HARVARD UNIVERSITY Library of the Museum of Comparative Zoology Brigham Young University Science Bulletin - Biological Series V. 10-12 (1968-71) 10:1. TICKS OF THE NATIONAL REACTOR TESTING STATION. Dorald M. Allred 10:2 NEW RECORDS AND SPECIES OF NEOTROPICAL BARK BEETLES (SCOLYTIDAE: COLEOPTERA). PART IV. Stephen L. Wood 10:3. A STUDY OF THE WEEVIL TRIBE CELEUTHETINI OF THE SOLOMON ISLANDS (COLEOPTERA: CURCULIONIDAE) . Vasco M. Tanner 10: A. NESTING ECOLOGY OF RAPTORIAL BIRDS IN CENTRAL UTAH. J.R. Murphy, F.J. Camenzind, D.G. Smith, J.B. Weston 11:1. OSTEOLOGICAL AND MYLOGICAL COMPARISONS OF THE HEAD AND THORAX REGIONS OF CNEMIDOPHORUS TIGRIS SEPTENTRIONALIS BURGER AND AMEIVA UNDULATA PARVA BARBOUR AND NOBLE (FAMILY TEIIDAE). D.L. Fisher and W.W. Tanner 11:2. PINYON-JUMPER SUCCESSION AFTER NATURAL FIRES ON RESIDUAL SOILS OF MESA VERDE, COLORADO. James A. Erdman 11:3. DISTRIBUTION OF THE NATIVE TREES OF UTAH. Kimball S. Erdman 11:4. FLORA OF THE NATIONAL REACTOR TESTING STATION. N. Duane At wood 12:1. MITES AND LICE OF THE NATIONAL REACTOR TESTING STATION. Dorald M. Allred 12:2. A COMPARATIVE STUDY OF THE HEAD AND THORACIC OSTEOLOGY AND MYOLOGY OF THE SKINKS EUMECES GILBERTI VAN DENBURGH AND EUMECES SKILTONIANUS (BAIRD AND GIRARD) . D.F. Nash and W.W. Tanner 12:3. EVOLUTION OF THE IGUANINE LIZARDS (SAURIA, IGUANIDAE) AS DETERMINED BY OSTEOLOGICAL AND MYOLOGICAL CHARACTERS. D.F. Avery and W.W. Tanner 12:4. LAND USE IN WAH WAH AND PINE VALLEYS, WESTERN UTAH. J.R. Murdock and S.L. Welsh •5 «) & 1 n , 0 Brigham Young University Science Bulletin TICKS OF THE NATIONAL REACTOR TESTING STATION by DORALD M. ALLRED BIOLOGICAL SERIES — VOLUME X, NUMBER 1 JUNE 1968 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Acting Editor: Wilmer W. Tanner, Department of Zoology and Entomology, Brigham Young University, Provo, Utah Associate Editor: Earl M. Christensen, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: Ferron L. Andersen, Zoology Joseph R. Murdock, Botany J. V. Beck, Bacteriology Wilmer W. Tanner, Zoology, Chairman of tlie Board Stanley L. Welsh, Botany Robert W. Gardner, Animal Science Ex officio Members: Rudger H. Walker, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Director, University Press The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from University Press, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for hbrary exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utali 84601. Brigham Young University Science Bulletin TICKS OF THE NATIONAL REACTOR TESTING STATION by DORALD M. ALLRED BIOLOGICAL SERIES — VOLUME X, NUMBER 1 JUNE 1968 TABLE OF CONTENTS Page INTRODUCTION 1 PHYSICAL CHARACTERISTICS 1 STUDY AREAS 9 PROCEDURES 18 Vertebrate Collections 18 Invertebrate Collections 20 VERTEBRATES EXAMINED 20 Amphibians 20 Reptiles 20 Birds 21 Mammals 22 TICKS 23 Dermacentor andersoni 23 Haemaphysalis leporispalustris 25 Ixodes kingi 25 Other Species 26 DISCUSSION 27 Trapping Efficiency 27 Interaction of Tick Species 27 Ecological Distribution of Ticks 28 Radiation Effects 28 SUMMARY 29 REFERENCES 29 LIST OF FIGURES Figure Page 1. Location of the National Reactor Testing Station in relationship to some cities in southeastern Idaho ... 