3m HARVARD UNIVERSITY Library of the Museum of Comparative Zoology The Great Basin Naturalist VOLUME 35, 1975 Editor: STi:i'Hr.N L. W( Published at Brigham Young University, by Brigham Young University TABLE OF CONTENl S Volume 36 Number 1 - March 31. 1975 Evolution of the sceloporine lizards (Iguaniclae). Kenneth R. Larsen and Wilmer W. Tanner 1 New synonymy and new species of American bark beetles (Coleop- tera: Scolytidae) . Stephen L. Wood 21 Genetics, environment, and subspecies differences: the case of Polites sabuleti (Lepidoptera: Hesperiidae). Arthur M. Shapiro 33 Life history and ecology of Megarcys signata (Plecoptera: Perlodidae), Mill Creek, Wasatch Mountains, Utah. Mary R. Gather and Arden R. Gaufin 39 Records of stoneflies (Plecoptera) from Nevada. Marv R. Gather, Bill P. Stark, Arden R. Gaufin ' 49 Growth of Plecoptera (stonefly) nymphs at constant, abnormally high temperatures. Joseph M. Branham, Arden R. Gaufin, and Robbin L. Traver 51 Water balance and fluid consumption in the southern grasshopper m.ouse, Onychomys torridus. Vernon C. Bleich and Orlando A. Schwartz ... 62 A systematic study of Coenia and Paracoenia (Diptera: Ephvdridae). Wayne N. Mathis 1 65 Environmental factors in relation to the salt content of Salicornio pa- cifica var. utahensis. D. J. Hansen and D. J. Weber 86 New records of stoneflies (Plecoptera) from New Mexico. Bill P. Stark, Theodore A. Wolff, and Arden R. Gaufin 97 The authorship and date of publication of Siren intermedia (Amphibia: Caudata). Hobart M. Smith, Rozella B. Smith, and H. Lewds Sawin ..-. 100 New mites from the Yampa Valley (Acarina: Cryptostigmata: Ori- batulidae, Passalizetidae). Harold G. Higgins and Tvler A. Woolley ■ 103 The identity of Boucourt's lizard Eunieces capito 1879. Hobart M. Smith, Rozella B. Smith, and Jean Guibe 109 Studies in nearctic desert sand dune Orthoptera. Part XV. Eremog- raphy of Spaniacris with biological notes. Ernest R. Tinkham 113 Roosting behavior of male Euderma maculatum from Utah. Richard M. Poche and George A. Ruffner 121 The nest and larva of Diploplectron hrunneipes (Cresson) Hymenop- tera: Sphecidae). Howard E. Evans 123 Number 2 - .Tune 30, 1975 A revision of the Phacelia Crenulatae group (Hydrophyllaceae) for North America. N. Duane Atwood 127 Rodent populations, biomass, and community relationships in Arte- misia tridentata. Rush Vallev, Utah. D. W. Nichols, H. D. Smith, and M. F. Baker ." 191 Computerized reduction of meteorologic measurements from irrigated and nonirrigated plots in central Utah. Ferron L. Andersen and Paul R. Roper - 203 3-^ ' Clarence Cottani, 1899-1974; a distinguished alumnus of Brigham Young University. Vasco M. Tanner 231 Evolutionary divergence in closely related populations of Mimulus guttatus (Scrophulariaceae). Karen W. Hughes and Robert W. Vickery, Jr 240 Number 3 - September 30, 1975 Urosaurus and its phylogenetic relationship to Uta as determined by osteolog}' and myology (Reptilia: Iguanidae). Charles Fanghella, David F. Avery, and Wilmer W. Tanner 245 Distribution and adundance of the black-billed magpie {Pica pica) in North America. Carl E. Bock and Larry W. Lepthien 269 Nectar composition of hawkmoth-visited species of Oenothera (Ona- graceae). Robert E. Stockhouse, II 273 A revision of the nearctic species of Clinohelea Kieffer (Diptera: Cera- topogonidae). William L. Grogan, Jr., and Willis W. Wirth 275 Basidiomycetes that decay junipers in Arizona. R. L. Gilbertson and J. P. Lindsey 288 Body size, organ size, and sex ratios in adult and yearling Belding ground squirrels. Martin L. Morton and Robert J. Parmer 305 Photoperiodic responses of phenologically aberrant populations of pie- rid butterflies (Lepidoptera). Arthur M. Shapiro 310 Additional records of reptiles from Jalisco, Mexico. Philip A. Medica, Rudolf G. Arndt, and James R. Dixon 317 Invasion of big sagebrush {Artemisia tridentata) by white fir {Abies concolor) on the southeastern slopes of the Warner Mountains, California. Thomas R. Vale 319 Morpholog}' of ephemeral and persistent leaves of three subspecies of big sagebrush grown in a uniform environment. W. T. McDonough, R. 0. Harniss, and R. B. Campbell 325 Number 4 - December 31, 1975 Endangered, threatened, extinct, endemic, and rare or restricted Utah vascular plants. Stanley L. Welsh, N. Duane Atwood, and James L. Reveal ". 327 Utah plant novelties in Cymopterus and Penstemon. Stanley L. Welsh.... 377 The Zygoptera (Odonata) of Utah with notes on their biology. A. B. Provonsha :. 379 New synonymy and new species of American bark beetles (Coleop- tera: Scolytidae), Part II. Stephen L. Wood 391 Correlates of burrow location in Beechey ground squirrels. Donald H. Owings and Mark Borchert 402 Arachnids as ecological indicators. Dorald M. Allred 405 Notes on the genus Bomhylius Linnaeus in Utah, with key and descrip- tions of new species (Diptera: Bombyliidae). D. Elmer Johnson and Lucile Maughan Johnson 407 Breeding range expansion of the starling in Utah. Dwight G. Smith .... 419 Some parasites of paddlefish {Polydon spathula) from the Yellowstone River, Montana. Lawrence L. Lockard and R. Randall Parsons .... 425 Reproductive cycle of the Belding ground squirrel [Sperrnnphilus hel- dingi) : seasonal and age differences. Martin L. Morton and John S. Gallup 427 A new combination in Penstemon (Scrophulariaceae). Stephen L. Clark.. 434 Some relationships between water fertility and egg production in brown trout (Salmo trutta) from Montana streams. Lawrence L. Lockard.. 435 Some relationships between internal parasites and brown trout from Montana streams. Lawrence L. Lockard, R. Randall Parsons, and Barry M. Schaplow 442 Sexual dimorphism in malpighian tubules of Pteronarcys californica Newport (Plecoptera) . Ralph R. Hathaway 449 New records of the bat Plecotus phyllotis from Utah. Richard M. Poche 451 € GREAT BASIN NATURALISl ime35Na1 March 31, 1975 Brigham Young Universit: MUQ. COMP. ZOOU L.JBRARY JUN 1 1 1975 HARVARD UNIVERSITY i ^\ GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Zoology; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, dean. College of Biological and Agricultural Sciences; Ernest L. Olson, director, Brigham Young University Press, University Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been continuously published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only original, previously unpublished manuscripts pertaining to the biological natural history of the Great Basin and western North America will be accepted. Manuscripts are subject to the approval of the editor. Subscriptions. The annual subscription is $9 (outside the United States $10). The price for single numbers is $3 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 Press, Marketing Department, 204 UPB, Provo, Utah 84602. Scholarly Exchanges. Libraries or other organizations interested in obtaining this 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 the Great Basin Naturalist should be addressed to the editor as instructed on the back cover. The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University Volume 35 March 31, 1975 No. 1 EVOLUTION OF THE SCELOPORINE LIZARDS (IGUANIDAE) Kenneth R. Larsen^ and Wilmer W. Tanner^ Abstract. — Phylogenetic relationships among Sceloporine genera are briefly discussed. Species re- lationships witliin the genus Sceloporus are analyzed, and evolutionary lines of descent are proposed. The genus Sceloporus is composed of three monophyletic groups: Group I, the most primitive, prob- ably developed from Salor-\\ke ancestral stock in Miocene times. This group speciated from stock similar to Sceloporus gadoviae in southern Mexico to S. merriami in the North and contains 7 species in 3 species groups. We propose that these species be included in the genus Lysoptychus Cope. Group II arose from Group I and evolved from centrally located Sceloporus pictus in all directions throughout Mexico. This intennediate group contains approximately 19 species in 5 species groups. Group III also arose from the primitive stock of Group I and radiated from several desert refugia created by Pleistocene glaciation. Evolution of this group in Mexico was generally from north to south with Sceloporus malachiticus extending as far south as Panama. This group contains approximately 33 spe- cies in 5 species groups. In a previous paper (Larsen and Tan- ner, 1974) we presented our analysis of the species in the lizard genus Sceloporus. Numerical statistical methods were used to analyze the species in the genus Scel- oporus using cranial osteology, external meristic and numeric characters, karyol- ogy, display behavior, and geographic dis- tribution. A new classification for the genus was proposed with three major branches or groups. Group I contained 7 species in 3 species groups. Group II con- tained approximately 19 species in 5 spe- cies groups. Group III contained approxi- mately 33 species in 5 species groups. This classification was supported by the cluster analysis of several different sets of data. Cranial osteology, zoogeograph}', behavior, and karyology were shown to be taxon- omically significant as numeric charac- ters. Stepwise discriminate analysis showed that this classification of the spe- cies of Sceloporus into 3 major groups and 13 species groups was significant at the .999 confidence level. The purpose of this paper is to present our views on the evolution of the species in the genus Sceloporus. We also propose a ph3dogeny of closely related (Scelop- orine) genera. We are grateful for the assistance of H. M. Smith, C. C. Carpen- ter, W. P. Hall, and the following per- sons at Brigham Young University: A. L. Allen, F. L. Anderson, J. R. Murphy, M. S. Peterson, J. K. Rigby, N. M. Smith, D. A. White, and S. L. Wood. Intergeneric Phylogeny In 1828 Weigmann described several genera, including Sceloporus (S. torqua- tus) . He distinguished Sceloporus from the South American Tropidurus mainly on the basis of femoral pores (S'c^j/o^ thigh, porus=\)OYe) . In 1852 Baird and Girard described the genus Uta (U. stansburiana) which is distinguished from the smaller species of Sceloporus by its gular fold and granular dorsal scales. In 1854 Hallowell erected the genus Urosaurus (U. gracio- sus), which is similar to Uta but has sev- eral rows of enlarged, carinate, imbricate vertebrals or paravertebrals. Two years later Dimieril (1856) described the genus Phymatolepis (Urosaurus bicarinatus) on the basis of enlarged paravertebrals. In 1859 Baird placed Hallowell's genus Uro- saurus in synonymy with Uta, and in V07 North 500 West, Provo, Utah 84601 . -Department of Zoology, Brigham Young Universitj-. Provo. Utah 84602. GREAT BASIN NATURALIST Vol. 35, No. 1 1864 Cope did the same with Dmneril's Phymatolcpis. Boulenger (1885) raised Cope's Uta thalassina to generic status (Petrosauriis) , but Cope (1900) rejected this proposal and made Petrosaurus a third synon;y^n of Uta. In 1888 Cope erected the genus LrsoptycJius (L. Iateralus^= Sceloporus couchi) on the basis of a single specimen that appeared to have a well- developed gular fold. Subsequent investi- gation (Stejneger, 1904) showed the "gu- lar fold" to be an artifact of preparation on a single specimen which "was pre- served in such a manner as to make a fold across the neck, which formed the basis for the erection of the genus" (Smith, 1939, p. 242). Dickerson (1919) de- scribed the genus Sator (S. grandaevus) which has persisted despite Sator's close similarity to Uta, Urosaurus and Scel- oporus. In 1942 Mittleman resurrected the genera Urosaurus and Petrosaurus. He also erected the genus Streptosaurus based on Uta mearnsi, which is most similar to Petrosawus. He proposed that Uta, Uro- saurus, and Sator all arose independently from Sceloporus. He placed PJirynosoma with the above genera in a distinct group. Smith (1946) moved Sauromalus and Dipsosaurus to more primitive positions but otherwise retained Mittleman's ar- rangement. Savage (1958) placed Strept- osaurus in synonymy with Petrosaurus. He separated Uta from Urosaurus mainly on the basis of sternal and costal mor- phology. He placed Uta and Petrosaurus with the sand lizards (Holhrookia, Unia, and Callisaurus) , leaving Sceloporus, Sa- SCEIOPORUS GROUP III SCELOPORUS \ GROUP II COPHOSAURUS HOIBROOKIA \/ \ /►'^ElOPORUS / GROUP 1 Y SATOR UMA \ ~\ /^^CALIISAURUS 1 ^/ /Urosaurus PHRYNOSOMA^"-\^ "'* \ / ^x petrosaurus ANCESTRAL STOCK Fig. 1. Phylogeny of sceloporine genera and the three major groups in Sceloporus. tor, and Urosaurus together. Etheridge (1964) rejected Savage's wide separation of Uta and Urosaurus, and placed Uta, Urosaurus, Sator, and Sceloporus on one side and Uma, Holbrookia, and Callisau- rus on the other. Primitive to both groups was Petrosaurus. A sand lizard resur- rected by Clarke (1965) was Troschel's (1852) genus Cophosaurus (C. texanus, previously Holbrookia texana) . Presch (1969) rejected Etheridge's re- moval of PJirynosoma from the scelopor- ines and placed Phrynosoma with the sand lizards as a primitive member of that group. On the basis of scleral ossicles, Presch (1970) indicated that Petrosaurus is a primitive member of the Sceloporus branch. Ballinger and Tinkle (1972) pro- posed an early separation of the Uta and Petrosaurus stock from the ancestor of Urosaurus, Sator, and Sceloporus. Several characters suggest further modi- fication of the above arrangement. Our jiroposed phylogeny of sceloporine genera is illustrated in Figure 1. Urosaurus shows a tendency for enlarged scales near the midline of the dorsum. This trend is further developed in Sator, which has en- larged dorsals and granular laterals. The migration of enlarged scales around the sides of the body and the increase in scale size and degree of imbrication, mucrona- tion, and carination is a general trend along the chain of genera from Petrosau- rus to Sceloporus. The new phylogeny is also supported by the gradual decrease in development of the gular fold, which is completely lost in all species of Sceloporus in Group III. Most of the species in Group I have what Smith (1939) called a rudimentary gular fold. Some of the species in Group II show a less pro- nounced tendency to develop a gular fold, and Group III lacks it completely. The gradual loss of the gular fold in the Sceloporus complex is more probable than a loss (from Petrosaurus to Sceloporus) and subsequent redevelopment (from Sceloporus to Uta, Urosaurus, or Sator). This reversal of the phylogeny resolves a question raised by Smith (1946:178): "It is a curious fact that all genera that have sprung from Sceloporus have developed a gular fold — including Sator, a Baja Cali- fornia genus. The tendency to develop this fold a})pears to be restricted to the ])rimitive groups of Sceloporus . . . and these are the groups from which Uta, March 1975 LARSEN, TANNER: SCELOPORINE LIZARDS Urosaurus, and Sator independently ap- pear to have been derived." Although Smith pointed to this prob- lem, he nevertheless accepted Mittleman's arrangement of the sceloporine genera. More recently, Smith (per comm.): has agreed that Sceloporus ma}' be derived with respect to Uta^ Urosaurus. and Sator. This position has also been suggested by Hall (pers. comm.): "Inspection of the structure of the femoral pores and their surrounding scales, and the development of mucronation and carination of the body scales, to mention but two sets of charac- ters in various primitive Sceloporus and in other sceloporine genera, will suggest that Sceloporus is derived even in respect to Uta and Urosaurus. '' We suggest the following conclusions with regard to the new phylogeny and published data on hip ratios of displaying males (Purdue and Carpenter, 1972a, 1972b). The hip ratio (vertical hip move- ment to vertical eye movement) increased from Petrosaurus (0.68) to Uta (average 0.74) to Urosaurus (average 1.06). After the transition from Sator (no published data on hip ratios) to Sceloporus, the trend reversed and hip ratios decreased from an average of 1.21 in Group I to 0.66 in Group II to 0.34 in Group III (averages computed from Purdue and Carpenter, 1972b). Etheridge (1964) illustrated clavicles and scapulocoracoids of 8 sceloporine genera (excluding Phrynosoma) . If his drawings are superimposed on the new phylogeny (Fig. 2), two trends are ap- parent: (1) a gradual development of the scapular fenestra (top groove) from Petrosaurus to Sceloporus Group III, and (2) an increase in size of the clavicular hook. If Urosaurus and Uta were derived from Sceloporus, the scapular fenestra would have developed and then disap- peared from Petrosaurus to Sceloporus to Uta. This improbable reversal is similar to the problem with the gular fold. We are persuaded that the new phylogeny is more probable. Intrageneric Phylogeny The first ph^dogenetic schemes for the genus Sceloporus were proposed by Smith (1934, 1937a, 1937b, 1938, 1939). Other workers have recently modified the phy- logeny on the basis of karyology (Cole, 1970, 1971a, 1971b; Hall, 1971, 1973), and behavior (Bussjaeger, 1971). Larsen and Tanner (1974) redefined relationships among the species in the genus Sceloporus. We used Ward's clus- ter analysis (Wishart, 1968) to cluster 55 species on the basis of external characters, cranial osteology, karyology, behavior, and zoogeography (Fig. 3). We then used step- wise discriminate analysis (Dixon 1967) and found that the arrangement of groups and subgroups is significant at the .999 level of confidence (Table 1). Although Ward's cluster analysis pro- vides a phenetic dendogram, it does not give any indication as to which branch of a cluster is derived and which is primi- tive. In 1939 Smith said, "The most primitive form of this group is undoubted- ly lunaei which is closely related to for- mosus malachiticus'' (p. 60). In other words, lunaei is the most primitive form PETROSAURUS Fig. 2. Clavicles and scapulocoracoids of sev- eral sceloporines. All illustrations except Scelo- porus I, Sceloporus II, and Sceloporus III are from Etheridge (1964). 4 GREAT BASIN NATURALIST Vol. 35, No. 1 Tabt.k 1. — Groups and subgroups in the genus Sceloporus Group I (7 spp ) Group II (20 spp.) Group III (33 spp.) Subgroup A ( 1 spp.) Subgroup A (7 spp.) Subgroup A (9 spp.) gadoviae grammicus spinosus Subgroup B (2 spp.) pictus orcutti couchi megalepidurus clarki memami cryptus melanorhinus Subgroup C (4 spp.) shannonorum * magister maculosus heterolepis olivaceus parvus asper cautus jalapae Subgroup B (2 spp.) horridus ochotei-enae pyrocephalus edwardtaylori nelsoni Subgroup B (7 spp.) Subgroup C (3 spp.) formosus scalaris lunaei goldmani* nialachiticus aeneus acanthinus Subgroup D (4 spp.) Subgroup C (5 spp.) siniferus undulatus carinatus virgatus utifonnis woodi squamosus occidentalis Subgroup E (4 spp.) graciosus variabilis Subgroup D (4 spp.) cozumelae jarrovi teapensis lineolateralis chrysostictus ornatus dugesi Subgroup E (8 spp.) torquatus cyanogenys bulleri insignis* macdougalli mucronatus serrifer poinsetti * Species not examined in this study. ill the spinosus s])ecies group because it is most similar to a member of the next closest group (formosus). This statement by Smith is consistent with the following method of converting a phenetic dendro- gram into a phylogeny (Fig. 4): If "A" is primitive to "B" it is less derived from (more similar to) the stem species "G." The more primitive member of the other cluster ("C" or "D") will also be more similar to "G." The more primitive mem- bers of the two clusters will therefore be phylogenetically "closer" and phenotypi- cally more similar than any other com- bination from the two clusters. This rule can be applied objectively with a similar- ity matrix. When all possible pairs between adja- cent clusters are compared, the two most similar species are considered jjrimitive within their res|)ective clusters. This technique will convert a dendrogram into a phylogeny. Ward's cluster analysis and the above phylogeny techni(|ii(> were repeated sever- al times using external and osteological characters, distribution, karyology, be- havior, and combinations of the above. (See Larsen and Tanner, 1974, for a pre- sentation of results.) The differences among results were resolved subjectively to produce a composite phylogeny (Fig. 5). This ])rocedure is based on several assumptions which are admittedly vul- nerable. To restrict the scope of our study it was assumed that the alpha taxonomy is complete and correct. That is, it was assvimed that all species of Sceloporus are now named and correctly defined in the literature. Of course, this assumption may be incorrect. But the purpose of our study is to produce a general overview and not a detailed taxonomic review. The de- tails near the ends of branches are there- fore tonlativo and stibject to future re- view. In spite of the large number of charac- ters considered (over 80), these results are also subject to errors due to parallel- ism, convergence, varying rates of diver- March 1975 LARSEN, TANNER: SCELOPORINE LIZARDS Formosus __^ spinosus .^_^ Horndus _— Olivaceus Cqutus Adieri Molachiticus — Luna«i Lundalli ^^— Acantninus ~__ Edwprdtoylori. Orcutti —^— Magistcr Undulatus Occidentalis ^ Virgatus '— ^— Graciosus ^^— i Torquatus sernfer Mucronatus Cyonogenyi Bulleri Poinsetti jarrovi -^—— Linaolateralis- Ornotus Dugesi — — ^— Atper Heterolepis_ Grammicus Megolopidurus. Pictus ^^-^^^ Ochoterenae — Jalopae scolons Aeneus pyrocephalus- Nelsoni — - Melonorhinus. siniferus Connotui Utiformis Variabilis Cozumela* Teapensif '^^— Chrysostictus- squamosus^^ Parvus ^^.— Maculosus^— Couchi ■ MerriamI— Cadoviae ^^ ^ ^ ^ i ;=^ 0.5 16 Fig. 3. Dendrogram generated by Ward's cluster analysis of external, diaracters (82 characters). skull, and distribution gence, pleiotrophy, and other cases in which the phenotype is not a direct mani- festation of the genotype. All phylogene- tic conclusions are subject to these liinita- tions, and the systematist can do little more than acknowledge the circumstantial nature of his evidence. We propose that SceJoporus is derived from Uta through Urosaurus and Sator (see above). Smith (1938) suggested that tlie connection between these genera is from Urosaurus ornatus to Sceloporus couchi. Smith included couchi in the variabilis species group. Figure 6 shows the arrangement of species in Smith's variabilis, maculosus, and mcrrianii groups according to Smith (1939, Fig. 42) and the new phylogeny. Four of these species {couchi, parvus, maculosus, and merriami) are transferred to Group I. Smith may have allowed for this by placing these four species on one side of his tree next to Uta. If Uta {Uta, Urosaurus, and Sator) is considered primi- ti^'e to Sceloporus, then Smith's evidence supports our conclusion that Group I is primitive to the other two groups in Sceloporus. The remaining species in Smith's variabilis group {variabilis, coz- umelae, and teapensis) are placed in Group II. Smith (1939:239) allowed for the re- moval of parvus and couchi from the var- iabilis grouj) with this statement: That parvus and couchi are only dis- tantly related to the remainder of the group is shown by the widely different charac- ter of the ventral coloration in the males, smooth head scales, larger number of fem- oral pores, and general habitus. ... It is my belief that this section approaches more closely the ancestral stock of Uta than the other species of the variabilis group. Smith (p. 239) also associated merriami with Uta: "It w^ould appear that merri- ami is closely related to Uta. and that Uta GREAT BASIN NATURALIST Vol. 35, No. 1 B E 1 1 F 1 1 G IF A-C = l A-D=2 B-C = 2 B-D = 3 THEN-- Fig. 4. Phylogeny theory- K the phenetic distance between "A" and "C" is less than that between any other pair, then "A" and "C" are primitive members in clusters "E" and "F." arose from the forms now extinct which closed the present gap between couchi and merriami.'" Note that our new ar- rangement places merriami and couchi together. Another divergence from Smith's phylo- genetic tree is the addition of chrysostic- tus to the variabilis group. Smith (p. 239) supports this inclusion (and the close proximity of the siniferus group) : "An- other group close!)' related to the variab- ilis section is the siniferus series, which closely approaches the variabilis group through cuprous. . . . The chrysostictus group is also closely related." Thus it can be seen that Smith allowed for the possibility of removing parvus and couchi and adding chrysostictus, which changes his variabilis group into the new variabilis group. Smith stated that the siniferus group "closely approaches the variabilis group" and yet his illustration (1939, Fig. 3) has these groups separated by several other groups. In the new phylogeny they are adjacent. Figure 7 compares Smith's arrange- ment of his chrysostictus. utiformis and siniferus groups with the new arrange- ment of the same species. Besides the placing of chrysostictus in the variabilis group (which has already been ex- plained), the only major difference in Figure 4 is the removal of ochoterenae to place it in Group I. (The inclusion of utiformis in the siniferus group is mi- nor) . Smith listed 1 1 diagnostic characters of the siniferus group. In three cases he said "except ochoterenae'' and in another "except ochoterenae and cupreusT He (p. 301 ) said, "Postanals tending to be poorly developed (except ochoterenae and cupreus); two postrostrals (except ocho- terenae, without postrostrals) ; . . . ventral scales pointed or, at least not notched (ex- cept ochoterenae in which they are notched) . . . males without distinctive ventral coloration (except ochoterenae).'" If size is discounted, then ochoterenae is different in 4 of the 10 diagnostic char- acters for the siniferus group. S. ocho- terenae also has more femoral pores than any other species in Smith's siniferus group. Smith's conclusions, therefore, would not be seriously challenged if ochoterenae were removed from the sinif- erus group and placed in Group I next to jalapae. In fact, when describing ocho- terenae., Smith (p. 309) said, "three or four scales on anterior border of ear, not so large as in jalapae.'' So apparently he was comparing these two species. Smith included jalapae in his scalaris group, which is otherwise identical to the new scalaris group (Fig. 5). Removing jalapae from the scalaris group to place it in the primitive Group I is supported by the following statement b^- Smith (p. 331): The only species doubtfully inchuled in this group is jalapae, which differs from the remaining fonns in having lateral scales in distinctly oblique rows, and in lacking postrostrals [as does ochoterenae]. . . . 5. jalapae is clearly, the most primitive member of the group. S. scalaris, aeneus and goldmani are clearly more closely re- lated to each other than any one of these is to jalapae. March 1975 LARSEN, TANNER: SCELOPORINE LIZARDS HORRIDUS MELANORHINUS MALACHITICUS ACANTHINUS OCCIDENTALIS SCALARIS I UTirORMIS GOLDMANI VARIABILIS TEAPENSIS Fig. 5. Proposed phylogeny for the genus Sceloporus. (* = species not examined.) GREAT BASIN NATURALIST Vol. 35, No. 1 PARVUS MACULOSUS MERRIAMI TEAPENSIS COZUMELAE Fig. 6. Phylogeny of Smith's (1939) variabilis, maculosus. and merriami groups according to Smith (A) and the new phylogeny (B). CHRYSOSTICTUS VARIABILIS GROUP SCALARIS GROUP OGHOTERENAE CHRYSOSTIGTUS VARIABILIS GROUP SCALARIS GROUP OGHOTERENAE Fig. 7. Phylogeny of Smith's (1939) chrysostictus, utiformis, and siniferus groups according to Smith (A) and the new phylogeny (B). March 1975 LARSEN. TANNER: SCELOPORINE LIZARDS (iroup I includes: parvus, couchi, ma- ( ulosus, mcrriami, ochoterenat\ jalapae, aiul gadoviac. the most primitive. Smith ( p. 362) inchided gadoviac with nelsoni and pyrocephalus in the pyrocephalus group. But once again he outhned rea- sons why gadoviac could be removed and [)laced in Group I. "5. gadoviae differs widely from other members of the group in having very small dorsal scales, a large number of femoral pores, a postfemoral dermal pocket, very small scales on pos- terior surface of the thighs, and many other minor characters." S. gadoviae is also the only member of this group to have a vestigial gular fold as mentioned by Smith (p. 374): "scales immediately preceding gidar fold region somewhat re- duced in size." All of these characters are diagnostic of Group I, and this primi- tive placement is therefore natural. In fact. Smith (p. 363) said, "I assume gadoviae to be nearest the primitive type, as it retains certain characters of the variabilis group, from which I believe it was derived." The main character on which Smith (p. 363) based his inclusion of gadoviae with the pyrocephalus group is the strong com- pression of the tail: "That the group is a natural one is more or less assured by its compact range and by the common char- acter of the compressed tail, which is otherwise unknown in the genus." In view of the many characters supporting the placement of gadoviac in Group I, we propose that a compressed tail developed twice: once in the pyrocephalus group, and once in gadoviae. Smith (p. 363) gave further support to this placement of gadoviae: "The assumption that gadoviac is a remnant of a primitive stock is sup- ported by its secretive habits and its re- striction to a somewhat arid region." The most serious difference between the new phylogen^- and that of Smith is the placement of the gramniicus and me- galepidurus groups. In both phylogenies the species are arranged in a similar man- ner within these groups. But Smith placed these groups next to the jormosus group with the large-scaled, large-sized species, and we ha\e moved them to a primitive position in Group II. How- ever, we propose that the grammicus group (we have combined Smith's gram- micus and hetcrolcpis groups) is the most primitive in Group II. In fact, Smith (1938:552) said "the microlepidurus [our grammicus^ group is assumed to be the most primitive of these [the large-scaled, large-sized sjiecies], largely because of its very small scales." This greater separ- ation between the grammicus and jormos- us groups is further justified by the fact that the diploid number of chromosomes is 22 (derived) in the jormosus group and 32 (primitive) in the grammicus group. We propose, therefore, that some of, the similarities between grammicus and jor- mosus (coloration, dorsal-scale count, ovo- viviparity, and preference for an arboreal habitat) are a result of convergence as is true of gadoviae and the pyrocephalus group. The only remaining difference from Smith's jormosus group is his inclusion of asper, which we have moved to the gram- micus group. This move is justified by the fact that asper has 32 chromosomes, as do the other members of the grammicus group. If the grammicus grou]:) is re- moved from Smith's large-scaled, large- sized branch, the remaining species are the same as those included in Group III. This grouping (the omission of grammi- cus) was allowed by Smith (1938:552): The relatively small size of the species of the undulatus group must be assumed as a parallel development rather than a direct inlieritence of the small size of the ancestor in the variabilis group, for the close rela- tionship of the spinosus and undulatus groups cannot logically be disputed, nor is the close relationship of the spinosus, lor- qualus and formosus groups doubtful." Smith and Taylor (1950) included the following species within the undulatus group: undulatus, cautus, occidentalism and woodi. Since then, virgatus has been raised from subspecific to specific status (Cole, 1963). Smith (1939) placed fjrac/- osus adjacent to the undulatus group, so the only discrepanc}' between the two classifications is the placement of cautus, which we have moved to the spinosus group next to olivaceus. This mo^'ement is justified by the fact that there is a zone of intergradation between cautus and oli- vaceus (Hall, pers. comm.). Bussjaeger (1971:151) remarked: The relation of cautus and olivaceus and the undulatus group of Sceloporus has been questioned. Hall's data indicated that these two species were the same and limited data on their displays indicate that they are similar. If one accepts that they are syn- 10 GREAT BASIN NATURALIST Vol. 35, No. 1 onyms, then olivaceus (cautus) would be the connecting link between the spinosus and undulatus groups. However, rather than use these forms as a link between species groups, we have placed them together in the spijiosus group. Smith (1938:554) indicated that the torquatus group consited of 2 subgroups: "It appears that soon after the separation of the torquatus stock from the other groups of Sceloporus, there was a separa- tion into two divisions, one of which ex- hibited a tendency to develop small scales, the other large scales." We have recog- nized the small-scaled division as the jarrovii group. Figure 8 shows the phylogeny of the jarrovii group according to Smith (1938, Fig. 4) and the new arrangement. Al- though he placed lineolateralis further away from jarrovii in his diagram. Smith (p. 556) did say, "S". jarrovii appears to be most closely related to lineolateralis. From this species, or its ancestors, the re- maining species of the small-scaled divi- sion have obviously been derived." Figure 9 shows the phylogeny of the torquatus group according to Smith (1938, LINEOLATERALIS Fig. 8. Phylogeny of jarrovi group according to Smith (1938) (A) and the new phylogeny (B). Figs. 3-4) and the new arrangement. There seems to be little similarity here, except that torquatus is derived from serrifer, and poinsetti is derived from cyanogenys in both trees. Smith (1938: 555) raised a question about the ancestral position of serrifer: S. serrifer appears to be the oldest of the large-scaled species. The postulation that this species, which is one of the larger ones POINSETTI CYANOGENYS Fig. 9 Phylogeny of torquatus group according to Smith (1938) (A) and the new phylogeny (B). March 1975 LARSEN, TANNER: SCELOPORINE LIZARDS 11 of the genus, and one having large scales, is nearest to the ancestral type of the large- scaled division of the torquatus group may appear to be contradictory to the postula- tion that Sceloporus is derived from small species with small scales. However, my as- sumption seems to be justified by the fact that serrifer occupies a southern position on the periphery of the geographical area now occupied by the torquatus group. The reason for this paradox is that Smith assumed speciation in Group III was from south to north. The data in 1938 strongly supported this conclusion. Obviously, Smith did not believe that a peripheral location is necessarily primitive, because on the next page (556) he said, "S". mu- cronatus appears to be the nearest to the ancestral type of these three species {cy- anogenys, poinsetti and omiltemanus) de- spite the fact that it has larger scales than they. I so conclude because of its central- ized geographical position with relation to the area occupied by the other three forms." So the basic problems can be solved, and the trend is indeed from small to large size and small to large scales if this group was developed from north to south rather than south to north. Smith indi- cated a northward development from ser- rifer to torquatus to mucronatus to cyano- genys, and our phylogeny indicates a southward development from cyanogenys to mucronatus to serrifer to torquatus. An ancestral placement of cyanogenys is fur- ther supported by Smith (1939:209): "Species of this group are as a rule con- fined to rocky habitats. So far as I am aware, only cyanogenys tends to live on or near the ground." Thus, the new ]:)hylogeny indicates a trend in this group from small-sized, small-scaled ground dwellers to large-sized, large-scaled rock dwellers. With this reversal in direction, the remaining differences between the two phylogenies in Figtire 9 are negligible and the trends within this group fit the overall phylogeny of the genus. In the genus Sceloporus, the spinosus group has been the object of more system- atic study than any other. No less than four different phylogenetic trees have been proposed by Smith, Bussjaeger, Cole, and Hall. The confusion is further compound- ed by the fact that the spinosus group is the largest in number of species and sub- species. The four phylogenetic trees and our conclusions are presented in Figure 10. Smith (1939) included acanthinus, lunaei. and lundelli with this group. In 1950, he and Tavlor moved acanthinus Fig. 10. Phylogeny of spinosus group according to Smith (1939), Cole (1970), Bussjaeger (1971), Hall (pers. comm. 1973), and the new phylogeny (L and T). 12 GREAT BASIN NATURALIST Vol. 35, No. 1 and lunaei into the formosus group. How- ever, in 1939 Smith (p. 60) said, "The most primitive form of the group is un- doubtedly lunaei. which is closely related to formosus malachiticus. S. acanthinus is a near relative of lunaei. as is also lun- delli.'" It should therefore be acceptable to remove lundelli from the spinosus group and place it in the formosus group next to lunaei as we have done. Behavioral data also support this ar- rangement. Bussjaeger (1971:136) ob- served: The display-action-patterns of lundelli gaigei of the spinosus group and asper, acan- thinus acanthinus and a. lunaei of the formosus group were quite similar with peaked single units and multiple units. Sceloporus asper and lundelli seemed to share more elements. In his conclusions, Bussjaeger (p. 151) an- ticipated the new position of S. lundelli: The status of lundelli is questionable. . . . Its display-action-pattern was between acan- thinus and orcuiti; but the pattern was based on only one female. More data are needed to establish this species relationship. At present it should be left in the spinosus group, although it appears to be closer to the formosus group. Cole's (1970) phylogenetic tree would xiot allow the removal of lundelli from this group unless melanorhinus and clarki were placed elsewhere. Cole (p. 39, Fig. 17) showed how four centric fusions could change the melanorhinus-clarki karyotype into the typical pattern for this group. According to Cole's assumption that only fusions (i.e., no fissions) are possible, melanorhinus and clarki are primitive not only for this group, but also for the genus Sceloporus. and for the entire fam- ily Iguanidae! As demonstrated by Web- ster, Hall, and Williams (1972), chromo- somal evolution can occur by fission as well as fusion. We believe this is the only acceptable explanation for the karyo- type in melanorhinus and clarki. If fission is accepted as well as fusion, Cole's data provide support for our arrangement of orcutti, clarki. and melanorhinus. (They also confirm the primitive position of lundelli and permit its placement in the formosus group.) If clarki and melanorhinus are derived from orcutti and if lundelli is removed from the group, then the only difference between Cole's tree and ours is a minor shift in the position of edwardtaylori. The single remaining difference between Smith's tree and ours is the placement of edwardtaylori. The close relationship of edwardtaylori to spinosus and horridus has been proposed by Cole and also by Hall. The justification is that the species clustering on one side {olivaceus. cautus, edwardtaylori. spinosus. and horridus) all have 22 chromosomes, whereas orcutti has 34, magister has 26, and clarki and me- lanorhinus each have 40. Zoogeography The phylogeny of the genus Sceloporus can be considered with its present geo- graphical distribution to produce a theo- retical history of events in the speciation in this genus. We conclude that the an- cestral sceloporine was a tropical or sub- tropical lizard (as Smith reasoned) \vith a distribution somewhat matching the sub- tropical conditions of western America before the Madro-Tertiary revolution (Ballinger and Tinkle( 1972:^63). This dis- tribution was not restricted to southern Mexico, where Smith pro])osed the begin- ning of Sceloporus evolution, but covered a vast area in the western United States ex- tending as far north as Canada. Milstead (1960:76) said, "Formation of the western deserts is presumed to have begun in Miocene times and continued through Pliocene and into early Pleisto- cene times." Accordingly, the derivation of the Scelporine genera could have oc- curred in late Miocene and early Pliocene times during the development of the west- ern deserts (Ballinger and Tinkle, 1972). The formation of deserts trapped a mesic-adapted relict (Petrosaurus) in Baja California. The remaining sceloporine stock began adapting to the oncoming desert conditions with such characters as a lengthened, sinuous nasal passage and the behavior called "shimmy burial" (Stebbins, 1944). The separation of the generic lines of Uta, Urosaurus, Sator, and Sceloporus was accomplished during the initial stages of adaptation to desert conditions. As tropical conditions moved south- ward during middle and late Pliocene (Axelrod, 1948), the ancestral stock of Group I moved south almost as far as the Isthmus of Tehuantepec. Some popu- lations did not migrate, but remained and March 1975 LARSEN, TANNER: SCELOPORINE LIZARDS 13 adapted to more xeric conditions (Group III). The mountains of central and south- ern Mexico J)ro^ ided a barrier that sepa- rated the western Group I and eastern Group II populations. A relict genus (Sator) was isolated in Baja California at this time (Fig. 11). The subsequent de- velopment of Grou])s I and II was a mat- ter of adaptive radiation and centrifugal speciation (Brown, 1957). Figure 12 shows the routes of speciation in Group I. The eastern branch extended from gadoviac (in southern Michoacan, Guerrero, Morelos, southern Puebla, and northwestern Oaxaca) northward across the Oaxaca Upland, the Neovolcanic Pla- teau and into the Sierra Madre Oriental to parvus (in Nuevo Leon, southeastern Coahuila, San Luis Potosi, and Hidalgo). Speciation continued northward along the Sierra Madre Oriental to couchi (Nuevo Leon, eastern Coahuila, and southern Tex- as) and merriami (northern Coahuila and adjacent Texas). (Locality information in this discussion is from Smith and Tay- lor, 1950. Topographical terminology is from Raisz, 1964.) The second branch of Group I extended from parvus to jalapac (Veracruz, Pueb- la, and Oaxaca). This radiation then moved across the Mixtec Upland (along the northern border of Oaxaca) and north- ward along the western flank of the Sier- ra Madre del Sur (through Guerrero, Michoacan, Colima, and Jalisco) and Fig. 12. Speciation in Group I. further northward along the western flank of the Sierra Madre Occidental (through Nayarit and Sinaloa and into Durango) . The Durango populations became niaculo- sus, and most of the pathway is now oc- cupied by ochotcrenae. Figure 13 shows the initial radiation from the ancestral stock of Group II. This ancestral stock is now represented by pictus (in central Puebla and central western Veracruz). The first radiation involved four species in four directions: aencus to the north, pyrocephalus to the west, sinifcrus to the south, and cozumelac to the east. Subsequent radiation from these cen- ters is shown in Figure 14. Sceloporus aeneus (Puebla, Veracruz, Oaxaca, Hidal- go, Morelos, Mexico, (juanajuato, Micho- acan, and Jalisco) produced scalar is (ni Durango, Guanajuato, Hidalgo, Jalisco, Mexico, Michoacan, Puebla, and Zacate- cas). S. pyrocephalus (Guerrero, Michoa- can, and Colima) produced nrlsoni (in Chihuahua. Jalisco, Sinaloa, and NaA'arit). Fig. li. Isolation of early Sceloporus stocks response to desert formation in middle Pliocene. Fig. 13. Early radiation ui Group II. 14 GREAT BASIN NATURALIST Vol. 35, No. 1 Fig. 14. Second These two species occupy most of the western flank of the Sierra Madre Occi- dental. According to Hall, the separation of nelsoni and pyrocephalus occurs along a river in Nayarit (the Rio Grande de Santiago). Concerning this river, Hall (pers. comm., 1973; see also Hall, 1973: 115-125) said: Evidence from the fresh water fish fauna in the Rio Grande de Santiago (Salvador Contreras B.. pers. comm.) suggests that at one time this major river drained the greater part of the Mexican Plateau. Even now it is the outlet for Lake Chapala and the entire Rio Lenna e.xtending east as far as the western border of the Distrito Federal. Although rivers usually are not very effective natural barriers, the steep gradient of this river as it falls off the Plateau and the comparative narrowness of the costal plain probably would have made it an extremely effective barrier during the Pleistocene pluvial times, which would have provided ample opportunity for the splitting of the ^troio-nelsoni into two stocks. The southern speciation produced sin- iferus (in Oaxaca, Chiapas, and Guerre- ro), carinatus (in Chiapas), squamosus (along the Pacific slopes from Chiapas to Costa Rica), and utiformis (to the north along the Pacific slopes of Michoacan, Colima, Jalisco, Nayarit, and Sinaloa). The eastern branch to cozumclae (in the northern peninsular states of Yucatan and Quintana Roo) produced chrysostictus (in the entire Yucatan Peninsula), teapensis (in southern Veracruz, Tabasco, Cam- peche, Quintana Roo, northern Guate- mala, and British Honduras), and variabi- lis (which has developed subspecies along the Gulf Coast plain from south-central Texas, through Nuevo Leon, Tamaulipas, San Luis Potosi, Queretaro, Hidalgo, Tlax- cala, Puebla, and Veracruz, across the Isthmus of Tehuantepec, through Oaxaca and Chiapas, and into western Guatema- la). The central stock of Group II also pro- duced a second wave of speciation. A southern speciation from pictus produced cry plus in the Oaxaca highlands. A west- ern speciation resulted in asper (in the Si- erra Madre del Sur in Guerrero and Mi- choacan and extending as far north as the Sierra Madre Occidental in Nayarit). This branch also produced heterolepis in the coastal mountains of Jalisco. An eastern branch from pictus pro- duced megalepidurus in Northern Puebla on the eastern slopes of the Neovolcanic Plateau. The most recent derivation from the pictus stock is grammicus. This spe- cies has invaded most of the Plateau re- gions in Mexico. The distribution of grammicus is widespread, and Hall (1971) has suggested that there ma}' be as many as 6 cryptic species in the grammicus com- plex. Further discussion of this species must therefore be deferred .until the alpha taxonomy is more complete. March 1975 LARSEN, TANNER: SCELOPORINE LIZARDS 15 Speciation in Group III was more com- plex and probably more recent than in the others. Other workers have suggested that considerable speciation resulted from repeated glaciation in Pleistocene times (Savage, 1960; Ballinger and Tinkle, 1972). Each glacial period forced desert species into southern refugia from which they later speciated through adaptive radiation and centrifugal speciation. Group III remained originally in the north and adapted to the xeric conditions of the southwest during middle and late Pliocene, as did Uta and Urosaurus. Sub- sequent Pleistocene glaciation forced the desert-adapted populations into southern refugia with massive northern extinctions. The five refugia south of 30° latitude in- clude Baja California, the Sonoran Desert, the Mexican Plateau, the Gulf Coastal Plain, and Florida. Barriers include the Gulf of California, the Sierra Madre Oc- cidental, the Sierra Madre Oriental, and the Gulf of Mexico. Ballinger and Tinkle (1972) discussed the first three refugia in considerable detail with reference to the e^■olution of Uta. After each glacial period, the isolated populations expanded in all directions from their refugia. (A worldwide increase in rainfall would restrict the midlatitude deserts from both sides. A subsequent de- crease in rainfall would cause a movement of xeric conditions both northward and 'southward from a small latitudinal band.) Each southerly movement was preserved as the species adapted to subtropical con- ditions, but the northerly radiations would bo eliminated during the next glacial peri- od (southern rains could be tolerated bet- ter than northern snows) . Each invasion to the south required a secondary adaptation to the ancestral en- vironment. This explains why formosus has not yet lost a behavioral trait called "shimmy burial." Hall (pers. comm.; see also Hall 1973:99-102) said: One gathers from Cole's (1970) discus- sion that he uncritically accepts Smith's (1939) idea that the arboreal, tropical for- mosus group is primitive in the genus. Smith (pers. comm.) believed, not unreas- onably on the limited infoiTnation then available, that the closest primitive relatives of sceloporus were the South American trop- idurines (from which Weigmann separated Sceloporus), and that its close xeric adap- ted relatives (i.e. "Uta" =-- Petrosaurus, Urosaurus, and Uta) were derived from within tlie radiation of Sceloporus. The work of Savage (1958), Etheridge (1964), and Presch (1969) tends to refute this idea. . . . Furthermore, it is interesting to note that the behavioral trait of 'shimmy burial' . . . is also found in most other Sceloporines. . . . From this analysis, it would seem that all sceloporines above Petrosaurus at least primitively know how to use loose sand for escape and sleeping cover. It seems un- likely that this behavior would evolve in a supposedlj' primitive fomi like formosus, which lives in inountain rain forests where the lizards would rarely or never encounter a suitable substrate for shimmy burial. Its presence in this species probably indicates only that formosus has only very recently entered the rain forest habitat. On the other hand, shinnnj' burial would be selectively valuable to a species inhabiting dry plains or deserts where loose sand might fre- quently be the only cover available for escape or sleeping. This quotation explains why Smith (1939) and Cole (1970) proposed phylogenies from south to north. We propose a re- versal of these phylogenies, which means that most trends in Group III are from the north and that the Group III forms moved southward and adapted to a climate similar to the one in which the ancestors lived. The smaller size and greater isolation of Baja California have limiited the genetic potential of its populations. This has al- lowed continental species to move north from the Sonoran Desert and enter the peninsula to trap southern relicts (see Savage, 1960). Another possible explanation for relict species in Baja California is the separa- tion and westward drift of the peninsula in Miocene-Pliocene times. Concerning this movement, Moore and Buffington (p. 1241) said, "Therefore, from about 4 to 10 million years ago, during late Miocene and Pliocene times, a proto-Gulf of Cali- fornia existed. . . . The present cycle of spreading began about 4 million years ago." lanner (1966:191) stated that this same event could apply to the night snakes: Thus the distribution of Eridiphus stock may have reached southern Baja California by a shorter route before the present Gulf of California was formed. Assuming this to be correct, Eridiphus is a relic of a once more widespread group of snakes in West- ern Me.xico. Hall (1973) has suggested that such a mechanism is responsible for speciation 16 GREAT BASIN NATURALIST Vol. 35, No. 1 in Baja California and that the Cape re- gion was isolated from the rest of the peninsula as well as the mainland during an intermediate stage. The first glacial advance divided Scelo- porus into four refugia: an orcutti stock in Baja California, a formosus stock in the Sonoran Desert, a virgatus stock on the Mexican Plateau and a cyanogenys stock on the Gulf Coastal Plain. Subse- quent postglacial speciation is illustrated in Figure 15. The virgatus stock expanded northward and as far eastward as Florida. It also ex- panded westward into the Sierra Madre Occidental. Most of the expansion from this stock was reduced to refugia during a second glacial advance. The second gla- cial advance was less severe than the first (Ballinger and Tmkle, 1972:63) and a population survived in Florida (ivoodi) . The main virgatus stock was again con- fined to the 5lexican Plateau, but some of the mountain ])opulations moved west into the Sonoran refuge. This isolation produced graciosus. The subsequent northward migration of graciosus and the northern speciation of undulatus and occidentalis from virgatus is shown in Figure 16. The orcutti stock, which was confined to the Baja California refuge during the first glaciation, emerged with sufficient adaptive specialization to displace the formosus stock as far south as Guerrero. The displacement of a mainland ])opula- tion by a restricted peninsular ])opulation is explained by the assumption that for- mosus descended from the part of the Sce- loporus stem that had been adapting to the mountain habitat between the central plains and the western deserts. As the Pacific slopes became more and more arid following glacial retreat, the desert-adapt- ed orcutti stock displaced the mountain- adapted fonnosus stock. From the Pacific slopes in Guerrero, the formosus stock speciated southward, pro- ducing formosus (with subspecies in Guer- rero and the central uplands of Oaxaca), malachiticus (along the Pacific slopes from Chiapas to Panama), lunaei (in the up- lands of central Guatemala), lundeUi (in the central regions of the Yucatan Penin- sula), and tanneri in Oaxaca (Smith and Larsen, 1975). Farther north along the Pacific Coast, the orcutti stock produced clarki (from central Arizona, through the center of Sonora and down the Pacific Coast of Sinaloa to Nayarit) and melanorhinus (along the Pacific slopes from Nayarit Fig. 15. Eai'ly radiation in Group III. March 1975 LARSEN, TANNER: SCELOPORINE LIZARDS 17 Fig. 16. Second radiation in Group III. through Jalisco, Colima, Michoacan, Guer- rero, and Oaxaca to Chiapas). Hall's com- ments about the separation of nelsoni and pyrocephalus along the Rio Grande de Santiago are also appropriate for clarki and melanorhinus . Apparently this river was a geographic barrier for two groups speciating in opposite directions. Another branch from the orcutti stock produced the nuigister complex. The sub- sequent subspeciation of magister accord- ing to Phelan and Brattstrom (1955) was from central California southward into Baja California and southeastward into Arizona and New Mexico. However, or- cutti has 34 chromosomes, magister zos- teromus (and all other peninsular sub- species of magister) has 30, and m. magis- ter has 26. This supports Hall's ph^logeny with early speciation in Baja Cahfornia and subsequent emergence of two stems {orcutti and magister). A third and final branch from the or- cutti stock moved eastward through the interglacial deserts of Arizona, New Mex- ico, and Texas. This branch (olivaceus) became trapped in the Gulf Coastal Plain refuge during the second glacial period (Fig. 15). Speciation proceeded from oli- vaceus (central Texas, Tamaulipas, Nuevo Leon, and adjacent states) southward across the Central Meseta to spinosus (oc- cupying the entire Neo volcanic Plateau from Puebla and Veracruz on the east to the tip of Durango on the west), horridus (with subspecies along the entire southern flank of the distribution of spinosus), and edwardtaylori (in Oaxaca) (Fig. 16). A secondary speciation from olivaceus (to cautus) has been questioned by Hall (because of intergrades), but he (pers. comm., 1973) did make this observation: Most interestingly there seems to be al- most no question that cautus and olivaceus intergrade south and west of Monterrey (Nuevo Leon) with gene flow occurring presently through the dry valleys and passes. There might be an absolute classic circle of subspecies whose terminal popula- tions are fully sympatric. The last major speciation wdthin Scelo- porus started with cyanogcjiys in the Gulf Coastal Plain refuge (Fig. 15). The first branch produced jarrovi (in the northern plateaus and adjacent escarpments from Arizona on the northwest to Veracruz on the southeast), which in turn produced ornatus (in the ranges of southern Coahu- ila), lineolatcralis (restricted to the moun- tains of eastern Durango), and dugesi (with subspecies in the mountains of Gua- najuato, Michoacan, Colima, Jalisco, and Nayarit.) The second branch from cyanogenys moved westward to produce poinsetti (which occupies most of the northern 18 GREAT BASIN NATURALIST Vol. 35, No. 1 Plateau through southern New Mexico, southwestern Texas, and the Mexican states of Chihuahua, Coahuila, and Du- rango). The third branch extended across Mexico in a southwesterly direction and resulted in hullcri (in the mountains of Jalisco) . The final radiation from the cyano- genys stock extended southward and re- sulted in serrifer (occupying most of the Gulf Coastal Plain in Tamaulipas, San Luis Potosi, Veracruz, Tabasco, Cam- peche, and Yucatan), mucronatus (a mountain form in the Oaxaca Upland and other mountains in the state of Guerrero, Veracruz, Puebla, Mexico, and Hidalgo), and torquatus (which inhabits a large area in central Mexico, including parts of Hidalgo, Veracruz, Mexico, Distrio Fede- ral, Puebla, Morelos, Guanajuato, Micho- acan, Nuevo Leon, Jalisco, San Luis Poto- si, and Zacatecas). Conclusions When presenting his arrangement, Smith (1939) said, "Material from cer- tain areas is still lacking, and more direct evidence of relationships is frequently to be desired. The conclusions now ])resented are accordingly tentative." Smith's state- ment may still apply. Problem areas in- clude Baja California and the grammicus complex. Also several new species and subspecies are being considered by various workers. New kinds of data are now be- ing researched (microdermatoglyphics, for example). However, a point has been reached at which different sets of data reinforce similar conclusions. With over 80 characters, the new groups and sub- groups are distinct at the .999 level of confidence (Larsen and Tanner, 1974). With such a high level of confidence, we conclude that Figure 5 is a natural ar- rangement of species and that future ad- justments may be minor. When phylogeny and zoogeography are considered simultaneousl}-, several trends are evident in the evolution of SccJoporus: (1) the size altered from small to large; (2) the scales, once small, smooth, and granular, changed, becoming large, carin- ate, mucronate, and imbricate; (3) ini- tial movement and speciation was from north to south, and several secondary ra- diations were from southern centers north- ward and from northern centers south- ward; (4) the geography of Baja Cali- fornia created several relicts; (5) habitat preference changed from ground to rocks, cliffs, and trees; and (6) the ancestral stock, which originally was subtropical, adapted to arid conditions, and then sever- al groups returned to tropical or sub- tropical climates. Cope (1900) called SccJoporus the piece de resistance for the theory of derivation of species. This genus seem to show such principles as parallelism, convergence, di- vergence, genetic drift, geographical bar- riers, adaptive radiation, centrifugal spe- ciation, and waif and relict population development. In fact, the cape region of Baja California may provide examples of speciation by continental drift. Sceloporus also exhibits a high degree of chromoso- mal variation, including examples of Rob- ertsonian fission and fusion, and several formulae for sex determination. This ge- nus is extremely well suited for illustra- tion and discussion of evolutionary theory. We conclude that Sceloporus has re- cently speciated in an explosive manner. Because of this ra]:)id adaptive radiation, it is difficult to determine phylogenetic relationships with classical techniques. We are ])ersuaded, however, that the genus Sceloporus does contain three dis- tinct monophyletic groups. Grou]:) I is dis- tinct from the other tw^o groups in having (1) a postfemoral dennal pocket and less than 7 ^'entrals betw'een the femoral pore series or (2) (if the postfemoral dermal pocket is absent) a vestigial gular fold and no postrostrals. The rest of the spe- cies in the genus Sceloporus lack either a jiostfemoral dermal pocket or a vestigial gular fold. If they lack the vestigial gular fold, postrostrals are ])resent and there are more than 8 ^'entrals between the femoral pore series. In considering the systemat- ics of the entire complex, we believe that it is now feasible to recognize for Group I (Table 1) the Cope (1888) monotypic generic designation of Lysoptychus (L. lateralis:=Sceloporus couchi Baird, 1858). We have not by our methods been able to arrive at a satisfactory taxonomic divi- sion of Groups II and III, even though these groups become sej)arable and distinct by use of multivariate analysis. We be- lieve that Groups II and III represent a large assemblage of species that have evolved more recently but that although the characters between the groups are March 1975 LARSEN, TANNER: SCELOPORINE LIZARDS 19 showing indications of evolutionary separ- ation, they have not reached a point of distinction that permits the development of a workable taxonomic key. We there- fore choose at this time to retain them in the genus Sceloporus. Literature Cited AxELROD, D. I. 1948. Climate and evolution in western North America during the middle Pliocene time. Evolution 2:127-144. Baird, S. F. 1858. Description of new genera and species of North American lizards in the Museum of the Smithsonian Institution. Proc. Acad. Nat. Sci. Philadelphia 10:253-256. . 1859. Reptiles of the boundary, with notes by the naturalists of the survey. U.S. — Mex. Boundary Surv. (Emory). 3(2): 1-35. pis. 1-41. ., AND C. GiRARD. 1852. Reptilcs. In: Stansbury, Howard. Exploration and survey of the valley of the Great Salt Lake of Utah, including a reconnaissance of a new route through the Rocky Mountains. Philadelphia, Lippincott and Grambo. (32nd Congr., Spec. Sess.. March. 1851, Sen. Exec. Doc, 2(3)). 487 pp.. 54 pis.. 3 maps. (Reptiles, appendix C, pp. 336-365, 8 pis.) Ballinger, R.E., AND D.W. Tinkle. 1972, Sys- tematics and evolution of the genus Uta. (Sauria: Iguanidae). Misc. Pub. Mus. Zool. Univ. Michigan 145:1-83. BouLENGER, G.A. 1885. Catalogue of the lizards in tlie British Museum (Natural History). 2d ed. London, Tajdor and Francis 2, xiii. 497 pp., 23 pis. Brown, W.L., Jr. 1957. Centrifugal speciation. Q. Rev. Biol. 32:247-277. Bussjaeger, L.J. 1971. Phylogenetic signifi- cance of the comparative ethology of the Spinosus group of Sceloporus (Iguanidae). Ph.D. dissertation. Univ. of Oklahoma. Clarke, R.F. 1965. An ethological study of the iguanid lizard genera Callisawus. Copho- saurus. and Holbrookia. Emporia St. Res. Stud. 13 (4): 1-66. Cole, C.J. i963. Variation, distribution, and taxonomic status of the lizard Sceloporus un- dulatus virgatus Smith. Copeia 1963: 413-425. . 1970. Karyotypes and evolution of tlie spinosus group of lizards in the genus Scelo- porus. Amer. Mus. Novit. 2431:1-47. . 1971a. Karyotypes of the five mono- typic species groups of lizards in the genus Sceloporus. Amer. Mus. Novit. 2450:1-17. . 1971b. Karyotypes and relationships of the pyrocephalus group of lizards in the genus Sceloporus. Herpetologica 27(1):l-8. Cope, E. D. 1888. Catalogue of batrachia and reptilia brought by William Taylor from San Diego. Texas. Proc. U.S. Nat. Mus.. 11:395- 398. . 1900. The crocodilians. lizards and snakes of North America. In: Annual Report, U.S. Nat. Mus. 1898(2): 151-1294. Dickerson, M. C. 1919. Diagnoses of twenty- three new species and a new genus of lizards from Lower California. Bull. Amer. Mus. Nat. Hist. 41(10):461-477. Dixon, W. J. 1967. BMD Biomedical Computer Programs. Univ. of Calif. Pi-ess, Berkeley and Los Angeles, 600 pp. Dumeril. a. H. a. 1856. Description des rep- tilcs nouveaux ou impai-faitement connus de la collection du Museum d'Histoire Naturelle et remarques sur la classification et les car- acteres des reptiles. Arch. Mus. Hist. Nat. Paris 8:439-588. pis. 17-24. Etheridge, R. E. 1964. The skeletal moi-phol- ogy and sj'stematic relationships of sceloporine lizards. Copeia 1964(4) :610-631. Hall. W. P. 1971. Chromosome evolution of the iguanid genus Sceloporus. Herpetol. Rev. 3:106. . 1973. Comparative population cyto- genetics, speciation, and evolution of the crev- ice using species of Sceloporus (Sauria, Iguanidae). Ph.D. dissertation, Harvard. Hallowell, E. 1854. Description of new rep- tiles from California. Proc. Acad. Nat. Sci. Philadelphia 7:91-97. L.\rsen, K. R.. and W. W. Tanner. 1974. Nu- meric analysis of the lizard genus Sceloporus with special reference to cranial osteology. Great Basin Nat. 34(1): 1-41. Milstead, W. W. 1960. Relict species of the Chihuahua Desert. Southwestern Nat. 5(2): 75-88. MiTTLEMAN, M. B. 1942. A summary of the iguanid genus Urosaurus. Bull. Mus. Comp. Zool. Harvard 91 (2) : 106-181. Phelan, R.L.. and B.H. Brattstropii. 1955. Geo- graphic variation in Sceloporus magister. Herpetologica 11(1): 1-14. Presch, W. 1969. Evolutionary osteology and relationships of the horned lizard genus Phrynosoma (family Iguanidae). Copeia 1969 (2):250-275. " ; _ . 1970. Scleral ossicles in the sceloporine lizards, family Iguanidae. Herpetologica 26 (4): 446-450. Purdue, J. R.. and C. C. Carpenter. 1972a. A compai-ative study of the display motion in the iguanid genera Sceloporus, Uta. and Uro- saurus. Herpetologica 28(2) : 137-141. , AND . 1972b. A comparative study of the bodj- movements of displaying males of the lizard genus Sceloporus (Iguan- idae). Behavior 41:68-81. Raisz, E. 1964. Landforms of Mexico. A map prepared for tlie geography branch of the Office of Naval Research. Cambridge. Mass. 1 map. Savage. J. M. 1958. The iguanid lizard genera Urosaurus and Uta. with remarks on related groups. Zoologica 43 (2): 41 -54. . 1960. Evolution of a peninsular lierpe- tofauna. Syst. Zool. 9:184-212. Smith, H. M. 1934. On the taxonomic status of three species of lizards of the genus Scelo- porus from Mexico and Southern United States. Proc. Biol, Soc. Washington 47:121-133. . 1937a. A synopsis of the yarmft/fc group of lizard genus Sceloporus. with description of new subspecies. Occ. Pap. Mus. Zool. Univ. Michigan 358:1-14. . 1937b. A synopsis of the scalaris group of the lizard genus Sceloporus. Occ. Pap. Mus. Zool. Univ. Michigan 361:1-8. 1938. The lizards of the /o/-gt/«/U5 group of the genus Sceloporus Weigmann. Univ. Kans. Sci. Bull. 24:539-693. 20 GREAT BASIN NATURALIST Vol. 35, No. 1 — . 1939. The Mexican and Central Amer- ican lizards of the genus Scrloporus. Field Mus. Nat. Hist. Zool. Ser. 26:1-397 . . 1946. Handbook of lizaids. Comstock Publ. Co.. Ithaca. 557 pp. AND K. R. L.vRSEN. 1975. A new species of the jormosus group of the lizard genus Sceloporus. Copeia 1975 (l):47-50. , ,\ND E. H. Tayi-or. 1950. An annotated check list and key to the i-eptiles of Mexico exclusive of the snakes. Bull. U.S. Nat. Mus. 199:1-199. STEnuiNs, R. C. 1944. Some aspects of the ecol- ogy of the iguanid genus Vma. Ecol. Monogr. 14(3): 31 1-322. Stejneger, L. H. 1904. A new lizard from the Rio Grande Valley, Texas. Proc. Biol. Soc. Washington 17:17-20. Tanner, W. W. 1966. The night snakes of Baja California. Trans. San Diego Soc. Nat. Hist. 14(15):189-196. Trosciiee, F. H. 1850. (Pub. 1852). Cophosau- rus Tcxanus, neue Eidechsen-Gattung aus Texas. Arch. Naturgesch. 16(1 ) :388-394. Werster. T. p., W. p. H.\ll HI, and E. E. Will- LiAMS. 1972. Fission in the evolution of a lizard karyotype. Science 1 77(4049) : 61 1-613. Weigmann, a. F. 1828. Beitrage zur amphi- bienkunde. Isis von Oken, V. 21, 1828:369-370- WisiiART. D. 1968. A fortran H programme (CLUSTAN) for numerical classification. Computing Laboratory. St. Andrews, Fife, Scotland, 50 pp. NEW SYNONYMY AND NEW SPECIES OF AMERICAN BARK BEETLES (COLEOPTERA: SCOLYTIDAE)^ Stephen L. Wood" Abstract. — New synonymy is proposed as follows: Pityokteines Fuchs { = Orthotomides Wood), Araptus foveifrons (Schedl) { = inter jectus Wood), n. comb., Cryptocarenus seriatus Eggers { = boliv- ianus Eggers), Hylastes gracilis LeConte {=asper Swaine), Procryphalus utahensis Hopkins {=aceris Hopkins), Scolytodes maurus (Blandford) {=Hexacolus ellipticus Eggers), Scolytus hermosus Wood {=sylvaticus Bright), Xyleborus adelographus Eichhoff { = vitiosus Schedl), Xyleborus catulus Blandford {=intricatus Schedl), Xyleborus nepos Eggers {=signatus Schedl), Xyleborus titubanter Schedl {=dissidens Wood). The following species and subspecies are named as new to science: Cnesinus electus (Costa Rica), C. pilatus (Mexico), Hylastes asperatus (New Mexico), Scolytus toru- lus (Costa Rica), S. laetus (Mexico), Scolytodes amabilis and S. lepidus (Mexico), S. obesus (Pana- ma), Pseudothysanoes concentralis and P. tumidulus (Mexico), Thysanoes tuberculatus (Mexico), Pityokteines mystacinus (Washington), Ips pilifrons subsp. thatcheri (Nevada), Araptus attemiatus (Mexico), A. fossifrons (Mexico, Guatemala), Amphicranus parilis (Mexico), Corthylus cecropii (Costa Rica). On the following pages several newly discovered cases of synonymy and 1 7 spec- ies and 1 subspecies new to science are presented for American Scolytidae. The specific synonymy is presented in alpha- betical order for convenient reference. The species new to science represent the gen- era Cnesinus (2), Hylastes (1), Scolytus (2), Scolytodes (3), Pseudothysanoes (3), Thysanoes (1), Pityokteines (1), Araptus (2), Amphicranus (1), and Corthylus ( 1 ) . The new subspecies is of Ips pili- frons. The new species are from the fol- lowing countries: United States (2), Mex- ico (10), Costa Rica (3), Panama (1), Mexico and Guatemala ( 1 ) . The new subspecies is from the United States (Ne- vada). New Synonymy Pityokteines Fuchs Pityokteines Fuchs, 1911, Morphologische Studien iiber Borkenkafer: I. Die Gattungen Ips De- Geer und Pityogenes Bedel, p. 33 (Type- species: Ips curvidens GeiTnar, subsequent designation by Hopkins, 1914, Proc. U.S. Nat. Mus. 48:127) Orthotomides Wood, 1951, J. Ent. Soc. Kansas 24:32 (Type-species: Orthotomicus lasiocarpi Swaine, original designation) . New synonymy The discovery of mystacinus Wood, described below, closes the gap in charac- ters between Pityokteines and Orthoto- mides to such an extent that the latter name must be placed in synonymy even though lasiocarpi (Swaine) entirely lacks the long female frontal hair characteristic of Pityokteines. Female mystacinus have part of the frontal vestiture elongate and also have two sutures visible on the pos- terior face of the antennal club. This generic synonymy necessitates the transfer of lasiocarpi from Orthotomides to Pityokteines. Araptus foveifrons (Schedl), n. comb. Tharnnophthorus foveifrons Schedl, 1963, Ent. Arb. Mus. Frey 14:161 (Holotype, male; Guadalajara. Jalisco, Mexico; Schedl Coll.) Araptus interjectus Wood, 1974, Brigham Young Univ. Sci. Bull., Biol. Ser. 19(1) :44 (Holo- type, male; Volcan de Agua, Guatemala; Wood Coll.). New synonymy When the North and Central American Araptus were reviewed for the mono- graph, two closely related species were found, both of which fit the description of Tharnnophthorus foveifrons Schedl. It was assumed that the common, widely distributed species of these two was fovei- frons; the rare one was named interjectus Wood. Recently, I had the opportunity to examine the male holotype of foveifrons and to compare it to the male holotype of interjectus. They represent the same species. The common species with which it had been confused is named below as Araptus fossifrons. Cryptocarenus seriatus Eggers Cryptocarenus seriatus Eggers, 1933, Orig. Mem. Trav. Lab. Ent. Paris 1(1): 10 (Holotype, female; Nouveau Chantier, French Guayana; Paris Mus.) Cryptocarenus bolivianus Eggers, 1943, Mitt. Miinchner Ent. Ges. 33:356 (Holotype, fe- ■■Part of this research was sponsored hy the National Science Foundation. -Department of Zoology, Brigham Young University, Prove, Utah 84602. Scolytidae rontrihutioii No, 60 21 22 GREAT BASIN NATURALIST Vol. 35, No. 1 male; Cochabamba, Bolivia; U.S. Nat. Mus.). New synonymy The holotypes of Cryptocarenus seriatus Eggers and boUvianus Eggers were com- pared directly to my material fromi Bra- zil, Venezuela, and other areas. All rep- resent the same biological species. Hylastes gracilis LeConte Hylastes gracilis LeConte, 1868, Trans. Amer. Ent. Soc. 2:174 (Lectotype, female; Tahoe Valley, California; Mus. Comp. Zool., de- signated by Wood, 1971, Great Basin Nat. 31:145) Hylastes asper Swaine, 1917, Dom. Canada Dept. Agric. Ent. Br. Bull. 14(1): 19 (Holotype, female; Larimer Co., Colorado; Canadian Nat. Coll.). New synonymy The holotype of Hylastes asper Swaine is rather different from what the descrip- tion might lead one to believe. It is vir- tually identical to the type of longus Le- Conte and well within the range of vari- ability of gracilis LeConte. All three types were compared to my material and, in my opinion, represent the same species. The species I have previously treated as asper is described below. Procryphalus utahensis Hopkins Procryphalus utahensis Hopkins, 1915, U.S. Dept. Agric. Kept. 99:33 (Holotype, female; Alta, Utah; U.S. Nat. Mus.) Procryphalus aceris Hopkins, 1915, U.S. Dept. Agric. Kept. 99:33 (Holotype, female; Al- bany, Oregon; U.S. Nat. Mus.). New synon- ymy Procrophalus aceris Hopkins was named from a series of about six specimens taken from a limb that had been cut in the vicinity of Albany, Oregon. The limb was given to Hopkins who identified it as Acer macrophyllum (personal com- munication from W. J. Chamberlin). Be- cause the strial punctures were slightly larger than usual, I did not associate these specimens with utahensis Hopkins until an identical series was taken from Salix scouleriana at Dixie Pass, Oregon. This series was compared directly to the type series of utahensis and aceris and to other series from British Columbia, Quebec, Cal- ifornia, Colorado, South Dakota, and Utah. Only one species is represented by this material; all specimens are from Salix. Chamberlin, myself, and many others have searched both standing and cut Acer macrophyllum on numerous occasions without finding indications of bark beetle activity. The host identification by Hop- kins must have been erroneous. Scolytodes maurus (Blandford) Prionosceles maurus Blandford, 1897. Biol. Centr. Amer., Coleopt. 4(6): 178 (Syntypes; Cerro Zunil and Pantaleon, Guatemala; British Mus. Nat. Hist.) Hexacolus ellipticus Eggers, 1934, Ent. Blatt. 30:80 (Holotype, male; Turrialba, Costa Ri- ca; Institut fiir Pflanzenschutzforschung Kleinmachnow). New synonymy The syntypic series of maurus (Bland- ford) and the male holotype of ellipticus Eggers were compared to my material taken from southern Mexico to Panama. Only one common species is represented by this material. In all, 158 specimens were examined. Scolytus hermosus Wood Scolytus hermosus Wood, 1968, Great Basin Nat. 28:12 (Holotype, male; 2 km N Tlaxcala state line 18 km N Tlaxco, Puebla, Mexico; Wood Coll.) Scolytus sylvaticus Bright, 1972, Canadian Ent. 104:1489 (Holotype, male; Cerro Potosi, Nu- evo Leon, Mexico; Canadian Nat. Coll.). New synonymy The holotype and allotype of sylvaticus Bright and three other specimens from the type locality of that species were com- pared directly to the type series of hermos- us Wood. While the holotype has abdom- inal sternum 2 less strongly concave than normal, the other Cerro Potosi specimens and Bright's illustration of a male from the type series (Fig. 1, p. 1490) are nor- mal specimens of hermosus. For this rea- son, sylvaticus must be placed in synon- ymy. Xylehorus adelographus Eichhoff Xyleborus adelographus Eichhoff, 1868, Berliner Ent. Zeitschr. 11:400 (Syntypes, female; Bra- zil; Brussels Mus.) Xyleborus vitiosus Schedl, 1940, An. Esc. Nac. Cienc. Biol., Mexico 1:367 (Holotype, female; Mexico?; Schedl Coll.). New synonymy The syntype of adelographus Eichhoff in the Brussels Museum is 2.8 mm in length; the holotype of vitiosus Schedl is 3.5 mm (not 3.7 mm as given by Schedl). Seven specimens in my collection from Colombia are 3.3-3.4 mm, and one from southern Brazil is 3.5 mm. The specimens from Colombia are very slightly stouter, March 1975 WOOD: AMERICAN SCOLYTIDAE 23 the dechvity along the suture is flat, and the punctures on the declivital striae are slightly more deeply impressed. The holo- type of vitiosus and my Brazilian speci- men of adelographus are identical in every respect except for size. These specimens are almost identical to the syntype of adelographus and are regarded as con- specific. Because extensive collecting has not produced any specimens of this spec- ies from Mexico or Central America, be- cause the Mexican origin of the holotype of vitiosus is questioned on its locality la- bel, and because the holotype of vitiosus resembles specimens from southern Brazil much more closely than it does represent- atives of this species from Colombia, this species should be dropped from the faunal Hst of Mexico until more definite evidence of its occurrence in Mexico is found. Xyleborus catulus Blandford Xyleborus catulus Blandford, 1898, Biol. Centr. Amer., Coleopt 4(6):216 (Holotype, female; Volcan de Chiriqui, Panama; British Mus. Nat. Hist.) Xyleborus intricatus Schedl, 1949, Rev. Brasil. Biol. 9:274 (Holotype, female; St. Catarina, Brazil; Schedl Coll.). New synonymy Several specimens in the California Academ}^ of Sciences were identified by Schedl as Xyleborus intricatus Schedl. Since these specimens are of catulus Blandford, a loan of the holotype of in- tricatus was requested. The specimen re- ceived was labeled as a "paratj^pe" of intricatus from Nova Teutonia, Santa Ca- tarina, Brazil; however, this species was based on a unique female, and paratypes were not designated in the original des- cription. Because this specimen is also of catulus, intricatus is tentatively placed in synonymy until the holotype is located. Xyleborus nepos Eggers Xyleborus nepos Eggers, 1923, Zool. Meded. 7:198 Lectotype, female; Java; U.S. Nat. Mus., designated by Anderson and Anderson. 1971, Smithsonian Contrib. Zool. 94:21) Xyleborus signatus Schedl, 1949, Rev. Brasil Biol. 9:278 (Holotype, female; Mexico; Schedl Coll.). New synonymy The holotype of signatus Schedl was named from a unique female labeled "Mexico." This specimen is identical to a long series of nepos Eggers from Indonesia and the Philippine Islands, some of which were compared directly to the lectotype of nepos. Because extensive collecting in Mexico has not produced another speci- men of this species, the holotype of signa- tus is considered a mislabeled specimen and should be dropped from the ]\Iexican faunal list until more definite evidence of its occurrence there is found. Xyleborus titubanter Schedl Xyleborus titubanter Schedl, 1948, Rev. de Ent. 19: '578 (Holotype, female; Mexico; Schedl Coll.) Xyleborus dissidens Wood, 1972. Brigham Young Univ. Sci. Bull., Biol. Ser. 19(1) :41 (Holo- type, female; 9 km NE Teziutlan, Puebla, Mexico; Wood Coll.). New synonymy The holotypes of titubanter Schedl and dissidens Wood were compared directly to one anotlier and were found to repre- sent the same species. Taxa New To Science Cnesinus electus, n. sp. This species is distinguished from the female holotype of bisulcatus Schedl by the more narrowly spaced eyes (above), by the smaller, more abruptly elevated epistomal process (difference very slight), by the very slightly larger, deeper prono- tal punctures, by the larger, more widely spaced strial punctures, and by the coarser declivital setae with a complete row of setae on interstriae 1 . Female. — Length 2.5 mm, (paratypes 2.3-2.6 mm), 2.8 times as long as wide; color dark brown, elytra dark reddish brown. Frons strongly, transversely impressed at level of antennal bases, convex above, somewhat flattened below this point, with epistomal process poorly developed and devoid of granules or tubercles; eyes ap- proximate above, separated by a distance equal to 0.5 times greatest width of an eye (about twice this wide in bisulcatus) ; surface rugulose and rather coarsely punctured above eyes, rugose-reticulate below upper level of eyes, with a few fine punctures along lateral margins and on epistoma; vestiture limited to lateral and epistomal areas. Eyes very large; very coarsely faceted. Pronotum 1.1 times as long as wide; widest just in front of middle, almost straight from widest point to sharply de- fined basal angles, somewhat constricted 24 GREAT BASIN NATURALIST Vol. 35, No. 1 just behind narrowly rounded anterior margin; surface dull, subshining, more distinctly shining toward base, punctures almost round at base, becoming increas- ingly elongate anteriorly, occasionally confluent in anterior area; glabrous. Elytra 1.7 times as long as wide; sides almost straight and parallel on basal two- thirds, rather narrowly rounded behind; striae 1 rather strongly, others moder- ately, not abruptly impressed, punctures small, except on 1 not confluent, distinct- ly, rather strongly impressed, separated by distances slightly less than their own diameters; interstriae feebly convex, shin- ing, punctures varying from minute to two-thirds size of those of striae, in in- definite uniseriate rows. Declivity con- vex, except moderately impressed between interstriae 3; striae feebly impressed, punctures rather small but impressed; in- terstriae 1 abruptly elevated to apex, about half as high as wide, others almost flat, 2 about one and one-half times as wide as 1, twice as wide as 3; all interstri- ae armed by uniseriate rows of moderately large rounded setiferous granules, each granule about as high as wide, distinctly larger than in elegans. Vestiture confined to declivity, except on interstriae 1, con- sisting of rows of rather coarse, moderately long, interstrial bristles, and short, fine, strial hair. Female. — Similar to male except epis- tomal callus more prominent. Type Locality.— Five miles or 8 km SE Cartago, Cartago, Costa Rica. Type Material. — The female holo- tvpe, male allotype, and 7 paratypes were taken on 2-VIII-63, at 1500 m, No. 98, from twigs of Miconia (?), by S. L. Wood. Seven paratypes bear the same data except that they were from twigs of an unknown tree; five paratypes came from same locality on 29-VIT-63, from a woody vine; and seven paratypes are from Peralta, Cartago, Costa Rica, lO-III- 64, 500 m, tree seedling, S. L. Wood. The holotype, allotype, and parat5q3es are in my collection. Cnesinus pilatus, n. sp. This species is distinguished from strigi- coUis LeConte by the larger size, by the stouter body form, by the different female frons, by the absence of tubercles on de- clivital interstriae 2, and by other charac- ters noted below. Female. — Length 3.1 mm, 2.1 times as long as wide; color dark brown, elytra dark reddish brown. Frons as in strigicollis except area below carina slightly less strongly impressed, distance between eyes 0.8 times as wide as width at level of antennal insertion (as in strigicollis) , carina 0.5 times as wide as distance between eyes, median impunctate area above carina larger, extending above upper level of eyes, vestiture in lateral areas not extending above upper level of eyes and not on median third at vertex (extending well above eyes and almost to median line in strigicollis). Pronotum 0.9 times as long as wide; about as in strigicollis except more close- ly, slightly more coarsely strigose. Elytra 1.3 times as long as wide, 1.7 times as long as pronotum; similar to strigicollis except interstriae three times as wide as striae, declivity more broadly impressed, declivital interstriae 2 devoid of granules, vestiture about half as long, stouter. Interstriae 2 on declivity with a row of punctures, each puncture about two-thirds as large as those of adjacent striae. Type Locality. — Thirteen km or eight miles W El Palmito, Sinaloa, Mexico. Type Material.^ — The female holo- type was taken at the type locality on 7- Vin-64, by H. F. Howden. The holotype is in the Canadian Na- tional Collection. Hylastes asperatus, n. sp. This is a difficult species to recognize. The very large pronotal punctures are ir- regular in size, as in porculus Erichson, but much closer; the discal interstriae are slightly wider than in allied species, more nearly convex, and very finely, closely crenulate (usually not clearly apparent unless the light source is posterior to the specimen). This species has been treated as asper Swaine, but the type of asper is quite different. Female. — Length 4.2 mm (paratypes 4.0-4.6 mm), 2.7 times as long as wide; color dark brown. Frons as in gracilis LeConte with inter- ocular impression moderately strong, fine, low carina evidently always extend- March 1975 WOOD: AMERICAN SCOLYTIDAE 25 ing from this impression to base of episto- mal lobe where it forks as in related spec- ies. Pronotum 1.2 times as long as wide; sides on slightl)^ more than basal half straight and parallel, rather broadly round- ed m front; surface subshining, indica- tions of reticulation obscure but usually visible at high magnification toward an- terior or basal areas, punctures coarse, very close, deep, irregular in size in some specimens, interspaces usually equal to less than one-fourth diameter of a punc- ture; median line partly impunctate, not raised; glabrous. Elytra 1.8 times as long as wide, 1.8 times as long as pronotimi; outline as in gracilis; striae moderately impressed, punctures rather coarse, deep; interstriae as wide as striae, punctures moderately coarse, close, confused, their anterior mar- gins elevated into fine, crenulate, trans- verse ridges of variable height (this char- acter approached in allied species, but not to this degree). Declivity convex, steep; striae narrowly, deeply impressed, punc- tures somewhat obscure; interstriae about twice as wide as striae, armed by fine, confused tubercles. Vestiture confined to declivity, scalelike. Male. — Similar to female except slightly stouter, and last visible abdominal sternum medially impressed and pubes- cent. Type Locality. — New Mexico. Type Material. — The female holo- type, male allotype, and 18 para types were mounted on cards all on one pin bearing the label "New Mexico, F. H. Snow." One paratype in the Canadian National Collection labeled "Las Vegas Hot Springs, New Mexico, 7000 ft., Aug. '82, F. H. Snow" may be from the same series. One paratype from each of the following Arizona localities is labeled: Hanagan Camp Ground, Greenlee Co., 12- Vn-68, D. E. Bright; Santa Catalina Mts., 5-Vin-68, D. E. Bright. The holotype, allotype, and 18 para- types are in my collection; the three re- maining paratypes are in the Canadian National Collection. Scolytus torulus, n. sp. This species is distinguished from dimi- diatus Chapuis by the smaller average size, by the subvertical, somewhat convex ab- dominal sternum 2, by the absence of a tuft of hair on sternum 2 immediately posterior to the spine, by the very differ- ent male frons, and by other characters described below. Male. — Length 2.4 mm (paratypes 2.0-2.4 mm), 2.1 times as long as wide; color very dark brown to black, elytra slightly lighter. Frons rather weakly convex above, a distinct, moderately deep, transverse im- pression just above epistoma, deepest in median area, a distinct but weak, rounded, median elevation immediately above im- pressed area; surface strongly reticulate above, more shining and obscurely acicu- late in impressed area, punctvires fine, rather deep, sparse in median area, more numerous laterally; vestiture of fine hair on margins and impressed area, a few of them rather long. Pronotimi as in dimidiatus except punc- tures in lateral areas considerably larger. Elytra 1.1 times as long as wide, 1.2 times as long as pronotum; surface simi- lar to dimidiatus but much more finely punctured, few to many longitudinal lines or striations sometimes present. Sparse setae shorter and much stouter than in dimidiatus. Sternum 2 subvertical, moderately con- vex, spine similar to dimidiatus but smal- ler; surface dull, very coarsely, deeply, closely punctured; sterna 3-5 similar but more finely punctured; vestiture of very fine, short hair, without a specialized tuft posterior to spine on 2. Female. — Similar to male except frons without impression or elevation, not strig- ose, vestiture similar to but finer and about one-third as abundant as in dimidi- atus female; spine on sternum 2 about half as large as in male. Type Locality. — Rincon de Osa, Punt- arenas, Costa Rica. Type Material. — The male holotype, female allotype, and 10 paratypes were taken at the type locality on ll-Vin-66, 30 m. No. 72, from a leguminous tree, by me. The holotype, allotype, and paratypes are in my collection. Scolytus laetus, n. sp. This species is distinguished from toru- lus Wood by the flattened sternum 2 26 GREAT BASIN NATURALIST Vol. 35, No. 1 which is much more finely, sparsely punc- tured, with longer, more abundant hair, and by characters of the frons described below. Male. — Length 2.4 mm (paratypes 2.4-2.5 mm), 2.4 times as long as wide; color very dark brown. Frons as in torulus except elevation wdder, area above elevation more dis- tinctly, more broadly impressed, vestiture on lateral margins more abundant, much longer. Pronotum and elytra as in torulus. Sternum 2 vertical, almost flat, surface dull, punctures small, not clearly evident, spine as in torulus, vestiture much longer, some setae as long as spine. Female. — Similar to male except frons as in female torulus but with vestiture slightly more abundant and longer, par- ticularly in lower areas; sternum un- armed, spine absent, vestiture finer, more abundant. Type Locality. — Forty-eight km or 30 miles N Rosamorada, Nayarit, Mexico. Type Material. — The male holotype, female allotype, and three male para- types were taken at the type locality on 15-Vn-65, 1000 m. No. 257, from Inga paterno, by me. The holotype, allotype, and paratypes are in my collection. Scolytodes amabilis, n. sp. In general body features, this species is very similar to clusiavorus Wood, but the female frons is much more similar to volcanus Wood. It is distinguished from those species as indicated below. Female. — Length 1.6 mm (paratypes L4-1.6 mm), 2.5 times as long as wide; color very dark brown, almost black. Frons essentially convex, with a very shallow, subconcave area on median third just below upper level of eyes, this im- pression continued on median fifth as an abrupt, shallow sulcus to epistoma, median half of lateral areas bordering sulcus with numerous beadlike granules, remaining areas somewhat dull, with fine punctures; surface of sulcus shining, almost smooth; rather sparse vestiture limited to margins, of fine, long hair, those on dorsal margin attaining level of antennal insertion, shorter laterally and below. Pronotum and elytral outlines as in clusiacolens Wood; pronotum surface re- ticulate, punctures as in clusiavorus; strial and interstrial punctures similar to but slightly larger than in clusiavorus. Strial setae almost obsolete; interstrial setae al- most obsolete on even-numbered inter- striae, fine, rather short, and widely spaced on odd-numbered interstriae. Male. — Similar to female except frons convex, reticulate, vestiture very sparse, short, inconspicuous. Type Locality. — Mt. Tzontehuitz, Chiapas, Mexico. Type Material.— The female holo- type, male allotype, and 15 paratypes were taken at the type locality on 29-V- 69, at 3000 m, by D. E. Bright. Twenty- one paratypes bear the same data except 23-VI-69; two of them were taken from Quercus sp. The holotype, allotype, and most of the paratypes are in the Canadian National Collection; the remaining paratypes are in my collection. Scolytodes obesus, n. sp. This species is distinguished from the remotely related immanus Wood by the smaller size, by the stouter body form, by the pair of carinae on the female frons, and by other characters. Female. — Length 1.8 mm, 2.1 times as long as wide; color yellowish brown, anterior third of pronotum darker. Frons shallowly, broadl}^ concave from level of antennal insertion to upper level of eyes (upper area concealed by prono- tum), surface smooth and shining except subreticulate near margins; epistomal area from margin to level of antennal inser- tion longitudinally divided into equal thirds by a pair of rather strongly ele- vated carinae; premandibular epistomal lobe large, conspicuous, pubescent; vesti- ture mostly confined to margins of upper half of frontal area, consisting of a row of long, subplumose setae, longest setae equal to about one-half to two-thirds distance between eyes. Pronotum 0.97 times as long as wide; widest near base, sides weakly, arcuately converging on basal two-thirds, rather broadly rounded in front; anterior third moderately declivous, finely asperate; posterior areas reticulate, very finely, rather closely punctured. Glabrous ex- March 1975 WOOD: AMERICAN SCOLYTIDAE 27 cept for an occasional coarse seta on as- perate area. Elytra 1.15 times as long as wide, 1.25 times as long as pronotum; sides almost straight on basal half, slightly wider at base of declivity, rather broadl}' rounded behind; disc confined to basal half; striae not impressed, punctures small, rather shallowly impressed, spaced by distances equal to diameter of a puncture; inter- striae four times as wide as striae, smooth, shining, punctures small to minute, weak- ly impressed; interstriae 10 weakly carin- ate to level of sternum 5. Declivity con- vex, moderately steep; sculpture as on disc. Vestiture of sparse, moderately long bristles on odd-nvmibered interstriae. Protibia slender, lacking minute tooth on posterior face near tarsal insertion. Type Locality. — Barro Colorado Is- land, Panama Canal Zone. Type Material. — The female holo- type was taken at the type locality on 7- VIII-67, L. and C. W. O'Brien. The holotype is in my collection. Scolytodes lepidus, n. sp. This species is distinguished from amoe- nus Wood by the slightly larger average size, by the slightly larger elytral punc- tures, by the presence of a few elytral setae, particularly along sides, and by the very different female frons. Female. — Length 2.0 mm (paratypes 1.8-2.3 mm), 2.2 times as long as wide; color dark brown, base of pronotum usu- ally pale. Frons broadly flattened from epistoma to vertex, almost smooth, upper half and sides below coarsely, closely punctured and pubescent, median third on lower half slightly elevated, smooth, shining, impmictate, glabrous; vestiture long, moderately abundant, more widely dis- tributed than m amoenus. Pronotum 1.0 times as long as wide; as in amoenus except discal area reticulate (smooth to subreticulate in amoenus) and moderately pubescent at lateral margins (almost glabrous in amoenus). Elytra 1.2 times as long as wide, 1.3 times as long as pronotum; as in amoenus except strial and interstrial punctures larger, more completely confused and lateral areas with sparse setae (entirely glabrous in amoenus). Male. — Similar to female, with frons similar to male amoenus except more coarsely punctured, more protuberant in median area, with no gramiles. Type Locality.^ — Thirty-three km or 21 miles N Juchitlan, Jalisco, Mexico. Type Material. — The female holo- type, male allotype, and 25 paratypes were taken at the type locality on 3-Vn- 65, at 1300 m. No. 177, from Ficus, by me, from the same branches that con- tamed the type series of amoenus. The holotype, allotype, and paratypes are in my collection. Scolytodes genialis, n. sp. This species is distinguished from lepi- dus Wood by the absence of pmictures in the asperate area of the jDronotum and by the much finer punctures on the pronotal disc and on the elytra. Female. — Length 1.8 mm (paratypes 1.8-2.1 mm), 2.3 times as long as wide; color light brown, anterior half of prono- tLun darker. Frons as in lepidus except vestiture fin- er, very slightly shorter. Pronotmn as in lepidus except punctures very fine, shal- low. Elytra as in lepidus except strial punctures fine, shallow, in definite rows, interstrial punctures very small, confused, striae 1 not impressed on declivity, vesti- ture on sides of elytra minute. Type Locality. — Laguna Santa Maria, Nayarit, Mexico. Type Material.- — The female holo- type, male allotype, and six paratypes were taken at the type locality on 7-VII- 65, at 900 m. No. 197, irom Ficus with yellow bark, by me. Four paratypes are from 24 km or 15 miles S Mazamitla, Jalisco, Mexico, 22-VL65, 2500 m, No. 97, Ficus with yellow bark, by me. The holotype, allotype, and paratypes are in my collection. Pseudot/iysanoes concentralis, n.sp. This species is distinguished from quer- cinus Wood by the concentric, carinate pronotal asperities, by the more strongly impressed Irons, by the greatly reduced elytral punctures, and by the wider elyt- ral scales. Female. — Length 1.0 mm (paratypes 28 GREAT BASIN NATURALIST Vol. 35, No. 1 1.0-1.2 mm), 2.6 times as long as wide; color dark brown. Frons rather strongly concave on medi- an two-thirds of area below upper level of eyes, surface subrugose, with sparse granules except almost smooth on lower half of concavity; vestiture short, sparse. Antennal scape as wide as long, as long as pedicel, bearing a small tuft of long hair; club about as in quercinus. Pronotum 0.73 times as long as wide; outline as in quercinus; simimit at mid- dle; anterior slope on median third armed by six concentric, uniform, transverse carinae, other asperities absent, carina 1 submarginal, 6 at summit; posterior areas shining, almost smooth, punctures al- most obsolete. Vestiture consisting of a row of scales posterior to each of first five carinae, and rather sparse, coarse hair in remaining areas. Elytra 1.8 times as long as wide, 2.4 times as long as pronotum; outline as in quercinus; striae not impressed, punc- tures small, shallow; interstriae almost smooth, about twice as wide as striae, punctures almost obsolete; surface usually covered by a thin incrustation. Declivity convex, steep, sculpture as on disc but punctures even more obscure. Vestiture mostly abraded, consisting of widely spaced, short, interstrial scales, each scale one and one-half to two times as long as wide. Male. — Length 0.8 mm; similar to female except smaller, slightly stouter; frontal concavit}' not as deep; pronotal asperities not fused or clearly concentric. Type Locality.- — Ten km or 6 miles SE Totolapan, Oaxaca, Mexico. Type Material. — The female holo- type, male allotype, and 24 paratypes were taken at the type locality on 21 -VI- 67, 1000 m. No. 73, from a leguminous roadside shrub with a yellow flower that was presumed to be Cassia sp., by me. The holot^^pe, allotype, and paratypes are in my collection. Pseudothysanoes tumidulus, n. sp. This species is distinguished from the distantly allied graniticus Wood by the different female frons, by the smaller ely- tral scales, and by the very different male declivity as described below. Male. — Length 1.7 mm (paratypes 1.5-2.0 mm), 2.2 times as long as wide; color very dark brown. Frons flattened on lower two-thirds, with small concavity on median fifth, ascending slightly to epistomal margin, convex above; subshining and almost smooth on flattened area, more coarsely punctate-granulate above; vestiture con- fined to epistomal area and to convex area. Pronotimi similar to graniticus except anterior margin narrowly rounded and armed by six teeth and posterior areas rather strongly reticulate. Elytra 1.3 times as long as wide; sides straight and parallel on basal three- fourths, rather abruptly rounded, then broadly rounded behind; striae not im- pressed, punctures small, moderately deep, spaced by distances equal to diameter of a puncture; interstriae three times as wide as striae, almost smooth, punctures min- ute, granulate, uniseriate except confused near declivity. Declivity with basal mar- gin abrupt, basically convex except upper half flattened to striae 4, a moderate bul- la just below middle from interstriae 2-4; strial punctLu-es smaller and not as deep as on disc, closer, in indistinct rows, of same size and shape as confused interstrial granules, interstrial punctures obsolete; bulla covered by same surface sculpture as elsewhere. Vestiture of interstrial rows of scales, each slightly longer than wide on disc except scales confused, more abundant, much longer, and more slender at base of declivity; declivity glabrous; rows of fine, recumbent strial hair on disc. Female. — Similar to male except frons broadly, shallowly concave from epistoma to well above eyes, vestiture on upper area slightly more abundant (less abundant and shorter than in graniticus) ; anterior margin of pronotum more broadly round- ed, unarmed; declivity convex, sculpture as on disc except strial punctures obso- lete; rows of interstrial scales continued to apex, each scale equal in length to three-fourths distance between rows, more closely spaced within a row, each about three to four times as long as wide. Type Locality. — Highway 120, 129 km NE San Juan del Rio, Queretaro, Mexico. Type Material. — The male holotype, female allotype, and 27 paratypes were taken at the type locality on 9-VL71, at March 1975 WOOD: AMERICAN SCOLYTIDAE 29 2500 m, from mistletoe on oak, b}^ D. E. Bright. The holotype, allotype, and most of the paratypes are in the Canadian National Collection; the remaining paratypes are in my collection. Thysanoes tuberculatus, n. sp This species is distingLiished from gran- ulifer Wood by the smaller size and by the ver}" different el}^tral declivit}^ as described below. Male. — Length 1.8 mm, 2.7 times as long as mde; color rather dark yellowish brown. Frons and pronotum as in granulifer and berchemiae Blackman. Elytra 1.6 times as long as wide; disc as in granulifer except granules slightly larger. Declivity steep, convex; striae 1 and 2 with punctures minute, visible al- most to apex; suture slightly elevated, with a row of small granules on basal half; in- terstriae 2 with two or three tubercles on less than basal fourth, broadly impressed below and entirely devoid of punctures and granules; interstriae 3 moderately el- evated on middle third and anned wdth a row of six to eight rather coarse tuber- cles, lower third without punctures or granules; lateral interstriae each with a row of tubercles on basal area but none of them attaining normal apex for these interstriae. Vestiture of rows of interstrial scales, scales on disc largely abraded, lit- tle if any longer than wide, longer at base of declivity; those on 3 up to three times as long as on disc and four times as long as wide, slightly shorter on other inter- striae; declivital interstriae 2, 4, lower two-thirds of 1, and lower half of 3 glab- rous. Type Locality. — Eighty-five km or 53 miles S Valle Nacional, Oaxaca, Mexi- co. Type Material. — The male holotype was taken at the type locality on 24-V-71, at 3300 m, D. E. Bright. The holotype is in the Canadian Na- tional Collection. Pityokteines mystacinus, n. sp. This species is distinguished from minu- tus (Swaine) b}^ the smaller size, by the very different ornamentation of hair on the female frons, by the finer pronotal and elytral pmictures, and by other char- acters. It probably is much more closely related to lasiocarpi (Swaine). Female. — Length 2.1 mm (paratypes 1.7-2.1 mm), 3.0 times as long as wide; color brown. Frons similar to lasiocarpi except more broadly convex, surface not as smooth, somewhat dull, punctures average smaller and, on lower third, becoming almost ob- solete; vestiture on lower third abundant, rather long, epistomal brush very broad, rather dense; setae on upper half of fron- tal area sparse, short. Antennal club al- most as in lasiocarpi. Pronotum as in lasiocarpi except anteri- or margin more narrowly rounded; vesti- ture uniformly short as in lasiocarpi. Ely- tra as in lasiocarpi except punctures on disc slightly larger, very slightly more confused on basal half, and declivital striae 1 more strongly impressed, with punctures on striae 1 slightly larger, sub- apical transverse elevation at apex of sul- cus more distinct (but still rather ob- scure) ; position, number, and size of tub- ercles as in lasiocarpi. Vestiture similar in abundance, but ver}^ slightly longer than a lasiocarni. Male. — Similar to female except upper half of frons more strongly convex, vesti- ture on lower third greatly reduced in abundance and length; elytral declivity with sulcus slightly deeper (about as in male lasiocarpi) but narrower. Type Locality. — Mount Rainier Na- tional Park, Washington. Type Material. — The female holo- type, male allotype, and three female paratypes were taken at the tvpe localitv on 21-Vin-62, silver fir, D. E. Bright. ^ The holotype, allotype, and one para- type are in the Canadian National Col- lection and two paratypes are in my col- lection. Ips pilifrons thc/tcheii, n. subsp. This geographical race of pilifrons is distinguished from p. pilifrons Swaine, of northern Colorado, b}- the characters of the frons described below. Both subspecies are replaced throughout all of Utah by p. utahensis Wood in which frontal charac- ters are entirely different. The following comparisons are based on females having the maximum frontal elevation and piles- 30 GREAT BASIN NATURALIST Vol. 35, No. 1 ity for their race; occasional specimens of p. pilifrons almost overlap the maximum development of p. tJiatcheri. Female. — Length 4.6 mm (parat3^pes 3.9-4.9 mm), 2.5 times as long as wide; color very dark brown. Frons similar to p. pilifrons except less strongly, less extensively elevated, eleva- tion occupying lower 80 percent of median distance from epistoma to upper level of eyes (115 percent in p. pilifrons) and pubescent area occupying less than 50 percent of median area between eyes (80 percent in p. pilifrons) ; lateral areas much more sparsely, more finely granulate than in p. pilifrons. Pronotum, elytra, and other features essentially as in p. pilifrons. Male. — Similar to male p. pilifrons except frontal vestiture less abundant and shorter, particularly along epistoma. Type Locality. — Mt. Wheeler, Neva- da. Type Material. — The female holo- type, male allotype, and 48 paratypes were taken at the type locality on 10- Vin-74, at 10,000 ft., from Picea engel- manni^ by me. The holotype, allotype, and paratypes are in my collection. This subspecies is named for Dr. T. O. Thatcher who discovered it more than 30 years ago. Araptus attenuatus, n. sp. This species is distinguished from placa- tus Wood by the much coarser strial punctures, by the much longer female frontal vestiture, and by other characters cited below. Female. — • Length 1.6 mm (paratypes 1.4-1.6 mm), 3.0 times as long as wide; color dark brown. Frons feebl}' convex, flattened on medi- an half in some specimens; subshining, rather finely, closely punctured in periph- eral areas, central area minutely irregular, often with a few punctures; a weak medi- an carina on lower half; vestiture of fine, long, white, subplumose setae in punctured area at sides and above, lon- gest setae equal to two-thirds diameter of frons, shorter toward epistoma. Pronotum 1.2 times as long as wide; as in placatus except anterior margin more narrowly rounded, posterior areas more distinctly reticulate, wdth punctures al- most twdce as large; minute setae present on disc. Elytra 1.8 times as long as wide, 1.6 times as long as pronotum; similar to placatus except strial punctures almost twice as large, declivity more narrowly convex, interstriae 2 not impressed, 1 more feebly elevated. Minute strial setae visible on posterior half of disc and decliv- ity except 1 and 2, interstrial setae con- fined to declivity, absent on 2, similar to placatus except usually finer. Male. — Similar to female except frons more distinctly convex, punctures subru- gose, obscurely acicutate, a distinct, sub- tuberculate, median prominence at upper level of eyes and continuing toward ver- tex, its summit transversely etched. Type Locality. — Forty-eight km or 30 miles W Bajia de los Angeles, Baja Cali- fornia Norte, Mexico. Type Material. — The female holo- type, male allotype, and four paratypes were taken at the type locality on Sl-IIL 74, Hopk. 58650, by M. M. Furniss, from host plant No. 88. Twenty-two paratypes are from 32 km or 20 miles N Punta Prieta, Baja California Norte, Mexico, 29- in-73, Pedialanthus niacrocarpus, J. Doy- en. The holotype, allotype, and several para- types are in my collection; two paratypes are in the Canadian National Collection; the remaining paratypes are in the Uni- versity of California (Berkeley) Collec- tion. Araptus fossifrons, n. sp. This common species was thought to be foveifrons (Schedl) until the type of Schedl's species became available for study; however, this species has the elj'tral punctures much finer and the declivital sulcus much more weakly impressed. Male. — Length 2.2 mm (paratypes 1.8-2.4 mm), 2.5 times as long as wide; color rather dark reddish brown. Frons as in foveifrons except lower area more finely punctured. Elytral disc with punctures much finer, interstrial punc- tures usually more widely spaced (vari- able). Elytral declivity with striae 1 rather strongly impressed, interstriae 2 not impressed, with a row of punctures. Vestiture as in foveifrons. Female. — Similar to male except frons weakly convex, a weak median carina March 1975 WOOD: AMERICAN SCOLYTIDAE 31 from vertex to epistoma, surface closely, rather coarsely punctured, with fme abundant, moderately long hair imiformly distributed from epistoma to above eyes; frons concealed in only available female of foveifrons. Type Locality. — Lago Amatitlan, Guatemala. Type Material. — The male holotype, female allotype, and 48 paratypes were taken at the type locality on lO-VI-64, 700 m, No. 702, from the fruiting body of a climbing (Cucurbitaceae) vine, by me. Other parat^'pes were taken in Mexico as follows: 9 at 3 km (2 miles) SE Acatlan, Puebla, 15-VI-67, 1500 m, No. 37; 17 at 9 km (12 miles) SE Oaxaca, Oaxaca, 18- VI-67, No. 57; 4 from 24 km (15 miles) W Armeria, Colima, 30-VI-65, 30 m, No. 141; 9 from 1 km N Atenquique, Jalisco, 24-VI-65, 1000 m. No. 115; 2 from 8 km (5 miles S Atenquique, Jalisco, 25-VI-65, 1000 m. No. 115A; 3 from 8 km (5 miles) W Juchitlan, Jalisco, 2-VII-65, 1000 m. No. 174; 14 from 24 km (15 miles) S Mazamitla, Jahsco, 22-VI-65, 2500 m. No. 96; 13 from Tuxpan, JaHsco, 23- VI- 65, 1300 m, No. 99; 3 from 8 km (5 miles) N Ruiz, Nayarit, 14-Vn-65, 100 m. No. 245; all from fruiting pods of climbing vines or a small tree, by me. The holotype, allotype, and paratypes are in mv collection. Amphicranus parilis. n. sp. This species is distinguished from //'//- formis Blandford by the much smaller size, by the absence of minute crenula- tions on the base of the pronotal disc, and by differences in the elytral declivity in- dicated below. Male. — Length 2.0 mm, 4.0 times as long as wide; color pals yellowish brown, elytral declivity brown. Frons and pronotum as in filiformis ex- cept as noted in diagnosis. Elytra as in filiformis except less strongly explanate, sutural emargination only slightly deeper than wide (twice as deep as wide in fili- formis), declivital spine 2 smaller, less strongly pointed. Type Locality. — Six km or 4 miles W Tepic, Nayarit, Mexico. Type Material. — The male holotype was taken at the type locality on 1 3-VIL 65, 1000 m. No. 240, from a tree branch, by me. The holot}pe is in my collection. Cor thy III s cecropii, n. sp. This abberrant species is distinguished from all other species in the genus by the small antennal club, without sutures, by the absence of female pronotal asperities, and by the minute to obsolete punctures of the pronotum and elytra. Female. — Length 2.4 mm (paratypes 2.0-2.5 mm), 2.3 times as long as wide; color black. Frons uniformh% deeply concave from eye to eye, from epistoma to vertex; sur- face densely, imiformly, very finely punc- tured over entire surface; vestiture very fine, rather abundant, uniformly rather short over concave area, margin above eyes with a dense row of Aery long hair, a small tuft of longer hair at level of an- tennal insertion on lateral half. Anten- nal club 1.6 times as long as wide; asym- metrically obovate, aseptate, entire surface minutely pubescent; posterior face with a small tuft of hair extending about half of club length beyond apex. Pronotum 1.1 times' as long as wide; sides weakly arcuate on posterior half, broadly rounded in front, a distinct, sub- marginal, transverse constriction; anterior margin unarmed; summit indefinite, near middle; asperities absent; surface reticu- late, anterior half with sparse, minute granules, ])OSterior half with sparse min- ute punctures. Acute lateral margins more strongly developed than in other species. Glabrous. Elytra 1.3 times as long as wide, 1.2 times as long as pronotum; sides almost straight and parallel on basal two-thirds, obtusely subangulate behind; disc reticu- late, a few obscure, irregular lines indi- cated, punctures minute, mostly obsolete, apparently confused. Declivity occupying slightly more than posterior third, rather steep, convex; sculpture as on disc, except a few irregularly placed fine granules usually present. Vestiture confined to de- clivity, consisting of sparse, short, fine bristles apparently on odd-numbered inter- striae. Male. — Similar to female except frons convex, a distinct, transverse impression above epistoma, surface smooth, shining. 32 GREAT BASIN NATURALIST Vol. 35, No. 1 impunctate; anterior margin of pronotum distinctly produced toward median line and armed b}' two slender teeth; anterior slope of pronotum much steeper, asperate. Type Locality. — Tapanti, Cartago, Costa Rica. Type Material. — The female holo- type, male allotype, and 15 paratypes were taken at the type locality on 24-X- 63, 1300 m. No. 242", from fallen Cecropia peltata petioles, by me. Eight paratypes are from Turrialba, Cartago, Costa Rica, 5-VII-63, 700 m. No. 19, Cecropia petioles, by me. Three paratypes are from 6 km S San Vito, Puntarenas, 13-III-68, H. Ilespenheide. Three specimens not in- cluded in the type series are from El Laurel (Experiment Station), 12 km SW Caracas, Venezuela, l-V-70, 1300 m. No. 475, Cecropia petioles, by me. The holotype, allotype, and paratypes are in my collection. GENETICS, ENVIRONMENT, AND SUBSPECIES DIFFERENCES: THE CASE OF POLITES SABULETl (Lepidoptera: Hesperiidae) Arthur M. Shapiro^ Abstract. — Polites sabuleii is an example of an insect having a univohine, monophenic high- elevation subspecies and a multivoltine lowland one that produces similar phenotypes only in cold weather. When reared under conditions that induce the warm-weather phenotype in lowland stocks, the montane subspecies P. s. tecumseh continues to produce its usual phenotype, indicating that it has become genetically fixed. One variant of the persistent "genetics- environment" duality in biology concerns the nature of subspecies differences. The problem, as it applies to butterflies, was well simimarized in Klots's (1951) dis- cussion of geographic variation: To what degree much of the recorded geo- graphic variation is a matter of tempera- ture and humidity differences is something which we can only infer. In Papilio glau- cus . . . spring specimens tend to be small and pale. . . . As we go northward we find that in central Canada, where there is only one generation a year, the whole popula- tion looks similarly small and pale. In Can- ada this population has been named as a geographic subspecies, ''canadensis,''' i.e. a part of the species limited to a certain area and showing distinctive characteristics. The temptation is strong to attribute the whole thing to lowered temperatures alone. But suppose we brought a batch of eggs of canadensis down to Florida, and reared the butterflies in the conditions under which the very large, richly colored subspecies australis develops there. Would our cana- densis eggs develop as australis ... or would they develop into the same small pale specimens that their parents were? Twenty-three years later Ehrlich, Holm, and Parnell (1974) could only write that many butterflies have spring generations that are smaller and darker than their summer generations, the difference pre- sumably being due to the seasonal variation in the environment. However, in some northern parts of their range, [they] have only a single summer generation, which is small and dark and resembles the spring generation of southern localities. In the northern populations, the individuals are presumed to have genotypes that produce the dwarfing and darkening. Although the critical transfer experiments have not been done, the greater constancy of the northern forms in the face of environmental changes supports these presumptions [emphasis added] . The same problem was recognized in plants as far back as the 1920s, in the clas- sic work of Turesson (1922, 1925, 1929) later brilliantly expanded by Clausen, Keck, and Hiesey (1940, 1947, 1948, and other papers). This work firmly estab- lished the concept of the ecotype in plant ecology and genetics, a concept more or less readily generalizable to animals in cases like those discussed by Klots and Ehrlich et al. Turesson and Clausen et al. were able, by transplant experiments, to separate phenotypic variation produced directly by the physical environment from that produced indirectly through the se- lection of climaticall}' adapted genotypes. This paper is the second of a series re- porting on analogous studies of North American Lepidoptera. The Subspecies of Polites sabuleti Situations of the sort described above are not limited to populations separated by latitude; many Lepidopterans — like the ]3lants studied by Clausen and his col- leagues— have altitudinal variants, often described taxonomically at the subspecies level, and these are especially interesting because of the short ground distances be- tween the high- and low-elevation popula- tions and the possibility of investigating the nature of their contacts, if any. Polites sabuleti Boisduval is a small, largely tawny skipper (Hesperiidae), widely distributed in western North America. Three named subspecies occur in California: P. s. sabuleti, P. s. tecumseh Grinnell, and P. s. chusca Edwards. The last is a very pale desert population and has not been examined in this study. P. s. sabuleti and P. s. tecumseh are parapatric in northern and central California, occur- ring at low and high elevations respec- tively. 'Department of Zoology, University of California, Davis, California 95616. 33 34 GREAT BASIN NATURALIST Vol. 35, No. 1 P. s. sabuleti is very widespread on sandy soils, in saline and alkaline marsh- lands, and in urban vacant lots. It is usually closely associated with its normal larval host, alkali grass {DisticJilis spicata [L.] Greene, Gramineae), but may breed occasionally on Bermuda grass (Cynodon dactylon [L.] Pers.), an introduced lawn grass and weed. P. s. sabuleti is strongly multivoltine at sea level, with possibly as many as five generations a year. The flight season is Aery long, ranging from late March-April to mid- or late Novem- ber at Sacramento and Suisun City, Cali- fornia. P. s. tecumseh occurs in subalpine mea- dows that become dry in summer, and in alpine fell-fields in the high Sierra Neva- da. Its host plant is not known, but many collectors have noted an association with species of sedges {Carex^ Cyperaceae). Tilden (1959) reports P. s. tecumseh fly- ing from Jul}' to September in Yosemite National Park, which he interprets as in- dicating two broods. The more complete data given in Garth and Tilden (1963) do not support .this interpretation, as there is no wide spread of dates within a given year at a single locality. Farther north, at Donner Pass (7,000 feet) there is no indication of more than one brood. Em- mel and Emmel (1962) found it there from 19 June to 19 August, 1960; Sha- piro found it at the same localities from 1 1 July to 24 August, 1973, and 18 July to 12 August, 1974. The condition of Don- ner Pass specimens does not suggest even a partial second brood. Like many mon- tane butterflies, P. s. tecumseh emerges later at higher elevations; thus, at 10,000 feet it flies mainly in August and into ver}'^ early September. As with the altitudinal subspecies of Phyciodes campestris Behr (N^mphali- dae) previously studied (Shapiro, 1975a), those of Polites sabuleti are separated by a zone in which neither seems to occur. At the latitude of Sacramento, P. s. sabu- leti is unknown as a breeding resident above 1,500 feet and P. s. tecumseh is un- recorded below 5,000 feet. Tilden (1959) confuses the matter by indicating that P. s. sabuleti extends much higher at Yo- semite, but his data (given in Garth and Tilden, 1963) make it plain that this re- fers to the arid east slope only: the rec- ords are from Bridgeport (6,743 ft.) and Mono Lake (6,419 ft.). On the west slope the lowest Yosemite-area record of P. s. tecumseh is Gin Flat (7,036 ft.) and there are no records of P. s. sabuleti at all (al- though it is abundant on the floor of the San Joaquin Valley). The nature of the east-slope contacts has not been studied, but few habitats suitable for either sub- species occur on the abrupt Sierran es- carpment. Polites sabuleti tecumseh differs from summer P. s. sabuleti in being smaller, hairier, and more heavily marked, especi- ally on the hindwing ventrally. The dark markings on this wing are often described as being a "colder," grayer color than in P. s. sabuleti. Although a series of 23 tecumseh from the vicinity of Donner Pass shows considerable variation, no sea- sonal pattern is apparent. In P. s. sabuleti from Sacramento and Suisun City (over 450 specimens examined) there is marked seasonal variation: March-May and Sep- tember-November specimens are, on the average, smaller, darker, and hairier than summer ones, and some are superficially exceedingly similar to P. s. tecumseh, al- though there are minor ( but fairl}^ con- sistent) differences in certain details of the pattern. The phenotypes of wild specimens of both taxa are illustrated in Figures 1-3. Experimental Methods and Results Would stock of P. s. tecumseh reared under conditions that produce summer phenotypes of P. s. sabuleti produce the normal tecumseh phenotype, or would it be modified in the direction of the low- land, summer one? Ova were obtained from a female tecu?nseh collected at Don- ner Pass (7,000 ft), 24 July 1974, and from two female sabuleti collected in a salt marsh at Suisun City, Solano County (10 ft.), 6 August 1974. The resulting progeny were reared side by side in plastic Petri dishes (51/2" diameter X %") at comparable densities (5-8 larvae/dish) under continuous illumination from a 60w bulb at 25C (77F). All larvae were fed fresh cuttings of Bermuda grass (Cynodon dactylon), and mortality in both stocks was negligible. Thirty adult P. s. sabuleti (16 cf 14 ? ) and thirteen P. s. tecumseh (8 cf 5 ? ) were obtained. Continuous light was selected as a regime ecologically nonsignificant to both stocks but known March 1975 SHAPIRO: POLITES SABULETI 35 (i-,-.:^ fei*:^ \K,M£AJ 'Ls^mJ' 4 , * Fig. 1. Dorsal and ventral surfaces of summer specimens of Polites sabuleti sabuleti from the Central Valley of California. ■4/ ^T Fig. 2. Dorsal and ventral surfaces of early spring and late fall P. s. sabuleti from the Central Valley, approaching the phenotype of P. s. tecumseh. •f'''.^:; Fig. 3. Dorsal and ventral surfaces of P. s. tecumseh from Donner Pass, California (7,000 feet), July-August. 36 GREAT BASIN NATURALIST Vol. 35, No. 1 to inhibit diapause in P. sahuleti and re- lated species. The two stocks differed in several re- spects in the laboratory. First- and second- instar larvae oi P. s. sabuleti were yellow- ish green; in the third instar they turned purplish brown; and thereafter they re- mained that color. P. s. tecumseh larvae were purplish brown throughout their de- velopment. At corresponding points in the life cycle the early stages of P. s. sabu- leti were always larger than their high- altitude counterparts. The developmental rates of the two stocks differed very sig- nificantly, with little overlap: from egg to adult P. s. sabuleti took 39-76 days (weighted mean, 58.5 days) and P. s. tecumseh, 70-111 days (weighted mean, 86.0). No diapause was observed in either stock. Normally, high-elevation or -lati- tude stocks of Lepidoptera develop more rapidly than conspecific ones from more temperate climates when reared under uniform laboratory conditions; the rever- sal of this situation in Polites sabuleti is to my knowledge unique in Lepido])teran stocks in which diapause is not manifested in culture. The developmental differences noted above were not mirrored in larval be- havior or morphology. The adults, how- ever, were obviously different and "true" to their normal phenotypes (Figs. 4, 5): nondiapaused tecumseh reared at high temjieratures retained all of their distin- guishing characters, including size. It thus appears that the complex of characters present as a developmental option in low- n Fig. 4. Dorsal and ventral surfaces of representative bred Polites sabuleti sabiileti; continuous light, 25C. Fig. 5. Dorsal and ventral surfaces of representative bred P. s. tecumseh; same conditions as in Figure 4. March 1975 SHAPIRO: POLITES SABULETI 37 land, multivoltine populations is geneti- call}' fixed in P. s. tecumsch, confirming Ehrlich et al.'s prediction. Discussion Three sets of "altitudinal subspecies" have now been investigated in butterflies, representing three different and quite un- related families. They are Picris occident- alis Reakirt and its alpine representative, familiarly (but incorrectly) known as P. o. "calyce^' Edwards (Pieridae); PJiycio- des campcstris Behr and its montane sub- species montana Behr (Nymphalidae) ; and Politcs sahidcti. The first two are dis- cussed at length in Shapiro (1975a). Each species presents a jiicture different from the others. Pieris occidentalis shows xery little, if any, genetic differentiation of the univol- tine, monophenic and bivoltine, diphenic populations of high and moderate eleva- tions, respectively. The alpine stock re- tains the ability to produce an estival phenot}'pe and to develop without dia- pause, and its own phenotype is indis- tinguishable from the vernal one produced downslope. Their mating behavior in- volves male aggregations on mountain- tops, a behavior pattern conducive to gene flow, and in laboratory experiments no reproductive barriers have been found be- tween uni- and bivoltine populations. Phyciodes campestris shows a superfi- cially similar picture, in that the high al- titude subspecies montana, when reared under outdoor conditions at sea level, pro- duces the foothill phenotype rather than its own. However, the cold-season pheno- type of lowland populations is quite dif- ferent from montana, and the ability to produce the montana phenotype appears to be restricted to high-elevation popula- tions. In this case, then, phenotypic plas- ticity is not reciprocal, and the high- elevation population is genetically dif- ferentiated. Polites sabuleti shows the highest de- gree of differentiation yet encountered. The possibility that it has achieved repro- ductive isolation (i.e., speciation) between high- and low-elevation populations can- not be discounted. Because of the differ- ence in developmental time, no crosses between the stocks could be made. They are not known to intergrade anywhere, since the altitudinal discontinuity i)etween them seems to rim the length of the Sier- ra Nevada. These experiments have been duplicated with a latitudinal subspecies pair — Cali- fornia Pieris occidentalis and its subspe- cies P. o. nelsoni Edwards from Fairbanks, Alaska (Shapiro, 1975b). Their pheno- typic differences are clearly heritable, and the res]:)onse of both phenot}'pe and dia- pause to photoperiod has been observed in Fi and F:. hybrids. Studies of the Pieris occidentalis and P. napi L. species complexes (Shapiro, 1975 c) strongly imply that univoltinism is evolutionarily derivative from multivoltin- ism, accompanying the successful invasion of increasingly rigorous climates. High- altitude and -latitude populations of wide- spread species are probably derived from lowland sources, as has been well docu- mented for the Sierran alj^ine flora (Cha- bot and Billings, 1972). The overall pic- ture emerging from these studies supports the suggestion that seasonal ])henotypes of multivoltine populations may become fixed through selection of modifiers in- fluencing thresholds of develo]:)mental ex- pression ("genetic assimilation," Wad- dington, 1953). The somewhat unusual circumstances in Phyciodes campestris montana will be explored in another pa- per. Acknowledgments Collection of livestock for this study was funded by Grant D-804 from the Com- mittee on Research of the Academic Sen- ate, U. C. Davis. Steven R. Sims, John H. Lane, and Adriejine R. Shapiro assisted in field work. Aspects of this research benefited from conversations from Mich- ael Rosenzweig, William E. Bradshaw, and E. W. Jameson, Jr. LiTER' Ci Chabot, B. F., and W. D. Billings. 1972. Ori- gins and ecology of the Sierran alpine flora and vegetation. Ecol. Monographs 42:163- 199. Clausen, J., D. D. Keck, and W. Hiesey. 1940. Experimental studies on the nature of spe- cies. I. Effects of varied environments on western North American plants. Carnegie Inst. Wash., publ. 520. . 1947. Heredity of geographically and ecologically isolated races. Amer. Nat. 81: 114-133. . 1948. Experimental studies on the na- ture of species. III. Environmental responses 38 GREAT BASIN NATURALIST Vol. 35, No. 1 of climatic races of Achillea. Carnegie Inst. Wash., publ. 581. Ehrlich, p. R., R. W. Holm, and D. R. Parnell. 1974. The process of evolution, 2d ed. Mc- Graw-Hill, New York. 378 pp. Emmel, T. C, and J. F. Emmel. 1962. Eco- logical studies of Rhopalocera in a high Sier- ran community — Donner Pass, California. I. Butterfly associations and distributional fac- tors. J. Lepid. Soc. 16:23-44. Garth, J. S. and J. W. Tilden. 1963. Yosem- ite Butterflies. J. Res. Lepid. 2:1-96. Klots, a. B. 1951. A field guide to the but- terflies of North America, east of the Great Plains. Houghton Mifflin, Boston. 349 pp. Shapiro, A. M. 1975a. Ecotypic variation in montane butterflies. Wasmann J. Biol., in press. . 1975b. Photoperiodic control of devel- opment and phenotype in a subarctic popula- tion of Pieris occidcntalis (Lepidoptera: Pi- eridae). Canad. Entomol., in press. . 1975c. Developmental and phenotypic responses to photoperiod in uni- and bivoltine Pieris napi (Lepidoptera: Pieridae) in Cali- fornia. Trans. Roy. Ent. Soc. London, in press. Tilden, J. W. 1959. The butterfly associations of Tioga Pass. Wasmann J. Biol. 17:249- 271. TuRESsoN, G. 1922. The species and the variety as ecological units. Hcreditas 3:100-113. . 1925. The plant species in relation to habitat and climate. Hereditas 6:147-236. . 1929. Zur Natur und Begrenzung der Artenlieiten. Hereditas 12:323-334. Waddington, C. H. 1953. Epigenetics and evo- lution. Symp. Soc. Exptl. Biology 7:186-199. LIFE HISTORY AND ECOLOGY OF MEGARCYS SIGNATA (PLECOPTERA: PERLODIDAE), MILL CREEK, WASATCH MOUNTAINS, UlAH^ Mary R. Gather- and Arden R. Gaufin- Abstract. — During an investigation of some of the stoneflies (Plecoptera) of Mill Creek, Wasatch Mountains, Utah, Megarcys signata, a large omnivorous stonefly, was found to have a univoltine life history and a slow seasonal life cycle. Temperature appears to affect the growth rate of Megarcys signata. Warmer stream temperatures accompany the acceleration of the growth rate, whereas cooler stream temperatures apparently retard the growth rate. Periods of maximum absolute growth rate con-espond with maximum carnivorous feeding from August to September and March to April. Ghironomidae, Ephemeroptera, and Plecoptera, in that order, were the most abundant prey in the foreguts. Young n^-mphs ingested considerable amounts of diatoms, filamentous algae, and detritus but not as much animal matter as did older nymphs. Megarcys signata was unifonnly distributed throughout Mill Creek, except at the lowest station, where few nymphs were found. Emergence occurred in May and June, the peak occurring in June. The mean size of females and males decreased as emergence progressed. This report is part of an eighteen-month study of some of the stoneflies of Mill Creek, Wasatch Mountains, Utah. Be- cause a detailed description of the study area and the methods and materials is given in another paper (Cather and Gau- fin 1975, only a summary is included here. Mill Creek Canyon is located 11 km southeast of Salt Lake City, Utah, in the Wasatch Mountains of the Middle Rocky Mountain Province. Six stations were se- lected along a 12 km length of the stream in the Wasatch National Forest with ele- vations ranging from 1,605 to 2,280 m. The sampling stations were nimibered consecutively. Station I denoting the high- est elevation. The three lower stations (1,605-1,785 m) are easily accessible all year, but the three upper stations (1,995- 2,280 m) are accessible only in the simi- mer and fall. Average minimum and maximum daily flows were 0.3 m^/sec and 1.2 mVsec, respectively, during the study period. Depth averaged 11-45 cm, and current averaged 0.2-0.7 m/sec during the fall when measurements were taken. The substrate of the sampling area at all stations ranged from coarse sand to small cobbles. Minimum water and air tempera- tures recorded during adult emergence were 3 C and 9 C, respectively. Maximum water and air temperatures during this period were 13 C and 26 C, respectively. Water chemistry was similar at all sta- tions. Dissolved oxygen ranged from 6.0 to 8.5 mg/1 (70-120 percent saturation). calcium bicarbonate 109-189 mg/1, calci- vmi carbonate 0-2.4 mg/1, pH 7.5-8.3, total hardness 100-340 mg/1, and conduc- tivity 312-859 mhos/cm. Methods and Materials Nymphs of Megarcy signata were col- lected at least monthly from June 1971 to December 1972 at each of six stations. Additional nymphs were collected in spring 1973 for food habit studies. Hand- screens of mesh sizes 7 and 9 sq/cm were used, the smaller handscreen being used during the majority of the study in an attempt to collect the smaller instars. An area of about 80 cm- of the stream bottom was disturbed in an attempt to collect at least 100 nymphs monthly. All nymphs were preserved in 80 percent ethanol. Adults were collected weekly through- out the emergence period and biweekly during peak emergence using a sweep net and handpicking from vegetation, rocks, and bridges. All adults were preserved in 80 percent ethanol. The interocular distance of all nymphs and adults was measured to the nearest 0.1 mm using an ocular micrometer in a dissecting microscope for determining growth rates and to see if the mean size of the adults decreased as emergence pro- gressed. The nymphs were identified as males and females when possible. Foregut analyses were conducted on 200 nymphs collected from the field. Nymphs were selected from an upper (I) and a ^Study supported by Environmental Protection Agency Traineesliip Grant No. 5T2-WP-542-03. ^Department of Biology, University of Utah, Salt Lake City. 39 40 GREAT BASIN NATURALIST Vol. 35, No. 1 lower (IV) station during each season (Station V had to be substituted for Sta- tion I during winter and spring). The method used followed Swapp (1972). All prey animals were enumerated and identi- fied to order except where family or generic determination was possible. Three types of the family Chironomidae were recognized and designated as species a, b, and c (based on morphology of head capsule). These are discussed under re- sults and discussion but are not separated in Table 1. All algae were determined to genus where possible. Percentage compo- sitions of algae and detritus were esti- mated when present, and dominant items were recorded. A volume analysis was not conducted. Identification of nymphs and adults fol- lowed Gaufin et al. (1966), and nomen- clature followed lilies (1966) and Zwick (1973). Results and Discussion The only systellognathan stonefly pres- ent in Mill Creek in numbers large enough for analysis is Megarcys signata. This species exhibits a slow seasonal type of life cycle. Emergence and oviposition oc- cur in May and June with hatching soon after. Small nymphs (0.5 mm interocular distance) appear in July at the lower sta- tions. Nymphs of comparable size gen- erally appear for the first time in August, September, and October at the upstream stations. Rapid growth occurs from Aug- ust to emergence (Figs, la, b, c). The size lOOn 75 50- 25 2.5- -. 2.0- E o ].o^ 0.5 Month Fig. 1. Growth of Megarcys signata. Arrows indicate emergence. I. O. indicates interocular dis- tance: (a) Monthly mean size as a percentage of total mean size at Station IV; (b) Monthly mean cumulative growth at Station IV; (c) Monthly mean absolute growth rate (data pooled from all sta- tions). A indicates change in I. O. distance. March 1975 GATHER, GAUFIN: UTAH PLECOPTERA 41 frequencies of nymphs and the mean cumulative growth at each station are shown in Figs. 2 and 3, respectively. A comparison of the cumulative growth at all stations reveals that the most rapid growth occurs at the lowest (warmest) stations (Fig. 4). Thus, there seems to be a direct correlation between growth and temperature. Baumann (1967) found no direct correlation in this species in Mill Creek. Seasonally the most rapid growth occurs during the fall and early winter (September- January). Growth apparently slows, but does not stop, during the winter (January-March), increases from March to April, and then decreases from April to May, prior to emergence. Sheldon (1972) reported similar results in his study on the Arcynopteryx species complex he studied in California. However, Schwarz (1970) reported no growth at times during the winter in other Systellognatha. The corre- lation between growth and food habits will be discussed later. 2.0' 101 II J O 1,0 2.0- 1.0- 103 2 A 33 •l 'h 34 27 4 3 0 50 J A 30 ND J F MAM J J ASO Month "n ^ Sfa 2.0-| b IV 1.0-1 1 2.0- O 1.0- 10' AC 15 9 1 38 33 <^h°^n r ^^ ly ,3 1/ ? cf: J A S 5 S 5 ] F M A ?ir Month 9 6 10 5 'l'^ 2 3 I ' 50 5 6 N F Fig. 2. [Frequency distribution of nymphal size classes of M. signata. Number of individuals shown above each polygon; males and females as indicated: (a) Stations I, II, and III; (b) Stations IV, V, and VI. 42 GREAT BASIN NATURALIST Vol. 35, No. 1 Megarcys signata seems to show no preference for either the upstream or downstream stations. This species is even- ly distributed throughout the stream ex- cept at Station VI. Here the substrate is almost entirely cemented. Where the water is deep enough for this large insect, the current is too slow; where the current is fast enough, the water is too shallow and the substrate too homogeneous. More individuals were collected at Station IV than at any other station. The emergence of M. signata began in early May at the lower stations and lasted until late June at the higher stations (Fig. 5). Peak emergence was in June. Bau- mann (1967) found this species emerg- ing from late April to mid- July in Mill Creek. Emergence is progressively later as the elevation increases. Baumann (1967), Hynes (1970), and Nebeker (1971) reported similar results. They were first collected at Stations III and V in early May, when the water tempera- Sta Sta 2.0- O ).0- 2.0- 2.0- 1.0- o - la --HH J ASONDJ FMAMJ JASOND Month T~y ~r~T T— r Fig. 3. Mean cumulative growth of nymphs of M. signata. Vertical line represents size range of nymphs; shaded area represents standard deviation; unshaded area represents standard eiTor of the mean; solid line connects means; rectangle represents emergence period: (a) Stations I, II, and III; (b) Stations IV, V, and VI. March 1975 GATHER, GAUFIN: UTAH PLECOPTERA 43 J a 1 5" ^^ Fig. 4. Comparison of mean cumulative growth of nymphs of (D) M. signata at Stations I-V. Station I represented by (o); Station II by (•); Station III by (■); Station IV by (A); Station V by (A); and Station VI by (D): (a) July, 1971 through June, 1972; (b) July, 1972 through December, 1972. tures were 3 C and 5 C, respectively. Emergence ended in late May (water tem- perature 9 C) at Station V, but lasted until mid- June (12 C) at Station IV and late June (5.5-7 C) at the three upper stations. Megarcys signata is a secretive insect that hides in cracks under bridges or among vegetation to escape warm summer tem- peratures. Clusters of these stoneflies usually containing one female and several males were often collected in these hiding places. Brinck (1949) reported the same phenomenon in related species. The col- lection data probably reflect this secretive habit in that M. signata should have been collected earlier at Station IV. Females and males generally emerged together in a 1:1 ratio. Harper and Pilon (1970) reported similar findings. Only 55 100- 0 ? 100 0 d" 100 0 ^^ 9 rf- 100- 0 ■ d- 100- 0- 1 Arcynopteryx signata Perlodidae June-July Acroneuria pacifica Perlidae Mar.-Sept. >1 Claassenia sabulosa Perlidae June-Aug. Mill Creek, Salt Lake Co. Upper Provo R., Wasatch Co. Lower Provo R., Utah Co. Mill Creek, Salt Lake Co. S. Fork Provo R., Utah Co. Weber R., Summit Co. [•Gaufin, Nebeker, and Sessions, 1966 were 10-12 C, and the experimental groups were established in the first part of November, except where noted below. The specimens were maintained separ- ately in perforated plastic (polyethylene) drinking cups 9.5 cm in depth and 7 cm in top diameter, tapering to 5 cm at the bottom. The cups were suspended and w'ere about half submerged in stainless steel aquaria approximately 100 cm long, 17 cm wide, and 15 cm deep and having a 25.5 U volume. Nine such aquaria were suspended in two refrigerated water baths; four in a bath at 9.5 C and five in another bath at 13 C. Each aquarium was heated with a thermostatically regulated element. Tap water flowed through each aquarium at about 0.4 L/min. (or about one aquar- ium volume/hour). Each aquariimi was aerated with filtered compressed air and agitated wdth a paddle wheel that mixed the water in the aquarium and caused a predominantly up-and-down oscillation of water in the cups. Concentration of dis- solved oxygen remained greater than 90 percent of saturation in the cups. Temper- ature was maintained with a standard er- ror of ± 0.02 C and a range of about ± 0.5 C, at 10, 12, 14, or 16 C. These aquaria were lighted from about one meter above with two fluorescent bulbs on a 12-hr. light, 12-hr. dark cycle. Ambient room light, which was not excluded, varied somewhat. In a separate experiment to estimate the effect of light periodicity, three species were maintained in cups suspended in plastic aquaria flushed with aerated run- ning water (1 vol./hr.) but with no pad- dle wheel (dissolved O^, > 90 percent saturation). These were in a walk-in cold room that maintained the water tempera- ture at 11.5 C within the same limits as in the other aquaria. They were illumin- ated with fluorescent lights (2 bulbs) that delivered about 400 Lux at the water sur- face on either a 12L, 12D (long- day) or a 6L, 18D (short-day) periodicity (Beck, 1968). One or several small stones and several decaying leaves (cotton wood, Populus an- gustifolia; or maple, Acer grandidenta- tum) were kept in each cup. Herbivorous species were also supplied a few pellets of Purina® rabbit chow every few days, and the carnivorous ones were kept supplied with a mixture of small aquatic organ- isms, including amphipods, chironomids, ephemeroptera, oligochaetes, and flat- worms collected from near a local fish hatchery. Changes in weight were monitored either each day between 1 and 3 pm or each week on Thursday between 1 and 5 PM. Each animal was gently blotted with a Kim wipe® (Kimberly-Clark Cor- pororation), air-dried for half a minute, and weighed to the nearest 0.1 mg on a Mettler H6T balance. The standard er- ror of the method, determined by re- peatedly weighing the same animals about 20 times in an hour, was about 0.002 of the mean for animals about 200 mg, about 0.005 of the mean for ones about 150 mg, and about 0.01 of the mean for animals about 30 mg. Repeated weighing revealed a gradual but statistically significant de- crease in weight of each animal even though they were retiu-ned to the water between each determination, so the stan- dard error of single weighings each day or week may have been less than for about 20 weighings an hour. Natural growth rates were calculated for Pteronarcella badia and Arcynopteryx signata from approximately monthly col- lections of 22 specimens of each species from the initial site. The animals were weighed as above, within several hours of collection, and growth Was determined as the rate of change of the average weight. The data were evaluated by regression ' March 1975 BRANHAM, ET AL.: PLECOPTERA GROWTH 53 analysis (Rao, 1958; Bailey, 1959; Alder and Roessler, 1968), using a Hewlett Packard 9100 B computer and pro- grammed procedures supplied by the manufacturer. Growth trends of each mdi- vidual were analyzed from weight data by determining the correlation coefficient (r) and, from it, the possibility that changes in weight were correlated linearly with time, according to the calculated slope (m) which intercepted the axis representing weight at the hypothetical initial weight (b) independent of fluctuations at the be- ginning of the observations. Growth was judged to be positive, negative, or insignif- icant from the correlation coefficient (P > .05), and its magnitude was deter- mined from the slope. Absolute growth values, in weight units, were converted to relative ones for comparisons between animals by determining the rate of change as a percentage of the averaged initial weight (m x 100/b). The effects of vari- ous conditions were then evaluated by t- test or regression analysis, using the rela- tive growth rates. Short-term Fluctuations in Weight The weight of all stoneflies examined every day increased and decreased rhythmically, typically varying 3 to 25 percent of the body weight about every 4-6 days (Table 2; Figs. 1, 2, and 3). The magnitude of weight difference between high and low periods exceeded the prob- able error due to the method, and the trends apparent in the plotted data indi- cate that the fluctuations were not merely artifacts of the method. There was no apparent correlation between the rhyth- mic pattern and environmental conditions, nor were the cycles of different animals Table 2. Periodic variations seen with daily weighings (Average ± SE) calculated from the first six cycles. Period (days No. of between animals peaks) Amplitude* P. californica 5 5.9 ± .4 3.3 ± .3 P. badia 4 4.5 ±1.0** 6.4 ±1.6 A. pacifica 5 5.0 ± .4 7.2 ±1.0 C. sabulosa 1 4.7 5.7 A. signata 1 6.3 12.3 -^X Fig. 1. Weight changes of a Pteronarcys ca fornica kept under various conditions This md vidual was collected in December and acclunatu d to laboratory conditions for 15 days without too. It was then weighed daily while being starved A 14 days), fed rabbit chow B (15 days), and the leaves and rabbit chow C (21 days), while being ept at 10 C. It was then changed to 16 L an weighed daily while being fed leaves and rabbit how D (50 days) and finally weighed once eac week for 10 weeks (E). It did not molt and w s still alive after the 170-day observation period. in phase with each other, even though they were in the same aquarium. It was hypothesized that periodic feed- ing behavior caused the observed rhyth- mic weight changes. To test this, ten Pteronarcys californica (collected 5 De- •Difference between peaks and troughs, expressed as per- centase of average weight. >•, • •Significantly (p < .05) different only from P. calijormca •Significantly (p < .001) different only from P. cahlonuca Fig 2. Daily weight changes before and a: er molting (Pteronarcys californica and Acroneun pacifica) and before emergence (Arcynopteryx si nata) . The molted cuticle was found and the an mal weighed about six hours after the molt occui ed. 54 GREAT BASIN NATURALIST Vol. 35, No. 1 Fig. 3. Daily weight changes before death. cember 1972) were starved for 15 days without weighing; then weighed every day for 14 days, while starving; then fed either Purina rabbit chow or decaying leaves and weighed daily for 15 days; and, fin- ally, fed the other food and weighed daily. Starvation did not eliminate the rhythmic weight changes (Fig. 1), although all the animals were losing weight. The periodi- city remained unchanged upon feeding either or both foods. The amplitude was not affected significantly (P > .05) when leaves were supplied but was significantly increased over the starvation level after feeding rabbit chow, either alone or with leaves (Table 3). The rhythmic weight changes could have been associated with molting. Weight changes during molting were observed in a few specimens. Of the 16 animals weighed daily, five molted one time suc- cessfully, one emerged, and six died molt- ing. One Ps. californica and one Acro- neuria pacifica were weighed during a successful molt. The Ps. californica was 24.4 percent heavier at the time of molt- ing than just before, and the A. pacifica was 19.0 percent heavier. The ones that died molting were all considerably heavi- er than they had been (Table 8) . Molting generally occurred when the pattern of weight change indicated that a peak weight was due to occur. After molting, the period between weight maxima was usually shorter and the amplitude greater than it had been before the molt; but af- ter several cycles, the pattern began to resemble the premolt condition again. The one animal on a daily weighing schedule that emerged (an A?-, signata, Fig. 2) also molted and emerged at a peak in the weight cycle. The adult was 25 percent lighter than the nymph. Its gut was empty. The frequency of weighing affected the weight of stoneflies. Animals weighed a number of times in an hour lost weight during the course of the observations. Five animals that had been weighed every day gained weight appreciably when the weighing frequency was reduced to once a week. The grovvi;h rate (m) was usu- ally slightly greater with .the less frequent weighings, but the change in rate was not statistically significant (Fig. 1). Hand- ling could have caused the animals to contract and expel water or gut contents, and it likely interfered with feeding pat- terns. Long-term Growth Patterns The stoneflies under observation lived for various periods. Those that lived a month or more either gained, lost, or re- mained the same weight during their Table 3. Effects of food on periodic variation, seen with daily weighings of P. californica (aver- age ± SE). Period (days between peaks) f Amplitude (difference be- tween peaks & ti-oughs, expressed as a % of aver- age wt.) fEstimated from three peaks IfThree that did not respond to cither food were e.xcluded fr JOnly those fed rabbit chow after leaves responded •Significantly different (p > .01) from starved group only ♦•Significantly different (p > .001) from starved group only om tnese cali Gain after feeding (% of previous wt.) Starved 10 5.1 ± 1.0 2.2 ± .3 Fed leaves 3it 5.6 ± 1.0 3.1 ± .3 4.3 ± .3 Fed rabbit chow ^\ 5.3 ± 1.2 5.3 ± .4** 11.8 ± .6 Fed both 7+f 5.1 ± 1.2 4.4 ± .5* 8.7 ± .5t March 1975 BRANHAM, ET AL. : PLECOPTERA GROWTH 55 Table 4. Average (X ± SE (N) ) Weeks of life. Long Day Short Day IOC 12 C 14 C 16C (11.5 C) (11.5 C) p. californica\ 25.2±3.0(5) 17.2±2.1(6) 24.5 ±4.9(6) 23.8±4.6(6) P. badia 15.0 + 2.5(5) 17.7±1.3(6) 11.4±2.1(6) 10.7+1.6(6) 6.0±1.1(11) 6.7+ .6(11) A^. cinctipes 4.7± .9(6) 6.2±1.2(6) 2.5 ± .3(6) 4.7 ± .9(6) A. pacifica 18.9±5.8(5) 16.5 + 3.5(4) 18.0±4.3(6) 14.2±4.3(5) 15.6±1.9(11) 11.1±2.6(11) C. sabulosa 17.0± .1(2) 17.5±1.5(2) 9.7±1.1(3) A. signata 9.5 + 3.1(4) 9.3±1.7(4) 2.7+ .6(7) 2.8+ .4(12) 3.4± .9(7) 2.7± .6(7) fEight Ps. californica and three Ac. pacifica were still nlivo at the tune of wiiting, 33 weeks after the observations began. life. Some individuals lost weight for part of their life and gained in another part. It was important to consider the history, length of life, and ultimate fate as well as the overall growth patterns in assessing the effects of temperature on growth rate. Some species lived longer in the labora- tory than did others (Table 4). In gen- eral the smaller, univoltine species, Ne- moura cinctipes. Pa. badia, and Ar. signa- ta, did not live as long as the larger spe- cies, Ac. pacifica, Claassenia sabulosa, and Ps. califo7-nica, which probably live for several years as nymphs (Table 1). Pter- onarcella badia and Ar. signata lived for significantly (P > .05) less time at higher temperatures. Pteronarcella badia, Ar. sig- nata, and Ac. pacifica kept in the light- control chambers died significantly sooner than their counterparts in the tempera- ture experiments. The former were caught in winter from streams near 0 C and were kept without stirring. Day length had no significant effect on the life span of Ar. signata and Pa. badia. Weight changes prior to death followed three distinctive patterns, which, in some specimens, could have been related to cause of death (Fig. 3, Table 5). Often death occurred during a recognizable molt. Sometimes the growth curve turned sharp- ly up, as if molting, but there was no ex- ternal sign of molting. More often, death followed a diminuation in the amplitude of cyclical weight changes and was not marked by any sharp change in weight. Two Ps. californica that did not change in weight upon being supplied with either food showed this pattern, and it may have reflected starvation. The third pattern was marked by a sharp loss of weight at death. Adults weighed less after emerg- ing (Fig. 2), and such sharp terminal weight loss could have indicated unsuc- cessful emergence. The circumstances of death of all animals that died are pre- sented in Table 5. Periodic molting is a characteristic of Arthropod growth. However, many of the animals observed here apparently did not molt, while many others died in the process (Table 5). No A^. cinctipes and only one Ar. signata were observed to molt. The occurrence and frequency of molting for the other species are enumer- ated in Table 6. Some molts were prob- ably overlooked, but the analysis of growth curves suggests that most were detected by the presence of the cast cuticle. The times until first molt, between molts, and until death for ones that did not molt are compared in Table 6. Time before the first molt (considering ones that molted successfully or that died molt- ing) was quite variable and was probably a function of the condition of the animals at the time of capture. The time between molts was also highl}' variable. The aver- age period before the first molt was not significantly different from the time be- tween molts for any species. The aver- age length of life of individuals that did not molt was also the same as the average period before the first molt and the period between molts. The average period be- fore molt differed between species: Ps. californica ^= C. sabulosa > Ac. pacifica > Pa. badia (t-test, differences considered significant if p > .05). There was no cor- Table 5. Circumstances of death (percentage of N). N Mt It 2t 3t Et P. californica 31+f 8 26 50 15 0 P. badia 52 21 20 53 3 2 N. cinctipes 22 0 0 100 0 0 A. pacifica 40ft 26 11 52 10 0 C. sabulosa 8 57 0 28 14 0 A. signata 42 2 11 74 11 2 fl Growtli curve tiuTicd up at death, as prior to molting 2 Growth cun-e continued imchanged until death 3 Growth cun'c turned down at death, as prior to emergence M Died molting E Emerged IfFom- additional P. californica and two .'1. pacifica were accidentally killed. 