate is straight with the dor- sal margin sinuously downcurving caudally; thus, the plate is broader anteriorly than posteriorly. The caudal margin of the free flange of the pleurostemum is relatively straight except for a small, sharp, median triangular projection. The pronotum is a simple, collarlike sclerite. There is considerable disagreement among authors concerning the structure of the sternal and pleural areas of the me.so- and metathorax and of the lateral notal and episternal areas of the metathorax. Snod- grass (1946) considered the sternum to con- sist of a narrow strip lying between the coxae and fused with the epistemum ante- riorly, leaving the posterior portion of the sternum as a free arm. Jordan (quoted in Hopkins and Rothschild 1953) considered the sternum to have lateral elements fused to the epistemum. This divergence in view- point confuses terminology: the epistemum of Snodgrass is considered the sternum (at least in part) by Jordan; the lateral meta- notal area of Snodgrass is considered the epistemum by Jordan. Johnson (1957) follows the interpretation of Snodgrass, whose terminology is used here, and presents additional evidence for its support. She points out that the ventral margin of the epistemum in both the meso- and metathorax ends in a free flange and that the sterna are entirely separate from the epistema except where fused anteriorly. The free flanges of the epistema appear not only to extend below the sterna, but also to overlap a portion of the coxae. The mesothorax of Anomiopstjlhis is rela- tively immodified (Fig. 1). The notum is simple with the anterior phragma large and prominent in slide-mounted specimens. It is marked by the complete loss of internal pseudosetae beneath the mesonotal collar. These pseudosetae are found in other ano- miopsyllines and differentiate the Cerato- phylloidea from Pulicoidea. The mesepister- num and mesepimere are separated by a strong vertical pleural rod, firmly attached at both ends. This rod supports the pleural articulating sclerites of the coxa. The ven- tral margin of the mesepisternum curves downward and anteriorly from the base of the pleural rod, forming a free flange slightly overlapping the coxa and fusing an- teriorly with the sternum. The ventral mar- gin of the mesepimere is relatively straight. The mesepimere forms a free caudal flange, slightly overlapping the mete- pistemum near the point of attachment of the third link-plate or vinculum. Dorsally, it slightly overlaps the ventral edge of the me- sonotum except at the anteriormost corner. The metathorax of Anomiopsyllus (and all other Anomiopsyllinae) is marked by ex- treme fusion and a rather radical dis- placement of parts. The net effect is ante- rior fusion of the notum with the June 1977 Barnes et al.: Anomiopsyllus Fleas 143 Figs. 2-3. Anomiopsyllus amphibolus: 2, male terminalia; 3, female terminali; 144 Great Basin Naturalist Vol. 37, No. 2 metepisternum, loss of the lateral metanotal area, and loss of the pleural arch. These complex phenomena have been subject to some discussion by taxonomists, particularly Johnson (1957), and warrant further dis- cussion and clarification, especially with re- gard to the mechanics involved, evolution- ary trends, and possible adaptive significance. In order to imderstand what has taken place in the metathorax, it is necessary to review the work of Snodgrass (1946), espe- cially with regard to the strengthening rods and lateral metanotal area. In most fleas, the metanotum is a collar- shaped structure. It normally bears two transverse strengthening ridges, the anterior prephragmal ridge and the more posterior notal ridge. Near the lower end, the trans- verse notal ridge gives off anteriorly a short longitudinal ridge which usually continues to the anterior edge of the notum where it blends into a marginal thickening continued around the ventral edge of the notal plate. The accessory ridge thus sets off a small, variously shaped ventrolateral area of the metanotum, termed the lateral metanotal area (see Snodgrass, PI. 8., Figs. K-0; PI. 9, Figs. A-E). This area normally bears a small but distinct, free marginal flange, but in Pu- lex it is fused with the upper margin of the metepisternum. Contrary to the findings re- ported by Johnson (1957), Pulex still retains the lateral metanotal area: though fused with the metepisternum, it is still set off by the ventrolateral thickening extending from the lower anterior margin to the ventral ex- tension of the notal ridge. The notal flange normally extends into the pleural area, where it terminates in an oval expansion that overlaps and clasps the upper end of the pleural rod, thus forming the so-called pleural arch (see Snodgrass, 1946: 26, PI. 9, Figs. C, H-K). The union is a very firm one. Snodgrass states (p. 26): "It is evident, therefore, that this apparent notopleural 'articulation' of the metathorax is for the purpose of solid union along the line of the notal and pleural ridge, and is not for controlled movement. The flexibility of the united ridges allows the pleuron to be twisted somewhat on the notum, but per- mits of no swinging movement in a longitu- dinal plane." According to Snodgrass, Opisodasys and Dactylopsylla Jordan possess a pronounced posterior angulation between the notal flange and the axis of the pleural ridge. This condition applies in Orchopeas Jordan and most ceratophyllids. In these fleas, the pleural ridge has moved anteriorly to the line of the notal flange and is fused strongly to its angled ventral extension. In Anomiopsyllus and allies, much more radical changes have taken place. In Ano- miopsyllus, Megarthroglossus, Callistopsyllus, and Stenistomera, the notal ridge is strongly reduced in thickness and extent and the lower portion has disappeared above the notal margin. The accessory ridge is still present; its identity is established by the at- tachment of the link-plate or vinculum at its anterior end (Fig. 1, VC. II). This ridge is strongly developed anteriorly, but fades posteriorly to varying degrees in the ano- miopsyllines examined, and does not meet the ventral remnant of the notal ridge. The ventral extension of the notal ridge is ap- parently gone. The long metapleural rod has been displaced anteriorly and is appar- ently completely fused with the accessory ridge. There is no sign of a pleural arch. Along with this, the metepisternum and me- tanotum are completely fused, and in Ano- miopsyllus there is no vestige of the lateral metanotal area. A clue to what has happened is present in Megarthroglossus, and perhaps in Ste- nistomera and Callistopsyllus. In Megarth- roglossus a small thickening leads anteriorly from the upper portion of the pleural rod, but ends before it reaches the anterior mar- gin of the segment. Its position suggests that it represents the vestigial ventral thickening of the lateral metanotal area. A similar thickening exists in the other genera, but leads posteriorly from the anterior margin. In Stenistomera the thickening is short and knoblike and in Callistopsyllus it is longer. If the structure in Megarthroglossus is in- deed, as suggested, part of the "pleural rod," it must represent the ventral portion of the notal flange, which has completely fused with the pleural rod. The notal flange June 1977 Barnes et al.: Anomiopsyllus Fleas 145 4-5. Anomiopsyllus amphibolus: 4, metathoracic leg; 5, aedeagus. 146 Great Basin Naturalist Vol. 37, No. 2 has lost its direct connection with the notal ridge proper and has been displaced ante- riorly with the pleural rod. This would be correct, since the ventral thickening of the lateral metanotal area always connects with the ventral extension of the notal ridge in those fleas that possess the lateral metanotal area. In the ceratophyllid genus, Orchopeas, and perhaps others, we see a forward ang- ulation of the ventral end of the notal ridge and an anterior displacement of the pleural rod, but the elements essential to the pleur- al arch are present and highly developed. In Anorniopsyllus and allies these structures have been entirely lost or have lost identity by fusion. Johnson (1957:8) has offered an explana- tion of the adaptive significance of the loss of the lateral metanotal area and the pleur- al arch. According to her, the pleural arch serves as a strengthening device which acts as a "cushion" in jumping. Since Anorniop- syllus is a "nest flea" and consequently a "poor jumper," the pleural arch is no long- er necessary. She explains the means by which it was lost as ". . . probably loss due to lack of selection against any mutation tending toward reduction of the pleural arch which, if my premise is correct, is mainly an aid in jumping." The negative view that such a radical change involving major structures has arisen by mutations in the absence of selection pressures is difficult to accept. Such phe- nomena undoubtedly occur in isolated island populations where the gene pool is small and selection pressure slight, but they must be exceedingly rare in any continental group. In a rigorous continental environ- ment, one would not expect mutations to survive in the absence of some positive se- lective advantage to the mutant. One must postulate that any such radical change would have to offer a positive selective ad- vantage to be retained. Losses which may take place by mutation in the absence of se- lection could be termed degenerative evolu- tion. Thus, Anorniopsyllus may be dubbed a "degenerative form," a term disapproved of by Johnson (1957:16). The possible selective advantage of the anterior displacement and fusion of strengthening ridges in the metathorax of anomiopsyllines may be partially clarified by observations of adult fleas in the labora- tory imder both artificial and near-natural conditions. Anorniopsyllus is not a weak jumper. It does not jump at all! Callistop- syllus, Megarthroglossus, and Stenistomera are weak jumpers. This is probably due in part to correlated differences in meso- and metathoracic coxae as well as to changes in the metathorax. This becomes apparent with microscopic observations of flea move- ments through tangles of nest material. The abdomen of Anorniopsyllus is capable of sweeping movement in the lateral plane as the flea moves through nest material with a rapid, sinuous movement. Lateral flexibility is not observable in the more rigidly con- structed Hystrichopsylla and Atyphloceras Jordan and Rothschild. Anterior dis- placement of thoracic ridges and reduction of the notal ridge offer a considerably greater degree of flexibility in the metatho- rax. Legs Fig. 4 The legs of Anorniopsyllus are marked by prominent posteroapical sinuses on both meso- and metacoxae, above which are large subacuminate spurs (Figs. 1, SPUR). The prolegs are not distinctive. The meso- coxae possess a complete longitudinal break (Fig. 1, BR.), considered to be an advanced condition. A complete break is also found in Megarthroglossus, but not in Callistopsyllus or Stenistomera. This condition also is found among the Rhadinopsyllinae. In an anterior view, the mesocoxae are seen to fold inward at the break, forming a shield similar to that formed by the procoxae, as described by Snodgrass (1946). The shield is formed by pivoting the procoxae at the pleurocoxal articulations. Snodgrass postulated that the "shield" formed by the procoxae may be useful for forcing between hairs or may be used as a flat base for feeding. The second "shield" formed by the mesocoxae in Ano- rniopsyllus and Megarthroglossus suggests the former explanation and perhaps is needed to cope with the greater hetero- June 1977 Barnes et al.: Anomiopsyllus Fleas 147 geneity and density of nest material rather than hairs on the host. In Anomiopsyllus and other anomiopsyl- hnes, the exterior coxal ridge in both meso- and metacoxae is incomplete. In those fleas examined by Snodgrass (1946), including Pu- lex Linnaeus, Hystrichopsylla, Dactylopsylla, and Opisodasys, and in most other fleas, the ridge is complete and reaches the apex of the coxae. The meso- and metacoxae are ar- ticulated mesally to the sternum by a pair of prominent articulating sclerites (Fig. 1, ex. A). The upper mesal coxal surface is membranous, and the articulating sclerites serve as the only rigid connection between sternum and coxae. These sclerites normally articulate with a well-defined socket in the dorsal edge of the mesal sclerotized surface of the coxa. At the point of articulation, the coxa is strengthened by a horizontal thick- ening and a longitudinal rod extending to- ward the coxal apex (see Snodgrass 1946) and may or may not reach the apex of the coxa. This structure has been termed the mesal coxal ridge by Snodgrass. In Ano- miopsyllus, there is no hint of thickening at the point of articulation and the rod is en- tirely absent. Both thickening and rod are present in the related genera Callistopsyllus, Megarthroglossus, and Stenistomera. The reduction of the coxal ridge, absence of mesocoxal ridge, and complete mesocoxal break in Anomiopsyllus would seem to be further examples of specialization to the nest environment and to walking rather than jumping. Abdomen The unmodified abdominal segments (I- VI) are not remarkable except for the char- acteristic reduction in number and size of setae. Terga I and II each bear spinelets at the apical margin, the former usually 2 per side, the latter 1. Each tergum bears a row of thin setae, 3 to 5 on each side, plus 1 very minute guard seta below and anterior to each spiracle. Sterna II to VI are simple, usually bearing two setae ventrally, the up- per one usually the longer. Modified abdominal segments of MALES (Fig. 3): Segment VII is the first modified segment. The tergum bears one long antepygidial seta which is not reduced or altered in shape. Sternum VII is not modified differently from preceding sternal segments. Sternum VIII is expanded posteri- orly and dorsally to serve as the chief ven- trolateral shield about the genitalia and differs markedly from the more anterior segments. The tergum is marked at the pos- terior margin by rather prominent seti- gerous grooves containing the spiracles be- low which it expands caudally to serve as the main dorsolateral shield for the gen- italia. It is comparatively shallow, descend- ing ventrally only about half the distance of the preceding terga. Sternum VIII closely resembles that of the other Anomiopsyllinae by being expanded and ensheathing much of the proximal portion of sternum IX. Segment IX of males (Fig. 3) is exceed- ingly modified; included in its structure is the clasping apparatus of the external gen- italia. Its tergum is much reduced dorsally and consists of a narrow but complete strap anterior to the pygidium (Figs. 3-4, T. IX). The tergum is continuous with the upper anterior angle of the clasper lobes, very much as noted for Hystrichopsylla by Snod- grass (1946). Immediately below and ante- rior to the tergal strap is a small but strong tergal apodeme, reinforced by thickened bars that join the anterodorsal angles of the large manubria (Figs. 3, MN). The apodeme forms a strong dorsal arch between the manubria at this point from the anterior edge of the dorsal arch. The anterior mar- gin descends downward into the abdominal cavity, then curves dorsally and posteriorly to join the clasper lobe, thus forming a prominent triangular structure. The tergal clasping apparatus consists of the broadened clasper lobe composed of a more or less dome-shaped fixed process (P) and an articulated movable process (F). The fixed process is usually equipped at or near the apex with a long, heavy bristle proj- ecting over the apex of the movable process (lacking in A. novomexicanensis). Invariably, the movable process or "finger" is equipped with heavily pigmented, large spiniform setae on the mesal surface. The number and position of these setae have been used ex- 148 Great Basin Naturalist Vol. 37, No. 2 tensively in taxonomy. The posteroventral apex of the clasper lobe below the articu- lation of the movable process bears the ace- tabular seta. The articulation of the mov- able process is on the inner surface of the clasper lobe. The socket may be oval or nearly round (as in nudatus, novomexi- canensis and falsicalifornicus groups) or linear and nearly vertical (as in the amphi- bolus group). Sternum IX (Fig. 3) consists of two main components, the proximal and distal arms. Together, in lateral view, they form a prominent V-shaped structure easily seen in slide-mounted specimens. The proximal arms are platelike sclerites, character- istically shaped in some species but variable enough that taxonomic value is limited. In Anomiopsijlhis, they extend dorsad and an- terad almost to the anterodorsal angle of the manubrium. The distal arms form a single troughlike structure from the base to a point 20 to 25 percent of the distance from the apex, after which they separate and are free. The heavily reinforced lateral portions are interconnected by a more light- ly sclerotized, distally convex median plate variously adorned by setae. The pattern is characteristic of the species. The apices of the free arms are equipped with large, tack- like setae. Their number and size are of tax- onomic value. In some species, there is a lobe which forms a narrow sheath around the angle of the distal arm about three- fourths of its length. In others, this structure is greatly reduced and difficult to see. Aedeagus (Fig. 5): The aedeagus of Ano- miopsyllus is simple and basically very sim- ilar to that of Hystrichopsylla dippiei Rothschild, as described by Snodgrass (1946) and Traub (1950). It is long and narrow. The apodeme is broadly scimitar shaped with a prominent basal constriction or neck. The endchamber is a simple hood opened apically and ventrally. The crochets (CR) are reduced and immovable in the ventrola- teral wall. The aedeagal pouch is deep, ex- tending considerably cephalad of the struts. The sclerotized inner tube (S.I.T.) is promi- nent, tubular, and funnel shaped at the base with a prominent anteroventral spur to which is attached the apodemal rod. The crescent sclerite (C.S.) of Anomiopsyllus is prominent and attached to the anterodorsal flange of the tube. Penis rods are long and coiled. There are no specialized lobes pres- ent, indicating its primitiveness as compared to ceratophyllids and pulicids. Modified abdominal segments of FEMALES (Fig. 2): Tergum VII bears the single, long antepygidial bristle. Sternum VII is marked by a posterior expansion with its margin shaped variously in the several species. The upper margin of sternum VII, as pointed out by Snodgrass (1946), overlaps its tergum (Fig. 2, St. VII). Segment VIII is exceedingly modified. The tergum consists of two lateral plates narrowly joined dorsally, expanding ven- trally and posteriorly to cover most of the genitalia. Its posterior margin near the dor- sum contains the deep setigerous grooves of the most posterior spiracle. From that point, it expands posteriorly. The most pos- terior margin may be straight or lobed (Fig. 2, T VIII), depending on the species. Its usefulness as a taxonomic character is re- duced by intraspecific variation. Sternum VIII, as in other fleas, is reduced to a small plate between the two valves of its tergum and functions only as the ventral floor be- low the genitalia. Tergum VIII completely encloses the ex- ternal genital chamber and is guarded at its orifice by a group of very heavy, inward-di- rected mesal setae. The lateral walls of the genital chamber are heavily sclerotized in Anomiopsyllus, apparently forming a chan- nel. The bursa copulatrix is a simple sack at or near the anterior end of the "channel." The bursa, in some species, (notably no- vomexicanensis Williams and Hoff, Fig. 90) has minute areas of sclerotization, and in others it appears as a wholly membranous sack. The spermatheca is simple, with a more or less globular bulga (head) and a curving, broad hilla (tail) without a terminal appendage (Fig. 2). Segment IX in Anomiopsyllus is marked by well-developed tergal plates. As pointed out by Snodgrass (1946: 36), segment IX of most female fleas is virtually obliterated. Those groups retaining well-developed ter- gal plates on this segment include: Hys- 1977 Barnes et al.: Anomiopsyllus Fleas 149 Figs. 6-9. Finger and clasper: 6, Anomiopsyllus amphibolus; 7, A. nidiophilus; 8, A. montanus; 9, A. sinuatus. 150 Great Basin Naturalist Vol. 37, No. 2 trichopsylla (Snodgrass 1946); Ctenop- thahnus Kolenati, 1956; Parapsylla (Wagner, 1939); and Atyphloceras Jordan and Rothschild, 1915. Anomiopsyllus is sug- gestive of the more primitive hys- trichopsyllid genera in the retention of a well-developed tergum IX. These plates consist of teardrop-shaped valves on either side of the pygidium, well rounded dorsally, ventrally, and anteriorly but acute posteri- orly with points projecting beyond the anus, below the anal stylets, and outside the anal plate (Fig. 2, 3, T. IX). From the dorsal as- pect, the plates appear to be joined ante- riorly only by a membranous connective and are completely separated from tergum X by a sutvire. Segment X contains the prominent pygi- dium, the proctiger, and the anus. The py- gidium is framed by a narrow, strongly sclerotized ring somewhat open anteriorly but joined by a small plate. It bears 20 (10 per side) sensillial pits (Fig. 2, SN.), though 21 were noted on one specimen of A. /. congniens from Placerville, California. The number of sensillia is used as a key charac- ter by Hopkins and Rothschild (1956) to separate Pulicoidea from Ceratophylloidea. According to them, pulicoids bear 8 to 14 sensillial pits per side and ceratophyl- loids sometimes bear 14 but usually bear 16 or more. The reduced number of these pits in Anomiopsyllus, a definite ceratophylloid in other respects, is interpreted here as an example of convergence through loss and does not indicate pulicoid affinities. The proctiger (Fig. 2, DAL.) is prominent and well endowed with strong setae. The anal stylets are simple and equipped with one long terminal seta and one or two minute subterminal setae. Discussion The skeletal anatomy of Anomiopsyllus is most marked for its extreme reduction of parts and small size. This reduction has been accompanied in Anomiopsyllus by the loss of a number of behavioral character- istics common to fleas, particularly the abil- ity to jump. Undoubtedly these reductions are adaptations to its role as a nest flea. with Anomiopsyllus progressing further than any other flea group. The loss of the pleural arch and the moving forward of the pleural rod reduce the mobility of Anomiopsyllus as far as jumping is concerned, but probably enable the flea to move through hetero- geneous nest material at a more rapid rate. Modifications of the mesocoxae probably also serve this purpose. With respect to both male and female modified abdominal segments, Anomiop- syllus has retained the basic structure of its more primitive relatives in the Hys- trichopsyllidae. The aedeagus, for example is far less complex than that of any cerato- phyllid or pulicid, and less complex than that of the other Anomiopsyllinae, including Callistopsyllus and Megarthroglossus, to which it is most closely related. Never- theless, the relationships of the aedeagus of Anomiopsyllus to those of its hys- trichopsyllid relatives are made clear by comparison with that of Hystrichopsylla. In the female, retention of the ninth tergal plates further links Anomiopsyllus to its more primitive hystrichopsyllid relatives. The reduction in the total number of sensi- llial pits on the pygidium represents special- ization, probably accompanying overall re- duction in size. Characters Used in the Study The differences between Anomiopsyllus species and, to an even greater extent, be- tween subspecies are found almost entirely in the male external genitalia. Few tenable characters are available in females. For the most part, differences in configuration of sternum VII of females serve only to sepa- rate species groups. Configuration of the posterior margin of tergum VII used by Hopkins and Rothschild (1962) to separate species in their key does not hold when long series are studied, though trends may exist in any given population. In males, particular attention has been given structures of the ninth abdominal seg- ment, which is modified to form the clas- ping apparatus of the external genitalia. The shapes of the movable and immovable processes are somewhat variable within spe- June 1977 Barnes et al.: Anomiopsyllus Fleas 151 cies, but are nevertheless stable enough to constitute good species characters. Con- sistent differences in these structures also serve as species group characters as does the structure of the articulation of the mov- able process. Arrangement and number of setae of both the movable and immovable processes of the clasper serve as species and subspecies characters. In some of the species one of the prominent spiniforms of the movable process seems to be in the process of being lost and is absent in some speci- mens. Differences in the shape and setation of the distal arm of sternum IX are useful at the subspecific level and more so at the specific level. The general shape of the pro- ximal arms of sternum IX is useful as a sec- ondary character at the species level, but varies a great deal in detail and is not us- able in determining subspecies. Among sev- eral subspecies the distance between promi- nent spiniform setae has proven to correlate well with other characters, including geog- raphic distribution. The aedeagus varies less than do other structures of the genitalia; ne- vetheless, some variation does occur in con- figuration of the dorsal margin and in di- mensions. These are of some use at the species level. List of Abbreviations A. A.F. ae.a. A.B. A.R. A.S. AB.S. A.T.I B.C. C.S. CX.A. ex. I ex. II ex. Ill DA. D.A.L. (T.X.) D.S. F.L.L. F.M.L. FO.S. F.S. Antennae Antennal fossa Aedeagus apodeme Antepygidial bristles Apodemal rod Anal stylet Abdominal spinelets Abdominal tergum I Bursa copulatrix Crescent sclerite Coxal anticulating rod Coxa I (Pro) Coxa II (Meso) Coxa III (Meta) Distal arm of Sternum IX Dorsal anal lobe (Tergum X) Duct of spermatheca (Ductus seminalis) Finger (movable process) of clasper Fulcral lateroventral lobe Fulcral median lobe Ford sclerite Fossa spiracular FT. Frontal tubercle C.L. Genal lobe L.P. Labial palp M.D.L. Median distal lobe of aedeagus MSN. Mesonotum MTN. Metenotum M.L. Maxillary lacinia MN. Manubrium MP. Maxillary palp MPM. Mesepimeron MTM. Metepimere M. Maxilla P. Process (immovable) of clasper P.A. Proximal arm of Sternum IX PHR. Phragma PHY. Pharnyx PN. Pronotum PEN. Penis P.R. Penis rods PL.R. Pleural rod PSP. Pleurostemal plate SP. Spermatheca SN. Sensillium S.I.T. Scleritized inner tube SPIN. Spiniform S.S. Satellite sclerite ST. VII Sternum VII ST. VIII Sternum VIII ST. IX Sternum IX T. VII Tergum VII T. VIII Tergum VIII T. IX Tergum IX V. Vesicle V.A.L. (St. X) Ventral anal lobe (Sternum X) ve. I Vinculum I (Link-plate I) VC. II Vinculum II (Link-plate II) V.D. Virga dorsalis Y.S. "Y" sclerite Systematic Section Anomiopsyllus Baker Anomiopsyllus Baker, 1904, Proc. U.S. Nat. Mus. 27: .377 (Type Species: Typhlopsylla nudata Baker, 1898.); Oudemans, 1909, Nov. Zool. 16: 157; Jordan and Rothschild, 1915, Ectoparasites 1: 45; Dalla Torre, 1924, Natur. Med. Ver. Innsbruck 39: 17; Ewing, 1929, Manual of Ectoparasites: 162; Jellison and Good, 1942, Index to the literature of Siphonaptera of North Amer- ica: 21; Ewing and I. Fox, 1943, U.S. Dept. Agric. Misc. Publ. 500; Costa Lima and Hathaway, 1946, Monogr. Inst. Oswaldo Cruz 4: 522 pp.; Hubbard, 1947, Fleas of western North America: 56, 78; Hol- land, 1949, Siphonaptera of Canada: 96; Eads and Menzies, 1950, Fleas of Texas; Anon., 1950 (a & b). Public Hlth. Repts. 65: 454, 526; Traub and Tipton, 1951, J. Wash. Acad. Sci. 41: 267-268; Wiseman, 1955, Univ. Wvo. Publ. 19: 28 pp.; Finley, 1958, Univ. Kans. Publ. Mus. Nat. Hist. 10: 21.3-522; Parker and Howell, 152 Great Basin Naturalist Vol. 37, No. 2 1959, J. Parasitol. 45(6): 507-604; Stark, 1959, Si- phonaptera of Utah; Vargas, 1960 Med. Rev. Mexicana 40 (849): 1; Kartman et al., 1960, Zoon Res. 1(1): 178; Hopkins and Rothschild, 1962, Catalogue of the Rothschild Collection of Fleas III: 392; Holland, 1965, Canadian Ent. 97: 1051-1058; Jellison and Glesne, 1967, Index to the literature of Siphonaptera of North Amer- ica, Suppl. 2: 13-19; Tipton and Mendez, 1968, Pacific Insects 10(1): 179-182; Beck and Allred, 1966, Brigham Young Univ. Sci. Bull., Biol. Ser. 7(2): 13; Stark and Kinney, 1969, J. Med. Ent. 6(3): 290; Pratt and Stark, 1973, DHEW Publication 74-8267: 11; Lewis, 1974, J. Med. Ent. 11(2): 154. Anomiopsylla (sic): Augustson and Durham, 1961, Bull. South. Calif. Acad. Sci 60(2): 100. Anomiopsyllus is immediately separable from other Anomiopsyllinae by extreme re- duction in number and size of setae, lack of a pleural arch, and absence of a pronotal comb and from Eopsylla by the lack of in- ternal mesonotal setae (Hopkins and Rothschild 1962). Head.— Rounded anteriorly, clypeal tu- bercle present, prominent; eyes absent; trabecula centralis absent; labial and max- illary palpi 4-segmented, reaching to or slightly beyond protrochanter; terminal seg- ment of labial palpi not markedly asymmet- rical nor greatly longer than other seg- ments; maxillae acutely triangular, somewhat convex anteriorly; concave poste- riorly, genal lobe broadly roimded, short. Antennae of male much enlarged, antennal fossae of males beginning almost at dorsal margin of head, connected dorsally by a falx; second antennal segment not over- lapping base of third; antennal setation sparse, short, setae of female second anten- nal segment only one-fourth length of clava. Frons with one row of setae; occiput with a row of minute setae; antennal fossa with several setae along posterior margin. Thorax.— Pronotal comb absent, pseudo- setae absent. Pleural arch absent. Mete- pimeron partially fused with metanotum, latter with a small, free posteroventral flange. Metepistemum completely fused with metepimeron and anterior portion of metastemum. Posterior margin of mesocoxa and metacoxa deeply emarginate pre- apically, forming a deep sinus and sharp preapical spur, mesocoxa with a prominent longitudinal break from anterior base to preapical po.sterior sinus. Coxae, femora, and tibiae with few or no lateral setae, pro- coxae with row of long, thin setae on poste- rior margin. Femora and tibiae with weak posterior marginal setae, meso- and metafe- mora and tibiae with few very weak ante- rior marginal setae. Tarsi with four lateral pairs of plantar bristles, the basal pair moved ventrally on the pro- and mesotarsi. Abdomen.— Terga with one row of weak setae; tergum I with two to four apical spinelets; tergum II with one or two. Ab- dominal sterna each with two slender, ven- tral submarginal setae. One antepygidial bristle in both sexes. Female.— Tergum VIII posterior margin sloping posteroventrally, then ventrally to enclose the genitalia. Genital chamber marked by heavily pigmented area on ter- gum VIII, its gonotreme protected by four to six heavy, distomesad setae, forming an "ovipositor." Tergum IX large, broadly curving laterad around pygidium and pro- ctiger, tapering to an acute point post- erolaterad near anal stylet. Anal stylet ap- proximately 2.0 to 2.5 times as long as broad, with long, slender, curving terminal seta and one minute preapical, subdorsal seta. Anal plate broad at base, dorsal mar- gin linear, ventral margin gently curving to point. Male.— Tergum IX narrow but entire dorsally with a well-developed tergal apo- deme forming clasper lobe laterally. Clasper and manubrium roughly triangular, descend- ing deeply into abdominal cavity; fixed process well developed; movable process variable, always with two to four heavy, pigmented, distomesad spiniform setae. Ster- num IX V-shaped, proximal arm extending anterodorsally into body cavity, distal arm extending posterodorsally to form a post- eroventral sheath about external genitalia, fitted distally with two free lobes armed at subapical posterior margin with modified setae. Aedeagus large, apodeme broadly scimitar shaped, bearing a weak terminal appendage. Apodemal neck present and well developed. Endchamber without well- developed, specialized lobes or with none, covered by a simple, weakly sclerotized hood, open apically and ventrally, extending from apex cephalad to a point anterior to June 1977 Barnes et al.: Anomiopsyllus Fleas 153 Figs. 10-12. Finger and clasper: 10, A. nudatus hiemalis; 11, A. nudatus nudatus; 12, A. nudatus mextcanus. 154 Great Basin Naturalist Vol. 37, No. 2 aedeagal struts, then curving dorsally and anteriorly as lateral lobes of aedeagal apo- deme. Aedeagal pouch arising below trough, extending distally and dorsally toward base of sclerotized inner tube, latter strongly sclerotized, broad at base, ventral margin curving ventrally and cephally; narrowing to a well-developed tube posterally, apex without auxiliary sclerites. Crescent sclerite (C.S.) present. Crochets represented by very small, internal, unarticulated plates at ven- tral apex of endchamber. Penis rods long, coiled. Aedeagal apodemal rods present, well sclerotized. Ventral intramural rod present. The species, though closely related, may be separated into five species groups based on characters in the male genitalia. The characters used in the grouping are: (1) number of spiniforms on finger of clasper and (2) position of spiniforms on distal or proximal portion of finger curve in either vertical or oblique rows. The grouping, though artificial, may give an indication as to the phylogeny of this genus. The five groups are: 1. Spiniforms: 1 distal, 1 proximal: am- phibolus, montanus, and sinuatus. 2. Spiniforms: 2 distal, 1 proximal: nu- datus nudatus, n. mexicanus, and n. hiemalis. 3. Spiniforms: 3 distal, 1 proximal: (verti- cal row) martini and walkeri; (oblique row) falsicalifornicus falsicalifornicus and /. congruens (f. fahicalifornicus has apparently lost the proximal spin- iform in its evolution). 4. Spiniforms: 3 distal: novomexicanensis, traubi, oaxacae, and durangoensis (in durangoensis one of the distal spin- iforms has apparently been lost in its evolution). 5. Spiniforms: 1 distal, 3 proximal: nidio- philus. The most widely distributed group of this genus is represented by A. amphibolus and A. montanus in the north and A. sinuatus in the south. The northern species are most closely related to the genus Callistopsyllus. The nudatus group, represented by three subspecies, is distributed primarily on the Colorado Plateau in the Sonoran Desert, which extends into Mexico. The distribution of the fakicalifornicus group is coastal or subcoastal in the Pacific region of North America. The distribution of nidiophilus, traubi, and oaxacae is limited to Mexico. Anomiopsyllus princei has been collected only from New Mexico and martini only from Mexico; however, both of these spe- cies have been collected only from Sdurus aberti, suggesting that they could be the same species. Further collecting is needed to determine their relationship. Key to Male Anomiopsyllus 1. Finger of clasper with both distal and proximal spiniforms 2 Finger of clasper with only distal spiniforms 11 2( 1). Finger with 1 distal and 3 proximal spiniforms nidiophilus Finger with 1 or more distal but only 1 proximal spiniform 3 3( 2). Finger with 1 distal and 1 proximal spiniform 4 Finger with 2 or 3 distal and 1 proximal spiniform 6 4( 3). Finger triangular shaped (Fig. 7) amphibolus Finger boomerang shaped (Fig. 8,9) 5 5( 4). Finger broad; 3 heavy bristles on apex of St. IX (Fig. 33) sinuatus Finger narrow; 4 heavy bristles on apex of St. IX (Fig. 32) montanus 6( 3). Finger with 2 distal and 1 proximal spiniform 7 Finger with 3 distal and 1 proximal spiniform 9 7( 6). Distance between distal spiniforms under 20u nudatus nudatus Distance between distal spiniforms over 20u 8 June 1977 Barnes et al.: Anomiopsyllus Fleas 155 8( 7). Thin bristle at angle of divergence of St. IX (Fig. 35) nudatus hiemalis Thick bristle at angle of divergence of St. IX (Fig. 36) nudatus mexicanus 9( 6). Finger with distal spiniforms in oblique row in relation to posterior margin (Fig. 13) fahicalifornicus congruens Finger with distal spiniforms in vertical row in relation to posterior margin 10 10( 9). Angle of divergence of St. IX with prominent nob at angle; 1 or 2 thick bristles (Fig. 38) walkeri Angle of divergence without nob; 2 thick bristles (Fig. 39) martini 11( 1). Finger with 2 distal spiniforms (Fig. 26) durangoensis Finger with 3 distal spiniforms (Fig. 27) 12 12(11). Finger with oblique row of spiniforms (Fig. 13) fahicalifornicus fahicalifornicus Finger with vertical row of spiniforms (Fig. 27) 13 13(12). Process of clasper with long bristle (Fig. 27) 14 Process of clasper without long bristle (Fig. 26) novomexicanensis 14(13). Apex of finger distinctly acuminate (Fig. 28); posterior margin of St. IX with 1 thin, submarginal bristle trauhi Apex of finger rounded (Fig. 29); posterior margin with large number of evenly spaced bristles extending from heavy apical bristles, about % down the distal arm of St. IX oaxacae Anomiopsyllus nidiophilus Tipton and Mendez Figs. 7, 32, 51, 67, 78, 95 Anomiopsyllus nidiophilus Tipton and Mendez, 1968, Pacific Insects 10(1): 179.182; Lewis, 1974, J. Med. Ent. 11(2): 154. Diagnosis: Anomiopsyllus nidiophilus, though similar to A. martini, is the only species in the genus with one distal and three proximal spiniforms on the movable process of the clasper. Sternum VIII differs from that of all other species, except A. si- nuatus, by having a sinus in the caudodorsal margin. The distal arm of sternum IX has a subapical constriction bearing three short, thick setae. Female characters are not sufficiently dis- tinctive to separate nidiophilus from mar- tini. Type locality: Cerro Potosi, Nuevo Leon, Mexico, 3350 m. Type host: Nest of wood rat (probably Neotoma albigula leucodon). Type specimens: Male holotype and fe- male allotype, 38 male and 156 female paratypes, same data; 6 males and 6 females from Neotoma albigula leucodon, same lo- cality as above but with dates and eleva- tions as follows: 5 males and 2 females on 21 April 1964, 3450 m; 1 male and 2 fe- males on 23 April 1964, 3030 m; 1 female on 11 September 1964, 3140 m; 1 female on 23 September 1964, 2990 m. Holotype and allotype in U.S. National Museum; paratypes in British Museum (Natur. Hist.), Canadian National Collection, Rocky Moun- tain Laboratory at Hamilton, Montana; Es- cuela Nacional de Cienias Biologicas (I.P.N.), Mexico, D. F.; Brigham Young Uni- versity, Provo, Utah; Gorgas Memorial Lab- oratory, Panama; Bishop Museum and col- lections of R. Traub and V. J. Tipton. Discussion: The distribution of Neotoma albigula leucodon is limited to the states of Coahuila, Jalisco, Mexico, Nuevo Leon, San Luis Potosi, Tamaulipas, and Zacatecas, but siibspecies of Neotoma albigula occur in west Texas, New Mexico, Arizona, and sev- eral states in Mexico and extend northward into the southernmost areas of Colorado and Utah. Further collections are needed to de- fine more clearly the distribution of this species in Mexico and possibly the south- western U.S. 156 Great Basin Naturalist Vol. 37, No. 2 Anomiopsyllus amphibolus Wagner Figs. 1-6, 30, 52, 80, 95 Anomiopsyllus nudatus Stanford, 1931 (nee Baker, 1898), Proc. Utah Acad. Sci. 8: 153 {fide Jellison and Good (1942: 21)) Anomiopsyllus amphibolus Wagner, 1936, Zeit fiir Parasitol. 8: 654-655; Jellison and Good, 1942, Index to the literature of Siphonaptera of North America: 21- 24; Ewing and I. Fox, 1943, U.S. Dept. Agric. Misc. Publ. 500: 114; Stanford, 1943, Proc. Utah Acad. Sci., Arts, and Lett. 19-20: 173; Costa Lima and Hathaway, 1946, Monogr. Inst. Oswaldo Cruz 4: 127; Hubbard, 1947, Bull. S. Calif. Acad. Sci. 48(3): 116; Tipton, 1950, Great Basin Nat. 10: 63; Beck et al., 1953, Proc. Utah Acad. Sci., Arts, and Lett. 30: 50; Jellison, Locker, and Bacon, 1953, Index to the literature of Siphonaptera of North America, Suppl. 1: 10; Howell, 1957, J. Para- sitol. 43(5): 40, 42; Parker and Howell, 1959, J. Para- sitol. 45(6): 507-604. Stark, 1959, Siphonaptera of Utah: 105-106; Holland, 1965, Canadian Ent. 97: 1053; Beck and Allred, 1966, Brigham Young Univ. Sci. Bull, Biol. Ser. 7(2): 13; Jellison and Glesne, 1967, In- dex to the literature of Siphonaptera of North Amer- ica, Suppl. 2: 13-19; Allred, 1968, Great Basin Nat. 28(2): 75; Tipton and Saunders, 1971, Brigham Young Univ. Sci. Bull., Biol. Ser. 15(2): 18; Lewis, 1974, J. Med. Ent. 11(2): 154. Anomiopsylla amphibolus (sic): Howell, 1955, Great Basin Nat. 15: 40, 42; Augustson and Durham, 1961, Bull. S. Calif. Acad. Sci. 60(2): 100. Diagnosis: Anomiopsyllus amphibolus males may be distinguished from other Ano- miopsyllus species, except A. montanus and A. sinuatus, by having only two widely spaced spiniforms near the posterior margin of the movable process and a single row of four very strong, heavy setae on the apical posterior margin of sternum IX, and from A. montanus and A. sinuatus by the triangular-shaped movable process. The ar- ticulation of the movable process is linear and less heavily sclerotized than in A. nu- datus. The proximal arm of sternum IX is narrower than in nudatus and not parallel sided, but somewhat boomerang-shaped with the hook pointed dorsad and anterad. The distal arm of sternum IX is broad basad and gradually narrows to the apex. It is more than twice as broad at the base than at the apex, with a row of 4 heavy setae followed by 6 or 7 small setae extending ventrad two-thirds of the distance up the posterior margin from the apex. The ae- deagus has a distinct mid-dorsal lobe ex- tending from a point above the aedeagal struts one-half of the distance to the apex of the endchamber. Females are easily distinguished from oth- er species of Anomiopsyllus, except A. mon- tanus and sinuatus, by a deep sinus of ster- num VII and the straight posterior margin of tergum VIII. Anomiopsyllus amphibolus is distinguished from A. montanus and A. sinuatus by the presence of a bulbular ex- pansion of the terminal portion of the blind ductus seminalis (Figs. 2, 66). Length (average): 1.2-1.8 mm. Type locality: Salina, Sevier County, Utah. Type host: Neotoma desertorum Type specimens: Male holotype and fe- male allotype, male and female paratypes collected on 21 March 1929 by J. S. Stan- ford, deposited in Dr. Stanford's collection at Utah State Univ., Logan, Utah. Recorded distribution: Nevada: Beatty, Nye Co., (Hubbard, 1949); Utah: many re- cords (Howell, Stark). New distribution: Arizona: Mojave and Coconino counties (north of the Colorado River); Nevada: Clark and Lincoln counties. Material examined: Arizona: Coconino Co.: 1 female, ex Neotoma albigula, 3 km E of Williams (juniper woodland), 20 April 1949 (PHS); 2 males, ex Neotoma lepida, 55-65 km N of Williams, 2134 m (cedar), 22 April 1939 (PHS). Mojave Co.: 1 male, 1 fe- male, ex Neotoma lepida, 19 km NW of Hackberry, 11 m, 8 April 1939, (PHS); 1 fe- male, ex Neotoma lepida. Cutler Pockets, 10 March 1957, F. E. Durham (HF); 1 male, ex Neotoma lepida, Mt. Trumbull, 12 March, 1957, F. E. Durham (HF); 1 female, same host, Fulton Springs, 2 April 1959, F. E. Durham (HF); 1 female, ex Neotoma fus- cipes'\ no specific locality, 26 April 1938 (PHS). Nevada: Clark Co.; 1 male, 1 fe- male, ex Neotoma mexicana, 16 km N, 18 km W of Las Vegas, 2592 m, 19 March 1949; 1 female, ex Neotoma fuscipes, no specific locality, 14 April 1938 (PHS). Lin- coln Co.: 1 female, ex Onychomys leucogas- 'The record from Neotoma fuscipes is undoubtedly an error, since N. fuscipes is not recorded from Arizona. The .V. albigula.) rd should probably be ascribed to June 1977 Barnes et al.: Anomiopsyllus Fleas 157 Figs. 13-16. Finger and clasper: 13, A. walkeri; 14, A. falsicalifornicus falsicalifornicus; 15, A. falsicalifornicus congruens; 16, A. martini. 158 Great Basin Naturalist Vol. 37, No. 2 ter, 39 km SW of Caliente, 1220 m (cactus, sage), 4 May 1939 (PHS). Nye Co.: 6 males, 10 females, ex Neotama albigula, 27 km NE of Pahrump, 1982 m (rocky timber), 19 De- cember 1949, E. Kudka; 3 males, ex Per- omyscus maniculatus, same locality, (PHS); 2 males, 4 females, ex Neotoma lepida nest, 28 June 1965, Merino; 1 male, ex Neotoma lepida. Mercury, 12 March 1961; 1 female, ex Neotoma lepida nest. Mercury, 15 June 1965, Merino; 1 female, same host, same lo- cation, 1 July 1965, Merino; 1 male, ex Per- omyscus. Mercury, 6 October 1959, NAV- KAI. Utah: Beaver Co.: 1 female, ex Reithrodontomys megalotis, Wahwah Springs; 1 male, ex Neotoma lepida, 13-14 September 1950, D. E. Beck. Box Elder Co.: 1 male, 1 female, host not specified, Lucin small reservoir at site of abandoned CCC camp, 18 June 1952, Beck and Kill- pack; 2 males, 3 females, ex Neotoma le- pida, Lucin, 14 February 1953, Beck et al. Davis Co.: 4 males, 6 females, ex Neotoma nest, Roy, 13 October 1951, Kohls and Sar- gent (KM). Duchesne Co.: 1 female, ex Per- omyscus maniculatus, locality not stated (Bluebell, junipers), 21 March 1953, Kill- pack. Grand Co.: 1 male, ex Neotoma sp.. Devil's Garden Area of Arches National Monument, 10 May 1950, D. M. Allred; 6 males, 6 females, ex Neotoma lepida nest, Moab S side of U 93 near willow sweeps 4.8 km W of Arches National Monument (red sandstone), 1 September 1950, D. M. Allred; 4 males, 4 females, same host, Moab 1.6 km S of US 160 and U 93 Junction (Navajo sandstone environs), 9 September 1950, D. M. Allred; 9 males, 20 females, same host, .8 km S of US 160 and U 93 Junction (Navajo sandstone environs), 9 Sep- tember 1950, D. M. Allred. Iron Co.: 2 fe- males, ex Neotoma lepida nest, 14.5 km W of Parowan (canyon defile), 6 September 1950, Beck and Allred. Juab Co.: 1 male, ex Neotoma lepida, 1.6 km NW of Joy, 1 June 1951, Allred et al.; 4 males, 10 females, same host. Fish Springs mine shaft in foot- hills 4.8 km N of Harris Ranch House (rocky ledges), 15 June 1951, Beck et al; 3 males, 8 females, ex Neotoma Lepida nest, Lynndyl (fallen tree, consisting of juniper twigs and bark), 17 November 1951, Barn- um and Moore; 30 males, 40 females, ex Neotoma lepida, N of Lynndyl and E of old CCC camp, same date, Barnum and Moore; 2 males, 1 female, same host, Lynndyl (juni- per tree), same date, Barnum and Moore; 40 males, 39 females, ex Neotoma lepida nest, N of Lynndyl and E of CCC camp, 17 November 1951, Milander and Cloward; 151 males, 169 females, same host, E of Je- richo (juniper), 13 December 1952, Beck et al.; 136 males, 165 females, same host, Je- richo by US 6, 13 March 1954, Howell and Beck; 35 males, 74 females, same host, Je- richo E side of Hwy 50-6, 4.5 km N of train station (cedar), 11 April 1954, J. F. Howell; 23 males, 49 females, same host, same location, 8 May 1954. J. F. Howell; 1 male, 8 females, same host, N of Jericho on E side of Hwy 50-6, 9 June 1954, J. F. Howell; 2 males, 2 females, same host 4.5 km N of Jericho (juniper and sage), 8 July 1954, J. F. Howell; 1 female, same host, Je- richo (juniper and sage), 21 July 1954, How- ell and Beck; 2 males, same host, 4.8 km N of Jericho, 3 September 1954, J. F. Howell; 1 male, 4 females, ex Neotoma lepida lepida nest, 4.8 km N of Jericho on US 6, 29 Sep- tember 1954, J. F. Howell; 41 males, 68 fe- males, same host, same location, 8 October 1954, J. F. Howell; 81 males, 82 females, ex Neotoma lepida nest, Jericho E side of US 6, 4.5 km N of train station, 6 February 1954, Howell and Beck; 18 males, 39 fe- males, ex Neotoma lepida, 4.8 km N of Je- richo on E side of US 6, 29 October 1954, Howell and Jaussi; 65 males, 65 females, same host, same location, 12 November 1954, Howell; 207 males, 285 females, ex Neotoma lepida, same location, 2 December 1954, Howell and Nielson; 40 males, 39 fe- males, ex Neotoma lepida nest, 4.8 km NE Jericho, 7 January 1954, J. F. Howell; 75 males, 69 females, same host, 4.8 km NE of Jericho on E side of US 6, 25 January 1955, J. F. Howell; 12 males, 8 females, ex Neo- toma lepida, 4.8 km NE of Jericho, 3 March 1959, BYU. Kane Co.: 1 male, ex Dipo- domys ordii, 16 February 1972, NAV-KAI; 24 males, 7 females, ex Neotoma albigula, 5 December 1971, NAV-KAI; 1 female, ex Neotama lepida, Navajo Wells, 24 June 1951, BYU; 1 male, 1 female, same host, 8.8 June 1977 Barnes et al.: Anomiopsyllus Fleas 159 km W of Adairville in Catstaire Canyon (rocky area), 8 September 1951, Beck and Allred; 1 female, same host, Navajo Wells SW of Corral (rocky ledges), 7 September 1951, Beck and Allred; 1 male, 1 female, same host, same location, 6 September 1951, Beck and Allred; 1 male, 1 female. same host, W entrance to Catstaire Canyon (Upper Sonoran sagebrush), 9 May 1952, M. D. Coffey; 1 male, 2 females, ex Neotoma lepida nest, Catstaire Canyon (sagebrush), 8 May 1952, M. D. Coffey; 1 male, same host, W entrance to Catstaire Canyon (rocky ledge), same date, M. D. Coffey; 8 Figs. 17-25. Variations in spiniform placement on finger: 17, A. nudatus mexicanus; 18-25, A. falsicalifornicus congruens. 160 Great Basin Naturalist Vol. 37, No. 2 males, 9 females, ex Peromyscus crinitus, 11 December 1971, NAV-KAI; 1 male, 2 fe- males, ex Peromyscus maniculatus, 40.2 km E of Kanab about 457 m off road near Paria and King manganese mine roads junc- tion, 9 May 1952, M. D. Coffey; 1 male, 1 female, ex. Peromyscus truei, 17 February 1972, NAV-KAI. 4 males, 2 females, ex Neo- toma cinerea, 6.4 km N of Kanab, 1494 m, 14 April 1949 (PHS). Millard Co.: 11 males, 31 females, ex Neotoma lepido, Lynndyl, 17 November 1951, Melander and Cloward. San Juan Co.: 1 female, ex Neotoma alhi- gula, Montezuma Creek 4.8 km N of trad- ing post. Beck et al.; 1 male, 6 females, ex Neotoma mexicana, Blanding, 7 September 1956; 1 male, 1 female, ex Peromyscus ma- niculatus 32.2 km S of Moab (sandy wash cliff), 7 May 1951, Allred et al.; 3 males, 2 females, ex Neotoma lepida, Yuba Dam 1.6 km from Hwy 28, 182.8 m W on N side of reservoir, 18 April 1952, Beck and Beck; 9 males, 9 females, ex Neotoma lepida nest, Yuba Dam, 29 April 1960, Beck. Sanpete Co.: 2 males, 2 females, ex Neotoma lepida, Yuba Dam, 18 April 1952, D. E. Beck. Tooele Co.: 1 female, ex Neotoma lepida 3.2 km N Little Granite Mt., 10 March 1955, E. A. Shippee (U of U); 1 male, 1 fe- male, ex Neotoma sp. nest, Sandy Pars, Dugway Valley, 15 April 1954, D. Allred (U of U); 3 males, 1 female, ex Neotoma le- pida nest, Dugway Proving Grounds, 3 March 1955 (U of U); 10 males, 7 females, same host, same location, 14 April 1955 (U of U); 1 male, 1 female, same host, same lo- cation, 6 April 1955 (U of U); 17 females, same host, same location, 6 January 1955 (U of U); 13 males, 12 females, same host, same location, 7 February 1955 (U of U); 1 male, 1 female, same host, same location, 20 January 1955 (U of U); 10 males, 12 fe- males, same host, same location, 20 January 1955 (U of U); 10 males, 12 females, same host, same location, 16 March 1955 (U of U); 3 males, 2 females, same host, same lo- cation, 9 December 1954 (U of U); 6 fe- males, same host, same location, 27 October 1954 (U of U); 6 males, 1 female, same host, same location, 24 November 1954 (U of U); 1 female, ex Peromyscus truei. Cane Springs, Cedar Mts., Dugway Proving Grounds, 21 January 1953 (U of U); 1 male, ex Peromyscus crinitus. Camel Back Mt., Dugway Proving Grounds, 30 January 1953 (U of U). Uintah Co.: 1 male, ex Peromyscus maniculatus. Dinosaur National Monument 1.2 km SSW of headquarters on N bank of river (sandy wash), 14 August 1951, Beck et al. Utah Co.: 8 males, 14 females, ex Neo- toma, Chimney Rock Pass, 25 October 1968, Clark; 1 female, ex Neotoma nest, Provo, top of ladder in Buckley's mine in Rock Canyon, 24 November 1948, D. M. Allred; 1 female, ex Neotoma cinerea, Springville, 29 November 1948, V. J. Tip- ton; 2 males, ex Neotoma cinerea nest, Thistle (tunnel composed of rags, sticks, pa- per, plants, rope, and many bones), 2 No- vember 1951, Barnum et al.; 10 males, 16 females, same host, Spanish Fork Canyon, 2 November 1951, Barnum et al.; 3 males, 1 female, same host, Spanish Fork Canyon, cave N of Ma's and Pa's Cafe in Canyon, 5 November 1951, Barnum et al.; 1 female, 2 males, ex Neotoma lepida. Eureka, 17 No- vember 1951, Barnum and Moore (BYU); 1 female, ex Neotoma cinerea, Spanish Fork Canyon, small cave (dried plants), 2 Novem- ber 1951, W. Milander; 1 female, ex Neo- toma lepida, Elberta, Chimney Rock Pass, 15 October 1949, D. M. Allred; 19 males, 29 females, ex Neotoma lepida nest. Cedar Valley (juniper-rocky hillside), 22 February 1953, Beck and Coffey; 1 female, ex Neo- toma lepida, nest, NE Chimney Rock Pass near abandoned mine, 28 March 1953, Cof- fey and Liddiard; 1 female, ex Peromyscus maniculatus. Cedar Valley on hill NE of big juniper grove, 25 March 1951, Allred et al.; 1 male, 2 females, same host, 3.2 km W of Genola on main Hwv to Eureka (rocky cliffs), 13 January 1952,' D.E. Beck. Wash- ington Co.: 1 female, ex Neotoma lepida, Beaver Dam Mtns., 22 April 1953, Howell; 1 male, ex Dipodomys merriami, W side Beaver Dam Wash bottom (alfalfa fields of Terry Ranch), 21 December 1950, Beck and Allred; 1 female, ex Dipodoinys merriami, 3.2 km S of Rockville, 13 July 1953, Beck and Killpack; 1 female, ex Neotoma lepida, St. George, 25 November 1949, Allred and Barnum; 43 males, 63 females, ex Neotoma lepida nest, Beaver Dam Wash, .8 km NE June 1977 Barnes et al.: Anomiopsyllus Fleas 161 Figs. 26-29. Finger and clasper: 26, A. durangoensis; 27, A. novomexicanensis; 28, A. traubi (redrawn fom rera 1951 and Barnes 1963); 29, A. oaxacae. 162 Great Basin Naturalist Vol. 37, No. 2 Terry Ranch (joshua tree peat), 21 Decem- ber 1950, Beck and Allred; 2 males, 5 fe- males, same host, Beaver Dam Wash of Beaver Mtns. (joshua tree foothills), 23 Feb- ruary 1952, D. E. Beck; 1 female, ex Neo- toma lepida, Beaver Dam Wash (flat, gravel), 16 April 1952, Grant et al.; 1 fe- male, ex Peromyscus eremicus, S of Hwy 15 and Berry Springs (sandy area), 19 Decem- ber 1950, Beck and Allred; 1 female, same host. Berry Springs (lava rock), 19 Decem- ber 1950, Beck and Allred; 1 female, ex Per- omyscus maniculatus, 6.4 km W of Hwy 59 state line crossing at Utah-Arizona border, 12 July 1952, Beck and Killpack; 4 females, ex Neotoma lepida nest, Rush Valley (juni- per), 9 November 1951, Barnum et al; 2 fe- males, ex Neotoma lepida nest, Toquerville, 5 September 1951, Beck and Allred. Wyom- ing: Sweetwater Co.: 1 female, ex Neotoma cinerea, 24 km SW of Green River, 1920 m (rocky ledges), 23 August 1938 (PHS). Discussion: This species is most closely related to A. montanus. Characteristics of the male clasper, e.g., configuration of the fixed process of the clasper and presence of a row of setae above the acetabular seta (Fig. 6), show relationship between males of these two species. The shape of the sperma- theca and configuration of sternum VII (Fig. 80) indicate a relationship between the females of these two species. With the exception of A. montanus, am- phibolus occurs farther north than any other species of Anomiopsyllus, and it is the Great Basin representative of the genus. The distributional pattern includes north- western Arizona from whence it extends northward between the Rocky Mountains and the Sierra Nevada Mountains into Wyoming. The species has been recorded most frequently from Utah where much re- search has been conducted by Allred, Beck, Howell, and Tipton at Brigham Young Uni- versity and Stanford at Utah State Univer- sity. This species has also been recorded from Nye and Douglas counties, Nevada (Beck et al. 1953, Hopkins and Rothschild 1962). Its distribution may include those parts of eastern and northeastern California which have a habitat similar to that of the Great Basin and where Neotoma lepida oc- curs, but we have not collected A. amphi- holus from that area during this study. The primary host appears to be Neotoma lepida, although records include N. cinerea and Peromyscus in frequent numbers. Re- cords from fortuitous hosts are most often obtained during plague surveys. Stark (1959) has pointed out that A. am- phiholus may prefer dry habits, since nests of N. cinerea occurring at high elevations (noted in Beck et al. 1953) had fewer fleas per nest than those of N. lepida occurring at lower elevations. This fails to take into consideration possible host differences, which may be a significant factor. Neotoma cinerea has been placed in a separate sub- genus by Goldman (1910), Hall and Kelson (1959), and others, and may well differ sig- nificantly from the standpoint of flea host selection. Howell (1957) states that in the vicinity of Jericho, Juab County, Utah, A. amphi- bolus adults appear in nests of N. lepida in October and reach a peak in December. This seasonal distribution almost duplicates that of A. falsicalifomicus in California. Among the many specimens examined, very little variation was evident; A. amphi- bolus from northwestern Arizona did not differ in any significant detail from speci- mens collected in Utah. This is in distinct contrast to some Anomiopsyllus species, e.g. A. falsicalifomicus, which shows consid- erable variation both geographically and among individuals in a series. The apparent stability of A. amphibolus over a broad area may be explained in part by the rather con- tinuous distribution of its host, N. lepida, through the Great Basin (see Hall 1946). The climate of the Great Basin, though complex, changes primarily with latitude and altitude. Populations of A. amphibolus would be expected to be relatively contin- uous with much opportunity for uninter- rupted gene flow. The area occupied by A. falsicalifomicus is subject to more climatic variability resulting not only from very het- erogeneous topography and latitude, but also from proximity to or distance from the sea. Populations of A. falsicalifomicus are disjunct in their distribution (of both the to- tal population and especially the local pop- June 1977 Barnes et al.: Anomiopsyllus Fleas 163 Illations) which results in much gene flow interruption. Anomiopsyllus sinuattis Holland Figs. 9, 33, 54, 65, 79, 94 Anomiopsyllus sinuatus Holland, 1965, Canadian Ent. 97: 1053, 1058; Lewis, 1974, J. Med. Ent. 11(2): 154. Diagnosis: Anomiopsyllus sinuatus is closely related to A. montanus in that both have the movable process L-shaped with a large, blunt spiniform mesally inserted at the distal and proximal portions. Anomio- psyllus sinuatus has a broader, shorter proc- ess than montanus. The ninth sternum differs from that of A. montanus by the presence of three thickened setae followed by a row of thinner setae as compared to four thick setae in A. montanus. The sinus formed by the inner margins of the prox- imal and distal arms is deep and narrow in Figs. 30-33. Male ninth sternum: 30, A. amphibolus; 31, A. montanus; 32, A. nidiophilus; 33, A. sinuatus. 164 Great Basin Naturalist Vol. 37, No. 2 sinuatus and broadly rounded in montanus. Anomiapsijllus sinuatus, like A. montanus, has a sinus on the caudal border of sternum VIII. Female characters are not sufficiently dis- tinctive to separate sinuatus from mon- tanus. Type locality: 48 km W of Durango, Du- rango, Mexico, 1982 m. Type host: Nest of Peromyscus sp. Type specimens: Male holotype and fe- male allotype, 4 male and 11 female para- types collected on 5 June 1961 by J. E. H. Martin, deposited in the Canadian National Collection. Discussion: Anomiopsyllus sinuatus and A. montanus present an interesting distribu- tional problem in that montanus has only been collected at the northern distribution of Anomiopsyllus and sinuatus has only been collected at the southern portion. Col- lections so far have not indicated that there are any intermediate forms between these two populations; however, further speci- mens should be collected with the view of hnking these geographically separated spe- Anomiopsyllus montanus Collins Figs. 8, 31, 53, 66, 81, 94 Anomiopsyllus montanus Collins, 1936, Ent. News 47: 128-130; Ewing and I. Fox, 1943, U.S. Dept. Agric. Misc. Publ. ,500: 115; Hubbard, 1947, Fleas of western North America: 82-83; Holland, 1952, Canadian Ent. 84(3): 65-72; Holland, 1965, Canadian Ent. 97: 1053, 1055, 1058; Senger, 1966, J. Kans. Ent. Soc. 39(1): 106; Jellison and Glesne, 1967, Index to the literature of Siphonaptera of North America, Suppl. 2: 1.3-19; Jelli- son and Senger, 1973, Mont. Agr. Exp. Sta. Res. Rept. 29: 18-19; Lewis 1974, J. Med. Ent. 11(2): 154. Arctopsylla montanus: Jellison, Kohls, and Mills, 1943, Mont. State Board Ent. 2: 2 (error for Anomio- psyllus). Anomiopsyllus nudatus: Dunn and Parker, 1923, Public Hlth. Rept. 38: 2774-2775 (misidentification). Diagnosis: Anomiopsyllus montanus is closely related to A. sinuatus but differs in that there are four heavy setae near the apex of sternum IX instead of three as in si- nuatus. The movable process of A. mon- tanus is narrower and longer with a deeper sinus than in sinuatus. Female characters are not sufficiently dis- tinctive to separate montanus from si- nuatus. Type locality: Canyon Creek, W of Ham- ilton, Montana. Type host: Bubo virginianus occidentalis Type specimens: Male holotype and fe- male allotype collected in 1922 by L. H. Dunn and R. R. Parker, deposited in the National Institutes of Health. Discussion: This flea was first collected from a horned owl {Bubo virginianus) near Hamilton, Montana, and identified as Ano- miopsyllus nudatus by Dunn and Parker (1923). Collins (1936) rectified the mis- determination and described the species as new. Subsequent records have been few. We have examined only material from near Hamilton, Montana, from Bubo virginianus (part of the original series) and Neotoma cinerea. Holland (1952) recorded the species from near Banff, Alberta, Canada. No other records are available. Anomiopsyllus montanus has the north- ernmost distribution of any species in the genus. We suspect that the primary host is a species of Neotoma, possibly N. cinerea since no other Neotoma species occurs as far north as Hamilton, Montana, and Al- berta, Canada (Hall and Kelson 1959). The original discovery of this flea on the horned owl should not be considered too unusual, as fleas are often found on the predators of their hosts. In addition, horned owls often nest in cliffs, which are a preferred nesting site of A^. cinerea. Anomiopsyllus nudatus nudatus (Baker) Figs. 11, 36, 55, 68, 84, 94 Tijphlopsylla nudata Baker, 1898, J. N. Y. Ent. Soc. 6:56. Anomiopsyllus nudatus Baker, 1904, Proc. U.S. Nat. Mus. 27:425, 426, 452; Fox, 1914, Hyg. Lab. Bull. 97: 25; Jordan and Rothschild, 1915, Ectoparasites 1:45; Dunn and Parker, 1923, PubUc Hlth. Repts. 38:2774-2775; Fox, 1929, Ent. News 40:218; Hirst, 1926, Ceylon J. Sci. 1: 261; Wagner, 19,30, Mag. de Parasitol. 1: 1.35; Stanford, 1931, Proc. Utah Acad. Sci. 1931: 153; Collins, 1936, Ent. News 47: 129-130; Eskey and Haas, 19,39, Public Hlth. Repts. 54: 1472; Eskey and Haas, 1940, Public Hlth. Bull. 254:29-74; Hub- bard, 1940, Pac. Univ. Bull. 37(4); Augustson, 1943, Bull. South. Calif. Acad. Sci. 42(2): 73; Ewing and I. June 1977 Barnes et al.: Anomiopsyllus Fleas 165 Fox, 1943, U.S. Dept. Agric. Misc. Publ. 500: 115 Hubbard, 1947, Fleas of western North America: 79-80 Eads and Menzies, 1948, J. Kans. Ent. Soc. 21: 135; Hubbard, 1949, Bull. South. Calif. Acad. Sci. 48: 125 Pollitzer, 1952, Bull. WHO 7:231-342, Annex 1; Au- gustson and Wood, 1953, Bull. South. Calif. Acad. Sci. 52: 48-56; Wehrle, 1953, Pan-Pacific Ent. 24: 39; Hold- enreid and Morlan, 1955, J. Inf. Dis. 96: 135; August- son, 1955, Bull. South. Calif. Acad. Sci. 54(1): 36-39; Knudsen, 1956, Bull. South. Calif. Acad. Sci., 55(1): 1- 63; Beer et al., 1959, J. Parasitol. 45(6): 605-623; Stark, 1959, Siphonaptera of Utah: 107; Holland, 1965, Cana- dian Ent. 97: 1051-1058; Barnes, 1965, Pan-Pacific Ent. 41(4): 272-280; Jellison and Glesne, 1967, Index to the literature of Siphonaptera of North America, Suppl. 2: 13-19; Tipton and Saunders, 1971; Brigham Young Univ. Sci. Bull., Biol. Ser. 15(2): 18; Pratt and Stark, 1973, DHEW Publ. 74-8267: 11; Haas et al, 1973, J. Med. Ent. 10(3): 282; Lewis, 1974, J. Med. Ent. 11(2): 154. Anomiapsyllus californicus: Baker, 1904, Invert. Pac. 1: 39-40; Fox, 1926, Pan-Pacific Ent. 2: 183-184. Diagnosis: The three subspecies of A. nti- datus: nudatus, mexicanus, and hiemalis can only be distinguished from each other by characters in the males. This separation is based on characters of the movable process and sternum IX. Anomiopsyllus nudatus nu- datus can be separated readily from the other two subspecies by the presence of two closely spaced (less than 20 u) spiniforms on the distal portion of the movable process and the presence of a thin bristle at the arc Figs. 34-36. Male ninth sternum: 34, A. nudatus hiemalis; 35, A. nudatus mexicanus; 36, A. nudatus nudatus. 166 Great Basin Naturalist Vol. 37, No. 2 of curvature of the distal arm of sternum IX. The movable process of the clasper in nudatus nudatus is narrow (M = 68 u, range 58 to 74 u) at base measured at the widest point and relatively short (M = 109 u, range 102 to 119 u). Female characters are not sufficiently dis- tinctive to separate the subspecies of nu- datus. Length: (average)— Male, 1.2 mm; female, 1.8 mm. Type locality: Tucson, Pima Co., Arizona. Type host: Neotoma albigula. Type specimen: Female, U.S. National Museum. Material examined: Arizona: Cochise Co.: 1 male, ex Neotoma lepida, 29 October 1938 (PHS); 1 male, ex Neotoma lepida, no date (PHS). Pima Co.: 1 female, ex Neotoma lepida, 7 February 1939 (PHS); 1 female, ex Neotoma lepida, 3 November 1938 (PHS); 1 female, ex Neotoma lepida, 4 November 1938 (PHS); 1 female, ex Dipodomys mer- riami, 7 February, 1939 (PHS); 1 male, ex Spermophilus harrissi, 8 February 1939 (PHS). California: Kern Co.: 1 male, ex Neotoma sp. Burbank Girl Scout Camp, Frazier Mtn. (oak, pine), 2 June 1958, Campbell (BVC); 1 female, ex Neotoma fus- cipes nest, 0.8 km E of Frazier Park (oak woodland), 1 April 1959, Barnes (BVC); 17 males, 37 females, ex Neotoma fuscipes nests, 6.9 km N, 3.7 km W of Willow Springs (juniper woodland), 19 February 1959; 25 males, 54 females, same host, same locality, 22 January 1960. Los Angeles Co.: 13 males, 34 females, ex Neotoma fuscipes nests, 7.7 km N, 3 km W of Sandburg (oak woodland), 1 April 1959, Barnes (BVC); 10 males, 28 females, ex Neotoma fuscipes nests, 4.5 km S, 1477 km W of Fairmount (juniper woodland), 1 April 1959, Barnes (BVC); 8 males, 17 females, ex Neotoma fuscipes, 2.2 km S of Gorman (oak, yucca), 1 April 1959, Barnes (BVC); 1 male, 1 fe- male, ex Neotoma fuscipes. School Canyon 5.9 km E of Glendale (oak woodland), 28 January 1959, D. Rohe (BVC); 1 male, 1 fe- male, ex Neotoma fuscipes, Burbank (oak woodland), 6 April 1959, D. Rohe (BVC); 3 males, 2 females, ex Neotoma lepida. Big Tujunga Canyon, 4 January 1942, J. Couffer (HE); 1 male, 3 females, ex Neotoma lepida, San Gabriel Canyon, 15 January 1942, G. Augustson (HE); 3 males, 1 female, ex Neo- toma nest, Chilao, 16 August 1942, G. Au- gustson (HE). Orange Co.: 1 female, ex Neo- toma sp., Irvine (cactus, Nicotiana), 19 April 1955, J. Poll (BVC); 1 male, 1 female, ex Neotoma sp., same locality (cactus, Nico- tiana), 26 March 1957, E. Perry (BVC); 5 males, 6 females, ex Neotoma sp., same lo- cality (cactus, Nicotiana), 19 November 1952, J. Poll (BVC); 12 males, 20 females, ex Neotoma fuscipes nest, Santiago Canyon (cactus, Nicotiana), no date, G. Augustson (HE). Riverside Co.: hundreds, ex Neotoma fuscipes, San Timoteo, many dates, R. Ryck- man; 3 females, ex Neotoma sp., 1.6 km E of McAfee Ranch, San Timoteo Canyon, 18 March 1955, J. Poll (BVC); 6 females, ex Neotoma nest, Matthew Lake, 3 February 1951, Ames; 4 females, ex Neotoma lepida, Matthew Lake, 3 February 1951, Ames; 2 males, 9 females, ex Neotoma lepida, Mat- thew Lake, 8 February 1951, Ames. San Bernardino Co.: 2 males, 1 female, ex Neo- toma lepida, San Bernardino (oak woodland and chaparral), 21 March 1957, Barnes (BVC); 1 female, ex Neotoma lepida, 5.4 km N, 6.4 km E of Pioneer Town (Joshua tree, yucca), 3 December 1956, K. Murray (BVC); 8 males, 3 females, ex Neotoma le- pida, 3.2 km to 3.8 km NW of Yucca Val- ley (yucca desert), 4 December 1956, K. Murray (BVC); 68 males, 144 females, ex Neotoma fuscipes nest, 1.6 km E of Cajon (heavy chaparral), 22 December 1957, Barnes (BVC); 1 male, 1 female, ex Neotoma fus- cipes nest, 3.5 km SE of Desert Springs (juniper woodland), 22 January 1959, Barnes (BVC); 1 female, ex Neotoma sp., George Air Force Base, Victorville (juniper wood- land), 25 October 1951, J. Poll (BVC); 1 male, 2 females, ex Neotoma sp., Big Bear Lake (oak, pine), 30 May 1951, J. Poll (BVC); 1 female, ex Neotoma sp., Seven Oaks, 10 May 1936 (RM); 1 male, 8 May 1936 (RM); 1 female, ex Neotoma lepida, Batista Canyon, 28 February 1951, Ames. San Diego Co.: 5 males, 9 females, ex Neo- toma fuscipes nests, 3 km E of Banner (juni- per, Rhus ovata woodland), 23 March 1960, Barnes (BVC); 10 males, 9 females, ex Neo- June 1977 Barnes et al.: Anomiopsyllus Fleas 167 toma fuscipes nest, 16 km E of Ramona (oak woodland), 23 March 1960, Barnes (BVC); 65 males, 122 females, ex Neotoma fuscipes nest, Alvarado Canyon, San Diego (oak wood and chaparral), 21 March 1960, Barnes (BVC); 1 male, ex Neotoma fuscipes nest, 7 km S of Julian, 1219 m (oak wood- land), 22 March 1960, Barnes (BVC); 4 males, 1 female, ex Neotoma nest, 3 km W of Guatay (oak woodland), 2 September Figs. 37-40. Male ninth sternum: 37, A. falsicalifornicus falsicaliforniciis; falsicalifornicus congruens. walkeri; 39, A. martini; 40, A. 168 Great Basin Naturalist Vol. 37, No. 2 1955, H. Stark (PHS); 1 female, ex Neotoma sp., San Diego, 1936, Eskey (PHS). Ventura Co.: 2 males, 7 females, ex Neotoma fus- cipes nests, Alder Creek 987 km N, 2.9 km W Fillmore, 28 January 1960, Barnes (BVC); 1 male, 1 female, ex Neotoma sp.. Ozena Public Camp 2.5 km E of Hwy. 399, 26 March 1957, J. Poll (BVC); 1 male, ex Neotoma sp.. Sycamore Canyon, 29 June 1955, E. Perry (BVC). Nevada: Nye Co.: 1 male, ex Neotoma lepida, Beatty, no date. Utah: San Juan Co.: 1 male, ex Peromyscus manicukitiis, 32 km S of Moab, 8 May 1951, (Tipton and Allred); 1 male, 2 fe- males, ex Neotoma alhigiila. Recapture Can- yon, 1738 m, 16 August 1939 (PHS); 1 male, 1 female, ex Neotoma mexicana, Westwater Canyon, 3.2 km S of Blanding, 1830 m, 19 August 1939 (PHS). New distribution: Arizona: Cochise Co. (PHS); California: Kern Co. (BVC), Riv- erside Co. (BVC, Ryckman), San Bernardino Co. (BVC), San Diego Co. (BVC and PHS), Ventura Co. (BVC). Discussion: C. F. Baker (1898) described A. nudatus on the basis of ". . . two females collected by Mr. Hubbard from the inner nest of Neotoma albigula at Tucson, Ari- zona," placing the species in the genus TypJilopsyUa, pending better knowledge of its relationships. Later, Baker (1904) de- scribed the species further and created the new genus Anomiopsyllus to contain it and the concurrently described A. californicus from Claremont, Los Angeles County, Cali- fornia (based on a series of females). Baker, at the time of the original descrip- tion of nudatus, did not designate either of his two female specimens as types, but both were deposited in the U.S. National Mu- seum. Since both specimens were collected in the Sonoran Desert at Tucson, Arizona, the subspecies occupying that area becomes the nominate subspecies. Jordan and Roths- child's male allotype, unfortunately, falls into another subspecies occupying the Colo- rado Plateau from eastern Arizona through New Mexico. Though their specimen was not available for study, the figure in the original description (Jordan and Rothschild 1915) is quite clearly that of the A. n. mexi- canus. The same figure is reproduced in Hopkins and Rothschild (1962). Never- theless, their specimen is the first described male for the genus and for the species. Jordan and Rothschild (1915) examined Baker's types and declared californicus a synonym of nudatus on the basis of sim- ilarities in the spermathecae and in charac- ters of the metatarsal setae. Jordan and Rothschild described and figured a male of nudatus from Paradise, Arizona, designating their specimen as allotype. Although their specimen is of nudatus, the "allotypic" lo- cality falls within the range of A. n. mexi- canus. Their figure of the movable process is clearly that of mexicanus. Carroll Fox (1926) acquired a series of Anomiopsyllus from Claremont, California, the type locality of californicus, which in- cluded a male differing from that described as nudatus by Jordan and Rothschild. He, therefore, concluded the latter authors to be in error concerning the synonomy, and de- scribed the new male as allotype of Baker's revived A. californicus. Jordan, on exam- ining Fox's series, found two species; one "true californicus'^ identical to Baker's types, the other a new species requiring a name (Fox 1929). Fox (1929), therefore, named the species A. falsicalifornicus, based on his 1926 description. Collins (1936), in separating A. montanus from nudatus, agreed with Jordan and Rothschild's syno- nomy and figured the female sternum VII of both nudatus and falsicalifornicus. Neither the characters used by Baker nor those used by Jordan and Rothschild (1915) are tenable. No character involving tarsal setation or shape of spermatheca has been found reliable in this study. Nevertheless, sternum VII of females offers some degree of reliability which, together with distribu- tional evidence, confirms Jordan and Rothschild's synonomy. As treated here, A. nudatus includes three subspecies, differentiated on the basis of male morphological characters, geogra- phic distribution, and habitat. Anomiop- syllus hiemalis Eads and Menzies (1948) is reduced to subspecific rank under A. nu- datus (Baker 1898); A. hiemalis mexicanus (Holland 1965) is also transferred here as a subspecies of A. nudatus. June 1977 Barnes et al.: Anomiopsyllus Fleas 169 Phenotypic characters used to differen- tiate these subspecies are length and width of the movable process, distance between spiniforms of the movable process, and pres- ence or absence of modified setae on the posterior margin of sternum IX. Measure- ments were made by means of a calibrated ocular grid at 430 X; the grid units were then converted to microns. Length and width measurements of the movable process were made along the longitudinal axis from the tip of the process to the "notch." The width was measured at an approximate right angle from the "notch" across the broadest part of the process. Measurements between the distal pair of spiniform setae were from center to center at the base of each spiniform. AnomiopsyUiis nudatus mexicanus Holland, n. comb. Figs. 12, 35, 57, 70, 83, 94 Anomiopsyllus nudatus: Holdenreid and Morlan, 1955, J. Inf. Dis. 96: 135; Stark, 1959, Siphonaptera of Utah: 107. Anomiopsyllus hiemalis mexicanus Holland, 1965, Canadian Ent. 97: 1056; Lewis, 1974, J. Med. Ent. 11(2): 154. Diagnosis: Males of A. n. mexicanus are separable from A. n. nudatus and A. n. hiemalis by the presence of a heavy bristle at the arc of curvature of sternum IX. The movable process of the clasper is longer than in nudatus or hiemalis (M = 118 u, range 100 to 133 u) and about equal to nu- datus in width. The distance between the distal spiniform pair is greater than in nu- datus, but less than in hiemalis (M = 29 u, range 23 to 35 u). Female characters are not sufficiently dis- tinctive to separate the subspecies of A. nu- datus. Type locality: Mesa del Huracan (180° 15' W, 29° 40' N), Chihuahua, 2256 m. Type host: nest of Neotoma sp. Type specimens: Male holotype and male paratype collected on 21-25 July 1964 by J. E. H. Martin, deposited in Canadian Na- tional Collection. Material examined: Arizona: Apache Co.: 1 male, 2 females, ex Neotoma lepida, 44 km W of Springerville, 2302 m (cactus), 1 October 1938 (PHS); 1 male, ex Neotoma sp., 24 km W of Springerville, 1829 m (grass), 26 September 1938 (PHS). Coconino Co.: 1 female, ex Neotoma alhigula. Pine Forest, 9.7 km SE of Red Lake, 2073 m (grass, cedars), 30 April 1938 (PHS); 1 male, 2 females, ex Neotoma alhigula, 29-32 km SW of Winslow, 1463 m (chaparral), 23 Au- gust 1939 (PHS); 1 female, ex Peromyscus maniculatus. Pine Forest, 9.7 km SE of Red Lake, 2073 m (grass, cedars), 30 April 1938 (PHS); 1 female, ex Peromyscus manicu- latus, 55-65 km N of Williams, 2134 m (ce- dar), 22 April 1938 (PHS); 1 male, ex Neo- toma lepida, 55-60 km N of Williams, 2134 m (cedar), 22 April 1939 (PHS); 1 female, 1 male, ex Neotoma lepida, 3-6 km SW of Winona, 2073 m (pinon), 27 April 1939 (PHS); 1 female, ex Neotoma lepida 29-32 km SW of Winslow, 1463 m (chaparral), 22 August 1939 (PHS). Mojave Co.: 1 male, ex Neotoma lepida, S. entrance. Grand Canyon, 5 June 1942, R. L. Rutherford (HE). Navajo Co.: 1 male, 1 female, ex Neotoma lepida, 32-41 km SE of Winslow, 1463 m (sage, cactus), 25 August 1939 (PHS); 1 male, 1 female, ex Neotoma lepida, 4-11 km NW of Snowflake, 1829 m, 11 May 1939 (PHS); 1 female, ex Neotoma lepida, 35-42 km SW of Holbrook, 1798 m, 10 May 1939 (PHS); 3 females, 3-8 km W of Snowflake, 1829 m, 12 May 1939 (PHS); 2 females, ex Neotoma lepida, 29-34 km SW of Holbrook, 1799 m (junipers), 9 May 1939 (PHS). Yavapai Co.: Figs. 41-46. Variations in apex of male ninth sternum of A. falsicalifornicus congruens. 170 Great Basin Naturalist Vol. 37, No. 2 4 males, 3 females, ex Neoknna cinerea, 22 km E of Seligman, 22 April 1938 (PHS); 1 male, ex Neotoma cinerea, 6 km S of Ash- fork, 1585 m (rocky), 27 April 1938 (PHS); 1 male, 1 female, ex Peromi/scus sp., 23 km E of Seligman, 1616 m, " 22 April 1938 (PHS); 1 male, 1 female, ex Peromysctis sp., 8 km W of Seligman, 1573 m (juniper), 19 April 1938 (PHS); 3 males, ex Peromijscus sp., 6 km S of Ashfork 1585 m (rocky), 27 April 1938 (PHS); 2 males, 2 females, ex Neotoma lepida, 16 km SW of Seligman, 1585 m, 20 April 1938 (PHS); 4 males, 3 fe- males, ex Neotoma lepida, 8 km NW of Clarkdale, 1067 m, 21-22 April 1938 (PHS); 3 females, ex Neotoma lepida, 10 km SE of Cottonwood, 23 April 1938 (PHS); 2 fe- males, ex Neotoma lepida, near Wickenburg, 19 April 1938 (PHS); 2 females, ex Neotoma alhigula, 16 km N of Paulden, W side of Verde River, 22 April 1938 (PHS), 1 male, 1 female, ex Neotoma alhigula, 5 km NW of Dewey, 1524 m (grass, clay), 16 April 1937 (PHS) 1 male, 1 female, ex Neotoma le})ida, 823 m, 13 April 1938 (PHS). Colo- rado: La Plata Co.: 1 female, ex Neotoma lepidii. Hillside 10 km N of Durango, 2012 m (oaks), 13 June 1939 (PHS). Las Animas Co.: 2 males, 1 female, ex Neotoma sp., 5 June 1943 (PHS); 1 female, ex Neotoma al- bigula. Las Animas, 20 June 1941 (PHS). Kansas: Logan Co.: 1 male, 2 females, ex Neotoma sp., 30 May 1945 (PHS). Nebraska: Morrill Co.: 3 males, 2 females, ex Neotoma alhigula, 20 May 1949 (PHS). Neiv Mexico: Bernalillo Co.: 2 females, ex Neotoma alhi- gula, Cedro Canyon, 3 km SE of Tijeras, 1677 m (sage, juniper), 20 May 1939 (PHS). Catron Co.: 3 males, 3 females, ex Neotoma alhigula, 13 km N of Alma, 1524 m (juni- per), 20 May 1939 (PHS); 1 male, ex Neo- toma alhigula, 3.2 km E of Alma, 1402 m (sage), 17 May 1939 (PHS); 2 males, 2 fe- males, ex Neotoma alhigula, 8-16 km N of Glenwood, 1402 m, 16 May 1939 (PHS); 1 male, ex Dipodomys ordii, 3 km E of Alma, 1402 m (sage), 17 May 1939 (PHS); 2 males, ex Neotoma lepida, same locality, 17 May 1939 (PHS); 1 female, ex SpermophUus vari- egatus, between Pleasanton and Mogollon, 1829-2134 m (grass), 25 April 1939 (PHS); 1 female, ex Peromyscus truei, 1.6-4.8 km S of Pleasanton, 1402 m (juniper), 19 May 1939 (PHS); 1 male, 2 females, ex Neotoma mexi- cana, 10 km NE of Adams Diggings, 2287 m (prairie, junipers) 16 August 1938 (PHS); 1 female, ex Neotoma mexicana, Cibola For- est, 10 km W of Datil, 2439 m O'uniper), 20 August 1938 (PHS); 1 female, ex Neotoma mexicana, 1.6 km SW of Datil, 2378 m (range, juniper), 24 August 1938 (PHS). Col- fax Co.: 1 female, ex Neotoma sp., 4 June 1943 (PHS). Grant Co.: 2 males, ex Per- omyscus eremicus, 3-11 km NW of Silver City, 1829 m (grass), 25 April 1939 (PHS); 1 male, 9 females, ex Neotoma alhigula. Fort Bayard Reservation, 1829 m (grass), 28 April 1939 (PHS); 1 male, 7 females, ex Neotoma alhigula, 3-11 km NW of Silver City, 1829 m (brush), 29 April 1939 (PHS); 6 males, ex Neotoma alhigula, 19-22 km SE of Silver City, 1768 m (juniper), 27 April 1939 (PHS), 1 male, 1 female, ex Neotoma alhigula, 6-8 km NW of Silver City, 1829 m (cactus), 22 April 1939 (PHS); 1 female, ex Dipodomys merriami, 19-22 km SE of Silver City, 1768 m (juniper), 27 April 1939 (PHS). Lincoln Co.: 2 males, 2 females, ex Neotoma alhigula, 43 km E of Capitan, 1936 m (rocky timber), 18 April 1949 (PHS). Rio Arriba Co.: 1 male, ex Per- omyscus sp., 5 km W of Coyote, 2591 m (timber), 10 June 1949 (PHS). San Miguel Co.: 1 male, 1 female, ex Peromyscus sp., Pecos, 22 February 1941, McMurray (RM). Sante Fe Co.: 14 males, 6 females, ex Neo- toma alhigula nests. Steel Ranch, November and December 1951, H. B. Morlan, 5 males, 10 females, ex Neotoma alhigula, 23-25 May 1939 (PHS); 1 male, ex Peromyscus manicu- latus, same locality, 23 May 1939 (PHS). Sierra Co.: 1 male, ex Neotoma alhigula, 8- 9.6 km S of Hillsboro, 1555 m (sandy), 28 June 1939 (PHS); 1 female, ex Neotoma al- higula. Canyon, 6 km W of Hillsboro, 1677 m (cactus), 18 April 1939 (PHS); 2 males, 6 females, ex Neotoma alhigula. Range, 3 km E of Winston, 1677 m (juniper), 21 April 1939 (PHS). Socorro Co.: 1 male, 2 females, ex Neotoma alhigula, 4.8-8 km NW of Mag- dalena, 2403 m (juniper), 23 June 1939 (PHS) (type series); 1 male, ex Neotoma al- higula, 3-6 km NW of Magdalena, 2403 m (juniper), 22 June 1939 (PHS); 1 male, 1 fe- June 1977 Barnes et al.: Anomiopsyllus Fleas 171 Figs. 47-50. Male ninth sternum: 47, A. durangoensia; 48, A. trauhi (redrawn from Barrera 1951 and Barnes 1963); 49, A. novomexicanensis; 50, A. oaxacae. 172 Great Basin Naturalist Vol. 37, No. 2 male, ex Neotoma albigula, 16 km W, 9.6 km N of Magdalena, 2348 m, 6 May 1948 (PHS). Taos Co.: 1 female, ex Neotoma mex- icana, 27 km S of Taos, 1829 m (sage, juni- per), 17 August 1948 (PHS). Valencia Co.: 1 male, 1 female, ex Neotoma nest, 11 km SE of El Morro (lava outcrop), 14 June 1950 (PHS). Utah: Beaver Co.: 2 males, ex Neo- toma sp., 8 May 1945 (PHS); 1 female, ex rodent nest, 16 km W of Beaver, 2134 m, 7 May 1949 (PHS). Garfield Co.: 1 male, 1 fe- male, 6 km S of Panguitch, 2043 m (juni- per), 29 April 1949. Wyoming: Albany Co.: 1 male, 1 female, ex Neotoma cinerea, 10 July 1944 (PHS); 3 females, ex Neotoma sp., 8 July 1943 (PHS). Discussion: Anomiopsyllus n. starki, a no- men nudum described by Barnes in his doc- toral dissertation in 1963, is the same in all morphological characters and measurements as A. n. mexicanus. Anomiopsyllus n. mexi- canus represents a form intermediate be- tween A. n. nudatus and A. n. hiemalis, as indicated by Barnes in his description of A. n. starki. Measurements (males) given by Barnes were based on specimens of A. n. starki. Anomiopsyllus nudatus hiemalis Eads and Menzies n. comb. Figs. 10, 34, 56, 69, 82, 94 Anomiopsyllus hiemalis Eads and Menzies, 1948, J. Kans. Ent. Soc. 21(4): 133-136; Eads and Menzies, 1950, Texas State Hlth. Dept. 53; Jellison, Locker, and Bacon, 1953, Index to the literature of Siphonaptera of North America, Suppl. 1: 11; Vargas, 1960, Med. Rev. Mexicana 40 (849) 1-4; Poorbaugh and Gier, 1961, J. Kans. Ent. Soc. 34(4): 200; Jelhson and Glesne, 1967, Index to the hterature of Siphonaptera of North Amer- ica, Suppl. 2: 13-19; Pratt and Stark, 1973, DHEW Publ. 74-8267: 11. Anomiopsyllus hiemalis hiemalis: Holland, 1965, Ca- nadian Ent. 97: 1055-1056; Lewis, 1974, J. Med. Ent. 11(2): 154. Diagnosis: Males of A. n. hiemalis are distinguished from A. n. nudatus by the wide space between the distal pair of spin- iforms on the movable process (greater than 20 u), and from A. n. mexicanus by the presence of a slender bristle at the arc of curvature of sternum IX. The movable process in hiemalis is shorter than in nu- datus or mexicanus (M = 107 u, range 97 to 112 u), but broader than both (M = 81 u. range 73 to 94 u). The distance between the distal pair of spiniforms closely overlaps that of mexicanus (M = 35 u, range 16 to 42 u), but the two species may be distinguished by the thin bristle on sternum IX. Female characters are not sufficiently dis- tinctive to separate the subspecies of A. nu- datus. Type locality: Gaines County, Texas. Type Iiost: Neotoma micropus Baird Type specimens: Male holotype and fe- male allotype collected on 23 and 9 (re- spectively) April 1948 by V. I. Miles, depos- ited in U.S. National Museum. Material examined: Colorado: Baca Co.: 1 male, 1 female, ex Neotoma albigula, 6 km N of Pritchett, 1341 m, 11 May 1948 (PHS). Pueblo Co.: 1 male, ex Neotoma sp., 6 August 1948 (PHS). Yuma Co.: 1 male, 1 female, ex Neotoma albigula, 1.6 km S, 6 km W of Wray, 1068 m, 10 July 1948 (PHS). Kansas: Barber Co.: 1 male, 1 fe- male, ex Neotoma micropus, 5.6 km S, 1.6 km W of Aetna, 13 April 1949, Loomis and Lipovsky (RM); 18 males, 23 females, ex Neotoma micropus, 8 km S of Aetna, no date, Hopla and Loomis (U.O.); 41 males, 89 females, ex Neotoma micropus, Hopla (U.O.). Clark Co.: 1 male, 1 female, ex Neo- toma micropus, 15 May 1942 (PHS). Gove Co.: 3 females, ex Neotoma sp., 7 Septem- ber 1946 (PHS). Meade Co.: 1 female, ex Neotoma micropus, 12 May 1942 (PHS); 1 female, ex Sigmodon hispidus, 19 km S of Meade, 10 May 1950 (PHS). Seward Co.: 1 male, 1 female, ex Neotoma micropus, 15 May 1942 (PHS). New Mexico: Chavez Co.: 2 males, 2 females, ex Neotoma albigula, 8 km E of Haggerman, 1037 m (mesquite), 13 April 1948 (PHS); 1 male, 1 female, ex Neo- toma albigula, 11 km E of Haggerman, 1158 m, 8 April 1948 (PHS). Eddy Co.: 2 males, 2 females, ex Neotoma albigida, 8 km E of Artesia, 1158 m (mesquite), 24 March 1949 (PHS); 1 female, ex Neotoma albigula, 3 km W of Loco Hills, 1006 m (yucca, mes- quite), 16 February 1950 (PHS). Lea Co.: 1 male, ex Neotoma albigula, 6 km N, 31 km E of Lovington, 1220 m, 25 March 1949 (PHS); 4 males, 4 females, ex Neotoma nest, .8 km N of Hobbs, 1006 m (mesquite), 22 February 1950 (PHS). Oklahoma: Caddo June 1977 Barnes et al.: Anomiopsyllus Fleas 173 Figs. 51-54. Male eight sternum: 51, A. nidiophilus; 52, A. amphibolus; 53, A. montanus; 54, A. sinuatus. 174 Great Basin Naturalist Vol. 37, No. 2 Co.: 2 males, 2 females, ex Neotoma micro- pus nest, April 1958, T. Emerson (U.O.). Herman Co.: 5 males, 16 females, ex Neo- toma micropus nests, April 1960, C. J. Mit- chell (U.O.). Texas Co.: 1 male, ex Neotoma albigula, 9.7 km E of Hardesty, 945 m (yucca, sagebrush), 9 May 1950 (PHS). Texas: Bailey Co.: 1 female, ex Neotoma sp., Muleshoe, 3 March 1957, Simmons. Brew- ster Co.: 1 male, ex Neotoma mexicana, 20 April 1942 (PHS); 2 females, ex rodent nest, Big Bend National Park, 31 October 1963, V. J. Tipton; 1 male, ex Neotoma micropus nest. Big Bend National Park, 2 November 1948, V. J. Tipton; 1 male, 1 female, ex Per- omyscus pectoralis. Big Bend National Park, 10 April 1963, V. J. Tipton. Coke Co.: 3 fe- males, 1 male, ex Neotoma albigula, 29 April 1942 (PHS). Crane Co.: 1 male, ex Neotoma albigula, 29 April 1942 (PHS). Dawson Co.: 1 male, 1 female, ex Neotoma micropus, 11 February 1948 (PHS); 1 fe- male, ex Neotoma micropus, 2 November 1948, Miles. Ector Co.: 2 males, 1 female, ex Neotoma floridana, 26 km W of Odessa, 884 m (mesquite, yucca), 15 April 1960 (PHS). Hartley Co.: 4 males, 3 females, ex Neotoma micropus, October 1944 (PHS). Hutchinson Co.: 3 males, 1 female, ex Neo- toma micropus, 31 May 1944 (PHS); 1 male, 1 female, ex Neotoma micropus, 9.7 km W of Electric City (mesquite), 2 May 1950 (PHS). Jeff Davis Co.: 1 male, 1 female, ex Neotoma mexicana, 2 May 1942 (PHS). Knox Co.: 1 male, 1 female, ex Neotoma mexicana, 16 October 1946 (PHS). Motley Co.: 1 male, 1 female, ex Neotoma albigula, 3 October 1946 (PHS). Pecos Co.: 1 male, ex Dipodomys spectabilis, 21 April 1942 (PHS). Terrell Co.: 1 male, ex Neotoma al- bigula, 1 April 1942 (PHS). Tom Green Co.: 3 males, 1 female, ex Neotoma sp., 11 April 1947 (PHS). Ward Co.: 2 males, 2 fe- males, ex Neotoma albigula, 5 April 1948 (PHS). Mexico: Coahuila D. F.: 2 males, 2 females, ex Neotoma micropus, Sabinas, 20 September 1944, A. Dampf (RM); 1 male, 1 female, same locality, no date, T. Mazzotti (RM); 2 males, 2 females, same locality, 30 August 1944 (PHS); 1 male, 1 female, ex Neotoma nest, same locality, 1 October 1944 (RT). Discussion: Eads and Menzies (1948) in describing A. hiemalis noted the close rela- tionship to nudatus, and differentiated their species primarily by the greater distance be- tween the distal spiniforms of the movable process in hiemalis. They stated that the movable process is ". . . broad and conical." The breadth of the process was considered diagnostic. Stark (1959) reported the existence of atypical nudatus in western Utah, noting the greater distance between spiniforms in his atypical specimens as compared to the typical nudatus. Stark also noted that ster- num IX of males in his atypical specimens varied from typical nudatus, but did not give any details. Anomiopsyllus nudatus mexicanus is intermediate between A. nu- datus nudatus and A. nudatus hiemalis with respect to distance between spiniforms, but distinct from either by virtue of several modified setae dorsomedially on the posteri- or margin of male sternum IX. It is also in- termediate geographically, its range falling between the eastern hiemalis and the west- ern nudatus. In regard to length and breadth of the movable process, A. n. mexi- canus is more like nudatus than like hie- malis. The process of mexicanus is appar- ently longer in relation to width than that of hiemalis, thus less "conical." Exhaustive examination of specimens available for study has not resulted in find- ing characters other than those mentioned. The morphology of the populations is sub- ject to remarkably little variation except in the male external genitalia. Other possible characters thoroughly investigated were: se- tation of head, thorax, and abdomen; shape of thoracic segments and setation of legs; potential difference in proportion of tarsal segments; and both male and female gen- italia. No other consistent variations were found. The females are apparently not sepa- rable, although a trend exists toward a deeper sinus in the posterior margin of ter- gum VIII of A. n. hiemalis. The sinus is used as a key character for hiemalis by Hopkins and Rothschild (1962). To examine the relationships of the three groups, a series of measurements was taken as follows: (1) width of the movable process June 1977 Barnes et al.: Anomiopsyllus Fleas 175 Figs. 55-57. Male eight sternum: 55, A. nudatus nudattis; 56, A. ntidatus hiemalis; 57, A. nudatus mexicanus. 176 Great Basin Naturalist Vol. 37, No. 2 from the "notch" above the point of articu- lation across the broadest portion of the lower lobe; (2) length of the process from its tip to the "notch"; (3) distance from cen- ter to center between the spiniforms. The manner in which measurements were made is illustrated in Figure 93. The measure- ments were taken with a calibrated ocular grid, using lOX eyepiece and 43X objective with grid units converted to microns (0.1 grid units = 1.624 u). Due to the allopatric nature of the popu- lations, mexicanus, the geographically cen- tral population, was compared first to hie- rnalis and then to nudatus. If each of the two populations is related to mexicanus, then they are related to one another. Anomiopsylliis n. mexicanus and hiemalis are closely linked by the overlap of dis- tances between spiniforms, yet separated by length and width of the movable process. Anomiopsylliis n. mexicanus and nudatus are sharply distinguished by distance be- tween spiniforms, but with significant over- lap in length of process, and they almost completely overlap in width of process. Anomiopsylliis n. mexicanus is further dis- tinguished from both hiemalis and nudatus by presence of modified setae (Figs. 24, 25) of sternum IX in males. The A. nudatus complex ranges from western Kansas, Oklahoma, and Texas south to Sabinas, Coahuila, Mexico, and west to the Pacific Coast of southern California. The subspecies A. nudatus nudatus, the westernmost representative of the three sub- specific populations, has a primarily Sono- ran Desert distribution (Shreve and Wiggins 1951; Map 2). Its range is not limited to the Sonoran Desert, but occurs in a variety of habitats occupied by its wood rat hosts; e.g., in pine-fir forest at 21.34 m elevation in the San Bernardino Mountains of southern Cali- fornia. In southern California, A. n. nudatus is found in coastal chaparral and oak wood- land bordering the sea as well as in its typi- cal desert habitat. In the Santa Monica Mountains and parts of the San Gabriel Mountains of Los Angeles County, it is re- placed by A. falsicalifornicus in moister oak woodland and chaparral situations. In the Los Angeles area, the distributions of A. n. nudatus and A. falsicalifornicus inter- digitate: nudatus occurs in drier areas where yucca and cactus grow, and fahicali- fornicus occurs in moister areas character- ized by Rhus diversiloba and Photinia. This situation continues northward to Ventura County, California, where nudatus occupies the arid habitat in the rain shadow south and east of the southern Santa Ynez Moun- tains. From Ventura County eastward, nudatus is recorded from the area of Frazier Park, Kern Coimty, and the Antelope Valley of northern Los Angeles County. It is replaced above the lower, winter snow line by A. walkeri in the Tehachapi and southern Sierra Nevada Mountains. From this point it occurs across the Mojave and Sonoran deserts, its distribution including the south- west comer of Nevada and Sonoran Desert areas of Arizona. Areas where the distribution of n. nu- datus is doubtful include the northern por- tions of the Mojave Desert and arid areas on the west side of the southern San Joa- quin Valley in California where collections of wood rat nests made by the senior author have been negative for Anomiopsylliis. Sev- eral females, collected by R. Ryckman from the San Pedro Martir Mountains of Baja California, are probably A. n. nudatus. A collection of 20 wood rats {Neotoma alhi- gula) near Hermosillo, Sonora, Mexico, by K. Murray and H. Johnson during March and April, 1960, failed to produce Anomio- psylliis. Anomiopsylliis n. mexicanus was collected in central Mexico by J. E. H. Martin during July 1964 from a nest of Neotoma sp. In Arizona, the distribution of A. n. nu- datus is sharply interrupted along the es- carpment of the Colorado Plateau by the presence of a central subspecies, A. n. mexi- canus. In San Juan County, southeastern Utah, A. n. nudatus makes a deep incursion into the area occupied by A. n. mexicanus (Map 2), a situation further discussed under that subspecies. The distribution of A. nudatus mexicanus lies within the Basin and Range Province as defined by Shreve and Wiggins (1951). In June 1977 Barnes et al.: Anomiopsyllus Fleas 177 Figs. 58-61. Male eight sternum: 58, A. falsicalifornicus congruens; 59, A. walkeri; 60, A. fahicalifornicus calif ornictis; 61, A. martini. 178 Great Basin Naturalist Vol. 37, No. 2 Arizona, its boundaries coincide with the edge of the Sonoran Desert and the occur- rence of A. nudatus nudatus. In the east, its range is interrupted by the western edge of the Great Plains (High Plains, according to Webb 1951) and the presence of A. n. hiemalis. Its northern range is indented by the central Rocky Mountain mass. A narrow band just east of the Rockies carries the subspecies through far eastern Colorado and western Kansas to Morrill County, Nebras- ka, and Albany County, Wyoming. In south- central New Mexico, it is absent from the lower Chihuahuan Desert habitat, where it is replaced by A. novomexicanensis, as A. n. mexicanus appears to occupy the higher elevations. In Utah, a confusing picture is presented by the occurrence of nudatus nudatus in San Juan County in the southeastern corner of the state. Stark (1959) reported atypical nudatus from Millard, Beaver, and Garfield counties in Utah and typical nudatus in San Juan County. Examination of his specimens shows the former to be n. mexicanus and the latter n. nudatus. It is suspected that barriers such as the Colorado River in the west and discontinuous wood rat distribu- tion to the east may explain the disjunct distribution observed in n. nudatus. In San Juan County, only n. nudatus has been col- lected; however, west of the Colorado River in Utah, only n. mexicanus has been collect- ed. Two of the San Juan n. nudatus collec- tions are from high elevations in habitats similar to those preempted by n. mexicanus along the Colorado Plateau escarpment where the two subspecies are known to meet. Collections in the rugged area of southcentral Utah and northcentral Arizona are needed to clarify the n. mexicanus dis- tribution. Anomiopsyllus n. hiemalis replaces A. n. mexicanus at lower elevations at the eastern and southwestern edge of the n. mexicanus distribution. The distributions of the two subspecies interdigitate where plains are in- tersected by spurs of the Rockies (see no- vomexicanensis for further discussion). Inter- grades between n. mexicanus and n. hiemalis occur in Santa Fe and San Miguel counties. New Mexico, and specimens as- signable to each subspecies have been taken in the same area in Santa Fe County by U.S.P.H.S. field workers. In eastern New Mexico and Colorado where apparent over- lap occurs, n. hiemalis occupies the habitat below 1341 m elevation and n. mexicanus occupies the higher elevations. The range of A. n. hiemalis falls primari- ly into two biotic areas defined by Webb (1951) as the High Plains and Trans-Pecos areas and by Dice (1943) as the Kansan and Tamaulipan. These areas are characterized by moderate elevation (generally below 1220 m) and low annual rainfall (50 cm or less). Where vegetation type has been noted on n. hiemalis slides examined, mesquite {Prosopis sp.) predominates. Geographically, the area occupied by n. hiemalis includes southwestern Kansas, western Oklahoma and Texas, extreme eastern Colorado, and New Mexico south to Sabinas, Coahuila D. F., Mexico. The northernmost record noted is Yuma County, Colorado, and the eastern- most are Barber County, Kansas, and Caddo County, Oklahoma. As was speculated earlier for n. nudatus, the ranges of all three subspecies probably extend south into Mexico. The lack of speci- mens from Mexico precludes an accurate es- timate, but the habitats occupied in the United States also occur in Mexico. Shreve and Wiggins (1951) shows Sonoran Desert, the habitat occupied by n. nudatus, as oc- curring west of the Sierra Madre Occidental as far south as the Rib Yaqui. The upland habitat preferred by n. mexicanus extends south along the continental divide (Brand 1936, Leopold 1950). The mesquite scrub, characteristic of n. hiemalis habitat in Texas, extends south into eastern Mexico. The apparent replacement of one sub- species by another with great changes in to- pography and habitat is strongly suggestive of similar phenomena noted with other Anomiopsyllus species. In California, where extensive field collecting was done in this study, A. nudatus was found to be replaced by A. walkeri at higher elevations in the Tehachapi Moimtains and by A. falsicalifor- nicus in more mesic environments (i.e., rain- fall over 50 cm /year). The phenomenon of displacement may be understood ultimately June 1977 Barnes et al.: Anomiopsyllus Fleas 179 Figs. 62-64. Male eight sternum: 62, A. novomexicanensis; 63, A. durangoensis; 64, A. oaxacae. 180 Great Basin Naturalist Vol. 37, No. 2 Figs. 65-67. Aedeagus: 65, A. sinuatus; 66, A. montanus; 67, A. nidiophilus. June 1977 Barnes et al.: Anomiopsyllus Fleas 181 Figs. 68-70. Aedeagus: 68, A. nudatus nudatus; 69, A. nudatus hiemalis; 70, A. nudatus mexicanus. 182 Great Basin Naturalist Vol. 37, No. 2 through laboratory studies or the range of tolerance for physical climate factors or by further field studies involving several of the species. The response to climate is not the only limiting factor involved in distribution of AnomiapsyUus species. This is illustrated by those species or subspecies that are ca- pable of occupying a broad spectrum of cli- matic conditions in the absence of related species or subspecies. Anomiopsijllus n. nu- datus occurs at elevations up to 2195 m in California and is displaced by A. n. mexi- canus at higher elevations in Arizona and A. walkeri in California as noted above. Hosts: Anomiopsijllus nudatus subspecies are restricted to wood rats {Neotoma spp.) as major hosts and seldom occur on other rodents. In the material examined, fortuitous hosts are probably more numerous than nor- mally occur in nature. Many of these re- cords result from plague surveys in which most of the specimens were ground and in- oculated into test animals, and collectors of- ten tended to retain specimens from odd hosts for the record. Within the habitats occupied by A. nu- datus, its subspecies occur indiscriminately on available wood rat species, with the ex- ception of Neotoma cinerea which Goldman (1910) and Hall and Kelson (1959) place in a separate subgenus, Teonoma. Neotoma floridana is primarily a resident of moist eastern deciduous forest communities. Its distribution extends through western Kansas and northeastern Colorado where it oc- cupies situations characteristic of N. albi- gula such as cactus huts (Warren 1942). In Colorado, Finley (1958) has recorded Ano- miopsijllus sp. from N. floridana, but his specimens have not been available for study. On the other hand, Rainey (1956), and Ellis (1955) did not record Anomiop- sijllus from N. floridana in eastern Okla- homa. The range of A. nudatus or any of its subspecies never coincides with the range of any one wood rat species. Thus, in southern California, nudatus nudatus occurs on both N. lepida and N. fuscipes in arid situations, while in less arid areas the same two hosts are parasitized by different Anomiopsijllus species. In Arizona and New Mexico the range of N. mexicana appears to limit the range of A. n. mexicanus, but the N. mexi- cana range is exceeded by the range of A. n. mexicanus in western Utah where the flea occurs on N. lepida. Within the range of the host genus Neotoma, A. nudatus sub- species are apparently limited by factors other than host occurrence as are other Anomiopsijllus spp. Anomiopsijllus falsicalifornicus Fox Figs. 14, 15, 37, 40, 58, 60, 71, 72, 85, 86, 95 Anomiopsijllus californicus: Fox, 1926 {nee Baker, 1904), Pan-Pacific Ent. 2: 183; Wagner, 1930, Mag. Parasitol. Moscow 1: 135. Anomiopsijllus fal-sicalifornieus Fox, 1929, Ent. News 40: 218; Collins, 1936, Ent. News 47: 130; Hubbard, 1947, Fleas of western North America: 78-80; Linsdale and Davis, 1956, Univ. Calif. Publ. Zool. 54: 306-308; Holland, 1965, Canadian Ent. 97: 1052; Barnes, 1965, Pan-Pacific Ent. 41(4): 278; Coultrip et al., 1973, J. Med. Ent. 10(3): 306. Diagnosis: Anomiopsijllus falsicalifornicus is separable from other species (males) by the suboblique row of three spiniform setae near the apex of the movable process coupled with the presence of only 0-3 spin- iforms at the apex of the distal arm of ster- num IX. The movable process is roughly triangular with always some degree of con- cavity (shallow to extreme) on the posterior margin below the distal spiniform row. The anterior margin is linear, tipped slightly an- terad at the subacute apex with three heavy spiniforms near the apex with or without a small spiniform near the posterior margin of the basal lobe. The proximal arm of ster- num IX is similar to nudatus. The distal arm is variable in shape and the apex rounded or flattened with 0-3 heavy setae at the apex and a row of 4-6 moderately heavy setae mesad on the posterior margin. Females are similar to nudatus but can be separated by the presence of the sclero- tized portion of the blind ductus seminalis (Fig. 72, 76). Female characters are not suf- ficiently distinctive to separate subspecies of fa hica liforn icus. Average Length: Male 1.6 mm; female 1.9 mm. Type locality: Claremont, Los Angeles Countv, California June 1977 Barnes et al.: Anomiopsyllus Fleas 183 Figs. 71-72. Aedeagus: 71, A. walkeri; 72, A. martini. 184 Great Basin Naturalist Vol. 37, No. 2 Type host: Neotoma fuscipes and Spilo- gale phenax Type specimens: Male holotype deposited in the U.S. National Museum. Fox (1926) first described the male of this species, believing it to be A. californicus Baker, a species that had been described from near Claremont, California. Jordan and Rothschild (1915) examined Baker's syntypic series of females and entered californicus into synonomy with nudatus. Fox (1929), acting on information from Jordan, then gave his male specimen the name falsicali- fornicus. Anomiopsyllus fahicalifornicus is treated here as two subspecies: A. /. fahicalifor- nicus, A. /. congruens. Anomiopsyllus fahicalifornicus congruens Stewart Figs. 15, 40, 58, 71, 85, 95 Anomiopsyllus congruens Stewart, 1940, Pan-Pacific Ent. 16: 17-18; Hubbard, 1940, Pac. Univ. Bull. 37(4); Hubbard, 1943, Pac. Univ. Bull. 39(8): 1-12; Augustson, 1943, Bull. South. Calif. Acad. Sci. 42:69; Hubbard, 1947, Fleas of western North America: 81-82; Jellison, Locker, and Bacon, 1953, Index to the literature of Siphonaptera of North America, Suppl. 1: 10; Linsdale and Davis, Univ. Calif. Publ, Zool. 54(5): 306-308; Hol- land, 1965, Canadian Ent. 97: 1052, Barnes, 1965, Pan- Pacific Ent. 41(4): 278. Anomiopsyllus fahicalifornicus congruens Hopkins, 1952, J. Wash. Acad. Sci. 42(11): 365; Holland 1965, Can. Ent. 97: 1053; Jellison and Glesne, 1967, Index to the literature of Siphonaptera of North America, Suppl. 2-13: 19; Lewis 1974, J. Med. Ent. 11(2): 154. Diagnosis: The males of A. /. congruens are distinguished from A. /. fahicalifornicus by the narrow sternum IX with an angulate, truncate apex. The fixed process of the clas- per is broader than long. Sternum IX is only one-third to one-half as broad mesally as in /. fahicalifornicus and is equipped apically with 1, 2, or 3 heavy setae. The movable process is similar to /. fahicalifor- nicus but possesses a proximal spiniform. Female characters are not sufficiently dis- tinctive to separate subspecies of fahicali- fornicus. Type locality: Frances Simes Hastings Natural History Reservation near James- burg, Monterey County, California. Type host: Spilogale gracilis ssp. Type specimens: Holotype male and allo- type female collected on 19 November 1938 by R. Holdenried, deposited in Mu- seum of the California Insect Survey, Univ. of California, Berkeley, California. Recorded distribution: California: Mon- terey Co. (type locality), San Luis Obispo Co. (Linsdale and Davis). New distribution: From Santa Ynez Mountains of Santa Barbara Co. north to Siskiyou Co., thence south to the western Sierra Nevada foothills to Tulare Co., Cali- fornia. Material examined: California: Alameda Co.: 1 male, 1 female, ex Neotoma fuscipes, Oakland (oak woodland), 1 July 1936 (RM); 27 males, 43 females, ex Neotoma fuscipes nests, Berkeley (oak woodland), 17 October 1959, Barnes (BVC). Amador Co.: 1 male, 3 females, ex Neotoma fuscipes nest, 1.6 km N, 5 km E of Plymouth (oak woodland), 14 January 1959, Barnes (BVC). Contra Costa Co.: 3 males, 7 females, ex Neotoma fus- cipes nests, Orinda (oak woodland), 18 Oc- tober 1959, Barnes (BVC). El Dorado Co.: 27 males, 43 females, ex Neotoma fuscipes nest, Placerville vicinity (oak woodland), many dates, Barnes (BVC); 110 males, 110 females, ex Neotoma fuscipes nest. Shingle Springs vicinity, many dates, Barnes (BVC). Fresno Co.: 4 males, 10 females, ex Neo- toma fuscipes nest, 3 km N, 2 km W of Au- berry (oak woodland), 7 January 1958, Barn- es (BVC); 2 males, 6 females, ex Neotoma fuscipes nest, 1.6 km SW of Prather (oak woodland), 13 November 1958, Barnes (BVC). Lassen Co.: 1 male, 4 females, ex Neotoma fuscipes nest, 5 km NW of Little Valley (oak, Ceanothus, sagebrush), 6 May 1960, Barnes (BVC). Monterey Co.: 1 fe- male, ex Neotoma fuscipes, Hastings Nat. Hist. Res., 14 July 1938 (RM); 1 male, 2 fe- males, ex Neotoma fuscipes, 4 km E of Car- mel, 21 December 1948, Holm (BVC); 1 fe- male, ex Eutamias merriami, Hastings Nat. Hist. Res., 8 February 1947 (PHS); 1 male, ex Perognathus californicus, same locality, 18 January 1947 (PHS). Napa Co.: 1 male, ex Neotoma fuscipes, 4 km N, 2 km E of Calistoga (oak woodland), 24 October 1956, Murray (BVC). Nevada Co.: 3 males, 9 fe- males, ex Neotoma fuscipes nest, 19 km S of Grass Valley (oak woodland), 15 January June 1977 Barnes et al.: Anomiopsyllus Fleas 185 Figs. 73-74. Aedeagus: 73, A. falsicalifornicus congruens; 74, A. falsicalifornicus falsicalifo 186 Great Basin Naturalist Vol. 37, No. 2 Figs. 75-77. Aedeagus: 75, A. durangoensis; 76, A. novomexicanensis; 77, A. oaxacae. June 1977 Barnes et al.: Anomiopsyllus Fleas 187 Figs. 78-81. Female seventh sternum and spermatheca: 78, A. nidiophilus; 79, A. sinuatus; 80, A. amphiboltis; 81, A. montanus. 188 Great Basin Naturalist Vol. 37, No. 2 1959, Barnes (BVC). Plumas Co.: 17 males, 33 females, ex Neotonia fuscipes nest, 3 km W of Virginia, 17 October 1961, Barnes. San Benito Co.: 2 males, ex Neotoma fus- cipes, T14S R6E Section 2, 13 February 1951, Kirkwood (BVC). San Francisco Co.: 1 male, ex Spemiophilus beecheyi, San Fran- cisco (laboratory), no date (PHS). San Luis Obispo Co.: 7 males, 19 females, ex Neo- toma fuscipes nest, 5 km N, 7.2 km E of Santa Margarita (oak woodland), 2 Novem- ber 1956, Barnes (BVC); 1 male, 5 females, ex Neotoma fuscipes, 1.6 km N, 6 km E of Santa Margarita (oak woodland), 15 June 1956, Barnes (BVC); 1 male, 1 female, ex Neotoma fuscipes, same locality (oak wood- land), 5 March 1957, Barnes (BVC); 1 male, 5 females, ex Neotoma fuscipes, 1.6 km N, 6 km E of Santa Margarita (oak woodland), 1 November 1956, Murray (BVC); 1 male, 6 females, ex Neotoma fuscipes nest, 5 km N of Pozo (chaparral, woodland), 20 Novem- ber 1956, Barnes (BVC); 3 males, 1 female, ex Neotoma fuscipes. La Panza Camp (chaparral), 3 June 1955, Murray (BVC); 1 male, 1 female, ex Neotoma fuscipes, San Luis Obispo, 1 June 1935 (RM); 1 male, ex Peromyscus boylei, 1.6 km N, 4 km E of Santa Margarita (oak woodland), 15 June 1956, Murray (BVC); 1 male, 2 females, ex Peromyscus truei, same locality (oak wood- land), 15 June 1956, Murray (BVC); 1 male, ex Peromyscus truei, 3 km N, 6 km E of Pozo (chaparral), 7 March 1957, Barnes (BVC); 1 female, ex Microtus californicus, 3 km E of Cayucos, 1 June 1947, Ryan and Durham (HF). San Mateo Co.: 1 female, ex Neotoma fuscipes. La Honda, 26 March 1949, Prince (PHS). Santa Barbara Co.: 3 males, 4 females, ex Neotoma lepida. Buck- horn Canyon, 6 km E of Santa Maria (chap- arral), 24 April 1957, Barnes, Murray (BVC); 1 male, ex Neotoma fuscipes, Lom- poc (oak woodland, chaparral), 9 February 1961, Rohe (BVC); 31 males, 53 females, ex Neotoma fuscipes nests, same locality, 9 February 1961, Rohe (BVC); 11 males, 18 females, ex Neotoma fuscipes nests, 47 adults reared from larvae 9 km N, 11 km W of Santa Barbara (oak, chaparral), 8 May 1961; 43 males, 74 females, ex Neotoma fus- cipes nests, San Marcos Pass, 12 km N, 13 km W of Santa Barbara (oak woodland), 8 May 1961, Barnes. Santa Clara Co.: 1 male, 1 female, ex Neotoma fuscipes, Los Gatos, 24 June 1936 (RM). Santa Cruz Co.: 2 males, 1 female, ex Neotoma fuscipes nest, Boulder Creek, 305 m, 30 May 1939, Good (BVC); 1 male, 5 females, ex Neotoma fus- cipes, same locality, 4 July 1940, Good (PHS). Siskiyou Co.: 2 males, 7 females, ex Neotoma fuscipes nest, 5 km W of Dorris (oak, pine), 6 April 1960, Barnes (BVC); 3 males, 6 females, 8 km S, 10 km W of Ga- zelle (juniper, oak), 2 April 1960, Barnes (BVC). Tulare Co.: 1 male, 2 females, ex Neotoma fuscipes nest, 1 km N, 5 km E of Lemon Cove (oak woodland), 2 April 1959, Barnes (BVC). Tuolumne Co.: 8 males, 17 females, ex Neotoma fuscipes nests. La Grange, 11 March 1958, Barnes and Keh (BVC). Discussion: Stewart (1940) described A. congruens as a full species related to A. fal- sicalifornicus. Hopkins (1952) reduced con- gruens to subspecific status under fakicali- fornicus, but his reasons weren't given. Linsdale and Davis (1956) presented a sub- stantial case for full specific status for A. congruens, demonstrating constant mor- phological differences between the two forms. Hopkins and Rothschild (1962) held to the view of Hopkins (1952) and kept congruens at the subspecific level. The view of Hopkins is amply supported by morphological, distributional, and repro- ductive data from this study. This, with the perspective gained from study of the genus as a whole, calls for subspecific assignment of congruens. Morphologically, males of A. /. congruens and A. /. falsicalifornicus are immediately separable by the shape of the distal arm of sternum IX. In all of the /. congruens exam- ined, this structure was narrow and ang- ulate, truncate apically. In Anomiopsyllus f falsicalifornicus, this structure is broader and gently rounded at the apex (Fig. 31). In the south coastal area of California where distributions of the two forms nearly meet, sternum IX of /. congruens has two large spiniform setae at the apex and /. falsicali- fornicus has only one spiniform of moderate size with occasionallv a small tacklike seta June 1977 Barnes et al.: Anomiopsyllus Fleas 189 Figs. 82-84. Female seventh sternum and spermatheca: 82, A. nudatus hiemalis; 83, A. nudatus mexicanus; 84, A. nudatus nudatus. 190 Great Basin Naturalist Vol. 37, No. 2 immediately below it. In the central Sierra Nevada Mountains populations of /. con- gruens, the number of apical spiniforms on this structure is reduced until, finally, in Tulare County, there are none in some local populations. Specimens with three spin- iforms on the finger are occasionally found in more northern coastal populations. The shape of sternum IX remains constant. Since Fox's (1926) description of fahicali- fornicus and Stewart's (1940) description of congruens, the shape of the movable process has been considered diagnostic for the two groups. Stewart's description of congruens emphasized the lack of an apparent distal dilation of the process that he had observed in fahicalifornicus. In reahty, the apparent dilation is due to a broad concavity of the posterior margin of the movable process, below which there is a broad, lightly sclero- tized lobe (Figs. 12, 16). This concavity oc- curs to some degree in both /. congruens and /. fahicalifornicus. There is a tendency for more concavity in the process of /. fahi- californicus, but some specimens of /. con- gruens are as deeply concave. A number of authors have erred in drawing and describ- ing this structure, including Fox (1926), Stewart (1940), and Hubbard (1947). These authors failed to include the proximal lobe, an oversight that changes the appearance of the structure considerably. Linsdale and Davis (1956) simply did not illustrate its lower portion. Wagner (1936) illustrated the structure correctly, as did Collins (1936) in his supplementary description of fahicalifor- nicus. The overlap in shape of the movable process reduces its importance as a tax- onomic character. The number and placement of the distal spiniforms were considered to be important by Linsdale and Davis (1956) in dis- tinguishing these two forms. In 73 speci- mens of /. congruens from the type locality, 36 had a full complement (three distal, one proximal on both sides), 19 had one prox- imal missing on one side, 7 had one prox- imal moved up near the distal row, and 11 had no proximal spiniforms at all. A series of 17 /. fahicalifornicus from Los Angeles had no proximal spiniforms. Of 54 males of /. fahicalifornicus examined in the present study, 7 had proximal spiniforms on at least one side and the remainder had none. Of the many /. congruens examined, 72 percent had at least one proximal spiniform and 63 percent had proximal spiniforms on both sides. Only an occasional specimen (1 per- cent) had one or more spiniforms moved up to or near the distal row. The proximal spiniform is evidently in the process of being lost in both forms, but the process has progressed further in /. fahicalifornicus than in /. congruens. Its presence in some /. fahicalifornicus and absence in some /. con- gruens reduces its value as a character. The fixed process of the clasper is used by Linsdale and Davis (1956) and by Hop- kins and Rothschild (1962) to separate fahi- californicus and congruens. In material ex- amined in the present study, the general shape was found subject to variation. That of /. fahicalifornicus is always longer than broad, and that of /. congruens broader than long. In females, sternum VII varies in contour within populations both in regard to breadth of the ventral lobe and depth of the posterodorsal sinus. The sinus varies from prominent in some specimens to ab- sent in others. The range of variation is completely overlapping between the two subspecies. The posterior margin of the female ter- gum VIII has been used to separate species by Hopkins and Rothschild (1962). In /. fal- sicalifornicus, there is considerable variation in its shape. Specimens of /. congruens from coastal California may or may not possess a dorsal lobe. Often there is no production of the posterodorsal margin, and rarely does this lobe exceed the ventral production of tergum VIII. Consequently, the sinus be- tween these lobes is either very shallow or absent. Specimens of /. congruens from the Sierra Nevada foothills show a definite, broad posterodorsal lobe of tergum VIII which extends beyond the ventral portion of the tergum forming a broad, shallow sinus. Females of /. fahicalifornicus from Los An- geles County differ only slightly from coast- al /. congruens in this respect. One of the oddities found by the study of many specimens, particularly /. congruens. June 1977 Barnes et al.: Anomiopsyllus Fleas 191 Figs. 85-88. Female seventh sternum and spermatheca: 85, A. fahicalifornicus congruens; nicus falsicalifornicus; 87, A. walkeri; 88, A. martini. A. fahicalifor- 192 Great Basin Naturalist Vol. 37, No. 2 is that specimens from more humid, coastal areas (e.g., Santa Margarita, San Luis Obispo County) are darker, i.e., more heavi- ly pigmented. This character is also found in the wood rat hosts (Hooper 1938) and is used here as a subspecific character, but it has little value as a taxonomic character in fleas. Anomiopsyllus falsicalifornicus falsicalifornicus Fox Figs. 14, 37, 60, 72, 86, 95 Anomiopsyllus californicus: Fox, 1926 (nee Baker, 1904), Pan-Pacific Ent. 2: 183; Wagner, 1930, Mag. Parasitol. Moscow 1: 35. Anomiopsyllus falsicalifornicus: Fox, 1929, Ent. News 40: 218; Collins 1936, Ent. News 40: 137; Hub- bard, 1947, Fleas of western North America: 78-80; Jel- lison, Locker, and Bacon, 1953, Index to the literature of Siphonaptera of North America, Suppl. 1: 10; Lins- dale and Davis, 1956, Univ. Calif. Publ. Zool. 54(5): 306-308; Holland, 1965, Canadian Ent. 97: 1052; Barnes, 1965, Pan-Pacific Ent. 41(4): 278; Jellison and Glesne, 1967, Index to the literature of Siphonaptera of North America, Suppl. 2: 13-19; Coultrip et al., 1973, J. Med. Ent. 10(3): 306. Anomiopsyllus falsicalifornicus falsicalifornicus Hop- kins, 1952, J. Wash. Acad. Sci. 365; Knudsen, 1956, Bull. South. Calif. Acad. Sci. 55(1): 1-6; Hopkins and Rothschild, 1962, Catalogue of the Rothschild Collec- tion of Fleas III: 402-403; Holland, 1965, Canadian Ent. 97: 1053; Lewis 1974, J. Med. Ent. 11(2): 154. Diagnosis: The males of A. falsicalifor- nicus falsicalifornicus are distinguished from A. /. congruens by the shape of the distal arm of sternum IX and the fixed process of the clasper being longer than broad. Ster- num IX is two to three times as broad me- sally as in /. congruens and has the apex gently rounded and equipped at its posteri- or margin near the apex with one or two stout setae. The movable process is similar to /. congruens but lacks the proximal spin- iform setae. Female characters are not sufficiently dis- tinctive to separate subspecies of falsicali- fornicus. Type data: That of the species. Recorded distribution: California: Clare- mont, Los Angeles County, west to Beverly Glen, Los Angeles County. Material examined: California: Kern County: 3 males, 4 females, ex Neotoma le- pida, Poso Creek, 3.5 km S, 8.8 km W of Granite Station, 23 May 1957, (BVC 1781). Los Angeles Co.: 6 females, ex Neotoma fuscipes, Sepulveda Canyon, Los Angeles, 11 November 1941, G. Augustson (HF41- 148c); 3 males, 7 females, ex Neotoma fus- cipes, Beverly Glen, June 29, 1941, G. Au- gustson (HF41-45 GA); 4 females, ex Neo- toma fuscipes, Los Angeles, 27 April 1941, G. Augustson (HF41-45 GA); 1 female, ex Neotoma fuscipes, same locality, 16 Novem- ber 1940, G. Augustson (HF40-146 GA); 3 males, 3 females, ex Neotoma fuscipes, Se- pulveda and Mission Roads, 5 May 1959, D. Rohe (BVC 15754); 2 males, 3 females, ex Neotoma fuscipes, 4 km N of Bel Air, 17 November 1959, Murray and Rohe (BVC 14703); 17 males, 27 females, ex Neotoma fuscipes nests, plus 33 females, 14 males reared from larvae, Sepulveda and Mission Road, 28 March 1962, D. Rohe; 1 male, 4 females, ex Neotoma fuscipes nest, Malibu Creek Cornell, 9 May 1961, A. Barnes; 11 males, 19 females, ex Neotoma fuscipes nests, San Gabriel Canyon, 13.2 km N of Azusa, 17 December 1956, A. Barnes. Anomiopsyllus walkeri Barnes Figs. 13, 38, 59, 74, 87, 94 Anomiopsyllus walkeri Barnes, 1965, Pan-Pacific Ent. 41(4): 272-280; Lewis, 1974, J. Med. Ent. 110: 154. Diagnosis: Barnes (1965) indicated that A. walkeri was related to A. nudatus, but may be distinguished on the basis of four spin- iforms on the movable process of the clas- per with three on the distal portion and one on the proximal portion. Since Holland's de- scription of A. martini, it would appear that walkeri would be more closely related to martini than nudatus. The aedeagus of A. ivalkeri differs markedly from nudatus in that walkeri has a heavily sclerotized dorsal margin and a well-defined dorsomedian "hump" above and slightly anterad to the aedeagal struts. The neck of the aedeagus curves ventrad from the longitudinal axis and the apodeme curves dorsad. Female characters are not sufficiently dis- tinctive to separate walkeri from nudatus and martini. Type locality: Male holotype, female allo- type from 11 km S, 9 km E of Tehachapi, Kern Co., Calif., ex Neotoma fuscipes nest, 19 February 1959 (Barnes). Designated June 1977 Barnes et al.: Anomiopsyllus Fleas 193 Figs. 89-92. Female seventh sternum and spermatheca: 89, A. durangoensis; 90, A. novamexicanensis, 91, A. traubi (redrawn from Barrera 1951 and Barnes 1963); 92, A. oaxacae. 194 Great Basin Naturalist Vol. 37, No. 2 paratypes: California: Kern Co., 7 males, 19 females, 19 February 1959, 2 females, 2 April 1959; 22 males, 30 females, 22 Janu- ary 1960; 1 male, 8 females 10 km SE of Tehachapi, 19 February 1959; 7 males, 8 fe- males, 3 km NW of Keene, 18 February 1959; 2 males, 3 females, 3-6 km SW of Glenville, 17 February 1959; 2 males, 1 fe- male, 3 km N, 1 km W of Weldon, 16 March 1960; 5 males, 13 females, 5 km E of Lake Isabella, 18 March 1960. Tulare Co., 2 males, 2 females, 8 km S, 10 km W of Little Lake (Inyo Co.). Type host: Neotorna fuscipes Type specimens: The male holotype and female allotype are in the U.S. National Museum. Paratypes are in the following col- lections: University of California, Berkeley; Bureau of Vector Control, California De- partment of Public Health, Berkeley, Cali- fornia; British Museum (Natural History), Tring, Hertfordshire; Rocky Mountain Lab- oratory, Hamilton, Montana; U.S. Public Health Service Field Station, San Francisco, California; private collection of Dr. Robert Traub, Bethesda, Maryland. Discussion: Barnes (1965) draws attention to the close morphological and distribution- al relationships between A. walkeri and A. nudatus, but provides ecological data which supports retention of walkeri as a full spe- cies. AnoiniopsyUus walkeri replaces A. nii- dutus in the Tehachapi and southern Sierra Nevada Mountains of California. This type of replacement is typical of AnomiopsyUiis species distribution, with no instance known of two Anomiopsyllus species inhabiting the same nest, or even the same wood rat colo- ny. The area of contact between the two spe- cies was examined with this in mind. Spe- cial attention was given to the possibility of morphological intergradation and to the habitats in which the two species were found. Substantial numbers of wood rat {N. fuscipes) nests were collected from the type locality and adjacent areas during the win- ter and spring of 1959 and 1960. Fleas were removed from nests by means of a modified Berlese funnel and were examined. Habitat notes were taken along with notes on weather conditions at the time of collection. The type locality of A. walkeri is on the east slope of Double Mountain, the eastern- most peak of the Tehachapi chain, and is at 1463 m to 1585 m. The dominant vegeta- tion consists of juniper {Juniperus occiden- talis) and Joshua tree {Yucca brevifolia) woodland, sagebrush {Artemisia sp.) and sev- eral species of Eriogonum. This vegetation characterizes a desert habitat in that area. Such habitats usually have some persistent snow (at least in patches) during midwinter. The average annual rainfall is approx- imately 30 cm. In the south, alluvial fans slope into Antelope Valley. The plain, which is relatively flat, slopes gradually from about 1067 m in the west to 762 m in the south and east. The valley is largely un- der cultivation, but on the alluvial fans and in the remaining natural areas on the valley floor juniper-Joshua tree woodland also ex- ists as habitat for N. fuscipes along with creosote bush {Larrea tridentata), a shrub characteristic of moderate elevation and gentle slopes in the Mojave Desert. Rare winter snows do not persist even for a day in Antelope Valley or on the slopes of Double Mountain below 1220 m. Average annual precipitation is 13 cm at Lancaster and 16 cm at Mojave, two nearby weather stations on the valley floor. Nest collections were made at and near the type locality on slopes at lower eleva- tions and in Antelope Valley on 19 Febru- ary 1959; 1-2 April 1959 and on 22 January 1960. A total of 51 nests was collected, with 34 containing fleas, of which 24 con- tained Anomiopsyllus spp., either walkeri or nudatus. Anomiopsyllus walkeri alone was removed from 11 of 22 nests collected at 1402 m or above in juniper-Joshua tree woodland. Anomiopsyllus nudatus alone was removed from 13 of 29 nests collected in juniper-Joshua tree-creosote bush at 1280 m or below. A total of 31 male walkeri was collected at the higher elevations, and 52 male nudatus from the lower. The females were not considered since they are in- distinguishable. No evidence of morpholo- gical intergradation was fovmd. It appears that A. walkeri is better adapt- ed to areas having some persistent winter June 1977 Barnes et al.: Anomiopsyllus Fleas 195 snow and higher rainfall. Anomiopsyllus nu- datus is better adapted to lower elevations characterized by no winter snow, much less rainfall, and the occurrence of creosote bush. In neither case is the rainfall great. It must be pointed out that the rainfall in the habitat of ivalkeri is roughly twice that for the valley below, where nudattis occurs. Competitive exclusion in the zone of con- tact may occur between Anomiopsyllus nii- dcihis and A. ivalkeri in the larval stage, when both species occupy the same nest. Further evidence of this competitive exclu- sion between A. nudatus and A. walked is noted when analyzing the collection data outside their zone of contact. A. nudatus has been collected elsewhere in California from wood rat hosts and nests far above the winter snow line at elevations up to 2287 m, and in areas with average annual precip- itation up to 89 cm or more (e.g.. Big Bear, California, with a 94 cm annual average). Collections of nudatus in the Sawtooth and Liebre Mountains near Sandburg, south of Antelope Valley, and near Desert Springs, San Bernardino County, California, were made in habitats characteristically occupied by walked in the zone of contact. Anomio- psyllus nudatus is never found within the range of ivalkeri. In view of the morphological differences and the lack of intergradation at a point of close contact, A. ivalkeri must be considered a full species related to nudatus. These con- center to center distance between _spiniforms Fig. 93. Diagram of clasper showing measurements used on features of the movable process in the nudatus group. All measurements made by means of calibrated eyepiece grid. 196 Great Basin Naturalist Vol. 37, No. 2 siderations further suggest that walkeri has evolved in isolation from nudatus and has subsequently been brought into contact with the parent species. Anomiopsyllus martini Holland Figs. 16, 39, 61, 73, 88, 95 Anomiopsyllus martini Holland, 1965, Canadian Ent. 97: 1054-1056; Tipton and Mendez, 1968, Pacific In- sects 10(1): 179; Lewis, 1974, J. Med. Ent. 11(2): 154. Anomiopsyllus princei Barnes, 1965, Pan-Pacific Ent. 41(4): 276-278 (new synonomy); Haas at al., 1973, J. Med. Ent. 10(3): 282. Diagnosis: AnomiopsyUus martini has been considered a close relative of A. nu- datus and A. ivalkeri (Holland 1965, Barnes 1965). The males can be separated by the presence of four spiniforms on the movable process rather than three as in nudatus and from walkeri by two long, closely placed thick bristles at the arc of curvature of ster- num IX. AnomiopsyUus martini also differs from walkeri in that it has only three heavy setae on the apex of sternum IX, whereas walkeri has a fourth smaller seta. Female characters are not sufficiently dis- tinctive to separate martini from nudatus and walkeri. Type locality: 16 km W of El Salto, Du- rango, Mexico, 2744 m. Type host: Nest of Sciurus aberti Wood- house. Type specimens: Male holotype and fe- male allotype, 55 male and 59 female para- types on slides, and 187 male and 207 fe- male paratypes in alcohol collected on 21 June 1964 in a hollow in a dead pine tree; 3 male and 2 female paratypes, same data, but in another tree, on 11 July 1964 by J. E. H. Martin and deposited in the Canadian National Museum. Discussion: The recorded geographical distribution of this species is limited to two collections: one from El Salto, Durango, Mexico, at an elevation of 2744 m, and one from Catron County near Luna, New Mexi- co, at an elevation of 2195 m. Both collec- tions were from Sciurus aberti and its nest. Anomiopsyllus martini and A. durangoensis are the only species of the genus, of which we are aware, that are not associated with wood rats. We list A. princei Barnes as a synonym of A. martini Holland, even though there are some slight differences in the two popu- lations. In both species there are three dis- tal and one proximal spiniforms on the movable process of the clasper, and the dis- tal arm of sternum IX bears three subapical thick bristles with the posterior margin broadly rounded in the middle. There are several isolated populations of S. aberti squirrels associated with stands of yellow pine in Arizona, New Mexico, and Utah. Further collecting is required to de- termine if the distribution of A. martini is consistent with that of its apparent host, S. aberti. AnomiopsyUus durangoensis Holland Figs. 26, 47, 63, 75, 89, 95 Anoiniopsyllus durangoensis Holland, 1965, Cana- dian Ent. 97: 1056-1058; Lewis, 1974, J. Med. Ent. 11(2): 154. Diagnosis: Anomiopsyllus durangoensis is apparently sympatric with A. martini but associated with Peromyscus sp. Holland (1965) indicated that A. durangoensis is closely related to A. amphibolus (males) in that both have a triangular-shaped movable process with two spiniforms, but in duran- goensis both spiniforms are distal. On this basis durangoensis would appear to be more closely related to A. novomexicanensis, A. traubi, and A. oaxacae, which possess only distal spiniforms, whereas A. amphibolus also possesses one proximal and one distal spiniform. The distal arm of sternum IX is truncate to subtruncate and bears three short, heavy apical setae; the remaining ventral two-thirds bears fewer and thinner setae than in martini and A. n. hiemalis. Female characters are not sufficiently dis- tinctive to separate durangoensis from inar- tini or n. hiemalis. Type locality: 5 km W of Durango, Du- rango, Mexico, 1982 m. Type host: Mouse nest (possibly Per- omyscus sp.) Type speciinens: Male holotype and fe- male allotype, 11 male and 12 female para- types collected on 24 June 1964 in a nest in a yucca plant by J. E. H. Martin and depos- ited in Canadian National Museum. Discussion: The known geographical dis- June 1977 Barnes et al.: Anomiopsyllus Fleas 197 WESTERN NORTH AMERICA :^ DISTRIBUTION OF THE GENUS ANOMIOPSYLLUS ■ A.f. falsicalifornicus ° A.f.congruens • A.novomexicanensis o A. amphibolus * A. martini '^ A.nidiophilus « A. oaxacae ,3 Fig. 94. Map showing the distribution of Ano7niopsyUus species. 198 Great Basin Naturalist Vol. 37, No. 2 tribution is limited to the type locality near Durango, Durango, Mexico. Intensive col- lecting in Arizona and New Mexico may re- veal the presence of A. durangoensis. Anomiopsylhis novoniexicanensis Williams and Hoff Figs. 27, 49, 62, 76, 90, 95 Anoviiopsyllus novoniexicanensis Williams and Hoff, 1951, Proc. U.S. Nat. Mus. 101(3278): 307-309; Traub and Hoff, 1951, Amer. Mus. Nov. 1530: ,3-4; Vargas, 1960, Med. Rev. Mexicana 40(849): 4; Holland 1965, Canadian Ent. 97: 1053; Jellison and Glesne, 1967, In- dex to the literature of Siphonaptera of North Amer- ica, Suppl. 2: 13-19; Rail et al., 1969, J. Med. Ent. 6(1): 93; Miller et al., 1970, J. Med. Ent. 7(6): 698; Graves et al, 1974, J. Med. Ent. 11(4): 490-497. Diagnosis: Anomiopsylhis novomexica- nensis, though similar in many taxonomic characters to the nudatus group, is placed with A. traubi and A. oaxacae on the basis of having only distal spiniforms. Anomio- psylhis novoniexicanensis males can be dis- tinguished from other Anomiopsylhis species by the loss of the long bristle on the fixed process of the clasper. Females are readily identified from other Anomiopsylhis species by the presence of a prominent comma-shaped sclerotization in a portion of the bursa copulatrix. Type locality: 9.7 km E of Albuquerque, Bernalillo County, New Mexico. Type host: Neotoma micropus canescens Allen. Type specimens: Male holotype and fe- male allotype, .332 male and 605 female paratypes collected on 10 October 1948; 62 male and 126 female paratypes on 28 Feb- ruary 1948; 16 male and 4 female paratypes in December 1947 by L. A. Williams and C. C. Hoff, deposited in U.S. National Mu- seum. Material examined: Arizona: Cochise Co.: I male, 1 female, ex Neotoma lepida, 19 km W of Bowie on Tucson Rd., 914 m (cactus), II October 1938 (PHS); 1 female, ex Neo- toma lepida, 8 km SW of San Simon, 914 m (cactus), 13 October 19.38 (PHS); 1 male, ex Neotoma lepida, 9 km S of Bowie, 974 m (cactus), 15 October 1938 (PHS). Graham Co.: 3 males, 3 females, ex Neotoma lepida, 32 km NW of Duncan, 1128 m (grass), 28 Februarv 1939 (PHS); 1 male, ex Neotoma lepida, 32 km E of Safford, 975 m (cactus), 5 October 1938; 1 male, ex Neotoma lepida, 24 km S of Safford on Bowie Rd., 1113 m (cactus), 6 October 1938 (PHS); 1 female, ex Neotoma lepida, 38 km NW of Duncan, 1128 m (grass), 1 March 1939 (PHS); 2 fe- males, ex Neotoma lepida, 24 km E of Saf- ford, 1067 m (grass), 4 October 1938 (PHS); 1 female, ex Neotoma lepida, 40 km S of Safford on Bowie Rd., 1128 m (cactus), 8 October 1938 (PHS); 1 female, ex Neotoma lepida, 45 km NW of Duncan, 1128 m (grass), 2 March 1939 (PHS). Greenlee Co.: 3 males, ex Neotoma lepida, 6 km SE of Duncan, 1097 m (grass), 4 March 1939 (PHS); 1 male, ex Neotoma lepida, 13 km SE of Duncan at N. Mexico line, 1113 m (grass), 6 March 1939 (PHS); 1 male, 1 fe- male, host unknown (Onychomys leticogaster capitidatns?), no locality, 2 December 1938 (PHS). New Mexico: Bernalillo Co.: 1 fe- male, ex Neotoma rnicropus, 10 October 1948, L. Williams (RM); 3 males, 3 females, ex Neotoma alhigula, 24 km W of Albu- querque, (juniper, cactus— range), 16 May 19.39 (PHS); 4 males, 4 females, ex Neotoma sp., 10 October 1948, L. Williams (RM); 24 males, 1 female, ex Neotoma canescens, Al- buquerque, 10 October 1948, Williams and Hoff; 1 male, 1 female, ex Neotoma canes- cens, Albuquerque, 28 February 1948, Wil- liams and Hoff. Dona Ana Co.: 2 males, 2 females, ex Neotoma alhigula, 16 km W of Las Cnices, 1097 m (cactus, mesquite— desert), 15 April 1939 (PHS); 3 males, 3 fe- males, ex Neotoma alhigula, 11-16 km W of Hatch, 1280 m (yucca, grass), 12 May 1939 (PHS); 1 female, ex Neotoma alhigula, 8-11 km S of Hatch, 1280 m (chaparral), 10 May 1939 (PHS); 1 female, ex Neotoma alhigula, 3-6 km S of Salem, 1280 m (mesquite), 11 May 1939 (PHS); 1 female, ex Neotoma al- higula. Las Cruces, February 1939 (PHS); 1 female, ex Ontjcliomys leucogaster, 11-16 km W of Hatch, 1280 m (grass), 12 May 1939 (PHS); 1 female, ex Dipodomys ordii, 16 km W of Las Cruces, 1097 m (cactus- desert), 15 April 1939 (PHS). Grant Co.: 3 females, ex Neotoma alhigula, 19 km NE of Lordsburg, 1372 m (cactus, mesquite), 12 April 1939 (PHS). Hidalgo Co.: 1 male, 4 females, ex Neotoma alhigula, 19-22 km June 1977 Barnes et al.: Anomiopsyllus Fleas 199 ^ WESTERN NORTH AMERICA DISTRIBUTION OF THE GENUS ANOMIOPSYLLUS A. n. nudaius 'A. n. mexicanus A. n. hiemalis A. montanus 'A. sinuatus ^A. traubi A. durangoensis ' A. walkeri .:3 Fig. 95. Map showing the distribution of Anomiopsyllus species. 200 Great Basin Naturalist Vol. 37, No. 2 NW of Lordsburg, 1292 m (cactus), 11 April 1939 (PHS); 2 females, ex Neotoma al- bigula, 4.8 km S of Lordsburg, 1292 m (brush), 14 April 1939 (PHS); 5 males, 5 fe- males, ex Neotoma sp., 23 km NW of Lordsburg, 1292 m (mesquite), 8 April 1939 (PHS); 1 male, 1 female, ex Neotoma lepida, 21 km N of Rodeo, 1341 m (cactus), 20 Oc- tober 1938 (PHS); 2 males, 2 females, ex Neotoma lepida, 32 km N of Rodeo, 1402 m (cactus), 21 October 1938 (PHS); 2 males, 2 females, ex Neotoma lepida nest, 8 km N of Rodeo, 1341 m (cactus), October 1938 (PHS); 1 female, ex Peromysciis manicu- latiis, 32 km N of Rodeo, 1402, (grass, cac- tus), 21 October 1938 (PHS). Lincoln Co.: 1 male, 2 females, ex Neotoma albigula, 6 km NE of Carrizozo, 1646 m (yucca), 15 April 1949 (PHS). Luna Co.: 2 males, 5 females, ex Neotoma albigula, 5 km E of Deming, 1372 m (chaparral), 2 May 1939 (PHS); 2 males, 2 females, ex Neotoma albigula, 5-8 km S of Deming, 1372 m (chaparral), 5 May 1939 (PHS); 1 female, ex Dipodomys ordii, 5-8 km N of Deming, 1372 m (yucca, mesquite), 6 May 1939 (PHS). Otero Co.: 1 male, 1 female, ex Neotoma albigula, 10 km E of La Luz, Lincoln Nat. Forest, 1372 m, 22 March 1949 (PHS); 1 male, 1 female, ex Neotoma albigula, 11 km NE of La Luz, Lincoln Nat. Forest, 1463 m, 24 April 1949 (PHS); 1 male, 1 female, ex Neotoma albi- gula, 4 km E of Tularosa, 1524 m, 25 March 1949 (PHS). Rio Arriba Co.: 1 fe- male, ex Neotoma albigula, 3 km SW of Es- panola, 1982 m (grass, sage, cactus), 30 May 1939 (PHS). Sandoval Co.: 2 males, 3 fe- males, ex Neotoma mexicana, 3 km E of Bernalillo, 1524 m (juniper, sagebrush), 23 July 1949 (PHS); 1 male, 4 females, ex Neo- toma albigula, 8 km NW of Bernalillo, 1585 m (sage, juniper), 18 May 1939 (PHS). San Juan Co.: 1 male, 1 female, ex Neotoma al- bigula, 8 km SE of Aztec, 2134 m (juniper), 28 May 1949 (PHS). Sierra Co.: 1 male, 1 female, ex Neotoma albigula, 32 km N of Engle, 1280 m, 24 March 1949 (PHS); 2 males, 2 females, ex Neototyia albigula, 13 km E of Hot Springs, 1280 m, 25 March 1949 (PHS); 1 male, 3 females, ex Neotoma albigula, 19 km S of Hot Springs, 1220 m (cactus, sage-desert), 19 April 1939 (PHS); 1 female, ex Neotoma albigula, 8-10 km S of Hillsboro, 1555 m (sage, mesquite— plateau), 28 June 1939 (PHS). Socorro Co.: 4 males, 7 females, ex Neotoma albigula, 5 km N of Bernardo, 1433 m (grass, sage, mesquite-range), 12 May 1939 (PHS); 4 males, 4 females, ex Neotoma albigula, Abo- Pass, 10 km W of Schoole, 1738 m (grass, oaks, juniper), 11 May 1939 (PHS); 2 males, 2 females, ex Neotoma albigula, 3 km SW of San Antonio (range), 6 May 1939 (PHS); 1 male, 4 females, ex Neotoma albigtda, 3 km N of Escondida, 1433 m (near irrigated crops-gravel banks), 2 May 1939 (PHS); 1 male, 1 female, ex Neotoma albigula, 16 km W of Socorro, 1524 m (grass, sage— range), 4 May 1939 (PHS); 1 male, 5 females, ex Neotoma albigula, 16 km E of San Antonio, 1372 m (sage, cactus— range), 5 May 1939 (PHS); 2 females, ex Neotoma albigula, 7 km N of Polradera, 1463 m (sage, mes- quite), 3 May 1939 (PHS); 1 female, ex Neo- toma albigula, 12 May 1939 (PHS); 1 fe- male, ex Neotoma albigula, 3 km W of Socorro, 1402 m (cactus, mesquite-range) 2 May 1939 (PHS). Valencia Co.: 4 males, 5 females, ex Dipodomys merriami, 3 km W of Belen, 1463 m (grass, sage— dump grounds), 9 May 1939 (PHS); 1 female, ex Neotoma albigtda, same locality, 1463 m (grass, sage— dump grounds), 9 May 1939 (PHS); 2 females, ex Neotoma albigula, 13 km W of Belen, 1524 m (juniper, sage- range), 9 May 1939 (PHS); 1 female, ex Neotoma albigula, 31 km W of Hoshunas, 1494 m (sage-range), 10 May 1939 (PHS); 1 female, ex Neotoma albigula, 24 km S of Correo, 1524 m (sage, juniper), 25 April 1939 (PHS); 1 male, ex Peromyscus truei, 12.9 km W of Belen, 1524 m (juniper- range), 9 May 1939 (PHS); 1 male, ex Per- omyscus boylei, Harrington Ranch, 1524 m (sage, juniper), 26 April 1939 (PHS). Texas: Hudspeth Co.: 1 female, ex Neotoma, 19 September 1940 (RM); 3 males, 1 female, ex Neotoma nest, Allamore, 19 September 1940 (RM). Culberson Co.: 2 males, 2 females, ex Neotoma albigula, 31 March 1942 (PHS). Discussion: In their remarks following the description of A. novomexicanensis Williams and Hoff (1951) grouped this species with A. n. hiemalis on the basis of its having June 1977 Barnes et al.: Anomiopsyllus Fleas 201 three spiniforms on the movable process. They considered the genus to contain two species groups: those with two such spin- iforms (i.e., A. amphibolus) and those with three (i.e., niidatiis hiemalis). Anomiopsyllus falsicalifornicus with four (usually) was not mentioned. Characteristics of the fixed process in- cluded in the diagnosis are not found in Anomiopsyllus species and would apparent- ly place A. novomexicanensis in a mon- otypic species group. The shape, articu- lation, and adornment of the movable process in novomexicanensis would seem to indicate its closeness to A. traubi and prob- ably place it in the group which includes A. traubi and A. oaxacae. Hosts: Anomiopsyllus novomexicanensis has been collected from four wood rat hosts: Neotoma albigula, N. micropus (type host), N. mexicana, and N. lepida stephensi. Occasional hosts include Peromyscus, Dipo- domys, and a few sciurids with about the same infestation rate as for other Anomiop- syllus species. There were 52 wood rat col- lections in the material examined, of which N. albigula predominated (33), followed by N. lepida stephensi (15), N. micropus (4), and N. mexicana (1). The known range of A. novomexicanensis falls within the distri- bution of N. albigula and iV. mexicana, but occupies only a small portion of either. The collection records for N. lepida are all from eastern Arizona. The distribution of iV. rni- cropus does not extend west beyond New Mexico (Hall and Kelson 1959). As is un- doubtedly the case with other Anomio- psyllus species for which considerable data exist, the distribution of A. novomexica- nensis is limited by factors other than that of host distribution. Distribution and habitat (Fig. 95): The known distribution of A. novomexicanensis includes most of southern and central New Mexico, a small portion of southeastern Ari- zona (Greenlee and Graham counties) and the westernmost corner of Texas (Culberson and Hudspeth counties). To the north and west its distribution interdigitates with that of A. nudatus mexicanus and to the east (Texas) with A. nudatus hiemalis. Collection records, where habitat clues and elevations are given, show that A. no- vomexicanensis is a flea of the Chihuahuan amphibolus ID-IP _ montanus sinuatus 1 2D-1P 2D-1P _ n. nudatus - n. mexicanus - n. hiemalis _ f. falsicalifor - f- congruens D and P 3D 3D-1P spiniforms 0-row licus 3D-1P V-row 3D-1P 3D-1P 2D - durangoensis - novomexicanensi only d spiniforms >. - traubi - oaxacae - nidiophilus 1D-3P spiniforms D-Dista P-Proxi 1 mal 0-obli V-vert que ical Diagram 1. Cladogram illustrating the relationships of species and subspecies in the genus Anomiopsyllus, based on the spiniforms of the movable process on the clasper. 202 Great Basin Naturalist Vol. 37, No. 2 biotic province. It occupies the desert plains and the arid grassland-mesquite belt mentioned by Bradt (1932) and Benson (1933). It also inhabits the low wastelands where its hosts occur, but it primarily oc- cupies the Chihuahuan roughlands belt of Blair (1940) and the lowest edges of the pinyon-juniper belt (sabinal belt of Dice 1942) at elevations of 1220 to 1524 m. Ex- treme southern New Mexico in Luna, Dona Ana, and Otero counties, where novome- xicanensis is the only AnomiopsijUus record- ed, also falls within the Chihuahuan pro- vince (Dice 1943). Elements of the Chihuahuan desert interdigitated with por- tions of the Colorado plateau to the north (Dice 1943, Fenneman 1931). Here, A. no- vomexicanensis is found at lower elevations and A. n. mexicaniis at higher elevations. Of 52 collections of novomexicanensis, 94 percent were made below 1577 m (x = 1357 m), but 83 percent of the A. n. mexicaniis collections were made above 1524 m and 72 percent above 1692 m (x = 1829 m). Further evidence that novomexicanensis occurs primarily in the lower desert habitats is offered by clues to vegetation type in Public Health Service collection records. These records, where available, have been included in the collection data under mate- rial examined. References to cactus, yucca, and mesquite are much more common to the Chihuahuan province where novomex- icanensis occurs, and references to piny on, juniper, and sagebrush are much more com- mon to the Colorado Plateau where A. n. mexicanus occurs. Differences in habitat occupied by the two species are probably dependent on two factors: (1) climate, particularly humidity or available moisture as in the observed habi- tat division between A. falsicalifornicus and A. nudatus nudatus or between n. nitdatus and walkeri, and (2) competitive exclusion resulting from two related species utilizing the same host niche. Altitudinal overlap in the records shown here may be explained on the basis of differences in microclimate brought about by slope exposure, rain shad- ow, and precipitation pattern. AnomiopsijUus traubi Barrera Figs. 28", 48\ 9r, 94 Anomiapsyllus traubi Barrera, 1951, Ciencia, 12(6- 9): 197; Vargas, 1960, Med. Rev. Mexicana, 40(849): 1- 4; Hopkins and Rothschild, 1962, Catalogue of the Rothschild Collection of Fleas III: 401-402; Holland, 1965, Canadian Ent. 97: 1054; Lewis, 1974, J. Med. Ent. 11(2): 154. Diagnosis: A. traubi males can be distin- guished from A. novomexicanensis and A. oaxacae in that the distal arm of sternum IX has 5-7 stout apical setae. The posterior margin of sternum IX is bare except for a subbasal row of 5-7 long, stout bristles in- termixed with long, slender bristles and a single submedian seta. The movable process is broadly triangular and rounded at the posteroventral angle containing a row of three spiniforms placed vertically along the posterior margin. Type locality: Cueve de Belen, near Zimapan, Hidalgo, Mexico. Type host: Nest of rodent. Type specimen: Male holotype collected on 2 November 1950 by A. Barrera, depos- ited in Alfredo Barrera's collection. Discussion: Barnes (1963) examined only two male paratypes of this species: one pro- vided by William L. Jellison and one pro- vided by Robert Traub. These specimens had been mounted in Hoyer's mounting me- dium, and the exoskeletons were fractured by shrinkage of the medium, resulting in the destruction of a number of characters of taxonomic value, particularly in the ae- deagus. Hopkins and Rothschild (1962) be- lieve A. traubi to be most closely related to A. novomexicanensis, but the fixed process of A. novomexicanensis is unique in the genus whereas that of traubi is most like A. oaxacae. The distal arm of sternum IX in traubi and oaxacae differ by virtue of the subbasal angle of the posterior margin of this structure. The true relationships of traubi and oaxacae will remain imcertain until additional specimens are available. "Figures 28, 48, and 91 were redra of the lack of a specimen. Barrera (1951) and Barnes (1963). The aedeagiis and Vlll of the male were not illustrated because June 1977 Barnes et al.: Anomiopsyllus Fleas 203 Anomiopsijllus oaxacae Barnes Figs. 29, 50, 64, 77, 92, 95 Anomiopsyllus oaxacae Barnes, 1965, Pan-Pacific Ent. 41(4): 278-279; Lewis, 1974, J. Med. Ent. 11(2): 154. Diagnosis: Barnes (1965) indicated that A. oaxacae is most closely related to A. /. falsi- californicus but very different from that species and all other species of Anornio- psyllus in that it possesses more and gener- ally longer bristles. Anomiopsyllus oaxacae is probably closer to A. trauhi than to /. fahicaliforniciis, based on the presence of only distal spiniforms on the movable proc- ess in A. trauhi, whereas /. congniens still retains a proximal spiniform. The movable process is broad almost to the point of being rectangular in shape and is equipped with three mesad spiniform setae parallel to the posterior margin of the process. The proximal arm of sternum IX is similar to that of /. falsicalifornicus, but the distal arm is long, narrow, and almost parallel sided and armed apically with two posterad rows of heavy, spikelike setae. The posterior margin of the distal arm has a row of 20 evenly spaced, long, thin bristles extending from the base of the lowest apical seta to three-fourths of the distance of the arm. The aedeagal hood extends anterad far beyond the aedeagal struts, and the length of the hood is about equal to the length of the apodeme beyond the anterior margin of the hood. Length: 1.7 mm. Type locality: 3 km SE of Matatlan, Oax- aca, Mexico Type host: Baiomys musculus (Merriam) Type specimen: Male holotype collected on 24 July 1953 by R. H. Baker, deposited in U.S. National Museum. Discussion: This species was described from a single male specimen that was loaned to the senior author by Traub and was originally noted as new by Traub. This specimen represents the southernmost re- cord for the genus. Thus, the range of Ano- miopsyllus extends into tropical America from whence it reaches northward to Banff, Alberta, Canada. Although Baiomys mus- culus is the type host, the normal host is probably Neotoma mexicana, the distribu- tion of which includes Oaxaca, Mexico. Literature Cited Allred, D. M. 1968. Fleas of the national reactor test- ing station. Great Basin Nat. 28(2): 78-87. Anonymous. 1950a. Plague infection in Lea County, New Mexico. Public Hlth. Repts. 65: 454. 1950b. Plague in Lea County, New Mexico. Public Hlth. Repts. 65: 526. AucusTsoN, G. F. 1943. Preliminary records and dis- cussion of some species of Siphonaptera from the Pacific Southwest. Bull. South. Calif. Acad. Sci. 42(2): 69-96. 1955. Records of fleas from the Pacific South- west. Bull. South. Calif. Acad. Sci. 54(1): ,36-.39. AuGUSTsoN, G. F., AND F. E. Durham. 1961. Records of fleas (Siphonaptera) from northwestern Ari- zona. Bull. South. Calif. Acad. Sci. 60(2): 100. AuGusTsoN, G. F., AND S. F. WooD. 1953. Notes on California mammal ectoparasites from the Sierra Nevada Foothills of Madera County. Bull. South. Calif. Acad. Sci. 52(2): 48-56. Baker, C. F. 1898. Notes on Siphonaptera, with de- scriptions of four new species. J. N.Y. Ent. Soc. 6: 53-56. 1904. A revision of American Siphonaptera, or fleas, together with a complete list of the group. Proc. U.S. Nation. Mils. 27(1361): .365-469. Barnes, A. M. 196.3. A revision of the genus Anomio- psyllus Baker 1904 (Siphonaptera: Hys- trichopsyllidae) with studies on the biology of Anomiopsyllus falsicalifornicus. Ph.D. Dis- sertation, University of California, Berkeley, Cali- fornia. 1965. Three new species of the genus Ano- miopsyllus (Siphonaptera: Hystrichopsyllidae). Pan-Pacific Ent. 41(4): 272-280. Barrera, a. 1951. Notas sobre Sifonapteros. II. De- scripcion de Anomiopsyllus trauhi nov. sp. (Siph., Hystrichops.). Ciencia. 12(6-9): 197-200. Beck, D. E., A. H. Barnum, and L. Moore. 1953. Ar- thropod consorts found in nests of Neotoma cine- rea acraia (Ord) and Neotoma lepida lepida Thomas. Proc. Utah Acad. Sci., Arts, and Lett. 30: 43-52. Beck, D. E., and D. M. Allred. 1966. Siphonaptera (fleas) of the Nevada Test Site. Brigham Young Univ. Sci. Bull., Biol. Ser. 7(2): 1-27. Beer, J. R., E. F. Cook, and R. G. Schwab. 1959. The ectoparasites of some mammals from the Chiri- cahua Moimtains, Arizona. J. Parasitol. 