2 2. Principal study areas at the National Reactor Testing Station 3 3. Lemhi Mountains at the northwestern edge of the National Reactor Testing Station 4 4. Big Lost River during time of spring run-off 4 5. Sinks area in the nortliwestem part of the station 5 6. Mouth of volcanic cave, study area 19 5 7. Mouth of volcanic cave, study area 21 6 8. Inside of volcanic cave, study area 21. Height of ceiHng at this point is approximately 12 ft 6 9. Inside of volcanic cave, study area 33. Height of ceiUng at this point is approximately 7 ft. Mounds at front and back center, and right foreground are piles of woodrat excreta, each several feet deep 7 10. Major plant types as designated by McBride. Reproduced by permission of Ray McBride, Atomic Energy Comm., Idaho Falls 8 11. Study area 1. Predominant plants are Chnjsothamnus and Artemisia 9 12. Study area 2, with twin buttes in the background. Predominant plant is Artemisia 10 13. Study area 3, with twin buttes in the background. Predominant plant is Elymus 11 14. Study area 4. Predominant plants are Ortjzopsis and Stipa 11 15. Study area 5. Predominant plant is Juniperus 12 16. Study area 6, with Lemhi Mtns. in the background. Predominant plants are Chrysothamnus, Tetradymia, and Artemisia 12 17. Study area 7. Predominant plants are Chrysothamnus and Artemisia 14 18. Study area 8, with Lemhi Mtns. in tjie background. Predominant plants are Artemusia and Atriplex 14 19. Study area 9, with twin buttes in the background. Predominant plants are Chenopodium and Eurotia .... 15 20. Study area 10. Predominant plant is Artemisia 15 21. Study area 11. Predominant plants are Chrysothamnus, grasses, and Tetradymia 16 22. Study area 12. Predominant plants are Juniperus, Chrysothamnus, Eurotia, and Artemisia 16 23. Sand dunes (study area 18) with predominant fringing plant, Salsola 17 24. YAW live-catch trap 18 25. Modified Hubbard live-catch trap 18 26. Components of a can pit-trap 19 27. Museum-Special break-back trap 19 28. Oneida-Victor carnivore trap 19 29. Trap arrangement used in the major study areas. P, metal post; C, can pit-traps; S, Museum- Special traps; H, Hubbard traps; Y, YAW traps 19 30. Berlese funnel 20 LIST OF TABLES Page 1. Total number of animals examined in each study area 23 2. Total numbers of ticks collected by species and stage of development 23 3. Hosts, degree of infestation, and general seasonal occurrence for three developmental stages of Dermacentor andcrsoni 24 4. Hosts, degree of infestation, and general seasonal occurrence for three developmental stages of H aemaphysalis leporispalustris 25 5. Hosts, degree of infestation, and general seasonal occurrence for three developmental stages of Ixodes kingi 26 6. Interaction relative to occurrence of Dermacentor andersoni and Ixodes kingi on common hosts 27 7. Frequency index of occurrence of Dermacentor and Ixodes in selected study areas 28 8. Frequency index of ticks in representative plant community types and areas 28 9. Relative infestation and abundance of ticks in irradiated and non-irradiated control areas 28 TICKS OF THE NATIONAL REACTOR TESTING STATION' by Dorald M. Allred INTRODUCTION Ecological investigations by Brigham Young University were begun at the National Reactor Testing Station in June, 1966, and were continu- ous until September, 1967. Although emphasis was on the study of ectoparasites, other related studies conducted as time permitted yielded ad- ditional data on the ecology of the vertebrate hosts and some of the free-living arthropods. These studies are part of an inventory and eco- logical survey of the native animals which occur on and adjacent to nuclear testing stations. These investigations, those begun at the Nevada Test Site by BYU in 1959 (Allred, Beck, and Jorgen- sen, 1963), and those being carried on by other