HARVARD UNIVERSITY LIBRARY OF THE Museum of Comparative Zoology The Great Basin Naturalist VOLUME 33, 1973 Editor: Stephen L. Wood Published at Brigham Young University, by Brigham Young University TABLE OF CONTENTS Volume 33 Number 1 — March 31, 1973 Helminths of Sceloporus lizards in the Great Basin and upper Colorado plateau of Utah. Richard C. Pearce and Wilmer W. Tanner 1 Miscellaneous chromosome counts of western American Plants — II. James L. Reveal and Eloise L. Styer 19 Incidence of spotted fever in wood ticks of Utah recreational sites. C. Selby Herrin 26 Notes on the nesting behavior of Steniolia elegans (Hy- menoptera: Sphecidae). Howard E. Evans 29 A Taxonomic revision of Physaria (Cruciferae) in Utah. Sheldon B. Waite 31 Two new species of Gymnodamaeus from Colorado (Aca- rina: Cryptostigmata, Gymnodamaeidae). Tyler A. Woolley and Harold G. Higgins 37 Neartic desert Decticidae (Orthoptera). Part II. A new genus and species from Arizona. Ernest R. Tinkham 43 An unusual population of spiders in Utah. Dorald M. Allred .... 51 Some helminths from mink in southwestern Montana, with a checklist of their internal parasites. Delbert L. Bar- ber and Lawrence L. Lockard 53 Number 2 — June 30, 1973 Three new species of Palmoxylon from the Eocene Green River formation, Wyoming. William D. Tidwell, David A. Medlyn, and Gregory F. Thayn 61 On the taxonomic status of Platypodidae and Scolytidae (Coleoptera). Stephen L. Wood 77 Studies on Utah stoneflies (Plecoptera). Richard W. Bau- mann 91 The male brachycistidine wasps of the Nevada Test Site (Hymenoptera: Tiphiidae). Marius S. Wasbauer 109 Notes on aquatic and semiaquatic Hemiptera from the southwestern United States (Insecta: Hemiptera). John T. Polhemus 113 A new subfruticose Eriogonuni (Polygonaceae) from west- ern Colorado. James L. Reveal 120 Records of Coreidae (Hemiptera) from the Nevada test site. Dorald M. Allred 123 Density, growth, and home range of the lizard Uta Stans- buriana stejnegeri in southern Dona Ana County, New Mexico. Richard D. Worthington and Edward R. Arviso 124 Small bones of the hypsilophodontid dinosaur Dryosaurus alius from the upper Jurassic of Colorado. Peter M. Galton and James A. Jensen 129 Number 3 — September 30, 1973 Ecology of Sceloporus magister at the Nevada Test Site, Nye County, Nevada. Wilmer W. Tanner and John E. Krogh 133 Further studies on the wasps of Jackson Hole, Wyoming (Hymenoptera, Aculeata). Howard E. Evans 147 Additional records of mutillid wasps from the Nevada Test Site. Dorald M. Allred 156 The effects of soil texture on species diversity in an arid grassland of the eastern Great Basin. John W. Wyckoff 163 New synonymy in American bark beetles (Scolytidae: Coleoptera). Part III. Stephen L. Wood 169 Undescribed species of Neartic Tipulidae (Diptera). XI. Charles P. Alexander 189 Neartic desert Decticidae (Orthoptera). Part III: The true tjrmpanum in certain genera, with key. Ernest R. Tinkham 197 Notes on reproduction in Lampropeltis triangulum and Coluber constrictor in Utah. William L. Grogan and n Lloyd C. Pack, Jr 202 Courtship behavior among white-tailed and black-tailed jackrabbits. Del F. Blackburn 203 Number 4 — December 31, 1973 Local distribution and interspecies interactions in micro- tines. Grand Teton National Park, Wyoming. Tim W. Clark 205 Notes on the occurrence and distribution of Pteronarcys calif ornica Newport (Plecoptera) within streams. John A. Elder and Arden R. Gaufin 218 Prehistoric bighorn sheep in the northern Sierra Nevada, California. Peter D. Schulz and Dwight D. Simons 221 Artemesia arbuscula, A. longiloba, and A. nova habitat types in northern Nevada. B. Zamora and P. T. Tueller .... 225 Significant bird records from Utah. William H. Behle 243 Small mammals of the National Reactor Testing Station, Idaho. Dorald M. Allred 246 Scorpions of the National Reactor Testing Station, Idaho. Dorald M. Allred 251 Density changes and habitat affinities of rodents of shad- scale and sagebrush associations. Earl J. Larrison and Donald R. Johnson 255 New species of American Microcorthylus (Coleoptera: Scolytidae). Stephen L. Wood 265 The nominal snake genera Mastigodryas Amoral, 1834, and Dryadophis Stuart, 1839. Hobart M. Smith and Ken- neth R. Larsen 276 Vokune 33, No. 1 March 31, 1973 The iARVARD Great Basin Published by Brigham Young University GREAT BASIN NATURAUST Editor-. Stephen L. Wood, Department of Zk)ology, Brigham Young UniversitT, Provo, Utah >. I ixuey L. Welsh, Botany, Chairman; Wilmer W. r, Zoology; Joseph R. Murdock, Botany; Vernon J. 1 ipton, Zoology; Ferron L. Andersen, Zoology Ex officio Editorial Board Members: A. Lester Allen, Dean, College of Biological and AgricuJ- tural Sciences Ernest L. Olson, Director, University Press, University Editor Tlic Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been continuously published from one to four times a yeor since then by Brigham Young University, Provo, Utah. In !j:{\uora], only original, previously impublished manuscripts pertain- ing to Xho biological, natural history of the Great Basin and western North Ariierica will be accepted. Manuscripts are subiect to the ap- ! roval of the editor. ipTiON: The annual subscription is $5.00 (outside the lies $5.50). The piice for single numbers is $2.00 each. Iters pertaining to the purchase of subscriptions and back Jinnl,^ be directed to Brigham Young University Press, 205 UPB, Provo, Utah 84601. Libraries or other ' d in obtaining this journal through a contin- ;irly publications should contact the Brigham rian. The Library, Provo, Utah y for the Greet to ttie editor. Contributors i>n tlie back cover of a recent The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University Volume 33 March 31, 1973 No. 1 HELMINTHS OF SCELOPORUS LIZARDS IN THE GREAT BASIN AND UPPER COLORADO PLATEAU OF UTAH Richard C. Pearce^ and Wilmer W. Tanner^ Abstract. — Fifty-five Sceloporus lizards representing the four species oc- curring in Utah were dissected. For each lizard specimen and species, helminth parasites were tabulated and classified. New host and locality records were recorded. An appendix containing literature citations of reptilian parasitology for the American Southwest is provided. The literature dealing with the parasitic fauna of the south- western United States contains little information on helminths of Sceloporus lizards. Previous studies are concerned more with a single genus or species of parasite than with the total parasitic fauna of a particular host. Even more scarce are studies of parasitic fauna from hosts in defined geographical areas. Literature on reptilian parasites from Sceloporus lizards of the Great Basin and Upper Colorado Plateau of Utah is sparse and scat- tered. Grimdmann (1957) examined 47 lizards belonging to six Utah species, one of which was S. occidentalis; he reported the para- sites to be host specific. Grundmann (1959) later reported specific names of hosts and parasites, and locations of parasites within the host. Woodbury (1934) studied three species of Utah reptiles, in- cluding S. graciosus. He recovered only one species of nematode, probably because he studied only the stomach. Burkholder (personal communication) examined over 300 stomachs of S. graciosus and found no nematodes. He did, however, recover a cestode from the intestines of each of two lizards. Parasites from Sceloporus of neighboring areas have been ex- amined and provide useful comparisons with Great Basin and Upper Colorado Plateau populations. Waitz (1961) studied the parasites of Idaho reptiles, including S. graciosus and S. occidentalis. He re- covered Oochoristica scelopori from the former and Skrjabinoptera phrynosoma from the latter. Harwood (1932) studied the helminth parasites of reptiles neai? Houston, Texas, and found in S. undulatus the nematode Oswaldocruzia pipiens. His paper was the first major study on reptilian parasites from a specific geographical area in the United States. Tanner and Hopkin (1972) studied the ecology of 'Department of Zoology, Brigham Young University, Provo, Utah 84601. 2 GREAT BASIN NATURALIST Vol. 33, No. 1 S. occidentalis at the Nevada Test Site and reported 41 of 145 lizards with unclassified round worms in the stomach. Telford (1970) reported on 1485 lizards of southern California, the coastal islands, and Mexico, including S. magister, S. graciosus, and S. occidentalis. He lists many helminths, but he generally did not report the degree of infestation or the location of the parasite in the host. Stebbins and Robinson (1946) analyzed lizard populations in parts of California which included S. graciosus. They found three species of helminths in nine of the 15 animals examined. The para- sites were classified but were described as being only from the digestive tract. The present project was designed as a survey of helminth infesta- tions in Sceloporus lizards of the Great Basin and Upper Colorado Plateau of Utah. It also contains a bibliography of literature on reptile parasitology in the southwestern United States (Appendix). We gratefully acknowledge the help and advice of Drs. Ferron L. Andersen and Glen Moore, Departments of Zoology and Botany, respectively, Brigham Young University. Mr. Gary L. Burkholder supplied data from his study on 5. graciosus from central Utah. Methods and Procedures Host lizards were noosed, removed to the laboratory, and either anesthetized with ethyl ether or pithed. They were opened from the throat to the vent and microscopically examined for parasites in the coelom, mesenteries, and intact organs. The liver, lungs, esophagus, stomach, and small and large intestines were then removed and placed in a 75 percent physiological saline solution in preparation for dissection. Approximately 25 minutes after dissection, the organs were reexamined and placed in numbered vials. The dissected lizards were hardened in 1 0 percent formalin and added to the herpe- tology museum. The method outlined by Meyer and Penner (1962) for processing helminths was followed. Worms from preserved lizards were simply removed and placed in numbered vials. Para- sites were cleared in a solution of lactophenol for identification. Literature used in the classification of helminths include llyman (1951) and Yorke and Maplestone (1926) for the higher taxonomic categories. For the specific identification of helminths of the order Oxyuroidea, Gambino (1957a, 1957b), Lucker (1951), Read and Amrein (1953), Telford (1965), and Thapar (1925) were used. Spiruroid worms were identified by means of Babero and Kav (1967), Morgan (1942), Ortlepp (1922), and Telford (1965). Results Results of this study are presented in four sections, one for each species of Sceloporus studied. They are based on the dissection of 55 Sceloporus, 49 of which were anesthetized; the remaining six had previously been preserved. Sceloporus graciosus Baird and Girard. — Literature review re- veals S. graciosus to be parasitized by four species of helminths — March 1973 pearce, tanner: lizard parasites 3 one cestode and three nematodes (Table 4). The three nematodes represent two orders: Spiruroidea {Skrjabinoptera phrynosoma) and Oxyuroidea {Pharyngodon giganticus and Cyrtosomum penneri). Of 22 dissections, 11 were of males and 11 females. The only helminth recovered from the small intestine of S. graciosus was the tapeworm Oochoristica scelopori. Although this worm was as long as the entire small intestine (about 40 mm), it was contained within the duodenum of its host (about 10 mm) . All worms in the cecum were of Cyrtosomum and are considered pin worms of reptiles (Gambino, 1957b). Sceloporus graciosus is a new host for C. readi and C. heynemani, and, in addition, C. penneri is reported for the first time from Utah lizards. The worms were living free in the cecum of the large intestine and were in greatest numbers around and inside of fecal material on which they ap- peared to feed. Cyrtosomum species were not recovered from other regions of the intestine. Only 60.6 percent of the total helminth population in cecums were adults. The remaining 39.4 percent were larval Cyrtosomum not identifiable as to species. The S. graciosus specimens used in this study were from Provo Canyon and from the Little Sahara sand dunes southwest of Jericho, Utah. Of nine specimens from the Provo area, seven harbored helminths. It was in this group that O. scelopori was found. Six lizards harbored only Cyrtosomum. Four of the 13 lizards from the Jericho area were infested by Cyrtosomum. With the exception of O. scelopori, helminths of all species were recovered from lizards of these two areas. There were some differences in population composi- tion in the worms recovered from the two areas (Table 3) . Sceloporus undulatus elongatus Stejneger. — Three species of nematodes, one from the stomach and two from the large intestine, have been reported from S. undulatus as summarized in Table 1. Oswaldocruzia pipiens belongs to the order Spiruroidea, and the two species of Cyrtosomum are Oxyuroidea. The results for this species are based on the dissection of 11 adults, 8 males and 3 females. Nine of these were noosed; the other two had been preserved. Data for individual species of nematodes, with the host being considered as one unit, is recorded in Table 4; total helminth infestation per lizard is considered in Table 2. The stomach worms, Skrjabinoptera phrynosoma, caused deep welts in the stomach lining by imbedding their heads, which contain a pair of intemolateral teeth. This mode of attachment was so strong that the entire weight of the gastrointestinal tract could be supported by a single worm when held with forceps. Cecal worms varied in their life styles. Physaloptera retusa had a similar mode of attachment to that of Sk. phrynosoma. Pharyngo- don giganticus were free in the cecum and appeared to feed on fecal material. Although P. retusa and P. giganticus both occur in the cecum, the two did not occupy the same cecum. Skrjabinoptera phrynosoma occurred in one individual w^ith P. retusa and in another GREAT BASIN NATURALIST Vol. 33, No. 1 ^■6 3 in ^ o a ffi ffi 2 2 •5 *- c 2 tn ;0 (^ rO <£) (M xn ^~t^ -^ ^o t^ -^ t»~ T-H ^-\o-^^ ^^ ^^ ^•^ T^ b>.t^ ^ c 3 3 3 u U 0) U ^ QJ QJ Q^ 0) ^ 1^ u o t>0 So o -C . . . 3 3- O C <2 3 ;'-c: "to bo o. to ^ , C3 O "" "^ >< p 3 ^ 3 ^ -t^ ^ ^ ^ c c ~ C 3 ~ a o ^ c 3 acio agis. agis. cide cide mw acic agis undul ■yngodi gracio occide O <^ o L. ~ ~ o o (» fc fc o o o r- r (^ Co Oo J^ Co Co 00 to Co Scoco March 1973 pearce, tanner: lizard parasites 5 host with P. giganticus. These were the only cases of multispecies infestations in S. undulatus. All three lizards from the Zion National Park area harbored helminths. One of these contained both the stomach worm Sk. phrynosoma and the cecal worm P. giganticus. All lizards from Arches National Park carried Sk. phrynosoma, and one a female P. retusa in the cecum. None of the hzards from the St. George- Santa Clara area had helminths. The one from Price, Utah, har- bored S. phrynosoma (Table 3). Sceloporus occidentalis biseriatus Hallowell. — Literature reveals S. occidentalis to harbor six species of helminths (Table 4). Oocho- ristica scelopori was the only cestode recovered. The nematodes were Skrjabinoptera phrynosoma and Physaloptera retusa, order Spiruroidea; and Cyrtosomum penneri, Pharyngodon giganticus, and Thelandros iguanae, order Oxyuroidea. Seven anesthetized and four preserved specimens were dissected. Three species of helminths were recovered: O. scelopori, P. retusa, and P. giganticus. Nine larval nematodes {Skrjabinoptera or Physa- loptera) were recovered from a single cecum. Table 1 provides data for the individual species of nematodes, whereas the total helminths per infested host is given in Table 2. The stomach worm Physaloptera retusa was always found in the anterior third of the stomach. The mode of attachment was similar to that previously noted for Sk. phrynosoma in Sceloporus undulatus. There was, however, a major structural difference in that P. retusa had the cuticle reflected over the lips, and the teeth were dissimilar Table 2. Total helminths per lizard and infestations of host organs ex- pressed in maximum, minimum, and average for the four species of Sceloporus occurring in Utah. Maximum Minimum Average Per Infested Host Helminths Per Lizard S. graciosus S. magister S. occidentalis S. undulatus 250 .240 14 12 1 2 1 1 140 102 7 5.7 Helminths Per Organ Cecum S. graciosus S. occidentalis S. undulatus Duodenum 250 8 10 10 1 1 154 4 5 S. graciosus S. occidentalis S. undulatus Large Intestine S. magister Small Intestine 1 1 8 200 1 1 1 150 1 1 4.4 150 S. magister Stomach 100 10 55 S. magister S. occidentalis 30 13 2 1 10.4 5 GREAT BASIN NATURALIST Vol. 33, No. 1 Co N lniinth species recovered. The following is a discussion of each helminth species. Oochoristica scelopori. — This is a tapeworm of reptiles. Two specimens were recovered, one from the duodenum of S. graciosus March 1973 pearce, tanner: lizard parasites 9 and one from the duodenum of S. occidentalis. Both hosts were adult females. This cestode has been reported from 5. graciosus and S. occi- dentalis in southern California (Telford, 1970) and S. graciosus in Idaho (Waitz, 1961). A single cestode was found in one of the nine S. graciosus from Provo Canyon. Burkholder (personal communica- tion) autopsied more than 300 Utah specimens of S. graciosus and found one cestode each in two lizards also from Provo Canyon. The probable explanation of this apparent nidus, or nested distribution, lies with the unknown intermediate host. The intermediate host presumably is an arthropod having a narrower environmental tol- erance than S. graciosus. This possibility is strengthened by the absence of this parasite in 13 lizards from the Little Sahara sand dunes southwest of Jericho, Utah. Telford (1970) attempted to measure the effect of altitude on frequencies of parasitic infestations. He reported that the occurrence of cestodes in lizards was due to the sympatric occurrence of inter- mediate and definitive hosts and that the presence of intermediate hosts was dependent on the altitude. Telford's measurement of alti- tude was actually the sum total of climatic factors. The infested specimens of S. graciosus from Provo Canyon are from an area where water and vegetation are relatively abundant compared to dryer areas west of the Wasatch Front such as the Little Sahara sand dunes. Therefore, the unknown, intermediate host is probably an arthropod inhabiting cooler, moister habitats. The habitat of O. scelopori is seemingly the duodenum of the small intestine. Since this is the main organ of digestion (Villee and Dether, 1971), this parasite has an advantage in securing nutrients formed by the digestive processes of the host. The effects of O. scelopori on the hosts were apparently negli- gible. Neither S. graciosus nor S. occidentalis appeared to have structural damage. Although there may have been some competi- tion between hosts and parasites for the nutrients, the hosts were adults with fat bodies in the coelum, indicating an excess of nutri- ents. It is, of course, possible that multiple infestations of this worm would damage the host. In both host lizards there were no other helminths harbored. The possibility exists that O. scelopori has some mechanism which limits or interferes with other helminths. The literature is inconclusive on this point, and laboratory experimentation would be necessary to establish this hypothesis. A wide distribution is shown by O. scelopori. Telford (1970) reports this cestode in lizards of Iguanidae and Xantusiidae, for which families it is seemingly host specific. Skrjabinoptera phrynosoma. — This species is the best known stomach worm of reptiles in the southwestern United States. We have recovered 47 specimens from two species of Sceloporus. Seven of 11 specimens of S. undulatus and three of 11 specimens of S. magister harbored this worm. This appears to be the first report of this parasite in these species. 10 GREAT BASIN NATURALIST Vol. 33, No. 1 A possible nidus distribution exists for Sk. phrynosoma within the Great Basin and the Upper Colorado Plateau of Utah. Six of seven lizards, representing both host species, from Arches National Park were parasitized. In the St. George-Santa Clara area, however, only one of eight hosts harbored the worm. In infested stomachs of anesthetized lizards, Sk. phrynosoma was always located in the first third of the stomach with the head deeply buried in the stomach lining. The reason for this anterior attachment is not known. There may be less movement of the stomach anteriorly, or it may allow the parasite to extend without moving into the environment of the small intestine where the pH is quite different. Externally, lizards with Sk. phrynosoma could not be distin- guished from those without. Internally, however, the worms caused great welts in the stomach lining. There were no signs of tissue in- flammation or disintegration, but infestation was light. In some horned lizards, reported infestations are as high as 500 worms, with an average of 50 worms per lizard, and tissue damage is severe (Babero and Kay, 1967). However, the maximum number of worms in S. magister was 10, and in S. undulatus eight; the average for both species was about five worms per host. It seems that Sk. phrynosoma is not as abundant in Sceloporus as it is in Phrynosoma platyrhinos. The latter is a notorious ant eater, and since ants are intermediate hosts (Lee, 1957), we suspect that one or more species of ants may be involved in the parasitic cycle. Skrjabinoptera phrynosoma is limited to the family Iguanidae but is widely scattered through many of its genera (Telford, 1970). Physiological host specificity exists at the intermediate host level (Lee, 1957), but apparent preference for certain definitive hosts is ecologically determined by the dietary specializations of the host (Telford, 1970). Physaloptera retusa. — This species was recovered from S. occi- dentalism S. undulatus, and S. magister, the latter two constituting new host records. Physaloptera retusa is typically a stomach worm of S. occidentalis (Telford, 1970), but in both of the new hosts P. retusa was recovered from the cecum. In this study, specimens of 5. occidentalis were captured from the foothills west of Utah Lake and S. undulatus and S. magister were taken from Arches National Park. Sceloporus occidentalis was first reported to harbor P. retusa by Grundmann (1959), who examined seven specimens from Utah. Since then, Telford (1970) has reported this nematode to be the most common stomach worm i)arasitizing S. occidentalis in southern California. Perha[)s it is now justifiable to say that P. retusa is the most abundant stomach worm of 5". occidentalis in the southwestern states. Sceloporus magister and 5. undulatus harbored five and ten specimens respectively of the stomach worm Skrjabinoptera phryno- soma. It is interesting to note that P. retusa occurs in the cecum of these species and in the stomach of iS. occidentalis. March 1973 pearce, tanner: lizard parasites 11 The effects of P. retusa on its Sceloporus hosts appeared negli- gible. Attachment of worms to host stomachs was similar to that described for Sk. phrynosoma. Grundmann (1959) thought P. retusa was host specific for S. occidentalis, but Telford (1970) has shown it to be in genera as diverse as Crotaphytus, Uta, Callisaurus, Eu- meces, and Gerrhonotus. He suggested that host specificity of P. retusa is dependent on ecological factors similar to those in Sk. phrynosoma, the main factor being the diet of host lizards, which explains the distribution of this parasitic worm. Telford attributed the lack of P. retusa in S. magister from southern California to the absence in their diet of an unknown intermediate host. The presence of P. retusa in both 5. magister and S. undulatus from Arches Na- tional Park indicates that the intermediate host is a common item in these diets. Thubunaea iguanae. — This spiruroid nematode was found in the stomachs of two specimens of S. magister. One, from Arches National Park, had 30 worms; the other, from the Price area, had six. This is a sizable range extension for this nematode. Telford (1970) originally reported its occurrence in S. magister. The effects of T. iguanae on its host appeared negligible. The head of the worm was imbedded, but the attachment did not appear as strong as that reported above for Sk. phrynosoma and P. retusa. Both hosts were adults, apparently in good health. The effects of T. iguanae on other species of helminths could not be determined; however, there is a possibility that Sk. phrynosoma may have af- fected the distribution of T. iguanae. These helminths were not recovered from the same host. As noted above, P. retusa was not recovered from stomachs containing Sk. phrynosoma. These two facts support a theory that Sk. phrynosoma affects the presence and/or distribution of other nematodes within the host. This nematode is harbored by several species from each of the following families of reptiles: Iguanidae, Xantusiidae, Teiidae, and Gekkonidae (Telford, 1970). Pharyngodon giganticus. — This nematode belongs to the order Oxyuroidea, and was found in the cecum. It was recovered from two specimens of S. undulatus from Zion National Park, as well as from four specimens of S. occidentalis- — three from west of Utah Lake and one from St. George, Utah. Sceloporus undulatus repre- sents a new host record for P. giganticus. The effects of P. giganticus upon hosts were apparently negli- gible. This helminth lives in the fecal material of the host's cecum. The host-worm relationship appears to be commensal. There is the possibility that under heavy infestations this worm may be injurious to the host; however, such was not noted by us. The effect of P. giganticus on other oxyuroid helminths is a possibility, since there were no other oxyuroid worms sharing the cecum with it; however, there were no other oxyuroid species found in specimens of S. undulatus or S. occidentalis. 