45(6): 605- 623. Benson, S. B. 1933. Concealing coloration among some desert rodents of the southwestern United States. Univ. Calif. Publ. Zool. 40: 1-70. Blair, W. F. 1940. A contribution to the ecology and faunal relationships of the mammals of the Davis Mountain Region, southwestern Texas. Misc. Publ., Mus. Zool., Univ. Mich. 46: .37. 204 Great Basin Naturalist Vol. 37, No. 2 Bradt, G. W. 1932. The mammals of the Malpais, an area of black lava rock in the Tularosa Basin, New Mexico. J. Mammal. 13: 321-328. Brand, D. D. 1936. Notes to accompany a vegetation map of northwest Mexico. Univ. N. M. Bull., Biol. Ser. 4(4): 3-27. Collins, B. J. 1936. A new species of Anomiopsyllus from Montana (Siphonaptera). Ent. News. 47: 128- 130. Costa Lima, A. da, and C. R. Hathaway. 1946. Pul- gas, Bibliografia, catalogo e animais por Elas Su- gados. Monogr. Inst. Oswaldo Cruz. 4: 1-522. CouLTRip, R. L., R. W. Emmons, L. H. Legter, J. D. Marshall, Jr. and K. F. Murray. 1973. Survey for the arthropod vectors and mammalian hosts of Rocky Mountain Spotted Fever and Plague at Fort brd, California. J. Med. Ent. 10(3): 303-309. Dalla Torre, C. G. 1924. Aphaniptera Orbis Terra- rum (Synopis Praecursoria). Naturw. Med. Ver. Innsbnick. ,39: 1-29. Dice, L. E. 1942. Ecological distribution of Per- omyscus and Neotoma in parts of southern New Mexico. Ecology 23(2): 199-208. 1943. The biotic provinces of North America. Univ. Mich. Press, Ann Arbor. 78 pp. Dunn, L. H. and R. R. Parker. 1923. Fleas found on wild animals in the Bitterroot Valley, Montana. Public Hlth. Repts. 38: 2763-2775. Eads, R. B., and G. C. Menzies. 1948. An undescribed Anomiopsyllus Baker from the pack rat, Neotoma micropus Baird. J. Kans. Ent. Soc. 21(4): 133-136. 1950. The fleas of Texas. Texas State Hlth. Dept. Bull. 85 pp. Ellis, L. L. 1955. A study of the ectoparasites of cer- tain mammals in Oklahoma. Ecology. 36(1): 12-18. EsKEY, C. R., AND V. H. Haas. 19.39. Plague in the western part of the United States. Infection in ro- dents, experimental transmission by fleas, and in- oculation tests for infection. Public Hlth. Repts. 54(32): 1467-1481. 1940. Plague in the western part of the United States. Public Hlth. Bull. 254: 1-83. EwiNG, H. E. 1929. Notes on the Siphonaptera genus Catallagia Rothschild, including the description of a new species. Proc. Biol. Soc. Wash. 42: 125-128. EwiNG, H. E., AND I. Fox. 1943. The fleas of North America. Classification, identification, and geo- graphic distribution of these injurious and dis- ease-spreading pests. U.S. Dept. Agric. Misc. Publ. 500: 128 pp. Fenneman, N. M. 1931. Physiography of the western U.S. McGraw-Hill, New York. 534 pp. Finley, R. B., Jr. 1958. The woodrats of Colorado: Distribution and ecology. Univ. Kans. Publ. Mus. Natur. Hist. 10(6): 21.3-552. Fox, C. 1914. I. Some new Siphonaptera. II. A further report on the identification of some Siphonaptera from the Philippine Islands. III. The taxonomic value of the copulatory organs of the females in the order Siphonaptera. Hygenic Laboratory Bull. 97: 1-25. 1926. New Siphonaptera from California. Pan- Pacific Ent. 2(4): 183. 1929. Some new Siphonaptera. Ent. News. 40(7): 218. Goldman, E. A. 1910. Revision of the woodrats of the genus Neotoma. U.S. Dept. Agric. Biol. Survey, N. Amer. Fauna. 31: 1-124. Graves, N. G., W. C. Bennett, J. R. Wheeler, B. E. Miller, and D. L. Forcum. 1974. Sylvatic plague studies in southeast New Mexico. I. Flea- host relationships from six years' study. J. Med. Ent. 11(4): 488-498. Haas, G. E., R. P. Martin, M. Swickard, and B. E. Miller. 1973. Siphonaptera-mammal relationships in northcentral New Mexico. J. Med. Ent. 10(3): 281-289. Hall, E. R. 1946. Mammals of Nevada. Univ. Calif. Press, Berkeley. 710 p. Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. Vol. II. The Ronald Press, New York. pp. 681-1083. Hirst, L. F. 1926. Researches on the parasitology of plague. Part I. Ceylon J. Sci., Sect. D. Med. Sc. 1: 155-276. Holdenreid, R. and H. B. Morlan. 1955. Plague- infected fleas from northern New Mexico wild ro- dents. J. Inf. Dis. 96: 1.3.3-137. Holland, G. P. 1949. Siphonaptera of Canada. Do- minion of Canada-Dept. Agric. Publ. 817, Tech. Bull. 70, 306 pp. 1952. Notes on some Siphonaptera from Can- ada. Canadian Ent. 84(3): 65-73. 1965. New species and subspecies of Anomio- psyllus Baker from Mexico (Siphonaptera: Hys- trichopsyllidae). Canadian Ent. 97: 1051-1058. Hooper, E. T. 1938. Geographic variation in wood rats of the species Neotoma fuscipes. Univ. Calif. Publ. Zool. 42: 213-246. Hopkins, G. H. E. 1952. Notes on synonomy in Siphonaptera. J. Wash. Acad. Sci. 42(11): 36.3-365. Hopkins, G. H. E., and M. Rothschild. 1953. An il- lustrated catalogue of the Rothschild Collection of fleas (Siphonaptera) in the British Museum (Natural History). Vol. I. Tungidae and Pulicidae. Cambridge Univ. Press, 361 pp. 1956. An illustrated catalogue of the Roths- child Collection of fleas (Siphonaptera) in the British Museum (Natural History). Vol. II. Cop- topsyllidae, Vermipsyllidae, Stephanocircidae, Ischnopsyllidae, Hypsophthalmidae and Xiphio- psyllidae. Cambridge Univ. Press, 445 pp. 1962. An illustrated catalogue of the Roths- child Collection of fleas (Siphonaptera) in the British Museum (Natural History). Vol. III. Hys- trichopsyllidae. Cambridge Univ. Press, 560 pp. Howell, J. F. 1955. A study of the aspectional varia- tions of Siphonaptera associated with nests of the Thomas wood rat, Neotoma lepida lepida Thomas. Great Basin Nat. 15: .35-49. 1957. Fleas associated with nests of the desert wood rat. J. Parasitol. 43(5): ,566-573. Hubbard, C. A. 1940. A checklist of the fleas of the Pacific Northwest (Washington, Oregon, northern California and northwest Nevada) with notes from southern California. Pacific Univ. Bull. 1977 Barnes et al.: Anomiopsyllus Fleas 205 37(4): 1-4. 1943. The fleas of California (with checkhsts of the fleas of Oregon, Washington, British Co- lumbia, Alaska, Idaho, Nevada, and Arizona). Pa- cific Univ. Bull. 39(8): 1-12. 1947. Siphonaptera of western North America. Iowa State College Press, 533 pp. 1949. Fleas of the state of Nevada. Bull. South. Calif. Acad. Sci. 48(3): 115-128. Jellison, W. L., and N. E. Good. 1942. Index to the literature of Siphonaptera of North America. U.S. Public Hlth. Serv., Nat. Inst. Hlth. Bull. 178:1- 193. Jellison, W. L., G. M. Kohls, and H. B. Mills. 1943. Species and host list of Montana fleas. Montana State Board of Ent. Misc. Publ. 2: 22 pp. Jellison, W. L., B. Locker, and R. F. Bacon. 1953. Index to the literature of Siphonaptera of North America. Supplement 1, 1939-1950. U.S. Public Hlth. Serv., Nation. Inst. Hlth., Nation. Microbio. Inst., Rocky Mountain Lab., Hamilton, Montana. 246 pp. Jellison, W. L., and L. Glesne. 1967. Index to the literature of Siphonaptera of North America. Sup- plement 2, 1951-1960. U.S. Public Hlth. Serv., Na- tion. Inst. Hlth., Nation. Inst. Allergy and Infect. Dis., Rocky Mountain Lab., Hamilton, Montana. 406 pp. Jellison, W. L., and C. M. Senger. 1973. Fleas of Montana. Montana Agric. Exp. Sta., Montana State Univ., Bozeman, Montana, Res. Rept. 29: 75 pp. Johnson, P. T. 1957. A classification of the fleas of South America. Mem. Ent. Soc. Wash. 5: 299 pp. Jordan, K., and N. C. Rothschild. 1915. Contribution to our knowledge of American Siphonaptera. Ec- toparasites 1(1): 45-60. Kartman, L., F. Prince, S. Quan, and H. Stark, 1960. The role of rabbits in sylvatic plague epide- miology with special attention to human cases in New Mexico and use of the fluorescent antibody technique for detection of Pasteurella pestis in field specimens. Zoon. Res. 1(1): 1-27. Knudsen, J. W. 1956. Pseudoscorpions, a natural con- trol of Siphonaptera in Neotoma nests. Bull. South. Calif. Acad. Sci. 55(1): 1-6. Leopold, A. S. 1950. Vegetation zones of Mexico. Ecology 31(4): 507-518. Lewis, R. E. 1974. Notes on the geographical distribu- tion and host preferences in the order Siphonap- tera. Part 3. Hystrichopsyllidae. J. Med. Ent. 11(2): 147-167. LiNSDALE, J. M., AND B. S. Davis. 1956. Taxonomic ap- praisal and occurrence of fleas at the Hastings Reservation in central California. Univ. Calif. Publ. Zool. 54(5): 293-370. Mendez, E. 1956. A revision of the genus Megarthro- glossus Jordan and Rothschild 1915 (Siphonaptera: Hystrichopsyllidae). Univ. Calif. Publ. Ent. 11(3): 159-192. Miller, B. E., D. L. Forcum, K. W. Weeks, J. R. Wheeler, and C. D. Rail. 1970. An evaluation of insecticides for flea control on wild mammals. J. Med. Ent. 7(6): 697-702. Oudemans, a. C. 1909. Neue Ansichten iiber die Mor- phologic des Flohkoptes sowie viber die Onto- genie, Phylogenie und Systematic der Flohe. Nov. Zool. 16:133-178. Parker, D. D., and J. F. Howell. 1959. Host-flea rela- tionships in the Great Salt Lake Desert. J. Para- sitol. 45(6): 507-604. PoLLiTizER, R. 1952. List of the wild rodent fleas which have been found plague infected in nature or proved to be susceptible to experimental in- fection. Bull. WHO 7: 318: 322, Annex 1. PooRBAUGH, J. H., and H. T. Gier. 1961. Fleas (Siphonapteral of small mammals in Kansas. J. Kans. Ent. Soc. .34(4): 198-204. Pratt, H. D., and H. E. Stark. 1973. Fleas of public health importance and their control. DHEW, Publ. No. (CDC) 74-8267: 42 p. Rail, E. D., D. L. Forcum, J. R. Wheeler, and E. B. Miller. 1969. Wild mammals and fleas of Red Bluff Ranch, New Mexico. J. Med. Ent. 6(1): 92- 94. Rainey, D. G. 1956. Eastern woodrat, Neotoma florid- ana: Life history and ecology. Univ. Kans. Publ., Mus. Nat. Hist. 8(10): 535-646. Senger, C. M. 1966. Notes of fleas (Siphonaptera) from Montana. J. Kans. Ent. Soc. 39(1): 105-109. Shreve, F., and I. Wiggins. 1951. Vegetation of Sono- ran Desert. Carnegie Inst, of Wash. Publ. 591, Washington, D.C. Vol. 1, 192 pp. Snodgrass, R. 1946. The skeletal anatomy of fleas (Siphonaptera). Smithson. Misc. Coll. 104: 1-89. Stanford, J. S. 1931. A preliminary study of Utah Siphonaptera. Proc. Utah Acad. Sci. 8: 153. 1943. More Utah Siphonaptera. Proc. Utah Acad. Sci., Arts, and Lett. 19-20: 17.3-178. Stark, H. E. 1959. The Siphonaptera of Utah. U.S. Dept. H.E.W., Public Hlth. Serv. 239 pp. Stark, H. E., and A. R. Kinney. 1969. Abundance of rodents and fleas as related to plague in Lava Beds National Monument, California. J. Med. Ent. 6(3): 287-294. Stewart, M. A. 1940. New Siphonaptera from Califor- nia. Pan-Pacific Ent. 16: 17-18. Tipton, V. J. 1950. New distributional records for Utah Siphonaptera. Great Basin Nat. 10: 1-4. Tipton, V. J., and R. C. Saunders. 1971. A list of ar- thropods of medical importance which occur in Utah with a review of arthropod-borne diseases endemic in the state. Brigham Young Univ. Sci. Bull., Biol. Ser. 15(2): 1-31. Tipton, V. J., and E. Mendez. 1968. New species of fleas (Siphonaptera) from Cerro Potosi, Mexico, with notes on ecology and host-parasite relation- ships. Pacific Insects 10(1): 177-214. Traub, R. 1950. Siphonaptera from Central America and Mexico. A morphological study of the ae- deagus with descriptions of new genera and spe- cies. Fieldiana: Zool. Mem. 1: 1-127 (Chicago Na- tur. Hist. Mus.) 1953. Systematic notes on eyeless fleas. J. Wash. Acad. Sci. 43: .352: .353. Traub, R., and C. C. Hoff. 1951. Records and de- 206 Great Basin Naturalist Vol. 37, No. 2 scriptions of fleas froin New Mexico (Siphonap- tera). Amer. Mus. Nov. 1530: 23 pp. Traub, R., and V. J. Tipton. 1951. Jordanopsylla all- redi, a new genus and species of fleas from Utah. J. Wash. Acad. Sci. 41(8): 264-270. Vargas, L. 1960. Encuentro de Anoniiopsyllus hiemalis Eads y Menzies, 1948. Med. Rev. Mexicana. 40(849): 1-4. Wagner, J. 1930. Keys to the Aphanptera Hving on Muridae. Mag. Parasitol. 1: 97-192. 1936. Neve norda amerikanische Floharten. Zeits fur Parasitol. en Kunde, 8(6): 654-655. Warren, E. R. 1942. The mammals of Colorado. Univ. Oklahoma Press, Norman, Oklahoma. 330 pp. Wiseman, J. S. 1955. The Siphonaptera (fleas) of Wyoming. Univ. Wyo. Publ. 19(1&2): 28 pp. Webb, W. L. 1951. Biogeographic regions of Texas and Oklahoma, Ecology 31(3): 426-433. Wehrle, L. p. 1953. A host index of Arizona fleas. Pan-Pacific Ent. 24(1): 37-41. Williams, L. A., and C. C. Hoff. 1951. Fleas from the upper Sonoran Zone near Albuquerque, New Mexico. Proc. U.S. Nation. Mus. 101(3278): 305- 313. NEW SYNONYMY AND NEW SPECIES OF AMERICAN BARK BEETLES (COLEOPTERA: SCOLYTIDAE), PART IV' Stephen L. Wood" Abstract.— New synonymy is proposed as follows: Pityophthorus Eichhoff ( = Pityophthoroides Blackman), Scolytodes Ferrari { = Ctenophorus Chapuis), Araptus pallidus (Blackman), n. comb. (? = Neodryocoetes portori- censis Schedl, Neodryocoetes devius Schedl), Corthylus letzneri (Ferrari) ( = CorthyIus strigilatus Eggers, Corthylus columbianus Schedl, Corthylus ater Schedl), Corthylus spinifer Schwarz { = Corthylus guayanensis Eggers), Hy- pothenemus brunneus (Hopkins) { = Hypothenemus cryphalomorphus Schedl, Stephanoderes bituberculatus Eggers), Monarthrum parvum (Eggers) { = Monarthrum praeclarum Wood), Phloeoborus punctatorugosus Chapuis { = Phloeoborus breviusculus Chapuis), Phloeotrihus pilula Erichson { = Phloeotribus australis Schedl), Phloeotribus setulosus Eichhoff {Phloeotribus dubius Eichhoff, Phloeotribus spinipennis Eggers), Pityophthorus, pulicarius (Zim- mermann) { = Pityophthorus cubensis Schedl), Pityophthorus subsimilis Schedl { = Pityophthorus subimpressus Bright), Premnobius cavipennis Eichhoff { = Premnobius latior Eggers), Scolytus propiguus Blandford { = Scolytus penicillus Schedl), Xyleborus dryographus (Ratzeburg) { = Xyleborus linearis Schedl), Xyleborus horridus Eichhoff [ = Xyleborus flohri Schedl). Xyleborus tonsus (Hagedom), n. comb., was transferred to Xyleborus from Dryocoetes. Pityophthorus aquilus Blackman and Pityophthorus cascoensis Blackman, n. status, were removed from synonymy. Scolytodes chapuusi, n. n., is proposed for Scolytodes { = Ctenophorus) laevigatus Chapuis, 1869 (nee Ferrari 1867). The following species are named as new to science: Araptus crassus (Panama), A. obesus (Brazil), Cnesinus pro- minulus (Mexico), Conophthorus conicolens (Mexico), Microcorthylus vicinus (Mexico), Pityophthorus alnicolens (Mexico), P. assitus (Mexico), P. concinnus (Mexico), P. indigens (Colorado), P. leechi (California), P. solatus (Mex- ico), P. speciosus (Mexico), Pseudopityophthorus denticulus (Texas), Pseudothysanoes securus (Mexico), P. turnbowi (Texas), Pycnarthrum fulgidum (Colombia), Scolytodes fulmineus (Venezuela), S. radiatus (Costa Rica), Xylebo- rinus tribuloides (Mexico), Xyleborus devexus (Florida, Puerto Rico), and X. laticeps (Venezuela). On the following pages several newly dis- covered cases of synonymy and of species new to science are presented for American Scolytidae. The 21 species new to science represent the genera Cnesinus (1), Pycnarth- rum (1), Scolytodes (2), Pseudothysanoes (2), Xyleborus (2), Xyleborinus (1), Pseudopi- tyophthorus (1), Araptus (2), Microcorthylus (1), Conophthorus (1), and Pityophthorus (7), and they are from the following countries: United States (5, including one which also occurs in Puerto Rico), Mexico (10), Costa Rica (1), Panama (1), Columbia (1), Vene- zuela (2), and Brazil (1). New Synonymy Pityophthorus Eichhoff Pityophthorus Eichhoff, 1864, Berliner Ent. Zeitschr. 8:39 (Type-species: Bostrichus lichtensteini Ratze- burg, subsequent designation by Hopkins, 1914, Proc. U. S. Nat. Mus. 48:127). Pityophthoroides Blackman, 1942, Proc. U.S. Nat. Mus. 92:199 (Type-species: Pityophthoroides pudens Blackman, original designation). New synonymy In a review of the North and Central American Pityophthorus and allied genera, Blackman's Pityophthoroides pudens was placed in the genus Pityophthorus following a study of the type series. The characters on which Pityophthoroides was based inter- grade completely with those of Pityoph- thorus. Blackman's name, therefore, must be placed in synonymy. Scolytodes Ferrari Scolytodes Ferrari, 1867, Die Forst- und Baumzuchtschadlichen Borkenkafer, p. 77 (Type- species: Scolytodes laevigatus Ferrari, Monobasic) Ctenophorus Chapuis, 1869, Synopsis des Scolytides, p. 49 (Type-species: Ctenophorus laevigatus Chapuis, monobasic). New synonymy Each type series of Scolytodes laevigatus Ferrari and Ctenophorus laevigatus Chapuis consists of two males, all four of which were 'Part of this research was sponsored by the National Science Foundation. 'Life Science Museum and Department of Zoology, Brigham Young University, Prove, Utah 84602. Scolytidae contribution No. 64. 207 208 Great Basin Naturalist Vol. 37, No. 2 examined. It is clear that they are con- generic, but that they represent different species. Since this generic synonymy creates homonymy, Chapius's species is renamed below. Araptus pallidus (Blackman), n. comb. Neodryocoetes pallidus Blackman, 1942, Proc. U.S. Nat. Mus. 92(3147): 103 (Holotype, male; Cayamas, Cuba; U.S. Nat. Mus.) Neodryocoetes portoricensis Schedl, 1951, Dusenia 2:109 (Holotype, male; Portorico; Schedl Coll). Probable synonymy Neodryocoetes devius Schedl, 1972, Koleopt. Rund- schau 50:61 (Holotype, male; Valle Ancon, Pinar del Rib, Cuba; Schedl Coll.). Probable synonymy The male holotypes of Neodryocoetes pal- lidus Blackman, portoricensis Schedl, and devius Schedl were all examined at different times and compared to my series of closely allied Araptus tenellus (Schedl). All are 1.7 mm in length, of pale color, and fit the same descriptive notes. Although they have not been compared directly to authentic material, there is a strong possibility of syn- onymy. Corthylus letzneri (Ferrari) Pseudocorthylus letzneri Ferrari, 1867, Die Forst- und Baumzuchtschadlichen Borkenkafer, p. 59 (Syn- type, male; Venezuela, apparently from Colonia Tovar; Vienna Museum). Corthylus strigilatus Eggers, 1933, Trav. Lab. d'Ent. Mus. Nat. d'Hist. Nat., Paris, Mem. 1:20 (Holo- type, male; Colonia Tovar, Venezuela; Paris Mus.). New synonymy Corthylus columbianus Schedl, 1950 (nee. Hopkins, 1894), Dusenia 1:158 (Holotype, female; Comatan, Colombia; Schedl Coll.). New synonymy Corthylus ater Schedl, 1952, Dusenia 3:345 (Replace- ment name for columbianus Schedl). New synon- ymy A male syntype of Pseudocorthylus letz- neri Ferrari, the male holotype of Corthylus strigilatus Eggers, and the female holotype of Corthylus columbianus Schedl were all compared directly to my .series from Merida and Rancho Grande (Aragua), Venezuela. All clearly represent the same species. Corthylus spinifer Schwarz Corthylus spinifer Schwarz, 1891, Proc. Ent. Soc. Washington 2:114 (Syntypes, females; Key West, Florida; U.S. Nat. Mus.) Corthylus guayanensis Eggers, 1933, Trav. Lab. d'Ent. Mus. Nat. d'Hist. Nat. Paris, Mem. 1:22 (Syntype, male; Camopi, Franz. Guayana; Paris Mus.). New synonymy The male type and female cotype of Cor- thylus guayanensis Eggers from Camopi were compared to my series from Florida, Central America, and Venezuela, some of which were compared to the syntypes of spinifer Schwarz. All represent the same common species. Hypoth hrunneus (Hopkins) Stephanoderes brunneus Hopkins, 1915, U.S. Dept. Agric. Rept. 99:31 (Holotype, female; Brownsville, Texas; U.S. Nat. Mus.). Hypothenemus cryphalomorphus Schedl, 19.39, Proc. Roy. Ent. Soc. London 8(1): 14 (Holotype, female; Trinidad, British West Indies; British Mus. Nat. Hist.). New synonymy Stephanoderes bituberculatus Eggers, 1940, Arb. Morph. Taxon. Ent. Berlin 7:126 (Holotype, fe- male; Env. de Trois-Rivieres, Guadeloupe; Paris Mus.). New synonymy The female holotypes of Stephanoderes brunneus Hopkins, Hypothenemus cryphalo- morphus Schedl, and Stephanoderes bitu- berculatus Eggers were compared directly to my long series from Florida, Texas, and Mexico. Since only one species is represent- ed, the names cryphalomorphus and bitu- berculatus must be placed in synonymy. Monarthrum parvum (Eggers) Anchonocerus parvus Eggers, 1933, Trav. Lab. d'Ent. Mus. Nat. d'Hist. Nat., Paris Mem. 1:22 (Holo- type, male; Nouveau Chantier, Guyane Frangaise; Paris Mus.) Monarthrum praeclarum Wood, 1968, Great Basin Nat. 28:6 (Holotype, male; Manaka, British Guiana; British Mus. Nat. Hist.). Netc synonymy Because my concept of Anchonocerus was quite different from that of Eggers, I over- looked the possibility that any of his species in that genus might be allied to my Mon- arthrum praeclarum. The male holotype of parvus was directly compared to my male paratype of praeclarum. They are identical. Phloeoborus punctatorugosus Chapuis Phloeoborus punctatorugosus Chapuis, 1869, Synopsis June 1977 Wood: American Scolytidae 209 des Scolytides, p. 14 (Holotype, male Nouvelle Grenade; Mus. Roy. Nat. Beige) Phloeobortis breviusculus Chapuis, 1869, Synopsis des Scolytides, p. 14 (Holotype, male; Cayenne; Brvis- sels Mus.). New synonymy The male holotypes of Phloeobortis punc- tatorugosus Chapuis and breviusculus Cha- puis were directly compared to my series. Except for size they are identical. The name punctatorugosus is more widely known in the literature and is retained for this species. Phloeotribus pilula Erichson Phloeotribiis pilula Erichson, 1847, Archiv Naturgesch. 13(1):1.38 (Lectotype, male; Peru; Zool. Mus. Ber- lin, designated by Wood, 1973, Great Basin Nat. .33:181) Phloeotribus australis Schedl, 1953, Mem. Queensland Mus. 13:80 (Holotype, male; Queensland, Austra- lia; Schedl Coll.). New synonymy Phloeotribus australis Schedl is based on a unique male that is labeled "Queensland." Since this is the only reported member of this genus from Australia, and because the original description and accompanying illus- tration suggest a South American and not an oriental affinity, the type of this species was of unusual interest from zoogeographic- al and phylogenetic points of view. The holotype is identical in every respect to specimens compared by me to the lectotype of pilula Erichson. This species is widely distributed in Central and South America in Brosmium. The Schedl specimen was appar- ently intercepted in Australia or was mis- labeled. Phloeotribus setulosus Eichhoff Phloeotribus setulosus Eichhoff, 1868, Berliner Ent. Zeitschr. 12:149 (Lectotype, male; Colombia; Brussels Mus., designated by Wood, 1973, Great Basin Nat. 33:182) Phloeotribus dubuis Eichhoff, 1868, Berliner Ent. Zeitschr. 12:150 (Holotype?, male; Columbie; Ber- lin Mus.). New synonymy Phloeotribus spinipennis Eggers, 1930, Ent. Blatt. 26:168 (Holotype, male; Columbie, Moritz; Berlin Mus.). New synonymy The male holotypes of Phloeotribus dubuis Eichhoff and spinipennis Eggers were compared directly to one another and to my material that had previously been compared to the lectotype of setulosus Eichhoff. Except for the pale, callow color of the type of dubuis, they are identical. For this reason the names dubuis and spin- ipennis must be placed in synonymy. Pityophthorus aquilus Blackman, n. status Pityophthorus aquilus Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:.33 (Holotype, female; Kaibab National Forest, Arizona; U.S. Nat. Mus.) Bright (1977, Canadian Ent. 109:514) placed in synonymy under Pityophthorus ca- rinulatus Swaine, 1925, the names aquilus Blackman, 1928, caelator Blackman, 1928, opimus Blackman, 1928, and aristatae Bright, 1964. A review of the holotypes of all of these names plus an additional 442 specimens of these taxa, indicates that two species are represented. Pityophthorus ca- rinulatus { = opimus) is larger (2.0-2.5 mm) and it has the pronotum and male head ret- iculate. The sulcus on the elytral declivity is wider, not as deep, with the tubercles on interstriae 1 and 3 finer, and the host is Picea. The other species, designated by the senior name aquilus Blackman { = caelator, aristatae), is smaller (1.8-2.1 mm), the pro- notum and male head lack reticulation, the declivital sulcus is slightly deeper, with the tubercles on interstriae 1 and 3 larger, and the host is Pinus. Both are rather widely distributed in western North America. Pityophthorus cascoensis Blackman, n. status Pityophthorus cascoensis Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:99 (Holotype, female; Peak Island, Maine; U.S. Nat. Mus.) Bright (1977, Canadian Ent. 109:515) placed in synonymy under Pityophthorus in- textus Swaine, 1917, the names cascoensis Blackman, 1928, tonsus Blackman, 1928, kenti Blackman, 1928, and pilifer Schedl, 1931. A review of the holotypes and other type material of the above names and 1,121 additional specimens in these taxa indicates the presence of at least two and probably three .species. One, represented by the type of cascoensis {= pilifer), is of larger average size (1.6-2.2 mm), the male frons is less strongly impressed and may have a weak 210 Great Basin Naturalist Vol. 37, No. 2 median carina, the female frontal setae nev- er appear scalelike, the discal interstriae are sparsely pimctured, and declivital interstriae 1 bears a row of regularly spaced setae. The second species, represented by the type of intextus {=shepardi, tonsus), is smaller (1.4- 2.0 mm), the male frons is rather strongly, transversely impressed and never has a me- dian carina, the female frontal vestiture ap- pears to include short scales when viewed with the light source coming from the di- rection of the beetle's vertex (an illusion caused by an abrupt curve near the base of each long frontal seta), the discal interstriae are impimctate, and the setae on declivital interstriae 1 are irregularly, widely spaced or absent. The type series of kenti is in poor condition, but is considered to be syn- onymous with ornatiis Blackman, a species very closely related to intextus. Pityophthorus puHcarius (Zimmermann) Crypturgus pulicarius Zimmermann, 1868, Trans. Amer. Ent. Soc. 2:144 (Syntypes?; Lake Superior, Southern States, Tampa, and Illinois: Mus. Comp. Zool.) Pityophthorus cuhensis Schedl, 1972, Koleopt. Rundsch. 50:65 (Holotype, male; Vinales, Pinar del Rib Prov., Cuba; Schedl Coll.). New synony- my The male holotype of Pityophthorus cu- hensis Schedl and the 10 syntypes of puH- carius (Zimmermann) were compared to my series of this species. Since only one species is represented by this material the name cu- hensis must be placed in synonymy. Pityophthorus suhsimiUs Schedl Pityophthorus subsimilis Schedl, 1955, Zeit. angew. Ent. 38:25 (Lectotype, female; Ciudad Guate- mala, Guatemala; Schedl Coll.) Pityophthorus subimpressus Bright, 1977, Great Basin Nat. 36:441 (Holotype, female; 32 miles or 53 km S Valle Nacional, Oaxaca, Mexico; Canadian Nat. Coll.). New synonymy The female lectotype of Pityophthorus subsimilis Schedl was compared directly to the type series of subimpressus Bright. Although the lectotype of subsimiUs is in rather poor condition and frass obscures part of the pronotal asperities, it is clear that the asperities are essentially in obscure concentric rows. Because I see no differen- ces in other characters, they are considered to be synonymous. A long series of this species taken by me from a nonconiferous shrub is identical to the lectotype of subimilis. Several series of attenuatus Blackman from a variety of hosts in Mexico exhibit minor differences in the female frons and have the declivital punc- tures variable, but they are almost always smaller than in the lectotype of subsimiUs. Because of this variability, I have regarded subsimilis as a minor geographical variant of attenuatus. However, it appears desirable to await the collection of additional mate- rial before proposing synonymy with atten- uatus. Premnobius cavipennis Eichhoff Premnobius cavipennis Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:404 (Syntypes; Cap bonae spei in Africa meridionalis, and Colombia; lost with Hamburg Mus.) Premnobius latior Eggers, 1933, Trav. Lab. d'Ent. Mus. Nat. d'Hist. Nat. Mem. 1:9 (Holotype, female; St. Jean du Maroni, French Guiana; Paris Mus.) The female holotype of Premnobius latior Eggers was compared directly to my series of this species from Venezuela, some of which were compared to Eggers' homotypes of cavipennis Eichhoff. They are obviously of the same species. Scolytodes chapuisi, n. n. Ctenophorus laevigatus Chapuis, 1869, Synopsis des Scolytides, p. 49 (Syntypes, two males; Colombie; Brussels Mus.) As indicated above, the synonymy of Scolytodes and Ctenophorus has created homonyms of their type species. It is, there- fore, necessary to rename the junior homo- nym. I propose the new name, Scolytodes chapuisi, as a replacement for S. laevigatus (Chapuis). Scolytus propinguus Blandford Scolytus propinguus Blandford, 1896, Biol. Centr. Amer., Coleopt. 4(6): 121 (Syntypes; Mexico, Guatemala, Cuba: British Mus. Nat. Hist.) Scolytus peniciUus Schedl, 1973, Papeis Avulsos Zool. 26:165 (Holotype, male; Veracruz, Mexico: Schedl Coll.). New synonymy June 1977 Wood: American Scolytidae 211 This species is rather common in a varie- ty of host trees from the states of Nayarit and Veracruz in Mexico to Costa Rica. Sev- eral long series in which males and females are definitely associated are in my collec- tion. Several females were directly com- pared to the holotype of Scohjttis prop- inguus Blandford and are identical; the males from that series are identical to the male holotype of penicillus Schedl. Since these names represent the opposite sexes of the same species, penicillus must be placed in synonymy. Xtjlehonis dryographus (Ratzeburg) Bostrichiis dryographus Ratzeburg, 1837, Die Forstin- sekten, p. 167 (Syntypes, female; apparently Ger- many; presumably Berlin Mus., not examined) Xyleborus linearis Schedl, 1949, Rev. Brasil. Biol. 9:273 (Lectotype, female; Am. Bor.; U.S. Nat. Mus., present designation). New synonymy Xijlebortis linearis Schedl was named from a syntypic series in the Eggers collec- tion, which Eggers had described but did not publish. The syntype in the U.S. Nation- al Museum is labeled "Am. Bor.; Xyleborus linearis n. sp. Type, Eggers det. 1927." Since it was apparently intended to be the type by Eggers and since the Schedl syn- types are on loan from the Eggers collec- tion, I here designate the above "type" specimen in the U.S. National Museum as the lectotype of Xyleborus linearis Schedl. It is a specimen of the European X. dryo- graphus (Ratzeburg) that was either mis- labeled or intercepted in America and is not known to be established anywhere in America. The lectotype was compared di- rectly to my series of five specimens from Europe, which were determined by Dr. A. Pfeffer. Xyleborus horridus Eichhoff Xyleborus horridus Eichhoff, 1869, Berliner Ent. Zeitschr. 12:282 (Syntypes, female; Teapa, Mexi- co; Brussels Mus.) Xyleborus fiohri Schedl, 1972, Koleopt. Rund. 50:69 (Holotype, female; Mexico; Schedl Coll.). New synonymy The female holotype of Xyleborus flohri Schedl and the syntypes of X. horridus Eichhoff in the Chapuis collection were both compared directly to my series of this species. Since all represent the same species, Schedl's name mu.st be placed in synonymy. Xyleborus tonsus (Hagedorn), n. comb. Dryocoetes tonsus Hagedorn, 1905, Bull. Mus. d'Hist. Nat., Paris 6:412 (Holotype, female; environs de Saint-Georges, Oyapock, Guyane frangaise; Paris Mus.) Dryocoetes tonsus Hagedorn must be transferred to Xyleborus. It represents a spe- cies group within the genus previously un- known to me. New Species Araptus crassus, n. sp. This species represents a unique group in the genus that is characterized by the very stout body form which has the general habitus of Coccotrypes, by the similar frons in both sexes, by the shallow, equally large strial and interstrial punctures, by the unique antennal club, and by the broad tibiae. Female(?).— Length 2.2 mm (paratypes 2.0-2.2 mm), 1.9 times as long as wide; col- or dark brown. Frons evenly convex, epistomal margin with a weak, narrow, median suggestion of a premandibular process; surface smooth, shining, a very few minute, isolated gran- ules. Antennal club obovate, about as long as scape, about 1.15 times as long as wide, and about equally divided by rather strong- ly procurved sutures 1 and 2, these weakly grooved and marked by rows of setae, su- ture 1 septate on lateral half. Pronotum 0.94 times as long as wide; widest near ba.se, sides strongly arcuate on basal half, rather strongly constricted, then narrowly rounded in front; anterior margin armed by about four small, indefinite gran- ules; summit indefinite, on basal third; ante- rior slope rather gradual armed by low, iso- lated, abundant asperities extending almost to base in lateral areas, surface between as- perities shining, almost smooth; posterior areas shining, not smooth, punctures very fine, indefinite, sparse. Vestiture of short, fine, recumbent hair. Elytra 1.01 times as long as wide, 1.2 212 Great Basin Naturalist Vol. 37, No. 2 times as long as pronotum; sides straight and parallel on basal half, very broadly roiinded behind; striae not impressed, punc- tures moderately large, distinctly impressed, spaced by diameter of a puncture; inter- striae smooth, shining, almost three times as wide as striae, punctures as large as those of striae, a bit more widely spaced. Declivity broadly convex, very steep; sculpture about as on disc. Vestiture of minute strial hair, length of each less than diameter of its puncture, and erect, flattened, narrowly spatulate, closely set interstrial setae, each equal in length to about two-thirds distance between rows. Protibia much broader than normal for this genus, armed by three major and about six minor denticles; posterior face with sev- eral granules. Sexual differences not apparent, although both sexes apparently represented. Type Locality.— Eight km NE Cerro Jefe, Panama Pro v., Panama. Type Material.— The holotype (female?) and four paratypes were taken at the type locality on 27-III-1976, 700 m, by H. P. Stockwell. The holotype and two paratypes are in the Canadian National Collection; the other two paratypes are in my collection. Araptus obesus, n. sp. This species is distinguished from the very similar crassus Wood by the smaller size, by the absence of a median epistomal lobe, by the more strongly impressed decliv- ital striae, and by the yellowish brown col- or. Female.— Length L8 mm (paratypes L5- L8 mm), 2.1 times as long as wide; color yellowish brown. Frons as in crassus except surface more nearly subreticulate, median epistomal area with no indication of a lobe. Pronotum as in crassus except asperities largely absent from posterolateral areas, punctures on posterior areas distinctly larger, setae much stouter. Elytra as in crassus except declivital striae weakly impressed and interstrial pimctures on 1 and 3 replaced by minute granules, interstrial setae slightly wider. Type Locality.— Eighteen km NE Orixi- mina, Brazil. Type Material. The holotype (female?) and seven paratypes were taken at the type locality on 13-14-XI-1969, by J. M. and B. A. Campbell. The holotype and fovir paratypes are in the Canadian National Collection; three paratypes are in my collection. Cnesinus prominulus, n. sp. This species is distinguished from the closely allied minax Schedl by the larger size, by the more slender body, by the more abundant, shorter vestiture, and by other minor details. Female.— Length 3.0 mm (paratypes: fe- male 2.7-3.3 mm, males 2.4-3.1), 2.7 times as long as wide. As in minax except as indicated above. Type Locality.— Volcan Colima, Jalisco, Mexico. Type Material.— The holotype, allotype, and 29 paratypes were taken at the type lo- cality on 23-VI-1965, 2500 m. No. 107, from twigs of a shrub, by me. The holotype, allotype, and most of the paratypes are in my collection. Two para- types are in the British Museum (Natural History). Conophthorus conicolens, n. sp. This species is distinguished from apa- checae Hopkins and ponderosae Hopkins by the slightly stouter body form, by the more strongly, more broadly impressed declivity, with the tubercles on interstriae 3 larger, by the larger punctures on dechvital striae 3, and by the smoother elytral disc, with the punctures averaging larger and somewhat confused. Female.— Length 3.7 mm (paratypes 3.3- 3.8 mm), 2.2 times as long as wide; color very dark brown. Frons and pronotum as in mexicanus June 1977 Wood: American Scolytidae 213 Wood except crenulations on pronotal disc distinctly smaller. Elytra about as in ponderosae except dis- cal punctures larger, slightly confused, de- clivital sulcus distinctly deeper, lateral con- vexities distinctly higher than suture, tubercles on interstriae 3 larger. Male.— Similar to female except punc- tures on frons apparently larger. Type Locality.— Eight miles or 13 km W Texmelucan, Puebla, Mexico. Type Material.— The female holotype, male allotype, and 10 paratypes were taken at the type locality on 13-VI-1967, 2500 m. No. 25, from Piniis cones, by me. Other paratypes from Mexico include: 4 from Tlalmanalco, Mexico, 24-XI-1949, #13A, Pinii.s leiophijlla, J. P. Perry, Jr.,; 2 from Tulancingo, Hidalgo, 24-VI-1953, Pinus, by me. The holotype, allotype, and paratypes are in my collection. Microcorthijliis vicinus, n. sp. This species is distinguished from min- imus Schedl by the larger size, by the larger epistomal tubercle, by the more strongly reticulate elytra, and by the slightly narrower declivital impression. Female.— Length 2.0 mm (paratypes 1.5- 2.0 mm), 2.9 times as long as wide; color brown. Frons as in minimus except epistomal tu- bercle distinctly larger and more nearly carinate. Pronotum and elytra about as in mini7nus except elytra strongly reticulate, strial punc- tures more distinctly indicated, declivity less strongly, more narrowly impressed, lateral elevations ending more remote from costal margin. Male.— Similar to female except antennal club without long setae on posterior face. Type Locality.— Thirty-two miles or 51 km S Valle Nacional, Oaxaca, Mexico. Type Material.— The female holotype, male allotype, and 34 paratypes were taken at the type locality on 21-V-1971, by D. E. Bright. Additional paratypes from Mexico include: 9 from 3 miles or 5 km N Such- ixtepec, Oaxaca, on Highway 175, 4-VI- 1971, 7000 ft., Salix, D. E. Bright; 20 from 8 miles or 13 km E San Cristobal, Chiapas, 6-VI-1969, D. E. Bright; 1 from Mt. Tzon- tehuiz, Chiapas, 12-VI-1969, Quercus, D. E. Bright. The holotype, allotype, and most of the paratypes are in the Canadian National Col- lection; the remaining paratypes are in my collection. Pityophthorus alnicolens, n. sp. This species is distinguished from alni Blackman by the smaller size, by the reti- culate, dull pronotum, by the smaller, less deeply impressed strial punctures, and by the proportionately wider discal interstriae which have numerous, irregular, impressed lines. Male (?).— Length 1.5 mm (paratypes 1.5-1.7 mm), 2.6 times as long as wide; col- or dark brown. Frons broadly convex, flattened on lower half, a very feeble median carina from epis- toma to upper level of eyes; epistomal mar- gin slightly elevated; vestiture sparse, short, inconspicuous. Antennal club as in alni ex- cept slightly smaller, slightly more slender. Pronotum much as in alni except discal area strongly reticulate, punctures coarser, margins not asperate; asperities absent from posterior third. Elytra much as in alni except striae not impressed, punctures much smaller, not as deep, interstriae three times as wide as striae, with fine, irregular, impressed lines and points, with fine, rather sparse, uniseri- ate punctures. Declivity about as in alni ex- cept punctures much finer. Vestiture about as in alni except much longer. Type Locality.— Highway 131, 115 miles or 184 km S Oaxaca, Oaxaca, Mexico. Type Material.— The holotype (male?) and four damaged paratypes in poor condi- tion were taken at the type locality on 27- 30-V-1971, 6000 ft., in Alnus, by D. E. Bright. 214 Great Basin Naturalist Vol. 37, No. 2 The holotypes and two paratypes are in the Canadian National Collection; two par- atypes are in my collection. Pityophtfwrns a.ssitus, n. sp. Tliis species is distinguished from morosus Wood by the larger size, by the stouter body, by the strongly reticulate head and pronotum, and by the more strongly im- pressed declivital interstriae 2. Female.— Length 2.1 mm (paratypes 1.9- 2.2 mm), 2.6 times as long as wide; color dark brown. Frons and pronotum as in morosus except reticulate. Elytra as in morosus except strial punc- tures smaller, not as deep, in slightly irregu- lar rows, a few interstrial punctures present on posterior half of disc, declivital inter- striae 2 more strongly impressed on its lat- eral half, strial setae slightly longer. Male.— Similar to female in all respects. Type Locality.— Lagos des Colores, Chiapas, Mexico. Type Material.— The female holotype, male allotype, and 59 paratypes were taken at the type locality on 17-V-1969, by D. E. Bright. Six specimens from 15 miles or 24 km S Valle Nacional, Oaxaca, Mexico 20-V- 1971, 4000 ft., D. E. Bright, are not includ- ed in the tvpe series. The holotype, allotype, and most of the paratypes are in the Canadian National Col- lection; the remaining paratypes are in my collection. Fityopfithorus concinnus, n. sp. This species is distinguished from coro- nariiis Blackman by the larger size, by the more strongly impressed male frons, and by the longer, darker, more abundant setae on the female frons. Female.— Length 1.9 mm (paratypes 1.7- 2.3 mm), 2.7 times as long as wide; color dark yellowish brown. Frons similar to coronarius except a bit more strongly impressed; vestiture reddish brown, longer, much more abundant; long- est setae equal in length to half distance be- tween eyes. Pronotum and elytra as in coronarius ex- cept lateral convexities on declivity con- spicuously higher than suture. Male.— Similar to female except frons rather strongly impressed from epistoma to upper level of eyes, upper margin of im- pressed area marked by a conspicuous, transverse carina. Type Locality.— "Plaquepaque, Mexico." Type Material.— The female holotype, male allotype, and 14 paratypes were inter- cepted at Nogales, Arizona (No. 49236) in imidentified wood from the type locality on 21-X-1941, Lot No. 41-18984. Four para- types are labeled: Mexico, 22-III-1962, Cnimp Coll., unidentified wood, Boston 27132, No. 62-11725. The holotype, allotype, and 11 paratypes are in the U.S. National Museum; two para- types are in the Canadian National collec- tion; and live paratypes are in my collec- tion. Pitijophthorus indigens, n. sp. This species is distinguished from fusciis Blackman by the shorter vestiture on the frons, by the more smooth, shining elytra, with smaller punctures, and by the deeper declivital sulcus. Female.— Length 2.2 mm (paratypes 2.0- 2.3 mm), 2.8 times as long as wide; color very dark brown. Frons broadly convex, a weak, median, subcarinate tubercle on epistomal margin; surface smooth, shining, punctures coarse, deep, uniformly distributed; vestiture min- ute, much shorter than in fuscus. Pronotum about as in fuscus except disc smooth, shining, punctures replaced by iso- lated, rounded granules. Elytra about as in fuscus except smooth, shining, discal punctures minute, almost ob- solete, declivital sulcus distinctly deeper. Male.— Similar to female except median carina on frons very weakly indicated from epistoma half distance to upper level of eves. June 1977 Wood: American Scolytidae 215 Type Locality.— Two miles or 3 km E Gould, Jackson Co., Colorado. Type Material.— The female holotype, male allotype, and three paratypes were taken at the type locality on 12-VI-1968, No. 8, from a Pinus contorta branch, by me. Pityophthorus leechi, n. sp. This species is distinguished from scalptor Blackman by conspicuous characters of the female frons as indicated below. Female.— Length L4 mm (paratypes 2.0- 2.5 mm), 2.8 times as long as wide; color reddish brown. Frons tranversely impressed on area be- low upper level of eyes, very feebly con- cave on median half of impressed area; sur- face smooth, shining, punctures moderately coarse, almost uniformly distributed, inter- spaces about equal to diameter of a punc- ture; a weak epistomal process clearly in- dicated; vestiture much less abundant than in scalptor, almost uniformly distributed, rather short, only slightly longer in marginal areas, longest setae about equal to one-third distance between eyes. Pronotum and elytra as in scalptor, per- haps less brightly shining and elytral vesti- ture very slightly longer. Male.— Similar to male scalptor except transverse frontal impression not as strong, median carina less strongly elevated. Type Locality.— Two miles or 3 km NNE Angwin on N side of Howell Mt., Napa Co., California. Type Material.— The female holotype, male allotype, and 27 paratypes were taken at the type locality on 5- VI- 1977, reared from a shaded-out branch of Pinus ponder- osa, by H. B. Leech. The holotype, allotype, and some para- types are in the California Academy of Sci- ences Collection; the remaining paratypes are in the Canadian National Collection and in my collection. Pityophthorus solatus, n. sp. This species is distinguished from solers Blackman by the different frons and elytra as described below. Female.— Length 2.1 mm (paratypes 1.9- 2.2 mm), 2.7 times as long as wide; color very dark brown. Frons similar to solers except plano- convex, punctured pubescent area extending to well above eyes, vestiture slightly more abundant and longer. Pronotum and elytra as in solers except surface of elytral disc smoother, declivital sulcus slightly wider and deeper, its surface more nearly microreticulate, declivital vesti- ture distinctly longer. Male.— Similar to female except frons as in male solers, declivity similar to male so- lers except sulcus narrower, lateral elevation distinctly higher. Type Locality.— Fifty-one miles or 81 km NW Oaxaca, Oaxaca, Mexico. Type Material.— The female holotype, male allotype, and 12 paratypes were taken at the type locahty on lO-V-1971, 7500 ft., Quercus, by D. E. Bright. Since all allied species are from con- iferous hosts, the type was probably either mislabeled or its occurrence in Quercus was accidental. Since Bright also collected in potential hosts at Cerro Potosi (Nuevo Leon) and in the Popocatepetl-Iztaccihuatl area (Mexico), it is possible that the type series was taken at a locality north of that designated; the relationship to solers sug- gests such a possibility. The holotype, allotype, and most of the paratypes are in the Canadian National Col- lection; the remaining paratypes are in my collection. Pityophthorus speciosus, n. sp. This species is distinguished from assitus Wood by the smooth, shining frons which is transversely impressed on its lower half and by differences on the pronotum and elytra as described below. Female.— Length 1.8 mm (paratypes 1.7- 216 Great Basin Naturalist Vol. 37, No. 2 2.0 mm), 2.5 times as long as wide; color reddish brown. Frons smooth, shining, punctures coarse, close, deep; transversely impressed from epistoma to slightly below upper level of eyes, upper area convex; vestiture short, in- conspicuous. Pronotum about as in assitus except transversely much more broadly arched, smooth, shining, with numerous, rather large, impressed points, punctures very small. Elytra about as in assitus except surface smooth, brightly shining, declivital inter- striae 2 strongly impressed and narrower. Male.— Similar to female in all respects. Type Locality.— Fifteen miles or 24 km S Valle Nacional, Oaxaca, Mexico. Type Material.— The female holotype, male allotype, and 7 paratypes were taken at the type locality on 20-V-1971, 4000 ft., by D. E. Bright. One paratype is from 92 miles or 147 km N Oaxaca, Oaxaca, Mexico, 18-V-71, at blacklight, D. E. Bright. The holotype, allotype, and most of the paratypes are in the Canadian National Col- lection; the remaining paratypes are in my collection. Pseiidopitijophthorus denticulus, n. sp. This species is distinguished from gran- ulifer Wood by the different frons, by the less definite discal striae, by the more wide- ly spaced, more slender interstrial bristles, and by the near absence of granules and bristles on declivital interstriae 2. Female.— Length 1.9 mm (paratypes 1.6- 2.0 mm), 2.9 times as long as wide; color dark brown. Frons as in granulifer except more shal- lowly, more broadly concave, punctures smaller, more widely spaced. Pronotum and elytra as in granulifer ex- cept strial punctures on disc in less definite rows, erect interstrial setae more slender, more widely spaced, spaced by distance equal to length of a seta, setae and granules absent from declivital interstriae 2, one or two granules sometimes present near base or apex on 2. Male.— Similar to female in all respects except antennal club sometimes more slen- der. Type Locality.— Boot Springs, Big Bend National Park, Brewster Co., Texas. Type Material.— The female holotype, male allotype, and 17 paratypes were taken at the type locality on 20- VII- 1974, from Quercus, by D. E. Bright. The holotype, allotype, and most of the paratypes are in the Canadian National Col- lection; the remaining paratypes are in my collection. Pseudothijsanoes securus, n. sp. This species is distinguished from quer- cicolens Wood by the much longer, more slender, flattened process on the female scape, by the more slender, more acutely pointed antennal club, by the more slender, more isolated pronotal asperities, and by many other characters. Female.— Length 1.5 mm (paratype 1.5 mm), 2.9 times as long as wide; color black, vestiture white. Frons about as in quercicolens except sur- face rugose-reticulate. Scape acutely triangular, twice as wide as long, orna- mented by a conspicuous tuft of rather long hair; club small, twice as long as wide, acutely pointed at apex, sutures obscure. Pronotum about as in quercicolens except anterior margin unarmed, asperate area smaller, asperities smaller, more slender. Elytra about as in quercicolens except strial punctures not as deep, interstriae more irregular, punctate granules larger, closer; declivital striae 1 and 2 with punc- tures reduced, interstriae 2 narrower, flat, 3 with weak elevation not reaching junction with 9. Vestiture white, closer, interstrial scales on disc one and one-half to two times as long as wide, on declivity about as long as wide. Type Locality.— Five miles or 8 km W Tulancigo, Hidalgo, Mexico. June 1977 Wood: American Scolytidae 217 Type Material.— The female holotype and one female paratype were taken at the type locality on ll-VI-1967, 2400 m. No. 12, from a Quercus branch, by me. The holotype and paratype are in my collection. Psetidothysanoes turnbowi, n. sp. This species is distinguished from recamis Wood by the smoother, more brightly shin- ing pronotum and elytra, by the less deeply, more extensively excavated female frons which has punctures and small, stout setae in the excavated area, and by the more slender scales on the male declivity. Female.— Length 1.3 mm (paratypes 1.2- 1.4 mm), 2.3 times as long as wide; color very dark brown. Frons similar to recavus except less strongly, more extensively impressed, sur- face of impressed area smooth, shining, with sparse punctures and stout setae. Pronotum as in recavus except surface on posterior areas smooth, not reticulate. Elytra as in recavus except smoother, in- terstrial setae slightly longer, more slender. Male.— Similar to male of recavus except differing by smoother frons, pronotum, and elytra and by the much more slender declivital setae, each about three times as long as wide. Type Locality.— Bentsen-Rio Grande State Park, Hidalgo Co., Texas. Type Material.— The female holotype, male allotype, and 67 paratypes were reared from Prosopis glandulosa branches by Robert H. Turnbow, Jr., with emergence dates ranging from 29-31-XII-1975, 9-13, 21-23 (type), 27-29-1, 1-5, 11-18-VII, 2-8- VIII- 1976. One paratype is labeled; Hidalgo Co., Texas, 26-III-57, D. J. and J. N. Knull. The holotype, allotype, and most of the paratypes are in my collection, other para- types are in the University of Georgia col- lection and the Canadian National Collec- tion. Pijcnarthrum fulgidum, n. sp. This species is distinguished from lucidum Wood by the larger size, by the less regu- larly punctured pronotum, by the smaller, less strongly impressed punctures on the elytral striae, and by the much finer vesti- ture. Male.— Length 1.8 mm, 2.0 times as long as wide; color brown. Frons shallowly, broadly, transversely im- pressed, not concave; surface shining, slightly irregular, punctures fine, rather sparse, not clearly defined; vestiture short, rather sparse, almost hairlike. Eyes more finely faceted and more widely separated than in lucidum. Pronotum much as in lucidum except slightly more declivous on anterior third, surface not as smooth, pmictures slightly larger, not as regular in size. Elytra some- what as in lucidum except striae not im- pressed, punctures much smaller, less strongly impressed, interstriae three to four times as wide as striae, ground vestiture very sparse, hairlike, as fine as strial setae, except interstrial setae slightly longer and much more slender than in lucidum, not stouter on declivity. Female.— Similar to male except frons convex. Type Locality.— Eight km S Colonia, Valle de Cauca, Colombia. Type Material.— The male holotype, fe- male allotype, and seven paratypes were taken at the type locality on 9- VII- 1970, 30 m. No. 650, from Guarea trichiloides, by me. The holotype, allotype, and paratypes are in my collection. Scolytodes fulmineus, n. sp. This species is distinguished from ama- bilis Wood by the larger size, by the more strongly reticulate pronotum, with finer punctures, and by the very different female frons. Female.— Length 1.8 mm (paratypes 1.6- 1.8 mm), 2.4 times as long as wide; color black. Frons weakly convex from epistoma to above upper level of eyes, median third 218 Great Basin Naturalist Vol. 37, No. 2 STnooth, shining, impunctate, lateral thirds rather finely, closely punctured; vestiture confined to punctured areas, fine, short, longest setae about equal in length to diam- eter of three facets of eye. Pronotum about as in amabilis except surface strongly reticulate, punctures much smaller but slightly larger than in clusiae Wood. Elytra as in ambilis except vestiture con- fined to odd-numbered interstriae, not more than three or four setae on each of these. Male.— Similar to female except frons more strongly convex, punctures rather sparse, uniformly distributed, vestiture sparse, inconspicuous. Type Locality.— La Carbonera Experi- mental Forest, about 50 km (airline) NW Merida, Merida, Venezuela. Type Material.— The female holotype, male allotype, and 11 paratypes were taken at the type locality on 14-X-69, 2500 m, No. 50, from Clusia branches, by me. Scolytodes radiatus, n. sp. This species is distinguished from clusiae Wood and volcanus Wood by the slightly larger size, by the much coarser pronotal punctures, and by the very different female frons. Female.— Length 1.7 mm (paratypes 1.6- 1.9 mm), 2.4 times as long as wide; color al- most black. Frons convex and virtually glabrous as in males of allied species. Pronotum much as in volcanus except punctures much larger, each equal to from half to full diameter of a strial puncture (variable). Elytra about as in amabilis Wood except punctures larger and vestiture less abun- dant; striae not impressed, punctures rather coarse, deep; interstriae slightly wider than striae, smooth, shining, punctures uniseriate, less than half diameter of those of striae. Subglabrous, a few fine setae on odd-num- bered interstriae on posterior half. Male.— Similar to female in all respects. Type Locality.— La Georgiana, 79 km SE San Jose, Costa Rica. Type Material.— The female holotype, male allotype, and 13 paratypes were taken at the type locality on 31-VII-1965, from Quercus, by L. S. Otvos. The holotype, allotype, and paratypes are in my collection. Xyleborinus tribuloides, n. sp. This species is distinguished from bicorna- tulus Wood by the uniformly reticulate pro- notum and by the much shorter, less strong- ly impressed, very different declivity. Female.— Length 2.1 mm (paratype 2.3 mm), 2.9 times as long as wide; color dark brown. Frons and pronotum about as in bicorna- tidus except pronotal disc uniformly, strong- ly reticulate. Elytra 1.7 times as long as wide, 1.5 times as long as pronotum; declivity con- fined to posterior third; striae not im- pressed, punctures fine, shallow; interstriae smooth, shining, three to four times as wide as striae, punctures finely granulate. De- clivity gradual, shallowly subconcave; punc- tures on striae 1 and 2 slightly larger than on disc; interstriae 1 and 2 imarmed except for small granules at base, 3 armed one- third declivital length from base by a mod- erately coarse, pointed tubercle, as high as wide, a larger, blunt spine two-thirds of declivital length from base, longer than wide, its length about equal to width of an interstriae, a small tubercle at base above spine 1 and another at apex below spine 2; lateral summit at striae 4, lateral areas armed by about a dozen moderately coarse tubercles (smaller and less numerous than in hicornatulus. Vestiture of minute strial hair and distinctly longer interstrial hair, regu- larly, closely spaced to base, each interstrial seta about equal in length to width of an interstriae. Type Locality.— Fifteen miles or 24 km S Valle Nacional, Oaxaca, Mexico. Type Material.— The female holotype and one female paratype were taken at the June 1977 Wood: American Scolytidae 219 type locality on 20-V-1971, from a Cecropia petiole, by D. E. Bright. The holotype is in the Canadian National Collection; the paratype is in my collection. Xyleborus devexus, n. sp. This species is distinguished from ob- liquus (LeConte) by the smaller size, by the coarser, deeper elytral punctures, and by the absence of tuberct?^ on declivital inter- striae 1 and 3. Female.— Length 1.9 mm (paratypes 1.8- 2.1 mm), 2.5 times as long as wide; color very dark brown. Frons and pronotum as in obliquus. Elytra as in obliquus except declivity slightly steeper, less strongly convex; punc- tures on disc larger, deeper, interstriae twice as wide as striae; interstrial punctures near declivity not granulate. Declivity more distinctly impressed between interstriae 3; interstriae 1-3 without granules. Vestiture as in obliquus. Type Locality.— Homestead, Florida. Type Material.— The female holotype and three female paratypes were taken at the type locality on 27-X-1957 on Burdiken palm. Additional female paratypes were taken as follows: 3 from Orange Heights, Florida, 3-V-1914, Hopk. US 9925i, pecan, E. W. Berger; 1 from Lake Co., Florida, 21-1-1930, B. L. Smith; 1 from Manatee Co., 2-1 V- 1930, R. F. Tinker; 1 from Or- ange Co., Florida, 24-1-1930, W. M. Loe; 1 from Orange Co., Florida, 22-IIM930, J. E. Sadler; 2 from Pinellas Co., Florida, 22-III- 1930 and 12-IV, B. P. Moore; 15 from Rib Grande, Puerto Rico, 26-1 V- 1940, Cedrella mexicana, Hopk. US 33100-V-l, D. De- Leon; 4 from Vega Alta, G. N. W. (presum- ably Puerto Rico), 16-XI-1915, ace. no. 170- 15. The holotype and several paratypes are in my collection; the remaining paratypes are in the U.S. National Museum of Natural History. Xyleborus kiticeps, n. sp. This species is distinguished from retusus Eichhoff by the slightly smaller size, by the nonreticulate, much more coarsely punc- tured pronotal disc, and by the different declivity as described below. Female.— Length 3.5 mm (paratypes, fe- males 3.1-3.6 mm, males 1.7-2.0 mm), 1.7 times as long as wide; color black. Frons as in retusus except more coarsely pimctured. Pronotum as in retusus except posterior areas usually smooth, shining, coarsely, closely punctured, interspaces in discal area equal to about half diameter of a puncture (reticulate and equal to two to four diame- ters in retusus). Elytra as in retusus except discal punc- tures slightly larger, margin of declivity with a subacutely elevated circumdeclivital ring forming a complete circle (forming about two-thirds of a circle in retusus), punctures on declivital face at least twice as large, surface with few to many irregu- lar, serpentous impressed lines (these lines absent in retusus). Male.— Similar to female except dwarfed, deformed; pronotum and elytra more gradually declivous; all features less sharply defined. Type Locality.- Twenty km SW El Vigi'a, Merida, Venezuela. Type Material.— The female holotype, male allotype, and 33 paratypes were taken at the type locality on 21-XI-69, 50 m. No. 145, from Jacaranda poles, by me. (The al- lotype and 17 of these paratypes were reared at and incorrectly labeled Merida, 22-IX-69, 1700 m.) Additional paratypes from Venezuela include: 12 from 9 km S Barancas, Barinas, l-X-69, 150 m. No. 24, Protium tenuifoliwn, 9 more with the same data except 5-XI-69, Inga sp., Spondias mombin, or Dendropanax arboreum, 1 more with the same data except 2-XII-69, Meli- coccous bijugata; 1 from 17 km SE Miri, Barinas, 17-XII-69, 150 m. No. 195, Pro- tium; 1 from 10 km SE Miri', 8-n-70, 150 m, No. 295, Inga; 3 from 8 km SW Bum- bum, Barinas, 11-11-70, Moraceae, Protium tenuifolia, or "Tartaguito"; 6 are from 40 km E Canton, Barinas, 8-III-70, from various 220 Great Basin Naturalist Vol. 37, No. 2 hosts or in flight; all were taken by me. J. L. Saunders. Three paratypes are from Valle de Choroni, The holotype, allotype, and paratvpes are Venezuela, 3-1 V- 1964, Theohroma cacao, in my collection. SUPPLEMENTARY NOTE ON VENEZUELAN BAT FLIES (DIPTERA: NYCTERIBIIDAE) Lindolpho R. Guimaraes' Abstract.— Basilia dtinni Curran (1935) is reported from Venezuela for the first time, and comments are made on four other indigenous species. After the publication of Venezuelan Nyc- teribiid Bat Flies (Guimaraes 1972) I re- ceived 17 additional vials with 33 speci- mens of nycteribiids from Dr. Vernon J. Tipton. They were collected in the same area where the previous collection was made. The cumulative total number of specimens collected by the personnel of the Smithsonian Venezuelan Project (SVP) is 166, from 94 host specimens. There are five species represented in the new material, of which one had been recorded previously from Venezuela and one, Basilia diinni Cur- ran (1935), is reported for the first time. Basilia anomala Guimaraes and D'Andretta, 1956 Fig. 1 Basilia anomala: Guimaraes, 1972: 1 Present record.— One female ex RJio- geessa tumida (SVP 11109), Miranda: 5 km E Rio Chico, near Puerto Tuy, 6-XI-1966. Remarks.— This is the second known fe- male of the only South American species with three tergal plates. The abdomen of the female figured in the original description (Guimaraes and D'Andretta 1956) was swollen, and so some- what different from the specimen figured herein, chiefly with regard to the shape of tergal plates I and III. In this specimen, the abdomen is rather shrunken, the lateral margins of tergal plate I are rolled up, and the posterior margin looks bilobate, due to a notch on the midline; viewed under the stereomicroscope, the midline of the tergite appears depressed. Tergal plate III is more regular: the lateral margins converge to- ward the back, and the posterior margin is slightly emarginate. id D'Andretta, Basilia dubia Guimaraes 1956 Basilia dubia: Guimaraes, 1972: 3 Present records.— Two females, ex Myotis albescens (SVP 16173), Amazonas: \ 1 de Zoologia, Universidade de Sao 1 Fig. 1. Basilia anomala, abdomen of female, dorsal view (SVP 11109). 221 222 Great Basin Naturalist Vol. 37, No. 2 Belem, Rio Cunucununia, 150 m elev., l-II- 1967; 3 females and 2 males, ex Myotis al- bescens (SVP 16210), Amazonas: Belem, Rio Cunucunuma (mouth Caiio Culebra), 150 m elev., 2-II-1967; 3 females, ex Myotis albes- cens (SVP 19515 and 19634), Amazonas: 1 km down Casiquare Canal from Capibara, 130 m elev., 6-VI-1967 and 13-VI-1967. Remarks.— The characters that dis- tinguish this species from B. carteri show some variability. The females mentioned in my 1972 paper had a less cordiform tergal plate I than specimens previously known; one specimen, from Apure, had no pustulate setae on the lateral connexivum. Among the females now examined, the majority of which have tergal plate I perfectly cordi- form, I found one specimen with the mar- gin truncate, and one specimen with a much more elongate plate than the others. In some specimens the pustulate setae of the lateral connexivum are so few (3-4) and so small that they are hardly visible. Basilia ferrisi Schuurmans-Stekhoven, 1931 Basihi ferrisi: Guimaraes, 1972: 7 Present records.— One female (SVP 43462) ex Myotis simus, Monagas: Mata de Bejuco, 54 km SE Maturin, 18 m elev., 3- VI-1968; two females (SVP 44232), ex Myotis nigricans. Falcon: Capatarida, 40 m elev., 26-VI-1968. Basilia dunni Curran, 1935 Fig. 2 Bmilia dunni Curran, 1935: 3, Figs. 1, 2; Scott, 1936: 497 (citation); Del Ponte, 1944: 118, 124 (citation and key); Guimaraes, 1946: 16, 20 (citation and key); Guimaraes and D'Andretta, 1956: 26 and 95, Figs. 152, 153 (key and redescription of holotype); Maa, 1965: 380 (included in the subgenus Pseude- lytromyia); Guimaraes, 1966: 399, Fig. 37 (com- ments and description of male) Guimaraes, 1968: 101.2 (citation). Previous records and hosts.— Panama: Santa Rosa, ex Myotis nigricans (holotype); Canal Zone, Juan Mina, ex Myotis n. nigri- cans; Darien, Rio Tuira, ex Myotis albes- cens. Present records.— Five females (SVP 29146 and 29149), ex Myotis albescens, Amazonas: almost directly across Rio Mana- piare from camp, 155 m elev., 26-VII-1967; two females (SVP 30603), ex Myotis albes- cens, Amazonas: Paria, 25 km SEE Puerto Ayacucho, 114 m elev., 13-IX-1967. Remarks.— B. dunni is here recorded for the first time from Venezuela; it was known only from Panama. The holotype was found on Myotis nigricans; all other records were from Myotis n. nigricans (1) and from Myotis albescens (4). The two figures of the female of this species (Curran 1935, Guimaraes and D'Andretta 1956) were based on the holo- type. In the specimen now figured the dis- cal setae of tergal plates I and II are fewer than in the holotype. The male was de- scribed and figured by me in 1966. In my key to females of Venezuelan spe- cies of Basilia (Guimaraes 1972) B. dunni Fig. 2. Bmiliii view (SVP 29146). ditnt abdomen of female, dorsal June 1977 GuiMARAEs: Bat Flies 223 should appear in the last couplet, after B. ferrisi, as follows: "Tergal plate II with dis- cal setae in irregular rows converging obliquely toward the midline; longer setae of posterior margin of tergal plate I not reaching the posterior margin of tergal plate II." Basiliu otiizi Machado-Allison, 1963 Figs. 3, 4, 5 Ba.siha ortizi: Giiimaraes, 1972: 2 Present records.— Five males and seven females ex Eptesicus (SVP 25562A, 26765, 26767, 26768, 26770, 26771, and 26773), Fig. ,3. BiViiUa ortizi, spiniform setae of the posterior margin of the fourth visible stemite of the male. Amazonas: San Juan, Rio Manapiare, 155 m elev., 9 and 15-VII-1967. Remarks.— As previously noted (Gui- maraes 1972), this species closely resembles B. bequaerti. It is indeed difficult to believe that it is a good species, but only the exam- ination of -Specimens from the same host and area will allow a firm decision. On the basis of the specimens seen, it seems to me that the setae of the lateral connexivum, especially the outer ones, are a little longer in B. ortizi than in B. bequaerti. The posterior processes of tergal plate II are narrower in ortizi, although not as thin as those of B. wenzeli. The number of setae on this process is also quite variable. Guimaraes and D'Andretta (1956) state that B. bequaerti has from four to six spiniform setae and from two to four long and robust setae. Fig. 4. Basilia ortizi, genitalia of the male, lateral. Fig. .5. Basilia ortizi, abdomen of the female, dorsal view (SVP 26771). 224 Great Basin Naturalist Vol. 37, No. 2 Thev illustrate one specimen with six spin- iforni setae and three long setae on one of the processes and, respectively, four and four on the other. The drawing of the type by Machado-Allison (1963) shows three spiniforms on one side and four on the oth- er, and two long setae on either side. The specimens I am identifying as ortizi have at most two long setae; the specimen now fig- lued has two spiniform and two long setae on one side, respectively, and seven and one on the other side. Differences in the posterior elevation of the mesonotum and in the shape of the anal segment are probably due to preservation. In the figured female the genital and adanal plates are similar to those of B. be- quaerti. The aedeagus of the figured male resem- bles that of B. beqiiaerti in the serration of the dorsal margin, but its distal end is thick- er, while the parameres are much thinner distally than in B. beqiiaerti. However, since little is known of the variability of the male genitalia in Basilio, the significance of the differences mentioned is not clear. Literature cited CuRRAN, C. H. 1935. New species of Nycteribiidae and Streblidae (Deptera). American Museum Novi- tates No. 765: MI, 12 figs. Del ponte, E. 1944. Basilia romahai, nobis (Diptera, Nycteribiidae) de la Argentina. Annales del In- stitute de Medicina Regional, Tucuman, Argen- tina I (1):117-128, ,3 figs. GuiMARAEs, L. R. 1946. Revisao das especies sulameri- canas do genero Basilia (Diptera, Nycteribiidae). Arquivos de Zoologia do Estado de S. Paulo, Bra- sil 5:1-88, 98 figs., 1 map. 1966. Nycteribiid batflies from Panama (Dip- tera, Nycteribiidae), p. 393-404, figs. .35-37. In R. L. Wenzel and V. J. Tipton, editors. Ectopara- sites of Panama, Chicago, Illinois). 1968. Family Nycteribiidae, p. 1-8. In A Cata- logue of the Diptera of the Americas South of the United States. Museo de Zoologia, Universidade de Sao Paulo, No. 101. 1972. Venezuelan Nycteribiid Batflies (Diptera, Nycteribiidae). Brigham Young University Sci. Bull. Biol. Ser. 17 (I): 1-2, 2 figs. GuiMARAEs, L. R., AND M. A. d'Andretta. 1956. Si- nopse dos Nycteribiidae (Diptera) do Novo Mun- do. Arquivos de Zoologia do Estado de S. Paulo, Brasil 10:1-184, 232 figs., 3 tables, 5 maps. Machado-Allison, C. E. 1963. Un nuevo Nycteri- biidae (Diptera) de Venezuela. Acta Biologica Venezuelica 3:455-459, 6 figs. Schuurmans-Stekhoven, J. H., Jr. 1931. Eine seltene, ungeniigend beschriebene Basilia-Ar^ (Diptera Pupipara) aus Venezuela. Zeitschrift fiir Para- sitenkund 3 (2):205-219, 6 figs. Scott, H. 1936. Descriptions and records of Nycteri- biidae (Diptera Pupipara), with a discussion of the genus Basilia. Linnean Society's Journal— Zo- ology 39 (267): 479-505, 11 figs. Theodor, O. 1967. An illustrated catalogue of the Rothschild Collection of Nycteribiidae (Diptera) in the British Museum (Natural History) with keys and short descriptions for the identification of subfamilies, genera, species, and subspecies. British Museum (Natural History), vii -t- 506 pp., 898 figs., 5 pis., 6 maps, London. THE SYSTEMATICS OF CROTAPHYTUS WISLIZENI, THE LEOPARD LIZARDS. PART IIL THE LEOPARD LIZARDS OF THE GREAT BASIN AND ADJOINING AREAS, WITH A DESCRIPTION OF A NEW SUBSPECIES FROM THE LAHONTAN BASIN Wilmer W. Tanner' and Benjamin H. Banta^ Abstract.— A general analysis of the populations of Crotaphytus wislizeni in the Great Basin is presented. A new subspecies, C. wislizeni maculosus, for the Lahontan Basin, Nevada, is described and comparisons of color pattern, some scale patterns, and skull measurements are presented. In Part I of this .series we (Tanner and Banta 1963) reported the taxonomic status of Crotaphytus wislizeni wislizeni and de- scribed the subspecies C. iv. punctatus from the Upper Colorado River Basin. Part II (Banta and Tanner 1968) dealt with the leopard lizards of Baja California, Mexico, with the description of the subspecies C. w. neseotes from Cedros Island. The present study is concerned with the taxonomy of the populations occurring in the Great Ba- sin and adjoining areas west of the Colorado River. With a few exceptions, leopard liz- ards from this large area have been referred to as C. w. wislizeni. However, we note real differences between some populations in this area and those seen from areas to the east (Arizona and eastern Utah), the south (Baja California), and the west (Central Cal- ifornia). The Great Basin is a large geographical area lying between the Rocky Mountains and high plateaus of central Utah on the east and the Sierra Nevada Mountains to the west. Ingress into this area has occurred from the south and southeast with lizards from the southern deserts extending their ranges northward either along the desert ranges or the alluvial valleys. The leopard lizards have used the desert valleys to ex- tend their ranges as far north as southern Oregon and southwestern Idaho. Specimens from the Great Basin seen by us while preparing the two previous reports were reexamined, and the data are now being added to that pertinent to this report. We have received on loan from Dr. John Wright, Los Angeles County Museum (LACM), a large series of specimens from the southern parts of the Great Basin. These and those seen from other collections (see previous reports) form the basis for this study. Several collections have been made by the authors or their students, in areas not well represented by existing collections, for the purpose of procuring live material. We are grateful to the following for mate- rials sent on loan from critical areas: Mr. James Davis, Indio, California; Dr. Steven C. Anderson, Pyramid Lake, Nevada; Dr. Nathan M. Smith, southern Idaho and southern California; and Mr. M. L. Abts, south central Oregon. Specimens from the Nevada Test Site were obtained by the sen- ior author as a part of the Atomic Energy Commission Grant AT (11-1)-819. Other colleagues and students have aided in the field work and have provided data and sug- gestions for which we are most apprecia- tive. We are grateful to Drs. Robert C. Stebbins and David Wake for the privilege of examining the extensive collections at the University of California at Berkeley (MVZ), and to Dr. Alan E. Leviton for the opportu- Life Sciences Museum. Brigham Young Universit; 421 Santa Helena, Solana Beach. California 9207 Provo. I'tah 84602. 225 226 Great Basin Naturalist Vol. 37, No. 2 nitv to examine or receive on loan speci- mens from the California Academy of Sci- ences (CAS). We are also grateful for the privilege of studying specimens from the Field Museum of Natural History (FMNH), University of Michigan (UM), University of Kansas (KU), and the U.S. National Museum (USNM). Perhaps the first report dealing with the leopard lizards of the Great Basin was the notes of C. Hart Merriam (Stejneger 1893), which include observations of these lizards from southwestern Utah, southern Nevada, and east, central, and southern California. Since the report by Stejneger (1893), many articles have referred to the leopard lizards of the Great Basin. However, in none of these is there any change in the taxonomy, since all references refer to Crotaplujtus wislizeni. Aside from our previous reports (parts I and II) most other studies have included the leopard lizard as a part of a distribution list or as a part of ecological or natural history notes (Taylor 1912, Richardson 1915, Van Denburgh 1922, Knowlton et al. 1934, 1936, 1946, AUred, Beck, and Jorgensen 1963, Banta 1963, Jorgensen and Tanner 1963, Stebbins 1966, Fitch 1970). Mon- tanucci (1970) analyzed the relationship be- tween the leopard lizards of the central val- ley of California (C. w. silus) and those adjoining populations (C. w. wislizeni) in northern Los Angeles and eastern Kern counties, California. In this report on this species we will discuss and characterize those populations occurring in the Great Basin and attempt to indicate the zones of intergradation, as well as the presumed lanes of migration into the present area of distribution. Those populations in southern California (Imperial, Riverside, and San Bernardino counties) include individuals with varying expressions of the color pattern seen in C. w. wislizeni in Arizona and New Mexico. There is a strong tendency for the spots to be smaller and for the rings of white spots to be reduced or absent so that in some large females the pattern is greatly faded and the spots reduced in size and number. In adjacent northeastern Baja California the few specimens seen are similar and do not indicate any influence from C. w. copei. However, those examined from western So- nora (Kino Bay) have dorsal patterns as typ- ical for C. w. wislizeni as those from south central Arizona. In southern and eastern Nevada the trend is toward smaller spots. This also occurs in southwestern Utah, where many individuals are patterned similar to C. w. punctatus of eastern Utah. There is a reduction in the numbers of postmentals from southeastern Utah to southwestern Utah and adjoining eastern Nevada. This same pattern exists in much of east central Nevada and the Great Basin of western Utah. An explanation of the lanes of dispersion used in reaching this large area and the tax- onomic explanation of the population diver- sities are not clear. If we assume that the species had its origin in the desert plateaus of central or north central Mexico, then the following may be an initial attempt at an explanation of what we now see: at the close of the last thrust of the Pleistocene ice age the distribution of Crotaphytus wislizeni must have been restricted primarily to the drier, warmer areas of northern Mexico and perhaps some adjoining areas of the south- western United States. We have no evi- dence that the leopard lizard populations now extant were isolated as a result of the several ice flow advances or extensive plu- vial lake formations. Because of their gener- al morphological similarity and the zones of intergradation occurring between popu- lations, we consider the present distribution of this species to have been etablished since the Pleistocene. As the areas to the north and west warmed and dried, these lizards dispersed into the desert valleys formed between the mountain ranges south of the high plateaus in central Arizona and western New Mexico. The first range extensions into this area must have been into the lower valleys extending west- ward to the Colorado River and northward into the Rio Grande Valley. As the wanning trend continued, higher elevations and the more northern areas became occupied. Per- haps at this time the range was rapidly ex- tended up the Rio Grande Valley in New 1977 Tanner, Banta: Leopard Lizards 227 Mexico, and thus onto the plateaus of New Mexico, Arizona, western Colorado, and Utah. By entering such areas through low areas in the terrain, some populations were removed by distance and partially isolated fom the main body of the species. This re- duced gene flow and the new and different habitat resulted in adaptive changes in some of the fringe populations, which we now recognize as sufficiently distinct to warrant subspecies designation. Two basic color patterns exist east of the Colorado River and in northwestern Mexico. Those populations in the low valleys north and south of the U.S.-Mexican border are C. w. wislizeni, and those in the high plateaus (mostly upper Colorado River drainage north and east of Grand Canyon) are C. w. pitnctatus. The northeastern population of C. w. pitnctatus has reduced pigmentation on the dorsvim; small, widely dispersed spots; narrow, light cross bands; and an increase in the number of postmentals. If one travels south from the San Juan River of Utah into Arizona and New Mexico the characteristics are modified by an enlargement of the spots and a decrease in the number of post- mentals. The zone of intergradation be- tween C. It;, wislizeni and C. w. punctatus is wide, and no attempt is made to define its precise limits. The dorsal spots do in- crease in size from southeastern Utah across the plateaus south and east into Arizona and New Mexico. In spite of the steep can- yons and swift rivers in the Upper Colorado Basin, these leopard lizards have succeeded in crossing them and are established on the west side of the Colorado and Green Rivers in Utah and northern Arizona. From this position they have moved westward from the Colorado River through the gap be- tween the Kaibab Plateau to the south and Fig. 1. Possible routes of leopard lizard movement from the ancestral stock to their present distribution since the Pleistocene. 228 Great Basin Naturalist Vol. 37, No. 2 the Paunsaugunt Plateau to the north, to reach the Virgin River drainage in south- western Utah and southeastern Nevada. If the above hypotheses are correct, the Great Basin was invaded by leopard lizards originally derived from two populations: 1) Those moving westward through the desert valleys of southern Arizona which crossed the Colorado River, entered southern Cali- fornia, and extended their range south into Baja California Norte and west and north into the Great Basin. 2) Those moving west in southern Utah entered Nevada, where they also expanded south, west, and north into the Great Basin (Fig. 1). An examination of the populations now extant in southwestern Utah, southeastern Nevada, and southern California show a wide zone of intergradation between these two basic population types (C. w. wislizeni and C. w. punctatus). A large series from the Nevada Test Site (Mercury, Frenchman, Jackass, and Yucca valleys) contains exam- ples of both of the subspecies indicated above and many color patterns intermediate between them (Fig. 2). The populations in southern Nevada apparently represent a ma- jor area of intergradation. As one samples populations to the north and east into ex- treme eastern Nevada and the Great Basin of Utah, the characteristics of C. iv. punc- tatus are strongly evident but with an occa- sional representative of C. w. wislizeni pat- tern and individuals with intermediate characteristics. To the south particularly, and southwest of the Nevada Test Site, indi- viduals with C. w. wislizeni characteristics are seen more often. We are also impressed with the general differentiations in these populations brought about perhaps not only by introgression (hybridization) but also by the natural selection processes. Although most of the valleys in the Great Basin are interconnected either through low passes in the mountain chains or are connected at one or both ends of the discontinuous ranges, there are factors which bring about varying degrees of isolation. In some valleys the effects of isolation and thus some differ- entiating characteristics are more apparent. This is most obvious in the fringe areas such as the Lahontan and Humboldt valleys in northwestern Nevada, the adjoining areas in northeastern California, southern Oregon, I Fig. 2. Individuals showing color pattern variations. June 1977 Tanner, Banta: Leopard Lizards 229 and the Snake River Valley of southwestern Idaho. The population in Antelope Valley (Los Angeles County) also shows some dis- tinct color pattern characteristics which seem to represent some degree of inter- gradation with the C. w. silns population in the San Joaquin Valley. Montanucci (1970) contends that there is sufficient differentiation between silus and the adjoining populations to the south and east to justify full species rank for siltis. He found that the skull in the silus population was shorter than for other C. wislizeni pop- ulations, and the color pattern is distinct. Furthermore, few intergrades were found nor were there recognizable areas of over- lapping for the populations in question. We have examined a large series of speci- mens from the south end of the San Joaquin Valley as well as available specimens from along the Tehachapi Mountains from Mo- jave, both north and south. West from Lan- caster and Palmdale, Antelope Valley slowly rises in elevation and narrows as it ap- proaches Frazier Park (at about 3,500 feet) just north of Tejon Pass. On the basis of specimens seen from Antelope Valley, Fra- zier Park, Grapevine Canyon, and to the west and north of Wheeler Ridge, there is apparently a narrow route into the south end of the San Joaquin Valley. Furthermore, at Frazier Park we find intergrades (CAS 12787-8 and 1243). In northeast Kern Coun- ty a specimen (CAS 93427) from 1.3 mi W of Cane Wells (7 mi SE of Weldon) has a color pattern very similar to individuals from Washoe County, Nevada. There are other low passes along the Tehachapi Mountains through which entrance into the San Joaquin Valley may have occurred. We agree with Montanucci (1970) that intergrades are not common between C w. silus in the Central Valley and those popu- lations of wislizeni to the south and east. This does not suggest, however, that the two populations will not produce fertile off- spring in nature or (and) that the adaptive changes in the color pattern and length of the skull are entirely the result of a gene flow so reduced as to produce reproductive isolation. There is no question that the pop- ulation in the San Joaquin Valley, once es- tablished, received only a limited gene flow from the adjacent populations. The nature of the terrain was apparently the principal factor responsible for this reduction, al- though in recent years human activities south and east of Bakersfield, Kern County, have so altered the habitat that this lizard no longer occurs in wide areas. By elimi- nating leopard lizards fom the areas be- tween the Tehachapi Mountains and Bak- ersfield, that portion of the population in the San Joaquin Valley most likely to show intergrading characters was also eliminated. There appears to be only one possible area in which there might still be contac?^ be- tween the two populations. That area is from the west end of Antelope Valley, then along the foothills westward to the Pleito Hills and Wheeler Ridge. Even this contact may have been seriously disrupted or closed by recent highway construction. The entire area extending westward from southern Nevada serves as a potential route of dispersion toward the low passes from the Mojave Desert and into the southeastern end of the San Joaquin Valley of California. This route leads directly into Antelope Val- ley and the areas south and east of Bak- ersfield. Extensive agriculture in the south end of the valley has effectively established an artificial barrier between these popu- lations. All of this is, however, very recent, certainly within the last century. There is no reason to suspect that noticeable genetic changes in these populations have occurred in the time since man intervened. It should be noted that Montanucci (1970) reported some overlap of the skull character between silus and other popu- lations of wislizeni. Smith and Tanner (1974) found that the collared lizards in the Great Basin (C. collaris bicinctares) also was distinguishable from other C. collaris sub- species on the basis of cranial anatomy (ra- tio of width to length). Similarly, we must recognize the possibility that there is pro- portional variation in the length and width of the skull in the genus Crotaphytus. Fur- thermore, we question the validity of spe- cies/subspecies based on the estimated amount of gene flow between populations. If this is important (and we agree that it 230 Great Basin Naturalist Vol. 37, No. 2 plays a role in such deliberations), then what should be done with the numerous in- sular subspecies in Baja California where gene flow is nil or absent? We have weighed the pros and cons of species vs. subspecies in this case and can recognize strengths in both alternatives. The occurrence of recognizable intergrades re- ported by Montanucci (1970) and among the specimens we have seen (even though the intergrades in each study are few in number) is a strong deterrent to the recog- nition of silus as a species. Furthermore, what is to be gained by splitting? We are persuaded, as was Simpson (1945), that such splitting may tend to destroy or dim the true relationship between these populations rather than to establish the basic position of each to each other. We, therefore, retain the leopard lizards in the San Joaquin Val- ley of California as a subspecies of Crota- phiftus wislizeni until sufficient data become available to justify the elevation of silus to full species status. The populations in the Great Basin exhib- it considerable variation if one examines material from the Salton Sea Basin north into the Lahontan Basin of northwestern Nevada. There are, in fact, two populations with rather distinct color patterns. Those seen from the Coachella Valley south have a faded pattern. This is particularly true for adult females, in which some have few or no spots or bars. In males the pattern is more evident and often reflects traces of the pattern seen in Arizona specimens. Ob- viously a distinct pattern has not been es- tablished, and what we are seeing should be considered, for the present at least, as a de- veloping pattern resulting from adaptive change in the new habitat resulting from partial isolation, but with some influence from the populations to the north and east. The color pattern throughout a wide area in parts of northern and eastern Riverside County, most of San Bernardino County, eastern Inyo County, California; and in southern Nevada is extremely variable. Nearly all of the color patterns seen in this species have been observed in specimens from the Nevada Test Site, except for the typical silus pattern. This wide area of vari- ability is apparently maintained by contin- uous gene flow from the northeast (C. w. punctatus) and from the southeast (C. w. wislizeni). The phenotypes in Fig. 2 are in- dicative of a large and varied gene pool, which under selective environmental condi- tions has the potential to produce a wide variety of color patterns. In contrast to the south and central parts of the Great Basin, the north and western areas comprising the Lahontan Basin and the Snake River Basin from Ada County south and east to at least Bannock County, Idaho, have evolved a distinct color pattern. The most striking variation is in the size and shape of the spots. Because of the large distinct spots, we propose it be known as: Crotaphijtus wislizeni maculosus, subsp. Figs. 3, 4d HoLOTYPE.-An adult male, BYU 32685, taken approximately 200 m W of the look- out point along Nevada Highway 33, west side of Pyramid Lake, Washoe County, Ne- V'\)S^. ■?. I^orsal view of the holotvpe of Crotiiphytufi u. tiiarulosus. fiVU 32fi8o, taken on the west side of Pyramid Lake, 24 July 1970. June 1977 Tanner, Banta: Leopard Lizards 231 vada, bv Wilmer W. Tanner, on 24 July 1970. Paratypes.— Nevada: Washoe County: Topotypes (BYU 32686-7); Pyramid Lake, near south end (BYU 36405-8); Pyramid Lake (CAS 40507-9, 40516-26, 40533-5, and 44157). Idaho: Owyhee Co.: 17 mi SW Nampa, Hwy 78 (BYU 40676-7, 40773, 40815), 14 mi S. Bruneau (BYU 32706); Gooding County: 5.5 mi S. Gooding (BYU 32699-701), 5 mi N. Wendell (BYU 32697-8 and 32704-5); Ada Co.: Foothills N. Boise (SDHNM 1450-2, 23636 and CAS 45426-27, 53775-80). Diagnosis.— A subspecies of Crotaphytiis wislizeni most closely related to C. w. piinc- tatiis and C. w. wislizeni to the south and east of its distribution. Distinguished from C. IV. punctatiis and C. w. wislizeni in that the dorsal spots are greatly enlarged in size and often quandrangular rather than round. C. IV. maculosus is distinct from C. w. punctatiis in having fewer postmentals and greatly enlarged spots and from C. w. wisli- zeni in that there is not a circle of white dots around the larger dorsal spots. In con- trast to C. w. silus the enlarged dorsal spots are distinct on each side of the middorsal line and between the transverse bars. Indi- viduals have not been seen with the entire dorsal area between the bars pigmented as in most C. w. silus. Description of type.— An adult male, snout to vent 94 mm, total length 283 mm, tail total length ratio 1.49; dorsal scales (oc- cipital to base of tail) 187; ventrals 93, no- ticeably larger than laterals and dorsals; scales around middle of body 162, middorsal rows not noticeably enlarged, beadlike; supralabials 16-17; infralabials 16-18; femor- al pores 19-20, 2 or 3 small scales on poste- rior margin of pore; 20 scales between fe- moral pore series; head scales from rostral to occiput 22, smooth, occiput largest dorsal head scale, but with row of 8 enlarged scales from rostral posteriorly; postmentals 2-2; scales on anterior two-thirds of tail smooth, posterior scales weakly keeled. Head scales smooth, platelike, some raised medially, none imbricate; gulars enlarged near labials, smaller medially, and slightly longer than wide; gular fold prominent; scales posterior to fold enlarged and these in about 40 longitudinal rows. Head distinct, 22 mm long, 16.5 mm wide; rostral to ear 22 mm; body slightly flattened, wider than high; longest toe 20 mm. Color pattern consisting of a series of six rows of large dark brown spots extend- ing from head and shoulders to base of tail, middorsal rows and one lateral row extend- ing onto tail; a series of 9 spots in middor- sal rows from nape to base of tail, spots from 10 to 12 mm long, from 8 to 10 wide, and involving from 12 to 16 rows of dorsal scales; lateral spots smaller; transverse bars not distinct, confused with light areas sur- rounding the enlarged spots; gular region marked with several parallel dark, longitu- dinal stripes extending from labials to poste- rior edge of ear, a few spots between me- dian strip and first lateral strips; legs marked with large spots; body ground color cream to light gray. Range.— Lahontan Basin in western Ne- vada northwest into northeast California, north into southern Oregon, and northeast into the Snake River Valley of southwestern Idaho. Crotaphijtus wislizeni wislizeni Baird and Girard Specimens examined.— Arizona, Yuma Co., CAS 33486, 34204-8 and 33490; SDNHM 16730 and 17123; MMZ 71080(4) and 71080(2). California, Imperial Co., BYU 41177; LACM 4007, 15649, and 37787-8; LBSC 1471-2; MMZ 122632(2); SDNHM 1879, 7143, 7847, 10937, 11346, 13352, 13911, 18596, 20967, 28762, 36541, and 39735. Inyo Co., CAS 65117, 65318-20, 65336, 65381, 65486-8, 65529-31, 65583-4, and 65634; LACM 26804-6, 36668-9, and 52876; SDNHM 2503 and 12243. Kern Co., CAS 21234; LACM 4013, 26803, 53883-4, 63807, 63809-10, and 94709; MVZ 26026 and 70372; SDNHM 5877, 6047, 15877, 19483-5, 37438, 40005, and 41803. Los An- geles Co., LACM 4009, 15628-30, 15631, 15632-33, 15634, 15635, 15636, 15637, 15638, 15639, 15640, 15641-2, 15643, 15644, 15645, 15646-7, 15710, 26802, 52878, and 52879; MVZ 873 and 19652-4. Mono Co., SDNHM 28941. Riverside Co., 232 Great Basin Naturalist Vol. 37, No. 2 BYU 40941-3 and 40945; CAS 45369-70; LACM 4011, 15648, 22218-9, 26807-8, and 52880-82; SDNHM 4999-5000, 8040, 11344, 19486, 20700-3, 22352, 29448, 31428, 31906-7, 39747, 39850, and 41339. San Ber- nardino Co., BYU 3108 and 40942; CAS 43181-4; LACM 4012, 4014, 15653-61, 21646-49, and 52877; SDNHM 2489, 2498, 4408, 5875-6, 7798, 9013-5, 20887, 23636, 25344, 25641-2, 28939-40, 28942, 29660, 31453, 38889, and 41201. San Diego Co., LBSC 1473-4;LACM 15651-2; SDNHM 44, 8041, 9952, 11099, 11345, 18593-5, 22328, 23411, 24036, 29516, 33943, and 38418. Nevada, Clark Co., BYU 460, 504, 556, and 1541-2; LACM 15681-95; MMZ 84941(2), 90699-01; SDNHM 7626-7, 31718, 38675, and 40655-7. Elko Co., CAS 40933. Esmeralda Co., CAS 37801. Lander Co., BYU 2918; MMZ 84945 and 91868. Lincoln Co., BYU 9816; CAS 37014-18 and 38019- 23. Mineral Co., MMZ 84948 and 117360. Nye Co., BYU (Nevada Test Site including Mercury and Rock valleys and areas in and associated with Frenchman, Jackass, and Yucca Flats) 3037-51, 5084-7, 3093-98, 17276-9, 17308-22, 17944, 18969, 18984, 21740-55, 23620, 23966, 30083, 30085-7, 30121, 30539, 30546, 30589-90, 30595, 30608-9, 31425, 31810-14, 32571, 32632-4, 33571, 33609, 40127, and 41452-5; CAS 37017-8, 37512-3, 37688-92, 37996, and 38019-23; LACM 15696 and 26810; MMZ 84942(2), 34943(4), 34946, 84951, 84953-6, 84957(2), and 117336. White Pine Co., BYU 558. Utah, Beaver Co., BYU 580, 4302, 11349- 53, and 12001-2. Iron Co., BYU 478, 2371, 2374, 2376-7, and 2899; MMZ 59573, UU 1659. Juab Co., BYU 2731, 3020, 3027, 10244, 11295-6, and 12496; MMZ 91870(2). Millard Co., BYU 481, 580, 4308, 9099, 10179, 11354, and 14843-53; CAS 893-4, 1530-2, 38031, 85399-00, and 54151-2; MMZ 70553, 70554(6), 70555(14), 70556(2), 70557(2), 70558-60, 70561(2), 60562-3, 70564(3), 70565(4), 70566, 70567(5), 70568(2), 70569(3), 70570(3), 84950, and 91869; SDNHM 893-4, 1530-2, and 26399- 02. UU 3298-01. Salt Lake Co., UU 18. Se- vier Co., SDNHM 37960, 38315-6, 38244- 50, 38328-31, 38381-2, 39016-8, 39191, 39397-9, and 40571-3. Tooele Co., BYU 480, 1555, 14688, and 14843-53; MMZ 69432(2), 69433-4, 69435(4), 69436-7, and 91823; SDNHM 22998, 24988-9, and 26399- 402. Utah Co., BYU 483, 576, 1578, 1633, 2210-11, 8496, 12196, 14690-1, 16601, 21506, 22474, and 22835; UU 1-4, 2a, and 3563-7. Washington Co., BYU 515, 680, 1635, 8464, 11382, 23657, 31970-1, and 31972-8; CAS 54085-6; SDNHM 21078, 24993, and 25643. Mexico: Baja California Norte, BYU 23336; CAS 57446 and 90256; Sonora, BYU 3175; CAS-SU 17049-50; SDNHM 38251-4, 38605-6, 38888, and 40601; USNM 146455. Crotophytus wislizeni maciilosus subsp. nov. Specimens examined.— California, Modoc Co., CAS 63912; LACM 4010. Idaho, Ada Co., CAS 41366-71, 45417-25, and 53781-3. Boise Co., MMZ 54072, 68245(2), 68246-8, and 93084(4). Bingham Co., CAS 41275-6. Butte Co., BYU 30773. Canyon Co., MMZ 68249. Elmore Co., MMZ 60250; SDNHM 1452. Owyhee Co., BYU 2835; CAS 55261, 64151-4, and 64161-2; MMZ 68251(2), 122787(2), 122790, and 125694. Oregon, Harney Co., BYU 41445, 41454- 71; Malheur Co., MMZ 124618(2). Nevada, Churchill Co., MMZ 72653, 84944(2), and 84949; Humboldt Co., MMZ 43171-7, 60029, 77996, 85618(5), and 91865-7. Lyon Co., LACM 15696. Ormsby Co., SDNHM 38417. Pershing Co., MMZ 84947 and 91825(2). Washoe Co., CAS 38016-7, 40510-15, and 40527-32; MMZ 91824, 91826(2), 91827(2), and 91828-9. Crotaphytiis wislizeni silus Stejneger Specimens examined.— Fresno Co., CAS 22713, 22716-8, 22857-8, 23212-3, 23250, 23271, 23292, and 85416; King Co., MVZ 116442; Kern Co., CAS 3046, 39000, 43266, and 47996; LACM 4008, 15662-9, 15671-6, 15679-80, and 26809; San Luis Obispo Co., CAS 32195, 23200, and 23203-4; MVZ 2751, 8817, and 61077. MVZ 2725, 2733, 2735, 2737-43, 2747, 2749, 3775-81, 4891-2, 4894-8, 6851, 43142, and 72353-4. Specimens showing one or more inter- June 1977 Tanner, Banta: Leopard Lizards 233 grading characters with C. w. wislizeni: CAS 1243, 12787-8 (Frazier Park), 38999 (Tehachapi Mts.); LACM 15650 (Mt. Pinus), 15670 (25 mi S. Bakersfield), 15671 (Wheel- er Ridge), 15677 (McKittrick), 52879 (Black Butte), 15630-3 (Lovejoy Sp.), 15635 (Piute Butte): MVZ 43139 (3 ^ni SW Kicks Cor- ner). MoRPHOLOG"i Variati These attractive lizards of moderate size have only recently been critically examined. Since Smith (1946) noted the considerable variation in patterns, we have observed a correlation between color patterns and geo- graphic distribution. Tanner and Banta (1963) described and figured the pattern for C. t^\ wislizeni in Arizona and New Mexico and for C. w. punctatus in the upper Colo- rado River Basin. Montanucci (1970) dis- cussed in detail the color patterns of C. w. silus in the San Joaquin Valley of Califor- nia. Although these and other studies have made some comparisons with those popu- lations occurring in the Great Basin, a criti- cal examination of a large series has not been made. It is hoped that this study will yield the results necessary to provide some conclusions for the leopard lizards of this large and varied area. Color pattern types Spotting.— In the western United States and northern Mexico there are five distinct spotting patterns: 1. A pattern of moderately sized spots, with a circle of light spots surrounding the larger dark spots. This pattern is prominent Fig. 4. Dorsal views of (a) C. w. punctatus, (b) C. w. wislizeni, (c) C. wislizeni NTS intergrade, (d) C. w. macu- hsus. 234 Great Basin Naturalist Vol. 37, No. 2 on the dorsal aspect of the trunk, base of the tail, and hind legs. There may also be smaller spots interspersed between the larger ones and on the lateral parts of the trunk and tail. The white transverse bars may or may not be apparent; when present they are more obvious posteriorly on the trunk and base of tail (Fig. 4b). 2. A pattern of small spots, widely dis- persed on the trunk, tail, and legs. The background is lighter than in those with other patterns. Individuals with this pattern show no white dots in association with the small dark spots. Furthermore, there is a greater uniformity in the size of spots than in other patterns. The white bars are dis- tinct but narrow and less prominent than in populations occurring west of the Colorado River (Fig. 4a). 3. A pattern of moderate to large spots, with light centers in juveniles but not in adults. The light center is apparently lost by a gradual darkening and blending with the ground color. As the blending of colors oc- curs, the spots are subdivided to form from two to four closely associated or slightly connected spots. This pattern is character- istic of C. w. neseotes from Cedros Island and many individuals from Baja California, Mexico. 4. A pattern in which the areas between the white transverse bars are from a solid dark brown to a nearly black color. In juve- niles there are spots present, but these are lost in adults by a suffusion of dark pigment that obscures all spotting. The transverse white bars are distinct and much wider than in other patterns. This pattern occurs in the populations in the lower parts of the San Joaquin Valley. We note, however, that the spotted pattern of juvenile C. w. silus is very similar to that of adults from Antelope Valley (Fig. 5). 5. The subspecies C. w. macitlosus has a pattern of large dark spots between the transverse white bars and a lighter contin- uous cycle around the larger spots. This is reminiscent of the pattern in C. w. wislizeni except for larger spots and a light circle rather than a series of small light spots. The transverse white bars are moderate in size, being narrower than in C. w. sihis but wider than in C. w. pimctatus. In some in- dividuals the transverse bars become dif- fused and blended with the light circles, which produces an irregular netlike con- figuration of light markings (Fig. 4d). Montanucci (1970) referred to those light dots, circles, and other additions to the spotted pattern as adornments. This is seem- ingly an appropriate way to treat all addi- tions to the basic pattern of spots. Only a few populations are without adornments. Specimens of C. w. punctatus from south- eastern Utah have a light ground color and small dark brown spots (Tanner and Banta 1963). Those from near Bakersfield, Kern County, and north in Fresno County, Cali- fornia, have as adults reduced the pattern to a uniform dark color between the light transverse bars, thus eliminating the basic spot pattern. An examination of hatchlings and small juveniles demonstrates that C. wislizeni has Fig. 5. Dorsal view of specimens of C. w. silus (A-C; D and E, intergrades, Antelope Valley). LACM tags. June 1977 Tanner, Banta: Leopard Lizards 235 tliroughout its wide range maintained a spotted pattern in its young. This juvenile pattern is modified in adults of all color pattern tvpes. The similarity of hatchling patterns is unique, and we are impressed with this unifying character throughout the species. Adjacent populations with very dif- ferent adult patterns such as Antelope Val- ley (Los Angeles County), with its spotted pattern and the dark nonspotted pattern in the Bakersfield population (Kern County), have similar spotted patterns in hatchlings and young juveniles. Populations occurring between those populations with patterns as described above (1-5) show many variants. This is most evident in populations occur- ring in southern Nevada, in which a series from valleys such as Frenchman Flat or Mercury may have individuals with patterns similar to all other populations except that pattern found in the San Joaquin Valley (Fig. 2). In patterns 3 and 5 the white transverse bars may be absent on all or part of the trunk or so intertwined with the light areas between the spots as to lose their identity. Scale Patterns PosTMENTALS.— Only a few scale patterns vary noticeably within this species. As noted in Part I (Tanner and Banta 1963), the postmentals show significant variations between C. w. wislizeni and C. w. piinc- tatiis. We have reexamined this character in 101 specimens from the Upper Colorado River area of southeastern Utah and north- eastern Arizona, and our findings indicate an average postmental count of 5.71 in con- Table 1. Variation in the postmentals in populations occurring in northern Arizona and southeastern Utah. Area Number Range Mean Utah Populations Grand Coimty 34 4-7 5.7 Emery County 17 5-7 6.0 Garfield County 15 4-7 5.9 San Juan County 6 4-6 5.5 Northeast Arizona 6 4-7 5.8 Kane County 23 4-6 5.2 trast to the 5.72 reported by us in 1963 (Table 1). This figure includes a series from Kane County, Utah, which has a lower av- erage (5.17), and we consider Kane County to be a lane of di.spersal into southwestern Utah and eastern Nevada. If the specimens from Kane County are removed, the post- mentals for C. IV. pimctatus average 5.81. In Table 2 we have averaged the post- mentals for Washington County, and for populations in the Great Basin of west cen- tral Utah, Nevada, California, Oregon, and in the Snake River Valley of southwestern Idaho. These figures show an average de- crease in the numbers of postmentals from east to west. However, this decrease is ac- companied by an increase in the range of variation. We consider this a strong in- dicator of intergradation, particularly in those populations occurring in Washington County, Utah, the Great Basin of west cen- tral Utah, at the Nevada Test Site, and in southern Nevada. Those populations occurring along the northern limits of distribution (Table 2: Harney County, Oregon, and southwestern Idaho) show a noticeable increase in the number of postmentals when compared to other Great Basin populations. These same populations also show a decrease in the range of variation, with none of the Oregon Table 2. Variation in the postmentals in the Great Basin and adjacent areas in Utah, Nevada, Oregon, Idaho, and California. Area Number Range Mean Washington County, 15 4-5 4.5 Utah Great Basin Western Utah 57 2-6 4.1 Nevada Test Site 56 2-7 4.2 Lahontan Basin 29 2-6 4.1 Harney County, 9 4-6 5.0 Oregon San Bernardino, 77 2-6 4.2 Riverside County Inyo County 23 2-6 4.4 Imperial County 21 3-6 4.4 Antelope Valley, 31 2-6 4.4 Los Angeles County Snake River Valley Southwest Idaho 34 .3-6 4.6 101 5.71 352 236 Great Basin Naturalist Vol. 37, No. 2 specimens with less than four and only one Idaho specimen with three postmentals. Al- though our samples are small for some pop- ulations (Oregon), they show consistency ex- cept for Washington County, Utah, which we include with the large and varied inter- grading populations occurring in south- western Utah and much of central and southern Nevada. The increase in the num- ber of postmentals for the northern popu- lations of the subspecies maculosiis, and for that matter punctotus, is not explainable ex- cept as a response to the climatic changes in the northern localities. An examination of the scale pattern sum- maries in Table 3 indicates that there are only minor variations. A few trends are ap- parent, such as a higher range and average for femoral pores and the scale rows around the body in the populations of Baja Califor- nia Sur, Mexico. In neither of these patterns does the lower limits of the range show as few pores or rows of scales as in other pop- ulations. Only in the higher averages, par- ticularly in the scale rows, are the varia- tions between the Baja California Sur and the continental populations obvious. In all populations and subspecies there is an over- lapping in the ranges of all scale patterns examined. In most patterns this overlapping is extensive. The Nevada Test Site population shows greater variation than other adjoining popu- lations in the number of scale rows, ven- trals, and postmentals. In the scale rows the average is low, with only C. w. punctatus lower. The ventrals average the lowest of all populations studied, and the range for the postmentals from two to seven is great- er than in other populations. Skull Measurements Skull length/width ratio.— The ratio of the skull width into its length (Fig. 6) substantiates the data presented by Mon- tanucci (1970) that the skull is shorter in the San Joaquin population {silus) than oth- er subspecies and populations. There is, however, an overlapping with other popu- lations of approximately 25 to 50 percent when compared with other populations ex- amined (Fig. 6). Discussion On the basis of color and color pattern, we must assume that C wislizeni expanded San Joaquin Valley 1.23 1.42 Reno, Pyramid Lake Nevada Test Site Emery-Grand Co. Ut. Western, Utah 1.31 l.,54 1.35 1.59 1.37 1.59 1.36 1- 1.55 1.20 1.25 1.30 1.35 1.40 1.45 1.50 1.55 1.60 Fig. 6. Ratio of width into skull length (from rostral to median posterior of skull). Specimens used and data available on request. 1977 Tanner, Banta: Leopard Lizards 237 o 9- So S CO ic ?^ § S - 2 5 5 •2 .S ? X •= -73 Z U c« D CQ W Z H 238 Great Basin Naturalist Vol. 37, No. 2 its ransji;e rapidly at the close of the Pleisto- cene and occupied the low valleys in south- ern New Mexico, southern Arizona, north- ern Sonora, and southern California, perhaps before entering the areas of the high plateaus. Such an hypothesis would have much of southern California, southern Nevada, and the low valleys along the Cali- fornia-Nevada line occupied before the in- vasion of C. IV. ptinctatus occurred in west- ern Utah and east central Nevada. This hypothesis is based on the fact that color patterns in southern ('alifornia, the Ante- lope and San Joaquin valleys of California, and the populations in northern Nevada (nuinilostis) do not seem to be in any way similar or related to the punctatits pattern. The invasion of western Utah and south- eastern Nevada by pimrtatus seems to have occurred more recently. This is based on the fact that there is a real jimibling of pat- tern in southern Nevada (NTS); and from this general area a diminishing effect is seen as one examines specimens from the south- west and north. To the northeast, that is from southern Nevada to Washington Coun- ty, Utah, the influence of punctdtus increas- es. If populations of wislizcni had occupied these areas before punctatits arrived, they have or are in the process of a continuing and extensive introgression with a strong punctatus influence. This is most obvious throughout southern and extreme eastern Nevada. There is reason to suspect that the main- How of leopard lizard migration at the clo.se of the Pleistocene was from southern Cali- fornia, through the low valleys of eastern California north into the Lahontan Basin of west central Nevada. As the higher valleys of western Utah and eastern Nevada be- came environmentally available, they were occupied. Furthermore, with improving cli- matic conditions, the terrain (with its north- south-oriented mountains) is well suited for a rapid movement to the north (or south) through the broad alluvial valleys. Migration into the Bonneville Basin of western Utah and the high valleys of cen- tral and eastern Nevada was slower. Eleva- tion barriers ranging up to 5,000 feet may have remained cool and were probably most responsible for slowing the northward movement. This is still effective for some species such as Sceloporus ma^ster and Cal- lisauru.s centralis, which are not in the Bon- neville Basin but have reached the Lahon- tan Basin. The early arrival and partial isolation in the Lahontan Basin provided a greater opportunity for differentiation and also time to extend their ranges into south- ern Oregon, northeastern California, and southwestern Idaho. Although those popu- lations now in the Bonneville Basin are par- tially isolated, they appear to have been de- rived too soon to have differentiated as much as those occupying the Lahontan Ba- sin. In western Utah, southern Nevada, and adjoining California (San Bernardino and Inyo counties) there is a large area of inter- gradation between those populations exist- ing in the area or those that moved north from southern California (C. w. wislizeni) and those moving into the area from south- eastern Utah and northern Arizona (C. w. punctatus). Because of the size of the area of intergradation, one may assume that the gene flow from these populations has been strong and steady. As one samples popu- lations radiating from this central area, the populations are more homogeneous. The one area of intergradation most difficult to interpret is the populations in the Bonne- ville Basin of western Utah and eastern Ne- vada. Within this basin (from Iron County to Box Elder County, Utah) the populations are more homologous than those from the Nevada Test Site, but they still provide an occasional individual with either a wislizcni or a punctatus pattern. Apparently, introgression had occurred before the Bonneville Basin was occupied, or there was a mingling of the two as they moved northward at about the same time. Once movement was slowed and population stability was achieved, differentiation oc- curred. This is seen in such populations as those in the Lahontan Basin and San Joa- quin Valley, and to an extent in Antelope Valley (Los Angeles County) and the Coa- chella Valley. Differentiation has occurred primarily on the fringes of distribution, with the large central area in southwestern Utah, June 1977 Tanner, Banta: Leopard Lizards 239 southern Nevada, and much of southeastern CaHfornia occupied by a population with variable degrees of intergrading character- istics. Perhaps the most difficult populations to understand occur in the areas of western San Bernardino County (west of Barstow), eastern Kern County (near Mojave), and northern Los Angeles County (Palmdale- Lancaster area). Populations in these areas are far enough removed from the inter- grading population of southern Nevada to reflect some pattern characteristics of their own. This is particularly true for Antelope Valley in northern Los Angeles County, where a darker ground color obtains and the spots are darker and more uniform in size and color. Although our samples are small, those seen reflect a slow but steady change from C. w. wislizeni to C. w. silus from western San Bernardino County through Antelope Valley to the areas near Fraser Park and on westward into the low hills and valleys of northern Ventura and Santa Barbara counties. The data concerning C. w. sihis presented by Montanucci (1970) agrees generally with our findings. Any differences are in the area of interpretation. We see in adjoining popu- lations (Antelope Vallev, Los Angeles Coun- ty, and those near Weldon, San Bernardino County) strong influences of the adult and juvenile color patterns of C. w. silus. We also recognize a more sudden change in pattern from the foothills surrounding the San Joaquin Valley to the valley floor. In other areas of the extensive leopard lizard range ecological changes are more gradual, or they occur in smaller geograph- ical areas which do not seemingly provide sufficient space for isolation and thus slowly induce genetic adaptive change. This gradu- al change in the environment has provided for wide areas of introgression in much of the Great Basin and is perhaps unique in that the central area of distribution (south- ern and eastern Nevada, southern Califor- nia, and western Utah) represents a large area of genetic variation as deduced from the phenotype. As indicated above, within this wide area of diversity subspecific pat- terns (except for silus, copei, and neseotes) appear in some individuals. Within the pop- ulations considerable individual variation occurs and is so extensive in some areas that few lizards show the same phenotypic pat- tern. Only in the fringe areas where some degree of isolation obtains do we find sub- species or incipient subspecies. Literature Cited Allred, D. M., D. E. Beck, and C. D. Jorgensen. 1963. Biotic communities of the Nevada Test Site. Brigham Young Univ. Sci. Bull. Biol. Ser. 2(2): 1-52. B.\NTA, B. H. 196.3. Preliminary remarks upon the zoogeography of the lizards inhabiting the Great Basin of the western United States. Wasmann Jour. Biol. 20(2): 25.3-287. Banta, B. H., and W. W. Tanner. 1968. The system- atics of Crotaphytiis wislizenii, the leopard lizards (Sauria: Iguanidae). Part II. A review of the stat- us of the Baja California peninsular populations and description of a new subspecies from Cedros Island. Great Basin Nat. 28:183-194. Fitch, H. S. 1970. Reproductive cycles in lizards and snakes. Univ. Kansas Mus. Nat. Hist. Misc. Pub. 42:247, p. 16 fig. Jorgensen, C. D., and W. W. Tanner. 1963. The ap- plication of the density probability fimction to determine the home range of Uta stansburiana stansburiana and Cnemidophoriis tigris tigris. Her- petologica 19(2): 105-1 15. Knowlton, G. F., and W. L. Thomas. 1934. Notes on some insectivorous Utah lizards. Proc. Utah Acad. Sci. Arts, Lett. 11:257-259. 1936. Food habits of Skull Valley lizards. Copeia 19.36:64. Knowlton, G. F., D. R. Maddock, and S. L. Wood. 1946. Insect food of the sagebrush swift. J. Econ. Ent. 39:382-.383. Montanucci, R. R. 1970. Analysis of hybridization be- tween Crotaplit/ttis uislizenii and Crotaphytus .vi/i/s (Sauria: Iguanidae) in California. Copeia 1:104-12,3. Richardson, C. H. Jr. 1915. Reptiles of northwestern Nevada and adjacent territory. Proc. U.S. Nat. Mus. 48:403-4.35. Stebbins, R. C. 1966. .\ field guide to western reptiles and amphibians. Houghton Mifflin Co., Boston. Stejneger, L. 1890. Annotated list of reptiles and bat- rachians collected by Dr. C. Hart Merriam and Vernon C. Bailey on the San Francisco Mountain plateau and desert of the Little Colorado, Ari- zona, with descriptions of new species. N. Am. Fauna 3:103-118. Smith, H. M. 1946. Handbook of lizards. Comstock Publ. Co., Ithaca, N.Y. Tanner, W. W., and B. H. Banta. 1963. The system- atics of Crotaphytus wislizenii, the leopard liz- ards. Part I. A redescription of Crotaphytus uisli- zenii, wislizenii Baird and Girard, and a description of a new subspecies from the Upper 240 Great Basin Naturalist Vol. 37, No. 2 Colorado River Basin. Great Basin Nat. 23:129- features of that region. Univ. Calif. Publ. Zool. 7:319-436. TAVLor'w. P. 1912. Field notes on amphibians, rep- Van Denburgh, J. 1922. The reptiles of western North tiles and birds of northern Humboldt County, America. Occ. Pap. Calif. Acad. Sci. 10:1U2». 148. ,OR, tiles, Nevada, with a discussion of some of the faunal SYZYGIOXYLON CHHINDWARENSE, A NEW FOSSIL WOOD FROM THE DECCAN INTERTRAPPEAN SERIES OF INDIA E. M. V. Nambudiri' and William D. Tidwell' Abstract.— A petrified wood related to genera of the family Myrtaceae was collected from the Deccan Inter- trappean Series of India. On comparison with fossil woods of this family, it most closely resembles Syzygioxylon Ingle (1973). The specimen differs from described species of Syzygioxylon and, therefore, it is proposed as the new species Syzygioxylon chhindwarense. Fossil wood resembling species of Eu- genia (Myrtaceae) has been collected from a black chert in the Deccan Intertrappean beds near the Mohgaon Kalan locality in Chhindwara District, Madliya Pradesh, In- dia. Myrtaceous woods have been described previously from these formations by Ingle (1973) and Patil and Singh (1974). Prakash (1965) reviewed reports on dicotyledonous woods from India and the Far East which included those from this locaHty. Since then, additional fossil woods from the Moh- gaon Kalan locality have been described by Chitaley and Patel (1971), Chitaley and Pa- til (1972), Chitaley and Kate (1972), and Nambudiri and Tidwell (1975). Description The specimen is 37 mm in diameter and consists of secondary xylem. The xylem is diffuse porous, and two growth rings are faintly visible to the unaided eye. Vessels are numerous, small, and faintly visible to the unaided eye, although they are distinct under a hand lens. They are round to oval, with a tangential diameter of 77.5 jum to 116 jum and a radial diameter of 145 jUm to 164 jum. They are mostly solitary, occasion- ally in pairs and rarely in radial multiples of 2 to 4 vessels (Figs, la, 2a). Distribution of these vessels is 85 per sq cm. They have simple, oblique perforation plates (Fig. lb). The vessel wall is 2.5 jum in thickness. In- tervascular pit pairs are round to oval, al- ternate, and vestured (Fig. Id). They have a mean diameter of 4.2 jitm. Parenchyma is paratracheal, either scanty or vasicentric (Fig. 2b). Generally, they form one or two layers around the vessels. These are 41 jxm long and 7 ju,m broad. Vessel to parenchyma pits were not ob- served. Rays (Fig. Ic) are extremely fine to mod- erately fine, faintly visible to the unaided eye. They are heterogeneous and uniseriate as well as 2-4 seriate. Some of the multi- seriate rays are fused by their tips. Unise- riate rays range from 68 jum to 240 /xm long. Multiseriate rays are from 435 jum to 659 jixm in length. The former has a mean width of 9 jLim, whereas the latter are 30 jLim in width. Vessel to ray pits (crossfield pitting) are similar to intervascular pit pairs (Fig. 2d). Fibres are libriform and nonlibriform and thin walled. They are 388 jum long and 14 /xm broad. Fibres are polygonal in cross sec- tion and are septate as well as nonseptate (Fig. 2c). Discussion The fossil wood of Syzygioxylon chliindwarense is similar to members of the Ochnaceae, Sonneratiaceae, Vochysiaceae, Punicaceae, Dipterocarpaceae, and Myr- taceae families. Wood of Ochnaceae plants with very scanty parenchyma differ from the occasional occurrence of the vasicentric Department of Botany and Range Science, Young University, Provo, Utah 1 241 242 Great Basin Naturalist Vol. 37, No. 2 type in the present wood. The ahform con- fluent parenchyma in genera of the Vochy- siaceae separates the xylein of this family from S. chhindivarense. The xylem of Son- neratiaceae has exclusively uniseriate rays. Xvlem of punicaceous genera lacks paren- chyma, whereas woods of Dipterocarpaceae contains apotracheal parenchyma rather than the paratracheal type of S. chhindwa- rense. Thus our specimen differs from these families. However, it is similar to several taxa of the Myrtaceae. The paratracheal scanty or vasicentric parenchyma, as ob- served in the present wood, is contained in the myrtaceous genera Eucalyptus, Goni- desia, Xanthostemon, and Eugenia. In con- trast, vessel multiples are fewer in S. chhindivarense than in Eucalyptus. Rays of Gonidesia and Xanthostemon differ from those of our specimen. They are up to 8 cells wide in Gonidesia and from 4 to 6 cells wide in Xanthostemon. Eugenia is, therefore, the closest extant genus to S. chhindivarense. Vessel arrangement, nature of the inter- vascular pit pairs, and type of rays in Eu- genia correspond to that of the present wood. Eugenia Linn, is one of the largest genera of tropical plants, with about 1000 species— of which some 150 are represented in the lowlands and mountains of the Ma- laysian Peninsula (Desch 1954) and four are commercial timbers of India (Pearson and Brown 1932). The common name for this genus is Kelat in Malaysia and Makaasin in the Philippines. Species of Eugenia range in size from dwarfed and shrubby treelets of the mountain tops to the medium-sized or quite large trees in the lowland forests. The anatomical structure of the wood of Eu- genia is similar, but there is an appreciable variation in the amount and, to some de- gree, the distribution of wood parenchyma in different species. Some species of Eu- genia have apotracheal parenchyma, where- as in others it is aliform confluent. In Eu- genia nitidula Ridl., as in other species of this genus, the parenchyma is paratracheal vasicentric and only occasionally confluent. Thus no one species is entirely like our specimen. Other similar species are E. long- iflora (Presl.) Vill., E. polyantha Wight, E. rhamphiphylla Craib, and E. tetraptera var. pseudotetraptera Herd. S. chhindivarense varies from the above mainly in its parenchyma distribution. It lacks either the confluent or aliform paren- chyma that is present in the others. How- ever, it is similar in having more distinct growth rings, which also occur in £. cumingiana Vidal and E. rhamphiphylla. This feature stands out because one charac- teristic of the genus is the general lack or inconspicuous nature of the growth rings. Two features that are often present in some species of Eugenia but lacking in our fossil form are frequent tyloses and traumatic gum ducts, the latter occurring horizontally in the rays of some species of Eugenia. The present wood demonstrates closest affinities to described species of the fossil genus Syzygioxylon Ingle. However, there are sufficient differences between them and our specimen to propose ours as the new species, Syzygioxylon chhindivarense. Ingle (1973) described Syzygioxylon mandlaense from the Mandla District in Madhya Pradesh, India. This species shows both paratracheal and metatracheal paren- chyma, a condition differentiating it from the present specimen. In proposing Syzy- gioxylon as a new genus. Ingle (1973) dem- onstrated similarities between the living genus Syzygium and his taxon. However, many species of Syzygium with which he compared his specimen have been placed in Eugenia by Pearson and Brown (1932). From the Mohgaon Kalan locality, Patil and Singh (1974) reported Syzygioxylon mohgaoense. This species has metatracheal parenchyma along with the paratracheal vasicentric type. In S. mohgaoense the vasi- centric parenchyma becomes aliform. Kra- mer (1974) described a myrtaceous wood from the Tertiary beds of Java under the name Syzygioxylon bataviae. However, he Fig. 1. Syzygioxylon chhindivarense sp. nov.: a) Transverse section illustrating vessel distribution (X45). b) Ves- sels with simple perforation plates (X50). c) Tangential section showing narrow rays (XllO). d) Vessel exhibiting alternating, vestured intervascular pitting (XQOO). June 1977 Nambudiri, Tidwell: Fossil Wood 243 244 Great Basin Naturalist Vol. 37, No. 2 was not aware of the institution of the genus Syzygioxylon by Ingle (1973) and treated Syzijgioxylon as a new genus in his paper (Kramer 1974). Syzygioxylon bataviae has paratracheal aliform parenchyma. The short lateral extensions of the aliform paren- chvnia in this species unite and form dis- continuous confluent strands. Regardless of these differences, S. el iJiindwa reuse shows resemblances to S. mandlaense Ingle (1973) and S. bataviae Kramer (1974). The similarities are in the arrangement of vessels and in the septate and nonseptate fibres. However, the wood is closer to S. mohgaoense Patil and Singh (1974) than to the other two species. The characteristics common to S. mohgaoense and S. chhindwarense are the nature and ar- rangement of vessels, some united rays, and the septate or nonseptate fibres. However, they differ from each other in the total ab- sence of metatracheal diffuse parenchyma and the presence of vestured pits in the present specimen. Shallom (1960) described a similar fossil wood, Baningtonioxylon dec- canense, from the Intertrappean beds of Mahurzari. She assigned it to the family Le- cythidaceae, whereas Chowdhury (1965) considered it to be a member of the Myr- taceae. However, Chowdhury (1965) sugges- ted that B. deccanense be reinvestigated. Myrtaceae is probably the oldest repre- sentative of Myrtales (Muller 1970) and pos- sibly extended back into Cenomanian time (Penny 1969). From the Eocene Greeir Riv- er Formation of northwestern Colorado and northeastern Utah, MacGinite (1969) report- ed a leaf impression Eugenia americana. E. arenaceafonnis was described earlier from the Oligocene Florissant flora of Colorado (MacGinite 1953). Myrtaceous fossils are also known from Eocene of Europe (Krutzsch 1967) and Cretaceous of Argen- tina (Menendez 1972). The family is esti- mated to contain over 70 genera and about 2,800 species of trees and shrubs, widely distributed throughout the tropical and sub- tropical regions of the world. Members of this family occur especially in the hot zones of low rainfall bordering on the tropics rather than in the rain forest belt (Desch 1954). Raven and Axelrod (1974) noted that they are abundant in tropical Asia, South America, and Australia. Several researchers (Dadswell and Ingle 1947, Pike 1956, Chattaway 1959, Schmid 1972) have studied the taxonomic problem of Eugenia and Syzygium. They, generally, followed Merril and Perry (1938) in sugges- ting that genus Eugenia be restricted to species of the New World and Syzygium be utilized for the majority of species of this taxon in the Old World. In comparing the fossil species of Syzy- gioxylon with the criteria used by Dadswell and Ingle (1947) for separating Eugenia and Syzygium, two species (S. chliindwarense and S. mandlaense) have more characters in common with Eugenia than Syzygium. This is interesting considering that their collec- tion localities are in the Old World and not the New. This paradox can be partially ex- plained by speculating upon the origin of Eugenia and Syzygium. The origin of these two genera may have been from a common ancestor such as Syzygioxylon. Diagnosis of Species Syzygioxylon chhindwarense, no v. sp., wood diffuse porous, two growth rings faintly visible; vessels numerous; solitary, in pairs or in radial multiples of 2 to 4; tan- genial diameter 77.5 jum to 116 jum radial diameter, 145 jum to 164 jum, vessel wall 2.5 /xm thick; perforation simple, plates oblique- ly placed; intervascular pit pairs round to oval, alternate, 4.2 jum in diameter, ves- tured; parenchyma paratracheal, vasicentric or scanty; rays extremely to moderately fine, uniseriate rays 68 /im to 240 jum long and 9 jum broad, multiseriate rays 435 fim to 659 jtim in length and 30 jxm in width; vessel ray pits similar to intervascular pit pairs; fibres libriform and nonlibriform, mostly aseptate but few septate, 388 /im long and 14 jum broad, polygonal in cross view. HoLOTYPE.-MOG 69/N (To be deposited at the Birbal Sahni Institute of Paleobo- tany, Lucknow, India.) Locality.— Mohgaon Kalan Horizon.— Deccan Intertrappean Series Age.— Early Eocene June 1977 Nambudiri, Tidwell: Fossil Wood 245 Fig. 2. Syzygioxylon chhindwarense sp. nov.: a) Close up of radial multiple of two vessels (X145). b) Vessel in transverse section with vasicentric parenchyma (X340). c) Tangential section showing nonseptate fibres (X450). d) Vessel to ray pits (crossfield pitting) on vessel. Note their similarity to the intervascular pitting (X620) (See Fig. Id). 246 Great Basin Naturalist Vol. 37, No. 2 Literature Cited Chattaway, M. M. 1959. The Anatomy of Bark. VII .species of Eugenia (sens. lat.). Trop. Woods ,3:1- 14. Chitaley. S. D., and M. Z. P.^tel. 1971. A fossil dico- tyledonous stem from the Deccan Intertrappean cherts of Mohgaon Kalan, India. Jour. Biol. Sci. 14:50-57. Chitaley, S. D., and U. R. Kate. 1972. A petrified Rhamnaceous wood from the Deccan Inter- trappean Beds of Mohgaon kalan. Botanique 3:41- 44. Chitaley, S. D., and G. V. Patil. 1972. Ebenaceous fossil wood infected with Deuteromycetous fun- gus from the Deccan Intertrappean Beds of India. Botanique 3:99-106. Chowdhury, K. a. 1965. The Tertiary flora of India and probable disposition of continents. Paiaeobot- anist 14:172-184. Dadswell, H. E., and H. D. Ingle. 1947. The wood anatomy of the Myrtaceae, I. A note on the gen- era Eugenia, Syzijgium, Acmena, and Cleistocalyx. Trop. Woods 90:1-7. Desch, H. E. 1954. Manual of Malayan Timbers 2. Malayan Forest Records, no. 15. Ingle, S. R. 1973. Syzygioxylon mandlaense gen. et. sp. nov., a fossil dicotyledonous wood from Mandla District of Madhva Pradesh, India. Botanique 4:71-76. Kramer, K. 1974. Die tertiaren Holzer Siidost-Asians (Unter Ausschluss der Dipterocarpaceae) I. Teil. Paleontographica. 144 B, 45-181. Khutzsch, W. 1967. Der Florenusechsel im Alttertiar Mitteleuropa.s auf Gnmd von Sporenpalaontolo- gischen Untersuchungen. Ahh. Zentr. Geol. Inst. 10:17-37. MacGinite, H. D. 1953. Fossil plants of the Florissant Beds, Colorado. Carnegie Inst. Wash. Publ. 599, 153 p. 1969. The Eocene Green River Flora of north- western Colorado and northeastern Utah. Univ. Calif. Publ. Geol. 83:1-140. Menendez, C. a. 1972. Estudios paleobotanicos en la Argentina, avances, problemas y perspectivas. Mem. Svmp. I Congr. Latino-Amer. Bot., p. 61- 97. Merrill, E. D., and L. M. Perry. 1938. On the In- dochinese species of Syzygiitm Gaertner. Jour. Ar- nold Arboretum 19:99-116. MuLLER, J. 1970. Palynological evidence on early dif- ferentiation of angiosperms. Bio. Rev. 45:417-450. Nambudiri, E. M. v., and W. D. Tidwell. 1975. Elaeocarpus chitaleyi, sp. nov., from the Deccan Intertrappean Beds of India. Brigham Young Univ. Geol. Stud. 22:29-37. P,\til, G. v., and R. B. Singh. 1974. An infected stem from the Deccan Intertrappean Beds of Mohgaon Kalan. Botanique 5:141-145. Pike, K. M. 1956. Pollen morphology of Myrtaceae from the Southwest Pacific area. Austral. Jour. Bot. 4:13-53. Pearson, R. S., and H. P. Brown. 1932. Commercial timbers of India. 1 & 2. Calcutta. Penny, J. S. 1969. Late Cretaceous and Early Tertiary Palynology, p. 331-376. In R. H. Tschudy and R. A. Scott (editors). Aspects of Palynology. Wiley- Interscience, N.Y. Prakash, U. 1965. A survey of the fossil dicotyle- donous woods from India and the Far East. Jour. Paleontol. ,39:815-827. Raven, P. H., and D. I. Axelrod. 1974. Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Card. 61:539-673. ScHMiD, R. 1972. A resolution of the Ettgenia-Syzygium controversy (Myrtaceae). Amer. Jour. Bot. 59:423- 436. Shallom, L. J. 1960. Fossil dicotyledonous wood of Lecythidaceae from the Deccan Intertrappean Beds of Mahurzari. Jour. Indian Bot. Soc. 39:197- 203. i BIRDS OF GUNNISON AND CUB ISLANDS GREAT SALT LAKE, UTAH F. L. Knopf and V. C. Bachman' Abstract.— a total of 87 avian species were recorded on islands of Great Salt Lake by Behle, with 17 ob- served on Gunnison and Cub islands. This paper presents sightings and salvaged carcasses recorded on Gunnison and Cub islands, 1972-1974. Of the 112 species observed, 95 are first records for Gunnison and Cub islands and 49 are first records on or near any island of Great Salt Lake. Over 90 percent of species observed on the islands were transients. Behle (1958) reported a total of 87 avian species occurring on the islands of Great Salt Lake, Utah. Those records resulted from collecting expeditions to larger islands, plus field observations of species during brief visits to smaller islands. This paper re- ports observations of species recorded dur- ing prolonged stays on two of the smaller islands, 1972-1974. Nine islands occur within Great Salt Lake (for map, see Marshall 1940). Gunnison and Cub islands lie about 25 m apart in the northwest arm of the lake, approximately 14 km north of Lakeside, Utah, 40 km from Promontory Point and 13 km from the west shoreline. Water depths around the islands average 3 m, and these plus the distance to the western shoreline vary with annual fluc- tuations in water levels of the lake (Knopf 1974). The two islands become a single land mass during years when the lake level drops approximately one meter below the "nor- mal" of 1,280 m above sea level. Gunnison Island is about 1.6 km long by 0.8 km wide with its long axis extending in a north-south direcion. This island has an ir- regular shoreline of 4.5 km with four small bay areas, rises 85 m above mean lake level, and includes 66 ha. The substrate is loose soil with protruding rocks and cliffs. The shorelines are rock except for sandy areas along the bays. Vegetation is typical of the Great Basin desert, with June Grass {Bromiis tectorwn) predominating. Opiintia fragilis, Salsola kali, and Bassia hyssopifolia are also widely dispersed and common. Sandy areas along the shoreline support pure or mixed stands of Sarcohatus vermiculatus, Atriplex sp., and Chnjsothamnus nauseosus, with the latter also occurring on north- and east-fac- ing slopes at higher elevations. Cub Island lies just north and east of Gunnison, is comma shaped, and comprises 3.2 ha. The physiography, substrate, and ve- getation of Cub are similar to Gunnison, al- though larger vegetative forms are less com- mon. Sarcohatus is absent. This latter species was used by many transient birds (Tyrannidae, Sylviidae) on Gunnison Island, birds not seen elsewhere on the islands. The nearest fresh water is at Locomotive Springs, about 40 km north of the islands. Sightings of birds were recorded between 15 April and 4 August 1972; 6 April and 2 August 1973; and 5 April and 25 July 1974. We spent 11 days on the islands in 1972, and over 70 days each in 1973 and 1974. We surveyed the islands for birds at least one hour daily and also recorded all sight- ings of species observed at other times. Each sighting reported herein was verified at the time of the observation by at least one other person. The specific area sur- veyed included the islands plus the lake sur- face within 25 m of shoreline. List of species Behle (1958) lists 17 species of birds re- ported on Gunnison Island, with no separate 'Sthoul uf Biolo(iKal Sciences. Oklahoma State University. Stillwater, Oklahoma 74074. ^285 North 2(X) East, Price. Utah 84,501. 247 248 Great Bsin Naturalist Vol. 37, No. 2 account for Cub Island. We recorded 15 of those species from 1972 to 1974 also. The following list (a preliminary listing was re- ported in Rawley 1976) includes the dates of sight records for 95 additional species (denoted by " or °°), giving a total of 112 species recorded on Gunnison and Cub is- lands. In addition, 47 of the species (de- noted by ''° only) observed in these efforts represent new records for any island in Great Salt Lake, bringing the total to 134 species occurring on all islands. Carcasses of Great Egret {Casmerodiiis albiis) and An- cient Murrelet (SyntliUboramphus antiquus) found on Gunnison Island may have washed ashore after death and have not been in- cluded in the Gunnison /Cub Island list. They do represent species recorded for the first time for Great Salt Lake, bringing the lake total to 136, (49 new records in this paper). For information on the murrelet, see Knopf (1976). °Podiceps nigricoUis. Eared Grebe. 8 June 1973, flock of 25 birds 29-30 April 1974. Abundant on lake in migration. ° ° Aechmophorus occidentalis. Western Grebe. 14 April 1974. Common on lake in migration. Pelecanus enjthrorhijnchos. White Pelican. Nesting and abundant (Behle 1958), with 5,000 to 5,200 birds estimated for each season, 1972-1974 (Knopf 1975). " Phalacrocorax auritus. Double-crested Cor- morant. 16 April 1972, 15 April 1974. Ardea herodias. Great Blue Heron. Formally nesting on Gunnison Island (Behle 1958). One pair nested on Cub Island each season 1972-1974. 'Botaurus lentiginosus. American Bittern. 14 May 1973. "Plegadis cliihi. White-faced Ibis. Flock of 3 birds 16 May 1973. °Branta canadensis. Canada Goose. 24 April 1972, 14 April 1974, 6 May 1974. "Anas acuta. Pintail. Flock of 11 birds 29 May 1973. "Anas crecca. American Green-winged Teal. 15 April 1972, 6 April 1973. ""Anas discors. Blue-winged Teal. 14 Mav 1973. '"Anas cyanoptera. Cinnamon Teal. 23 April 1973, 15 May 1973, 14 April 1974. ""Anas americana. American Wigeon. 1 May 1974, 3 birds in one flock. ""Melanitta deglandi. White-winged Scoter. Two birds flew across Gunnison Island 15 April 1972. ""Oxyiira jamaicensis. Ruddy Duck. 14, 18 May 1973; 9, 15 May 1974. ""Mergiis senator. Red-breasted Merganser. 16 April 1974. ""Cathartes aura. Turkey Vulture. 23 May 1973. ""Accipiter gentilis. Goshawk. 25 April 1973. ""Accipiter cooperii. Cooper's Hawk. Two birds on Gunnison Island 6-7 May 1974. Buteo swainsoni. Swainson's Hawk. 11 June 1948 (Behle 1949), 9 June 1974. "Aquila chrysaetos. Golden Eagle. 21 May 1974. "Circus cyaneus. Marsh Hawk. 4 August 1972, 6 April 1973, 15 May 1973, 24 July 1974. Falco mexicanus. Prairie Falcon. One pair successfully nested on cliffs at the north end of Gunnison Island each year, 1972-1974. The species has apparently nested at the site annually since 1944 (Behle 1958). ""Falco columharius. Pigeon Hawk. 6 May 1973. "Falco sparverius. American Kestrel. 26 April 1972, 24 July 1974. ""Fulica americana. American Coot. 4, 23 April 1973, 14 May 1973. "Charadrius vociferus. Killdeer. 4, 7 May 1973, 15 April 1974. "Pluvialis squatarola. Black-bellied Plover. 23-25 April 1973. ""Capella gallinago. Common Snipe. 23 April 1973, 4 May 1973. "Numenius americanus. Long-billed Curlew. 20 April 1972, 3 May 1973. ""Tringa macularia. Spotted Sandpiper. 6, 15-16 May 1973; 22-23 May 1974. "Catoptrophorus semipalmatus. Willet. 3, 8 May 1974. A total of 10 birds in 3 flocks. ""Calidris hairdii. Baird's Sandpiper. 15 May 1973. A total of 26 individuals in a mixed flock with Calidris pusilla. June 1977 Knopf, Backman: Utah Birds 249 ""Calidris pusiUa. Semipalmated Sandpiper. 15 May 1973, 3 individuals. °°Lwiosa fedoa. Marbled Godwit. 3 May 1974 and 24 July 1974, a total of 36 in- dividuals. "Recunirostni americana. American Avocet. 16 April 1974. Himantopus mexicanus. Black-necked Stilt. Previously recorded on Gunnison Island (Behle 1958). Twenty-eight individuals seen in one flock 6 May 1974. ° Phalaropus tricolor. Wilson's Phalarope. 20 June 1973. Lams californicus. California Gull. Gun- nison Island is a traditional nesting site for California Gulls (Behle 1958). Num- bers of the birds were estimated to be about 15,000 individuals annually, 1972- 1974. °°Larus pipixcan. Franklin's Gull. 4 May 1973. 'Sterna caspia. Caspian Tern. A single Cas- pian Tern "mobbed " investigators on Gunnison Island 4 August 1972. A nest was not located. Four nests (with in- complete clutches) were discovered at the same location 10 June 1974. A fifth nest was also observed 11 June 1974. ""Columba livia. Rock Dove. 5 May 1973. "Zenaida macroura. Mourning Dove. Late April through mid June annually. No nests were located. Asio flammeus. Short-eared Owl. Permanent resident of Gunnison Island (Behle 1958). This species was present 1972- 1974, with as many as 4 individuals being seen simultaneously. One in- vestigator was harassed on Gunnison Is- land 20 May 1974 by a single bird, but no nest was found. ° °Selasphorus platycercus. Broad-tailed Hummingbird. 4, 14, 16 May 1973; 1, 23 May 1974. "Megaceryle alcyon. Belted Kingfisher. 23 April 1973. 'Colaptes auratus. Common Flicker. 6 April 1973. '"Tyrannus verticalis. Western Kingbird. 2 May 1974. ""Tyrannus vociferans. Cassin's Kingbird. 5 May 1973. This is the northernmost re- cord of the Cassin's Kingbird in Utah. Myiarchus cinerascens. Ash-throated Fly- catcher. 11 June 1949 (Behle 1958), 8, 22 May 1974. Sayornis saya. Say's Phoebe. 11 June 1949 (Behle 1958), 6 May 1974. " ° Empidonax hanimondii. Hammond's Fly- catcher. 1 May 1974. This bird was considered E. hammondii based upon the contrast between yellowish under- parts and a grayish crest, by olive- colored tinge on the back, and by tail- flicking behavior while perched. How- ever, no definite means of discerning between E. hammondii and E. oherhol- seri is reliable in the field, and the clas- sifying of this sighting as hammondii must be considered speculative. Empidonax oberholseri. Dusky Flycatcher. This species was reported 8 June 1937 (Marshall and Leatham 1942), but its identity was questioned by Behle (1958). This species failed to appear in Rawley (1976) and was incorrectly list- ed as not observed on Gunnison or Cub Island. No sightings were available 1972-1974. °'' Empidonax wrightii. Gray Flycatcher. 14 May 1973; 15^ 21 May 1974. °° Empidonax difficiUs. Western Flycatcher. 22-23, 28 May 1974. "Contopus sordidulus. Western Wood Pewee. 4-14 May 1973, 14-22 May 1974. " Eremophihis alpestris. Horned Lark. 23 April 1973, a flock of 8 individuals. ""TacJiycineta thahissina. Violet-green Swallow. 1 May 1974. ""Tachycineta hicolor. Tree Swallow. 6 April 1973, 14 May 1974. °°Riparia riparia. Bank Swallow. 1 May 1974. ° " Stelgidopteryx riificoUis. Rough-winged Swallow. 22 May 1974. "Hinindo nistica. Barn Swallow. 25 April 1973; 5-6 May 1973; 1, 8-10, 15, 22 May 1974. ° " PetrocheUdon pyrrJwnota. Cliff Swallow. 25 April 1973,' 10 May 1974. Corviis corax. Common Raven. A common nesting species (Behle 1958). One pair nested annually 1972-1974. "Corvus hrachyrhynchos. Common Crow. 250 Great Basin Naturalist Vol. 37, No. 2 16 April 1972. ''°Giimnor}iinus cyanocephalus. Pinon Jay. 15 May 1974, a single flock of 24 birds. ""Parus gambeli. Mountain Chickadee. 22 May 1974, 3 individuals were seen in one flock in Atriplex bushes. °''Cinchis mexicanus. Dipper. 12 May 1972, a lone bird sitting among rocks and in apparent good health. Salpinctes obsoletus. Rock Wren. Nesting and abundant (Behle 1958), 1972-1974. "Mimus poly glottis. Mockingbird. 28 May 1974. °Du7netella carolinensis. Gray Catbird. 23 April 1973. "Oreoscoptes montanus. Sage Thrasher. 30 April 1974; 6, 15, 28 May 1974; 11 June 1974. "Tardus migratorius. American Robin. 23 April 1973, 4 May 1973, 29 April 1974, 2 May 1974. "Cathanis guttatus. Hermit Thrush. 4-5, 8, 14 May 1973; 29 April 1974; 2 May 1974. ""Catharus ustulatus. Swainson's Thrush. 16 April 1972; 15, 22 May 1974. ""Myadestes townsendi. Townsend's Soli- taire. 6 May 1973. "Polioptila caerulea. Blue-gray Gnatcatcher. 12 April 1973; 5-6 May 1973; 29 April 1974; 7, 21 May 1974. °°Reguhis calendula. Ruby-crowned King- let. 15 April 1972; 4, 8, 14 May 1973; 1-2, 8, 15 May 1974. ""Anthus spinoletta. Water Pipit. 23-24, 26- 27 April 1973; 15, 29 April 1974; 1 May 1974. "Lanius ludovicianus. Loggerhead Shrike. Two pairs nested in 1973 with one nest resulting in 4 fledglings. A single sight- ing on 29 April was the only 1974 re- cord. "Vireo gilvus. Warbling Vireo. 29 May 1974. ""Vennivora celata. Orange-crowned War- bler. 5 May 1973. Dendroica petechia. Yellow Warbler. Pre- viously recorded on islands (Behle 1958). Common 8-13 Mav 1973, 6-24 May 1974. "Dendroica coronata auduhoni. Yellow- rumped Warbler. 25 April 1973; 4 May 1973; 30 April 1974; 1, 7, 11, 15, 22, 29 May 1974. D.c. coronata. 16 May 1973. ""Dendroica nigrescens. Black-throated Gray Warbler. 9-10 May 1974. ""Geothlypis tohniei. MacGillivray's War- bler. 6, 8 May 1973; 2, 14, 22-24, 28 May 1974. ""Wiisonia pusilla. Wilson's Warbler. 8 May 1973; 6, 10-11, 22-23, 28 May 1974. "Setophaga ruticilla. American Redstart. 21 May 1974. "Sturnella neglecta. Western Meadowlark. 12 May 1972, 25-26 April 1973, 5 May 1973, 1-2 May 1974. Xanthocephalus xanthocephalus. Yellow- headed Blackbird. Previously recorded on Gunnison Island (Behle 1958). 28 May 1972; 23-24 April 1973; 5, 14, 23 May 1973; 22-23, 29 May 1974. "Agelaius phoeniceus. Red-winged Black- bird. 15-16 April 1972; 24 April 1973; 3, 5-6 May 1973. "Icterus galhula. Northern Oriole. 4 May 1973, 22 May 1974. "Euphagus cyanocephalus. Brewer's Black- bird. 24, 26-27 April 1973; 3-8, 14, 23 May 1973; 15 April 1974; 3, 23 May 1974. Molothrus ater. Brown-headed Cowbird. Previously recorded on Gunnison Island (Behle 1958). 24 April 1973; 22-24, 29 May 1974; 24-25 July 1974 Pheucticus melanocephalus. Black-headed Grosbeak. Previously recorded (Behle 1958), but not seen 1972-1974. "Passerina amoena. Lazuli Bunting. 6 May 1973. "Carpodacus mexicanus. House Finch. 16 April 1972, 10 May 1974. "Pipilo chlorura. Green-tailed Towhee. 4-8, 14, 23 May 1973; 1-2, 7-10, 14-15, 22- 23, 29 May 1974. Pipilo erythrophthalmus. Rufous-sided tow- hee. 15-16 April 1972, 4 May 1973, 7 May 1974. "Passerculus sandivichensis. Savannah Spar- row. 29-30 April 1974. "Pooecetes gramineus. Vesper Sparrow. 4-6 May 1973, 30 April 1974, 9 May 1974. "Chondestes grammacus. Lark Sparrow. 4-5, 1977 Knopf, Backman: Utah Birds 251 7 May 1973; 7, 10, 14, 21-22 Mav 1974. "Aimophila bilineata. Black-throated Spar- row. 2-3 May 1973. °°Aimophila belli. Sage Sparrow. 16 April 1972. ""Junco Injemalis. Dark-eyed Junco. 5-6, 23- 24, 26-27 April 1973;' 13 May 1973; 15 April 1974. ""Junco caniceps. Gray-headed Junco. 8 May 1973, 8 May 1974. ""Spizella arborea. Tree Sparrow. 21 May 1974. Spizella passerina. Chipping Sparrow. Pre- viously recorded on Gunnison Island (Behle 1958). 13 May 1973; 30 April 1974; 1, 6-7, 14-15, 22, 24, 29 May 1974. ""Spizella pallida. Clay-colored Sparrow. 9 May 1974. "Spizella breweri. Brewer's Sparrow. 5-6, 8 May 1973; 30 April 1974; 9-10, 15, 23- 24 May 1974. "Zonotrichia leiicophnjs. White-crowned Sparrow. 15-16 April 1972; 23 April 1973; 3, 5-6 May 1973; 29 April 1974; 2, 6-10, 14-15, 23-24 May 1974. ""Zonotrichia melodia. Song Sparrow. 15 April 1972. Comments on the Avifauna The avifauna of Gunnison and Cub is- lands consisted primarily of transient spe- cies. Of 112 species, 104 (93 percent) oc- curred as transients. Transients which found abundant food on the islands (e.g., individ- uals of the Columbidae and Fringillidae) were present in larger numbers and for longer periods of time than those not find- ing food (e.g., individuals of the Anatidae and Scolopacidae). None of the species oc- curring as transients were present long enough to be considered seasonal residents. The Short-eared Owl was the only non-nest- ing species considered to be a resident. Species observed on the two islands were typical of those occurring in similar habitats throughout northern Utah. Some notable ex- ceptions were the "sedentary" Dipper and Mountain Chickadee, both being 40 km from their typical habitats. The Black-billed Magpie {Pica pica), which is abundant in the region, was not seen on the islands. Few species of birds actually nested on Gunnison and Cub islands relative to the surrounding Great Basin desert. Many of the desert species (Mourning Dove, Horned Lark, Lark Sparrow, Brewer's Sparrow) were observed on the islands in large flocks during spring migration. The impoverished community of breeding birds may reflect poorer nesting habitat for these species. Habitats for Loggerhead Shrike appeared marginal (as developed by Johnson 1965:121), with two pairs breeding in 1973, where none bred in 1972 or 1974. Acknowledgments These observations were recorded in- cidental to projects conducted under the su- pervision and funding of the Utah Coopera- tive Wildlife Research Unit, Logan, Utah, and the Rob and Bessie Welder Wildlife Foundation, Sinton, Texas. Special thanks go to Jessop B. Low for his encouragement and assistance. We thank the Utah Division of Wildlife Resources for access privileges to the islands. Literature cited Behle, W. H. 1949. Report on the colonial nesting birds of Great Salt Lake, 1947-49. Condor 51:269-270. 1958. The bird life of Great Salt Lake. Univ. Utah Press. Salt Lake City. 203 p. Johnson, N. K. 1965. The breeding avifaunas of the Sheep and Spring ranges in southern Nevada. Condor 67:93-124. Knopf, F. L. 1974. Fluctuating levels of Great Salt Lake: Effects on island wildlife. Proc. Utah Acad. Sci., Arts, and Letters 51 (2):34-41. 1975. Spatial and temporal aspects of colonial nesting of the white pelican, Pelecanus enjthro- rhynchos. Ph.D. Dissertation. Utah State Univer- sity, Logan. 77 p. 1976. Ancient Murrelet in Utah. Western Birds 6:27. Marshall, W. H. 1940. A survey of the islands in Great Salt Lake, Utah. J. Mammal. 21:144-159. Marshall, W. H. 1940. A survey of the mammals of the islands in Great Salt Lake, Utah. J. Mammal. 21:144-159. R\WLEY, E. V. 1976. Small islands of Great Salt Lake. Utah Div. Wildl. Resour. Pub. No. 76-19. 109 p. VARYING PREVALENCE OF TAENIA TAENIAEFORMIS STROBILOCERCI IN MICROTUS PENNSYLVANICUS OF MONTANA Richard H. McBee, Jr.' Abstract.- Of 42 Microtits pcnnsijhaniciis trapped 1 km from Bozeman, Montana, 14 (33.3 percent) were in- fected hv strobilocerci of Taenia taeniae fonnis, while only one (2.1 percent) of 47 was infected at a niral site 16 km from Bozeman. The higher incidence of the primary host, domestic cats, is believed to account for the higher rate of infection near Bozeman. Microtits pennsylvanicus has previously been reported as an intermediate host for Taenia { = Hydatigera) taeniaeformis by Rausch and Tiner (1949, Am. Midi. Nat. 41:665-694), who found 43 of 570 voles in the north central U.S. infected, and by Kin- sella (1967, Can. J. Zool. 45:269-274), who found strobilocerci in 2 of 25 voles from western Montana. Survey data collected in southwestern Montana in 1971 indicated that the preva- lence of infection of M. pennsylvanicus with T. taeniaeformis strobilocerci may vary widely according to the proximity of the voles to large numbers of primary hosts such as the domestic cat. Voles were trapped from boggy areas on two different sites near the city of Boze- man, Gallatin County, Montana. The first area was located just outside the city limits and within easy range of numerous house cats. The second area was located approx- imately 16 km (10 mi) from Bozeman in farm land, with the nearest residence more than 1 km (.6 mi) away. A total of 42 M. pennsylvanicus was trapped from the first area, and their livers were examined for the strobilocerci of T. taeniaeformis. Infected voles totaled 14 (33.3 percent positive), with several animals having multiple cysts (an average 1.86 cysts per infected animal). A total of 47 voles was examined from the second area and, of these, one animal (2.1 percent) was found to be infected, with its liver harboring a single cyst. The low incidence of infection of the ru- ral M. pennsylvanicus with T. taeniaeformis is attributed to the lack of large numbers of primary hosts in the area. 