56 GREAT BASIN NATURALIST Vol. 35, No. 1 Table 6. Occurrence and frequency of molts Number observedf Weeks (average ± SE (range ) molting Before first Of life of ones Ox Ix 2x 3x molt Between molts not molting P. californica P. badia A. pacifica C. sabulosa 16 ± 3 (12-30) 5 ± 1 ( 1-18) 9 ± 1 ( 2-21) 15 ± 1 (13-17) 15 ± 3 (10-19) 7 ± 1 ( 1-11) 9 ± 1 ( 2-18) 17 ± 2 (3-32) 5 ± 1 (1-19) 8 ± 1 (3-22) 6 ± 3 (3- 8) fMolting recognized by finding a cast cuticle or found in the process of molting. relation between size and the length of time before or between molts either with- in or between species. There was also no correlation between the length of time before molts and the temperature or light period for any species examined. Changes in weight accompanied molting (Table 7). Most animals weighed during a molt (alive or dead) showed a dramatic increase over the premolt weight. Usually this increase was much greater than the longer-term increase (determined by com- paring the average weight the month be- fore molt wdth the average weight for the month afterward) and could have been as- sociated with the mechanism of molting. Animals that molted several times, and thus apparently were adapted to labora- tory conditions, grew appreciably between molts. Some of the ones that died with- out molting grew about the same amounts as did others between molts, suggesting that death could have resulted from fail- ure to molt. For each animal observed, the growth rate (calculated as the regression coef- ficient [m] ) , for the four weeks prior to molting usually was different from that for the month after molt (Table 8). There was a great deal of variability between individuals: some lost before and gained after; some gained before and lost after; some gained or lost more rapidly before than after; and vice versa. One Ps. cali- fornica lost 1.0 percent/ week for 20 weeks, molted, and then gained 1.1 percent/week for 15 weeks. The growth patterns in Fig. 2 are from apparently normal animals that lived many months in the laboratory. The number that molted and lived at least a month afterward was small, so averages and limits (Table 8) do not in- dicate significant (p > .05) differences in average pre- and postmolt rates. There was no significant correlation with the controlled parameters of temperature or light. The overall growth of each animal un- til death was evaluated by calculating the correlation coefficient (r) and the regres- Table 7. Change in weights with molting. Percent change in weight (average ± SE (range) ). N During moltf N With moltfl N Between moltsf N Total, ones not moltingft P. californica 4- 26 ± 2(22-30) 6 25±5(9-41) 3 22.6±4.9(13.1-29.1) 28 19.4± 10.1 (-22.8-174.3) P. badia 10 65 ± 6(36-85) 10 12±4( 5-32) 5 9.8±5.7( -1.2-20.4) 30 21.6± 6.9(-25. 1-144.4) A. pacifica 12 22± 4( 3-47) 12 2±2(-9-19) 8 15.6±5.5( 0.7-48.6) 15 -3.8± 1.6(-14.4- 9.2) C. sabulosa 4 36±14(18-78) 3 1±7(-8-15) 0 110.0 fAnimals weighed during a successful molt or found dead in the process of molting. Weight during molt as a percentage of the last premolt weight. tfAverage weight for the month after molt as a percentage of the average for the month preceding the molt. JCalculated from the growth curve. Slope multiplied by the time and e.\pressed as a percentage of the intercept (i.e., the hypothetical initial weight) . ttTotal growth calculated as m x weeks of life x 100/b. Table 8. Growth rates (percentage change/week) before and after molt (m ± SE (range) ). Month before molt Month after molt P. californica P. badia A. pacifica All Long day Short day C. sabulosa -.16 2.73 .84 (-3.35- 2.05) 1.34 (-2.37-10.69) .54 ± .41 (-2.68- 6.32) .96 ± .99 (-3.68- 3.46) -.02 ± .50 (-2.53- 1.69) .38 ± 2.07 (-2.71 - 4.33) 2.79 ± .81 ( .09- -.39 ± 1.29 (-9.76 5.75) 5.75) 1.17 ± .56 (-3.23-10.98) 3.23 ± "1.20 (- .05-10.96) -.30 ± 1.00 (-3.23- 1.69) -2.04 ± .78 (-3.57- -1.08) March 1975 BRANHAM, ET AL.: PLECOPTERA GROWTH 57 sion coefficient (m) in 0.1 mg change/ week. The overall growth rate was some- times influenced strongly by the final phase of the growth curve as the animal expired. Growth during the first two months was therefore calculated separate- ly. To simplify comparison between spe- cies and experimental groups (Table 4) the growth rate (m) was expressed as a percentage of the calculated initial weight (b, the intercept). The number of animals was so small and the range of values so great within each group that averages and standard errors are of little meaning. Re- gression analysis of rates of change against temperature, however, indicated that tem- perature had a significant effect on the growth rates of some animals (r signifi- cant at p > .05). The proportionate (per- centage) effect of temperature on growth rate, exclusive of experimental error, is expressed as the coefficient of determina- tion (r- X 100) (Alder and Roessler, 1968). The relationship between differ- ence in growth rate per degree Celsius is expressed by m. These observations were apparently val- id only for the animals collected in Octo- ber and November, when the stream tem- peratures were still about 10 C. The few animals collected later in the year from colder streams died in a shorter time and generally grew at rates different enough from the others' that it seemed best to ex- clude them from the preceding calcula- tions. Light periodicity could also have af- fected development in the laboratory. Significantly more Pa. badia molted under long- than short-day conditions. There was no significant difference in length of life or growth rate between animals of either species held under long- or short- day conditions. Acroneuria pacifica lived longer under long-day conditions (15.5 dr 2.0 vs 10.2 ± 2.3 weeks) but molted sooner under short days (7.1 ± 1.5 vs 9.1 ±1.5 weeks). Significantly more molted under long than short days. Their growth rate to death was significantly greater under long days (1.9 ± .8 percent/week) than under short days (-1.0 ± .3 per- cent/week) (Table 6). These animals were not strictly comparable to the ones used for the temperature experiments: the water in their tanks was not stirred, and they were collected later in the year (Pa. badia and Ac. pacifica in November and Ar. signata in February). Pteronar- cella badia and Ar. signata kept under equal periods of light and dark died signif- icantly sooner than ones under similar light and temperature conditions but col- lected earlier and kept in stirred water (Table 4). Laboratory conditions were quite differ- ent from nature, so it was desirable to compare growth of laboratory and wild animals from natural populations where possible. Pa. badia and Ar. signata per- sisted at the initial collecting sites in popu- lations of relatively uniform-sized indi- viduals. Their growth in nature was quite linear for both species between October and May 9 (Fig. 4). The average per- centage of weight increase per week of wild Pa. badia was 6.26 ± 0.5 (r = .998), and that of Ar. signata was 10.17 ± .07 (r = .938). Such an analysis was im- practical with the larger species because of the simultaneous existence of several year classes and the effect of investigator selection. Wild Ar. signata emerged na- turally in May (water temperature 5-8 C) and Ps. californica. Pa. badia, and Ac. pa- cifica in May (water temperature 10-11 C). Discussion Stonefly naiads varied in weight as time passed. Some of the variation was associa- Fig. 4. Growth of Pteronarcella badia and . rcynopteryx signata in the stream. Means (wit SE) were calculated from periodic samples of 22 nimals and converted to percentage of the averag initial weight. ^H GHKAT BASIN NATURALIST Vol. 55, No. 1 tcrj wilh short-torm IJuftu^jtiori in state, vvhiJo some represented long-term "growth." Variation in gut content rapid- ly altered the weight of experimental ani- ]nals--as much as 10 percent or more n^'ig. 1, Table 3j. Cyclical weight changes, on the order of 5 percent variation every 5 days (Tables 2 and 3), occurred even in the absence of feeding and (ould have resulted from the amount of water in the animal, in either its gut or its tissue. This cyclical weight change could have been .issociated with molting. Each animal I hat was observerl daily and that molted did so wlieii a predicted peak wciight liould have occurred, and animals weighed ( onsiflerably more wlien mfjiling llian just before or after the molt riabb; 7). Such rai)id wciight gain could be rc- spoiisibb; for splitting the f)id cuticle and rilling out the new ont! (Wiggbisworth, l')f)5j. The laihifc of molting was a common (ause of death of animals in th(? labora- tory. Many animals died during or just after molting. Others flied at a time of peak weight, considerably above the pre- vious average, as if about to molt (b^ig. 1 Table 5). Most of the; animals died with- out any molts iil()s(/ ri'able fi). On the average, llios(> tbat did not molt died approximately wlicu they should have moltcnl, as judged iioiri the average time before or Ix'tween molls, '['heir average growth was also about the same as the average growth be- tween molts of animals that survived in som(! s[)(H;ies (Table 7). No A^. cinclipcs and only oru; Ar. siirnala wer(> observed to molt, and their averag(i life was bvss tlian lor tlu! other species (Table 4). It seems likely that failure oi some as])cct of molt- ing was a common cimse of death of stonefli(!s kept in the laboratory. Relatively few animals lived long enough with repealed molts (Table 1) to be con- si in the next 8 weeks, and emefged in .fanuary, 5 months i)rema~ tin-ely. Its growth was almost linear from the l)eginning of the observations (r =: .<)9H) and slightly less (m = 5.2 percent/ week) than the average for the species in nature. Most of the P. hatlia that molted successfully and then lived for at least a month gained weight prior to, and as a re- sult of, the molt, but then lost weight (Ta- \ bles 6, 7j. Food may have been deficient, or the newly molted ones may have been less tolerant of laboratory conditions. Two P. californica that lived for the entire 55- week observation period molted twice each. Both lost weight the month before the first molt and showed no significant change the month before the second molt. Both gained considerable weight at both molts, remained at a higher weight, and ( on tinned to grow at an accelerated rate after the molt (Fig. 2, Tables 6, 7, and 8). They apparently did not feed before the molt, expanded in volume during the molt, and, at the larger volume, hardened and began to eat. The four Ac. pacifica that lived for the duration of the observations usually were not growing significantly prior to molt, expanded appreciably dur- ing the molt, but then declined to a weight considerably below the premolt weight within a day or so. They then grew rap- idly for several weeks, until the previous weight was reached, after which they did not grow significantly again until after anolh(!r molt. Some regressed at molting i and lost weight overall (Table 7) (Fig. 2, Table 9). Probably their pattern was not normal but reflected subsistence in an un- natural environment (Beck and Bharad- waj, 1972). The aniicipatcHi |)remature emergence (Nebeker, 1971 j was not found. Of the animals coIUhUhI in October, only one (Pa. hadia) emerged and only a few died with a pattern of weight changes even sug- gesting em(>rgence (Table 4). This could hav(^ l)(HMi because the animals were col- lected early in the autumn and kept at constant temperatures and long-daylight conditions at or above those existing at the time of collection. Perhaps some en- vironmental cue was absent. Or possibly the small j)lastic cups in which the ani- mals were kept were too confining to per- mit normal behavior. A number of P. calijornica collected from streams near O (' in January, acclimatized to 16 C, and kept (onnnunally in fish-breeding nets in the laboratory did emerge in March, three months before the wild population (>merg(Hl naturally. One Ar. signata, col- lected in February, emerged 22 days after being put in a cup and kept at 10 C (Fig. 2). The lenglli of lile of Pa. hadia and Ar. March 1975 BRANHAM, ET AL.: PLECOPTERA GROWTH 59 22^ O o, +1+1 +1 +1 +1 rO 00 05 00 in -^ !>■ 00 T-! ro c>i '■ ?s T-< ^ CO csi ^ ^' Cs)^ ^ ^ -I- ^ vno^ o-i^o vn^^ (Tj^o (oxoo i^'^^ ^^o oooo oo xnxo 00 to 00 00 to to ^ +1+1 +1 +1 +1 +1 8^^ 00 00 in s ^csic4 fOOO vn ^ CO'^T-i tOQDO roO- tJ- O O ^ ^ O O +1 +1 +1 +1 00 -^ ^ ^ o inxn o +1 +1 fv^ oo>n 00 '^' oi 1,3^ Tj-^o in^«-> i^ oooo in rh 00 CS) (^ to too s § ^ 04 in at ^cr, to t^ ro ss 00 CO ro -- 04 +1 +1 +1 +1 +1+1 +1 +1 +1 +1 +1 +1+1 +1 +1+1 +1 li 04 CO ro O 04 00 ;: So^ OOO inoi ^ ^ ^^§ §¥8 cr> O) 04 01 o to '^ '■ doi oOfO -^ ■«^ ^ CO ' ^^ iniri CNi t0 0.4fO tOtOO ^^-^ ■^-^O ■4 '^ in in CT) 'J- OO 00 o +1 +1 +i po in 00 00 o O^ in 00 t>-c. (1942, 1944, 1946, 1949), whose papers reviewed most Nearctic gen- era of the subfamilies Psilopinae, Noti- philinae, and Parydrinae. His mitimely death precluded the completion of this series and left the synoptic study of the subfamily Ephydrinae largely unfinished imtil Sturtevant and Wheeler's review in 1954. This review was the last compre- hensive treatment of the genera considered here. Prior to Sturtevant and Wheeler's pa- per, the American species of Coenia or Paracoenia had not been treated together. Coquillet (1902) and Curran (1927) de- scribed the first Nearctic species, Coenia bisetosa and C. turhida respectively, and Johnson (1925) included C palustris (Fal- len) in his list of Ephydridae from Massa- chusetts. Johnson's identification of the latter species as C. palustris is question- able, and Sturtevant and Wheeler include this citation under C. curvicauda Meigen. After studying the Ephydrinae in the Naturhistorisches Museum, Wien, Cres- son (1930) reviewed the European species of Coenia. He distinguished C. curvicauda from C. palustris and designated a lecto- type for C. curvicauda. Cresson (1935) described a new genus, Paracoenia^ that included two new species, platypelta and fumosalis^ in addition to Coenia bisetosa and C. turbida, described previously from North America, and two European species, C. futnosa (Stenhammar) and C. beckeri (Kuntze). C. curvicauda and C. palustris were left in the geiuts Coenia. Sturtevant and Wheeler described one additional species, C. paurosoma. in their review of 1954. Cresson delimited Paracoenia from other genera based on comparative dif- ferences he noted in the dimensions of the head, in the number of dorsocentral bristles (three in Coenia, four in Para- coenia), and in the presence {Paracoenia) or absence {Coenia) of well-defined hum- eral bristles. Most specialists have con- tinued to recognize both Coenia and Para- coenia c[S waWA genera. Dahl (1959) stud- ied the male genitalia of both genera and suggested that the marked differences be- tween them supported Cresson's view. The two genera were also recognized by Wirth (1965) in the catalog of North American Diptera. Sturtevant and Wheeler, how- ever, recognized only Coenia, relegating Paracoenia to subgeneric status. This revision is based primarily upon a comparative study of the male postabdo- men and a reevaluation of previously rec- ognized characters. From these studies, I generally concur with Cresson's concept of Coenia and Paracoenia; however, I now recognize three subgenera in the lat- ter genus, Paracoenia, Calocoenia, and 'Department of Entomology, Oregon State University, Con-allis, Oregon 97331 65 66 GREAT BASIN NATURALIST Vol. 35, No. 1 Leptocoenia. The basis for these proposals will be treated in greater detail in the sections on ])hylogcny and under the ap- propriate generic discussions. Four new s])ecies are described, three in Parncoenia and one in Cocnia. Acknowledgments This study was initially suggested b}- Dr. Stephen L. Wood while I was an un- dergraduate at Brighani Young University. Many of the preliminary observations were" completed then, although the stud}' has since been expanded as new species were discovered and specimens were ex- amined from a wider geographic area. I thank Dr. Wood for his encouragement and for sponsoring two summers of field work. Special thanks are extended to Dr. Paul H. Arnaud. California Academy of Scien- ces, to Dr. Willis W. Wirth, Systematic Entomology Laboratory, ARS, USDA-Na- tional Museum of Natural History, Wash- ington, D.C., and to Mr. Guy E. Shewell, Canadian National Collection of Insects, for their extensive cooperation and assist- ance. T^'pe specimens were borrowed from the Academy of Natural Sciences of Phil- adelphia (Drs. David C. Rentz and Selw>n S. Roback); the National Museum of Na- tural History (Dr. W. W. Wirth); and the Museum of Comparative Zoology (Mrs. Janice C. Scott and Dr. John F. Lawrence) . In addition to the above, the following institutions and curators kindly loaned specimens, wdthout which this study could not have been completed: LTniversity of Minnesota (Dr. Philip J. Clausen); Kent State University (Dr. B. A. Foote); Uni- versity of California, Riverside (Dr. Saul I. Frommer) ; Los Angeles County Mu- seum of Natural History (Dr. Charles L. Hogue); Iowa State University (Dr. Rob- ert E. Lewis); Cornell University (Dr. L. L. Pechuman) ; University of Califor- nia, Davis (Dr. R. 0. Schuster); Wash- ington State University (Dr. William J. Turner) ; Florida State Collection of Ar- thropods (Dr. Howard V. Weems, Jr.); and Brigham Young UniAorsity (Dr. Ste- phen L. Wood). Dr. Marshall R. Wheeler generously loaned specimens from his pri- vate collection. I also wish to gratefully acknowledge Drs. John D. Lattin, Carol A. Musgrave, Paul Oman, and Paul O. Ritcher, Oregon State University, for their technical and editorial assistance and for many stimu- lating discussions on systematics. The stereoscan electron micrograph was taken by Mr. Alfred Soeldner, Oregon State Uni- versity. Dr. Willis W. Wirth, USDA- USNM was also kind enough to review this study. Methods and Discussion of Characters All observations were made using a stereomicroscope; a filar micrometer was used for the measurements. Morpholog- ical characters, especially those of the male postabdomen, w^ere illustrated using an ocular grid. All illustrations were drawn to the same scale on mylar draft- ing film. Preparation of the male or fe- male postabdomen for study involved its removal and heating in a 10 percent so- dium hydroxide solution to remove ex- traneous tissue. The abdomen was then washed, further dissected, and compared. For permanent storage the abdomens were preserved in plastic microvials filled with glycerin and attached to the appro- priate specimen. During the course of the study, I ex- amined approximately 4,000 specimens, including the type specimens of all but Cocnia curvicauda. and I did examine European specimens of this species. The type specimens examined formed the basis for the species descriptions. Species descriptions are purj)osefully brief; for the most part they summarize specific differences or additions not found in the more detailed generic descriptions. The diagnoses will differentiate the spec- ies from similar taxa. For previously rec- ognized species, the descriptions also con- lain any newly acquired information for comparative purposes. Polymorphic and polytypic variations are included under re- marks. Characters considered in this study are ffom all bod}^ tagma and have been quan- lifi(>d where appropriate. Ratio values are based on an average of ten specimens se- lected because of obvious size differences. Head. — Eye-to-cheek ratio. This is the ratio of genal height to^ eye height. Meas- urements are taken from the head in pro- file. This ratio is a convenient character for some species groups. March 1975 MATHIS: AMERICAN EPHYDRIDAE 67 Width-to-height ratio. This ratio is cal- culated as head height to head width; measurements are made from a cephalic orientation. Eye-width-to-foce-length ratio. This ratio is hased on measurements from the head in profile and is calculated as face length to eye width. Height-to-length ratio. This is the ra- tio of the height of the head in profile to its length, measured from the most ante- rior surface of the face to the posterior margin of the eye. Cresson first used this character when describing Paracoenia. Aristal -pectinations. The length of the pectinate branches on the dorsum of the arista is compared with the base width of the arista. This character is sometimes difficult to use and is best seen in well- preserved specimens. Interfoveal hump and marginal bristles. This character is correlated with the eye- to-cheek ratio. It is the comparison of the hump height to the length of the bristles along the oral margin. Hump height is related to genal height. Facial color. This character is subject to considerable variation in many species, especially species of Paracoenia, but it is of some diagnostic value in others. The species of Coenia and Calocoenia ha^e rel- atively constant facial color. Postocular bristles. The development of the dorsalmost postocular bristles is useful in distinguishing Paracoenia from Coenia. Thorax. — Acrostichal hairs. The ar- rangement and degree of development of the acrostichal hairs have been overlooked as a diagnostic character other than at the species level. These characters are important in distinguishing the subgenera of Paracoenia. Dorsocentral bristles. The number of dorsocentral bristles has been extensively used as a major character and was ac- corded significance at the generic level by Cresson. Humeral bristles. The presence or ab- sence of well-developed humeral bristles also was used by Cresson to delimit these genera. I have followed Cresson in at- tributing generic importance to this char- acter as well as to the number of dorso- central bristles. Halters. The color of the halters can be used to distinguish some groups of spec- ies. Cresson (1930) mentioned that this character is usually variable and is not important as a key character. Costal vein ratio. This is the ratio of distance along the coastal margin between Ri and R:,+,; to the distance between R2+3 and Ro+i. All measurements are the maxi- imum straight-line distances. Mi+2 ratio. This is the ratio of the dis- tance of the Mi+o anterior to the posterior cross vein to the distance posterior to the posterior cross vein. Costal bristles. The presence or absence of costal bristles along the dorsal and/or ventral surface is diagnostic of some gen- era. Calocoenia is the only taxon of Scatel- lini with prominent bristles on both sur- faces, a character found in many species of Eph3drini. Femoral comb. The femoral comb is a sexually dimorphic character restricted to the males of Paracoenia s. str. Abdomen. — Male postabdomen. The male genitalia previously have not been used as characters at the species or generic level; I have found them extremely use- ful at both levels. These characters are discussed more fully in the generic and specific descriptions. Female ventral receptacle. The shape of this structure seems to be of consider- able diagnostic value, especially at the generic level. Phylogeny and Classification Both Coenia and Paracoenia belong to Scatellini as it is presently characterized. The tarsal claws are curved and short, and the pulvilli are developed normally. The tribal concepts, however, have not been reassessed since Wirth (1948, 1970, 1971), Oliveira (1954a, 1954b, 1957), and others (Collin, 1963; Steyskal, 1970) be- gan incorporating characters of the male postabdomen in their treatments of vari- ous Ephydrinae genera. This is especially evident with the annectant genus Austro- coenia Wirth from South America as well as several und escribed genera from the neotropics, which will require further evaluation before a reliable classification of the higher categories can be achieved. Biological information and systematic studies of the immature stages would al- so be most useful. Because biological and morphological data of the larvae are lacking in many re- 68 GREAT BASIN NATURALIST Vol. 35, No. 1 lated genera and in some of the taxa here- in considered, I have elected to recognize a conservative classification in this study. Taxa above the species level but within the generic limits of Paracoenia as Cres- son described it are given subgeneric stat- us. But this status is provisional, await- ing the accumulation of additional in- formation as outlined above and further assessment. The subgenera of Paracoenia are pri- marily based on characters of the male postabdomen and correlated external fea- tures. The resulting concepts are suffi- ciently distinct to be easily recognizable as delimited in the diagnoses and as seen by reviewing the figures. Paracoenia {Paracoenia), for example, is the largest subgenus with eight species, yet each known taxon belonging to this category can be readily placed without difficulty. Further, most of the diagnostic charac- ters are apomorphus and define mono- phyletic groups. On the other hand, the relationships be- tween subgenera are somewhat obscure and the generic concept is not as neatly circumscribed. More reliance is ])laced on chaetotaxy characters of doubtful signifi- cance. However, coupled with our mea- ger knowledge regarding biology, habitat, etc., I feel that the genus is convenient and does reflect a cohesive unit. Coenia has only three species, all of which are evidently closely related. Ex- cept by association with males and in some instances with locality, the females of one species are generally indistinguish- able from those of another. Similarly, the males closely resemble each other, al- though their genitalic characters are con- sistent and constant, a fact that facilitates identification and classification. This group in particular needs biological study. Taxonomy Key to Coenia and Paracoenia Four pairs of dorsocentral bristles; humeral bristles well developed; dorsal- most postocular bristles subequal to verticals .. Paracoenia Cresson Three pairs of dorsocentral bristles; no developed humeral bristles; dorsal postocular bristles much smaller than verticals Coenia Robineau-Desvoidy Genus Paracoenia Cresson Paracoenia Q-esson, 1935, Trans. Amer. Ent. Soc. 61:356. Type-species, Coenia bisetosa Coquil- lett. by original designation. Sturtevant and Wheeler, 1954. Trans. Amer. Ent. Soc. 79:164- 166 (review of Nearctic species as subgenus of Coenia). Wirth. 1965, USDA Agricultural Handbook No. 276, pp. 755-756 (catalog). Diagnosis. — Members of this genus are similar to those of Coenia but can be distinguished from the latter as follows: Postocular bristles immediately posterior to the vertical bristles subequal to verti- cals; at least one humeral bristle well de- veloped, much larger than the surround- ing setae; foiu- pairs of dorsocentral ])rist- les. Description. — Small to large, length 2.1 to 4.4 mm, females usually larger than males; dark species, often with sub- shining metallic reflections; head with characteristic arched prefrons. Head. Front (postfrons) rectangular, wider than long; margins of mesofrons directed inward anteriorly; mesofrons subshining with metallic reflections, setu- lose; ocellar triangle equilateral, concolor- ous with fronto-orbital areas, dull, micro- sculptured. One large pair of proclinate diverging ocellar bristles; postvertical bristles various; two pairs of fronto-orbi- tal bristles; both inner and outer vertical bristles well developed; two pairs of strong postocular bristles immediately pos- terior to vertical bristles. Antennae dark brown to black, pollinose; second segment setulose, especially on median and ventral surfaces; dorsum of arista pectinate, pec- tinate branches up to three times the width of aristal base, sometimes equaling third antennal segment width. Pruinose face (])refrons) protruding, arched with inter- foveal hump, setulose; setae descending from hump and along ventral margin strongest. Eye suboval; gena variously do veloped with genal bristle toward ventral margin. Mouthparts dark; prementum large, bulbous. Thorax. Dorsum of mesonotum sub- shining to dull, generally concolorous, sometimes with discernible median and lateral stripes. Acrostichal hairs in ap- proximately six irregidar rows to two dis- tinct rows; four pairs of dorsocentral brist- March 1975 MATHIS: AMERICAN EPHYDRIDAE 69 les; one pair of intra-alars; two to three pairs of humeral bristles; two pairs of notopleurals; one pair of presuturals; one pair of supra-alars; two pairs of post-alars; dorsum of scutellum convex to flat, setu- lose; at least two well-developed pairs of lateral scutellar bristles; one ])air each of mesopleural and sternopleural bristles. Wings transparent to infuscated; costal setae weak to strong, on dorsal and/or ventral margins. Legs dark, pollinose to subshining. Abdomen. Abdomen of males with five visible tergites, females with six to seven, subshining to pollinose; most species with pollinose band near the posterior margin of each tergite; fifth abdominal tergite of males with anteroventral ])rojection in some species of Paracoenia s. str. Female postabdomen with three complete seg- ments, six, seven, eight; ninth segment with tergite not fused dorsally, with one pair of long spines on ventral margin; sternite of ninth segment also with one pair of spines; female abdomen terminat- ing with cerci. Ventral receptacle vari- ously shaped. Male postabdomen sym- metrical, reduced; sixth segment with spiracles only although European species of Paracoenia s. str. have a transverse sclerotized band posterior to fifth sternite which could represent sixth sternite; seventh and eigth segments absent. Spir- acles one through six present, sixth spir- acle in membrane between fifth abdomin- al tergite and epandrium, all other spira- cles in ventral margin of respective terg- ites. Epandrium suboval with scattered setae; dorsum of epandrial plate bearing two setulose cerci; ventrally epandrium terminates at juncture with prominent, projecting surstyli of various shapes. Hy- pandrium extending dorsally and ven- trally, attaching to epandrium, lateral hypandrial process sheathing aedeagus; aecteagus well sclerotized, pointed apically. Discussion. — For purposes of classifi- cation the species of Paracoenia are ar- ranged in three subgenera, Paracoenia, Calocoenia, and Lepiocoenia. The latter two subgenera might be accorded generic status by future revisers since the male postabdomens and other characters are fairly distinctive. But before the generic classification of these taxa is assessed, the higher classification of the subfamily Ephydrinae should be revised on a world- wide basis to insure consistency in generic concepts, especially the distinguishing gap. Such a review will necessarily entail a great deal of descriptive work because of our spotty knowledge of many faunal areas, i.e., the Neotropical Region, which are replete with undescribed species. The following key works best for male specimens and includes both European species. Illustrations of the male genitalia will facilitate accurate identification. Key to Paracoenia subgenera and species Acrostichal hairs in two rows; dorsum of scutellum slightly convex to flat; posteroventral margin of mesofemora without row of comblike bristles in males 2 Acrostichal hairs in four to six irregular rows; dorsum of scutellum con- vex; posteroventral margin of mesofemora with dense row of bristles in males subgenus Paracoenia Cresson 3 Length over 3.25 mm; eye-to-cheek ratio 1:0.25 or larger; well-developed costal bristles projecting anteriorly from ventral and dorsal surfaces subgenus Calocoenia, Caloceonia platypelta (Cresson) Length under 3.00 mm; eye-to-cheek ratio 1:0.25 or less; bristles along cos- tal margin not developed subgenus Leptocoenia^ Lepiocoenia paurosoma (Sturtevant and Wheeler) Fifth abdominal sternite of male deeply U-shaped, Figs. 11, 12; Palearctic .... 4 Fifth abdominal sternite broadly U-shaped, Figs. 1-6; Nearctic 5 Length approximately 5 mm; dorsum of thorax and abdomen grayish-blue to light brown, dull; pleura gray P. beckeri (Kuntze) Length usually less than 4 mm; thorax and abdomen darker, bluish-olive green to greenish-gray; subshining dorsally P. fumosa (Stenhammar) 70 GREAT BASIN NATURALIST Vol. 35, No. 1 5. Postocellars small, no longer than their distance apart at base; facial pru- inosity yellowish-gold with greenish-blue metallic reflection showing through around interfoveal hump; ventral projection of fifth abdominal tergite in males blunt, parallel to remainder of ventral margin .... P. fumosalis Cresson Postocellars larger, longer than their distance apart at base, facial pruin- osity various; ventral projection of fifth abdominal tergite absent or not as above 6 6. Process of fifth abdominal tergite in males not developed 7 Process of fifth abdominal tergite developed into a projection of various shapes 8 7. Subshining with metallic blue reflections; eye-to-cheek ratio 1:0.45 or larger; associated with hot sulfur springs P. calida n.sp. Subshining with metallic green reflections; eye-to-cheek ratio 1:0.3 or less; associated with thermal or cool water P. turbida (Curran) 8. Process of fifth abdominal tergite of male broadly produced; acrostichal hairs few; male genitalia as in Fig. 1 ; presently known only from vicin- ity of Los Angeles, California P. ampla n. sp. Process of fifth abdominal tergite not as broadly produced; acrostichal hairs stronger; surstyli of male genitalia not pointed distally or bare 9 9. Surstyli thickened basally, at least one-half total length; sheathing projec- tion of hypandrium broadly rounded apically; smaller, length 3.1-3.6 mm; setation less well developed, especially on face and mesonotum; male genitalia as in Fig. 6 P. ivirthi n. sp. Basal expansion of surstyli less than one-third total length; lateral process of hypandrium pointed; length 3.7-4.4 mm; facial and mesonotal hairs well developed P. bisetosa (Coquillett) Subgenus Paracoenia Cresson hump, strongly arched; eye-to-cheek ra- n ■ n ,n:>r- rr . t7 . c tio usually 1:0.22-0.45; width-to-height Paracoenia Cresson, 19i5, Irans. Amer. Ent. boc. ^- a r\ rn • ^^^ \ r i ..i 61:356. Type-species Coenia bisetosa Coquil- ratio 1 : 0.67; eye-width-to-face-length ra- lett, by original designation. Sturtevant and tio 1:0.45 (1:0.85 in P. callda) ; height- Wheeler, 1954, Trans. Amer. Ent. Soc. 79:164- to-length ratio 1:0.93. Thorax. Dorsum subshining although Diagnosis. — This subgenus is similar subdued in aged specimens, generally to Calocoenia but may be distinguished as shining more posteriorly. Acrostichal follows: acrostichal hairs in several ir- hairs in four to six irregular rows; other- regular rows, no prominent hairs; dor- wise chaetotaxy as in generic description; sum of scutellum convex; costal setae scutellum convex. Costal vein ratio 1:0.2; weak, in a single row; postero ventral mar- Mi+.. vein ratio 1:0.90; costal setae gen- gin of mesofemora with distinct row of erally weak, developed setae on dorsal bristles in males; many species with an margin only. Mesofemora of males with anteroventral projection of the fifth ab- j)osteroventral row of comblike bristles, dominal tergite; epandrium of male with Abdomen. Subshining to pollinose, if an anteromedian triangular projection; shining, reflection somewhat less than aedeagal apodeme subquadrate. dorsum of scutellum. Female ventral re- Description. — Moderately large, ceptacle with operculum, wider than high, length 3.1-5.0 mm; dark species, often extending process not longer than oper- with subshining metallic reflections. cukmi length. Surstyli of male postab- Head. Front rectangular; mesofrons domen projecting from lateral margins subshining to shining with metallic re- <^f epandrium, long, variously shaped; a flections; pectinate branches of arista up triangular process lies between surstyli, to throe times the width of aristal base, ^"me species, with a median groove (see often equaling third antennal segment figures of included species), width. Face with prominent interfoveal Discussion. — The species included in March 1975 MATHIS: AMERICAN EPHYDRIDAE 71 this taxon form a fairly homogeneous group based mostly on apomorphous char- acters (see diagnosis). The joint posses- sion of these characters dehmits the mono- phyletic grouping here understood as Para- coenia s. str. and determines the sub- generic concept. Two species previously included here, P. paurosoma (Sturtevant and Wheeler) and P. platypelta Cresson, are sufficient!}' distinct to fomi the basis for new subgenera. Paracoenia s. str. is Holarctic, but no one species is presently known to occur in both Eurasia and North America. How- ever, many species of the subgenus are widely distributed, and it is not uncom- mon to collect two or more species from the same general locality-. Other species such as the Nearctic P. calida and P. am- pla and the Palearctic P. beckeri are known only from very localized geo- graphic areas. The subgenus contains eight species; six are Nearctic and two are Palearctic. All of the Nearctic species except P. fum osalis are found principally in western North Ainerica. A more detailed zoogeo- graphic account, aside from the general distributions indicated under the appro- priate species, will not be possible until more collection data become available. The paucity of biological studies pre- cludes a detailed accounting for the en- tire subgenus. For the Nearctic region, only P. turhida has been studied in any detail (Brock et al, 1968, 1969). How- ever, Dr. B. A. Foote and associates at Kent State University and Dr. Karl W. Simpson of Cornell University are cur- rently engaged in studies of various eph- ydrid species that will greatly enhance our biological knowledge. Most species can tolerate harsh environ- ments, especially aquatic habitats with high concentrations of various salts. These shore flies are often abundant, for exam- ple, along the margin of Great Salt Lake or associated with hot sulfur springs in Yellowstone National Park, Wyoming. Scheiring and Foote (1973) further re- port finding larvae in the shoreline mud of alkaline lakes and in sewage-impreg- nated mud. The larva and pupa of P. fumosa. a Palearctic species, were de- scribed by Beyer (1939). Map 1. — Distribution of Paracoenia wirihi, filled stars; Paracoenia platypelta, filled circles; Paracoenia ampla, open circle; and Paracoenia calida, open star. Paracoenia (Paracoenia) ampla, n. sp. Fig. 1, Map 1 Diagnosis. — Although this species is quite similar to P. hisetosa, it can be readily distinguished from the latter by comparing male postabdomens. The sur- styli of P. ampla are sinuate, bare, and strongly narrowed apically. Further, the shape of the lateral hypandrial process is distinctive, especially the clavate ex- tension. This species is larger than most P. bisetosa and in general is less setulose. The acrostichal hairs, in particular, are weak and widely scattered. Description. — Length approximately 4.0 mm (the abdomen was removed for dissection before measurements were tak- en); generally subshining with bluish- green metallic reflections. Head. Width-to-height ratio 1:0.7; height-to-length ratio 1:1; eye-width-to- face-length ratio 1 : 0.44; facial pruinosity brownish-gold. Thorax. Setae generally weak, scattered; costal vein ratio 1:0.17; Mi+o ratio 1:1. Abdomen. Fifth tergite ventrally pro- duced into broadly based processes that extend anteriorly to basal margin of fused 72 GREAT BASIN NATURALIST Vol. 35, No. 1 U-shaped fourth and fifth abdominal sternites, processes pointed apically; fifth abdominal sternite thin, weak, deeply U- shaped with parallel arms; epandrium subquadrate; medial triangular projection proportionately small to epandrial size; surstyli directed inward, apically nar- rowed, sinuate, bare; hypandrium in pro- file almost rectangular, with extending process slightly clavate; aedeagus broad basally, tapering rapidly, curved and pointed apically. Male genitalia as in Fig. 1. Distribution. — Los Angeles, Cali- fornia. Types. — Male holotype with the fol- lowing label data: Los Angeles, Cal., Apr 29th, 1915, M. VanDuzee; a determina- tion label, Coenia hisetosa Coq., 1919, Cresson; a blue M C VanDuzee collection label. The type will be deposited with the California Academy of Sciences, type number 12032. Remarks. — This species is known only from the unique male holotype. Recogni- tion of the specimen as representing a new species is justified in view of the very distinctive male postabdomen. Un- fortunately, P. ampla inay already be ex- tinct due to the tremendous and rapid urban growth in the Los Angeles area since 1915. I have examined several Para- coenia specimens from Los Angeles County but none were ampla. Paracoenia {Paracoenia) hisetosa (Coquillett) Fig. 4, Map 3 Coenia hisetosa Coquilllett. 1902. J. N. Y. Ent. Soc. 10:183 Caenia [sic] hisetosa: Aldrich, 1905, Smithson. Misc. Coll. 66(1444): 631 Paracoenia hisetosa: Cresson, 1935, Trans. Amer. Ent. Soc. 61:356 Coenia {Paracoenia ) hisetosa: Sturtevant and Wheeler, 1954, Trans. Amer. Ent. Soc. 79:164 Types. — Male holotype, Salt Lake, Utah, 25 June, E. A. Schwarz collector. The type is deposited with the National Museum of Natural History, Washington, D.C., type number 6644. This specimen is in relatively good condition, although the wings are ragged and torn. Diagnosis. — P. hisetosa is similar to P. turhida. but the former is larger, more setulose, and more brownish. The poste- rior margin of the fifth abdominal stern- | ite is sclerotized and of uniform thickness throughout. The Aentral process of the fifth abdominal tergite is well developed and pointed. Basally, the surstyli are en- larged but narrow quickly, making the lateral margin sinuate. The hypandrial process usually tapers evenly to a rounded point. Outwardly, this species might be confused with P. ampla, but the male genitalia of P. hisetosa differ sufficiently from the latter that recognition of either species should not be difficult. Description. — Length 3.7-4.4 mm; dark greenish-brown dorsally; laterally quite pollinose. Head. Mesofrons with bronze metallic reflections. Eye-to-cheek ratio 1:0.34; width-to-height ratio 1:0.69; height-to- length ratio 1:0.9; eye-width-to-face- length ratio 1:0.5. Thorax. Dorsum with pollinose an- terior, becoming subshining posteriorly; pleural areas largely pollinose. Wings infuscated with light brown. Abdomen. As in diagnosis and Fig. 4. Specimens examined. — 1881. Distribution. — Like P. turhida, this species is primarily a western North American taxon, although collecting data indicate eastern extensions to New York (5 miles W Cardiff), Pennsylvania (Phil- adelphia), Delaware (Bombay Hook), and Virginia (Saltville). The Northwest Ter- ritories (Nyarling River), Canada, is the northernmost collection site, and the spec- ies ranges from there southward through most of the Midwest to Texas (Buffalo Spring Lake) and into Mexico (Guada- lupe Can., B. Calif.). Westward, speci- mens have been collected in every state and Canadian pro\ince west of the 100th ])arallel. Remarks.^ — This is the most common species of the genus and among the most widespread. It is also one of the most variable. Facial pruinosity color runs from bright brownish-orange to silver, and the general body color varies from shining greenish-brown to a subdued, grayed green. Age polymorphism is also apparent; older specimens are often more brownish and are worn. Dr. Willis W. Wirth has made several collections of P. hisetosa from aquatic habitats with varying concentrations of both alkaline and saline salts. March 1975 MATHIS: AMERICAN EPHYDRIDAE 73 Map 2. — Distribution of Paracoenia fumosalis, filled circles; Paracoenia turbida, filled stars; and Coenia alpina, enclosed stai-s. Paracoenia {Paracoenia) calida, n. sp. Fig. 3, Map 1 Diagnosis. — This is the most distinc- tive Nearctic species and is easily separa- ted from all others of the genus. Exter- nally, the blue metallic reflections from the dorsum, the protruding prefrons, and the eye-to-cheek ratio are diagnostic. The male postabdomen resembles that of P. bisetosa but differs in the shape of the hypandrial process, which is more or less of uniform thickness and has a noticeable taper just before the apices. Additionally, the fifth abdominal tergite does not have a ventral extension, although the margin is pointed. P. calida and P. bisetosa are approximately the same size. Description. — Length 3.4-4.3 mm, holotype male 3.9 mm, generally dark, gray pollinose on head and thoracic plurae with subshining blue metallic reflections dorsally. 74 GREAT BASIN NATURALIST Vol. 35, No. 1 Head. Eye-to-cheek ratio 1:0.45; height- to-length ratio 1:1; width-to-height ratio 1 : 0.71 ; eye- width-to-face-length ratio 1:0.85. Fronto-orbital and ocellar triangle areas blackish-gray, concolorous with mar- gins of frons; postocellar bristles weak in some specimens; pruniose face grayish- tan. Thorax. Pleural areas dull, pollinose; dorsmn subshining to shining, brownish- blue; halters dark, reddish-bro\^^l to black; wings completely infuscated, brown to smoky. Tarsal claws well developed, as long as third or fourth tarsomere; pulvil- lar pads proportionately small to claw size. Abdomen. Dorsum of all segments with distinct blue to purplish-blue reflections, reflections stronger in general than else- where on body; ventral margin of fifth abdominal tergite broadly pointed, with- out lobelike projection; fifth abdominal sternite broadly U-shaped, narrow pro- jecting arms forming obtuse angle; setae along posterior margins of tergites much larger than rest of setae, at least twice as long, in some female specimens three to four times as long. Surstyli of male genitalia with more or less gradual taper, not sinuate or pedunculate; aedeagus short; hypandrial process as described in diagnosis. Distribution. — Wilbur Hot Springs, Colusa Co., California. Types. — Male holotype, allotype, and all paratypes are from the type locality. Two male and 9 female paratypes, H. J. Jacob; 1 male and 1 female para type, 27 June 1950, L .W. Quate; holotype, allo- type, and 205 male and 96 female para- types, 25 June 1974, W. N. Ma this. Pri- mary types will be deposited with the U. S. National Museum of Natural History, type number 72975. Male and female paratypes will be deposited with the Cali- fornia Academy of Sciences, Canadian National Collection, Academy of Natural Sciences of Philadelphia, Kent State Uni- versity, Washington State University, and Oregon State University. The remaining paratypes are in my collection. Remarks. — Of all the Nearctic species of Paracoenia s. str., P. calida is perhaps the most remarkable. Its known distribu- tion is limited to a hot sulfur spring in the foothills just east of Clear Lake, California. The larvae develop and mature in all but the hottest water where they can easily be collected in great numbers. The adults are also abundant and were often ob- served to congregate in large clumps near the spring source where shaded or pro- tected areas could be found. Empty pu- paria that are scattered on the surface of the effluent and along its margins are often utilized as oviposition sites. Figure 13 is a stereoscan electron micrograph of the egg of P. calida. The effluent of the spring emptied into a small creek around which swarming numbers of other ephydrids were encoun- tered on emergent grasses and in quieter eddies on the water's surface. P. calida, however, was not common there, and only an occasional collection was made away from the hot springs. It is also of interest that a new saldid species was recently described from the same locality (J. T. Polhemus, 1967). The specific name, calida, is descriptive of the habitat. Paracoenia {Paracoenia) fumoscdis Cresson Fig. 2, Map 2 Paracoenia fumosalis Cresson, 1935, Trans. Amer. Ent. Soc. 61:356 Coenia {Paracoenia) fumosalis: Sturtevant and Wheeler, 1954, Trans. Amer. Ent. Soc. 79:164 Types. — Male holotype, Rockport, Massachusetts, 28 August 1913, C. W. Johnson collector. The type is deposited with the Boston Natural History Society (Museum of Comparative Zoology, Har- vard University, Cambridge, Massachu- setts), type number 31759. One male and two female paratypes are topotypical. Four female paratypes were collected on Nantucket Island, 13 July 1926 by C. W. Johnson. All paratyj)es are deposited with the Academy of Natural Sciences of Phil- adelphia. Diagnosis. — Cresson (1935) stated that this species is similar to P. hisetosa and P. funiosa, which is Palearctic. My observa- tions agree with Cresson's, although in many respects P. fumosalis is unique among Paracoenia species. The males are most easily distinguished from similar taxa by the rounded, fingerliko projection of the fifth abdominal tergite. This pro- cess is parallel to the ventral margin of the tergite that is deeply incised and of uniform thickness before the slightly en- larged, rounded apex. The fifth abdomin- March 1975 MATHIS: AMERICAN EPHYDRIDAE 75 Map 3. — Distribution of Paracoenia bisetosa, enia curvicauda, filled circles. al sternite is subrectangular without ex- tending arms from the posterolateral mar- gins. The surstyli are narrowly S-shaped, and the median triangular process often has a median groove that is cleft apically. The hypandrial process is much longer than the aedeagus, and apically it is trun- cate. Externally, P. fumosalis differs from all others in the length of the post- ocellar bristles, which are usually shorter than their distance apart at the base. filled stars; Coenia paurosoma, open stars; and Co- Further, the tannish-bronze color of the pruinose face seems to be constant. Description. — Length 3.5-4.0 mm; dark brown, lightly pollinose to subshin- ing dorsally; some greenish-blue metallic reflections. Head. Mesofrons shining with bluish reflections; fronto-orbital areas subshining brown; face concolorous with mesofrons, pruinose, tannish-bronze. Eye-to-cheek ratio 1:0.3; width-to-height ratio 1:0.64; 76 GREAT BASIN NATURALIST Vol. 35, No. 1 height-to-length ratio 1:0.87; eye-width- to-face-length ratio 1:0.41. Thorax. Dorsally subshining, purplish- blue reflections, dark brown. Pleural areas pollinose except dorsal margin. Wings in- fuscated \\dth brown. Abdomen. Concolorous with dorsum of thorax. Male postabdomen as in diagno- sis and Fig. 2. Specimens examined. 368. Distribution. — P. fumosalis is pre- dominately a nothern and eastern North American species. I have examined ma- terial from Alaska (Matanuska and Eagle River flats) and from all of the Canadian provinces except British Columbia and the Yukon Territory. In the continental United States, P. fumosalis ranges west- ward to Montana (Libby), eastward through the Great Lake states, some mid- western states (Nebraska, Iowa), and in- to the Northeast. It has also been col- lected as far south as Florida (Archbold Biological Station, Lake Placid), and it presumably occurs between Florida and the Northeast. Wirth (1965) lists Cali- fornia as the westernmost extension of P. fumosalis, but I have not seen specimens from California. Remarks. — Examination of over 350 specimens of this species from a wide selection of localities within its distribu- tion revealed very little morphological variation. This species is very uniform except for slight artificial size differences reflecting the mode of preservation. The facial coloration seems to be a consistent diagnostic character unlike other wide- spread species of Paracoenia. I suspect that the species is quite vagile and that the apparent uniformity is due to exten- sive genetic exchange. Although I do not know of any locali- ties in which this species is associated with hot springs, it has been collected around alkaline and saline habitats. Scheiring and Foote (1973) reared specimens and report finding larvae in mud impregnated with organic sewage. Paracoenia {Paracoenia) turhida (Curran) Fig. 5, Map 2 Caenia [sic] turbida Curran, 1927, Can. Ent. 59:91 Coenia {Paracoenia) turbida: Sturtevant and Wheeler, 1954, Trans. Amer. Ent. Soc. 79:165 Paracoenia turbida: Wirth, 1965, USDA Agri. Handbk. No. 276:756 Types. — Male holotype and allotype, Old Faithful, Yellowstone National Park, Wyoming, 30 September 1924, N. Crid- dle. Two male and one female paratype have the saine label data as the type. All types are deposited with the Canadian Na- tional Collection, type number 2370. Diagnosis. — Externally, this species re- sembles P. ivirthi and is sometimes con- fused with P. hisetosa. However, it differs from both in the shape of the hypandrial process, which apically narrows more abruptly although the apex is rounded. Also, the lateral margins of the surstyli do not taper apically as rapidly as P. hi- setosa nor are they enlarged basally as in P. wirthi. Outwardly, P. turbida is smal- ler than P. bisetosa but of approximately the same length as P. ivirthi. However, the acrostichal hairs are weaker in turbi- da. The fifth abdominal sternite is more similar to that of bisetosa, although the more sclerotized posterior margin is not as wide nor as uniform in thickness as it is in bisetosa. Description. — Length 3.25-3.75 mm; dark greenish-brown, subshining dorsally. Head. Fronto-orbital areas dark brown, pollinose to subshining. Eye-to-cheek ratio 1:0.3; width-to-height ratio 1:0.65; height- to-length ratio 1:0.93; eye-width-to-face- length ratio 1:0.44. Thorax. Anterior area of dorsum slight- ly pollinose, becoming subshining to shin- ing posteriorly; pleural areas in general more subdued than dorsum, especially along margins. Wings nearly transparent to light brown. Abdomen. As in diagnosis and Fig. 5. Specimens examined. — 674. Distribution. — The majority of col- lection localities are west of the Rocky Mountains, although they do extend east to Nebraska (Cherry Co., Big Alkali Lake), Iowa (Ames), and Ohio (Kent, 5.6 miles SE). Specimens have been col- lected as far north as Alaska (Circle Hot Springs) and southward through Canada (British Columbia to Manitoba) and the western United States into Mexico (60 km S Tijuana). Remarks. — Strength of setation, gen- oral body color, especially the facial pru- inosity, and overall size show considerable variability. Polymorphism within a sin- March 1975 MATHIS: AMERICAN EPHYDRIDAE n Figs. 1-5. — Male and female genitalia. 1 a,b Paracoenia ampla, 2 a,b,c Paracoenia funiosalis; 3 a,b,c,d Paracoenia calida; 4 a,b,c Paracoenia bisetosa; 5 a,b,c Paracoenia turbida. Fig. a, ventral view of cerci, epandrium, and surstyli; Fig. b, lateral view of cerci, epandrium (epn), surstyli (sur), aede- agal apodeme (aeg ap), hypandrial process (hyp pr), and aedeagus (aeg); Fig. c, ventral view of male fifth abdominal steniite; Fig. d, lateral view of female ventral receptacle. gle population of P. turbida seems to var}^ as greatly as the total species variance. Character displacement was not apparent in areas where turbida occurs sympatri- cally with other Paracoenia species. Brock, et al. (1968, 1969) have studied the biology of turbida from near the type locality in Yellowstone National Park, Wyoming. They found that both larvae and adults feed on blue-green algae and the filamentous bacteria of mucilaginous mats, which develop under a variety of hot spring flow conditions. Fly activity is generally restricted to cooler mats (from 30-35C), where most egg laying occurs. Above 40C the eggs fail to hatch and first instars die. At 35C the life cycle takes approximately 14 days (egg to egg), 78 GREAT BASIN NATURALIST Vol. 35, No. 1 and mature females can produce 100 or more eggs a day. Turbida can exploit transient islands of available resource quickly, and the lar^'ae soon decimate the optimum habitat (Weigert and Mitchell, 1973). Wiegert and Mitchell also an- alyzed the interactions between the algal mats and turbida and between turbida and a mite parasite Partnuniella thermalis Viets. Mitchell and Redmond (1974) de- scribe the egg of turbida (several stereo- scan electron micrographs) and suggest inechanisms of respiration under varying environmental conditions. Paracoenia turbida, however, is not endemic to hot springs, and the details of their feeding habits and habitat preferences under dif- ferent conditions could vary considerably. More notes on the biology of this species may be found in Scheiring and Foote (1973). Paracoenia {Paracoenia) wirthi, n. sp. Fig. 6, Map 1 Diagnosis. — This species resembles both P. bisetosa and P. turbida and in many respects is intermediate. As in P. bisetosa, the ventral margin of the fifth abdominal tergite is produced into a pointed extension, but the projection is not as long as that of P. bisetosa. The general coloration and setal characters more resemble P. turbida; however, males of P. wirthi are destinct from either spec- ies in the shape of the surstyli, the hy- pandrial process, and the fifth abdominal sternite. Basally, the surstyli are wide, a condition that extends to about one-half their total length. The hypandrial pro- cess is broadly produced apically with a bluntly rounded apex. The fifth abdomin- al sternite is broad, and the projecting arms are subparallel and short. Description. — Length 3.1-3.6 mm; subshinning with bluish-green to green metallic reflections. Head. Mesofrons greenish-blue, shin- ing; pruinose face grayish-tan; eye-to- cheek ratio 1:0.22; width-to-height ratio 1:0.6; height-to-length ratio 1:0.87; eye- width-to-face-length ratio 1:0.44. Thorax. Acrostichal hairs in three to four rows anteriorly, becoming irregular with five to six rows posteriorly. Pleural areas although subdued not grayed. Wings in many specimens almost transparent, in others smoky infuscate. Abdomen. Metallic reflections green to olive green. Male genitalia as in diagnosis and Fig. 6. Distribution.^ — This species is found in the Sonoran desert from southern Cali- fornia and northern Baja California east- ward into Arizona. It extends south into Mexico (Distrito Federal, Mixquic). Types. — Male holotype, allotype and 21 paratypes (7 males, 14 females), Cali- fornia, Inyo Co., 1 mile N Tecopa Hot Springs, 24 June 1974, Wayne N. Mathis. Thirty-nine paratypes as follows: Cali- fornia, Inyo Co., Tecopa Hot Springs, 16 May 1965, W. F. Barr, 1 male; Inyo Co., Shoshone, 24 June 1974, W. N. Mathis, 1 male, 5 females; Inyo Co., Shoshone, 1 October 1935, A. J. Basinger, 1 male, 3 females; San Diego Co., Mtn. Palm Springs, Anza Desert, 2 March 1964, 1 male. Arizona, Pima Co., Lowell Ranger Station, 6-20 June 1916, 32° 18.5' N, 110° 49' W, ca. 2,700', 1 male, 1 female; Bill Williams Forest, August, F. H. Snow, 1 male, 1 female. Mexico, Baja California, Guadalupe Canyon, 19 May 1957, F. X. Williams, 1 male, 3 females; Distrito Fed- eral, Mixquic, 9 km SW Chalco, 4 August 1965, K. R. Valley, 4 males, 16 females. The type, allotype, and 8 paratypes will be deposited in the California Academy of Sciences, type number 12033. The re- maining paratypes will be deposited with the U. S. National Museum of Natural History, Cornell University, the Canadian National Collection, The Academy of Na- tural Sciences of Philadelphia, and my collection. Remarks. — Although P. wirthi ap- pears to be intermediate in many charac- ters, their consistency, especially the male genitalia, justifies recognition as a new species. As with other species of the genus, P. wirthi is sometimes associated with hot springs, usually along the mar- gins of the effluent. P. wirthi is named to honor Dr. Willis W. Wirth for his contribution to the systematics of the Ephydridae and for his helpful, encouraging responses to my many questions. Calocoenia, n. subgen. Type-species: Paracoenia plalypelta Cresson, 1935, monobasic Diagnosis. — Although similar to Para- coenia s. str. and Leptocoenia, Calocoenia March 1975 MATHIS: AMERICAN EPHYDRIDAE 79 Figs. 6-10.— Male and female genitalia. 6 a,b,c,d Paracoenia ivirthi; 7 a,b,c,d,e Paracoema plaly- pelta; 8 a,d,e Coenia curvicauda; 9 a,h,c4,e Paracoenia paurosoma; 10 a,b,c,d,e Coenia alpina. Fig. e, lateral view of internal genitalia; otliers as in caption of 1-5. may be distinguished from either as fol- lows: Externally, Calocoenia differs from Paracoenia s. str. in the arrangement of acrostichal hairs, which are in two dis- tinct rows, by the absence of a postero- ventral comb of bristles along the male mesofemora, by the flattened scutellum, by the prominent costal bristles project- ing anteriorly from both dorsal and ven- tral margins, and by the generally uni- form, concolorous abdomen which in male specimens is more noticeably narrowed apically. The most apparent difference between Calocoenia and Lcptocoenia is size; Calocoenia is nearly twice the length of most Leptocoenia. Further, the eye-to- cheek ratio of Calocoenia is at least 1:0.2 and the M1+2 vein ratio is under 1:0.8. The male postabdomen is symmetrical; the epandrium is subeUiptical with closely fused surstyli ventrally and with a median groove. The hypandrial process and aede- 80 GREAT BASIN NATURALIST Vol. 35, No. 1 agus are tusklike, long, and well sclero- tized. The aedeagal apodeme is crescent shaped. Description. — Length 3.4-4 mm; sub- shining to shining, metallic brown to greenish-brown; pollinose, gra^' ventrally. Head. Mesofrons shining, bronze-gold metallic reflections; pectinate branches of arista not more than twice aristal width at base; pruinose face tan; interfoveal hump not as prominent as Paracoenia s. str., dorsally sloping; eye large, subcircu- lar, width in profile double the length of projecting face in profile; eye-to-cheek ratio 1:0.25; width-to-height ratio 1:0.66; height-to-length ratio 1:0.9. Chaetotaxy of head and thorax like Paracoenia s. str. except acrostichal hairs. Thorax. Acrostichal hairs in two rows; dorsum pollinose to subshining; pleural areas concolorous with mesonotum cen- trally, becoming pollinose, grayed mar- ginally; halters yellow. Male mesofemora without comb of bristles. Wings with costal bristles on dorsal and ventral mar- gins; costal vein ratio 1:0.2. Abdomen. Subshining to shining, brown metallic reflections; fifth abdominal ter- gite of male more or less truncate, with- out anteroventral process; fifth abdominal sternite with three posteriorily oriented prongs. Female postabdomen similar to Paracoenia s. str. Male postabdomen as in diagnosis. Fig. 7. Ventral receptacle with operculum wider than high, extending process considerably longer than opercu- lum. Discussion. — Calocoenia is a mono- typic subgenus known only from the No- arctic Region. Nothing is known about the biology of the included species. Although the type-species of Calocoejiia was originally described in Paracoenia s. str., its inclusion with the latter subgenus would form a paraphyletic grouping since the sister group of Calocoenia is Lepto- coenia. The sister-group relationship with Leptocoenia is deduced from the joint pos- Figs. 11-13. — Male genitalia and egg. 11 a,b,c Paracoenia beckeri; 12c Paracoenia fumosa; 13 stereo- scan electron micrograph of Paracoenia calida egg, 100 X. Figures as in caption of 1-5. March 1975 MATHIS: AMERICAN EPHYDRIDAE 81 session of the following apomorphous fea- tures not found in Paracoenia s. str.: eye- to-cheek ratio less than 1:0.3; interfoveal hump not as pronounced as in Paracoenia s. str.; and the fifth abdominal tergite lacking an anteroventral process. Paracoenia (Calocoenia) platypelta (Cresson) Fig. 7, Map 1 Paracoenia platypelta Cresson, 1935, Trans. Amer. Ent. Soc. 61:356 Coenia {Paracoenia) platypelta: Sturtevant and Wheeler, 1954, Trans. Amer. Ent. Soc. 79:165 Types. — Male holotype, Pine Lake, So. Cal., Johnson. The type specimen also has a small label with the male sex sym- bol, Cresson' s pink type label, and a red USNM type label, number 51110. The type is deposited with the U. S. National Museum of Natural History. Cresson's original description also lists a topotypi- cal female paratype. I have examined this latter specimen, presently with the Academy of Natural Sciences of Philadel- phia, and determined it as Paracoenia turbida. Diagnosis and Description. — See generic description. Specimens examined. — 516 Distribution. — This species is known only from the West (US). I have ex- amined specimens from Washington to Alberta (Laggan) and southward to New Mexico, Arizona, and California. I have not seen specimens from Montana or Wyoming, although collection attempts have been made. Remarks. — C. platypelta is a very homogeneous species exhibiting little morphological variation. There is some color polymorphism, but this could repre- sent age polymorphism. As mentioned previously, nothing is known regarding the biology or larvae of this species. Leptocoenia. n. subgen. Type-species. — Coenia paurasoma Sturtevant and Wheeler, monobasic Diagnosis. — Leptocoenia resembles Calocoenia but the body size is much smaller, length 2.1-2.6 mm, and the male postabdomen of Leptocoenia differs con- siderably. The surstyli are well separated apically with a small median triangular process between them, which is very simi- lar to a comparable structure in Para- coenia s. str. rhe triangular process in Paracoenia s. str., however, is better de- veloped in comparison with the lateral surstyli. Externally, Leptocoenia is simi- lar to Calocoenia. and both share the fol- lowing character states: The acrostichal hairs are in two distinct rows; the scutel- Imn is slightly flat; the ventral margin of the fifth abdominal tergite is not pro- duced into a lobe; and the posteroventral surface of the mesofemora does not bear a row of comblike bristles. Description. — Length 2.1-2.6 mm; dark brown, pollinose. Head. Fronto-orbital areas, mesofrons nearly concolorous, the later subshining; pectinate aristal branches at most two and one-half times aristal width at base; in- terfoveal hump not prominent, without pronounced dorsal indentation; pruinose face light tan; longest bristles along ven- tral margin of face approximately three- fourths length of interfoveal himip height; genal bristle weak, subequal to humeral bristles. Eye-to-cheek ratio 1:0.175; width-to-height ratio 1:0.65; height-to- length ratio 1:0.93; eye-width-to-face- length ratio 1:0.3. Thorax. Lightly pollinose dorsally. Acrostichal setae in two rows; four pairs of dorsocentral bristles; humeral bristles present; halters yellowish-brown to brown. Costal setae weak, developed only on dorsal margin. Abdomen. Male postabdomen as in Fig. 9; ventral receptacle as in Fig. 9d. Discussion. — In many respects, this subgenus is pivotal, linking Paracoenia with Coenia. This is evident in characters associated with size and dimension, but the annectant role of Leptocoenia is best evidenced by the shape of the female ven- tral receptacle. The ventral receptacle closely resembles those of Coenia species and is probabl}^ less likely to be affected by selective pressure, which would bring about convergence in dimension and size of external characters except by pleio- trophy. I attribute considerable import- ance to this feature and the relationship with Coenia that it demonstrates. The internal male genitalia of Lepto- coenia also reflect the intermediate posi- tion of this subgenus with Coenia. This is best seen by comparing Figs. 9e and lOe. 82 GREAT BASIN NATURALIST Vol. 35, No. 1 Paracoenia {Leptococnia) pawosoma (Sturtevant and Wheeler) Fig. 9, Map 3 Coenia paurosoma Sturtevant and Wheeler, 1954, Trans. Amer. Ent. Soc. 79:165 Paracoenia paurosoma: Wirth, 1965, USDA Agri. Handbook No. 276, p. 567 Types. — Female holotype, three para- types (one male, two females), Lander, Wyoming, 16 August 1950. The holo- type also bears a collector label, M. R. Wheeler, and type number 6696. A fourth paratype (female). Rainbow Lake, Colo- rado, ^elev. 10,200', 50 hi. The original description indicates that K. W. Cooper was the collector of the fourth paratype, but no collector label accompanied the specimen. The holotype is deposited with the Academy of Natural Sciences of Phil- adelphia, two paratypes with the U.S. National Museum of Natural History, and two paratypes with M. R. Wheeler. Di.\GNOsis AND Description. — See generic description. Specimens examined. — 41 Distribution. — Paurosoma has been collected from Colorado north through Wyoming, Alberta (Laggan), and into Alaska (Matanuska Flats). I have also examined five specimens from Sweden (Norrbotten, 3 km N Messaure). Although paurosoma is Holarctic, it was the most recently discovered species. Moreover, nothing is known about its bi- ology. The habitat of this species is mon- tane; at lower latitudes it is found at ele- vations up to 10,500 feet (Rainbow Lake, Colorado) . Genus Coenia Robineau-Desvoidy Coenia Robineau-Desvoidy, 1830, Essai sur les Myodaires 2:800. Type-species Coenia caricicola Robineau-Desvoidy i = Ephydra palusiris Fal- len), by monotypy; Sturtevant and Wheeler, 1954, Trans. Amer. Ent. Soc. 79:164-166 (re- view of Nearctic species as subgenus of Coenia) ; Wirth. 1965, USDA Agri. Handbook No. 276, pp. 755-756 (catalog) Caenia emendation: Walker, 1853, Insecta Britan- nica Diptera ?:259 (preoccupied-Newman, 1853, Entomological notes. Art X. Ent. mag., pp. 372-402, Coleoptera) Diagnosis. — Coenia species resemble those of Paracoenia^ especially the sub- genus Leptocoenia. but they are differen- tiated as follows: Dorsalmost postocular bristles not subcqual to the verticals; no well-developed humeral bristles; two pairs of dorsocentral bristles; halters brownish- yellow to dark brown; and fifth abdomin- al sternite of males longer than wide. The size of Leptocoenia is within the dimen- sions of Coenia^ and superficially they ap- pear very similar. However, the chaeto- taxy characters readily distinguish either and set the Coenia species apart from any Paracoenia subgenus. Description. — Length 2.2-2.8 mm; dark brown, pollinose to subshining. Head. Mesofrons subshining with me- tallic reflections; postocular bristles weak, at most slightly larger than their distance apart as base; postorbital bristles usually weak; pectinate aristal branches various; bristles of face comparatively large, bristles along ventral margin subequal in length to interfoveal hump height; facial pruinosity various; chaetotaxy of head similar to Paracoenia except as noted. Eye relatively large, subcircular although higher than wide; width-to-height ratio 1:0.62; height-to-length ratio 1:0.95; eye- width-to-face-length ratio 1:0.3. Thoj-ax. Dark brown, generally con- colorous; strength of pollinose covering various. Acrostichal hairs in two rows, in- distinct in some species. Three or four pairs of dorsocentral bristles; humeral bristles various; otherwise chaetotaxy as in Paracoenia. Wings without prominent costal bristles, at most weakly developed on dorsal margin; costal vein ratio 1:0.3; Mi+_. vein ratio 1:0.7. Legs uniformly dark brown; male mesofemora without posteroventral comb. Halters as in diag- nosis. Abdomen. Subshining to shining dark brown; male with five visible tergites; female with six to seven; male with i\\e sternites; fifth sternite narrowed to small strip; spiracle arrangement as in Para- coenia. Male postabdomen with surstvli closely apposed basally, becoming well se])arated on apical 2/3; aedeagal apo- deme broad to narrowly crescent-shaped; aedeagus curved, pointed apically. Ven- tral receptacle with small operculum, ex- tending process C-shaped. Discussion.- — Of the three known Co- enia species, one is aj)parently endemic to the Nearctic Region, a second is limited to the Palearctic, and a third species is Holarctic. Cresson's comments regarding the European species and their distinguish- March 1975 MATHIS: AMERICAN EPHYDRIDAE 83 ing characteristics should bo consulted for tions of various salts. Dahl (1959) and species separation from that fauna. Scheiring and Foote (1973) report on as- Species of Coenia are not usually asso- sociations of Coenia curvicauda (Meigen) elated with aquatic environments that are with mud shore habitats and to a lesser contaminated or contain high concentra- extent with the limnic wrack. Key to Coenia species 1. Male genitalia large, exposed ventrally; epandrium over twice as long as wide, with median suture - C. curvicauda (Meigen) Male genitalia more compact, not exposed; epandrium less than twice as long as wide, without median suture --.. C. alpina n. sp. Coenia alpina, n. sp. Fig. 10, Map 2 Di.'\GNosis. — C. alpina most closely re- sembles C. palustris, a Palearctic species. It is distinguished from the latter by dif- ferences in male genitalic structures. The surstyli are proportionately shorter to the epandrial length, the aedeagus is deeper and Aentrall}' rounded, the aede- agal apodeme is longer, and the hypandri- al processes are thinner and project mesad to the hypandrial connection with the aedeagus. The females of both species are very similar. Description. — Length 2.1-2.6 mm; dark brown with some purplish metallic reflections dorsally. Head. Interfoveal hump more or less prominent; pruinose face brown; pectin- ate branches of arista long, approximately three times aristal width at base; post- orbital setae not developed dorsally. Eye- to-cheek ratio 1:0.12; width-to-height ra- tio 1:0.6; height-to-length ratio 1:0.96; eye-width-to-face-length ratio 1:0.33. Thorax. Acrostichal setae weak, in two rows; three pairs of dorsocentral bristles; humeral bristles absent; halters dark brown. Abdomen. Subshining to shining with some purplish reflections; ventral recep- tacle as in Fig. lOd; male genitaha as in Fig. 10. Distribution. — C. alpina has been collected in Colorado (Rabbit Ears Pass), in the Northwest Territories (Aklavik), and in Labrador (Cartwright). Types. — Male holotype, allotype, and 21 para types (8 males, 13 females). Cart- wright, Labrador, 29 June 1955, E. F. Cashman; 17 paratypes with same data as type except as follows: 5 males, 3 fe- males, 3 July 1955; 4 males, 2 females, 2 July 1955, E. E. Sterns; 1 male, 6 August 1955, E. E. Sterns; 1 male, 12 August 1955. The type, allotype, and 32 para- types will be deposited wdth the Canadian National Collection, type number 13435. A male and female paratype also will be deposited with the California Academy of Sciences, the U.S. National Museum of Natural History, and in my collection. In addition to the type series, I have examined 31 specimens of this species from the following localities: 9 males, 17 females, Aklavik, Northwest Territories, May-August 1930 and 1931; 2 males, 3 females, Rabbit Ears Pass, Colorado, 11 June 1968, S. L. W. Remarks. — This species is alpine, which accounts for the specific name. Al- though the known distribution is based on minimal data that is rather disjunct, I feel that C. alpina is distributed through- out the Rocky Mountains at higher ele- vations and across northern Canada. From my study, I liaAe found very httle variation except for minor size dif- ferences as indicated in the description. Otherwise, C. alpina seems to be a very uniform species. Coenia curvicauda (Meigen) Fig. 8, Map 3 Ephydra curvicauda Meigen, 1830, Syst. Beschr. 6:116 Coenia curvicauda: Macquart, 1835, Hist. Nat. Ins. Dipt. 2:530 Types. — Cresson (1930) designated a lectotype for this species from material in the Naturhistorisches Museum, Wien. According to Cresson, the male lectotype has the following label data: "curvicauda Coll. Winth," "curvicauda." A second male specimen with similar data was de- signated a parat3^pe b}' Cresson. Diagnosis. — C. curvicauda is similar to C. alpina and to C. palustris but differs 84 GREAT BASIN NATURALIST Vol. 35, No. 1 from either by the well-developed male postabdomen which })rotrudes from the venter of the abdomen. The epandrium plus surstjdi are over twice as long as the epandrial width, and the epandrium is divided by a median groove. The sur- styli arms are longer than their base, the aedeagal apodeme is slender and C-shaped in profile, and the aedeagus is broadly developed basally and curves forming a J-shaped structure. Description. — Length 2.3-2.8 mm; dark brown, subshining to shining. Head. Mesofrons shining, dark brown; fronto-orbital areas subshining; pectinate aristal branches subequal to third anten- nal segment width; face lightly pollinose, mostly dark brown; chaetotaxy as in C. alpina; eye-to-cheek ratio 1:0.11; width- to-height ratio 1:0.6; height-to-length ra- tio 1:1; eye-W'idth-to-f ace-length 1:0.3. Thorax and abdomen as in C. alpina except as given in diagnosis. Specimens examined. — 150 Distribution. — Coenia cwvicauda is a Holarctic, boreal species. In the Nearc- tic region it is found in Montana (Big- fork) and east through the Midwest to the Northeast. It extends north to Alaska (Tonsina) and east to Quebec (Cross Point) . Remarks. — Although I did not ex- amine the lectotype, I have studied Euro- pean specimens of this species. The aedeagus in some specimens does not curve apically to the extent found in Ne- arctic specimens, but I did not find any other major differences. I consider all specimens I examined to be conspecific. Literature Cited Aldrich, J. M. 1905. A catalogue of Nortli American Diptera. Smithsn. Inst.. Smithsn. Misc. Collect. 46(2-pub. 14-44) : 1-680. Beyer, A. 1939. Morphologische, Okologische und Ph3-siologisc.hc Studion an den Larven del- Fliegen: Ephydra riparia Fallen. E. mi- cans Haliday und Cacnia jiauosa Stenhani mar. Kieler Meereforschungen 3:265-320. Brock, T. D. .^nd M. L. Brock. 1968. Life in a hot-water basin. Nat. Hist. 77:47-54. Brock, M. L., B. G. Wiegert, and T. D. Brock. 1969. Feeding by Paracoenia and Ephydra (Diptera: Ephydridae) on the microorgan- isms of hot springs. Ecologj- 50:192-200. Coi.i.iN. J. E. 1963. The British species of Ephydra (Dipt., Ephydridae). Entomol. Mon. Mag. 99:147-152. CoQUiLLETT, D. W. 1902. Now acalyptrate Diptera from North America. Jour. N. Y. Ent. Soc. 10:177-191. Cresson, E. T., Jr. 1930. Studies in the Dip- terous Family Ephydridae. III. Trans. Amer. Ent. Soc. 56:93-131. . 1935. Descriptions of genera and spec- ies of the dipterous family Ephydridae. Trans. Amer. Ent. Soc. 61:345-372. . 1942. Synopses of North American Ephydridae (Diptera). I. Trans. Amer. Ent. Soc. 68:101-128. . 1944. Synopses of North American Ephydridae (Diptei-a). la and II. Trans. Amer. Ent. Soc. 70:227-240. . 1946. Synopses of North American Ephydridae (Diptera). III. The tribe Noti- philini of the subfamily Notiphilinae. Trans. Amer. Ent. Soc. 72:227-240. 1949. A systematic annotated arrange- ment of the genera and species of the North American Ephydridae. IV. The subfamily Napaeinae. Trans. Amer. Ent. Soc. 74:225- 260. Curran, C. H. 1927. Descriptions of nearctic Diptera. Canad. Ent. 59:79-92. D.\HL, R. G. 1959. Studies on Scandinavian Ephydridae (Diptera. Brachycera). Opusc. Ent.^ (Soc. Ent. Lund.). Sup. 25:1-225. H.\LiD.\Y, A. H. 1839. Remarks on the generic distribution of the British Hydromyzidae (Diptera). Ann. Nat. Hist. 3:217-224. Johnson. C. W. 1925. Fauna of New England. List of the Diptera or two-winged flies. Occas. Papers Boston Soc. Nat. Hist. 7(15) :1- 326. LoEW, H. 1860. Die Europaeischen Ephydrini- dae und die Bisher in Schlesien Beobachteten Arten Derselben. Neue Beitrage zur Kennt- nis der Diptereu. part 7. Mittler und Sohn, Berlin. M.\CQU.\RT, J. 1835. Histoire Naturelle des In- sectes Dipteres. vol. 2. Roret. Paris. Meigen, J. W. 1830. iSj-stematische Beschrei- bung der bekannten europaischen zweiflugel- igen Insekten. vol. 5. Hamm. Mitchell, R. M. ,\nd B. L. Redmond. 1974. Fine structure and respiration of the eggs of two ephydrid flies (Diptera: Ephvdridae). Trans. Amer. Micros. Soc. 93:113-118. Newm.\n, E. 1838. Entomological notes. Art. XL. Ent. Mag., pp. 372-402. Oliveir.\, S. J. 1954a. Contribuicao para o con- hecimento do genero ''Dimecoenia" Cresson. 1916. I. ''Dimecoenia lenii" sp. n. encontrada no Chile (Diptera, Ephydridae). Rev. Brasil. Biol. 14:187-194. . 1954b. Contribuigao para o conheci- mento do genero ''Dimecoenia'" Cresson, 1916, II. Sobre 3 especies novas do Brasil (Diptera. Ephydridae). Rev. Brasil. Biol. 14:269-278. . 1957. Contribuigao para o conhecimen- to do genero ''Dimecoenia' Cresson, 1916. III. Sobre uma especie nova do Estado de Sao Paulo. Brasil (Diptera. Ephydridae). Rev. Brasil. Biol. 17:305-308. PoLHEMUs, J. T. 1967. A new saldid from California (Hemiptera: Saldidae). Proc. Ent. Soc. Wash. 69:346-348. Rorineau-De-svoidy. a. J. "B. 1830. Essai sur les Myodaires. Memoires da I'Academie Rovale des Sciences de ITnstitut de France. 2:1-813. March 1975 MATHIS: AMERICAN EPHYDRIDAF. 85 ScHEiRiNG. J. F. AND B. A. FooTE. 1973. Hab- itat distribution of the shore flies of north- eastern Ohio (Diptear: Ephydridao). Ohio Jour. Sri. 7^:152-166. Stenh.\mm.\r. C. 18-H, Forsok till gruppering och revision af de svenska Ephydrinae. K. Vetensk. Akad. Handl. 1843:75-272. Steyskal, George C. 1970. The species of the genus Dimecoenia (Diptera: Ephydiidae) in America north of Panama, with the descrip- tion of a new species. Ann. Ent. Soc. Amer. 63:462-465. SxURTEyANT, A. H. AND M. R. WlIEEI.ER. 1954. Synopses of nearctic Ephydridae (Diptera). Trans. Amer. Ent. Soc. 79:151-261. Walker, F. 1849. List of the specimens of dip- terous insects in the collection of the British Museum. Vol. 4. pp. 689-1172. . 1853. Insecta Britannica. Diptera. Vol. II, 297 pp. WiEGERT, R. G. and R. Mitciiei.e. 1973. Ecol- ogy of Yellowstone thermal effluent systems: intersects of blue-green algae, grazing flies (Paiaroenia, Ephydridae) and water mites {Part/iuniclla, Hydrachnellae). Hydrobiol- ogia M: 25 1-271. WiRTir, W. W. 1948. A taxonomic study of Hawaiian Ephydridae (Diptera) related to Scalclla Robineau-Desvoidy. Proc. Hawaiian Ent. Soc. 13:277-304. — -. 1965. Ephydridae. Pages 734-759 in Alan Stone, C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson, eds. A catalog of the diptera of America north of Mexico. U. S. Dep. Agr. Handb. No. 276. . 1970. A new genus and species of shore fly (Diptera. Ephydridao) from southern Pata- gonia. Act. Zool. Lill. 26:1-8. . 1971. The brine flies of the genus Ephydra in NoiUi America (Diptera: Eph- ydridae). Ann. Ent. Soc. Amer. 64:357-377. ENVIRONMENTAL FACTORS IN RELATION TO THE SALT CONTENT OF SALICORNIA PACIFIC A VAR. UTAHENSIS' D. J. Hansen- and D. J. Weber ABSTII.A.CT. — The stability of the salt content in Salicornia pacifica Standi, var. utahensis (Tide- strom) Munz in relation to environmental changes was investigated. Salicornia pacifica communities have a characteristic soil pH of 7.5 to 8.0 ± 0.2 and a constant subsurface soil moisture level of 25 to 35 percent. The ion content in the tissue of S. pacifica remained constant despite increased moisture stress throughout the growing season. The concentrations of the salts were significantly higher in the surface soil layers than in the subsurface layers around the roots. Normal metabolic processes in the tissues of S. pacifica appear to occur even though some fluctuations in the ionic balance and concen- tration of ions in the plant occur. Introduction Halophytic plants are among the few species of higher plants that can with- stand high sahne soil conditions without detrimental effects (Waisel, 1972). The is mainly associated with an increase in the chloride content of the tissues (Adri- ani, 1958; Bernstein, 1961). Steiner (1935) found that chloride ions accounted for 67 to 88 percent of the increase of osmotic potential of different species of salt marsh plants, whereas other osmotic- ally active substances had only a negligi- ble effect. Chloride accounted for 80 per- cent of the total osmotic potential in 5^//- cornia ambigua, 91 percent in S. stricta, and 93 percent in S. mucronata (Arnold, 1955). Harward and McNulty (1965), on the other hand, found that chloride accounted for less than 49 percent of the osmotic potential in S. ?ubra. Osmotic values for leaf saps of a num- ber of herbaceous halophytes including S. herbacea, ranged from 25 to 75 atm when grown in salt marshes (Yabe et al., 1956). Seasonal changes in osmotic potentials in tissues of 5". rubra ranged from 40 atm to over 100 atm over a two-month period (Harward and McNulty, 1965). Higher osmotic potential values have been record- ed for a number of halophytes; for ex- ample, Waisel (1972) reported that os- motic potentials of Rhizophora and Ain- cennia leaves reached values of 148 and 163 atm, respectively. According to Bowen and Rovira (1966), salt ions can cause toxicity in the follow- ing ways: (1) acting as antimetabolites, (2) binding or precipitating various me- tabolites, (3) catalyzing rapid decomposi- tion of essential elements, (4) combining with cell membranes and affecting their permeability, and (5) displacing essential elements but failing to fulfill their func- tions. Bowen and Rovira (1966) sug- gested that salt injury is not due to a direct effect of the salts but to the indirect effects of one or more of the above men- tioned metabolic disturbances. Waisel (1972) singled out nitrogen metabolism as an important area affected by high salts. Salt-induced growth retardation leads to an accumulation of unused substances that may be toxic (Gauch and Eaton, 1942). The degree of salt injury or tolerance of plants may be affected by a nmnber of environmental factors such as water- logged soils. Some plants, however, have adapted to waterlogged conditions. Sali- cornia foliosa not only tolerates water- logged conditions but appears to benefit from them, because of increased capacity to obtain iron under these conditions thereby avoiding chlorosis (Adams, 1963). In habitats with marked fluctuations in salt concentration, only the species with high osmotic shock resistance {Salicornia sp.) can survive (Levitt, 1972). Some of the facultative halophytes, such as Sali- cornia rubra, are found at the highest salinities 3^et are capable of growing nor- mally in low to nonsaline environments (Ungar et al., 1969). Since species of Salicornia are among the most salt tolerant forms of higher plants (Chapman, 1960), S. pacifica Standi, var. utahensis (Tidestrom) Munz., a halophyte conunon to inland salt playas of northern Utah, was selected for this investigation. Due to the lack of basic in- formation about environmental fluctua- 'This research was supported in part by NSF Grant No. GB3I0G7 and a -Department of Botany, Brighani Young University, Prove, Utah 84602. It from Brigham Young University. 86 March 1975 HANSEN, WEBER: SALICORNIA 87 tions in S. pacifica habitats and difficulties in providing an artificial environment that parallels the natural environment, investigations were undertaken to corre- late fluctuations of the natural environ- ment with physiological and morphologi- cal characteristics in relation to salt con- tent of 5'. pacifica. Methods and Materials This investigation was conducted dur- mg a typical growing season for Salicor- nia pacifica (April to August 1972) . Three sites were selected to compare soil and moisture factors in relation to changes in salt content of S. pacifica. Sites 1 and 2 were six miles (9 km) north and Site 3 was about one-half mile (1 km) east of Goshen, Utah. Site 1 was especially se- lected because it was an ecotone between a stand of S. pacifica and a stand of Dis- tichlis stricta. It was hoped that the data from this site could be used to explain some of the environmental factors respon- sible for separating the Salicornia com- munities from the Distichlis communities. The ground at Site 1 was covered by a thin layer of dried algae of the genus, Oscillatoria. This covering formed a sur- face mulch which increased the moisture of the soil surface. Site 2 was about 100 m west of Site 1 on the opposite side of a large drainage basin. Site 3 (Fig. 1 ) was 6 miles (9 km) from Sites 1 and 2 adja- cent to a natural drainage system. Site 3 was selected because of reduced fluctua- tions in soil moisture throughout the grow- ing season. Soil and plant samples were taken every two weeks at all three sites. A standardized hygrothermograph unit was used to continuously monitor temper- ature and relative humidity. The housing unit for the hygrothermograph was loca- ted three inches above the ground about 100 m from Sites 1 and 2. Measurements were recorded from 1 May through 20 September 1972. The percentage of plant cover for Sites 1, 2, and 3 was determined using eight randomly distributed rectangular 1/4 m- quadrats. Dry weight production in each site was determined by clipping four ran- domly distributed rectangular %. m- quad- rats. Soil samples were taken with a soil core borer. Each core measured 1 inch (2:54 cm) in diameter and was extended to a depth of 10 inches (25.4 cm). The cores were separated to provide surface (upper Fig. 1. Site 3. located one mile east of Goshen, Utah, sliowi pacifica (Photograph courtesy of W. M. Hess.) (! (if Salicornia 88 GREAT BASIN NATURALIST Vol. 35, No. 1 2 inches or 5 cm of the core) and subsur- face (lower 6-10 inches or 15-25 cm of the core) soil samples. Five to eight core samples were taken at each site and pooled to obtain the soil sample. The samples were placed in plastic bags, sealed, and immediately taken to the laboratory for analysis. Samples were weighed to the nearest one-hundredth of a gram and dried in an oven at HOC for 48 hours. The samples were weighed again, and the percentage of moisture was calculated. Measurements of the soil pH were de- termined from saturated soil paste sam- ples using the Sargent- Welch pH Meter, Model PAX, with a combination electrode. Osmotic potentials of soil samples were determined from saturated soil pastes by freezing-point depression methods accord- ing to the procedure outlined by Hansen and Weber (1974). Soil samples (5 g) were leached of ex- changeable cations and anions by flush- ing with four 25 ml volumes of IN CH3- C-NHo, pH 7.0. Each volume was allowed to drain before the next was applied. The cations (sodium, potassium, magnesium, and calcium) were detected in the filtrate according to procedures outlined bv Per- kin-Elmer (1971) on the Modef 290B Atomic Absorption Spectrophotometer. The anion, chloride, was detected in the filtrate according to the procedure outlined by Marius/Fiske (1972) using a Mari- us/Fiske Chlor-o-counter. Osmotic potential measurements for Salicornia were determined by freezing- point depression techniques as described by Gary and Fisher (1969, 1971) and Fisher (1972). The circuitry was modi- fied by replacing the two 1.35 vdc Hg bat- tery cells with an alkaline 9 vdc battery. A lOK 1-tum potentiometer was installed to compensate for voltage drop that oc- T.ABLE 1. Percentage moisture, dry weight production, and percentage cover of Salicornia pacifica in three salt desert playas. Dry weight Percentage production Percentage Site moisture g/m' cover 1 79.0 73.1 56.0 S. pacifica 2.5 S. rubra 80.6 26.7 7.6 S. pacifica 1.0/1. occidentalis 6 80.6 141.2 65.0 S. pacifica 2.5 S. rubra curred with time and usage. Measure- ments were made at each internode for several plants. Measurements were occa- sionally made using a vapor pressure os- mometer. Model 301 Mechrolab Inc., to verify freezing-point measurement values. The percentage of crude protein in each plant sample was determined by the micro- Kjeldahl method (Horwitz, 1970). Results Climatic and Growth Factors Continuous hygrothermograph monitor- ing of climatic factors indicated that tem- peratures fluctuated consistently through- out the growing season. The average temperatures gradually increased through May and June. A peak was reached dur- ing July, followed by a gradual decrease from August to September. The average of the daily highs during July was 34C. The hottest temperature in July was 38G on 12 July. The coolest temperature re- corded in July was 6G on 25 July. Lowest daily means (calculated on an hourly basis) for relative humidity oc- curred from 15 July through 20 August. A series of rain storms increased the rela- tive humidity during late August and early September. Light rain was also common diu-ing late May and early June. No quantitative data concerning the amount of rain was taken. Relative hu- midity reached 100 percent every night except for about six days during the grow- ing season. The dry weight of the standing crop of S. pacifica plant tissue was the highest on Site 3 (141.2 g/m-) and the lowest on Site 2 (26.7 g/m-) as shown in Table 1. The percentage cover was also highest on Site 3 (65 percent) and lowest on Site 2 (7.5 percent). The percentage moisture of the plants in all three sites was about 80 percent. Soil Moisture The subsurface and surface soil mois- ture readings for Site 2 were highest dur- ing June, gradually decreasing through- out the growing season (Fig. 2). The sub- surface soil moisture for this site was less than the surface soil moisture from 1 April through 15 July. This may have been due to the heavy mulch of algae on the surface. However, from mid-July through the rest of the season, this trend March 1975 HANSEN, WEBER: SALICORNIA 89 Fig. 2. Soil moisture for three salt desert playas (Sites 1, 2, and 3). Solid lines indicate the plot of percentage values for surface soil sam- ples (upper 2"). Dashed lines indicate the plot of percentage values for subsurface soil samples (6-10"). was reversed. In Sites 2 and 3 the subsur- face soil moisture was consistently higher than the surface soil moisture throughout the growing season, except for one week in June when Site 2 had an increase in the surface soil moisture due to rainfall. Sur- face and subsurface soil moisture percent- ages at Site 3 were well above moisture percentages of Sites 1 and 2. From June through July soil moisture in Site 3 was above 30 percent, whereas soil moisture of Sites 1 and 2 was well below 30 per- cent. This was probably due to under- ground seepage from the nearby drainage system adjacent to Site 3. Soil moisture was lowest in all three sites during May. Soil pH The surface soil pH of the three sites was generally higher and fluctuated more than the corresponding subsurface pH (Fig. 3). The subsurface pH of these sites changed very little during the grow- ing season. The decrease in the pH of the soil surface appeared to be correlated to the amount of rainfall. This effect was probably due to the transporting of solu- ble salts into the subsurface layers by the percolating rain. A decrease in the pH of the surface generally was inversely proportional to the increase in the subsur- face pH. Ion Content of the Soil Site 1 Osmotic potential measurements of the surface soil were high during Jul}' and September. Osmotic potential values were Fig. 3. Soil pH as recorded for three salt desert playas (Sites 1, 2, and 3). The solid lines indicate the plot of tlie pH for surface samples (upper 2"). The dashed lines indicate the plot of the pH for subsurface samples (6-10"). as high as 135 atm (Fig. 4). Osmotic potential measurements of subsurface soil samples were considerably lower than os- motic potential measurements of the siu-- face. Osmotic potential values for the sub- surface soil samples gradually increased. The highest value reached was 48 atm. This value was recorded on 20 September, when the study terminated. Values dur- ing the hottest month did not exceed 23 atm. Individual ion analysis of soil samples from the surface (Fig. 5) and the subsur- face layers (Fig. 6) showed that sodium and chloride were the two ions responsi- ble for most of the osmotic potential. The increasing ion accumulation of the soil closely paralleled the increase in osmotic potential previously described. The con- 's— ik — to — it «»"• LIST I UPTiytl* Fig. 4. Osmotic potential of saturated soil paste for a salt desert playa (Site 1) expressed as atmospheres pressure. The solid line indicates the plot of osmotic potential values for the sur- face soil samples (upper 2"). The dashed line indicates the plot of osmotic potential values for tlie subsurface samples (6-10"). 90 GREAT BASIN NATURALIST Vol. 35, No. 1 Fig. 5. Ion content of the soil for the surface layer (upper 2") of a desert playa (Site 1) ex- pressed as a percentage of the dry soil. The solid line indicates the percentage of chloride. The dashed line indicates the percentage of sodium. The dash-dotted line indicates the percentage of calcium. The solid vertical-barred line indicates the percentage of potassium. centratioiis of sodimn and chloride were highest during July and September. The concentration levels of calcium and potas- sium ions were fairly constant through- out the season. Concentration values sel- dom exceeded 0.5 percent of the dry soil weight in either the surface or the subsur- face layers. Site 2 Osmotic potential values for the surface layer samples were uniform and high (over 130 atm) throughout the season with the exception of a substantial de- crease on 29 August, when values dropped to 80 atm (Fig. 7). The subsurface values steadily increased throughout the growing season to a high of about 96 atm on 20 September. In most cases osmotic poten- tial values of the subsurface soil samples were 50 to 100 atm lower than osmotic potential values of the surface soil sam- ples. Thus, roots near the surface would SUBSURFACe LAYEn Fig. 6. Ion content of the soil for the sub- surface layer (6-10") of a salt desert playa (Site 1) expressed as a percentage of the dry soil. The solid line indicates the percentage of chloride. The dashed line indicates the percentage of sodium. The dash-dotted line indicates the percentage of calcium. The solid vertical-barred line indicates the percentage of potassium. V) o 0 VZ 4T"6 k I lb 12 114 16 APRIL* MAY ' JUNE ' JULY ' AUO OSMOTIC POTENTIAL OF SOIL PASTE Fig. 7. Osmotic potential of satui-ated soil paste for a salt desert playa (Site 2) expressed as atmospheres pressure. The solid line indicates the plot of osmotic potential values for the surface soil samples (upper 2"). The dashed line indi- cates the plot of osmotic potential values for the subsurface soil samples (6-10"). be in an environment of higher moisture stress than deeper roots. Ion analysis of the surface layer soil samples showed that sodium and chloride ions were responsible for most of the os- motic potential of the soil samples (Fig. 8). The calcium ion concentration of the surface layer for this site was consider- ably higher than for Site 1. The subsur- face calcium and potassium ion concen- trations were comparable with other sites and rarel}^ exceeded 0.5 percent of the soil dry weight (Fig. 9). The combined ion concentration in the subsurface layer increased gradually over the growing sea- son and reached a peak of about 6 per- cent soluble salts on 20 September. Site 3 Osmotic potential measiu-ements of soil samples on this site gradually increased in both the surface and subsurface layers. Measurements were highest on 20 July and 20 September (Fig. 10) and were comparable with measurements from Site 2, both of which were considerably higher than Site 1 . Chloride ion content from the surface layer fluctuated considerably, but the so- dium content was more stable (Fig. 11). The major increase in osmotic potential of the subsurface layer from 20 July through 20 September was due chiefly to the chloride ion content. The calcium ion concentration level of "this site was con- siderably higher than Site 1 and not as high as Site 2. The potassiimi ion con- centration level was less than 0.5 percent March 1975 HANSEN, WEBER: SALICORNIA 91 ~!2 WEEKS SEPTEHBEK Fig. 5. Ion content of the soil for the surface layer (upper 2") of a salt desert playa (Site 2) expressed as a percentage of the dry soil. The solid line indicates the percentage of chloride. The dashed line indicates the percentage of sodi- um. The dash-dotted line indicates the percentage of calcium. The solid vertical-ban-ed line indicates the percentage of potassium. APRILI U*r I JUNE I JULY I AUGUST I SEP Fig. 9. Ion content of tlie soil for the sub- surface layer (6-10") of a salt desert playa (Site 2) expressed as a percentage of the dry soil. The solid line indicates the percentage of chloride. Th dashed line indicates the percentage of sodium. The dash-dotted line indicates the percentage of calcium. The solid vertical-barred line indicates the percentage of potassium. and did not fluctuate significantly. The concentration level gradually increased to a peak of 0.5 percent on 20 July and then gradually decreased to 0.3 percent on 20 September. Concentrations of so- dium and chloride ions in the subsurface soil samples gradually increased (Fig. 12). Calcium and potassium ion concentra- tions in this layer remained low and rela- tively constant through the growing sea- son; these ion concentrations were com- parable to ion concentrations of the other sites. Ion Content in Salicornia Site 1 The chloride ion content in Salicornia tissues remained constant throughout most of the growing season but increased slightly in April (Fig. 13). This increase was followed by a stabilization of the con- centration at about 12 percent of the dry weight. Sodimn ion concentration gradu- ally increased from about 4.2 to 9 percent, while potassium ion concentration de- creased throughout the season from 2.2 to 1 percent. Site 2 The chloride content gradually in- creased from 14.2 to 16.1 percent at the end of the growing season (Fig. 14). In- crease in the sodium content was ]:)ropor- tional to the increase in the chloride con- tent and was 10.2 percent at the end of the growing season. The potassium ion concentration increased from 4.5 to 7.5 percent and was closely correlated to de- creases in the sodium ion concentration. n — • I 'o ,,« |i — ^ — f I JUNE I JUtT I AUOUST I ~iz WEEKS Fig. 10. Osmotic potential of saturated soil paste for a salt desert playa (Site 3) expressed as atmospheres pressure. The solid line indicates the plot of osmotic potential values for the surface 5oil samples (upper 2"). The dashed line indi- cates the plot of osmotic potential values for the subsurface samples (6-10"). SEPTEHSEI) Fig. 1 1 . Ion content of the soil for the sur- face layer (upper 2") of a salt desert playa (Site 3) expressed as a percentage of the dry soil. The solid line indicates the percentage of chloride. The dashed line indicates the percentage of so- dium. The dash-dotted line indicates the percent- age of calcium. The solid vertical-barred line indicates the percentage of potassium. 92 GREAT BASIN NATURALIST Vol. 35, No. 1 Fig. 12. Ion content of the soil for the sub- surface layer of a salt desert playa (Site 3) ex- pressed as a percentage of the diy soil. The solid line indicates the percentage of chloride. The dashed line indicates the percentage of sodium. The dash-dotted line indicates the percentage of calcium. The solid vertical-barred line indicates the percentage potassium. irr Fig. 13. Ion content in Salicornia pacifica (Site 1) expressed as a percentage of the dry weight. The solid line indicates tlie plot of the percentage values for chloride ions. The dash- dotted line indicates the plot of the percentage values for sodium ions. The dashed line indicates the plot of the percentage values for potassium Site 3 The ion content in Salicornia tissues in this site showed a decrease in chloride, so- dium, and potassium throughout the sea- son (Fig. 15). The chloride content ranged from 16 percent on 1 May to 12.4 percent on 20 September. The sodium content ranged from 1 1 percent on 1 May to 7.6 percent or 20 September. The po- tassium content ranged from 4.8 percent on 1 May to 2.1 percent on 20 September. Osmotic Potentials in Salicornia Early in the investigation it was dis- covered that each internode of the plant was osmotically different from other in- temodes of the same plant. Freezing- point depression measurements showed that osmotic potential values increased from the base of the aerial shoot upward to the top of the plant (Table 2). The lowest intemodes near the base generally had a lower osmotic potential than inter- nodes near the middle of the plant. In some cases there was as much as 15 atm difference between two regions (inner and outer) of the cortex, although usually the difference was only about 5 atm (Table 2). Because of the complexity of such differences in osmotic potential read- ings, only periodic measurements were made on the plants. The measurements ranged from a low of about 80 atm in May to about 130 atm to 150 atm in mid- July and August. However, the average was about 90 to 100 atm. Crude Protein Analysis Crude protein analysis showed a gradu- al decrease in the total crude protein con- tent in the plants from all three sites from 1 April through 30 July. Crude protein decreased from 20 to 8 percent on a dry weight basis. From 30 July through 20 September the content remained about 8 percent. Phenology and Morphology Growth of 5". pacifica usually begins with development of the subterranean or ION CONTENT IN SALICORNIA .... -iSf Fig. 14. Ion content in Salicornia pacifica (Site 2) expressed as a percentage of the dry weight. The solid line indicates the plot of the percentage values for chloride ions. The dash- dotted line indicates the plot of the percentage values for sodium ions. The dashed line indi- cates the plot of the percentage values for po- tassium ions. IST Ho n wtEW Fig. 15. Ion content in Salicornia pacifica (Site 3) expressed as a percentage of the dry weight. The solid line indicates the plot of the percentage values for chforide ions. The dash- dotted line indicates the plot of the percentage values for sodium ions. The dashed line indicates the plot of the percentage values for potassium March 1975 HANSEN, WEBER: SALICORNIA 93 Table 2. Osmotic potentials of four different 5. pacifica plants showing the values (atm) of the outer and inner "cortex" tissues from the bottom of the shoot (Node 1) to the top of the shoot (Nodle 5). Node Plant A Inner Outer Plant B Inner Outer Plant C Inner Outer Plant D Inner Outer 1 76 76 81 81 73 73 84 89 2 69 89 57 75 61 80 64 74 3 71 86 93 86 83 96 71 74 4 67 67 96 106 68 83 73 79 5 106 130 96 106 103 108 84 88 6 89 92 93 108 7 106 112 near-subterranean axillary buds of older shoots that start to grow in March and emerge in late April or early May. Flow- ering occurs in mid-June and is generally completed in July. By late August lower internodes begin to wither and die sequentially from the base of the shoot to the tip (Fig. 16). Oc- casionally internodes may become injured or for other reasons may wither and die. The central stele continues to function in a normal manner. Seed-producing inter- nodes are the last to die (Fig. 16). It is not known whether the internodes die strictly by senescence, whether accumu- lation of additional salts in these areas causes death, or whether death is related to some other phenomenon. The seeds are shed in October and No- vember. A few seeds adhere to the mother plant until rain or heavy snows separate them. Seeds are protected by a bulky, lightweight seed coat that aids in dis- persal and absorption of moisture. Seasonal Variations of the Environment Temperatures were highest during July, while relative humidity was lowest dur- ing July and August. These high tem- peratures would cause increased moisture stress. The toxicity of salts increases with the temperature, according to Kaho (1926) and Waisel (1972). The high moisture content of the soil and the subsequent evaporation may have had a cooling ef- fect upon the temperature of the soil. Both lower temperatures and high rela- tive humidity would have a favorable effect upon plant survival. At all three sites there was an increase in moisture stress throughout the season primarily due to increases in salt from underground sources and slight decreases in soil moisture. Soil moisture of 25 to 35 percent is perhaps an important fac- tor in maintaining S. pacifica in the en- vironment. Sodium and chloride were the two prin- cipal ions responsible for increases in os- motic potential of soil samples. They were also responsible for 85 to 95 percent of the osmotic potential of S. pacifica. Har- ris (1915) considered NaCl the most toxic of several soluble salts but concluded that salt mixtures were not as toxic in soils as in culture solutions. The salt concentration of the soil sur- face layers was considerably higher than the salt concentration in the rooting lay- ers. Soil surface layers were often en- crusted with deposits of white salt, which gave the impression that the plants were surrounded by extremely high concentra- tions of salt when, in fact, the rooting zones or layers were only moderately sa- line. This observation was also reported by Wiesel (1972). The wicking action caused by evaporation of water resulted in salt crystallization at the surface, which removed salts from the rooting layers and provided a unique environmental niche for S. pacifica to occupy. Succulents such as Salicornia appear to lose the function of portions of their fleshy cortex and leaves (Fig. 16), yet the vascular system continues to function for the upper fleshy tissue (Fig. 16). The thick cuticle and waxy layer of the plants and the active phellogen of the central stele appear to protect the upper shoots from the more severe environmental stresses. The soil pH was relatively stable throughout the growing season in all three sites. The surface soil pH of each site was about one-half pH unit higher than the subsurface pH. Decreases in the pH of the surface layers were proportional to increases in pH of the subsurface layer. Conversely, increases in the pH on the surface layer were proportional to de- 94 GREAT BASIN NATURALIST Vol. 35, No. 1 Fig. 16. Shoots of Salicornia pacifica showing healthy fleshy portions of stem above dead fleshy sections of cortex. creases in the pH of the subsurface layer. Calcium ion concentrations in the surface layers were considerably higher at Sites 2 and 3 than at Site 1. Soil pH at Sites 2 and 3 was significantly lower than at Site 1. The pH may have been indirectly affected by the calcium concentration or may haAe been related to the algal cover- ing of the soil surface in Site 1. Ecological Interactions and Plant Distribution There appear to be two groups of thought explaining the restriction of Sali- cornia to a saline environment. New- wohner (1938) stated that halophytes fail to succeed in fresh-water habitats because of competition with other species. This idea was supported by Stalter and Batson (1969), who stated that survival and growth rate data of transplanted salt marsh vegetation suggest that several species of halophytes (S. virginica includ- ed) can tolerate conditions not found in their usual zones. Montfort et al. (1927), on the other hand, claimed that this ex- planation of plant distribution must be displaced by the concept of "direct ecolog- ical salt action." He pointed out that S. herbacea has an optimum growth between 1.5 and 3 percent salt. Webb (1966) al- so pointed out that Salicornia plants grown without salt soon die. Waisel (1972) stated that the proportion of chlorides to sulfates and the total salt content were important in determining distribution. For example, S. herbacea was more sensitive to a high proportion of sulfates than was Aster tripolium. The fact that Site 1 is an ecotone be- tween S. pacifica and Distichlis stricta suggests that differences in this site com- pared with the other two sites would provide information regarding some par- ameters of S. pacifica. Distichlis stricta in this site appears to tolerate a higher pH than S. pacifica and may survive in pH values from 8.0 to 9.0. The soil pH apparently affects the availability of nu- trients. At pH values over 7 the avail- ability of manganese, iron, copper, molyb- denum, and zinc declines (Buckman and Brady, 1969). At high pH values phos- phorus forms insoluble" complexes of cal- cium that are unavailable to the plants (Buckman and Brady, 1969). Levitt (1972) also pointed out that Allenrolfea March 1975 HANSEN, WEBER: SALICORNIA 95 occidentalis and S. suhtcrminalis are highly resistant to salt but sensitive to al- kalinity. A change from saline to sodic soils may kill these plants. Site 3, where S. pacifica cover and production were the highest, had an average pH of 7.7 and a moisture content of 32 percent. At Site 1 reduction in the soil moisture and the in- crease in the total concentration of salts of the subsurface soil layers during the season apparently also favored growth of D. stricta over S. pacifica. Distichlis stric- ta appears to be a better competitor for low soil moisture than S. pacifica. It seems probable that during the initial adaptation of Salicornia to the saline en- vironment competition played a major role in determining plant distribution. As natural selection of Salicornia occurred, adaptations allowing the plant to maintain itself in an optimal salt concentration were reinforced. Plasticity could be re- duced and that portion of the gene pool lost that allowed the plant to move back into non-saline areas. As such selective forces would be localized, it would be im- proper to generalize and say that all spe- cies of Salicornia have optimiun growth at high salt concentration. The variations in salt concentrations that yield optimum growth of Salicornia would bear this out (Halket, 1915; Webb, 1966; Levitt, 1972; Waisel, 1972). The Nature of the Fleshy Stem Anatomical investigations with light microscopy indicate that as internodes mature or become injured a phellogen that produces fibers and suberized cells is formed in the central stelar region (un- published results). These cells seal off the conductive tissues from water and nutritive loss as the internodes wither and die. This development would protect the plant when the salt and moisture stresses are the greatest near the surface in July and August. Osmotic potential measurements of the plants indicated that each internode is probably osmotically independent from other internodes. This would allow the plant to survive in the environment if a few of the internodes were attacked and destroyed by insects or disease. Hill (1908) showed that osmotic adaptation differs not only between species but also between organs and even between cells. In his investigation root hairs varied greatly in this respect, even in the same individual plant. Measurements of freez- ing-point depression of S. pacifica inter- nodes indicated that at least two areas of osmotically different tissues were present within one internode, corresponding to the inner "cortex" and outer chloren- chyma tissues. Osmometer measurements on dissected portions of these tissues indi- cated that the chlorenchyma tissue had a higher osmotic potential. Scholander et al. (1966) suggested that xylem sap of S. pacifica may contain very little salt as freezing-point depression of S. pacifica sap was found to approach freezing-point depression of pure water. Since chloride ions were equally distributed in these two tissues (unpublished data), the difference was probably due to sodium ions or sug- ars and other organic molecules that were produced by chloroplasts in the immediate vicinity. Steiner (1935) has shown that chloride ions account for 67 to 88 percent of the increase in osmotic potential of dif- ferent species of salt marsh plants, where- as other osmotically active substances had only negligible effects. This increase in chloride was shown to account for in- creases in the osmotic potential of the en- tire plant. This does not rule out the pos- sibility that differences in osmotic poten- tials of individual tissues might be due to other osmotically active substances. Increase in the chloride and sodiimi con- tent of the succulent tissues appears to be the primary means of osmotic adapta- tion whereby the plant can survive under increased salt and moisture stress. Sodium and chloride ions account for 85 to 95 percent of the osmotic potential of S. pacifica. These results agree with the findings of Harward and McNult}' (1965) and Scholander et al. (1966). Ion analysis of S. Pacifica tissues in Site 1 indicated that increases in osmotic potential were due to increases in Na+ but not CI". In- creases in osmotic potential at Site 2 were due to both Na+ and C1-. However, at Site 3 there was a decrease in both Na"^ and CI". As the concentration of the soil salts increased, causing an increase in moisture stress, osmotic potential of the plant also increased. This increase was due chiefly to ions other than Na+ and CI" or sugars and other organic molecules. Ion analysis of S. pacifica tissues showed that the chloride ion was present 96 GREAT BASIN NATURALIST Vol. 35, No. 1 in higher concentrations than sodium or potassiiim ions. Azizbekova and Babaeva (1970) found that the amount of absorbed Na+, CI", and Mg"^"" in Salicornia increased with increasing salt concentrations. During the period of greatest moisture stress, from July through September, the percentage of crude protein in S. pacifica for all three sites remained constant at about 8 percent of the dry weight. The percentage ion content of sodium, potas- sium, and chloride also remained relative- ly constant throughout the growing sea- son, even when environmental stresses fluctuated considerably. The metabolism of S. pacifica func- tioned even with changes in the ionic balance and concentration of ions in the plant. The changes, however, were gradu- al. References Adams, D. A. 1963. Factors influencing vas- cular plant zonation in North Carolina salt marshes. Ecology 44:^145-456. Adriani, M. J. 1958. Halophyten. Encycl. Plant Physiol. 4:709-736. Arnold, A. 1955. Die Bedeutung der Chlori- onen fiir die Pflanze. Guxtav Fischer. Jean. Bernstein, L. 1961. Osmotic adjustment of plants to saline media. I Steady state. Amer. J. Bot. 48:909-918. BowEN, G. D., AND A. D. RoviRA. 1966. Mi- crobial factor in short-term phosphate uptake studies with plant roots. Nature 211:665-666. BucKMAN O.. AND C. Brady. 1969. The nature and properties of soils. Macmillan Company, London. C.-\RY, J. W., AND H. D. Fischer. 1969. Plant moisture stress: a portable freezing-point meter compared with the psychrometer. Agron. J. 61:302-304. . 1971. Plant water potential gradients measured in the field by freezing point. Physiol. Plant. 24:396-401. Chapman, V. J. 1960. Salt marshes and salt deserts of the world. Lenard Hill Books Ltd.. London. Fisher. H. D. 1972. An ine.xpensive meUiod of determining plant moisture stress using freez- ing-point depression. Soil Sci. 113: 383-385. Gauch, H. G., AND F. M. E.ATON. 1942. Effect of saline substrate on hourl}' levels of carbo- hydrates and inorganic constituents of barley plant. Plant Physiol. 17:422-434. Halket, a. C. 1915. The effect of salt on the growth of Salicornia Ann. Bot. 29:143-154. Hansen, D. J., and D. J. Weber. 1974. Ap- plication of freezing point depression for the detemiination of osmotic potential of solu- tions and soils in saline areas. Soil Sci. 117:191-193. Harris, F. S. 1915. Effect of alkali salts in soils on the germination and growth of crops. J. Agr. Res. 15:287-319. H.\rward, M. R., and L McNulty. 1965. Sea- sonal changes in ionic balance in Salicornia rubra. Proc. LTtah Acad. Sci.. Arts. Letters 42:65-69. Hill, T. G. 1908. Observations on the osmotic properties of the root hairs of certain salt marsh plants. New Phytol. 7:133-142. Horwitz, W. 1970. Official methods of analysis of the association of official analytical chem- ists. Association of Official Analytical Chem- ists, Washington, D.C. Kaho, H. 1926. Uber den Einfluss der Tem- peratur auf die koagulierende Wirkung ein- iger Alkalisalze auf das Pflanzenplasma. VIII Biochem. Z. 167:182-194. Levitt, J. 1972. Responses of plants to environ- mental stress. Academic Press, New York. Manus/Fiske. 1972. Chlor-o-counter instruc- tions for use. Fiske Associates, Inc., Massa- chusetts. MoNTFORT C. AND W. Brandrup. 1927. Physi- ologische and Pflanzengeographische Seesalz- wirkungen. I. Okologische Studien uber Kei- mung and erste Entwicklung bei Halophyten. Jahrb. Wiss. Bot. 66:902-946. Neuwohner, W. 1938. Der tagliche Verlant von Assimilation und Atmng bein emign Halophyten. Planta 28:644-679. As abstracted by Biological Abstracts. 1939. Entry No. 1842. Perkin-Elmer. 1970. Analytical methods for atomic absorption spectrophotometry. Nor- walk, Connecticut. ScnoLANDER, P. F., E. D. Bradstreet, H. T. Ham- MEL, AND E. A. Hemmingsen. 1966. Sap concentrations in halophytes and some other plants. Plant Physiol. 41:529-532. Stalter, R., and W. T. Batson. 1969. Trans- plantation of salt marsh vegetation, George- town. South Carolina. Ecology 50:1087-1089. Steiner, M. 1935. Zur Oekologie der Salz- marschen der nordostlichen Vereinigten Staat- en von Nordamerika. Jahrb. Wiss. Bot. 81:94- 202. Ungar, I. A.. W. HoGAN, AND M. McClelland. 1961. Plant communities of saline soils at Lincoln. Nebraska. Amer. Midi. Natur. 82:564-577. Waisel, Y. 1972. Biology of halophytes. Aca- demic Press, New York. Webb, K. L. 1966. NaCl effect on growth and transpiration in Salicornia bigelovii, a salt marsh halophyte. Plant Soil 24:261-265. Yabe, a., G. Kuse, T. Mur.\ta, and H. Takada. 1965. Physiology of halophytes I. The os- motic value of leaves and the osmotic role of each ion in cell sap. Physiol. Ecol. 13:25-33. NEW RECORDS OF STONEFLIES (PLECOPTERA) FROM NEW MEXICO^ Bill P. Stark ,= Theodore A. Wolff ' and Arden R. Gaufin' Abstract. — Distributional data are presented for 30 species of New Mexico Plecoptera including 8 species new to the state list. Previouslj', many of the included species were known from a single lo- cation in the state. A revised checklist of 46 confirmed species is presented for the state. Distribtitional data on the New Mexico stonefly fauna have accumulated slowly because of the isolated nature of much potential stonefly habitat. Records of win- ter-emerging forms are particularly mea- ger, with three sjiecies of Capniidae listed for New Mexico in a review of southwest- ern stoneflies by Stewart et al. (1974). Two species were recorded from a single location and the other from two locations in the state. In this paper we report eight additional species, Malenka flexura (Claassen), Pod- mosta delicatula (Claassen), Zapada haysi (Ricker), Capnia confusa Claassen, Cap- nia gracilaria Claassen, Paraleuctra rick- eri Nebeker and Gaufin, Diura knowltoni (Frison), and Isogenoides zionensis Han- son; new distributional data for species previously reported are also given. Two of us (Stark and Wolff) collected all ma- terial unless otherwise noted. The authors thank M. R. Cather for providing records from her personal col- lection, and R. W. Raumann for sending records from the United States National Museum. Taenionema nigripennis (Ranks). — New Records: Sandoval Co., Santa Clara Can- yon, Wem Povi Pond, elev. 6,900', 20-10- 73, nymphs. Santa Fe Co., Rig Tesuque Creek, Rig Tesuque Campground, elev. 9,700', 21-IV-73, nymphs; Rio Enmidio, Hyde Park Ski Rasin, lO-VI-74, 1 d^. Malenka coloradensis (Ranks). — New Records: Catron Co., Willow Creek, Wil- low Creek Campground, 11-VT74, 2 d" 1 ? . Sandoval Co., Tschicoma Pond, Santa Clara Canyon, 13-VI-74, 3 d' 9 9 . Malenka flexura (Claassen). — Distribu- tion: Taos Co., West Fork Red River, A^heeler Peak Wilderness Area, 10 miles iouth of Red River, 9-VI-74, 25 c^ 14 9 • Podmosta delicatula (Claassen). — Dis- tribution: Rio Arriba Co., Rio Chama, Chama, elev. 7,870', 8-VT74, 9 d" 19 9 . Taos Co., Red River, 2 miles east of Red River, 9- VI- 74, 1 9 . Prostoia hesametsa (Ricker). — New Records: Santa Fe Co., Rio Santa Cruz, near Espanola, elev. 5,600', 17-III-74, 1 d" ; Rig Tesuque Creek, Rig Tesuque Campground, elev. 9,700', 17-III-74, ex- uvium. Zapada cinctipes (Ranks). — New Rec- ords: Sandoval Co., Santa Clara Canyon, Wem Povi Pond, elev. 6,900', 20-111-73, 1 9 . Santa Fe Co., Rig Tesuque Creek, Rig Tesuque Campground, elev. 9,700', 21-IV-73, 5 d' 10 9. Zapada frigida (Claassen). — New Rec- ords: Santa Fe Co., Rig Tesuque Creek, Rig Tesuque Campground, elev. 9,700', 10-VT74, 1 d. Zapada haysi (Ricker). — Distribution: Lincoln Co., North Fork Ruidoso River, 17-V-72, S. M. Fiance, 1 9, nymphs. Santa Fe Co., Rig Testique Creek, Rig Tesuque Campground, elev. 9,700', 10- VT74, 2 d" 5 9 ; same location, 21-IV-73_, nymphs; Rio Enmidio, Hyde Park Ski Rasin, 10-VT74, elev. 10,560', 2 d' 7 9 . Taos Co., Wheeler Peak, 15 to 25-VT60, Rurks and Kinzer, 1 d" 1 9 ; 4 miles north of Arroyo Seco, 22-VI-61, S. G. Jewett, Jr., 2 d" 1 9 ; Rio Hondo, Taos Ski Valley, 22-in-67, R. W. Raumann, nym])hs; Rio Trampas, above El Valle, lO-VI-74, 2 c^ 8 9 ; West Fork Red River, Wheeler Peak Wilderness Area, 10 miles south of Red River, 9-VI-74, 3 d" 4 9 . Capnia confusa Claassen. — Distribtttion: Rio Arriba Co., Rio Chama, Chama, elev. 7,870', 19-IV-73, 13 cf 9 9 . Sandoval Co., Santa Clara Canyon, Wem Povi Pond, elev. 6,900', 20-IIT73, 2 d- San Miguel Co., Pecos River, Tererro, elev. 7,600', 21- IV-73, 26 d" 10 9 . Taos Co., Rio Tram- pas, 1 1/2 miles southeast of El Valle, elev. 7,800', 20-IV-73, 1 9 . Capnia gracilaria Claassen. — Distribu- Tesuque Creek, tion: Santa Fe Co., Rig Study supported by EPA Grant 3053-3G4 and NSF Grant G98G-600. ■Department of Biology, University of Utah, Salt Lake City. 97 98 GREAT BASIN NATURALIST Vol. 35, No. 1 Big Tesuque Campground, elev. 9,700', 21-IV-73, 3 d" 13 ? ; same location, 17- III-74, 2 cf ; Rio Enmidio, Hyde Park Ski Basin, eler. 10,560', lO-VI-74, 2 9 . Taos Co., West Fork Red River, Wheeler Peak Wilderness Area, 9-VI-74, 1 cf 1 $ • Eucapnopsis brevicauda Claassen. — New Records: Santa Fe Co.. Big Tesuque Creek, Big Tesuque Campground, elev. 9,700', 21TV-73, \ d- Paralcuctra rickeri Nebeker & Gaufin. — Distribution: Taos Co.. West Fork Red River, Wheeler Peak Wilderness Area, 10 miles south of Red River, 9 -VI- 74, 12 d 26 9. Paralcuctra vershina Gaufin and Rick- er. — This species has been previously re- ported from the state as P. sara (Claas- sen). New records: Rio Arriba Co.. Rio Chama, Chama, elev. 7,870' 8-VI-74, 2 cT 2 9 . Taos Co.. Rio Trampas, above El Valle, 9-VI-74, 6 c? H ?• Pteronarcella badia (Hagen). — New Records: Sandoval Co.. Santa Clara Can- yon, Wem Povi Pond, elev. 6,900', 20-III- 73, nymphs. Diura knowltoni ( Prison). — Distribu- tion: Rio Arriba Co.. Rio Chama, Chama, elev. 7,870', 8-VI-74, 2 d' 6 9 ; same loca- tion, 25-V-74, M. and E. Cather, nymphs. Taos Co.. Rio Trampas, IV2 miles south- east of El Valle, elev. 7,800', 20-IV-73, n}TTiphs. Isogenoides elongatus (Hagen). — New^ Records: San Miguel Co.. Pecos River, Pecos, 24-V-74, M. and E. Cather, 5c^ 5 9 . _ Isogenoides zionensis Hanson. — Distri- bution: Catron Co.. San Francisco River, Glen wood, 19-11-70, M. Suavely, nymph. McKinley Co.. McGaffey Lake, lO-iV-70, n^niph. Rio Arriba Co.. Rio Chama, Chama, elev. 7,870', 19-IV-73, 1 d (reared) ; same location, 8-VI-74, 6 cT 2 9 . Taos Co.. Red River, Red River, 5- III-61, A. R. Gaufin, nymph; Red River, 3 miles east of Cuesta, '9-VI-74, nymphs. Kogotus modestus (Banks). — New Rec- ords: Taos Co., Red River, 2 miles east of Red River, 9-VI-74, nymphs. Megarcys si gnat a (Hagen). — New Rec- ords: Santa Fe Co.. Big Tesuque Creek, Big Tesuque Campground, elev. 9,700', 21-IV-73, nymphs; same location, 10-VI- 74, 46 cf 1 ? ; Rio Enmidio, Hyde Park Ski Basin, elev. 10,560', 17-III-74, nymphs. Taos Co., Rio Trampas, 11/2 miles southeast of El Valle, elev. 7,800', 20-IV-73, iiymphs; West Fork Red River. Wheeler Peak Wilderness Area, 9-VI-74, nymphs. Skwala parallel a ( Frison ) . — New Rec- ords: 7V/O.S- Co.. Rio Hondo, Hwy. 3, 5-III- 61, A. R. Ciaufin, nymphs. Isoperla ebria (Hagen). — New Records: Santa Fe Co., Big Tesuque Creek, Big Tesuque Campground, elev. 9,700', 10- VI-74, 1 cf; same location, 21-IV-73, nymphs; Rio Enmidio, Hyde Park Ski Basin, elev. 10,560', 17-III-74, nymphs. Isoperla fulva Claassen. — New Records: Santa Fe Co.. Big Tesuque Creek, Big Tesuque Campground, elev. 9,700', 21- IV- 73, n}Tnphs. Isoperla mormona Banks. — New Rec- ords: Catron Co.. Whitewater Creek, Cat- walk Picnic Area, ll-VI-74, 1 d ; West Fork Gila River, 17-V-74, M. and E. Cather, 15 d' 6 9 . Grant Co., Gila River, 31 miles north of Pinos Altos, Hwy 15, ll-VII-73, 1 (S (reared); same location, 17-V-74, M. and E. Cather, 26 d" 17 9 . Claassenia sabulosa (Banks). — New Records: San Miguel Co.. Pecos River, Tererro, elev. 7,600', 21-IV-73, nymphs. Hesperoperla pacifica (Banks). — New Records: Catron Co.. Whitewater Creek, 16-V-74, M. and E. Cather, exuvium. Sweltsa coloradensis (Banks). — New Records: Catron Co., Willow Creek, Wil- low Creek Campground, ll-VI-74, 8 d 7 9 . Rio Arriba Co.. Rio Chama, Chama, elev. 7,870', 8-VI-74, 1 d' 3 9 . Sandoval Co., Tschicoma Pond. Santa Clara Can- yon, 13-VI-74, 2 9. Sweltsa lamba (Needham & Claassen). — New Records: Taos Co.. West Fork Red River, Wheeler Peak Wilderness Area, 10 miles south of Red River, 9-VI-74, 1 d- Triznaka diver sa (Frison). — New Rec- ords: Santa Fe Co.. Big Tesuque Creek, Big Tesuque Cami)ground, elev. 9,700', 10- VI- 74, 31 d^ 24 9 . Triznaka pintada (Bicker). — New Rec- ords: Taos Co.. Rio Pueblo, Penasco, 9-VI- 74, 2 d' 1 3 9 . Triznaka signata (Banks).— New Rec- ords: Rio Arriba Co.. Rio Chama, Chama, elev. 7.870', 8-VI-74, 17 d 10 9 . Ni.w Mi:xic:o List Below is a checklist of 46 stonefly spe- cies that lune been confirmed for New Mexico. Sjiecies regarded as questionable March 1975 STARK, ET AL.: NEW MEXICO PLECOPTERA 99 or unconfirmed by actual specimens and distributional data have been omitted. Taeniopterygidae Taenionema nigripennis (Banks) T. pallida (Banks) Taenioptcryx sp. Nemouridae Amphinemura mogollonica Baumann and Gaufin Malenka coloradensis (Banks) M. flexura (Claassen) Podmosta delicatiila (Claassen) Prostoia besametsa (Ricker) Zapada cinctipes (Banks) Z. frigida (Claassen) Z. haysi (Ricker) Capniidae Capnia confusa Claassen C. fibula Claassen C. gracilaria Claassen Eucapnopsis brevicauda Claassen Mesocapnia frisoni (Baumann and Gaufin) Leuctridae Paraleuctra rickeri Nebeker and Gaufin P. vershina Gaufin and Ricker Perlomyia utahensis Needham and Claassen Pteronarcidae Pteronarcella badia (Hagen) Pterormrcys califnrnica Newport Perlodidae Cultus aestivalis (Needham and Claassen) Diura knoivtoni (Prison) Isogenoides elongatus (Hagen) /. zionensis Hanson Kogotus modestus (Banks) Megarcys signata (Hagen) Skwala parallela (Prison) Isoperla ebria (Hagen) . fulva Claassen /. longiseta Banks /. mormona Banks /. patricia Prison /. phalerata (Smith) / quinquepunctata (Banks) Perlidae Claassenia sabulosa (Banks) Hesperoperla pacifica (Banks) Neoperla clymene (Newman) Chloroperlidae Paraperla frontalis Banks Suwallia pallidula (Banks) Sweltsa borealis (Banks) S. coloradensis (Banks) S. lamba (Needham and Claassen) Triznaka diver sa (Prison) T. pintada (Ricker) T. signata (Banks) Literature Cited Stewart, K. W.. R. W. Baumann, and B. P. Stark. 1974. The distribution and past dispersal of southwestern United States Ple- coptera. Trans. Amer. Entomol. Soc. 99:507- 546. THE AUTHORSHIP AND DATE OF PUBLICATION OF SIREN INTERMEDIA (AMPHIBIA: CAUDATA) Hobart M. Smith^, Rozella B. Smith-, and H. Lewis Sawin^ Abstract. — However "just" it might be to credit LeConte, 1828. with Siren intermedia, both Harlan, 1826 (not 1827 as often cited), and Barnes, 1826, antedate LeConte's proposal of the name. As the earliest, Barnes, 1826, stands credited with it. In analysis of precedent for these conclusions, types of taxonomic plagiarism (calculated vs. innocent, homoplagiarism vs. heteroplagiarism) and the distinctions between nomina nuda and nomina dubia are reviewed, giving examples of each category. does not nullify applicability of Art. 50 of the International Code of Zoological No- menclature (ICZN, 1964: 49), which states, "The author ... of a scientific name is . . . the person . . . who first pub- lishes it in a way that satisfies the criteria of availability, unless it is clear from the contents of the publication that . . . some other person is alone responsible for both the name and the conditions that make it available" (italics ours). LeConte obviously was responsible for the name but equally clearly was not re- sponsible for the "description" that "satis- fies the criteria of availability." Harlan obviously wrote the description; and de- spite his aj)parent wish to the contrary, the present rules would require that he be regarded as author of the name in zoo- logical nomenclature if indeed his ac- count were the earliest to have appeared. There is ample precedent for crediting the immediate source of an}" given name and its characterization, however ques- tionable may be the derivation of either, for that name. This policy unfortunately rewards plagiarism with permanence un- less the International Commission on Zo- ological Nomenclature intercedes. On the other hand, plagiarism seldom occurs, either inadvertently or deliberately. Nev- ertheless, it does occur on occasion, and the Code requires that the perpetrator bear responsibility for his act, whether it be innocent or calculated. Examples of cal- culated plagiarism are provided by Thom]:)son's three privately printed not- ices of 1912; the first two antedated Van Denburgh's competitive advance diagnosis of 1912, and although Thompson's de- scri])tions are sourced directly from Van Denburgh's manuscript, insofar as they antedate Van Denburgh's descriptions Martof (1973: 1-3), in the most recent review of Siren intermedia, notes that the earliest full description of the species in LeConte (1828: 133-134, pi. 1) actually was antedated by a brief but nominally occupying characterization, credited to Le- Conte, that appeared in a work b}" Har- lan (1826: 322), dated 1827 by Schmidt (1953: 14) and others. Two points merit observation in this context: (1) the particular page on which the "description" of Siren inter- media appeared in Harlan's work was ac- tually published in 1826, fide the 1913 In- dex to the Scientific Contents of the Journal and Proceedings of the Academy of Natural Sciences of Philadelphia, p. viii; and (2) there is reason to accept the author of this "description" as Harlan, not LeConte. Harlan "read" his paper at the 12 Dec. 1826 meeting of the Academy, and accordingly- the pages published in 1826 (pp. 317-324) must have appeared sometime after 12 Dec, the remainder (pp. 325-372) in February 1827, accord- ing to the Index. The article was com- pleted in no. 1 of vol. 6 of the Journal (pp. 7-38) appearing in March 1827 fide the same source. The author of the description appearing in Harlan (1826: 322) is clearly Harlan, not LeConte, despite the fact that Harlan attributed the name to LeConte and stat- ed (in a footnote) that the material on this species was sourced from "manuscript notes." The characterization obviously was written by Harlan, not LeConte, as becomes evident when one consults Le- Conte's formal description that appeared in 1828. Harlan seemingly saw the Le- Conte ms. and published in his own words the name and certain characters cited in the ms. The acknowledgment of source 'Department of Environmental. Population, and Organist Biology and Museum, University of Colorado, Boulder ^Department of EPO Biology and Center for Computer Research in the Iluma 'Center for Computer Research in the Humanities, University of Colorado. lies. University of Colorado. 100 March 1975 SMITH, ET AL.: AMPHIBIAN NOMENCLATURE 101 they are accepted under the Code as valid (see Barbour, 1917, for details). Examples of innocent taxonomic plagi- arism fall into two categories: self -plagi- arism (or homoplagiarism) and hetero- plagiarism. A medium for frequent homo- plagiarism is Dissertation Abstracts, wherein summaries of doctoral disserta- tions occasionally include sufficient in- formation with a new name or a new com- bination to occupy them; for example Walker's abstract (1967) includes suffi- cient information on two new names {Cnernidophorus gularis rciuni, C. g. semi- annulatus) to occupy both, whereas it was intended that these names not be en- tered into nomenclature before full docu- mentation could be provided (full descrip- tions have not even yet appeared). A similar case occurred in another journal (Harris, 1974), wherein a photograph and brief notice of some characteristics of a new subspecies of rattlesnakes was ac- companied by a name {Crotalus ivillardi obscurus), thus occupying the name in advance of the intended date and work which was then in press. An example of heteroplagiarism oc- curred with inadvertent mention of Palm- atotriton by Smith (1945), who used the name under the impression that his for- mer professor, E. H. Taylor, had a ms. in press establishing the genus, and that the casual mention in the popular journal would be meaningless. Unfortunately Taylor had decided against erection of the genus, and, more regrettably. Smith's use of the name was accompanied by a few incidental comments inadvertently serving to occupy the name nomenclatur- ally. It was necessary to appeal to the International Commission on Zoological Nomenclature to "deoccupy" Palmatotri- ton as of Smith, 1945, making the name available for use by anyone else, in any desired sense (ICZN, 1956). In all these examples, including that of Harlan, it is clear that intent has noth- ing to do with result; only the briefest characterization, in but a few words, may serve to occupy a name even when not so intended, and the person responsible is the one presenting those words, even though he may not have intended to re- ceive that responsibility. In this context it is important to recog- nize that a name may be occupied even though its characterization may be inade- quate for definitive allocation to its prop- er taxon in nature; such names are nom- ina diibia despite the fact that they are occupied names. There is a rather wide misapprehension that a full characteriza- tion is required in order to occupy a new name, but this is not so. Nomina dubia are often rendered identifiable (i.e., nom- ina clara) by subsequent provision of further details, as is true in the case of Siren intermedia. Harlan's description, although adequate to occupy the name were it the original usage, would not alone have sufficed for allocation; but with Le- Conte's full account, no doubt remains. As of Harlan, Siren intermedia is a no- men dubiiim; as of LeConte, it became a nomen clarum although occupied at an earlier date by another author. Harlan's usage was not of a nomen nudum, which is nonexistent nomenclaturally, because it did provide some distinguishing informa- tion. The Code makes clear (Art. 13) that any "statement that ]:)urports to give char- acters differentiating the taxon" (italics ours) suffices to occupy an accompanying name, and practice has conformed with this liberal rule. In the case of Siren intermedia, how- ever, the comedy of errors did not really begin with Harlan, even of 1826. There is a still earlier usage that occupied the name. Barnes (1826: 269, footnote) saw or otherwise knew of LeConte's ms and rendered the name Sir-eii intermedia avail- able in almost precisely the same way that Harlan's work would have done had it been the earliest usage. The Barnes foot- note follows: ''Additional note communi- cated by the author, Aug. 15, 1826. The delay in the printing of this paper has given the author an opportunity of an- nouncing, in this place, the discoverv of ANOITTER NEW SPECIES OF SIREN, by Capt. LECONTE. It belongs to this section, and is called by its discoverer Siren intermedia. In its color it resembles the Lacertina, and in its gills, the Striata; but it has peculiar characters of its own, which will be explained at length in a paper soon to be published in the Annals of the Lyceum. Length about one foot, inhabits the Southern states in large num- bers. Specimens are preserved in the Cabinet of the Lyceum. Fig. Annals of the Lyceum, Vol. 2, fig. 1." That Barnes knew of LeConte's description long be- 102 GREAT BASIN NATURALIST Vol. 35, No. 1 fore its publication is not surprising, in- asmuch as he was the "Recording Secre- tary of the New York L}xeum," as indi- cated (p. 268) in his 1826 paper. Only by the close familiarity permitted by such an association could he have known some two years in advance of publication that LeConte's account would appear in Vol- ume 2 and incorporate Figure 1 of the Annals of the Lyceuin! The Harlan and Barnes works were both dated 1 826, but the Harlan paper appeared very late in the year — certainly after December 12 — whereas the Barnes paper, read before the Lyceum in July 1825, was surely published shortly after 15 August 1826^ when Barnes inserted his footnote on S. intermedia. We have not been able to pinpoint the exact date of publication of either work, but the evidence that Barnes' work preceded that of Harlan is overwhelming. The same generalities })ertinent to Har- lan's use of the name SireJi intermedia are equally pertinent to the earlier Barnes usage. Barnes actually must be regarded as the author of Siren intermedia (which accordingly dates from 1826), unless the case is appealed to the ICZN, asking for rejection of the contributions of both Barnes and Harlan on that species, giving LeConte (1828) priority. The effort is not warranted, however, since the significance of the case is grossly inadequate to justify the protracted, laborious protocol involved in ICZN action. Custom dictates that sus- pension of the rules be requested only for names of relatively broad familiarity among zoologists; the present certainl}- does not fall into that category. We are accordingly left with the con- clusion that the proper citation for the species under consideration is Siren inter- media Barnes, 1826. Literature Cited B.^RBOUR, T. 1917. A most regrettable tangle of names. Occ. Pap. Mus. Zool. Univ. Michigan 44:1-9. Barnes, D. H. 1826. An arrangement of the genera of batracian [sic] animals, with a de- scription of the more remarkable species; in- cluding a monograph of the doubtful reptils [sic]. Am. J. Sci. 11:268-297. Harl.^n, R. 1826-27. Genera of North American Reptilia and a synopsis of the species. J. Acad. Nat. Sci. Philadelphia 5:317-372; ibid., 6:7-38 (pp. 317-324. Dec, 1826; 325-372, Feb., 1827; 7-38, Mar., 1827). Harris, H. S., Jr. 1974. The New Mexican ridge-nosed rattlesnake. Nat. Parks Cons. Mag. 48(3): 22-24, 3 figs. International Commission on Zoological Nomen- clature. 1956. Opinion 425: Addition to the "Official index of rejected and invalid generic names in zoology" of the name ''Pabnatotri- ton' Smith (H. M.). 1945 (Class Amphibia). Opin. Decl. Int. Comm. Zool. Nom. 14:243- 256. . 1964. International code of zoological nomenclature adopted by the fifteenth inter- national congress of zoology. London, Intern. Trust Zool. Nomencl. xviii, 176 pp. LeConte, J. E. 1828. Description of a new species of Siren. Ann. Lyceum Nat. Hist. New York 2:133-134, pi. 1. Martof, B. S. 1973. Siren intermedia. Cat. Am. Amph. Kept. 127:1-3. map. Schmidt, K. P. 1953. A check list of North American amphibians and reptiles. Chicago, Am. Soc. Ichth. Herp. vii. 280 pp. Smith, H. M. 1945. Herpetological collecting in banana fields of Mexico. Ward's Nat. Sci. Bull. 19(l):3-7. figs. 1-6. Thompson. J. C. 1912. Prodrome of a descrip- tion of a new genus of Ranidae from the Loo Choo Islands. Herpetological Notices 1:1-3, 1 pi. . 1912. Prodrome of descriptions of new species of Reptilia and Batrachia from the Far East. Herpetological Notices 2:1-4. . 1912. On reptiles new to the island arcs of Asia. Herpetological Notices 3:[i-ii], 1-5. Van Denrurgh. J. 1912. Advance diagnoses of new reptiles and amphibians from the Loo Choo Islands and Formosa. Privately Printed. 5 pp. Walker. J. M. 1967. Morphological variation in the teid lizard Cnemidophorus gularis. Diss. Abst. Int., B 28:1738-1739. NEW MITES FROM THE YAMPA VALLEY^ (ACARINA: CRYPTOSTIGMATA: ORIBATULIDAE, PASSALOZETIDAE) Harold G. Higgins- and Tyler A. Woolley^ Abstract. — A study was made of the soil mites from under different vegetative types near a coal-burning power plant in the Yampa Valley near Hayden, Colorado. The following new species of oribatids were found: Zygoribatula apletosa n.sp., Multoribates haydeni n.sp., Paraphauloppia cordylin- osa n.sp., Passalozetes moniles n.sp. Concentrated collections have been made near a coal-burning power plant in the Yampa Valley near Hayden, Colora- do. A number of new and unrecorded species of soil mites for Colorado have been found in the project area. The col- lections were made with reference to the soil forms and to vegetative types with which they were found. As might be ex- pected, many species appear to be more abundant at one season of the year than at another or may be more closely as- sociated with certain vegetative types than with others. This concentrated collecting over a two- year period has given new insight as to the importance of the microclimate in the biology of oribatids. For example, depend- ing on the amount of moisture, sunlight or shade, and slope, many species may be more abundant under one side of a bush than under the other. Preliminar}^ studies also indicate that destruction of vegetation and distiu-bances of the soil in such changes as the formation of spoil banks or strip-mining and pollution from coal- burning power plants seriously depletes the numbers and kinds of soil arthropods. Following are descriptions of four new species representing two families of orib- atids found in the Hayden area. Family Oribatulidae Zygoribatula apletosa, n.sp. Figs. 1 and 2 Diagnosis. — Large size, larger than any known Zygoribatula; with 14 pairs of large, setose notogastral setae; rostral hairs further apart than lamellar hairs; lamel- lae curved inward with distinct translam- ella; areae porosae Aa located near the small shoulder projections. The trivial name apletosa is modified from the Greek, and implies "immense," referring to the size of these oribatids. Description. — Color reddish-brown; rostrimi rounded; rostral hairs heavy, reaching beyond tip of rostrum by about half their lengths, hairs farther apart than lamellar setae; lamellae of almost uniform width throughout, length curved inward toward anterior tip, slightly less than one-half as far apart at tip as at base; translamella narrower than lamel- lae; lamellar hairs similar to rostral hairs but about one-third longer inserted in anterolateral ends of lamellae; interlamel- lar hairs situated midway between inser- tions of lamellar hairs and pseudostig- mata, closer to inner margin of lamellae; pseudostigmata cuplike with edge erected above surface of prodorsiun; sensillum with broad, rounded setose head and short pedicel, about half as long as interlamel- lar hair; exobothridial hair rather heavy and stiff. Hysterosoma longer than broad, widest near middle, with tapering posterior end; dorsal surface with 14 pairs of heavy, long, spined setae, many extending beyond body outline as shown in Figure 1; areae porosae all large, Aa much longer than broad and located near small humeral process. Camerostome oval in outline; ventral surface with apodemata and setae as shown in Figure 2; genital and anal aper- tures far apart, smaller genital opening more than twice its length anterior to larger anal aperture; each genital cover with four setae; aggenital setae as shown in Figure 2; each anal cover with two setae (2 of 12 specimens with 3 anal se- tae) ; two adanal setae present. Legs all about equal in size; heterotri- dactylous, median claw larger than later- als. ^Yampa Valley Project, Ecologj- Consultants, Inc. -Biology- Department, Granger High School, Granger, Utah. ^Department of Zoologj- and Entomology, Colorado State University, Fort Collins, Colo. 80523. 103 104 GREAT BASIN NATURALIST Vol. 35, No. 1 Fig. 1. Zygoribatula apletosa, dorsal aspect, legs omitted. Measurements. — length .705 mm; width, .495 mm. (Range .853-.705 mm X .600-. 