12 GREAT BASIN NATURALIST Vol. 33, No. 1 Oxyuroid nematodes show a greater degree of host specificity than do the spiruroids previously mentioned. Telford (1970) reports P. giganticus to be restricted to Sceloporus, with the exception of two widely separated island populations of Uta, which he suggests may be a relict distribution. Cyrtosomum penneri, C. heynemani, and C. readi. — These three oxyuroids are considered together because of the many similarities in host, habitat, and life style. The Cyrtosomum species were usually found in association with either one or both of the other Cyrtosomum species. All three species were recovered from specimens of Scelopo- rus graciosus taken from Provo Canyon and the Little Sahara sand dunes. Cyrtosomum penneri and C. readi were recovered from one specimen of S. magister taken from Arches National Park and also from one specimen from the Price area. This study reports 5. graciosus as a new host for C. readi. Gambino (1957a, 1957b) describes and discusses the morphology of these nematodes. All three Crytosomum nematodes infested the ceca of their hosts. In only two specimens of S. magister did these worms occur outside of the cecum; since both lizards were preserved specimens, this occurrence could be explained by a migration at the time of preser- vation. Dissections and observations of infested hosts revealed that Cyrtosomum worms probably live exclusively in the cecum. Ecologi- cal barriers such as peristaltic movement, scarcity of food, and re- striction in space may be limiting factors. Physiological barriers in the large intestine may limit worm distribution past the cecum, where the colon removes most of the water and concentrates meta- bolic wastes. These would cause osmotic and pH gradients that might impair the survival of parasites. The effects on the host by Cyrtosomum nematodes appeared negligible. At times, 300 worms occurred in a single host. Such heavy infestations of Cyrtosomum worms often caused the walls of the cecum to move as the worms pushed against it. This may have caused irritation; however, there was no tissue damage noted. Worms were seemingly feeding on the fecal matter, and thus were not competing with the host for food but living in a commensal relationship. The presence or absence of other nematodes showed no effect on the presence or abundance of Cyrtosomum worms. Cyrtosomum penneri was the most commonly occurring Cyrtosomum. It shared all the infested lizards harboring either C. readi or C. heynemani and was found in two 5. magister lizards by itself. Cyrtosomum heynemani was the only species which did not occur in S. magister. This may not be significant, because some experts are not certain that C. readi is a distinct species from C. heynemani (Babero and Kay, 1967). Cyrtosomum are host specific to the family Iguanidae, infesting many genera including Dipsosaurus, Crotaphytus, Sauromalus, Phrynosoma, Uta, Callisaurus, and Sceloporus (Telford, 1970). March 1973 pearce, tanner: lizard parasites 13 Conclusions and Summary Dissections of 55 lizards representing four species of Sceloporus resulted in the recovery of eight species of helminths. These in- clude: a cestode, Oochoristica scelopori; three species of spiruroid nematodes, Skrjabinoptera phrynosoma, Physaloptera retiisa, and Thubunaea iguanae; and four species of oxyuroid nematodes, Pharyngodon giganticus, Cyrtosomum penneri, C. heynemani, and C. readi. Eight new host records were reported, and the geographic distribution was extended for most of the parasitic species. Host specificity was not restrictive enough to indicate phylo- genetic relations between host species. The occurrence of helminths in Sceloporus lizards apparently was correlated with the distribution of intermediate hosts and the diets of definitive hosts. The effects of helminths on their Sceloporus hosts were apparently slight; how- ever, infestations of cestodes or spiruroid nematodes was light. The possibility exists that lizards with high infestations have a lower survival rate. Oxyuroid nematodes did occur in great numbers, but in all cases the hosts appeared healthy. The effects of helminths on other helminths in the same host were indicated. Stomachs were parasitized by a single species of nematode even when other helminths, usually inhabitants of the stomach, were recovered from the same lizard. Skrjabinoptera phrynosoma was apparently dominant to both Physaloptera retusa and Thubunaea iguanae. There were some differences in number and types of parasites found in the same host from different collecting sites. Specimens from the St. George-Santa Clara area were almost free of helminths, whereas those from Arches National Park were all infested. This nested distribution evidently resulted from different diets and the presence or absence of intermediate hosts in the locality. Appendix The literature available on reptilian parasitology from the American Southwest is presented as an addition to this study. We recognize the bibliographic contributions of Dr. Sophie Jakowska. Alexander, C. G., and E. P. Alexander. 1957. Oochoristica crotalicola, a new anoplocephalid cestode from California rattlesnakes. J. Parasitol. 43:365-366. Allred, D. M. 1958. A new species of pit mite (Acarina: Ophioptidae) infest- ing snakes. Herpetologica 14:107-112. Allred, D. M., and D E. Beck. 1962. Ecological distribution of mites on lizards at the Nevada Atomic Test Site. Herpetologica 18:47-51. Amrein, Y. U. 1951. The intestinal entozoa of the night lizards of California and their mode of transmission. Unpublished Ph.D. dissertation, Univ. Cali- fornia, Los Angeles, California. . 1952. A new species of Isospora, /. xantusiae, from Southern California lizards. J. Parasitol. 38:147-150. 1953. The mode of transmission of non-cyst-forming flagellates in the ovoviviparous Yucca night lizard Xantusia vigilis. Ecology 34:243-245. Andersen, F. L., and N. D. Levine. 1961. Tritrichomonas batrachorum in a hog-nosed snake {Heterodon platyrhinos) . J. Parasitol. 47:877-878. 14 GREAT BASIN NATURALIST Vol. 33, No. 1 Ash, L. R., and P. C. Beaver. 1962. A restudy of Ophidascaris labiatopapillosa occurring in the stomach of northern American snakes. J. Parasitol. 48(2): 41. Abstr. Babero, B. a., and D. Matthias. 1967. Thubunaea cnemidophorus n. sp. and other helminths from lizards, Cnemidophorus tigris, in Nevada and Arizona. Trans. Amer. Microsc. Soc. 86(2) : 1 73-177. Ball, G. H. 1944. Sarcosporidia in Southern California lizards. Trans. Amer. Microsc. Soc. 58(2): 144-148. Banta, B. H. 1962. Beetles attacking lizards. Brit. Jour. Herpetol. 3:39. Barrow^, J. H., Jr., and J. J. Stockton. 1960. The influence of temperature on the host-parasite relationship of several species of snakes infested with Entamoeba invadens. J. Protozool. 7:377-383. Baylis, H. a. 1926. On a new species of the nematode genus Thubunaea. Ann. Mag. Nat. Hist. 18(9): 361 -364. BoNoiRis, J. S., AND G. H. Ball. 1955. Shellackea occidentalis n. sp., a blood- inhabiting coccidian found in lizards in Southern California. J. Protozool. 2:31-34. BovEE, E. C. 1961. Entozoic amoebas from feral reptiles. Amer. Zool. 1:439. . 1966. Eimeria urosuris sp. n., a coccidium from the lizard Urosaurus graciosus Hallowell, in California. J. Protozool. 13(4) :607-610. BovEE, E. C, AND S. R. Telford, Jr. 1965a. Eimeria sceloporus and Eimeria molochis spp. n. from lizards. J. Parasitol. 51:85-94. . 1965b. Eimeria noctisauris sp. n., a coccidian from the lizard, Klau- berina riuersiana. J. Parasitol. 51:325-330. Brennan, J. M., AND E. K. JoNES. 1954. A report on the chiggers (Acarina: Trombiculidae) of the Frances Simes Hastings Natural History Reservation, Monterey County, California. Wasmann, J. Biol. 12:155-194. Caballero, E. 1938. Nematodes of the reptiles of Mexico. H Ann. Trop. Med. Parasitol. 32:225-229. Cable, R. M., and W. B. Hopp. 1954. Acanthocephalan parasites of the genus N eoechenorphynchus in North American turtles, with the description of two new species. J. Parasitol. 40:674-680. Camin, J. H. 1953. Observations on the life history and sensory behavior of the snake mite, Ophionyssus nastricis (Gervais) (Acarina: Macronyssidae) . Chicago Acad. Sci. Spec. Publ. 10:1-75. Chitwood, B., and E. Wehr. 1934. The value of cephalic structures as char- acters in nematode classification, with special reference to the superfamily Siruroidea. Z. Parasitenk. 7:273-335. Cobb, N. 1928. Amphids of the nema Physaloptera phrynosoma. J. Parasitol. 15:70. Davidson, J. A. 1958. A now species of lizard mite and a generic key to the family Pterygosomidae (Acarina: Anystoidea). Proc. Ent. Soc. Washington 60:75-79. Diamond, L. S. 1960. The axenic culture of two reptilian parasites, Enta- moeba terrapinae, Sanders and Cleveland, 1930, and Entamoeba invadens, Rodhain, 1934. J. Parasitol. 46:484. Doge, H. R. 1955. Sarcophagid flies parasitic on reptiles. Proc. Ent. Soc. Washington 54(4) : 183-187. Edgerly, R. H. 1952a. Two new species of Nematoda, Strongyluris riversi- densis and Pharyngodon mearnsi, from lizard, Streptosaurus mrarnsi. Trans. Amer. Microsc. Soc. 71:288-292. 1952b. A new oxyuroid nematode. Alaeuris yumanae n. sp., from the lizard, Dipososarus dorsalis. Trans. Amer. Microsc. Soc. 71:341-343. Eglis, a. 1959. Tick infestations of western spur-thighed tortoises. Herpe- tologica 15:125. Evans, F. R. 1959. Protozoa from the intestine of some desert lizards. Proc. Utah Acad. Sci., Arts, Letters 36:175-176. Abstr. EwiNG, H. E. 1926. The common box turtle, a natural host for chiggers. Proc. Biol. Soc. Washington 39:19-20. Fantham, H. B., and a. Porter. 1954. The endoparasites of some North American snakes and their effect on the Ophidia. Proc. Zool. Soc. London 123:867-898. March 1973 pearce, tanner: lizard parasites 15 Fowler, J. A. 1946. Reptiles and amphibians as a source of blood parasites in protozoology and parasitology. Turtox News 24(4): 73-75. Gambino, J. J. 1957. Relationships of North American Iguanidae, as suggested by analysis of their pinworm parasites. Anat. Record. 128:553. Geiman, Q. M., and H. L. Radcliffe. 1936. Morphology and life cycle of an amoeba producing amoebiasis in reptiles. J. Parasitol. 28:208-228. George, J. E. 1960. Notes on the parasitic mites of some west Texas reptiles. Southwestern Nat. 5:105-106. Goodman, J. D. 1951. Some aspects of the role of parasitology in herpetology. Herpetologica 7:65-67. Goodwin, M. H. 1954. Observations on the biology of the lizard mite Gecko- biella texana (Banks) 1904 (Acarina: Pterygosomidae) . J. Parasitol. 40: 54-59. Gould, D. J. 1956. The larval trombiculid mites of California (Acarina: Trom- biculidae). Univ. California Pub. Ent. 11:1-115. Hannum, C. a. 1943. Nematode parasites of Arizona vertebrates. Theses Ser. Univ. Washington 7:229-231. Hegner, R., and R. Hewitt. 1940. A new genus and new species of amoebae from Mexican lizards. J. Parasitol. 26:319-321. HiLMAN, J. L., AND R. W. Strandtmann. 1960. The incidence of Hepatozoon serpentium in some Texas snakes. Southwestern Nat. 5:226-228. Honigberg, B. M. 1950. Intestinal flagellates of amphibians and reptiles. Unpublished Ph.D. dissertation, Univ. California, Berkeley, California. . 1953. Structure, taxonomic status, and host list of Tritrichomonas hatrachorum (Perty). J. Parasitol. 39:191-208. . 1955. Structure and morphogenesis of two new species of Hexamasdx from lizards. J. Parasitol. 41:1-17. 1963. Evolutionary and systematic relationships in the flagellate order Trichomonadida Kirby. J. Protozool. 10:20-63. Hopp, W. B. 1954. Studies on the morphology and life cycle of Neochinor- hynchus emydis (Leidy), an Acanthocephalan parasite of the map turtle, Graptemys geographica (Le Sueur). J. Parasitol. 40:284-299. Hughes, R. C, J. W. Higginbotham, and J. W. Clay. 1941. The trematodes of reptiles. Part H. Host catalogue. Proc. Oklahoma Acad. Sci. 21:35-43. Hughes, R. C, J. R. Baker, and C. B. Dawson. 1942a. The trematodes of reptiles. Part I. Systematic section. Amer. Midi. Nat. 27:109-134. . 1942b. The tapeworms of reptiles. Part III. Proc. Oklahoma Acad. Sci. 22:81-89. Hyland, K. E., Jr. 1950. The copperhead snake as a host for the chigger mite Trombicula (Eutrombicula) alfreddugesi. J. Parasitol. 36:494. Irwin-Smith, V. 1921. Note on nematodes of the genus Physaloptera with special reference to those parasitic in reptiles. Part I. Proc. Linn. Soc. N. S. W. 46(4). Sydney. . 1922a. Note on nematodes of the genus Physaloptera, with special reference to those parasitic in reptiles. Part II. A review of the Physalop- tera of lizards. Proc. Linn. Soc. N. S. W. 47(2). Sydney. 1922b. A review of the Physaloptera of lizards. Proc. Linn. Soc. N. S. W. 47:232-244. Janakidevi, K. 1961. Hexamastix dobelli n. sp., a new trichomonad parasitic in the starred tortoise. J. Protozool. 8(3):294-296. King, W., and J. V. Griffo, Jr. 1958. A box turtle fatality apparently caused by Sarcophaga cistudinis larvae. Florida Ent. 41:44. Lawrence, R. F. 1953. Two new scale-mite parasites of lizards. Proc. U. S. Natl. Mus. 103:9-18. Lee, J. J. 1960. Hypotrichomonas acosta (Moskowitz) gen. nov. from reptiles. I. Structure and division. J. Protozool. 7:393-401. Lee, S. H. 1955. The mode of egg dispersed in Physaloptera phrynosoma Ortlepp (Nematode: Spiruroidea), a gastric nematode of Texas horned toads, Phrynosoma cornutum. J. Parasitol. A'l:70-7'\-. Lehmann, D. L. 1960. Blood parasites of West Coast amphibians and reptiles. Amer. Philos. Soc. Yearbook 1959:244-246. 16 GREAT BASIN NATURALIST Vol. 33, No. 1 Lent, H., and J. F. Teixeira de Freitas. 1948. Una colegao de nematodeos, parasites de vertebrados, do Museu de Historia Natural de Montevideo. Mem. Inst. Oswaldo Cruz. 46:1-71. Loewen, S. L. 1940. On some reptilian cestodes of the genus Oochoristica (Anoplocephalidae). Trans. Amer. Microsc. Soc. 59:511-518. LooMis, R. B. 1964. A new species of chigger (Acarina: Trombiculidae) from lizards of western North America. Great Basin Nat. 24:13-17. LooMis, R. B., AND D. A. Crossley. 1963. New species and new records of chiggers (Acarina: Trombiculidae) from Texas. Acarologia 5:371-383. McMuLLEN, D. B. 1940. Cutaneous myiasis in a box turtle. Proc. Oklahoma Acad. Sci. 20:23-25. Meerovitch, E. 1960. Thermal barrier to the infectivity of Entamoeba inva- dens in snakes. Nature 185:631. . 1961. Infectivity and pathogenicity of polyxenic and monoxenic Entamoeba invadens to snakes kept at normal and high temperatures, and the natural history of reptile amoebiasis. J. Parasitol. 47:791-794. Mockerras, M. J. 1953. Lizard Filaria: transmission by mosquito of Oswaldo- filaria chlamydosauri (Breinl) (Nematode: Filarioidae). Queensland Inst. Med. Res., Brisbane, Australia. Biol. Abstr. 27:3154 (33157). Morgan, B. B. 1940. The Physalopterinae (Nematoda) of North America. J. Parasitol. 26:40. . 1941a. A summary of the Physalopterinae (Nematoda) of North America. Proc. Helm. Soc. Washington 8:28-30. . 1941b. Additional notes on North American Physalopterinae (Nema- toda). Proc. Helm. Soc. Washington 8:63-64. . 1942. The Physalopterinae (Nematoda) of North American verte- brates. Summary of doctoral dissertation, Univ. Wisconsin 6:88-91. 1943. The Physaloptera (Nematoda) of reptiles. Natural. Canad. 70:179-185. Moskowitz, N. 1951. Observations on some intestinal flagellates from rep- tilian hosts (Squamata). J. Morphol. 89:257-321. Ortlepp, R. 1922. The nematode genus Physaloptera Rud. Proc. Zool. Soc. London 999-1107. . 1937. Some undescribed species of the nematode genus Physaloptera, together with a key to the sufficiently known forms. Onder. J. Vet. Sci. 9:71-84. Pelaez, D., and R. p. Reyes. 1958. Piratuba Prolifica sp., parasita de un Sceloporus mexicano (Nematoda: Filar.). An. Esc. Nat. Cienc. Biol. 9:49-50. Peters, J. A. 1948. The box turtle as a host for dipterous parasites. Amer. Midi. Nat. 40:472-474. Powder, A., and R. B. Loomis. 1962. A new species and new records of chig- gers (Acarina: Trombicuhdae) from reptiles of Southern California. J. Parasitol. 48:204-208. Read, C. P., and Y. U. Amrein. 1952. Some new ox^uroid nematodes from southern California. J. Parasitol. 38:379-384. Reichenback-Klinke, H., and E. Elkan. 1965. The principal diseases of lower vertebrates. Academic Press, London and New York. RoTHMAN, A. 1954. A new species of Strongyluris (Nematoda) from a South- ern California lizard. J. Parasitol. 40:673 674. RouDABUSH, R. W. 1937. Some coccidia of reptiles found in North America. J. Parasitol. 23:351-359. Saxe, L. H., AND E. M. Schmidt. 1953. Trilmitus parvus Grasse (Protozoa: Mastigophora) from a garter snake, Thamnophis radix. Proc. Iowa Acad. Sci. 60:754-758. ScHiFFER, P. 1929. Mites on 5ce/oporw5. Copeia 1171:45-46. Seurat, L. G. 1914a. Sur un nouvel Oxyure des reptiles. C. R. Sernc. Soc. Biol. 77:96. Paris. . 1914b. Sur deux Physalopteres tetrahysteriens des reptiles. C. R. Seme. Soc. Biol. 77:433-436. Paris. Skinner, R. V. 1957. Acarine parasite on sand lizard {Lacerta agilis) . Brit. Jour. Herpet. 2:93. Steelman, G. M. 1939. A new cestode from Texas horned lizard. Tr. Am. Micr. Soc. 58:452-455. March 1973 pearce, tanner: lizard parasites 17 Stiles, C. W., and A. Hassall. 1894. A preliminary catalogue of the parasites contained in the collection of the U.S. Bur. of Animal Industry, U.S. Army Med. Mus., Biol. Dept. of the Univ. Penn. (Coll. Leidy) and in Coll. Stiles and Coll. Hassall. Vet. Mag. 1:245-253, 331-354. Telford, S. R., Jr. 1965a. New species of the oxyuroid nematode genus Pseu- doalaeris from California lizards. Japan. J. Exper. Med. 35:99-109. . 1965b. A new nematotaeniid cestode from California lizards. Japan. J. Exper. Med. 35:301-303. . 1965c. Some Thelandros (Nematoda: Oxyuroidea) from California lizards. Japan. J. Exper. Med. 35:463-472. 1966. Morphological observations on haemosporidian parasites of some California and Mexican lizards. Japan. J. Exper. Med. 36:237-250. Thomas, L. A., and C. M. Eklund. 1960. Overwintering of western equine encephalotyelitis virus in experimentally infested garter snakes, and trans- mission to mosquitoes. Proc. Soc. Exp. Biol. Med. 105:52-55. Thompson, P. E., and C. G. Huff. 1942. Exoerythrocytic schizogony in a new species of saurian Plasmodium. J. Parasitol. 28(suppl.) :15. . 1944a. A saurian malarial parasite, Plasmodium mexicanum, n. sp., with both elongatum- and gallinaceum-types of exoerythrocytic stages. J. Inf. Dis. 74:48-67. . 1944b. Saurian malarial parasites of United States and Mexico. J. Inf. Dis. 74:68-79. Vincent, I. 1948. Studies on the endoparasites of the Texas horned lizards Phrynosoma cornutum. (Harlan). Proc. Texas Acad. Sci. 30:250-252. Walton, A. 1927. A revision of the nematodes of the Leidy collections. Proc. Nat. Acad. Sci. Philadelphia 79:49-163. Walton, A. C. 1941. Distribution of the genus Thelandros (Nematoda: Oxyuroidea). Proc. Helminth. Soc. Washington 8:15-18. WiECzoROWSKi, Elsie. 1939. Parasite lesions in turtles. J. Parasitol. 25:395-399. WiTENBERG, G. 1934. Studies on the cestode genus Mesocestoides. Arch. Zool. Ital. 20:467-509. Wood, W. F. 1935. Some observations on the intestinal protozoa of California lizards. J. Parasitol. 21 : 165-173. Literature Cited 1. Babero, B. B., and F. R. Kay. 1967. Parasites of horned toads {Phryno- soma spp.), with records from Nevada. J. Parasitol. 53:168-175. 2. Gambino, J. J. 1957a. Cyrtosomum readi n. sp. and Cyrtosomum heyne- mani n. sp. (Oxyuroidea: Atractidae) two new pinworms of iguanids. J. Parasitol. 44:439-445. 3. . 1957b. Cyrtosomum penneri n. sp. (Oxyuroidea: Atractidae). J. Parasitol. 43:76-80. 4. Grundmann, a. W. 1957. A preliminary study of the intestinal parasites of some Utah lizards. Proc. Utah Acad. Sci. 34:147-148. 5. . 1959. Parasites recovered from six species of Utah lizards. J. Parasitol. 45:394. 6. Harwood, p. D. 1932. The helminths parasitic in the amphibia and reptilia of Houston, Texas, and vicinity. Proc. U.S. Nat. Mus. 81:1-67. 7. Hyman, L. H. 1951. The invertebrates: Acanthocephala, Aschelminthes, and Entoprocta. Vol. Ill, 1st ed. McGraw-Hill Book Co., New York. 8. Lee, S. H. 1957. The life cycle of Skrjabinoptera phrynosoma (Ortlepp) Schulz, 1927 (Nematoda: Spiruroidea), a gastric nematode of Texas horned toads, Phrynosoma cornutum. J. Parasitol. 43:66-75. 9. Lucker, J. T. 1951. Some new Thelandros (Nematoda: Oxyuridae) from the island night lizard, Xantusia riversiana reticulata Smith, from San Clemente Island, California. J. Parasitol. 37(suppl.): 14-15. 10. Meyer, M. C, and L. R. Penner. 1962. Laboratory essentials of parasi- tology. Wm. C. Brown Co., Dubuque, Iowa. 11. Morgan, B. B. 1942. The nematode genus Skrjabinoptera Schulz, 1927. Lloydia 5:314-319. 18 GREAT BASIN NATURALIST Vol. 33, No. 1 12. Ortlepp, R. 1922. The nematode genus Physaloptera Rud. Proc. Zool. Soc. London 72:999-1107. 13. Read, C. P., and Y. U. Amrein. 1953. North American nematodes of the genus Pharyngodon Diesing (Oxyuridae). J. Parasitol. 39:365-370. 14. Stebbins, R. C, and H. B. Robinson. 1946. Further analysis of a popula- tion of the lizard Sceloporus graciosus gracilis. Univ. Calif. (Berkeley) Pubis. Zool. 48(3): 149-168. 15. Tanner, W. W., and J. M. Hopkin. 1972. Ecology and life history of Sceloporus occidentalis longipes Baird and Girard on Rainier Mesa, Nevada Test Site, Nye County, Nevada. Brigham Young Univ. Sci. Bull., Biol. Series 15(4): 1-39. 16. Telford, S. R., Jr. 1965. A new species of Thubunaea (Nematode: Spiruroidea) from California lizards. Japan. J. Exper. Med. 35:111-114. 17. . 1970. A comparative study of endoparasitism among some South- ern California lizard populations. Amer. Midi. Nat. 83:516-533. 18. Thapar, G. S. 1925. Studies on the Oxyuroid parasites of reptiles. J. Helminthol. 3:83-150. 19. ViLLEE, C. A., and v. G. Dether. 1971. Biological principles and pro- cesses. W. B. Saunders Co., Philadelphia, London, and Toronto. 20. VoGE, M., and W. Fox. 1950. A new anoplocephalid cestode, Oochoristica scelopori n. sp. from the Pacific Fence Lizard, Sceloporus occidentalis occidentalis. Trans. Amer. Microsc. Soc. 69:236-242. 21. Waitz, J. A. 1961. Parasites of Idaho reptiles. J. Parasitol. 47:51. 22. Woodbury, L. A. 1934. Notes on some parasites of three Utah reptiles. Copeia 1:51-52. 23. YoRKE, W., AND P. A. Maplestone. 1926. The nematode parasites of vertebrates. P. Blakiston's Son and Co., Philadelphia. MISCELLANEOUS CHROMOSOME COUNTS OF WESTERN AMERICAN PLANTS— IP James L. Reveal- and Eloise L. Styer* Abstract. — Chromosome numbers are reported for 19 species of western American plants. Reports for previously uncounted species include Nitrophila mohavensis Munz and Roos, n=18; Eriogonum contiguum (Reveal) Reveal, n = 16; Mentzelia leucophylla Brandeg., n=18, and M. torreyi A. Gray, n = 12; Prunus virginiana L. var. melanocarpa (A. Nels.) Sarg., n = 8; Astragalus beatleyae Barneby, n = ll; Angelica kingii (S. Wats.) Coult. and Rose, n = 22; Gilia nyensis Reveal, n==9; and Arnica parryi A. Gray var. sonnei (Greene) Cronq. in Ferris n==19. New ploidy levels are reported in Oxystylis lutea Torr. and Frem., n=10; Lupinus aridus Dougl. ex Lindl., n = 12; and Castilleja mar- tinii Abrams var. clokeyi (Pennell) N. H. Holmgren, n = 12. Seven additional counts are given which confirm those published by others. These are Opuntia chlorotica Engelm. and Bigel., n = ll; Eriogonum inf latum Torr. and Frem. var. inf latum, n = 16, E. trichopes Torr., n = 16, and E. bifurcatum Reveal, n = 20; Stanleya pinnata (Pursh) Britt. var. inyoensis (Munz and Roos) Reveal, n = 28; Lupinus argenteus Pursh, « = 24; and Castilleja viscidula A. Gray, n = 12. In this series of papers, of which this is the second, chromosome counts of randomly gathered western American plants will be re- ported as part of the Intermountain Flora Project (headed by Arthur Cronquist of the New York Botanical Garden and aided by Arthur H. Holmgren, Noel H. Holmgren, and James L. Reveal) and the proposed Southwest Flora Project (headed by Noel H. Holmgren and James L. Reveal). These counts will be presented from time to time by Reveal and various others of his colleagues or students and will be based on materials gathered by him or others. The present paper reports on some of the miscellaneous plants gathered in south- ern Nevada in 1970 and 1971. Flower buds were collected in developmental series from plants growing in their native habitats. The buds were fixed in ethanol and glacial acetic acid (3:1) and refrigerated; the buds were not transferred to 70 percent ethanol after 24 hours, as is the usual tech- nique. Anthers were squashed in acetocarmine and illustrations drawn using a camera lucida mounted on a Wild M20 research microscope. Most of the counts reported here were made by Reveal. Voucher specimens are deposited in the herbarium at the Nevada Test Site, with duplicates deposited in several major herbaria. Chromosome numbers of 19 species of western American plants are reported here, nine for the first time, with three counts of new ploidy levels. One count is given for a species already reported in the literature but not figured, and six counts confirm those made previously by others. Opuntia chlorotica Engelm. and Bigel. 72=11. Figs. 1, 2. This count confirms those made by others (Stockwell, 1935; Pinkava and 'The first paper in this series was published in the Southwestern Naturalist. ^Department of Botany, University of Maryland, College Park 20742; National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560. ^Department of Botany, University of Maryland, College Park 20742. 19 20 GREAT BASIN NATURALIST Vol. 33, No. 1 McLeod, 1971). The voucher is Beatley and Reveal 11231, west side of Spring Mountains [Charleston Mountains], Clark Co., Nevada, 17 June 1970. Nitrophila mohavensis Munz and Roos. « = 18. Figs. 3, 4. This rare species is known only from a small site in extreme eastern Inyo Co., California, where it is a locally dominant species. It would appear that A^. mohavensis is a tetraploid, since Covas and Schnack (1947) have reported a count of 72 = 9 for the South American species A^. australis Chod. and Wile, although a second North American species, A^. occidentalis (Nutt.) Moq., should be counted to confirm this. Nevertheless, it seems possible to state that the base number for Nitrophila is t = 9. The voucher collection is Reveal 2291, Amar- gosa Desert, southern Ash Meadows, along the road between Ash Meadows and Death Valley Junction on the saline flats of Carson Slough, Inyo Co., California, 5 May 1971. Eriogonum inf latum Torr. and Frem. var. inf latum. «=16. Fig. 5. This count confirms those made by Stone and Raven (1958) and Reveal (1965). The voucher is Reveal 2290, Stewart Valley, 2 miles north of Nevada Highway 52 on road to Ash Meadows, Nye Co., Nevada, 5 May 1971. Eriogonum contiguum (Reveal) Reveal. n=\6. Figs. 6, 7. When this plant was counted and found to equal that of E. inflatum and E. trichopes, and when fieldwork showed no hybridization be- tween the three, it was decided to elevate E. inflatum var. contiguum to the species rank (Reveal, 1972). The voucher of this new count is Reveal 2288, Stewart Valley, 2 miles north of Nevada Highway 52 on road to Ash Meadows, Nye Co., Nevada, 5 May 1971. Eriogonum trichopes Torr. n=16. Figs. 8, 9. This count con- firms that made previously (Reveal, 1965). The voucher is Reveal 2289, Stewart Valley, 2 miles north of Nevada Highway 52 on road to Ash Meadows, Nye Co., Nevada, 5 May 1971. Figs. 1, 2: Opuntia chlorotica, ti=\\ — metaphase I. Figs. 3, 4: Nitrophilia mohavensis, n=\S — diakinesis. Fig. 5: Eriogonum inflatum var. inflatum. n = 16 — telophase II. Figs. 6, 7: Eriogonum contiguum. n = \6 — anaphase I, Fig. 6; metaphase I, Fig. 7. Figs. 8, 9: Eriogonum trichopes, « = 16 — diakinesis. Fig. 8; anaphase I, Fig. 9. Fig. 10: Eriogonum bifurcatum. n^20 — diakinesis. Figs. 11, 12: Mentzelia leucophylla, n=\S — diakinesis. Figs. 13, 14: Mentzelia torreyi, „ = 12— telophase II, Fig. 13; anaphase II, Fig. 14. Figs. 15, 16, 17, 18: Oxy- stylis lutea, n=10 — metaphase I. Figs. 15, 17, 18; diakinesis, Fig. 16. Figs. 19, 20: Stanleya pinnata var. inyoensis, n^28 — diakinesis. Fig. 19; anaphase I, Fig. 20. Figs. 21, 22, 23: Prunus virginiana var. melanocarpa, n = 8 — metaphase I, Figs. 21, 22; anaphase I, Fig. 23. Figs. 24, 25: Astragalus beatleyae. n = ll — anaphase I. Figs. 26, 27: Lupinus argenteus, n = 24 — metaphase I, Fig. 26; anaphase I, Fig. 27. Figs. 28, 29: Lupinus aridus. n=\2 — metaphase I, Fig. 28; metaphase II. Fig. 29. Figs. 30, 31: Angelica kingii. n = 22 — anaphase I, Fig. 30; metaphase I, Fig. 31. Fig. 32: Gilia nyensis, n = 9 — anaphase I. Figs. 33, 34: Casiilleja viscidula, n=\2 — anaphase I, Fig. 33; telophase II, Fig. 34. Figs. 35, 36: Castilleja martinii var. clokeyi, n = l2 — anaphase I. Figs. 37, 38: Arnica parryi var. sonnei, n=19 — anaphase I. March 1973 reveal, styer: plant chromosomes 21 22 GREAT BASIN NATURALIST Vol. 33, No. 1 Eriogonum bifurcatum Reveal. « = 20. Fig. 10. When this species was proposed (Reveal, 1971), the chromosome number was noted but a figure not provided. The voucher of this count is Reveal 2283, Pahrump Valley, 1 mile east of the California-Nevada state line just south of Nevada Highway 52, Nye Co., Nevada, 1 3 June 1970. M entzelia leucophylla Brandeg. n=18. Figs. 11, 12. As current- ly understood, this is a rather rare species of eastern California and adjacent southern Nevada. The count indicates the species to be a polyploid (most likely a tetraploid), but one would like more counts of the species from throughout its range to confirm this point. The voucher is from the type location in Ash Meadows, Nye Co., Nevada, based on Reveal 2247, 11 June 1970. Mentzelia torreyi A. Gray. n—\2. Figs. 13, 14. This count is most unusual, since this number is unknown (at least to us) for the genus. A more reasonable number would be n=ll, but our report was confirmed by a number of smears. Bud material was obtained by Reveal and Beatley on 14 June 1970 from the same place Reveal made a collection earlier. The voucher is Reveal 1980, Kawich Val- ley, Nye Co., Nevada, 31 August 1968. Oxystylis lutea Torr. and Frem. n=10. Figs. 15, 16, 17, 18. Raven, Kyhos, and Hill (1965) recently reported the number of 2a2 = 20 pairs for this species, but our counts would indicate their plants to have been tetraploids, since our count is just half theirs. Based on these data, the base number for the genus would be x=10. Our voucher is Reveal 2248, west of Ash Meadows, along a dirt road southwest of the California-Nevada state line from California High- way 127 toward the abandoned Tonopah and Tidewater Railroad tracks, Inyo Co., California, 11 June 1970. Stanley a pinnata (Pursh) Britt. var. inyoensis (Munz and Roos) Reveal, stat. nov., based on S. pinnata ssp. inyoensis Munz and Roos, Aliso 3:115. 