'Department of Zoology, Montana State Univ 97470. iity, Bozeman, Montana .59715. Current address: Roseburg Senior High School, Roseburg, Oregon 252 AN ANNOTATED LIST OF THE AQUATIC INSECTS OF SOUTHEASTERN IDAHO, PART II: TRICHOPTERA Robert L. Newell' and G. Wayne Minshall' Abstract.— Distribution records for the Trichoptera occurring in the southeastern one-third of Idaho are sum- marized based on published records and collections by the authors and their colleagues. The study revealed rep- resentatives of 13 families, 37 genera, and 84 species. Several species were collected for the first time in Idaho, including: Hesperophijlax rnagnus, Limtiephilus frijole and L. thorns, Psychoglypha ormiae, Ceraclea annnlicornis and C. copha, Pohjcentropus crassicornis, Tinodes provo, and Agnjpania dextra. The Trichoptera (caddisflies) comprise one of several purely aquatic orders of in- sects. This small order is composed of some 30 families and 5,000 species distributed throughout the world. This study revealed representatives of 13 families, 37 genera, and 84 species. Some genera were captured, but a species determination was not possible and these are not listed. The study area in- cludes the southeastern one-third of Idaho (Fig. 1). Note that this figure corrects and replaces the one given in Newell and Min- shall (1976). Information on geographical distribution of caddisflies in Idaho is widely distributed. Local investigations that included distribu- tional records are Andrews (1972), Givens (1976), Newell and Potter (1973), Nimmo (1971), Kroger (1970), Smith (1965, 1968a, 1968b, 1969), Terch (1972), Winger (1968) and Wold (1974). Other Idaho records are available in Denning (1948, 1950, 1954, 1956, 1958, 1965a, 1965b, 1970, 1973), Denning and Blickle (1972), Gordon (1974), Milne and Milne (1938), Hornung and Barr (1970), Ross (1941, 1944, 1947, 1951, 1952, 1956), and Schmid (1968, 1970). Identi- fication of immature caddisflies will be aided greatly by the key of Wiggins (1977). Several of the above-listed papers include taxonomic keys to adult caddisflies. The tax- onomic treatment used throughout this pa- per is that of Ross (1967). Several species were collected for the first time in Idaho, and these include: Hes- perophijlax itiagnus, Limnephihis frijole and L. thorns, Psychoohjpha onniae, Ceraclea annnlicornis and C. copha, Polycentropus crassicornis, and Tinodes provo and Agr- ijpania dextra. Individuals of the family Hydropsychidae were the most frequently encountered. Indi- viduals of Hydropsijche occidentalis were extremely abundant in the main Snake Riv- er, often visible in swarms of thousands of adults. Several species were encountered only once, either because of their small populations or because their habitats were difficult to collect (i.e., high altitude streams). The large number of species collected is probably indicative of the diversity of avail- able habitats, ranging from spring seeps to large mountain rivers and from silty low- land streams and irrigation reservoirs to constant temperature springs. The authors would like to thank the many people who contributed time, effort, and specimens for this study. They would also like to thank Dr. Norman H. Anderson and Dr. A. P. Nimmo for their suggestions concerning the manuscript and Dr. Donald G. Denning for his help in identifying spec- imens. Family hydropsychidae 1. Arctopsyche grandis (Banks).— Bear Lake Co.: St. Charles Cr. Blaine Co.: Big Stearns-RoHer. P.O. Box .5888, Denver, Colorado 80217 Department of Biology, Idaho State University. Poeatello, Idaho 8.3209 253 254 Great Basin Naturalist Vol. 37, No. 2 Wood R. Bonneville Co.: Pine Cr., Rainey Cr., Snake R. (S. Fk.). Butte Co.: Little Lost R. Caribou Co.: Portneuf R., Tin Cup Cr. Clark Co.: Birch Cr., Medicine Lodge Cr. Custer Co.: Big Lost R. 2. CheumatopsycJie campijla Ross.— Bon- neville Co.: Idaho Falls. Franklin Co.: Pres- ton. Gooding Co.: Niagara Springs, Snake R. Twin Falls Co.: Rock Cr. 3. Chemnatopsijche enonis Ross.— Bonne- ville Co.: Idaho Falls. Twin Falls Co.: Salmon Falls Cr. 4. Cheumatopsyche pettiti (Banks).— Bear Lake Co.: Dingle. Bonneville Co.: Brock- man Cr. Caribou Co.: Lane's Cr. Oneida Co.: Deep Cr. 5. Hijdropsijche bifida Banks.— Bannock Co.: Mink Cr. 6. Hydropsyche califomica Banks.— Ban- nock Co.: Portneuf R. Bingham Co.: Spring Cr. Gooding Co.: Niagara Springs. Oneida Co.: Deep Cr. Owyhee Co.: Snake R. Pow- er Co.: Snake R. 7. HydropsycJie cockerelli Banks.— Bonne- ville Co.: Snake R. Teton Co.: Teton R. 8. Hydropsyche occidentalis Banks.— Ban- nock Co.: Portneuf R. Caribou Co.: Port- neuf R. Fremont Co.: Falls R. Gooding Co.: Niagara Springs. Oneida Co.: Deep Cr. Power Co.: Snake R. 9. Hydropsyche oslari Banks.— Bannock Co.: Mink Cr. Bonneville Co.: Pine Cr., Rainey Cr. Teton Co.: Trail Cr. 10. Parapsyche ahnota Ross.— Bannock Co.: Mink Cr. 11. Parapsyche elsis Milne.— Blaine Co.: Big Wood R. Bonneville Co.: Rainey Cr. Butte Co.: Little Lost R. Franklin Co.: Cub R. Lemhi Co.: Little Lost R. Family polycentropodidae 12. Pohjcentropus cinereus Hagen.— Bing- ham Co.: Spring Cr. 13. Pohjcentropus crassicornis Walker.— Bingham Co.: Spring Cr. Family psychomyiidae 14. Psychomyia flavida Hagen.— Fremont Co.: Falls R., Robinson Cr., Snake R. (Hen- ry's Fk.). Power Co.: Snake R. 15. Tinodes provo Ross.— Gooding Co.: Niagara Springs. Oneida Co.: Deep Cr. Family philopotamidae 16. DolophUodes aequaUs (Banks).— Ban- nock Co.: Mink Cr., Rapid Cr. Bonneville Co.: Rainey Cr. 17. Wonnaldia nock Co.: Mink Cr. gahriella (Banks).— Ban- Fig. 1. An outline map of Idaho showing the location of the study area (shaded portion) in relation to the major river drainage systems. Note: This figure re- places the one given in Newell and Minshall (1976) in which the shaded area was lost in the printing process. Family rhyacophilidae 18. Rhyacophila acropedes Banks.— Ban- nock Co.: Mink Cr. Bonneville Co.: Pine Cr., Rainey Cr. Butte Co.: Little Lost R. Clark Co.: Birch Cr., Medicine Lodge Cr. Gooding Co.: Niagara Springs. Teton Co.: Trail Cr. 19. Rhyacophila alberta Banks.— Bonne- ville Co.: Rainev Cr. Teton Co.: Trail Cr. June 1977 Newell, Minshall: Idaho Trichoptera 255 20. Rhyacophila angelita Banks.— Bonne- ville Co.: Rainey Cr. 21. Rhiiacophila hifilo Banks.— Fremont Co.: Falls Cr. 22. Rhyacophila coloradensis Banks.— Bon- neville Co.: Pine Cr., Rainey Cr. Gooding Co.: Hagerman Springs, Niagara Springs. Teton Co.: Trail Cr. 23. Rhyacophila hyalinata Banks.— Butte Co.: Little Lost R. 24. Rhyacophila narvae Navas.— Butte Co.: Little Lost R. 25. Rhyacophila pellisa Ross.— Bannock Co.: Mink Cr. Bonneville Co.: Rainey Cr. Lemhi Co.: Little Lost R. 26. Rhyacophila vaccua Milne.— Clark Co.: Birch Cr. Lemhi Co.: Little Lost R. Teton Co.: Trail Cr. 27. Rhyacophila vagrita Milne.— Bonne- ville Co.: Rainey Cr. 28. Rhyacophila vao Milne.— Bannock Co.: Mink Cr. Blaine Co.: Big Wood R. 29. Rhyacophila verrula Milne.— Bonne- ville Co.: Rainey Cr. Lemhi Co.: Little Lost R. Teton Co.: Trail Cr. 30. Rhyacophila vocala Milne.— Bonne- ville Co.: Rainev Cr. Fam ily glossosomatidae 31. Agapetus montanus Denning.— Clark Co.: Beaver Cr. 32. Anagapetus debilis Ross.— Custer Co.: Wildhorse Cr. 33. Glossosoma alascensis Banks.— Bonne- ville Co.: Pine Cr. Custer Co.: Wildliorse Cr. Fremont Co.: Robinson Cr. Teton Co.: Trail Cr. 34. Glossosoina idaho Ross.— Fremont Co.: Falls R. Gooding Co.: Niagara Springs. 35. Glossosoma intennedium (Klapalek).— Teton Co.: N. Fk. Teton R. 36. Glossosoma montana Ross.— Bonne- ville Co.: Snake R. (S. Fk.). Fremont Co.: Falls R. 37. Glossosoma schiihi Ross.— Bannock Co.: Mink Cr. 38. Glossosoma traviatum Banks.— Fre- mont Co.: Falls R. 39. Glossosoma velona Ross.— Bannock Co.: Mink Cr. Bingham Co.: Spring Cr. Caribou Co.: Portneuf R. Teton Co.: Teton R. 40. Glossosoma verdona Ross.— Bonneville Co.: Rainev Cr. 41. Protoptila coloma Ross.— Fremont Co.: Warm R. Fam ILY HYDROPTILIDAE 42. Ochrotrichia logana (Ross).— Gooding Co.: Niagara Springs. 43. Ochrotrichia stylata (Ross).— Power Co.: Snake R. Family lepidostomatidae 44. Lepidostoma pluvialis (Milne).— Ban- nock Co.: Mink Cr. 45. Lepidostoma podager McLachlan.— Bannock Co.: Mink Cr. 46. Lepidostoma strophis Ross.— Bingham Co.: Spring Cr. Gooding Co.: Niagara Springs. Oneida Co.: Daniels Spring. 47. Lepidostoma unicolor (Banks).— Ban- nock Co.: Mink Cr. Bonneville Co.: Pine Cr. Fam ILY LIMNEPHILIDAE 48. Amphicosmoecus canax (Ross).— Bing- ham Co.: Spring Cr. Bonneville Co.: Rainey Cr. Custer Co.: Little Lost R. Fremont Co.: Snake R. (Henry's Fk.). Teton Co.: Trail Cr. 49. Apatania chasica (Denning).— Blaine Co.: Big Wood R. Bonneville Co.: Rainev Cr. 50. Apatania comosa Denning.— Bonne- ville Co.: Pine Cr. 51. Chyranda centralis (Banks).— Bannock Co.: Mink Cr. Custer Co.: Wildhorse Cr. 52. Ecclisomyia maculosa Banks.— Lemhi Co.: Little Lost R. Teton Co.: Trail Cr. 53. Ecclisomyia simtdata Banks.— Bonne- ville Co.: Rainey Cr. 54. Hesperophylax consimilis (Banks).— Bannock Co.: Mink Cr. Bonneville Co.: Rainey Cr. Caribou Co.: Portneuf R. 55. Hesperophylax incisi4S Banks.— Ban- nock Co.: Mink Cr. Bonneville Co.: Rainey Cr. Fremont Co.: Buffalo R. 56. Hesperophylax magnus Banks.— Bonne- ville Co.: Rainey Cr. 256 Great Basin Naturalist Vol. 37, No. 2 57. Lenarchus brevipennis (Banks).— Fre- mont Co.: Pond near Upper Mesa Falls. 58. Limnephihis challisa Denning.— Blaine Co.: Hyndman Cr. 59. Limnephilus extractus Walker.— Fre- mont Co.: Snake R. (Henry's Fk.). 60. Limnephilus frijole Ross.— Oneida Co.: Deep Cr. 61. Limnephihis spinatus Banks.— Bannock Co.: Hawkins Res. Bonneville Co.: Pond near Palisades Res. Caribou Co.: Grey's Lake. Fremont Co.: Snake R. (Henry's Fk.). Oneida Co.: Daniels Res. 62. Limnephihis tJiorus Ross.— Bingham Co.: Clear Cr. 6.3. Neophylax occidentis Banks.— Lemhi Co.: Firebox Cr. 64. Neophylax splendens Denning.— Ban- nock Co.: Mink Cr. Bonneville Co.: Rainey Cr. 65. Neothremmo ahcia Banks.— Teton Co.: Trail Cr. 66. Ohgophlebodes minutiis (Banks).— Ban- nock Co.: Gibson Jack Cr., Mink Cr. 67. Ohgophlebodes riithoe Ross.— Camas Co.: Big Lost R. 68. Oligophlebodes sierra Ross.— Bear Lake Co.: Bloomington Cr., St. Charles Cr. Bonneville Co.: Rainey Cr. 69. Onocosmoecus atripes (Hagen).— Ban- nock Co.: Mink Cr. Bingham Co.: Spring Cr. Bonneville Co.: Rainey Cr. 70. Onocosmoecus unicolor (Banks).— Ban- nock Co.: Mink Cr. Bingham Co.: Spring Cr. Bonneville Co.: Rainev Cr. Teton Co.: Trail Cr. 7L Psychoglypha ormiae (Ross).— Bonne- ville Co.: Rainey Cr. 72. Psychoglypha subhorealis (Banks).— Bannock Co.: Mink Cr. Bonneville Co.: Rainey Cr. Family phryganeidae 73. Agrypania dextra Ross.— Bannock Co.: Pond in Indian Rocks St. Park. Family brachycentridae 74. Amiocentrus aspilus (Ross).— Bingham Co.: Clear Cr., Spring Cr. Bonneville Co.: Rainev Cr. Teton Co.: Teton R. 75. Brachycentnis americanus (Banks).— Bannock Co.: Mink Cr. Bingham Co.: Clear Cr. Bonneville Co.: Pine Cr. Caribou Co.: Portneuf R. 76. Brachycentnis occidentalis Banks.— Bingham Co.: Spring Cr. Bonneville Co.: Rainey Cr. Butte Co.: Little Lost R. Clark Co.: Beaver Cr., Birch Cr., Camas Cr., Medicine Lodge Cr. Family helicopsychidae 77. Helicopsyche borealis (Hagen).— Bing- ham Co.: Clear Cr. Caribou Co.: Blackfoot R., Lane's Cr., Portneuf R. Clark Co.: Birch Cr., Camas Cr. Teton Co.: Warm Spring Cr. Fam ILY LEPTOCERIDAE 78. Ceraclea annulicornis (Stephens).— Power Co.: Snake R. 79. Ceraclea copha (Ross).— Power Co.: Snake R. 80. Ceraclea tarsipunctata (Vorhies).— Bingham Co.: Snake R. 8L Ceraclea transversa (Hagen).— Cassia Co.: Small stream near Malta. 82. Mystacides alafimbriata Griffin.— Te- ton Co.: Pond near Teton R. 83. Nectopsyche diarina (Ross).— Blaine Co.: Big Wood R. Oneida Co.: Deep Cr. 84. Triaenodes frontalis Banks.— Bingham Co.: Spring Cr. Oneida Co.: Deep Cr. Literature cited Andrews, D. A. 1972. An ecological study of the lost streams of Idaho with emphasis on the Little Lost River. Unpubl. Master's Thesis, Idaho State Univ., Pocatello. 57 p. Denning, D. G. 1948. A review of the Rhyacophilidae (Trichoptera). Can. Ent. 80:97-117. 1950. Records and descriptions of nearctic caddisflies. Part II. J. Kansas Ent. Soc. 23:115- 120. 1954. New species of western Trichoptera. J. Kansas Ent. Soc. 27:57-64. 1956. Several new species of western Trichop- tera. Pan-Pac. Ent. 32:73-80. 1958. New western Trichoptera. Pan-Pac. Ent. 34:93-98. 1965a. New Trichoptera from United States and Mexico. Pan-Pac. Ent. 41:262-272. 1965b. New rhyacophilids and limnephilids June 1977 Newell, Minshall: Idaho Trichoptera 257 (Trichoptera: Rhyacophilidae and Limnephilidae). Can. Ent. 97:690-700. 1970. The genus Psychoghjpha (Trichoptera: Limnephihdae). Can. Ent. 102:15-30. 1973. New species of Trichoptera. Pan-Pac. Ent. 49:132-143. Denning, D. C, and R. L. Blickle. 1972. A review of the genus Ochrotrichia (Trichoptera: Hydro- ptihdae). .Ann. Ent. Soc. Amer. 65:141-151. GivENS, D. R. 1976. Synopsis of western Arctopsy- chinae (Trichoptera: Hydropsychidae). Unpubl. Master's Thesis, Central Washington St. College, Ellensburg. Gordon, A. E. 1974. A synopsis and phylogenetic out- line of the nearctic members of Cheumatop syche. Proc. Acad. Nat. Sci. Phil. 126:117-160. HoRNUNG, D. S., .\ND W. F. Barr. 1970. Insects of Craters of the Moon National Monument, Idaho. Univ. of Idaho, Col. of Agri., Misc. Ser. No. 8. 118 p. Kroger, R. L. 1970. An annotated list of stream in- vertebrates collected from Grand Teton National Park, Wyoming. Univ. Wyoming Publ. 36:1-14. Milne, L. J., and M. J. Milne. 19.38. The Arctopsy- chidae of continental America north of Mexico (Trichoptera). Bull. Brooklyn Ent. Soc. 33:97-110. Newell, R. L., and G. W. Minshall. 1976. An anno- tated checklist of the aquatic insects of south- eastern Idaho. Part I. Plecoptera. Great Basin Nat. 36(4): 501-504. Newell, R. L., and D. Potter. 1973. Distribution of some Montana caddisflies (Trich.). Proc. Montana Acad. Sci. 33:12-21. NiMMO, A. P. 1971. The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta and east- ern British Columbia and their past glacial origin. Quaestiones Entomologicae 7:3-234. Ross, H. H. 1941. Descriptions and records of North American Trichoptera. Trans. Amer. Ent. Soc. 67:.35-126. 1944. The caddisflies or Trichoptera of Illinois. Bull. 111. Nat. Hist. Surv. No. 23. 136 p. 1947. Descriptions and records of North Amer- ican Trichoptera, with synoptic notes. Trans. Amer. Ent. Soc. 73:125-168. 1951. The caddisflv genus Anagapetii.s. Pan- Pac. Ent. 27:140-144. 1952. An annotated key to the nearctic males of Limnephilus (Trichoptera, Limnephilidae). Amer. Midi. Nat. 47:435-455. 19.56. Evolution and classification of the moun- tain caddisflies. Univ. Illinois Press. Urbana. 213 p. 1967. The evolution and past dispersal of the Trichoptera. Ann. Rev. Ent. 12:169-206. Schmid, F. 1968. La famille des .Arctopsychides (Tri- choptera). Mem. Soc. Ent. Que. No. 1:1-84. 1970. Le genre Rhyacophila et la famille des Rhyacophilidae (Trichoptera). Mem. Soc. Ent. Can. 66:1-230. Smith, S. D. 1965. Distributional and biological re- cords of Idaho caddisflies (Trichoptera). Ent. News 76:242-245. 1968a. The Rhyacophila of the Salmon River drainage of Idaho with special reference to lar- vae. Ann. Ent. Soc. Amer. 61:655-674. 1968b. The Arctopsychinae of Idaho (Trichop- tera: Hydropsychidae). Pan-Pac. Ent. 44:102-112. 1969. Two new species of Idaho Trichoptera with distributional and taxonomic notes on other species. J. Kansas Ent. Soc. 42:46-53. Terch, J. C. 1972. Bionomics of the caddisfly (Tri- choptera) fauna of Rainey Creek, Bonneville Co., Idaho, with special reference to the life histories of five selected species. Unpubl. Master's Thesis, Idaho State Univ., Pocatello. 53 p. Wiggins, G. B. 1977. Larvae of the North American caddisfly genera. Univ. of Toronto Press, (in press). Winger, P. V. 1968. The invertebrate drift in two mountain streams. Unpubl. Master's Thesis, Idaho State Univ., Pocatello. 76 p. Wold, J. 1974. Systematics of the genus Rhyacophila (Trichoptera: Rhyacophilidae) in western North America with special reference to the immature stages. Unpubl. Master's Thesis, Oregon St. Univ., Corvallis. 229 p. A NEW STATUS FOR PENSTEMON CYANANTHUS HOOK. SSP. SUBGLABER PENNELL (SCROPHULARIACEAE) Stephen L. Clark' Abstract.— Penstemon cyananthus Hook. ssp. subgkiber Pennell, a plant found frequently in northwestern Utah, southeastern Idaho, and southeastern Wyoming, is elevated to species status. A new name, Penstemon holm- grenii Clark is proposed because P. subglaber is preoccupied. glaber is densely covered with long yellow hairs, and the two taxa are not sympatric. These differences suggest to the writer that the subspecies should be elevated to species status. The following new name is proposed to replace the epithet Penstemon cyananthus Hook. ssp. subglaber Pennell: Penstemon holmgrenii Clark nom. nov., based on P. cyananthus Hook. ssp. subglaber Pennell in Scrophulariaceae of the Rocky Mountain States. Contributions from the United States National Herbarium 20(9):313-381. Not P. subglaber Rydb. Named in honor of Arthur H. Holmgren, a friend who first guided me Department of Botany, Weber State College, Ogden, Utah 84408. intO StudiCS iu PenStemOU. Penstemon cyananthus Hook. ssp. sub- glaber Pennell is a tall, densely puberulent to pubescent, blue-flowered plant found commonly within sage and pinyon-juniper areas of northwestern Utah, southeastern Idaho, and southwestern Wyoming. Pennell described it in 1920 and separated it from P. cyananthus ssp. cyananthus on the basis that the leaves of the subspecies are more lanceolate and more densely puberulent. In- vestigations by this writer suggest, however, that other differences exist. Penstemon cya- nanthus ssp. cyananthus is never pubescent. The staminode of P. cyananthus ssp. sub- lb^ RANGE EXTENSION OF TRIONYX SPINIFERUS EMORYI INTO UTAH (REPTILIA) Carlos Y. Roby' and Earl C. Loveless' Abstract.— Four Texas Spiny Softshell Turtles {Trionyx spinifertts emoryi) were collected near St. George, Utah. This represents a range extension of 65 km northward into Utah from .Nevada. Of the six subspecies of Trionyx spinifer in the United States, only the Texas Spiny Softshell Turtle, Trionyx spinifenis e7noryi (Agassiz), is known to occur in the Colorado River drainage. Miller (Copeia 1946:46) sug- gested that the species was introduced into this area around the turn of the century when a dam of an artificially stocked pond on a ranch in Grant County, New Mexico, gave way. This permitted the softshell ac- cess to the Gila River, a tributary of the Colorado River. On 24 July 1976, four T. s. emoryi were collected in a small pond within 50 m of the Virgin River, approximately 6 km (4 mi) north of St. George, Utah. The specimens inchide three females and one male (depos- ited at Brigham Young University, BYU 41646-9), with plastral lengths of 12.20, 11.0, 9.5, and 9.1 cm. Seven others were sighted in this pond and thirteen others in the Virgin River. This represents a northern extension of the known range (Webb, 1962, Univ. Kansas Publ. Mus. Nat. Hist. 13(10):429-611) of ap- proximately 65 km (40 mi) from the pre- vious record 2 km (1.5 mi) above Mesquite, Clarke County, Nevada, and a total exten- sion of approximately 720 km (450 mi) since its introduction into the Gila River. We wish to thank Dr. Andrew H. Barnum for his assistance and the Associated Stu- dents of Brigham Young University for funding this project. We also thank Dr. Wilmer W. Tanner for his suggestions. Life Sciences Mu igham Young University, Pv 259 THE LIFE CYCLE OF HYDROPERLA CROSBY I (PLECOPTERA: PERLODIDAE)' Reed Y. Oberndorfer- and Kenneth W. Stewart- ABSTRA.CT.— Hydroperla croshiji in Texas exhibited a univoltine, fast hfe cycle over the three-year study period. Adults emerged in February-March when mean daily stream temperature reached ca. 15 C. Reared females in the lab deposited up to three egg masses. Mean fecundity of dissected females was 787 eggs/female. Oviposition in the field was observed and described. Eggs were triangular in cross section, brown, and measured 400 jiim X 535 jum. They underwent an ca. seven- month diapause until mean daily stream temperature decreased to 18 C. Eyespots appeared, and hatching fol- lowed in two to three weeks. First instar nymphs were measured and described. Male and female nymphs under- went 12 and 14 instars, respectively, and could be sexed by the sixth. Fast growth occurred in the coldest season and Simuliidae and Chironomidae larvae were preferred food throughout development. Eggs contained a mean of 6.21 cal/mg. Ash-free mean caloric value of last instar nymphs was 6.0 cal/mg. .\dult males and females lost 33.8 percent and 57.6 percent, respectively, of their caloric pool through their ca. 12 days of life. The life cycles and ecology of North American Plecoptera are relatively un- known. The classic works of Needham and Claassen (1925), Claassen (1931), Frison (1929, 1935, 1942), and Kicker (1952) were primarily taxonomic, with a few largely de- scriptive notes on morphology, life cycle events, and ecology. Detailed early studies were made by Wu (1923) for Soyedina val- licularia (Wu) and Holdsworth (1941a, b) for AUonarcijs proteus (Newman). Recent papers by Minshall and Minshall (1966), Harper and Pilon (1970), Schwarz (1970), Tarter and Krumholz (1971), Radford and Hartland-Rowe (1971a, b). Harper and Hynes (1970, 1972), Harper (1973a, b), Vaught and Stewart (1974), Stanford (1975), Gather and Gaufin (1975), Hynes and Hynes (1975), and Hynes (1976) are indicative of a resurgent interest in Plecoptera life cycles as supportive knowledge for structural and functional studies at the population and community level. The only detailed autecological study of a southwestern Nearctic stonefly was reported by Vaught and Stewart (1974) for Neoperia chjtnene (Newman), a widespread species Study supported in part bv U.S. Env Grants .34866 and 34905. 'Department of Biological Sciences, North Te ham Young University, Provo, Utah 84602. occurring only in large, permanent rivers. Life cycles of southwestern species should differ markedly from those of boreal species (Harper and Hynes 1972, Harper 1973a, b), due to post-Pleistocene adaptation to drier, warmer climates, and the effects of stream permanence and physicochemical condi- tions. Hydroperla crosbyi (Needham and Claas- sen) is a large perlodid stonefly that is wide- ly distributed throughout the Mississippi Valley from Indiana and Illinois to Arkansas and Oklahoma (Ricker 1952). Stewart et al. (1974) first reported the .species from Texas. It is common in streams across the Black- land Prairie into the Edwards Plateau and Eastern and Western Cross Timbers in Texas, and it appears to be a dominant spe- cies in the macrobenthic community of Clear Creek in Denton County, Texas, where this study was made. Presence of small nymphs in December, and larger nymphs and adults in March and April, had indicated a fast cycle (Hynes 1970). No pre- vious life cycle studies have been published for any Hydroperla species. Stewart and Stark (1977) described the reproductive mor- Asency Traiiieeship T90()115 and North Texas State University Faculty Research Fund Texas 76203. Oberndorfer's current address: Department of Zoology, Brig- 260 June 1977 Oberndorfer, Stewart: Stonefly Biology 261 phology and mating behavior of H. crosbyi, and Prison (1935) reported nymphal micro- distribution and food habits. The purpose of this research was to make a thorough aute- cological investigation of H. crosbyi, with emphasis on its hfe cycle. Study stream All sampling and observations were done at several sites along Clear Creek in Denton County, Texas, from Highway 455, ca. 0.5 km west of Bolivar, Texas, to the bridge on Highway 428, ca. 4 km northeast of Den- ton. Clear Creek originates in Montague County, Texas, and flows through Cooke and Denton counties, merging with Elm Fork of the Trinity River ca. 5 km above Lake Lewisville. Stream discharge is highly variable, depending upon season and rain- fall, with the highest levels usually occur- ring in spring and lowest levels occurring during the summer. In dry years, the stream becomes intermittent during July-Septem- ber. The substrate consists mainly of sand, with gravel and rock rubble riffles. Packs of leaves and other debris collect on large rocks and fallen tree limbs in riffle areas. The streamside forest is dominated by Cot- tonwood {Populus deltoides), sycamore {Pla- tanus occidentalis), black willow {Salix nigra), and several species of elm {Ulmus spp.). As leaves become conditioned by mi- crobes, a successive invasion by macro- benthos, including shredders, collectors, scrapers, and predators, ensues (Cummins 1974). The role of H. crosbyi in this se- quence was suspected to be that of a domi- nant macropredator. Materials and methods Seasonal growth, instar analysis, food habits, sex ratios, and numbers/m- were de- termined from quantitative and qualitative samples taken from January 1974 through March 1976. Biweekly qualitative samples were taken from October through March in each year except in October and November 1974, when heavy fall rains prevented ac- cess to the stream. A fine (363 jum) mesh net was used when earlier life stages were present, and a larger (1 mm) mesh kick net was used for later instars. Specimens were either preserved in 70 percent isopropyl al- cohol or transported live to the laboratory for rearing studies. Six quantitative samples were taken each month from November 1975 to March 1976 by kicking up an area of 25 cm- in front of a fine (363 jum) mesh net; care was taken to sample all typically available habitat types, including leaf debris and rocky substrate. These were preserved in 70 percent isopropyl alcohol and taken to the laboratory for sorting and identi- fication of ail organisms. Last instar exuviae were collected in March 1976, to aid in a more precise estimate of the adult sex ratio since adults were difficult to obtain in the field. In an attempt to find eggs or smaller instars, 5 liters of damp substrate from areas with no surface flow and from pools were collected monthly during the period of March through September 1974, preserved in 99 percent isopropyl alcohol, and brought to the laboratory for elutriation (Stewart 1975). Aspects of adult emergence, mating, and oviposition were determined from field and laboratory observations. The longevity of laboratory-reared adults was determined by holding them in 8-dram, cotton-stoppered, glass shell vials at room temperature, ca. 24 C. Caloric values of last instar male and fe- male nymphs and their exuviae, newly emerged virgin males, spent (postulated) males, newly emerged virgin females with ova and after ova removal by dissection, spent females (after oviposition), and depos- ited egg masses were determined and com- pared. Energy allocated to reproduction (Pr) was equated to the caloric contents of the total amount of eggs deposited by a female. An instantaneous measure of reproductive eff^ort (IRE) was expressed as a percentage of energy going to production of eggs (Pr) and to growth (Pg). After obtaining live weights, individuals were dried in a vacuum oven at 80 C (-15 psi) for 24 hours, weighed, ground with a mortar and pestle, and pressed into pellets of known weights (10-20 mg).The pellets were combusted in a Gentry-Weigert modification of a Phillipson 262 Great Basin Naturalist Vol. 37, No. 2 microbomb calorimeter (Phillipson 1964). The drv weight of material not combusted was considered to be ash weight; therefore, caloric data were expressed as calories/mg ash-free dry weight. Organisms were not bombed individually, but pooled to produce the pellets. Groups of eggs deposited in March by reared, mated females were incubated in 1974 and 1976. They were held in loosely covered 8-dram vials at 22 ±2 C until Octo- ber (1974) and September (1976), then transferred into a Percival E-50 environ- mental chamber, lowered 1 C per day down to 18 C. Eggs "in uteri" and deposited eggs were counted, photographed, measured, and de- scribed, with special attention given to any gelatinous layers and/or other attachment structures and chorionic sculpturing (Stew- art et al. 1969). Scanning electron micro- graphs were made using an IS! Mini-SEM. First instar nymphs, hatched from eggs obtained in March 1974, were preserved in 70 percent isopropyl alcohol for drawings and descriptions. The interocular distance of these and field-collected nymphs was meas- ured using a calibrated ocular micrometer, and sex was determined when possible. Live nymphs were placed in styrofoam cups with creek water and kept in the environmental chamber at simulated creek temperature and photoperiod to determine size changes and to rear them to adults. After one sam- pling date in February 1976, 40 last instar nymphs were divided into three groups and kept at three constant temperatures, 7 C, 15 C, and 25 C, respectively, to determine the effect of temperature on the time re- quired for development and emergence. Size-frequency histograms, constructed from interocular distances of field-collected nymphs, and the methods of Cassie (1954) and Janetschek (1967) were used to inter- pret growth and instar development. To supplement these analyses and aid in inter- pretation, field-collected nymphs of differ- ent sizes were reared to successive instars. Food habits were determined by foregut analysis (Richardson and Gaufin 1971). For- age ratio (FR) (Hess and Swartz 1941) and electivity (E) (Ivlev 1961) were calculated for determination of food preferences. Stream temperature was taken on each sampling date with a total immersion ther- mometer. A seven-day continuous recording thermograph (Tempscribe remote bulb re- corder) was placed in the stream in June 1975 and left for the remainder of the study. Stream flow was taken on each sam- pling date from August 1975 to March 1976, using a Kahl Pygmy Flow Meter cali- brated at 0.4714 meters per revolution. Sea- sonal water samples were taken and ana- lyzed for pH, conductivity, and alkalinity according to standard methods (American Public Health Association 1971). Results and Discussion During the study, pH varied from 7.6 (summer 1975) to 8.1 (winter 1976), con- ductivity 530 iumhos (fall 1975) to 1,180 jumhos (summer 1975), and total alkalinity 212 ppm (fall 1975) to 408 ppm (winter 1976). Stream flow was highly variable, with the highest level observed in October- November 1974 and no flow in June-August 1974. From August 1975 to March 1976, stream flow remained relatively constant at ca. 1.4 m/sec. Temperature ranged from 30 C in June 1974 to 2 C in February 1976. Such highly variable and intermittent condi- tions of Clear Creek are typical of moder- ate-sized streams on the western edge of the temperate deciduous forest. Life cycle ad- justments of stoneflies such as H. crosbyi and Pcrlesta pkicida (Hagen) through natu- ral selection, which have enabled success in such harsh environments, were of prime in- terest in this study. Other dominant insects associated with H. crosbyi in Clear Creek included Isomjchia sicca manca (Eaton), Choroterpes (Neo- choroterpes) mexicanus (Allen), P. pkicida, Corijdalus cornutus (Linnaeus), Hijdropsijche simulans (Ross), Cheumatopsyche campyla (Ross) and C. lasia (Ross). Adult.— H. crosbyi displayed a relatively spontaneous emergence, with adults occur- ring from mid-February through March dur- ing the three-year study period. A similar recurrent emergence pattern has been re- 1977 Oberndorfer, Stewart: Stonefly Biology 263 ported for several stonefly species (Nebeker 1971b, Miiller 1973, Kerst and Anderson 1974). Average daily stream temperatures at the onset of emergence were ca. 15 C with 19 C maxima and 13 C minima. Emergence was essentially over and nymphs gone when average daily stream temperatures reached 22 C. As emergence progressed, there was a corresponding decline in nymphal popu- lations. These observations and results of laboratory holding of mature nymphs in en- vironmental chambers suggest that the mean daily stream temperature cue for emergence in H. croshiji is ca. 15 C. Field- collected, mature nymphs held at 7, 19, and 25 C showed a highly significant difference (P<0.01) in the mean number of days to emergence, with most time(X = 44.8 days) at 7 C, shortest successful emergence (X = 25.6 days) at 19 C, and 100 percent mortal- ity at 25 C. The emergence of some stonefly species is apparently dependent on the number of de- gree days or temperature accumulation dur- ing nvmphal development (Cummins 1974, Stanford 1975). Several investigators have indicated that events such as emergence and hatching result when a certain temperature cue is attained during periods of increasing or decreasing temperatures (Sheldon and Jewett 1967, Heiman and Knight 1970, Har- per and Pilon 1970, Nebeker 1971a, b, Rad- ford and Hartland-Rowe 1971, Sheldon 1972, Harper 197.3a, b). The emergence re- sponse range (tolerance) to such cues is nar- row in nymphs such as H. croshiji and oth- ers, effectively synchronizing emergence into a short time period (Macan 1958, Nebeker and Gaufin 1967, Nebeker 1971c, Hynes and Hynes 1975). Since accurate temperature data were not available over successive seasons, and a major egg-in- cubation experiment was lost due to envi- ronmental chamber malfunction, it could not be determined whether temperature ac- cumulation (Baskerville and Emin 1969) in- fluenced the development and emergence of H. croshiji. Collections of cast exuviae indicated that mature nymphs crawled out of the water onto rocks or debris above the surface of the water or along the bank for transforma- tion. Similar emergence has been reported for Paragneiina media (Walker) (Tarter and Krumholz 1971) and Isoperla transmarina (Newman) (Harper 1973a). Adults then ei- ther fly or walk to streamside vegetation or debris near the water. Preemergent nymph- al sex ratios were ca. 1:1, but collections of exuviae in the middle of the emergence pe- riod in March 1976 favored females, 1.0 cf :1.5 ? , indicating at least a slightly protandrous emergence. Schwarz (1970), Heiman and Knight (1970), Sheldon (1972), and Harper (1973a, b) have reported similar findings for other species. A seasonal size variation was exhibited in adult H. croshiji. The interocular distance decreased as emergence continued from a mean of 2.0 mm to 1.6 mm for females, and from 1.7 mm to 1.4 mm for males. Khoo (1968a), Schwarz (1970), Sheldon (1972), and Cather and Gaufin (1975) have shown similar variation, indicating that the emer- gence cue hurries smaller specimens so that full size is not attained (Khoo 1968a). Adults did not feed, but they were ob- served to drink water; Hynes (1976) suggested this is probably true of all adult Systellognatha. Dissection indicated that the digestive tract was much reduced. A period of two to five days is required for maturation of eggs in adult females. Iso- perla clio (Newman) (Harper 1973a), Iso- perla goertzi lilies, and Diiira hicaudata (Linnaeus) (Schwarz 1970) require a similar egg maturation period. Males find females by a tactile or possibly by a pheromone re- sponse, and mating takes place on stream- side vegetation or debris. Reproductive morphology, mating, and sperm transfer have been described bv Stewart and Stark (1977). Sperm transfer in this species is ex- ternal, and the aedeagus does not function as an intromittent organ. The species exhib- its polygamous mating habits. Mated fe- males flew to tree branches from 5 to 10 m above the stream on ca. the fifth day after emergence. An egg mass was extruded from the genital opening located at the rear of the abdominal sternum 8. Between ca. one to one-half hour before sunset, the female began her oviposition flight, approaching the site from upstream. She glided down 264 Great Basin Naturalist Vol. 37, No. 2 and rested briefly on smooth water from 2 to 10 m from the head of a riffle. Contact with water and a slight movement of the abdomen resulted in separation of the mass, which then sank. The female was carried by the current ca. 1 m during this process, then rose off the water, flying back into adjacent trees. Laboratory-held females extruded up to two more egg masses over the next three to five days. Mean life span of mated adult males and females held in the laboratory at simulated light and temperature conditions was 11.4 and 11.6 days, respectively. Mean numbers of eggs in successive mas- ses for reared females were 330, 235, and 172, respectively. Fecundity of dissected, gravid females was from 442 to 1,418 eggs/ $ , with a mean of 787/ ? . Smaller females generally contained fewer eggs. Deposited eggs had the highest caloric value (6.21 cal/mg) of any stage in the life cycle (Table 1). Mean caloric value of ca. 6.0 cal/mg ash-free for late instar H. crosbiji nymphs generally agree with those of Cummins and Wuycheck (1971) for im- matures of various aquatic insects, McDif- fett (1970) for Pteronarcys scotti, Lawton (1970, 1971) for Pyrrhosoma mjmphula and its prey, Hofsvang (1973) for Tipula excisa, and Brown (1974) for Corydalus cornutus. Mean ash-free cal/mg of males were signifi- cantly different from females in all stages (Student's t; P<0.05). Few calories (16.34) appear to be lost in transformation, through loss of the exuvium (mostly epicuticle). Males lose 33.8 percent (27.13) of their ca- loric pool as metabolic heat through their ca. 12 days of life (Table 1). A mean of 58.99 calories (deposited egg masses) constituted the energy allocated to reproduction (Pr) by females. However, a difference of 110.45 calories between newly emerged virgin females with ova and newly emerged virgin females with ova removed by dissection indicated that 51.46 calories might be resorbed by the female from un- deposited eggs to be used for maintenance and metabolic costs involved in mating and oviposition. Energy allocated to growth (Pg) was estimated as the mean biocontent of a newly emerged virgin female with ova re- moved (106.95 calories), since the first instar was very small and its calories negligible. An instantaneous reproductive effort (IRE) was thus calculated as: Pr (58.99)/[Pr (58.99) -I- Pg (106.95)] = 35.5 percent. An- other measure of IRE is the ratio of Pr to female carcass calories. Expressed this way, IRE of H. crosbyi was: Pr (58.99) /female carcass calories (106.95) = 55.1 percent. These values compare favorably with RE's determined by Brown (1974) for C. com- Table 1. Caloric values of major stages in the life cycle of H. Crosbyi. Material X dry wt. X cal/mg N° X Biocontent (mg) Ash-free (cal) Male, last instar 19.4 6.01 10 116.59 Female, last instar 36.0 5.95 10 214.20 Last instar exuviae 3.0 5.43 10 16.34 Male, newly emerged 14.2 5.66 10 80.37 (virgin) Female, newly emerged 36.6 5.94 11 217.40 (virgin with ova) Female, newly emerged 18.6 5.75 7 106.95 (virgin— ova removed) Male, spent (postmated) 10.2 5.22 8 53.24 Female, spent 18.7 4.93 8 92.19 (after oviposition) Egg masses 9.5 6.21 2 58.99 (total deposited/female) number of determinations June 1977 Oberndorfer, Stewart: Stonefly Biology 265 titiis. A later discussion of food habits of H. crosbyi will show that its general food re- sources are similar to those of C. cornutus. Females lose 57.6 percent (125.21) of their caloric pool as metabolic heat and Pr dur- ing their adult life. Egg.— Eggs are oval shaped, triangular in cross section, brown in color, and an aver- age 400 jum wide by 535 jum long (Fig. 1). They are similar to descriptions of other perlodid eggs such as Hydroperla fugitans (Frison 1935), Isogenoides frontalis and Isogenoides zionensis (Knight et al. 1965), Perlodes microcephala and Diura hicaudata (Schwarz 1970), and Pictetiella expansa and Isogenoides zionensis (Baumann 1973). The chorion surface generally lacks the sculptur- ing so evident in perlids (Stark and Gaufin 1976a, b), the Isoperlinae, and other stone- flies. The crown ridge is reduced and from two to four micropyles occur near the middle on each of the three sides of the egg- Each egg was enclosed by a gelatinous envelope, whose possible functions include: (1) prevention of desiccation after extrusion and prior to deposition, (2) action as a ce- menting substance to retain eggs in masses, and (3) attachment of individual eggs to substrate near the site of oviposition (Brinck Fig. 1. Hydroperla crosbyi egg; SEM, 400x. 1949, Hynes 1970). A gelatinous mushroom- shaped attachment structure, emerging from the crown area of the egg, was prominent for at least the first 48 hours after depos- ition. Its "carapace" portion appeared as a shriveled mass in older eggs in water and those preserved in alcohol (Fig. 1). Both the envelope and the attachment mushroom were very effective in gluing eggs to glass or other substrate. Eggs of H. crosbyi underwent a long, ca. seven-month diapause from February- March, when deposited, to October-Novem- ber, when fall rains occurred and mean daily water temperature declined to ca. 18 C. Elutriation (Stewart 1975) of sand sub- strate from an area where there was no sur- face flow in July 1974 yielded one H. crosbyi egg and several Perlesta placida eggs that appeared turgid and alive. The dia- pause and development of eggs were similar to those of Diura bicaudata and Brachyp- tera risi (Khoo 1968b, c) and Arnphinemura delosa, A. Undo, A. nigritta, and Prostoia completa (Harper 1973b), in that eyespots appeared only two to three weeks before hatching. One group of March 1974 eggs held at 25 ±2 C until September, when the temper- ature was reduced 1 C every three to five days, hatched spontaneously at 19 C. This and early November appearance of first in- star nymphs in Clear Creek suggest that rejuvenation of streams by fall rains and de- clining temperatures cue the termination of diapause. Another group of 2,000 March 1976 eggs, held in the laboratory at a con- stant 23 ± 1 C, hatched over an extended time period of five to eight months. The diapause is a great advantage since it allows survival through hot summer, when water temperatures might be lethal to yoimg nymphs and many of the intermittent streams inhabited by this species become dry. Several species such as Zealeuctra ar- noldi, Z. claasseni, and Z. hitei and Perlesta placida that are endemic to, or extend their ranges into, the ecotone between the tem- perate deciduous forest and grassland are apparently capable of similar fast cycles. This adaptation is similar to several species reported by Hynes and Hynes (1976) in 266 Great Basin Naturalist Vol. 37, No. 2 semiarid regions of Australia. Further work on their life cycles should provide some in- teresting comparisons with fauna of similar regions of the two continents. Many stone- flies undergo an egg or nymphal diapause to avoid adverse effects of extended cold or hot temperatures (Hartland-Rowe 1964, Khoo 1968a, b, c, Harper and Hynes 1970, 1972, Harper 1973a). Eclosion (Fig. 2) follows the same general pattern reported by Heiman and Knight (1970) and Vaught and Stewart (1974); the chorion is torn, and the nymph pushes the hinged cap away and crawls out. Hatching in the field is apparently synchronous, since first instar nymphs in November 1975 were found only over a four-week period. Macan (1958) postulated that synchronous hatching is a disadvantage since competition for food and space would be very intense. This would only apply, however, in situations where numbers approach or exceed the car- rying capacity. This theoretical asymptote (K) has not been defined for stream systems, where it would be expected to vary geo- graphically and from year to year. Also, drift might play a large role in dispersing synchronously hatched nymphs. Although both H. crosbyi and Perlesta placida occupy similar habitats, have similar diets, undergo an over-summer diapause, and have a relatively synchronous egg hatch (October-November), differential growth rates appear to separate them temporally. Nymphs of H. crosbyi grow faster through winter, leading to an emergence in Febru- ary-March at the time when early instars of P. placida are just beginning their growth spurt, leading to an April-May emergence. A thorough comparison can be made in southwestern streams such as Clear Creek, when another current study of the life cycle of P. placida is completed (Rosalyn Snellen, personal correspondence). Nymphs.— After hatching in October, nymphs grow rapidly through the winter until just prior to February-March emer- t t t t Fig. 2. Eclosion sequence; Htjdroperla croshiji. Fig. 3. Seasonal cycle and growth of Hydroperla crosbyi, March 1974' to March 1976, Clear Creek, Denton County, Texas: Vertical line = range, number at top of line = niunber measured, solid circle = mean, solid square and triangle within bars = mean female and male interocular distance, respectively, ver- tical bar =-!-&- ISD, arrow = emergence. June 1977 Oberndorfer, Stewart: Stonefly Biology 267 gence (Fig. 3), when water temperatures are coldest (Fig. 4). The species thus fits Hyne's (1961) classification as an F^ (fast) univoltine cycle. Similar cycles have been reported for Isoperia clio and Isogenus decisiis (Minshall and Minshall 1966); Atnphinemura delosa, Prostoia completa, and Shipsa rotunda (Har- per 1973b); and Capnia confusa, Utacapnia trava, U. columbiana, Doddsia occidentalism and Taenionema pacifica (Stanford 1975). This emphasizes the wide range of adapta- tion in Plecoptera nymphal growth charac- teristics since many species such as Zapada columbiana, (Hartland-Rowe 1964), Tae- niopteryx neuhulosa, Nemura meijeri, A. borealis, N. cinerea, Leuctra hippopus, Capnia atra, Diura nanseni, Isoperia dif- formis, I. grammatica (Svensson 1966), Isoperia goertzi, Diura bicaudata (Schwarz 1970), Paragnetina tyiedia (Heiman and Knight 1970), Pteronarcys dorsata, Isoperia frontalis (Nebeker 1971b), Brachyptera risi, Protonemoura meyeri, Amphinemura sulci- collis, A. borealis, A. standfussi, Capnia atra, Siphonoperla burrneisteri (Benedetto 1973), Neoperla clymene (Vaught and Stew- art 1974), Megarcys signata (Gather and Gaufin 1975), and Pteronarcella badia, Skwala parallela, Diura knowltoni, Isoperia 30 fulva, I. patricia, Sweltsa coloradensis, Su- wallia pallidula, and Zapada columbiana (Stanford 1975) exhibit slow cycles, with most growth occurring at moderate to warmest seasonal water temperatures. Tem- perature compensation (Gummins 1974) is thus manifested in this species through rap- id winter growth, early spring emergence, and diapause for protection against drying or lethal temperatures in summer. This growth pattern is a selective advantage in allowing avoidance of competition for food and space with such species as Corydalus cornutus (Brown 1974) and Perlesta placida. First instar nymphs were obtained from egg-hatching experiments and Glear Greek. They were unpigmented, had 9 antennal and 4 cereal segments, and the mean inter- ocular distance was 0.18 mm (Fig. 5). Hynes (1976) indicated that many first-instar Ple- coptera exhibit these characters. Distinctive color patterns became evident through the middle and later instars, and numbers of cereal and antennal segments in- creased to 30 and 50, respectively. Mouth- parts of first-instar nymphs are shown in Fig. 6. Life cycle studies of stoneflies should give attention to good descriptions of first instars to facilitate accurate identifications u 25 ^ 20 S 15 10 EMERGENCE - J J A S O N D J F M « 1975 ^ 1'^^^^^ ► Fig. 4. Mean weekly stream temperature, June 1975 to March 1976. 268 Great Basin Naturalist Vol. 37, No. 2 for future ecological studies. Comparison of Figs. 6 and 7 show the major changes that take place in presence and pattern of setal arrangement and shape of mouthparts be- tween first and last instars; number of pal- pal segments remain unchanged. Nymphs can first be sexed at the sixth in- star, by differences in the posterior setal ar- rangement of the eighth abdominal sternum. The fringe is interrupted in females and continuous in males. Dimorphism in setal arrangement, nymphal size, and even even- tual presence of the male epiproct, becomes more distinct in postsixth instars. The sex ratio of 870 postfifth instar nymphs sexed throughout the study was 1.0 j* :1.1 9 . The larger size of female nymphs appears typical of all stoneflies and has been report- ed by Holdsworth (1941a, b), Heiman and Knight (1970), Tarter and Krumholz (1971), Brittain (1973), and Vaught and Stewart (1974). Estimation of nymphal instar number was determined from 971 field-collected nymphs by using a size-frequency histogram (Fig. 8) and the methods of Cassie (1954) and Ja- netschek (1967) (Figs. 9 and 10, respective- ly). An expanded discussion of application of these methods is given by Harper (1973b) and McClure and Stewart (1976). These methods indicated 12 and up to 14 instars for males and females, respectively. Similar dimorphism in instar number has been re- ported by Holdsworth (1941b), McDiffett (1970), and Vaught and Stewart (1974). The size change between most instars was con- firmed and these estimates substantiated by rearing field-collected nymphs of various sizes through at least one molt. Of course, 0.5 mm Fig. 5. Hydroperla crosbyi first instar nymph. 0.1 mnn Fig. 6. Hydroperla crosbyi first instar nymphal mouthparts: A = labrum, B = mandible, C = max- illa, D = labium. June 1977 Oberndorfer, Stewart: Stonefly Biology absolute determination of instars can only be made through rearing individuals from egg to adult, under various physical condi- tions (Harper 1973b). Stanford (1975) sug- gested that temperature may influence the number of instars in hemimetabolous in- sects. Seasonal food habits of nymphs are shown in Table 2. All calculations are based on numbers of dietary items in the foregut and environment. Gut contents were examined using methods of Richardson and Gaufin (1971), and food availability was calculated from quantitative samples. Larvae of Sim- uliidae and Chironomidae always made up over 85 percent of the diet (Table 2), and electivity (Ivlev 1961) and forage ratio (Hess and Swartz 1941) consistently in- dicated that these organisms were preferred or more available. Nymphs of Isonychia sic- ca manca were the third most common dietary item, but preference for them was always negative (or they were less available 1.0 mm Fig. 7. Hydroperla crosbyi last instar female nymph- al mouthparts: A = labrum, B = mandible, C = maxilla, D = labium. ^^'9 y 50 o LU m 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 2.2 2.4 INTEROCULAR DISTANCE (mm) Fig. 8. Interocular distance— frequency of 971 Hydroperla crosbyi nymphs, illustrating instars. 270 Great Basin Naturalist Vol. 37, No. 2 as 5.0 10 ■ >^^> 20 - 40 - 60 - - /.^2 80 - 90 - /''' 95 - /'' 99 0.2 OA 0.6 0.8 1£) 1.2 1.4 1£ 1£ 2.0 2.2 2.4 INTEROCULAR DISTANCE (mm) Fig. 9. Instar analysis of 971 Hydroperla croshiji nymphs by Cassie (1954) method. for feeding). Drusinus (Limnephilidae), Chi- marra (Philopotamidae), Perlesta (Perlidae), and unidentified aquatic Lepidoptera occa- sionally appeared in stomachs but were con- sidered of little importance. Prison (1935) reported large numbers of chironomids in the diet of H. croshiji nymphs. Similar pref- erences for dipteran larvae have been re- ported for carnivorous insects (Minshall and Minshall 1966, Tarter and Krumholz 1971, Sheldon 1972, Stewart et al. 1973, Vaught and Stewart 1974, and Gather and Gaufin 1975). Simuliids were predominant to chi- ronomids in the diet from November- Janu- ary, with electivity or preference greater until January (Table 2). A shift in dietary occurrence became evident in February, with the two components occurring in equal proportions, then simuliids again pre- dominated in March; electivity for sim- Table 2. Seasonal food habits of 110 Hydroperla croshiji nymphs November 1975 to March 1976, Clear Creek, Denton County, Texas. Food organisms S S % of Diet 17.2 72.4 10.3 Nov. % of Env. 9.2 6.6 17.6 1975 E 0.30 0.83 0.26 FR 1.87 10.97 0.58 % of Diet 13.6 84.1 1.1 Dec. % of Env. 8.3 16.4 15.0 1975 E 0.24 0.67 0.86 FR 1.64 5.13 0.07 % of Diet 21.0 77.2 1.7 Jan. % of Env. 6.9 42.1 10.0 1976 E 0.,50 0.29 0.71 FR 3.04 1.83 0.17 % of Diet 42.8 42.8 8.3 Feb. % of Env. 22.9 17.2 13.2 1976 E 0.30 0.43 -0.23 FR 1.87 2.49 0.63 % of Diet 28.2 70.0 Mar. % of Env. 25.0 9.5 1976 E 0.06 0.76 FR 1.13 7.37 3.7 0.0 16.0 11.1 13.3 21.0 25 27 30 June 1977 Oberndorfer, Stewart: Stonefly Biology 271 uliids was greater in both these months even though in February their numbers were de- creasing because of emergence. Proportion of empty stomachs was highest just prior to emergence (Table 2). Molting nymphs al- ways had empty stomachs. Similar findings were reported by Hanson (1960), Tarter and Krumholz (1971), and Gather and Gaufin (1975). Fat stores are probably relied on for necessary energy from one to five days prior to emergence and for transformation. Finni (1975) reported preemergent fat reserves for AUocapnia granulata. Mean nymphal standing crops were 8.0, U CO o UJ DQ 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 2.2 2.4 Fig. five of INTEROCULAR DISTANCE (mm) 10. Instar analysis of 971 Hijdropcrla crosbiji nymphs by Janetschek (1967) method: A = running the frequencies, B = modal periodicity of instars (difference between Fig. 8 histogram and A). mean of 272 Great Basin Naturalist Vol. 37, No. 2 25.6, 40.0, and 21.3/m- each month from December 1975 to March 1976. The lower number in December was probably due to sampling error, peculiarities in micro- distribution, or difficulty in detection of tiny first-instar nymphs. Decreases from Febru- ary-March were likely an indication of sur- vivorship and emergence. Literature Cited American Public Health Association, Inc. 1971. Standard methods for the examination of water and wastewater. Amer. Publ. Health A.ssn., Inc., New York. 874 pp. Baskerville, G. L., and P. Emin. 1969. Rapid estima- tion of heat accumulation from maximum and minimum temperatures. Ecology 50:514-517. Baumann, R. W. 1973. Studies on Utah stoneflies (Ple- coptera). Great Basin Nat. .33:91-108. Benedetto, L. A. 1973. Growth of stonefly nymphs in Swedish Lapland. Entomol. Tidskr. 94:15-19. Brinck, p. 1949. Studies on Swedish stoneflies (Plecop- tera). Opusc. Entomol., Supplementum 11:1-250. Brittain, J. E. 1973. The biology and life cycle of Ne- moura avicularis Morton (Plecoptera). Freshwat. Biol. 3:199-210. Brown, A. V. 1974. Ecological energetics of the dob- son fly, Conjdahis cornuttis. Ph.D. dissertation, North Texas State Univ. Cassie, R. M. 1954. Some uses of probability paper in the analysis of size frequency distributions. Aust. J. Mar. Freshwater Res. 5:513-522. C.-vTHER, M. R., and a. R. Gaufin. 1975. Life history and ecology of Megarcijs signata (Plecoptera: Per- lodidae). Mill Creek, Wasatch Mountains, Utah. Great Basin Nat. 35:39-48. Claassen, p. W. 1931. Plecoptera nymphs of America (north of Mexico.) Thomas Say Found. 3:1-199. Cummins, K. W. 1974. Structure and function of stream ecosystems. BioScience 24:631-641. Cummins, K. W., and J. C. Wuycheck. 1971. Caloric equivalents for investigations in ecological ener- getics. Mitt. Int. Ver. Limnol. Mitteilungen Com- munications No. 18:1-158. FiNNi, G. R. 1975. Feeding and longevity of the winter stonefly, Allocapnia granulatu (Claassen) (Plecop- tera: Capniidae). Ann. Entomol. Soc. Amer. 68:207-208. Prison, T. H. 1929. Fall and winter stoneflies, or Ple- coptera of Illinois. Illinois Nat. Hist. Surv. Bull. 18:.340-4()9. 1935. The stoneflies, or Plecoptera of Illinois. Illinois Nat. Hist. Surv. Bull. 20:281-471. 1942. Studies of North American Plecoptera, with special reference to the fauna of Illinois. Il- linois Nat. Hist. Surv. Bull. 22:231-355. Hanson, J. F. 1960. Visible changes preceding molting in Plecoptera. Verb. XI. Int. Kongr. Entomol. Wein. 3:271. Harper, P. P. 1973a. Emergence, reproduction, and growth of setipalpian Plecoptera in southern On- tario. Oikos 24:94-107. 1973b. Life histories of Nemouridae and Leuctridae in southern Ontario (Plecoptera). Hy- drobiologia 41:309-356. Harper, P. P., and H. B. N. Hynes. 1970. Diapause in the nymphs of Canadian winter stoneflies. Ecol- ogy 5i:925-927. 1972. Life histories of Capniidae and Taeniop- terygidae (Plecoptera) in southern Ontario. Arch. Hydrobiol., Suppl. 40:274-314. Harper, P. P., and J. G. Pilon. 1970. Annual patterns of emergence of some Quebec stoneflies (Insecta: Plecoptera). Can. J. Zool. 48:681-694. Hartland-Rowe, R. 1964. Factors influencing the life histories of some stream insects in Alberta. Verb. Int. Ver. Limnol. 15:917-925. Heiman, D. R., and a. W. Knight. 1970. Studies on the growth and development of the stonefly Para- gnetina media Walker (Plecoptera: Perlidae). Amer. Mid. Nat. 84:274-278. Hess, A. D., and A. Swartz. 1941. The forage ratio and its use in determining the food grade of streams. Trans. 5th N. Amer. Wildlife Conf. 1940:162-164. HoFsvANG, T. 1973. Energy flow in Tipula excisa Sebum. (Diptera: Tipulidae) in a high mountain area, Finse, South Norway. Norw. J. Zool. 21:7- 16. Holdsworth, R. p., Jr. 1941a. The life history and growth of Pteronarcys proteus Newman (Pteronar- cidae: Plecoptera). Ann. Entomol. Soc. Amer. 34:495-502. 1941b. Additional information and a correction concerning the growth of Pteronarcys proteus Newman (Pteronarcidae: Plecoptera). Ann. Ento- mol. Soc. Amer. 34:714-715. Hynes, H. B. N. 1961. The invertebrate fauna of a Welsh mountain stream. Arch. Hydrobiol. 57:344- 388. 1970. The ecology of running waters. Univer- sity of Toronto Press, Toronto. 555 p. 1976. Biology of Plecoptera. Ann. Rev. Ento- mol. 21:135-153. Hynes, H. B. N., and M. E. Hynes. 1975. The life his- tories of many of the stoneflies (Plecoptera) of southeastern mainland Australia. Aust. J. Mar. Freshwat. Res. 26:11.3-153. Ivlev, V. S. 1961. Experimental ecology of the feeding of fishes. Yale University Press, New Haven and London. .302 pp. Janetschek, H. 1976. Growth and maturity of the springtail Gomphiocephalus hodgsoni Carpenter, from South Victoria Land and Ross Island. Antarc- tic Res. Serv. 10 Entomology of Antarctica, pp. 295-305. Kerst, C. D., and H. H. Anderson. 1974. Emergence patterns of Plecoptera in a stream in Oregon, U.S.A. Freshwat. Biol. 4:205-212. Khoo, S. G. 1968a. Experimental studies on diapause in stoneflies. I. Nymphs of Capnia bifrons (New- June 1977 Oberndorfer, Stewart: Stonefly Biology 273 man). Proc. Roy- Entomol. Soc. London Ser. A. 43:40-48. 1968b. Experimental studies on diapause in stoneflies. II. Eggs of Ditira Bicaudata (L.). Proc. Roy. Entomo!. Soc. London Ser. A. 43:49-56. 1968c. Experimental studies on diapause in stoneflies. III. Eggs of Brachyptera risi (Morton). Proc. Roy. Entomol. Soc. London Ser. A. 43:141- 146. Knight, A. W., A. V. Nebeker, and A. R. Gaufin. 1965. Further descriptions of the eggs of Plecop- tera of Western United States. Entomol. News. 76:233-239. Lawton, J. H. 1970. Feeding and food energy assimila- tion in larvae of the damselfly Pyrrhosoma nym- phula (Sulzer) (Odonata: Zygoptera). J. Anim. Ecol. 39:669-689. 1971. Ecological energetics studies on larvae of the damselfly Pyrrhosoma nymphula (Sulzer) (Odonata: Zygoptera). J. Anim. Ecol. 40:385-423. Macan, T. T. 1958. Causes and effects of short emer- gence periods in insects. Verh. Int. Ver. Limnol. 13:845-849. McClure, R. G., and K. W. Stewart. 1976. Life cycle and producion of the mayfly Choroterpes (Neochoroterpes) rnexicatius Allen (Ephemerop- tera: Leptophlebiidae). Ann. Entomol. Soc. Amer. 69:134-144. McDiFFETT, W. F. 1970. The transformation of energy by a stream detritivore, Pteronarcys scotti (Ple- coptera). Ecology 51:975-988. MiNSHALL, G. W., AND J. N. MiNSHALL. 1966. Notes on the life history and ecology of Isoperla clio (New- man) and Isogeniis decisus Walker (Plecoptera: Perlodidae). Amer. Mid. Nat. 76:340-350. MtJLLER, K. 1973. Life cycles of stream insects. Aquilo Ser. Zool. 14:105-112. Nebeker, A. V. 1971a. Effect of temperature at differ- ent altitudes on the emergence of aquatic insects from a single stream. J. Kans. Entomol. Soc. 44:26-35. 1971b. Effect of high winter water temper- atures on adult emergence of aquatic injects. Wa- ter Research. Pergamon Press. 5:777-783. 1971c. Effect of water temperature on nymph- al feeding rate, emergence, and adult longevity of the stonefly Pteronarcys dorsata. J. Kans. Ento- mol. Soc. 44:21-26. Nebeker, A. V., and A. R. Gaufin. 1967. Factors af- fecting wing length and emergence in the winter stonefly Capnia nana. Entomol. News. 78:85-92. Needham, J. C., AND P. W. Claassen. 1925. Plecop- tera or stoneflies of America north of Mexico. Thomas Say Found., Lafayette, Indiana. .397 pp. Phillipson, H. 1964. A miniature bomb calorimeter for small biological samples. Oikos 15:130-139. Radford, D. S., and R. Hartland-Rowe. 1971a. Emergence patterns of some Plecoptera in two mountain streams in Alberta, Can. J. Zool. 49:657-662. 1971b. The life cycles of some stream insects (Ephemeroptera and Plecoptera) in Alberta. Can. Entomol. 103:609-617. Richardson, J. W., and A. R. Gaufin. 1971. Food habits of some western stonefly nymphs. Trans. Amer. Entomol. Soc. 97:91-121. RiCKER, W. E. 1952. Systematic studies in Plecoptera. Indiana University Publications, Science Series No. 18. 200 pp. ScHWARZ, P. 1970. Autokologische Untersuchungen zum Lebenszyklus von Setipalpia-Arten (Plecop- tera). Arch. Hydrobiol. 67:103-140. Sheldon, A. L. 1972. Comparative ecology of Arcy- nopteryx and Diura (Plecoptera) in a California stream". Arch. Hydrobiol. 69:521-546. Sheldon, A. L., and S. G. Jewett, Jr. 1967. Stonefly emergence in a Sierra Nevada stream. Pan-Pac. Entomol. 43:1-8. Stanford, J. A. 1975. Ecological studies of Plecoptera in the upper Flathead and Tobacco rivers, Mon- tana. Ph.D. dissertation. University of Utah. 241 pp. Stark, B. P., and A. R. Gaufin. 1976a. The Nearactic species of Acroneuria (Plecoptera: Perlidae). J. Kans. Entomol. Soc. 49:221-253. 1976b. The Nearactic genera of Perlidae (Ple- coptera). Misc. Pub. Entomol. Soc. Amer. 10:1- 77. Stewart, K. W. 1975. An improved elutriator for sep- arating stream insects from stony substrates. Trans. Amer. Fish. Soc. 104:821-823. Stewart, K. W., G. L. Atmar, and B. M. Solon. 1969. Reproductive morphology and mating be- havior of Perlesta placida (Plecoptera: Perlidae). Ann. Entomol. Soc. Amer. 62:1433-1438. Stewart, K. W., R. W. Baumann, and B. P. Stark. 1974. The distribution and past dispersal of southwestern United States Plecoptera. Trans. Amer. Entomol. Soc. 99:507-546. Stewart, K. W., G. P. Friday, and R. E. Rhame. 1973. Food habits of hellgrammite larvae, Cory- dahis corniitits (Megaloptera: Corydalidae), in the Brazos River, Texas. Ann. Entomol. Soc. Amer. 66:959-963. Stewart, K. W., and B. P. Stark. 1977. Reproductive system and mating of Hydroperia crosbyi (Need- ham and Claassen); a newly discovered method of sperm transfer in Insecta. Oikos 28:84-89. SvENSsoN, P. 1966. Growth of nymphs of stream living stoneflies (Plecoptera) in northern Sweden. Oikos 17:197-206. Tarter, D. C., and L. A. Krumholz. 1971. Life his- tory and ecology of Paragnetina media (Walker) (Insecta: Plecoptera) in Doe Run, Meade County, Kentucky. Amer. Mid. Nat. 86:169-180. Vaught, G. L., and K. W. Stewart. 1974. Life history and ecology of the stonefly Neoperla clymene (Newman) (Plecoptera: Perlidae). Ann. Entomol. Soc. Amer. 67:167-178. Wu, C. G. 1923. Morphology, anatomy, and ethology of Nemttra. Bull. Lloyd Libr. 23:1-81. SPRING WATERFOWL MIGRATION IN THE UINTA BASIN OF NORTHEASTERN UTAH Mary E. Sangster' Abstract.— Spring waterfowl migrations at Ouray National Wildlife Refuge and Pelican Lake, Uintah County, Utah, were monitored during 1975 and 1976. Peak numbers occurred during the first week in April 1975 when 11,440 birds were counted. In 1976 the highest count, 9,650 birds, was made on 15 March. Most numerous early- season migrants were Mallards and Pintails. Important midseason migrants included Blue-winged and Cinnamon Teal, Canvasbacks, and Redheads. Late season migrants were Gadwall and Ruddy Ducks. Surveys conducted only at Ouray Refuge or at Pelican Lake would not adequately have reflected species composition of spring migration. Results point up the advisability of sampling large open water areas to obtain a representative view of waterfowl species composition in a given area during spring migration. The Unita Basin, situated in northeastern Utah, is a structural depression lying be- tween the Uinta Mountains to the north and the Tavaputs Plateau to the south. The ba- sin is dissected by many streams and guUeys and the topography is broken and rough. Climate of this area is arid, with light snowfall and an average precipitation of less than 18 cm. Winters in the Uinta Basin are long and severe, with temperatures often falling below —18° C. Summers are short and cool and, although temperatures of 38° C are common, such temperatures do not last for prolonged periods. The Green River transects the Uinta Ba- sin from north to south and is the master stream for the entire drainage system of the basin. The Green River has long been known to serve as an important migratory route for waterfowl. From the south the main migratory routes follow the Colorado River to its junction with the Green. Water- fowl continue along the Green River through the Book Cliffs and the Uinta Mountains and then into Wyoming. To the west, the Provo River provides a passage from the western section of the Uinta Basin. Low passes through the Wasatch Mountains create routes into the Great Salt Lake drainage system (Twomey 1942). In his paper on Colorado River Basin birds. Hay ward (1967) stated that migrating waterfowl formerly depended almost entire- ly on the Green River and its tributaries for resting and feeding. More recently. Hay- ward noted, reservoirs constructed to pro- duce electric power or for storage of irriga- tion waters have become important areas for migrating birds. The purpose of this paper is to present information on the species and numbers of waterfowl migrating through the Uinta Ba- sin and the chronology of spring migration in 1975 and 1976. Waterfowl counts were conducted at weekly intervals in 1975 and at 10-day intervals in 1976 from 5 March through 17 May. Waterfowl counts were made on Ouray National Wildlife Refuge in the southeast portion of the Uinta Basin. The 4,860 ha refuge is situated astride the Green River for 12 km. There are six sepa- rate bottoms along the length of the refuge which have been formed by ancient oxbows in the river. A system of shallow ponds and marshes has been constructed in these bot- toms, and water pumped from the river fills these impoundments in the spring and fall. Vegetative zones on the refuge are com- ponents of the Mixed Desert Shrub Commu- nity (Twomey 1942). Waterfowl censuses were also made at Pelican Lake, a reservoir designed for irrigation of adjacent land.s, lo- cated 1.6 km west of the refuge. This paper documents in detail the use of this newly Gavlord Memorial Labo 274 June 1977 Sangster: Waterfowl Migration 275 created reservoir and Ouray Refuge im- poundments by waterfowl of the Uinta Ba- sin. The migration data presented here were collected as part of a larger study of ecol- ogy of waterfowl on Ouray National Wild- life Refuge (Sangster 1976). The study was supported by the Gaylord Memorial Labo- ratory (School of Forestry, Fisheries and Wildlife, University of Missouri, cooperating with the Missouri Department of Con- servation) and by the Missouri Cooperative Wildlife Research Unit (U.S. Fish and Wild- life Service, Wildlife Management Institute, Missouri Department of Conservation, and the University of Missouri— Columbia coop- erating). The author was an Edward K. Love Fellow. I thank Leigh H. Fredrickson for guidance and criticism during this study and Thomas S. Baskett for critically review- ing this manuscript. Results A few Canada Geese- and Mallards winter along the Green River as long as some open water is present. As more open water be- comes available in March, ducks and geese begin moving into the Basin. In 1975 open water appeared at the edges of Pelican Lake the last week in February, and on 5 March a concentration of 2,500 ducks, mostly Mallards and Pintails, marked the be- ginning of spring migration. On 5 March 1975, pumps started moving water into im- poundments on the refuge, and a week later over 2,000 birds were recorded using the refuge. In 1976 open water conditions were considerably different in the Uinta Basin due to prolonged cold weather. Pelican Lake was frozen except for a few small openings at the edge until 1 April, and ref- uge pumps did not begin filling the im- poundments until April. Most of the early migrants in March 1976 were recorded on the Green River. The spring migration peak in 1975 oc- curred during the first week in April when 11,440 birds were counted. In 1976 the All scientific names of waterfowl according to the An Its' Union (1957, 1973) are presented in Table 1. highest count, 9,650 birds, was made on 15 March. Early migrants.— Early migrants were Mallards and Pintails, which reached peak numbers during the fourth week in March 1975 and two weeks earlier in 1976 (see Table 1 for peak migration numbers and dates for all species). The peak for Ameri- can Green-winged Teal was recorded the last week in March in both years, although in 1975 a second high was witnessed the second week in April. The American Wi- geon, another early migrant in 1975, was most common during the third week in March. In 1976 peak numbers of wigeon occurred the second week in May. In 1975 over 75 percent of the wigeon observed were on Pelican Lake. The fact that Pelican Lake was frozen until April the following year may account for the high numbers of wigeon observed later in the season in 1976. MiDSEASON MIGRANTS.— Midseason mi- grants included Blue-winged Teal, Cinna- mon Teal, Canvasback, Redhead, Buffle- head, and Shoveler. Small numbers of Lesser Scaup, Common Goldeneye, Ring- necked Ducks, and Common Merganser also were present at midseason. In 1975 peak numbers of Cinnamon Teal, Blue-winged Teal, Canvasback, Bufflehead, Ring-necked Ducks, and Common Mer- ganser were recorded during the second week in April and the Common Goldeneye during the third week of April. In 1976 Blue-winged Teal peaked during the same period but the peaks for the other species varied. The highest numbers of Bufflehead, Ring-necked Ducks, and Common Gold- eneye were recorded the fourth week in March and Canvasback and Common Mer- ganser were most common the first week in April. Cinnamon Teal numbers remained low until 1 May 1976. Redhead and Lesser Scaup populations were highest during the third week in April 1975. In 1976 Lesser Scaup peaked at the same period, but peak numbers of Redheads occurred the first week in April. In 1975 Shoveler populations showed two peaks, one the third week in April and the other the first week in May. The first week in May 276 Great Basin Naturalist Vol. 37, No. 2 was also a peak period for Shoveler popu- lations in 1976. Late-season migrants.— The Gadwall and Ruddy Duck were late migrants to the Uinta Basin. The Gadwall and Ruddy Duck reached peak numbers the fourth week in April 1975 and the first week in May 1976. Some Gadwall were in the area as early as the end of March but the Ruddy Duck did not appear until the second week in April. Over 85 percent of all Ruddy Ducks were observed at Pelican Lake. In 1975 a total of 1,610 Ruddy Ducks were counted, but in 1976 only 178 were observed (Tables 2 and 3). Although there were two more census periods in 1975 than in 1976, this did not appear to account for the great differences in observed niunbers of migrating Ruddy Ducks through the Uinta Basin. Canada Geese.— Large flocks of Canada Geese do not appear to use the Green River as a migration route. In 1975 the highest concentration of geese, 265 birds, occurred on 14 March. On 15 March 1976 a peak number of 500 geese was recorded. Other weekly census figures fluctuated between 100 and 160 geese. I believe these counts represent resident geese along the Green River. Refuge records (1965-1972) show that the average peak concentrations of geese in the spring has been 180 birds. Data for 1975 and 1976 appear to be consistent with refuge records. Band recovery information for Canada Geese banded and released at Ouray Refuge in March 1965 yielded information on the movements of 25 birds. Of the 15 birds re- covered, 12 were recovered in the Uinta Basin and 3 along the lower Colorado Riv- er. Reports on 10 birds came from retrap- ping information at Wheatland Reservoir in southeast Wyoming. Records for geese trapped at Wheatland Reservoir during June banding operations indicate that birds using the Ouray Refuge may move to Wyoming reservoirs to molt (unpublished records, Ouray National Wildlife Refuge). Trautin and Low (1975) proposed that the harvest of geese in northeastern Utah is de- pendent upon out-of-state-produced birds because the ratio of out-of-state-banded birds to Utah-banded birds is 9.00:1.00. Most of the birds banded out of state have come from Wyoming, where they were banded as molting adults. The production areas that these geese came from is un- Table 1. Peak numbers and the dates recorded for migrating waterfowl in 1975 and 1976. Data for Ouray Na- tional Wildlife Refuge and Pelican Lake are combined. Species 1975 Peak Number Date 1976 Peak Number Date Mallard (Anas platyrhynchos) Pintail {Anas acuta) Gadwall (Anas strepera) Shoveler (Anas clypeata) American Green-winged Teal (Ancvi crecca) Cinnamon Teal (Anas cyanoptera) Ruddy Duck (Oxyiira jarnaicensis) Canada Goose (Branta canadensis) Redhead (Aythya arnericana) American Wigeon {Anas arnericana) Bufflehead {BttcephaUt albeola) Lesser Scaup (Aythya affinis) Common Merganser (Mergus merganser) Blue-winged Teal (Anas discors) C^anvasback (Aythya valisineria) Ring-necked Duck (Aythya collaris) Common Goldeneye (Bucephala clangula) Snow Goose (Chen cuendescens) Whistling Swan (Olor columbianus) 3,725 3/28 4,750 3/15 4,930 3/28 4,120 3/15 1,110 4/24 285 5/1 635 4/16 255 5/1 775 3/28 805 3/24 365 4/9 225 5/1 405 4/23 80 5/1 265 3/14 500 3/15 315 4/16 530 4/2 345 3/20 225 5/11 345 4/9 215 3/24 275 4/16 300 4/21 330 4/9 130 4/2 175 4/9 120 4/11 175 4/9 140 4/2 145 4/9 215 3/24 90 4/16 124 3/24 17 3/14 June 1977 Sangster: Waterfowl Migration 277 known, but the limited band recovery data from Ouray Refuge indicate that some of these out-of-state-banded birds may actually be geese produced in the Uinta Basin. Other species.— On 14 March 1975, 17 Snow Geese were sighted on Pelican Lake. Snow Geese are considered rare in the Uinta Basin (Hayward 1967). In 1976 100 Whistling Swans were re- corded during four census periods in March and April. Hayward (1967) lists this species as an uncommon transient in the upper Col- orado River Basin. American Coots {Fulica americana) were the most abundant migrant recorded in 1975 and 1976. Peak migration levels for coots in 1975 occurred the third and fourth weeks of April, with counts of over 6,000 birds each week. In 1976 peak numbers of coots were recorded the third week in April and the second week in May, with counts of approximately 4,500 birds. Species composition and local dis- tribution.—During spring migration, Mal- lards comprised 28.9 percent of the migra- tory population in 1975 and 33.0 percent in 1976; Pintails 25.2 percent and 27.2 per- cent; teal 10.8 percent and 13.3 percent; Gadwall 8.1 percent and 4.1 percent; Shovelers 6.3 percent and 2.5 percent; and Canada Geese 3.0 percent and 4.6 percent (Tables 2 and 3). Over 80 percent of the waterfowl recorded in 1975 and 1976 were dabbling ducks. Waterfowl use of river bottom marsh and reservoir habitat was nearly equal in 1975 but species use varied widely (Table 3). The open water of Pelican Lake was used exten- sively for feeding and resting by diving ducks, especially Canvasbacks, Redheads, Ruddy Ducks, Bufflehead, and American Goldeneye, as well as by wigeon and coots. In 1976 the reduced use of Pelican Lake by migrating waterfowl is readily apparent (Table 3). This can be partially explained by the frozen condition of the lake throughout March and high winds in April that made the lake less attractive to migrating water- fowl. These conditions should not have af- fected the numbers of late arriving migrants such as Gadwall, Ruddy Ducks, Shovelers, and Buffleheads. Apparently the riparian Table 2. Comparison of species composition at Ouray National Wildlife Refuge and Pelican Lake and species composition for Ouray NWR and Pelican Lake combined, spring 1975. Ouray NWR Pelican Lake Combined Species Number Percent of Total Number Percent of Total Number Percent of Total Mallard 7,560 33.7 6,155 24.7 13,715 28.9 Pintail 4,435 19.8 7,520 30.2 11,955 25.2 Gadwall L470 6.6 2„350 9.4 3,820 8.1 Shoveler L060 4.7 1,930 7.7 2,990 6.3 American Green- L960 7.7 575 2.3 2,535 5.4 winged Teal Cinnamon Teal L430 6.4 405 1.6 1,835 3.9 Ruddy Duck 210 0.9 1,430 5.8 1,640 3.5 Canada Goose 1,060 4.7 350 1.4 1,410 3.0 Redhead 465 2.1 880 3.5 1,345 2.8 American Wigeon 305 1.4 980 3.9 1,285 2.7 Bufflehead 310 1.4 810 3.3 1,120 2.4 Lesser Scaup 285 1.3 745 3.0 1,0.30 2.2 Common Merganser 970 4.3 35 0.1 1,005 2.1 Blue-winged Teal 560 2.5 140 0.6 700 1.5 Canvasback 105 0.5 300 1.2 405 0.8 Ring-necked Duck 230 1.0 150 0.6 380 0.8 Common Goldeneye 10 tr. 150 0.6 160 0.3 Snow Goose — 17 0.1 17 0.1 TOTAL 24,922 100.0 47,347 100.0 278 Great Basin Naturalist Vol. 37, No. 2 Table 3. Comparison of species composition at Ouray National Wildlife Refuge and Pelican Lake and species composition for Ouray NWR and Pelican Lake combined, spring 1976. Species Ouray NWR Pelican Lake Combined Number Percent Number Percent Number Percent of Total of Total of Total 8,775 38.9 275 5.6 9,050 33.0 6,810 30.2 665 13.5 7,475 27.2 885 3.9 240 4.9 1,125 4.1 505 2.2 170 3.5 675 2.5 1,685 7.5- 500 10.2 2,185 8.0 935 4.1 75 1.5 1,010 3.7 40 0.2 140 2.9 180 0.6 1,265 5.6 1 tr. 1,265 4.6 45 0.2 875 17.8 920 3.3 510 2.3 50 1.0 560 2.0 120 0.5 380 7.7 500 1.8 20 0.1 670 1.3.6 690 2.5 335 1.5 25 0.5 360 1.3 405 1.8 50 1.0 455 1.6 55 0.2 240 4.9 295 1.1 80 0.4 345 7.0 425 1.5 215 4.4 215 0.8 100 0.4 ... ... 100 0.4 Mallard Pintail Gadwall Shoveler American Green- winged Teal Cinnamon Teal Ruddy Duck Canada Goose Redhead American Wigeon Bufflehead Lesser Scaup Common Merganser Blue-winged Teal Canvasback Ring-necked Duck Common Goldeneye Whistling Swan TOTAL 22,570 100.0 4,916 100.0 27,485 habitat of Ouray Refuge compensated for the lack of open water on Pelican Lake for the early migrants. Mallard and pintail peaks were similar both years, but the dis- tribution between the refuge and Pelican Lake varied (Tables I, 2, and 3). A migration survey only at Ouray Refuge or only at Pelican Lake would not have presented a complete picture of the species composition of the spring migration. These differences in waterfowl species distribution indicate the advisability of sampling large, open water areas to obtain the species com- position of the migration population for any given area. LITERATURE CITED American Ornithologists' Union. 1957. Check-list of North American birds, 5th ed. Baltimore, Amer. Omithol. Union, xiii & 691 pp. 1973. Thirty-second supplement to the American Ornithologists' Union check-list of North American birds. Auk 90:411-419. Hayward, C. L. 1967. Birds of the upper Colorado River Basin. Brigham Young Univ. Sci. Bull., Biol. Ser. 9 (2): 1-63. Sangster, M. E. 1976. Migration and ecology of wa- terfowl on Ouray National Wildlife Refuge, Utah. M.S. thesis. University of Mo., Columbia, v & 91 pp. Tautin, J., AND J. B. Low. 1975. Harvest trends of Canada Geese in Utah. Tran. N. ,\m. Wildl. Nat. Resources Conf. 40:.334-343. TwoMEY, A. C. 1942. The birds of the Uinta Basin, Utah. Ann. Carnegie Mus. 28:,341-490. NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be di- rected to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be sim- ilarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. They should be typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. Use a recent issue of either journal as a format, and the Council of Biology Edi- tors Style Manual, Third Edition (AIBS 1972) in preparing the manuscript. An abstract, about 3 percent as long as the text, but not exceeding 200 words, written in accordance with Biological Abstracts guidelines, should precede the introductory paragraph of each ar- ticle. Authors may recommend one or two reviewers for their article. All manuscripts re- ceive a critical peer review by specialists in the subject area of the manuscript under con- sideration. Manuscripts that are accepted and that are less than 100 printed pages in length will automatically be assigned to the Great Basin Naturalist. Those manuscripts larger than 100 printed pages in length will be considered for the Memoirs series. Illustrations and Tables. All illustrations and tables should be made with a view toward having them appear within the limits of the printed page. Illustrations that form part of an article should accompany the manuscript. Illustrations should be prepared for reduction by the printer to either a single-column (2 5/8 inches) or double-column (5 1/2 inches) width, with the length not exceeding 7 1/2 inches. Costs Borne by Contributor. Contributors to the Great Basin Naturalist should be pre- pared to donate from $10 to $30 per printed page toward publication of their article (in addition to reprint costs outlined in the schedule below). Authors publishing in the Great Basin Naturalist Memoirs may be expected to contribute $35 per printed page in addition to the cost of printed copies they purchase. No printed copies are furnished free of charge. A price list for reprints and an order form are sent with the galley proof to contributors. Reprint Schedule of the Great Basin Naturalist 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each 100 copies $20 $24 $28 $32 $36 $40 additional 200 copies 28 32 36 40 44 48 2 pp. 300 copies 36 40 44 48 52 56 $4 Great Basin Naturalist Memoirs No. 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodburry, H. H. Frost. $10. No. 2 Intermountain biogeography: A symposium. By K. T. Harper, J. L. Reveal, et al. $15. GREAT BASIN NATURALIST Editor. Stephen L. Wood, Life Science Museum, Brigham Young University, Provo, Utah 84602. Assistant Editor. Stanley L. Welsh, Life Science Museum, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; William D. Tidwell, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agri- cultural Sciences; Ernest L. Olson, Director, Brigham Young University Press, Univer- sity Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been pub- lished from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only previously unpublished manuscripts of less than 100 printed pages in length and pertaining to the biological natural history of western North America are ac- cepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manu- scripts are subject to the approval of the editor. Subscriptions. The annual subscription rate of the Great Basin Naturalist is $12 (outside the United States the rate is $13). The price for single numbers is $4 each. All back num- bers are in print and are available for sale. All matters pertaining to the purchase of sub- scriptions and back numbers should be directed to Brigham Young University Press, Mar- keting Department, 204 UPB, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same office at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either jour- nal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts or other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back 1978 Brigham Young University Press. All rights reserved Printed in United States of America 650 3-78 29240 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 37 September 30, 1977 No. 3 UTAH FLORA: BRASSICACEAE (CRUCIFERAE) Stanley L. Welsh' James L. Reveal- Abstract.— The mustard family, Brassicaceae (Cruciferae) is revised for the state of Utah. Treated are 155 spe- cies and 44 varieties, including 37 species of introduced weeds or escaped cultivated plants. A key to the genera and species is included, along with detailed descriptions, distribution data, and pertinent comments for many of the taxa. Proposed new varieties are: Lepidium montanum Nutt. var. stellae Welsh & Reveal; L. montanum var. neeseae Welsh & Reveal; Lesquerella hemiphysaria Maguire var. liicens Welsh & Reveal; Physaria aciitifolia Rydb. var. purpurea Welsh & Reveal; and, Tfielypodhtm sagittatum (Nutt.) Endl. in Walp. var. vennicularis Welsh & Reveal. The following new combinations are made: Arabis confinis S. Wats. var. interposita (Greene) Welsh & Reveal; Descurainia pinnata (Walt.) Britt. var. paysonii (Detl.) Welsh & Reveal; D. richardsonii (Sweet) O. E. Schulz var. brevipes (Nutt.) Welsh & Reveal; Draba asprella Greene var. zionensis (C. L. Hitchc.) Welsh & Reveal; Draba densifolia Nutt. ex Torr. & Gray var. daviesiae (C. L. Hitchc.) Welsh & Reveal; Lesquerella alpina (Nutt.) S. Wats. var. parvula (Greene) Welsh & Reveal; Lesquerella kingii S. Wats. var. parvifolia (Maguire & Holmgren) Welsh & Reveal; Rorippa islandica (Oed.) Borbas var. glabra (O. E. Schulz) Welsh & Reveal; Thelypo- diopsis divaricata (Rollins) Welsh & Reveal; Thelypodium integrifolium (Nutt.) Endl. var. affine (Greene) Welsh & Reveal. This paper, dealing with the Brassicaceae, There are 155 species and 44 varieties. Ad- is one of a series of proposed revisions ventive weedy species and cultivated plants which will result ultimately in a definitive known to escape and persist constitute 37 treatment of the flora of Utah. Not since species and two varieties. Thus, almost 20 the work of Tidestrom (1925) has the family percent of the taxa included are introduced, been treated in its entirety, although a par- It is unlikely that any of the other large tial treatment is to be found in the work by families of vascular plants in Utah, such as Welsh and Moore (1973). The family has Asteraceae, Poaceae, Fabaceae, or Rosaceae not been neglected by monographers, how- will contain such a large proportion of ever. That observation is supported by the weedy and adventive taxa. numerous references listed following the ge- Generic limits in this family are often dif- neric descriptions in the taxonomic treat- ficult to define, and because of this well- ment. known problem, some taxa have been The family is both large and tax- shifted from one genus to another, or even onomically complex. In the present treat- among several genera, depending on the au- ment, some 199 taxa are recognized as oc- thority one chooses to follow. One author, curing (or probably occuring) in Utah. Krause, writing in Sturm's Flora von 'Department of Botany and Range Science, and Life Science Museum, Brigham Young University, Provo, Utah 84602. 'Departments of Botany, University of Maryland, College Park, Maryland 20742, and National Museum of Natural History, Smithsonian Institution, Washington, DC. The support provided by National Science Foundation grant BMS 75-13063 is acknowledged. The work reported here was conducted while on sabbatical leave to Brigham Young University from January to July 1976. 279 280 Great Basin Naturalist Vol. 37, No. 3 Deutchland in 1902, proposed an inclusive genus, Crucifera, for plants treated in this paper as belonging to Conringia, Diplotaxis, Erysimum, Brassica, Lepidium, Hesperis, Draba, Barbarea, Hutchinsia, Baphanus, Si- symbrium, and Thlaspi. Such an approach is even more frustrating than treating taxa with obscure taxonomic limits; it is neither realistic nor is it practical. In the Utah flora, problems of generic limits fall mainly within the purportedly primitive members of the family, such as Streptanthus, Thelypodium, Tlielypodiopsis, Caulanthus, Chlorocrambe, and Sisymbrium. The approach taken herein is somewhat conservative, as we have tended to accept the various genera which appear to be more or less distinct, not only in Utah, but in other western states as well. Even so, the species treated in Caulanthus do not appear to represent a natural group, and some spe- cies of Tlielypodiopsis appear to be more closely related to taxa in other genera, espe- cially in Thelypodium, than they are to each other. The species in Caulanthus seem to show affinities to Thelypodium, Tlielypo- diopsis, Chlorocrambe, and Streptanthus. In large part, we have tended to follow the published opinions of Dr. Reed C. Roll- ins of Harvard University, long-time student of the family, but we feel that much more work -is needed in ascertaining relationships and hope that students will investigate some of the more difficult elements in the future. Some solutions might be forthcoming from studies of fine structure in various mor- phological features of the species, especially those to be derived from pollen mor- phology, from a review of phytochemistry beyond that which has already been done, and from a detailed review of cytology of the troublesome genera. Continued reliance upon floral and fruit- ing characteristics, features upon which taxa have been and are currently based, is a part of the problem of arriving at a realistic sys- tem of classification of genera. An entity represents the sum of its features; yet in the Brassicaceae, much emphasis, sometimes to the exclusion of other features, has been placed on features of flowers and fruit, and, until work is directed toward an attempt to summarize all features, our understanding of the genera in this family will remain con- fused. Genera and species, in the present treat- ment, are arranged alphabetically. The key to genera is divided into seven different, morphologically based subkeys. These arti- ficial subkeys provide leads to groups of species or individual taxa within the various genera. Literature Gited Holmgren, P. K., and W. Keuken. 1974. Index her- barionim-I. Edition 6. Regnum Veg. 92: 1-397. Lawrence, G. H. M., A. F. G. Buchheim, G. S. Daniels, and H. Dolezal. 1968. B-P-H, Bo- tanico-Periodicum-Huntianum. Pittsburgh: Hunt Botanical Library. 1063 pp. Stafleu, F. a., and R. S. Cowan. 1976. Taxonomic literature. Volume 1: A-G. Regnum Veg. 94: 1- 1136. TiDESTROM, I. 1925. Flora of Utah and Nevada. Contr. U.S. Natl. Herb. 25: 1-665. Welsh, S. L., and G. Moore. 1974. Utah plants, Tracheophyta. Provo: Brigham Young University Press. 474 pp. Brassicaceae Burnett Mustard Family Annual to perennial herbs or rarely suf- frutescent, often with a pungent watery juice, glabrous to pubescent with simple or more commonly branched to stellate hairs, often glaucous; leaves alternate or basal and still alternate, entire or lobed, simple to compound, exstipulate, the cauline leaves usually sessile and auriculate; flowers per- fect, regular or nearly so, hypogynous, borne in ebracteate or rarely bracteate racemes, spikes, or corymbs, rarely solitary and terminal on leafless scapes; sepals 4, erect to somewhat spreading, greenish to colored, the outer 2 often somewhat bulged or saccate at the base; petals 4, rarely lack- ing, yellow, white, or pink to blue or purple, commonly clawed, the blade rounded to occasionally bilobed and spread- ing in the form of a cross (hence Cruci- ferae); stamens 6 with the outer 2 in.serted lower and shorter than the other 4 (tetrady- namous), rarely 4 or 2; nectar glands com- monly 4; ovary superior, 2-loculed or, less commonly, only 1-loculed, usually with a Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 281 thin partition (replum) between the two the repkiin or, less commonly, flattened marginal placentae from which, when ma- contrary to the replum; seeds (1) 2 to sever- ture, the valves usually separate, the stigma al in a single row per locule (uniseriate) or rather small, entire to shallowly 2-lobed; more or less distinctly arranged in 2 rows fruit usually nonstipitate but occasionally so (biseriate), smooth to striate or pitted and with a slender stipe, typically dehiscent or plump to flattened or even wing margined, if indehiscent then constricted between the exalbuminous. seeds (tonilose) and sometimes breaking A large, worldwide family of perhaps 375 transversely, several times longer than broad genera and 3000 species, with many plants (silique) to much shorter and not more than of economic value as ornamentals, impor- 2 or 3 times as long as broad (silicle), the tant vegetables, or noxious weeds, sides flattened or compressed parallel with 1. Plants with cauline leaves both sessile and auriculate (at least some), or auriculate and petiolate (in some Barbarea, Nasturtiuyn, and Rorippa) 2 Plants without cauline leaves or with cauline leaves not auriculate and ses- sile, either petioled or merely sessile 4 2(1). Petals yellow; plants glabrous or with simple hairs, rarely with some mal- pighian hairs in Thelypodiopsis Key I Petals white, pink, lavender, chestnut, or purple, but not yellow (cream colored in Camelina and Arahis); plants glabrous or variously pubescent 3 3(2). Plants glabrous or with simple trichomes only Key II Plants pubescent with at least some malpighian, branched, or stellate hairs Key III 4(1). Petals yellow, sometimes fading white or pinkish to purplish 5 Petals white, pink, lavender, purple, or chestnut, but not yellow 6 5(4). Plants glabrous or with simple hairs only Key IV Plants pubescent with malpighian, branched, or stellate hairs Key V 6(4). Plants glabrous or with simple hairs only Key VI Plants pubescent with at least some malpighian, branched, or stellate hairs Key VII Key I Cauline leaves auriculate; petals yellow 1. Uppermost cauline leaves falsely perfoliate-clasping; basal leaves finely dis- sected, usually lacking at anthesis; seeds 2, one in each locule; silicles about as broad as long (L. perfoliatum) Lepidium Uppermost cauline leaves various, but not perfoliate-clasping, more than twice as long as broad; basal leaves pinnatifid to dentate or subentire; seeds several to many, or reduced to one in Isatis; siliques or silicles more than twice as long as broad 2 2(1). Cauline leaves lyrate-pinnatifid or falsely petiolate above the auriculate base 3 Cauline leaves entire, ovate to oblong or lanceolate, undulate or rarely toothed, sessile 4 282 Great Basin Naturalist Vol. 37, No. 3 3(2). Styles (0.5) 1-2 mm long, abruptly contracted at the stigma; fruit 15 mm long or more, many times longer than broad; plants not rhizomatous Barbarea Styles 0.5-0.8 (1.3) mm long, or if longer, then plants rhizomatous, tapering to the stigma; fruit less than 10 mm long (rarely longer), 1 to several times longer than broad Rorippa 4(2). Cauline leaves hastately lobed; pedicels deflexed in fruit; slilicles cuneate, winged-flattened 1-seeded, 1-loculed Isatis Cauline leaves clasping-auriculate; pedicels spreading-ascending to ascend- ing in fruit; siliques linear, not or only somewhat flattened, not winged, many-seeded, 2-loculed 5 5(4). Cauline leaves broadly rounded to truncate apically, strictly entire, elliptic- oblong to lance-oblong Conringia Cauline leaves rounded to acute apically, more or less dentate to entire, ta- pering from base to apex 6 6(5). Plants biennial; weeds of cultivated or disturbed sites; basal leaves lyrate- pinnatifid {B. campestris) Brassica Plants perennial or biennial, rarely annual; indigenous plants of clay soils; basal leaves dentate to entire 7 7(6). Stamens exserted; siliques long-stipitate, the stipe 1-2 cm long or more (S. viridiflora) Stanleya Stamens included; siliques sessile or only short-stipitate Thelypodiopsis Key II Cauline leaves auriculate; petals white, pink, lavender, or chestnut; herbage glabrous or with simple trichomes only 1. Leaves pinnately compound or pinnatifid; plants aquatic, glabrous (or nearly so); flowers white Nasturtium Leaves simple, entire or merely toothed; plants terrestrial; flowers white, pink, lavender, or chestnut 2 2(1). Uppermost cauline leaves commonly rounded to emarginate or truncate apically, the lower ones dentate apically; flowers chestnut to brown-purple or purple; siliques 3-5 (6) mm broad Streptanthus Uppermost cauline leaves attenuate to acute apically, the lower ones vari- ous but not apically dentate only; flowers white, pink, or lavender 3 3(2). Plants annual or winter annual; pedicels recurved in fruit; fruit indehiscent, winged-flattened, 1-seeded, 1-loculed; restricted to Washington Co. Thysanocarpus Plants annual, biennial, or perennial; pedicels spreading-ascending to erect, rarely descending in some Thlaspi; fruit dehiscent or indehiscent in Car- daria, winged-flattened to subterete, 2 or more seeded; distribution vari- ous 4 4(3). Limb of petal 4-6 (7) mm long; sepals mostly 4-7 mm long; siliques (at least some) more than 5 cm long 5 Limb of petal 2-3 mm long or less; sepals mostly 2-4 (7) mm long; siliques or silicles less than 5 cm long except in Arabis 6 Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 283 5(4). Petals obovate, constricted at the base, the claw broad; stigma expanded and deeply bilobed; plants of eastern Utah (T. elegans) Thelypodiopsis Petals oblanceolate, tapering to a slender claw; stigma narrowed, not deep- ly bilobed; plants mostly of north-central and western Utah (T. sagitta- tiim) Thelypodium 6(4). Fruit 10-30 times longer than broad or more, linear or narrowly oblong in outline 7 Fniit 1-4 times longer than broad or less, clavate to obcordate, ovate, or cordate-reniform in outline 9 7(6). Pedicels mostly 8-12 mm long, erect or nearly so; plants of middle altitudes in mountains (A. hirsuta) Arabis Pedicels 2-7 mm long, spreading-ascending to descending; plants of low to moderate elevation 8 8(7). Plants 10 dm tall or more, biennial, the basal leaves often withered at an- thesis; cauline leaves numerous, somewhat hastately and acutely auriculed; known only from central northern Utah {T. rolUnsii) Thelypodium Plants 4 dm tall or less, annual or winter annual, the basal leaves not with- ered at anthesis; cauline leaves few to several, cordate-auricled to merely sessile; known only from Washington Co. (C. cooperi) Caulanthus 9(6). Seeds and ovules 3 to many per fruit, usually 2 or more in each locule; fruit conspicuously winged or more than twice as long as broad Thlaspi Seeds and ovules 2 per fruit, 1 in each locule; fruit not or only somewhat winged 10 10(9). Fruit broader than long, indehiscent; upper cauline leaves ovate; racemes 2-5 cm long in fruit, numerous Cardaria Fruit longer than broad, dehiscent; upper cauline leaves lanceolate; ra- cemes 5-10 cm long or more in fruit, few (L. campestre) Lepidium Key III Cauline leaves auriculate; petals white, pink, or lavender, (except in Camelina and Arabis); herbage pubescent with malpighian, branched, or stellate hairs 1. Plants indigenous biennials or perennials of broad distribution in native plant communities; fruits siliques, several to many times longer than broad; flowers pink, lavender, or white (cream in Arabis glabra) 2 Plants adventive annuals or winter annual of disturbed or cultivated places; fruits silicles, less than 3 times longer than broad; flowers usually white 3 2(1). Plants pubescent with mixed simple, forked, and branched hairs; siliques subquadrangular; known only from the northern foothills of the Uinta Mts Halimolobos Plants variously pubescent but not as above nor of that distribution; siliques distinctly flattened Arabis 3(1). Plants flowering in early springtime; siliques triangular-obcordate, com- pressed Capsella Plants flowering in late springtime and summer; siliques obovoid, terete or nearly so Camelina 284 Great Basin Naturalist Vol. 37, No. 3 Key IV Cauline leaves sessile or petiolate, not auriculate, or if hastately lobed then petiolate; flowers yellow; herbage glabrous or with simple hairs only 1. Cauline leaves both hastately lobed and petiolate, the leaf-blades triangular- ovate or lanceolate, entire; plants mostly 8 dm tall or more; of middle elevations often in dense vegetation Chlorocrambe Cauline leaves pinnatifid to entire, the leaf-blades not hastately lobed; plants low to tall, but if tall then seldom if ever of middle elevations in dense vegetation 2 2(1). Leaves all simple and entire or sparingly toothed; plants perennial from a caudex 3 Leaves (at least some) pinnatifid or definitely and regularly toothed (or, if all entire, then plant not arising from a caudex); plants annual, biennial, or perennial, sometimes with a distinct caudex 6 3(2). Plants low, less than 1 dm tall; cauline leaves lacking; silicles lanceolate in outline, 1-4 times longer than broad (D. densifolia) Draha Plants 1-5 dm tall or more; cauline leaves present; siliques linear, many times longer than broad 4 4(3). Stamens long-exserted from the flower; pedicels spreading; siliques long- stipitate on stipes 10-25 mm long or more Stanleya Stamens not exserted beyond the flower; pedicels ascending to suberect; siliques sessile, subsessile, or on a short stipe less than 1 mm long 5 5(4). Siliques 10-20 mm long, 2-3 mm wide; plants known only from Uintah Co. Glaucocarpwn Siliques 25-65 mm long, 0.8-1.2 mm wide; plants widespread and often weedv Schoencratnbe 6(2). Plants growing in mud, along beaches, or in or near streams; petals usually less than 3 mm long; siliques usually less than 12 mm long Rorippa Plants seldom if ever in perennially moist sites; petals more than 4 mm long or siliques more than 12 mm long, or both 7 7(6). Leaves glaucous, thickened; perennial herbs from an often woody caudex; stamens long-exserted; siliques long-stipitate on stipes 10-25 mm long Stanleya Leaves not glaucous, or, if so, not especially thickened; annual, biennial, or perennial herbs without a distinct caudex (except Schoencrambe); stamens not exserted; siliques not stipitate 8 ' 8(7). Plants perennial, rhizomatous, indigenous, of lower elevations in native plant communities Schoencrambe Plants annual or biennial, adventive, of disturbed or cultivated places 9 9(8). Fruits dehiscent to the apex; pedicels spreading-ascending to ascending, or, if erect (as in S. officinalis), the petals only 3-4 mm long Sisymbrium Fmits with long, indehiscent sterile apices; pedicels ascending, or, if ap- pressed (as in B. nigra), the petals 5-8 mm long or more 10 10(9). Plants mostly 2-5 dm tall; leaves mainly basal; fruiting racemes longer than the leafy stems; seeds biseriate Diplotaxis Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 285 Plants mostly 4-10 dm tall or more; leaves basal and cauline; fruiting ra- cemes shorter than the leafy stems; seeds uniseriate Brassica Key V Plants with cauline leaves sessile or petiolate or lacking; petals yellow; herbage pubescent with malpighian, branched, or stellate hairs 1. Leaves once to twice or rarely thrice pinnately dissected or compound; plants annual or winter annual Descurainia Leaves simple and entire or merely toothed or lobed; plants annual, bien- nial, or perennial 2 2(1). Cauline leaves lacking, all leaves basal Draba Cauline leaves present, at least one, the basal leaves present or absent 3 3(2). Plants pubescent with appressed, Y-shaped and/or malpighian hairs; siliques many times longer than broad Erysimum Plants pubescent with branched, dendritic, or appressed stellate hairs; silicles from about as broad as long to about 5 times longer than broad 4 4(3). Pubescence of branched or dendritic hairs, rarely with some stellate; silicles lance-ovate to oblong or elliptic, usually more than twice as long as broad Draba Pubescence of appressed stellate hairs only (except for Alyssum saxatile); silicles orbicular in outline or subglobose to bladdery inflated, not over twice as long as broad 5 5(4). Silicles compressed, lens-shaped; plants annual Alyssum Silicles either subglobose or greatly inflated and terete or didymous; plants annual, biennial, or perennial 6 6(5). Basal leaf blades often exceeding 20 mm in width; silicles (excluding style) 8-10 mm long or more, didymous and bladdery inflated, often cordate at the base Physaria Basal leaf blades rarely up to 20 mm in width; silicles (excluding style) 3-6 mm long, not didymous, seldom bladdery inflated, not cordate at the base , Lesquerella Key VI Plants with cauline leaves sessile or petiolate or lacking; petals white, pink, lavender, purple, or chestnut; herbage glabrous or with simple hairs only 1. Herbage stipitate-glandular or glabrous and the plants scapose and with large flowers; plants either alpine perennials or winter annuals of low elevations 2 Herbage glabrous or simple hirsute; plants of various habits, habitats, and duration 3 2(1). Plants scapose alpine perennials; leaves basal; petal blades 8-10 mm long or more; siliques flattened, dehiscent Parrya Plants winter annuals; leaves cauline and basal; petal blades 2-4 mm long; siliques terete, indehiscent Chorispora 286 Great Basin Naturalist Vol. 37, No. 3 3(1). Blades of cauline leaves hastately lobed, otherwise entire, triangular-ovate to triangular-lanceolate; plants 8-10 din tall or more; siliques short-stipitate, the stipe 2-7 mm long Chlorocrambe Blades of cauline leaves, if any, not both hastately lobed and entire; plants low to tall; siliques or silicles subsessile or sessile on the pedicel (except in Lunaria) 4 4(3). Plants dwarfed scapose perennials; leaves often less than 10 mm long; scapes often less than 10 cm tall Draha Plants caulescent, at least some leaves cauline; leaves mostly more than 10 mm long; stems usually over 10 cm tall 5 5(4). Plants slender annuals, 1-2.5 dm tall; leaves entire or merely serrate (at least the cauline ones); fruits obovate in outline and several-seeded, or, if linear, less than 15 mm long 6 Plants annual, biennial, or perennial, 1.5-10 dm tall or more; leaves vari- ously toothed, lobed, or entire; fruits obovate in outline and 2-seeded, or lance-ovoid, or, if linear, mostly more than 15 mm long 7 6(5). Petals 1.2 mm long or less; silicles obovate, compressed, 2-3 mm long Hutchinsia Petals 2-2.9 mm long; siliques linear, terete, 9-14 mm long Arabidopsis 7(5). Basal leaves long-petiolate, the blades oblong to lanceolate, mostly 10-30 (50) cm long or more; cauline leaves lance-oblong, irregularly crenate- serrate; petals white; silicles less than twice longer than broad, abor- tive; plants cultivated but escaping and often persistent Arrnoracia Basal leaves sessile, poorly developed, or if long-petiolate, the blades sel- dom as above, usually less than 10 cm long; cauline leaves not both lance-oblong and irregularly crenate-serrate; petals pink, white, or chest- nut purple; fruit various; plants indigenous or cultivated 8 8(7). Petal blades 8-12 mm long, pink to lavender; silicles more than 20 mm wide; styles 6-8 mm long; leaves cordate-ovate, irregularly toothed Lunaria Petal blades usually less than 8 mm long; silicles or siliques much less than 10 mm wide; styles less than 3 mm long; leaves various but not cordate- ovate (except in Cardamine cordifolia) 9 9(8). Seeds and ovules 2, 1 in each locule; silicles obcordate, orbicular, or ellip- tic, usually about as long as broad Lepidhmi Seeds and ovules more than 2, usually more than 2 in each locule; silicles lance-oblong or siliques linear and 3 to many times longer than broad 10 10(9). Plants annual, with thickened tuberous roots; flowers white or pink, the petal limb more than 5 mm long; siliques terete, indehiscent Raphanus Plants perennial, biennial, or, if annual, not from thickened roots; flowers variously colored, but if white or pink then the petal limb less than 5 mm long; siliques or silicles terete or flattened, dehiscent 11 11(10). Cauline leaves ovate, cordate-ovate, or pinnately compound; flowers white; plants of moist sites Cardamine Cauline leaves pinnatifid or entire to subentire and linear to elliptic, but not as above; flowers white or brown-purple, lavender or with purple veins; plants of various habitats 12 12(11). Flowers minute, less than 1.2 mm long; silicles lance-oblong, less than 10 Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 287 mm long; leaves lyrate-pinnatifid (R. tenerrima) Rorippa Flowers 2-6 mm long or more; siliques linear to oblong, much more than 10 mm long; leaves pinnatifid to toothed or entire 13 13(12). Flowers subsessile, the pedicels less than 2 (4) mm long; leaves, at least some, definitely pinnatifid; petals mostly brown-purple Caulanthus Flowers pedicellate, the pedicels usually over 2 mm long, or the leaves en- tire; petals variously colored, but, if brown-purple, pedicels definitely longer than 2 mm 14 14(13). Plants annual; pedicels curved-descending; siliques compressed, the sutures parallel, not tonilose Streptanthella Plants biennial or perennial; pedicels spreading to spreading-ascending (de- clined in Thelypodium wrightii); siliques with parallel sides or else torulose 15 15(14). Petals strongly purple-veined; leaves linear; plants from a caudex, known only from Uintah Co. (T. argillacea) Thelypodiopsis Petals not or seldom purple veined; leaves various; plants with or without a caudex, of various distribution 16 16(15). Plants erect biennials; basal rosette prominent, but usually withered at an- thesis; pedicels spreading or declined; siliques torulose; flowers numerous to very numerous Thelypodium Plants ascending to erect perennials; basal rosette not especially prominent; pedicels spreading-ascending; siliques with parallel sides; flowers several to many (A. nuttallii) Arabis Key VII Plants with cauline leaves sessile, petiolate, or lacking; petals white, pink, lavender, or purple; herbage pubescent with malpighian, branched, or stellate hairs 1. Leaves pinnately lobed; herbage cinereous pubescent; plants of high eleva- tions Srnelowskia Leaves entire to serrate or sinuate-dentate; herbage green or glaucous (cin- ereous in Dithyrea); plants of low to high elevations 2 2(1). Plants scapose, the leaves basal, mostly less than 1 dm tall Draba Plants with at least some leaves cauline, often more than 1 dm tall 3 3(2). Petal blades more than 6 mm long; sepals definitely saccate, 6-8 mm long; cauline leaves sinuate-dentate; plants 5-10 dm tall or more, cultivated and frequently escaping Hesperis Petal blades less than 6 mm long; sepals not especially saccate, 1-6 mm long; cauline leaves entire or sinuate-dentate; plants (0.5) 1-4 dm tall, in- digenous or adventive, not cultivated (except Lobularia) 4 4(3). Styles (1.5) 2-3 mm long; ovaries appressed stellate pubescent, about twice as long as broad, dehiscent; seeds several Berteroa Styles 0.2-1 (1.5) mm long; ovaries variously pubescent, but if appressed stellate then much broader than long, dehiscent or indehiscent; seeds 1 to many 5 5(4). Pedicels 5-21 mm long, spreading; silicles with stellate or dendritic pub- 288 Great Basin Naturalist Vol. 37, No. 3 escence, more than twice as broad as long, 2-seeded Dithyrea Pedicels less than 8 mm long or not spreading, or, if spreading, silicles or siliques not with stellate or dendritic hairs and broader than long; seeds 1, 2, or more 6 6(5). Petals minute, less than 2 mm long; seeds 1 or 2; fruit indehiscent; plants annual or winter annual 7 Petals mostly 2-10 mm long or more; seeds 2 to many; plants annual, bien- nial, or perennial 8 7(6). Ovaries and fruit with dendritic pubescence; fruit ovoid, with a curved sty- lar beak up to 1 mm long, 2-seeded Eiiclidiiim Ovaries and fruit with simple hairs; fruit orbicular, compressed, lacking a stylar beak, 1-seeded Athysanus 8(6). Pedicels less than 2 mm long, appressed or ascending; leaves sinuate-den- tate; siliques ascending, 33-63 mm long, pubescent with dendritic or branched hairs, the stigma oblique Malcolmia Pedicels 2-10 mm long, descending to spreading or ascending; silicles or si- liques various, sometimes pubescent with dendritic or branched hairs, the stigma terminal 9 9(8). Pedicels descending; fruit pendulous, 13-65 mm long; plants perennial Arabis Pedicels spreading to ascending; fruit erect or spreading-ascending, 2-14 mm long; plants annual or perennial 10 10(9). Leaves mainly cauline, strigose with malpighian hairs; fruit about as broad as long; seeds 2; plants cultivated and often escaping Lobularia Leaves mainly basal, with stalked dendritic or stellate hairs; seeds usually more than 2; plants indigenous or adventive 11 11(10). Fruit linear to narrowly oblong, terete or subterete, less than 1 mm broad; plants slender annuals Arabidopsis Fruit lance-ovate to lanceolate or oblong in outline, flattened, 1.5-3 mm broad; plants annual or perennial Draba Alyssum L. cream or white), emarginate; stamens 6, at Plants stellate-pubescent annuals or pe- least the two shorter filaments with a whit- rennials from taproots; leaves alternate, ish process near base; style slender, stigma simple, entire, tapering to base, not auricu- capitate; fruit a sessile silicic, less than late; flowers in racemes, pedicels spreading- twice longer than broad, broadly elliptic to ascending to ascending or erect, not sub- oval in outline, compressed parallel to the tended by bracts; sepals 4, deciduous or septum, valves veinless; seeds 1 or 2 per lo- persistent; petals 4, yellow (often fading cule. 1. Cauline leaves, at least some, more than 4 cm long; flowers bright yellow, fading cream; petal blades obcordate, about as broad as long; plants perennial, cultivated and occasionally escaping A. saxatile Cauline leaves less than 4 cm long; flowers pale yellow (fading white); pet- al blades cuneate, much longer than broad; plants adventive annuals of arid sites 2 2(1). Silicles glabrous or essentially so, orbicular in outline; styles 0.5-0.8 mm long, persistent A. desertorum Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 289 Silicles stellate-pubescent, orbicular to elliptic in outline; styles various 3 3(2). Silicles 4-5 mm broad; styles 0.8-1.2 mm long; pubescence of coarse spreading-ascending stellate hairs A. minus Silicles 2.8-4 mm broad; styles 0.3-0.6 mm long; pubescence of delicate ap- pressed-stellate hairs 4 4(3). Silicles emarginate at the style, about as broad as long; hairs of fruit mi- nute, not or seldom overlapping A. ahjssioides Silicles truncate at the style, longer than broad; hairs of fruit ample, usually overlapping A. szowitsianum Alyssum alyssioides (L.) L. (Chjpeoki alyssioides L.) Alyssum. Plants annual, stems ascending to erect, simple or branched from near base (3) 6-25 cm tall; leaves spatulate to oblanceolate, 5-20 (37) mm long, 2-6 mm broad, stellate and green on both surfaces; pedicels spreading-ascending, 1.5-4 (5) mm long; sepals 2.1-2.8 (3) mm long, green, stel- late-pubescent, persistent; petals 3-4.2 mm long, yellow, fading white, cuneate, surpass- ing sepals; silicles 3-4 mm long and about as broad, valves minutely stellate-pubescent, emarginate at style; styles persistent, mostly 0.3-0.6 mm long. Foothills, roadways, and other dry sites at low elevations in Cache, Salt Lake, Tooele, Utah, and Wasatch coun- ties, and to be expected elsewhere; wide- spread in North America; adventive from Europe. Alyssum desertorum Stapf. Desert Alys- sum. Plants annual, stems decumbent to as- cending or erect, simple or branched from near the base, (3) 4-20 cm tall; leaves spatu- late to oblanceolate or almost linear, 4-22 mm long, 2-3 mm wide, stellate and green on both surfaces; pedicels spreading-ascend- ing, 1.8-2.5 mm long; sepals 1.2-1.9 mm long, often suffused with red, stellate- pubescent; petals 2.3-2.8 mm long, yellow, fading white, cuneate, surpassing sepals; si- licles 2.9-3.8 mm long and about as broad, valves glabrous or rarely with scattered stel- late hairs, emarginate at the style; styles persistent, 0.5-0.8 mm long. Foothills, road- sides, and other dry sites at low elevations in Morgan, Salt Lake, Sanpete, Tooele, Utah, and Weber counties, and to be ex- pected elsewhere; Washington, Oregon, Idaho, and Montana; adventive from Asia Minor and adjacent southeastern Europe (Rollins, R. C. 1941. Contr. Dudley Herb. 3: 183). Alyssum minus (L.) Roth. {Clypeola minor L.). Plants annual, stems ascending to erect, simple or branched from near base, 0.9-2.5 dm tall; leaves spatulate to oblanceolate, 4- 25 mm long, 2-7 mm broad, coarsely stel- late and green on both surfaces; pedicels spreading-ascending, 2.8-4.9 mm long; se- pals (1.5) 1.7-2.5 mm long, often suffused with red, stellate-pubescent, caducous; pet- als 2.7-3.6 mm long, yellow, fading white, surpassing sepals; silicles (3.5) 4-5.2 mm long and about as broad or broader, valves coarsely stellate with ascending rays, emarginate at style; styles persistent, 0.8-1.2 mm long. Foothills and roadsides in Salt Lake Co., and to be expected elsewhere; California and Colorado; adventive from the Old World. Alyssum saxatile L. Sweet Alyssum. An- nual, stems sprawling-decumbent to ascend- ing or erect, branched from base and above, 20-35 cm tall; leaves oblanceolate to elliptic or oblong, 90-135 mm long, 2-17 mm wide, minutely stellate and with long, forked, and simple hairs, and green on both surfaces; pedicels spreading-ascending, 3.5-8 mm long; sepals 1.7-3 mm long, green or cream, loosely stellate to glabrate, caducous; petals 4-5 mm long, yellow, fading cream, much surpassing sepals; silicles 3.5-4.5 mm long or more and about as broad, valves glabrous, truncate to rounded at style; styles per- sistent, 0.5-1 mm long. Cultivated ornamen- tal, occasionally escaping; introduced from southern Europe Alyssum szowitsianum, Fisch. & Meyer Szowits Alyssum. Plants annual, stems de- cumbent-ascending to erect, usually 290 Great Basin Naturalist Vol. 37, No. 3 branched from near base, 5-15 cm tall; leaves oblanceolate, 8-25 mm long, 1-5 mm broad, stellate and green or suffused with red on both surfaces; pedicels ascending to erect, 2.7-4.2 mm long; sepals 0.9-1.2 (2) mm long, often suffused with red, stellate- pubescent, caducous; petals 1.7-2 mm long, yellow, fading white, only slightly surpass- ing the sepals; silicles 4-5 (6) mm long, longer than broad, valves densely stellate, truncate at style; styles persistent, 0.5-0.6 mm long. Dry foothills, in Salt Lake Coun- ty, and to be expected elsewhere; adventive from Europe. Arabidopsis (DC.) Schur Plants annual, glabrous, or pubescent with simple or branched hairs, from tap- roots; leaves alternate or basal (and still al- ternate), simple, entire, or remotely serrate, tapering to l3ase, not auriculate; flowers in racemes, pedicels spreading-ascending, not subtended by bracts; sepals 4, deciduous; petals 4, white, not emarginate; stamens 6, at least the two shorter filaments subtended by a semicircular gland; style very short, ta- pering, stigma not enlarged; silique sessile, several times longer than broad, subterete, valves with conspicuous mid-nerve; seeds several to many in each locule. Arabidopsis thaliana (L.) Schur. Mouse- ear Cress. {Arabis thaliana L.). Plants slen- der annuals, stems erect or nearly so, usual- ly branched throughout, 8-30 (40) cm tall; leaves mainly basal, spatulate to oblong, 3- 30 (50) mm long, 2-8 mm broad, remotely toothed to subentire, pubescent with simple or 2- to 4 (5) -rayed hairs, green; cauline leaves much smaller than basal ones, lance- oblong, sessile or nearly so, 5-20 mm long, 2-5 mm wide; pedicels very slender, 2.5-10 mm long; sepals 1.2-1.7 mm long, green to cream or reddish tinged, sparingly long- hairy; petals 2-2.9 mm long, white, spatu- late, surpassing sepals; siliques 9-14 mm long, 0.5-0.8 mm broad, valves glabrous; styles 0.2-0.3 mm long. Dry hillsides at lower elevations in Salt Lake and Weber counties, and to be expected elsewhere; widespread in temperate North America; adventive from Europe. Arabis L. Rock-Cress Plants biennial or perennial, glabrous or pubescent with simple, branched, or stellate hairs; leaves alternate and basal, simple, en- tire, dentate, serrate, or sinuate, tapering to base or cauline blades sessile and usually au- riculate; flowers in racemes, the pedicels erect, ascending, spreading-ascending, spreading, descending, or reflexed, not sub- tended by bracts; sepals 4, deciduous; petals 4, white, pink, lavender, or purple (cream in A. glabra); stamens 6, at least the two shorter filaments subtended by glands; styles prominent to lacking, the stigmas entire to lobed; fruit a sessile or substipitate silique many times longer than broad, laterally flat- tened, valves usually with a midnerve; seeds numerous. A large genus of perhaps 100 species of Eurasia and North America. Rollins, R. C. 1941. Monographic study of Arabis in western North America. Rho- dora 43: 289-325, 348-411, 425-481. Re- printed at Contr. Gray Herb. 138. 1. Cauline leaves usually attenuate to rounded basally, either petiolate or ses- sile but not auriculate (rarely so in A. pendulina) 2 Cauline leaves auriculate, at least some 6 2(1). Petals 9-18.5 mm long; petal limb divaricate; ovary and silique densely pubescent; herbage pubescent with minute dendritic hairs A. pulchra Petals (4.5) 5-8 (9) mm long; petal limb ascending to erect; ovary and si- lique glabrous or sparingly hairy only; herbage glabrous or with simple or branched hairs 3 3(2). Pedicels spreading-ascending to erect; siliques ascending to erect 4 Pedicels divaricate to descending; siliques descending to pendulous 5 Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 291 4(3). Pedicels and siliques erect; siliques 3-6 cm long; plants wide- spread in northern and central Utah A. hirsuta Pedicels and siliques merely ascending; siliques 1-3 cm long; plants known only from Cache County A. nuttallii 5(4). Seeds biseriate; siliques 2-3 mm broad; plants of central and southern Utah A. pendulina Seeds uniseriate; siliques 1.5-2 mm broad; plants of Daggett and Uintah counties A. demissa 6(1). Lower leaves and/or stems pubescent exclusively with malpighian hairs (at least some); flowers white to pink; pedicels and siliques erect; plants widespread A. drummondii Lower leaves and /or stems glabrous or variously pubescent but not exclu- sively of malpighian hairs only, or, if so, otherwise various; flowers white, pink, lavender, or purple (cream in A. glabra); pedicels and si- liques variously disposed but sometimes erect 7 7(6). Basal leaves more or less hirsute and usually ciliate with long simple or forked hairs, not both hirsute and with dendritic hairs on the blade surfaces .... 8 Basal leaves more or less densely pubescent with dendritic (rarely malpig- hian) hairs, rarely also with a few long simple or forked hairs along the leaf bases 11 8(7). Flowers cream to white or pinkish; pedicels and siliques ascending to erect 9 Flowers pink to lavender or purple, rarely white; pedicels and siliques spreading to descending 10 9(8). Stigmas expanded. 0.8-1.1 mm broad, much wider than the style base; out- er sepals not gibbous at the base; petals cream to rarely pinkish; siliques not strongly compressed; plants from Utah County northward A. glabra Stigma not obviously expanded, 0.3-0.6 mm broad, not much wider than the style base; outer sepals gibbous at the base; petals white to pink; siliques definitely compressed; plants of a broader distribution A. hirsuta 10(8). Stems solitary or few, 2.5-6 dm tall; cauline leaves 1-4 cm long; seeds bise- riate A. fendleri Stems several to many, 1-3 dm tall; cauline leaves 0.5-1 cm long; seeds uniseriate A. demissa 11(7). Fruiting pedicels (but not necessarily the siliques) ascending to erect; siliques erect to spreading or even curved-pendulous 12 Fruiting pedicels spreading to descending, pendulous, or appressed down- ward along the axis of the raceme 16 12(11). Leaves, stems, and pedicels (and sometimes even the siliques) hoary with minute soft hairs; plants of central western Utah A. shockleyi Leaves variously pubescent; stems and pedicels glabrous or puberulent, or the stems hairy near the base only; plants of various distributions 13 13(12). Pedicels erect; siliques erect or steeply ascending; plants 0.5-2.5 dm tall, of alpine sites at high elevations A. lyallii Pedicels spreading-ascending to ascending; siliques spreading to more or less pendulous; plants (2) 2.5-5 dm tall or more, of moderate to low ele vations 14 292 Great Basin Naturalist Vol. 37, No. 3 14(13). Stems usually solitary, 3-10 dm tall (usually pubescent with malpighian hairs); cauline leaves 10 or more, well developed, closely positioned, and often overlapping A. confinis Stems usually more than one, 2.5-5 dm tall; cauline leaves usually fewer than 8, poorly developed, commonly widely spaced and only the lower- most overlapping 15 15(14). Stems arising from between the basal rosette and a tuft of ascending leaves; basal leaves mostly 2.5-5 cm long or more A. selbyi Stems arising from the basal rosette, a secondary tuft of leaves lacking or poorly developed; basal leaves mostly 0.5-2.5 cm long A. microphylla 16(11). Lowermost leaves entire, poorly developed, smaller than the main cauline ones; cauline leaves linear to narrowly oblong; siliques more or less finely pubescent A. pulchra Lowermost leaves in a rosette, more or less well developed, usually oblan- ceolate to spatulate and broader than the main cauline ones, often toothed; siliques not or rarely pubescent (except in A. puberula) 17 17(16). Stems usually 3 or more, arising between basal rosette and an ascending- erect tuft of leaves; lowermost leaves more or less toothed; pedicels spreading-descending; siliques more or less pendulous A. perennans Stems usually solitary, rarely 3 or more but then branches not arising from between a basal rosette and a tuft of ascending leaves, lowermost leaves entire or toothed 18 18(17). Leaves, stems, pedicels and fruit densely pubescent with minute hairs; plants of north central and perhaps western Utah A. puberula Leaves, stems and pedicels variously pubescent but seldom if ever all of them hairy at once 19 19(18). Stems usually numerous, less than 2 dm tall; flowers bright pink to lav- ender or purple; plants of high elevations in Wasatch and Uinta moun- tains A. lemmonii Stems usually solitary or, if more, over 3 dm tall; flowers pale pink to lav- ender or white; plants of lower to moderate elevations, not of alpine sites 20 20(19). Petals 8-12 mm long or more; pedicels and siliques merely spreading; plants evidently rare in northern Utah A. sparsiflora Petals 4-9 mm long; pedicels and siliques spreading-descending to reflexed 21 21(20). Basal leaves thickish, usually entire; pedicels merely spreading-descending ... A. lignifera Basal leaves thin, often sharply toothed; pedicels curved-decending to re- flexed; our most common species A. holboellii Arabis confinis S. Wats. [Turritis brachij- Rupr.; A. oblanceokita Rydb.; A. pmtincola carpa Torr. & Gray; A. drummondii var. Greene; A. neniophila Greene; A. dacotica brachycarpa (Torr. & Gray) A. Gray; A. Greene; A. brevisilu]ua Rydb.; A. stokesiae confinis var. brachycarpa (Torr. & Gray) Rydb.; A. drummondii var. pratincola Wats. & Coult.; A. divaricarpa A. Nels.; A. (Greene) Hopkins]. Plants biennial or short- brachycarpa (Torr. & Gray) Britton, not lived perennial; stems (2.5) 3-9 (10) dm tall. Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 293 solitary, or, when more than one, not aris- ing from between basal rosette and a tuft of ascending leaves; herbage subglabrous to pubemlent with malpighian or 3- to sever- al-rayed hairs at least on lower stems and leaves; basal leaves (1.3) 2-7 cm long, 0.3- 0.9 cm wide, oblanceolate, subentire to den- tate, usually acute; cauline leaves auriculate, numerous, mostly 2.2-6.5 cm long and 0.3- 0.8 cm wide; pedicels 3-14 mm long, glabrous, spreading-ascending or even (rarely) descending; sepals 3-5 mm long, sparsely pubescent or glabrous; petals 6-10 mm long, pink to lavender, spatulate, erect or ascending; siliques 20-85 mm long, 1.5- 2.5 mm wide, divaricate to subpendulous, valves glabrous, nerved to the middle or above; styles short or obsolete; seeds uni- or biseriate.— Moist sites at middle elevations in the mountains of Cache, Carbon, Gar- field, Millard, Rich, Salt Lake, Sevier, Sum- mit, Utah, Washington, Wayne, and Weber counties, and to be sought elsewhere; Alaska and southern Yukon east to the At- lantic and south to California and Colorado. Our material is var. confinis. We are reluctantly taking up the name Arabis confinis for one of North America's more common species of the genus and placing the long-used name, A. divaricarpa A. Nels., in synonymy. The nomenclatural problems surrounding A. confinis were in- itially discussed by Fernald (Rhodora 5: 225- 231. 1903) and reviewed again by Hopkins (Rhodora 39: 63-98, 106-148, 155-186. 1937) and Rollins (1941). Fernald correctly noted that Watson based his description on, and cited representative specimens of, two dif- ferent kinds of Arabi, one being A. drum- mondii A. Gray, and the second A. brachy- carpa (Torr. & Gray) Britton. What Fernald did not realize, apparently, was that the name A. brachycarpa had already been used by Ruprecht in an 1869 pub- lication. Thus, Hopkins and Rollins took up the next available species name, A. divari- carpa, which was published by Nelson in 1900. Neither Hopkins nor Rollins at- tempted to typify the Watson name on one or the other of these elements (A. drwn- mondii or A. divaricarpa) and simply placed A. confinis in synonymy under both. We have reviewed the original Watson publication (Proc. Amer. Acad. Arts 22: 466. 1887) and have noted that under the cur- rent International Code, it is clear that Watson was proposing a new name for Tur- ritis brachycarpa Torr. & Gray (not Arabis brachycarpa Rupr.), and, in spite of Wat- son's confused description and citation of specimens, one must accept A. confinis as a new name for T. brachycarpa and its type. Thus, A. confinis is the earliest available name for T. brachycarpa, and it is not a synonym of A. drummondii as proposed by Fernald, Hopkins, and Rollins. In addition to var. confinis, there is also Arabis confinis var. interposita (Greene) Welsh and Reveal, comb, nov., based on A. interposita Greene, Leafl. Bot. Observ. 2: 78. 1910. Arabis demissa Greene. [A. rugocarpa Os- terh. A. aprica Osterh. ex Nels. in Coult. and Nels.] Plants perennial, stems 1-3 dm tall, solitary or 2-5 from a simple caudex, these arising from between the basal rosette and a tuft of ascending leaves, subglabrous to hirsute with simple or forked hairs at least below; basal leaves 1.5-3.5 cm long, 0.2-0.5 cm wide, narrowly to broadly oblan- ceolate, entire, hirsute marginally and often on surface with simple hairs; cauline leaves sessile and non-auriculate (except in var. lanugida), mostly 0.5-1 cm long and 0.1-0.4 cm wide, oblong to lanceolate, hirsute to glabrous; pedicels 3-7 mm long, glabrous, slender, arched downward; sepals 2-3.5 mm long, sparsely pubescent; petals 4.5-6.5 mm long, white to pink, spatulate, erect or as- cending; siliques 20-40 mm long, 1.5-2 mm wide, pendulous, valves glabrous, nerved to about the middle; styles obsolete; seeds imiseriate.— Foothills and drainages in the Wasatch Plateau and eastern portion of the Uinta Mountains, Utah; southern Wyoming and central Colorado. The species is closely allied to both A. fendleri and A. penduUna, differing from the former in smaller size of stems and leaves, from the latter in the more slender siliques, and from both of them in the uniseriate seeds. 294 Great Basin Naturalist Vol. 37, No. 3 1. Cauline leaves auriculate; basal leaves, at least the outer ones, oblanceolate .... var. lanugida Cauline leaves merely sessile, not auriculate; basal leaves merely linear to narrowly oblanceolate var. russeola Var. lanugida Rollins. Low hills and drainages, sagebrush and juniper commu- nities, at about 1500 to 1700 m elevation, Daggett and Emery counties; southern Wyoming. Var. russeola Rollins. Low hills and val- leys in juniper communities, at about 1700 m elevation, Daggett and Uintah coimties; southern Wyoming. These two variants are only weakly dis- tinguishable and may not deserve separa- tion. The var. demissa is restricted to the mountains of central Colorado. Arabis drummondii A. Gray. [Tiirritis stricta Graham, not A. stricta Huds.; Strep- tanthus angustifolins Nutt. ex Torr. & Gray, not A. angustifolia Lam.; A. connexa Greene; A. oxyphylla Greene; A. albertina Greene; A. philonipha A. Nels. ex Rydb.; Tiirritis drummondii (A. Gray) Lunell; A. drummondii var. connexa (Greene) Fern.; A. drummondii var. oxyphylla (Greene) Hop- kins]. Plants biennial or short-lived per- ennial, the stems (0.8) 1.2-9 (9.7) dm tall, solitary or 2 to several from a simple or branched caudex, not arising from between basal rosette and tuft of ascending leaves, usually glabrous throughout or strigose with malpighian hairs at base only; basal leaves (1.1) 1.5-7 (8) cm long, 0.2-1.2 cm wide, ob- lanceolate, entire, subglabrous or pubescent with malpighian hairs; cauline leaves usual- ly longer than intemodes, numerous, (1) 1.5- 6 cm long, 0.2-1.5 cm wide, oblong to lan- ceolate, usually entire, sessile and auricu- late, usually glabrous; pedicels 7-15 mm long in fruit, erect, glabrous; sepals (3) 3.3- 5.7 mm long, glabrous; petals 6.5-10.5 mm long, white to pink, spatulate, ascending to erect; siliques (27) 35-95 (110) mm long, (1.2) 1.5-2 (3) mm wide, erect, valves glabrous, nerved to middle or above; styles short or obsolete; seeds biseriate.— Wood- lands and meadows at moderate to high ele- vations, in Beaver, Cache, Duchesne, Gar- field, Grand, Iron, Juab, Kane, Millard, Piute, Rich, Salt Lake, San Juan, Sanpete, Sevier, Summit, Uintah, Wasatch, Washing- ton, Wayne, and Weber counties; Alaska and Yukon east to the Atlantic and south to California, Arizona, and New Mexico. Arabis fendleri (S. Wats.) Greene. Plants perennial or rarely biennial, stems (1.4) 2-6 dm tall, solitary or 2-5 or more from a simple, or rarely branched, caudex, not aris- ing from between the basal rosette and a tuft of ascending leaves, variously hairy to subglabrous with coarse simple or branched hairs at base only; basal leaves (1) 1.5-6 cm long, (0.2) 0.3-1.3 (1.5) cm wide, oblanceol- ate, coarsely dentate to subentire, ciliate with simple or forked hairs, surfaces sim- ilarly hairy to subglabrous; cauline leaves usually longer than internodes, several to numerous, (0.5) 1-3.5 (4) cm long, 0.3-0.7 cm wide, oblong to lanceolate, entire or rarely toothed, sessile and auriculate, glabrous or lower ones hairy; pedicels 4-18 mm long in fruit, ascending-spreading to de- scending, glabrous; sepals 2.3-5 mm long, pubescent or glabrous, more or less gibbous; petals 5-7 (8) mm long, white to pink, spatulate, ascending to erect; siliques 30-46 (60) mm long, 1.5-2.1 (2.5) mm wide, divari- cate to curved pendulous, valves glabrous, nerved to middle or above; styles obsolete; seeds biseriate.— Woodlands, shrublands, and grasslands from low to moderate elevations, in eastern and southern Utah; Wyoming and Colorado west to Nevada, and south to Texas and Mexico. 1. Basal leaves dentate, definitely oblanceolate; petals usually pink A. fendleri var. fendleri Basal leaves entire, narrowly oblanceolate; petals white A. fendleri var. spatifolia Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 295 Var. fendleri. [A. holhoellii var. fendleri S. Wats, in Gray] Garfield, Millard, Piute, San Juan, Sevier, and Washington counties; range of the species. Var. spatifolia (Rydb.) Rollins. [A. spati- folia Rydb.] Known only from Daggett County; Wyoming, Colorado, and New Mexico. Arabis glabra (L.) Bernh. Plants biennial or rarely perennial, stems 3-8 (10) dm tall, solitary or 2 or 3, from a taproot, not aris- ing from between the basal rosette and a tuft of ascending leaves, usually hirsute with simple or forked hairs, rarely with ap- pressed dendritic hairs, at base only; basal leaves 3-10 (15) cm long, 0.8-3.5 (5) cm wide, oblong-oblanceolate to spatulate, si- nuate-dentate to entire, more or less ciliate with forked or dendritic hairs, surfaces (es- pecially the veins) hirsute with simple or forked hairs; cauline leaves usually longer than internodes (at least below), numerous, 1.5-9 (12) cm long, 0.4-2.3 (3.5) cm wide, lanceolate, denticulate to entire, sessile and auriculate, glabrous at least above; pedicels 4-12 mm long in fruit, erect, often ap- pressed, glabrous; sepals 3.3-4.5 mm long, glabrous, often tinged reddish-purple; petals 4.7-6 (7) mm long, cream or rarely pinkish, narrowly spatulate, ascending to erect; si- liques 40-90 mm long, 1-1.5 mm wide, strictly erect, valves glabrous, nerved to the middle or above; styles about 0.5 mm long, stigma 0.8-1.1 mm broad; seeds more or less biseriate.— Sagebrush, oak, and woodland communities at low to moderate elevations in northern Utah; widespread in North America; Europe. 1. Pvibescence of stem base appressed, of several-branched hairs A. glabra var. furcotipilis Pubescence of stem base hirsute with simple or merely forked hairs A. glabra var. glabra Var. furcatipilis Hopkins. Cache and Salt Lake counties, and to be expected else- where; Utah and California. Var. glabra. [Turritis glabra L.; A. per- foliata Lam.; T. macrocarj)a Nutt. ex Torr. and Gray; A. macrocarpa (Nutt.) Torr.] Box Elder, Millard, Tooele, Uintah, Utah, Wasatch, and Weber counties; range of the species. Arabis hirsuta (L.) Scop. [Turritis hirsiita L.] Plants biennial or perennial, stems (0.9) 1.5-6 (7) dm tall, solitary or more com- monly 2-6 or more, from a simple or branched caudex, not arising from between the basal rosette and a tuft of ascending leaves, hirsute with simple or forked hairs, at base at least; basal leaves (0.8) 1.5-6 (8) cm long, 0.3-1.8 (3) cm wide, elliptic to ob- long or oblanceolate, entire or more or less dentate, ciliate with simple or forked hairs. surfaces glabrous or more or less hirsute with simple or forked hairs; cauline leaves usually longer than internodes at least be- low, numerous, (0.6) 1-5 cm long, 0.2-2 cm wide, oblong to lanceolate, toothed to en- tire, sessile and auriculate, rarely merely sessile, hirsute or glabrous; pedicels 3-18 mm long in fruit, erect, appressed, glabrous; sepals 2.2-4 (4.5) mm long, glabrous or spar- ingly hairy, seldom tinged reddish-purple; petals 3.2-7 (9) mm long, white or pink, ob- long to spatulate, ascending to erect; si- liques 30-55 (60) mm long, 1-1.5 (2) mm wide, valves glabrous, erect, nerved to above the middle; styles 0.3-1 mm long, stigma 0.3-0.7 mm broad; seeds uniseriate.— Moist woods at moderate to high elevations; Alaska and Yukon east to the Atlantic and south to California, Arizona, and New Mex- ico; circumboreal. 1. Flowers 5-9 mm long; siliques often more or less divergent ....A. hirsuta var. glabrata Flowers 3-5 mm long; siliques strictly erect A. hirsuta var. pijcnocarpa Var. glabrata Torr. & Gray. [Turritis pestris Nutt. ex Torr. & Gray; A. pycno- spathulata Nutt. ex Torr. & Gray; A. ru- carpa var. glabra (Torr. & Gray) Hopkins]. 296 Great Basin Naturalist Vol. 37, No. 3 Raft River Mts., Box Elder Co. (US), and Deep Creek Mts., Juab Co., Utah; British Columbia south to California, east to Wyoming and Utah. Var. pycnocarpa (Hopkins) Rollins. [Tur- ritis ovata Pursh?; A. hirsiita var. ovata (Pursh) Torr. & Gray?; A. pycnocarpa Hopkins] Mountains in northern and central Utah; Daggett (US), Duchesne, Emery, Gar- field, Kane, Piute, Salt Lake, Sevier (US), Uintah (US), Utah, and Wasatch counties; widespread in North America. The name ''ovata' may have priority over the name ''pycnocarpa" , but, until the type of the Pursh name can be discovered, the description is inadequate to know exactly which species of Arahis Pursh had in mind, and thus the name cannot be ap- plied to our plants. Arahis holboellii Homem. Plants biennial or perennial, stems (1) 2-11.5 dm tall, soli- tary or less commonly 2-6, from a simple or branching caudex, not arising from between the basal rosette and a tuft of ascending leaves, pubescent with appressed or spread- ing hairs, at least at base; basal leaves (1) 1.5-5.5 cm long, 0.2-8 mm wide, elliptic to oblanceolate, entire to dentate, with den- dritic hairs on margins and usually on sur- faces, rarely with some simple or forked hairs near petiole base; cauline leaves usual- ly longer than internodes, at least below; numerous, 1.2-6.5 cm long, 0.2-0.9 cm wide, oblong to lanceolate, entire or some toothed, sessile and auriculate, pubescent to glabrous; pedicels 5-23 mm long in fruit, re- flexed to loosely descending, glabrous or pubescent; sepals 2.9-5 mm long, pubescent or glabrous, often tinged reddish; petals 4-9 (10) mm long, pink to lavender or white, spatulate, erect or ascending; siliques (20) 25-70 mm long, 1-2.5 mm wide, reflexed to loosely pendulous, the valves glabrous, nerved to below or above middle; style ob- solete or very short, stigma not much en- larged; seeds biseriate.— Foothills and moun- tains from lower to moderate elevations among shrubs or in woods; Alaska and Yu- kon east to Greenland and south to Califor- nia, Utah, Colorado, and Nebraska. 1. Pedicels gently curved downward; pods pendulous, often somewhat curved in- ward A. holboellii var. pinetonim Pedicels abruptly curved at the base, deflexed; siliques strictly reflexed to de- scending and often straight A. holboellii var. secunda Var. pinetorum (Tidestr.) Rollins. [A. pi- netonim Tidestr.] Widespread in Beaver, Box Elder, Cache, Daggett, Duchesne, Emery, Garfield, Juab, Kane, Millard, Salt Lake, Sevier, Summit, Uintah, Utah, Wasatch, and Washington counties; Sas- katchewan to British Columbia south to California, Colorado, and Nebraska. Var. secunda (Howell) Jeps. [A. retro- fracta Grab.; Turritis retrofracta (Grab.) Hook.; Streptanthus virgatus Nutt. ex Torr. and Gray; A. secunda Howell; A. arcuata var. secunda (Howell) Robins.; A. holboellii var. retrofracta (Grab.) Rydb.; A. rhodantha Greene; A. exilis A. Nels.; A. tenuis Greene; A. lignipes A. Nels.; A. consanguinea Greene; A. kochii Blankinship, not Jordan; A. sparsiflora var. secunda (Howell) Piper; A. caduca Greene; A. macdougalii Rydb.] Widespread and common in Beaver, Cache, Daggett, Davis, Duchesne, Grand, Millard, Piute, Salt Lake, San Juan, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, Washington, and Weber counties; Quebec to Alaska and British Columbia, south to California and Colorado. The name var. secunda is taken up for our most common species of the genus as the varietal rank was first occupied by "se- cunda' (1895), one year ahead of Rydberg's publication of var. retrofracta. Arahis lemmonii S. Wats. [A. canescens var. latifolia S. Wats.; A. latifolia (S. Wats.) Piper; A. bracteolata Greene; A. egglestonii Rydb.] Plants perennial, the stems 0.5-2 dm tall, several to many from a branching cau- dex, often with tufts of ascending leaves, pubescent to glabrous; basal leaves 0.8-2 (2.5) cm long, 0.2-0.5 cm wide, spatulate to oblanceolate, usually entire, marginally pub- escent with dendritic hairs, rarely with some hairs simple or forked on the petiole Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 297 bases, surfaces densely pubescent with den- dritic hairs; cauhne leaves shorter or longer than internodes, few, 0.4-1.5 cm long, 0.1- 0.3 cm wide, elliptic-oblong to lanceolate, entire, sessile and auriculate (at least some), pubescent to glabrous; pedicels 2-6 mm long in fruit, ascending to spreading, glabrous or pubescent; sepals 2-3.5 mm long, glabrous or pubescent, often purplish; petals 4-5.5 (6) mm long, pink to lavender, spatulate, erect to ascending; siliques mostly 20-50 mm long, (1.5) 2-2.5 (3) mm wide, as- cending, spreading or somewhat pendulous, valves glabrous, nerved to middle; seeds uniseriate.— Mountain summits at high ele- vations, mostly 3300-4000 m elevation, in Duchesne, Summit, and Salt Lake counties; Alaska and Yukon south to California, Colo- rado, and Montana. Our plant is var. lemmonii. Arabis lignifera A. Nels. Plants perennial, the stems 1.5-6 dm tall, solitary or 2-5 from a simple or branching caudex, not arising from between basal rosette and a tuft of as- cending leaves, more or less densely pub- escent with dendritic hairs, at least below; basal leaves 1.5-4 (5) cm long, 0.3-1 cm wide, spatulate to oblanceolate, entire or nearly so, densely pubescent with dendritic hairs, rarely with long simple or forked hairs on the petiole base; cauline leaves from longer to shorter than internodes, nu- merous, 0.8-3.5 cm long, 0.2-1 cm wide, ob- long to lanceolate, sessile and auriculate, pubescent to glabrous; pedicels (3) 5-15 mm long in fruit, spreading to arching down- ward, glabrous or pubescent; sepals 3-5.6 mm long, pubescent or glabrate; petals 5.2- 9.1 mm long, pink to lavender, spatulate, erect or ascending; siliques 30-60 (65) mm long, 1.5-2 mm wide, more or less spread- ing-pendulous, the valves glabrous, nerved to middle or above; style short or obsolete, stigma not or only slightly expanded; seeds uniseriate to more or less biseriate.— Foot- hills and desert ranges at moderately low to low elevations in desert shrub, sagebriLsh, and juniper-pinyon woodlands in Beaver, Box Elder, Carbon, Daggett, Duchesne, Iron, Juab, Kane, Millard, Salt Lake, San Juan, Summit, Tooele, and Uintah counties; Wyoming and Idaho south to Nevada and Arizona. Arabis lyallii S. Wats. [A. drummondii var. alpina S. Wats.; A. oreophila Rydb.; A. artnerifolia Greene; A. densa Greene; A. multiceps Greene; A. drummondii var. hjallii (S. Wats.) Jeps.; A. drummondii var. oreophila (Rydb.) Hopkins]. Plants perennial, stems 0.3-2.5 dm tall, few to many from a branching caudex, often with tufts of as- cending leaves, glabrous; basal leaves 0.5-2.5 cm long, 0.2-0.6 cm wide, oblanceolate to spatulate, entire, glabrous or margin and surfaces sparingly pubescent with dendritic hairs; cauline leaves shorter to longer than internodes, few, 0.6-2 cm long, 0.2-0.6 cm wide, ovate to lanceolate or oblong, entire, sessile and at least some auriculate, usually glabrous; pedicels 4-10 (13) mm long, erect- ascending, glabrous; sepals 3.3-4.5 mm long, glabrous, not or only slightly gibbous, often purplish; petals 7-9 (10) mm long, bright pink to lavender, spatulate, erect or ascend- ing; silique 20-60 mm long, 2-3 mm wide, erect-ascending, valves glabrous, nerved to middle; style short, stigma not or slightly enlarged; seeds uniseriate or more or less biseriate.— Alpine tvmdra, krumholz, glacial moraines and among subalpine conifers, at 2600-3700 m elevation in Cache, Daggett, Duchesne, Millard, Salt Lake, Uintah, Utah, Wasatch, and Weber counties; Yukon and British Columbia south to California, Ne- vada, northern Utah, and western Wyoming. Our material is var. lyallii. Arabis microphylla Nutt. ex Torr. & Gray. Plants perennial, stems 0.5-3 (5) dm tall, several to many from a branching cau- dex, tufts of ascending leaves lacking or poorly developed, glabrous throughout or hairy below with simple or forked hairs; basal leaves 0.7-2.5 cm long, 0.2-0.5 cm wide, oblanceolate to spatulate, entire or rarely toothed, margins with dendritic or merely sessile and forked hairs, densely pu- bescent with minute dendritic hairs; cauline leaves often shorter than internodes, few, 0.6-2 cm long, 0.1-0.6 cm wide, oblong to lance-linear, entire or some toothed, sessile and auriculate, glabrous or lowermost pu- bescent; pedicels 4-13 (15) mm long, as- cending to spreading-ascending, glabrous or 298 Great Basin Naturalist Vol. 37, No. 3 pubescent; sepals 2.5-4 mm long, glabrous base only; style developed or obsolete, or pubescent, often purplish; petals 5-8 mm stigma not expanded; seeds uniseriate.— long, pink to lavender, spatulate, ascending Foothills and mountains at moderate eleva- to erect; siliques (20) 25-60 mm long, 1.2-2 tions; British Columbia to Montana, south mm wide, erect to spreading, nerved at to Oregon, Nevada, and western Wyoming. 1. Stems mostly 2.5-5 dm tall; pedicels and siliques spreading A. microphyUa var. macoiinii Stems mostly less than 2 dm tall; pedicels divaricate, the siliques erect A. microphyJla var. micropJiylla Var. macounii (S. Wats.) Rollins. [A. ma- coiinii S. Wats.; A. densicaulis A. Nels.] Mill Creek Canyon, Salt Lake County, and Uintah County, and to be expected else- where; Montana to British Columbia, south Idaho, Wyoming, and Utah. Var. microphyUa. [A. tenuicola Greene] Low to moderate elevations, often on rock outcrops or talus. Cache, Grand (?), Salt Lake, Utah and Weber counties; Montana to Washington, south to Nevada, Utah, and Wyoming. Arabis nuttallii Robinson in Gray. [A. spathulata Nutt. ex Torr. & Gray, not DC; Erysimum nuttallii (Robinson) Kuntze; A. hridgeri M. E. Jones; A. macella Piper] Plants perennial, stems 0.9-3 din tall, sever- al to many from a branching caudex, tufts of ascending leaves lacking, glabrous throughout or hirsute below with simple or forked hairs; basal leaves 1-4 cm long, 0.4- 1.2 cm wide, oblanceolate, usually entire, ciliate and hairy on lower surface (at least) with long simple and often with some forked hairs; cauline leaves shorter than in- ternodes, few, 0.5-2 cm long, 0.1-0.7 cm wide, elliptic to oblong, lanceolate or oblanceolate, entire, sessile or subsessile, not auriculate, hairy or the upper glabrous; pedicels 5-20 mm long, spreading-ascending, glabrous; sepals 3-4 mm long, glabrous or sparingly hirsute, usually green or cream; petals (5) 6-8 mm long, white or lavender, spatulate, more or less spreading; siliques (8) 12-20 mm long, 1-1.5 mm wide, erect to spreading, valves glabrous, nerveless or faintly nerved; style about 1 mm long, stigma not especially expanded; seeds unise- riate.— Meadows and shrublands from low elevations to mountain summits in Cache and Wasatch counties, and to be expected elsewhere; Alberta south to Wyoming and northern Utah. Arabis pendulina Greene. [A. setulosa Greene; A. diehlii M. E. Jones; A. neva- densis Tidestr.] Plants perennial, the stems 0.6-3.5 (4) dm tall, (one) several to many from a simple or branched caudex, arising from between basal rosette and a usually well-developed tuft of ascending leaves, hir- sute with simple hairs below or glabrous throughout; basal leaves 0.8-4 cm long, 0.1- 1 cm wide, spatulate to narrowly oblanceo- late, entire, ciliate and hairy on one or both surfaces with long simple hairs or totally glabrous; cauline leaves much shorter than internodes, few, 0.3-1.2 cm long, 0.1-0.6 cm broad, oblong to lanceolate, entire, sessile and usually not auriculate (but sometimes slightly clasping), hairy or the upper glabrous; pedicels 3-6 (10) mm long, arched downward, glabrous; sepals 2.5-3.6 (4) mm long, usually purplish; petals 4.8-6.8 mm long, pink to lavender, spatulate, erect or ascending; siliques (10) 13-40 mm long, 1.6- 2.3 (3) mm wide, pendulous, valves glabrous, nerved below middle; style obso- lete or very short, stigma not enlarged; seeds biseriate.— Open knolls to forest ground layer, from 1670 to 3400 m eleva- tion, in pinyon-juniper, ponderosa pine, and spruce-fir communities in Emery, Garfield, Kane, Piute, Sevier, Wayne, and Washing- ton counties; eastern and southern Nevada east to Utah. Plants from some sites have clasping leaves but otherwise seem not to differ in any other way from the more typical speci- mens with non-auriculate leaves. Arabis perennans S. Wats. [A. arcitata var. perennans (S. Wats.) M. E. Jones; A. gracilentu Greene; A. eremophila Greene; A. Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 299 recondita Greene; A. aii^^ulata Greene ex Woot. & Standi.] Plants perennial, stems 0.9-5.5 (6) dm tall, (one) several to many from simple or branching herbaceous to woody caudex, arising from between basal rosette and a tuft of ascending leaves, pu- bescent with dendritic hairs on margin and on surfaces, rarely with simple or forked hairs along petiole base; cauline leaves lon- ger than internodes at least below, several, 0.7-4 cm long, 0.2-0.8 cm wide, oblong to lanceolate, entire to toothed, hairy or the upper glabrous; pedicels 4-24 mm long, spreading to arched downward, glabrous or pubescent; sepals 3-4 (4.5) mm long, often purplish, usually dendritic hairy; petals 5-7 (9) mm long, pink to lavender, spatulate, erect or spreading; siliques (20) 27-55 (60) mm long, 1.2-2 mm wide, spreading to pendulous, valves glabrous, nerveless or nerved at the base; style obsolete or very short, stigma not enlarged; seeds unise- riate.