institutions will provide a basis for evaluation of the effects of nuclear testing on native ani- mals relative to the economy and welfare of man. I am indebted to my former associate, D Elden Beck, who contributed a great deal be- fore his untimely death in August, 1967. The field studies were efficiently performed by Clyde L. Pritchett, formerly of Ricks College and now a faculty member of the Department of Zoology and Entomology, BYU; Robert L. Amoureux, formerly a biology teacher in the Payette and Pocatello public schools; Gerald Richards, formerly a graduate student at BYU, now at Wisconsin State University, Whitewater; Eugene McKennie, formerly a graduate student at BYU, now with the Westinghouse Corpora- tion, Idaho Falls; and Duane Atwood, graduate student in the Botany Department, BYU. C. Lynn Hayward, Department of Zoology and En- tomology, BYU, assisted in field studies of the birds, and identified reference collections of the birds and mammals. Wilmer W. Tanner, De- partment of Zoology and Entomology, BYU, verified the identifications of the amphibians and reptiles. Joseph R. Murdock, Department of Botany, BYU, identified the plant types in our major study areas. The ticks were identified by Jose Merino, University of Wisconsin, Milwaukee, and Rich- ard Ashley, Department of Zoology and Ento- mology, BYU. Some identifications and verifi- cations were made by Carleton Clifford, Rocky Mountain Laboratory, Hamilton, Montana. Financial support for the project was pro- vided by the U. S. Atomic Energy Commission, Contract AT( 11-1) -1559. Logistics were ar- ranged by personnel of the AEC operations of- fice in Idaho Falls and at the National Reactor Testing Station, specifically Courtney D. Scott, Preston Brimhall, and Ralph Batie. I am indebted to many laboratory tech- nicians, particularly Jane Ashley, Sylvia Child, and Connie Donaldson, for preparation of the ectoparasites for identification. PHYSICAL CHARACTERISTICS The National Reactor Testing Station (NRTS) is situated in southeastern Idaho, with its southeastern boundary approximately 30 miles west of Idaho Falls ( Fig. 1 ) . Most of the station lies in Butte County, with extensions into Bingham, Bonneville, Jefferson, and Clark Coun- ties (Fig. 2). It covers an area of approximately 'BYU-AEC Report No. COO-1559-r 894 square miles, and extends for about 39 miles north and south and 36 miles from east to west in its longest dimensions ( U. S. AEC, 1965). The station is situated on a level plain with an average elevation of 486.5 ft. This is part of the Snake River Plain section of the Columbia Plateaus province. The station is bordered on Brigham Young Univehsity Science Bulletin j DUBOIS BLACKFOOT Fig. 1. Location of the National Reactor Testing Station in relationship to some cities in southeastern Idaho. Ticks of Reactor Testing Station LEGEND Fig. 2. Principal study areas at the National Reactor Testing Station. Brigham Young University Science Bulletin '^""•^^.itt^ %, ■5* Fig. 3. Lemhi Mountains at the northwestern edge of the National Reactor Testing Station, Fig. 4. Big Lost River during time of spring run-off. Ticks of Reactor Testing Station ';*T/tv-^/*;-u;,v 'Hi^^^SC Fig. 5. Sinks area in the northwestern part of the station. Fig. 6. Mouth of volcanic cave, study aJ-ea 19. Brigham Young University Science Bulletin ^^.if^ ut ^"JiT ^,J.-.d.^.=r ^i'*. <'r\r i-^M^ '^■,2\.