495 mm). The type (a male) and 6 paratypes (6 females) were taken under serviceberry about 1/4 mile NE power plant, Hayden, Colorado, 7 Oct. 1971, by H. G. Higgins; 2 specimens, females, were taken under rosebush, I/2 mile N power plant, Hayden, Colorado, 21 June 1972; 2 specimens (males) were taken i^ mile N power plant, Hayden, Colorado, 1 Aug. 1971; 1 specimen (male) was taken under aspens, 4 miles S Seneca Road, Hayden, Colorado, 1 Aug. 1971; all by H. G." Hig- gins. Discussion. — This species stands a- part from other known North American Zygoribatula by its large size, its long, heavy, setose body setae, and by its big, long, areae porosae Aa located near the shoulder. In general appearance Z. aple- tosa n.sp. resembles Z. lata Hammer but differs in the much larger size as well as in the size and locations of areae porosae. Fig. 2. Z. apletosa, ventral aspect. To date, this species has always been found associated with rather heavy, moist litter under dense vegetation. This species shows interesting variation in the width of the translamellae and lo- cation of body setae. Also, as pointed out earlier, 2 of 12 specimens have 3 pairs, rather than 2 pairs, of anal setae. Multoribates haydeni, n.sp. Fig. 3 Diagnosis. — Similar to Multoribates chavinensis Hammer, 1961, but larger, and with only 11 pairs of dorsal setae; lacks the ventral keel on femur II. The trivial name is indicative of location. Description. — Large size; color yel- lowish to light brown; body egg-shaped with the pteromorphs hardly projecting beyond lateral outline of body; prodorsum triangular in outline with rostrum fairly pointed, often hyaline; rostral setae in- serted posteriorly on lateral margins of propodosoma, much wider apart than la- mellar hairs; lamellae narrow, tapering slightly anteriorly; translamellae absent; lamellar hairs stiff, setose about same length as lamellae; interlamellar hairs March 1975 HIGGINS, WOOLLEY: COLORADO MITES 105 /f ^^ Fig. 3. Multoribates haydeni, dorsal aspect, legs omitted. heavy, nearly equal in length to lamellar hairs; pseudostigmata cuplike, rim project- ing beyond the body level; sensillum with narrow stalk and broad, setose head that is bent backwards; dorsosejugal suture curved anteriorly. Hysterosoma longer than broad with small pteromorphs that project only slightly beyond the outline of body; 11 pairs of fine, simple dorsal hairs visible as shown in Figure 3; areae porosae absent, but replaced with chitinous pores; muscle scars and markings visible round edge of hysterosoma as indicated in Figure 3. Var- iations occur in the locations of body se- tae. Ventral surface similar to M. chavinen- sis with only a few minor exceptions; gen- ital plates separated from larger anal plates by approximately twice their length, each plate with four setae; ana] plates much larger than genital plates, situated near posterior end of body, each anal plate with two hairs; adanal and aggenital setae place similarly to M. chav- inensis; fissure iad located near antero- mediad margin of anal plates. Legs about equal in size; all legs hetero- tridactylous, median claw larger than lat- erals; femur II without visible keel. Measurements. — Length, .45 mm; width, .26 mm. The type, a gravid fe- male was collected at Seneca #2, Hayden, Colorado, under serviceberry, 10 April 1971, by H. G. Higgins and T. A. Wool- ley. Additional specimens are as follows: I specimen at Seneca :^2, Hayden, Colo- rado, 9 June 1971, in sagebrush, by T. A. Woolley and H. G. Higgins; 3 specimens from under aspens associated with bitter- brush (Purshia), 8 June 1971, 4 miles S Seneca Road, Hayden, Colorado, by H. G. Higgins and T. A. Woolley; 6 specimens from under bitterbrush, 5 miles S Seneca Road, Hayden, Colorado, 1 Aug. 1971, by H. G. Higgins; 8 specimens from under bitterbrush, 5 miles S Seneca Road, Hay- den, Colorado, 8 Oct. 1971, by H. G. Hig- gins; 1 specimen from under bitterbrush, 4 miles S Seneca Road, Hayden, Colorado, 21 June 1972, by H. G. Higgins. Discussion. — In general appearance M. haydeni, n.sp. resembles M. chavinen- sis Hammer but is larger, lacks the ventral keel on femur II, and has 11 rather than 14 pairs of dorsal setae. Preliminary study seems to indicate that although this new species is found in several habitats, it pre- fers the microhabitat under bitterbrush in rather arid conditions. Paraphauloppia cordylinosa, n.sp. Fig. 4 Diagnosis. — Similar in outline to Para- phauloppia novazealandica Hammer, 1967, but with much larger lamellae and II pairs of notogastral hairs. The name cordylinosa refers to the clublike sensillum of the new species. Description. — Color yellowish; ros- tral setae large, rough, situated on the an- terolateral margins of the propodosoma; lamellae quite large, extended more than half the length of propodosoma, about equal in width throughout theirs lengths, with a small spur (prolamella) located anteromediad; lamellar hairs inserted in anterior tip of lamellae, extending to tip of rostrum, heay>', barbed, and about equal in length to rostral hairs; interla- mellar hairs three-fourths as long as la- mellar hairs, inserted mediad, closer to pseudostigmata than to tip of lamellae ex- tending to the sides of prodorsum; pseudo- stigmata cuplike with a short stalk and expanded, rounded, setose head. Hysterosoma oval, tapering anteriorly; dorsosejugal suture greatly arched; ptero- 106 GREAT BASIN NATURALIST Vol. 35, No. 1 Fig. 4. Paraphauloppia cordylinosa, dorsal as- pect, legs omitted. morphs absent; 11 pairs of simple smooth, dorsal hairs as shown in Figure 4. Areae porosae visible but not noticeably en- larged as in P. novazealandica; a line of light marking, probably muscle scars, vis- ible mediad of lateral margins. Anal opening much larger than geni- tal opening and situated near posterior end of body, each cover with two setae; genital opening smaller, situated about twice its length in front of anal opening, three pairs of visible hairs on each plate; aggenital setae posterior to genital plate; aggenital and adanalsetae located similarly to P. novazealandica, adi ])osterior to anal plate, ado situated near the middle of plate along lateral side, and ads situated anteri- or to anal plate. All legs about equal in size, hetero- tridactylous, with median claw larger than laterals. Measurements. — Length, 282^1,; width, 132/i. The type (a female) and a para- type from sagebrush, Yampa Airport, Hayden, Colorado, 9 June 1971; 6 speci- mens from sagebrush, 2 miles S Yampa Airport, Hayden, Colorado, 9 June 1971; all by H. G. Higgins and T. A. Woolley. Discussion. ^ — In general appearance this species resembles P. novazealandica Hammer but differs in having much lar- ger, heavier lamellae and 11 rather than 10 pairs of dorsal setae. Although this taxon differs somewhat from Paraphaulop- pia in the number of dorsal setae, it ap- pears to be nearer this genus than to Phauloppia. We hesitate, at this time, to describe a new genus based on these minor differences and because in the small sample of mites collected there is great variation in the exact location of the dor- sal setae, and the lengths of the lamellar hair. It is interesting that although col- lections were made at the same general site several times a year, and over a two- year span, specimens of this species were found only once, in June 1971, and those in rather dry sagebrush habitat. Prelimin- ary postulations attribute this to the pos- sible influence of pollutants in the area. Family Passalozetidae Passcdozetes moniles, n.sp. Figs. 5-6 Diagnosis. — This species is readily separated from P. linearis, the only other known North American species, by the banded, beadlike pattern of dorsal and ventral integuminal and by the smooth sensillum with a pointed tapered head. The trivial name moniles is modified from the Latin meaning "necklace" and refers to the beaded appearance of the integu- ment. Description. — Yellowish in color; pro- dorsimi slightly wider than long; rostrum blunt, rounded; rostral hairs simple, in- serted near tip of rostrum, curved medi- ally toward tip of rostrimi; lamellae ab- sent; lamellar hairs small, simple, slightly longer than rostral hairs, curved down to- ward tip of rostrum; interlamellar hairs threadlike, simple, inserted anteromediad of pseudostigmata, adjacent to coalesced median section of dorsosejugal suture; l)seudostigmata cuplike, separated from each other by a little more than length of sensillum; sensillum with a narrow curved base and tapering into a narrow pointed head. Hysterosoma oval, anterior margin ex- tended forward, coalesced medially with March 1975 HIGGINS, WOOLLEY: COLORADO MITES 107 ^rv/,l;M^»\ Fig. 5. Passalozetes moniles, dorsal aspect, legs omitted. ted. Fig. 6. P. moniles, ventral aspect, legs omit- dorsum of propodosoma beyond level of interlamellar hairs; dorsosejugal suture in- terrupted by this middle projection; lenti- culus clear, round, surrounded by lines; dorsum with simple hairs as shown in Figure 5. Integument of fine lines with darker pigmented areas resembling strings of beads (the beadlike cerotegument may be removed by soaking in lactophenol) ; two pairs of area porosae and a glandular fissure as seen in Figure 5. Camerostome with rather parallel sides, longer than wide; ventral plate as seen in Figure 6; genital covers each with four pairs of genital setae; aggenital setae in- serted about twice their lengths postero- laterad of genital aperture; anal aper- ture nearly one-third larger than anal opening; anal aperture in the posterior end of ventral plate, each cover with two setae; adanal setae difficult to find in the cerotegument, visible setae and glands as shown in Figure 6. All tarsi heterobidactylous, heavier of the two claws toothed (median or lateral) . Measurements. — Length, .36 mm; width, .15 mm. The type (a male) and 3 para types were taken from under bitterbrush and squaw- brush, 6 miles E Craig, Colorado, 14 June 1972; 4 specimens from burned-over area at Seneca #3, Hayden, Colorado, 21 June 1972; all by H. G. Higgins. Discussion. — As is to be expected, there is considerable variation in the ar- rangement of setae and pigmentation in specimens examined. Those examples from the burned-over area were more heavily pigmented and have a wider hys- terosoma than those taken near Craig. The only previously described Passalozetes from this western area was taken from a dry woodrat nest in Tooele Co., Utah, several hundred miles to the west of the location of this new species. Both North American species of Passalozetes have been taken from dry desert sands, which implies that they are found in xeric habi- tats. References AoKi, J. 1961. On six new oribatid mites from Japan. Jap. J. Sanitary Zool. 12(4): 233-238. . 1964. Some oribatid mites (Acarina) from Laysan Island. Pacific Insects 6(4): 649-664. Balogh, J. 1965. A synopsis of the world ori- batid (Acari) genera. Acta Zool. 11(1-2): 5- 99. CoETZER, A. 1967-68. New Oribatulidae THOR, 108 GREAT BASIN NATURALIST Vol. 35, No. 1 1929 (Oribatei, Acari) from South Africa, new combinations and a key to the genera of the family. Mems. Inst. Invest. Ciest. Mocamb. 9, Serie A: 13-126. EwiNG, H. E. 1913. New Acarina. Bull. Amer. Mus. Nat. Hist. 32(5): 93-121. . 1917. New Acarina. Bull. Amer. Mus. Nat. Hist. 37(2): 149-172. Hammer, M. 1961. Investigations on the oriba- tid fauna of the Andes Mountains. II. Peru. Biol. Skr. Dan. Vid. Selsk. 13(1): 1-150 + plates. . 1967. Investigations on the oribatid fauna of New Zealand, Part II. Biol. Skr. Dan. Vid. Selsk. 15(4): 1-64 + plates. Higgins, H. G., .\nd T. a. Woolley. 1962. A new species of Passalozetes from Utah with notes on the genus. (Acarina: Oribatei). Great Basin Nat. 22(4): 93-100. Jacot, a. p. 1961. Journal of North American moss-mites. J. N. Y. Ent. Soc. 45(3-4): 353- 375. WiLLMANN, G. 1931. Moosmilben oder Oribati- den (Oribatei). In: Tierwelt Deutschlands 22(5): 79-200. Woolley, T. A. 1957. Redescription of Ewing's oribatid mites. Ill — Family Eremaeidae (Acarina: Oribatei). Ent. News 68(4): 147- 156. . 1961. Redescriptions of Ewing's oriba- tid mites. Ill — Family Oribatulidae (Acarina: Oribatei). Trans. Amer. Microscop. Soc. 80(1): 1-15. Higgins, H. G., and T. A. Woolley THE IDENTITY OF BOCOURT'S LIZARD EUMECES CAPITO 1879 Hobai't M. Smith\ Rozella B. Smith\ and Jean Guibe" Abstract. — Eumeces capito Boucourt, 1879, is a senior synonym of Eumeces xanthi Giinther, 1889. No exception to application of the Law of Priority is recommended in this case. The type locality of E. capito as originally published ("La cote oriental des Etats-Unis") is erroneous. Undoubt- edly the correct locality is China, but it is not restricted at present. In preparation of "Synopsis of the Herpetology of Mexico," a problem has arisen: whether or not to apply the name Eumeces capito Bocom-t (1879:429-431, pi. 22D, Figs. 8, 8a-8c) to some Mexican species. The origin of the only specimen (holotype, no. 5531 of the Museum Na- tional d'Histoire Naturelle, Paris) is in doubt, for although the published locality is "la cote oriental des Etats-Unis," the several registers in the Museum in Paris give still other indications: "Mexique" and "Amerique septentrionale." The col- lector is also unknown, for M. S. Bracon- nier, by whom Bocourt {loc. cit.) indicat- ed the specimen was "donne," was merely an assistant in the Museum, never par- ticipating in any expedition; he simply made this and other specimens available for study by Bocourt and other scientific personnel. Taylor's exhaustive monograph (1936: 28, 231-2, 506) of Eumeces sheds no light upon the problem of the identity of E. capito, as the type was not examined, and by description alone the name could not be allocated. Taylor did note the possi- bility that the name applies to his Eu- meces inexpectatus, 1932, although he thought it "probable that it is based upon an aberrant specimen of jasciatus" (p. 232). We have thus been motivated to re- examine the only specimen constituting the hypodigm for Eum.eces capito^ being made available by the junior author for study by all of us. It proves to be a typi- cal, mature example of Eumeces xanthi Giinther (1889:218). The specimen is in excellent condition, slightly softened, but not notably damaged, discolored, or faded. Salient data are: snout- vent 72 mm, hind leg 28 mm, foreleg 19 mm, axilla-groin 40 mm, tail 69 mm (30 mm regenerat- ed); 24 scale rows around midbody; 54 scales from parietals to above anus; 16-16 lamellae under 4th toe; a conspicuous patch of enlarged postfemoral scales; one postnasal; two postmentals; parietals nar- rowly separated posteriorly by tip of in- terparietal; frontonasal broadly contacting frontal; upper secondary temporal quad- rangular, dorsal and lateral edges nearly parallel, separated from 7th (posterior) supralabial by contact of primary tem- poral and lower, subtriangular secondary temporal; two pairs of nuchals; median preanals overlapped by lateral scales; a somewhat modified, slightly keeled lateral postanal; median subcaudals twice as wide as adjacent scales; a distinct, brown later- al stripe on 4th scale row and edges of adjacent 3rd and 5th rows; a dorsolateral light stripe occupying most of the 3rd scale row; and a lateral light stripe occu- P3'ing the lower half of the 5th scale row; no evidence of a median light stripe on head, and its only evidence on trunk the absence of dark pigment on the adjacent halves of the 2 median scale rows; other dorsal scale rows with some dark flecking on the base of each scale. Other features as indicated in the accompanying figures. Most of the characters of this specimen conform with those detailed by Taylor (1936:239-243, Fig. 33, PL 15) for Eu- meces xanthi, and indeed the holotype of E. capito closel}' resembles one of the syn- types figured on his Plate 15 (Fig. 3), although the pigment loss has not attained the level there depicted for a 76 mm speci- men. Critical are the modified lateral postanals, the enlarged postfemorals, the low number (24) of scale rows, and the position of the dorsolateral light stripe on the 3rd scale row, in addition to the single postnasal and paired postmentals. No American species of Eumeces have en- larged postfemorals; only a few Asiatic species possess them, and all except E. ^Department of En\-ironmental, Population, and Orgnnismic Biology, University of Colorado, Boulder, 80302, U.S.A. ^Museum National d'Histoire Naturelle. 25 Rue Cuvier. Paris. 75005, France. 109 110 GREAT BASIN NATURALIST Vol. 35, No. 1 Fig. 1. Holotype of Eumeces capita Bocourt, dorsal view. xanthi are eliminated from consideration by number of scale rows or by the post- nasal-postmental characters. The only notable discrepancy between the holotype of E. capita and Taylor's account of E. xanthi is the low number of dorsals (54) in the former, as compared with the range (56 to 60) for the latter. However, northern examples of E. xanthi tend to have fewer dorsals (56-59, com- pared with 59-60 for southern examples) ; nevertheless, northern examples tend to have 22 scale rows, whereas southern ones usually have 24. We suspect that the type of E. capita was taken in more northern parts of the species' range, con- forming more closely with the geographic variant that bears the name Eumeces pekinensis Stejneger (1924:120), type lo- cality Hsin-Lung-Shan district, imperial hunting grounds, Chihli Province, 665 mi N Peking, China, than with the geograph- ic group represented by Eumeces xanthi (type locality Ichang, Hupeh, China). The contact of frontonasal with frontal that occurs in the type of E. capita con- forms with Stejneger's description and figures (1925:49-51, fig. 2) for the three types of E. pekinensis and with Taylor's figure (1936:242, fig. 33), in which the two prefrontals are in contact and there- fore separate frontal and frontonasal. However, the figured specimen is from the same district as the types of E. pe- kinensis; presumably the character is not taxonomically significant. On the con- trary, the number of dorsals and number of scale rows may indeed be significant. /x\ I Fig. 2. Dorsal view of head of the holotype of Eumeces capita Bocourt. March 1975 SMITH, ET AL.: LIZARD IDENTITY 111 Fig. 3. Lateral view of the head of the holo- type of Eumeces capita Bocourt. The minute spheres abundantly evident in this figure, and less abundant in Figs. 2 and 4, are air bubbles. All photos were taken of the specimen under water. at least subspecifically; more material will be required to establish the nature of the variation that occurs in the species. Cer- tainly the low number of dorsals in the type of E. capita casts no doubt upon proper allocation with E. xanthi^ although it may be important in naming the geo- graphic races of that species at some time in the future. We are not aware of more recent studies of E. xanthi that would shed any hght upon the geographic variation of that species. The related species E. tamdaoen- sis Bourret (1937:19-21, fig. 5) is very similar and may well be referable to E. xanthi as a geographic race, but the origi- nal description does not note presence or absence of enlarged postfemorals; a pe- culiar head pattern of juveniles may be distinctive, as well as the greenish color above and below in life. Eumeces coreen- sis Doi and Kamita (1937:211-215, figs.) is not closely similar, being related more closely to E. chinensis (no postnasal, no enlarged postfemorals). However, the wide range of E. xanthi (and its close relatives E. tamdaoensis, E. elegans and E. tunganus) suggests that a polytypic species or a species complex may be in- volved, the nomenclature of which is far from stable. Because of this primitive state of knowl- edge of variation in the xanthi subgroup (unique in having enlarged postfemorals) of the Asiatic members of Taylor's fascia- tus group, we regard it unwise to restrict the type locality of E. capita; clearly the published designation of eastern United States is in error, and China probably em- braces the lizard's actual origin, but the final fixation remains in the hands of future workers. In like fashion we are reluctant to sug- gest that the name Eumeces capita be sup- pressed in order to preserve the name Eumeces xanthi despite the facts that (1) E. xanthi has been used for 85 years whereas (2) E. capita has never been used, except for its types, since it was pro- posed 95 years ago. Even E. xanthi was not clearly fixed with a recognizable spe- cies until 1936, however, when Taylor demonstrated that it is the same as the more familiar (even though more recently described) Eumeces pekinensis Stejneger (1924). Present decisions are not limited an}^ longer by a rigid (and ambiguous) namen ablitum rule. That rule is replaced by this statement: "A zoologist who con- siders that the application of the law of Priorit}' would in his judgment disturb stability or universality or cause confusion is to maintain existing usage and must re- fer the case to the Commission for a de- cision under the Plenary Powers." It is to be noted that the requirement to justify suspension of the Law of Priority — use of xanthi by at least five • different authors in at least 10 publications after Taylor's 1936 fixation— would be difficult to meet (see ICZN, 1972: 185-186). Convinced that stability of nomenclature is not a fac- tor to be considered in the present context, we recommend that E. xanthi be replaced as a species name b}'^ its senior synonym E. capita, recognizing that it is quite likely that the name E. xanthi may well be util- ized in the future for a subspecies of E. capita. Since E. capita is the earliest name applied to any member of the subgroup characterized by enlarged postfemorals, we Fig. 4. Posterior view of hind leg of the holo- type of Eumeces capita Bocourt, showing the patch of enlarged scales on thigh. 112 GREAT BASIN NATURALIST Vol. 35, No. 1 suggest that it be designated the capito subgroup. Our conclusion not to recommend sup- pression is reinforced to a certain extent by realization that the failure of recogni- tion of the identit}' of E. capito for almost a hundred years is not to be attributed wholh' to Bocourt, for his description is exemplary in detail and illustration (even though the critical postfemoral scale char- acter was not noted) and appeared in a widely known work. The erroneous lo- cality was, of course, the misleading fac- tor, but certainly not a unique one; many species have been properly allocated des- pite totally misleading type localities. A succession of extraordinary circmnstances that prevented subsequent workers from reexamining the holotype is responsible for the name's long histor}' as a nomcn duhium. Had the name been proposed with a totally inadequate description or in a very obscure outlet onl}- recently dis- covered, it might be construed as appro- priate that it remain in oblivion; but un- der the circumstances as they actually exist, it is fully appropriate that Bocourt's name be given its impartial place in no- menclature, subject only to the automatic provisions of the Code. The present de- ficienc}^ of an effective tjpe locality can readily be remedied at the appropriate time, when an arbitrary designation can be proposed in conformance with detailed knowledge of geographic variation that is now lacking. Literature Cited Bocourt, Marie-Firmin. 1879. Etudes sur les reptiles. Miss. Sci. Mexique, Rech. ZooL. Livr. 6:361-440, pis. 21-22, 22A-22D. BouRRET. R. 1937. Notes lienDetologiques sur rindochine francaise. Bull. Gen. Inst. Publ. 9:1-26, figs. 1-5. pi. Doi, H., .AND T. K.AMITA. A new species of Eumeces from West Corea. Zool. Mag., Tokyo 49:211-215, ill. GiJNTHER, A. 1889. Third contribution to our knowledge of reptiles and fishes from the Upper Yangtsze-Kiang. Ann. Mag. Nat. Hist. (6)4:218-229. ICZN (International Commission on Zoological Nomenclature). 1972. Seventeenth Interna- tional Congress of Zoology. Monaco, 24-30 September. 1972. Bull. Zool. Nomencl., 29: 168-189. Stejneger, L. 1924. Herpetological novelties from China. Occ. Pap. Boston Soc. Nat. Hist. 5:119-121. . 1925. Chinese amphibians and reptiles in the United States National Museum. Proc. U.S. Nat. Mus. 66:1-115, figs._ 1-4. Taylor, E. H. 1936. A taxonomic study of the cosmopolitan scincoid lizards of the genus Eumeces with an account of the distribution and relationships of its species. Kansas Univ. Sci. Bull. 23:1-643, figs. 1-84, pis. 1-43. STUDIES IN NEARCTIC DESERT SAND DUNE ORTHOPTERA. PART XV. EREMOGEOGRAPHY OF SPANIACRIS WITH BIOECOLOGICAL NOTES Ernest R. Tinkham^ Abstract. — Four decades of the author's records indicate that Spaniacris deserticola (Bruner) is confined within the periphery of the Colorado Desert. It is usually found, near or within a few hundred feet of sea level, marking the shore line of ancient Lake Cahuilla (e.xcept for the Dale Lake record). The preferred host plant is Coldenia palmeri growing on the lower fringes of bajadas, with C. plicata on drift sand being second in preference. Spaniacris can tolerate sand and rock tem- peratures of 60 C. (believed to be a maximum for Colorado Desert life). Mating takes place at that and lower temperatures. When they are disturbed while on the tops of host plants, their flight is low and direct and of short duration, and they come to rest on the torrid soil for long periods of time. The female, much larger than the male, can sustain the male in flight while mating. The study verified spatial longevity of Spaniacris at Indio, California, after approximately 70 years and for the Kane Springs area after 52 years. Spaniacris deserticola (Bruner, 1906), one of the rarest of Nearctic Desert grass- hoppers, was based on a single female collected at Indio by H.F. Wickham. In 1931, Morgan Hebard (1937:376) instructed me to make special efforts to locate S. deserticola. Late in the after- noon of 25 August 1931, a large colony was located on a low sand ridge some ten miles east of Coyote Wells and about a mile or so east of Plaster City, Cali- fornia. A torrid day had been spent hunting for the elusive creature on the blistering mesas. One had to blink con- stantly to keep one's eyes somewhat moist, and it was so hot that the ubiquitous grass- hopper Trimerotropis p. pallidipennis flew from the top of one creosote bush to another. At 8:00 that night, after sun- down, it was still 122 F in El Centro. Hebard records that I collected 13 males and 7 females for him that day and that my brother collected 12 males and 9 fe- males for me. In 1940 the colony was still there, and on 14 August I took 8 males and 10 females. In May 1961 I could not find any trace of the colony. During those intervening years the High- way Department had made a barrow pit out of the sand ridge and had destroyed the habitat. During the period 1949-1973 I took 2 males and 1 female in early June 1953 at or near the motith of Palm Canyon at a location northward and across High- way 111 from the Smoke Tree Ranch (both locations had disappeared by 1972 due to residential developments) and P. H. Timberlake took a female here on 24 June 1952 and a male on 21 June 1953. On 22 May 1954, Dr. John Goodman and I were collecting on the east side of dry Dale Lake, 25 miles east of Twenty Nine Palms and found a female nymph, probably in the last stadium, on barren playa, where sand drifted across the grav- el road. In May 1973 inspection showed this area so blasted and eroded by vio- lent sand storms, that the only surviving vegetation was some ancient, dying creo- sotes with their crowns supported, like mangroves, on long exposed roots. It is believed that this colony has been ex- terminated. It was the only one at a considerable elevation, as all others lie close to sea level. On 14 June 1964, I examined a rather level sweep of sand (air temperature 119 F) well covered with sand mat {Coldenia plicata) some miles west of Rice, California, and one several miles east of the turnoff to the Iron Mountain Pimiping Station. The first plant ex- amined contained a female Spaniacris (juietly resting ainong the leaves not more than several inches above the torrid sands. On 21 June 1964, along the old high- way about 4 miles west of Thousand Palms, I found a single male. On 29 July 1974 I reexamined this area with sand mat margining the south edge of the road and scattered on the sand but could find no Spaniacris, although that week I had located six new colonies some miles east of Thousand Palms. On 29 June 1970 Jim Davis took a pair of Spaniacris in Thousand Palms Canyon. The next day I found Spania- cris at that locality confined to Coldenia palmeri bushes bordering several hun- m-H\ Date Palm Avenue, Indio, California 02201. 113 114 GREAT BASIN NATURALIST Vol. 35, No. 1 dreds of feet of an old gravel road near the base of the gravelly hills. I took 2 females and one male. Associated were: Anconia integra on Atriplex canescens var. linearis and A. polycarpa; Xeracris minimus on sandpaper weed (Petalonyx thurbcri) ; Ciholacris parviceps on the road and Tytthotyle maculata, the Mal- pais lubber, nearby among the boulders of the brief bajada at the foot of the hills. One male was taken there 1 July 1971, and one male and one female on 3 July 1972; there were no Spaniacris there in 1973, a very dry year. In 1974 this area was designated Colony No. 6. (I shall re- port on Colony No. 6 later.) On the hot evening of 11 June 1973, Jim Davis and I made a night collecting trip along the roads to the Borrego Desert. One female Spaniacris was taken on the road 6 miles west of Salton City, just east of the radar tower; one pair was taken one mile west of the tower; and one crushed female was taken two miles west. In the arroyo area of a broad Pleistocene valley, about 13.5 miles southeast of Bor- rego Springs by road and within half a mile of Highway 78, another female was found on the paved road at night. In the late evening of 10 Aug. 1974, I surveyed the adjacent arroyo area, both sides of the road, but found no evidence of Spania- cris. The Borrego area, based on the parched conditions and poor showing of flowers, apparently got little of the day- long drizzle that drenched Coachella Valley on 7 Jan. 1974. Theodore J. Cohn contributed some of his collection records as follows: "Imperial Co., Ca., 3 mi. E Plaster City, at the Oy- ster Beds Turnoff, 25 June 1965, T. J. Cohn, 1 male, 2 females. On ground near road around noon, hot as a firecracker. I have stopped there many times since and have never seen them again. Lots of Anconia around in other years. "Sonora, Mexico, 22.3 mi. SE San Luis Rio Colorado, 20 June 1965, T. J. Cohn, No. 25. Base of sand dunes but not in them. 1 juv. female, last instar. Xeracris and Coniana also found here. These are the first set of dunes east of San Luis. "Sonora, Mexico, 76 mi. SE San Luis Rio Colorado (14 mi NW Los Vidrios). 14 June 1966, T. J. Cohn, No. 16. Very sparse vegetation on moderately sandy soil, not far from stabilized dunes. This was around a corral a few hundred yards south of the road, near the next set of dunes, 1 female. Lots of what I think is Coniana here." The sand dunes 22.3 miles SE San Luis are in large part in Arizona; the Inter- national fence terminating on some rocky hills astride the International Boundary. The best dunes are in Arizona immed- iately north of these hills. I have collec- ted there on many occasions but have never found evidence of Spaniacris. This should be the best location to discover Spaniacris in Arizona. At the 76-mile location, a high ridge of aeolian dunes sweeps far inland into the area from the southwestern coastal regions. Notes on the 1974 Survey Colony No. 1. Found, 25 July 3 miles north of Indio, California, on Monroe Street on C. palmeri on the north side of the dyke protecting the Coachella Valley branch of the All American Canal within fifty feet of the paved road. The Colony was at the southern edge of a con- siderable bajada that gently slopes down from the Indio Hills over an area of many miles. This location is directly south of the Curtis Desert Palms Oasis, whose ex- istence is threatened by the huge Massey Rock and Sand gravel pit. The plant life is typical of that described fully under No. 2 below but is not as extensive. Thus, the presence of Spaniacris deserticola for Indio was verified almost 70 years after Bruner described it in 1906; I have not been able to find the date of Wickham's collections prior to 1906. Colony No. 2. That afternoon, at the powerline crossing of Washington Street, 4 miles north of Interstate 10 and about 10 miles northwest of Indio, I found a much larger colony, here designated No. 2. The location was similar to that of No. 1, representing the southern edge of the long bajada margining the south side of the Indio Hills. The desert vegetation consists of shrubs, living ephemerals, and the dead skeletons of spring ephemerals. The desert shrubbery was composed of many widely scattered Coldenia palmeri growing along the narrow, poorly paved powerline road as well as along the mar- gins of a shallow dry- wash with fewer scattered clumps of creosote {Larrea di- varicata), sandpaper weed (Petalonyx thurheri), Burrobush (Franseria dumosa), March 1975 TINKIIAM: DESERT GRASSHOPPER 115 ' union of the placentae. Four ovules are commonly produced; however, sometimes one is reduced in size or, less frequently, lacking altogether. This condition ajipears only si)oradically and is probably influenced by environ- mental and nutritional factors. Calyx: The calyx is five-parted to the base, or nearly so. The lobes vary in size and shape from species to species. There is consistent variation in flowering and fruiting calyces with those in fruit being larger and sometimes scarious, as in P. scariosa. Seeds The seeds are geminate, elliptic to ob- long and ovoid, and generally cymbiform in shape. Size, shape, and surface mark- ings are diagnostically important. The seeds are unique in having the ventral surface excavated on one or both sides of a prominent ridge. However, the seeds of P. bakeri have the dorsal surface flat, with only a faint longitudinal groove down the center. On the ventral surface, the raphe is elevated above the normally excavated portions, thus giving the seed a triangular shape in cross section. In other taxa, the dorsal surface may be transversely ridged, as in P. arizonica, 130 GREAT BASIN NATURALIST Vol. 35, No. 2 P. palmeri, and P. popei, and reticulate to scabrous in P. congesta and P. rupestris. P. howelliana, P. serrata, and P. utahen- sis haAe the dorsal surface smooth and shiny ^^^th faint alveolations, while the seeds of P. pediccllata are tuberulate. The remaining species are alveolate (pitted). The ridge is corrugated on one side in over half the taxa, while the remainder lack corrugations. The seed margins can be entire as in P. alba, P. denticulata, and others, or corrugated along part or all of the marginal edge. Corrugated mar- gins are well represented in seeds of P. bombycina, P. coerulea, and P. corrugata. Color variations occur, but brown pre- dominates as in P. neomexicana and P. pediccllata. In such taxa as P. constancei, P. pallida, and P. palmeri the seeds are black, while in P. bombycina, P. coerulea, and P. formosula, they are dark brown. Reddish or reddish browTi seeds are typ- ical of P. glandulosa, P. rafaelensis, and P. utahensis. The smallest seeds occur in P. coulteri (1.6 mm long), and the largest known are in P. denticulata (4 mm long) . The light- colored, glutinous thickened band spoken of by Voss (1937) is a feature that de- velops during the ontogeny of the seeds. When immature, the seeds are either dark and turn light in color through a mottling pattern, or they are light and become dark when mature. The descriptions of seeds in this treatment are based on ma- ture examples. They are considered the most important single feature in delin- eating taxa, and on the basis of the size, shape, and surface characters, several dis- tinct groups can be arranged in an appar- ent phylogonetic order. These groupings are supported by other morphological fea- tures as well, namel}' leaf shape, pub- escence, corolla shape and color, and du- ration of the plant. The branching pat- tern of the stem and inflorescence, the type of calyx segments, and the stamens and style, whether included or exserted, are also useful features. Vesture Consi(lerabl(> confusion exists as to the terminology used in describing the ves- ture of plants. Ihe terms employed here- in are defined in Ai)pen(Hx II to lend uni- formity to their interpretation. The pub- escence often consists of two or more types of intermixed hairs. There are two main types of trichomes, each exhibiting vari- ation in size: (1) simple, unicellular tri- chomes, which vary in length and rigidity and may be erect, straight, or appressed; and (2) a stipitate-glandular type, which is usually multicellular. The stalk in the stipitate-glandular type varies in length and in number of cells. Sometimes the gland is sessile, or nearly so, and the stalk is often flattened. Phylogeny Constance (1963) indicates that the family appears to be a collection of mor- phological and geographical odds and ends, held together by floral and capsular features. He states, "I am not prepared to offer a complete system for Phacelia." However, Constance (1963) appears to have arrived at the most natural grouping of the genus Phacclia by recognizing three subgenera, Cosmanthus, Howellanthus, and Phacelia. The latter is the largest and most complex of the three and has been subdivided by Constance (I.e.) into the following species-groups: Crenulatae, Eu- glypta, Gymnobythus, Miltitzia, Pulchel- lae, Tanacetifoliae, and Whitlavia. The Crenulatae group, revised by Voss (1937), was the most complete study of the group ])rior to the ])resent work. Gillett (1960b) indicates. "The current infrageneric clas- sification of PJuwclia is generally consid- ered to be inadequate . . . and that con- siderably more evidence must be accumu- lated before the various species groups can be accorded classification that properly re- lates them to each other." The author agrees that natural generic and infra- generic relationships cannot be ])roposed imtil additional morphological, distribu- tional, and cytological data have been ac- cumulated. Howe^■er, he is ]irepared to offer a tentative phyogenetic sunmiary of relationships within the Crenulatae grouj). These data are subject to change as ad- ditional research may warrant. Those species occurring from Mexico to South America present a problem in th(> formulation of a complete phylogen- etic scheme. These southern taxa a])pear to !)(> the most jirimitiAO and are certainly file least understood of all the Cremdatae grou]). Most are known only from the t^■pe collections. Il wo"uld seem likely that the Cremdatae group has been derived June 1975 AT^VOOD: I'HACELIA CRENULATAF. CiROUP 131 congesta crenulata welshi neomexicana Fig. 1 A phylogenetic arrangement of the complexes in the Crenulatae group. from some form of Phacelia, past or pres- ent, somewhere in Mexico or South Amer- ica. The modern sj^ecies suggest several avenues of migration from Mexico, which have contributed to the ])resent diversity in morphology and distribution. Further- more, the morphological, cytological, and distributional relationships of the subgen- eric and sectional groups of Phacelia sug- gest either a polyphyletic origin or, if a monophyletic one, then a derivation pos- sessing several major lines of development. Those main lines of development occur- ring within the Crenulatae group are out- lined in Figures 1-7. The species are grouped together and arranged on the basis of similar morphological features and distribution. The following discussion is given to indicate which characters are considered to be advanced or primitive in this group. Seeds provide the most important char- acters in differentiating entities. The most primitive species, which occur in Mexico, all have small seeds, which suggests that large seeds are probably a derived feature. This character seems to follow a south-to- north trend with the largest seeds occur- ring to the north. There have been several avenues of specialization with the primi- tive seeds having more surface markings and being thicker and narrower. The seeds of most taxa, except in P. bakeri and P. argillacea, uniformly have the ventral surface excavated on both sides of a prom- inent ridge. These latter entities have the raphe elevated above the usually exca- vated portions and would appear to have diverged from the more typical form. Seeds with a corrugated ridge appear to be primitive, and those with pitted (alveo- late) and entire margins appear to be ad- vanced. Light brown seeds are apparently primitive, and dark brown, black, and reel- dish types are apparently derived. Retic- ulate, transversely ridged, and smooth-sur- faced seeds are also probably derived. The corolla has developed along three basic lines. Primitive plants are those having blue to purple colored campan- ulate corollas and exserted stamens and styles. However, some of the less advanced species have small, pale, campanulate corollas with included stamens and styles. Thirdly, pale to white tubular corollas are present in the more advanced entities. These advanced forms have less attractive flowers and long exserted stamens and styles. The corolla lobes have developed along two major lines, with the P. neo- mexicana complex having denticulate or erose margins and the remainder having entire margins. The former feature is bakeri glandulosa ^ argillacea formosub denticulata coulter Fig. 2. A phylogenetic arrangement of the species in the neomexicana complex. 132 GREAT BASIN NATURALIST Vol. 35, No. 2 splendens rafaelensis utahensis Fig. 3. A phylogenetic arrangement of the species in the welshii complex. probably advanced, entire margins being primitive. The scarious calyx segments of P. scar- iosa and P. pedicellata seem to be an ad- vanced feature. This is supported by the fact that the calyx lobes are persistent in fruit and probably aid in dispersal. Small, narrow calyx segments are considered to be primitive. palMeri The spicate thyrsus type of inflores- cence of the perennial, biennial, and some robust annual species is apparentl}" ad- vanced, while the variously branched sys- tems developed in most annual and some biennial taxa appear to be primitive. The primitive taxa do not follow the generalization that the perennial habit is more primiti^ e than the biennial or an- nual type. The majority of entities are ro- bust annuals, probably an inherited fea- ture; biennial and perennial types ap- pear to be derived. The primitive taxa integrifolia howel liana onelsonii intePMedia coerulea boMbycina constcncei robusta pallida Fig. 4. A phylogenetic arrangement of the species in the palmeri complex. Fig. 5. A phylogenetic arrangement of the species in the crenulata complex. possess an erect, usually branched, stout stem with a compound scorpioid inflores- cence. Some advanced biennial and pe- reimial entities have become specialized in the development of a spicate thyrsus in- florescence. Entire or subentire leaves are appar- ently derived from compound leaves. The narrowly revolute type exhibited by P. constancei is considered to be a special- ization, while the basal rosette common in the biennials and robust annuals appears to be a feature that has been retained during the phylogenetic development of leaves. Members of the Crenulatae group are probably monophyletic and have devel- oped along six major lines. These are June 1975 AT WOOD: PHACELIA CRENULATAE GROUP 133 rupestris congesta \7 infundibuliforMis Fig. 6. A phylogeiietic arrangement of the species in the congesta complex. treated as complexes but are not accorded taxonomic status. The hypothetical ancestor (s) of the Crenulatae group were apparently robust, densely glandular annuals with a branched stem, compound leaves and in- florescences, nonscarious sepals, and broadly campanulate blue or purple cor- ollas. The seeds were light brown, small, excavated on both sides of the corrugated ridge, with thick entire margins, cymbi- form, and elliptic to oblong in shape. Bi- ennial and peremiial types developed later in the phylogeny of the group. The prim- itive members of the neoniexicana complex are apparenth" the most primitive and are considered to be closest to the ancestral forms. The palmeri complex arose some- what later and extended more to the west of the neomexicana complex in its migra- tion northward. The congesta complex had its origin somewhere in north central Mexico and ])ossibly arose from the multi- ovulate P. infundibuliformis or some sim- ilar form. The crenulatae and scariosa complexes probably arose from taxa now extant in western Mexico. These six complexes are discussed and outlined in Figures 1-7. pattern of this complex has been north- ward out of Mexico through New Mexico to Wyoming and Montana. P. popei and P. arizonica have developed from a com- mon ancestor, as indicated by their sim- ilarity in seed, pubescence, and vegetative features. The same is true of P. fonnosula^ P. glandulnsa, P. bakeri, and P. argillaceci; however, the latter two have become spe- cialized in the development of noncorru- gated seeds. This is the only specializa- tion away from the typically excavated seed type present in the rest of the Cren- ulatae group. Welshii complex This complex is characterized by the large reddish seeds, showy corollas, and generally long exserted stamens and style. All taxa are narrowly restricted endemics occurring in Utah, Arizona, and western Colorado. P. welshii is considered to be the most primitive on the basis of its smaller, somewhat brownish seeds and branched habit. P. utahensis and P. splen- dens probably had a common ancestor but have adapted to different edaphic situa- tions and have therefore been isolated and selected out; judged on its robust bran- ching habit and glandular pubescence, the former is probably more primitive. P. rafaelensis is related to P. utahensis and may have been derived from it. P. serrata is the most advanced species as characterized by its smaller, lighter-col- ored corollas, and shortly exserted sta- mens and styles. Palmeri complex P. vossii and P. pallida are the most primitive and, along with P. robusta, are restricted to the south central part of the U.S. and adjacent Mexico. The remaining two species, occurring in Utah, Arizona, Neomexicana complex This complex is characterized by non- corrugated seeds, densely glandular pub- escence, light brown seeds (except in P. glandulosa), compound leaves, and branched habit. P. coulteri is closest to the ancestral species. P. alba, P. denticulata, and P. neomexicana are related to P. coulteri but possess the advanced features of small, white to pale-colored corollas, and less robust habit. The migrational pedicel lata Fig. 7. A phylogenetic arrangement of the species in the scariosa complex. 134 GREAT BASIN NATURALIST Vol. 35, No. 2 and Nevada, are disjunct from the others. The species are distinguished by their pale tubular corollas, small black seeds, and perennial or biennial habit. P. robusta is related to P. pallida but is considered to be advanced on the basis of its larger, reddish seeds. P. palmeri pos- sesses the advanced features of less-divided leaves and thin-margined seeds. Crenulata complex Although not lacking in a glandular pubescence, this complex displays more divergence from this primitive feature than does any other complex. There ap- pears to be a bilateral development, with P. crenulata and its relatives becoming specialized with a mixed pubescence of long stipitate glands and short hairs and dark brown seeds. P. anelsonii and P. coerulea have small corollas with included stamens and style. The former is more ad- vanced and possesses a thyrsoid inflores- cence and more or less scarious sepals. The other line, of which P. corrugata is the primitive taxon, is characterized by light brown or dark brown seeds, short stipitate glands, and yellowish stems. P. hoivellicma and P. integrifolia are the most highly developed species in this line, the latter apparently being the most ad- vanced on the basis of its lavender corollas and large, noncorrugated seeds. The former has large, dark brown seeds and bicolored corollas. The entire complex, with the exception of P. intergrifolia, po- ssesses distinctly corrugated seeds. Congesta complex This complex is related to and has pos- sibly been derived from /-•. injwidihuli- formis or some form close to it. P. infuii- dibuliforniis cHffers from other taxa in this complex only in its multiovulate, nar- rowly oblong capsule and overall vege- tative appearance. The small white cor- ollas, barely exserted stamens an(] style, and perennial habit of P. rupestris in- dicate that it is the most advanced species in this complex. The species of the con- gesta complex occur in the cast conlral part of the range of the Crenulatae gronjx Scariosa complex This most advanced complex is char- acterized by distinctly scarious sepals, bi- colored corollas, and large, corrugated. transversely ridged seeds. The species oc- curs in the southwesternmost part of the range of the grouj). Distribution and Ecology Members of the Crenulatae group oc- cur mostly in western North America. The remaining species, P. boliviano Brand, P. pinnatifida Griseb. ex Wedd., and possibly others, occur in Peru, Bolivia, and Argentina. The center of distribution in North America, based on the greatest concentration of taxa, is Arizona and New Mexico. In general, members of this group are desert plants that occur from near sea level to 5,000 feet elevation (up to 11,000 feet) . Some are restricted to a particular geologic formation, such as P. utahensis, which grows only on the Arapian Shale formation. P. baker i is restricted to mon- tane or subalpine regions on talus or al- pine slopes in Colorado, whereas P. integ- rifolia occurs mostly in deep sand. P. splendens is endemic to gypsiferous soil in western Colorado and northwestern New Mexico. The majority of taxa occur in the Lower Sonoran zone and are restricted to an isolated mountain range or valley. P. corrugata, P. denticulata, P. glandulosa, and others occur in the Upper Sonoran zone and generally have a wider distri- bution. P. denticulata is limited by the continental divide, occurring only on its eastern side. P. congesta, P. pedicellata, P. arizonica, P. denticulata, and P. rupe- stris are able to survive in the shade of overhanging ledges or as an understory of trees and shrubs. An important isolating mechanism that helps to account for the wide distri- bution is the seasonal ^ariati()n in phe- nology. The palmeri (omplex flowers in late sunnner and fall, whereas members of the welshii com])lex flower in spring and early summer. The foetid odor of some species is known to attract beetles, while bees and other insects are imj:)ortant pollinating agents in other species. The light, cymbiform seeds are prob- ably wind-dispersed. Tn addition to wind, birds are probabh' an important dispers- ing agent. (Cytology Cave aufi Constance (1042. 1944. 1947. 1950. 1959) and Constance (1963) have June 1975 atwood: phacelia crknulatae (;roup 135 made chromosome comits on about half alternate, entire to bipinnate, sometimes the Crenulatae group, all of which are revolute, sessile to long petiolate; in- 11=11. The uncounted members are florescence of terminal, axillary, or thyr- mostly narrowly restricted endemic soid, compound, scorpioid cymes; calyx plants and include the following: P. and- divided nearly to the base, elliptic to lin- sonii. P. hakeri. P. boUviana. P. bomby- ear, oblanceolate or spatulate, variously cina, P. constancei, P. coulteri, P. formo- jnibescent and sometimes accrescent; co- sula, P. glandulosa, P. howcUiana, P. in- rolla white or lavender to blue, tubular, frgrifolia, ^ ar. tcxana, P. intermedia. P. campanulate to rotate-campanulate, a pair pallida. P. serrata. P. utahensis, P. vossii. of variously shaped scales attached to the and P. welshii. base of each filament, these partially free from or completely attached to the tube, filament, or adjacent scale; stamens ex- Taxonomic Treatment ^^^^^j ^^ included within the tube, and Phacelia Kiis. Gen. 129. 1789. inserted at the base of the corolla tube; Subgenus Phacelia Constance. Britt. 15:278. Style exserted or included withm the tube, 1963. bifid 1/2-3/4 its length, capsule nearly Section Phacelia Brand. Das Pflanzenreich bilocular by union of the placentae, OVoid IV. 251:72. 1913 jq subglobose, variously pubescent and Group Crenulatae Constance. Britt. 15:279. niostly glandular; mature seeds 4 (1, 2 or 4 in P. amabilis and P. congesta)^ Annual, biennial, or j:)erennial herbs light brown to black, favose, reticulate, from a taproot; stems simple to much entire to corrugated or tranversely ridged, branched, erect, ascending or prostrate, excavated on both sides of a prominent leafy, puberulent to hispid, strigose or ridge (except in P. bakeri) and mostly \ariously glandular; leaves jirevailingly cymbiform. Key to the species of the Crenulatae group la. Stamens and style included or nearly so ----- 2 lb. Stamens and style exserted 2 mm or more - - 4 2a. Corolla tubular, light blue, lobes denticulate; plants of Colorado and Wyoming 15. P. denticulata 2b. Corolla campanulate or rotate-campanulate, lobes entire or at most crenulate - — 3 3a. Plants brittle, breaking easily; corolla 3-4 mm long, pale mauve to light blue; mature seeds dark brown 9. P. coerulea 3b. Plants not brittle; corolla ca. 6 mm long, lavender or white; seeds brown -- 4. P. anelsonii 4a. Pedicels shorter than the calyx; sepals not scarious in fruit 6 4b. Pedicels filiform (at least as long as the calyx lobes); sepals scarious in fruit; leaves pinnately compound, the divisions broad 5 5a. Sepals less than 3 times longer than broad; mature seeds 2.5 mm long or less; plants of lower Baja California and southwestern Sonora, Mexico - - - 30. P. scariosa 5b. Sepals 3 or more times longer than broad; mature seeds 2.5 mm long or more; plants of central Baja California north to California, Ari- zona, and Nevada 25. P. pedicellata 6a. Corolla over 4 mm long, white or variously colored 13 6b. Corolla small (4 mm long or less), white, blue, or lavender 7 7a. Plants prostrate, diffusely branched (at the base); mature seeds 1.8-1.9 mm long, ovate, transversely ridged; corolla white - 6. P. arizonica 136 GREAT BASIN NATURALIST Vol. 35, No. 2 7b. Plants erect, mature seeds mostly over 2 mm long, if smaller then not with the above combination of characters 8 8a. Corolla lobes entire; mature seeds with the ridge corrugated or the dor- sal surface reticulate 11 8b. Corolla lobes erose or denticulate; mature seeds pitted, margins and ridge entire 9 9a. Corolla white or pale colored, 3-4 mm long 1. P. alba 9b. Corolla blue or purple, 4-5 mm long 10 10a. Stems thick, robust; corolla 4-5 mm long, bluish purple; mature seeds 1.6-1.9 mm long; plants endemic to the states of Hidalgo and Zacatecas, Mexico 13. P. coulteri 10b. Stems weak; corolla 4 mm long, blue; mature seeds 3.2-3.3 mm long; plants of Arizona and New Mexico 22. P. neomexicana 11a. Corolla white; mature seeds 2.1-2.7 mm long, the ridge not corrugated, dorsal surface reticulate; plants from southeastern Arizona eastward 29. P. rupestris lib. Corolla blue to light violet; seeds 2.5-3.2 mm long, the ridge corrugated _._._ 12 12a. Corolla bicolored (tube white, lobes blue to lavender), campanulate; seeds ovate, ridge corrugated, margins entire; plants from western Ar- izona westward 3b. P. ambigua var. minutiflora 12b. Corolla not bicolored, light blue to lavender, rotate; seeds elliptic to ob- long, ridge and margins corrugated; endemic to Coconino County, Arizona 31. P. serrata 13a. Corolla distinctly tubular, white or pale colored 14 Hb. Corolla campanulate, purple, blue, lavender, or white (appearing tu- bular in some pressed specimens) -. 19 14a. Plants annual or biennial (possibly perennial in P. pallida) ; north of Nu- evo Leon, Mexico — -. 15 14b. Plants perennial; endemic to the state of Nuevo Leon, Mexico 34. P. vossii 15a. Seeds brown or reddish brown, 2.9 mm long or more 16 15b. Seeds black, 2.9 mm long or less 17 16a. Seeds 3.5-4 mm long; cauline leaves sessile (or nearly so), auric- ulate; plants of Utah -.. 27. P. rafaelensis 16b. Seeds 2.9-3.7 mm long; cauline leaves distinctly petiolate, not auric- ulate; plants of Kansas, Oklahoma, Texas, and Mexico 28. P. rolmsta 17a. Inflorescence thyrsoid; stems solitary or if branched then near the base ..^. 24. P. palmeri 17b. Inflorescence open; stems branched throngiiont, especially at base 18 18a. Leaves revolute, narrowly linear or lanceolate (less than 1.5 cm wide); plants of Utah and Arizona 11. P. constancei 18b. Leaves not revolute. broadly oblong or lanceolate (mostly over 1.5 cm wide); plants of Texas and adjiKciit Mfwico 23. P. pallida 19a. Leaves pinnately or bipinnately compound, fiiu^iy dissected 20 19b. Leaves simple or if compound not fiiu^l\' so. the divisions broad (over 5 mm wide) 25 June 1975 atwood: phacelia crenulatae group 137 20a. Corolla violet; plants endemic to Jackson County, Colorado 16. P. formosula 20b. Plants not as above 21 21a. Pubescence of the leaves mostly unicellular, j)uberulent to hispid; plants native from western and southern New Mexico, southward and east- ward (except in P. orgillacea which is endemic to Utah) 22 21b. Leaves mostl}' with multicellular, stipitate-glandular hairs; native from central New Mexico northward and westward 24 22a. Flowers violet to light blue; seeds more or less excavated on one side of the ventral ridge; ])lants endemic to the Green River Shale formation in Utah County, Utah 5. P. argillacea 22b. Flowers blue to purple; seeds excavated on both sides of the ventral ridge - 23 23a. Seeds 1.8 mm long, ovate; ultimate leaf divisions mostly less than 4 mm wide - 26. P. popei 23b. Seeds over 1.8 mm long, elliptic to oblong; ultimate leaf divisions mostly over 5 mm wide 10. P. congesta 24a. Seeds not excavated ventrally; plants east of the Continental Divide in Colorado and New Mexico 7. P. hakcri 24b. Seeds excavated ventrally; plants west of the Continental Divide in Wyoming, Montana, and Idaho 17. P. glandulosa 25a. Corolla not distinctly bicolored, blue, purple, or white 28 25b. Corolla distinctly bicolored, the tube white or yellow, the lobes blue 26 26a. Cauline leaves sessile, auriculate, plants robust, 0.8-5.8 dm tall, en- demic to Sanpete and Sevier counties, Utah 33. P. utahcnsis 26b. Cauline leaves distinctly petiolate; plants not especially robust, less than 2.7 dm tall, more eastern in distribution 27 27a. Stems branched at base; leaves simple, strigose and glandular; corolla tube white; seeds corrugated on the margins and ridge, dorsal sur- face smooth 18. P. howelliana 27b. Stems simple or branched above; leaves essentially glabrous, some of the lower usually compound; corolla tube yellowish; seeds essen- tially lacking corrugations, dorsal surface dee])ly j)itted 32. P. splendens 28a. Corolla white; plants endemic to Saline Valley, Inyo County, Califor- nia :: 2. P. amabilis 28b. Corolla pale blue, purple, or lavender (rarely white); plants wide- spread or if endemic not as above 29 Corolla lavender; seeds lacking ventral corrugations 19a. P. integrifoUa var. integrifolia Corolla pale blue to purple; seeds corrugated ventrally 30 Stamens and style exserted 4 mm or less; mature seeds 2.9 mm long or less 31 Stamens and style exserted over 4 mm; mature seeds over 2.0 mm long 32 31a. Mature seeds 2.2-2.5 mm long; dark brown; plants very brittle an- nuals 8. P. bombycina 31b. Mature seeds 2.7-2.9 mm long; brown; plants not especially brittle 20. P. intermedia 138 GREAT BASIN NATURALIST Vol. 35, No. 2 32a. Mature seeds corrugated only on the ridge; pubescence of the stems densely hispid, glandular above 3a. P. ambigua var. ambigua 32b. Seeds with the margins and ridge corrugated; pubescence of the stems mostly glandular, sometimes finely so 33 33a. Glandular pubescence of the stems and herbage mostly multicellular 34 33b. Glandular pubsecence of the stems and herbage stipitate but not multi- cellular 35 34a. Corolla lavender to purple, 4.5-6 mm long; anthers blue green; stems often reddish; mature seeds reddish brown, 2.4-3 mm long; plants of southeastern New Mexico and adjacent Texas 19b. integrifolia Torr. var. tcxana 34b. Corolla blue, 6 mm long or longer; anthers always yellow; stems green or yellow green; mature seeds light brown, 3.1-4 mm long; plants of northwestern New Mexico, Arizona, Colorado, and Utah 12. P. corrugata 35a. Leaves narrowly oblong, mostly less than 1 cm wide, glandular pub- escence of the leaves and lower part of the stems short stipitate, the nonglandular hairs mostly fine, retrorse .. 14a. P. crenulata var. angustifolia 35b. Leaves mostly well over 1 cm wide, glandular pubescence of the leaves and stems stipitate-multicellular, nonglandular hairs spreading 36 36a. Stems usually reddish at least below; mature seeds with a dark cen- ter dorsally and lighter margins; plants of western Utah and Ar- izona westward to California and Nevada 14b. P. crenulata I'orr. var. crenulata 36b. Stems green; mature seeds uniform in color dorsally; plants endemic to Coconino County, Arizona '. 37 37a. Corolla lavender to white, campanulate, 5-6 mm long; leaves un- dulate to regularl}' dentate, oblong to lanceolate; seeds uniformly brow2i; plants endemic to Kane and Garfield counties, Utah 21. P. mammillarensis 37b. Corolla dark blue, tubular to funnelform, 5-8 mm long; leaves irreg- ularly crenate to dentate, lanceolate; seeds brown ventrally and reddish dorsally; plants endemic to Coconino County, Arizona .... 35. P. ivelshii 1. Phacelia alba Rvdberg July 1899, Townsend and Barber, 129, (ny); -p-^. o ' Isotypes (cAs, mexu, pom, rm, uc, uc, us). ^" Phacelia glandulosa Nutt. ssp. eu-glandulosa Phacelia alba Rydb.. Bull. Torr. Bot. Club 28:30. Brand var. elatior Brand. Das Pflanzcnreich 1901. IV. 251:82-83. 1913. in part. Holotype: Wyo- Holotype: Colorado: Costilla Co.: Sangre de ining-' Albany Co.: Jolm, 12 August 1900, Christo Creek, 2 July 1900, Rydberg and A. Nelson. 8053. (gh, pom, rm). Vroeland, 5755, (ny?); Isotype (rm/). Paratypes: Colorado: Clear Creek County: Plants annual. 0.5-7 (hn tall; stems Valley of Upper Arkansas River, 1873. J. simple to much branched, erect or as- ™f«-„^9' (NY); head waters of Clear Creek, ,o,Kli,,o. loafy, puberulent, setose to hir- 1861. C. Parry, 314 (gh, ny); New Mexico: ^ ^ , ^. •. ■ ' , i i • n • Lincoln Co.: " Ruidoso Creek. White Moun ''^'^^ •'"^' stipitate-glandular, especially in tains, 1 July 1895, E. Wooton, s.n.. (ny). the ijiflorescence; leaves irregularly lobed Phacelia neomexicana Thurber ox Torr. var-. to bipiniiate, 2-10 (. ni wide, strigose tO alba (R.ydb.) Brand. Das Pflan/.eiueich IV. setose, slightly to moderately stipitate- 251:83. 1913. glandular, long petiolate below to sessile Phacelia neomexicana Thurl^er ex To'. . var. ;,j. ^ubsessile above; inflorescence of dense couUeri subvar. folisissima Brand. Das Pflan- . . , , - • . , zenreich IV. 251:84. 1913. Holotype: Mexico: terminal compound scorpioid cymes, state of Chihuahua: near Colonia Garcia. 13 densely glandular and puberulent to hir- June 1975 ATWOOD: PHACELIA CRENULATAK GROUP 139 sute, the cymes 1-2 cm long in flower to 8 cm long in fruit, pedicels 0.8-1 mm long; sepals linear to oblanceolate, 3.5-4 mm long, 0.5-1 mm wide, finely glandular and somewhat setose; corolla campanulate, white (sometimes pale purple), 3-4 mm long and broad, lobes pubescent and den- ticulate; capsule ovoid to subglobose, 3- 3.3 mm long, 2.5-2.9 mm wide, puber- ulent and quite glandular (the partition oblanceolate) ; mature seeds elliptic to ob- long, light to dark brown, 2.4-3 mm long, 1.4-1.5 mm wide, uniformly alveolate throughout and cymbiform, the ventral surface shallowly excavated on both sides of the ridge and lacking corrugations, the margins thick and entire (Fig. 9). Col- lections: 207 (5); representative: C. Parry 314, 1861 (gh, ny); D. Atwood 1975, 1962, 1963a (bry. ny. us); M. Jones 511 (pom). Habitat. — Dry clay-loam or sandy draws and flats, fields, meadows, and gravelly hillsides. From 6,000 to 9,500 feet. Growing on the short grass prairie, sagebrush belt, and pinyon-juniper com- munities at the lower elevations and as- sociated with aspen, spruce, fir, or pine forests at the higher elevations. Late May to early October. Distribution. — Laramie and Albany COS., Wyoming, southward through cen- tral Colorado, New Mexico, and adjacent Chihuahua, Mexico, westward to Gra- ham and Apache cos., Arizona, and Sevier, Wayne, Garfield, and Washington cos., Utah (Map 1). The material cited by Brand (1913) for P. glandulosa ssp. eu-glandulosa var. Fig. 8. Phacelia alba Rydberg. L. Higgins 2229 (bry). Fig. 9. Dorsal and ventral view of the seeds of P. alba Rydberg. D. Atwood 1975 (bry). 140 GREAT BASIN NATURALIST Vol. 35, No. 2 elatior Brand (Nelson 8053) belongs to P. alba. County, California, along Hunter Creek at an elevation of 1,800 feet. 2. Phacelia amabilis Constance Phacelia amabilis Constance. Madrono 7:56-59. 1943. Holotvpe: California: Inyo County: Saline Valley, 21 April 1942, A. Alexander and L. Kellogg 2681 (uc!); Isotype (uc). Plants annual; stems stout, branched above, stipitate-glandular, puberulent, and hispid; leaves petiolate, oblong to oblong- ovate, 8-15 cm long, 3-5 cm wide, pin- natifid, upper leaves reduced and less deepl}" divided; inflorescence of com- pound scorpioid cymes, the cymes 5-12 cm long, pedicels 2-3 mm long; sepals lanceolate, 3-5 mm long, 1-2 mm wide; corolla broadly campanulate, white, 7-8 mm long, 8-12 mm wide; stamens and style exserted 5 mm or more; capsule ovoid, 3-4 mm long, 2-3 mm wide; im- mature seeds apparently 2 or 4, 3-4 mm long, thin and pale, ventral surface ex- cavated on each side of the prominent ridge; collections: 1 (0); representative: A. Alexander and L. Kellogg 2681 (uc). Distribution and Habitat. — Appar- ently endemic to Saline Valley, Inyo 3. Phacelia amhigua Jones Plants annual, 0.2-5.7 dm tall; stems simple to much branched, hispid, pub- erulent, and stipitate-glandular; leaves simple to pinnately compound, petiolate to sessile above, the margins various, stri- gose to hispid and stipitate-glandular, 0.5- 13 cm long, 0.5-4.5 cm wide; inflores- cence of compound scorpioid cymes, the cymes elongating to 12 cm in fruit, pub- escence as for the stem; sepals elliptic to oblanceolate, 2.7-5.1 mm long, 1-1.3 mm wide, puberulent, hispid, and stipitate- glandular; corolla campanulate to rotate- campanulate, purple or dull lavender, 4-10 mm long and broad, pubescent; sta- mens and style exserted 2-10 mm; style bifid, pubescent below; capsule globose to subglobose, 3-3.5 mm long, 2.5-3.4 mm wide, puberulent and glandular; mature seeds 4, ovate, reddish to brown, 2.5-3.3 mm long, 1.5-1.8 mm wide, alveolate, cymbiform, the ventral surface excavated on both sides of the ridge, the ridge cor- rugated on one side. 1. Corolla 4 mm long or less; style 6.5 mm than 4 mm long 1. Corollas over 4 mm long; style 9 mm mm long Key to the varieties of P. amhigua long or less; calyx in fruit less var. minutiflora more; calyx in fruit 4 var. amhigua 3a. var. amhigua Phacelia ambigua Jones. Contr. West. Bot. 12:52. 1908. Holotype: California: San Bernardino Co.: Needles, 5 May 1884, M. Jones 3822. (pom!); duplicates (ny, rm, uc, us). Paratypes: Ari- zona: Coconino Co.; Hole in the Bock. 13 April 1894. M. Jones s.n.. (?): Nevada: Lin- coln Co.: Calienfe. 29 April 1904, M. Jones, s.n., (?). Phacelia crenulata Torr. in Wats. var. am- bigua (Jones) Macbride. Contr. Gray Herb. 49:25. 1917. Plants annual, 0.5-5.7 dm tall; stems simple to much branched, usually more leafy toward base; leaves 0.5-13 cm long, 0.5-4.5 cm wide, slrigose to hispid, us- ually only with scattered glands, reduced from the base upward; cymes elongating to 12 cm in fruit; sepals 3.2-5.1 mm long, 1-1.3 mm wide; corolla campanulate. purple to blue, 5-10 mm long and wide, pubescent; stamens and style exserted 9 mm or more; style bifid 2/3 its length, puberulent and glandular below; capsule 3.3-3.5 mm long, 3-14 mm wide, puber- ulent and stipitate-glandular; mature seeds 3.3 mm long, 1.5 mm wide. Col- lections: 250 (30); representative: E. Pal- mer 625 (ny); M. Jones 5018 (ny, pom, RM. uc. us); C. Pringle s.n. (cas, gh, ny); J. Howell 3504 (rsa); D. Atwood 2210, 2220a, 2294, 2296, 2303, 2310, 2319, 2353 (bry). Habitat. — Growing on a wide \ ariety of soils in the lower Sonoran Desert from 490 to 5,000 feet elevation. February to mid-June. Distribution. — vSouthern Nevada and south wostoni Utah in Washington Co., June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 141 N ^^" n 1 : -J ••• 1 L ^ • / t • 1 V 1 ) • / ? Map 1. Southern Wyoming, Utah, Colorado. Arizona, New Mexico, and Chihuahua, Mexico. Distribution of P. alba Rydberg. southward through Arizona (except for Navajo and Apache cos.,) and southeast- ern California (Map 2). Phacelia arnbigua has been treated as a variety of P. crenulata, which it closely resembles. However, the former species appears to be more uniform throughout its range than does the latter. Also, to treat P. arnbigua at infraspecific rank would require the inclusion of other closely re- lated taxa, namely P. bombycina W. & S. and P. amabilis Constance. In order to understand the complete relationships be- tween these taxa additional field and greenhouse studies are needed. These studies may demand nomenclature changes, but until such studies are carried out the present treatment provides a more uniform arrangement of the entities in- volved. 3b. var. minutiflora (Voss in Munz) Atwood comb. nov. Fig. 10 Phacelia minutiflora Voss in Munz, Man So. Calif. Bot. 409, 600. 1935. Holotype: California: Imperial Co.: 2 miles north of Cargo Muchacho Mountains. 5 April 1932, P. Munz and L. Hitchcock 12141 (pom!). Phacelia crenulata Torr. in Wats. var. minuti- flora (Voss) Jeps., Fl. Calif. 3:266. 1943. Plants annual, 0.2-4.5 dm tall; stems simple or variously branched, hispid, puberulent and stipitate-glandular (es- pecially in the inflorescence) ; leaves 0.5- 11 cm long, 0.5-3.5 cm wide, strigose to hispid and often quite glandular, the lower with longer petioles than the up- per reduced leaves, cymes elongating to 7 cm in fruit; sepals 2.7-3.8 mm long, 1 mm wide, corolla rotate-campanulate, tube white, lobes lavender to blue, pub- escent, 4 mm long and broad; stamens and style exserted 2 mm or less; style bifid 2/3 its length, puberulent and glandular below; capsule 3-3.2 mm long, 2.5-3 mm wide, puberulent and glandular; mature seeds 2.5-3.2 nun long, 1.3-1.8 mm wide (Fig. 11). Collections: 89 (4); repre- sentative: E. Palmer 626 (ny); T. Kear- ney and R. Peebles 10941, 10963, 11016 (us); D. Atwood 2320, 2355, 2352, 2341 (bry); I. Wiggins 9669 (rsa, uc). Habitat. — Sandy to rocky desert flats, washes and slopes from near sea level to 2,200 feet. Common in Larrea, Ambrosia. Atriplex, Fouguieria, and Ccrcidiuin com- munities. Late December to late April. Map 2. Utah, Nevada, Arizona, California, and adjacent Mexico. Distribution of P. ambigua Jones: O var. ambigua; var. minutiflora. # 142 GREAT BASIN NATURALIST Vol. 35, No. 2 .^ ^v( ;i^ \ m Fig. 10. Phacclia ambigua Jones yav. minutiflora (Voss in Muiiz) Atvvood. D. Atwood 2341 (bry). Distribution. — Southwestern Arizona in Maricopa, Pima, and Yuma cos., and westward to San Bernardino Co., Califor- nia, south into Baja California and Sonora, Mexico (Map 2). 1. Phacclia anclsonii Macbride Fig. 12 Phacelia anelsonii Macbride, Contr. Gray Herb. 49:26. 1917. Holotype: Nevada: Lincoln Co.: Meadow Val- ley Wash. 28 April 1902, L. Goodding 635 (rm!). Erect annual, 1-5.5 dm high; stems terete, usually simple covered \\dth brown- ish stipitate glands, leafy throughout; leaves narrowly to broadly oblong, pin- natcly cleft, 1.5-8 cm wide, pubescence brownish, stipitate-glandular, with a few j non-glandular hairs, jiedicels from 3 cm long on the lower part of the stem to nearly sessile on the upper part, the pin- June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 143 plant which Goodding, when he des- cribed his P. foetida . . . took to represent P. palmeri Wats., a very different plant with exserted stamens and corrugated Fig. 11. Dorsal and ventral view of the seeds of P. ambigua Jones var. minuiiflora (Voss in Munz) Atvvood. D. Atwood 2320 (bry). nae somewhat reduced toward the base of leaf, margins crenate; inflorescence racemose to paniculate, usually terminal on the upper half of the stem, some- times on leafy lateral branches, individual cymes 1-5 cm long, setose and glandular pubescent; corolla light violet or white, rotate-campanulate, 6 mm long and wdde; sepals oblanceolate to spatulate, 3-6 mm long, 1-2 mm wide, setose to glandular, 1-2 mm longer than the capsule; stamens included, anthers yellow; style included, 3.5-4.8 mm long, shorter than the stamens, cleft 2/3 its length, glandular and puber- ulent at the base; capsule oval, 3.3-3.7 mm long, glandular spotted throughout and pilose on the upper half; mature seeds 4, oblong, 2.7-3.4 mm long, 1-1.3 mm wide, light browai, margins entire, ventral surface strongly alveolate, divided by a prominent ridge, ridge corrugated along one side, dorsal surface aheolate (Fig. 13). Collections: 30 (1); representative: L. Goodding 635 (rm); R. Barneby 2937 (cAS, rsa) : S. Welsh, D. Atwoood, and E. Mathews 9542 (bry); L. Higgins 499 (bry). Habitat. — Commonly in shady places at the base of sandstone and limestone cliffs or among rocks and in sandy to grav- elly washes, 2,000 to 5,000 feet elevation. Usually locally scattered, April to May. Distribution. — Lincoln Co., Nevada, southward to Washington Co., Utah, Inyo and San Bernardino cos., California (Map 3). Macbride (1917) in his original des- cription of P. anelsonii savs, "this is the Fig. 12. Ripley and R. Phacelia anelsonii Macbride. H. Barneby 3496 (cas). 144 GREAT BASIN NATURALIST Vol. 35, No. 2 seeds." It is not known from whence Macbride drew this conclusion, but it supports the observations of the author that P. foetida is the same entity as P. palmeri. P. anelsonii is related to P. crenulata and P. amabilis as indicated by both leaves and seed characters. P. anelsonii is easily distinguished from them by the included stamens and style, a feature which it has in conunon with P. coerulea and P. denticulaia. Detailed observations of the type specimens, as well as other collections, revealed the presence of cor- rugations along one side of the ridge in some seeds of P. anelsonii. Both P. cren- ulata and P. amabilis have the corrugated ridge. Macbride (1917) and Voss (1937) have indicated that the seeds lack cor- rugations. John Thomas Howell was the first to report this taxon for Washington Co., Utah, and San Bernardino Co., California (1941), and for Inyo Co., Cahfornia (1942). Two collections from Washing- ton Co., Utah (R. Barneby 2937 and B. Wood 140), are more robust in habit and have unusually large parts. 5. Phacelia argillacea Atwood Phacelia argillacea Atwood. Phytologia 26(6): 437. 1973. Phacelia glandulosa Nutt. var. argillacea At- wood in Welsh & Moore, nomen. nudum. Holotype: Utah Co.: Spanish Fork Canj-on. Clear Creek ca. 6 mi west of Soldier Summit. 18 August 1971. D. Atwood et al. 3091 (bry); Isotypes, to be distributed; Pleasant Valley Junction (Colton). Wasatch Moun- N 1 • I ^^\^ 1 • • • • ^s^ k • • Fig. 13. Dorsal and ventral view of tlie seeds of P. anelsonii Macbride. R. Barneby 2937 (CAs). Map 3. Southwestern Utali, southern Ne- vada, and California. Distribution of P. anelsonii Macbride. tains, August 1883, M. E. Jones s.n. (us!); Isotypes (c.^s, cas, ny, ny, pom, uc, uc). Paratypes: Utah: Utah Co.: Clear Creek near Soldier Summit. 6 July 1894, M. E. Jones 5591 (ny, pom, uc). Plants annual, 1-3.6 dm tall; stems finely })ubescent; leaves oblong in outline, pinnatifid, 0.8-5 cm long, 0.5-1.5 cm wide, strigose, petiolate; inflorescence of com- pound scorpioid cymes, stipitate-glandular and setose to hirsute, pedicels 0.7-1 mm long, cymes elongating to 7.5 cm in fruit; sepals elliptical to oblanceolate, 2-3.8 mm long, 1 mm v\dde, stipitate-glandular and hirsute to setose; corolla campanulate. bluish violet, ca. 5 mm long and broad, lobes pubescent; stamens and style ex- serted en. 7 nmi; capsule subglobose, 3.2- 3.3 nun long, 2.3-2.4 mm wide, glandular and setose; mature seeds 4, brown, ovate to elliptic, 2.4 mm long, 1.1 mm wide, pitted, the ridge curved and more or less excavated on one side. Collections: 3 (1); representative: M. Jones s.n. (cas. NY, POM, RM. uc, us); M. Jones 5591 (ny, pom, uc); D. Atwood 3091 (bry). Habitat. — The species ap])arentl^ grows on gravelly hillsides of the Green- river formation between (i.5()0 and 7.000 feet, July to August. DisTiuBUTiON. — Known only from Utah Co., Utah. This species is related to both P. inland- iilosa and to P. bakeri but can be (Hstin- guished by its more nearly glabrous herb age, smaller capsule, flowers, and dil ferent seeds. June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 145 6. Phacelia arizonica A. Gray Fig. 14 Phacelia arizonica A. Gray, Syn. Fl. II, 1:394. 1878. Lectotype: Arizona: Maricopa Co.: plains of the upper Gila, 15 April 1880, E. Greene s.n. (gh!); duplicate (c^s). Phacelia popei Torr. & Gray var. arizonica (Gray) Voss, Bull. Torr. Bot. Club 64:94-95. 1937. Low diffusely branched (at the base), prostrate annual, 0.2-2.1 dm tall (up to 4 dm in extreme forms); stems slender, 1- many, hispid to villous and glandular, often reddish; leaves larger and denser at the base, oblong to linear in outline, sessile above to short petiolate below deeply lobed to pinnatifid, 1-5 cm long, 0.5-2 cm wide, densely strigose, inflores- cence terminal, cymes densely flowered, the flowers subsessile (pedicels to 0.5 mm long), densely covered with small gland- ular and short simple hairs, with some longer simple hairs intermixed; sepals oblong to elliptic, 2.5-4 mm long, 1 mm wide, hirsute; corolla campanulate, white or rose white, (often reddish in bud), 3-4 mm long and broad, lobes pubescent and shallowly erose; stamens and style ex- serted 3-4.5 mm; style bifid 3/4 its length, the lower 1/2 puberulent and sometimes finely glandular; mature seeds 4, ovate and more or less cymbiform, brown, 1.8- 1.9 mm long, 1.2-1.3 mm wide, alveolate, ventral surface excavated on both sides of the ridge, dorsal surface alveolate and transversely ridged (Fig. 15). Collections: 104 (2); representative: C. Pringle s.n. (ny); L. Goodding 1035 (ny, uc, us); M. Jones 28501, 28503 (pom); D. Atwood 2200a (bry, cas, ny, wts, b); L. Higgins 2814 (bry); D. Atwood 2186 (b, bry, cas, NY, wts). Habitat. — Common along roadsides, sandy flats, and gravelly hillsides, from 1,600 to 2,500 feet. Often growing with Prosopis. Quercus, Junipcrus, or grass com- munities, mid-February to late June. Distribution. — Arizona from Mari- copa Co. southward into Sonora, Mexico, and eastward to Sierra and I^una cos.. New Mexico (Map 4). This species was treated as a variety of P. popei T. and G. by Voss (1937), but is easily distinguished as a species on the basis of its smaller, white, glandless co- rolla, prostrate habit, less-dissected leaves. and nearly leafless stems. However, the two are similar in seed characters and in the much-branched habit. P. popei has erect to ascending and more rigid stems in contrast to the slender, prostrate stems of Phacelia arizonica. Gray (1878) did not designate a type specimen. Therefore, the author has chosen the collection of Greene, "on the plains of the upper Gila," as the lectotype. 7. Phacelia bakeri (Brand) Macbride Fig. 16 Phacelia bakeri (Brand) Macbride. Contr. Gray Herb. n. ser. 49:24. 1917. Phacelia crenulata Torr. e.x S. Wats. var. bakeri Brand. Das Pflanzenreich IV. 251:78. 1913. Holotype: Colorado: Ouray Co.: Ouray, 10 August 1901, C. F. Baker 758 (gh!); duplicates (ny. pom, rm, uc, us). Phacelia glandulosa Nutt. subsp. eu-glandu- losa Brand var. australis Brand, Das Pflan- zenreich IV. 251:82-83. 1913, in part. Lecto- type: Colorado: Conejos Co.: Cumbers. 7 Sep- tember 1899. C. F. Baker 549 (us!); dupli- cates (gh. ny. pom, rm, rm). Paratypes: Colorado: El Paso Co.: Manitou, 15 July 1903. C. Clements 47.1 (rm), in part; Hins- dale Co.: Lake Fork River, Lake City. July 1893. C. A. Purpus 618 (uc, uc). Annual, 0.5-4.8 dm tall; stems simple or branched, with multicellular stipitate glands, pilose to somewhat hirsute; leaves pinnately divided, the pinnae irregularly crenate to dentate, 2-8 cm long, 0.5-3 cm wide, reduced upwards, petioles 0.5-4 cm long, dorsal surface strigose, ventral sur- face strigose, glandidar and setose along • N • ••• • • % Map 4. Soutliern Arizona and adjacent So- nora, Mexico, and western New Mexico. Distribu- tion of P. arizonica Gray. 146 GREAT BASIN NATURALIST Vol. 35, No. 2 Fig. 14. Phacelia arizonica Gray. L. Higgins 2814 (bry). June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 147 Fig. 15. Dorsal and ventral view of the seeds of P. arizonica Gray. D. Atwood 2186 (bry). the veins; inflorescence of compound scorpioid c>TTies, terminal on the main stem and lateral branches, setose to puber- ulent and glandular, pedicels 1-2 mm long; sepals oblanceolate to narrowly spat- ulate, 1-1.5 mm longer than the capsule, setose to puberulent and glandular; co- rolla campanulate, violet to dark blue, 7-8 mm long, 5-7 mm wide, pubescent; stamens exserted 5-9 mm, anthers green- ish, filaments bluish; style exserted bifid 2/3 its length and pubescent on the lower 1/3; capsule oblong to oval, 2.5-4 mm long, 3-3.2 mm wide, setose and gland- ular; mature seeds elliptic, brown, 2.7-3 mm long, 1.3-1.6 mm wide, ventral sur- face pitted with a central ridge gradually tapering toward the margins, excavations lacking on each side of ridge, dorsal sur- face flattish with a faint longitudinal groove down the center, pitted (Fig. 17). Collections: 83 (0); representative: T. S. Brandegee 1139 (uc); M. Jones 511 (pom); C. Baker 549 (gh, ny, pom. rm, us); L. Higgins 2235, 2236, 2244, (bry, WTs) ; A. Nelson 9812 (rm, uc) ; W. Web- er 9416 (rsa, uc. ut); B. Hartman 2782 (bry. rm). Habit.\t. — Gravelly and sandy soils and talus slopes from 7,050 feet elevation upward to tiniberline. Commonly in open tundra and grassy alpine slopes of spruce, fir, pine, or aspen communities. Some- times growing as a weed along roadsides and in waste field, July to September. Distribution. — Mostly along or east of the Continental Divide in central and south central Colorado, south to Colfax Co., New Mexico (Map 5). Fig. 16. Phacelia bakeri (Brand) Macbride. R. Hartman 2782 (bry). This taxon was misinterpreted by Brand (1913). The type specimen is the only material cited by him that belongs to bakeri in a strict sense. Macbride (1917) and Voss (1937) present additional in- formation on Brand's treatment of this entity. P. bakeri is related to P. gland- iilosa Nutt., differing in the lack of excavations on the ventral surface of the seeds, usually greener herbage, later flowering time, and distribution. 148 GREAT BASIN NATURALIST Vol. 35, No. 2 ^: Fig. 17. Dorsal and ventral view of the seeds of P. bakeri (Brand) Macbride. C. Baker 549 (gh). 8. Pfiacelia hombycina Wooton & Slandley Fig. 18 Phacelia bombycina Wooton & Standley. Contr. U.S. Natl. Herb. 16:163. 1913. Holotype: New Mexico: Carton Co.: on gravelly banks at Mangas Springs, March or April 1880, H. Rusby 276 (us!); Isotypes (ny, ny, uc, us). Paratypes: New Mexico: Grant Co.: Bear Mountains near Silver City, no date, C. Metcalfe 75 (ny, pom, uc, us, us). Phacelia tenuipes Wooton & Standley. Contr. U.S. Natl. Herb. 16:163. 1913. Holotype: Carizalillo Spring, 17 April 1902, E. Mearns 91 (us!). Annual 0.9-4.1 dm tall; stems 1 -sev- eral, often branched throughout, erect and very brittle, setose to puberulent and glandular at least in the inflorescence; basal leaves petiolate (the petiole up to 5 cm long), oblong to nearly orbicular, pinnatifid into oblong to ovate, crenate lobes, setose and often glandular on both surfaces, 1.5-8 cm long, 0.5-2.5 cm wide, upper reduced, short petiolate, lobed; in- florescence paniculate, narrow, with a few branches; individual cymes with numerous, crowded flowers, pedicels short, stout, setose to puberulent and glandular; sepals oblong to elliptical, up to 3.5 mm long, setose to glandular; corolla blue to violet, 5-6 mm long and wide, campan- ulate, lobes pubescent; stamens exserted, anthers yellow, filaments bluish violet; style exserted, bifid to about the middle, pubescent below the middle, bluish vio- let; capsule 2.5-2.7 mm long and broad, globose, pilose and glandular especially at the apex; mature seeds oblong to ellip- tical, cjTnbiform, 2.2-2.5 mm long, 1-1.4 mm wide, dark brown, ventral surface Map 5. Southern Colorado and northern New Mexico. Distribution of P. bakeri (Brand) Macbride. pitted and divided by a prominent ridge, the ridge corrugated on one side, margins corrugated, lighter than the center, dorsal surface pitted (Fig. 19). Collections 89 (9); representative: L. Goodding 2230 (rm, uc); W. Cottam 10198 (uc) ; D. Demaree 42048 (uc); L. Higgins 2877 (bry, wts); H. Ripley and R. Barnebv 4218 (rsa); D. Atwood 2195, 2241, 2256, 2280 (bry) ; D. Atwood 2250, 2253, 2255 (bry, CAS, NY. WTs). Habitat. — Sandy, gravelly, or lava sloj)es and mesas from an elevation of 1,500 to 7,500 feet. Commonly found in the Larrea, Prosopis, and other Lower Sonoran mixed shrub communities, late March to late May. Distribution. — Sierra Co., New Mex- ico, southward to Chihuahua and Sonora. Mexico, westward through southern and central Arizona to Yavapai and Coconino COS. (Map 6). The character differences that Wooton and Standley (1913) used to separate P. tenuipes and P. bombycina var}' de- pending on the maturity of the plants June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 149 Fig. 18. Phacelia bombycina Wooton & Standley. D. Atwood 2253 (bry). and environmental conditions. Of the two taxa, the original description of the latter better fits the entity concerned. For these reasons P. tenuipes is placed in synonomy. The holotype data indicate that P. bom- bycina was collected in March and April; however, an isotype in the U.S. herbarium Rives the exact data as 25 March 1880. This entity has been confused with both P. intermedia and P. crenulata. It is roacHly distinguished from the former by the exserted stamens, larger blue lavender corollas, very brittle and less glandular stems, and smaller seeds. It differs from the latter by its brittle stems and smaller, darker seeds. 150 GREAT BASIN NATURALIST Vol. 35, No. 2 Fig. 19. Dorsal and ventral view of the seeds of P. bombycina Wooton & Standley. D. Atwood 2255 (bry). 9. Phacelia coerulea Greene Fig. 20 Phacelia coerulea Greene, Bull. Torr. Bot. Club 8:122. 1881. Lectotype: bluffs of the Gila (New Mexico), 23 April 1881, E. Greene s.n. (gh!). Phacelia invenusta Gray, Proc. Amer. Acad. 20:303. 1885. Lectotype: Arizona: Pima Co.: Sierra Tucson, 12 April 1884, C. G Pringle s.n. (gh!); duplicates (us), in part Phacelia intermedia Wooton, in part. Para types: New Mexico: Socorro Co.: San An tonio, Quitman Mountains, 14 March 1852 C. Wright 1579 (gh, gh, gh, ny); Texas El Paso Co.: El Paso, March 1851, Thurber 11 (gh, gh, NY, ny). Annual, 0.5-6 dm tall; stems erect, branched throughout, reddish, puberulent to setose and sparsely to densely stipitate- glandular, leafy throughout; leaves oblong to ovate, gradually reduced from the base upward, upper deeply sinuate, lower pin- natifid, dorsal surface with setose ap- pressed hairs, dorsal surface setose to glandular, 0.6-8 cm long, 0.3-2.5 cm wide, petioles from 5 cm long at the base to nearly sessile at apex, margins crenate; inflorescence terminal, commonly loosely paniculate or cymose, setose, stlpitate- glandular and puberulent; scorpioid cymes compact in flower but loosening in fruit, 1.5-7 cm long; flowers nearly sessile in flower to 1 mm long in fruit; corolla campanulate, lobes pale mauve to blue (turning white in fruit), tube yellowish, 3-4 mm long and broad, glabrous; sepals narrowly oblanceolate to elliptical, 2.5-4 mm wide, setose to brownish stipitate- glandular, 3/4 as long as the corolla, shorter or sometimes longer than the cap- Map 6. Southwestern New Mexico, Arizona, and adjacent Sonora, Mexico. Distribution of P. bombycina Wooton and Standley. sule; stamens mostly included to slightly exserted, anthers yellow, ovate, filaments bluish; style included to slightly exserted, equaling the stamens; capsule globose, 2.5-3.5 mm long, 1-1.8 nun wide; mature seeds dark brown, ventral surface pitted and divided by a prominent ridge, the ridge corrugated on one side, margins usu- ally corrugated, dorsal surface pitted, 0.3- 0.4 mm of the margin slightly elevated and smoother than the pitted center (Fig. 21). Collections: 112 (6) ; representative: M. Jones s.n. (pom); W. Wooton s.n. (ny); C. Parry 934 (ny); E. Greene s.n. (gh); C. Wright 1579 (gh, gh, gh, ny) ; D. Atwood 2137, 2152, 2196a, 2573 (bry); D. Atwood 2197, 2281 (bry, gas, NY, wTs); L. Higgins 3126, 3134, 2978, 2999 (bry. wts). Habitat. — Gravelly and arid calcar- eous hills and banks, sandy-gravelly stream beds, and rocky ledges from 2,000 to 6.000 feet. Commonly associated with the paloverde and creosote mixed shrub communities. Usually locally scattered, late February to early July. June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 151 Fig. 20. Phacelia coerulea Greene. D. Atwood 2196a (dry). Distribution. — Southern Nevada in Clark Co., southeastern Cahfornia in San Bernardino Co., eastward through Mohave Co. to southern Arizona and southern New Mexico from Socorro and Lincoln COS. to EI Paso, Presidio, and Brewster COS., Texas, and adjacent Mexico in the state of Chihuahua. Onlv one collection 152 GREAT BASIN NATURALIST Vol. 35, No. 2 Fig. 21. Dorsal and ventral view of the seeds of P. coerulea Greene. H. Ripley and R. Barneby 3361 (c.as). is knowTi from Nevada and California (Map 7). Greene cited no material on which he based his original description in November 1881. Voss (1937) noted this fact and se- lected Greene's collection at the Gray Herbarium as the type, since he had col- lected P. coerulea in the spring (23 April Map 7. Southern Nevada, southeastern Cali- fornia, Arizona, New Me.xico, western Texas, and adjacent Chihuahua, Mexico. Distribution of P. coerulea Greene. 1881) of the same year. However, Howell (1943), in Sertulum Greeneanum, indi- cates that a part of the type collection chosen by Voss is present in the Greene Herbarium at Notre Dame and that this specimen should be the type. Since Voss selected the specimen at the Gray Her- barium as the type, it should probably stand as such even though Greene's orig- inal is at Notre Dame. This taxon is most closely related to P. bombycina and secondarily to P. cren- ulata. It differs from these species in its included stamens and smaller corollas. Jones (1908) contended that P. coerulea and P. invcnusta Gray were separate species. The author believes, as did Gray (1886) and Voss (1937), that the latter is the same entity as the former. Howell (1941) reported P. coerulea from San Bernardino Co., California (H. Ripley and R. Barneby 3361 (cas, rsa) . To my know- ledge this is the only collection from California. No type was selected by Gray when he described P. invenusta; therefore I have selected the Pringle collection (gh) as the lectotype. Duplicates are located at cas and a fragment at us. 10. Phacelia congesta Hooker Fig. 22 Phacelia^ congesta Hooker. Bot. Mag. 62: t. 3452. 1835. Holotype: Texas: Galveston Co.: Gal- veston Bay. Drummond 303 (gh!). Phacelia congesta Hooker var. typica Voss. Bull. Torr. Bot. Club 64:133. 1937. Torr. Bot. Club 64:133. 1937. Phacelia congesta Hooker var. dissecta Gray. Syn. Fl. 11:1 SuppL, 415. 1886. Holotype: Texas: Dallas Co.: shaded rocks. Dallas, May and July 1880, Reverchon s.n. (gh!); dupli- cates (C.-^S, GH, GH, NY, Ny). Phacelia dissecta (Gray) Small. Fl. South- eastern U.S. pp. 972, 1337. 1903. Phacelia conferta D. Don. Gen. Syst. Gard. 4:397. 1837. Erect, often robust, annual plants, 1-10 dm tall; stems simple or diffusely branched throughout, with midticellular stipitate glands and puberulent to hispid unicellular hairs; leaves oblong in out- line, pinnately compound, the 3 ter- minal lobes usually not completely lobed to the midrib, often somewhat larger than the lower, usually petiolate lobes, 1-12 cm long, 0.5-4 cm wide, strigose and sometimes glandular: inflorescence of June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 153 J '< 'fl§- terminal, compound, scorpioid cymes, the individual cymes 1.5-15 cm long, pedicels 2.5 mm long or less, pubescent; sepals narrowly linear, 3-4.5 mm long, 0.5-0.7 mm wide, setose to hispid and often gland- ular; corolla campanulate, blue (rarely white), 4-6 mm long and wide, the lobes pubescent; stamens and style exserted, ca. 2-4 mm long, anthers pale yellow, filaments purplish; style 7-8 mm long, bifid 3/4 its length, lower 1/4 pubescent; capsule subglobose to oval, 2.3-3.6 mm long, 2.3-3 mm wide, puberulent and often glandular; mature seeds usually 4, sometimes 1 or 2, 2.6-3.2 mm long, 1.2- 1.4 mm wide, elliptical to oblong, brown, reticulate to scabrous, ventral surface ex- cavated on both sides of the ridge (Fig. 23). Collections: 222 (21); representa- tive: V. Cory 28660 (gh); R. McVaugh 7780 (uc); E. Palmer 33743 (ny, us); H. Ripley and R. Barneby 11107 (c^s) ; E. Tyler s.n. (us); L. Higgins 2671, 3162 (bry); D. Atwood 2048a, 2049, 2063, 2098, 2099, 2104-2107, 2111, 2117 (bry). Habitat. — Commonly associated with Prosopis, Larrecu Acacia^ and Opuntia in sandy to sandy loam, rocky limestone, or sandstone flats and outcrops. Along the coast of southern Texas it grows on low shoreline dunes and is usually as- sociated with Quercus, February to Sep- tember, 300 to 7,000 feet. Distribution. — Scattered throughout most of central and southern Texas, west to Eddv and Dona Ana cos., New Mexico; northeastern Mexico and Caddo and Com- Fig. 22. Phacelia congesta Hooker. wood 2046 (bry). D. At- Fig. 23. Dorsal and ventral view of the seeds of P. congesta Hooker. D. Atwood 2117 ( BRY ) . 154 GREAT BASIN NATURALIST Vol. 35, No. 2 manche cos., Oklahoma. Specimens from Florida, Massachusetts, and Sweden are presumably cultivated (Map 8). Some plants from Tamaulipas, Zacate- cas, and adjacent Nuevo Leon, Mexico, are fall-flowering and differ in sufficient morphological features to warrant further investigation. They are apparently rhiz- omatous perennials and possess a con- gested inflorescence and small, pale lav- ender flowers. This taxon varies throughout its range in pubescence and in leaf size and shape. The number of seeds per capsule was used by former workers as an important character in separating var. dissecta from var. congesta. However, the author has examined capsules from the type ma- terial of var. dissecta and found that they possess 4 seeds. Other material ex- amined varies in the number of seeds per capsule. This variation probably results from environmental conditions and is hardly consistent enough to warrant tax- onomic recognition. The seeds of P. congesta are dark when immature and are light brown upon reaching maturity. A more or less mottled pattern can be observed in the different stages of development. 11. Phacelia constancei Atwood Fig. 24 Phacelia constancei Atwood. Rhodora 74(800): 451-454. 1972. Holotype: Arizona: Coco- nino Co.: 1 mi north of Fredonia, 27 Mav 1968, D. Atwood, 1385a (bry!). Isotypes (aRIZ, bry, CAS, NY, UC, Us). Erect biennial herb, 1.5-4.3 dm tall, leafy throughout; stems stout, simple or branched throughout, reddish, from hir- sutulous to hirsute, and finely glandular; leaves mostly petiolate, 1-10 cm long, 0.3- 1.5 cm wide, revolute, from undulate to pinnatifid, linear to lanceolate, upper sur- face hirsutulous with scattered glandular hairs; inflorescence of compound scorpioid cymes; pedicels to 1 mm long; sepals el- liptic to oblanceolate, 3-4 mm long, hir- sutulous to hirsute and stipitate-glandular; corolla tubular, whitish, 5-6 mm long; sta- mens exserted 3-4 mm; style bifid, lower 1/3 pubescent, exserted 3-4 mm longer than the stamens; capsule subglobose, glandular, and hirsutulous throughout. Map 8. Texas, adjacent New Mexico, Okla- homa, and Mexico. Distribution of P. congesta Hooker. shorter than the sepals, mature seeds 4, black, 2.5-2.8 mm long, 1-1.2 mm wide, el- liptic, the margins corrugated, ventral surface finely pitted, excavated, and di- vided by a prominent ridge, the ridge cor- rugated on one side, the dorsal surface finely pitted. Collections: 12 (10); rep- resentative: H. Ripley and R. Barneby 4836 (cAs, rsa); D. Atwood 1529 (bry, GH, uc); D. Atwood 1785 (ariz, bry, cas, NY, uc, us, uTc); D. Atwood 1792b (bry, CAS, GH, NY, POM. RM. US). Habitat. — Alkaline clay bluffs and flats of the Moenkopi formation, 5,500 feet. Late May to early August. Distribution. — Mohave Co., Arizona, and Kane Co., Utah (Map 9). Taxonomically P. constancei appears to be most closely related to P. palmeri Torr. ex Wats, but is distinguished from that species by the smaller growth form, nar- rower and more revolute leaves, the red- dish stems with shorter and fewer hairs, and the leafier lateral inflorescence branches. June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 155 Fig. 24. Phacelia constancei Atwood. D. Atwood 1 835a (bry). 156 GREAT BASIN NATURALIST Vol. 35, No. 2 Map 9. Southern Utali and northern Ari- zona. Distribution of P. constancci Atwood. 12. Phacelia corrugata A. Nelson Fig. 25 Phacelia corrugata A. Nelson, Bot. Gaz. 34:26. 1902. Holotype: Colorado: Garfield Co.: Rifle, 23 June 1900. G. E. Osterhout 2129 (rm!); Isotypes (ny, rm). Paratypes: Colorado: Mesa Co.: Palisades, 14 May 1898, C. S. Crandall 4174 (rm, rm, us). Phacelia crenulata Torr. ex Wats. var. cor- rugata (A. Nels.) Brand, Das Pflanzenreich IV, 251:79. 1913. Phacelia orbicularis Rydberg, Bull. Torr. Bot. Club 40:479. 1913. Holotype: Utah: Wayne Co.: Marvine Laccolite. 22 July 1894, M, Jones 4663 (us!). Plants annual or winter annuals, 0.5- 4.3 dm tall; stems greenish, from puber- ulent to finely stipitate-glandular and sometimes with a few longer hairs inter- mixed; leaves 1-10 cm long, setose to stri- gose and stipitate-glandular; sepals 4-5.5 mm long; corolla campanulate, deep blue, 6 mm long or more; stamens and style exserted over 3 mm, filaments and style blue; capsule elliptic, 3.8-4.5 mm lon^; mature seeds oblong to elliptic, light brown, 3.1-4 mm long, 1.3-1.7 mm wide, pitted, the ventral surface corrugated on the margins and one side of the ridge (Fig. 26). Collections: 216 (44); representa- tive: R. Barneby 13033 (cas, ny) : C. Par- ry s.n. (cas, uc) ; A. Eastwood and J. Howell 9358 (cas, gh, utc); L. Higgins 3304, 3303, 3305 (bry, wts); D. Atwood 1314, 1489, 1856, 2539, 2523, 2583, 2581, 2618 (bry). Habitat. — This species grows in a large number of habitats from dry, grav- elly hillsides and flats, sandy soil, and red shaly clay to heavy clay soils. It grows with Atriplex, Sarcobatus^ Ephedra, Coleogyne or grass, from 3,500 to 7,000 feet elevation. Late April to mid- July. Distribution. — Eastern Nevada, east- ward to Colorado from Garfield and Gun- Fig. 25. Phacelia corrugata A. Nelson. D. Atvvood 2457 (bry). 1975 ATWOOD: PHACELIA CRENULATAE GROUP 157 ^B .Lj ! Fig. 26. Dorsal and ventral viev/ of the I seeds of P. corrugata A. Nelson. D. Atwood 2583 i (bry). nison cos., southward through Ouray and Montezuma cos. to northwestern New Mexico and northern Arizona (Map 10). This species is related to P. crenulata and apparently intergrades with it in eastern Nevada and western Utah. In general, however, it is easily distinguished from the latter by its greenish yellow stems, finer pubescence, and lighter, nar- rower seeds. 13. Phacelia coulter i Greenman Phacelia coulteri Greenman. Proc. Amer. Acad. 41:241. 1904. Holotype: Mexico: state of Hidalgo: fields about Buena Vista Station. 4 August 1904, C. Pringle 8988 (gh!); Iso- types (cAS, MEXU, ny, ny, pom, uc, us, us). Paratypes: Mexico: Vera Cruz: Real del Monte, T. Coulter, no date 921 (?). Phacelia neoniexicana Thurber ex Torr. var. coulteri (Greenman) Brand. Das Pflanzen- reich IV. 251:84. 1913, Phacelia grandulosa Hem. Biol. Cent. Amer. Bot. 2:359. 1882. Stout annual, 3.4-5.4 dm tall; stems solitary or branched, brownish to red- dish, hirsute and provided with a softer indument; leaves pinnately compound, finely dissected, 0.2-1 dm long, 3-4.5 cm wide, hispid, with small dark glands, mid- rib and some of the lateral veins prom- inent ventrally and therefore producing a furrowlike appearance dorsally, the mar- gins of the pinnae thick ventrally, lower leaves petiolate, the pedicels gradually re- duced upward; inflorescence of terminal compound scorpioid cymes, hispid, with brownish glandular hairs, the cymes con- gested, up to 7 cm long, pedicels 1 mm long or less; sepals linear to oblanceolate, Map 10. New Mexico, Utah, and adjacent Nevada. corrugata A. Nelson. Arizona, Colorado, Distribution of P. 3.3-3.6 mm long, 0.5-1.3 mm wide, hir- sute and brownish glandular; corolla campanulate, bluish purple, 4-5 mm long and broad, finely pubescent petals barely fimbriate; stamens and style exserted 1- 3.5 mm, the anthers yellow, filaments bluish; style ca. 6 mm long, bifid 1/2 its length, lower 1/2 glandular and puber- ulent; capsule oval, 2.4-3 mm long and broad, glandular and puberulent; mature seeds 4, oblong, brown, 1.6-1.9 mm long, 0.9-1 mm wide, pitted over the entire sur- face, the ridge level with the margins and deeply excavated on both sides (Fig. 27). Collections: 3 (0); representative: C. Pringle 8988 (cas, gh, mexu, ny, pom, uc, us); G. Rzedowski 16995 (mexu); Tinsley et al. 8 (uc). Habitat. — A weed of fields and road- sides as well as meadows and well-vege- tated areas in the mountains, 7,350 to 8,500 feet. July to August. Distribution. — Known only from the states of Hidalgo and Zacatecas, Mexico. Greenman did not select a holotype but Pringle 8988 was cited and is probably the type material. 158 GREAT BASIN NATURALIST Vol. 35, No. 2 foliis angustioribus, corollis violaceis, stam- inibus et stylis excertis 5-6 mm, capsulis subglobosis 3-4 mm longis 2-3 mm latis, seminibus ellipticis 2-3 mm longis, 1-1.75 mm latis. Annuals, 1.4-3.9 dm tall; stems simple to more commonly branched from the base, finely and densely stipitate-glandu- lar, setose and puberulent, reddish purple especially below; leaves 1.1-9 cm long, 0.2- 1 cm wide (up to 2.2 mm wide), stipitate- glandular and setose, petiolate; pedicels 1 mm long; corolla campanulate, violet, 7-10 mm long and broad; stamens and style exserted 5-6 mm; capsule subglobose, 3-4 Fig. 27. Dorsal and ventral view of the seeds of P. coulteri Greenman. C. Pringle 8988 (mexu). Phacelia coulteri is related to P. alba but differs from that taxon in having smaller seeds (2 mm long) and larger campanulate corollas (4.5 mm long). The stamens and style are exserted up to 3.5 mm, the corolla lobes are only shallowly erose, and the leaves are more finely dis- sected. 14. Phacelia crenulata Torr. in Wats. Plants annual, 0.25-8.3 dm tall; stems 1 -several, simple to much branched, stip- itate-glandular, setose or puberulent, red- dish purple to green; leaves 0.4-1.2 dm long, 0.2-4.0 cm wide, strigose to setose and stipitate-glandular, sessile to petiolate; inflorescence of compound scorpioid cymes, setose and stipitate-glandular, the pedicels 0.5-2 mm long; sepals cleft to near the base, the lobes elliptic to oblanceolate, 3-3.5 mm long, 1-1.4 mm wide, glan- dular and setose; corolla campanulate to rotate-campanulate, violet, blue, or pur- ple, pubescent, 4.5-10 mm long and broad; stamens and style exserted 5.5-11 mm; style bifid 3/4 its length, glandular be- low; capsule globose to subglobose, 2.6-4.1 mm long, 2-3 mm wide, puberulent and glandular; seeds 4, elliptic to oblong, 2-3.6 mm long, 1-2 mm wide, the dorsal surface with a dark center and lighter margins, the ventral surface corrugated. 14a. Phacelia crenulata vnr. angustifolia At wood, var. no v. Fig. 28 Phacelia crenulatae Torr. in Wats. var. crenu- Fig. 28. Phacelia crenulata Ton-, in Wats. latae affinis sed brevioribus et ramosioribus. var. angustijolia Atwood. Atwood 2523 (dry). June 197: ATWOOD: PHACELIA CRENULATAE GROUP 159 mm long, 2-3 mm wide; seeds 4, elliptic, 2-3 mm long, 1-1.75 mm wide. Type. — Arizona; Coconino Co.: Small mesas just north of Wupatki National Monument Headquarters, sandy soil covered with volcanic ash, 18 May 1970, N.D. Atwood 2597 (Holot>ioe: bry; Iso- types: to be distributed). Additional materials examined: Ar- izona: Coconino Co.: D. Atwood 1784, 2602, 2600, 2606, 2604, 3650, 4559, 4555, 2597 (bry); L. Higgins 5187, 5396 (bry); L. Williams 5993 (bry); L. Cureton 45 (bry). Utah: Beaver Co.: D. Atwood and L. Higgins 3839 (bry). Garfield Co.: D. Atwood 1356 (bry). Kane Co.: D. Atwood 1532B, 3603, 3612 (bry); L. Higgins and D. Atwood 5247 (bry); R. Allen 211, 243 (bry). Habitat. — Sandy, clay, or rocky ground in the can3'ons and benches be- low 5,000 feet elevation, April- June. Distribution. — Coconino Co., Arizona, north to Kane, Garfield, and Beaver cos., Utah (Map 11). Map 1 1 . Utah, Nevada. Arizona, and Cali- fornia. Distribution of P. crenulata Torr. in Wats, var. crenulata #; var. augustifolia *. 14b. Phacelia crenulata var. crenulata Phacelia crenulata Torr. in Wats., Bot. King Exped. 251. 1871. Holotype: Nevada: Per- shing Co.: Trinity Mountains, May 1868, S. Watson 873 (ny!); duplicates (gh, us). Phacelia crenulata Torr in Wats. var. vul- garis Brand, Das Pflanzenreich IV. 251:78. 1913. Phacelia crenulata Torr. in Wats. var. funerea Voss in Munz. Man. So. Calif. Bot. 409, 600. 1935. Holotype: California: Mono Co.: Black Canyon, White Mountains, 12 May 1930, V. Duran 561 (pom!); Isotypes (b, CAS, GH, NY, UC, Us). Plants annual, 0.25-8.3 dm tall; stems 1 -several, simple or branched, puberulent, pilose, setose and stipitate-glandular, red- dish purple to green; leaves 0.4-1.2 dm long, 0.5-4 cm wide, strigose to setose and stipitate-glandular, sessile to petiolate; in- florescence of compound scorpioid cymes, puberulent to setose and stipitate-gland- ular, the pedicels 0.5-2 mm long; sepals elliptic to oblanceolate, 3-5.3 mm long, 1-1.4 mm wide, setose and stipitate-gland- ular; corolla campanulate to rotate-cam- panulate, blue, pale purple, or violet, pub- escent, 4.5-7 mm long and broad; stamens and style exserted 5.5-11 mm; style bifid 3/4 its length, glandular pubescent below; capsule globose to subglobose, 2.6-4.1 mm long, 2.3-3.2 mm wide, puberulent and glandular; seeds 4, elliptic to oblong, 2.8- 3.6 mm long, 1.2-2 mm wide, the dorsal surface with a dark center and light mar- gins, the ventral surface corrugated (Fig. 29). Collections: 124 (13); representa- tive: Lemmon s.n. (us); C. Purpus 5976 (uc); J. Howell 26588 (cas); C. Purpus s.n. (uc); D. Atwood 2597, 2600, 2623 Fig. 29. Dorsal and ventral view of the seeds of P. crenulata Torr. in Wats. var. crenu- lata. D. Atwood 2623 (bry). 160 GREAT BASIN NATURALIST Vol. 35, No. 2 (bry); p. Munz 14790 (cas, pom); A. Nelson 3433 (rm). Habitat. — Rock slides, limestone talus, lava flows, gravelly and sandy soil of the foothills and canyons, from 4,600 to 8,000 feet elevation. Late February to early July. Distribution. — Nevada, eastward to western and southern Utah (except Wash- ington Co.), and southward to Mohave and Coconino cos., Arizona, and eastern California from Nevada Co. southward to San Bernardino Co. (Map 11). Phacelia crenulata was described from immature specimens (lacking seeds) in the northern part of its range in north- western Nevada. This has resulted in some confusion as to its relationship to such other closely allied species as P. corrugata A. Nels. and P. ambigua Jones. Examination of specimens from Pershing Co., Nevada, as well as sur- rounding counties, reveals that P. cren- ulata is quite distinct from both P. cor- rugata and P. ambigua. However, there is some overlap in morphological charac- ters where the species grow together. For the most part P. crenulata differs from P. ambigua in its reddish, usually simple and only slightly setose stems, and its distinctive seed characters. P. corrugata is easily distinguished by its yellowish green stems, uniformly fine-glandular pubescence, and distinctive seed. The type of var. funera Voss appears to be rnorphologically the same as the type of P. crenulata. The author has been unable to study this complex in the field; when field observations and greenhouse studies have been accomplished, a change in taxonomic status may be necessary. However, until such studies are carried out, the present arrangement seems the most natural. 15. Phacelia denticulata Osterhout Fig. 30 Phacelia denticulata Osterhout. Torrcya 16:70. 1916. Holotypc: Colorado: Larimer Co.: Tlie Glades, Owl Canyon, between Fort Collins and Livermore. 18 June 1915, G. Osterhout 5233 (rm!); Isotypes (ny. rm. rm, rm). Phacelia glandiilosa Nutt. ssp. eu-glandulosn Brand var. australis Brand, Das Pflanzenreich IV. 251:82-83. 1913, in part. Lectotype: Colorado: El Paso Co.: Manitou. 15 July 1903. F.E. and F.S. Clements 47.1, in part (us!); duplicate (gh). Phacelia neomexicana Thurber ex Torr. var. microphylla Brand, Das Pflanzenreich IV, 251:84. 1913. Lectotype: Colorado: Can- yon, 18 July 1878, central Colorado, T. Brandegee s.n. (gh). Annual plants, 0.5-5.4 dm tall; stems erect, simple or sometimes branched, se- tose and stipitate-glandular; leaves oblong to oblanceolate in outhne, pinnately cleft or divided, 1-7.5 cm long, 0.5-4.5 cm wide, Fig. 30. Phacelia denticulata Osterhout. J. Ewan 18154 (uc). June 1975 ATWOOI): PHACELIA CRENULATAE GROUP 161 strigose and stipitate-glandular; inflores- cence of terminal scorpioid c}Tiies, the longer cymes becoming 10 cm long in fruit; sepals narrowly linear to oblanceo- late, 2.5 mm long in flower, 5-6 mm long in fruit, 0.8-0.9 mm wide, setose and stip- itate-glandular; corolla tubular, light blue, 3.5-4.5 mm long, 2.3 mm wide, the lobes short, denticulate; stamens included; style included. l)ifid, glandular at base; capsule ovoid, 5 mm long, 3-3.5 mm wide, ]:)ilose and glandular; mature seeds 4, el- liptical to oblong, brown, 4 mm long, 1.7 mm wide, ventral surface slightly exca- vated on each side of the curved ridge, alveolate, dorsal surface alveolate (Fig. 31). Collections: 61 (7); representative: H. Ripley and R. Barneby 7525 (cas, ny); W.^ Weber 5974 (c.\s); C. Shear 3306 (ny); a. Nelson 1361 (rm); D. At- wood 1941, 1946, 1959, 1973 (bry). Habit.at. — Gravelly, sandy, or clay banks, draws and flats of the prairie to higher mountain slopes from 6,800 to 9,700 feet. Commonly as understor}' of Quercus^ Cercocarpus. Artemisia, and Fi- nns edulis. Frequently associated with Populus trcmuloidcs and Pseudotsuga at the higher elevations, .Tune to September. • ' ^ • \ • i ; ^\ • i \ Y' \ 1 1 Map 12. Colorado and southern Wyoming. Distribution of P. dendculata Osterhout. Fig. 31. Dorsal and ventral view of the seeds of P. dendculata Osterhout. D. Atwood 1973 (bry). Distribution. — Albany and Laramie COS., Wyoming, southward through cen- tral Colorado (Map 12). Phacelia dendculata is most closely re- lated to P. neomexicana, from which it dif- feres in having included stamens, a denser glandular pubescence, larger seeds, and a more robust, erect habit. The holotype of P. neomexicana var. ?nicrop/iylla Brand was deposited in the Berlin Herbarium (Brand 1913). The author has seen the entire collection of the Crenulatae group at Berlin. Apparently most of their collection was destroyed during the war. This has made it neces- sary to select a lectotype for var. micro- phylla; the author has chosen the Brand- egee collection at GH as the lectotype. Brand (1913) failed to select a holotype for P. neomexicana var. australis. Most of the material cited by him belongs to P. bakeri (Pur]uis 838" Baker 549, and Clements 47.1, in part). The Clements collection consists of several sheets, most of which are P. bakeri. However, the Clements collection at the U.S. National Herbarium and Gra}^ Herbarium includes the only sheets which represent var. australis. I designate the specimen at the U.S. National Herbarium as the lectot>T)e. 16. Phacelia formosula Osterhout Fig. 32 Phacelia formosula Osterhout. Bull. Torr. Bot. Club 46:54. 1919. Holotype: Colorado: Jackson Co.: North Park near Waldren. along the road descending to Michigan Creek, 6 August 1918, G. Osterhout 5794 (rm!); Isotypes: (rm, rm, rm). 162 GREAT BASIN NATURALIST Vol. 35, No. 2 f -'f Fig. 32. Phacelia formosula Osterhout. D. Atvvood 1971 (bry). Annual, 1.5-2.2 dm high, up to 3 dm broad; stems single or branched through- out (especially at base), glandular and hirsute, somewhat grayish; leaves lanceo- late or elliptical, 3-7 cm long (up to 4.5), 1-3 cm wide, pinnately divided, strigose, hirsute and glandular; inflorescence of compound scorpioid cymes, cymes up to 5 cm long in fruit, and more densely glandular than the stems; sepals 3.2-3.8 mm long, 0.5-0.8 mm wide, glandular and hirsute; corolla campanulate, violet, 6 mm long, 6 mm wide, slightly glandular and pilose; stamens and style long exserted; style ca. 2 mm longer than the stamens and puberulcnt throughout; capsule ob- long to oval, 3.5 mm long, glandular and hirsute; mature seeds 4, oblong, dark brown, 2.5-3 mm long, 1.2-1.4 mm wide, excavated ventrally on each side of the ridge, pitted, dorsal surface pitted, margins rounded and smooth. Collections: 7 (2); representative: D. Keck 889 (cas, uc); H. Ripley and R. Barneby 9008 (cas); D. Atwood 1977, 1977a (bry). Habitat. — Loose sandy soil of sand- stone bluffs at an elevation of 8,300 feet. Associated with Artemisia and Tetrady- mia, July to August. Distribution. — Apparently confined to Jackson Co., Colorado. This species appears to be most closely related to P. glandulosa Nutt. but can be distinguished from that species by its usu- ally much-branched, erect to spreading habit, less exserted stamens and style, darker seeds, narrower calyx lobes, and more pubescent style. P. glandulosa is a somewhat variable species, and P. for- mosula may, perhaps, best be treated as a variety of it; however, additional mate- rials and field work are necessary before this suggestion can be confirmed. 17. Phacelia glandulosa Nutt. Fig. 33 Phacelia glandulosa Nutt.. Journ. Acad. Phil. n. ser. 1:160. 1848. Holotype: Wyoming: Lin- coln Co.: about Ham's Fork, Colorado, of the West, July, T. Nuttall 93 (gh?). Eutoca glandulosa (Nutt.) Hook., Kew Journ. Bot. 3:293. 1851. Wyoming: Sweetwater Co^: Greenriver. 31 May 1897. A. Nelson 3050 (rm!); Isotypes (gh, ny, rm). Phacelia glandulosa Nutt. subsp. eu-glandu- losa Brand var. deserla Brand. Das Pflanzenreich IV, 251.82. 1913. Plants annual or possibly biennial, 0.6- 3.6 dm tall; stems simple or branched, erect, densely stipitate-glandular and hir- sute; leaves lanceolate to oblong in out- line, pinnatifid, 1-7 cm long, 0.5-3 mm wide, glandular and densely hirsute, the lower petiolate, the upper subsessile; in- florescence of congested terminal com- pound scorpioid cymes, stipitate-glan- dular and hirsute, 1-1.5 mm long, cymes elongating to 6.5 cm in fruit; sepals el- liptical to oblanceolate, 3-4 mm long, 1.2- 1.4 mm wide; corolla campanulate, pur- ]ile to bluish, 5-7 mm long and broad, the lobes pubescent and often more or less crenate; stamens and style exserted 5-9 June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 163 .::-^ Fig. 33. Phacelia glandulosa Nutt. Hitch- cock 10804 (uc). min, the style bifid 3/4 its length, the lower 1/4 pubescent; capsule subglobose, 3.5-4 mm long, 3.2-3.3 mm wide, glan- dular and setose; mature seeds elliptic to oblong, reddish brown, 2.4-3.3 mm long, 1.1-1.4 mm wide, pitted, the ventral sur- face excavated on both sides of the ridge (Fig. 34). Collections: 25 (1); representa- tive: E. Graham 9774 (cas, us) ; R. Davis 585 (us); H. Ripley and R. Barneby 8826 (cas, ny); S. Watson 281 (gh, us); A. Rudvalis 70 (bry); H. Fitch s.n. (cas, NY, pom); a. Williams s.n. (ny). Fig. 34. Dorsal and ventral view of the seeds of P. glandulosa Nuttall. Habitat. — Rock slides, sandy talus slopes, and clay knolls from 5,000 to 7,050 feet. Mid-June to early August. Distribution. — West of the Continen- tal Divide in Rio Blanco Co., Colorado, northward to southwestern and western Wyoming, southwestern Montana, and central Idaho in Lemhi and Custer cos. (Map 13). For a discussion of this taxon see P. formosula. 18. Phacelia howcUiana Atwood Fig. 35 Phacelia howelliana Atwood. Rhodora 74(800): 456-462. 1972. Holotype: Utah: San Juan Co.: ca. 0.4 mi north of Bluff on Utah high- way 163, in mouth of canyon, 13 May 1970, D. Atwood 2454 (bry); Isotypes (ariz, asc, B, BRY, CAS, GH, NY, UC, US, UTc). Plants annual, 0.9-2.3 dm tall; stems mostly branched and leafy toward the base, glandular and hirsute; leaves broadly oblong to oval, 2.0-6.0 cm long, 1.0-2.5 cm wide, irregularly crenate to lobed, strigose and slightly glandular, the petiole up to 5 cm long; inflorescence of compound scorpioid cymes; pedicels up to 2 mm long; sepals linear to narrowly oblanceolate, 3.5-4.5 mm long, 1.0-1.2 mm wide, glandular and hirsute; corolla 5-6 mm long, 6-7 mm wide, rotate to funnelform, the lobes light violet to blue, the tube white; stamens and style ex- serted 3-4 mm, style shorter than the stamens, bifid 3/4 its length, lower 1/4 pubescent; capsule oblong to subglobose, glandular and hirsutulous, especially to- ward the apex; seeds 4, brown, 3.2-4 mm 164 GREAT BASIN NATURALIST Vol. 35, No. 2 Fig. 35. Phacelia howelliatia Atvvood. D. Atwood 2454 (bry). long, 1.4-1.8 mm wide, elliptical, the mar- gins corrugated, involute to flattened, ventral surface pitted, excavated, and di- vided by a ])rominent ridge, the ridge sometimes curved to one side and barely corrugated, dorsal surface reddish brown, smooth and surrounded by a lighter mar- gin (Fig. 36). Collections: 16 (3): rep- resentative: S. Welsh, D. Atwood and G. Moore 9957 (bry); A. Eastwood s.n. (ny); B. Harrison 11244 (bry); C. Han- sen 101 (bry); M. .lones s.n. (pom); D. Atwood 2511 (ARIZ. ASC, B. BRY, CAS- DIXTE. Gil. JEPS. NY. POM, RM, RSA, UC. US. ITT, UTC. wsc); ,T. Howell 24689 (CVS). Habitat. — Red sandv, gravellv. or clay soils at ca. 4,500 to 5,000 feet. ' Distribution. — Known only from San .Tuan and Grand cos., -Utah. It probabh OTows in (Colorado near Moab and also ATWOOD: PHACELIA CRENUL.\TAE GROUP 165 Map 13. Idaho, Montana, Wyoming, and Utah. Distribution of P. glandulosa Nuttall. Monument Valley in Arizona, although no specimens have been seen from either area (Map 14). This entity is related to P. corrugata A. Nelson which grows throughout most of Utah and extends to Colorado, northern New Mexico, and northern Arizona. It is distinguished from P. corrugata by its low, much-branched growth form and smaller corolla with a white tube. The leaves are mostly basal, and the seeds are larger and reddish brown. 19. Phacelia integri folia Torr. ex Watson Plants annual (possibly biennial in ^ar. te.vana) , 1.2-6 dm tall, stems stout, green to reddish brown, simple or branched, puberulent, stipitate-glandular and hirsute; leaves simple, crenate to somewhat cleft, oblong to ovate or lanceo- late, strigose, finely glandular, setose, lower leaves long petiolate, the upper short pctiolate to sessile; inflorescence of com- ()ound scorpioid cymes, the cymes elon- ij,ating to as much as 2.1 dm in fruit, ped- i( els 0.5-2 mm long; sepals elliptical to nl)]anceolate, 2.5-6.5 mm long, 0.9-2.8 mm wdde, often reddish, puberulent, hir- sute and stipitate-glandular; corolla cam- panulate, purplish to lavender, 4.5-6.5 Fig. 36. Dorsal and ventral view of the seeds of P. howelliana Atwood. J. T. Howell 24687 (c.^s). mm long and broad, the lobes crenulate, pubescent; stamens and style exserted 4- 6.5 mm; style bifid 2/3-3/4 its length, pubescent below, the filaments purplish, the stamens bluish green; capsule ovoid to globose, 2.6-5.3 mm long, 1.1-3.5 mm wide, glandular and puberulent; mature seeds ovate or elliptic to oblong, reddish brown or dark brown to black, 2.4-4.5 mm long, 1.3-2.2 nun wide, the dorsal surface pitted and transversely ridged, the margins more finely pitted than the Map 14. San Juan and Grand counties, Utah. Distribution of P. howelliana Atwood. 166 GREAT BASIN NATURALIST Vol. 35, No. 2 excavated portions (these tending to have surface excavated on both sides of the transversely elongate pits), the ventral ridge, corrugated or corrugations lacking. la. lb. Key to the varieties of P. integrifoUa Mature seeds 3 mm long or less, 1.4 mm wide or less, ventrally cor- rugated; capsule 3.1 mm long or less; plants of southeastern New Mexico and adjacent Texas var. texana Mature seeds 3.1 mm long or more, 1.7 mm wide or more, ventral corrugations lacking; capsule 3.2 mm long or more; plants wide- spread in rocky to sandy soil var. integrifoUa 19a. var. integrifoUa Fig. 37 Phacelia integrifoUa Torr. in Wats., Ann. Lye. New York 2:222, t. 3. 1826. Holotype: on the Platte, 25 June 1820. Dr. James s.n. (ny!). Phacelia arenicola Brandegee. Univ. Calif. Pub. Bot. 4:185. 1911. Holotype: Mex- ico: Coahuila: El Tore near Movano. July 1910, C. A. Purpus 4458 (us!). Phacelia integrifoUa Torr. in Wats. var. arenicola (Brandegee) Brand. Das Pflanzen- reich IV, 251:82. 1913. Plants annual, 1.6-6 dm tall; stems pu- berulent, finely to densely stipitate-glan- dular and hirsute; leaves 1-13 cm long, 0.5- 3 cm wide; cymes elongating to 2.1 cm in fruit, pedicels 1 mm long; sepals ob- lanceolate to elliptic, 3.5-4.5 mm long (4.4-6.5 mm in fruit), 1-1.8 mm wide (1.1-2.8 mm in fruit); corolla 5-6.5 mm long and broad; stamens and style ex- serted 5-6 mm; capsule ovoid to globose, 3.2-5.3 mm long, 3-3.5 mm wide; mature seeds oblong to elliptic, dark brown to black, 3.1-4.5 mm long, 1.7-2.2 mm wide, transverse ridges on the dorsal surface cpiitG distinct, the ventral surface lacking corrugations, the ridge often curved to one side (Fig. 38). Collections: 113 (23); rep- resentative: L. Higgins 3138, 3129, 3131 (bry); D. Atwood 2555, 2556, 2278, 2275, 2273, 2169 (b, bry, cas, ny. wts) ; D. At- wood 2265, 2263, 2557a,' 2274, 2171 (bry). Habit.at.— Sandy hills and flats, rocky hillsides of Larrca, Yucca, Quercus, Coleo- gyne. and grass communities. From 3.750 to 7,500 feet, late March to mid-Ser)tem- ber. Distribution. — Southeastern Utah in Kane and San Juan cos., southward through northeastern Arizona, eastward through much of Now Mexico to western Texas and Chihuahua, Mexico (Map 15) Fig. 37. Phacrlia integrifoUa Torr. e.\ Wats. var. integrifoUa. L, Higgins 3131 (bry). June 1975 ATWOOD: PHACELIA CRENLLATAE GROUP 167 Fig. 38. Dorsal and ventral view of the seeds of P. integrifolia Torr. ex Wats. var. in- tegrifolia. D. Atwood 2556 (bry). Considerable confusion has existed as to the relationships of this taxon to other species. This has probably resulted from the inadequate type material and initial misunderstandings that have been per- petuated and even enlarged upon by sub- sequent authors, Brand fl913) and Voss (1937). These misunderstandings have, in part, come about through lack of field work and by the fact that the corollas in most herbarium specimens fade to white Map 15. Parts of southwestern United States and adjacent Mexico. Distribution of P. integrifolia Torr. ex Wats.: # var. integrifolia; var. O. tex- ana. and appear to be tubular. Var. integrifolia has a broad distribution and is easily dis- tinguished b}' its large, broad, noncor- rugated seeds. Plants from the higher elevations in western New^ Mexico and eastern Arizona are different morpho- logically. Additional field work is nec- essar)' to decide whether these differ- ences are sufficient to warrant taxonomic recognition. 19b. var. texana (Vossj Atwood, new- comb. Phacelia texana Voss. Bull. Torr. Bot. Club 64:141. 1937. Holotvpe: Texas: Hudspeth Co.: Fin- lay, 5 May 1931, M. E. Jones 28500 (pom!); Isotypes (rm. uc); photo at (bry. ny. uc, us). Plants annual or possibly biennial. 1.2- 4.3 dm tall: stems puberulent, densely covered with short stipitate glands (usu- ally 0.2 mm long or less; and sometimes ^^dth a few scattered longer, simple hairs; leaves 1-10 cm long, 0.3-2 cm wide, stip- itate-glandular (0.2 mm long or less), and puberulent; cymes elongating to 1.4 dm in fruit, pedicels 0.5-1.2 mm long; sepals elliptical to oblanceolate, more or less heteromorphic ^two narrow and three broad), 2.5-3 mm long (3.5-5.5 in fruit), 0.9-1.4 mm wide (1.2-2 in fruit) puber- ulent, stipitate-glandular, and setose; co- rolla 4.5-6 mm long and broad; stamens and style exserted 4-6.5 mm; capsule globose, 2.6-3.2 mm long, 1.1-2.6 mm wide; mature seeds ovate, reddish bro\Mi, 2.4-3 mm long, 1.1-1.4 mm wide, trans- verse ridges on the dorsal surface only fairly distinct, the central surface cor- rugated on the ridge and part of the mar- gin. Collections: 38 (Oj; representative: C. Cory 37574 (uc); H. Wilkens 2209 (uc); L. Higgins 3157 (bry, wts) ; T. Collins 1182 (uc); U. Waterfall 4558 (gh, CAS, NY); R. McVaugh 8163 (uc). H.ABiTAT.^ — Gypsum, limestone, and cal- careous soils in mixed shrub commun- ities. Commonly associated with Larrea. Foquieria, and Acacia, 2,550 to 6,850 feet elevation. Late April to early October. Distribution. — Southeastern New Mexico and adjacent Texas (Map 15). Var. texana differs morphologically from var. integrifolia in seeds and size of capsule. The corollas of var. texana ap- pear to be lavender in contrast to the pur- 168 GREAT BASIN NATURALIST Vol. 35, No. 2 plish corollas of the latter. Jones (1931) indicated that he had collected the type at Findlay, correctly spelled Finlay. The correct collection number is 28500 rather than 285ae as cited by Voss (1937). 20. Phacelia intermedia Wooton Fig. 39 Phacelia intermedia Wooton. Bull. Torr. Bot. Club 25:457. 1898. Holotype: New Mex- ico: Dona Ana Co.: mesa west of the Organ Mountains. 10 April 1893, E. O. Wooton (us!); duplicates (uc, us). Plants annual, 0.6-3.6 dm tall, stems simple or branched, often reddish, puber- ulent with short stipitate glands; leaves narrowly oblong, elliptic to ovate-lanceo- late, sinuate to deeply lobed or pinna- tifid, 0.5-8 cm long, strigose and stipitate- glandular or glandular, petiolate to ses- sile above; inflorescence of compound scorpioid cymes, terminal on the main stem and lateral branches, the cymes up to 1.5 dm long in fruit, pedicels subses- sile to 1 mm long in flower, slightly longer in fruit; sepals linear to oblanceolate, 2.5- 3.7 mm long, 1.3-1.5 mm wide, setose and glandular; corolla bluish violet, 6 mm long or less, campanulate, pubescent, the lobes crenate to entire; stamens and style exserted 3 mm or less, filaments violet, anthers yellow, style violet, bifid 2/3 its length, the lower 1/3 glandular and puberulent; capsule oval, 3 mm long and broad, glandular and puberulent; ma- ture seeds ovate, 2.7-2.9 mm long, 1.5- 1.6 mm wide, dark brown, pitted, ventral surface corrugated on one side of the ridge, both margins and partly to com- pletel}^ across the excavations (Fig. 40) . Collections: 34 (7); representative: E. Wooton s.n. (us); D. Atwood 2554, 2560, 2570, 2565, 2572. 2557. 2170, (bry); L. Higgins 3118, 3114 (bry, wts) ; H. Bob- isud 149 (nm). Habit.at. — Sandy to gravelly or c\i\\ soils of foothills and higher mesas from 3,750 to 5,000 feet. Often associated with Larrea and Prosopis, March to May. Distribution. — Central New Mexico, southward to western Texas and northern Mexico (Map 16). This taxon is related to P. corruania, Follugia. and Atriplex. Distribution. — Clark Co., Nevada, eastward to Washington and Iron cos., Utah, and Mohave Co., Arizona (Map 19). P. palmeri is related to P. constancei Atwood but differs in having taller stems, larger and less revolute leaves, and coarser and longer hair. The lateral branches are fewer and less leafy, and the stems lack the reddish color characteristic of the latter. Both species occur on the Moenkopi formation but have different ranges. A form from Lake Mead, Arizona (E. U. Clover 6470 & 6230), has seeds that are not corrugated and are smaller and narrower (2.0 mm long and 1.2 mm wide), dark dorsally but brown ventrally with darker glands. However, in the ma- r Fig. 43. Dorsal and ventral view of the seeds of P. palmeri Torr. ex Wats. D. Atwood 1723 (bry). Map 19. Southwestern Utali, northwestern Arizona, and southeastern Nevada. Distribution of P. palmeri Torr. ex Wats. terial from Glendale, Nevada, the seeds are dark dorsally and lighter ventrally and are less corrugated than in typical material. 25. Phacelia pcdiceUata Gray Fig. 44 Phacelia pedicellata Gray. Syn. Fl II. 1:160. 1878. Holotype: Mexicor Baja California: Lower California, 1875, Dr. Streets s.n. (gh!); Isotype (us). Plants annual, 1-6 dm tall; stems branching or sometimes simple, brittle, villous to setose with multicellular stalked glands; leaves suborbicular to oblong, pin- nately compound with 3-9 pinnae, 0.3-1.3 cm long, 1.5-11.5 cm wide, villous to setose and glandular; inflorescence of compound cymes, somewhat dichotomously branched; pedicels filiform, 2.6 mm long, densely setose to hirsute; sepals linear to oblance- olate, 2.8-7.9 mm long, 0.6-2 mm wide, setose to hirsute and glandular; corolla lobes lavender, violet, or white, the tube white, ca. 5 mm long and broad; stamens and style exserted, style pubescent, upper 174 GREAT BASIN NATURALIST Vol. 35, No. 2 Fig. 44. Phacelia pcdicellata Gray. W. Jcp- son 12482 (c.\s). Habitat. — Dry gravel and sandy washes, often in the shade of large boul- ders, limestone cliffs, and as understory of larger plants, below sea level in Death Valley to 5,000 feet elevation. It has been collected in the middle of February in Mexico but usually flowers from March to the middle of June. Distribution. — Southern Nevada in Nye and Clark cos., southward through San Bernardino to central Baja California, eastward to Coconino^ Graham, Gila, Pi- nal, and Pima cos., Arizona (Map 20). Phacelia pedicellata is most closely re- lated to P. scariosa h\xt differs in having narrower, longer, and more pubescent calyx lobes, a more compound and con- gested inflorescence, and a heavier, glan- dular, villous, and setose pubescence. The leaves have 3-9 pinnae, whereas those of P. scariosa have only 3-5 lobes, and the style is more pubescent. That they are related is indicated by the similar seeds, corolla, pedicels, and brittle stems. 26. Phacelia popei Torr. & Gray Fig. 46 Phacelia popei Torr. & Gray. Pacific Rail. Rep. Explor. Mississippi 2:172. 1885. Holotype: Te.xas: Llano Estacado, no date, Captain Pope s.n. (gh!). Paratype: Te.xas: Pecos Co.: C. Wright 1578 (gh, ny, us). Phacelia popei Torr. & Gray var. typica Voss. Bull. ToiT. Hot. Club 64:94. 1937. Phacelia similis Wooton & Standley. Bull. Torr. Bot. Club 36:111. 1909. Holotype: New Me.xico: Sierra Co.: on the plains near Nutt Station, 12 May 1905, O. Metcalfe 1665 (ny!). Phacelia popei Torr. & Gray var. similis 1/3 bifid; capsule globular, 3-3.4 mm long, 2.3-2.5 mm wide, pilose to glan- dular; mature seeds 4, elliptical, 3 mm long, 1.1-1.8 mm wide, ventral surface ex- cavated, pitted to tuberculate, the ridge corrugated on one side, dorsal surface tu- berculate and pitted, margins corrugated (Fig. 45). Collections: 155 (3); represen- tative: A. Eastwood 17400 (cas, ny, us); M. .Tones s.n. (pom, utc); W. Cottam 13125 (ut); T. Brandegee s.n. (ny); H. Ripley and R. Barneby 2952 (rsa); .1. Howell 3952 (utc); D. Atwood 2339 (bry) ; S. Welsh, D. Atwood and E. Matt- hews 9633 (bry). Fi^'. 45. Dorsal and \entral view of the seeds of P. pedicellata Gray. M. Beal (jeps). Tune 1975 AT WOOD: PHACELI V \ N •• • •\4 VN ^ \^_t - •• • • 1 S c»^ ■^"^^^ ^ ^ Map 20. Part of southwestern United States and adjacent Baja California, Mexico. Distribution of P. pedicellata Gray. (Wooton & Standley) Voss. Bull. Torr. Hot. Club 64:94. 1913. Phacelia glandulosa A. Gray in Brand, Das Pflanzenreich IV, 251:84. 1913, in syn- onomy. Phacelia depauperata Wooton & Standley. Contr. U. S. Natl. Herb. 16:163. 1913. Holotype: New Mexico: Caves Co.: Arroyo Ranch near Roswell, 1903, D, Griffiths 4249 (us!). branched from the base, 0.5-3.6 dm tall, with simple spreading or bent hirsute and somewhat glandular hairs intermixed with a finer pubescence; leaves narrowly ob- long, pinnate to bipinnate, with linear or lanceolate divisions, 2-15 cm long, 1-3 cm wide, petiolate, strigose to glandular, ex- cept on the petioles and then like that on the stems; inflorescence of compound scorpioid cymes, the cymes crowded, ter- minal, up to 10 cm long in fruit, glan- dular and hirsute, flowers nearly sessile (pedicels ca. 0.5 mm long) ; corolla cam- panulate, blue to purplish, 3.5-7 mm long and broad, pubescent; sepals oblanceolate to spatulate, somewhat keeled (at least at. the base), 2.3-3.9 mm long, 1-1.8 mm wide, glandular, hirsute; stamens and 175 style exserted, style exserted ca 2 mm longer than the stamens, bifid 2/3 its length, lower 1/3 pubescent; capsule glo- bose, 2.3-2.5 mm long, 2.4-3.1 mm wide, glandular and pilose; mature seeds 4, cymbiform, ovate, brown, 1.8 mm long, 1.4 mm wide, favose to reticulate, ventral surface deeply excavated on both sides of the ridge, dorsal surface reticulate and transversely ridged (Fig. 47). Collections: 150 (11); representative: D. Atwood 2096a, 2095, 2268, 2266, 2153, 2159, 2131, 2133, 2134 (bry); L. Higgins 3083, 3025 (bry, wts); R. Barneby 12593 (cas, ny) ; M. Jones 25750 (pom); A. Hershey s.n. (cas). Habitat. — Commonly in sandy or sandy clay soil of roadsides. Less com- monly in limestone or rocky soil and as- sociated with Larrea, Prosopis^ Yucca, or short grass prairie. Mid-February to late May. Fig. 46. Higgins 2931 Phacelia popei Torrey & Gray. L. (bry). 176 GREAT BASIN NATURALIST Vol. 35, No. 2 Fig. 47. Dorsal and ventral view of the seeds of P.popei Torrey & Gray. V. Cory 13616 (gh). Distribution. — West central Texas from Dickens and Lubbock cos. soutli- ward to southwestern Texas, westward to New Mexico from Roosevelt Co., west- W'ard to Sierra and Luna cos., Nuevo Leon, Mexico (Map 21). Wooton and Standley (1909), in their original description, indicated that P. si- milis was most closely related to P. arizo- nica but failed to mention P. popei. Com- parison of the type material of the two entities leaves little doubt that they are identical. Also, P. arizonica is very dis- tinct from P. popei. Torrey and Gray (1885), in their original description of P. popei., indicated that the corolla was "apparently white." I have seen this tax- on in the field in Texas and New Mexico and find that the corolla is not white or only seldom so. The flowers of some specimens turn white upon drying, and this may have been the case with the type material. 27. Pliacclia rafachmsis Atwood Fig. 48 Phacelia rafaelcnsis Atwood. Rhodora 74(800): 454-457. 1972. Holotype: Utah: Wayne Co.: Capitol Reef National Monument, 12 June 1969, D. Atwood and L. Higgins 1834 (bry!). Map 21. New Mexico, Texas, and Nuevo Leon, Mexico. Distribution of P. popei Torrey & Gray. Fig. 4b. Phacelia rafaelcnsis Atwood. D. At- wood and L. Higgins 1834 (liin'). June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 177 Erect biennial herb, 0.8-5.4 dm tall; stems stout, simple or sometimes branched at the base, olive green to brownish glan- dular, and hirsute; basal leaves petiolate, dentate, crenate to pinnatifid, 2-7 cm long; 0.5-1.5 cm wide, strigose to hirsute, cau- line leaves sessile, undulate to crenate or dentate, oblong-lanceolate, 1-10 cm long, 0.5-3.5 cm wide, strigose to hirsute and sparsely stipitate-glandular; inflores- cence mainly terminal, paniculate, some axillary, flowers nearly sessile; sepals oblanceolate to spatulate, 5-4 mm long in flower, 5-6 mm long in fruit, 1-1.7 mm wide, glandular and hirsute; corolla tub- ular, pale and grooved with the lobes somewhat spreading, 5-6 mm long; sta- mens and style exserted only 3-5 mm, anthers dull in color, style bifid 3/4 its length, the lower half pubescent; capsule globose, 4-5 mm long, stipitate-glandular and hirsute; mature seeds 4, elliptic to oblong, 3.5-4 mm long, 1.5-2 mm wide, ventral surface alveolate, lighter than the dorsal surface, excavated and divided by a prominent ridge, the ridge sometimes cor- rugated along one side, the margins usu- ally entire, dorsal surface brown and less deeply pitted, the surface often smoothish (Fig. 49). Collections: 33 (19); repre- sentative: M. Jones s.n. (pom); H. Ripley and R. Rarneby 4362 (rsa); W. Cottam 13313 (ut); D. Atw^ood 1530, 1853, 1843, 1855, 1847, 1703, 1417, 1698, 1860 (bry); S. Welsh, D. Atwood, and G. Moore 9846, 9844, 9903 (bry); D. Atwood 1390 (bry). Habit.at. — Clay hills of the Moenkopi formation. May to June. Distribution. — Emery Co., Utah, southward to Kane Co., Utah, and Mo- have Co., Arizona, eastward to Washing- ton Co., Utah (Map 22). Phacelia rafaelensis is related to P. uta- hensis but differs in having a slightly tub- ular and grooved corolla, with the lobes somewhat spreading and the stamens and style exserted only 3-5 mm. The filaments and stamens are dull in color, and the ridge is sometimes corrugated. 28.Phocelia robusta (Macbr.) Johnst. Fig. 50 Phacelia robusta (Macbr.) Johnst. Journ. Arnold. Arb. 24:97. 1943. Phacelia integrifolia Torr. e.x Wats. var. robusta Macbride. Contr. Gray Herb. 49:25. Fig. 49. Dorsal and ventral view of the seeds of P. rafaelensis Atwood. S. Welsh et al, 9903 (bry). 1917. Holotype: Texas: Presidio Co.: Chinati Mountains, no date, Harvard 250 (gh!). Robust, viscid annual or biennial, 4.5-12 dm tall; stems branched at the base or simple, brownish, puberulent, pilose and densely glandular, the glandular hairs flattened, stipitate, unicellular to multi- cellular; leaves broadly ovate to orbicular, irregularly crenate to sinuate, sometimes with a single lobe below the leaf blade, 2-11.5 cm long, 1.5-9 cm wide, gradually Map 22. Southern Utah and northwestern Arizona. Distribution of P. rafaelensis Atwood. 178 GREAT BASIN NATURALIST Vol. 35, No. 2 i \f L^' ' d r Fig. 50. Phacelia robusta (Macbride) ston. I. Higgins 3268 (bry). John- reduced upwards, puberulent, stipitate- glandular, setose to hirsute, the lower long- petiolate to subsessile above; inflorescence of compound scorjnoid cymes, the chines up to 1.3 dm long in fruit, pubescence the same as that of the stem, pedicels 1-1.7 mm long; sepals spatulate, 4.5-5.8 mm long, 1.5-2.5 mm wide; corolla salver- form, pale lavender, 5-6 mm long, ca. 4 mm wide, pubescent; stamens and style exserted 4-6 mm. style bifid 3/4 its length, lower 1/2 pubescent; capsule subglobose, 3.9-4.1 mm long, 2.8-3mm wide, puber- ulent to strigose; mature seeds oblong to ovate, reddish brown, 2.9-3.7 mm long, 1.1-1.7 mm wide, ventral surface exca- vated on both sides of the corrugated ridge, pitted, the margins at least partly corrugated (Fig. 51). Collections: 36 (0); representative: C. Pringle 255 (iic. rsa) : E. Palmer 34077 (ny): L. Hincklev 829 (gh, ny); U. Waterfall 7316, 8255, 8283 (uc); C. Smith 289 (uc); G. Stevens 1636 (gh, ny); L. Higgins 3256, 3182 (bry, wts). Habitat. — Gravelly sand bars, clay slopes, and rocky hills from ca. 3,500 to 6,000 feet elevation, March to August. Distribution. — Barber Co., Kansas, southward through Woods, Blaine, Custer, Washita, and Horman cos., Oklahoma, to north central and southwestern Texas and Chihuahua, Mexico (Map 23). This taxon has been confused with P. inter grifolia, but it is easily recognized by the white, tubular corollas, robust habit, and larger, less glandular leaves. In seed characters, P. robusta is similar to P. integrifolia var. texana. There appear to be several distinct entities included within this taxon. The material in north central Texas is disjunct in distribution from that in Presidio and Brewster cos., but additional material is needed to deter- mine if there are sufficient morphological characters to delineate the populations. Also, specimens from southern Colorado and adjacent New Mexico appear to be different and need to be investigated further. 29. Phacelia riipestris Greene Phacelia rupestris Greene. Leaflets 1:152. 1905. Holotype: New Me.xico: Sierra Co.: south end of the Black Range, 1 mi west of Hills- boro. 25 June 1904. O. Metcalfe 1012 (gh!); Isotypes (cvs. ny, pom. uc. uc, us, us, us). Phacelia congesta Hook. var. rupestris (Greene) Macbride. Contr. Gray Herb. 49:25. 1917. . '-H»^*V Fig. 51. Dorsal and ventral view of the seeds of P. robusta (Macbride) Johnston. L. Hinckley 829 (ny). ATWOOD: PHACELIA CRErsULATAE C.ROUP 179 Map 23. Southern Kansas. Texas, and ad- joining states. Distribution of P. robusta (Mac- bride) Johnston. Perennial plants (sometimes flowering the first year), 1-6 dm tall; stems 1-many, usually from a woody caudex. pubescence of whitish hairs, these hispid to setose and finer, sometimes glandular (these not multicellular glands); leaves pinnately compound, the terminal part incompletely 3-5 lobed and larger than the lower pin- nae, 1.5-10 cm long, 1-5 cm wide, setose to densely pilose; inflorescence of ter- minal compound scorpioid c}Tnes, individ- ual oTtnes 1.5-4.5 cm long, pedicels 1.5-2.5 mm long; sepals linear to oblanceolate, 3-4.8 mm long, 0.5-0.8 mm w^ide, setose to hirsute; corolla campanulate, white, 2-4 mm long and broad, lobes pubescent; sta- mens and style exserted up to ca. 2 mm, anthers dull blue; style 5-6 mm long, bifid over 1/2 its length, the lower un- branched portion pubescent; capsule oval to ovate, 2.6-3 mm long, 2.4-2.5 mm wide, finely pubescent with a few long hairs near the apex; mature seeds 4, brovsn, 2.1- 2.7 mm long, 1-1.1 mm wide, elliptical to oblong, reticulate scabrous, ventral sur- face excavated on both sides of the ridge (Fig. 52). Collections: 78 (1); repre- sentative: L. Goodding 2330 (gh, rm, uc): H. Rusby s.n. (cas. us); B. Maguire 11204 (bry);' a. Nelson 1248 (ny); C. Pringle 162 (mexu, ny, us); E. Wooton s.n. (ny); D. Dunn 8525 (uc). Habit.\t. — Growing on coarse, sandy soil and gravel bars, moist shady crevices of limestone cliffs, and ledges in canyons and arroyos, at 2,100 to 6,500 feet ele- vation. Flowering commonly occurs from late June to late August but some- times as early as mid-March and as late as the last of October. Usually associated with Larrea, Acacia. Lippia. Fallugia. Chilopsis, and Opuntia at lower elevations and with Qucrcus. Cercocarpus. Juniperus, Pinus, and Fra.rinus at higher elevations. Distribution. — Southern New Mex- ico from Socorro Co. westward to Pinal, Pima, and Cochise cos.. Arizona, and southward to southwestern Texas and ad- jacent Mexico (Map 24). This entity is closely related to P. con- gesta but differs in having smaller, white corollas, shorter and fewer flowered cymes, a perennial habit, and the ab- scence of multicellular glands. At times it apparently flowers in the first year, at least in the more southern parts of its range. 30. Pliacelia scariosa T. S. Brandegee Fig. 53 Phacelia scariosa T. S. Brandegee. Proc. Calif. Acad. Sci. 2:185. 1889. Holotype: Mexico: Baja California: Lower California, 12 Janu- ary 1889, Brandegee s.n. (uc!); duplicates (ny, us). ► Fig. 52. Dorsal and ventral view of the seeds of P. rupestris Greene. W. Eggleston 16341 (us). 180 GREAT BASIN NATURALIST Vol. 35, No. 2 • I ^ Map 24. Southeastern Arizona, New Mexi- co, southwestern Texas, and adjoining Mexico. Distribution of P. rupestris Greene. Plants annual, 0.5-4 dm tall; steins branching from the base (sometimes dichotomously), finely glandular and pi- lose; leaves ovate to oblong, deeply cleft to more often pinnately divided with 3-5 pinnae, terminal leaflet usually trilobed and larger than the other leaflets, 1.5-8 cm long, 1-5.5 cm wide, strigose and some- what glandular; inflorescence paniculate to racemose, glandular to villous, cymes elongate, open, up to 11 cm long, pedicels filiform, 3 mm long in flower, up to 8 mm long in fruit; corolla broadly cam- panulate, bluish to lavender, with white throat and tube, 4 mm long, 4-5 mm wide, pubescent; sepals obovate, 2-3 mm long in flower, becoming broadly obovate and conspicuously enlarged and scarious in fruit, 4.5-7.6 mm long, 2.3-3.7 mm wide, glandular to villous; stamens and style exserted 1-2 mm, style cleft 1/3 its length, lower 1/3 pubescent; ca])sule globular, 3.3-3.7 mm long, 2.6-3.5 mm wide, glan- dular to pilose; mature seeds 4, brown, el- liptical, 2.5-3 mm long, 1.2-1.3 mm wide, ventral surface di\ided by a prominent ridge, pitted, margins corrugated, the ridge corrugated on one side, dorsal surface curved, pitted (Fig. 54). Collections: 42 (0); representative: C. Orcutt 13 (cas, NY, us); M. Jones 24069 (pom, rm); T. Brandegee s.n. (uc); R. Moran 3890 (uc); D. Porter 236 (cas, mexu); F. Shreve7023 (us). Habitat. — Sandy, gravelly washes, rocky hillsides, and lava flows from 200 to 5,000 feet elevation. Apparently a winter annual, flowering from late October to mid-June. Distribution. — Lower California and adjacent Sonora, Mexico (Map 25). Phacelia scariosa is apparently related to P. pedicellata and is discussed under that species. Fif,'. 53. Phacelia scariosa T. S. Brandegee. Wiggins 7887 (us). June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 181 1884, Lemmon and wife (uc, uc. us); Sun- set Mountain, Flagstaff. 21 August 1915, H. Rusby s.n. (ny). Phacelia macdougalii Heller in Brand. Das Pflanzenreich IV. 251:80. 1913. as synonym. Annual plants, 1-3.4 dm tall; stems erect, simple or branched at the base, with simple setose, hirsute and multicellular Fig. 54. Dorsal and ventral view of the seeds of P. scariosa Brandegee. I. Johnston 3884 (c.^s). *^^ :Sj^ Map 25. Baja California and Sonora. Mexi- co. Distribution of P. scariosa Brandegee. 31. Phacelia serrata Voss Fig. 55 Phacelia serrata Voss. Bull. Torr. Bot. Club 64:88-89. 1937. Holotype: Arizona: Coco- nino Co.: San Francisco Mountains, May- October 1900, C. A. Purpus 8064 (pom!); Isotypes (uc, us). Paratypes: Arizona: Coco- nino Co.: vicinity of Flagstaff, 15 July 1898. Macdougal 288 (ny, rm, us, us); volcanic scoria, San Francisco Mountains, September Fig. 55. Phacelia serrata Voss. J. T. Howell and G. True 45184 (c.\s). 182 GREAT BASIN NATURALIST Vol. 35, No. 2 glandular hairs; leaves lanceolate, serrate, dentate to shallowiy lobed, 1.4 cm long, 0.5-2 cm wide, with a basal leaf cluster, especially when young, gradually reduced upward, the upper sessile or nearly so, the lower with a petiole up to 1.5 cm long, setose, hirsute and glandular; inflores- cence of compound scorpioid cymes, se- tose, puberulent with multicellular stip- itate glands, pedicels up to 1 mm long; sepals elliptical to oblanceolate, more or less keeled at the base, 3.5-6.5 mm long, 1.5-2 mm wide, setose, puberulent and stipitate-glandular; corolla rotate (appear- ing tubular in some pressed specimens), blue to light violet, 3-4 mm long and broad, pubescent; stamens and style ex- serted, style bifid 3/4 its length, lower 1/4 pubescent; capsule subglobose, 2.8- 3.5 mm long, 2-2.5 mm wide, glandular and puberulent; mature seeds 4, elliptical to oblong (sometimes unequally so when one of the margins is involute), dark brown, 3-3.2 mm long, 1-1.3 mm wide, excavated and divided by a prominent ridge, the ridge corrugated on one side, the margins corrugated, pitted and often one or both involute, dorsal surface smooth and shiny to somewhat dull, sometimes faintly pitted, the tip and margins darker for part of their length (or at least dif- ferent in appearance from the rest of the dorsal surface). Fig. 56. Collections: 15 (2); representative: .1. Hill s.n. (us); L. Goodding 1526 (uc); D. Atwood 2586 (bry); D. Dunn 12644 (rsa); H. Hansen 615 (rm); J. Howell and G. True (bry. cas) . Habitat. — Confined to volcanic scoria slopes of open yellow pine forest and ju- niper flats and hills. Flowering from late June to mid-September. 5,900 to 7,150 feet. Distribution. — In and around Sunset Crater National Monument and San Francisco Mountains north of Flagstaff. Coconino Co., Arizona. The relationships of this entity to other taxa in this group are questionable at the present time. Part of the paratype ma- terial cited by Voss (1937) belongs to P. palmeri. The specimen in question is Palmer 335 (gh, ny) and was probably collected in southern Utah or the extreme northern part of Arizona in Mohave Co. The label bears the data southern Utah- northern Arizona. Brand (19H) inad- vertently listed P. macdougalii as a syno- nym of P. integrifoUa Torr. even though it was only a manuscript name and had never been published. I have seen the specimens in question and conclude that they are referable to P. serrata Voss. Voss (1937) described P. serrata but failed to mention P. macdougalii. 32. Phacelia splendens Eastwood Fig. 57 Phacelia splendens Eastwood. Zoe 4:9. 1893. Holotype: Colorado: Mesa Co.: Grand Junc- tion, 19 May 1892. A. Eastwood s.n. (cas?); Isotypes (uc. uc, us. sketch at ny). Phacelia glandulosa Nutt, subsp. splendens (Eastwood) Brand. Das Pflanzenreich IV, 251:83. 1913. Plants annual, 0.5-2.7 dm tall; stems erect, simple or branched leafy, puber- ulent, with scattered stipitate-glandular hairs; leaves pinnatifid, 2-7.5 cm long, 0.7- 4 cm wide, petiolate, leaf blade essen- tially glabrous (pubescent only on the pet- iole and rachis or lower portion of the pinnae); inflorescence terminal on each branch and the main stem, cymes compact and densely flowered, pedicels short but lengthening to as much as 1.7 mm in fruit, slightly more pubescent than the stem; sepals linear to narrowly oblanceolate, 2.5- 3 mm long in flower, 4-4.4 mm long in fruit, 0.6-1 mm wide, hirsute and with a few scattered glandular hairs; corolla cam- panulate, the lobes bright blue, the tube yellowish, 4-8 mm long and broad, glab- rous to sparsely pubescent; stamens and style exserted 7-11 mm, the filaments blue, anthers yellow, style bifid ca. 2/3 Fig. 56. Dorsal and ventral view of the seeds of P. serrata Voss. J. T. -Howell and G. True 45184 (bky). ATWOOD: PHACELIA CRENULATAE CROUP 183 Fig. 57. Phacelia splendens Eastwood R. Barneby 12797 (cas). feet elevation. Commonly associated with Atriplci\ but in Mesa Verde National Monument it was collected in pinyon- juniper. Mid-May to mid-July. Distribution. — Known only from western and southwestern Colorado and northwestern New Mexico (Map 26). This species is related to P. corrugata and P. utahensis and may be a link be- tween the two complexes. It can be dis- tinguished from the former by its nearly glabrous and more-divided leaves, yellow- ish corolla tube, and different-textured and less-corrugated seeds. From the latter, it differs in having a shorter and less- glandular indument, a less-robust and less- branched habit, and different seeds. The seeds were reported by both Eastwood (1893') and Voss (1937) as lacking cor- rugations. Observations of mature seeds of P. splendens demonstrate that there definitely are evident corrugations on one side of the ridge and sometimes on one of the incurved margins. 33. Phacelia utahensis Voss Fig. 59 Phacelia utahensis Voss. Bull. Torr. Bot. Club 64:135. 1937. Holotype: Utah Sanpete Co.: Gunnison. 7 ,Iune 1910, M. Jones s.n. (pom). Plants stout, erect annuals, 0.8-5.8 dm tall; stems usually simple, sometimes branched at the base, brow'nish to yellow^- ish, densely glandular and finely pubes- cent; leaves linear to narrowly lanceolate, strigose to ciliate on the margins, wdth scattered glands (especially the upper), 1.5-12 cm long, 0.5-1.5 cm wide, the mar- its length, the undivided portions puber- ulent and glandular; capsule subglobose, 4-4.5 mm long, 3-3.5 mm long, 1.5 mm wide, finely favose, the ventral surface ex- cavated on both sides of the ridge, the ridge with evident corrugations on one side, the margins more or less revolute (Fig. 58). Collections: 23 (4); representa- tive: W. Weber 7509 (cas, rm, rsa, uc); D. Atwood 2532 (bry) ; D. Atwood and L. Higgins 1814 (bry); L. Iliggins 3302 (bry, wts); S. Welsh 756 (bry); R. Bar- nebv 12743 (cas, ny, rsa); E. Payson 671 (gh). Habitat. — Apparently confined to the Mancos Shale formation, 4,500 to 6,000 Fig. 58. Dorsal and ventral view of the seeds of P. splendens Eastwood. D. Atwood and L. Higgins 1814 (bry). 184 GREAT BASIN NATURALIST Vol. 35, No. 2 1 — ' N 1^ ,...../-' r- / — ^ -' 1 I i 1 1* .--f--. 1 <' 1 V--^. — •« y ■*--\y*-i /' /^ -| r- / "v •/ r--^- • / /' • / •• Map 26. Southwestern Colorado and adjoin- ing states. Disti'ibution of P. splendens Eastwood. gins often revolute, crenate, undulate to irregularly dentate, basal ones petiolate and dense, the upper sessile, aiu-iculate to cordate; inflorescence thyrsoid, u]) to 3.4 dm long, often with a few lateral, leafy inflorescence branches below, stipitate- glandular and finely pubescent, cymes mostly in pairs, (or 1-3), up to 4 dm long in fruit, densely flowered, the ped- icels, 1-1.5 mm long; sepals oblanceolate, 3-4 mm long, 0.8-1.1 mm wide, glandular and hirsute; corolla rotate to campan- ulate, the lobes bluish to violet, the tube yellowish, ca. 3-4 mm long, ca. 6 mm broad, glabrous; stamens exserted 9-10 mm, filaments violet, anthers yellow; style exserted ca. 10 mm, bifid 3/4 its length, the lower 1/4 setose and glandular; cap- sule globose to subglobose, 3.5-4.1 mm long, 2.6-3.5 mm wide, glandular and setose; mature seeds 4, elliptical, dark (reddish), the dorsal surface faintly pitted. the ventral surface excavated on both sides of the ridge, often lighter than the dorsal surface, pitied with the markings in the excavations longer (transversely) than those of the ridge or margins, the ridge .sometimes faintlv corrugated on one '\!b/=" Y J/'*^. Fig. 59. PhaccUa utahcnsis Voss. D. wood and L. Higgins 1624 (bry). At- side. Collections: 22 (13); representative: .1. Howell and G. True 44640 (bry, cas) ; L. Higgins 1624 (bry); D. Atwood 1520, 1893, 1835, 1684, 1892, 1894, 1895, 1526, 1519, 1518, 1525. 1528, 1527 (bry). Habitat. — Endemic to the Arapian Shale formation. Distribution. — Sevier and Sanpete COS., Utah, from 5,500 to 5,700 feet, April to June (Map 27). 34. PhaccUa vossii Atwood Fig. 60 PhaccUa vossii Atwood. Rhodora 74(800) :462- 465. 1972. Sierra Madre Oriental, calcite and limestone hills bej'ond Pablillo toward Santa Clara, ca. 15 mi southwest of Galeana, 18 July 1934, C. H. Mueller 1075 (gh!); Isotype (mexu). Paratype: Mexico: Nuevo Leon: Hacienda Pablillo, Galeana. 13 August 1936. M. Taylor 156 (ny, uc). Peremiial ]>lants, 2.1-7.^ chn high; stems erect from a woody caudex. with hirsute to setose and stipilatf^-ghuukdar hairs; leaves June 1975 ATWOOD: PHACELIA CRENULATAE GROUP 185 N \ Map 27. Central Utah in Sanpete and Sevier counties. Distribution of P. utahensis Voss. linear to lanceolate, 2-11 cm long, 0.5-2.2 cm wide, revolute, ventral surface heavily glandular, dorsal surface hirsute, with scat- tered stipitate-glandular hairs, margins dentate to irregularly toothed, petiolate, the petiole up to 1.5 cm long; inflorescence axillary to terminal, scattered along the stems for as much as 1/2 its length or less, consisting of simple to compound scorpioid cymes, individual cymes up to 7.5 cm long in fruit, pedicels up to 2 mm long in fruit, glandular and hirsute; co- rolla tubular to short campanulate, pale lavender (rarely white), 5.5 mm long; sepals oblanceolate to spatulate, 4.6 mm long, 1.5-2 mm wide, glandular and hir- sute; stamens exserted, filaments purple, anthers greenish blue; style exserted ca. 2 nun longer than the stamens ca. 9 mm long, bifid for 3/4 its length, lower 1/4 pubescent; capsule ovate, 3.7 min long, 2.5-2.7 mm wide, glandular and strigose; inmiature seeds 4, elliptical to oblong, brown, 2.5-3.1 mm long, 1-1.4 mm wide. Collections: 4 (1); representative: known only from the type collections and D. At- wood and ,1. Reveal 5985, 25 September 1973 (bry). Fig. 60. Phacelia vossii Atwood. G. Mueller 1075 (me.xu). 186 GREAT BASIN NATURALIST Vol. 35, No. 2 Fig. 61. Phaceiia welshii Atwood. D. Atwood 2605 (bry). June 1975 ATWOOD: PIIACF.LIA CRENULATAE (IROUP 187 1 Iabitat. — Apparently endeiriic to cal- (itc and limestone soils. Distribution. — Known only from the type locality near Galeana in the state of Nuevo Leon, Mexico. The corollas of the type specimens have faded to white, which is a common oc- currence in several species of this group. This taxon appears to be related to /•*. pinnatifida Griseb., which, according to Brand (1913), occurs m the Andes of South America in Peru, Bolivia, and Ar- gentina. 35. Phacelia ivelshii At wood Fig. 61 Phacelia welshii Atwood. Rliodora 74(800) :465- 468. 1972. Holotj'pe: Arizona: Coconino Co.: along highway 89 iust north of Gray Mountain, 19 May 1970. D. Atwood 2608 (bry!); Isotypes (.ariz, asc, b, bry, c.-\s, colo, DIXIE, GH. NY, POM, RM, RS.