1965 « = 28. Figs. 19, 20. Our counts are in agree- ment with those recently obtained from root tips by Rollins and Riidenburg (1971). The voucher is Reveal 2277, Stewart Valley, Inyo Co., California, 12 June 1970. Prunus virginiana L. var. melanocarpa (A. Nels.) Sarg. « = 8. Figs. 21, 22, 23. This represents a new report, since no number for var. melanocarpa has been previously published. It is also a new count for the species, since the only report of var. virginiana we know of is n = 16 (Sax, 1931), and such a count may raise some questions as to the relationship between the western American en- tities, var. melanocarpa and var. demissa (Nutt.) Sarg., and the eastern chokecherry, var. virginiana. More data are needed and hopefully can be obtained during the next few years. Our voucher is Beatley and Reveal 11208, Upper Eden Creek Canyon, east slope of the Kawich Range, Nye Co., Nevada, 15 June 1970. Astragalus beatleyae Bameby. n=\\. Figs. 24, 25. This new count joins those of two other species in subsection Aridi of the March 1973 reveal, styer: plant chromosomes 23 section Inflati; all are n—\l (Barneby, 1964), which is the most common number in the North American species of the genus (see bolkhovskikh et al., 1969). Our voucher of this recently proposed species (Barneby, 1970) is Beatley and Reveal 10907, flatrock area -Z.d, miles north of Pahute Mesa Road on the Plateau Road, Pahute Mesa, Nye Co., Nevada, 4 June 1970. Lupinus argenteus Pursh. n = 24. Figs. 26, 27. This complex species is found throughout much of the western United States and is variously divided into species, subspecies, or varieties by several authors. C. P. Smith (1944) most likely called this plant L. corym- bosus Heller. Later, these plants were called L. alpestris A. Nels. (Clokey, 1951) or L. argenteus var. tenellus (Dougl. ex G. Don) D. Dunn (Dunn, 1956). More recently, these plants have been re- ferred to L. argenteus var. stenophyllus (Nutt. ex Rydb.) R. J. Davis (Beatley, 1969). We are still not clear what name should be applied to the collections from southern Nevada; L. argenteus is certainly the oldest name, although L. alpestris would have priority in our vicinity. The voucher is Beatley and Reveal 10909, south rim of Pahute Mesa, Nye Co., Nevada, 4 June 1970. Lupinus aridus Dougl. ex Lindl. /2=12. Figs. 28, 29. This species has been variously defined in recent years. C. P. Smith (1944) and Dunn (1956) have considered this form a distinct spe- cies, while Detling (1951), who monographed the group, referred L. aridus to a subspecific rank under L. lepidus Dougl. ex Lindl.; this latter treatment was followed by that of Hitchcock (1961), who used the varietal rank instead. To date, all entities referred to this species complex have had a single chromosome number, « = 24. Among the related species are L. lyallii A. Gray [or L. lepidus var. lobbii (A. Gray) C. L. Hitchc], « = 24 (Phillips, 1957); L. caespito- sus Nutt. ex Torr. and Gray [or L. lepidus var. utahensis (S. Wats.) C. L. Hitchc], ^2 = 24 (Heiser, 1963); and L. lepidus itself, with 72 = 24 (Phillips, 1957). Dunn has seen our voucher and other specimens from the same site and has identified our material as L. aridus. However, according to Detling's monograph, our material would be far out of the known range of the L. lepidus complex, the nearest confirmed site being in the high Sierra Nevada of California (L. lobbii A. Gray and L. lyallii) or in southeastern Oregon (L. aridus). One collection from northern Nye Co., Nevada, was tenta- tively assigned to L. aridus by Detling, but he was uncertain as to the exact identity of the collection. Based on the unusual chromo- some number, geographical disjunction, and the observable morpho- logical differences seen by both Beatley and Reveal, it would seem the plants from southern Nye Co., Nevada, may represent a new taxon. Our voucher is Beatley and Reveal 11138, southern Kawich Valley just below Gold Flat, Nye Co., Nevada, 14 June 1970. Angelica kingii (S. Wats.) Coult. and Rose. n = 22. Figs. 30, 31. This represents a new count for the species. Our voucher is Beatley and Reveal 11284, Cold Creek, eastern slope of Spring Mountains [Charleston Mountains], Clark Co., Nevada, 17 June 1970. 24 GREAT BASIN NATURALIST Vol. 33, No. 1 Gilia nyensis Keweal. n = 9. Fig. 32. A new count. This recently proposed species (Reveal, 1969), found in sandy places in southern Nye Co., Nevada, is related to G. hutchinsifolia Rydb., which also is reported to be /2 = 9 (Munz and Keck, 1959). The voucher is Beatley and Reveal 11147, Kawich Valley, Nye Co., Nevada, 14 June 1970. Castilleja viscidula A. Gray. ^2=12. Figs. 33, 34. Our count agrees with that recently published by Holmgren (1971). Our voucher is the southernmost population of the species: Beatley and Reveal 11068, Kawich Range, northwest of Kawich Peak, Nye Co., Nevada, 6 June 1970. Castilleja martinii Abrams var. clokeyi (Pennell) N. H. Holm- gren. n = 12. Figs. 35, 36. Our count represents a new ploidy level for var. clokeyi. Holmgren (1971) reported n = 24 for plants gathered on the east slope of the Spring Mountains (Holmgren and Reveal 2988), while our count, w^hich was obtained from specimens gathered on the west side of the same range, indicates a diploid situ- ation. This polyploid condition is to be expected, since Holmgren reports /2=12, 24, and 36 for var. martinii. The voucher is Beatley and Reveal 11282, Trough Springs, north end of Spring Mountains, Clark Co., Nevada, 17 June 1970. Arnica parryi A. Gray var. sonnei (Greene) Cronq. in Ferris. 72=19. Figs. 37, 38. Our count is the first reported for this taxon and represents a new ploidy level in the species. Ornduff et al. (1963) have reported var. parryi to be n = csi. 36, so it is possible that the typical variant might be n = 38, or twice the value found for var. sonnei. Our voucher is Beatley and Reveal 10975, near the stream at Longstreets Ranch, west of Kawich Peak, Kawich Range, Nye Co., Nevada, 6 June 1970. Laboratory work on this project was supported by NSF Grant GR-22645 to the senior author. Fieldwork in Nevada was conducted under Contract No. AT (04-1) Gen- 12 between the University of California and the Division of Biology and Medicine, United States Atomic Energy Commission, under the direction of Janice C. Beatley, research ecologist. We are grateful to her for her aid with this work. Literature Cited Barneby, R. C. 1964. Atlas of North American Astragalus. Mem. New York Bot. Gard. 13:1-1188. . 1970. A new Astragalus from Nye County, Nevada. Aliso 7:161-163. Beatley, J. C. 1969. Vascular plants of the Nevada Test Site, Nellis Air Force Range, and Ash Meadows. UCLA 12-705. Laboratory of Nuclear Medi- cine and Radiation Biology, University of California, Los Angeles. BoLKHovsKiKH, Z., V. Grif, T. Matvejeva. AND O. Zakharyeva. 1969. Chromo- some numbers of flowering plants. Academy of Sciences of the USSR, Lenin- grad. Clokey, L W. 1951. Flora of the Charleston Mountains, Clark County, Nevada. Univ. Calif. Publ. Bot. 24:1-274. Covas, G., and B. Schnack. 1947. Estudios cariologicos en Antofitas. Revista Argent. Agron. 14:224-231. DetlinGj L. E. 1951. The caespitose lupines of western North America. Amer. Midi. Nat. 45:474-499. March 1973 reveal, styer: plant chromosomes 25 Dunn, D. B. 1956. Leguminosae of Nevada, II — Lupinus. Contr. toward Flora Nevada 39:1-64. Heiser, C. B. 1963. Numeracion cromosomica de plantas ecuatorianas. Cienc. y Naturaleza 6:2-6. Hitchcock, C. L. 1961. Leguminosae. In Hitchcock, C. L., et al., Vascular plants of the Pacific Northwest. Univ. Wash. Publ. Biol. 17(3): 194-377. Holmgren, N. H. 1971. A taxonomic revision of the Castilleja viscidula group. Mem. New York Bot. Gard. 21 (4) : 1-63. MuNz, P. A., AND D. D. Keck. 1959. A California flora. University of Cali- fornia Press, Berkeley. Ornduff, R., p. H. Raven, D. W. Kyhos, and A. R. Kruckeberg. 1963. Chromosome numbers in Compositae. III. Senecioneae. Amer. J. Bot. 50: 131-139. Phillips, L. L. 1957. Chromosome numbers in Lupinus. Madrono 14:30-36. PiNKAVA, D. J., AND M. G. McLeod. 1971. Chromosome numbers in some cacti of western North America. Brittonia 23:171-176. Raven, P. H., D. W. Kyhos, and A. J. Hill. 1965. Chromosome numbers of Spermatophytes, mostly Califomian. Aliso 6:105-113. Reveal, J. L. 1965. Eriogonum. In Documented chromosome numbers of plants. Madrono 18:124. . 1969. New species in Eriogonum and Gilia from southern Nevada. Bull. Torrey Bot. Club 96:476-484. . 1971. A new annual Eriogonum (Polygonaceae) from southern Ne- vada and adjacent California. Aliso 7:357-360. 1972. Descriptions of new species and combinations in Eriogonum (Polygonaceae). Phytologia 23:163-178. Rollins, R. C, and L. Rudenberg. 1971. Chromosome numbers of Cruciferae. II. Contr. Gray Herb. 201:117-133. Sax, K. 1931. The origin and relationships of the Pomoideae. J. Arnold Arbort. 12:3-22. Smith, C. P. 1944. Lupinus. In Abrams, L., Illustrated flora of the Pacific States. Vol. II. Stanford University Press, Stanford. California. Stockwell, p. 1935. Chromosome numbers of some of the Cactaceae. Bot. Gaz. 96:565-570. Stone, D. E., and P. H. Raven. 1958. Eriogonum. In Documented chromo- some numbers of plants. Madroiio 14:237. INCIDENCE OF SPOTTED FEVER IN WOOD 1 ICKS OF UTAH RECREATIONAL SITES' C. Selby Herrin^ Abstract. — In 1964, 135 male and 223 female Dermaccntor andersoni Stiles were collected at 48 national forest recreation areas in Utah. Using guinea pigs immunological tests were performed on each of 62 samples to determine the presence of the Rocky Mountain spotted fever rickettsia. Thirteen of the samples (21 percent) were positive. These 13 samples contained 35 males and 34 females yielding an infection rate of at least 3.6 percent, assuming one infected tick was in each sample. The Rocky Mountain wood tick, Dermacentor andersoni Stiles, has been recognized as the principal vector of Rocky Mountain spot- ted fever (RMSF) in the Rocky Mountain states since shortly after the turn of the century. Much human suffering and many deaths have resulted from this disease in Utah as well as throughout the western United States. The presence of D. andersoni in recreational sites of the foothills, canyons, and mountains of the West offers a potential threat to the health of man. Expanding human population and increasing use of recreational facilities enhances this potential. A previous paper (Herrin, 1968) dealt with the prevalence of D. andersoni adults in national forest recreational sites of Utah. This research note reports the incidence of Rickettsia rickettsi (Wol- bach) in adult ticks collected from these recreational areas. Using a white flannel drag, 358 adult D. andersoni (135 males and 223 females) were collected from 48 recreational sites during the spring and summer of 1964. Ticks from 54 collections were divided into 62 samples, rinsed several times in sterile physiological saline, and preserved in vials of sterile, nonfat skim milk at below -30 C. Subsequently, the ticks were thawed, removed from the milk, triturated, and diluted with 10 ml of sterile, nonfat skim milk. One guinea pig per sami)le was inoculated with 1 ml of supernatant. Guinea pigs were observed for scrotal reactions, and rectal tempera- tures were taken every other day for 28 days. If the guinea pig died, a second was subsequently inoculated with the original super- natant. After 28 days, blood was taken and serum was extracted from the guinea pigs. Four complement fixation (CF) screen tests were performed on each sample, followed by an additional CF test to determine the antibody titer of positive samples. As a final test, all guinea pigs were inoculated with a lethal dose of virulent R. rickettsi to determine if they were immune to RMSF. In this test, one guinea pig per sample was challenged, and immunity was judged solely on survival. Abnormal temperatures in guinea pigs resulted after injections of triturated tick material from 18 samples. Seven showed abnormally >This paper is taken from a master's thesis submitted to the Department of Zoology and the Graduate School of Brigham Young University (1966). ^Center for Health and Environmental Studies, Brigham Young University, Provo, Utah 84001. 26 March 1973 HERRIN: SPOTTED FEVER 27 Table 1. Collections of D. andersoni adults determined to be positive for R. rickettsi. Number Complement National forest Collection and sex fixation Immunity Recreational site date in sample titer test* Cache Friendship and Spring 7 July 2M, 2F 32 + Fish Lake Bowery 4 June 2M, 2F 64 + City Creek 2 June 8M 128 + 7F 0 — Monrovian Park 2 June 6M 128 + 12F 0 Ponderosa 2 June 7M 0 — lOF 128 + Manti-Lasal Manti Community 10 July 7M 128 + 8M 0 lOF 0 — lOF 0 — Uinta Altamont 25 June 2M, 4F 64 + Aspen Grove 25 June 2M, 5F 64 Mutual Dell 25 June 2M, 3F 128 + Silver Lake Flat 25 June IM, 4F 64 Timpooneke 25 June 4M 128 + IIF 0 Whiskey Springs 10 June 2M, 3F 128 + Wasatch Sunset 7 July IF 128 +^ * Legend: + = immune; — = susceptible. high temperatures immediately and throughout the observation period, whereas 1 1 had an increase in temperature after several days. None demonstrated scrotal reactions or necrosis. Eight guinea pigs died within 3 to 14 days after inoculation. Thirteen of 62 serum samples (21 percent) yielded positive CF tests (see Table 1). The titers of these samples ranged from 1:32 to 1:128, whereas con- trols gave titers of 1:512. Eleven of 62 guinea pigs challenged with virulent spotted fever organisms survived, indicating immunity to RMSF. These 11 correspond with 11 of the 13 serum samples yield- ing positive CF results. In two cases of positive CF tests, guinea pigs were susceptible to the challenge. This close correlation be- tween results of the CF tests and the immunity tests suggests that 11, and probably 13, tick samples contained R. rickettsi. Three of the CF positive guinea pigs showed slightly abnormal temperatures. The remaining 15, and possibly 18, instances of abnormal tempera- tures are not attributable to spotted fever, since there was no correla- tion between samples that showed positive CF and immunity tests. Of the eight guinea pigs that died, two were from samples deemed positive by CF and immunity tests. Observations made at necropsy indicated that death was probably caused by bacterial infection. 28 GREAT BASIN NATURALIST Vol. 33, No. 1 The 13 samples of ticks positive for spotted fever represented 69 ticks (35 males and 34 females). If each positive sample yielded one tick infected with spotted fever, then 3.6 percent of all ticks col- lected were infected. Infected ticks were from sites in the northern half of Utah near human population centers (Provo, Salt Lake City, Ogden, and Logan). Infected ticks were collected from early June to late July and from every elevation range at which collections were made (6000 to 8800 ft). Further studies relative to incidence and virulence of R. rickettsi in wood ticks would help to determine the enzootic status of RMSF in Utah. Such studies should include identification of R. rickettsi by fluorescent antibody staining and recovery of rickettsiae in egg cultures prior to immunological tests with guinea pigs. Literature Cited Herrin, C. S. 1968. Dermacentor Andersoni in national forest recreation sites of Utah. Great Basin Nat. 28(1): 30-43. NOTES ON THE NESTING BEHAVIOR OF STENIOLIA ELEGANS (HYMENOPTERA: SPHECIDAE) Howard E. Evans ^ Abstract. — The nesting behavior of a small aggregation of the digger wasp Steniolia elegans near Fort Collins, Colorado, is described. The prey was found to consist primarily of bee flies (Bombyliidae), the egg being laid erect on the first fly placed in the cell. Nests were shallow, unicellular, and were closed at all times when the female was away. Several years ago Evans and Gillaspy (1964) reviewed what is known of the nesting behavior of digger wasps of the genus Steniolia (tribe Bembicini). One of the more common and widely distributed species, elegans Parker, was largely omitted from this review, since little was known concerning it. The present report is based on a two-day study of a nesting aggregation of this species just west of the city of Fort Collins, Colorado. This aggregation was discovered on 4 July 1971 on the slag pile of an abandoned mine on a south-facing slope. There were an esti- mated 50 individuals of each sex. Males were most active during the morning (8:30 a.m. -12: 30 p.m.), flying close to the ground in irregular patterns and landing here and there with their legs out- stretched and their antennae extended rigidly forward. Now and then they hovered, with their middle legs extended, over females occupied at their nests, and on several occasions attempted copula- tions were observed. The nests of the females were located in sloping, moderately fri- able but very stony soil. During the morning (9:30-11:30 a.m.) each female reopened her nest, first hovering over the entrance briefly and then digging through the closure, remaining inside for 20-60 seconds, then emerging and making a fresh closure. Then she hovered over the nest, flew off, and returned one or more times to hover again over the nest. These were evidently "inspection trips" serving to inform the wasp of the needs of the larva and perhaps to reinforce her memory of the nest site. Within a few minutes to an hour or two following the inspection, each female returned with her first prey, carried with the middle legs in the usual manner of Bembicini. Prey-laden females pro- duced a loud whine and often hovered briefly over the nest before digging through the closure. They remained in the nest very brief- ly (usually less than a minute) before coming out and restoring the closure. The prey consisted mostly of bee flies, as is common in the genus. The following species, in the numbers indicated, were taken from nests or from provisioning females (determinations by L. V. Knutson) : Bombyliidae Anthrax irroratus Say 1 Poecilanthrax signatipennis (Cole) 5 'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138. 29 30 GREAT BASIN NATURALIST Vol. 33, No. 1 Systoechus vulgaris Loew 4 Villa sinuosa Wiedemann 2 Villa spp. 8 Asilidae Holopogon atripennis Back 1 Syrphidae Eupeodes volucris Osten Sacken 1 Microdon coarctatus Loew 1 Volucella sp. 1 Provisioning was found to be fully progressive, the number of flies brought in per day being determined by the size of the larva. Nests dug out in the morning, before provisioning began, contained only the remains of flies. Several females were seen starting new nests in the afternoon, after having made the final closure of their previous nests. Much use was made of the mandibles in breaking through the soil and in dragging pebbles from the burrow and depositing them on the mound. From time to time, the female backed out of the entrance scraping soil, then remained on the mound for a few seconds, turn- ing to one side or the other while still scraping soil. As the burrow deepened, the wasp appeared less frequently (at 5- to 10-minute in- tervals), each time she appeared taking either a brief flight in a small loop or a much larger loop with a diameter of 3-6 meters. I observed no leveling movements following completion of the nest, and the mound of earth at the entrance was left intact, measuring about 3 cm wide by 4 cm long and 0.5-1.0 cm deep. Several hours were required for completion of a nest. When the nest was finished, each female made a thick closure from the outside by digging into that portion of the mound close to the entrance. Thus, newly completed nests could be recognized by the fresh mound which had been dug away on one side, facing the covered entrance. The egg was found to be laid erect on the side of the initial fly placed in the cell, in the common manner of Bembicini. Six nests were excavated, and all were found to be unicellular — a simple, oblique burrow leading to a terminal cell 4.5-9 cm deep (mean 6.6 cm). Some burrows were straight, while others were curved in various ways to avoid stones. Burrow length varied from 7 to 17 cm (mean 11 cm). None of the nests excavated aj)peared to be parasitized. It was evident that wasps did not spend nights or rainy periods inside their nests, and it is known that this species forms clusters on vegetation in the manner of other Stcniolia species (Evans and Gillaspy, 1964). However, an extensive search of surrounding ter- rain during the evening failed to reveal any such clusters. Literature Cited Evans, H. E., and J. E. Gillaspy. 1964. Observations on the ethology of digger wasps of the genus Steniolia (Hymenoptera: Sphecidae: Bembicini). Amer. Midi. Nat. 72: 257-280. A TAXONOMIC REVISION OF PHYSARIA (CRUCIFERAE) IN UTAH Sheldon B. Waite^ Abstract. — The Physaria species which occur in Utah are revised. This study is based on the treatment of tliis genus by Rollins (1939) and Mulligen (1967). Keys, descriptions, distribution maps, herbarium specimen citations, and discussions of the taxonomy of the genus are presented. The genus Physaria is confined to western North America. It occurs in the upper sonoran, transition, montane, and lower Canadi- an life zones, chiefly on high plateaus and lower mountain elevations (Rollins, 1939). This distribution extends from the southwestern United States to the Canadian border and from the Great Plains to the Sierra Nevada and Cascade mountain ranges. The genus, accord- ing to Rollins (1939), is of very recent origin. Mulligen (1967) verifies this view with studies he conducted using chromosome counts. He infers from his chromosome studies that several species are so closely related that a single species could include them all. One such cluster includes three of the four species which occur in Utah: P. acutifolia, P. chambersii, and P. newberryi. The other closely related species which belong to the group are P. didymocarpa, P. bellii, P. rollinsii, P. brassicoides, P. condensata, and P. floribunda. Since morphological characteristics distinguish these groups and all species within them are virtually isolated geographically, Mulligen (1967) believes they should be retained as individual species. Some integration can be observed morphologically in southern Utah between P. chambersii and P. acutifolia. Integration is also evident between P. chambersii and P. newberryi in the extreme southern counties of Utah. Mulligen (1967) corrected the name applied by Rollins (1939) to the Physaria of eastern Utah from P. australis to P. acutifolia. This was due to specimens studied from Colorado which have since been named P. rollinsii (Mulligen, 1966). Rollins described P. chambersii var. membranacea from speci- mens taken from Red Canyon, 16 miles west of Rryce Canyon Na- tional Park, Garfield County, Utah. The classification was based on the membranaceous nature of the siliques. Plants taken from this region and cultivated in different soils and environmental conditions showed a wide variation in the texture of the valves. Mulligen (1967) concluded that the specimens treated as var. membranacea did not, therefore, warrant varietal status. Specimens examined in this study are in the herbaria of Brigham Young University (bry), Utah State University (utc), and the University of Utah (ut). Those labeled only as Physaria sp. were lacking fruits and were not included in citations. ^Department of Botany, Brigham Young University, Provo, Utah 84601 . 31 32 GREAT BASIN NATURALIST Vol. 33, No. 1 Physaria inhabits dry barren regions where sunlight is intense and competition is minimal. The adaptation of this plant to survive xeric conditions is reflected in the heavy pubescence on the gray green leaves. Physaria (Nutt.) Gray Perennial, caespitose, silvery stellate; stems simple, arising later- ally on a somewhat elongated caudex; basal leaves usually numerous, often terminating the caudex or its branches in rosette form, petiolate oblanceolate to obovate or the blade rotund, entire dentate, or divided into segments; cauline leaves present, usually few, entire or dentate; inflorescence congested to somewhat elongated, usually elongating in fruit; pedicels rigid; sepals linear-oblong, pubescent, often cucul- late at apex; petals yellow or rarely purplish, usually spatulate, glab- rous; siliques didymous, pubescent, often highly inflated, apical sinus present; ovules 2-6 in each loculus; style persistent; seeds brown wingless. Key to the Species Known in Utah 1. Style less than 3.5 mm long P. newberryi Style more than 4 mm long 2 2(1 ) . Sinuses of silique equal above and below, valves nearly orbicular P. acutifolia Sinuses of silique equal above and below, valves not as above - 3 3(2). Silique highly inflated, 1.5-3 cm wide, valves mem- branaceous P. chambersii Silique moderately inflated less than 1.5 cm, plants loosely pubescent P. grahamii Physaria acutifolia Rydb. Physaria acutifolia Rydb., 1901, Bull. Torr. Bot. Club 28:279. Physaria didymocarpa (Hook.) Gray var. australis Payson, 1918. Ann. Gard. 5, 144. Physaria australis (Payson) Rollins, 1939, Rhodora 11, 408. Description. — Small, tufted })eronnial with a deep, rather slender taproot; basal leaves numerous 1.5-2 cm long, oblanceolate or obovate, acute entire or slightly wavy and very finely stellate stem; leaves rather few, oblanceolate; flowering stems 4-6 cm long, ascending or depressed; fruit small, obtuse or slightly cordate at the base, deeply divided above; cells inflated almost spherical, 4-6 mm in diameter; style about 5 mm long. This species differs from P. didymocarpa in its smaller acute leaves and in its smaller fruit, which is more deeply divided above as well as below. The siliques are characteristic features of this species, being highly inflated and strongly contracted toward the replum. The March 1973 WAITE: PHYSARIA 33 1. Physaria acutif olia Rydb. 2 , Physaria newberryi Gray 3. Physaria chambersii Rollins 4. Physaria srahamii Morton Figs. 1-4. Physaria distribution in Utah: 1, P. acutif olia Rydb.; 2, P. newberryi Gray; 3, P. chambersii Rollins; 4, P. grahamii Morton. 