— Warm desert sliRib, pinyon-juniper, sagebrush, ponderosa pine, and oakbrush communities at low to moderate elevations in -Beaver, Carbon, Emery, Garfield, Grand, Kane, Millard, Piute, San Juan, Sanpete, Se- vier, Wayne, and Washington counties; Col- orado and New Mexico to Nevada, Califor- nia, and Baja California. Our specimens are far from uniform, with some approaching A. Ugnifera on the one extreme, and A. selhiji on the other. Arabis puherula Nutt. ex Torr. & Gray. [A. beckwithii S. Wats.; Erysimum pu- henilum (Nutt.) Kuntze; A. siibpinnatifida var. beckwithii (S. Wats.) Jeps.; A. arida Greene; A. lignipes var. impar A. Nels.; A. sabulosa M. E. Jones; A. sabulosa var. frig- ida M. E. Jones; A. sabulosa var. colorata M. E. Jones; A. subpinnatifida var. ijnpar (A. Nels.) Rollins] Plants perennial or in- frequently biennial, stems (0.7) 1-3 dm tall or more, solitary or few from a simple cau- dex, not arising from between a basal ro- sette and a tuft of ascending leaves, pu- bescent throughout with dendritic hairs, rarely glabrous above; basal leaves 1-2.5 (3) cm long, 0.2-0.6 cm wide, oblanceolate, en- tire or toothed, pubescent with minute den- dritic hairs; cauline leaves usually longer than internodes, several to many, 1-3 cm long, 0.2-0.8 cm wide, toothed to entire, sessile and at least some auriculate, hairy like the basal leaves; pedicels 2-7 mm long, arched downward, pubescent; sepals 3.5-6 mm long, dendritic hairy; petals 7-11 mm long, pink to white, spatulate, erect or as- cending; siliques (25) 30-50 (60) mm long, 2-3 mm wide, pendulous to reflexed, valves copiously hairy to glabrate, nerved to below middle; style obsolete, stigma not enlarged; seeds uniseriate.— Oakbrush community at moderate elevations in Salt Lake County {Rose 1048, BRY), and to be sought else- where; Washington and Idaho south to Cal- ifornia and Nevada. A closely related species, Arabis cobrensis M. E. Jones, is to be sought in extreme northern Utah. The species occurs just north of the state line in Uinta Countv, Wyoming, and in northeastern Elko County, Nevada, but as yet it is not known from Utah. Arabis cobrensis is a well-defined perennial with a well-branched caudex, the basal leaves are only 0.1-0.3 cm wide, the few cauline leaves are remote and only 0.1-0.3 cm wide, and the petals are 4-6 mm long. It is to be sought in Daggett and Box Elder counties. Arabis pulchra M. E. Jones. Plants pe- rennial, stems 1.5-6 dm tall, solitary or sev- eral from a branching herbaceous to woody caudex, not arising from between basal ro- sette and a tuft of ascending leaves, pu- bescent with dendritic hairs throughout or glabrous above; basal leaves in poorly de- veloped rosettes, 1-6 cm long, 0.2-0.6 cm wide, narrowly oblanceolate to spatulate, entire, densely hairy with minute hairs; cauline leaves shorter or longer than inter- nodes, many, 1.2-6 cm long, 0.1-0.5 cm wide, entire, sessile and mostly not auricu- late, hairy like the basal leaves; pedicels 5- 18 mm long, recurved to pendulous in fruit, pubescent to glabrate; sepals 5-8.2 mm long, more or less gibbous, dendritic- pubescent, often purplish; petals 9-18.5 mm long, pale pink to white or pink to lavender or purple, spatulate to obovate-spatulate, spreading or ascending to erect; siliques 35- 55 (65) mm long, 1.8-2.5 (3) mm wide, pendulous to reflexed, valves copiously hairy to glabrate, nerved to middle or 300 Great Basin Naturalist Vol. 37, No. 3 above; style obsolete or very short, stigma not expanded; seeds biseriate.— Desert shrub, pinyon-juniper, and sagebrush com- munities from 650 to 2000 m elevation in the southern two-thirds of Utah; common from Colorado and New Mexico west to California and Baja California. 1. Flowers lavender to purple, 9-11 mm long; petals ascending to erect A. pulchra var. munciensis Flowers pale pink or white, less commonly lavender, 10-18.5 mm long; petals spreading A. pulchra var. pollens Var. munciensis M. E. Jones. Desert shrublands in Beaver, Juab, Iron, Kane, Mil- lard, Tooele, and Washington counties; Ne- vada and California. Var. pollens M. E. Jones. [A. fomwso Greene] Desert shrublands to pinyon-juniper zone in Duchesne, Emery, Garfield, Grand, Kane, San Juan, Uintah and Wayne coun- ties; Colorado, New Mexico, and Arizona. Arobis selbyi Rydb. Plants perennial, the stems 1.5-4 (5) dm tall, usually several from a simple or branching caudex, arising from between basal rosette and an ascending tuft of leaves, pubescent below with dendritic or forked hairs; basal leaves 1.5-4 (6) cm long, 0.2-1 cm wide, oblanceolate to spatulate, usually entire, densely to sparsely pubescent with dendritic hairs on margin and surfaces, sometimes with simple or branched hairs along petiole base; cauline leaves usually shorter than internodes, few to several, 0.3- 2.5 (3) cm long, 0.1-0.4 cm wide, narrowly oblong to lanceolate, entire, hairy or glabrous; pedicels 4-15 mm long in fruit, spreading-ascending, glabrous or pubescent; sepals 2.6-3.5 (4) mm long, often purplish, usually dendritic hairy; petals 5-7 (8) mm long, pink to lavender, spatulate, erect or ascending; siliques 30-55 (60) mm long, 1.2- 1.8 mm wide, spreading-ascending to more or less pendulous, valves glabrous, nerved at base or nerveless; style obsolete or very short, stigma not enlarged; seeds unise- riate.— Sagebrush and pinyon-juniper com- munities mostly between 1500 and 2200 m elevation in Carbon, Duchesne, Emery, Gar- field, Grand, San Juan, Uintah, and Utah counties; western Colorado and north- western New Mexico. Only the slightly more ascending pedicels and usually entire or subentire leaves serve to distinguish this entity from Arobis per- ennans. Possibly, it would be treated better at an infraspecific level within that species. Arobis shockleyi Munz. Plants perennial or infrequently biennial, stems 1.2-3.5 (4) diu tall, solitary or 2-4 from a simple or branching caudex, not arising from between a basal rosette and a tuft of ascending leaves, densely pubescent throughout with minute dendritic hairs, often somewhat less densely so above; basal leaves 1.6-3 (3.5) cm long, 0.5-0.9 cm wide, oblanceolate to spatulate, entire, densely pubescent through- out with dendritic hairs only; cauline leaves much longer than internodes, numerous, 1.2- 3 cm long, 0.5-1.2 cm wide, ovate-lanceo- late to lance-attenuate, entire or nearly so, sessile and auriculate, hairy as on basal leaves; pedicels 12-17 mm long in fruit, as- cending, pubescent; sepals 5.5-7 mm long, often reddish, dendritic hairy; petals 7.5-10 mm long, pink to lavender, spatulate, erect or ascending; siliques 42-65 mm long, 1.2- 1.8 mm wide, ascending to spreading, valves glabrous, nerved to about middle; style up to 0.8 mm long, stigma not much expanded; seeds biseriate.— Desert shrublands and pi- nyon-juniper communities mostly 1430 to 1900 m elevation in Juab, Millard, and Tooele counties; north-central and western Nevada, and in eastern and southern Cali- fornia. Arobis sporsifloro Nutt. ex Torr. and Gray. Plants perennial, stems 2.3-10 dm tall, solitary or less commonly 2 or more from a simple or branching caudex, not arising from between the basal rosette and a tuft of ascending leaves, pubescent with appressed or spreading hairs at least below; basal leaves 3-7 (9) cm long, 0.3-0.6 (0.9) cm wide, oblanceolate, entire or dentate, pub- escent with coarse dendritic hairs usually on both surfaces; cauline leaves longer than in- Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 301 ternodes, numerous, 1.5-6 cm long or more, 0.3-0.7 cm wide, oblong to lanceolate, en- tire or the lower toothed, sessile and auricu- late, hairy like the basal leaves or wholly glabrous; pedicels 5-15 mm long, ascending- spreading, glabrous or pubescent; sepals 4.7- 6 mm long, dendritic hairy; petals 8-12 mm long, pink to lavender or purple, spatulate, ascending to erect; siliques 60-100 (120) mm long, 1.2-2 mm wide, ascending to curved- descending; style obsolete or nearly so, stigma not expanded; seeds imiseriate.— Foothills and canyons at low to moderate elevations in oakbrush, sagebrush, and streamside forest communities of northern Utah; British Columbia and Alberta south to California, Utah, and Wyoming. 1. Basal leaves entire, narrowly oblanceolate; pedicels ascending-spreading A. sparsiflora var. sparsiflora Basal leaves dentate, oblanceolate; pedicels spreading A. sparsiflora var. subvillosa Var. sparsiflora. [A. peramoena Greene; A. sparsiflora var. peramoena (Greene) Rol- lins] Foothills and stream courses in Cache, Salt Lake, and Weber counties; California, Oregon, and Idaho. Var. subvillosa (S. Wats.) Rollins. [A. ar- ciiata var. subvillosa S. Wats.; A. perelegans Nels. in Coult. and Nels.] Foothills and mountain slopes in Davis, Uintah, and We- ber counties; Washington to Montana south to Wyoming, Utah, Nevada, and California. Armoracia Gaertn. Horse-Radish Plants glabrous perennials from tuberous- thickened taproots; leaves alternate, simple, crenately toothed or lobed, petiolate to sub- sessile, not auriculate; flowers racemose, the pedicels ascending, not subtended by bracts; sepals 4, deciduous; petals 4, white, not emarginate; stamens 6, at least the 2 short stamens subtended by glands; style short, the capitate stigma hemispheric; fruit a silicic, bilocular, about as long as broad, obovoid-ellipsoid, the valves with an incon- spicuous midnerve; seeds apparently never developing. A genus of 3 species of Eurasia. Armoracia rusticana Gaertn. [Cochlearia armoracia L.; Nasturtium armoracia (L.) Fries; Rorippa armoracia (L.) A. S. Hitchc; Radicula armoracia (L.) A. Gray; Armoracia armoracia (L.) Cockerell] Plants perennial, stems 6-10 dm tall or more, few to many arising from summit of root crown; basal leaves oblong to oblong-lanceolate, blades 15-50 cm long, 10-15 cm wide, base cordate to rounded, petioles 0.6-4 dm long or more; cauline leaves reduced and only short petiolate to subsessile upwards, lanceolate to elliptic and crenately toothed to lobed, glabrous; pedicels mostly 8-11 mm long, as- cending in fruit; sepals 2-2.5 mm long, ca- ducous, greenish, glabrous; petals 4.2-4.5 (5) mm long, white, obovate-spatulate, surpass- ing sepals; silicles 3-6 mm long and about as wide, valves glabrous; styles 0.2-0.3 mm long, hemispheric stigma to 0.5 mm broad or more.— Cultivated and occasionally es- caping and persistent as in Cache, Salt Lake, Summit, Utah, and Weber counties, and most likely elsewhere in agricultural re- gions of the state; widespread in North America; introduced from Eurasia. Athysanus Greene Plants annuals with typically dendritic and with some simple hirsute hairs, arising from slender taproots; leaves alternate, simple, few-toothed, sessile, and not auricu- late; flowers racemose, pedicels recurved, not subtended by bracts; sepals 4, de- ciduous; petals 4, white, not emarginate; stamens 6; style short, stigma capitate; fruit a silicle, unilocular, about as broad as long, orbicular, compressed, valves nerveless; seeds solitary. A monotypic genus. Athysanus pusillus (Hook.) Greene. [Thy- sanocarpus pusillus Hook.; A. pusillus var. glabior S. Wats.] Plants annual, stems erect or ascending, usually branched from near base, 0.8-3 dm tall; leaves ovate to obovate or spatulate (especially the lowermost), den- 302 Great Basin Naturalist Vol. 37, No. 3 dritic hairy on both surfaces; pedicels re- curved, 1-4 mm long; sepals about 1 mm long, cream to purplish, glabrous; petals 1-2 mm long, white, oblong-spatulate, surpass- ing sepals; silicles 1.5-2.5 mm long and about as wide, valves hirsute with simple, imcinate hairs, emarginate at style; styles persistent, 0.2-0.3 mm long.— Dry sites at low elevations, not definitely known from Utah but to be expected in Washington County; British Columbia south to Califor- nia and Arizona. Barbarea R. Br. Plants glabrous to sparsely hirsute bien- nials or rarely annuals from taproots; leaves alternate, lyrate-pinnatifid to pinnately compound, cauline leaves auriculate-clas- ping and often falsely petiolate above clasp- ing base; flowers racemose, pedicels ascend- ing to erect, not subtended by bracts; sepals 4, deciduous; petals 4, yellow, truncate to rounded apically; stamens 6, filaments lack- ing glandular processes; style stout, abruptly contracted to capitate stigma; fruit a si- lique, many times longer than broad, linear, only slightly compressed, more or less con- tracted between the seeds, valves 1 -nerved; seeds numerous, uniseriate. About 20 species of the northern temper- ate regions of the world. Barbarea orthoceras Ledeb. [B. ameri- cana Rydb.; Campe orthoceras (Ledeb.) Hel- ler; B. orthoceras var. dolichocarpa Fern.] Plants erect, stems 1.5-10 dm tall, glabrous; basal leaves lyrate-pinnatifid to pinnately compound, rarely reduced to terminal lobe, mostly (1.5) 4-15 (20) cm long, 1-2.5 (4) cm wide, glabrous or petiole and lower lobes sparsely hirsute; cauline leaves reduced up- wards, auriculate-clasping; pedicels 2-4 mm long, glabrous, ascending; sepals 2.5-3.5 mm long, yellowish, glabrous; petals 4-5.5 mm long, yellow, spatulate-oblanceolate, ascend- ing-spreading; siliques (15) 20-50 mm long, 1.5-2.5 mm wide, erect or ascending, valves glabrous, prominently nerved to apex; style beaklike, (0.5) 1-2 mm long, abniptly con- tracted to stigma; seeds uniseriate, pitted.— Moist meadows, roadsides, and openings in woods from 1350 to 2500 m elevation in Box Elder, Salt Lake, Utah, Wasatch, and Washington counties, and to be expected al- most throughout the state; Alaska and Yu- kon east to the Atlantic and south to Cali- fornia, Nevada, Utah, and Colorado; Eurasia. Berteroa DC. Plants stellate-pubescent annuals or in- frequently winter annuals from taproots; leaves alternate and basal, simple, entire, re- duced upwardly and sessile, not auricula te; flowers racemose, pedicels erect-ascending, not subtended by bracts; sepals 4, de- ciduous; petals 4, white, deeply emarginate and often bilobed; stamens 6, the filaments lacking glandular processes; style long, slen- der, stigma capitate; fruit a silicle, 1-3 times longer than broad, compressed parallel to septum, valves 1-nerved or nerveless; seeds several. About 8 species of worldwide distribu- tion. Berteroa incana (L.) DC. [Ahjsswn in- canum L.] Plants erect, stems 3-10 dm tall or more, appressed stellate-hairy; basal leaves oblanceolate, 3-5 cm long, entire, petiolate, usually withered at anthesis; caul- ine leaves reduced upwards, sessile or short- petiolate below, stellate-hairy; pedicels erect or ascending, 4-10 mm long, stellate and sometimes more or less hirsute; sepals 2-3 mm long, greenish to whitish, stellate- hairy; petals 4-6 mm long, white, deeply bi- lobed; silicles 5-7 mm long, 2-3 mm wide, moderately inflated, stellate hairy; styles (1.5) 2-3 mm long, persistent.— Roadsides and other disturbed places, not definitely known from Utah but to be expected; wide- spread in North America; Europe. Brassica L. Plants glabrous or hirsute annuals from taproots; leaves alternate and basal, various- ly lobed to entire, basal ones often lyrate- pinnatifid, reduced upwardly and petiolate to sessile or auriculate; flowers racemose, pedicels erect or ascending, not subtended by bracts; sepals 4, deciduous; petals 4, yel- low; stamens 6, filaments lacking glandular processes; styles slender to thick, mostly well developed, stigma capitate; fruit a si- lique, several to many times longer than Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 303 broad, linear, terete or nearly so, often weedy adventives distinguished below. They more or less constricted, valves 1— 3-nerved, are: Brassica caitlorapa Pasquale (kohlrabi); apical portion producing a stout 1—3- B. napohmssica Mill, (rutabaga); B. oleracea nerved beak; seeds several to many, unise- L. var. botnjtis L. (cauliflower), var. capi- riate. tatci L. (cabbage), var. gemifera Zenker Several cultivated members of this genus (brussel sprout), and var. italica Plenck are present in our region in addition to the (broccoli); and B. rapa L. (turnip). 1. Cauline leaves sessile, auriculate-clasping, glaucous and entire or nearly so B. campestris Cauline leaves petiolate and not auriculate, or, if rarely so, falsely petiolate above the clasping base 2 2(1). Valves of fruit, and often the pedicels and raceme rachis, hirsute with course, spreading hairs; plants cultivated and escaping B. hirta Valves of fruit, pedicels, and raceme rachis glabrous; plants adventive weeds 3 3(2). Silique with a flattened, 2-edged or angular beak, valves and beak strongly 3 (5)-nerved B. kaber Silique with a cylindrical or rarely slender-conic beak, valves and beak with 1 (rarely with 2 additional delicate) nerves 4 4(3). Pedicels 2-6 mm long; siliques 1-2.5 cm long, 1-1.8 mm wide, ascending- appressed B. nigra Pedicels mostly (5) 10-15 mm long; siliques 2-4 cm long, 2-3 mm wide, as- cending to erect but not appressed B. jimcea Brassica campestris L. [B. rapa of authors, not L.; B. napus of authors, not L.] Plants erect, glabrous or with very few hairs, stems 2.5-10 dm long or more, simple or branched; basal leaves lyrate-pinnatifid, 5- 18 cm long, terminal lobe mostly 2-5 cm wide, crenate-dentate; lower cauline leaves similar to basal ones, reduced upwards, be- coming auriculate-clasping and dentate to entire; pedicels 7-20 mm long, slender, as- cending, glabrous; sepals 4.5-6 mm long, yellowish to greenish; petals 6-10 mm long, yellow; siliques 30-70 mm long, (1.5) 2.5-3.5 mm thick, beak 8-15 mm long, 1-nerved, valves conspicuously 1-nerved and with 2 more or less delicate lateral nerves, glabrous.— Cultivated fields, roadsides, and waste places in Summit, Utah, and Wash- ington counties, though probably cosmopoli- tan in the state; widespread in temperate regions of the world; adventive from Eu- rope. Brassica hirta Moench. [Sinapsifi alba L.; B. alba (L.) Rabenh., not Gilib.] Plants erect, pubescent with coarse, descending hairs at least below, the stems 2-10 dm tall. usually branched; basal leaves lyrate-pin- natifid, mostly 5-15 cm long, the terminal lobe 3-10 cm wide, obscurely crenate-den- tate; cauline leaves reduced upwards, usual- ly all petiolate, becoming merely lobed, not auriculate; pedicels 5-10 mm long, slender or stout, spreading, often hirsute; sepals 4-5 mm long, yellowish, glabrous; petals 7-10 mm long, yellow; siliques (20) 30-50 mm long, 3-4.5 mm wide, the beak 8-16 mm long, 3 (5) nerved, the valves conspicuously 3-nerved, hirsute.— Cultivated white mus- tard of commerce and rarely escaping, but potentially a noxious weed of cultivated land and to be expected in agricultural re- gions of the state; widespread in the west- ern United States; adventive or introduced from Europe. Brassica juncea (L.) Czernj. [Sinapsis jim- cea L.] Indian Mustard. Plants erect, glabrous or hirsute, the stems 3-10 dm tall or more, usually branched; basal leaves lyr- ate-pinnatifid, 8-25 cm long, terminal lobe 5-15 cm wide, crenate to dentate or lobed; cauline leaves reduced upwards, short-pe- tiolate to sessile, not auriculate-clasping; 304 Great Basin Naturalist Vol. 37, No. 3 pedicels 8-17 mm long, slender to stout, as- cending, glabrous; sepals 4-6 mm long, yel- lowish, glabrous; petals (5.5) 7-12 mm long, yellow; siliques 20-50 mm long, 2-3 mm wide, beak 6-12 mm long, 1 -veined, valves 1 (or lightly 3) nerved, glabrous.— A weed of disturbed soils (Washington County) but not yet common in the state; introduced from Asia. Brassica kaber (DC.) Wheeler. [Sinapsis arvensis L.; B. arvensis Rabenh., not L.; S. kaber DC] Charlock. Plants erect, pub- escent with coarse spreading hairs at least below, stems 3-10 dm tall or more, simple or branched; basal leaves lyrate-pinnatifid to merely dentate, 5-20 cm long, 3-10 cm wide; cauline leaves reduced upwards, short-petiolate or sessile, not auriculate-clas- ping, or, if apparently so, falsely petiolate or leaves sinuate-dentate; pedicels 2-6 mm long, ascending, stout, glabrous; sepals 4-5 mm long, yellowish, glabrous; petals 8-14 mm long, yellow; siliques 30-50 mm long, 2-3 mm thick, beak 7-15 mm long, 3- veined, valves 3 (5)-nerved, glabrous.— Road- sides, fields, and ditch banks in Summit, Utah, and Washington counties, but prob- ably in all coimties of the state; widespread in temperate portions of the world; adven- tive from Europe. Brassica nigra (L.) Koch in Rohling. [Si- napsis nigra L.] Black Mustard. Plants erect, glabrous or more usually sparsely to densely hirsute-hispid at least near base, stems 3-12 dm tall or more, usually branched; basal leaves lyrate-pinnatifid to lobed or serrate-dentate, 5-25 cm long, 2-15 cm wide; cauline leaves reduced upwards, short petiolate to sessile, not auriculate; pedicels 2-6 mm long, erect, stout, glabrous; sepals 3-4 mm long, yellowish, glabrous; petals (5) 7-12 (15) mm long, yellow; si- liques 10-25 mm long, 1-2 mm wide, beak 1-5 mm long, 1-veined, valves with 1 mid- nerve and two faint lateral ones, glabrous.— Roadsides, fields, and other disturbed places in Garfield, Juab, Salt Lake, San Juan, Utah, Wasatch, Washington, Weber, and perhaps all Utah counties; widespread in North America; adventive from Europe. This is a common weed of grainfields in northern Utah. Camelina Crantz False Flax Plants pubescent with forked or stellate hairs, annual, from taproots; leaves alter- nate, simple, entire, auriculate-clasping ba- sally; flowers racemose, pedicels ascending, not subtended by bracts; sepals 4, de- ciduous; petals 4, pale yellowish; stamens 6, filaments lacking glandular processes; styles slender, stigma capitate; fruit a silique, less than twice longer than broad, obovoid, somewhat compressed parallel to septum, valve 1 -nerved; seeds several per locule, biseriate. A Eurasian genus of 8-10 species. Camelina microcarpa Andrz. ex DC. Plants erect, stems (0.8) 1.5-8 dm tall or more, hirsute to subappressed with simple and forked to stellate hairs at least near base; leaves mainly cauline, basal mostly 1-7 cm long, entire or obscurely toothed, usual- ly withered by late anthesis; cauline leaves reduced upward, at least upper ones auricu- late; pedicels spreading-ascending, (6) 8-18 mm long, glabrous; sepals 2-2.7 mm long, often reddish, more or less villous; petals 3- 4 (5) mm long, white or nearly so, apex rounded; siliques 5-6.5 mm long, 3-4 mm wide, moderately inflated, glabrous; styles 1-2 (2.5) mm long, persistent.— Roadsides, foothills, gardens, and other disturbed moist to dry sites in Box Elder, Juab, Millard, Salt Lake, Summit, Tooele, Utah, and Weber coimties; widespread in North America; ad- ventive from Asia. Capsella Medic. Nom. Cons. Plants stellate-pubescent and often with coarse simple hairs also, annual, from tap- roots; leaves alternate or basal, simple, den- tate or variously toothed or lobed to entire, cauline ones auriculate-clasping; flowers racemose, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, white; stamens 6, the filaments lacking glandular processes; style short, stigma capitate; fruit a silicle, less than twice longer than broad, cuneate- obcordate in outline, compressed at right angles to the septum, valves reticulately veined, strongly keeled; seeds many per lo- cule. Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 305 A genus of European plants of perhaps 5 rubella Renter) and a white flower form species, now widely disseminated as weeds. with the style 0.5-1 mm long (C. thracica Capsella bursa-pastoris (L.) Medic. Velen.). [Thkispi bursa-pastoris L.; Bursa pastoris Cardamine L Weber in Wigg.; Bursa bursa-pastoris (L.) Bitter Cress Britton] Sheperd's Purse. Plants erect, stems ^^^^^^ glabrous or sparsely pubescent 1-5 dm tall, stellate pubescent and more or ^.^^ ^. ^^^ ^^.^^^ ^^^^^^ ^j^^^^^^^j ^^ less hirsute; basal leaves oblanceolate in ^^^j^j f^^^ ^^ ^^^^^ ^^ rhizomes; leaves al- outline 2.5-16 (20) cm long, 0 5-2.8 (4) cm ^^^^^^^ sometimes with basal rosettes, wide, lyrate-pinnatifid to merely toothed or ^-^ j^ ^^ pinnately compound, petiolate, subentire; cauline leaves much reduced up- ^^^ auriculate; flowers racemose or rarely wards sessile and auricailate; sepals 1.2-2.5 s^bcorymbose, pedicels spreading-ascending mm long, often reddish, pubescent or to ascending, not subtended by bracts; sep- glabrous; petals 2-4 mm long, white to ^j^ 4 deciduous; petals 4, white to pinkish; pinkish, apex rounded; silicles 4 5-8 mm ^^^^^^^ g^ filaments lacking glandular pro- long, 3-5 (6) mm wide, cuneate-obcordate, ^^^^^^. ^ j^ ^^^^^^^ ^^. capitate; fruit a si- glabrous; styles 0 3-0.6 (1) mm long, per- ^. ^^^^^^^ ^^ ^.^^^ ^ ^^^^ si^stent.- Disturbed sites at low to moderate ^road, sUghtly compressed parallel to sep- elevations in Cache, Garfield, Juab, Kane, ^^^^ ^^j^^^ obscurely 1 (3)-nerved or nerve- Salt Lake, Uintah, Utah, Wasatch Washing- j^^^. ^^^^^ ^^^^^^1 ^^ ^^^^ uniseriate. ton, and Weber counties (probably m all ^ ^^^^ ^f j^^ ^50 ^^.j^^ ^f ^^^ counties); widespread in North America; in- ^^^^^ temperate regions of the world, troduced from Europe. We may have, in our material from Utah, Schulz, O. E. 1903. Monographic der Gat- two other species from Europe which are tung Cardamine. Bot. Jahrb. Syst. infrequently recognized: Flowers pinkish (C. 32: 280-623. 1. Leaves all simple, cordate-ovate to orbicular; petals 7-12 mm long; plants of stream and seep margins at middle and higher elevations C. cordifolia Leaves pinnately compound, at least the lower ones; petals 2-7 mm long; plants of spring and seep margins, occasionally elsewhere, at middle to low elevations 2 2(1). Leaflets usually 3-5, terminal leaflet, at least, more than 10 mm wide; upper leaves simple, with broadly ovate blades; petals 3-7 mm long; siliques 1-2 mm wide C. breweri Leaflets usually 6-11 (rarely 3-5), terminal leaflet usually less than 10 mm wide; upper leaves compound; petals 2-3 mm long; siliques 0.7-1 mm wide 3 3(2). Cauline leaves mostly 2-4 cm long, with narrow nondecurrent lateral seg- ments 1-3 mm wide, linear to linear-spatulate or narrowly oblong; rare C. parviflora Cauline leaves mostly 4-8 cm long, with broad decurrent lateral segments 3-8 mm wide, commonly oval to broadly oblong; infrequent C. pensylvanica Cardamine breweri S. Wats. [C. vallicola simple, mostly 1-7 cm long, lateral leaflets Greene] Plants perennial, rhizomatous, erect mostly 10-25 mm long and 0.4-1.2 mm or descending, stems (2) 2.5-5 (6) dm tall, wide, terminal segment 12-35 mm long and glabrous or pubescent with simple hairs 13-30 mm wide, subentire to sharply toothed, near the base; leaves mostly cauline, pin- ovate to orbicular, glabrous or sparsely hir- nately compound with 3-5 (rarely more in sute; pedicels 4-10 mm long or more, ours) leaflets, or basal and upper ones glabrous, ascending; sepals 1.5-2.5 mm long. 306 Great Basin Naturalist Vol. 37, No. 3 whitish, glabrous or sparingly simple-hairy; petals 3-7 mm long, white, rarely pinkish, spatulate-obovate, spreading; siliques 17-30 mm long, 1-1.8 mm wide, erect or ascend- ing, valves glabrous, obscurely 1 (3)-nerved; styles 0.5-2 mm long, tapering to stigma; seeds 1-1.5 mm long, smooth.— Stream sides and seep margins at middle elevations in Wasatch and Utah counties, and imdoubt- edly elsewhere; Alaska and British Colum- bia south to California, Nevada, Utah, and Colorado. Our plant is var. breweri. Cardamine cordifolia A. Gray. [C. cordi- foUa var. pubescens A. Gray; C. infausta Greene; C. uintahensis F. J. Hermann] Plants perennial, rhizomatous, erect or as- cending, stems (1.5) 2-6 dm tall, glabrovis to more or less densely pubescent with simple hairs near the base; leaves mostly cauline, all simple, the blade mostly (1.5) 2-6 (8) cm long and 1.3-5 (7) cm wide, cordate-ovate or broader, usually sinnate-crenate, glabrous or rarely pubescent; pedicels mostly 10-20 mm long, glabrous or hairy, ascending- spreading; sepals 3-5 mm long, greenish, glabrous or sparingly hairy; petals 7-12 mm long, white, obovate-spatulate, spreading; si- liques 20-35 mm long, (1) 1.5-3 mm wide, ascending to erect, valves glabrous, ob- scurely 1 -nerved; styles 0.5-2 mm long or more; seeds 1.5 mm long or more, smooth.— Stream sides and seeps at middle to higher elevations in Beaver (US), Duchesne (US), Garfield (US), Iron, Piute, Salt Lake, San Juan (US), Sanpete (US), Sevier (US), Sum- mit, Utah, Wasatch, Washington, Wayne (US), and Weber counties, and to be ex- pected elsewhere; British Columbia to Wyoming south to California, Nevada, Utah, and New Mexico. Our plants is var. cordifolia. Cardamine parviflora L. Plants annual or biennial from a taproot, stems erect, usu- ally solitary, 1-3 dm tall, glabrous; basal leaves with 3-5 pairs of oblong to cuneate- obovate leaflets, lateral leaflets 0.2-0.4 (0.5) cm long and 0.2-0.3 cm wide, entire or slightly lobed, terminal leaflet broadly cord- ate to orbicular, 0.3-0.8 (1) cm long and 0.5-1 (1.2) cm wide; cauline leaves reduced upwards, mostly 2-4 cm long, with 3-6 pairs of lateral leaflets, segments similar to basal ones only lateral leaflets slightly narrow- er,linear to linear-spatulate or narrowly ob- long, terminal segment linear to cuneate-ob- long, entire or toothed, 0.5-1.2 cm long and 0.1-0.3 cm wide, not decurrent; pedicels 3-7 mm long, spreading-ascending, glabrous; sepals 1.2-1.6 mm long, greenish, glabrous; petals 2-3 mm long, white, oblanceolate, spreading; siliques 12-30 mm long, 0.5-1 mm wide, erect or nearly so, glabrous, style 0.3-0.6 mm long; seeds 1-1.5 mm long, smooth.— Rare and local, Duchesne County; widespread and common in the eastern United States; Europe. Our plant is var. arenicoki (Britton) O. E. Schulz. Cardamine pensylvanica Muhl. ex Willd. [C. flexuosa ssp. pensylvanica (Muhl.) Schulz; C. hirsuta var. pensylvanica (Muhl.) Graff] Plants annual or biennial from a taproot, stems erect, usually solitary, 1.5-3.5 dm tall, glabrous or pubescent; basal leaves with 7-11 pairs of oval to lanceolate or ob- lanceolate leaflets, lateral leaflets 0.3-1.5 cm long and 0.2-1.2 cm wide, entire or lobed, terminal leaflet orbicular to cuneate-oblan- ceolate, 0.4-2 cm long and 0.3-1.5 cm wide; cauline leaves reduced upwards, mostly 4-8 cm long, with 3-5 pairs of lateral leaflets, segments broadly oblong to oval, terminal segment cuneate-obovate, entire or toothed, (0.5) 1-3 cm long and 0.5-2 cm wide, decur- rent; pedicels 3-10 mm long, spreading-as- cending, glabrous; sepals 1.2-1.8 mm long, pinkish, glabrous; petals 2-3 mm long, white, oblanceolate, spreading; siliques 15- 30 mm long, 0.7-1 mm wide, erect, glabrous, style 0.4-0.8 mm long; seeds 1-1.5 mm long, smooth.— Stream sides and other moist areas at middle to low elevations in Duchesne and Summit counties; widespread in North America. This species is only weakly distinct from Cardamine oligosperma Nutt. ex Torr. and Gray, and that species might better be treated as a variant of C pensylvanica. The difficulty with this proposal is that C. pen- sylvanica itself may only be a minor phase of the European C. hirsuta L. We cannot express an opinion on the taxonomy of this species complex based upon our provincial studies, but suspect that our material might best be considered a subspecies of the Eu- Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 307 ropean plant based upon what we have seen in herbaria. Cardaria Desv. White-top Plants pubescent rhizomatose perennials; leaves alternate, sinuate-dentate, auriculate- clasping; flowers in clustered (paniculate) racemes, pedicels spreading-ascending, not subtended by bracts; sepals 4, caducous; petals 4, yellow, the apex rounded; stamens 6, filaments lacking glandular processes; style slender, prominent, stigma capitate; fruit a silicle, usually broader than long, compressed at right angles to septum, in- dehiscent or tardily so, valves reticulately veined; seeds 1 (rarely 2) per locule. As here defined, a genus of 4 species of Europe and Asia. Rollins, R. C. 1940. On two weedy cruci- fers. Rhodora 42: 302-306. 1. Silicles obcordate in outline, glabrous; plants widespread and common C. draba Silicles orbicular in outline, pubescent or glabrous; plants uncommon 2 2(1). Silicles puberulent, 1.5-2.5 mm long and about as wide C. pitbescens Silicles glabrous, 3-4 mm long and about as wide C. chalepense Cardaria chalepensis (L.) Hand.-Mazz. [Lepidium chalepense L., L. draba ssp. chalepense (L.) Thell., C. draba ssp. chale- pensis (L.) O. E. Schulz] Plants decumbent to ascending or erect, 2-6 dm tall, glabrous; leaves elliptic to oblong or lanceolate, 0.8- 10 cm long, 0.3-2 cm wide, sinuate-dentate to entire, the upper sessile and auriculate; pedicels 2-8 mm long in fruit, spreading- ascending, glabrous; sepal 1.2-1.8 mm long, greenish to whitish, glabrous; petals 2-3 mm long, white, spatulate, spreading; silicles (ex- cluding the style) 3-4 mm long, 3-4 mm wide, erect, glabrous, style 0.7-1 mm long; seeds 1 or 2. Moist soil, at 1500 to 1700 m in the Oquirrh Mountains, Salt Lake Co. (Rokich s.n. UT, BRY), and to be expected elsewhere; adventive from Europe. Cardaria draba{L.) Desv. [Lepidium draba L.; Cochlea ria draba (L.) L. Phys- olepidium repens Schrenk ex Fisch. and Meyer; L. repens (Schrenk) Boiss.; C. repens (Schrenk) Jarmolenko] Plants decumbent to ascending or erect, stems (1.2) 1.5-6 dm tall, puberulent to hirtellous with usually de- scending simple hairs; leaves elliptic to ob- long, ovate, or oblanceolate, 0.9-9.8 cm long, 0.6-3.5 cm wide, sinnate-dentate to ir- regularly toothed, lower ones petiolate, up- per sessile and auriculate, puberulent to hir- tellous with usually retrorse simple hairs; pedicels 5-12 mm long in fruit, spreading- ascending, glabrous or puberulent; sepals 1.2-2 mm long, greenish, usually glabrous; petals 2-3.5 (4) mm long, white, broadly spatulate, spreading; silicles (excluding the style) 2-3.8 mm long, 3.5-5.7 mm wide, erect, glabrous, style 0.6-1.2 mm long; seeds 1-2 mm long.— Cultivated and waste places at lower elevations in Beaver, Duchesne, Juab, Salt Lake, Sanpete, Sevier, Utah, Washington, and Weber counties (and prob- ably in most counties of the state); wide- spread in the United States and Canada; ad- ventive from Europe. Cardaria pubescens (C. A. Meyer in Le- deb.) Jarmolenko. [Hymenophysa pubescens C. A. Meyer in Ledeb.] Plant ascending to erect, stems 1.5-4 dni tall, puberulent to hirtellous with usually descending simple hairs; leaves elliptic to oblong or oblanceol- ate, 0.6-6 cm long, 0.3-1.5 cm wide or long- er, irregularly sinuate-dentate, the lower pe- tiolate, upper sessile and auriculate, puberulent to hirtellous with usually simple hairs; pedicels 6-10 mm long in fruit, as- cending, hairy; sepals 1.8-2 mm long, green- ish, hairy; petals 3.5-4 mm long, white, broadly spatulate, spreading; silicles (exclud- ing the style) 1.5-2.5 mm long and about as wide, erect, puberulent, style 0.7-1.2 mm long.— Agricultural lands and disturbed places, Salt Lake Co., and to be expected elsewhere; widespread in the United States and Canada; adventive from Asia. Our plants are var. elongata Rollins. 308 Great Basin Naturalist Vol. 37, No. 3 As here defined, the genus Cardaria in- cludes Hymenophysa. Except for an occa- sional publication, this seems to be strictly an American concept as most workers in Europe and Asia prefer to distinguish be- tween the two. Caulanthus S. Wats. Plants glabrous or pubescent with simple hairs, rarely with some malpighian ones, an- nual to perennial, from taproots; leaves al- ternate or mostly basal, simple, lyrate-pin- natifid, pinnatifid, toothed or subentire, petiolate or sessile and auriculate; flowers racemose, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, white, yellow, or chestnut-brown to purple; stamens 6, fil- aments lacking glandular processes; style obsolete or slender and conspicuous, stigma capitate and sometimes distinctly bilobed; fruit a sessile or subsessile silique, many times longer than broad, terete or more or less compressed, valves 1 (3)-nerved; seeds several to many, uniseriate. A genus of 10-12 species of the western United States, and mainly of California. Payson, E. B. 1922. A monographic study of Thelypodium and its immediate allies. Ann. Missouri Bot. Card. 9:233-324. Rollins, R. C. 1971. Protogyny in the Cruci- ferae and notes on Arabis and Cau- lanthus. Contr. Gray Herb. 201:3-10. 1. Cauline leaves sessile and auriculate at the base; plants of southwestern Utah . C. cooperi Cauhne leaves petiolate or sessile but not auriculate; plants of broad distribu- tion and sometimes of southwestern Utah 2 2(1). Stems usually conspicuously inflated, glabrous or nearly so; plants perennial ... C. crassicaulis Stems not inflated; hispid (at least below); plants annual or biennial 3 3(2). Pedicels very short, 1-2 mm long, soon recurved; siliques up to 4 cm long, descending; known only from Washington County C. lasiophyllus Pedicels 3-7 mm long, spreading-ascending; siliques 4.5-13.5 cm long, ascend- ing to curved-pendulous; known only from Millard and Tooele counties.. C. pilosus Caulanthus cooperi (S. Wats.) Payson. [Thelypodium cooperi S. Wats.; Guillenia cooperi (S. Wats.) Greene] Plants annual, erect or sprawling, stems not inflated, 1-7.5 dm tall, glabrous and often glaucous or sparsely pubescent with simple or malpig- hian hairs; leaves mainly cauline, lower ones 1-7 cm long, 0.4-2 cm wide or more, obscu- rely sinuate-dentate, glabrous, cauline ones reduced upwardly, mostly 1-7.5 cm long and 0.2-1.2 cm wide, mostly entire with at least the uppermost auriculate; pedicels 1-4 mm long, soon recurved, glabrous; sepals 5- 6.5 (7) mm long, green or reddish, glabrous; petals 6-9 mm long, yellowish, suffused with purple, narrowly spatulate, ascending; an- thers 1-1.3 mm long; siliques 20-45 mm long, 2-3 mm wide, sessile, descending, compressed, glabrous; styles 1-2.5 mm long, stigma not expanded, shortly bilobed.— iMr- rea and Joshua tree communities in Wash- ington County, Utah; Arizona, Nevada, and California. Caulanthus crassicaulis (Torr.) S. Wats. Plants perennial, erect, stems usually strong- ly inflated, (2) 3.2-9.7 (10.8) dm tall, glabrous and glaucous; leaves mainly basal, lower ones 3-12 (17) cm long, 0.3-3 cm wide, lyrate-pinnatifid to entire, glabrous, cauline ones much reduced upwardly, linear to narrowly oblanceolate, petiolate and not auriculate; pedicels 1-4 mm long, stout, as- cending, glabrous or more commonly hir- sute at least apically; sepals (7) 9-13 mm long, brown to brown-purple, narrowly spatulate, spreading-ascending; anthers 3.9- 6.3 mm long; siliques 70-140 mm long, 1.5- 2 mm wide, sessile, ascending to erect, glabrous; style obsolete, stigma more or less expanded, lobes up to 0.8 mm long.— Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 309 Pinyon-juniper, sagebrush, shadscale, and ponderosa pine woodlands from moderate to low elevations in the southern two-thirds of Utah; Oregon and Idaho to Colorado south to California and Arizona. As here defined, we are treating Cau- lanthus crassicaiilis in a broad sense and not recognizing C. glaher nor C. major. These three sympatric entities, segregated on fea- tures which tend to show some apparent in- tergradation in Utah, seem to be very dis- tinct elsewhere, particularly in Nevada and California. The var. crassicaiilis is reported to have a haploid number of n= 12, where- as as one of us has shown (Reveal & Styer, Southw. Naturalist 18:397-402. 1974) the var. glaher is n= 14 based on a collection from southwestern Nevada. The two vari- ants of the species must be carefully in- vestigated in the field to see if the plants found outside Utah really belong to the taxa we have in the state. 1. Sepals more or less hirsute; plants of broad distribution C crassicaulis var. crassicaulis Sepals glabrous or with a few hairs; plants of various distributions 2 2(1). Stigmas deeply divided; plants common C. crassicaulis var. glaher Stigmas shallowly lobed; plants evidently rare C. crassicaulis var. major Var. crassicaulis. [Streptanthus crassicaulis Torr.; C. senilis Heller] Widespread and lo- cally common in Box Elder, Carbon (US), Daggett, Duchesne, Emery, Juab, Kane, Sanpete, Sevier, and Uintah (US) counties and likely elsewhere— the type is from the east side of the Great Salt Lake; Idaho, Ne- vada, Colorado, Arizona, and California. Var. glaher M. E. Jones. [C. glaher (M. E. Jones) Rydb.] Widespread and locally com- mon in Beaver, Garfield, Kane, Millard, Piute, San Juan, Sevier, and Washington counties— the type is from Sink Valley in Kane County; southern Nevada. Var. major M. E. Jones [C. 7najor (M. E. Jones) Payson; C. procerus of authors, not S. Wats.] Rare and local, perhaps not sepa- rable from var. glaher in Utah where known only from Garfield County— the type is from Bromide Pass in Henry Mountains; Nevada and California where perfectly dis- tinct from both var. glaher and var. crassi- caulis. Caulanthus hsiophyllus (Hook. & Arn.) Payson. [Turritis lasiophylla Hook. & Arn.; Thehjpodium lasiophijllum (Hook. & Arn.) Greene; Sisymhrium lasiophyllum (Hook. & Arn.) K. Brandegee; Guillenia lasiophylla (Hook. & Arn.) Greene] Plants annual, stems not inflated, 1-8 (12) dm tall, more or less hirsute with simple or rarely forked hairs; leaves mainly cauline, these 0.7-15 cm long, 0.1-5.5 cm wide, irregularly pinnatifid. petiolate and not auriculate; pedicels 1-2 mm long, deflexed in fruit, glabrous or spar- ingly hirsute; sepals 2-3 mm long, often purplish, glabrous; petals 3-5 mm long, yel- lowish, oblong-spatulate, not constricted at juncture of blade and claw, ascending- spreading; anthers 0.6-1 (1.5) mm long; si- liques 25-45 (60) mm long, 0.8-1.1 mm wide, sessile, terete, reflexed-descending, glabrous; styles 0.8-1.3 mm long, stigma small, obscurely lobed.— Sandy or gravelly soils in the Larrea community in Washing- ton County; California and Nevada south into Arizona and Mexico. Our material has been designated as var. utahensis (Rydb.) Payson [Thelypodium utahense Rydb.; T. lasiophyllum var. utahense (Rydb.) Jeps.], basically a Mojave Desert phase which oc- curs in northwestern Arizona, southern Ne- vada, and southeastern California. Caulanthus pilosus S. Wats. [Streptanthus pilosus (S. Wats.) Jeps.] Plants biennial or infrequently annual, stems not inflated, mostly 4-10 dm tall or more, hirsute with simple hairs at least below; leaves mostly basal, lower ones 3-15 cm long, 0.5-3.5 cm wide, irregularly pinnatifid, hirsute, cauline ones only slightly reduced and shorter up- wardly, petiolate and not auriculate; pedi- cels 4-9 mm long, spreading-ascending, glabrous or nearly so; sepals 5-7 (9) mm long, often purplish, glabrous or hairy; pet- als 7.5-9 (10) mm long, white, suffused with 310 Great Basin Naturalist Vol. 37, No. 3 purple or pink, spatiilate-lanceolate, con- stricted at juncture of blade and claw, as- cending-spreading; anthers 2-4 mm long; si- liques 70-115 mm long, 0.8-1 (1.5) mm wide; style short, stigma bilobed.— Sandy or gravelly soils in the shadscale-winter fat communities of Millard and Tooele coun- ties; Oregon and Idaho south to C^alifornia and Nevada. Chlorocramhe Rydb. Plants glabrous perennials from a stout caudex; leaves alternate and mainly cauline, simple, more or less hastate, entire or si- nuately lobed, petiolate, not both sessile and auriculate; flowers racemose, pedicels not subtended by bracts; .sepals 4, deciduous; petals 4, white; stamens 6, filaments lacking glandular processes; style obsolete or up to 0.5 mm long, stigma small, entire; fruit a stipitate silique, many times longer than broad, subterete, valves 1 (3-5)-nerved; seeds imiseriate. A monotypic genus. Chlorocramhe hastatus (S. Wats.) Rydb. [Caulantluis hastatus S. Wats.] Plants erect, stems 6-18 dm tall, usually simple, glabrous and glaucous; leaves with slender petioles 1- 16 cm long, blades hastate to ovate or lan- ceolate, 3-13.5 cm long, 1-8.5 cm wide, more or less hastate, entire or sinuate-lobed; pedicels spreading to reflexed, 5-10 mm long, glabrous; sepals usuallv siupassing pet- als, their tips coiled, yellowish-green, glabrous; petals 4.5-8 mm long, white, most- ly 4-6 mm long, blade constricted at junc- ture with claw, ascending-spreading; siliques 40-105 mm long, 1.8-2.5 mm wide, spread- ing to curved descending, glabrous, stipe 2- 7 mm long; style up to 0.5 mm long, stigma not lobed.— Thickets, woodlands, and less commonly in openings from 1900 to 2800 m elevation in Davis, Salt Lake, Tooele, Utah, Wasatch, and Weber counties; known only from northern Utah and the Wallowa Mountains of northeastern Oregon. Chorispora R. Br. ex DC. Nom. Cons. Plants stipitate-glandular and infrcfjuently also hirsute annuals, from taproots; leaves alternate and basal, simple, sinuate-dentate to pinnatifid or entire, cauline ones petiol- ate to sessile but not auriculate; flowers racemose, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, pink to laven- der; stamens 6, filaments lacking glandular processes; style apical on a slender sterile beak, stigma minute, bilobed; fruit a silique, many times longer than broad, terete, in- dehiscent, breaking at maturity into 1 -seed- ed segments, valves 1 (3 or more)-nerved; seeds uniseriate. A genus of about 9 species, mainly of central Asia. Chorispora tenella (Pall.) DC. [Raphanus tcncllus Pall.; Cliorispennum teneUum (Pall.) R. Br. ex Ait.] Plants deciuiibent-ascending to erect, stems 0.2-4.5 dm tall, simple or branched from base, stipitate-glandular and often simple hirsute at least at base; leaves mainly cauline, 0.5-8.5 cm long, 0.1-2.8 cm wide, sinuate-dentate, pinnatifid, or entire, petiolate or sessile but not auriculate; pedi- cels spreading-ascending, 2-6 mm long, stipitate-glandular and often sparingly hir- sute; sepals 4.4-6.7 mm long, reddish or purplish, stipitate-glandular; petals 9-12.5 mm long, pink to lavender, apex rounded, spreading; silicjues 30-45 mm long, curved- ascending, stipitate-glandvilar, beak 8-22 mm long; style obsolete or very short, stigma minute.— Roadsides, foothills, and other dis- turbed sites at lower elevations in Box El- der, Emery, Grand, Kane, Salt Lake, San- pete, Utah, Washington, Wayne, and Weber counties, and probably throughout the state; Washington and Idaho south to California, Arizona and Colorado; adventive from Asia. Conrincua Adans. Plants glabrous and glaucous annuals or biennials, from taproots; leaves alternate and basal (and still alternate), simple, entire, tapering to ba.se or cauline sessile and au- riculate-cla.sping; flowers racemose, pedicels ascending to curved-erect, not subtended by bracts; sepals 4, deciduous; petals 4, yellow or cream; stamens 6, filaments lacking glandular appendages; styles stout, stigma lobed; fruit a sessile, slender silicjue many times longer than broad, quadrangular, valves 1— 3-nerved; seeds numerous, unise- riate. Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 311 About 6 species of Eurasia. Descurainia Webb and Berthel. Conringia orientalis (L.) Diimort. [Bras- Nom. Cons. sica orientalis L.] Hare's-ear Mustard. Plants Plants stellate-pubescent, stipitate-glandu- annual or winter annual, stems 1.9-5 (7) dm lar, or glabrate annuals or biennials from tall, solitary or 2 or 3 from base, glabrous; slender to stout taproots; leaves basal and basal leaves 3-6 (9) cm long, 1.2-3 cm wide, cauline, alternate, 1-3 times pinnately com- entire, glabrous; cauline leaves several, 1.7- pound or pinnatifid, not auriculate basally; 12 cm long, 0.8-5.7 cm wide, ovate to ob- flowers racemose, pedicels not subtended by long to elliptic, shorter to longer than inter- bracts; sepals 4, deciduous; petals 4, yellow nodes, glabrous, entire; pedicels 5-14 mm to cream; stamens 6, filaments lacking long, ascending to curved-erect, glabrous; glandular processes; style short or obsolete, sepals 4.3-6 (8) mm long, glabrous, often stigma capitate; fruit a silique more than (3) reddish tinged, acute; petals 6.2-10 (12) mm 5 times longer than broad, linear to oblong long, yellow to cream, spatulate, spreading- or clavate, terete or nearly so, valves 1- ascending; siliques 70-100 (130) mm long, nerved, glabrous; seeds several to many, 1.5-2 mm thick, erect, valves glabrous; uniseriate or biseriate. styles up to 1 mm long, stigma small; seed Perhaps 30 species mainly of the Ameri- numerous.— Slopes, roadsides, and other dis- cas, a few in Europe. turbed sites at middle and lower elevations Detling, L. E. 1939. Descurainia in North in Utah and Washington counties (and to be America. Amer. Midi. Naturalist 22:481- expected elsewhere); widely distributed in 520. North America; adventive from Europe. Schulz, O. E. 1924. Descurainia. Pflanzenr. IV. 105 (Heft 86): 481-520. 1. Upper leaves bi- or tripinnate; siliques narrowly linear, mostly about 20 (10- 30) mm long; seeds usually more than 20, uniseriate; replum 2-3-nerved; tall to low plants of low elevations D. sophia Upper leaves once-pinnate; siliques clavate, elliptic, or, if linear, less than 20- seeded and less than 15 mm long; replum nerveless or 1-nerved 2 2(1). Siliques clavate or linear to elliptic, rounded to pointed above; seeds often in 2 rows or at least partially so; replum usually nerveless; plants of middle to lower elevations D. pinnata Siliques linear or elliptic, usually pointed above; seeds in one row; replum 1- nerved; plants of middle and higher elevations 3 3(2). Siliques 7-14 mm long, linear or less commonly ellipsoid; pedicels appressed- erect or ascending; seeds mostly 4-10 per locule D. richardsonii Siliques 3.3-7.2 mm long, ellipsoid; pedicels ascending to spreading; seeds mostly 1-3 per locule D. californica Descurainia californica (A. Gray) O. E. ing to spreading-ascending, glabrous; sepals Schulz. [Smelowskia californica A. Gray] spreading, 0.8-1.3 mm long, yellow or Plants annual or winter annual to biennial, greenish, glabrous; petals 1.1-1.6 mm long, stems 4-13 dm tall or more, simple or more yellow; siliques 3.3-7.2 mm long, 0.9-1.3 commonly profusely branching from most of mm wide, erect or ascending, not ap- upper leaf and bract axils, minutely dendrit- pressed; styles 0.3-0.7 mm long; seeds unise- ic-stellate pubescent to almost or quite riate, 1-3 per locule.— Woodlands, espe- glabrous below, glabrous above; leaves basal cially in aspen but also in ponderosa pine, and cauline, 2-7 cm long, lower once-pin- spruce-fir, and less commonly in mountain nately compound with 2-4 pairs of entire to brush communities from 1750 to 3100 m incised pinnae, upper reduced, once-pinnate elevation in Beaver, Carbon, Garfield, Mil- or pinnatifid; pedicels 3-7 mm long, spread- lard, Piute, Salt Lake, Utah, Washington, 312 Great Basin Naturalist Vol. 37, No. 3 Wayne, and Weber counties; Wyoming, Colorado, and New Mexico westward to California. This taxon is a mirror-image congener of Descurainia richardsonii (q.v.), especially of those taxa of that species with ascending pedicels. Descurainia pinnata (Walter) Britton [Erysiinwn pinnatum Walter; Sisymbrium pinnatum (Walter) Greene; Sophia pinnata (Walter) Howell] Pinnate Tansy- Mustard. Plants annual or winter annual, stems 1-10 dm tall, stellate-pubescent and sometimes also stipitate-glandular at least below, simple or highly branched above; leaves basal and cauline, 2-10 cm long, once to twice pinnatifid, segments linear to oblong, often toothed, upper reduced and usually once pinnatifid; pedicels 3-24 mm long, spreading, stellate-pubescent to glabrous; sepals 1-2.2 mm long, yellowish to greenish or violet, stellate-pubescent to glabrous; petals 1.5-3 mm long, cream to yellow; si- liques 3-15 (rarely 20) mm long, 1-2 mm wide, clavate to oblong or linear, very rarely ellipsoid; styles up to 0.3 mm long; seeds 1-20 per locule, biseriate in part.— Widespread and common in arid to sub- mesic regions of North America; composed of numerous freely intergrading variants, several of which occur in Utah. 1. Pedicels (17) 18-24 mm long; siliques mostly 12-18 mm long; petals yellow, over 2 mm long; terminal leaflet of uppermost leaves linear, entire, and more than 2 cm long; plants of Grand, San Juan, and Uintah counties D. pinnata var. paysonii Pedicels usually less than 15 mm long; siliques often less than 12 mm long (up to 20 mm in var. filipes); terminal leaflet various but not or seldom as above; petals yellow to cream, shorter or longer than 2 mm in length; dis- tribution various 2 2(1). Stems moderately to densely stipitate-glandular 3 Stems stellate-pubescent to glabrous, not at all stipitate-glandular 4 3(2). Flowers with calyx usually rose-colored; corolla 1.5-2 (2.2) mm long; siliques usually 10 mm long or less D. pinnata var. osmiarum Flowers with calyx yellowish; corolla 2-3 mm long; siliques often more than 10 mm long D. pinnata var. filipes 4(2). Siliques usually shorter than pedicels, mostly 10-21 mm long, not or only somewhat clavate in outline D. pinnata var. osmiarum Siliques usually subequal to or longer (rarely shorter) than pedicels, mostly 3- 12 mm long, more or less clavate or elliptic in outline 5 5(4). Siliques 3-8 mm long, borne on pedicels 4-12 mm long; plants of warm deserts in Washington County D. pinnata var. glabra Siliques 8-10 mm long, or, if only 4-8 mm long, not of Washington County; pedicels various 6 6(5). Pedicels 4-6 (8) mm long; flowers 1-1.5 mm long D. pinnata var. nelsonii Pedicels (6) 8-12 mm long; flowers 2-3 mm long D. pinnata var. intermedia Var. filipes (A. Gray) M. E. Peck. [Sisyin- brium incisum var. filipes A. Gray; Sisym- brium longipedicellata Fourn. Sophia filipes (A. Gray) Heller; Sisymbrium gracilis Rydb.; D. rydbergii var. eglandulosa O. E. Schulz; D. longipedicellata (Fourn.) O. E. Schulz; D. longipedicellata var. glandulosa O. E. Schulz; Sisymbrium glandifera Osterh.; Si- symbrium longipedicellata var. glandulosa (b. E. Schulz) St. John; D. pinnata ssp. fi- lipes (A. Gray) Detling] Common and wide- spread from 975 to 2300 m elevation in desert shrub, mountain brush, pinyon-juni- per, and ponderosa pine communities in Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 313 Beaver, Emery, Grand, Juab, Kane, Millard, Salt Lake, San Juan, Sevier, Tooele, Utah, Washington, and Wayne counties; British Columbia and Alberta south to California, Arizona, and Colorado. Var. glabra (Woot. & Standi.) Shinners. [Sophia glabra Woot. & Standi.; D. pinnata ssp. glabra (Woot. & Standi.) Detling] Local common from 750 to 1200 m elevation in Washington County; Arizona and New Mexico west to California and northern Mexico. Var. intermedia (Rydb.) C. L. Hitchc. [Sophia intermedia Rydb.; D. intermedia (Rydb.) F. P. Daniels; Sisymbrium inter- medium (Rydb.) Garrett; D. pinnata ssp. in- termedia (Rydb.) Detling] Locally common in pinyon-juniper, sagebrush, and semidesert shrublands from 1200 to 1850 m elevation in Garfield, Kane, Millard, Uintah, and Wayne counties; British Columbia and Al- berta south to California, Nevada, Utah, and Colorado. Var. nelsonii (Rydb.) M. E. Peck. [Sophia nelsonii Rydb.; D. brachycarpa var. nelsonii (Rydb.) O. E. Schulz] Widespread but ap- parently imcommon from 1200 to 3250 m elevation in Duchesne, Garfield, San Juan, Sevier, Uintah, and Wasatch counties; Washington east to Montana and south to Nevada, Utah, and Colorado. Var. osmiarum (Cockerell) Shinners. [Sophia andrenarum var. osmiarum Cock- erell; S. halictonim Cockerell; S. andrena- rum Cockerell; Sisymbrium halictorum (Cockerell) K. Schum.; D. halictorum (Cock- erell) O. E. Schulz; D. halictorum var. an- drenarum (Cockerell) O. E. Schulz; D. ha- lictorum var. osmiarum (Cockerell) O. E. Schulz; D. andrenarum (Cockerell) Cory; D. pinnata ssp. halictorum (Cockerell) Detling] Our most common variant, in sagebrush, pinyon-juniper, salt-desert shrub, grasslands, and warm desert shrublands from 850 to 2450 m elevations in Box Elder, Davis, Duchesne, Emery, Garfield, Iron, Juab, Kane, Millard, Piute, San Juan, Sevier, Uin- tah, Utah, Washington, and Wayne coun- ties; Oregon to Wyoming south to Mexico, Oklahoma, and Arkansas. Var. paysonii (Detling) Welsh & Reveal, Stat, nov., based on Descurainia pinnata ssp. paysonii Detling, Amer. Midi. Naturalist 22:515. 1939. Uncommon in pinyon-juniper and sagebrush zones at lower elevations in Grand, San Juan, and Uintah counties; Wyoming and Colorado south into Arizona. Descurainia richardsonii (Sweet) O. E. Schulz. [Sisymbrium canescens Richards., not D. canescens Nutt.; S. richardsonii Sweet; S. canescens var. mapr Hook.; Soph- ia richardsoniana Rydb.] Richardson Tansy- mustard. Plants annual or winter annual to biennial, stems (1.5) 3-12 dm tall or more, simple or more commonly profusely branch- ing from upper leaf and bract axils, minute- ly dendritic-stellate pubescent and some- times also minutely stipitate-glandular to almost or quite glabrous below, glabrous or glandular above; leaves basal and cauline, 1,5-8 cm long, the lower once- to twice-pin- natifid with 2-4 pairs of toothed or lobed or subentire pinnae, the upper reduced and only once-pinnatifid; pedicels (2.5) 3-9 mm long, spreading to erect, pubescent or glabrous; sepals spreading, 0.9-1.7 mm long, yellow or greenish, glabrous or hairy; petals 1.3-2.8 mm long, yellow; siliques 7-14 mm long, 0.8-1.2 mm wide, ascending or erect, often appressed; styles 0.2-0.6 mm long; seeds imiseriate, (1-3) 4-10 per locule.— Spruce-fir, aspen, mountain meadows, or sagebrush communities from 2150 to 2900 m elevation in central and eastern Utah; British Columbia and Alberta south to Cali- fornia, Arizona, and Mexico. Three weakly defined variants can be at least arbitrarily distinguished as in the following key: 1. Pedicels and siliques erect, more or less appressed to raceme axis D. richardsonii var. brevipes Pedicels ascending, siliques erect or ascending, neither one appressed to 2 raceme axis 2(1). Plants not glandular D. richardsonii var. sonnet Plants stipitate-glandular D. richardsonii var. viscosa 314 Great Basin Naturalist Vol. 37, No. 3 Var. brevipes (Nutt. ex Torr. & Gray) Welsh & Reveal, comb, nov., based on Si- si/mbriian canescens var. brevipes Nutt. ex forr. & Gray, Fl. N. Amer. 1:92. 1838. [Sophia procera Greene; Sisymbrium pro- ceritm (Greene) K. Schum; Sophia brevipes (Nutt.) Rydb.; D. richardsonii var. macro- spenna O. E. Schulz, type from Alta, Salt Lake Coimty; D. richardsonii ssp. procera (Greene) Detling; D. richardsonii var. pro- cera (Greene) Breitung] Locally common from 2450 to 2900 m elevation in Duch- esne, Garfield, Iron, Piute, Salt Lake, Sum- mit, and Wasatch counties; Idaho and Mon- tana south to New Mexico. Var. sonnei (B. L. Robinson) G. L. Hitchc. [Sisymbrium incisum Engelm. in Gray; D. incisa (Engelm.) Britton; S. in- cisum var. sonnei B. L. Robinson; Sophia sonnei (B. L. Robinson) Greene; Sophia in- cisa (Engelm.) Greene; Sophia leptophylla Rydb.; Sophia serrata Greene; Sophia pur- purascens Rydb; Sisymbrium leptophyllum (Rydb.) Nels. & Macbride; D. serrata (Greene) O. E. Schulz; D. incisa var. lepto- phylla (Rydb.) O. E. Schulz; D. richardsonii ssp. incisa (Engelm.) Detling] Widespread and locally common from 1800 to 2750 m elevation in Duchesne, Garfield, Grand, Salt Lake, Utah, and Wasatch counties; Idaho and Montana southward to Mexico. This phase of Descurainia richardsonii closely simulates D. californica (q.v.). Var. viscosa (Rydb.) M. E. Peck. [Sophia viscosa Rydb.; Sisymbrium viscosum (Rydb.) Blankinship; D. rydbergii O. E. Schulz; D. richardsonii ssp. viscosa (Rydb.) Detling] Local and common from 2450 to 2900 m elevation in Duchesne and Wasatch coun- ties; British Columbia and Alberta south to California, Arizona, and New Mexico. The var. richardsonii is known to occur just north and east of Utah in southwestern Wyoming and northwestern Colorado. It differs from var. brevipes in being canescent as opposed to the moderately pubescent to nearly glabrous condition of var. brevipes. Descurainia sophia (L.) Webb. [Sisym- brium Sophia L.; Sophia sophia (L.) Britton; Sophia parviflora Standi.] Plants annual or infrequently winter annual, stems 1.7-8.5 (10) dm tall or more, simple or more com- monly branched above, softly dendritic- or mixed simple- and dendritic-hairy at least below; leaves basal and cauline, 1-12 cm long, the lower 2-3 times pinnately com- pound to pinnatifid, with 2-6 pairs of pin- natifid pinnae, the upper ones smaller and usually twice pinnately compound or pin- natifid; pedicels 4-17 mm long, ascending, puberulent or glabrous; sepals erect, 2-3.1 mm long, yellowish, glabrous or hairy; pet- als 2.2-3 mm long, cream; siliques (10) 12- 27 (30) mm long, 0.8-1.2 mm wide, ascend- ing-erect; styles 0.1-0.3 mm long; seeds uniseriate, mostly 10-25 mm long.— Road- sides, corrals, agricultural lands, foothills, and other disturbed sites from 750 to 2450 m elevation in Beaver, Emery, Garfield, Grand, Juab, Millard, San Juan, Summit, Uintah, Washington, and Weber counties, and probably cosmopolitan in Utah; wide- spread in North America; adventive from Europe. DiPLOTAXIS DC. Plants glabrous or simple-hirsute annuals or biennials, from taproots; leaves alternate, mostly basal, pinnatifid to irregularly toothed, reduced upwards and petiolate to merely sessile but not auriculate; pedicels ascending or ascending-spreading, not subtended by bracts; sepals 4, deciduous; petals 4, yellow or sometimes fading rose; stamens 6, fila- ments lacking glandular processes; styles stout, well developed, stigma capitate; fruit a silique, many times longer than broad, lin- ear, somewhat flattened parallel to parti- tion, valves 1-3-nerved, beak not nerved; seeds numerous, biseriate. A genus of perhaps 20 species of Eurasia. Diplotaxis mutatis (L.) DC. [Sisymbrium murale L.] Plants erect or ascending, pub- escent with coarse, descending hairs at least below, stems 0.7-5 dm tall, usually branched; basal leaves lyrate-pinnatifid to irregularly lobed, mostly 2.5-9.5 cm long, 0.5-3.5 cm wide; cauline leaves usually much reduced upwards, all petiolate, not auriculate; pedicels 6-23 mm long or more, slender, ascending to spreading-ascending, often hirsute; sepals 3-4.5 mm long, purplish tinged, glabrous or hirsute; petals 4.5-7.5 mm long, yellow or sometimes fading rose; Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 315 siliques 17-33 mm long, 1.5-2.8 mm wide; style 1.5-2.5 mm long, valves glabrous, lightly 1-3-nerved.— Fields and disturbed sites in Cache County, and to be expected elsewhere; scattered in North America; ad- ventive from Europe. DiTHYREA Harv. Plants stellate or dendritic hairy, annuals or winter annuals, from taproots; leaves al- ternate, simple, subentire to toothed, lobed or pinnatifid, petiolate to sessile but not au- riculate; flowers racemose, pedicels spread- ing, not subtended by bracts; sepals 4, de- ciduous; petals 4, white; stamens 6, filaments lacking glandular processes; style broad and stout, stigma enlarged-capitate; fruit a silicle, more than twice longer than broad, spectaclelike, compressed at right an- gles to the replum, valves reticulately veined; seeds 1 per locule. A genus of 2 species of western North America. Dithyrea wislizenii Engelm. in Wisliz. Spectacle-pod. Plants erect or ascending, pubescent with soft stellate or dendritic hairs, stems 0.7-5 dm tall, simple or branched; basal leaves often withered by anthesis; cauline leaves 1.2-9.5 cm long, 0.2- 2.5 cm wide, pinnatifid, sinuately dentate, irregularly lobed, or entire, moderately to densely hairy like stems, reduced upwards; pedicels 5-21 mm long in fruit, spreading, dendritic hairy; sepals 3-5.5 mm long, greenish, yellowish, or purplish, usually den- dritic hairy; petals 4.8-8 mm long, white, claws occasionally lavender, blades orbicular to spatulate, 2.5-5.5 mm wide; silicles 4-6.5 mm long (from apex of the short stipe), 9- 14 mm wide, valves pubescent; seeds 1 per locule.— Sandy sites in warm desert shrub. Draba L. Plants with stellate, dendritic, forked, or simple hairs, or glabrate, annual, biennial, or perennial, from taproots and often with grassland, and pinyon-juniper communities from 750 to 1700 m elevation in Emery, Garfield, Grand, Kane, San Juan, and Wash- ington counties; Colorado, Oklahoma, and Texas westward to Utah and Arizona. There is much variation in leaf form, but this does not seem to be correlated with other features or with ecological variations, well-developed caudices; leaves all basal or some cauline, alternate, simple, entire or toothed, tapering to the base or rounded, not auriculate; flowers racemose, pedicels usually not subtended by bracts; sepals 4, deciduous; petals 4, white, yellow or cream, apex rounded to bifid; stamens 6, filaments lacking glandular processes; style obsolete to prominent and slender, stigma obscurely bilobed; fruit a silicle or infrequently a short silique, mostly 1-10 times longer than broad, ovale to ovate, lanceolate or linear, compressed parallel to septum, plane or twisted, straight or curved, valves obscurely 1 -nerved or nerveless; seeds biseriate, usual- ly numerous. A genus of 250 to 300 species mainly of the northern temperate region of the world. Hartman, R. L., et al. 1975. Biosystematics of Draba cuneifolia and D. platycarpa (Cniciferae) with emphasis on volatile and flavonoid constitutents. Brittonia 27:317-327. Hitchcock, C. L. 1941. A revision of the drabas of western North America. Univ. Wash. Publ. Biol. 11:1-132. Payson, E. B. 1917. The perennial scapose drabas of North America. Amer. J. Bot. 4:253-267. 1. Plants scapose, leaves all basal, mostly depressed-cespitose perennials (ex- cept in D. verna which has deeply bilobed petals) Key 1. Plants with one to many cauline leaves in addition to basal ones, annual (and the petals not bilobed), biennial, or perennial plants Key II. Key I. Plants scapose 1. Plants annual, flowering in springtime; style scarcely if at all evident (up to 0.1 mm long) D. verna 316 Great Basin Naturalist Vol. 37, No. 3 Plants perennial, flowering in springtime and in summer; styles 0.15-2.5 mm long 2 2(1). Petals white 3 Petals yellow 6 3(2). Plants pubescent throughout with doubly pectinate hairs; known only from Daggett County D. oligospemia Plants variously hairy, but either stellate or with simple or forked hairs only, seldom or nearly pubescent throughout; known from Uinta and Wasatch mountains and plateaus of southern Utah 4 4(3). Leaves with simple hairs only; petals 2-3 mm long; styles up to 0.2 mm long D. fladnizensis Leaves variously pubescent; petals mostly 3-5 mm long; styles 0.2-0.5 mm long 5 5(4). Leaves cinereous with appressed hairs, sometimes stellate; plants of high elevations in Wasatch and Uinta mountains D. lonchocarpa Leaves glabrous above and below, ciliate with simple or forked hairs; plants of plateaus of southern Utah D. subalpina 6(2). Leaves glabrous or with unforked hairs only, often merely ciliate 7 Leaves pubescent, at least some of the hairs forked, stellate, or doubly pec- tinate 9 7(6). Styles 1-2.5 mm long; petals 5-6 mm long; plants of Box Elder and Weber counties D. maguirei Styles usually less than 1 mm long; petals less than 5 mm long; distribution various 8 8(7). Styles not over 0.2 mm long; plants biennial or short-lived perennial, leaves not densely imbricated; petals 1.5-3 mm long D. crassifolia Styles 0.2-1 mm long; plants perennial, densely cespitose, with closely im- bricated leaves; petals often more than 3 mm long D. densifolia 9(6). Lower side of leaves (at least) with appressed, pectinately branched hairs; plants of Wasatch and Uinta mountains D. oligospemia Lower side of leaves glabrous or pubescent, but, if so, with merely stellate or forked hairs, not pectinate 10 10(9). Leaves almost glabrous, ciliate with forked or dendritic hairs; plants of Box Elder and Weber counties D. maguirei Leaves more or less pubescent on one or both surfaces 11 11(10). Stems glabrous, at least above 12 Stems pubescent throughout 13 12(11). Basal leaves 1-3.5 cm long, 0.3-1 cm wide; silicles 7-14 mm long, 2-4 mm wide; plants of Washington County D. asprella Basal leaves 0.6-1.2 cm long, 0.9-1.4 cm wide; siliques 4-10 mm long, 1.5-3 mm wide; plants of Cache County D. maguirei 13(11). Leaves densely cinereous pubescent, individual hairs almost in- distinguishable; known from Uinta Mountains D. ventosa Leaves not densely cinerous pubescent, individual hairs apparent; plants of Garfield and Piute counties D. sobolifera Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 317 Key II. Plants with 1 or more cauline leaves 1. Plants annual; styles obsolete or rarely up to 0.2 mm long 2 Plants biennial or perennial; styles mostly 0.2-1 mm long or more 7 2(1). Upper portion of stem, including pedicels, pubescent 3 Upper portion of stem glabrous 4 3(2). Flowers white; plants widely distributed at low elevations D. cuneifolia Flowers yellow; plants of montane regions in northern Utah D. rectifructa 4(2). Upper leaf surfaces usually glabrous; cauline leaves 1 or 2, rarely lacking .... D. crassifolia Upper leaf surfaces usually pubescent; cauline leaves 1-5 or more 5 5(4). Petals white; leaves entire or nearly so; silicles less than 2 mm wide; plants of low elevations D. reptans Petals yellow, sometimes fading whitish; silicles at least 2 mm wide or the plants of montane places 6 6(5). Pedicels usually at least 1.5 times longer than the silicles; plants usually of low elevations D. nemorosa Pedicels rarely up to 1.5 times longer than the silicles; plants usually of montane places D. stenoloba 7(1). Petals white; plants of high elevations 8 Petals yellow or cream, sometimes fading whitish; plants of low to moder- ate or high elevations 10 8(7). Styles obsolete or nearly so, less than 0.2 mm long D. fladnizensis Styles 0.2-0.8 mm long 9 9(8). Silicles glabrous or merely ciliate; cauline leaves 1 or 2 D. lonchocarpa Silicles pubescent on valves; cauline leaves 1-10 D. lanceolata 10(7). Cauline leaves solitary; plants of low elevations, restricted to Washington County D. asprella Cauline leaves usually 2-20, or if solitary then not of Washington County; plants not or rarely of Washington County 11 11(10). Styles obsolete or nearly so, up to 0.2 mm long D. stenoloba Styles 0.2-2.5 mm long 12 12(11). Petals 2.8-3.8 mm long; pubescence of leaves stiff, stalked, 2— 5-rayed hairs; silicles pubescent with simple or stalked and forked hairs D. brachystylis Petals (3.8) 4-8 mm long; pubescence of leaves various; silicles glabrous or pubescent with stellate to simple hairs 13 13(12). Leaf surfaces glabrous, ciliate or margins smooth; silicles glabrous D. crassa Leaf surfaces with stalked 2— 4-rayed, forked, or simple hairs, margins ci- liate or not; silicles glabrous or hairy 14 14(13). Leaves solitary, subtending lowermost flower or branch of inflorescence; plants of Box Elder County D. incerta Leaves 2-20; plants of various distribution 15 15(14). Leaves bright green to somewhat grayish; cauline leaves often denticulate; 318 Great Basin Naturalist Vol. 37, No. 3 silicles plane or slightly contorted; plants of Grand and San Juan counties D. spectabilis Leaves grayish-green; cauline leaves entire; silicles usually contorted; plants of broad distribution D. aurea Draba asprella Greene. Plants perennial, cespitose, stems 0.5-1.4 (2) dni tall, arising from a branching caudex, hirsute with mixed simple and forked to dendritic hairs; leaves all basal, rarely with one cauline, 1-3.5 cm long, 0.3-1 cm wide, oblanceolate to spatu- late, entire or obscurely denticulate, green, surfaces more or less pubescent with usually stalked and 4-rayed hairs; racemes simple, 10— 30-flowered, elongating in fruit; pedi- cels 3-15 mm long, ascending, glabrous; sep- als 1.8-2.5 mm long, greenish, stellate-hairy; petals 3.4-5 (6) mm long, yellow to yellow- orange, obovate-spatulate, rounded; silicles 7-14 mm long, 1.5-4 mm wide, lance-ellip- tic, glabrous or ciliate; styles 0.8-1.1 mm long; seeds 12-20.— Talus slopes and low hills from 1050 to 1700 m elevation in Zion Canyon and south of St. George {Higgins s.n., 12 Apr 1961; DIX), Washington Coun- ty; Arizona. Our material belongs to var. zi- onensis (C. L. Hitchc.) Welsh & Reveal, Stat. & comb, nov., based on Draba zion- ensis C. L. Hitchc, Univ. Wash. Publ. Biol. 11:49. 1941. The var. asprella is restricted to Arizona. Draba aurea Vahl in Hornem. [D. luteola Greene; D. surculifera A. Nels.; D. aiirei- fortnis Rydb.; D. iiber A. Nels.; D. mccallae Rydb.; D. decwnbens Rydb.; D. aurea var. luteola (Greene) O. E. Schulz; D. aurea var. aureifonnis (Rydb.) O. E. Schulz; D. aurea var. deeumbens (Rydb.) O. E. Schulz; D. aureifonnis var. leiocarpa Payson & St. John] Plants perennial, not cespitose, stems 0.7-4 (5) dm tall, simple or few from a branching caudex, pubescent throughout with simple hairs often intermixed with forked ones; basal leaves 0.8-4 cm long, 0.2- 1.3 cm wide, oblanceolate, entire or serru- late, green or grayish, surfaces pubescent with stalked 4-rayed hairs; cauline leaves mostly 3-20, lanceolate to ovate or oblan- ceolate, 0.5-3 cm long, 0.3-1.2 cm wide, en- tire or less commonly denticulate, pubescent like the basal ones; racemes simple or branched, several- to many-flowered, much elongating in fruit; pedicels 3-15 (20) mm long, ascending to erect, pubescent; sepals 2-3.5 mm long, greenish, pubescent; petals (3.5) 4-6 mm long, yellow, spatulate to obo- vate, rounded to emarginate; silicles 8-17 mm long, plane or more commonly con- torted, 2-4 mm wide, ovate-lanceolate to lanceolate or elliptic, pubescent with simple, branched, or stellate hairs; styles (0.3) 0.8-1.3 (1.5) mm long; seeds 20-50.- Woodlands, grasslands, and among shrubs from 2300 to 3800 m elevation in Beaver, Carbon, Daggett, Duchesne, Garfield, Piute, Salt Lake, San Juan, Sevier, Summit, Uin- tah, and Utah counties; Alaska and Yukon east to Labrador and Greenland, south to Arizona and New Mexico. This is a highly variable taxon in our region. Draba brachystylis Rydb. Plants (annual) biennial or short-lived perennial, not cespi- tose, stems (1) 1.3-2.5 (3) dm tall, usually branched, pubescent throughout with simple, branched, or stellate hairs; basal leaves 1-5.3 cm long, 0.3-1.5 cm wide, ob- lanceolate, entire or denticulate, surfaces pubescent with stalked 2— 5-rayed hairs; cauline leaves (1) 2-8, ovate to obovate, el- liptic, or lanceolate, 0.6-2.8 cm long, 0.2-1.1 cm wide, denticulate to entire, pubescent like the basal ones; racemes simple or branched, several- to many-flowered, elon- gating in fruit; pedicles 1-10 mm long, spreading to spreading-ascending, pub- escent; sepals 2-2.7 mm long, yellowish, pubescent; petals 2.8-3.8 mm long, yellow, spatulate, rounded to emarginate; silicles (7) 10-15 mm long, 2-3.5 mm wide, plane, obliquely oblong-elliptic, pubescent with simple and branched hairs; styles 0.4-0.8 mm long; seeds 20-30.— Woodlands and shrublands from 1675 to 2130 m elevation in Salt Lake and Utah counties; Spring Mountains, Clark County, Nevada. This is a poorly known and rarely collected plant with affinities to both D. aurea and D. rec- tifructa. The type is from the Wasatch Moimtains. Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 319 Draba crassa Rydb. [D. chn/santha var. crassa (Rydb.) O. E. Schulz] Plants per- ennial, not cespitose, stems 0.8-1.8 dm tall, arising from a thickened caudex clothed with numerous marcescent leaf-bases, mod- erately hairy with simple or branched hairs, at least below; basal leaves 1.5-8 cm long, 0.3-1.2 cm wide, elliptic to oblanceolate, entire or obscurely toothed, glabrous or merely ciliate; cauline leaves 2-8, ovate to elliptic or obovate, 0.5-2 cm long, 0.2-0.8 cm wide, entire or nearly so, usually glabrous; racemes few- to many-flowered, elongating in fruit; pedicles 3-10 mm long, ascending, softly villous; sepals 2.3-3 mm long, greenish, suffused with purple, pub- escent; petals 4-8 mm long, yellow, obovate, rounded; silicles 8-15 mm long, 2.5-4 (5) mm wide, plane or contorted, ovate to lan- ceolate or elliptic, glabrous; styles 0.8-1.2 mm long; seeds 14-26.— Alpine tundra from 3500 to 3800 m elevation in the Uinta Mountains of Summit County but to be ex- pected in Daggett, Duchesne, and Uintah coimties; Montana, Wyoming, and Colo- rado. Draba crassifolia R. Graham. [D. parriji Rydb.; D. crassifolia var. parnji (Rydb.) O. E. Schulz] Plants (annual) biennial or short- lived perennial, not cespitose, stems 0.2-1.2 (2) dm tall, arising from a resulate tuft of leaves, usually glabrous except for a few hairs near base; basal leaves 0.3-1.5 (2.3) cm long, 0.1-0.3 cm wide, narrowly spatulate, entire, surfaces usually glabrous, sometimes ciliate; cauline leaves lacking or 1 or 2, very small; racemes 2- to several-flowered, congested or elongating in fruit; pedicels 2- 10 mm long, curved-ascending, glabrous; sepals 1-1.4 mm long, greenish, glabrous; petals 1.7-2.5 mm long, yellow but rarely fading white, elliptic-spatulate, emarginate; silicles 5-10 mm long, 1.5-2.5 mm wide, plane, glabrous, lance-elliptic; styles up to 0.5 mm long; seeds 10-60.— Alpine tundra from .3050 to 3800 m elevation in Duchesne (US), Grand, Piute, Salt Lake, San Juan, Sanpete (US), Sevier, and Summit counties; widespread from Alaska and Yukon east to Greenland and south to California, Arizona, and Colorado; Europe. This entity appar- ently grades with some phases of D. steno- loba in Utah, from which it is difficult, if not impossible, to segregate all specimens. Draba cuneifolia Nutt. ex Torr. & Gray. Plants annual, not cespitose, 0.1-1.5 (2) dm tall, very short leafy stems arising from a taproot, simple or branched, more or less dendritic-hairy throughout; basal leaves 0.5- 4 cm long, 0.2-2.7 cm wide, suborbicular to oblanceolate or cuneate-spatulate, dentate to entire, surfaces hirsute with stalked 2—4- rayed hairs, sometimes intermixed with simple hairs; cauline leaves few to several, usually much reduced, pubescent like the basal ones; racemes 3- to many-flowered, congested or elongatirig in fruit; pedicels 1- 7 mm long, spreading to ascending, dendrit- ic-hairy; sepals 1.5-2.5 mm long, greenish, pubescent; petals 3-4.5 (5) mm long, occa- sionally small or even lacking in cleisto- gamous flowers, white, spatulate, rounded to emarginate; silicles 4-13 (15) mm long, 1.8-3.8 (5) mm wide, plane, strigose, oblong- elliptic; styles up to 0.2 mm long; seeds 20 or more.— Warm desert shrublands upwards to mountain brush, pinyon-juniper, and pon- derosa pine communities, from 750 to 2300 m elevation in much of Utah; Washington and Idaho south to Mexico, Texas, and Ar- kansas. Two varieties are present. 1. Racemes compact in fruit, seldom half as long as plant height D. cuneifolia var. cuneifolia Racemes much elongated in fruit, commonly at least half plant height D. cuneifolia var. platijcarpa Var. cuneifolia. [D. helleri Small; D. am- mophila Heller; D. cuneifolia var. helleri (Small) O. E. Schvilz; D. cuneifolia var. leio- carpa O. E. Schulz] This is our common variant, being in Beaver, Box Elder, Grand, Kane, Millard, San Juan, Sevier, Uintah, Utah, and Washington counties; Colorado west to California and south to Mexico and Texas. Var. platycarpa (Torr. & Gray) S. Wats. 320 Great Basin Naturalist Vol. 37, No. 3 [D. plattjcarpa Torr. & Gray; D. viperensis St. John] Our few specimens from Washing- ton Coimty are tentatively assigned to this taxon, but they differ inter alia in having longer and narrower silicles; Arkansas west- ward to California, north to Washington. Draba densifolia Nutt. ex Torr. & Gray. Plants perennial, pulvinate-cespitose and matted, scapose, arising from compacted caudex branches clothed with marcescent leaf-bases, scapes 0.5-1.5 dm tall, glabrous to pubescent throughout; leaves 0.2-0.9 cm long, mostly 0.1-0.3 cm wide, oblong to ob- lanceolate, surfaces glabrous or with few- forked or dendritic hairs beneath, more or less ciliated with stiff, coarse, simple or forked hairs; racemes 2- to 10 (or more)- flowered, not elongated in fruit; pedicels 0.5-2 mm long, ascending, glabrous; sepals 2-3 mm long, greenish, glabrous or pub- escent; petals 2-6 mm long, yellow, obovate, truncate to emarginate; silicles 2-7 mm long, 2-3.5 mm wide, ovate to elliptic, glabrous; styles 0.2-1 mm long; seeds 2-12.— Alpine tundra from 3050 to 3800 m eleva- tion of Wasatch and Uinta mountains in Daggett, Duchesne, Salt Lake, Summit, and Uintah counties; British Columbia and Mon- tana south to California, Nevada, Utah, and Wyoming. Two weakly defined, sympatric variants are recognized. 1. Styles 0.2-0.5 mm long; plants glabrous except for cilia of the leaves D. densifolia var. daviesiae Styles 0.5-1 mm long; plants sometimes hairy on the lower leaf surfaces and on the scape D. densifolia var. densifolia Var. daviesiae (C. L. Hitchc.) Welsh & Reveal, comb, nov., based on Draba apicu- lata var. daviesiae C. L. Hitchc, Univ. Wash. Publ. Bot. 17(2):489. 1964. [D. apicu- lata C. L. Hitchc] This is a poorly differen- tiated phase of alpine sites in Duchesne, Salt Lake, Summit, and Uintah counties; Wyoming, Montana. The type of D. apicu- lata is from LaMotte Peak, Uinta Moun- tains, Utah. Var. densifolia. [D. glacialis var. pecti- nata S. Wats.; D. midfordae Payson; D. nel- sonii Macbride & Payson; D. globosa Pay- son; D. sphaerula Macbride & Payson; D. pectinata (S. Wats.) Rydb.; D. caeruleomon- tana Payson & St. John; D. caeruleomon- tana var. piperi Payson & St. John; D. den- sifolia f. nelsonii (Macbride & Payson) O. E. Schulz; D. globosa var. sphaerula (Macbride & Payson) O. E. Schulz] This is the com- mon phase of the species in alpine sites of Daggett, Duchesne, Salt Lake, Summit, and Uintah counties; distribution of the species. Draba fladnizensis Wulfen in Jacq. [D. pattersonii O. E. Schulz; D. pattersonii var. hirticaidis O. E. Schulz; D. pattersonii var. dasijcarpa O. E. Schulz] Plants perennial, not cespitose, stems 0.2-0.9 dm tall, glabrous or pubescent at least near the base with simple or forked hairs; basal leaves 0.3-1 cm long, 0.1-0.2 cm wide, oblanceol- ate, surfaces glabrous or moderately hairy with 1— 2-forked hairs, cilitate; cauline leaves 1 or 2, greatly reduced; racemes 3-to several-flowered; pedicels 1-3 mm long, as- cending to spreading, glabrous; sepals 1.2- 1.8 mm long, greenish, glabrous; petals 1.8- 2.5 mm long, white (rarely pink), spatulate, rounded to retuse; silicles 3-6 mm long, 1.2- 2 mm wide, oblong-ovate, glabrous; styles essentially lacking; seeds 10-20.— Alpine tundra in Uinta and La Sal mountains above 3100 m elevation in Daggett (?), Grand, San Juan, and Uintah (?) counties; Alaska to Mackenzie south to Colorado and Utah. Our material is difficult to interpret from the dwarf alpine specimens of D. stenoloba (q.v.), but apparently the white petals are diagnostic. Draba incerta Payson. [D. laevicapsula Payson; D. incerta var. laevicapsuki (Pay- son) Payson & St. John] Perennial, cespitose but loosely so, 0.2-1.5 (2) dm tall, stems pubescent with stellate or dendritic and sometimes simple hairs; basal leaves 0.5-1.8 (2.5) cm long, 0.1-0.3 cm wide, narrowly oblanceolate, surfaces with at least some doubly pectinate hairs, often intermixed with other types of pubescence, margin ci- liate with simple to pectinately branched Sept. IQ?' Welsh, Reveal: Utah Flora, Brassicaceae 321 hairs; cauline leaves 1 or lacking; racemes 5- to many-flowered, elongating in fruit; pedicels 2-12 mm long, ascending, usually hairy; sepals 2.5-3.5 mm long, greenish or suffused with purple, pubescent; petals 4-5.5 mm long, yellow but fading cream, cuneate- obovate, broadly emarginate; silicles 6-10 mm long, 2.5-3.5 mm wide, ovate to lan- ceolate, plane, pubescent or glabrous; seeds 8-14.— Alpine sites above 3050 m elevation in Raft River Mountains of Box Elder Coun- ty; Alaska and Yukon south to Washington, Utah, and Wyoming. Draba lanceolata Royle. [D. cana Rydb.; D. valida Goodding] Plants perennial, loose- ly cespitose, caudex simple or branched, stems 0.5-3.5 dm tall, pubescent throughout with soft many-branched hairs; basal leaves 0.5-4 cm long, 0.1-0.4 cm wide, oblanceo- late, entire, pubescent of overlapping, stel- late or branched hairs; cauline leaves sever- al, commonly toothed; racemes several- to many-flowered, sometimes with solitary flowers in upper leaf axils; pedicels 2-9 mm long, erect, usually appressed to rachis, pub- escent; sepals 1.5-2 mm long, sparsely pi- lose; petals 2.2-4 mm long, white, cuneate- obovate, more or less emarginate; silicles 5- 12 mm long, 1.5-2.5 mm wide, narrowly lanceolate to oblong, plane or contorted, softly pubescent, rarely glabrous; styles 0.2- 0.8 mm long; seeds 20 or more.- Alpine meadows and krumholz in the Uinta and Henry mountains mostly above 3000 m ele- vation, in Daggett, Duchesne, Garfield, Summit, and Uintah counties; Alaska and Yukon south to Nevada, Utah, and Colo- rado. The type of D. valida is from Dyer Mine, Uintah County. Draba lonchocarpa Rydb. Plants per- ennial, loosely to densely cespitose, caudex usually branched, scapose or rarely with one cauline leaf, scape 0.1-1.2 dm tall, glabrous or pubescent with soft many-branched hairs; leaves 0.5-1.5 cm long, 0.1-0.4 cm wide, pubescent with usually overlapping or stel- late, rarely some simple, hairs, marginal pubescence entirely stellate or with some simple hairs; racemes 3— 12-flowered, con- tracted or elongating in fruit; pedicels 1-6 (11) mm long, ascending to erect, glabrous; sepals 1.5-2 mm long, glabrous or pub- escent; petals 2.5-4 mm long, white; silicles 5-14 mm long, 1-2 mm wide, linear to lance-linear or oblong, plane or twisted, glabrous or pubescent; styles 0.2-0.5 mm long; seeds 8-30.— Alpine tundra in Wasatch and Uinta mountains; Alaska and Yukon south to Oregon, Utah, and Colorado. Our material has been treated as portions of an expanded D. tiivalis Lilj., from which it differs in technical features of pubescence and silicle characteristics. Two sympatric variants have been designated among the Utah material. 1. Silicles mostly less than 7 mm long, elliptic to linear D. lonchocarpa var. exigua Silicles mostly more than 10 mm long, linear to narrowly elliptic D. lonchocarpa var. lonchocarpa Var. exigua O. E. Schulz. [D. nivalis var. exigua (O. E. Schulz) C. L. Hitchc] Alpine tundra in Uinta and La Sal mountains of Duchesne, Grand, and Summit counties; Wyoming and Colorado. Var. lonchocarpa. [D. nivalis var. elon- gata S. Wats.; D. lonchocarpa var. dasij- carpa O. E. Schulz; D. lonchocarpa var. ves- tita O. E. Schulz; D. lonchocarpa var. semitonsa St. John] Alpine sites from 3050 to 3900 m elevation in the Wasatch, Uinta, and La Sal mountains of Cache, Duchesne, Grand, Salt Lake, San Juan, and Summit counties; Alaska south to Oregon, Utah, and Wyoming. Draba maguirei C. L. Hitchc. Plants per- ennial, cespitose, with substoloniferous branches, scapose, scapes 0.2-2 dm tall, glabrous or with a few forked hairs near base; leaves 0.3-1.5 cm long, 0.1-0.4 cm wide, oblong-oblanceolate to obovate-oblan- ceolate, surfaces glabrous or nearly so, ci- liate with simple, forked, or 4-rayed, shortly stalked hairs; racemes few- to several-flow- ered, elongating in fruit; pedicels 2-10 (15) mm long, ascending, glabrous; sepals 2-3 mm long, yellowish, glabrous; petals 4.5-6 mm long, yellowish, spatulate, rounded; si- licles 4-9 mm long, 2-3.5 mm wide, ovate to lanceolate, oblique, glabrous or scabe- 322 Great Basin Naturalist Vol. 37, No. 3 nilous; styles 1-1.5 mm long; seeds 2-8.— tenuous and somewhat arbitrary varieties Talus slopes and rocky outcrops from 2600 have been designated, to 2900 m elevation in northern Utah. Two 1. Hairs of leaves all simple or rarely some forked D. maguirei var. burkeri Hairs of leaves all branched, mostly 4-rayed D. maguirei var. maguirei Var. burkeri C. L. Hitchc. Cottonwood Canyon, Wellesville Mountains, Box Elder County (type locality), and mountains east of Ogden, Weber County; endemic. Var. maguirei. Apparently restricted to the Bear River Range, Cache County (type from Mt. Naomi); endemic. Draba nemorosa L. [D. dictyota Greene] Annual, from a slender taproot, stems simple or branched, 0.5-2.5 dm tall, pub- escent with mixed forked and stellate hairs, or less commonly with some simple ones, or even glabrate; leaves 0.3-3 cm long, 0.2-0.8 cm wide, oblanceolate to lanceolate, ovate or oblong, entire or toothed, pubescent with branched or simple hairs; racemes few- to many-flowered, much elongating in fruit; pedicels 5-25 mm long, spreading-ascending, glabrous; sepals 1-1.5 mm long, green to yellowish or suffused purple, pilose to glabrous; petals 1.2-4 mm long, yellow to white; silicles 4-10 mm long, 1.5-3 mm wide, oblong to oblanceolate or elliptic, plane, glabrous; styles obsolete; seeds 25 or more.— Roadsides, foothills, and dry exposed sites from 1375 to 2250 m elevation in Box Elder, Cache, Salt Lake, Summit, Uintah, Utah, Weber, and perhaps all other coun- ties; common throughout much of North America and Eurasia. Draba oligospermia Hook. Perennial, ces- pitose, the caudex much branched, scapose, the scapes 0.1-1 dm tall, pubescent through- out with pectinate or stellate hairs or glabrous, at least below; leaves 0.3-1.2 cm long, 0.1-0.2 cm wide, linear to spatulate or oblong, surfaces (one or both) pubescent with sessile, appressed, doubly pectinate- branched hairs, commonly ciliate with at least some pectinately branched hairs; ra- cemes 2— 15-flowered, only moderately elongating in fruit; pedicels 1-10 mm long, ascending, glabrous or pubescent; sepals 1.5- 2.5 mm long, yellowish, pubescent; petals 3- 5 mm long, yellow or white, obovate, rounded to emarginate; silicles 3-8 mm long, 2-4 mm wide, ovate to oval or oblong, plane, glabrous to pubescent; styles 0.1-1.2 mm long; seeds 2-10.— Ponderosa pine and Douglas fir woodlands upwards to alpine tundra in the Wasatch and Uinta Mountains from 2290 to 3800 m elevation; British Co- lumbia and Alberta south to California, Utah, and Wyoming. Two varieties are known. 1. Petals yellow; pubescence of silicles, when present, of simple or forked hairs only; plants widespread D. oligospertmi var. oligosperma Petals evidently white; pubescence of silicles at least in part of doubly pecti- nate hairs; plants of Daggett County D. oligospermo var. pectinipila Var. oligosperma. [D. oligosperma var. andina (Nutt.) ex Torr. & Gray: D. andina (Nutt.) A. Nels.; D. saximontana A. Nels.; D. oligosperma var. microcarpa Blankinship; D. oligosperma var. saximontana (A. Nels.) O. E. Schulz; D. oligosperma var. leiocarpa O. E. Schulz. Widespread in the Wasatch and Uinta mountains in Cache, Daggett, Duchesne, Summit, Uintah, and Utah coun- ties; range of the species. Var. pectinipila (Rollins) C. L. Hitchc. [D. pectinipila Rollins] Vicinity of Flaming Gorge, Daggett County, where possibly ex- tirpated; Wyoming. Draba rectifructa C. L. Hitchc. [D. mon- tana S. Wats., not Bergeret] Annual, from a slender taproot, stems simple or branched, 1-2.2 dm tall, pubescent throughout al- though sparsely so above in some with branched and/or simple hairs; leaves 0.5-3 cm long, 0.1-0.7 cm wide, lanceolate to ob- lanceolate, entire, pubescent with branched Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 323 and simple hairs intermixed; racemes sever- al- to many-flowered, much elongating in fruit; pedicels 2-6 mm long, spreading to curved-ascending, pubescent; sepals 1.2-1.7 mm long, greenish, pubescent; petals 2-4 mm long, yellow, narrowly spatulate, usual- ly emarginate; silicles (4) 6-10 mm long, 2- 2.5 mm wide, obliquely oblong, pubescent; styles obsolete; seeds 40 or more.— Road- sides, foothills, meadows, and woodlands, mostly from 2200 to 3050 m elevation in Carbon, Duchesne, Garfield, Grand, Piute, Salt Lake, and Wasatch counties (and to be expected elsewhere); Colorado, New Mexi- co, and Arizona. Draba reptans (Lam.) Fern. Annual, from a slender taproot, 0.2-1 (2) dm tall, very short leafy stem simple or sometimes branched, pubescent with simple to stellate hairs below, glabrous above; basal leaves 0.3-1.8 cm long, 0.1-1 cm wide, spatulate to ovate or obovate, usually entire, surfaces pubescent with branched or forked hairs or upper surface only of simple hairs; cauline leaves few, usually reduced, pubescent like basal ones, or upper surface with simple hairs; racemes (1-) several- to many- flowered, compact in fruit; pedicels 1-6 mm long, ascending, glabrous; sepals 1.5-2.5 mm long, greenish or yellowish, usually pub- escent; petals 3-5 mm long, white, ovate, rounded, sometimes smaller or lacking in cleistogamous flowers; silicles 5-20 mm long, 1-2 mm wide, oblong, nearly erect, pubescent or glabrous; styles up to 0.15 mm long; seeds 15 or more.— Dry exposed sites from 1200 to 2000 m elevation in the northern two-thirds of Utah (but possibly cosmopolitan); throughout much of northern North America. Two more or less arbitrary varieties are present. Draba reptans is sim- ilar to, and not always distinct from, D. ciineifolia (q.v.). Upper surface of the cauline leaves and one or both surfaces of the basal leaves pubescent with mostly simple hairs D. reptans var. reptans Upper surface of all leaves predominantly pubescent with forked or branched hairs D. reptans var. steUifera Var. reptans. [Arabis reptans Lam.; D. ca- roliniana Walter; D. micrantha Nutt. ex Torr. & Gray; D. coloradoensis Rydb.; D. reptans var. micrantha (Nutt.) Fern.] Range of the species. Var. SteUifera (O. E. Schulz) C. L. Hitchc. [D. carolinana f. steUifera O. E. Schulz; D. carolinana ssp. steUifera (O. E. Schulz) Payson & St. John] Sympatric with the former. Draba sobolifera Rydb. [D. uncinalis Rydb.; D. soboUfera var. uncinalis (Rydb.) O. E. Schulz] Perennial, cespitose, caudex branched, scapose, scapes 0.1-0.6 dm tall, pubescent with intermixed stellate, iDranched, and simple hairs; leaves 0.8-2 cm long, 0.2-0.5 cm wide, obovate to oblanceol- ate, pubescent with stalked stellate or 4- rayed hairs at least beneath, and usually ci- liate with simple hairs at base; racemes (2-) 5— 20-flowered, compact to elongating in fruit; pedicels 3-8 mm long, stellate or with branched hairs; sepals 1.8-2.5 mm long; pet- als 4-5 mm long, yellow, obovate; silicles 3- 8 mm long, 2.5-4 mm wide, ovate to ellip- tic, pubescent to glabrous; styles 0.4-1 mm long; seeds 4-12.— Sagebrush communities upwards to alpine tundra, from 2290 to 3660 m elevation in Tushar Mountains and on Markagaunt Plateau in Beaver, Garfield, and Piute counties; endemic. The types of both Draba sobolifera and D. uncinalis were collected by Marcus E. Jones in the Delano Peak area west of Marysvale, Piute County. Draba spectabilis Greene. [D. spectabilis var. glabrescens O. E. Schulz] Perennial, not cespitose, caudex branched, stems mostly 1- 4 dm tall, usually simple, pubescent with simple or forked hairs; basal leaves (0.5) 1-4 cm long, 0.2-1 cm wide, obovate to spatu- late, subentire to denticulate, green, surfaces with subsessile 4-rayed or forked hairs, in- frequently with the upper surface glabrous; cauline leaves mostly 3-15, ovate to lanceo- late, subentire to sharply toothed, 0.5-2 cm long, 0.2-1.5 cm wide, pubescent like basal ones; racemes several- to many-flowered. 324 Great Basin Naturalist Vol. 37, No. 3 much elongating in fruit; pedicels 5-15 mm long or more, ascending, glabrous or hairy; sepals 2-3.5 mm long, yellowish, pubescent; petals 4.5-7 mm long, yellow but fading white, elliptic, rounded; silicles 5-14 mm long, 2-3 mm wide, lanceolate to ovate, plane, glabrous or pubescent; styles 0.8-2.5 mm long; seeds 10-20.— Opening in woods, at moderate to high elevations in La Sal and Abajo mountains of Grand and San Juan counties; Colorado, New Mexico, Ari- zona. Our material is var. spectahilis. The type of var. glabrescens was obtained in the La Sal Mountains. Draba stenoloba Ledeb. [D. nemorosa var. stenoloba (Ledeb.) M. E. Jones; D. ni- tida Greene; D. deflexa Greene; D. nitida var. nana O. E. Schulz; D. nitida var. prae- longa O. E. Schulz] Plants (annual?) biennial or short-lived perennial, from a taproot, caudex more or less developed, stems 0.3- 2.5 dm tall, glabrous or sometimes hirsute below; basal leaves 0.3-4 cm long, 0.2-0.8 cm wide, entire to denticulate, pubescent with simple to forked or branched hairs, or with one or both surfaces glabrous on some leaves; cauline leaves (0) 1-8, ovate to lan- ceolate or elliptic, entire or denticulate, similar to the basal ones in all respects; racemes several- to many-flowered, elongat- ing in fruit; pedicels 1-14 mm long, ascend- ing, glabrous; sepals 1-2.2 mm long, green- ish, glabrous; petals 2-4.5 mm long, yellow to cream or fading white, spatulate, rounded to emarginate; silicles 6-18 mm long, 1.5-2.2 mm wide, linear to oblong or elliptic, usually glabrous; styles up to 0.2 mm long; seeds 16 or more.— Woods, mead- ows, sagebrush communities and along stream banks from 2075 to 3350 m eleva- tion in Beaver, Box Elder, Cache, Daggett, Duchesne, Emery, Garfield, Piute, Salt Lake, Sevier, Summit, Uintah, Utah, Wasatch, and Washington counties; Alaska and Yukon south to California, Nevada, Utah, and Colorado. Draba subalpina Goodman 6c Hitchc. [D. oriebata of Utah references, not Macbride & Payson] Perennial, cespitose, caudex simple or branched, clothed with marcescent leaves, scapose, scapes 0.1-1.2 dm tall, rarely with one cauline leaf, glabrous throughout or pubescent with simple or forked hairs at least near base; leaves 0.3- 1.8 cm long, 0.1-0.4 cm wide, oblong to spatulate, surfaces glabrous or sparingly hir- sute, ciliate with coarse simple or less com- monly forked or branched hairs; racemes few- to many-flowered, only moderately elongating in fruit; pedicels (1) 2-10 mm long, usually purplish, glabrous; petals 4-5 mm long, white, cuneate-spatulate, emargi- nate; silicles 3-8 mm long, 2-4 mm wide, ovate to elliptic, plane or more or less con- torted, glabrous; styles 0.6-1 mm long; seeds 6-12.— SpRice-fir, Douglas fir, or bristlecone pine woodlands mostly on the Pink Lime- stone member of the Wasatch Formation, from 2130 to 3050 m elevation in Garfield, Iron, and Kane counties; endemic. The type is from Cedar Breaks, Iron County. Draba ventosa A. Gray. Perennial, cespi- tose, the caudex usually branched, more or less clothed with marcescent leaves, scapose, scapes 0.2-0.4 dm tall, pubescent with simple and forked or sometimes stellate hairs; leaves 0.5-1.2 cm long, 0.2-0.4 cm wide, elliptic to lanceolate, surfaces pub- escent with simple, forked, or branched to stellate hairs; racemes 3- to many-flowered, little elongating in fruit; pedicels mostly 4-8 mm long, ascending, densely pilose to stel- late; sepals 2-2.5 mm long, greenish or yel- lowish, pilose; petals 4-5 mm long, yellow, obovate; silicles 5-8 mm long, 3.5-5.5 mm wide, oval to ovate, plane, densely hairy; styles 0.6-1.2 mm long; seeds 10-16.- Al- pine tundra from 3050 to 3800 m elevation in the Uinta Mountains, in Summit (and perhaps elsewhere) County; Wyoming. This species is poorly known and is sel- dom collected. Our description is tentative at best, being drawn on only limited mate- rial. The Utah plants are supposedly assign- able to var. ventosa. Draba verna L. Diminutive annual from a slender taproot, scapose, scapes 0.2-0.5 (1.2) diT» tall, glabrous throughout or pub- escent near base only; leaves 0.1-1 (2) cm long, 0.08-0.3 cm wide, spatulate to oblan- ceolate, entire or toothed, pubescent with branched hairs; racemes few- to many-flow- ered, elongating in fruit; pedicels 2-12 mm Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 325 long, ascending, glabrous; sepals 0.6-1.1 mm long, greenish or tinged purplish, glabrous or pubescent; petals 1.5-2.5 mm long, white, deeply bilobed; silicles 2-10 mm long, 1-4 mm wide, elliptic to obovate, glabrous; styles up to 0.2 mm long.— Dry open sites from 1375 to 1680 m elevation in Salt Lake and Weber counties; widespread in North America; Asia. Our material is scanty and cannot be seg- regated into the two variants reported for North America. Erysimum L. Plants pubescent with 2-3 (4)-rayed hairs, annual to perennial, from taproots; leaves alternate or basal and still alternate, simple. entire to toothed, not auriculate; flowers racemose, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, yellow to or- ange pr burnt-orange to purple; stamens 6, filaments lacking glandular processes; style prominent, short to elongate, stigma bi- lobed; fruit a silique, many times longer than broad, compressed parallel to partition or subterete, valves 1- to several-nerved; seeds uniseriate, many per locule. A genus of 100 species, mainly of Eu- rope. Rossbach, G. B. 1958. The genus Erysimum (Cruciferae) in North America north of Mexico— A key to species and varieties. Madrorio 14:261-267. 1. Petals (10) 12-20 mm long or more; styles mostly 1.5-3 mm long; plants bien- nial or perennial; widespread in indigenous plant communities E. asperu7n Petals 3.5-11 mm long; styles usually less than 1.5 mm long (longer in E. re- pandum); plants annual or biennial to short-lived perennial; restricted and weedy or less commonly in indigenous plant communities 2 2(1). Petals 3.5-5 mm long; siliques 12-27 mm long; plants uncommon, annual weeds of moist sites E. chieranthoides Petals mostly 5-11 mm long; siliques (15) 25-100 mm long; plants common, annual, biennial or perennial, of various habitats 3 3(2). Pedicels usually more than 5 mm long, more slender than fruit; siliques as- cending to erect, less than 50 mm long; plants indigenous biennial or short- lived perennial E. inconspicuum Pedicels 2-5 mm long, almost or quite as thick as fruit; siliques spreading to curved-ascending, at least some often more than 50 mm long; plants adven- tive, annual weeds E. repandum Erysimum asperum (Nutt.) DC. Wallflow- er. Biennial or short-lived perennial, with simple or less commonly branched caudex, stems 1.2-8.5 (10) dm tall or more; basal leaves 2-10 (12) cm long, 0.2-1.4 cm wide, sublinear to elliptic or spatulate, entire or denticulate, grayish to green, pubescent with malpighian or Y-shaped appressed hairs; cauline leaves 1.1-10.4 cm long, 0.1- 1.5 cm wide, much reduced to little if at all reduced upwards, variously shaped, entire or toothed, pubescent like basal ones; race- mes much elongating in fruit; pedicels 3-17 mm long, spreading-ascending to ascending, usually more slender than the fruit; sepals 7.5-14 mm long, yellowish or purplish; pet- als 12-28 mm long, yellow to yellow-orange or burnt-orange; siliques (17) 20-115 mm long, 1-2.5 mm thick, subquadrangular to somewhat flattened, ascending to erect or less commonly spreading-ascending; styles 1- 4 (5) mm long; seeds wingless or winged only near tip, 1.5-2.3 mm long.— Warm desert shrub, cool desert shrub, mountain brush, pinyon-juniper, ponderosa pine, as- pen, spruce, fir, and Douglas fir to alpine tundra communities, throughout Utah; Yu- kon Territory south to California and Ari- zona, and eastward to Oklahoma, Kansas, and Minnesota. 326 Great Basin Naturalist Vol. 37, No. 3 Flowers yellow or yellowish; fruits erect or nearly so; common E. asperwn var. purshii Flowers orangish to burnt-orange; fruits ascending to spreading-ascending; common E. aspenim var. amoenum Var. amoenum (Greene) Reveal. [Chei- ranthus nivalis var. amoenus Greene; Chei- rinia amoena (Greene) Rydb.; E. wheeleri Rothrock; Cheirinia wheeleri (Rothrock) Rydb.] Common throughout southern por- tions of state; California eastward to Colo- rado. Var. purshii Durand. [Cheiranthus capi- tatus Dougl. ex Hook.; E. elatum Nutt. ex Torr. & Gray; E. capitatum (Dougl.) Greene; Cheiranthus elatus (Nutt.) Greene; Cheiranthus asperrimus Greene; Cheiranthus argillosus Rydb.; Cheiranthus bakeri Greene; Cheirinia elata (Nutt.) Rydb.; E. asperrimum (Greene) Rydb.; E. oblanceolatum Rydb.; E. bakeri (Greene) Rydb.; E. aridum A. Nels.; E. capitatum var. argilhsum (Greene) R. J. Davis] Common throughout most of state, but particularly common in northern por- tion; Yukon Territory south to California and Arizona, east to Montana, Wyoming, Colorado, and New Mexico. Erysimum cheiranthoides L. [Cheiritiia cheiranthoides (L.) Link; Cheiranthus chei- ranthoides (L.) Heller] Annual, stems simple or branched, 2-12 dm tall; leaves 2-8 cm long, 0.2-1.5 cm wide, linear to oblong, lan- ceolate or oblanceolate, entire or denticu- late, green, pubescent with malpighian or Y-shaped hairs; racemes much elongating in fruit; pedicels 4-15 mm long, spreading-as- cending, very slender; sepals 2-3 mm long, yellowish or greenish; petals 3-5 mm long, pale yellow; siliques 12-27 mm long, about 1 mm wide, subterete, ascending to erect; styles 0.8-1 mm long; seeds 1-1.2 mm long, not winged.— Moist places, in meadows and along roadsides in Cache and Wasatch counties, and to be expected elsewhere; widespread in North America; adventive from Eurasia. This plant is evidently un- common to rare in Utah. Erysimum inconspicuum (S. Wats.) Mac- Millan. [E. asperum var. inconspicuum S. Wats.; E. parviflorum Nutt. ex Torr. & Gray, not Pers.; E. syrticohim Sheldon; Cheiranthus inconspicuus (S. Wats.) Greene; Cheirinia inconspicua (S. Wats.) Rydb.; Cheirinia syrticola (Sheldon) Rydb.] Biennial or short-lived perennial with usually un- branched caudex, stems mostly 2-10 dm tall, usually simple; leaves 1.5-8 cm long, 0.2-0.8 cm wide, linear to oblong, elliptic, lanceol- ate or oblanceolate, pubescent with malpi- ghian or Y-shaped hairs; racemes elongating in fruit; pedicels 3-8 mm long, ascending; sepals 4-7 mm long, greenish or purplish; petals (6) 7-10 (11) mm long, pale to bright yellow; siliques 15-50 mm long, 1-2 mm thick, quadrangular, erect or ascending; styles 1-1.5 mm long; seeds about 1.5 mm long, not winged.— Sagebrush and aspen- spruce communities at middle elevations in Garfield, Wasatch, and Washington coun- ties, and to be expected elsewhere; Alaska and Yukon south to Oregon, Utah, and Col- orado, and east to central Canada and the north-central states. This species resembles E. asperum in general habit, but is more re- stricted in its distribution in Utah. Erysimum repandum L. [Cheirinia re- panda (L.) Link] Annual, stems 0.8-3 (5) dm tall, simple to much-branched; leaves 0.7-11 (15) cm long, 0.1-0.8 (1.2) cm wide, oblong, oblanceolate to linear, pubescent with mal- pighian and Y-shaped hairs; pedicels 2-5 mm long, spreading, almost or quite as thick as the fruit; sepals 3.5-5.8 mm long, yellowish or greenish; petals 5.2-8 mm long, yellow; siliques 26-85 mm long, 0.9-1.5 mm thick, quadrangular, spreading to ascending, rarely descending; styles 1-3 mm long; seeds ca 1 mm long, not winged.— Disturbed sites from 1300 to 1950 m elevation in Beaver, Grand, Juab, Millard, Salt Lake, Sanpete, Utah, and Washington counties, and to be expected elsewhere; widespread in North America; adventive fom Europe. EucLiDiuM R. Br. Nom. Cons. Plants pubescent with forked hairs, an- nual, from taproots; leaves alternate, simple, entire to remotely serrulate, petiolate to Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 327 subsessile, not auriculate; flowers solitary and axillary or subaxillary, or borne in elon- gate racemes, pedicels sometimes subtended by bracts; sepals 4, caducous; petals 4, white, minute; stamens 6; style very short atop beak of fruit, stigma bilobed; fruit a si- licic, bilocular, tardily dehiscent; seed 1 per locule. A Eurasian genus of 2 species. Euclidium syriacum (L.) R. Br. [Anasta- tica sijriaca L.] Plants 0.4-5 dm tall, simple to much-branched, pubescent with forked hairs; leaves mainly cauline, 0.7-6.5 cm long, 0.2-1.7 cm wide, oblanceolate to ellip- tic or lanceolate, pubescent; pedicels 0.5-1 mm long, ascending to erect, pubescent; sepals 0.7-0.9 mm long, tinged with purple; petals 0.8-1.1 mm long, white, spatulate, erect; silicles (including beak) 2.8-4 mm long, body 1.2-2 mm wide, pubescent with simple or forked hairs, beak 1.2-1.6 mm long; style short or obsolete.— Roadsides, va- cant lots, and dry foothills in Box Elder, Salt Lake, Summit, and Utah counties, and likely elsewhere; Idaho, Washington; adven- tive from Europe. Glaucocarpum Rollins Plants glabrous and glaucous, perennial from a branching caudex; leaves alternate, simple, entire or sparingly dentate, not au- riculate; flowers racemose, pedicels not sub- tended by bracts; sepals 4, deciduous; petals 4, yellow; stamens 6, paired ones united by filaments and anthers their entire length; style short, stigma capitate, entire, unex- panded; fruit a subsessile to sessile silique, many times longer than broad, slightly flat- tened, valves strongly 1 -nerved; seeds unise- riate. A monotypic genus endemic to Utah. Rollins, R. C. 1938. Glaucocarpum, a new genus in the Cniciferae. Madroiio 4:232- 235. Glaucocarpum suffrutescens (Rollins) Roll- ins. [Thelypodium suffrutescens Rollins in Graham] Plants perennial from a branching caudex forming small clumps, stems 1-2.5 dm tall, glabrous, lacking basal leaves; leaves (0.7) 1-2.5 cm long, 0.3-1 cm wide, elliptic to lanceolate or oblanceolate, short petiolate to subsessile, not auriculate; pe- dicles (2.5) 3-12 mm long, curved-ascending to erect, glabrous; sepals 4-6 mm long, yel- lowish or greenish; petals 9-11 mm long, yellow, spatulate, ascending to spreading; si- liques 10-20 mm long, 1.2-3 mm wide, flat- tened, erect, glabrous; styles (0.5) 1-2 mm long, stigma capitate, entire; seeds 4-8 per locule.— Local and infrequent on calcareous shale of the Green River Formation, west of Willow Creek in the vicinity of Big Pack Mountain, from 1645 to 1825 m elevation, Uintah County; endemic. This genus is the only one strictly endem- ic to the state. Halimolobos Tausch Plants pubescent with simple, forked, and branched hairs, (annual) biennial or per- ennial, arising from a taproot; leaves alter- nate or basal and still alternate, simple, dentate to subentire, tapering to base or cauline sessile and auriculate; flowers race- mose, pedicels ascending, not subtended by bracts; sepals 4; petals 4, white; stamens 6; styles prominent, stigma small, entire; fruit a sessile, slender silique, many times longer than broad, terete to quadrangular, valves strongly 1 -nerved; seeds biseriate, numerous. A genus of perhaps 15 species of North and South America. Rollins, R. C. 1943. Generic revisions in the Cniciferae: Halimolobos. Contr. Dudley Herb. 2: 241-265. Rollins, R. C. 1952. A note on Halimolobos. Rhodora 54:161-163. Halimolobos virgata (Nutt.) O. E. Schulz. [Sismybrium virgatwn Nutt. ex Torr. & Gray; Hesperis virgata (Nutt.) Kuntz; Ste- nophragma virgatwn (Nutt.) Greene; Arabis brebneriana A. Nels.; Pilosella virgata (Nutt.) Rydb.; P. stenocarpa Rydb.; Arabi- dopsis virgata (Nutt.) Rydb.; Arabidopsis stenocarpa (Rydb.) Rydb.] Biennial but occa- sionally flowering the first year, stems 1-3.5 dm tall, simple or branched, pubescent with mixed simple, forked, and branched hairs at least below; basal leaves 3-6 cm long, 0.5- 1.8 cm wide, oblanceolate to lanceolate, denticulate to dentate, rarely entire; cauline leaves several, reduced upwardly, at least uppermost both sessile and auriculate; pedi- cels 7-11 mm long, ascending, glabrous or 328 Great Basin Naturalist Vol. 37, No. 3 puberulent; sepals 2.5-3 mm long, greenish, pubescent; petals 4-4.5 mm long, white, veins often suffused with pink or purple, usually erect; siliques 15-40 mm long, most- ly 1-1.5 mm wide, subquadrangular, erect, valves glabrous, strongly nerved; styles 0.2- 0.5 mm long; seeds irregularly biserrate.— Grassy meadows and shrublands along the north slope of the Uinta Mountains in Dag- gett (US) and Summit counties; Colorado, Wyoming, Idaho, and northward to Alberta and Saskatchewan. The number of generic segregates listed above are an indication of the difficulties involved in placing this unusual species in a genus. It simulates an Arabis, from which it is outwardly separable only with difficulty, but on technical grounds is probably more closely allied to Sismybrium in a broad sense. Our Utah material, and that from southwestern Wyoming, differs slightly from that found elsewhere and may deserve vari- etal recognition. Hesperis L. Plants pubescent with simple and forked hairs, perennial, from taproots; leaves alter- nate, simple, sinuate-dentate or serrate, mainly cauline, petiolate to subsessile, not auriculate; flowers racemose, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, pink to lavender or white; stamens 6, filaments lacking glandular processes; styles obsolete, stigma bilobed, massive; fmit a silique, many times longer than broad, subterete, tardily dehiscent, valves 1 (3)-nerved; seeds uniseriate. A Eurasian genus of perhaps 24 species, many cultivated. Hesperis matronalis L. Dame's Violet; Sweet Rocket. Plants perennial with one to several stems, these simple or branched, mostly 5-12 dm tall or more, pubescent with mixed forked and simple hairs; leaves 2-15 (20) cm long, 0.6-3.5 (4) cm wide, ovate-lanceolate to elliptic or lanceolate, si- nuate-dentate to serrate, pubescent; pedicels 8-21 mm long, ascending to spreading, pub- escent; sepals 6.5-8 mm long, often suffused with red or purple, pubescent; petals 15-25 mm long, pink to lavender or white, obo- vate, spreading; siliques 31-100 mm long, 1- 2 mm wide, subterete, erect or ascending, puberulent; styles obsolete, stigma deeply bilobed; seeds numerous in each locule.— Cultivated ornamental, persisting and es- caping, now widely established in cultivated lands especially along irrigation canals at mid elevations in Cache, Utah, and Summit counties, and expected to be established elsewhere; introduced from Europe. HuTCHiNsiA R. Br. Plants glabrous, annual or winter annual, from taproots; leaves alternate, simple, en- tire to pinnatifid, petiolate to sessile, not auriculate; flowers racemose, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, white; stamens 6, filaments lacking glandular processes; styles obsolete or very short, stigma capitate, entire; fruit a silicle, only somewhat longer than broad, strongly compressed at right angles to septum, valves reticulately veined; seeds several. A genus of 6 Eurasian and 1 North Amer- ican species. Hutchinsia procumbens (L.) Desv. [Lepi- dium procumbens L.; Hijmenolobus divari- catus Nutt. ex Torr. & Gray; Hymenolobus erectiis Nutt. ex Torr. & Gray] Slender, di- minutive annuals, stems glabrous, simple or more commonly branched, 0.5-3 dm long, erect or prostrate; leaves basal and cauline, 0.5-3 cm long, 0.1-1.3 cm wide, ovate to lanceolate, oblanceolate, or nearly linear, entire to pinnatifid, petiolate to subsessile; pedicels 3-8 mm long, ascending to spread- ing-ascending, glabrous; sepals 0.7-1.1 mm long, greenish or purplish, glabrous; petals 0.8-1.3 mm long, white, spatulate, rounded to retuse; silicles 2.4-4.2 mm long, 1.5-2 mm wide, elliptic to obovate, truncate, rounded, or somewhat emarginate, glabrous; styles up to 0.2 mm long; seeds several per locule.— Moist to dry sites, hanging gardens, drain- age banks, sidewalks, roadsides, playas, and peat bogs from 850 to 1830 m elevation in Garfield, Salt Lake, San Juan, Utah, and Washington counties; widely distributed in North America; Eurasia. ISATIS L. Plants pubescent with long simple hairs at least below, biennial or short-lived per- Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 329 ennial, from strong taproots; leaves simple, alternate, basal petiolate, cauline hastately- auriculate; flowers racemose or paniculate, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, yellow; stamens 6, fila- ments lacking glandular processes; styles ob- solete, bilobed stigma sessile; fruit a silicle, samaroid, indehiscent, flattened at right an- gles to the plane of the partition (which is lacking), more or less reticulate; seeds soli- tary. A genus of about 30 Eurasian species, many in cultivation. /softs tinctoria L. Dyer's Woad. Plants erect, stems 3.5-10 dm tall or more, glabrous throughout or hirsute with long simple hairs at base; basal leaves 3.5-15 (18) cm long, 0.8-4 cm wide, oblanceolate to el- liptic, subentire to crenulate, ciliate to pi- lose with simple hairs; cauline leaves gradu- ally reduced upwards, lanceolate to elliptic, entire, hastately-auriculate, glabrous or pi- lose on veins beneath and often ciliate; pedicels 4.5-9 mm long, reflexed, glabrous; sepals 1.8-2.3 mm long, yellowish, glabrous; petals 3-4.2 mm long, yellow, spatulate, rounded; silicles mostly 10-18 mm long, 4-7 mm wide, cuneate-oblong to oblanceolate, more or less truncate-rounded apically, glabrous; stigmas sessile.— Roadsides, aban- doned fields, and dry foothills from 1350 to 1980 m elevation in Box Elder, Cache, Davis, Sanpete, Summit, Utah, and Weber counties; widely established in the United States; adventive from Europe. This plant is the source of a blue dye and was thus widely cultivated in the recent past. It seems to be spreading outwardly from its initial foci in Box Elder County, where it has been established as a weed since at least 1947. Lepidium L. Plants glabrous or with simple hairs, an- nual, biennial or perennial, from taproots; leaves alternate or basal and still alternate, simple and entire or variously toothed or bi- or tripinnatifid, petiolate or sessile, au- riculate in some; flowers racemose, pedicels lacking bracts; sepals 4, caducous or per- sistent; petals 4, yellow or white, in- frequently lacking; stamens 6, rarely 2 or 4, filaments lacking glandular processes; style obsolete or well developed, stigma capitate; fruit a silicle, usually less than twice longer than broad, compressed at right angles to septum, dehiscent, valves more or less reti- culately veined; seeds 1 per locule. (Note: Measurement of silicle length includes style.) A complex genus of perhaps 150 species found throughout the world, many as weeds. Hitchcock, C. L. 1936. The genus Lepidium in the United States. Madrono 3:265-320. Mulligan, G. A. 1961. The genus Lepidium in Canada. Madroiio 16:77-90. 1. Cauline leaves perfoliate-clasping or auriculate 2 Cauline leaves petiolate to sessile but not auriculate or clasping 3 2(1). Petals 2-2.5 mm long, white; cauline leaves lanceolate to lance-oblong, merely auriculate L. campestre Petals 1-2 mm long, yellow; cauline leaves oval to ovate, pseudoperfoliate .... L. perfoliatum 3(1). Plants arising from thickened, well-developed caudices; leaves all entire; stems seldom more than 2.5 dm tall; plants rare, south-central to north- ern Utah 4 Plants from simple or branched caudices, these seldom thickened; leaves en- tire, toothed or pinnatifid; stems short to long; plants widespread 5 4(3). Leaves linear; petals more than 3 mm long; plants of xeric shales, known only from Duchesne County L. barnebyanum Leaves oblanceolate to elliptic; petals less than 3 mm long; plants of moist meadows from Beaver County northward but not of Duchesne County 330 Great Basin Naturalist Vol. 37, No. 3 L. integrifolium 5(3). Styles 0.3-1 mm long; plants perennial or biennial 6 Styles obsolete or only up to 0.3 mm long; plants annual, rarely if ever biennial 7 6(5). Stems woody well above base; petals mostly 3 mm long or more; silicles 4.5- 7.5 mm long, obovate; plants of Washington County L. fremontii Stems slightly if at all woody above base; petals mostly less than 3 mm long (but sometimes longer); silicles 2-4 mm long, ovate; plants almost cosmo- pohtan in Utah L. montanum 7(5). Sepals persistent, enclosing mature silicle; silicles plainly reticulate; pedicels narrowly wing-margined; plants evidently rare L. strictwn Sepals deciduous at or shortly following anthesis; silicles plainly to slightly reticulate; pedicels terete to strongly flattened; plants at least locally common 8 8(7). Fruits emarginate and with prominent, acute, and divergent toothlike apices L. dictijotiim Fruits emarginate but lobes on each side of sinus neither acute nor diver- gent 9 9(8). Pedicels much flattened, about twice broader than thick 10 Pedicels not strongly flattened, not twice as broad as thick 11 10(9). Plants short-hirsute to hispid hairy; silicles pubescent L. lasiocarpum Plants softly hairy; silicles glabrous or pubescent L. densiflonim 11(9). Petals lacking, or, if present, generally shorter than sepals; silicles oblong- obovate to obovate L. densiflonim Petals present and usually surpassing sepals; silicles elliptic-rotund to orbicu- lar L. virginicum Lepidium barnebyanum Reveal. [L. mon- stems 1.5-6 dm tall, hirtellous throughout tanitm ssp. dernissum C. L. Hitchc, not L. with simple hairs; basal leaves 3-12 cm demissum C. L. Hitchc] Perennial, densely long, 0.8-1.5 cm wide, oblanceolate, entire pulvinate-cespitose, from a thickened or variously lobed; cauline leaves numerous, branched caudex, clothed with marcescent reduced upwards, becoming sessile and au- leaf bases; stems erect, 0.7-1.2 dm tall, sub- riculate, usually denticulate; pedicels mostly glabrous or minutely recurved scaberulous, 4-7 mm long, spreading, slightly flattened, simple or branched from the upper axils; hirtellous; sepals 1.3-2 mm long, greenish or basal leaves 1-4 cm long, 0.1-0.2 (0.3) cm variously tinged, hirtellous; petals 1.7-2.5 wide, somewhat flattened and minutely mm long, white, spatulate, ascending; sta- wing-margined, scaberulous; sepals 2-2.3 mens 6; silicles 5-6 mm long, oblong-ovate, mm long, greenish, glabrous or puberulent; glabrous or puberulent, concave, wingless, petals 3.5-4.2 mm long, white, obovate, slightly emarginate; styles 0.2-0.6 mm spreading; stamens 6; silicles 3-6.2 mm long, long.— Roadsides and disturbed sites from 3-4 mm wide, lanceolate to elliptic, 1300 to 2000 m elevation in Salt Lake, glabrous, plane, wingless; styles 0.5-1.2 mm Summit, and Utah counties, and probably long.— White (Green River) shale outcrops, widespread in the state; widely established Indian Canyon, Duchesne County; known in North America; Asia, only from the type locality; endemic. Lepidium densiflonim Schrad. Annual, Lepidium campestre (L.) R. Br. [Thlaspi lacking a caudex, stems 0.3-5 dm tall or campestre L.] Annual, lacking a caudex, more, densely finely hairy to puberulent Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 331 throughout; basal leaves 1.2-11 cm long, 0.3-2.2 cm wide, oblanceolate, entire and more commonly pinnately lobed; cauline leaves several to many, reduced upwards, petiolate to sessile, not auriculate, lobed, toothed or entire; pedicels 1.5-3 mm long, spreading to ascending, subterete to con- spicuously flattened, glabrous to puberulent; sepals 0.6-1.1 mm long, often purplish, glabrous; petals 0.7-1 mm long, white, nar- rowly oblong, sometimes lacking; stamens 2; silicles 2.5-3.5 mm long and about as wide, elliptic to oval or obovoid, glabrous or rarely pubescent, shallowly notched api- cally, teeth rounded; style lacking.— Wide- spread from low to moderate elevations in Utah; North America; Eurasia. Our material is separable into three more or less dis- tinctive variants. 1. Silicles pubescent, the pedicels usually somewhat flattened, especially on lower side; plants evidently rare L. densiflorum var. pubicarpum Silicles glabrous, pedicels definitely flattened or else subterete; plants com- mon 2 2(1). Pedicels subterete, silicles averaging less than 3 mm long; plants commonly of waste places L densiflorum var. densiflorum Pedicels definitely flattened, especially on upper side, silicles averaging at least 3 mm long; plants often growing in indigenous communities L. densiflorum var. ramosum Var. densiflorum [L. neglectum Thell.; L. hourgeauanwn Thell.; L. densiflorum var. bourgeauanum (Thell.) C. L. Hitchc] Weedy species usually of waste places, less commonly of indigenous plant communities, of Duchesne, Grand, Iron, Kane, Millard, Piute, Sevier, Uintah, Utah, and Washington counties, possibly cosmopolitan; range of species. This variety is abundant in Utah; we have not been able to distinguish the var. bourgeauanum from among this material. Var. pubicarpum (A. Nels.) Thell. [L. pubicarpum A. Nels.] Uncommon in dry sites in Salt Lake and Sanpete counties, and undoubtedly elsewhere; Washington to Montana and south to California. Var. ramosum (A. Nels.) Thell. [L. ramo- sum A. Nels.; L. densiflorum var. pubicaule Thell.] Common, usually in indigenous com- munities, in Emery, Garfield, Grand, Kane, San Juan, Sanpete, Uintah, and Utah coun- ties, and probably elsewhere; Wyoming south to New Mexico west to California. Lepidium dictyotum A. Gray. [L. dictyo- tum var. macrocarpum Thell.] Annual, lack- ing a caudex, stems 0.2-1.5 dm tall, pub- escent; basal leaves usually pinnatifid with 2-5 pairs of linear lobes; cauline leaves mostly entire, reduced upwards; pedicels 1.5-3.5 mm long, flattened, spreading to re- flexed; sepals 0.7-1 mm long, greenish, pub- escent; petals 1-1.2 mm long, white, oblong, or usually lacking; stamens typically 4; si- licles 3.2-4.5 mm long, 2-2.5 mm wide, ovate, glabrous or hirtellous, notched api- cally, teeth prolonged and usually divergent, acute; style lacking.— Evidently rare in Utah, reported from Salt Lake County by Hitchcock (1936, cited above). Lepidium fremontii S. Wats. Perennial subshrubs, definitely woody above base, stems mostly 3-8 dm tall, glabrous and glaucous; leaves cauline, mostly 1.5-5 (8) cm long, pinnatifid into linear-oblong, acute lobes up to 2 mm wide, becoming simple above, not auriculate; bracts subtending to curved-ascending, glabrous; sepals 1.6-2.1 mm long, green, glabrous; petals 3-4.5 mm long, white, obovate; stamens typically 6; silicles 4.5-7.5 mm long, 5.2-6.5 mm wide, obovate, glabrous, slightly notched apically; styles 0.4-1.3 mm long.— Blackbrush, creos- ote brush, and other warm desert shrub communities from 600 to 1525 m elevations in Washington County; Arizona, Nevada, and California. This species is obviously related to L. montanum (q.v.), but differs in being woody, having distinctive bracts in the in- 332 Great Basin Naturalist Vol. 37, No. 3 florescence, and in having larger, obovate silicles. Lepidium integrifolium Nutt. [L. iita- hense M. E. Jones; L. zionis A. Nels.; L. montanwn var. integrifolium (Nutt.) C. L. Hitchc] Perennial, cespitose, from a thick- ened, usually branched caudex, more or less clothed with marcescent leaf bases, stems decumbent-ascending, 1.5-2.5 dm tall, min- utely puberulent, simple or branched from upper axils; basal leaves 3-8.5 cm long, 0.6- 2.5 cm wide, elliptic to oblanceolate, entire; cauline leaves mostly 1-4 cm long, 0.2-1.2 cm wide, gradually reduced upwards, glabrous or nearly so; pedicels 5-8.5 mm long, spreading to ascending, puberulent; sepals 1.5-2 mm long, greenish, glabrous or pubescent; petals 2.7-3.1 mm long, white, obovate; stamens typically 6; silicles 3-4.2 mm long, 1.7-2.7 mm wide, ovate to lance- ovate, glabrous, plane, wingless; styles 0.4- 0.7 mm long.— Saline meadows at low to moderate elevations in Beaver, Rich, Sevier, and Uintah counties; Wyoming. This distinctive entity has been collected only rarely possibly because the habitat type has been exploited as marginal pas- tureland in Utah and Wyoming. Lepidium lasiocarpum Nutt. ex Torr. & Gray. Annual, lacking a caudex, stems 0.2-3 dm tall or more, hispidulous to distinctly hispid with simple hairs; basal leaves mostly 1-8.5 cm long, 0.3-0.9 cm wide, oblanceol- ate, pinnatifid or merely lobed or toothed; cauline leaves few to several, reduced up- wards, petiolate to sessile, not auriculate; pedicels 1.5-5 mm long, spreading to as- cending, conspicuously flattened, usually puberulent to hispidulous; sepals 1-1.2 mm long, greenish or reddish to purplish, glabrous; petals 1-1.3 mm long, white, nar- rowly oblong, sometimes lacking; stamens 2 or 4; silicles 3.4-4.5 mm long, 2.7-4 mm wide, elliptic to rotund or obovate, hispi- dulous, notched apically, teeth sometimes elongate-winged, rounded; style lacking or nearly so.— Widely distributed mostly in the southern two-thirds of Utah; California east- ward into New Mexico and Texas. Plants with somewhat attenuate apical teeth on the silicles approach L. dictijotum, and possibly they account for reports of that entity from Utah. Two scarcely distinguishable varieties are known from Utah: 1. Pedicels glabrous or merely puberulent on lower side; stems, at most, hispi- dulous L. lasiocarpum var. georginum Pedicels hispidulous on lower side; steins definitely hispid L. lasiocarpum var. lasiocarpum Var. georginum (Rydb.) C. L. Hitchc. [L. georginum Rydb.; L. lasiocarpum ssp. georgi- num (Rydb.) Thell.] Warm desert shrublands and lower pinyon-juniper woodlands in Grand, Kane, San Juan, Tooele, Uintah, and Washington counties; Arizona westward to California. Var. lasiocarpum [L. pahneri S. Wats.; L. lasiophyllum Brandegee; L. lasiocarpum ssp. pahneri (S. Wats.) Thell.; L. lasiocarpum ssp. lasiophyllum (Brandegee) Thell.] Warm to cool desert shrublands in Kane, Millard, Utah, and Washington counties; Arizona westward to California. Lepidium montanum Nutt. Perennial or less commonly biennial herbs or rarely suf- frutescent, seldom definitely woody above base, stems 0.3-12 dm tall or more, glabrous or variously puberulent to hirtellous; leaves basal or basal and cauline, 0.5-12.5 cm long, 0.1-2.5 cm wide, variously shaped, entire to pinnatifid, often at least uppermost cauline ones simple; bracts subtending branches of inflorescence seldom more than 1 cm long; pedicels 3-8 mm long, spreading to ascend- ing, glabrous or puberulent; sepals 1.2-1.7 mm long, green or variously tinged, glabrous or puberulent; petals 2.5—3.5 mm long, white, obovate to spatulate; stamens typically 6; silicles 2.8-4.1 mm long, 2.1-2.5 mm wide, ovate to elliptic, glabrous or rarely puberulent; styles 0.3-0.9 mm long.— Low to high elevations throughout Utah; Oregon to Wyoming southward to Califor- nia, Arizona, New Mexico, and Texas. There is an amazing amount of variation Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 333 within L. montanum as interpreted herein. all of these entities at the specific level, but The more stable phases have been treated the convenience of having an inclusive spe- previously either as species or as varieties, cies with several variants seems to best rep- There is some justification for recognizing resent the situation within Utah. 1. Plants mostly 6-12 dm tall; cauline leaves simple, toothed or entire, often more than 4 mm wide; flowering from midsummer to autumn; eastern to southeastern Utah L. montanum var. spathulatum Plants usually less than 6 dm tall; cauline leaves various, if simple and entire then usually less than 4 mm wide; flowering in springtime and midsummer (at higher elevations); distribution various 2 2(1). Basal leaves well developed, usually over 5 cm long and 0.6 cm wide; cauline leaves few, reduced upwards; plants usually montane 3 Basal leaves poorly, if at all, developed, usually less than 5 cm long and 0.6 cm wide; cauline leaves numerous to few, gradually reduced upwards; plants seldom montane 4 3(2). Stems puberulent almost or quite throughout L. montanum var. alpinum Stems glabrous almost or quite throughout L. montanum var. heterophyllum 4(2). Stems glabrous or essentially so (rarely puberulent); plants somewhat woody at the base 5 Stems puberulent almost or quite throughout; plants herbaceous or somewhat woody at the base 6 5(4). Plants (0.7) 1-6 (10) dm high; basal leaves divided, not entire; widespread in eastern, southeastern and southern Utah L. montanum var. jonesii Plants 0.2—0.5 (0.7) dm high; basal leaves entire or some divided; rare and local, Aquarius Plateau, Garfield County L. montanum var. neeseae 6(4). Plants perennial, the caudex woody; silicles puberulent; stems 0.5-1 dm tall; restricted to Kane County L. montanum var. stellae Plants biennial or perennial, the caudex lacking or only slightly woody; si- licles glabrous; stems often over 1 dm tall; widespread in western Utah L. montanum var. montanum Var. alpinum S. Wats. [L. scopulprum M. Garfield, Grand, Kane, San Juan, Uintah, E. Jones; L. heterophyllum M. E. Jones] Utah, Washington, and Wayne counties; Plants of rocky or sandy places at moderate Arizona and Nevada eastward to Colorado, to high elevations in Box Elder and Salt Var. montanum. [L. corymbosum Hook. Lake counties; endemic. & Arn.; L. utaviense Regal; L. brachyhot- Var. heterophyllum (S. Wats.) C. L. ryum Rydb.; L. philonitrum Nels. & Macbr.; Hitchc. [L. integrifolium var. heterophyllum L. albiflorum Nels. & Macbr.; L. montanum S. Wats.] Oak brush community. Cedar var. stenocarpum Thell.; L. scopulorum f. Canyon, Iron County. Endemic. nanum Thell.; L. montanum var. canescens Var. jonesii (Rydb.) C. L. Hitchc. [L. (Thell.) C. L. Hitchc] The phase distin- jonesii Rydb.; L. crandallii Rydb.; L. tortum guished as var. canescens by Hitchcock L. O. Williams; L. alyssioides var. jonesii passes into var. montanum through a series (Rydb.) Thell.; L. alyssioides var. stenocar- of morphological intermediates; thus it is pum Thell.] This is the abundant spring- placed here in synonymy with var. mon- flowering form of L. montanum of the Col- tanum. The var. montanum is a plant of orado drainage system. Mostly of middle to lower elevations in Box Elder, Duchesne, low elevations in Carbon, Duchesne, Emery, Iron, Juab, Millard, Piute, Sevier, Summit, 334 Great Basin Naturalist Vol. 37, No. 3 Tooele, Utah, and Washington counties; Idaho and Wyoming south to Arizona, Ne- vada, and California. Var. neeseae Welsh & Reveal, var. nov. Planta similis Lepidio montano var. stellae sed differt in caudicibus non ligneis, foliis basalibus et integris vel lobatis, et siliquis glabris. Holotype: Utah, Garfield Co., T.32S., R.3E., sec. 35, 14 mi due north of Escalante at Hell's Backbone, 2745 m, Neese & White 3332, 23 June 1977 (BRY, Isotypes to be distributed). Paratype: Gar- field Co., T.33S., R.3E., sec 35, 1 mile due southeast of Hell's Backbone, 2745 m, Neese & White 3944, 3 Aug. 1977 (BRY). The variety is named to honor Elizabeth Neese, ardent collector and student of the Henry Mountains flora. Var. spathulatum (B. L. Robinson) C. L. Hitchc. [L. scopiilonim var. spathulatum B. L. Robinson. L. spathulatum Vasey ex Rob- inson, not Philippi; Thehjpodium crenatum Greene; L. crenatum (Greene) Rydb.; L. vaseijanwn Thell.; L. montanum var. east- ivoodiae sensu Utah material; L. montanum var. ahjssioides sensu Utah material] This is the tall phase of the species which flowers in midsummer and autumn in southeastern Utah, in Garfield, Grand, Kane, San Juan, and Uintah counties; Colorado. This variant is sufficiently distinct from the body of the remaining varieties of L. montanum to be recognized at the specific level. If that is done, the correct name at the specific level would be L. crenatum. Var. stellae Welsh & Reveal, var. nov. Plantis similis Lepedio montano var. mon- tano sed differt in siliculis puberulentis et caudicibus bene evolutis. Type: Kane Coun- ty, Utah, along Cottonwood Wash Road, 6.4 miles south of Cannonville, on reddish clay soil near white barren shale outcrops south of the road, associated with Arte7nisia, Juni- perus, and Lesquerella tumulosa at about 1700 m elevation, 29 May 1976, Reveal & Reveal 4454. Holotypus, BRY. Isotyi, 15 to be distributed from US. Paratvpe: Welsh & Welsh 12841 (BRY). The variety is named to honor Stella L. T. Welsh, the cocoUector, with the senior author, of this plant in June of 1975. Lepidium perfoliatum L. Peppergrass. Plants annual, lacking a caudex, stems 0.7-4 (6) dm tall, hirtellous below, glabrous and glacous above; basal leaves 0.8-15 cm long, 0.2-2.8 cm wide, bi- or tripinnatifid into narrow segments; cauline leaves with lower- most like basal ones, these transitional up- wardly into entire and falsely perfoliate- clasping ones; pedicels 3-6.5 mm long, spreading-ascending, glabrous, subterete; sepals 0.7-1.2 mm long, often purplish, glabrous or pubescent; petals 0.8-1.5 mm long, yellow, narrowly spatulate, ascending; stamens usually 6; silicles 3.5-4.3 mm long, 3.2-3.6 mm wide, oval to elliptic, glabrous, plane, wingless or essentially so, slightly emarginate; style 0.1-0.3 mm long.— Dis- turbed sites and dry slopes, flats, and other places at low elevations in Box Elder, Cache, Iron, Juab, Kane, Salt Lake, San Juan, Sanpete, Tooele, Utah, Washington, and Weber counties, and probably cosmo- politan; widespread in the United States; adventive from Europe. Lepidium strictum (S. Wats.) Rattan. [L. oxycarpum var. strictum S. Wats.; L. reti- culatum Howell; L. puhescens of authors, not Desv.] Annual, lacking a caudex, stems 0.5-2 dm long, prostrate to rarely erect, pubescent; basal leaves mostly 3-7 cm long, 1-2 cm wide, bipinnatifid; cauline leaves less divided, uppermost sometimes entire; pedicels mostly 2-3 mm long, ascending, flattened, and narrowly wing-margined; sep- als 1-1.5 mm long, persistent, purplish, pub- escent; petals minute or lacking; stamens 2; silicles 2.2-3.5 mm long, 2-3 mm broad, oval to oblong-ovate, planely reticulate, slightly winged, concave, notched apically; style lacking.— Apparently rare in Utah and possibly not a part of the continuing flora; cited by Hitchcock (1936) from Castle Gate, Carbon County, collected by Grant in 1900. Lepidium virginicum L. Plants annual, lacking a caudex, stems 1.5-7 dm tall, pub- escent throughout or glabrous above; basal leaves 1.3-15 cm long, 0.3-3.5 (5) cm wide, coarsely toothed to lobed; cauline leaves re- duced upwards, uppermost usually entire; pedicels 2-6 mm long, spreading, terete or nearly so, pubescent or glabrous; sepals 0.6- 1 mm long; petals 0.7-1.5 (3) mm long, white, spatulate, rarely lacking; silicles 2.5-4 Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 335 mm long, 2.2-3.5 mm broad, elliptic to or- bicular, usually glabrous, plane, notched apically; style lacking.— Roadside, trails, foothills, stream sides, and similar disturbed places from low to moderate elevations in Salt Lake, Utah, Wasatch, Washington, and Weber counties, and likely elsewhere; wide- ly distributed in North America. Our material belongs to var. puhescens (Greene) Thell. [L. intennediinn var. pub- escens Greene; L. hirsutum Rydb.], the combination made by Thellung annodating that by Hitchcock by thirty years, and thus making the change of authorship necessary here. Because it seems probable that other varieties will be found in the state, the fol- lowing key is provided: 1. Silicles usually consistently longer than broad; stems usually hirtellous throughout; plants of eastern United States, adventive in our region L. virginicu7n var. virginicitm Silicles usually about as long as broad; stems glabrous above or minutely pu- berulent throughout; plants indigenous 2 2(1). Pedicels and raceme-rachis minutely puberulent L. virginicum var. puhescens Pedicels and raceme-rachis glabrous L. virginicum var. medium Var. m^edium (Greene) C. L. Hitchc. [L. 4, yellow, rounded; stamens 6, filaments intermedium A. Gray, not Richard; L. me- lacking glandular processes; style slender, dium Greene; L. glaucum Greene; L. ida- stigma capitate; fruit a sessile to substipitate hoense Heller] Reported from all states sur- silicle, less than twice longer than broad, rounding Utah on north and east, and to be varying in outline, compressed parallel or expected here. contrary to septum, valves veinless; seeds 2- Var. puhescens (Greene) Thell. Distribu- 10 per locule, biseriate. tion as given above. A genus of about 80 species found in Var. virginicum. Reported from scattered North and South America and just entering sites in the West and to be expected in Asia. Utah. Maguire, B., and A. H. Holmgren. 1951. Botany of the intermountain region— II. Lesquerella S. Wats. LesquereUa. Madrono 11:172-184. Plants stellate pubescent annual (biennial) Payson, E. B. 1921. A monograph of the or perennial, arising from taproots; leaves genus LesquereUa. Ann. Missouri Bot. basal and cauline, alternate, simple, taper- Gard. 8:1-3-236. ing to base or merely sessile, nto auriculate; Rollins, R. C., and E. A. Shaw. 1973. The flowers racemose, pedicels straight, sigmoid, genus LesquereUa in North America, reflexed, spreading or ascending, not sub- Cambridge, Mass.: Harvard University tended by bracts; sepals 4, deciduous; petals Press. 288 p. 1. Lowermost leaves narrow, mostly 1-5 mm wide, blade and petiole indistinct, or, if so, as in L. rectipes, plants tall and of low elevations, tufted at stem base; basal and cauline leaves alike in general shape Key I Lowermost leaves often over 5 mm wide, blade distinct from petiole, forming a rosette; basal leaves clearly of a different shape from cauline leaves Key II Key I. Basal and cauline leaves alike 1. Silicles and ovaries glabrous 2 Silicles and ovaries stellate pubescent 4 2(1). Stems mostly 0.6-1.8 dm tall; silicle bodies 4 mm long or more; plants of San Juan County L. fendleri 336 Great Basin Naturalist Vol. 37, No. 3 Stems 0.1-0.5 dm tall; silicle bodies usually less than 4 mm long; plants of south-central Utah 3 3(2). Plants densely pulvinate-cespitose, cushionlike, with usually numerous caudex branches; restricted to white shale outcrops southeast of Cannonville, Kane County L. tumulosa Plants cespitose in small compact to loose clumps, with one to several caudex branches; restricted to limestones on Paunsa gaunt Plateau L. rubricundula 4(1). Plants definitely pulvinate-cespitose, usually less than 1 dm tall; styles mostly 1-2 mm long; known from Washington and Kane counties L. arizonica Plants various, but if pulvinate-cespitose and less than 1 dm tall, then styles usually 2-4 mm long or more, or distribution otherwise 5 5(4). Plants usually less than 1 dm tall; basal leaves linear to narrowly spatulate; usually of middle to high elevations 6 Plants usually 1.5-2 dm tall or more; basal leaves spatulate to oblanceolate; usually of middle to lower elevations 7 6(5). Pedicels usually strongly S-shaped; inner basal leaves usually flat L. alpina Pedicels usually straight or only slightly curved; inner basal leaves usually in- volute L. intermedia 7(5). Pedicels generally recurved or arched in fruit, or less commonly almost straight; blades of basal leaves rarely more than 4 mm wide L. ludoviciana Pedicels more or less S-shaped; blades of basal leaves often more than 4 mm wide L. rectipes Key II. Basal and cauline leaves different 1. Plants slender annuals, stems erect or ascending, mostly 1-4 dm tall; restricted to Washington County L. tenella Plants cespitose perennials, stems decumbent to erect, mostly less than 1 (1.5) dm tall; distribution otherwise 2 2(1). Plants with caudex definitely branched; known from low to moderate eleva- tions of Duchesne, Emery, and Uintah counties L. siibumbellata Plants with caudex simple or rarely branched; distribution various, but usual ly of higher elevations 3 3(2). Silicles ellipsoid to subglobose or obovoid to obdeltoid, compressed contrary to septum 4 Silicles of various shape, but, if as above, compressed parallel to septum 5 4(3). Racemes loose to elongated in fruit, often secund; silicles sparingly pubescent with hairs 0.2-0.25 mm broad; plants of Wasatch and Uinta mountains L. utahensis Racemes short and dense in fruit, not at all secund; silicles moderately pu- bescent with hairs 0.2-0.35 mm broad; plants of Wasatch Plateau south- ward L. wardii 5(3). Silicles ovoid, compressed at apex or margins; plants known from northern Wasatch Mountains, from Mt. Nebo northward L. occidentalis Silicles various but not compressed at apex or margins; plants of various dis tribution 6 Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 337 6(5). Basal leaves angular, blades usually deltoid or hastate, narrowed abruptly to petiole; plants of Rich and possibly Summit counties L. prostrata Basal leaves not angular, blades elliptic to obovate, narrowed gradually to pe- tiole; plants of various distribution 7 7(6). Silicles either definitely compressed at apex or truncate to emarginate api- cally; plants from Utah County southward 8 Silicles more or less acute apically; plants from Utah County northward 9 8(7). Silicles obcordate to obdeltoid, sparingly pubescent; largest basal leaves usual- ly less than 1 cm wide; plants of Carbon, Emery, Sanpete, and Utah counties L. heyniphysaria Silicles ellipsoid, densely pubescent; largest basal leaves usually more than 1 cm wide; plants of Garfield, Kane, Sevier, and Tooele counties L. kingii 9(7). Pedicels loosely S-shaped; silicles sparsely pubescent with appressed or erect hairs; caudex branches lacking conspicuous leaf scars and bases; plants of Cache and Rich counties L. multiceps Pedicels straight or curved; silicles densely pubescent with erect hairs; caudex branches with conspicuous scars and/or leaf bases; plants of Davis, Salt Lake, Utah, and Wasatch counties L. garrettii Lesquerella alpina (Nutt.) S. Wats. Per- ennial, arising from simple or branched caudices, these often clothed with marces- cent leaf-bases; herbage pubescent with stel- late, 4-7-branched hairs; stems 0.1-1 dm tall, rarely more, erect, simple; basal leaves mostly 0.5-5 cm long, 0.1-0.4 cm wide, lin- ear to very narrowly spatulate, gradually narrowed to base; cauline leaves 0.5-2 cm long (or more), 0.1-0.3 cm wide; pedicels 4- 10 mm long or more, straight, curved, or S- shaped; sepals 3.5-6 (7) mm long; petals 4-8 mm long, yellow, spatulate; silicles (exclud- ing style) mostly 3-5 mm long, sessile or subsessile, ovoid, more or less compressed apically, valves pubescent externally and sometimes internally also; styles mostly 2-4 mm long; ovules 2-6 per locule.— Ridge tops and rocky slopes from 1830 to 3050 m elevation in central and northeastern Utah. Two varieties which lack definite diag- nostic criteria are represented among our materials. 1. Leaves spatulate, at least some; perhaps not distinct from the next L. alpina var. alpina Leaves uniformly narrow, linear to linear-spatulate L. alpina var. parvula Var. alpina. [Vesicaria alpina Nutt. ex Torr. & Gray; L. .spatidata Rydb.; L. ciir- vipes A. Nels.; L. alpina var. spatukita (Rydb.) Payson; L. condensata var. laevis Payson; L. alpina var. laevis (Payson) C. L. Hitchc] Ridgetops and alpine areas from 2200 to 3050 m elevation in Uintah and Sanpete counties; widespread northward to Alberta and Saskatchewan. Var. parvula (Greene) Welsh & Reveal, Stat, nov., based on Lesquerella parvula Greene, Pittonia 4:308. 1901. [L. alpina ssp. parvula (Greene) Rollins & Shaw] Ridgetops and alpine to subalpine areas from 1830 to 2600 m elevation in Duchesne and Daggett counties; Colorado and Wyoming. It seems probable that all the Utah mate- rial designated as L. alpina might belong only to var. parvula. The var. condensata (A. Nels.) C. L. Hitchc. is to be sought in extreme northern Daggett County; it is cur- rently known from just north of the Utah line in Uinta County, Wyoming. The var. condensata is a small, tufted plant with the stems barely exserted beyond the leaves. Lesquerella arizonica S. Wats. [L. arizo- nica var. nudicaulis Payson] Perennial, more or less pulvinate-cespitose, with few 338 Great Basin Naturalist Vol. 37, No. 3 to several (or many) caudex branches, these often with marcescent leaf-bases; herbage densely pubescent with stellate hairs; stems 0.2-1 dm tall, rarely more, erect, simple; basal leaves 0.5-3.5 cm long, 0.1-0.4 cm wide, oblanceolate to spatulate, gradually narrowed to base; cauline leaves 0.5-2.5 cm long or more, 0.1-0.3 cm wide; pedicels 3- 10 mm long or more, straight or curved, as- cending; sepals 4-6.5 mm long; petals 5.5-7 mm long, yellow, suborbicular; silicles (ex- cluding style) 3-5 mm long, sessile or sub- stipitate, ovoid to ellipsoid, rounded and compressed apically, valves pubescent exter- nally; styles 0.5-2 (4) mm long; ovules 2-5 (8) per locule.— Mountain brush, sagebrush, and pinyon-juniper communities from 1280 to 2750 m elevation in Kane and Washing- ton counties; Arizona. Lesquerella fendleri (A. Gray) S. Wats. [Vesicaria fendleri A. Gray; V. stenophyUa A. Gray; L. foliacea Greene; L. stenophyUa (A. Gray) Rydb.; L. praecox Woot. & Standi.] Perennial, cespitose, caudex simple or few- to several-branched, branches naked or with few marcescent leaf-bases; herbage usually densely pubescent with stellate hairs; stems 0.5-2.5 dm tall, rarely more, erect to decumbent, simple; basal leaves 1-4 cm long, 0.1-0.6 cm wide, elliptic to oblan- ceolate, gradually tapering to the base; cauline leaves 0.4-2 (2.5) cm long, mostly 0.1-0.5 mm wide; pedicels 7-15 (20) mm long, straight or curved to S-shaped, spread- ing to suberect; sepals 4.5-8 mm long; pet- als 6-12 mm long, yellow, obovate; silicles (excluding style) 4.5-8 mm long, sessile or subsessile, ellipsoid to ovoid, rounded api- cally, the valves glabrous; styles 2-6 mm long; ovules 6-16 per locule.— Warm desert, salt desert, and lower juniper-pinyon com- munities in San Juan County; Colorado and Arizona eastward to Texas and south to Mexico. Lesquerella garrettii Payson. Perennial, cespitose, caudex simple or few- to several- branched, branches clothed with marcescent leaf-bases and scars; herbage pubescent with stellate hairs; stems 0.5-1.5 dm tall, decum- bent-spreading to suberect, simple; basal leaves 1-3.5 (4) cm long, 0.2-0.7 cm wide, elliptic to obovate, differentiated into blade and petiole; cauline leaves 0.3-1.3 cm long, 0.1-0.4 cm wide; pedicels 3.5-7 mm long, straight or curved, spreading-ascending; sep- als 4.5-6 mm long; petals 6-9 mm long, yel- low, spatulate to obovate; silicles (excluding style) 3.5-4 mm long, short-stipitate, sub- globose or obovoid, valves pubescent exter- nally, glabrous within; styles 4-7 mm long; ovules 2-4 per locule.— Rocky sites at high elevations, from 2900 to 3350 m elevation in Salt Lake, Utah, and Wasatch counties; endemic. The type was collected in Big Cottonwood Canyon, Salt Lake County. Lesquerella hemiphysaria Maguire. Per- ennial, cespitose, caudex simple or few- branched, branches more or less clothed with marcescent leaf-bases; herbage pub- escent with stellate hairs; stems 0.3-1 (1.5) dm tall, decumbent-spreading or sometimes rarely erect; basal leaves 0.5-3.5 (5.5) cm long, 0.4-1 (1.5) cm wide, obovate to ellip- tic, differentiated into blade and petiole; cauline leaves 0.3-1.5 cm long, 0.2-0.5 cm wide; pedicels 3-7 mm long, spreading, as- cending, or recurved, S-shaped or curved; sepals 4-7 mm long; petals 5-7 mm long, yellow, oblanceolate; silicles (excluding style) 3-7 mm long, sessile or substipitate, obcordate in outline, valves more or less pubescent externally, glabrous within; styles 3-6 mm long; seeds 4-8 per locule. Silicles glabrous or nearly so; plants of the West Tavaputs Plateau L. hcmiphi/saria var. hicens Silicles uniformly pubescent throughout; plants of the Wasatch Plateau L. hemiphysaria var. hemiphysaria Var. hemiphysaria. Meadows and ridge- Var. lucens Welsh & Reveal var. nov. O tops from 2600 to 3200 m elevation in Car- Lesquerella hemiphysaria var. hemiphysaria bon, Emery, Sanpete and Sevier counties; differt in sili(}uis glabris vel fere glabris. endemic. Holotype: Utah, Carbon Co., 7 miles due Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 339 northeast of Sunnyside, Range Creek Can- yon, S. Welsh & K. Taylor 15139, 23 June 1977 (BRY, iso types to be distributed). Para- type: Utah, Carbon Co., 6 miles north- northeast of Sunnyside, S. Welsh & K. Tay- lor 15122, 22 June 1977 (BRY). Sagebrush and woodland communities at 2700 to 2800 m on the West Tavaputs Plateau, Carbon Co.; endemic. Lesquerella intermedia (S. Wats.) Heller. [L. alpina var. intermedia S. Wats.] Per- ennial, cespitose, caudex simple or few- branched, branches more or less clothed with marcescent leaf-bases; herbage pub- escent with stellate hairs; stems 0.2-1.5 (2.5) dm tall, erect or ascending, simple; basal leaves 1-5 cm long, 0.1-0.2 cm wide, hnear or only slightly expanded apically, inner usually invokite, tapering gradually to base; cauhne leaves 0.8-3.5 cm long, 0.1-0.3 cm wide; pedicels 4-12 mm long, spreading to ascending, straight or curved; sepals 2.8-6.5 mm long; petals 5.5-10.5 mm long, yellow, spatulate; silicles (excluding style) 4-7 mm long, sessile or substipitate, ellipsoid or ovoid, acute and somewhat flattened api- cally, pubescent externally, glabrous or pub- escent within; styles (2) 3-4.5 mm long; seeds (4) 6-8 (10) per locule.— Aspen, mixed conifer, ponderosa pine, and pinyon-juniper communities, from 1525 to 2900 m eleva- tion in Beaver, Emery, Garfield, Sevier, Washington, and Wayne counties; Arizona, New Mexico. This taxon approaches L. alpina on the one hand and L. rectipes on the other. De- finitive features to separate alpine phases of L. intermedia from L. alpina are subject to interpretation. Only arbitrary separation seems possible. Lesquerella kingii S. Wats. Perennial, cespitose, caudex usually simple, densely clothed with marcescent leaf-bases; herbage pubescent with stellate hairs; stems 0.3-1.8 dm tall, decumbent to suberect, simple; bas- al leaves 1.5-4.5 (6) cm long, 0.4-2 cm wide, blade spatulate to oval, obovate or ovate, sharply differentiated from petiole; cauline leaves 0.5-2 cm long, 0.1-0.7 cm wide; pedi- cels 4-10 mm long or more, curved to straight or S-shaped, ascending to descend- ing; sepals 4-8 mm long; petals 5.5-12 mm long, yellow, spatulate; siliques (excluding style) 3.5-9 mm long, sessile to substipitate, ellipsoid, compressed apically, valves pub- escent externally, pubescent to glabrous within; styles 2-5 mm long; seeds 4-8 per locule.— Ponderosa pine and juniper-pinyon communities at low to moderate elevations in Garfield, Millard, Sevier, Tooele, and Washington counties; Nevada, California and Oregon. Our material, which is not always dis- tinguishable from L. wardii, belongs to var. parvifolia (Maguire & Holmgren) Welsh and Reveal, comb, nov., based on Les- querella occidentalis var. parvifolia Maguire & Holmgren, Madrofio 11: 179. 