\, US, US, UT, UTC, WSC, WTS). Annual, 1-5.5 dm tall; stems more or less yellowish to green, simple or branched, leafy, hirsute and densely co\ered with multicellular stipitate glands; leaves ob- long to lanceolate, 1.5-8 cm long, 0.5-2.7 cm wide, hirsute and densely glandular, the margins often revolute, undulate and dentate, the basal leaves clustered, petio- late, the petiole 2 cm long or less, cauline leaves sessile or nearly so, often cordate at the base; inflorescence of compound scorpioid cymes, these terminal at the ends of the main stem and lateral branches, densely glandular and hirsute, the individual cymes congested, but loos- ening in fruit, up to 10 cm long, pedicels up to 1.5 mm long; sepals spatulate to oblanceolate, 3.5-4.5 mm long, 0.7-1.7 mm wide, hirsute and stipitate-glandular; co- rolla campanulate, purplish to blue, 5-6 mm long and broad, pubescent; stamens exserted ca. 8-10 mm, filaments the same color as the corolla, anthers yellow; style exserted ca. 8 mm, bifid 3/4 its length, the lower 1/4 setose and glandular; cajj- sule oval, 3-3.2 mm long, 2.9-3.1 mm wide, hirsute and glandular; mature seeds 4. oblong, brown ventrallv to reddish dor- sally, pitted, 2.8-3.4 mm long, 1.3-1.5 mm wide, the ventral surface lighter than the dorsal surface, the ridge corrugated on one side, the margins corrugated and more or less revolute. Collections: 15 (6); repre- sentative: D. Atwood 2591 (bry. cas); J. Howell 24397 (c..\s) ; D. Atwood 2598 (aRIZ, B, bry, CAS, COLO, DIXIE, GH, NY) ; D. Demaree 43982 (uc); D. Atwood 2601 (asc. bry. c.\s. lts, wsc). Habitat. — Red shale formation. Distribution. — Coconino Co., Arizona. Phacelia welshii is probably most closely related to P. utahensis and P. corrugata. It can be distinguished from the former by its broader, dark brown seeds, long (up to 1.3 mm long), flattened, multicellular, stipitate glands, and broadly lanceolate leaves. The more open inflorescence, cor- rugated seeds, and broader, shorter, densely glandular leaves easily separate P. welshii from P. corrugata. Appendix I Synonyms The following is a list of synonyms in the Phacelia Crenulatae group. The names in the left column are the synonyms, and the number to the right is the reference to the numbered taxa in the present treatment. EUTOCA E. glandulosa Hook 17 PHACELIA P. arenicola Brandegee 19a P. conferta D. Don .■ 10 P. congesta yar. dissecta Gray 10 P. congesta yar. rupestris (Greene) Macbride 29 P. congesta yar. typica Voss 10 P. dissecta (Gray) Small 10 P. crenulata yar. amhigua (Jones) Macbride 3a P. crenulata var. bakeri Biand 7 P. crenulata yar. corrugata (Nels.) Bi-and .... 12 P. crenulata yar. funerea Voss in Munz .... 14b P. crenulata yar. minutiflora (Voss) Jeps 3b P. crenulata yar. vulgaris Brand 14b P. depauperata W. & S 26 P. deserta Nels 17 P. foetida Goodding 24 P. glandulosa Gray in Brand, pro syn 26 P. glandulosa Hemsley 13 P. glandulosa ssp. eu-glandulosa Brand var. australis Brand, in part 15 P. glandulosa ssp. eu-glandulosa Brand yar. australis Brand, in part 7 P. glandulosa ssp. eu-glandulosa Barnd yar. deserta Brand 17 P. glandulosa ssp. eu-glandulosa 1 P. glandulosa ssp. splendens (Eastwood) Brand 32 P. glandulosa yar. neornexicana (Thurber ex Torr.) Gray 22 P. intcgrifolia var. arenicola (Brandegee) Brand ..._ 19a P. intcgrifolia var. palmeri (Torr. ex /Wats.) Gray 24 P. integrifolia var. rohusta Macbr 28 P. intermedia Wooton, in part 9 P. invenusta Gray 9 188 GREAT BASIN NATURALIST Vol. 35, No. 2 P. macdougalli Heller in Brand, pro. syn 31 P. neomezicana var. alba (Rydberg) Brand .. 1 P. neomezicana var. coulteri (Greenman) Brand - ■- 13 P. neomezicana var. coulteri subvar. folisis- sima Brand 1 P. neomezicana var. eu-neomezicana Brand 22 P. neomezicana var. microphylla Bi-and 15 P. palmeri var. typica Voss 24 P. petiolata Johnston 23 P. popei var. arizonica (Gray) Voss 6 P. popei var. similis (W. & S.) Voss 26 P. popei var. typica Voss 26 P. similis W. & W 26 P. tenuipes W. & S 8 P. tezana Voss 19b Appendix II Glossary Alveolate. Honeycombed; pits in the surface of the seed. Auriculate. With earlike appendages. Corrugated. Wrinkled or folded. Cymbiform. Boat shaped. Cyme. A detenninate flov^er cluster in which the first flower is terminal on the main axis and the central flowers open first. Denticulate. Slightly and finely toothed. Favose. Honeycombed; pits in the surface of the seeds. Fimbriate. Fringed with elongate, slender pro- cesses or lobes on the margins of the corolla lobes. Geminate. In pairs, as regarding the seeds. Glandular. A globose-secreting stinicture borne on the surface and estipitate. Gypsiferous. Containing gypsum. Hirsute. Pubescent with stiff, coarse hairs. Hispid. Pubescent with long, very stiff hairs, these able to penetrate the skin. Pilose. Pubescent with soft, slender hairs point- ing the same direction as if combed. Pitted. Having little depressions or pits. Puberulent. Pubescent with very short hairs, not stiff. Reticulate. Net-veined. Revolute. Rolled backward from both margins, toward the inside. Scabrous. Rough to the touch owing to the pres- ence of short stiff hairs. Scarious. Thin, dry. and membranous, not green. Scorpioid. A unilateral inflorescence circinately coiled in bud and anthesis. Setose. Pubescent with short, rather stiff hairs, these not able to penetrate the skin. Stipitate-glandular. A globose, stipitate, secre- tory structure borne on the surface of vegeta- tive parts. Strigose. Pubescent with short, straight appressed hairs. Tuberculate. Having small knoblike projections. Villous. Pubescent with long and weak, tangled, but not matted, hairs. Bibliography Abrams, L. 1954. Illustrated flora of the Pa- cific States. Stanford Univ. Press, Stanford, Calif. Bailey, L. H. 1949. Manual of cultivated plants. The Macmillan Company, New York. Ballion. H. E. 1890. Hydrophylleae in His- toire des Plantes 10:397-402. Benson, L. 1962. Plant taxonomy. The Ronald Press Company. New York. Bentham, G. 1837. Review of the order Hy- drophyllaceae. Trans. Limi. Soc. 17:267-282. , and J. D. Hooker. 1876. Genera Plan- tarum. Vol. I. Reeve and Company, London. Blake. S. F.. and A. C. Atwood. 1942. Geo- graphical guide to the floras of the world. U.S. Govenmient Printing Office, Washing- ton, D.C. Bradley. R. A. 1950. The vascular flora of Moffat County, Colorado. 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Colorado Ag- riculture College Expt. Sta.. Ft. Collins. . 1913. Studies on the Rocky Mountain flora. Bull. Torr. Bot. Club 40:479. . 1917. Flora of the Rocky Mountains and adjacent plains. Published by the author. New York. . 1954. Flora of the Rocky Mountains. Hafner Publishing Company, New York. Shinners, L. H. 1958. Spring flora of the Dallas-Fort Worth area. Published by the author, Dallas, Texas. Tidestrom, I. 1913. Novitate Florae Utahensis. Proc. Biol. Soc. Wash. 26:122. . 1925. Flora of Utah and Nevada. Contr. U.S. Natl. Herb. 25:1-665. , AND KiTTELi,. 1941. A flora of Arizona and New Mexico. The Catholic Univ. of America Press, Washington, D.C. ToRREY, J. 1827. Some account of a collection of plants made during a journey to and from the Rocky Mountains in the summer of 1880, by E. P. James. M.D. Hist. N.Y. 2:225. , .\ND A. Gray. 1838. Flora of North America. Vol. 1. Wiley and Putnam. Ne\v York. . 1854. Botany in Marcy. Explorations of the Red River. U.S. Dept. of Interior. Washington. D.C. . 1859. Botany in W. Emory, Report on U.S. and Mexican boundary survey. Vol. 2. U.S. Dept. of Interior. Washington, D.C. . 1860. Botanical appendix in J. C. Ives. Report upon the Colorado River of the west. C. Wendell. Printers, Washington, D.C. Voss, J. W. 1934. A revision of the Phacclia hispida group. Bull. So. Calif. Acad. Sci. 33: 169-177. . 1937. A revision of the Phacclia crcnu- lata group for North America. Bull. Torr. Bot. Club 64:80-81. 133-144. Watson, S. 1871. Botany U.S. Geol. Expl. of the fortieth parallel. Vol. 5. Government Printing Office. Washington. D.C. We.\ver, J. E. '1954. North American prairie. .Tolinsen Publishing Company. Lincoln, Ne braska. Welsh. S. L,, M. Treshow, and G. Moori . 1964. Guide to the common Utah plants Brigham Young Uniy. Press. Provo, Utali. Wilson. K. A. 1960. The genera of Hydro phyllaceae and Polemoniaceae in the south- eastern United States. Jour. Arnold Arb. 41:197-212. WooTON. E. O.. AND P. C. Standlev. 1913. De- scription of new plants preliminary to a re- port on the flora of New Mexico. Contr. U.S. Natl. Herb. 16:1. . 1915. Flora of New Mexico. Contr. U.S. Natl. Herb. 19:545. RODENT POPULATIONS, BIOMASS, AND COMMUNITY RELATIONSHIPS IN ARTEMISIA TRIDENT AT A. RUSH VALLEY, UTAH D. W. Nichols\ H. D. Smith"', and M. F. Baker' Abstract. — Three desert Artemisia tridentata communities in Rush Valley. Utah, were trapped for small rodents during the summer of 1970, and population densities were estimated for each popula- tion category using Lincoln's index. Animals were weighed and rodent biomass calculated by species throughout the summer. Population, biomass. and other data were then analyzed to gain an under- standing of the community relationships of the three study areas to each other as well as to the A. Iridentata community types of the Great Basin. Peromyscus maniculatus. Eutamias minimus, and Reithrodontomys megalolis were common to area 1, whereas P. maniculatus. E. minimus, and Perognathus parvus were common to areas 2 and 3. The peak estimated standing crops were 182.8 (74.0). 143.1 (57.9), and 129.7 g/acre (52.5 g/ha) for areas 2, 1, and 3 respectively. The population and biomass of area 2 peaked in midsum- mer, area 1 early summer, and area 3 late summer. Introduction Big sagebrush {Artemisia tridentata Nutt.) covers an estimated total area of 226,364 square miles (586,283 km-) in the Great Basin and associated areas of the western United States (Beetle, 1960) and is the most abundant plant species over much of this area. According to Hiro- naka (1963), ''A. tridentata has the widest distribution of all the sagebrushes and oc- curs across the entire moisture gradient of the sagebrush zone." Passey and Hugie (1962) found A. tridentata occupying a greater number of soil types than any other sagebrush species. A. tridentata is ecologically significant in that it provides not only food and cover for some species but competes against other desirable food and cover species. For economic reasons, however, many peo- ple consider sagebrush to be a highly un- desirable plant. As a result, much re- search done in the sagebrush community has been directly concerned with control- ling its spread and decreasing its abun- dance. Treatment resulting from such re- search has sometimes been temporarily effective. Some treated areas after 14 years may have more sagebrush on them than adjacent untreated areas (Johnson, 1969). The mean useful life cycle of spraying projects throughout Wyoming has been estimated to be about 15 years (Kearl, 1965). The economic importance and ecolog- ical impact of such control measures make research leading to an understand- ing of the A. tridentata community im- perative. The objective of this study is to establish baseline data by estimating the comparative small rodent density and bio- mass of A. tridentata communities in low- intermediate- and high-altitude desert areas of the Great Basin and using this estimation to compare the three commun- ities. Rush Valley, Utah, was chosen for the area of research because (1) it is a desig- nated grazing research area of the Inter- mountain Forest and Range Experimental Station, which funded the project; (2) a large part of the valley is covered by A. tridentata; (3) the data gathered will augment that of current and past research in the valley; and (4) the data collected will aid future studies and management of the valley. No literature relating rodent density with biomass or energy flow has been published concerning sagebrush commun- ities. There are works, however, that have been reported for other terrestrial com- munities. Densities of rodents have been studied in Rush Valley, Utah, primarily in piny on- juniper and reseeded areas (Baker, 1969). Woodbury (1955) re- ported on the small mammal distribution in Cedar Valley which borders Rush Valley on the east. Vest (1962) reported on the small mammal distribution in Dug- way Valley which borders Rush Valley on the west. Rodents of •sagebrush com- munities in both valleys were discussed. Although literature concerning biomass in A. tridentata communities is sparse, ^Department of Biology, Northland Pioneer College, Show Low, Arizona. ^Department of Zoology, Brigham Young University, Provo, Utah. ^Wildlife biologist, Intemiountain Forest and Range Experiment Station, Provn, Utah. 191 192 GREAT BASIN NATURALIST Vol. 35, No. 2 much descriptive material is available on the plant and its community type. A mor- phological life history of A. tridentata was written by Diettert (1938), and Beetle (1960) published a taxonomic and distrib- utional study of all the north American sagebrush taxa. Other references concern- ing A. tridentata communities in Utah may be found in Christensen (1967). Materials and Methods Study Areas Three 14.5 acre (5.87 ha) study areas located in the southern part of Rush Valley, Tooele County, Utah, were selec- ted and are described in Table 1. Area 1, elevation 5,100 feet (1,554 m), is lo- cated in the southwest quarter of Section 10, Township 7 south. Range 5 west, Tooele County, Utah. It contains irreg- ularly scattered A. tridentata interspersed with Chrysothamnus puberulus, the grass Distichlis striata, and bare ground. Dis- tichlis striata, an indicator of alkaline soil conditions, is abundant in a number of small areas with poor drainage as are numerous A. tridentata plants 1-3 inches (2.5-7.6 cm) tall. This short sagebrush is possibly stunted by an accumulation of soil salts. The area has a wash 1-2 feet (0.3-0.6 m) deep and five feet (1.5 m) wide that runs across the west side. Area 2, elevation 5,700 feet (1,737 m) is located in the northwest quarter of Sec- tion 35, Township 8 south, Range 6 west, Tooele County, Utah, and is characterized by scattered A. tridentata with some Sar- cobatus vermiculatus and much bare ground. The west boundary of this area is a dirt road, beyond which there is an extensive stand of S. vermiculatus mixed with the grass Agropyron cirstatuni. Area 3, elevation 6,500 feet, (1,981 m) is located in the west half of Section 4, Township 9, Range 5 south, Tooele County, Utah, and is covered with irreg- ularly scattered A. tridentata with a dense understory of a perennial lupine and var- ious grasses, principally Agropyron dasy- stachyuni and Sitanion Jiystrix. There is little bare ground except on the east and west borders which were exposed when adjacent land was cleared of A. tridentata and some Juniperus osteosperma in the fall of 1969. Vegetation sampling on all areas con- sisted of measuring (1) absolute ground cover, (2) percent species cover compo- sition, (3) frequency and density of pe- rennial species other than grasses, (4) fre- quency of all grasses, both annuals and perennials, lumped together, and (5) fre- quency of all annuals, except grasses, lumped together. A modified line-point method of sampling was used to determine cover, and small quadrats were used for frequency and density data collection (Cain and Castro, 1959). These data are available in Nicholes (1972). Trapping Each of the quadrat study areas, 14.5 acres (5.87 ha), was equally divided into 25 squares with a trap station located in the center of each square where three Sherman aluminum live traps were Table 1. General comparisons of the study areas. Characteristics Area 1 Area 2 Area 3 ^Soil, TYPE Slope Water runoff Erosion ^Climate Mean annual temp. Mean annual pupc. Principai. cover Principal rodents Elev.\tion deep silt-clay alkali soils of the arid & semi- arid valley bottoms 0-2% slow slight to moderate 51 F 8-10 inches Artemisia tridentata Chrysothamnus puberulus Distichlis stricta Peromyscus maniculatus Rutamias minimus Reithrodontomys megalotis 5.100 ft. (1,554 m) deep silt-loam soils of the semiarid valley bottoms 1-5% slow to medium high deep, loamy, dry soils of the dry subhumid alluvial fans 1-25%, most less tlian 10% slow to medium moderate 48 F 45-47 F 8-12 inches 12-15 inches Artemisia tridentata Artemisia tridentata Sarcobatus vermiculatus Lupine sp. various grasses various grasses Peromyscus maniculatus Peromyscus maniculatus Rutamias minimus Rutamias minimus Perognathus parvus Perognathus parvus 5,700 ft. (1.737 m) 6.500 ft (1.981 m) ^(Harvey and Woodward, 1969) June 1975 NICHOLS ET AL.: RODENT POPULATIONS 193 placed. The trapping stations were 160 feet (48.8 ni) apart. Rolled oats were used as bait. The traps were set in the afternoon, checked each morning, and closed until the afternoon resetting. Each trapping period covered five consecutive nights, every other week from 2 June to 14 August 1970 (Table 2). The three areas were trapped simultaneously for six trapping periods. To offset bias prior to each trapping period, the sequence in which each of the three areas would be checked and reset was determined ran- domly. This sequence was maintained throughout a trap }:)eriod. At the end of six trapping periods each area was "kill trapped" to compare with live trapping success during the previous periods. A "kill trapping" consisted of one night of live trapping with one live trap set at each of the regular trapping sta- tions, one live trap ])laced at the corners of each square, and one live trap placed at the middle of each side of the squares, for a total of 121 traps. Before the second trap night each live trap was replaced by two museum special snap traps for a total of 242 traps per quadrat. This trapping pattern continued three to four nights until the number of previously marked animals caught was reduced to none or nearly none. The "kill trapping" did not occur simultaneously for each area be- cause of the large number of traps in- volved (Table 2). Animals were toe clipped for identifica- tion. Data recorded for each individual animal handled during the study included (1) species, (2) sex, (3) age, (4) weight, (5) trapping station, and (6) notes con- cerning the animal's general condition, Table 2. Schedule of trapping periods. Areas 'Dates 1 2 3 Live Trapping 1. 2-6 June X X X 2. 15-19 June X X X 3. 29 June-3 July X X X 4. 13-17 July X X X 5. 27-31 July X X X 6. 10-14 Aug. X X X "Kill Trapping" 7. 24-27 Aug. X 8. 31 Aug.-4 Sept. X 9. 6-9 Sept. X ^Dates extend from the first mnming traps were rhocked to the last morning traps were checked during a trap period. such as pregnancy, parasitism, injuries, and others. Age classes of juvenile, sub- adult, and adult were determined primar- ily by pelage color and molt patterns; but the appearance of genitals, behavior of animals, and, in cases where age is ex- tremely difficult to determine, i.e., chip- munks, the weights of the animals were considered. Animals were weighed using a spring-operated scale accurate to the nearest 0.5 g. Population and Biomass Estimation For each species caught and recaptured in sufficient numbers, population esti- mates were made at the end of each trap- ping period using Lincoln's, Hayne's, and Jolly's indices (Giles, 1969). Estimates were made for ( 1 ) the total species pop- ulation, (2) the population of each age class within the species, and (3) the pop- ulation of each sex within the species. Nichols (1972) presents this data. Com- parison of the three estimators showed Lincoln's and Hayne's to be similar, but Lincoln's estimates were used in the bio- mass calculations. Population estimates for "kill trapping" periods were made after the first two nights of trapping, since two nights of "kill trapping" may have caused abnormal immigration into the areas. All other population estimates were calculated using five days of live trapping data. Species biomass was calculated by mul- tiplying the mean species weight by the estimated population number of that species for each trapping period. Previous experience had shown that animals re- peatedly caught during a trapping period tended to lose weight, likely due to trap- ping stress. To compensate for this, the mean weights were calculated two differ- ent ways: (1) using only the weight of an animal taken the first day during a given trapping period and (2) using the weight of an animal taken every day during its captivity for a given trapping period. The greatest of these mean weights for any population category was used in this study. In most instances method one was used. Results Four species of rodents were recaptured in sufficient numbers to be considered in 194 GREAT BASIN NATURALIST Vol. 35, No. 2 detail, but only three were prevalent in any given area. These species were — Area 1 Area 2 Area 3 Peromyscus maniculatus XXX Pcrognathus parvus X X Reithrodontomys megalotis X Eutamias minimus XXX The Lincoln population estimate for each species caught during each trapping period and study area is given in Table 3. The combined total number of rodents estimated, of all species considered, for each area is graphically shown in Figure 1. The following is a general breakdown of each area's trapping according to spe- cies. Detailed charts and tables of data are given in Nichols (1972). Area 1 Peromyscus maniculatus. The greatest number of P. maniculatus, 47 (Lincoln's estimate 50), appeared during trap period 3 with a male-female ratio of nearly 2:1. There were 16 juveniles, 30 subadults, and 1 adult. The least number, 24 (Lin- coln's estimate 25), composed of 3 ju- veniles, 17 subadults, and 4 adults with a male-female ratio of 5:3, appeared during period 1. The greatest mean species weight, 19.6 g, occurred during period 1 and the least, 15.5 g, during period 5. Reithrodontomys megalotis. In area 1 R. megalotis was captured least often of the main species. The greatest number, 14 (Lincoln's estimate 18), appeared during period 2 with a male-female ratio of 3:4. There were no juveniles, 2 sub- adults, and 12 adults. During periods 4, 5, and 6 no mice of this species were caught. The greatest mean species weight, 13.9 g, occurred in period 3 and the least, 11.1 g, in period 2. Eutamias minimus. The greatest num- ber of E. minimus, 33 (Lincoln's estimate 33), appeared during period 1 with a male-female ratio of approximately 3:2. There were 16 juveniles, 14 subadults, and 3 adults. The least number, 10 (Lin- coln's estimate 8), com})osed of 0 juv- eniles, 9 subadults, and 1 adult with a male-female ratio of nearly 1:1, appeared during period 6. The greatest mean spe- cies weight, 30.5 g, occurred during period 5 and the least, 28.4 g, during period 4. Area 2 Peromyscus maniculatus. The greatest number of P. maniculatus, 49 (Lincoln's estimate 43), appeared during trap period 6 with a male-female ratio of nearly 3:2. There were 5 juveniles, 38 subadults, and 6 adults. The least number, 13 (Lin- coln's estimate 12), composed of 6 ju- veniles, 5 subadults, and 2 adults with a male-female ratio of 5:8, appeared during period 2. The greatest mean species weight, 18.2 g, occurred during period 5 and the least, 15.3 g, during period 1. Perognathus parvus. In area 2 P. par- vus was captured least of the three main species. Both periods 5 and 6 yielded the greatest number, 10 (Lincoln's estimates 9 and 10), with male-female ratios of 7:3 and 3:1. The respective age distributions were 0 and 2 juveniles, 9 and 7 subadults, 1 and 1 adults, llie period of least cap- ture was period 1 when one adult female was captured. The greatest mean species weight, 18.7 g, occurred during period 2 and the least, 14.5 g, during period 3. Eutamias minimus. The greatest num- ber of E. minimus, 43 (Lincoln's estimate 46), appeared during period 2 with a male-female ratio of approximately 4:3. There were 20 juveniles, 16 subadults, and 7 adults. The least number of chip- munks, 14 (Lincoln's estimate 13), com- posed of 1 juvenile, 12 subadults, and 1 adult with a male-female ratio of nearly 4:1, appeared during period 3. The greatest mean species weight, 31.1 g, oc- curred during period 5 and the least, 27.7 g, during period 1. Area 3 Peromyscys maniculatus. The greatest nimiber of P. maniculatus, 55 (Lincoln's estimate 49), appeared during trap period 3 with a male-female ratio of approx- imately 3:5. There were 16 juveniles, 31 subadults, and 8 adults. The least num- ber, 29 (Lincoln's estimate 26), composed of 2 juveniles, 18 subadults, and 9 adults with a male-female ratio of 3:4, appeared during period 1. The greatest mean spe- cies weight, 19.9 g, occurred during period 1 and the least, 17.5 g, during period 6. Perognathus parvus. The greatest num- ber of P. parvus, 47 (Lincohi's estimate 54), appeared during period 6 with a June 1975 nichols et al.: rodent populations Table 3. Estimated population numbers and biomass for the three study areas. 195 Trap Period Lincoln's N no./ 14.5 acres (5.86 ha) Estimated biomass (g/14.5 acres) (5.86 ha) Species Area 1 Area 2 Area 3 Area 1 P.M. E.M. P.P. R.M. 25 33 58 P.M. E.M. P.P. R.M. 44 39 18 101 P.M. E.M. P.P. R.M. 50 31 1 P.M. E.M. P.P. R.M. 82 48 23 71 P.M. E.M. P.P. R.M. 41 5 46 P.M. E.M. P.P. R.M. 35 8 43 P.M. E.M. 33 12 45 P.M. E.M. P.P. P.M. P.P. 112 67 101 490.0 943.8 833.9 488.4 337.2 825.6 Area 2 275.4 941.8 2650.4 1067.5 85.2 1152.7 Area 3 517.4 530.0 51 1433.8 1217.2 1047.4 36 752.4 199.2 698.4 1123.2 1311.0 40 199.8 18.7 836.0 76 2075.8 1528.9 1534.4 49 875.0 283.2 886.9 914.5 374.4 29 13.9 29.0 559.7 78 1803.4 686.6 1446.6 55 811.2 434.2 1078.0 653.2 848.4 36 148.0 655.2 91 1464.4 1430.6 1733.2 58 635.5 491.4 1044.0 2 152.5 746.4 68.4 38 162.9 767.6 98 788.0 1400.7 1880.0 47 591.5 • 722.4 822.5 242.4 1770.0 54 158.0 1015.2 1837.7 705.2 91.5 794.3 1591.0 Key: P.M. — Peromyscus maniculatus; E.iNI. — Eutamias minim.i \\_y\—lirillnn,l.., male-female ratio of approximately 7:5. There were 4 juveniles, 21 subadults, and 22 adults. The least number, 19 (Lin- coln's estimate 25), composed of 0 ju- veniles, 3 subadults, and 16 adults with a male-female ratio of nearly 2: 1 appeared during period 1. The greatest mean spe- cies weight, 21.2 g, occurred during period 1 and the least, 18.2 g, during ]:)eriod 4. Eutamias minimus. In area 3 E. mini- mus was captured the least of the three main species. The greatest number, 6 (Lincoln's estimate 0), appeared during period 6 with a male-female ratio of 1:2. There were 0 juveniles, 3 subadults, and 3 adults. No chipmunks were caught during ])eriod 3. The weight, 39.0 g, of one adult female, the only animal caught 196 GREAT BASIN NATURALIST Vol. 35, No. 2 during period 1, represents the greatest mean species weight. The least mean species weight was 29.5 g and occurred during period 3. Estimated Small Rodent Biomass The estimated biomass of each species for every period and study area, along with total biomass per area, is given in Table 3. In addition Table 3 lists the Lincoln population estimates used in the biomass computations. Kill-trap data were not included in calculating the greatest, least, and mean area biomass as given in the following description of ro- dent biomass by areas, but they are includ- ed in Table 3. The kill-trap biomass for any of the three areas fell within the limits of the estimates for that particular area from the six previous trapping pe- riods. Area 1. The greatest estimated small rodent biomass, 143.1 g/acre (57.0 g/ha), for area 1 occurred during period 2. The least, 54.3 g/acre (22.0 g/ha), occurred during period 5. The mean biomass of area 1 over the six trapping ])eriods was 96.5 g/acre (39.1 g/ha). The mean was approached during periods 1 and 4. Pe- riods 5 and 6 were well below the mean as was kill-trap period 7. Area 2. The greatest biomass, 182.8 g/acre (74.0 g/ha), for area 2 occurred during period 6. The least, 47.4 g/acre (19.2 g/ha), occurred during period 3. The mean biomass of area 2 over the six trapping periods was 102.5 g/acre (41.5 g/ha). The mean was approached during periods 2, 4, and 5. Area 2 had the greatest and the lowest biomass of all three areas during the study. Area 3. The greatest biomass, 129.7 g/acre (52.5 g/ha), for area 3 occurred during period 5 and the least, 72.2 g/acre (29.2 g/ha), occurred during period 1. The mean biomass of area 3 over the six trapping periods was 109.9 g/acre (44.5 g/ha). The mean was approached during period 2 and kill-trai) period 8. Discussion To understand an A. tridentata com- munity a knowledge of the associated vertebrates and vegetation must be ob- tained. An analysis of each A. tridentata study area is thus important in a discus- sion relating the rodent population and biomass of the individual areas to each other to establish the picture for the com- munity type. Peromyscus majiiculatus and E. mini- mus were two of the three predominant rodent species in each area. This could be exj)ected for P. maniculatus because of its geographic range over most North American biomes (Burt and Grossenhei- der, 1964; King, 1968), including sage- brush regions of the Great Basin. Eutam- ias minimus also has a broad geograph- ical range, including the Great Basin and much of Canada. Its appearance on all study areas should also be expected since it is characteristic of sagebrush commun- ities (Gordon, 1943; Burt and Grossen- heider, 1964). Out of 43 vertebrate spe- cies noted during the study, 15 were com- mon to all three areas, an indication that the study areas were similar. Table 1, however, indicates that notable differ- ences existed between them. These dif- ferences resulted from the distribution of the 28 vertebrates ^vhich were not com- mon to the three areas (Nichols, 1972). Area 1 had 4, area 2 none, and area 3 12 unique species of vertebrates present. The large number of unique species for area 3 can be explained by the presence of a more diverse habitat than in the other areas. This di^'ersity was likely due to more favorable climatic and edaphic factors (Table 1). Area 2 was interme- diate in soil, slope, temperature, and pre- cipitation (Table 1). The lack of verte- brate species unique to area 2 also indi- cates that it was an intermediate area. Area 1 exhibited the least vertebrate diversity, area 2 was transitional but closer to area 1, and area 3 was the most diverse. Most of the rodent biomass for areas 1 and 2 came from E. minimus, while their (ontribution in area 3 was small (Table 3). This supports the ap- parent (loser relationship of area 2 to area 1 . It should also be kept in mind that areas 1 and 3 are the farthest apart geo- grajihically and altitudinally, with area 1 (elev. 5,100 ft.; 1,554 m) near the valley floor, area 3 (elev. 6,500 ft.; 1.981 m) at the base of the valley-forming mountains, and area 2 (elev. 5,700 ft.; 1,737 m) on the benchland between. them. The ground cover of each area was doniin;il(Hl h\ A. tridentata. although each June 1975 NICHOLS ET AL.: RODENT POPULATIONS 197 area had a different predominant under- story species (Nichols, 1972). The di- versity pattern reflected was one of low plant species diversity for areas 1 and 2 and high diversity for area 3. Area 1 had 14, 2 had 15, and 3 had 30 plant species present. This was similar to the verte- brate diversity pattern for the respective j areas. This similarity in vertebrate and j plant diversity ])atterns was likely due j to the edaphic and climatic factors affec- j ting the plants which in turn affected the i vertebrates. The lack of complete simi- I larity between vertebrate and plant pat- terns, however, may have been due to the I physiognomic differences of the cover spe- cies present in the areas rather than the amount of diversity. It is suggested that the amount, distri- bution, and physiognomy of dominant vegetative cover in A. tridentata com- munities may have a greater influence than the diversity of cover species in de- termining vertebrate presence. Turner (1950) supports this idea, especially for Peromyscus distribution, in his study of 10 vegetative types, including 3 having A. tridentata as the dominant or co- dominant plant. Rosenzweig and Wina- kur (1969) have hypothesized from stud- ies in the lower-Sonoran desert scrub vegetation that "the spatial variations in density of some species [rodent] are re- sponses to spatial characteristics of their environment. Important among these en- vironmental characteristics tend to be mea- sures of the presence and/or absence of vegetation of various physiognomies." The absolute cover of A. tridentata in area 1 was 17.6 percent; 2 was 15.5 per- cent; and 3 was 23.2 percent. Percent cover composition was 44.2 percent (area 1), 48.0 percent (area 2), and 38. .0 per- cent (area 3). Area 2 had the least ab- solute cover of A. tridentata with the most bushes less than 2 feet (0.6 m) high but had the greatest percent cover composi- tion of A. tridentata for the three areas. This cover pattern may have been why area 2 had no unique vertebrates, thus functioning as a limiting factor to diver- sity. Area 1 had four species but was no more diverse than area 2 in terms of plant species, indicating that in this case cover type had a greater effect than did plant diversity. Area 3, in terms of A. tridentata cover, was opposite area 2. i Area 3 had the greatest absolute coverage of A. tridentata with most bushes greater than 2 feet (0.6 m) high but had the least A. tridentata cover composition for the areas. This showed the opposite ef- fect on vertebrate presence than the pat- tern in area 2. Instead of having no unique vertebrate species as in area 2, there were 12, including 2 rodents. The relationship for area 3, however, was not as pronounced because that area had nearly twice the plant species diversity of areas 1 and 2; and the greater plant species diversity may be responsible for the greater vertebrate diversity. If this is true, it is a direct reversal of the results obtained by Rosenzweig and Winakur (1969) in the lower-Sonoran desert scrub vegetation. They found that "the varia- tion in plant s])ecies diversity failed to ex- plain the \ariation in animal species di- versity and that some of the most faunal- ly diverse areas had the fewest species of plants." The three areas reported in this study, however, are in the Great Basin cokr desert, which varies considerably from the Sonoran hot desert. It is possible that the amount and distribution of A. tridentata cover is only important in determining vertebrate distribution in areas with low total absolute cover as in areas 1 and 2 but not in area 3. According to Pearson (1965a, 1965b) and Beatley (1969), primary productivity of A. tridentata communities peaks in late si)ring and early summer. Peaks in rodent populations in A. tridentata communities occur during the early summer to fall period (Turner, 1950; Sullivan, 1961). Trojan (1970) has shown that in a Polish grassland the rodent biomass increase during the summer is four times as great as during the winter and two times as great as during the spring. Summer and autumn increases accounted for 89 per- cent of the annual increase. He stated that "winter increases are of almost no importance to assessment of energy flow (3.2 percent)." His results may be appli- cable to the Great Basin sagebrush zone because its increased elevation could par- tially compensate for the higher latitude and much lower elevation of Poland. The Polish study was done in a grassland, but the areas studied in Rush Valley, Utah, were probably grassland before the valley's settlement (Christensen and Hutchinson, 1965). Winter production 198 GREAT BASIN NATURALIST Vol. 35, No. 2 may be important in some sagebrush areas, but no winter data were taken in this study due to inaccessibility. If (1) the annual peak primary pro- duction of A. tridentata communities was late spring and early summer, (2) ro- dent populations in A. tridentata com- munities peaked in the early summer to fall, and (3) there was little increase in rodent biomass during the previous winter months, then the population, mean species weights, and biomass data collec- ted during this study should illustrate the dynamic relationships of the three stud}' areas. Estimated Populations The total population estimates of all species are shown in Table 3 and Figure 1. Area 1 had a definite early summer population peak followed by a steady de- cline and a leveling off in the fall, but area 3 had three different population peaks during the summer with the high- est population occurring in late summer (approx. August 13). Area 3 had a grad- ual increase in rodent numbers, peaking in late sunmier (approx. August 13) and declining by September 3. Mean Species Weights Mean species weight is not only im- portant in calculating the estimated rodent biomass, but Walkowa (1970) pointed out the importance of species weights as an exploitation compensation mechanism in rodent j)opulations. He found that re- production operates as a compensating mechanism only if exploitation exceeds 31 percent. When exploitation was 0-30 percent, an increase in exploitation caused an increase in the production of biomass. In this study there appeared to be no un- usual predatory or disease exploitation of populations above 30 percent; thus the species were likely reacting to exploita- tion by increasing biomass without in- creasing reprodiutioii. This was further evidenced by the hu k of high-poi)ulation densities. The greatest estimated rodent density was eight rodents per acre (3.24 /ha) in area 2 during its summer peak. In addition to rodent weights varying with population exploitation, they also vary with the animal's dailv activities. Tevis (1955) showed that the £?ross bodv Are a 2 n |-| Trapping Periods Fig. 1. Number of rodents estimated in all areas. weight increase in chipmunks going from an empty stomach to a full stomach averaged 6-8 percent, whereas Evans (1949) observed that voles increased 20 percent in body weight within five min- utes after water consumption. It is evi- dent, therefore, that an accurate biomass estimation requires use of weights from the particular time and population being considered. Mean species weights used in this study were calculated from what was believed to be the most accurate weight according to the above criteria, but errors may have occurred. Area 1. The mean species weight of P. manicuhttus was greatest in the early spring and then oscillated between lower weights throughout the study. The high weight during period 1 reflected the pop- ulation structure at that time. There were few juvenile and subadult animals compared with subsequent periods. After [)(>rio(l 1 there were increased numbers of juNcniles emerging from the nests, causing a sharp decrease in mean species weight during trap period 2. For the re- maiiidof of the slud\- fe])rocluction caused June 1975 NICHOLS ET AL.: RODENT POPULATIONS 199 oscillations in the mean species weights between trapping periods. These oscil- lations were expected since P. maniculatus is polyestrous (Asdell, 1964). The E. min- imus mean species weights generally ap- peared to increase throughout the study to a peak near the end of the summer during trap period 5. This increase paralleled a gradual decrease in population numbers throughout the summer. The E. minimus reproductive pattern, one litter in the spring and subsequent growth of the young during the summer (Asdell, 1964), was responsible for the inverse relation- ship. No pattern was evident for R. mega- lotis because of insufficient captures. Area 2. The mean species weight pattern of P. maniculatus in area 2 oscillated for the same reason as their observed oscilla- tions on area 1. but the greatest mean species weight occurred in period 5. There was an apparent slow period in repro- ductive activity during June and July compared wdth area 1. This undoubtedly allowed the summer mean species weight of the population to peak later than in area 1. The mean species weights of E. minimus gradually increased dm-ing the summer because of their reproductive pattern. Too few P. parvus were caught in this area to show any definite patterns, but the same slow reproductive activity as noted in P. maniculatus on this area was also noted for P. parvus. Area 3. The mean species weight pat- tern of P. maniculatus showed reproduc- tion occurring throughout the summer. The P. parvus pattern in area 3 showed increased population numbers associated with decreased mean species weights throughout the study. This was exj^ected since P. parvus is polyestrous (Asdell, 1964). Too few E. minimus were caught on this area to show any definite pattern. Biomass The total estimated seasonal biomass in each area (Table 3 and Fig. 2) was similar to the population estimate for the area (Fig. 1). In terms of biomass, however, there was clearly a closer relationship be- tween areas 1 and 3 than had previously been proposed. Biomass in area 2 showed that secondary production was rather un- stable compared with areas 1 and 3. Area 2 2600 Area A 2400 / \ / \ 2200 . / \ 2000. /\ -r\ 1800 . / / \ \ ^, 1600- / / \/' ^\ / A~- ---'' \- \ \ UOO / ' \ Ar 1200- / \ / ^ / \ / 1000- \ / \ / \ \ 800 V 600 - 400 200 - Fig. 2. Estimated total small rodent biomass by trapping periods. Area 1. An early summer increase of 50 percent in rodent biomass was evi- denced in area 1 during a 13-day interval between trap periods 1 and 2. which made trap period 2 aj^pear as the peak period of summer rodent biomass production for area 1 (Fig 2). There was then a steady drop in biomass over a 42-day period fol- lowed by a leveling off. The data indi- cates, however, that the number of E. minimus and R. megalotis ma}' have been overestimated during period 2. There were 39 E. minimus estimated during period 2, but only 29 were handled. There were also 4 more R. megalotis esti- mated than were actually handled. A high number of unmarked animals being caught at the end of a trap period would cause a high unmarked-to-marked animal ratio to occur. This high ratio would in turn cause an overestimation of the popu- lation. This high ratio may be caused by immigration of animals into the area, by new animals emerging from their nests, or by various other factors. In this in- stance the possible overestimation may have been caused by new juvenile male chipmunks becoming available to the traps. If the actual number of rodents handled 200 GREAT BASIN NATURALIST Vol. 35, No. 2 during period 2 is used in the biomass calculation, then the summer peak on area 1 did not occur during trap ])eriod 2 but during trap period 3. The possible cor- rection is shown in Figure 3. The total pattern for the summer, however, is still one of an early peak in rodent biomass followed by a steady drop and then a leveling off for the remainder of the study. This is what could be expected for a com- munit}- in poor soil with low plant spe- cies diversity, as area 1 was earlier shown to be. Secondary production peaked early in the summer, after the observed peak primary production, thus placing greater demands upon the primary production of the community to support the increased secondary production. The plant com- munity, beyond its peak production and with very little diversification, likely could not produce more food for the iii- creased rodent population, so the rodent population rapidly declined to a level re- flecting the probable carrying capacity of the community and then remained there through the end of the summer. Be- cause of poor soil and other factors, the carrying capacity was low, approximately 55 g of rodent biomass per acre (22.27 1400 1200 1000 Trapping Periods Fig. 3. Estimated total small rodent bio- mass by trapping periods using possible correc- tions. g/ha). This low carrying capacity could liaAe supported no more than one E. min- imus and one P. mcmicidatus per acre (.24/ha). Area 3. The rodent biomass of area 3 showed an entirely different pattern than area 2 throughout the summer (Fig. 2). Secondary rodent productivity increased rapidly after spring reproduction and was then followed by a short period of reduc- tion before rising again. This reduction may have been due to a number of fac- tors but was most likely caused by cold temperatures. The P. parvus estimate for the period of biomass drop (trap period 3) was considerably less than the previous estimate. This rodent, as a protective be- havioral ada])tation, regularly goes into a state of torpor when it encounters cold (Bartholemew and Cade, 1957; Beer, 1961; Morrison and Ryser, 1962; Chew et al., 1965, 1967; Tucker, 1962, 1963, 1965a, 1965b, 1966). During the second and third nights of trap period 3 the coldest summer temperatures were re- corded for area 3, and four P. parvus that j)robably would ha^'e been caught had the nights been warmer were not recap- tured. This could have led to a low pop- ulation estimate and thus a low biomass estimate for the period. If this were the case, then the biomass of area 3 would have shown an increase from trap period 1 to its peak in trap period 5. The pos- sible correction is shown in Figure 3. At the end of the summer peak biomass grad- ually declined. Because of the diversity of the }ilant community in area 3, fa- vorable climatic conditions and primary production were jirobably sufficient to su])ply food for the gradual summer in- crease in secondary production. Area 2. The graphing of rodent biomass for area 2 showed no pattern similar to areas 1 or 3. It did, however, show an earl}' sininner decline in standing crop, a rapid reco^•ery with leveling off for a trap period, a midsummer biomass increase to a peak higher than in either areas 1 or ^, and a late summer decline in standing crop of the same rate as that which oc- curred in the early summer. Close examination of the data, how- ov(n-. for tli(> 1\. minituus popidation showed a large trap mortality during [x'riod f). This c auscd T.iiu oin's index to June 1975 NICHOLS ET AL.: RODENT POPULATIONS 201 overestimate the population. It estimated 20 more E. minimus than were actually handled, which would represent an over- estimation of 1,060.5 g in hiomass. Thus, at traj) jieriod 6 the high peak shown in Figure 2 would probably be much lower. The possible correction is shown in Fig- ure 3. With this correction the biomass was below that of area 3 for the same period. Closer examination of the data re- vealed another possible error. In trap period 9 there were 29 different E. min- imus handled, but because this period was a kill-trap period, the diurnal chipmunks were not caught during the one live trap night. As a result, none were marked, and no population estimates were made on day 2 of the kill-trap period. There were 17 different chipmunks handled on days 1 and 2 of this period. If the biomass of these rodents were added to the total bio- mass of this period, it would raise the total b}' 879.7 g. This possible correction is also shown in Figure 3. The above corrections change consider- ably the estimated summer biomass pat- tern of area 2. The pattern now becomes one of an early summer decline in bio- mass, a recovery period, a period of no increase or decrease in biomass, and a steady rise in biomass that may not have peaked before the study ended. In terms of stability the community of area 2 ap- peared to be less stable than those of areas 1 and 3. The corrected pattern showed that in terms of biomass area 2 was more similar to area 3 than area 1 as had been previously proposed. Conclusions It appeared that when the characteris- tics of ( 1 ) vertebrate presence and dis- tribution, (2) plant presence and distri- bution, (3) comparative rodent popula- tion numbers, and (4) comparative rodent biomass of the areas were considered, the three areas were in three different cli- matic or edaphically induced successional stages or conditions. Area 2 had the least diverse vertebrate presence. This may have resulted from the A. tridcntata cover being composed of small bushes represent- ing the greatest percent cover composi- tion of the three areas. Area 2 was the least stable in terms of the annual cycle for rodent populations and biomass in- crease, probably a result of the poor plant species diversity. Area 3 had the most di- verse vertebrate presence and was the most stable in terms of the annual cycle for rodent population and biomass in- crease. This may have been due to the A. tridentata cover pattern, which was op- ])Osite that found in area 2, and/or to the increased plant species diversity found in area 3. Area 1 appeared to be interme- diate, in its vertebrate presence and A. tri- dentata cover patterns, to areas 2 and 3. Its annual cycle in terms of rodent pop- ulation and biomass increase showed the earliest peak of the three areas. This ma}' have been due to poorer soil and climatic conditions causing an early peak in pri- mary productivity, a subsequent early peak in rodent productivity, and an ex- tended period through the rest of the sum- mer when conditions did not favor either primary or secondary production. Literature Cited AsDELL, S. A. 1964. Patterns of mammalian re- production. Cornell Univ. Press, Ithaca, New York. Baker, M. F. 1969. Unpublished data on rodent populations in the pinon-juniper zone of Rush Valley, Utah. Work-Unit #D.C. 101.2, U.S. Fish and Wildl. Sei^. B.\RTHOLOMEW, G. A., .\ND T. J. C.-^DE. 1957. Temperature regulation, hibernation, and aestivation in the little pocket mouse, Pe- rognathus longimembris. J. Mammal. 38: 60-72. Be.\tley, J. C. 1969. Biomass of desert winter annual plant populations in southern Nevada. OIKOS 20:261-273. Beer, J. R. 1961. Hibernation in Perognathus flavescens. J. Mammal. 42:103. Beetle, A. A. 1960. A study of sagebioish, the section Tridentatae of Artemisia. Wyoming Agric. Expt. Sta. Bull. 368:1-83. Burt, W. H., and R. P. Grossenheider. 1964. A field guide to the mammals. Houghton Mifflin Co.. Boston. Cain, S. A., and G. M. de Oliveire Castro. 1959. Manual of vegetation analysis. Harper and Brothers, New York. Chew, R. M., R. G. Lindbert, and P. Hayden. 1965. Circadian rhythm of metabolic rate in pocket mice. J. Mammal. 46:477-494. . 1967. Temperature regulations in the little pocket mouse, Perognathus longimem- bris. Comp. Biochem. Physiol. 21:487-505. Christensen, E. M. 1967. Bibliography of Utah botany and wildlife conservation. Brig- ham Young Univ. Sci. Bull., Biol. Ser., vol. 9, no. 1. Christensen, E. M.. and M. A. Hutchinson. 1965. Historical obsers'ations on the ecology of Rush and Tooele Valleys, Utah. Proc. Utah Acad, of Arts and Sci. 42:90-105. 202 GREAT BASIN NATURALIST Vol. 35, No. 2 DiETTERT, R. A. 1938. The morphology of Artemisia tridentata Nutt. Lloydia 1:3-74. Evans, R. C. 1949. A population study of house mice {Mus musculus) following a period of local abundance. J. Mammal. 30:351-363. GoLLEY, F. B. 1960. Energy dynamics of an old field community. Ecol. Monogr. 30:87-206. Gordon, K. 1943. The natural history and be^ havior of the western chipmunk and the mantled ground squirrel. College Press, Ore- gon St. College, Coi-vallis. H.ARVEY, J. L., .AND L. Woodward. 1969. Spe- cial soils report on eastern Tooele County. Utah. Soil Conserv. Ser. USDA, Portland, Ore. HiRONAKA, M. 1963. Plant-environment rela- tions of major species in sagebrush-grass vegetation of southern Idaho. Ph.D. dissei- tation. Univ. Wisconsin. Green Bay. (Diss. Abstr. 24:45.46.) Johnson, W. M. 1969. Life expectancy of a sagebrush (Artemisia) control project in cen- tral Wyoming. J. Range Manage. 22:177-182. Kearl, W. G. 1965. A survey of big sage- brush control in Wyoming: 1952-1964. Wyo- ming Agric. Expt. Sta. Mim. Cir. 217. 42pp. King. J. A. (ed.) 1968. Biology of Peromys- cus (Rodentia). Special Publication No. 2. The Amer. Soc. of Mammalogists. Morrison, P., and F. A. Ryser. 1962. Hypo- thermic behavior in the hispid pocket mouse. J. Mammal. 43:529-533. Nicholes, D. W. 1972. Small rodent popula- tions and biomass in three sagebrush com munities of Rush Valley. Utah. Master's thesis. Brigham Young Univ.. Provo. Utah. Odum, E. p. 1959. Fundamentals of ecologv. W. B. Saunders Co., Philadelphia. Passey. H. B., and V. K. Hugie. 1962. Sage- brush on relict ranges in the Snake River Plains and Northern Great Basin. J. Range Manage. 15:273-278. Pearson, L. C. 1965a. Primary productivitv in a northern desert area. OIKOS 15:211-228. . 1965b. Primarj' production in grazed and ungrazed desert communities of eastern Idaho. Ecology 46:278-285. ROSENZWEIG, M. L., AND J. WiNAKUR. 1969. Population ecology of desert rodent com- munities: habitats and environmental com- plexity. Ecology 50:558-572. Shelford, V. E. 1963. The ecology of North America. Univ. Illinois Press, Urbana. Sullivan, John O. 1961. Population structure of Peromyscus maniculatus in two areas in Green Canyon, Cache County, Utah. M.S. thesis. Utah St. Univ.. Logan. Tevis, L., Jr. 1955. Observations on chipmunks and mantled squirrels in northeastern Cali- fornia. Amer. Midi. Nat. 53:71-78. Trojan, P. 1970. Energy flow through a popu- lation of Microtus arvalis in an agrocenosis during a period of mass occurrence. Pages 267-279 in Energy flow through small mam- mal populations. Warszawa. Tucker, V. A. 1962. Diurnal torpidity in the California pocket mouse. Science 136:380-381. . 1963. The energetics of the torpor cycle in the California pocket mouse, Pe- rognathus californicus. Ph.D. dissertation. Univ. California Los Angeles. (Diss. Abstr.) . 1965a. Oxygen consumption, thermal conductance, and torpor in the California pocket mouse, Perognathus californicus. J. Ce. Comp. Physiol. 65:393-403. . 1965b. The relation between the tor- por cycle and heat exchange in the Cali- fornia pocket mouse, Perognathus californi- cus. J. Cell. Comp Physiol. 65:405-414. . 1966. Diurnal torpor and its relation to food consumption and weight changes in the California pocket mouse, Perognathus cali- fornicus. Ecology 47:245-252. Turner, G. C, Jr. 1950. Peromyscus popula- tions as related to seasons and vegetative types at the hardware ranch. Cache County, Utah. M.S. thesis. Utah St. Univ., Logan. Vest, E. D. 1962. The plant communities and associated fauna of Dug^vay Valley in west- ern Utah. Ph.D. dissertation. Univ. of Utah, Salt Lake City. Walkowa, W. 1970. Operation of compensation mechanisms in exploited populations of white mice. Pages 247-253 in Energy flow through small mammal populations. Warszawa. Woodbury, L. 1955. An ecological and distri- butional study of small mammals of Cedar Valley, Utah. M.S. thesis. Brigham Young Univ., Provo, LTtah. COMPUTERIZED REDUCTION OF METEOROLOGIC MEASUREMENTS FROM IRRKiATED AND NONIRRIGATED PLOTS IN CENTRAL UTAH' Ferron L. Andersen" and Paul R. Roper'' Abstract. — Two Fortran iv computer programs were developed to facilitate reduction of mete- orologic data from iri-igated and nonirrigated plots at Provo, Utah. The first program compiles and tabulates daily, monthly, and jearly summaries of precipitation as rain and/or snow, snowfall, total snow cover, soil moisture, dew, lelative humidity, potential evaporation, cloud cover, and wind. Temperature values are tabulated for measurements taken in a standard weather shelter. 5 cm beneath soil surface under grass cover, at soil surface under grass cover, and on bare ground. The second program enables complete computerized (Calcomp) construction, labeling, and graph- ing of 10 different meteorologic measurements and 3 calculated comparisons of temperature means. Advantages of the first computer program relate generally to that obviously noticeable with any computerized tabulation. Those of the second j)rogram relate more specifically to the greatly reduced cost of computerized graphs compared with those produced manually, as well as to the marked reduction of eirors compared with the number frequently associated with the usual tedious and laborious plotting of voluminous weather data. Meteorologic data collected for the year demonstrated the beneficial effect of irrigation in the creation of microenvironmenfs for living organisms. Introduction A comparison of meteorologic measure- ments fromi irrigated and nonirrigated plots in Provo, Utah, for 1970 through 1972 was published recently by Andersen, Wright, and Fox (1974). Included in their report was a detailed description of the study area, instrumentation employed, method of handling meteorologic clata, and a series of manually rej^roduced graphs depicting the results for those three years. The current report is designed as a companion article to the one referred to above. It extends the meteorologic measurements through 1973 for an ad- jacently located plot watered via sprink- ling as opposed to flood irrigation used in the previous project. Furthermore, it emphasizes those changes that have been incorporated to enable the graphing of 10 different meteorologic measurements and 3 comparisons of calculated means by a Calcomp plotter. The importance of irrigation in creating optimum microenvironments for biolog- ical organisms has been substantiated for such invertebrates as mosquitoes (Rainy and Hess, 1967; Reeves and Hammon, 1962), snails (World Health Organization, 1950), and nematode larvae (Furman, 1944; Honess and Bergstrom, 1966; Fox, Andersen, and Hoopes, 1970; and Wright and Andersen, 1972). The importance was elaborated upon further by Andersen ^This project was supported in part Ijy Public ticallli Seri ^Department of Zoology, Brigham Yoiuig l^niversity. I'n ^Eyring Researcli Institute, Provo, Utali. et al. (1974) and, thus, will not be dis- cussed here. Description of Experimental Plot The experimental plot measured 10 x 30 m and was located at the Brigham Young LTniversity Animal Science Farm, Provo, Utah, adjacent to the plot used in the study by Andersen et al. (1974). The coordinates of the station site, elevation, description of vegetative cover, soil type and drainage, instrumentation, and meth- od of collecting data were the same as re- ported earlier. The plot was divided into an irrigated and a nonirrigated section separated by an elevated dike. Addition- al diking, a])roximately 8 in (20 cm) high, was also placed around the upper border and sides of the plot to prevent any flood irrigation waters from entering the experimental area. Beginning in May the irrigated portion was sprinkled each Monday with a total of 1 in (2.5 cm) of water as measured in the rain gauge. This was applied at an approximate rate of 0.5 in/hr. In June the amount was in- creased to 1.5 in (3.25 cm) and in July to 2 in (5.0 cm). The amounts were re- versed for August, September, and Oc- tober, after which time sprinkling ceased. This regimen for 1973 added a total of 39 in (97.5 cm) of water to the irrigated plot over the six month period. 203 204 GREAT BASIN NATURALIST Vol. .55, No. 2 Descriptions of Programs Two FORTRAN IV programs, one for data tabulation and a second for Calcomp graphing, were developed for this project. Figure 1 outlines the collation and hand- ling of meteorological data and Figures 2 and 3 illustrate flow charts for the data tabulation and Calcomp graphing pro- grams respectively. All meteorologic data for the year were entered on specially de- signed worksheets (Figs. 4 and 6), key- punched on 80-column IBM cards, and handled as diagrammed. All mensural data not already in the metric system were so transposed by appropriate con- ifii WEATHERSTATION version formulae. Also, the printing of all negative or zero values was suppressed whenever such data were not meaningful. The first program compiles and tab- idates daily, monthly, and yearly sum- maries of all meteorologic data collected. Tables 1, 2, and 3 are sample printouts of one month's data (May 1973), and Tables 4 and 5 are the two-page annual sum- mary sheets. The second program developed for the project enables complete computerized PRINT MONTHLY HEADING . >' READ DATA FOR ONE DAY , CALCULATIONS a CONVERSIONS ^/^C 0F\. NO VONTHLY SUMS, AVERAGES 6 EXTREMES PRNT MONTHLY D(\TA(3PAGES) STORE MONTHLY TOTALS FOR YEARLY SUMMARY Pig. 1 Diiigrarn showing how metoorologir data were collated and handled. Fig. 2. Flow chart for the data tabulation program. June 1975 ANDERSEN, ROPER: METEOROLOGIC DATA 205 plotting of 10 different meteorolocric measurements and 3 calculated mean tem- perature comparisons. Each deck of ]iro- gram data for any one year must be ])re- ceded by a control card identifying the year and number of days in that year. Also included on the control card for the Calcomp plotting program is a list of identifying nmubers which permit selec- tion of desired gra])hs for that year. These graphs are then comjileted sequentially as selected. Figures 6 through 18 depict comj5uter- ized reproduction of 10 different metero- logic measurements taken during 1973 and 3 calculated comparisons of means as indicated on each individual legend. Table 6 gives information for the 1973 data regarding the a]:)proximate run times and current costs for the IBM 360/65 in- stallation at Brigham Young University. The complete printout of all daily, monthly, and yearly data is available uj)on request for the cost of reproduction. Both com])uter j)rograms are printed here- in (Appendix) for researchers who might find sections or subroutines applicable to their specific needs in meteorolog\- or biology. Regarfling specific handling of meteoro- ( BEGIN j READ CONTROL CARDS POSITION TO YEAR'S DATA READ YEARLY DATA BRANCH TO DESIRED GRAPH NUMBER INCREMENT TO NEXT GRAPH NUMBER >rl3 SCALE DATA DRAW AXES 8 LABEL MOfvTTHS DRAW AXES 8 LABEL MONTHS LABEL LEGENDS a UNITS LABEL LEGENDS 8 UNITS PLOT DATA I i i i i PLOT DATA Fig. 3. Flow chart for the Calcomp plotting program. 206 GREAT BASIN NATURALIST Vol. 35, No. 2 Table 1. Sample monthly summary of measurements on precipitation, soil moisture, relative umiditv. potential evapoiation, cloud cover, and wind — May 1071 PRECIPITATION (f SOIL MOISTURE 0.3 0.3 PERCENT DEfc TOTALS MEANS EXTREMES HIGH L28.6 23 TE Cf IRRIGATION (MAY THROUGH OCTOBER! Table 2. Sample monthly summary of temperatures measured from a standard weather shelter and at 5 cm beneath soil surface under grass cover — May 1973. •EATHEP SHfl THERMOMETEf FEMPEBATUReS HYGPOTHERMOGRAPH L SURFACE NON-IRRIGATfD MAX. MIN. M 12.0 17. T 11. 5 ll.O 35.0 12.0 26.0 13.3 27. ) 15.1 MEANS EXTREMES EMPERATURES IN DEGREES CELSIUS June 1975 ANDERSEN, ROPER: METEOROLOGIC DATA 207 Table 3. Sample monthly summary of temperatures measured from a standard weather shelter, at soil surface under grass cover, and on bare soil surface — May 1973. SOIL SURF4CE UNDER GRASS COVER 8«RE SOIL SURFS THERMOMETER IRRIGATED NON-IRRIGATED X. HI N. MEAN .3 -3 5.0 7 14 14 12 12 12 10 K 13 15 17 15 15 17 17 'o 10 20 .8 K .1 10 18 U H .7 IC 18 8 6 13 15 15 17 5.8 14.3 25.0 25.0 32.0 .0 .0 11. 13. .0 16. .0 18. .0 15. .0 16. .0 17. .0 .0 I7I 17. !o 21. .0 24. .0 24. .0 26. .0 25. .0 27. .0 27. 28. !o 31. .0 30. .0 24. .0 26. .0 2<>. .0 30. .0 It. .0 21. .0 22. .0 21. .0 25. .0 25. .0 26. 32.0 9.0 20.5 17.0 10.0 13.5 24.0 9.0 16.5 24.0 11. 0 17.5 26.0 12.0 19.0 22.0 8.0 15.0 28.0 10. 0 19.0 30.0 10. 0 20.0 30.0 8.0 19.0 35.0 9.0 22.0 35.0 9.0 22.0 36.0 8.0 22.0 35-0 8.0 21.5 36.0 9.0 22.5 39.0 10.0 24.5 41.0 16.0 28.5 43.0 12.0 27.5 37.0 13.0 25.0 38.0 9.0 23.5 43.0 9.0 26.0 40.0 13.0 26.5 45.0 14.0 29.5 IT.O 7.0 12.0 27.0 7.0 17.0 31.0 8.0 19.5 36.0 10.0 23.0 39.0 10.0 24.5 39.0 12.0 25.5 38.0 14.0 26.0 .0 0 0 14.0 18.3 .0 5 0 24.0 .0 6 0 25.5 .0 5 23.0 .0 6 0 24.0 .0 6 0 26.5 .0 2 0 20.0 .0 0 24.5 .0 3 0 27.5 .0 8 0 23.5 .0 8 0 29.0 .0 0 30.5 .0 8 0 31.0 .0 7 0 31.0 .0 9 0 32.5 .0 10 0 34.5 .0 15 0 37.5 .0 12 0 36.5 .0 13 0 32.5 .0 9 0 20.5 .0 9 0 27.0 .0 14 9 32.0 .0 9 0 30.5 .0 7 0 12.0 .0 6 0 18.5 .0 6 0 21.5 .0 9 0 21.5 .0 9 0 23.5 .0 11 0 27.0 .0 14 0 30.0 25.5 27.5 24.0 ALL TEMPERATURES IN DEGREES CELSIUS Table 4. Yearly summary of measurements on precipitation, soil moisture, relative humidity, potential evaporation, cloud cover, and wind — 1973. MONTH TOTAL PREC PRECI PAIN pREr PITATTON SNCW PREC (MM) SNOW FALL SNOW COVER SOIL MOISTURE (PERCENT! IRRG NON-IRRG RELATI HUMIDI JANUARY TOTAL MEAN 29.7 1.0 0.0 0.0 2';. 7 1.0 297.2 3302.0 106.5 NO NO 98 FEBRUARY TOTAL MEAN 33.8 1.2 31.7 l.l 2.0 Q.l 2T.3 0.7 363.2 13.0 NO NO 98 MARCH TOTAL MEAN 86.6 2.8 58.7 1.9 27.9 0.9 279.4 2S7.0 9.3 UD NO 98 APRIL TOTAL KEAN 1.5 33. U 1.1 I 1.4 0.4 114.3 3.8 114.3 J. 8 NO NO 98 IS OF POT AUGUST SEPTEMBFP CCTOBER NOVEMBER CECEMPER OTAL EAN 34.0 1. I 34.0 1. L 0.0 0.0 OTAL 18.3 0.6 18.3 0.6 0.0 0.0 OTAL EAN 19.8 0.6 19.8 0.6 0.0 0.0 OTAL 51.8 51.8 T.O tOTAL "FAN lOTAL 13.2 0.4 17.0 0.0 0.0 0.0 0.0 0.0 0.0 9. 8. 0.0 0.0 0.0 0.0 <,'. 0.0 0.0 0.0 0.0 11. 0.0 0.0 oio 12. 0.0 0.0 0.0 0.0 25.7 10. CLOUD COVER WINO (0-101 (KM) 1183 6 38 1144 5 40 2012 fc 64 1901 4 63 1 15 a FEpruflffT npwcn . prr^iL rwr juwe , jult wjcust sEnEweEw, octopew , wovEr^pew pecEWBEr SNOW COVER PROVO. LiTBH. 1973 250- 225-] cco-j © ^.. infi 6 ^ !H lA M IN3 JHNUflflY FEBRURRY HflRCH ^ flFRlL . ^»Y jUnF . JULY , PJJGOST ^ SFPTEMBCR DCTQBEfl . NOVEMBER . DFCgUBFR I RELATIVE HUMIDITY IN WEATHER SHELTER JHNUfWY FEBRUflRY , MARCH fiPRlL JUNE . JULY , BUGUST , SEPTEMBER. JHOBER . NOVEMBER , DECEMBER 210 GREAT BASIN NATURALIST Vol. 35, No. 2 jUNf , Mir , AUGUST . SLPieMPeR. OCTOBgB . NPYEM6FH . DCCEMeew ffwusT , scrTEfipEn, ocToeE^ , wovEnoEff , oEcempeh TOTAL WIND 1 METER ffiOVE GROUND PSDVO. U7PH, 1913 E 160. i JBNUWY .rCBBUOSy . MflBCH . fyWIL . HftY . jOnE . JULY . RUGUST , SCPIfrBER. OCTDBER . NOVFWER . oecCMBfK . June 1975 ANDERSEN, ROPER: METEORO LOGIC DATA 211 TEMPERATURE IN WEATHER SHELTER PRDVD. UTftH. 1973 15 1 15 1 15 1 15 1 15 1 15 1 IS 1 15 ) 15 1 15 1 IS ) IS JHNUPRV .FfBBI^RY . MftRCH , WRU . mf . JUNE . JUIV . flUCDST . 5fP1FM6fR. OCTQefB . NO»fWBfH . DfCCM8fR COMPARISON OF MAXIMUM AND MINIMUM TEMPERflTLIR&S'l , 5 CM OEfP In soil L'NDfR 10 O CRASS COVER , fyj ^ ', ii ^_ ON IRRIGfiTED ftNO NON-lflRICfllfO PLOTS PROVO% UTflHi 1973 MAX NON-IRR MIN NON-IRR HftX IRRIGWED HIN IRRIGATED JflNUtWY EEBflUflRY ^ MfWCri ^ ftPRIL . WiY flOGbST . SEPTfMeER. OCTOBER . NDVEMeER . DECEneER COMPRRISON OF DFllLY MEAN TEMPERATURES S Cfl DEEP IN SOIL UNPEB id CM DPflSS UIVFR ON IRHIOPTFD AND NDN-IBR lOflTED TLOTS rrOVO. UTRH. 1973 212 GREAT BASIN NATURALIST Vol. 35, No. 2 COMPARISON OF MAXIMUM AND MINIMUM TEMPfRRTUP£:,S!: r _..^: WW NON-IRR 50- ON IRBJCflTfO AND N0N-IRP1&H7CD PLOTS . . PflOVO, UTAH, 1973 . ^ :,■ jj^ ii^lji': MIN IRRIGOTED MO- " 30- M Wi %ij 1 /: h L,¥fts:i ^^n^ t\'' 10- ri>^Y^ V ■^\W%v temll A 0 ID Am^i a' JMi./W"^ ■ f •■ ■. ^1 i L' v )?(/f\/|;;;^!5t[(c^ ^^^^^^^^^^^^^^(V^jV^I •» '■■"■■■"'■■ I's i is ') ')5 I 15 1 15 1 is 1 is 'i is i I'S 1 15 i 15 i I'S QPMUflBT . FtBBUflRT RUCUST . SFFTFfiBFR. 0CT06CB . NOVCMSfH . DECfUBCH COMPfiRISDN OF OfllLY MCflN TEMPERATURES m sou suRFHCf UNDfn lo i>i Offlss ON IRfilGRTfO fiND NON-lRRIGfllfO PtmS PROVO, UTBH, 1973 JfiNUPRT f-fBRUW SFPlffWfR OClOefR NDVEwerR . OCCfllBCR COMPARISON OF MAXIMUM AND MINIMUM TEMPERATURES OT SOIL SURFRCC ON BflRf GROUND ON IRRlGflTCD AND NON- IRRIGATED PLOTS j PROVO. UTAH. 1973 « ■"/ HflX NON-]Rfl WIN NON-IRfl NftX IRfllGRTEO (•IN IRRIGfllfO I'S i Ts flocusT . sfPTfnefR. ocToefR . NOVC^eCR . occfnBffl . .hiiie 1975 ANDERSEN, ROPER: METEOROLOGIC DATA 213 logic iiiforniatioii from the i)asture plots as well as certain calculations and con- versions b}' the computer, the following clarifications may be warranted. All measurements of precipitation as rain are measured and entered in fractions of inches and then converted before tabu- lation into millimeter units. New snow and total snow cover are similarly han- dled. Precipitation as snow is calculated as one-tenth that of total snowfall for any one day. Soil moisture samples were de- termined as described previously by An- dersen et al. (1974), with the exception that two samples were taken each week from the irrigated plot — one just prior to sprinkling of the plot and the second 24 hr later. Only one weekly sample was taken from the nonirrigated plot. The weight in grams of each sample was en- tered as an original wet weight and then as a dry weight determined after 24 hr storage at 105 C. Programmed formulae calculated the percent soil moisture by dividing the dif- ference in the wet and dr}- weight for each sample by the dry weight value. This per- cent was then printed on the first print- out page for each of the six months that irrigation was used. The presence or ab- sence of dew on each plot was noted for each morning during those six months and recorded as a " + " when present. Irrigation by sprinkling was performed each Monday during the six-month period and also indicated with a " + " on the ap- propriate dates. Relative humidity (RH) maxima and minima, as well as the total number of hours each day at which 98-100 percent RH occurred, were entered and printed out directly. The e\aj)orating pan on the instnnnent that measured potential evap- oration was filled each morning, and the daily water loss noted for the following 24 hr was entered in mm and also printed out directly. Since evaporation could not be read in freezing weather, the measure- ment was taken only during May through October, those same months during which the (^ne plot was irrigated. A difficulty ensued whenever rain occurred because the e\aporating pan held only 20 nun of water and refilled partially or completely on any day during which rain fell. Thus, any potential evaporation which may ha^e occurred on such days was invariably ne- gated to some degree by the rain that col- lected in the evaporating pan. Neverthe- less, the recording evaporimeter used in the study gave a much more accurate re- flection of the potential evaporation with its shallow pan (20 mm) than do the large evaporation tanks used by the U.S. Weather Bureau. In those tanks the water level is frequently 4-6 in (10-15 cm) below the ujiper rim of the pan and hence protected markedly from the evapo- rating effect of wind currents. Any evaluation of daily cloud cover was determined visually each morning and recorded in tenths. The evaluation indi- cated the approximate portion of the sky that was covered sufficiently with clouds to cast a shadow at the time the instru- ments were read. It was thus the most subjective of all measurements taken but, nevertheless, provided some estimation of cloud cover in this region. Wind totals were entered in mile units read from the anemometer dial each day, calculated as the difference from the value of the pre- 214 GREAT BASIN NATURALIST Vol. 35, No. 2 ceding day, and converted and printed out as kilometers. The final daily wind total for any preceding year is listed as a starting value and included on the control card for the main program. Maximum and minimimi temperatures monitored from a standard weather shel- ter, from 5 cm beneath soil surface under grass cover, from soil surface under grass cover, or from bare ground were recorded, converted to Celsius if not already in those units, and ])rinted onto the second and third sheets for each month's data. Means and extremes for all values were stored for eventual calculation of totals and grand means for the annual smnmary pages. Daily maximum and minimimi tem- peratures were measured in the weather shelter by mercury- and alcohol-filled thermometers as well as by a standard hygrothermograph. Differences noted in the recorded temperatures relate mainly to the longer time lag required by the bi- metallic sensor within the thermograph unit. Discussion The main objective of this paper was to present the computer programs devel- oped for our research on the effect of irrigation on pasture microenvironments. These programs have proved extremely satisfactory to us, and hopefully some sections or subroutines will be of value to others engaged in related research pro- jects. Mitchell and Andersen (1969) re- ported on a computer program, deAoloped at the Uni\ersity of Illinois at Urbana, for handling meteorologic data collected from grass plots. Certain similarities exist between that program and the one re- ported here, since the choice of meteoro- logic instruments and the overall research projects at the two institutions were closely correlated. The program at Illi- nois, however, was designed to handle some additional measurements not taken in the current study, such as solar radi- ation. The Illinois study also gave em- phasis to conversion data for a series of soil-moisture and soil-temperature mea- surements obtained through moisture-cell leads (wafers) and built-in thermistor units. The programs developed at BYU use data on soil moisture only from simj)- lified gravimetric measurements but in- clude the techniques designed for Cal- comp graphing as well. The advantage of this plotting program is not only in the funds saved through not having to man- ually plot, trace, and label all such graphs but more particularly in the marked re- duction of errors that invariably accom- l)an3" the tedious and laborious tasks en- countered in manually plotting daily weather data. The impact of irrigation on the moisture and temperature profiles in central Utah during 1973 was essentially the same as that reported for 1970-72 by Andersen et al. (1974). Grand mean temperatures for 5 cm beneath soil surface under grass cover, at soil surface under grass cover, or on bare soil surface were consistently lower on the irrigated plot than on the nonirrigated area. These temperature dif- ferences were most apparent during the warm summer months when irrigation is connnonly employed throughout the re- gion. In July, for example, the month during v.hich most yearly maxima were recorded, the average monthly maximum temperatures measured 5 cm beneath soil surface under grass cover on irrigated and nonirrigated plots differed by 6.4 C, by 15.2 C for those measured at soil sur- face under grass cover, and by 17.9 C on bare ground. During the six months when no irrigation occurred, the differences were not so apparent; hence the grand means for the year do not give an ac- curate reflection of these temperature ranges for the irrigation season. The contrast between soil moisture measurements in the two plots was read- ily apparent for the six months during | which soil samples were gravimetrically analyzed for moisture content. The grand mean for the percent soil moisture for that period was 22.7 percent for the samples removed from the irrigated section and 9.4 percent for those from the nonirrigated area. Other meteorologic measurements which were taken, such as the precipitation pat- tern for rain or snow, relative humidity, ]iotential evaporation, and wind, were monitored for both sections combined, ' since the instruments used could not de- tect flifferences from microenvironments. The general pattern of these values was similar to that recorded for pre\'ious years. The collective data for 1973 emphasize the marked beneficial impact of irrigation in creating favorable microenvironments for Ha inij; organisms. June 1975 ANDERSEN, ROPER: METEOROLOGIC DATA 215 0002 0005 0006 0007 0008 0OO9 OOLO OOll 0012 0013 0014 0015 0016 0017 0018 0019 0020 0021 0022 0023 0024 0025 0026 0027 0028 0029 0030 0031 0032 0033 0034 0035 0036 0037 Appendix * weather data analysis, part i ♦ * DEVELOPED BY FERRON ANDERSON, BYU * PROGRAMMED BY PAUL ROSS ROPER, ER I 4< ^L'^^if^m************************ ************************* *^L^:t ****** INTEGER NDAYS, DAY, CLCV, WIND, OYR, OEWl, IRRG, DEW2, HHUM, LHUM, T OLOW, S2(26,5), M0N(12), HOLD, MONTH REAL •MARC , •OCTO' , 'L ', •0 ME* , SNOW, •APR I •NOVF 3*« 8*« SNCV, MOSI, MOSN, PEVP, Sl(26,8), S3(13,24), •MAY ' •DECE' •ST ' •R " 'JUNE' 'GRAN' •EMBE' 3*' • CCMMON THRS(35) 'JANU', 'FEBR •AUGU*. 'SEPT •UARY', 'H •MBER'f 'MBER TPRE, RAIN, PRSN IRDW, NIWW, NIDW, T(34,16) PAGE2(34,12). PAGE3(34,15) NOAYS, 0AY(3l», DYR(3l), TPRE(35), RAIN(35), PRS k SN0W(35), SNCV(35>, M0SI(35), 0EW1(32), IRRG(32) ) M0SN(351, DEW2I32), HHUM(35), LHUM(35) : PEVP(35), CLCV(35), WIN0(35) DATA S1,S2,S3 /208*0.0, 130*0, 312*0.0/ TAPE ID DATA ITAPE /5/ CENT(X) = (X-32.0)*5. 0/9.0 REWIND ITAPE LOOK FOR CORRECT YEAR 1 REAO<5,200,END=999» NYEAR 2 READ( ITAPE, 200, END=999I lYEAR, IFdYEAR.EQ.NYEARI GO TO 4 DO 3 I=l,IOAYS READ! ITAPE,200,END=999) 3 CONTINUE GO TO 2 HRS, (13,31/ •JULY' , •ARY ', 'BER ', 'AN •/ IRWW, IDAYS, MON, OLDW PROCESS DATA PAGE BY PAGE 4 DO 900 11=1,12 CALL HEAD ( I YE AR , II , 1 , MONTH) NOAYS = MON( II I WRITE PAGE 1 HEADING WRITE(6,120I DO 10 1=1, NOAYS READ( ITAPE, 20H DAYd), DYR ( I ) , RAIN(I), SNOW(I), SNCVM), IRWW, A IRDW, OEWKI), IRRG(I), NIWW, NIDW, 0EW2<1). B HHUMdJ, LHUM(I), THRS(I), PEVP(I), CLCV(I), C WINO(I), (T( I ,J),J=1, 16) PRSN( I ) = SNOWd ) ♦ 0. I TPRE( I) = RAIN( I) ♦ PRSN( I ) IF(OLOW.GT.WINDd)) OLDW = OLDW - 1000 HOLD = WINDd ) WIND(U = (WIND(I) - OLDW) * 1.6093 ♦ 0.5 OLDW = HOLD MOSId) = 0.0 MOSN( I) = 0.0 IFdRWW.NE.0.0) MOSKI) = dPWW - IRDW) * 100 / IRDW IF(NIWW.NE.O.O) MOSN(I) = (NIWW - NIDW) ♦ 100 / NIOW CALL PRINTL (1,11) = CENT(T(I,l)) = CENT(T(I,2) ) = CFNT(Td,3) ) = CENTITd ,4) ) = T( 1,5) = T( 1,6) = T( I, ID PAGE2d ,1) PAGE2( 1,2) PAGE2( 1,4) PAGE2d,5) PAGE2d,7) PAGE2( 1,8) PAGE2( I. 10) 216 GREAT BASIN NATURALIST Vol. 35, No. 2 0038 0039 0040 0041 0042 0043 0044 0045 0046 0047 0048 0049 0050 005L 0052 0053 0054 0055 0056 0057 0058 0059 0060 0061 0062 0063 0064 0065 0066 0067 0069 0070 0071 )072 0C73 0C74 0075 0C76 0077 0C78 0079 0080 0081 0082 0083 PAGE2(I.in = T(I 12) PAGE3(I,I> = PAGE2( I, I) PAGE3(I,2) = PAGE2(I,2) PAGE3(It4) = T(I 7) PAGE3( 1,5) = T(I 8) PAGE3( 1,7) = T(I 13) PAGE3(I,8) = T( I 14) PAGE3( 1,10) = T(I 9) PAGE3(I,ll) = T(I 10) PAGE3( 1,13) = T( I 15) PAGE3( 1,14) = T( I 16) c 10 CONTINUE c CALL SUMR (TPRE) CALL SUMR (RAIN) CALL SUMR (PRSN) CALL SUMR (SNOW) CALL SUMR (SNCV) CALL SUMR (PEVPI CALL SUMZ (MOSI) CALL SUMZ (MOSN) CALL SUM! (HHUM) CALL SUM! (LHUM) CALL SUMI (THRS) CALL SUM! (CLCV) CALL SUMI (WIND) CALL SUMC ( IRRG) CALL SUMC (OEWL) c c c CALL SUMC (DEW2) PAGE 1 OF WEATHER DATA IFdl .LT.5.GR.II.GT.10) GOTO 20 WRITE(6,l2l) TPRE(32), A M0SI(32), B PEVP(32), WPITE(6,122) (TPRE(I), A MOSI(I), PRSN(32), SN0W(32), SNCV(32), IRRG(32), M0SN(32), 0EW2(32), RAIN(32) 0EW1(32) WIND(32) RAIN(I), PRSN(I), SNOW(I), SNCVd), MOSNd), HHUM(I), LHUM(I), THRS(I), F) PEVPJn, CLCV(I), V2 FORMATCO ' , 3 A4 , ' T HTAL ' f 5F 8. I , 5( 6X , ' -- • ) , F 8. I ,oX , • -- ' , I 8 , / , A 15X,'MEAN ' ,7F8. I, 3I8,F8.1,2I8) 0160 163 FORMAT('0 ' , 3A4 , • TOTA L ' , 5F8 . I , 5( 6X , ' -- • ) , 6X, ■ ND ' , 6X, ' -- ' , I 8 , / , A 15X,'MEAN • , 5F6. 1,2(6X,'ND' ) ,318,6X,«ND' ,218) 0161 164 FORMATI'- YEARLY TOTAL ' , 5F 8. I ,5 ( 6X , ' — •), F 8. 1 , ' *' , 5X , ' -- ' , A l8,/,'0 GRAND MEAN • , 5F 8 . 1 , I X ,2 ( F 7 . 1 , ' * ' ) , I 7, 21 8 , 3 F8. l,'*', 17, 18,/,'- ND = NOT DETERMINED',/,' * CALCULATE CD 1 MAY THROUGH 31 OCTOBER ONLY') 0162 165 FORMAT('l ',14,' SUMMARY - TE MPER ATURES • , 95X , • P ART 2',///,5lX, A 14, • SUMMARY OF METEOPOLOGIC 0 AT A • , / , 55X , ' BYU PARASITOLOGY STATIC BN* ,/ ,61X,' PPOVO, UTAH',//) 0163 166 FORMAT! 34X, 'WEATHER SHELTER TEMPERATURES', 32X,'5 CM BENE AATH SOIL SURFACE',//,28X,'THERMnMETFR',16X, 'HYGR CTHERMOGR AP H' , 16X, B' IRRIGATED' ,18X, 'NON-IRRIGATED' ,/, '0 MONTH ' , 8 X , 4 ( 8X, ' MAX . MIN. C MEAN ' ) ,/) J164 167 FORMAT (12(' ' , 3A4 , 4 ( 5X , 3F 8 . I ) / ) , • 0 • , 3 A4 , 4( 5X , 3FS . 1 ) , // ) 0165 168 FORMAT( '-' ,32X,' SOI L SURFACE UNDFR GRASS C OVER ', 34X ,' BARE SOIL SUR AF ACE ',//, I IX, 2( 18X ,' IRRIGATED' , 18X, ' NON- I RR IGAT ED ' ),/,'0 MONTH', B 7X,4(9X, 'MAX. MIN. MEAN'),/) 0166 169 FORMAK'- ALL TEMPERATURES IN DEGREES CEL S lUS • , / , ' I ' ) U67 200 F0PMAT(20I4) 01^8 201 F0RMAT(A2,A3,F3.2,4F3. l,2Al,2F3.1,Al,2I3,I2,F3.l,I2,I3, A /,5X,16F3.0) 0 169 END JOOl SUBROUTINE HEAD ( 1 YE AR , M, I PG, MONTH ) C C WRITE A HEADING AND PAGE NUMBER FOR EACH MONTH C 0002 INTEGER NOAYS, LAY, DYR, OEWl, IRRG, DEW2, HHUM, LHUM, THRS, A CLCV, WIND, LINE(IO), LINl(2), LIN2(2). M0NTH(13,3) C003 RE»L TPRE, RAIN, PRSN, SNOW, SNCV, MC S I , MOSN, PEVP 0004 roMMnN NDAYS, DAY(31), DYR(31), TPRE(35), RAIN(35), P«SN(35), A SNOH(35), SNCV(35), MnSI(35), 0EW1(32), IRRG(32), 8 MOSN(35), DEW2(32), HHUM(35), LHUM(35), THRS(35), C PEVP(35), CLCV(35). WIND(35) June 1975 ANDERSEN, ROPER: METEOROLOGIC DATA 219 WRITE(6,100) (MONTH(M,I ) ,1=1,3) , lYEAR, \ lYEAR RETURN IPG, {MONTH(M, I ) , 1= 1,3), ENTRY PRINTL (1,11) PRINTL FORMATS DAILY DATA FOR PRINTING INCLUDES SCALING AND SUPPRESSING ZERO RESULTS CALL ALPHA (TPRE(I), LINE(1», 25.4) CALL ALPHA (RAIN(I), L I NE ( 3 > , 25.'*) CALL ALPHA (PRSN(!), LINE(5), 25.4) CALL ALPHA (SNOW(I), L ! NF ( 7 ) , 25.4) CALL ALPHA = .TRUE. GOTO 1 END SUBKJUTINE SUMI (lY) PROCESS INTEGER DATA IN ARRAY lY AND RETURN SUM IN IY{32», ME4N IN IY(33), MINIMUM IN IY(34I, AND MAXIMUM IN IY(35). INTEGER IY(35), PLUS/'*'/ ANDERSEN, ROPER: METEORO LOGIC DATA 221 CCMMON NDAYS IY(32» = 0 IY( 3^) = 0 IY(35) = 1000 DO 13 I=l,NOAYS I Y(32) = I Y(32» ♦ IY(I ) IF(IY(34» .LT. IY( I ) ) IY{34) IF( I Y(35).GT. IY( ! ) ) IY{35) 13 COM iNue I Y( 33) = IY(32) • / NDAYS ♦ RFTURN ENTRY SUMC (lY) lYtl) IY(I ) PROCESS CHARACTER DATA IN ARRAY lY AND RETURN THE NUMBER OF IN IY(32). IY( 32) = 0 DO I't 1 = 1, NDAYS IF(IY( D.EC.PLUS) IY(32) = IY(32) ♦■ 1 l^ CCNTINUE RFTURN FND WEATHER DATA ANALYSIS, PART II PROGRAMMED BY PAUL RCSS ROPER EYRING RESEARCH INSTITUTE, 197^ Hc*t*^f****j^i^****^t^^*t^t^f7t:^^it:tti¥** ********* *********************** DECK FORMAT I. CONTROL CARD 2. DATA HEADER COMMON /PLOTO /PLOTl /PL0T2 /PL0T3 /PL0T4 /PL0T5 /PL0T6 /PLOT? /PL0T8 /PLQT9 /PLOTIO/ /PLOTll/ /PL0T12/ /PL0T13/ COMMON /LABELS/ CCNTROL CAPO DATA HEADER DATA (ON TAPE (ON TAPE IF DESIRED) IF DESIRED) COLUMNS DESCRIPTION 1-A YEAR 5-8 DESIRED GRAPH BY NUMBER 9-12 ... 13 GRAPHS POSSIBLE 1-4 YEAR OF DATA 5-e NUMBFP OF DAYS IN YEAR 9-12 LAST WIND READING OF PREVIOUS YEAR SEE PUBLICATION FOR CARD COLUMNS AND DATA DESCRIPTION. XPLT( RAIN( SNOW( YHHD( HUMO( EVAP( WIND( YMAXC YH5C( YHAK YHSS( YHA2( YHBG( YHA3( AXl(3 BX(4) RC1(7 RC3(l RC5(1 RC7(l 366) 366) 366) 366) 366) 366) 366) 366) 366) 366) 366) 366) 366) 366) 7), ), !)• l)t 2), ITAPE XIRR(366), YIRR(366), YNIR(366) , YMIN(366) , YL5C(366) , YLA 1(366) , YLSS(366) , YLA2(366) , YLBG(366) , YLA3(366) AYl(37), AX2(73) ZH5C(366) , ZL5C(366) ZHSS(366) , ZLSS(366) ZHBG(366) , ZLBG(366) AY2(73) BY(4), XL(4), YL(4), LCl(7), RC2( 13), LC2( 13) LC3( ID , RC4(13) , LC4( 13), LC5(18), RC6(8), LC6( 16) Lr7(24) CREATE A DEGREE SYMBOL 222 GREAT BASIN NATURALIST Vol. 35, No. 2 0003 0004 0005 0006 0007 0008 0009 0010 0011 0013 0014 0015 0016 0017 0018 0019 0020 0021 0022 0023 0024 0025 0026 0027 0028 0029 0030 0031 0032 0033 0034 0035 0036 0037 1 J38 n39 0040 J HI D JA2 •)043 0C44 0045 0046 0047 0048 DIMENSION DEG(3)f 0(3), NGRAPH(14) DATA DEG /Z05061727, DATA 0 /ZC1061737, CALL SYMBL5 (J, 9, DEG) CALL SYMBL5 (K,9,0) REWIND ITAPE READ CONTROL CARD 100 PFA0(5,200,EN0=999) NYEAR, N 200 FORMAT( 1514) NUM = 0 READ ONE YEAR'S DATA CALL READC ( NYE AP , NCAYS ) CONTROL LCOP 300 NUM = NUM ♦ 1 M = NGRAPH(NUM) GO TO (1,2,3,4,5,6,7,8,9,10,11,12,13) GO TO 100 999 CALL PLOT (12.0,0.0,-3) STOP Z36352414, ZC506I727/, J /Z9l/ Z46413010, Z01061737/, K /ZD6/ C****:****!^* ******* PLOT I ♦********«*****♦****»»*****»*******♦*♦** ******* PRECIPITATION AND SCIL MOISTURE ( 3. 0,RC I, 4, 0.5, 7, I, LCI, 4, 0.5, 7, I) (RAIN,NCAYS,3.00, 3.0, 0.0, 2) CALL SYMBL4 ( -0. 24 , I .2 , 0. 07, 'M I LL I MET ERS • , 90. J , II ) CALL SYMBL4 ( 0. 2 , 2. 8 ,0. 1 , • PR EC I PI T ATI ON • , 0 . 0, 13 ) CALL SYMBL4 ( I .39 , 2 . 8, 0. 07 , • AND • , 0. 0, 3 ) CALL SYMBL4 ( I .64 , 2 . 8, 0. 1 , • SCIL MO I ST URE • ,0.0 , 13 ) (NYEAR ,0.2,2.65) ( XPLT,RAIN,NOAYS) (8.21, 1.3,0.0 7,' INCHES ',90. 0,6) CALL AXES CALL SCALE CALL DATE CALL BAR CALL SYMBL4 CALL MOIST GO TO 303 C C C********»********PLOT 2*****************"****************************** SNOW COVER (3.0,RC2,4,0.2 5,13, 1,LC2,4,0.25,13,1) (SNOW,N0AYS,3.00, 12.0, 0.0, 3) CALL SYMBL4 ( -0 . 2 7 , I .2 , 0. 0 7 , • MI LL I MET E RS' , 90. 0, 1 1 ) CALL SYMBL4 ( 0.2 , 2 . 8 ,0 . 1 , ' SNOW COVER ' ,0.0 , 10) (NYFAR ,0.2,2.65) (XPLT,SNCW,NDAYS) CALL SYMBL4 ( 8. 2 3, 1 .3, 0. 07 , • INCHE S' ,90.0 , 6) 2 CALL AXES CALL SCALE CALL DATE CALL BAR GO TO 300 C C C**********> C c c c*****PI^OT 3*********************************************** RELATIVE HUMIDITY IN WEATHER SHELTER 3 C ML CALL C'^LL AXES ( 3.0, Rr 3,^,0.25, II, I, LC 3,4,0.25. I 1 , I ) SCALE ( YHHL ,NCAYS,2.49, ICO.O, 0.0, 't ) SCALE ( YLHr,NCAYS,2.49, loo.o, 0.0, 5) CALL SY^BL4 { -0 . -^ 5 , I . 1 , 0. 0 7 , ' PE PC (" N T • , 90. 0 , 7) CALL SY'^PLA ( 1.2 , ^.8 ,T. I, 'PhLAT! VE HUMIDITY IN WEATHER SHELTER \ 0.0,3b) CALL DATE ( fJYF /iP , 0 .2 , 2 . 65 ( C^LL LEGtrjD ( 1,7. J, ^.6) CALL LINES ( XPL T , YHHD ,ND AYS , 0 . QI , 2 ) CALL OASH ( XPLT , YLHD .KCAYS, J. ri) CALL SYMRL4 ( 0 . 2 5 , I . 1 , '' . ) 7 , ' PF P CF^J T • , 9 ■) . J , 7 ) GG TO 300 ANDERSEN, ROPER: METEOROLOGIC DATA 223 C 04TI Y nURATION OF RbLATIVF HUMIDITY C r ^ CALL AXES < 3.0,PC^,4,0.21, 13, l.LC^, -^,0.21, 13, 1) CALL SCALE < HUMO , NDAY S , 2 . ^*9 , 2^.3, 0.3, 6) CALL SYM8L4 ( -0 . 2 2 , I . 5 , 0. 0 7 , • HCUR S • , 9 0. 3 , 5 J CALL SYM8L4 ( 0 . 2 , 2 . 8 , 0. I , ' D A JL Y OHRATlnN OF RELATIVE HUMIDITY AT 9 A8-100(', ).1,^0) CALL DATE ( NYF AR , C.2 , 2. 6 5 ) CALL BAR ( XPLT,HUMO,NOAYS) CALL SYMBL4 ( 8 . 23 , 1 . 5, 0 . 0 7 , ' HOURS ' , 9 ) .0 , 5 ) GO TO 300 C C C«*«*«* ««:(<*♦* *«***PLCT 5«*****»** «•****«*♦*«♦ **♦*♦♦«♦*:**♦«****«♦*»***«♦* C POTENTIAL EVAPCRATICN 5 CALL AXES ( 3.0,RC5,4,0.28, 11, l,LC5, 5,0.3b,l8,2) CALL SCALE ( E VAP , MCAY S , 3 . 0 ) , 21. T, ■■^.0, 7) CALL SYMBL4 ( - J. 2 3 , I -2 , 0.0 7 , • M I LL I MFT FRS • ,90. 0 , 1 I ) CALL SYMBLA ( 0 . 2 , 2. 8 ,0. I, ' POTEN TI AL E V APORAT I ON' , 0. 0, 2 I ) CALL DATE (NYEAR,0.2,2.6t)) C\LL SYMBL4 ( 2 .0 , 0 . 5 ,0. C7 , • ST AR TED • , D . T ,7 ) J = NDAYS - 244 CALL LINES ( XPLT ( J ) ,E VAP ( J ) , 184, : . 01 , 2 J CALL SYMBL4 ( 7.0 , 0. 5 ,0 . C7 , • STOP PED • ,0.0 , 7 ) CALL SYMBL4 ( 8 . 30 , I . 3, 0.07 , ' I NCHF S • ,90. 0 , 6 ) GO TO 3J0 C r f «***«**♦***♦ j(* + **PLOT 6***************** *"■**♦**«"*****♦*♦*♦**■«• i"* ***♦♦*♦ C TOTAL WIND I METFR ABOVE GROUND C C 6 CALL AXES ( 3.0,RC6,4,0.42,e, I ,Lr6,5,0.42, 16,2) CALL SCALE ( W I ND ,NDAY S ,2. 94 , 1 7 5. 0 , 0.0, 8) CALL SYMBL4 (-0. 29, 1 .2 , 0. 07, 'KI LOMETE RS • , 90 .0 , 10 ) CALL SYMBL4 ( 0. 2 , 2. 8 ,0. 1 , • TOTAL WIND 1 METER ABOVE GROUND ', 0. T , 3 1 ) CALL DATE ( NYE AR , 0 .2 , 2. 65 ) CALL LINES ( XPLT , W I ND , ND A YS ,0.0 1 , 2 ) CALL SYMBL4 ( 8 . 3, 1 . 4 , 0- 07 , • M I L E S ' , 90. 0 , 5) GO TO 3)0 C C C ♦**«*♦****<.**♦**«? LOT 7*** ********************************* *«**♦*♦»*** C TEMPERAT'JRE IN WEATHER SHELTER C C 7 CALL AXES ( 4. 0 , R C 7 ( I ) ,4, 0. 5, 9 , I , Lf 7 ( I) , 5 , 0. 5 , 1 3 , 2 ) CALL SCALE ( YMAX , NC AYS ,4. 00 , 122 . 0 , -22 . 0, 9) CALL SCALE ( YM IN , NC AY S ,4. 00 , 122 . 0 , -22 . 0 , I 0 ) CALL SYMBL4 ( -0. 25 , 1 .2 , ). 07 , • TE r'PE R ATURE ( JC ) • , 90 .0, 16 ) CALL SYMBL4 ( 0. 2 , 3. 7, 0. 1 , • TEMPE R ATURE IN WEATHER SHEL TER • , 3 . 0, 301 CALL DATE ( NYE AR , 0 .2 , 3. 55) CALL LEGEND (1,7.0,3.7) CALL ZERO (1.5) • CALL SYMBL4 { 8 . 32 , I .2 , 0. 07 , • TE MPE RA TURE ( JF ) • , 90 . 0 , 16 ) CALL LINES ( XPL T , YM AX , NOA YS , 0. 01 , 2 ) CALL DASH (XPLT, YMIN,NDAYS, 0.02) GO TO 3J0 C C (-*♦♦*♦♦*♦*♦♦♦*♦♦♦♦ PLOT 8 ************************************ ****♦***♦*♦ C COMPARISON OF MAX AND MIN TEMP 5 CM UNDER 10 CM GRASS C C 8 CALL AXES ( 3.0, RC7(3) ,4, 0.6, 6,1, LC7(5) ,5,0.6,12,2) CALL SCALE ( YH5C , ND AY S , 3. 00, 40.0,-10.0,11) CALL SCALE ( YL5r , NCAY S , 3 . 00, 40.0,-10.0,12) CALL SCALE ( ZH5C , NC AY S . 3. 00, 4 1.0,-10.0,13) CALL SCALE (ZL5C,NDAYS,3.00, 40.0,-10.3,14) CALL SYMBL4 ( -0. 25 , 1 . 0 , 0. 07, • T E MPER ATURE ( JC) ' , 90. 0, 1 6) CALL SYMBL4 ( 0 . 2 , 2 . 8 ,0 . 1 , ' COMPAR I SON OF MAXIMUM AND MINIMUM TEMPER AATURES', 0.0,46) 224 GREAT BASIN NATURALIST Vol. 35, No. 2 0096 0C97 0C98 0099 0 100 Old 0102 0103 0104 0 10 5 0106 0107 0108 0109 oin Olll 0112 0113 0114 0115 0116 0117 0118 0119 CALL SYMRL4 ( 0 .2 , 2. 7 ,0. 07 , • 5 CM DEEP IN SOIL UNDER 10 CM GRASS GOV AER't 0.0f41) CALL SYMBL4 (0 .2 , 2 . 6 , 0. 07 , ' ON IRRIGATED AND NON- IRR IGATE 0 PLOTS', A 0-0,36) CALL DATE ( NYE AR , 0 .2 , 2. 45 » CALL LEGEND (2,6.75,2.8) CALL ZERO (0.6) CALL SYMBL4 ( 8. 32 , 1 . 0, 0. 07 , ' TE MPER ATURE ( JF ) • , 90 . 0, 16 ) CALL LINES ( XPL T , YH5C , NDA YS , 0. 01 , 3 » CALL LINE (XPLT, YL5C,N0AYS,n CALL DASH (XPLT, ZH5C ,NDAYS, 0.04) CALL DASH ( XPLT , Z L5C , NOAYS , T . 02 ) GO TO 300 C C C c COMPARISCN OF MEANS 5 CM UNDER 10 CM GRASS 9 CALL AXES ( 3.0 , RC7 ( 3 ) ,4 , 0. 6 , 6 , I , LC7 ( 5 ) , 5 , 0. 6 , 1 2 ,2 ) CALL SCALE ( YHA I , NC AYS , 3. 00, 40.0,-10.0,15) CALL SCALE ( YL A I , NC AY S , 3 . 00, 40.^,-10.0,16) CALL SYMBL4 ( -0. 2 5 , 1 .0 , 0. 0 7 , « TEMPE R AT URE ( JO • , 90 . 0, 16 ) CALL SYMBL4 ( 0. 2 , 2. 8 ,0 . I , • CCMPA R J SON OF DAILY MEAN TEMPERATURES" A CALL AER', CALL A CALL CALL CALL CALL CALL CALL SYMBL4 SYMBL4 DATE LEGEND ZERO LINES DASH SYMBL4 0.0,37) (0.2,2.7,0.07, '5 CM DEEP IN SOIL UNDER 10 CM GRASS COV 0.0,41) (0.2,2.6,0.07, 'ON IRRIGATED AND NON- I PR IGATE 0 PLOTS', 0.0,36) (NYEAR ,0.2,2.45) (3,6.5,2.8) (0.6) ( XPLT, YHA 1 , NO AYS, 0.01, 2) ( XPLT.YLAl ,NDAYS,0.02) (8-32, 1.0,0.07, 'TEMPERATURE ( JF ) • , 90 . 0 , 16 ) GO TO 300 .«««**«******»PLGX 10***»**** ***♦♦♦*«■=• **♦**♦*•*♦♦♦♦♦♦♦♦♦♦♦ ***♦***> COMPARISON OF MAX AND MIN TEMP AT SOIL SURFACE 0120 0121 0122 0123 0124 0125 0126 0127 0128 0129 0130 0131 0132 0133 0134 0135 0136 0137 0 138 0139 0140 0141 0142 ")143 10 CALL CALL CALL CALL CALL C«LL CALL AATUR CALL A* . AXES SCALE SCALE SCALE SCALE SYMRL4 SYM6L4 ES' , SYMBL4 CALL SYMBL4 CALL CALL CALL CALL CALL CALL CALL CALL GO T DATE LEGEND ZERO SYMBL4 LINES LINE CASH DASH 0 3 )0 (3.5,RC7(3),4,C.5,8,l,LC7(5),5,0.5,16,2) (YHSS,N0AYS,3.50, 60.0,-10.0,17) (YLSS,NCAYS,3.50, 60.0,-10.0, 18) ( ZHSS,NDAYS,3.50, 60.0,-10.0,19) (ZLSS,NCAYS,3.50. 60.0,-10.0,20) (-0.25, I. 3, J. 0 7, 'TEMPERATURE (JC)',9J.0,16) (0.2, 3. 3,0. 1, 'COMPAR! SON OF MAXIMUM AND MINIMLIM T^ 0.0,46) (0.2,3.2,0.C7, 'AT SOIL SURFACE UNDER 13 CM GRASS C 0.0,39) ( T. 2, 3. 1,0. 07, 'ON IRRIGATED AND NON- I RR IGATE 0 PLOT 0.0,36) (NYEAR ,0.2,2.95) (2,6. 75,3. 3) (0.5) (8. 32, I. 3, 0.0 7, 'TEMPERATURE (JF)' ,90.0, 16) ( XPLT ,YHSS,NDAYS,0.01,3) (XPLT, YLSS .NDAYS, 1) (XPLT.ZHSS ,NDAYS,0.04) (XPLT, ZLSS,NDAYS, 0.02) MPER OVER S' , i^:^*** A««*> '♦♦***PLOT 11* ♦♦«*******♦*«**♦«****** ♦**♦*♦♦♦«***♦♦: COMPARISON OF MEANS AT SOIL SURFACE 11 CALL AXES CALL CALL CALL CALL (3.'>,RC7(3),4,C.5,7,l,LC7(5),5,0.5,l4,2) ( YHA?. NDAYS . 3. "10. 5 ') . n . - 1 '1. U ? 1 1 AXES (3.'>,RC7(3),4,C.5,7,l,LC7(5),5,0.5,l4,2) SCALE ( YHA2,NDAYS,3.00, 50. n , - I '). 0 . 2 1 ) SCALE (YLA2, NCAYS,3.00, 50.0,-10.0,22) SYM6L4 (-0.25, 1.0,0.07, 'TEMPEPATUPt: ( JC ) ' ,90 . 0 , 16 ) SYMRL4 ( 0.2,2.8 ,0. I, 'CCNPARISHN PF DAILY MEAN TEMPERATURE 0.0,37) CALL SYMRL4 ( 0 . 2 , 2 . 7 ,0 . 07 , ' AT SOIL SURFACE UNDER 10 C '^ GRASS', 0.0,33) June 1975 andersen, roper: meteorologic data 225 01'*'^ CALL SYMBL4 (0 . 2 , 2. 6 , 0. 07 , • OM IRRIGATED AND NGN- I KR I G ATFn PLOTS', A ).l,3b) 0145 CALL DATb ( NYE 6P , 0 .2 , 2. 4 5 ) 0146 CALL LEGEND (3,6.5,2.8) 0147 CALL ZERO <').5) 0148 CALL SYMBL4 ( 8 . 32 , I . 0, 0. 0 7 , • TE MPEP A TUR E ( J F ) • , 90 , 0 , 16 ) 0149 CALL LINES ( XPL T , YH A2 , NDA YS , 0. 11 , 2 I 0150 CALL CASH ( XPLT , YL 42 , NC AY S , 0 . 02 ) 0151 GO TO 300 C C C ♦**,«**«**»«* «*«««PLfjT I 2********'*^ ****************<'*********** ********** C CUMP/KISLN OF MAX AND MIN ON BARE GROUND C c 12 CALL AXES ( 5. 0 , P C 7 ( 2 ) ,4 , 0 . 5 , 1 1 , 1 , LC 7( 3) , 5 , 0 . 5 , 22, 2) CALL SCALE ( YHBG , N F. AY S , 5 . 00 , 8 ). 0 , -2'^ . 0, 2 3 ) CALL SCALE < YLBG , NC AY S , 5 . 00, 80.0,-20.0,24) CALL SCALE ( ZHRG, NC AYS , 5. 00, 80.0,-20.0,25) CALL SCALE ( ZLBG,NCAYS,5.00, 8 1. ) , -2 0. 0, 26 ) CALL SYMBL4 ( -0 . 25 , 2.0 , 0. 0 7 , • TE MPb P ATURE ( JC ) • , 90 . 0 , I 6 ) CALL SYMBL4 ( 0. 2 ,4 . 8 , 0. I , ' C CMPA R I SON OF MAXIMUM AND MINIMUM TEMPER \A TURFS', 0.0,46) CALL SYMBL4 ( 0 . 2 , 4 . 6 fc, 0. 08 , • AT SOIL SURFACE UN BARE GROUND CN', A 0.0,33) CALL SYMBL4 ( 0 . 2 ,4 . 52 , C. 0 8 , • 1 P P I G AT ED AND NON- I RR I GAT ED PLOTS', A 0.0,33) r&LL DATE {NYFAR,0.2,4.37) CALL LEGEND (2,6.75,4.8) CALL ZERO (l.O) CALL SYMBL4 ( 8 .32 , 2 . 0, 0. 07 , ' TE MPER ATURE ( JF ) • , 90 . D , 16 ) CALL LINES ( XPLT , YHBG ,NDAYS, 0. 01 , 3 ) CALL LINE ( XPLT, YLBG, NOAYS, 1) CALL DASH ( X PL T , ZHBG , NDAYS , 0. 04) CALL DASH ( XPLT, ZLBG, NOAYS, 0.02) GO TO 3 JO r c C *♦ 4 »*♦♦♦♦♦ »*****+PLOT 13**«*4**«******** ***♦♦♦♦*****♦♦*♦**♦* «♦******♦*♦ r COMPARISON OF MEANS OF BARE GROUND C c 13 CALL AXES ( 3. 0 , R C 7( 3 ) , 4 , 0 . 5 , 7 , I , LC7 ( 5 ) , 5 , 0. 5 , I 4 , 2 ) CALL SCALE ( YH A3 , NC AY S , 3. 00, 50.0,-10.0,27) CALL SCALE ( YL A 3 , NC AY S , 3. 00, 50.0,-10.0,28) CALL SYMBL4 ( -0. 25 , I .0 , 0. 0 7, ' TE MPE R ATURE ( JC ) • ,90 .0 , I 6 ) CALL SYMBL4 ( 0 . 2 , 2 . 8 ,0 . I , ' C CMPARI SON OF DAILY MEAN TEMPERATURES', A 0.0,37) CALL SYMBL4 ( 0. 2 , 2. 7 , 0. 07 , • ON BARE GROUND CN I RR IGATED' , 0 .0 , 2 7 ) CALL SYMBL4 ( 0. 2 , 2. 6 ,0. 07 , ' AND NON- I RR I GA TED PLOT S ' ,0. 0 , 2 3 ) CALL DATE ( NYE AR , 0 .2 , 2. 45 ) CALL LEGEND (3,6.5,2.8) CALL ZERO (0.5) CALL SYMBL' CALL LINES ( XPL T , YHA3 , NOA YS , 0. 01 , 2 ) CALL CASH (XPLT, YLA3,NDAYS, 0.03) GJ TO 300 E^D BLOCK DATA COMMON /PLOTO / XPLT(366) , ITAPE A /PLCTl / RAIN(366) , XIPR(366) , YIRR(366) , YNIR(366) B /PL0T2 / SNCW(366) C /PL0T3 / YHHO( 366) , YLHD( 366) 0 /PL0T4 / HUM0{366) E /PLQT5 / EVAP(366) F /PL0T6 / WINO( 366) G /PL0T7 / YMAX( 366) , YMIN( 366) H /PL0T8 / YH5C(366) , YL5C(366) , ZH5C(366) , ZL5C(366) I /PL0T9 / YHAK 366) , YLAl(366) J /PLOTIO/ YHSS(366> , YLSS(366) , ZHSS(366) , ZLSS(366) K /PLUTll/ YHA2( 366) , YLA2(366) L /PLGT12/ YHBG(366) , YLBG(366) , ZHBG(366) , ZLBG(366) M /oi nri 4/ VWA -Xl ^AA> - VI A ^r i<^A« 226 0003 0004 0005 0006 GREAT BASIN NATURALIST 0012 0013 0014 Vol. 35, No. 2 COMMON /LABELS/ AXl(37), AYI(37) DATA RC2.LC2 DATA RC3,LC3 DATA RC'«,LC4 DATA RC5,LC5 DATA RC6,LC6 DATA PC7,LC7 DATA BX.BY DATA XL.YL DATA AXl.AYl DATA AX2 DATA AY2 BY(4) , Lri(7) TAPE UNIT NUMBER DATA ITAPE DATA RCl.LCl BX(4I , RCl(7) RC3(ll), LC3(ll) RC5{ll), LC5(18) PC7(12), LC7(24) /5/ AX2(73)t AY2(73), XL(4), YL(4), RC2(13) , LC2( 13) , RC4( 13) , LC4( 13) , RC6(8), LC6(16), /• 0 •, • _t , • 25-', ' -• » • 50-', ' -• t • 75-', • 0 ', • - ' , ■- I ' , '- ' , '- 2 ', ' - * » t 3 1/ /' 0 ', • 25-', • 50-', ' 75-' , '100-', •125-', • 150-', •175-', '200-', •225-' , •25D-', '275-', •300-«, • 0 ', •- I ' , •- 2 •, •- 3 ', '- 4 ', •- 5 ', •- 6 • , '- 7 ' , •- 8 ' , i_ 9 • , '- 10' , •- 11', •- 12'/ /' 0 ', • 10-', ' 20-', ' 30-' , • 40-', • 50-', • 60-', • 70-', ' 8 0-', ' 90-' , • 100-' , ' 0 ', •- 10', •- 20«, •- 30', '- 40' , '- 50', '- 60', •- 70', •- 80», '- 90' , •-100'/ /• 0 •, . 2-', • ^,-1 , • 6-' , ' 8-' , • 10-', • 12-', • 14-', ' 16-', ' 18-', ' 20-', • 22-«, • 24-', • 0 ', •- 2 • , •- 4 ' , '- 6 ' , '- 8 ', •- 10', •- 12', '- 14', •- 16' , •- 18', '- 20', '- 22^, •- 24'/ /• 0 ', ■ 2-', • ^_t, • 6-' , • 3-' , ' 10-', • 12-', ' 14-', • 16-', ' 18-', ' 20-', ' 0.', •0 ', •- 0.', •1 •- 0.' , •2 •- 0.', •3 •, '- 0.', '4 ' , '- 0.' , •5 '- 0.', •6 •, •- 0.', •7 ', •- 0.', •8 '/ /• 0 •, • 40-', ' 8 0-', •120-' • 160-' , •200-', •240-', '280-', t 1 , '0 •- 2' , *5 • , •- 5* , '0 ', •- 7', •5 • , '- rv. •0 •- 12* , •5 ', '- 15' , •0 • , •- 17', • 5 •/ /•-30-', '-20-', •-10-' , • 0 ' • 10-', • 20-', • 30-', • 40-', ' 50-', • 60-', ' 70-', • 80-', ._ _2., '2 • - -' , '4 ' •- I' , •4 ', •- 3'. '2 '- 5', '0 • •- 6' , '8 ' , •- 8', •6 ' , '- 10' , '4 • '- 12' , • 2 • , •- 14«, •0 •- 15' , •8 • , '- 17' , '6 '/ /2*0.0, 2*7.94, 4*0.0/ /3.99, 2*0.0, -0.04, 2*0.0, 2*1.34/ / 2*7. 94, 3* 7. 2 7, 3*6. 6 1,3* 5. 94, 3*5. 29, 3*4. 61 , 3*3.9 4,3*3.29,3*2.61,3*1.96,3*1.2 8,3*0.67,2*0.0, 0.03,2*0.0,0.03,2*0.0,0.03,2*0.0,0.03,2*0.0, 0.0 3,2*0. 1,0.03,2*0.0,0.33,2*0.0,0.03,2*0.0, 0.0 3,2*0. 0,0.03,2*0.0,0.0 3,2*0.0,0.03,2*0.0, 0.03/ /2*7.S4, 3*7.60,3*7.2 7,3*6.94 3*5.9 5, 3*5.62,3*5.2 9,3*4.95 3*3.94,3*3.62,3*3.29,3*2.95 3*1.96,3*1.62,3*1.2 8,3*0.98 /- 0.03, 2*0. 0,-0. 0 3, 2*0. 0,-0. -0.03,2*0.0,-0.03,2*0.0,-3. -0.03,2*0.0,-0.0 3,2*0.0,-0. -0.0 3,2*0.0,-0.0 3,2*0.0,-0. -0.03,2*0.0,-0.03,2*0.0,-0. -0.03,2*0.0,-0.0 3,2*0.0,-0. -C.03/ ,3*6.61, 3*6 28 1 ,3*4.61, 3*4 28 ,3*2.61, 3*2 29 ,3*0.67, 3*0 34 '2*0 3/ 03 2*0.0 ,-0 03 ,2*0 0, 03 2*0.0 ,-0 03 2*0 0, 03 2*0.0 ,-0. 03 2*0 0, 03 2*0.0 ,-0 03 2*0 0, ^1> 2*0.0 ,-3 33 2*0 0, 0 3 2*0.0 ,-0. 03 2*0.0, 0002 0003 SUBROUTINE AXES HT,X,NXC,DX,N1,M1,Y,NYC,0Y,N2,^'2) GRAPH A BOX OF HEIGHT HT WITH THE MONTHS PLOTTED ALONG THE BOTTCM. ALSO LABEL THE BOX WITH APPAY X ON THE RIGHT AND Y ON THE LEFT. UX IS THE DISTANCE BETWEEN EACH LABLE. Nl IS THE NUMBER OF LABELS ANO Ml IS THE STEP BETWEEN LABLES. THE SAME IS TRUE FOR ARRAY Y WITH CY, N2, AND M2. DIMENSION X(N1), v(N2) COMMON /LABELS/ axi(37), A BX(4J , DATA NCCUNT /I/ AY1(37) , BY(4) AX2(73), AY2(73) June 1975 ANDERSEN, ROPER: METEOROLOGIC DATA 227 0005 0006 3007 0008 0009 0010 0011 0012 0013 0C14 3015 0016 0C17 0018 0019 0C20 0021 0022 0023 0024 3025 0026 0027 3028 0029 0J33 0031 0032 0033 0034 0035 0036 0037 !F = HT BY(3) = HT CALL SYMBL*, (3.0,0.03,0.07,' JANUARY APRIL MAY JUNE JULY BT08ER NOVEMBER DEC EMBER • ,0. 0, 1 3 1 ) CALL LINE ( AX1,AYI,37, 1) CALL PLOT (0.0,0-27,-3) CALL SYMBL4 ( 0- 0 ,-0. 1 1 , 0. 0 7 , • 1 15 1 A 15 I 15 1 15 1 15 1 B 15 I 15 1 15", 3. 0,123) CALL LINE ( AX2,AY2,73, 1 ) CALL LINE (BX,BY,4,1) OYY = -0.03 DO 12 I=1,N2,M2 CALL SYMBLA ( 7.9 ,DYY ,0. 07 , Y 1 I ) , 0.0 , NYC » OYY = OYY ♦ DY 12 CONTINUE DXX = -0.03 DO 13 I=l,Nl,Ml CALL SYMBL4 ( -0 . 22 , DXX , 0. 07 , X ( I) , 0 . 0 , NXC ) OXX = DXX *■ DX 13 CONTINUE GU TO 14 DRAW ZERO LINE AT HEIGHT HT. ENTRY ZERO (HT) BY(2) = HT BY(3) = HT CALL LINE (BX( 2) ,RY(2) ,2,1) 14 RETURN END FEBRUARY AUGUST MARCH A SEPTEMBER OC SUBROUTINE DASH (X.Y.N.S) C C C C SUBROUTINE 0*SH WILL DRAW X = X ARRAY c Y = Y ARRAY c N = NUMBER CF POINTS c c S = LENGTH OF DASHES 3002 DIMENSION X(N), YCN), A(2) 0003 SS = S ♦ 2.0 0004 DO 30 1=2, N 0005 DX = X( I) - X( I-l) 0006 DY = Y( n - Y( I-l) 0007 H = SQRT(DX*DX «■ OY*DY ) 0C08 DX = S * DX / H 0009 DY = S » DY / H 0010 A( I) = X( I-l) 0011 B( I) = Y( I-l) 0012 n CONTINUE 0013 IF(H.LE.S) GO TO 20 0014 A(2) = A( 1) + DX 0015 8(2) = B( I) ♦ DY 0016 CALL LINE (A,B,2,1) 0317 IF(H.LE.SS) GO TO 30 0018 A( I) = A{2) + DX 0019 B( I) = B(2) ♦ DY 0020 H = H - SS 0021 GO TO 10 3322 20 A(2) = X( I) 0023 6(2) = Y( I) 0024 CALL LINE (A,B,2,1) 3025 30 CONTINUE 228 0026 GREAT BASIN NATURALIST Vol. 35, No. 2 0027 0028 0029 0030 0031 0032 0033 J035 0036 0037 0038 C039 DOAO 0041 0042 0 343 0044 0045 SUBROUTINE LINES WILL MAKE THE LINE DARKER AND WIDER BY DRAWING N NUMBER OF LINES X = X ARRAY Y = Y ARRAY N = NUMBER OF POINTS D = DISTANCE BETWEEN LINES M = NUMBER OF LINES ENTRY LINES (X,Y,N,0,M) DY =0.0 DO 40 1=1, M CALL LINE (X.Y.N.I) CALL PLOT (0.0,0,-3) OY = OY - D 40 CONTINUE CALL PLOT (0.0,DY,-3) GO TO 99 ENTRY BAR (X,Y,N) SS = 0.0 CALL PLOT (X(1),SS,3) DO 50 1=2, N CALL PLOT (X( I-l),Y( I) ,1) CALL PLOT (X( I », Y( I ) ,1) 5J CONTINUE CALL PLOT (X(NI),SS,I) 99 RETURN END 0001 SUBROUTINF DATE (NYR.X.Y) GRAPH DATE SPECIFIED BY NYR AT POINT (X,Y) 0002 0003 0004 0005 0006 3007 J008 0C09 oon 0311 0012 0013 0014 0016 >3017 0018 0319 JJ2T 0021 0 322 0023 0024 3026 J 326 0027 0028 CALL SYMBL4 ( X , Y , 0. C 7 , ' PRC VD , UT AH , • , 0. 0 , 12 ) N = NYR-1969 GOTO (1,2,3,4,5), N 1 CALL SYMBL4 ( X+0. 7 6, Y, 0. 07 , ' 1970 • , 0. 0 ,4) RETURN 2 CALL SYMeL4 ( X ♦•O . 7 8 , Y , C. C 7 , • 1 9 7 1 • , 0. 0 , 4 ) RETURN 3 CALL SYMBL4 ( X+0. 78 , Y, 0. 0 7 , • 1972 • , 0. 0 ,4) RFTUPN 4 CALL SYMBL4 ( X + J . 78 , Y , 3. 0 7 , • 1 973 ' , 0 . D , 4) RETURN 5 CALL SYMBL4 ( X* 3 . 7 8 , Y , 0. 0 7 , ' I 9 74 ' , 0 . "> , 4 ) RETURN ENTRY LEGEND (N,X,Y) GRAPHS ONE OF FH'JR LEGENDS SPECIFIED BY N AT POINT (X.YI G3TQ ( 10, 1 I, 12, 13) ,N 10 CALL SYMPL4 ( X , Y , 0 . 07, ' f- A X I MUM • , 0 . 0 , 7 ) CALL SYMBL4 ( X , Y-0. 1 , 0. 07 , • M I N I MU^' • , n . 3 , 7 ) RETURN 11 CALL SY'^eL4 ( X , Y ,0 . 0 7 , • MA x NCN- I RR • , 0 . 0 , 1 1 ) CALL SYMBL4 ( X , Y-0. 1 , 0. 07 , • M IN NON- I RP • ,0 . 0 , 1 I ) CALL SYMBL4 ( X , Y-0 . 2 2, 0. 07 , • f A X I PR I GA TED • , 0. 0 , 1 3 ) CALL SYMBL4 ( X , Y-0. 32, 0.0 7 , ' M I N I PR I GATED ' , 0. 0, 13 ) RETURN 12 CALL SYMBL4 ( X , Y ,0 . 3 7, • ME AN NOr'- I RR IG A TED • , 0. 0 , 1 8 ) CALL SYMBL4 ( X , Y-0. 1 ,0 . 37 , • Mf AN I RP IG ATED* , J. 3 , L 4 ) 13 RETURN END 0001 SUBROUTINE SCALE ( X , N, S , YM AX , YM I N , I D) S'"ALE DATE IN A^KAY X DIMENSICNFO BY N AND RETURN SCALEO OMA IN X. S IS THE MAXIMUM HEIGHT OF SCALED DATA, YMAX HAS MAXIVUM ALLOWED DATA AND YMN THE KIMMUM. SHOULD DATA EXC>=FD THESE BOUNDS, THE ID NUMBER WILL BE PRINTED ALONG WITH THE DATA. June 1975 andersen, roper: meteorologic data 229 O0T2 DIMENSION X < f' ) 0003 SS = S / ( YMAX-YMNJ TJ34 on 10 1=1, N 00^5 IF(X( I ) .LT.VMINJ WRITE(6,l-jn ID, X(I), YMIN 0006 IF(X( n.GT.YMAX) VIRITE(&,100) ID, X(T), YMAX 0007 X( I ) = SS » ( X( I) - YMIN) 0CC8 10 CCNTINUE 3:39 RFTURN 0010 no FORMAT!' CALL • , 1 2 . 5 X , F I 0. 2 , • EXCFEOED '.Fg.D 0011 lOI FORMAT(« CALL • , I 2 , 5 X, F 10 . 2 , • WAS BELCW '.FQ.l) 3312 END OCjI SJ'ShLuTI.^L .M-AuC ( I Y£A^ ,i\CAYS) C r, KcAu Ifi'z Y£Ak'b uiTA SPECIFIEU BY lYEAR. THE FIRST CARD IS i CATA C HEAL.Lr, LUi\,TA INI M:, THE Y£AK, NUMBER OF DAYS IN YEAR AND LAST C kECukDEG WlNb VALUE CE PRtVlCUS Yi^AR. RETURim NJMOlR UF DAYS IN C NO AYS. C CrC2 CLM^.-JN /PLCK / XPLTOto), ITAPt A /P^GTl / TPRE(3oo), XlRROfab), YlRRCjOb), YNIP(36d) P /PLCT2 / Si\CV(366) C /PLLT3 / Hho^(3cfc), LHUN(3o6) D /PLCT4 / THPS<'3o6) E /PLLT5 / PEVP(j.6fcJ F /PLCTo / wINC(j)o6) G /PLCT7 / YMAX(366), YMlN(3b6) H /PLJTo / YH5C(JC6), YL5C(j6t), LhbCiibti, ^LbC(36b) I /PLUT9 / YHAl(366), YLA1(366) J /PLCTIO/ YpSS(366), YLSS(366), ZHSS136o), ZLSS(36o) K /PLuTU/ YHi2(:>ob), YLA^(366) L /PLJTi2/ YhfcG(366), YLBG(366), ZhBG(36o), ZLB3(366) M /PLOUi/ YHA3<366), YLA3(366) 0C03 CGMHCN /LABELS/ DUNY(22b), XL(A), YL(4J CCO^ REAL CbUl) /• 0 ', • -', • 10-', • -', • 2 3-', ' -', A ' 30-' , • -', • ^0-', • -• , ' 50 •/ Jjyj REAL FHUM, LhLf'', M k Vv , M C V. , IRnW, IROW ,OLDw OCCo AVE(X,Y) = (X+YJ/^.C C ->j7 1 PEAu( ITaPl ,2J J,LND=?9>7 ) I YF , NDA YS , CLUV^ Ci-Oe 200 F3RKATi21A,F^.O) ::39 ifciyeak.ew.iyr) gu to 3 OCIO 00 ^ I=l,NCAYS DCll 2 KcAOd TAPE ,200, END = SS9) CC12 Gu TO 1 C C 0C13 3 DO ^ I=i,NoAYS 3314 REAC( ITAPc ,201 ,c,mD=';99) RAIN, SNOkJ, SNCVCI), IK^«, IRDW, NlViU, A NIDU, HHUK(I), LHUM(I), THRS(I), PtVP(I), B LLCV, UINC(I), YMAX(I), YMINCH, C YH5C(!), YL5C(I), YHSS(I), YLSS(I), YHBGd), D YLBG(I), ZH5C(I), ZL5C(I), ZHSS(I), ZLSS(I), E ZhBG(I), ZLBGd) ) n-J 2 31 FQRNAT(bX,F3.2,4F3. 1,2X,2F3.1,1X,2F3.C,F2.0,F3.1,F2.0,F3.0,/, A 5X,^F3.0,bX, 12Fi.O) CClo XPLT(I) = I .:317 TPRE(n = RAIN •»• SNLVs*C.l 0C13 YIRR(I)=-i.J 0319 YMR< I ) = -1.0 0C20 IFINI wW.NE . J.O J YNIR(I) = < N IWW-N I DW ) * IJO . j/NIJW C021 IFdRww.NE.O.O) YIRR(I) = d RWW- I RDW ) * I JO. J/IRCW ::22 lF(uLJW.GT.ksINOd ) ) CLClv = OLDW - ICOO.O 0C23 HOLD = WIND(I) 0C24 wIND(I) = (wINi.( n-CLDW) * l.6093«-0.05 3025 DLOW = HOLD 0026 YHAKIJ = AVE( YHijC< I ),YLbCd)) 3327 YLAKIJ = A VE ( ZH5C ( I ) , ZL5C ( I ) > ^C2d YHi2d) = iVt ( YHSS( I), YLSSI n ) ^C^9 YLi^«I) = AVE(ZHoS( n.ZLSSd) J 3)jJ YhAj)(I) = AVt< Yhoo( I ), YL'3Gd ) I CC31 YlAjJI) = AVE< ZHB.:,( I ),ZL&G( n ) 230 GREAT BASIN NATURALIST Vol. 35, No. 2 0C3^ 4 COMIi\Ub JCiJ Y3AVS = MJ>S 0CJ4 CALL SCfiLL tXPLl ,\[./^YS ,7.94,YCiYS, I.'., U r-Tss -itTLKN J-J6 9 99 ^»o ! Tfc (6,5 )j) 0C^7 500 FCkMAK •-• »**'»»-*i''( LRHCR IN INPUT DATA fhClA F AP c: * v** *• , / / ) ::3o C srjp CC39 c c.n;TKY must d:;^j CALL PLOT (u. 0,1. 2, -3) :j4i N = 0 0C42 DJ ZL 1=12 1,^06 J J-ifi iFiYikh (I) .LT. J..) g: tc z: cc^^ N = K + 1 CC45 YIFR(NJ = Y1K5( I ) CC46 XIRk (N) = XPLT ( I ) 0C47 20 CONTINUE 0C4b CALL SCALE (YIRF ,,>j, l.^d,50.0,C.0,29) CC49 CALL LINt (XlKR,YlKr,,\,l) CC5C K = C J Obi OC JJ I=U1,jOo 0C52 1 F( YMR<1 > .LT.C. 0) GC TO 30 j:i)3 N = N + 1 jC54 )ttam was a bibliophile; he built up a complete set of ornithological jour- nals and textbooks on American birds for the use of students at the refuge. He was also concerned with the migration, breed- ing, and feeding of game birds and cam- paigned for the j)rotection of rare and vanishing species, such as hawks, owls, eagles, whooping cranes, and brown pel- icans. He constantly emphasized the ne- cessity of ha\ing a broad, well-grounded program, based upon verifiable facts, when dealing with conservation problems. Clarence also found time to coauthor, with Dr. Angus M. Woodbury and ,Tohn Sudgen, a manuscrijit dealing with the birds of Utah. The untimoh', accidental death of Dr. Woodbury, the senior author, delayed the publication of this important treatise. Through the efforts of Dr. Cot- tam this manuscript was turned over to Dr. C. Lynn Hayward of the Department of Zoology at BYU to review, edit, and publish. Clarence had the help and companion- ship of a gracious, artistic helpmate and wife, Margery Brown Cottam, whom he married on May 20, 1920, while they were both teaching at Alamo, Nevada. The}^ were the parents of four girls: Glenna, Mrs. Ivan L. Sanderson of San Francisco, California; Margery, Mrs. Grant Osborne, Amherst, Massachusetts; Josephine, Mrs. Douglas S. Day, Salt Lake City, Utah; and Caroline, Mrs. Dwayne Stevenson, McClaine, Virginia. They had 23 grand- children and 4 great grandchildren. Mar- gery was an active member of the LDS Church, serving as stake YWMIA pres- ident and stake Relief Society president. She died February 28, 1975, and was buried beside Clarence at Orem, Utah. Always religious and devoted to the LDS Church, Dr. Cottam was a pillar of strength in the LDS Corpus Christi (Texas) Stake. He served as first coun- selor in the San Antonio Stake, after which he served for nine years as president of the Corpus Christi Stake. At the time of his death, March 30, 1974, he was pa- triarch of the Corpus Christi Stake. He was the essence of tolerance in his dealings with his fello\MTien. As an edu- cator, administrator, researcher, and inter- mediary between scientific groups and the the sports public. Dr. Cottam was most successful. 234 GREAT BASIN NATURALIST Vol. 35, No. 2 Dr. Clarence Cottam's name is indelibly inscribed on the roster of distinguished alumni of Brigham Young University. For the past 50 years I have enjoyed \vatching him become one of the most honored biologists of the nation. The long list of his publications which follows reveals his wide experience and insight in providing solutions to man's mistakes in dealing with his animate en- vironment. Published Biographical Reports of Clarence Cottam Anonymous. 1974. Clarence Cottam: Nation- ally acclaimed conservationist. National Parks Conservation I^lagazine 28:26. Bolen, Eric G. 1975. In memoriam: Clarence Cottam. Auk 92:118-125. Bibliography of Clarence Cottam 1928-1975 For the preparation of the following list of publications by Dr. Clarence Cot- tam, I am obligated to Mr. Douglas S. Day, Dr. Cottam's son-in-law. 1928. 1928. 1928. 1929. 1929. 1929. 1929. 1929. 1930. 1930. 1931. 1931. 1931. 1932. Christmas bird census, Provo, Utah. 1927. Bird Lore 30(1): 65. Killdeer swimming on Green River. Utah. Auk 45 (2): 207-208. White pelicans and great blue herons win- tering in northern Utah. Condor 30:160. April. Christmas bird census, Provo. Utah. 1928. Bird Lore 31(1): 61-62. A shower of grebes. Condor 31 (2): 80-81. March. The status of the ring-necked pheasant in Utah. Condor 31 (3):117-123. May. The fecunditv of the English sparrow in northern Utah. Wilson Bull. 41:193-194. September. A studv of the water birds of Utah Countv. Utah. Proc. LTtah Acad. Sci. 6:8-11. July. Food habits of the shoal-water and diving ducks of Florida. Florida Woods and Wa- ters 1(2): 37-38. Summer. A friendly humming bird. Bird Lore 32 (5):352. September-October. Montlil}" and yearly percentages of food, page 124; Animal food, pages 158-159; In- dex, pages 547-559 in H. L. Stoddard, The bobwhite quail: its habits, preservation, and increase. Charles Scribner's Sons, New York City. Some new and uncommon bird records for South Dakota. Wilson Bull. 43 (4): 311. Birds and motors cars in South Dakota. Wilson Bull. 43(4):313-334. (with F. M. Uhler, Leon Kelso, and E. R. Kalmbach). Christmas bird census. Port Tobacco, Maryland. 1931. Bird Lore 34 (1):49. 1932. (with Leon Kelso, and W. H. Ball). The Louisiana heron in Washington. D.C., region. Proc. Biol. Soc. Washington 45:207. 1932. Nocturnal habits of the chimnev swift. Auk 49(4): 479-481. October. 1933. (with F. M. Uhler. A. L. Nelson, and Leon Kelso). Christmas bird census. Port To- bacco, Maryland. 1932. Bird Lore 35 (1):30. 1933. (with Phoebe Knappen). Oil gland usually tufted in Hj'dranassa tricolor ruficollis. Auk 50(l):94-95. 1933. (with Leon Kelso). An incubating wood- cock. Auk 50(2): 170-173; 4 pi. 1933. Night migration of eastern chipping spar- rows. Bird Banding 4(l):54-55. 1933. Feeding habits of the lesser scaup duck. Condor 35 (3): 1184 19. 1933. Winter records for the coastal region of North Carolina. Auk 50(2) : 231 -232. 1933. A pot cock pheasant. Bird Lore 35(3): 148. 1933. Disappearance of eelgrass along the At- lantic coast. Plant Disease Reporter 17(6): 46-53. (Mimeographed, Bureau of Plant Industry.) 1933. The blue goose in North Carolina. Auk 50(3): 353. 1933. Recent observations on eelgrass conditions. Plant Disease Reporter 17(10) :119-120. 1933. Further reports on eelgrass. Plant Disease Reporter 17(11). 1933. Nelson's sparrow and tree swallow winter- ing in New Jersej-. Bird Banding 4(2): 115. 1933. Spring migration of the great blue heron. Auk 50(4):427-428. 1934. Summer bird records for North Cai-olina. Auk 51(0:94-96. 1934. Incursion of American scoters in Norfolk. Connecticut. Auk 51 (2) : 228-229. 1934. Possible e.xtension of regular winter range of the great black-backecl gull. Auk 51(3): 376. 1934. Past periods of eelgrass scarcitv. Rhodora 36(427) :261-264. 1934. The eelgrass shortage in relation to water- fowl. Trans. American Game Conf. 20:272- 279. 1934. Eelgrass disappearance has serious effects on waterfowl and industry. Yearbook of Agriculture (1430) : 191-193. 1934. Summary of reports of baiting and other gunning practices. Bureau of Biological Survev 17 pp. (Unsigned, mimeographed.) 1935. (withT. H. Scheffer). The crested myna, or Chinese starling, in the Pacific North- west. USDA Tech. Bull. 467. 27 pp; 3 pi., 2 fig. 1935. The present situation regarding eelgrass (Zns/eia marina). Wildlife Research and Management Leaflet BS-3. 7 pp. 1935. (with F. M. Uhler). Mosquito control and its effects on aquatic wildlife. Proc. Amer. Game Conf. 21:291-294. 1935. The effects of arsenic, as used in poison- ing grasshoppers, upon birds. .Xuk 52(1): 118-119. 1935. Wasting disease of Zoslrra marina. Nature 135(3408):, 306. 1935. Late migration of tree swallows and mourning doves. Auk 52(2): 189. 1935. Umjsual food habits of California gulls. Condor 37(3): 170-171. Juno 1975 TANNER: CLARENCE COTTAM 235 1935. The eelgrass situation in 1934. Proc. Anier. Game Conf. 21:295-301. 1935. Winter notes from coastal North Carolina. 1937. Auk 52(3):318-319. 1935. Waterfowl problems rlarified by stud\' of 1937. gunning practices. Yearbook of Agriculture (1519):328-330. 1935. The present eelgrass situation along the American Atlantic Coast. Plant Disease 1937. Reporter 19( 14) :230-231. (Presented by Neil Stevens at the International Botanical Conference at Amsterdam. August. 1937. 1935. Further notes on past periods of eelgrass scarcity. Rhodora 37(440) : 269-271. 1935. Blue and snow geese in eastern United 1937. States in the winter of 1934-35, with notes on their food habits. Auk 52(4) : 432-441. 1935. (with F. M. Uhler). Bird records new or 1938. uncommon to Maryland. Auk 52(4) .-460- 461. ^ 1938. 1935. Review of Food preferences and require- ments of the white-tailed deer in New York State, bulletin by L. A. Maynard. 1938. Gardiner Bump, Robert Darrow, and J. C. Woodward. J. of Mammal. 16(4) :330-331. 1936. (with I. N. Gabrielson. A. L. Nelson, and 1938. E. R. Kalmbach). Christmas bird census. Port Tobacco. Maryland. Bird Lore 38(1). 1936. (with F. M. Uhler). The role of fish- eating birds. Progressive Fish Culturist. 1938. M. 1-131 (14.3202): 1-14. (Multigraphed. Bureau of Fisheries.) 1936. Earlj- migration of the great black-backed gull. Auk 53(1):81. 1938. 1936. Economic oinithology and the correlation of laboratory and field methods. Wildlife Research and Management Leaflet BS-30. 1938. 13 pp.; 1 fig. 1936. Food of the limpkin. Wilson Bull. 48(1): 1938. 11-13. 1936. Food of arctic birds and mammals col- lected by the Bartlett Expeditions of 1931- 32-33. J. Washington Acad. Sci. 26(4) : 165- 177. 1938. 1936. Notes on tlie birds of Nevada. Condor 38 (3): 122-123. 1936. The place of food habits research in wild- 1938. life management. Utah Juniper 7:16-19. 1936. Status of the black-backed gull. Auk 53 (3):332-333. ., 1938. 1936. Broken wing ruse in the yellow warbler. Auk 53 (4): 481. " 1939. 1936. (with H. R. Lewis). Eelgrass and other waterfowl foods: present status and future 1939. prospects. Proc. North American Wildlife Conf.. Wildlife Restoration and Conser- vation, pp. 498-500. 1936. Food habits of North American diving 1939. ducks. George Washington Univ. Bull. pp. 66-70. (Summarv of doctoral disserta- tion.) 1939. 1937. (with L N. Gabrielson, A. L. Nelson, and C. F. Smith). Christmas bird census. Port 1939. Tobacco. Maryland. Bird Lore 39(1 ):45. 1937. Review of Home life and economic status of the double-crested cormorant, bv H. L. 1939. Mendall. Auk 54(2) :21 3-214. 1937. An unusual concentration of blue jays. 1939. Bird Banding 8 (2): 79-80. 1937. Speed of the gray fo.x. J. Mammal 18(2): 1939. 240-241 . 1937. (with F. M. Uhler). Birds in relation to fishes. Wildlife Research and Manage- ment Leaflet BS-83. 16 pp. American egret and black-bellied plover in Delaware in winter. Auk 54(3) :382. (with J. J. Lynch). Status of eelgrass {Zostera marina) on the north Atlantic Coast, January. Wildlife Research and Management Leaflet BS-94. 15 pp. (with A. L. Nelson and C. S. Williams). Uncommon winter birds in coastal North Carolina. Auk 54(4): 548. (with A. L. Nelson). Winter nesting and winter food of the barn owl in South Carolina. Wilson Bull. 49(4) : 283-285. (with J. E. Shillinger). The importance of lead poisoning in waterfowl. Proc. North Amer. Wildlife Conf. 2:398-403. Coot swallowed b}' fish. Wilson Bull. 50 (1):60. (with W^ S. Bourn). Some effects of mos- cjuito control on wildlife. Conserv^ation 4 (2): 20-22. Review of Utah birds in control of certain insect pests, by G. F. Knowlton. Utah Acad. Sci. 14:159-166. 1937. Auk 55(2) :300. Review of Biological control of the beet leafhopper in Utah, by G. F. Knowlton, Utah Acad. Sci. 14:111-139. 1937. Auk 55 (2): 303. Status of eelgrass [Zostera marina) on the north Atlantic Coast. February. Wildlife Research and Management Leaflet BS-110. 7 pp. (with H. C. Hanson). Food habits of some arctic birds and mammals. Zool. Ser. Field Mus. Nat. Hist. .20 (31): 405-426. A fatal combat between a heron and a snake. Wilson Bull. 50(2): 140. The coordination of mosquito control with wildlife conservation. Proc. New Jersey Mosquito Exterm. Assn. 25:217-223. Wild- life Research and Management Leaflet BS- 119. 6 pp. (with W. S. Bourn). What's wrong with mosquito control? Trans. North Amer. Wildlife Conf. 3:81-87, 98-99. (with A. L. Nelson). Why study the food of fur animals? Trans. North Amer. Wild- life Conf. 3:527-530. Nesting of an eastern kingbird in a de- serted oriole nest. Condor 40(6) :259. Great blue heron swimming. Condor 41 (1): 37. (with I. N. Gabrielson, A. L. Nelson, and A. C. Martin). Christmas bird census. Port Tobacco, Marvland. 1938. Bird Lore 41 (l):22-23. (with Phoebe Knappen). Food of some un- common North American birds. Auk 56(2): 138-169. Late occurrence of nighthawk in Connect- icut. Auk 56(2):188. (with William Vogt, Victor Cahalane, and Aldo Leopold). Report of Committee on Bird Protection. Auk 56(2) :212-219. Food habits of North American diving ducks. USDA Tech. Bull. 643. 140 pp. The eelgrass situation on the American Pacific Coast. Rhodora 41 (487) :257-260. (with A. L. Nelson and T. E. Clarke). Nptes on early winter food habits of the black bear in the George Washington 236 GREAT BASIN NATURALIST Vol. 35, No. 2 National Forest. .T. Mammal. 20(3) :310- 1942. 314. 1939. (with W. S. Bourn). Need mosquito con- 1942. trol be incompatible with wildlife? Trans. North Amer. Wildlife Conf. 4:121-130. 130-140, 143. 1942. 1939. (with W. S. Bourn). The effects of lower- ing water levels on marsh wildlife. Trans. North Amer. Wildlife Conf. 4:343-350. 1939. (with F. M. Uhler and T. E. Clarke). 1942. Food of snakes of the George Washington National Forest. Trans. North Amer. Wild- 1942. life Conf. 4:605-622. 1939. (witli C. S. Williams). Food and habits of 1942. some birds nesting on islands in Great Salt Lake. Wilson Bull. 51 (3): 150-155. 1939. (with W. S. Bourn). Marshes of the past 1942. and future. Appendix F, pages 471-488 in Van Campen Heilner. A book on duck shooting. Penn Publishing Company. Phila- 1942. delphia. 1940. (with I. N. Gabrielson, A. C. Martin, and A. L. Nelson). Christmas bird census. Port 1942. Tobacco, Maryland. 1939. Bird Lore Suppl. 42(l):89-90. 1940. (with V. H. Cahalane, Aldo Leopold, and 1942. W. L. Finley). Report of Bird Protection Committee in 1939. Auk 57 (2): 279-291. 1942. 1940. Obituary of John William Sugden. Sr. Auk 57(3): 448. 1942. 1940. (with A. L. Nelson, and L. W. Saylor). The chukar and hungarian partridges in America. Trans. North Amer. Wildlife 1942. Conf. 5:432. Modern Game Breeding and Hunting Club News 10(7) :6-7, 12. Wild- 1942. life Research and Management Leaflet BS- 159. 6 pp. 1940. (with F. M. Uhler). Birds as a factor in controlling insect depredations. Wildlife 1942 Leaflet BS-162. 6 pp. 1941. Incubation feeding of calliope humming- 1943 bird. Auk 58(1): 59-60. 1941. (with L N. Gabrielson. A. C. Martin, and 1943. A. L. Nelson). Christmas bird census. Port Tobacco, Maryland. 1940. Audubon Magazine Supplement 43(1): 99. 1943. 1941. The eelgrass situation, fall 1940. Plant Dis- ease Reporter 25 (2): 46-52. 1943. 1941. LeConte's sparrow in Utah. Condor 43(2): 116-117. 1943. 1941. Indigo bunting and band-tailed pigeon in Utah. Condor 43 (2): 122. 1941. (with Phoebe Knappen). Eskimo curlew 1943. food note corrected. Auk 58(2) :256. 1941. Color attractive to hummingbirds. Auk 58(2) :261. 1943. 1941. (with V. H. Cahalane, W. H. Finley. and Aldo Leopold). Report of the Committee on Bird Protection. 1940. Auk 58 (2): 292- 1943 298. 1941. California cuckoo in southeastern Nevada. 1943. Condor 43(3): 160. 1941. How fast can a fox squirrel run? J. Mam- 1944. mal. 22(3): 323. 1941. (with C. S. Williams). Wilson snipe perches on telephone pole. Condor 43(6): 1911. 293. 1941. European starling in Nevada. Condor 43(6):293-294. 1941. (with James Moffitt). Eelgrass depletion 1944. on the Pacific coast and its effect upon black brant. Wildlife Leaflet 204. 26 pp. European widgeon at the upper Souris National Wildlife Refuge. Auk 59(1): 104. (with V. H. Cahalane, Aldo Leopold, and W. L. Finlev). Report of the Committee on Bird Protection. Auk 59(2) :286-300. (with I. N. Gabrielson, F. M. Uhler, and A. L. Nelson). Christmas bird census. Port Tobacco. Marvland. 1941. Audubon Maga- zine 44(1 ):27. Supplementary notes on the food of the limpkin. Nautilus 55(4) : 125-128. Records from extreme northeastern Neva- da. Condor 44(3): 127-128. (with C. S. Williams, and C. A. Sooter). Flight and running speeds of birds. Wilson Bull. 54(2):121-131. (with Seth Low, and R. E. Griffith). Glaucous gull in Oklahoma. Wilson Bull. 54(2): 139-140. (with C. S. Williams, and C. A. Sooter). Cooperative feeding of white pelicans. Auk 59(3):444-445. (with C. A. Sooter, and R. E. Griffith). The European starling in New Mexico. Condor 44(4): 182. Slate-colored junco in Nevada. Condor 44 (4): 185. Food of the ruddy turnstone. Auk 59(4): 581. (with C. A. Sooter. and R. E. Griffith). The yellow rail and the Caspian tern in New Mexico. Condor 44(5) :230. Coyote without external ears. J. Mammal. 23(4): 450. (with C. S. Williams, and C. A. Sooter). Some unusual winter visitors or late mi- grants to the Bear River Marshes. Utah. Great Basin Naturalist 3(2): 51 -53. New or uncommon Utah bird records. Wilson Bull. 54(4):254-255. Bullock's oriole as a fighter. Auk 60(1): 94-95. (with V. H. Cahalane, W. L. Finley, and Aldo Leopold). Report of Committee on Bird Protection. Auk 60(1): 152-162. (with C. S. Williams). Speed of some wild mammals. J. Mammal. 24(2) : 262-263. Greater vellow-legs as a fish-eater. Wilson Bull. 55(2): 128. Is the starling population decreasing in northeastern United States? Auk 60(3): 439-440. (with C. S. Williams, and G. H. Jensen). Some birds not commonlv observed in Utah. Condor 45(4): 159-160. (with L. E. Givens, and D. V. GrajO- Vermilion flvcatcher at St. Marks, Florida. Wilson Bull. 55 (3): 192. Ihiusual feeding habits of grnckles and crows. Auk 60(4) : 594-595. Tlie least bittern at Long Lake. North Dakota. Wilson Bull. 55(1): 54. (wath A. M. Woodbury). Type locality of Pcrisoreus canadensis capitalis Ridgway. Auk 61(1): 131 -132. (with J. J. Lynch, and A. L. Nelson). Food habits and management of American sea brant. J. Wildlife Mgmt 8(1): 36-56; 3 pi. - fwith C. C. Sperry). The greater and lesser yellow-legs as fish caters. Wilson Bull. 56(1):45. Iiiiie 1975 TANNER: CLARENCE CX)TTAM 237 1944. Birds and the west Tennessee River Lake. Migrant 15( 1):L5; 2 maps. 1944. Gulls as vegetarians. Condor 46(3) : 127-128. I 9 17. 1944. Eastern lark sparrow and upland plover in western Tennessee. Migrant 15(2) :29. 1944', Starlings feeding on the liacks of cattle. Migrant 15(2) : 24-25. 1947. 1944. The role of impoundments in 'post-uar plaiming for wateifowl. Trans. Noitli 1947. Amer. Wildlife Conf. 9:288-295. 1945. Diving habits of the shoveller duck. Condor 1947. 47(1 ):39. 1945. (with A. L. Nelson, and W. S. Bourn). 1947. Red fox breeding in salt marsh. J. Mam mal. 26(1): 91-92. 1948. 1945. Speed and endurance of the covote. J. Mammal. 26(1): 94. 1945. (with J. H. Steenis). A progress report on 1948. the marsh and aquatic plant jnohlcm. Reel- foot Lake. .1. Tennessee Acad. Sci. 20(1): 6-19. 1948. 1945. The ruddv turnstone in Ltah. Condor 47(2): 79. 1948. 1945. Feeding habits of the Clark's nutcracker. Condor 47(4): 168. 1948. 1945. Eelgrass conditions along the Atlantic sea- board of North America. Plant Disease 1949. Reporter 29(12) : 302-310. 1945. Some records of birds in LTtah. Condor 47 (4): 172-173. 1949. 1945. California gulls feeding on midges. Condor 47(5):216.' 1945. Research problems on the LI.S. National l')4<». Wildlife Refuges. Trans. North Amer. Wildlife Conf. 10:347-355. 1949. 1945. American eider in Delaware. Auk 62(4): 634. 1945. The whistling swan in Maine. Auk 62(4): 1949. 634-635. 1945. (with H. S. Zim). Bad news for brother rat. Saturdav Evening Post 2I8(19):17. 1949. 113-116. 1946. Abundance of wildlife in Illinois a century ago. Illinois Conservation. Winter 1945-46. p. 41. 1949. 1946. (with Elmer Higginsj. DDT and its effect on fish and wildlife. .1. Econ. Ent. 39(1): 19-1-9. 44-52. 1946. Late nesting of Caspian tern in Utah. 1949. Condor 48(2): 94-95. 1946. (with Clifford Presnall). Partnership in conservation of renewable resources. State 1949. Government 19(6) : 153-157. 162. ,Iune. 1947. Missouri River Basin development: Its 1949. effects on fish and wildlife. Outdoor Amer- ica 12(3). Februaiy. Pi'oc. 36th Conv. Int. Asso. Game, Fish, Cons. Comm. September 1950. 9-11, 1946. pp. 200-206. North Dakota Out- doors. July. pp. 12-14. Wyoming Wild- 1950. life 11 (6): 20-27. June. 1947. Unusual flight of American egret. Migrant (Published by Tennessee Ornithological 1950. Society). June. p. 27. 1947. Zone-tailed hawk feeds on rock squirrel. Condor 59(5) :210. September-October. 1947. Late nests in Yellowstone National Park. '^'^O- Wilson Bull. 59 (3): 172-1 73. September. 1947. Some improvements needed in wildlife re- 1950. search. J. Wildlife Mgmt. 1 1 (4) : 339-347. October. 1950. 1947. Waterfowl at the crossroads. Trans. 12th North Amer. Wildlife Conf. Ivhruary. pp. 67-85. Piesent eelgrass condition and problems on the Atlantic Coast of North America. Trans. 12th North Amer. Wildlife Conf. February, pp. 387-398. Some bird records for southern Nevada. Condor 49(6): 2+4. November-December. Utah swans and geese. Utah Magazine 9(9): 10-13, 25-29. September. Famous geese of Utah. LTtah Magazine 9(10): 12-15. 24-27. October. The puddle ducks of LUah. Utah Magazine 9(10):20-23. October. Unusual consumption of fish bv three species of birds. Wilson Bull. fi0?2):117- 118. June. Our waterfowl problena and program. Proc. 19-1-8 Northeastern Game Conf. pj). 125-131. Aquatic habits of t.lie Norwav rat. J. xMammal. 29(3):299. August. The mourning dove in Alaska. Wilson Bull. 60(3): 188-189. September. Yellow-headed blackbird on Long Island. Auk 65 (4): 605. The trumpeter swan, greatest of American waterfowl, stages a comeback. The Field, London. England. 26 March: 350-351. (with Angus M. Woodbury, and John W. Sugden). Annotated checklist of the birds of Utah. Bull. Univ. Utah 39(16). March. Swinmiing pigeons. Condor 51 ( 3) : 150-1 51. May- June. Does stocking pay? Texas Game and Fish 7(6) :6. 33. May. Wildlife in North Caro- lina 8(5): 118-120. Limiting factors of present waterfowl knowledge. Trans. 14th North Amer. Wild- life Conf. j)p. 42-57. The effects of uncontrolled introductions of plants and animals. Proc. Papers Inter- nal. Tech. Conf. Protect. Nat.. Lake Suc- cess. N.Y. August, pp. 408-413. The eelgrass story. Virginia Wildlife 10 (10): 16-18. Wildlife research pays off. Sports Afield. November, pp. 34-35, 58-60. Conservation in Mississippi. Mississippi Game and Fish 13(4) :4-5. 12. October. 13(5): 10-12. November. Further needs in wildlife research. J. Wildlife Mgmt. 13(4):333-341. October. Conservation in New Zealand and LTtah. The Pioneer, Salt Lake Citj-. Utah. Sep- tember, pp. 5-6. Rehabilitation in the wild. Auk 67(2): 238. April. (with James Silver). The last stand of the key deer. National Parks Magazine 24(102) : 100-103. July-September. Planned enterprises and their effects on the balance of nature. Proc. and Papers of Internal. Tech. Conf. Protect. Nat.. Lake Success. N.Y. August 1949. pp. 323-325. Our federal government and consei-vation. Proc. Utah Acad. Sci. 25:59-67. 1948. The waterfowl regulations. Maryland Con- servationist 27(2) :4-5, 28. Summer. The why of migrator)^ waterfowJ regula- tions. Proc. International Assoc, of Fish & 238 GREAT BASIN NATURALIST Vol. 35, No. 2 Game Commissioners. Texas Game and Fish 9(l):4-7, 31. 1951. (with Warren S. Bourn). Some biological effects of ditching tidewater marshes. Re- search Report 19. January. 1951. (with Herbert Zim). Insects. A guide to familiar American insects. Simon & Schue- ster. New York. 1949. A New Zealand appraisal. The Wood Thrush 5(2). November-December. Au- dubon Society, District of Columbia. 1951. Waterfowl's future depends upon manage- ment. Trans. 16th North Amer. Wildlife Conf. March 5-7. pp. 109-121. 1951. Snow goose record. The Oriole 16(2) :23. June. 1951. Fifty years of progress and handicaps in wildlife management in the U.S. Trans. 41 Con. Int. Asso. Game, Fish and Cons. Commissioners, Rochester. N.Y. Wyoming Wildlife 15(10) :4-ll, 31-34. October. 16 (l):22-29. January 1952. 1952. Present status of migratorj' game birds. Atlantic Nat. 9(3) : 118-123. 1952. Management of our waterfowl. Happy Hunting Ground (magazine of Kentucky fish and game division). Januar3\ 1951. Conservation of our wildlife and other renewable resources. The County Officer. December, pp. 301-307. 1952. Chemical controls vs. wildlife. Ducks Un- limited Quarterly 15 (2): 7. Spring. South Dakota Conservation Digest. May. pp. 14- 15. North Dakota Outdoors. June. pp. 10- 11. Tennessee Conservationist 17 (7):6-7. 18. July Iowa Conseivationist 11(7). July. 1952. (with Warren S. Bourn). Coastal marshes adversely affected by drainage. Presented at North Amer. Wildlife Conf. March 17- 19. Published in Proceedings and in Florida Nat. 25(2). April. 1953. Jamaica Bay sanctuary. Bulletin to the Schools (Arbor and Wild Life Dav issue) 39(7):213-217. March. University of the State of New York. 1953. (with Philip DuMont). What has hap- pened to the key deer? National Parks Magazine, p. 82. April-June. 1953. Wildlife and chemicals. Virginia Wildlife 14(5):10-12. 1953. Conservation of America's faunal re- sources. Proc. Symposium on Cons, of Re newable Nat. Res. of the French-Canadian Assoc, for the Adv. of Sci., Laval Univ.. Quebec, Canada. October 1952. 1953. Report of committee on wildlife conser- vation planning. Proc. Int. Assoc, of Game. Fish & Cons. Comm. pp. 60-65. 1953. (with Geo. A. Rounsefell, and W. Harry Everhart). Fishery science: its methods and applications. .lohn Wiley & Sons. Inc.. New York. 1953. Does upland game beai- its fair share of hunting pressure? Proc. Midwest Assoc, of Fish, Game. Cons. Conmi. pp. 184-206. 1954. Letter to the editor of journal wildlife management. Re Carp removal at Lake Mattemuskeet. A controversv 1«(1):134- 135. 1954. Twin opportunities: conservation and en- gineering. Proc. Amer. Soc. Civil Engineers 80(419). March. 1954. (with David A. Munro). Eelgrass status and enviromnental relations. J. Wildlife Mgmt. 18(4): 449-460. 1954. Bird records for Nevada. Condor 56(4): 223-4. 1955. The Welder Wildlife Foundation. Inter. Assoc. Game. Fish. Cons. Comm. Proc. 45: 133-134. 1955. Progress in wildlife restoration and train- ing. Natl. Wildl. Conf. Pub. 20th Trans. North Amer. Nat. Wildl. Conf. 1955. On the food habits of the diving ducks in Denmark. J. Wildlife Mgmt. 19(4). Oc- tober. 1955. Conservation and engineering partnership. Virginia Wildlife. 1955. The Welder Wildlife Foundation. Inter- national Proceedings. 1956. Little thing may become big. Instructor 91(1):9. 1956. History of waterfowl management in America. Univ. Minnesota. Processed Proc. by Univ. in 1957. 1956. Water and waterfowl. Minnesota Conser- vation Federation. January 21. 1956. Who's doing what in wildlife research. Minnesota, North Carolina State Wildlife Societv. January 20. 1956. The Welder Wildlife Foundation. Atlantic Naturalist. Washington, D.C. 1U4):163- 168. March and April. 1956. Prairie ducks: a study of behavior, ecology, and management. Review. Wilson Bull. 68(l):86-87. 1956. Seeing America's wildlife in oia- national refuges, a review. Ecology. 1956. Chemical controls in relation to wildlife. Virginia Wildlife. March and April. 1956. Marking birds for scientific purposes. Ecol- ogy 37 (4): 675-681. 1956. The problem of wildlife introductions: its success and failures. Proc. Int. Assoc, of Game, Fish, Cons. Comm. 1956. Some wildlife research problems of the southeast. Proc. 1956 Virginia Wildlife, and other states. 1957. Chemical controls in relation to wildlife. Texas Game and Fish Magazine. Con- gressional Record. Appendix pp. A4974- A4976. 1957. The role of piivate agencies in conser- vation. Proc. Int. Assoc, of Game, Fish. & Cons. Comm. 1957. Future i-ole of federal agencies in forest wildlife management. Proc. Soc. Amer. Foresters. Syracuse. N.Y. pp. 97-100. 1057. Late nesting in south Texas. Texas Acad. S(i. (processed). Deceinber H. 1958. (with W. C. Glazencr). Exotics. Texas Game and Fish 16(l):-1'-7. Jaiuiar\ . 1958. Wildlife: its lole in our dynaniii e(oiiom>'. Conference of Southwest Foundations (processed). April. 1958 Waterfowl research: whv research is needed. Naturalist 9(1):28-31. 1958. Wildlife and water conservation. J. Soil and Water Cons. 1-3(2) :65-69. 1958. Science and resources in America. Pro- cessed 23rd No. Amer. Wildl. Conf. Vir- ginia Wildlife. May. Line 1975 TANNER: CL.\RENCE COTTAM 239 1958. Coturnix quail in America. Proc. Int. 1960. Assoc. Game, Fish, Cons. Comm. pp. 111- 119. 1958. A commentary on the fire ant problem. 1960. Proc. Symposium on Fire Ant, Southeast- ern Assoc, of Game and Fish Commis- sioners. Columbia, South Carolina. October 1960. 19-22. 1959. The Welder Wildlife Foundation ap- proach to research and education. Gazette 1961. 8(1): 17-19. 53-54. January. Salt Lake City. Utah. 1959. The whooping crane. Gazette 8(2): 14-20. 1961. February. 1959. (with thadis W. Bo.x). Wildlife in rela- tion to trends in range management. Proc. 1961. Amer. Soc. Range Management. Presented January-. 1959. Knowns and unknowns about l)ird mi- 1961. gration. Naturalist 10(l):2-8. 1959. Chemical pesticides and consei-vation problems. Proc. National Wildlife Federa- 1962. tion. Presented February 27. New York. 1959. (with W^ C. Glazener). Late nesting of 1964. water birds in south Texas. Proc. 24tli North Amer. Wildl. Conf. Presented March 3. New York. 1959. A conservationist's views on the new in- 1965. secticides. Seminar on Biological Problems in Water Pollution, Cincinnati. Ohio. April 21. 1970. 1959. Future research needs in pollution abate- ment and water quality. Seminar on Bio- 1972. logical Problems in Water Pollution. Trans. U.S. Public Health Service. April 24. 1959. Summary of the panel on the effects of pesticides on aquatic life. Seminar on Bio- 1974. logical Problems in Water Pollution, Cin- cinnati, Ohio. April 24. 1975. 1959. Water waste. Water for Texas Confer- ence. September 10. College Station, Texas. 1959. The unconti-olled use of pesticides in the southeast. Southeastern Association of Fish, 1975. Game and Conserv. Commissioners. 13:9- 18. 1959. (with W. C. Glazener. and Gerald W. . Raun). Notes on the food habits of the snakes on the Welder Wildlife Foundation. Texas Game and Fish. June 1960. Pesticides and wildlife. 9th Annual Texas Agricultural-Aviation Conf. February 23. College Station, Texas. Conservation is everybody's business. Pre- sented before State Wildlife Federation, Park Rapids. Minnesota. August 20. Pesticides, chemicals, and pollution. Proc. Nat. Water Pollution Conference, pp. 222- 235. 239-244. Pesticides and wildlife in Canada. Re- sources for Tomorrow Conference, pp. 919- 930. October. Planning for population pressures: a cri- tique. Proc. North Amer. Wildlife Con- ference, pp. 514-524. March 8. Some effects of sprays on crops and live- stock. Soil Conservation Society of Amer- ica, LaFayette, Indiana. August 1. Twinning and other unusual records in gallinule reproduction. Gazette 10(12): 9- 10. Is the Attwater prairie chicken doomed? Audubon Magazine 64(6) : 328-330. (with Eric. G. Bolen. and Burruss Mc- Daniel). Ecological aspects of the black- bellied tree duck. Southwestern Naturalist 9(2): 78-88. (with Milton Caroline). The black-tailed prairie dog in Texas. Texas J. Science 17 (3): 294-302. September. Vasco M. Tanner: a great teacher. Great Basin Nat. 30:200-202. (with Lee Otteni. and Eric G. Bolen). Predator-prey relationships and reproduc- tion in the barn owl in south Texas. Wil- son Bull. 84(4):434-H8. Pesticide pollution. National Parks and Conservation Magazine 43(266): 4-9. (with Eric G. Bolen). Notes on the color phases of the reddish egret {Dichromanas- sa rufescens) in Texas. Southwestern Nat- uralist 20(1): 133-1 34. (witli Eric G. Bolen). Sabine's gull on south Texas coast. Southwestern Natural- ist 20(1):134-135. (with Eric G. Bolen, and James W. Ben- nett). Some ecological effects of lotus on small lake biota in southern Texas. South- western Naturalist. In press. EVOLUTIONARY DIVERGENCE IN CLOSELY RELATED POPULATIONS OF MIMULUS GUTTATUS (SCROPHULARIACEAE) ' Karen W. Hughes- and Robert K. Vickery. Jr.^ Abstract. — The evohitionary divergence of five closely related populations of the 3'ellow monkey flower. Mimulus guttatus. was compared with the length of time the populations could have occu- pied the site and with the ability of the populations to intercross. Populations from the younger sites were found to be morphologically intermediate to i)opulations from the older sites. Ability to intercross was not correlated with morphological divergence at this stage of evolution. Glaciation patterns of the Wasatch Mountains of Utah provide an estimate of the geological age of existing plant pop- ulations. If evolutionary relationships be- tween isolated populations can be deter- mined, an estimate of rates of evolution for these populations may be obtained. The purpose of this investigation is to deter- mine the amomit of divergence between populations from previously glaciated and unglaciated sites and to relate this di- vergence to the age and possible origin of the populations. Divergence is estimated by calculating a Prim network (Prim, 1957), based on morphological charac- teristics of the populations, and by analy- zing the ability of the populations to intercross. The Prim network was calcu- lated for 24 morphological traits and for 6 selected traits to determine w^iether larger numbers of characters make a sig- nificant difference in the network. Methods and Materials Five populations of Mimulus guttatus Fischer ex D.C., the yellow- monkey flower, from two adjacent canyon drain- ages were selected for investigation (Table 1 ) . M. guttatus grows in small isolated populations near streams or springs and is thus ideal for studies of evolutionary divergence. Plants were grown from seeds in the greenhouse. Quantitative data were gathered for 24 morphological traits on an average of 16 plants per population*. We were unable to select a larger number of traits because of the close morphological relationships of the populataions. Traits selected include height, dr}' weight, degree of pigmentation, timing and extent of flowering and seed set, etc. (Crook, 1964) . ^This invesligation was supported in part by a Public Heal I Division of General Medical Sciences, Public Health Service. -Department of Botany. University of Tennessee, Knoxvill ^Department of Biolopy. University of Utali. Salt I-akc Cil Plants from each population were inter- crossed in all comfjinations but one, i.e., diallel design. The estimated age of the populations was determined from the period of time each site was available, geologically, for oc- cupation by the populations. The Thou- sand Springs and Storm Mountain sites have never been glaciated and apparently have been available for occupation for 50,000 years or more, while the remaining three sites were glaciated and have been available only since the retreat of the Wis- consin period glaciers (Table 2). Pop- ulations may not be as old as the site and may have migrated up and down the moinitainsides with changing environmen- tal conditions at the end of the glacial epoch. Table 1. Populations of Mimulus guitalus Culture number Location 6648 Big Water Gulch, Millcreek Canyon. Along stream south of parking lot at end of road. Altitude 7,680 feet. 5840 Thousand Springs, Millcreek Canyon. Hillside adjacent to roacl. Site is marked with a road sign. Altitude 7,200 feet. 6649 Brighton Loop, Big Cotton- wood Canyon on Clayton Peak drainage. East edge of Brigh- ton Loop at top of canvon. Al- titude 8.760 feet. 5839 Spruces, Big Cottonwood Can- yon. Stream and marsh areas of Spruces Campground. Al- titude 7.360 feet. 6127 Storm Mountain, Big Cotton- wood Canyon. Stream in small canyon south of Storm Moun- tain Campground. Altitude 6,240 feet. Service Followsliip 1 No. D-Fl-GM- 3/920. 841 12 from the 240 June 1975 HUGHES, VICKERY: MIMULUS GUTTATUS 241 Table 2. Maximum estimates of time available for occupation of sites by Mimulus guttatus pop- ulations. Site Most recent glaciation Brighton Spi-uces Storm Mountain Big Water Gulch Thousand Springs 9,000 j'ears before present* 11.000 j'ears before present* Not glaciated 11,000 years before present* Not glaciated • Deglaciation occuitchI later lluin ihesc (lales. The 24 morphological traits were ana- lyzed by the principal components me- thod of factor analysis as outlined by Harman (1964) to obtain factors composed of mutually correlated groups of traits. Six major factors were obtained with eigenvalues greater than one (Crook, 1964). The morphological similarities of the populations were determined by calcula- tions of a shortest distance network de- veloped by Prim (1957) and adapted to evolutionary studies by Edwards and Ca- valli-Sforza (1964). In this method the sum of the normalized character differ- ences between the populations is the esti- mate of the difference. Populations with large numbers of differing traits are widely separated on the network and vice versa. The assimiption implicit in the use of the Prim network for evolutionary studies is that the sum of the character differences is jDroportional to the evolu- tionar}' differences between the popula- tions. It should be noted that Edwards and Cavalli-Sforza made assiunptions of independence and selective neutrality not made here. The assumption of neutrality is not necessary, as we are interested in divergence, whether random or the re- sult of selective pressures. The 24 mor- phological traits are clearly not indepen- dent, as they may be combined into 6 groups of mutually correlated traits by factor analysis. Some of the 24 variables are represented in more than one factor (Crook, 1964). The 6 traits are as inde- pendent as can be obtained in that they represent high factor loading on one fac- tor and very low loadings on the other factors. Prim networks were calculated for all 24 characters and for the 6 char- acters obtained from factor analysis. Results and Discussion The 6 major factors derived from the factor analysis of 24 characters can be identified as follows. Factor 1 is a compos- ite factor representing general bushiness of the plant, with high factor loadings on the number of leaves and branches and on measurements related to the age of the plant. Factor 2 represents dry-weight measurements, wdth emphasis on roots and runners. Factor 3 represents almost entirely pigment measurements, with negative loadings on age. Factor 4 is a general measure of height. Factor 5 is a general measure of flower size. Factor 6 has the highest loadings on duration of flowering and seed set. The characteris- tic with the highest factor loading in each factor was selected for analysis by the Prim network (Table 3). The Prim networks based on 24 mor- phological characters and on the 6 se- lected morphological characters are given in Figures 1 and 2. Both networks are identical in pattern, but the distances be- tween populations vary somewhat. •H U PQ o iH P o u sUilus ol' the f;;eiius rrosctunis Baird and Giiaid as piest'iitcd in i)ro\ ions studit and tln-oat osteology and myology are included ii ratios detei'mined. and the position noted. Five dis Uta and Urosaurus. and it is concluded that both of generic status. Vta is considered to be phylogenetic The climbing utas {Urosaurus) and the the ground utas [Uta) are two genera which have had an uncertain sy.stematic relationship. Studies by Mittleman (1942), Savage (1958), Etheridge (1962), and Larsen and Tanner (1975) have used various sets of morphological data to ex- plain their affinities, but myological char- acteristics have been mostly neglected. Other studies involving taxonomy l)y Bal- anger and Tinkle (1973) and I'anner and Jorgensen (1963), ecology and external anatomy by Smith ( 1 946 ) \ Tinkle ( 1 967 ) , Turner et al. (1970), and Tanner (1972), of Uta and Urosaurus and other related genera have been done. However, the va- lidity of the genus Urosaurus and its po- sition in the phylogeny of the Igiianidae has not been completely established. The present study is designed to in^-estigate the anterior osteological and myological anatomy of these genera in order to de- termine their relationship to each other and to show their j)hylogenetic ])osition in the family Iguanidae. The literature dealing with the anatomy of iguanid lizards has been reviewed by Avery aiul Tanner (1971); therefore, we will confine our study primarily to prob- lems relating to phylogeny. Data from previous studies, as indicated abo\e, will be added to our myological findings. |i In the genus Ula l.il.i lidni the head muscle is measured, are noted between different to warrant Ilallowell and its i-elalinnsh s NC> is represented by the genus Phryjjo- sonia. which lacks xiphisternal ribs. The ()tbc>r group, with the utiform sterum bearing xijihisternal ribs, contains the genera CaUisaurus. Holbrookia. Unia. and Uta. Within Uta.. the subgenus Petro- saurus is considered by Savage to be the most primitive, although highly adapted for a rock habitat. The genera Urosaur- us. Sator. and Sceloporus are closely allied ,uid differ from the other sceloporines in ha\ ing a urosaurine type of sternum which possesses xiphisternal ribs. Thus in Savage's studies. I)ased upon the type of sternal arrangements. Uta and Urosaurus are distinctly different genera. Presch (19()9) reports that the osteo- logical characteristics indicate that the horned lizards (Phrynosoma) form a highly s])ecialized genus within the Sce- loporus grou{) of genera (Sceloporus. Sa- lor. Uta, Urosaurus. Uma, CaUisaurus, llolhrookia. and Petrosaurus). Phryno- soma is distinguished from all other mem- bers of \\\o familv in having both a large sternal f()iit and femoral pores. He lists Pe/r(jsaurus as primitive because of its incideratelv sized sternal f()ntan(dle and Sept. 1975 FANOIIELLA. ET AL: LI/.ARn ANATOMY 247 the presence of four sternal ribs. Of the two groups (leriA'ecl from Pctrosaui us. the least altered are Urosaurus, Uta^ Scclop- orus, and Sator, which have hooks on the clavicle and a covering oxer the antero- lateral processes of the frontal in some species, while Holbrookia. Unia. and (\iUi- saurus are the most higlilA ('\nlved. They have lost the lacrimal and postfrontal bones and the first ])air of cervical ribs. Also, the interclavicle is shortened, and the anterolateral processes of the frontal are covered. Thus Uta and Urosaurus are placed in the same group but in distinct- ly different genera. Etheridge (1964) claims tthat osteo- logical comparisons do not {irovide a strong enough argument for or against the recognition of Uta and Urosaurus as separate genera. However, the few osteo- logical differences between sceloporines that do exist suggest that three sul)groups might be recognized: (1) Holbrookia.. Cal- lisaurus, and Uma with two cervical ribs (three in all others); (2) Uta. Urosaurus. Sator, and Sccloporus with cla\icular hooks present (absent in all others); and (3) Petrosaurus with four sternal ribs (two or three in others). Hotton (1955) in his studies of den- tition and food habits has implied that although Uta and Urosaurus are inter- preted as direct but independent descen- dants of sceloporines, the dentition and diet of the utas are similar to Callisaurus. Low^e (1955) studied the problem of gen- eric status of Uta and Urosaurus using ecological relationships. He w^as able to recognize genera on the basis of ecologic divergence alone, without the support of any other character. On this ecological concept Lowe and Norris (1955) based their classification of the assemblage of lizards formerly placed in the genus Uta. They confirmed Mittleman's arrangement of these species because of supporting ecological differ- ences between and similarities within the groups involved. As a result of their studies, they recognized the following taxonomic arrangement: genus Petro- saurus with subgenus Streptosaurus- genus Uta and genus Urosaurus. Petrosaurus and Streptosaurus \^•ere placed together because of their cliff- dwelling habits. Urosaurus was retained as a distinct genus because the species within the group are plant dwellers and clind)ers. lUa was distinguished from the other two genera \)\ \\<. gi-()und dwelling I de-style. MittlemaiTs c lassification of these igua- nids has ]iot been generally accej)ted by Savage and others Ixnause he failed to present lonxincing e\ idence that the sev- eral groups were mor})hologically dif- ferent from one another. The most strik- ing morphological feature listed by Mit- tleman as se]Kirating Uta from Urosaurus was the homogeneous scutellation of the former and the differentiation of the para- \ertebral scales in the latter. We extend our gratitude to those who have helped us in ihe preparation of this pa])er. We are grateful to Dr. Ernest Williams, at the Dei)artment of Herpe- tology, Museum of ComJlarati^■e Zoology at Harvard, for providing us with various ])re])ared skeletons of Uta anrl Urosaurus. We also thank Mr. (diester .1. Bosworth aiid Dr. Dwight G. Smith, who have been so kind as to read and criticize this study, distribute necessary literature, and make suggestions. We are grateful to Kenneth R. Larsen and Wilmer W. Tanner for making available a copy of the manuscript of Larsen and Tanner (1975). Lastly we thank Southern Connecticut State College for financial aiti and the loan of materials and s]:)ace for ])art of this study, and Brig- ham Young LTniAersity for editorial and l)ublication courtesies. Ma'ikrials and Methods Skeletons used in the study were bor- rowed from the Museum of Comj)arative Zoology at Harvard (MCZ), and ])reserved speciinens were borrowed from Southern Connecticut State College (SCSC). One skeleton of Urosaurus ornata wrighti and three of Uta s. stanshuriana were prepared by carefully stripping away the skin, connective fascia, and large mus- (le the first (\i\\ of skeletonizing. After dr^'ing, the remaining tissues were re- moved by stri])])ing and picking until the skeletons were clean. All measurements were taken iji milli- meters with an ocidar micrometer mounted in a chssecting microscope. All measurements were taken from the ex- treme points of the width ntaoe \\as computed be- tween length and width and compared with similar data for both genera. Mea- surements and ratios were taken for iden- tical bones in both genera. Those bones haAing an average mean greater than 40 points are jiresented in Tables 1 and 2. venience of reference the skull has been subdivided into a posterior occipital unit and <\\\ anterior maxillary unit. The skulls are strej)tostylic with a freeh' movable (luadrate bone which ar- T.\Bi.E 1. — Mininuiui. mean, and nuixinnmi measurcnu'nts Uta. Length Name of structure Min. Mean Max. Basisphenoid 1.22-1.35- 1,46 Basioccipital 1.46- 1.75-2.14 Pterygoid 3.95 - 4.58 - 5.12 Ectopterygoid 2.20 - 2.26 - 2.'U1' Vomer 1 .76 - 1 .92 - 2.04 Palatine 1.7! - 1.92 -2.10 Premaxilla 1.12- 1.86- 2. 4-!' Maxilla \^.7^' - 5.08 - V.SO Nasal - 1 .80 - 1 .97 - 2.01- Prefrontal -... 2.39 - 2.62 - 2.88 Lacrimal 487 - .574 - .681' Frontal 3.90 - -k07 - k35 Postorbital 1.07- 1.92-2.62 Jugal 6.10-6.58- 7.22 Parietal 3.24 - 3.86 - 4.10 Postfrontal 487 - .682 - .926 Squamosal 2.30-2.60 2.93 Quadrate 735 ■ .81 i .')75 Supratemp. Fossa 2.78 - 2.91 3.0 I Orbit 4.35-4.41 - 4.50 Nasal Opening 975 - 1.38 - 1.6! Dentarv 6.60-7.08 - 7.80 Articular 3.16- 3.46 - 3.86 Angular Pr 5.70 - 6.89 - 9.50 Surangular 2.20 - 3.17 - 4.1 :• Splenial 2.20-2.42 -2.58 Angular 5.70 - 6.89 - 9.50 Coronoid 2.20 - 3.23 - t.40 Pvriform Becess 2.68 - 3. 1 7 3.42 Parasplienoid Pr 925- 1.2! 1.46 Entu-e Skull 11.2-11.6-11.9 ,kuil uctures of Mi Width Width-length ratio . Mean Max. Min. Mean Max. 1.7i 2.20 1.27 1.61 .780 .975 1.1-6 1.9", .486 .490 .390 3.66 1.66- 1.7') .440- .610 5.15-5.66 .098 - . 1 95 .68') .8 11 1.83- 2.46 1.62- 1.77- .830 1.18 1.67 - 1.00 .658 - 1.41 • .57 3- 1.02 .58() .140 .10-, 1.13 1 .46 2.04 .487 6.35 1.1 1 .720 6.07 .',98 .I'^l 1 .88 1.67 2 17 ~m 7.26- 1.9', 2.78 2.30 1.9', .880 1.22 2.21- 2.04 .975 1 .90 .732 4.6 1- .7 32 5.00 .20 3 1.07 2. 1 1 3. 17 1.42 1.22 .880 7.\-^ ,780 ,585 2.30 2.01- 2.1 1 .487 8.05" .715 .610 .274 .700 .100 .500 .600 .352 .239 .202 .072 .8 12 ,',',0 ,070 ,()',() .10 1 .720 ,388 ,700 331 566 ,/6/ - 77 1 ,701 - .770 . 354 - .445 .704 -.885 .431 - .472 .579 - .687 .733-1.00 .396 -.41 6 .3 32 -.476 .433- .710 .101 -.150 ,003- ,940 ,711 - .925 .092 ,680 288 H) .101- , H)0 . V)0 ,410 .505 ,762 -.820 .770 - .995 .163 -.185 .207 - .243 .824 - .855 .185 -.214 . 1 74 - .226 .280 - .334 .560 - .926 .722 -.765 .424 - .525 .623 - .675 Sept. 1975 Table 2. — Minimum, nu\ saurus. FANGHELLA. ET AL: LI/ARD ANATOMY 249 u and mnxinunn measiurements and ratios for tlie skull structures of Uro- Length Widtli Name of structure Min. Mean Max. Min. Mean Max. Basisphenoid 1 .56 Basioccipital 1 .22 Pterygoid 4.00 Ectoptervgoid 1 .41 Vomer 1 .22 Palatine \ .46 Premaxilla 1,22 Maxilla 5.26 Nasal 2.04 Prefrontal 2.74 Lacrimal 487 Frontal 3.80 Postorbital 2.44 Jugal 5.37 Parietal 3.90 Postfrontal 585 Squamosal 2.44 Quadrate 930 Supratemp. Fossa 2.78 Orbit 3.90 Nasal Opening 1.49 Dentary 6.84 Articular 3.26 Angular Pr 5.70 Surangular 2.68 Splenial 2.20 Angular 5.70 Coronoid 2.68 Pvrifomi Recess 2.44-2.76-3.18 1.76 Parasphenoid Pr 925-1.25-1.42 .440 Entire Skull 11.2-11.6-12.4 6.75 Width-lengtli ratio Min. Mean Max. 1.71 - 1.95 1.57- 1.80 4.74-5.37 1.84-2.24 1.76-2.-M. 1.84-2.20 1.44-1.56 5.84- 6.10 2.32 - 2.78 2.91- .615- 4.15- 2.75- 6.00- 4.08- .791 - 2.55- 1.07- 3.14 4.59- 1.52- 7.15- 3.44 6.30- 7.15 2.86 - 3.26 2.53 - 2.92 6.30-7.15 2.96-3.18 2.76-3.18 1.25- 1.42 11.6- 12.4 3.12 .737 4.40 3.18 6.35 4.30 .975 2.93 1.32 3.36 4.90 1.61 7.60 3.76 1.90 2.68 ■1.17 1.17 .733 .975 1.71 2.20 .830 1.07 .487 4.40 1.71 .540 5.35 .024 .780 2.20 1.32 3.42 .585 .810 .974 5.70 .684 .2-H 1.45 -2.01 - 2.89 -1.32- -1.50 -.811 - - 1 .24 - 2.23 - - 2.21 . - .946 - - 1 22 - .615 - 4.60 1.86- .635 5.64 .190 .940- 2.34 1.54 3.68 .975 .889 1.39 6.77 .794- .478 1.03 1.73 2.08- .597 7.38 2.14 3.17 1.46 1.80 .930 1.71 2.58 2.34 1.07 1.36 .732 4.87 1.95 .732 5.96 .-440 1.07 2.64 1.76 3.90 1.22 .928 1.85 8.10 .880 .585 1.90 1.95 2.44 .925 7.80 .800 .455 .280 .655 .340 .445 .565 .360 .360 .344 .073 .780 .538 .088 .661 .024 .327 .405 .430 .720 .388 .118 .300 .705 .225 .084 .252 .400 .635 .360 .582 .872- 1.00 .555 - .675 .304 -.366 .753 - .880 .488 - .680 .848 - 1 .00 .656 - .730 .405 - .437 .4 10 -.464 .419 - .462 .286- 1.00 .902 - .990 .688 - .800 .106-. 