34 GREAT BASIN NATURALIST Vol. 33, No. 1 apical and basal sinuses are well developed and almost equal. The valves are rounded. Distribution. — This species grows in eastern Utah in dry, sparsely populated plant communities at ranges from 2,000 feet up to 9,000 feet in elevation. Members of the genus are found outside of Utah in northwestern New Mexico, western Colorado, southeast- ern Idaho, and most of Wyoming. Specimen citations include the following counties: Grand, B. F. Harrison 5968, 9-V-1933 (bry); San Juan, S. L. Welsh 2898, 31-V- 1964 (bry); Uintah, S. L. Welsh 49, 29-IV-1955 (bry); Emery, B. F. Harrison 8076, 8-V-1936 (bry); Kane, J. R. Murdock 403, 3-V-1962 (bry); Carbon, N. D. Atwood 1300, 29-IV-1968 (bry); Garfield, B. Maguire 19,098, 25-VI-1940 (utc); Duchesne, A. O. Garrett 7784, 19-VII-1938 (ut); Salt Lake, A. O. Garrett 6355a, 28- VI-1933 (ut); Tooele, J. Reveal 191 A, 15-VI-1961 (utc); Rich, J. W. Harrison no number, 24-VI-1927 (ut) (labeled as Cache County); and Cache, S. Flowers 223, VI-1924 (ut). Note: The University of Utah specimens used here were labeled by the pre- viously used name of P. australis or, in some instances, by the name of P. didymocarpa, which is not known to grow in Utah. Physaria newberryi Gray Physaria newberryi Gray, 1860, in Ives' Report Colo. River, pt. 4, 6. Coulterina newberryi (Gray) O. Kuntze, 1891, Revis. Gen. 2, 931. Physaria didymocarpa (Hook.) Gray var. newberryi (Gray) Jones, 1895, Proc. Calif. Acad. Sci. 2, 5. Description. — Perennial, caespitose, silvery stellate throughout; caudex simple branched; stems several to numerous, erect, simple, arising laterally, 0.5-1 dm long including the fruiting raceme; basal leaves obovate, incised, or merely dentate with broad teeth, slender petioled, 4-8 cm long, 3-4 mm wide; petals yellow ligulate, often truncate at apex, 10-13 mm long, 2-3 mm wide; fruiting raceme dense, 3-5 cm long; pedicels rigid, straight, divaricate, 5-10 mm long; siliques didymous, highly inflated, apical sinus broad, the shoulders angular and evenly compressed with appressed stellae; valve keeled on both outer margins, each valve 8-12 mm wide, 12-16 mm long; replum linear, acute at apex, 8-10 mm long, 1-1.5 mm wide; style 2-3 mm long; ovules 2-4 in each locule; seeds obovate, light brown marginless, 2-3 mm wide, 3-4 mm long. Distribution." — This species is found only in the extreme south- ern part of Utah growing in dry, open areas. It is found in north central and northern Arizona and in the extreme northwestern part of New Mexico. The distinctive characteristics of this species are its V-shaped apical sinus, short style, and straight-sided silique. Specimen citations include the following counties: Washington, S. L. Welsh and G. Moore 6865, 6-IV-1968 (bry); Iron, W. P. Cot- tarn 4724, 22-IV-1930 (ut); San Juan, B. F. Harrison 11600, 19-V- March 1973 waite: physaria 35 1950 (bry); Kane, Cottam 4289, 12-VI-1929 (bry); and Beaver, Cottam 8032, 8-VI-1940 (ut) (labeled as P. didymocarpa) . Physaria chambersii Rollins Physaria chambersii Rollins, 1939, Rhodora 41, 403. Physaria chambersii var. membranacea Rollins, 1939, Rhodora 41, 405. Description. — Perennial, caespitose, silvery stellate throughout; stems numerous from a simple caudex, arising laterally, erect or very often decumbent, simple, 5-15 cm long including the fruiting raceme; radial leaves entire or dentate, obovate to orbicular, slender petioled, 3-6 cm long, 1-2 cm broad; cauline few, entire, spatulate, often acute, 1-2 cm long, 3-6 cm wide; inflorescence rather lax; sepals linear-oblong, pubescent 6-8 mm long, 1 mm wide; petals yellow, spatulate, 10-12 mm long, 3-4 mm wide; fruiting raceme congested, 2-10 cm long; pedicels divaricate, slightly sigmoid, 8-15 mm long; siliques didymous, greatly inflated, evenly and often densely pubescent, often purplish at maturity, obtuse to slightly cordate at base; apical sinus deep and open, crests rounded; valves subreniform, each valve 1-1.5 cm long, 1 cm wide; style 6-8 mm long; ovules 2-6 (mostly 4) on each side of the replum; seeds orbicu- lar, flattened brown, 2-3 mm broad, 2-4 in each loculus margin. P. chambersii is related to P. newberryi but differs in having round-sided siliques and apical margins, a cordate or nearly truncate base, sinus crests rounded, style 8-13 mm long and replum 3-6 mm long compared to keeled apical margin siliques, truncate base, sinus crests decidedly angular, style 2-3 mm long, replum 8-10 mm long with acute apex and straight-sided valves for P. newberryi. Distribution.- — This species is found in the western half of Utah, most of Nevada, northwestern Arizona, extreme southeastern Oregon, and southwestern California. Specimen citations include the following counties: Utah, B. F. Harrison 8326, 18-V-1938 (bry); Juab, W. P. Cottam 7187, 28-V- 1937 (ut);* Tooele, C. Edwards 123, 15-V-1968 (bry); Garfield, W. O. Stanton 153, 28-VI-1930 (bry); Sanpete, A. O. Garrett no number, 29-IV-1911 (bry); Kane, Cottam 4289, 12-VI-1929 (ut); Summit, S. L. Welsh 6263, 20-VI-1967 (bry); Millard, W. P. Cot- tam 7347, 14-V-1939 (ut);* Washington, P. Plummer 5429, 9-V- 1939 (ut);* Sevier, R. Stevens 134, lO-IV-1966 (bry); Beaver, M. Milner 9212, 30-IV-1946 (ut); Iron, Cottam 3922, 31-V-1926 (bry); Piute, W. P. Cottam 7005, 15-V-1936 (ut);* and Salt Lake, S. Flowers 1390, 1927 (ut). Plants marked with an asterisk(*) were mislabeled as P. didymocarpa. A specimen labeled A. O. Garrett 8333a, 9-VI-1940 (ut), from Duchesne County, was sent to Reed Rollins, who stated in a letter now in the herbarium that this is a new species closely related to P. oregoni or a variety of the same. However, P. oregoni is found no farther south than central Idaho. No further information was avail- able concerning this specimen. 36 GREAT BASIN NATURALIST Vol. 33, No. 1 Physaria grahamii Morton Physaria grahamii Morton, 1937, in Ann. Carneg. Mus. 26:220. Description. — Perennial, caespitose, densely pubescent through- out with spreading stelae; stems simple, somewhat decumbent, about 1.5 dm long; basal leaves numerous, broadly oblanceolate to broadly spatulate, obtuse irregularly pinnatifid, 10-15 cm long, about 3 cm broad, distal lobes large and variable, cauline, few, dentate or rarely entire; pedicels divergent, 5-15 mm long; sepals linear-oblong, pubes- cent, about 5 mm long; petals yellow, spatulate, 6-8 mm long; siliques erect, didymous, inflated but not highly so, shallow sinus below, deep sinus above; replum linear-oblong, somewhat constrict- ed, ovules 2 on each side; style 6-8 mm long; mature seeds unknown. The type of this species is not altogether satisfactory because the fruits are immature. Its distinctiveness rests upon the fact that the entire plant is covered with loose, spreading stelae and large basal leaves deeply lobed along the margins. P. grahamii is at present known only from the type collection. Distribution.— This species is known from specimens collected at Chandler Canyon, Uinta Basin, Uintah County, Utah, 3-VIII- 1935. Graham 9976 (us type). References Index Kewensis. 1895 to present. Clarendon Press, Oxford, England. Kearney, T. H., and R. H. Peebles. 1951. Arizona Flora. University of California Press, Berkeley and Los Angeles. MuLLiGEN, G. A. 1966. Two new species of Physaria (Cruciferae) in Colo- rado. Can. J. Bot. 44:1661-1665. . 1967. Cytotaxonomy of Physaria acutifolia, P. chamber sii, and P. newbe-ryi (Cruciferae). Can. J. Bot. 45:1887-1898. Rollins, R. C. 1939. The cruciferous genus Physaria. Rhodora 41:392-415. TiDESTROM, I. 1925. Flora of Utah and Nevada. Government Printing Office, Washington, D.C. Welsh, S. L., and M. Treshaw and G. Moore. 1964. Guide to common Utah plants. Brigham Young University Press, Provo. Utah. TWO NEW SPECIES OF GYMNODAMAEUS FROM COLORADO (ACARINA: CRYPTOSTIGMATA, GYMNODAMAEIDAE)^ Tyler A. Woolley- and Harold G. Higgins'^ Abstract. — Gymnodamaeus plokosus and G. leurolomasus, n. spp., are de- scribed from soil at an ecological study site near Hayden, Colorado. Their rela- tionship to allied species is discussed. A review of the literature indicates that the first species of Gymnodamaeus were described by Berlese (1910, 1916). Woolley (1972) reviews the literature of known soil mites in the genus Gymnodamaeus and describes a new species, Gymnodamaeus chala- zionus, from the motmtainous areas of northern Colorado. The two new species described below are additions to this genus. The specimens of the first species described below were taken in soil beneath sage and grass along the undisturbed margin of a spoil bank near one of the coal strip-mining sites of the Hayden, Colorado, power plant. Comparisons of these specimens with others in the collections of the writers and with descriptions in the literature indi- cate that the species is new. It is described below and compared with other known species. Those species of Gymnodamaeus from North America with which this new species were compared are G. veriornatus Higgins, 1961, and G. chalazionus Woolley, 1972. The new species is smaller in size than the above species but averages very close to the size of G. gildersleeveae Hammer, 1952, and G. elegantulus Hammer, 1958; it is smaller than G. minor Banks, 1947, slightly larger than G. pearsei Banks, 1947, and much larger than G. quadricaudiculus Jacot, 1937. The details of this size comparison are given below with collection data and measurements of the new species. Gymnodamaeus plokosus, n. sp. (Figs. 1, 2) Diagnosis." — Most easily recognized by the crenulated posterior margin of notogaster with four nubbins, each bearing an ornate, curved hair (Figure 1); differs from G. gildersleeveae Hammer, 1952, in this feature as well as in the marginal placement of the pos- terior notogastral hairs; without reticulate pattern on dorsum of notogaster as in gildersleeveae, but with an arched, transverse bar on prodorsum just anterior to interlamellar hairs which is not present in gildersleeveae. Differs from G. quadricaudiculus Jacot, 1937, in the absence of the dorsal oval sculpturing with rays. The inter- lamellar hairs of the new species are short and most nearly resemble ^Research supported by the Yampa Project; Ecology Consultants, Inc. -Department of Zoology and Entomology, Colorado State University, Fort Collins, Colorado. ^Biology Department, Granger High School, Granger, Utah. 37 38 GREAT BASIN NATURALIST Vol. 33, No. 1 those of G. chalazionus Woolley, 1972. The sensillum of the new species is most similar to G. gilder sleeveae, broadly flabelliform and spined for its entire length. The trivial name plokosus is from the Greek, meaning "a lock of hair," or "curled," and refers to the four ornate, posterior notogas- tral hairs that arise from the nubbins and are so characteristic of the species. Description. — Color golden brown with a tuberculated, yellow- ish cerotegument giving a halo effect around the margins of body and legs; this cerotegument on prodorsum, notogaster, venter, legs, prodorsal hairs (except for interlamellar hairs), and notogastral hairs; prodorsum less than half as long as notogaster, broadly trian- gular in shape, with blunt, squarish rostrum; rostral and lamellar hairs of about equal length, decurved, covered with tuberculated integument; rostral hairs inserted in slight notches in anterolateral margin of prodorsum; lamellar hairs inserted in short apophyses at lateral margins, a transverse, sclerotized bar extending between in- sertions, hairs also covered with tuberculated integument, but tubercles of minute size compared to those of prodorsum; an arched, slightly U-shaped bar in posterior third of prodorsum immediately anterior to interlamellar hairs, extending in width about distance between interlamellar hairs; interlamellar hairs short, simple, in- Figs. 1-2. Gyninodaniaeus plokosus: 1, dorsiil aspect, legs partiallj emitted; 2, ventral aspect, legs omitted. March 1973 woolley, higgins: new mites 39 serted in inverted V-shaped apophyses posterolaterad of arched bar; pedotecta I and II as in Figure 1, with tuberculated surface due to cerotegument; pseudostigmata rounded, cup-shaped, with lip raised above surface of prodorsum, covered externally and internally with small tubercles; sensillum flabelliform, flattened in one plane, head broad, pedicel narrow, spined throughout length. Notogaster oval in shape, with nearly straight anterior margin; surface covered with tuberculated cerotegument; 8 to 10 large, more flattened tubercles near median area of anterior margin, other tu- bercles of surface smaller, more rounded; four short, sclerotized nub- bins posteriorly, forming a crenulated posterior margin; each nubbin with an inserted curved, ornate hair covered with small tubercles, a similar, less curved hair anterolaterad of nubbins on each side (placement of these six hairs apparently a specific characteristic in species of Gymnodamaeus) ; other hairs and surface features as in Figure 1. Camerostome somewhat triangular, with heavily sclerotized mar- gins; ventral setae, apodemata as seen in Figure 2; ventral surface with tubercles, but ventral tubercles only a third as large as dorsal tubercles; ventral setae simple, short; apodemata IV arched anterior- ly over genital opening as in other species of Gymnodamaeus; geni- tal and anal openings contiguous throughout widths; genital covers with cerotegument of tubercles, each cover with six genital setae; aggenital setae short, simple, laterad of genital opening; anal covers with tubercles, each cover bearing two simple anal setae; adanal setae ornate, covered with cerotegument of smaller tubercles (Figure 2). Legs heterotridactylous; surface of legs covered with cerotegument similar to dorsum; hairs of legs also covered with tuberculated cero- tegument, but tubercles of proportionately smaller size. Collections and measurements. — Ten males and two females of G. plokosus were collected by the writers at the Seneca site num- ber 2, Hayden, Colorado, lO-IV-1971. The males ranged in size from 432-372 i.l x 216-186 /t, with an average of 399 /i x 207 ^u. Four females and three males were collected from the same site on 8-VI- 1971 by the writers. These males ranged in size from 420-408 jx x 210-204 fi, averaging 410 /x x 208 /*,. The females ranged in size from 444-420 jx X 238-210 jx, with an average size of 432 /x x 228 fx. Additional specimens were taken in the project areas as follows: 3 males (425-415 /x x 212-205 /x) from under scrub oaks, 4 miles N Seneca Road, l-VIII-1971, by H. G. Higgins; 2 males (425-430 /t x 210-205 jx) from under chokecherries, one-fourth mile N Hayden Power Plant, l-VIII-1971, by H. G. Higgins; 1 male specimen (395 IX X 205 ix) under serviceberry on spoil bank, lO-IV-1971, by H. G. Higgins and T. A. Woolley; 1 male (380 /x x 210 /x) from beneath sagebrush, two miles S Yampa Valley Airport, 9-VI-1971, by H. G. Higgins and T. A. Woolley. Three specimens (2 females and 1 male) of this species were collected at Terry Falls, Trail Lake Ranch, Dubois, Wyoming, 20-Vn-1964, by H. and M. Higgins. They ranged in size from 432-420 /x x 210-204 /x. A single specimen was 40 GREAT BASIN NATURALIST Vol. 33, No. 1 also collected by H. and M. Higgins at Soapstone, Wasatch County, Utah, on 4-IX-1955. The specimen is so broken, however, that sex and size are not distinguishable. Two females were collected by T. A. Woolley in South Dakota, one (426 /<. x 204 ix) at Black Hills, 28-VIII-1968, and one (426 f/. x 316 /i) in the Nordbeck Game Pre- serve, 28-VIII-1968. The measurements of the width of all measurable specimens were taken at the level of legs IV. Generally speaking, females ap- pear to be larger and more elongated in shape than males. The drawings were made of a male specimen collected 10-IV- 1971. The type and a para type specimen will be deposited in the U.S. National Museum. The second new species was collected in soil beneath aspens and scrub oak in an undisturbed area about four miles south of the spoil banks mentioned above. Compared to G. plokosus, it is much larger. Gymnodamaeus leurolomasus, n. sp. (Figs. 3, 4) Diagnosis. — Larger size (.70 mm) median edge of genital plate smooth, lacking the interlocking dentes found in both G. chalazionus and G. veriornatus; each genital cover with seven setae; entire body and setae covered with a fine, granular cerotegument. The words leuros 'smooth' and loma 'border,' from the Greek, refer to the smooth median edge of the genital covers. This feature, along with the seven pairs of genital setae, distinguishes the new species from the other larger representatives of Gymnodamaeus from North America. Description.— Color dark reddish brown, entire dorsal and ventral surfaces covered with a granular cerotegument; prodorsum about half as long as notogaster, broadly triangular in shape; rostral and lamellar setae about equal in length, rostral setae inserted dorsal- ly, posterior to rostral margin but anterior to transverse sclerotized line between lamellar hairs; interlammellar hairs short, simple, coni- cal, each inserted near tip of curved, sclerotized, median ridge an- terior to pseudostigmata; in some specimens, a faint, transverse, curved sclerotized bar can be seen antoriomediad of interlamellar hairs (Figure 3); costulae absent; pseudostigmata rounded, with edge erected above surface of prodorsum; sensillum clavate, surface of club spined; entire organ slightly longer than prodorsum. Hysterosoma longer than wide, with a wrinkled surface; posterior margin with six setae and two pairs of dorsal setae of different sizes, with muscle scars and j)ores as shown in Figure 3; no distinct dorsal ornamentation as seen in G. veriornatus Higgins or G. ornatus Hammer. Camerostome oval in outline, montum broad, rutella narrowed anteriorly; ventral surface with apodemata and setae as shown in Figure 4; genital and anal apertures contiguous; genital covers smooth on medial edge, each cover with seven simple setae; anal covers much longer than wide, each cover with two simple setae; March 1973 WOOLLEY, HIGGINS: NEW MITES 41 Figs. 3-4. Gymodamaeus leurlamasus: 3, dorsal aspect, legs omitted; 4 ven- tral aspect, legs omitted. entire ventral surface covered with a granular cerotegument that obscures the location of coxistemal setae. Legs long, surface and hairs covered with cerotegument; all legs heterotridactylous, median claw larger than laterals. Collections and measurements.- — Length, .65-. 70 mm; width, .36-.41 mm. The type, a male, and eight other paratype specimens (six males, two females) were taken from under quaking aspens, 4 miles S Seneca Road, Hayden, Colorado, 8-X-1971, by H. G. Hig- gins. Five specimens were taken from under scrub oaks associated with aspens, 4 miles S Seneca Road, Hayden, Colorado, 1 -VIII- 1971, by H .G. Higgins. The type and one paratype will be deposited in the U.S. National Museum. Discussion and Ecological Notes It is interesting that two species of smaller Gymnodamaeus, G. gildersleeveae and G. plokosus, should be found in the project area. The latter species was first found under sagebrush along the edge of a spoil bank in rather dry soil. Subsequent collections have shown that this species is fairly well distributed throughout the western United States (Colorado, North Dakota, Utah, and Wyoming) as well as being found in the litter of several plant communities. For example, G. plokosus has been taken in rather dry soil and litter 42 GREAT BASIN NATURALIST Vol. 33, No. 1 under scrub oaks, serviceberry, sagebrush, and chokecherry in the Hayden project area. G. gildersleeveae has been found only under quaking aspens or in more moist scrub oak litter closely associated with aspens. This suggests that the populations of G. gildersleeveae prefer a more humid habitat, while G. plokosus is found mainly in a drier habitat. Gymnodamaeus leurolomasus must be listed among the larger North American members of this genus, but it is smaller than either G. veriornatus or G. chalazionus. It is nearer the size of G. ornatus Hammer from Canada but lacks the dorsal ornamentation so well known for that species. It differs from both G. veriornatus and G. chalazionus in that it lacks the interlocking dentes on the median edges of the genital plates and possesses seven pairs rather than six pairs of genital setae. This disparity in the number of genital hairs is considered a specific characteristic, inasmuch as the generic fea- tures match other representatives known from North America. Literature Cited Banks, N. 1947. On some Acarina from North Carolina. Psyche 54(2) : 110-141. Berlese, a. 1910. Brevi diagnosi di generi e specie nuovi di Acari. Redia 6: 346-388. — . 1916. Centuria prima di Acari nuovi. Redia 12: 19-67. Hammer, M. 1952. Investigations on microfauna of Northern Canada. Acta Arctica 4:108p. . 1958. Investigations of Oribatid fauna of Andes Mts.-I. The Argentine and Bolivia. Biol. Skr. Dan. Vid. Selsk. 10(1):1-129. HiGGiNs, H. G. 1961. A new beetle mite from Utah (Oribatei: Gymnodama- eidae). Great Basin Nat. 21 (1-2) :27-28. Jacot, a. p. 1937. Journal of North American moss mites. J. New York Entomol. Soc. 45(3-4) : 353-375. WooLLEY, T. A. 1972. A new species of Gymnodamaeus from Colorado. Great Basin Nat. 32(2):97-103. NEARTIC DESERT DECTICIDAE (ORTHOPTERA). PART II. A NEW GENUS AND SPECIES FROM ARIZONA Ernest R. Tinkliami ABSTRACT. — Plalyoplus, n. gen., is erected for P. gilaensis, n. sp., from the Gila Mountains, Yuma Co., Arizona. The method of preservation for these large, soft-bodied insects is described. The genus and species herein described was discovered in the Gila Mountains, 20 miles east of Yuma, Arizona, on 26 April 1958. The first specimen, found in a crevice of a giant boulder on a rocky ridge, eluded capture; however, other specimens were taken later. Platyoplus, n. gen. This new eremophilous, xerophilous, and petrophilous genus portrays characters that would place it between Ateloplus Scudder and Inyodectes Rentz and Birchim, with closest relationships indi- cated to Ateloplus. There appears to be little relationship shown to Eremopedes Cockerell and Pediodectes Rehn and Hebard. A review of the cereal features found in Ateloplus shows that four species, namely, notatus, luteus, hesperus, and splendidus have rather slender cerci with inner apical uncinate hook, while minor and schwarzi have shorter, quadrate cerci with inner apical tooth. The cerci of Platyoplus, with an inner subapical projection with uncinate tooth, is somewhat like that in Oreopedes Rehn and Hebard but is amply distinct in many ways. On the other hand, the features of the pronotum are quite ateloploid in nature; but even here the diagnostic differences are the shallower lateral lobes, which are quite outwardly flared and with those portions of the lateral lobes of the metazona tumidly raised and semitransparent above the greatly enlarged tym- panum, which in itself is diagnostic for the genus. Such features, the writer believes, warrant generic recognition. Experience based on many years of study convinces the author that any member of a particular genus must have the same characters as the other species of that genus. Thus the cercus of Ateloplus has an inner apical tooth, whereas in other genera such as Platyoplus and Eremopedes the inner cereal tooth is subapical or intermediate in position, al- though in some Eremopedes the cercus is also undulate and quite distinctive in character. Likewise, the pronotum in the various genera such as Platyoplus, Ateloplus, Eremopedes, Pediodectes, In- yodectes, and Oreopedes are different in each genus. Added to these and other characters not mentioned is the true tympanum, which is distinctive in the various genera mentioned above and portrays its greatest development in Platyoplus. Description. — Head broad and rather shallow, face with great- est breadth just below compound eyes, its breadth 1^4 times depth '81-441 Date Palm Avenue, Indio, California 82201 43 44 GREAT BASIN NATURALIST Vol. 33, No. 1 from vertex to clypeal suture. Fastigium with frontal costa broad, flat, abruptly terminated intermediately between the antennal sockets. Pronotum diagnostic, flat and broad, its breadth equal to its length, with shallow, reflexed lateral lobes; metazona shorter than prozona, its lateral lobes tumidly swollen over upper half of greatly enlarged true tympanum. Penultimate abdominal notite, with shallow U-shaped median groove. Cerci moderately slender, with an inner quadrate subapical prominence bearing an uncinate hook. Legs average, strong; caudal femora stout and shorter than in related genera, with both ventral keels bearing six widely spaced teeth in the apical half. Caudal tibiae fully spined, with numerous teeth along the entire ventral keels and with four pairs apically on the upper carinae of the caudal tibiae. Ovipositor stout, apical portion very slightly recurved, apex barely surpassing geniculae of caudal femora. Subgenital plate with a small, V-shaped median notch situated apically in the male; in female, deep median emargination, its margins very strongly raised with keels continuing basadly and convergent with an additional very high and narrow median keel connecting from base of deep, U-shaped emargination to base of plate; this character quite diagnos- tic for genus and not observed in related genera. Coloration pale brownish with very fine mottling of pale purplish dots and with genicular areas of caudal femora black. Type species. — Platyoplus gilaensis Tinkham, by monotypy. This genus is named after the very broad and flat pronotum which partially hides the very large true tympanum. The species is named after the Gila Mountains, that barren range of rocks about 20 miles east of Yuma, Arizona. Platyoplus gilaensis, n. sp. Description. — Male: head concolorous, unmarked; eyes sub- globose, their depth about equal to their breadth. Antennae fully twice body length. Head broad and short, facial breadth just below compound eyes 1 1/4 times depth from vertex to clypeal suture. Fastium angularly rounding into flat and smooth frontal costa, its margins convergent forward and roundly terminated medianly be- tween antennal sockets. Pronotum diagnostic, quite flat with breadth equal to length, very shallowly rounded into flaring lateral lobes; metazona nearly flat due to tumid swelling of metazonal lobe above upper half of greatly enlarged true tympanum; tympanum broadest in its apical half under swollen metazonal flange. Metazona shorter than pro- zona; prozona with a prominent but shallow, very broadly V-shaped sulcus, about two-fifths of prozonal length posteriad of anterior mar- gin. Anterior margin of the pronotum very smoothly arcuate for its entire breadth to the anterior lateral angle of lateral lobes; posterior margin almost squarely truncate, with at most only a slight sugges- March 1973 TINKHAM: DESERT ORTHOPTERA 45 9 p^^^f ,.1-7- ^^«^ropZi/5 gilaensis Tinkham: 1, Male holotype, profile view oiaSTl^'t. l"""^" ^°^°'^P" showing notite, cerci, titillator? 