-' J' Fig. 7. Mouth of volcanic cave, study area 21. Fig. 8. Inside of volcanic cave, study area 21. Height of ceiling at this point is approximately 12 ft. Ticks of Reactor Testing Station the north and west by the Bitterroot, Lemhi, and Lost River Mountains ( Fig. 3 ) . On the east and south the topography is typical of the plainHke area of the Snake River drainage and the Great Basin. Little Lost River, Birch and Camas Creeks barely reach and drain into the northern areas of the station. Big Lost River enters from the southwest, traverses more than half the north-south distance, and drains into the sinks of the north-central portion (Figs. 4 and 5). The central and southern parts of the station are typified by basalt flows which are exposed or covered by only a few feet of soil. The north- ern section is primarily lake and eolian deposits, and basalt flows are less common. The basalt flows are from earliest Pleistocene to Recent, some dating perhaps as recent as 2000 years (Bates, 1965). Rising above the basalt flows at the southern part of the station are three promi- nent buttes towering 1000 ft or more above the landscape — East, Middle, and Big Southern Buttes. Typical of many volcanic areas, the basalt flows are honeycombed with caves of various sizes and degrees of accessibility (Figs. 6 to 9). Annual precipitation at the station averages less than 10 inches. The average annual tem- perature is 42 degrees, with extremes of 102 and minus 43 degrees F (U. S. AEC, 1965). Prevail- ing breezes are typical of the area. The vegetation of the NRTS is characteristic of the cool, northern desert shrub type biome. The most conspicuous plant over most of the area is sagebrush, Artemisia tridej^tata. Other predominant plants are rabbitbrush, Chryso- thamnus sp., and grasses of several genera — Agropyron, Elymus, Oryzopsis, and Stipa. Mc- Bride prepared a vegetation map of the NRTS on which he designated 19 major plant communi- ties (Fig. 10). Detailed plant analyses of our permanently established study areas vary some- what from McBride's designations; however, ref- erence is made to his types for correlation with his map. '■^rc-'fi Fig. 9. Inside of volcanic cave, study area 33. Height of ceiling at this point is approximately 7 ft. Mounds at front and back center, and right foreground are piles of woodrat excreta, each several feet deep. Brigham Young University Science Bulletin NRTS VEGETATION TYPE MAP LEGEND NRTS BOUNDABV HIGHWAY INTERMITTENT STREAMS INSTALLATIONS TOWNSHIPS RANGE LINES BIG SAGEBBUSH-BLUeeUNCH WHEATGRASS-RABB I TBRUSH Bl G SAGEBRUSH-RABBI TBRUSH- FOXTA I L BIG SAGEBRUSH-THICKSPI KE WHEATGRASS- NEEOLEGRASS e IG SAGEBRUSH- WIN TERF A T-R ABB J TBRUSH BIG SAGEBRUSH-WINTERFAT-SALTSUSH BIG SAGEBRUSH-SHADSCALE-RABBITBRUSH Bl G SAGEBRUSH-RICEGHASS-NEEDLEGRASS BLACK SAGEBRUSH-BIG SAGEBRUSH- SHAOSCALE I SMALL SAGESHUSH-SHAOSCALE-FOXTAI L CRESTED WHEATGRASS (SEEDED) BLUEBUNCH WHEaTGRASS-TKREETIP S AGE - RABB I TBHU SH LUESTEM WHEATGRASS-WIREGRASS- IVA Rl CEGRASS NEEDLEGRASS- RABB I TBRUSH- CACTUS CANADA WILD R YE- RABBI T BRUSH-B I G SAGEBRUSH ' JUNIPER-8IG SAGEBRUSH-BLUEBUNCH WHEATGRASS H0BSEBBUSH-RAB8IT BRUSH-BIG SAGEBRUSH HABBITBRUSH-8IG SAGEBRUSH-GRASS SALT9USH-WINTERFAT- INDIAN RICEGRASS XED SHRUB I SAND DUNES Fig. 10. Major plant types as designated by McBride. Reproduced by permission of Ray McBride, Atomic Energy Comm., Idaho Falls. Ticks of Reactor Testing Station STUDY AREAS Twelve primary sites (Areas 1 to 12) were studied periodically in the major vegetation types of the NRTS, and 28 secondary areas were studied at less regular intervals (Fig. 2). These