127 .723 - .765 .255 - .500 .370 - .440 .460 - .590 .448 - .525 .806 - .875 .656 - .897 .124- .131 .399 - .507 .875-1.00 .271 - .328 .156 -.244 .284 -.333 .591 -.726 .760 -.910 .474 - .650 .633 -.670 ticulates dorsall}' with the paroccipital process and ventrally with the quadrate process of the pterygoid. Thus, they form a compact, light, and strong cage for the brain and sense organs. The actual shape of the skull is either elongated and flattened dorsoventrally, as in Uta, or shortened and lateraHy com- I)ressed, as in Urosaurus. Measurements of the length of the skull were from the top of the premaxillary bone to the most pos- terior extension of the occipital condyle. Measurements of the width were from the widest extension between the sub- orbital bars in the area of the orbit. The means in Tables 1 and 2 indicate that Uta has a slightly lower skull ratio (.623) than Urosaurus (.633). For con- The occipital jiortion forms a median wall for the attachment of the neck and articulation for the remainder of the skull. It consists of two parts: (a) braincase (basisphenoid, basioccipital, prootic, exoc- cipital, supraocci])ital) (b) foramen magnum (enclosed by the basioccipital, exoccipitals, and supraoccipital). A tripar- tate occipital condyle is located on the pos- terior end of the basioccipital and the lateral exocci])ital in all iguanine genera. Basisphenoid: Length is from the suture between basisj)henoid and basioccijntal, to the beginning of the paras])henoid ])ro- cess (Fig. 1). Wichh is the distance be- tween the widest expansion of the basio- {)tygoid ])rocesses. The lowest ratio mean is in Uta {.767), the highest in Urosaurus (.872). A low ratio indicates that the bone is much longer than it is wide, whereas the higher ratios indicate bones with lengths and widths more nearly equal. Basioccipital: Length is from the suture between the basisphenoid and basioccipital to the posterior tip of the occipital condyle (Fig. 1); and width is between the tips of the lateral extensions of the sphenoccipi- tal tubercles. The ratio in Uta is .701 and Urosaurus .555. Pterygoid: Length (Figs. 1, 2, 3) is be- tween the anterior [portion of the i)terygoid where it sutures with the palatine and the most j)osteri()r tij) of the quadrate process. Width is between the articulation with the basipterygoid process of the basisphen- 250 GREAT BASIN NATURALIST Vol. 35, No. 3 BO — Basioccipital BS — Basisphenoid EC — Ectopterygoid FE — Fenestra e.xonarina FEO — Fenestra e.xochoanal FR— Frontal FVE — Fenestra vorner- onassalis e.xterna JU—Jugal MX— Maxilla NA— Nasal OB -Orbit PAL — Palatine PAR— Parietal PF- -Pineal foramen PM- -Premaxilla POT— Postorbital PP -Parasplienoid process PR Pyriforni recess PRF— Prefrontal PT— Pterygoid PTF— Postfrontal QU— Quadrate STF — Supratemporal fossa SQ — Squamosal VO- Vomer Fig. Dorsal and ventral views of skulls. A ,ind C: l' la. B and 1) / Sept. 1975 FANGHELLA, ET AL: LI/.ARD ANATOMY 251 AR — Articular JU — Jugal CO — Coronoid LA- -Lacrimal DE— Dentary MX— Maxilla EC- -Ectoptorygoid NA — Nasal EP— Epipterygoid OB— Orbit FE-- PVnestra exonarina FM — Premaxilh FR— Frontal PO -Postorbital Fig. 2. Lateral view of skull aiul saurus. ial view of niandib PRE— Prefrontal PR— Parietal PT— Pterygoid PTF— Postfrontal QU— Quadrate SP— Splenial SR Surangular A and C f'!a. and I) Uro- 252 GREAT BASIN NATURALIST Vol. 35, No. 3 CC — Constrictor colli EP — Episternocleidoniastoideus GE — Genioglossus lAP — Intormandibularis anterior profundus IAS — Intermandibularis an- terior superficialis NP — Intermandibularis pos- terior MHI— Mandiliulohyoideiis I OM — Omohyoideus PE— Pectoralis SH — Sternohyoideus ST — Sternothyroidcu: Fig. 3. Ventral view of thioat musculature; superfi Uta; B. Urosaurus. 'pth left aud fi ;pth •igh oid and the suture with the ectopterygoid. The ratio is Urosaurus .304 and Uta .354. Ectopterygoid: Length (Figs. 1, 2) is between the suture with the pterygoid and the suture with the jugal and maxilla. The greatest (Hameter is at its point of union with the jugal and maxilla. The lower ratio is in Uta (.704) and the higher in Urosaurus (.753). Vomers: Length (Fig. 1 ) is from the anterior suture with the premaxilla to the most posterior point of the suture with the palatine. Width is between the median border of the vomer at the ventral mid- line and the most lateral border where it attaches to the maxilla. The ratio in Urosaurus is .488 and in Uta .431. The vomers possess a small blunt projection which jirotrudes from its lateral border into the ojiening of the fenestra exocho- analis and fenestra vomeronasalis externa and divides the opening. This anterolateral projection is seen in both genera. Palatine: Length is from the anterior suture with the \()mer at the midline to the most posterior extension of the suture with the pterygoid (Fig. 1). Width is from the skull's midline to the lateral su- ture between the palatine and the maxilla. The ratio in T^rosaurus is .848 and in Uta .57<». Pn/nan//ae: Length (Figs. 1, 2) is from its anteroventral tip to its dorsal union with the nasal at the dorsal midline. Wi(hh is between th{>- lateral sutures shared b\ the |)reniaxillae with the maxil- Sept. 1975 FANGHELLA. ET AL: LIZARD ANATOMY 253 la on the ventral surface of the jiremaxilla. The ratio in Uta is .7 3^ and in T^rosaurus .656. Maxillac-.l.ewgXh. (Figs. 1. J) is from the most anterior extension of the j)re- maxillarv process to the })osterior-most extension of the maxillae where it sutured with the jugal and ectopterygoid. Width is the vertical distance from the ventral border of the maxillae to the dorsal-most extension at the ]ioint of suture with the nasals and prefrontals. The ratio in Via is 3.96 and in Urosaurus .405. Nasal: Length (Figs. 1, 2) is from the tip of the ventral border as it forms the fenestra exonarina to the posterodor- sal extension that sutures with the jire- frontal. Width is from its medial suture with its opposite member to its most lat- eral extension where it sutured with the maxilla and prefrontals. The ratio in T'ro- saurus is .410 and in Uta .332. Prefrontal: Length (Fig. 1) is from the suture between the ])refrontal and lacrimal bones at the anterior lip of the orbit, to the suture between the prefroiital and frontal. Width is from the suture be- tween the prefrontal and lacrimals to the median point wdiere the frontal, nasal, and prefrontal bones suture together. The ratio in Uta is .433 and in Urosaurus .419. Lacrimal: Length is from the antero- dorsal border as it sutures with the pre- frontal and maxilla to the posterior border on the rim of the orbit as it sutures with the jugal (Fig. 2). Width is the distance between the dorsal border of the lacrimal at the rim of the orbit to its ^'entral border at its suture wdth the maxilla. The ratio in Uta is .101 and in Urosaurus .286. Frontal: Length (Figs.l, 2) is from the most anteromedian suture shared with the parietal. Width is between the most lateral j)osterior projections which suture with the parietal and })ostfrontal. The ratio in Uta is .903 and in Urosaurus .902. Postfrontal: Length (Figs. 1. 2) is the extremities of its longest axis. Width is the distance between the parallel borders on the axis at right angles to the length. The ratio in Urosaurus is .255 and in Uta .288. Jugal: Length (Figs. 1, 2) is between its most anterior projections as it sutures with the lacrimal and maxillae, to the posterior j)rojection which sutures to the anteroventral border of the postorbital. Width is the distance between the two parallel borders at right angles to the length. The ratio in I'ta is .092 and in Urosaurus .106. Parietal: Two measurements (Figs. 1, 2) were taken. The anterior two-thirds of the bone was subjected to length-width measurements, with the length being the distance along the midline, from the an- terior suture with the frontal to the suture between the ]iarietal and the supraoccipi- tal. Width is the distance between the two anterolateral ])rojections that suture with the ]iostorbital and jiostfrontal. The ratio in Urosaurus is .723 and in Ufa .680. Postorbital: Length (Figs. 1, 2) is between the anteroventral and postventral projections. Width is from the ^•entral border to the tij) of the dorsal ]irojection where it sutures with the ])arietal and postfrontal bones. The ratio in Uta is .711 md rosaurus .688. Squamosal: Length ( Fig. 1 ) is between the most anterior and ])osterior extrem- ities. Width is between the parallel bor- ders on an axis at right angles to the length. Ratio in Urosaurus is .370 and in Uta .^25. Quadrate: Length (Fig. 2) is from its dorsal border where it attaches to the squamosal and the \('ntral extremity of the condyle which articulates with the articular. Width is between its medial and lateral borders. The ratio in Urosaurus is .460 and in Uta .^46. Supratemporal fossa: Its length (Fig. 1) is the inside distance on the longest axis and width the inside distance on the long- est axis at right angles to the length. The ratio in ['ta is . 1()7 and in Urosaurus .488. Orbit: Length (Figs. 1, 2) is between the lacrimal and jiostorbital. Width is be- tween jugal and frontal bones .The ratio (most circular o])ening) in Urosaurus is .806 and (most (41i]itical opening) in Uta .762. Feru'stra e ronarina: Length (Fig. 1) is the nitenial distance between the lateral jirojection of the |)remaxilla and maxilla and the sutiu'e between the nasal and maxilla. Width is the inside distance be- tween the lateral border of the premaxilla and the anterior border of the maxilla. The ratio (most circular opening) in Uta is .770 and (most elliptical opening) in Urosaurus .65(). The lower jaw consists of two paired rami united anteriorly in a mental sym- ])h\ sis. Each articidates posteriorly with 254 GREAT BASIN NATURALIST Vol. 35, No. 3 the quadrate. The dentarv of each ramus bears a single row of pleurodont teeth, whereas the remaining bones (articular, surangular, angular, splenial, and coro- noid) are edentate. Dentary. Length is from the anterior tip to the posterior-most projection on the lateral surface of the mandible (Fig. 2). \\'i(hh is the Acrtical distance bewteen the to]) and the bottom of the mandible, immediately in front of the coronoid. The ratio in Uta is .163 and in Urosaurus .124. Articular: Length (Fig. 2) is from it most anterior projection on the median surface where it sutured to the coronoid and splenial to the most posterior tip of the retroarticular process. Width is from the most ventromedial projection of the angular ]:)rocess to the opposite border of the articular where it sutured with the surangular on the lateral surface. The ratio in Urosaurus is .399 and in Uta .207. The angular process of the articular bone differs in shape and size in each genus. This projection was also subjected to length-width measurements. The length is the greatest length of the mandible and was contrasted with the width of the artic- ular, which in part is a result of the size of the angular process. Urosaurus .875 has the greatest ratio (shortest, widest) while the smallest (longest, narrowest) is in Uta .824. Surangular: Length is the longest an- terior-posterior axis on the lateral surface of the mandible (Fig. 2). Width is the longest dorsal-ventral axis in the area of the anterior sutures wdth the dentary and coronoid on the lateral surface. The ratio f shortest, widest) is in Urosaurus .271 and in Uta .185. Spenial: Length is the longest anterior- posterior axis and the greater dorsal-ven- tral axis is the width. The ratio (shortest, widest) is in Uta .174' and in Urosaurus .156. Angular: The angular is roughl}' fusi- form; its length is between the most an- terior and most posterior projections. Width is between the opi)osit(> borders on an axis at right angles to \\\v length. The ratio (shortest, widest) in Urosaurus is .284 and in Uta .280. Coronoid: Length is ftoin the dorsal tip of the bone to tbe tip of the ventral- most projection on the later-al surface (Fig. 2). Width is betw(MMi anterioi- and ])osterior borders where they contact the dorsolateral surface of the mandible. The ratio in Urosaurus is .591 and in Uta .560. Ah()LO(A- To a\()id (onfusion, the terminology used lor the following descriptions of the muscles is that of Robison and Tanner (1962), .Jenkins and Tanner (1968). and Avery and Tanner (1954, 1971). The musculature also follow^s the basic iguanid pattern described h\ the aboAe. Only deviations will be noted in the test. Throat Muscidature M . /u/crniandibularis anterior superfic- ialis is tonstant in both genera examined with the following exception: slightly broader in Urosaurus than Uta (Fig. 3). However, in both genera the muscle is sheetlik(> with the width at least half the length. M. Iut(-rniandil>ularis anterior profun- dus is relatively consistent in its location; however, in Uta it is a wide band of mus- cle attached to the intermandibularis an- terior superficialis. In Urosaurus it is a thin sheet separated anteriorly from the intermandibularis anterior superficialis and posteriorly from the intermandibularis posterior by a thin membrane (Fig. 3). M. Intermandibularis posterior is con- tinuous posteriorly with the constrictor colli from wlii( h it can be delineated by a natural sej)aration of the muscle fiber bundles (Figs. 3, 9 and 10). The posses- sion of this separation is varied in the genera examined. In Urosaurus the con- strictor colli and intermandibularis j)os- lerior are (■l()sol^■ associated along their entire (onnnon border. In Uta the two muscles are separated totally laterally but are continuous for a short distance near the midlin(> raphe. M. Mandihulohyoideus I in Uros(nirus has ap[)r()\iinatcly one-half of its body covered l)\ the omohyoideus. whereas in the Uta ()nl\ a small posterior ])ortion is covered i b'ig. > ) . M . Ma/idil>ul(jlnoideus II ^vas described l)V AM'r\- and Tanner (1971) for other ignanids. I l(t\\('\ cf. we were iniable to lo- in (Mther Ufa or Urosaur- us^. M. Mfnidihuinhyoideus III in both Uta and Urosaurus arises from the ventre- Sept. 1975 FANGHELLA. ET AE: EIZARD ANATO!\IY 255 medial surfaces of the dentary and angu- lar between the anterior and posterior mvohoid foramina (Fig. 4). Tiio narrow insertion is on the lateral surfaic of the ceratohyal distal to its midpoint. M. Genioglossus is a thick bandlike muscle in both generji which occupies a large area between the mandibular rami (Figs. 3 and 4). Its position is ventral to the tongue and anterior to the basihyal. The first, second, and third mandibulo- hyoideus and the intermandibnlari^ muscles are all dorsal to it. M. Hyoglossus is as described in ollu>r iguanids. M. Branchiohyoideus in Uta (as in Sau- romalus) has a narrow insertion on the first ceratobranchial, whereas in Urosaur- us the insertion covers over half the (hstal portion of the first ceratobranchial (Fig. 4 ) . M. stcrnohyoidcus. as reported in the literature, is subject to considerable con- fusion concerning its limits (Figs. 3, 4, 6). Davis (1934:19) considers the super- ficial layer to be divisible into three parts in (^rotaphytus. One of these muscles he calls the omohyoideus. Robison and Tan- ner (1962:6) consider this muscle con- tinuous in the same genus. Oelrich (1956:51-52) treats this muscle in Cteno- saura as being continuous, but owing to the different origin and direction of the fibers he separates the layers into omo- hyoideus and sternohyoideus. Kesteven (1944:245-246) studied the agamid, Phy- signathus, suggesting a separation in young sj)ecimens and treats these layers as con- BH — Branchiohyoideus GE — Genioglossus Fig. 4. Ventral view of throat miisculaturr A. Uta; B. Urosaurus. 256 (,KI.\r BAMN NAirUAl. Vol. 35, No. 3 sisting of three patis which he considers to represent the similar, though distinct, divisions present in \'ar(inus. In the igua- nines Avery and Tanner (U)71i treated the sternohyoid(His complex as three sej)- arate muscles, sternohvoideus, sterno- throideus, and omohyoideus. This arrange- ment is followed here. In both genera ex- amined, the sternohyoid ens forms a broad elongated sheet of muscle covering the posterior portion of the mandibulohyoideus I muscle. \1. omohyoideus is sheetlike and forms the lateral extension of the sternohvoideus complex (Figs. 3, 5, 10). In both genera it originates medially from the lateral tip of the transverse ]irocess of the interclavi- cle with some fibers of the episternocleido- niastodeus. Laterallv, the omohyoideus originates from the anterolateral surface of the cla\ i( Ic and anterior border of the su])rascapnla. Its fibers pass obliquely an- terior to insert on the posterior margin of the first ceratobraiK hial and the ])roximal end of the second c cratohranchial cartil- ages. In both geiiera the niecHan border is separated from the lateral border of the sternohvoideus. 7'he delineation of both muscles must b(^ made l)y comparing the origins and insertions. In U rosaurus it is easil^• se])arated, as the fibers of this mus- c ie pass oblicjue to those of the sterno- hvoideus covering most of the mandibulo- hyoideus I. In Uta it is a thin band just lateral to the sternohvoideus. M. Stcrnotliyroidcus is the most medial extension of the sternoh}-oideus complex Ix- 7*— , C;L Claviclp IC- -Interclavicle LX — Larynx Fig. 5. Ventral view A. I'to; B. IJrosaurus. oM {)nniii\.ii(i(Mis rr. I'M I'h.nvn-..,,! uu-mUam- PT Pl(MVK()i(lin,in.lihul,ins •i\\ iiius< iihitin-o; foiirlh en the basihvoid and hypohyal. In Urosaurus the lateral bordc>r of the sternothyroideus and the mechau l)ord(M- of the sternohyoideus are difficidt to de- termine. In Uta their sei)aratiou is chs- tinguishal)le since the three muscles have fibers obliciue to one another as described scured b\ the more superfic for Saurornnlus by A^■er^' and Taimer tor colli. (1971). M. Levator scapulae superficiaHs is not is. \. t. 9). it IS s| of the later-al siir- / /<)^(/urus. A nari"o\N us HI / '/a. h'.pistcriiix lctil(tin(i.\t third bundle ( cervicomandibularis) in I Id and Urosaurus is com])leteh' ob- onstric- AM—Adductor ir.andib e.\tei-nus niedius CC — Constrictor colli DM— Depressor iiian(lii)ul, LS — Levator scapulae siipoi-ficialis PS- -I^seudoteiiiporali superficiaHs TR Trapezius Fig. (3. Dorsal view of head and neck musculature; supeifu ial dejitli at left and fi at right. A. Via; 15. Urosaurus. ie|)tli 258 (JREAT BASIN N ATT HAL. L^ Vol. 35, No. 3 as fan shaped in I'tci as in I'rosdurus (Figs. 6, 7, 10 ,11, and 12). M. Levator scapular profundus has a more superficial j)osition in ltd than in Vrosaurus (Figs. 7, 12). Temporal Mnsc ulature .1/. Ptery^oniandihularis does not de- viate from the txpical ionatiid pattern (Figs. 4, 5). .1/. Levator angularis oris thffers m size in Urosaurus where it covers over half the intratemporal fossa (Fig. 9). In Uta it is narrower, covering apjiroximately one- third of the fossa. .1/. Adductor mandihularis externus su- perficialis is similar to that of other igua- nids (Figs. 9, 10). M. Adductor itiaiidihuhiris externus niedius is also with the t\j)ical iguanid pattern (Figs. 6, 9, 10, 11)." U. Adductor mandihularis etternus projundus is as in otht^r iguanids (Fig. 12). M. Pseudotemporalis superficial is does not deviate from other iguanids (Figs. 6, 12). ^L Pseud()t('nip()r(dis profundus is more ohscured by the levator pterygoideus in Urosaurus than in Uta (Fig. 13). 1/. Adductor mandihularis posterior shows some \ ariations in the two genera, particularly in the location of the muscle with reference to the auditory meatus ( Fig. 13). In Urosaurus it is located both Aentral and anterior to the meatus, whereas in Uta the nuiscle is found slishtlv ventral to the meatus. F,P Episternoclcidoniiistnicleus I.S Icn I.P Levator scapulao inofiindus supcii Fig. 7. Dorsal view of liead iiml right. A. IJla; B. Urosaurus. Sept. 1975 FANGHELLA. KT AL: Ll/ARD ANATOMY 259 AI. Levator ptcrygoidrus is as seen in other iguanids (Fig. 14"). M. Protractor ptcrygoidrus lias a nuu h larger insertion on the medial (rest of the {[uadrate in Urosaurus than in Uta (Figs. 13, 14). DiSClTSSlON A study of the anterior osteology and myology of Uta and Urosaurus reveals some distinct anatomical differences be- tween the two genera. Mittleman (1942) consider(Hl the rela- tionship between Uta, Urosaurus, and the iguanines (Sauromalus, Dipsosaurus, and Ctenorsaura) . He regarded Uta and Z7ro- saurus as distinct genera. Savage (1958) outlined the iguanine characteristics and included Crotaphytus in that evolutionary line. He also determined some structural differences between Uta and Urosaurus. The differences cited by Savage include Urosaurus possessing a ])ectf)ral girdle of the urosanrine l\pe: lateral xiphisternal ribs present and no supranasal scales. Uta is (lislhutK different in having a pectoral girdle of the utiform type, no lateral xiphisternal ribs, and supranasal scales separating nasals from internasals. Avery and Tanner (1964) ])resent several myological differences between Sauromal- us an(l Crotaphytus and indicate these two genera are not in the same evolutionary line, indicating that at least two major subdivisions exist in the family Iguanidae. Etheridge, in 19(54, also examined the iguanines and se])arated Crotaphytus from them based on osteological differences. He states that osteological comparison sug- gests that three subgroups of sceloporines may exist: (1) Holbrookia. Callisaurus, and Urna possessing the scapular fenestra; (2) Uta, Urosaurus, Sator, and Sceloporus demonstrating the absence of the scapular IE— Iiiteicostalis e.xtciiii SD- Sf Fig. 8. Dorsal view of head rip;ht. A. Uta; B. I'msaurus. itus (dorsal pat SP- Spinus dorsi iicrU musculature; fourth dei)t.l left and fifth depth at 260 GREAT BASIN NATURALIST Vol. 35, No. 3 ! fenestra; and (3) Pctrosaurus possessing very few osteological comparisons to the other sceloporines. Presch (1969) reported that the osteo- logical chararteristics indicate that the horned lizards (Phrynosoma) form a highly specialized genus within the sce- lojiorine group of genera. Phrynosoma is distinguished from all other members in having a large sternal fontanelle. Petro- saurus. he states, is clearly primitive with its moderately sized sternal fontanelle and four sternal ribs. Of the two groups de- rived from Petrosdurus. the least altered are Unjsaurus. Via. Sccloporus, and Sator. with hooks on the clavicle and a covering o\er the anterolateral ])rocesses of the frontal. T'hus Presch places Ufa and Uro- saurus in the same grou]i but as distinct genera. Zug (1971 ) studied arterial patterns in many iguanids and found differences be- tween Ufa and Urosaurus. In his Figures 10 and 15 he illustrates these differences. Particularly significant is the representa- tion of separate phylogenetic lines for these genera. Recentl} Purdue and Carpenter (1972) AM — Adductor rnan(lit)ulaiis extornus medius AS — Adductor mandil)ularis e.xtemus superficialis AU — Auditory meatus Fig. 9. Lateral view of liciid (',(] (Joiistrii tor coll i DM Dciiicssor ma Klil.ulai KI^ Epistciiioc li'ido iiatdidi" IP Intrnniiiulilnila IS posterior 1 aiirl neck niuscul ■iture; I..\ Levator annularis us I'H I'lapezius superficial depth. A. I'ta; B. Urosaur Sept. 1975 rAN(;Hi-.i.i.,\. i:t al: lizard anatomy 261 have studied the relation- orus. Uta. and / 'rosauru^ h\ tlieir dis])la\- motions i- \ sn,u,u(>st that (HstiiKt genera •i\('d fi-oni one group of Scrloporus, wliile L'/a is nujre closely related to Pctrosaurus. Larsen and Tanner (1975) have })re- sented a new j^hylotreny for the scelop- orines hased on external characteristics in- cluding the development of specialized scales and structure of the gular fold, and ( ters ni( hi(H 1114 hip anc Thev nid Hat • th.il Seel op 1 and leri\ed h'oni one hnc intei-nal (hi shoulder rati 0/7/V is adxan of s( ("lopofiiuvs. iiK Jnd I Id. and /\ir(}\auf i/s and the s,,nd ii/.anjs. Iisdurus. I lolhroohid. comprise a separate Tliey separate the primitive species of Sccloporus from that genus and resurrect Cope's genus Lysoptychus for them. They also consider lUd and Urosaurus to he closely related, with Ufa being the more primitive. ig Sa/of\ Urosaurus, uhile Phrynosoina irluding ( ' iiid. (d/- and Cophosdurus. lin(> of evolution. AM- -Adductor iiianililuilaris e.xtenius inedius AS — Adductor mandihularis e.xternus superficial is DM- Depressor niandilDularis EP- Episternocleidomastoideus IP — Interinandiliularis jiosterior LS- Levatoi superficia OM— Oniohvoideus SH Sternoliyoideu pulae Fig. 10. Lateral view of head let k musculature ^t depth. A. I'ta: B. rrosaurus. 262 GREAT BASIN NATHHAUST Vol. 35, No. 3 Ostcologv As stated eai'lier. length-width measure- ments of hones and bone shaj)Os were utilized to analyze the osteological rela- tionships between the two genera. The ratio means in Tables 1 and 2 were used to make these relationships clear. Utilizing the method of Avery and Tanner (1971), one can assume that a chfference of .40 or fewer j)ercentage points (.20-. 60) between means of the same bone indicates a close relationship. The possession of bones with similar shape is also an indicator of close relationship. Based on comparisons of the skulls of iguanine lizards, Avery and Tanner (1971) indicate osteological characters of the skulls of iguanid lizards to be stable within generic limits. This osteological sta- bility is also demonstrated by the skulls of Uta and Urosaurus. Re^ iewing Tables 1 and 2, the 35 char- acteristics and corresj)onding mean ratios indicate 24 structures with mean ratios differing bv .40 or more percentage points (.2()-.65). ■ As indicated by Etheridge (1964), the difference between the skull length and AP — Adductor niiindihul; externus profundus LP — Levator scapulae profundus LS Levator s: suprrfuialis Fig. 11. Lateral view of head ipulae dati PS - Pseudotoniporalis supcrfirialis d df|)tli. A. ltd: B. rrosaunis. Sept. 1975 FANGHKLLA. ET AL: LIZARD ANATOMY 263 width ratio in Utd and Urosaurus is al- most negligible. I fowever, oiu- measure- ments of the basisphenoid and basioccipi- tal bones show differences of 1.05 and 1.46 points respectively, along with the more posterior location of the suture be- tween the two structures. Considering the \entral bone structures (Fig. 1). the follwoing mathematical dif- ferences exist. Both the jiterygoid and octo- pterygoid bones differ by more than .40 j)oints (Tables 1 and 2); however, the ectopterygoid possessed by Urosaurus shows anterior wings extending to the maxilla, not seen in Uta. The primary differences found between the vomer and pahitine bones in both genera is mainly the j)ositioning of their common suture. In Urosaurus the suture extends antero- medially from the inferior orbital foramen to the fenestra exochioanalis, whereas in Uta the suture is found extending later- ally from the anterior ])ortion of the pyri- form recess to the maxilla. In reference to the nasal capsule (nasal, prefrontal, lacrimal, and se{)tomaxilla), there are found ratio differences in the nasal and lacrimal (slight difference in the prefroiital) and practically no struc- tural peculiarities. The premaxilla and AM — Adductor maiidilmlaris externus medius EP — Episternocleidomastoideus LS — Levator scapulae superficialis Fig. 12. Lateral view of head and neck niuscula PM— Pharyngeal membrane thinl depth. A. Uta; B. Lrosaurus. 264 (iUKAT BASIN NATURALIST Vol. 35, No. 3 maxilla of the palate coni])lex show a discrepancy only in the premaxilla. The premaxillary bon(>s of the two genera dif- fer by .n points \vith the premaxilla of Uta being narrower anteriorly than in JJrosaurus. The parietal, scpiamosal, and (juadrate (of the temporal fenestra portion of the maxillary segment) all differ in size. shape, and ratio. The parietal in Uta is much more rectangular and broad than in Urosaurus and co^■ers more of the pos- terior portion of the braincase. The differ- ence between the squamosal and quadrate in both genera is mostly in ratio (shape- size) rather than in location. On the dorsal area of tlu> skidl a striking difference is noticed in the size and shape of the fenestra exonarina. In Uta the fe- nestra is oval and quite similar to that of SauroT?iaIus o. multiforaminatus (Avery and Tanner, 1964), whereas in Urosaurus the anterior portion of the fenestra is ex- panded anterolaterally. Turning to the mandible, we see several differences. The articular differs by 1.92 points in I'ta and is only half the length seen in Urosaurus. The largest mandib- AM- -Adductor mandihulnris posterior PP- -Protractor pterygoideus FT Pseudotomporali^ profundus SI) — Spinus dorsi SS--.Su[)iasrapuli Fig. 13. Lateral doi)th of hoad and neck inus< uhiturc; fourth ptii A, t'la; B. f'rosnurus. Sept. 1975 lANGHELLA. ET AL: LIZARD ANATOMY 265 ular difference is in the size and location of the surangular, wliere th(^ ratio (hf- fcrential is .86 jioints and lh<> [)osition in Urosaurus is directly Aciilral lo Uic (oro- noid. In Uta, however, it is [i()st(>r()\(Mitral to th(^ coronoid with onlx an anterior \^in^• making conlat I \Nith the (oronoid anteri()rl\'. This arrani^enicnt of the snr- angnhu- and coronoid I'onnd in Uta is siniihir to Affihlyr/iy/u /lus ( risfatus\ Bra- chylophus faciatus. (lialarodon niadagas- (■(ircTisis. Conolophus pallidus. Clrnosaura pcctinatd. Cyclura nuicclryi. Dipsosdurus (lorsalis. Iguana igiKnia. Opiurus scbac. and Saurornalus ohcsus ( A^ ('r\' and Tan- ner, 1971 ). None Urosaurus. lese are sinnlar to :\lvol..-v The anicr-ior anatniny in I'la wIkmi (()in|)ai-ed with dial in Urosaurus sliows ohserx.ihie ni\ nloLjic ns. and M. slernothroidens, one can see that all three mnscles were distinctly separated in Uta as they were reported for the iguanids Cyclura uuchalis and Sauro- nudus ohcsus bv Aver\' and Tanner ( 1971 ). where. resembles flight I \ that of Brachylophus fasriafus. J' Ia-vh 'P Proti tor ptervgoideus SD- Spiiius (lorsi SK--Sonatus iciorsal part) Fig. 14. Latei-al \ unv of iicad atid iiort; muscula liftli (Irnt.li. A. rta; 266 GREAT BASIN NATLTRALIST Vol. 35, No. 3 The dorsal nius( ulaturo at the first depth shows a distinct difference between IJta and Urosaurus. The M. le\ator scap- ulae superficial is. which is evident in all species studied h}- Avery and Tanner (1971) as well as in Uta. wvis overlaid by the trapezius complex in Urosaurus. Another distinction of the ventral muscu- lature is that seen in the M. episterno- cleidomastoideus. In Uta, as in the genera studied by Avery and Tanner, the M. epi- sternocleidomastoideus and the levator scapulae profundus are found anterior to the A I. levator scapulae superficialis, whereas in Urosaurus the jwsition of these two muscles is posterior to M. levator scap- ulae superficialis. Lateral musculature shows some simi- larities between Urosaurus, Chalarodon madagascarensis , and Oplurus schac, while the similarity of Uta to Sauromalus obesus and Cyclura nuchalis (Avery and Tanner 1971) is noticeable. In Urosaurus the M. episternocleidomastoideus is overlaid by the trapezius, whereas in Lha it is not covered. The M. episternocleidomastoideus and M. levator scajndae superficialis of Urosaurus are buried beneath the second depth of muscle tissue, while in Uta these muscles are mostly superficial. The orientation of the M. levator scap- ulae superficialis and M. le^'ator scapulae ])rofundus at the third depth is the same in Urosaurus and Chalarodon rnadagascar- ensis, while these muscles in Uta resemble Sauromalus obesus and Cyclura nuchalis. In Urosaurus the M. le^■ator scapulae su- perficialis covers the M. levator scapulae profundus ])osteriorly as seen in Chalar- odon rrmdagascarcnsis (Avery and Tanner, 1971 ) . In Sauromalus obesus the common border of the M. levator sca]:)ulae pro- fundus and M. levator sca])ulae super- ficialis is similar to that in Uta. The temporal musculature of Cyclura nuchalis (Avery and Tanner, 1971), M. protractor ptervgoideus, M. ])seudotem- poralis ])rofundus, and M. levator ptery- goidous appears to be similar to that of Uta. whereas in Urosaurus: these muscles are seemingly very similai- to iliose of Chalarodon. based on internal and external characters. We agree with this analysis of Uta's re- lationshi}) to Urosaurus. We believe that Uta is more })rimitive for the following reasons: 1. The ectopterygoid of Uta is simple ill structure and shape, while that of Urosaurus is greatly expanded and more complex in shape. 2. The fenestra exon- arina of Uta is similar to that of Sauro- malus. a primitive iguanine lizard. 3. The structure of the surangular and coro- noid bones in Uta is similar to that seen in the more primitive iguanines and the Madagascar iquanids. 4. The omohyoi- deus muscle complex in Uta may be sub- divided into three distinct muscle bun- dles as seen in the primitive Cyclura and Sauromalus, wheras Urosaurus resembles the omohyoideus configuration seen in the specialized Brachylophus. 5. In Uta the levator scapulae superficialis, levator scap- ulae profundus, and the episternocleido- mastoideus muscles have a configuration similar to the more primitive iguanines. In Urosaurus the muscle pattern shows considerable deviation. 6. In Uta the leva- tor scapulae superficialis is superficial while it is overlain by the tra])ezius com- plex in Urosaurus, indicating to us a higher degree of sjiecialization. The similarity of these two genera to the Madagascar iguanids Chalarodon and Oplurus remains a confused question. If ^ Oplurus is ancestral to the iguanine lizards j as indicated by Avery and Tanner (1971), is Clialarodon also ancestral to the scelop- orine lizards? Ap]:)arently not since both Uta and Urosaurus share some character- istics with Chalarodon and Oplurus with- out establishing a consistent pattern of relationshij). These similarities are more likely the result of parallelism than a close phylogenetic relationship. All four genera are desert animals and Chalarodon sui)erficially resembles both Uta and Uro- saurus externally. Perhaps future comparisons should be inade between Phrynosoma and Petro- saurus and the Madagascar iguanids to determine the phylogenies of the main lines of iguanid evolution and the Mad- agascar iguanids. Phylog(Mieti( Helationshijis Larson and Tainier ( 1975 ) consider Ita to be more primitive than Urosaurus and both more primiti\e than Sceloporus. Conclusions and vSummary Data (leri\(Ml From the preceding ob- servations strongly indic-ate that Uta and Urosaurus are (listinct genera. These con- Sept. 197-5 FANGIIELLA, ET AL: I,T/AH1) Ai\ATf):\IY 267 elusions are based on the distinct osteo- logical and nivological ( liafacteristics found in the head and thioat anatomy. Osteological differences are simunarized as follows: 1. Bones showing major dif- ferences in size and shajie are the basi- sphenoid. basioccipital, pterygoid, ecto- pterygoid. premaxiUa, articular, and sur- angular. 2. The ectoptervgoid possesses an extended anterior wing to the maxilla in Urosaurus which is not present in Uto. 3. The parietal in Uta is much more rt>c- tangidar and broader than in Urosaurus. 4. The fenestra exonarina in Uta is o^al, whereas in Urosdurus the anterior portion is expanded anterolaterallw 5. The posi- tion of the surangular in Urosaurus is di- rectly ventral to the coronoid, but in Uta it is ])osteroventral to the coronoid with only an anterior wing making a narrow contact with the coronoid. Several noticeable generic my' Herr A. Forrer in Mexico. Ann. Mag. Nat. Hist., ser. 5, 11:342-344. ' . 1885. Catalogue of the lizards in the British Museum ( Natui'al History). London, pi'inted bv order of the trustees. 2. xiii plus 4')7 pp.. 23 pis. Camp. C. L. 1921 Classification of the lizards. Bull. Amer. Mus. Nat. Hist. 48(11): 289-481. Cope, E. D. 1865. Descriptions of new Ameri- can sfiuamata in the museum of the Smith- sonian Institution. Proc. Acad. Nat. Sci. Phila. 1863:100-106. . 1864. Contributions to the herpetology of tropical America. Proi . Acad. Nat. Hist. Sci. Phila. 16: 166-181. . 1887. Catalogue of hatrachians and rep- tiles of Central America and Mexico. Bull. T:.S. Nat. Mus. 52: 1-98. Davis, D. D. 1954. The collared lizard, a labora- tory guide. Macmillan. N.Y. 57 p. DuMERii., A. 1856. Description des reptiles nouveaux ou imparfaitement connus de la collection du Museum d'Histoire Naturelle et i-emarques sur la classification et les carec- tcres des reptiles. Arch. Mus. Hist. Nat. Paris. 8: 459-588, pis. 17-24. Ethkrhjoe, R. 1962. Skeletal variation in the iguanid lizard Salor ^raiuhicrus. Copeia 1962 (4): 615-614. . 1964. The skeletal morphologv and svstematic relationships of S(eloi>orine lizards. Copeia 1964(4): 610-651. . 1965. The abdominal skeleton of liz- ards in the famih Iguanidae. Herpetologica 21(3): 161-l()7. 268 GREAT BASIN NATURALIST Vol. 35, No. 3 1967. Lizard (Inl Copei 1067(41: 699-721. FisciiKii. .1. G. 1882. Heipetologisch(> i)omci- kuiigiMi vorzugsvveise ubei" stucko dci- samni- liing (les Naturhistorirlion Musoiini in Bremen. Aliliand. Natiuwiss. Ver. BieiiuMi 7. 225-238. pis. 14-17. H.\i, LOWELL, E. 18)4. Desciii)tioti of new re]v tiles from California. Pnu . Acad. Nat. Sri. Phila. 7: 91-97. HoTTON. N. \9'^j. A survey of adaptive rela- tionships of dentition to diet in the North American Iguanidae. Anier. Midi. Nat. 5 3 (1): 88-114. Iknkins. R. L.. and W. W. Tannf.h. 196S. Os- teology and myology of Phrynosotna p. platyrhinus Girard and Phrynosotna p. her- nandisi Girard. Brigham Young Univ. Sci. Bull.. Bio. Ser.. 9(4): 1-34. Kesteven, H. L. \9-VV. The evolution of the skull and the replialic muscles. A compar- ative study of tlieir development and adult morf)holog\'. Memoir .Australia Mns. ,S\dne\- 8(3): 237-264. I.ARSEN. K. R.. AND W. W. Tanner. 1975. Evo lution of the sceloporine lizards (Iguanidae). Great Basin Nat. 35: 1-20. Lowe. K. H.. and K. S. Norrl'-. 1955. Analysis of tlie herpetofauna of Baja California. Mex- ico. IIL Herpetologira. 11: 89-96. I.UNDELius, E. L. I'liT. ,Ske!(^t,d adapt, ilions \n two species of Str/d/ioi us. I'Aolutiou II: 65-83. MiTTLEM.\N. M. B. 19-1-2. .\ sumniarv of the iguanid genus I rosaurus. Bull. Mus. Comj). Zool. Han-ard 91(2): 106-181. NoRRis, K. S.. AND C. H. Lowe. 1951. A stu. Handbook of lizards of the United St.ites and Canada. Comstock Puhl. Co. Ithaca. NY. 557 p. S.MiTii. H. M.. AND K. H. TvYLoR. 1950. An annotated (lieiklist and kev to the reptiles of Mexico exclusive of the snakes. Bull. U.S. Natl. Mus. 199: 1-253. Stebbins, R. C. 1954. Amphibians and reptiles of Western Nortii America. McGraw-Hill Book Co. 5 36 p. Strjneger, L.. AND T. Barbour. 1943. A check- list of Nortli .\meiiian amphibians and rep- tiles. 5th ed. Harvard I'mv. Press, Cambridge. Mass. 260 I). Van Denbur(;h. .1, l'»J2, The rejjtiles of west- cnx Nortii Amen, a. 1. Lizar.ls Ore. Pap. Calif. .\(ad. S, 1. 10: 1-611. pK. 1 57. /i (,. (t. H. I'»71. Tiio (hstnhiitioii and patterns of till' major arterie;, of the iguanids and comments on the intergeneric relationships of iguanids (Reptilia: Lacertilia). Smith- sonian Contributions to Zoology. 83: 1-23. DISTRIBUTION AND ABUNDANCE OF 1111*: BLACK-BILLED MAGPIE [PICA PICA) IN NORTH AMERICA Carl E. Bock^ and I. W. I, Abstract. — Analysis of Auduhnu Society Christmas bird mi variables shows the degree to whidi the Bla(k-l)illed Magpie is America. The abundance and distiil)ution of this species apiicn barriers: increasing sunnner temperatures in the Southwest ami precipitation on the central plains. tid certain environmental arid regions in North d b, sumnie iditv and The Black-billed Magpie ( Pica pica ' is one of the most conspicuous ])asscri]io birds on the western plains and in the Great Basin. In this paper we are con- cerned ^^'ith what factors determine the abundance ]iattern and southern and east- ern limits of distribution in this species. Linsdale (iji Bent 1946:134) noted that ■"a rather striking relation to climate ex- hibited by this bird has not been clearly explained" or, specifically, that Pica pica in the New World seems restricted to the "cold type steppe dry climate" region characteristic of that portion of the ITiited States north of approximately 35'^ latitude, west of about 100° longitude, and east of the Sierra Nevada-Cascade Range. With- in this area Pica pica is widely distributed, breeding at most elevations u]) to 10,500 ft. (Ligon, 1961). The one universal characteristic of mag- pie habitat is an association of thickets or riparian areas, necessary for breeding anrl roosting, with open meadows, grassland, or sagebrush fields suitable for its method of foraging (Linsdale, 1937; (labrielson and .lewett. 1940; Grinnell and Miller, 1944; Jones, I960; Erpino, 1968). With- out doubt these habitat requirements ex- plain the absence of magpies from the closed boreal forests in the north. How- ever, this sort of habitat mosaic occurs along watercourses in the Great Plains and Southwest, so that it is not immediately apparent why this species does not occup\ a larger part of the Lhiited States. We have analyzed the winter abun- dance ])attern of Pica pica using data from the annual Audubon Society Christmas bird counts. Although magpies may wan- der somewhat in winter (Jewett et al., 1953), the bulk of the population seems to stay within the breeding range. Christ- mas count data were compared with cer- tain climatic variables taken from maps of 50-\c>ar ( liniati( iiicaiis published \)\ the U.S. be|)arlnicMil of Agric uhur(> (1911). Results pi-()\ ide insight nito those' eii- \'iroinneiital tactors which directly or in- direct]\' influence the abundance and dis- tribution of the Black-billed Magpie. Mi'.TilODS Each (JhrisOnas (ount is a standardized one-day census conikicted inside a pre- scribecl circle 15 miles in diameter. Hun- dreds of such counts are made aimually in North America. Data gathered include the nuniher of each sj)ecies seen and the nund)CH' of ■■|)arty-hours" of fieldwork as a measure^ of c ciisus effort. Published re- ^idts of the cotints provide an index to c ontinent-\\i(h' patterns of bird distribu- tion and abundance (Bock and Lepthien, 1974; Bystrak, 1974). Details of techniques for computerized data storage, retrieval, and analysis have been described previously (Bock and Lep- thien, 1974). In this case we retrieved data from the 1969-70, 1970-71, and 1971 -72 Christmas coiuits (2,743 individual censuses), sorle*! thi-se In- blocks of five degrees of latitude and longitude, and computed mean nundier of birds ])er party-hour lor all Kiunts within each l)lock I Eig. I ). Similar maps of climatic data ucrc^ constructed for all latitude- longitude blocks at least partiall}^ \^dthin thc^ L^nited States, adapted from the maps in the U.S. Department of Agriculture (1941). I'he statistics |)rogram B:\ID-02R (Dix- on. 1971 1 was used to compute stepwise regression of magpie^ mimbers against the series of ( limatic \ariables mapped. Rl'.Sl'LTS Eigiu'e 1 shows the winter abiuidance [lattern of the Black-billed Magj)ie. Christ- 'Departmcnt of Etivironmental, Population, and Organisniic Biology, Univcrsily of (Jol( Boulder 80302 269 270 GREAT BASIN NATURALIST Vol. 35, No. 3 Fig. 1. Winter abundance pattern of the Black-billed Magpie, based on Christmas count data. Open blocks = no birds observed; four de- grees of shading represent > 5.0, 3.0-4.9. 1.0-2.9, and < 1 .0 birds per party-hour-, respectively. mas count data indicate that this species is restricted to the region described by Linsdale (19 37) but that it is not nni- formly distributed \yithin that area. High- est densities were found in the northern Great Basin and on the northwestern plains of Montana, Alberta, and Saskatch- ewan. Densities appeared lower in the southern and eastern portions of the range. Figure 1 suggests that those same fac- tors limiting the distribution of the Black- billed Magpie also may be influencing its abundance within that range. Table 1 shows correlation coefficients between bird abundance and various climatic factors. It is evident that magpie densities are negatively correlated with a variety of temperature and moisture variables. whether or not one includes blocks out- side the range of the species. Tables 2 and 3 show the results of step wise multi[)le regression of magpies and the climatic \ariables listed in Table I. When data for all latitude-longitude bhx ks are used, only two variables (.Jul}' hu- midity and maximum temperature) made a meaningful contribution to ihe regres- sion equation, but these a((i)nnte(l lor W- percent of the \ariation in magpie abun- dance. Restricting the analysis lo llie s[)(>- Table 1. Correlation coefficients between Black-billed Magpie abundance and certain en- vironmental valuables. Bird data are from 1969- 70, 1970-71. and 1971-72 Christmas counts, grouped by blocks of latitude and longitude (see I'ig. 1 ) ; environmental data are 50-year means from USD A (1941). Variable X aiuiual t(>ni > Maximum Im p Mininunn ti'in > No. frost -IVoo la AuTuial precipil,itii>ii Sunmier ]iroci]iitation Winter l)recipitation X Julv humid tv Correlation coefficient Blocks within ,'\11 blocks (n = 47) or adjacent to magpie range (n = 26) -.38* -.20 -.39* -,43* -.41* -.40* -.36 -.51* -.45* -.33 -.45* -.31 -.24 -.59* -.12 -.50* cies' range resulted in a stepwise regres- sion including four independent variables and accounting for 59 percent of the pat- tern of magpie density (Table 3). Frost- free days entered as the best predictor (negative) within the range. This, how- ever, is simply another parameter of tem- perature I'egime. Table ^ shows that the same famih' ol \ariables is involved with- in the sj)ecies range — namely, an inverse relationship to temperature and moisture (luring the warm season of the year. Tablk 2. Stepwise nuiltiple regression of eiglit environmental variables against winter alnuidance of Black-billed Magpies. Based upon Christmas count data and climate variables for 17 latitude-longitude blocks in the United States ( see text and Fig. 1 ) . Step no. Variable entered R* R=** 1 .luly humiihty 2 Maximum temperature 0.59 0.74 0.35 0.54 \V = iiiiillipU; (oncl.iUnii ,(.f-fririoiil I\-" r= 1 (H'ltiriciil (if (Iclorniinntinii. (^(| uv;il0Mt t .1 .,1 rnrl percent ■;tep. 'i'\m r. ;. Same , upon 26 latitude-Ion States within or adii range (see Fig. 1 ). Table 2. except based only ude blocks in the United •ut to Black-billed Magpie Step no. Variable entered 1 No. frost-free davs 0.51 0.26 2 .Fulv humidity 0.71 0.51 5 Summer precipitation 0.75 0.56 4 Maximum temperature 0.77 0.59 Sept. 1975 BOCK, lepthii:n: aia(;pii. disikibtttion 271 Discussion and ('oxc.i.i mons Results of this study show [\\o dcgroo to which the Black-billed Magpie is a bird of cool arid climates in the United Slates. This does not necessarily mean that mag- pie distribution and abundance actually are determined by climate, or r\ou h\ the influence of climate on food. It could be that the species is restricted by babitat availability or by the appearance of close competitors coincidental with changes in climatic regime. We can ])ro\ ide no dc- finitiye answer to this dilennna. but a \v\\ considered speculations are in order. ])ar- ticularly as they might stinudate more research. First, there is no clear break in suitable habitat in the United States wlncli shotdd Imiit magpie distribution, except in the West where the closed coniferous forests constitute a sharp boundary. Riparian habitat along such watercourses as the Rio Grande and the North Platte. South Platte. Arkansas, and Colorado rivers all support Black-billed INIagpie ])opulations; yet. Pica pica become scarce and disajipear along these rivers when they reach the hot Southwest or the more humid central plains. Subtle habitat changes may occur, but it is not clear wdiy such an ojiportu- nistic species should be limited by them. Concerning competitors, it is obvious that Pica pica is replaced by the closely related P. nuttalli in interior California. Magpies forage opportunistically on in- vertebrates (especially grasshoppers), car- rion, and various other items (Linsdale, 1937; Verbeek, 1973). It is very difficult to describe the foraging niche of such a species, but the Common Crow (Corvus brachyrhynchos) and White-necked Ra- ven (C. cryptoleucus) appear geherall^ similar in food and habitat requirements (Bent, 1946). The White-necked Raven is a bird of the Southwest. The Common Crow is distributed all across the United States, but is especialh' abundant in cen- tral and eastern regions (Bystrak. 1974). While these two species of (^onnis do roughly circumscribe the range of the Black-billed Magpie, we would be rebu- tant to conclude, without more field evi- dence, that they are invohed in a com- petitive exclusion. First, the White-necked Raven is uncommon (Bystrak. 1974) and more typical of southwestern grasslands than the actual hot desert country (Phil- lips et al., 1964) marking the boundary )l magpie (bsl fibul ioi i. Second, tlie (oni- inou (j-ow is. in Die I. \^i(lel\■ s\ lupalfic uiib ilie P.lac k I. died Mag[.ie. hnisdale and lid ( l')-57 ' i-e|Mii-ls ,,Mi\ (ic lasi iiilefadioiis belweeu (i-o\\s and niagjiies. Vei-beek ( I'iZ )! ohsecNcd a luimber of in- teractious between ci-o\\s and Pi((/ iiut- tdlli: bc)\ve\c'f. tb(>se oc c urred onh .u-ounci llie uesi and nia\ liaxc imohed a response to the crows as potential Jiest predators. At other seasons the two species were loleccuit of each other. Finally, Pica pica is Ilolarc tic in distribution and in Great Pjiitaiu and luirope is sympatric with four species of Corvus with generally similar haliitat recpnrements (^Bannerman, 1953). ^^ (' would like to conclude by returning to the cpie.iic f climate. It seems very likel^ Ihal ihe Black-billed Magpie (and/ or their in\ (>rtebrate prey) cannot tolerate the extreme tem])eratures of a warm des- ert. Verbeek (1972:571) noted that Pica nuttalli forage in summer mainly in the morning, partly because midday heat even in California stops the activity of inverte- brate prey, and ])artly because "this same heat sevcM'ly limits the birds in their feed- ing." In the Old World magpies are dis- tributcMl throughout Europe and into North Africa as far as the edge of the Sahara ( Baimerman. 195 3 ) . In New Mex- ico Pica pica breeds only in the northern third of the state; yet in the cooler winter season birds may wander down the Rio Grande Valle\ considerable distances (Ligon, 1961). While the eastern distributional limits of the Black-billed Magpie do not fit \'S'ith striking pin siographic or habitat changes, the\' clo coincide' \\ith major changes in climatic regime which appear to have general avifaunal significance. For ex- am])le. Salt (1952) concluded that the House Finch [Carpodacus rncxicanus) does not breed eastward in the Great Plains because of its intolerance of high summer humichty. The eastern limits of this sj)ecies iwv \qy\ sinnlar to those of the magpi(\ The eastward disappearance of Pica pica also is generally coincidental with rather steep zones of introgression between east- ern and western ])opulations of several bird species. The most familiar of these is the /one of ■'h\])ridization" between the western [cafcr] and eastern (auratus) subspecies of the Common Flicker {Colap- les auratus), analyzed by Short (1965). 272 (;ri:at basin xaii hai Vol. ^5, No. 3 Interestingly, a third subspecies, (\ a. chrysoides. replaces cafer in the South- west, although this zone is somewhat south of the limits of magpie distribution. .h)lu^ son (1969:229) reviewed the situation in flickers and made these comments; One of llio niosl sliikiii^ ivn .'lalions tliat ranie to me upon exaniiiiiiig Slioi't's generalized maps of tlie geof^iajihic dis- tribution of flickers in Nortii America is the great coincideiKc of l)llenot^•pic change in the various Imnis willi major climatic boundaries on the (onlinent. Rising (1969), in fact, studied the com- parative physiologies of Northern Orioles {Icterus galbula) which, in the same part of the western Great Plains as the flickers, undergo marked phenot^^pic change. He found that the westerti "Bullock's Oriole" (/. g. huUocki) is better adapted to hot dry climate than is the eastern "Baltimore Oriole" (/. g. galbula). We suggest that climatic fac tors related to temperature and humidity may limit the abundance and distribution of the Black-billed Magpie, either directly or by their effects upon the availability of in- vertebrate prey. Future research on this interesting bird coidd profitably include ecophysiological studies as \\{A\ as more fieldwork on the relationships of this spe- cies to subtle habitat changes or the ])res- ence of competitors, especially the Com- mon Crow^ Acknowledgments. — We are grateful to the University of Colorado Computing Center and Taximetrics Laboratory for technical assistance, and to the many stu- dents who have helped to build the Christ- mas count data banks. This ])roject was supported by grant GB 368f)() from the National Science Foundation. Literature Cited B.\NNERMAN, D. A. 1953. The birds of the British Isles. Vol. I. Oliver and Boyd. Edin- burgh. 356 pp. Bent, A. C. 1946. Life histories of North American jajs, crows, and titmice. V.S. Nat. Mus. Bull. 191. Bock, C. E., .\nd L. W. Leptiiien. 1974. Win- ter jiatterns of bird species diversity and abundance in the United States and southern Canada. Am. Birds 28:556-562. Bysthai-;. 1). I'*7K Wintering areas of bird s|i(>(ics |)(]|cnliall\- hazardous to aiicraft. Na- tional Audubon Soc. N.Y. 15G pp. Dixon. W. ,I. (ed). 1971. BMD Biomedical cominitei' programs. I'niv. Calif. Publ. in Automatic Computations no. 2. 600 pp. EariNo. M. .1. 1968. Nest-related activities of Black-billed Magpies. Condor 70:154-165. G.\nRiEr,soN. L N.. .\nd S! G. .Tewett. 1940. Birds of Oregon. Oregon St. College. Cor- vallis. 650 pp. Grinnele, .T., .\nd a. H. Mieeer. 1941-. The distribution of the birds of California. Pac. Coast Avifavuia no. 27. Tewett. S. G., W. P. T.weor, W. T. See\av. .\nd .1. W. Aedrich. 1953. Birds of Washing- ton State. TTniv. Washington Pi-ess. Seattle. 767 pp. Johnson, N. K. 1969. Review: three papers on variation in flickers (Colaptes) by Lester L. Slioit. Jr. Wilson Bull. 81:225-230. Jones. R. E. 1960. Activities of the magpie dur- ing the breeding period in southern Idaho. Northwest Sci. 34:18-24. Eicon, J. S. 1961. New Mexico birds and where to find them. Univ. New Mexico Press. .Mbuciuerque. 360 pp. EiNsnsTi:. J. M. 1937. The natural history of magpies. Par. Coast Avifauna no. 25. PlIIEEII'S. \. R.. J. T. M.\RSHAEE, .\ND G. MoNSON. 1964. The birds of Arizona. TTniv. Arizona Pi-ess. Tucson. Rising. J. D. 1969. A comparison of metabo- lism and e-vajiorative watei- loss of Baltimore and Bullock Orioles. Comp. BincJieuE Phvsiol. 31:915-925. Sait. G '\'\' 105 2. Tlie relation of metaboli'^m to (limat(> and distribution in three finches of the gtMius Carpndacus. Ecol. Monogr. 22: 121-152. Short. E. E.. Ti-. 1965. Hvbiidization in the flickers (Colaptes) of North America. Bull. Am. Mus. Nat. Hist. 129:307-428. U.S. Dep.\rtment op Agriculture. 1941. Cli- mate and man (yearbook of agrii ultuie). TJSDA. Washington. D.C. Verbeek, N. A. M. 1972. Daily and annual time budget of the Yellow-billed Magpie. Auk 89:567-582. 1973. The exploitation svsteni of the Yellr.w billed Magpie. TTniv. Calif. Publ. Zool. 99:1-58. NECTAR COMPOSITION OF HAWKMOTH-VISITED SPECIES OF OENOTHERA (ONAGRACEAE) Robert E. Stockliouse, IF Abstract.- Noctars of 11 hawkmotlis-visited taxa of Ocnolhcra were studied. Qualitatively the ;ars of all taxa ucre identical. The amount of nectar prochjced \)vv night among the taxa was quite •iable. Potential energy availabh- from nectar of an average flowei- of O. caespitosa was deter- ned 1,. l.c 42.1 , ab.nes. There is litth' iiifoi-inalioii on the amount of nectar prochued. its composi- tion, or nutritive j)oteiitial foi- hawkmoth- visited flow'ers. (Gregor^', Aliso '5:357- 419), 1963/64); (Heinrich and Raven, Science 176:597-602, 1972); (Handel et al., Am. Jour. Bot. ;59: 1030-1032, 1972); Baker and Baker, Sttidies of nectar — con- stitution and ])olhnator — plant coevolu- tion. Pages 100-140 in Coevolution of Ani- mals and Plants). The nectars of 10 hawkmoth- visited taxa of OcJiothcra were studied to determine quantity and sugar composition available to nocttirnal visitors. Methods. — Nectar was collected \u capillary tubes (5 microliter) in the field or from plants grown in the greenhouse. The component sugars were determined using ])aper chromatography. Whatman No. 1 filter paper (5" x 18") was spotted with '5 microliters of nectar from each species and three standard sugars. Each chromatogram was run in butanol, etha- nol, and w'ater (10:6:4 v/v/v) for 44 hours until the solvent had nearly reached the end of the paper. The chro- matograms were allowed to (h'v and wer(> spra\ed ^^ ith a sugar spra\' ( (SO n\\ of 95% ethanol, 10 m\ of 40% TCA, and 10 nd of glacial acetic acid, which was sattirated with benzidine dihydrochloride),' which made the sugars visible. The j)ercentage sugar was determined with s Average height of nectar in hypanthium (cm) Average length of Species Average in microliters Range in microliters hypanthium (cm) caospitosa cximia 32.5 35.0 26.0 35 20 20 8 18-69 5-32 17-23 4-9 6.2 4.0 3.0 3.0 9.4 5.5 29.5 16.0 priniiveris 34.0 5.0 xylocarpa 33.5 The amount of jiotetitinl energy avail- able from an average flower of O. caes- pitosa was calculated. \\\e flowers av- eraged 35 microliters of nectar ])er night, of which a]i[)roximately 32.5% was sugar. [Of the 35 microliters approximately 25.5 (73%) were available when the flower opened at sunset, 6 additional had been produced by 9:15 p.m., the remainder by 8 a.m. the following morning.} Assuming there are approximatel^' 3.7 calories per mg of glucose (Fleinrich and Raven. Science 176: 597-602, 1972) there are 42.1 calories per flower per night avail- able for hawkmoths [35 mg nectar flower X 0.325 (sugar concentration) x 3.7 calories mg sugar = 42.1 calories per flower.] Even as a rough approximation, it is clear that each flower is a large po- tential energy source for hawkmoths. E^■en in small populations with only 20- 50 flowers open on a given night, 42 cal- ories per flower would offer a large energy reward for the hawkmoth pollinators. Acknowledgments. — I wish to thank Steve and Susan Chaplin and Pat Wells for helpful comments on this manu- script. This study was supported in part by a grant-in-aid of research from the Scientific Research Society of North America. A REVISION OF THE NEARCTIC SPECIES OF CLINOHELEA KIEFF1<:R (DIPrERA: CERAIOPOGONIDAE) William L. Giogaii. Jr.' and Willis W. Wirtlr Abstract. — The seven species of Clino/ielea known to inhabit North America are described and illustrated, and a key is provided for identification. Two species groups are recognized: the unimacu- lata group and the bimaculatd group. Clinohclca longitheca and C. pscudonubifera are new. Clino- Iielea nebiilosn (Malloch) is a synonym of C. rurrici (Coquilh^tt) new synonymy. ClijioJielca Kioffer is a fairh lis of ceratopogoiiids. worldwid mall t^eii- in (listri- i)iition. Little is known of their bioloo^•; hut the adult females are predaceous ou other insects, and the larvae are aquatic. The fiAe ])reviously known North Amer- ican species were described bv Loew (1861), Adams (1903), Coquillett (1905). Malloch (1915), and Wirth (1952). Most of these species were originally described in the genus Ccratopoiron INleigen and later transferred to Palpomyia Meigen, .Johannseniella Williston, or J nliannscno- rnyia Malloch. Although .lohannsen (1943) correctly placed these species in Clinohelea, the North American species have needed comprehensive revision and a good key for identification. In the present paper seven Nearctic species of CUnohclea, two of which are new, are described and illustrated. Two species groups are recognized and given the names of the oldest named species in their group. All of the types of CUnohelca species from North America have been examined, as well as examples of 17 spe- cies from other parts of the world. All specimens examined unless otherwise noted are part of the collection of the Na- tional Museum of Natural History (USNM) in Washington. In the Jists of speciuKMis examined, slide-moimted spe- ( imens are denoted (S), and pinned spe- ( imens as (P). The types of our new spe- cies will be deposited in the USNM. Measurements and other data are based on slide-moiuited specimens and are re- corded in the manner of Chan and LeRoux ( 1965 ) . When j^ossible, 10 females of each s[)ecies were critically measured. The data are presented in the following man- ner: mean value (mininumi value - maxi- mum value, n = number of measure- ments), except in the case of new species, where the actual values are given for the holotype, and the mean, minimum-maxi- mum, and number of measurements are given in the variation section. Numerical characters for female Nearctic Clinohelea are presented in Table 1. For general terminology of Ceratopogo- nidae see Wirth (1952) and Chan and Le- Roux (1965). The following special terms are used in the descriptions of fe- males. Wing length is measured from the basal arculus to the wing tip. Antennal proportions (AP) are the relative lengths of each flagellomere; antennal ratio (AR) is the length of the proximal 8 flagello- nieres, divided into the length of the distal 5 flagellomeres. Palpal ratio (PR) is the length of the 3rd palpal segment divided T.\BLE 1.- - Numeiical charac te rs of female Nearctic Clinohelea (minimum-maximum values) . Wing length (mm) Wing breadth Costal (inm) ratio Palpal ratio Antennal Femoral spines Species ratio Fore Mid Hind Unimacui,.\ta Group curriei 2.53-3.23 nubifera 2.32-2.42 pseudonubifera .... 2.03-2.19 BlMACLILATA GrOUP bimaculata 1.65-2.68 dimidiata 2.32-2.74 usingeri 2.50-2.74 longitheca 1.97-2.00 0.81-0.94 0.71-0.77 n.65-0.69 0.50-0.74 0.68-0.87 0.70-0.81 0.61 0.81-0.87 0.85-0.86 0.82-0.83 0.82-0.86 0.76-0.82 0.80-0.81 0.82-0.84 3.20-4.50 4.00-4.36 2.86-3,00 2.60-140 3.17-3.80 3.17-3.67 2.89-3.11 1.50-1.66 1.41-1.46 1.35-1.38 1.17-1.36 1.25-1.40 1.29-1.35 1.17-1.36 0-3 0 0 0 0 0 0 0-2 1-3 0-3 2-4 0-1 0-1 'Department of Enlomologv, University of Marvlanii. College Park. Maryland 20742. ^Systematic Entomology laboratory. IIBIII. .\p.iA He Sfry . USD.\. i/o U.S. National Mi 20560. 0-3 0 0 0 Washington, D.C. 275 276 GREAT BASIN NATLHiAMST Vol. 35, No. 3 by its greatest breadth. 'J erniiiiology deal- ing \\'ith male genitalia follows that of Snodgrass (1957) and Chan and LeRonx (1965). All female genitalia and sper- mathecae have been (h-awn to the same scale. Types ha\e been illustrated when- ever possible. We are especially indebted to Mrs. Ethel L. Grogan for preparation of the illustrations. Thanks are also extended to the following jiersons and their institu- tions for the loan of type and other ma- terial or information regarding s])ecimens in their collections: Donald W. Webb, Illinois Natural History Survev, Urbana (INHS); George W. Byers, Snow Ento- mological INIusemn, University of Kansas. Lawrence (KU); Wilford .1. Hanson, Utah State University, Logan (LTSLT) ; and .Tanice C. Scott, Museum of Comparative Zoology. Cambridge, ?slassachusetts (MCZ). Gemis Clijiohclca Kieffer C/ino/irIra Kieffer. I'd 7: 205. Tvi)e-speries. Crrn topogon variegatus WiTincrtz, by original designation. Di.\GNOsis. — Moderately large, shining ceratopogonids, usually with infuscated wings; body nearly bare, rather slender; plenron usually with transverse sihery band. Eyes bare; widely separated. An- tenna slender; flagellomeres 1-8 long, fla- gellomeres 9-13 elongate in female; fla- gellomeres 11-13 elongate in male; plume sparse in male. Palpus slender; 3rd seg- ment slender, lacking a pit. Female man- dible with coarse teeth. Mesonotum mod- erately robust, without himieral pits, a short anterior tubercle sometimes present. Femora slender, occasionally with up to four spines; 4th tarsomere of at least mid and hind legs deeply bilobed, each lobe ending in a stout blunt spine and smaller spines; fore 5th tarsomere greatly swollen in both sexes; fore claws equal, mid and hind very unequal in female; all claws equal in male. Wing long, without macro- trichia; costa extending to 0.75 but not more than 0.90 of wdng length; two radial cells present, 2nd much longer than 1st; no intercalary fork; medial fork broadly sessile. Female abdomen without eversible glands or gland rods; genital sclerotization small, simple; two well-developed sper- mathecae. Male genitalia with 9th ster- luun short, broad; 9th tergum tapered \vitli large cerci; basimere and telomere relatively long and slender; aedeagus with low anterior arch, distal portion broad, underlying membrane extending beyond tip; clasjiettes usually divided, each por- tion slender withan elongated bulbous tip. Iaimaturf. stages. — Larvae are aquat- ic. Wirth (1951) described the pupa of C. hirnaculata. which he reared from the sandy margin of a small stream in Vir- ginia. This is apparently the only Nearc- tic sjiecies that has been described in an innnature stage. Adult habits. — Adults can be found on ^ egetation bordering water, and Gro- gan has taken them at flowers and from a small grove of trees in Utah. Downes (1960,^1971) stated that adult females are [iredaceons on other small insects that are captured in flight, but did not give specific examples. Key to the Nearc tic Species ui Clinoliclca (])rimarily Females) 1. b'ore 5th tarsomere solid bioxMi; wing with two sjiots. one centered over 1st radial cell, second iicvir tip of (osta { hirudcuhitd group) 2 bore 5lh tarsomere with pale b.iiid: \^ill^ with 1 spot (entered over 1st ra- dial (('II or eiitii-('l\ iiiliisc ,il('(l ^ liuinuicnUitd group) 5 2. Spermathe(ae large, elongated, ellipsoid longitJwca n. sp. Spermathecae small, spheroid to ovoid 3 3. Legs pre(loniiii.nitl\ \cllow. dislid one-Fouiili of hind fetnur brown hirriaculata (Loew) Legs prcHJonnn.niilx hfouinsh. disi.d one-hidf ol hind fenuu' brown 4 4. Tibiae entirely biown; basnl arms of nnde a(>(leagus separated dimidiata (Adams) Tibiae yellowish in midportion. basal an(lapi(al portions brown; basal arms of male aede.igus inta( t usingeri V^irXh. Sept. 1975 (JHOCAN. WIRTH: CERATOPOC.OIVIDAE 277 5. Wing with narrow dark infuscation extending from apex of costa to tip nubifera (Coquillett) Wing without narrow dark infuscation at tij) . 6 (). Fore 6th tarsoniere with pale hand twice as long as width of tarsomere curriei (Coquillett) Fore 5th tarsoniere with pale hanrl nnich shorter than width of tarso- .... mere pseudonubifera n. sp. Unimaculata Group Clinohclca curriei (Coquillett) Wing with infuscation usually centered \ ^S- ■< ^) over 1st radial cell or entirely infuscated. Ccratopogon curriei Coquillett, 1905: 62 (female; Fore 5th tarsomere with pale band. At British Columbia) least hind femur wdth spines usually Palpomyia curriei (Coquillett); Malloch 1914: present. Species examined in this group 219 (combination, description; key) X r i\T 4.1 A • /^ • 7 J Llinonelea curriei (Coquillett); Johannsen, 1943: not from North America: C. umnmculata 733 ^combination); Wirth, 1965: 136 (distri- (Macquart), Europe. bution) Fig. 1. Clinohelea curriei ( Coquillett] female: a. antenna; b. leg pattern; c. variations in hind leg pattern; d, wing; e. genitalia. 278 GREAT BASIN NATURALIST Vol. 35, No. 3 Palpomyia nebulosa Malloch. 1915: ^22 (female; Miciiignn). new synonymy Clinohelea nebulosa (Malloch); Johannsen, 1943: 783 (combination); Wirth, 1965: 136 (distri- bution) Diagnosis. — Distinguished from all other Nearctic Clinohelea by the following combination of characters: fore 5th tarso- niere with pale band longer than width of 5th tarsomere; wing with infnscation cen- tered over 1st radial cell or entirely in- fuscated. Female.— Wing length 2.78 (2.53- 3.32, n = 10) mm; breadth 0.85 (0.81- 0.94, n - 10) mm. Head: Brown. Antenna (Fig. la) slen- der; pedicel yellow to pale brow^i; basal flagellomere with j)roximal two-thirds j)ale, distal one-third brown; remaining flagellomeres brown; AP 22-10-10-10-10- 10-11-12-30-28-29-29-31 (n = 10); AR 1.57 (1.50-1.(56, n = 10). Palpus brown; 3rd segment longer than 5th; PR 4.03 (3.20-4.50, n = 10). Mandible like that of C. bimaculata (Fig. 4c). Thorax: Mesonotum, scutellum, post- scutellum dark browTL Fegs (Fig. lb) yellow; apex of fore tibia, mid femorotibial area, fore distal 3 tarsomeres, mid and hind 4th and 5th tarsomeres brown; hind leg pattern variable, most common form (Fig lb) with distal one-sixth of femur and tibia brown, other hind leg patterns as in Figure Ic; fore 5th tarsomere with pale band longer than width of tarsomere; 0-3 fore, 0-2 mid, and 1-3 hind femoral spines. Wing (Fig. Id) usually with infuscated area centered over 1st radial cell or en- tirely infuscated; veins brown; CR 0.84 (0.81-0.87, n = 10). Halter pale. Ahdorrien: Brown. Genitalia as in Figure le with a ])air of slender, pos- teriorly directed sclerotized arms arising anteriorly from a lighter sclerotized area. Spermathecae small, spheroid to ovoid, subequal to unequal with short nee k^. Male. — Similar to female with follow- ing differences: smaller; antennal pedicel flark brf)wn. flagellum brown; legs more diffused with brown; 0-1 fore, 0-1 mid, and 0-2 hind f(>moral spines. Genitalia form and shape like that of C. bimaculata 'Fig. 4f ) ; aedeagus as in Figure 6a. Distribution. — Alaska and California to Newfoundland and Florida (locality records plotted in Figure 3). Types. — Holotype female of C. curriei, Kaslo, British Columbia, 17 .Tune 1903, R. P. Currie (Type no. 8361, USNM); holo- tyj)e female of C. nebulosa, Grand Junc- tion, Van Buren Co., Michigan, 15 July 1914, C. A. Hart (INHS). Specimens examined. — 82 slides, 218 pinned specimens from: ALASKA: Anchorage (Aldrich) ; Matanuska (Chamberlin). CALIFORNIA: Eldorado Co., Lu- ther Pass (Schlinger. Univ. Calif. Davis). CON- NECTICUT: Fairfield Co.. Redding (Melander); Tolland Co.. Storrs (Melander). DELAWARE: New Castle Co., Delaware City. FLORIDA: Ala- chua Co.. Gainesville (Wirth). IDAHO: Ada Co., Boise (INHS); Nez Perce Co., Sweetwater (Aid- rich). INDIANA: Porter Co., Mineral Springs (INHS). IOWA: Hancock Co., Pilot Knob St. Park (Gaud). MAINE: Hancock Co., Bar Harbor (Johnson). MASSACHUSETTS: Franklin Co., Row." (Cohcr); Middlese.x Co., Bedford (Wirth); Concord (Wirth); Suffolk Co.. Boston (Melander). MICHIGAN: Cheboygan Co. (Dreisbach), Doug- las Lake (Williams) ; Clare Co. (Dreisbach) ; Iron Co. (Dreisbach); Lake Co. (Dreisbach); Livingston Co., George Reserve (Sabroskj-, Steys- kal); Manistee Co. (Dreisbach); Midland Co. (Dreisbach); Nottawa (Sabrosky. Dreisbach); Missaukee Co. (Dreisbach); Osceola Co. (Dreis- bach); Roscommon Co. (Dreisbach); Van Buren Co., Grand Junction (Hart, holotype of nebulosa); Wexford Co.. (Dreisbach) . MINNESOTA: Ram- sey Co. (Wall). NEBRASKA: Cherry Co.. Hack- berry Lake (Wirth), Pelican Lake (Wirth). NEW HAMPSHIRE: Grafton Co., Stinson Lake (Wirth). NEW YORK: Chautauqua Co., S. Day- ton (Wirth); Erie Co.. East Aurora (Van Duzee), East Concord Bog (Wirth); Franklin Co.. Adiron- dacks (Melander); Lew^is Co.. Brantingham Lake (Wirth). Letchworth St. Park (Wirth), Whet- stone Gulf (Wirth); Monroe Co., Braddock Bay (Wirth); Orleans Co.. Albion (Wirth); St. Law- rence Co.. Cranberry Lake (Wirth); Suffolk Co., Cold Spring Harbor (Melander); Tompkins Co., Ringwood Reserve (Wirth). OHIO: Summit Co. (Lipovsky, KU). UTAH: Cache Co.. Hvrum (Grogan). VERMONT: Caledonia Co.. Lyndon (Melander). VIRGINIA: Alexandria (Wirth); Fairfax Co.. Dead Run (Wirth). WISCONSIN: Polk Co. (Baker, paratype of nebulosa, INHS). WEST VIRGINIA: Pocahontas Co.. Cranberry Glades (Wirth. Sabrosky). BRITISEt COLUM- BIA: Kaslo (Currie. holotype of curriei). Quebec: Meach Lake (Wirth). NEWFOUNDLAND: Squire's Mem. Park (Alexander). NOVA SCOTIA: Baddeck (Fairchild). ONTARIO: Al- gonquiti Park (Wirth); Kemptville (Wirth); Ot- tawa (Melander. Wirth); Toronto (Van Duzee); Waubamick (Melander). Disci TssioN. — The Palaearctic species, C. unimaculata (Macquart) closely re- sembles C. curriei. However, the hind tibia is pale except for the narrow base and apex; the apices of the fore and mid fe- mora are conspicuously dark, narrowly on the fore leg but more broadly on the mid leg; atifl the hind femur lacks any Sept. 1975 CROCAN, WIHTH: CKKATOPOCON IDAi: 279 trace of infuscation except tlie conspicuous a[)ical (lark band. Clitiohclcd nuhifcra ( Coquillett) (Fig. 2a. c. e. g; fib) Ceratopogon iiubifer Coquillett. 1905: f)l li'eniale; Florida) Palpomyia nubifera (Coquillett); Mallocb. 1914: 217 (combination; key) Clinohelea nubifera (Coquillett); .lohannsen, 1943: 783 (combination); Wn-tli, 1965: 136 (distribution) Diagnosis. — Distinguished from all other Nearctic Clinohelea by the following combination of characters: wing with in- fuscation centered o^■er 1st radial cell and a narrow infuscated band extending from apex of costa to wing tip; and fore 5th tarsomere with pale band. Female.— Wing length 2.35 (2.32- 2.42, n = 3) mm; breadth 0.74 (0.71- 0.77, n = 3) mm. Head: Vertex and proboscis brown, frontoclypeus lighter brown to yellowish. Antenna (Fig. 2a) slender; pedicel yellow to light brown; proximal 5-8 flagellomeres j)ale on basal portions, distal portions light brown; distal 5 flagellomeres brown; AP 26-12-1 1-1 1-1 1-1 1-1 1-12-32-29-30-30-30 (n = 3); AR 1.43 (1.41-1.46, n = 3). Palpus Fig. 2. Female Clinohelea: a,c,e,g, C. nubifera (Coqudlett); ,b, antennae; c,d, leg patterns; e,f, wings; g,h, spermathecae. d.f.h. C. pscudonubifera n. sp. 280 GREAT BASIN NATURALIST Vol. 35, No. 3 brown; 3rcl segment nearly twice as long as 5th; PR 4.12 (4.()()-4.36" n = 3). Man- dible like that of C. himaculata (Fig. 4c). Thorax: Mesonotuni, scutellnm, post- scutellimi dark brown. Legs (Fig. 2c) yellow; proximal jiortions of mid and hind coxae, distal i)ortion of hind femur, apices of fore and mid femora, all of hind tibia, proximal one-half of fore and mid tibiae, and 4th and 5th tarsomeres brown; fore 5tli tarsomere with })ale band shorter than width of tarsomere; 0-2 mid and 2-4 hind femoral sj)ines. Wing (Fig. 2e) veins brown; infuscated area centered over 1st radial cell, and narrow infuscated band extending from apex of costa to tip; CR 0.86 (0.85-0.86. n - 3). I4alter stem brownish; knob pale. Abdomen: Brown. Sjiermathecae (Fig. 2g) small, ovoid. sid)equal with short ]iecks. Male. — Unknown. A female specimen from Santa Rosa Co., Florida, had male genitalia moimted with her on a slide. These male genitalia were remounted in an attempt to examine them in detail. The overall shape and form of the genitalia are like that of C. himaculata (Fig. 4f ) ; aedeagus as in Figure 6b. Dlstkibution. — Florida, New York (locality records plotted in Figure 3). Type. — Holotype, female, Jacksonville, Florida, Mrs. A. T. Slosson (Type no. 8357, USNM, pinned). Specimens examined. — From the fol- lowing localities: FLORIDA: Alachua Co., Gainesville (Blan- ton), 2 females (S); Jacksonville (Slosson, holo- type female. P); Highland Co., Sebring (Wirth), 1 female (P); Santa Rosa Co., Blackwater River (Fairchikl), 1 female (S). NEW YORK: Suffolk Co., Cold Spring Harbor (Melander), 1 female (P). Clinoheh'a pseudoimhifera Grogan and Wirth, n. sp. (Fig. 2b, d, f, h; 6c) Clinohelea species 1; Wirth, 1951: 321 (females; Virginia). Diagnosis. — Most closely related to (\ uulnfera. and can be distinguished from all other Nearctic Cliiiohdea by the fol- lowing combination of characters: fore 5th tarsomere with very short, pale band, legs mostly yellow wdth hind tibia and distal five-sixths of hind femur brow^n, and wing nubiTera C. pseudonubifera n Fig. 3. North American localitv records for Clino/irlra of tiic iiniiuaculatn group. Sept. 1975 r,RO(;.\X. WIHTII: ceratopogonidae 281 with iiifuscatioii ((Mitci-cd nxcr 1st r-adial cell. Female iioloiyim:. — \^illl^ Iciit^tli 2.0^ mm; breadth ().(y5 mm. Head: Vertex and [jroboscis brown; frontoclypeiis lighter brown. Antenna (Fig. 2b) slender; pethcel yelloNv. j^roximal 4 flagellomeres pale, distal 9 flagellomeres brown; AP 17-9-9-8-9-9-9-10-21-21-21-21- 26; AR \.W. Pal|)ns brown; ^rd segment about as long as 1th; I^R 100. Mandible Hke that of ('. hiffit/culafa (Fig. 4c). lljorai: Mcsoiiotnm, scutellum, j)ost- scutellum dark lirown. Legs (Fig. 2d) yellow; ])roximal jiortions of coxae, (hs- tal five-sixths of hind femur, distal one- fifth of mid fenuir, mid and hind tibiae, and 4th and '5th tarsomeres of tarsi browu; fore tibia very light brown; fore 5th tarso- mere with ver^' short, pale baud; mid and hind femora with 1 spine. Wing (Fig. 2f) with dark infuscation centered over 1st radial cell; \eins brown; CR 0.8 3. Hal- ter pale with dark brown spot on knob. Abdomen: Rrown. Spermathecae (Fig. 2h) small, o^•oid. subequal, with short necks. Male .allotype. — Similar to female holot^"pe with the following differences: smaller; antennal ])edicel dark brown, flagellum brown; femora lacking spines. Genitalia sha{)e and form like that of C. himacidata (Fig. 4f); aedeagus as in Figure 6c. Etymology. — The iiame pseudotinhi- fera refers to the resembhnue to ('. nuhi- fera. Varl\tion. — The following characters were recorded for the single female to])o- t^•pe: wing length 2.19 mm; breadth 0.69 mm. AR 7.35. PR 2.86. CR 0.82 The general coloration of all of the paratypes is like that of the holotype. Femoral spines ranged from 0-1 mid. and 0-1 hind. Distribution. -Ontario to North (>aro- lina (locality records plotted in Figure 3). Types. — Female holotyjje, male allo- type, 1 female j)aratype. Snow I [ill. Worcester Co., Maryland, 2 June 1968. W. H. Anderson, light tra]) (Tvpe no. 66495, USNM). Other paratyj)es, 4 pinned females as follows: NORTH CAROLINA: Macon Co., Highlands. 15 .Tune 1957, .1. R. Vockeroth. 1 female (Canada Nat. Coll.). ONIARIO: Ottawa, Mer Rleue. 23 .June 1952 G. E. Shewell, 1 female (CNC). VIRGINIA: Fairfax Co., Falls Church, 4 July 1950, W. W. Wirth, 2 females. Discussion. — Wirth (1951) in refer- ence to the two female specimens from Falls Church, Virginia, stated that they were close to C. nubifera and C. dimidiata but declined to name them at the time. Bimaculata Group \\'ing with two nifuscated areas, one (entered o\er 1st radial cell, the other near tij) of costa. Fore 5th tarsomere uniformly brown. Femora usually lack- ing spines, or if present, only on hind femur. Species examined in this group not from North America: C. barrettoihane and Duret, Brazil; horacioi Lane, Brazil; rieirai Lane. Brazil; nigripes Macfie, Bra- zil; pachydactyla Kieffer, Singapore; nibriceps Kieffer, Paraguay; saltanensis Lane and Duret, Argentina; townesi Lane, Brazil; toivnseudi Lane, Brazil. (linohelea bimaculata (Loew) (Fig. 4, 6d) Ceratopogon bimaculatus l.oew, 1861: 311 (fe- male; Wasliington. D.C.). Johaiinsrniella bitnaculain (Loew); Malloch, 1914: 226 (combination; description; kejO- J ohannsenomyia bimaculata (Loew); Malloch, 1915: 352 (combination; l^ey). Clinohelea bimaculata (Loew); Kieffer, 1917: 317 (combination; key; fig. tarsus); Wirth, 1951: 321 (description and fig. pupa); Johannsen, 1952: 164 (key, fig. tarsus); Wirth, 1965: 136 (distribution). Diagnosis. — Distinguished from all other Nearctic Clinofielea by the following combination of characters: legs mainly yellow with a dark subapical band on the hind femur, two-spotted wings, and an- tennal pedicel yellow. Female.— Wing length 2.06 (1.65-2.68, n 10) nun; breadth 0.63 (0.50-0.74, n = 10 ) mm. Head: Frontovertex brown; j)roboscis and palpus pale yellow. Antenna (Fig. la ) slender; pedicel yellow, proximal 8 flagellomeres brown, distal 5 flagellomeres lighter brown; AP 20-11-10-10-10-10-11- 12-24-23-23-23-25 (n = 10); AR 1.24 (1.17-1.36, n = 10). Palpus with 3rd seg- ment slightly longer than 5th; PR 2.87 (2.60-3.40, n = 10). Mandible (Fig. 4c) 282 GREAT BASIN NATURALIST Vol. 35, No. 3 Fig. 4. Clinohelea bimaculata (Loew): a-e, female; f, male genitalia; g-i, pupa; a. antenna; b, wing; f, mandible; d, leg pattern; e. genitalia; g. operculum; li. respiratory organ; i. anal segment. heavily sclerotized; inner margin with six to nine large coarse teeth; outer margin with four or five small teeth. Thorax: Mesonotum, scutellum, post- scutellum brown. Legs (Fig. 4d) yellow; proximal portion of hind coxa, subapical band on hind femur, and 1th and ^th tar- someres brown; hind f(Mnur witii 0-3 spines. Wing (Fig. lb i veins brown; in- fuscations centered ov(n- 1st raihal cell and just before tip of costa; occasionally, in- fuscated areas joined posteriorly; CR 0.84 (0.82-0.86, n = 10). Halter' stem pale yellowish; knob white. Abdomen: Brown to reddish brown. Genitalia as in Figure 4e wdth a pair of slender, sclerotized, anteriorly directed arms arising from shorter, thicker arms. Spermathecae small, spheroid to ovoid, subequal to unequal, with short necks. Mali.. — Similar to female with the following differences: smaller; antennal pedicel brown; hind femur lacking spines. Genitalia as in Figure 4f. Ninth sternum about three times broader than long, base shghtly curved with a caudomedial exca- vation; 9th tergum tapered distally to a rounded tip. cerci short, not reaching apex Se])t. [97 1 CKOCAIV. WIRTH: CllHATOPOGONIDAE 283 of basimeres. Basiniorc sli^htl^ curved. 2.5 times longer tliaii broad; telomere slightly longer than iiasimer(\ luiNcd. tapered distally ^^ith poiiiti^d. hooked lip. Aedeagus (Fig. ()d i hea\ il\ selerolized, triangular, aliout as broad a>> long: basal arm recurved about 'M) degret^s. hea\ily sclerotized; (hstal portion with iilinit pointed ti]i; underl^ ing membrane ex- tending beyond ti]). rounded with a dark spot. Clasj)ettes di^ided; basal arm heavily sclerotized, recm'\ed; (hstal portion more lightly sclerotizecb ti]) (>longate. bid- bous. PrPA. — Length 3.5 nmi; color light brown. 0])ercuhmi (Fig. 4g) narrow; 0.9 times as broad as long with a ]iair of rounded tubercles bearing long seta; sur- face with fine tubercles, those on lateral margin sharp and setose. Respiratory horn (Fig. 4h ) moderately long and slen- der, about iWo times longer than broad with 10 aj)ical spiracular paj^Uae. Anal segment (Fig. 4i) about twice as long as broad; surface covered with fine tubercles; apicolateral processes about one-third of total length, with subapical fine tubercles; tips heavih' sclerotized and sharply ])oint- ed. Distribution. — Michigan and Texas to New Hampshire and Florida (locality records plotted in Figure 7 ) . Type. — Holotype, female, Washington, D.C., Osten-Sacken coll. (Type no. 10379, MCZ). Specimens examined. — 177 slides, 158 pinned specimens from: ALABAMA: Mobile Co.. Mobile ( Blantou, Cannon). CONNECTICtJT: Litchfield Co.. Lake Waramaug (Melander). DLSTRICT OF COLUM- BIA: Washington (Coquillett ) . FL0RII3A: Ala- chua Co., Gainesville (Blanton. Wirth); Baker Co., Olustee (Blanton) ; Bay Co.. Panama City Beach (McElvey); Calhoun Co.. Blountstown (Blanton); Collier Co.. Collier Seminole St. Park (Wirth), Ochopee (Blanton); Escambia Co., Bratt (Blanton); Glades Co.. Palmdale (Irons); Gulf Co., 2 mi. N Beacon Hill (Blanton). Wevva- hitchka (Blanton); Hardee Co., Ona (Irons); Highlands Co., Archbold Biol. Sta. (Wirth). Lake Placid (Layne), Sebring (Wirth); Indian River Co., Fellsmere (Wirthj. Vero Beach (Wirth); Duval Co.. Jacksonville (Knight) ; Jefferson Co., Monticello (^'V^^itcomb) ; Lake Co., Leesburg (Braddock); Leon Co., 3 mi. N Tallahassee (Blan- ton); Liberty Co.. Torreya St. Park (Blanton, Fairchild. Weems, Wirth); Marion Co., Juniper Springs (Wirth); Orange Co.. Lake Magnolia Park (Irons), Rock Springs (Wirth); Palm Beacli Co.. W. Palm Beach (Hardy, KU); Putnam Co., Lon's Lake (Blanton); Sarasota Co., Myakka i^iver St. Park (Wirth); Suwanee Co., Suw,'anee Springs (Beamer, KU); Wakulla Co., Ocklocko- iiee River St. Park (Wirth); Walton Co. (Butler). (iKORGTA: Charlton Co., Okefenokee Swamp i Beamer. KU); .Mitchell Co., Newton (Pratt); iiiomas Co.. ThomasviUe (Palmer). ILLINOIS: llenrv Co.. Algonquin (INHS); Champaign Co., Urbana (Malloc.h. INHS); Piatt Co., Monticello i.Malloch, INHS); Pulaski Co., Pulaski (Malloch, INHS). INDIANA: Tippecanoe Co., Lafayette (Aldrich. Melander). LOUISIANA: East Baton Rouge Parish. Baton Rouge (Wirth). MARY- LAND: Anne Arundel Co.. Mayo (Wirth); Cal- v(Mt (ji.. (;hesapeake Beach (Shannon. Knab); (^harh's C>o., Nomomonee (Wirth); Frederick Co., Thurmont (Steyskal); Montgomery Co., Glen Echo (Malloch); Prince Georges Co.. Beltsville (Malloch); Worcester Co.. Snow Hill (Wirth). MASSACHUSETTS: Middlese.x Co., Bedford (Wirth). Concord (Wirth). MICHIGAN: Lapeer Co.. Deerfield (Stevskal); Livingston Co., George Reserve (Stevskal); Midland Co.. (Dreisbach); Wayne Co., Detroit (Steyskal). NEBRASKA: Nemaha Co., Peru (Harmston). NEW HAMP- SHIRE: Grafton Co., Stinson Lake (Wirth). NEW YORK: Franklin Co.. Adirondacks (Melan- der). NORTH CAROLINA: Durham Co., Nel- son (Beamer. KLT); Onslow Co.. Jacksonville (Bo- hart. USU). SOUTH CAROLINA: Georgetown Co.. Hobcaw House (Henry). TENNESSEE: Lake Co., Reelfoot Lake (Snow). TEXAS: Col- lin Co., Piano (Tucker); Kerr Co.. Hunt (Wirth), Kerrville (Bottimer). VIRGINIA: Alexandria (Wirth); Fairfax Co., Falls Church. Montgomery Co.. Blacksburg (Messersmith). WEST VIR- GINIA: Pocahontas Co., Cranberry Glades (Wirth. Sabroskv) ; Taylor Co., Grafton (Steys- kal). CUnohclcd dimidiata (Adams) (Fig. 5a, e, g; 6e) Ceratopogon diinidiatus Adams, 1903: 27 (female; Arizona). Johannseniella dmndiata (Adams) ; Malloch: 226 (combination; key). ] ohannsenomyia dimidiata (Adams) ; Malloch, 1915:332 (combination; key). Clinohelea dimidiata (Adams); Johannsen, 1943: 783 (combination); Wirth, 1965: 136 (distri- bution). Diagnosis. — Distinguished from all other Nearctic ClinoJielea by the two- spotted wings and dark brown tibiae; males with basal arms of aedeagus sep- arated. Female.— Wing length 2.54 (2.32-2.74, n = 5) mm; breadth 0.76 (0.68-0.87, n = 4) nmi. Head: Rrown. Antenna (Fig. 5a) slen- der; brown, proximal two-thirds of basal flagellomere pale; AP 17-10-9-9-9-10-10- 11-22-21-23-23-25 (n = 4); AR 1.35 (1.25-1.40, n - 4). Palpus with 3rd seg- ment slightly longer than 5th; PR 3.43 (3.17-3.80, n = 3). Mandible like that of C. biinaculata (Fig. 4c). 284 GREAT BASIN NATUKALIST Vol. 35, No. 3 c:ocii3Gr]c Fig. -5. Fenial thcca ri. sp.; ;i-c. ;: (Uinnhclca: a.e.p. C. dimidiata (Adams); b.d.h. C. usingeri Wirth; r.f.i. C. longi- tciinac; d-f. \v^ pattcfns; g. genitalia; h. i. sponnathecae. Thorax: Mesoiiotuni, scutelluni, post- scutellum dark brown. Legs (Fig. 5ej brown; yellow on fore coxa, tli.stal one- fourth of mid nnd Iniid coxae, trochanters, most of fore tcimu-. proximal five-sixths of mid femur, proximal half of hind fe- mur, and mid and hind 1st and IwA tarso- meres. Wing like that of ('. hima( uhita (Fig. 4bj. Halter stem pale; knol) white. Abdomen: Brown. Genitalia as in h'igure 5g with three small pairs of anteriorly directed, lightly scleroti/.ed arms. Sper- mathecae small, ovoid, subequal wdth short necks. M\Li.. -Siiuilar to the female with the following differences: smaller; flagellum (Milir-el\ brown; legs more diffused vsdth l)i(>\Mi. Genitalia shape and form like that of (\ hunacnUttd (Fig. 4f ) ; aedeagus as in j'igure (ie with basal arms apparently sep- arated, and an anteriorlv directed point on the anterior membrane. Sept. 197-5 ;k()(,\:\. wiktii: ci-.matoi' 285 Distribution. — Aj-i/oiin, !\'o\v Mexico. Utah (locality recoi-ds plotted in l''i,mire 7). IVPES. — Female lee lotApe, ) leniale paralectotyi^es, (iraiid C^aiiNon, Coconino Co., Arizona, C. F. Adams, (KU), here designated. Specimens ex.\mined. — From the fol- lowing localities: ARIZONA: Apachf Co.. Spniigeiville (Wirth). I male (P); Cocomno Co., Grand Can- yon (Adams, types). 4 females (KII) (P). NEW "MEXICO: Taos Co., Rio Grande (Wirth), 3 fe- males (P). UTAH: Cache Co., Hyrum (Grogan). I females (S); Box Elder Co.. Brigham City (Hardy, Stains. USU). I male, 1 female (SJ. 1 female (P); Weber Co.. Huntsville (Hardy, USU). 1 female (P); Wasatch Co.. Heber City (Dreisbach). I female (P). Discussion. — The syntype series was labeled "G. Zmii R.. Ariz., 7-27." which Adams ])ublished as "Grand Canon, Ari- zona" for the type locality. Clinohelca usingeri Wirth (Fig. 2b, d. h; 6f) Clinohelea usingeri Wirth, 1952: 209 (female; Cahfornia); Wirth. 1965: 136 (distribution). Diagnosis. — Distinguished from all other N(Nn-( li( (Tuiohelca by the two- spotled winn and the legs mainly yellow with distal li, df of hind femur and apices of tihiae hrowii: males with basal anus of aed(>agns intact. Fe.m.\li:.— Wing length 2.65 (2.50-2.74, 11 - 4) mm; breadth 0.76 (0.70-0.81, n 3) mm. Head: Brown. Antenna (Fig. 2b) mod- erately slender; basal one-half of proximal flagellomere lighter brown than remain- der of flagellum; AP 17-10-9-9-9-9-9-10- 23-22-22-22-20 n = 3); AR 1.33 ri.29- 1.35, n -- 3). Palpus with 3rd segment longer than 5th; PR 3.33 (3.17-3.67, n 3 ). Mandible like that of C. bimaculata (Fig. 4c). Thorax: Mesonotum, scutellum, post- scutellum dark brown. Legs (Fig. 2d) yellow; proximal one-third of fore coxa, most of mid and hind coxae, distal one- half of hind femur, femorotibial areas of mid and hind fore legs, apices of tibiae, and distal 4 tarsomeres brown. Wing like that of C. bimaculata (Fig. 4d). Halter })ale to whitish. Abdomen: Brown. Spermathecae (Fig. 2h) small, spheroid, subequal with short necks. Fig. 6. Aedeagi of male Clinohelea-. a, C. curriei; b, C. nubifera; c, C. pseudonubifera; d, C. bimaculata; e. C. dimidiata; f. C. usingeri. 286 (,REAT BASIN NATURALIST Vol. 35, No. 3 Male. — Similar to female with the fol- lowing differences: smaller; flatrellum entirely brown; femora and tibiae entirely brown. Genitalia shape and form like that of C. bimaculata (Fig. 4f ) ; aedeagus as in Figure 6f. Distribution. — Arizona, (California (locality records j)lotted in Figure 7). Type. — Holoty]:)e, female, Black Lake Canyon. San Luis Obispo Co., California. 22 August 1943, W. W. Wirth (Type no. 59949, USNM). Specimens examined. — From the fol- lowing localities: ARIZONA: Cochise Co., Sunnvside Canyon (Hardy, KU). 1 male. 3 females. CALIFORNIA: San Diego Co.. Live Oak Park (Melander) 1 female (P); San Luis Obispo Co., Black Lake Canyon (Wirth, type series). 1 male. 2 females (P), 3 females (S). Discussion. — Present records indicate that C. dimidiata is an inhabitant of the Great Basin and the LT]:)per Colorado Pla- teau, while C. usingeri is an inhabitant of the Mojave and Sonoran deserts. Further collecting is necessary to determine wheth- er the geographic ranges of these two closely related s])ecies oyerlap, or if they are separated by altitude. Cliuohclca longitheca Grogan and Wirth n. sp. (Fig. 5c, f, i) Diagnosis. — Distinguished from all other Nearctic CUnohelea by the very large, unequal, elongate, ellipsoid sper- mathecae, the legs mainly yellow with hind tibia and distal fourth of hind femur brown, and the two-spotted vyings. Female holotype. — Wing length 2.00 mm; breadth 0.61 mm. Head: Brown; frontoclypeus lighter brown. Antenna (Fig. 5c) slender, brown; AP 17-11-10-10-10-10-10-11-21- 21-21-20-20; AR 1.17. Palpus brown; 3rd segment longer than 5th; PR 3.11. Man- dible like that of C. bimaculata (Fig. 4c). Thorax: Mesonotum, scutellum, post- scutellum dark brownish black. Legs (Fig. 5f) yellow; most of mid and hind coxae, distal one-fourth of hind femur, hind tib- ia, and distal 3 tarsomeres brown; distal one-fourth of fore tibia light brown. Wing like that of C. bimaculata (Fig. 4d) with an infuscation over 1st radial cell and just before tip of costa. Halter light brown. Abdomen: Brown. Spermathecae (Fig. 5i) very large, unequal, elongate ellipsoid. Male. — Lin known. C. bimaculata C. dimidiata C. usi ngeri C. lonqitheca Fig. 7. North American locality records for Clinohrlra of the himaculata group. Sei)t. 1975 CROGAN. WIKTH: C1:HAT0P0G0NIDAE 287 Etymology. — The name longitheca is derived from the Latiii longus (long) and t/ieca (sac) and refers to the very large, elongate, ellipsoid sj^ermathecae that are characteristic of this sj)ecies. Variation. — The following characters were recorded for the single female to])o- t^•])e: wing length 1.97 mm; hreadth ().()! mm. AR'i.36. PR 2.89. CR 0.84. I'he general coloration is like that of the liolo- type. Distribution. — Florida (type locality plotted in Figure 7 ) . Types. — Female holotype, 1 female })aratype (S), A. & M. Riological Station, Blackwater River State Forest, Santa Rosa Co., Florida, 21 May 1971, G. B. Fairchild, hlack light trap (T\pe no. 66496, USNM). Literature Cited Ad.-mnis. C. F. 1903. Dipterological contributions. Kansas Univ. Sci. Bull. 2:21-47. Chan, K. L., and E. J. LeRoux. 1965. Descrip- tion of Forcipomyia (Neoforcipomyia) saun- dersi n. sp. and redescription of Forcipomyia (Neoforci])07nyia) ccjues (Johannson) (Dip- tera: Ceratopogonidae) with an account of the digestive and reproductive systems. Phy- toprotection 46: 74- 1 04. CoQUiLLETT. D. W. 1905. New Nematocerous Diptera from North America. Jour New York Ent. Soc. 13:56-69. DowNEs, J. A. 1960. Feeding and mating, and their interrelationshiji in the insectivorous (>eratoi)ogoninae (Diptera). Verh. XI Int. Kongr. Ent. Vienna 1:618. . 1971. The ecology of blood sucking Diptera: an evolutionary perspective. Pages 232-258 in A. M. Fallis, Ecology and physi- ology of parasites, a symposium. Univ. Toronto Press. JoHANNSEN. O. A. 1943. A generic synopsis of the Ceratopogonidae (Heleidae) of the Ameri- cas, a bibliography, and a list of the North American species. Ann. Ent. Soc. Amer. 36: 763-791. . 1952. Guide to the insects of Connecti- cut. Part 6. The Diptera or true flies. Fasc. 5. Midges and gnats. Heleidae (Cera- topogonidae). Bull. Conn. St. Geol. Nat. Hist. Surv. 80: 149-175. KiEFFER, .1. J. 1917. Chironomides d'Amerique conserves au Musee National Hongrois de Budapest. Budapest Magyar Nemzeti Muz., Ann. Hist. Nat. 15:292-364. LoKW, H. 1861. Diptera Americae septentriona- lis indigena. Centuria prima. Berlin Ent. Ztschr. 5:307-359. Mali.och, J. R. 1914. Notes on North Ameri- can Diptera. Bull. Illinois St. Lab. Nat. Hist. 10:213-243. . 1915. Tlie Chironomidae or midges of Illinois. Bull. Illinois St. Lab. Nat. Hist. 10: 275-543. Snodgrass, R. E. 1957. A revised interpreta- tion of the external reproductive organs of male insects. Smithson. Misc. Colls. 135:1-60. WiRTii, W. W. 1951. New species and records of Virginia Heleidae. Proc. Ent. Soc. Wash- ington 53:313-326. . ] 952. The Heleidae of California. Univ. California Publ. Ent. 9:95-266. . 1965. Family Ceratopogonidae (Helei- dae). Pages 121-142 in A. Stone et al., A catalog of the Diptera of America north of Mexico. U.S. Dept. Agr. Handbook 276. 1696 pp. BASIDOMYCETKS THAI DECAY JUNIPERS IN ARIZONA' R. L. Gilheitsoir and .) . F. Liiulsey- Abstract. — Twenty-seven spin ies dl \v(i in Arizona. A key to the spei i(>s. iies( i-i|)t ii vide.l. Pymfonirs dcmidnjjii. Daralra junipri licartwood in living tiecs. H yphodi'niut il< rnuing an.l Ii /. Pruia ticoUi ( :et('s are reported to decay junipers ;s of nnrroscopic characters are pro- id I'hrllinus texanus cause decay of I.iuds. is described as new. Species of the genus .liniipcru\ of th(^ Cupressaceae are among the most (oii- spicLious members of the flora of Arizona. Eive species occur in the state: Junipcrus (Icppeana Steud. ( aUigator juniper), Juni- pcrus scopulorum Sarg. (Rocky Mountain juniper), Juniperus monospernu/ ( En- gehn. ) Sarg. (one-seed juniper), ,/z//?//;(77/\ ostcospcrrna (Torr.) Little (Utah juniper), and Juniperus communis L. (common jimiper). The first four sj)ecies listed at- tain tree size and, with pinyon (Pinus cdulis Engelm.), occupy a large area of Arizona in the pinyon- juiii|)(n- woodland vegetation type from kiOO to 7,500 feet in elevation ( little. 100(S ) . I lo\^ever, juni- |)ers occiu' over a wider eknational range, from the u]iper desert grass vegetation type at 3,000 feet to the ponderosa })ine forest at approximately 8,500 feet. The amuial precijiitation in the piii\(>ii-|iuiiper woodland is only 12-20 int hes. most of which occurs in the summer rainy season in .July and August and during a winter [)eriod in December or .lainiar\ . During the remainder of the year it is usually dry in the zones where the junipers are found, and conditions are unfavorable for the development of wood-rotting fungi. A relatively small number f)f wood-rot- ting fungi have been foinid on junipers. This is probably due to the pres(>iK(^ of fungistatic compounds in juniper wood. Species of oaks and pines, connnonly as- sociated with junipers and expost'd to the same environmental ( oiiditions. are sid) strata for many wood-rotting fungi, .luni- pers are widely used in the Southwest for fence posts because of their high resis tance to decay and their ahiuidancc in most areas. .Junipers are aUd (oninionix used for fireplace wood. Hearti-dl in ]i\ing .Arizona juni[i('fs is maiidy attributed to loin- spfu ics in llic Polyporaceae. These are PyrofoiiK-^ dcini- dofjii. Dacdalea juniperina. /'on a ri//ise are the only host-fungus relation- shi|)s gi\en by Shaw that are not recorded for tli(> Southwest. The USDA host index lists Fomes roseus (Alb. et Schw. ex Fr.) ( ke. and ( Oniophora corrugis Burt on juni- per in .Arizona, but we have not seen \(Mu her sjx'c imens to support the records. uinal .\ill().N. ijxdm.'i : dkcay fungi 289 Key to 1^'iiiigi That I)(Hay .Iiiiiipers in Arizona 1. Hviiienojihore ( oiisistiiii^ of imited tubes 2 Hymenophore Miiooth. hydiiac (H)us. oi- lamellate — 16 2(1). Tissue brown, beconniiu pernianetitlv blackeiuMi in KOH solution .... _ Phellinus texanus Tissue white or pale colored, not permanently blackening in KOH so- lution 3 3(2). Basidiocarps centrally stipitate Polyporus arcularius Rasidioc arps s(^ssile or resujiinate 4 4(3). Basidiocar])s with brick red (onte.xt tissue Pyrofomes demidoffii Basidiocarps with context not brick red 5 5(4). Basidiospores thick walled, dextrinoid in Melzer's reagent Fames fraxinophilus Basidiospores thin w^allcd. not dextrinoid in Mel/.er's reagent 6 6(5). Basidiocarps always resu])inate 7 Basidiocarps sessile or reflexed 11 7(6). Pore surface often rose jiiid^; h^■phae siniple-sej)tate. no clanip comiec- tions ])resent .. Poria tarda Pore surface white to pale buff; generative hyphae with clamp connec- tions ; 8 8(7). Basidiospores subglobose to broadi^' ellii)soid Poria apachcriensis Basidiospores cylindric to allantoid 9 9(8). Basidiospores allantoid; cystidia present .-. Poria rimosa Basidiospores cylinrlric; c^■stidia not jiresent 10 10(9). Basidiospores 4-5.5 x 1.5-2 /an . Poria sinuosa Basidif)S]X)res gnlai'. tubes splitting to Fur in a lamellate hymenophore; tissue brown 15 15(14) . UppcM- surface^ usually distin(tl\- zoiiate with bright yellowish or red- dish brown zones; hymenophon^ strongl\- lamellate (ilocophyllurri sacpiarium Up[)er surlai e .i/onatc or indistin( tl\ /onale. dull brown; liNineno- pliore ponod to lamellate Cjlorophylliun trabeurn 1()(1). Basidiocarps stipit.ite; b\ nicnoplidro r.!(liall\ lamellate .._. Panus fulridus Basidiocar])s sessile or r(>supiiiate; hymenophore smooth to hydna- ceous 17 290 CHI-AT BASIN NATIRALIST Vol. 35, No. 3 17( 1()). Basidiocarps c ii|mlat('. up to 2 nun wide LachncUa alhoviolascens Basidiocarps ccsupiiialc 18 18(17K HynuMiophnrc smooth ... 19 H\nuMiophor(> liMlnac (M)ns - 25 19( 18). Basidia \erticall\ .septate - ....- Exidiopsis calcca Basirlia nonsoptatt' 20 20( 19). Hvnieiiial surtacc oin accous; hasidiosporcs thick walled, dextrinoid in Melzers reai^ciit ... Coniophora erctnop/nla H3'menial suita((> not olivaceous; basidiospores thin walled, negative or amyloid in Melzer's reagent 21 21 (20) . Basidios]X)res amyloid in Melzer's reagent; ac'anth()hy})hi(Ha and glo- eocystidia present 22 Basidios])ores negative in Melzer's reagent; acanthohyphidia and glo- eocystidia not j)resent 23 22(21 ). Basidiospores ().i-8 x 4-5 //m; liymenial surface becoming bluish gray .. Alcurodiscus lividocoeruleus Basidiospores 9.5-11.5 x 5.5-7 /-m; h^^^lenoph()re remaining pale buff .... Alcurodiscus cerrusatus 2^(21 ). Basidios])ores globose to subglobose; dendroh\ ])hidia i)resent Dendrothele incrustans Basidiospores cylindric to ellij)Soid; dendroh\phidia absent 24 24(23). Cystidia subulate, thin walled, (S-8 /an wide; aggregates of golden crys- talline material present .. Hyphoderrna pallidum Cystidia cylindric. with a rooted base, thick walled, 10-12 /im wide; golden crystalhne material absent Hyphoderrna descrticola 25(18). tlymenophorc pa|)illate. papillae with an amber-tolored, beadlike drop- let at the aj)ex .. Dacryobolus sudans Hymenophore distinctly hydnaceous with cylindric or flattened teeth 26 26 (25 J. Sagittate cysitha present Hyphodontia arguta Sagittate cysti(ha not present Hyphodontia spathulata J'lxiDiopsis CALCKA ( Pers. ) \V(dls, My- basidia to 55 /xm long; basidiospores (Fig. cologia 5 5(4): 548. 19f)l. Ic) cylindric, curved to slightly allantoid, 777r/,7;Wr/ r«/rr« Pers.. Mv.oi. Km. I: I -.1 182:2. smooth, hyahue, germinating by repe- tition, negative in Melzer's reagent, 14- Basidiocarps resupinate, thin. hard, arid- o() ^ (^.^ .,-^^1 wax^■. at first de^eloi)ing as small patches. ^ Exidiopsis calcea decays dead branches then coalescing to become widely ef used. ,,^- ^^,^/^^^^^ ^^d shrubs and occurs from cracking laterally to expose the substra- „^,. Sonoran Desert to high-elevation coni- tum; hymenial surface white to grayish ,•.,. ,,,^.,.^j^ j^ ^^ associated with a white white, smooth, shining; margin minutel\ .^ . fimbriate. sf)rnetim(>s abrupt; subicidar ., , . t- n /- r- i n -^ hvphae of tw.) tvpes. some narrow. \ "u. her specimen: E. R Canfield, / 1- branched and s,mious 1 1 ') „ni (ham '''• "'" ''H".^''!"!" juniper. Scotia Canyon, aseptate, others branched, with damp Hu.i.hu. a \ Its.. Cochise Co., AZ (ARIZ) . connections, 3-5 "in diain ( b"ig. la), tlu^se giving rise to basidia; basidia (Fig. Ibi Fachxi.lla alboviolascens (Alb. et with a basal clamp connection, hypo- Sc hw. ex Fr. i Fr., Sum. Veg. Scand. p. basidia subglobose to ellipsf)id, becoming ^65. 1849. longitudinallv septate and foiUMelled at /,^,^-.^, alhoriolasccus Alb. et Schw. e.x Fr., Syst. maturity. 12-1) x r)-20 /uii. the tour (^pi- Myc. 1: 'ifj. 1822. Sept. 1^)75 CII.HI'.RTSON. I.INDSI:Y: DF.CAY IM'NOI 291 Fig. 1. Rxidiopsis calcca (ERG 71-131), a, subinilar hyphae; h. hasidia; c. basidiospores. Basidiocai'ps cupulate. gregarif)iis. de- \plopiiig directly from the substratiiiii with a small patch of basal white my- ( (diiim. separate or crowded together, to 1 mm diam. sessile; outer surface co\ ered with a white, wooly-matted layer of hy- phae; hymenial surface pale pinkish brown, smooth, obscured on (h'ied spe- cimens by the inrolled margin; hyphal system dimitic, generative hyphae of the outer layer (Fig. 2a ) thick walled, hyaline, with occasional clamp coimections. 6-7 /un diam, some with an extremely narrow. caj)illarv lumen; skeletal hyphae (Fig. 2b) of outer layer thick walled, aseptate, to 7 ,"m diam; subhymenial hyphae moderately thick walled, with abundant clamp con- nections, 2-4 /ail diam; basidia (Fig. 2c ) broadly clavate, with thickened walls i()-<)() i,m long and 13-l(i /an at the apex, four-sterigmate, the sterigmata 2-3 /im in basal (ham; basidiospores (Fig. 2d) broadly ellipsoid, hyaline, negati\e in .Melzer\s reagent, 14-18 x 8.5- 10 /an. This fimgus is branches of a mu (les,>rl shrubs nnd brown rot. Voucher spe( imei i.r Ml (in d(Vi s(iiilb\\('st(n- i.ilcil with RLC; 10193. on o seed juniper. Black Oak Cemetery, (^anelr R.L. Santa Cruz Co.. A/, f ART/ ) . ■I Cnl- ( ioNIOI'lIOHA l.KI.MOl' III.A 1 .inds. bcrts.. iM\cotaxon 1: -SI). 1 0 7'). Basifhocarj)s Irai^il '. eas \\ se[ occurring in small pa tclics ( r of 111- cm; hymenal surbn ' srno< dh. \n Brownish Olixc from inass('( basid ited. as they mature; hymenial layer soft and flo( ( os(> oyer a white arachnoid subiculum; margin with fine, white mycelial strands radiating from subiculum; subicular hy- phae simple-septate, hyaline, often lightly incrusted, some (Fig. 3a) thin walled, 2-5 /an diam, others thin to thick walled (Fig. 3b), to 10 /im diam; cystidia none; basidia (Fig. 3c) utriform to clavate, usually sin- uous, 40-(S() X 6-10 /im, four-sterigmate, the sterigmata to 7 /an long; basidiospores (Fig. 3d) brownish olive in mass, pale yellow in KOH, cyanophilous, dextrinoid in Melzer's reagent, thick walled, ellipsoid to subglobose, 9-11 x 5-8.5 /xm, with an apical germ })ore and prominent peglike apiculus. Coniophora cremophila has also been found on several Sonoran Desert trees and shrubs and is associated with a brown rot. Voucher specimen: RLG 7400. on one- seed juniper. Gallinas Mts., Lhicoln Co., NM (ARIZ). Aij:i!rodiscus CKRUssATUs (Bres.) Hoelin. et Litsch., K. Acad. Wiss. Wien Math.- Nal. Kl. Sitzungsb. 116: 807. 1907. Corticium cerussalum Bres., Fung. Trid. II. 37. 1892. il I. Fig 2. Lac/tnrlla alboviolascens (RLG 101931, a. grnerative tiyphae; b. skeletal hypha; . hasidia; d. basidiospores. 292 (iHi:.\T BASIN NATlKALIsT Vol. 35, No. 3 Biisidiocarps in small |) oily ((Hiteiits. some moiiililorm. 10-70 x 10-1 ^ //in; weakh positixc in sulphoheir/.- aldehyd(>; acalltlloph^ ses ( Mg. l-c ) api- cally thick walled, thin walled at the base, cylindric. 30-55 x >l /nn; with a basal clamj) connection; mature hasidia (Fig. 4d) four-sterigmatc. 5-9 x 45-hO /an; basidiospores ( Fig. 4e) broadly cylindric to ellii)soid. thin walled, smooth, amyloid in -Melzer's reagent, inostly collajised or fragmenterl. 9-11 x 5-7 /an. AIriir()(lis( us ct-russa/us grows on dead branch(>s of other shruhs and tre(>s in the Southwest. It is associated with a white rot. Vcnu htu" s])ecimen: RFG 8551. on one- seed junij^er. Cliiricahna Nat. Monnment. Chiricahna Mts., Cochise Co.. AZ fARIZ). ALF.riu)i)isc:rs li\ nj()f:()KKrM:is ( Karst. ) Femke. Can. .T. Rot. 12: 252. 1964. Corliciuni UvidocoerulcuTn Karst., Not. Siilsk. Faun, et IHor. Fonn. Forli. 9: 570. 1868. Rasidiocarj)s resupinate. originating as small, separate ])atclies, then becoming confluent and widely effused, w-axy; mar- gin abrupt and fertile; hymenial surface cream colored or Pale Pinkish Buff to blu- ish gra}' (Light Neutral Gray to Dark Plumbeus), often rimose with age; hyphal system monomitic; subicular hyphae (Fig. 5a) thin walled, with clamp connections, 3-6//mdiam; gloeocvstidia (Fig. 5b) abun- dant, positive in sulphobenzaldehyde, em- bedded or slightly projecting, some moni- liform or mammillate, 20-90 x 6-F3 /i.m; acanthophyses (Fig. 5c ) abundant, acule- ate o\(>r the terminal [)ortioii. thin to thick walled at the ajKw. with a basal clamp connection. 15-^)0 x ^)-7 /an; hasidia (Fig. 5d) clavate. -10-50 x 10-11 /an. ste- rigmata not seen; basidiospores ( |'"ig. 5e) broadly ( ylindrir to ellipsoid. Inaline. smooth, amyloid in Mel/er's reagent, thin walled and collapsing readily. f)-S x ').5-5 /'.m. This fungus occurs thronghonl the Rocky Mountain region on niaii\ (onifers. It is associated with ;i \\hil(' i-ot. \^)n(her specimens: RLG 10777, on al- ligator iunij)er, Turkey Creek, Chiricahna \lts., Cochise Co., AZ; ERC 71-234, on alligator juniper. General Hitchcock Pic- nic Area, Santa Catalina Mts., Pima Co., AZ (ARIZ). Di xDHorin-.LK ijxcRusTANs ( Lcmke) Hike. !\>rso()nia ^: MM^ 1965. ■Ucurocorliciuin incrus/ans Lcmke, Can. J. Bot. 42: 7' Basidi ')f)4. |)s small, usually in patches tip to 5 mm wide, sometimes confluent; margin abru])t, fertile; hymenial surface smooth, \^hite to Cartridge Buff or cine- reous, rimose on drying; subicular hj'phae ( Fig. ()a ) slender, with frequent branch- ing, with inconspicuous clamp connections, 1-2.5 /an diani; cystidia absent; hyaline oias_ Fig. 3. Coniophora eremophila (RLG 7400), a. narrow subicular hyphae; b, broacl subicular li>pha; c. basidia; d. basidiospores. Fig. 4. Aleurodiscus cerrusatus (RLG 8551), a, subicular hyphae; b, gloeocj'Stidia; c, acantho- physes; d, basidia; e. basidiospores. Fig. 5. Aleurodiscus Uvidocoeruleus (RLG 10777), a, subicular hypha; b, gloeocystidia; c, ai anthophyses; d, immatui-e -basidia; e, basidio- Sept. 1075 (;ir,Bi:HTs()%'. NDSl.^ : ni.CAl I'UNGI 293 (londrohy|)lii(lia i Fig. ()1) i ahiitidaiil iti h^ menial region, ultimate hiaiic lies | /,iii (liam or less; hasidia i I''ig. fxl i (la\at(> lo cvlindric. some snmiIIcii al the hase. de- veloping from imbedded hasidioles (Fig. 6c), 30-60 X 8-10 /-ni. four-storigniate, with a basal clamp connection; basidiospores (Fig. 6e) hyaline, smooth, negative in Melzer's reagent, globose to suhglohose. 9-11 X 8-9.5 /-m. sometimes adhofing in groups of two to four. Dendrotlicle 'uKrustdiis aj)parently util- izes bark tissue as a substratum. It also occurs on bark of se^•eral other southern Arizona trees. inchuHng oaks and Arizona madroiK* [Arhutiis arlznnica ( Cjray ) Sarg.). Lendse ( l<)6 1i reports this fungus (as Aleurocorticium inr rustans } from sev- eral localities in the Pacific Northwest, but it has not been rejiorted previously from the Southwest. Voucher spe Fl. 9: 12- .riTNiPERiNA Murr., N. Amer. 1908. Fig. 14. Coriolellus sepium (RLG 10129). a, contextual generative liyphae; b. contextual skele- tal hyphae; c. basidia; d. l)asidiospoies. tomentose, azonate, shallowl\' sukate, smooth or slightly rugose; tiuirgiti con- coloroiis, rouiidecl, fertile or narrowly sterile below; pore surface Cinnaniou Buff to Buckthorn Brown, the pores cir- cular to angular, 2-3 per mm, the dissep- iments thick, entire, eventually becoming lacerate; context ivory, azonate, corky, to 4 mm thick; the tube layer concolor- ous and continuous with context, to 1 cm thick; odor sometimes fragrant; hyphal system dimitic; contextual generative hy- [)hae (Fig. 14a) hyaline in KOH, mostly thin walled, rareh' branched, with abun- dant clamp connections, 2.5-5.5 /ini diami; contextual skeletal hyphae (Fig. 14b) thick walled, aseptate, 3-5.5 n.m diam; tramal hyphae similar; cystidia none; basidia (Fig. 14c) clavate, four-sterigmate, 20-25 X 8-10 /-,m; basidiospores (Fig. 14d) hyaline, negati^■e in Melzer's reagent, cy- lindric, 9.5-14 x 3-4 /an. Coriolellus scpiu/n is associated with a brown cubital rot with thin, arachnoid mycelial felts. Voucher sjx'cinuMis: RL( i 72()l.()n al- ligator juniper, Rucker Canytjii. (diirica- hua Mts., Cochise Co., AZ;^RLG 10045, on juniper post, Washington Camp. Pata- gonia Mts.. Santa Cruz Co.. A/; RI.G 10048, on alligator JLuiipcr. Montezuma Pass, Huachuca Mts.. (^odiise Co.. AZ; RI/i 10198, on juniper post, Sunn\'side, Cochise Co., A/; mXr 10129, on alligator juniper, Parker Canvon Lake, C>)chise Co., AZ; RLG 108 38, on alligator juniper, Indian Creek, Animas Mts., Hidalgo Co., NM; ERC 70-L on alliuator juniper. Rust- ler Park, Chiri(ahii,i Mtv. Co. Ium" Co. AZ; ERC 71-129. on alligator jumper-. Scotia Canyon, Ilii,i(lui(a Mts.. {d.jiisc Co., AZ (ARIZ ) Basidiocarps annual or perennial, ef- fused-reflexed or resupinate; pilei solitary or imbricate, f)fteii laterally fused, to 5 X 10 x 9 cm; surface of the pileus weath- ering Light Buff to Cinnamon Buff, to gray or blackish, indistinctly zonate, be- coming glabrous and incrusted; pore sur- face Light Buff, rough, pores large, dae- daloid, often more than 1 mm diam; dis- sepiments thick, entire, splitting with age to form lamellae or spines; context yel- lowish ivory, soft-corky, faintly zonate, to 2 cm thick; tube layers concolorous and ( ontinuous with context, to 6 cm long; hvjihal system dimitic; contextual gener- ative hyphae (Fig. 15a) thin walled, with rare branching, with abundant clamp con- nections, 3-fi //m diam; contextual skeletal hvphae (Fig. 15b) hyaline in KOH, thick walled, with rare branching, aseptate, 3-7 //m diam; tramal hyphae similar; cystidia none; basidia (Fig. 15c) clavate, 20-25 x 6-7 nin. four-sterigmate; basidiospores K. !■ Dacdalca juniper ina (JPL 328, RLG IDfjOh. a. contextual generative liyphae: b, con- icxiM.il skeletal hvphae; c. basidia; d. basidiospores. Fig. 16. Fomes fraxinophilus (ERC 71-25), a. subicular generative hyphae; b, subicular skele- tal hyphae; c. basidia; d, basidiospores. 298 GREAT BASIN NATURALIST Vol. 35, No. 3 (Fig. 15d) hyaline, negati-ve in Melzer's reagent, smooth, cylinclric, 6.5-9 x 2.5- 3.5 /.(.m. Daedaica jiniipcriiia is associated with a brown cubical heartrot of living juni- pers. Thick, buff colored mycelial felts develop in shrinkage cracks of decayed wood. Voucher specimens: RLG 6940, on one- seed juniper. Salt River Canyon, Gila Co., AZ; RLG 9945 and ERC 7^1-26, on one- seed juniper, Sycamore Canyon, Atascosa Mts., Santa Cruz Co., AZ; P. D. Keener, on alligator juniper, Mingus Mt., Yavapai Co., AZ; RLG 10604, on Utah juniper, Seegmuller Mt., Mohave Co., AZ; ,IPL 328, on alligator juniper, Rucker Cannon, Chiricahua Mts., Cochise Co., AZ; KJM 335, on one-seed juniper. Canyon del Oro, Santa Catalina Mts., Pinal Co., AZ (ARIZ). FoMEs FRAXiNOPHiLus (Pk.) Cke., Gre- villea 15: 51. 1886. Polyporus fraxinophilus Pk.. Bot. Gaz. 7: 43-44. ■l882. Basidiocarps perennial, resupinate, ef- fused to 6 cm; pore surface Pale Ochrac- eous Buff to Pinkish Buff, the pores 3-4 per mm; dissepiments thick, minutely to- mentose; margin narrowly sterile, 0.5 mm wide. Light Buff; subiculum ])ale buff, to 1.5 mm thick; hyphal system dimitic; generative hyphae (Fig. 16a) in subiculum thin walled, 2-3 /mi diam, with inconspicuous clamp connections, these more apparent in marginal tissue; subi- cular skeletal hyphae (Fig. 16b) hyaline, moderately thick walled to thick walled, with occasional branching, aseptate or rarely with simple septa, 2.5-5 /'.m diam; tramal hyphae similar to subicular skele- tal hyphae but mosth' 2-3 /mi diam; cys- tidia none; basidia (Fig. 16c) broadly clavate, four-sterigmate, 20-25 x 9-11 /'m; basidiospores (Fig. 16d) broadly ellipsoid to subglobose, truncate at the apex, hya- line, dextrinoid in Melzer's reagent, 9-10.5 x 6.5-8 jim, thick walled at maturity with a germ pore at the truncate apex. Fomes fraxinophilus causes a wbit(» heartrot of living trees and is common in southern Arizona on Frnxinus rclufina Torr. (Arizona ash). Small resupinate basidiocarps are also commonly found on ash. Clamp corniections of F. frarinoph- ilus are abundant and conspicuous in (h- karyotic cultures of the fungus but are difficult to discern in basidiocarp tissue. Voucher specimens: ERC 71-25, on one-seed juniper. Sycamore Canyon, Atas- cosa Mts., Santa Cruz Co., AZ; J. L. Lowe 9091, on one-seed juniper, Silver Creek Rd., Chiricahua Mts., Cochise Co., AZ (ARIZ). Gloeophyllum saepiarium (Wulf. ex Fr.) Karst., Finl. Hattsv. 2: 80. 1879. Daedalea sepiaria Wulf. e.v. Fr., Syst. Myc. 1: 333. 1821. Basidiocarps annual, effused-reflexed, sessile, or occasionally resupinate; upper surface Warm Sepia to Bister or blacken- ing, hirsute to coarsely strigose, concen- trically zonate; lower surface Sudan Brown to Amber Brown, pores present at the margin of some specimens, but tubes typically splitting to form a radially lamellate hymenophore; context Honey Yellow to Clay Color, darkening to Sepia, azonate, up to 4 mm thick; hyphal system dimitic; contextual generative hyphae (Fig. 17a) hyaline, thin walled, with abundant clamp connections, 3-5 /im diam; contextual skeletal hyphae (Fig. 17b) thick walled, pale yellowish brown, asep- tate, with occasional branching, 2.5-5 /im diam; cystidia (Fig. 17c) thin to thick walled, cylindric, not incrusted, to 85 /xm long and 2.5-6 /.i.m diam; basidia (Fig. 17d) narrowly clavate, with a greatly elongated base, 60-80 x 7-8 /mi; basidio- spores hyaline, smooth, cylindric, slightly curved, negati\e in Melzer's reagent, 8.5- 11 X 3-4 /mi. G/oeophyiluni saepiarium causes a brown cubical rot of conifers and hard- woods in Arizona but is rarely found on junijier. Voucher sj)ecimen: RLG 10918, on al- ligator juniper, Gardner Canyon, Santa Rita Mts., Santa Cruz Co., AZ (ARIZ). Gloeophyllum trabeum (Pers. ex Fr.) Murr., N. Amer. Fl. 9: 129. 1908. Daedalea Irabea Pers. ex Fr., Syst. Myc. 1: 335. 1821. Basidiocarps annual, sessile, effused-re- flexed or occasionally resupinate in early stages of do\elopment; pilei dimidiate to elongated, often imbricate and confluent; upper surface yellowish - brown, weath- ering to tan or grayish, tomentose and be- Sept. 1975 GII.BF.RTSON. LTiyDSK^': DKCAY FUNGI 299 Fig. 17. Gloephyllurn sacpiariutn ( RIXi 10918), a, contextual generative hyphae; b. con- textual skeletal hyphae; c, cystidia; d. basidia; e, basidiospores. Fig. 18. Gloephyllurn Irabeum ( RLCi 10128), a, contextual generative hyphae; b. con- textual skeletal hypliae; c, cystidia; d, basidia; e, basidiospores. coming glabrous. f;iiiitl\ zoiiate; lower surface i)ale brownish, poroid at first and in some specimens remaining poroid witli ])ores 1-^ j)er nmi. radially elongated, in others becoming ratlially lamellate l)\ splitting of dissepiments; (ontext pale brown, soft-felt}, a/ojiate, up to 12 nnn thick; tube layer or lamellae ]iale brown, up to 4 mm thick; hyphal system diniitic; contextual generative hyjihae (Fig. IHa) hyaline, thin walled, with abundant (lam]) connections, 2-15 nn\ diani; contextual skeletal hy])hae (Fig. IHb) pale \ellowish brown, thick walled, aseptate, with rare branching, 3-5.5 /im diam; tramal hyphae similar; cystidia (Fig. 18c) cylindric, thin walled, 35-50 x 3-5 /an; basidia (Fig. 18d) clavate, foi.u--sterigmate, 30-45 x 5-7 //.m; basidiospores (Fig. 18e) tylindric . h\ aline, smooth, negative in Mc'l/cr's icagent, 7.5-9 X 3-15 nm. Glocophyllum Iralx-iini is a^so( iated with ».. AZ ARIZ). PoLYPORUs ARCuLARH's Batscli ex Fr., Syst. Myc. L 342. 1821. Basidiocarps annual, centrally stii)itate; pilei circular, solitary, uj) to 2.5 cm diam and 0.3 cm thick; surface of the pileus straw colored to dark brown, a/.onate, glabrous, smooth to rugose; margin ciliate. acute, sterile below; stripe central, con- colorous with pileus, glabrous, up to 3.5 cm long and 0.4 cm thick; pore surface cream colored to buff, dull, rough, the pores large, he.xagonal, radially aligned, 1-2 per mm, the dissepiments' thin, be- coming lacerate; context whitish to buff, azonate, tough, less than 1 nun thick; tube layer concolorous and continuous with context, up to 2 mm thick; hyphal sys- tem dimitic; generative hyphae (Fig. 20b) hyaline in KOH, thin walled, often branched, with abundant clamp connec- tions, 2.5-5 ,am diam; skeletal hvphae (Fig. 20c, Fig. 20d) thick walled,' asep- tate, with occasional branching, 2-11 nxn diam; tramal hyphae similar, not readily separable; hyphae on pileus surface (Fig. 20a) slender, thin walled, with clamp connections, 1-1.5 /-.m diam; basidia (Fig. Fig. 20. Polyporus arcularim i.IPI, 278 and HLG 790^ j, a. livjihae from ])il('us siirfact'; h. generative hvpliac; <. broad skeletal livi)hao; d. narrow skeletal lispliar. ,.. Iwisidui; I. b.isidin S|l<)|cs 20e) four-sterigmate, clavate, 25-35 x 5-6 /an; cystidia none; basidiospores (Fig. 2()f) h^'aline, negative in Melzer's re- agent, smooth, cylindric, straight or slight- ly curved, 7-9 x 2.5-3.5 /tm. Poly par us circular ius causes a white rot and is especially common on dead oak wood throughout the oak woodland vege- tation zone in southern Arizona. It is oc- casionall}' found on other substrata. This report is based on a field observation of P. arcularius on one-seed juniper at Sunnyside, Iluachuca Mts., Cochise Co., AZ by RLG. PoRiA APACHKRiENsis Gilberts, et Canf., Mycologia 65: 1117. 1973. Basidiocarps annual, effused up to 10 cm. adnate, soft-fibrous; pore surface white to Cream Color or Light Buff; sterile margin tomentose; pores circular to daedaloici, up to 1 nxn diam, mostly 2-3 per mm; tube layer soft-fibrous, white to cream colored, up to 2 mm thick; subi- culiun soft, white, less than 0.5 mm thick; hyphal system monomitic; subicular hy- phae (Fig. 21 a j with abundant clamp con- nections, thin to moderately thick walled, with occasional branching, 2-4 //.m diam; tramal hyphae similar, incrusted at dis- sei:)iment edges; cystidia thin walled, of two types, some acicular or cylindric (Fig. 21b), smooth to lightly incrusted, 45-60 X 2.5-5 ,um, others capitate (Fig. 21c), 3-5 /un diam with swollen apex up to 8 /im diam, 40-50 /tm long; basidia (Fig. 21 d) with a median constriction, four-steriginate, 18-30 x 6-7 /xm; basidio- spores (Fig. 21 e) hyaline, smooth, nega- \\\i' in MeTzer's reagent, broadly ellipsoid to subglobose. 5-6.5 x 1.5-5.5 //in. Poria apachcricusis occurs on a number of Sonoran Desert plants, including the Saguaro cactus, Carnegiea gigantea (En- gelin. ) Britt. et Rose (Lindsey and Gilbert- son. 1975), and causes a white rot. It I'ruitb aroinid the base of recently killed junipers at the ground line, suggesting it may cause a root rot in Hving trees. Vou.her spc(ini(Mis: RLG 10047 and l"',KC LSI. (in alligator juniper, Monte- /.inna l*)chise Co., AZ (ARIZ). PoHi A 1 I iu).\ Long et Baxter, Papers Mich. Acad. S(i. 25: 1 19. 1()40. Basidiix ;ir[is perennial, becoming wide- \\ fffuscil; [lore surface (cartridge Buff to l^ndsish P.utr nr I'ale Orange Yellow, the Sr,.l 501 N u Fig. 21. Porid iiimchericfisis iJPL 407 and 409). a. sul)i(ular liyphae; b. cvliiidiic rvstidia; c. capitate cystidia; d. basidia; e. basidiospoies. pores circular to .lugular. 4-() per tnni; margin abrupt, fertile or narrowly sterile, blackening with age; subiculum less than 1 nnn thick, cream colored to jjale buff; tube layers stratified, single layers u\) to 1 mm thick; hyphal system dimitic; su- bicular generative hyphae (Fig. 22a) in- conspicuous, thin walled, with clamp con- nections. 2-3 ii.m diam; subicular skeletal hyphae (Fig. 22b) hyaline, thick walled, aspetate, with rare branching, 5-5 //.m diam; tramal hyphae similar; fusoid cys- tidioles (Fig. 22c ) present, barely pro- jecting, 11-18 .X 5.5-7 /an; basidia (Fig. 22d), broadly clayate, four-sterigmate. 14-18 X 8-9 /(.m; basidiospores (Fig. 22e ) cylindric, subfusiform, hyaline, smooth, negative in Melzer's reagent. 8-10 x 5-4 ,um. Poria ferox causes a brown c ubical rot with consjHcuous cream colored to pale buff mycelial felts in the shrinkage cracks. Voucher specimens: RLG 10126. on al- ligator iuniper. Parker CauAon, Huacliuca Mts., Santa Cruz Co.. AZ; RLG 10817 and 10822, on alligator juniper, Indian Creek, Animas Mts.. Hidalgo Count^■. NM; RLG 10127. on alligator juniper. Parker Can^•oll Lake. Cochisf Ca).. AZ ( ARIZ ) . PoRi.\ RiMos.y ATurr.. Mycologia 12: 91. 1920. Basidiocarps perennial, sometimes de- veloping as scattered j)atclies on a cottony mycelial mat f)n the surface of the sub- stratum, often becoming rinios(> with age; pore surface Light Ochraceous Ruff to Cinnamon Buff, pores 5-7 jier mm. angu- lar; hyphal pegs present; dissei)iments at first appearing tottientose under a 30X lens, llni Is. Ixm oming thin and splitting with age; lube layer ivory to straw colored, up to 2 mm thick; context con- colorous with tubes, soft, fibrous, up to 0.5 nun tln( k. taste mild; hy{)hal system di- mitic; subic ular- gcMierative hyphae (Fig. 2 5a) thin wallecl, with abundant clamp c ounce tions. often branched, 2-5 yxn diam; subicular skeletal hyphae (Fig. 23b) thick walled, apparently nonseptate, 2-3 /'m diam; tramal hyj)hae predominantly thin walled, with clamp connections, 2-3 ;:x\\ diam; cystidia (Fig. 23c) scarce to conmion, not incrusted, thin walled, hy- phoid, up to 2.5 /an diam and projecting up to 25 N.m beyond the hymenium; ba- sidia (Fig. 25d) clavate, four-sterigmate, 10-13 X 4-6 ,"m; basidiospores (Fig. 23e) hyaline, smooth, negative in Melzer's re- agent, allantoid, 4-5.5 x 1-2 ,um. Poria riifiosa causes a white heartrot of living trees and is also found on dead standing and fallen junipers. In the ad- vanced stages the rot is characterized by small em])ty pockets. It has a wide dis- tribution in western North America (Gil- bertson, 1961). Voucher sjiecimens: RLG 6961, on al- ligator juniper. Sponsellor Lake Rd., Nava- jo Co.. AZ; RLG 6963, on one-seed juniper, between Show Low and Snow- flake, Navajo Co., AZ; RLG 7560, on cme-seed juniper, Stoneman Lake Rd., Co- conino Co.. AZ; K. D. Butler, on alligator juniper, Mingus Mt., Yavapai Co., AZ; .ILL 9106, on one-seed juniper. Silver Creek, near Portal, Cochise Co., AZ; RLG 10605, on LTtah juniper, Seegmuller Mt. area near Wolf Hole, Mohave Co., AZ (ARIZ). PoRi.'^ siNuos.^ (Fr.) Cke., Grevillea 14: 115. 1886. Pnlrporus sinuosus Fr.. Syst. Myc. 1: 381. 1821. Basidiocarps annual, often widely ef- fused, tough, corky, easily separable, taste resinously bitter; margin fertile or narrowly st(>rile. Light Buff, soft, fim- briate, to 1 nmi wide; pore surface Cream Color or in Melzer's re- agent, 4-6 X 1.5 2 i>n\. Poria si nuosa causes i\ brown cubic al rol of conifer logs and slash. It is clisiriljni,.,! throughout western conilcrons iof(vsts. Voucher speciiiieii: II. M. Burdsall 11 ^1 on Juuipcrus s}).. Valoiu ia (^o.. Cibola Nat. Forest Rd. No. HSO near ( Irani s. NM (ARIZj. Porta ta]U)a (Berk.) Cke., Grevillea 14: 109. 1S8(). Polyporus tardus Berk., London J. Bot. 4: 56. 1845. Basidiocarps annual, usually adnate, pore surface rose pink to cream, usually drying Pinkish Buff to Light Buff; tubes originating as isolated cupules and then iniiting; pores 3-5 per mm; sterile margin usually rather wide, thinning out; context white to cream, soft, thin, Iwphal system monomitic; subicular hyphae (Fig. 25a) hyaline, thin walled, simple-septate, oc- casionally ampullate at the septa, fre- quently branched at right angles, some with crystalline incrustation, 2.5-6 jxva diam; tramal hyphae similar; cystidia none; basidia (Fig. 25b) clavate, four- sterigmate, 15-20 x 4-5 ixva; basidiospores (Fig. 25c) oblong to cylindric-ellipsoid, hyaline, smooth, negative in Melzer's reagent, 4-5 x 2-2.5 /im. Poria tarda is a common fungus in southern Arizona and has been found on dead wood of many trees and large shrubs from the Sonoran Desert up to the pond- erosa pine forest. It causes a white rot. Voucher specimens: RLG 10135, on al- ligator juniper, Sunnyside, Huachuca Mts.. Cochise Co., AZ; RLG 10197, on al- ligator iimiper, Canelo Rd., Santa Cruz Co., AZ; FRC 71-118, on alligator juniper, Suimyside, Huachuca Mts., AZ (ARIZ). Pyrofomes demidoffii (Lev.) Kotl. et Pouz., Repert. nov. Spec. Regn. veg. 69: 140. 1961. Polyporus demidoffii Lev. in Demidoff, Voy. Russ. Merid. 2: 92. 1842. Basidiocarps ])erennial, sessile, solitary, tmgulate, often becoming columnar, to 15 cm wide, 7 cm thick, and 10 cm high; up- per surface brownish and tomentose in young specimens, becoming blackened and rimose with age, concentrically sul- cate; margin rounded. Warm Buff to Ochraceous Buff, finely tomentose to blackened and rimose in older specimens; pore surface^ Light Ochraceous Buff to Ocbracc'ons I'uif. smooth, the pores i-oinidc'(l. J ) |)C'i- mm; dissepiments thick, entire; (oiUc.xl Orange Ciimamon to Cin- namon Ridons. woody, azonate; tube layers Ochraceous Buff to Antimony Yel- low at first, e\entually becoming filled with mycelium and concolorous with con- Sept. 1975 (;ilbi:kts()in. li;\'1) i)i;(; x^ I'l' ;\(; 303 'ig. 26. Pyrofotncs demuhffii {«L(i lUbOO). a. liypliae from mycelial felts; b, contextual skeletal hyphae; c, much-branched skeletal hyphae from trama; d. contextual generative hvphap; e. fiisoul fvstidioles: f. basidia; g. basidiospores tt^xt, iiidisliiK tl\- stiiitifii'd. each hiA or to 7 mm tliick; hy|)hal system dimitic; con textual generative hy])hae (Fig. 26d) dil ficult to discern, thin walled, hy^aline in KOH and Melzer',s reagent, with clamp connections, 2.5-4 /an diam; contextual skeletal hyphae ( Fig. 26b) moderately thick walled, rarely septate, with rare branching, pale brownish in KOH and dextrinoid in mass in Melzer's reagent, 3-5.5 /an diam; tramal tissue similar but with some much-branched skeletal h\phae (Fig. 26c), 2-5 /'m diam; tramal tissue distinctly dextrinoid in Melzer's reagent; hyphae of mycelial felts (Fig. 26a) in wood frequently branched, some thick walled, aseptate. 1-^ /an diam. others thin walled. simple-sej)la1e or with (xca^ional clamp connections. 1.5-15 /an diam; in- conspicuous fusoid cystidiolirs (Fig. 26e) in hymenium. thin walled, not incrusted, 2()-30 X 3-5 /an; basidia (Fig. 26f) broad I v clavate from a narrow base, four-sterig- mate. with a basal (lamp ( otmection. 22- 35 X 8-10 /an; basidiospores i Fi-. 26g i pale brownisli. slightly dextrinoid in \lel zer's reagent. ()\oid lo IxoiuJlx ellipsoid or tiiore elongated, angular, thick walled, most truncate at apex with an inconspicu- ous germ ])ore, 6-12 x 5-7 /xm. Pyrojoiurs dcinidoffii is probably the most important heartrot fungus in western junipers. It ( anses a white rot with abun- dant my((>lial lelts in the decayed wood. It has been referred to as Fomes juniperi- nus (yon Schrenk) Sacc. et Syd. in most American literature. Voucher specimens: KLG 6960, on one- seed juniper, between Show Low and Snowflake, Navajo Co., AZ; RLG 7384 and 7815, and ERC 71-28 and 71-326, on one-seed juniper, Sycamore Canyon, Ata- scosa Mts., Santa Cruz Co., AZ; RLG 7559, Stoneman Lake Rd., 7 mi E of High- way 79, Coconino Co., AZ; RLG 7562, on one-seed juniper, Dry Creek, 6 mi W of Sedona, Coconino Co., AZ; RLG 7890, on Utah juniper. South Rim, Grand Canyon Nat. Park Coconino Co.. AZ; RLG 9875, on LTtah juniper, Mt. Trumbull, Mohave Co., AZ; JLL 9070, on one-seed juniper. Silver Creek, Chiricahua Mts., Cochise Co., AZ ( ARLZ ) . 56. 1900. Rasidi.Harps to gregarious; diam; npper >ii iiamou Ikifl to fibrillar scales; to Ruffy Rrowi Rres., Fung. Trid. XL p. (('ntrall\ stipitate, single jjileus ( irmlar, 0.5-3 cm ilacc pale brownish (Cin- (>lay Color) with radial sti})e Light Ruff at apex 1 at the base, glabrous to minutely pubescent or scaly% to 3 mm diam and 2 cm long; gills pinkish cream to pale buff when dried, distant, free to adnate, eflges siimous and distinctly granu- lose under a 30X lens; contextual h}'phae variable, some (Fig. 27a and b) simple- septate, with occasional branching, thin to slighth' thick walled. 2.5-6 /an diam, others (Fig. 27c I very thick walled to almost solid, aseptate. rarely branched, 5-9 /im diam; |)letu-oc\ stidia (Fig. 27d) fusoid, barely projecting, 35-60 x 6-8 /an; cheilo- ( ystidia similar; basidia (Fig. 27e) clav- ate, four-sterigmate, simple-septate at base, 38-60 x 9-13 /-m; basidiospores (Fig. 2; li) br( )adl^ r c\ dindric, si lightly curved. ii; valine. sm( .olh'. negative in Melzer's re- agent. 11 2-16 X 5. 5-7 /an. Pr/rn/s fu/, ' vV/?/. s causes a browat cubical r-( )t and i onitt ion finigii< ; on dead stand- ii igand ! la Me n ini :ii[i(^rs and also on juniper fe ■nee po sts 1 n soi itliern Arizona. 304 GREAT BASIN NATURALIST Vol. 35, No. 3 Fig. 27. Panus fulridus (ERC 71-1^2 aiul 71-158), a, thin-walled contextual livphao; b. slightly thick-walled contextual liyphae; c, thick walled to solid contextual liyphae; d. fusoid pl(;urocystidia; e. basidia: f. basidiosjiores. Voucher specimens: ERC 71-1 ^2, on al- ligator juniper, Scotia Canyon, Huachiua Mts., Cochise Co.. AZ; RTX; 100 Ik on junij)er Nmkc Pataooiiia \I|> 10258, on jiii Canyon, Ala^ AZVARIZ). Lr po^l. VVashinoion (Jamp, . Santa Cm/. Co.. AZ; RLG i\)cv fence post. Sycamore nsa Mt'^.. Santa Cruz Co., :r.\ti !RK C I ted Index of plant diseases in s. TT.S. Dept. Agr. Handb. Anony.aious. 1960. the United Sta No. 163. VJl p. Eriksson, .!.. \xi) L. Rvvakdf.n. 1973. The Corticiaceae of North Europe. Vol. 3. Fungi- flora. Oslo, Norway, pp. 287-546. Gir,BERTS0N, R. L. 1961. Notes on western poly- pores. Pap. Mich. Acad. Sci. 46:209217. Gir.BERTsoN. R. J,.. K. ,T. M.artiiv. and .1. p. LiNDSEY. 1974. Annotated check list and host index for Arizona wood-rotting fungi. Univ. Ariz. Agr. Exp. Sta. Tech. Bull. 209. 48 p. Hedgcock. G. G.. and W. H. I.ong. 1912. Pre- liniinaiA" notes on three rots of juniper. Mycologin 4:109-1 13. IjEMKe, p. a. 1964. The genus Aleurndiscux (sensLi lafo) in Norlh America. Can. .1. Bot. 42:72^-768. LiNDSEY, ,T. P.. 'Wood-inhal) gnai'o in A I.iTir.K. ]•:. I... TT.S. n.M)t. ^ND tig RroGWAY. R. 19 nDineiu laluie. by the author. Sii.vw, C. G. 197=; Pacific NoT'tiiv Univ. Agr. Exj R. L. (ill.RERTSON. 1975. lomobasidionivcetes on sa- ■izona. Mycotaxon 2:83-103. ,Tr. 1930. Soufliweslern frees. Agr. Handb. No. 9. 109 p. Color standards and color Washington. D.C. Published Host fungus index for the >est -I. Hosts. Wash. State ). Sta. Bull. 763. 121 p. BODY SIZE, ORCJAiN SIZE, AND SEX RAJ'IOS IN ADULT AND YEARLINC; BELDINC; OROUND SQUIRRELS Marl L. M( WnlH-n .1. I'a AiJSTKACT.-^ A fivcMMi- sIikK uT I',i ■]( I i 1 1^' ^^Miiinil s(iiiii-rcls was roi Sierra Nevada. Bodv \N.M-hl and l.ndv len^illi vane.! seasunalK dcpe. deposition cycle, age. aiul ,se.\. Aduil males tended to he l;ea\ ht -jihI h ticularly in the last half of the active season. A similar pattern was squirrels were often distinguishable from adults on tlie basis ol ImiU greater in adults throughout the season, and mean body lengths weie giealc first half of the season. Adults also had larger internal organs than yearlings , season. In liver and heart this difference was sustained. Se.x ratios in arluh'- 1:1 hut thei-e was c(insid(Mal)le sp.itial and temporal as\nnnetr\' in distiihulK tended to live in aieas pci iphnai to lush nicailow s cm i upied 1)\ [em.ih's .iiid \( lu( ted at liigh altitude in the hug upon the fat depletion- igci- liian adidt females. i)ar- K'siMit in \earlings. Yearling i/(\ MiMii lM)(f\- weights wei-e gii'alcr IN aihilts through the llic l)eginiung of the iid in yearlings were (if the se.xes. Males The Belding ground sc|iiirrel [Spermo- ph'dus heldingi bcldingi) is a hiberiiator that Hves in the central Sierra Nevada Mountains mainly from the eastern di- vide to the edge of the Great Basin at altitudes between 1,825 m and 3,650 m (Storer and Usinger, 1970). During a five-year, mark-release study of 5. b. bcldingi our records of retrapj)ed animals enabled us to compile data on individuals of known age and sex for prolonged periods. In the course of this study it became clear that three functional groups, based upon age. existed within the pojDulation: juveniles, yearlings, and adults. Characteristics of juveniles have been previously reported (Morton, Max- well, and Wade, 1974). Herein we rejiort on seasonal changes in body size, organs, and on sex ratios in both yearling and adult S. b. bcldingi. Methods The study \vas conducted from 1969 through 1975 in meadows and their bor- dering areas in Lee Vining Canyon, Mono County, California. Most of our infor- mation stems from work done at Big Bend (elevation ca 2.100 m) and es])ecially at Tioga Pass (elevation (a 3,000 m). The active seasons are similar in duration for populations at both areas but may begin >^ix weeks or more a})art due to chmatic differences associatc^d with altitude (Mor- ton, 1975). All data reported on body weights, bodv lengths, and sex ratios of squirrels of known age are from Tioga Pass animals. Data on organ weights were combined for the two populations at 10-day intervals throughout the active season in order lo bolster sample size. Squirrels were ca])tured alive in Toma- hawk wire-mesh traps baited with peanut butter. Those to be released were toe- clipjDod, and those retained for specimens were ethcrizecL In some cases specimens were collected with a .22 caliber rifle. Body weights were measured to the near- est b.l g on a pan balance. Body lengths were taken with calipers to the nearest 0.1 ( ni. Wet weights of freshly excised and debrided organs were measured to the nearest 0.01 g on an anal^^ical pan balance. Certain small meadows or sections of large meadows were used only for mark- release studies. Sej)arate data logs were maintained for each toe-clipped animal. In our terminology juveniles are the young of the year, yearlings were born in the preceding year, and adults are all animals older than Aoarlings. Resi^lts The first S. b. bcldingi to emerge each season were adult males. Within a few days, however, some adult females and yearlings could be found. The pace of emergence varied somewhat from year to year, depending upon snow cover. Adults tended to enter hibernation earlier than yearlings. On the average each individual was active above ground for about three months fMorton. 1975). BoDi wEicHT. — There were large sea- sonal differences in bod^' weight due pri- marily to fat depletion or deposition and to sex and age differences (Fig. 1). ^Biology Dcpartmcn -School of Mcdirine Odidcntnl (.'ollogc, I.os iVngcIcs, iiivei'sity ci[ Cnlifoinia. San Diego 505 306 GREAT BASIN NATURALIST Vol. 35, No. 3 Fig. 1. Seasonal change in mean body weight of Spermophilus beldingi beldingi at Tioga Pass. Data were accumulated over five seasons. 1969-73. Numerals indicate sample size; vertical bars de- note ± 2 S.E. Upon emerging in mid-May adult fe- males at Tioga Pass were lighter than adult males, but during pregnancy they became heavier than males. In late July and for the remainder of the active season mean weights of adult males were signifi- cantly greater than those of adult females (P<0.05). The sexes of yearlings were not different in weight until late July. Thereafter, as in adidts, males were heavier. Seasonal trends in body weight were much the same for each sex, par- ticularly in the second half of the season. During the first half of the season year- lings were still growing rapidly. As a group, yearlings never achieved adult weight. Adults were significantly heavier (P<0.05) than their yearling counter- parts of the same sex at every class in- terval throughout the season. The large weight gain observed in all animals dur- ing the last half of the season was due to fat deposition (Morton, 1975). Body length. — Growth in yearlings, as indicated by body length, occurred throughout the season (Fig. 2), but from 23 22 21 20- 19 18 A Yearling Males O Yearling Females 10 20 30 10 20 30 10 20 30 10 20 30 10 20 May June July Aug Sept Fig. 2. Seasonal change in mean ho{l\ length of Spermophilus beldingi brldingi at Tioga Pass. Data were accumulated over five seasons. 1969-73. Numerals indicate sample size; vertical bars denote ± 2 S.E. Sept. 1975 MOHTON. PARMER: SQUIRRELS 307 mid- July on yearling females as a group were indistinguishable from adult females in body length. The same was true of males except that the smaller males handled in August and September invari- ably were yearlings. Body length increased in adult males as the season progressed, suggesting that maximum size in S. b. bcldingi males ma\' not be reached until be\'oiid their second year of life. Organ weights. — Liver weights in- creased rapidly following emergence in all animals (Fig. 3), but the increase was more rapid in females than in males. Between the third and sixth weeks of activity livers of females were larger than those of males (P<0.05). Liver hyper- trophy in females was coincident with lactation. By the twelfth week of the season yearlings had livers of adult size. Livers for all ages and sexes were smaller at the end of hibernation than at the beginning. Ap])arently this organ atrophied during hibernation. The heart, kidneys, and spleen, were larger in adults than in yearlings during the first part of the season (Fig. 4). This difference was particularly noticeable and ])rolonged in heart weight. Sex Ratios. — During the five years of this study a total of 341 yearhngs (170 males and 171 females) and 484 adults (238 males and 246 females) were handled. The sex ratio for either age group did not differ from 1 : 1 according to a chi-square tost (P>0.5()i. Discussion Body size. — Although yearling ground squirrels often represent a sid)stantial ])or- A Adult Males • Adult Females A Yearling Males O Yearling Females 60 Days Fig. 3. Seasonal clianges in mean liver weight of Sprrnwphilus bcldingi beldingi from Big Bend and Tioga Pass. Numerals indicate sample size. Day 0 of abscissa refers to time first squirrels emerged from hibernation. 308 GREAT BASIN NATl^RALIST Vol. 35, No. 3 Days Fig. 4. Seasonal clianges in mean weight of spleen, kidneys, and heart in Spcrmophilus bcl- dingi beldingi from Big Bend and Tioga Pass. Numerals indicate sample size. Symbols as in Figure 3. Day 0 of abscissa refers to time first squirrels emerged from hibernation. tion of the population and may interact in unique ways with other niembers (Michener and Michener. 1973), there is little published evidence that they differ externally from older animals. Even in such large-bodied species as S. undulatus juveniles reach adult size, or nearly so. by the time they are ready for hiberna- tion (Mayer and Roche. 1954). A priori this might be predicted since ca]itive \\\- A'eniles of hibernatory Spcrmophilus. par- ticularly those from high latittide or high altitude, tend to be precocious and to have e.xceptionallv high growth rates (Clark. 1970; Morton and Tung. 1971). In the case of S. h. /)('hli?is:i. at least, by the time they enter hibernation feral juveniles have foot and tail lengths indistinguish- able from those of adults (Morton and Tung, 1971). Nonetheless, neither maxi mum body weight nor itiaxinuun boch length is achieved in .S'. h. hcldingi until well into the second or possibly even third year of life. Furth(M-mor(\ we have found that yearling males are sexualh- inunatuir and do not reproduce (Morton and (lal- lup. unpubl.). Yearling females do vv- p/roduce. Similar age (hfferences in re jiroductive capacity have been found in 5. arrnatus (Slade and Balph. 1971 i. The differences in body size noted in age classes of S. h. beldingi probably are not unique among ground squirrels. Such differences are likely to be overlooked un- less the investigator examines large num- bers of animals of known age over a span of several consecutive seasons. Sex r.\tios. — An unbalanced sex ratio in favor of females has been reported for many ground squirrel populations. This ratio may approach or exceed 3: 1 (Mc- Carlev. 1966; Michener and Michener, 1971;^Sheppard. 1972; Turner, 1972). In a few cases, however, the ratio foimd did not differ significantly from 1:1 (Clark. 1970; Murie, 1973; present study). In his study of S. h. orcgonus. Turner (1972) foimd that the sex ratio was 1:1 in juveniles. He suggests that because ju- venile males tend to wander and explore more than females they incur greater mortality, resulting in an unbalanced sex ratio in older animals. In S. h. hcldingi the sex ratio in juveniles is also 1:1; males probably wander more than females in that they have larger home ranges than females and are more likely to expand their range late in the season (Morton. Maxwell, and Wade, 1974). Although this behavior would seem to make juve- nile males more susceptible to predation. we have no evidence that it does. To the ( ontrarv. males and females occur in equal tnunbers in both yearlings and adidts. The sexes are not distributed ran- domly throtighout the habitat occupied, however. In our trap]iing at burrow sys- tems located in lush meadow areas re- served for mark-release studies, adult fe- males outnumbered males by 1.3:1 when all data were summed. The ratio tended to fluctuate seasonally, however, and was sometimes near 3:1, particularly at mid- season. We often captured adult males at a particular burrow system oidy a few times per season, during the first days or weeks following emergence and again at th(^ \ erv end of the season. This suggests that some males were forced from the colony \)\ conflicts associated \\dth repro- (hiction and were able to return only in time to (Mitcn- their trachtional hibernacu- liun. If this is correct, it follows that dis|)laced males should be found in areas jx'ripheral to main colonies. We found this to be the (as(>. When males were de- sii-ed for spin imens. they cotdd usually be found b\- collecting" animals scattered in lodgepole pine stands fringing mea- Sept. 1975 MORTON, PARMER: SQUIRRELS 309 flows, and in rock)- outcro])s and talus slopes in steep areas above the meadows A tendency for males to live in areas peripheral to the main colonies has been observed previously in S. armatus (Balph and Stokes, 1963) and in S. richardsonii (Quanstrom, 1971). Turner (1972) did not believe, however, that this occurred in S. b. oregonus. The spatial distribution of the sexes could vary enormously de- pending upon such factors as habitat physiography and upon intrinsic charac- teristics of the population itself. Undoubtedly unbalanced sex ratios ex- ist in many ground squirrel populations. Such ratios may be a function of inter- populational or interspecific differences in breeding systems and social organization (Murie, 1973). However, an investigator who confines his work to locations with maximum animal density or to those of easy accessibility could obtain an inac- curate measurement of sex ratio. Acknowledgments. — We wish to thank John Gallup, Roland Leong, Cath- erine Maxwell, Allan Tway, and Charles Wade for assistance in trapping opera- tions. Southern California Edison Co. pro- vided housing for two seasons. Financial support was provided by Occidental Col- lege and by National Science Foundation Grant GB 29146X1. Literature Cited Balph, D. R., and A. W. Stokes. 1963. On the ethology of a population of Uinta ground squirrels. Am. Midi. Nat. 69:106-126. Clark, T. W. 1970. Richardson's ground squir- rels {Spermophilus richardsonii) in the Lara- mie Basin, Wyoming. Great Basin Nat. 30: 55-70. Mayer, W. T., and E. T. Roche. 1954. De- velopmental patterns in the Barrow ground squirrel, Spermophilus undulalus barrowensis. Growtli 18:53-69. McCarley, H. 1966. Annual cycle, population dynamics and adaptive behavior of Citellus tricedemlineatus. J. Mammal. 47:294-316. MiCHENER, D. R.. AND G. R. MiCHENER. 1971. Sex ratio and interyear residence in a popu- lation of Spermophilus richardsonii. J. Mammal. 52:853. MiCHENER, G. R., AND D. R. MiCHENER. 1973. Spatial distribution of yearlings in a Richard- son's ground squirrel population. Ecology 54:1138-1142. Morton, M. L. 1975. Seasonal cycles of body weights and lipids in Belding ground squir- rels. Bull. So. Calif. Acad. Sci. In press. , C. S. Maxwell, and C. E. Wade. 1974. Body size, body composition, and behavior of iuvenile Belding ground squirrels. Great Basin Nat. 34:121-134. . and H. L. Tung. 1971. Growth and development in the Belding ground squirrel {Spermophilus beldingi beldingi) . J. Mam- mal. 52:611-616. Murie, J. O. 1973. Population characteristics and phenology of a Franklin ground squirrel '{Spermophilus franklinii) colony in Alberta, Canada. Am. Midi. Nat. 90:334-340. Quanstrom, W. R. 1971. Behavior of Richard- son's ground squirrel Spermophilus richard- sonii richardsonii. Anim. Behav. 19:646-652. Sheppard, D. R. 1972. Reproduction of Rich- ardson's ground squirrel {Spermophilus rich- ardsonii) in southern Saskatchewan. Can. J. Zool., 50:1577-1581. Slade, N. a., and D. F. Balph. 1974. Popula- tion ecology of Uinta ground squirrels. Ecol- ogy. 55:989-1003. Storer, T. I.. AND Usinger, R. L. 1970. Sierra Nevada natural history. Univ. Calif. Press, Berkeley. 374 pp. Turner, L. W. 1972. Autecology of the Beld- ing ground squirrel in Oregon. Ph.D. thesis, Univ. of Arizona, Tucson. 149 pp. PHOTOPERIODIC RESPONSES OF PHENOLOGICALLY ABERRANT POPULATIONS OF PIERID BUTTERFLIES (LEPIDOPTERA) Arthur M. Shapiro' Abstr.'^CT. — T^vo local pierid populations in western Norlh Ainerit.i showing regi(jnalh' aix'rrant l)henologies were investigated in the lahoratory. Neither a parlialK hivoltine Piciis iiapi from the Sierra Nevada foothills in El Dorado County. California i sui rnuiidi'd hy luiivoltine populations), nor a vcrnal-univoltine P. occidenlalis from a foothill outlier of llie (Colorado Front Range (below hi- voltine populations) showed unusual resjwnses to controlled developmental regimes in the labora- tory. Their unusual phenologies are hypothesized to be the produi t of microclimate. Failure to under- go genetic adaptation to unusual microclimates is discussed witii ])articulai- reference to the i)res- ence or absence of gene flow from neaib\' normal populations. The timing of life-history phenomena in an insect population is (leterminetl by physiological responses to en\ironniental stimnli. These proximate controls reflect a genetic basis Ijelievecl to be the jiroduct of natural selection for seasonal cycles appropriate to the environment of the population. In the western LTnited States topography has a dramatic impact on cli- mate, and great differences may occur over short ground distances. How closely can insect populations ada])t to their im- mediate climates on a microgeograj)hic scale? Phenological adaptation is merely one case of the more general jiroblem of population differentiation (cf. Ehrlich and Raven, 1969; Ehrlich et al., 1975). In most organisms, at least prior to the advent of electrophoretic genetics, popu- lation differentiation was assessed on the basis of visible phenotypic characters. Such characters, like the enzyme systems studied by electrophoresis, are often not translatable into specific selection pres- sures. In markedly seasonal climates the nature of selective pressures acting on phenology may be very apparent. Where local deviations from the broad geograph- ic pattern of voltinism are observed in a species, the potential exists for the demon- stration of microgeograj)hic (or ecotypic) differentiation. This is the fourth paper in a series exploring the evolution of sea- sonality in the butterfly genus Pieris in western North America. In various multi\oltine Pieridae both phenotype and diapause are under photo- periodic control. The two sets of develop- mental options (diapause /direct develop- ment; vernal /esti\ a 1 j)henotype) may be physiologically coupled {Pieris napi Lin- 'Departmcnt of Zoology. Univcisily n[ Cnlifoniin. D.ivi';. Ca naeus complex) or not (P. protodice Bois- duval & LeConte, P. occidciitdUs Reakirt). Recent st tidies have shown that univol- tinism in both grouj)s is derivative from multivoltinism, accompanying invasion of a short-summer climate (P. occidcn- ta/is, Sha})iro, 1975a) or persistence in a progressively drier one (P. napi, Shapiro, 1975b). Such patterns are defined over broad geographic areas. California P. napi, for example, is differentiated into a commonly biv'oltine, heavily pigmented subspecies in the coastal summer-fog belt and a univoltine, more lightly marked subspecies in the interior, where summers are clear and hot. The transition between the subspecies apjjears to be in the form of a stee]) cline through the central Coast Ranges (Shapiro, in preparation). Recently Lees and Archer (1974) have reported the existence of phenological dif- ferences among napi populations on a mvich finer scale. They have found ap- parently relict univoltine populations in suitable (bog-heath) habitats completely surrounded by multivoltine ones in the British Isles. Their preliminary interpre- tation of this situation is that it provides (n ideiice for midtiple invasions of Britain h\ napi stocks having different pheno- logical characteristics and source regions. In th(> course of recent work on pierid |)hen()logy and e\olution. the existence of regionally aberrant populations has b(>en i)r()ught to my attention in both the napi and protodicc-occidcntalis groups. In both ( ases the populations appear to be uniciiie, rather than forming a repeating pattern as in British P. napi. They would therefore seem to be good candidates for 310 Sept. 1975 311 local genetic tliflcreiitiatioii under al\[j ical microclimates he Sierra Foothills ricns iiapi ii Pieris napi from interior (-alitornia are. as noted above, luiivoltine and moiio- phenic in nature. Under laboratory (au- ditions they can lie reared without dia- pause; then they produce the eslixai phenoty|)(» ''castoria'' Airtually unknown in the wild in the interior (Sha])iro. 1975b). In June 1974 Mr. William Pat- terson of Sacramento, California, took se\-eral wild 'Vy/.s/o/vV/"" of hoth sexes in the canyon of the American Ri\ er below Auburn in the Sierra Nevada foothills (El Dorado County, 650 feet). The oc- currence of a second brood there was con firmed in 1975. P. napi is connnon in the canyon, producing its usual \ernal pheiid type in March, llie second brood, whi( li is much scarcer, unlike the first is ex- tremely localized ^vithin llu^ camon -al present being known from onh' two densely shaded ravines where the intro- duced cruciferous weed watercress (Nas- lurliuni offn iiialc W. Br. turliu!ih(i(jii(ih(nni Schii gr'ows in [xTincUient stn Rorippa na.s- :. & Thell.) ims (Fig. 1). hutterflies are bred Sierran Most .,r ih,> uiid ,1,1, idonlicil to lal)orat( "cdstnrid' I h'ig. 1 i . On J<) Man h l<)7) niii(> in.de and three lemale lirst-hrood, vernal phenotyjie napi were colIectcMl in one of these ravines. These included two coj)ulating j)airs in which the females were soft-winged, in- dicating that they had developed in the ravine^ itself. The eggs from these females were used in photoperiod experiments ( Table 1 ) . ( Rearing methods are de- scribed m Shaj)iro. 'l975a and 1975b.) Hie results are entirely typical for Sier- ran stock and do not suggest that x-Xmer- i(an Ri^er material has a greater pro- pel isit\' to flevelop directly than do stocks tVoju purely uniAoltiiie localities, at least under our laboratory regimes. However, ihis is not particularly surprising. The second brooci of napi in the American Ri^■er gorge is nukh rarer than the first, indicating that is is onl}' partial; its num- bers also fluctuate from year to year. B B (^5SS£S^IJ4;>rSjgx%,c-;.v|f^Vr^ Fig. 1. Locations of ravines ("B") where !)ivoItiiie Pieris tuipi univoltine napi are generally distriluited at low (Icnsity. l'S(;S 7. Ill the American River gorge; life "Aulnun"' quadrangle. 