'arfd subgemt'l plate d, Lateral view of pronotum of male holotype; 4, Tympanum of male holo- rtv?eVTatf l'"- '^P'f^ ^' P-t--/ P0^*^°" ^f -bgeSital plat oFfemale 46 GREAT BASIN NATURALIST Vol. 33, No. 1 tion of emargination. Lateral lobes, seen from above, arcuately rounded, metazonal area outwardly bowed due to convexity of meta- zonal lateral lobe area above enlarged tympanum — a feature dis- tinguishing Platyoplus from all other desert decticids. Tympanum very large. Entire body surface smooth. Abdomen with ultimate notite bearing a moderate, median, V-shaped notch. Cercus moderate in build, broadest at base and very slightly tapered to apical two-thirds, inner margin of this area with a quadrate projection bearing a small, inner, slightly uncinate tooth. Subgenital plate with posterior lateral margins straight and with a median, posterior, V-shaped notch flanked outwardly with a short stylus. Plate distinguished further by a prominently rounded ridge extending anterior-laterally from base of each stylus, the ridges merging with basal portions of plate. Wings exposed for a length equal to that of the metazona. Forelegs with prominent procoxal spur; profemora with one to three very small inferior spines on exterior margin (leg forward) and four slightly larger spines on inner ventral keel. Protibiae with six or seven usually paired larger spines on ventral keels and two outer external spines dorsally, one of which is basal. Mesoleg with- out mesocoxal spine; mesofemora with five outer and two inner sub- apical small spines on ventral keels (legs in backward position); mesotibiae with six pairs of ventral spines and two pairs dorsally, as in protibiae. Hind legs with caudal femora bearing six spines on each ventral keel. Caudal tibiae heavily spined; 27-28 external and 24-26 internal strong spines on ventral keels and eight outer and inner attingent, widely spaced, slender spines on upper keels (ven- tral in position on folded legs). All tarsi three-segmented. Living Coloration (notes made 1965): Head with face faintly purple gray; eyes with incomplete cross-striations of purplish brown. Antennae reddish brown, nodes narrowly ringed in darker reddish brown. Pronotum dull grayish brown, tinged more definitely with burnt umber on central portions of metazona and more narrowly on front central edge of prozona. Tegmina pale purplish gray, with cells in exposed central portions piceous. Abdomen above citrus yellow, profusely and finely mottled with dark j)urplish brown, pos- terior marginal areas with about 12 washed or running spots of dark reddish brown. Sternites of thorax and abdomen citrus yellow. Fore and middle legs pale reddish brown; caudal femora pale reddish brown, outer pagina with a single central piceous band heaviest pos- teriorly; geniculae and pregenicular area dark piceous or almost black; spines of lower keel dark reddish brown. Caudal tibiae pale reddish brown, piceous at extreme base with both dorsal spine rows tipped in apical half with reddish brown; fewer scattered ventral spines similarly colored. Female: Slightly larger than male, but otherwise closely similar in coloration and spination of legs. Ovipositor slightly shorter than length of abdomen, rather heavy in build and very gently recurved, apex of ovipositor barely exceeding apices of caudal femora. Sub- genital plate diagnostic, with a large and deep quadrangular emargi- March 1973 tinkham: desert orthoptera 47 Fig. 8. Photograph of living Platyoplus gilaensis Tinkham on rock of native habitat. nation occupying two-fifths of total length of subgenital plate. Lateral margins of emargination prominently raised and extending basadly to terminate in an arcuate fashion about half the distance to base of plate, plus a prominent median keel running from center of quadrangular emargination to base of subgenital plate. HoLOTYPE. — Male, Telegraph Pass, Gila Mountains, Yuma Coun- ty, Arizona, 20 miles E Yuma, Arizona, 800 ft elevation, 30-V-1965 (night collecting on Creosote Larrea divaricata on steep rocky slopes; Ernest R. Tinkham). Caliper measurements in mm: body length 22.0; pronotum 7.75 long x 7.2 broad; tegmina exposed 2.0; caudal femora 24.3; antennae 51.5. Holotype deposited in the Tinkham Eremological Collection. Female allotype: Same data as holotype. Caliper measurements in mm: body length 24.9; body length to apex of ovipositor 39.2; ovipositor 16.0 x 1.6 in middle; pronotum 7.7 x 7.7:, antennae 51.6; caudal femora 24.9 x 5.2. Allotype in the Tinkham Eremological Collection. Male para types: Twelve males same data as holotype; 2, 25-1 V- 1960. Range in calipered measurements (mm): body length 21.6- 28.2; pronotum 7.2-8.1 (length) x 7.2-7.2 (breadth); caudal femora 23.1-24.0. Paratypes identical to the holotype male in every respect including coloration. Female paratypes: Twelve females same data as allotype. Range in calipered measurements (mm): body length 23.6-27.6; total body length to apex of ovipositor 38.5-42.0; pronotum 8.2-8.8 x 7.2-7.8 (breadth); caudal femora 24.1-26.0; ovipositor 15.7 x 15.7. 48 GREAT BASIN NATURALIST Vol. 33, No. 1 Paired para types will be presented to such major orthopterologi- cal museums as the following: Academy of Natural Sciences of Philadelphia, British Museum, California Academy of Sciences, Uni- versity of Michigan Museum of Zoology, Smithsonian Institution, and the Tinkham Eremological Collection. Biology.- — With the advent of the necessary winter rains in the Colorado Desert, it is believed that the nymphs hatch out either in the late fall or early winter. By late May the nymphs are in their last nymphal stadia and become adults in late June or very early July. The author has kept them alive in his home until late Novem- ber, but whether they would survive the torrid heat of their rocky, inhospital habitat in the Gila Mountains is questionable. Faunal Designation: Platyoplus gilaensis is a member of the Colorado Desert fauna, since the Gila Mountains represent the east- ern periphery of that desert. The area ranging from just east of the Gila Mountains and extending 100 miles to Gila Bend is Gila Desert. Both are eremological components of the Great Sonoran Desert, which is composed of seven different deserts. At or in the region of Gila Bend, the Gila Desert merges with the Sahuaro Desert. Floral Characteristics: In addition to Larrea divaricata, and Franseria dumosa growing sparsely on boulder-strewn slopes, Bursera microphylla is here more commonly represented than in any other desert sierra in the United States. Occasional Cercidium microphyl- lum and Hyptis emoryi, Hoffmanseggia microphylla, Franseria ilici- folia, Ditaxis lanceolata, Eneclia farinosa, and rarely Carnegica gigantea are all characteristic of these barren, but very interesting, mountains. Orthopteran associates: The orthopteran fauna is naturally quite impoverished, even in good years when up to three inches of rain may fall. However, since drought years have prevailed for the past decade, it is likely that some of the members present in the early forties are now extinct. Thus, Tanaocerus reported as koebe- lei in 1947 has not been seen since 1942. Likewise, Oedomerus coral- lipes, discovered in 1942 and reported for the first time in the United States in 1947, may also be close to extinction. Other orthop- terans taken in the forties, including Arphia aberrans, Tytthotyle maculata the Malpais Lubber, and even Trimerotropis p. pallidi- pennis, have not been seen since 1949. Two other orthopterans are known from the Gila Mountains, namely, Schistoccrca vaga and Capnobotes fuliginosus, both considered very rare. Host Plants: Platyoplus gilaensis was first discovered hiding in a crevice of a giant boulder by day, in 1958, and it was not until 1960 that the first specimens were taken by night collecting. The subadult nymphs were found feeding in creosote bushes, about the only plant available in that habitat. On another occasion, also on creosote, a fine colony was found in a small mountain canyon. As far as is known, creosote is the host plant in nature. Song: The stridulated note of Platyoplus is a soft "zee-zee-zee," long continued but scarcely audible beyond ten feet. The very large March 1973 tinkham: desert orthopter a 49 tympanum characteristic of Platyoplus may be necessary in order to hear the feebly stridulated song. Specimen preservation. Considerable has been written about the preservation of desert decticids, which are admittedly hard to pre- serve while maintaining colors that are close to those existing in the living creature. The usually large bodies filled with body fluids and the delicate coloration make good preservation difficult. The secret to fine preservation of body and color is the immediate gutting of the specimen upon its death. The decticids are collected alive in the field in specially designed cages, which the author has designed for the occasion. In this particular case, the large series of subadults was collected in the mountains in late May and removed to the author's home for proper study. Many biological notes were taken while the subadult specimens were developing into adults. In August, when the adults had been imagos for a month and all tissues were well hardened, small series at a time were killed with potassium cyanide. Each specimen killed was then immediately gutted by removing the alimentary tract and the gonads through a snip centrally or laterally on the three or four basal abdominal stemites. A little careful swab- bing should be done to remove excess liquid, but care should be taken not to damage or disturb the subcutaneous color structure. A small fluff of cotton can then be inserted through the incision, although this is not necessary. If cotton is used, care should be taken that it not be rolled, because if a roll of cotton were pushed into the tho- racic cavity, insertion of the insect pin could cause the whole to be pushed through the thoracic sternites, to the ruination of the speci- men. The specimens are now ready to be pinned, with legs and an- tennae arranged with the use of additional pins, on a sheet of balsa or white plastic cellulose. When the legs have been positioned by the pins and the antennae pulled back and positioned so that they lie along the dorsolateral line of the body, the creatures are ready for drying. The pinning block is now placed in a gas or electric range. If gas, the oven should be only at pilot heat; that is, no burners on and the temperature no more than 125 F. Here they are watched and removed after several hours when dry. Care must be taken not to use more than gentle heat, since color injury can result. After several hours the specimens should be dry and the coloration in life almost perfectly preserved. Living color notes should be made of the freshly killed creatures. One other method, relatively new, is also available. This is the freeze-dry method, where the creature is frozen immediately upon killing or death and left frozen for a long period of time, during which time it is completely desiccated by the cryoprocess. If this method is used, it may be necessary to relax the specimen or speci- mens overnight unless the specimens were pinned and legs arranged for the cryoprocess, which all depends upon the space available in the freezer. 50 GREAT BASIN NATURALIST Vol. 33, No. 1 References Hebard, M. 1925. XV. Expedition of the California Academy of Sciences to the Gulf of California in 1921. Dermaptera and Orthoptera. Proc. California Acad. Sci., Series 4, 12(15) :319-340, figs. 1-13. . 1934. Studies in Orthoptera which occur in North America north of the Mexican boundar}\ Part III. Revisionary data and new North American Decticinae. Trans. Amer. Ent. Soc. 30:31-54, pis. II, III. 1935. Studies in the Orthoptera of Arizona. Part II. A list of the Dermaptera and Orthoptera, with new records and corrections of the litera- ture subsequent to 1909. Trans. Amer. Ent. Soc. 61:269-316. Rehn, J. A. G., AND M. Hebard. 1920. Descriptoins of new genera and species of North American Decticinae (Orthoptera: Tettigoniidae) . Trans. Amer. Ent. Soc. 46:225-265, pis. 8-11. Rentz, D. C, and J. D. BuRCHiM. 1968. Revisionary studies in the Nearctic Decticinae. Memoirs Pacific Coast Ent. Soc. 3:1-173, 36 figs. TiNKHAM, E. R. 1941. Zoogeographical notes on the genus Atlanticus, with keys and descriptions of seven new Chinese species. Notes d'Entomologie Chinoise, Musee Heude, L'Aurore Univ., Shanghai 8(5) : 187-243, 1 map, 3 pis. . 1944. Biological, taxonomic and faunistic studies on the shield-back katydids of the North American deserts. Amer. Midi. Nat. 31 (2):257-328, figs. 1-28. . 1947. New species, records, and faunistic notes concerning Orthoptera in Arizona. Amer. Midi. Nat. 38(1) : 127-149, pis. 1-4. . 1948. Faunistic and ecological studies on the Orthoptera of the Big Bend region of Trans-Pecos Texas, with especial reference to the Orthopteran zones and faunae of midwestern North America. Amer. Midi. Nat. 40(3): 521-663, figs. 1-37. . 1960. Studies in desert sand dune Orthoptera. Part I. A new species of Plagiostira from eastern New Mexico, with key and notes. Great Basin Nat. 20(1 & 2): 39-47, 10 figs. . 1962. Studies in Nearctic desert sand dune Orthoptera. Part VII. A new dwarf race of Plagiostira gillettei from a Utah dune, with generic key. Great Basin Nat. 22(4) : 105-107. . 1972. Nearctic desert Decticidae (Orthoptera). Part I. A new genus from California. Great Basin Nat. 32(3) : 176-179. . 1972. Nearctic desert sand dune Orthoptera. Part XIV. A new Eremopedes (Decticidae). Great Basin Nat. 32(4): 223-228. AN UNUSUAL POPULATION OF SPIDERS IN UTAH Dorald M. Allred' An unusual population of spiders belonging to the species Neo- scona oaxacensis (Keyserling) was observed on West Mountain in Utah County, Utah, by Miss Mary Fenley and her mother, Mrs. Ed. J. Fenley, of Provo, Utah, while on a Labor Day outing in 1971. They kindly directed my attention to the phenomenon. West Mountain is situated in a north-south axis along the south- eastern side of Utah Lake. Its three major peaks have elevations of 6083, 6813, and 6904 feet, respectively. The level of the lake is at approximately 4487 ft. The predominant vegetation consists of various grasses along the foothill areas, and sagebrush and rabbit brush merging with a scattering of juniper trees at the higher eleva- tions. The spiders were densely distributed over several acres in the sagebrush and rabbit brush about midway up the mountain on the northern slopes. Only adults were present, and the females were predominant in number. The webs were not ornate as is typical of some of the other species of orb weavers. The radii of the webs were almost exclusively situated in some part of the individual shrub rather than in the spaces between shrubs. However, bridge lines frequently extended between separate plants. The bridge lines were of unusual strength, causing the limbs of plants to which they were attached to bend toward one another. As I walked between the plants and consequently broke the bridge lines, the tension of each line against my legs was almost like that of a lightweight twine. Several random counts were made of the unusually high popula- tion of spiders. These varied from 10 to as many as 60 individuals per square meter. In more than 20 years of extensive field investi- gations in temperate desert areas I have never observed such a high, localized population of spiders of any species. Some of my colleagues stated that they had noted that populations of spiders of other species in Utah were much higher this year than they had ever seen pre- viously; but they had not observed such a density as reported here. Dr. Willis J. Gertsch, who kindly identified the spiders and pro- vided some information on their biology, indicated that many years ago the same species occurred in great numbers on the old Saltair Pavilion on the shore of Great Salt Lake west of Salt Lake City. Incidentally, Ralph V. Chamberlin at that time named the species Neoscona saleria, now a synonym. According to Dr. Gertsch, Neoscona oaxacensis is a common orb weaver of the southwestern United States, ranging southward through Mexico to Panama. The species has had several names, probably the most familiar of which is A^. vertehrata McCook, based on specimens from California. Neoscona oaxacensis is probably the 'Department of Zoology, Brigham Young University, Provo, Utah 84601. 51 52 {}REAT BASIN N.'.TURALIST Vol. 33, No. 1 largest of the species of the "smaller" round-shouldered araneas with elongate oval abdomens. It is a species that often lives in clusters, with males, females, and the young stages living close together in webs. However, they are not necessarily tied by social habits, and their webs are single and not communal ones. Continued annual observations of this species on West Mountain in order to determine cyclical population fluctuations related to environmental influences would be desirable. SOME HELMINTHS FROM MINK IN SOUTHWESTERN MONTANA, WITH A CHECKLIST OF THEIR INTERNAL PARASITES Delbert L. Barber^'- and Lawrence L. Lockard^ Abstract. — Thirty-four percent of 100 mink examined from Gallatin and Madison counties, Montana, revealed the presence of Perostrongylus pridhami An- derson, 1962 (Anderson, 1963), in the lungs. This is the first report of P. prid- hami in the United States. Thirteen percent of tlie mink were infected with Taenia mustelae Gmelin, 1790. This is the first report of T. mustelae in south- western Montana. A checklist of internal parasites of Mustela vison is included. Parasites of mink in North America have been reported by nu- merous authors. No previous parasite surveys have been conducted on Mustela vison in southwestern Montana. The mink necropsied in this study were trapped in the Madison River, upper Gallatin River, and headwaters region of the Missouri River drainages during the winters of 1969 and 1970. A total of 100 mink were examined. These animals were eviscer- ated and the viscera were placed in plastic bags and frozen. An NaCl fecal flotation was prepared to determine the presence of cestode and nematode eggs or larvae prior to examination of the animals. The gastrointestinal tract was dissected using an enterotome device (Figure 1). The contents were washed onto a 200-mesh screen. Fig. 1. Enterotome Device. Seam ripper modified with plastic bead placed on end of point to facilitate intestinal incisions. then transferred to an illuminated tray for examination (Figure 2). Cestodes were fixed in 10 percent formalin, stained in Delafield's hematoxylin, dehydrated in ethanol, cleared in beechwood creosote, and mounted in HSR (Hartman-Leddon Co.). Lungs, liver, and kid- neys were dissected and each was placed in a jar with water and agitated on a mechanical shaker for 20 minutes. The contents were poured onto a 200-mesh screen, washed, transferred to an illumi- nated tray, and examined. Lung tissues that appeared to contain cysts or capsules were pressed between glass plates and observed under a dissecting microscope. Adult nematodes removed from lung parenchyma were fixed in 70 percent alcohol-5 percent glyc- erol and mounted in glycerol. Skulls, when available, were ex- amined for nasal nematodes. ^Present address; 3131 E. 12th, Casper, Wyoming 82601. ^Department of Zoology and Entomology, Montana State University, Bozeman, Montana 59715. 53 54 GREAT BASIN NATURALIST Vol. 33, No. 1 IHBJH^^^^HH^H ^ ■ - " f I'^BhHKI H ^^HB'l ^1 iiHmiii HHR^^^^v I^H ^ flHJ^^H ^^^1 Fig. 2. Illuminated tray used in postmortem recovery of helminth parasites. Results and Discussion Fecal examinations revealed that 34 percent of 100 mink were passing first-stage nematode larvae. Three male nematodes and portions of two females were dissected out of the lung tissue; larvae were removed from the gravid females. These larvae were compared with those found in the fecal samples and it was determined that they were of the same type. The adult males were identified as Table 1. Checklist of parasites reported from inink. Parasite Bibliography number Treniatoda (Flukes) Alaria freundi 85 A. minuta 18 A. mustelae 14, 15, 30, 31, 61 Apophallus muehlingi 86 Baschkirovitrema incrassatum 61 Cephalophallus obscurus 53 Cryptocotyle concava 57 C. lingua 57, 58 Enhydridiplostomum alarioides 61 Euparyphium beaveri 61 E. incrme 1, 51, 69, 90 ¥.. mclis 11, 30, 31, 38, 39, 46, 51 Euryhelmis monorchis 4, 30, 31, 61 E. pacificus 61, 80 E. pyriformis 61 E. squnrnula 38, 54, 61 Fasciola hepatica 58 March 1973 BARBER, LOCKARD: MINK PARASITES 55 Table 1 (continued) Fibricola cratera Metagonimoides oregonensis Metorchis conjunctus Nanophyetus salmincola Neodiplostomum lucidum Paragonimus kellicotti P. westermani Parametorchis canadensis Procyotrema marsupiformis Sellacotyle mustelae S. vitellosa Tocotrema lingua Troglotrema acutum 73 47, 61, 81 30, 31, 35 10, 38, 46, 78 73 2, 3, 12, 30, 31, 35, 38, 49, 61, 79, 93 37 38 61 30, 31, 61, 94 61, 84 86 38 Cestoda (Tapeworms) Diplogonoporus tetrapterus 71 Mesocestoides litteratus 30, 31 Moniezia sp. 58 Taenia mustelae { = tenuicollis) 5, 29, 30, 31, 32, 33, 44, 45, 52, 59, 61, 66, 70, 72, 77, 83, 92 Centrorhynchus conspectus Corynosoma hadweni C. semerme C. strumosuni C. sp. Macracanthorhynchus ingens Eimeria mustelae Isospora bigemina A( :anthocephala 61 58 28, 67 I 28, 58, 67 9 17, 61 Protozoa 43 79 Nematoda (Roundworms) Aelurostrongylus falciformis Ascaris sp. Capillaria mustelorum Crenosoma hermani Dictyocaulus filaria Dioctophyme renale Dranunculus insignis D. medinensis Epomidiostomum sp. Eustrongylus gigas Filaroides bronchialis F. mustelarum ( =martis) Gnathostoma spinigerum Heterakis isolonche Molineus patens Muslelivingylus skrjabini Muellerius capillaris Perostrongylus pridhami ( = Aelurostrongylus) Physaloptera sp. Seurocyrnea sp. Skrjabingylus nasicola Soboliphyme baturini Strongyloides sp. Trichinella spiralis 6, 8 1,35, 51, 61, 90 30, 31, 35, 58, 61, 68, 79 6, 26 58 1, 12, 21, 24, 30, 31, 35, 36, 40, 50, 55, 56, 60, 61, 75, 76, 79, 95, 96, 97, 98 19, 22, 35, 42 13, 20, 30, 31, 62 61 38 34, 79 1, 6, 25, 30, 31, 40, 48, 61, 88, 89 17, 38, 40, 99 61 30, 31, 46, 61, 64, 82 46, 74 58 6, 7, 8, 87, 88 30, 31, 63, 65 61 34, 41, 79, 91 46 1, 51, 58, 90 16. 23, 27, 35,40, 100 56 GREAT BASIN NATURALIST Vol. 33, No. 1 Perostrongylus pridhami Anderson, 1962 (Anderson, 1963), by Dr. Roy C. Anderson (personal correspondence). This parasite was previously reported by Anderson (1962) in Ontario, Canada. To our knowledge this is the first report of this species in the United States. Thirteen percent of the M. vison were infected with the cestode Taenia mustelae Gmelin, 1790. Identification of these specimens was confirmed by Dr. Gerald D. Schmidt (personal correspondence). This is the first report of adult T. mustelae in mink from Gallatin and Madison counties in southwestern Montana. Acknowledgments The authors express their appreciation to Dr. Roy C. Anderson for identification of the lung nematodes, to Dr. Gerald D. Schmidt for identifying the cestode, to Marvin Donahue for providing the mink viscera, to Dr. David E. Worley for assistance in preparing the manuscript, and to Donald H. Fritts for photographs. Bibliography 1. Allen, J. A. 1934. Parasites of fur-bearing animals. Proc. 5th Pan- Pacific Sci. Congr. 4:2981-2989. 2. Ameel, D. J. 1931. Life history of North American lung flukes of mammals. J. Parasitol. 18:261-268. 3. . 1934. Paragonimus: Its life history and distribution in North America and its taxonomy (Trematoda: Troglotrematidae). Am. J. Hyg. 19:279-317. 4. . 1938. The morphology and life cycle of Euryhelmis monorchis, n. sp. (Trematoda) from the mink. J. Parasitol. 24:219-223. 5. . 1942. Two larval cestodes from muskrat. Trans. Am. Micro. See. 61:267-271. 6. Anderson, R. C. 1962. The systematics and transmission of new and previously described metastrongyles (Nematoda: Metastrongylidae) from Mustela vison. Can. J. Zool. 40(5) :893-920. 7. . 1963. Further studies on the ta.xonomy of Metastrongyles (Nema- toda: Metastrongyloidea) of Mustelidae in Ontario. Can. J. Zool. 41:801-805. 8. Annderson, R. C, and A. Fyvie. 1961. Observations on Aelurostrongylus falciformis (Schlegel) of Mustela vison in Ontario. J. Parasitol. 47(4, sec. 2): 43. 9. Arro, I. V. 1966. Symposium on parasites and diseases of fish and aqua- tic invertebrates. (In Russian) Izchitestvo "Nauka." p. I 10. Raker, G. A. 1950. Troglolrema salniincola in mink. .1. Parasitol. 36:503. 11. Re.wer, p. C. 1941. Studies on tlie life iiistorv of Eupai yhiuin nielis (Trematoda: Echinostromidae). J. Parasitol. 27:35-44. 12. Beckett, J. V. and V. Gallicchio. 1966. Occurrence of the lung fluke, Paragonimus kellicotii Ward, 1908. in Ohio mink. J. Parasitol. 52:511. 13. BenBrook, E. a. 1940. The occurrence of the guinea worm. Dranun- culus medinensis, in a dog and in a mink, with a review of this para- sitism. J. Am. Vet. Med. Assoc. 96:260-263. 14. Bosma, N. J. 1931. Alaria mustelae, sp. nov.. a trematode requiring four hosts. Sci. 74:521-522. 15. . 1934. The life history of the trematode Alaria mustelae, sp. nov. Trans. Am. Micro. Soc. 53:116-154. 16. Chaddock, T. T. 1947. Veterinary problems of the fur ranch: Occur- rence of trichinosis in mink. Vet. Med. 42(8) :283. March 1973 barber, lockard: mink parasites 57 17. Chandler, A. C, and D. M. Melvin. 1951. A new cestode, Oochoristica pennsylvanica, and some new or rare helminth host records from Pennsylvania mammals. J. Parasitol. 37:106-109. 18. Chandler, A. C, and R. Rausch. 1946. A study of Strigeids from Michi- gan mammals, with comments on tlie classification of mammalian strigeids. Trans. Am. Micro. Soc. 65(4) : 328-337. 19. Cheatum, E. L., and A. H. Cook. 1948. On the occurrence of the North American guinea worm in mink, otter, raccoon, and skunk in New York State. Cornell Vet. 38(4) :421-423. 20. Chitwood, B. G. 1933. Does the guinea worm occur in North America? J. Am. Med. Assoc. 100:802-804. 21. Crichton, V. J. 1970. Dioctophyme renale in Manitoba mink. Can. J. . Zool. 48(3): 591-592. 22. Crites, J. L. 1963. Dracontiasis in Ohio carnivores and reptiles, with a discussion of dracunculid taxonomic problems. Ohio J. Sci. 63(1) :l-6. 23. Czarnowski, A., and G. Chylinski. 1964. Wlonsnica u lisow i norek [Trichinella in foxes and mink}. Medy. Wet. 20(9) :535. 24. Dearborn, N. 1946. Miscellaneous notes. J. Mammal. 27 (2): 178. 25. Dougherty, E. C. 1943. The genus Filaroides van Beneden, 1858, and its relatives. Preliminary note. Proc. Helminthol. Soc. Wash. 10:69-74. 26. . 1945. A review of the genus Crenosoma Molin, 1861 (Nema- toda: Trichostrongylidae) : Its history, taxonomy, adult morphology, and distribution. Proc. Helminthol. Soc. Wash. 12:44-62. 27. DuBNiTSKi, A. A. 1956. Prevent the occurrence of trichinelliasis at state mink farms. Karakulevodstvo i Zverovodstvo. 9(3) :60. 28. . 1957. The infestation of fur-bearing animals by acanthocephalan parasites. (In Russian) Karakulevodstvo i Zverovodstvo. 10(4):52-54. 29. DuNAGAN, T. T. 1957. Helminth parasites of Alaskan muskrats. Trans. Am. Micro. Soc. 76:318-320. 30. Erickson, a. B. 1944. Helminths of Minnesota Canidae in relation to food habits, and a host list and key to the species reported from North America. Am. Midland Natur. 32:358-372. 31. . 1946. Incidence of worm parasites in Minnesota Mustelidae and host lists and keys to North American species. Am. Midland Natur. 36(2): 494-509. 32. Freeman, R. S. 1956. Life history studies on Taenia mustelae Gmelin, 1790. and the taxonomy of certain taenioid cestodes from Mustelidae. Can. J. Zool. 34:219-242. 