1951 [L. latifolia A. Nels.; L. barnebyi Maguire; L. kingii ssp. latifolia (A. Nels.) Rollins & Shaw]. The var. cordiformis (Rollins) Ma- guire & Holmgren is to be sought in the deserts of extreme western Utah. Lesquerella ludoviciana (Nutt.) S. Wats. [Myagrum argenteum Pursh; Ahjssum ludo- vicianum Nutt.; L. argentea (Pursh) Mac- Millan, not L. argentea (Schauer) S. Wats.] Perennial, loosely cespitose, caudex simple to many-branched, branches clothed with marcescent leaf-bases; herbage pubescent with stellate hairs; stems 0.7-3 (4) dm tall, ascending to erect or outer decumbent, simple; basal leaves 1-9 cm long, (0.1) 0.2- 0.8 cm wide, spatulate to oblanceolate or appearing linear when folded; cauline leaves 1-6 cm long, 0.1-0.6 cm wide; pedi- cels 5-15 mm long or more, straight or curved, ascending to recurved; sepals 4-7 mm long; petals 5-10 mm long, yellow, ob- lanceolate to obovate; silicles (excluding style) 3-6 mm long, sessile or nearly so, sub- globose or obovoid, valves pubescent exter- nally and usually pubescent within; styles 3- 5 mm long; seeds 2-8 per locule.— Pinyon- juniper, sagebrush, and upwards to spruce- fir woodlands, from 1370 to 2750 m eleva- tion in Daggett, Duchesne, Garfield, Kane, and Uintah counties; Minnesota and Illinois westward to Montana, and southward to Nevada, Colorado, and Kansas. Lesquerella multiceps Maguire. Perennial, cespitose, caudex simple or several- to many-branched, more or less clothed with marcescent leaf-bases; herbage pubescent 340 Great Basin Naturalist Vol. 37, No. 3 with stellate hairs; stems 0.3-2 dm long, rarely more, prostrate to erect; basal leaves 1-6 cm long, 0.4-1.5 (2) cm wide, blades el- liptic to obovate, differentiated from long slender petioles; cauline leaves 0.4-2 cm long, 0.1-0.5 cm wide; pedicels 3-10 mm long, S-shaped; sepals 4.3-7 mm long; petals 5-10 mm long, yellow, spatulate; silicles (ex- cluding style) 3-5.5 mm long, sessile or nearly so, ovoid, valves pubescent exter- nally, glabrous within; styles 3-6.5 rm long; ovules 2-4 per locule.— Ridges and upper slopes from 1830 to 3050 m elevation in Cache, Davis, and Rich counties; Idaho, Wyoming. Lesquerella occidentalis S. Wats. Per- ennial, cespitose, caudex simple or few- branched, more or less clothed with marces- cent leaf-bases; herbage pubescent with stel- late hairs; stems 0.2-1.5 dm long, rarely more, prostrate, decumbent, ascending or erect, usually simple; basal leaves 0.5-8 cm long, 0.3-1.5 cm wide, blades spatulate to oval or obovate, differentiated from pe- tioles; cauline leaves 0.4-1.5 cm long, 0.1- 0.8 cm wide; pedicels 3-10 mm long or more, sigmoid or straight, ascending; sepals 4-7 mm long; petals 6-10 mm long, yellow, spatulate; silicles (excluding style) 4-6 mm long, ellipsoid, usually compressed at apex, valves pubescent externally and usually pub- escent within; styles 2-5 mm long; seeds 2-8 per locule.— Limestone cliffs and talus slopes from 1525 to 3660 m elevation in Box Elder and Juab (Mt. Nebo) counties; Idaho and Utah to California and Oregon. Our material belongs to var. cinerascens Maguire and Holmgren [L. occidentalis ssp. cinerascens (Maguire & Holmgren) Rollins & Shaw]. Lesquerella prostrata A. Nels. Perennial, cespitose, caudex simple or few-branched, branches more or less clothed with marces- cent leaf-bases; herbage pubescent with stel- late hairs; stems 0.3-1.5 dm tall, decumbent to ascending, simple; basal leaves 1-5 cm long, 0.5-1 (1.5) cm wide, blades deltate to hastate, more or less angular, differentiated from slender petioles; cauline leaves 0.4-1.5 cm long, 0.1-0.5 cm wide; pedicels 4-10 mm long, curved to straight or S-shaped, spreading; sepals 5-6 mm long; petals 5-9 mm long, yellow, spatulate; silicles (exclud- ing style) 4-7 mm long, sessile or nearly so, ovoid to ellipsoid, not flattened apically, valves pubescent externally, glabrous or sparsely hairy within; styles 3-6 mm long; seeds 2-4 per locule. —Sagebrush commu- nity, especially on dry shaly slopes from 1830 to 2440 m elevation in Rich (and to be sought in Summit) County; Wyoming and Idaho. Lesquerella rectipes Woot. & Standi. Pe- rennial, loosely cespitose, caudex simple or few-branched, branches more or less clothed by marcescent leaf-bases; stems 0.4-4 dm tall or more, decumbent to ascending or erect, simple or branched; basal leaves 1-8 (10) cm long, 0.3-1.2 cm wide, oblanceolate to elliptic, entire or toothed, tapering grad- ually to base; cauline leaves 0.6-3 cm long, 0.2-1 cm wide; pedicels 4-10 mm long or more, straight or S-shaped, spreading to as- cending or recurved; sepals 4-8 mm long; petals 6-10 mm long or more, yellow, spatulate to obovate; silicles (excluding style) 4-7 mm long, substipitate to sub- sessile, ovoid to ellipsoid, pubescent exter- nally and glabrous or pubescent within; styles 2-7 mm long; seeds 4-8 per locule.— Pinyon-juniper, grasslands, warm desert shrub, and ponderosa pine communities from 1220 to 2440 m elevation in Carbon, Emery, Garfield, Grand, Kane, San Juan, Uintah, Washington and Wayne coimties; Utah and Arizona eastward to Colorado and New Mexico. This is the most common of the tall spe- cies of Lesquerella in southeastern Utah. Lesquerella rubricundula Rollins. [L. hitchcockii ssp. rtibiciindula (Rollins) Ma- guire & Holmgren] Plants perennial, loosely cespitose, not matted, caudex simple or with few to several branches, branches clothed with marcescent leaf bases; herbage pubescent with stellate hairs; stems 0.1-0.5 dm tall, erect, simple; leaves mainly basal, 0.3-1.2 cm long, 0.1-0.2 cm wide, not differ- entiated into blade and petiole; pedicels 1-6 mm long, straight or curved, ascending; se- pals 3.2-4.5 mm long; petals 4.5-7 mm long, yellow, spatulate; silicles (excluding style) 3- 5 mm long, sessile to substipitate, ovoid, valves glabrous externally and within; styles Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 341 2.8-3.5 mm long; ovules 2-4 per locule.— Mixed conifer woodlands, on White and Pink limestone member of the Wasatch Formation from 2440 to 3050 m elevation in Garfield (type locality at Red Canyon) and Iron coimties; endemic. Lesquerella subumbellata Rollins. Pe- rennial, cespitose, caudex several-branched, branches clothed with marcescent leaf bases and often with leaves as well; herbage pu- bescent with stellate hairs; stems 0.1-0.8 (1) dm tall, ascending to erect, simple; basal leaves 0.8-3 cm long, 0.2-0.6 (1) cm wide, at least outer and usually most of them obo- vate-spatulate, gradually tapering to petiole; cauline leaves 0.3-2 cm long, 0.1-0.4 cm wide; pedicels 2-5 mm long, straight or curved, ascending; sepals 3-4.5 mm long; petals 5-6 mm long, yellow, spatulate; silicles (excluding style) 3.5-5 mm long, sub- sessile, ellipsoid, valves pubescent exter- nally; styles 1.5-2.5 mm long; seeds 2-6 per locule.— Pinyon-juniper and sagebrush com- munities from 1680 to 1980 m elevation in Duchesne and Uintah (type locaity 18 miles north of Vernal) counties; Colorado, Wyo- ming. Rollins and Shaw (1973) place this taxon within the range of variation of Lesquerella alpina. That disposition ignores the basic continuity of L. subumbellata, and likewise denies the apparent relationship of L. al- pina in quite another direction, i.e., with L. intermedia. The uniformly broad blades of basal leaves, although tenuous as a diagnos- tic feature, are on the order of other diag- nostic features in this critical genus. Lesquerella tenella A. Nels. [L. gordonii var. sessilis S. Wats.; L. palmeri of Utah ref- erences, not S. Wats.] Annual, herbage pu- bescent with stellate hairs; stems 1-5 dm tall or more, spreading-decumbent to erect, simple or branched; basal leaves 1.5-6 cm long, 0.2-1.5 cm wide, blades elliptic, some- times toothed, differentiated from petiole; cauline leaves 0.4-4 cm long, 0.2-1 cm wide; pedicels 4-12 (18) mm long or more, S-shaped, spreading to recurved; sepals 3.5- 7 mm long; petals 5-10 mm long, yellow to orange, orbicular to obovate; silicles (ex- cluding style) 3.5-5 mm long, sessile to sub- stipitate, globose to obovoid, valves pu- bescent externally and internally; styles 2- 4.5 mm long; ovules 2-6 per locule.— Black- brush, cresote brush, and Joshua tree com- munities at lower elevations in Washington County (the type of var. sessilis is from St. George); Nevada and California south to Mexico. Lesquerella tumulosa (Bameby) Reveal. [L. hitchcockii ssp. tumulosa Barneby] Pe- rennial, pulvinate-cespitose, densely matted, caudex many-branched, branches clothed with numerous marcescent leaves and leaf bases; herbage pubescent with stellate hairs; stems 0.1-0.4 dm tall, erect, simple; leaves mainly basal, 0.2-1 (1.5) cm long, (0.07) 0.1- 0.15 (0.2) cm wide, not differentiated into blade and petiole; pedicels 2-5 mm long, straight or S-shaped, spreading to ascending; sepals 2.8-4 mm long; petals 5-7 mm long, yellow, spatulate; silicles (excluding style) 2.7-3.8 mm long, substipitate, ovoid, valves glabrous externally and internally; styles 1.9-2.7 mm long; ovules 2-4 per locule.— White, bare shale knolls (Winsor Member of the Carmel Formation) among scattered juniper in a Bouteloua grassland, Kane Coimty; endemic. This plant was included with L. rubicun- dula by Rollins and Shaw (1973), but is dis- tinct morphologically, spatially, and sub- strate-wise from that taxon as noted by Reveal (Great Basin Nat. 30:94-98. 1970). Lesquerella utahensis Rydb. Perennial, cespitose, caudex usually simple, more or less clothed with persistent leaf-bases; her- bage pubescent with stellate hairs; stems 0.2-1.5 dm tall or more, decumbent to as- cending or erect; basal leaves 1.2-6.5 cm long, 0.1-0.5 cm wide; pedicels 3-10 mm long, sigmoid or curved, ascending; sepals 4.5-5.7 (7) mm long; petals 6.5-9 mm long, yellow, spatulate; siliques (excluding style) 3.5-4.5 (6) mm long, substipitate to sessile, globose-ellipsoid, compressed contrary to partition, valves pubescent externally, glabrous internally; styles 4-5.5 (6.5) mm long; seeds 3-6 per locule.— Openings among spruce-fir woodlands from 2600 to 3350 m elevation in Duchesne, Salt Lake, Summit, Uintah, Utah (the type is from American Fork Canyon), and Wasatch counties; en- demic. 342 Great Basin Naturalist Lesquerella wardii S. Wats. Perennial, caudex usually simple, clothed with marces- cent leaf-bases; herbage pubescent with stel- late hairs; stems 0.2-2 dm tall, rarely more, decumbent to ascending or erect, simple; basal leaves 1-4.5 (6) cm long, 0.3-1.5 (2) cm wide, blades deltoid to orbicular or el- liptic, differentiated from slender petiole; pedicels 2-7 mm long, straight, curved, or S-shaped, ascending; sepals 4-6.3 (8) mm long; petals (5.5) 6-9 (11) mm long, yellow, spatulate; silicles (3.5) 4-6.8 (8) mm long, substipitate to sessile, ellipsoid or ovoid, not compressed apically, pubescent externally and usually so internally; styles (1) 2.2-4 (7) mm long; seeds 2-8 per locule.— Spruce-fir woodlands, meadows, pinyon-juniper, and desert shrub commimities from 1830 to 3400 m elevation in Beaver, Garfield (type area is Aquarius Plateau), Iron, Kane, Mil- lard, Piute, Sevier, and Washington coun- ties. Lesquerella wardii is most similar to the partially sympatric L. kingii var. parvifolia from which it is distinguished inter alia in the silicles being rounded, not compressed, apically. Lobularia Desv. Nom. Cons. Plants annual or rarely biennial, pu- bescent with malpighian hairs, arising from taproots; leaves alternate, simple, entire, petiolate or sessile, not auriculate; flowers racemose, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, white, pink, or lavender; stamens 6, filaments lacking glandular processes; style short, stigmas capitate; fruit a silicle, about as broad as long, compressed parallel to septum, dehis- cent, valves 1-nerved; seeds 1 per locule. A European genus of 5 species. Lobularia maritima (L.) Desv. [Chjpeola maritima L.; Alys.sum maritimum (L.) Lam.] Sweet Alyssum. Annual, stems spreading-de- cumbent or ascending to erect, usually much branched, 0.8-3 dm long or more; leaves linear-oblanceolate, 0.8-3 (4) cm long, 0.1-0.4 cm wide, strigose on both surfaces; pedicels spreading-ascending, 5-9 mm long; sepals 1-1.5 (2) mm long, green or purplish, pubescent; petals 2.5-4 mm long, white. Vol. 37, No. 3 pink or lavender, the blades obovate; silicles 2.5-3.5 mm long and about as broad, valves sparingly strigose; styles 0.3-0.6 mm long.— Ornamental border plant with sweetly scented flowers, escaping and occasionally persistent; introduced from Europe. LUNARIA L. Plants annual or biennial, pubescent with simple or with some forked hairs, arising from taproots; leaves usually alternate and basal, simple, dentate to lobed, petiolate, not auriculate; flowers racemose, pedicels not subtended by bracts; sepals 4, de- ciduous; petals 4, pink to lilac or blue or purplish; stamens 6, filaments lacking glandular processes; style elongate, per- sistent, stigma capitate, bilobed; fruit a stipitate silicle, about as broad as long, compressed parallel to septum, dehiscent, valves reticulately veined; seeds 3-5 per lo- cule. A small Eurasian genus of 2 or 3 species. Lunaria annua L. Honesty; Moonwort; Satin-flower. Plants with one to several stems, these simple or branched, mostly 4- 10 dm tall or more, pubescent with simple, and rarely with some forked, hairs; leaves 2.5-10 (15) cm long, 1.5-8 cm wide, rarely more, ovate to lance-ovate or cordate, den- tate to lobed, pubescent with simple or rarely forked hairs; pedicels 10-22 mm long, spreading-ascending, pubescent; sepals 6-8 mm long, greenish or variously suffused, sparsely pubescent; petals 14-20 mm long, pink to lilac or blue to purplish, obovate; silicles 32-45 mm long, 25-35 mm wide, borne on stipes 7-12 mm long, much flat- tened, valves glabrous; styles 6-8 mm long. —Ornamental, cultivated for the showy flowers and distinctive fruiting in- florescences, occasionally escaping; in- troduced from Europe. Malcomia (L.) R. Br. Nom. Cons. Plants pubescent with forked or 3-rayed hairs, annual, from taproots; leaves alternate and basal, simple, sinuate-dentate, cauline ones petiolate to sessile, not auriculate; flowers racemose, pedicels seldom sub- tended by bracts; sepals 4, tardily de- Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 343 ciduous; petals 4, pink to lavender; stamens 6, filaments lacking glandular processes; style tapering, stigma oblique; fruit a si- lique, many times longer than broad, sub- terete, dehiscent, valves nerveless; seeds uniseriate. About 30 species of Europe and Africa, with 2 or 3 species in North America as weeds. Malcolmia africana R. Br. in Ait. Plants decumbent-ascending to erect, stems 0.3-4 dm tall, simple or branched almost through- out, pubescent with forked or 3-rayed hairs; leaves mainly basal to mainly cauline, 1.2-9 cm long, 0.3-2.3 cm wide, oblanceolate to elliptic, sinuate-dentate, petiolate to sessile, not auriculate; pedicels spreading, 1-2 mm long, pubescent; sepals 3.7-5.2 mm long, of- ten reddish or purplish, pubescent; petals 6.2-9.5 mm long, pink to lavender, rounded apically, spreading; siliques 33-66 mm long, straight, pubescent; style up to 1 mm long, stigma oblique.— Roadsides, foothills, and other disturbed sites at lower elevations in Cache, Carbon, Emery, Garfield, Grand, Juab, Kane, Millard, Salt Lake, San Juan, Sevier, Tooele, Uintah, Utah, and Washing- ton counties, and probably throughout the state; widespread throughout the Great Ba- sin and adjacent areas of the western United States; adventive from Africa. Nasturtium R. Br. Plants glabrous perennials from sub- rhizomatous stolons; leaves alternate, simple or some pinnately compound, petiolate and auriculate; flowers racemose, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, white; stamens 6, filaments lacking glandular processes; styles stout, well devel- oped, stigma capitate, bilobed; fruit a si- lique, several times longer than broad, ob- long, somewhat compressed parallel to septum, the valves 1 -nerved; seeds several to many, biseriate. Perhaps 50 species of moist places in the temperate zones. Nasturtium officinale R. Br. in Ait. [Si- symbrium nasturtium-aquaticum L.; Ro- rippa nasturtium-aquaticum (L.) Schinz & Thell.] Watercress. Plants submersed or emergent, succulent stems (0.3) 1-8 (10) dm long or more, glabrous; leaves 1-10 cm long, terminal lobe usually largest, falsely petiolate and narrowly auriculate-clasping basally; pedicels 5-13 (20) mm long, spread- ing to spreading-ascending, glabrous or pu- bescent; sepals 2-3 mm long, green or tips white, glabrous; petals 3-4.7 mm long, white, rarely with veins purplish, oblanceo- late; siliques 10-18 (25) mm long, 1.8-2.4 mm wide; styles 0.7-1.2 mm long.— Seeps, springs, and sluggish streams, usually in flowing water at moderte elevations in Cache, Garfield, Grand, Iron, Millard, Salt Lake, San Juan, Summit, Utah, Wasatch, Washington, Wayne, and Weber counties, and probably throughout the state; widely established in North America; introduced from Europe. Watercress is used as a salad plant. Parrya R. Br. Plants perennial, glabrous, pubescent with stipitate-glandular hairs, arising from tap- roots and with simple to branched caudices; leaves mainly basal, simple, dentate, taper- ing basally, not auriculate; flowers race- mose, pedicels not subtended by bracts; sep- als 4, deciduous; petals 4, pink to lavender; stamens 6, filaments lacking glandular pro- cesses; styles stout, persistent, stigmas bi- lobed; fruit a silique, usually several times longer than broad, oblong, constricted be- tween seeds, strongly compressed parallel to septum, valves 1-nerved; seeds 1 to several per locule. An Asian and North American genus of perhaps 25 species. Parrya rydbergii Botsch. [P. platycarpa Rydb., not Hook. f. & Thomas] Plants rosu- late, low perennials, caudex clothed with marcescent leaf bases; stems scapose, 0.7-1.2 dm tall, 2-8 cm to base of raceme, herbage stipitate-glandular; leaves 3-10 cm long, 0.6- 2 cm wide, oblanceolate to elliptic; flowers racemose, 3-7 (10); pedicels 4-20 mm long, stout, steeply ascending; sepals 7.2-9.3 mm long, purplish, stipitate-glandular or glabrous; petals 16-23 mm long, pink to lav- ender, cuneate-spatulate, emarginate; si- liques 25-47 mm long, 3-3.5 mm wide, straight or curved, midnerve prominent, glabrous or stipitate-glandular; style 0.3-0.6 344 Great Basin Naturalist Vol. 37, No. 3 mm long, stigma deeply bilobed; seeds 1-4 per locule.— Talus slopes, krumholz commu- nities, and alpina tundra sites in the Uinta Mountains from 3200 to 3720 m elevation in Daggett, Duchesne, Summit, and Uintah counties; endemic. This is one of our most unique and at- tractive Utah endemic species, and one that may be worthy of cultivation if low-eleva- tion cultivars could be developed. Physaria (Nutt.) A. Gray Plants stellate pubescent perennials, aris- ing from taproots; leaves mainly basal, caul- ine ones much reduced, alternate, simple, tapering basally or merely sessile, not au- riculate; flowers racemose, pedicels straight, curved, sigmoid, spreading or descending, not subtended by bracts; sepals 4, de- ciduous; petals 4, yellow, rarely purple, rounded; stamens 6, filaments lacking glandular processes; style slender, stigma capitate; fruit a sessile, bladdery-inflated si- licic, often broader than long, varying in outline, compressed (if at all) contrary to septum, valves veinless; seeds 2-6 per lo- cule, biseriate. A genus of perhaps 15 species restricted to western North America. The diagnostic features used to segregate entities of this genus are mainly based on the shape of the mature silicic and the length of the style. These features are sub- ject to variation within rather broad limits, and not all specimens will fit neatly into the following key. Immature specimens are particularly difficult to place, and strictly fruiting collections are often more readily definable than flowering collections. Mulligan, G. A. 1967. Cytotaxonomy of Physaria acutifolia, P. chambersii, and P. newberryi (Cruciferae). Canad. J. Bot. 45:1887-1898. Payson, E. B. 1918. Notes on certain Cruci- ferae. Physaria. Ann. Missouri Bot. Gard 5:143-147. Rollins, R. C. 1939. The cruciferous genus Physaria. Rhodora 41:392-415. Waite, S. B. 1973. A taxonomic revision of Physaria (Cruciferae) in Utah. Great Ba- sin Nat. 33:31-36. 1. Styles 2-3 (4) mm long; valves of silicles sharply angled at maturity; plants of Grand, Kane, San Juan, and Washington counties P. newberryi Styles mostly 4-8 mm long or more; valves of silicles obtusely angled to rounded at maturity; distribution various 2 2(1). Basal leaves sinuate-dentate to lobed, sometimes with some entire; cauline leaves often toothed; plants endemic to Uintah Basin P. grahamii Basal leaves entire or rarely sinuate-dentate; cauline leaves entire; plants of broad distribution 3 3(2). Sinuses of silicles indented above and below, when immature equally rounded to apex and base; valves rounded, not angled at maturity; plants mainly of the Colorado River drainage system P. acutifolia Sinuses of silicles only slightly indented to rounded below, deeply indented above, when immature tapering to base and obcordate in outline; valves obtusely angled at maturity; plants mainly of Great Basin and valleys of Virgin River drainage system P. chambersii Physaria acutifolia Rydb. Plants pe- rennial, cespitose, caudex usually simple, clothed with marcescent leaf bases; stems 0.3-2 (2.5) dm long, decumbent to ascending or erect, simple; basal leaves 1.7-9.5 cm long, 0.7-3 cm wide, blade orbicular to ovate or obovate, less commonly lance-ellip- tic, entire, angular, or toothed; cauline leaves greatly reduced; pedicels mostly 7-16 mm long, ascending to descending; sepals 5.2-8.7 mm long; petals 7.5-12.5 mm long, yellow or purple, spatulate; silicles blad- dery-inflated, sinuses indented above and somewhat less so below, valves 7-15 mm Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 345 long in fruit, surface rounded in cross-sec- tion, papery to membranous; styles 3.8-7 mm long. —Desert shrub, pinyon-juniper, sagebrush, and oak communities from 1220 to 2440 m elevation in Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Kane, San Juan, Sevier, Uintah, and Wayne coun- ties; Idaho and Utah eastward to Wyoming, Colorado, and New Mexico. This very critical species contacts P. chambersii to the west where apparent in- termediates are known; a similar situation occurs southward involving P. newberryi. Two varieties are known. 1. Flowers clear yellow, rarely fading purplish; silicles definitely indented below; plants widespread P. acutifolia var. acutifolia Flowers purple or tinged purple; silicles only slightly indented below; plants known only from the Book Cliffs area of Grand County P. acutifolia var. purpurea Var. acutifolia. [P. didijinocarpa var. aus- tralis Payson; P. australis (Payson) Rollins] Plants widespread, our common phase. Var. purpurea Welsh & Reveal, var. nov. Plantis similis P. acutifolia var. acutifolia sed petalis purpureis et siliculis pauci in- dentibus inferioribus. Type: Grand County, Utah, Sego Canyon, 5 miles north of Thompson, 1 May 1968, S. L. Wehh 6902. Holotypus, BRY. Paratype: Utah, Grand Coimty, about 3 miles north of Thompson, 5 May 1971, S. L. Welsh 10966 (BRY). Known only from the type locality. Physaria chambersii Rollins. [P. cham- bersii var. membranacea Rollins] Plants pe- rennial, cespitose, the caudex usually simple, clothed with marcescent leaf bases; stems 0.3-2.5 dm long, decumbent to ascending or erect, simple; basal leaves 1-7.5 (13) cm long, 0.3-3.5 cm wide, blade orbicular to ovate, obovate, elliptic or oblanceolate, en- tire, angular or toothed; cauline leaves greatly reduced; pedicels mostly 4-18 mm long, ascending to descending; sepals 6-7.2 mm long; petals 8-13 mm long, yellow, spatulate; silicles bladdery-inflated, upper sinus deeply indented, lower shallow or lacking, valves 11-17 mm long in mature fruit, surface roughly angled at edges, an- gles obtuse, papery to membranous; styles 4-8 mm long. —Desert shrub, sagebrush, oak-juniper, pinyon-juniper, aspen-mixed conifer, ponderosa pine, and spruce-fir com- munities from 1200 to 2900 m elevation in Beaver, Emery, Garfield, Iron, Juab, Kane, Millard, Piute, Sanpete, Sevier, Summit, Tooele, Utah, and Washington counties; Arizona and Utah west to California and Oregon. Physaria grahamii Morton. Plants pe- rennial, cespitose, caudex simple, clothed with marcescent leaf bases; stems 0.5-1.5 dm long, ascending to erect, simple; basal leaves 2-15 cm long, 0.8-5 cm wide, blade oblanceolate to spatulate, toothed to pin- natifid; cauline leaves often toothed, much reduced; pedicels 6-20 mm long, ascending; sepals 4.5-5 mm long; petals about 6.5 mm long, yellow, spatulate; siliques strongly in- flated, cordate, to 9 mm long and 10 mm wide; styles 3.5-7 mm long. Duchesne, Uin- tah, and Grand Cos.; Colorado. This distinctive taxon is still only poorly understood. We have seen only the holo- type (Chandler Canyon, 3 Aug 1935, Gra- Imm 9976, US), plus fruiting material gath- ered in August 1976 {Welsh et al. 14414, BRY). A second collection made bv Welsh (Welsh & Higgins 6239, BRY) from Garfield County, Colorado, seems to belong to this taxon. The species appears to be endemic to soils dferived from the Green River Shale Formation in the southern portion of the Uinta Basin and along the escarpments south of the Basin proper. Physaria newberryi A. Gray. [Coulterina newberryi (A. Gray) Kuntze; P. didymocarpa var. newberryi (A. Gray) M. E. Jones] Plants perennial, cespitose, caudex simple, clothed with marcescent leaf bases; stems 0.4-2.2 dm long, decumbent to ascending or erect; basal leaves 2-7.5 cm long, 0.8-4 cm wide, blade obovate to orbicular, ovate, or spatu- late, angled, indistinctly toothed or more commonly entire; cauline leaves much re- 346 Great Basin Naturalist Vol. 37, No. 3 duced, entire; pedicels 5-17 mm long, as- cending to descending; sepals 6-8.5 mm long; petals 6.5-12 mm long, yellow, spatu- late; silicles bladdery-inflated, upper sinus deeply indented, lower sinus shallow or lacking, valves 8-11 mm long or more in fruit, sharply angled at margins, papery; styles 2-3.5 mm long. —Warm desert shrub upwards to ponderosa pine communities from 1200 to 2000 m in Grand, Kane, San Juan and Washington counties; Arizona and New Mexico. Where this species contacts either P. chambersii or P. aciitifolia there are transi- tional specimens. Style length forms a con- tinuum from one entity to another, and er- rors in determination are possible, especially in material which lack mature fruits. Raphanus L. Plants pubescent with simple hairs, an- nuals, from tuberous taproots; leaves alter- nate and basal, simple, lyrate-pinnatifid, cauline ones petiolate to subsessile, not au- riculate; flowers racemose, pedicels not sub- tended by bracts; sepals 4, deciduous; petals 4, white or pink to lavender; stamens 6, fil- aments lacking glandular processes; styles apical on a tapering sterile beak, stigma minute, bilobed; fruit a silique, many times longer than broad, terete, indehiscent, breaking irregularly at maturity into seg- ments, valves several-grooved; seeds unise- riate. About 6 Eurasian species. Raphanus sativus L. Radish. Plants erect, stems mostly 4-10 dm tall, simple or more commonly branched, more or less hispid with simple hairs; leaves basal and cauline, mostly 2-18 cm long, 0.5-6.5 cm wide, ly- rate-pinnatifid, hispid, cauline ones reduced upwardly; pedicels 10-20 mm long or more, spreading to ascending; sepals 7-9 mm long, green to reddish or purplish, glabrous; pet- als 12-20 mm long, white to pink or laven- der; siliques 30-60 mm long, 5-10 mm wide, beak 10-25 mm long. —Cultivated for the edible roots, rarely escaping but usually not persisting; introduced from Europe. The yellow-flowered species, R. ra- phanistrum L., is an occasional weed which Ls to be expected in the state. RoRipPA Scop. Plants glabrous to hirsute with simple hairs, annual, biennial or perennial, from taproots or rhizomes; leaves alternate and basal, lyrate-pinnatifid, sinuately toothed, lobed, or uppermost subentire, petiolate to sessile, auriculate in some; flowers race- mose, pedicels not subtended by bracts; sep- als 4, deciduous; petals 4, yellow, fading pinkish in some, truncate to rounded api- cally; stamens 6, filaments lacking glandular processes; style prominent to almost lacking, stigma capitate; fruit a sessile silique or si- licic from one to several times longer than broad, terete or somewhat flattened; seeds several to numerous, mostly biseriate. A genus of perhaps 40 species on all ma- jor continents, but mainly in the northern temperate regions of the world. Stuckey, R. L. 1972. Taxonomy and distri- bution of the genus Rorippa (Cruciferae) in North America. Sida 4:279-430. 1. Plants perennial, with creeping rhizomes; petals 2.5-5 mm long 2 Plants annual or biennial, rarely perennial, from taproots; petals 0.7-2.5 (3) mm long 3 2(1). Plants 3-9 dm tall; sepals 1-2 mm long; fruits globose, usually seedless; intro- duced weeds of cultivated lands R- austriaca Plants 1-3 dm tall; sepals 2-4 mm long; fruit ovoid to oblong, bearing seeds; indigenous plants of saline valley bottoms R- siniiata 3(1). Siliques globose, 1-1.3 times longer than broad; partition circular in outline .... R. sphaerocarpa Siliques cylindroid or tapering at the apex, 2-5 times longer than wide or more; partition oblong to triangular in outline 4 Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 347 4(3). Plants prostrate to decumbent, stems 2 dm long or less; siliques tapering to apex, minutely papillose R. tenerrima Plants prostrate to decumbent or erect, stems often over 2 dm long; siliques cylindroid or tapering to apex, smooth 5 5(4). Plants erect, stems mostly 3-10 dm long; pedicels usually as long as fruit, mostly 4-12 mm long R. islandica Plants prostrate, decumbent, ascending, or erect, mostly 2-5 dm tall; pedicels usually shorter than fruit, mostly 2-4 mm long R. ctirvipes Rorippa austriaca (Crantz) Besser. [Nas- turtium austriacwn Crantz; Radiciila austri- aca (Crantz) Small] Austrian field-cress. Pe- rennial, from thickened rhizomes, stems 3-9 dm tall, erect, slender, finely puberulent; leaves 3-6 cm long, oblong to oblong-ovate, unequally serrate, glabrous, narrowed to a petiolelike auriculate base; racemes 7-12 cm long, in terminal panicles; pedicels 4-10 mm long in fruit, spreading-ascending; sepals 1-2 mm long; petals 3-15 mm long, yellow; si- licles subglobose, 1.5-3 mm long. —Occa- sional to rare weed of cultivated places, re- ported for south-central Utah by Stuckey (1972). Rorippa curvipes Greene. Annual or short-lived perennial, arising from taproots, stems (1) 2-5 diTi long, prostrate to decum- bent, ascending or erect, glabrous; leaves (2) 4-8 (12) cm long, 0.3-1.5 cm wide, oblong, obovate, spatulate or oblanceolate, entire, crenate, irregularly serrate, or lower ones pinnatifid, petiolate to sessile, often auricu- late; racemes 4-15 cm long, in axillary and terminal racemes; pedicels mostly 2-5 mm long in fruit, spreading to ascending or de- scending; sepals 0.8-1.7 mm long; petals 0.5- 2.8 mm long, yellow, rarely fading pinkish; silicles 1.8-8.7 mm long, 0.6-2.3 mm wide; style 0.2-1.3 mm long. —Moist sites from low to high elevation throughout much of Utah; western North America from southern Canada to Arizona and New Mexico. Three more or less distinct varieties are recognized. 1. Petals 1.2-2.8 mm long, longer than sepals; siliques usually acute; plants pros- trate to decumbent R. curvipes var. alpina Petals mostly 0.5-1.5 mm long, about as long as or shorter than sepals; sili- ques acute or obtuse; plants prostrate to decumbent or erect 2 2(1). Siliques 1.4-5 mm long, acute to somewhat obtuse; petals 0.5-1 (1.3) mm long R. curvipes var. curvipes Siliques 3.5-8 mm long or more, obtuse apically; petals 1-1.5 mm long R. curvipes var. integra Var. alpina (S. Wats.) Stuckey. [Nastur- tium ohtusum var. alpinum S. Wats.; R. ob- tusa var. alpina (S. Wats.) Britton; jR. al- pina (S. Wats.) Rydb.; Radicula alpina (S. Wats.) Greene] Bogs and seeps from 1980 to 3050 m elevation in Cache, Duchesne, Garfield, Summit (type from head of Bear River Canyon), and Utah counties; Idaho, Wyoming, and Colorado. Var. curvipes. [Cardamine palustris var. jonesii Kuntze; R. underwoodii Rydb.; Radi- cula curvipes (Greene) Greene; Radicula underwoodii (Rydb.) Heller; Rorippa obtusa and Rorippa curvisiliqua of Utah authors] Ponds, streams, and seep margins from 1370 to 2440 m elevation in Garfield, Grand, Kane, Piute, Salt Lake (type of var. jonesii from City Creek Canyon), Sanpete, Sevier, Utah, Wasatch, and Washington covmties; Alberta and Saskatchewan east to Wisconsin and south to California, Arizona, New Mex- ico, and Kansas. Var. integra (Rydb.) Stuckey. [R. integra Rydb.; Radicula integra (Rydb.) Heller; Ro- rippa obtusa var. integra (Rydb.) Marie-Vic- torin] Stream banks and beaches from 2440 348 Great Basin Naturalist Vol. 37, No. 3 to 3050 m elevation in Box Elder, Piute, spread throughout the Northern Hemi- Summit, Utah, and Wasatch (the probable sphere. type area) counties; Alberta south to Cali- Worldwide, or even within North Ameri- fornia and east to Montana and Wyoming. ca, this species is exceedingly complex. We have selected a conservative approach to Rorippa islandica (Oeder) Borbas. Annual the species, prefering to maintain R. po- or biennial, arising from taproots, stems 3- lustris (L.) Besser [Sisymbrium amphibium 10 dm tall, erect, glabrous or pubescent; var. pahistre L.; Sisymbrium pahistre (L.) leaves mostly 2-10 cm long, 0.6-2.5 cm Pollich; Radicula palustris (L.) Moench; wide, more or less pinnatifid, or cauline Nasturtiuiii pahistre (L.) DC] within the ones merely toothed, petiolate to sessile and broad definition of R. islandica [Sisymbrium more or less auriculate; racemes 3-10 cm islandicum Oeder], and thereby differing long or more, in axillary and terminal ra- from the recent treatment of the species cemes; pedicels mostly 4-10 mm long in complex in North America by Stuckey fruit, ascending to spreading or descending; (1972). Within the North American popu- sepals 1.2-2.5 mm long; petals 0.8-3.5 mm lations, Stuckey recognizes several sub- long, yellow, fading pinkish or purplish; si- species and varieties, denoting a bewilder- liques 3-8 mm long, rarely more, 2-3 mm ing array of morphological and wide, valves glabrous; styles 0.2-1.2 mm geographically reinforced infraspecific vari- long. —Lake shores, stream banks, springs, ation. In Utah, we have only 2 of the 11 and seeps, mostly in northern Utah; wide- entities he recognizes. 1. Leaves glabrous beneath; stems glabrous or pubescent only at base R. islandica var. glabra Leaves hairy beneath; stems pubescent usually to apex R. islandica var. hispida Var. glabra (Q. E. Schulz) Welsh & Re- stems 0.8-5 dm long, often more or less de- veal, comb, nov., based on Nasturtium pa- cumbent, glabrous to minutely hairy; leaves lustre var. glabrum O. E. Schulz, Symb. An- (1) 2-8 cm long, oblong in outline, pinnati- till. 3:516. 1903. [Radicida glabra (O. E. fid, petiolate to sessile and somewhat au- Schulz) Britton; R. palustris ssp. glabra (O. riculate; racemes 3-15 (25) cm long, in ax- E. Schulz) Stuckey] Moist places in Garfield illary and terminal racemes; pedicels 5-11 and Utah counties; Alberta south to New mm long in fruit, ascending to recurved; Mexico, with a disjunct series of populations sepals 2.7-4.5 mm long; petals 3.5-5.5 mm on Cuba (the type area). long, yellow, fading light yellow; siliques 5- Var. hispida (Desv.) Butters & Abbe. 12 mm long, about 1.5 mm wide, glabrous [Brachylobus hispidus Desv.; Sismybrium to roughened, narrowed to a style; styles 1- hispidum (Desv.) Poir.; Nasturtium hispidum 2.5 mm long.— Stream sides and other moist (Desv.) DC.; N. palustre var. hispidum sites at lower elevations, not definitely (Desv.) A. Gray; Radicula hispida (Desv.) known from Utah but reported from all of Britton; Radicula palustris var. hispida the surrounding states. (Desv.) B. L. Robinson; Rorippa palustris Rorippa sphaerocarpa (A. Gray) Britton. ssp. hispida (Desv.) Jonsell] Moist sites [Nastuiiium sphaerocarpum A. Gray; N. ob- froml370 to 2600 m elevation in Cache, tusum var. sphaerocarpum (A. Gray) Wats. Duchesne, Grand, Salt Lake, Summit, Utah, ex Allen; Radicula sphaerocarpa (A. Gray) and Weber counties; widespread in North Greene; Radicula obtusa var. sphaerocarpa America. (A. Gray) B. L. Robinson; Rorippa obtusa Rorippa sinuata (Nutt.) A. S. Hitchc. var. sphaerocarpa (A. Gray) Cory] Annual, [Nasturtium sinuatum Nutt. ex Torr. & arising from taproots, stems 1-4 dm long. Gray; Radicula sinuata (Nutt.) Greene] Pe- decumbent to erect, sparingly hirsute be- rennial, arising from a slender rhizome, low; leaves 3-10 cm long, 0.3-1.5 (3) cm Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 349 wide, entire, crenate, serrate or pinnatifid; racemes 2-10 cm long, in axillary and termi- nal racemes; pedicels 1.5-4.2 mm long, as- cending to recurved, often secund; sepals 0.7-1.3 mm long; petals 0.6-1.2 mm long, yellow, fading pinkish; siliques subglobose, 1-2.5 mm long and as wide or 1-1.3 times longer than broad; styles 0.4-0.7 mm long. —Moist sites at lower elevations in Garfield and Utah counties where evidently rare; Arizona and New Mexico northward to Wyoming. This species approaches both R. curvipes and R. islandica var. hlspkla. Despite the apparent convergences, R. sphaerocarpa ap- pears to exist as a fimctional entity and not just as an extreme morphological phase of either of the two taxa it approaches. Rorippa tenerrima Greene [Radicula te- nerrima (Greene) Greene] Annual, arising from taproots, the stems 0.4-2 dm long, de- cumbent to prostrate, glabrous; leaves 1-5 (8) cm long, 0.5-1.5 cm wide, lyrate-pinnati- fid to subentire; racemes 2-10 cm long, in axillary and terminal racemes; pedicels 1-3 (4) mm long, ascending to spreading; sepals 0.7-1.2 mm long; petals 0.6-0.8 mm long, yellow; siliques 3-8 mm long, 0.8-2 mm wide, tapering to apex; styles 0.3-1 mm long. —Marshy sites at lower to mid eleva- tions in Cache, Garfield, Kane, Iron, Salt Lake, Sevier, and Washington counties; Washington eastward to North Dakota and southward to Mexico. Schoencrambe Greene Plants glabrous, perennial, arising from a caudex and creeping rhizome; leaves alter- nate and some often basal, pinnatifid below, becoming entire above or all entire, petiol- ate to sessile, not auriculate; flowers ra- cemose, the pedicels not subtended by bracts; sepals 4, deciduous; petals 4, yellow; stamens 6, filaments lacking glandular pro- cesses; style almost lacking, stigma ex- panded, bilobed; fruit a sessile silique, many times longer than broad, terete, valves in- distinctly 1 -nerved; seeds uniseriate. A western American genus of 2 species. Schoencrambe linifolia (Nutt.) Greene. [Nasturtium linifolium Nutt.; N. pumilum Nutt., not St. Hilaire; Sisymbrium linifolium (Nutt.) Nutt ex Torr. & Gray; Sisymbrium pygmaeum Nutt. ex Torr. & Gray; Erysi- mum glaberrirnum Hook. & Am.; Schoen- crambe pygmaea (Nutt.) Greene; Schoen- crambe pinnata Greene; Schoencrambe decumbens Rydb.; Schoencrambe linifolia var. pinnata (Greene) A. Nels. in Coult. & Nels.; Sisymbrium decumbens (Rydb.) Blan- kinship; Sisymbrium linifolium var. pinna- tum (Greene) O. E. Schulz; Sisymbrium lini- folium var. decumbens (Rydb.) O. E. Schulz] Perennial, glabrous and usually glaucous, arising from a simple or more usually branched caudex, this from a creep- ing, deeply placed rhizome, stems (1.5) 2-9 dm tall, erect or less commonly ascending to decumbent, simple or branched; leaves 1.3-9.3 cm long, 0.1-2.5 cm wide, entire to deeply pinnatifid, basal ones often de- ciduous by midanthesis; pedicels 2-9 (10) mm long in fruit, ascending to spreading; sepals 4.3-6 mm long, yellowish; petals 7.5- 11 mm long, yellow, spreading; siliques 25- 65 (75) mm long, 0.8-1.2 mm wide, terete, sessile, erect or ascending; styles 0.3-0.6 mm long. —Salt desert shrub, sagebrush, and pi- nyon-juniper communities in Carbon, Dag- gett, Duchesne, Emery, Garfield (probable type location of Schoencrambe pinnata). Grand, Kane, Sanpete, Sevier, Summit, Uin- tah, Wasatch, and Wayne counties, and probably elsewhere; British Columbia east to Montana and south to Nevada and New Mexico. The indecision as to the generic position of this plant is indicated by the numerous generic names associated with it. The plants simulate Sisymbrium in a broad sense but do not show apparent relationship with the introduced annual or biennial weedy species of that genus. It seems best to place this plant within its own genus even though the principle diagnostic features involve vegeta- tive instead of floral characteristics. Sisymbrium L. Plants glabrous or hirsute annuals or rarely biennials, arising from taproots; leaves alternate and basal, variously lobed to entire, lower ones usually pinnatifid, re- duced upwards, petiolate to sessile, not au- riculate; flowers racemose, pedicels spread- 350 Great Basin Naturalist Vol. 37, No. 3 ing to erect, not subtended by bracts; sepals linear to tapering, terete, valves usually 3- 4, deciduous; petals 4, yellow; stamens 6, nerved; seeds several to many, uniseriate. filaments lacking glandular processes; styles A complex genus of perhaps 75 species in almost lacking, stigma bilobed; fruit a ses- temperate regions of the Old World and sile silique many times longer than broad. South America. 1. Leaves strongly dimorphic, lower lyrate-pinnatifid, uppermost with linear-fili- form lobes; pedicels ascending, siliques 50-90 mm long, spreading-ascend- ing to erect, not appressed; petals 6.3-8.5 mm long P. altissimum Leaves not dimorphic, upper and lower lobed about same; pedicels ascending to erect, siliques 10-45 mm long, erect or ascending, appressed or not; pet- als 2-4 mm long 2 2(1). Siliques and pedicels ascending to spreading, not appressed to rachis, siliques (20) 25-45 mm long S. ino Siliques and pedicels appressed-erect, siliques 10-15 mm long S. officinale Sisymbrium altissimum L. [Norta altis- Sisymbrium officinale (L.) Scop. [Erysi- sima Britton] Jim Hill Mustard; Tumbling mum officinale L.] Hedge Mustard. Annual, Mustard. Annual, stems 2.5-10 dm tall or stems 2.5-8 dm tall or more, hispid-hirsute more, sparingly to densely hirsute near base, throughout; leaves 1.5-20 cm long or more, usually glabrous above; leaves 1-20 cm long lyrate-pinnatifid to pinnatifid, not especially or more, petiolate, lower ones pinnatifid or dimorphic, upper ones merely reduced; merely lobed, becoming pinnatifid into line- pedicels 2-3 mm long, stout, erect, tip filiform segments upwardly; pedicels 4-10 about as thick as the silique; sepals 1.5-2.2 mm long, stout, almost or quite as thick as mm long, green or yellow; petals 3-4 mm the silique, ascending to spreading; sepals long, yellow, fading white, narrowly oblan- 3.6-5 mm long, often yellowish; petals 6.3- ceolate, ascending; siliques 8-15 mm long, 8.5 mm long, yellow, fading white, obovate appressed-erect, tapering to beaklike tip, to spatulate, spreading; siliques 50-90 mm valves 3-nerved. —Uncommon weedy plant long, 1-1.5 mm wide, terete, spreading to of disturbed sites in Utah County and to be ascending, valves evidently 3-nerved. — Usu- expected elsewhere; widespread in North ally in open disturbed sites at lower and America; adventive from Europe, middle elevations in Cache, Davis, Duch- esne, Emery, Iron, Juab, Kane, Millard, Smelowskia C. A. Meyer Piute, Salt Lake, San Juan, Tooele, Utah, Plants pubescent perennials, pulvinately Wasatch, Washington, Wayne, and Weber cespitose, arising from a branching caudex, counties, and probably cosmopolitan; wide- pubescence with branched and often with spread in North America; adventive from some simple hairs; leaves alternate or Europe. chiefly basal, usually pinnatifid, petiolate to Sisymbrium irio L [Norta irio (L.) Brit- sessile, not auriculate; flowers subcorymbose ton] Annual, stems 2-8 dm tall, erect, to racemose, pedicels not subtended by glabrous; leaves 1.5-10 (20) cm long, run- bracts; sepals 4, deciduous; petals 4, white, cinate-pinnatifid to pinnatifid, reduced up- rarely purplish; stamens 6, filaments lacking wardly; pedicels 6-10 mm long, slender, as- glandular processes; styles prominent, cending; sepals 2-2.5 (3) mm long, greenish stigma expanded; fruit a silique, 3- to sever- or yellowi.sh; petals 3-4 mm long, yellow, al times longer than broad, subterete or oblanceolate, spreading-ascending; siliques compressed, valves 1-nerved; seeds few to (20) 25-45 mm long, 0.8-1 mm wide, as- several per locule. cending. —Weed of dry sites in Beaver and A genus of perhaps four species of Asia Washington counties where locally com- and North America. Smeloivskia holmgrenii mon; California to Arizona; adventive from Rollins clearly does not belong to this Europe. genus, and this Nevada endemic probably Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 351 ought to be placed in its own monotypic south to Nevada, Utah, and Colorado. Our genus. material is var. americana (Kegel & Herd.) Drury, W. H., and R. C. Rollins. 1952. The Drury & Rollins [Hutchinsia cahjcina var. North American representatives of Sme- americana Regel & Herd.; S. americana lowskia. Rhodora 54:85-119. (Regel & Herd.) Rydb.; S. lineariloha Rydb.; Smelowskia calycina C. A. Meyer. Plants S. lineariloba f. virescens O. E. Schulz]. perennial, cespitose, with short to elongate caudex branches, these clothed with marces- Stanleya L. cent leaf bases; stems 0.4-2 dm tall, pub- Perennial, glabrous to simple pubescent escent with short, branched and long, herbs or subshrubs, arising from taproots or simple or branched hairs; basal leaves 0.5-5 caudices; leaves alternate and basal, pinnati- cm long, 0.4-1.6 cm wide, pinnately di- fid to entire, petiolate to sessile, auriculate vided; cauline leaves reduced upwardly; in some; flowers showy, racemose, pedicels pedicels mostly 3-8 mm long in fruit, as- not subtended by bracts; sepals 4, de- cending to spreading-ascending, pubescent; ciduous; petals 4, yellow to greenish-yellow; sepals 2-3.2 mm long, often tinged purplish, stamens 6, filaments lacking glandular pro- pubescent; petals 3-8 mm long, white to cesses, long-exserted; styles lacking to prom- cream or tinged pink to lavender, ovate, inent, stigma small; fruit a long-stipitate si- spreading; siliques 5-9 mm long, slightly lique, linear, terete to compressed, valves 1- flattened parallel to the septum or terete; or more-nerved; seeds numerous, uniseriate. styles 0.2-1 mm long; seeds 6-10. —Alpine A western North American genus of sev- tundra from 2900 to 3900 m elevation in en species. Beaver, Daggett, Duchesne, Grand, Iron, Piute, Salt Lake, Uintah, and Utah counties; Rollins, R. C. 1939. The cruciferaceous Alaska Yukon and Northwest Territories genus Stanleya. Lloydia 2:109-127. 1. Middle and upper cauline leaves sessile, auriculate; leaves all entire; petal- claw glabrous on both surfaces S. viridiflora Middle and upper cauline leaves petiolate to subsessile, not auriculate; leaves various; petal-claw pubescent on the inner or outer surfaces 2 2(1). Leaves entire or merely toothed, basal ones ovate to elliptic, cauline ones ovate to lanceolate or oblanceolate; plants of east-central and northeast- ern Utah S. integrifoUa Leaves, at least some, pinnatifid, only upper ones entire and usually lanceo- late to lance-linear; distribution various 3 3(2). Plants woody at base, caudex well developed; blades of petals mostly 1-3 mm wide; our most common and widespread species of the genus S. pinnata Plants herbaceous, caudex not developed; blades of petals 3-6 mm wide or more; an uncommon to rare plant of southeastern Utah S. albescens Stanleya albescens M. E. Jones. Biennial, hairy within; siliques 30-60 mm long, 1.5- caudex neither woody nor well developed; 2.5 mm wide, subterete, stipes 10-15 mm stems 2.5-10 dm tall, glabrous and glaucous, long. — Semidesert shrublands upwards to simple or branched; leaves mostly 5-15 cm pinyon-juniper woodlands, to be expected in long, 1-5 cm wide, lyrate-pinnatifid or up- Grand or San Juan counties, not presently per ones reduced in size and subentire to known from the state; Colorado and Ari- entire, glabrous and glaucous; pedicels zona. mostly 5-10 mm long, spreading, glabrous; Stanleya integrifoUa James. [S. pinnati- sepals 10-15 mm long, white with green fida var. integrifoUa (James) B. L. Robinson; tips, glabrous, reflexed; petals 10-16 mm S. glauca var. latifolia Cockerell; S. pinnata long, blade 3-6 mm wide or more, claw var. integrifoUa (James) Rollins] Perennial, 352 Great Basin Naturalist Vol. 37, No. 3 caudex well developed and more or less woody; stems 2.5-9 dm tall or more, spar- ingly hairy, glaucous, simple or branched; leaves 0.8-12.3 cm long, 0.3-3.5 cm wide, elliptic to ovate or lanceolate, all entire or lower ones merely toothed, puberulent, glaucous; sepals mostly 9-19 mm long, yel- lowish, reflexed, glabrous; petals 10.5-15.5 mm long, blade 1.4-2.6 mm wide, claw hairy within; siliques 33-75 mm long, 1.2- 1.8 mm wide, subterete, stipes 10-25 mm long. —Clay soils in desert shrub and pi- nyon-juniper communities from 1530 to 2000 m elevation in Daggett, Duchesne, Emery, and Uintah counties; Utah and Wyoming eastward to Kansas. Our material is distinctive and about equivalent in diagnostic features found among the other species of Stanleya in Utah. Stanleya pinnata (Pursh) Britton. [Cleome pinnata Pursh; S. pinnatifida Nutt.; S. het- erophyUa Nutt. ex Torr. & Gray; S. friiti- cosa Nutt.; S. arcuata Rydb.; S. canescens Rydb.; S. glauca Rydb.] Prince's Plume. Perennial, caudex well developed and slightly to very woody; stems (2.5) 3.5-12 (15) dm tall or more, glabrous to pilose, glaucous, simple or branched; leaves mostly 5-18 cm long, 2-5 cm wide or more, lan- ceolate to elliptic in outline, pinnatifid or upper usually entire and narrowly lanceo- late to elliptic, not auriculate, glabrous or sparsely pilose; pedicels 4-13 mm long, spreading; sepals 11-22 mm long, yellowish, reflexed, glabrous; petals 11-17 mm long, blade 1-3.8 mm wide, claw hairy within; si- liques (30) 35-70 (80) mm long, 1.2-2 mm wide, subterete to flattened, stipes 12-24 mm long, puberulent at base. — Seleniferous soils derived from shales, mudstones, and siltstones in many geological formations, from 850 to 2900 m elevation in Beaver, Box Elder, Carbon, Emery, Garfield, Grand, Iron, Juab, Kane, Millard, San Juan, San- pete, Sevier, Uintah, Utah, Washington, and Wayne counties; Idaho east to North Da- kota south to California, Arizona, and New Mexico. This plant is a primary indicator of the poisonous element selenium, because the plant is common in such soils. Our material belongs to var. pinnata, but the var. gibbe- rosa Rollins, known from Uinta County, Wyoming, is to be sought in Summit Coun- ty, Utah. Its leaves are all bipinnate. The var. inyoensis (Munz & Roos) Reveal may be encountered in Washington County; it differs from var. pinnata in being a woody subshnib. Stanleya viridiflora Nutt. ex Torr. & Gray. [S. collina M. E. Jones] Desert Plume. Perennial, caudex simple or branched, some- what woody at base; stems 3-12 dm tall or more, glabrous and glaucous, simple or branched; leaves 7-30 cm long, basal ones 1.8-8.7 cm wide, petiolate, blades united or somewhat nmcinate-pinnatifid, upper ones entire, sessile and auriculate, glabrous; pedi- cels mostly 6-12 mm long, spreading; sepals 11-20 mm long, yellow, reflexed, glabrous; petals 10-19 mm long, blade 0.8-1.5 (3) mm wide, not much wider than glabrous claw; siliques 35-70 mm long, subterete, stipe 14- 25 mm long. — Gypsiferous or clay soils in sagebRish and pinyon-juniper communities from 1680 to 2450 m elevation in Du- chesne, Emery, Salt Lake (?), Summit, Uin- tah, and Wayne counties; Idaho and Wyom- ing southwestwardly to Nevada and Oregon. Streptanthella Rydb. Plants glabrous, annual or winter annual, arising from taproots; leaves alternate, en- tire to shallowly dentate, tapering to base, not auriculate; flowers racemose, pedicels spreading-recurved to recurved, not sub- tended by bracts; sepals 4, deciduous; petals 4, white or purplish; stamens 6, the fila- ments lacking glandular processes; style ob- solete, small capitate stigma borne atop a beaklike extension of fruit; fruit a subsessile, slender silique many times longer than broad, strongly compressed, valves 1-nerved, dehiscent at base only; seeds several to many per locule, uniseriate. A monotypic genus of western North America. Streptanthella longirostris (S. Wats.) Rydb. [Arabis longirostris S. Wats.; Strep- tanthiis longirostris (S. Wats.) S. Wats.; Eu- klisia longirostris (S. Wats.) Rydb.; Thelypo- dium longirostris (S. Wats.) Jepson] Annual, Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 353 steins erect or ascending, usually branched throughout, 1-5 dm tall; leaves 1.5-8.5 cm long, 0.1-1.2 cm wide, oblanceolate to ellip- tic or lance-linear, sinuate-dentate to entire, lower ones usually deciduous by anthesis, reduced upwardly; pedicels 1.5-6 mm long, curved in fruit; sepals 2-4.8 mm long, greenish or purplish, with scarious margins; petals 5-8 mm long, white with purplish veins; siliques 30-60 mm long, 1.5-2 mm wide, reflexed-descending, valves narrowed apically into an indehiscent beak 3-7 mm long; style obsolete. —Sandy, clayey, or gravelly soils in desert shrub, pinyon-juni- per, and low elevation grassland commu- nities from 900 to 1900 m elevation in Bea- ver, Carbon, Emery, Garfield, Grand, Kane, Millard, San Juan, Sevier, Uintah, Washing- ton, and Wayne counties; Washington east to Wyoming and southward to Baja Califor- nia and New Mexico. Streptanthus Nutt. Plants glabrous, perennial, arising from taproots, rarely rhizomatous; leaves alter- nate and basal, simple, dentate to entire, basal ones petiolate, upper ones becoming sessile, entire, and auriculate upwardly; flowers racemose, pedicels not subtended by bracts; sepals 4, deciduous; petals 4, chest- nut to brown-purple or purple; stamens 6, often in 3 pairs as regards length and posi- tion, filaments lacking glandular processes; styles conspicuous, expanded, stigma bi- lobed; fruit a short-stipitate silique many times longer than broad, much flattened, valves with one main nerve and more or less reticulate lateral nerves; seeds several to many per locule, uniseriate. A genus of perhaps 25 species of the western United States. Streptanthus cordatus Nutt. ex Torr. & Gray. [Euklisia cordata (Nutt.) Rydb.; Car- tiera corclata (Nutt.) Greene; S. crassifolia Greene] Perennial, stems not inflated, 1.8- 5.7 (8) dm tall, glabrous and glaucous; basal leaves 1.5-8 (15) cm long, 0.5-2 (5) cm wide, spatulate to obovoid or oblanceolate, variously dentate, often sharply ciliate at least basally; cauline leaves becoming sessile and auriculate, ovate to oval or lanceolate; pedicels 4-9 mm long, ascending, glabrous; sepals (5) 7-10.5 mm long, usually purplish, glabrous except apically; petals 10-14.5 mm long, purple to chestnut, broad claw not constricted at juncture of blade, ascending to recurved; siliques 50-85 mm long, 3-5.8 mm wide, ascending to erect, glabrous; styles 1-3 mm long, expanded upwards, the stigma bilobed. —Sagebrush and pinyon- juniper communities from 1200 to 2600 m elevation in most of the counties of Utah; Oregon to Wyoming south to California, Arizona, and New Mexico. A specimen collected near Kanosh, Mil- lard County, by Pickford {130-OGDF) is definitely rhizomatous. Thelypodiopsis Rydb. Plants glabrous or pubescent with simple hairs, annual or biennial to perennial, aris- ing from taproots; leaves alternate or alter- nate and basal, simple, oblong-oblanceolate to lanceolate or ovate, toothed to entire, petiolate or sessile and auriculate or merely sessile; flowers in racemes, pedicels ascend- ing or spreading-ascending, not subtended by bracts except in some lowermost flowers; sepals 4, deciduous; petals 4, white, pink, lavender, or yellow; stamens 6, filaments lacking processes; style expanded upwards, stigma bilobed; fruit a sessile to stipitate si- lique, many times longer than broad, sub- terete to terete, valves 1 (3-5)-nerved; seeds uniseriate. A genus of western United States and perhaps northern Mexico of perhaps seven species. The species included herein have been treated as members of Streptanthus, Thely- podium, an expanded Sisymbrium, and even Caulanthus. The generic problem has been reviewed by Al-Shebaz (1973; see Thehjpo- dium), but the solution adopted by that au- thor, and followed in part here, is not alto- gether acceptable for our species. We have placed the wholly discordant Caulanthus di- varicatus Rollins in Thehjpodiopsis, and as Thelypodiu7n sagittatum mirrors both The- lypodiopsis elegans and T. amhigua that en- tity might well be placed in the latter genus also. 354 Great Basin Naturalist Vol. 37, No. 3 1. Leaves merely sessile, not auriculate, all cauline; plants perennial, from a branching caudex T. argillacea Leaves auriculate, at least the cauline; plants biennial or winter annual 2 2(1). Petals yellow 3 Petals pink, lavender, or white 4 3(2). Pedicels, sepals, and siliques glabrous; petal blade not especially constricted at juncture with petaloid claw T. atirea Pedicels, sepals, and siliques sparingly villous; petal blade conspicuously con- stricted at juncture with broadened membranous claw T. divaricata 4(2). Pedicels, sepals, and siliques glabrous; basal leaves often over 6 times longer than broad; plants known from Kane County T. ambigua Pedicels, sepals, and siliques sparingly villous; basal leaves usually less than 5 times longer than broad; plants of eastern Utah T. elegans Thelypodiopsis ambigua (S. Wats.) Al- Shebaz. {Thehjpodium ambiguiim S. Wats.). Biennial or short-lived perennial, stems not inflated, 2-10 (12.5) dm tall, glabrous throughout; leaves basal and cauline, basal ones 3-15 (20) cm long, 0.6-2.8 cm wide, si- nuate-dentate to entire, cauline ones re- duced upwards, becoming entire and lance- ovate to lanceolate, 0.5-5.5 cm long, 0.3-1.8 cm wide, sessile and auriculate; pedicels 4- 12 cm long, curved-ascending to spreading- ascending, glabrous; sepals 5-7 mm long, of- ten pinkish-hyaline, glabrous, erect; petals 9-12 mm long, blade 3-4 mm wide, pink to lavender or white, claw broad, slightly con- stricted at juncture of blade; anthers 2.5-3.5 mm long; siliques 55-75 mm long, 1.2-2 mm wide, short-stipitate, terete, ascending to erect; styles 1-2.5 mm long, expanded up- wards, stigmatic lobes developed. —Clay to sandy and gravelly soils in pinyon-juniper community from 1530 to 1830 m elevation in Kane '(Eastwood & Howell 9300-US; Welsh & Atwood 9706-BRY) and Washing- ton (Palmer 27, southern Utah, 1877-US) c-ounties; Mohave Coimty, Arizona. Thelypodiopsis argillacea Welsh & At- wood. Plants perennial, glabrous, glaucous; stems 1.3-3 dm tall, simple or branched above, arising from a subligneous caudex; leaves sessile, all cauline, not auriculate, lin- ear, 0.9-3 cm long, 0.8-2 mm wide, some- what fleshy, entire, acute to rounded; ra- cemes (5) 8- to 22-flowered; pedicels 7-13 mm long, ascending; sepals 4.2-6.5 mm long, purplish, margins hyaline; petals 7.8- 10.9 mm long, 2.5-3.2 mm wide, white to lilac, with conspicuous purplish veins, claw not differentiated from the blade; anthers 1.7-2.5 mm long; siliques 18-25 mm long, 1- 1.2 mm wide, subsessile, terete, ascending to erect; styles 0.5-1 mm long, expanded up- wards, stigmatic lobes developed. Desert shrublands on Green River Shale, at lower elevations in Uintah County; endemic. Thelypodiopsis aurea (Eastw.) Rydb. [Thehjpodium aureum Eastw.; Sisymbrium aureum (Eastw.) Payson] Biennial or rarely short-lived perennial, stems not inflated, 1-7 dm tall, glabrous throughout or somewhat pubescent below with flattened, flexuous hairs; leaves basal and cauline, basal ones 2.5-6.5 cm long, 0.5-1.3 cm wide, irregu- larly toothed to subentire, cauline ones re- duced upwardly, becoming entire and lan- ceolate to ovate-lanceolate, 1.3-6.5 cm long, 0.4-2.3 cm wide, sessile and auriculate; pedicels 5-9 mm long, spreading-ascending, glabrous; sepals 5-9.3 mm long, yellowish, glabrous, ascending; petals 6.9-12.5 mm long, blade not expanded, 1.5-3.5 mm wide, yellowish, claw very broad, ascending limb not especially constricted at juncture with claw; anthers 2.7-3.7 mm long; siliques 40- 70 mm long, 1-1.5 mm wide, stipitate, stipe 2-7 mm long, terete, ascending to erect, glabrous; styles 1-3 mm long, expanded up- wards, stigmatic lobes well developed. — Clay to sandy soils in semidesert shrub and pinvon-juniper communities from 1500 to 2450 m elevation in San Juan County; Col- orado. Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 355 Thelypodiopsis divaricata (Rollins) Welsh & Reveal, comb, nov., based on Cimlanthiis divaricatus Rollins, Contr. Gray Herb. 201: 8. 1971. Biennial or winter annual, erect, simple, stems not inflated, 2-11 dm tall, her- bage more or less pubescent with long, tan- gled hairs at least below; leaves basal and cauline, basal ones 1.5-10 cm long, 0.4-2.8 cm wide, irregularly toothed or rarely pin- natifld, subentire or dentate at apex only, cauline ones reduced upwardly, becoming entire and ovate-lanceolate, 0.7-9 cm long, 0.4-2.8 cm wide, sessile and auriculate; pedicels 5-21 mm long, spreading-ascending, sparingly villous to glabrate; sepals (3.5) 4.8-7 mm long, yellowish, sparingly villous; petals 7-10 mm long, yellowish, claw very broad, ascending limb constricted at junc- ture with claw; anthers 2.5-3.8 mm long; si- liques 40-80 (90) mm long, 1-1.5 mm wide, sessile or subsessile, terete, ascending to erect, sparingly villous when young, becom- ing glabrous with age; styles 1-2 mm long, with stigma reduced or expanded and lobes not or only moderately well developed. — Clay soils in shadscale, blackbrush, and pi- nyon-juniper communities from 1280 to 1700 m elevation in Carbon, Emery, Gar- field, Grand, San Juan, and Wayne counties; endemic. Thelypodiopsis elegans (M. E. Jones) Rydb. [Thehjpodiitm elegans M. E. Jones; Sisijmbritmi elegans (M. E. Jones) Payson; Streptanthus wyorningensis A. Nels.; The- lypodiopsis wyomingensis (A. Nels.) Rydb.] Biennial or rarely a winter annual or a short-lived perennial, stems 1.2-9.5 dm tall, almost or quite glabrous to densely pu- bescent below with flattened, flexuous hairs; leaves basal and cauline, basal ones 1-6.5 cm long, 0.3-1.5 cm wide, sinuate-dentate to irregularly toothed or entire, cauline ones reduced upwardly, becoming entire and lan- ceolate to ovate-lanceolate or oblong, 0.3- 8.5 (12) cm long, 0.3-2 cm wide, sessile and auriculate; pedicels 5-18 mm long, curved- ascending to spreading-ascending, glabrous or sparingly villous, erect; petals 11-14.5 mm long, blade 3-6 mm wide, pink to lav- ender or white with pinkish veins, claw very broad, not constricted at juncture with blade; anthers 2.6-3.2 mm long; siliques 45- 75 mm long, 1.2-1.8 mm wide, sessile or subsessile, terete, ascending to erect, glabrous or sparingly villous; styles 1.8-3 mm long, expanded upwards, stigmatic lobes well developed. —Clay, sandy, or gravelly soils in pinyon-juniper, sagebrush, and desert shrub communities from 1500 to 2300 m elevation in Duchesne, Grand, and Uintah counties; Wyoming, Colorado. It has not been possible to segregate T. wyomingensis from the bulk of T. elegans, and that species is hereby reduced to synon- ymy. Thelypodium Endl. Plants glabrous or pubescent with simple hairs, annuals, biennials, or short-lived per- ennials, arising from taproots; leaves alter- nate and basal, simple, lyrate-pinnatifid, toothed or entire, petiolate or sessile, au- riculate in some; flowers racemose, pedicels not subtended by bracts; sepals 4, de- ciduous; petals 4, white, pink, lavender, or purple; stamens 6, filaments lacking glandu- lar processes; styles slender to stout, cylin- dric to somewhat expanded upwards, stigmas small, entire or somewhat bilobed; fruit a stipitate silique many times longer than broad, terete or somewhat flattened, valves 1 (3)-nerved; seeds uniseriate. A genus of about 10 species of western North America. Pennellia micrantha (A. Gray) Nieuwl. [Thelypodium micranthum (A. Gray) S. Wats.; Heterothrix micrantha (A. Gray) Rydb.] has been repeatedly reported from Utah, but we have failed to find any speci- mens definitely known from the state in any of the herbaria we have consulted. It differs from the species of Thelypodium reported here in having stellately pubescent leaves. Al-Shebaz, I. A. 1973. The biosystematics of the genus Thelypodium (Cruciferae). Contr. Gray Herb. 204 :3-148. Payson, E. B. 1922. A monographic study of Thelypodium and its immediate allies. Ann. Missouri Bot. Card. 9: 233-324. 1. Cauline leaves definitely auriculate 2 Cauline leaves petiolate or sessile but not auriculate 3 356 Great Basin Naturalist Vol. 37, No. 3 2(1). Cauline leaves linear to oblong or oblong-lanceolate, appressed to stem; basal leaves 1-7 cm long, 0.4-1.8 cm wide T. rollinsii Cauline leaves oblong to lanceolate, not appressed to stem; basal leaves 6-20 cm long, 0.6-5 cm wide T. sagittatum 3(1). Plants robust biennials, mostly 5-15 dm tall or more; racemes borne in co- rymbose panicles T. integrifolium Plants various, but if (as rarely) as above, then racemes seldom if ever borne in corymbose panicles 4 4(3). Racemes mostly 1-8 dm long or more; petals 8 mm long or more, differen- tiated into a blade and claw; plants known only from Box Elder County T. millefolium Racemes mostly less than 1 dm long; petals 4-8 mm long, hardly differen- tiated into a blade and claw; plants not of Box Elder County 5 5(4). Sepals ascending to spreading; petals 1.5-3.5 mm wide; eastern Utah T. laxiflorum Sepals spreading to reflexed; petals 1-2 mm wide; Henry Mountains, Garfield County T. wrightii Thelypodium integrifolium (Nutt.) Endl. glabrous; petals 5-10 mm long, mostly 1-2 Biennial, stems mostly 5-30 dm tall, simple mm wide, white to lavender or purple; an- or branched, glabrous and glaucous; basal thers 1-3 mm long; siliques 10-45 mm long, leaves 5-30 cm long, 1.5-10 cm wide or 1-2 mm wide, stipitate, stipes up to 2 mm more, spatulate to lanceolate, ovate or obo- long; styles slender, 0.5-3 mm long; stigma vate, entire to denticulate; cauline leaves 2- entire. —Widespread at the lower elevations 12 (15) cm long, 0.2-2 cm wide, oblanceo- in Utah; common throughout much of the late to elliptic or lanceolate, sessile, not au- American West. riculate; pedicels 3-10 mm long, variously Four more or less distinctive varieties are spreading, strongly to moderately flattened known, at base; sepals 3-7 mm long, often purplish, 1. Mature fruiting pedicels whitish, stout; flowers white; plants of southwestern Utah T. integrifolium var. affine Mature fruiting pedicels not whitish or only seldom so, slender or stout; flow- ers pink to lavender or white; plants not of southwestern Utah 2 2(1). Mature fniiting pedicels not or only somewhat flattened at base; plants most- ly of south-central to north-central Utah T. integrifolium var. integrifolium Mature fniiting pedicels strongly flattened at base; plants of eastern and southeastern or northwestern Utah 3 3(2). Racemes mostly 1-8 cm long in fruit; stipes often less than 1 mm long; plants of northwestern Utah T. integrifolium var. complanatum Racemes mostly more than 8 cm long in fruit; stipes often more than 1 mm long; plants of eastern and southeastern Utah T. integrifolium var. gracilipes Var. affine (Greene) Welsh & Reveal, springs, irrigation canals, and stream sides comb, nov., based on Thehjpodium affine from 850 to 1100 m elevation in Kane, Mil- Greene, Pittonia 4:314. 1901. [T. rhomhoi- lard, and Washington counties; Nevada, deum Greene; Pleurophragma rhomhoideum California. (Greene) O. E. Schulz; T. integrifolium ssp. Var. complanatum (Al-Shebaz) Welsh & affine (Greene) Al-Shebaz]. Saline seeps, Reveal. [T. integrifolium ssp. complanatum Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 357 Al-Shebaz]. Uncommon, at lower elevations in Beaver and Box Elder counties; Utah westward to California and Oregon. Var. gracilipes B. L. Robinson. [T. graci- lipes (B. L. Robinson) Rydb.; Pleurophragiiia gracilipes (B. L. Robinson) Rydb.; P. platy- podiurn Rydb., the type is from Moab; T. rhomboidewn var. gracilipes (B. L. Robin- son) Payson; T. integrifolitim ssp. gracilipes (B. L. Robinson) Al-Shebaz]. Canyon bot- toms, terraces, and hanging gardens at low elevations in Garfield, Grand, Kane, San Juan, Uintah, and Wayne counties; Arizona, Colorado, and New Mexico. Var. integrifolium. [Pachypodium integri- folitim Nutt. ex Torr. & Gray; T. lilacinum Greene; Pleurophragma integrifoliimi (Nutt.) Rydb.; Pleurophragma lilacinum (Greene) Rydb.; T. lilacinum var. subumbellatum Payson]. Marshes, seeps, stream sides, and other moist sites in Cache, Iron, Piute, Salt Lake, Sevier, and Utah counties; Washing- ton east to the Dakotas and south to Ore- gon, Utah, Colorado, and Nebraska. Thelypodium laxiflorum Al-Shebaz [T. ivrightii var. tenellum M. E. Jones; Stan- leyella ivrightii var. tenella (M. E. Jones) Payson] Biennial, stems 1.5-15 dm tall or more, simple, glabrous throughout or hispid near base; basal leaves 3-15 cm long, 1-6 cm wide, pinnatifid to toothed; cauline leaves 3-9 cm long, 0.2-2 cm wide, sub- entire to entire, not auriculate; pedicels 4- 13 mm long, spreading to descending, some- what flattened at base; sepals 2.5-6 mm long, white to lavender, glabrous ascending to spreading; petals 4-8 mm long, 1.5-3.5 mm wide, white to lavender; anthers 1-3 mm long; siliques 20-75 mm long, 0.8-1.2 mm wide, stipitate, stipes 0.2-2 mm long; styles 0.5-2 mm long. —Mountain brush and forest communities from 1700 to 2450 m elevation in Carbon, Duchesne, Iron, Kane, San Juan, Utah (type from Slate Canyon), and Washington counties; Nevada to Colo- rado. Closely related to T. ivrightii. Thelypodium milleflorum A. Nels. [T. la- ciniatum var. milleflorum (A. Nels.) Payson] Biennial, stems mostly 4-12 dm tall, almost always hollow, simple or branched, glabrous and glaucous; basal leaves 6-15 cm long, 1-7 cm wide, oblong to lanceolate or ovate, toothed or pinnatifid; cauline leaves similar to the basal ones only gradually reduced upwardly, petiolate, not auriculate; pedicels 2-6 mm long, curved-ascending, slightly flat- tened at base; sepals 4-9 mm long, creamy- white, glabrous; petals 8-15 mm long, 1-2 mm wide, white; anthers 2.5-5 mm long; si- liques 25-85 mm long, 0.8-1.8 mm wide, stipitate, stipes 0.5-5 mm long; styles stout, 0.5-2 mm long; stigma entire. —Dry slopes and saline flats in Box Elder County; British Columbia south to California and east to Idaho and Utah. Thelypodium rolUnsii Al-Shebaz. Bien- nial, the stems (4) 5-16 (20) dm tall, simple or branched, glabrous and glaucous; basal leaves 1.3-7 cm long, 0.4-1.8 cm wide, spatulate to oblanceolate, toothed to sub- entire; cauline leaves 1-6 cm long, 0.1-0.8 cm wide, linear to narrowly lanceolate or oblong, erect, not flattened; sepals 4-7 mm long, lavender to purplish, glabrous; petals 6-10 mm long, 1.2-3 mm wide, lavender to purplish; anthers 2-4 mm long; siliques 20- 60 mm long, 0.7-1 mm wide, stipitate, stipe 0.5-6 mm long; styles 0.5-2 mm long; stigma entire. —Stream terraces, saline flats, and seeps in greasewood and saltgrass com- munities from 1370 to 1700 m elevation in Carbon, Juab, Millard, Piute, Sanpete, and Sevier counties; endemic. Thelypodium sagittatum (Nutt.) Endl. in Walpers. Biennial or short-lived perennial, stems 2-10 dm tall, rarely more, simple or more usually branched, glabrous or hirsute with simple hairs below; basal leaves 2-25 cm long, 1-5 cm wide, ovate to oblanceo- late or oblong, entire; cauline leaves 0.7-10 (14) cm long, 0.2-2.8 cm wide, ovate to lan- ceolate or lance-oblong, entire, glabrous or puberulent especially on the veins, auricu- late; pedicels 2.5-15 (20) mm long, spread- ing to spreading-ascending; sepals 2.5-10 mm long, green to purplish, glabrous; petals 5-15 mm long, 1-4 mm wide, white to lav- ender or purple; anthers 1.5-5 mm long; si- liques 10-65 mm long, 0.5-1.2 mm wide, stipitate, stipes 0.3-2 mm long; styles 0.5-3 mm long. —Widespread in the state; Wash- ington south to California and eastward to Montana and Wyoming. 358 Great Basin Naturalist Vol. 37, No. 3 This species simulates Thehjpodiopsis ele- is divisible into three more or less dis- gans in some features and care should be tinctive varieties in Utah, taken in distinguishing the two. The species 1. Petals linear to oblanceolate, 1.5 mm wide or less; pedicels 2.5-9 mm long; plants of southwestern Utah T. sagittatum var. ovalifolium Petals oblanceolate to spatulate, 1-4 mm wide; pedicels 5-15 mm long or more; distribution various 2 2(1). Petals pink to lavender or purplish, or white fading lavender to purplish or white; siliques straight to curved, not or only slightly tonilose T. sagittatum var. sagittatum Petals white to pink, fading yellowish on drying; siliques flexuous-contorted and definitely torulose, wormlike T. sagittatum var. vermicularis Var, ovalifolium (Rydb.) Welsh & Reveal. [T. ovalifolium Rydb., type from Panguitch Lake; T. palmeri Rydb., type from southern Utah; T. sagittatum ssp. ovalifolium (Rydb.) Al-Shebaz]. Clay soils in Garfield, Iron, and Kane counties; Nevada. Var. sagittatum. [Streptanthus sagittatus Nutt.; Pachypodium sagittatum (Nutt.) Nutt. ex Torr. & Gray; T. nuttallii S. Wats.; T. amplifolium Greene; T. torulosum Heller; T. macropetalum Rydb., type from Davis County; Thelypodiopsis nuttallii (S. Wats.) O. E. Schulz; Thehjpodiopsis torulosa (Hel- ler) O. E. Schulz]. Clay or silty, often sa- line, soils from 1200 to 2300 m elevation in Box Elder, Cache, Davis, Duchesne, Salt Lake, Summit, Tooele, Utah, and Wayne counties; Washington and Montana south to Nevada, Utah, and Colorado. Var. vermicularis Welsh & Reveal, var. nov. A Thelypodio sagittato var. sagittato petalis discoloriter flavidis et siliquis flex- uosis et torulosis differt. Holotype: Utah: Sevier County, about 4 miles southeast of Sigurd along Utah Highway 24, on the Ara- pien Shale Formation in a greasewood com- munity, 29 May 1972, Welsh & Atwood 11718. Holotypus, BRY. Paratypes: Utah: Box Elder County, Grouse Creek, 27 May 1973, Albee 1355; Juab County Fish Springs Range, 9 May 1968, Reveal & Thomas 984 (BRY, US);' Millard County, west of Hinckley, 19 Apr 1930, Cottam 4622; 2 miles south of Black Rock Station, 14 May 1968, R. C. Holmgren 534; Sevier County, Glenwood, 18 May 1875, Ward s.n. (US); Utah County, Coyote Pass, 1.2 mile west of Utah Highway 68, 15 May 1968, Weston 71 (all BRY imless otherwise indicated); endem- ic to Utah. Thelypodium wrightii A. Gray. [Stan- leyella wrightii (A. Gray) Rydb.] Biennial, stems 1.5-10 din tall or more, simple or more commonly branched, glabrous throughout or hispid near the base; basal leaves 3-15 cm long, 1-6 cm wide, pinnati- fid to lyrate-pinnatifid; cauline leaves 3-9 cm long, 0.2-2 cm wide, pinnatifid to si- nuate-dentate, rarely subentire, not auricu- late; pedicels 4-13 mm long, spreading to descending, somewhat flattened at base; sepals 2.5-6 mm long, white to lavender, glabrous, spreading to reflexed; petals 4-8 mm long, 1-2 mm wide, white to lavender; anthers 1-3 mm long; siliques 20-75 mm long, 1-1.5 mm wide, stipitate, stipes 0.2-2 mm long; styles 0.5-1 mm long. —Rare and isolated in Utah well away from the bulk of range of this species, known only from a M. E. Jones collection from the Henry Moun- tains, Garfield County; Colorado and Ari- zona eastward to Texas and northern Mexi- co. Thlaspi L. Plants glabrous, annual or perennial, aris- ing from taproots; leaves cauline or cauline and basal, alternate, simple, entire to den- tate or lobed; flowers racemose, pedicels not subtended by bracts; sepals 4, de- ciduous; petals 4, white, sometimes pinkish or lavender; stamens 6, filaments lacking glandular processes; fruit a sessile silicle, compressed contrary to septum, often more or less wing-margined; style obsolete or Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 359 slender and conspicuous, stigma capitate; seeds 2 to several per locule, uniseriate. A genus of perhaps 50 species, mostly of Eurasia. Holmgren, P. K. 1971. A biosystematic study of North American Thkispi mon- tanwn and its allies. Mem. New York Bot. Card. 21:1-106. Payson, E. B. 1926. The genus Thlaspi in North America. Univ. Wyoming Publ. Bot. 1:145-186. 1. Plants annual; styles obsolete or up to 0.2 mm long; fruit orbicular in outline, conspicuously winged T. arvense Plants perennial; styles 0.3-2.5 mm long or more; fruit oblanceolate to obcor- date or obovate in outline, narrowly wing-margined or not winged at all .... T. montaniim Thlaspi arvense L. Annual, stems mostly 1-7 dm tall; basal leaves usually deciduous by anthesis; cauline leaves 1-8 cm long, 0.2- 2.5 cm wide, elliptic to lanceolate or oblan- ceolate, sinuate-dentate to pinnatifid or sub- entire, uppermost sessile and auriculate; pedicels 5-12 mm long or more, spreading to curved-ascending; sepals 1.5-2.5 mm long, green with whitish margins; petals 3- 4.5 mm long, white; silicles 10-17 mm long, 7-12 mm wide, strongly compressed, wing- margined all around; styles almost obsolete. —Weedy species of roadsides, meadows, fields, and other disturbed places from 1370 to 2450 m elevation in Box Elder, Cache, Davis, Grand, Rich, Salt Lake, Utah, Wasatch, and Weber counties, and to be ex- pected elsewhere; widespread in North America; adventive from Europe. Thlaspi montanum L. [T. alpestre of au- thors, not L.; T. cochlearifonne DC; r. nuttallii Rydb.; T. alpestre var. glaucum A. Nels.; T. glaucum (A. Nels.) A. Nels.; T. coloradoense Rydb.; T. purpurascens Rydb.; T. alpestre var. purpurascens (Rydb.) Os- tenf.; T. glaucum var. pedunculattim Pay- son; T. glaucum var. hesperiuin Payson; T. hesperium (Payson) G. N. Jones; T. fendleri var. coloradoense (Rydb.) Maguire; T. fend- leri var. tenuipes Maguire, type from San- pete Gounty; T. fendleri var. hesperium (Payson) G. L. Hitchc] Perennial, with simple or branched caudex branches, stems 0.2-4 dm tall; basal leaves 0.9-3.5 cm long or more, 0.2-1 (1.5) cm wide, oval to oblong or spatulate-oblanceolate; cauline leaves 0.5- 2.5 cm long, 0.1-1.2 cm wide, sessile and jjriculate; pedicels 2-15 mm long, spread- ■^ -^r spreading-ascending; sepals 1.5-3.5 mm long, greenish to purplish; petals 3.5- 7.5 mm long, white, pinkish or lavender, spatulate; silicles 3-8 mm long, rarely more, 1.5-5 mm wide, obovate to obcordate, wing- ed or not so; styles 0.5-2.5 mm long. —For- est, knmiholz, and alpine tundra commu- nities from 2150 to 3965 m elevation in Beaver, Box Elder, Davis, Daggett, Duch- esne, Emery, Garfield, Grand, Iron, Kane, Piute, Salt Lake, San Juan, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, Washington, and Weber counties; Washing- ton east to Montana and south to Galifor- nia, Arizona, New Mexico, and Texas; Eu- rasia. Our material belongs to var. montanum. Some specimens from the southern part of the state approach the more southern var. fendleri (A. Gray) P. K. Holmgren, which has styles longer than 2.5 mm in length, petals 6-13 mm long, and silicles 7-16 mm long and 4-9 mm wide. Thysanocarpus Hook. Plants glabrous or pubescent with simple hairs, annual, arising from taproots; leaves cauline, alternate, simple, oblong-lanceolate to linear, toothed to entire; flowers race- mose, pedicels recurved and not subtended by bracts; sepals 4, deciduous; petals 4, white or tinged pink or purple; stamens 6, the filaments lacking glandular processes; fruit a strongly compressed, unilocular, 1- seeded, indehiscent silicle, margin winged. About 5 species of western North Ameri- ca. Thysanocarpus curvipes Hook. Annual, stems 1-5 dm tall or more, simple or branched, sometimes hirsute below, other- 360 Great Basin Naturalist Vol. 37, No. 3 wise glabrous; leaves 1.2-5 cm long, 0.1-1 cm wide, lance-oblong to elliptic or linear, sinuate-dentate to entire, transitional up- wards to smaller sessile and auriculate blades; pedicels 3-7 mm long, recurved in fruit; sepals 1-1.5 mm long, often purplish; petals 1.5-2 mm long, white or tinged purplish or pinkish; silicles 4.5-8 mm long, 3-4 mm wide, ovate to obovate, often plano-convex, glabrous or pubescent, wing crenate and sometimes perforate; styles 0.4- 0.6 mm long. —Sand and gravelly soils in the warm desert shrub communities from 670 to 975 m elevation in Washington County; British Columbia south to Califor- nia and eastward to Utah and Arizona. Our material belongs to var. eradiattis Jepson. General References Arnow, L. a. 197L Vascular flora of Red Butte Can- yon, Salt Lake County, Utah. Unpublished mas- ter's thesis. Salt Lake City: University of Utah Li- brary. 388 pp. Graham, E. H. 1937. Botanical studies of the Uinta Basin of Utah and Colorado. Ann. Carnegie Mus. 26: 1-432. Harrington, H. D. 1954. Manual of the plants of Col- orado. Denver: Sage Books. 666 pp. Hitchcock, C. L. 1964. "Cruciferae." In: C. L. Hitch- cock, et al.. Vascular plants of the Pacific North- west. Univ. Wash. Publ. Biol. 17(2): 430-557. Holmgren, A. H. 1948. Handbook of the vascular plants of the northern Wasatch. San Francisco: Lithotype Process Co. 202 pp. Kearney, T. H., and R. H. Peebles. 195L Arizona flora. Berkeley: University of California Press. 19.32 pp. McDouGALL, W. B. 1973. Seed plants of northern Ari- zona. Flagstaff: Museum of Northern Arizona. 594 pp. MuNZ, P. A., and D. D. Keck. 1959. A California flora. Berkeley: University of California Press. 1681 pp. Welsh, S. L., and G. Moore. 1974. Utah plants, Tracheophyta. Provo: Brigham Young University Press. 474 pp. Index Alyssum alyssioides, 289 desertorum, 289 incaniim, .302 ludovicianum, .339 maritimum, 342 minus, 289 saxatile, 289 szowitsianum, 289 Anastatica syriaca, 327 Arabidopsis stenocarpa, 327 thaliana, 290 virgata, 327 Arab is albertina, 294 anguUita, 299 aprica, 293 arcuata, perennans, 298 secunda, 296 subvillosa, 301 arida, 299 artnerifolia, 297 heckwithii, 299 hrachycarpa, 292 hracteoUitu, 296 hrebneriana, 327 brevuiiliqiui, 292 bridgeri, 298 caduca, 296 canescens, 296 latifolia, 296 cobrensis confinis, 292 brachycarpa, 292 confinis, 293 interposita, 293 connexa, 294 consanguinea, 296 dacotica, 292 demissa, 293 demissa, 294 lanugida, 294 russeola, 294 densa, 297 densicaulis, 298 diehlii, 298 divaricarpa, 292 drummondii, 294 alpina, 297 brachycarpa, 292 connexa, 294 lyallii, 297 oreophila, 297 oxyphylla, 294 pratincola, 292 egglestonii, 296 eremophila, 298 exilis, 296 fendleri, 294 fendleri, 295 spatifolia, 295 fomiosa, 300 glabra, 295 furcatipilis, 295 glabra, 295 gracilenta, 298 hirsuta, 295 glabrata, 295 ovata, 296 pycnocarpa, 296 holboellii, 296 fendleri pinetorum, 296 retrofracta, 296 secunda, 296 interposita kochii, 296 latifolia, 296 lemmonii, 296 lemmonii, 297 lignifera, 297 lignipes, 296 impar, 299 longirostris, 352 lyallii, 297 lyallii, 297 macdougalii, 296 tnacella, 298 macounii, 298 macrocarpa microphylla, 297 macounii, 298 microphylla, 298 multiceps, 297 nemophila, 292 nevadensis, 29^ Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 361 nuttallii, 298 oblanceolata, 292 oreophila, 297 oxypfiylla, 294 pendulina, 298 peramocna, 301 perelegans, 301 perennans, 298 perfoliata, 295 philonipha, 294 pinetorum, 296 pratincola, 292 puberula, 299 pulchra, 299 munciensis, 300 pallens, 300 pijcnocarpa, 295 glabra, 295 recondita, 299 reptans, 323 retrofracta, 296 rhodantha, 296 rugocarpa, 293 ritpestris, 295 sabulosa colorata, 299 /rtgida, 299 secundOy 296 selbyi, 300 setulosa, 298 shockleyi, 300 sparsiflora, 300 peramoena, 301 ifcunda, 296 sparsiflora, 301 subvillosa, 301 spathulata, 298 spatifolia, 295 stokesiae, 292 subpinnatifida beckwithii, 299 impar, 299 tenidcola, 298 tenuw, 296 thaliana, 290 Armoracia arrnoracia, 301 rusticana, 301 Athysanus pusillus, 301 giabtor, 301 Barbarea, 302 americana, 302 orthoceras, 302 dolichocarpa, 302 Berteroa incana, 302 Brachylobus hispidus, 348 Brassica a/ba, 303 arucnsis, 304 campestris, 303 hirta, 303 juncea, 303 kaber, 304 napus, 303 nigra, 304 orientalis, 311 rapa, 303 Bursa bursa-pastoris, 305 pastoris, 305 Camelina microcarpa, 304 Campe orthoceras, 302 Capsella bursa-pastoris, 305 rubella, 305 thracica, 305 Cardamine breweri, 305 cordifolia, 306 cordifolia, 306 piibescens, 306 flexuosa, 306 pensylvanica, 306 hirsuta, 306 pensylvanica, 306 infausta, 306 palustris, 347 jonesii, 347 parviflora, 306 arenicola, 306 pensylvanica, 306 uintahensis, 306 vallicola, 305 Cardaria chalepensis, 307 draba, 307 chalepensis, 307 pubescens, 307 elongata, 307 repens, 307 Cartiera -. cordata, 353 Caulanthus cooperi, 308 crassicaulis, 308 crassicaulis, 309 glaber, 309 major, 309 divaricatus, 355 giaber, 309 hastatus, 310 lasiophyllus, 309 utahensis, 309 major, 309 pilosus, 309 procerus, 309 senilis, 309 Cleome pinnata, 351 Cheiranthus argillosus, 326 asperrimus, 326 bakeri, 326 capitatus, 326 cheiranthoides, 326 efatus, 326 tnconsptcuus, 326 niua/is, 326 amoenus, 326 Cheirinia amoena, 326 cheiranthoides, 326 e/ata, 326 inconspicua, 326 repanda, 326 syrticola, 326 wheeleri, 326 Chlorocrambe hastatus, 310 Chorispermum tenellum, 310 Chorispora tenella, 310 Cleome pinnata, 352 Clypeola alyssioides, 289 maritima, 342 minor, 289 Cochlearia armoracia, 301 draba, 307 Conringia orientalis, 311 Coulterina newberryi, 345 Descurainia andrenarum, 313 brachycarpa, 313 nelsonii, 313 califomica, 311 canescens, 313 halictorum, 313 andrenarum, 313 osmiarum, 313 tnctsa, 314 leptophylla, 314 intermedia, 312 longipedicellata glandulosa, 312 pinnata, 312 filipes, 313 glabra, 313 halictorum, 313 intermedia, 313 nelsonii, 313 osmiarum, 313 paysonii, 313 richardsonii, 313 brevipes, 314 incisa, 314 macrosperma, 314 procera, 314 sonnei, 314 362 Great Basin Naturalist Vol. 37, No. 3 viscosa, 314 rydbergii, 312, 314 egjandulosa, 312 serrata, 314 Sophia, 314 Diplotaxis muralis, 314 Dithyrea wislizenii, 315 Draba ammophila, 319 andina, 322 apiculata, 320 dcwiesiae, 320 asprella, 318 asprella, 318 zionensis, 318 aurea, 318 aureifomiis, 318 decumbens, 318 luteola, 318 aureifomiis, 318 leiocarpa, 318 brachystylis, 318 caeruleomontana, 320 piperi, 320 cana, 321 cawliniana, 323 stellifera, 323 chrysantha, 319 crassa, 319 coloradoensis, 323 crassa, 319 crassifolia, 319 parryi, 319 cuneifolia, 319 cuneifolia, 319 M/eri, 319 leiocarpa, 319 platycarpa, 319 decumbens, 318 deflexa, 324 densifolia, 320 daviesiae, 320 densifolia, 320 nelsonii, 320 dictyota, 322 fladnizensis, 320 glacialis, 320 pectinata, 320 globosa, 320 sphaerula, 320 /jg/feri, 319 incerta, 320 laevicapsula, 320 laevicapsula, 320 lanceolata, 321 lonchocarpa, 321 exigua, 321 dayscarpa, 321 lonchocarpa, 321 semitonsa, 321 cesrita, 321 /utpofa, 318 maguirei, 321 burkeri, 322 maguirei, 322 nmccallae, 318 micrantha, 323 montana, 322 mulfordae, 320 nelsonii, 320 nemorosa, 322 stenoloba, 324 n/hda, 324 nana, 324 praelonga, 324 nivalis, 321 elongata, 321 exigua, 321 oligosperma, 322 andina, 322 leiocarpa, 322 microcarpa, 322 oligosperma, 322 pectinipila, 322 saximontana, 322 oriebata, 324 pattersonii, 320 dasycarpa, 320 hirticaulis, 320 parryi, 319 pectinata, 320 pectinipila, 322 platycarpa, 320 rectifructa, 322 reptans, 323 micrantha, 323 reptans, 323 stellifera, 323 saximontana, 322 sobolifera, 323 uncinalis, 323 spectabilis, 323 glabrescens, 323 spectabilis, 324 sphaerula, 320 stenoloba, 324 subalpina, 324 surculifera, 318 ufoer, 318 uncinalis, 323 ua/tda, 321 ventosa, 324 verna, 324 viperensis, 320 cjonensts, 318 Erysimum aridum, 326 asperrimum, 326 asperum, 325 amoenum, 326 inconspicuum, 326 purshii, 326 fta^en, 326 capitatum, 326 argillosum, 326 cheiranthoides, 326 elatum, 326 glaberrimum, 349 inconspicuum, 326 nuttallii, 298 oblanceolatum, 326 officinale, 350 parviflorum, 326 pinnatum, 312 puberulum, 299 repandum, 326 syrticolum, 326 wheeleri, 326 Euclidium syriacum, 327 Euklisia cordata, 353 longirostris, 352 Glaucocarpum suffrutescens, 327 Guillenia cooperi, 308 lasiophylla, 309 Halimolobos virgata, 327 Hesperis matronalis, 328 virgata, 327 Heterothrix micrantha, 355 Hutchinsia calycina, 351 americana, 351 procumbens, 328 Hymenolobus divaricatus, 328 erectus, 328 Hymenophysa pubescens, 307 Isatis tinctoria, 329 Lepidium albiflorum, 333 alyssioides jonesii, 333 stenocarpum, 333 bamebyanum, 330 bourgeauanum, 331 brachybotryum, 333 campestre, 330 chalepense, 307 corymbosum, 333 crandallii, 333 crenatum, 334 densiflorum, 330 bourgeauanum, 331 densiflonnn, 331 pubicaule, 331 pubicarpum, 331 ramosum, 331 dictyotum, 331 macrocarpum, 331 Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 363 draba, 307 chalepense, 307 fremontii, 331 georginum, 332 glaucum, 335 heterophyllum, 333 hirsiitum, 335 idahoense, 335 integrifolium, 332 heterophyllum, 333 intermedium, 335 pubescens, 334 jonesii, 333 lasiocarpum, 332 georginum, 332 lasiocarpum, 332 lasiaphyllum. 332 palmeri, 332 lasiophyllum, 332 medium, 335 montanum, 332 alpinum, 333 alyssioides, 334 canescens, 333 demissum, 330 eastwoodiae, 334 heterophyllum, 333 integrifolium, 332 jonesii, 333 montanum, 333 neeseae, 334 spathulatum, 334 stellae, 334 stenocarpum, 333 neglectum, 331 oxycarpum, 334 strictum, 334 palmeri, 332 perfoliatum, 334 philonitrum, 333 procumbens, 328 pubescens, 334 pubicarpum, 331 ramosum, 331 repens, 307 reticulatum, 334 scopuloTum, 333 nanum, 333 spathulatum, 334 spathulatum, 334 strictum, 334 tortum, 333 utahense, 332 utaviense, 333 vaseyanum, 334 virginicum, 334 medium, 335 pubescens, 334, 335 virginicum, 334, 335 ;:ionts, 332 Lesquerella, 335 alpina, 337 alpina, 337 condensata, 337 intermedia, 339 faeuis, 337 parvula, 337 spatulata, 337 argentea, 339 arizonica, 337 nudicaulis, 337 barnebyi, 339 condensata, 337 iaeuis, 337 curvipes, 337 fendleri, 338 foliacea, 338 garrettii, 338 gordonii, 341 sessilis, 341 hemiphysaria, 338 hemiphysaria, 338 lucens, 338 hitchcockii rubicundula, 340 tumulosa, 341 intermedia, 339 kingii, 339 cordiformis, 339 lati folia, 339 parvifolia, 339 latifolia, 339 ludovicinana, 339 multiceps, 339 occidentalis, 340 cinerascens, 340 parvifolia, 339 palmeri, 341 parvula, 337 praecox, 338 prostrata, 340 rectipes, 340 rubricundula, 340 spatulata, 337 stenophylla, 338 subumbellata, 341 " tenella, 341 tumulosa, 341 utahensis, 341 wardii, 342 Lobularia maritima, 342 Lunaria annua, 342 Malcolmia africana, 343 Myagrum argenteum, 339 Nasturtium amioracia, 301 austriacum. Ml hispidum, 348 linifolium, 349 palustre, 348 glabrum, 348 hispidum, 348 pumilum, 349 obtusum, 347 alpinum, 347 sphaerocarphum, 348 officinale, 343 pumt/um sinuatiwi, 348 sphaerocarpum, 348 iVorta altissima, 350 irio, 350 Pachypodium integrifolium, 357 sagittatum, 358 Parrya platycarpa, 343 rydbergii, 343 Pennellia micrantha, 355 Physaria acutifolia, 344 acutifolia, 345 purpurea, 345 australis, 345 chambersii, 345 membranacea, 345 didymocarpa, 345 australis, 345 newberryi, 345 grahamii, 345 newberryi, 345 Physolepidium repens, 307 Pilosella stenocarpa, 327 virgata, 327 Pleurophragma gracilipes, 357 integrifolium, 357 lilacinum, 357 platypodium, 357 rhomboideum, 356 alpina, 347 armoracia, 301 austriaca, 347 curvipes, 347 g/afora, 348 hispida, 348 integra, 347 ofcfusa, 348 sphaerocarpa, 348 palustris, 348 hispida, 348 sinuata, 348 sphaerocarpa, 348 tenerrima, 349 underwoodii, 347 Raphanus raphanistrum, 346 sativus, 346 tenellus, 310 Rorippa 364 Great Basin Naturalist Vol. 37, No. 3 armoracia, 301 glandulosa, 312 longirostris, 352 alpina, 347 murale, 314 Streptanthus austriaca, 347 nasturtium-aquaticum, 343 angustifolius, 294 curvipes, 347 officinale, 350 cordatus, 353 alpina, 347 palustre, 348 crassicaulis, 309 curvipes, 347 pinnatum, 312 crassifolia, 353 integra, 347 procerum, 314 longirostris, 352 curvisiliqua, 347 pygmaeum, 349 ptfosus, 309 integra, 347 richardsonii, 314 sagittatus, 358 islandica, 348 sop/iia, 314 virgatus, 296 glabra, 348 virgatum, 327 t/;i/omingensts, 355 hispida, 348 uiscosum, 314 Thelypodiopsis nasturtium-aquaticum, 343 Smelowskia ambigua, 354 obtusa, 347 americana, 351 argillacea, 354 alpina, 347 calif ornica, 311 aurea, 354 integra, 347 calycina, 351 divaricata, 354 sphaerocarpa, 348 americana, 351 elegans, 355 palustris, 348 lineariloba, 351 nuttallii, 358 gUihra, 348 utrescens, 351 torulosa, 358 hispida, 348 Sophia If i/orntngensis, 355 sinuata, 348 andrenarum, 313 Thelypodium sphaerocarpa, 348 osmiarum, 313 a//tne, 356 tenerrima, 349 brevipes, 314 ambiguum, 354 underwoodii, 347 /t/ipes, 312 amplifolium, 358 Schoencrambe glabra, 313 aureum, 354 decumbens, 349 halictorum, 313 coopert, 308 linifolia, 349 incisa, 314 crenatum, 334 pinnata, 349 intermedia, 312 elegans, 355 pinnata, 349 leptophylh, 314 gracilipes, 357 pygmaea, 349 nelsonii, 313 integrifolium, 356 Sinapsis parviflora, 314 affine, 356 a/fea, 303 pinnata, 312 complanatum, 356 aruensis, 304 procera, 314 gracilipes, 357 /tmtea, 303 purpurascens, 314 integrifolium, 357 fcafoer, 304 richardsoniana, 313 laciniatum, 357 nigra, 304 serrato, 314 millefhrum, 357 Sisymbrium sonnet, 314 lilacinum, 357 altissimum, 350 sop/ita, 314 subumbelkitum, 357 palustre, 347 viscosa, 314 lasiophyllum, 309 amphibium, 348 Stanleya utahense, 309 paZusfre, 348 albescens, 351 laxiflorum, 357 aureum, 354 arcuata, 352 longirostris, 352 canescens, 313 canescens, 352 macropetalum, 358 brevipes, 314 co//tna, 352 micranthum ma/or, 313 fruticosa, 352 milleflorum, 357 decumbens, 349 glauca, 352 nuttallii, 358 elegans, 355 iari/oZta, 351 ovalifolium, 358 glandifera, 312 heterophylla, 352 pa/meri, 358 graci/is, 312 integrifolia, 351 rhomboideum, 356 halichirum, 313 pinnata, 352 gracilipes, 357 hispidum, 348 integrifolia, 351 rollinsii, 357 jncisum, 312, 314 gibberosa, 352 sagittatum, 357 /i/ipes, 312 inyoensis, 352 ovalifolium, 358 sonnet, 314 pinnata, 352 sagittatum, 358 intermedium, 313 pinnatifida, 352 vermicularis, 358 irio, 350 integrifolia, 351 suffrutescens, 327 isZandicum, 348 viridiflora, 352 torulosum, 358 lasiophyllum, 309 Stanleyella utahense, 309 leptophyllum, 314 tvrightii, 358 wrightii, 358 linifolium, 349 fpne/^, 357 tenellum, 357 decumbens, 349 Sfenop/iragma Thlaspi pinnatum, 349 tirgafum, 327 alpestre, 359 bngipedicellata, 312 Streptanthella glaucum, 359 Sept. 1977 Welsh, Reveal: Utah Flora, Brassicaceae 365 purpurascens, 359 arvense, 359 hursa-pastoris, 305 campestre, 330 cochleariforrne, 359 cohradoense, 359 fendleri color adoense, 359 hesperium, 359 tenitipes, 359 glaucum, 359 hesperium, 359 pedunculatum, 359 hesperium, 359 montanum, 359 fendleri, 359 montanum, 359 nuttallii, 359 purpurascens, 359 Thysanocarpus curvipes, 359 eradiatus, 359 pusillus, 301 Turritis brachycarpa, 292 drummondii, 294 glabra, 295 hirsute, 295 lasiophylla, 309 macrocarpa, 295 ouata, 296 retrofracta, 296 spathulata, 295 sfricta, 294 Vesicaria alpina, 337 fendleri, 338 stenophylla, 338 TYPHLODROMUS MCGREGORI CHANT (ACARINA: PHYTOSEIIDAE) AND ITS CONTROL OF PHYTOPHAGOUS MITES IN UTAH AND SOUTHERN CALIFORNIA APPLE ORCHARDS' B. A. Croft- and C. D. Jorgensen' Abstract.— A review of published and unpublished data for the predatory mite Typhlodromits mcgregori in Utah and southern California apple growing areas is presented to summarize its role. Typhlodromiis mcgregori was not found resistant to pesticides and, although usually present with Bryobia rubrioculus, only when Acultis schlech- tendali was also present was there significant predation on B. rubrioculus. Aculus schlechtendali was considered the primary host and B. rubrioculus secondary for T. mcgregori. Typhlodromus occidentalis was always found as- sociated with Tetranychus spp. and often with Eotetranychus and Pronematus ubiquitus in unsprayed orchards. Certain phytoseiid mites are specialized predators of phytophagous mites and are capable of controlling or suppressing their populations at low densities (Collyer 1964, Hoyt 1969, Croft and Barnes 1971). This potential for biological control has prompt- ed studies of the native species in many ag- ricultural areas of the world (Huffaker et al. 1969), and, although predator-prey relation- ships vary in different geographic regions, similar associations are not uncommon in areas where ecological conditions are sim- ilar. During 1967-70, Jorgensen and co-work- ers (Jorgensen 1968, Nelson and Jorgensen 1968, Croft and Jorgensen 1969, Leetham and Jorgensen 1971, and Duke et al. 1970) investigated predator-phytophagous mite relationships in Utah apple orchards. They reported Typhlodromus mcgregori Chant and T. occidentalis Nesbitt in predatory associ- ation with Bryobia rubrioculus Scheuten and Tetranychus mcdanieli McGregor, re- spectively. Croft (1970) found predator-prey relation- ships in mountain apple orchards of south- ern California (Oak Glen, near Yucaipa, Ca- lif.) almost identical to those reported in Utah. Because of this and similar native vegetation, he concluded that the Oak Glen environment was an extension of the eco- logical type reported for Utah, and atypical of other apple areas in California. His con- clusions were also influenced by the com- mon occurrence of Tet. mcdanieli through- out Utah and at Oak Glen. This species is not reported from other apple areas of Cali- fornia (Barnes and Madsen 1961). These similarities allow for the summary of both areas in this single analysis and should pro- vide a general understanding of the role oc- cupied by T. mcgregori. Although different techniques were used, both Jorgensen's group and Croft studied the ecology, bionomics, and control effec- tiveness of T. mcgregori. This paper presents additional data and a summary of the pre- dator-prey relationships, along with the ap- parent role of T. mcgregori in both areas. Although T. occidentalis was an effective natural enemy of Tet. mcdanieli in both re- gions, references to this predator are limited to comparisons of predator effectiveness or possible niche overlaps with T. mcgregori and their respective relationships to the to- tal ecology of mites in the orchards studied. General Methods Investigations in southern California were conducted at Oak Glen, an apple growing area ca 12.9 km north of Yucaipa, Califor- nia, at the eastern edge of the Los Angeles basin, and included about 162 ha of com- Piiblished as Journal Article No. 8027 of the Michigan State Agricultural, Experin ■Department of Entomology. Michigan State University, East Lansing 48823. Department of Zoology, Brigham Y'oung University, Provo, Utah 84602. .366 Sept. 1977 Croft, Jorgensen: Mite Predation 367 mercial apples grown between 1341-1463 m in the San Bernardino Mountains. Pre- dominant apple varieties in both areas in- clude Delicious, Rome Beauty, and Jona- than. Vegetation types in the southern California study area were typical of those in valleys of more northern latitudes in Utah, i.e., Acer negundo. Primus virginiana, Pinus ponderosa, and Symphorocarpiis albiis. Studies on the effects of pesticide treat- ments on, and the biologies of, certain pest species had previously been conducted in the Oak Glen orchards (Barnes and Madsen 1961). During the summers of 1968-70, sam- pling studies emphasizing both pedator and prey mite relationships were made. Seven unsprayed, neglected orchards and 11 sprayed orchards, all of the Delicious varie- ty, were sampled at two-week intervals from 13 June to 24 Oct 1968. Spur samples (50 spurs/ tree) with leaves were processed using Berlese extraction methods and mites mounted for identification. During the summers of 1969 and 1970 a series of orchards, including one which had not been sprayed for 10 years, three which had not been sprayed for 3-5 years, and one under a commercial chemical control pro- gram, were sampled at two-week intervals to determine population trends of both pest and predatory mites. In samples from five orchards, 20 leaves were collected from five locations of the four compass zones in each of five trees (80 leaves/tree) during the summer. All population counts were made by direct observation through a binocular microscope or from plates of the mite- brushing machine. In addition to the field studies in Califor- nia, evaluations of the oviposition rates and developmental times for T. mcgregori from egg to adult, when reared on several erio- phyid rust mite species, were made in the laboratory. Oviposition rates/female pre- dator for an eight-day period (replicated 10 times ea) when confined on excised apple, peach, or pear leaves containing popu- lations of Aculiis schlechtendali (Nalepa), A. coronutus (Banks), or Eriophyes pyri (Pgst.), respectively, at densities of ca 200 rust mites per leaf were determined. Devel- opmental rates for T. mcgregori, using four replications with 10 females/ replicated leaf when feeding only on A. schlechtendali at densities of 200/ leaf, were also measured. In all experiments the temperatures were24 ± 2 C and the RH 30-60 percent. Li Utah, two types of studies were con- ducted. First, an extensive examination of mite distributions and species associations was made (Jorgensen 1968, Nelson and Jor- gensen 1968, Leatham and Jorgensen 1971, Duke et al. 1970); however these were only correlative comparisons, and no experiments were done to measure specific associations between the various predator and prey mite populations. Attention was not given to the possibility that rust mites were preferred by T. mcgregori. Second, the life history of T. mcgregori was studied by Croft and Jorgen- sen (1969), using B. rubrioculus, Tet. mcda- nieli, and Panonychus ulmi Koch as experi- mental food bases. Again, rust mites were not used as food in these experiments. Results and Discussion Mite Fauna at Oak Glen Mite species collected commonly from unsprayed Oak Glen apple orchards were: Tetranychidae— Tef. mcdanieli. B. rubri- oculus, and Eotetranychus uncatus Garman; Phytoseiidae— T. mcgregori, T. occidentalis, and T. pomi (Parrott); and Tydeidae— Prone- matus ubiqidtus (McGregor); Eriophyidae— A. schlechtendali; and Erythraeidae— Be- laustium sp. Tetranychus mcdanieli was col- lected predominantly from commercial or- chards which were intensively sprayed with pesticides. Barnes and Madsen (1961) noted the absence of P. ulmi from Oak Glen, an observation that was supported during our work. Unsprayed for Ten Years.— Population levels of phytophagous and predatory mites were extremely variable among trees in this orchard during 1969 and 1970, and for this reason separate plots for two different tree groupings are reported. In three trees dur- ing 1969 (Fig. lA), B. rubrioculus attained high population levels of ca 40/leaf while only very low populations of T. mcgregori developed in late season. In 1970, mites in these same trees showed similar population 368 Great Basin Naturalist Vol. 37, No. 3 trends for these two species; however, brown mites only increased to ca 15/leaf while T. mcgregori was more abundant and reached 0.8/leaf in August (Fig. IB). During 1969 (Fig. IC) and 1970 (Fig. ID), in two other trees in this orchard, pop- ulations of A. schlechtendali were present in addition to B. rubrioculus and T. mcgregori. In these instances, B. rubrioculus attained lower peak levels and T. mcgregori exhibited a more marked numerical response as com- pared with the situation when A. schlech- tendali was not present (i.e., compare data of lA and B with IC and D). Although Tet. mcdanieli was occasionally found in un- sprayed blocks, it was not collected in this orchard, nor was T. occidentalis. Unsprayed from Three to Five Years.— Figures 2A and 2B show the mean densities of Tet. mcdanieli-Eotetramjchus uncatus (collectively referred to as Tet. mcdanieli in Figures 2A and 2B because E. uncatus was only present at low densities) and B. rubr- ioculus populations in two apple orchards for the 1969 season. These species were sep- arated temporally; B. rubrioculus, with an early season distribution, reached its peak density in July, whereas the Tet. mcdanieli- E. uncatus populations began to increase in July and peaked in late August or early 40 - 30 - 20 10 5 —^^•^y V- c 40 30 L 20 o 10 A 5 • ^^ 1 ,^-'' / • Fig. 1. Densities of plant-feeding and predaceous mites in an apple orchard unsprayed for 10 years at Oak Glen, California: pooled values for three trees during the 1969 (A) and 1970 seasons (B); pooled values for two other trees during the 1969 (C) and 1970 (D) season. Sept. 1977 Croft, Jorgensen: Mite Predation September. The apple nist mite, A. schlech- tendali, was present in the trees, but ahnost at undetectable levels. Although both T. mcgregori and T. occidentalis occurred in these orchards, T. mcgregori did not respond markedly to increasing densities of any tet- ranychid species, whereas T. occidentalis populations appeared to increase with in- creases in the densities of Tet. mcdanieli-E. uncatus populations and eventually over- came their populations before they exceed- ed three to five mites per leaf. In the other orchard unsprayed for three to five years, B. nibrioctilus occurred only at very low densities (2 mites/ leaf) and T. mcgregori was seldom found (Fig. 2C). Rust mites were only occasionally observed in this block (not plotted in Fig. 2C). In Fig- ure 2C the tetranychid mites, Tet. mcdanieli and E. uncatus began increasing in late July and early August, with an apparent numer- ical response by T. occidentalis. Predator populations increased to nearly 3 /leaf be- fore declining to a lower density as prey became scarce. The tydeid, P. ubiquitus, in- creased to a density of 0.6/ leaf during the period when the populations of tetranychids were greatest. The presence of both tet- ranychids and tydeids appeared to provide prey for the observed numerical increase in 40 A 30 ■ 20 10 ■S 1 / • / ' ^t* 'l-tij /\ B \ 40 C 30 20 10 5 1 M A S • aB. rubrioculus a AJ. mcdanieli ■ •'>>.«■ P.ublquitus A A Aschlechtendal O oT. mcgregori O o T. occidentalis Fig. 2. Densities of plant-feeding and predaceous mites in four apple orchards unsprayed for 3-5 years at Oak Glen, California, during 1969 (A, B, and C) and 1970 (D): represents pooled values for 5 trees/orchard. 370 Great Basin Naturalist Vol. 37, No. 3 the predator populations, as T. occidentalis were frequently observed feeding on the ty- deids. In Figure 2D, P. ubiquitiis populations attained 2.6/ leaf and were the major prey supporting the populations of T. occiden- talis, which increased to 0.8/ leaf on 19 Sep- tember. In this orchard, Tet. mcdanieli-E. uncatus populations did not exceed 0.3/leaf during the entire season and virtually no B. rubriocuhis were observed. Commercially Managed Orchards.— Fig- ures 3A and 3B illustrate the population densities of Tet. mcdanieli throughout the summers of 1969 and 1970, in a sprayed or- chard without any phytoseiid mites. Tetra- nychus mcdanieli exceeded 100/leaf in total orchard samples and, on occasion, exceeded 1,000 mites on individual leaves. If miticide sprays were not applied, intraspecific com- petition for food finally limited their in- creases, but only after severe damage to the apple foliage had been sustained. During the summers of 1968 and 1969, over 200 samples (50 leaf spurs/sample) were taken from 19 other sprayed apple or- chards. Less than 50 phytoseiid mites were collected from the foliage of apple trees at these sites, approximately equal numbers of T. mcgregori and T. occidentalis. Most of these predators were taken from peripheral trees which were near native, unsprayed vegetation. Predators were encountered only late in the growing season after pesti- cide residues were dissipated. Although the selection of sites and the presentation of data in Figures 1-3 over- simplify the ecological differences between sprayed and unsprayed orchards at Oak Glen, California, the following species as- sessments were concluded from these data: Bryohia nibriocidus was collected only in unsprayed orchards and was most abimdant during the early summer. Acidns schlechten- dali occurred at low levels in unsprayed plantings and varied greatly among loca- tions from year to year. It was also ob- served at low densities in commercial or- chards. Unsprayed orchards were never observed where A. schlechtendali occurred without B. rubriocuhis, although some or- chards supported B. rubriocuhis only. Tetra- nychiis mcdanieli was found most com- monly in sprayed apple orchards in late summer when populations attained extreme- ly high densities and caused serious damage to apple foliage in the absence of sufficient phytoseiid mites or adequate chemical con- trol. This species and low densities of E. uncatus were also found in imsprayed trees where the tydeid mite, P. ubiquitus, was sometimes observed. Typhlodromus mcgregori was always found in imsprayed orchards where B. rubr- 100 50 30 10 5 B ,A A AT. mcdanieli ; / / J \ / \ 1 A / \ 1 1 \ 1 / / ■ /'' M M Fig. 3. Densities of Tetranychtis mcdanieli in a sprayed commercial apple orchard at Oak Glen, California, during 1969 (A) and 1970 (B); represents pooled values for 5 trees. Sept. 1977 Croft, Jorgensen: Mite Predation 371 ioculus occurred. The predators appeared to prey on this species, but they did not sup- press its densities significantly. When A. schlechtendali was present with B. rubrioc- ultts, T. mcgregori rapidly increased in re- sponse to the rust mites and seemed to sup- press B. rubrioctdiis populations to a greater extent than when rust mites were absent. With the exception noted above, T. occiden- talis was collected almost exclusively from unsprayed orchards where it was most closely associated with population com- plexes of Tet. mcdanieli, and E. uncatus, which were effectively maintained at low levels by the predator. Pronematus uhiqiii- tus also appeared to be an adequate alter- nate host for T. occidentalis in several un- sprayed orchards. Although Hoyt (1969) and Burrell and McCormick (1964) reported A. schlechtendali as a favorable prey species for T. occidentalis, the predator was never found in orchards at Oak Glen where only Rist mites or B. rubriocidus were present. Only in one case in Utah was T. occiden- talis found associated with B. rubrioculus and A. schlechtendali, and it was in a mixed block of apples, plums, and peaches along with high populations of rust mites and Eo- tetramjchus carpini borealis Pritchard and Baker. We concluded from these data, as did Burrell and McCormick (1964), that B. rubrioculus probably is not an acceptable prey for T. occidentalis. These observations and conclusions in re- lation to the bionomics of these species fol- low closely with those of Croft and Jorgen- sen (1969) and Duke et al. (1970), who reported the relationship of T. mcgregori in close association with B. rubrioculus in un- sprayed apple orchards in Utah; and yet T. mcgregori did not exert any significant con- trol of the prey. The association of T. occi- dentalis with Tet. mcdanieli and/or E. un- catus or closely related species was reported earlier by Hoyt (1969) for T. occidcntalis- Tet. mcdanieli, Schuster and Pritchard (1963) for T. occidentalis-Tet. pacificus McGregor or Eotetranychus willamettei (McGregor), and Flaherty (1967) for T. occi- dentalis-T. pacificus or E. williamettei. In no instance, has T. occidentalis been report- ed as a predator of B. rubrioculus, yet this predator occurs in many geographical re- gions where B. rubrioculus is found. One additional result which provides in- sight into the role of T. mcgregori as a pre- dator of orchard pests relates to its relative- ly high oviposition rate /female /day when feeding on each of the three eriophyids: A. schlechtendali = 2.0 ± 0.2, A. coronutus = 1.9 ± 0.3, and E. pijri = 2.0 ± 0.1. The greatest average egg production observed for a 24-hour period was 2.3, which prob- ably is close to the maximum for this spe- cies. Mean developmental time (egg-egg) for four replications of 10 T. mcgregori/ replication feeding on A. schlechtendali were 8.5, 8.7, 8.0, and 8.5 days for an over- all mean of 8.42 ± 0.3 hrs. The role of Typhlodromus mcgregori: a reevaluation Typhlodromus mcgregori occurs through- out North America north of Mexico and has been collected from several native plants in addition to walnut, grape, fig, apple, and citrus (Schuster and Pritchard 1963, Duke et al. 1970). Reported prey for this pedator includes Aegijptobia nothus P. and B., Tet. pacificus, Eoteteranijchus boudreauxi P. and B., Eriophyes vites Pgst., E. williamettei and B. rubrioculus (Poe and Enns 1969, Schuster and Pritchard 1963, Duke et al. 1970). Croft and Jorgensen (1969) studied the life history of T. mcgregori while it fed on B. rubrioculus, P. ulmi and T. urticae and observed that it preferred B. rubrioculus as compared to P. ulmi and T. urticae. They concluded that T. mcgregori did not assist substantially in controlling the latter two pests which occurred primarily in sprayed apple orchards in Utah. Even when feeding on B. rubriocidus, which T. mcgregori seem- ed most closely associated with in un- sprayed orchards, the rate of development and oviposition rates were considerably lower than reported values for most other phytoseiid species which are efficient pre- dators of tetranychid mites (McMurtry et al. 1970). Duke et al. (1970) examined the distribu- tion of T. mcgregori: (1) among apple or- chards in Utah, (2) within apple orchards, 3' (iuKAi' IVvsiN N.\riiH.\i,isr Vol. 37, No. 3 [l\) within appU" tri"i"s, ;uk1 [■{) upon appU" U'.iNt's. riu'N also roportoil tlu> .soatvhinu; he- ha\ ior ot /'. inr^if^i>h on appir loavo.s. Vic- ilati>is \Noiv loiuul primipalh in nnspiaNcnl onliauls, at;nt nonnni lonnK npon U\i\i\s, and thi"\ st-anhfil liM\i>s ni a ri'sliiilivo niannor. In all la.scs. (h<" jModatoi (.lisliihulion and .soarc-hinu; hv- ha\ ioi laiK'd to ilosfh roiii'spi>nd with niOMMUiMits and disti ilnilions ol />. nihiiih-- tiltis. Nelson and |ori;i«nsiMi y h)(iS^ tniihiM stnd- \i\\ [\\c disporsal anil ilistrihnlional associ- ations hi'twoiMi /'. iiu-^if^i>n and B. nthriiH- tiltis anil tomul litlli- or no lorndation lu'twoiMi pmlators and pro\ on spnis and U'a\i"s limine, iho rntiio Nrar or o\im a sini;lr 2 l-honr poiiod. Hn/ohid nihiiiHiihis was cxliiMnrh n\ohilr and piosont o\ im a >;M'aliM portion ol tlu" a[>plo Ivcv than was To sninuKiri/i", Ioru,iMisiMi and lo workiMs listed throe taetois which eontiilniteil to the i>hser\ed inability ot 7". mcpr^^ori to appre- eiabh' atteet or I'ontrol populations of />. nihriociilus in the field: ^l"! /". DU'^rr^ori was liniiteil in its reprotlnetive and ilesttiieti\ e ability when eonipaieil to the reprodneti\e potential ot" B. nihrioculus [Croh and ]or- i;ensen 1970); ['!) l\ nu-^ir^ori had limited [n>tential tor eontiolling B. nthhoculus be- eanse ot ditteienees in respeetive dislribn- tii>ns in orehards, trees, and lea\es and the predators' restricted searching beha\ iiM vHnke et al. 1970^ and ^3) T. mr^ir^oh had dissimilar dispers;d habits and spring distri- bntions as compared to />. nthrioctihis, which was more active anil present o\er a greater portion ot foliage and wood) areas ot apple trees ^^Nelson and jorgensen UHSSl (Considering the three factors suggested In jorgensen and co-workers and in light of the data presented herein, w c> ean now ree\alnato the role of /'. ;/j(gr8) pointed ont that the spa- tial occnrrence (^distribution and dispersal mo\'ement) of 7'. inc^ir^ori was onU' looselv associateil with that of B. nthrioculus. If the preilalor-[)re\ as,sociation was primar\, this incongrnity wonid be an important factor in limiting effective predation. \V'e now qnes- tion the a.ssnmption of primar\- a.s.sociation altogether. It is ajiparent that neither the I'tah nor sonthern C'alifornia .strain of /'. ;;i(gng()n are [Mimarib adapted to the spider mites i^e.g. /'. ulmi, Tct. itiiicac, Tct. incdiinicli) which most connnonlv occnr in sprayed or- chards. I'eeding stndics b\ Caoft and ]or- gen.sen ^19(i9') suggested a lack of a.ssoci- ation with /'. ulnti or 7"«7. tirticoc for the I'tah strain and popnlation studies in both areas indicated little or no association with l\t. lucihniiili. .Mso, colonies of 7'. //ug/c- g(>r( i.\o poorl\ in the laboratory when reared on l\t. pacifinis (B. .\. (aoft, nn- pnblished ilatal The relationship between 7". luc^ir^oii and B. nthrioculus is nnilear. since both species were consistent 1\ fonnil together in nnspruved orcharils dnring the growing sea- son. It is not known whether this is a pri- mar\ association and B. ruhriocuhis is the preferred pre\ , oi a sc>i'ondar\' a.ssociation allowing for the maintenance of 7'. /ncgrr- g(>n' populations at low levels, or an as.soci- ation which de\elops alter [McdatiMS re- spond [o other speiies i^g-. A- srhliThti-tuUili). The latter question seems most likeK- considering the reprodnctive rates reported herein for 7'. //Kgngcn when feeding on three rnst mite species, inclnding -A. schli-chtcmiali. Caoft and Jorgensen ^^19(S9) reported 11.5 days (egg-egg) for do- Sept. 1977 ( juii'i, Joi<(,i;nsi;n: Mrii; I'ummiion 373 V(;lf)|)rn(;nlal timcj and 1.2 <;gj^s/(]ay/f(;irial<; lor 7'. }nehavior, and biology of H. ruhrioculus may account ff>r this immu- nity, inclu(Jing the large si/e of tli<; a(Jult mites (.McMurtry et al. 1970; and their higfily mobile behavior, particularly the a(Jults which spen(J a large portion of their time on the woo(Jy parts of \\i<: plant, where they lay many '-ggs (.\c-ls(jn and jor- gensen 1968;. Tijphlodrornus rncs^raiuni was sfdd(;m foun(J on sprayefJ plantings in eitfier study area, altfiough A. schUichleruldli occurred in both unspraylc orcliards of Utah or south(;rn Califor ni;i ;iiif| its appli(;ation undrjubt(;dly contrib ul<;s to the exclusion u\ I', tncijrciuiri from sprayed orchards in b(;th areas. l-ni;j|>l<; in Califrjrnia: Univ. Califf>rnia CJirc. .'>()2, raiifornia CJorp, Kxl. S<;r. 'J I pp. liimtKi.i,, H. W,, ANi> W. J. .McCoiiMK K, Vm. Tijphh- drornus and Amhlyfieiw) fArarina; I'hylosciidac; as predators on orctiard miles. Arm iMilomol Sdr(nnuH occidanUiliH ,\esbitt ^Acarina: Pfiytoseiidaej. \'\\.\). Diss., (Jniv. fjalifornia, liiv- erside. 92 pp. (au)KI, B. a., a.so .M. M, IUmnks. I'j7). f,om(>arative studies of four strains of TtjjthludnmiUH t/ccUUin- UilU. III. Kvaliiations <>\ releav;s of insn. iW) f,om(>arative studies of four strains of 'lyphl/jdnmiwi Dcculan- UiHh. II. \^\>Ty toxicity of 10 compounds common to apple pest control. J. f>;on. Kntomol. 6.'i: ]r,2M\r,:i\ r;»«>Ki, li. A., ANo (, i) ]<,!'.<, y.SHV.s. ]im. Life history of TyphlodromuH rnfurefiori ^Acarina; Fhyto- vjiidae;. Ann. Kntornol. Vx;. Arn. 62: J26I-J267. I>r;KK, K. ,M., B. A. (AV)ri, ss\> C D. ]<>n<,y.SHKS. 1970. Distribution and searching behavior of Typhl/jdri/muH rru:i^rej:^ori. \. Kcon. Kntornol. 0.3:221-227. Fi.AJJKK'ry, D. 1967. The «jgy of tetranychid mites and their natural control, Ann. H«fV. Knto- mol, J4; 12.5-174. 374 Great Basin Naturalist Vol. 37, No. 3 JoRGENSEN, C. D. 1968. Ecology of Utah apple mites: a statement of the problem. Proc. Utah Acad. Sci., Arts and Letters. 44: 265-274. Leetham, J. M., AND C. D. JoRGENSEN. 1971. Over- wintering phytoseiid mites in central Utah apple orchards. Great Basin Nat. 29: 96-104. McMuRTRY, J. A., C. B. HUFFAKEB, AND M. VAN DE Vrie. 1970. Ecology of tetranychid mites and their natural enemies: A review I. Tetranychid enemies: their biological characters and impact of spray practices. Hilgardia: 40: 331-390. Nelson, E. E., and C. D. Jorgensen. 1968. Dispersal of Bryobia spp. and Tijphlodromus rncgregori Chant within apple trees in central Utah. Proc. Utah Acad. Sci., Arts and Letters. 54: 168-181. PoE, L. S., AND W. R. Enns. 1969. Predaceous mites (Acarina: Phytoseiidae) associated with Missouri orchards. Trans. Missouri Acad. Sci. 3: 69-82. Schuster, R. O., and A. E. Pritchard. 1963. Phyto- seiid mites of California. Hilgardia 34: 191-285. HOST-INDUCED MORPHOLOGICAL VARIATIONS IN THE STRIGEOID TREMATODE POSTHODIPLOSTOMUM MINIMUM (TREMATODA: DIPLOSTOMATIDAE). III. ORGANS OF ATTACHMENT James R. Palmieri' Abstract.— A variety of amphibian, reptilian, avian, and mammalian hosts were used in experimental devel- opment of Posthodiplostomum minimum. As a result of this study much host-induced morphological variation was noted in several organs of attachment. Variations in the oral sucker ranged from a well-developed muscular organ to a weakly developed oral slit. Acetabular development ranged from well-developed to reduced forms lacking apical musculature, tegumental spines, and sensory structures. The holdfast organ showed marked reduction in most poikilothermic hosts. Members of the family Diplostomatidae Poirier, 1886, constitute a group of trema- todes characterized by a distinct flat or spoon-shaped forebody containing an oral sucker, acetabulum, and bulbous tribocytic or holdfast organ. Throughout the literature, when investigators describe strigeoid trema- todes, measurements of the oral sucker, ace- tabulum, and holdfast organ are character- istically given. Little emphasis has been placed on the effects of the definitive host upon the development of these organs. A variety of amphibian, reptilian, avian, and mammalian hosts were experimentally employed in the development of adult Post- odiplostomwn minimum, a strigeoid trema- tode characteristically found in a variety of piscivorous avian hosts (Palmieri 1975). As a result of these experimental studies, great variations in the size, shape, and complexity of the oral sucker, acetabulum, and holdfast organ were recorded and are subsequently described in this paper. Material and Methods The experimental hosts employed and the techniques used in experimental infections of P. minimum to these hosts, as well as specimen preparation for examination by scanning electron microscopy, are explained in detail in part II of this .study. Results and Discussion Oral sucker.— Five variables were ana- lyzed for the oral sucker of P. miniminn. (length, width, cross sectional area, ratio of length to width, and ratio of oral sucker- acetabulum cross sectional areas) and are listed - in Table 1. Various morphological modifications of the oral sucker are com- pared according to hosts (Fig. 1-6). Of all the morphological features of the adult worm examined during this in- vestigation, the oral sucker shows the great- est amount of morphological variation. In ecologically normal definitive hosts (piscivorous birds), as well as mammalian hosts, the oral sucker is well developed, muscular, and situated in the anteromost portion of the forebody (Figs. 5-8, 14). In certain avian and mammalian hosts, how- ever, it appears to lie below the surface of the anterior end of the forebody (Figs. 3-4, 9-11, 15). In such instances, the cavity of the oral sucker appears collapsed, although the musculature retains its identity. In am- phibian and reptilian definitive hosts, an oral sucker appears to be lacking or may be so markedly reduced that only an oral slit appears (Figs. 1-2, 11-13). In cases where the oral slit is present, it does lead into the pharynx. In other than avian hosts, it is not atypical to find some specimens with well- Department of Zoology, Iowa State University, Ames, Iowa 50010. Present addn 14-Malaysia. UCICMR. tute for .Medical Research, Pahang Road, Kuala Lumpur, 02- 375 Great Basin Natvralist Vol. 37, No. 3 Figs. 1-6. Morphological \-ariations of the oral sucker of adult P. minimum from vertebrate hosts. .\11 specimens 72 hours old. >Seale: f inch = .030 mmV 1. Host: Btifo americanits, note presence of an oral slit: 2, Host: Ratui pipiens, note presence of an oral slit; 3-4. Host: Ig^taita igiwxw. note the infolding of the oral sucker: 5. Host: Co- lomha livia, note the well-developed oral sucker: 6, Host: Felis catus, note the well-developed oral sucker. Sept. 1977 Palmieki: Trkmatouk Variatkjns 377 developed oral suckers and others with oral slits, even among worms recovered from a single definitive host. Ac;etabulum.— Six variables were ana- lyzed in studying the acetabulum (length, width, cross sectional area, ratio of the length to width, acetabular index, ratio of the acetabulum length to the body length, and the acetabulum width to the body width). Length and width were measured along the major axes of the body. An analysis of the above data appears in Table 2. Morphological variations of the acetabulum are illustrated in Figures 16-21. Throughout the four classes of definitive hosts experimentally used for development of adult P. minimum, complexity in struc- ture and overall size of the acetabulum vary greatly depending on the group of defini- tive host used. Within some hosts (especially avian) the acetabulum is large, consisting of a full base well endowed with tegumental spines and an expanded muscular surface re- gion covered with two rows of presumed sensory structures. Two views of the ace- tabulum recovered from the avian host Ijirus ar^entatus clearly show the above structures (Figs. 18-19). In other hosts, the acetabulum is reduced in size (Fig. 21) and neither the base nor the apical region are fully developed (Figs. 17-25). In one speci- men of P. minimum recovered from a frog (Rana pipiens) (Fig. 16), the base of the acetabulum is not distinguishable, and there is no evidence of apical musculature, well- developed tegumental spination, or sen.sory structures. In a few abnormally developed worms recovered from Amhystoma tigri- num, the acetabulum has apparently degen- erated to a point where it is no longer pres- ent. Holdfast or tribocytic; organ.— Four variables (length, width, cross sectional area, and ratio of length to width) were analyzed in studying the holdfast organ of P. min- imum. An evaluation of the above data is presented in Table 3. Structure of the hold- fa.st organ is shown on Figures 22-27. The holdfast or tribocytic organ of P. minimum occupies a large portion of the Table 1. Statistical analysis of oral sucker measurements of P. Miniynum from experimental definitive hosts. Variable Mean of Combined Host Class Standard Deviation Amphibian Reptilian Aves Mammalia Oral Sucker Length (OSL) Oral Sucker Width (OSW) Cross-sectional Area of Oral Sucker (AOSLYOSW) Ratio of Oral Sucker Length to Width (ROSLOSW) Ratio of Oral Sucker to Ace- tabulum Cross- sectional Area (ROSXXA) 0.033 0.028 O.OlO 0.029 0.049 0.0.38 0.0.36 0.524 0.009 0.001 0.393 L552 0.026 0.025 0.031 0.029 O.OOI 0.001 0.001 0,001 1.04.5 0.946 L2.32 L244 0.684 0.388 0.429 0.371 "All measurements in mm. 378 Great Basin Naturalist Vol. 37, No. 3 posterior region of the forebody. Although its size is consistent in specimens recovered from most vertebrate hosts, it is somewhat reduced in amphibian and reptilian hosts. The relative constancy in size of the hold- fast organ in worms from all vertebrate hosts make variations in the shape of this structure difficult to ascertain. Furthermore, the shape of the holdfast is probably affect- ed by the underlying musculature. Figs. 7-12. Morphological variations of the oral sucker depicting the normally developed oral sucker and its re- duction to an oral slit. Figs. 13-15. Morphological variations of the oral sucker, vitelline gland, and testes. Sept. 1977 Palmieri: Trematode Variations 379 - ■ • ---. II ii—i '■fe. —Wf Figs. 16-21. Morphological variations of the acetabulum of adult P. minimum recovered from vertebrate hosts Ail specmiens 72 hours old. 16, Host: Ram, fupicm. note the loss of the base of the acetabulum and the reduc- ?r,a J''"'^ structures and tegumental spmes. (.Scale: 1 inch = .018 mm); 17, Host: Merione, unguiculatus; 18-19, Host: Larm argentatus, note the well-developed tegumental spines, sensory structures and muscular apical region. (Scale: 1 inch = .018 mm); 20, Host: Bufo americanus (Scale: 1 inch = .018 mm); 21, Host: Chrmemm ptcta, note the relationship of the acetabulum to the forebody and holdfast organ. (Scale- 1 inch - ""== ^ .055 mm). 380 Great Basin Naturalist Vol. 37, No. 3 y/rm \ pr 1 ^, s •i'-4 ®fe JHj y \ @] Figs 22-27. Morphological variation of the holdfast (tribocytic organ) of adnlt P. mxnxmum. All specimens 72 r'u if ^^'^^' "°'*' ^^''''' '"""' ""**" ''"■'^'' well-developed holdfast; 24-25, Host: Chnisemys picta, two views of the holdfast and acetabulum, note the well-developed tegumental spination (Fig. 25); 26, Host; Canis famiUaris note the effects of the underlying musculature on the shape of the holdfast; 27, Host: Meriones unEuiciikitus note the contracted appearance of the holdfast. Sept. 1977 Palmieri: Trematode Variations 381 Table 2. Statistical analysis of acetabulum measurements of P. Minimum from experimental definitive hosts." Variable Mean of Combined Host Class Standard Deviation Means Amphibian Reptilian Aves Mammalia Acetabulum Length (AL) 0.047 Acetabulum Width (AW) 0.054 Cross-sectional Area of Acetabulum (AALYAX) 0.00.3 Ratio of Acetabulum Length to Width (RALAW) 0.888 Acetabulum Index of Length (ACEINDL) 0.095 Acetabulum Index of Width (ACEINDW) 0.205 Ratio of Oral Sucker Cross- sectional Area to Acetabulum Cross- sectional Area (ROSXXA) 0.524 0.009 0.010 0.001 0.193 0.024 0.045 1.552 0.050 0.191 0.237 0.049 0.047 0.060 0.058 0.002 0.002 0.8.32 0.794 0.083 0.( 0.212 0.202 0.388 0.429 0.371 Table 3. Statistical analysis of holdfast organ measurements of P. Minimum from experimental definitive hosts." Variable Mean of Combined Host Class Standard Deviation Means Amphibian Reptilian Aves Mammalia Holdfast Length (HL) Holdfast Width (HW) Cross-sectional Area of Holdfast (AHLXHW) Ratio of Holdfast Length to Width (RHLHW) 0.090 0.018 0.841 0.003 0.245 0.087 0.060 0.106 0.104 0.019 0.104 0.072 0.127 0.116 0.009 0.005 0.014 0.012 0.854 0.672 0.872 0.912 'All measurements in mm. 382 Great Basin Naturalist Vol. 37, No. 3 Figures 22-23 show the appearance of the holdfast organ in the extended appearance, whereas Figures 26-27 demonstrate the holdfast organ being pulled inward by the underlying musculature. Examination of the holdfast organ shows that it is well endowed with tegumental spines (Fig. 25). Such spines do not vary ap- preciably in size and shape as do tegumen- tal spines from other body regions, as re- ported in part II of this study. Acknowledgments The author thanks Dr. Martin J. Ulmer for his time and guidance throughout this research project, Dr. D. Cox and Paul Du- bose for data analysis, Dr. Darwin Wittrock for aid in fieldwork, and Mr. James Amrine for specimen preparation for S.E.M. Appre- ciation is also extended to the Iowa State Conservation Commission, the Federal Bu- reau of Sport Fisheries for animal collec- tion, and the Iowa State University Depart- ments of Zoology, Genetics, and Statistical Computations. This study was supported in part by the National Science Foundation Grant GB-23057; the Thomas H. MacBride Scholarship from the Iowa Lakeside Labora- tory; Brigham Young University Depart- ment of Zoology; and the Department of International Health, School of Medicine, University of California, San Francisco, from the National Institute of Allergy and Infectious Diseases, National Institutes of Health, U.S. Public Health Service GrantAI 10051 (UC ICMR). Literature Cited Palmieri, J. R. 1975. Host-Parasite Relationships and Intraspecific Variation in the Strigeoid Treinatode Posthodiplostomum ininimtnn (Trematoda: Diplos- tomatidae). Proc. Utah Academy of Science, Arts and Letters 52(1) 68-69. NEW SYNONYMY AND NEW SPECIES OF AMERICAN BARK BEETLES (COLEOPTERA: SCOLYTIDAE), PART V Stephen L. Wood^ Abstract.— New synonymy of American Scolytidae is proposed as follows: Phloeohorus Erichson ( = Phloeot- rtipes Erichson), Phheotribus Latreille { = Eiilytocerus Blandford), Scohjtudes Ferrari { = Epornaditts Blandford), Carphohoms pinicolens Wood ( = Carphohorm tubercukitus Bright), Coccotrypes indicus (Eggers) { = Dryocoetes subimpressus Eggers), Coccotrypes surinamensis Schedl { = Coccotrypes brevipilosus Eggers), Conophthonis con- iperda Schwarz { = Conophthonis clunicus Hopkins, Conophthorus taedae Hopkins), Conophthonis ponderosae Hopkins { = Conophthonis scopulorum Hopkins, Conophthorus contortae Hopkins, Conophthorus monticolae Hop- kins, Conophthonis flexilis Hopkins, Conophthorus lambertianae Hopkins), Conophthorus resinosae Hopkins ( = Conophthorus virginianae Hopkins), Cryphalomorphus knabi (Hopkins) { = Cryphahmorphus mimitissimus Schedl), Ips grandicollis (Eichhoff) ( = Toinicus cribricoUis Eichhoff), Ips perturhatus (Eichhoff) ( = Tomicus inter- punctits Eichhoff), Ips tridens engehnanni Swaine { = lps yohoensis Swaine, Ips semirostris Hopping, Ips amisk- wiensis Hopping), Leperisinus aculeatus (Say) { = Hylesinus imperialis Eichhoff), Leperisinus californicus Swaine { = Leperisinus hoferi Blackman), Phheosimis serratus (LeConte) { = Phloeosinus neotropicus Schedl), Phoeotribus championi (Blandford) ) = Eulytocerus substriatus Schedl), Pseudohylesinus sericeus (Mannerheim) { = Pseudohyle- sinus yasanuitsui Nobuchi), Pityophthorus boycei Swaine { = Pityophthorus siouxensis Bright), Pityophthonis de- letus LeConte { = Pityophthorus inquietus Blackman, Pityophthorus monophyllae Blackman, Pityophthorus prael- atus Bright, Pityophthorus brucki Bright), Scolytus opacus Blackman ( = Scolytus abietis Blackman), Scolytus tsugae Swaine { = Scolytus reflexus Blackman, Scolytus wickhami Blackman), Scolytus uniipinosus LeConte { = Scolytus fiskei Blackman). The following species are named as new to science: Phheotribus hirtus, P. ingae, P. levis, P. minor, P. nebulosus, P. simplicidens (Colombia), P. amplus, P. fici, P. squamiger, P. tetricus, P. vesculus (Venezuela), Pityophthorus discretus (Mexico). On the following pages several newly dis- covered cases of synonymy and species new to science are presented for American Scolytidae. The species new to science rep- resent the genera Phloeotribus (12) and Pi- tyophthorus (1). They are from Mexico (1), Colombia (7), and Venezuela (5). New Synonymy Phloeohorus Erichson Phloeoborus Erichson, 18.36, Arch. Naturgesch. 2(1): 54 (Type-species: Phloeoborus rudis Erichson, sub- sequent designation by Hopkins, 1914, Proc. U.S. Nat. Mus. 48: 126). Phloeotrupes Erichson, 18.36, Arch. Naturgesch. 2(1): .53 (Type-species: Phloeotrupes grandis, subsequent designation by Hopkins, 1914, Proc. U.S. Nat. Mus. 48: 127). Netv synonymy The holotype and 21 other specimens of Phloeohorus rudis Erichson, the holotype and two other specimens of Phloeotrupes grandis Erichson, and 10 other species in this complex were examined and compared to one another. The characters of the tibiae, antennae, and palpi on which the separation of genera was based, are so variable that no combination of them will distinguish genera. I propose that they be placed in synonymy. Although the name Phloeotrupes has page priority, I exercise the option of the first revisor and select Phloeoborus as the valid name for the genus, because it is much more widely known in the literature and has been applied to a much larger number of species. Phheotribus Latreille Phheotribus Latreille, 1796, Precis des caracteres generiques des insects, p. 50 (Type-species: Hyh- sinus ohae Fabricius = Scolytus scarabaeoides Bernard, subsequent inclusion by Latreille, 1802/3, Histoire naturelle, generale et particu- laere des Crustaces et des Insects 3: 203; spelling emendation from Phhiotrihus, Latreille, 1804, Mag. f. Insekenk. 3: 108) Eulytocerus Blandford, 1897, Biol. Centr. Amer., Coleopt. 4(6): 161 (Type-species: Eulytocerus championi Blandford, monobasic). New synonymy The holotype of Eulytocerus championi Blandford was examined and was found to 'Part of the research was sponsored by the National Science Foundation. ^Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. Scolytidae contribution 383 384 Great Basin Naturalist Vol. 37, No. 3 be congeneric with Phloeotribus Latreille. All characters of apparent generic value in championi are shared by most species of Phloeotribus. Scolytodes Ferrari Scolytodes Ferrari, 1867, Die Forst- und Baumzuchtschadlichen Borkenkafer, p. 77 (Type- species: Scolytodes laevigatus Ferrari, monobasic) Epomadius Blandford, 1897, Biol. Centr. Amer., Coleopt. 4(6): 179 (Type-species: Epomadius culci- tatus Blandford, monobasic). New synonymy Both female syntypes of Epomadius cidci- tatus Blandford were examined. The only unique character on which the genus Epo- madius was based was the conspicuous pi- lose impression on each side of the pro- thorax. There are no other significant characters that separate this species from Scolytodes. While the species culcitatus is unique, it must be transferred to Scolytodes where allied species are placed. Carphoborus pinicolens Wood Carphoborus pinicolens Wood, 1954, Canadian Ent. 85: 512 (Holotype, female; Logan Dry Canyon, Utah; U.S. Nat. Mus.) Carplwborus tuberculatus Bright, 1964, Pan Pacific Ent. 40: 165 (Holotype, female; Crooked Creek, White Mts., Mono Co., California; California Acad. Sci.). New synonymy The large type series of Carphoborus pi- nicolens Wood and the holotype and 48 paratypes of C. tuberculatus Bright were ex- amined and compared to one another. The tubercle at the center of the frons in tu- berculatus is represented in only 77 percent of the females in the type series; it is also present in 70 percent of the females in the type series of pinicolens. For this reason, because no other characters distinguish these populations, and since there is contin- uous distribution between the type localities of this widely distributed species, Bright's name must be placed in synonymy. Coccotrypes indicus (Eggers) Thamnurgides indicus Eggers, 1936, Ann. Mag. Nat. Hist. (10)17:631 (Holotype, female; Sakalaspur, Mysore, India; British Mus. Nat. Hist.) Dryocoetes subimpressus Eggers, 1940, Arb. Morph. Tax. Ent. Berlin-Dahlem 7: 127 (Holotype, fe- male; Trois Rivieres, Guadeloupe; deposited in Eggers Collection, apparently on loan to Schedl). New synonymy The holotype of Dryocoetes subimpressus Eggers is a female of indicus (Eggers) hav- ing a normal head, prothorax, the lateral parts of the elytra, and ventral parts. The median areas of the elytral disc and declivi- ty have the interstriae and spaces between the strial punctures in each row abnormally roughened and distorted. The distortion is similar to that seen in specimens injured by siblings during the pupal stage. Since this is an aberration of a recognizable common species, the name subimpressus must be placed in synonymy. Coccotrypes surinamensis Schedl Coccotrypes surinamensis Schedl, 1949 (1948), Tijdscr. Ent. 91: 116 (Syntypes; Surinam; Zool. Mus. Am- sterdam and Schedl Coll.) Coccotrypes brevipilosus Eggers, 1951, Ent. Blatt. 45- 46: 150 (Holotype, female; Blumenau, Brazil; de- posited in Eggers Coll., presumably on loan to Schedl). New synonymy Four specimens, presumably syntypes, of Coccotrypes surinamensis Schedl exhibit var- iability in length of the interstrial setae from very short to normal length for this genus. In one of the specimens these setae, although a bit less consistent in length, are almost indentical to the very short setae on the holotype of C. brevipilosus Eggers. Two of the specimens are intermediate between brevipilosus and C. surinamensis and the fourth specimen is virtually indistinguishable from surinamensis. Since virtually all Amer- ican Coccotrypes are known to have been introduced from Africa or the Indo-Austra- lian region, it is very likely that the sus- pected synonymy with sundaensis (Eggers, 1923) will be established as soon as ade- quate material is available for study. The entire genus Coccotrypes is in a state of tax- onomic chaos because so many "species" have been named from extraterritorial (in- troduced) populations without any apparent attempt to establish their origins that identi- fication of most of them is virtually impos- sible. Conophthorus coniperda (Schwarz) Pityophthorus coniperda Schwarz, 1895, Proc. Ent. Soc. Washington (Lectotype, male; Marquette, Mich- igan; U.S. Nat. Mus., present designation) Conophthorus clunicus Hopkins, 1915, J. Washington Acad. Sci. 5: 432 (Holotype, female; 13, 122, To- Sept. 1977 Wood: American Scolytidae 385 micus cluriicus Fitch, Det. No. 12 Hopk., Collec- tion Fitch; U.S. Nat. Mus.). New synonymy Conophthorus taedae Hopkins, 1915, J. Washington Acad. Sci. 5: 431 (Holotype, sex?; Ft. Monroe, Va.; U.S. Nat. Mus.). New synonymy Of the three syntypes of Pitijophthonis coniperda Schwarz examined at the U.S. National Museum, two are from Marquette and one is from Eagle Harbor, Michigan. I here designate a male syntype from Marquette as the lectotype of Pityophthonis coniperda Schwarz. These three syntypes were compared to the holotypes of C. clu- nicus Hopkins and C. taedae Hopkins and to 113 other specimens. Only one species is represented by this material. The smaller size, steeper elytral declivity, tuberculate declivital interstriae 1, and other characters clearly distinguish it from the allied resi- nosae Hopkins. Conophthorus ponderosae Hopkins Conophthorus ponderosae Hopkins, 1915, J. Washing- ton Acad. Sci. 5: 431 (Holotype, female; Ashland, Oregon; U.S. Nat. Mus.) Conophthorus scopulorum Hopkins, 1915, J. Washing- ton Acad. Sci. 5: 431 (Holotype, female; Flagstaff, Arizona; U.S. Nat. Mus.). New synonymy Conophthorus contortae Hopkins, 1915, J. Washington Acad. Sci. 5: 432 (Holotype, female; Newport, Oregon; U.S. Nat. Mus.). New synonymy Conophthorus monticolae Hopkins, 1915, J. Washington Acad. Sci. 5: 432 (Holotype, female; Priest River, Idaho; U.S. Nat. Mus.). New synonymy Conophthorus flexilis Hopkins, 1915, J. Washington Acad. Sci. 5: 4.33 (Holotype female; Mount Manitou, Colorado; U.S. Nat. Mus.). New synony- my Conophthorus kimhertianae Hopkins, 1915, J. Washing- ton Acad. Sci. 5: 433 (Holotype, female; Hilt, Cal- ifornia; U.S. Nat. Mus.). New synonymy The female holotypes of Conophthorus ponderosae Hopkins, C. scopulorum Hop- kins, C. contortae Hopkins, C. monticolae Hopkins, C. flexilis Hopkins, and C. lamher- tianae Hopkins were compared directly to one another and .studied in conjunction with 784 other specimens. Although a limited amount of variability occurs in all series, I see no basis for recognizing more than one species from this material. Because the name ponderosae has been used more wide- ly in the literature, it was .selected to desig- nate this species. In view of the sibling spe- cies resinosae and banksianae that are distinguishable only from their habits there is a po.ssibility that sibling species occur in the material treated here as ponderosae. Ex- tensive biological work will be required to determine whether or not this is the case. Conophthorus resinosae Hopkins Conophthorus resinosae Hopkins, 1915, J. Washington Acad. Sci. 5: 431 (Holotype, male; Conoph. resi- nosae Hopkins, 1 Harring., Conophthorus resi- nosae; U.S. Nat. Mus.) Conophthorus virginianae Hopkins, 1915, J. Washing- ton Acad. Sci. 5: 431 (Holotype, male, not a fe- male as labeled; Huttonsville, West Virginia; U.S. Nat. Mus.). New synonymy The male holotypes of Conophthorus resi- nosae Hopkins and C. virginianae Hopkins were compared directly to one another and were studied with 166 other specimens. Al- though minor variation exists in this mate- rial, only one species is represented. It is distinguished from the allied C. coniperda (Schwarz) by characters listed above under that species. Except for size and host I see no means of distinguishing it from C. bank- sianae McPherson, although both appear to be valid species. Cryphalomorphus knahi (Hopkins) Ernoporides knahi Hopkins, 1915, Dept. Agric. Rept. 99: 34 (Holotype, female; Cordoba, Mexico; U.S. Nat. Mus.) Cryphalomorphus minutissimus Schedl, 1951, Dusenia 2; 97 (Holotype male; Env. de Trois Rivieres, Guadeloupe; Schedl Coll.). New synonymy The male holotype of Cryphalomorphus minutissimus Schedl is somewhat smaller than average for the species and the basal half of the elytra have been rubbed, but there is no doubt that it represents C. knahi (Hopkins), a very common and widely dis- tributed .species in the Caribbean region. The type of minutissimus when received on loan for study had fallen from its mount and was crushed between the wall and pin- ning surface of the shipping box. The frag- ments were recovered and re-mounted by me; no other possible contaminating parts were in the box. Ips grandicollis (Eichhoff) Tomicus grandicollis Eichhoff, 1868, Berliner Ent. Zeitschr. 11:402 (Syntypes; Amerique boreali; presumably lost with Hamburg Mus.) Tomicus cribricollis Eichhoff, 1869, Berliner Ent. 386 Great Basin Naturalist Vol. 37, No. 3 Zeitschr. 12: 273 (Holotype, male; Mexico; Brus- sels Mus.). New synonymy This species is easily recognized from the description, although the type series of To- micus grandicollis Eichhoff is lost. More than 1,000 specimens from the United States and eastern Canada were examined in addition to the holotype of Toinicus cribricollis Eichhoff and about 500 speci- mens from Mexico and Central America. Although geographical variation is apparent, it does not appear to be correlated with characters reported by Lanier (1970, Cana- dian Ent. 102: 1151) nor can I confirm all of the characters he used to distinguish grandicollis and cribricollis. Until a more comprehensive analysis can be made, cribri- collis should be regarded as a junior syno- nym of grandicollis. Ips perturbatus (Eichhoff) Tomicus perturbatus Eichhoff, 1869, Berliner Ent. Zeitschr. 12: 274 (Syntypes?; Amerique boreali; presumably lost with Hamburg Mus.) Tomicus interpunctus Eichhoff, 1878, preprint of Mem. Soc. Roy. Sci. Liege (2)8: 241 (Syntypes?; Sitka, Alaska; presumably lost with Hamburg Mus.). New synonymy The type series of both Tomicus per- turbatus Eichhoff and T. interpunctus Eich- hoff were lost when the Hamburg Museum was destroyed. Prior to that time Eichhoff had examined part of the original series of T. hudsonicus LeConte and referred them to perturbatus. The identity of interpunctus has not been clarified. According to the original description, interpunctus was 4.0 mm long, it had a pair of transversely ar- ranged tubercles near the center of the frons, and the basal parts of the interstriae were impunctate. The only Sitka Ips of this size, with paired frontal tubercles and ba- sally impunctate interstriae is perturbatus. It is also the most common Ips in the Sitka area. Although pini (Say) has frontal tu- bercles in some parts of its range, it does not have them in areas adjacent to Sitka and there are no records of it from Sitka. For these reasons, interpunctus is placed in synonymy under perturbatus. Ips tridens engelrnanni Swaine Ips engehmmni Swaine, 1917, Dom. Canada Dept. Agric. Ent. Br. Tech. Bull. 14(1): 30 (Lectotype, female; Rogers Pass, British Columbia; Canadian Nat. Coll., designated by Bright, 1967, Canadian Ent. 99: 675) Ips yohoensis Swaine, 1917, Dom. Canada Dept. Agric. Ent. Br. Tech. Bull. 14(1): 31 (Lectotype, female; Yoho Valley, British Columbia; Canadian Nat. Coll. designated by Bright, 1967, Canadian Ent. 99: 676). Neiv synonymy Ips semirostris Hopping, 1963, Canadian Ent. 95: 213 (Holotype, female; Kenai Penninsula, Alaska; Ca- nadian Nat. Coll.). New synonymy Ips amiskwiensis Hopping, 196.3, Canadian Ent. 95: 216 (Holotype, female; Amiskwi River Yoho Nat. Pk., British Columbia; Canadian Nat. Coll.). New synonymy The types and most of the type series of Ips engelrnanni Swaine, /. yohoensis Swaine, I. semirostris Hopping, and /. amiskwiensis Hopping and 580 other specimens of this species were examined and compared. The female frons is unusually variable and ap- pears in forms varying from almost uni- formly convex to moderately protuberant to exceedingly protuberant, with or without abundant vestiture. The various forms may appear in pure or mixed series and are known to be morphological expressions of various gene combinations and do not ex- press specific differences. Leperisinus aculeatus (Say) Hylesinus aculeatus Say, 1824, J. Acad. Nat. Sci. Phila- delphia 3: 322 (Syntypes?; Missouri; evidently lost) Hylesinus imperialis Eichhoff, 1868, Berliner Ent. Zeitschr. 12: 149 (Syntypes; Wisconsin and Georgia; lost with Hamburg Mus.). New synony- my LeConte's series of Hylesinus aculeatus Say was either part of the Say series or was compared to it; his series of H. imperialis Eichhoff was either part of Eichhoff s series or was compared to it. Both of LeConte's series are of the same species. The size, de- scriptions, and distributions are such that there is virtually no question that both names apply to the same species. In view of these facts and since imperialis has been a doubtful species since it was first proposed, these names are considered synonymous. Leperisinus californicus Swaine Leperisinus californicus Swaine, 1916, Canadian Ent. 48: 190 (Holotype, female; San Diego, California; Canadian Nat. Coll.) Sept. 1977 Wood: American Scolytidae 387 Leperisintis hoferi Blackman, 1943, Proc. U.S. Nat. Mus. 93: 394 (Holotype, female; Sabino Canyon, Arizona; U.S. Nat. Mus.). New synonymy The holotypes of Leperisinus californicus Swaine and L. hoferi Blackman and 242 other specimens from Oregon and North Dakota to CaHfornia and Chihuahua were examined and compared. From this material it is apparent that only one species is pres- ent. Apparently Blackman was not familiar with Swaine's species when hoferi was named. Phloeosinus serratus LeConte Hyleaintis serratus LeConte, 1868, Trans. Amer. Ent. Soc. 2: 170 (Holotype, male; Middle States; Mus. Comp. Zool. Phloeosinus neotropiciis Schedl, 1939, Proc. Roy. Ent. Soc. London 8(1): 12 (Holotype, female; Jamaica; British Mus. Nat. Hist.). Netv synonymy The male holotype of Htjiesinus serratus LeConte was examined and compared to several examples of my specimens. It repre- sents a common species from Washington and Idaho to Durango (Mexico) and Texas. The female holotype of Phloeosinus neo- tropicus Schedl was examined, but, as is the case with many female Phloeosinus, it could not be associated with any other material at that time. The male "paratype" in the Schedl collection, however, is an easily di- agnosed normal specimen of serratus. Once this association was made, the female holo- type could then be placed as a synonym of serratus. There is doubt as to whether or not this species is established in Jamaica. Phloeotrihus championi (Blandford), n. comb. Eulytocerus championi Blandford, 1897, Biol. Centr. Amer., Coleopt. 