312 GREAT BASIN NATURALIST Vol. 35, No. 3 Table 1. Incidence of diapause ( D) and non- diapause (ND) pupae in bivoltine (American River, 650') and univoltine (Placerville, 1800') Pieris nopi from El Dorado Co., Calironia, reared on watercress at 27 C under two photoperiods. Photophase: Continuous 15 hr Stock: Pupae: D ND D ND American River Placerville 15 29 16 10 23 18 (In 197:5 three trips by Patterson and Shapiro in season turned up only tNvo males and one female. A later search of the host plant at the optimum time failed to turn up ;ni\ napi immatures, although ten Pieris rapac lar^'ae were found.) These circumstances suggest that the production of a second brood here is ac- cidental, resulting from the peculiarly cool and moist conditions within the ra- \'ines. There is no evidence that the bi- Aoltine sites are in any sense isolated from adjacent univoltine ones, nor is it clear that there is successful re])roduction by the second brood in all years nor even that there is genetic continuity from year to year in the ravines; perhaps a few pu- t ^P'- Fig. 2. Wild second-brood Pieris napi from the American River gorge, collected by W. Patterson in June 1974 fmales at top; dorsal (left) and ventral (right) surfaces). The heavily marked female is atypical for an inland population. Fig. 3. Phenotypes of representative lab-reared nondiapause Pirris napi from the Ameiican Riv( stock; 27 C. continuous light; dorsal (left) and ventral (right). Sept. 1975 SHAPIRO: BUTTERFLIES 313 pae will develop directly there whenever any female napi happens to colonize them. Experienced California collectors (R. L. Langston, B. Walsh) agree that even near the coast some localities produce second-brood napi every year and others only rarely or sporadically. Exj)eriments have shown both developmental and phenotypic differences between coastal and inland stocks but not among the coastal stocks themselves. Watercress is known to be host of P. napi in various Sierran sites up to about •5,000 feet (Shapiro, 1975c). The only other record of a Sierran ^'castoria" known to me is a fresh male taken flying Table 2. Incidence of diapause (D) and non- diapause (ND) pupae in two split broods of a Barbarea verna-ieeAin^ univoltine Pieris napi (Gates Canyon, Inner Coast Ranges, Solano Co., 750') reared at 27 C on continuous light. None of the differences was significant. Brood Host Pupae: Brassica kabet-^ 7 Nasturtium officinale^' 5 Brassica kaber''^ 6 Lepidiuni latifolium^'-'' 5 ND develop- mental time in days 21 25.2 11 25.3 14 25.6 16 24.6 Notes: [a] Tops, (b) Elongating rosettes. Ui In subsequent experiments mature tops did not supirart development. Butterflies in this brood were stunted. among first-brood vernal napi at Lang Crossing, Nevada County, 4,500 feet, 9 .June 1975. At this locality napi feeds on both watercress and native vernal cru- cifers. There are several possible explan- ations of this odd individual, but to test the hypothesis that watercress feeding itself inhibits dia})ause, split-brood experi- ments were conducted in 1975 using an Inner Coast Range stock (Gates Canyon) with no previous exposure to the plant. No evidence of a dietary influence on the incidence of diapause was found in this univoltine strain (Table 2). Pieris occidcntalis in Colorado Haystack Mountain (5,589 feet) is an isolated hill eight miles northeast of Boul- der, Boulder County, Colorado (Fig. 4), where Dr. Ray E. Stanford of Denver has for several years taken small, dark vernal ''calycc' phenotypes of Pieris oc- cidentalis indistinguishable from the single brood above treeline in midsmnmer (Fig. 5). He has no summer records of P. occidentalis from Haystack Mountain but finds its lowland sibling P. protodice there in summer instead. Because P. pro- todice winters only very locally but colo- nizes widely in summer, this is not sur- prising; it does however, raise the pos- sibility that P. occidentalis has undergone a phenological shift to univoltmism in 7 r^- r> t Fig. 4. Location of Haystack Mountain, Boulder County, Colorado. USGS 7.5-minute "Boulder' and "Niwot" quadrangles. 314 GREAT BASIN NATURALIST Vol. 35, No. 3 Fig. 5. Wild Pieris occidentalism vernal phenotype C'calycc''). from Haystack Mountain, collected by R. E. Stanford. Males at left; dors;d (left) and ventral (right) surfaces. response to competition from P. protodice (although no such phenomenon is known at other localities where the two are sym- patric). Haystack Mountain is probably the lowest elevational record for P. oc- cidentalis in Colorado. In the Rockies proper it is bi\oltine at middle elevations (perhaps locally trivoltine) and univol- tine in the Alpine zone (cf. Brown, Eff, and Rotger, 1957) and has two seasonal phenotypes (Shapiro, 1975d). A laboratory stock was established from ova laid by five females collected by Stanford on 6 April 1975. Under labora- tory conditions their developmental and phenotypic responses (Table 3 and Fig. 6) were identical to both Sierran multi- voltine and Colorado Alpine stocks (Sha- piro 1974, 1975d). Once again we have no experimental evidence for the evo- lution of a phenological ecotype and are therefore forced to look for micro- climatic explanations. Since Haystack Mountain is effectively in the Great Plains climatic regime, which is hotter and drier than the usual regime of P. occidentalis, it may not be surprising that conditions there would be associated wdth summer dormancy. This (juestion can be settled only b}' laboratory duplica- tion of Haystack Mountain conditions or by testing the developmental responses of multivoltine P. occidentalis stock from elsewhere reared at Haystack Mountain. We hope to carry out such experiments within the next couple of seasons. Phenological differences are known be- tween plains and lower montane popu- lations of a inunber of Colorado butter- flies (.I.A. Scott, R.E. Stanford, pers. comm.), but they may go in a direction o])posite to those observed in Pieris oc- cidentalis. Two species (Colias olexandra Edwards, Pieridae; Plebeius icarioides complex, Lycaenidae, both Legume feed- ers) are bivoltine on the plains and uni- voltine in the mountains. The basis for these differences is uninvestigated. With no evidence for genetic differen- tiation of Haystack Mountain occidentalis. the attractive hypothesis of competitive Tahi.e 1 Incidence of diapause iD) and non- diapause (ND) pupae in veiiial-univoltine (Hay stack Mountain, Colordao, o.oSO') and bivoltine (Donnor Pass, Colorado, 7,000') Pieris occident- alis reared on Brassica kaher at 27 C. Stock: Photophase: Continuous 15 hr Pupae: D ND D ND Haystack Mountain. 1975 Donner Pass. 1973 0 22 0 16 Sept. 1975 SHAPIRO: BUTTERFLIES 315 ^ '. -^_f ^- -1 Fig. 6. Phenotypes of representatives lal)-reared nondiapause Picris occidcntalis from the Haystack Mountain stock; 27 C. continuous light. Estival phenotvi)es characteristic of nnitlivoltine. Males at left; dorsal (left) and venti-al (right) surfaces. seasonal displacement with protodicc must be set aside. The host plants of both species on Haystack Mountain are un- identified. Both prefer species of pepper- grass, Lepidium, throughout their ranges. On the plains most crucifers are vernal species, as in lowland California. DiSCEISSION Many instances are on record of eco- typic differentiation on a microgeogjraphic scale, particularly in plants, which have more versatihty in developing isolating mechanisms than do animals (Jain and Bradshaw, 1966). As noted above, the lack of a genetic basis for biovoltinism in Sierran Picris napi is not very surprising, granted the extremely restricted habitat and the ex- tensive distribution of univoltine butter- flies, with ample opportunity for gene flow. The failure of the Haystack Moun- tain P. occidcntalis to differentiate is more intriguing. It is, of course, possible that it has differentiated and that the lab rearing regimes were too crude or inap- propriately selected to show it. It is cer- tain that experiments to date, involving simple manipulation of constant rearing temperatures and unchanging day- lengths, have gi^en an oversimplified picture of the developmental versatility of pierids in the field. If microclimate determines aberrant voltinism in these stocks, it is very likely that humidity, for exam])le, may interact with photo- period and temperature in controlling de- \'elopment in natural populations. The same genetic information may allow Picris occidcntalis to respond ap])ropriate- ly to regimes as diverse as those at Hay- stack Mountain (5,589 feet) and Love- land Pass (12,400 feet). Given such jilasticity, we may wonder whether the Baldwin effect (Simpson, 1953) might not come into play in popu- lations in extreme environments. Briefly, the Baldwin effect postulates the buildup by selection of a genetically obligate basis for the ada})tations produced via develo])- menlai plasticity. In an atypical hut pre- dictable climate like Haystack Mountain, might not the developmental flexibility chara( t(>ristic of montane poj)ulations be lost? (^Alaskan Picris occidcntalis nclsoni seem to be evohing in this direction; Shai)ir(>, 1975a.) One important counter- 316 GREAT BASIN NATURALIST Vol. 35, No 3. vailing force would be gene flow, which is almost certainly operating on high- elevation univoltine occidcnUdis in Colo- rado (Shapiro, 1975d). Haystack Moun- tain is about 30 air miles from timberline and much closer than that to the montane zone, but how isolated it actually is is quite unknown. Nor is there any infor- mation bearing on how long occidentalis has been there — whether it is a Pleisto- cene relict or a recent colonization. There are much more isolated, certainly relict occidentalis populations in other localities east of the Front Range — the Black Hills of South Dakota and perhaps the Pine Ridge of northwestern Nebraska — which deserve study in this regard. Acknowledgments Of the collectors who provided vital in- formation and who have been credited in the text, special thanks are due Mr. William Patterson and Dr. Ray Stanford. without whose help these experiments would have been impossible. Mr. Mark Kauzer assisted in field w^ork and Mrs. Adrienne R. Shapiro in rearing. This re- search is part of a larger study of colo- nizing ability and the evolution of season- ality in Pieris funded by the Committee on Research, UCD, under grant D-804. LiTER.-^TURE Cited Brown. F. M., D. Eff, .\nd B. Rotger. 1957. Colorado butterflies. Denver Museum of Natural History. Denver. 368 pp. Ehrlich. p. R.. and p. H. Rwen. 1969. Dif- ferentiation of populations. Science 165: 1228-1233. EiiRLicH. P. R.. R. R. White. M. C. Singer. S. W. McKechnie. and L. E. Gilbert. 1975. Checkerspot butterflies: a historical perspective. Science 188:221-228. Jain. S. K.. and A. D. Bradshaw. 1966. Evo- lutionary divergence among adjacent pop- ulations. I. The evidence and its theoretical analysis. Heredity 21:407-441. Lees, E.. and D. M. Archer. 1974. Ecology of Pieris napi (L.) (Lepidoptera. Pieridae) m Britain. Ent. Gazette 25:231-237. Shapiro. A. M. 1974. Photoperiodic control of seasonal polyphenism in Pieris occidentalis Reakirt (Lepidoptera: Pieridae). Wasmann J. Biol. 31:291-299. . 1975a. Photoperiodic control of de- velopment and phenotype in a subarctic population of Pieris occidenlalis (Lepidoptera: Pieridae). Canad. Ent. . 1975b. Developmental and phenotypic responses to photoperiod in uni- and bivoltine Pieris napi (Lepidoptera: Pieridae) in Cali- fornia. Trans. Rov. Ent. Soc. London 127: 65-71. . 1975c. The role of watercress, Nas- lurtiuni officinale, as a host of native and introduced pierid butterflies in California. J. Res. Lepid. . 1975d. Ecotvpic variation in montane butterflies. Wasmann J. Biol. 32:267-280. Simpson. G. G. 1953. The Baldwin Effect. Evolution 7:110-117. ADDITIONAL RECORDS OF Ri:i?TILl^:S I'ROM .lAIJSCO. MEXICO Philip A. Mcdira'. Rii,l,,ir (i. Ain.ll-. ,ni.l l.nurs I',. I)iv,,n^ Abstract. — Notewortliy ri-cords ol tr|ililc^ iVoi first state recoi'ds of Colennyx clrga/is /u-nioi tills, llic notes on the distribution ;ind roprodiK tinti of F.umrcc l.diMii. Affxito. are presented. The KM 111 d for Gcophis tarasra<\ and if/duhl/us are discussed. A small collection of amphibians and reptiles from Jalisco, Mexico. \ icldc^l several noteworthy specimens. Coleonyx elegans nemoralis I\laiil)(M'. One male (88 nnn snout to Aciit lo]i<>th. SVj from 28.1 km SW Autlan, .lali'sco. (Texas Cooperative Wildlife Collection, i^exas A & M University, TCWC 48035 ). collected on 19 May 1974. elevation 518 m. There are three known sjiecimens from Jalisco. The first, a male (Los An- geles Coimty Musemn, LACM 37568 ) 83 mm SV, from 74.2 km SW Antlan. Jalis- co, was collected by J. R. Dixon and R. Heyer on 23 July 19fv. This specimen has 10 preanal pores; 7-7 supralabials; h-7 infralabials; 9 scales nostril to nostril; 0 gulars contacting mental; 4/4 gulars con- tacting first infralabials; 21 rows of tuber- cles across at midbody, and 16 fourth toe lamellae. Our specimen (TCWC 48035 ) has 11 preanal })ores; 8-8 supralabials; 7-8 infralabials; 9 scales nostril to nostril; 5 gulars contacting mental; 1/5 gulars con- tacting first infralabials; 20 rows of tuber cles across at midbody, and 1 7 fourth toe lamellae. The third specimen ( Brigham Young University, BYLT 41299). a male 85 mm SV, from 14.5 km N Barra de Navidad, Jalisco, was collected by J. Ott- ley on 30 October 1974. This spetimen has 10 preanal pores; 6-7 supralabiaL; 7 -S infralabials; 9 scales nostril to nostril!; (> gulars contacting mental; 2/2 gulars con- tacting first infralabials; 21 rows of tu- bercles across at midbody, and 16 fourth toe lamellae. All specimens are within the range of variation described by Klauber (1945). These specimens extend X\\v known range of this species inland some 107 km N from the coastal area of Cf)- lima, the heretofore northenmiost portion of its known range (Klauber. 1945; Kluge. 1975). Eunieces brevirostris iudiibitus Taylor. A series of 20 specimens, including 9 jm c- 1U.C.I..A., P. 0. Box 495, Mercury, Xcvnda SOUJ !. ^Faculty of Natural Sciences & Mnthematics, Slofkl-u Sl.il.' ^Dcparimcnt of Wildlife Sciences, Texas A Sc M T"ui\ci^ii v niles (2l-)0 mm SV. x -- 26.7 mm) and n adidts (52-75 mm SV, x - 61.2 mm), collected 25 km '>>¥. Autlan, Jalisco (J'CWC ISO 50- 18055 I. on 17 May 1974. J'his siic is appro\iniat(4\ 50 km W of the ne.ucsl i-eporled hxalitv and about 100 km SSE of the northernmost locality for this subspecies, both in Jalisco, thus par- tially filling the hiatus in the distribution described 1)\ Dixon (1969). Our speci- mens (>xhil)it large female (75 mm SV. weight ().()7 g) that bore six live young between llie time of capture and the next morning. Iheir r.uige in length was 24- 28 mm S\' (X 2<). > mm) and in weight from 0. )7-0. I() g (X 0.11 gi. All Kumeces \^('re Found in pine-oak woodland imder rocks within 20 m of a |)ermanent stream, elevation 1.1-3 3 m. It is of interest to note (hat in this area there had been no ap])re- ciable rainfall since October 1973, and most decidous vegetation was devoid of lea\es. Despite |he apparent lack of mois- lure. /''u/ncccs h. 'mdiihitus was capable of reprochu ing. perliaps o\\ing to its ovo^'i- \iparous nature. (h'ophis ididsciw flartweg. One female from l-\ km S1-: Autlan. Jalisco (TCWC l'79l-8), collected on 17 May 1974, eleva- tion 1,1'33 m. J1hs apparently is but the Fifth known sj)ecimen oF this species and the stMond from th(> slate of Jalisco. Downs (1967) rej)ortecl on three speci- mens From tli(> ty])e locality at Uruapan, \lichoa(an; Dixon (19()8i reported one spcH imen From Ne\,ido de Colima some FS km 'o [he !■'.. Our s[)ecimcMi has 15 nll..,^,.. P.iiiiM,,.,. Xcw Jci>ey nS2-IU^ 317 318 gri:at basin nai Vol. 35, No. 3 scale rows; 146 ventrals; 41 caiidnls; 0 + 1 temporals; 6 supralabials; 6 infralabials; 1 loreal; 1 postocular; no preocular, and 10/10 maxillary teeth. The first pair of chin shields is twice the length of the second }:)air; sujiraocular distinct, larger than loreal; internasals (H^ided and dis- tinct. The dorsum and tail are dark gray with blackish crossbands (40 on the body, 13 on tail) which do not extend across the venter; anterior crossbands 2-3 scale rows wide and bordered on the anterior and posterior by white-edged scales, pos- terior crossbands 1 scale row wide and highly irregular, frequently broken later- ally and dorsally; head distinct from neck; color of rostral and ])renasals similar to adjacent scales; eye twice into snout length; ^'enter greenish white in life, heavily spotted with black. The total length is 236 mm, and the tail is 45 nnn. This specimen has fewer Aentrals than reported by Downs (1967) and Dixon (1968). Habitat was as described for Eumeces brevirostris indul>itus above. The authors are indebted to Dr. An- tonio Landazuri Ortiz, Director General de Fauna Silvestre of Mexico, for pro- Aiding the necessary collecting ])ermits. We also tlijuik Drs. Wilmer W. Tanner ,iM(l Idlni W. Wright for loan of speci- mens in llieir care, and are grateful to Plnli[) I. M(Mlica for his untiring assis- tance in the field. l.ri i;i! MLKI'. ClTKO Dixo'V, I. I'. I"I()S. Notes on tlu- snake genus Gcf)f)/iis. Iiom Nevado de Colima Jalisco, Me\i,,K Sniiiliwest. Nat.. 13(,+):452-454. I '((]'). I'axononiic review of the Mex- ican skinks nl the Eumeces brevirostris group. I.os Auiivlv^ Coimtv Mus. Contrih. Sci. No. Ifi8: !-')(). Downs, I''. I.. l')f)7. Intrageiieii( I'ehitionships ;iiiioiig- (ohihiid snakes of the s<"iius Geophis Wauh'i-. Misc. I'uhh Mus. /ool. I'liiv. Mich. Ki.AUHi.H. I.. M. I'*!-"). 'I'lie fie, kos of tlie genus Colc'i/nv uith (les( liptions of new suljspecies. 'I^-.iiis. San Dh'Ko So,. Nat. Hist. lOi'll): I >')-2Mi. Kir, a;. ;\. (.. l')7'). l^ln l,ifj;eiieti, lelationsiiips ,inil ,'\i)hihiiiiai \ triTifls in tlie eublepliarine ll/ai-,1 f;,Mins Cnlrnnrx. Copeia 1975 (l):24-35. ■|'\Ni\i;H. \\'. \\'. IM'xS. Two new skinl^s from Dinanf;,). Me.xi,,). Great Basin Nat. 18(2): ")74)2. INVASION OK BIG SAC^EBRUSH [ARIEMISIA rRlDENlATA) BY WHITE FIR {^ABIES CONCOLOR) ON THE SOUTHEASTERN SLOPES OF THE WARNER MOUNTAINS, CALIFORNIA rii IS R. Vale Abstract.- - \Vliito I'ir [ Ahics concolor) appears to he i sagebrush (Artemisia tridcniata) on the southeastern slopes northeastern California. The time of initial tree establishment increment borings. Possible causes of the invasion involving s tory. and grazing use of the area during years of the (>stahlisii ('omestic livestock, particularly sheep, appears respousil)l<' f tation and allowing tree invasion. i\a(hng \('g('tation (Joniiiiat(>(l b\- big if the ^^'al■ner Mountains of e.xtrenie within the shrubs was detennined by ic(ili( (limalic conditions, fire his- iiciil were cxidoied. Heav\- grazing by V .iltcrnig the sagi-brush-grass vege- The Wariicf Moiiiitaiiis of cxtrGme northeastern California rise to 3,000 ni above sea level atid snj)port a forest dominated by white fir {Ahics concolor) and ])onderosa pine {Pinus ponderosa) (Fig. 1). The adjacent valleys lie at 1.220 ni and, where not irrigated for pas- ture or hay crojjs, are co^'ered b}' thick stands of big sagebrush {Artemisia tri- dentata) with an understory of herbac- eous plants. In the southeastern portion of the range, the transition zone between coniferous forest and shrubs is charac- terized b}' populations of small fir trees within the brush, suggesting a recent downslope movement of the trees (Fig. 2). The cause of this invasion of sage- brush by white fir in the Warner Moun- tains is the focus of this paper. Relation to Other Studies Many observers have noted invasions of sagebrush by tree sj)ecies in various parts of the Intermountain West. Al- though pinyon pines (Pinus rnonophylla and P. edulis) and junipers {Juriiperus spp.) are the most common invaders of sagebrush (Arnold et al., 1964; Black- burn and Tueller, 1070; Burkhardt and Tisdale, 1969; Cottam and Stewart, 1940; Wright and Fisser, 1968), lodgepole pine (Pinus contorta) also has been found ex- panding into stands of Artemisia (Patten, 1969). Moreover, young trees of ponder- osa pine in eastern Oregon and Jeffrey pine {Pinus jeffrcyi) in eastern Califor- nia may be readily observed within sage- brush areas along forest edges. Although all the above vegetation changes imply a recent establishment of trees in environ- ments formerly unsuited to them, studies 'Assistant Professor, Department of Geogmphy, University- (i lia\'e not yet documented the in^ asion of the relati^el}' xeric big sagebrush by so mesic a species as white fir. In these previous studies, the initiation of tree establishment is often found to correlate with periods of intense livestock grazing. Cattle may deplete the herbac- eous plants in the vegetation, thereby re- ducing the full utilization of the habitat's resources (e.g., soil moisture); this "open- ing" of a formerly "closed" plant cover public domain lands administered by the Bureau of Land Manageme STUDY SITE 35 Fig. 1. The Waiiier Mountains area of e.v- tieme noitheastern California. 319 320 GREAT BASIN NATURALIST Vol. 35, No. 3 Fig. 2. Eastern slope of the Warner Mountains with the study area in the middle background. may permit the establishment of species previously excluded (Robertson and Pearse, 1943). In a lodgepole phie inva- sion of grassy meadows, Vankat (1970) determined that most invasive trees were established immediately after, rather than during, a time of grazing by sheep. Trampling by the animals apparently killed seedling trees, but the heavy browsing and grazing did make the mea- dows susceptible to successful invasion after the sheej) were removed. Another cause sometimes imoked to exj)lain tree advances into sagebrush is climatic fluctuation. Patten (1969) and Arnold et al. (1964) both suggest that, during abnormally wet j)eriods. trees may become established in shrub environ- ments typically too dry for them. John- sen (1962) offers a \ariation on this theme by speculating that long drouglits may initiate the invasion by rechicing the cover of brush and herbaceous ])lants; when moist conditions return, trees arc able to sprout and survive in the "opened'" vegetation stand. This sec[uence of events is analogous to the "opening" of a "closed" plant community by livestock grazing. Fire suppression is often invoked to ex- plain the iiiA'asion of woody growth, usually trees, into grasslands in the south- ern Intermountain West (e.g., Foster, 1917; Pearson, 1931; .Tohnsen, 1962). Blackburn and Tueller (1970), moreover, suggest that a decrease in fires, together with overgrazing, accounts for the inva- sion of pinyon pine and juniper into brush in eastern Nevada. Abundant evi- dence exists suggesting that fires retard the spread of woody plants in the south- ern Intermountain West, although inter- pretation of tree invasion into sagebrush as a resj)onse to fire su])pression is com- plicated hv the fact that both the trees and the slwub', suffer from freipieiit fires. MiniioDs A site V) km soulli of the town of (>edar\ille was selec IcmI (oi- sampling the age striK liii-(> of ihe \oiiiil; lre(>s. The site appeared to he represeiital i\ e of the ex- posure, slope. cUkI \ (^i^etatioii character- ized b\ invading white fir in the south- eastern Warner Moinita.ins. Five plots, each 50 m l)\ 60 m, were located at in- tervals of 0.5 km along the lower limit Sept. 1975 ^^^LK•. white fik I!NV\sTC)^^ 321 of young trees: an atlditional plot was lo- cated in a stand of young fir trees at a higher elevation where a southeastern (>\- posure caused a high(>r forest-shruh tr.ni- sition. The selection of plots along the lower Hmit of these Ivvv^ wa^ designed to estahlish the [jerioil during \vhicli white fir invaded that portion of the area usually considered least hospitahle to it. AA'ithin each plot, all trees greater than 20 cm dbh were cored with an increment borer, while the time elapsed since tree establishment was estimated to be (ught years plus those indicated by the tree rings. Trees with diameters smaller than 20 cm were recorded by estimated height. Results and Discussion Although invasion began between 1915 and 1919, most white fir became estab- lished between 1925 antl 1944 (Table 1). After 1944 a decrease in successful tree establishment is suggested by a gap in the age structure. More recent repro- duction seems improved, judging from the relatively large number of trees between 0.5 and 2 meters in height. The absence of dead trees of any size precludes the possibility that older cohorts suffered mortality, an e^-ent which would compli- cate iu\-< nation of the initial tree Climate. — The years correlated with I lie beginning of tree invasion constitute the driest ])eriod in the historical record, i.e.. since the 18()0s (Fig. 3). Consequent- ly, the first white fir sprouted and sur- vived, not when climatic conditions would ha\(' l)e(>n most favorable for this Tablk 1. Age structure of trees in sample plots and numbers of seedlings by height. Trees whose diameters exceed 20 cm Year of establishment Number of trees Prior to 1915 0 1915-1919 1 1920-1924 1 1925-1929 6 1930-1934 8 1935-1939 10 1940-1944 6 1945-1949 2 1950-1954 1 1955-1959 4 Trees wliose diameters are less than 20 cm Height HI m Number of trees 2-3 8 Less than 2 39 1870-71 Fig. .3. Running five-year means of tation years (I July- 30 June), for Cedai source: U.S. Weather Bureau. Precipitation Year F the percentage of average precipitation, based on precipi- le and Fort Bidsvell in Surprise Valley. California. Data 522 (,KEAT BASIN NATURALIST Vol. 33, No. 3 niesic species, but when drought should have hiuderod ils ostablishnieut. The notion that dry conditions might have favored tree invasion l)\ reducing the sagebrush-grass cover is untenai)le, be- cause a drought so severe as to decrease the xerii sagebrush toidd not possibh' siniuhaneouslv inc reas(> the niesic white fir. The coincidence of drought and initial establishment of the white fir in this study, then, suggests that the trees in- vaded in spite of the weather rather than because of it. Moreover, it is apj^arent that nonclimatic factors prevented tree invasion during the more moist periods existing in the study area prior to 1910. Fire supprkssion. — Although the Modoc Forest Reser^ e was established in 1904, wildfires in the Warner Mountains apparently continued to be connnon un- til after the creation of the Civilian Con- servation Corps in 1933; the year 1924, for example, is reported to have been a particularly serious fire year (Cook, n. d.). More specifically, information from the U.S. Forest Service indicates that the area immediately adjacent to the study plots was burned by wild fires l)etween 1921 and 1930, aiid that fires were com- mon within the national forest during that decade (U.S. Forest Service. 197-1). Supporting the impression that fire su]i- [)ression was not well developed in the region by the time of tree invasion, the rangeland outside of the forest resei-\'e, and in which the white fir became estab- lished, was not given official protection until passage of the Taylor Grazing Act in 1934. The Bureau of Land Manage- ment office in Susanville, California, out of which the federal rangelands in the region are administered, reports no rec- ords of fire sui)pression activities on the east slope of the Warner Mountains prior to 1930 (U.S. Bureau of Land Man- agement, 1974). The State of (California was not. and is not, res})onsible for fire control in the study area, although it presently protects private rangelands west of the Warner Mountains (California Division of For- estry, 1975). Moreover, the state did not have any system for fire supj)ression during the initial years of tree establish- ment, and its fire protection policies can- not be considered to have been effective until after 1943 (Clar, 1969; Davis, 1963). It seems \ali(l to conclude that suc- cessful fire suppression on the east slope of the southern Warner Mountains was not effective until after the period of initial tree establishment. Although sub- sequent fire control has probably aided the maintenance of the trees, it cannot be invoked to exjilain the initiation of tree advanc(^ dowiislope. Grazing. — The intensity of grazing by domestic livestock in the southeastern Warner Mountains has varied greatly since the initial settlement of Surprise Valley, inniiediately east of the Warner Range, in 1(S(>1. The number of cattle on raiichcs in Modoc County as a whole was little changed between 'lS90 and 1945, but it doid)letl in the following twenty years; resident sheep, by contrast, in- creased rapidly between 1890 and 1930, but by 1940 the\' had declined j)recip- itously (Fig. 4). Much of Modoc Count}' is heavily for- ested, thus restricting grazing by domestic livestock to areas of brush or grass. Such habitats in the Warner Mountains and ad- jacent A'alleys have supported large num- bers of animals, particularly sheep, over the last century. Contributing to the heavy grazing of these rangelands was the seasonal migration of sheep from the inountains in summer to the semiarid lowdands of northern Nevada in winter, a pattern well established by the 1870s (Olmsted, 1937). Moreover, sheep drives from Idaho and Oregon to shipping points in western Nevada jiassed through the Warner Mountains (Olmsted, 1937). Fig. !•. Niiinl)('i-> 1)1 uitll.t" and slu'cj) on i-iuicliofi in Mcxloc (bounty. Data soiirco: IT.S. Bureau of Census. Sept. 1975 IITF. FIU IN\-ASI()X 323 By 1900 the ranges of Modoc County are said to have been greatl\- overgrazed (Brow^i, 1951; Pease. 196-3). Establishment of f(>deral forest res(>r\('s (later to become national forests) in the early 19()0s apj^arently (hd littl(> to re- (hice innnediately the grazing j)ressures in the higher elevations of Modoc County, including the Warner Moun- tains. TYansient sheej) continued to be driven across national forest land from Oregon initil the Forest Service banned such use in 1914 (Tierney, 1946). Pease (1965) suggests that the elimination of grazing by transients promjited the estab- lishment of new sheep ranches, with resi- dent flocks, in Surjirise Valley; this con- tributed to the ra])id increase of resident sheep in Modoc County l^etween 1910 and 1920. Also accentuating the heavy grazing pressures at this tiine, the Forest Service intentionalh' allowed overstock- ing on national forest lands in north- eastern California during World W^ar I to help meet war demands for food and wool; even after the war, heavy stocking con- tinued because it was felt a]n-u])t reduc- tions in livestock numbers might ha^e created economic hardship for area ranchers (Tierney. 1946). Even while the national forest lands were under nominal regulation, the pub- lic domain continued to be completely free and open range. These latter federal lands in northwestern Nevada served, in part, as wintering grounds for sheep that were moved from California during the autumn season. Olmsted (1957) claims that 150,000 sheep were grazed in Surprise Valley in 1920, and. when com- j)ared to resident sheep rejiorted on ranches in all of Modoc Coimty in that year (109,000), it is apparent that much use of Surprise Valley ranges, including much ])ublic domain acreage, was by transient flocks. The peak in grazing pressure by sheep in the 1920s was ap- |)arently even greater than that suggc^sted by the numbers of resident sheej). The end of uncontrolled sheep grazing on the public domain came with the pas- sage of the Taylor Grazing Act in 1934 (Olmsted, 1957; Pease, 1965). With more stringent regulations against tran- sient flocks, the public lands coidd jio longer be used by migratory sheep herd- ers. The grazing history suggests that the white fir iii\asion (oinc ided with the ])eak. and tli(> period immediately fol- lowing the peak, (jf >lieep grazing in the region. Sheep may have reduced the cox- erage of grass and shrubs, thereb}' in- creasijig the a^■aiiabilit}' of soil moisture and allowing the establishment of seed- ling trees. Sheep browse shrubs, hut not conifers, on ^^ inter range, thus encour- aging the tree invasion. Conclusions The evidence suggests that grazing b}" domestic livestock altered the sagebrush- grass vegetation on the east slope of the Warner 1\ fountains and allowed the estab- lishment of white fir seedlings. Yet, while grazing seems responsible for the initia- tion of tree in^'asion, the accelerated rate of tree establishment during the period 1935-1944 may have been the result of increased precipitation on the openings in the vegetation cover ])roduced by grazing. Such an explanation would ac- count for the decrease in tree establish- ment after 1944 because, by that time, the plant cover would have sufficiently recovered from the effects of the earlier heavy grazing to minimize the avail- ability of suitable seedbeds and soil mois- ture. The cause of the present abundance of seedling trees is more obscure, but may be related to a resurgence of grazing pres- sure due to increasing numbers of cattle in recent years (Fig. 4). In the northern Intermountain West generally, grazing by domestic livestock may be adequate to explain the vsdde- spread invasion of trees into sagebrush- grass vegetation dialing the late nine- teenth and early twentieth centuries. The example from California, discussed in this paper, suggests that fire suppression has been too recent, except perhaps lo- cally, to correlate with these tree invas- ions. (Control of fire may account for tre(^ establishment in other vegetation types, however, during this time.) Re- gional (dimatic fluctuations, by them- selves, also seem inadequate to account for the (>x])ansion of trees into sagebrush- grass vegetation during the latter half of the 1800s, a time characterized by "fluc- tuating Init below axerage moisture" in western North America (Fritts, 1965). This portrayal of climate does not suggest conditions sufficiently wet to favor tree 324 (iKEAT BASIN NATUKALIST Vol. .35, No. 3 growth in formerly xeric brush. More- over, tree invasion in the Warner Moun- tains began, not during a wet period, but during an extended time of below average precipitation. Grazing by domestic live- stock, then, remains the most likely gen- eral cause, apjilicable on a regional basis, to account for the widespread ijivasion of sagebrush vegetation by tree species. Periods of extended drought may, in cer- tain ])laces, accentuate the plant Vover de- terioration caused by grazing, thus en- couraging tree estab'lishment once more moist conditions returii. Literature Cited Arnold, J.. D. J.\]\ieson, and E. Reid. 1964. The pinyon-iuniper typo of Arizona: ef- fects of grazing, fire and tree-control. U.S. Dept. Ag. Prod. Res. Rept. No. 84. 28 p. Blackburn, W.. and P. Tufxler. 1970. Pin- yon and juniper invasion in Ijlack sage- brush communities in east-central Nevada Ecol. 51:841-848. Brown, W. 1951. California Northeast: The bloody ground. Oaklaiifl. California. Bio- books. 207 p. Burkhardt. J., and E. Tisdale. 1969. Nature and successional status of western juniper vegetation in Idaho. .]. Range Manage 22- 264-270. California Division of Forestry. 1975. I^et- ter to author. CiAR. C. 1969. Evolution of California's Wild- land Fire ProtcM tion System. Sacramento. St^ate of Califonna. The Resources Agency. ?5 p. Cook. F, n. d. History of Modoc Counlv. Vol- cano. California. California Tiavelei- Inc 104 p.. CoTTAM, W.. AND G. Stewart. 1910. Plant suc- cession as result of grazing of meadow desic- cation by erosion since settlement in 1862 J. For. 38:613-626. Davis. L. 1965. The economics of wildlife pro- k'ction with emphasis on fuel break svstems Sacramento, State of California. The Re- sources Agencv. 166 p. Iuxster. ,J. 1917. The spread of timbered areas HI central Te.xas. ,J. For. 15:442-145. luuTTs. H. 1965. Tree-ring evidence for cli- matic ciumgcs in western North America Mon, Weather Rev. 93:421-443. .loiiNSEN. T. 1962. One-seed juniper invasion of northern An/.ona grasslands. Ecol. Monog. 32:187-207. ^ Oimsted, p. 1957. The Nevada-California-Ore- gon border triangle: a study in sectional his- tory. M.A. tliesis, Univ. Nevada, Reno. 284 p. Patten, 13. 1969. Succession from sagebrush to mixed conifer forest in the northern Rocky Mountains. Am. Midi. Nat 82- 229-240. Pe.\rson. G. 1931. Forest types in the south- west as determined bv climate and soil. U S Dept. Ag. Tech. Bull. 247. Pe.\se. R. 1965. Modoc County: a geographic time continuum on the California volcanic tableland. Cniv. California Publ. Geog Vol 17. 304 p. ^'' Robertson, J., and C. Pearse. 1945. Artificial reseeding and the closed communitv. North- west Sci. 19:58-66. Tierney. H. 1946. Modoc County: past and present. Alturas, California, Modoc County Superintendent of Schools. 139 p. ir.S. Bureau of Land Management. 1974 Letter to author. U.S. Forest Service. |i)7k Letter to author. Vankat, ,1. 1970. Vegetation cliange in Se- quoia National Park. California. Ph.D. dis- sertation. Univ. California. Davis. 197 p. Wright. .[.. and H. Fisser. 1968. Juniperus osteospeiiua in northwestern Wyoming: their (bstribution and ecology. Sci. Monograph No. 7. Laramie. Wyoming. LTniv. Wvoming Ag. Exper. Stat. 31 p. MORPHOLOGY OF EPHEMERAL AND J^ERSISTENT LEAVES OF TFIREE SUBSPECIES OF BIG SAGEBRUSH GROWN IN A UNIFORM ENVIRONMENT W. T. McDonough. R. (). Hainl (I I',, 15. Canipbe: Abstract.- Measui-ciiipnts uere made of inorj)h()li)gi( al ( liai a( tciistic s of ephenieral and j)er- sistent leaves of three subspecies of big sagebrush (Arlc/iiisia Iri/lr/ila/a Nutt.) grown in a uni- form environment. No morphological feature clearh' separaterl the subspecies. Genetic-environ- mental interactions on leaf morphology apparently reduce its \alue as a sole criterion for (hstin- guishing the subspecies. Taxonomic subdivisions of a species may differ in morphological and physio- logical characteristics, and these (hffer- ences may be genetically (jr einirf)n- mentally controlled to varying extents. There is considerable interest in subspe- cies of big sagebrush (Artemisia tridcnt- ata Nutt.) because of differences in palat- ability, habitats occuj^ied, and aggressive- ness-characteristics of interest to range managers fMcArthur et al., 1974). Iden- tification is based chiefly on the growth form and morphology of leases and on the inflorescence of plants collected from their normal habitats (Winward, 1970). Biochemical differences have also been noted in reproducti^•e and vegetative parts (Winward and Tisdale, 1969; Hanks and Jorgensen, 1973; Stevens and McArthur, 1974). Biochemical patterns are less subject to enviromnental mod- ification (Winward, A. H., 1975. Per- sonal communication. Oregon State Univ.. Corvallis) but are inconvenient for use in field indentification. To determine the extent to which leaf morphology of big sagebrush subs])ecies is genetically rather than environmen- tally controlled, we grew plants of three subspecies under the same greenhouse conditions to identify distinguishing leaf characteristics that persist in a uniform environment. Methods Potted year-old plants of each of three subspecies, basin big sagebrush (tridcnt- ata Nutt.), mountain big sagebrush (vasc- yana Rydb.), and Wyoming big sage- brush {wyomingensis Beetle), were estab- lished from seeds collected the fall of 1973 near the Sheep Ex})erimental Range, Du- 'Plant physiologist, range scientist, and range tcclmici and Range Experiment Station, Ogden, Utali 84401. stati tained in cooperation with Utah State University, Logan. bois, Idaho. The (25 cm) pots were ran- domly arranged on a greenhouse bench and given routine care. From the 50 ]jlants of each subspecies, 6 plants and 10 mature leaves (ephemeral and per- sistent; from each plant were randomly selected for determinations of length, width, length /width, number and depth of lobes. Lobes were distinguished from occasional minor dentations by the pres- ence of a secondary Acin. Leaves were fixed to cards, photographed, and the 35-mm transparencies projected for mea- surements. At the time of sampling, growing plants averaged 27 cm in height and had from 6 to 1 7 branches. Significance of dif- ferences at the 5 percent level was eval- uated by variance analysis and multiple range tests. Results Representative leaves are shown in Figure 1; mean dimensioius, in Table 1. Only limited differentiation was observed for both types of lea\(vs among the sub- species. In the e|)hemeral leaves, variation in lobe tium])er was not significant. Width and length/width differed only in subsp. vascyana. Subsp. wyomingensis differed from subsp. trident at a in lobe length and from subsp. vaseyana in lobe depth. In the jiersistent leaves, length dis- tinguishes subs]). wyomingensis from the other subspecies and length/width dis- tinguishes wyomingensis from tridentata. Number of lobes was uniformly three in all subs]iecies, and lobing generally was so shallow that attempted measurements were uiu'eliable. ui, respective! V. USiXV I'orest Service. Interniounlain P'orcl med in I.ogan. l.'tah. at l-"orestry Sciences Lal)or,'ilory. mam- 325 326 GREAT BASIN NATURALIST Vol. 35, No. 3 T.VBLE 1 . brush grown Mean dimensions' ± standard devi 1 a unifonn environment. of til iul)species of l)ig sage- Subspecies Length (mm) Width (mm) Length/Widtli Number of lobes Lobe deptli (mm) Ephemeral Leaves Tridcnta/a Vaseyana Wyomingeusis 50.9=1 ±7.1 47.9=1'' ±5.4 43.3'' ±4.7 19.3=1 15.6" 19.6=1 ±6.8 ±5.9 ±6.1 5.0=1 ±1,2 3.6" ± 1 .8 2.5=^ ±1.0 Persistent Leaves 4.0=1 3.8=1 4.0=1 ±1.1 ±0.9 ±1.0 9.8=1" ±^j 8.4" ±3.4 11.4=1 ±3.6 Tridentata Vaseyana Wyomingeusis 14.3=1 ±2.5 14.9=1 ±3.0 11.9>^ ±2.3 3.1=1 3.4=1 3.0=1 ±0.7 ±0.7 ±0.8 4.9=1 ±1.2 4.5=1" ±0.6 4.1" ±0.8 3=1 3=1 3=1 - 'Means for anv measurement same letter in superscript Fig. 1. Ephemeral (above) atid persistent (below) leaves of sagebiiish grown in a uniform environment — subsp. tridentata (T). vaseyana (V). and wyomingeusis (W). Wiiiward (197()j exaniiiiofl only per- sistent leaves of field-grown plants. Only differences in length/width were con- sidered usefnl in separating the subspe- cies. Values of 5.6, 4.0, and 3.1 were as- signed, respectively, to subsp. tridentata, vaseyana, and wyomingensis. I.ea^-es of plants grown under uniform conditions are in the same sequence but have some- what divergent values that do not differ- entiate subsp. vaseyana (Table 1). Conclusions Even with careful measurements, no characteristic of ephemeral or persistent leaA'es from plants growai under uniform conditions was found to clearly separate the three subspecies. Only subsp. vase- yana is distinguishable by two character- istics of ephemeral leaves and subsp. wyomingensis by one characteristic of persistent leaves. Distinguishing leaf mor- phology may be so dependent upon en- vironmental inodification that it is not uniformly useful in identification. Al- ternatively, distinct morphological dif- ferences may be obscured by genetic in- trogression among subspecies and hybrids (Plummer, A. P., 1975. Personal com- munication. U.S. Dep. Agric. For. Serv., Intermt. For. and Range Exp. Stn., Og- flen, Utah) or because insufficient num- bers of plants are sampled from the same or other areas. r.ITER VrURF. CiTF.n Hainks. I). I... \M) K. R. .JoRr.ENSEN. 1973. Chroniatograpliic identification of big sage- brush seed. ,1. Range Manage. 26:304. McArtiiur. E. D.. B. C. Giunta. and A. P. Pr.uMiMER. 1974. Shrubs for restoration of depleted ranges and disiurbed areas. Ltali Sci. 35:28-33. Stevens, R.. and E. D. McAruiur. 1974. A siini)le field teduiiciue for identification of some sagebrusii taxa. J. Range Manage. 27: '>25- 526. \\'iN\\ARD. A. H. l'>70. Taxonomy- and ecology of big sagebruslr Ph.D. Thesis. Univ. Idaho. Moscow. 80 ]). \\IN\VARD. A. H.. \.M) K. W. TlsDAI.E. 1969. .A simplified .hrninal iiiriho.l for sagebrusii id(Mitifi(.iti.)ii. I iiiv. Id.ih.. Stn. Note 11. 2 p. 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 directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. Manuscripts. Two copies are required, 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 this journal as a format, and the Council of Biological Editors 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, and written in accordance with Bio- logical Abstract guidelines, should precede the introductory paragraph of each article. Authors may recommend one or two reviewers of their article. Illustrations and Tables. All illustrations and tables should be made with a view toward having them appear within the limits of the printed ])age. Illustrations that form a part of an article should accompany the manuscript. Illustrations should be prepared for reduction by the printer to either a single-column (2% inches) or double- column (5^/2 inches) width, with the length not exceeding 71/2 inches. Costs Borne by Contributor. Authors should be prepared to contribute $10 per printed page toward publication of their article (in addition to reprint costs). No re- prints are furnished free of charge. A price list for reprints and an order form are sent with the proof. Reprint Schedule of the Great Basin Naturalist 2pp. 4pp. 6pp. 8pp. 10pp. 12pp. Each 100 copies $20 $23 $26 $29 $32 $35 Additional 200 copies 26 29 32 35 38 41 2pp. 300 copies 32 35 38 41 44 47 $3 TABLE OF CONTENTS Urosaurus and its phylogenetic relationship to Uta as determined by oste- ology and myology (Reptilia: Iguanidae). Charles Fanghella, David F. Avery, and Wilmer W. Tanner 245 Distribution and adundance of the black-billed magpie (Pica pica) in North America. Carl E. Bock and Larry W. Lepthien 269 Nectar composition of hawkmoth-visited species of Oenothera (Onagraceae). Robert E. Stockhouse, II 273 A revision of the nearctic species of Clinohelea Kieffer (Diptera: Ceratopo- gonidae). William L. Grogan, Jr. and Willis W. Wirth 275 Basidiomycetes that decay junipers in Arizona. R. L. Gilbertson and J. P. Lindsay 288 Body size, organ size, and sex ratios in adult and yearling Belding ground squirrels. Martin L. Morton and Robert J. Parmer 305 Photoperiodic responses of phenologically aberrant populations of pierid butterflies (Lepidoptera). Arthur M. Shapiro 310 Additional records of reptiles from Jalisco. Me.xico. Philip A. Medica. Rudolf G. Arndt. and James R. Dixon 317 Invasion of big sagebrush (Artemesia Iridentata) by white fir {Abies con- color) on tlie southeastern slopes of the Warner Mountains, California. Thomas R. Vale --- 319 Morphology of ephemeral and persistent leaves of three subspecies of big sagebrush grown in a uniform environment. W. T. McDonough. R. O. Harniss, and R. B. Campbell 325 IE GREAT BASIN NATURALIS me 35 No.4 December 31, 1975 Brigham Young Universi ■OOL ** •^1 - ^iSF^' ^^Sl^ ^^-y^W GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Zoology; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, dean. College of Biological and Agricultural Sciences; Ernest L. Olson, director, Brigham Young University Press, University Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been continuously published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only original, previously unpublished manuscripts pertaining to the biological natural history of the Great Basin and western North America will be accepted. Manuscripts are subject to the approval of the editor. Subscriptions. The annual subscription is $9 (outside the United States $10). The price for single numbers is $3 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 Press, Marketing Department, 204 UPB, Provo, Utah 84602. Scholarly Exchanges. Libraries or other organizations interested in obtaining this journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Lih-arian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts and other copy for the Great Basin Naturalist should be addressed to the editor as instructed on the back cover. The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University Volume 35 December 31, 1975 No. 4 ENDANGERED, THREATENED, EXTINCT, ENDEMIC, AND RARE OR RESTRICTED UTAH VASCULAR PLANTS Stanley L. Welshi, N. Duane Atwood-, and James L. ReveaP Abstract. — The status of 382 vascular plant taxa with distribution in Utah is presented. Some 66 species are possibly endangered, 198 threatened, 7 extinct, and 20 extirpated within the state; 4 spe- cies have questionable taxonomic status. Included in the list are nearly 225 species of endemic plants, many of which are among the possibly endangered, threatened, and extinct or extirpated plants. Bibliographic citations, type locality, status, and distribution by counties is included for each species or infraspecific taxon. Two new species are described: Psoralen pariensis and Eriogonum natum. One new name. Astragalus barnebyi, is proposed; and one new variety, Eriogonum umbel- latum var. deserticum, is proposed. The following new combinations are made: Cycladenia hum- ilis var. jonesii; Aralia racemosa ssp. bicrenata; Heterotheca jonesii; Hymenoxys depressa; Xantho- cephalum sarothrae var. pomariense; Thelypodium integrifolium var. complanatum; Thely podium sagittatum var. ovalifolium; Arenaria kingii var. plateauensis; Psorothamnus thompsonae; Najas caes- pitosus; Oenothera gouldii; Eriogonum corymbosum var. revealianum; Penstemon humilis var. ob- tusifolius; Penstemon lentus var. albiflorus; and Viola purpurea var. charlestonensis. The vascular plant flora of Utah is both large and complex. Its components are diverse, representing numerous flor- istic elements from many parts of North America, including unique and provin- cial elements restricted to the state. Spe- cies of many major geographical group- ings of plants occur within the multiplic- ity of habitats available within Utah, all to a greater or lesser degree of their entire range. Some of these species are at the edge of their total range, and these occur in smaller portions of the state. Other taxa occur only in one or few peculiar, limited edaphic situations or habitats, while others are more widespread and cover a broad altitudinal or latitudinal expanse of Utah. Those plants that occur only within the state, or within the natural basins that overlap the artificial political boundaries of the state, are known as local endemics. Their range can be widespread within Utah, but more often they are restricted in distribution to very limited areas. These are plants that are of much inter- est to scientists, because they present liv- ing proof of the origin and evolution of species, origin of floras, and indications of relationships of plant species. These plants are those entities which have orig- inated here or are mere remnants of spe- cies which have had a much broader area of distribution in the past. The impress of man and his activities onto the natural habitats of Utah has re- duced the area available to most native or indigenous plants. Those species of broad extent and wide ecological toler- ances have withstood these activities best, with only a reduction in their range and number. Less frequently their range has increased in size as less-tolerant plants of adjoining areas have been reduced. How- ever, many of the most unique species have areas of distribution that are very small, with only a few known individuals. In some examples the species have ap- parently ceased to exist in the Utah flora. In most cases the new habitats made ^Department of Botany, Brigham Young Universily, Provo, Utah 84602. -Bm-eau of Land Management, Cedar City, Utah 84720. ^Departments of Botany, University of Maryland, College Park, Maryland 20742, and National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560. 327 328 GREAT BASIN NATURALIST Vol. 35, No. 4 available by the acthdties of man have been occupied by introduced, cultivated, and adventive plants from the Old World. These are the crop plants and weeds of modern agricultural and industrial so- ciety. The phalanxes of intolerant native plants have been retreating under the im- pacts of agriculture and grazing for more than a century. Industrial development during most of that period was limited in extent, if not in effect. In Utah most of these activities were restricted mainly to the broad valleys and river basins, where plant communities that were rela- tivel}^ fragile are now almost totally lacking within the state. However, these sites apparently contained few of the nar- rowly endemic plant species. With the advent of the second half of the twentieth century, there has occurred a resurgence of economic activities, mineral explora- tion, and a greater use of the public lands which hitherto had been considered as useful (if considered useful at all) only for grazing and watershed. The rapid spread of industrial development into pre- viously undeveloped, low-elevation, arid lands in the southern portions of the state is all the more impressive when one con- siders that most of the narrowly restric- ted plants occur in those areas (Figs. 1 and 2). Plant species which were once remote from the impacts of civilization — industrial, agricultural, or recreational ac- tivities— are now threatened not only by the effects of ranching, construction, and off-road travel, but even by the very ageiicies of government which are estab- lished by law to oversee in the public trust the proper use and protection of the public lands. At the present time, hardly a part of Utah, even that set aside as national parks, monuments, or wilderness areas, is safe from degradation by masses of people or by those seeking to exploit the very natural resources and features these unique areas were established to protect. Inroads into the most remote and most arid portions of the state now guarantee further reduction of the unique flora of Utah. Naturally, those entities that will suffer greatest from the commercializa- tion of the state will be those which have specific and naturally restricted areas of Fig. 1. Distributional incidence of endangered, threatened, extinct, or extirpated Utah plants by county; total number is greater than the total for the state due to overlap. Fig. 2. Distribution of endangered, threatened, extinct, and rare or restricted Utah plants, by phytogeographical subdivision. Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 329 distribution. Only the most enlightened management and protection from un- reasonable exploitation will ensure the continuation of the rich indigenous flora that these plants represent. Commerical exploitation need not bring about the demise of species, although the basic nature of the flora will change. It is a fact of biology that in even the most enlightened and carefully planned devel- opment, there will be a reduction in the native vegetation. Reclamation attempts will be made with Old World introduc- tions, or with selected ecotypes of indig- enous plants, and not with those that oc- curred in the region prior to its modifi- cation. Both of these types of plants can hardly replace the quality of those cleared and destroyed by construction activities. Revegetation of a disturbed site can be more or less productive from an economic standpoint, but natural plant succession will require generations before any real resemblance of the natural vegetation will again be possible, and then, depend- ing on the nature of the disturbance, the native vegetation occupying the site can be of an entirely different composition than formerly. Indigenous taxa with specific habitat requirements will not survive if those habitats are altered, and there is no known technology which can simulate some of the peculiar habitats present in the state — which are now oc- cupied by narrowly restricted plants — and no technology can ever replace an extinct species (Reveal 1973b). Because of the rate at which Utah is being impressed by all the forces of a modern society, it seems important that the endangered, threatened, extinct, en- demic, and rare or restricted vascular plants of Utah be inventoried, and that their known or historic areas of distribu- tion be outlined. That task is the basic goal of this paper, but this is only a pioneering effort. Much work remains to be done. While the present paper was in prep- aration, the secretary of the Smithsonian Institution, S. Dillon Ripley, transmitted to the Congress of the United States a "report on endangered and threatened species of the United States." This report (cited herein as "Ripley 1975") lists num- erous plants from Utah as either threat- ened, endangered, possibly extinct, or probably extinct. Each category was ab- breviated as T, E, PoEx, or PrEx, respec- tively. That report was prepared during the calendar year of 1974 as mandated by the Endangered Species Act of 1973 (Pub- lic Law 93-205) in which the secretary of the Smithsonian was to report to Congress within one year on all of the "species of plants which are now or may become en- dangered or threatened" in the United States (sec. 12). The Congress provided no funding for the preparation of this re- port, and the time restrictions prevented a detailed field examination of all spe- cies included in the list. Some states, notably California and Texas, with active offices concerned with endangered plant species, were able to provide precise data; other states were able to supply some in- formation, and still others had little or no input into the final report, except that gathered by the committee established by the secretary to prepare a statement for the Congress. The data for the state of Utah was provided by a number of indi- viduals (including the authors of this paper, and Reveal served on the Smith- sonian committee), but time did not per- mit the type of critical examination of each taxon which has been largely pos- sible for the present paper. The Smithso- nian report (Ripley 1975) was published in the Federal Register (Schreiner 1975) as a "notice of consideration," and we understand that the Department of In- terior will submit a revised listing in the near future. Disagreement as to the desig- nation of degree of endangerment be- tween this paper and that of Ripley (1975) represents the results of a more detailed and concentrated survey of the Utah flora, and is based on detailed literature search and personal information of the present authors and that of their col- leagues who have reviewed the manu- script. These differences have largely been resolved and will appear in the Smithso- nian's revised list to be submitted early in 1976 to the Secretary of the Interior. Much of the information has been ac- quired through many years of investi- gation of the Utah flora, both in the field and in the herbarium, and has been stim- ulated by the Ripley (1975) report. The designation of the status of an in- dividual taxon as belonging to a partic- 330 GREAT BASIN NATURALIST Vol. 35, No. 4 ular category is subjective. Still, it is based on the best information available to us at the present time (a requirement of the Endangered Species Act). A plant species is considered as "endangered" when its known area of distribution is very small, and when the expected de- velopment or exploitation of the area oc- cupied has already occurred or is immi- nent. A "threatened" plant is one of some- what larger known areal extent, but which has experienced or is now exper- iencing a reduction of its natural distribu- tional area. These are the two major cat- egories required by law to be designated. Plants that are "rare or restricted" are those which have been collected only oc- casionally or which are known from very limited regions of the state; they may or may not be widely distributed elsewhere outside the political confines of Utah. In- troduced species are excluded from this category, even though they might be re- stricted or even rare. Plants considered to be "endemic" are those whose entire distributional area is within Utah or within one of the natural drainage basins which overlap the politi- cal boundaries of the state. Some plants in this category especially are poorly known taxonomically and biologically, and in these cases we have so indicated the need for a careful systematic evalu- ation of the taxon to determine the status of the plant entity. All statements about status of those plants not endemic to Utah are with re- gard to the occurrence of those plants within Utah. In many, if not most of these cases, the species are widespread and com- mon to abundant in other portions of their area of distribution. If they are limi- ted, rare, or possibly extirpated from Utah, only that portion of their range is considered in making the designation. This follows the guidelines established by California and Texas in which the en- dangered and threatened lists are based solely upon the situation of the plant in question within the confines of the state boundaries (Table 1). Plants listed in one of the categories designated above by Ripley (1975) are included here, whether or not they are considered as something other than threat- ened, endangered, or extinct on the new list that will be published in 1976. Ob- vious errors, which will not be repeated in the future, such as Lewisia maguirei and Penstemon decurvus, to mention only two, are excluded. Voucher specimens for some of the re- ports are cited in the distribution state- ments (e.g., Harrison 6370, for Cymop- terus basalticus) . Bibliographic citations in support of distributional data are in- cluded for many species, especially for those which are obscure or are poorly rep- resented in herbaria. We are not making any attempt to provide precise location data (except as might be obtained from published type localities) in fear of com- mercial exploitation of some plants (i.e., catci and orchids) and because of the pos- sibility of destruction of selected popula- tions by those having a vested interest in ridding parcels of land of any species of plant that might fall under the protection of the provisions of the Endangered Spe- Table 1. Numerical summary of the endan- gered, threatened, extinct, extirpated and rare or endemic species in Utah. o T3 t 0) 0) 'S 1 60 CO 0) c 1 -o U u S JS c« County W H M w tf Beaver 1 7 0 1 11 Box Elder 1 5 0 0 1 Cache 2 8 0 0 3 Carbon 2 6 0 0 11 Daggett 3 9 0 1 4 Davis 0 0 0 0 4 Duchesne 3 9 1 1 14 Emery 4 19 1 0 28 Garfield 9 39 1 1 38 Grand 6 17 0 1 30 Iron 2 15 1 0 9 Juab 0 7 0 3 10 Kane 12 31 0 4 26 Millard 2 9 1 0 14 Morgan 0 0 0 0 0 Piute 0 12 0 1 8 Rich 1 1 0 0 1 Salt Lake 0 8 0 1 8 San Juan 7 25 0 2 27 Sanpete 1 6 1 0 9 Sevier 2 13 0 0 7 Summit 2 3 0 3 6 Tooele 0 3 0 3 7 Uintah 7 17 1 1 14 Utah 1 7 0 1 13 Wasatch 0 1 1 0 5 Washington 12 41 0 3 24 Wayne 5 14 0 0 29 Weber 0 7 0 0 4 Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 331 cies Act of 1973. Nevertheless, those agen- cies charged with protection of endan- gered or threatened plants must deter- mine the precise localities of those entities in order to plan for their survival. Maps are presented (Figs. 1 and 2) which demonstrate the unequal distribu- tion of these unusual plants. In Figure 1 the number of taxa known of each county is indicated. The total appears to be very large, but this is due to many of the spe- cies being listed in several counties when area of distribution occurs outside a single county. Phytogeographic regions of Utah are plotted in Figure 2, and the number of taxa considered to be unusual is cited for each. The number of taxa totals are less than for the entire state because some dis- tributional data are so indefinite as to not be plotable. Also, the figures represent an attempt to plot plants within their main area of distribution; double representation has been avoided. This accounts for the ap- parent discrepancies between the two maps. Recommendations Land use planning should take into ac- count the presence of the unusual plants present in the state of Utah. If and when the Secretary of Interior proclaims any species of plants found in Utah as en- dangered or threatened, they will fall un- der the protective provisions of the En- dangered Species Act which, in section seven, calls upon all federal departments and agencies "to insure that actions auth- orized, funded, or carried out by them do not jeopardize the continued existence of such endangered species and threatened species or result in the destruction or modification of habitat of such species." Those which are protected by law must be determined, and their areas of distri- bution should become known to the agen- cies responsible for their protection. Questionable taxonomic units should be investigated in order to determine their nature and area of distribution. Plants thought to be extinct should be sought in carefully coordinated field studies. The results of these studies should be reported to the Secretary- of the Interior so that in- formation about the various species may be updated from time to time, and so the status of each taxon can be reviewed. When new species are described from Utah, their status should be evaluated, and if they should prove to be endangered or threatened, that information should be presented to the Secretary so that such species may be considered for the federal endangered species list. In a recent issue of the Federal Register, Green wait (1975) proposed a "rule- making" for various amendments to the Endangered Species Act of 1973. These amendments introduce the term "plant" into many sections of the act. However, as Lachenmeier (1974) has pointed out, there are several legal and constitutional questions about the act that need to be resolved still, and from a botanical point of view, there still exist certain discrim- inatory differences between "wildlife" and "plants." These must be resolved in the future. In and for the state of Utah, we recom- mend that a review board be appointed to oversee the protection of Utah's most unique plants, and funding should be forthcoming to allow a real understand- ing not only of the endangered and threat- ened species, but the entire state's flora. The state should adopt the federal act and include within its own listing those spe- cies of the state which are endangered or threatened within its borders. Policies should be adopted that will determine basic land use with regard to these unique plants, and "critical habitats" as provided by the act should be investigated through- out the state (see also Green wait & Geh- ringer 1975). Above all, prompt actions must be taken by the state to preserve and protect the state's unique flora, and the members of the botanical community must be ready to aid and assist all levels of government in this important endeavor. Note In the following list, we are abbrevi- ating journals according to the abbrevi- ations listed by Lawrence et al. (1968) and herbaria according to listing given by Holmgren and Keuken (1974). We have attempted to give the type information as published in the original place of publica- tion; however, we are not attempting to typify any name, nor is the herbarium cited considered a statement of lectotyp- ification. This information is given for the convenience of the reader. For the most part, the abbreviations of the authors fol- 332 GREAT BASIN NATURALIST Vol. 35, No. 4 low the unpublished suggestions of the Index Kewensis staff. Acknowledgments The authors wish to acknowledge the critical reviews supplied by our colleagues in the study of Utah's plant taxonomy. Especially important help and criticism was supplied b}' Arthur H. Holmgren, Lois Arnow, and Andrew H. Barnum. Apiaceae Angelica wheeleri S. Wats., Amer. Nat- uralist 7: 301. 1873. Type: Utah, Wheeler s.n. (gh). Status: Endemic, evidently rare and local, possibly threatened. Distribution: Northern and central Utah (Mathias & Constance 1945). Cymopterus basalticus M. E. Jones, Contr. W. Bot. 12: 6. 1908. Type: Half-way Station, west of Wa Wa, Millard or Beaver Co., Utah, 7.000 feet, 15 May 1906, M. E. Jones s. n. (pom). Status: Restricted and locally abun- dant, neither threatened nor endan- gered (Ripley, T) . Distribution: Western Utah (Mil- lard Co., Harrison 6370; Matthews 14 [bry]), and adjacent Nevada (Math- ias & Constance 1945). Cymopterus coulteri (M. E. Jones) Math- ias, Ann. Missouri Bot. Gard. 17: 276. 1930. based on C. corrugatus var. coul- teri M. E. Jones, Contr. W. Bot. 12:19. 1908. Type: Juab, Juab Co., Utah, 4,000 feet, 30 Apr 1880. M. E. Jones 1691 (us). Status: Endemic, rare and restric- ted, threatened (Ripley, T). Distribution: Western Utah (San- pete Co., Mabey 5300 [bry]). Cymopterus duchesnensis M. E. Jones. Contr. W. Bot. 13: 12. 1910. Type: Among loose rocks on south- ern slopes of mesas, Myton, Duchesne Co., Utah, 5,000 feet, 20 May 1908, M. E.. Jones s.n. (pom). St.\ti:s: Endemic, rare and restric- ted, threatened (Ripley, E). Distribution: Duchesne and Uintah counties, Utah {Welsh 180 [bry]): Holmgren & Reveal 1887 [ny, utc]; Holmgren & Holmgren 5169 [bry, NY, utc]). Cymopterus jonesii Coult. & Rose, Rev. N. Amer. Umbell. 80. 1888. Type: Frisco, Beaver Co., Utah, 8,000 feet, 22 Jun 1880, M. E. Jones 1808 (US). Status: Rare and restricted. Distribution: Southwestern Utah and Nevada (Mathias & Constance 1945). Cymopterus higginsii Welsh, Great Basin Nat. 35: 377. 1976. Type: Shadscale dominated bajada, on gravelly pedimental fan east of None Butte, ca 17 miles east of Glen Canyon City, Kane Co., Utah, 31 May 1975, 5. L.^ Welsh 12740 (bry). Status: Endemic, local; possibly threatened. Distribution: Eastern Kane Co., Utah. Cymopterus minimus (Mathias) Mathias, Brittonia 2: 245. 1936, based on Aulo- spermum minimum Mathias, Ann. Mis- souri Bot. Gard. 17: 353. 1930. Type: On the upper part of the "Breaks" at Cedar Breaks, Iron Co., Utah, ca 10,500 feet, Mathias 723 (mo). Status: Endemic, rare and restric- ted to Cedar Breaks, possibly endan- gered (Ripley, E). Distribution: Cedar Breaks, Iron Co., Utah. Cymopterus newberryi (S. Wats.) M. E. Jones, Zoe 4: 47. 1893, based on Peu- cedanum newberryi S. Wats., Proc. Amer. Acad. Arts 11: 145. 1876. Status: Widespread and at least lo- cally abundant, neither threatened nor endangered (Ripley, T). Distribution: Garfield, Grand, Kane, Millard, San Juan, Uintah, Washington, and Wayne counties, Utah (bry, utc), and northern Arizona (Mathias & Constance 1945). Cymopterus rosei M. E. Jones, Contr. W. Bot. 12: 17. 1908. Type: Richfield, Sevier Co., Utah, 18 Jun 1898, M. E. Jones 30 (us). Status: Endemic, rare and local, pos- sibly threatened (Ripley, T) . Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 333 Distribution: Iron, Sanpete, Sevier, and Washington counties, Utah {Ma- guirc 1913U Holmgren 10932 [utc]). Ligusticum porteri Couh. & Rose var. brevilobum (Rydb.) Mathias & Con- stance, Bull. Torrey Bot. Club 68: 123. 1941, based on L. brevilobum Rvdb.. Fl. Rocky Mts. 613, 1064. 1917. " Type: Aquarius Plateau, Utah :5 Aug 1905, Rrdberg & Calrton 7473 (nyj. Status: Endemic, rare and local; possibly threatened. Distribution: Aquarius Plateau, Garfield Co., Utah (Mathias & Con- stance 1941). Lomatium latilobum (Rydb. ) Mathias, Ann. Missouri Bot. Card. 25: 281. 1937, based on Cynomarathriim latilobum Rydb., Bull. ToiTey Bot. Club 40: 73. 1913. Type: Proposed dam site, near Wil- son Mesa, Grand Co., Utah, Rydberg & Garrett 8371 (ny). Status: Endemic, rare and local, possibly threatened. Distribution: Grand Co., Utah (bry,- utc). Lomatium megarrhizum (A. Nels.) Mathias, Ann. Missouri Bot. Gard. 25: 282. 1937, based on Peucedanum megar- rhiza A. Nels., Bull. Torrev Bot. Club 26: 130. 1899. Status: Rare and local, possibly threatened. Distribution: Emery Co., Utah {Higgins 1268 [bry]) and southwest- ern Wyoming (Mathias & Constance 1945; Higgins 1972b). Lomatium minimum Mathias, Ann. Mis- souri Bot. Gard. 25: 273. 1937. Type: Near the hotel, on dr}' slopes bordering Bryce Canyon, Garfield Co.. Utah, 8,600 feet, Mathias 670 (mo). Status: Endemic, local, possibly threatened (Ripley, T) . Distribution: Garfield, Iron, and Kane counties, Utah (bry, utc). Lomatium scabrum (Coult. & Rose) Ma- thias, Ann. Missouri Bot. Gard. 25: 275. 1937, based on Cynomarathrum scab- rum Coult. & Rose, Contr. U.S. Natl. Herb. 7: 247. 1900. Type: Frisco, Beaver Co., Utah, 2,400 meters, Jones 1864 (us). Status: Endemic (but possibly in Mohave Co., Arizona), locally common, not threatened nor endangered. Distribution: Beaver, Iron (ut), Millard, and Washington counties. Utah (bry) . Musineon lineare (Rydb.j Mathias, Ann. Missouri Bot. Gard. 17: 265. 1930, based on Daucophyllum lineare Rydb., Bull. Torrey Bot. Club 40: 69. 1913. Type: Near Logan, Cache Co., Utah, 9 Aug 1895, Rydberg s.n. (ny). Status: Endemic, possibly threat- ened. Distribution: Cache Co., Utah (Holmgren 3595 [utc]). Apo(;yn.\ceae Cycladenia humilis Benth. var. jonesii (Eastw.) W^elsh & Atwood, stat. et comb, nov., based on C. jonesii Eastw., Leafl. W. Bot. 3: 159. 1942. Type: San Rafael Swell, Emery Co., Utah. 19 May 1914, M. E. Jones s.n. (C.'^S). Status: Endemic, rare and endan- gered (Ripley, E). Distribution: Emery and Grand counties, Utah. This entity is known in Utah from three basic collections: the type as cited above, a collection by Cottam (ut) also from the San Rafael Swell, and a collec- tion from Castle Valley. Grand Co., Utah (Welsh 1970b). Araliaceae Aralia racemosa L. ssp. bicrenata (Woo- ton & Standley) Welsh & Atwood, Stat. & comb, no^ ., based on A. bicre- nata Wooton & Standley, Contr. U.S. Natl. Herb. 16: 157. 1913. Status: Rare; restricted to Zion Narrows, Washington Co., Utah. Distribution: Washington Co.. Utah {Welsh 12366 [bry] ), and from south- eastern Canada south to Georgia, west- ward to Arizona and northern Mexico (Kearney & Peebles 1951); the sub- species is from Utah and Arizona. ASCLEPIADACEAE Asclepias cutleri Woodson, Ann. Missouri Bot. Gard. 26: 263. 1939. Status: Rare, threatened. 334 GREAT BASIN NATURALIST Vol. 35, No. 4 Distribution: Grand {Cottam 5799 [bry] and San Juan {Holmgren 3144 [uTc]) counties, Utah, and from north- eastern Arizona. Asclepias engelmanniana Woodson, Ann. Missouri Bot. Card. 28: 207. 1941. Status: Rare. Distribution: Grand (Rydberg & Garrett 8504 [ut]) and San Juan (Welsh et al. 2930 [bry]) counties, Utah; from Nebraska to Texas and Arizona. Asclepias labriformis M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 708. 1895. Type: Capital Wash, near the Henry Mts., Wayne Co., Utah, 5,000 feet, in sandy gulch, 19 Jul 1894, M. E. Jones 5650 (pom). Status: Endemic to the state but common, neither threatened nor en- dangered. Distribution: Emery, Garfield, San Juan, Uintah, and Wayne counties, Utah (bry). Asclepias ruthiae Maguire & Woodson, Ann. Missouri Bot. Gard. 28: 245. 1941. Type: Frequent, sandy soil, vicinity of water tanks, Calf Spring Canyon, 5,000 feet, San Rafael Swell, 18 miles southeast of Castle Dale. Emery Co., Utah, 10 May 1940, Maguire & Ma- guire 18310 (uTc). Status: Endemic, common in dis- junct populations on the San Rafael Swell; possibly threatened (Ripley, T). Distribution: Emery (Cottam 5500, Harrison 8067, Higgins & Reveal 1285a [bry], Grand [see Woodson 1954), and Wayne (Harrison 11223 and 11891 [bry]) counties, Utah. Asteraceae Chamaechaenactis scaposa (Eastw.) Rydb., Bull. Torrey Bot. Club 33: 156. 1906, based on Chaenactis scaposa Eastw., Zoe 2: 231. 1891. Status: Restricted, rare except lo- cally, not threatened nor endangered. Distribution: Carbon, Duchesne, Emery, San Juan, Uintah, and Wayne counties, Utah, and western Colorado (bry. utc). Cirsium rydbergii Petrak, Beih. Bot. Centralbl. 35 (2): 315. 1917. Type: Along the San Juan River near Bluff, San Juan Co., Utah, 1,200- 1,500 meters, 1-2 Jul 1911, Rydberg 10001 (g?). Status: Habitat specific, in hanging gardens, restricted and possibly threat- ened. Distribution: Garfield, Grand, Kane, San Juan, and Wayne counties, Utah (bry), and Black Mesa, Apache Co., Arizona (Howell & McClintock 1960). Enceliopsis nutans (Eastw.) A. Nels., Bot. Gaz. 47: 433. 1909, based on Encelia nutans Eastw., Zoe 2: 230. 1891. Status: Habitat specific, in heavy clay soils, restricted and rare. Distribution: Carbon, Duchesne, Emery, Grand, Uintah, and Wayne counties, Utah (bry, utc), and western Colorado (Harrington 1954). Erigeron abajoensis Cronq., Brittonia 6: 168. 1947. Type: Abajo Mts. (eastern range), 3,000-5,300 meters, San Juan Co., Utah, 17 Aug 1911, Rydberg & Garrett 9755 (ny). Status: Endemic, rare and threat- ened (Ripley, T). Distribution: Garfield, Kane, and San Juan counties, Utah (bry). Erigeron arenarioides (D. C. Eaton) Rydb., Fl. Rocky Mts. 1067. 1917, based on Aster arenarioides D. C. Eaton ex A. Gray, Proc. Amer. Acad. Arts 8: 647. 1873. Type: Rocky gulch above Cottonwood Canyon, Wasatch Mountains, Salt Lake Co.,' Utah, 8,000-9,000 feet, Aug 1869, Waston 547 (us). Status: Endemic and rare, restric- ted to rock crevices, threatened. Distribution: Salt Lake, Tooele, Utah, and Weber counties, Utah (bry, UTC; Cronquist 1947). Erigeron cronquistii Maguire, Brittonia 5: 201. 1944. Type: Cliffs, north side of Logan Canyon l^ mile below forks. Bear River Range, Cache Co., Utah, 5,800 feet, 20 May 1939, Maguire 16681 (ny). Status: Endemic, rare and threat- ened (Ripley, T). Distribi'tion: Cache Co.. Utah (Cronquist 1947). Dec. 1975 WELSH, ET AL.: ENDANCJERED UTAH PLANTS 335 Erigeron garrettii A. Nels., Manual Bot. Centr. Rocky Mts. 526. 1909. Type: Big Cottonwood Canyon, Salt Lake Co., Utah. 28 Jun 1905, Garrett 1310 (rm). Status: Endemic, rare and possibly threatened (Ripley, T). Distribution: High altitudes in Box Elder (ut). Salt Lake and Utah coun- ties, Utah (bry; Cronquist 1947). Erigeron flagellar is A. Gray var. triloba- tus Maguire ex Cronq., Brittonia 6: 258. 1947. Type: Canyon to Cedar Breaks, 12 miles east of Cedar City, Iron Co., Utah, 5 Aug 1934, Maguire 14947 (ny). Status: Endemic and rare, endanger- ed (Ripley, E). Distribution: Iron Co., Utah; known only from the type locality (Cronquist 1947). Erigeron kachinensis Welsh & Moore, Proc. Utah Acad. Sci. 45: 231. 1968. Type: Hanging gardens and seeps, near Kachina Natural Bridge, Natural Bridges N.M., San Juan Co., Utah, 13 Aug 1963, Welsh & Moore 2398 (bry). Status: Endemic, rare and endan- gered (Ripley, E). Distribution: San Juan Co., Utah; known only from the type locality (Welsh & Moore 1968). Erigeron maguirei Cronq., Brittonia 6: 165. 1947. Type: Dry rocky sandy canyon bot- tom. Calf Spring Wash, 1.5 mile up San Rafael Swell, 16 Jun 1940, 5,500 feet, Maguire 18459 (ny). Status: Endemic, rare and possibly extinct (Ripley, E). Distribution: Emery Co., Utah; known only from the type locality. Erigeron mancus Rydb., Fl. Rocky Mts. 902, 1067. 1917. ' Type: LaSal Mts., Grand or San Juan Co., Utah, 7 Jul 1911, Rydberg & Garrett 8671 (ny). Status: Endemic, rare and possibly threatened (Ripley, T) . Distribution: LaSal Mts., Grand and San Juan counties, Utah (Cronquist 1947.) Erigeron religiosus Cronq., Brittonia 6: 258. 1947. Type: Clear Creek, Zion N.P., Wash- ington Co., Utah, 8 Sep 1938, Eastwood 8z Howell 6339 (cas). Status: Endemic, rare and endan- gered (Ripley, E). Distribution: Kane [Harrison 11085 [bry] and Washington counties, Utah (Cronquist 1947). Erigeron sionis Cronq., Brittonia 6: 258. 1947. Type: Zion N.P., Washington Co., Utah, 1-3 Aug 1925, Pilsbry s.n. (ph). Status: Endemic, very rare and en- dangered (Ripley, E). Distribution: Washington Co., Utah; known only from type locality. Flaveria campestris J. R. Johnston, Proc. Amer. Acad. Arts 29: 287. 1903. Status: Rare, restricted, but not ap- parently threatened or endangered. Distribution: Grand Co., Utah (Ma- guire 1937; Harrison et al. 1964); Colo- rado, New Mexico and northern Mex- ico. Gaillardia flava Rydb., N. Amer. Fl. 34: 139. 1915. Type: Lower Crossing of the Price River (see Jones 1965), Emery Co., Utah, 2 Jul 1898, M. E. Jones 6412 (us). Status: Endemic, rare and endan- gered. Distribution: Emery Co., Utah (Cottam & Hutchings 2176 [bry]). Gaillardia spathulata A. Gray, Proc. Amer. Acad. Arts 12: 59. 1876. Type: Rabbit Valley, Wayne Co., Utah, 7,000 feet, 1875, Ward s.n. (gh). Status: Endemic, common through- out its range, neither threatened nor en- dangered. Distribution: Carbon, Emery, Gar- field, Grand, and Wayne counties, Utah (bry, utc). Haplopappus scopulorum (M. E. Jones) Blake in Tidestrom, Contr. U.S. Natl. Herb. 25: 546. 1925, based on Bigelovia menziesii var. scopulorum M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 692. 1895. Type: Near Cedar City, Iron Co., Utah, 6,000 feet, 10 xMay'l894, M. E. Jones 5204v is the first of two collec- tions cited (pom). Status: Uncommon, habitat specific, 336 GREAT BASIN NATURALIST Vol. 35, No. 4 neither endangered nor threatened (Ripley, T). Distribution: Iron, Kane (?), and San Juan (Welsh & Moore 2447; Welsh 8813; Atwood 4100 [bry]) counties, Utah, and Coconino Co., Arizona (Mc- Dougall 1973). Helianthus anomalus Blake, J. Wash. Acad. Sci. 21: 333. 1931. Type: Desert between Hanksville and Henry Mts., Wayne Co., Utah, 5 Jul \9^0^, Stanton 4806 (us). Status: Restricted to dunes and sandy washes, where locally abundant, neither threatened nor endangered. Distribution: Emery, Garfield, Grand, Juab, Kane, Millard, San Juan, Tooele, Uintah, Washington, and Wayne counties, Utah, and northern Arizona (Blauer 1966). Helianthus canus (Britton) Wooton & Standley, Contr. U.S. Natl. Herb. 6: 190. 1913, based on Helianthus petiolar- is var. canus Britton, Mem. Torrey Bot. Club 5: 334. 1894. Status: Rare, possibly threatened. Distribution: San Juan Co., Utah (Cottam 2544' [bry] ), and from western Texas to southern California and north- ern Mexico. Helianthus deserticolus Heiser, Proc. In- diana Acad. Sci. 70: 209. 1961. Type: 3.3 miles west of Hurricane, Washington Co., Utah, 29 Jun 1957, Stoutamire 2574 (ind). Status: Rare, restricted and possibly threatened. Distribution: Washington Co., Utah, Mohave Co., Arizona, and Clark Co., Nevada (Heiser 1961). Heterotheca jonesii (Blake) Welsh & At- wood, comb. no\'., based on Chrysopsis jonesii Blake in Tidestrom, Contr. U.S. Natl. Herb. 25: 536. 1925, a substitute name for C. caespitosa M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 694. 1895, not Nutt. Type: Springdale, Washington Co., Utah, 4,000 feet, 16 May 1894, M. E. Jones 5249u (pom). Status: Endemic, rare, local, and en- dangered. Distribution: Washington and (jar- field {Harrison 12345 [bry]) counties, Utah. Heterotheca grandi flora Nutt., Trans. Amer. Philos. Soc. II, 7: 315. 1840. Status: Rare, local and restricted. Distribution: Washington Co., Utah {Welsh et al. 9530 [bry]); also in Cali- fornia and Arizona. Heterotheca subaxillaris Britton & Rusby, Trans. New York Acad. Sci. 7: 10. 1887. Status: Rare, possibly endangered. Distribution: Grand Co., Utah {Welsh & Moore 2744 [bry]); wide- spread elsewhere in the United States. Hulsea heterochroma A. Gray, Proc. Amer. Acad. Arts 7: 359. 1868. Status: Local, rare and possibly threatened. Distribution: Washington Co., Utah [Higgins & Atwood 1410 [bry]; Hig- gins 1972b); California and southern Nevada. Hymenopappus filifolius Hook. var. al- pestris (Maguire) Shinners, Rhodora 61: 155. 1959, based on H. nudipes var. alpestris Maguire, Amer. Midi. Nat- uralist 37: 144. 1947 (includes var. nudipes (Maguire) Turner). Type: Cedar Breaks rim. Iron Co., Utah, 23 Jun 1940, Maguire 19023 (ny). Status: Restricted but locally com- mon, not threatened nor endangered. Distribution: Duchesne, Garfield, Iron, Kane, Piute, Sevier, Summit, and Utah counties, Utah, and Lincoln Co., Wyoming (Turner 1956). Hymenopappus filifolius Hook. var. to- mentosus (Rydb.) Turner, Rhodora 58: 237. 1956, based on H. tomentosus Rydb., Bull. Torrey Bot. Club 27: 633. 1900. Type: St. George, Washington Co., Utah, 1877, Palmer 270 (ny). Status: Endemic, rare and restricted, threatened (Ripley, T). Distribution: Kane and Washington counties, Utah (Turner 1956). Hymcnoxys depressa (Torr. & Gray ex Gray) Welsh & Reveal, comb, nov., based on Actinclla depressa Torr. & Gray ex Gray, Mem. Amer. Acad. Arts II, 4: 100. 1849. Status: Rare and . local, possibly threatened. Distribution: Emery (bry) and Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 337 Garfield (ut) counties, Utah, and from Colorado. Lepidosparluni latisquamum S. Wats., Proc. Amer. Acad. Arts 25: 133. 1890. Status: Restricted and rare, possibly threatened. Distribution: Millard Co., Utah (bry), and adjacent Nevada west to eastern California. Lygodesmia grandiflora (Nutt.) Torr. & Gray var. striata Maguire, Amer. Midi. Naturalist 37: 145. 1947. Type: Frequent on adobe clay, 1 mile south of Price, Carbon Co., Utah, 5 Jun 1940, Maguire 18417 (ny). Status: Endemic, rare, restricted and endangered. Distribution: Carbon Co., Utah; known only from the type locality. Machaer anther a glabriuscula (Nutt.) Cronq. & Keck var. confertifolia Cronq., Leafl. W. Bot. 10: 11. 1963. Type: Barren clay slopes in pinyon- juniper zone, 11 miles northeast of Henrieville, Garfield Co., Utah, 7,000 feet, 31 May 1961, Cronquist 9164 (ny). Status: Endemic, rare and threat- ened. Distribution: Garfield and Kane counties, Utah (bry, utc) . Machaeranthera grindelioides (Nutt.) Shinners var. depressa (Maguire) Cronq. & Keck, Brittonia 9: 237. 1957, based on Haplopappus nuttallii var. depressa Maguire, Amer. Midi. Natural- ist 37: 144. 1947. Type: Warm Point, 5 miles southwest of Desert Range Experiment Station headquarters, Millard Co., Utah, 10 Jun 1941, Maguire 20859 (ny). Status: Rare and restricted, neither threatened nor endangered (Ripley, T) . Distribution: Beaver and Millard counties, Utah (bry), and Eureka, Lin- coln and White Pine counties, Nevada (Maguire 1947). Machaeranthera kingii (D. C. Eaton) Cronq. & Keck, Brittonia 9: 238. 1957, based on Aster kingii D. C. Eaton in King, Rep. Geol. Explor. 40th Parallel 5: 141. 1871. Type: Wasatch Mts., above Cotton- wood Canyon, Salt Lake Co., Utah, 1869, 9,000 feet, Watson 507 (gh). Status: Endemic, rare and restricted, possibh' threatened. Distribution: Cache, Salt Lake and Utah counties, Utah (bry, utc). Malacothrix clcvelandii A. Gray. Bot. Calif. 1: 433. 1876. Status: Local and infrequent. Distribution: Washington Co., Utah [Ativood 4966 [bry, wts]); also in Nevada and California. Partheniuni ligulatum (M. E. Jones) Barneby, Leafl. W. Bot. 5: 20. 1947, based on P. alpinum var. ligulatum M. E. Jones, Contr. W. Bot. 13: 16. 1910. Type: On nearly bare clayey and gravelly knolls on ridges, Theodore [now Duchesne], Duchesne Co., Utah, 6,000 feet, M. E. Jones s.n. (pom). Status: Endemic, rare and restricted, threatened (Ripley, T). Distribution: Duchesne and Emery counties, Utah (bry, ny). Senecio dimorphophyllus Greene var. intermedins Barkley, Trans. Kansas Acad. Sci. 65: 363." 1962. Type: Edge of swampy places. Geys- er Pass, LaSal Mts. Utah, 10,500 feet. Pay son & Pay son 4097 (mo). Status: Endemic, rare, restricted and threatened (Ripley, T) . Distribution: San Juan Co., Utah; known only from the type area. Sphaeromeria capitata Nutt., Trans. Amer. Philos. Soc. II, 7: 402. 1841. Status: Local and rare, threatened. Distribution: Grand Co., Utah (utc); Wyoming and southern Mon- tana. Townsendia aprica Welsh & Reveal, Brit- tonia 20: 375. 1968. Type: Ca 6 miles south of Fremont Junction along Utah Highway 72, on low, rolling exposed gray clay slopes, among scattered igneous boulders, Se- vier Co., Utah, ca 6,500 feet, 1 May 1966, Reveal & Welsh 721 (bry). Status: Endemic, rare and restricted, endangered (Ripley, E) . Distribution: Sevier Co., Utah (bry; Welsh & Reveal 1968). Townsendia mensana M. E. Jones, Contr. W. Bot. 13: 15. 1910. 338 GREAT BASIN NATURALIST Vol. 35, No. 4 Type: Benches of the Uinta Mts., near Theodore [now Duchesne], Du- chesne Co., Utah, 14 May 1908, M. E. Jones s.n. (pom). Status: Endemic, locally restricted, possibl}' threatened. Distribution: Duchesne and Uintah counties, Utah (Reveal 1970b). Townsendia minima Eastw., Leafl. W. Bot. 1: 206. 1936. Type: Bryce Canyon, Garfield Co., Utah. 19 Jun 1933, Eastwood & Howell 727 (cAs). Status: Endemic, edaphically restric- ted, possibly threatened. Distribution: Garfield and Kane counties. Utah (Beaman 1957; Reveal 1970b). Viguiera soliceps Barneby. Leafl. W. Bot. 10: 316. 1966. Type: Locally plentiful on gumbo- clay knolls and bluffs, lower Cotton- wood Canyon near its confluence with Paria River, about 41 miles southeast of Cannon ville, Kane Co., Utah, 4,500 feet, 12 Jun 1966, Barneby 14435 (ny). Status: Endemic, local on Tropic Shale formation, threatened (Ripley. E). Distribution: Cottonwood Canyon east to Last Chance Canyon, Kane Co., Utah (bry, utc). Xanthocephalum sarothrae (Pursh) Shin- ners var. pomariense (Welsh) Welsh, comb, nov., based on Gutierrezia saro- thrae (Pursh) Britton & Rusby var. pomariensis Welsh. Great Basin Nat. 30: 19. 1970. Type: Sand}' flat, base of sandstone cliffs, Frontier formation, mouth of Orchard Creek Draw, Dinosaur N.M.. Uintah Co., Utah, 2 Oct 1969, Welsh et al. 9471 (bry). Status: Endemic, rare and threat- ened. Distribution: Uintah Co., Utah (Welsh 1970b). Berberidaceae Berberis fendleri A. Gray, Mem. Amer. Acad. Arts II, 4: 5. 1849. Status: Rare, floristically restricted to seeps and hanging gardens. Distribution: San .luan (Maguirc 5904, Holmgren 13850 [utc]; Moore 204a; Welsh & Moore 3839 [bry]; Ma- guire 1937); southern Colorado and New Mexico. Betul.'^ceae Betula X utahensis Britton, Bull. Torrey Bot. Club 31: 165. 1904. Type: City Creek Canyon, Salt Lake Co., Utah, 18 Apr 1900, Stokes s.n. (ny). Status: Rare; a putative hybrid be- tween B. occidentalis Hook and B. papy- rifera Marsh. Distribution: In Utah known only from the t}'pe locality; the hybrid is widespread north of Utah (Dugle 1966). Ostrya knowltonii Coville, Gard. & Forest 7:' 114. 1894. St.atus: Rare, disjunct in hanging gardens, along seeps and on slickrock in sandstone canyons. Distribution: Grand [Cottam 2145 and 556 [bry, utc]), Kane {Welsh & Toft 11871 [bry]), San Juan (Welsh et al. 2939. 2961 and 3721; Moore 336; Welsh 11893; Welsh & Moore 11783; Welsh & Atwood 11693 and 11711; Atwood 4103 [bry]); northern Arizona, southwestern New Mexico and western Texas (Little 1953; Correll & Johnston 1970). Boraginaceae Cryptantha barnebyi I. M. Johnston, J. Arnold Arbor. 29: 240. 1948. Type: Ca 30 miles south of Ouray on white shale knolls, Uintah Co., Utah, 17 Jun 1947, Ripley & Barneby 8748 (gh). Status: Endemic, rare, threatened (Ripley, T). Distribution: Uintah Co., Utah (Higgins 1971). Cryptantha brevi flora (Osterh.) Payson, Ann. Missouri Bot. Gard. 14: 318. 1927, based on Oreocarya breviflora Osterh. ex Payson, Univ. Wyoming Publ. Sci., Bot. 1: 169. 1926. Type: 6.5 miles north of Jensen, Uintah Co., Utah, 19 Jun 1925, Oster- hout 6414 (rm). St.-vtus: Endemic. . locally common ill the Uinta Basin, neither threatened nor endangered (Ripley, E). Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 339 Distribution: Daggett, Duchesne and Uintah counties, Utah (Higgins 1971). Cryptantha capitata (Eastw.) I. M. Johnston, J. Arnold Arbor. 21: 66. 1941, based on Oreocarya capitata Eastw., Leafl. W. Bot. 1: 9. 1937. Status: Rare and obscure. Distribution: Garfield, Kane, Washington, and Wayne counties, Utah (Higgins 1971), and in Coconino Co., Arizona. Cryptantha compacta Higgins, Great Ba- sin Nat. 28: 196. 1968. Type: Ca 8 miles west of Desert Range Experiment Station headquar- ters, along Utah Highway 21, Millard Co., Utah, 18 Jun 1968/ Higgins 1613 (bry). Status: Endemic, rare and local, threatened (Ripley, T). Distribution: Millard Co., Utah (Higgins 1971). Cryptantha elata (Eastw.) Payson, Ann. Missouri Bot. Card. 14: 285. 1927, based on Oreocarya elata Eastw., Bull. Torrey Bot. Club 30: 241. 1903. Status: Rare and possibly threatened (Ripley, T). Distribution: Grand Co., Utah, and adjacent Mesa Co., Colorado (Higgins 1971). Cryptantha grahamii I. M. Johnston, J. Arnold Arbor. 20: 391. 1939. Type: On bench west of Green River, north of the mouth of Sand Wash, Uintah Co., Utah, 28 May 1933, Graham 7924 (gh). Status: Endemic, locally abundant, on white shale outcrops along Willow Creek, possibly threatened (Ripley, E). Distribution: Uintah Co., Utah (Higgins 1971). Cryptantha johnstonii Higgins, Great Ba- in Nat. 28: 195. 1968. Type: 15 miles west of U.S. High- way 50-6 along the road from Wood- side to Castle Dale, Emery Co., Utah, 25 May 1968, Higgins 1510 (bry). Status: Endemic, locally common, threatened (Ripley, E). Distribution: Emery Co., Utah (Higgins 1971). Cryptantha jonesiana (Payson) Payson, Ann. Missouri Bot. Gard. 14: 323. 1927, based on Oreocarya jonesiana Payson, Univ. Wyoming Publ. Sci., Bot. 1: 168. 1926. Type: San Rafael Swell, Emery Co., Utah, 15 May 1914, M. E. Jones s.n. (pom). Status: Endemic, rare except in clay barrens in the San Rafael Swell, threat- ened (Ripley, E). Distribution: San Rafael Swell, Emery Co., Utah (Higgins 1971). Cryptantha longiflora (A. Nels. ) Payson, Ann. Missouri Bot. (7ard. 14: 326. 1927, based on Oreocarya loniflora A. Nels., Erythea 7: 67. 1899. Status: Rare and restricted in Grand Valley. Distribution: Grand Co., Utah, and west central Colorado (Higgins 1971). Cryptantha rnensana (M. E. Jones) Pay- son, Ann. Missouri Bot. Gard. 14: 333. 1927, based on Krynitzkia rnensana M. E. Jones, Contr. W. Bot. 13: 14. 1910. Type: Emery, Emery Co., Utah, 16 May 1894, M. E. Jones 5445p (pom). Status: Rare and restricted to clay soils; threatened. Distribution: Carbon, Emery, and Grand counties, Utah (Higgins 1971). Cryptantha ochroleuca Higgins, Great Ba- sin Nat. 28: 197. 1968. Type: On an outcrop 100 meters south of Red Canyon Campground, along Utah Highwav 12, 21 Jul 1968, Higgins 1788 (bry)^ Status: Endemic, rare and endan- gered (Ripley, E) . Distribution: Garfield Co., Utah (Higgins 1971) . Cryptantha osterhoutii (Payson) Payson, Ann. Missouri Bot. Gard. 14: 329. 1927, based on Oreocarya osterhoutii Pavson, Univ. Wyoming Publ. Sci., Bot. 1:" 167. 1926. Status: Rare or obscure. Distribution: Disjunct in Duchesne, San Juan, and Wayne counties, Utah, and in Mesa Co., Colorado (Higgins 1971). Cryptantha paradoxa (A. Nels.) Payson, Ann. Missouri Bot. Gard. 14: 330. 340 GREAT BASIN NATURALIST Vol. 35, No. 4 1927, based on Oreocarya paradoxa A. Nels., Bot. Gaz. 56: 69. 1913. Status: Rare, San Rafael Swell, threatened. Distribution: Emery Co., Utah, western Colorado and northwestern New Mexico (Higgins 1971). Cryptantha rollinsii I. M. Johnston, J. Arnold Arbor. 20: 391. 1939. Type: Shale hillside on Thome's Ranch, near Willow Creek, ca 22 miles south of Ourav, Uintah Co., Utah, 16 Jun 1937, Rollins 1715 (gh). Status: Endemic, locally common, neither threatened nor endangered. Distribution: Uinta Basin, in both Duchesne and Uintah counties, and on the San Rafael Swell, Emeny' Co., Utah (Higgins 1971). Cryptantha semiglabra Barneby, Leafl. W. Bot. 3: 197. 1943. Status: Rare and restricted, threat- ened (Ripley, T) . Distribution: Vicinity of Fredonia, Coconino, and Mohave counties, Ari- zona, and in Washington Co., Utah (Higgins 1971). Cryptantha striata (Osterh.) Pay son, Ann. Missouri Bot. Card. 14: 264. 1927, based on Oreocarya stricta Osterh., Bull. Tor- rey Bot. Club 50: 217. 1923. Status: Rare, but neither threatened nor endangered (Ripley, T) . Distribution: Daggett, Summit, and Uintah counties, Utah; also in Moffat Co., Colorado, and Carbon Co., Wyo- ming (Higgins 1971). Cryptantha tenuis (Eastw.) Payson, Ann. Missouri Bot. Gard. 14: 327. 1927, based on Oreocarya tenuis Eastw., Bull. Torrey Bot. Club 30: 244. 1903. Type: Court House Wash, near Moab, Grand Co., Utah, 25 May 1892, Eastwood s.n. (cas). St.\tus: Endemic, common and wide- spread. Distribution: Emery, Grand, San Juan, and eastern Wayne counties, Utah (Higgins 1971). Cryptantha wether illii (Eastw.) Payson, Ann. Missouri Bot. Gard. 14: 324. 1927, based on Oreocarya wetJierillii Eastw., Bull. Torrey Bot. Club 30: 242. 1930. Type: Court House Wash, near Moab, Grand Co., Utah, 25 May 1892, Eastwood s.n. (cas). Status: Endemic, common and wide- spread. Distribution: Carbon, Emery, Gar- field, Grand, and Wayne counties, Utah (Higgins 1971). Hackelia patens (Nutt.) I. M. Johnston var. harrisonii J. L. Gentry, Southw. Naturahst 19: 140. 1974. Type: Pine Valley Mts., ca 1.5 miles up Forsyth Trail from Pine Valley, 30 May 1968, 7,300 feet, Gentry 2002 (ny). Status: Endemic, restricted but lo- cally common. Distribution: Utah, Wasatch, and Washington counties, Utah (Gentry 1974). Mertensia arizonica Greene, Pittonia 3: 197. 1897. Type: "Arizona," without definite locality, but more likely from southern Utah, 1869, Palmer s.n. (us). Status: Endemic (?), locally com- mon; the var. arizonica is not known from Arizona in modern times. Distribution: Beaver, Garfield, Iron, Piute, and Washington counties, Utah (Higgins 1972a). Mertensia viridis A. Nels. var. cana (Rydb.) L. O. Williams, Ann. Missouri Bot. Gard. 24: 115. 1937, based on M. cana Rydb., Bull. Torre^' Bot. Club 36: 698. 1909. Status: Rare and restricted, possibly threatened. Distribution: Bald Mtn., Summit Co., Utah (Maguire 14699; Weber 5894 [uTc] ) and north central Colorado (Williams 1937; Higgins 1972a). Mertensia viridis A. Nels. ^-ar. dilatata (A. Nels.) L. O. Williams, Ann Mis- souri Bot. Gard. 24: 113. 1937, based on M. coriacea var. dilatata A. Nels., Bull. Torrey Bot. Club 29: 403. 1902. Status: Rare, known from a single collection from the Uinta Mts. Distribution: Daggett Co., Utah {Williams 599 |rm]); southeastern Wyoming and adjacent Colorado (Wil- liams 1037; Higgins 1972a). Dec. 197'. WELSH. ET AL.: ENDANGERED UTAH PLANTS 341 Brassicaceae Arabis deniissa Greene ^ar. languida Rol- lins, Rhodora 43: 388. 1941. Status: Rare and possibly threat- ened (Ripley, T). Distribution: Daggett Co., Utah (Rollins 1941); Albany and Sweet- water counties, Wyoming. Arabis dcmissa Greene \c\r. russcola Rol- lins, Rhodora 43: 387. 1941. Type: Ca 18 miles north of Vernal, Uintah Co., Utah, Jun 1937, Rollins 1757 (gh). Status: Rare and threatened (Rip- ley, T). Distribution: Daggett and Uintah counties, Utah, and Albany Co., Wyo- ming (Rollins 1941). Caulanthus divaricatus Rollins. Contr. Gray Herb. 201: 8. 1971. Type: Ca 75 miles west of Blanding and 10 miles east of Hite, San Juan Co., Utah, 16 May 1961, Cronquist 9033 (NY). Status: Endemic, rare though wide- spread, neither threatened nor endan- gered. Distribution: Carbon, Emery, Gar- field, Grand, San Juan, and Wayne counties, Utah (bry. utc). Draba maguirei C. L. Hitchc, Univ. Wash. Publ. Biol. 11: 71. 1941. Type: Rocky soil on east slope of Mt. Naomi, Bear River Range, Cache Co., Utah, 9,600 feet, Maguire et al. 14161 (wtu). Status: Endemic, restricted and possibly threatened. Distribution: Cache Co.. Utah (Hitchcock 1941). Draba maguirei C. L. Hitchc. var. burkei C. L. Hitchc, Univ. Wash. Publ. Biol. 11: 72. 1941. Type: Cottonwood Canyon, Wells- ville Mts., Box Elder Co., Utah, 25 May 1932, Burke 2968 (utj. Status: Endemic, restricted and threatened. Distribution: Box Elder (Hitch- cock 1941) and Weber {Clark 2332 [bry, wsco]) counties, Utah. Draba pectinipila Rollins, Rhodora 55: 231. 1953. Status: Endangered or possibly ex- tirpated. Distribution: Flaming Gorge, Dag- gett Co., Utah, and adjacent Sweet- water Co., Wyoming (Rollins 1953). Draba sobolifera Rydb., Bull. Torrev Bot. Club 30: 251. 1903. Type: Marysvale, Piute Co., Utah, above timber line, 1894, M. E. Jones 5936 (us). Status: Endemic, rare and threat- ened (Ripley, T). Distribution: Garfield and Piute counties, Utah (Hitchcock 1941). Draba subalpina Goodman & Hitchc, Ann. Missouri Bot. Card. 19: 77. 1932. Type: Cedar Breaks, Iron Co., Utah, Goodman & Hitchcock 1622 (mo). Status: Endemic, infrequent and possibly threatened (Ripley, T). Distribution: Garfield, Iron and Kane counties, Utah (bry, utc). Draba zionensis C. L. Hitchc, Univ. Wash. Publ. Biol. 11: 49. 1941. Type: Zion Canyon, Washington Co., Utah, 7 May 1923, M. E. Jones s.n. (pom). Status: Endemic, rare and threat- ened (Ripley, T). Distribution: Juab (Cottam 7201 UT) and Washington counties, Utah (Hitchcock 1941). Glaucocarpum suffrutescens (Rollins) Rollins, Madrono 4: 233. 1938, based on Thelypodium suffrutescens Rollins in Graham, Ann. Carnegie Mus. 26: 244. 1937. Type: West of Willow Creek, on Thome's Ranch, eastern slope of Big Pack Mtn., Uintah Co., Utah, 23 May 1935, Graham 8950 (gh). Status: Endemic, rare and endan- gered (Ripley, E). Distribution: Uintah Co., Utah; known only from the tApe locality (Rollins 1938). Lepidium barnebyanum Reveal, Great Ba- sin Nat. 27: 178. 1967, based on L. mon- tanum Nutt. ex Torr. & Gray ssp. de- missum C. L. Hitchc, Madrofio 10: 157. 1950. Type: Indian Creek Canyon, on white shale ridge tops, ca 4 miles south- west of Duchesne, Duchesne Co., Utah, 342 GREAT BASIN NATURALIST Vol. 35, No. 4 15 Jun 1947, Ripley & Barneby 8699 (WTU). Status: Endemic, rare and endan- gered (Ripley, E). Distribution: Duchesne Co., Utah; knowTi onl}' from the type locality (Reveal 1967). Lesquerella garrettii Payson, Ann. Mis- souri Bot. Gard. 8: 213. 1921. Type: Big Cottonwood Canyon, Salt Lake Co., Utah, 28 Jun 1908, Garrett 1544 (mo). Status: Endemic, rare and possibly threatened (Ripley, T). Distribution: Salt Lake, Utah and Wasatch counties, Utah (Rollins & Shaw 1973). Lesquerella hemiphysaria Maguire, Amer. Midi. Naturalist 27: 456. 1942. Type: Frequent, limestone breaks, south side of Middle Fork Park, Wa- satch Plateau, 10 Aug 1940, 10,800 feet, Maguire 20053 (utc). Status: Endemic, locally common but restricted. Distribution: Carbon, Emery, San- pete, and Utah counties, Utah (Rollins & Shaw 1973). Lesquerella rubicundula Rollins, Contr. Dudley Herb. 3: 178. 1941. Type: Red Canyon, Garfield Co., Utah, 2,300 meters, 6 Jul 1912, Eggle- ston 8198 (na, the type now trans- ferred to us). Status: Endemic, rare and threat- ened (Ripley, T) . Distribution: Garfield, Kane, and Piute counties, Utah (Reveal 1970a; Rollins & Shaw 1973). Lesquerella tumulosa (Barneby) Reveal, Great Basin Nat. 30: 97. 1970, based on L. hitchcockii Munz ssp. tumulosa Barneby, Leafl. W. Bot. 10: 313. 1966. Type: Bare white shale knolls, 6.5 miles southeast of Cannonville, Kane Co., Utah, 12 Jun 1966, Barneby 14424 (NYj. Status: Endemic, rare and endan- gered. Distribution: Kane Co., Utah; known only from the type locality (Reveal 1970a). Lesquerella utahensis Rydb., Bull. Torre v Bot. Club 30: 252. 1903. Type: American Fork Canyon, Utah Co., Utah, 31 Jul 1880, M. E. Jones 1354 (ny). Status: Endemic, rare to locally com- mon. Distribution: Davis, Duchesne, Juab, Salt Lake, Summit, Tooele, Uintah, Utah, Wasatch and Weber counties, Utah (RoUins & Shaw 1973). Parry a rydbergii Botsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk SSSR 17: 178. 1955, based on P. platy- carpa Rydb., Bull. Torrey Bot. Club 39: 326. 1912, not Hook. f. & Thomas. Type: Uinta Mts., Aug 1869, Wat- son 54 (ny). Status: Endemic, rare and threat- ened (Ripley, T). Distribution: Crest of the Uinta Mts., Duchesne {Murdoch 54 [bry]) and Daggett-Uintah counties {Waite 252 and 297 [bry]). Physaria grahamii Morton in Graham, Ann. Carnegie Mus. 26: 220. 1937. Type: Chandler Canyon, Uintah Co., Utah, 3 Aug 1935, 6,000 feet, Graham 9976 (us). Status: Endemic, rare and possibly extinct (Ripley, PoEx). Distribution: Uintah Co., Utah; known only from the type collection as recent efforts to recollect this spe- cies have failed (Waite 1973). Thely podium integrifolium (Nutt.) Endl. var. complanatum (Al-Shehbaz) Welsh & Reveal, stat. nov., based on T. in- tegrifolium ssp. complanatum Al-Sheh- baz, Contr. Gray Herb. 204: 105. 1973. Status: Local and infrequent. Distribution: Box Elder Co., Utah (uTC; Al-Shehbaz 1973); found to the west of LTtah. Thelypodium rollinsii Al-Shehbaz. Contr. Gray Herb. 204: 97. 1973. Type: On alkaline soil in the vicinity of the Sevier RiAer. ca 12 miles north of Scipio along U.S. Highway 91, 29 Jul 1969, Al-Shehbaz &■ Al-Shehbaz 6913 (gh). Status: Endemic, locally common, neither threatened nor endangered. Distribution: Carbon, Juab, Mil- lard, Piute, Sanpete, and Sevier coun- ties, Utah (Al-Shehbaz 1973). Dec. 1975 WELSH. ET AL.: ENDANGERED UTAH PLANTS 343 Thely podium sagittatum (Nutt.) Endl. ex Walp. var. ovalifoUum (Rydb.) Welsh & Reveal, stat. & comb, nov., based on T. ovaJifoUum Rydb., Bull. Torrey Hot. Club 30: 253. 1903. Type: Pangnitch Lake, Garfield Co., Utah, 7 Sep 1894, M. E. Jones 601 5e (us). Status: Rare and restricted, possibly threatened. Distribution: Garfield and Iron counties, Utah, and White Pine Co.. Nevada (Al-Shehbaz 1973). Cactaceae Echinocereus engelmannii (Parry) Le- maire var. purpurcus L. Benson, Cact. Succ. J. (Los Angeles) 41: 127. 1969. Type: Mojavean desert north of St. George, Washington Co., Utah, 5 May 1949, 2,900 feet, Benson 15637 (pom). Status: Endemic, rare and endan- gered (Ripley, E). Distribution: Washington Co., Utah; known only from the vicinity of the type locality (Benson 1969b). Ferocactus acanthodes (Lemaire) Britton & Rose, Publ. Carnegie Inst. Wash. 248 (3): 129. 1922, based on Echinocactus acanthodes Lemaire, Cact. Gen. Nov. Sp. 106. 1839. Status: Restricted and endangered. Distribution: Washington Co., Utah (bry); southern Nevada, Arizona and California (Benson 1969b). Mammillaria tetrancistra Engelm., Amer. J. Sci. Arts II, 14: 337. 1852. Status: Restricted and rare, possibly threatened. Distribution: Washington Co., Utah; southern Nevada, California, and Arizona (Benson 1969b). Opuntia pulchella Engelm., Trans. Acad. Sci. St. Louis 2: 201. 1863. Status: Rare, restricted and possibly threatened. Distribution: Millard Co., Utah (bry); Nevada and Arizona (Benson 1957). This species includes those en- tities included in Micropuntia by Das- ton (1946). Opuntia whipplei Engelm. & Bigelow var. multigeniculata L. Benson, Cact. Ariz. ed. 3, 38. 1969. Status: Restricted and rare. Distribution: Washington Co., Utah, and adjacent Nevada and Arizona (Benson 1969b). Pediocactus silcri (Engelm.) L. Benson, Cact. Succ. .1. (Los Angeles) 33: 53. 1961, based on Echinocactus sileri En- gelm. ex Coulter, Contr. U.S. Natl. Herb. 3: 376. 1896. Type: Cottonwood Springs and Pipe Springs, supposedly from southern Utah but more likely from Arizona, 1883, Siler s.n. {mo)/ Status: Arizona Strip endemic, rare, local and endangered (Ripley, E). Distribution: Washington Co., Utah {Welsh 12712 [bry]), and northern Arizona (bry; Benson 1969b). Sclerocactus glaucus (K. Schum.) L. Ben- son, Cact. Succ. J. (Los Angeles) 38: 53. 1966, based on Echinocactus glaucus K. Schum., Gessammtb. Kakt. 438. 1898. Status: Rare, possibly extirpated from Utah (Ripley, E). Distribution: Duchesne and Uintah counties, Utah, and Delta Co., Colorado (Benson 1966). Sclerocactus pubispinus (Engelm.) L.Ben- son, Cact. Succ. J. (Los Angeles) 38: 103. 1966, based on Echinocactus pubi- spinus Engelm., Trans. Acad. Sci. St. Louis 2: 199. 1863. Status: Rare and threatened (Rip- ley,T). Distribution: Box Elder, Beaver, and Sevier counties, Utah, and Elko Co., Nevada (Benson 1966). Sclerocactus pubispinus (Engelm.) L. Benson var. sileri L. Benson, Cact. Ariz. ed. 3, 23. 1969. Type: Southern Utah, 1888, Siler s.n. (PH). St.\tus: Rare and now extirpated from Utah. Distribution: Kane Co., Utah, and Coconino Co., Arizona (Benson 1969b). Sclerocactus whipplei (Engelm. & Bige- low) Britton & Rose var. spinosior (Engelm.) Boissev. in Boissev. & Da- vidson, Colorado Cacti 51. 1940, based on Echinocactus whipplei var. spinosior Engelm., Trans. Acad. Sci. St. Louis 2: 199. 1863. 344 GREAT BASIN NATURALIST Vol. 35, No. 4 Type: South of the Great Sah Lake Desert, 19 Jul 1859, H. Engelmann s.n. (mo). Status: Rare to locally common, neither threatened nor endangered. lau fliul no indication that such a made. According to Benson (1966), this name is a synon.^^n of .V. inibispiiuis (Engohii.i I.. Benson combination h var. pubspinus. Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 345 rather common in Arizona, New Mexi- co and Mexico (Hanson 1692b). Atriplex welshii C. A. Hanson, Stud. Syst. Bot. 1: 1. 1962. Type: 4 miles south of Cisco along Utah Highway 128, Grand Co., Utah, 5 Jul 1961, Hanson 522 (bry). Status: Endemic, restricted and pos- sibly threatened. Distribution: Carbon and Grand counties, Utah (bry) . Cucurbitaceae Cucurbita foetidissima H.B.K., Nov. Gen. &Sp. 2: 123. 1817. Status: Restricted, but locally com- mon and often weedy, neither threat- ened nor endangered. Distribution: San Juan {Welsh et al. 2992 [bry]), and Washington (bry) counties, Utah, and widespread else- where. CUSCUTACEAE Cuscuta warneri Yuncker, Brittonia 12: 38. 1960. Type: Vicinity of Powell, 15 miles west of Fillmore, Millard Co., Utah, 10 Sep 1957, on Phyla cuneifolia, War- ner s.n. (uTc). Status: Endemic, possibly extinct (Ripley, PoEx). Distribution: Millard Co., Utah; kno\vn only from the type collection (Yuncker 1960, 1965). Cyperaceae Car ex curatorum Stacey, Leafl. W. Bot. 2: 213. 1937. Status: Rare and restricted to spec- ialized habitats; threatened. Distribution: San Juan Co., Utah {Welsh 12425 [bry]), and in Grand Canyon, Arizona (Stacey 1937). Cladium californicum (S. Wats.) O'Neill in Tidestr. & Kittell, Fl. Arizona & N. Mex. 773. 1941, based on C. mariscus var. californicum S. Wats., Bot. Calif. 2: 224. 1880. Status: Rare, restricted and endan- gered. Distribution: Kane {Welsh & At- wood 11709; Welsh & Moore 11780; Welsh & Toft 11870 [bry]) and San Juan {Welsh & Toft 11874 [bry]), and from northern Arizona, southern Ne- vada, and southern California to Cen- tral America (Kearney & Peebles 1951). Ela EAGNACEAE Elaeagnus commutata Bemh., Allg. Thij- ring. Gartenzeitung 2: 137. 1843. Status: Restricted, local and possibly extirpated from Utah. Distribution: Daggett {Julander J56 [bry]) and Summit {Welsh & Moore 6724 [bry] ) counties, Utah; widespread northward to Alaska (Welsh 1974b). Ericaceae Arctostaphylos pringlei Parry, Bull. Calif. Acad. Sci. 2: 494. 1887. Status: Local, rare, possibly threat- ened. Distribution: Washington Co., Utah {Higgins 665 and 3414 [bry]; Higgins 1972b). EUPHORBIACEAE Euphorbia nephradenia Barneby, Leafl. W. Bot. 10: 314. 1966. Type: Lower Cottonwood Canyon, about 41 miles southeast of Cannonville, Kane Co., Utah, 12 Jun 1966, 4,500 feet, Barneby 14434 (ny). Status: Endemic, restricted edaph- ically, threatened (Ripley, T). Distribution: Emery, Kane, and Wayne counties, Utah (bry). Fabaceae Astragalus adanus A. Nels., Bot. Gaz. 53: 222. 1912. Status: Rare and possibly extirpated from Utah. Distribution: Juab Co., Utah; wide- spread in Idaho (Barneby 1964). Astragalus adsurgens Pallas ssp. robus- tior (Hook.) Welsh, Iowa State Coll. J. Sci. 37: 357. 1963, based on A. ad- surgens var. robustior Hook., Fl. Bo- reali-Amer. 1: 149. 1831. Status: Restricted, rare and possibly threatened. Distribution: Daggett Co., Utah {Hanson 283a [bry]); widespread east of the continental divide north- ward to Alaska (Welsh 1974b). 346 GREAT BASIN NATURALIST Vol. 35, No. 4 Astragalus alpinus L., Sp. PI. 760. 1753. Status: Rare and local, possibly threatened. Distribution: Grand {Holmgren et al. 232 J [bry, ny, utc] ) and Salt Lake (ut, without collector) counties; wide- spread in North America and Eurasia (Welsh 1974b). Astragalus ampullarius S. Wats., Amer. Naturalist 7: 300. 1873. Type: Kanab, Kane Co., Utah, 1872, Thompson s.n. (gh). Status: Arizona strip endemic, edaphically restricted and though locally common, probably threatened (Ripley, T). Distribution: Kane and Washington counties, Utah, and northernmost Co- conino and Mohave counties, Arizona (bry,- Rarneby 1964). Astragalus asclepiadoides M. E. Jones, Zoe2: 238. 1891. Type: Cisco, Grand Co., Utah, 21 Jun 1889, M. E. Jones s.n. (pom) Status: Edaphically restricted al- though locall}' common, neither threat- ened nor endangered. Distribution: Carbon, Duchesne, Emery, Garfield, Grand, Sanpete, Se- vier (ut), Uintah, and Wayne coun- ties, Utah (bry, utc), and western Colorado (Rarneby 1964). Astragalus aus trails Lam., Fl. Frang. 2: 637. 1778. Status: Possibly extirpated from Utah. Distribution: Piute (?) Co., Utah (Rarneby 1964). Astragalus barneby Welsh & Atwood, nom. nov., based on A. desperatus M. E. Jones var. conspectus Rarneby, Leafl. W. Rot. 5: 87. 1948. Status: Rare, restricted and threat- ened (Ripley, T). Distribution: Garfield Co., Utah (bry), and in Navajo and Coconino counties, Arizona (Rarneby 1964). This remarkable milkvetch is adequate- ly distinct on account of size of flower and parts, and because of stature to segregate it from its near and mirror-imaged cog- ener A. desperatus. The name change honors the author of the monumental "Atlas of North American Astragalus," Rupert C. Rarneby of the New York Ro- tanical Garden. Astragalus bodinii Sheldon, Minnesota Rot. Stud. 1: 122. 1894. Status: Rare and endangered. Distribution: Wayne Co., Utah (bry); widespread in Alaska, Canada, Colorado, Wyoming, and one record from Nebraska (Rarneby 1964). Astragalus brandegei Porter in Porter & Coulter, Syn. Fl. Colo. 24. 1872. Status: Rare and obscure, probably threatened. Distribution: Carbon, Emery, Gar- field, Piute, and Wayne counties, Utah (bry) ; also in Colorado, New Mexico, and Arizona (Rarneby 1964). Astragalus bryantii Rarneby, Proc. Calif. Acad. Sci. IV, 25: 156. 1944. Status: Probably extirpated from Utah. Distribution: Kane Co., Utah, in Glen Canyon, the collection area inun- dated by Lake Powell; also in northern Arizona (Rarneby 1964). Astragalus callithrix Rarneby, Leafl. W. Rot. 3: 102. 1942. Status: Rare and restricted, possibly threatened (Ripley, T) . Distribution: Millard Co., Utah (bry); otherwise known only from two locations in Nye Co., Nevada. Astragalus castaneiformis S. Wats. var. consobrinus Rarneby, Amer. Midi. Naturalist 41: 496. 1949. Type: Southeast of Ricknell, Wayne Co., Utah, 10 Jun 1947, 7,600 feet, Ripley & Barneby 8605 (cas). Status: Endemic, rare and obscure, possibly threatened. Distribution: Garfield, Piute, Se- vier, and Wayne counties, Utah (Rar- neby 1964). Astragalus chloodes Rarneby, Leafl. W. Rot. 5: 6. 1947. Type: Ca 6 miles southeast of Jen- sen, Uintah Co., Utah, 7 Jun 1946, Ripley & Barneby 7797 (g.\s). Status: Endemic, rare and threat- ened (Ripley, T). Distribution: Uintah Co., Utah (BRY. utc). Astragalus coltonii M. E. Jones, Zoe 2: 237. 1891. Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 347 TYPE: Canyons of the Coal Range east of Castle Gate, 27 .Tun 1889, M. E. Jones s.n. (pom) . Status: Endemic, locally abundant, neither threatened nor endangered. Our plant is var. coltonii. Distribution: Carbon, Emery, Gar- field, Kane, Sevier, and Wayne coun- ties, Utah (bry, ut. utc) . Astragalus convaUarius Greene var. finiti- mus Barneby, Leafl. W. Bot. 7: 192. 1954. Type: Ca 3 miles south of Enterprise, Washington Co., Utah, 11 Jun 1942, Ripley & Barneby 4767 (gas). Status: Rare and local, threatened ^(Ripley, T). Distribution: Washington Co., Utah, and adjacent Nevada (Barneby 1964). Astragalus cottamii Welsh, Rhodora 72: 189. 1970. Type: Ca 4 miles east of Clay Hills divide, San Juan Co., Utah, 1 May 1966, Welsh 5207 (bry). Status: Restricted and local, possibly threatened (Ripley, T). Distribution: San Juan Co., Utah, and adjacent Monument Valley, Ari- zona (bry). Astragalus cronquistii Barneby, Mem. New York Bot. Gard. 13: 258. 1964. Type: In desert along west side of Comb Wash, 9 miles west of Bluff, San Juan Co., Utah, 27 May 1961, Cronquist 9123 (ny). Status: Endemic and very restricted, endangered (Ripley, E). Distribution: San Juan Co., Utah (bry, utc). Astragalus cymboides M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 650. 1895. Type: Huntington, Emery Co., Utah, 16 Jun 1894, at 5,000 feet, M. E. Jones 5464] (pom). Status: Endemic, locally common to abundant, neither threatened nor endangered. Distribution: Carbon, Emery, Grand, and San Juan counties, Utah (bry). Astragalus deserticus Barneby, Mem. New York Bot. Gard. 13: 634. 1964. Type: Common on slopes near In- dianola, 17 Jun 1909, Tidestrom 2249 (GH). Status: Endemic and possibly ex- tinct (Ripley, PrEx). Distribution: Sanpete Co., Utah (Barneby 1964). Astragalus detri talis M. E. Jones, Contr. W. Bot. 13: 9. 1910. Type: Ca 4 miles above Theodore [ Duchesne] on the Colton road, Du- chesne Co., Utah, 11 May 1908, M. E. Jones s.n. (pom). Status: Uinta Basin endemic, local and possibly endangered (Ripley, E) . Distribution: Duchesne and Uintah counties, Utah, and Rio Blanco Co., Colorado (bry). Astragalus diversifolius A. Gray, Proc. Amer. Acad. Arts 6: 230. 1864. Status: Rare, chsjunct and possibly extirpated from Utah. Distribution: Juab and Tooele coun- ties; also in southern Idaho where rare (Barneby 1964). Astragalus duchesnensis M. E. Jones, Contr. W. Bot. 13: 9. 1910. Type: Ca 1 3 miles below Theodore [Duchesne] toward Chepeta Well, Du- chesne Co., Utah, 23 May 1908, M. E. Jones s.n. (pom). Status: Endemic, restricted and threatened (Ripley, T). Distribution: Duchesne and Uintah counties, Utah (bry). Astragalus eastwoodae M. E. Jones, Zoe 4: 368. 1894, based on A. preussii var. sulcatus M. E. Jones, Zoe 4: 37. 1893. Type: Westwater, Grand Co., Utah (incorrectly given as "Colorado"), 6 May 1891^ M. E. Jones s.n. (pom). St.^tus: Rare, disjunct and possibly threatened. Distribution: Emery, Grand, and San Juan counties, Utah; also in west central Colorado (Barneby 1964). Astragalus emoryanus (Rydb.) Cory, Rhodora 38: 406. 1936, based on Hamosa emoryana Rydb., Bull. Torrey Bot. Club 54: 237. 1927. Status: Rare and endangered. Distribution: Kane {Atwood 4629 [bry] ) Co., Utah; Arizona, New Mex- ico, Texas, and Mexico (Barneby 1964). Astragalus ensiformis M. E. Jones, Rev. Astrag. 226. 1923. 348 GREAT BASIN NATURALIST Vol. 35, No. 4 Status: Rare and local; threatened (Ripley, T). Distribution: Washington Co., Utah, and Mohave Co., Arizona (bry). Astragalus eucosmus B. L. Robinson, Rho- dora 10: 33. 1908. Status: Possibly extirpated. Distribution: Summit Co. , Utah; widespread in northern North America (Barneby 1964; Welsh 1974b). Astragalus eurekensis M. E. Jones, Contr. W. Bot. 8: 12. 1898. Type: Eureka, Juab Co., Utah, 1891, M. E. Jones s.n. (pom). Status: Endemic, locall}' common to abundant, neither threatened nor endangered. Distribution: Juab, Iron, Millard, Sanpete, Tooele, and Utah counties, Utah (bry,- Barneby 1964). Astragalus flavus Nutt. ex Torr. & Gray var. argillosus (M. E. Jones) Barneby, Mem. New York Bot. Card. 13: 401. 1964, based on A. argillosus M. E. Jones, Zoe2: 241. 1891. Type: Green River, Emery Co., Utah, 7 May 1890, M. E. Jones s.n. (pom). Status: Endemic, restricted edaph- ically but locally common, neither threatened nor endangered. Distribution: Emer\ , Garfield, Grand, and Wayne counties, Utah (bry). Astragalus gilviflorus Sheldon, Minne- sota Bot. Stud. 1: 21. 1894, based on A. triphyllus Pursh, Fl. Amer. Sept. 2: 740. 1814, not Pallas. Status: Rare or possibly extirpated from Utah. Distribution: Summit Co., Utah (uTc); widespread east of the conti- nental divide (Barneby 1964). Astragalus hallii A. Grav var. fallax (S. Wats.) Barneby, Leaff. W. Bot. 9: 91. 1960, based on A. fallax S. Wats., Proc. Amer. Acad. Arts 20: 362. 1885. Status: Rare and local, possibly threatened. Distribution: Garfield and Kane counties, Utah (bry). Astragalus hamiltonii C. L. Porter, Rho- dora 54: 159. 1952. Type: On the Wasatch formation 5 miles south of Vernal, Uintah Co., Utah, 24 May 1950, Hamilton &■ Beath s.n. (rm). Status: Endemic, rare and local, en- dangered (Ripley, E). Distribution: Uintah Co., Utah (bry). Astragalus harrisonii Barneby, Mem. New York Bot. Card. 13: 271. 1964. Type: Wash below the Natural Bridge, near Fruita, Wayne Co., Utah, 8 Jun 1961, Barneby 15131 (cas). Status: Endemic, rare and endan- gered (Ripley, E). Distribution: Wa3'ne Co., Utah; known only from the type area (bry) . Astragalus iodanthus S. Wats, in King, Rep. Geol. Explor. 40th Parallel 5: 70. 1871. Status: Rare and possibly extirpated from Utah. Distribution: Tooele Co., Utah; widespread to the west of Utah (Bar- neby 1964). Astragalus iselyi Welsh, Great Basin Nat. 34: 305. 1974. Type: Brumley Bridge, ca 1.5 miles north of Pack Creek Ranch, San Juan Co., Utah, 5 May 1971, Welsh 10970 (bry). Status: Endemic, edaphically re- stricted, endangered. Distribution: Grand and San Juan counties, Utah (bry,- Welsh 1974a). Astragalus jejunus S. Wats, in King, Rep. Geol. Explor. 40th Parallel 5: 73. 1871. Status: Rare and restricted, possibly threatened. Distribution: Rich Co., Utah (Bar- neby 1964); southwestern Wyoming and in an isolated location in White Pine Co., Nevada. Astragalus kentrophyta A. Gra} var. coloradoensis M. E. Jones, Contr. W. Bot. 10: 63. 1902. Status: Navajo Basin endemic; rare to locally common but probably threat- ened . Distribution: Garfield, Kane, San Juan, and Wayne counties, Utah (bry), and adjacent northern Arizona (Bar- neby 1964). Astragalus lance arius A. Gra>-, Proc. Amer. Acad. Arts 13: 370. 1878. Status: Rare and local in generalized Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 349 habitats; possibly threatened (Ripley, T). Distribution: Kane (bry) and Washington counties, Utah and in Co- conino and MohaA'e counties, Arizona (bry,- Barneby 1964). Astragalus lentiginosus Dougl. ex Hook, var. chartaceous M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 673. 1895. Type: Ephraim, Sanpete Co., Utah, 13 Jul 1894, 6,000 feet, M. E. Jones 5627m (pom). Status: Rare and local, possibly threatened (Ripley, PoEx, Tax?).^ Distribution: Daggett, Juab, San- pete, Sevier, Summit, and Tooele coun- ties, Utah, and widespread from Colo- rado, Wyoming, Idaho, Oregon, Cali- fornia, and Nevada (Schoener 1975). Astragalus lentiginosus Dougl. ex Hook, var. fremontii (Gray ex Torr.) S. Wats, in King, Rep. Geol. Explor. 40th Paral- lel 5: 66. 1871, based on A. fremontii Gray ex Torr. in Whipple, Explor. & Surv. Railroad Route from Mississippi River to Pacific Ocean 4(5): 80. 1857. Status: Rare and ephemeral, pos- sibly threatened. Distribution: Washington Co., Utah (bry,- Schoener 1974); widespread in southern Nevada and adjacent Cali- fornia (Barneby 1964). Astragalus lentiginosus Dougl. ex Hook, var. scorpionis M. E. Jones, Rev. Astrag. 124. 1923. Status: Rare and local; possibly threatened. Distribution: Juab Co., Utah (Schoener 1975), and Nevada (Barne- by 1964). Astragalus lentiginosus Dougl. ex Hook, var. stramineus (Rydb.) Barneby, Leafl. W. Bot. 4: 122. 1945, based on Cystium stramineum Rydb., N. Amer. Fl. 24: 409. 1929. Type: Southern Utah, 1870, Palmer s.n. (ny). Status: Locally common, restricted, neither threatened nor endangered. Distribution: Washington Co., Utah (bry), Mohave Co., Arizona, and Clark Co., Nevada (Schoener 1975). Astragalus lentiginosus Dougl. ex Hook, var. ursinus (A. Gray) Barneby, Leafl. W. Bot. 4: 133. 1945, based on A. ursinus A. Gray, Proc. Amer. Acad. Art 13: 367. 1878. Type: Bear Valley, in south central Utah, 1877, Palmer s.n. (gh). Status: Endemic, possibly extinct. Distribution: Iron (or possibly Se- vier) Co., Utah; perhaps the specimens are mislabeled." Astragalus limnocharis Barneby, Leafl. W. Bot. 4: 236. 1946. Type: Gravelly beach of Navajo Lake, at Spruce Forest Camp, Iron Co., Utah, 13 Jul 1940, Maguire 19474 (ny). Status: Endemic, rare and restricted; threatened. Distribution: Iron and Kane (bry) counties, Utah (Barneby 1964). Astragalus loanus Barneby, Mem. New York Bot. Gard. 13: 661. 1964, based on A. newberryi var. wardianus Bar- neby, Amer. Midi. Naturalist 37: 481. 1947. Type: Canyon east of Glen wood, Se- vier Co., Utah, 1875, Ward 223 (gh). Status: Endemic, rare and threat- ened (Ripley, E) . Distribution: Garfield, Piute, Sevier and Wayne counties, Utah (bry). Astragalus lutosus M. E. Jones, Contr. W. Bot. 13: 7. 1910. Type: White River, Uintah Co., Utah, 25 May 1958, 5,200 feet, M. E. Jones s.n. (pom). Status: Uinta Basin endemic, rare and edaphically restricted; endangered (Ripley, E). Distribution: Uintah Co., Utah, and Rio Blanco Co. Colorado (Barneby 1964). Astragalus malacoides Barneby, Mem. New York Bot. Gard. 13: 500. 1964. Type: Northeast slope of Kaiparowits Plateau, south of Willow Tank, Kane Co., Utah, 9 May 1939, fJarrison 9069 (us). 'Schooner (1975) has deteirmined that this name includes var. platyphrllidius (Rydb.) M. E. Peck, and therefore de- fines the taxon differently from Ripley (1975). "This variety was reduced to sj-nonjnny luider var. palans (M. E. .Tones) M E. Jones bj- Schoener (1975), but that taxon is currentlj' known only from the valleys of the Colorado and Virgin rivers. The exaction location of "Bear Valley" has not been determined (McVaugh 1956). 350 GREAT BASIN NATURALIST Vol. 35, No. 4 Status: Endemic, restricted though locallv common; threatened (Riplev, E). " ' , ^ Distribution: Kane Co., Utah (bry). Astragalus minthorniae (Rydb.) Jeps. var. gracilior (Barneby) Barneby, Amer. Midi. Naturalist 55: 493. 1956, based on A. ensiformis var. gracilior Barne- by Proc. Calif. Acad. Sci. IV. 25: 158. 1944. Type: Ca 5 miles south of Veyo, Washington Co., Utah, 4,900 feet, Rip- ley & Barneby 4951 (cas). Status: Endemic (?), locally com- mon in disjunct populations, possibly threatened (Ripley, E). Distribution: Washington Co., Utah (bry), and possibly in Mohave Co., Arizona. Astragalus monumentalis Barneb^'. Leafl. W. Bot. 7: 35. 1953. Type: White Canyon, 25 miles south- east of Hite, San Juan Co., Utah, 18 May 1950. Harrison 11595 (us). Status: Endemic, rare and restricted edaphically, possibly threatened. Distribution: Garfield and San Juan counties, Utah (bry) . Astragalus musiniensis M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 671. 1895. Type: San Rafael Swell. Emery (?) Co., Utah, 12 May 1914, \1. E. Jones s.n. (pom). Status: Endemic, rare to common, neither threatened nor endangered. Distribution: Carbon. Emery, Gar- field, Grand, and Wayne counties, Utah (bry). Astragalus nidularius Barnebv. Leafl. W. Bot. 8: 16. 1956. Type: Near the head of White Can- yon, 2 miles below the Kachina Bridge of Natural Bridges N.M.. Barneby 12778 (cas). Status: Endemic, restricted and lo- cal, disjunct in several main locations, neither threatened nor endangered. (Ripley, T) . Distribution: Ciarfield, San Juan, and Wayne counties, Utah (bry). Astragalus oophorus S. Wats. var. loncho- caly.r Barneby. Leafl. W. Bot. 7: 194. 1954. Status: Rare and threatened (Rip- ley,T). Distribution: Iron Co., Utah (bry), and adjacent Nevada (Barneby 1964). Astragalus pardalinus (Rydb.) Barneby, Mem. New York Bot. Gard. 13: 883. 1964, based on Phaca pardalina Rvdb.. N. Amer. Fl. 24: 352. 1929. Type: Cedar Mtn., Emery Co., Utah, 20 May 1915, M. E. Jones s.n. (ny). Status: Endemic, restricted though locally common, neither threatened nor endangered (Ripley, E). Distribution: Emery, Garfield, and Wayne counties, Utah (bry). Astragalus perianus Barneby, Mem. New York Bot. Gard. 13: 973. 1964. Type: Mountains north of Bullion Creek, near Marvsvale, Piute Co., Utah, 23 Jul 1905, Rydberg & Carlton 7104 (ny). Status: Endemic and threatened (Ripley, PoEx). Distribution: Piute (Barneby 1964) and Garfield (bry) counties. Utah." Astragalus pinonis M. E. Jones, Contr. W. Bot. 8: 14. 1898. Type: Frisco, Beaver Co., Utah, 22 Jun 1880, ca 8,000 feet. M. E. Jones s.n. (pom). Status: Rare and possibly extirpated. Distribution: Beaver and Juab counties. Utah, and in east central Ne- vada (Barneby 1964). Astragalus platytropis A. Gray. Proc. Amer. Acad. Arts 6: 526. 1865. Status: Rare and local, both edaph- ically and altitudinally restricted. Distribution: Western Beaver (bry) and Tooele counties. Utah, and much of Nevada and adjacent Cali- fornia (Barneby 1964). Astragalus rafaelensis M. E. Jones, Rev. Astrag. 146. 1923. Type: Near Cedar Mtn., Emery Co., Utah, 19 May 1915, M. E. Jones s.n. (pom). Status: Endemic, restricted edaph- ically, threatened (Ripley, T). Distribution: San Rafael Swell, Emery Co., Utah (bry).. "This species, previously known only from the type coUeclioii. was rediscovered in 1073; one of many species though be extinct but found diunng 1975 as a result of the interest generated by the Smithsonian Report. Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 351 Astragalus sabulosus M. E. Jones, Zoe 2: 239. 1891. Type: Cisco, Grand Co., Utah, 2 May 1890, M. E. Jones s.n. (pom). Status: Endemic, rare and edaph- ically restricted; threatened. Distribution: Grand Co., Utah (bry; Welsh 1974a). Astragalus saurinus Barneby, Leafl. W. Bot. 8: 17. 1956. Type: Dinosaur N.M., 6 miles north of Jensen, Uintah Co., Utah, 26 Jun 1953, Holmgren & Tillett 9527 (ny). Status: Endemic, rare and edaph- ically restricted; threatened (Ripley, E). Distribution: Uintah Co., Utah (bry. utc). Astragalus serpens M. E. Jones. Proc. Calif. Acad. Sci. II, 5: 641. 1895. Type: Loa Pass, Wayne Co., Utah, 17 Jul 1894, M. E. Jones 56391 (pom). Status: Endemic, local and period- ically abundant in disjunct populations, neither threatened nor endangered (Ripley, E). Distribution: Garfield, Piute and Wayne counties, Utah (bry). Astragalus stocksii Welsh, Great Basin Nat. 34: 307. 1974. Type: Henry Mts., Penellen Pass, Garfield Co., Utah, 30 May 1972, Welsh 11740 (bry). Status: Endemic, rare and local; threatened. Distribution: Garfield Co., Utah; known only from the type area (bry). Astragalus striatiflorus M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 643. 1895. Type: Above Springdale, Washing- ton Co., Utah, 25 Sep 1894, 4,000 feet, M. E. Jones 6080k (pom). Status: Rare and edaphically re- stricted; endangered (Ripley, T). Distribution: Kane and Washing- ton counties, Utah, and Coconino Co., Arizona (bry) . A main locality of distribution for this plant is the Coral Pink Dunes region where the plants grow in the interdune valleys. The use of that region for rec- reation possibly spells the doom of this plant in that section. Astragalus tephrodes A. Gray var. brachy- lobus (A. Gray) Barneby, Amer. Midi. Naturalist M -. 466. 1947, based on A. shortianus var. brachylobus A. Gray, Proc. Amer. Acad. Arts 13: 367. 1878. Status: Rare and possibly extirpated from Utah. Distribution: Washington Co., Utah (Barneby 1964). Astragalus tctrapterus A. Gray, Proc. Amer. Acad. Arts 13: 369. 1878. Type: Ca 25 miles north of St. George, Washington Co., Utah, 1877, Palmer 111 (gh). Status: Restricted but locally com- mon to abundant, possibly threatened. Distribution: Beaver, Iron, Kane, and Washington counties, Utah (bry) ; also in northwestern Arizona (bry), Nevada, and Oregon (Barneby 1964). Astragalus wardii A. Gray, Proc. Amer. Acad. Arts 12: 55. 1876. Type: On the edge of the Aquarius Plateau, Sevier Co., Utah, 1875, 8,700 feet. Ward 424 (gh). Status: Endemic, local and disjunct, neither threatened nor endangered. Distribution: Garfield, Kane, Piute, and Sevier counties, Utah (bry). Astragalus wetherillii M. E. Jones, Zoe 4: 34. 1893. Status: Possible extirpated from Utah (Ripley, T). Distribution: Grand Co., Utah, and west central Colorado (Barneby 1964). Astragalus woodruffii M. E. Jones, Rev. Astrag. 77. 1923. Type: On the sandy foot of the San Rafael Swell, Emery (?) Co., Utah, 17-18 May 1914, M. E. Jones s.n. (pom). Status: Endemic, Local and edaph- ically restricted, possibly threatened. Distribution: Emery, Wayne, and possibly Garfield counties, Utah (bry). Dalea epica Welsh, Great Basin Nat. 31: 90. 1971. Type: Ca 10 miles east of Halls Crossing, San Juan Co., Utah, 30 Apr 1966, Welsh 5205 (bry). Status: Endemic, restricted and rare, and possibly threatened. Distribution: San Juan Co., Utah (bry); known only from the type lo- cality. 352 GREAT BASIN NATURALIST Vol. 35, No. 4 Hedysarum boreale Nutt. var. gremiale (Rollins) Northstrom & Welsh, Great Basin Nat. 30: 125. 1970, based on H. gremiale Rollins, Rhodora 42; 230. 1940. Type: West of Vernal, Uintah Co., Utah, 16 Jun 1937, Rollins 1735 (gh). Status: Endemic, local and restric- ted, possibly threatened. Distribution: Duchesne and Uintah counties, Utah (bry). Hoffmanseggia repens (Eastw.) Cocker- ell, Muhlenbergia 4: 68. 1908, based on Caesalpinia repens Eastw., Zoe 4: 116. 1893. Type: Court House Wash, near where it comes into the Grand [Colo- rado] River, near Moab, Grand Co., Utah, 26 May 1892, Eastwood s.n. (CAs). Status: Endemic, edaphically re- stricted but locally common, neither threatened nor endangered, but appar- ently extirpated from the type area (Harrison et al. 1964). Distribution: Emery (bry), Grand, and Wayne (bry) counties, Utah. Lathyrus brachy calyx Rydb. ssp. zionis (C. L. Hitchc.) Welsh, Proc. Utah Acad. Sci. 52: 217. 1966, based on L. zionis C. L. Hitchc, Univ. Wash. Publ. Biol. 15: 36. 1952. Type: Ten miles east of the east en- trance of Zion N.P., Kane (?) Co., Utah, 30 May 1949, Hitchcock 19013 (WTU). Status: Rare to locally abundant in disjunct populations, neither threatened nor endangered. Distribution: Grand, Kane, San Juan, and Washington counties, Utah, and Coconino Co., Arizona (bry) ; re- portedly more widespread in Arizona (McDougall 1973) . Lotus longebracteatus Rydb., Bull. Tor- rey Bot. Club 30: 254. 1903. Type: Southern Utah, possibly Wash- ington Co., Utah, 1877, Palmer 94 (us). Status: Endemic (?), restricted but locally abundant, neither threatened nor endangered. Distribution: Kane and Washing- ton counties, Utah (bry,- Ottley 1944). Lupinus jonesii Rydb., Bull. Torrey Bot. Club 30: 256. i903. Type: Silver Reef, Washington Co., Utah, 3 May 1894, M. E. Jones 5143 (us). Status: Endemic, rare and threat- ened. Distribution: Washington Co., Utah (bry). Lupinus marianus Rj-^db., Bull. Torrey Bot. Club 34: 41. 1907. Type: Along Bullion Creek above Marysvale, Piute Co., Utah, 21 Jul 1905, Rydberg & Carlton 7024 (ny). Status: Endemic, restricted and threatened (Ripley, T). Distribution: Piute Co., Utah (bry). Oxytropis jonesii Barneby, Proc. Calif. Acad. Sci. IV, 27: 215. 1952. Type: Red Canyon, Garfield Co., Utah, 7 Jun 1947, 7,150 feet, Ripley & Barneby 8550 (gas). Status: Endemic, edaphically re- stricted and threatened. Distribution: Emery (us), Garfield (bry), Iron (bry, us) and Uintah (bry, NY, UTc) counties, Utah. Oxytropis multiceps Torr. & Gray, Syn. Fl. N. Amer. 1: 341. 1838. Status: Rare and local, possibly en- dangered. Distribution: Daggett Co., Utah, and in Wyoming and Colorado (Barne- by 1952).' Oxytropis obnapiformis C. L. Porter, Ma- drono 9: 133. 1947. Status: Rare and local, possibly threatened (Ripley, E). Distribution: Daggett Co., Utah; also in Wyoming and Colorado (bry). Peter ia thompsonae S. Wats., Amer. Nat- uralist 7: 300. 1873. Type: Kanab, Kane Co., Utah, 1872, Thompson s.n. (gh). St.\tus: Restricted in disjunct pop- ulations, neither threatened nor endan- gered (Ripley, T). Distribution: Emery, Grand, Kane, San Juan, and Washington counties, Utah; Mohave Co., Arizona, Nevada, and Idaho (bry,- Porter 1956). Psoralea aromatica Pavson, Bot. Gaz. 60: 379. 1915. Dec. 1975 WELSH. ET AL.: ENDANGERED UTAH PLANTS 353 Status: Edaphically restricted, local and rare; possibly threatened. Distribution: Emery and Grand counties, Utah, and Montrose Co., Colorado (bry; Ockendon 1965). Psoralea castorea S. Wats., Proc. Amer. Acad. Arts 14: 291. 1879. Type: Near Beaver City, Beayer Co., Utah, 1877, Palmer 96 (gh).'^ Status: Unknown from Utah in contemporary collection, although a specimen without collector or date labeled P. castoria is on file at ut; either extirpated from the state, or, and more likely, never from Utah. Distribution: Arizona and adjacent southern California (Kearney & Peebles 1951; Munz &Keck 1959).' Psoralea epipsila Barneby, Leafl. W. Bot. 3: 193. 1943. Type: Ca 17 miles east of Kanab to- ward Jepson Springs, Kane Co., Utah, 6 Jun 1942, Ripley & Barneby 4832 (gas). Status: Mohave corridor endemic, rare and endangered (Ripley, T). Distribution: Kane Co., Utah, and adjacent Coconino Co., Arizona (Bar- neby 1943). Psoralea juncea Eastw., Proc. Calif. Acad. Sci. II, 6: 286. 1896. Type: Epsom Creek, San Juan Co., Utah, 13 Jul 1895, Eastwood 21 (gas). Status: Navajo Basin endemic, edaphically restricted but abundant to common, neither threatened nor endan- gered. Distribution: Garfield, Kane, and San Juan counties, Utah (bry-), and Coconino Co., Arizona. Psoralea lanceolata Pursh var. steno- phylla (Rydb.) Toft & Welsh, Great Basin Nat. 32: 85. 1972, based on P. stenophylla Rydb., Bull. Torrey Bot. Club 40: 46. 1913. Type: Proposed dam site near Wil- son Mesa, Grand Co., Utah, 1 Jul 1911, Rydberg & Garrett 8367 (ny). Status: Endemic, locally common and neither threatened nor endangered. Distribution: Garfield, Grand, Kane, San Juan and Wayne counties, Utah (Toft & Welsh 1972). 'It seems likely that Palmer mislabled the type material, a Arizona. The species is not known from Utah. Psoralea pariensis Welsh & Atwood, spec, nov. P. megalanthae Wooton & Stand- ley proxime affinis sed floribus brevior- ibus et venas supra valde albo-strigosas. Plantae habentes caudices bene-evo- luti et radices tuberosae, 2-8 cm altae; caules 0.5-3 (4.5) cm longi, internodiis 1-3 elongatis, strigosi; stipulae 4-7 (10) mm longae, ovatae vel obovatae, strigosae; foliola 3-5, 9-23 (25) mm longa, 7-22 mm lata, obovata vel orbicularia, cuneata, rotundata ad truncata vel emarginata apicaliter, utrinque glandulifera; petioli 1.3-6.3 (7) cm longi, strigosi, pilis ap- pressis vel adscendentibus; pedunculi 0.5- 2.2 (2.8) cm longi, pilis appressis vel ad- scendentibus; bracteae 4-6 (8) mm longae, ovatae vel obovatae, acuminatae abrupte, pilosae rigide; pedicelli (1) 2-3.8 mm longi, pilis adscendentibus; calyces plus minusve gibbosi basim, tubus 3.3-4 (4.6) mm longus, dentes inaequales, infimis 5.3- 6.8 mm longis, circa duplo latioribus quam lateralibus; corolla 8.8-10.5 (12) mm lon- ga, dente infimo calycis leviter longiori- bus; alae vexillo subaequalis, purpureo- maculatae ad apicem; fructus usque ad 9 mm longum et seminum usque ad 5.2 mm longum. Type: UTAH: Garfield Co.: Bryce Canyon National Park, in ponderosa pine woods, as ground layer, at ca 8,000 feet elevation, 26 Jun 1975, Welsh & Murdoch 12859. Holotype, bry, Isotypes to be distributed. Additional spegimens examined: UTAH: Garfield Co.: East Creek, 3 miles south of Inspiration Point, Bryce Canyon N.P., in black sagebrush area, 11 Jun 1970, Buchanan 1494 (bry, wsGo); Paria View, Bryce Canyon N.P., in open ponderosa pine woods, ca 8,000 feet elevation, 12 Jun 1975, S. L. & S. L. Welsh 12810 (bry); East Creek, 9 Jun 1931, Weight B-31/ 6-305 (bcnp, us). Kane Co.: Hackberry Can- yon in Cottonwood Wash, ca 10 miles north of U.S. Highway 89, sec. 31, T. 40S., R. IW., 26 Apr 1972, Atwood 3684 (bry). Status: Endemic, rare and threat- ened. Distribution: Kane and Garfield counties, Utah. I tliat it was in leality from "Beaverdam." Mohave Co.. 354 GREAT BASIN NATURALIST Vol. 35, No. 4 Parryella filifolia Torr. & Gray in Gray, Proc. Amer. Acad. Arts 7: 397. 1868. Status: Restricted and rare; threat- ened. Distribution: Grand and San Juan counties, Utah (bry); also in northern Arizona and New Mexico. Psorothamnus thompsonae (Vail) Welsh & Atwood, comb, nov., based on Paro- sela thompsonae Vail, Bull. Torrey Bot. Club 24: 18. 1897. Type: Kanab, Kane Co., Utah, or pos- sibly Mohave Co., Arizona, 1872, Thompson s.n." (ny). Status: Endemic (?); restricted but locally common and neither threatened nor endangered. Distribution: Emery, Garfield, Kane, San Juan, and Wayne counties, Utah (bry and personal observations). Robinia neomexicana A. Gray, Mem. Amer. Acad. Arts II, 5: 314. 1855. Status: Restricted and rare, possibly threatened. Distribution: Garfield (ut) and Washington counties, Utah (bry) ; widespread elsewhere. Gentianaceae Gentianella tortuosa (M. E. Jones) Gil- lett, Ann. Missouri Bot. Gard. 44: 248. 1957, based on Gentiana tortuosa M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 707. 1895. Type: Panguitch Lake, Garfield Co., Utah, 7 Sep 1894, 8,400 feet, M. E. Jones 6008 (pom). Status: Restricted but locally com- mon, neither threatened nor endan- gered. Distribution: Garfield, Iron, Kane, Sanpete counties, Utah (bry), and Clark Co., Nevada (Gillett 1957). Geraniaceae Geranium marginale Rydb. ex Hanks & Small, N. Amer. Fl. 25: 16. 1907. Type: Aquarius Plateau at the head of Poison Creek, Garfield Co., Utah, 4 Aug 1905, Rydberg & Carlton 7401 (ny). Status: Endemic, restricted and possibly threatened (Ripley, T). Distribution: Garfield, Sevier, and Wayne counties, Utah (Nebeker 1974). Hydrophyllaceae Hydrophyllum fendleri (A. Gray) A. A. Heller,' PI. World 1: 23. 1897, based on H. occidentale var. fendleri A. Gray, Proc. Amer. Acad. Arts 10: 314. 1875. Status: Restricted and rare, possibly threatened. Distribution: San Juan Co., Utah (uT. UTc); widespread from Washing- ton and Oregon to Wyoming, Colorado and New Mexico, with var. fendleri restricted to the Rocky Mountain area (Constance 1942). Nama retrorsum J. T. Howell, Leafl. W. Bot. 5: 149. 1949. Status: Restricted and local, possibly threatened (Ripley, T) . Distribution: Garfield, Grand, and Kane counties, Utah (bry), and north- eastern Arizona (Atwood, in press). Nemophila parviflora Dougl. ex Benth. var. austinae (Eastw.) A. Brand, Pflanzenr. IV (Heft 251): 55. 1913, based on A^. austinae Eastw., Bull. Tor- rey Bot. Club 28: 143. 1901. Status: Restricted and local, possibly threatened. Distribution: Weber Co., Utah; widespread in Idaho, Nevada, Califor- nia, Oregon, and Washington (Atwood, in press) . Phacelia anelsonii J. F. Macbride, Contr. Gray Herb. 49: 26. 1917. Status: Restricted and local; threat- ened (Ripley, T) . Distribution: Washington Co., Utah (bry) ; southern Nevada and adjacent California (Atwood, in press). Phacelia argillacea Atwood, Phvtologia 26: 437. 1973. Type: Clear Creek, in Spanish Fork Canyon along the railroad on the east side of U.S. Highway 50-6, Utah Co., Utah, Atwood et al. 309i (bry). Status: Endemic, rare and endan- gered (Ripley, E). 'It is likely that sonic of llic collections sent to (iiay at Haivaid University iwho, in turn, i^avd tlicm to \\ntson) niay not liavc been collected by Mrs. Thompson, hut others with John Wesley Powell. It is known that l)otii Powell and Bishoj) collected plants and gave them to Ellen Thompson who did not collect outside of the Kanab area of Kane Co., Utah (Cron- quist ct al. 1972). Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PEANTS 355 Distribution: Utah Co., Utah (At- wood 1973, in press). Phacelia cephalotes A. Gray, Proc. Amer. Acad. Arts 10: 325. 1875. Type: Valley of the Virgin River, Washington Co., Utah, May 1874, Parry 179 (gh). Status: Restricted and local, pos- sibly threatened (Ripley, T). Distribution: Kane and Washing- ton counties, Utah, and Mohave and Navajo counties, Arizona (Atwood, in press). Phacelia constancei Atwood, Rhodora 74: 451. 1972. Status: Edaphically restricted and local; threatened (Ripley, T). Distribution: Emery, Garfield, Kane, and San Juan counties, Utah, and Mohave Co., Arizona (Atwood 1975, in press). Phacelia crenulata Torr. ex Wats, in King var. angustifolia Atwood, Great Basin Nat. 35: 158. 1975. Status: Restricted and in disjunct populations, neither threatened nor endangered. Distribution: Beaver, Garfield, and Kane counties, Utah, and in Coconino and Mohave counties, Arizona (At- wood 1975, in press). Phacelia demissa A. Gray var. heterotri- cha J. T. Howell, Amer. Midi. Nat- uralist 29: 8. 1943. Type: Marysvale, Piute Co., Utah, 4 Jun 1894, M. E. Jones 5388a (pom). Status: Endemic, restricted and pos- sibly threatened (Ripley, T). Distribution: Piute, Sevier, and Wayne counties, Utah (Atwood, in press). Phacelia howelliana Atwood, Rhodora 74: 456. 1972. Type: Ca 0.4 miles north of Bluff along Utah Highway 163, San Juan Co., Utah, 13 May 1970, Atwood 2454 (bry) . Status: Endemic, restricted but lo- cally abundant, neither threatened nor endangered. Distribution: Grand and San Juan counties, Utah (bry; Atwood 1975, in press). Phacelia indecora J. T. Howell, Amer. Midi. Naturalist 29: 12. 1943. Type: Bluff, San Juan Co.. Utah, 24 May 1919, M. E. Jones s.n. (gas). Status: Endemic, rare and endan- gered (Ripley, T). Distribution: Wayne and San Juan counties, Utah (Atwood, in press). Phacelia mammillarensis Atwood, Phyto- logia 26: 437. 1973. Type: Ca 6 miles east of Glen Can- yon City, along the road to Warm Creek, Kane Co., Utah, 6 May 1970, Welsh & Atwood 9809 (bry). Status: Endemic, edaphically restric- ted and endangered (Ripley, E). Distribution: Garfield and Kane counties, Utah (Atwood 1975, in press). Phacelia rafaelensis Atwood, Rhodora 74: 454. 1972. Type: Capitol Reef N.M., Wayne Co., Utah, 12 Jun 1969, Atwood & Hig- gins 1834 (bry). Status: Restricted and local, threat- ened (Ripley, T). Distribution: Emery, Washington, and Wayne counties, Utah, and Coco- nino Co., Arizona (Atwood 1975, in press). Phacelia salina (A. Nels.) J. T. Howell, Leafl. W. Bot. 4: 16. 1944, based on Emmenanthe salina A. Nels., Bull. Torrey Bot. Club 24: 381. 1898, a sub- stitute name for E. foliosa M. E. Jones, Zoe 4: 278. 1893, not P. foliosa Phillip. Type: Deep Creek Valley, above Furber, Tooele Co., Utah, 8 Jun 1891, M. E. Jones s.n. (pom). Status: Rare and possibly extirpated from Utah. Distribution: Tooele Co., Utah, and adjacent Nevada and southern Wyo- ming (Atwood, in press). Phacelia tetramera J. T. Howell, Leafl. W. Bot. 4: 16. 1944, based on Emme- nanthe pusilla A. Gray, Proc. Amer. Acad. Arts 11: 87. 1876, not P. pusilla Buckley nor Torr. Status: Rare and possibly threatened. Distribution: Weber Co., Utah (Arnow 3985 [bry, ut]); also in Ne- vada, California and Oregon (Howell 1944). Phacelia utahensis E. G. Voss, Bull. Tor- rey Bot. Club 64: 135. 1937. 356 GREAT BASIN NATURALIST Vol. 35, No. 4 Type: Gunnison, Sanpete Co., Utah, 7 Jun 1910, M. E. Jones s.n. (pom). Status: Endemic, edaphically re- stricted and threatened (Ripley, T). Distribution: Carbon (ut), Sanpete, and Sevier counties, Utah (Atwood 1975, in press). JUNCACEAE Juncus bryoides F. J. Hermann, Leafl. W. Bot. 5: 117. 1948. Status: Restricted, rare and possibly threatened. Distribution: Salt Lake and Wash- ington counties, Utah, and in Califor- nia (Hermann 1948). Lamiaceae Stachys rothrockii A. Gray, Proc. Amer. Acad. Arts 12: 82.1876. Status: Restricted, rare and possibly endangered. Distribution: Near Orderville, Kane Co., Utah (bry,- Barneby 1944), north- ern New Mexico and Arizona. LiLIACEAE Agave utahensis Engelm. in King, Rep. Geol. Explor. 40th Parallel 5: 497. 1871. Type: About St. George, Washing- ton Co., Utah, Palmer s.n. (mo). Status: Restricted and rare except locally; commercially exploited and possibly endangered. Our plant is var. utahensis. Distribution: Washington Co., Utah (bry) ; also in Coconino and Mohave counties, Arizona (Breitung 1960). Allium passeyi Holmgren & Holmgren, Brittonia 26: 309. 1974. Type: Southwest part of Howell Valley, sec. 7, T.llN., R.5W., Box Elder Co., Utah, 13 Jun 1960, 4,800 feet, Holmgren et al. 15125 (ny). Status: Endemic, rare and endan- gered (Ripley, E). Distribution: Box Elder Co., Utah; known only from the type locality (Holmgren & Holmgren 1974). Eremocrinum albomarginatum (M. E. Jones) M. E. Jones, Zoe 4: 53. 1893, based on Hesperanthes albomarginata M. E. Jones, Zoe 2: 251. 1891. Type: Green River, Emery Co., Utah, 9 May 1890, M. E. Jones s.n. (pom). Status: Navajo Basin endemic, edaphically restricted but locally com- mon to abundant and neither threat- ened nor endangered. Distribution: Emery, Garfield, Grand, Kane, San Juan, and Wayne counties, Utah, and Apache and Nava- jo counties, Arizona (MacDougall 1973). Nolina microcarpa S. Wats., Proc. Amer. Acad. Arts 14: 247. 1879. Status: Apparently extirpated from Utah; last seen in Zion N.P. in 1925 (Cottam 1974). Distribution: Washington Co., Utah, to northern Mexico, eastward to western Texas. This plant has mistakenly been called Nolina parry i (Kearney & Peebles 1951) or A^. bigelovii (Cottam 1974). Yucca brevifolia Engelm. in King, Rep. Geol. Explor. 40th Parallel 5: 496. 1871. Status: Restricted but locally abun- dant; possibly threatened by commer- cial exploitation. Distribution: Southwestern Wash- ington Co., Utah; western Arizona, southern Nevada and adjacent Califor- nia (McKelvey 1938; Weber 1953). Yucca toftiae Welsh, Great Basin Nat. 34: 308. 1974. Type: Three Garden, ca 1 mile north of the confluence of San Juan and Colo- rado rivers. Lake Powell, San Juan Co., Utah, 4 Jun 1973, Welsh 11955a (bry) . Status: Endemic, local and uncom- mon; possibly threatened. Distribution: Kane and San Juan counties, Utah (bry); much of the known range of this entity has been inundated by the water of Lake Powell. Zigadenus vaginatus (Rvdh.") J. F. Mac- bride, Contr. Gray Herb. 53: 4. 1918, based on Anticlea vaginata Rvdb., Bull. Torrey Bot. Club 39: 108. 1912. Type: Armstrong Canyon, near the Natural Bridges, San Juan Co., Utah, 4-6 Aug 1911. Rydberg & Garrett 9407 (ny). Status: Endemic, restricted and pos- sibly endangered. Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 357 Distribution: Grand, Kane, and San Juan counties, Utah (bry). This distinctive species was placed in synonymy under the Mexican species, Zigadenus volcanicus Benth., PI. Hart- wegianus 96. 1840, by I'idestrom (1925), Tidestrom and Kittell (1941), and fol- lowed by Holmgren and Reveal (1967). This plant is related to Z. elegans Pursh, but differs in having an elongated panicu- late inflorescence and smaller white to greenish flowers borne in the late summer and early fall. LOASACEAE Mentzelia argillosa J. Darlington, Ann. Missouri Bot. Gard. 21: 153. 1934. Type: Vermillion, Sevier Co., Utah, 16 Jul 1894, 5,600 feet, M. E. Jones 563^ (mo). Status: Endemic, edaphically re- stricted and threatened. Distribution: Sevier Co., Utah (bry). Mentzelia obscura Thompson & Roberts, Phytologia 21: 284. 1971. Status: Local and infrequent. Distribution: Washington Co., Utah; southern Nevada and western Arizona westward to California and Baja California, Mexico. Petalonyx parry i A. Gray, Proc. Amer. Acad. Arts 10: 72. 1874. Type: St. George, Washington Co., Utah, "within a stone's throw of the great Mormon Temple" (see Parry 1875), 1874, Parry 75 (gh). Status: Rare and possibly endan- gered. Distribution: Washington Co., Utah (bry), northern Arizona and southern Nevada (Davis & Thompson 1967). LOGANIACEAE Buddie j a utahensis Coville, Proc. Biol. Soc. Wash. 7: 69. 1892. Type: Near St. George, Washington Co., Utah, 1877, Palmer s.n. (us). Status: Restricted, rare and possibly threatened. Distribution: Washington Co., Utah; also in northwestern Arizona, southern Nevada and southeastern Cali- fornia (Munz 1974). Loranthaceae Phorandendron californicum Nutt., J. Acad. Nat. Sci. Philadelphia II, 1: 185. 1848. Status: Restricted but locally abun- dant; neither threatened nor endan- gered. Distribution: Washington Co., Utah; widespread in Arizona, Nevada, southern California, and northern Mex- ico (Kearney & Peebles 1951). Malvaceae Sphaeralcea caespitosa M. E. Jones, Contr. W. Bot. 12: 4. 1908. Type: Wah Wah, Beaver Co., Utah, 1906, 6,000 feet, M. E. Jones s.n. (pom) . Status: Endemic, rare and threat- ened. Distribution: Beaver and Millard counties, Utah (bry; Jefferies 1972). Martyniaceae Proboscidea louisiana (Miller) ThelL, Mem. Soc. Sci. Nat. Cherbourg 3: 480. 1912, based on Martynia louisiana Mil- ler, Gard. Diet, ed 8, Martynia No. 3. 1768. Status: Restricted and uncommon, but neither rare nor endangered. Distribution: Washington Co., Utah (bry); widespread in the southwestern states and elsewhere. Najadaceae Najas caespitosus (Maguire) Reveal, stat. & comb, nov., based on Najas flexilis ssp. caespitosus Maguire in Maguire & Jensen, Rhodora 44: 7. 1942. Type: Common in shallow water. Pelican Point, Fish Lake, Sevier Co., Utah, 3 Aug 1940, 8,600 feet, Maguire 19888 (ny). Status: Endemic and endangered. Distribution: Sevier Co., Utah; known onl}' from the type area (Ma- guire & Jensen 1942). Najas guadalupensis Morong, Mem. Tor- rey Bot. Club 3: 60. 1893. Status: Rare and restricted. Distribution: Cache Co., Utah (uTC; Maguire & Jensen 1942); wide- spread elsewhere. 358 GREAT BASIN NATURALIST Vol. 35, No. 4 Nyctaginaceae Boerhaavia torreyana (S. Wats.) Stand- ley, Contr. U.S. Natl. Herb. 12: 385. 1909, based on B. spicata var. torrey- ana S. Wats., Proc. Amer. Acad. Arts 24: 70. 1889. Status: Local, rare and highly re- stricted; possibly threatened. Distribution: Kane Co., Utah {At- wood 3389 [bry, wts]). Hermidium alipes S. Wats. var. pallidum C. L. Porter, Rhodora 54: 158. 1952. Type: Ca 5 miles south of Vernal, Uintah Co., Utah, 3 Jun 1950, 5,200 feet, Porter 5308 (rm). Status: Endemic, rare and endan- gered. Distribution: Uintah Co., Utah (Porter 1952). Oleaceae Menodora scabra A. Gray, Amer. J. Sci. Arts II, 14: 44. 1852. Status: Rare, in disjunct populations, and possibly threatened. Distribution: Garfield and Wash- ington counties, Utah (bry) ; wide- spread in the southwestern states and northern Mexico (Kearney & Peebles 1951). Onagraceae Epilobium nevadense Munz, Bull. Torrey Bot. Club 56: 166. 1929. Status: Rare and threatened (Rip- ley, T). Distribution: Washington Co., Utah (Higgins 1972b), and Clark Co., Nevada. Oenothera brevipes A. Gray var. pallidula Munz, Amer. J. Bot. 15: 229. 1928. Status: Restricted and local, possibly threatened. Distribution: Washington Co., Utah (bry) ; also in northwestern Arizona, southern Nevada and southern Califor- nia (Raven 1969). Oenothera gouldii (Raven) Welsh & At- wood, comb, nov., based on Camissonia gouldii Raven, Contr. U.S. Natl. Herb. 37: 368. 1969. Type: Steep slope of volcanic cone among loose cinders, Diamond Valley, 12 miles north of St. George, Wash- ington Co., Utah, 15 Oct 1941, 3,500 feet, Gould 1423 (pom). Status: Restricted and rare, pos- sibly threatened. Distribution: Washington Co., Utah, and Coconino Co., Arizona (Ra- ven 1969). Oenothera megalantha (Munz) Raven, Univ. Calif. Publ. Bot. 34: 111. 1962,. based on O. heterochroma var. mega- lantha Munz, Leafl. W. Bot. 3: 52. 1941. Status: Restricted and rare, endan- gered (Ripley, E). Distribution: Kane Co., Utah (bry), and Nye Co., Utah (Raven 1969; Beat- ley 1973). Oenothera parryi S. Wats., Amer. Nat- uralist 9: 270. 1875. Type: Near St. George, Washington Co., Utah, 1874, Parry 72 (gh). Status: Restricted but locallv abun- dant (Ripley, T). Distribution: Washington Co., Utah, and adjacent Arizona (bry). Ophioglossaceae Botrichium boreale (Fries) Milde var. obtusilobum (Rupr.) M. Braun, Index N. Amer. Ferns 33. 