33. Gibes, H. C. 1957. On the role of rodents in the epidemiology of hydatid disease in the MacKenzie river basin. Can. J. Comp. Med. Vet. Sci. 21:287-289. 34. Gobel, F. C. and a. H. Cook. 1942. Notes on nematodes from the lungs and frontal sinuses of New York fur-bearers. J. Parasitol. 28: 451-455. 35. GoRHAM, J. R., AND H. J. GRIFFITHS. 1952. Diseascs and parasites of minks. Farmer's Bull. U.S. Dept. of Agri. No. 2050:41. 36. Graves, E. F. 1937. Dioctophyme renale in mink. J. Am. Vet. Med. Assoc. 90:531-532. 37. . 1937. Paragonimus westermani in mink. J. Am. Vet. Med. Assoc. 90:667-668. 38. Grini, O. 1933. Die Krankheiten des Minks. Pelztierzucht 9(2): 23-28. 39. Gupta, S. P. 1962. A redescription of Euparyphium melis (Schrank, 1788) Dietz, 1909, and Echinostoma revolutum (Froelich, 1802) Looss, 1899, parasitic in the intestine of mink {Mustela vison) and muskrat (Ondatra zibethica) from Canada. Indian J. Helminthol. 14(2):77-85. 40. Hanson. K. B. 1932. Parasites of mink and their control. U.S. Dept. Agric. Bureau Bio. Survey Bi-1235 (Mimeographed). 41. Hansoon, I. 1968. Cranial helminth parasites in species of Mustelidae. I. Frequency and damage in mustelids from Sweden. Oikos. 19(2): 217-233. 42. Hugghins, E. J. 1958. Guinea worms in a raccoon from South Dakota and in mink from Minnesota. J. Parasitol. 44:41-42. 58 GREAT BASIN NATURALIST Vol. 33, No. 1 43. KiNGScoTE, A. A. 1934. Eimeria mustelae, n. sp., from Mustela vison. J. Parasitol. 20:252-253. 44. KiNSELLA, J. M. 1967. Helminths of Microtinae in western Montana. Can. J. Zool. 45:269-274. 45. . 1968. Helminths of the western jumping mouse (Zapus princeps) in western Montana. J. Parasitol. 54:309. 46. KoNTRiMAVicHus, V. L. 1966. The helminth fauna of American mink acclimatized in the Khabarovsk Territory. In Jerusalem: Israel Pro- gram for Scientific Translations, pp. 241-242. 47. Lang, B. Z. 1970. Metagonimoides oregonensis from Mustela vison in Michigan. J. Parasitol. 56(5): 1031-1032. 48. Lankester, M. W., and R. C. 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Pathological changes in the kidney of mink due to infection with Dioctophyme renale (Goeze, 1782), the giant kidney worm of mammals. Trans. Am. Micro. Soc. 67:257-261. 57. McTaggart, H. S. 1958. Cryptocotyle lingua in British mink. Nature (London) 181 (4609) :651. - 58. . 1960. Helminths from the alimentary canal of farm mink in Britain. J. Helminthol. 34(3-4) :247-258. 59. Meggitt, F. J. 1924. Cestodes of mammals. London. 60. Meyer, M. C, and J. F. Witter. 1950. The giant kidney worm {Diocto- phyme renale) in mink in Maine, with a summary of recent North American records. J. Am. Vet. Med. Assoc. 116(878) :367-369. 61. Miller, G. C, and R. Harkema. 1964. Studies on helminths of North Carolina vertebrates. V. Parasites of the mink, Mustela vision. J. Para- sitol. 50:717-720. 62. Mohler, J. R. 1933. Zoological Division. Reports of the chief of the Bureau of Animal Industry, U.S. Dept. of Agriculture, pp. 42-47. 63. Morgan, B. B. 1941. A summary of the Physalopterinae (Nematoda) of North America. Proc. Helminthol. Soc. Wash. 8:28-30. 64. . 1943. New host records of nematodes from Mustelidae (Carni- vora). J. Parasitol. 29:158-159. 65. — . 1944. The Physaloptera (Nematoda) of carnivores. Trans. Wis. Acad. Sci. 36:375-388. 66. MoRozov, F. N. 1939. The parasitic worms of Mustelidae in Gorkiy's region. (In Russian) Trudy Gor'Kousk. Gosudarst. Pedagog. Inst. 4:3-44. 67. NuoRTEVA, p. 1966. Corynosoma strumosum (Rud.) and C. semerne (Forssell) (Acanthocephala) as pathogenic parasites of farmed mink in Finland. J. Helminthol. 40(1-2): 77-80. 68. Petrov, a. M., AND L. V. Panisheva. 1952. Phenothiazine therapy against Capillaria infections of the stomach and bladder of mink and sable. (In Russian) Karakulevodstvo i Zverovodstvo 5(1): 70-71. 69. Price, E. W. 1932. The occurrence of Euparyphium inerme (Fuhrmann) in North America. J. Parasitol. 18:311. March 1973 barber, lockard: mink parasites 59 70. Rausch, R. L. 1952. Studies on the helminth fauna of Alaska. XI. Hel- minth parasites of microtine rodents — taxonomic considerations. J. Parasitol. 38:415-444. 71. . 1964. Studies on the helminth fauna of Alaska. XLI. Observa- tions on cestodes of the genus Diplogonoporus Lonnberg, 1892 (Diphyl- lobothriidae). Can. J. Zool. 42(6): 1049-1069. 72. Rausch, R. L., and J. Tiner. 1949. Studies on the parasitic helminths of the North Central States. II. Helminths of voles {Microtus spp.). Preliminary report. Am. Midland Natur. 39:728-747. 73. Read, C. P. 1948. Strigeids from Texas mink, with notes on the genus Fibricola Dubois. Tr. Am. Micro. Soc. 67(2) :165-168. 74. Romanov, I. V. 1962. Mustelivingylus skrjabini, n.g., n. sp., from Mus- telidae. (In Russian) Lab. 12:94-97. 75. Sadighian, a., and F. Amimi. 1967. Dioctophyme renale (Goeze, 1782) Stiles, 1901, in stray dogs and jackals in Shahsavar Area, Caspian Region. Iran. J. Parasitol. 53:961. 76. ScHACHER, J. F., and E. C. Faust. 1956. 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A redescription of Taenia tenuicollis Rudolphi, 1819, and its larvae Cysticercus talpae Ruldolphi, 1819. Parasitol. 27:175-185. 84. Soganaders-Bernal, F. 1961. Sellacotyle vilellosa, a new Troglotrematid trematode from the mink in Louisiana. J. Parasitol. 47(6) :911-912. 85. Sprehn, C. E. W. 1932. Lehrbuch der Helminthologie, pp. 341-342. 716-717. 86. Sprehn, C. 1956. Seltene Helminthen in deutschen Farmnerzen. Deut. Tierarztliche Wochenschrift 63(33-34) : 342-346. 87. Stockdale, p. H. G. 1970. The development, route of migration, and pathogenesis of Perostrongylus pridhami in mink. J. Parasitol. 56:559- 566. 88. . 1970. Pulmonary lesions in mink with a mixed infection of Filaroides martis and Perostrongylus pridhami. Can. J. Zool. 48:757-759. 89. Stockdale, P. H. G., and R. C. Anderson. 1970. The development, route of migration, and pathogenesis of Filaroides martis in mink. J. Para- sitol. 56:550-558. 90. Swales, W. E. 1933. A review of Canadian helminthology I, II. Can. J. Res. 8:468-482. 91. . 1938. Skrjabingylus nasicola (Leuckart, 1842) Petrow, 1927, a nematode parasitic in the frontal sinuses of American Mustelidae. Livro Jub. Travassos, pp. 455-458. 92. Thienemann, J. W. 1906. Untersuchungen iiber Taenia tenuicollis Rudolphi, 1819, Mit Beriicksich tigung der iibrigen Musteliden Tae- nien. Arch. Naturgeschichte 1:227-248. 93. Wallace, F. G. 1931. Lung flukes of the genus Paragonimus in Ameri- can mink. J. Am. Vet. Med. Assoc. 31:229-234. 94. . 1935. A morphological and biological study of Sellacotyle mus- telae, n.g., n. sp. J. Parasitol. 21:143-164. 95. Woodhead, a. E. 1941. The life cycle of Dioctophyme renale, the giant kidney worm of man and many other mammals. J. Parasitol. (suppl.) 60 GREAT BASIN NATURALIST Vol. 33, No. 1 96. ■ — . 1945. The life history cycle of Dioctophyme renale, the giant kidney worm of man and many other mammals. J. Parasitol. (suppl.) 31:12. 97. . 1950. Life history cycle of the giant kidney worm, Dioctophyme renale (Nematoda), of man and many other mammals. Trans. Am. Micro. Soc. 69(l):21-46. 98. WooDHEAD, A. E., AND C. W. McNeil. 1939. Dioctophyme renale, the giant kidney worm occurring in mink, from the southern counties of Michigan. J. Parasitol. (suppl.) 25:23. 99. YosHiDA, S. 1934. Observation on Gnathostoma spinigerum Owen, 1836, cause of esophageal tumor in Japanese mink (Lutreola itatsi itatsi Temmenick, 1844), with especial reference to its life history. Japan. J. Zool. 6:113-122. 100. ZiMMERMANN, W. J., E. D. HUBBARD, L. H. SCHWARTE, AND H. E. BlESTER. 1962. Trichinella spiralis in Iowa wildlife during the years 1953 to 1961. J. Parasitol. 48:429-432. GRFAT BASIN NATURALIST Notice to Contributors Original manuscripts in English pertaining to the biological .atural history of western North America and intended for pubiica- on in the Great Basin Naturalist should be directed to Brighani oung University, Stephen L. 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Contributor ' ; the insi printed on the back cover of a recent The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University Volume 33 June 30, 1973 No. 2 THREE NEW SPECIES OF PALMOXYLON FROM THE EOCENE GREEN RIVER FORMATION, WYOMING William D. Tidwell,^ David A. Medlyn/ and Gregory F. Thayn^ Abstract. — Silicified, upright axes of Palmoxylon are abundant in the Green River Formation in Eden Valley, Wyoming. Three new species of Pal- moxylon, as well as the previously described Palmoxylon macginitiei Tidwell et al., were collected. These new species are P. edenense, P. contortum, and P. colei. They are compared to P. macginitiei and other anatomically similar Palmoxylon species. Palmoxylon edenense appears to be the most abundant species at this locality. Silicified remains of palm axes are very abundant in the Green River Formation in Eden Valley, Wyoming, near the collection site of Palmoxylon macginitiei (Tidwell et al., 1971). For the most part, these remains consist of axes in growth position surrounded by tough, silicified Chlorellopsis algae. However, one axis (P. colei) was col- lected lying down, as though it had fallen prior to fossilization. Weathered fragments eroded from these axes are found on the ground surface throughout the area. The upright specimens occur in groups of threes and are ar- ranged in either a triangular or linear alignment. Although the in- vestigated specimens in these groupings were of the same species (either P. macginitiei or P. edenense), no rhizomous or soboliferous connections were uncovered between them. With the exception of Palmoxylon contortum, the specimens vary from 6 to 12 inches in diameter and are generally 2 to 3 feet high. The incomplete horizontal stem of P. colei is nearly four feet in length. The preservation of the specimens is essentially the same as that discussed for P. macginitiei (Tidwell et al., 1971 ) . Palmoxylon edenense Tidwell, n. sp. Figures 1, 2, 5A, 9 Stem Central Zone: This zone is characterized by the usual ran dor arrangement of its vascular bundles. The bundle"^ vary in size from 700 to 900 jx high by 500 to 600 /i wide. There are approximately 85 bundles per cm-. The f/v ratio of these bundles is 3:1, although 'Department of Botany and Range Science, Bnglcdm Young University, Provo, Utah 84002 61 62 GREAT BASIN NATURALIST Vol. 33, No. 2 Fig. 1. Pnlmoxylon cdencnsc. Cros.s-sortions illustrating buiullos of flio (A) subdermal and (B) dermal zones (30X each). June 1973 TIDWELL ET AL: FOSSIL PALMS 63 \f- <%- ^V^iTi*^^ •/' *»«ii yfr --^^ Fig. 2. Palmoxylon edenense. A. Cross-section of the central zone (SOX). B. Enlarged bundles of the central zone (60X). 64 GREAT BASIN NATURALIST Vol. 33, No. 2 this may vary slightly (3.5:1.0). The fibrous bundle cap is oval to flabellate, having a shallow median sinus with rounded auricular lobes. Auricular sinuses are shallow to absent (Figure 2B). The cap is encased by one or two layers of tabular parenchjTna, whereas radial parenchyma surrounds the vascular tissue. The bundles are typically bivascular, with metaxylem elements 78 to 91 jx in diameter and protoxylem elements varying from 25 to 35 /x. The phloem is not structurally preserved. The ground tissue is tightly compacted, consisting of elongated parenchyma cells. Numerous fibrous bun- dles, 350 per cm", varying in diameter from 52 to 104 /j. are present but lack stegmata. SuBDERMAL ZoNE: Bundles of this zone tend to be irregularly oriented near the central zone and more or less regularly aligned close to the dermal zone. These bundles are similar in overall shape to those of the central zone, although their bundle caps are larger. The f/v ratio of the bundles is 5:1. There are approximately 100 bundles per cm^ ranging in size from 600 to 700 /x high to 400 to 500 /x wide. The reniform bundle cap is usually as high as it is wide. The bundles are bivascular, and the size and shape of their vessel elements is similar to those of the central zone. Cortical and Dermal Zone: The bundles are all regularly oriented with their caps towards the stem periphery. The bundles are more tightly compacted than are those of the subdermal zone, although they are not contiguous. There are approximately 205 bundles per cm-. Their fibrous bundle caps tend to be radially elongated. The bundle f/v ratio of this zone is 7:1. The bundles, which are 600 to 800 /i high by 280 to 470 /x wide, are smaller than those in the other zones. As in the other zones, the fibrous cap is surrounded by tabular parenchyma, and radial parenchyma en- sheaths the vascular tissue. The xylem contains two metaxylem ele- ments that average about 50 /x in diameter. The radial and tabular parenchyma composing the ground tissue is compact. Numerous fibrous bundles and leaf traces appear throughout his zone. Repository. — Brigham Young University, 916 (Holotype) Locality. — Eden Valley, Wyoming Horizon. — Green River Formation Age. — Eocene Palmoxylon colei, Tidwell, n. sp. Figures 3, 4, 5B Stem Central Zone: The vascular bundles of the central zone are irregularly oriented and loosely compacted. This zone has approxi- mately 115 bundles per cm-. The bundles range in size from 500 to 600 fjL in both height and width. Their f/v ratios vary from 2.5:1 to 3:1. The bundle cap fits Stenzel's Rcniforrnia group (Stenzel, 1904) by having rounded auricular lobes with shallow auricular sinuses. The median sinus is only slightly indented, giving the June 1973 TIDWELL ET AL: FOSSIL PALMS 65 t^ ^ % t Fig. 3. PalmoxyLon colei. Cross-sections of the (A) subdermal and ;,B) dermal zones (30X). 66 GREAT BASIN NATURALIST Vol. 33, No. 2 * - m'- V •^'» ". ~". ■ 'i^Ai •.Mt] % V ;'^| Fig. 4. Palmoxylon colei. A. Cross-section of the central zone (30X). B. Longitudinal section of a fibrous bundle demonstrating tJie stegmata along its surface (240X). June 1973 tidwell et al: fossil palms 67 bundle cap its characteristic kidney shape (Figure 4A). The fibrous caps are encased by one or two layers of tabular parenchyma, but the vascular portion is encircled by radial parenchyma. The bun- dles are typically bivascular and the metaxylem elements are 65 to 85 /x at their widest diameters. The protoxylem, when present, varies from 30 to 40 [x in diameter. The phloem is not structurally preserved. The ground tissue consists of thin-walled, tightly compacted, tabular and radial parenchyma, and has approximately 290 fiber bundles per cm-. These bundles exhibit characteristic stegmata, and their diameters vary from 78 to 120 /x (Figure 5B) . SuBDERMAL ZoNE: The bundles of this zone are irregularly aligned near the central zone and regularly aligned near the dermal zone. There are approximately 160 bundles per cm" in this region. The bundles are slightly larger than those of the central zone, being approximately 600 to 750 /x high and 500 to 650 /x wide. They have an f/v ratio of about 6:1, although this may vary slightly from bundle to bundle. The bundles are basically the same shape as those of the central zone, although the fibrous cap may approach a sagitate form in some. The presence of departing leaf traces that are at- tached to the vascular portion of several of the bundles is also notable. The bundles are commonly bivascular but may be trivascular. The metaxylem vessels are generally about 75 fi in diameter, although they vary from 65 to 85 /i. Fibrous bundles with their characteristic stegmata are present. Cortical and Dermal Zone: The bundles of the cortical and dermal zone are tightly compacted, with approximately 230 bundles per cm-. They are regularly aligned and have an f/v ratio of about 8:1. The bundle caps are reniform in shape but are elongated radially. The caps have rounded lobes and the auricular sinuses of the cap are shallow to nearly absent. The bundles are approximately 650 [X high and 400 /x wide. There are commonly two, occasionally three, metaxylem elements per bundle. Each element has an average diameter of 52 /x. This specimen was named for its collector, Mr. Walter N. Cole, of Provo, Utah. Repository. — Brigham Young University, 917 (Holotype) Locality. — Eden Valley, Wyoming Horizon. — Green River, Formation Age. — Eocene Palmoxylon contortum Tidwell, n. sp. Figures 6, 7 Stem The vascular bundles of all the zones are essentially the same size and shape, with the only observable difference being the degree of compaction. The number of bundles per cm- varies from 350 in the central zone to 500 in the dermal zone. They are tightly ap- 68 GREAT BASIN NATURALIST Vol. 33, No. 2 Fig. 5. A. Cross-section of a fibrous bundle of Palmoxylon edcnense. B. ibrous bundle of P. colei illustrating the stegmata surrounding the fibers. A fib Both are 160X. June 1973 TIDWELL ET AL: FOSSIL PALMS 69 Fig. 6. Palmoxylon contortum. Cross-sections of the (A) dermal and (B) central zones showing the extreme closeness of the bundles causing their distor- tion (30X). 70 GREAT BASIN NATURALIST Vol. 33, No. 2 J^i^y-W ^^^ I b #i'v /I I ^ v< .vT.l^ ^\' *•-■- -^ ^••t* -- ^V ,-^ ^» Fig. 7. Palmoxylon contortum. A. Cross-section of the dermal zone (SOX). B. Longitudinal section of scalariform thickenings on some protoxylem vessels (240X). June 1973 tidwell et al: fossil palms 71 pressed in all three zones but are distorted in the subdermal and dermal zones due to extreme bundle compaction. Where not com- pressed, the bundles are blocky to ovate in appearance. They typi- cally lack auricular lobes and sinuses but have definite median sinuses. These bundles commonly contain more vascular than fibrous tissue, with an f/v ratio between 1:1 and 1:1.5. Both the vascular and fibrous cells are surrounded by tabular parenchyma. Bundle size varies from 320 to 400 jx high and 300 to 500 /x wide. The bundles are characteristically bivascular, with the large rneta- xylem elements averaging about 55 /x in diameter. The ground tissue is tightly compact, consisting mostly of thin-walled columnar paren- chyma. Numerous sclerotic bundles ranging from 40 to 50 /x in diameter, without stegmata, are present. Specimens of this species were donated by Mrs. Marian Whitehead of Salt Lake City, Utah. They were approximately two inches in diameter. Repository. — Brigham Young University, 918 (Holotype) Locality. — Eden Valley, Wyoming Horizon. — Green River Formation Age. — Eocene Comparisons Palmoxylon macginitiei Tidwell et al. is similar in general ana- tomical features to both P. edenense and P. colei. P. contortum is dissimilar due to the disfiguration of the bundles and to the larger number of the vascular bundles per cm^ and its overall size. All of these species have scalariform perforation plates on their metaxylem elements. In order to delineate the above species, to give additional infor- matipn, and to correct an error in the previous description of Palmo- xylon macginitiei, a brief description of that species is included. P. macginitiei is characterized by the shape and size of its fibrovascular bundles (Figure 8A). They are oval, rarely rounded, and have a reniform bundle cap. The vascular portion of the bundle is blocky in appearance and is typically bivascular with tyloses common in the metaxylem elements (Figure 8B). The metaxylem vessels have scalariform perforation plates. The bundle size varies from 600 to 700 /A high by 450 to 500 /x wide in the central zone to approximately 420 /x high by 600 /x wide in the cortical and dermal zone. The numlDer of bundles per cm^ ranges from 104 in the central portion to 212 in the outer dermal zone. Palmoxylon edenense, although similar to P. macginitiei, should be considered a separate species because of the difference in degree of bundle compaction and bundle shape. P. edenense has an fV ratio of 3:1 to 3.5:1 in the central zone, whereas the ratio for P. mac- ginitiei is 1.5:1 to 2:1. Therefore, the bundle caps of P. edenense are larger than those for P. macginitiei. Hence, more definite auricular lobes and sinuses are present than in P. macginitiei, and the overall outline of the vascular portion is more rounded and not as blocky in appearance. The fibrovascular bundles of P. edenense are 72 GREAT BASIN NATURALIST Vol. 33, No. 2 . ^^''T)^-M Fig. 8. Palmoxylon macginitiei. A. Cross-section of the central zone (30X). B. Enlarged cross-section of bundles showing tyloses in the vessels (60X). June 1973 tidwell et al: fossil palms 73 less compacted, with 85 bundles per cm" in the central zone to 205 bundles per cm- in the cortical and dermal zone. Palmoxylon colei is unique in that it possesses stegmata, an ana- tomical structure which sets it apart from P. macginitiei. It also differs from the latter in bundle shape, with less vascular tissue in the central zone. Also both P. colei and P. edenense characteristically lack tyloses in the metaxylem elements. Palmoxylon edenense is similar to P. macginitiei. Therefore, the comparisons between P. macginitiei and other previously described Palmoxylon species (Tidwell et al., 1971) would also be valid for P. edenense. Although further study has shown that P. crassipes Unger is similar, it differs in that it has a complanate bundle cap and less vascular tissue in the central zone. The bundle cap of P. edenense tends to be less rounded and more flabellate with a deeper median sinus. P. hungaricum Greguss (Greguss, 1959) shows a close affinity to P. edenense but has larger and more widely spaced fibrovascular bundles. The presence of stegmata (stegmata is an anatomical feature found only in fossil palms [Stenzel, 1904]) in Palmoxylon cold restricts the number of similar species for comparison. Stenzel (1904) described three species exhibiting stegmata: P. densum, P. confertum, and P. astrocaryoides. These all differ from P. colei in their characteristic bundle shape, with P. densum also having dis- tinctly smaller bundles. Sahni (1964) described P. pondicherriense, a palm with stegmata from India. This species is dissimilar from P. colei in that the bundles in the latter are less numerous and that they have a reniform bundle cap instead of the lunarian form of P. pondicherriense. Palmoxylon contortum is a unique fine-bundled palm, and only a few species of Palmoxylon are even similar. P. liebigianum Shenk, which Sahni regards as synonymous with P. kvishna, also exhibits compact bundles, 150 per cm- in the central zone to 250 per cm- in the dermal zone, although the fibrovascular bundles of P. liebigianum are not appressed and disfigured, and its vascular bundles have lunarian-shaped bundle caps rather than reniform. P. ceylanicum Unger was considered by Stenzel (1904) as a variety of P. liebigi- anum. However, Sahni (1964) concluded that P. ceylanicum should be maintained as a separate species. He based this on the vascular bundles of P. ceylanicum being thinner and more crowded than in P. liebigianum. P. contortum differs from the above by its distinc- tive bundle shape, its degree of compaction, and its characteristic bundle distortion. Palmoxylon colei varies from P. simper i Tidwell, P. pristina Tidwell, and P. gustavsoni Tidwell et al. by having stegmata which the latter species lack; and P. contortum varies from them by its smaller size and by having more compressed, distorted bundles. Discussion In comparing Palmoxylon edenense and P. macginitiei with P. simperi Tidwell and P. gustavsoni Tidwell et al., it is a case of com- 74 GREAT BASIN NATURALIST Vol. 33, No. 2 paring species representing the basal portion of palm stems, or the so-called "stemless" varieties, from the Green River Formation with the distal portions of P. simperi and P. gustavsoni. Tomlinson and Zimmermann (1967) illustrated the different vessel types they had mascerated from a species of Sabal. The base of the stem had rather long vessels with scalariform perforation plates similar to the Eden Valley taxa; whereas those near the middle and top of the same axis had short vessels with simple pores like P. simperi and P. gustavsoni. P. simperi and P. gustavsoni, whose stem bases have not been ob- served, are from the same geographical area in central Utah (Tid- well et al., 1972). Scott et al. (1972) implied that all specimens from this area are of the same species; however, they most probably represent the middle to upper portions of P. simperi and P. gustav- soni. Thus, in comparing these latter two species to each other at about the same stem elevation and from the same zone (central), they are quite distinct and probably portray more than a delineation between form species. As further discussed by Tomlinson and Zimmerman (1967), the apical portion of a palm stem is always softer than the basal part. Consequently, when dead, they generally rot from the apex to base. The anatomy of the upper portions of the stem of living palms is easily sectioned and thus studied, whereas the basal portions are most often fossilized. Therefore, the matching of the anatomy of fossil and living forms becomes a dubious procedure, and comparisons be- tween the different parts of the stem may result in rather inaccurate conclusions. Therefore, comparisons between species representing the basal portion of the palm stems with those of the upper part would not be conclusive. The specimens of Palmoxylon macginitiei and P. edenense ap- pear to be complete palm axes. They do not indicate any truncation of the stems as would be anticipated if the stem had been originally taller (Figure 9). The bases of these specimens are surrounded with roots, whereas the middle and upper portions of these same axes have numerous attached petiole bases. The apices of these specimens consist of overlapping petiole bases. In progressing up the stem, each successive base becomes smaller and closer to the center of the axis. These stems may represent either very young stems or the so- called "stemless" palms similar to Nipa, some species of Acantho- cocos and Serenoa. Several forms of Palmae do not have any trunks above ground and thus are similar to the specimens from Eden Valley. In Serenoa repens, leaves appear to come right out of the ground, whereas Attalea cohune grows for many years before its trunk shows and has been mistaken as trunkless (McCurrah, 1960). Living Nipa palms balance enormous stemless rosettes on the treach- erous semiliquid mud of estuaries by means of a stout horizontal trunk (Corner, 1966). These spread by means of branching root- stocks. In the Philippine Islands, a Nipa marsh has been reported covering approximately 20,000 acres (McCurrah, 1960). The particularly numerous upright specimens from Eden Valley June 1973 TIDWELL ET AL: FOSSIL PALMS 75 Fig. 9. A specimen of Palmoxylon edenense drawn after collecting. Note the attached petioles. Roots are present in the lower portion of the specimen. 76 GREAT BASIN NATURALIST Vol. 33, No. 2 appear to represent the stemless types rather than the younger forms of taller trunked species. With the number of specimens at this locality, it would seem that intermediate stages of the trunked forms would also be present and these have not been observed. However, it is anticipated that further collecting will provide additional infor- mation concerning these forms and their possible placement. Acknowledgments The authors wish to acknowledge the assistance of the following: Naomi Hebbert and Paul Smith for aiding in preparing the illustra- tions and collecting, Mrs. Marian Whitehead of Salt Lake City and A. Daniel Simper of the University of California at Davis for their help in collecting and thin-sectioning of the specimens, Mr. Walter Cole of Provo for donating a specimen of Palmoxylon colei, and Dr. J. Keith Rigby of the Department of Geology and Dr. S. R. Rushforth of the Department of Botany at Brigham Young University for re- viewing the manuscript. References Corner, E. J. H. 1966. The natural historj- of palms. Univer. Calif. Press, Berkeley and Los Angeles. Greguss, p. 1959. A palmtrunk from the Miocene Coal Basin of Salgotarjan. Palaeobotany 8:19-21. McCuRRAH, J. C. 1960. Palms of the world. Harper and Brothers, New York. Sahni, B. 1964. Revisions of Indian fossil plants. Part III — Monocotyledons. Monogr. Birbal. Sahni Inst. Palaeobot. 