latter areas were chosen because of their atypical nature in relationship to the extensive vegetative types. Area 1. Two miles west of Big Lost River bridge along highway US 20-26, thence 0.7 mile north-northwest along old paved highway, thence 100 yards north of road (Fig. 11). This is designated as a Chrysothamnus-Artemisia community, and occurs in McBride's Big Sage- brush-Bluebunch Wheatgrass-Rabbitbrush type. Plant analysis: Chrysothamnus 38%; Artemisia 24%; grasses {Oryzopsis, Sitanion, Stipa) 15%; forbs (Allium, Astragalus, Descurainia, Eriogo- num, Lappula, Lupinus, Umbelliferae, Ziga- dcnus) 10.5%; Opuntia 6%; Tetradymia 1.5%; bare ground 5%. Area 2. Six miles northeast of junction of highways Idaho 22-88 and US 20-26 along high- way Idaho 22-88, thence 125 yards east of road ( Fig. 12 ) . This Artemisia-Chrvsothamnus com- munity occurs in McBride's Big Sagebrush- Winterfat-Rabbitbrush type. Plant analysis: Artemisia 50%; Chrysothamnus 15%; grasses (Agropyron, Oryzopsis, Poa, Stipa) 15%; Euro- tia 10%; forbs (Astragalus, Castilleja, Cryptan- tlia, Erigeron, Eriogonum, Lupinus) 7%; bare ground 3%. ''>'iF-^>''SP\ . <« ■%p'^Mm^ . ^i Fig. 14. Study area 4. Predominant plants are Oryzopsis and Stipa. 12 Brigham Young UNrvEHsixY Science Bulletin ■:U:.:^^ ".r-^tir*^.- •?. ' ^:j - Fig. 15. Study area 5. Predominant plant is Juniperus. h:--- ■■^■: J-j-' •*•. Oi*«it!f >^ ^ ■*■■ • ^ l.^*' ' ■ ;*:- " --^l; v;, ■'4.-V*f ■•«" »-■■■* ,^c,.>.- m ; Eurotia 9%; forbs and grasses (misc. grasses and Salsola kali) 4%; bare ground 16%. Area 8. One mile northeast of junction of highways Idaho 22 and 88 along Pole Line Road, thence north 3 miles along old road (Fig. 18). Artemisia-Atriplex community occurs in Mc- Bride's Big Sagebrush-Winterfat-Rabbitbrush type. Plant analysis: Artemisia 30%; Atriplex 26%; Chrysothamnus 5%; forbs and grasses (misc. grasses, Penstemon, Phlox) 1%; bare ground 38%. Area 9. Two and eight-tenths miles north of junction of highways Idaho 22 and 88 along highway 22, thence 0.2 mile east of highway (Fig. 19). This Chenopodium-Eurotia commu- nity occurs in McBride's Rabbitbrush-Big Sage- brush-Grass type. Plant analysis: Chenopodium 40%; Eurotia 35%; Artemisia and Chrysotham- nus 2%; Salsola 2%; forbs 1%; bare ground 20%. Area 10. Five miles northeast of NRTS facility NRF along Lincoln Blvd., thence east 1.8 miles along Shell Road, thence 125 yards south of road (Fig. 20). This Artemisia com- munity occurs in McBride's Big Sagebrush-Rice- grass-Needlegrass type. Plant analysis: Arte- misia 68%; Opuntia 7%; grasses {Agropyron, Oryzopsis, Stipa) 7%; forbs (Astragalus Sphae- ralcea, Umbellif erae ) 5%; bare ground 13%. Area 11. Along Lincoln Blvd., 0.5 mile south of its junction with highway Idaho 88, thence 220 yards east of Lincoln Blvd. (Fig. 21). Chrysothamnus-Grass-Tetradymia community oc- curs in McBride's Ricegrass-Needlegrass-Rabbit- brush-Cactus type. Plant analysis: Chrysotham- nus 30%; grasses (Agropyron, Oryzopsis, Stipa) 26%; Tetradymia 22%; Artemisia 5%; Opuntia 5%; forbs (Allium, Astragalus, Comandra, Erio- gonum, Cilia, Lupinus, Phlox) 5%; Atriplex 2%; bare ground 5%. Area 12. Along highway Idaho 22, 1.5 miles north of its junction with highway Idaho 88, thence 175 yards wesit of highway (Fig. 22). Juniper (with understory) community occurs in McBride's Juniper-Big Sagebrush-Bluebunch Wheatgrass type. Plant analysis: Juniperus 30%; Chrysothamnus 15%; Eurotia 15%; Artemisia 13%; grasses ( Oryzopsis, Stipa) 10%; forbs (Chenopodium, Eriogonum, Mentzelia, Salsola) 6%; Opuntia 1%; bare ground 10%. Area 13. Nuclear waste burial ground 2 miles southwest of NRTS facility EBR-I. This is in McBride's Big Sagebrush-Bluebunch Wheat- grass-Rabbitbrush type. Area 14. One-half mile west of Lincoln Blvd. along Big Lost River south of NRTS fa- cility TRA. This is in McBride's Big Sagebrush- Thickspike Wheatgrass-Needlegrass type. Area 15. Grassy sinks area west of Big Lost River, near its junction with Shell Road, 3.5 miles west of Lincoln Blvd. This is in McBride's Bluestem Wheatgrass-Wiregrass-Iva type. Area 16. Nuclear waste burial ground 0.5 mile north of NRTS facility EBR-II. This is in McBride's Big Sagebrush-Rabbitbrush-Foxtail type- Area 17. Along highway US 26 one mile southeast of its junction with highway US 20, thence 0.6 mile south-southeast along old Black- foot Highway, thence west to lava flow. Situat- ed in McBride's Mixed Shrub type. Area 18. The sand dunes area is situated immediately north of highway Idaho 88 approxi- mately 0.6 mile west of its junction with Lincoln Blvd. (Fig. 23). Area 19. This cave is situated south of NRTS facility EBR-II (Fig. 6). Proceed north from highway US 20 for 0.2 mile from its junc- tion with Grant Road. At this point take the left fork and proceed for an additional 0.8 mile to the cave which is on the east side of the road. This is in McBride's Big Sagebrush-Rabbitbrush- Foxtail type. Area 20. The west-facing side of a rocky outcrop may be reached by proceeding west- ward along a dirt road that extends to the top of the hill west of the junction of highways Idaho 22 and 88. This is in McBride's Juniper- Big Sagebrush-Bluebunch Wheatgrass type. Area 21. This cave is situated slightly to the north of and between East and Middle Buttes (Fig. 7). Proceed south from highway US 20 along a dirt road, which junctures near highway US 20 mileage post 295, until an eastward road extending along the northern sides of the buttes is reached. Turn east and proceed for approxi- mately 0.4 mile to a road which extends south- ward for a short distance to the cave. Area 22. A reservoir is situated adjacent to NRTS facility TRA. This is in McBride's Big 14 Brigham Young Unineksity Science Bulletin ,^.| tifi"- J' V l\ ■^^'*^^. ^^^ ' ^5^^f^i^.^.i| ^, f .^v- \>' •* Fig. 17. Study area 7. Predominant plants are Chrysothamnus and Artemisia. ^^-^^-3^-^^^Sfc^ C ^ ■ .. . V •■'• •«•;■ ■ ■i Fig. 18. Study area 8, with Lemhi Mtns. in the background. Predominant plants are Artemisia and Atriplex. Ticks of Reactor Testing Station 15 .'^■ ij"V*t.i -^«aMMt<'' 3 ^/-.^* --W.'' sf^ ,. . -i.-^' .>««>,• Fig. 19. Study area 9, with twin buttes in the background. Predominant plants are Chenopodium and Eurotia. SijS^i»5SfeiJ^«*i»B«s*«*«vi->>.^ ,, S-^*;- ■?':-•■ Fig. 20. Study area 10. Predominant plant is Artemisia. 16 Brigham Young Untv'eksity Science Bulletin ■> '" d ' f- 4:y 4 .^i&- ■' / r U-i Fig. 21. Study area 11. Predominant plants are Chrysothamnus, grasses, and Tetradymia --' f. 'T^ >-* yi^-'^ o« Fig. 22. Study area 12. Predominant plants are Juniperus, Chrysothamnus, Eurotia, and Artemisia. i Ticks of Reactoh Testing Station 17 '*-.