1938. Based on B. crassinervum var. obtusilobum Rupr., Beitr. Pflanzenk. Russ. Reiches 11: 42. 1959. Status: Rare; possibly threatened. Distribution: Summit Co., Utah according to Flowers 1944; not reported from Utah by Cronquist et al. 1972); widespread in northwestern America. Botrychium lanceolatum (S. G. Gmelin) Angstrom, Bot. Not. 1854: 68. 1854, based on Osmunda lanceolata S. G. Gmelin, Novi Comment. Acad. Petrop. 12: 516-1768. St.\tus: Rare; possibly threatened. Distribution: Juab Co., Utah (Flowers 1944; Maguire & Holmgren 1946); widely distributed in northern North America (Welsh 1974b). Botrichium matricariifolium A. Braun in Koch var. hesperium (Maxon & Clau- sen) M. Braun, Index N. Amer. Ferns 39. 1938, based on B. matricariifolium ssp. herperium Maxon & Clausen, Mem. Torrey Bot. Club 19: 38. 1938. Status: Rare; possibly threatened. Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PL.'\NTS 359 Distribution: Salt Lake Co., Utah (Flowers 1944; not reported by Cron- quist et al. 1972 as, in their opinion, Flowers' identification was in error); also in Colorado. Orchidaceae Cypripcdiurn calceolus L. xar. parviflo- rum (Salisb.) Fern., Rhodora 48: 4. 1946, based on C. parviflorum Salisb., Trans. Linn. Soc. London 1: 77. 1791. Status: Extirpated or endangered. Distribution: Cache (utc) and Utah (bry) counties, LTtah; widespread in northern North America. Cypridedium fasciculatum Kellogg ex Wats., Proc. Amer. Acad. Arts 17: 380. 1882. Status: Rare and endangered. Distribution: Salt Lake (bry) and Summit (Tidestrom 1925) counties, Utah; widespread and rare in Wash- ington, California, Idaho, Montana, Wyoming, and Colorado. Spiranthes cernua (L.) Rich., Mem. Mus. Hist. Nta. 4: 59. 1818, based on Ophrys cernua I.., Sp. PI. 946. 1753. Status: Extirpated from Utah. Distribution: Salt Lake Co., Utah (M. E. Jones 1908 [pom]); widespread in the eastern United States (Correll 1950). Papaveraceae Papaver radicatum Rottb., Skr. Ki0ben- havnske Selsk. Laerd. Elsk. 10: 455. 1770. Status: Rare and restricted. Distribution: Duchesne Co,, Utah (bry) ; \\ddespread in northern North America, circumboreal (Welsh 1974b). Arcotmecon humilis Coville, Proc. Biol. Soc. Wash. 7: 67. 1892. Type: St. George, Washington Co., Utah, 1874, Parry s.n. (gh). Status: Restricted and rare, endan- gered (Ripley, E) . Distribution: Washington Co., Utah, and Mohave Co., Arizona (bry). PiNACEAE Pinus longaeva D. K. Bailey, Ann. Mis- souri Rot. Card. 57: 243. 1970. Status: Restricted but hardly rare; possibly threatened. Distribution: Beaver (personal ob- servations). Carbon, Duchesne, Gar- field, Iron (personal observations), Kane, Millard, and Washington coun- ties, Utah; also in Nevada and eastern California (Bailey 1970). Poaceae Andropogon glomeratus (Walter) Britton, Sterns & Poggenb., Prelim. Cat. Antho. & Pterid. within 100 mi. New York City 67. 1888, based on Cinna glomerata Walter, Fl. Carol. 59. 1788. Status: Rare and possibly extirpated from Utah, previously known only along Lake Powell. Distribution: Kane and San Juan counties, Utah (bry); widely distrib- uted in southern and southeastern United States, West Indies, Yucatan and Central America. Blepharidachne kingii (S. Wats.) Hackel in DC, Monogr. Phanerog. 6: 261. 1889, based on Eremochloe kingii S. Wats, in King, Rep. Geol. Explor. 40th Parallel 5: 382. 1871. Status: Rare and local. Distribution: Beaver, Millard, and Tooele counties, Utah, and adjacent Nevada (bry). Festuca dasyclada Hackel ex. Beal, Grasses N. Amer. 2: 602. 1896. Type: Joe's Valley, Emery Co., LTtah, 1875, Parry s.n. (us). Status: Endemic and possibly en- dangered (Ripley, PoEx). Distribution: Emery and Sanpete counties, Utah (N. H. Holmgren, per- sonal communication), and Colorado (N. West, personal communication).^" Festuca sororia Piper, Contr. U.S. Natl. Herb. 16: 197. 1913. Status: Rare and restricted. Distribution: Southeastern Utah (Hitchcock & Chase 1950). Helictotrichon mortonianum (Lams.- Scribn.) Henrard, Blumea 3: 429. 1940, based on Avena mortonianum Lams.-Scribn., Bot. Gaz. 21: 133. 1896. Status: Rare and endangered. Distribution: Utah (Hitchcock & ^"Until 1975 this species was known only from the type. 360 GREAT BASIN NATURALIST Vol. 35, No. 4 Chase 1950) ; also in Colorado and New Mexico. Imperata brevifolia Vasey, Bull. Torrey Bot. Club 13: 26. 1886. Status: Rare, restricted and endan- gered or possibly extirpated from Utah. Distribution: San Juan Co., Utah (bry) ; California, Nevada, and Mexico (Hitchcock & Chase 1950). The known localities in Utah where this grass occurred have been inundated by Lake Powell. Muhlenbergia arsenei Hitchc, Proc. Biol. Soc. Wash. 41: 161. 1928. Status: Rare and local. Distribution: Garfield (bry), Kane (bry), San Juan (ut), and Washing- ton (utc) counties, Utah; New Mexico and Cahfomia (Cottam et al. 1940). Muhlenbergia curtifolia Lams.-Scribn., Bull. Torrey Bot. Club 38: 328. 1911. Type: Between Kanab and Carmel, Kane Co., Utah, M. E. Jones 6047 (us). Status: Restricted and local, pos- sibly threatened. Distribution: Garfield, Kane, San Juan, and Washington (ut) counties, Utah (bry, utc) ; southern Nevada and northern Arizona (Hitchcock & Chase 1950). Muhlenbergia minutissima i(Steudel) Swallen, Contr. U.S. Natl. Herb. 29: 207. 1947, based on Agrostis minutis- sima Steudel, Syn. PI. Glum. 1: 171. 1854. Status: Local and rare. Distribution: Washington Co., Utah (Atwood & Higgins 5528 [bry]). Muhlenbergia wrightii Vasey in Coult., Manual Bot. Rocky Mt. Region 409. 1885. Status: Rare and local. Distribution: Utah (Hitchcock & Chase 1950); Arizona, Colorado, Okla- homa, New Mexico, and northern Mexico. Puccinellia simplex Lams.-Scribn., U.S.D.A. Div. Agrostol. Circ. 16: 1. 1899. Status: Rare and possibly threatened. Distribution: Weber Co., Utah {Arnow 3986 [bry]); also in Califor- nia (Hitchcock & Chase 1950). Sporobolus pulvinatus Swallen, J. Wash. Acad. Sci. 31: 351. 1941. Status: Rare and possibly threat- ened. Distribution: San Juan Co., Utah (bry); New Mexico, Arizona, Texas and northern Mexico (Hitchcock & Chase 1950). POLEMONIACEAE Gilia caespitosa A. Gray, Proc. Amer. Acad. Arts 12: 80. 1876. Type: Rabbit Valley on barren cliffs of sandstone, Wayrte Co., Utah, 1875, 7,000 feet, Ward s.n. (gh). Status: Endemic, rare and endan- gered (Ripley, E). Distribution: Wayne Co., Utah (bry, utc). Gilia lad folia S. Wats, ex Parry, Amer. Naturalist 9: 347. 1875. Type: Valley of the Virgin, near St. George, Washington Co., Utah, 1874, Parry 188 (gh). Status: Rare and local; possibly threatened. Distribution: Kane, Wayne, and Washington counties, Utah (bry) ; Ari- zona, southern Nevada and southern California (Matthews 1971). Gilia mcvickerae M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 712. 1895. Type: Marysvale, Piute Co., Utah, 7,000 feet, M. E. Jones 5378 (pom). Status: Endemic, rare and local; threatened (Ripley, T). Distribution: Piute, Sevier, and Uintah counties, Utah (bry). Gilia stenothyrsa A. Gray, Proc. Amer. Acad. Arts 8: 276. 1870. Type: In a clear forest, Uinta Mts., Duchesne or Uintah counties, Utah, 1844, Fremont s.n. (gh). Status: Endemic, locally common and neither threatened nor endangered. Distribution: Carbon (ut), Emery, Duchesne and Uintah counties, Utah (bry, utc). Phlox cluteana A. Nels., Amer. J. Bot. 28: 24. 1922. St.a>tus: Rare and local; possibly threatened (Ripley, T). Distribution: San .Juan Co., Utah (bryj, and northern Arizona (Kearney & Peebles 1951). Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 361 Phlox gladiformis (M. E. Jones) E. Nels., Rev. West. N. Amer. Phloxes 21. 1899, based on P. longifolia var. gladiformis M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 711. 1895. Type: Cedar City, Iron Co., Utah, 11 May 1894, 6,500 feet, M. E. Jones 5208c (pom). Status: Rare and local; possibly threatened (Ripley, T). Distribution: Garfield, Iron, and Washington counties, Utah (bry), and adjacent Nevada. Phlox grahomii WherrA', Brittonia 5: 63. 1943. Type: Talus slopes on west side of Green River, south of the mouth of Sand Wash. Uintah Co., Utah, 27 May 1933, Graham 7884 (cm). Status: Endemic, rare and local; threatened (Ripley, T). Distribution: Uintah Co., Utah; known only from the type locality. Phlox jonesii Wherry, Notul., Nat. Acad. Nat. Sci. Philadelphia 146: 8. 1944. Type: Zion Canyon, Washington Co., Utah, 7 May 1923, M. E. Jones s.n. (us). Status: Endemic, rare and threat- ened (Ripley, T) . Distribution: Washington Co., Utah; known only from the type lo- cality (Wherry 1955). Polygonaceae Eriogonum ammophilum Reveal, Phyto- logia 23: 163. 1972. Type: Ca 1.3 miles northwest of Ibex Warm Point, on a dry sandy flat, Millard Co., Utah, 4 Aug 1970," 5,270 feet, Holmgren & Holmgren 4650 (us). Status: Endemic, rare and local; endangered (Ripley, E). Distribution: Millard Co., Utah (bry, DERM, UTC). Eriogonum aretioides Barneby, Leafl. W. Bot. 5: 154. 1949. Type: Bare limestone gravel benches in the foothills of the Escalante Range at Widtsoe, Garfield Co., Utah, 8 Jun 1947, 7,750 feet, Ripley & Barneby 8570 (c.-^s). Status: Endemic, edaphically re- stricted and endangered (Ripley, E) . Distribution: Garfield Co., Utah (BRY, UT, UTC). Eriogonum hatemanii M. E. Jones, Contr. W. Bot. 11: 11. 1903. Type: Price, Carbon Co., Utah, 29 Jun 1898, M. E. Jones s.n. {pom). Status: Restricted and local, neither threatened nor endangered. Distribution: Carbon, Duchesne, Emery, Garfield, and Uintah counties, Utah, and Rio Blanco Co., Colorado (Reveal 1973a). Eriogonum brevicaule Nutt. var. cottamii (S. Stokes) Reveal, Great Basin Nat. 32: 113. 1972, based on E. tenellum ssp. cottamii S. Stokes, Gen. Eriog. 70. 1936. Type: Canyons in bottoms of the slopes of West Mtn., Utah Co., Utah, 20 Aug 1925, Cottam 411 (bry). Status: Endemic, restricted and rare. Distribution: Juab, Millard, and Utah counties, Utah (bry. ny, utc). Eriogonum brevicaule Nutt. var. wa- satchense (M. E. Jones) Reveal, Great Basin Nat. 32: 113. 1972, based on E. wasatchense M. E. Jones, Contr. W. Bot. 11: 11. 1903. Type: American Fork Canyon, Utah Co., Utah, 27 Jul 1880, M. E. Jones 1877 (pom). Status: Endemic, restricted and rare. Distribution: Davis, Juab, Millard, Salt Lake, Utah, and Weber counties, Utah (bry, ds, gh, ut, utc). Eriogonum clavellatum Small, Bull. Tor- rey Bot. Club 25: 48. 1898. Type: Barton Range, San Juan Co., Utah, 13 Jul 1895, Eastwood 132 (ny). Status: Rare and highly restricted; threatened (Ripley, T). Distribution: San Juan Co., Utah (bry, utc), and Montezuma Co., Colo- rado (cs). Eriogonum contortum Small ex Rydb., Agric. Exp. Sta. Agric. Coll. Colorado Bull. (Fl. Colorado) 100: 107. 1906. Status: Infrequent but neither threatened nor endangered. Distribution: Grand Valley en- demic in Grand Co., Utah (bry, utc), and Garfield and Mesa counties, Colo- rado. 362 GREAT BASIN NATURALIST Vol. 35, No. 4 Eriogonum corymbosum Benth. in DC. var. davidsei Reveal, Great Basin Nat. 27: 216. 1968. Type: Ca 0.7 miles south of U.S. Highway 50-6 at Wellington, just south of the Price River bridge, 9 Sep 1967, Reveal & Davidse 956 (utc). Status: Endemic, restricted and lo- cal; endangered (Ripley, E). Distribution: Carbon Co., Utah; known only from type locality. Eriogonum corymbosum Benth. in DC. var. divaricatum Torr. & Gray in Beck- with, Explor. & Surv. Railroad Route from Mississippi River to Pacific Ocean 2: 29. 1857. Type: Near Green River, Emery Co.. Utah, 1 Oct 1853, Creutzfeldt s.n. (ny). Status: Endemic, local and restric- ted; neither endangered nor threatened. Distribution: Emery, Garfield, and Grand counties, Utah. This form of Eriogonum corymbosum differs from var. corymbosum in having small, usually crenulate leaves, smaller more compact stature, and compact inflo- rescences. Eriogonum corymbosum Benth. in DC. var. revealianum (Welsh) Reveal, stat. & comb, nov., based on E. revealianum Welsh, Great Basin Nat. 30: 17. 1970. Type: Gravelly, boulder-strewn, east- facing slope near the head of the can- yon at milepost 26 south of Antimony along Utah Highway 22, Garfield Co., Utah, 4 Sep 1969, S. L. & S. L. Welsh 9389 (bry). Status: Endemic, rare and local; en- dangered. Distribution: Garfield Co., Utah; known only from the type locality. This form of Eriogonum corymbosum differs from var. corymbosum in having elongated, entire leaves concentrated near the base of elongated flowering stems and open, spreading inflores- Eriogonum cronquistii Reveal, Madrono 19: 289. 1969. Type: Loose decomposed granite talus slopes on the west side of Bull Mtn., Henry Mts., Garfield Co., Utah. 14 Aug 1967, 8,300 feet, Holmgren & Reveal 3010 (utc). Status: Endemic, restricted and local; endangered (Ripley, E). Distribution: Garfield Co., Utah; known only from the type locality (bry, NY, utc). Eriogonum desertorum (Maguire) R. J. Davis, Fl. Idaho 246. 1952, based on E. chrysocephalum ssp. desertorum Ma- guire, Leafl. W. Bot. 3: 11. 1941. Status: Restricted and rare; possibly threatened. Distribution: Box Elder and Tooele counties, Utah; northeastern Nevada and southern Idaho (Reveal 1973a). Eriogonum ephedroides Reveal, Madroiio 19: 295. 1969. Type: Ca 10 miles south of Bonanza along Utah Highway 45 south of the White River, 25 Jul 1965, Holmgren et al. 2265 (utc). Status: Uinta Basin endemic, re- stricted and rare; endangered (Ripley, E). Distribution: Uintah Co., Utah (bry, utc), and Rio Blanco Co., Colo- rado (ny). Eriogonum eremicum Reveal, Ph-s^tologia 23: 165. 1972. Type: Ca 17 miles southeast of Gar- rison along Utah Highway 21, Millard Co., Utah, 23 Jul 1965, Holmgren et al. 2247 (utc). Status: Endemic, restricted and rare; threatened (Ripley, T). Distribution: Millard Co., Utah (bry, utc). Eriogonum fasciculatum Benth. var. poli- folium (Benth. in DC.) Torr. & Gray, Proc. Amer. Acad. Arts 8: 169. 1870, based on E. polifolium Benth. in DC, iProdr. 14: 12. 1856. Status: Local and common to abun- dant, neither threatened nor endan- gered. Distribution: Emery (bry, ut,- where rare) and Washington (where common) counties, Utah; widespread and common in Arizona, Nevada, Cali- fornia, and Baja California, Mexico. Eriogonum grayi Reveal, Phvtologia 25: 193. 1973. Type: Lake Blanche, Salt Lake Co., Utah, 15 Aug 1947. Holmgren et al. 7121 (uTC). Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 363 Status: Endemic, disjunct and infre- quent. Distribution: Juab, Salt Lake, Utah, and Weber counties, Utah (bry, wsco. UTC; Reveal 1973a). Eriogonum humivagans Reveal, Madrono 19: 219. 1969. Type: Ca 13.5 miles east of Monti- cello, 13 Aug 1966, 6,800 feet, Holm- gren & Reveal 3001 (utc). Status: Endemic, restricted and rare; endangered (Ripley, E). Distribution: San Juan Co., Utah; known on!}' from the type locality. Eriogonum hylophilum Reveal & Brother- son, Great Basin Nat. 27: 190. 1968. Type: Along Utah Highway 53 in Gate Canyon, 2.7 miles southwest of the summit of the Badlands Cliffs, Du- chesne Co., Utah, 15 Aug 1966, 6,500 feet, Holmgren & Reveal 3017 (utc). Status: Endemic, rare and restricted; endangered (Ripley, E) . Distribution: Duchesne Co., Utah (Reveal 1968a; 1973a). Eriogonum intermontanum Reveal, Ma- drono 19: 293. 1969. Type: Ca 1.5 miles south of the Uin- tah Co. line at the head of Middle Canyon of West Water Creek drain- age in Roan Cliffs, Grand Co., Utah, 27 Jul 1965, 8,400 feet, Holmgren et al. 2278 (uTc). Status: Endemic, rare and local; en- dangered (Ripley, E). Distribution: Grand Co., Utah (bry, utc). Eriogonum jamesii Benth. in DC. var. rupicola Reveal, Phytologia 25: 202. 1973. Type: Along Utah Highway 15, 4.9 miles west of the east entrance to the park on Checkerboard Mesa, Zion N.P., Washington Co., Utah. 12 Aug 1972, Reveal & Reveal 2874 (us). Status: Endemic, restricted and rare; threatened (Ripley, T) . Distribution: Kane and Washing- ton counties, Utah (Reveal 1973a). Eriogonum lancifolium Reveal & Brother- son, Great Basin Nat. 27: 188. 1968. Type: On low hills 5 miles east of Wellington, Carbon Co., Utah, 9 Sep 1967, Reveal & Davidse 957 (utc). Status: Endemic, restricted and lo- cal; threatened (Ripley, E). Distribution: Carbon Co., Utah (bry, us, utc). Eriogonum leptocladon Torr. & Gray in Beckwith, Explor. & Surv. Railroad Route from Mississippi River to Pacific Ocean 2: 129. 1857. Type: Near the Green River, Emery Co., Utah, 1 Oct 1853, Creutzfeldt s.n. (ny). Status: Endemic, edaphically re- stricted but locally abundant and neither threatened nor endangered. Our plant is vai . leptocladon. Distribution: Emery, Garfield, Grand, San Juan, and Wayne counties, Utah (bry. UTC; Reveal 1966). Eriogonum leptophyllum (Torr. & Gray) Wooton & Standley, Contr. U.S. Natl. Herb. 16: 118. 1913, based on E. ef- fusum var. leptophyllum Torr. in Sitgr., Rep. Exped. Down Zuni & Colorado rivers 168. 1853. Status: Local and rare; threatened. Distribution: San Juan Co., Utah {Harrison 12163 [bry] ) ; southwestern Colorado, northeastern Arizona and ad- jacent New Mexico (Reveal 1968a). Eriogonum loganum A. Nels., Bot. Gaz. 54: 149. 1912. Type: Logan, Cache Co., Utah, 26 Jun 1909, Smith 1704 (rm). Status: Endemic, extremel}^ restric- ted and rare; endangered (Ripley, E). Distribution: Cache Co., Utah (bry. utc). Eriogonum nanum Reveal, Phytologia 25: 194. 1973. Type: Talus slopes and limestone outcrops south of Willard Peak, Box Elder Co., Utah, 31 Aug 1964, 9,500 feet. Reveal & Holmgren 665 (us). St.\tus: Endemic, restricted and rare; threatened (Ripley, T). Distribution: Box Elder and Weber counties, Utah (bry, utc, wsco). Eriogonum natum Reveal, spec. nov. A Eriogono hrevicaule Nutt. differt foliis ellipticis, 2-2.5 (3) cm longis et (8) 10-13 (15) mm latis, lanatis, inflores- centiis cymoso-umbellatis, floribus flavis, 2-2.5 (3) mm longis, glabris. Spreading herbaceous perennials 1-3.5 364 GREAT BASIN NATURALIST Vol. 35, No. 4 dm high, 1-4 dm across, with a short woody caudex arising from a stoutish, woody taproot; leaves essentially basal, the leaf-blade elliptic, 2-2.5 (3) cm long, (8) 10-13 (15) mm wide, densely to- mentose below, somewhat less so and greenish-tomentose above, the petiole (1) 2-3 cm long, tomentose; flowering stems erect to spreading, slender, 1-2 (2.5) dm long, white to greenish-tomentose; inflo- rescences cymose-umbellate. 3-10 (15) cm long, 3-5 (8) cm wide, trichotomously branched throughout, tomentose; bracts scalelike to foliaceous, ternate, the former 1-3 mm long, tomentose to floccose with- out, tomentose within, the latter 1-3 per node, linear-lanceolate to lanceolate, 5-10 (12) mm long, (1.5) 2-4 (5) mm wide, tomentose; peduncles lacking; involucres solitary or infrequently in groups of 2, turbinate-campanulate, 2.5-4 mm long, 2-3 mm wide, thinly to densely tomentose without, glabrous within, the 5 acute teeth 0.5-0.8 mm long, usually with a membranaceous margin, the bractlets linear-oblanceolate, 1.5-3 mm long, fringed with gland-tipped cells, the pedi- cels 2.5-5 mm long, glabrous; flowers bright yellow with golden yellow bases and golden to greenish midribs, 2-2.5 (3) mm long, glabrous, the tepals oblong to obovate, distinctly keeled at the base and along the midrib of each tepal, united about 14 to 1/5 the length of the flower; stamens exserted, 2.5-4 mm long, the fila- ments sparsely pilose basally, the anthers yellow, 0.3-0.5 mm long, oblong to oval; achenes light brown, 2-3 mm long, the globose base tapering to a long, 3-angled, slightly roughened beak. Type: UTAH: Millard Co.: Along U.S. Highway 50-6, 46.2 miles east of the Nevada state line and about 43 miles west of Delta, on low white alkaline clay out- crops 50-300 meters north of the highway, ca 0.2 miles east of the (hrt road junction to the Antelope Spring-Black Hill Well roads, north-northwest of Sevier Lake. 1 3 Aug 1975, Reveal & Reveal 3924. Holo- type, us! Isotypes, ariz, asu, bry, cas, COLO, GH, ISC, K, MARY, MO, NY, OKL, osc. ph. rm, rsa, smu, tex, uc, utc, wtu! Additional specimens examined: UTAH: Millard Co.: Ca 43 miles west of Delta, 30 Aug 1975, Reveal & Reveal 3999 (bry, CAS, GH, NY, OKL, RSA, US, UTc) ; ca 29.8 miles west of Delta, 30 Aug 1975, Reveal & Reveal 4000 (ariz, asu, BRY, CAS, GH, ISC, MARY, MO, NY, OKL, OSC, RM, RSA, SMU, TEX, US, UTC, WTu) . Eriogonum natum belongs to the large species group t}pified by E. brevicaule and is seemingly most closely related to E. brevicaule var. cottamii (S. Stokes) Re- veal, a narrowly restricted variant of the pinyon-juniper woodlands of north central Utah. The new species differs from var. cottamii in having longer and broader el- liptical leaves, a longer but less branched inflorescence, and smaller flowers. The new species is restricted to the white alka- line beaches of Sevier Lake and is cur- rently known for the two locations cited above. Eriogonum natum is named to honor its discoverer, Mark L. Reveal (1961- ). Status: Endemic, rare and threat- ened. Distribution: Millard Co., Utah. Eriogonum nummulare M. E. Jones, Contr. W. Bot. 11. 13. 1903. Type: Dutch Mtn., Tooele Co., Utah, 15 Jun 1900, M. E. Jones s.n. (pom). Status: Endemic, rare and seeming- ly local. Distribution: Juab, Millard, and Tooele counties, Utah (Reveal 1973a). Eriogonum ostlundii M. E. Jones, Contr. W. Bot. 11: 12. 1903. Type: Near Joseph City, Sevier Co., Utah, 13 Jun 1898, M. E. Jones s.n. (pom). Status: Endemic, restricted and lo- cal; threatened (Ripley, T). Distribution: Piute and Sevier counties, Utah (Reveal 1973a). Eriogonum panguicense (M. E. Jones) Reveal, Proc. Utah Acad. Sci. 42: 291. 1966, based on E. pauciflorum var. panguicense M. E. Jones. Contr. W. Bot. 11: 9. 1903. Type: Panguitch, Garfield Co., Utah, 24 Jun 1890, M. E. Jones s.n. (pom). Status: Endemic, restricted and lo- cal. Distribution: Garfield, Iron, Kane, Sevier, and Washington counties, Utah (Reveal 1966; 1973a). Eriogonum panguicense (M. E. Jones) Reveal var. alpestre (S. Stokes) Reveal, Proc. Utah Acad. Sci. 42: 292. 1966, Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 365 based on E. chrysocephalum ssp. al- pestre S. Stokes, Gen. Eriog. 93. 1936. Type: Cedar Breaks N.M., Iron Co., Utah, 18 Jul 1930, Goodman & Hitch- cock 1601 (cAs). Status: Endemic, rare and local; threatened (Ripley, T). Distribution: Iron Co., Utah (bry, us, UTC). Eriogonum pharnaceoides Torr. in Sitgr. var. cervinum Reveal, Great Basin Nat. 34: 245. 1974. Type: Foothills south of Pinto on the north slope of the Pine Valley Mts., 18 Aug 1973, Atwood & Higgins 5895 (us)- Status: Rare, in disjunct populations. Distribution: Millard and Washing- ton counties, Utah, and in Mohave Co., Arizona, and Lincoln Co., Nevada (Re- veal 1974). Eriogonum plumatella Dur. & Hilg., J. Acad. Nat. Sci. Philadelphia II, 3: 45. 1855. Status: Probably extirpated from Utah. Distribution: "Utah" {Palmer s.n. [gh] ) ; infrequent and widely scat- tered in northwestern Arizona, south- ern Nevada and southeastern California. Eriogonum saurinum Reveal, Great Basin Nat. 27: 197. 1968. Type: Along the Island Park road, 10 miles east of Vernal along Brush Creek on steep hillsides on the ridges, Uintah Co., Utah, 15 Aug 1966, 5,200 feet, Holmgren & Reveal 3019 (utc). Status: Edaphically restricted; threatened (Ripley, T). Distribution: Uintah Co., Utah, and adjacent northwestern Colorado (Re- veal 1973a). Eriogonum smithii Reveal, Great Basin Nat. 24: 202. 1968. Type: Between Little Flat Top and Big Flat Top, San Rafael Desert, ca 10 miles southeast of Utah Highway 24, Emery Co., Utah, 14 Aug 1966, 5,500 feet, Holmgren & Reveal 3012 (utc). Status: Endemic, edaphically re- stricted and local; threatened (Ripley, T). Distribution: Emery Co., Utah (bry, utc) . Eriogonum spathulatum A. (jray, Proc. Amer. Acad. Arts 10: 76. 1874. Type: Lower Valley of the Sevier River, Sevier Co., Utah, Jul 1874, Parry 245 (gh). Status: Endemic and scattered in isolated populations, but neither threat- ened nor endangered. Distribution: Beaver, Iron, Millard, Piute, Sanpete, and Sevier counties, Utah (Reveal 1973a). Eriogonum thompsonae S. Wats., Amer. Naturalist 7: 302. 1873. Type: Sandstone cliffs near Kanab, Kane Co.. Utah, 1872. Thompson s.n. (gh). Status: Arizona stri]) endemic, re- stricted and rare; threatened (Ripley, T). Distribution: Kane and Washington counties, Utah (bry, mary, utc), and Mohave Co., Arizona (bry. cas, us, utc). Eriogonum thompsonae S. Wats. var. albiflorum Reveal, Madrono 19: 299. 1969. Type: Ca 3 miles west of Virgin, Washington Co., Utah, 11 Aug 1966, 3,700 feet, Holmgren & Reveal 2991 (utc). Status: Endemic, edaphically re- stricted and threatened (Ripley, T). Distribution: Washington Co., Utah (bry, utc). Eriogonum tumulosum (Barneby) Reveal, Phytologia 23: 173. 1972, based on E. villiflorum A. Gray var. tumulosum Barneby, Leafl. W. Bot. 5: 153. 1949. Type: Sandstone ledges and rock- pavement on Red Plateau, southwest of Woodside, Emery Co., Utah, 13 Jun 1947, Ripley & Barneby 8678 (cas). Status: Restricted and very local in disjunct populations. Distribution: Duchesne and Emery counties, Utah, and in Moffat Co., Colorado (cs). Eriogonum umbeUatum Torr. Var. deser- eticum Reveal, var. nov. A var. umbel- lato foliis glabris et floribus stramineis differt. Type: UTAH: Utah Co.: Along the Timpooneke Road, 1 mile northwest of Utah Highway 80, near Timpooneke Campground, east of Mt. Timpanogos, 366 GREAT BASIN NATURALIST Vol. 35, No. 4 associated with Quercus, Populus and Artemisia at about 7,600 feet, 10 Jul 1974, Reveal 3702. Holotype, us! Iso- typeS, BRY, CAS, GH. MARY, MO, NY, OKL, UTC! Status: Endemic, locally common, but neither threatened nor endangered. Distribution: Juab, Salt Lake, San- pete, Tooele, Utah, and Wasatch counties, Utah. This form of Eriogonum umbellatum has been confused with var. dichro- cephalum Gandoger which has leaves pubescent at least on the lower surface. Eriogonum viridulum Reveal, Proc. Utah Acad. Sci. 42: 287. 1966. Type: Ca 8 miles east of Duchesne along U.S. Highway 40, Duchesne Co., Utah, 2 Sep 1964, Reveal 675 (utc). Status: Uinta Basin endemic, re- stricted and local; threatened (Ripley, T). Distribution: Duchesne and Uintah counties, Utah, and Moffat Co., Colo- rado (Reveal 1973a). Eriogonum zionis J. T. Howell, Leafl. W. Bot. 2: 253. 1940. Type: Zion N.P. along the Mt. Car- mel highway in the canyon of Clear Creek, Washington Co., Utah, 8 Sep 1938, Eastwood & Howell 6344 (cas). Status: Endemic, rare and local; en- dangered (Ripley, E). Distribution: Kane and Washington counties, Utah (for var. zionis), with var. coccineum J. T. Howell restricted to northern Arizona. Polygonum utahense Brenckle & Cottam, Bull. Univ. Utah, Biol. Ser. 4 (4): 3. 1940. Type: Ca 6 miles north of Escalante, Garfield Co., Utah, 17 Sep 1935, Cottam 6507 (ut). Status: Endemic; species of uncer- tain taxonomic status. Distribution: Garfield Co., Utah (bry, ut) ; known only from the type locality. POLYPODIACEAE Asplenium adiantum-nigrum L., Sp. PI. 1081. 1753. Status: Rare and local; status un- certain within Utah as not collected since the 1930s (Flowers 1944). Distribution: Washington Co., Utah; widespread in Eurasia, known only from three locations in the United States (Cronquist et al. 1972). Asplenium resi liens Kunze, Linnaea 18: 331. 1844, based on A. parvulum Mar- tens & Galeotti, Mem. Foug. Mex. 60. 1842, not Hook. Status: Local and rare; possibly threatened. Distribution: San Juan Co., Utah (Flowers 1965); widespread in North and South America. Asplenium septentrionale L., Sp. PL 1068. 1753. Status: Rare and obscure; possibly endangered. Distribution: Daggett iWieholdt 1460 A [utc]) and Grand (Maguire 1935) counties, Utah; circumboreal. Notholaena jonesii Maxon, Amer. Fern. J. 7: 108. 1917. Status: Restricted and rare; possibly threatened. Distribution: Washington Co., Utah (Maxon 1917); Arizona and southern California (Flowers 1944; Cronquist et al. 1972). Portulacaceae Calyptridium monandrum Nutt. ex Torr. & Gray, Fl. N. Amer. 1: 198. 1838. Status: Rare and restricted; possibly threatened. Distribution: Washington Co., Utah (bry); also Arizona and California, and Baja California, Mexico. Talinum valid ulum Greene, Leafl. Bot. Observ. Crit. 2: 270. 1912. Status: Rare and obscure; possibly threatened. Distribution: Emery Co., Utah (bry); Coconino Co., Arizona. Primulaceae Primula incana M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 706. 1895. Type: Beaver Co-op Ranch, at the head of the South Fork of the East Fork of the Sevier Ri^cr, Garfield Co., Utah, in cold bogs, 7,000 feet, M. E. Jones 531 2av (i'om). Status: Rare and possibly extir- pated in the type area; threatened. Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 367 Distribution: Daggett and Garfield counties, Utah (Cosgriff 1968); wide- spread in northwestern North America (Welsh 1974b). Primula maguirei L. 0. Williams, Amer. Midi. Naturalist 17: 747. 1936. Type: Damp overhanging rock ledges and cracks, 5 miles up Logan Canyon, Cache Co., Utah, 19 Apr 1932, Maguirc & Maguire 3650 (mo). Status: Endemic, rare and threat- ened (Riple}^ T) . Distribution: Cache Co., Utah (UTC). Primula specuicola Rydb., Bull. Torrey Bot. Club 40: 461. 1913. Type: Along the San Juan River near Bluff, San Juan Co., Utah, 25-29 Aug 1911, Rydberg 9882 (ny). Status: Restricted habitatwise, local and threatened (Ripley, T). Distribution: Garfield (ut). Grand, Kane, San Juan, and Wayne counties, Utah, and in northern Arizona (Cos- griff 1968; McDougall 1973). Ranunculaceae Aquilegia micrantha Eastw., Proc. Calif. Acad. Sci. II, 4: 559. 1895. Type: Near Bluff, San Juan Co., Utah, Jul 1894, Wetherill s.n. (cas). Status: Restricted habitatwise, local but not threatened nor endangered. Distribution: Emery, Garfield, Grand, Kane, and San Juan counties, Utah (bry) ; also in Arizona and Colo- rado. Ranunculus acriformis A. Gray var. aesti- valis L. Benson, Amer. Midi. Natural- ist 40: 43. 1948. Type: Meadow at springs just east of U.S. Highway 89 and 300 yards west of the Sevier River, 8.3 miles north of the principal intersection in Panguitch and about 1.5 miles south of the inter- section with Utah Highway 20 leading to Parowan, Garfield Co., Utah, 29 Aug 1948, 6,400 feet, Benson 13420 (pom). Status: Endemic and presumed to be extinct (Ripley, PoEx). Distribution: Garfield Co., Utah (bry) ; known only from the type lo- cality. ROSACEAE (Viamacrhodos erecta (L). Bunge in Ledcb., Fl. Altaica 1: 430. 1829, based on Sihbaldia erecta L., Sp. PI. 1: 284. 1753. Status: Rare and local, altitudinally restricted. Distribution: Piute and Wayne counties, Utah (bry) ; also from Colo- rado, North Dakota and Michigan, north to Yukon and Alaska; Asia (Welsh 1974b). Crataegus chrysocarpa Ashe, North Caro- lina Agric. Exp. Sta. Techn. Publ. 175: 110. 1900, based on C. rotundifolia Moench, Verz. Ausl. Baume Stand Weiss 29. 1785, not Lam. Status: Rare and threatened. Distribution: Cache Co., Utah (Maguire 1937); widespread elsewhere. Crataegus succulenta Schrader ex Link, Handbuch2: 78. 1831. Status: Local and rare; threatened. Distribution: Utah Co., Utah (bry, UTC; Barnes 1943); widespread to the east of Utah (Little 1953). Ivesia sabulosa (M. E. Jones) Keck, Lloydia 1: 124. 1938, based on Ponten- tilla sabulosa M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 680. 1895. Type: Head of the Sevier River, prob- ably in Garfield Co., Utah, 11 Sep 1894, 8,000 feet, M. E. Jones 6032 (pom). Status: Rare and local, possibly threatened. Distribution: Garfield and Wash- ington counties, Utah (bry), and Nye Co., Nevada (Keck 1938b). Ivesia utahensis S. Wats., Proc. Amer. Acad. Arts 17: 371. 1882. Type: On the summit of Bald Moun- tain, in Wasatch Range, above Alta, Salt Lake Co., Utah, Aug 1879, over 12,000 feet, M. E. Jones 1231 (gh). Status: Endemic, rare and local. Distribution: Salt Lake, Summit, and Utah counties, Utah (Keck 1938b). Rubus neomexicanus A. Gray, Smithso- nian Contr. Knowl. 5: 55. 1853. Status: Local, rare and threatened. Distribution: San Juan Co., Utah (bry) ; New Mexico, Arizona, and northern Mexico. 368 GREAT BASIN NATURALIST Vol. 35, No. 4 Most of the known range of this species has been destroyed by Lake Powell. RUBIACEAE Galium multiflorum Kellogg var. wat- sonii A. Gray, Syn. Fl. N. Amer. 1 : 40. 1884. Type: Wasatch Mts., Utah, 1869, Watson 484 (gh) . Status: Endemic, neither threatened nor endangered. Distribution: Box Elder, Cache, Davis, Tooele, and Utah counties, Utah (Dempster & Ehrendorfer 1965). Galium scabruisculum. (Ehrendorfer) Dempster & Ehrendorfer, Brittonia 17: 312. 1965, based on G. hypotrichium ssp. scabjuisculum Ehrendorfer, Contr. Dudley Herb. 5: 13. 1956. Type: Calf Springs Wash, San Rafael Swell, Emery Co., Utah, Maguire 18457 (gh). Status: Endemic, neither threatened nor endangered. Distribution: Carbon and Emery counties, Utah. RUTACEAE Ptelea trifoliata L. ssp. pallida (Greene) V. L. Bailey, Brittonia 14: 23. 1962, based on P. pallida Greene, Contr. U. S. Natl. Herb. 10: 70. 1906. Status: Rare and possibly extirpated. Distribution: Garfield and Kane counties, Utah; this subspecies also in Arizona and Colorado (Bailey 1962). SCROPHULARIACEAE Castilleja aquariensis N. H. Holmgren, Bull. Torrey Bot. Club 100: 87. 1973. Type: Aquaris Plateau, 22 miles northwest-north of Escalante on the road to Bicknell, 0.5 mile north of Clay- ton Guard Station turnoff, Garfield Co., Utah, 11 Aug 1970, 9,600 feet, Holm- gren & Holmgren 4726 (ny). Status: Endemic, rare and local; en- dangered (Ripley, E). Distribution: Garfield Co., Utah; known only from the type area. Castilleja leonardii Rydb., Bull. 7V)rrey Bot. Club 34: 36. 1907. Type: Head of American Fork Can- yon, Utah Co., Utah, 1885, Leonard 151 (ny). Status: Endemic, locally common but neither threatened nor endangered. Distribution: Cache, Daggett, Davis, Duchesne, Salt Lake, Sanpete, Sununit, Tooele, Utah, and Wasatch counties, Utah (bry, ut) . Castilleja parvula Rydb., Bull. Torrey Bot. Club 34: 40. 1907. Type: Mountains north of Bullion Creek near Marysvale, Piute Co., Utah, 1905, Rydberg & Carlton 7158 (ny). Status: Endemic, rare and local; threatened (Ripley, T). Distribution: Piute Co., Utah (ny, utc). Castilleja revealii N. H. Holmgren, Bull. Torrey Bot. Club 100: 87. 1973. Type: Bryce Canyon N.P., along the road to Bryce Point, 0.5 mile from In- spiration Point turnoff, Garfield Co., Utah, 24 Jun 1965, 8,000 feet, Holm- gren & Reveal 2017 (ny). Status: Endemic, rare and local; en- dangered (Ripley, E). Distribution: Garfield Co., Utah; known only from the type locality. Mimulus eastwoodiae Rydb., Bull. Torrey Bot. Club 40: 483. 1913. Type: In crevices of perpendicular or overhanging rocks along the San Juan River near Bluff, 25-29 Aug 1911, Rydberg 9883 (ny). Status: Edaphically restricted, local and disjunct; not threatened nor endan- gered. Distribution: Grand, Kane, and San Juan counties, Utah (bry), and adja- cent northern Arizona (Kearney & Peebles 1951). Penstemon abietinus Pennell, Contr. U.S. Natl. Herb. 20: 276. 1920. Type: Ireland Ranch, head of Salina Canyon, Sevier Co., Utah, 15 Jun 1894, 2,400 m, M. E. Jones 5440 (us). Status: Endemic, rare and local; threatened (Ripley, T). Distribution: Iron (ut), Sevier, and Utah counties, Utah (bry; Keck 1937a). Penstemon acaulis L. O. Williams, Ann. Missouri Bot. Card. 21: 345. 1934. Status: Restricted, rare and threat- ened (Ripley, T). Dec. 1975 WELSH, F,T AL.: ENDANGERED UTAH PLANTS 369 Distribution: Daggett Co., Utah (ny, us, utc) and adjacent vSweetwater Co., Wyoming (Keck 1937a). Penstemon atwoodii Welsh, Great Basin Nat. 35: 378. 1976. Type: South end of Horse Mtn., ca 10 miles south-southeast of Canaan Peak, Kane Co., Utah, 14 Jun 1975, S. L. & S. L. Welsh 12820 (bry). Status: Endemic, local and threat- ened. Distribution: Garfield and Kane counties, Utah (bry, ny). Penstemon bracteatus Keck, Leafl. W. Bot. 1: 82. 1934. Type: Red Canyon, Garfield Co., Utah, 20 Jun 1933, Eastwood & Howell 783 (cAs). Status: Endemic, restricted and rare; possibly threatened. Distribution: Garfield Co., Utah (bry). Penstemon caespitosus Nutt. var. suffruti- cosus A. Gray, Syn. Fl. N. Amer. 2: 270. 1878. Type: Near Beaver, Beaver Co., Utah, 1877, Palmer s.n. (gh). Status: Endemic, restricted and lo- cal; threatened (Ripley, T). Distribution: Beaver, Garfield and Piute counties, Utah (Keck 1937a). Penstemon compactus (Keck) Crosswhite, Amer. Midi. Naturahst 77: 6. 1967, based on P. cyananthus ssp. compactus Keck, Amer. Midi. Naturalist 23: 615. 1940. Type: Stony slopes of Mt. Naomi, Cache Co., Utah, 18 Aug 1938, 2,900 'meters, Maguire 16148 (utc). Status: Endemic, restricted and threatened (Ripley, T). Distribution: Cache Co., Utah (bry, UTC, w^sco). Penstemon concinnus Keck, Amer. Midi. Naturalist 23: 608. 1940. Type: Tunnel Springs, northwest corner of Desert Range Experiment Sta- tion boundary, about 10 miles east of Garrison, Millard Co., Utah, 28 Jun 1933, 1,675 meters, Cottam 5655 (ds). Status: Endemic, restricted and rare; endangered (Ripley, E). Distribution: Beaver and Millard counties, Utah (bry, ny, us, utc). Penstemon garrettii Pennell, (^ontr. U.S. Natl. Herb. 20: 353. 1920. Type: Crevices in travertine rock, "Hot Pots," near Midwav, Wasatch Co., Utah, 6 Jul 1905, Carlrton & Garrett 6697 (ny). St.'Vtus: Endemic, restricted and pos- sibly extinct. Distribution: Duchesne (utj and Wasatch counties, Utah. Penstemon grahamii Keck in Graham, Ann. Carnegie Mus. 26: 331. 1937. Type: Talus slope on the west side of Green River, south of the mouth of Sand Wasli, Uintah Co., Utah, 27 May 1933, Graham 7883 (cm). Status: Endemic, rare, restricted and endangered (Ripley, E) . Distribution: Uintah Co., Utah (bry, utc; Keck 1938a). Penstemon humilis Nutt. ex Gray var. brevifolius A. Gray, Syn. Fl. N. Amer. 2: 267. 1878. Type: Cottonwood Canyon, Wasatch Mts., Salt Lake Co., Utah, 1869, 9,000- 10,000 feet, Watson 781 (gh). Status: Endemic, local and possibly threatened. Distribution: Cache, Juab, Salt Lake, Utah, and Weber counties, Utah (bry, ut). Penstemon humilis Nutt. ex Gray var. obtusifolius (Pennell) Reveal, stat. no v., based on P. obtusifolius Pennell, Contr. U.S. Natl. Herb. 20: 370. 1920. Type: Springdale, Washington Co., Utah, 16 May 1894, 1,600 meters, M. E. Jones 5249am (pom). Status: Endemic, rare and restricted; possibly threatened. Distribution: Beaver and Washing- ton counties, Utah (Keck 1945). Penstemon jonesii Pennell, Contr. U.S. Natl. Herb. 20: 338. 1920. Type: Springdale, Washington Co., Utah, 17 May 1894, M. E. Jones 5250, in part (us). Status: Endemic, rare and obscure; taxonomic status questionable. Distribution: 'Washington Co., Utah; known only from the tj^e lo- cality. Penstemon laevis Pennell, Contr. U.S. Natl. Herb. 20: 347. 1920. 370 GREAT BASIN NATURALIST Vol. 35, No. 4 Type: Red sand at Springdale, Wash- ington Co., Utah, 17 May 1894, 1,200 meters, M. E. Jones 5250^ in part (us). Status: Endemic, restricted and lo- cal. Distribution: Kane, Garfield, and Washington counties, Utah (bry, utc) . Penstemon leiophyllus Pennell, Contr. U.S. Natl. Herb: 20: 346. 1920. Type: Mammoth Creek, Garfield Co., Utah, 10 Sep 1894, 2,400 meters, M. E. Jones 6026b (us). Status: Endemic, restricted and lo- cal; threatened (Ripley, T). Distribution: Garfield, Iron, Kane, and Washington counties, Utah (bry). Penstemon lentus Pennell var. albiflorus (Keck) Reveal, stat. nov., based on P. lentus ssp. albiflorus Keck, Amer. Midi. Naturalist 23: 616. 1940. Type: Abajo Mts., ca 8 miles west of Blanding, near the "Bear's Ears," San Juan Co., Utah, 9 Jun 1938, 2,400 meters, C. L. Porter 1801 (rm). Status: Endemic, locally common and not threatened. Distribution: San Juan Co., Utah (bry). Penstemon leonardii Rydb., Bull. Torrev Bot. Club 40: 483. 1913. Type: Diehl's Grove, Wasatch Mts., possibly Utah Co., Utah, 1 Aug 1884, Leonard s.n. (ny). Status: Endemic, locally common and not threatened nor endangered. Distribution: Cache, Davis, Rich, Salt Lake, Summit, Uintah, Utah, Wa- satch, Washington, and Weber counties, Utah (bry. ut). Penstemon longiflorus (Pennell) S. L. Clark, Great Basin Nat. 35: 434. 1976, based on P. cyananthus ssp. longiflorus Pennell, Contr. U.S. Natl. Herb. 20: 353. 1920. Type: Beaver, Beaver Co., Utah, Palmer 376 (ny). Status: Endemic, restricted but lo- cally common. Distribution: Beaver, Millard, and Piute counties, Utah (bry). Penstemon nanus Keck, Amer. Midi. Nat- uralist 23: 607. 1940. Type: Desert Range Experiment Sta- tion, about 10 miles east of Garrison, Millard Co., Utah, 13 May 1939, 1,675 meters, Plummer 7313 (ds). Status: Endemic, restricted and lo- cal; threatened (Ripley, E). Distribution: Beaver and Millard counties, Utah (bry, utc). Penstemon parvus Pennell, Contr. U.S. Natl. Herb. 20: 345. 1920. Type: The Button, Aquarius Plateau, Wayne Co.. Utah, 11 Aug 1875, Ward 546 (us). Status: Endemic, restricted and rare; threatened (Ripley, T). Distribution: Garfield and Wayne counties, Utah, (bry; Pennell 1920). Penstemon sepalulus Rydb., Bull. Torrey Bot. Club 36: 690. 1909. Type: Canyons of the Wasatch Mts., Provo Canyon, Utah Co., Utah, Jul 1869, Watson 786 (ny). Status: Endemic, locally abundant. Distribution: Utah Co., Utah (bry; Keck 1932). Penstemon tidestromii Pennell, Contr. U.S. Natl. Herb. 20: 379. 1920. Type: "XL" Canyon, San Pitch Mts., Sanpete Co., Utah, 24 Jun 1908, 1,650 meters, Tidestrom 1296 (us). Status: Endemic, local and obscure; taxonomic status questionable. Distribution: Sanpete Co., Utah; known only from the type locality. Penstemon uintahensis Pennell, Contr. U.S. Natl. Herb. 20: 350. 1920. Type: Dyer Mine, Uinta Mts., Uin- tah Co., Utah, 50 Jun 1902, Goodding 1221 (ny). St.'vtus: Endemic, rare and restric- ted; threatened (Ripley, T). Distribution: Daggett. Duchesne, and Uintah counties, Utah (bry). Penstemon wardii A. Gra^'. Proc. Amer. Acad. Arts 12: 82. 1876. Type: Near Glenwood, Sevier Co., Utah, 4 Jun 1875, Ward 162 (us). Status: Endemic, restricted and lo- cal; threatened (Ripley, T). Distribution: Sanpete and Sevier counties, Utah (bry. utc). Synthris laciniata (A. Gray) Rydb. ssp. ibapahensis Pennell, Proc. Acad. Nat. Sci. Philadelphia 85: -92. 1933. Type: Wot gravelly slopes near ra- vine of snow on Mount Ibapah, Juab Dec. 1975 WELSH, ET AL.: ENDANGERED UTAH PLANTS 371 Co., Utah, 5 Jul 1932, 9,500-10,000 feet, Stanton 1000 (ph). Status: Endemic, restricted iind ob- scure; taxonomic status questionable. Distribution: Juab Co., Utah; known only from the type locality. SELAGINELL.A.CEAE Selaginella utahensis Flowers, Amer. Fern J. 39: 83. 1949. Type: South of St. George, Washing- ton Co., in a wash bottom, 5 Apr 1931, Cottam 5644 (ut). Status: Rare and local; possibly threatened. Distribution: Washington Co., Utah, and Spring Mts., Clark Co., Ne- vada (Cronquist et al. 1972). Verbenaceae Aloysia wrightii (A. Gray) A. A. Heller, Muhlenbergia 1: 147. 1906, based on Lippa wrightii A. Gray, Amer. J. Sci. II, 16: 98. 1853. Status: Local and rare; possibly en- dangered. Distribution: Washington Co., Utah (Higgins 615 [bry]; Higgins 1972b); from Texas to California and in north- ern Mexico. Violaceae Viola purpurea Kellogg var. charlestonen- sis (Baker & Clausen) Welsh & Reveal, Stat. & comb, nov., based on V . charles- tonensis Baker & Clausen in Clokey, Madrono 8: 58. 1945. Status: Restricted, local and rare; threatened (Ripley, T). Distribution: Washington Co., Utah, and Clark Co., Nevada (Clokey 1945; Eastmond 1969). Literature Cited Al-Shehbaz, I. A. 1973. The biosystematics of the genus Thely podium (Cruciferae). Contr. 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Naturalist 17: 741-748. . 1937. A monograph of the genus Mer- tensia in North America. Ann. Missouri Bot. Gard. 24: 17-159. Woodson. R. E., Jr. 1954. North American spe- cies of Asclepias. Ann. Missouri Bot. Gard. 41: 1-195. YuNCKER, T. G. I960. Two new species of Cuscula from North America. Brittonia 12: 38-40. . 1965. Cuscuta. N. Amer. FI. II. 4: 1-51. UTAH PLANT NOVELTIES IN CYMOPTERUS AND PENSTEMON Stanley L. Welshi Abstract. — Cymopterus higginsii nnd Penstetnon atwoodii are named and described from ma- terials collected in the Kaiparowits Plateau vicinity of eastern Kane County. Utah. Habitat, distri- bution, and probable affinities are outlined. Examination of specimens obtained from the Kaiparowits Plateau region of eastern Garfield and Kane counties in southern Utah has revealed the existence of two previously undescribed entities, one each in Cymopterus (Apiaceae) and Penstemon (Scrophulariaceae). Existence of these taxa is not surprising when one considers the historic remoteness of much of that great region. The discovery and the extent of the range of each taxon must be credited to the extensive field work- allowed under the baseline studies of the Navajo-Kaiparowits enA'ironmental pro- ject directed by personnel from Brigham Young University (Welsh, Murdock, and Wood 1975). The Cymopterus species is known from saline soils of the Tropic Shale formation and associated pedimental gravels on fans and bajadas below the Straight Cliffs for- mation in that portion of Kane County from the Paria River eastward to the Last Chance Creek vicinity. Apparent rela- tionships of C. higginsii seem to lie with C. fendleri from which it differs inter alia in the rose to purple flowers with evident pedicels and wider wings on the fruit. The pseudoscape is poorly developed. The corollas in the Penstemon species are glandular hairy externally, and the taxon seems to belong with those species treated by Pennell (1920) as Section Crista ti and by Keck (1938) illegitimately as Section Aurator. The nearest ally ap- pears to be P. jamesii from which P. at- woodii differs as noted in the diagnosis. P. atwoodii is known only from middle elevations of the Kaiparowits region, where it grows on the Cretaceous forma- tions in juniper-piny on woodland. Both species, the Cymopterus and the Penstemon, are plants of very restricted range. They are in areas which are now subject to commercial exploitation, and both should be considered as threatened species. Cymopterus higginsii Welsh sp. nov. Plantae acaulescentes non caespitosae pseudoscapis non vel non nisi evolutis in- fermis, pubescentes parse pili complanti: folia ovata vel subelliptica in circum- scriptem, laminis 1.8-7.7 cm longis 1.5- 6.0 cm latis hi- vel tripinnatis vi- ridibus foliolosis longior quam latis pinnatis ad bipinnata, lobi obtusi ad ro- tundatos vel acutos raro, petiolis 1.8-14 cm longis; pedunculi folia longior ad ex- tremum 2-12 cm longi, purpura scentes; involucrum vaginans margine scariosa; involucellum bracteolarum brevior quam floras, lobis aliquot dentatis acutis vel acuminatis; umbellae compactae, radiis 3-5, 1-10 mm longis, umbellula centrali sessili; pedicelli 1-6 mm longi; flores rosei ad purpurascens; fructus ovalis ad ellipticum 7-10 mm longus 5-8 mm latus, alis corpus subaequalis incrassatis spon- giosis. C. fendleri affinis sed floribus roseis ad purpurascens pedicellis evidentibus et alis corpus subaequalis. Habitat and Distribution. — Tropic shale and pedimental covering derived from Straight Cliffs and other formations, on saline soils, from East Clark Bench eastward to Last Chance Canyon, at least 30 miles east of Glen Canyon City, east- ern Kane County, Utah. Type: Utah: Kane Co., Shadscale dom- inated bajada, on gravell}' pedimental fan, east of None Butte, ca 17 miles east of Glen Canyon Citv, S. L. Welsh 12740, 31 May 1975 (Holotype BRY; Isotypes to be distributed). Paratypes: Utah: Kane Co., Site 9, Navajo-Kaiparowits Project, base of Smoky Mt., 2 miles from Ahl- strom Point junction, Atriplex-Kochia- 'Department of Botan.v ;ind Range Science, Brigham Young Univeisily. 377 378 GREAT BASIN NATURALIST Vol. 35, No. 4 Artemisia community, N. D. Atwood 3439, March 1972 (BRY) ; ca 2 miles north of Church Wells, on bench between Coyote Creek and Wahweap Creek, N. D. Atwood et al. 3493, 23 March 1972 (BRY) ; ca 30 miles east of Glen Canyon City, on Tropic Shale formation, N. D. Atwood 4549, 23 April 1973 (BRY). The species is named to honor Larry Charles Higgins, student of Boragmaceae, especially of Cryptantha. and specialist in western botany generally. Penstemon atwoodii Welsh sp. nov. Herbae perennes 1.4-5.3 cm altae; caules pauci vel multi e caudicibus ramifi- cantibus glabri infra medium ])ilis patulis glanduliferis super; folia glabra, basalia oblanceolata ad spathulata vel ovata ad elliptica raro Integra 2.8-9.0 cm longa (0.2) 0.4-0.8 (1.4) cm lata, caulina lanci- linearia ad oblonga vel spathulata, ± auriculata super Integra vel serrata re- mota raro, 3.0-7.0 cm longa 0.3-0.8 (1.4) cm lata; inflorescentia verticillastorum distinctarum plurium; bracteae foliaceae; calyces 6.5-8.5 mm longi lobis lanceolatis herbaceis purpureis pilis glanduliferis; corollae pilis glanduliferis externis cyanae vel cyano-caesiae 13-16 mm longae ex- pansae distales 5-6 mm latae glabrae intus praeter ad orficium labium inferum; an- therarum thecae glabrae divaricatae vel explanatae; staminodium barbatum pilis luteo lineare; capsulae glabrae. P. jamesii sensu lato affinis sed floribus parvioribus et glabris intus praeter ad orficium labium inferum. Habitat and Distribution. — Kaipar- owits, Wahweap, and Straight Cliffs for- mation at 6,200 to 8,000 feet elevation in pinyon-juniper woodland on the Kai- parowits Plateau of eastern Garfield and Kane counties, Utah. Type: Utah: Kane Co., Gray sand of Kaiparowits formation, pinyon-juniper community, south end of Horse Mountain, ca 10 miles south-southeast of Canaan Peak, S. L. & S. L. Welsh 12820, 14 June 1975 (Holotype; BRY; Isotypes to be distributed). Paratypes: Utah: Garfield Co., Death Ridge ca 16 miles southwest of Escalante, N. D. Atwood 5177, 30 May 1973; do S. L. Welsh & J. R. Murdock 12866, 28 June 1975. Kane Co., 4 miles southeast of sum- mit of Collets Wash, Kaiparowits Plateau, N. D. Atwood s. n., 19 June 1969; Pin- yon-juniper woods on basal Wahweap formation, ca 6 miles north of junction of Escalante road with head of Last Chance Creek, Kaiparowits Plateau, S. L. Welsh & J. R. Murdock 12793, 4 June 1975; do S. L. Welsh & J. R. Murdock 12973a, 4 June 1975. This species is named to honor Nephi Duane Atwood, student of Hydrophyl- laceae, field botanist extraordinary, and first to recognize the unique nature of this taxon. References Keck. D. D. 1938. Studies in Penstemon VI. The section Aurator. Bull. Torrev Bot. Club 65: 233-255. Mathi.\s, M. E. .\nd L. Constance. 1945. Cymopterus Raf. N. Amer. Fl. 28B: 170-183. Pennell, F. W. 1920. Scrophulariaceac of the central Rockv Mountain states. Contr. U. S. Nat. Herb. 20: 313-381. Welsh, S. L., J. R. Murdock, and B. W. Wood. 1975. Navajo-Kaiparowits Environmental Baseline Studies. Unpublished mss, 800 p. THE ZYGOPTERA (ODONATA) OF UTAH WITH NOTES ON THEIR BIOLOGY^ A. V. Provonsha^ Abstract. — Detailed distribution maps of Utali are provided for each of the 33 state species of Zygoptera. Notes on the general range, habitat preference, reproductive behavior, emergence data, and general biology of each species are also included. As noted by Kormondy (1957), pub- lished data on the geographical distribu- tion of western Odonata is scarce. Ken- nedy (1915) gave a partial list of the Odonata of Washington and Oregon, and in 1917 he published records from central California and Nevada. The Washington list has recently been updated by Paulson (1970) and a complete list and keys to the California Odonata is given by Smith and Pritchard (1956). Bick and Hornuff (1972) published man}^ new Odonata records for northwestern Wyo- ming. Although Larsen (1952) and Mus- ser (1962) added considerably to our knowledge of Utah Anisoptera, only one paper (Brown 1934) dealing exclusively with the Odonata of Utah included a treatment of the Zygoptera. Brown's re- port consisted of an annotated checklist including some 25 currently valid zygop- teran species. During the summers of 1970 and 1971 the state of Utah was extensively collected in an effort to gain an increased under- standing of the distribution and species composition of the damselfly fauna of that area. Some 152 localities throughout the state were sampled, and the collections at the University of Utah, Utah State Uni- versity, Brigham Young University, and Dixie College were examined. As a result, the list of Zygoptera known to occur in Utah has been expanded to 33. In ad- dition, field work and rearing has added considerably to our knowledge of the bi- ologv of many species (also see Provonsha and McCafferty 1973). No attempt has been made to cite in de- tail all collection records. Although county records may be adequate for some states where counties are numerous and rela- tively small, they are meaningless for Utah, where several counties contain over Fig. 1. County map of Utah. 5,000 square miles and in many cases more than one distinct biotic region. In- stead, distributions based on all collecting localities known to the author are plotted for each species.^ For those instances where counties are mentioned in the text, the reader may refer to Fig. 1 for their specific location. Where possible, notes on general distribution, habitat prefer- ences, emergence data, and reproductive behavior are included in the text. For keys to the zygopteran species known to occur in Utah, the author re- commends the following publications: Johnson (1972), Smith and Pritchard (1956), and Walker (1953). ^Complete collecting data may be obtained from the author upon request. ^Published with the approval of the Director of the Purdue University Agricultural Experimeni Series No. 6034. ^Department of Entomology, Purdue University, West Lafayette, Indiana 47907. Station as Journal 379 380 GREAT BASIN NATURALIST Vol. 35, No. 4 Calopteryx aequabilis Say, 1839 Fig. 2 This species is known to occur through- out most of Canada east of Saskatchewan and the north central and northeastern United States, with isolated pockets in Colorado, California, and the Columbia River drainage system. The major popula- tion of C. aequibilis in Utah is found at the Raft River, Box Elder Co., which is the only major river in Utah flowing north as part of the Columbia River drain- age system. One other small isolated pop- ulation occurs at Far West, Weber Co. A single male was collected at Goshen, Utah Co., on 19 May 1969, by P. V. Winger. Subsequent collections at that locality have failed to provide additional specimens, and it is doubtful that a popu- lation is established there. The species is restricted to streams where the nymphs cling to debris and submerged roots along the banks. The above record from Goshen is the earliest Utah record I have; species have been taken at the Raft River through mid-September. This species does not ovi- posit in tandem; however, the male re- mains in close proximity to the female and actively wards off other intruding males. Martin (1939) and Walker (1953) observed females descending as much as one foot below the water to deposit eggs. In Jidy 1971 I observed several females ovipositing at the Raft River. On this oc- casion none submerged but completed oviposition just below the water line in algal mats in shallow water close to the bank. It must be noted, however, that the water level was lower than usual and there was little suitable vegetation in the deeper portions of the river. Hetaerina amcricana (Fabricius, 1788) Fig. 2 With the exception of the far eastern and northwestern states and Florida, H. americana has a general distribution throughout the United States and extends southward through Mexico into Guate- mala. It is found throughout Utah in rivers and streams below 5,800 feet ele- \ation which have a moderate flow and sufficient marginal vegetation for ovipo- sition and nymphal development. The earliest emergence date I have for the state is 29 May. The nymphs emerge over most of the summer and can be found in the adult stage to the end of September. The female oviposits singularly and com- pletely submerge to deposit their eggs, while the male, which perches nearby, faces the submerged female and actively fends off intruding males (Johnson 1961 and Bick and Sulzback 1966). Hetaerina vulnerata Hagen, 1853 Fig. 2 H. vulnerata is restricted to the south- western United States and Mexico, enter- ing only the southwest corner of Utah in Washington County, which is part of the Mohave Desert Lower Sonoran. Like H. americana, the nymphs are found on roots, vegetation, and debris in streams with a moderate current. Although these two species were found at nearby streams, they were never taken at the same lo- calities (Provonsha and McCaffertv 1973). FIG, 2 Calopteryx aequibile Hetaerina americana Hetaerina vulnerata ]> Emergence begins around the first of June and is probably completed by mid-July. I am not aware of any reproductive stud- ies conducted on this species, and I have never observed any in copulation. How- ever, it is assumed that .the method of oviposition is similar to that of H. amer- icana. Dec. 1975 PROVONSHA: ZYGOPTERA OF UTAH 381 Archelestes grandis (Rambur, 1842) Fig. 3 Although this species ranges over much of the United States, in Utah it is restric- ted to the southern deserts. The n5nnphs are active swimmers and can be found in ponds and slow desert streams. All Utah collecting sites were below 4,600 feet. A. grandis is the largest of all Nearc- tic damselflies, the females having a wing span of approximately 40 mm. Like most other lestids, it is a late emerger. The earliest Utah record I have is 9 July. Bick and Bick (1970) reported that in Okla- homa emergence is much earlier and that oviposition is common by mid-June. In that area the eggs reportedly hatch ap- proximately 16 days after oviposition, and the nymphs overwinter in a fairlj' late stage of development. However, there is some evidence that in the western des- erts, where emergence tends to be later and where there is often a winter drought, eclosion may be delayed until the fol- lowing spring. Oviposition takes place in tandem, and the eggs are deposited as high as 13 m above water in woody plants overhanging the water. This unique re- productive behavior has enabled this spe- cies to colonize certain habitats, such as desert streams, where marginal vegeta- tion may be plentiful but vegetation within the water may be wanting. T^m Archelestes grandis Lestes congener ■m- Lestes congener Hagen, 1861 Fig. 3 This species is found over much of the United States and is by far the most com- mon and wide-ranging Lestes in Utah. All collecting sites were at permanent and semipermanent ponds or "pondlike" ex- pansions of slow streams at altitudes rang- ing from 4,200 to 7,000 feet. The nymphs are free swimmers and seem to prefer shallow ponds with some areas free of vegetation. Emergence begins in early July and continues through late August. As in most other Lestes species, the eggs are usually deposited some distance above water. However, on one occasion during flood conditions I observed two females submerge their abdomens almost to the base to deposit eggs below the water line. Lestes disjunctus disjunctus Selys, 1862 Fig. 4 Although primarily found in Canada, Alaska, and the northern regions of the United States, this species does follow the mountains south through Utah and Colo- rado into Arizona. In Utah it was found mainly at permanent and semipermanent, richly vegetated ponds in mountainous areas between 5,000 and 7,000 feet. Emer- gence begins around mid-July and con- tinues through mid- August. Although ovi- FiG. ^ Lestes disjunctus ^ 382 GREAT BASIN NATURALIST Vol. 35, No. 4 position usually takes place well above water, on two separate occasions in 1970 I observed paired adults completely sub- merge to ovipost. These observations were made during flood conditions when more than the usual amount of vertical stems were below water. These observa- tions coincide for the most part with ob- senations by Bick and Bick (1961) for L. d. nustralis Walker. Lestes dry as Kirby, 1890 Fig. 5 This species is Holarctic, occurring not onh' in North America but Europe and Asia as well. It is found most commonly at permanent and semipermanent ponds and less frequently in marshy areas. It was the only Lestes species taken above 8,000 feet in Utah. Although the locality records are scattered, L. dryas tended to be the dominant species at these localities. Emergence begins about the last week in June and continues through mid-August. Lestes unquiculatus Hagen, 1861 Fig. 6 The range of L. unquiculatus is trans- continental in the northern United States and southern Canada. It was found at only a few localities in Utah and never in any 7W FIG, 6 Lestes unquiculatus l> large numbers. Specimens were collected primarily at temporary and semiperm- anent marshy areas between 5,000 and 6,350 feet. Emergence data for the state is insufficient. However, Walker (1953) reported that in Canada they fly mainly in .Inly and August. Argia alberta Kennedy, 1918 Fig. 7 A. alberta is restricted to the western United States. Although it occurs in most regions of Utah at altitudes between 4,000 and 6,500 feet, it was never taken in any large numbers. The nymph has not been described, and none were taken during this study. With the exception of the San Juan River in San Juan Co., all adults were taken in association with small, slow flowing streams or marshy springs. The earliest collecting record I have for the state is 4 June and the flight period extends through the end of August. As with most Argia species, oviposition usu- ally takes place in tandem. Argia emma Kennedy, 1915 Fig. 8 This species is found in the western United States and British Columbia. In Utah it is found in the mountainous areas Dec. 1975 PROVONSHA: ZYGOPTERA OF UTAH 383 I^ FIG, 7 • Argia alberta * Argia sedula 'm- FIG, 8 Argia emma -^m ^ in the northern two-thirds of the state in rivers and streams with gentle to moderate currents at altitudes between 4,200 and 5,800 feet. The nymphs may be found on stones but were most frequently taken on roots and debris near the bank. Emergence begins around the first of June and con- tinues through mid-July and oviposition usually takes place in tandem. Argia fumipennis violacea (Hagen, 1861) This species is known from Guatemala north through the southwestern United States, most of the central and eastern United States, and eastern Canada. How- ever, I have seen only one specimen from Utah (Univ. of Mich. Coll.). The label read: "Utah E. M. Legard #55." Although specific local information was lacking, it is speculated, based on overall distribu- tion, that this specimen was collected in the Lower Sonoran region of Washington Co. It is reported that A. f. violacea pre- fers small lakes and shallow streams with moderate currents and exposed rocks (Walker 1953). Argia lugens (Selys, 1854) Fig. 9 This species is found only in Mexico and the southwestern United States. The only Utah records I have are from Zion National Park and Leeds Canyon (both in Washington Co.) where the nymphs in- habit permanent desert streams. I have no emergence data for the state. All spec- imens examined were collected during the month of July. .■ *' N ,IV;. FIG, 9 , . 7 \, ;^- • Argia lugens <3 J^-;-- * Argia moesta ^v\y^-v L,.V-A^^r.;,.;-;-.^?^ .A f --J:','-fl^4^__:__S^----- '• •"' ;;----r- r'-y 384 GREAT BASIN NATURALIST Vol. 35, No. 4 Argia moesta (Hagen, 1861) Fig. 9 Although common throughout North America wdth the exception of the north- western states and western Canada, this species has been taken in Utah only from the Colorado River, where the nymphs can be found on rubble and in debris near the bank. Emergence begins in late May and continues through most of June. Walker (1953) reports that females oviposit either in tandem or unattended by the male and that they may completely submerge to deposit their eggs. Unlike most western species of Argia, which prefer to light on bare ground, all individuals of A. moesta observed in Utah rested on willows along the river's edge. Argia nahauna Calvert, 1901 Fig. 8 This species is known only from the southwestern United States, entering Utah only in Washington Co. It is found at small streams with moderate currents and occasionally in permanent desert springs. At most localities it was found in association with A. sedula. The earliest Utah record I have is 5 June and the latest is 9 September. Argia sedula (Hagen, 1861) Fig. 7 The reported range of A. sedula in- cludes most of the southern portion of the United States, northern Mexico, the mid- western states, and southern Ontario. In Utah it was found only in Washington Co. at altitudes between 2,800 and 4,400 feet. The nymphs prefer small streams with gentle current and a rich growth of vegetation. Emergence begins in mid -May and continues through mid- June. Argia vivida Hagen, 1865 Fig. 10 A. vivida is restricted to the western United States and southwestern Canada. It is fairly common in Utah and can be found at most rivers and streams with moderate currents. During this study they were taken at altitudes between 2,800 and 6,500 feet. Although they can be found in debris along the stream banks, the nymphs prefer stones and rubble well FIG. 10 • Argia vivida ??-iu:. within the main current of the stream. Emergence begins around the first of June and continues through mid-July. Ovipo- sition takes place in tandem, and eggs are laid just below the water surface in a wide variety of aquatic plants. Like most other Argia species, the males will fre- quently balance in a vertical position during oviposition when the immediate environment does not provide a good ob- ject for them to grasp. Amphiagrion abbreviatum (Selys, 1876) Fig. 11 This species is known from the western United States and southwestern Canada. I found this species at a great many lo- calities throughout the state, but never in any large numbers. The habitats were varied, ranging from moderately flowing streams with gravel bottoms, to heavily vegetated ponds and springs, to large lakes. However, as Whitehouse (1941) foiuid, they tend to prefer shallow, sunlit marshy areas with little or no current. They were taken at altitudes between 4,200 and 7,500 feet. Emergence begins in early May and the flight period ex- tends through the end of August. Ovipo- sition usually takes place" in tandem, but I have observed unattended females prob- ing algal mats. Dec. 1975 PROVONSHA: ZYGOPTERA OF UTAH FIG. 11 • Amphiagrion abbreviatum 385 fU''\ m^.^-. • .-- • Telebasis salva (Hagen, 1861) Fig. 12 T. salva ranges from the southwestern United States to Venezuela. I have seen only three specimens taken in Utah. They were all collected at a small spring-fed pond near St. George, Washington Co.: two in 1941 (Univ. of Utah Coll.) and one in 1959 (Dixie College Coll.). Re- cently the area has been converted into a golf course, greatly changing the origi- nal habitat. It is possible that this species no longer occurs in the state. Coenagrion resolutum (Hagen, 1876) Fig. 12 This species is known from throughout most of Canada and Alaska, the northeast- ern states, and higher elevations in the West. In Utah it was found at small ponds and slow-flowing high meadow streams above 7,000 feet, where they emerge dur- ing the month of July. Evidently all ovi- position is in tandem. EnaUagma anna Williamson, 1900 Fig. 13 E. anna is restricted to the western United States. This species is fairly com- mon throughout the mountainous areas Coenagrion resolutum Telabasis salva ^■^^^ FIG, 13 of northern Utah at altitudes between 4,200 and 7,000 feet. Nymphal develop- ment takes place in rivers and streams with slow to moderate currents. The flight period ranges from mid-May through the first of September, and, like most other species of EnaUagma, ovipo- sition takes place in tandem. 386 GREAT BASIN NATURALIST Vol. 35, No. 4 Enallagma boreale Selys, 1875 Fig. 14 The range of this species includes most of Canada and the northern United States with isolated populations at higher al- titudes in the southwest. Next to Ischnura perparva Selys, it is the most common species in Utah, being absent only from the desert areas in the southeast. It is most frequently found at lakes and ponds and rarely at slow-flowing streams. Col- lection sites ranged from 4,000 to 9,000 feet. It is an early emerger, and the first record I have for the state is 8 May. None were taken after mid-August. Oviposition usually takes place in tandem. Enallagma carunculatum Morse, 1895 Fig. 15 With the exception of a break along the Rocky Mountains, this species is found transcontinentally in southern Canada and the northern United States, extending southward in the West into northern Mexico. This is one of the more common and widely adapted species in Utah, often occurring in great numbers. Although it was most frequently found at small ponds, it was also taken at streams, rivers, and some of the larger lakes and reservoirs at altitudes between 3,000 and 8,000 feet. E. carunculatum is one of the few species which can develop in brackish water, such as that occurring in drainage ditches west of Salt Lake Cit}^ near the Great Salt Lake, where it was found in close as- sociation with E. clausum Morse. This as- sociation has also been reported for other brackish waters such as Pyramid Lake, Nevada (Kennedy 1917). Emergence be- gins in late May, and oviposition takes place in tandem. vr^ FIG, 15 • Enallagma carunculatum imt ^ Enallagma civile (Hagen, 1861) Fig. 16 With the exception of the far northwest, E. civile has a wide range over most of the United States and extends as far south as the West Indies and Colombia. Al- though this is one of the most common species in North America, it was taken at only a few scattered localities in Utah. Locally abundant populations were taken in the Sevier River drainage system in Millard Co., but only small numbers were taken at most other collecting sites. Nymphs were taken in lakes, ponds, and streams and rivers with slow currents at altitudes from 2,800 to 6,000 feet. Like its close relative E. carunculatum, emergence begins in late May and continues through most of the summer. Dec. 1975 PROVONSHA: ZYGOPTERA OF UTAH 387 FIG, 16 • Enallagma civile * Enallagma ebrium W Enallagma clausum Morse, 1895 Fig. 17 This species is found in the western United States and southwestern Canada with isolated populations in Ontario and Quebec. In Utah it is restricted to the western half of the state at altitudes be- tween 4,200 and 6,300 feet. The nymphs W] • Enallagma clausum M& may be found in slow streams and lakes with some wave action. Enallagma clau- sum is tolerant to saline conditions and was frequently found in association with E. carunculatum at brackish waters in the western desert. UnHke most species of Enallagma, adults prefer to rest on bare ground rather than on vegetation; when they are at rest the wings are frequently held at a slight angle away from the body. Emergence begins in late May and con- tinues through most of the summer. Enallagma cyathigerum (Charpentier, 1840) Fig. 18 This is a Holarctic species, being found in Europe, Asia, Canada, and most of the northern United States. It is one of the more common species in Utah and is ab- sent only from the western deserts. Al- though most common at small ponds, it was frequently taken at streams wdth slow to moderate currents. With the ex- ception of Coenagrion resolutum, it was the only species taken above 9,000 feet. Emergence begins in mid-June, and adults can be found through early September. Enallagma ebrium (Hagen, 1861) Fig. 16 The range of this species is reported as Boreal North America. It has been taken at only two localities in Utah. Fourteen specimens were collected at the Weber River, West Weber, Weber Co., by C. J. D. Brown (1934), and two adults were collected during this present study at marshy areas along the Bear River in Cache Co. Both of the above-mentioned collections were made in July. Walker (1953) records the flight period in Canada to be from 30 May through 24 August. Enallagma praevarum (Hagen, 1861) Fig. 13 This species ranges from the southern United States to southern Mexico. In Utah it is restricted to desert streams and springs at 2,800 to 5,800 feet elevation in the unique southwest corner of the state. Although very closely related to E. anna, at no place did their ranges overlap. Emergence begins in mid-May and con- tinues through the month of June. 388 GREAT BASIN NATURALIST FIG, 18 • Enallagma cyathigerum Ischnura barberi Currie, 1903 Fig. 19 Ischnura barberi is known only from the western United States. All collecting sites in Utah were at ponds in the west- ern portion of the state at altitudes rang- ing from 2,800 to 4,730 feet. This species appears to be rare in numbers, since no more than two specimens were taken at any one locality. Oviposition was never observed. All collections were made dur- ing the months of June and July. Ischnura cervula Selys, 1876 Fig. 20 Southwestern Canada, western United States, and northern Mexico are included in the range of this species. It is fairly common and was frequently found at ponds and slow streams with a preference for shallow marshy areas. Samples were from altitudes ranging from 4,200 to 7,000 feet. Oviposition does not usually take place in tandem. Emergence begins in early May and continues through most of the summer. Ischnura damula Calvert, 1901 Fig. 21 This species is known in southern Canada from Manitoba west and the west- Vol. 35, No. 4 Ischnura barberi Ischnura demcrsa )> ern United States. Except for the warm springs in Tooele Co., its distribution in Utah is restricted to the southern portion of the state. All specimens were collected from small ponds and springs with dense stands of cattails along the margins. All sites were between an altitude of 4,225 and 5,825 feet. Females were observed ovipos- FIG. 20 Dec. 1975 PROVONSHA: ZYGOPTERA OF UTAH FIG. 21 • ISCHNURA DAMULA 389 FIG. 22 SCHNURA DENTICOLLIS l^-f iting singularly and in tandem. The ear- liest record I have for the state is 23 May. It is probable that emergence be- gins in early May and continues through early July. Ischnura demorsa (Hagen, 1861) Fig. 19 Ischnura demorsa is reported from the western United States and Mexico. This close relative of /. perparva is rare in Utah, however. Only two specimens were taten during this study, both from Mc- Cracken Spring in San Juan Co. at an al- titude of 4,900 feet. I have no emergence data for this species. Ischnura denticollis (Burmeister, 1839) Fig. 22 This species is found only in the south- western United States and Mexico. In Utah it is restricted to the cold deserts in the western half of the state. It is equally suited to both ponds and streams and is tolerant to a wide range of altitudes, being found from 2,700 to 6,300 feet. Oviposition usually takes place in tan- dem. This species tends to emerge some- what later than most other Utah isch- nurans. No specimens were taken before the first of June, and 15 August was the last date they were encountered. Ischnura perparva Selys, 1876 Fig. 23 This species is known from the western United States and southwestern British Columbia. This is by far the most common damselfly in Utah. It was found at 87 of the 152 localities sampled during this FIG. 23 SCHNURA PERPARVA 390 GREAT BASIN NATURALIST Vol. 35, No. 4 Study (almost twice as many as any other species). Its great abmidance can be at- tributed, in part, to its ability to survive in many diverse habitats. It was found in most aquatic situations between 4,200 and 7,500 feet where there was sufficient vege- tation and a high enough minimum tem- perature to support zygopteran forms. Al- though oviposition may take place in tan- dem, it is usually accomplished unattend- ed by the male. Emergence begins in early May and continues throughout most of the summer. Acknowledgments The author wishes to thank Dr. George F. Edmunds, Jr., University of Utah; Dr. Wilford J. Hanson, Utah State University; Dr. Vasco M. Tanner, Brigham Young University; and Dr. Andrew H. Barnum, Dixie College, for the loan of specimens. Gratitude is also extended to Mrs. Leonora K. Gloyd, Museum of Zoology, University of Michigan, and Dr. Minter J. Westfal'l, Jr., University of Florida, for verification of identifications and advice given during this study. Special appreciation is ex- pressed to Dr. W. P. McCafferty, Purdue University, for his encouragement, advice, and assistance both during the study and in the preparation of the manuscript. References BiCK, G. H., AND J. C. BicK. 1961. An adult population of Lestes disjunctus australis Walker (Odonata: Lestidae). Southwestern Nat. 6(3-4): 111-137. . 1970. Oviposition in Archilestes gran- dis (Rambur) (Odonata: Lestidae). Ent. News 81:157-163. BiCK, G. H., AND L. E. HoRNUFF. 1972. Odo- nata collected in Wyoming, South Dakota, and Nebraska. Proc. Entomol. Soc. Wash. 74(l):l-8. BiCK, G. H., AND D. SuLZBACK. 1966. Repro- ductive behavior of the damselfly, Hetaerina americana (Fabricius) (Odonata: Calopterygi- dae). Anim. Behav. 14:156-158. Brown, C. J. D. 1934. A preliminary list of Utah Odonata. Univ. Mich. Occas. Pap. Mus. Zool. no. 291, 17 pp. Johnson, C. 1961. Breeding behavior in Hetae- rina americana (Fabricius) and H. titia i(Drury) (Odonata: Agriidae). Canadian Ent. 93:260-266. . 1972. The damselflies (Zygoptera) of Texas. Bull. Florida State Mus., Biol. Sci. 16(2):55-128. Kennedy, C. H. 1915. Notes on the life history and ecology of the dragonflies (Odonata) of Washington and Oregon. Proc. U.S. Natl. Mus. 99(2107) :259-345. . 1917. Notes on the life history and ecology of the dragonflies (Odonata) of cen- tral California and Nevada. Proc. U.S. Natl. Mus. 52(2192) :483-635. KoRMONDY, E. J. 1957. Records of western Odo- nata with notes on Amphiagrion abbreviatum (Selys). J. Kansas Ent. Soc. 30(3) : 108-110. Larson, W. P. 1952. The dragonflies (Anisop- tera) of Utah. Unpublished master's thesis, Dept. of Zoology, Univ. Utah, Salt Lake City, 95 pp., 30 plates. Martin, R. D. C. 1939. Life history of Agrion aequabile and Agrion maculatum. Ann. Ent. Soc. Amer. 32:601-618. MussER, R. J. 1962. Dragonfly nymphs of Utah (Odonata: Anisoptera) Univ. Utah Biol. Sen, vol. 12, no. 6, Oct. 1962. Paulson, D. R. 1970. A list of the Odonata of Washington with additions to and dele- tions from the state list. Pan-Pacific Ent. 46(3): 194-198. Provonsha, a. v., and W. P. McCafferty. 1973. Previously unknown nymphs of western Odonata (Zygoptera: Calopterygidae, Coenagrionidae) . Proc. Entomol. Soc. Wash- ington 75(4): 449-454. Smith, R. F., and A. E. Pritchard. 1956. Odonata. pp. 106-153. In: Aquatic insects of California. R. L. Usinger, ed. Univ. Calif. Press, Berkeley, and Los Angeles. Walker, E. M. 1953. The Odonata of Canada and Alaska. Vol. 1, Univ. of Toronto Press, Toronto XI + 292 pp. Whitehouse, F. C. 1941. British Columbia dragonflies (Odonata) with notes on distribu- tion and habits. Amer. Midland Nat. 26: 488-557. NEW SYNONYMY AND NEW SPECIES OF AMERICAN BARK BEETLES (COLEOPTERA: SCOLYTIDAE), PART IP Stephen L. Wood- Abstr..\ct. — New synoiiyiiij- is proposed as follows: Pityophthoius Eichhoff (= GmUhophorus Schedl. Gnathopfithorus Wood). Araptus confinis (Blandford) (= Neopilyophthorus glabricollis Schedl), A. eruditus (Srhedl) (= Neodryocoetes buscki Blackiiian), A. hymenaeae (Eggers) (= Neo- dryocoetes humilis Blackmail), A. schedli (Blackmail) (= Neodryocoetes lenis Blackmail), A. lenel- lus (Schedl) (= Ctenyophthorus mexicanus Schedl, Neodryocoetes granulatus Schedl, Araptus cuspi- dus Wood), Coccotrypes carpophagus (Hornung) (= Coccotrypes liberiensis Hopkins, Coccotrypes pimctatulus Eggers), C. daclyliperda (Fabricius) ( =- Coccotrypes bassiavorus Hopkins). C. robuslus Eichhoff (= Coccotrypes cylindricus Schedl). Cryptocarenus heveae (Hagedorn) (= Cryptocarenus caraibicus Eggers). Hypothenemus setosus (Eichhoff) (^ Stephanoderes congonus Hagedorn). Micro- corthylus minutus Schedl (= Microcorthylus minutissimus Schedl), Pseudopityophthorus limbatus Eggers (= Pseudopityophthorus rnicans 'Wood), Xyleborus obliquus (LeConte) (= Xyleborus gil- vipes Blandford, A', brasiliensis Eggers, illepidus Schedl). Hypothenemus javanus Eggers is a valid species. The genus Dacnophthorus. type-species Gnathophthorus clematus Wood, is described as new to science. The following species are described as new to science: Araptus consobrinus, A. micaceus, Pityophthorus explicitus, and P. inceptis (Mexico). P. costatus and P. mendosus (Costa Rica). P. de- gener and P. timidulus (Panama). P. amiculus (Mexico. Costa Rica), and P. dissolutus (Costa Rica, Panama). Xyleborus californicus (California). X. incultus. X. molestulus (Panama), and tristiculus (Brazil). On the following pages seAeral newly discovered cases of synonym}', one new genus, and 14 sjDecies new^ to science are presented for American Scolytidae. The specific synonymy is presented alphabeti- cally for convenience of reference. The species new to science represent the genera Araptus (2), Pityophthorus (8), and Xyleborus (4). They are from the fol- lowing countries: United States (1), Mexico (4), Costa Rica (2), Panama (4), Brazil (1), Mexico and Costa Rica (1), Costa Rica and Panama ( 1 ) . New Synonymy Pityophthorus Eichhoff Pityophthorus Eichhoff. 1864, Berliner Ent. Zeit. 8:39 (Type-species: Bostrichus lichtensteini Ratzeburg, subsequent designation by Hop- kins. 1914, Proc. U.S. Nat. Mus. 48:127) Gnathophorus Schedl, 1935 (nee Kirby, 1837). Rev. de Ent. 5:342 ( Type-species : Gnatho- phorus sparsipilosus Schedl. monobasic). New synonymy Gnathophthorus Wood, 1962, Great Basin Nat. 22:76 (Replacement name for Gnatho- phorus) . New synonymy The holotype of Gnathophorus sparsi- pilosus Schedl was examined and com- pared directly to a series of this species in my collection. It fits well within the limits of the genus Pityophthorus anatom- ^Part of this research \-\ -Department of Zoology . sponsored bj- the National Scienc Brigham Yoiing University, Prov ically as well as biologically. For this rea- son the names Gnathophorus and its re- placement, Gnathophthorus, must be placed in synonymy as indicated above. The five species subsequently assigned to this genus must be transferred to another genus that is described below. Araptus confinis (Blandford) Pityophthorus confinis Blandford, 1904, Biol. Centr. Amer. Coleopt. 4(6):241 (Lectotype, male; Jalapa. Veracruz. Me.xico; British Mus. Nat. Hist., present designation) Neopilyophthorus glabricollis Schedl, 1938, Archiv Naturgesch. 7(2): 181 (Holotype. male; Teo- pisca, Chiapas, Mexico; Schedl Coll.). New synonymy The first syntype in the type series of Pityophthorus confinis Blandford is here designated as the lectotype of that species. This lectotype was compared directly to my males from Guatemala City, Guate- mala, and was found to be identical in all respects. My series was later compared directly to the male holotype of Neopity- ophthorus glabricollis Schedl. Except for the loss of declivital and frontal setae, an apparent result of abrasion on the Schedl type, these specimens are also identical. As indicated above, the junior name must be placed in synonymy. The lectotype of confinis is labeled "Type" and has been considered as the type for many years. e t'oundation. o, Utah 84602. Scolytidae contribution No. Gl 391 392 GREAT BASIN NATURALIST Vol. 35, No. 4 Araptus eruditus (Schedl) Neopityophthorus eruditus Schedl. 1938, Archiv Naturgesch.. n. f., 7(2): 182 (Holotype, fe- male; Mexico; Schedl Coll.) Neodrrocoetes buscki Blackman, 1942, Proc. U.S. Nat. Mus. 92:192 (Holotype. female; Cabima, Panama; U.S. Nat. Mus.). New synonymy The female holotype of Neopityoph- thorus eruditus Schedl and the female holotype of Neodryocoetes buscki Black- man were compared direct!}- to my series. All represent the same species. The last visible abdominal sternum is armed by a subtuberculate callus which aids in the identification of this species. Araptus hymenaeae (Eggers) Neodryocoetes hymenaeae Eggers. 1933. Mem. Trav. Lab. d'Ent. Mus. Nat! d'Hist. Nat.. Paris 1(1) :9 (Holotype, female; Gourdon- ville, French Guiana; Paris Mus.) Neodryocoetes humilis Blackman, 1942, Proc. U.S. Nat. Mus. 92:188 (Holotype, female; Bonito, Pernambuco. Brazil; U.S. Nat. Mus.) New synonymy The \ariable frons of this species has led to the description of several synonyms. Following a study of the holotype and six cotypes of Neodryocoetes hymenaeae Eg- gers, the female holotype of Neodryocoetes humilis Blackman, and 76 other speci- mens, it was concluded that only one spe- cies was represented and that Blackman's name must be placed in synonymy. Araptus schedli (Blackman) Neodryocoetes schedli Blackman, 1942, Proc. U.S. Nat. Mus. 92:195 (Holotype, male; Tampico. Mexico; U.S. Nat. Mus.) Neodryocoetes lenis Blackman, 1942, Proc. U.S. Nat. Mus. 92:198 (Holotype. male; Cordoba. Veracruz. Mexico; U.S. Nat. Mus.). New synonymy The male holotypes of Neodryocoetes schedli Blackman and Neodryocoetes lenis Blackman were compared directly to one another and to several specimens inter- cepted at New York in seeds of Mexican origin. I am unable to detect any differ- ences among these specimens that might justify the continued recognition of two names. Araptus tenellus (Schedl) Neodryocoetes tenellus Schedl, 1951, Dusenia 2: 109 (Holotype. male; Chiapas, Mexico; Schedl Coll.) Ctenyophthorus mexicanus Schedl. 1963. Ent. Arb. Mus. Frey 14:162 (Holotype, female; Tram- paluz, Escarcaga. Campeche, Mexico; Schedl Coll.). New synonymy Neodryocoetes granulatus Schedl, 1964, Reichen- bachia 3:311 (Replacement name for Cteny- ophthorus mexicanus Schedl). New synon- ymy Araptus cuspidus Wood, 1974, Brigham Young Univ. Sci. Bull.. Biol. Ser. 19 (1):46 (Holo- type, female; 8 km E San Bias, Nayarit. Mex- ico; Wood Coll.). New synonymy The holotype of Neodryocoetes tenellus Schedl and the allotype of Araptus cus- pidus Wood were compared directly to one another. Except for minor differences in the convexity of the frons, they are identical. The peculiar abdominal stern- um 5 was not mentioned by Schedl. The Schedl holotype of Ctenyophthorus mexi- canus is a female, not a male as stated in the original description. It was compared directly to the holotype of cuspidus and was found to be identical. The two junior names and the replacement name, Neo- dryocoetes granulatus Schedl, must be placed in synonymy as indicated above. Coccotrypes carpophagus (Hornung) Bostrichus carpophagus Hornung, 1842, Stettiner Ent. Zeit. 3:116 (Syntypes; intercepted in Germany from "Betelniissen" of "Ostindien" origin) Coccotrypes liberiensis Hopkins, 1915, U.S. Dept. Agric. Kept. 99:47 (Holotype, female; Mount Coffee, Liberia; U.S. Nat. Mus.). New sy- nonymy Coccotrypes punctatulus Eggers, 1951. Ent. Blatt. 45-46:151 (Holotype, female; Insel St. Thomas, Virgin Islands; deposited in Eggers Coll., apparently on loan to Schedl). New synonymy The holotypes of Coccotrypes liberiensis Hopkins and C. punctatulus Eggers and the syntypes of Bostrichus carpophagus Hornung were all examined and com- pared directly to my series of this species. Only one species is represented by the three names. Coccotrypes dactyliperda (Fabricius) Bostrichus dactyliperda Fabricius, 1801, Systema Elcutheratorium 2:387 (Syntypes; in date pits intercepted in Europe; Copenhagen and Berlin museums) Coccotrypes bassiavorus Hopkins. 1915, U.S. Dept. Agric. Rept. 99:47 (Holotype, female; Wash- ington, D.C.; M.S. Nat. Mus.). New synon- ymy Several female specimens of Cocco- trypes dactyliperda (Fabricius) that were Dec. 1975 WOOD: AMERICAN SCOLYTIDAE 393 compared by Eggers to the Fabricius syn- types were compared directly to the fe- male holotype of C. hassiavorus Hopkins. Since they are identical in all respects, Hopkins's name must be placed in synon- >'my. Coccotrypcs robustus Eichhoff Coccotrrpes robuslus Eiclihoff. 1878. Mem. Soc. Roy. Sci. Li6ge (2)8:513 (Syntypes?; Cuba; presumably- lost witJi Hamburg Mus.) Coccotrrpes cylindricus Schedl, 1949, Tijdschr. Ent. 91:116 (Holotype, female; Crucos. Cuba; Schedl Coll.). New synonymy A female of Coccotrypes robustus Eich- hoff that was compared by Hopkins to a syntype and the holotype of C. cylindri- cus Schedl were compared to my series of this species. Since all represent the same distinctive species, Schedl's name must be placed in synonymy. Cryptocarenus heveae (Hagedom) Stephanoderes heveae Hagedorn, 1912, Rev. Zool. Afr. 1:338 (Syntypes, female; Eala, Congo; Tervuren Mus.) Cryptocarenus caraibicus Eggers. 1937, Rev. de lEnt. 7:82 (Holotype, female; Guadeloupe; U.S. Nat. Mus.). New synony?ny The four female syntypes of Stephan- oderes heveae Hagedorn and the female holotype of Cryptocarenus caraibicus Eg- gers were compared to my material. All of these specimens represent the same species. Schedl (1975, Ent. Blatt. 71:43) treated C. lepidus Wood as a synonym of C. car- aibicus. These species are easily confused but quite distinct. Hypothenemus setosus (Eichhoff) Hypoborus (?) setosus Eichhoff, 1867, Berliner Ent. Zeitschr. 11:391 (Syntypes, Guadeloupe; one syntype in U.S. Nat. Mus., others lost with Hamburg Mus.) Stephanoderes congonus Hagedorn, 1912, Rev. Zool. Afr. 1:337 (Lectotype, female; Eala, Congo; Tervuren Mus., present designation). New synonymy Stephanoderes congonus Hagedorn was based on a syntypic series. Four of those syntypes are in the Musee Royal de I'Afri- que Central, Tervuren, labeled as one "Holotypus" and three "Paratypus." These type designations, however, have not beeen mentioned in print and are con- sidered invalid. The "Holotypus" is se- \erely damaged but recognizable. All four specimens bear identical data labels: "Musee du Congo Beige; Eala; Rakusu (H?? 1140 M, 1911); D. Hevea brasilien- sis." Because the first specimen is severely damaged, I designate the third specimen as the lectotype. The above lectotype was compared to several of my series of setosus (Eichhoff) that previously had been compared di- rectly to the only known syntype of setosus. Only one species is represented by this material. It is a species distinct from javanus (Eggers), as noted below. Other synonyms of setosus include Stephano- deres obscurus Eichhoff (nee Ferrari) and S. depressus Eichhoff. Hypothenemus javanus Eggers, new status Stephanoderes javanus Eggers. 1908. Ent. Blatt. 4:215 (Lectotype. female; Java; U.S. Nat. Mus.. designated by Anderson and Ander- son, 1971, Smithsonian Contrib. Zool. 94:16) This species was placed in synonymy under setosus (Eichhoff) by Schedl (1962, Ent. Blatt. 58:204). Using the lectotype of javanus as a point of reference, I (Wood, 1972, Great Basin Nat. 32:51) added several additional synonyms. A re- examination of the types of javanus^ seto- sus, and all synon}Tns, and 172 other specimens assigned to setosus, clearly in- dicates the existence of two easily dis- tinguished species. Hypothenemus javanus ( = Stephanoderes obesus Hopkins, S. phil- ippinensis Id^opkins, S. bananensis Eggers, S. kalshoveni Schedl, 5. subagnatus Eg- gers) has a stouter body form (2.2 times as long as wide), only 12-18 coarse asper- ities on the anterior slope of the pronotum, 2-4 denticles on the anterior margin of the pronotum, and several minor differences in details of sculpture. It occurs in Indo- nesia, Philippines, Florida, Cuba, Jalisco (Mexico), Congo, Ghana, and the Cam- erouns. Hypothenemus setosus has the body 2.4 times as long as wide, more than 25 pronotal asperities, 6-8 denticles on the anterior margin of the pronotum, and other differences in details of surface sculpture. It occurs from Chiapas (Mex- ico) to Brazil, in the Congo, and in the Camerouns. Therefore, javanus must be restored as a valid name to designate the above species. 394 GREAT BASIN NATURALIST Vol. 35, No. 4 Microcorthylus minutus Schedl Microcorthylus minutus Schedl, 1950, Dusenia 1: 160 (Syntypes, females; Nova Teutonia, Santa Catarina. Brazil; Schedl and Plaumann colls.) Microcorthylus minutissimus Schedl, 1952, Dus- enia 3:361 (Syntypes; Jamaica; Schedl Coll.). New synonymy Female syntypes of minutus Schedl and minutissimus Schedl were compared di- rectly to one another and to 36 other spec- imens from Veracruz, Honduras, Costa Rica, Panama, and Brazil. Only one small, distinctive species is represented by this material. Pseudopityophthorus limbatus Eggers Pseudopityophthorus limbatus Eggers, 1930, Ent. Blatt. 26:169 (Holotype. female; "R. d. M.," Mexico; deposited in Eggers collection, evi- dently now on loan to Schedl) Pseudopityophthorus micans Wood, 1967, Great Basin Nat. 27:44 (Holotype. male; 96 km W. Durango, Durango, Mexico; Wood Coll.). New synonymy The female holotype of Pseudopityoph- thorus limbatus Eggers and the female al- lotype of P. micans Wood were compared directly to one another and to 85 other specimens of this species and were found to represent only one species. Xyleborus obliquus (LeConte) Pityophthorus obliquus LeConte, 1878, in Schwarz. Proc. Amer. Philos. Soc. 17:432 (Holotype. female; Enterprise, Florida; Mus. Comp. Zool.) Xyleborus gilvipes Blandford, 1898, Biol. Centr. Amer., Coleopt. 4(6):205 (Holotype, female; Zapote, Guatemala; British Mus.), Neu^ synonymy Xyleborus brasiliensis Eggers, 1928, Arch. Inst. Biol., Sao Paulo 1:96 (Lectotype, female; Blumenau, Santa Catarina. Brazil; U.S. Nat. Mus.). New synonymy Xyleborus illepidus Schedl. 1941. Rev. Zool. Bot. Afr. 34:402 (Holotype. female; Deutsch Ostafrika; Schedl Coll.). New synonymy This species has been reported from areas disturbed by human activities in a pattern that suggests introduction. Its ori- gin has not been established except that allied species are Neotropical. Its occur- rence in Africa appears to be recent. In establishing the above synonymy, the holotypes of obliquus (LeConte), gil- vipes Blandford, and illepidus Schedl, the lectotype of brasiliensis Eggers, and 78 U.S., 46 Neotropical (Mexico, Guatemala, Colombia, Brazil), and 14 African (Con- go) specimens were examined and com- pared directly. There is a certain amount of variability within this material, but the variation between series is no greater than it is within a series. In addition to the above, the synonymy of this species with Under ae (Hopkins) and mexicanus Eggers has already been established. Taxa New to Science Dacnophthorus. n. gen. This genus is distinguished from Pity- ophthorus Eichhoff by the very slender body form, b}^ the very large, coarsely faceted eyes, by the large antennal club, with two partly septate sutures, by the pronotal summit being anterior to the middle of the pronotum and lacking a transverse impression posterior to it, by the distinctive elytral declivity, and by the very different habits. Frons dimorphic, male convex above, impressed below, female uniformly shal- lowly concave to lower third, then weakly convex, upper area variously sculptured and ornamented by hair; eye very large, coarsely faceted, emarginate. Antennal scape slender, elongate; funicle 5-segmen- ted, some specimens apparently 4-segmen- ted; club large, much longer than scape, sutures 1 and 2 partly septate, 3 clearly indicated by setae and rather remote from apex. Pronotum elongate, summit anterior to middle, without a transverse impression behind simimit. Scutellum rather large, flat. Elytra striate; declivity rather short, steep. Legs as in Pityophthorus. Type-species. — Gnathophthorus de- mat us Wood. Note. — The five species previously I)laced by me in (rnathophthorus must be transferred to this genus. Araptus consobrinus^ n. sp. This species is distinguished from atten- uatus Wood by the evenly convex elytral declivity (attenuatus has striae 1 impres- sed and interstriae 1 weakly elevated) i\ui\, on the female frons, by the coarser, more abimdant, longer, yellowish ves- titnre. Mai,i:.— Length LI- mm (])aratypes 1.4-1.7 mm), 2.7 times as long as wide; ( olor \ery dark brown. Dec. 1975 WOOD: AMERICAN SCOLYTIDAE 395 Frons similar to attenuatus except more nearly convex, punctures not as coarse, no indications of aciculation, median cal- lus at upper level of eyes smaller, less strongly elevated. Pronotum as in attenuatus except an- terior margin armed by about eight ser- rations. Elytra as in attenuatus except surface less brightly shining, strial punctures slightl}" larger. Declivity more evenly convex, punctures much smaller; striae 1 not impressed, interstriae 1 not elevated. Female.^ — Similar to male except frons much as in female attenuatus except frontal hair much more abundant, coarser, slightly longer, yellowish in color. Type Locality. — Six km or 4 miles N Tepic, Nayarit, Mexico. Type Material. — The male holotype, female allotype, and 20 paratypes were taken at the type locality on 13-VII-1965, 1000 m, No. 241, from a shrub; seven paratypes are from 33 km or 21 miles N Juchitlan, Jalisco, Mexico, 3-VII-1965, 1300 m, No. 177, Ficus twigs 3 cm in diameter, all by me. The holotype, allotype, and paratypes are in my collection. Araptus micaceus, n. sp. This species is distinguished from obso- letus (Blandford) by the strongly im- pressed frons, with a pair of epistomal calluses at the anterior articulations of the mandibles, by the smooth surface be- tween punctures on the area above the eyes, and by the coarser strial punctures on the declivity. Male. — Length 1.4 mm (paratypes 1.3-1.4 mm), 2.6 times as long as wide; color light brown. Frons as in obsoletus except much more strongly impressed, punctures in im- pressed area smaller, surface on area above eyes reticulate; epistomal calluses at anterior articulation of mandibles much larger. Pronotum as in obsoletus except ser- rations on anterior margin much smaller (margin subcostate) and reticulation on posterior areas more strongly impressed. Elytra as in obsoletus except strial punc- tures much larger, interstriae one and one-half times as wide as striae on disc, about twice as wide on declivity; im- pressed points obsolete on declivity. Female. — Similar to male except frons planoconvex on median two-thirds from level of antennal insertion to slightly above eyes, finely, closely punctured ex- cept impunctate on median line to upper level of eyes, punctured area bearing fine, rather long, moderately abundant hair; hair shorter, less abundant and covering a smaller area than in female obsoletus. Type Locality. — Los Corchos, Nayarit, Mexico. Type Material. — The male holotype, female allotype, and seven paratypes were taken at the type locality on lO-Vn-1965, 7 m elevation. No. 222, from a recently cut vine, by me. The holotype, allotype, and paratypes are in my collection. Pityophthorus costatus. n. sp. This abberant species has a variable number of segments in the antennal funicle; it is also distinguished by the long, costiform pronotal asperities, by the stout body form, b}' the frons, and by other characters. Female. — Length 1.1 mm (paratypes, 1.1-1.2 mm), 2.3 times as long as wide; color dark reddish brown. Frons rather strongly convex, an abrupt, moderately deep impression immediately above epistomal margin, epistomal margin distinctly elevated except for small median notch; surface smooth shining, sparsely punctured, punctures rather coarse, area above eyes somewhat reticulate; vestiture very sparse, short. Antennal funicle variable, 3-5-segmented; club ovate, small, sutures almost straight, 1 and 2 septate only at margins. Pronotum 1.0 times as long as wide; widest at base, weakly arcuate on basal third rather strongly converging toward narrowly rounded anterior margin; an- terior margin continuously costate; sum- mit just behind middle, rather indefinite; asperities long, subcostate, confused, con- tinued to basal fourth in median area; posterior areas strongly reticulate, punc- tures small, not close. Glabrous. Elytra 1.5 times as long as wide, 1.6 times as long as pronotum; sides almost 396 GREAT BASIN NATURALIST Vol.35, No. 4 Straight and parallel on basal two-thirds, rather broadly rounded behind; striae not impressed, punctures small, deej), spaced by one to two diameters of a puncture; interstriae smooth, shining, two to three times as wide as striae, punctures minute, confused, rather close. Declivity steep, convex; striae not impressed, punctures greatly reduced to obsolete; interstriae as on disc except punctures reduced to obso- lete. Vestiture confined to declivity, of sparse, short, stout interstrial setae. Male. — Similar to female except epi- stomal impression usually more strongly impressed in lateral areas, with a weak median elevation dividing this impression. Type Locality. — Tapanti, Cartago, Costa Rica. Type Material. — The female holo- type, male allotype, and 12 paratypes were taken at the type locality on 2-VII- 1963, 1300 m. No. Ha, from a liana, by me. Two paratypes are from 13 km SE Cartago, Cartago, Costa Rica, 3-VII-1963, 1800, from the same species of liana, by me. The holotype, allotype, and paratypes are in my collection. Pityophthorus inceptis, n. sp. This species is distinguished by the simple male frons and declivity, by the female frontal vestiture and by the coarse pronotal and elytral punctures. It is not closely allied to other known species. Female. — Length 1.5 mm (male para- types 1.5-1.6 mm), 2.7 times as long as wide; color very dark brown. Frons basically convex except flattened on median half from epistoma to slightly above eyes; surface shining, punctures rather coarse, moderately close; vestiture rather sparse and short except on margins of upper half of flattened area forming a dense fringe of long yellow hair, longest setae equal in length to three-fourths dis- tance from their bases to epistomal margin. Antennal club broadly obovate, 1.2 times as long as wide, suture 1 almost straight, 2 moderately ])rocurved. Pronotum 1 .07 times as long as wide; widest on basal third, moderately arcuate from base to rather broadly rounded an- terior margin; anterior margin armed by about six to eight low, basally contiguous teeth; summit at middle; asperities rather coarse, confused; posterior areas subrug- ose-reticulate, punctures coarse, deep, mostly spaced by distances equal to one- half diameter of a puncture, median line impunctate. Vestiture of sparse semi- recumbent short hair in lateral and as- perate areas. Elytra 1.7 times as long as wide, 1.7 times as long as pronotum; sides almost straight and parallel on basal three- fourths, somewhat narrowly rounded be- hind; striae not impressed, punctures rather coarse, deep, occasional punctures not in row, spaced by less than diameter of a puncture; interstriae almost smooth, shining, impunctate except for an oc- casional puncture near declivity. Declivity steep, convex; striae not impressed, punc- tures reduced, about one-third as large as on disc, distinctly impressed; interstriae as on disc except 1 and 3 each with a row^ of fine punctures. Vestiture confined to sides and declivity, that on sides of minute strial hair, that on declivity of fine, rather short interstrial setae on odd-numbered interstriae. Male. — Similar to female except frons more uniformly convex, without brush of long hair; serrations on anterior margin of pronotum slightly larger. Type Locality. — Six km or four miles W Quiroga, Michoacan, Mexico. Type Material. — The female holo- type, male allotype, and one male para- type were taken at the type locality on 17-VI-1965, 2200 m. No. 72, from a shrub- by herbaceous ]:)lant, by me. The holotype, allotype, and paratype are in my collection. Pityophthorus timididus. n. sp. This species is distinguished from men- dosus Wood by the larger size, by the coarser pronotal punctures, and by slight differences on the elytral declivity. Both species are allied to mandihularis Schedl. Male. — Length 2.0 mm (paratypes 1.8-2.0 mm), 2.5 times as long as wide; color very dark brown. Frons convex above eyes, upper half of area below upper level of eyes rather abruptly, strongly, transversely impressed almost from eye to eye, a smaller trans- Dec. 1975 WOOD: AMERICAN SCOLYTIDAE 397 verse impression in lateral areas immed- iately above epistoma; surface shining, coarsely, rather closely punctured; ves- titure sparse exce])t on epistoma. Antennal club oval, 1.3 times as long as wide, su- tures 1 and 2 moderately arcuate, 2 at middle of club. Pronotum. 1.1 times as long as wide; sides on basal half almost straight, sub- parallel, rather broadly rounded in front; anterior margin armed by about 12 low serrations; summit at middle; asperities on anterior slope rather coarse, close, con- fused; posterior areas smooth, shining, with moderately abundant minute im- pressed points, punctures rather coarse, deep, moderately close, irregularly spaced by about one to two diameters of a punc- ture. Glabrous except a few setae on margins. Elytra 1.6 times as long as wide, 1.6 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; striae 1 weakly, others not impressed, punctures rather small, deep, spaced by diameter of one puncture; interstriae smooth, shining, a few small punctures on 1 near declivity, others impunctate. Declivity steep, con- vex, shallowly bisulcate; interstriae 1 dis- tinctly elevated, armed by a row of about seven rounded tubercles, 2 moderately impressed, slightly wider than 1, smooth, shining, devoid of punctures, 3 convex, as high as 1, armed as on 1, lateral areas with punctures somewhat confused. Ves- titure confined to declivity, consisting of rather short, moderately coarse, sparse, interstrial setae on odd-numbered inter- striae. Female. — Similar to male except frons below upper level of eyes shallowly, broadly, transversely impressed, surface regular, not granulate, punctures moder- ately fine, vestiture sparse, declivital sul- cus much less strongly impressed, gran- ules on interstriae 1 and 3 minute. Type Locality. — Volcan Chiriqui, Chiriqui, Panama. Type Material. — The male holotype, female allotype, and 12 paratypes were taken at the type locality on 11-1-1964, 1800 m, No. 407, from a sapling 4 cm in diameter, by me. The holotype, allotype, and paratypes are in my collection. Pityophthorus mcndosiis, n. s]). This species is distinguished from limidulus Wood by the smaller size, by the smaller pronotal punctures, by the shorter, stouter elytra 1 bristles, and by the ( omplete absence of tubercles on female declivital interstriae 1 and 3. Male. — Length 1.7 mm (paratypes 1.5-1.7 mm), 2.8 times as long as wide; color very dark brown. Frons as in timidulus except callus at level of antennal insertion usually more strongly develojied and with a small cusp at dorsomedian extremity. Pronotal punc- tures averaging slightly smaller than in timidulus. Elytra as in timidulus except declivital setae slightly shorter and dis- tinctly stouter. Female. — As in female timidulus ex- cept frons less distinctly impressed, decliv- ital granules absent, and declivital setae shorter and stouter. Type Locality. — San Isidro del Gen- eral, San Jose, Costa Rica. Type Material. — The male holotype, female allotype, and 18 j)ara types were taken at the type locality on 5-Xn-63, 1000 m. No. 282, from "Fosforo" leaf petioles, by me. The holotype, allotype, and paratypes are in my collection. Pityophthorus degener, n. sp. This species is distinguished from the allied timidulus Wood and mendosus Wood by the much more weakly im- pressed male frons, by the strongly im- pressed elytral declivity, and by the finer pronotal punctures. Male. — Length 2.0 mm (paratypes 1.7- 2.0 mm), 2.8 times as long as wide; color very dark brown. Frons convex, but with abrupt, irreg- ular, shallow, transverse impressions just below upper level of eyes and just above epistoma; surface subshining, coarsely, closely punctured; almost glabrous ex- cept at epistomal margin. Pronotum and elytral disc as in timidulus and mendosus except pronotal punctures much smaller, slightly closer; elytral declivity strongly bisulcate, interstriae 1 almost as high as wide, 3 higher than 1, each armed by a 398 GREAT BASIN NATURALIST Vol. 35, No. 4 row of moderately coarse tubercles. Ely- tral vestiture as in timidulus. Female. — Similar to male except me- dian half of frons flattened below upper level, surface smooth, with punctures rather fine, close, deep, with rather abun- dant, fine, moderately long hair; declivital sulcus half as deep, interstriae 1 and 3 unarmed. Type Locality. — Volcan Chiriqui, Chiriqui, Panama. Type Material. — The male holotype, female allotype, and seven paratypes were taken at the type locality on 11-T 1964, 1800 m. No. 384, from a tree limb 8 cm in diameter, by me. The holotype, allotype, and paratypes are in my collection. Pityophthorus amiculus, n. sp. This species is distinguished from deg- encr Wood by the absence of sexual di- morphism, and by the different frons, pronotum, and other characters. Male. — Length 1.8 mm (paratypes 1.7- 1.9 mm), 2.7 times as long as wide; color reddish brown. Frons convex above eyes, with median line smooth, impunctate, a median callus at upper level of eyes, area below upper level of eyes broadly, shallowly, trans- versely impressed to epistoma; surface almost smooth, punctures moderately coarse, close, deep; almost glabrous ex- cept along epistoma. Pronotum 1.2 times as long as wide; outline as in degener; asperate area con- tinued slightly into lateral portion of pos- terior half as weak rugae; posterior areas smooth, shining, with rather numerous impressed points, punctures rather fine, moderatel}" close, median line impunc- tate. Sparse setae confined to asperate area. Elytra 1.5 times as long as wide, 1.3 times as long as pronotum; outline as in timidulus Wood; striae not impressed, punctures rather small, deep, close, rows occasionally slightly confused; interstriae smooth, shining, slightly irregular, with a few impressed lines, impressed points moderately abundant, 1 with obscure sub- granulate punctures almost to base, others with an occasional similar puncture. De- clivity steep, strongly bisulcate, sulcus commencing slighth' behind middle of ely- tral length; striae 1 and 2 with punctures reduced in size but clearly impressed; interstriae 1 almost as high as wide, 3 higher than 1, each armed by a row of about eight subacute tubercles of moderate size, 2 wider than 1, its surface ascending laterally, unarmed. Vestiture consisting of interstrial bristles on declivity, continued to middle of disc on odd-numbered inter- striae; longest bristles rather slender, equal in length to twice width of an in- terstriae. Female. — Similar to male in all re- spects. Type Locality. — Guapiles, Limon, Costa Rica. Type M.a.terial. — The male holotype, female allotype, and 12 paratypes were taken at the type locality on 22-Vin- 1966, 100 m. No. 121, from a liana, by me. Eighteen paratypes are from Coatzocoal- cos, Veracruz, Mexico, 26-VL1967, 30 m. No. 103, liana, by me. The holotype, allotype, and paratypes are in my collection. Pityophthorus dissolutus. n. sp. This species is distinguished from ex- plicitus Wood by the finer frontal punc- tures, by the finer granules on the pro- notal disc, and by the shallower declivital sulcus that is armed by finer granules. Male. — Length 1.5 mm (paratypes 1.4-1.6 mm), 2.7 times as long as wide; color dark brown. Frons convex, a fine median tubercle on epistomal process; surface strongly re- ticulate, punctures rather fine, deep, spaced by diameter of a puncture or more; vestiture fine, sparse, inconspicuous. Pronotum 1.1 times as long as wide; widest on basal half, sides feebly arcuate, subjiarallel. rather narrowly rounded in front; anterior margin armed by about eight moderately coarse serrations; sum- mit at middle, indefinite; asperities rather fine, confused; posterior areas strongly re- ticulate, j)unctures rather fine, moderately close. Vestiture confined to marginal and asperate areas. Elytra 1.6 times as long as wide, 1.5 times as long as pronotum; sides almost Dec. 1975 WOOD: AMERICAN SCOLYTIDAE 399 straight and parallel on basal two-thirds, rather broadly rounded behind; striae 1 weakly, others not impressed, punctures fine, distinct, decreasing in size posterior- ly; interstriae almost smooth, with some indistinctly impressed lines, about three times as wide as striae on basal fourth, six times as wide near declivity. Declivity rather steep, shallowly bisulcate; striae 1 deejily impressed, jninctures small, indis- tinct, surface ascending gradually to lateral convexity, striae 2 obscure; inter- striae 1 almost as high as wide, almost smooth, with a row of fine tubercles, 2 and lateral areas shining, rather densely cov- ered by impressed points, 3 slightly higher than 1, similarly armed. Vestiture of minute strial hair, and, on posterior half, interstrial bristles on odd-numbered inter- striae; bristles sparse, rather fine, short. Female.^ — Similar to male except epi- stomal tubercle evidently absent, and de- clivital impressed points reduced or ab- sent. Type Locality. — Thirteen km SE Car- tago, Cartago, Costa Rica. Type Material. — The male holotype, female allotype, and 27 paratypes were taken on 24-IX-1963, 1800 m. No. 201, from a liana 1 cm in diameter. Two para- types are from Tapanti, Cartago, Costa Rica, 24-X-63, 1300 m. No. 244, liana; and six paratypes are from Volcan Chir- iqui, Chiriqui, Panama, 11-1-1964, No. 394, in a sapling; all were taken by me. The holotype, allotype, and paratypes are in my collection. Pityophthorus explicitus, n. sp. This species is distinguished from dissolutus Wood by the coarser frontal punctures, by the larger granules on the pronotal disc, and by the deeper declivital sulcus that is armed by coarser granules. Male. — Length 1.5 mm (paratypes 1.5-1.7 mm), 2.7 times as long as wide; color very dark brown. Frons as in dissolutus except punctures distinctly larger, spaced by less than di- ameter of a puncture. Pronotum as in dissolutus except granule on lateral mar- gins of discal punctures distinctly larger. Elytra as in dissolutus except declivital sulcus deeper, wider, interstriae 2 not ascending laterally on lower half. Female. — Female similar to male in all respects. Type Locality. — Nine km NE Teziut- lan, Puebla, Mexico. Type Material. — The male holotype, female allotype, and 14 paratypes were taken at the type locality on 2-Vn-1967, 1600 m. No. 143, from a liana, by me. The holotype, allotype, and paratypes are in my collection. Xyleborus calif ornicus, n. sp. This species almost certainly was intro- duced into California from another area, })ossibly from South America or south- eastern Asia. Among North American species it might be confused with pubes- cens Zimmermann, but it is distinguished by the smaller size, by the more abun- dant pubescence, and by the reticulate- granulate interiors of the strial punctures on the declivity. Female. — Length 2.0 mm (paratypes 2.0-2.2 mmj, 2.9 times as long as wide; color yellowish brown. Frons rather strongly convex; surface strongly reticulate, a few small granules from epistoma to upper level of eyes. Ves- titure of fine, sparse hair. Pronotum 1.2 times as long as wide; sides almost straight and parallel on basal two-thirds, rather broadly rounded in front; anterior margin unarmed; summit in front of middle; anterior slope steep, rather coarsely asperate; posterior areas strongly reticulate, punctures small, shal- low, rather close. Vestiture of fine, short, rather abundant hair. Elytra 1.7 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal two-thirds, broadly rounded behind; disc occupying basal three-fourths; striae not impressed, punctures small, shallow, distinct, in rows, spaced by diameter of a puncture; interstriae three to four times as wide as striae, almost smooth, shining, punctures fine, in definite rows in some specimens, distinctly confused on basal half in others. Declivity steep, convex, general contours as in pubescens; strial punctures large, shallow, distinct, their interior surfaces reticulate-granulate; interstriae only slightly wider than striae, their punctures mostly replaced by minute granules on 400 GREAT BASIN NATURALIST Vol. 35, No. 4 all interstriae, a few larger granules on 1, 3, and lateral areas; posterolateral mar- gin rounded, with an indefinite row of scattered granules. Vestiture of rather abundant, short, fine hair, distinctly longer on margins of declivity. Type Locality. — Stanford University, Palo Alto, California. Type Material. — The female holotype and five female paratopes were taken at the type locality on 15 -25 Til- 1944, by C. D. Duncan. One paratype is from Knight's Landing, Yolo Co., California, 10-M949, at light, by J. R. Fowler. The holotype is in the California Acad- emy of Sciences; three para types are in the Canadian National Collection, two are in my collection, and one is in the U.S. National Museum. Specimens of this species were sent to Bright, Browne, and Schedl in an effort to associate it with a named species from another area, but without success. All agree that its anatomical characters, its current distribution, and the limited period it has been known in California strongly suggest that it was introduced from another area. Xyleborus incultus. n. sp. This species is distinguished from pristis Wood by the larger size, by the more dis- tinctly impressed strial punctures, and by the declivital sculpture as described be- low. Female. — Length 2.3 mm, 2.6 times as long as wide; color dark brown. Frons and pronotum essentially as in micarius Wood except anterior margin of pronotum weakly produced at median line and armed by two larger serrations, punc- tures on posterior areas larger, at least twice as large as in micarius. Elytra 1.5 times as long as wide, 1.3 times as long as j)ronotum; sides almost straight and y)arallel on basal 60 percent, narrowly roundofl behind, a slight emargi- iiation at suture; striae not impressed, {junctures small, distinctly impressed, spaced within a row by one to two diam- eters of a puncture; interstriae almost smooth, punctures fine, less than half as large as those of striae, moderately con- fused on basal third of disc, uni.seriate be- hind. Declivity rather steep, convex; sur- face minutely granular on lower two- thirds, dull; interstriae 1 moderately ele- vated to near apex, armed by 15 or more small, pointed, confused denticles, 2 rather strongly impressed, armed on upper third by 4-6 similar, uniseriate denticles, 3 slightly elevated, armed as in 1 except denticles almost uniseriate, 4-9 each with a row of fine denticles, 3 and 9 join and continue submarginally to apex as a mod- erately strong elevation, its crest armed b}' a row of about four denticles. Vestiture largely confined to sides and declivity; of fine, short strial and slightly longer in- terstrial hair, longest setae about equal in length to width of an interstriae. Type Locality. — Fort Clayton, Canal Zone, Panama. Type Material. — The female holotype was taken at the type locality on 22-Xn- 1963, from a Cecropia branch, by me. The holotype is in my collection. Xyleborus molestulus. n. sp. This species is distinguished from cri- nitulus Wood by the interstrial granules on the disc, by the uniseriate rows of ely- tral setae, and by other characters. Female. — Length 2.0 mm (paratypes 2.0-2.2 mm), 2.2 times as long as wide; color dark brown. Frons as in crinilulus except less strong- ly reticulate, more sparsely punctured. Pronotum about as in crinitulus except anterior margin armed by eight serrations, ]:)osterior areas very finely, shallowly punctured. Elytra 1.2 times as long as wide, 1.3 times as long as pronotum; sides almost straight and parallel on basal two-thirds, broadly rounded behind; striae not im- pressed, ])unctures rather small, shallow, distinct, spaced by one to two diameters of a puncture; interstriae almost smooth, three times as wide as striae, each with a uniseriate row of fine tubercles from base to a])ex. Declivity occupying posterior half of elytral length, moderately steep, broadly convex; sculpture as on disc ex- cept striae 1 feebly impressed; postero- lateral margin acutely, subcrenulately elevated from apex to interstriae 7. Ves- titure of interstrial rows . of long, erect hair from base to apex, alternate setae in each row twice as long as distance be- Dec. 1975 WOOD: AMERICAN SCOLYTIDAE 401 tween rows and alternating with setae half that length on both disc and declivity. Type Locality. — Barro Colorado Is- land, Canal Zone, Panama. Type Material. — The holotype and six para types were taken at the type lo- cality on 27-XII-1963, No. 348, from a tree branch. Four paratypes are from Fort Clayton, Canal Zone, Panama, 22-XII- 1963, No. 363, from a tree branch; three paratypes are from Madden Forest, Canal Zone, Panama, 2-1-1964, 70 m. No. 365, from a tree seedling; and one paratype is from 13 km (8 miles) S El Hato del Vol- can, Chiriqiii, Panama, 7-1-64, tree seed- Hng. All were taken by me. The holotype and paratyjies are in my collection. Xyleborus tristiculus. n. sp. This species is distinguished from rno- lestulus Wood b}" the more slender body, by the larger pronotal punctures, and by the differences on the elytra described below. Female. — Length 2.0 mm (paratypes 1.9-2.1 mm), 2.4 times as long as wide; color brown. Frons about as in molestiilus except imiformly, more strongly reticulate. Pronotum 1.02 times as long as wide; as in rnolestulus except median part of anterior margin weakly produced, four median serrations distinctly larger, and punctures on posterior areas distinctly larger, spaced by at least two to six diam- eters of a puncture; vestiture finer, long- er, more abundant. Elytra 1.3 times as long as wide, 1.3 times as long as pronotum; similar to rnolestulus in outline except more nar- rowly rounded behind; about as in rno- lestulus except discal interstriae not as smooth, interstrial punctures very feebly if at all granulate, declivital striae slightly smaller, interstriae not as smooth, inter- strial tubercles more closely spaced, av- eraging smaller. Vestiture consisting of rather long, fine, semirecumbent strial hair on disc and declivity, and rows of erect interstrial setae on declivity and j:)Osterior half of disc, each seta coarser and distinctly longer than those of striae, distance between rows and between setae within a row about two-thirds length of a seta. Type Locality.— Brazil, 12° 49' S 51° 46' W (not 12° 31' S as given on label). Type Material. — The female holotype and four female paratypes were taken at the tvpe locality 8-XL1968, No. C-47, 22- XL1968, No. D-35, 24-IX-1968, Nos. 170, 172, 2-Xn-1968, No. E-18, all by R. A. Beaver. The holotype is in the British Museum (Natural History), one paratype is in Museo de Zoologia, Universidade de Sao Paulo, and three paratypes are in my col- lection. CORRELATES OF BURROW LOCATION IN BEECHEY GROUND SQUIRRELS Donald H. Owings^ and Mark Borchert' Abstract. — Partial correlation analysis indicates that Beechey ground squirrels show a strong pref- erence for digging burrows under and around large objects, ma>- show a weaker tendency to locate tJieir burrows under the cover of tree canopies, and avoid digging burrows in areas with both tree- canopy and ground cover (stones, logs). These relationships hold for large but not. small burrow sys- tems. The need for unobstructed visual surveillance and an autumn food supply are proposed to be detenninants of tliese preferences. Beeche}' ground squirrels {Spermoph- ilus beecheyi) appear to show preferences for areas with certain characteristics for the location of their burrows. At least three factors have been suggested to af- fect the choice of burrow sites: (1) bur- rows are often constructed under large objects such as stones or logs (Lins- dale 1946: 9); (2) good drainage is said to be iinportant, which is best provided by sloping terrain (Tomich 1962); (3) burrows often seem to be concentrated under the cover of tree canopies (Fitch 1948). One purpose of the research re- ported here was to quantitatively assess the relationship between burrow location and these three independent variables — ground cover (stones, logs), slope of ter- rain, and tree canopy cover. In addition, we felt that the effect of ground and tree cover might interact, or at least be ad- diti^'e, in areas in which these factors physically overlapped. We therefore ad- ded a fourth independent variable (com- mon cover) to the analysis which consis- ted of a measure of the amount of area having ground and tree cover in common. It is probably true, though, that bur- row systems of different sizes may dif- fer in their relationship with these var- iables. Small systems, for instance, may be established by young squirrels who have moved into less than optimal habitat during dispersal, or by adults for refuge from predators when feeding in the open (Fitch 1948; Carl 1971). A second pur- pose of this research was to divide the burrow systems into size categories for separate analysis. Study Area This study w^as done in the Department of Zoology Experimental Wildlife Area ^Departxiient of Psychology, University of California. Davi on the campus of the University of Cali- fornia at Davis (elevation about 15.85 m; 38° 32': N, 121° 47': W). The study plot was located in the 82-m wide origi- nal bed of Putah Creek which is now permanently dry. This area contained numerous trees (principally black wal- nut, Juglans hindsi. and valley oak, Quer- cus lobata), grasses (principally ripgut brome, Brornus rigidus^ with some Italian ryegrass, Lolium multiflorum, and wild oats, Avena fatua), and thistle (Centau- rea solstitialis and Silybum marianum)^ as well as logs, and included most of both sloping sides of this cross-section of the bed. A substantial population of squirrels inhabited this area: 44 were trapped and marked in 0.60 ha in the spring of 1973. This area was being mapped in prepara- tion for behavioral studies. Methods Our procedure was to lay out a grid of 9.14 X 9.14 m squares and to map on graph paper the location of all burrow entrances, the location and size (to scale) of ground cover, and the outer limits of tree canopies for each of 50 of these squares. We derived measures of ground cover area, tree canopy area, common area, and nimiber of burrow entrances from these maps. Slope was measured over uniform sections of the area; new mea- sures were taken wherever significant changes in slope occurred. We assumed that size of burrow sys- tem was positively correlated with num- bers of entrances. We used a portable blower (Steco Model DS-5) and non- toxic smoke bombs (Superior 0.5 min) to assess the number of entrances in a system by blowing smoke into one en- s. California 95010, 402 Dec. 1975 OWINGS. BORCHERT: GROUND SQUIRRELS 403 trance and designating as connected to that entrance all burrows emitting smoke. Results and Discussion In order to minimize the problem of correlated independent variables, we cal- culated partial correlation coefficients. This statistic provides a measure of the correlation of burrows with each inde- pendent variable, while holding the ef- fects of the remaining independent var- iables constant. The results of this analy- sis can be found in Table 1. The first (uppermost) row includes burrow sys- tems of all sizes and suggests that Beech- ey ground squirrels (1) show a strong preference for digging burrows under and around large objects, (2) may show a weaker tendency to locate their burrows under the cover of tree canopies, but (3) avoid digging burrows in areas with both tree-canopy and ground cover. In order to test the hypothesis that the larger systems were the ones most consistently associated with our indepen- dent variables, we successively separated increasing sizes of small systems and analyzed them separately from the larger systems. The results of these analyses are presented in the remaining rows of Table 1 . It is clear that there is little effect upon the relationship between burrows and ground cover of deleting from the analy- sis burrow systems with 1 to 4 entrances. However, deletion of 5-entrance systems from the large-system analysis caused a large drop in the correlation coefficient. Addition of 5-entrance systems to the small-system analysis results in the first significant correlation with ground cover. We conclude that small burrow systems tend not to be associated with ground cover, whereas larger systems (5 or more entrances) do. The negative correlation with common cover and the positive correlation with tree cover followed the same pattern when system size was manipulated. As a result, we conclude that it is principally the larger systems which tend not to be as- sociated with areas having overlapping ground and tree cover, but which are lo- cated under or near tree cover. Our behavioral observations indicate that squirrel concentration was correlated with burrow concentration, and G. L. Hunt (unpubl. data) has quantitatively docmnented this in another location (r= 0.80 - 0.85 between numbers of squirrels and burrow entrances). We assume then that our data indicate microhabitat se- lection by these ground squirrels. We feel that the strongest determinant of this propensity to live in the vicinity of ground cover is the visibility it pro- vides. Hunt (unpubl. data) has gathered considerable support for the hypothesis that Beechey ground squirrels in a much more open area tend to concentrate in locations in which their ground level view (probably of approaching predators) is least obstructed by locating themselves Table 1 Partial correlation coefficients between the numbers of burrow entrances from burrow system of various sizes (dependent variable) and the independent variables (1) slope. (2) ground cov- er, (3) tree cover, and (4) common cover. Size of burrow system is expressed as the number of en- trances to the system. Burrow system sizes Number ( Df entrances Partial correlation coefficients of entrances with included in anal lysis included in analysis Slope Ground cover Tree cover Common cover 1-39 261 -0.067 0.620* 0.231 -0.382* 1 23 0.010 0.012 0.325* -0.220 2-39 238 -0.067 0.613* 0.187 -0.354* 1-2 39 0.106 -0.116 0.051 0.069 3-39 222 -0.089 0.636* 0.225 -0.398* 1-3 54 -0.009 -0.089 0.054 0.013 4-39 207 -0.064 0.629* 0.216 -0.383* 1-4 74 0.109 0.053 0.029 0.043 5-39 187 -0.100 0.591* 0.211 -0.378* 1-5 92 0.079 0.407* 0.134 -0.250* 6-39 169 -0.089 0.484* 0.164 -0.271* 1-6 107 0.025 0.369* 0.122 -0.238* 7-39 154 -0.071 0.486* 0.164 -0.266* 1-7 127 0.027 0.316* 0.246* -0.233 8-39 134 -0.075 0,503* 0.104 -0.265* •■With x no. 24-hr periods operated Table 2. Species diversity of arachnids in three geographic areas. No. species collected Area Spiders Scorpions Solpugids Phalangids Total species collected Nevada Idaho Utah In both Nevada & Utah In both Nevada & Idaho In both Idaho & Utah Common to all areas 138 94 42 40 15 7 6 3 10 9 1 5 4 1 1 1 29 28 6 3 2 2 0 0 2 2 0 0 0 0 0 0 ^Deparlnicnt of Zoolog}-. Brjgliani Youjig University, Prove. Utah S4602. 405 406 GREAT BASIN NATURALIST Vol. 35, No. 4 Table 4. Ecological predominance of arachnid species in three geographic areas. Most Widespread Nevada Idaho Utah Nevada Utah Nevada Idaho Calilena restricta Calilena restricta Psilochorus utahensis Vaejovis confusus Paruroctonus boreus Hemerotrecha californica Eremobates septentrionis Spiders Scorpions Solpugids Most Abundant Psilochorus utahensis Schizocosa avida Psilochorus utahensis Vaejovis confusus Paruroctonus boreus Hemerotrecha californica Eremobates septentrionis Table 3. Arachnid abundance in predominant plant communities. Most Species Fewest Species Spiders Blackbrush Juniper Sagebrush Rabbitl)iusli Juniper Blackbiusli Nevada Idaho Utah Nevada Utah Nevadc Idaho Scorpions Mixed Juniper Blackbrush Sagebrush Solpugids Hopsage Blackbrush Rabbitbrush Goosefoot demonstrated, as between sjnders, scor- pions and solpugids, the occurrence of each group would be expected to differ within the same community. For example, in Nevada the spiders were most common in the blackbrush and mixed communities, whereas the solpugids were least common in the blackbrush and most common in the pinyon- juniper and hopsage- wolf berry where the spiders and scorpions were not abundant. In Idaho the spiders were most abundant in the sagebrush and least in the rabbitbrush, whereas the solpugids were most abundant in the rabbitbrush. In Utah the scor])ions were most abundant in the blackbrush, whereas the sjnders were least abundant there. The relationship of arachnid species to vegetative type is not a direct one, but an interrelationship between a variety of environmental-influencing factors such as cover, food, moisture, temperature, and edaphic conditions. Specific data on these influencing factors are not available for correlation with the results summarized here. Such data are necessary for com- plete ecological interpretations, and such correlative studies are desirable for a better understanding of these interrela- tionships. However, knowledge of species diversity alone in different geographic areas as well as predominant plant com- munities is sufficient to determine eco- logical indicators that may be monitored relative to changes in environmental quality. References Allred. D. M. 1965. Note of phalangids at the Nevada Test Site. Great Basin Nat. 25(1-2): 37-8. . 1968. Ticks of the National Reactor Testing Station. Brigham Young Univ. Sci. 'Bull. Biol. Ser. 10(1): 1-29. . 1969. Spiders of the National Reac- tor Testing Station. Great Basin Nat. 29(2): 105-108. . 1973a. Scorpions of the National Reac- tor Testing Station Idaho. Great Basin Nat. 33(4): 25 1-254. . 1973b. Effects of a nuclear detonation on arthropods at the Nevada Test Site. Brig- ham Young Univ. Sci. Bull.. Biol. Ser. 18(4): 1-20. At.lred. D. M., .\ni) D E. Beck. 1964. Arthro pod associates of plants at the Nevada Test Site. Brigham Young Univ. Sci. Bull.. Biol. Ser. 5(2): 1-16. . 1967. Spiders of the Nevada Test Site. Great Basin Nat. 27(1): 11-25. Allred, D. M., D E. Beck, and C. D. Jorgen- SEN. 1963. Biotic communities of the Ne- vada Test Site. Brigham Young Univ. Sci. Bull., Biol. Ser. 2(2): 1-52. Allred, D. M., and W. J. Gertsch. 1976. Spiders and scorpions from northern Arizona and southern TTtah. Great Basin Nat. (in press) . Allreu. I). M.. AND M. H. Muma. 1971. Sol- pugids of the National Reactor Testing Sta- tion. Idaho. Great Basin Nat. ^1 (3) : 164-168. fli.RTscH. W. J.. .\ND I). M. Allred. 1965. Scor- jiions of tiic Nevada Test Site. Brigham Young 1 luv. Sci. Bull.. Biol. Ser. 6(4): 1-15. Mil ma, M. H. 1963. Solpugida of the Nevada Test Site. Brigham Young TTniv. Sci. Bull.. Biol. Ser. 3(2): 1-15. NOTES ON THE GENUS BOM BY LI US LINNAEUS IN UTAH, WITH KEY AND DESCRIIH IONS OE NEW SPECIES (DIPTERA: BOMBYLIIDAE) D. Elmer Lucilc Maughaii .lohiisoii' Abstract. — A key to the Utah species of Bombylius is piesented. Bonibylius hrxinuiculalux. nigri- ventris. monianus. abdominalis. aeslirus. and auriferoides. new species, and lancifer kanabensis, sub- species new. are described from Utah. Bombylius lassenensis. new name, is proposed to replace the preoccupied B. pallescens .Johnson and Maughan. The first comprehensi\ e pajjer on Utah Bombyliidae (Maughan 1935) lists six species in the genus Bombylius Linnaeus, with the identity of a seventh left in doubt because of the small amount and poor condition of the material available for study. Additional collecting of spec- imens in good condition showed this spe- cies might be B. clio WilHston, but the specimens are not well enough preserved to include in our present key. The alter- nate species, B. fascialis Cresson, has been collected in abundance by us. These results were published in a subsequent paper (Maughan and Johnson 1936). Still further collecting has added much to our knowledge of this genus as it occurs in the northern Great Basin area of Utah, but its status in the southern Great Basin and in the Colorado River drainage sys- tem is still little known. We feel, never- theless, that publication of our available information at this time is worthwhile as it may stimulate future collectors to study this group of flies. In this paper we dis- cuss briefl}' the forms previously known to occur in the state, describe seven new taxa, and present a key for their iden- tification. In addition, we use this oppor- tunity to give a new name to the homo- nym Bombylius pallescens Johnson and Maughan. A large part of the work on this paper was completed eight to ten years ago. Unfortunately, changing circumstances and the illness and death of Elmer John- son slowed progress on its completion. We had completed the key and most of the descriptions prior to that time. In May 1975 I (Lucile) assumed the responsibil- ity, in cooperation with Dr. Vasco M. Tan- ner of Brigham Young University (BYU), of completing this paper so it could be published. I have completed the remaining species descriptions and revised the paper where it seemed advisable. The holotypes, allotypes, and ])aratypes dealt with in this paper are in the Brig- ham Young University entomological type collection. The extensive collection of bee flies which we built u]) over the past 40 years has been transferred to Brigham Young University where it has been carefully curated by Dr. Tanner and his associates and is available for study. Many of the genera represented in the collection in- clude apparently undescribed species which will hopefully be the subject for further research. Much of our collection over the years and all of the types we have named in other genera were pre- viously deposited in the Brigham Young University Life Sciences Museum. Nearly all of the material dealt with in this paper is from Utah, with a few specimens from neighboring states, large- ly collected by Mr. Johnson, some by both of us, and a few by other collectors. Sincere thanks are expressed to those who have aided and given encouragement toward the completion of this paper: to our family; especially to Dr. Tanner for his advice and active assistance in caring for our collection; and to Mrs. Reginald H. Painter, who has been a source of strength and inspiration. Key to the Species of Bombylius Discussed in this Paper Wings with the anterior half brown or blackish, this colored area with a sinuous and sharply defined posterior margin major Linneaus 117 I. Sircct. Salt Lake City, Utah 84103. D. Elmer Johnson died W Scptcmhcr 1973. 407 408 GREAT BASIN NATURALIST Vol. 35, No. 4 Wings with brown area not sharply defined, nearly hyaline, or spotted -- 2 2(1). Wings with five to seven distinct dark spots on veins and crossveins in hyaline portion 3 Wings at most faintly spotted on veins and crossveins other than the r-m crossvein and free part of vein Cui 4 3(2). Wings with dark spot near distal end of vein R. .,; heavily tomentose, female with golden tomentum on front; usually in the mountains or foothills albicapiUus albicapillus Loew Vein Ro , without dark spot near its end; sparsely tomentose, female with pale yellowish tomentum on front; usually found in the foothills and valley floors heximaculatus n. sp. 4(2). Face prominent, facial pile wholly black , 5 Face pollinose, facial pile at least in part pale 11 5(4). Pile of dorsum mostly pale yellow to golden; legs except bases of fe- mora and spots on knees yellow; female with large trapezoidal area on front shining; both sexes with median line of pale tomen- tum on abdominal dorsum; halteres pale brown to yellow fascialis Cresson Pile of dorsum not as above; usually with a more or less prominent line of brown or black hairs extending from base of wings forward onto occiput --- 6 6(5). Males - .- 7 Females - -. 9 7(6). Pile of venter black, that of dorsum pale gray; no tufts of black pile on sides of abdomen visible from above; relatively large species (9-12 mm); frequents the foothills nigriventris n. sp. Pile of at least first three ventral abdominal segments pale; tufts of black pile on sides of abdomen visible from above 8 8(7). Pale pile of dorsum dark yellowish gray, always several on posterior callosities black; line of dark hair from wing onto occiput usually broad and prominent; prominent tufts of black pile on hind angles of second and third terga; band of black ]Hle on front of two and often on three also; frequents mountains .- . lancifer lancifer Osten Sacken Pale pile of dorsum nearly white, on posterior callosities may be oc- casional black hairs; line of dark hairs from base of wing onto occi- put thin and inconspicuous; tufts on sides of terga prominent on anterior angles of third segment only, on fourth visible from ven- ter only - lancifer kanahensis n. ssp. 9(6). Relatively large species (9-12 mm); lower frontal patch almost ob- solete, the s])arse scales mostly pale brown; ]iatches of dense white or nearly white tomentum on second and following segments form a i)rominent median line on abdominal dorsum; tomentum of remainder of dorsum hairlike and relati\(4y sparse; frequents foothills -- nigriventris n. sp. Smaller species (7-10 mm); lower frontal patch ]:»rominent, of black hairlike scales; abdomen with crossbands of more or less dense pale scales on posterior half of second and following segments, median line of scales evident but not prominent 10 10(9). Pale hairs on dorsum mostly dark grayish yellow, many black hairs on mesonotal disc and posterior callosities; some of pale tomentum of abdominal dorsum noticeably pale brown; dorsum appears Dec. 1975 johnson. johnson: utah bomhyliidae 409 banded black and pale due to sparse dark tonientum in bands of black pile on anterior part of terga . lancifer laucifcr Osten Sacken Pale hairs of dorsum nearly white, few if an>" black hairs on nieso- notal disc and posterior callosities; tomentuni of ])Osterior half of second and following abdominal segments whitish gray, that of an- terior half of second and third segments tinged with tan, covering segments lancifer kanabensis, n. ssp. 11(4). Lower frontal patch white to grayish 12 Lower frontal jjatch yellow to golden 14 12(11). Facial pile wholly white; base of wings brownish; ])re- and postalar and scutellar bristles prominent, brown; female with many hea\ y bristles on upper mesopleura nionUinus n. sj). At least some black hairs below antennae and along facial orbits; base of wings blackish 13 13(12). Most of pile of face black, only that on oral margin yellowish; entire frontal triangle of male covered with dense appressed white scales; on the female these scales are found only between antennae and eyes; dark of wings fills anteroproximal one-third or more of wing; dark spots present on r-m crossvein and free jiart of vein Cui .... metopium Osten Sacken Most of pile yellow to white, only a narrow band of hairs below an- tennae and on facial orbits black; middle of frontal triangle of male devoid of dense appressed tomentum; only anteroproximal fifth or less of wing dark, no brown clouds on veins or cross veins; prom- inent crossbands of black pile on abdominal dorsum abdominalis n. sp. 14(11). At most bases of femora dark, rest of legs reddish tan 15 Legs black 16 15(14). Pile at base of wings golden; dark anterobasal part of wing with dis- tinctly reddish brown cast; male genitalia red, slightly darker than femora; late summer species (August) aestivus n. sp. Pile at base of wings grayish yellow to white, male genitalia brown to black 17 16(14). Abundant erect pile of front, first two antennal segments and wide band below lower frontal patch black and very long in both sexes, female with mixed pale and black on front; pale gray species, fe- male with nearly completely hyaline wings lassenensis new name No erect pile on front in male; pile of face, appressed hairlike tomen- tum on front of both sexes, and erect pile of female all same pale color; black pile on antennae and narrow band below lower frontal patch . . aurifer pendens Cole 17(15). Halteres yellow; third antennal segment of female widest at or be- yond middle, with sides nearly parallel for three-fourths of its length, then tapering abruptly to end; pile of front and ocellar tu- bercle of female wholly yellow; dark color in wing of male does not extend beyond tip of vein Ri aurifer aurifer Osten Sacken Halteres various shades of brown; third antennal segment of female widest before the middle, not evenly tapering toward end; pile of front and ocellar tubercle of female mixed black and yellow; shin- ing wliite scales form complete band above antennae between eyes; dark color in wing of male more or less fills all of cell Ri auriferoides n. sp. 410 GREAT BASIN NATURALIST Vol. 35, No. 4 Bombylius heximaculatus, ii. sp. Wings with five or six distinct black spots in the hyaline area or bordering the dark color. Legs mostly red. 9-12 mm in length. Female. — Black. Head gray pollinose, occiput densely so; lower frontal patch black, very sparse; tomentum of front sparse, golden; pile of antennae, ocellar tubercle, front, and face black, a few pale hairs around the oral opening; pile of occiput white or nearly so, a patch of brown to black near lower hind angle of eye; segments 1 and 2 of antennae about same width, 2 one-third as long as 1 ; segment 3 somewhat wider, about one- fourth longer than first two combined, widest at about first third, sides of apical third about parallel; proboscis about five- sixths as long as head and body combined. Mesonotal disc velvety black; pile long, dense on anterior one-third and sides; sparse, gray mixed with a little black on center and scutellum; with some sparse decumbent grayish yellow tomentum; pleura densely gray pollinose, lavender glints in some lights; a dense patch of mixed black, brown, and gray pile on upper mesopleura, just before wings, con- tinuous with dark hairs of lower occiput; most of rest of pleura naked or nearly so, the sparse pile mostly pale, a few black hairs intermixed. Wings with anteroproximal one-third smoky, remainder gray hyaline; dark spots on veins at r-m crossvein, base of R,, at junction of Mj + R,-„ at free part of M2, at cubital end of median crossvein, and at free part of Cui. Basicostal pile black, a few pale hairs on sides of costa. Halteres brown. Legs dark red, hind femora yellowish except knees, others brownish red, darker basally, and tarsi dark; pile of fore coxae pale yellowish gray, of middle coxae mixed black and pale, and on hind ones only a few black in the gray; of rest of legs black; bristles black; tomentum pale, like the pile. Abdomen faintly gray pollinose; pile dense on sides, lateral fringe mostly almost white on sides of first, anterior half of second, fifth, and following seg- ments, considerable black pile mixed in on sides of posterior half of second and on third and fourth; pile of dorsum most- ly short jjale yellowish gray, with many long black hairs on hind margins of sec- ond and following segments. Most of pale pile of venter black, a few pale hairs on first segTnent; sparse tomentum pale yellow. Type. — Female holotype: Alpine, Utah Co., Utah, 20 May 1955' (L. M. Johnson). Paratypes: 4 $ same data. Several males were collected but be- came damaged so no description will be attempted at this time. The males are paler, with less black and pale pile on ab- dominal dorsum and more black pile on sides. Pile of venter is yellowish gray. Females are more tomentose. There is some variation in both sexes in the amount of dark pile at hind angles of eye and on mesopleura between the wing and head. Bombylius nigriventris. n. sp. Black. Anteroproximal third of wings blackish, gradually evanescent behind. Most of pile pale yellowish gray, that of abdominal venter mostly black. Length 6-12 mm exclusive of proboscis. Proboscis long, three-fourths as long as to fully as long as body and head. Male. — Head black, face except upper and outer margin brown. Front and dark margin of face faintly grayish pollinose. Occiput and genae yellowish gray polli- nose. Pile of face, front, antennae, a row along u])per orbits on occiput and ocellar tubercle black. That of rest of head pale, faintly yellowish gray, a patch of bro\vn hair on lower occiput. Lower frontal patch of short, black hairlike scales. An- tennae cylindrical, segment 1 a little more than twice as long as broad, segment 2 same width, about as long as broad, seg- ment 3 somewhat narrower, slightly more than twice as long as first two com- bined, widest at its proximal one-fourth, sides of distal two-thirds almost parallel; first segment long pilose, segment 2 very short fine pilose, segment 3 with a few short fine hairs on its u])per side on distal half. Palpi black, black haired. Pile long and dense on anterior third, rolativel}- short and sparse on remainder of mesonotum, very pale yellowish gray, some dark hairs scattered among pale in center of mesonotal disc. Pile of pleura dense above, more sparse below, lower Dec. 1975 JOHNSON. JOHNSON: UTAH HOMBYLHDAE 411 pteropleura bare, mostly nearly white, that on sterno- and hypopleura slighth yellowish. A band of mixed brown and pale hairs extending from the base of wing forward to join with patch of brown hairs on lower occi])ut. Pile of scutellum same color, occasional darker hairs along hind margin. Dark of wings fills alula, basal third of axillary, two-thirds of anal cell, base of fourth posterior to base of discal past r-m cross vein to tip of subcostal cell. Re- mainder of wing gray hyaline, veins dark brown. Basicostal pile mostly black, longer and pale at base, a few pale hairlike scales beneath. Halteres brown. Coxae and base of femora dark brown, remainder of legs reddish brown; bristles black; pile of coxae pale, many dark hairs intermixed on middle pair, that on femora mixed pale and dark brown; tomentum of legs pale yellowish gray, a line of brown scales on facies of femora. Abdomen blue black; pile of dorsum long, dense, pale faintly yellowish gray, lateral fringe on second and third and a few hairs on hind margins of second and following segments black, somewhat more numerous on fifth and sixth terga. Pile of first two sternites pale, of rest of venter and on genitalia black, a very few pale yellowish hairlike scales intermixed. Female.