4(6): 161 (Holotype, male; Volcan de Chiriqui, Chiriqui, Panama; British Mus. Nat. Hist.) Euhjtocerus suhstriatus Schedl, 1935, Rev. de Ent. 5: 344 (Holotype, male; Turrialba, Costa Rica; Schedl Coll.). Neiv synonymy The male holotypes of Euhjtocerus cham- pioni Blandford, length 3.7 m, and E. suh- striatus Schedl, length 3.6 mm, were com- pared directly to one another. Although the frontal setae of the former are slightly long- er and the interstrial punctures of the latter are slightly deeper, the differences appear insignificant. Only one species can be rec- ognized in this material; consequently, Schedl's name is placed in synonymy. Pseudoliijlesinus sericeus (Mannerheim) Hylesinus sericeus Mannerheim, 1843, Bull. Soc. Imp. Nat. Moscou 16(2): 296 (reprint p. 124) (Holotype, female; Alaska; Univ. Zool. Mus., Helsinki) Pseudohylesinus yasamatsui Nobuchi, 1971, Bull. Gov. For. Expt. Sta., Tokyo 2.38: 160 (Holotype, male; Takanishi, Nagano, Japan; Gov. For. Expt. Sta., Tokyo). New synonymy Two paratypes of Pseudohylesinus yasa- matsui Nobuchi in the Canadian National Collection are identical to my series of se- riceus (Mannerheim) two of which were compared to the holotype at the Helsinki Museum by me. Nobuchi's name therefore, must be placed in synonymy. This is the first reported introduction of a member of this genus outside of western North Ameri- ca. Pityophthorus boycei Swaine Pityophthorus boycei Swaine, 1925, Canadian Ent. 57: 192 (Holotype, male; Cisco, Placer Co., Cali- fornia; Canadian Nat, Coll.) Pityophthorus siouxensis Bright, 1977, Great Basin Nat. 36: 439 (Holotype, female; Black Hills, South Da- kota; Canadian Nat. Coll.). Neiv synonymy The type series of Pityophthorus boycei Swaine, P. catulus Blackman, P. iniquus Blackman, and P. siouxensis Bright and 133 other specimens from Briti.sh Columbia and California eastward to South Dakota and Colorado were examined and compared di- rectly to one another. Although slight varia- tion within and between series is evident, it is clear that only one species is represented by this material. Pityophthorus deletus LeConte Pityophthorus deletus LeConte, 1879, U.S. Dept. Inte- rior, Geol., Geogr. Surv. Bull. 5: 519 (Lectotype, female; Veta Pass, Colorado; Mus. Comp. Zool., designated by Bright, 1976, Coleopt. Bull. 30: 185) Pityophthorus inquietus Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25: 46 (Holotype, female; Las Vegas Hot Springs, New Mexico; U.S. Nat. Mus.). New synonymy Pityophthorus monophyllae Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25: 47 (Holo- type, female; Argus Mts., California; U.S. Nat. Mus.). New synontjmy Pityophthorus praelatus Bright, 1966, Pan Pacific Ent. 388 Great Basin Naturalist Vol. 37, No. 3 42: 303 (Holotype, female; Mt. Shasta Ski Area, Siskiyou Co., California; California Acad. Sci.). New synonymy Pityophthorus brucki Bright, 1971, Pan Pacific Ent. 47: 63 (Holotype, female; San Bernardino, Califor- nia; Ohio State Univ. Coll.). New synonymy Two syntypes of Pityophthorus deletus LeConte, the holotypes and most of the type series of P. inquietus Blackman, P. monophyUae Blackman, and P. brucki Bright, and two para types of P. praelatus Bright, and more than 400 other specimens were studied and compared directly to one another and to examples of my material. Variations in the female frons are con- spicuous and are partly associated with ge- ographic origin. The type series of mon- ophyUae, from the Argus Mountains, California, and a series from Beaver, Utah contain some females with very long and others with very short or no frontal setae. Series from San Bernardino and Ventura counties, California contained females with the frons very strongly, extensively flattened in various patterns and with long hair in a variety of arrangements. In view of the lim- ited material available from California and the conspicuous variability within and be- tween series in the remainder of the range, from South Dakota to Durango (Mexico) and westward, it is considered advisable to treat all of this material as one species until a comprehensive analysis can be made. Scolytus opacus Blackman Scolyttis opacus Blackman, 1934, U.S. Dept. Agric. Tech. Bull. 431: 20 (Holotype, male; Ouray, Colo- rado; U.S. Nat. Mus.) Scolytus abietis Blackman, 1934, U.S. Dept. Agric. Tech. Bull. 431: 21 (Holotype, male; Sandpoint, Idaho; U.S. Nat. Mus.). New synonymy The male holotypes of Scohjtus opacus Blackman and S. abietis Blackman and more than one hundred other specimens were compared directly to one another. Speci- mens from the Pacific Coast states and northern Idaho have the spine on male ster- num 2 slightly smaller, with the summit more narrowly rounded and its crest acute and with the vertex of the head more strongly convex. Some males from the southern Rocky Mountain area have the summit of the spine on sternum 2 flattened or sulcate and the vertex less strongly con- vex. Since the types of opacus and abietis represent the extremes in these characters and because both characters intergrade in series from intermediate localities, it is nec- essary to place one of the names in synony- my. The name opacus was selected as the name to be used for this species. Scohjtus tsugae Swaine Scolytus tsugae Swaine, 1917, Dom. Canada Dept. Agric. Ent. Br. Tech. Bull. 14(1): 32 (Lectotype, female; Glacier, British Columbia; Canadian Nat. Coll., designated by Bright, 1967, Canadian Ent. 99: 674) Scolytus reflexus Blackman, 1934, U.S. Dept. Agric. Tech. Bull. 431: 13 (Holotype, male; Santa Cata- lina Mts., Arizona; U.S. Nat. Mus.). New synony- my Scolytus wickhami Blackman, 19.34, U.S. Dept. Agric. Tech. Bull. 431: 13 (Holotype, male; Buena Vista, Colorado; U.S. Nat. Mus.). New synonymy In an earlier paper, I (Wood, 1966, Great Basin Nat. 26: 30) placed Scolytus mon- ticolae Swaine in synonymy under S. tsugae Swaine. The examination of the holotypes of S. reflexus Blackman and S. wickhami Blackman and more than 100 other speci- mens from eastern Washington and northern Idaho to Arizona leaves little doubt that only one species exists in this area. How- ever, this poulation forms a longitudinal pa- rental gallery, while the coastal form appar- ently forms a transverse one. The absence of significant morphological characters sug- gests that only one species is represented by this material and that reflexus and wickhami must be placed in synonymy. The apparent difference in galleries suggests the necessity for a critical reexamination of this species as soon as adequate material for study is available. Scolytus unispinosus LeConte Scolytus unispinosus LeConte, 1876, Proc. Amer. Philos. Soc. 15: 372 (Lectotype, male; Oregon; Mus. Comp. Zool., present designation) Scolytus fiskei Blackman, 1934, U.S. Dept. Agric. Tech. Bull. 431:25 (Holotype, male; Capitan Mts., New Mexico; U.S. Nat. Mus.). New synony- my The male syntype of Scolytus unispinosus in the LeConte collection that is labeled "type" is here designated as the lectotype of LeConte's name unispinosus. That lecto- type, the holotype of S. fiskei Blackman, Sept. 1977 Wood: American Scolytidae 389 and 164 other specimens from British Co- lumbia to Arizona were examined. Although slight geographical differences are apparent, individual variation within series is such that it is impossible to recognize geographi- cal races or species. Therefore, fiskei must be placed in synonymy under the senior name, unispinosus. New Taxa Phloeotribus tetricus, n. sp. This species is distinguished from ve- nezuelensis Schedl by the larger average size, by the larger, deeper pronotal punc- tures, and by differences in the frons, elytra, and vestiture noted below. Male.— Length 1.8 mm (paratypes 1.7- 2.1 mm), 1.8 times as long as wide; color reddish brown. Frons transversely impressed slightly above level of antennal insertion, convex above, with a large, subfoveate impression on median fourth, lower area shallowly, rather broadly impressed, epistomal margin rather broadly, acutely carinate, carina low, indefinite laterally; surface shining, upper area coarsely punctured, rugose-reticulate in interspaces, lower area reticulate, with a few fine punctures; vestiture inconspicuous. Antenna about as in veneznelensis. Pronotum similar to venezuelensis except punctures coarser, deeper, vestiture more slender. Anterolateral areas unarmed in both species. Elytra similar to venezuelensis. except strial punctures considerably larger, deeper, interstriae half as wide as striae, tubercles on odd-numbered interstriae larger, high, narrow, pointed, continuing to lower half of declivity, those on even-numbered inter- striae obsolete except near base, vestiture more abundant, more slender, a few on cen- tral row of each interstriae longer. Female.— Similar to male except lower half of frons almost flat, carina not clearly evident, transverse impression less con- spicuous. Type Locality.— La Carbonera Experi- mental Forest, about 50 km (airline) NW Merida, Merida, Venezuela. Type Material.— The male holotype, fe- male allotype, and 25 paratypes were taken at the type locality on 27-X-1969, 2500 m. No. 93, from the bole of Eschiveilera sp., by me. Two paratypes were taken at the same locality on 12-1-1970, from a woody vine, by me. The holotype, allotype, and para- types are in my collection. Phloeotribus ingae, n. sp. This species is distinguished from simplex Wood by the smaller size, by the absence of reticulation on the pronotum, by the less strongly impressed striae and strial punc- tures, and by the more slender elytral setae. Male.— Length 1.5 mm (paratypes 1.4- 1.7 mm), 2.0 times as long as wide; color brown. Frons broadly, deeply concave from epis- toma to upper level of eyes, lateral margins subcostate on lower half; surface smooth, shining, punctures small, rather sparse; ves- titure short, rather sparse, inconspicuous. Pronotum as in simplex except surface smooth, shagreened, without any re- ticulation, punctures more sharply, rather shallowly impressed. Elytra as in simplex except surface slightly shagreened, strial punctures slightly smaller, interstriae one and one-half times as wide as striae, interstrial tubercles small- er in diameter, higher, vestiture slightly more slender, of similar length and abun- dance, strial punctures on declivity much smaller, less strongly impressed. Female.— Similar to male except frons al- most flat on lower half, convex above, punctures moderately coarse, shallow. Type Locality.— Bugo, Columbia. Type Material.— The male holotype, fe- male allotype, and 101 paratypes were taken at the type locality on 16- V- 1973, from two pods of Inga sp., by G. Ekis. The holotype, allotype, and paratypes are in my collection. Phloeotribus simplicidens, n. sp. This species is distinguished from ingae Wood by the much more deeply impressed male frons, by the finer punctures and deep median fovea on the female frons, and by other characters described below. Male.— Length 1.8 mm (paratypes 1.7- 390 Great Basin Naturalist Vol. 37, No. 3 1.9 mm), 2.0 times as long as wide; color yellowish brown. Frons as in ingae except impression dis- tinctly deeper, its surface reticulate. Pronotum as in ingae except surface con- tour more uniform, punctures slightly larger, deeper. Elytra as in ingae except strial punctures slightly larger, interstrial tubercles smaller, vestiture finer; interstrial tubercles on de- clivity almost obsolete on lower half. Female.— Similar to male except frons convex; as in female ingae except punctures very small, central area with a large, deep fovea. Type Locality.— Finca La Hermosa, Sa- lento, Caldas, Colombia. Type Material.— The male holotype, fe- male allotype, and eight paratypes were taken at the type locality on l-VI-1959, en guamo, by J. A. Garzon. The holotype, allotype, and paratypes are in my collection. Phloeotribus atnplus, n. sp. This species apparently represents a unique species group within the genus. It is distinguished by the large size, rather slen- der form, simple male frons, asperate pro- notum and simple elytra. It may be remote- ly related to the setulosiis group of species. Male.— Length 3.3 mm (paratypes 3.3- 3.8 mm), 2.1 times as long as wide; color dark brown. Frons with a large, strong, median fovea two-thirds distance from epistoma to upper level of eyes, lower area shallowly im- pressed, smooth, shining, closely, finely punctured, upper area convex, rather coarsely punctured, with a few small, isolat- ed tubercles; vestiture sparse, inconspicuous. Each segment of antennal club about three times as wide as long. Pronotum 0.8 times as long as wide; widest at base, sides accurately converging toward rather broadly rounded anterior margin; surface shining, rather finely, densely asperate except basal third on me- dian half closely, rather coarsely, deeply punctured. Vestiture short, fine, moderately abundant. Elytra 1.4 times as long as wide, 2.0 times as long as pronotum; sides straight and parallel on basal two-thirds, rather nar- rowly rounded behind; striae moderately impressed, punctures coarse, deep, very close; interstriae as wide as striae, shining, imiseriately crenulate except confused near base on 1 and 3, crenulations each half as wide as interstriae, transversely, acutely, moderately elevated, submarginal ones at base larger. Declivity steep, convex; striae and interstriae about half as wide as on disc, crenulations narrower, forming acutely pointed, small tubercles, interstriae 9 slightly elevated, more strongly serrate. Ves- titure of uniseriate rows of fine interstrial setae, each seta equal in length to three- fourths distance between rows. Female.— Similar to male except frons convex, with a distinct transverse impression and a conspicuous median fovea near middle. Type Locality.— La Carbonera Experi- mental Forest, 50 km (airline) northwest Merida, Merida, Venezuela. Type Material.— The male holotype, fe- male allotype, and 10 paratypes were taken at the type locality on 14-X-69, 2500 m. No. 59, from the bole of a large uniden- tified tree, by me. The holotype, allotype, and paratypes are in my collection. Phloeotribus nehulosus, n. sp. Although this species apparently belongs to the setulosus group, it lacks an epistomal male carina, the male scape is less densely setose, and declivital interstriae 9 is less strongly elevated than in most representa- tives of the group. Male.— Length 1.9 mm (paratypes 1.7- 2.0 mm), 1.9 times as long as wide; color very dark brown, almost black. Frons shallowly concave from epistoma almost to upper level of eyes, a transverse impression with a median fovea somewhat deeper two-thirds of distance from epistoma to upper level of eyes; epistomal carina not indicated; surface strongly reticulate, a few small, obscure punctures and granules on upper third; vestiture sparse, short, incon- spicuous. Antenna about as in setulosus Sept. 1977 Wood: American Scolytidae 391 Eichhoff except long setae on scape about one-third as numerous. Pronotum about as in setulosiis except as- perities smaller, punctures on basal half larger, more clearly impressed, vestiture much finer. Elytra 1.3 times as long as wide, 1.9 times as long as pronotum; much as in setu- losiis except interstriae slightly impressed at base, moderately impressed on posterior half of disc, punctures coarse, deep; interstriae as wide as striae, crenulations narrower than in setolosus; declivital sculpture about as on disc except striae and interstriae slightly narrower, tubercles narrower, pointed; de- clivital interstriae 9 much less strongly ele- vated than in setulosus, moderately high, rather coarsely, closely tuberculate. Vesti- ture of uniseriate rows of fine bristles, each slightly shorter than distance between rows. Female.— Similar to male except frons convex, strongly reticulate, obscure punc- tures of moderate size indicated, a median fovea slightly above middle. Type Locality.— Piedras Blancas, 1 1 km west of Medellin, Antioquia, Colombia. Type Material.— The male holotype, fe- male allotype, and 126 paratypes were taken at the type locality on 17-VII-1970, 2300 m, No. 693, from Croton guianensis, by me. The holotype, allotype, and paratypes are in my collection. Phloeotribus minor, n. sp. This species is distinguished from, armatus Blandford by the much smaller size and by numerous characters described below. Male.— Length 1.3 mm (paratypes 1.2- 1.4 mm), 2.1 times as long as wide; color very dark brown. Frons as in armatus except less strongly impressed, epistomal carina less strongly elevated. Pronotum as in armatus except sculpture slightly finer. Elytra as in armatus except striae much less strongly impressed, punctures more clearly defined, more distinctly impressed, interstrial crenulations not as high or as coarse, declivital sculpture much finer, in- terstriae 9 only moderately elevated, much less strongly, less closely serrate, vestiture much finer. Female.— Similar to male except frons convex, with an indefinite central fovea, surface reticulate, punctures shallow, ob- scure, moderately coarse; asperities on ante- rolateral areas of pronotum usually larger; declivital interstriae 2 and 4 sometimes with a few tubercles. Type Locality.— Twenty-seven km northeast of Montoya, Santander, Colombia. Type Material.— The male holotype, fe- male allotype, and 36 paratypes were taken at the type locality on 2- VII- 1970, 150 m, No. 601, from a Pseudoohnedia limb, by me. Cladoctonus boliviae Wood was using the entrance tunnels of this species to gain access to the cambium region. The holotype, allotype, and paratypes are in my collection. PJdoeotribus remorsus, n. sp. This species is distinguished from setu- losus Eichhoff by the much larger size, by the dull, uniformly, strongly reticulate pro- notum, and by other characters described below. Male.— Length 2.9 mm (paratypes 2.4- 3.2 mm), 1.7 times as long as wide; color dark brown. Frons as in setulosus except slightly high- er, slightly narrower, reticulation more uni- form more strongly impressed. Pronotum as in setulosus except surface dull, uniformly, strongly reticulate, punc- tures very small, inconspicuous. Elytra as in setulosus except strial punc- tures smaller, less sharply defined, declivital tubercles on interstriae 3, 5, and 7 slightly larger, almost obsolete on 6 and 8, 9 more acutely elevated. Female.— Similar to male except irregu- larly convex, uniformly, strongly reticulate, punctures and central fovea obscure; ante- rolateral crenulations on pronotum larger. Type Locality.— Piedras Blancas, 10 km east Medellin, Antioquia, Colombia. Type Material.— The male holotype, fe- male allotype, and 80 paratypes were taken at the type locality on 15-VII-1970, 2500 m. No. 654, from a Quercus humholdtii branch, by me. 392 Great Basin Naturalist Vol. 37, No. 3 The holotype, allotype, and paratypes are in my collection. Phloeotribiis vescuhis, n. sp. This species may be remotely allied to squamatiis Wood, but it is distinguished by the stouter body form, by the obscure later- al asperities on the pronotum, and by other characters described below. Female.— Length 1.3 mm, 1.9 times as long as wide; color very dark brown. Frons convex, strongly reticulate, a pair of calluses on median two-thirds slightly above level of antennal insertion. Segments of antennal club slightly more than twice as wide as long. Pronotum about as in squamatus except anterior constriction obsolete, a few obscure crenulations in lateral areas; surface uni- formly reticulate, punctures very shallow, coarse. Setae short, coarse, each widest on its distal half. Elytra 1.2 times as long as wide, 1.6 times as long as pronotum; sides straight and parallel on basal two-thirds, very broad- ly rounded behind; striae not impressed ex- cept 1 weakly, punctures coarse moderately deep, close; interstriae slightly narrower than striae, shining, with moderately abun- dant fine, confused punctures, each with a uniseriate row of pointed tubercles, tu- bercles spaced by distance equal to width of an interstriae. Declivity very steep convex; striae and interstriae narrower than on disc; tubercles obsolete on interstriae 1 and 2. Vestiture consisting of interstrial rows of erect bristles; each bristle stout, widest on its distal half, about six times as long as wide, about two-thirds as long as distance between rows. Type Locality.— Five km west of El Pino, Zulia, Venezuela. Type Material.— The female holotype was taken at the type locality on 20-X- 1969, 10 m. No. 140, from an Ochroma branch, by me. The holotype is in my collection. Phloeotrihus levis, n. sp. This species apparently is distantlv allied to squamatus Wood but is distinguished by the impressed striae, slender interstrial setae, and many other characters. Male.— Length 1.5 mm (paratypes 1.7 mm), 2.1 times as long as wide; color very dark brown. Frons moderately concave from epistoma to upper level of eyes, sulcus narrower and deeper near middle, lateral margins and epistoma not elevated or armed; surface ret- iculate, obscurely so on lower half; vestiture short, fine, sparse. Segments of antennal club about four times as wide as long. Pronotum about as in squamatus except surface reticulate, punctures moderately coarse, sharply, shallowly impressed, spaced by one-half to three diameters of a punc- ture. Vestiture of fine, short, inconspicuous hairlike setae. Elytral outline about as in squamatus; striae impressed, moderately near base, rather strongly near declivity, punctures rather coarse, deep, close; interstriae as wide as striae, rather strongly convex, smooth and shining, each armed by a unise- riate row of moderately high crenulations, each crenulation about two-thirds as wide as interstriae, crenulations spaced in row by distance slightly less than width of an inter- striae. Declivity steep, convex; sculpture about as on disc except striae and inter- striae distinctly narrower, crenulations nar- rowed to become rounded tubercles except slightly larger on interstriae 9. Vestiture of interstrial rows of slender setae each as long as distance between rows, much stouter (8- 10 times as long as wide) on declivity. Female.— Similar to male except frons convex, an obscure, shallow median fovea at center, punctures rather sparse, moderately coarse declivital setae not as stout. Type Locality.— Piedras Blancas, 10 km ea.st of Medellin, Antioquia, Colombia. Type Material.— The male holotype, fe- male allotype, and one female paratype were taken at the type locality on 15-VII- 1970, 2500 m. No. 658, from a branch of a tree known locally as Graptero, by me. The holotype, allotype, and paratype are in my collection. Phloeotrihus squamiger, n. sp. This species is distinguished from squam- Sept. 1977 Wood: American Scolytidae 393 atus Wood by the larger size and by nu- merous minor characters described below. Male.— Length 1.5 mm (para types 1.4- 1.7 mm), 2.1 times as long as wide; color black with white setae. Frons as in sciuamatus except slightly more strongly, more uniformly concave, me- dian epistomal tubercle much narrower, slightly higher. Pronotum as in sqiiamatus except shallow punctures more clearly impressed, scales stouter. Elytra as in squamatus except strial pimctures evidently slightly larger, scales in interstrial rows stouter, each almost as wide as long. Female.— Similar to male except frons convex, strongly reticulate, with a few mi- nute granules. Type Locality.— Colonia Tovar, Aragua, Venezuela. Type Material.— The male holotype, fe- male allotype, and 30 paratypes were taken at the type locality on 4-V-1970, 1700 m. No. 495a, from Inga branches, by me. Two paratypes are from La Carbonera Experi- mental Forest near Merida, Merida, Vene- zuela, 27-X-1969, 2500 m. No. 89, from a tree seedling, by me. The holotype, allotype, and paratypes are in my collection. Phloeotrihiis hirtus, n. sp. This species is distinguished from spin- ipennis Eggers by the smaller size, by the vmarmed lateral margins of the male frons, by the more coarsely punctured pronotum, and by the very different declivity and ves- titure. Male.— Length 2.0 mm (paratypes 1.9- 2.1 mm), 2.0 times as long as wide; color very dark brown to almost black. Frons moderately, broadly concave from epistoma to upper level of eyes, lateral margins moderately elevated, unarmed; sur- face smooth, shining, a few, fine punctures and granules evident; vestiture fine, sparse, inconspicuous. Antennal scape with a small tuft of hair; segments of club about six times as wide as long. Pronotum shape about as in spinipennis; basal half very coarsely, closely, deeply punctured, interspaces smooth, shining, without micropunctures, asperities on ante- rolateral areas very small, obscure. Vestiture of moderately long, fine hair. Elytra 1.4 times as long as wide, 1.8 times as long as pronotum; outline about as in spinipennis; striae moderately impressed, punctures coarse, deep, close; interstriae as wide as striae, rather strongly convex, smooth shining, tubercles uniseriate, rather small except one or two near margin of de- clivity rather coarse. Declivity steep, con- vex; sculpture much as on declivity except striae and interstriae slightly narrower, strial punctures half as large; most interstrial tu- bercles replaced by punctures, a rather coarse tubercle at base on 3, 5, and 7, at middle on 3 and near apex on 1, 9 acutely, rather strongly elevated and coarsely ser- rate. Vestiture of slender, erect interstrial hair, each seta at base twice as long as dis- tance between rows, up to four times as long at base of decHvity, slightly shorter on lower declivity. Female.— Similar to male except frons convex, median fovea indefinite, surface strongly reticulate, rather coarsely, closely punctured; tuft of hair on scape absent; setae on elytra slightly shorter. Type Locality.— Caparrapi, Cunadin, Colombia. Type Material.— The male holotype, fe- male allotype, and eight paratypes were taken at the type locality on 28- V- 1959, from Arbol de Sangregao, by A. Diaz. The holotype, allotype, and paratypes are in my collection. Phloeotrihiis fici, n. sp. This species is distinguished from hirtus Wood by the smaller size, by the more shal- lowly impressed frons, by the shorter elytral setae, and by the coarser declivital tu- bercles. Male.— Length 1.6 mm, 2.0 times as long as wide; color black. Frons shallowly impressed, almost flat, from epistoma to upper level of eyes, later- al margins weakly elevated at level of an- tennal insertions, an elongate fovea at middle; surface reticulate, punctures moder- ately fine, not close, about uniformly dis- 394 Great Basin Naturalist Vol. 37, No. 3 tributed; vestiture fine, short, inconspicuous. Pronotum about as in hirtus except punc- tures larger, asperities almost obsolete, ves- titure mostly abraded. Elytra about as in hirtus except strial punctures smaller, interstriae slightly wider than striae, interstrial tubercles on disc more rounded, those on odd-numbered de- clivital interstriae larger (those on 1 fine, extend to near apex, on 2 five extend to middle, on 3 two at base of declivity, on 7 three on basal third, on 8 four at base, on 9 strongly, acutely elevated, coarsely serrate, about nine serrations). Vestiture of rows of rather coarse, interstrial bristles, short on basal half, longest setae near declivity about as long as distance between rows, absent on even-numbered declivital interstriae. Type Locality.— Universidad de los Andes Campus, Merida, Merida, Venezuela. Type Material.— The male holotype was taken at the type locality on ll-IX-1969, 1700 m, No. 5, from Ficus bole, by me. The holotype is in my collection. Pityophthorus discretiis, n. sp. This species is distinguished from perotei Blackman by the larger average size, by the reticulate pronotum which lacks tubercles on the disc, by the more coarsely punctured frons, and by the steeper, more broadly sul- cate declivity. Female.— Length 2.0 mm (paratypes L6- 2.2 mm), 2.9 times as long as wide. Frons planoconvex as in perotei except more coarsely punctured, vestiture shorter (about two-thirds as long). Pronotum about as in perotei except pos- terior areas rather strongly reticulate, punc- tures moderately coarse, deep (tubercles ab- sent). Elytra about as in perotei except disc less shining, almost subreticulate in some speci- mens; declivity slightly steeper, sulcus slightly deeper and wider. Male.— Similar to female except frontal vestiture shorter and apparently less abun- dant. Type Locality.— Three miles or 5 km W El Salto, Durango, Mexico. Type Material.— The female holotype, male allotype, and one paratype were taken at the type locality on 7-VM965, 2500 m, No. 33, Pinus ayacahuite, by me. Other paratypes taken in Mexico by me include: 2 at 29 km or 18 miles W Quiroga, Michoa- can, ll-VI-1965, 2300 m. No. 70, Pinus; 1 at 9 km or 6 miles E Volcan Paricutin, Ja- lisco, 19- VI- 1965, 2500 m. No. 89, Pinus; 1 at 25 km or 14 miles W Texmelucan, Puebla, 14-Vn-1953, 2800 m, Pinus. The holotype, allotype, and paratypes are in my collection. BIOMASS OF FIFTY CONIFER FORESTS AND NUTRIENT EXPORTS ASSOCIATED WITH THEIR HARVEST' T. Weaver and F. Forcella' Abstract.— Biomasses of climax Rocky Mountain forests studied ranged from less than 50 to more than 300 tons/ha. Total biomass was approximately 1.5 times the biomass of normally merchantable boles. When com- pared with conventional bole harvest, the nutrient exports associated with harvest of all aboveground parts in these stands would apparently be at least three times higher for nitrogen, six times higher for phosphorus, four times higher for potassium, and three times higher for calcium. Tlie volume or biomass of a tree can be predicted from its diameter or from its di- ameter and height. Foresters routinely esti- mate the volume of logs using these dimen- sions and tables calculated on the assumption that tree trimks are truncated cones or parabolas (Forbes and Meyer 1961). Weights can then be estimated by multiplying their volumes by the density of the wood. In most organisms the size of one organ system is directly related to the size of other organ sy.stems (allometry, Sinnott 1960), so one can predict the volumes or weights of whole organisms from simple measurements. This has been demonstrated for a wide variety of trees including those of the conifer genera Abies, Finns, and Pseudotsuga (Ovington 1957, Ovington and Madgwick 1959, Kimura 1963, Baskerville 1965, Whittaker and Woodwell 1968, Moir 1972, Odegard 1974, Whittaker et al. 1974, and Long and Turner 1975). This paper presents a single equation which predicts total aboveground biomass of a tree, from its diameter, for any of the seven most common tree species of the northern Rocky Mountains. A comparison of this equation with others reported in the lit- erature emphasizes the similarity of diame- ter biomass relationships among conifers in general. Various shrubs may also have sim- ilar size-weight relationships (Weaver 1977). An equation predicting weight from mea- sures of both height and diameter is similar to published conifer D-H-weight relation- ships but less accurate than the diameter- weight relationship. Similar relationships are presented for weights of boles, branches, twigs, and leaves. The biomass-DBH relationship found was applied to the stand tables of Daubenmire and Daubenmire (1968) to estimate the aboveground arboreal standing crops of fifty forest stands representing the following cli- max associations: Pimis ponderosa-Festuca idahoensis, F. ponderosa-Sijmphoricarpos albiis, F. ponderosa-Fhysocarpiis malvaceus, Fseudotsiiga menziesii-Sytnphoricarpos albiis, F. menziesii-Fhysocarpus malvaceus, F. men- ziesii-Calamagrostis rubescens, and Abies lasiocarpa-Vaccinium scoparium. The nutrient content of the boles and of all aboveground parts was estimated so the nutrient export associated with conventional logging and complete tree utilization could be compared. Methods Thirty-nine trees of six coniferous species and one broadleaf species were felled in southcentral Montana. The trees were sec- tioned into shoot material greater than 10 cm diameter, shoots between one and 10 cm basal diameter, shoots less than 1 cm basal diameter, and leaves. Each class was This study was supported in part by the National Science Foundation grant BMS 73-02027-A02 to the Grassland Biome, U.S. International Biological Programme and U.S. Forest Service Grant 12-11-204-12 for the study of whitebark pine forests. ■'Department of Biology, Montana State University, Bozeman, Montana .59715. 395 396 Great Basin Naturalist Vol. 37, No. 3 weighed wet. Wet weights were converted to dry weights by multiplying by a con- version factor determined for each com- ponent of each tree. The conversion factor was computed as follows: dry weight (60 C to constant weight)/wet weight. The tree species and the numbers of trees per species sampled were Abies lasiocarpa (Hook.) Nutt., 7; Picea engelmannii (Parry) Engelm., 3; Finns contorta var latifoUa Engelm., 4; Finns ponderosa Dougl., 6; Finns albicanlis Engelm., 7; Fsendotsnga menziesii (Mirb.) Franco., 5; and Fopnhis tremnloides Michx., 7. The trees sampled ranged from 4 to 54 cm in diameter at breast height (D at 1.35 m) and from 3 to 20 m in height (H). The trees came from a wide variety of environ- mental conditions ranging from dry to moist sites and from open to closed stands. A list of the trees and their habitat types, sizes, weights, and ages is available from the au- thors. After a preliminary graphical analysis, all data were pooled, and simple and multiple regression analyses were made to determine which models best predicted tree biomass, independent of species. These included lin- ear, curvilinear, and log linear regressions of 12 dependent variables (including weight and logarithms of weights of total shoot and its components) against the independent variables D, D-, DH, and D^H. D--biomass relationships were used to es- timate the aboveground arboreal biomasses of 50 forest stands for which Daubenmire and Daubenmire (1968) have published stand tables. Since their tables list tree num- bers by 10-cm-diameter size classes, tree weights were estimated by two different methods. 1) The weight of all trees in a plot were summed. These weights were esti- mated by assuming that the diameter of each tree was equal to that of the midpoint of the 10-cm-diameter class in which it was recorded. 2) The weights of all trees in a plot were summed. These weights were esti- mated by pooling all plot data for a given habitat type on the assumption that it rep- resented a single (abstract) forest; assigning trees in this sample to 2.5-cm-size classes from a balanced distribution of tree size vs tree number (Meyer 1953); and estimating the weights of the trees as if their diameters were equal to the midpoint of the size classes. Regressions through logarithmically transformed data are said to underestimate biomass (Baskerville 1972); we used uncor- rected values because graphical analysis sug- gests that the imcorrected values represent our data better than corrected values do (cf Whittaker et al. 1974). Results and Conclusions Linear regressions of log^oD- and logioD'H against log weight generally pre- dict biomasses of Rocky Mountain trees well (Table 1 and Fig. la). (1) Both inde- pendent variables predict total biomass well (r- = 0.98 and 0.96, respectively). LogioD'H may be the safer predictor if our regressions are used in habitat types where taller trees than ours may be found. (2) LogioD'H vs logio weight is the best pre- dictor for woody material with diameters greater than 1 cm or greater than 10 cm (r- = 0.98 and 0.95, respectively). This is likely so because the conic or parabolic models of stem volume are good, weights are strongly correlated with volumes, most of the weight is in the larger classes, and branch volume is both correlated with and overridden by stem volume. (3) Log^oD- vs logi ight is the best predictor for leaves and branches with diameters less than 1 cm (r- = 0.84 and 0.79, respectively). This may be so because the capacities of current transporting tissues is highly correlated with diameter (cf Shinozaki et al. 1964). A graphical analysis of the leaf weight-diame- ter relationship suggests that the relatively low r' is partly due to between-species dif- ferences. (4) A regression of untransformed weights against D -I- D- was our best pre- dictor of branch weights (1 to 10 cm, r- = 0.97). This curve, not shown, is J-shaped be- cause in small trees the bole is in the branch-size class. (5) Regressions of untrans- formed variables (D, D\ DH, D-H) vs un- transformed biomass were not linear, were not used, and are not presented. Our regressions of logigD- or log^pD-H against log^o weight are remarkably similar to, but not identical with, those determined Sept. 1977 Weaver, Forcella: Conifer Biomass 397 in other regions (Fig. lb); predictions of weights from such relationships apparently become poorer as one considers species and environments increasingly different from those for which the relationships were de- veloped. The differences may be due to dif- ferences in form factor, wood density, branching patterns, and height-diameter relationships. For example, the tendency of deciduous trees (Monk 1970, Whittaker and Woodwell 1968) to outweigh conifers of similar diameters could be due to the great- er wood densities and branchiness of the de- ciduous trees. The aboveground arboreal biomasses of 50 stands are summarized in Fig. 2; the val- 500 ■D CO CO < o 5 10 DBH (cm) DBH (cm) 5 10 50 or D'H/IO' (cm' m) Fig. 1. Biomass-dimension relationships for coniferous species. (A) Diameter-biomass relations for seven species of the northern Rocky Mountains: a = total biomass, b = stem biomass (diameters greater than 1 cm), c = bole biomass (diameters greater than 10 cm), d = twig biomass (diameters less than 1 cm), e = branch biomass (di- ameters 1-10 cm), and f = leaf biomass. (B) PubHshed DBH-biomass and D'H-biomass relationships: a = Picea rubens (Whittaker et al. 1974), b = Pinus rigida (Whittaker and Woodwell 1968), c = seven species (Fig. la), d = Pinus contorta (Odegard 1974), e = Picea rubens (Whittaker et al. 1974), f = Pseudotsuga menziesii (Long and Turner 1975), g = Pinus contorta (Moir 1972), h = Pinus sylvestris (Ovington and Madgwick 1959), i = Pinus rigida (Whittaker and Woodwell 1968), j = seven species (Fig. la), k = Abies balsamea (Baskerville 1965),1 = Abies veitchii (Kimura 1963), and m = Pinus sylvestris (Ovington 1957). 398 Great Basin Naturalist Vol. 37, No. 3 lies presented were calculated by assuming that tree diameters were equal to the mid- points of their 10-cm-size classes, i.e., by method 1. One may conclude (1) that total arboreal biomasses of climax conifer stands of the northern Rocky Mountains are gener- ally in the 50-300 tons/ha range and (2) that those of specific forest series or associ- ations generally have a narrower range: Daubenmire's Pinus ponderosa-Festuca ida- hoensis stands had biomasses of 50-150 tons/ha; his P. ponderosa-Physocarpus mal- vaceus, Pseudotsuga menziesii-Sijin- phoricarpos albus, and Abies lasiocarpa-Vac- cinium scopariwn stands had biomasses of 100-250 tons/ha and his P. menziesii-Phys- ocai'pus malvaceus and P. menziesii-Calama- grostis nibenscens stands had biomasses of 100-350 tons/ha. One metric ton/ha = 1000 kg/ha. These estimates of standing crop do not include understory biomasses (perhaps 0.1 to 3 percent or possibly 7 per- cent of the arboreal biomasses: e.g., Oving- ton 1962, Whittaker et al. 1974, Tappeiner and John 1973) or belowground biomasses (perhaps 20-30 percent or possibly 7-50 per- cent of the arboreal biomasses: e.g., Oving- ton 1962, Rodin and Bazilevich 1965, Whittaker and Woodwell 1968). Despite the exclusion of understory and underground biomasses, the standing crops reported are larger than most of those re- ported in the literature (30-70 tons/ha; Ro- din and Bazilevich 1965, Art and Marks 1971, Whittaker et al. 1974). Reasons for the large standing crops observed are prob- ably two: Daubenmire selected stands much older than those commonly studied, and he selected the parts of those stands with the biggest trees. The accuracy of volume and biomass esti- mates can usually be increased slightly by recording narrower diameter classes (Meyer 1953). To achieve this we pooled the data from all stands occupying a given habitat type on the assumption that it represented multiple stands of one (abstract) forest; de- termined balanced size distributions (Meyer 1953), allocated trees to 2.5-cm-diameter classes, and estimated their weights by using Table 1. Regression constants and indices of error used in relating sizes and weights of important trees of the northern Rocky Mountains. Total Wood Wood Wood Wood Notes' Tree >1 cm > 10 cm 1-10 cm <1 cm Leaves D^ a 2 -1.05 -1.18 -1.40 0.60 -1.87 -1.91 b 2 1.19 1.21 1.25 0.75 1.16 1.13 r' 5 0.98 0.97 0.89 0.81 0.79 0.84 E 6 1.26 1.34 1.47 1.60 2.23 1.91 D^H a 3 1..35 -1.53 -1.84 -0.73 -1.99 -2.08 b 3 0.93 0.96 1.01 0.57 0.86 0.84 r- 5 0.96 0.98 0.95 0.76 0.69 0.77 E 6 1.36 1.27 1.30 1.72 2.64 2.19 D + D^ a 4 -4..51 + 0.08 -29.36 + 29.37 -3.02 + 1.16 b 4 -3.32 -2.96 + 0.71 -3.69 -0.57 + 0..39 c 4 + 0.85 + 0.71 + 0.49 + 0.21 + 0.15 + 0.06 r- 5 0.96 0.96 0.94 0.97 0.63 0.56 e 6 0.29 0.38 0.25 1.14 0.28 1.00 'Identification of symbol!; to the left. ■LxjgioWT = a + b logjo D' with weight in kg and diameter in cm. Log,,, WT = a + b logiu D'H with weight in kg, diameter in cm, and height in m. VVT = a + bD + cD' with weight in kg and diameter in cm. r is the correlation coefficient, r-' is the fraction of the variation explained by the regression. E is the antilog of the standard error of a logarithmic regression; expected errors lie between the predicted value x E and the predicted value/E. An estimate of relative error (e) is the standard error of a linear regression divided by the average weight (y) used (Whittaker and Woodwell 1968). Sept. 1977 Weaver, Forcella: Conifer Biomass 399 the midpoints of those classes as diameters (method 2). Forest weights estimated by method 2 ranged from 80 to 98 percent of the mean of stand weights determined with 10-cm-diameter classes (i.e., by method 1). They were: Pinus ponderosa-Festuca ida- hoensis 75 tons/ha, 80 percent; P. ponde- rosa-SympJioricai-pos albus 155 tons/ha, 82 percent; P. pondewsa-Physocarpos mal- vaceus 140 tons/ha, 98 percent; Pseitdot- suga menziesii-Symphoricarpos albus 180 tons/ha, 98 percent, P. menziesii-Calama- grostis ruhescens 336 tons/ha, 91 percent; and Ahies kisiocarpa-Vaccinium scoparium 178 tons/ha, 93 percent. 350 '^ 200 o < ^ 150 100 '^^ eP D «« . c t* ♦♦ ■a d ♦ ♦ 10 20 30 40 50 60 70 80 90 BASAL AREA (m2/ha) Fig. 2. Aboveground arboreal standing crops of 50 forest stands representing 7 habitat types: Piniif; pon- derosa-Festuca idahoerisis (A), P. ponderosa-Sym- phoricarpos alba (B), P. ponderosa-Physocarpus mal- vaceus (C), Pseitdotsiiga menziesii-Symphoricarpos alba (D), P. menziesii-Physocarpus malvaceus (E), P. men- ziesii-Calamatrostis rtibescens (F), and Abies kisiocarpa- Vaccinium scoparium (G). Calculations include all trees with DBH's greater than 5 cm; basal areas of stands 66, 101, 149, 173, 174, and 179 were recalcu- lated from stand table data. Biomasses in the leaf, twig, branch, and bole compartments are very nearly (SE ± 1 percent) 9 percent, 13 percent, 13 percent, and 66 percent of the arboreal biomass in the mature forests studied. Total harvest in most Rocky Movmtain forests should there- fore equal approximately 1.5 times the nor- mal (10 cm +) stem harvest (Fig. 2). Whittaker et al. (1974) give similar values. This is a consequence of both the allometric equations used and of the size class distribu- tions observed in the stands studied. Mineral nutrient export associated with total harvest is greater than 1.5 times that associated with normal bole harvest because needles and bark (which comprise a larger fraction of twigs and branches than of boles) are relatively rich in mineral ele- ments. Table 2 simimarizes published values for nitrogen, phosphorus, potassium, and calcium contents for needles, twigs, branches, and boles. One can estimate the amount of a nutri- ent element exported from a harvested stand by adding up nutrient export associ- ated with each component exported (Table 2). The nutrient export associated with har- vest of any component equals the total ex- port X the proportion (percent) of the total due to the component considered (e.g., nee- dles or bole) x the nutrient content of that component. Since needles, twigs, branches, and boles comprise relatively constant pro- portions of total abovegrovmd arboreal bio- mass (9, 13, 13, and 66 percent, respective- ly), we may conclude that nutrient export associated with total harvest— as opposed to bole harvest alone— is three to six times greater for nitrogen, six to nine times great- er for phosphorus, four times greater for potassium, and three times greater for cal- ciimi. Thus, total harvest of a Rocky Moun- tain conifer stand with a relatively low aboveground arboreal biomass of 100 tons/ha might remove 160-460 kg/ha of ni- trogen, 20-40 kg/ha of phosphorus, 80-180 kg/ha of potassium, and 120-500 kg/ha of calcium. Bole harvest alone would remove only 25-160 kg/ha of nitrogen, 2-6 kg/ha of phosphorus, 20-40 kg/ha of potassium, and 40-200 kg/ha of calcium. Exports from for- 400 Great Basin Naturalist Table 2. Typical nutrient contents of conifers. ^'^ Vol. 37, No. 3 Nitrogen kg/ % loot Phosphorus % kg/ loot Potassium % kg/ loot Calcium % kg/ loot Needles max 1.34 121 0.22 20 0.78 70 1.05 95 min 0.84 76 0.08 7 0.28 25 0.15 14 Twigs max 0.60 78 0.07 9 0.30 .39 1.10 143 min 0.31 40 0.05 7 0.14 18 0.27 35 Branches max 0.77 100 0.06 8 0.25 32 0.54 70 min 0.15 20 0.02 3 0.12 16 0.28 36 Bole max 0.24 158 0.01 7 0.06 40 0.30 198 min 0.04 26 0.00 2 0.03 20 0.06 40 Total max 457 43 181 506 min 162 18 79 125 'Percentage compositions reported are the highest and lowest values reported for a variety of conifers {Larix, Picea. Pinus. and Pseudotsuga) in papers by Odegard (1974), Overton et al. (1974), Stark (1973), and Rodin and Bazilevich (1965). 'Kg/lOOt contents refer to 100 tons of aboveground standing crop. They were calculated by multiplying 100 tons of standing crop x the approximate percentage of the standing crop contributed by the organ (9 percent needles, 13 percent twigs, 13 percent branches, and 66 percent boles) x the per- centage of the organ contributed by the nutrient considered (N, P, K, or Ca). est twice to three times as productive would be twice to three times as great. One wonders whether the forest product value of 'slash' will exceed its fertilizer val- ue. If inputs from nitrogen fixation process- es or from decomposition of rocky sub- strates are small, the fertilizer value of branch material with diameters less than 10 cm might exceed its forest product value. Literature Cited Art, H., and P. Marks. 1971. A summary table of bio- mass and net annual production in forest ecosys- tems of the world. In: International Union of Forest Research Organizations. Forest biomass studies. College of Life Science and Agriculture, University of Maine, Orono. 240 pages. Baskerville, G. 1965. Estimation of dry weight of tree components and total standing crop in conifer stands. Ecology 46: 867-869. 1972. Use of the logarithmic equation in the estimation of plant biomass. Can. J. Forestry Re- search 2: 49-53. Daubenmire, R., and J. Daubenmire. 1968. Forest vegetation of eastern Washington and northern Idaho. Washington Ag. Expt. Sta. Tech. Bull. 60, Pullman. 104 pages. Forbes, R., and A. Meyer. 1961. Forestry Handbook. Ronald Press, New York. 1200 pages. KiMURA, M. 1963. Dynamics of vegetation in relation to soil development in northern Yatsugatake Mountains. Japanese J. Bot. 18: 255-287. KiRA, T., and T. Shidei. 1967. Primary production and turnover of organic matter in different forest eco- systems of the western Pacific. Jap. J. Ecol. 17: 70-87. Long, J., and J. Turner. 1975. Aboveground biomass of overstory and understory in an age sequence of four Douglas fir stands. J. Applied Ecol. 12: 179- 188. Meyer, H. 1953. Forest mensuration. Penns. Valley Pub. State College, PA. 357 pages. MoiR, W. 1972. Litter, foliage, branch, and stem pro- duction in contrasting lodgepole pine habitats of the Colorado Front Range. In: J. Franklin, L. Dempster, and R. Waring (eds). Research on Coniferous Forest Ecosystems. USDA Forest Ser- vice, Pacific NW Forest and Range Expt. Sta., Portland, Oregon. 322 pages. Monk, C, G. Child, and S. Nicholson. 1970. Bio- mass, litter, and leaf surface estimates of an oak- hickory forest. Oikos 21: 138-141. Odegard, G. 1974. Nutrient and biomass distribution following clearcutting in a lodgepole pine forest ecosystem. Unpublished Ph.D. thesis, Colorado State University, Fort Collins. 182 pages. Overton, W., D. Lavender, and R. Hermann. 1973. Estimation of biomass and nutrient capital in stands of old-growth Douglas fir. In: H. Young (ed.) lUFRO Biomass Studies. College of Life Sci- ences and Agriculture, Univ. of Maine, Orono. 532 pages. Ovington, J. 1957. Dry matter production by Pinus sylvestris L. Ann. Bot. NS 21: 287-314. 1962. Quantitative ecology and the woodland ecosystem concept. In: J. Cragg (ed.). Advances in Ecological Research 1: 103-192. Ovington, J., and H. Madgwick. 1959. Distribution of organic matter and plant nutrients in a plantation Sept. 1977 Weaver, Forcella: Conifer Biomass 401 of Scotch pine. Forest Science 5:344-355. Pfister, R., B. KovALCHiK, S. Arno, and R. Presby. 1974. Forest habitat types of Montana. USDA, Forest Service, Intenntn. Forest and Range Expt. Sta., Ogden, Utah. 213 pages. Rodin, L., and N. Bazilevich. 1965. Production and mineral cycling in terrestrial vegetation. Oliver and Boyd, Edinburgh. 288 pages. Shinozaki, K., K. Yoda, K. Hozumi, and T. Kira. 1964. A quantitative analysis of plant form: the pipe model theory. Japanese J. Ecol. 14: 97-105. SiNNOTT, E. 1960. Plant Morphogenesis. McGraw Hill, New York. 550 pages. Stark, N. 1973. Nutrient cycling in a Jeffrey Pine eco- system. University of Montana, Institute for Mi- crobiology, Missoula. 389 pages. Tappeiner, J., AND H. JoHN. 1973. Biomass and nutri- ent content of hazel undergrowth. Ecology 54: 1342-1348. Weaver, T. 1977. Area-weight relationships for com- mon Montana shrubs. Proc. Mont. Acad, of Sci. 37: in press. Weaver, T., and D. Dale. 1974. Pinus albicaulis in central Montana: environment, vegetation, and production. Am. Midland Naturalist 92: 222-230. Whittaker, R., and G. Woodwell. 1968. Dimension and production relations of trees and shrubs in the Brookhaven Forest, New York. J. Ecol. 44: 233-252. Whittaker, R., F. Bormann, G. Likens, and T. SiccAMA. 1974. The Hubbard-Brook ecosystem study: forest biomass and production. Ecol. Mono. 44: 233-252. THE ALGAL FLORA OF THE CAMPUS OF BRIGHAM YOUNG UNIVERSITY, PROVO, UTAH Cheng Mou-Sheng' and Samuel R. Rushforth' Abstract.— The algal flora of the Brigham Young University campus is more diverse than previously thought. Sixty-eight genera containing 160 species of Chlorophyta, Euglenophyta, Chrysophyta, and Cyanophyta were col- lected and identified. The greatest number of species was obtained from a small pond in the arboretum, with progressively fewer species obtained in the irrigation canal partly surrounding the campus and in specialized hab- itats in the greenhouses. Taxonomic and ecological studies of algae in the Intermountain West and Utah in par- ticular have been rare. The earliest papers on Utah algae concerned the flora of the Great Salt Lake (Rothpletz 1892, Talmage 1900, Daines 1917). This unusual habitat has continued to be of interest to the present time and is currently under study by several biologists and water quality specialists. Other early papers on Utah algae were those of Norrington (^1925) on the algae of the lakes and streams of the Uinta and Wasatch Mountains of Utah; Harrison (1926) on the algae of Washington County, Utah; Tanner (1930, 1931) on the algae of Utah Lake; Snow (1932) on the algae of Utah Lake; Kirkpatrick (1934) on the biolo- gy of the Great Salt Lake; and Patrick (1936) on the diatoms in core samples from the edge of the Great Salt Lake. The algal flora found on the Brigham Young University campus has been under observation for several years, particularly for teaching purposes. However, nothing has been written concerning this flora until recently. The most important contribution to a knowledge of this flora was made by the senior author of the present paper in a study during 1971-1972 for the research re- quirement for the Master of Science degree (Mou-Sheng 1973). The junior author has continued to collect algae from the campus since that time. Methods Several collecting stations were estab- lished at selected sites on the Brigham Young University campus. Sites were estab- lished in the arboretum pond, the irrigation canal transversing the campus, and in the Department of Botany and Range Science greenhouses. Phytoplankton, attached algae, and algae in the sediments were all sam- pled. In addition several physical and chem- ical parameters were measured at the aquatic sites in order to provide an overall picture of the environment. Algae were returned to the laboratory, subsampled, and examined immediately for nondiatom algae. These were studied using a Zeiss RA microscope with Nomarski inter- ference phase accessories. Algae were iden- tified using standard reference texts. Following study of the nondiatoms, per- manent diatom slides were prepared by standard boiling nitric acid tenchiques. Dia- toms were mounted in pleurax diatom mountant. These slides are in the Brigham Young University diatom collection. Dia- toms were examined with the same equip- ment cited above and identified by us, using standard reference texts. Results Sixty-eight genera and 160 species of al- gae have been identified and described from Department of Botany and Range Sci Jrigham Young University 402 Sept. 1977 Mou-Sheng, Rushforth: Algal Flora 403 the campus. This flora consists of 27 genera containing 36 species of Chlorophyta, 2 genera containing 2 species of Eu- glenophyta, 32 genera containing 106 spe- cies of Bacillariophyceae, 1 genus contain- ing 2 species of Chrysophyta excluding diatoms, and 6 genera containing 14 species of Cyanophyta. Tlie Brigham Young University arboretum pond affords an excellent habitat for the growth of algae. This pond contains high levels of dissolved silica (27-81 PPM), bi- carbonates, carbon dioxide (8-68 PPM) and oxygen (5-12 PPM) which support a high population of diatoms throughout the win- ter and spring and Chlorophyta (especially Spirogyra diibia Kutzing, S. jitgolis [Dan.] Kutzing, S. neglecta [Hass.] Kutzing and S. porticalis [Muell.] Cleve) through the late spring and summer. The diatom flora of the arboretum pond is dominated by Melosira varians Agardh, Sijnedm parasitica var. subconstricta (Grun.) Hust., S. ulna (Nitz.) Ehr., S. ulna var. siib- aequalis (Grun.) v. Heur., Cocconeis pedi- cuhts Ehr., C. placentula var. lineata (Ehr.) V. Heur., Anomoeoneis spliaerophora (Kutz.) Pfitz., Cijmbelki cistula (Hemprich) Grun., Nitzschia linearis W. Sm., N. sigmoidea (Ehr.) W. Sm., and Cijmatopleura solea (Breb.) W. Sm. Several species of diatoms, especially of Navicula, CymbeUa, and Nitzschia have been found only in this pond on campus. These include Navicula cuspidata (Kutz.) Kutz., N. odiosa Wallace, N. placentula var. rostrata Mayer, N. pupula Kutz., N. pyg- maea Kutz., CymbeUa heteropleura (Ehr.) Kutz., C. mexicana (Ehr.) Schmidt., C. pro- strata (Berk.) CI., C. tumida (Breb.) v. Heur., C. twnidula Grim., C. turgida Greg., Nitzschia hungarica Grun., N. sigma (Kutz.) W. Sm., and N. tryblionella Hantz. Other species restricted to this locality were Melo- sira distans (Ehr.) Kutz., Fragilaria brevist- riata var. inflata (Pant.) Hust., Synedra capitata Ehr., Eunotia curvata (Kutz.) Lagerst, Caloneis ventricosa (Ehr.) Meist., Neidiuni iridis (Ehr.) CI., Goniphonema acu- minatum Ehr., Epithemia turgida (Ehr.) Kutz., E. turgida var. granulata (Ehr.) Gnm., and Rhopalodia gibba (Ehr.) O. Mull. The Brigham Young University botanical greenhouses represent rather specialized en- vironmental conditions. Thus, several pa- rameters such as light, water, and temper- ature are controlled and demonstrate little fluctuation. The algal flora of the green- houses is dominated by Protococcus viridis Agardh, Chlorococcum humicola (Naeg.) Ra- benhorst, Oscillatoria sancta (Kutz.) Go- mont. Amphora normani Rabh., and Han- tzschia amphioxys (Ehr.) Grun. Several species of algae have been co- lected on campus only from the green- houses. These include Lyngbya aestuarii (Mertens) Liebmann, L. martensiana Men- eghini, Oscillatoria angustissima West and West, O. animalis Agardh, O. cruenta Grim., O. limosa (Roth) Agardh, O. sancta (Kutz.) Gomont, O. tenuis Agardli, Phormi- dium inundatum Kutzing, Anabaena varia- bilis Kutzing, Tolypothrix penicillata (Ag.) Thur., Sticlwcoccus bacillaris Naegeli, S. scopulinus Hazen, S. subtdis (Kutz.) Kler- cker, and Chlorella vulgaris Beyerinck. Sixteen genera containing 33 species of diatoms have been collected from the soil in Table 1. Number of species of algae on the Brigham Young University campus by algal division and collection locality. Division Arboretum Pond Collection Locality Campus Stream Greenhouses Chlorophyta Euglenophyta Chrysophyta Cvanophyta TOTAL 404 Great Basin Naturalist Vol. 37, No. 3 the campus greenhouses. Hantzschia am- phioxijs (Ehr.) Grun. and Amphora normani Rabh. are the most common soil diatoms. Achnanthes lanceolata (Breb.) Grun., Nav- icula tripunctata (Mull.) Bory, Hantzschia amphioxys var. capitata Mull., and Nitzschia palea (Kutz.) W. Sm. are also quite com- mon. Pinnularia gentilis (Donk.) CI. is the only diatom restricted to the soil. The water in the irrigation canal that crosses the Brigham Young University cam- pus generally has higher levels of nitrates, phosphates, dissolved oxygen, alkalinity, and carbon dioxide than the arboretum pond. It contains near the same number of species of algae but has a lower standing crop due to the paucity of filamentous Chlorophyta. The flora of this stream is dominated by Stepluinodiscus niagare Ehr., Diatoma vul- gare Bory, D. tenue var. elongatum Lyngb., Cocconeis pedicuhis Ehr., C. placentula var. eughjpta (Ehr.) CI., Naviciiki tripunctata (Mull.) Bory, Gornphonerna oHvaceiim (Lyn- bye) Kutz., and Nitzschia sigmoidea (Ehr.) W. Sm. Species found only in this stream include Chaetophora incrassata (Huds.) Hazen, Dia- tonia hiemale var. mesodon (Ehr.) Grun., Gornphonerna acuminatum var. coronatum (Ehr.) W. Sm., G. angustata var. sarco- phagus (Greg.) Grun., Hannaea arcus var. amphioxys (Rabh.) Patr., Navicula capitata Ehr., Navicula laevissima Kutz., N. mutica Kutz., Neidium affine (Ehr.) Pfitz., Palmella mucosa Kutz., and Synedra ulna var. con- stricta Ostr. The following list contains those algae collected on the Brigham Young University campus. Chlorophyta Chlorophyceae Chlorococcales Chlorococcaceae Characitim ambiguum Hermann Chlorococcum humicola (Naeg.) Rabenhorst Palmellaceae Palmella mucosa Kutzing Sphaerocystis schroeteri Chodat Oocystaceae Chlorella vulgaris Beyerinck Scendesmaceae Scenedesmus denticulatus Lagerheim S. quadricauda var. quadrispina (Chod.) G. M. Smith Hydrodictyaceae Pediastrutn boryanum (Turp.) Meneghini P. tetras (Ehreb.) Ralfs Ulotrichales Protococcaceae Protococcus viridis C. A. Agardh Ulotrichaceae Stichococciis bacillaris Naegeli S. scopulinus Hazen S. subtilis (Kutz.) Klercker Ulothrix zonata (Weber & Mohr) Kutzing Chaetophorales Aphanochaetaceae Aphanochaete repens A. Braun Chaetophoraceae ChaeUyphora incrassata (Huds.) Hazen Stigeoclonium lubricum (Dillw.) Kutzing Oedogoniales Oedogoniaceae Oedogonium sp. Cladophorales Cladophoraceae Cladophora glomerata (L.) Kutzing Zygnematales Zygnemataceae Mougeotia genuflexa (Dillw.) C. A. Agardh Spirogyra dubia Kutzing S. jugalis (Fl. Dan.) Kutzing S. neglecta (Hass.) Kutzing S. porticalis (Muell.) Cleve Zygnema insigne (Hass.) Kutzing Desmidiaceae T.\BLE 2. Number of species of algae on the Brigham Young University campus restricted to selected collection localities. Algal Division Collection Locality Arboretum Pond Campu > Stream Greenhouses 4 8 0 0 18 3 0 12 22 23 Chlorophyta Euglenophyta Chrysophyta Cvanophyta TOTAL Sept. 1977 Mou-Sheng, Rushforth: Algal Flora 405 Closterium lanceolatttm Kutzing C. moniliferum (Bory) Ehrenberg Cosmariuvi hotrytis Meneghini C. perforatum Lund. Penium navicula Breb. EUGLENOPHYTA Euglenophyceae Euglenales Euglenaceae Eiiglena aciis Ehrenberg Phacus acuminata Stokes Chrysophyta Xanthophyceae Vaucheriales Vaucheriaceae Vaucheria geminata (Vaucher) DeCandolle V. sessilis (Vaucher) DeCandolle Bacillariophyceae Centrales Coscinodiscaceae Melosira distans (Ehr.) Kutz. M. granulata (Ehr.) Ralfs M. italica (Ehr.) Kutz. M. varians C. A. Ag. Cyclotella bodanica Eulenst. C. meneghiniana Kutz. Stephanodiscus niagare Ehr. Pennales Fragilariaceae Asterionella formosa Hassall Hannaea arcus var. amphioxys (Rabh.) Patr. Diatoma anceps (Ehr.) Kirch. D. hiemale var. mesodon (Ehr.) Grun. D. tenue var. elongatum Lyngb. D. vulgare Bory Fragilaria brevistriata var. inflata (Pant.) Hust. F. capucina var. mesolepta Rabh. F. construens var. venter (Ehr.) Grun. F. crotonensis Kitton F. leptostauron (Ehr.) Hust. F. vaucheria (Kutz.) Peters. Meridian circukire var. constrictum., (Ralf.) v. Heur. Synedra acus Kutz. S. capitata Ehr. S. fasciculata (Ag.) Kutz. S. parasitica (W. Sm.) Hust. S. parasitica var. subconstricta (Grun.) Hust. S. rumpens Kutz. S. idna var. subaequalis (Grun.) v. Heur. S. ulna var. constracta Ostr. S. idna var. rarnesi (Herib.) Hust. S. ulna var. ulna (Nitz.) Ehr. Eunotiaceae Eunotia curvata (Kutz.) Lagerst Achnanthaceae Achnanthes lanceolata var. dubia Grun. A. lanceolata var. lanceolata (Breb.) Grun. A. minutissima Kutz. Cocconeis pediculus Ehr. C. placentula var. euglypta (Ehr.) CI. C. placentula var. lineata (Ehr.) v. Heur. Rhiocosphenia curvata (Kutz.) Grun. ex Rabh. Naviculaceae Anomoeoneis sphaerophora (Kutz.) Pfitz. Caloneis ventricosa (Ehr.) Meist. Diploneis oblongella (Naeg. ex Kutz.) Ross Frustulia vulgaris (Thwaites) DeT. Gyrosigma spencerii (Quek.) Griff. & Henfr. Navicula capitata Ehr. N. cryptoccphala Kutz. N. cryptoccphala var. veneta (Kutz.) Rabh. N. cuspidata (Kutz.) Kutz. N. elginensis (Greg.) Ralfs N. laevissima Kutz. N. lanceolata (Ag.) Kutz. N. mutica Kutz. N. oblonga (Kutz.) Kutz. N. odiosa Wallace N. plancentula var. rostrata A. Mayer A^. pupula Kutz. N. pygmaea Kutz. N. rhynchocephala Kutz. N. tripunctata (O. F. Mull.) Bory Neidimn affine (Ehr.) Pfitz. N. binode (Ehr.) Hust. N. iridis (Ehr.) CI. Pinnularia brebissonii (Kutz.) Rabh. P. gentilis (Donk.) CI. P. viridis (Kutz.) Ehr. Stauroneis smithii Grun. Gomphonemaceae Gomphonerna acuminatum Ehr. G. acuminatum var. coronatum (Ehr.) W. Sm G. angustata var. sarcophagus (Greg.) Grun. G. constrictum Ehr. G. intricatwn Kutz. G. olivaceum (Lyngbye) Kutz. G. olivaceum var. calcarea CI. G. parvulum (Kutz.) Grun. G. parvulum var. micropus (Kutz.) CI. Cymbellaceae Amphora ovalis Kutz. A. nonnani Rabh. Cymbella affinis Kutz. C. as-pera (Ehr.) CI. C. cistula (Hemprich) Grun. C. ehrenbergii Kutz. C. heteropleura (Ehr.) Kutz. C. mexicana (Ehr.) A. Schmidt C. prustrata (Berk.) CI. C. tiimida (Breb.) v. Heur. C. tumidula Grun. C. turgida Greg. C. ventricosa Kutz. Epithemiaceae Epithemia sorex Kutz. £. turgida (Ehr.) Kutz. £. turgida var. granulata (Ehr.) Gnm. Rhopalodia gibba (Ehr.) O. Mull. Nitzschiaceae Hantzschia amphioxys (Ehr.) Gnm. //. amphioxys var. capitata Mull. Nitzschia amphibia Grun. iV. dissipata (Kutz.) Gnm. 406 Great Basin Naturalist Vol. 37, No. 3 N. fonticola Grun. N. httngarica Grun. N. linearis W. Sm. N. palea (Kutz.) W. Sm. N. sigrrui (Kutz.) W. Sm. N. sigmoidea (Ehr.) W. Sm. iV. tryblionella Hantz. Surirellaceae Cipmitopleura solea (Breb.) W. Sm. Surirella angustata Kutz. S. ovalis Breb. S. ovata var. pinnata W. Sm. S. robusta Ehr. Cyanophyta Myxophyceae Chroococcales Chroococcaceae Chroococcus rufescens (Kutz.) Naegeli Oscillatoriales Oscillatoriaceae Lyngbya aestuarii (Mertens) Liebmann L. martensiana Meneghini Oscillatoria amphibia C. A. Agardh O. angustissima West & West O. animahs C. A. Agardh O. cruenta Grun. O. limosa (Roth) C. A. Agardh O. princeps Vaucher O. sancta (Kutz.) Gomont O. tenuis C. A. Agardh Phormiditim imtndatttm Kutzing Nostocales Nostocaceae Anabaena variabilis Kutzing Scytonemataceae Tolypothrix penicillata (Ag.) Thur. Literature Cited Daines, L. L. 1917. On the flora of the Great Salt Lake. Amer. Natur. 51: 499-506. Harrison, J. W. 1926. A preliminary study of the freshwater algae of Washington Co., Utah. M.A. Thesis, Brigham Young University, Provo. KiRKPATRicK, R. 1934. The life of the Great Salt Lake, with special reference to the algae. M.S. Thesis, University of Utah, Salt Lake City. Mou-Sheng, Cheng. 1973. A taxonomic investigation of the algae of the Brigham Young University campus, Provo, Utah. M.S. Thesis, Brigham Young University, Provo. NoRRiNGTON, A. 1925. Phycological study of some of the mountain lakes and streams of the Wasatch and Uinta Ranges in Utah. Ph.D. Thesis, Univ. of Chicago. Patrick, R. 1936. Some diatoms of the Great Salt Lake. Bull. Torrey Bot. Club 63(3): 157-166. Rothpletz, a. 1892. On the formation of oolite. Bot. Centralb. 35. Snow, E. 19.32. A preliminary report of the algae of Utah Lake. Proc. Utah Acad. Sci. 9: 21-28. Talmage, J. E. 1900. The Great Salt Lake, Past and Present. Deseret Book Co., Salt Lake City. Tanner, V. M. 1930. Freshwater biological studies at Utah Lake, Utah. Proc. Utah Acad. Sci. 70: 60-61. 1931. Freshwater biological studies at Utah Lake, Utah, No. 2. Proc. Utah Acad. Sci. 8: 199- 203. OCCURRENCE OF TICKS ON MULE DEER IN CENTRAL UTAH Jordan C. Pederson' Abstract.— Two species of ticks were collected from mule deer and identified as Dertnacentor alhipictus (Packard) and Ixodes sp. The rate of occurrence of these ticks was found to be 99.6 percent and 0.4 percent, re- spectively. The infestation rate increased from 18.2 percent in January, to 87.5 percent in February, to 100.0 per- cent in March. From January through March 1976, a mule deer {Odocoileus hemionus Rafi- nescjue) trapping operation was conducted by Utah State University, the Cooperative Wildhfe Research Unit, and the Utah State Division of Wildhfe Resources. The princi- pal emphasis of this study was the gathering of deer reproduction data. Deer were live- trapped, weighed, and ear-tagged, and a laparotomy was performed on all females. Additionally, a blood sample was taken from each animal. These deer were also checked for external parasites. This paper reports the occurrence of ticks on mule deer in central Utah based on this study. Appreciation is extended to Dr. C. Selby Herrin for his help in the identification of the parasites collected. The help of Paul Tervort, Dick Worthen, and Phil Zwank is gratefully acknowledged. Thanks are also extended to Dr. S. L. Welsh and R. Drob- nick for their critical review of this manu- script. Harris (1945) reported associations of Dennacentor alhipictus (Packard) on mule deer in a South Dakota study dealing with malnutrition in deer. He states, "In a few cases the factors contributing to the death (deer) were predators or minor infestation of internal and external parasites, such as nose bot (Cephenomijia) and winter tick {Dennacentor alhipictus). '^ Edmunds (1951) gives an account of D. alhipictus on deer near Logan, Utah, collected by Clamaine in 1942. Beck (1955) regards D. alhipictus as localized in distribution and a potential vec- tor of Rocky Mountain spotted fever. Rich- ens (1967) found this parasite on mule deer in Daggett County, Utah, where the pro- portion of deer infested with this tick rose from 37 percent in January to 92 percent in March of 1960. This parasite was also re- ported on deer in Garfield, Grand, San Juan, and Wayne coimties in Utah by Pe- derson (1970); deer on the La Sal and Hen- ry mountains were shown to have an in- festation rate of 55 and 75 percent, respectively, or a combined occurrence of 63 percent. Members of the genus Ixodes have not previously been reported on deer in Utah, although they are found as juveniles during winter months in association with small mammals (Edmunds 1951). Methods and Procedures Live trapped deer were systematically ex- amined for external parasites. Those found were removed and placed in isopropyl alco- hol (70 percent). The site of occurrence or attachment was noted. Data recorded for the host animals included sex, age, ear tag number, weight, and an estimate of general body condition. Deer killed by motor ve- hicles were examined as part of this study. An effort was made to collect a representa- tive sample of ectoparasites at each site of attachment on the deer. All other ticks were removed to obtain data on reinfesta- tion rates on live-trapped deer. All deer examined during 1976 were from Utah County, specifically the areas of Long Hollow, Knoll Hollow, Lasson Draw, and Spanish Fork Canyon. Utah Division of Wildlife Resources, Environmental Studies, 1115 .North Main, Sprnigville, Utah 8466.3. 407 408 Great Basin Naturalist Vol. 37, No. 3 Results and Discussion Two species of ticks were collected and identified, Derrnacentor albipictus and Ixodes sp. The rate of occurrence of these ticks on mule deer was 99.6 percent and 0.4 percent, respectively. Dennacentor albi- pictus specimens collected were identified as 35.6 and 57.3 percent adult males and fe- males, respectively; 6.7 percent nymphs, and 0.4 percent larvae (Table 1). Adult fe- males made up the largest percentage of the population for January, with a 66.6 per- cent occurrence rate. Adult males were highest in the population in February at 39.8 percent and lowest in March at 27.1 percent. Nymphs were first collected during February and were highest during March (14.4 percent), with an average occurrence of 6.7 percent during this study (Table 1). Their occurrence rate varied from zero in January to 14.4 percent in March. During this study two female specimens of Ixodes sp. were collected, one in January and one in February. Both were collected in Long Hollow from two female mule deer older than five years. The number of deer infected by ticks var- ied by age class and month. The highest rate of infestation was in the fawn age class, where it averaged 95.1 percent during this study (Table 2). The deer of 1% and 2% age classes were very similar in their monthly and total infestation rates (Table 2). Mature deer (three years and older) had an in- festation rate exceeded only by fawns. They were the highest infested age class for Feb- ruary, with a 87.5 percent infestation rate (Fig. 2). All age classes had a 100 percent infestation during March. The infestation rate for all age classes for the duration of the study was 84 percent. Data gathered on the point of attachment of ticks on each deer showed that three spe- cific points of attachment accounted for 94.6 percent of the ticks. The largest num- ber of D. albipictus were attached around the anus (49.7 percent), the ears were the next highest (39.7 percent), the areas of the flank and udder accounted for 5.3 percent, and the remaining 5.3 percent were at- tached to other parts of the body. Ticks were usually found on the body areas of short and very sparse hair growth. Five of the deer captured during the study were retaken and reexamined during subsequent trapping periods. Three were fawns (1 male, 2 female) and two were ma- ture females. A female fawn trapped in Long Hollow on 24 January 1976 was ex- amined and found to be free of external parasites. When this deer was retrapped on 23 February 1976 three male and eight fe- male were found attached near the anus. She had lost 1.8 kg, a 5 percent loss in body weight, possibly due in part to the presence of ticks. The remaining four deer all had D. albipictus present on their ears and near the anus when first captured. All ticks were removed. When recaptured, all had D. albipictus present on their ears and near the anus. One of these, a female fawn retrapped 16 days after initial capture, had lost 1.4 kg. This is also a 5 percent loss in body weight. The other three deer, one male fawn and two mature does, expe- Table 1. Occurrence on mule deer of Derrnacentor albipictus by developmental stage, sex and month. The per- centage of ticks comprising each sex or life stage for each month appears in parentheses following the number found. Month Adult Male Adult (%) Female (%) Nymph (%) Larva Total January 2 (33.4) 4 (66.6) 0 February 155 (39.8) 220 (56.5) 12 March _53 (27.1) 114 (58.5) 28 Total 210 (35.6) 338 (57.3) 40 (0.0) 0 (0.0) (3.1) 1 (0.,3) (14.4) 0 (0.0) (6.7) 1 (0.4) 195 590 Sept. 1977 Pederson: Ticks on Mule Deer 409 Table 2. Infestation rate of Dermacentor albipictus on mule deer by month and age class. Month Deer Age Fawn P/4 Years 2% Years 3 Years and Over Summary 2 z 5' z 5 i i ^% 2 Z 5; Z 3 i I"! g z 3; z 3 i f"! 2 Z 5' z £ = ST c 3 3 S 3 z.% i 3 3 %S ^l January 2 1 50.0 2 0 0.0 1 0 0.0 6 1 16.7 11 2 18.2 Feiiniarv 30 29 96.6 8 5 62.5 8 5 62.5 38 34 89.4 80 70 87.5 March _9_ _9^ 100.0 _2_ 2_ 100.0 _L 1^ 100.0 16_ i6_ 100.0 28 28 100.0 Total 41 39 95.1 12 7 58.3 10 6 60.0 51 85.0 119 100 84.0 rienced no loss of body weight. During this study no deer mortahty could be attributed to infestation of ticks. Literature Cited Beck, D. E. 1955. Distributional studies of parasitic ar- thopods in Utah, determined as actual and poten- tial vectors of Rocky Mountain spotted fever and plague, with notes on vector-host relationships. Brigham Young Univ. Sci. Bull., Biol. Ser. 1(1) 1- 64. Edmunds, L. R. 1951. A checklist of the ticks of Utah. Pan-Pacific Ent. 27(1): 2.3-26. Harris, D. 1945. Symptoms of malnutrition in deer. J. Wild). Manage. 9(4): 319-322. Pederso.n, J. C. 1970. Productivity of mule deer on the La Sal and Henry mountains of Utah. Utah State Div. Fish and Game Publ. No. 70-2, 133 pp. RicHE.Ns, V. B. 1967. Characteristics of mule deer herds and their range in northeastern Utah. J. Wildl. Manage 31(4): 651-666. NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be di- rected to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be sim- ilarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. They should be typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. Use a recent issue of either journal as a format, and the Council of Biology Edi- tors Style Manual, Third Edition (AIBS 1972) in preparing the manuscript. An abstract, about 3 percent as long as the text, but not exceeding 200 words, written in accordance with Biological Abstracts guidelines, should precede the introductory paragraph of each ar- ticle. Authors may recommend one or two reviewers for their article. All manuscripts re- ceive a critical peer review by specialists in the subject area of the manuscript under con- sideration. Manuscripts that are accepted and that are less than 100 printed pages in length will automatically be assigned to the Great Basin Naturalist. Those manuscripts larger than 100 printed pages in length will be considered for the Memoirs series. Illustrations and Tables. All illustrations and tables should be made with a view toward having them appear within the limits of the printed page. Illustrations that form part of an article should accompany the manuscript. Illustrations should be prepared for reduction by the printer to either a single-column (2 5/8 inches) or double-column (5 1/2 inches) width, with the length not exceeding 7 1/2 inches. Costs Borne by Contributor. Contributors to the Great Basin Naturalist should be pre- pared to donate from $10 to $30 per printed page toward publication of their article (in addition to reprint costs outlined in the schedule below). Authors publishing in the Great Basin Naturalist Memoirs may be expected to contribute $35 per printed page in addition to the cost of printed copies they purchase. No printed copies are furnished free of charge. A price list for reprints and an order form are sent with the galley proof to contributors. Reprint Schedule of the Great Basin Naturalist 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each 100 copies $20 $24 $28 $32 $36 $40 additional 200 copies 28 32 36 40 44 48 2 pp. 300 copies 36 40 44 48 52 56 $4 Great Basin Naturalist Memoirs No. 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodburry, H. H. Frost. $10. No. 2 Intermountain biogeography: A symposium. By K. T. Harper, J. L. Reveal, et al. $15. TABLE OF CONTENTS Utah flora: Brassicaceae (Cruciferae). Stanley L. Welsh and James L. Reveal 279 Typhlodromus mcgregori Chant (Acarina: Phytoseiidae) and its control of phyto- phagous mites in Utah and southern California apple orchards. B. A. Croft and C. D. Jorgensen 366 Host-induced morphological variations in the strigeoid trematode Post- hodiplostomum minimum (Trematoda:Diplostomatidae). III. Organs of attach- ment. James R. Palmieri 875 New synonymy and new species of American bark beetles (Coleoptera:Scolytidae), Part V. Stephen L. Wood 383 Biomass of fifty conifer forests and nutrient exports associated with their harvest. T. Weaver and F. Forcella 395 The algal flora of the campus of Brigham Young University, Provo, Utah. Cheng Mou-Sheng and Samuel R. Rushforth 402 Occurrence of ticks on mule deer in central Utah. Jordan C. Pederson 407 HE GREAT BASIN NATURALIST ISSN 0017-3614 )lume 37 No. 4 December 31, 1977 Brigham Young University MUS. COMP. ZOOU. LIBRARY AUG 1 5 1978 HARVARD UrsllVEERfiilTY GREAT BASIN NATURALIST MEMOIRS Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agri- cultural Sciences; Ernest L. Olson, Director, Brigham Young University Press, Univer- sity Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only previously unpublished manuscripts of less than 100 printed pages in length and pertaining to the biological natural history of western North America are ac- cepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manu- scripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist is $12 (outside the United States $13). The price for single numbers is $4 each. All back numbers are in print and are available for sale. All matters pertaining to the purchase of subscriptions and back numbers should be directed to Brigham Young University, Life Science Museum, Pro- vo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same of- fice at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts and other copy for either the Great Basin Naturalist or the Great Basin Naturalist Me7noirs should be addressed to the editor as instructed on the back cover. © 1978 Brigham Young University Press. All rights reserved Printed in United States of America 5-78 650 31730 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 VOLI'ME 37 December 31, 1977 No. 4 POPULATION DYNAMICS OF THE FERRUGINOUS HAWK DURING A PREY DECLINE Neil D. Woffinden' and Joseph R. Murphy- Abstract.— a nesting population of Ferruginous Hawks (Buteo regalis) at the eastern edge of the Great Basin in west-central Utah was studied for three nesting seasons, 1972-1974, during which time a significant decline in jackrabbit numbers occurred. The total number of hawks and nesting pairs varied throughout the study. In 1972, 16 pairs occupied territories and only 1 pair failed to nest. By 1974, however, only 5 pairs and 2 single birds oc- cupied territories. The number of young fledged ranged from 31 in 1972 to 3 in 1974. The nesting phenology of the Ferruginous Hawk and the reproductive period of black-tailed jackrabbits are clearly correlated. Of the jack- rabbit remains collected from hawk nests, 90 percent were from rabbits younger than 13 weeks. The decline in hawk numbers is thought to be directly correlated with a drop in the jackrabbit population. How to get something to eat and how to keep from being eaten are among the most hindamental requirements of life. These fac- tors have a tremendous impact on the struc- ture, abundance, and community integration of an organism (Huffaker 1970: 327). Much has been written about predation; yet many questions remain unanswered. Brown (1970: 247) suggested that seasons and breeding cycles of certain raptor spe- cies have never been properly correlated with behavior and abundance of their prey. In this study we sought to determine the ef- fect of prey density on the nesting success of the Ferruginous Hawk {Buteo regalis) and to investigate the timing of predator and prey reproductive cycles and establi.sh, if possible, a correlation between these two events. Finally, since little was known at the time of this study about the Ferruginous Hawk, we felt that any general knowledge of this species gained in the study would be of value. Funding was provided in part by the Na- tional Audubon Society and the Department of Zoology, Brigham Young University, Pro- vo, Utah. Gerald Richards and Reed Ferris aided in field work, and D. M. Allred and C. M. White gave valuable criticism of the manuscript. Methods and Procedures The study was conducted from June 1972 through October 1974 within an area of ap- proximately 932 km-, involving portions of Utah and Tooele counties in northcentral Utah. Most of the field work was carried out during the nesting season, but we made monthly ob.servations from October 1973 through October 1974. During the periods of intensive study, several visits were made to the study area each week (Fig. 1). 'Present address: Division of Natural Sciences, University of Pittsburgh at Johnstown, Johnstown, Pennsylvania, 15904. 'Department of Zoology, Brigham Young University, Provo, Utah, 84602. 411 412 GREAT BASIN NATURALIST Vol. 37, No. 4 Fig. 1. Study area location. Dec. 1977 WOFFINDEN, MURPHY: FERRUGINOUS HAWK 413 A smaller portion of the overall study area (238 km-) was selected for intensive study. This area coincides approximately with the boundaries of Cedar Valley, one of the eastern valleys of the Great Basin valley-and-range complex. The intensive study area, henceforth re- ferred to as Cedar Valley, was system- atically searched for Ferruginous Hawk nests in 1972 and 1973. Previous studies (Murphy et al. 1969, Smith and Murphy 1973) and banding records were also utiliz- ed in nest location and production for 1972 and previous years. An effort was made to locate inactive as well as active nests in both Cedar Valley and the western part of adjoining Rush Valley. Prey remains and castings were collected regularly from the active nests. Identi- fication of prey items was made by com- parison to known specimens. The length of rabbit feet found in and around the nests was recorded and compared with tables in Haskell and Reynolds (1947) to estimate ages and weights. Black-tailed jackrabbits (Lepus califor- nicus) density indices were obtained by use of line transects. Five randomly distributed, square transects .25 km on a side (Gross 1967) were conducted periodically through- out the latter part of the study. The tran- sect leg and direct distance from the ob- server to the point of flush was recorded for each rabbit flushed. The angle formed by the transect line and a line from the observ- er to the point where the animal was lo- cated was measured to the nearest degree. Numbers of road-killed jackrabbits ob- served during regular visits to the study area as well as during winter raptor surveys were recorded. By expressing the number of kilometers traveled per road-killed jackrab- bit, an index of abundance was obtained. During the latter part of the study, hind foot lengths were recorded for each road- killed jackrabbit. Clutch size, fledging rates, and other pro- duction figures were determined by regular nest visits throughout the nesting season. Statistical tests used include student "t" analysis of variance, and chi square tests. The level of significance is 0.05 unless stated differently. Results The black-tailed jackrabbit was abundant in the study area during the 1972 nesting season. It was not uncommon, during a visit to the study area, to see as many as 15 jackrabbits flush before the vehicle. Active nests during this season were cluttered with jackrabbit remains. One nest contained re- mains of at least 24 jackrabbits. Unfortu- nately, we were imable to obtain objective population densities. Gross et al. (1974), however, gave jackrabbit density indices for a nine-year period (1962-1970) for their northern Utah study area. The indices rep- resented approximately the number of jack- rabbits per km- (Gross et al. 1974: 27). They recorded in the autumn of 1970 a peak den- sity of 71.4 jackrabbits per km-. Stoddart (pers. comm. and in Howard 1975) reported a spring 1972 density of 47.1 jackrabbits per km- in the same area. This value dropped sharply to 9.7 km- in the spring of 1973. We assume that population densities in Cedar Valley were similar to those in northern Utah during the 1972 nesting sea- son. We also observed a decline in jackrabbit numbers between the 1972 and 1973 nest- ing seasons. One nest active in 1973 con- tained no rabbit remains. This nest was in the area of peak 1972 nesting density. An area of 27,225 m- near this nest was sys- tematically searched four times for jackrab- bits. One rabbit was flushed in each of the first three searches but none on the last search. The three flushes may have repre- sented one individual. In 1974 five randomly selected transects located throughout the study area were walked to establish jackrabbit densities (Table 1). These transects were similar to those used by Gross et al. (1974). Only one area had a density greater than one (1.0) rabbit/km- (Table 1). The number of kilo- meters per rabbit sighted varied from 0.46 in the area of highest densities to 8.0 in an area where only one jackrabbit was sighted. 414 GREAT BASIN NATURALIST Vol. 37, No. 4 In addition to the transects, during the first eight months of 1974 the number of road-killed jackrabbits observed was record- ed. Over 19,948 km were traveled during this period and 122 jackrabbits were count- ed (Table 2), for an average number of 89.7 kilometers traveled per jackrabbit. In spite of a lack of quantification, we feel that jackrabbit numbers dropped signif- icantly in our study area, as they did in northern Utah, between the 1972 and 1973 nesting seasons. Numbers continued to re- main low in 1974. This decline in prey numbers had a significant impact on the Ferruginous Hawk population in Cedar Val- ley. The total number of birds and number of nesting pairs varied throughout the study. Table 1. Number of transects, total rabbits ob- served, and mean density index of jackrabbits for Ce- dar Valley, 1974. No. Density Trans- No. Index sects Rabbits Km/ Rabbits/ Location Walked Counted Rabbit Km^ Cat Hill 6 7 1.37 0.17 Long Point 10 35 0.46 1.33 Roadtop Hill 5 1 8.0 0.06 Carrol 11 16 1.1 0.32 Quarry 3 0 - - Table 2. Number of road-killed jackrabbits, January- August 1974. No. No. Km No. Km Month Roadkills Traveled per Roadkill Jan 4 1310.5 327.6 Feb 0 1145.2 - Mar 5 1163.7 232.7 Apr 4 569.1 142.3 May 8 1664.4 208.1 Jun 33 2002.5 60.7 Jul 46 2355.4 51.2 Aug 22 822.7 37.4 Totals 122 11033.5 90.4 Table 3 summarizes these data for an eight- year period. In 1972, 16 pairs occupied ter- ritories in the intensive study area. There were no single birds present. Only one pair failed to make a nesting attempt. A total of nine pairs occupied nesting territories in 1973; of these, two pairs failed to nest. No single birds were observed. In 1974 five pairs of birds occupied territories; two pairs failed to nest and two single birds were ob- served occupying territories (Fig. 2). The total population varied from 32 birds present in 1972 to 18 in 1973 and 12 in 1974. Single individuals were observed only in 1974. A marked decline occurred over the three-year study, with the peak popu- lation occurring in 1973 (32 birds) and a low in 1974 (12 birds). The average number of birds present (for the study period) was 20.7, with a range of 12-32. The average number of Ferruginous Hawks occupying territories from 1967 to 1970 was 19 (range 18-27). The average number for the study period did not vary significantly from the average for the four-year period 1967-1970. In 1972 there were 14.85 km' per pair of birds in the intensive study area. This value increased to 26.24 km- and 34.03 km' for 1973 and 1974, respectively. It should be noted, however, that much of the habitat of Cedar Valley is unsuitable for nesting. The previous values are thus misleading because nests were placed on the hills surrounding the valley and no nests were found on the valley floor proper. The relative changes in nesting density during the study, however, would remain the same. In 1972, 94 percent of the resident pairs nested. This value then dropped to 77.8 percent and 42.9 percent for 1973 and 1974. The average for the three year period was 71.6 percent. Clutch Size.— Disturbance of Ferru- ginous Hawks during incubation is a signifi- cant cause of nest desertion (Powers et al. 1973, Olendorff 1973), and therefore few nests were visited during incubation. Thus, an accurate clutch size was not recorded for all nests. However, by extrapolating back from the number of young hatched, or in some cases fledged, a minimum clutch size Dec. 1977 WOFFINDEN, MURPHY: FERRUGINOUS HAWK 415 wa.s obtained. Minimum clutch sizes will be given, but it should be kept in mind that the numbers presented probably under- estimate the true clutch size. Minimum clutch size varied from a high of 2.7 in 1972 to 2.1 and 2.4 in 1973 and 1974. The average clutch size for these three years is 2.4. More complete data on clutch size was obtained by Weston (Mur- phy et al. 1969) and Smith and Murphy (1973). Mortality.— Nestling mortality was high during the study. Data are not available for the 1972 nesting season, but, based on the number of young fledged from each nest, mortality appeared to be negligible. An av- erage nestling mortality of 49.5 percent was, however, calculated for the 1973 and 1974 nesting seasons. This value is signifi- cantly greater than the 13.3 percent aver- age calculated from the 1967-1970 data (Murphy et al. 1969, Smith and Murphy 1973). In 1973 when the nests were visited regu- larly during incubation, two of the three nests in the southeastern end of the study area failed. Four chicks were lost in these two nests. One of the two chicks in the third nest disappeared during the first weeks after hatching. This may have been a case of cannibalism, or the bird may have died and was then eaten. The agent respon- sible for the deaths of the other young birds is not known. The striped skunk (Mephitis mephitis) was observed in the area as was the coyote (Canis latrans). A cliff near the nests was apparently used by Great Horned Owls as a roost. They or other raptors in the area could have taken the young. One nest was blown out of a tree during the 1973 nesting season. Thus, three of the six nests (50 percent) were completely de- stroyed, contributing to a nestling mortality of 42 percent. High nestling mortality occurred during the 1974 nesting season. Three nests hatched a total of seven young and fledged only three. Each nest fledged one young hawk. One young bird large enough to leave the nest, approximately five weeks old, was found dead beside the nest. Exam- ination suggested that it died of starvation. Earlier, approximately three weeks after hatching, one nestling had disappeared from Table 3. Ferniginous Hawk territory occupation and nesting attempts, 1967-1974. Percent Year No. Pairs No. Single Birds No. Territories Occupied No. Nesting Attempts Nesting per Occupied Territories Area per Territory Location' 1967' 13 13 13 100 18.34 CV 1968' 14 - 14 14 100 17.02 cv 1969^ 13 1 14 12 85.7 17.02 CV 197(F 9 2 11 7 63.6 21.65 cv 1971^ 5 0 5 5 100 - cv 1972 16 0 16 15 94 14.85 cv 1973 9 0 9 7 77.8 26.24 cv 1974 5 2 7 3 42.9 34.03 cv 1972^ 8 0 8 8 100 _ RV 1973^ 3 0 3 3 100 - RV 1974 1 1 2 1 50 _ RV 1974 5 0 5 5 100 - DV 'Data from Weston (1969) -Data from Smith and Murphy (1973) 'Data from J. R. Murphy (pers. comm.) Less time was spent in the field during the 1971 nesting season than the other years here listed. 'Data from R. Ferris (unpubl. field notes) "Abbreviations: CV = Cedar Valley, RV = Rush Valley. DV = Dugway Valley 416 GREAT BASIN NATURALIST Vol. 37, No. 4 the same nest, perhaps a victim of cannibal- ism. Two of three young were lost in anoth- er nest, the second 11 days after the first. The nestling mortality in 1974 was 57 per- cent. D }irds Present not nesting I iirds not }^i,t-!,:ii,ii, present Fig. 2. Ferruginous Hawk nesting territories and young fledged, 1971-1974. Numbers within the sections of each circle represent the number of young fledged for that year. Nest desertion.— None of the nests vis- ited during the study were deserted by the nesting pair. However, visits were not made prior to laying or during incubation periods in two of the three nesting seasons. In 1973 regular visits were made to three nests dur- ing incubation. Although these nests were not deserted, our presence probably contrib- uted to their failure (see previous section). Renesting.— There were no renesting at- tempts during the study. In all cases when nests failed the pair continued to occupy the nesting territory. A visit to the territory after nest loss would still elicit normal nest defense behavior, although the degree of in- tensity was often reduced. Fledging Success.— The number of young fledging declined drastically from a high of 31 in 1972 to 7 in 1973 and only 3 in 1974. Figure 3 summarizes these and oth- 32. 19«7tl96«19«9tl9TO 1971 1972(1973 1974 a a b b c I972h973 197411974 d d CV RV DV Fig. .3. Number of young fledged per year in Cedar, Rush, and Dugway valleys, 1967-1974. •Data from Weston (1968) •■Data from Smith (1971) ■"Data from J. R. Murphy (pers. comm.) ''Data from R. Ferris (unpubl. field notes) .Abbreviations are CV = Cedar Valley, RV = Rush Valley, and DV = Dugway Valley Dec. 1977 WOFFINDEN, MURPHY: FERRUGINOUS HAWK 417 er data for an eight-year period. The num- ber of young fledging per nesting attempt ranged from 1.9 in 1972 to 0.7 in 1973, and 1.0 in 1974 (Figs. 2 and 4). Figure 4 gives the number of nests fledg- ing one, two, three, and four young, respec- tively. In 1969 and 1972 the number of nests fledging three or four young was much greater than the number fledging one or two. In 1973 and 1974 the reverse was true; in fact, the maximum number of young fledged from any nest in these two years was two. Age of Jackrabbits Taken as Prey.— Measurements of jackrabbit hind feet taken from nests were compared with tables in Haskell and Reynolds (1947) to determine ages (Fig. 5). The age distribution was tri- modal. Thirty-one (44 percent) of the 71 rabbits were from 2 to 6 weeks old, 32 (45 percent) of the total were from 8 to 12 weeks of age, and 7 (10 percent) were 16 weeks or older. Of the total rabbits found in Ferruginous Hawk nests, 90 percent were 12 weeks old or younger. Figure 6 gives the age distribution of road-killed rabbits during June, July, and August of 1974. Fifty-seven percent of these rabbits were less than 13 weeks of age. Nesting Phenology and Jackrabbit Reproduction.- Gross et al. (1974) found four reproductive peaks in the black-tailed jackrabbit, the first commencing approx- imately mid-January and others occurring at intervals of 40 days (Fig. 7). Figure 7 gives the approximate birth dates of these four litters and the average size of each. Super- imposed on the graph are important dates in the nesting phenology of the Ferruginous Hawk. It appears that the hawk's nesting period is closely correlated with the repro- ductive period of the jackrabbit. Discussion Commencing with a peak in 1972, black- tailed jackrabbit populations declined throughout the study period. Gross et al. (1974) found that in their Curlew Valley study area, jackrabbit numbers reached a peak in 1970, then declined in 1971 and 1972. Numbers then dropped precipitously in the spring of 1973 (C. Stoddart pers. *- UJ Z IL. o Of UJ ^- z 1 1 2 19 3 68 4 1 2 19 3 69 4 1 2 19 3 72 4 1 2 19 3 73 4 1 2I3 1974 M NUMBER OF YOUNG FLEDGED Fig. 4. Number of Cedar Valley Ferruginous Hawk nests fledging 1, 2, 3, and 4 young, 1968-1974. 'Data from Weston (Murphy et al. 1969) 418 GREAT BASIN NATURALIST Vol. 37, No. 4 We assume that jackrabbit numbers in our study area were similar to those in Cur- lew Valley, 192 km to the north, prior to 1973, inasmuch as jackrabbits were numer- ous in 1972 at least on the east side of Ce- dar Valley. The hawk nests located that year were cluttered with numerous rabbit remains, and hawk production was at its highest point of the study. Smith (unpubl. ms.) reported a drop in jackrabbit numbers between 1969 and 1970. His observation corresponds closely with those made by Gross et al. (1974). We observed a decline in rabbit numbers between the 1972 and 1973 nesting seasons. This decline coincided with a fivefold de- crease in jackrabbit densities for the same period in Curlew Valley (C. Stoddart pers. comm.). An area of 27,225 m- in the eastern portion of Cedar Valley was systematically searched four times for jackrabbits during 13- 12- 11- 10 9- 8- 7- 6 5- 4- 3- 2. 1- the 1973 nesting season. Only one rabbit was seen. An active nest near this area had no evidence of rabbit remains. During the 1974 nesting season, transects were made in various parts of the study area (Table 1). Only one area had a density greater than one rabbit per km-. The number of kilome- ters per rabbit sighted varied from a high of 0.46 to a low of 8.0 in an area where only one rabbit was sighted during the study. Smith (unpubl. ms.) flushed 4.3 rabbits per km during 1970 (0.23 km /rabbit), the year of lowest jackrabbit densities during his four-year study. This value is twice as large as the comparable one for our study. In the first eight months of 1974, the number of road-killed jackrabbits was re- corded. Over 10,000 km were traveled dur- ing this period, and only 122 jackrabbits were seen. The average number of km trav- eled per jackrabbit was 89.7 (Table 2). oTT 10 11 12 13 I 14 I 15 16 AGE IN WEEKS Fig. 5. Number and ages in weeks of jackrabbit prey remains from Fermginous Hawk nests, 1972-1974. Dec. 1977 WOFFINDEN, MURPHY: FERRUGINOUS HAWK 419 Adams and Adams (1959) made similar sur- veys and found the frequency of road-kill per km far exceeded the value we obtained. Breeding success of the Ferruginous Hawks was higher on the eastern side of Cedar Valley. The western side of the val- ley, although it does contain some small areas of good rabbit habitat, does not have as much prime habitat as the eastern side. In times of high densities this marginal hab- itat was utilized by jackrabbits, which in turn allowed hawks to utilize an otherwise unsuitable area. We believe this helped produce the peak Ferruginous Hawk populations in the late 1960s and early 1970s. We believe that rab- bits began to decline first in areas of mar- ginal habitat, where they were most vulner- able to predation, and finally in areas of suitable habitat. Hence, a decrease in rabbit numbers occurred first in the western part of Cedar Valley and then in the eastern sec- tions, producing a similar pattern in the Ferruginous Hawk populations. ihhM'l' AGE IN WEEKS Fig. 6. Age distribution of road-killed jackrabbits, 1974. Dependence on Jackrabbits.— Weston (Murphy et al. 1969) found that jackrabbits made up a major portion of the Ferruginous Hawk diet. Kangaroo rats were found more frequently as prey remains, but their bio- mass was not as great as that of the jack- rabbits. Howard (1975) found that jackrab- bits made up 86 percent of the biomass of the three main prey species. Smith and Murphy (1973) indicated that jackrabbits made up from 93 percent to 95 percent of the prey biomass in their study. In Rush and Cedar Valleys, jackrabbits are used ex- tensively as a food source when young hawks are in the nest. One active nest in Rush Valley in 1972 contained the remains of at least 24 rabbits. In 1973 no active nests were found in Rush Valley. Alternate Prey Species.— Ground squir- rels were scarce in both valleys. Only three white-tailed antelope ground squirrels (Am- mospetmophilus leucurus) were seen in the two valleys over the entire study period. Townsend ground squirrels (Citellus toivns- endii) were more common, but scarce. Jack- rabbits made up the only plentiful mamma- lian prey. When a decline in rabbit numbers occurred, alternate prey was thus not available in sufficient numbers for the hawks to maintain their numbers, and the evidence indicates that the decline we have outlined is due to the drop in jackrabbit numbers. Townsend ground squirrels were more numerous in Dugway Valley, which is ap- proximately 85 km west of Cedar Valley. Prey populations in this valley were not quantitatively sampled but ground squirrels were evident in large numbers during each visit to the area, and Ferruginous Hawks were correspondingly more successful as nesters. There, only three (20 percent) of 15 nestling hawks failed to fledge. One nest failure accounted for two of these three deaths. The nesting success of the Dugway Valley hawks was evidently due to the pres- ence of a broad prey base. Therefore, our results, perhaps, apply only to areas where jackrabbits are the staple food of this hawk. Population Dynamics.— From 1968 to 1974, the numbers of pairs of Ferruginous 420 GREAT BASIN NATURALIST Vol. 37, No. 4 Hawks in the study area declined steadily (Table 3). In 1972, 16 pairs were present and only one pair failed to nest. In 1973, 9 pairs occupied territories in the intensive study area, and 2 pairs failed to nest. Final- ly, in 1974 the number of hawks in the study area dropped to 12 individuals— 5 pairs and 2 single birds. Two of the 5 pairs failed to nest. R. Ferris (pers. comm.) documented a similar decline in Ferruginous Hawk num- bers in Rush Valley (Table 3). Swartz et al. (1973) observed an increase in the number of nesting pairs of Rough-legged Hawks (Buteo lagopus) in Alaska, from 35 pairs in 1968 to 82 pairs in 1970; then the number of nesting pairs plummeted to 10, and only 2 young were fledged in 1972. Tubbs (1974) documented a similar decline in the Com- mon Buzzard (Buteo buteo) in Great Brit- ain. In these studies, as in ours, prey num- bers declined drastically prior to the decline of raptor populations. The data imply a di- rect correlation between predator and prey fluctuations. Habitat Vacancy.— In general, birds of prey occupy the same nesting territory for more than one year (Craighead and Craig- head 1969). Nesting territories can thus be conveniently checked each year in the early spring and throughout the year for the pres- ence of nesting birds. In 1972, 16 nesting territories were occupied. Only 9 territories were occupied in 1973, and 7 in 1974. The number of territories that were occupied declined each year, and previously active nesting territories were vacant (Table 3). Keith (1966) outlined the extent of vacant habitat of snowshoe hares (Lepus american- iis) during a decline. A similar phenomenon of previously occupied habitat being vacant occurred with the Ferruginous Hawk in our study area (Fig. 2). Davis et al. (1969) found that woodchucks CD CO < 5- 4. Fig. 7. Number and mean litter sizes of jackrabbits in Curlew Valley, with important dates in Fermginous Hawk nesting cycle. Vertical lines represent jackrabbit litters. The horizontal line gives important events in Fer- ruginous Hawk nesting cycle. 1 = territory selection, 2 = mean laying date, 3 = mean hatching date, 4 = mean fledging date, and 5 = mean migration date. 'Department of Zoology. Brigham Young University. Provo, Utah, 84602. 'Present address: Division of Natural Sciences, University of Pittsburgh at Johnstown, Johnstown, Pennsylvania, 15904. Dec. 1977 WOFFINDEN, MURPHY: FERRUGINOUS HAWK 421 (Marmota monax) that were removed from one section of their study area were promptly replaced by immature wood- chucks apparently from neighboring areas. Carrick's (1963) work with Australian Mag- pies (Gijmnorhina tibicen) suggested that a group of mimated birds was present in his study area, and any vacancies in the breed- ing group were filled by members of this unmated group. Vacant territories were not filled by extra individuals nor was a group of nonbreeding birds observed in the pres- ent study with the Ferruginous Hawk. Although 31 young were fledged in 1972, no young hawks were seen in the study area in 1973. Whether young hawks return to the place of their birth their first year is not known. Mortality rates for this period are also unknown. Craighead and Craighead (1969) suggested that as many as 88 percent of young hawks may die their first year. Hickey (1952) found annual mortality to be approximately 59 percent in yearling Marsh Hawks (Circus cyaneiis) and 30 percent for the adults. Based on production of the pre- vious year, even if mortality rates were as high as those listed here, a stable population size could be expected. We suggest that prey densities rather than mortality are re- sponsible for the 1973 and 1974 population declines. In 1972 all of the birds present on the east side of Cedar Valley nested and suc- cessfully fledged young, while at least three pairs on the west side did not. One pair on the west side of the valley did not nest. Two pairs that laid one and two eggs, re- spectively, did not hatch young. In 1973, three pairs were present on the west side of the valley. One pair fledged two young, one pair failed to nest, and the other pair's nest was blown from the tree. On the east side of the valley four pairs were present, plus a single bird. Though all four pairs built nests and laid eggs, over 50 percent of the eggs failed to hatch and only four young were fledged from these four nests. In 1974 four nest sites were occupied on the west side of the valley. One pair laid one egg, but none of the other birds laid eggs, although some nest building occurred. In the same year on the east side of the val- ley two pairs and one single bird occupied territories. Each of the pairs successfully fledged one of the three young that hatched in each nest. Clutch Size.— The average clutch size was 2.7 in 1972, 2.1 in 1973, and 2.4 in 1974. The overall average of 2.4 is lower than the 3.3 average reported by Weston (Murphy et al. 1969) and Smith and Mur- phy (1973). Piatt (1971) reported an average clutch size of 3.5 for 11 pairs of Ferru- ginous Hawks in Curlew Valley, and How- ard (1975) reported an average clutch size of 2.78 and 2.77 for 1972 and 1973 in the same area. His method of gathering clutch size data was similar to that of our study. Smith and Murphy (1973) found that clutch sizes vary from two to four eggs. Olendorff (1973) and Howard (1975) report- ed clutch sizes from one to five eggs. Clutch sizes of from one to four eggs were observed during this study. Clutches con- taining one and two eggs occurred frequent- ly in areas of low prey numbers. Howard (1975) found 11.6 percent of the clutches examined contained only one egg, but cor- relation was not made between these nests and prey densities; the data imply that one- and two-egg clutches were correlated with low prey densities. Fledging Rate.— The number of young fledged per year declined over the study period (Fig. 3). Thirty-one young success- fully fledged in 1972, an average of 1.9 per active pair. This value dropped to one young per active pair in 1974; however, there were only three active pairs (Table 1). The overall average fledging rate was 1.2. The number of nests fledging two or more young decreased over the study period (Fig. 4). The reduction of young fledged per nest- ing pair was evidently correlated with a re- duction in prey numbers. Nestling Mortality.— The low prey abundance was an important factor in nest- ling mortality. The female from one nest was observed 5.2 km away actively hunting at a time when normally she should have been at the nest brooding the young. Nests 422 GREAT BASIN NATURALIST Vol. 37, No. 4 were visited periodically after hatching oc- curred. On a few occasions two nests con- taining small young were visited, but no adult birds were observed. Howard (1975) also examined nestlings without observing the adult birds in the area. On one occasion we observed a Swainson's Hawk (Buteo swainsoni) coursing low over an active Fer- mginous nest, but no adult Ferruginous Hawks were observed. The absence of adults from the nest at this time was unusu- al, as almost always when a nest was ap- proached at least one bird and usually two would begin to display and defend the nest as soon as the investigator left his vehicle. This apparent nest inattentiveness would enhance the chance of chick mortality both from predation and exposure. We suggest that the high nestling mortality rate in 1973 (43 percent) was due in part to this behav- ior. The behavior resulted, we feel, from the additional pressure placed on the female hawk to provide herself and young with food, since the male was unable to provide adequate amounts. C. White (pers. comm.) found that Rough-legged Hawks on the Alaskan tundra were not attentive at the nest when prey density was low. On one occasion he saw a Golden Eagle eating young Rough-legged Hawks at the nest while the adults soared over the tundra some distance away. The nestling mortality in 1974 was 57 percent. During this nesting season, three nests hatched a total of seven young and fledged only three. Responses to Lower Prey Densities.— Pairs in areas of low prey densities either failed to nest or laid smaller clutches. Single birds, although they occupied nesting terri- tories, did not mate during the nesting peri- od. In those pairs that laid larger numbers of eggs, the female as well as the male was forced to spend periods away from the nest to supply food for the young. As a result, in many cases, the number of eggs hatched and young fledged was low. Otteni et al. (1972) found that Bam Owls (Tyto alba) raised 1.5 times as many young per pair in years when prey was abundant as in years when prey populations were more limited. Mclnvaille and Keith (1974) reported a numerical increase in a Great Horned Owl (Bubo virginianus) population in response to a cyclic increase of snowshoe hares. Shelf ord (1943) observed that the population of Snowy Owls (Nyctea scan- diaca) in the Arctic is directly dependent upon the abundance of lemmings (Lenimus spp. and Dicrostonyx spp.). During years when lemming populations were low, the owls failed to nest or laid only one or two eggs. In years of high prey populations the percentage of owls nesting was high, and each pair raised many young (maximum of 12). Pitelka et al. (1955) recorded similar correlations for Snowy Owls and Jaegers (Stercorarius spp.). Southern (1970) found that the percentage of Tawny Owls (Strix aliico) that nested during low prey years was near zero; however, in years of abun- dant prey, nearly 100 percent of the owls nested. Murphy (1975) found that the Golden Eagle population in central Utah expe- rienced a decline in numbers of nesting pairs during the period of this study. He suggested that there was strong evidence to indicate that this trend was also related to population fluctuations of the black-tailed jackrabbit. Howard (1975) also reported a decline in Ferruginous Hawk numbers and attributed it to a drop in jackrabbit numbers. We sug- gest that the Ferruginous Hawk responds to fluctuations in prey densities in a similar fashion to the other species discussed above. Nest Abandonment and Renestinc— Olendorff (1973) and Powers et al. (1973) suggested that Ferruginous Hawks are quick to abandon their nests if disturbed during nest building or incubation. In 1967, West- on (Murphy et al. 1969) reported that three nests containing eggs and four nests ready for eggs were abandoned early in the nest- ing season after only one visit to each nest. The next year, although identical nest ob- servation procedures were conducted, no nests were abandoned. We visited three active nests regularly during the 1973 nesting season. None of these three nests was abandoned, nor did Dec. 1977 WOFFINDEN, MURPHY: FERRUGINOUS HAWK 423 we observe any nest abandonment during the entire study. However, the young of two of these nests were destroyed by some- thing. We feel that our presence at the nest was probably responsible to a large degree for these deaths, because some carnivores are known to follow human tracks. Willis (1973) reported the destruction of Ant Bird (Gytnnopithijs bicolor) nests by predators af- ter a routine visit. Ray (1968) suggested the use of napthalene crystals around the nest and on paths to destroy the scent trails made by the investigator. Olendorff (1973) outlined methods of handling and times and duration of visits that will have the least in- jurious effect on the research subjects. An investigator would be wise to contemplate the impact of the study on the population prior to commencing his or her work. The methods outlined in the references above may be valuable research tools that could decrease the amount of investigator-caused losses to yoimg birds. Many raptors will lay a new clutch of eggs if all eggs are taken shortly after com- pletion of the clutch (Olendorff 1971). No renesting occurred during this study even though several nests were destroyed, includ- ing one nest that was blown out of a tree shortly after the time when the clutch should have been completed. The pair from one of the aforementioned nests that had successfully hatched three young, only to have them destroyed, moved 0.4 km east and defended an alternate nest site. Less than a dozen sticks were piled on top of a rock at this new location. Despite this, the pair actively defended this token nest. Age of Rabbits Taken as Prey.— The hind legs of 71 jackrabbits were collected from active nests throughout the study. The length of the hind foot was compared with tables in Haskell and Reynolds (1974) to de- termine age. A trimodal distribution was noted (Fig. 5). Thirty-one (44 percent) of the 71 rabbits were from two to six weeks old, 32 (45 percent) of the total were be- tween eight and twelve weeks of age, and 7 (10 percent) were sixteen weeks or older. Of the total rabbits found in Ferruginous Hawk nests, 90 percent were less than thirteen weeks of age. Similar results were reported by Dvmnet (1957) in Australia where avian predators were selectively killing young rabbits. He found that when the rabbits reached three months of age they were no longer vulnerable to the intense avian pre- dation to which they had been subjected earlier. The adult Ferruginous Hawks were clear- ly utilizing the immature jackrabbits to a much greater degree than the older rabbits. This is not surprising, because during the nesting season there are more immature than mature rabbits available to the hawks, and the young rabbits would imdoubtedly be easier to catch, kill, and carry. Nesting Phenology and Jackrabbit Reproduction.— Gross et al (1974) found four reproductive peaks in the black-tailed jackrabbit, the first commencing approx- imately mid-January and others occurring at intervals of 40 days (Fig. 7). The first two Utters are born prior to the selection of ter- ritories by the Ferruginous Hawks. In some cases the hawks do not even return to the nesting area prior to the birth of these two Utters. During years of good rabbit produc- tion, litters one and two should provide an abundant prey source for the courting hawks. After the nest has been constructed, egg laying begins (Fig. 7). During egg de- velopment an ample supply of prey may be as important as when young are in the nest. Average clutch sizes from years of good rabbit numbers are larger than those calcu- lated for years of low rabbit numbers (Smith and Murphy 1973). The average clutch size for 1972 was 2.7, but for 1973 and 1974 it was 2.1 and 2.4, respectively. In 1972 one nest on the west side of Cedar Valley contained a single egg, another only two eggs. In both cases hatch- ing did not occur. Also, a pair in this same area failed to nest. There is a general lack of good rabbit habitat on the west side of the study area. Data for years other than 1972 indicate fewer rabbits in this area than the eastern side of the valley. Although the data do not unquestionably support the thesis that fewer eggs are laid when prey is scarce during periods prior to egg devel- 424 GREAT BASIN NATURALIST Vol. 37, No. 4 opment, it appears that such a trend did oc- cur in this study as well as in the in- vestigation conducted by Shelford (1943) on the relationship of Snowy Owl response to low prey densities. We suggest that the timing of the arrival of the Ferruginous Hawk in its nesting ter- ritory is correlated with the presence of a suitable prey base in the form of coinciding litters of raljbits that are available for prey during prelaying activities. Rabbit litter three is born just prior to egg laying by the hawks. This is the largest of the three lit- ters, averaging 5.2 young per pregnant fe- male (Gross et al. 1974). During incubation in many raptor species the female hawk does most of the brooding and the male supplies her with food (Brown and Amadon 1968). If the male has little trouble sup- plying both himself and his mate with food during this period, the female apparently needs to leave the nest infrequently during the incubation period. A large number of young jackrabbits of approximately 1 to 12 weeks of age would be available to a Ferru- ginous Hawk pair that occupied a suitable territory throughout the period of in- cubation during a year of good rabbit den- sities. This presumptive age distribution compares favorably with the distribution of the ages of rabbits actually taken from the nest (Fig. 5). Members of the first litter would be approximately 12 weeks of age at the time of laying and would still be avail- able to the hawks during incubation. Shortly after hatching, rabbit litter four is bom. Litter three would be approximately four weeks of age, and litter two approx- imately twelve weeks of age at this time. The three largest litters are thus available for food as the young hawks develop. The members of litter four are less than five weeks of age while the young hawks are in the nest. Based on ages of rabbits taken to the nest, three of the four litters (litters 2, 3, and 4) are available for use as food for the nestling hawks. The last two litters probably provide the majority of the rabbits utilized as prey during hawk development. After the young hawks leave the nest they remain in the area for approximately one month prior to migration. Rabbit litter four is approximately 10 weeks old at this time, and probably provides the major por- tion of the food supply of the young hawks. Young hawks likely cannot catch adult rab- bits or even young of that year other than members of litter four. The timing of de- parture could well be correlated with a de- cline in available prey. Data from rabbit transects made during the nesting season in 1974 indicate that this may be so, inasmuch as jackrabbit densities declined throughout the nesting season. Other raptors migrate at approximately the same time as the Ferruginous Hawk (August 1). A. Jenkins (pers. comm.) noted a peak in migrating hawks along the Wasatch Front in late August. Although some buteos were observed, most migrants were accipi- ters and falcons. Olendorff (1973) recorded the Ferruginous Hawk as a resident species in the Pawnee Grassland of Colorado. It is not known why this species should be mi- gratory in central Utah, yet resident in a lo- cation only about 800 km to the east. In both locations an influx of wintering raptors occurs, primarily Rough-legged Hawks and Bald Eagles (Haliaeetus leiicocephalus). In Utah the Ferruginous Hawks have migrated before these species appear. Competition does not seem, then, to be the factor of im- portance. We suggest that perhaps prey availability dictates time of migration. Fur- ther light could be shed on this hypothesis if more were known about the wintering areas and food habits of the Ferruginous Hawk. Literature Cited Adams, H. B., and L. Adams. 1959. Black-tailed jack- rabbit carcasses on the highways in Nevada, Idaho, and California. Ecology 40(4): 718-720. Brown, L., and D. Amadon. 1968. Eagles, hawks, and falcons of the world. McGraw-Hill Company Inc., New York. Brown, L. 1970. African birds of prey. Collins, Lon- don. Carrick, R. 1963. Ecological significance of territory in the Australian Magpie, Gymnorhina tibicen. Proc. XIII Intern. Omithol. Congr. 740-753. Craighead, J. J., and F. C. Craighead, Jr. 1969. Hawks, owls, and wildlife. Dover Publ, Inc., New York. Dec. 1977 WOFFINDEN, MURPHY: FERRUGINOUS HAWK 425 Davis, D. E., J. J. Christian, and F. Bronson. 1964. Effects of exploitation on birth, mortality, and movement rates in a woodchuck population. J. Wildl. Manage. 28(1): 1-9. DuNNET, G. M. 1957. Notes on avian predation on young rabbits, Oryctolagus cuniculus (L.). CSIRO Wildl. Res. 2(1): 66-68. Gross, J. E. 1967. Demographic analysis of a northern Utah black-tailed jackrabbit population. Unpub- lished doctoral dissertation. Department of Wild- life Sciences, Utah State University, Logan, Utah. Gross, J. E., L. C. Stoddart, and F. H. Wagner. 1974. Demographic analysis of a northern Utah jackrabbit population. Wildl. Mongr. No. 40: 1-68. Haskell, H. S., and H. G. Reynolds. 1947. Growth, developmental food requirements, and breeding activity of the California jackrabbit. J. Mammal. 28(2): 129-136. HiCKEY, J. J. 1952. Survival studies of banded birds. Spec. Sci. Rept. Wildl. No. 15. 117 p. Howard, R. P. 1975. Breeding ecology of the Ferru- ginous Hawk in northern Utah and southern Idaho. Unpublished master's thesis. Department of Wildlife Sciences, Utah State University, Lo- gan, Utah. HuFFAKER, C. B. 1970. The phenomenon of predation and its roles in nature. In: Dynamics of popu- lations, P. J. den Boer and G. R. Gradwell (eds.). Proc. Adv. Study Inst, on Dynamics of Numbers in Pop., Oosterbeek, Netherlands. Keith, L. B. 1966. Habitat vacancy during a snowshoe hare decline. J. Wildl. Manage. 39(4): 828-832. McInvaille, W. B., and L. B. Keith. 1974. Predator- prey relations and breeding biology of the Great Horned Owl and Red-tailed Hawk in central Al- berta. Canadian Field-Nat. 88(1): 1-20. Murphy, J. R. 1975. Status of a Golden Eagle popu- lation in central Utah, 1967-1973. In: Status of raptors. J. R. Murphy, C. M. White, and B. E. Harrel (eds.). Raptor Research Report No. 3, pp. 91-96. Raptor Research Foundation, Vermillion, South Dakota. Murphy, J. R., F. J. Camenzind, D. G. Smith, and J. B. Weston. 1969. Nesting ecology of raptorial birds in central Utah. BYU Sci. Bull., Biol. Ser. 10(4): 1-36. Olendorff, R. R. 1971. Falconiform reproduction: a review. Part 1. The pre-nestling period. Raptor Research Report No. 1. Raptor Research Founda- tion, Vermillion, South Dakota. 1973. The ecology of the nesting birds of prey of northeastern Colorado, USIBP Brassland Biome. Technical report No. 211. Oteni, L. C, E. G. Bolen, and C. Cottam. 1972. Predator-prey relationships and reproduction of the Bam Owl in southern Texas. Wilson Bulletin 84(4): 434-448. Platt, J. B. 1971. A survey of nesting hawks, eagles, falcons and owls in Curlew Valley, Utah. Great Basin Nat. 31(2): 51-65. Pitelka, F. a., P. Q. Tomich, and G. W. Treichel. 1955. Ecological relations of jaegers and owls as lemming predators near Barrow, Alaska. Ecol. Mongr. 25:85-117. Powers, L. R., R. Howard, and C. Trost. 1973. Pop- ulation status of the Ferruginous Hawk in south- eastern Idaho and northern Utah. In: Status of raptors, J. R. Murphy, C. M. White, and B. E. Harrel (eds.). Raptor Research Report No. 3, pp. 153-157. Raptor Research Foundation, Vermil- lion, South Dakota. Ray, T. D. 1968. Napthalene crystals as protection for nesting raptors. Hawk Chalk 7(1): 52-54, April. Shelford, V. E. 1943. The abundance of lemming in the Churchill area, 1929-1940. Ecol. 24: 472-484. Smith, D. G., and J. R. Murphy. 1973. Breeding ecol- ogy of raptors in the eastern Great Basin of Utah. BYU Sci. Bull., Biol. Ser. 18(3): 1-76. Smith, D. G. Breeding response of raptors to prey population fluctuations in the eastern Great Basin desert of Utah. Unpublished manuscript. Southern, H. N. 1970. The natural control of a popu- lation of Tawny Owls (Strix altico). J. Zool., Lon- don 162: 197-285. SwARTZ, L. G., W. Walker II, D. G. Roseneau, and A. M. Springer. 1973. Populations of Gyrfalcons on the Seward Peninsula, Alaska, 1968-1972. In: Status of raptors. J. R. Murphy, C. M. White, and B. E. Harrel (eds.). Raptor Research Report No. 3, pp. 71-75. Raptor Research Foundation, Vermillion, South Dakota. TuBHS, C. R. 1974. The buzzard. David and Charles Inc., North Pomfret, Vermont. Willis, E. O. 1973. Survival rates for visited and un- visited nests of Bicolor Antbirds. Auk 90(2): 263-267. A NEW SPECIES OF RHOPALOMYIA (DIPTERA: CECIDOMYIIDAE) FROM ARTEMISIA LUDOVICIANA NUTTALL (COMPOSITAE) IN UTAH Raymond J. Gagne' .\bstract.— Rhopalomyia (Diarthronomyia) subhumilis Gagne, a new species of Artemisia ludoviciana Nuttall (Compositae), is described and its taxonomically important structural features illustrated. The new species is com- pared to its most similar North American congeners. This paper provides a name for a species of Rhopalomyia whose biology is treated in a companion paper by Ranasinghe (1977). As pointed out in Jones, Gagne, and Barr (in press), Rhopalomyia species are so nu- merous and the structural differences so few and so slight that the species do not lend themselves well to description without bio- logical data. That condition is met for this new species by the information in Rana- singhe (1977). Rhopalomyia subhumilis n. sp. Male.— Wing length, 2.2-2.5 mm. Eyes separated at vertex by less than diameter of 1 eye facet. Antenna with 14-16 flag- ellomeres; neck flagellomere III, 0.50-0.86 times length node; neck V, 0.60-0.88 times length node; neck VII, 0.64-0.90 times length node. Fused labellae (Fig. 6) with ventral notch in frontal view, 0-4 setae per side. Palpus (Fig. 6) 2-segmented: segment I as wide as long, with 1-5 setae; segment II as long as I but tapered apically, with 1-3 setae. Frontoclypeus with 6-9 setae per side. Anepisternum with 4-8 setae. Claws toothed (Fig. 7). Empodia as long as claws. Abdo- men: tergum III with 8-22 setae per side, tergum V with 13-26, VII with 13-20 and occasional scales, and VIII with 8-20, the two groups on VII united mesally or sepa- rated by short distance; pleura with scales, 0 setae; sterna II-VII each with basal pair of trichoid sensilla, sternum VII with basal setal group continuous across sclerite. Ter- minalia (Figs. 3-5): basimere stout with 'Research Entomologist, Systematic Entomology Laboratorv. IIBIII, Agricultu NHB 168, Washington, D.C. 20560. short ventral lobe; telomere short-ovoid, the tooth wide; sternum X concave apically. Female.— Wing length, 1.8-1.9 mm. An- tenna with 15-16 flagellomeres; flagellomere necks not defined or very short to VI, unde- fined beyond VI. Other head and thoracic characters as in male. Abdomen: tergum III with 12-14 setae per side; V with 13-18, and VII with 5-12, the two groups on VII barely divided; pleura and sterna as in male; tergum VII 0.23-0.26 times length dis- tal half of ovipositor. Pupa.— Head (Figs. 1-2): antennal horns strongly developed, acutely pointed; frons with two prominent, pointed protuberances. Pronotum convex, not prominent, the two setae long. HoLOTYPE, male, ex subterranean stem galls on Artemisia ludoviciana Nutt., Phila- delphia Flat, 12 mi. E Ephraim, Sanpete Co., Utah, 9-VII-1975, R. B. Famsworth, USNM Type No. 66621. Paratypes: 14 males, 23 females, same data as for holo- type; 2 males, 6 females, same data as holo- type, except collected lO-VII-1976, S. K. Ranasinghe. Other material: pupal exuvia and galls associated with above collections. Representative collections of males, females, and pupal exuvia in Life Science Museum, Brigham Young University and California Academy of Sciences. Remainder of mate- rial in USNM. Remarks.— Rhopalomyia subhumilis be- longs in the subgenus Diarthronomyia as do all other Rhopalomyia that occur on Arte- misia and Chrysanthe7nu7n. The dis- iral Research Service, USDA. Mail address: c/o U.S. National Museum, 426 Dec. 1977 gagne: a new gall midge 427 Figure Legends Figs. 1-7, taxonomic characters of Rhopalomyia subhumilis Gagne: 1-2, pupal head; 3, male terminalia (dorsal); 4, same (ventral); 5, same (mesal); 6, labella and palpus; 7, claw and empodium. 428 GREAT BASIN NATURALIST Vol. 37, No. 4 tingiiishing structural feature uniting these species is the lack of all but caudal setae on the abdominal terga. The new species differs from all other Nearctic species of Diarthronomyia by the following com- bination of characters: palpus two-seg- mented, claws toothed, male basimere with ventral lobe, male telomere short, with wide apical tooth, pupal head with frontal protuberances and strongly developed an- tennal horns. Rhopalomyia siibhumilis will not run past couplet 5 in the key to 23 spe- cies of Rhopalomyia on Artemisia tridentata (Nuttall) in Jones, Gagne, and Barr (in press). The pupal head will readily dis- tinguish this species from the other keyed Rhopalomyia having 2-segmented palpi and toothed claws. Of the 9 other Nearctic Rhopalomyia (Diarthronomyia), only R. gnaphalodis Felt from a woolly leaf gall of "Artejiiisia gnaphaloides" and jR. chrysan- themi from a conical gall on green tissues of Chrysanthemum spp. have two-segmented palpi and toothed claws, but the pupal an- tennal horns are much more attenuate in both species, and the male telomere of R. chrysanthemum is much longer than in R. siibhumilis. The name "subhumilis" means "subterranean". Literature Cited Gagne, R. J. 1975. A redefinition of Diarthronomyia Felt as a subgenus of Rhopalomyia Riihsaamen (Diptera: Cecidomyiidae: Oligotrophidi). Ann. En- tomol. Soc. Amer. 68: 482-484. Jones, R. G., R. J. Gagne, and W. F. Barb. (In press.) A systematic and biological study of the gall midges (Diptera: Cecidomyiidae) of Artemisia tri- dentata Nuttall (Compositae) in Idaho. Univ. of Idaho, College of Agriculture Misc. Series. Ranasinghe, M. a. S. K. 1977. The biology of a spe- cies of Rhopalomyia on Artemisia ludoviciana Nutt. Great Basin Naturalist .37: 000. BIONOMICS OF A SUBTERRANEAN GALL MIDGE (DIPTERA: CECIDOMYIIDAE) FROM ARTEMISIA LUDOVICIANA^ M. A. S. K. Ranasinghe' Abstract.— Bionomics of a gall midge that emerged from nodulelike structures of herbaceous sage, Artemisia bidoviciana, was studied as a part of a larger investigation on possible nitrogen fixation by this plant. Infested plants collected from the field were regularly examined in the laboratory where some of them were grown in a liquid nutrient medium. In the laboratory, adult midges were reared from pupae and induction of infestation was attempted. Apparent nodulation of these plants is caused by the subterranean bud galls of a previously unknown gall midge, Rhopalomijia subhumilis Gagne. Life history of this midge is reported. These midges have one gener- ation per year in the study areas and overwinter as larvae. There were no indications of paedogenesis. These midges are parasitized by a species of Platygasteridae. Apparent nodulation on the underground parts of an herbaceous sagebrush, Artemisia ludoviciana Nutt., was reported by Farns- worth and Hammond (1968) in their in- vestigations on nitrogen fixation by non- leguminous plants. Later, nitrogenase activity, measured through acetylene- ethylene gas assay, was recorded from these nodulelike structures (Farnsworth and Claw- son 1972). When midges emerged from the stored mature bodies in the laboratory, it was hypothesized that they might act as vectors of the nitrogen-fixing micro- organisms (Farnsworth 1975). It was also possible that the presumptive nodules ac- tually were subterranean galls of a hitherto unknown midge that infested these plants. This study was imdertaken to investigate the bionomics of these midges in relation to the nodulelike structures of these sage plants, as a part of a larger investigation of possible nitrogen fixation by Artemisia lu- doviciana. The midge that emerged from the no- dulelike structures was found to be Rhopa- knnyia subhumilis Gagne, a species pre- viously imknown to science, in the family Cecidomyiidae (Gagne 1977). This large family includes most of the economically important gall midges (Mani 1964). Rhopa- lomijia subhumilis is the only subterranean gall midge reported from Artemisia. It is one of the few gall midges known to in- habit the subterranean parts of the host plant. Arte7nisia ludoviciana is an herbaceous plant with a perennial, subterranean, rhi- zomal stem. In Utah, the annual aerial growth of this plant persists from early spring to early fall. This plant is indigenous to much of the United States from western Canada and California eastward to Montana and the Dakotas. The objective of the present research is to study the life history of this gall midge in relation to the galls of the host plant. Rubsaamen (1892) erected the cosmopoli- tan genus Rhopalomijia which now contains over 200 known species (Gagne 1974), 56 of which are from North America north of Mexico (Foote 1965). Jones (1971) apparent- ly found an additional 35 species from the Artemisia tridentata complex in Idaho, and in relation to this work Gagne (1974) re- vised the subgenus Diathronomyia. Gagne (1975) recognized additional new Rhopa- ■Results of a research project for M.S. degree. 'Department of Zoology, Brigham Young University, Provo, Utafi, Florida, Gainesville, Florida, 32611. 84602. Current address: Department of Entomology and Nematology, University of 429 430 GREAT BASIN NATURALIST Vol. 37, No. 4 lomyia species. He has named and reviewed the taxonomic status of the gall midge re- ported in this paper (Gagne 1977). Rhopalomyia species are apparently host specific and they are responsible for a par- ticular kind of gall on some part or parts of the plant (Gagne 1974). Mani (1964) report- ed that galls of Rhopalomyia are largely confined to the Compositae, but a few make galls in plants belonging to other fam- ilies (Felt 1940). Several Artemisia species harbor galls of Rhopalomyia (Felt 1940), al- though none has been reported from Arte- misia htdoviciana. Most Rhopalomyia spe- cies show marked preference to flower and bud galls (Felt 1915). Rhopalomyia thomp- soni Felt was named from the root galls of Solidago nigosa (Felt 1907), and Rhopa- lomyia hirtipes O.S. from both aerial and subterranean bud galls of Solidago juncea (Felt 1915). Published records of sub- terranean galls on Artemisia were not found, although galls are known to occur on other parts of these plants (Felt 1911b, 1940). Earlier work on Rhopalomyia was mostly confined to records of rearing them from galls and to taxonomic descriptions (Felt 1907, 1911a, 1916). Cockerell (1909) de- scribed the biology of Rhopalomyia heth- eliana Ckll. from Artemisia frigida, and Lander (1951) reported a short description of an unidentified Rhopalomyia species from Artemisia tridentata Nutt., in Utah. The ecology of several apparent species of Rhopalomyia found on Artemisia tridentata complex in Idaho was studied by Jones (1971), and the biology of Rliopalomyia hir- tipes from Solidago juncea was reported in detail by Spence (1969). Materials and Methods The study areas are located in the Manti- La Sal National Forest within an elevation range of 2,440 to 3,200 m and between 22 to 30 km southeast of Ephraim, Sanpete Coimty, Utah, where an abundance of host material was available. This study was car- ried out from May 1976 to June 1977. In the laboratory, for each gall the loca- tion on the host plant was recorded and the diameter was measured using a vernier cali- per. Some of the galls were dissected under a stereoscopic microscope to record the number and stage of development of the midges found inside. An ocular micrometer was used for all linear measurements of the gall midges. The soil-free, infested plants, supported by cotton wads kept on either side of the gallbearing section, were grown in glass tubes (Fig. 1) containing White's liquid nu- trient medium (Paul 1970). Each tube was covered with black paper to prevent Hght penetration. Adult midges were obtained from the ma- ture galls by keeping them in plastic bags. Once emerged, the adult males and females were separated, their dimensions were re- corded, and the midges were preserved in 70 percent alcohol. The adult parasites col- lected were also preserved in 70 percent al- cohol. To find whether both the male and fe- male midges emerged from the same gall, 24 mature galls were kept, each in a sepa- rate plastic vial (8.0 x 2.5 cm), between cot- ton wads at the top and the bottom of the vial. Some infested plants were kept intact in rearing chambers (36 x 26 x 26 cm), modified from Spence (1969), to allow ob- servance of the adult behavior (Fig. 2). Some other infested plants were grown in a growth chamber at 35° C and 12 hours of daylight using White's liquid nutrient me- dium. Induction of the galls in the labora- tory was attempted by keeping 24 unin- fested plants in one half of a Plexiglas cage with a nylon-mesh-covered window, separated by a removable Plexiglas parti- tion from the other half of the cage con- taining 24 infested plants (Fig. 3). These plants were grown in plastic trays contain- ing cotton pads moistened with the liquid nutrient medium periodically. The partition was removed once the adult midges emerged and the uninfested plants were ex- posed to them. Later, these plants were ex- amined for the signs of gall formation. Dec. 1977 ranasinghe: gall midge bionomics 431 The photographs were taken using a Honeywell Pentax ® SP 1,000 camera, Vivi- tar extension tubes, and a Nikon ® stereo- scopic microscope with the ase of Pan- atomic (g) X or Plux X (g) films. Results Life History Egg.— The eggs the laboratory on were first observed in 17 June 1976 but were collected from the field from 24 June to 17 Figs. 1-3: 1, Glass rearing tube with an Artemisia ludoviciana plant, CW-Cotton wad, NM-Nutrient medium. 2, Single-compartment midge-rearing cage, CD-Cage door. 3, Twin-compartment midge-rearing cage, CD-Cage door. 432 GREAT BASIN NATURALIST Vol. 37, No. 4 July 1976. These eggs were normally found in or near the leaf axils of the lower part of the aerial shoots and rarely on the under- side of the leaves. In the laboratory the eggs were found even on the walls of the glass rearing tubes. Eggs were usually found in clusters varying from 12 to 49 eggs per cluster. Nine egg clusters included a total of 205 or an average of 23 eggs per cluster. The eggs were pale red in color. The av- erage measurements of 45 randomly se- lected eggs were 0.30 x 0.06 mm. Each egg was cylindrical in shape with tapering ends (Fig. 4). Under laboratory conditions the eggs hatched in three days at room temper- ature to produce pale red-colored, first in- star larvae. In the field the exact time of hatching could not be determined, but on one occasion eggs observed in the field on 7 July at Skyline Drive could not be found during the next visit a week later, in- dicating that the eggs most likely hatched in less than a week. Larva.— Earliest laboratory observation of the pale red-colored, first instar larvae was on 19 June 1976. Average measure- ments of 23 such larvae were 0.27 x 0.06 mm. When examined under the microscope, these larvae were found to crawl on the stem. However, the first instar larvae soon perished under laboratory conditions. In the field similar larvae were recovered from around the basal axillary buds of the shoots. On one occasion discarded pale red-colored larval skins were observed near a newly formed gall. The first larva to be recovered from a new young gall was found on 8 July 1976 at the Blue Bell area at an elevation of 2,700 m when the soil temperature at a depth of 7 cm was 18° C. These larvae were colorless and translucent. Each of these larvae was confined to a distally ta- pering larval chamber which was slightly larger than the larva. The average of 33 such larvae measured was 0.35 x 0.23 mm. The midge overwintered in this larval form. In the following spring few larvae similar to above were still found in the galls on 9 May 1977 at the Blue Bell area, soon after snow had melted. At Skyline Drive (eleva- tion 3,300 m) these young larvae occurred even in the first week of June. At the Blue Bell area these young larvae were inside thin-walled, liquid-containing larval cham- bers of the galls located at the bases of ae- rial shoots about 1.7 cm high, growing in soil where the temperature at a depth of 2.5 cm was 8° C. However, by this time some larvae had already begun to develop (Fig. 5). With the growth of the larva, the size and the wall thickness of the larval cham- ber increased progressively. But the distal end of the larval chamber remained com- paratively thin walled and was oriented away from the basal attachment of the gall to the host plant. The fluid content of the larval chamber gradually decreased with the growth of the larva. No solid materials were found in the larval chambers at any time. In the laboratory the young, growing lar- vae kept between the moist filter papers survived only up to three days, probably due to lack of nutrients. Hence the devel- opment of a given larva could not be fol- lowed through all the different stages by rearing them in the laboratory. Instead, midge galls regularly collected from the field during the growing season were dis- sected to follow the development of the lar- vae. The range and the average of the lengths of the larvae collected from the field during their growing season are shown in Table 1. Table 1. Larval lengths of Rhopalomijia subhumilis from Artemisia ludoviciana galls at Philadelphia Flat (1976). Date 1 Number of Length in mm larvae Average Range 03-VI-76 26 0.54 0.42-1.32 08-VI-76 47 0.78 0.42-1.38 17-VI-76 26 1.30 0.62-2.12 24-VI-76 26-VI-76 17 1.51 0.81-2.12 Ol-VII-76 07 1.47 1.00-1.70 09-VII-76 15 1.56 '0.40-2.10 'New young larvae Dec. 1977 ranasinghe: gall midge bionomics 433 During this period the length-to-width ra- tio of the larvae remained about 2:1. The largest larva observed was 2.12 x 1.00 mm. With growth, the originally colorless larva in the young gall changed to white, then yellow, and finally to brownish vellow at maturity. This mature larva (Fig. 6) was opaque compared to the translucent young larva. These midge larvae were rather char- acterless. The larval body consisted of 13 segments. The head capsule of the young larva was inconspicuous but became more 0-5MM Figs. 4-9. Developmental stages of Rhopabmyia subhumilis: 4, Eggs. 5, Young larva. 6, Lateral view of the ma- ture larva. 7, Lateral view of early pupa. 8, Ventrolateral view of late pupa. 9, Lateral view of female pupa just prior to eclosion. 434 GREAT BASIN NATURALIST Vol. 37, No. 4 visible with growth, due to pigmentation. The growing larvae were orientated with their heads toward the bases of the larval chambers until they reached the prepupal stage when their position reversed. The lar- vae were usually sluggish except for their feeding movements. When kept on a drop of water and subjected to intense light from a microscope lamp they moved away from the light by waves of body contractions. Pupa.— The appearance of a pair each of antennal and facial horns on the ventral side at the anterior end of the mature larva was the first externally visible sign of pupa- tion. At this time some of the other pupal structures were also visible through the cu- ticle of the prepupa. The horns were first visible on the laboratory-reared larvae on 9 June 1976, and the pupae were collected at Philadelphia Flat (elevation 2,900 m) on 15 June 1976. This early pupa was white in color with a slight dorsal curvature of the body at the anterior end (Fig. 7). By the end of the first 24 hours the distal spines of the antennal and facial horns became brown in color, due to pigmentation. The compound eyes, which were white in color, became progres- sively browner within the first 48 hours, be- ginning around the ommatidia. On the fourth day a black bar appeared connecting the two eyes on the dorsal side. The eyes became black in color by the end of the fifth day. The pigmentation of the antennae commenced at their distal ends by the third day and then gradually proceeded toward the bases, making the orange-colored an- tennae entirely black by the end of the fifth day. The white-colored thorax turned yellow within the first 24 hours and brownish red by the end of the third day. It became brown in color by the end of the fifth day, and by then two black bands appeared dor- solaterally at the posterior end of the tho- rax. The black coloration of the wing buds and the legs was initiated at their distal ends by the end of the third day and then gradually proceeded proximally to make them entirely black by the end of the fifth dav. The color of the pupal abdomen changed from white to pale yellow within the first 24 hours. In the male pupae this color in- tensified and then turned yellowish brown in the mature condition. But in the female, pupae the color of the abdomen gradually changed to pale red by the third day and then increased in intensity to become dark red upon maturation. The pupae were com- pletely mature by the end of the sixth day (Fig. 8). The setal bands of the adult were visible through the pupal case just before eclosion, and the ovipositor in the female pupa was apparent by this time (Fig. 9). The average length of 10 randomly se- lected mature male pupae was 1.92 mm, and that of 10 female pupae selected at random was 2.11 mm. In addition to this size difference the female pupae could be distinguished from the male pupae by the red-colored abdomen and comparatively shorter legs which extended only up to the end of the fifth abdominal segment. In con- trast, the male pupae had a yellowish- brown abdomen and longer legs, which ex- tended up to the end of the seventh abdom- inal segment. I was able to rear the pupae from the first day of pupation to the adult stage by keeping them between moist filter papers in petri dishes at room temperature. The first adult emerged in the laboratory on 16 June 1976. Duration of the pupal stage varied from six to seven days in the laboratory at room temperature. In the field, pupal exuvia were first observed on 24 June 1976 at the Blue Bell area when soil temperature was 18° C at a depth of 6.0 cm. Adults.— Immediately prior to eclosion the mature pupa became periodically active with bending movements of the body. The first signal of eclosion was the appearance of a longitudinal split along the mid-dorsal line of the pupal thorax. This thoracic split widened, exposing the adult thorax, and the adult head protruded through it with the eyes followed by the bases of the antennae. The white-colored adult thorax, bearing four dorsal bands of hair, emerged with the prothoracic legs coming out next and the halteres becoming free before the wings Dec. 1977 ranasinghe: gall midge bionomics 435 (Fig. 10). Once the head and thorax were free, the adult apparently paused for a few seconds and then vigorously wriggled to free the abdomen from the pupal case. Dur- ing eclosion the shaking of the free parts of the body perhaps helped to draw the en- cased parts out of the pupal case. In the laboratory eclosion was completed in 11 minutes for this pupa which was reared in a petri dish. The examination of the pupal exuviae indicated a similar process of eclosion under the field conditions. The pupal exuviae remained protruding out of the cavities of the gall surface (Fig. 11). The adults emerged in the laboratory on 16 June 1976. In the field adult emergence occurred from 24 June 1976 (lower eleva- tions) to 17 July 1976 (higher elevations). Adults were collected imtil 5 August 1976 from the infested plants reared in a growth chamber. When these plants were allowed to continue inside the growth chamber, five more .adults emerged from the galls on the new shoots; they presumably belonged to a second generation. But no second gener- ation adults were collected from the field. Laboratory observations on 106 adults gave a male to female sex ratio of 1:1.9. Twenty-five randomly selected males aver- aged 1.76 mm in length, within a range of 1.28 to 2.12 mm. For 25 randomly selected females the average length was 1.96 mm within a range of 1.54 to 2.30 mm. These measurements were taken from the anterior end of the head capsule to the end of the last abdominal segment, exclusive of the an- tennae and the terminalia. The adult male has a yellowish-brown abdomen, com- paratively long legs and a short pair of claspers (Fig. 12). The female could be rec- ognized by her shorter legs, broad, red-col- ored abdomen, and the long ovipositor (Fig. 13). A complete description of the adults is presented by Gagne (1977). Gravid females isolated from the males did not oviposit even after three days, but when adults of both sexes were present to- gether they laid the eggs during the first day after emergence. I did not observe the act of mating, but it presumably took place shortly after emergence and lasted for a brief period of time. Prior to oviposition the female probed over the host plant, apparently to select a suitable site. During oviposition the female was found near a leaf axil with her oviposi- tor extended. The longitudinal axis of her body was orientated to make a narrow angle with the stem. The pro thoracic legs grasped the stem and the other legs sup- ported the body. The metathoracic legs were observed to shake vigorously at inter- vals. The ovipositor was inserted into the leaf axil and the abdomen contracted peri- odically, accompanied by the wriggling movements of the ovipositor. This oviposi- tion lasted four minutes in the laboratory. The adults were never observed to feed, and their mouth parts were vestigial. The males were very active on the first day af- ter emergence and they flew rapidly. They became lethargic and died on the second day. The females usually walked over the host plants after emergence and invariably rested for a few hours underneath the leaves (Fig. 14). They did not fly as swiftly as the males, but were still active on the second day after emergence. The females lived up to three days in the laboratory. Parasites Twenty six of the 160 midge larvae, or 16.2 percent of the larvae examined, were parasitized by an undetermined species of Platygasteridae. However, this sample in- cluded 45 very young larvae, smaller than 0.5 mm, at which stage it was difficult to detect the presence of parasites. This para- site apparently laid its eggs on the midge eggs. The larvae of the parasite were first observed on 3 June 1976 when the host lar- vae had already begun their development in the larval chambers. I never observed more than one parasite per larval chamber. Ei- ther all of the larvae were found in a gall or only part of them were parasitized. The larva of the parasite was invariably found on the midge larva. The earliest para- site larva observed inside a midge larval chamber measured 0.3 x 0.06 mm. It was 436 GREAT BASIN NATURALIST Vol. 37, No. 4 quite active compared to the lethargic midge larva and moved away from light. The host larva was not attacked until the parasite larva reached an advanced stage. The parasitized midge larvae were appar- ently more fragile and damaged easily dur- ing handling. The parasite larva remained colorless up to its maturation, and just prior to pupation it bored into the midge larva near its ante- rior end. The first pupa of the parasite was observed on 30 June 1976. Six of these av- eraged 1.6 X 0.8 mm in dimensions (Fig. 15). The first adult parasite emerged on 18 ■f^« 'm 0»5MM 0-5MM Figs. 10-13: 10, Eclosion of the adult, HA-Haltere, PC-Pupal case, TH-Adult thorax. 11, Pupal exuvium pro- truding out of the gall surface, G.\-Gall, PE-Pupal exuvium. 12, Lateral view of the adult female (ovipositor re- tracted). 13, Lateral view of the adult male. Dec. 1977 ranasinghe: gall midge bionomics 437 June 1976 and they continued to emerge until 5 August 1976. I did not observe any inquilines in the 77 galls dissected in the laboratory. Galls New galls were first observed in the field on 8 July 1976 at the Blue Bell area (eleva- tion 2,700 m) in soil with a temperature of 18° C at a depth of 7.0 cm. The aerial shoots of most of the host plants were dying by this time, but a few young secondary shoots were still active. These young galls, covered by scale leaves, occurred at the bases of the dying aerial shoots and on the young rhizomal branches. Superficially, these appeared very similar to the axillary buds. The galls remained dormant through the fall and winter months to resume their growth the following spring. On 9 May 1977 most of the galls were already grow- ing at Blue Bell in soil at 8° C in temper- ature and at a depth of 2.5 cm. The spring growth of the galls and the host plants was concommitant till the galls reached matu- rity by the time of pupation of the midge, beginning early in June. The adults emerged as soon as the galls began to wither and dry, by late June (Fig. 16). Some of the old galls still remained attached to the host plant even in the following year. The galls were confined either to the bases of the aerial shoots (Fig. 17) or to the active young branches of the rhizome (Fig. 18). No galls were found on the lateral roots (Fig. 18) or on the old rhizome. On the shoots the galls were largely restricted to the basal subterranean areas, but a few were also found protruding just above the soil surface. The depth of the galls in soil varied, depending on the soil texture. In decaying organic matter the galls were closer to the surface than in the loose loamy soil where they were down to a depth of about 4.0 cm. Out of a total of 77 galls examined in the laboratory, 32 (45 percent) were mon- othalamous and the remaining 45 (55 per- cent) were polythalamous. Galls occurred either singly or in clusters (Fig. 19). Single galls were more or less globose and the largest single gall measured 8.5 mm in di- ameter. The largest cluster of galls reached 25.0 mm across. These galls were creamy white in color, but a few had violet-colored areas on them during the early stages of growth. The galls exposed to sunlight devel- oped a green color in the outer tissues. 15 1.0MM 0»SMM Figs. 14-15: 14, Adult female resting underneath a leaf, LE-Leaf. 15, Pupa of the parasite, RM-Remnant of the midge larva. 438 GREAT BASIN NATURALIST Vol. 37, No. 4 The gall tissues initially consisted of moist, turgid cells which became dry after the maturation of the gall. In the poly- thalamous galls there was only one larva per larval chamber. In a given gall all the midges were in about the same stage of de- velopment. However, different galls, even those found on the same host plant simulta- 16 y Innm l! ' 9i 5.0MM Figs. 16-19: 16, Partly dissected midge gall after the emergence of the adults, ST-Stem. 17, Artemisia ludovi- ciana plant bearing midge galls, MG-Midge galls. 18, Young rhizomal branch of Artemisia hidoviciana bearing galls, GA-Gall, YB-Young axillary bud. (Note absence of galls on lateral roots.) 