1:1-89. Scott, R. A., P. L. Williams, L. C. Cr-mg, E. S. B.arghoorn, L. J. Hickey, and H. D. MacGinitie. 1972. "Pre-Cretaceous" angiosperms from Utah: Evi- dence for Tertiary age of the palm woods. Amer. Jour. Bot. 59:886-896. Stenzel, K. G. 1904. Fossile Palmenholzer. Beitr. Palaeont. Geol. Ost. — Ung. 16:107-287. Tidwell, W. D., a. D. Simper, and D. A. Medlyn. 1971. A Palmoxylon from the Green River Formation (Eocene) of Eden Valley, Wyoming. Botanique 2:93-102. Tidwell, W. D., D. A. Medlyn, and G. F. Thayn. 1972. Fossil palm materials from the Tertiary Dipping Vat Formation of Central Utah. Great Basin Nat. 32:1-15. Tomlinson, p. B., and M. H. Zimmermann. 1967. The wood of monocotyle- dons. Bull. Intemat. Assoc. Wood Anatomists 2:4-24. ON THE TAXONOMIC STATUS OF PLATYPODIDAE AND SCOLYTIDAE (COLEOPTERA) Stephen L. Woodi Abstract. — The reduction of Plat;yT)odidae and Scolytidae to subfamihes of Curculionidae by certain recent authors is questioned. Several fundamentally im- portant anatomical characters that could not possibly have been derived from Curculionidae are cited. Platypodidae (including Coptonotidae) is recognized provisionally as a derivative of Scolytidae; Scolytidae, with subfamilies Hylesi- ninae and Scolytinae (including Ipinae), is also recognized as a family. Proto- platypus, new genus, for P. vetulus, n. sp., a primitive platypodid from New Guinea, and Proiohylastes, new genus, for Pr. annosus, n. sp., a primitive scolytid from Queensland, are described. Apparently commencing in 1954 with Crowson (1967:155), a number of coleopterists have reduced the traditionally recognized families Scolytidae and Platypodidae to the rank of subfamily within the Curculionidae without establishing a foundation for this action. The recent generic classification of the Platypodidae (Schedl, 1972) neglected either to acknowledge or to respond to this change. In reviewing more than a dozen classifications of higher cate- gories within the Curculionoidea, the most striking feature noted in them is the lack of agreement on even the most fundamental divi- sions of the group, apparently due to a lack of detailed knowledge on the morphology of this enormous superfamily. It would appear that only Crowson's (1967) major division, based on separate or confluent gular sutures, is the only phylogenetically sound division of the group that has been presented. His classification of families in the first division (couplets 1 to 5) is reasonable (although the Oxycorynidae and Proterhinidae are unknown to me). His classifi- cation of the second division, groups having only one median gular suture (couplets 6 to 8), is questioned. The basis for this question is an unpublished and incomplete comparative anatomical study of the Curculionoidea that was started many years ago but interrupted due to the lack of specimens for dissection in critical groups. The following comments relating to the pregula and pregular sutures were drawn from that study. In the superfamily Curculionoidea five families (Crowson, 1967) have at least the posterior indications of widely separated gular sutures. In the Belidae these sutures continue separately to near the ventral apex of the rostrum, where they diverge and direct their course to or at least toward the anterior tentorial pits above the an- terior articulation of the mandible as in other insects. In the An- thribidae these sutures are obsolete except for minute irregularities on the margin of the occipital foramen; in Nemonychidae and ap- parently in Oxycorynidae (not seen) and Proterhinidae (not seen), separate sutures (Crowson, 1967: Figures 201-202) extend anteriorly 'Department of Zoology, Brigliam Young University, Provo, Utah 84602. Scolytidae Contribii- tion No. 49. 77 78 GREAT BASIN NATURALIST Vol. 33, No. 2 to the posterior tentorial pits. In all other Curculionoidea the gular sutures are confluent, with only one median suture (usually) visible from near the occipital foramen to the single, median, posterior tentorial pit at the ventral base of the rostrum. Apparently all workers have overlooked the minute postgula on the margin of the occipital foramen at the base of the gular suture (Figures 1-19). Some workers, realizing that a pregula should be present (c.f. Hop- kins, 1909:16, and Stickney, 1923), either created pregular sutures for their illustrations of curculionids or misinterpreted longitudinal ridges on the rostrum as being pregular sutures. In a survey of several hundred genera of Attelabidae, Brenthidae, Apionidae, and Curculionidae (s. str.) I was unable to find any representative hav- ing pregular sutures accompanied by internal ridges that resulted from the inflection of the cuticle to form those sutures. However, pregular sutures are conspicuously present in all Platypodidae and in all Scolytidae except a few Micracini, etc., that bore into exceed- ingly hard wood (in which case the gular suture is also obliterated). It apparently has been traditional among those coleopterists who have considered the matter (Crowson, 1967:155; Schedl, 1972) to presume that the Scolytidae were derived from the tribe Rhyncolini of the subfamily Cossoninae and that the rostrum of the ancestral form was lost when scolytid habits were established. Yet in no truly primitive platypodid or scolytid is there a rostrum resembling that of Cossoninae. Furthermore, no representative of the Rhynco- lini or of any other cossonid known to me (including an apparently unnamed Australian cossonid genus with absolutely no rostrum whatever and large mandibles) has pregular sutures (Figure 4). In this latter character I have found no intergradation, either ex- ternally or in the much more complex internal structure. The recog- nition of this fact suggests that other indicators of phyletic relation- ship be examined. Primitive Platypodidae, primitive representatives of all primitive tribes of Scolytidae, and many Cossonini have interstriae 10 on the elytra broad throughout its length to near the elytral apex. In all Rhyncolini known to me the portion of interstriae 10 posterior to the level of the hind coxa is strongly constricted or entirely obsolete. Therefore, based on this character in living material examined, there is no possibility that the Rhyncolini could have given rise to either the Platypodidae or Scolytidae. Crowson (1967:158) indicated that all adult Curculionoidea hav- ing one gular suture, except Attelabidae, have the adult maxillary palpus 2- or 3-segmented. Browne (1971:49) reported a 4-segmented maxillary palpus in Austroplatypus BrowTie, a platypodid. It is very doubtful that this geiuis was derived from an ancestral form having a 3-segmented maxillary palpus. In all Rhyncolini and most (all?) Cossoninae known to me there is a conspicuous spine on the margin of the oral fossa that arises between the posterior margin of the mandible and the base of the maxilla (Figure 4). There is no com])arable structure or irregularity in this area in Platypodidae or Scolytidae (Figures 12, 14, 19). June 1973 wood: platypodidae and scolytidae 79 1. Stenoscelis 5. Hylurgops 2. Stenoscelis 6. Hylurgops 7. Hylurgops 10. Schedlarius 3. Stenoscelis 11. Schedlarius 12. Gnathotrupes 8. Platypus 4. Rhyncolus Figs. 1-12. Head capsules of beetles with the tentorial appartus indicated by dotted lines: 1-3, Stenoscelis brevis (Boh.), Cossoninae, 1 lateral, 2 posterior, and 3 dorsal aspects; 4, Rhyncolus knowltoni (Thatcher), Cossoninae, ventral aspect; 5-7, Hylurgops rugipennis (Mannerheim), Hylesininae, 5 lateral, 6 pos- terior, and 7 dorsal aspects; 8, Platypus lucasi Chapuis, Platypodinae, dorsal aspect; 9-11, Schedlarius mezicanus (Duges), Coptonotinae, 9 lateral, 10 posterior, and 11 dorsal aspects; 12, Gnathotrupes sp., Scolytinae, ventral aspect. 80 GREAT BASIN NATURALIST Vol. 33, No. 2 Figs. 13-19. Head capsules of beetles with the tentorial appartus indicated by dotted lines: 13-16, /p? mcxicanus (Hopkins). Scolytinao. H lateral. 11 ven- tral, 15 posterior, and 16 dorsal aspects; 17-19, Platypus lucasi Chapuis. Platy- podinao. 17 lateral, 18 posterior, and 19 ventral aspects. Kuschel (1966:6) suggested that a series of sj)iiies on the lateral margins of the tibiae in Scolytidae and in the Araucariini (Cosso- ninae) indicate a relationship between these groups. Evidently it was not recognized that the most primitive genera of Scolytidae lack these spines and have tibiae more nearly like the (>ossonini or even like certain Brenthidae. The supernumerary tibial spines in these June 1973 wood: platypodidae and scolytidae 81 groups apparently were acquired independently as they adapted to a common niche and, therefore, are not primitive. Kuschel also called attention to the similarity in habits of the Araucariini and Scolytidae. In the two species of Araucariini (presumably Coptoco- cynus spp.) examined in the field, the habits do not resemble in any way those of more than 2000 species (about 30 percent of the known fauna) of Scolytidae I have studied in the field. The habits of certain neotropical Rhyncolini (unidentified) are infinitely more similar to primitive scolytids. Larval characters that separate the Scolytidae from Curculionidae have not been reported, presumably due to inadequate research rather than to the absence of characters. Most larval Platypodidae have the clypeus reduced or absent and the labrum somewhat en- larged. The cercus-like processes in Trachyostus (Browne, 1967: Figure 20) and the tenth abdominal segment of Dolgopygus (Browne, 1967: Figure 23) require investigation. It is my contention that the Scolytidae and Platypodidae repre- sent a phyletic line that diverged from other Curculionoidea before pregular sutures were lost and before a rostrum developed. The Cossoninae (Figures 1-4) are true curculionids in the structure of the gular area, tentorial apparatus, head, legs, and body form, and probably resemble scolytids largely because they independently oc- cupied the same or a similar niche. While the scolytids and platypo- dids clearly fall within the Curculionoidea, it is as logical to recog- nize them as an independent group as it is any other family within this superfamily. There is great difficulty, however, in separating the Scolytidae from the Platypodidae, and I find it difficult to give independent family status to the latter group. In order to emphasize the significance of the above items, it is necessary to describe the most primitive platypodid and the most primitive hylesinine scolytid known to me. Protoplatypus, n. gen. The phloeophagous habit and normal tarsi of this remarkably primitive genus suggest that it should be placed in the family Scoly- tidae; however, the head, tibiae, pronotum, and other characters in- dicate a closer relationship to primitive Platypodidae. Although not closely related to either genus, its phylogenetic position probably lies between Mecopelmus Blackman (Platypodidae) and Craniodycticus Blandford (Scolytidae). For convenience of reference I tentatively place it in the Mecopelmini (Platypodidae) . Characters of particular significance in phylogeny found in this genus include the cylindrical, 3-segmentod maxillary palpus, an antenna intermediate between that of Platytarsulus (Platypodidae) and Craniodycticus (Scolytidae), a protibia intermediate between that of Schedlarius (Platypodidae) and Protohylastes (Scolytidae), an eye similar to that of platypodids, and tarsi similar to those of scolytids. The phloeophagous and polygamous habits are, for the most part, typical of scolytids. 82 GREAT BASIN NATURALIST Vol. 33, No. 2 Description. — Head about as in Schedlarius Wood; eyes sub- circular, moderately convex; pregula rather large, as in most Platy- podidae; antennal scape long, slender, funicle 5-segmented, club rather small, somewhat flattened, entirely devoid of sutures, gla- brous except for a marginal fringe of abundant, short hair (much as in Platytarsulus Schedl but much more restricted to actual mar- gin). Pronotum elongate, sides constricted as in most Platypodidae; precoxal area of prosternum elongate, almost half as long as entire prosternum, precoxae small, very widely separated. Scutellum sub- acutely pointed behind. Elytral bases rounded, not precipitous, striate, interstriae 10 attaining declivital area; posterior area decli- vous, sculpture simple. Protibiae of platypodid type; tarsi with seg- ment 1 only slightly longer than 2 or 3, very slightly shorter than 5, entire tarsus slightly longer than tibia, all segments cylindrical. Type species. — Protoplatypus vetulus Wood, described below. Protoplatypus vetulus. n. sp. Figures 20, 21, 26 This species is distinguished from other known forms by char- acters summarized in the above description of the genus. It could be placed in either the Platypodidae or Scolytidae, depending upon which characters are emphasized. Male. — Length 1.2 mm (paratypes 1.2-1.5 mm), 4.0 times as long as wide; color light brown. Frons strongly, rather evenly arched from epistoma to vertex, Figs. 20-21. Protoplatypus vetulus Wood: 20, dorsal aspect; 21, lateral aspect of head and prothorax. leg omitted. Figs. 22-24, Protohylastes annosus Wood: 22, lateral aspect; 23, dorsal aspect; 24. anterior aspect of left metathoracic tibia. June 1973 wood: platypodidae and scolytidae 83 rather broadly convex, somewhat intermediate between Schedlarius and Coptonotus Chapuis; surface reticulate, with sparse, fine, shal- low punctures; vestiture very sparse, hairlike; eye broadly oval to subcircular, mioderately convex; pregula rather large, as in Platy- podidae; maxillary palpi cylindrical, 3-segmented. Pronotum 1.5 times as long as wide; widest on basal third, sides conspicuously constricted just in front of middle, anterior margin only slightly narrower than base; surface shining, reticulate at base and sides, longitudinally, subreticulately strigose on discal area, punctures minute, shallow, moderately close. Glabrous. Elytra 2.3 times as long as wide, 1.6 times as long as pronotum; sides straight and parallel on basal three-fourths, broadly rounded behind; scutellum acute; elytral bases rather weakly, not precipi- tously, elevated; entire surface strongly reticulate; striae not im- pressed, punctures small, moderately deep, spaced within row by about two diameters of a puncture; interstriae twice as wide as striae, punctures very small, widely, regularly spaced. Declivity confined to posterior fourth, broadly convex; striae about as on disc; interstriae 1 and posterior half of 9 moderately, continuously elevated and meet at sutural apex, 3 less strongly elevated and ending before attaining 9. Vestiture of sparse, very minute, almost scalelike setae. Female. — Similar to male except frons with a triangular area on lower two-thirds smooth, shining, its upper margins marked by an irregular row of small punctures; scape with setae near apex distinctly longer. Type Locality. — Five miles or 8 km NW Bulolo, Morobe, New Guinea. Type Material. — The male holotype, female allotype, and 96 paratypes were collected at the type locality in the LATEP logging area, on 9-VIII-72 from the bole of a recently cut Har pallia pedi- cellaris, by S. L. Wood. The parental galleries were in the cambium region but did not engrave the wood; they were of the radiate type, and the beetles appeared to be polygamous. Larval mines were parallel to the grain of wood and were visible on the inner surface of peeled bark. The holotype, allotype, and several paratypes are in the Aus- tralian National Collection at Canberra; the other paratypes are in the British Museum (Natural History) and in my collection. Protohylastes, n. gen. This genus superficially resembles Hylurgops LeConte or Pseudo- hylesinus Swaine, except that the tibiae are completely different from any other genus of Hylesininae and the protibiae are more nearly like a curculionid (Curculionoinae) than a scolytid. The bases of the elytra are similar to those of Hylurgops, without a defi- nite marginal row of crenulations, except that the submarginal crenu- lations are even more poorly developed. 84 GREAT BASIN NATURALIST Vol. 33, No. 2 Diagnosis and Description. — Head much as in Pseudohyle- sinus; eye oval, short; antennal scape short, funicle 7-segniented, shorter than scape, club as long as scape, small, subconical but dis- tinctly flattened, entirely devoid of sutures. Prothorax about as in Pseudohylesinus except coxae rather widely separated, an acutely elevated ridge extending from anterolateral margin of coxa to antero- lateral margin of prothorax (as in Hylurgops). Elytra much as in Hylurgops except basal margins not armed and interstriae 10 ex- tends to declivity. Anterior tibia slender, entirely unarmed on margins, a terminal spine next to tarsal insertion at center of apex, a very small spine on lateral and median apical angles; meso- and metathoracic tibiae similar except obliquely truncate at apex, with a very short, blunt spine on both inner and outer angles (Figure 24). Tarsal segments 1 and 5 each about as long as 2 and 3 combined, 3 broadly bilobed. rvPE-SPECiEs. — Protohylastes annosus Wood, described below. Protohylastes annosus, n. sp. Figures 22-25 This species resembles a very large Pseudohylesinus, but it is distinguished from all previously known Scolytidae by the very different tibiae. Adult. — Sex not determined. Length 9.8 mm, 2.6 times as long as wide, color very dark brown, vestiture pale. Frons convex, epistomal area transversely impressed, a small, median impression at upper level of eyes; surface smooth, shining, punctures moderately coarse, close, their interiors apparently reticu- late; subglabrous, a few scales on lateral margins, a few hairlike setae in epistomal area. Eye oval, about 1.5 times as long as wide. Antenna as described above. Pronotum 0.80 times as long as wide; widest at base, sides arcu- ately converging to a rather strong constriction just behind very broadly rounded anterior margin; basal margin bisinuate; surface smooth and shining on most of discal area, becoming reticulate in all marginal areas and on sides; punctures moderately coarse, close, deep, weakly subcrenulate at base and in lateral areas. Vestiture of rather sparse scales, each scale about six times as long as wide. Elytra 1.7 times as long as wide, 2.3 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather narrow- ly rounded behind; striae weakly impressed, punctures small, close, deep; interstriae twice as wide as striae, surface finely rugose- subreticulate, with numerous small, confused, transverse crenula- tions, each up to one-third width of an interstriae; crenulations near basal margins much as in Hylurgops, but margin without a definite row of crenulations. Declivity moderately steep, occupying posterior third of elytra, broadly convex, with areas at apices of interstriae 5-7 slightly elevated; striae more deeply impressed; interstriae 10 reaching declivity. Vestiture of rather abundant, small scales, June 1973 wood: platypodidae and scolytidae 85 each scale very slightly longer than wide, their color pale except small patches of dark brown on declivity. Type Locality. — Eungella National Park, Queensland, Aus- tralia. Type Material. — The unique holotype was taken at the type locality on lO-XII-65, by G. Monteith. It is in the Queensland Museum. Platypodidae Most platypodids belong to a compact subfamily of highly modi- fied ambrosia beetles. In fundamental structure they differ from scoiytids only in degree, not in the kind of characters represented. They share with primitive scoiytids the same basic head structure, including a large, well-defined pregula (Figures 8, 17-19). The mouthparts in most of them are highly modified to the ambrosial habit, but primitive forms (Coptonotinae) are comparable to those of scoiytids. Virtually identical antennae and pronota are found among primitive forms in the two groups. The protibiae at first appear unique until it is noted that tubercles and ridges appear on the posterior face of the protibiae of some representatives of almost all groups of scolytid woodboring ambrosia beetles. If the minute ridges and tubercles on the posterior face are ignored and the spines and denticles on the apical and lateral margins are emphasized, or if only bark-infesting genera are examined, a graded series of steps apparently bridging the gap between primitive scoiytids and primi- tive platypodids can be demonstrated (Figures 25-33). Since socketed tibial spines characteristic of most scoiytids occur only in that group and not in platypodids, only unsocketed spines are considered primi- tive here. When the posterior face of the prothoracic tibiae are viewed from exactly the same aspect it is noted that the tarsal insertion is visible near the apex on all Platypodidae, including Coptonotinae, on Protohylastes, and on all Scolytini. In Protoplatypus, Mecopelmus, and Protohylastes the tarsus evidently is capable of movement through an arc of almost 180 degrees from the lateral to posterior to mesal positions. In Coptonotus and most Scolytini the action is simi- lar but the arc is somewhat smaller. In some Scolytini the tarsal insertion is closer to or even on the apical margin, and greater free- dom of movement may occur. In Schedlarius the lateral margin of the tarsal insertion is slightly elevated, thus restricting tarsal action more nearly to the posterior to mesal arc; in Platypodinae this arc evidently is even more strongly restricted. In Aricerus, Scolytopla- typus, and many of the higher Scolytinae, the tarsal insertion is on the apical or lateral margin, and tarsal action apparently is through an arc from the lateral to anterior and possibly posterior positions. In most Hylesininae the tarsal insertion is clearly on the anterior face, and tarsal action is restricted to an arc from the lateral to anterior position or less. A submarginal tubercle on the posterior face near the tarsal insertion in primitive forms appears to have special significance; it is absent in Scolytini and Cossoninae. 86 GREAT BASIN NATURALIST Vol. 33, No. 2 Coptonotus 31. Scolytodes 32. ^ 33. Diamerus Camptocerus Figs. 25-33. Posterior face of prothoracic tibiae: 25, Protohylastes annosus Wood, with tarsus. Hylastini; 26, Protoplalypus veiulus Wood, with tarsus, Coptonotinae; 27, Mecopelmus zeteki Blackman, with first segment of tarsus, Coptonotinae; 28. Schedlarius mexicanus Duges, wdth first segment of tarsus, Coptonotinae; 29. Coptonotus cyclops Chapuis, witli first and second tarsal seg- ments, Coptonotinae; 30, Tricolus peltatus Wood, Scolytinae; 31, Scolytodes sp., Scolytinae; 32, Diamerus impar Chapuis. Hylesininae; 33, Camptocerus auri- cnrnus Blandford, Scolytinae. All drawings were made at different scales so as to be reproduced at a uniform size. In platypodids the first tarsal segment is supposed to be as long as the remaining segments combined; in some it is actually less than half as long (Schedl, 1939). Representatives of the Coptonotinae (Coptonotidae of Schedl) have the tibiae and tarsi as in some Platy- podinae (Platypodidae of Schedl) except that there is greater vari- ability, particularly in the length of the first tarsal segment. One of these, Schedlarius^ has wood-boring habits, but the larvae, unlike Platypodinae, form independent mines that wander through deep xylem tissues; fungal activity definitely associated with the beetles alters the character of the wood, although there is no mycelial growth in the tunnels that could be used for food as occurs with ambrosia beetles. Another coptonotid, Mecopelmus Blackman, is phloeopha- gous; a large nuptial chamber is formed by the ]iarcnts in which clusters of eggs are deposited, and larvae then form individual mines that wander in a radiating pattern from this chamber. The genus Protoplalypus described above is anatomically rather closely allied June 1973 wood: platypodidae and scolytidae 87 to Mecopelmus except that the tarsi are typical of scolytids; as noted above, it is phloeophagous and polygamous. In most male platypodids, abdominal tergum 8 is of reduced size and is largely or entirely covered by tergum 7 as in the female. In Schedlarius, male tergum 8 is more nearly intermediate between the reduced state seen in other platypodids and the normal condi- tion of most scolytids and many curculionids. A reduced male ter- gum 8 also occurs in the scolytid tribes Ipini (Ips, etc.) and Carpho- dycticini {Craniodycticus, etc.). It is doubtful that this reduction in these two tribes indicates a close relationship to platypodids. The larvae of platypodids and scolytids are inadequately known, but most of them can be segregated using features of the labrum and clypeus. In platypodids the clypeus is broad and strikingly reduced in length or entirely absent and the labrum is proportionately lengthened. In Schedlarius and Protoplatypus of the Platypodidae and Carphodycticus Wood in Scolytidae, the labrum and clypeus are intermediate in size and shape. In my opinion the platypodids are an aberrant group of am- brosia beetles, as are most other groups of ambrosia beetles within the family Scolytidae, that arose long after scolytid-platypodid char- acters had been well established. The ambrosial habit has arisen repeatedly within the Scolytidae; for example, Camptocerus Erich- son (Scolytini), Scolytoplatypus Schaufuss (Scolytoplatypini), Both- rosternus Eichhoff (Bothrosternini), Hyleops Schedl (Hylesinini), and numerous genera in the more highly evolved tribes Xyleborini, Xyloterini, and Corthylini. For this reason, the appearance of the ambrosial habit very early in scolytid phylogeny is not unusual. Every character on which the family Platypodidae is based, whether anatomical, behavioral, or ecological, intergrades with the Scolytidae. The only justification I see, at present, for retaining it as a family separate from Scolytidae is tradition and the fact that most forms encountered in the field are easily recognized. Scolytidae As conceived here, the family Scolytidae consists of the two sub- families Hylesininae and Scolytinae (including Ipinae). The adult forms of Scolytidae ( 1 ) have a definite pregula and pregular sutures, (2) lack a spherical head and rostrum, (3) have a distinctive man- dibular articulation (quite different from curculionids?), (4) never have a petiolate point of labial articulation, and (5) have tibiae that are adapted to a bark- or wood-boring habit and, in primitive forms, are no more similar to Cossoninae than they are to those of certain Brenthidae. In part, the similarity between Scolytidae and Rhyn- colini is due to convergence or more probably to parallel evolution that commenced when both groups entered part of the same broad ecological niche. The short pseudorostrum of the Hylastini and allied forms of Hylesininae appears to have been acquired independently and differs in fundamental structural detail from that seen in Curculionidae (Figures 1-7). 