^ — Less pilose and more tomen- tose than male. Vertex, front, and dark upper and outer margin of face brown pollinose. Antennal segment 1 about two and one-half times as long as wide; seg- ment 2 same width, as long as wide; seg- ment 3 widest at proximal third, thence tapering evenly to short onion-shaped style. Tomentum of front yellow, a few brownish scales near orbit in lower fron- tal patch; that of occiput nearly white. Pile of head as in male, except that there are a few pale hairs on front along orbits. Mesonotum and scutellum with moder- ately dense, pale grayish yellow curly appressed hairlike tomentum. The band of darker pile extending from base of wing to occiput is yellowish brown, in- termixed with a few black hairs. Wings paler than in male, basicosta has more pale tomentum. Pile of abdomen much coarser than in male, pale hairs mostly more yellow; many more black hairs, that of fifth and sixth terga mostly black; black hairs of lateral fringe of second and third terga confined to hind angles; lateral fringe of fourth and fifth segments white. Dense median patches of faintly yellow to white tomentum on second and follow- ing terga form a prominent line down middle of abdominal dorsum. Pale hairlike tomentum on hind margins of first two and in median spots on following ster- nites; rest of sparse tomentum of venter black. Types. — Male holotype: Alpine, Utah Co., Utah 23-V-1955 (D. E. Johnson), female allotype, topotypical, 20-V-1955 (D. E. Johnson), paratypes 13cf 9, topo- typical, ll-V-1954, 16-V-1954, 18-V-1954, 22-V-1955, 19-VI-1955, (D. E. and L. M. Johnson); 2cf 1 5 Dry Canyon, Salt Lake Co., Utah; Id 1 9 Fort Douglas, Salt Lake Co., Utah, lO-V-1933, 6-V-1940 (W. Anderson); 2^ Oak Creek Canyon, Mil- lard Co., Utah, 24-V-1957 (D. E. John- son); 1 cT Mt. Carmel Junction, Kane Co., Utah, 21-V-1957 (D. E. Johnson). In addition to the variation in size, there is some variation in the amount of black pile on the mesonotal disc and hind margins of the abdominal terga in the males, and in the amount and color of tomentum on the abdominal dorsum of the females. In both sexes there is some variation in the amount and color of the dark pile which forms the conspicuous dark line from the base of the wrings for- ward onto the lower occiput. This species has the same type of face and wings as lancifet\ but can readil}^ be separated from that species by its larger size, by the absence in both sexes of the crossbands of black pile found on the hind margins of the second and third terga of lancifer, by the presence of most- ly clark vestiture on the abdominal venter, where lancifer is mostly pale, and by the presence in the female of the prominent median line of white tomentum on the abdominal dorsum. Moreover, lancifer is a species of the mountains, while nigri- ventris appears to be confined to the foot- hills and valley edges, and is on the wing about a month earlier in the season. While hovering, the females of this species produce a sound somewhat lower in pitch than the hum of a honeybee, and of a different quality, being more of a buzz than a hum. The hovering males produce a hum considerably higher than 412 GREAT BASIN NATURALIST Vol. 35, No. 4 that of the bee. This high-pitched hum was audible to us for a distance of forty or fifty feet, and was frequently used as a means of locating specimens. Frequent- ly males were found hovering or darting about some 7 to 12 or so feet above a given small area. We found our first female, as well as several subsequent ones, resting on the ground beneath such a hovering male. We did, indeed, use the males as guides to the females, collecting first the female and then the male. Another spe- cies of Bombylius heximaculatus n. sp., was present in the same habitat as nigri- ventris, and was found in far greater numbers. There was sufficient difference in the sounds produced by the two species to readily recognize and locate nigriven- tris specimens b}-^ the lower pitch. The types were taken about one mile west of the village of Alpine, at an eleva- tion of 5,340 feet. The site is a narrow unplowed strip alongside a road through agricultural lands. The whole area is foot- hills of the Wasatch Mountains above the highest level of Pleistocene Lake Bonne- ville. With the exception of Mt. Carmel .function, the other collection sites are also foothill areas of the Wasatch, at ele- vations between 5,300 and 6,000 feet. The elevation at Mt. Carmel Junction is listed as 5,241 feet. Although other species were available, the only flowers at which we saw these beeflies feeding were those of Phlox longi- folia. Wherever we have found nigriven- tris flying we have also found Phlox blooming. Bomhylius lancifer kanahensis. n. ssp. Like lancifer O.S. except it has far few- er black hairs on thorax and abdomen. Pale pile of dorsum glistens in reflected light, making the fly appear to shine. Length 8-10 mm. Male. — Head black, face shining brown, face and front from some angles lightly brownish gray pollinose, occiput and chin gray i)ollinose; pile of ocellar tubercle, front, face, antennae, and a few hairs and scales in a patch on occiput near lower corner of eye brown, with a few- black intermixed; lower frontal patch black, sparse hairlikc tomentum of front ])ale grayish yellow; pile of chin white, of occiput grayish yellow (same color as that of adjoining thorax). Antennae slender; segment 1 about three times as long as wide; segment 2 same width as first, about as long as wide; segment 3 not quite as wide as first two, its sides nearly parallel, not quite twice as long as first two combined, its style evenly long conical. Proboscis black, about four-fifths as long as head and body combined. Thorax black, bluish reflections in some lights, pleura thinly gray pollinose over shining surface; pile fairly long and dense, very pale yellowish gray, nearly white on pleura, a patch of dark brown with a few black hairs before the wing extending onto occiput; macrochaetae not strongly developed, nearly same color as pile. Two black hairs on left posterior callus, none on others. Wings brownish hyaline, anteroproxi- mal third brownish, color evanescent be- hind; veins brown; basicostal pile black, a few white hairs near base shading to brown along heavy vein; pile of alulae long, very pale yellowish. Halteres brown. Legs yellow, knees, tarsi, and bases of femora brown, first two pairs broadly so; pile of coxae nearly white, of remain- der of legs mostly dark brown to black; tomentum nearly white; bristles black. Abdomen blue black; pile of dorsum long and dense, faintly yellowish (paler than thorax), tufts on hind angles of second, on hind angles of third when viewed from below, and a few hairs on hind margins of second and succeeding .segments black; tomentum nearly white, a sj^arse jiatch in center of second and fairly dense crossbands on ])osterior third of tiiird to fifth segments, covering last segments. Pile of venter mostly white, a few black hairs on hind margins of last three segments; sparse tomentum whitish. Genitalia dull black, shading to brownish at tips. Female. — Much like the male. Less pilose, but much more tomentose than male. Front brown pollinose; pile black; tomentum golden, a few of the pale scales mixed into the lower frontal patch, especially on inner end. Face with a few scattered yellow hairlike scales scattered among pile. Sparse tomentum of occiput pale yellowish, a few black hairs mixed with pale buff pile near vertex, and a few black and dark buff hairs near lower inner corner of eye. Onl}^ a few black Dec. 1975 JOHNSON, JOHNSON: UTAH BOMBYLnDAE 413 hairs in dark band before the wing. Pos- terior two-thirds of mesonotuni and scutel- lum with sparse pale yellow curly tomen- tum. Wings only slightly paler than in male. Dark ])art of femora less extensive than in male. Bristly black hairs in tufts on hind angles of second and third terga similar to male, relatively more bristly black hairs on hind margins of second and succeeding terga. Second sparsely except in middle, and succeeding terga densely tomentose on posterior half of segments and in centers, forming a distinct nearly white median line; anterior half of segment 1 almost bare of tomentum, front of 3 and 4 with sparse tomentum. Most of pile on first three ventral segments white, on succeeding segments mostly black; tomen- tum pale yellow to white. Types. — Male holotype and female allotype: Six miles north of Kanab, Kane Co., Utah, 5-VI-1965 (D. E. & L. M. Johnson). Paratypes: 21 cJ", 12 9 same data. The type locality, six road miles north of Kanab, on U.S. Highway 89, is in the red sandstone hills of the Vermillion Cliffs area. The flies were on a red sandy sidehill among scattered Juniperus osteo- carpa and Artemesia tridentata. Asso- ciated with this species were males of Lordotus apicula Coq. The males were hovering and the females feeding be- tween 8:30 and 10:00 a.m. This series of flies is quite uniform. There is, however, some variation in the amount of black pile on the thorax and abdomen. But even the darkest specimens have very few black hairs. This subspe- cies is readily distinguishable from the nominal subspecies by this scarcity of black pile, and by the paler color of the pale pile, which in this subspecies is nearly white. On females there may be a few brown to black hairs on occipital or- bits near the vertex; tomentum of front may be pale yellow, on the occiput and the thorax in front of the wing there may be no dark pile; there may be no brown on the femora. Bombylius major Linnaeus This is the earliest bee fly on the wing in the spring in our area.. It appears as early as mid-March on the edges of the valley floors and in the foothills. We have not done any early collecting in the southern part of the state and have not seen this species in the collections of others from there. We have seen it in col- lections from Arizona. Bornhjiius nu'topiutri Osten Sacken This small species appears shortly after major around the edges of the valleys of the northern part of the state. As the seas- on progresses, metopium^ like se\eral others of its congeners, moves up into the canyons and onto the foothills. We have found it common in only a few places, but it is so inconspicuous that we may easily have overlooked it at other sites where we were not specifically seeking it. The shin- ing white tomentum on the male com- pletely obscures the front; on the female it is found only at the sides of the anten- nae. The specimens have a gray appear- ance. Bombylius lancifer lancifer Osten Sacken We have collected this species in the mountains abo^e 7,000 feet elevation from May into July. We have found them in the canyons and on northern slopes of practically every mountain range in which we ha^e sought them in the northern part of Utah. We do not know how far south they may be found. Specimens have been examined from Utah, Oregon, and California. Bombylius montanus. n. sp. Facial pile and lower frontal patch wholly white; macrochaetae well devel- oped. Length 9-11 mm. Male. — Head dark brown, densely gray pollinose, face and oral margin yel- lowish; pile white, a line of dark brown to black hairs on each side of bare pol- linose area on frontal triangle between antennae and vertex, some on ocellar tu- bercle, and upper and outer side of first antennal segment; tomentum of occiput and lower frontal patch at sides above antennae subshining white, a few brown decumbent hairlike scales at junction of inner orbits on upper front. Antennal segment 1 about twice as long as wide; segment 2 about as wide as first, as long as wide; segment 3 about as wide as first 414 GREAT BASIN NATURALIST Vol. 35, No. 4 two, about twice as long as first two com- bined, widest at about the middle, tapering evenly thence to blunt tip; style evenly conical. Proboscis brown, about two- thirds as long as body and head; palpi brown, pile yellow. Thorax brown, dorsum faintly brown- ish, pleura densely gray pollinose; pile very dense, pale gray and reddish brown on anterior one-third of mesonotum, sparse on posterior two-thirds except along sides; posterior callosities have dense tufts of pile, reddish at base, gray at ends, bordered next to mesonotum with shorter almost black pile. Sparse pile of mesono- tum has pale gray pile with hairlike gray tomentum beneath it in two indis- tinct longitudinal stripes, with a few black hairs intermixed in the darker contrasting brownish pile on the rest of the disc. Pile and tomentum of scutellum like that of posterior mesonotum; most of macrochae- tae brown to black, well developed, three or four on notopleura, a number on pos- terior callosities and along hind margin of scutellum. Pleura with pile moderately dense, yellowish gray on upper half, less dense and white on lower half. Anteroproximal half of wing brown, somewhat darker along veins, the color evanescent behind, becoming gray hya- line; veins brown, basicostal pile black, a few white hairs at extreme base of wings; pile of alulae dense, long, white. Halteres brown. Legs yellow, knees and tarsi brown; pile and tomentum pale yellowish to white, bristles black. Abdomen grayish brown, pollinosity almost same color; pile of first three terga pale brownish yellow on sides, nearly white in middle; dark brown to black bristly hairs form single rows on hind margins on outer third of first and across second and third segments and form prominent tufts on sides of second segment, with small tufts on sides of third; pile of rest of abdominal dorsum white, a few black hairs on hind margin of sixth segment; tomentum white, sparse on second and third, moderately dense on succeeding segments, more concentrated on posterior half and middle of second, third, and fourth; pile of venter mostly white, a few black hairs on terminal three segments; sparse tomentum pale yellow. Genitalia dark brown and reddish, pos- terior angles of ninth sternite produced into sharply pointed spinelike projections; pile yellow. Female. — Much like male but less pilose and much more tomentose. Wings almost completely hyaline. Pile of occi- put yellowish, that of front sparse, mixed black and white. Front and face densely pale gray pollinose. Dark pile of thorax paler than in male. Halteres paler than in male. Tomentum on abdomen nearly white, appearing pale brownish on mid- dles of second, third, and fourth terga be- cause less dense, rest of tomentum dense; pile mostly white, that on hind margins of segments two to four black and bristly, brown to pale on fifth and following seg- ments, brown tufts on sides of two and three. Pile of first three sterna mostly white, rest mostly black and brown, bris- tly on hind margins of segments. Types. — Male holotvpe: Aspen Grove, Utah Co., Utah, 19-VIU1953 (D. E. John- son) . Female allotype: Pebble Basin, Ban- nock Co., Idaho, 24-VII-1948 (D. E. Johnson). Paratypes: 1 cT same data as holotype; 1 d" Strawberry Valley, Wa- satch Co., Utah, 1()-VII-1969 (on flax and cinquefoil). In general appearance this species re- sembles eboreus Painter except that the dark of the wings is distinctly brown rather than blackish. However, the strong development of the macrochaetae sets this species apart from all of its congeners in our area. We have seen specimens from Pasadena, California, which may be this species, but more material will be neces- sary to be certain. Apparent variation shows pile of posterior callosities all pale gray. Slight rubbing destroys distinctive lines on mesonotum. Macrochaetae may be pale. The holotvpe and paratype specimens were collected on the Mt. Tim])anogos trail about a half mile from Aspen Grove. There were three males and one female alternately hovering and darting about the site, but only two of the males could be ca[)tured. Bomhylius ahdotninalis. n. sp. Black, pale pile of dorsum mostly yel- lowish gray. Length 9-12 mm exclusive of proboscis. Male. — Head black, densely gray pol- linose, oral margin vellow shading to Dec. 1975 JOHNSON. JOHNSON: UTAH BOMBYLHDAE 415 black below antennae. Appressed shining white tomentuni above antennae leaves center pollinose strip bare except for a ver}' few long white scales. Black erect pile on bare area next to white tomentum. Several rows of black pile below antennae and on facial orbits, dense yellowish white pile on lower face and oral margin. Tuft of black pile on ocellar tubercle, tomen- tum of occiput white, pile yellowish gray abo^•e, white when light reflects off it. Antennae with first two segments quite straight-sided, slightly wider than third; first more than twice as long as second, third about one and one-half times the length of first tw^o combined. Black pile on first two segments. Proboscis about three-fourths the length of body and head combined, black, as are palpi. The latter about one and one-half times as long as the width of the base of the proboscis, pile black. Thorax and abdomen black with pale pile from third tergite forward 3'ellowish gray, shining almost white in some lights, that of fourth and following tergites white. Dorsum of thorax wdth pile dense at sides and front third of mesonotum and on scutellum; shorter on rear two-thirds, sparse, with a few black hairs intermixed, and with scattered decumbent fine golden tomentum. Macrochetae yellowish. Sparse black pile on scutellum and post allar cal- losities. Wings hyaline except anterobasal fourth cloudy reddish brown, evanescent behind. Basicostal pile white, with black on front edge. Pile of alulae white, dense and long. Halteres brownish with part of knobs paler. Legs have black femora, tarsi dark yel- lowish brown. Vestiture white, except spines are black. Dorsum of abdomen with pale pile bushy, black pile in prominent band on rear of second tergite, narrower band on third and some black pile on rear of other segments. All pile slightly shorter on sides of second and third tergites. All to- mentum of abdomen is white; scant on second and third tergites, more dense in center of each segment, giving the effect of a stripe, wider in the white pile, very narrow in the black pile. Fourth and fol- lowing segments with dense tomentum. Venter of second and third segments wjth long white pile and dense long hairlike tomentum, following segments with black and white pile scant, and with short white tomentum also not dense, but forming a stripe at center of last three segments. Genitalia very dark, yet not black in color. Female. — Much like the male. Tufts of black pile on sides of wide upper front and a few fine golden scales on front. Mesonotum with disc back of long pile covered with decumbent fine golden to- mentum, more dense on scutellum, where it is arranged with the ends pointing to- ward center. Abdomnial dorsum much more white tomentose, longitudinal stripe more than twice as wide as on male and more continuous, with diamond-shaped effect on second and third segments. Much less white pile than male on dorsum, sides about same. Venter less pilose, heavily tomentose on second and third sternites, following ones with short tomentum forming a stripe on fourth to sixth, cover- ing seventh. Type with front femora dark, other femora partly so, and tarsi paler than male. Types. — Male holotype and female al- lotype: Stansbur}' Mountain, Tooele Co., Utah, 23-V-1969 (D. E. Johnson). Para- types: 236^, 39, topotypical, 23-V-1967, 18-V-1969, 23-V-1969; 8d, Hickman Canyon, Stansbury Mountain, Tooele Co., Utah, 2-VI-1957; 6d, H ? Clover Creek, Tooele Co., Utah, 9-VI-1957; 16cf , Alpine, Utah Co., Utah, 18-V-1954, 24-V-1969 (D. E. and L. M. Johnson); Id Little Mt., Salt Lake Co., Utah, 17-VL1968. Other specimens have been collected from Lark, Salt Lake Co., Utah, 9-VL1953. In addition to variations in size, there may be little black pile on the thorax; knobs of halteres are sometimes mostly dark; three segment stripe of white to- mentum on posterior sternites of male faint or not ])resent; some females have completely pale yellowish brown femora. The description of this species was in- complete at the time of Mr. Johnson's death. lie did not record any information about the ecology of the Stansbury Mountains, Hickman Canyon, or Clover Creek areas, which are all within a 10- mile distance on the east side of the Ona- qui Mountains. However, the Alpine spec- imens were collected in the same strip be- tw^een cultivated fields as nigriventris in the foothills of the Wasatch Mountains, 416 GREAT BASIN NATURALIST Vol. 35, No. 4 and all the other areas are wild habitat at similar altitudes and with similar vege- tation. Bombylius aestivus^ n. sp. Similar to aurifer O.S. but has all red femora, and pile of face, front, and pleura pale tawny instead of nearly white. Length 5-7% nun. Male. — Dark brown; yellowish gray pollinose on front, face, first and second antennal segments, mesonotum, pleura, first tergum, and venter of abdomen. Antennae black, segment 1 about twice as long as wide; segment 2 about same width, almost round; segment 3 almost twice as long as other two together, not quite as wide, almost parallel sided. Pro- boscis black, about two and one-half times as long as head; palpi yellow at base, black about half of length, somewhat long- er than width of proboscis. Pile long and shaggy, moderately dense on face, occiput, thorax, and sides of ab- domen; color of pile shining pale tawny, somewhat paler on pleura and abdominal venter, with black pile found in a row below lower frontal patch, a few on an- tennal segments 1 and 2, and sparse tufts of hairs on anterior lateral angles of third and fourth terga. Tomentum hairlike, shining, of about the same color as pile, moderately dense on thorax and abdomen, a dense appressed patch above each an- tenna. Genitalia pale red. Legs red, knees narrowly black, tarsi darkening distally; pile and tomentum as on remainder of body, bristles black. Wings gray hyaline, brown on anterior proximal one-third, color evanescent be- hind and distally; pile on base of wings and alulae pale tawny, long and shaggy. Halteres brown, knobs pale. Female. — Very much like the male. Fewer black hairs on face and on anterior angles of abdominal segments than in male. Abdomen relatively broader, more densely tomentose; front with pile and to- mentum as in remainder of body. Types. — Holotype male: Provo Can- yon, Wasatch Co., Utah, 14-Vin-1966 (D. E. .Johnson). Allotype female: Tryol Lake, Uintah Mts., Summit Co., Utah (no other information, but collected before 1931). Paratypes: 4cf, 1? topotypical with holotype. Some damaged specimens in the col- lection show some variation in the amount of black pile on the face, on antennal seg- ment 2 and on the abdomen. Some speci- mens, particularly females, have only a very few black hairs, or even none at all in these places. This species may be readily separated from aurifer aurifer O.S. which may occur in the same area by the red femora and by the darker color of the pile of the face, occiput, and pleura, which in aurifer aurifer are nearly white. Cole's aurifer pendens has only partially red femora. Whereas the other species of Bombylius in this area are flies of the spring and very early summer, this species is apparently not on the wing until much later in the season and is not in evidence until long after the others have disappeared from even the high country that seems to be home for aestivus. The type locality is in the Wasatch Momitains about one mile southeast of the Deer Creek Resenoir in Provo Can- yon. The flies were collected on a north- facing slope in open areas between patches of scrub oak, Quercus gambeli^ and choke cherries, Prunus melanocarpa. They were feeding on Helianthus sp. and Aster sp. The elevation is between 7,000 and 8,000 feet above sea level. The allotype has only a locality label, but Tryol Lake is a similar ecological habitat in the Uintah Mts. Other species of bee flies common in the area at the same time include Poccil- anthrax signatipennis (Cole), P. sackenii (Coq.), P. willistoni (Coq.), Sparnopolius coloradensis Grote, Villa edititoides Paint- er, Aphoebantus mormon Melander, and Lor dolus gibbus Loew. Bombylius lassenensis, new name At the time Bombylius pallescens John- son and Maughan (1935) was described, we were unaware that Hesse (1938:226) had used the name for an African spe- cies. Since our species is a junior homo- nym of Bombylius pallescens Hesse, the new name Bombylius lassenensis is pro- ])osed as a replacement. This species, col- lected in Lassen Co., California, is in- cluded in the key in this paper, even fhouerh it is not from Utah. Dec. 1975 JOHNSON, JOHNSON: UTAH BOMBYLHDAE 417 Bombylius auriferoides, n. sp. Black, pile mostly pale yellowish gray; similar to aurifer 0. S. but with brown halteres. 4.5-10 mm in length exclusive of proboscis. Male.^ — Head black, gray pollinose; pale pile faintly yellowish gray, that of ocellar tubercle, antennae, and outer face mostly black; lower frontal patch covered by dense, appressed, shining, nearly white scales which nearly completely obscure the front. Antennal segment 1 about twice as long as wide; segment 2 same width as first, about as long; segment 3 about one and one-half times as long as first two combined, about as wide at its widest part as first two, widest at its basal third; pile of segment 2 mostly pale, that of segment 1 black. Proboscis about half as long as head and body combined. Thorax faintly grayish pollinose on the mesonotal disc, densely so on pleura; pile long and dense, that on pleura nearly white; short, moderately sparse shining hairlike scales on mesonotum and scutel- lum. Wings brown on anteroproximal half, gray hyaline behind, color extends to distal end of cell R,, evanescent behind; veins brown, darker distally; pile of squa- mae and extreme base of wing pale, rest of pile of costa black. Halteres brown. Legs black, dense tomentum and most of pile pale, a few dark hairs beneath on first and second femora; bristles black. Abdomen moderately densely pilose, three tufts of black hairs at anterior an- gles of third, fourth, and fifth terga; to- mentum of abdomen of short, curly, ap- pressed hairlike scales, same color as pile. Genitalia brown, members yellowish dis- tally. Female. — Much like male, somewhat less pilose and more tomentose. Upper half of front and vertex brownish polli- nose, very sparsely tomentose; lower half of front gray pollinose and densely pale yellow tomentose in an uninterrupted band between eyes; pile of ocellar tu- bercle, front, and antennae with black and pale yellow pile, pale pile being mostly on posterior part of tubercle, along orbits, and on upper part of an- tennae. Wings somewhat paler than in male. Fewer black hairs at sides of ab- domen. Types. — Male holotype and female allotype: Mt. Timpanogos, Utah Co., Utah, elevation 8,600 ft., 30-VI-1957 (D. E. Johnson). Paratypes: Idaho: Ban- nock Co., 1 cf Lava Hot Springs, 23-VL 1935. Utah: IScT, same data as types; Utah Co., Utah: 37cr, 18? American Fork Canyon; IcT, 1 ? Prove ; 2cf Aspen Grove; 8cf, 2$ Pay son Canyon. Salt Lake Co., Utah: 1 J" Lambs Canyon; 4cr, 2 $ Parley's Canyon; 28 cf, 7 5 L'ttle Mountain. All sj)ecimens collected by D. E. .Johnson. In addition to the remarkable range in size, there is considerable variation in some other characters. The color of the halteres varies from rather pale to Aery dark brown, with the females usually being somewhat paler than males. The color of pile varies from nearly white to distinctly yellowish, particularly on the dorsum of some s])ecimens. The smaller specimens are usually paler than the larg- er. The very early season flies are usually paler in all respects than later season specimens. The amount of black pile on the sides of the abdomen varies from three distinct tufts in some specimens to only a few inconspicuous black hairs in others. Some males have no black hairs on either the ocellar tubercle or antennae, while others have a few on either or both. There is variation in the number of black hairs on the antennae, front, and ocellar tuber- cle of the females; but always there are at least same black hairs at all three of these locations, and always there are some pale hairs along the orbits on the lower front. There is some \ariation in the relative length of the proboscis. The third anten- nal segment varies somewhat in shape in drying, but always, in both sexes, there is some constriction between the base of the segment and its widest point, which is nearly always at the basal third. Also, this segment is always narrowest just be- fore the tip, flaring more or less to the end where the onion-shaped style is at- tached. This species and aurifer 0. S. are so similar in general appearance that it was not until we were studying a long series of specimens of what we assumed were aurifer, from a single locality to determine the intraspecific variation, that we re- alized we were dealing with more than one taxon, and that the small differences we had noted consistently separated the 418 GREAT BASIN NATURALIST Vol. 35, No. 4 whole into two distinct groups. Inasmuch as the two forms have sympatric distri- bution they can only be considered as distinct species. The following characters may be used to separate them: In aurifer the halteres are always pale yellow; in auriferoides they are some shade of brown; the pile of the vertex and front of the females of aurifer is always wholly yellow, while in auriferoides it is always mixed black and yellow. The sides of the third antennal segment of aurifer fe- males are always nearly parallel for the proximal half, and then evenly tapering to the end; this segment in auriferoides is always more or less restricted near the base and before the tip, and the widest part is usually before the middle. The dark color in the wing of the males of aurifer does not extend beyond the tip of vein Ri; in auriferoides it more or less fills all of cell Ri- The females are more readily separated than the males. Reexamination of the material listed in the earlier paper by Maughan (1935) shows that all specimens studied are auriferoides rather than aurifer. The specimens are in too poor condition to include as paratypes. We have found auriferoides to be more numerous than aurifer in the study area. Literature Cited Hesse. A. J. 1938. A revision of the Boniby- liidae (Diptera) of southern Africa. Ann. So. Afr. Mus. 34(1053): 332, text figures. Maugii.xn, L. 1935. A systematical and morph- ological study of Utah Bombyliidae with notes on species from inteniiountain states. .1. Kansas Ent. Soc. 8( 1-2) :27-80. 4 pis. Maughan, L.. and D. E. Johnson. 1936. Notes on Utah Bombvliidae (Diptera). Proc. Utah Acad. Sci., Arts. Lett. 13:197-201. .Johnson, D. E, and L. Maughan. 1953. Stu- dies in Great Basin Bombvliidae. Great Basin Nat. 13 (1-2): 17-27. BREEDING RANGE EXPANSION OF THE S J ARLING IN UTAH Dwight G. Sniitli' Abstract. — The discover^' and observation of colonies of starlings nesting in the eastern Great Basin desert indicates further expansion of the breeding range of this species in ITtah. Data on nest site se- lection, nesting productivity, food habits, and relationships with other avian s])ecies are presented. The dispersal and establishment of star- Hngs (Sturnis vulgaris) in North Ameri- ca has been well documented (Bent 1950; Davis 1960). In Utah information on the initial spread of this species was sum- marized b}- Behle (1954). While study- ing raptor populations in central Utah from 1967 to 1972 I observed several small colonies of starlings nesting in des- ert habitats. I believe these colonies show the establishment of starlings as mem- bers of the breeding avifauna of the cen- tral Utah Great Basin desert. As such, they represent a breeding range expan- sion which reflects the general adapta- bility of this species. History of the St.^rling in Utah The range expansion of starlings in Utah was initially gradual. Starlings were first observed in the state in February 1939 near Salt Lake City. In the next year small winter flocks were observed in other parts of Salt Lake Valley and a lone individual was reported near Lehi, Utah County. In January 1941 a flock of 200 was observed at Alt. Carmel, Kane County, in south central Utah (Behle 1958). From 1941 to 1947 small winter flocks were frequently observed near feedlots and ranches in Salt Lake Valle}^ In 1948 their winter range again expand- ed; a flock of 1,000 individuals was seen northward in Davis County; and a single individual was found in Kanab, Kane County. Their winter range expansion continued in 1950, when they were re- corded for the first time northward in Box Elder County and westward in Tooele County. During the decade 1950- 1960 winter flocks of starlings increased tremendously. Bailey (1966) reported flocks as large as 100,000 and noted that they constituted an important agricultural pest of feedlots and orchards in 16 ^Dcpl. of Biology. Soullicrn Coniicclicul Stntc C;ollegc, Nc\ counties of the state, from Washington Count}' in the south tf) Box Elder Coimty in the north. The first nest of the starling in Utah was found on 25 May 1949 in an old woodpecker or flicker hole on the west side of Salt Lake City. In the following year a nest was discovered in a shed at Randolph, Rich County (Behle 1954). Within six years starlings were nesting at many localities in the central, popu- lated valleys of the state. In 1956 a star- ling nest containing young was found in a shed at Lynn, in the northwestern cor- ner of the state (Behle 1958). Starlings have since spread throughout most of the state and now are a sizable com- ponent of the breeding bird populations of the towns, settlements, and ranches of the northern Great Basin area (Hay- ward 1967). During my study of raptor populations in Cedar Valley, LTtah County, and Rush Valley, Tooele County, I found starlings and house sparrows (Passer domesticus) to be the most com- mon breeding birds in small towns such as Fairfield and Cedar Fort. In these set- tlements they typically nested in a variety of buildings and in holes in cottonwoods and willows. They were also frequently observed nesting in wind breaks border- ing agricultural fields and pastures. Here they usuall}^ nested in holes in living and dead trees, but unused bulky stick nests of hawks and magpies [Pica pica) were also appropriated. My observations in other settled areas of Tooele, Juab, and Millard counties showed similar choices of nesting site selection and habitats by starlings. I belie^■e that my observations of star- lings nesting in desert habitats warrant particular interest because this suggests invasion and adaptation to a new habitat as well as a further range extension by this species in Utah. 419 420 GREAT BASIN NATURALIST Vol. '35, No. 4 Nesting Localities and Nest Sites I observed nine breeding sites located in three counties, all judged to be ex- amples of a starling breeding range ex- pansion into habitats of the northern Great Basin desert. It should, however, be mentioned that each spring and summer I frequently observed starlings in addition- al localities, and the breeding range ex- pansion of this species is undoubtedly oc- curring over a much wider area than my records indicate. Four of the nine sites supported small but regular breeding populations during the six-year study. The other sites were irregularly active. Starlings displayed their well-known adaptability in choosing nest sites (Kes- sel 1957), and a comparison of nesting habitats and nest site selection indicates some degree of opportunism. Although Michael (1971) considered his observa- tions of starlings nesting in rocky cliffs in Kentucky and West Virginia to be significant, I found 18 of 42 (42.9 per- cent) of the nests I actually located to be placed in crevices in the sheer walls of quarries and cliffs. Of the remainder, 9 (21.4 percent) nests were located in some type of abandoned mining structure, 8 (19 percent) were found in holes in juni- pers {Juniperus osteosperma) ^ and 7 (16.7 percent) in stick nests of hawks, owls, and ravens (Corvus corax). The choice of nesting sites reflected the di- versity of the nesting habitat. Usually pairs of a colony were found nesting in a variety of sites where available. Several sites will be described in some detail to facilitate comparison with ])os- sible future breeding locations. Utah County. — Five sites were found, all in the western section of the county. A colony was located in the vicinity of an abandoned mine and clay pit oper- ation at Five Mile Pass, which is about six miles west of Fairfield. The habitat at this locale is desert scrub intermixed with widely scattered junipers. Four mine structures are still standing and there are six large quarries and clay pits. From 1967 to 1972 a total of 17 nests were found at this site as follows: 2 in 1967; 4 in 1968: 5 in 1969; 3 in 1970; 1 in 1971 and 2 in 1972. Several additional pairs were seen each year after 1968. and some may have been nesting. Of the nests found, seven were placed in crevices in the rock walls of clay pits, four in holes in junipers, three in the walls of a mine shack, two in mine bunkers, and one in the beam sup- ports of a mine tunnel. A second colony active during all six breeding seasons was located in the vi- cinity of the abandoned Little Topliff quarry at Ten Mile Pass. This site was approximately 14 miles southwest of Fair- field and 5 miles northwest of Allan's Ranch. The habitat of this site is a mix- ture of grasses and desert scrub. Fourteen nests were found at this site: 2 in 1967, 2 in 1968, 5 in 1969, 3 in 1970, 1 in 1971, and 1 in 1972. Again, additional pairs were observed each 3'ear after 1968. Eight of the nests were located in creA'ices in the sheer rock face of the quarry, two in an unused golden eagle {Aquila chrysae- tos) nest, one in an abandoned prairie falcon {Falco mexicanus) nest, and three in cracks in a wooden retaining wall. Another Ten Mile Pass site was located in the abandoned Big Topliff quarry which is about one mile east of Little lopliff. This quarry, one of the largest in the area, is bordered entirely by desert scrub communities. Two nests were found in 1970, both constructed in an un- used golden eagle nest. Several individ- uals, some of which may have been paired, were observed in 1971. In 1969 a third colony was found at Ten Mile Pass, about two miles east of Big Topliff quarry and three miles north- west of Allan's Ranch. Several individuals and two nests each were found in 1969 and 1970 in a large limestone cliff line 400 feet above the valley floor. The sur- rounding habitat is sparse desert scrub de- void of trees. Three of the four nests were located in crevices in the cliff face. The fourth was in an unused red-tailed hawk {Butco ja- maicensis) nest constructed in a large crevice. A fifth colony was found in the vicinity of the abandoned Tintic Empire Mine, located in the foothills of the Boulter Mountains approximately four miles north of Eureka and two and one-half miles southeast of Allan's Ranch. The surrounding habitat of this colony was a large stanrl of widely spaced junipers. Two nests, both located in holes in juni- pers, were found in 1969; other pairs Dec. 1975 SMITH: STARLING IN UTAH 421 were present. Onl}- one bird was seen when this site was rechecked in 1970, and none were observed in 1971. I was unable to visit this site during the 1972 breeding season. Tooele County. — I found nests at two separate sites in this county and evidence of nesting activity at one ad- ditional site. In 1968 and again in 1969 I found a starling nest in a hole in a juniper near the entrance to Black Rock Canyon. This nest site was approximately seven iniles east of Vernon. This site was unusually interesting because it was located in a juniper which also supported an active great horned owl {Bubo virginianus) nest during the two nesting seasons. Both starling and great horned owl nesting attempts were successful during the two nesting seasons. No nest was present in 1970 or 1971, but starlings were observed in May and June in the same juniper stand, although about one-half mile above the original nesting site. On 15 June 1969 I observed several starlings in a juniper stand in the north- ern foothills of Simpson Mountains. This site is approximately 16 miles northeast of Simpson Springs. One nest was found in a hole in a juniper, and, judging by the behavior of the other birds, additional nests may have been present. Juab County. — Murphy et al. (1969) observed 12 starlings near a golden eagle nest located in sandstone cliffs at Yuba Dam State Park and presumed them to be nesting. On 15 May 1970 a pair of starlings was observed approximately six miles north of Trout Creek in the foothills of the Deep Creek Mountains. One carried nesting material and was seen in the vi- cinity of a small stand of junipers. We were, however, unable to locate a nest. Reproductive Chronology and Success Starlings were not found in the vi- cinity of the desert nesting sites during the fall and winter months (September through February), although small flocks were frequently observed in nearby set- tlements. They began appearing in the future nest site vicinity in early March, and the majority were present by late March and early April. During this time they were frequently seen ins])ecting di- lapidated mine buildings and holes in junipers and often reacted to my pres- ence near these sites by protesting vigor- ously. Adults carrying nesting material were seen in late March and throughout April. Most nests were constructed of grass, pri- marily wheat grass (Agropyron spica- tum), and frequently lined with feathers. Four of the nests I found were decorated with juniper greenery. Nests containing eggs were found from late April through mid- June. Dates of the 24 nests with eggs which we found are as follows: 27, 30 April; 1, 2, 4(3), 10 (2), 17(4), 19, 21(2), 30 May; 4, 9(3), and 19 June. I found no evidence of at- tempts to raise a second brood in July and August. The clutch size of these nests averaged 4.2 - 1.2 eggs (range, 3-7; mode, 4). A clutch size comparison with other areas is presented in Table 1. Utah clutch size averaged significantly smaller than those of New York and Holland (t = 3.95, 2.84 respectively; P<0.05 for both) but not significantly different from northwestern England (t=1.5, P>0.90). Young were in the nest from mid-May through mid- July. My earliest and latest dates for nests with young are 14 May and 28 July. The brood size of 17 nests was 5.9 ±1.1 young (range, 2-7; mode, 4). Interestingly, there was no significant difference among brood sizes of Utah, Table 1. Clutch and brood size comparison of central Utah nests.* Location No. No. eggs No. No. young Author clutches per clutch broods per brood Central Utah 24 4.5 ± 1.2 17 3.9 ± 1.1 Present Study Ithaca, New York 199 5.5 ± 0.9 230 4.3 ± 1.3 Kessell (1957) Holland 1592 5.2 ± 1.0 1377 4.4 ± 1.3 Kluijver (1933) NW England 105 4.9 ± 1.1 913 4.2 ± 1.1 Lack (1948) •Data is average ^ one standard deviation. 422 GREAT BASIN NATURALIST Vol. 35, No. 4 New York, Holland, and England nests, indicating perhaps a somewhat higher overall hatching success of Utah nests. Overall reproductive success was high. Of 13 nests on which I was able to obtain complete information, 12 successful!}' hatched young; and of these, 11 nests fledged young. Two nests were aban- doned; one containing three eggs and one with five young. Neither jDair at- tempted to renest. Overall hatching and fledging success was 94.2 percent and 84.6 percent, respectively. Both percent- ages are slightl}"^ higher than reported from previous studies in other areas. Foraging and Food Habits Information on foraging and food habits is limited and w^as obtained from morning observation of three nests, two in 1969 and one in 1970, all located in the Five Mile Pass nesting colony. Adults were observed from a parked vehicle with a 40X spotting scope attached to a window mount. Only those food items brought to the nest site which could be identified are included in the results presented in Table 2. Adults foraged predominantly in the sagebrush-wheatgrass {Artemisia- Agropy- ron) associations which were the common plant communities in the nesting site vi- cinities. They spent considerably less time in the ground layer vegetation of pinyon- juniper (Pinus-Juniperus) communities and among the rubble-strewn floor of quarries. Over 86 percent of the arthropod food items brought to the nest were insects. Of these, Orthoptera comprised 56 per- cent and Colcoptera almost 27 percent. Araenids were the only other animal food which was taken in significant quantities. In a food habit study in eastern Texas Table 2. Arthropod food of starlings in the eastern Great Basin. Item No. indv. % Freq. Acrididae 31 47.0 Tettigoniidae 6 9.1 Carabidap 11 16.7 Tenebrionidae 3 k") Scarabidac 3 4.5 Cicadellidae 2 10 Formicidae 1 1.5 Araneidae 9 13.6 Totals 66 99.9 based on stomach contents, Russell (1967) found Orthoptera and Coleoptera, particu- larly Carabidae, to comprise 84 percent of the total insects eaten and 68 percent of the total food, with other arthropods and some plant material constituting the remainder of the diet. I did not identify any utilization of plant material for food, but results are undoubtedly biased be- cause the small nesting populations pre- cluded collection of adults and young for stomach contents analysis. Both Killpack and Crittenden (1952)' and Bailey (1966) noted the extensive use of such plant ma- terials as grain and corn silage by win- tering flocks of starlings. Starlings are undoubtedly opportunistic in their feed- ing habits and utilize the most available food. This is reinforced by a comparison of the food habits of these desert nesting starlings with the result of Fautin's (1946) investigations of the invertebrate populations of the sagebrush community. Analysis reveals that, with the exception of Formicidae, starlings utilized the most prevalent ground layer invertebrates in the sagebrush community. Relations with Other Species Starlings appear to be the predominant avian species in the vicinity of their breeding locales. Other birds observed in the same locale included the house spar- row, pinyon jays {Gyrnnorhinus cyano- cephala)^ scrub jays {Aphelocoma coe- rulescens), mourning doves (Zenaidura macroura), common nighthawks (Chor- deiles minor), and mountain bluebirds (Sialia currucoides) . Only house spar- rows were common nesting associates, and at two sites. Big and Little Topliff quar- ries, they outnumbered the starlings. At these and other sites the two species ap- peared to mutuall}' tolerate one another and no aggressive interactions were ob- served. We did observe starlings inter- acting aggressively with scrub jays (twice) and mourning doves (twice) which had perched in the immediate vi- cinity of the nesting site. In each in- stance the starlings displaced the intrud- ing birds. The only direct evidence of nest displacement which we obsened concerned a mountain bluebird nest which contained six eggs when discovered on 15 May 1970. When this nest was re- checked five days later we found an Dec. 1975 SMITH: STARLING IN UTAH 423 adult starling sitting on a clutch of six eggs. No trace of the mountain bluebird was found, and the adults were not ob- served again in the nesting territory. In a previous study of the food habits of raptors breeding in the eastern Great Basin Desert, starlings were occasionally recorded as prey of several hawk and owl species (Smith and Murphy 1973). It is possible that their plumage, size, and aggressive habits make them conspicuous targets and hence more liable to be preyed upon by raptors. Discussion Within the relatively short time span of 30 years the starling has become a pre- dominant component of Utah's avian fauna. Its successful invasion and estab- lishment can be roughly delineated into three segments, each of approximately 10 years duration. In the first 10-year period (1939-1948) following their ap- pearance in the state, starlings were ob- served only as individuals or small flocks of winter visitants. In the next 10 years (1949-1958) these winter flocks increased tremendously in size and greatly ex])and- ed their winter range to include most of the populated central j)ortions of the state. The first nesting attempts were reported early in this period and by the end of the decade small nesting populations were found in many widely separated towns and settlements of the state. In the third 10-year period (1959-1968) both winter and summer starling populations had in- creased in size to the point at which the starling had achio\ed the status of a major pest species. During this time the starling became (with the possible ex- ception of the house sparrow in some areas) the most abundant bird throughout most of the settled portions of the state. Although most common in cities and towns, they were also found in widely separated settlements and ranches. In winter they formed large flocks which, through their feeding and roosting ac- tivities, became an economic nuisance to feedlot owners and fruit growers through- out the state. At the present time, so far as is known, starlings nest in all the settlements and towns in the state and have recently ex- tended their breeding range into desert habitats. Although initiallj^ sporadic. their occupation of distinctly desert habi- tats for nesting pur])oses has recently be- come more widespread, suggesting that they have been able to adapt to a new habitat type. In analyzing the range expansion and establishment of starlings in Utah I con- sider the following to be of significance: (1) mobility, (2) suitable climate, (3) suitable habitat, and (4) sufficient popula- tion })ressure. The four factors are, in fact, a measure of the starling's adaptability and, when considered together, explain the success of this species. Although not specifically investigated in this study, their rapid range expansion across North America indicates that starlings have adequate powers of mobility and wide climatic tolerances. While largely non- migratory, their behavorial adaptation of forming large, mobile winter flocks which break up and disperse to fa\orable nesting areas in spring undoubtedly allows some exploratory activity which may in turn lead to further range expansion. Water, but not food, may pose the only j)otential limiting factor for starling range expansion into the Great Basin desert. Starlings at their desert breeding sites were observed drinking water from ephe- meral rain puddles and from liA'estock watering troughs. The adaptability of starlings to new habitats is well known (Kessell 1957). A review of the pattern of starling in- vasion in Utah suggests that a habitat with some form of human modification provides a favorable impetus for range ex- pansion. Thus, widely separated towns and settlements were occupied within 20 years, but the interAening areas of desert were not invaded until after starling populations were well established in near- Ijy settlements. Only after starling popula- tions had occupied these locales did a further range expansion into the upper Great Basin desert take place. It is quite ])ossible that the human habitats pro- vided breeding sites, probabl}' due to the well-known breeding behavior adaj)ta- tions of this species with reference to man, which resulted in local population increases. Pressures resulting from these local population increases may have en- couraged starlings to examine the adja- cent habitats of the upper Great Basin desert. 424 GREA'l' BASIN NATURALIST Vol. 35, No. 4 The short time span ol their successful utilization of this new habitat suggests behavorial adaptations rather than genetic changes within the population. However, the possibility of future evolutionary changes in populations occui)ying such habitats may warrant further study. Literature Cited Bailey. E. P. 1966. Abuiidanre and activity of starlings in winter in northern lUah. Con- dor 68:152-162. Behi.e, W. H. 1954. Changing status of tiie starling in Utah. Condor 56:49-50. . 1958. The birds of tlie Raft River Mountains, northwestern Utah. Univ. Utah •Biol. Ser. 11(6): 1-49. Bent, A. C. 1950. Life histories of North American wagtails, shrikes, vireos and their allies. U.S. Natl. Mus. Bull. 197:182-214. D.wis. D. E. 1960. Comments on the migration of starlings in eastern United States. Bird- Banding 31:216-219. F.AUTiN, R. W. 1946. Biotic communities of the northern desert shrub biome in Western Utah. Ecol. Monogr. 16:251-310. H.\YW.-\RD, C. L, 1967. Birds of tlie Upper Colo- rado Rivei- Basin. Brigham Young Univ. Sci. Bull.. Biol. Ser. 9(2): 1-64. Kessei.i., B. 1957. A study of the breeding biolo- gy of the European starling (Slurnis vul- garis) in North America. American Midi. Nat. 58:257-331. Kii.i.PACK, M. C, AND D. N. Crittenden. 1952. Starlings as winter residents in the Uinta Basin, Utah. Condor 54:338-344. Kluijver, I. H. N. 1933. Bijdrage tot die bio- logic en do ecologie van den spreeuw (Stur- nus vulgaris vulgaris L.) gedurende zijn voortplantingstiid. Versl. Meded. Planten- ziekt (Wageningen) 69:1-145. Lack. D. 1948. Natural selection and family size in the stalling. Evolution 2:95-110. Michaei,. E. D. 1971. Starlings nesting in rocky cliffs. Bird-Banding 42:123. Murphy, J. R., F. J. C.mviinzand, D. G. Smith, and ,I. B. Weston. 1969. Nesting ecology of raptorial birds in central Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. 10:1-36. RussELi,. D. N. 1971. Food habits of the star- ling in eastern Texas. Condor 73:369-372. Smith, D. G., and J. R. Murphy. 1973. Breed- ing ecology of raptors in the eastern Great Basin of Ihdh. Brigham Young Univ. Sci. Bull.. Biol. Ser. 18(3): 1-76. SOME PARASITES OF PADDLEFISII (POLYODON SPAT HULA) FROM THE YEELOWSTONE RIVER, MONTANA Lawrence I^. Lockai nd R. Randall Parsons- Abstract. — One species of copepod (Ergasilus elongatus), one tiematode (Dichholhriuiu liaiuu- laturn), two nematodes (Camallanus oxrcephalus am\ Contracaecum sp.). and two cestodos (Marsi- pomctra hastata and M. pnrva) were recovered from 17 i)addlefisli [Polyodun spalhula) collected from the Yellowstone River. Montana, on 11 and 18 May 1971 The male paddlefish averaged 60.4 nematodes and 157.6 cestodes per infected fish while the females averaged 12.0 nematodes and 415.7 cestodes. The higher intensity of cestode infection in female paddlefish was attributed to their larger size and consequent greater intake of food resulting in more exposure to the cestode intermediate hosts (Cyclops bicuspidalus) . The gastrointestinal tract was dissected, and the contents were washed onto a 2()()-niesh screen, then transferred to an illuminated tray (Barber and Lockard 1973) for examination. Cestodes were fixed in AFA (alcohol-formalin-acetic acid) and stained with Delafield's hema- toxylin. Nematodes were placed in a mix- ture of 70 percent alcohol and 5 percent glycerine and later mounted in glycerine. The liver, heart, gall bladder, spleen, and gills were dissected and each was placed in a jar wdth water and agitated on a mechanical shaker for 5-10 minutes. The contents were poured onto a 200- mesh screen, washed, transferred to an illuminated tray, and examined. Trema- todes and copepods were placed in a mix- ture of 70 percent alcohol and 5 per- cent glycerine. The ovaries, testes, and gas bladder were examined grossly and observed abnormalities checked micro- scopicall}'. Results There have been relatively few reports on the parasites of paddlefish (Polyodon spat hula). Linton (1898) found ta])e- worms in paddlefish from the Ohio River, Ohio. Stockard (1907) mentions large numbers of cestodes in paddlefish from the state of Mississippi; and Wilson (1914) found parasitic copepods on paddlefish from the Mississippi River in Illinois and Iowa. Simer (1929 and 1930) and Beaver and Simer (1940) examined 171 paddlefish from the Tallahatchie River, Mississippi, and reported on tre- matodes and cestodes from them. Bang- ham and Vernard (1942) found one pad- dlefish from Reelfoot Lake, Tennessee, infected with trematodes, nematodes, and cestodes. Meyer (1946) discovered leech- es parasitizing paddlefish while Causey (1957) examined one paddlefish and found parasitic copepods. Meyer (1960) studied cestodes from paddlefish collected from the Mississippi River in Iowa and the Missouri River in South Dakota. Hugghins (1972) examined one paddle- fish from Fort Randall Resenoir on the Missouri River in South Dakota and found it parasitized with tapeworms and nema- todes. Weisel (1973) reported uniden- tifted tapeworms and nematodes from 3 paddlefish from the Yellowstone River, Montana. The present study was under- taken in an effort to gain information on the parasites of paddlefish from Montana. Methods This report was based upon the necrop- sy of 17 paddlefish collected from the spawning run in the Yellowstone River near Intake, Montana, on 11 and 18 May 1973. The fish were eviscerated, and the viscera and gills were placed in plastic bags and frozen for later examination. The ten male paddlefish had an av- erage weight of 10.6 kilograms and range of 6.8 to 13.6 kilograms, while the seven females had an average weight of 22.8 kilograms and range of 20.0 to 27.2 kilo- grams. Table 1 lists the rates of infection with copepods, trematodes, nematodes, and cestodes recovered from paddlefish during this study. The parasitic copepod Ergasilus elongatus was identified by Dr. Z. Kabata of the Fisheries Research Board of Can- ada. The trematode Diclybothrium ham- ulaturn was identified by Dr. Fred P. Meyer of the Bureau of Sport Fisheries and Wildlife. Identifications of the nema- todes Camallanus oxycephalus and Con- tracaecum sp. w^ere confirmed by Dr. E. J. J312 S. Grant Ave.. Picirc. SD -Department of Biology. Montana State Universily. Bozcmnn. Montana 425 426 GREAT BASIN NATURALIST Vol. 35, No. 4 Table 1. Incidence of parasites of Pohdou spathula collected from the Yellowstone River. Montana. Percent infected Mean no. of parasites (range) Parasite Male Female Male Female Ergasilus elongatus Diclybothrium hamulatum Carnal lanus oxycephalus Contracaecum sp. Nematode cysts Marsipometra hastala M. parva 30 14 100 100 80 86 80 86 100 100 100 100 90 100 69.4 (12-302) 12.0 (4-25) 157.6 (34-356) 415.7 (37-1013) Hugghins of South Dakota State Univer- sity and Dr. G. L. Hoffman of the United States Fish and Wildlife Service, respec- tively. Spherical cysts (1-3 mm in diam- eter) containing larval nematodes were present on the surface of the stomach, pyloric caecum, and the intestine of all fish examined. All 17 paddlefish ex- amined were infected by tapeworms identified from Beaver and Simer (1940) and Meyer (1960) as Marsipometra has- tata and M. parva. Discussion It was felt that the handling procedures greatly reduced chances of discovering parasitic copepods as well as monogenetic flukes which may have been present on the gills of the fish. Causey (1957) says statements of incidence of infection for copepods have little value. Thus, the in- cidence of infection of copepods found in Table 1 may not be indicative of the num- ber of infected fish actually present. Like- wise, quantitative analysis on the pres- ence of the monogenetic fluke would be of little value. Differences in intensity of nematode infections (Table 1 ) between male and female paddlefish were noted but were difficult to interpret because of the pres- ence of immature Contracaecum sp. in the intestine. Simer (1930) and Beaver and Simer (1940) name three species of cestodes in- fecting paddlefish: Marsipometra hastata, M. parva, and M. confusa. Meyer (1960) examined tapeworms from these studies and his own material and concluded that M. hastata and yi. confusa were synony- mous. He also found Cyclops hicuspidatus to be the intermediate host of M. hastata. In this study no attempt was made to distinguish M. confusa from .M. hastata. Females had a higher intensity of tape- worm infection than male paddlefish, and this may be attributable to larger size of the females. Females weighed more than twice as much as males and thus would have ingested more of the inter- mediate hosts of these parasites while feeding. Acknowledgment The authors express their appreciation to Jeffrey Bagdanov for assisting with the necropsy of the paddlefish. Literature Cited B.^NGH.^M. R. V., .and C. E. Vernard. 1942. Studies on parasites of Reelfoot Lake fish. IV. Distribution studies and checklist of parasites. J. Tcnn. Acad. Sci. 17(1): 22-38. Barber. D. L., and L. L. Lockard. 1973. Some helminths from mink in southwestern Mon- tana, with a checklist of their internal para- sites. Great Basin Nat. 33(1): 51-60. Beaver, P., and P. H. Simer. 1940. A restudy of the three existing species of the cestode genus Marsipometra Cooper (Amphicoty- lidae) from the Spoonbill. Polyodon spathula (Wal.), Tr. Am. Micr. Soc. 59: 167-182. Causey, D. 1957. Parasitic Copepoda from Louisiana freshwater fish. Am. Midi. Nat. 58(2): 378-382. HuGGiNs. E. I. 1972. Parasites of fishes in South Dakota. South Dakota Exper. Sta. Bull. 484: 1-73. Linton, E. 1898. Notes on cestode parasites of fishes. Proc. U.S. Nat. Museum 20: 423-456. Meyer. F. P. 1960. Life history of Marsipo- metra hastala and the biology of its host. Polyodon spathula. Iowa State ITniv. Library, Ames, Iowa (unpubl.). Meyer, M. C. 1946. Further notes on the leeches (Piscicolidae) living on freshwater fishes of North America. Tr. Am. Micr. Soc. 65(3): 237-249. Simer. P. H. 1929. Fish trematodes from the lower Tallahatchie River. Am. Midi. Nat. 11(12): 563-588. Simer, P. H. 1930. A preliminary study of the cestodes of the spoonbill. Polyodon spathula (Wal.). Tr. 111. State Acad. Sci. 22: 139-145. SiocKARD. C. R. 1907. Observations on the nat- ural history of Polyodon spathula. Amer. Nat. 41: 753-766. Weisei.. G. E. 1973. Anatomy and histology of the digestive system of the paddlefish {Polyodon spathula) J. Morph. 140(2): 243-251. Wii.soN, C. B. 1914. Copepod parasites of fresh- water fislies and theii- economic relations to mussel glochidia. Bull. F.S. Bur Fish. 34(1914): 331-374. REPRODUCTIVE CYCLE OF THE BELDINC; GROUND SQUIRREL {SPERMOPHILUS BELDINGl BELDINGI) -. SEASONAL AND AGE DIFFERENCES Martin L. Morton^ and John S. Gallup- Abstract. — The reproductive cycle in Belding ground squirrels was studied in the Sierra Nevada Mountains at two locations, one at 2,100 ni elevation, the other at 3,000 m. Adults emerged from hibernation completely prepared physiologically for reproduction. Males tended to emerge slightly ahead of females and yearlings tended to emerge later than adults. Year- ling females were fertile but produced smaller litters than adults, 4.48 vs. 6.31. Yearling males were infertile. They exhibited a slight seasonal cycle in testicular growth but did not reach sexual maturity. Testicular growth and spermatogenesis were incipient in many adults and in yearlings prior to hiber- nation. Hibernation and seasonal breeding are important survival strategems of rodents living at high latitude or high altitude. Typically in these environments there are extreme seasonal oscillations in ambient conditions. Winters tend to be long and cold and summers brief and sharply de- limited. Dormancy is employed as a means of bridging the long gap of energy shortage in winter, and breeding is co- incident with the clement weather and abundant food of surmner. Both respon- ses require advance preparation and ac- curate timing to be maximally adaptive. The physiology of hibernation is cur- rently a viable, active field of study, whereas seasonal breeding has aroused less interest and its complexities, especial- ly in wild populations, are poorly under- stood (Chapman 1972). Herein we report on seasonal changes in reproductive functions of the Belding ground squirrel {Spertnophilus beldingi beldingi), a hibernator that lives at high altitude in the Sierra Nevada Mountains of California. Methods This study extended from 1969 to 1973 and was on S. b. beldingi living principal- ly in meadows of Lee Vining Canyon, Mono County, California. A number of squirrels were live-trapped or shot at Big Bend (elevation ca. 2,100 m), but most data are from those living near Tioga Pass (elevation ca. 3,000 m). At both areas we carried on an extensive mark- release program throughout the time squirrels were active above ground. This program enabled us (1) to follow seasonal changes in body weights and dimensions and external appearance in individuals of known age and (2) to collect animals of known age for specimens. Live-trapping was conducted with Tomahawk wire mesh traps baited with peanut butter. A few specimens were col- lected with a .22 caliber rifle. Animals trapped for the first time were toe-clipped in a standard pattern never involving more than one toe per foot. All animals were examined externall}^ for appearance of vulva and mammae or of scrotal pig- mentation and position of testes. Freshly excised reproductive organs were fixed in Bouin's solution and trans- ferred to 70 percent ethanol. At the time of transfer they were debrided, blotted, and weighed to the nearest 0.1 mg. Tis- sues were sectioned at 7 or 1 0/t and stained with hematoxylin and eosin. Measure- ments of seminiferous tubules were taken with an ocular micrometer. Results Arousal schedule.^ — The sequence and pace of events in the active season were alike at the two study areas, but the active season at Big Bend usually be- gan at least six weeks in advance of that at Tioga Pass. Snowcover was not com- parable at the two sites at the beginning of the season. At Big Bend most burrow sites were clear of snow when emergence occurred. At Tioga Pass emergence tended to begin on knolls that were the first areas to become snow free, but many in- dividuals tunneled out at sites covered by snow up to a depth of 2 m. Similar ef- ^Biology Department, Occidental College, Los Angeles, California 90041. -Alaska Department of Fish and Game, Homer, Alaska. 427 428 GREAT BASIN NATURALIST Vol. 35, No. 4 fects of snowpack on emergence of S. columbianus were noted by Shaw (1925). There was considerable annual variation in snow conditions and the schedule of S. b. beldingi was affected accordingly (see beyond). Adult males were the first animals seen above ground at a given location, but a few adult females and an occasional year- ling could be found within a few days thereafter. Testes. — At emergence adult males had scrotal testes weighing about 2 to 3 g, the maximum weight seen during the entire active season (Fig. 1). The scrotum was darkly pigmented. Within a month after emergence testicular weight of adults began to decrease noticeably and testes had become inguinal or abdominal in position and scrotal pigmentation was decreasing. Six weeks after emergence tes- ticular weight had decreased to a seasonal minimimi that was maintained thereafter at Big Bend until onset of hibernation (Fig. 1, upper). At Tioga Pass, however, considerable increase in testicular weight of adults occurred during the last few weeks of the season (Fig. 1, lower). ' ' ■ Big Bend . ■ • •.1 : t : ° " o I I r I .> . :»• *° 8 • ^11 ' "I .8 ° + Fig. 1. Paired testes weights of Spermophilus beldingi beldingi throughout tlie active season at two study areas. Note log scale on ordinate. Cross on lower right corner of Tioga Pass data indicates mean testicular weight of juveniles prior to hibernation. In histological perspective, testes of adults showed intense spermatogenic ac- tivity from time of emergence through onset of weight collapse. During this time seminiferous tubules were of large diam- eter (150 to 250/i) with spermatozoa fill- ing the liunina. As testicular weight de- creased spermatogenesis ceased and tu- bule diameter decreased. During June at Tioga Pass, for example, diameters went from about 150 to 60/j.. Beyond June, lumina in seminiferous tubules were ab- sent. Recrudescing testes of adults col- lected in late August at Tioga Pass had a thickening germinal layer with num- erous primary spermatocytes and a few secondary spermatocytes present. Semi- niferous tubules had enlarged slightly to a diameter of 80 to lOO/i. Yearling males tended to emerge later than adult males, were of smaller body size (Morton and Parmer, in press), and had considerably smaller abdominal testes (Fig. 1). This was most clearly ob- served at Tioga Pass where we had a larger pool of marked animals to collect from. Testes of yearlings were about three times heavier at emergence than when they entered hibernation the previous fall as juveniles. A decrease in testicular weight occurred soon after emergence, fol- lowed by an increase toward the end of the season. Two yearlings collected at Big Bend on 18 April 1973 had paired testes weighing 920 and 940 mg. Seminal vesicles of these animals were 37.4 and 34.2 mg, respec- tively. Although well above resting level, neither set of glands approached those of sexually active adults in weight or cel- lular maturation. There was some evidence of a cycle in spermatogenic activity in yearling testes in that a few spermatocytes were pro- duced early in the season. Diameter of seminiferous tubules remained small (be- low lOO/i) throughout the season, and no advanced stage of spermatogenesis was found in any yearling testis. As in adults at Tioga Pass, tubule diameter increased slightly in concert with increased testic- ular weight, and spermatoc>'te numbers increased just prior to hibernation. Seminal vesicles. — Seminal vesicles of adults tended to increase in weight for a few days following emergence, re- mained at maximum size for a few Dec. 1975 MORTON, GALLUP: GROUND SQUIRREL 429 weeks, then decreased to minimum size for the rest of the season (Fig. 2). These glands were only slightly enlarged in yearlings early in the season and tended to decrease in weight thereafter. As shown in S. lateralis by McKeever (1964), sem- inal vesicle growth is controlled by tes- ticular hormone. The near-maximum size of seminal vesicles in recently emerged S. h. heldingi suggests that upon final arousal they have fully secretory testes. In microscopic appearance the heaviest seminal vesicles of adults had a distended mucosal epithelimn and lumina filled with seminal fluid in a colloidal state. As involution occurred the colloid disap- peared, the mucosal layer became shrun- ken and folded, and the lumina nearly disappeared. This appearance was main- tained through onset of dormanc}'. Sem- inal vesicles of yearlings were without de- tectable cellular change throughout the active season. 5exual cycle of females. — Adult fe- males appeared to be sexually receptive almost immediately after emergence as judged by their swollen, open vulvae and enlarged, turgid uteri. Copulation was never observed, but additional evidence that mating occurred soon after emer- gence is that a few adults were already lactating during the fifth week after the first active females were seen. Gestation period in S. beldingi is thought to be 27 to 31 days (Turner 1972). Yearlings were in estrous later in the season than adults because they tended to emerge later and because estrous appeared to be delayed in smaller yearlings until additional body growth had occurred. Seasonal changes in ovarian weight were about two-fold and were similar for the two age groups. For the first six weeks after emergence ovaries weighed 20 to 30 mg. Weight then decreased to 8 to 15 mg for the remainder of the season. Anovulatory follicles and corpora lutea tended to enlarge during gestation and reached maximum diameters at partu- rition. These structures shrank in post- partum females. By the end of lactation follicular cavities were much reduced or absent and corpora lutea were becoming indistinct. No ovarian recrudescence was observed in yearlings or adults prior to hibernation, but follicular enlargement 1 • Big Bend • AcJuM . , o Yeorlmg . • ; 8 .. •• ; » oo i ° '« o Fig. 2. Paired seminal vesicle weights of Spermophilus beldingi beldingi throughout the active season at two study areas. Note log scale on ordinate. began in juveniles during their last weeks of activity after older animals had al- ready become dormant. Emergence of females at Tioga Pass occurred over a span of several weeks. As a result parturition dates were spread out and the percentage of females lac- tating at a given time was almost never 100 percent (Fig. 3). As far as we know, all females reproduced, although some of the smaller yearlings collected in late June at Tioga Pass had unscarred, thin, virgin-like uteri. Note that testicular atrophy was well advanced in adult males at that time (Fig. 1, lower). It is possible that a few yearling females did not bear young. Another possibility is that implan- tation was delayed in those of small body size. There are indications in other studies that female Spermophilus may become impregnated well after testicular collapse has begun in males (Wells 1935; Tomich 1962; McKeever 1966). If these smaller females do have young, they will be born relatively late in the season (we have observed considerable disparity in size of juveniles at the end of the season; Morton, Maxwell, and 430 GREAT BASIN NATURALIST Vol. 35, No. 4 Fig. 3. Mean percentage of female Spermophilus beldingi beldingi thought to be lactating at spe- cific 5-clay intervals at Tioga Pass. Numerals indicate sample sizes. Wade 1974), and if these young survive, a self-perpetuating cycle of late repro- duction by small yearling females will have been established. Litter size. — Our specimens and rec- ords of frequently retrapped females in- dicate that they have one litter per sea- son. This was true even of the earlist fe- males to breed. We obtained measure- ments of litter size through counts of placental scars and implanted embr3'os in collected specimens and through litters born in captivity (Table 1). The differ- ence in means between counts of scars and of embryos was not different ac- cording to a t test (P> 0.05) but some prenatal loss did occur. Ten of the 228 embryos examined (4.4%) were partially reabsorbed. This is similar to the re- absorption rate found in S. lateralis (McKeever 1964; Skryja and Clark 1970) and in S. richardsonii (Sheppard 1974). The lowest estimate of litter size was ob- tained from births in captivity. Cannibal- ism of their young by confined females Table 1 . Measurements of litter size in Spermo- philus beldingi beldingi. Mean S.D. N Placental scars Adults Yearlings 6.88 5.00 1.22 17 2 Implanted embryos Adults Yearlings 6.33 4.75 2.01 1.07 21 20 Young per captive Adults Yearlings 5.71 4.00 1.45 1.00 17 5 Total, all measurements Adults Yearlings 6.31 4.48 1.67 1.40 55 27 was noted in a few cases but all cases may not have been detected. When all measurements were summed, mean litter size of adults (6.31) was sig- nificantly larger (P< 0.01) than that of yearlings (4.48). This is quite similar to age differences in litter size observed in S. armatus (Slade and Balph 1974). Dec. 1975 MORTON. GALLITP: OHOUNI) SQl'IKKEL 431 Discussion In most published reports there is little indication that yearling Sperniophilus of either sex are functionally or visually- separable from older animals (compare Mayer and Roche 1954; McKeever 1963, 1964; Carl 1971; Zimmerman 1972), al- though they may constitute a considerable portion of the breeding population. How- ever, yearlings engage in unique behav- ioral interactions with older animals as they are integrated into the conmnniity of reproducers ( Michener and Michener 1973) and should be the focus of more study. Even in the larger-bodied sciurids such as Marmota monax (Christian, Stein- berger, and McKinney 1972) and Cy- nomys leucurus (Bakko and Brown 1967) some males are sexually mature as year- lings. The lack of I'eproductive ca])acity in yearling male but not in female S. b. hel- dingi is an interesting contrast that indi- cates the operation of sex-specific selec- tion factors. As pointed out by Conaway (1971) the nonpregnant cycle is a rarity and cannot be afforded by most natural populations. The breeding season is de- layed slightly in yearling female S. b. beldingi by their emergence schedule and the apparent necessity for some to attain greater body size before pregnancy. Nev- ertheless, all or nearly all do reproduce. In males, however, there is greater total body size to be reached than in females (Morton 1975) and selection seems to have favored the strategy of diverting energy expenditure from reproductive activities toward growth. Indeed from the last half of the yearling season onward males are significantly larger than fe- males (Morton and Parmer, in press). We have additional evidence that delayed sexual development in male S. b. beldingi is tied directly to body size. Two males born in captivity and kept under class- room conditions for display became ex- cessively obese in their first autumn, went through bouts of estivation for several months, and possessed scrotal testes soon after resuming normothermia at about nine months of age. Growth rates of captive S. b. beldingi juveniles are comparable to those of other hibernatory Sperniophilus, but there are indications that growth is much slower in feral animals than in captives even be- fore weaning (Morton and Tung 1971). It may be that it is unusually difficult for S. b. beldingi living at high altitude to achieve adult size and reproductive i)o- tential simultaneously. We have sug- gested (Morton and Tung 1971) and now shown clearly (Morton and Parmer, in press) that iS. b. beldingi do not reach maximum body size until late in their second year of life or beyond. Likewise, Sheppard (1972) found that yearling S. richardsonii aged on the bases of eye lens weight, tooth wear, and epiphyseal clo- sure had smaller mean body weights than older animals. The habitual lack of a reproductive cycle among certain male members of a population could not be tolerated unless a mating system were employed that as- sured impregnation of all receptive fe- males. A priori one might expect repro- ductively active males in such a system to be polygamous. The details of the mat- ing system of S. b. beldingi should be elucidated shortly by behavioral studies in progress at Tioga Pass by Paul Sher- man of the University of Michigan. The social system of a ground squirrel population has recently been implicated in sexual development of yearling males. Slade and Balph (1974) found that year- ling male S. armatus seldom had scrotal testes and rarely, if ever, bred. After the population was artificially reduced, how^- CAer, many yearling males were sexually active. Slade and Balph associate this precocity with low harassment as juve- niles, early arousal from hibernation, and decreased encounters with aggressive squirrels following emergence. They did not report on body size in these animals. It is possible but improbable that we have been studying a ground squirrel population with unique growth patterns and mating system. It is important to recognize that accurate aging in many studies has not been possible due to their brevity or to lack of history on recogniz- able individuals. For example, in his study of S. b. oregonus conducted in Las- sen County, California, at 1,370 to 1,730 m elevation, McKeever (1963) refers only to adults and jmeniles. Body weights shown for adult S. b. oregonus are 5 to 10 percent lower throughout the season than those of S. b. beldingi (Morton 1975), but juvenile 5. b. oregonus are at least 432 GREAT BASIN NATURALIST Vol. 36, No. 4 20 percent heavier than S. b. hcldingi (Morton, Maxwell and Wade 1974) be- fore entering hibernation. Mean maxi- mum testicular weight occurs in both subspecies at the beginning of the active season, but it is about three times greater in S. b. beldingi than in S. b. oregonus. This seems anomalous to us and, coupled with bod}' weight data and McKeever's observation that only 70 percent of adult male S. b. oregonus were sexually active, suggests that at least some j^earling S. b. oregonus could be distinguished from older animals on the bases of body size or reproductive capacity if ages were known . Finally, it should be recognized that unusual ecological conditions exist at high altitude. Many factors, both terres- trial and extraterrestrial in origin, could function as inhibitors of growth and de- velopment in young animals. Seasonal breeding. — Gro\^i;h of the reproductive tract during the whole hi- bernation period is well known among Spermop/iilus. Remarkably, gonadal re- crudescence, at least of testes, begins even prior to dormancy in several spe- cies with short active seasons such as S. undulatus (Mitchell 1959; Hock 1960), 5. b. oregonus (McKeever 1963), S. later- alis (McKeever 1964), S. richardsonii (Clark 1970), and 5. b. beldingi (pres- ent study). McKeever (1963) noted that S. b. oregonus emerged with testes of max- imum size. Usually the final stages of spermatogenesis in hibernatory Spermo- philus are not reached until ten days or more after emergence even in species at high latitude (Hock 1960). The total read- iness of 5. b. beldingi to reproduce at time of emergence is undoubtedly a response to conditions imposed by the short .sum- mers of high altitude. Seasonal breeding is highly adaptive in that young are born at a time favorable for their survival. At high altitufle this favorable season is comprcss(Hl temporally and has rather sliarply drawn boundaries. To cope successfully with these circum- stances the cycle of sexual maturation in S. b. beldingi is completed during dor- mancy. A corollary is that reproductive preparation must ha\(' a precise })hase relationship to average snowTnelt patterns and related euA ironmental effects. There is sensitivity, however, on the part of newly emerged animals to ambient con- ditions. At Tioga Pass considerable an- nual variation in snowpack, schedule of snowmelt, and emergence of vegetation were documented (Morton, Maxwell, and Wade 1974; Morton, in press). The schedule of S. b. beldingi was affected ac- cordingly. In 1969, for exam])le, snow- pack was about 240 percent above nor- mal, whereas in 1972 it was 35 percent below normal. As judged by subsequent emergence times of juveniles and their growth curves, reproduction occurred about three weeks later in 1969 than in 1972 (Morton, Maxwell, and Wade 1974). Reproduction is not delayed in- ordinately, however, even in heavy snow years, because about 25 percent of prehibernatory fat reserves still remain at emergence, ])ro-\iding a buffer to food requirements during the first weeks of activity (Morton 1975). Acknowledgments. — Many Occiden- tal College students participated in por- tions of this study. We are grateful to all of them but would like to acknowledge in particular Phil Rekey, Cassie Cusick, Shena Huang, Roland Leong, Cathy Maxwell, Robert Parmer, and Charlie Wade. Financial support was provided by Occidental College and by National Science Foundation Grant GR 29146X1. Literature Cited Bakko. v.. B., and I.. N. Brown. 1967. -Breed- ing Ijiology of the white tailed prairie dog, Cynomys leucurus. in Wyoming. J. Mam- mal. 48:100-112. (Jarl, E. a. 1971. l^opulation control ni arctic ground squirrels. Ecology 52:395-413. Chapman. D. I. 1972. Seasonal changes in the gonads and accessory glands of male mam- mals. Mammal. Rev. 1:231-248. Christian, .1. J.. E. Steinberger, and T. D. Mc- KiNNEY. 1972. Annual cycle of spermato- genesis and testis morpholog\' in \M)od(hncks. I. Mammal. 53:708-716. CiARK. T. W. 1970. Richardson's ground squiricl (Spermophilus richardsonii) in the T.aramie Basin, Wyoming. Great Basin Nat. 30:55-70. CoNAW.AY. C. H. l')71 Ecological adai)tation and manmialian reproduction. Binlog\ of Re I)roducti(m 4:239-247. Hock. R ,1. 1960. Seasonal yaiialions in physio logic functions of aiitic ground sciuiicels and black bears. Mus. Conip. /ool Bull.. Harvard 124:155-169. Mayer. W. V.. and E. R. Roche. 1954. Devel- opmental patterns in the Barrow ground sciuirrel. Sprrmofihilus utnlulalus barrowen- sis. Growth 18:5^-69. Dec. 1975 MORTON, GALLUP: GROUND SQUIRREL 433 McKeever, S. 1963. Seasonal changes in body weight, reproductive organs, pituitary, adrenal glands, thyroid gland, and spleen of the Belding ground squitTel {Citellus bel- dingi). Amer. J. Anat. 113:153-173. . 1964. The biology of the golden- mantled ground squirrel. Citellus lateralis Ecol. Monogr. 34:383-401. 1966. Reproduction in Citellus beldingi and Citellus lateralis in northeastern Cal fornia. Symp. Zool. Soc, London 15:365-385 MiCHENER. G. R., AND D. R. MiCHENER. 1973 Spatial distribution of yearlings in a Rich- ardson's ground squiiTel population. Ecology. 54:1138-1142. Mitchell, O. G. 1959. The reproductive cycle of the male arctic ground squirrel. J. Mam- mal. 40:45-53. Morton, M. L. Adaptive strategies of Zonotric- hia breeding at high latitude or high altitude. Proc. 16th Intern. Ornithol. Congr., in press. . 1975. Seasonal cycles of body weights and lipids in Belding ground squirrels. Bull. So. Calif. Acad. Sci., in press. Morton, M. L., C. S. Maxwell, and C. E. Wade. 1974. Body size, body composition, and behavior of juvenile Belding ground squirrels. Great Basin Nat. 34:121-134. Morton, M. L., and H. L. Tung. 1971. Growth and development in the Belding ground squirrel (Spermophilus beldingi bel- dingi). J. Mammal. 52:611-616. Morton, M. L., and R. J. Parmer. Body size, organ size, and sex ratios in adult and year- ling Belding ground squirrels. Great Basin Nat., in press. Shaw, W. T. 1925. The seasonal differences of north and south slopes in controlling the ac- tivities of the Columbian ground squirrel. Ecology 6:157-162. Sheppard, D. H. 1972. Reproduction of Rich- ardson's ground squirrel (Spermophilus rich- ardsonii) in southern Saskatchewan. Canadian J. Zool. 50:1577-1581. Skryj.-^, D. D., and T. W. Cl.ark. 1970. Repro- duction, seasonal changes in body weight, fat deposition, spleen and adrenal gland weight of the golden-mantled ground squirrel, Spermophilus lateralis lateralis (Sciuridae) in the Laramie Mountains, Wyoming. South- western Nat. 15:201-208. Sl-^de, N. a., and D. F. Balph. 1974. Popu- lation ecology of Uinta ground squirrels. Ecology 55:989-1003. ToMicH, P. Q. 1962. The annual cycle of the California ground squirrel Citellus beecheyi. Univ. Calif. Pub. Zool. 65:213-282. Turner, L. W. 1972. Autecology of the Bel- ding ground squirrel in Oregon. Ph.D. thesis, LTniv. Arizona, 149 pp. Wells, L. J. 1935. Seasonal sexual rhythm and its experimental modification in the male of the thirteen-lined ground squirrel {Citellus tridecemlineatus) . Anat. Rec. 62: 409-447. A NEW COMBINATION IN PENSTEMON ( SCROPHULARI ACEAE) Stephen L. Clarki Abstract. — Field and herbarium studies of Penstenion cyananthus Hook. ssp. longiflorus Pennell suggest that this taxon be elevated to species rank. Penstemon cyananthus Hook. ssp. longi- florus Pennell is a tall, attractive blue- flowered plant presently known only from Beaver, Millard, and Piute counties of southern Utah. It was first collected by Edward Palmer (376, NY) near Beaver City, Utah, and was distributed as P. glaber cyananthus, a synonym of P. cyan- anthus. Pennell recognized that Palmer's col- lection differed from the species as fol- lows: the calyx lobes are broader than in the species, the flowers longer, the stems and lower leaves are puberulent, and the inflorescence is strongly secund. In ad- dition, the distributions are verj' differ- ent. Penstemon cyananthus ssp. cyanan- thus has never been collected from south- ern Utah, but it is one of the most fre- quenty encountered species of Penstemon in the Wasatch mountains of northern Utah, southeastern Idaho, and southwest Wyoming (Fig. 1). On the basis of these morphological and geographical differences, Pennell (1920) described and named the subspecies P. cyananthus ssp. longiflorus. It is the writer's opinion that differ- ences of such magnitude warrant the ele- vation of this taxon to specific status, hence the following new combination is proposed: Penstemon longiflorus (Pennell) Clark comb, nov., based on P. cyananthus Hook, ssp. longiflorus Pennell in Contr. from the U.S. Nat. Herb., vol. 20, part 9, p. 353, 1920. Penstemon longiflorus can easily be recognized by its puberulent lower stem and leaves, broad sepals, longer corolla, secund inflorescence, and its blue tipped staminode. Both taxa have a chromosome number of 2n = 16. \ 1 "T (Xrif'* AS> • ; 1^ V|,..i.i-i 1 — .... — — ^^ — ^__ .._ ■}■/ y "\ ...Jr Fig. 1. Distribution of P. cyananthus (circles) and P. longiflorus (squares). 'Department of Botany, Weber State College, Ogden. Utah 84043. 434 SOME RELATIONSHIPS BETWEEN WATER FERTILITY AND EGG PRODUCTION IN BROWN TROUT {SALMO TRUTTA) FROM MONTANA STREAMS Lawrence L. LockarcP Abstract. — Relationships between water fertility (as measured by conductivity and alkalinity) of 17 Montana streams and the attainment of sexual maturity and fecundity of their resident female brown trout were studied. Fish from the streams having conductivity and alkalinity levels greater than 100 micromhos/cm and ppm CaCO,^, respectively, were younger at sexual maturity than fish from waters with lower levels. The attainment of earlier sexual maturity in fish from the former streams could not be completely explained on the basis of greater growth rates. Fish from the stream having the highest levels of conductivity had the slowest growth rate but became sexually mature at the youngest age. A positive relationship was found between chemical fertility of streams and the fecundity- of their fish. However, in the stream having the highest levels of conductivity, fish were the least fecund. It was concluded that the chemical fertility of these streams is generally related to the age at sexual maturity and fecundity of fish. The size and age at sexual maturity and the fecundity of female fish appear to be related to features of their environ- ment. In Pennsylvania brown trout {Salmo trutta) from infertile waters had a smaller proportion of mature fish per age class and smaller weight of eggs than comparable fish from fertile waters (Mc- Fadden, Cooper, and Anderson 1965). Scott (1962) and Bagenal (1969) dem- onstrated that rainbow trout {Salmo gaird- neri) and brown trout, respectively, brought a lower number of eggs to ma- turity under reduced nutritional levels than fish on higher levels of nutrition. This study is an attempt to determine the relationships between the conduc- tivities and alkalinities of Montana streams and (1) the size and age at sex- ual maturity and (2) the fecundity of the brown trout in those streams. Field collections were made from 8 September to 23 October 1972 and from 1 Septem- ber to 19 October 1973. Methods A total of 449 female brown trout were collected by electrofishing at sites on streams in the Clark Fork of the Co- lumbia River and in the Yellowstone and Missouri River drainages (Fig. 1). These streams had a wide range of physical and chemical conditions (Table 1). At least one fall, winter, and summer measurement of conductivity and alka- linity was made at each collecting site. The field measurements from each stream were averaged with the }'early conduc- tivity and alkalinity averages obtained from Water Resources Data for Montana (U.S.G.S. 1972) where available. Dis- charge values were obtained by aver- aging available yearly values from the above U.S.G.S. records with values measured or estimated by fisheries biolo- gists of the Montana Fish and Game De- partment. All fish were collected during Sep- tember and October of 1972 and 1973 (Table 2). Fish taken were preserved in 10 percent formalin and later washed in water and stored in 40 percent isopropyl alcohol. Fixation in formalin causes spe- cimens to shrink about 3-4 percent in length and increase 5-12 percent in weight (Parker 1963). After preservation, fish were measured, weighed, and scale samples were removed for age determi- nations. Each fish was classified as ma- ture or immature according to the condi- tion of the eggs in its ovaries. Mature fish containing eggs in a gradient of sizes were not used in the fecundity anal- yses because the number of eggs is re- duced by resorption throughout the ma- turation period, and regressing eggs could not be distinguished from maturing eggs in these fish. Only fish having distinct re- cruitment and maturing eggs without in- tervening size classes of eggs were used in fecundity work. The ovaries from these fish were removed and the number of maturing eggs determined by actual count. The streams from which collections ^Current address: 312 South Grant Avenue, Pierre, SD 57501. 435 436 GREAT BASIN NATURALIST Vol. 35, No. 4 Table 1. Selected chemical and physical features of streams sampled. Ck)lIection site no.' Location of site Conductivity (micromhos/cm) Alkalinity (ppm CaCOa) Discharge (C.F.S.) 1 Rock Cr. 2 St. Regis R. 3 Big Hole R. 4 W. Gallatin R. 5 Madison R. 6 Baker Cr. 7 O'Dell Cr. '8 L. Prickley Pear Cr. 9 E. Gallatin R. 10 Shields R. 11 Flagstaff Cr. 12 Beaverhead R. 13 16 Mile Cr. 14 So. F. Musselshell R. 15 Little Blackfoot R. 16 Bluewater Cr." 17 Big Horn R. 18 Bluewater Cr.^ 70 49 169 80 51 555 207 117 1,125 230 T18 791 249 107 1,409 317 154 70 348 167 100 358 195 69 360 195 400 402 221 159 405 197 5 521 193 405 522 195 50 561 243 83 612 188 105 798 209 18 805 188 3,500 1,387 214 28 'See Figure 1 ^Section above Bluewater Fish Hatchery 'Section below Bluewater Fish Hatcherj' Fig. 1. Map showing location of collecting sites. were made were grouped into classes pri- marily on the basis of similarities in con- ductivities followdng the technique used by McFadden et al. (1965). Streams from which collections 1 and 2 were taken each had less than 100 units of conductivity and alkalinity and formed Class I. Streams from which collecting sites 3 through 18 were located had alkalinity values above 100: thus conductivities were used as the primary indicator of water fertility. Class II contained streams on which collection sites 3 through 11 were located. These streams had conduc- tivities ranging from 207-405 micromhos/ cm at 25 C. Streams of collecting sites 12 through 15 had conductivities of from 521-612 and comprised Class III except for the analysis of size and age at sexual maturity in which Stream 17 was includ- Dec. 197; LOCKARD: BROWN TROUT 437 Table 2. The location, date, and number of fish collected. 1972 1973 Collection Number Collection Number Total Collection site date fish date fish fish 1 (Rock Cr.) Oct. 13 16 Sept. 14 17 33 2 (St. Regis R.) Sept. 1 22 22 3 (Big Hole R.) Oct. 9 17 17 4 (W. Gallatin R.) Oct. 18 17 Sept. 25 11 28 5 (Madison R.) Sept. 21 13 Sept. 20 19 32 6 (Baker Cr.) Oct. 10 17 Sept. 26 9 26 7 (O'Dell Cr.) Sept. 22 11 Sept. 21 8 19 8 (L. Prickley Pear Cr.) Oct. 23 16 Oct. 2 17 33 9 (E. Gallatin R.) Sept. 18 15 Oct. 4 16 31 10 (Shields R.) Oct. 6 11 Sept. 24 6 17 11 (Flagstaff Cr.) Oct. 9 9 9 12 (Beaverhead R.) Sept. 26 14 Oct. 17 "9 23 13 (16 Mile Cr.) Sept. 11 13 Oct. 11 14 27 14 (So. F. Musselshell R.) Sept. 24 13 13 15 (Little Blackfoot R.) Oct. 1 28 28 16 (Bluewater Cr.)' Sept. 8 T3 Sept. 12 14 27 17 (Big Horn R.) Sept. 15 14 14 18 (Bluewater Cr.)' Sept. 8 '\3 Sept. 12 37 50 ^Section above Bluewater Fish Hatchciy -Section below Bulewater Fish Hatchery ed. The streams of Class II and III were combined into Class IV because their fish had similar relationships to conduc- tivity. Class V was made up of Bluewater Creek on which collecting sites 16 and 18 were located. These collecting sites were grouped together primarily because of their high conductivities. The fish in stream classes were statis- tically compared by selected procedures and techniques from "Statistical Meth- ods" (Snedecor and Cochran 1971) and "Statistical Methods" (Arkin and Colton 1972). Additional techniques were pro- vided by Dr. R. E. Lund, Alathematics De- partment, Montana State University. Results Size and Age of Sexually Mature Female Brown Trout Generally the attaimnent of sexual ma- turity of fish is dependent on size and age. Inspection of age groups within stream classes indicated an apparent ten- dency for a higher proportion of the larger females to be sexually mature (Table 3). To test the linearity of this trend, regressions were made on fish from age groups in stream classes showing an increase in sexual maturity with increas- ing length. In age group I, fish from Class V streams showed a significant positive linear relationship between length and sexual maturity (P= 0.001). In age group II, a significantly higher proportion of larger fish were sexually mature in Stream Classes I, II, III, and IV with P values of less than 0.05. In age group III + , fish from Stream Classes II and IV had significant positive linear relation- ships between length and sexual maturity (P<0.05). McFadden et al. (1965) found a tendency within a given year class for a higher percentage of larger than smaller fish to be sexually mature. The effect of age on the attainment of sexual maturity in fish was determined by comparing the proportions of sexual- ly mature fish between age groups by a technique of R. E. Lund. Only 2 percent of age group I fish 8.0-13.9 inches long from Class IV streams were mature, while 34 percent of comparably sized fish in age group II were mature. The difference in proportions was significant with a P = 0.001. There were significantly fewer ma- ture 6.0-8.9 inch fish from Class V streams in age group I than in age group II (P = 0.08). These combined probability values demonstrated a significantly (P= 0.001) higher proportion of age II fish were ma- ture than age I fish. Significantly more of size group 10.0-19.9 inch fish from Class IV streams were mature at age III+ than age II (P= 0.001). This rela- tionship of a higher percentage of older females being sexually mature than youn- ger females in the same size group has been reported by McFadden et al. (1965). 438 GREAT BASIN NATURALIST Vol. 35. , No. 4 Table 3. Size and age of sexually mature female brown t rout by stream classes Length (inches) Stream classes Age I II III IV V group #Fish %Mat. #Fish %Mat . #Fish re Mat. #Fish i%Mat. #Fish -^cMat. I 4.0-4.9 0 0 0 0 5 0 5.0-5.9 0 0 0 0 3 33 6.0-6.9 3 0 0 0 0 16 31 7.0-7.9 4 0 1 "o 0 1 0 17 47 8.0-8.9 >1 0 7 0 2 0 9 0 6 100 9.0-9.9 1 0 6 0 5 0 11 0 0 10.0-10.9 0 11 0 10 0 21 0 0 ll.O-n.9 0 0 7 0 7 0 0 12.0-12.9 0 0 2 0 2 0 0 13.0-13.9 Total 0 0 2 50 2 50 0 II Q 0 25 0 28 4 53 2 47 43 6.0-6.9 u 0 0 0 1 100 7.0-7.9 0 0 0 0 6 67 8.0-8.9 2 0 1 0 0 1 0 6 83 9.0-9.9 9 33 5 0 2 0 7 0 5 100 10.0-10.9 6 33 16 13 9 11 25 12 1 100 n.0-11.9 1 100 29 31 3 0 32 28 1 100 12.0-12.9 2 100 23 61 9 22 32 50 4 100 13.0-13.9 T 100 9 67 7 57 16 63 0 14.0-14.9 0 19 84 3 100 22 86 0 15.0-15.9 0 3 67 5 80 "s 75 0 16.0-16.9 0 1 ilOO 4 100 5 100 0 17.0-17.9 0 1 100 3 100 4 100 0 18.0-18.9 0 0 2 100 2 100 0 19.0-19.9 Total 0 0 1 100 T 100 0 21 43 107 48 48 50 155 48 24 88 III + 8.0-8.9 0 0 0 0 I 100 9.0-9.9 0 0 0 0 1 100 10.0-10.9 3 67 1 0 0 1 -9 0 11.0-11.9 8 75 ^ 67 1 100 4 75 2 100 12.0-12.9 1 100 4 100 2 100 6 100 0 13.0-13.9 5 100 12 92 6 100 18 94 0 14.0-14.9 2 50 n 92 4 75 17 88 1 100 15.0-15.9 2 100 16 100 9 100 25 100 1 100 16.0-16.9 2 100 14 93 1 100 15 93 0 17.0-17.9 1 100 8 100 2 100 10 100 0 18.0-18.9 1 100 5 100 3 100 8 100 0 19.0-19.9 0 3 100 1 100 4 100 0 20.0-20.9 Total 0 1 100 0 1 100 0 25 84 80 94 29 97 109 95 6 100 Grand Total 55 55 212 59 105 51 317 57 77 61 Com])arisons were made of the pro- portions of sexually mature female brown trout between stream classes using a meth- od of Arkin and Colton (1972). No sig- nificant difference (0.05 level) was found in the proportion of mature females in Class II and III streams either by age group or grand total so further compari- sons by this method were made between the fish of Stream Classes I, IV, and V. There was no significant difference be- tween Class I and IV streams in the pro- portions of mature females in age grouji II; however, Class IV streams had a sig- nificantly higher ]:)roj)ortion of mature females in age group III+ than did Class I streams (P= 0.054). Class V streams had a higher proportion of sexually ma- ture females than both Class I and IV streams in both age group I (P= 0.014 and 0.001, respectively) and age group II (P= 0.001 and 0.001, respectively). Fisher's randomization test (Bradley 1968) was used to further test the hypoth- esis that maturation increases as conduc- tivity increases. The probability of obtain- ing the iiu reased proportions of mature lish ill all age groups with the increasing Dec. 1975 LOCKARD: BROWN TROUT 439 conductivities in Stream Classes I, II, III, and V (Table 3) is P= 0.00014. The class I and IV streams in this stud}' were similar in conductivit}" and alkalin- ity to the infertile and fertile streams in the studies of McFadden and Cooper (1962) and McFadden et al. (1965). In the latter study, fish from fertile waters attained maturity at an earlier age than those from infertile waters. This was at- tributed partially to a greater growth rate of fish in fertile waters; however, the authors also found higher proportions of fish of the same size and age were sexual- ly mature in fertile streams. In the present study, this latter rela- tionship was not observed among fish from Class I and IV streams. Instead, higher proportions of females of a given size and age tended to be mature in the less fertile Class I streams. The differences in age at maturity between fish from Stream Classes I and IV, therefore, seem closely related to differences in growth rate. The distribution of sizes of speci- mens of given age groups do indicate faster growth rates in Class IV streams (Table 3). McFadden and Cooper (1962) also reported positive correlations between growth rates of brown trout and environ- mental fertilit}^ Class V streams had higher conductiv- ity and alkalinity values than any of the streams studied by McFadden et al. (1965). In the more fertile waters (Class V), fish matured at younger ages than in less fertile waters (Classes I through IV); however, this early maturity in Class V streams was not due to a faster growth rate in fertile waters. That is, the smallest fish in each age group are found in the Class V streams (Table 3). Therefore, some factor other than grow1;h rate or chronological age apparent!}' influenced the size and age at which sexual maturity was reached by fish from the very fer- tile (chemically) waters of Class V. Fecundity Regression analyses for the number of mature eggs in a fish (dependent variable) on fish length (independent variable) were applied to the fish of the individual streams and stream classifications. T tests for the regression of numbers of eggs on lengths were statistically significant at the 0.05 level for fish from all streams and stream classifications with most probabil- ity values being less than 0.01. Flagstaff Creek and the Big Horn River were omit- ted from analyses because of an insuf- ficient number of mature fish. The regression lines of numbers of eggs regressed on fish length in stream classifications are shown in Figure 2. The regression lines with steeper slopes show a greater increase in number of eggs per increment of length than lines with lesser slopes. Regression coefficients, slopes of the re- gression lines, were calculated for these regressions on each stream and stream classification (Table 4). The slopes of the stream classification regressions were tested for significant differences by a method of R. E. Lund (Table 5). Six of the 8 comparisons of slopes of regressions were significantly different at the 0.05 level. Fish from Stream Class I were less fe- cund than fish from Stream Class II (Fig. 2 and Table 4). The difference between these stream classes was statistically sig- nificant at the 0.05 level (Table 5). This relationship of increased fecundity with increased conductivity is similar to that Table 4. Regression coefficients (slopes) of streams and stream classifications. Stream or No. of eggs regressed on fish length classifi- Regression Std cation coefficient error N Rock Cr. 213 22 18 St. Regis R. 173 47 12 Stream Class I 254 22 30 Big Hole R. 284 40 7 W. Gallatin R. 426 83 14 Madison R. 457 64 16 Baker Cr. 164 53 13 O'Dell Cr. 248 51 14 L. Prickley Pear Cr. 236 43 16 E. Gallatin R. 218 29 19 Shields R. 252 43 16 Stream Class II 325 20 115 Beaverhead R. 172 98 15 16 Mile Cr. 143 48 15 So. F. Musselshell R. 315 112 6 Little Blackfoot R. 249 65 15 Stream Class III 286 39 51 Stream Class IV 318 18 166 Bluewater Cr.^ 170 20 19 Bluewater Cr." 113 16 21 Stream Class V 147 13 40 ^Section above Bluewater Fish Hatchery -Section below Bluewater Fish Hatchery 440 GREAT BASIN NATURALIST Vol. 35, No. 4 "I I I I r 11 12 13 14 IS Fish Length (inches) 20 21 ■Fig. 2. The regression lines of number of eggs on length for fish in stream classifications. Table 5. Comparisons between stream classes by slopes of regressions of number of eggs on fish length. Slope Number of eggs regressed on fish length comparison T df P I vs II 2.42 77 0.018* I vs III 0.72 46 0.475 I vs IV 2.25 73 0.028* I vs V 4.18 47 0.000* II vs III 0.90 75 0.371 II vs V 7.61 149 0.000* III vs V 3.39 60 0.001* IV vs V 7.80 174 0.000* •Significant at the 0.05 level found by McFadden et al. (1965) in fish from infertile and fertile streams having conductivities and alkalinities similar to those of Class I and II streams in this study. Fish from Stream Class III appeared to be more fecund than fish from Stream Class I (Fig. 2 and Table 4). This rela- tionship of increased fecundity with in- creased conductivity was not statistically significant at the 0.05 level (Table 5). Stream Class III contained streams with higher levels of conductivity than those reported by McFadden et al. (1965). Fish from Stream Class IV (Stream Classes II and III combined) represent fish from a broad category of chemically fertile streams with conductivities from about 200 to 600 micromhos/cm. In gen- eral, these fish were more fecund (P= Dec. 1975 LOCKARD: BROWN TROUT 441 0.028) than fish from Class I streams, which represent chemically infertile waters. Fish from Stream Class V, which had the highest conductivity, had the lowest fecundity. The conductivity values of this stream were about three times great- er than the highest values reported by McFadden and Cooper (1962). The above results suggest some factor other than conductivity is determining the fecun- dity of fish in this stream class. Summary An inverse relationship between chemi- cal fertility and age at sexual maturity was found in brown trout from streams of Montana in this study. This same re- lationship between the chemical fertility of streams, as measured by conductivities and alkalinities, and the age of sexual ma- turity of brown trout from Pennsylvania has been reported by McFadden et al. (1965). They suggested this relationship was due partially to fish in fertile streams having greater growth rates. Growth rate may account for the age at maturity in fish from 1 6 of the 1 7 streams in this study, but cannot explain the age at ma- turity in fish from Bluewater Creek. Fish from Bluewater Creek attained sexual ma- turity much earlier than fish from less fertile streams; howe\er, these fish from the stream with the highest conductivity had the poorest growth rates of all the fish studied. This shows growth rate was not the determining factor in the attain- ment of sexual maturity for fish from Bluewater Creek. McFadden et al. (1965) found a posi- tive relationship between the chemical fertility of streams and the fecundity of their fish. A similar relationship was found between chemical fertility and fe- cundity in fish of this stud}' from streams having conductivities similar to those studied by McFadden et al. (1965). How- ever, fish from Bluewater Creek, chemi- cally the most fertile stream, were the least fecund. Generally the age at sexual maturity of fish from all stream classes and the fecundity of fish from Stream Classes I, II, III and IV appeared to be related to the chemical fertility of their streams. Literature Cited Arkin, H., and R. R. Coi.ton. 1972. Statistical methods. Barnes and Noble Books, New York. 344 pp. Bagenal, T. B. 1969. The relationship between food supply and fecundity in brown trout Salmo trutta L. J. Fish Biol. 1:167-182. Bradley, J. V. 1968. Distribution-free statistical test. Prentice-Hall, New Jersey. 399 pp. McFadden. J. T.. and E. L. Cooper. 1962. An ecological comparison of six populations of brown trout (Salmo trutta). Trans. Am. Fisheries Soc. 91:53-62. McFadden, J. T.. E. L. Cooper, and J. K. An derson. 1965. Some effects of environment on egg production in brown trout (Salmo trutta). Limn, and Ocean. 10(l):88-95. Parker. R. R. 1961 Effects of formalin on length and weight of fishes. J. Fish. Res. Bd. Canada 20(6) : 1441-1455. Scott. D. P. il962. Effect of food quantity on fecundity of rainbow trout. Salmo gairdneri. ,J. Fish. Res. Bd. Canada 19:715-731. Snedecor, G. W.. and W. G. Cochran. 1971. Statistical methods. Iowa St. Univ. Press. Iowa. 593 pp. United States Geological Survey. 1972. Water resources data for Montana. Part 2. Water ([uality records. 218 pp. SOME RELATIONSHIPS BETWEEN INTERNAL PARASITES AND BROWN TROUT FROM MONTANA STREAMS Lawrence L. Lockard^, R. Randall Parsons^, and Barry M. Schaplow^ Abstr.'^ct. — Forty-five percent of 306 brown trout from 16 Montana streams were infected with one or more of the nematodes Cystidicoloides salvelini, Bulbodacnids globosa, Rhabdochona sp., and Eustrongylides sp. The relationships between incidence and intensity of nematode infections and age and sexual maturity of the host fish were studied. Generally, se.xuallj' mature female brown trout had a higher rate of infection and had more nematodes per infected fish than immature fe- male brown ti'out. Higher incidence and intensity of infection in sexually mature fish was attributed to more aggressive feeding behavior leading to more exposure to the intermediate hosts (mayflies) of the nematode parasites. Nimierous parasitological surveys of trout in North America have been re- ported; however, few have included in- formation on the parasites of brown trout {salmo trutta) . Van Cleave and Mueller (1934) studied 13 brown trout from Oneida Lake, New York, and 3 from one of its tributary streams. Fifty-eight Wis- consin brown trout were studied by Bang- ham (1946) and Fischthal (1947a, 1947b, 1950, and 1952), with 54 originating from streams. In the western geographic region of the United States the studies of Bangham (1951), Hugghins (1959), Al- exander (1961), and Fox (1962) include information on the parasites of brown trout. In these studies a total of 55 brown trout were examined: 51 from lakes and 4 from streams. Heckmann (1971) ex- amined 28 brown trout from Montana for blood parasitism. This study is an attempt to examine some relationships between internal para- sites, brown trout, and the fish's stream habitat. It is based on the necropsy of 306 brown trout collected from 17 sites on 16 streams in south and western Montana. The specimens were collected in early September through October (1972 and 1973) just prior to and during the spawn- ing seasons. According to Van Cleave and Mueller (1934), fish are most heav- ily parasitized in the summer season when they are most actively feeding. Since Fox (1962) found nematode infections in Montana brown and rainbow^ trout (Salmo gairdneri) highest in August and September, these findings may be indica- tive of near maximum parasite infec- tions for brown trout in Montana streams. Methods and Materials All fish were collected by use of electro- fishing gear described by Vincent (1967). Collecting sites were selected to reflect a wide range of geographic locations (Fig. 1), and physical and chemical conditions (Table 1). Collections were taken on both sides of the Continental Divide from streams belonging to the Clark Fork of the Coltmibia, Yellowstone, and Missouri River drainages. At least one fall, winter, and summer measurement of conductivi- ty and alkalinity was made on each stream. These field measurements were averaged with the available yearly con- ductivity and alkalinity averages ob- tained from Water Resources Data for Montana (USGS 1972). Discharge values were obtained by averaging available yearly values from the above USGS rec- ords with values measured or estimated by fisheries biologists of the Montana Department of Fish and Game. The streams sampled varied in average dis- charge from 5 cubic feet per second to 1,409 cubic feet per second, in average conductivity from 70 to 1,387 micromhos /cm at 25 C., and in alkalinity from 49 to 221 ppm CaCO'. Ecologically the streams are diverse although they all support nat- urally reproducing populations of browoi trout. Immediately after capture the fish were preserved in 10 percent formalin, washed in water and stored in 40 percent isopro- pyl alcohol for later examination. Parker (1963) indicates fish shrink about 3-4 percent in length and gain between 5 and 12 percent in weight when preserved in formalin. About three to eight months after collection fish were individually ^Fish and Wildlife Service, U.S. Department of the Interior, P.O. Bo.\ 250, Picnc, South Dakota 57501. 442 Dec. 1975 LOCKARD, ET AL.: BROWN TROUT 443 Fig. 1. Map showing location of collecting sites. Table 1. Selected physical and chemical characteristics of 16 Montana streams and the incidence and intensity of nematode infections in brown trout from them. Stream Ave.# Collection dis- ' # % worms site no.i Stream charge Cond. Alk. Fish Par. par. /fish 1 Madison R. 1409 249 107 13 100 68.7 2 Big Hole R. 1125 207 117 19 100 14.3 3 W. Gallatin R. 791 230 118 17 100 21.7 4 St. Regis R. 555 80 51 22 14 1.7 5 Beaverhead R. 405 521 193 14 43 1.7 6 E. Gallatin R. 400* 360 195 15 87 3.9 7 Rock Cr. 169 70 49 16 63 3.4 8 Shields R. 159 402 221 11 18 1.5 9 L. Blackfoot R. 105 612 188 28 21 1.2 10 O'Dell Cr. 100* 348 167 11 46 3.0 11 So. F. Musselshell R. 83 561 243 13 8 1.0 12 Baker Cr. 70* 317 154 28 39 1.9 13 L. Prickley Pear Cr. 69 358 195 21 5 1.0 14 16 Mile Cr. 50* 522 195 13 0 0 15 Bluewater Cr.- 28 1387 214 27 56 2.1 16 Bluewater Cr.^ 18 798 209 22 77 3.7 17 Flagstaff Cr. 5* 405 197 16 0 0 ♦Estimated by fisheries biologists of the Montana Departm en of Fish md Game. ^See Figur B 1 =Section below Bluewater Fish Hatchery ^Section above Bluewater Fish Hatchery measured and weighed, and scale samples were taken for age determinations. The ovaries were removed for an unrelated fecundity study (Lockard 1974) and con- sequently were not included in this nec- ropsy. Also, the heart and gas hladder were excluded because they were either mutilated or lost when the ovaries were removed. The procedure for necropsy was as follows. The gills and external body of the fish were examined for ecto- jjarasites and fluke metacercaria, although the preserving and handling procedures greatly reduced the chances of discovering j)arasitic copepods, leeches, or mono- genetic flukes. After removal of the re- maining internal organs, the liver was dis- sected into small pieces, placed in a jar 444 GREAT BASIN NATURALIST Vol. 35. No. 4 with water, and put on a mechanical shaker for 5-10 minutes. The contents were then washed onto a 200-mesh col- lecting screen and examined in an il- luminated tray (Barber and Lockard 1973). The gastrointestinal tract was opened and its component parts scraped and placed with their contents in jars of water. After 5-10 minutes on the me- chanical shaker, the contents of each jar were examined in the illuminated tray. The kidney and testes w'ere examined grossly, and observed abnormalities checked microscopically. Recovered nema- todes were placed in 70 percent alcohol-5 percent glycerine and later mounted in glycerine. Results Parasites were found throughout the upper digestive tract from the esophagus to the pyloric caeca. The examinations revealed that 45 percent of the 306 brown trout were infected with one to four genera of nematodes. Table 2 lists the occurrence of each taxon in a given stream. Identification of Cystidicoloides salvelini, Bulbodacnitis globosa, and Rhab- dochona sp. was confirmed by Dr. James R. Adams, University of British Columbia (personal correspondence) . The Rhabdo- chona sp. apparently is a new species and has been submitted to Dr. Morovec in Prague, Czechoslovakia, for further identification. Inspection of Table 1 indicated an ap- parent relationship between the size of stream discharges, the percent parasitism (incidence) in fish and average number of worins per parasitized fish (intensity). Regression analyses were performed to test these relationships. The ])ercentage of fish parasitized regressed on stream size was found to be correlated (R- = 0.46; T test P= 0.003). The average number of nematodes per parasitized fish regressed on stream discharge was found to be high- ly correlated (R— 0:68; . T test P= 0.00004) . The 306 fish used for this study ranged from 6.4 to 20.7 inches (16.3 to 52.6 cm) in total length and from 0.11 to 3.90 pounds (49.9 to 1769.0 grams) in weight with the mean length and weight being 12.3 inches (31.2 cm) and 0.93 pounds (421.8 grams). To test for a possible correlation between length of fish and in- tensity of parasitism, a regression of num- ber of nematodes per infected fish on fish length was made on fish from the three collections with the highest percent parasitism (Table 1). Fish from each stream were tested individually to com- pensate for varying intensities of parasit- ism and differences in mean size of fish between streams. There was no significant correlation at the 0.5 level between num- ber of parasites and length of host in fish from the Madison River, Big Hole River, and the West Gallatin River. The relationship of fish age to the in- cidence and intensity of parasitism is shown in Table 3. Differences in the per- cent of infected fish between age classes Table 2. Occurrence of nematode genera in brown trout from 16 Montana streams. Rhabdo- Bulbodac- Cystidi- Eustrong- ColL action site chona nitis coloides ylides 1 (Madison R.) X X X X 2 (Big Hole R.) X X X 3 (W. Gallatin R.) X X X 4 (St. Regis R.) X X 5 (Beaverhead R.) X X 6 (E. Gallatin R.) X X 7 (Rock Cr.) X X 8 (Shields R.) X 9 (L. Blackfoot R.) X 10 (O'Dell Cr.) X X 11 (So. F. Musselshell R.) X 12 (Baker Cr.) X X X 13 (L. Prickley Pear Cr.) X 14 (16 Mile Cr.) 15 (Bluewater Cr.)i X X 16 (Bluewater Cr)- X X 17 (Flagstaff Cr.) 'Section above Bluewater Fish Hatchery -Section below Bluewater Fish Hatchery Dec. 1975 LOCKARD, ET AL.: BROWN TROUT 445 Table 3. Incidence and intensity of parasitism in age classes of brown trout. Age class # Fish Infected Ave. # worms/ infected fish I 58 II 137 III+ 111 Total (ave.) 306 50 38 62 (45) 2.4 9.1 21.3 (12.8) were tested by a method of Arkin and Colton (1972) with results showing no significant differences between age class- es I and II, and I and III+ at the 0.05 level. Although the difference in the per- cent of fish infected in age classes II and III+ was statistically significant (P = 0.028) it was not considered to be bio- logically significant. As the age of the fish increased the average number of worms per infected fish also increased (Table 3). These differences in the in- tensity of parasitism between age classes shown in Table 3 were significant at the 0.05 level. The relationship of incidence and inten- sity of parasitism to age and sexual ma- turity in brown trout is given in Table 4. Significantly (0.05 level) more mature fish than immature fish in age classes I and II and in the overall total were in- fected. However, there was no significant difference between the incidence of para- sitism in mature and immature fish in age class III + . Mature fish in age classes II and III+ and the overall total also had a significant!}' higher number of worms per fish than immature fish. In age class I there were no significant differences in the average number of worms in mature and immature brown trout. Tests were made to determine the re- lationship of the chemical productivity of each stream with the degree of parasit- ism in its fish. No significant (0.05 level) correlations were found when percent parasitism and average number of worms per parasitized fish were regressed on stream conductivities and alkalinities. In the five streams (Big Hole River, Baker Creek, Little Prickley Pear Creek, Bluewater Creek, and Flagstaff Creek) from which the 48 males were collected, there was no significant difference be- tween their rate of infection (45.8 per- cent) and that of females from the same streams (48.2 percent). Discussion Incidence of parasitism in brown trout varies with species of parasite and with habitat of the host fish. Fox (1962) found 30 brown trout from Meadow Lake, Mon- tana, infected with fluke metacercaria of Bolbophorus confusus. Five of the 30 brown trout had immature stages of the nematode Eustrogylides sp. encysted with- in the body and 1 of the 30 had an un- identified nematode in its digestive tract. Van Cleave and Mueller (1934) examined 13 brown trout from Oneida Lake and 3 collected from Black Creek about one mile from its confluence with Oneida Lake, New York. The stream-inhabiting brown trout had the nematode Cystidi- coloides hardwoodi exclusively, whereas the lake-inhabiting brown trout had the nematode Spinitectus gracilis and never C. hardwoodi. In over 1,000 fish from the lake C. hardwoodi was never taken; thus it was probably limited to streams. In correlating parasitism to habitat. Van Cleave and Mueller (1934) conclude "that in fishes of wide range through a variety of environmental types, the type of parasitism is more or less closely cor- related with the habitat from which the fish is taken." Bangham (1951) states "the fact that there were not many dif- ferent types of habitats limited the num- bers of various parasite species." Stream habitat may not favor the intermediate hosts of many lake parasites, i.e., cestodes T.\BLE 4. Incidence and intensitj- of parasitism related to sexual maturity and age of brown trout. Ave. # worms/ Age # Fish Se X. Mat. "Ji Infected infected fish class Se X. mat. Sex. imm. Sex. mat. Sex. imm. I 58 19 82 43 2.1 2.5 II 137 51 47 28 11.0 5.9 III + 111 88 52 54 23.7 4.4 Total 306 (60) (53) (34) (17.1) (4.2) (Ave.) 446 GREAT BASIN NATURALIST Vol. 35, No. 4 (copepods), trematodes (snails), acantho- cephalans ( Crustacea). Parasitic cope- pods and monogenetic flukes not com- mon in streams may be favored by the lake environment. 1'his would partially explain wh}' nematodes were the only parasites found in these brown trout from cold water stream habitats. Hugghins (1959) found no parasitism in four brown trout from Black Hills streams in South Dakota. Van Cleave and Mueller (1934) classified the fre- quency of occurrence of six species of parasites in brown trout as "occasional" based on a scale of abundant, common, oc- casional, and rare. Forty-one percent of 58 brown trout from Wisconsin streams and ponds were parasitized (Bangham 1946, and Fischthal 1947a, 1947b, 1950, 1952). This is comparable to the 45 per- cent incidence of parasitism found in this 5tudy. The correlation of size of stream with incidence and intensity of parasitism was shown in Table 1. It can be seen from Table 2 that large streams have a greater diversity of species of nematodes than small streams. It would be expected in a large river with diverse habitat niches that the chance of intermediate and de- finitive hosts for a certain parasite exist- ing together would be greater than in a smaller stream with a more restricted type of habitat. Fox (1962) states that, in general, the longer the trout, the greater the number of individual parasites and kinds of para- sites it contained. This is related to longer exposure time to parasitism, since the body length is generally determined by age. Woodbury (1940) summarized 12 previous investigations and reported that the longer fish had greater numbers of parasites in nine of these studies. Con- versely, Langlois (1936) and Hubbs (1927) found shorter fish the most heavi- ly parasitized. In this study no correlation between fish length and intensit}^ of in- fection was found in fish from the three streams with 100 percent })arasitism. How- ever, the fish in these streams were not proportionately representative of all age classes; thus these results do not rule out a relationship between age and intensity of infection. This study did not show^ that older fish were generally more likely to be infected than younger fish (Table 3). However, it can be seen from Table 3 that as the age of the fish increases, so does the average number of worms per in- fected fish. This could be related to the older fish having more exposure time to })arasites than the younger ones do. The relationships of incidence and in- tensity of infection to feeding aggressive- ness of fish as expressed by sexual ma- turity was examined. A higher percentage of mature fish was parasitized than im- mature fish in the overall total and for age classes I and II (Table 4). The dif- ference between infection rates between mature and immature fish in the overall total could be attributed to older mature fish having more exposure time to the parasites than younger fish do. However, the differences in infection rates between mature and immature fish within age classes I and II could be the result of the fastest-growing (most-aggressive) fish in an age class reaching sexual maturity be- fore less-aggressive fish. Bagenal (1969) fed different quantities of food to two groups of brown trout and found that the more aggressive of the under-fed fish took more than their share of the food. Conse- quently, the more aggressive of the starved fish had growth comparable with that of the well-fed fish. He also found that more of the better-fed fish were ma- ture than the under-fed fish. There were no significant differences (0.05 level) be- tween mature and immature fish in the percent infected for age class III + . Niel- son (1953) noted that brown trout in Cali- fornia began a shift in feeding habits from strictly aquatic invertebrate forms to partial diets of fish in the third year of life. An explanation for the similar per- cent infections for mature and immature age class III+ fish (Table 4) could be this change in feeding habits in which the parasite intermediate hosts (mayflies) would be preyed upon less by the larger mature fish. McFadden, Cooper, and Anderson (1965) state that perhaps some combin- ation of chronological age and growth rate (or the factors which determine growth rate), a "physiological age", de- termines the age of sexual maturity. One factor which influences growth rate and thus age at sexual maturity is the feed- ing aggressiveness of a fish. If sexually mature fish feed more actively than sex- Dec. 1975 LOCKARD. ET AL.: BROWN TROUT 447 ually immature fish, then mature fish of a given age would be larger than the im- mature fish in that age class and would have more exposure to parasitism through their aggressive feeding habits. The fol- lowing is a comparison of mean lengths of mature and inmiature female brown trout within age classes. In age class I, only the Bluewater Creek collections con- tained mature females, and they had a mean length of 8.0 inches while the im- mature females from Bluewater Creek were 7.5 inches in mean length. Mature female fish from all streams of age class- es II and 111+ had mean lengths of 13.3 and 15.1 inches, respectively; while the immature females from these age classes had mean lengths of 11.0 and 12.9 inches, respectively. Thus sexually ma- ture fish v\dthin an age class are larger than sexualh' immature fish. This could be the result of more aggressive feeding habits. The most actively feeding fish would have more chances to encounter the intermediate hosts (mayflies, Hoff- man 1967) of these nematode parasites {Cystidicoloides sp. and Rhabdochona sp. were 99.4 percent of all worms found) and thus more of them would be infected. This theory to explain differences in infection rates between mature and im- mature fish within age classes is rein- forced by inspection of the average num- ber of worms per parasitized fish in age classes (Table 4). The mature fish in age classes II and III+ and for the overall total have a higher average number of worms per fish than the immature fish. This suggests that the more actively feed- ing mature fish within an age class are exposed more to parasitism by their feed- ing habits than less actively feeding im- mature fish. In age class I (Table 4) there was no significant difference (0.05 level) be- tween mature and immature brown trout in the average number of worms j)er in- fected fish, possibly because exjiosure time was too short for significant differences to be expressed . Fox (1962) found little relationship be- tween incidence of parasitism and con- dition of trout. He states that possibly ". . . analysis of fecundity will show that parasite incidence does effect some host conditions." Comparison of the findings of this study with those of a fecundit} study on the same fish (Lockard 1974) indicates that no general relationship be- tween parasitism and fecundity exists. Table 1 shows the Madison and West Gallatin rivers were highly parasitized; in the above fecundity study they had the highest fecundity (steepest slopes of the regressions of number of eggs on fish length for a stream) . Thus parasitism ap- parently did not adversely affect the fe- cundity of infected fish. Acknowledgments The authors express their appreciation to Dr. Da^'id E. Worley of Montana State University for providing laborator}" space and equipment, and to Dr. William R. Gould of Montana State University and Mr. George D. Holton of the Montana Department of Fish and Game for re- viewing the manuscript. Bibliography Alexander, C. G. 1961. A survey of parasites of Oregon trout. Report to Oregon State Game Commission. 34 pp. Arkin, H.. .-^nd R. R. Colton. 1970. Statisti- cal metliods. Barnes and Noble Books, New York. 344 pp. Bagen.\l, T. B. i1969. The relationship between food supply and fecundity in brown trout Salmo trutta. L. J. Fish Biol. (1969) 1:167- 182. Bangham, R. V. 1946. Parasites of northwest Wisconsin fishes. Trans. Wisconsin Acad. Sci.. Arts and Lett. 36:291-325. . 1951. Parasites of fish in the upper Snake River drainage and in Yellowstone Lake, Wyoming. Zoologica, Scient. Contrib., New York Zool. Soc. 36(3) :213-217. Barber. D. L.. .vnd L. L. Lockard. 1973. Some helminths from mink in southwestern Mon- tana, with a checklist of their internal para- sites. Great Basin Nat. 33(l):53-60. FiscHTHAL. J. H. 1947a. Parasites of northwest Wisconsin fishes. L The 1944 survey. Trans. Wisconsin Acad. Sci. 37:157-220. . 1947b. Parasites of Brule River fishes. Brule River survey: Rep. no. 6. Tians. Wis- consin Acad. Sci. 37:275-278. . 1950. Parasites of northwest Wiscon- sin fishes IL The 1945 survey. Trans. Wis- consin Acad. Sci. 49(1) :87-l 13. 1952. Parasites of northwest Wiscon- sin fishes in. The 1946 sui-\ey. Trans. Wis- consin Acad. Sci. 41:17-58. Fox. A. 4962. Parasite incidence in relation to size and condition of trout from two Mon- tana lakes. Trans. Amer. Micr. Soc. 81(2): 179-184. Hec;kmann. R. A. 1971. Blood parasitism of some fishes fioni Montana and Yellowstone National Park. I. Wildlife Diseases (1971) 7:3-4. 448 GREAT BASIN NATURALIST Vol. 35, No. 4 Hoffman, G. L. 1967. Parasites of North American freshwater fishes. Univ. Calif. Press. Berkelej- and Los Angeles. 486 pp. HuBBS, C. L. 1927. The related effects of a parasite on a fish. J. Parasitol. 14:75-84. HuGGHiNs. E. J. 1972. Parasites of fishes in South Dakota. South Dakota Exper. Sta. Bull. 484:1-73. L.'^NGLois. T. H. 1936. Bass tapeworm infec- tion in a rearing pond. Trans. Amer. Fish Soc. 66:364-366. LocK.\RD, L. L. 1974. Some environmental in- fluences on the egg production in brown trout {Salmo irutta) from Montana streams. M. S. thesis. Montana State Univ. 28 pp. McFadden, J. T., E. L. Cooper, and J. K. An- derson. 1965. Some effects of environment on egg production in brown trout {Salmo trutta). Limn, and Ocean. 10(l):88-95. NiELSoN. R. S. 1953. Should we stock brown trout? Prog. Fish Cult. 15:125-126. Parker, R. R. 1963. Effects of formalin on length and weight of fishes. J. Fish. Res. Bd. Canada 20(6) : 1441-1455. United States Geological Survey. 1972. Water resources data for Montana. Part 2. Water quality records. 218 pp. Van Cleave, H. J., and J. F. Mueller. 1934. Parasites of Oneida Lake fishes. Part IIL A biological and ecological survey of the worm parasites. Roosevelt Wildlife Ann. 3(3&4): 161-334. Vincent. E. R. il967. Southwest Montana fishery study-evaluation of river fish popu- lations. D. J. Completion Report. Project F-9-R-15. Mont. Fish and Game Dept. 15 pp. Woodbury, L. A. 1940. A quantitative study of parasites of fishes with special reference to Clinostomum marginatum in the perch of Walsh Lake, Michigan. Ph.D. thesis. Univ. Mich., Ann Arbor. SEXUAL DIMORPHISM IN MALPIGHIAN TUBULES OF PTERONARCYS CALIFORNICA NEWPORT (PLECOPTERA) Ralph R. Hathaway 1 Abstr.'VCt. — The Malpighian tubules in female nymphs and adults of Pteronarcys californica New- port are larger and whiter than in males. This difference is detectable in nymphs as small as 300 mg in live weight (final instar nymphs weigh 800 to 1600 mg depending on sex), and is most pro- nounced in late nymphs and adults. These differences are thought to be related to the increased ex- cretory load imposed by the synthesis of ooplasm in the late female nympli and adult. The excretory systems of Plecoptera reflect the relatively primitive taxonomic level of this order. The Malpighian tu- bules are usually numerous and un- branched and have an uncomplicated re- lationship to the gut, joining it at the level where the midgut empties into the hind- gut. Apart from this connection, the tu- bules are mainly free and unattached, ex- tending throughout the body cavity. This situation is seen with textbook clarity in Pteronarcys californica. Much current interest in the Malpi- ghian tubules centers on their ultra struc- ture and on their function as a homeo- static organ (Berridge and Oschman 1969; Phillips and Maddrell 1974). They are involved in the elimination of salts and amines, and in water balance. Most studies have been done on terrestrial forms in which water retention and ion transport are achieved in specializations of the tubules and, at times, inclusion of tubules in the wall of the hind gut. The only studies on excretion in Plecoptera are those of Colby (1972), who used in vivo measurements to demonstrate the formation of a dilute urine in the aquatic nymphs of Pteronarcys californica. There have been no detailed morphological stud- ies of Plecopteran Malpighian tubules. In a study of the growth of internal organs (Branham and Hathaway 1975), I have observed that there is a distinct sexual dimorphism in the appearance of Malpighian tubules in P. californica, which has not to my knowledge been reported earlier. Methods Stoneflies were collected in the Provo River in Utah and transported alive to the laboratory. Several hundred fresh and preserved animals from all size classes ^Biology Department, University of Utah, Salt Lake City, Utah were examined. The dimorphisms were generally well preserved in animals fixed in 70 percent ethanol and other killing and fixing fluids. A few animals were used for histological preparations in which 10/t sections were stained with Delafields hematoxylin. Weights of live animals were taken on an analytical balance after thorough blotting to remove excess water. Observations Male nymphs in the final instars weigh, on the average, about 800 mg, whereas the corresponding female nymphs are about twice as big, or 1600 mg. The Mal- pighian tubules in the female nymphs in all size classes above 300 mg are larger and whiter than those in the males. This difference becomes more pronounced as the animals grow larger and is at its maxi- mum in the final instars and adults (com- pare Figs, la & c). The differences are not obvious in animals weighing less than 300 mg. It is at about 300 mg of body weight that the body-size differences between male and female nymphs becomes dis- cernable; therefore the onset of differ- ences in the Malpighian tubules corre- sponds with this event. It is not merely size, however, that is involved in the sex- ual dimorphism in the Malpighian tu- bules. In females the tubules are boldly conspicuous because of their intense whiteness, whereas the male tubules are at first difficult to find because of their translucent, colorless or pink-tinged ap- pearance. The female tubules, moreover, are of more uniform diameter throughout their length, while the diameter of male tubules is irregular. Females that are smaller and at least a year yoimger than final instar males still have tubules that 449 450 GREAT BASIN NATURALIST Vol. 35, No. 4 are larger and whiter (compare Figs, la &b). About 20 percent of the tubules in ani- mals of both sexes are connected ante- riorly to the midgut by their distal ends (Fig. la). This union is formed from a delicate strand of solid connective tissue, so there is no confluence at this point be- Fig. 1: Malpighian tubules in Pteronarcys californica: (a) Male final-year njrmph. Live wt. = 840 mg. Isolated gut with anterior end down. Some of the tubules are anchored to the gut by their dis- tal ends. Scale bar equals 2 mm. (b) Female penultimate-year nymph. Live wt. = 620 mg. Isolated gut with anterior end down. Tubules are larger and whiter than in (a). Anchored distal ends of some of the tubules are again visible. Magnification same as in (a), (c) Female final-year nymph. Live wt. = 1560 mg. Isolated gut with anterior end down. Magnification same as in (a) and (b). (d) Fe- male final-year n>-mph. Cross sections of tubules. Scale bar = 0.1 mm. (e) Male final-year nymph. Cross sections of tubules. Magnification is same as in (d). Dec. 1975 HATHAWAY: SEXUAL DIMORPHISM IN PLECOPTERA 451 tween the lumina of the gut and the tu- bules. The lumina of these tubules drain into the gut at the same level as all the other tubules, i.e. at the point of junction between the midgut and the hindgut. Be- tween their proximal and distal ends, these tubules are totally free of the gut wall; the only obvious difference from the majority of the Malpighian tubules is that their distal ends are anchored to the midgut at a point about two-thirds of the way from its anterior end, while the dis- tal ends of the other tubules are free. Discussion The differences described above must have a functional basis relating to sex. Gonadal development can be seen in both sexes as they begin their final year of nymphal life. At this time males weigh about 500 mg and females weigh 900 to 1000 mg. Gonaflal enlargement and sperm production are initiated early in the male; maximum gonadal size is reached midwa\' through the final year. Corresponding events in the female do not occur until the final six or eight weeks of nymphal life. During this period a large part of metabolic effort in females is directed toward the synthesis of oo- plasm. There is evidence that oogene- sis utilizes substances from the fat body (Branham and Hathaway 1972). This effort continues into the adult stage, in which gamete production in the ovaries can take place for at least four weeks. Thus, one of the more obvious explana- tions for sexual differences in Malpighian tubules is that the redirection of metabo- lism associated with egg production re- el uires more elaborate excretory organs. Another ])ossibility is that the Malpi- ghian tubules in the female are involved in a secondary function such as the stor- age of energy-rich substances. The pri- mary storage organ is the fat body, but this decreases in size during the time of egg production. Thus the tubules may serve to store additional reserves to be utilized during the non-feeding late- nymph and adult stages. It seems likely that the observations re- corded here have some general signifi- cance, especially in Plecoptera. Sexual di- morphism in Malpighian tubules also oc- curs in Pteronarcella hadia (Pteronarci- dae), which differs in many life historical respects from P. calif ornica. Further ob- servations on other species would prob- ably be rewarding. Literature Cited BeRIUDGE. M. .].. AND ,1. L. OSCHMAN. 1969. A structural basis for fluid secretion by Mal- pigbian tubules. Tissue and Cell 1:247-272. BiiANHAM, .r. M.. AND R. R. Hathaw.xy. 1975. Sexual differences in the growth of Pteron- arcys californica Newport and Pteronarcella badia (Hagen) (Plecoptera). Can. J. Zool. 5^:501-506. Coi.BY. C. 1972. Salt and water ijalance in stoneflv naiads. Pteronarcys californica New- j)ort. Comp. Biochem. Physiol. 41:851-860. Piiii.LiPS, .J. E., AND S. H. P. Maddrell. 1974. Active transport of magnesium by the Mal- pighian tubules of the larvae of the mosquito, Aedes campeslris. J. Exp. Biol. 61:76-771. NEW RECORD OF THE BAT PLECOTUS PHYLLOTIS FROM UTAH Richard M. Poche' Abstract. — In June 1974 a scrotal male Mexican big-eared bat [Plecotus phyllotis) was netted near Hurricane. Utah. This account represents the third report of the species from Utah and e.x- tends the range approximately 330 km west of previous accounts. Adjacent state records are located approximately 64 km to the southeast in Arizona and 80 km southwest in Nevada. The Mexican big-eared bat has been re- ported from Utah by Black (1970) and Armstrong (1974). Both records were from San Juan County about 8 km N Blanding and 65 km NW Monticello, re- spectively. A specimen was reported from Pipe Springs National Monument area in Arizona by Genoways and Jones (1967 ). and one from the Mesquite, Neva- da, region by M. J. O'Farrell (personal communication). On 25 June 1974 a scrotal male Mexi- can big-eared bat was obtained in <; mist net over Gould Wash, NW ^4, SW 14, sec. 19, T. 42 S, R. 12 W, Washington Co., Utah. The locality lies approximateh' 7 air km southeast of the town of Hurri- cane. The capture of this speci.men ex- tends the range in Utah approximately 330 km to the west and represents only the third report from that state. The Arizona report is from an area about 64 km southeast of Hurricane and the Ne- vada account 80 km to the southwest. General similarity in habitat in the area between Hurricane and the southeastern reports suggests that P. phyllotis probably inhabits most of southern Utah. The Mexican big-eared bat obtained near Hurricane, Utah, was netted over one of three small ponds remaining in the wash, striking the net approximately 1 m above the surface of the water be- tween 0100 and 0530 hours. It was ex- amined for ectoparasites, of which there were none, marked (with a 3 mm hole in the plagiopatagium, near the pollex, between the 4th and 5th phalanges of the right wing), and released. The bat flew west along the wash and disap- 'School o{ Forcsty & Cnnscrvnlinn. University of Cnlifom i peared into the canyon. Six male scrotal Pipistrellus hesperus also were captured in the same net and were released. Gould Wash drains into the Virgin River 14 km northwest of the netting site. Vegetation in the wash includes Tamarix and Salix. and that on the ad- jacent upland is comprised primarily of blackbrush (Coleogyne ramosissima) , and infrequent pinon pine {Pinus edulis) and juniper [Juniperus osteosperma) . Grass cover is predominantly Bromus and Hi- laria. Average elevation for the area is 1,250 m with annual precipitation less than 200 mm. The Hurricane Cliffs are located 5 km west of the collection area and descend over 500 m to Warner Val- ley, which is covered with creosotebush {Larrea tridentata) , blackbrush, and snakeweed {Gutierrezia microcephala) . The surrounding region consists of num- erous steep rises and mesas, canyons, and cliffs. I thank Geoff Baillie for assistance with netting activities. David Armstrong pro- vided helpful comments in reviewing this paper. This study was financed by the Nevada Power Company. LiTF,R.'\TURE Cited Armstrong. D. A. 1974. Second record of the Mexican big-eared bat in Utah. Southwest- ern Nat. 19(1):114-115. Black. H. L. 1970. Occurrence of the Mexican big-eared bat in l^tah. J. Mammal. 15(1); 190. Genoways. H. H.. and J. K. Jones. Jr. 1967. Notes on distribution and variation in the Mexican big-eared bat. Plecotus phyllotis. Southwestern Nat.. 12:477-480, Beikcle 047nj, 452 INDEX TO VOLUME 35 The genera and species described as new to science in tiiis volume appear in bold type in this index. Additional records of reptiles from Jalisco, Mexico, p. 317. Allred, Dorald M., article by, p. 405. Amphicranus parilis, p. 31. Andersen, Ferron L., and Paul R. Roper, article by, p. 203. A new combination of Penstemon (Scrophu- lariaceae), p. 434. Arachnids as ecological indicators, p. 405. Araptiis attenuatus, p. 30. Araptus consobrinus, p. 394. Araptus fossifrons, p. 30. Araptus micaceus, p. 395. A revision of the nearctic species of Clino- helea Kieffer (Diptera: Ceratopogonidae), p. 275. A revision of the Phacelia Crenulatae group ( Hydrophyllaceae ) for North America, p. 127. Arndt, Rudolf G., Philip A. Medica, and James R. Dixon, article by, p. 317. A systematic study of Coenia and Paracoenia (Diptera: Ephydridae), p. 65. Atwood, N. Duane, article by, p. 127. Atwood, N. Duane, Stanley L. Welsh, and James L. Reveal, article by, p. 327. Avery, David F., Charles Fanghella, and Wil- mer W. Tanner, artcile by, p. 245. Baker, M. F., D. W. Nichols, and H. D. Smith, article by, p. 191. Basidomycetes that decay junipers in Arizona, p. 288. Bleich, Vernon C, and Orlando A. Schwartz, article by, p. 62. Bock, Carl E., and Larry W. Lepthien, article by, p. 269. Body size, organ size, and sex ratios in adult and yearling Belding ground squirrels, p. 305. Bombylius abdominalis, p. 414. Bombylius aestivus, p. 416. Bombylius auriferoides, p. 417. Bombylius heximaculatus, p. 410. Bombylius lancifer kanabensis, p. 412. Bombylius lassenensis, p. 416. Bombylius montanus, p. 413. Bombylius nigriventris, p. 410. Borchert, Mark, and Donald H. Owings, p. 402. Breeding range expansion of the starling in Utah, p. 419. Branham, Joseph M., Arden R. Gaufin, and Robbin L. Traver, article by, p. 51. Calocoenia (subgenus), p. 78. Campbell, R. B., W. T. McDonough, and R. 0. Harniss, article by, p. 325. Gather, Mary R., and Arden R. Gaufin, article by, p. 39. Gather, Mary R., Bill P. Stark, and Arden R. Gaufin, article by, p. 49. Clarence Cottam, 1899-1974, a distinguished alumnus of Brigham Young University, p. 231. Clark, Stephen L., article by, p. 434. Cliriohelea pseudonubifera, p. 280. Cnesinus electus, p. 23. Cnesinus pilatus, p. 24. Coenia alpina, p. 83. Computerized reduction of meteorologic measurements from irrigated and nonirri- gated plots in central Utah, p. 203. Correlates of burrow location in Beechey ground squirrels, p. 402. Corthylus cecropil, p. 31. Cymopterus higginsii, p. 377. Dacnophthorus, p. 394. Distribution and abundance of the black-billed magpie {Pica pica) in North America, p. 269. Dixon, James R., Philip A. Medica, and Rudolf G. Arndt, article by, p. 317. Endangered, threatened, extinct, endemic, and rare or restricted Utah plants, p. 327. Environmental factors in relation to the salt content of Salicornia pacifica var. utahensis, p. 86. Evans, Howard E., article by, p. 123. Evolutionary divergence in closely related pop- ulations of Mimulus guttatus (Scrophulari- aceae), p. 240. Evolution of the sceloporine lizards (Iguani- dae), p. 1. Fanghella, Charles, David F. Avery, and Wil- mer W. Tanner, article by, p. 245. Gallup, John S., and Martin L. Morton, article by, p. 427. Gaufin, Arden R., and Mary R. Gather, article by, p. 39. Gaufin, Arden R., Bill P. Stark, and Theodore A. Wolff, article by, p. 97. Gaufin, Arden R., Joseph M. Branham, and Robbin L. Traver, article by, p. 51. Genetics, environment, and subspecies dif- ferences: The case of Polites sabuleti, p. 33. Gilbertson, R. L., and J. P. Lindsey, article by, p. 288. Grogan, WilHam L., Jr., and Willis W. Wirth, article by, p. 275. Growth of Plecoptera (stonefly) nymphs at constant, abnormally high temperature, p. 51. 453 454 GREAT BASIN NATURALIST Vol. 35, No. 4 Guibe, Jean, Hobart M. Smith, and Rozella B. Smith, article by, p. 109. Hansen, D. J., and D. J. Weber, article by, p. 86. Harniss, R. 0., W. T. McDonough, and R. B. Campbell, article by, p. 325. Hathaway, Ralph R., article by, p. 449. Higgins, Harold G., and Tyler A. Woolley, article by, p. 103. Hughes, Karen W., and Robert K. Vickery, Jr., article by, p. 240. Hylastes asperatus, p. 24. Invasion of big sagebrush (Artemesia triden- tata) by white fir {Abies concolor) on the southeastern slopes of the Warner Mountains, California, p. 319. Ips pilifrons thatcheri, p. 29. Johnson, D. Elmer, and Lucile Maughan Johnson, article by, p. 407. Johnson, Lucile Maughan, and D. Elmer John- son, article by, p. 407. Larsen, Kenneth R., and Wilmer W. Tanner, article by, p. 1. Lepthien, Larry W., and Carl E. Bock, article by, p. 269. Leptocoenia (subgenus), p. 81. Life history and ecology of Megarcys signata (Plecoptera: Perlodidae), Mill Creek, Wa- satch Mountains, Utah, p. 39. Lindsey, J. P., and R. L. Gilbertson, article by, p. 288. Lockard, Lawrence L., article by, p. 435. Lockard, Lawrence L., and R. Randall Par- sons, article by, p. 425. Lockard, Lawrence L., R. Randall Parsons, and Barry M. Schaplow, article by, p. 442. Mathis, Wayne N., article by, p. 65. McDonough, W. T., R. 0. Harniss, and R. B. Campbell, article by, p. 325. Medica, Philip A., Rudolf G. Arndt, and James R. Dixon, article by, p. 317. Morphology of ephemeral and persistent leaves of three subspecies of big sagebrush grown in a uniform environment, p. 325. Morton, Martin L., and John S. Gallup, article by, p. 427. Morton, Martin L., and Robert J. Parmer, article by, p. 305. Multoribates haydeni, p. 104. Nectar composition of hawkmoth-visited spe- cies of Oenothera (Onagraceae), p. 273. New mites from the Yampa Valley (Acarina: Cryptostigmata: Oribatulidae, Passalozeti- dae), p. 103. New record of the bat Plecotus phyllotis from Utah. p. 452. New records of stoneflies (Plecoptera) from New Mexico, p. 97. New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), p. 21. New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part n, p. 391. Nichols, D. W., H. D. Smith, and M. F. Baker, article by, p. 191. Notes on the genus Bombylius Linnaeus in in Utah, with key and descriptions of new species (Diptera: Bombyliidae), p. 407. Owings, Donald H., and Mark Borchert, p. 402. Paracoenia iParacoenia) ampla, p. 71. Paracoenia {Paracoenia) calida, p. 73. Paracoenia {Paracoenia) wirthi, p. 78. Parahauloppia cordylinosa, p. 105. Parsons, R. Randall, and Lawrence L. Lock- ard, article by, p. 425. Parsons, R. Randall, Lawrence L. Lockard, and Barry M. Schaplow, article by, p. 442. Parmer, Robert J., and Martin L. Morton, article by, p. 305. Passalozetes moniles, p. 106. Penstemon atwoodii, p. 378. Phacelia crenulata var. angustifolia, p. 158. Photoperiodic responses of phenologically aberrant populations of pierid butterflies (Lepidoptera), p. 310. Pityokteines mystacinus, p. 29. Pityophthorus amiculus, p. 398. Pityophthorus costatus, p. 395. Pityophthorus degener, p. 397. Pityophthorus dissolutus, p. 398. Pityophthorus explicitus, p. 399. Pityophthorus inceptis, p. 396. Pityophthorus mendosus, p. 397. Pityophthorus timidulus, p. 396. Poche, Richard M., article by, p. 452. Poche, Richard M., and George A. Ruffner, article by, p. 121. Provonsha, A. V., article by, p. 379. Pseudothysanoes concentralis, p. 27. Pseudothysanoes tumidulus, p. 28. Records of stoneflies (Plecoptera) from Ne- vada, p. 49. Rreproductive cycle of the Belding ground squirrel {Spemophilus beldingi beldingi): seasonal and age differences, p. 427. Reveal, James L., Stanley L. Welsh, and N. Duane Atwood, article by, p. 327. Rodent populations, biomass, and community relationships in Artemisia tridentata. Rush Valley, Utah, p. 191. Roosting behavior of male Euderma macu- latum from Utah, p. 121. Roper, Paul R., and Ferron L. Andersen, article by, p. 203. Ruffner, George A., and Richard M. Poche, article by, p. 121. Sawin, H. Lewis, Hobart M. Smith, and Ro- zella B. Smith, article by, p. 100. Schaplow, Barry M., Lawrence L. Lockard, and R. Randall Parsons, article by, p. 442. Schwartz, Orlando A., and Vernon C. Bleich, article by, p. 62. Sexual dimorphism in malpighaian tubules of Pteronarcys califomica Newport (Plecop- tera), p. 449. Scolytodes amabilis, p. 26. Dec. 1975 455 Scolytodes genialis, p. 27. Scolytodes lepidus, p. 27. Scolytodes obesus, p. 26. Scolytus laetus, p. 25. Scolytus torulus, p. 25. Shapiro, Arthur M., article by, p. 33, 310. Smith, Dwight G., article by, p. 419. Smith, H. D., D. W. Nichols, and M. F. Baker, article by, p. 191. Smith, Hobart M., Rozella B. Smith, and H. Lewis Sawin, article by, p. 100. Smith, Hobart M., Rozella B. Smith, and Jean Guibe, article by, p. 109. Smith, Rozella B., Hobart M. Smith, and H. Lewis Sawin, article by, p. 100. Smith, Rozella B., Hobart M. Smith, and Jean Guibe, article by, p. 109. Some parasites of paddlefish (Polydon spat- hula) from the Yellowstone River, Montana, p. 425. Some relationships between internal parasites and brown trout from Montana streams, p. 442. Some relationships between water fertility and egg production in brown trout {Salmo trutta) from Montana streams, p. 435. Stark, Bill P., Mary R. Gather, and Arden R. Gaufin, article by, p. 49. Stark, Bill P., Theodore A. Wolff, and Arden R. Galfin, article by, p. 97. Stockhouse, Robert E., II., article by, p. 273. Studies in nearctic desert sand dune Orthop- tera. Part XV. Eremogeography of Spini- acris with bioecological notes, p. 113. Tanner, Vasco M., article by, p. 231. Tanner, Wilmer W., Charles Fanghella, and David F. Avery, article by, p. 245. Tanner, Wilmer W., and Kenneth R. Larsen, article by, p. 1. The authorship and date of publication of Siren intermedia (Amphibia: Caudata), p. 100. The identity of Boucourt's lizard Eumeces capita 1879, p. 109. The nest and larva of Diploplectron brunneipes (Cresson) (Hymenoptera: Sphecidae), p. 127. The Zygoptera (Odonata) of Utah with notes on their biology, p. 379. Thysanoes tuberculatus, p. 29. Tinkham, Ernest R., article by, p. 113. Traver, Robbin L., Joseph M. Branham, and Arden R. Gaufin, article by, p. 51. Utah plant novelties in Cymopterus and Pens- temon, p. 377. Urosaurus and its phylogenetic relationship to Uta as determined by osteology and myology (Reptilia: Iguanidae), p. 245. Vale, Thomas R., article by, p. 319. Vickery, Robert K., Jr., and Karen W. Hughes, article by, p. 240. Water balance and fluid consumption in the southern grasshopper mouse, Onychomys torridus, p. 62. Weber, D. J., and D. J. Hansen, article by, p. 86. Welsh, Stanley L., article by, p. 377. Welsh, Stanley L., N. Duane Atwood, and James L. Reveal, article by, p. 327. Wirth, Willis W., and William L. Grogan, Jr., article by, p. 275. Wolff, Theodore A., Bill P. Stark, and Arden R. Gaufin, article by, p. 97. Wood, Stephen L., articles by, p. 21, 391. Woolley, Tyler A., and Harold G. Higgins, article by, p. 103. Xyleborus californicus, p. 399. Xyleborus incultus, p. 400. Xyleborus molestulus, p. 400. Xyleborus tristiculus, p. 401. Zygoribatula apletosa, p. 103. 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 directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. Manuscripts. Two copies are required, typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. 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No re- prints are furnished free of charge. A price list for reprints and an order form are sent with the proof. Reprint Schedule of the Great Basin Naturalist 2pp. 4pp. 6pp. 8pp. 10pp. 12pp. Each 100 copies $20 $23 $26 $29 $32 $35. Additional 200 copies 26 29 32 35 38 41 2pp. 300 copies 32 35 38 41 44 47 $3 TABLE OF CONTENTS Endangered, threatened, extinct, endemic, and rare or lestricted Utah vas- cular plants. Stanley L. Welsh, N. Duane Atwood. and James L. Reveal 327 Utah plant novelties in Cymopterus and Pensternon. Stanley L. Welsh 377 The Zygoptera (Odonata) of Utah with notes on their biology. A. Provon- sha 379 New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part II. Stephen L. Wood 391 Correlates of burrow location in Beechey ground squirrels. Donald H. Owings and Mark Borchert 402 Arachnids as ecological indicators. Dorald M. Allred 405 Notes on the genus Bombylius Linnaeus in Utah, with key and descrip- tions of new species (Diptera: Bombyliidae). D. Elmer Johnson and Lucile Maughan Johnson 407 Breeding range expansion of the starling in Utah. Dwight G. Smith 419 Some parasites of paddlefish (Polydon spathula) from the Yellowstone River, Montana. Lawrence L. Lockard and R. Randall Parsons 425 Reproductive cycle of the Belding gorund squirrel (Spermophilus beldingi): seasonal and age differences. Martin L. Morton and John S. Gallup .... 427 A new combination in Pensternon (Scrophulariaceae). Stephen L. Clark .... 434 Some relationships between water fertility and egg production in brown trout (Salmo trutta) from Montana streams. Lawrence L. Lockard 435 Some relationships between internal parasites and brown trout from Mon- tana streams. Lawrence L. Lockard, R. Randall Parsons, and Barry M. Schaplow 442 Sexual dimorphism in malpighian tubules of Pteronarcys californica New- port (Plecoptera). Ralph R. Hathaway 449 New records of the bat Plecotus phyllotis from Utah. Richard M. Poche .... 452 Index to Volume 35 453 '""" ' "Illll 3 2044 072 231 053 Date Due