19, Clusters of subterranean bud galls of Rhopalomyia subhumilis from Artemisia ludoviciana, LE-Close view of the host plant leaf. Dec. 1977 ranasinghe: gall midge bionomics 439 neously, contained midges growing in differ- ent stages. The regression coefficient for the relationship between the size of the mature galls and the number of midges per gall is highly significant. This relationship is shown in Figure 20. Out of the 24 galls kept individually in the plastic vials, only female adult midges emerged from seven galls, only male adults from three galls and both male and female adults emerged from two galls. From five other galls only adult parasites emerged. Both female adult midges and adult para- sites emerged from two galls, and none emerged from the remaining five galls. The attempt to induce gall formation by exposing uninfested plants to the newly emerged midges in the laboratory was not successful. However, when a number of in- fested host plants were allowed to grow in- side a growth chamber, new galls were ob- served on the new shoots on 2 September 1976 and adults, possibly belonging to a second generation, were collected from these galls on 22 October 1976. §■ 9' ^6 \i NUMBER OF MIDGES PER GALL Fig. 20. Relationship between the number of midges per gall and the average size of the galls, (1976-77). Discussion These results indicate that the midge galls are modified axillary buds found either at the bases of the aerial shoots or on the new branches of the rhizome. Since only some of the aerial shoots and rhizomal branches had galls on them they are not likely to be the natural outgrowths of the host plants. On the contrary, the young galls and the ax- illary buds are so similar in appearance, it is not possible to distinguish between them without dissection under the microscope. Both the galls and the axillary buds are cov- ered with scale leaves in the early stages of development and become activated about the same time in the spring. The spring and summer activation of the growth is charac- teristic of most of the other bud galls (Mani 1964). The absence of the galls either on the lateral roots or on the old rhizome, both of which are devoid of any active buds, fur- ther supports the view that these galls are modified buds. This view is also in accord with the preference of RJiopalomyia species for the bud galls (Felt 1915, 1940). These growing buds may be preferred by the gall midges because they are well pro- tected, readily supplied with the nutrients, and are in an active meristematic state— all of which are helpful for the formation of the galls (Felt 1936). Growing buds also may offer easily penetrable tissue for the young midge larvae. Subterranean buds in particular provide a microhabitat explored by relatively few other insects and appar- ently by none others in the case of Arte- misia. The comparatively low rate of para- sitism and apparent lack of inquilinism also indicate that .subterranean galls could pro- vide better protection for these midges. The presence of the larvae of these midges in each and every gall dissected in the laboratory indicates that they may be the causative agents. The appearance of the galls in those buds close to the soil surface shows that they are caused by an aerial spe- cies (Mani 1964). Furthermore, the growth patterns of the galls and the midges are closely related. Although the laboratory in- duction of the galls by these midges was 440 GREAT BASIN NATURALIST Vol. 37, No. 4 not successful, their ability to induce galls was indicated by the emergence of a pos- sible second generation of adults in October from those infested plants grown inside a growth chamber. The failure to induce galls was partly due to the asynchrony between the emergence patterns of the limited num- ber of the adult males and females in the cage which resulted in their deaths without mating and the decaying of some of the galls brought from the field, probably due to their continuous contact with the nutri- ent medium. Attempts by Spence (1969) to induce galls of Rhopalomyia hirtipes in the laboratory also failed. Association of bacterial and other types of microorganisms with the galls has been reported by others (Mani 1964). But the only report of bacterial microorganisms from Rhopalomyia galls is that by Farns- worth and Hammond (1968) for the galls of the midges under discussion. The attempts to find either the nitrogen-fixing bacteria by electron microscopy or to confirm the nitro- genase activity from these midge galls failed (Kent, unpublished data). The eggs of these midges are laid in the leaf axils, which apparently provides them with shelter and protection from dessica- tion. The female ovipositor is not adapted to lay the eggs into the host plant tissues and, consequently, eggs were never ob- served inside the host tissues. But the laying of some eggs on the wall of the glass tubes may be an indication of laying them on other inert surfaces like soil. But this was observed imder the artificial conditions in the laboratory, and the eggs of this midge were never recovered from soil in the field. The eggs are comparatively smaller than in the related species and were visible to the unaided eye only when found in clusters. In the field the eggs were found over a period of three weeks, which apparently is respon- sible for the overlap of the different devel- opmental stages. The first stage larva was mobile and pre- sumably induced the galls by invading the axillary buds. The exact mechanism of gall formation is not known. Since these young larvae lack well-developed mouth parts it is unlikely they bored into the host plant tis- sues. The passage of digestive enzymes from the larva to the gall tissues has been report- ed for other midge larvae (Bronner 1970), and it is possible for these midge larvae to secrete enzymes which could digest the host plant tissues, enabling them to enter the ax- illary buds. The larva is the sole trophic stage in the life cycle since the adults were never observed to feed. None of these midge larvae contained any smaller larvae inside them anytime, unlike in the case of the paedogenetic ones. Also, the adults of both sexes emerged from the same gall, which indicated again the lack of paedoge- nesis. Monothalamous galls are unlikely for the paedogenetic larvae, when only one lar- va is present in the mature state, as was found for these midges. Inhabitants of a given gall were at about the same stage of development, which in- dicated the possibility of gall initiation by several larvae simultaneously. But even one larva is capable of gall initiation, as shown by the presence of the monothalamous galls. The young, first instar larvae presumably invaded the axillary buds at random. But since the number of axillary buds suitable for gall initiation would be limited at any given time, it is possible for the larvae from the same egg cluster to attack the same ax- illary bud and, therefore, to be found in about the same stage of development later. The size of a given gall was directly pro- portional to the number of midges found in- side, which indicated a cumulative effect by the larvae which appeared to be the main influencing factor. The gall clusters may be the result either of multiple invasions by the young larvae at different intervals of time or an unusually large invasion by the young midge larvae on the same bud simul- taneously. As expected by the relatively long period of occurrence of the eggs in the field, dif- ferent galls growing at the same time had larvae of different stages. The number of larval stages could not be determined, since the growing larvae did not survive outside the galls. But for most Cecidomyiidae, the Dec. 1977 ranasinghe: gall midge bionomics 441 number of larval stages has been reported to be three (Gagne 1968). Pupation occurred inside the galls as in most other Rhopalornyia species. The pro- trusion of the pupal exuviae perhaps facil- itated the emergence of the newly hatched adult through the soil, although this has been observed for aerial species of midges as well (Spence 1969). This need for the adult to emerge through the soil imposes another restriction on the possible depth of the gall in soil. The vigorous preemergence movements of the pupa could help the pas- sage of the pupal case containing the adult through the gall and soil under the field conditions. The presence of some galls with adult midges belonging to one sex could be due to chance, since the females outnumber the males 2:1. Sex induction by the environ- ment is another possibility, but no apparent relationship between the occurrence of this phenomenon and any one of the likely ex- ternal factors could be found. Jones (1971) JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC l-D^ Fig. 2L Seasonal occurrence of Rhopahmyia sub- humilis developmental stages and aerial shoots of Arte- misia ludoviciana at Philadelphia Flat in 1976. observed a similar relationship between the adult sex and the galls for a related midge species. The short adult life span is common to most gall midges. This short life span, lack of feeding by the adults, and apparent need for mating prior to oviposition could all contribute to the need for quick oviposition and the localized pattern of infestation ob- served in the field. The sex ratio indicates the possibility of multiple matings for the male. Apparently there is one generation of midges per year in the study areas since the eggs and the other developing stages were observed only in one continuous period dur- ing the year and the host plants had one relatively short growing season (Fig. 21). Acknowledgments I gratefully acknowledge the valuable help given by Dr. Stephen L. Wood, Dr. Gary M. Booth, and Dr. Bruce N. Smith. Dr. Raymond B. Farnsworth helped collect the study materials and Dr. Raymond J. Gagne of the Systematic Entomology Labo- ratory, Agriculture Research Service, USDA, identified the specimens. Literature Cited Bronner, R. 1970. Cecidologie— Localisations histo- chemiques de I'invertase et du glucose dans le tis- su nourricier d'une galle de Cecidomyide: Perrisia urticae Ferris sur Urtica dioica L. Note de Mme Roberte Bronner presentee par M. Lucien Plan- tetol. Acad. Sci. Compt. Rend. Ser. D. 270 (4): 641-64.3. CocKERELL, T. D. A. 1909. A new gall-gnat on Arte- misia. Can. Entomol. 41: 150-151. Farnsworth, R. B. 1975. Nitrogen fixation in shrubs. Proc. Symp. and Workshop on wildland shrubs, Provo, Utah 1: ,32-71. Farnsworth, R. B. and M. W. Hammond. 1968. Root nodules and isolation of endophyte on Artemisia ludoviciana. Utah Acad. Sci., Arts, and Letters. 45 (1): 182-188. Farnsworth, R. B. and M. A. Clawson. 1972. Nitro- gen fixation by Artemisia ludoviciana determined by acetylene-ethylene gas assay. Amer. See. Agron. Abs. 96. Farnsworth, R. B., E. M. Romney, and A. Wallace. 1976. Implications of symbiotic nitrogen fixation 442 GREAT BASIN NATURALIST Vol. 37, No. 4 by desert plants. Great Basin Nat. 36 (1): 65-80. Felt, E. P. 1907. Appendix: New species of Cecido- myiidae. N.Y. State Mus. Bull. 110:97-165. 1911a. A generic synopsis of the Itonidae. J. N.Y. Entomol. Soc. 19: 31-62. 1911b. Hosts and galls of American gall midges. J. Econ. Entomol. 4: 451-475. 1915. Appendix: A study of gall midges III. N.Y. State Mus. Bull. (1916).' 180: 127-288. 1916. New western gall midges. N.Y. Entomol. Soc. J. 24: 175-196. 1936. The relation of insects and plants in gall production. Ann. Ent. Soc. Amer. 292: 694-700. 1940. Plant ealls and gall makers. Comstock Publishing Company, Inc. Ithaca, New York. FooTE, R. H. 1965. Family Cecidomyiidae (Itonidae). In: Stone, A. et al. A catalog of the Diptera of America north of Mexico. U.S. Dept. of Agr. Handb. 276. Gagne, R. J. 1968. A taxonomic revision of the genus Asteroniyia (Diptera: Cecidomyiidae). Ent. Soc. Amer. Misc. Pub. 6 (1): 1-40. 1974. A redefinition of Diathronomyia Felt as a subgenus of Rhopalomyia Rubsaamen (Diptera: Cecidomyiidae: Oligotrophidi). Ann. Entomol. Soc. Amer. 68 (3): 484-484. 1975. The gall midges of ragweed, Ambrosia, with descriptions of two new species (Diptera: Cecidomyiidae). Proc. Entomol. Soc. Washington 77 (1): 50-55. 1977. A new species of Rhopalomyia (Diptera: Cecidomyiidae) from Artemisia ludoviciana Nutt. (Compositae), in Utah. Great Basin Nat. 37 (4). Jones, R. G. 1971. Ecology of Rhopalomyia and Dia- thronomyia gall midges (Diptera: Cecidomyiidae) on sagebrush Artemisia spp. in Idaho. Unpub- lished dissertation, Dept. Entomol., University of Idaho, Moscow, Idaho. Kent, S. S. Jr. 1977. Personal communication. Lander, S. 1951. Galls and gall insects of Arteinisia tri- dentata Nutt. Unpublished M.A. thesis, Brigham Young University, Provo, Utah. Mani, M. S. 1964. Ecology of plant galls. Dr. W. Junk. Publishers, The Hague. Paul, J. 1970. Cell and tissue culture, 4th ed. Wil- liams and Wilkins Company, Baltimore. Rubsaamen, E. H. 1892. Die gallmucken des Konigl. Museums fiir Naturkunde zu Berlin. Berlin Ent. Zeitschr. 37: 321-411, pis. 7-18. Spence, a. P. 1969. Biology of the gall midge Rhopa- lomyia hirtipes O. S. (Diptera: Cecidomyiidae) on Solidago juncea Ait. Cornell Univ. Agr. Exp. Sta. Mem. 409. THE ALPINE BUTTERFLIES OF CASTLE PEAK, NEVADA COUNTY, CALIFORNIA Arthur M. Shapiro' .-Vbstract.— The west slope of Basin and Castle peaks and the ridge between them displays a wind-generated tree line below 8500 ft, with dry alpine fell-field vegetation above. The butterfly fauna, although depauperate compared to that of the High Sierra farther south, includes some truly boreal elements such as Oeneis ivallda, Pleheius shasta, and Hesperia nevada. These species commonly occur 1500 to 4000 ft higher in the southern Sierra. The Sierra Nevada of California is about 360 miles long, from the Lassen Peak area in the north to Walker Pass in the south. Along the crest the summits become pro- gressively higher southward— from 6000- 8000 ft in Plumas and Sierra counties to 10,000 ft west of Lake Tahoe, 13,000 ft in Yosemite National Park, and 14,000 ft near Mount Whitney. The term "High Sierra" is properly applied to the region from slightly north of Yosemite to Cottonwood Pass. This area, averaging 20 miles wide over its 150- mile length, includes about 90 percent of the Arctic-Alpine life zone in California. Tree line drops below 10,000 ft from Pyra- mid Peak northward, reaching about 9000 ft on the north side of Lassen Peak (where it is ill-defined due to the rocky substrate), but there are few areas of alpine vegetation in the northern Sierra, and they have not been well studied by biologists. The most exten- sive of them south of Lassen Peak is on the west side of the andesitic mass forming Ba- sin Peak (9015 ft). Castle Peak (9103 ft), and the ridge connecting them. Castle Peak is the second highest in Nevada County and in the Sierras proper, north of Donner Pass. The Sierran Arctic-Alpine butterfly fauna is best known around Tioga Pass (9941 ft), where fell-fields are readily accessible in a day's hike from a good road. Tilden (1958) and Garth and Tilden (1963) have reported in detail on its composition and ecological characteristics. Another fairly well-known, though unpublished, alpine fauna is that of Sonora Peak (11,429 ft), again because of its proximity to a road. Lepidopterists have visited other alpine areas including some to the north, but have not reported on their findings. Although Castle Peak (long known as Stanford Peak or Mt. Stanford) has been collected by botanists for over a century, I can find no record of its having been in- vestigated for butterflies before 1972. This is surprising, since it is an easy day's hike from Interstate Highway 80 (formerly U.S. 40); perhaps the forbidding south face of the mountain, which gives it its name, has given it a more difficult reputation than it deserves (Fig. 1). Again, it has not been widely recognized that an alpine fell-field was present on this mountain; True (1973) states that the Arctic-Alpine life zone does not occur in Nevada County. This is a mat- ter of interpretation, since so many organ- isms are distributed as if it does. The vege- tation, as described below, demonstrates that the summit community is easily consid- ered an alpine fell-field, and not even a very depauperate one. Smith (1973) notes that around Lake Tahoe "scarcely any of the lower peaks between 8500 and 9500 ft are forested at their summits and these sup- port an illusion if not, in fact, the actual plant associations of summit screes and al- pine fell-fields." The Alpine Vegetation Tree line is developed only on the west side and is obviously related to the pre- Department of Zoology, University of California, Davis, California 9561(: 443 444 GREAT BASIN NATURALIST Vol. 37, No. 4 vailing westerly wind. This was noted as early as 1921 by Smiley (p. 57). No mete- orological data are available for Castle Peak, but its orientation and elevation far above the surrounding terrain, the persistent pressure gradient across Donner Pass, and its location overlooking the great canyon of the South Yuba River predispose it to high winds. In my experience it is a very unusual day when the midaftemoon wind on the 4^-- '" '-1., ' - -t*^ Fig. 1. Southwest face of Castle Peak seen from near Castle Pass, showing eroded "badlands" and unvegetated talus. This is the aspect of the mountain visible from the highway. Fig. 2. Trees along the jeep trail below Basin Peak, showing shaping by prevailing westerly wind at about 8000 ft. Fig. .3. Upper edge of subalpine forest belt, showing krwnmhoh of lodgepole pine and mountain hemlock be- low the ridge top south of Basin Peak. Fig. 4. Tree line is clearly visible on the west side below the ridge; note "crawling tree." Photographed from near summit of Basin Peak. Dec. 1977 SHAPIRO: ALPINE BUTTERFLIES 445 ridge top is less than 20 mph. The trees show abundant evidence of battering by the west wind (Figs. 2-4). Matted krwnmholz, consisting of Pinus murrayana Grev. & Balf., P. albicaulis Engelm., and especially Tsuga mertensiana (Doug.) Carr., occurs as much as 350 ft below the ridge top on the west side, and farther down the tops of the trees are commonly bent to the east. On the east side erect, pyramidal trees come almost up to the ridge top. A few nearly prostrate trees occur at low points along the ridge, and one large individual (known locally as the "crawling tree") is pressed almost flat against a 45 degree slope just below the crest. The extensive treeless area which I am calling a fell-field (Figs. 5-8) includes about 100 species of vascular plants. Seventy of these are listed as alpine fell-field in- habitants by Munz (1970) and five are listed only from that community. Of the 24 "in- dicator" plants listed for alpine fell-fields by Mimz (1970: 10-18), 6 occur on Basin and Castle peaks. Smiley (1921) lists some addi- tional alpine "indicators" which occur on Basin-Castle peaks. The alpine flora in- cludes the following distinctive taxa: Arte- misia tripartita Rydb., A. arbuscula Nutt., Railardella argentea (Gray) Gray, Hap- lopappus macronenia Gray, Castilleja am- higua Jones, C. lemmonii Gray, Polemoniiim pulcherrimimi Hook., Gentiana newberryi Gray, Primula suffrutescens Gray, Cassiope mertensiana (Bong.) D. Don., Epilobium oh- cordatum Gray, E. anagallidifoUum Lam., Linum lewisii Pursh., Lupinus lohbii Gray, Astragalus whitneyi Gray, A. austinae GraLy, A. bolanderi Gray, Ivesia gordonii (Hook.) T. & G., Potentilla drummondii ssp. Bruceae (Rydb.) Keck, Sibbaldia procumbens L., Pur- shia tridentata (Pursh.) DC, Saxifraga apri- ca Greene, S. bryophora Gray, Draba crassi- folia Grah., D. densifolia Nutt., Erysimum perenne (Wats.) Abr., Ranunculus eschs- choltzii var. oxynotus (Gray) Jeps., Silene sargentii Wats., Lewisia pygmaea (Gray) Rob., Oxyria digyna (L.) Hill, Eriogonum ovalifolium var. nivale (Canby) Jones, Salix anglorum var. antiplasti Schneid., Carex hel- leri Mkze., etc. In Table 1 the 232 species of vascular plants collected by True (1973) and myself have been categorized by the plant communities in which they are re- corded by Munz (1970). The boreal charac- ter of the vegetation emerges clearly from these statistics. Although only 18 plant spe- cies are shown as being of east-slope distri- bution, the Castle Peak alpine fell-field (like many in the High Sierra) shows a marked east-slope aspect, accentuated by the con- spicuousness of Artetnisia and Purshia. Much of the alpine flora of the Sierra is be- lieved to be evolutionarily derived from the high desert (Chabot and Billings 1972). Af- ter snowmelt the thin, porous alpine soil dries out rapidly; occasional air-mass thun- derstorms in summer temporarily replenish surface moisture. Such thunderstorms are rather more frequent on Castle Peak than in the surrounding area, presumably because its greater elevation promotes the growth of cumulus. I have collected the Basin-Castle peaks area 13 times from 1972 through 1977. The entire area above tree line was collected on two dates: 28 July 1976 and 8 July 1977. The general Donner Pass region has been visited over 100 times since 1972. The Butterfly Fauna Family Satyridae Oeneis ivallda (Mead).— The most charac- teristic Sierran alpine butterfly, ivallda oc- curs from about 7000 ft in the east end of Donner Pass (presumably as a stray) to 14,000 ft in the southern Sierra. Breeding Table L Occurrence of the 232 plant species re- corded from Basin-Castle peaks by plant communities recognized in the Sierra Nevada by Munz (1970). The mean number of categories (not communities) per spe- cies is 2.65. Number Community of Species Sagebmsh scrub (east slope) Yellow pine forest 18 95 Red fir forest 138 Lodgepole pine forest Subalpine forest Alpine fell-field 149 123 70 Many communities (widespread) 22 446 GREAT BASIN NATURALIST Vol. 37, No. 4 colonies exist about 8000 ft on Mt. Judah and Anderson Peak in Donner Pass south of Interstate 80, but the one on Basin-Castle is the only colony known north of that road. There may be a colony on the smaller al- pine summit of Mt. Lola (9143 ft) a few miles to the north. Oeneis ivallda was ex- tremely abundant all along the crest on 8 July 1977, with 250 to 300 individuals seen. It was most numerous in the alpine sage- , ^^^ luf^^^"*'' ^^' ^■^.. •<**' .at It . Fig. 5. Habitat of Oeneis ivallda just below summit of Basin Peak; turf of sedges, grasses, Eriogonwn species, and Astragalus whitneyi. Fig. 6. Habitat of Oeneis ivallda and Hesperia nevada in alpine sagebrush, grasses, and composites below Basm Peak. Fig. 7. Ridge top showing trees on east side and alpine vegetation on west. Large-leaved plant m foreground is Wyethia mollis (Compositae). Fig. 8. Ridge top looking northwest from Castle Peak, showing extent of area above tree line. Dec. 1977 SHAPIRO: ALPINE BUTTERFLIES 447 brush areas and around isolated rock out- crops. It is said to have a two-year Hfe cycle and to be absent in many colonies in even-numbered years, but it was taken on Castle Peak in 1976. Variation is marked, especially in males. The host plant is un- known; 49 species of grasses and sedges are known from its habitat here. Cercijonis oetus (Boisduval).— An east- slope element, this small brown Satyrid is common long after Oe. ivallda in the same places, flying mostly in August and on into early September after most other species have disappeared. It is presumably a grass feeder. Family Nymphalidae Speijeria egleis (Behr).— Males are abun- dant hilltoppers along the crest in July and August, but no females have been collected. This is the only silver-spotted fritillary seen in the alpine fell-fields, which are too dry for S. morrnonia arge (Strecker). Phyciodes campestris montana Behr.— Oc- casional, June to August, but not known to breed. Much more common on dry or moist slopes in the lower subalpine areas. Pohjgonia zephyrus (Edwards).— In- frequent; worn specimens of both sexes (hi- bernators) on the crest in July. Breeding status unknown. This is a very common, widespread species around Donner Pass. Nymphalis californica (Bdv.).— Occasion- al, especially in migration years. There are no Ceanothus (Rhamnaceae) on the ridge top, but several species used by this insect occur in subalpine areas below. Nymphalis milberti (Godart).— Three seen (one taken) on the ridge top, apparently hilltopping, 8 July 1977. The host plants {Urtica spp.) do not occur above tree line. A female found on Basin Peak, 9 August 1977. Vanessa virginiensis (Dniry).— Occasional, July to September. Three potential host plants are recorded, but no larvae have been found. Vanessa annabella (Field).— Occasional, July to August. A female was taken on 8 July 1977, and the usual host plant at Soda Springs, Sidalcea glaucescens Greene (Mal- vaceae), occurs on Basin Peak. Family Lycaenidae Satyrium fuUginosum (Edwards).— Com- mon all along the ridge top, second half of July. The butterflies scarcely fly at all un- less disturbed, and they may be seen walk- ing on Eriogoniim and yellow composite flowers on windy afternoons. Four species of lupines occur on the summit, but I have been unable to establish an association. Callophrys lemberti Tilden.— Frequent, mid-June to mid-July, among Eriogonum plants and, like the last species, seldom seen flying. The host is presumably an Eriogo- num; eight taxa are recorded here. Lycaena nivalis (Bdv.).— Occasional on the ridge top among the Eriogonum or on composites, but less common than in the subalpine areas along the trail to Castle Pass. Late June to early August. Lycaena heteronea Bdv.— Frequent in "rock gardens," second half of July, presum- ably feeding on an Eriogonum. Females are quite dusky. Everes amyntula (Bdv.).— In the northern Sierra and the Trinity Alps occurs a sub- alpine-alpine amyntula which is larger and duller than the montane coniferous forest populations associated with native Vicia and Lathyrus at 1000 to 5000 ft. This animal, which occurs at 7500 ft in Trinity County and above 8500 ft in the Donner Pass area (Mt. Judah, Anderson Peak, Basin-Castle) is associated with Astragalus whitneyi Gray (Leguminosae) and oviposits in the in- florescences. Like other lycaenids on Castle Peak, these animals seldom fly and may have to be scared up from the plants. They occasionally visit flowering Eriogonum. Everes amyntula is done flying before Plebeius shasta (Edw.) begins flying in the same places; its peak flight is in early July. Philotes enoptes (Bdv.).— A single aberrant male was taken on 8 July 1977 along the ridge top. It was not associated with an Eriogonum. This species is so sedentary that a breeding colony must be assumed to exist. Celastrina argiolus echo (Edw.).— Occa- 448 GREAT BASIN NATURALIST Vol. 37, No. 4 Fig. 9. Characteristic alpine butterflies from Castle Peak area: a, b, light and dark male Oeneis ivallda; c, fe- male Oe. ivallda; d, male Hesperia nevada; e, female Satyriwn fuliginosum; f, g, Plebeius shasta male and female; h, i, Pieris occidentalis male and female. Dec. 1977 SHAPIRO: ALPINE BUTTERFLIES 449 sional, June to mid-July, both sexes. Breed- ing status uncertain. Much commoner in the subalpine zone lower on the mountain, where its hosts (especially Ceanothus) are common. Plebeius saepiolus Bdv.— Occasional, July. Two clovers {Trifolium monanthum Gray, T. productum Greene) occur on Castle Peak. This is one of the most common spe- cies in the meadow at Castle Valley. Plebeius icarioides (Bdv.).— Common tliroughout July. On the ridge associated with Lupinus arhustus Dough, ssp. silvicola (Hell.) D. Dunn. Plebeius shasta (Edw.).— Like Oeneis ivallda, this is a characteristic Sierran alpine butterfly. It is incredibly abundant from mid-July through mid-August, and very var- iable, although consistently larger than ma- terial from Inyo and Mono counties. It oviposits on pedicels of Astragalus whitneiji Gray, often on the same plants on which E. amyntula laid three weeks before. Plebeius acmon (Westw. & Hew.).— This ubiquitous, weedy species occurs as a stray and may breed occasionally, but does not overwinter. Recorded from July to Septem- ber. Plebeius lupini (Bdv.).— Common in "rock gardens," where it is associated with Eriogo- num umbellatwn var. covillei (Small) Munz & Reveal, which is also the host plant in Donner Pass. Late June to mid-July. Glaucopsyche lygdamus behrii (Edw.).— Resident but not common, late June to early July. Associated with Lupinus pohj- phyllus Lindl. ssp. superbus (Hell.) Munz. in a seep area on Basin Peak. females are not uncommon. Very variable. Eggs and larvae have been collected on the following Cruciferae: Arabis holboelli Horn, vars. pinetorum (Tids.) Roll, and retwfracta (Grah.) Rydb., A. platyspemia Gray, Descu- rainiu richardsonii ssp. viscosa (Rydb.) Detl., and Draba crassi folia Grah. This is the only resident butterfly on Castle Peak that may be partially double-brooded (some years). (Note: P. protodice Bdv. & LeC. has not been seen on Castle Peak but was recorded in the subalpine zone in 1972 and again in 1977.) Anthocaris sara Stella Edwards.— Occa- sional as a hilltopper, males only seen; breeding status uncertain. Here, as in Don- ner Pass (Emmel and Emmel 1962, Shapiro, unpublished), occasional white males in- distinguishable from A. s. sara Lucas are found. Eastward and lower, at Truckee, only the yellow form occurs. This situation deserves further attention. Euchloe ausonides Lucas.— One male, 8 July 1977 on the ridge top. This small race of ausonides is about the same size as the next species, flies with and a bit after it, and is commonly confvised with it. I have it from the Trinity Alps to Eldorado County, 5000 to 9000 ft. It may be recognized by the thinner bar at the end of the forewing cell; somewhat duller white color; less in- tense green mottling below; and more angu- lar hindwing. It seems always to be rare. Euchloe hyantis (Edw.).— A common hill- topper, mid-June to mid-July. It is not un- common to find hyantis, sisymbrii, and occi- dentalis flying together around the same rocky crag on Castle Peak. Family Pieridae Colias eurytheme Bdv.— A common flyup, perhaps breeding but never overwintering; all season. Pieris sisymbrii Bdv.— Frequent, late June to mid-July. Males hilltopping; females not seen, and breeding status uncertain; females have been collected in Donner Pass. Pieris occidentalis Reakirt.— The most common butterfly on the ridge top, flying mid-June to September. Males hilltop, but Family Papilionidae Papilio zelicaon Lucas.— Common. Males hilltop, and females are occasionally en- countered (especially on the ridge from Castle Pass to Castle Peak). Late June to early August. Host plant unknown; presum- ably one or more of the six recorded Um- bellifers. Papilio indra Reakirt.— Frequent. Males hilltop, and females are occasionally seen on the crest and also on the ridge to Castle 450 GREAT BASIN NATURALIST Vol. 37, No. 4 Fig. 10. Characteristic alpine butterflies from Castle Peak area: a, male Papilio zelicaon; b, female Papilio in- dra; c, male Pieris sistjinbrii; d, male Euchloe ausonides; e, female Nyrnphalis milberti; f, g, Everes amijntula male and female; h, female Lycaena heteronea; i, male Polites sahtdeti tecumseh. Dec. 1977 SHAPIRO: ALPINE BUTTERFLIES 451 Pass. Late June to mid-July. Oviposition ob- served on Loynatiiim dissectum var. midti- fidiim (Nutt.) Math. & Const. Papilio eunjmedon Lucas.— Occasional male hilltopper, July; the host plants do not occur above tree line. Parnassius clodius baldur Edwards.— Oc- casional, both sexes, July, probably flyups from the subalpine area where it is quite common. Family Hesperiidae Hesperia juba (Scudder).- Occasional, June and September; breeding status un- known. A very widespread species around Donner Pass. Hesperia )mrpalus ijosemite Leussler.— Only male hilltoppers thus far recorded, second half of July, occasional. Hesperia nevada (Scudder).— Occasional on the ridge top and crags, both sexes, first half of July. This very dark, small, shaggy skipper is a true alpine species in the Don- ner Pass region. The Emmels (1962) found it on the summit of Mt. Judah (8243 ft). Polites sabideti teciimseh (Grinnell).— In- frequent, grassy area below ridge top, first half of July. Much more common in the subalpine meadows, where it oviposits on Agrostis scabra Willd. and perhaps other grasses. Pyrgus communis (Grote).— Common, first half of July, ovipositing on Sidalcea glauces- cens Greene (Malvaceae). Pyrgus niralis (Bdv.).— Infrequent; grassy open places on the ridge mid-June to early July. Host plant unknown. Erynnis propertius (Scud. & Burg.) — Oc- casional hilltopper, males only. July. Com- mon in subalpine areas. Thorybes nevada Scudder.— Occasional on the ridge as a hilltopper, probably not breeding; late June to mid-July. Abundant in Round and Castle valleys. Comparisons to Tioga Pass Tilden's Tioga Pass study area is much more ecologically diverse than Basin-Castle Peak. It incTudes a spread of over 3000 ft in elevation, moraines, boggy areas, and wet meadows above tree line. He records 43 species, as against 40 in the present study; 5 are alpine obligates, as against 3 at Castle Peak. Missing at Castle Peak are the alpine species Chlosyne damoetas malcolmi (Com- stock), Colias behrii (Edwards), and Hesperia miriamae MacNeill. All of these are High Sierran endemics unrecorded north of Yose- mite. Some species found at Tioga Pass above tree line are strictly subalpine species far- ther north around Donner Pass. These in- clude Speyeria mortnonia arge (Strecker), Lycaena editha (Mead), L. mariposa Reakirt, L. cupreus (Edwards), and Agriades glandon podarce (Felder and Felder). One species, Lycaena rubidiis (Behr), is a common mem- ber of the alpine fauna at Sonora Peak, oc- curs in Donner Pass, and may yet turn up at Castle Peak where at least one of its known hosts, Oxyria digyna (L.) Hill (Poly- gonaceae), occurs (see Johnson and Balogh, 1977). It is basically an east-slope animal. Another species found on Sonora Peak but imrecorded at either Tioga Pass or Castle Peak is the "alpine" member of the Plebeius melissa (Edwards) complex. At Sonora Peak and also on Gumboot Butte, far to the north in the Trinity Divide, it occurs on As- tragalus ivhitneyi Gray, which is one of the most common plants on the Basin-Castle ridge top. Farther northwest, on Packer's Peak in the Trinity Alps, it is replaced on this plant by an ecotype of P. argyrognomon anna (Edwards), but in the Donner Pass re- gion this insect occurs in subalpine marshy habitats on Lupinus polyphyUus Lindl. and has never been found at Castle Peak or on Astragalus. Plebeius melissa is pre- dominantly an east-slope species. Multi- voltine "Great Basin" melissa has been found in Donner Pass (Shapiro, unpub- lished). Another east-slope species, Satyrium behrii (Edw.), which occurs at Truckee and Donner Pass, has not been found on Basin- Castle although its host, Purshia, is common there; it is not usually an alpine butterfly in the northern Sierra. The Donner Pass fauna as a whole is about 70 to 75 percent univoltine. Tilden claims that of the resident (as against colo- 452 GREAT BASIN NATURALIST Vol. 37, No. 4 nizing, nonwintering) tauna of Tioga Pass only Polites sabuleti tecumseh is bivoltine, but this claim is spurious (see Shapiro 1975). At Castle Peak one breeding resident iPieris occidentalis) and one whose status is un- known {Hesperia juba) appear bivoltine. Both are probably bivoltine at least in the subalpine zone at Tioga Pass as well. Of the remaining 38 species at Castle Peak, about two-thirds are "early" season emergents (e.g., Oeneis ivallda, Everes arnyntula, Hes- peria nevada), appearing about three to five weeks after bare ground appears. The re- maining third are "late" {Plebeius shasta, Lycaena heteronea, Cercyonis oetus). In the case of E. arnyntula and P. shasta, which have the same host plant, this seasonal dif- ference may facilitate coexistence. The flight season in a "normal" year is about 100 days, from late Jime to late September. Only one species, Pieris occidentalis, a fa- mous hilltopper, covers most of that period. One alpine species found elsewhere in the Donner Pass vicinity {Parnassiiis phoebus behrii Edw.) has not been found on Basin- Castle Peak, apparently because its host, Se- dum obtusatiim Gray, is extremely rare on the andesite substrate. It is common on granite, and the butterfly is thus effectively edaphically restricted. It ranges north to Plumas County, where it occurs at consid- erably lower elevations. The isolated character of alpine "islands" in the northern and north-central Sierra makes them ideal subjects for biogeographic analysis. As Lepidopterists become less au- tomobile-bound, increasing amounts of data may become available which will help in the eventual formulation of a biogeographic synthesis of the sources of our high-eleva- tion fauna. In the meantime, the elevational and associational flexibility of so many Sier- ran butterflies underscores the applicability of the "individualistic" approach to their distributions, rather than a rigidly "zonal" approach such as is commonly found in the butterfly literature. Acknowledgments Some of the data included here were ob- tained by Adrienne R. Shapiro, Frances Sharpies, Cheryl Palm, and Kenneth Ma- suda. Help was also received from Dr. E. W. Jameson, Jr., Allen Allison, and Dr. Jack Major. This research was partially sup- ported by grant D-804 from the Committee on Research, U.C. Davis. Literature Cited Chabot, B. F., and W. D. Billings. 1972. Origins and ecology of the Sierran alpine flora and vegetation. Ecol. Monogr. 42: 163-199. Emmel, T. C, and J. F. Emmel. 1962. Ecological stud- ies of Rhopalocera in a high Sierran community— Donner Pass, California. I. Butterfly associations and distributional factors. J. Lepid. Soc. 16: 23-44. Garth, J. S., and J. W. Tilden. 1963. Yosemite but- terflies. J. Res. Lepid. 2: 1-96. Johnson, K., and G. Balogh. 1977. Studies in the Lycaeninae (Lycaenidae). 2. Taxonomy and evolu- tion of the Nearctic Lycaena rubidus complex, with description of a new species. Bull. AUyn Mus. 43: 1-62. Munz, p. a. 1970. A California flora. Univ. Calif. Press, Berkeley, Los Angeles and London. 1681 pp. Shapiro, A. M. 1975. Genetics, environment, and sub- species differences: the case of Polites sabuleti (Lepidoptera: Hesperiidae). Great Basin Nat. 35: 33-38. Smiley, F. J. 1921. A report upon the boreal flora of the Sierra Nevada of California. Univ. Calif. Publ. Bot. 9: 1-423. Smith, G. L. 1973. A flora of the Tahoe Basin and neighboring areas. Wasmann J. Biol. 31: 1-231. Tilden, J. W. 1958. The butterfly associations of Tioga Pass. Wasmann J. Biol. 17: 249-271. True, G. H. 1973. The ferns and seed plants of Ne- vada County, California. Privately circulated ms, 63 pp. GEOGRAPHIC VARIATION IN CONTOPUS SORDIDULUS AND C. VIRENS NORTH OF MEXICO M. Ralph Browning' ABSTRA.CT.— Geographic variation in plumage color between populations of Contopus sordidulus is minor. The differences in color between the races described by Burleigh (1960) is largely the result of postmortem color changes of museum skins. Variation in wing chord and length of tail between samples does not support recogniz- ing additional races of C. sordidulus. Contopus virens is monotypic. The relationships between sordidulus and vi- rens is unclear. In a review of Contopus sordidulus (Western Wood Pewee) Burleigh (1960) proposed four nomenclatural subdivisions for the populations of the species north of Mexico. He recognized saturatus Bishop, 1900, and subdivided veJiei Coues, 1866, into three races, veliei and two new races, cimplus and siccicola. This subdivision was considered untenable by Mayr and Short (1970), who suggested that the only valid race north of Mexico is veliei. The AOU (1957) Check-list recognized both veliei and saturatus as races of the northern popu- lations of C. sordidulus. To determine the extent of geographic variation in the northern populations of C. sordidulus I compared Burleigh's type series of amplus and siccicola and other specimens for variation in color and size. Specimens used in this study include those collected in June. Although birds collected at this time and earlier could be migrants, those collect- ed later are usually too badly worn to per- mit meaningful color comparisons (Phillips and Parkes 1955, Phillips et al. 1966: 169, pers. observ.). All measurements (Table 1) are in millimeters. Nomenclatural history of contropus sordidulus veliei and c. s. saturatus Coues (1866) indicated that two color types were among his series from the south- western United States and that Baird be- 'National Fish and Wildlife Laboratory, U.S. National Museum of Na lieved these to represent two species. Al- though Coues was not so convinced, he nevertheless introduced the name Contopus veliei for the southwestern form and gave a diagnosis to distinguish it from C. richard- sonii, as C. sordidulus was then known (see Phillips and Parkes 1955). The name veliei as currently used is the equivalent of C. r. richardsonii of older usage. Parenthetically, Burleigh (1960) is incorrect in stating that Phillips and Parkes had overlooked Coues' lectotype of veliei. As pointed out by Phil- lips and Parkes (1955) and Parkes {in litt., 1976), the only locality in Coues' original description is Colorado, from whence Velie sent the first example. The ex post facto se- lection of a lectotype has no bearing on the fact that Coues had already stated that the first example came from Colorado. Thus, the specimen from Fort Whipple, Arizona (chosen by Coues at Richmond's prompt- ing), considered by Deignan (1961) as the lectotype of veliei, cannot be regarded as a primary type. The type locality of veliei is the mountains of Colorado as stated by Phillips and Parkes (1955). The change in the type locality of veliei has no nomencla- tural effects. Contopus sordidulus veliei was recognized by Phillips and Parkes (1955) and the AOU (1957) Checklist as the race breeding from central eastern Alaska south through south- ern Mackenzie, northeastern Alberta, north- western and central Saskatchewan, and cen- tral Manitoba, the eastern parts of North History, Washington, DC. 20560. 453 454 GREAT BASIN NATURALIST Vol. 37, No. 4 and South Dakota to central and western Texas. The AOU Checklist gives the west- ern boundry of veliei as from Alaska south through eastern British Columbia to north- ern Mexico. This agrees with Webster (1957), who considered placens van Rossem, 1940, type locality in Arizona, a synonym of veliei. The second northern race, Contopus sor- diduhis saturatus, was described by Bishop (1900) as darker dorsally than the other forms of the species. Bishop considered sat- uratus to be the race breeding from the Yu- kon Valley in Alaska south along the coast ranges to British Columbia. Ridgway (1907) did not recognize saturatus because dark birds occurred outside the range ascribed to saturatus, that is, within the breeding range of what is currently known as veliei. Al- though recognized by van Rossem (1945), saturatus was not generally accepted as a distinct race imtil subsequent studies by Aldrich {in Jewett et al. 1953) and Phillips and Parkes (1955). The breeding range of saturatus in the AOU (1957) Checklist is given as southwestern Alaska south through western British Columbia and west from the Cascade Mountains in Washington and Ore- gon. Variation in Color Burleigh (1960) subdivided veliei into three races on the basis of size (see below) and color. In so doing he restricted the range of veliei to northern California, Ne- vada, Utah, and Colorado south through Arizona, New Mexico, western Texas, north- ern Baja California, eastern Sonora, and northern Chihuahua. He claimed that the two new races breed north of this region. Contopus s. siccicola was said to breed from southern British Columbia east of the coast ranges, and east of the Cascades in Wash- ington and Oregon to western Montana west of the Rocky Mountains. The second new race, arnplus, was said to breed from central Alaska, southern Mackenzie, Sas- katchewan, and southern Manitoba south through British Columbia, northeastern and eastern Montana to Wyoming, and in west- em North and South Dakota. Contopus s. siccicola was described by Burleigh as the palest of the four northern forms. The up- per parts in siccicola were said to be washed with gray, whereas Burleigh consid- ered amplus to be browner, and his veliei to be fuscous-olive. The imderparts of siccicola were also said to be more gray and the bel- Table 1. Measurements of adult males of Contopus sordidiihis and C. virens. Wing Chord TaU range C.V. range saturatus 11 83.4-88.1 86.11 1.53 1.78 62.4-67.8 64.89 1.71 2.64 California' 20 82.4-89.6 85.63 1.87 2.18 61.6-69.7 64.77 2.18 3..37 siccicola 12 83.3-89.0 86.97 1.65 1.90 64.6-69.7 66.28 1.58 2.38 Nevada 8 85.0-88.7 86.60 1.51 1.74 65.0-70.0 66.70 1.59 2.38 Arizona 18 85.2-88.8 86.82 1.21 1.39 63.4-68.9 65.92 1.52 2.31 amplus- 14 83.6-90.6 87.69 2.22 2.53 62.6-71.1 67.09 2.82 4.20 northwestern C. virens' 15 82.0-88.3 85.17 2.28 2.68 63.5-71.8 68.20 2.39 3.50 eastern C. virens' 14 82.0-88.5 84.59 1.76 2.08 ft3.9-70.6 66.95 2.16 3.23 'Specimens from south of latitude 40 degrees 'Montana and Wyoming 'North and South' Dakota, Minnesota, Wisconsin 'East of Appalachij m Mountains Dec. 1977 browning: bird variation 455 ly more white (less yellow) than in veliei and ampins. Burleigh described ampins as differing from both siccicola and veliei by being darker and "distinctly" more brown and less olivaceous dorsally, and differing from his veliei by being paler and less yel- low ventrally. Burleigh probably believed siccicola to be paler and grayer than the other forms of C sordiduhis for the reasons mentioned in Mayr and Short (1970)— the museum ages of the type series of ampins and siccicola and the comparative series of his veliei are not similar (cf. Burleigh 1960: 143-146). The two dark races, ampins and veliei, were col- lected, on the average, 25 years earlier than birds assigned by Burleigh to the paler series of siccicola. The most recently col- lected specimens in Burleigh's series were collected in the 1950s with only 7 percent of these specimens assigned to veliei and 11 percent assigned to ampins, whereas 44 per- cent of the specimens assigned to siccicola were collected after 1950. As would be ex- pected, because of the nature of post- mortem color changes, the darker and browner specimens were collected much earlier than the cleaner and paler speci- mens. Comparison of the type series of ampins with similarly old specimens of veliei used by Burleigh does not reveal sufficient differ- ences in coloration to justify subspecific separation. Comparison of the more recent- ly collected specimens making up the type series of siccicola with the series of ampins and veliei used by Burleigh reveals ampins to be, on the average, slightly paler. Using specimens of C. sordidnlns of similar mu- seum age from the ranges of the three named races, however, reveals that the dif- ferences in palor between siccicola and veliei and ampins is the result of post- mortem color changes. Specimens represent- ing the three forms cannot be separated on the basis of dorsal coloration. I found that the underparts of siccicola are not noticeably paler than the other forms of Contopns sordidnlns {contra Bur- leigh 1960). Variation in ventral coloration in C. sordidnlns appears to be clinal from west to east, with the coastal form, satnrat- ns, being slightly darker and more yellow. The paleness increases gradually eastward toward the Rocky Mountains. Specimens from the northwestern slope of the northern Rocky Mountains, however, are slightly darker. Nonetheless, ventral coloration has not proven to be useful for distinguishing between the northern forms of C. sordi- dnlns. In a recent study of the races of migrant C. sordidnlns in Utah Behle (1967) consid- ered siccicola and ampins as valid races. Behle followed Burleigh's suggestion that some individuals of southern populations be- gin nesting while other populations are still en route to more northern localities. This was a reasonable assumption based on his limited sample and is a convenient means of disposing of the considerable amount of in- dividual variation in dorsal coloration. For example, I found that slightly pale gray birds and darker examples occur in adjacent areas in June. Although some individuals of C. sordidnlns may be migrating north as late as June, the lack of consistent geo- graphic variation in color among Burleigh's forms of the species argues against recogni- tion of additional races. The relationships of Contopns sordidnlns populations to those of Contopns virens (Eastern Wood Pewee), a monotypic species of eastern North America, not mentioned by Burleigh (1960), were also examined. These two species are morphologically similar. Phillips et al. (1966) have pointed out that adult virens are typically olivaceous and that sordidnlns is slightly darker and brown- er in plumage coloration and usually has a darker lower mandible. I compared speci- mens of sordidnlns collected in eastern Montana and Wyoming with a series of vi- rens from eastern North Dakota and Wis- consin. The two series may be readily sepa- rated by the browner upper parts of sordidnlns. However, both series are con- sistently olivaceous above. Specimens of C. sordidnlns from west and south of Montana and Wyoming are not as olivaceous as spec- imens of C. virens and C. sordidnlns from eastern Montana and Wyoming. 456 GREAT BASIN NATURALIST Vol. 37, No. 4 Variation in Mensural Characters Measurements of wing chord and length of tail of adult male Contopus sordiduhis and C. virens (Table 1) show considerable overlap in these characters with nearly identical means for the geographical sam- ples. Burleigh's (1960) characterization of amphis as longer-tailed than other forms of C. sordiduhis is not upheld. Conclusions Geographic variation in Contopus sordi- duhis north of Mexico is slight. The amount of variation in color and size is shown to be insufficient for recognizing amphis and sic- cicola. These names, along with pkicens, should be considered synonyms of C. sordi- duhis veUei. The form saturatus should be recognized, although it is a weakly marked race. There is no pronounced geographic variation in C. virens. Overlap in coloration between the north- western samples of C. virens and represen- tatives of C sordiduhis from eastern Mon- tana and Wyoming suggest a close relationship between the two species. The breeding ranges of these species overlap in western Manitoba, Nebraska (Phillips et al. 1966), and in Kansas (Barlow and Rising 1965). Van Rossem (1940, 1945), Rand (1948), Aldrich (in Jewett et al. 1953), and Meyer de Schauensee (1966) have consid- ered virens and sordiduhis conspecific. Evi- dence of interbreeding is inconclusive, but hybridization is mentioned by Mayr and Short (1970) as occurring in the Great Plains. It is premature at this time to con- sider virens and sordiduhis as conspecific. Field studies, urged by Phillips and Parkes (1955), where the two species come in con- tact, are still needed. Such studies should be conducted on territorial individuals in late May and June, during which time birds are retaining plumage suitable for color com- parisons (A. R. Phillips, in htt.). Acknowledgments I am indebted to the American Museum of Natural History for providing the op- portunity to examine specimens. 1 am espe- cially grateful to Kenneth C. Parkes and Al- lan R. Phillips who read the manuscript and offered many helpful suggestions. I also thank John Farrand, Jr., Alfred Gardner, and Roger Clapp, who read an earlier draft. Literature Cited American Ornithologists' Union. 1957. Check-list of North American Birds. 5th ed. Baltimore, Mary- land. 691 p. Barlow, J. C, and J. D. Rising. 1965. The summer status of wood pewees in southwestern Kansas. Kansas Omithol. Soc. Bull. 16: 14-16. Behle, W. H. 1967. Migrant races of Western Wood Pewee in Utah. Auk 84: 133-134. Bishop, L. B. 1900. Descriptions of three new birds from Alaska. Auk 17: 113-120. Burleigh, T. D. 1960. Geographic variation in the Western Wood Pewee (Contopus sordidulus). Proc. Biol. Soc. Washington 73: 141-146. CouES, E. 1866. List of the birds of Fort Whipple, Arizona: with which are incorporated all other species ascertained to inhabit the Territory; with brief critical and field notes, descriptions of new species, etc. Proc. Acad. Nat. Sci. Philadelphia: .39-96. Deignan, H. G. 1961. Type specimens of birds in the United States National Museum. U.S. Natl. Mus. Bull. 221. 718 p. Jewett, S. G., W. P. Taylor, W. T. Shaw, and J. W. Aldrich. 1953. Birds of Washington State. Univ. Washington Press, Seattle. 765 p. Mayr, E., and L. L. Short. 1970. Species taxa of North American birds. Nuttall Omithol. CI., No. 9, 127 p. Meyer de Schauensee, R. 1966. The species of birds of South America and their distribution. Acad. Nat. Sci. Philadelphia. 577 p. Phillips, A. R., and K. C. Parkes. 1955. Taxonomic comments on the Western Wood Pewee. Condor 57: 244-246. Phillips, A. R., M. A. Howe, and W. E. Lanyon. 1966. Identification of the flycatchers of eastern North America, with special emphasis on the genus Empidonax. Bird-Banding 37: 153-171. Rand, A. L. 1948. Birds of southern Alberta. Natl. Mus. Canada Bull. No. 111. 105 p. Ridgway, R. 1907. Birds of North and Middle Ameri- ca. Pt. 4. U.S. Natl. Mus. Bull. 50. 973 p. VAN RossEM, A. J. 1940. Notes on some North Ameri- can birds of the genera Myiodynastes, Pitangus, and Myiochanes. Trans. San Diego Soc. Nat. Hist. 9: 79-86. 1945. A distributional survey of the birds of Sonora, Mexico. Occ. Papers Mus. Zool., Loui- siana State Univ., No. 21. .379 p. Webster, J. D. 1957. A new race of wood pewee from Mexico. Indiana Acad. Sci. 66: 337-340. DISTRIBUTIONAL PATTERNS OF MAMMALS IN UTAH David M. Armstrong' Abstract.— Based on a synthesis of recent work on distribution of mammals in Utah, the hierarchy of eco- geographic distributional units proposed by Durrant (1952) is reevaluated by numerical methods. Areographic faunal elements, distinguished on the basis of shapes of distributional ranges in North America, are identified. Relationships are shown between ecogeographic faunal units and areographic faunal elements, and their historical implications are discussed. Biogeography seeks to describe patterns in the landscape and to understand their ev- olution. Utah provides a study area of con- siderable interest to the zoogeographer in- terested in faunal movements and effects of corridors, barriers, and isolation. The state is large (nearly 85,000 sq mi or 220,000 km') and includes a wide range of ecological conditions, from hot desert to alpine tundra. Mean elevation is roughly 6100 ft (1860 m) and the range of relief is from about 2000 ft (610 m where Beaverdam Wash leaves the state in the southwest) to nearly 13,500 ft (4115 m at the summit of Kings Peak in the Uinta Mountains). A north-south "archi- pelago" of mountains and high plateaus di- vides Utah roughly in half. The eastern part is drained by the Colorado River and its tributaries, which have carved horizontal sedimentary formations into an intricate landscape of basins and canyons. West of the central highlands lies the Great Basin, a complex area of minor mountain ranges and internal drainage, dominated by the vast bed of Pleistocene Lake Bonneville. The only previous zoogeographic analysis of mammals of Utah is that of Durrant (1952), who distinguished "faunal areas" in the state. These were ecogeographic units, roughly comparable to the biotic provinces of Dice (1943). Faunal areas were recog- nized subjectively, by the coincidence be- tween mammalian distributions and phys- iographic units. Durrant (1952: 480) pointed out that faunal areas tended to be centers of differentiation for subspecies. Kelson (1951) discussed the influence of the Colo- rado River and its major tributaries on dif- ferentiation and distribution of rodents, re- fining Durrant's work on faunal areas. Marshall (1940) studied ecological biogeo- graphy of mammals on islands in the Great Salt Lake. Lee (1960) investigated the mon- tane mammals of several mountain ranges in southeastern Utah in an effort to under- stand faunal relationships among the high- land faunas and the effects of Pleistocene climatic change on the patterns observed. Brown (1971, in press) studied montane mammals of Utah as an example of insular biogeography. Armstrong (1973) discussed zoogeographic relationships of mammals in Canyonlands National Park, which lies as- tride the confluence of the Colorado and Green rivers. This work in southeastern Utah suggested some intriguing local pat- terns of ecological and historical biogeo- graphy, but the existing literature was in- adequate to place the area in a broader context. The present paper is meant as a partial answer to this need. Its purpose is to refine ecogeographic analyses of previous authors and to provide an areographic anal- ysis of the mammals of Utah. Methods Analyses of range limits in Utah were based on maps of 92 species. Seventeen spe- cies range essentially statewide in suitable 'Department of Integrated Studies and University Museum, University of Colorado, Boulder, Colorado 80309. 457 458 GREAT BASIN NATURALIST Vol. 37, No. 4 habitat and hence were not mapped: Myotis hicifiigiis, Myotis volans, Myotis evotis, Myotis leihii, Lasionycteris noctovagans, Ep- tesicus fitscus, Lasiiirus cinereus, Plecotiis townsendii, Peromysciis maniculatus, Erethi- zi)n dorsatwn, Canis latrans, Bassariscus as- tutus, Mustela frenata, Taxidea taxus. Mephitis mephitis, Spilogale putorius, and Lynx riifus. (These species are inckided in analyses of faiinal areas.) Twelve species are known from too few localities to allow their ranges in Utah to be outlined with any con- fidence: Sorex merriaini, Sorex nanus, No- tiosorex crawfordi, Myotis velifer, Euderma maculatum, Idionycteris phyllotis, Tadarida macrotis. Ondatra zibethicus, Procyon lotor, Maries pennanti, Liitra canadensis, Odo- coileus virginianus. Three species {Canis hipus, Ursiis arctos, and Bison bison) have been extirpated in Utah over the last 125 years; limits of their former ranges are un- known. In addition, the natural ranges of Antilocapra americana and Ovis canadensis have been altered to an unknown extent since the advent of European civilization in Utah. Range maps for mammals in Utah gener- ally were based on those published by Dur- rant (1952) and refined using distributional records from the following more recent pa- pers: Anderson 1955, 1959, Armstrong 1972b, 1974, Behle and Hansen 1951, Ben- son 1954, Black 1970, Brown 1971, in press, Dearden and Lee 1955, Durrant and Han- sen 1954, Durrant and Newey 1953, Dur- rant and Lee 1955, 1956, Durrant, Lee, and Hansen 1955, Durrant and Dean 1959, 1960, Easterla 1965, 1966, Egoscue 1961, 1966, Egoscue and Lewis 1968, Hansen 1953, Harris 1974, Hayward and Killpack 1956, 1958, Hennings and Hoffmann, in press, Genoways and Jones 1967, Keegan 1953, Killpack 1955, Knitsch and Heppens- tall 1955, Lee 1960, Lee and Durrant 1960, Lowery and Egoscue 1968, Miller and Kel- logg 1955, Musser and Durrant 1960, Nich- ols et al. 1975, Poche 1975, Poche and Bail- ie 1974, Poche and Ruffner 1975, Rasmussen and Chamberlain 1959, Shippee and Egoscue 1958, Stock 1970, Thaeler 1972, Wauer 1966, White 1953, Wood 1958. An attempt was made to make range maps at least roughly comparable in their level of resolution. As a check on the re- drafted range maps, range limits in Utah were compared to those reported in adja- cent states of Idaho (Davis 1939), Nevada (Hall 1946), Arizona (Cockrum 1960), Wyoming (Long 1965), Colorado (Arm- strong 1972a), and New Mexico (Findley et al. 1975) and refined where appropriate. Maps of continental ranges were based on those in Hall and Kelson (1959) with limits in Utah and adjacent states refined on the basis of more recent publications (Cockrum 1960, Long 1965, Armstrong 1972a, Findley et al. 1975) and maps of ranges in Utah prepared for the present paper. Nine species have been documented in Utah since Durrant's (1952) checklist was published: Sorex cinereus (Durrant and Newey 1953), Sorex nanus (Durrant and Lee 1955), Notiosorex crawfordi (Wauer 1966), Myotis thysanodes (Krutsch and Hep- penstall 1955), Idionycteris phyllotis (Black 1970), Sperrnopfnlus richardsonii (Hansen 1953), Perognathus penicillatus (Stock 1970), Perognathus fasciatus (Hayward and Kill- pack 1956), and Odocoileus virginianus (Miller and Kellogg 1955). Two other kinds, Eutamias umbrinus (White 1953) and Tho- mojuys idahoensis (Thaeler 1972) have been accorded specific status. This brings to 126 the total number of species of recent mam- mals known from Utah. Most of these spe- cies were included by Sparks (1974) in a re- cent popularized checklist of the mammals of the state. Nomenclature in this paper fol- lows Jones et al. (1975) and Hennings and Hoffmann (in press). Work on mammals of Canyonlands Na- tional Park, of which the present study is a by-product, has been supported by the So- ciety of the Sigma Xi, the Penrose Fund of the American Philosophical Society, the Council on Research and Creative Work of the University of Colorado, and the Can- yonlands Natural History Association. I thank Dr. G. L. Kirkland, Jr., for helpful comments on an earlier draft of this paper. The paper is dedicated to the memory of Dec. 1977 ARMSTRONG: UTAH MAMMAL DISTRIBUTION 459 Stephen D. Durrant, late dean of mamnia- logists of the Intermountain West. Results and Discussion Distributional limits of mammalian spe- cies form complex but recurring patterns. These patterns may be described at various levels of resolution and by various means. In a local area, for example, one might be interested in the pattern of habitat require- ments of a single species' population, in re- curring communities of organisms, or in dis- tribution along altitudinal or other gradients. On a broader scale, concern might be with distribution of species through ecological community types, with ecogeographic units (like the biotic pro- vinces of Dice, 1943) that summarize re- gional ecological pattern, or with the shapes of species' ranges (areographic analysis). Emphasis here is on ecogeographic and areographic analysis, and the relationships between them. Ecogeographic description summarizes broad environmental patterns; areographic analysis may provide historical clues to the evolution of regional faunas (Armstrong 1972a). Ecogeographic Considerations.— J. A. Al- len (1892) pioneered ecogeographic studies of North American mammals, and Kendeigh (1954) reviewed subsequent work. Dice (1943) developed the concept of the Biotic Province to describe coherent units of re- gional landscape. Hagmeier and Stults (1964) and Hagmeier (1966) derived "mam- mal provinces" in North America by nu- merical methods, based on range maps in Hall and Kelson (1959). Hagmeier (1966) in- cluded parts of Utah in six mammal pro- vinces, arranged in his hierarchy of eco- geographic units as follows: I. Coniferan Subregion A. Mountain Superprovince 1. Coloradan Province II. Sonoran Subregion A. Navajo Superprovince 1. Navajonian Province 2. Uintian Province B. Mapimi Superprovince 1. Kaibabian Province C. Columbia Superprovince 1. Columbian Province 2. Artemisian Province The fact that the six provinces in Utah were arranged in four different super- provinces underscores the patent faunal het- erogeneity in Utah detailed below. Durrant (1952) outlined "faunal sub- divisions" of Utah as follows (also see Fig. 1): I. Middle Rocky Mountain Faunal Area A. Wasatch Mountain Province B. Uinta Mountain Province C. High Plateau Province 1. Northern High Plateau Subcenter 2. Southern High Plateau Subcenter II. Southern Rocky Mountain Faunal Area A. Coloradan Province 1. La Sal Mountain Subcenter 2. Abajo Mountain Subcenter III. Colorado Plateau Faunal Area A. Canyonlands Province 1. Kaiparowits Subcenter 2. San Rafael Subcenter 3. Grand Valley Subcenter 4. San Juan Subcenter 5. Painted Desert Subcenter a. Monument Valley District b. Navajo Mountain District B. Virgin River Valley Province 1. Beaverdam Wash Subcenter 2. St. George Subcenter ^y^ 1 M 0 20 40 to MilM 6A IT^^^ 5 x^fi ('"/ J i ^^^^^^ ^ 3AI / 3A4 b!| 3B2 [ Fig. 1. Mammalian faunal subdivisions of Utah (after Durrant, 1952). For key to names of units, see text, p. 461. 460 GREAT BASIN NATURALIST Vol. 37, No. 4 IV. Columbia Plateau Faunal Area V. Great Basin Faunal Area VI. Northern Great Plains Faunal Area A. Bridger Basin Province B. Uinta Basin Province 1. Duchesne Subcenter 2. Uintah Subcenter Durrant (1952) recognized these areas as distinctive because (1) certain species were restricted there and (2) because they acted as centers of differentiation for subspecies (p. 480). Boundaries of faunal areas were based on physiography. Although based mostly on distribution of mammals, Cur- rant's faunal areas are analogues of L. R. Dice's "biotic province," an ecogeographic imit that "... covers a considerable and continuous geographic area and is charac- terized by the occurrence of one or more ecologic associations that differ, at least in proportional area covered, from the associ- ations in adjacent provinces" (Dice 1943: 3). Dice's units are based on distributional pat- terns in the biota as a whole, Durrant's on patterns in distribution and differentiation of mammals. Boundaries between both faun- al areas and biotic provinces are zones of relatively rapid biotic change, zones in which limits of species tend to be concen- trated (Armstrong 1972a). This fact was used in order to evaluate boundaries be- tween faunal areas proposed by Durrant (1952: 480). A number of quantitative meth- ods have been used in recent years to deter- mine boundaries of faunal units, but faunal change in Utah is so rapid that useful units can be identified by inspection. Limits of ranges of 92 mammalian species were su- perimposed (Fig. 2). The concurrence be- tween zones of rapid faunal change and ecogeographic boundaries is apparent from comparison of Figures 1 and 2. Each of the boundaries indicated in Durrant's map is marked by a concentration of limits in Fig- ure 2. In particular, note the dense cluster of limits which outlines the central high- lands and separates the Colorado Plateau on the east from the Great Basin in the west. In the northwestern corner of the state the Raft River Mountains are outlined clearly, as are the La Sal and Abajo Moimtains in the southeast. A concentration of limits in the northeast suggests the distinctiveness of mammals of the Bridger Basin. In the west- central part of the state, the Deep Creek Mountains are highlighted by a concentra- tion of limits. Were sufficient data available on mammals of this and other isolated mountain ranges of the Great Basin— com- parable to those provided by Lee (1960) for the highlands of southeastern Utah— sub- divisions of the Great Basin Faunal Area might be recognizable. It should be reemphasized that this evaluation of the boundaries of Durrant's faunal units is based on species' limits, whereas Durrant used two criteria, species' occurrence and differ- entiation of subspecies; this difference in technique has little bearing on the faunal imits recognized. Relationships among various faunal divi- sions of Utah were analyzed by a method used previously for Coloradan mammals (Armstrong 1972a). Occurrence of 109 mammalian species (92 species for which limits in Utah can be drawn with reason- able confidence and 17 species that range virtually statewide) was tabulated in each of Fig. 2. Superimposed distributional limits in Utah of 92 mammalian species. Dec. 1977 ARMSTRONG: UTAH MAMMAL DISTRIBUTION 461 the 19 faunal units recognized by Durrant. Faunal resemblance was calculated by the formula 2C / N^ + Ng, where C is the number of species held in common between two units and N^ and No are total numbers of species in the two units. Resemblance values (which ranged from 0.438 to 0.964) were clustered by the unweighted pair- group method (Sokal and Sneath 1963: 309). Figure 3 is the result. In large measure, the dendrogram sub- stantiates the hierarchy of units suggested by Durrant (1952: 481-see p. 00 above), de- spite wide differences in technique and re- cent additions to knowledge of Utah mam- mals. Units of the Colorado Plateau form a tight cluster, with faunas of the Canyon- lands and the Virgin River Valley separa- ting into distinct subclusters, and units with- in the Canyonlands forming further subclusters east and west of the Colorado River. In the central mountainous core, two subclusters are evident. The Wasatch and Uinta mountains are more closely related than all but one other pair of faunal units, and the Northern High Plateaus and South- ern High Plateaus form a distinct, albeit weaker, subcluster. The isolated La Sal and Abajo mountains form a distinct unit, allied (on average) more closely with the Colo- rado Plateau than with the central high- lands of Utah. Their nearest contact with a well-developed highland fauna is via the high plateaus of western Colorado, not the Middle Rocky Mountains in Utah. The Duchesne and Uintah units of the Uinta Ba- sin form a tight cluster and are distinct from faunal units of the Colorado Plateau south of the Tavaputs Plateaus, but they are more closely related to the Bridger Basin. The Great Basin and the Columbia Plateau (Raft River Mountains) tend to be faunally distinct from each other and from the rest of the state. Comparison of Durrant's faunal units with the diagram in Figure 3 suggests that levels of the hierarchy (faunal area, pro- vince, subcenter, district) are not used quite consistently, at least at the specific level of analysis employed here. In particular, faunas of the Wasatch and Uinta Mountains are more closely related than most other pairs of units, yet they are distinguished as provinces. Recent studies (e.g., Durrant and Dean 1959) suggest that the Navajo Moun- tain and Monument Valley units are more distinctive than Durrant (1952) supposed. Indeed, Monument Valley seems to be more closely related to the San Juan area (which lies north of the San Juan River) than to the Navajo Mountain unit. Based on relation- ships in the dendrogram (Fig. 3), I suggest the following rough guidelines for levels of the hierarchy of ecogeographic imits: aver- age resemblance between faunal areas, less than 0.800; average resemblance between provinces within faunal areas, 0.800 to about 0.900; average resemblance between subcenters within provinces, greater than about 0.900. Using these criteria, the fol- lowing slightly revised list of faunal sub- divisions in Utah is suggested: I. Central Highlands Faunal Area A. Middle Rocky Mountains Province 1. Wasatch Mountains Subcenter 2. Uinta Mountains Subcenter B. High Plateaus Province 1. Northern High Plateaus Subcenter 2. Southern High Plateaus Subcenter T .800 900 MEAN FAUNAL RESEMBLANCE Fig. 3. Mean resemblance among mammalian faunal subdivisions of Utah. (For explanation of index, see text.) 462 GREAT BASIN NATURALIST Vol. 37, No. 4 II. Southern Rocky Mountains Faunal Area A. La Sal Mountains Subcenter B. Abajo Mountains Subcenter III. Colorado Plateau Faunal Area A. Canyonlands Province 1. Kaiparowits Subcenter 2. San Rafael Subcenter 3. Grand Valley Subcenter 4. San Juan Subcenter 5. Monument Valley Subcenter 6. Navajo Mountain Subcenter Virgin River Valley Province 1. Beaverdam Wash Subcenter 2. St. George Subcenter IV. Columbia Plateau Faunal Area V. Great Basin Faunal Area VI. Wyoming Basin Faunal Area A. Bridger Basin Province B. Uinta Basin Province 1. Uinta Subcenter 2. Duchesne Subcenter Changes in names of two faunal areas (Middle Rocky Mountains to Central High- lands, Northern Great Plains to Wyoming Basin) follow Durrant's lead in naming faun- al units to correspond with physiographic units (here following Fenneman, 1931). Areographic Patterns.— Having considered distributional patterns of mammals in Utah and arrived at a set of ecogeographic faunal areas, let us turn to broader, continental patterns of distribution. The pattern of spe- cies' ranges, irrespective of extant ecologic pattern, may suggest historic affinities of the fauna. Udvardy (1969: 282) noted that the constituent species of a faunal list "... fall into groups with respect to the shapes of their geographic areas." These groups of species may be called "faunal elements." Polunin (1960: 212) suggested sorting out elements in a local flora as follows: (1) re- move exotic and occasional species; (2) re- move widespread species; (3) remove en- demic species; and (4) sort out the remainder according to the shapes of their ranges. Such a procedure was used here, re- sulting in nine distinctive, areographic faun- al elements and a group of widespread spe- cies. The implications of these areographic faunal elements are complex. The area oc- cupied summarizes the interaction of spe- cies' limits of tolerance with the mosaic of regional landscapes; both are subject to change over time. Hulten (1937) argued strongly that areographic analysis leads to historical insights in plant geography, pro- mulgating the concept of "progressive equi- formal areas, " the common centers of which point to areas of origin (or refugia from en- vironmental purturbations). Broad areogra- phic studies of animals (e.g., Dunn 1931, for North American amphibians and reptiles, Mayr 1946, for birds, and Simpson 1947, and Burt 1958, for mammals) have used ranges— usually of higher taxonomic cate- gories—as a basis for historical conclusions. A few studies have sought to use areogra- phic patterns of species as clues to local faunal history. For example. Miller (1951: 582) recognized four avifaunal ele- ments in California, ". . . on the basis of strong or repeated associations of species which have similar centers of distribution and probably also similar areas of origin." Armstrong (1972a: 333) discussed areogra- phic patterns of Coloradan mammals, draw- ing tentative historical conclusions. Clearly, faunal elements may have a degree of his- toric integrity, reflecting centers of origin and dispersal, although recent ecologic his- tory may have distorted older patterns. De- spite problems in interpretation, the at- tempt to sort out areographic faunal elements is important. In the absence of a fossil record, such an exercise may provide the only clues to the development of the extant fauna. Seventeen species (14 percent of the Uta- han mammalian fauna) have ranges center- ing on the Middle Rocky Mountains (see Fig. 4A) and are called Cordilleran species. These are: Sorex Dograrw— Vagrant Shrew Sorex monticoliis— Dusky Shrew Sorex nanus— Dwarf Shrew Ochotona princeps—Pika Eutamias amocnus— Yellow-pine Chipmunk Eutamiwi umfortnws— Uinta Chipmunk Mannota /7at)jt)entrts— Yellow-bellied Marmot Spermophilus armatus—V inta Ground Squirrel Spermophihis /atera/is— Golden-mantled Ground Squirrel Tliomomys talpoides— Northern Pocket Gopher Neotoma cinerea— Bushy-tailed Woodrat Dec. 1977 ARMSTRONG: UTAH MAMMAL DISTRIBUTION 463 Microtus montaniw— Montane Vole Microtus fongtcaudus— Long-tailed Vole Arvicola ric/iardsoni— Water Vole Zapus princeps— Western Jumping Mouse Ovis canadensis— Bighorn Sheep Bison fotson— Bison j CHIHUAHUAN (30) Fig. 4 A-D. Superimposed continental distributions of species of four areographic faunal elements. 464 GREAT BASIN NATURALIST Vol. 37, No. 4 Ranges of most of these species extend from the Canadian Rockies southward, often in an increasingly insular pattern, to Ari- zona and New Mexico. They are restricted at their southern limits to mountainous country. Habitat of many of the species is mountain meadows, streamsides, and forest- edge situations. Two of the species (Sorex obscurus and Spemiophihis lateralis) have isolated montane populations in Mexico. However, four other species (Sorex vagrans, Eutamias amoenus, Spemwphilus annatus, and Arvicola richardsoni) do not extend far- ther south than the Middle Rocky Moun- tains, and they do not reach Colorado. The American bison of Utah were members of the subspecies Bison bison athabascae (see Hall and Kelson 1959: 1025) and hence are included as Cordilleran rather than wide- spread species. The following 15 species of Utahan mam- mals (12 percent of the fauna) share a dis- tributional pattern that may be called Boreo-Cordilleran: Sorex cinereus— Masked Shrew Sorex palustris— 'Water Shrew Lepus americaniis—Snowshoe Hare Eutamias minimus— Least Chipmunk Tamiasciurus hudsonicus—Red Squirrel, or Chick- aree Glaucomijs safcrinus— Northern Flying Squirrel Clethrionornys gapperi— Southern Red-backed Vole Phenacomys infermedius— Heather Vole Microtus pennsylvanicus— Meadow Vole Maries ainericana— Marten Maries pennanfi— Fisher Mustela ^rminea— Ermine Gulo gnfo— Wolverine Felis lynx— Lynx Alces akes— Moose These species are distributed in mountain- ous parts of western North America and also eastward across the continent, mostly in forested areas (Fig. 4B). In an areographic sense, these species contrast markedly with those of the Cordilleran Faunal Element. Unlike Cordilleran species, many Boreo- Cordilleran mammals range throughout for- ested parts of Alaska and several species (in- cluding Sorex cinereus, Mustela enninea, Gulo gulo, Felis lynx, Alces aloes) occur on both sides of Bering Strait. In addition. there is a tendency for Boreo-Cordilleran mammals to be associated with heavy for- est. There is some suggestion that Eutamias minimus, perhaps the most euryecious mammal in this faunal element, may in- clude more than a single species (Sutton and Nadler, 1969). Thirty species (24 percent of the fauna) comprise the Chihuahuan Faunal Element: Notiosorex crawfordi— Desert Shrew Myotis ca/i/ornicus— California Myotis Myotis t/umanensis— Yuma Myotis Myotis velifer— Cave Myotis Myotis gyofis— Long-eared Myotis Myotis thysanodes— Fringed Myotis Pipistrellus hesperus— Western Pipistrelle Plecoius iownsendii—T awnsend' s Big-eared Bat Idionycteris phyllotis—AWens Big-eared Bat Antrozous pa//tdui— Pallid Bat Tadarida forasi/iensis— Brazilian Free-tailed Bat Tadarida macrotis— Big Free-tailed Bat Sylvihgus audubonii— Desert Cottontail Lepus ca/i/ornicus— Black-tailed Jackrabbit Eutamias dorsalis—Clifi Chipmunk Spermophilus spilosoma—Spotted Ground Squirrel Spermophilus variegatus— Rock Squirrel Perognathus flavus—Si\ky Pocket Mouse Dipodomys ordii—Ord's Kangaroo Rat Reithrodontomys megalotis— Western Harvest Mouse Peromyscus fooy/ii— Brush Mouse Peromyscus fruei— Pinyon Mouse Peromyscus difficilis— Rock Mouse Neotoma a/foigwia— White-throated Woodrat Neotoma mexicana— Mexican Woodrat Microtus ?nexicanus— Mexican Vole Vtdpes macrotis— Kit Fox Urocyon cinereoargenteus— Gray Fox Bassariscus astutus— Ringtail Spilogale graci/is— Western Spotted Skunk These species mostly occur in arid to semiarid grasslands or in rocky, broken brushlands or woodlands. They share a cen- ter of coincidence in the basin and range region of Chihuahua and Coahuila, Mexico, and Trans-Pecos Texas (Fig. 4C). Of this faunal element, four species {Plecotus towns- endii, Tadarida brasiliensis, Reithrodontomys megalotis, and Urocyon cinereoargenteus) range east of the Mississippi River. How- ever, their identification in the southwest with this faunal element is clear. The two species of free-tailed bats, Tadarida brasi- liensis and T. macrotis, occur widely in South America, as does Urocyon cine- Dec. 1977 ARMSTRONG: UTAH MAMMAL DISTRIBUTION 465 reoargenteus. These might have been segre- gated as a Neotropical Faunal Element, in- asmuch as only four of the remaining Chihuahuan species (Mijotis velifer, Per- omijsctis boylii, Neotoma mexicana, Spilo- gale gracilis) range farther south than the Isthmus of Tehuantepec. Note that 12 of the Chihuahuan species (40 percent) are bats. Despite their capacity for flight these species are of restricted distribution, present patterns perhaps reflecting the historical in- tegrity of this faunal element. A Nevadan Faunal Element (Fig. 4D), comprised of the following 14 species, with a center of coincidence in Nevada, contrib- utes 11 percent of the Utahan fauna: Sorex merriami— Merriam's Shrew Myotis Doians— Long-legged Myotis Euderma niaculatum—Spotted Bat Sylvilagus idahoensis— Pygmy Rabbit Sylvilagus nutta//ii— Nuttall's Cottontail Lepiis totfnsendti— White-tailed Jackrabbit Spermophilus townsendii—T ownsend's Ground Squirrel Spermophilus foeWingi— Belding's Ground Squirrel Perognathus parvus— Great Basin Pocket Mouse Microdipodops inegacephalus—DarV. Kangaroo Mouse Dipodomys microti— Chisel-toothed Kangaroo Rat Onychomys leucogaster— Northern Grasshopper Mouse Lagurus curtatus— Sagebrush Vole Antilocapra amencana— Pronghorn This is a complex distributional element. At a finer level of analysis it might be sub- divided profitably. Some species are re- stricted to arid interior basins of the west- ern United States (e.g., Spermophilus toivnsendii, Microdipodops megacephaliis, Dipodomys microps), whereas others inhabit sagebmsh steppe (e.g., Sorex merriami, Syl- vilagus idahoensis, Sylvilagus nuttallii. La- gurus curtatus). Lepus townsendii, Ony- chomys leucogaster, and Antilocapra americana have an additional center of co- occurrence on the central Great Plains and might be considered a part of a Camp- estrian Faunal Element (Armstrong 1972a: 356), although this designation would not be particularly meaningful with respect to these species as they occur in Utah, in- asmuch as communication with the Great Plains is indirect. Twelve species (10 percent of the fauna) have continental distributions that might be called "Yuman," for their center of coinci- dence is in the Mojave Desert and along the Lower Colorado River in California, Arizona, Sonora, and Baja California (Fig. 4F). These species are: AmmospermophUus Zeucurivs— White-tailed Ante- lope Squirrel Tliomomys bottae—Botta's Pocket Gopher Perognathus longimembris— Little Pocket Mouse Perognathus /omiosus— Long-tailed Pocket Mouse Perognathus penicillatus— Desert Pocket Mouse Perognathus intermedius— Rock Pocket Mouse Dipodomys deserti— Desert Kangaroo Rat Dipodomys merriami— Merriam's Kangaroo Rat Peromyscus eremicus— Cactus Mouse Peromyscus crinitus— Canyon Mouse Onychomys torridus—Southern Grasshopper Mouse Neotoma lepida— Desert Woodrat In the vicinity of the center of coincidence, AmmospermophUus leucurus and Per- ognathus fonnosus only occur west of the Colorado River, and Perognathus inter- medius is known only east of the river. Yu- man mammals generally do not range south- ward in Mexico as far as Chihuahuan species, with only five reaching Jalisco, Guanajuato, and San Luis Potosi. On the north, most species reach no farther than southeastern Oregon and southwestern Idaho, and several reach limits in Nevada and Utah. Note that half the species in this faunal element are heteromyid rodents, whereas the predominant rodents in the Chihuahuan element are cricetids. This fact underscores the historical integrity of these faunal elements. Five species, comprising 4 percent of the Utahan mammalian fauna, constitute an Arizonan Faunal Element (Fig. 4F). These are: Eutamias quadrivittatus— Colorado Chipmunk Cynomys gimnisoni— Gunnison's Prairie Dog Sciurus aberti—Ahert's Squirrel Perognathus apache— Apache Pocket Mouse Neotmna stephensi— Stephens' Woodrat These are species of the Four Corners area (although Neotoma stephensi is as yet un- known from Colorado, Armstrong 1972a: 312). In Utah, Arizonan species generally 466 GREAT BASIN NATURALIST Vol. 37, No. 4 are restricted to the southeastern part Three mammals of the Great Plains of the state; all but E. quadrivittatus occur (Spemiophilus tridecemlineatiis, Perognathus only east of the Colorado and Green rivers. fasciatus, Mustela nigripes) have limited CAMPESTRIAN (3) \ -a Fig. 4 E-H. Superimposed continental distributions of species of four areograhic faunal elements. Dec. 1977 ARMSTRONG: UTAH MAMMAL DISTRIBUTION 467 ranges in Utah and constitute a Camp- estrian Faunal Element (Fig. 4G). All occur in grasslands of the eastern part of the state. Another three species (Cynomys leucurus, Thomomys idahoensis, and Spennophilus richardsonii) share a center of coincidence in the Bridger Basin of southwestern Wyoming (Fig. 4H) and are herein called the Bridgeran Faunal Element. Armstrong (1972a: 354) placed S. richardsonii as a member of the Cordilleran Faunal Element, but its range is somewhat discordant with that group of species. Recent studies (Nad- ler et al. 1971) suggest that the population in Wyoming, Utah, and Colorado, known as S. r. elegans, is in fact specifically distinct from S. richardsonii. The range of elegans fits closely with the Bridgeran Faunal Ele- ment. A single species of the mammalian fauna of Utah is obviously endemic there, Cy- nomys parvidens, the Utah Prairie Dog. Ac- cording to Pizzimenti and Collier (1973), C. parvidens is more closely related to C. leuc- urus (a species of the Bridgeran Faunal Ele- ment) than to other prairie dogs; Collier and Spillett (1975) concluded that the range of the C. parvidens once covered large por- tions of the Great Basin. Of mammals of Utah, the following 26 kinds (21 percent of the state's fauna) are sufficiently widespread (Fig. 41) that they cannot be identified with any one areogra- phic faunal element: Myotis lucifugus— Little Brown Bat Myotis feifoti— Small-footed Bat Lasionycteris nocfiuagans— Silver-haired Bat Eptesicus fuscus— Big Brown Bat Lasiurus cinereus— Hoary Bat Lasiurus borealis— Red Bat Castor canadensis— Beaver Peromyscus manicitlatits— Deer Mouse Ondatra zibethicus—Muskrat Erethizon dorsatum— Porcupine Canis latrans— Coyote Canis lupus— Gray Wolf Vulpes vulpes—Bed Fox Ursus americanus—B\acV Bear Ursus arctos— Grizzly Bear Procyon fotor— Raccoon Mustela /renata— Long-tailed Weasel Mustela vison—MinV Mephitis mephitis—Striped Skunk Taxidea taxus— Badger Lutra canadensis— River Otter Felis concofor— Mountain Lion Felis rufus— Bobcat Cervus elaphus— Wapiti, or American Elk Odocoileus /lemionus— Mule Deer Odocoileus DirgJnianuA— White-tailed Deer Many of these species are rather large in size and many have broad habitat toler- ances. It is perhaps noteworthy that half of these eurychores are members of a single order, Camivora; carnivores are at least one step removed from direct dependence on the vegetation for food and generally are less narrowly restricted to particular habi- tats than are herbivores. Thirteen of the 23 carnivores known to occur in Utah are widespread species. In addition, three of four Utahn cervids are widespread on a continental scale. Three highly specialized aquatic species, the beaver, the muskrat, and the mink, appear on the list. This is hardly surprising, since aquatic habitats pro- vide corridors of uniform habitat for dis- persal through otherwise highly distinctive regions. Fewer than one-third of Utah's bats are widespread species; nearly 60 percent Fig. 4 \. Superimposed continental distributions of 26 widespread species. 468 GREAT BASIN NATURALIST Vol. 37, No. 4 are Chihauhuan kinds. Were distribution of bats better known, they might provide quite useful data for zoogeographers, contrary to the conventional wisdom. The least broadly distributed species on the above list are the small-footed myotis {Myotis leibii) and the grizzly bear {Ursiis arctos). The range of the former species is similar to that of those labelled "Nevadan," but it extends also across the Great Plains to the Ohio Valley and the East Coast. The former range of the grizzly bear extended eastward across the Northern Great Plains and Central Great Plains; otherwise the range is that of a Cordilleran species. Figure 5 indicates cumulative percentage composition of the mammalian faunas of 19 ecogeographic faunal units by species of the 10 faunal elements identified by areogra- phic analysis. Also indicated, for com- parison, is the composition of the fauna of Utah as a whole, based on a total of 126 species. Only 109 species were tabulated in the faunal units, because of inadequate dis- tributional data on 17 species (see Methods, above). Those species too poorly known to map represent five faunal elements: Cordil- leran (three species), Boreo-cordilleran (one), Chihuahuan (four), Nevadan (three), and widespread (six). Thus, poorly known spe- cies are sufficiently well distributed across the major faunal elements that they do not bias the remarks that follow. Note first in Figure 5 the consistent im- portance of widespread species at about 35 percent (28 to 38) through each of the faun- al elements. It is the differential occurrence of species of well-defined areographic ele- ments that makes the faunal units dis- tinctive. Cordilleran species are most impor- tant in faunal units of the central mountain core of the state, somewhat less important in the La Sal Mountains and the Raft River Mountains of the Columbia Plateau. In oth- er faunal units, their contribution falls to 10 percent or less. Boreo-cordilleran species are even more narrowly restricted to moun- tainous areas, although one species, Eii- tamias minimus, is sufficiently euryecious that it occurs in most faunal areas. "Boreal" mammals of Utah discussed by Brown (in press) mostly are Cordilleran and Boreo- cordilleran species as defined here. The Chihuahuan Faunal Element is the largest distinctive areographic element in Utah, constituting about one quarter of the fauna. Given this prominence statewide, the element is under-represented in the central highlands faunal areas and on the Columbia Plateau. Over most of the Colorado Plateau south of the Tavaputs Plateaus, the Chihua- huan element contributes some 40 percent of the fauna. North of the Tavaputs Plateaus, in the southeastern mountains, and in the Great Basin, the importance of Chi- huahuan species is diminished. Yuman mam- mals are most important on the southern Colorado Plateau, especially in the Virgin River Valley. They are absent from the Middle Rocky Mountain Province and from the Northern High Plateaus. Nevadan spe- cies, on the other hand, are most important in the Great Basin, with modest representa- tion over most of the rest of the state (ex- cept on the Colorado Plateau, where they tend to be imder-represented). The minor faunal elements tend to show rather narrow distribution across the state. Campestrian species occur in eastern Utah. Arizonan spe- cies occur mostly in the Canyonlands Pro- vince of the Colorado Plateau and in the Uinta Basin. Bridgeran species occur only in northeastern and east-central parts of the state. Having considered the composition of the faunas of the ecogeographic imits, it might be useful (if only as a check on the integri- ty of the faunal elements) to look briefly at the extent to which members of the 10 faunal elements occupy the various units (Table 1). Most of the units include 60 to 70 percent of the widespread species, with highest percentages occurring in mountain- ous areas. As might be expected, the units of the Central Highlands Faunal Area ac- commodate most members of the Cordille- ran and Boreo-cordilleran elements, with occurrence attenuating southward. The Col- orado Plateau includes the ranges of the highest proportion of Chihuahuan species. Yuman species, by contrast, occur strongly only in the Virgin River Valley, and Neva- Dec. 1977 ARMSTRONG: UTAH MAMMAL DISTRIBUTION 469 dan mammals are well represented only in Arizonan species east of the Colorado River the Great Basin. The minor faunal elements (and south of its confluence with the Green also are strongly represented only in local River), the Bridgeran species in the Bridger areas— Campestrian species in the northeast. Basin. STATEWIDE TOTAL 00 CUMULATIVE PERCENTAGE Fig. 5. Cumulative percentage composition of mammalian taunas of ecogeographic faunal areas by members of ten areographic faunal elements. 470 GREAT BASIN NATURALIST Vol. 37, No. 4 Historical Implications.— One of the ma- jor goals of zoogeographic analysis is to gain some insight into the evolution of local or regional faunas. Indeed, in the absence of an adequate fossil record, extant patterns of distribution and differentiation provide the only historical data that we have to work with. Durrant (1952) looked at patterns of subspecific differentiation in fossorial ro- dents as a clue to the Pleistocene history of the Bonneville Basin. Lee (1960) studied the distribution and differentiation of montane mammals of southeastern Utah and drew in- ferences about past faunal movements. Findley (1969) presented a strong argument on historical implications of such patterns in New Mexico and adjacent areas. Brown (1971, in press) analyzed distribution on sev- eral mountain ranges in Utah of mammals typical of woodland (and higher elevation) biotic communities, arguing convincingly that extant distributional patterns stem from local extinction of populations that reached the montane islands during the late Pleisto- cene. Armstrong (1972a) relied on evidence at both the specific and subspecific level to reach tentative historical conclusions about Coloradan mammals, and hypothesized ac- cess to the state by several faunal com- ponents under a diversity of environmental conditions which prevailed at various times in the past. Extant zoogeographic patterns in western North America are a post-Pleistocene devel- opment. The Pleistocene Epoch was marked by pronounced climatic fluctuations. Warm, dry periods were interrupted by cool, moist glacio-pluvial intervals. Evidences of zoo- geographic patterns in the earlier Pleisto- cene were obliterated by the last major gla- cial interval, the subsequent Hypsithermal Interval (Deevey and Flint 1957), and the development of the existing climatic regime (Armstrong 1972a). Extant patterns may provide clues to these most recent events. In the absence of a coherent or extensive fossil record, distributional patterns of re- cent species are the only clues available to Table 1. Percentage of members of areographic faunal elements occurring in ecogeographic faunal units. 72 5 P 1 Z S < 69 82 93 23 29 77 71 80 17 - 29 - - - 7.3 71 79 37 - 36 - - - 65 65 47 37 8 36 33 _ 33 69 47 40 43 25 21 — — — 62 24 27 43 25 29 _ 40 _ 6.5 6 — 63 50 21 — 20 33 65 6 — 60 33 21 _ 20 33 65 6 — 63 33 14 - 20 33 65 6 7 73 .33 14 33 60 — 65 6 7 73 33 14 - 60 — 62 6 7 77 42 14 — 80 _ 62 6 7 67 92 29 — — — 58 6 7 60 75 29 33 — _ 69 53 27 37 25 36 — — — 58 18 7 57 50 71 _ _ — 62 24 7 25 17 36 _ _ 100 65 24 7 50 25 36 67 20 33 62 29 7 57 33 29 67 40 .33 Wasatch Mountains Uinta Mountains Northern High Plateaus Southern High Plateaus La Sal Mountains Abajo Mountains Kaiparowits San Rafael Grand Valley San Juan Monument Valley Navajo Beaverdam St. George Columbia Plateau Great Basin Bridger Duchesne Uintah 100 100 Dec. 1977 ARMSTRONG: UTAH MAMMAL DISTRIBUTION 471 US. This is the case in Utah. Conspicuously lacking from the abundant literature on Pleistocene environments of Utah is any re- cord of a mammalian local fauna. Even fragmentary fossils are few. When a nearly adequate fossil local fauna does become available (the Hogup Cave deposits from Box Elder County described by Durrant, 1970, and dated at 8500 years B.P.) the mammals that are present are those that would be expected in the vicinity of the cave today. For purposes of discussion of Pleistocene conditions, Utah is conveniently divisible into three broad sections: the Great Basin, the central mountainous core, and the Colo- rado Plateau. The Great Basin records evi- dence of a series of pluvial periods sepa- rated by intervals of dessication (for details, see Morrison 1965). Lake levels fluctuated in synchrony with alpine glaciers in the higher mountains of central and northern Utah. This glaciation strongly influenced the modern landscape, lowering regional snow- lines some 4000 ft, and producing summer temperatures perhaps 16 F cooler than today (Richmond 1965). On the Colorado Plateau, glacial intervals were marked by erosion, and warmer, drier periods produced sedimentary deposition (Kottlowski, Cooley, and Ruhe 1965). These events and condi- tions set an environmental baseline against which present distributional patterns of mammals have developed. During the last glacio-pluvial stage, conditions probably prevailed that were beyond the tolerance of many species in the fauna today. Previous patterns of distribution would have been obscured. With a depression of zonal biotic communities, forested situations that are highly disjunct today would have been more nearly continuous, and semidesert and desert community-types would have seen concommitant restriction and fragmentation. The present-day fauna of the highlands of Utah provides better evidence of past envi- ronmental change than do faunas of the Colorado Plateau and the Great Basin. Today, a number of the state's mountain ranges and high plateaus appear as "islands" in a "sea" of nonmontane habitats. Commu- nication among them by nonvolant mam- mals restricted to forested habitats is prob- ably impossible. Nonetheless, some of the highlands have reasonably complete high- land faunas (Brown 1971, in press). Altitu- dinal depression of zonal biotic communities by 2000 to 4000 ft would unite many of these areas with corridors of suitable habi- tat. The La Sal Mountains would have been connected with the Uncompahgre Plateau and the San Juan Mountains (and nearby ranges), and the Abajo Mountains probably would have been joined with the La Sals by more-or-less continuous forest or woodland corridors. Navajo Mountain, south of the San Juan River, and the Henry Mountains probably have been isolated (or nearly so) throughout Pleistocene times (although Lee, 1960, noted some affinity of mammals of the Henrys with those of the Aquarius Plateau). The distribution of forest-dwelling mammals along the mountains of central Utah suggests that Pleistocene conditions there provided a continuous corridor for movement. Isolated ranges of the Great Ba- sin generally support depauperate highland faunas (Brown 1971, in press), although the definitive study of mammals of the minor ranges of southwestern Utah (House Range, Wah Wah Mountains, etc.) remains to be made. The fact that faunas of the more isolated ranges on either side of the central high- lands corridor have variously depauperate faunas probably reflects local extinction rather than selective or chance dispersal across barriers. The fact that such extinction has been more pronounced on the smaller uplifts may reflect the greater susceptibility of small populations to extinction (Brown 1971, in press). Such small populations would have been constricted still further by the climatic changes of the Hypsithermal Interval (warmer and effectively or abso- lutely drier than at present) which would have resulted in upward movement of zonal biotic commimities with consequent restric- tion of the higher zones. At lower elevations in Utah, extant distri- butional patterns strongly reflect present- day physiography, suggesting the efficacy of 472 GREAT BASIN NATURALIST Vol. 37, No. 4 existing boundaries throughout the period of development of the fauna. Ranges of many Chihuahuan and Yuman species, for ex- ample, are limited on the Colorado Plateau by the major rivers and their canyons, de- spite the fact that seemingly suitable habitat often exists on the other side of the barrier. On a north-south axis, the Wasatch Moun- tains and the high plateaus today form an effective barrier to communication between the Great Basin and the Colorado Plateau. This barrier seems to have been generally effective throughout the period of evolution of the fauna. If the Hypsithermal Interval had significantly reduced its effectiveness, one would expect to find several Nevadan species on the Colorado Plateau. However, only the most euryecious species are found on both sides of the mountains. The Uinta Movmtains also seem to have persisted as an effective barrier as faimal patterns have evolved. Bridgeran species are not found on the Colorado Plateau to any significant ex- tent, and most Chihuahuan species are lim- ited south of the Uintas. A barrier to one faunal element or spe- cies may well be a corridor for the move- ment of another. The central mountainous corridor has been discussed in this context. The river systems of the Colorado Plateau seem generally to have been ineffective as corridors. Often they are entrenched deeply; riparian habitats tend to be fragmented or nonexistent. Species adapted to the broken habitats of the canyons seem to have found the river systems more effective corridors than have species of desert grasslands (such as several Yuman species). Kelson (1951) suggested that the Colorado-Green system becomes an increas- ingly less effective zoogeographic barrier northward. East of the Colorado River and north of the San Juan, there is no strong barrier to northward faunal movement until one reaches the Book Cliffs. West of the Colorado, the Kaiparowits Plateau and the canyon of the Escalante River may con- stitute major barriers, but the mammals of this area remain poorly known except in the immediate vicinity of Lake Powell. In summary, scrutiny of broad patterns of distribution of mammalian species provides few clues to the conditions of the past that influenced the composition of the extant fauna. Ranges of montane mammals suggest the occurrence of more continuous highland biotic communities in late Pleistocene time, and the depauperate faunas of isolated ranges may reflect the efficacy of the Hyp- sithermal Interval in constricting highland communities more severely than at present. In the lowlands, distributions correspond to extant barriers. Truly relict populations of lowland, xeric-adapted species are lacking. The overall conclusion must be that barriers to distribution in the present also were bar- riers to distribution in the past. Summary Distributional patterns of the 126 mam- malian species native to Utah were ana- lyzed and discussed. Ecogeographic faunal areas, proposed by Durrant (1952), were reevaluated. These correspond strongly with physiographic subdivisions of Utah. Areogra- phic analysis indicated that several faunal elements contribute to the total mammalian assemblage within the political confines of Utah. These faunal elements contribute dif- ferentially to the several ecogeographic units, and they differ in their response to barriers and corridors for dispersal. With the exception of the montane fauna of the highlands of the state— which reflects both a cooler, moister late Pleistocene climate, and a warmer, effectively drier Hypsithermal climate— clues are lacking to suggest the va- garies of Quaternary history. Data still are inadequate to allow a satisfactory picture of some aspects of mammalian zoogeography in Utah. Areas of particular interest are the south-central part of the state (Kaiparowits Subcenter) and the isolated basins and ranges of the southern part of the Great Ba- sin Faunal Area. Literature Cited Allen, J. A. 1892. The geographical distribution of North American mammals. Bull. Amer. Mus. Nat. Hist. 4: 199-243. Anderson, S. 1955. An additional record of Mustela er- minea from Utah. J. Mamm. 36: 568. 1959. Distribution, variation, and relationships Dec. 1977 ARMSTRONG: UTAH MAMMAL DISTRIBUTION 473 of the montane vole, Microtus montanus. Univ. Kansas Publ, Mus. Nat. Hist. 9: 415-511. Armstrong, D. M. 1972a. Distribution of mammals in Colorado. Monogr., Univ. Kansas Mus. Nat. Hist. 3: X + 1-415. 1972b. Mammals of Canyonlands National Park, Utah, I. Preliminary report of in- vestigations, June-July 1972. Processed report for National Park Service, U.S. Dept. of Interior, 22pp. 1973. Preliminary observations on mammalian faunal relationships, Canyonlands National Park, Utah. J. Colorado-Wyoming Acad. Sci. 7(4): 41-42. 1974. Second record of the Mexican big-eared Basin. Anthro. Papers, Univ. Utah 48: 209-235. DuRRANT, S. D., AND R. M. Hansen. 1954. Taxonomy of the chickarees (Tamiaschirus) of Utah. J. Mamm. 35: 87-95. DuRRANT, S. D., AND M. R. Lee. 1955. Rare shrews from Utah and Wyoming. J. Mamm. 36: 560-561. 1956. A new pocket mouse from southeastern bat in Utah. Southw. Nat. 19: 114. Behle, W. H., and R. M. Hansen. 1951. Notes on the short-tailed weasel in Utah. J. Mamm. 32: 226-227. Benson, S. B. 1954. Records of the spotted bat (£u- demia inaculata) from California and Utah. J. Mamm. 35: 117. Black, H. L. 1970. Occurrence of the Mexican big- eared bat in Utah. J. Mamm. 51: 190. Brown, J. H. 1971. Mammals on mountaintops: non- equilibrium insular biogeography. Amer. Nat. 105: 467-478. In press. The theory of insular biogeography and the distribution of boreal birds and mam- mals. Great Basin Nat. Mem. 2: 209-227. Burt, W. H. 1958. The history and affinities of the re- cent land mammals of western North America. Publ, Amer. Assoc. Adv. Sci. 51: 131-154. CocKHUM, E. L. 1960. The recent mammals of Ari- zona: their taxonomy and distribution. Univ. Ari- zona Press, Tucson, viii -I- 276 pp. Collier, G. D., and J. J. Spillett. 1975. Factors in- fluencing the distribution of the Utah prairie dog, Cynoniys parvidens (Sciuridae). Southw. Nat., 20: 151-158. Davis, W. B. 1939. The recent mammals of Idaho. Caxton Printers, Caldwell, 400 pp. Dearden, L. C, and M. R. Lee. 1955. A new sage- brush vole, genus Lagurus, from Utah. J. Mamm. 36: 270-273. Deevey, E. S., Jr., and R. F. Flint. 1957. Postglacial Hypsithermal Interval. Science 125: 182-184. Dice, L. R. 1943. The Biotic Provinces of North Amer- ica. Univ. of Michigan Press, Ann Arbor, viii + 78 pp. Dunn, E. R. 1931. The herpetological fauna of the Americas. Copeia 1931: 106-119. DuRRANT, S. D. 1952. Mammals of Utah, taxonomy and distribution. Univ. Kansas Publ., Mus. Nat. Hist. 6: 1-549. 1970. Faunal remains as indicators of Neo- thermal climates at Hogup Cave. Anthro. Papers, Univ. Utah 93: 241-245. DuRRANT, S. D., AND N. K. Dean. 1959. Mammals of Glen Canyon. Anthro. Papers, Univ. Utah 40: 7.3-106. 1960. Mammals of Flaming Gorge Reservoir Utah. Proc. Biol. Soc. Washington 69: 183-186. Durrant, S. D., M. R. Lee, and R. M. Hansen. 1955. Additional records and extensions of ranges of mammals from Utah. Univ. Kansas Publ., Mus. Nat. Hist. 9: 69-80. Durrant, S. D., and J. P. Newey. 1953. Masked shrew, Sorex cinereus, in Utah. J. Mamm. ,54: 116-117. Easterla, D. a. 1965. The spotted bat in Utah. J. Mamm. 46: 665-668. 1966. Yuma myotis and fringed myotis in southern Utah. J. Mamm. 47: 350-351. Egoscue, H. J. 1961. Small mammal records from western Utah. J. Mamm. 43: 122-123. 1965. Records of shrews, voles, chipmunks, cottontails, and mountain sheep from Utah. J. Mamm. 46: 685-687. 1966. Gray fox records from western Utah. Southw. Nat. 11: 14,3-144. Egoscue, H. J., and T. J. Lewis. 1968. An albino long- tailed pocket mouse from Utah. J. Mamm. 49: 319. Fenneman, N. M. 1931. Physiography of Western United States. McGraw-Hill Book Co., New York, xiii + ,534 pp. FiNDLEY, J. S. 1969. Biogeography of southwestern bo- real and desert mammals. Misc. Publ., Univ. Kan- sas Mus. Nat. Hist. 51: 11,3-128. FiNDLEY, J. S., A. H. Harris, D. E. Wilson, and C. Jones. 1975. Mammals of New Mexico. Univ. New Mexico Press, Albuquerque, xiii + 360 pp. Genoways, H. H., and J. K. Jones, Jr. 1967. Notes on distribution and variation in the Mexican big- eared bat, Plecotus phyllotis. Southw. Nat. 12: 477-480. Hagmeier, E. S. 1966. A numerical analysis of the dis- tributional patterns of North American mammals, II. Re-evaluation of the provinces. Syst. Zool. 15: 279-299. Hagmeier, E. S., and C. D. Stults. 1964. A numerical analysis of the distributional patterns of North American mammals. Syst. Zool 13: 125-155. Hall, E. R. 1946. Mammals of Nevada. Univ. Califor- nia Press., Berkeley, xi -I- 710 pp. Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. Ronald Press Co., New York, 2 vols. Hansen, R. M. 1953. Richardson ground squirrel in Utah. J. Mamm. 34: 131-1,32. Harris, A. H. 1974. Myotis yumanensis in interior southwestern North America, with comments on Myotis lucifugus. J. Mamm. 55: 589-607. Hayward, C. L., and M. L. Killpack. 1956. Occur- rence of Perognathus fasciatus in Utah. J. Mamm. 37: 451. 474 GREAT BASIN NATURALIST Vol. 37, No. 4 1958. Distribution and variation of the Utah poulation (sic) of the Great Basin pocket mouse.Great Basin Nat. 18: 26-30. Hennings, D., and R. S. Hoffmann, in press. A re- view of the taxonomy of the Sorex vagrans spe- cies complex from western North America. Occas. Papers, Mus. Nat. Hist., Univ. Kansas. HuLTEN, E. 1937. Outhne of the history of the Arctic and boreal biota during the Quaternary period. Bokforlags Aktiebolaget Thule, Stockholm, 168 pp. Jones, J. K., Jr., D. C. Carter, and H. H. Genoways. 1975. Revised checklist of North American Mam- mals north of Mexico. Occas. Papers Mus., Texas Tech Univ. 28: 1-14. Keegan, H. L. 1953. Collections of parasitic mites from Utah. Great Basin Nat. 13: 35-42. Kelson, K. R. 1951. Speciation in rodents of the Colo- rado River drainage. Biol. Ser., Univ. Utah. 11(3): vii -I- 1-125. Kendeigh, S. C. 1954. History and evaluation of vari- ous concepts of plant and animal communities in North America. Ecology 35: 152-171. KiLLPACK, M. L. 1955. Notes on the thirteen-lined ground squirrel in Utah. J. Mamm. 36: 296-297. KOTTLOWSKI, F. E., M. E. COOLEY, AND R. V. RuHE. 1965. Quaternary geology of the Southwest. In: H. E. Wright, Jr., and D. G. Frey (eds.). Qua- ternary of the United States, pp. 287-298. Prince- ton Univ. Press, Princeton. Krutsch, p. H., and C. a. Heppenstall. 1955. Addi- tional distributional records of bats in Utah. J. Mamm. 36: 126-127. Lee, M. R. 1960. Montane mammals of southeastern Utah— with emphasis on the effect of past cli- mates upon occurrence and differentiation. Un- pubhshed doctoral dissertation, Univ. Utah, 204 pp. Lee, M. R., and S. D. Durrant. 1960. A new jumping mouse (Zapus princeps Allen) from Utah. Proc. Biol. Soc. Washington 73: 171-174. Long, C. A. 1965. The mammals of Wyoming. Univ. Kansas Publ., Mus. Nat. Hist. 14: 493-758. Long, C. A., and C. F. Long. 1964. Range extension of the gray fox, Urocyon cinereoargenteus. Southw. Nat. 9: 108-109. LowERY, E. J., AND H. J. Egoscue. 1968. The ermine in western Utah. Great Basin Nat. 28: 47. Marshall, W. H. 1940. A survey of the mammals of the islands in Great Salt Lake, Utah. J. Mamm. 21: 144-159. Mayr, E. 1946. History of the North American bird fauna. Wilson Bulk 58: .3-41. Miller, A. H. 1951. An analysis of the distribution of the birds of California. Univ. California Publ. Zool. 50: 531-644. Miller, G. S., Jr., and R. Kellogg. 1955. List of North American recent mammals. Bull. United States Nat. Mus. 205: vii -I- 1-954. Morrison, R. B. 1965. Quaternary geology of the Great Basin. In: H. E. Wright, Jr., and D. G. Frey (eds.). Quaternary of the United States, pp. 265-285. Princeton Univ. Press, Princeton. MussER, G. G., and S. D. Durrant. 1960. Notes on Myotis thysanodes in Utah. J. Mamm. 41: .393-394. Nadler, C. R., R. S. Hoffmann, and K. R. Greer. 1971. Chromosomal divergence during evolution of ground squirrel populations (Rodentia: Spermo- philus). Syst. Zool. 20: 298-305. Nicholas, D. W., H. D. Smith, and M. F. Baker. 1975. Rodent populations, biomass, and commu- nity relationships in Artemesia tridentata. Rush Valley, Utah. Great Basin Nat., 35: 191-202. Pizzimenti, J. J., and G. D. Collier. 1975. Cynamys parvidens. Mamm. Species 52: 1-3. PocHE, R. M. 1975. New record of the bat Plecotus phyllotis from Utah. Great Basin Nat. 35: 452. PocHE, R. M., and G. L. Baile. 1974. Notes on the spotted bat {Euderma maculatnm) from southwest Utah. Great Basin Nat. 34: 254-256. Poche, R. M., and G. a. Ruffner. 1975. Roosting be- havior of male Euderma maculatnm from Utah. Great Basin Nat. 35:121-122. Polunin, N. I. 1960. Introduction to plant geography. McGraw-Hill Book Co., New York, xix -\- 640 pp. Rasmussen, D. r, AND N. V. Chamberlain. 1959. A new Richardson's meadow mouse from Utah. J. Mamm. 40: 53-56. Richmond, G. M. 1965. Glaciation of the Rocky Mountains. In: H. E. Wright, Jr., and D. G. Frey (eds.). Quaternary of the United States, pp. 217-230. Princeton Univ. Press, Princeton. Shippee, E. a., and H. J. Egoscue. 1958. Additional mammal records from the Bonneville Basin, Utah. J. Mamm. 39: 275-277. Simpson, G. G. 1947. Holarctic mammalian faunas and continental relationships during the Cenozoic. Bull. Geol. Soc. America 48: 613-687. SoKAL, R. R., AND P. H. A. Sneath. 1963. Principles of numerical taxonomy. W. H. Freeman Co., San Francisco, xiv + 359 pp. Sparks, E. A. 1974. Checklist of Utah wild mammals. Publ., Utah Div. Wildl. Resources 74-3: x + 1-33. Stock, A. D. 1970. Notes on mammals of southwestern Utah. J. Mamm. 51: 429-433. Sutton, D. A., and C. F. Nadler. 1969. Chromosomes of the North American chipmunk genus Eu- tamius. J. Mamm. 50: 524-535. Thaeler, C. S., Jr. 1972. Taxonomic status of the pocket gophers, Tliornomys idahoensis and Tfio- momys pygmaeus (Rodentia, Geomyidae). J. Mamm. 53: 417-428. Udvardy, M. D. F. 1969. Dynamic zoogeography with special reference to land animals. Van Nostrand, Reinhold Co., New York, xviii -I- 445 pp. Wauer, R. H. 1965. Genus and species of shrew new for Utah. J. Mamm. 46: 496. White, J. A. 1953. Taxonomy of the chipmunks, Eu- tamias quadrivittatus and Eutamias umbrinus. Univ. Kansas Publ., Mus. Nat. Hist. 5: 563-582. Wood, S. L. 1958. A wolverine in Utah. Great Basin Nat. 18: 56. A NEW FOSSIL SPONGE FROM THE ORDOVICIAN GARDEN CITY LIMESTONE OF SOUTHEASTERN IDAHO J. Keith Rigby' and James K. Gilland' Abstract.— The new choristid demosponge, Loganiella johnsoni, is described from the upper part of the Lower Ordovician Garden City Formation of southeastern Idaho. The new sponge is a relatively thin walled, low, sub- cylindrical form. The skeletal net is composed of tracts of curved rodlike spicules, but details have been lost on all specimens because of extensive silicification. Impressions, however, suggest a close relationship to the choristid Dystactospongia Miller, 1889. Several specimens of a new sponge were collected from the Ordovician Garden City Limestone by Victor Church during a study of the Preston Quadrangle in southeastern Idaho in 194 L He was assisted by Chester O. Johnson of Gustavus Adolphus College during the study. Dr. Johnson kindly loaned us the collection of sponges he made at that time. A collection of sponges was also given to Utah State University by Mr. Church. These sponges were loaned to us by Dr. J. Stewart Williams and incorporated in this study. The Garden City Formation was named for exposures in the Bear Lake region of northern Utah and southeastern Idaho by G. B. Richardson (19L3: 407-408, 1914: 13-16) and includes the Lower Ordovician beds be- tween the underlying Cambrian St. Charles Limestone and the overlying Swan Peak Quartzite. The formation is approximately 900 feet thick in the type section, which is approximately 15 miles east of the locality where the sponges were collected. The Garden City Limestone was later restudied in detail by R. J. Ross, Jr. (1949, 1951). He described the fauna and zoned the formation utilizing the moderately abundant trilobites and other fossils and es- tablished an alphabetic zonation, parallel to that utilized by Hintze (1951, 1952) for equivalent beds of the Pogonip Group in western Utah and eastern Nevada. The fossils were collected from E V2, NW 1/4, SE 1/4 Section 30, Township 16 South, Range 42 East, in southern Franklin Coun- ty, southern Idaho (Map 1). The locality is along the west face of Crab Ridge near the ridge crest, .2 mi. north of the Utah-Idaho boundary. Crab Ridge is a prominent linear feature along the east side of the head- waters of Logan River. Johnson (1962, pers. comm.) notes that there are no major outcrops in the wooded slope along the east side of Logan River valley near where the sponges were collect- ed from talus fragments. There seems little question, however, that the material came from near the crest of Crab Ridge because of the location of the talus on the slope. The sponges are silicified and some of the better specimens were etched free of matrix by dissolving the limestone in dilute acetic acid. Many small, spherical, HindiaAike sponges(?) were recovered in the residue, along with numerous conodonts. The con- odonts were sent to Dr. Raymond L. Eth- ington of the Department of Geology, Uni- versity of Missouri for identification and zonation. Ethington and Clark (Hintze 1977) have established a conodont zonation for the Pogonip Group in western Utah and eastern Nevada and have tied this to the trilobite zones of Hintze and Ross (1950). On the basis of the conodonts the sponges came from rocks of zone J to L (Ethington, 1977, pers. comm.). Conodonts recovered from the matrix around the holotype (BYU 1090) include the following forms, identified by Ethington: Department of Geology, Brigham Young University, Provo, Utah 84602. 475 476 GREAT BASIN NATURALIST Vol. 37, No. 4 ?Drepanoistodus sp. Microzarkodina aff. M. flabelhis (Lindstrom) "Scandodus" robustus Serpagli Scolopodus gracilis Ethington and Clark ?Walliserodus aff. W. compttis (Branson & Mehl) New Genus A of Sweet, Ethington, & Barnes, 1971 New Genus B of Sweet, Ethington, & Barnes, 1971 Those conodonts recovered from matrix around the figured paratype (BYU 1091) in- ckide the following forms, identified by Ethington: ?Acodus sp. ?DrepanoistodH.s sp. Microzarkodina aff. M. flabellus (Lindstrom) Oistodus multicorrugatus Harris Protopanderodus sp. s. f. Scohpodus gracilis Ethington & Clark New Genus A of Sweet, Ethington, & Barnes, 1971 New Genus B of Sweet, Ethington, & Barnes, 1971 Ethington (1977, pers. comm.) notes that Map 1. Index map to the type locality of the new Ordovician sponge Loganiella johnsoni Rigby and Gil- land. The sponge came from upper beds of the Garden City Formation, from talus blocks, on the east side of Logan River canyon. ''Oistodus multicorrugatus and Scohpodus gracilis have rather long ranges and are of little help for stratigraphic interpretation. Fortunately some of the forms are quite di- agnostic. In particular. New Genus A, New Genus B, Microzarkodina aff. M. flabelhun, and 'Scandodus' robustus occur together in the Wah Wah and Juab Formations. New Genus A and 'Scandodus' robustus are pres- ent in the basal Whiterockian {Orthidiella Zone) in the Monitor Range, central Ne- vada, which is the equivalent of Zone L in the Ibex area of western Utah. . . . Clearly these conodonts represent a stratigraphic level near the top of the Canadian Series or near the bottom of the Whiterockian Stage. Until this series boundary between the Lower and Middle Ordovician has been de- fined more precisely, one cannot be more specific. In terms of the Hintze-Ross zonal scheme for shelly fossils they represent the general range of Zones J, K, and L." Systematic Paleontology Class Demospongia Sollas, 1875 Order Choristida Sollas, 1888 Family uncertain Loganiella, n. gen. Diagnosis.— Cylindrical to subcylindrical sponges with a flat base, a moderately thick wall, and a simple wide-open spongocoel. Relatively small canals feed radially, through the wall into the spongocoel. The skeletal net is composed of curved oxeas(?) that produce tracts with reticulate pattern that has a dominant radial structure. Discussion.— At first glance, these forms appear similar to flat-crested Receptaculites, but the skeletal structure and the canal pat- tern is so distinctly dissimilar that the forms are clearly unrelated. Perhaps the most closely related genus is Dystactospongia (Miller 1882: 42). In Mil- ler's original description the spicule struc- ture was poorly defined, but in a sub- sequently discovered specimen of Dystactospongia madisonenses Foerste, de- scribed by Rigby (1966), the spicule pattern is preserved. In this latter specimen, the skeletal net appears as a confused mass of Dec. 1977 RIGBY, GILLAND: FOSSIL SPONGE 477 fibrous spicules at first glance, but is com- posed of closely spaced spicules which in- tertwine—almost like a batch of spaghetti. Individual spicules line the canal openings, much as in the present specimens. Dystac- tospongia is a massive lobate, or digitate, to semiglobular form, however, in contrast to the simple, open, thin-walled, bowl shape of the present sponge. If our interpretation of the spicule pattern on the Idaho specimens is correct and the forms are related to Dys- tactospongia from the Ordovician of the midcontinent, Loganiella should be included in the Order Choristida in the Class Demos- pongia. Etymology.— The genus Loganiella is named after the Logan River, where the type material was recovered. Type species.— Loganiella phnsoni. Loganiella phnsoni, n. sp. Figs. 2-6 Description.— Bowl-shaped to cup- shaped subcylindrical sponges (Fig. 1) with a flattened base and nearly vertical to slightly inclined walls that surround a simple, broad, open spongocoel. The base of the sponge is slightly concave to nearly flat, but with a rounded margin. From the rounded margin of the base, the walls ex- tend vertically to slightly inclined inward for approximately 3 cm to the upper edge of the fragment. The holotype (BYU 1090; Figs. 3, 5, 6) is nearly circular, when viewed from above, and has a maximum diameter of 72 mm. The base is slightly warped, possibly during growth because the spicule net apparently is solidly fused and the sponges appear to show httle distortion. Wall thickness is moderately uniform, ranging from 3.7 mm at the thin oscular(?) margin up to a max- imum of 4.5 mm where the flat base curves upward into the subcylindrical wall. A paratype (BYU 1091; Figs. 2, 4) is of the same general shape and has a maximum diameter of 70 mm and is from 20 to 22 mm high. The base of the sponge is approx- imately 10 mm thick. The wall thickens to a maximum of 15 mm at the lower-most margin of the spongocoel but then thins to only 5 to 6 mm at the upper edge of the preserved fragment. In the paratype a mod- erately long section of what is considered to be the upper oscular rim is preserved. It has a rounded, although somewhat chan- neled, upper surface. The wall structure shows well completely around the upper edge. Where the wall has been broken and irregularly silicified the radial canal pattern shows moderately well. The canal pattern shows best in the holo- type where four different sizes of openings are present. The largest of these occur on the exterior of the base and they are par- ticularly well defined in an area newly etched free of matrix. These large canals are circular to elliptical. They are particu- larly elliptical where two canals appear to converge to form one circular opening within the interior. Circular single canals range from 0.20 mm to 0.6 mm in diame- ter. These canals are separated by relatively thick skeletal tracts that range from 0.3 to 1.5 mm across. This particular canal series is spaced moderately irregularly over the surface, but in some areas occurs in an al- most rectangular pattern. In a characteristic area 5 mm square there are from 20 to 25 of the larger canal openings. Where the pattern is moderately regular or linear they are spaced approximately 1 mm apart. There may be five or six of the larger canal openings per 5 mm, as measured along any one single series. A second, smaller series of canals ranges from 0.15 to 0.30 mm in diameter. These occur irregularly within the skeletal tracts and appear to parallel or possibly converge with the larger canal openings in the interi- or of the wall. Where best preserved, these small canals are spaced from 0.4 to 0.6 mm apart along an individual skeletal tract but, in general, they occur in the dense skeletal net. Some of these openings are irregular and may be produced, in part, by irregular- ities of silicification. Others, however, are sufficiently circular and so well defined that they must indicate original openings in the net of the sponge. These canals are spaced from 0.1 to 0.2 mm apart in cross sections 478 GREAT BASIN NATURALIST Vol. 37, No. 4 ^L'l^^ !>A :v^^; Dec. 1977 RIGBY, GILLAND: FOSSIL SPONGE 479 through the wall and, like the large canal series, appear to pass directly through the wall and to be almost straight. The inner surface of the spongocoel is somewhat more irregular than the exterior and the canals there are poorly defined. Circular canals ranging from 0.25 to 0.6 mm in diameter occur on the silicified or encrusted interior wall. Most canal openings here range from 0.25 to 0.30 mm across, are terminations to the straight radiating canal series, and connect to the larger ostia on the exterior. These spongocoel openings, in general, are separated by skeletal tracts from 0.4 to 0.6 mm across. These tracts, like those on the exterior, have rounded cross sections so that the diameter of the canal openings flare at the immediate spongocoel surface but narrow a few tenths of a millimeter into the wall from the spon- gocoel margin. Smaller openings, from 0.15 to 0.3 mm across, are also common and may represent inner termination of some of the large radi- ating canals. The spongocoel margin has a somewhat frosted or rounded appearance, in contrast to the much more sharply defined canals and skeletal tracts of the exterior. The gas- tral or interior surface of the wall is irregu- larly porous and there is practically no pat- tern of canal placement, such as the linear pattern seen on the exterior. Because of silicification, spicules in the interior of the tracts are not preserved, but impressions of the spicules are evident on the exterior of a few tracts, particularly on the spongocoel margin. They are curved to moderately straight rodlike spicules which appear to be diactines. It is impossible to tell in this preservation whether these are rhizoclone spicules with a smooth exterior facing toward the canals or whether they are oxeas or simple curved spicules as in Dystactospongia. These spicules are from 0.02 to 0.035 mm in diameter and up to 0.25 mm long, although on none can the to- tal length be recognized because the tips are either broken or lost in the ropy fabric. Spicules are curved to outline the circular canals and lie tangential to the main fibers. On some tracts, where the spicules show particularly well, there are from 10 to 15 spicules coating the exterior. This would suggest that individual tracts may have as many as 50 to 75 spicules in a single cross section. The spicules generally are elongate, parallel to the long dimensions of the some- what reticulate mesh. In the cross- connecting bars, for example, they are tan- gential to the spongocoel wall and cross- connect the tracts which are extending di- rectly through the sponge wall. The spicule pattern is less clearly preserved on the exte- rior—a few spicules' impressions here and there— and the generally ropy appearance of the tracts is poorly expressed. Etymology.— The species is named in honor of Dr. Chester O. Johnson, who rec- ognized the sponge origin of the material and made most of the collection available. Type materials and occurrence.— The Figs. 1-6. Fossil sponges from the Garden City Limestone of southeastern Idaho: 1. Small spherical Hindia{?) associated with the holotype of Loganiella, from the Garden City Limestone, Logan River locality; 87 percent of natural size. Figs. 2-6. Loganiella johnsoni. 2, 4, Paratvpe, BYU 109L 2, Flat, coarsely silicified base of the paratype with some calcareous matrix in the sponge wall. 4, View from above into the simple spongocoel of the coarsely silici- fied paratype showing the moderately thick walls of this particular specimen. 3, 5, 6, Holotype BYU 1090. 3, View of the flat base of the holotype showing the somewhat rectangularly spaced skeletal tracts of the sponge. The opening at the lower right is an artifact of preservation. Small light triangular area in the upper right center is a residum of matrix. Spicules show under the microscope near the base of this remnant. 5, View down into the flat bottom of the spongocoel of the holotype, here partially encrusted by matrix. The thin wall of this specimen shows above, in the wall fragment in the upper part of the specimen. The distinctly nonpatterned openings of the canals on the interior of the sponge contrast sharply with the distribution seen in the basal part of the exte- rior. The artificial opening through the sponge wall here is at the top of the specimen. 6, Side view showing the characteristic flat base, rounded margins, subcylindrical to broadly conical walls, and the moderate regularity of canal spacing on the upper flanks of the sponge; 87 percent of natural size. 480 GREAT BASIN NATURALIST Vol. 37, No. 4 sponges are known, at present, only from the upper part of the Garden City Forma- tion, in rocks of zone J to L, in southern Idaho. Holotype, BYU 1090, and paratype, BYU 1091. Literature Cited Hi.NTZE, L. F. 1951. Lower Ordovician detailed stra- tigraphic sections for western Utah. Utah Geol. Min. Survey Bull. 39: 100. 1952. Lower Ordovician trilobites from west- em Utah and eastern Nevada. Utah Geol. Min. Survey Bull. 48: 249. Miller, S. A. 1882. Description of two new genera and eight new species of fossils from the Hudson River Group, with remarks upon others: J. Cin- cinnati Soc. Nat. Hist. 5: .34-44. Richardson, G. B. 1913. The Paleozoic section in northern Utah. Amer. J. Sci. 36 (4): 406-416. 1941. Geology and mineral resources of the Randolph quadrangle, Utah-Wyoming. U.S. Geol. Survey Bull. 923: 55. RiGBY, J. K. 1966. Microstructure and classification of an Ordovician sponge, Dtjstactospongia nuidiso- nensis Foerste from Indiana. J. Paleont. 40: 1127-1130. Ross, R. J. Jr. 1949. Stratigraphy and trilobite faunal zones of the Garden City Formation, northeastern Utah. Amer. J. Sci. 247: 472-491. 1951. Stratigraphy of the Garden City Forma- tion in northeastern Utah and its trilobite faunas. Yale Peabody Mus. Bull. 6: 161. HOST-INDUCED MORPHOLOGICAL VARIATIONS IN THE STRIGEOID TREMATODE POSTHODIPLOSTOMUM MINIMUM (TREMATODA: DIPLOSTOMATIDAE). IV. ORGANS OF REPRODUCTION (OVARY AND TESTES), VITELLINE GLAND, AND EGG' James R. Pahnieri- Abstract.— a variety of amphibian, reptilian, avian, and mammalian hosts were used in experimental devel- opment of Posthodiplostomiim minimum. Results of this study indicate that the organs of reprodviction (testes and ovary) as well as the vitelline gland and egg underwent host-induced morphological variations. Due to the lack of host specificity of P. minimwn as well as the overlap of reported egg and body sizes of species of Post- hodiphstomum, it is apparent that several reported species are not valid. Throughout the Uterature, members of the trematode family Diplostomatidae have been classified and divided based upon the size and shape of the anterior and posterior testes, extent of vitelline gland, position of the ovary relative to the testes and the size of the egg (Dubois 1937, 1952, 1964, 1968, 1970). In the above cases, Dubois consid- ered the strong host specificity supposedly demonstrated by strigeoids to be the basis for his extensive taxonomic revisions. In re- cent years several investigators have shown that strigeoid trematodes are not as phys- iologically host specific as previously sug- gested. Ulmer (1961) emphasized the need for additional experimental data relative to host specificity in order to assess the valid- ity of Dubois's use of it as a major criterion for establishing taxonomic relationships. Pal- mieri (1973, 1975, 1976) clearly demon- strated that Posthodiplostomiim minimum is able to develop to a gravid state in many host species within all vertebrate classes ex- cept fishes. It is doubtful, however, that host specificity in a strict sense is of value in differentiating species of Posthodiplostomum. Analysis of data presented in this paper clearly indicates that testes number, size and shape, ovary position, vitelline gland distribution and egg size undergo great morphological variations when experimen- tally developed in amphibian, reptilian, avian, and mammalian hosts. Similar results dealing with morphological variation have been reported by Watertor (1967), Blankes- poor (1971), Kinsella (1971), Campbell (1973), and Palmieri (1973, 1975). Material and Methods The material and method section of this paper has been reported by Palmieri (1976). Results and Discussions OvABY.— Five variables were analyzed in studying the ovary of adult P. minimum, namely: (length, width, cross sectional area, ratio of length to width, and the variation in the position of the ovary relative to ante- rior and posterior testes). A complete analy- sis of the above data can be found in Table 1. Variations in ovary position of P. min- imum are found in Figures 1-6 and Table 2. The ovary of P. minimum commonly var- ies in position from anterior to the anterior testis (Figs. 1-2) to lateral (Fig. 3) or poste- rior (Figs. 4-6) to it. In worms recovered from all four classes of experimentally fed definitive hosts, the ovary was situated most commonlv to the left of the anterior testis 'See Part II of this study. Great Basin Nat. 37:137, 1977. 'Department of Zoology, Iowa State University, Ames, Iowa 50010. Present address. Department of Zoology, Brigham Young University, Provo, Utah 84602. 481 482 GREAT BASIN NATURALIST Vol. 37, No. 4 Table 1. Analysis of ovary measurements of P. ininiinwn from experimental definitive hosts." Variable Means Mean of Combined Standard Host Class Deviation Amphibian Reptilian Aves Mammalia Ovary Length (OL) Ovary Width (OW) Cross-Sectional Area of Ovary (AOLXOW) Ratio of Ovary Length to Width 0.027 0.007 0.029 0.013 0.031 0.030 0.029 0.008 0.030 0.014 0.033 0.031 0.001 0.882 0.001 0.241 0.001 0.001 0.542 0.001 0.940 0.001 0.976 'All measurements in Figs. 1-6. Variations in position of ovary as depicted in Table 2. Dec. 1977 PALMIERI: TREMATODE VARIATION 483 (Fig. 3). It was not uncommon to find the ovary in several of the above positions, however, within worms recovered from any given genus of cold-blooded definitive host. Variations such as these are much more common in poikilothermic than in homio- thermic definitive hosts. Testes.— Ten variables were used in de- scribing the size, number, and relationships of the anterior and posterior testes (anterior testes length and width, posterior testes Table 2. Variation in ovary position relative to test- es in adult Pasthoclipbstomum minimum recovered from experimental amphibian, reptilian, avian, and mammalian Hosts. Experimental Host Class Amphibia Reptilia Aves Mammalia Ovary Position Corresponding to Figures 1-6 2,3,5 2,3,6 1,2,3,4,5 2,3,4,5 Table 3. Analysis of testes measurements of P. minimum from experimental definitive hosts. Variable Mean of Combined Standard ^^^"' Host Class Deviation Amphibian Reptilian Aves Mammalia Anterior Testis Length (ATL) Anterior Testis Width (ATW) Posterior Testis Length (PTL) Posterior Testis Width (PTW) Cross-Sectional Area of Anterior Testis (AATLXPTL) Cross-Sectional Area of Posterior Testis (APTLXPTW) Ratio of Anterior Testis Length to Width (RATLPTW) Ratio of Posterior Testis Length to Width (RPTLPTW) Ratio of Cross- Sectional Area of Anterior Testis to Posterior Testis (RATXXPT) 0.048 0.017 0.042 0.031 0.066 0.150 0.056 0.136 0.087 0.198 0.050 0.017 0.048 0.027 0.( 0.138 0.072 0.123 0.079 0.( 0.008 0.004 0.006 0.002 0.013 0.008 0.004 0.006 0.003 0.013 0.355 0.153 0.332 0.446 0..3.38 0.408 0.,399 0.,527 0.251 0..371 1.016 0.658 1.174 0.522 1.067 0.057 0.182 0.053 0.010 0.010 0.352 0.308 'All measurements in mm. 484 GREAT BASIN NATURALIST Vol. 37, No. 4 length and width, cross-sectional area, ratio of testes length to width, ratio of the cross- sectional area of the testes, and testes num- ber). An analysis of the above data can be found in Tables 3 and 4. A comparison of the morphological variation is drawn at the light level on Figures 9-21.- The appearance of the testes, including their size, shape, and number, varied widely in specimens of P. minmium recovered from various vertebrate hosts. A definite trend is noticeable in accordance with the specific class of vertebrate host employed. In worms developed within avian hosts, anterior and posterior testes occupy a large area of the hindbody (Figs. 14, 16-) and are large, well Table 4. Variations in testes number in adult Post- hodiplostomum minimum recovered from experimental amphibian, reptilian, avian, and mammalian hosts. Experimental Host Class Testes Number Amphibia Reptilia Aves Mammalia 0,2 1,2 2 1,2 Table 5. Variation in vitelline gland distribution in adult Posthodiphstomum minimum recovered from ex- perimental amphibian, reptilian, avian, and mamma- lian hosts. Experimental Host Class Vitelline Gland Distribution" Amphibia Reptilia Aves Mammalia 0,2,3 0,2,3 1,2 0,2,3 •0. No vitellaria present 1. From posterior region of the holdfast organ to region between ace- tabulum and oral sucker (corresponds to Fig. 13, Part III of this study). 2. From midregion of anterior testis to region between acetabulum and oral sucker (corresponds to Fig. 14, Part III of this study). 3. From midregion of holdfast organ to anterior region of acetabulum (corresponds to Fig. 15, Part III of this study). developed, and approximately twice as wide as they are long. In mammalian hosts, worms have less normal-looking testes (Fig. 18"), but the latter are about three to four times as wide as long (Figs. 17, 21-), and are "dumbbell" shaped. Within poikilother- mic hosts, testes of P. minimum appear even wider and sometimes show twists and diverticula (Figs. 11,13'). In several reptilian and mammalian hosts, no posterior testis ap- pears. Several worms recovered from am- phibian hosts showed no signs of testes de- velopment. Throughout all classes of hosts experimentally infected, worms from the more ecologically abnormal hosts tend to develop extremely wide testes with trends toward reduction in size of the posterior testis. Vitelline gland distribution.— Only the distribution of vitelline glands in the forebody and hindbody was considered in the adult P. minimum recovered during this investigation. This data was reported and diagrammed in Figures 13-15, Part III of this study. In most specimens of P. minimum recov- ered from avian hosts, vitelline follicles are large and granular. Follicles are most heavi- ly concentrated in an area extending from a region equidistant between acetabulum and oral sucker to the anterior half of the ante- rior testis. In some amphibian, reptilian, and mammalian hosts, vitelline di.stribution is identical to that found in the avian hosts, but follicles are smaller and less granular. In worms recovered from some reptilian and avian hosts {Chnjsemys, Cohimba, and Cal- lus), vitelline distribution is reduced to an area surrounding only the holdfast organ and the acetabulum. Worms recovered from amphibian hosts demonstrate a further re- duction in vitelline distribution to a point where only a few follicles scattered around the holdfast are present. In some cases (am- phibian, reptilian, and mammalian hosts) a complete absence of follicles resulted, even though age of the worms was sufficient to have permitted full development of these follicles. 'See Part II of this study. Great Basin Nat. .37:137. 1977. Dec. 1977 PALMIERI: TREMATODE VARIATION 485 Egg.— Six variables were analyzed in evaluating measurements of the eggs of P. minimum. Linear measurements consisted of egg length and width from worm specimens within a single host class. Two relationships of the above data were also studied, namely the ratio of the egg length means to the egg width means and the mean value for the cross-sectional area. A complete analysis of the above data can be found in Table 6. Egg shape and development of P. min- imum vary among experimental poikilother- mic and homiothermic definitive hosts uti- lized in this investigation. P. minimum eggs from amphibian and reptilian hosts are small and round. Those recovered from am- phibian hosts were not viable and no mira- cidial development could be detected. Egg .size was largest from worms recovered from homiothermic hosts. The largest and most viable eggs of P. minimum are recovered from piscivorous avian and carnivorous mammalian hosts, and such eggs are charac- teristically oval and operculate. Egg number per worm varies from 1-2 in amphibian and reptilian to 1-5 (typically 3) in avian and mammalian hosts. One of the principal factors contributing to the complex problems of speciation with- in the genus Posthodiplostomum and per- haps other genera of digenetic trematodes has been the lack of experimental studies providing experimental data on host- induced variation at the intraspecific and interspecific levels. This study, ba.spd upon careful examination of over a thousand specimens of Posthodiplostomum minimum recovered from experimental hosts, empha- sizes the pronounced morphological varia- tion resulting from the rearing of P. min- imum in different vertebrate definitive hosts, from any individual host class. Char- acters undergoing a significant change (F .01 level of significance or higher) include egg length and width, both ratio and area of the egg length means and the egg width means, and body length. These are the poorest characters for taxonomic identi- fication of P. minimum since they show a significant morphological variation due to influences of the class of host in which de- velopment takes place. This study provides evidence that the only significant morphological criteria for the determination of Posthodiplostomum minimum developed within any definitive host class are the ratio of the body length to body width, oral sucker cross-sectional area, acetabular index of length, acetabulum length, oral sucker width, and acetabulum width. Ratios between holdfast length and width and anterior testis length and width, Table 6. Analysis of egg measurements of P. minimum from experimental definitive hosts." Mean of Combined Standard Means Variable Host Class Deviation Amphibian Reptilian Aves Mammalia Egg Length (EL) Egg Width (EW) Cross-Sectional Area of Egg (AELXEW) Ratio of Egg Length to Width (RELEW) 0.078 0.007 0.034 0.025 0.084 0.085 0.030 0.054 0.028 0.020 0.058 0.057 0.004 0.005 L445 0.095 0.013 L324 0.001 0.005 0.004 1.481 'AH measurements i 486 GREAT BASIN NATURALIST Vol. 37, No. 4 as well as the oral sucker length and width, also aid in the taxonomic identification of this species. It is apparent that, on the basis of those variables studied in this investigation, avian definitive hosts are by far the most suitable ones for P. minimum. Internal organs are best developed with respect to size, and their position remains more constant than do those of worms reared in other verte- brate classes. Mammalian definitive hosts also serve as "normal hosts" to a lesser de- gree, for the extent of the vitellaria and the nature of the oral sucker are somewhat Table 7. Comparison of body and ported in this study." measurements of reported species of Posthodipbstomum with those Species of Posthodiplostomum Body Length Min. Max. Body Width Min. Max. Egg Length Min. Max. Egg Width Min. Max. P. australe (Dubois 1937) P. bi-ellipticiim (Dubois 1958) P. botauri (Vidyarthi 1938) P. boydae (Dubois 1969) P. brevicaudatum (Nordmann 1832, Wisniewski 1958) P. cheni (Yang 1959) Odening 1962 P. cuticola (V. Nordmann 1932, Dubois 1936) P. grande (Diesing 1850) Dubois 1936 P. grayi (Verma 1936) Dubois 1938 P. huesingi Odening, 1962 P. impraeputiatwn (Dubois 1934) Dubois 1936 P. larai (Refuerzo et Garcia 1937) P. macrocotyle Dubois, 1937 P. microsicya Dubois, 1936 P. milui Fotedar et Reina 1965 P. nanum Dubois, 1937 P. obbngum Dubois, 1937 P. opisthosilya Dubois, 1969 P. podicipitis (Yamaguti 1939) P. prosostoinum (Dubois et Rausch) (Dubois 1952 comb. emend.) P. scardinii (Shulman in EXibinin 1952) 2.5.30 0.091 0.450 0.500 0.082 0.090 0.052 0.0,52 0.062 1.190 1.230 0.530 0.560 0.078 0.078 0.056 0.056 1.200 1.562 0.256 0.720 0.080 0.080 0.064 0.064 0.990 1.140 0.120 0.180 0.097 0.097 0.058 0.058 1..300 1.920 - - 0.090 0.110 0.050 0.065 1.269 1.7.35 0.268 0.454 - - - - 0.670 2.310 0.220 0.920 0.073 0.110 0.051 0.088 1.620 2.250 0.250 1.8.30 0.070 0.086 0.045 0.055 0.830 1.077 0.210 0.390 0.070 0.093 0.042 0.074 1.080 2.600 0.300 1.600 0.073 0.110 0.051 0.080 1.200 2.110 0.160 0.430 0.092 0.103 0.055 0.065 0.810 1.280 _ _ 0.070 0.081 0.053 0.074 0.890 1.200 0.200 0.290 0.077 0.089 0.050 0.057 1.200 1.990 0.190 0.290 0.060 0.072 0.038 0.045 0.410 0.950 0.090 0.610 0.062 0.076 0.042 0.048 2..340 2.700 0.300 0.540 — _ — _ 0.910 1.160 - - 0.092 0.096 0.057 0.070 0.660 1.100 0.200 0.310 0.090 0.093 0.057 0.066 0.058 P. suni N. T. Lee, 1964 1.840 1.890 0.301 0..301 0.073 0.073 0.053 0.053 P. minimum (MacCallum 1921) 0.840 1.750 0.160 0.600 0.073 0.091 0.048 0.064 This study Mean Mean Mean Mean P. minimum (Amphibia) 0.517 0.269 0.039 0.028 P. minimum (ReptiUa) 0.4.35 0.230 0.025 0.020 P. minimum (Aves) 0.614 0.288 0.084 0.058 P. minimum (Mammalia) 0.589 0.296 0.085 0.057 'All measurements in mm. Dec. 1977 PALMIERI: TREMATODE VARIATION 487 more variable than in avian hosts. Poikilo- thermic hosts demonstrate marked variation and abnormal development. Especially among the Reptilia are extreme effects no- ticeable in developing worms, as evidenced by great reduction in body and organ size when compared with individuals reared in other host classes. Throughout the literature, extensive syn- onymy exists for many species of Post- hodiplostomum. Neither Dubois (1968, 1970) nor Yamaguti (1971) agree as to the number of valid species of this genus. As indicated by this investigation, few characters can be effectively utilized in separating adults of P. minimum reared in a great variety of poi- kilothermic and homiothermic hosts. Experi- mentally developed P. minimum from one or more classes of vertebrate hosts demon- strate such a wide degree of morphological variation that one doubts the validity of the number of species currently placed in the genus Posthodiplostomum as well as related genera in this strigeoid group. Certain char- acters such as body shape, ovary position relative to the testes, extent of vitellaria, relative size of fore- and hindbody, oral sucker shape, and host specificity have been and continue to be used in separating gen- era of the Diplostomini. Even if one accepts Postliodiplostomum as a valid genus isolated morphologically and genetically from other related genera, the problem of speciation within the genus is even more confusing. Characters used by Dubois (1968, 1970) and Yamaguti (1971) (such as "testes tandem," "anterior testis oval," "posterior testis larger than the ante- rior testis") appear to be of little value, for this study has shown that host influences upon the adult P. minimum render such characters much too "plastic" to be of tax- onomic importance. Because of inconsis- tencies in published accounts of egg sizes and descriptions of the 23 species of Post- hodiplostomum, it is difficult to make a comparative analysis of them in relation to experimental results derived from this in- vestigation. However, four variables (body length, body width, egg length, and egg width) have been compared (Table 7). Ex- amination of data in this table makes it ap- parent that several reported species of Post- hodiplostomum are probably not valid because of the considerable overlap in mea- surements of these structural features. Acknowledgments The author wishes to thank Dr. Martin J. Ulmer for his time and guidance throughout this research project. Dr. D. Cox and Paul Dubose for data analysis. Dr. Darwin Wit- trock for aid in fieldwork, and Mr. James Amrine for specimen preparation for SEM. Appreciation is also extended to the Iowa State Conservation Commission and the Federal Bureau of Sport Fisheries for ani- mal collection and to the Iowa State Uni- versity Departments of Zoology, Genetics, and Statistical Computations. This study was supported in part by the National Sci- ence Foundation Grant GB-23057, by the Thomas H. MacBride Scholarship from the Iowa Lakeside Laboratory, by the Brigham Young University Department of Zoology, and by the Department of International Health, School of Medicine, University of California, San Francisco, from the National Institute of Allergy and Infectious Diseases, National Institutes of Health, U.S. Public Health Service Grant AI 10051 (US ICMR). Literature Cited Blankespoor, H. 1971. Host-parasite relationships of an avian trematode, Plagiorchis noblei Park, 1936. Ph.D. Dissertation. Iowa State University Library. (Library Congr. Card No. Mic. 71-14, 205). Uni- versity Microfilms, Ann Arbor, Mich. (Diss. Abstr. .31:12). Campbell, R. 1973. Studies on the host-specificity and development of the adult strigeoid trematode Cotylurus flabelliformis. Trans. Am. Microsc. Soc. 92: 256-265. Dubois, G. 1937. Sur guelques strigeides (Notes pre- liminaires). Rev. Suisse Zool. 44: .391-396. 1952. Revision de quelques strigeides (Trema- toda). Bull. Soc. Neuchatel. Sc. 75: 169-178. 1964. Du Statut de quelques Strigeata La Rue 1926 (Trematoda). Bull. Soc. Neuchatel. Sc. 87: 27-71. 1968. Synopsis des strigeidae et des diplos- tomatidae (Trematoda). Bull. Soc. Neuchatel. Sc. 1-258 pp. 1970. Synopsis des strigeidae et des diplos- 488 GREAT BASIN NATURALIST Vol. 37, No. 4 tomatidae (Trematoda). Bull. Soc. Neuchatel. Sc. 259-727 pp. KiNSELLA, J. 1971. Growth, development and in- traspecific variation of Quinqueserialis quinquese- rialis (Trematoda: Notocotylidae) in rodent hosts. J. Parasitol. 57: 62-70. Palmieri, J. 1973. Additional natural and experimental hosts and intraspecific variation in Post- Iwdiplostomum 7ninimitm (Trematoda: Diplos- tomatidae). J. Parasitol. 59: 744-746. 1975. Host-parasite relationships and in- traspecific variation in the strigeoid trematode Posthodiplostomum minimum (Trematoda; Diplos- tomatidae). Utah Acad. Sci., Arts, and Letters. 52: 68-69. 1976. Host-parasite relationships and in- traspecific variation in the strigeoid trematode Posthodiploshnnum minimum (Trematoda: Diplos- tomatidae). Great Basin Nat. 36(3): 334-346. Ulmer, M. 1961. Passerine birds as experimental hosts for Posthodiplostomum minimum (Trematoda: Diplostomidae). J. Parasitol. 47: 608-610. Watertor, J. 1967. Intraspecific variation of adult Te- lorchis bonnerensis (Trematoda: Telorchiidae) in amphibian and reptilian hosts. J. Parasitol. 53: 962-968. Yamaguti, S. 1971. Synopsis of digenetic trematodes of vertebrates. Vol. 1. Keigaku Publishing Co. Tokyo, Japan. 1074 pp. A SURVEY OF BREEDING AND MIGRATORY BIRDS SOUTHWEST OF FARMINGTON, NEW MEXICO Diiane A. ToUe' Abstract.— The relative abundance and habitat affinities of breeding and migratory waterfowl were documented for a 16,556 ha (40,880 acre) area of the Navajo Indian Reservation, San Juan County, New Mexico. The objec- tive of the 1973-1974 study was to obtain baseline data for use as part of an environmental assessment for coal gasification plants and an associated strip mine proposed for an area not previously described in ornithological journals. Roadside breeding-bird surveys resulted in the observation of 26 species of birds, of which the horned lark was most abundant. Additional breeding-bird observations were made during afternoon reconnaissance surveys around stock ponds and arroyos, and by a helicopter survey of cliff-nesting raptors. Nine of 30 raptor nests located were active. Migratory waterfowl surveys were made in September and November at four stock ponds and a marsh on the study area. Peak waterfowl numbers were present in September, when teal and shovelers were the most abun- dant species. The importance of stock ponds to breeding and migratory birds and the significance of cliffs to nesting raptors were discussed. Details of observations made for five species of birds designated "threatened" or "status undeter- mined" were presented. Avifauna of a 16,556 ha (40,880 acre) area (hereafter called the study area) lo- cated in San Juan County, New Mexico, about 48.3 km (30 miles) southwest of the city of Farmington were studied during four- to seven-day periods in mid-November 1973 and late February, early June, late September, and mid-November 1974. This overgrazed grassland ecosystem is located in the Navajo Indian Reservation bordering the Chaco River at an elevation of about 1615 m (5300 ft). It is intersected by arroyos which provide a maximum relief of about 122 m (400 ft). The climate of the study area is arid, with an average of 16.8 cm (6.6 inches) of precipitation per year. The period from July to October normally accounts for half of this precipitation. The study area includes plants and ani- mals typical of both the grassland and basin sagebrush biociations (Kendeigh 1961). Al- though vegetation is sparse, it includes grasses, forbes, and low-growing shrubs. A few trees and larger shrvibs are located in the arroyos and near ponds. The vegetation has been heavily grazed by livestock, in- cluding sheep, cows, horses, and goats. Plants requiring more water are located at the small, man-made stock ponds scattered over the area. The objective of this study was to docu- ment relative abundance and habitat af- finities of species of breeding birds and mi- gratory waterfowl, using an area not previously described in ornithological jour- nals, where coal gasification plants and an associated strip mine are planned. These two general groups of birds were considered to be the most likely to be affected by this major industrial development. The resultant baseline data on avifauna of the area was used as part of an environmental impact re- port. A substantial number of ducks are present in the San Juan River Valley during the mi- gratory season, due in part to the concen- trated production of food crops in this val- ley (Huey 1967). Bellrose (1971) considers the San Juan River Valley to be a major (nationwide) waterfowl concentration area. Undoubtedly, the attractiveness of the San 'Ecology and Ecosystems Analysis Section, Battelle's Columbus Laboratories, 505 King Avenue, Columbus, Ohio 43201. 