88 GREAT BASIN NATURALIST Vol. 33, No. 2 The division of the Scoiytidae (including Platypodidae) into three major lines of development occurred very early m its phyletic history but well after the basic characteristics of the group were established. The earliest living genera representing those lines, Pro- toplatypus and possibly Mecopelmus for Platypodidae, Protohylastes, and, doubtfully, certain Cnemonyx (galeritus Eichhoff) for Hyle- sininae, could appropriately be placed as primitive Scolytinae since they are intermediate and lack some of the most diagnostic characters of the groups I presume they represent. The argument for the above classification, and also for the re- tention of Platypodidae and Scoiytidae as families, must be con- sidered tentative and biased. It is based largely on a limited number of external adult characters. The larvae, habits, and internal char- acters of most primitive genera of concern are totally unknown. Collecting experience in tropical areas has demonstrated that speci- mens of these primitive genera are exceedingly rare and suggests that several more equally rare genera await discovery. It should also be mentioned that in a review of this problem careful attention should be given to the sporadic occurrence of a very short antennal scape, broadly bilobed tarsi, scalelike vestiture, gnathal, and other characters commonly found in primitive genera of these groups. The significance of a complete gula, with two totally separate gular sutures found in several species of the neotropical genus Gnatho- trupes (Scoiytidae) (Figure 12), and the possible cercus-like struc- tures and the tenth abdominal segment of larval platypodids, cited above, must also be investigated. Convergent and parallel evolution among wood-boring Curculionoidea must be examined very carefully in order to sift the meaningful from the misleading superfluous char- acters found in these families. Among primitive Coleoptera the tentorial structure arises from two separate tentorial pits on the gular sutures about midway be- tween the foramen magnimi and the oral fossa (Stickney, 1923). From these pits the posterior tentorial bridge arises, forming a clear- ly identifiable internal landmark. The anterior tentorium extends forward from this point for some distance, then branches to form (1) the dorsal arms that extend to but do not fuse with the wall of the head near the anteromesal margin of the eye (connected by liga- ments only) and (2) the anterior arms that extend to the anterior tentorial pits near the anterior articulation of the mandibles. In all Curculionoidea having an anterior tentorium the anterior arms ex- tending to the anterior tentorial pits are entirely absent. The dorsal arms vary from well-develo[)ed to obsolete. The posterior tentorium in Curculionoidea is carried inward on a Y-shaped apodeme having a median basal portion arising from the median gular suture from the posterior tentorial bridge to the postgula; the two arms of this basal piece branch from its inner margin. The anterior extremity of this internal gular structure forms a single median ])illar in all Curculionoidea having a single gular suture and lacking pregidar sutures (Figures 1-4). In all Platypodidae and Scoiytidae this an- terior tentorial structure forms two pillars that continue antero- June 1973 wood: platypodidae and scolytidae 89 laterally along the strongly inflected apodemal ridges formed by the pregular sutures (Figures 5-19). In my opinion this character alone is sufficiently unique and striking, in combination with the external pregular sutures, to warrant family recognition of the group repre- sented by the Platypodidae and Scolytidae. Summary and Conclusions The tibial and other characters used as a basis for reducing the Platypodidae and Scolytidae to subfamilies of Curculionidae (Crow- son, 1967; Kuschel, 1966) were adaptive characters not found in primitive representatives of the group. A definite pregula clearly defined by sutures associated with an internal inflection of the cuticle occurs in Platypodidae and Scoly- tidae but not in any other Curculionoidea having only one gular suture. It is postulated that pregular sutures occur only in those groups in which a rostrum is lacking and has never been developed; therefore, since pregular sutures are absent in all groups with a rostrum, or in which it was secondarily lost, their absence indicates a specialization. One genus of Platypodidae has the adult maxillary palpus 4- segmented. Since no known representative of the Curculionidae (s. str.) has more than a 3-segmented palpus, the ancestral stock from which the Platypodidae were derived must predate that of the Curculionidae. Three of the most primitive genera of Platypodidae for which habits are known are either phloeophagous or xylophagous and are not associated with an ambrosial fungus. Since the ambrosial habit has arisen independently many times in the Scolytidae on almost every major phyletic line, it is postulated that the platypodids arose very early from the Scolytidae and now constitute an aberrant group within or very near that family. Tibial, tarsal, and gnathal char- acters in primitive genera also tend to intergrade in the two groups. Therefore, three major, equally distinctive phyletic lines of develop- ment are recognized in Platypodidae (including Coptonotidae), Hylesininae, and Scolytinae (including Ipinae). Platypodidae is tentatively retained as a family separate from Scolytidae for reasons of tradition until exhaustive studies clearly indicate the need for a change. References Browne, F. G. 1972. Larvae of the principal old world genera of the Platy- podinae (Coleoptera: Platypodidae). Trans. Roy. Ent. Soc. London. 124:167- 190. Crowson, R. a. 1967. The natural classification of the families of Coleoptera. Classey: Hampton, England. Reprinted from Entomologists Monthly Maga- zine, 1950-1954, and from Lloyd, London, 1955. Hopkins, A. D. 1909. The genus Dendroctonus. U.S. Dept. Agric, Bur. Ent. Tech. Bull. 17(1):1-164, 8 pi. Kuschel, G. 1966. A cossonine genus with bark-beetle habits, with remarks on relationships and biogeography (Coleoptera Curculionidae). New Zealand .1. Sci. 9(l):3-29. 90 GREAT BASIN NATURALIST Vol. 33, No. 2 ScHEDL, K. E. 1939. Die Einteilung und geo^aphische Verbreitung der Platy- podidae. Proc. VII Internat. Cong. Ent., Berlin, p. 377-410. . 1972. Monographie der Faniilie Platvpodidae Coleoptera. Junk, Den Haag. 322 p. Stickney, F. S. 1923. The head-capsule of Coleoptera. Illinois Biol. Monogr. 8(1):1-105. STUDIES ON UTAH STONEFLIES (PLECOPTERA) Richard W. Baumann^ Abstract. — Questionable Utah distribution records are reviewed and new state records are given. The female of Capnia cygna Jewett is described and figured and an allotype 9 designated. Detailed synonomies and nomenclatural notes are provided for Capnia vernalis (Newport), Utacapnia poda (Nebeker and Gaufin) and Oemopteryx fosketti (Ricker). Descriptions are given of the female, nymph, and egg of Isogenoides zionensis Hanson and an allotype 5 designated. "Hie male, nymph, and egg of Pictetiella expansa (Banks) are described and an allotype 5 designated. Sweltsa gaufini, sp. nov., is described for the male, female, and egg stages and a holotype $ designated. Descriptions are either supported with original figures or reference is made to acceptable figures in the literature. Besides line drawings and halftones, useful photographic figures are provided which were prepared by using a scanning electron microscope. An annotated list of Utah species is given following the revised nomenclature of lilies (1966) and Zwick (1973). The publication of a monograph on the stoneflies of Utah (Gaufin, Nebeker, and Sessions, 1966) contributed greatly to the knowledge of intermountain Plecoptera. Since this time, however, additions and corrections have been discovered. This study contains this information and includes an annotated list of Utah species. Detailed synonymies are given for recent nomenclatural changes and where special clarification is necessary. The listing of type specimens and their depositories indicates that they were studied. For further information and complete taxonomic treatment, see lilies (1966) and Zwick (1973). Malenka flexura (Claassen) NemouTa flexura Claassen (1923:284). Boulder, Colorado. This species was recorded by Gaufin et al. (1966) as occurring in Utah. The record was based on one female in the collection of Dr. William E. Ricker, which was collected at Huntsville, Weber Co. The specimen was examined as part of this study and found to belong to Malenka californica (Claassen). This does not preclude the possibility that M. flexura might be found in the future, but no confirmed records are presently available. Podmosta decepta (Prison) Nemoura decepta Prison (1942:13). Estes Park, Colorado. This species is found in most of the Intermountain states. It was first collected in Utah at the following locality in the Uinta Moun- tains: small creek 7 miles N Mirror Lake, Hwv. 150, Simimit Co., 21-VII-1967, R. W. Baumann, 1 d (NMNH). 'Department of Entomology, Smitlisonian Inslltiilion. Wnsliington. D.C. 20560. 91 92 GREAT BASIN NATURALIST Vol. 33, No. 2 Podmosta delicatula (Claassen) Nemoura delicatula Claassen (1923:285). Boulder, Colorado. Specimens of this species were collected by the author during a survey of the stoneflies of the Wasatch Mountains. Since the species was fairly common in small creeks at high altitudes, a closer study was made of the collection at the University of Utah. This resulted in the discovery of other Utah specimens previously included with Prostoia besametsa (Ricker). The confirmed records are as follows: Davis Co., Farmington Canyon Creek, 23-VIII-1965, R. W. Bau- mann, 8 9? (UU). Duchesne Co., Mirror Lake, 12-VII-1947, L. T. Nielsen, 5 5 9 (UU). Rich Co., Allen Canyon, 19-VI-1972, G. F. Knowlton, 1 9 (NMNH). Salt Lake Co., Big Cottonwood Creek, Brighton, 16-Vn-1952, A. R. Gaufin, 6 d^ c?, 3 9 9 ; 16-VIII- 1962, 1 d'; 6-Vm-1965, R. W. Baumann, 1 9 ; 14-VII-1966, 1 9 ; 29-VII-1967, 1 d (UU) (NMNH). Summit Co., Beaver Creek, Beaver Creek Campground, 1 -VIM 959, A. R. Gaufin, 11 cTc?, 29 9 9 (NMNH); Hoop Lake, 30-VM958, D. B. Cahill, 1 9 (USU); Provo River, below Trial Lake, 5-Vin-1947, A. R. Gaufin, 1 9 ; 6- VIII-1962, 1 cT (UU); Smith Morehouse Creek, South Fork Guard Station, 19-Vn-1961, A. R. Gaufin, 4 cf cf, 3 9 9 (UU); Weber River, junction Smith-Morehouse Creek, 30-VI-1959, A. R. Gaufin, 12 cT cf, 29 9 9 (NMNH). Wasatch Co., Provo River, Soapstone, 21-Vn-1947, A. R. Gaufin, 2 9 9; 6-Vn-1959, 9 cfc^, 15 9 9 (UU) (NMNH). Zapada frigida (Claassen) Nemoura frigida Claassen (1923:285). Sitka, Alaska. Zapada frigida is never abundant but has a scattered distribution throughout most of the western states. A single male from the Manti- La Sal National Forest is the first Utah record: Johnson Creek, 19 miles N Blanding, Abajo Mountains, San Juan Co., 18-VI-1946, S. B. Muliak (NMNH). Zapada oregoncnsis (Claassen) Nemoura oregonensis Claassen (1923:288). Blitzen Valley, Harney Co., Oregon. This species was recorded from Utah by Ricker (1952), Gaufin (1955, 1964), Gaufin et al. (1966), and Baumann and Gaufin (1971). The author examined all available specimens of Z. ore- gonensis while studying the Rocky Mountain Nemouridae and found them all to bo ZMpada haysi (Ricker). The species has been coiifirmed from Idaho, Wyoming, and Colorado, so it is possibly also present in Utah. Capnia cygna Jewett Capnia cygna Jewett (1954:546). Washington (?). The type locality of C. cygna is believed to be Washington. Nebe- ker and Gaufin (1966a) recorded it from Idaho and gave a brief June 1973 baumann: utah plecoptera 93 description and drawing of a female dissected from a mature nymph. Mature females have since been collected, including a single speci- men from Mueller Park, Davis Co., Utah, 25-11-1949, R. B. Selander, which is here designated as the allotype $ (NMNH). A detailed description and new drawing are included because of the teneral condition of the specimen studied earlier. Female. — Macropterous. Length of forewings 8-10 mm; length of body 8-10 mm. Body and appendages dark brown almost black, broad membranous dorsal stripe on abdominal tergites 1-8. Wings hyaline; venation typical of genus, with 1-3 crossveins between Ri and R2 beyond cord. Eighth stemite with subgenital plate quite simple, heavier sclerotization on both median margins, posterior median margin produced but not extending beyond distal margin of segment, produced portion broadly rounded or slightly angular (Figure 2). Capnia elongata Claassen Capnia elongata Claassen (1924:56). Caribou, Plumas Co., California. The confirmed distribution of this species is along the Pacific Coast, so it probably does not occur in Utah. The specimens that contributed to the Utah records by Knowlton and Harmston (1938), Gaufin (1955), and Gaufin et al. (1966) were found to be Capnin gracilaria Claassen. Capnia vernalis (Newport) ^ Capnia vernalis Newport (1851:451). Lectotype, male; Albany River, Ontario. Canada (BMNH). Capnia limata Prison (1944:155). Holotype, male; South Platte River, Littleton, Colorado (INHS). Capnia vernalis: Zwick (1973: ?). The name Capnia limata has been given to specimens from the western United States, while the name Capnia vernalis is used in Canada and the northeastern United States. After I examined both type specimens, it was apparent that a single, widely distributed species was involved. This synonymy is recorded in Zwick (1973). Isocapnia hyalita Ricker Isocapnia hyalita Ricker (1959:648). Hyalite Creek, Gallatin Co., Montana. The discovery of this species in Utah is a large range extension, since all previous records are from Montana. It appears to be quite common if sought in the correct habitat at the right time. All records to date are from mountain streams during the months of April and May. The new Utah records are: Utah Co., American Fork Creek, near Timpanogos Cave National Monument, 8-IV-1967, C. D. Bjork and B. R. Oblad, 2 long- winged d"c?, 1 9; lO-IV-1967, 2 long- winged d'cf, 2 short-winged d'cf, 2 9 9; 12-IV-1967, 1 short- winged d", 1 9 (NMNH). Tooele Co., South Willow Creek, Upper 94 GREAT BASIN NATURALIST Vol. 33, No. 2 Figs. 1, 3-5, 8. Sweltsa gaufini, n. sp.: 1, male abdomen, dorsal view; 3, female terminalia, ventral view; 4, epiproct, a. lateral, b. dorsal, c. ventral; 5, leaflike structure on aedeagus; 8, head and pronotum. Fig. 2. Capnia cygna Jewett: Female tentninalia, ventral view. Fig. 6. Sweltsa albertensis (Needham and Claassen): Leaflike structure on aedeagus. Fig. 7. Sweltsa lamba (Needham and Claassen): Leaflike structure on aedeagus. Fig. 9. Pictetiella expansa (Banks): Right wings of male. June 1973 baumann: utah plecoptera 95 Narrows, 15-V-1967, R. W. Baumann and B. R. Oblad, 5 long- winged cf cT , 6 short- winged dd,^ ? 9 (NMNH). Utacapnia poda (Nebeker and Gaufin) Capnia poda Nebeker and Gaufin (1965:475). Holotype, male; Gunnison River, Colorado (UU). Capnia (Utacapnia) poda Nebeker and Gaufin (1967:226). Utacapnia poda: Zwick (1973:?). This species is widely distributed in Colorado, and the collection of a specimen in Utah near the border is not too surprising: Green River, Jensen, Uintah Co., 24-11-1968, R. W. Baumann, 1 9 (NMNH). Oemopteryx fosketti (Ricker) Brachyptera fosketti Ricker (1965:475). Holotype, male; South Saskatchewan River, Clarksboro, Saskatchewan, Canada (CNC). Brachyptera zelona Ricker (1965:477). Holotype, male; My ton, Utah (INHS). Oemopteryx fosketti: Zwick (1973:?). Oemopteryx zelona was described from one male collected at the Duchesne River in Myton, Utah, which Ricker (1965) erroneously listed as the Green River. Since the holotype was a single male, a special effort was made to collect additional specimens. The exami- nation of several males from Utah led to the synonymy of O. zelona under O. fosketti included in Zwick (1973). The following records have become available since the original description: Duchesne Co., Duchesne River, Myton, 24-11-1968, R. W. Baumann, 1 ? (NMNH) ; Duchesne River, near Randlett, 24-11-1968, R. W. Baumann, 1 cf (NMNH). Uintah Co., Uinta River, Hwy. 40, near Roosevelt, 5-III-1963, A. R. Gaufin, 1 $ (UU); Green River, Dinosaur Na- tional Monument, 25-III-1967, S. L. Jensen, 3 $ 2 (NMNH); Green River, Jensen, 24-11-1968, R. W. Baumann, 9 d'cf, 48 9 9 (NMNH); Green River, Ouray, 24-11-1968, R. W. Baumann and G. Z. Jacobi, 9 d^c^,8 9 9 (NMNH). Isogenoides zionensis Hanson Isogenoides zionensis Hanson (1949:109). Holotype, male; Zion National Park, Utah (UMA). Female. — Macropterous. Length of forewings 21-23 mm; length of body 20-22 mm. Body dark brown dorsally, yellow ven- trally; legs brown. Pronotum brown, with broad yellow median stripe. Subgenital plate large, extending 1/3 length of eighth sternite, posterior margin mostly straight, sometimes with small, rounded median projection, lateral corners slightly rounded, forming nearly right angles. Nymph. — Length of mature d" 22-25 mm; mature 9 26-29 mm; antennae 7-9 mm; cerci 10-12 mm; with single submental gills 6-7 times as long as wide. Body and legs light brown; occipital ridge 96 GREAT BASIN NATURALIST Vol. 33, No. 2 Fig. 10. Isogenoides zionensis (Hanson): Mature nymph. June 1973 baumann: utah plecoptera 97 heavy on lateral margins, becoming somewhat thinner towards median line; thin rows of long, light hairs along middorsal line of thorax and abdomen; faint color pattern with small setae on dorsal surface of head and thorax (Figure 10). Mouthparts similar to /. elongatus (Hagen). Labium typical in shape for genus, para- glossae covered with long, dense hairs. Maxillae elongate; lacinia with two short teeth, one apical and one subapical, inner margin with sparse row of thin spinules; galea long and narrow; tip rounded and bearing tuft of short spines on ventral half (Figure 130), ex- tending to base of subapical tooth; palpus five-segmented (Figure 13P). Mandibles rather similar, with two apical and one subapical tooth on each cusp, ventral basal margin of outer cusp with row of stout denticles, narrow patch of fine hairs extending from denticles to base of mandible (Figure 14Q,R), basal margin of inner cusp with dense tuft of long, thin hairs, dorsal surface with rectangular patch of hairs at base of teeth (Figure 14S,T) . Egg. — Average width 350 /(,, average length 550 /x. Oval and somewhat elongate, triangular in cross-section, sides equilateral. Collar located at one end, round in shape, with three stabilizing ribs, one at each angle, with broadly rounded, membranous anchor plate when fresh. Microstructure of surface composed of numerous round- ed knobs of almost equal size. Micropiles situated at midline in rows of 4-6 running perpendicular to long axis, present on all three sides (Figure 12A,B,C,D). Isogenoides zionensis was described from five males collected at the Virgin River in Zion National Park. Gaufin et al. (1966) gave figures of a cT abdominal tip, 5 subgenital plate, and nymphal mouth parts but did not include descriptions. They also gave excel- lent figures of the male genitalia, which were mislabeled as being those of Isogenus elongatus. Their figure of the mandible fails to show the denticles on the basal margin of the outer cusp. The labium is shown as bare without the conspicuous covering of dense hairs. The egg is figured in Knight et al. (1965) . The nymph is almost indistinguishable from /. elongatus (Hagen) and keys out as such in Ricker (1952), the only known difference being the very light brown almost yellow color of /. zionensis as compared to the darker brown of /. elongatus. Nymphs are very common throughout the Virgin River drainage of southwestern Utah, but few adults have been collected. The author, after many attempts, was finally able to obtain 2 cf c^ and 4 ? 9 from a culvert at the following locality: East Fork Virgin River, Hwy. 89, near Glendale, Kane Co., 23-V-1970, R. W. Baumann (UU) (NMNH). One of the females is here designated as the allotype 9 (NMNH) . Pictetiella expansa (Banks) Perla expansa Banks (1920:317). Holotype, female; Grant, Colorado (MCZ), 10,816. Perla expansa: Needham and Claassen (1925:81, 313, 325) description of $ and 9; figures of wings. $ and 9 genitalia {$ incorrectly associated). 98 GREAT BASIN NATURALIST Vol. 33, No. 2 Fig. 11. Pictetiella expansa (Banks): Mature nymph. June 1973 baumann: utah plecoptera 99 Perla expansa: Claassen (1931:55) description of nymph (incorrect nymphal association) . Isogenus (Pictetia) expansus: Ricker (1952:120-122) description of $ and nymph; figures of nymphal maxilla and $ genitalia. Pictetiella expansella: lilies (1966:375) (incorrect specific name). Isogenus {Pictetia) expansus: Gaufin, Nebeker, and Sessions (1966:62, 64) figures of $ and 9 genitalia. Male. — Slightly brachypterous. Length of forewings 9-12 mm; length of body 13-16 mm. Body dark brown, last two abdominal segments yellow dorsally; legs yellowish brown; antennae brown; cerci yellow. Head as broad as prothorax, ocellar triangle equilateral, posterior ocelli slightly closer to eyes than to each other, anterior area light yellow, dark brown U-shaped bands connecting lateral ocelli to anterior ocellus, bands extending forward beyond anterior ocellus and laterally from lateral ocelli to form rocking "H," pos- terior median area yellow, lateral posterior corners brown, palpi brown. Pronotum slightly wider than long, brown with broad yel- low median stripe, rugosities rather coarse and restricted to inner %, marginal groove present only at anterior and posterior margins, forming broad anterior and posterior bands. Wings hyaline, with small infuscated area near cord, veins brown, venation slightly aber- rant (Figure 9). Abdominal segments normal to ninth, which is narrowed dorsally and greatly extended ventrally at apical margin, ventral extended margin broadly rounded and clothed with fine brown hairs; tenth tergite nearly bisected by deep, sclerotized groove which shields epiproct when not extended. Hemitergites set off from remainder of tergite by angular membranous areas, anterior half of hemitergites covered with long, fine hairs. Epiproct long and narrow with pointed apex, mostly membranous with thin, sclerotized areas visible on dorsal and ventral median areas, patches of small setae on dorsolateral margins. Nymph. — Length of mature cT 15-16 mm; mature 9 17-21 mm; antennae 8-9 mm; cerci 7-8 mm; with single submental gills 3-4 times as long as wide. General color brown, but quite strikingly patterned; legs mostly yellow; antennae slightly darker at apex; apical half of cerci very dark brown almost black. Head as wide as pronotum; maxillae barely visible from above; hind ocelli closer to eyes than to each other; occipital ridge as few sparse hairs near lateral margins. Thin rows of hairs running along middorsal line of thorax and abdomen. Head pattern light on dark background: bulb- shaped structure in ocellar triangle, M-line, front of head, lateral tubercles and elongate spot near each eye. Pronotum transversely oval, almost twice as wide as long, outer borders with narrow, dark band fringed with long setae, marginal groove complete but dark along anterior and posterior margins, with distinct light reticulations on dark background. Wing pads on mesonotum and metanotum well developed, outer margins running slightly convergent to body axis, with distinct patches of dark spines along lateral anterior borders, light reticulate markings medially. Abdominal segments fringed with row of dark hairs on posterior margins; faint, narrow, dark 100 GREAT BASIN NATURALIST Vol. 33, No. 2 Fig. 12. Eggs. AD, Isogenoides zionensis Hanson: A, collar, end view (240X); B, lateral view (200X); C, micropiles (620X); D, micropile, closeup (6,000X). E-H, Picteticlla expansa (Banks): E. collar, end view (270X); F. dorsal view (195X); G, ventral view (180X); H, collar opening, closeup (385X). June 1973 baumann: utah plecoptera 101 bands on anterior margins which are interrupted medially and al- most fade out on terminal segments. Cerci with fringes of short hairs at apex of each segment, fringe of long hairs running dorsally throughout entire length (Figure 11). Mouthparts quite distinctive. Labium typical of family Perlodidae, paraglossae covered with long dense hairs. Maxillae elongate; lacinia with two long, narrow teeth, one apical and one subapical, inner margin with few long thin spinules; galea short but elongate, tip pointed and bearing tuft of short spines on ventral half (Figure 13L), extending to base of sub- apical tooth, palpus five-segmented (Figure 13M,N). Mandibles rather similar, with one subapical and two apical teeth on each cusp, middle teeth with serrated margins, ventral basal margins of outer cusp smooth and bare except for large patch of fine hairs at base of cusp (Figure 14U,V), basal margin of inner cusp with dense tuft of long, thin hairs, dorsal surface with rounded patch of hairs at base of teeth (Figure 14W,X). Egg. — Average width 300 /x, average length 550 /i. Dorsal aspect nearly oval, axillary lateral ridges extending from median ridge at broadly rounded end to form skewed, cross-like structure, collar end enlarged and rounded. Triangular in cross-section, almost equi- lateral, but ventral side longest. Collar modified as flat, hood-like structure, convex dorsally, nearly flat ventrally with large opening, anchor plate large, flat, and covering entire ventral surface. Micro- structure of surface composed of numerous rounded knobs of almost equal size, knobs arranged in loose, almost circular, designs. Micro- piles situated near midline in rows of 5-7 running perpendicular to long axis, those on ventral surface not as obvious. Surface of un- cleaned egg covered with small puffball-like structures which, ac- cording to Brinck (1949), are special adhesive bodies (Figure 12E,F, G,H). Pictetiella expansa was named from Colorado from a single female. Since the time of the original description, much confusion has existed as to the identity of this species in both the adult and nymphal stages. Most specimens previously assigned to this species were examined as part of this study, and four genera were repre- sented. This indicated a need for the detailed descriptions of the male, nymph, and egg which are included here. Kicker's (1952) description of a nymph and of teneral male genitalia seem to be correctly associated but are rather general. The figures of the male and female genitalia by Nebeker (Gaufin et al., 1966) were done from specimens used in this study and are excellent. Saether (1970) figures a nymphal maxilla labeled Isogenus (Pictetia) expansiis which does not appear to belong in the genus Pictetiella. The genus Pictetiella was monotypic and recorded as nearctic until the recent description of Pictetiella asiatica from Siberia by Zwick et al. (1972). Records of P. expansa have been confirmed from Colorado, Montana, Utah, and Wyoming. The Utah localities are: Salt Lake Co., Parley's Creek above Mountain Dell Reservoir, 5~IV-1963, A. V. Nebeker, nymph (UU); Lambs Canyon Creek, 102 GREAT BASIN NATURALIST Vol. 33, No. 2 19-VI-1966, R. W. Baumann, nymphs; 18-VIII-1966, exuviae (UU); Mill Creek, Aug. and Sept. 1965, R. W. Baumann, 12 d' cT, 15 9 $ , nymphs; July and Aug. 1966, 7