489 490 GREAT BASIN NATURALIST Vol. 37, No. 4 Juan River Valley influences waterfowl use of ponds on the study area. The San Juan River is part of the Upper Colorado River Basin, which is known as an important migratory route for waterfowl and shorebirds, as well as smaller passerine birds. The Upper Basin has been the site of continued development of aquatic habitat in the form of lakes and reservoirs. This has materially changed the status of waterfowl and shorebirds thoughout the entire area (Hayward, 1967). Morgan Lake and Navajo Reservoir are examples of the water bodies which have recently been created in the San Juan River area of New Mexico. Methods Avifauna of the study area were surveyed by three methods during the breeding sea- son and by two additional methods during the migratory seasons. All common names of birds are reported in the form designated by the Fifth AOU checklist (Wetmore 1957) and the twenty-third supplement (Eisen- mann, 1973) (See Appendix I.). Breeding bird survey methods.— Road- side surveys: Breeding birds were surveyed during 1 to 7 June 1974 by the following three methods: (1) roadside surveys, (2) re- connaissance surveys along stock pond ar- royos, and (3) a helicopter survey of cliff- nesting raptors. The roadside survey devel- oped by the U.S. Department of the Interi- or (1973a) was conducted on four mornings to obtain an index of abundance of breeding birds. The only variance from the standard operating procedure was the seven-hour pe- riod required for completion of the survey. Four and one-half hours are normally al- lotted to reach the end of a 40.2 km (25 mile) route, with three-minute stops made 0.8 km (0.5 miles) apart to count all birds within a 0.4 km (0.25 mile) radius. On this study area, the additional time was neces- sary due to the rough roads and back- tracking required to reach a representative portion of the area. All four roadside surveys were nm on mornings without precipitation or fog. Wind speed was less than Beaufort 3 (8-12 mph) during all survey periods except for the finish of the 2 and 3 June surveys. Midafternoon reconnaissance surveys: Af- ternoon surveys were conducted at five stock ponds (only one had water during the survey period) and their adjacent vegeta- tion, plus five arroyos. Walking surveys were made by a party of two observers for a total of eight hours after each of the four roadside surveys, and during the afternoon of 5 June. A special effort was made to lo- cate nests. Helicopter survey of raptor nests: Four hours were spent searching for cliff nests of large raptors from a Bell B-1 helicopter on 4 June 1974. A total of 112.7 km (70 miles) of suitable cliff-nesting areas were searched, using the helicopter flight patterns sug- gested by White and Sherrod (1973) to avoid disturbance of adult and nestling rap- tors. The flight was made at a time when most of the raptors in the area had finished incubation, but before the majority of the nestlings had fledged. During this period the adult raptors are least likely to desert the nest. One of the two observers plotted nest lo- cations on topographic maps. The number and description of eggs or nestlings, plus nest materials, diameter, and height were recorded for identification of inactive nests or nests where adults were absent. Photos were taken of nearly all of the nests. Migratory-bird survey methods.— Wa- terfoivl surveys: Waterfowl surveys were made during the following four periods: (1) 13-16 November 1973, (2) 20-24 February 1974, (3) 22-25 September 1974, and (4) 11- 14 November 1974. The November and February survey periods were chosen to coincide with the peak period of both wa- terfowl numbers and variety of species mi- grating through San Juan County. The Sep- tember period was chosen to coincide with peak populations of early migrants (J. L. Sands, pers. comm.). Five waterfowl observation sites were se- lected on the study area. Of these five sites, four were at man-made stock ponds created by damming arroyos, and one was at a marsh created by overflow from a spring- Dec. 1977 TOLLE: new MEXICO BIRDS 491 fed stock pond. When any one of the ponds was dry or frozen during a survey period, no counts were made. Since all ponds were frozen in February 1974, no waterfowl sur- veys were made. During all other survey periods, 0.5-hour observation periods were conducted at each of the sites. General observation of non-waterfowl birds: Non-waterfowl bird species were also recorded during each of the four waterfowl survey periods listed above. An accurate census during these periods was not pos- sible, due to the nonterritorial, flocking be- havior of most nonbreeding birds. However, the total number of each species observed was recorded. These data were used to pre- pare tables of the relative frequency of ob- servation of all nonwaterfowl species. Small birds not associated with the playas or marsh were likely missed. A special effort was made to record all raptors observed while driving between survey sites. Survey Results Relative abundance and distribution OF breeding birds.— Species observed along roads: Twenty-six species of birds were heard or observed during 200 three-minute stops made over a four-day period (Table 1). The most abundant species was the Horned Lark, followed by the Mourning Dove and Black-throated Sparrow. The number of stops where a species was ob- served is an indication of how widely dis- tributed it is over the survey area. The commonness of the Horned Lark probably resulted from the fact that the majority of the study area is relatively flat, mesa habi- tat with short vegetation which is the pre- ferred habitat for the Horned Lark (Ligon 1961, Fautin 1946). Species of birds observed at the only marsh on the area were considerably differ- ent from birds seen at other survey stops. Nine species of birds were recorded only at a marsh created by overflow from a spring- fed stock pond. These nine species recorded during roadside surveys included the Great Egret, Mallard, Pintail, Green-winged Teal, Cinnamon Teal, Killdeer, Spotted Sand- piper, Tree Swallow, and Brown-headed Cowbird. Also, the following five-bird spe- cies were seen at this marsh after the three- minute survey period was over: Savannah Sparrow, Hummingbird (spp. undetermined), American Coot, Western Meadowlark, and Rough-winged Swallow. Observations of all of the above 14 species were restricted to the overflow marsh. Species observed in arroijos below ponds: Midafternoon reconnaissance surveys around stock ponds and along arroyos resulted in the observation of 21 species of birds (Table 2). Although the time of day prevented an accurate census of the avifauna, a rough es- timate of relative abundance between spe- cies indicated that the mourning dove, horned lark, and house finch were more abundant around stock ponds and arroyos than other bird species. Very few nests were located during mid- afternoon surveys, in spite of a careful search of the vegetation by two observers during eight hours of survey. This searching resulted in the location of active nests of the four following species: Red-tailed Hawk, Ferruginous Hawk, Great Homed Owl, and Cliff Swallow. The Red-tailed Hawk and Cliff Swallow nests were located in cliffs along the Chaco River. Two inactive nests believed to be House Finch nests were lo- cated at one of the dry stock pond sites. Cliff-nesting raptors: Thirty large raptor nests were located during the helicopter survey (Table 3). Nine of these nests were active (Table 4), with the majority of the occupants either Golden Eagles (two nests), Great Horned Owls (two nests), or Red- tailed Hawks (two nests). The Ferruginous Hawk, Prairie Falcon, and Common Raven were recorded at only one nest per species. One of the Golden Eagle nests may have been abandoned, since only one egg and a rock were observed in the nest, and no adult eagle was seen near the nest. Only four of the six species of raptors had successfully raised young by the survey date (Table 4). Nestling or fledgling Great Horned Owls, Red-tailed Hawks, Ferru- ginous Hawks, and Common Ravens were observed. 492 GREAT BASIN NATURALIST Vol. 37, No. 4 During the helicopter search for nests, a record was kept of all large birds (Table 5). In addition to the six cliff-nesting raptors, eight Mourning Doves, a Burrowing Owl, and two Common Nighthawks were ob- served during the four-hour flight. The Common Nighthawk was not seen during any of the other surveys; however, no eve- ning surveys were conducted. Relative abundance of migratory BIRDS.— Waterfowl at stock ponds: Ten spe- cies of waterfowl were observed during the three survey periods (Table 6). The greatest variety (6) of species were recorded during the November 1973 survey period, but by far the greatest number (268) of individuals were seen during the September 1974 sur- vey period. This large number of individuals was due mainly to the presence of Blue- winged and Cinnamon Teal. These two spe- cies of teal and the Northern Shoveler nor- mally migrate south before November in Table 1. Summary of roadside breeding-bird survey on the study area, June 1974* Four-Day Totals Daily Totals Total Stops per Species'" Common Name 1 2 3 6 Individuals Homed Lark 77 71 98 74 320 121 Mourning Dove 17 27 18 17 79 53 Black-throated Sparrow 6 23 8 7 44 27 Rock Wren 7 6 15 2 30 25 House Finch 2 14 7 7 30 20 Loggerhead Shrike'' Mockingbird Red-winged Blackbird 3 4 4 7 7 3 11 4 5 8 4 7 25 22 20 8 17 4 Say's Phoebe 3 6 3 3 15 13 Cinnamon Teal 6 2 - - 8 4 Common Raven 1 2 3 1 7 2 Killdeer 1 4 2 - 7 6 Brown-headed Cowbird _ 3 4 7 2 Green-winged Teal - 1 2 1 4 3 Ferruginous Hawk Spotted Sandpiper Mountain Plover'^ - 3 1 1 3 3 3 2 2 1 2 Mallard - 2 - 2 Lark Sparrow 2 _ _ 2 Great Egref* Pintail*^ - 1 1 Golden Eagle American Kestrel - 1 1 Burrowing Owl Empidonax Flycatcher Tree Swallow 1 1 1 - Totals 134 173 190 140 637 Average Ind./Stop 2.68 3.46 3.80 2.80 3.18 *Fifty three-minute stops were made at 0.8 km (0.5-mile) intervals beginning at 0.5 hour before sunrise on 1, 2, 3, and 6 June 1974. The route was re- versed on two days. ''Indicates the number of stops, out of a possible 200, when one or more individuals of a species were observed. 'Eggs, nestlings, or fledglings observed. ""Probably a late spring migrant. Dec. 1977 TOLLE: new MEXICO BIRDS 493 New Mexico (Chattin 1964, Ligon 1961). Most of the Blue-winged and Cinnamon Teal observations were lumped together be- cause the eclipse molt made positive field identification difficult during September. Female Blue-winged and Cinnamon Teal are inseparable through binoculars all year long. General observations of nonivaterfowl birds: As expected, the September 1974 sur- vey resulted in observation of the greatest number (32) of nonwaterfowl species out of the four waterfowl survey periods (Table 7). Three-fourths of these species were either shorebirds or passerines. Since the bulk of the shore and passerine bird migration takes place in late April and May and again in Table 2. General reconnaissance survey around stock ponds and along arroyos in the study area, June 1974^. Common Name Frequency of Observations'' Red-tailed Hawk<= Ferruginous Hawk*" Killdeer Mourning Dove Great Homed OwF Burrowing Owl Western Kingbird Ash-throated Flycatcher Say's Phoebe Empidonax Flycatcher Western Wood Pewee Horned Lark Cliff Swallow<^ Common Raven Rock Wren Mockingbird Northern Oriole Brown-headed Cowbird Blue Grosbeak House Finch Black-throated Sparrow ■Represents eight hours of midaftemoon walking surveys by two observ- ers on 1, 2, 3, 5, and 6 June 1974. Includes surveys at five stock ponds (only one pond had water), and five arroyos (including the dry Chaco River) on the study area. The approximate number of birds observed are indicated as follows: A = Abundant, 20+; C = Common, 8-19; U = Uncommon, 3-7; and R = Rare, 1-2. 'Adult in nest, eggs, nestlings, or fledglings observed. late August through early October (Hay- ward 1967), the September survey period was the most likely to include species in these two large families. The fewest (12) species of nonwaterfowl birds were recorded during the February 1974 survey period, when most migrants are still on their wintering grounds. The total bird species seen during each November survey (1973 and 1974) was intermediate in number between species lists for the Sep- tember and February surveys. The most frequently observed species during all four survey periods was the Horned Lark. This permanent resident of San Juan County (Hubbard, 1970) was con- sidered abundant during all survey periods except November 1974, when it was consid- ered common relative to other birds ob- served (Table 7). Four nonwaterfowl species were observed during seasons not previously recorded by Hubbard (1970) and/or Alan P. Nelson (unpubl. county checklist). These species in- cluded one Black-throated Sparrow ob- served during November 1973, one Swain- son's Hawk and 53 Sage Sparrows observed during February 1974, and four Pectoral Sandpipers observed during September 1974. Discussion Importance of stock ponds to desert AVIFAUNA.— Breeding birds: Some desert birds seen on the study area are dependent on drinking water from the 10 temporary and 4 permanent ponds on the area. In June, 1974, the unusually dry weather re- duced the normally "permanent" ponds to only 2. This reduced water supply was un- doubtedly a limiting factor for the gran- ivorous birds. Smyth and Coulombe (1971) observed that all of the birds which drank regularly at a desert spring in California were granivorous; three of these species of birds (Mourning Dove, House Finch, and Black-throated Sparrow) were also seen on the study area. Another granivorous bird which requires water every day is the Scaled Quail (Sprunt and Zim 1961). Al- 494 GREAT BASIN NATURALIST Vol. 37, No. 4 though the House Finch and Black-throated Sparrow are grain eaters, they may be able to live without drinking when succulent vegetation or insects are available (Smyth and Bartholomew 1966). The Mourning Dove, however, needs water to drink at least once a day (Sprunt and Zim 1961). This may account for the groups of 5 to 10 mourning doves recorded every day at the overflow pond. Insectivorous, carnivorous, or frugivorous birds were observed to drink very little or not at all at a California desert spring (Smyth and Coulombe 1971). These birds may be limited more by their food resource than by a lack of drinking water. Leopold (1969) suggests that water is no problem for insect feeders and raptors, since the flesh they eat is made up mostly of water. Smyth and Coulombe (1971) listed the following three species which were never seen drink- ing at the California spring: Ash-throated Flycatcher, Rock Wren, and Loggerhead Shrike. They also listed the Say's Phoebe and Mockingbird as drinking infrequently. All of the above five species were observed Table 3. Helicopter survey of raptor nests on cliffs in the study area, 4 June 1974^ Common Name Nest Number Nest Diameter (Ft.) Nest Height Above Ground (Est. in Ft.) Nest Materials Great Homed Owl ? 1 2 2 (scrape) ? 30 40 None (scrape) Sticks Golden Eagle Golden Eagle? Golden Eagle? Golden Eagle? ? 3 4 5 6 7 4-6 4-6 4-6 4 3 150 150 100 100 50 Twigs +? Twigs -I-? Twigs -I-? Twigs, rock Sticks Red-tailed Hawk 8 2 50 Sticks and grass ■? 9 p 30 Sticks ■) 10 ? 50 Sticks p 11 ? 50 Sticks Golden Eagle? 12 ? 15 Sticks Prairie Falcon ? 13 14 1.5 (scrape) ? 40 None (scrape) Sticks Common Raven 15 2 40 Sticks Common Raven? 16 2.5 55 Sticks Common Raven? 17 2.5 55 Sticks ? 18 3 55 Sticks and grass ? 19 3 60 Sticks ? 20 2 70 Sticks ? 21 2.5 60 Sticks Golden Eagle? 22 3.5 80 Sticks Ferruginous Hawk? 23 4 50 Sticks, grass, and Yucca stems Red-tailed Hawk Falcon? Golden Eagle? 24 25 26 2 scrape 3-5 40 30 40 Sticks (1 rabbit leg) None (scrape) Sticks Falcon? Ferruginous Hawk 27 28" scrape 3 50 0 (on brow of 15' hill) None (scrape) Sticks, grass, and Yucca stems ? 29 2 40 Sticks (in crevice) Great Homed Owl 30 .scrape 45 None (scrape) 'Flying time was four hours in a Bell B-1 helicopter. Two men made observations. ""Discovered in low rolling hill area from observation of adults during roadside surveys. Dec. 1977 TOLLE: new MEXICO BIRDS 495 on the study area, and none of them are considered granivorous (Martin et al. 1961). The abihty to obtain water through their food may account for the relative abun- dance of these birds (excluding the Ash- throated Flycatcher) in the roadside survey (Table 1). The overflow pond which is adjacent to a spring-fed stock pond had the greatest bird diversity of any area of comparable size on the study area. Thirteen species of birds were seen during the June survey only at this location. These species alone represent more than one-quarter of the breeding spe- cies recorded. The overflow pond is the only pond with emergent aquatic vegetation on the mine lease, and thus is the only hab- itat suitable for breeding waterfowl. The four species of dabbling ducks observed at the overflow pond (Table 1) were the only waterfowl observed on the mine lease dur- ing the June surveys. Constant disturbance by livestock drink- ing at the pond may have prevented water- fowl and other birds from nesting, since no nests were located during a search of the vegetation around the pond. Fautin (1946) suggested that disturbance by livestock at a spring in Utah may have prevented birds from nesting in the area. Also, Hensley (1954) suggested that the constant influx of birds coming to water sources in the Sono- ran desert of Arizona may have reduced the number of birds nesting around those limit- ed sources of water. The number of birds, particularly Mourning Doves, coming to the overflow pond to drink, may have increased the disturbance caused by livestock, and Table 5. Flying birds observed during helicopter survey of Wesco Mine Lease, 4 June 1974^*. Common Name Number Observed Red-tailed Hawk'' Golden Eagle'' Prairie Falcon'' American Kestrel Mourning Dove Great Homed Owl'' Burrowing Owl Common Nighthawk Common Raven'' 'Flying time was four hours in a Bell B-1 helicopter. Two men made ob- servations. nestlings, or fledglings obse Table 4. Active raptor nests located during a helicopter survey, 4 June 1974" Number and Nest Number and Number and Size" of Common Name Number Color of Eggs Size*" of Nestlings Fledglings Red-tailed Hawk 8 2 _ Red-tailed Hawk 24 1 (white w/dark blotches) — 1 (% Ad.) Ferruginous Hawk 28^ 1 (white w/reddish brown spots) 3 (W Ad.) " Golden Eagle 3 - 1 or 2 (1/4 Ad.; dead) — Golden Eagle? 6 1 (white) - — Prairie Falcon 13 2 (light brown w/dark spots) 1 (hatchling dead?) ~ Great Homed Owl 1 — 3 (1/2 Ad.) — Great Homed Owl 30 _ — 1 (% Ad.) Common Raven 15 — 5 (1/6 Ad.) ~ "Numbers correspond to nest descriptions in Table .3. Traction of adult (Ad.) size. 'Discovered in low rolling hill area during roadside survey. 496 GREAT BASIN NATURALIST Vol. 37, No. 4 thus added to the unsuitabihty of this area for nesting. Migratory Waterfowl: Edminster (1964) has noted that the stock ponds in the high plains of New Mexico are not only used for nesting and rearing of young waterfowl, but were visited by surprisingly large numbers of waterfowl and shorebirds during migra- tory periods. Ponds with aquatic plants and /or ponds with shores that are not bare or overgrazed were most heavily used. Also, dugouts or pit-type ponds are less useful to waterfowl than ponds formed by damming arroyos. Since the overflow pond had grad- ually sloping sides and emergent vegetation, it is not surprising that many more ducks were seen on it than any other stock pond. An abundant food supply of aquatic in- sect larvae and mollusks was probably one reason that the overflow pond was heavily used by migrant Blue-winged Teal, North- em Shovelers, and Bufflehead. During the September 1974 survey, teal accounted for 75 percent of all waterfowl recorded on the study area (Table 6), and all but four ducks recorded were observed at the overflow pond. In addition, 15 of the 17 Bufflehead seen on the study area during both Novem- Table 6. Migratory waterfowl'' observations at four stock ponds and a marsh on the study area. Number of Observations Nov. Sept. Nov. Common Name 1973 1974 1974 Mallard 8 1 Gadwall 2 Pintail 12 Green-winged Teal 2 20 Cinnamon Teal 3 Blue-winged or Cinnamon Teal 198 Northern Shoveler 35 Bufflehead 11 6 Ruddy Duck 2 American Coot 1 TOTAL 26 268 7 •Although coots are not in the waterfowl family (Anatidae), they were counted along with waterfowl. ber surveys were seen on the overflow pond (Table 6). About one-quarter of the Blue- winged Teal and Northern Shovelers diet and over three quarters of the Bufflehead's diet are aquatic insects and mollusks (Mar- tin et al. 1961). Importance of cliffs to nesting RAPTORS.— Cliffs are important as nesting sites for many of the raptors on the study area, because large trees are nearly nonex- istent. Eight species of large raptors (hawks, eagles, falcons, owls, and ravens) were ob- served during the June surveys (Table 7), and five of these raptors were found nesting on cliff ledges (Table 4). In addition, Ameri- can Kestrels were observed around cliffs, and probably used them as nesting sites. The American Kestrel is a cavity nester (Li- gon 1961), which would make its nest diffi- cult to locate by the helicopter survey used in this study. There was evidence of old Ferruginous Hawk nests on cliffs, although the only active nest found in this study was on the ground. Ligon (1961) reported that Prairie Falcons, Great Horned Owls, and Common Ravens prefer to nest on cliff ledges. He also observed that Red-tailed Hawks, Golden Eagles, and American Kes- trels normally nest on cliff ledges or in trees. Therefore, the absence of large trees on the study area, and the fact that most of the large raptor nests located on the study area were on cliff ledges suggests that six of the eight raptor species found on the study area are dependent on the cliffs for nesting. Although cliff ledges are extremely im- portant to many of the raptors nesting on the study area, they do not appear to be a major limiting factor to raptors in this area. Not only were 21 inactive cliff nests located (Tables 3 and 4), but also distance between existing nests was greater than for three other western study areas. Two nests each were located for the Red-tailed Hawk and Great Horned Owl in this study (Table 4). Distances between these nests were 8.1 km (5 miles) for the Red-tailed Hawk and 10.5 km (6.5 miles) for the Great Horned Owl. However, Craighead and Craighead (1969) found that the maximum diameter of nest- Dec. 1977 TOLLE: new MEXICO BIRDS 497 ing ranges for both the above raptors was 2.4 km (1.5 miles) at their semiwilderness study area near Moose, Wyoming. Also, Smith (1969) fomid that great horned owl nests averaged 1.6 km (1 mile) apart in the Great Basin desert of central western Utah. Maximum and minimum distances between nearest owl nests were 4.8 km (3 miles) and 1.2 km (0.75 miles). In contrast, Hensley (1954) studied the Red- tailed Hawk in the lower Sonoran desert of Arizona and found that the greatest distance between two nests was about 9.7 km (6 miles), while the clos- est distance was about 4.8 km (3 miles). It appears that nesting distances are greatest in the cold desert of this study area, least in the Wyoming wilderness area and Great Ba- sin desert of Utah, and intermediate in a warm desert habitat. Peak waterfowl migration periods.— No set patterns have been established when any one species can be counted on to occur in peak numbers in the San Juan River val- ley (J. L. Sands, pers. comm.). The peak population of each species migrating through San Juan County depends on weather conditions and brood population. However, some of the species observed dur- ing this study appeared to peak during the portion of the season considered normal for that species. Blue-winged Teal, Cinnamon Teal, and Northern Shovelers are considered Table 7. Nonwaterfowl birds observed during four migratory waterfowl surveys" on the study area. Frequency of Frequency of Observation'' Observation'' during during Survey Periods Survey Periods Common Name 1 2 3 4 Common Name .1 2 3 4 Turkey Vulture R Common Raven C u u u Cooper's Hawk R Rock Wren - R - Red-tailed Hawk R R R U Sage Thrasher - R - Swainson's Hawk - R - American Robin - R - Rough-legged Hawk - R - Blue-gray Gnatcatcher - U - Ferruginous Hawk R R U Ruby-crowned Kinglet - R - Golden Eagle - R R Loggerhead Shrike R R - U Marsh Hawk R R R Yellow-mmped Warbler - R - Prairie Falcon R R R R (Audubon's Var.) Merlin R - Wilson's Warbler - U - American Kestrel R R Western Meadowlark - - - R Scaled Quail C . Yellow-headed Blackbird - U - Killdeer - U Red-winged Blackbird R - - R Pectoral Sandpiper R - Cassin's Finch R American Avocet U R House Finch R R R R Wilson's Phalarope R - Green-tailed Towhee - U - Mourning Dove R Vesper Sparrow - - R - Great Homed Owl R - R Black-throated Sparrow R - - - Burrowing Owl R Sage Sparrow U C R U Say's Phoebe - R Dark-eyed Junco - U - - Western Wood Pewee R Clay-colored Sparrow - R - Homed Lark A A A C Brewer's Sparrow - - U - Bank Swallow R White-crowned Sparrow R - - R Barn Swallow - R Number of Species 17 12 32 15 'Surveys were made during the following four periods: (1) 13-16 November 1973, (2) 20-24 February 1974, (3) 22-25 September 1974. and (4) 11-14 No- vember 1974. "These birds were not censused. Letters indicate frequency of observation during waterfowl surveys. The approximate number of observations are in- dicated as follows: A = Abundant, 200-t-; C = Common, 30-199; U = Uncommon, 5-29; R = Rare. 1-4. 498 GREAT BASIN NATURALIST Vol. 37, No. 4 early fall migrants (Chattin 1964, Bellrose 1976). These ducks were relatively abundant during the September survey but were ab- sent during the November surveys (Table 6). The Mallard is hardy, and normally mi- grates southward late, when it is forced to move south by the fall freeze (Chattin 1964). Since the Mallard is a permanent resident in San Juan County (Hubbard 1970), the fall migration is usually apparent as a steady increase in numbers. In surveys on the study area, Mallards were relatively common only in November of 1973 but were absent in September (Table 6). Peak waterfowl use of stock ponds on the study area probably occurs during late Sep- tember due to the preference of early mi- grants for this habitat type. The normal peak of fall waterfowl migration through San Juan County is in mid-November (J. L. Sands, pers. comm.). However, early mi- grants hke Cinnamon Teal, Blue-winged Teal, and Northern Shovelers prefer to feed in shallow water, and thus make frequent use of small ponds (Sprunt and Zim 1961). On the other hand, ducks such as the Mal- lard constitute a major portion of the wa- terfowl migrants present along the San Juan River during November (J. L. Sands, pers. comm.). Because these birds often feed on waste grain (Bellrose 1976), they are at- tracted to the fields along the river and probably do not utilize desert ponds in San Juan County to the degree that teal and shovelers do. The effect on fall waterfowl migration is that the major wetland areas in San Juan County (along the San Juan River) are heavily used by late-migrating species, and desert ponds (like those on the study area) are primarily used by early-migrating species. Threatened or status undetermined SPECIES.— Four species of birds observed on the study area during the June breeding bird surveys deserve detailed discussion due to their federal designation as "threatened" (Prairie Falcon) or "status undetermined" (Ferruginous Hawk, Mountain Plover, and Burrowing Owl) (U.S. Department of the Interior 1973b). Two Prairie Falcons were observed in separate locations at the cliffs bordering the Chaco River. A Prairie Fal- con nest with eggs was located at one of these sites on a tall rock column (Tables 3 and 4). A pair of Ferruginous Hawks (plus their nest. Tables 3 and 4) and three Moun- tain Plovers (plus one nest) were located on open mesa habitat, where the vegetation varies from 0.6 m (2 feet) tall to nonexistent and the soil is sandy with some gravel (Tolle, 1976). Four Burrowing Owls were observed at three different Gunnison's prairie dog towns on the mine lease. In ad- dition, a Merlin (which is considered "status undetermined") was seen during migratory surveys in November 1973. The Ferruginous Hawk nest located on the study area was slightly less than 1.6 km (1 mile) from a Gunnison's prairie dog town. Because the prairie dog is one of this hawk's major food sources (Ligon 1961), the prairie dog town may have influenced the nest location. Acknowledgments Alan P. Nelson, Steven E. Pomeroy, and C. Greg Schmitt deserve thanks for their as- sistance with the field work. James L. Sands supplied information on waterfowl migra- tion in the San Juan River valley. Clayton M. White assisted with selection of raptor survey techniques, and both White and John P. Hubbard reviewed early drafts of the manuscript. Project funding was provided by WESCO (Western Coal Gasification Company) and UII (Utah International). Dec. 1977 tolle: new mexico birds Appendix I. Scientific nomenclature of birds observed on the study area. 499 Common Name" Scientific Name" Common Name" Scientific Name" Great Egret Mallard Gadwall Pintail Green-winged Teal Blue-winged Teal Cinnamon Teal Northern Shoveler Bufflehead Ruddy Duck Turkey Vulture Cooper's Hawk Red-tailed Hawk Swainson's Hawk Rough-legged Hawk Ferruginous Hawk Golden Eagle Marsh Hawk Prairie Falcon Merlin American Kestrel Scaled Quail American Coot Killdeer Mountain Plover Spotted Sandpiper Pectoral Sandpiper American Avocet Wilson's Phalarope Mourning Dove Great Homed Owl Burrowing Owl Common Nighthawk Western Kingbird Say's Phoebe Casmerodius alhus Anas platyrhynchos Anas strepera Anas acuta Anas crecca Anas discors Anas cyanoptera Anas clypeata Biicephala albeola Oxyiira jamaicensis Cathartes aura Accipiter cooperii Buteo jamaicensis Buteo swainsoni Buteo lagopus Buteo regalis Aquila chrysaetos Circus cyaneus Falco mexicanus Falco columbarius Falco sparverius Callipeph squamata Fulica americana Charadrius vociferus Charadrius montanus Actitis inacularia Calidris meUinotos Recurvirostra americana Steganopus tricolor Zenaida macroura Bubo virginianus Speotyto cunicularia Chordeiles minor Tyrannus verticalis Sayornis say a Ash-throated Flycatcher Western Wood Pewee Horned Lark Tree Swallow Bank Swallow Barn Swallow Cliff Swallow Common Raven Rock Wren Mockingbird Sage Thrasher American Robin Blue-gray Gnatcatcher Ruby-crowned Kinglet Loggerhead Shrike Yellow-rumped Warbler Wilson's Warbler Western Meadowlark Yellow-headed Blackbird Red-winged Blackbird Northern Oriole Brown-headed Cowbird Blue Grosbeak Cassin's Finch House Finch Green-tailed Towhee Vesper Sparrow Black-throated Sparrow Lark Sparrow Sage Sparrow Dark-eyed Junco Clay-colored Sparrow Brewer's Sparrow White-crowned Sparrow Myiarchus cinerascens Contopus sordidulus Eremophila alpestris Iridoprocne bicolor Riparia riparia Hirundo rustica Petrochelidon pyrrhonota Corvus corax Salpinctes obsoletus Mimus polyglottos Oreoscoptes montanus Turdus migratorius Polioptila caerulea Regulus calendula Lanitts ludovicianus Dendroica coronata Wilsonia pusilla Sturnella neglecta Xanthocephalus xanthocephalus Agelaius phoeniceus Icterus galbula Moloihrus ater Guiraca caerulea Carpodacus cassinii Carpodacus mexicanus Chlorura chlorura Fooecetes grarnineus Amphispiza bilineata Chondestes grammacus Amphispiza belli junco hyemalis Spizella pallida Spizella breweri Zonotrichia leucophrys "Common and scientific names are from the Fifth AOU Check-Hst (Wetmore, 1957) with from the 32d Supplement (Eisenmann, 1973). Literature Cited Bellrose, F. C. 197L Migration corridors of water- fowl in the United States. Prep, for Fed. Aviation Admin., Dept. of Trans., Washington. 18 pp. 1976. Ducks, geese, and swans of North Amer- ica. Stackpole Books, Harrisburg, Pa. 544 pp. Chattin, J. E. 1964. Pacific flyway. pp. 23.3-252. In: J. P. Linduska (ed.) Waterfowl Tomorrow. U.S. Dept. Int., Fish and Wild. Serv., Washington. 770 pp. Craighead, J. J., and F. C. Craighead, Jr. 1969. Hawks, owls, and wildlife. Dover Publ., Inc., New York. 443 pp. Edminster, F. C. 1964. Farm ponds and waterfowl, pp. 399-407. In: J. P. Linduska (ed.) Waterfowl Tomorrow. U.S. Dept. Int., Fish and Wild. Serv., Washington. 770 pp. Eisenmann, E. (Chairman). 1973. Thirty-second supple- ment to the American Ornithologists' Union check-list of North American birds. Auk 90:411-419. Fautin, R. W. 1946. Biotic communities of the north- em desert shrub biome in western Utah. Ecol. Monogr. 16(4): 251-310. Hayward, D. L. 1967. Birds of the upper Colorado River basin. Brigham Young University Sci. Bull., Biol. Ser. 9(2): 1-64. 500 GREAT BASIN NATURALIST Vol. 37, No. 4 Hensley, M. M. 1954. Ecological relations of the breeding bird population of the desert biome in Arizona. Ecol. Monogr. 24(2): 185-207. Hubbard, J. P., 1970. Checklist of the birds of New Mexico. New Mexico Ornithological Soc, Publ. No. 3. McLeod Printing Co., Albuquerque. 108 pp. HuEY, W. S. 1967. Migratory game birds, pp. 153-164. In: New Mexico Wildlife Management. New Mexico Dept. Game and Fish, Santa Fe. 250 pp. Kendeich, S. C. 1961. Animal ecology. Prentice-Hall, Inc., Englewood Cliffs, N.J. 468 pp. Leopold, A. S. 1969. The desert. Time-Life Books, New York. 191 pp. LiGON, J. S. 1961. New Mexico Birds: and Where to Find Them. Univ. of New Mexico Press, and New Mexico Dept. Game & Fish, Albuquerque. 360 pp. Martin, A. C, H. S. Zim, and A. L. Nelson. 1961. American wildlife and plants: a guide to wildlife food habits. Dover Publ., Inc., New York. 500 pp. Smith, D. G. 1969. Nesting ecology of the great horn- ed owl {Bubo virginianus). Brigham Young Univ. Sci. Bull., Biol. Series 10(4): 16-25. Smyth, M., and G. A. Bartholomew. 1966. The water economy of the Black-throated Sparrow and the Rock Wren. Condor 68: 447-458. Smyth, M., and H. N. Coulombe. 1971. Notes on the use of desert springs by birds in California. Con- dor 73: 240-243. Sprunt, a., IV, AND H. S. Zim. 1961. Gamebirds: a guide to North American species and their habits. Golden Press, New York. 160 pp. Tolle, D. a. 1976. A westward extension in the breeding range of the Mountain Plover. Wilson Bull. 88(2):358-359. U.S. Department of the Interior. 1973a. Cooperative breeding bird survey of North America. Mig. Non-game Bird Studies, U.S. Fish and Wild. Serv., Laurel, Maryland. 4 pp. Mimeogr. 197.3b. Threatened wildlife of the United States. Bureau of Sport Fisheries and Wildlife. Resour. Pub. 114. 289 pp. Wetmore, a. (Chairman). 1957. Check-list of North American birds, 5th ed. American Ornithologists' Union, 691 pp. White, C. M., and S. K. Sherhod. 1973. Advantages and disadvantages of the use of rotor-winged air- craft in raptor surveys. Raptor Research 7 (3/4): 97-104. A NEW MONOTYPIC GENUS OF CHIGGERS AND FOUR NEW SPECIES OF QUADRASETA FROM VENEZUELA (ACARI: TROMBICULIDAE)' M. L. Goff and J. M. Brennan' Abstract.— Curebureia, n. gen., and its type-species, C. johnstoni, n. sp., are described. The genus Quadraseta Brennan, 1970, is redefined and a key to the 11 included species given. Four new species, Quadraseta tachirensis, holotype ex Akodon sp., Q. rotstieri, ex Proechhnys guyannensis, Q. mirandae, holotype ex Oryzomys albigularis, and Q. fakonensis, ex SylviUigus floridanus, are described. Four new species of Quadraseta Brennan, 1970, and a new monotypic genus, all from Venezuela, are described and illustrated be- low. Hosts are small mammals. Collections were made under the direc- tion of Dr. Charles O. Handley, Smithsonian Institution, Dr. Vernon J. Tipton, Brigham Young University, and Dr. Conrad E. Yun- ker, Rocky Mountain Laboratory. Holotypes are in the collection of the U.S. National Museum of Natural History (currently housed at the Bishop Museum, Honolulu, Hawaii) and paratypes are there and in the collection of the Bishop Museum. All mea- surements are given in micrometers of the holotypes followed in parentheses by the means and extremes of the type series. Quadraseta Brennan Quadraseta Brennan, 1970: 1695; Brennan & Reed, 1975: 62. Type-species.— Euschoengastia pazca Brennan & Jones, 1964: 700. Diagnosis.— Trombiculine larvae parasitic on small mammals and, occasionally, birds; legs 7-segmented; coxae unisetose; 2-3 gen- ualae I, genuala II and III; tibiala III; sub- terminala and parasubterminala I; 6 branched setae in basal whorl of tarsus I. Scutum sparsely punctate; subrectangular, with sinuous anterior and posterior margins; 5 setae; sensilla clavate to capitate. Eyes 2/2. Cheliceral blades with tricuspid cap; palpal tarsus with 4B; palpal claw 3- pronged. Quadraseta tachirensis, n. sp. Fig. 1 Type data.— Holotype and 7 paratypes from Tachira, San Cristobal, 21-III-1968, ex Akodon sp. (RML 53653); 2 paratypes, same data except 15-III-1968, ex Akodon bogo- tensis (RML 55933). Diagnosis.— Palpal setal formula B/B/BBB/4B; palpal claw 3-pronged; ga- leala B; 3 genualae I; 54 dorsal body setae; 2 pairs of humeral setae; PL>AM>AL; PW/SD = 1.83. Description of species.— Larvae. Idiosoma: Measuring 290 x 235 in engorged specimen. Eyes 2/2, anterior larger, on ocular plate. Two pairs of humeral setae, measuring 44- 46; 54 dorsal body setae, measuring 32-40, arranged 6-10-14-10-6-6-2; 2 pairs of sternal setae, anterior 37-40, posterior 28-31; 42 preanal setae, 20-24; 20 postanal setae, 28- 33; total body setae 124. Gnatho.sorna: Pal- pal setal formula B/B/BBB/4B; palpal claw 3-pronged; galeala B; chela (27) with tricus- pid cap. Scutum: Sparsely punctate; ante- rior margin biconcave, posterior margin bi- convex; AM base in line with AL bases; SB anterior to PL bases; PL>AM>AL; sensilla capitate, head with setules; PW/SD = 'Studies upon which this paper is based were supported in part by Department of Army contract DA-49-193-MD-2788 with the Smithsonian In- stitution. 'Department of Entomology. Bishop Museum, P.O. Box 6037, Honolulu, Hawaii 96818, USA. 501 502 GREAT BASIN NATURALIST Vol. 37, No. 4 1.83. Scutal measurements: AW 47 (48, 47- 49); PW 66 (66, 62-69); SB 22 (19, 17-22); ASB 22 (22, 20-23); PSB 14 (14, 12-16); AP 25 (25, 24-27); AM 25 (29, 25-32); AL 23 (26, 23-30); PL 41 (45, 41-48); sens. 34 x 15 (35, 33-37 X 14-15). Legs: All 7-segmented terminating in a pair of claws and a claw- like empodium. IP = 670-710. Leg L 225- 238; coxa with 1 branched seta (IB); tro- chanter IB; basifemur IB; telofemur 5B; genu 4B, 3 genualae, microgenuala; tibia 8B, 2 tibialae, microtibiala; tarsus (56 x 21) 19B, tarsala (14-15), microtarsala, sub- terminala, parasubterminala, pretarsala. Leg U: 205-220; coxa IB; trochanter IB; basife- mur 2B; telofemur 4B; genu 3B, genuala; tibia 6B, 2 tibialae; tarsus (50 x 19) 14B, tarsala (12-14), microtarsala, pretarsala. Leg lU: 240-255; coxa IB; trochanter IB; basife- mur 2B; telofemur 3B; genu 3B, genuala; tibia 6B, tibiala; tarsus (65 x 16) 15B. Remarks.— Qitadraseta tachirensis is sim- ilar to Q. pazca (Brennan & Jones, 1964) in having galeala B and 3 genualae I, but differs in having eyes on a distinct ocular plate, PW/SD = 1.83 (1.51 in Q. pazca), and arrangement of body setae. Additional specimens examined (40).— Fig. 1. Larva of Quadraseta tachirensis: (a) scutum; (b) leg I distal 3 segments showing specialized setae (mea- surements in micrometers) and bases of branched setae; (c) leg II, as above; (d) leg III, as above. Dec. 1977 GOFF, BRENNAN: VENEZUELAN MITES 503 Guarico, 6.1.1968, 1 Zygodontomys sp. (7). Tachira, San Cristobal, 2-17-III-1968, 2 Akodon sp. (20), 1 Akodon bogotensis (2), 1 Onjzomys sp. (3), 1 Oryzomys albigularis (1), 2 Oryzomys minutus (6). Yaracuy, W of San Felipe, 8-XII-1967, 1 Oryzomys capito Quadraseta rotstieri, n. sp. Fig. 2 Type data.— Holotype and 8 paratypes from Falcon, E of Maracaibo, ex Proechimys guyannensis (RML 53271), 13-V-1968. Diagnosis.— Palpal setal formula B/B/BBB/4B; palpal claw .3-pronged; ga- leala N; 2 genualae I, genuala II and III; genu I with 5 branched setae; tibiala III; 58 dorsal body setae, arranged 8-8-10-6-10-4-6- 6; sensilla clavate; PW/SD = 1.90-1.97. Description of species.— Larvae. Idiosoma: Measuring 400 x .305 in engorged specimen. Eyes 2/2, of equal size, on ocular plate. Two pairs of humeral setae, measuring 46- 48; 58 dorsal body setae, measuring 32-41, arranged 8-8-10-6-10-4-6-6; 2 pairs of ster- nal setae, 30; 28 preanal setae, 20-26; 30 postanal setae, 30-35; total body setae 124. Fig. 2. Larva of Quadraseta rotstieri: (a) scutum; (b) leg I distal 3 segments showing specialized setae (measure- ments in micrometers) and bases of branched setae; (c) leg II, as above; (d) leg III, as above. 504 GREAT BASIN NATURALIST Vol. 37, No. 4 Gnathosoma: Palpal setal formula B/B/BBB/4B; palp'al claw 3-pronged; ga- leala N; chela (35-38) with tricuspid cap. Scutum: Sparsely punctate; anterior margin biconcave, posterior margin biconvex; AM base posterior to AL bases; SB anterior to PL bases; PL>AL>AM; sensilla clavate, head with setules; PW/SD = 1.90-1.97. Scutal measurements: AW 56 (56, 52-59); PW 73 (76, 73-80); SB 22 (24, 22-27); ASB 23 (24, 20-27); PSB 14 (16, 14-17); AP 28 (30, 28-35); AM 36 (36, 33-38); AL 38 (38, 34-40); PL 55 (52, 48-56); sens, missing in holotype (43 x 13, 42-45 x 13). Legs: All 7- segmented, terminating in a pair of claws and a clawlike empodium. Onychotrictes absent. IP = 690-717. Leg L 235-245; coxa IB; trochanter IB; basifemur IB; telofemur 5B; genu 5B, 2 genualae, microgenuala; tibia 8B, 2 tibialae, microtibiala; tarsus (57 x 20) 20B, tarsala (15), microtarsala, sub- terminala, parasubterminala, pretarsala. Leg U: 218-220; coxa IB; trochanter IB; basife- mur 2B; telofemur 4B; genu 3B, genuala tibia 6B, 2 tibialae; tarsus (51 x 19) 14B tarsala 15; microtarsala, pretarsala. Leg III 238-252; coxa IB; trochanter IB; basifemur 2B; telofemur 3B; genu 3B, genuala; tibia 6B, tibiala; tarsus (67 x 17) 15B. Remarks.— Quadraseta rotstieri is similar to Q. antillarum (Brennan, 1967) in having 2 genualae I and 4 humeral setae, but differs in having palpal tibia BBB (BNB in Q. antillarum), PL>AL, and 5 branched setae on genu I. Additional specimens examined (18).— Falcon, E of Maracaibo, 13-15-V-1968, 3 Akodon urichi (18). Quadraseta mirandae, n. sp. Fig. 3 Type data.— Holotype and 9 paratypes from Miranda, 1 km N of Caracas, Que- brada Chacaito, ex Oryzomys albigularis (RML 52518), 18-V-1967. Diagnosis.— Palpal setal formula B/B/BBB/4B; palpal claw 3-pronged; ga- leala N; 3 genualae I; 68 dorsal body setae; 2 pairs of humeral setae; PL>AM>AL; PW/SD = 1.80. Description of species.— Larvae. Idiosoma: Measuring 290 x 225 in engorged specimen. Eyes 2/2, anterior larger, on ocular plate. Two pairs of humeral setae, measuring 49- 55; 68 dorsal body setae, measuring 40-50, arranged 10-6-8-10-10-6-10-6-2; 2 pairs of sternal setae, 34-40; 38 preanal setae, 30-35; 22 postanal setae, 34-42; total body setae 136. Gnathosoma: Palpal setal formula B/B/BBB/4B; palpal claw 3-pronged; ga- leala N; chela (37-40) with tricuspid cap. Scutum: Sparsely punctate; anterior margin shallowly biconcave; posterior margin shal- lowly convex; AM base slightly posterior to AL bases; SB anterior to PL bases; PL>AM>AL; sensilla clavate, head with coarse setules; PW/SD = 1.80. Scutal mea- surements: AW 61 (62, 60-64); PW 80 (81, 73-90); SB 28 (28, 23-31); ASB 29 (28, 26- 29); PSB 17 (17, 14-18); AP 38 (37, 34-39); AM 44 (48, 44-52); AL 35 (35, 33-37); PL 55 (60, 55-64); sens. 55 x 13 (50, 47-55 x 12-14). Legs: All 7-segmented, terminating in a pair of claws and a clawlike em- podium. IP = 883-910. Leg L 300-312; coxa IB; trochanter IB; basifemur IB; te- lofemur 5B; genu 4B, 3 genualae, micro- genuala; tibia 8B, 2 tibialae, microtibiala; tarsus (75 x 22) 19B, tarsala (18-20), micro- tarsala, subterminala, parasubterminala, pre- tarsala. Leg U: 270-280; coxa IB; trochanter IB; basifemur 2B; telofemur 4B; genu 3B, genuala; tibia 6B, tibiala; tarsus (68 x 22) 16B, tarsala (18-20), microtarsala, pretarsala. Leg lU: 305-317; coxa IB; trochanter IB; basifemur 2B; telofemur 3B; genu 3B, gen- uala; tibia 6B, tibiala; tarsus (82 x 18) 15B. Remarks.— Quadraseta mirandae is similar to Q. flochi in having 3 genualae I, clavate sensillae and PL>AL, but differs in having tarsalae I and II longer (18-20, 14 and 15, respectively, in Q. flochi), posterior scutal margin evenly convex (biconvex in Q. flochi) and longer dorsal body setae (40-50 in Q. mirandae and 25-30 in Q. flochi). Additional specimens examined (134).— NW of Orania, 1 Proechimys guyannensis (1). BohVar, E of Caicara, 1 P. guyannensis (2). Falcon, E of Mirimire, 2 Heteromys anomalus (2), 3 Monodelphis brevicaudata (20), 1 Zygodontomys sp. (2). Lora, N of El Tocuyo, 1 Echymys sp. (1). Miranda, Ca- Dec. 1977 GOFF, BRENNAN: VENEZUELAN MITES 505 racas Country Club, 4 Didelphis marsupialis (36); 1 km N of Caracas, Quebrada Cha- caito, 1 H. anornahts (8), 6 Oryzomys al- bigiilaris (18). Sucre, E of Carupano, 1 H. anornalus (11), 1 M. hrevicaudata (5), 1 P. guyannensis (4), 1 Zygodontomys microtinus (1). Yaracuy, W of San Felipe, 2 M. hrevi- caudata (8). Zulia, Sierra de Perija, 1 P. guyannensis (2), 2 Proechimys sp. (3). Quadraseta falconensis, n. sp. Fig. 4 Type data — Holotype and 12 paratypes from Falcon, W of Coro, 17-VII-1968, ex Sylvikigus floridanus (RML 53360). Diagnosis.— Palpal setal formula B/B/BNB/4B; palpal claw 3-pronged; ga- leala N; 3 genualae I; 34 dorsal body setae; 2 pairs of humeral setae; AL>PL>AM; PW/SD = 1.84. Fig. 3. Larva of Quadraseta mirandae: (a) scutum; (b) leg I distal .3 segments showing specialized setae (mea- surements in micrometers) and bases of branched setae; (c) leg II, as above; (d) leg III, as above. 506 GREAT BASIN NATURALIST Vol. 37, No. 4 Description of species.— Larvae. Idiosoma: Measuring 430 x 300 in engorged specimen. Eyes 2/2, anterior larger, on ocular plate. Two pairs of humeral setae, measuring 25- 28; 34 dorsal body setae, measuring 19-26, arranged 8-10-8-6-2; 2 pairs of sternal setae, 25-27; 18 preanal setae, 15-18; 18 postanal setae, 21-23; total body setae 78. Gnatho- sorna: Palpal setal formula B/B/BNB/4B; palpal claw 3-pronged; galeala N; chela (25) with tricuspid cap. Scutum: Sparsely punc- tate; anterior margin deeply biconcave, pos- terior margin biconvex; AM base in line with or slightly posterior to AL bases; SB anterior to PL bases; AL>PL>AM; sensilla capitate, head with fine setules; PW/SD = 1.84. Scutal measurements: AW 44 (41, 37- 44); PW 61 (57, 52-61); SB 12 (12, 10-13); ASB 22 (22, 20-25); PSB 9 (9, 8-10); AP 26 (25, 22-27); AM 20 (19, 18-20); AL 37 (36, 34-38); PL 32 (31, 28-32); sens, missing in holotype (24, 21-26, head 15 x 16). Legs: All 7-segmented, terminating in a pair of claws and a clawlike empodium. IP = 540- 560. Leg I: 190-195; coxa IB; trochanter IB; basifemur IB; telofemur 5B; genu 4B, 3 Fig. 4. Larva of Quadraseta falconensis: (a) scutum; (b) leg I distal .3 segments showing specialized setae (mea- surements in micrometers) and bases of branched setae; (c) leg II, as above; (d) leg III, as above. Dec. 1977 GOFF, BRENNAN: VENEZUELAN MITES 507 genualae, microgenuala; tibia 8B, 2 tibialae, microtibiala; tarsus (43 x 17) 19B, tarsala (10-11), microtarsala, subterminala, para- subterminala, pretarsala. Leg II: 160; coxa IB, trochanter IB; basifemur 2B; telofemur 4B; genu 3B, genuala; tibia 6B, 2 tibialae; tarsus (37 x 15) 14B, tarsala (13-14), micro- tarsala, pretarsala. Leg III: 190-205; coxa IB; trochanter IB; basitemur 2B; telofemur 3B; genu 3B, genuala; tibia 6B, tibiala; tar- sus (55 X 12) 14B. Remarks.— Quadraseta falconensh shares the character of a nude lateral palpotibial seta with Q. antillamm (Brennan, 1967) but differs in having 3 genualae I. Key to Species of Quadraseta 1. Two genualae I 2 Three genualae I 5 2(1). Palpal setal formula B/B/BNB antiUanim (Brennan, 1967) Palpal setal formula B/B/BBB 3 3(2). Galeala branched azulae (Brennan & Jones, 1964) Galeala nude 4 4(3). Genu I with 5 branched setae rotstieri, n. sp. Genu I with 4 branched setae trapezoides (Brennan & Jones, 1964) 5(1). Galeala branched 6 Galeala nude 7 6(5). PW/SD>1.75 tachirensis, n. sp. PW/SD<1.55 pazca (Brennan & Jones, 1964) 7(5). Two humeral setae mackenziei (Yunker & Brennan, 1964) Four humeral setae 8 8(7). Sensilla clavate; more than 60 dorsal body setae 9 Sensilla capitate, fewer than 50 dorsal body setae 10 9(8). Tarsala I 18-20; dorsal body setae measuring 40-50 mirandae, n. sp. Tarsala I 14; dorsal body setae measuring 25-30 flochi (Brennan & Jones, 1960) 10(9). Lateral palpotibial seta nude falconensis, n. sp. Lateral palpotibial seta branched macarenae (Brennan & Jones, 1964) Carebareia, n. gen. Type-species.— Carebareia johnstoni Goff & Brennan, n. sp. Diagnosis.— Trombiculine larvae. Palpal tarsus 4B; palpal subterminala absent; pal- pal claw 3-pronged; galeala N; chela with tricuspid cap; sensilla capitate; genuala I, genuala II and III absent; tibiala III; sub- terminala and parasubterminala I; no masti- setae on leg III. Remarks.— Carebareia is similar to Quad- raseta Brennan, 1970, but may be separated by having only 1 genuala I (2-3 in Quad- raseta), lacking genuala II and III and hav- ing slightly expanded dorsal body setae. Single genuala I and absence of genuala II and III also serve to separate Carebareia from Aitkenius Brennan, 1970, Ectonyx Brennan, 1960, Cheladonta Lipovsky, Crossley & Loomis, 1955, and Trombe- wingia Fonseca, 1955. Carebareia is similar to Kayella Vercammen-Grandjean, 1960, in having 1 genuala I and lacking genuala II and III, but differs in having both sub- terminala and parasubterminala I. Carebareia phnstoni, n. sp. Fig. 5 Type data.— Holotype and 5 paratypes from Venezuela, Falcon, W of Coro, ex Marmosa robinsoni (RML 53544), 25.VII.1968. Diagnosis.— Palpal setal formula B/B/BBB/4B; palpal claw 3-pronged; ga- leala N; chela with tricuspid cap; eyes 2/2; 1 genuala I; tibiala III; subterminala and 508 GREAT BASIN NATURALIST Vol. 37, No. 4 parasubterminala I; 54 slightly expanded dorsal body setae; capitate sensilla; PW/SD = 2.10-2.30. Description of species.— Larvae. Idiosoma: Measuring 315 x 220 in partially engorged specimen. Eyes 2/2, anterior larger, on ocu- lar plate. Two pairs of humeral setae, slightly expanded, measuring 26-29; 54 slightly expanded dorsal body setae, measur- ing 20-22, arranged 8-8-8-6-8-6-6-4; 2 pairs of sternal setae, 28-30; 22 preanal setae, 20- 22; 20 postanal setae, similar in form to dorsal body setae, 23; total body setae 104. Gnathosoma: Palpal setal formula B/B/BBB/4B; palpal claw 3-pronged; chela (25) with tricuspid cap; galeala N; gnatho- base lightly pimctate, bearing a pair of branched setae. Scutum: Sparsely punctate; anterior margin shallowly biconcave, poste- rior margin biconvex; AM base in line with AL bases; SB slightly anterior to PL bases; PL>AM>AL; sensilla capitate, head with Fig. 5. Larva of Carebareia johnstoni: (a) scutum; (b) leg I distal 3 segments showing specialized setae (mea- surements in micrometers) and bases of branched setae; (c) leg II, as above; (d) leg III, as above. Dec. 1977 GOFF, BRENNAN: VENEZUELAN MITES 509 coarse setules; PW/SD = 2.10-2.30. Scutal measurements: AW 45 (45, 40-50); PW 55 (59, 55-60); SB 10 (15, 10-16); ASB 16 (17, 16-20); PSB 10 (11, 10-12); AP 18 (20, 18- 23); AM 26 (27, 26-29); AL 23 (22, 20-24); PL 30 (33, 30-35); sens. 23 x 15 (26 x 15, 23-28 X 15). Legs: All 7-segmented, termi- nating in a pair of claws and a clawlike empodium. Onychotrictes absent. IP = 559- 604. Leg L 192-204; coxa with 1 branched seta (IB); trochanter IB; basifemur IB; te- lofemur 5B; genu 4B, genuala, micro- genuala; tibia 8B, 2 tibialae, microtibiala; tarsus (47 x 18) 15B; tarsala (10-11), micro- tarsala, subterminala, parasubterminala, pre- tarsala. Leg U: 170-190; coxa IB; trochanter IB; basifemur 2B; telofemur 4B; genu 3B; tibia 6B, 2 tibialae; tarsus (38 x l6) 15B, tarsala (13), microtarsala, pretarsala. Leg IIL 197-210; coxa IB; trochanter IB; basifemur 2B; telofemur 3B; genu 3B; tibia 6B, tibiala; tarsus (52 x 14) 14B. Remarks.— This species is named in honor of Dr. Donald E. Johnston, Acarology Labo- ratory, Ohio State University, in recognition of his many significant contributions to acarology. Literature Cited Brennan, J. M. 1970. Two new genera and a new spe- cies of Neotropical chiggers (Acarina: Trombicu- lidae). Ann. Entomol. Soc. Am. 63: 1694-1696. Brennan, J. M. and J. T. Reed. 1975. A list of Vene- zuela chiggers, particularly of small mammalian hosts (Acarina: Trombiculidae). Brigham Young Univ. Sci. Bull., Biol. Series 20(1-2): 45-75. A NEW NAME FOR GALIUM SCABRIUSCULUM (RUBIACEAE) OF UTAH Lauramay T. Dempster' and Friedrich Ehrendorfer- Abstract.— A new name, Galium emeryense, is proposed to substitute for the preempted name G. scabriuscu- lum. The name Galium scabriusculum (Eh- rend.) Damp. & Ehrend., used in 1965 for a Utah species of the G. niultiflorum complex, was discovered by us to have been preemp- ted, and therefore invahd. The name was pubhshed first as a subspecies of G. hypotri- chium by Ehrendorfer, later as a variety of G. coloradoense by Dempster, and eventu- ally as a species in our joint publication. We have since become aware that a Ga- lium scabriusculum was published in 1912: Galium scabriusculum (H. Braun) Dalla Torre & Samth., Fl. Tirol 6(3): 378 (1912) = G. nitidulum /S scabriusculum H. Braun ap. Oborny, Fl. Mahren Schlesien 1: 737 (1885!). This is now a synonym of G. pumi- lum Murr., but was validly published as a species. It is therefore necessary to give a new name to the Utah species. The name we propose refers to Emery Co., Utah, which contains the type locality. Full syn- onymy follows: Galium emeryense Demp. & Ehrend. nom. nov. G. hypotrichium ssp. scabriusculum Ehrend., Contr. Dudley Herb. 5: 13. 1956. G. coloradoense var. scabriuscu- lum Demp., Brittonia 11: 120. 1959. G. scabriusculum Demp. & Ehrend., Brit- tonia 17: 312. 1965. Type: Calf Springs Wash, San Rafael Swell, Emery Co., Utah, Maguire 18437 (GH. Isotypes CAS, NY, UC, UTC, WTU). Galium emeryense ssp. protoscabriusculum Demp. & Ehrend. comb. nov. G. sca- briusculum ssp. protoscabriusculum Demp. & Ehrend., Brittonia 17: 312. 1965. Type: side canyon ca 1 mile NW of Castle Gate, Carbon Co., Utah, Eh- rendorfer ir Stutz 5954 (UC). 'Jepson Herbarium, Department of Botany, University of California, Berkeley. 'Botanical Institute and Botanic Garden of the University of Vienna, Austria. 510 NEW SYNONYMY AND NEW SPECIES OF AMERICAN BARK BEETLES (COLEOPTERA: SCOLYTIDAE), PART VI' Stephen L. Wood' Abstract— . New synonymy of American Scolytidae is proposed as follows: Corthylus Erichson ( = Pseudocor- thylus Ferrari), Hylesinus Fabricius { = Leperisinus Reitter), Monarthrum Kirsch {=Anchonocerus Eichhoff), Cor- thylus spinifer Schwarz { = Metacorthylus af finis Fonseca, Corthylus affinis Fonseca), Cryphalus ruficollis Hopkins { = Cryphalus amabilis Chamberlin, Cryphalus coioradensis Wood), Gnathotrichus retusus (LeConte) ( = Gnathot- richus alni Blackman), Gnathotrichus sulcatus (LeConte) { = Gnathotrichus aciculatus Blackman), Hypothenemus eruditus Westwood ( = Hypothenemus germari Eichhoff, Stephanoderes myrmedon Eichhoff, Stephanoderes inter- setosus Eggers), Hypothenemus seriatus Eichhoff ( = Stephanoderes nitidulus Hopkins, Stephanoderes subopacicollis Hopkins), Hypothenemus obscurus (Fabricius) { = Stephanoderes asperulus Eichhoff, Stephanoderes cassiae Eich- hoff), Pityophthorus annectens LeConte { = Pityophthorus citus Blackman), Pityophthorus balsameus Blackman ( = Pityophthorus patchi Blackman), Pityophthorus briscoei Blackman ( = Pityophthorus mundus Blackman), Pi- tyophthorus cariniceps LeConte { = Pityophthorus cognatus Blackman), Pityophthorus confertus Swaine { = Pityoph- thorus agnatus Blackman, Pityophthorus comptus Blackman), Pityophthorus confusus Blandford ( = Pityophthorus bellus Blackman), Pityophthorus consimilis LeConte { = Pityophthorus nudus Swaine), Pityophthorus fuscus Black- man { = Pityophthorus smithi Schedl), Pityophthorus immanis Blackman { = Pityophthorus sulcatus Bright), Pityoph- thorus murrayanae Blackman { = Pityophthorus gracilis Swaine, Pityophthorus cutleri Swaine, Pityophthorus exilis Swaine, Pityophthorus depygis Blackman, Pityophthorus watsoni Schedl, Pityophthorus aurulentis Bright), Pityoph- thorus ornatus Blackman { = Pityophthorus limatus Wood), Pityophthorus pseudotsugae Swaine { = Pityophthorus thatcheri Bright), Pityophthorus pullus Zimmermann { = Pityophthorus bisulcatus Eichhoff), Scolytodes levis (Black- man) { = Ctenophorus laevigatus Chapuis, Scolytodes chapuisi Wood). The following species are named as new to science: Pityophthorus crotonis (Venezuela), Scolytodes constrictus, S. festus, S. praeceps (Colombia), S. canal- iculus, S. contractus, S. jucundus, S. opacus, S. opimus, S. serenus, S. suturalis, and S. varius (Venezuela). On the following pages several newly dis- covered cases of synonymy and 12 species new to science are presented for American Scolytidae. The species new to science rep- resent the genera Scolytodes (11) and Pi- tyophthorus (1) and were taken in Colombia (3) and Venezuela (9). New Synonymy Corthylus Erichson Corthylus Erichson, 1836, Archiv. Naturgesch. 2(1):64 (Type-species: Bostrichus compressicornis Fabri- cius, subsequent designation by Ferrari, 1867, Die Forst- und Baumsuchtschadlichen Borkenafer, p. 48) Pseudocorthylus Ferrari, 1867, Die Forst- und Baumsuchtschadlichen Borkenkafer, p. 59 (Type- species: Pseudocorthylus letzneri Ferrari, sub- sequent designation by Hopkins, 1914, Proc. U.S. Nat. Mus. 48:128). New synonymy The name Pseudocorthylus has appeared in the literature for more than a century in either an uncertain status or as a subgenus of Corthylus Erichson. The types of the type-species of both genera were examined. Pseudocorthylus letzneri is a normal repre- sentative of the genus Corthylus. Con- sequently, Pseudocorthylus must be placed in synonymy under the older name as in- dicated above. Hylesinus Fabricius Hylesinus Fabricius, 1801, Syst. Eleuth. 2:390 (Type- species: Hylesinus crenatus Fabricius, subsequent designation by Westwood, 1838, Synopsis of the genera of British insects, p. 39) 'Part of the research was sponsored by the National Science Foundation. 'Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. 5i; 512 GREAT BASIN NATURALIST Vol. 37, No. 4 Leperisinus Reitter, 1913, Wiener Ent. Zeit. 32(Bei- heft):41 (Type-species: Bostrichus fraxini Pan- zer = Bostrichus varius Fabricius, subsequent des- ignation by Swaine, 1918, Dom. Canada Dept. Agric. Ent. Br. Tech. Bull. 14(2):70). New synony- my The genus Hylesinus Fabricius contains a diverse group of species from Europe to Australia, with a few species in north Africa and North America. Leperisinus was erected for a group of species, found mostly in Fra- xinus, that differed from Hylesinus crenatus Fabricius by the conspicuously scalelike ves- titure. After examining most of the species of Hylesinus from Europe and Asia and many others from Australia, Indonesia, and the Philippines, it became apparent that the scalelike or hairlike vestiture intergrades to such an extent that it is not a meaningful generic character. If Hylesinus were to be divided, the division should come elsewhere in the genus. In the absence of other gener- ic characters, those species previously as- signed to Leperisinus must be grouped with H. crenatus in the genus Hylesinus. Thus, Leperisinus must be placed in synonymy. Monarthrum Kirsch Monarthrum Kirsch, 1866, Berliner Ent. Zeitschr. 10:213 (Type-species: Monarthrum chapuisi Kirsch, monobasic) Anchonocerus Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:431 (Type-species: Anchonocerus ru- fipes Eichhoff, monobasic). New synonymy The holotypes of Monarthrum chapuisi Kirsch and Anchonocerus rufipes Eichhoff, a long series of the latter species, and more than 70 other species in this genus were ex- amined. It is quite clear that rufipes falls well within the generic limits of Mon- arthrum. For this reason the name Anchono- cerus must be placed in synonymy. Corthylus spinifer Schwarz Corthylus spinifer Schwarz, 1891, Proc. Ent. Soc. Washington 2:114 (Syntypes, females; Key West, Florida; U.S. Nat. Mus.) Metacorthylus affinis Fonseca, 1925, Commissao de Es- tudio e Debella^ao da Praga Cafeeira, Pub. 12:3 (Syntypes; Itatiba, Sao Paulo, Brazil; not lo- cated?). New synonymy Corthylus affinis Fonseca, 1927, Rev. Mus. Paul. 15(1):585 (Syntypes; Itatiba, Sao Paulo, Brazil; not located?). New synonymy Corthylus affinis Fonseca, 1927 is an ob- vious duplication of the validation of Meta- corthylus affinis Fonseca, 1925. Four pre- sumed syntypes of Fonseca's species and numerous other specimens from Brazil clearly match the excellent illustrations of this species that accompany the original de- scription. Part of this Brazilian material was compared to the syntypes of Corthylus spin- ifer Schwarz and other material from Flor- ida, Central America, and northern South America. All represent one species. It is very probable that this species reached Florida through commerce from tropical America. Cryphalus ruficollis Hopkins Cryphalus ruficollis Hopkins, 1915, U.S. Dept. Agric. Rept. 99:40 (Holotype, female; Alta, Utah; U.S. Nat. Mus.) Cryphalus amahilis Chamberlin, 1917, Canadian Ent. 49:321 (Lectotype, female; Elk Lake, Oregon; Ca- nadian Nat. Coll., designated by Bright, 1967, Canadian Ent. 99:681). New synonymy Cryphalus ruficollis coloradensis Wood, 1954, Univ. Kansas Sci. Bull. 36(2): 1008 Holotype, female; 7 miles N Grand Canyon Nat. Pk., Arizona; Snow Coll., Univ. Kansas). New synonymy The type series of Cryphalus ruficollis Hopkins, C. amahilis Chamberlin, and C. ruficollis coloradensis and more than 300 other specimens were examined and com- pared. The supposed geographical character gaps observed by me in 1954 have been completely eliminated by subsequent collec- ting. Therefore, it is necessary to place amahilis and coloradensis in synonymy un- der the senior name as indicated above. Gnathotrichus retusus (LeConte) Cryphalus retusus LeConte, 1868, Trans. Amer. Ent. Soc. 2:155 (Syntypes; California; Mus. Comp. Zool.) Gnathotrichus alni Blackman, 1931, J. Washington Acad. Sci. 21:271 (Holotype, female; Hoquiam, Washington; U.S. Nat. Mus.). New synonymy Gnathotrichus alni Blackman was named from a population that breeds in Alnus in Washington and Oregon. Except for the host, I have been unable to find characters that distingui.sh it from Gnathotrichus re- tusus (LeConte), a common species in con- Dec. 1977 wood: AMERICAN BARK BEETLES 513 iferous hosts in that area. Until biological characters can be found that distinguish alni from retiisus, it should be placed in synony- my as indicated above. Gnathotrichus sulcatiis (LeConte) Cryphalus stikatus LeConte 1868, Trans. Amer. Ent. Soc. 2:155 (Holotype, female; middle California; Mus. Comp. Zool.) Gnathotrichus aciculatus Blackman, 1931, J. Washing- ton Acad. Sci. 21:272 (Holotype, female; Cloud- croft, New Mexico; U.S. Nat. Mus.). Neiv synony- my Gnathotrichus aciculatus Blackman was named from specimens having a deeply, narrowly sulcate elytral declivity. After ex- amining 594 specimens of G. sulcatus (Le- Conte) from throughout its range, it was concluded that the aciculatus specimens represent an aberrant form of the species that intergrades completely with normal sulcatus specimens. Until biological or other characters can be found that justify recogni- tion of this form, it should be placed in syn- onymy as indicated above. The type series of both were examined. Hypothenernus eruditus Westwood Hypothenemus eruditus Westwood, 1836, Trans. Ent. Soc. London 1(1):34 (Syntypes; England?; some in British Mus. Nat. Hist.) Stephanoderes gennari Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:159 (Syntypes?; Mexico; lost with Hamburg Mus.). New synonymy Stephanoderes myrmedon Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:159 Holotype, female; Colom- bia; Inst. Roy. Sci. Nat. Belgique, Brussels). New synonymy Stephanoderes intersetosus Eggers, 1928, Archiv. Inst. Biol. Def. Agric. Anim. 1:85 (Lectotype, female; Sao Paulo, Brazil; U.S. Nat. Mus., designated by Anderson and Anderson, 1971, Smithsonian Con- trib. Zool. 94:16). New synonymy The female holotype of Stephanoderes myrmedon Eichhoff and the female lecto- type of Stephanoderes intersetosus Eggers were examined and compared directly to part of my series that was compared to syn- types of Hypothenemus eruditus Westwood. It is quite obvious that only one species is represented by this material. The type series of Stephanoderes germari Eichhoff was lost with the Hamburg Museum during World War II; consequently, the only means of fix- ing the identity of this species, other than from the generalized description, is through the examination of specimens compared to the types before they were lost. Five such series were examined and all are conspecific with eruditus. For this reason, it was placed in synonymy as indicated above. Hypothenemus seriatus (Eichhoff) Stephanoderes seriatus Eichhoff, 1872, Berliner Ent. Zeitschr. 15:133 (Lectotype, female; New Orleans, Louisiana; Inst. Roy. Sci. Nat. Belgique, Brussels, designated by Wood, 1973, Great Basin Nat. 33:177) Stephanoderes nitiduhts Hopkins, 1915, U.S. Dept. Agr- ic. Rept. 99:29 (Holotype, female; Cayamas, Cuba; U.S. Nat. Mus.). New synonymy Stephanoderes subopacicollus Hopkins, 1915, U.S. Dept. Agric. Rept. 99:.30 (Holotype, female; Cayamas, Cuba; U.S. Nat. Mus.). New synonymy The female holotypes of Stephanoderes nitiduhts Hopkins and S. subopacicollis Hopkins were examined and compared to my material that was previously compared by me to the female lectotype of Hypoth- enemus seriatus (Eichhoff). All of these specimens fall well within the range of vari- ability of seriatus. Hypothenemus obscurus (Fabricius) Hylesinus obscurus Fabricius, 1801, Syst. Eleuth. 2:395 (Lectotype, female; Essequibo, British Guiana, published as America meridionalis; Copenhagen Mus.) Stephanoderes asperulus Eichhoff, 1872 (nee. LeConte, 1868), Berliner Ent. Zeitschr. 15:133 (Lectotype, female; Cassia, probably from northern South America; Inst. Roy. Sci. Nat., Bnissels, present designation). New synonymy Stephanoderes cassiae Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2)8:152 (Replacement name for S. as- peruhts Eichhoff). New synonymy Three female syntypes of Stephanoderes asperulus Eichhoff are in the Chapuis Col- lection at the Brussels Museum. The first and third .specimens were compared directly by me to my homotypes of Hypothenemus obscurus (Fabricius) and are conspecific. The second specimen is of Hypothenemus seriatus Eichhoff. I here designate the third syntype, which is in the best condition, as the lectotype of asperulus Eichhoff. This ac- 514 GREAT BASIN NATURALIST Vol. 37, No. 4 tion places Eichhoffs name in synonymy under obscurus (Fabricius), as indicated above. Pityophthorus annectens LeConte Pityophthorus annectens LeConte, 1878, Proc. Amer. Philos. Soc. 17:622 (Lectotype, female; Tampa, Florida; Mus. Comp. Zool., designated by Bright, 1976, Coleopt. Bull. 30:185) Pityophthorus citits Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:137 (Holotype, female; Chiricahua Mts., Arizona; U.S. Nat. Mus.). New synonymy LeConte's series of syntypes of Pityoph- thorus annectens and 148 other specimens from the southeastern United States, Black- man's type series of P. citus from Arizona, 16 specimens from near Durango, Durango, Mexico (and several other Mexican speci- mens), and 22 specimens from Honduras were examined and compared directly. Af- ter much study and hesitation, it was con- cluded that all represent the same species. Consequently, the name citus must be placed in synonymy as indicated above. Pityophthorus balsameus Blackman Pityophthorus balsameus Blackman, 1922, New York St. Coll. For., Syracuse, Tech. Pub. 16:119 (Holo- type, female; Orono, Maine; U.S. Nat. Mus.) Pityophthorus patchi Blackman, 1922, New York St. Coll. For., Syracuse, Tech. Pub. 16:120 (Holotype, female; Orono, Maine; U.S. Nat. Mus.). Netv syn- onymy The female holotypes of Pityophthorus balsameus Blackman and P. patchi Black- man and 108 other specimens were exam- ined and compared. The female frons is var- iable and intergrades completely between the two named forms. Since patchi repre- sents only a minor variant, it is placed in synonymy as indicated above. Pityophthorus briscoei Blackman Pityophthorus briscoei Blackman, 1922, New York St. Coll. For., Syracuse, Tech. Pub. 16:123 (Holotype, female; Bnmswick, Maine; U.S. Nat. Mus.) Pityophthorus mundus Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:86 (Holotype, female; Littleton, New Hampshire; U.S. Nat. Mus.). New synonymy The female holotypes of Pityophthorus briscoei Blackman and P. mundus Blackman and 145 other specimens of this species were examined. The female frons is unusu- ally variable and includes several forms, with all stages of intermediacy, in addition to those that have been named. Since 77iun- dus represents no more than a form that is well within the limits of variability for this species, it must be placed in synonymy as indicated above. Pitijophthorus cariniceps LeConte Pityophthorus cariniceps LeConte, 1876, Proc. Amer. Philos. Soc. 15:353 (Holotype, female; Detroit, Michigan; Mus. Comp. Zool.) Pityophthorus cognatus Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:69 (Holotype, female; Davidson's R., North Carolina; U.S. Nat. Mus.). New synonymy Blackman's series of Pityophthorus cog- natus was composite. When two series of P. crassus Blackman, bearing only collection numbers on their labels, were removed from consideration, it became apparent that the male declivity was exactly the same as one variation found in male P. cariniceps Le- Conte. While the female frons of cognatus is usually radically different from cariniceps, a few specimens in a very long series form New Brvinswick were as in cognatus, while others exhibited various stages of inter- mediacy or one of two forms common in cariniceps. In view of the extreme varia- bility in the female frons and male declivity in this species and in the allied species of this group, it appears advisable to place cognatus in synonymy as indicated above. Pityophthorus confertus Swaine Pityophthorus confertus Swaine, 1917, Dom. Canada Dept. Agric. Ent. Br. Tech. Bull. 14(1):27 (Lecto- type, female; Adams Lake, British Columbia; Ca- nadian Nat. Coll., designated by Bright, 1967, Canadian Ent. 99:678) Pityophthorus agnattts Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:125 (Holotype, female; Cloudcroft, New Mexico; U.S. Nat. Mus.). Neiv synonymy Pityophthorus comptus Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:127 (Holotype, female; Santa Catalina Mts., Arizona; U.S. Nat. Mus.). New synonymy Dec. 1977 wood: AMERICAN BARK BEETLES 515 The female holotypes of Pityophthorus confertus Swaine, P. agnatus Blackman, and P. comptus Blackman and more than 3,000 other specimens of this species were exam- ined. Very slight differences in the southern areas of the distribution were noted, repre- sented by agnatus and co7nptus, but these are not sufficiently distinct to warrant sub- specific recognition. For this reason they were placed in synonymy as indicated above. Pityophthorus confusus Blandford Pityophthorus fuscus Blackman Pityojihthorus fuscus Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:32 (Holotype, female; Glacier National Park, Montana; U.S. Nat. Mus.) Pityophthorus smithi Schedl, 1931, Canadian Ent. 63:163 (Holotype, female; Copper Mountain, Brit- ish Columbia; Canadian Nat. Coll.). New synony- my The female holotypes of Pityophthorus fuscus Blackman and P. smithi Schedl and 16 other specimens were examined and compared directly to one another. Only one rare species is represented by this material. Pityophthorus confusus Blandford, 1904, Biol. Centr. Amer., Coleopt. 4(6):237 (Lectotype, female; San Geronimo, Guatemala; British Mus. Nat. Hist., designated by Bright, 1976, Coleopt. Bull. 30:184) Pityophthorus beUus Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:123 (Holotype, female; Monongalia Co., West Virginia; U.S. Nat. Mus.). New synonymy Blandford's syn types of Pityophthorus con- fusus and 62 other specimens from Mexico, Guatemala, Honduras, and northwestern Nicaragua were examined and compared to the holotype and 306 specimens of P. beUus Blackman from the southern United States. They obviously represent the same dis- tinctive species in which I see no basis for recognizing geographical races. Pityophthorus immanis Blackman Pitijophthorus immanis Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:98 (Holotype, female; Chiricahua Mts., Arizona; U.S. Nat. Mus.) Pityophthorus sulcatus Bright, 1977, Canadian Ent. 109:528 (Holotype, female; Mt. Lemon, Pima Co., Arizona; Canadian Nat. Coll.). New synonymy The female holotype and seven paratypes of Pityophthorus immanis Blackman from the Chiricahua Mts. (one paratype from Flagstaff, Arizona is of P. crassus Blackman) and the type series of 86 specimens of P. sulcatus Bright were examined. Following a careful study of this material it was con- cluded that only one species is represented. It is closely allied to P. abiegnis Wood. Pityophthorus consimilis LeConte Pityophthorus consimilis LeConte, 1878, Proc. Amer. Philos. Soc. 17:622 (Lectotype, male; Marquette, Michigan; Mus. Comp. Zool., designated by Bright, 1976, Coleopt. Bull. 30:185) Pityophthorus nudus Swaine, 1917, Dom. Canada Dept. Agric. Ent. Br. Tech. Bull. 14(1):30 (Lecto- type, male; Ste. Anne de Bellvue, Quebec; Cana- dian Nat. Coll., designated by Bright, 1967, Ca- nadian Ent. 99:678). New synonymy The male lectotypes of Pityophthorus con- similis LeConte and P. nudus Swaine and 382 other specimens were examined. These names were based on the presence or ab- sence of frontal vestiture in the female. Since the character is variable and complete intergradation in series is common, nudus must be placed in synonymy as indicated above. Pityophthorus murrayanae Blackman Pityophthorus murrayanae Blackman, 1922, New York St. Coll. For., Syracuse, Tech. Pub. 16:138 (Holo- type, female; Grand Lake, Colorado; U.S. Nat. Mus.) Pityophthorus gracilis Swaine, 1925, Canadian Ent. 57:195 (Holotype, female; Grant Co., Oregon; Candian Nat. Coll.). New synonymy Pityophthorus cutleri Swaine, 1925, Canadian Ent. 57:195 (Holotype, female; Merritt, Midday Valley, British Columbia; Candian Nat. Coll.). New syn- onymy Pityophthorus exilis Swaine, 1925, Canadian Ent. 57:196 (Holotype, female; Ochoco N. F., Oregon; Canadian Nat. Coll.). New synonymy Pitijophthorus depygis Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:128 (Holotype, female; Clyde, Colorado; U.S. Nat. Mus.). New synonymy Pityophthorus watsoni Schedl, 1930, Canadian Ent. 62:197 (Holotype, female; Nictor Lake, New Brunswick; Canadian Nat. Coll.). New synonymy 516 GREAT BASIN NATURALIST Vol. 37, No. 4 Pityophthorus aurulentus Bright, 1966, Pan Pacific Ent. 42:301 (Holotype, female; Shell ridge at Walnut Creek, Contra Costa Co., California; Cal- ifornia Acad. Sci.). New synonymy The female frons of this species varies from convex to extensively flattened, with punctures moderately coarse to minute, and with or without vestiture. The vestiture, when present, may be minute to very long. Any combination of these variations may be found in pure culture in a local area or all may occur there. Following the study of the holotypes and type series of Pityophthorus murrayanae Blackman, P. gracilis Swaine, P. cutleri Swaine, P. exilis Swaine, P. depygis Blackman, P. watsoni Schedl, and P. auru- lentus Bright, more than 2,000 other speci- mens, and numerous field observations, it was concluded that all represent the same species. Similar variability of the female frons occurs in Pityophthorus cariniceps Le- Conte, the P. balsameus complex of species, Ips tridens (Mannerheim), and 7. pilifrons Swaine. Pityophthorus ornatus Blackman Pityophthorus ornatus Blackman, 1928, New York St. Coll. For., Syracuse, Tech. Pub. 25:102 (Holotype, female; Manitou, Colorado; U.S. Nat. Mus.) Pityophthorus limatus Wood, 1964, Great Basin Nat. 24:65 (Holotype, female; Sanford Canyon, Dixie N. F., Utah; Wood Coll.). New synonymy The female holotypes of Pityophthorus or- natus Blackman and P. limatus Wood and 111 other specimens were examined. Of the 10 series examined, 4 from Colorado and 6 from Utah, the host was Picea pungens for 8 of them; 2 were from Pinus and both ap- peared to be accidental associations. The population in Colorado has the vestiture on the female frons longer, particularly on the margins, and apparently less dense. The host and differences on the female frons led to the description of limatus. It is now rec- ognized as a slight geographical variant and should be placed in synonymy as indicated above. Pityophthorus pseudotsugae Swaine Pityophthorus pseudotsugae Swaine, 1918, Dom. Can- ada Dept. Agric. Ent. Br. Tech. Bull. 14(2):99 (Holotype, female; BX Mountain, Vernon District, British Columbia; Canadian Nat. Coll.) Pityophthorus thatcheri Bright, 1977, Great Basin Nat. 36:442 (Holotype, female; Big Sandy Meadow, S28, T5S, R2E, California; Wood Coll.). New syn- onymy The female holotypes of Pityophthorus pseudotsugae Swaine and P. thatcheri Bright and more than 500 other specimens of this species were examined and compared di- rectly to one another, including the entire type of series of P. thatcheri and several other specimens taken in series with it. It is apparent that the series named P. thatcheri was either accidental in Pinus lamhertiana or else an error occurred in recording the host. These specimens are normal represen- tatives of P. pseudotsugae and must be placed in synonymy as indicated above. Pityophthorus pidlus (Zimmermann) Crypturgus pullus Zimmermann, 1868, Trans. Amer. Ent. Soc. 2:143 (Holotype, female; South Caro- lina; Mus. Comp. Zool.); Bright, 1976, Coleopt. Bull. 30:187 (?Lectotype) Pityophthortis bisulcatus Eichhoff, 1869, Berliner Ent. Zeitschr. 12:274 (Holotype, sex?; America bo- realis; apparently lost with Hamburg Mus.). New synonymy The type series of Pityophthorus bisul- catus Eichhoff apparently was lost in the destruction of the Hamburg Museum. Eich- hoff (1878, Mem. Soc. Roy. Sci. Liege (2)8:185) indicated possible synonymy of his species with P. pullus Zimmermann. The size of bisulcatus was given as "% lin." (1.7 mm) in 1869, but 1.5 mm in 1878. If the 1.5 mm measurement is correct, it would almost certainly place bisulcatus in synony- my with annectens; if the 1.7 mm measure- ment is correct it might place it in synony- my with either pullus or confusus Blandford. Since the Eichhoff description does not treat anatomical areas that would distinguish bisulcatus from confusus or an- nectens (the only other species of com- parable size from the southeastern United States where Eichhoff received specimens), it appears advisable to as.sume that Eich- hoff, 1878, was correct and place his species in synonymy under pullus. Dec. 1977 wood: AMERICAN BARK BEETLES 517 Scolytodes levis Blackman Ctenophonis laevigatus Chapuis, 1869 (nee. Ferrari, 1867), Synopsis des Scolytides, p. 49 (Syntypes, males; Colombie; Brussels Mus.). Preoccupied Hexacolus levis Blackman, 1943, Proc. U.S. Nat. Mus. 94(3174):382 (Holotype, female; Paraiso, Canal Zone, Panama; U.S. Nat. Mus.) Scolytodes chapiiisi Wood, 1977, Great Basin Nat. 37:210 (Replacement name). New synonymy Ctenophonis laevigatus Chapuis was based on two male syntypes. These were compared to my long series from Colonia Tovar, El Laurel, and Ranch Grande, Vene- zuela, and apparently agree in all details. Females from my series were compared to the female holotype of Hexacolus levis Blackman and were found to be conspecific. Assuming that the association of males is correct, the name levis has priority and should be used to designate this species. Ni Species Pityophthorus crotonis, n. sp. This large species is remotely related to Conophthocranulus blackmani Schedl, but is distinguished by numerous characters de- scribed below. Male.— Length 3.8 mm (paratypes 3.2- 3.9 mm), 2.5 times as long as wide; color very dark brown. Frons convex, an indistinct, feeble, trans- verse impression near middle; surface smooth, shining, coarsely, closely punctured; vestiture short, inconspicuous. Pronotum 1.03 times as long as wide; sides almost straight and parallel on basal half, rather broadly rounded in front; ante- rior margin armed by 10 low serrations; summit at middle; anterior half with low, dense, confused asperities; posterior half smooth, shining, impressed points fine, abundant, punctures fine, rather close. Ves- titure sparse, confined to marginal areas. Elytra 1.4 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal two-thirds, broadly rounded (feebly bi-emarginate) be- hind; striae 1 weakly impressed; surface smooth, shining, punctures moderately coarse, close, confused. Declivity very steep. rather broadly bisulcate; striae 1 very strongly, 2 strongly impressed, punctures in rows, fine, strongly impres.sed; interstriae 1 moderately elevated, broadly roimded, with dense impressed points and a few punctures, armed at base by a row of about four fine denticles, 2 slightly wider than 1, weakly convex, smooth, shining, with numerous im- pressed points, 3 elevated as high as 1, armed by a row of about 6-10 coarse den- ticles, lateral areas rather coarsely punc- tured. Vestiture apparently derived from odd-numbered interstriae, very long, absent on declivital interstriae 1. Female.— Similar to male except appar- ently smaller, denticles on anterior margin of pronotum and on elytral declivity small- er, strial punctures on disc usually in dis- tinguishable rows. Type Locality.— Thirty km east of Me- rida, Merida, Venezuela. Type Material.— The male holotype, fe- male allotype, and 93 paratypes were taken at the type locality on 8-1-1970, 2500 m. No. 220, Croton boles and limbs, by me. Two paratypes are from 20 km W Merida, Merida, Venezuela, lO-X-1969, No. 45, same elevation, host, and collector. The holotype, allotype, and paratypes are in my collection. Scolytodes canalicidus, n. sp. This species is distinguished from the re- motely allied levis Blackman by the larger size and by numerous other characters de- scribed below. Female.— Length 3.0 mm (paratypes 2.5- 3.1 mm), 2.4 times as long as wide; color yellowish brown to light brown. Frons uniformly, moderately convex; sur- face reticulate except smooth, shining, and impunctate on a sulDtriangular area extend- ing from median third of epi.stomal area to an indefinite median point two-thirds of dis- tance to upper level of eyes, reticulate area with rather numerous, fine punctures near margins of .shining area; vestiture of fine, rather sparse, long hair on area below eyes. Eyes separated above by less than twice width of an eye. Antennal scape orna- 518 GREAT BASIN NATURALIST Vol. 37, No. 4 mented by a tuft of about 20 long setae. Pronotum 1.0 times as long as wide; about as in levis; anterior slope unarmed, surface reticulate, punctures fine, rather close but distinctly larger than in levis. Elytra 1.4 times as long as wide, 1.6 times as long as pronotum; sides straight on basal two-thirds, very slightly wider at base of declivity; striae not impressed except 1 moderately from base, punctures moderately coarse, rather deep; interstriae shining, al- most smooth (obscurely reticulate in some areas), almost three times as wide as striae, punctures fine, close, uniseriate on 2 and 3, moderately confused on other interstriae. Declivity steep, shallowly sulcate; striae 1 strongly impressed; interstriae 1 convex, lat- eral areas ascending from striae 1 rather abruptly to rounded summit at striae 3; sculpture about as on disc. Vestiture of a few minute strial and interstrial setae (short- er than distance equal to diameter of a puncture), mostly on posterior half, and sparse rows of rather long interstrial setae on and near declivity. Male.— Similar to female except frons uniformly reticulate, shining area and most long setae absent, antennal scape without tuft of long hair. Type Locality.— La Carbonera Experi- mental Forest, about 50 km (airline) west of Merida, Merida, Venezuela. Type Material.— The female holotype, male allotype, and 140 paratypes were taken at the type locality on 9-XII-1969, 2,500 m, No. 177, from Clusia branches, by me. The holotype, allotype, and paratypes are in my collection. Scolytodes constrictus, n. sp. This species is distinguished from varius Wood by the much finer pronotal and elyt- ral punctures, by the greatly reduced fe- male frontal pubescence, by the more nearly flattened elytral declivity, and by other characters described below. Female.— Length 2.7 mm (paratypes 2.5- 2.8 mm), 2.8 times as long as wide; color al- most black, some specimens with elytra uni- formly dark brown. Frons flat on lower two-thirds, moder- ately convex above; smooth and shining on flattened area, reticulate above, punctures fine, distinctly impressed; vestiture sparse, rather short, inconspicuous. Pronotum 1.17 times as long as wide, sides widest one-fourth pronotum length from anterior margin, rather strongly con- stricted on posterior half; summit in front of middle, anterior slope finely, closely aspe- rate; posterior areas mostly smooth, shining, some areas of weak reticulation in some specimens; punctures mostly fine, rather close, distinctly impressed. Vestiture of fine, long, moderately abundant, hairlike setae. Elytra 1.7 times as long as wide, 1.5 times as long as pronotum; outline about as in varius; striae not impressed, punctures fine, spaced by twice diameter of a punc- ture; intestriae three times as wide as striae, smooth, shining, punctures slightly smaller and spaced equal to those of striae. Declivi- ty rather steep, very broadly convex; sculp- ture similar to disc, but all punctures much smaller, striae not impressed. Vestiture as in varius, but slightly shorter. Male.— Similar to female except frons more strongly convex, with punctures finer. Type Locality.— Piedras Blancas, 11 km west of Medellin, Antioquia, Colombia. Type Material.— The female holotype, male allotype, and 27 paratypes were taken at the type locality on 17-VII-1970, 2300 m. No. 688, from Baccharus, by me. The holotype, allotype, and paratypes are in my collection. Scolytodes contractus, n. sp. This species is distinguished from varius Wood by the slightly more slender body form, by the sparse, shorter, female frontal pubescence, and by the smaller pronotal and elytral punctures. It is distinguished from constrictus Wood by the brownish col- or, by the coarser pronotal and elytral punctures, and by the more convex declivi- ty, with the striae clearly impressed. Female.— Length 2.8 mm (paratypes 2.5- Dec. 1977 wood: AMERICAN BARK BEETLES 519 3.1 mm), 2.7 times as long as wide; color brown, anterior half of prothorax and head usually dark brown. Frons as in constrictus except lateral areas reticulate to epistoma, punctures dis- tinctly larger. Pronotum as in constrictus except punc- tures distinctly larger. Elytra as in constrictus except striae feebly impressed, punctures larger, closer, spaced by less than diameter of a puncture, interstriae twice as wide as striae, punctures closer, declivity more strongly convex, with striae weakly impressed and punctures slightly larger. Vestiture about as in varius. Male.— Similar to female except lower frons more distinctly convex. Type Locality.— Thirteen km southwest of El Vigi'a, Merida, Venezuela. Type Material.— The female holotype, male allotype, and 34 paratypes were taken at the type locality on 22-X-1969, 100 m. No. 95, from a large vine (liana), by me. The holotype, allotype, and paratypes are in my collection. Scolytodes festus, n. sp. This species is distinguished from atratus Blandford by the different frons in both sexes, by the much coarser punctures on the pronotum and elytra, and by the longer elytral setae. Female.— Length 2.8 mm (paratypes 2.5- 2.8 mm), 2.0 times as long as wide; color black. Frons almost flat to feebly concave on lower half, about as in jucundus Wood ex- cept reticulation less distinct, punctures slightly larger, less abundant. Pronotum about as in atratus except punctures much larger, deep, interspaces equal to about twice diameter of a punc- ture. Glabrous except near margins. Elytra about as in atratus except striae not impressed, strial and interstrial punc- tures slightly larger, mostly in rows, declivi- tal striae not impressed, punctures much smaller than on disc, declivital tubercles smaller. Vestiture longer than in atratus, half as close, each seta on disc as long as distance between rows, not longer on de- clivity; short recumbent strial hair present on lower and lateral areas of declivity. Male.— Similar to female except pronotal asperities larger, more abundant, frons very different. Frons similar to male jucundus except upper margin of transverse impres- sion gradually elevated, lower margin abrupt, crest more uniformly elevated, me- dian summit not evident. Type Locality.— Piedras Blancas, 11 km west of Medellin, Antioquia, Colombia. Type Material.— The female holotype, male allotype, and nine paratypes were taken on 17-VII-1970, 2,300 m. No. 691, from petioles of fallen Cecropia leaves. The holotype, allotype, and paratypes are in my collection. Scolytodes jucundus, n. sp. This species is distinguished from atratus Blandford by the larger size and by the dif- ferent frons in both sexes and declivity as described below. Female.— Length 3.3 mm (paratypes 3.0- 3.5 mm), 2.1 times as long as wide; color very dark brown to almost black. Frons about as in atratus except surface between punctures reticulate, median callus entirely absent on lower half. Pronotum 1.1 times as long as wide; about as in atratus except reticulation slightly more strongly impressed, punctures slightly more numerous. Elytra 1.3 times as long as wide, 1.5 times as long as pronotum; about as in at- ratus except striae less distinctly impressed, all punctures slightly smaller, granules on declivity slightly smaller, setae more widely spaced, apparently shorter. Male.— Similar to female except frons above level of antennal insertion convex, reticulate, rather finely punctured; area at level of antennal insertion broadly, rather strongly, transversely impressed, lower mar- gin of impression forming a transverse crest above flattened epistomal area, crest with a median and a pair of lateral low summits; pronotal asperities larger and more numer- ous than in female. 520 GREAT BASIN NATURALIST Vol. 37, No. 4 Type Locality.— Ranch Grande, Aragua, Venezuela. Type Material.— The female holotype, male allotype, and 17 paratypes were taken at the type locality on 9-IV-1970, 1,100 m, No. 403, from petioles of fallen Cecropia leaves, by me. The holotype, allotype, and paratypes are in my collection. Scolytodes opacus, n. sp. This species is distinguished from opimus Wood by the smaller size, by the darker color, and by other characters described be- low. Female.— Length 3.0 mm (paratypes 2.5- 3.0 mm), 2.0 times as long as wide; color very dark brown. As in opimus except frons more distinctly convex, granules on elytral disc slightly larger, punctures on sides of elytra slightly larger, deeper, elytral vestiture slightly stouter, shorter. Type Locality.— Merida, Merida, Vene- zuela. Type Material.— The female holotype, male allotype, and 93 paratypes were taken at the type locality between Teleferico Sta- tions 2 and 3 on 27-11-1970, near 3,000 m. No. 331, from the 10 cm bole of a tree with extremely large, simple leaves, by me. The holotype, allotype, and paratypes are in my collection. Scolytodes opimus, n. sp. This species represents the extreme ex- pression of characters in a group of species allied to punctatus Eggers. Superficially it resembles Gymnochilus, but fundamental characters clearly place it in Scolytodes. It is distinguished by the large size, by the stout form, and by the contour and sculp- ture of the elytra. Some of Eggers' "Pro- blechilus" species are closely related. Female.— Length 3.3 mm (paratypes 2.7- 3.5 mm), 1.9 times as long as wide; color brown. Frons almost flat below level of antennal insertion, an indistinct, transverse impression above this point, weakly convex above; a low, oblique carina from margin of antennal insertion to lateral part of epistomal mar- gin; surface strongly reticulate, punctures very fine, moderately close, those above level of antennal insertion finely granulate; vestiture of short, fine, moderately abun- dant, inconspicuous hair. Pronotum 1.0 times as long as wide, sub- circular in outline; evenly, rather strongly arched from anterior margin to base; ante- rior margin armed by a row of about a doz- en fine serrations; surface reticulate; ante- rior area armed by numerous small, isolated asperities, these decrease in size but not in density, disappear on basal fifth, and are re- placed by fine, close, moderately deep punctures. Vestiture fine, short, inconspic- uous, covering entire surface. Elytra 1.15 times as long as wide, 1.4 times as long as pronotum; sides weakly ar- cuate and subparallel on basal two-thirds, broadly rounded behind; declivity com- mencing on basal third near suture, near middle in lateral areas; striae not impressed, punctures very minute, distinct; interstriae smooth, shining, about 6 to 10 times as wide as striae, punctures similar to those of striae, confused, those near declivity granu- late (subvulcanate). Declivity gradual on up- per half, steep below, convex except shal- lowly sulcate toward striae 1; striae 1 moderately, rather broadly impressed, as- cending gradually to striae 3; sculpture as on disc, most punctures on interstriae 1 and 2 weakly, irregularly subvulcanate. Vestiture of moderately abundant hair, setae on de- clivity short, longer on disc, with many in- terstrial setae on disc exceedingly long (these long setae abraded on older speci- mens. Male.— Similar to female in all respects. Type Locality.— La Carbonera Experi- mental Forest, about 50 km (airline) west of Merida, Merida, Venezuela. Type Material.— The female holotype, male allotype, and 27 paratypes were taken at the type locality on 9-XII-1969, 2,500 m. No. 172, from an unidentifiable vine (liana) 5-8 cm in diameter, by me. Dec. 1977 wood: AMERICAN BARK BEETLES 521 The holotype, allotype, and paratypes are in my collection. Scolytodes praeceps, n. sp. This species is distinguished from canal- iculus Wood by the very different frons and elytra as described below. Female.— Length 2.8 mm (paratypes 2.4- 2.8 mm), 2.4 times as long as wide; color dark brown. Frons weakly convex, median half from epistoma to three-fourths distance to upper level of eyes smooth, shining, impunctate, lateral and upper areas finely, rather closely punctured; vestiture of moderately abun- dant, very long hair on punctured areas. Eyes rather widely separated above. Anten- nal scape with about a dozen long setae. Pronotum about as in canaliculus except punctures larger, closer, interspaces slightly wider than diameter of a puncture. Elytral outline about as in canaliculus; striae not impressed, punctures rather small, deep; interstriae smooth, shining, three times as wide as striae, punctures mostly uniseriate, a few of those on 1 and 2 near declivity finely granulate. Declivity very steep, broadly convex; striae 1 moderately, 2 and 3 weakly impressed; sculpture about as on disc except interstriae 1-3 each with a row of fine granules on upper two-thirds. Vestiture of minute strial hair and rows of stout, erect, rather closely spaced bristles; each bristle on disc about as long as dis- tance between rows, on declivity about two-thirds this length. Male.— Similar to female except frons more strongly convex, uniformly reticulate, vestiture sparse, inconspicuous. Type Locality.— Piedras Blancas, 10 km east of Medellin, Antioquia, Colombia. Type Material.— The female holotype, male allotype, and 11 paratypes were taken at the type locality on 15-VII-1970, 2,500 m. No. 660, from a Clusia branch, by me. The holotype, allotype, and paratypes are in my collection. Scolytodes serenus, n. sp. This species is distinguished from levis (Blackman) by the much larger size, by the weak pronotal asperities, and by the larger pronotal and elytral punctures. Female.— Length 2.8 mm (paratypes 2.6- 3.0 mm), 2.5 times as long as wide; color pale brown, elytra usually yellowish brown. Frons as in levis except more strongly flattened to vertex (not receding). Pronotum similar to levis except anterior third very finely asperate (asperities visible only when light comes from appropriate angle); reticulation on posterior areas coarser, punctures slightly larger. Glabrous. Elytra similar to levis except strial and interstrial pvmctures larger and deeper; in- terstriae about four times as wide as striae, punctures spaced by one to two diameters of a puncture, about as large as those of striae, in almost definite rows; sculpture on declivity about as on disc except all punc- tures slightly smaller. Male.— Similar to female except frons weakly convex on upper two-thirds, sub- glabrous, pronotal asperities distinctly larger. Type Locality.— La Mucuy, 20 km west of Merida, Merida, Venezuela. Type Material.— The female holotype, male allotype, and 37 paratypes were taken at the type locality on lO-X-1969, 2,500 m, No. 46, from a Martiiio branch (presumably Meriana sp.), by me. The holotype, allotype, and paratypes are in my collection. Scolytodes suturalis, n. sp. This species is distinguished from atratus (Blandford) by the slightly smaller size, by the different frons in both sexes, by the slightly larger pronotal punctures, and by the very different elytra. Male.- Length 2.7 mm (paratypes 2.5- 2.7 mm), 2.1 times as long as wide; color very dark brown. Frons about as in jucundus Wood except punctures fine, transverse impression larger, much deeper, transverse crest much higher, subacute, almost uniformly elevated; upper areas reticulate, rather coarsely punctured; vestiture inconspicuous. Pronotum about as in atratus except punctures slightly larger, interspaces aver- 522 GREAT BASIN NATURALIST Vol. 37, No. 4 aging three to four diameters of a puncture. Elytra resembling atraUis. Striae 1 strong- ly impressed on posterior three-fourths, oth- ers narrowly, weakly impressed, punctures very fine, subconfluent, rows not straight; interstriae about five times as wide as striae, smooth, shining, punctures mostly replaced by rounded granules almost to base, unise- riate. Declivity slightly steeper than in at- ratus, striae narrowly, rather deeply im- pressed, punctures as large as on disc but not as close, interstriae more narrowly con- vex, granules slightly closer, more regular, vestiture much stouter, slightly longer, ex- tending to base. Female.— Similar to male except lower frons about as in jiiciindus except more dis- tinctly concave, reticulation absent; strial impressions not as strong as in male and granules smaller, particularly on disc. Type Locality.— El Laurel Experimental Farm, 12 km southwest of Caracas, Vene- zuela. Type Material.— The male holotype, fe- male allotype, and five paratypes were taken at the type locality on l-V-1970, 1,300 m. No. 461, from petioles of fallen Cecropia leaves, by me. The holotype, allotype, and paratypes are in my collection. Scolytodes varius, n. sp. This species apparently is allied to piinc- tatus Eggers, but it is distinguished by the slender body form, by the narrower posteri- or third of the pronotum, and by numerous other characters described below. Female.— Length 2.7 mm (paratypes 2.5- 3.1 mm), 2.5 times as long as wide; color very dark brown except yellowish brown on intestriae 2-4 to 2-6 and declivity; males yellowish brown except anterior third of pronotum darker. Frons broadly flattened to feebly concave from epistoma to upper level of eyes; sur- face smooth, shining, rather coarsely, close- ly, almost uniformly punctured except a median, impunctate callus on median fourth at level of antennal insertion; vestiture mod- erately abundant, fine, very long. Pronotum 1.12 times as long as wide; widest on anterior half, sides weakly con- stricted, anterior margin semicircularly rounded, unarmed; anterior slope finely, closely asperate; posterior areas coarsely, rather closely punctured, surface obscurely, irregularly subreticulate. Vestiture fine, very long, rather abundant. Elytra 1.5 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; striae not impressed, punctures moderately coarse, deep, spaced by diameter of a puncture; in- terstriae shining, almost smooth, punctures uniseriate, as close as and almost as large as those of striae. Declivity steep, broadly con- vex; sculpture as on declivity except punc- tures very slightly smaller. Vestiture of fine, long strial and interstrial hair, some inter- strial setae three or more times as long as distance between rows. Male.— Similar to female except color imiformly pale over most of body, frons dis- tinctly convex on upper half, its vestiture sparse, inconspicuous. Type Locality.— La Mucuy, 20 km west of Merida, Merida, Venezuela. Type Material.— The female holotype, male allotype, and 116 paratypes were taken at the type locality on 12-XI-1969, 2,500 m. No. 129, from a tree branch, by me. The holotype, allotype, and paratypes are in my collection. NECTAR-SUGAR CONCENTRATIONS AND FLOWER VISITORS IN THE WESTERN GREAT BASIN Larry J. Gut,', Robert A. Schlising,- and Carol E. Stopher- Abstract.— Nectar-sugar concentrations and major flower visitors were determined for 15 species of plants in the Eagle Lake area of Northeastern California. Sugar concentrations for 12 of these are reported for the first time, with means ranging from a low of 10 percent in Mentzelia laevicauUs to a high of 63 percent in Ranunculus un- cinatus. The utilization of the various nectar concentrations varied with the type of flower visitor as well as with the habitat and distributional ranges of the plant and/or animal. Hummingbirds and hawkmoths were not observed visiting the flowers they typically visit in other areas (e.g. Aquilegia and Ipomopsis, or Oenothera), but here pre- ferred more concentrated nectar (Cirsium spp., with x of 57 percent sugar). Specialization in nectar use is report- ed at the generic and specific level in Hymenoptera and Lepidoptera; solitary bees, as a whole, used slightly less concentrated nectar (x = 38 percent sugar) than butterflies (x = 44 percent sugar). Numerous .studies dealing with plant- animal interactions report the importance of flower characteristics such as shape, color, and odor in determining which animals visit a particular species. Recent studies have shown that a correlation also exists between the type of animals which visit a plant and its nectar composition, including the volume of nectar (Heinrich and Raven 1972), types of sugars (Percival 1961, 1965, Wykes 1952), concentration of sugars (Watt, Hoch, and Mills 1974, Baker 1975), and other nec- tar constituents such as amino acids and proteins (Baker and Baker 1975). In this pa- per we present data on nectar-sugar con- centrations in several nectars utilized by different classes of flower visitors in our study area at the south end of Eagle Lake, Lassen County, northeastern California. The area is characterized by open forests of western juniper {Junipeni.s occidentalis) and Jeffrey pine {Pinus jeffreyi), and by more open areas dominated by big sagebrush (Ar- temisia tridentata) and rabbitbrush {Chnjso- thamnus nauseosm). The plant populations used in this study (June to August 1976) were located at elevations between 1530 and 1800 m above sea level; plant names are according to Munz and Keck (1968). Methods and Materials Nectar was collected in the field with 10 jLtl microcapillary pipettes (Drummond Scientific Co.). For extraction from narrow, tubular flowers, the pipettes were drawn out into fine points. Approximately 24 hours prior to nectar extraction the flowers were covered with sheets of porous lens tis- sue (15 x 20 cm) to keep flower visitors from removing the nectar. The percentage of sugar was determined in the field with a Bellingham and Stanley pocket refractome- ter, which read up to 50 percent. For nec- tars more concentrated than this, and for samples smaller than 3-4 ju,l, an equal amount of distilled water was measured in a second calibrated pipette, and mixed with the nectar sample on the stage of the ref- ractometer. The reading obtained is based on the refractive index of the solution. Nec- tar .sugars in flowering plants consist mainly of sucrose, fructose, and gluco.se in varying proportions; sucrose is the most widespread and usually predominates (Percival 1961). Sucrose, fructose, and gluco.se give similar refractive index readings for equal percent solutions by weight (Wykes 1952); there- fore, we report our readings as "nectar-.sug- ar," "sucro.se," or simply "sugar." 'Department of Entomology, Oregon State University, Corvallis 97331. •Department of Biological Sciences, California State University, Chico ! 523 524 GREAT BASIN NATURALIST Vol. 37, No. 4 Flower visitor data were obtained through observations and collection rather than by consulting the literature. On several occasions flower visitors were observed and collected during three time periods (morn- ing, afternoon, and evening), with approx- imately equal time being spent at each plant species. Flower visitors referred to be- low as "major" are those which were ob- served on the flowers on each day (although not necessarily during all three time peri- ods). Representative insect specimens are on file in the Entomology Museum, California State University, Chico. Results and Discussion The nectar-sugar concentrations of the 15 species sampled are presented in Table 1. Means of our readings agree fairly closely with those previously reported for three species: Aquilegia formosa, 25 percent (vs 32 percent by Baker 1975); Ipornopsis ag- gregata, 25 percent (vs 22 percent by Watt, Hoch, and Mills 1974, and ca 23 percent by Hainsworth 1973); Oenothera hookeri, 32 percent (vs 26 percent by Stockhouse 1975). The mean percent sugar contained in the nectars ranged from a low of 10 percent in Mentzelia kievicaulis to a high of 63 per- cent in Ranunculus uncinatus. A large vari- ation in nectar-sugar concentrations was also observed within most species. Asclepias fascicularis, the most extensively studied species, had nectar-sugar readings ranging from 16 to 72 percent. The openly exposed nectar of this species makes it highly sus- ceptible to environmental factors which cause evaporation and/or dilution; these factors can account, in part, for the wide range of nectar concentrations observed (Stopher, Schlising, and Gut, ms in prepara- tion). Table 2 is a summary of the major flower visitor types found on flowers of the 15 spe- cies of plants studied. Wasp, fly, beetle, and ant visitor types are listed here (and wasps and ants again in Table 3), but since there are no detailed data available for these types they will not be further discussed in Table 1. Fifteen plant species studied near Eagle Lake, June to August 1976, arranged according to nectar- sugar concentrations. Major flower visitors observed are also listed for each species, roughly in order of impor- tance. Visitor code letters refer to the abbreviations listed in Tables 2-4. Plant names are from Munz and Keck (1968). Percent Sugar No. Flowers Species Mean Range Sampled Flower Visitors Raminctthis uncinatus 6,3 62-63 3 Sb Wp Bf Dl Ad Vp Ly Sy Fl Bt At Cirsium californicum 59 44-74 20 Sb Bb Wp Bf Hm Xy An Os Ag Vp Sp Pp Sv Ml Lm Fl Bt At Cirsium breweri 54 35-66 28 Sb Bb Hk Hm An Mg Dl Sp Agastache parvifolia 50 41-76 14 Sb Bf An Os Dl Sy Bt Asclepias fascicularis 47 16-72 365 Sb Ap Bb Wp Bf Xv An Mg Dl Sh Hy Vp Sp Dn Pp Lv Lm Fl Bt Scrophularia califomica 43 39-49 5 Sb Wp Dl Hv Vp Fl At Asclepias speciosa a3 21-42 66 Sb Ap Wp Bf Cr An CI Mg Dl Hy Vp Sp Sn Pp Ly Fl Bt Oenothera hookeri 32 23-38 25 Monardella odoratissima 31 17-44 12 Sb Bf Os Sy Ml Bt Ipcnmypsis aggrcgata 25 17-36 33 Fl Acfttilegia fomwsa 25 14-40 94 SbCr Schoenolirion album 22 16-.33 14 Sb Ap Bb Wp Bf Cr CI Hy Vp Ph Bt At Pentstemon speciosus 21 14-28 8 Sb Cr Os Dl Hy At Nocotiana attenuate 21 14-33 27 Mentzelia laevicaulis 10 7-18 27 Sb Ap Dl Fl Overall 36 7-76 741 Dec. 1977 GUT ET AL.: NECTAR-SUGAR CONCENTRATIONS 525 this paper. All bees other than the honey- bee and bumblebees have been grouped un- der solitary bees. Solitary bees comprise the largest number of flower visiting species in the area and were found on all but three of the plant species studied. With the exception of Asclepias fascicu- laris (mean nectar concentration of 47 per- cent sugar), honeybee visits were restricted to the flowers with the more dilute nectars (i.e., concentrations below 35 percent). All of the other flower visitor types preferred more concentrated nectars. However, all of these visitor types had representative spe- cies which were found on flowers contain- ing a dilute nectar. Schoenolirion album, with an average nectar-sugar concentration of 22 percent, was an especially utilized di- lute nectar source. Possible reasons for this were easy accessibility to the nectar, many flowers per raceme, and the fact that this species was one of the very few species lo- cally in flower at the time. Two average sugar concentrations are given for both hawkmoth and hummingbird visitor types due to discrepancies between the flowers they typically visit and the flowers they vis- ited in the study area (see below). Hummingbirds.— There was a large differ- ence in the concentration of sugars found in the nectar of the four "hummingbird flow- ers" but these plants can be grouped into two pairs with similar concentrations (Table 2). One pair consists of two "typical" hum- mingbird flowers (e.g., Grant and Grant 1968), Ipomopsis aggregata and Aquilegia fonnosa, both of which had mean sugar concentrations of 25 percent here, but were not visited by birds. The other two species, Cirsium californicurn and C. breweri, had 59 percent and 54 percent sugar, respectively; these were heavily visited by hummingbirds Table 3. Major families and genera of Hymenoptera and the nectar-sugar concentrations utilized (of the 15 plant species studied) near Eagle Lake, Lassen County, California. No. Table 2. Major flower-visitor types and the nectar- Species of Flowers Visited sugar concentrations utilized (of the 15 plant species Percent Sugar studied) near Eagle L ,ake, Lassen County, , California. Visitors Mean Range Anthophoridae 8 39 No. 21-59 Species of Xtjlocopa (Xy) 2^ 53 47-59 PpFfpnt Ceratina (Cr) 4 25 21-33 Flowers Visited Sugar Anthophora (An) Apidae Apis (Ap) 5 6 4 49 38 28 33-59 10-59 10-47 Visitor Type Mean Range Bombus (Bb) 4 46 22-59 Solitary bee (Sb) 12^ 38 10-63 Megachilidae 8 40 21-59 Honeybee (Ap) 4 28 10-47 Osmia (Os) 4 40 21-59 Bumblebee (Bb) 4 46 22-59 Chelostomoides (CI) 2 28 22-33 Wasp (Wp) 6 45 22-63 Megachile (Mg) 3 45 33-54 Butterfly (Bf) 7 44 22-63 Halictidae 9 42 10-63 Hawkmoth (Hk) 1 54 Dialictus (Dl) 8 40 10-63 (Hawkmoth 2" 27 21-32) Agapostemon (Ag) 1 59 Hummingbird (Hm) 2 57 54-59 Sphecodes (Sh) 1 47 (Hummingbird 2^- 25 25-25) CoUetidae 5 .33 21-47 Fly (FI) 7 38 10-63 Hylaeus (Hy) 5 ,33 21-47 Beetle (Bt) 7 40 22-63 Andrenidae 1 &3 Ant (At) 6 38 25-59 Andrena (Ad) 1 63 Overall 15 36 10-ft3 Formicidae (At) 6 38 21-59 Vespidae (Vp) Sphecidae (Sp) 6 44 22-63 ^Abbreviations by visitor type names are listed in Table I to show 4 48 ,33-59 which species of flowers were visited. Overall 12 38 10-63 "No visitors were observed, but these plants are expected to be hawk- moth visited (see text). "■No visitors were observed, but these plants are expected to be hum- mingbird visited (see text). ".Abbreviations by bee genus and by nonbee family names are listed in Table 1 to show which species of flowers were visited. 526 GREAT BASIN NATURALIST Vol. 37, No. 4 in the study area. Moldenke (1976) does list hummingbirds as especially important pol- linators for the genus Cirshim. Exact identi- fications could not be made, but anna's, cal- liope, and rufous hummingbirds are all known to occur in the area (R. Lederer, personal communication 1977). Since hummingbirds are high-energy de- manding animals, it is advantageous for a cross-pollination-dependent plant to produce a less concentrated nectar and "force" the potential pollinator to visit the maximum number of flowers in order to fulfill its own energy needs (Baker 1975, Heinrich and Raven 1972). Previous studies have found that hummingbirds prefer the less concen- trated nectars (Baker 1975, Hainsworth 1973, Hainsworth and Wolf 1972). This is contradictory to our findings, in which hummingbirds did not utilize "their typical flowers" with less concentrated nectars but preferred richer food sources. This has also been found by Stiles (1976), who showed that experimental anna's hummingbirds pre- ferred sucrose and glucose in the highest concentrations available, up to 60 percent. Factors which favor the use of nectar with a weak sugar content may include the dis- advantage of a viscous nectar to a bird which must hover while feeding (Weymouth, Lasiewski, and Berger 1964), the difficulty of imbibing and swallowing a more viscous nectar (Hainsworth 1973, Weymouth et al 1964), and the need the birds have for a free water source (Baker 1975). Then what factors, other than nectar concentration, were important in determin- ing hummingbird flower selectivity reported here? Three important factors may be plant population size and density, growth habit of the plants, and the concentrations of the different sugars in the nectar. Investigations on the energetics of forag- ing by tropical hummingbirds have shown that large numbers of flower visits are re- quired each day (Wolf, Hainsworth, and Stiles 1972). Gass, Angehr, and Centa (1976) reported that temperate zone hummingbirds defend a territory containing 239 floral units of resource value which is equivalent to that produced by 239 Aquilegia fomwsa flowers. The population of A. forrnosa in the present study contained no more than 100 flowers on any one day. Since no hum- mingbirds were seen visiting Aquilegia here, but were common in the area, it seems den- sity of plants may help determine nectar source. Gass et al (1976) noted that rufous hummingbirds regulate the size of their feeding territories in order to maintain food supplies at a level approximating their metabolic requirements. Also, migratory species feed in a wide variety of habitats each year under a wide range of phys- iological stresses. Thus, when a territory supplies insufficient energy, the birds' strat- egy is to seek food elsewhere. This may be the case with hummingbirds we saw, where the populations of Cirsitim hreweri and C. califomicwn were both much larger than that of Aquilegia forrnosa and were in fact the only plants seen visited by the birds. Gass et al (1976), however, also noted that hummingbirds will feed on the nectar of a preferred species first, even if the pop- ulation size is too small to meet the birds' energy requirements, and then turn to an alternate source. The fact that we did not observe any foraging on Aquilegia may be partially explained by noting that the plants of Aquilegia were growing closely inter- spersed with shrubs, while plants of C. hre- weri especially were located in the open only about 10 m from the Aquilegia. The flowers of Aquilegia may not have been as readily accessible and energetically favor- able for nectar foraging. The single population of Ipomopsis aggre- gata studied was also small (about 50 flow- ers at any time), and had the same nectar- sugar concentration as Aquilegia and per- haps even a similar floral unit of resource value. This may again suggest that popu- lation density could be an important factor in determining the nectar utilization strate- gy seen in this study. Yet another possible factor contributing to the nectar selection of the hummingbirds is the proportions of the different sugars in the nectars. The kinds of sugars present were not determined for our Eagle Lake plants; but Cirsium califomicwn sampled Dec. 1977 GUT ET AL.: NECTAK-SUGAR CONCENTRATIONS 527 elsewhere was found to have a rather high percent of ghicose (58 percent) in the sug- ars present (I. Baker, personal commu- nication, 1976). Stiles (1976) reported the following hierarchy of sugar preferences in several species of hummingbirds: sucrose over glucose over fructose, with an equal- parts mixture of the three falling some- where in the middle of the preference or- der. A nectar containing a large percentage of glucose may be less viscous than one containing a large percentage of sucrose. Thus the high percentage of total sugars in the nectar of C. californicum (a potentially highly viscous nectar) may be compensated for by the high percentage of glucose; this may make the nectar less viscous, which, as noted earlier, is advantageous to the feeding of the birds. Hawkmoths.— a situation similar to that with the hummingbirds existed for the hawkmoths and the flowers they visited in our study area (Table 2). No hawkmoths were collected, but common species in the area include Sphinx perelegans (Hy. Edw.) and Hyles Hneata (Fabr.) (from collections at California State University, Chico, and Eagle Lake Field Station). Of the three spe- cies of potential hawkmoth flowers present, Nicotiana attenuata (Wells 1959) and Oeno- thera hookeri (Baker 1961, Stockhouse 1975) are "typical" hawkmoth-visited flowers (see also Moldenke 1976). They both had rather low average nectar concentrations of 21 percent and 32 percent, respectively. Baker (1975) also found hawkmoth flowers to have lower nectar concentrations similar to those of hummingbird flowers. At Eagle Lake, as with the hummingbird flowers, the hawk- moths were not seen visiting "their typical flowers." Instead, the moths preferred the flowers of Cirsium breweri, with a concen- trated nectar (x = 54 percent). Hawkmoths are homeothermic, hence high energy-demanding ing insects, and in this relatively cool region may require a more concentrated energy source. Also, plant population size was pos- sibly a limiting factor. The two typical hawkmoth plants were represented by very small populations of about 10 individuals each, with 25 open flowers per evening; Cirsium breweri heads were very abundant. (Stockhouse [1975] however, found that a single flower of Oenothera caespitosa pro- duces an average of 42.1 calories per flower [35 microliters of nectar per night of which approximately 32.5 percent is sugar], which he believes is a large enough potential energy source for hawkmoths even in a small population, with only 20-50 flowers open on a given night. Thus our few flow- ers of O. hookeri could also be a serviceable food source.) Hymenopterans.— a summary of the major flower-visiting Hymenoptera and the con- centrations of nectars they feed on is given in Table 3. Considering the first four fami- lies listed (all bees), it is seen that at the family level the mean sugar concentration utilized varied little— from only 38 percent to 42 percent. More specific trends toward nectar selectivity can perhaps be seen with- in these families. The Anthrophoridae and Megachilidae each had two genera which preferred a more concentrated nectar and a third genus which preferred a more dilute nectar. Although not entirely con.sistent throughout the data, the genera containing the smaller-bodied species seemed to feed on the less concentrated nectars, and the larger-bodied seemed to utilize the more concentrated nectars. However, at the fam- ily level these four means do reflect bee us- age, overall, of fairly concentrated nectars. Baker (1975) found the mean percentage of nectar-sugar for 60 species of California na- tive bee flowers to be only 31 percent. In this study bees of a fifth family (Colletidae) preferred nectar with 33 percent sugars, while bees in a sixth family (Andrenidae) fed exclusively on the richer nectar (63 per- cent sugars) of a single plant species. The Family Andrenidae in this study was represented by two species, Andrena (Di- andrena) cuneilabris Viereck and A. (£u- andrena) caerulea Smith (formerly A. com- plexa). The.se bees restricted their foraging to the flowers of Ranuncuhi.s uncinatus, which produced the most concentrated nec- tar (63 percent) found in our samples. In a .study done in the Coast Ranges of Califor- nia, A. complexa was found to feed exclu- 528 GREAT BASIN NATURALIST Vol. 37, No. 4 sively in the flowers of Ranunculus califor- nicus (Linsley and MacSwain 1959). Other species of Andrena have also been found to feed almost exclusively on various species of Ranunculus (Thorpe 1969, Linsley and MacSwain 1959). Host-specific relationships such as these have usually been described in terms of habitat, flower morphology, and /or pollen source specificity. If nectar con- centration and sugar content values are "conservative characters," at least at the genus level (Percival 1961), then the high nectar-sugar concentration in R. uncinatus and perhaps other species of Ranunculus may also be a factor influencing these host- specific plant-insect interactions. Butterflies.— Data for the four major fam- ilies of butterflies found in this study are given in Table 4. The range of means for the nectar is only from 40 percent to 48 percent. In general, butterflies were feeding on a concentrated nectar source. Although data are not given. Baker (1975) reported that nectars of butterfly flowers are slightly less concentrated than those of bee flowers. Our study suggests the reverse: flowers vis- ited by bees had a less concentrated nectar (x = 38 percent) than that of flowers vis- ited by butterflies (x = 44 percent). Table 4. Major families and genera of Lepidoptera and the nectar-sugar concentrations utilized (of the 15 plant species studied) near Eagle Lake, Lassen County, California. No. Species of Percent Flowers Visited Sugar Visitors Mean Range Danaidae (Dn) 2" 40 33-47 Papilionidae (Pp) 3 46 33-59 Lycaenidae (Ly) 3 48 33-63 Nymphalidae 6 45 22-63 Speyeria (Sy) 4 51 31-63 Melitaea (Ml) 2 45 31-59 Limenitui (Lm) 2 5.3 47-59 Phycoides (Ph) 1 22 Overall 7 44 22-63 "Abbreviations by visitor names are listed in Table 1 to show which spe- cies of flowers were visited. In their study on plant resources and adult butterflies in a subalpine environment. Sharp, Parks, and Ehrlich (1974) found that butterfly foraging seemed to show opportu- nistic use of what was available through the season, and no species-specific nectar source relationships were found. Baker and Baker (1975) reported that nectars of butterfly- pollinated flowers are high in amino acids and suggested this may be a factor involved in butterfly foraging strategies, because in most cases this is their only source of these essential chemicals as adults. In this study we did find an apparent specific relation- ship between a butterfly and its local nectar source. The genus Phycoides was represent- ed here by a single species, P. mylitta my- litta (Edw.), which restricted its feeding to the flowers of a single species, Schoenolirion album. During this study adults were only seen foraging while this plant was in flower, although they are known to visit other flower species (personal observations; Ben- seler 1975). Tlie nectar produced by this species was much more dilute (x = 22 per- cent) than the nectar of any of the other butterfly-visited flowers. Elsewwhere, Colias butterflies have been found to prefer more dilute nectars, probably due to their severe problems of water loss associated with small size (Watt, Hoch, and Mills 1974). Individ- uals of Phycoides mylitta mylitta are also small and may have severe water loss prob- lems in the Eagle Lake area where midday readings of 35 percent relative humidity were of common occurrence. The less con- centrated but abundant nectar of Schoenoli- rion may also be an evolutionary trend to early seasonal appearance of both the plant and its visitors. The lack of other flowers "competing" for this potential pollinator may also allow early-flowering Schoenolirion album to somewhat limit its nectar-sugar production without lowering chances for cross pollination here. Conclusions This study emphasizes several points with data collected in 1976. Data on just 15 spe- cies sampled show there was a wide range of nectar-sugar concentrations available to flower visitors found in the Eagle Lake Dec. 197-; GUT ET AL.: NECTAR-SUGAR CONCENTRATIONS 529 area. The concentrations produced not only varied from species to species, but they also varied considerably within flowers of the same species. Some of this variation was doubtless caused by local and immediate en- vironmental conditions, but it was still rep- resentative of the range of nectar-sugars available to flower visitors there. Various nectar-sugar concentrations were utilized by different types of flower visitors. Data on Hymenoptera and Lepidoptera visitors at the family level show some preferences in the concentrations utilized, but more pro- nounced preferences were found at the genus or species level of visitor. Species- specific, plant-insect interactions were seen in both of these visitor types in the Eagle Lake area, and the concentration of nectar sugars may be one of several factors in- fluencing these relationships. Factors such as plant population size and density, flower accessibility, and the degree of physiological stress may vary from location to location and affect the utilization of nectar sources by foragers in a given area. This seemed es- pecially true for the hawkmoth and hum- mingbird flowers, which were not being vis- ited by these animals at Eagle Lake, who preferred more abundant, more concen- trated, or more readily accessible nectar. Acknowledgments We would like to thank Robbin Thorp for identifying the bees, and Irene Baker and Roger Lederer for supplying unpub- lished information. We would also" like to thank Tim Spira for his critical reading of the manuscript and Mark Stopher for his field assistance. We are grateful for the sup- port of a National Science Foundation Un- dergraduate Research Participation Grant (SMI 76-03206) that was administered through the Eagle Lake Field Station. Literature Cited Baker, H. G. 1961. Nocturnal and crepuscular pollina- tors. Quart. Rev. Biol. 36: 64-73. 1975. Sugar concentrations in nectars from hummingbird flowers. Biotropica 7: 37-41. Baker, H. G., a.nd I. Baker. 1975. Studies of nectar- constitution and pollinator-plant coevolution. In: L. E. Gilbert and P. H. Raven (eds.). Animal and plant co-evolution, pp. 100-140. Univ. Texas Press, Austin. Benseler, R. W. 1975. Floral biology of California buckeye. Madrono 23: 41-S3. Gass, C. L., G. A.ngehr, a.nd J. Centa. 1976. Regu- lation of food supply by feeding territoriality in the rufous hummingbird. Can. J. Zool. 54: 2046-2054. Grant, K. A., and V. Grant. 1968. Hummingbirds and their flowers. Columbia Univ. Press, .\ew York. 1 15 p. HLviNswoRTH, F. R. 1973. On the tongue of a hum- mingbird: i'lS role in the rate and energetics of feeding. Comp. Biochem. Physiol. 46.'\: 65-78. Hainsworth, F.R., and L. L. Wolf. 1972. Crop vol- ume, nectar concentration and hummingbird energetics. Comp. Biochem. Phvsiol. 42 A: .359-366. Heinrich, B., and p. H. R.\ven. 1972. Energetics and pollination ecology. Science 176: 597-602. Linsley, E. G., and J. W. MacSwain. 1959. Ethology of some Ranunculus insects with emphasis on competition for pollen. Univ. Calif. Pubis. Ent. 16: 1-46. Moldenke, a. R. 1976. California pollination ecology and vegetation tvpes. Phytologia .34: 305-361 . MuNZ, P.A., and D. D. Keck. 1968. .\ California flora and supplement. Univ. Calif. Press, Berkeley. 1681 -I- 224 p. Percival, M.S. 1961. Tvpes of nectar in angiosperms. New Phytol. 60: 235-281. 1965. Floral biology. Pergamon Press, London. 243 p. Sharp, M.A., D. R. Parks, and P. R. Ehrlich. 1974. Plant resources and butterfly habitat selection. Ecology 55: 870-875. Stiles, F. G. 1976. Taste preferences, color prefer- ences, and flower choice in hummingbirds. Con- dor 78: 10-26. Stockhouse, R. E., II. 1975. .Nectar composition of hawkmoth-visited species of Oenothera (Ona- graceae). Great Basin Nat. 35: 27.3-274. Thorpe, R. W. 1969. Systematics and ecology of bees of the subgenus Diandrena (Hymenoptera: Andre- nidae). Univ. Calif. Pubis. Ent.' 52: 1-146. Watt, VV. B., P. C. Hoch, and S. G. Mills. 1974. Nectar resource use by Colias butterflies. Oeco- logia 14:353-374. Wells, P. V. 1959. An ecological investigation of two desert tobaccos. Ecology 40:626-644. Weymouth, R. D., R. C. Lasiewski, and A. J. Berger. 1964. The tongue apparatus in hummingbirds. Acta Anat. 58:252-270. Wolf, L. L., F. R. Hainsworth, and F. G. Stiles. 1972. Energetics of foraging: rate and efficiency of nectar extraction by hummingbirds. Science 176:1351-1352. Wykes, G. R. 1952. An investigation of the sugars present in the nectar of flowers of various spe- cies. New Phytol. 51:210-215. NEW GRASS DISTRIBUTION RECORDS FOR NEW MEXICO AND THE UNITED STATES' Stephan L. Hatch- Abstract.— New distribution records are given for 13 grasses now found in New Mexico. One of these, Urochloa panicoides Beauv., is a new distribution record for the United States. Recent collections have revealed the oc- currence of 13 taxa new to New Mexico and one taxon new to the United States. These occurrences are extensions to the known distributions of these species as re- corded by Hitchcock (1951) and Gould (1951, 1975a, 1975b). A recent checklist for New Mexico by Martin and Castetter (1970) was used for grass distribution records for the State of New Mexico. Voucher speci- mens for these new records have been de- posited in the New Mexico State University Herbarium (NMC). Brachiaria ciliatissima (Buckl.) Chase Eddy Co.: Thirty-two km east of Carls- bad, Sandia Radioactive Waste Storage Project, 18 September 1976, Wilson sn. (NMC). This collection is a new record for New Mexico. The species has been reported on sandy soils in Arkansas, Oklahoma, and Texas (Gould 1975b). Bromus diandrus Roth Catron Co.: Eight km west of Glenwood, elevation 2621 m, 17 October 1976, Stock- man sn. (NMC). This is a new record for New Mexico. Previously reported from Ari- zona (Gould 1956) and Texas (Gould 1975b). Grant Co.: Locally abundant 25 km west of Silver City, 25 April 1977, Bodenchuk 20 (NMC). Bromus mollis L. Dona Ana Co.: Infrequent I km south of the junction of University Ave. and Inter- state 25, elevation 1189 m, 4 March 1977, Kolo 7 (NMC). Growing between the fron- tage road and the golf course in Las Cruces. This is a new record for New Mexico. Pre- viously reported from Arizona (Gould 1956) and Texas (Gould 1975b). Bromus rubens L. Dona Ana Co.: Locally abundant on roadsides at the junction of Interstate 25 and highway 70 N.E. of Las Cruces, 10 April 1977, Hatch 2399 (NMC). This collec- tion is a new distribution record for New Mexico. This introduced species has been reported from the adjacent states of Arizona (Gould, 1951) and Texas (Gould 1975a). Otero Co.: Locally abundant on disturbed sites, 1 km south of the White Sands Missile Range, 5 March 1976, Williams sn. (NMC). Eremopijron triticeum (Gaertn.) Nevski San Juan Co.: Locally abundant as a roadside weed, 4.7 km east of Shiprock along highway 55, 12 August 1976, Hatch 2242 (NMC). This collection is a new re- cord for New Mexico. This introduced spe- cies from southern Russia has been reported in Montana, Idaho, and Washington (Hitch- cock 1951) and collected in Utah's Box El- der County and Emery County. Muhlenhergia villosa Swallen Otero Co.: Rare in a small patch, east of Orogrande on Otero Mesa, 24 September 1976, Spellenberg 4565 (NMC). This collec- tion is a new record for New Mexico. The taxon was previously reported as endemic to Texas (Gould 1975b). Journal Article 639, Agricultural Experiment Station, New Mexico State University, Las Cruces, New Mexico, 88003. 'Assistant Professor. Department of Animal and Range Sciences, New Mexico State University, Las Cruces, New Mexico, 88003. 530 Dec. 1977 hatch: new grass distribution 531 Panicum texanum Buckl. Dona Ana Co.: Locally abundant as a weed on ditchbanks and in cultivated fields, 3.5 km south of Las Cruces along highway 142, 9 October 1976, Flacker sn. (NMC). This is a new record for New Mexico. Pre- vious reports of collections have been made in Arizona (Gould 1951) and Texas (Gould 1975a). The species is adapted to disturbed conditions of cultivated land and was prob- ably introduced with crop seed from other states. Schisrniis arabiciis Nees Dona Ana Co.: Abundant locally as an adventive grass on the New Mexico State University campus at Las Cruces, 5 March 1976, Williams sn. (NMC). This is a new re- cord for New Mexico. This introduced grass has been reported from Arizona, California (Gould 1951), and Nevada (Hitchcock 1951). Hitchcock (1951) reported the grass to be a good winter forage grass that was "appar- ently" spreading rapidly. Setaria adhaerans (Forsk.) Chiov. Dona Ana Co.: Locally abundant as an adventive on disturbed sites throughout New Mexico State University campus and Las Cruces, 29 September 1976, Hatch 2321 (NMC). Reported from the Trans-Pecos re- gion of Texas, Bermuda, Cuba, Mexico, and Central America (Gould 1975b). This record is new for New Mexico and is the second state having a distribution record in the United States. Setaria ramiseta (Scribn.) Pilger Chavez Co.: An abundant, native pe- rennial grass, growing 40 km east of Hager- man along highway 31 on the caprock, 10 August 1976, Ellsworth 16 (NMC). New re- cord for New Mexico. Previously reported from Texas and northeastern Mexico (Gould 1975b). Triplasis purpurea (Walt.) Chapm. Eddy Co.: Thirty-two km east of Carls- bad, Sandia Radioactive Waste Storage Project on sandy soils, 18 September 1976, Wilson sn. (NMC). Reported from west Texas and Oklahoma (Gould 1975b). This taxon is new to New Mexico. Urocliloa panicoides Beauv. Dona Ana Co.: Occasional weed in lawns at New Mexico State University, Las Cruces, 26 August 1976, Spellenherg 4480 (NMC). This is a new distribution record for New Mexico and the United States, the spe- cies occurs in Africa and South America. The specimen was identified by Jose F. M. Valles from Texas A & M University, Col- lege Station, Texas. Vulpia mytiros (L.) K. C. Grelin Dona Ana Co.: Frequent between "A" Mountain and Interstate 25 southeast of Las Cmces, 10 April 1977, Hatch 2365 (NMC). New record for New Mexico. Previously re- ported from Texas (Gould 1975b) and Ari- zona (Gould 1956). Acknowledgments I thank Jose F. M. Valles, Department of Range Science, Texas A & M University, for the identification of Urochloa and Dr. R. F. Spellenberg, Department of Biology, New Mexico State University, for bringing the Urochloa and Muhlenheroia collections to my attention. Literature Cited GouLu, F. W. 1951. Grasses of southwestern United States. University of Arizona Press, Tucson. 352 P- 1975. Texas plants, a checklist and ecological summary. Texas Agricultural Experiment Station, MP-585,' rev. ed. 121 p. 1975. The grasses of Texas. Texas A & M Press, College Station. 653 p. Hitchcock, A. S., 1951. Manual of the grasses of the United States. USDA Miscellaneous Publication 2(K), rev. by Agnes Chase. 1051 p. Martin, W. C, and E. F. Castetter, 1970. A checklist of gymnosperms and angiosperms of New Mexico. Published privately. STILT SANDPIPER AND HUDSONIAN GODWIT IN NEVADA C. S. Lawson' This note reports the first specimens of Stilt Sandpiper (Micropalama himantopus) and Hudsonian Godwit {Limosa haemastica) collected in Nevada. All specimens were taken at the tailing ponds in Las Vegas Wash, 24 km southeast of Las Vegas, Clark County, Nevada. One specimen of each is now in the Museum of Biology, University of Nevada, Reno, Reno, Nevada (UNMB) and one Hudsonian Godwit specimen is now in the Nevada State Museum (NSM) collection, Carson City, Nevada. Stilt Sandpiper.— Prior to this report there were only two published records of sightings of Stilt Sandpiper in Nevada. In northern Nevada, three birds were observed 30 July 1939 in Churchill County, 24 km northwest of Fallon near Hazen by Slipp (1942, Murrelet 22: 61-62). In south-central Nevada, seven birds were observed at the Nevada Test Site in Nye County 8 October 1961 (Hayward, et. al, 1963, Brigham Young Univ. Sci. Bull. Vol. 3, No. 1: 10). I collected one bird 17 September 1976 (UNMB #1813, immature male, skull not fully ossified, gonad minute, moderate fat, weight 59.25g). Three other sight records, all from Las Vegas Wash, include the fol- lowing: I observed one bird 28 April 1972 and Vince Mowbray observed two birds on two separate occasions, 16 September 1973 and 7 September 1976. Hudsonian Godwit.— There are no pre- vious sightings of this species in Nevada. I collected two of five birds present 15 May 1976, one male and one female. The male is UNMB #1808, adult, skull fully ossified, left testes 8x3mm, light fat, weight 231.05g. The female is now NSM #1616, adult, skull fully ossified, ovary granular, largest ova lx2mm, heavy fat, weight 279.05g. The presence of these birds had been discovered earlier in the day by Vince Mowbray. I would like to thank Vince Mowbray for use of his previously unpublished records. 'Department of Biology, Nevada State Museum, Carson City, Nevada 89710. Mailing address; P.O. Box 5475, Las Vegas, Nevada 532 Dec. 1977 533 INDEX TO VOLUME 37 The genera and species described as new to science in this vokune appear in hold tvpe in this index. A mountain Cerocarpus population— revisited, p. 97. A new fossil sponge from the Ordovician Gar- den City Limestone of southeastern Idaho, p. 475. A new monotypic genus of chiggers and four new species of Qtiadraseta from Venezuela (Acari: Trombiculidae), p. 501. A new name for Galium scahriusculum (Ru- biaceae) of Utah, p. 510. A new species of Rhopalomyia (Diptera: Ce- cidomyiidae) from Artemisia ludoviciana Nuttall (Compositae) in Utah, p. 426. A new status for Penstemon cyananthus Hook, ssp. subglaber Pennell (Scrophulariaceae), p. 100. A survey of breeding and migratory birds south- west of Farmington, New Mexico, p. 489. An annotated list of the aquatic insects of southeastern Idaho, Part II: Trichoptera, p. 253. An undescribed species of Thelypodiopsis (Brassicaceae) from the Uinta Basin, Utah, p. 95. Anderson, Christine Minton, article by, p. 24. Araptus crassus, p. 211. Araptiis obesus, p. 212. Armstrong, David M., article by, p. 457. Astragalus atwoodii, p. 103. Atwood, N. Duane, and Stanley L. Welsh, ar- ticle by, p. 95. Au, P., A. T. Vollmer, and S. A. Bamberg, ar- ticle by, p. 81. Austin, Dennis D., Philip J. Umess, and Michael L. Wolfe, article by, p. 101. Bachman, V. C, and F. L. Knopf, article by, p. 247. Bamberg, S. A., A. T. Vollmer, and F. Au, ar- ticle by, p. 81. Banta, Benjamin H., and Wilmer W. Tanner, ar- ticle by, p. 225. Barnes, A. M., V. J. Tipton, and J. A. Wildie, article by, p. 138. Biomass of fifty conifer forests and nutrient ex- ports associated with their harvest, p. 395. Bionomics of a subterranean gall midge (Dip- tera: Cecidomyiidae) from Artemisia ludovi- ciana, p. 429. Bionomics of two aspen bark beetles, Try- pophloeus populi and Procryphalus mucro- natus (Coleoptera: Scolytidae), p. 105. Birds of Gunnison and Cub islands, Great Salt Lake, Utah, p. 247. Breeding biology of the Double-crested Cormo- rant on Utah Lake, p. 1. Brennan, J. M., and M. L. Goff, article bv, p. 501. Brvmson, John T., Richard N. Seaman, and Don- ald J. Nash, article by, p. 77. Carebareia, p. 507. Carebareia johnstoni, p. 507. Cattail decline at Farmington Bay Waterfowl Management Area, p. 24. Cheng, Mou-Sheng, and Samuel R. Rushforth, article by, p. 402. Clark, Stephen L., article bv, p. 100. Clark, Tim W., article by, p. 91. Cnesinus prominulus,, p. 212. Conophthorus conicolens, p. 212. Cox, Douglas C, and Wilmer W. Tanner, ar- ticle by, p. 35. Croft, B. A., and C. D. Jorgensen, article by, p. 366. Crotaphytus wislizei maculosus, p. 230. Dempster, Lauramay T., and Friedrich Ehren- dorfer, article by, p. 510. Distributional patterns of mammals in Utah, p. 457. Fames, Michael, Don C. Erman, and Kenneth B. Roby, article by, p. 57. Egoscue, Harold J., article by, p. 75. Ehrendorfer, Friedrich, and Lauramay T. Dempster, article by, p. 510. Food of Townsend ground squirrels on the .\rid Land Ecologv Reserve (Washington), p. 128. Forcella, F., and T. Weaver, article by, p. 395. Gagne, Raymond J., article by, p. 426. Geographic variation in Contopiis sordidulus and C. virens north of Mexico, p. 453. Gilland, James K., and J. Keith Rigby, article bv, p. 475. Goff', M. L., and J. M. Brennan, article by, p. 501. Guimaraes, Lindolpho R., article by, p. 221. Gut, Larry J., Robert A. Schlising, and Carol E. Stopher, article by, p. 523. Hatch, Stephan L., article by, p. 530. Home range of the western jumping mouse, Zapus princeps, in the Colorado Rocky Mountains, p. 87. Host-induced morphological variations in the strigeoid trematode Posthodiplostoinum min- imum (Trematoda: Diplostomatidae). II. Body measurements and integument, p. 129. Host-induced morphological variations in the strigeoid trematode Postfwdiplostotnum 7nin- imum (Trematoda: Diplostomatidae). III. Or- gans of attachment, p. 375. Host-induced morphological variations in the strigeoid trematode Posthodiplostomum min- 534 GREAT BASIN NATURALIST Vol. 37, No. 4 imum (Trematoda: Diplostomatidae). IV. Or- gans of reproduction (ovary and testes), vitel- line gland, and egg, p. 481. Hydrological features of a California coastal fen, p. 57. Interpopulational variation of blood proteins in pika {Ochotona princeps saxatilis), p. 77. Introduced and exported American Scolytidae (Coleoptera), p. 67. Johnson, Mark K., article by, p. 128. Jorgensen, C. D., and B. A. Croft, article by, p. 366. Knopf, F. L., and V. C. Bachman, article by, p. 247. Lawson, C. L., article by, p. 532. Lepidium montanum var. stellae, p. 334. Lesquerella hemiphysaria var. lucens, p. 338. Loganiella johnsoni,, p. 477. Loveless, Earl C, and Carlos Y. Roby, article by, p. 259. Microcorthijlus vicinus, p. 213. Minshall, G. Wayne, and Robert L. Newell, ar- ticle by, p. 253. Mitchell, Ronald M., article by, p. 1. Murphy, Joseph R., and Neil D. Woffinden, ar- ticle by, p. 411. Nambudiri, E. M. V., and William D. Tiwell, article by, p. 241. Nash, Donald J., John T. Brunson, and Richard N. Seaman, article by, p. 77. Nectar-sugar concentrations and flower visitors in the western Great Basin, p. 523. New grass distribution records for New Mexico and the United States, p. 530. New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part IV, p. 207. New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part V, p. 383. New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part VI, p. 511. Newell, Robert L. and G. Wayne Minshall, ar- ticle by, p. 253. Nixon, E. S., article by, p. 97. Oberndorfer, Reed Y., and Kenneth W. Stewart, article by, p. 260. Observations on the distribution of micro- organisms in desert soil, p. 81. Occurrence of ticks on mule deer in central Utah, p. 407. Osteology and myology of the head and neck regions of Callisaurus, Cophosaurus, Hol- brookia, and Uma (Reptilia: Iguanidae), p. 35. Palmieri, James R., articles by, p. 129, 375, 481. Pederson, Jordan C, article by, p. 407. Petty, Jerold L., article by, p. 105. Phlocotrihtis amplus, p. 390. Phloeotrihm fici, p. 393. Phleotribus hirtus, p. 393. Phleotribus ingae, p. 389. Phleotribus levis, p. 392. Phloetribus minor, p. 391. Phleotribus nebulosus, p. 390. Phleotribus remorsus, p. 391. Phleotribus simplicidens, p. 389. Phleotribus squamiger, p. 392. Phleotribus tetricus, p. 389. Phleotribus vesculus, p. 392. Pityophthorus alnicolens, p. 213. Pityophthorus assistus, p. 214. Pityophthorus conicinnus, p. 214. Pityophthorus erotonis, p. 517. Pityophthorus discretus, p. 394. Pityophthorus indigens, p. 214. Pityophthorus leechi, p. 215. Pityophthorus solatus, p. 215. Pityophthorus speciosus, p. 215. Plants of Arizona: a new species of Astragalus from the Kaibab Plateau, p. 103. Population dynamics of the Ferruginous Hawk during a prey decline, p. 411. Preliminary mammal survey of Fossil Butte Na- tional Monument, Wyoming, p. 91. Pseudopityophthorus denticulus, p. 216. Pseudothysanoes securus, p. 216. Pseudothysanoes turnbowi, p. 217. Pycnarthrum fulgidum, p. 217. Quadraseta falconensis, p. 505. Quadraseta mirandae, p. 504. Quadraseta rotstieri, p. 503. Quadraseta tachirensis, p. 501. Range extension of Trionyx spiniferus emoryi into Utah (Reptilia), p. 259. Reveal, James L., and Stanley L. Welsh, article by, p. 279. Rhopalomyia subhimilis, p. 426. Rigby, J. Keith, and James K. Gilland, article by, p. 475. Roby, Carlos Y., and Earl C. Loveless, article by, p. 259. Roby, Kenneth B., Don C. Erman, and Michael Eames, article by, p. 57. Rushforth, Samuel R., and Cheng Mou-Sheng, article by, p. 402. Sangster, Mary E., article by, p. 274. Schlising, Robert A., Larry J. Gut, and Carol E. Stopher, article by, p. 523. Seaman, Richard N., John T. Brunson, and Don- ald J. Nash, article by, p. 77. Scolytodes canaliculus, p. 517. Scohjtodes constrictus, p. 518. Scolytodes contractus, p. 518. Scohjtodes festus, p. 519. Scolytodes fulmineus, p. 217. Scolytodes jucundus, p. 519. Scolytodes opacus, p. 520. Scolytodes opimus, p. 520. Scolytodes praeceps, p. 521. Scolytodes radiatus, p. 218. Scolytodes serenus, p. 521. Scolytodes suturalis, p. 521. Dec. 1977 535 Scolytodes varius, p. 522. Shapiro, Arthur M., article by, p. 443. Spring waterfowl migration in the Uinta Basin of northeastern Utah, p. 274. Stewart, Kenneth W., and Reed Y. Obemdorfer, article by, p. 260. Stilt Sandpiper and Hudsonian Godwit in Ne- vada, p. 532. Stinson, Nathaniel, Jr., article by, p. 87. Stopher, Carol E., Larry J. Gut, and Robert A. Schlising, article by, p. 523. Supplementary note on Venezuelan bat flies (Diptera: Nycteribiidae), p. 221. Syzugioxylon chhindivarense, a new fossil wood from the Deccan Intertrappean series of In- dia, p. 241. Syzygioxylon chhindwarense, p. 241. Tanner, Wilmer W., and Benjamin H. Banta, ar- ticle by, p. 225. Tanner, Wilmer W., and Douglas C Cox, ar- ticle by, p. 35. The algal flora of the campus of Brigham Young University, Provo, Utah, p. 402. The alpine butterflies of Castle Peak, Nevada County, California, p. 443. The influence of predator control on two adja- cent wintering deer herds, p. 101. The life cycle of Hydroperla crosbyi (Plecop- tera: Perlodidae), p. 260. The sagebrvish vole flea, Megabothris clantoni princei, in western Utah, with comments on the distribution of Megabothris in the Bonne- ville Basin, p. 75. The subfamily Anomiopsyllinae (Hystri- chopsyllidae: Siphonaptera). I. A revision of the genus Anomiopsyllns Baker, p. 138. The systematics of Crotaphytus wislizeni, the leopard lizards. Part III. The leopard lizards of the Great Basin and adjoining areas, with a description of a new subspecies from the Lahontan Basin, p. 225. Thelypodiopsis argiltacea, p. 95. Thehjpodhirn sagittuium var. ovalifolium, p. 358. Thome, Kave H., and Stanlev L. Welsh, article by, p. 103. Tidwell, William D., and E. M. V. Nambudiri, article by, p. 241. Tipton, V. J., A. M. Barnes, and J. A. Wildie, article by, p. 138. Tolle, Duane A., article by, p. 489. Typhlodromus macgregori Chant (.\carina: Phytoseiidae) and its control of phytophagous mites in Utah and southern California apple orchards, p. 366. Umcss, Philip J., Dennis D. Austin, and Michael L. Wolfe, article by, p. 101. Utah flora: Brassicaceae (Cruciferae), p. 279. Vollmer, A. T., F. Au, and S. A. Bamberg, ar- ticle by, p. 81. Weaver, T. and F. Forcella, article by, p. 395. Welsh, Stanley L., and Kave H. Thome, article by, p. 103. Welsh, Stanlev L., and N. Duane Atwood, ar- ticle by, p. '95. Welsh, Stanlev L., and James L. Reveal, article by, p. 279.' Wildie, J. A., A. M. Barnes, and V. J. Tipton, article by, p. 138. Woffinden, Neil D., and Joseph R. Murphy, ar- ticle by, p. 411. Wolfe, Michael L., Dennis D. Austin, and Phil- ip J. Urness, article by, p. 101. Wood, Stephen L., articles by, p. 67, Xyleborimis tribuloides, p. 218. Xylehorus devexus, p. 219. Xyleborus laticeps, p. 219. 383, 511. The Great Basin Naturalist VOLUME 37, 1977 Editor: Stephen L. Wood Published at Brigham Young University, by Brigham Young University TABLE OF CONTENTS Volume 37 Number 1 - March 31, 1977 Breeding biology of the Double-crested Cormorant on Utah Lake. Ronald M. Mit- chell 1 Cattail decline at Farmington Bay Waterfowl Management Area. Christine Minton Anderson 24 NOTICE TO CONTRIBUTORS Original mamiscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Xatiirali.st should be di- rected to Brigham Young University, Stephen L. Wood, Editor, Great Basin Xaturalist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be sim- ilarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. They should be typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. Use a recent issue of either journal as a format, and the Council of Biolo^ij Edi- tors Style Manual, Third Edition (AIBS 1972) in preparing the manuscript. An abstract, about 3 percent as long as the text, but not exceeding 200 words, written in accordance with Biological Abstracts guidelines, should precede the introductory paragraph of each ar- ticle. Authors may recommend one or two reviewers for their article. All manuscripts re- ceive a critical peer review by specialists in the subject area of the manuscript under con- sideration. Manuscripts that are accepted and that are less than 100 printed pages in length will automatically be assigned to the Great Basin Naturalist. Those manuscripts larger than 100 printed pages in length will be considered for the Memoirs series. Illustrations and Tables. All illustrations and tables should be made with a view toward having them appear within the limits of the printed page. Illustrations that form part of an article should accompany the manuscript. Illustrations should be prepared for reduction by the printer to either a single-column (2 5/8 inches) or double-column (5 1/2 inches) width, with the length not exceeding 7 1/2 inches. Costs Borne by Contributor. Contributors to the Great Basin Naturalist should be pre- pared to donate from $10 to $.30 per printed page toward publication of their article (in addition to reprint costs outlined in the schedule below). Authors publishing in the Great Basin Naturalist Memoirs may be expected to contribute $35 per printed page in addition to the cost of printed copies they purchase. No printed copies are furnished free of charge. A price list for reprints and an order form are sent with the galley proof to contributors. Reprint Schedule of the Great Basin Naturalist 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each 100 copies $20 $24 $28 $32 $36 $40 additional 200 copies 28 .32 36 40 44 48 2 pp. 300 copies 36 40 44 48 52 56 $4 Great Basin Naturalist Memoirs No. 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodbury, H. H. Frost. $10. No. 2 Intermountain biogeography: A symposium. By K. T. Harper, J. U. Reveal, el al. $15. TABLE OF CONTENTS Population dynamics of the Ferruginous Hawk during a prey decline. Neil D. Woffinden and Joseph R. Murphy 411 A new species of Rhopalomyia (Diptera: Cecidomyiidae) from Artemisia liidovi- ciana Nuttall (Compositae) in Utah. Raymond J. Gagne 426 Bionomics of a subterranean gall midge (Diptera: Cecidomyiidae) from Artemisia ludoviciana. M. A. S. K. Ranasinghe 429 The alpine butterflies of Castle Peak, Nevada County, California. Arthur M. Sha- piro 443 Geographic variation in Contopus sordidulus and C. virens north of Mexico. M. Ralph Browning 453 Distributional patterns of mammals in Utah. David M. Armstrong 457 A new fossil sponge from the Ordovician Garden City limestone of southeastern Idaho. J. Keith Rigby and James K. Gilland 475 Host-induced morphological variations in the strigeoid trematode Post- hodiplostornum rninimtim (Trematoda: Diplostomatidae). IV. Organs of re- production (ovary and testes), vitelline gland, and egg. James R. Palmieri ... 481 A survey of breeding and migratory birds southwest of Farmington, New Mexico. Duane A. Tolle 489 A new monotypic genus of chiggers and four new species of Quadraseta from Venezuela (Acari: Trombiculidae). M. L. G off and J. M. Brennan 501 A new name for Galium scahriusculum (Rubiaceae) of Utah. Lauramay T. Demps- ter and Friedrich Ehrendorfer 510 New synonymy and new species of American bark beetles (Coleoptera: Scoly- tidae), Part VI. Stephen L. Wood 511 Nectar-sugar concentrations and flower visitors in the western Great Basin. Larry J. Gut, Robert A. Schlising, and Carol E. Stopher 523 New grass distribution records for New Mexico and the United States. Stephan L. Hatch 530 Stilt Sandpiper and Hudsonian Godwit in Nevada. C. S. Lawson 532 Index to Volume 37 533 Date Due cflcme Bookbinding Co., Inc. 300 Summer Street Bwton, Mass. 02210 2044 072 231 038