-• - * •* ' vtf^ ■ .1-^- ■ *. «>- '" . »■ Fig. 23. Sand dunes ( study area 18 ) with predominant fringing plant, Salsola. Sagebrush - Thickspike Wheatgrass - Needlegrass type. Area 23. Frenchman's Well is situated ad- jacent to a cabin, west-northwest of and approxi- mately 0.5 mile from Big Southern Butte. This may be reached by proceeding from the area of NRTS facility EBR-I to the peripheral road which encircles Big Southern Butte and its junc- tion with the road which proceeds to the top of the butte. Area 24. Webb Springs is situated on the east-northeast slope of Big Southern Butte. The canyon where the spring occurs has a dense growth of aspen and other trees. It may be reached by proceeding along the peripheral road which encircles the butte. Area 25. Top and slopes of Middle Butte. This butte is entirely basalt with mixed brush and junipers. It occurs in McBride's Juniper- Big Sagebrush-Bluebunch Wheatgrass type. Area 26. A lava outcrop is situated 3.5 miles along Shell Road east of its junction with Lin- coln Blvd. This is in McBride's Big Sagebrush- Rioegrass-Needlegrass type. Area 27. A volcanic sink may be reached by proceeding eastward along Shell Road for 4.1 miles from its junction with Lincoln Blvd., thence along the right fork for an additional 0.5 mile. This is in McBride's Big Sagebrush-Rice- grass-Needlegrass type. Area 28. A burned area situated in Mc- Bride's Big Sagebrush-Bluebunch Wheatgrass- Rabbitbrush type. Proceed south along a dirt road, which junctures with highway US 20-26 east of mileage post 279, until the road crosses a railroad track. Turn right and proceed along the dike until the road crosses a dam on the Big Lost River. Approximately 300 yards further west the burned area is situated on the south side of the river. Area 29. A lava outcrop is situated along Shell Road, 0.4 mile east of its junction with Lincoln Blvd. This is in McBride's Big Sage- brush-Tliickspike Wheatgrass-Needlegrass type. Area 30. The Experimental Dairy Farm may be reached by proceeding north from NRTS facility OFA along Lincoln Blvd. for 5.8 miles, thence east along Stage Road for 1.7 miles. This is in McBride's Big Sagebrush-Thickspike Wheat- grass-Needlegrass type. Area 31. Cinder Butte is situated a short distance east of Lincoln Blvd., about 0.5 mile south of its junction with highway Idaho 88. This is on the hne between McBride's Ricegrass- Needlegrass-Rabbitbrush-Cactus and Big Sage- brush-Thickspike Wheatgrass-Needlegrass types. Area 32. At the north edge of NRTS facility CFA is an area occupied by huge concrete blocks formerly used as targets for naval ordnance. The plants are predominantly Artemisia, Chryso- thamnus, and Salsola, with few annuals. This occurs on the border between McBride's Big Sagebrush-Rabbitbrush-Foxtail and Big Sage- brush-Thickspike Wheatgrass-Needlegrass types. Area 33. This volcanic cave is situated ap- proximately 200 yards north of highway US 20, three miles east from its junction with the road 18 Brigham Young University Science Bulletin that proceeds north to NRTS faciUty EBR-II (Fig. 9). This is in McBride's Big Sagebrush- Bluebunch Wheatgrass-Rabbitbrush type. Area 34. This area is approximately 300 yards north of study area 3. It is on the border- line of the Canada Wild Rye-Rabbitbrush-Big Sagebrush and Big Sagebrush-Rabbitbrush-Pox- tail types of McBride. Area 35. The Salix community is situated 200 yards southwest of the reservoir near NRTS facility TRA. It occurs in McBride's Big Sage- brush-Thickspike Wheatgrass-Needlegrass type. Area 36. This area extends over a linear dis- tance of approximately six miles, but is centered alongside the old Blackfoot Highway, about 4 miles north of Atomic City. The predominant vegetation is Ehjmus, Artemisia, and Chrijso- thamnus. It is situated in McBride's Crested Wheatgrass type. Area 37. Proceed nine and one-half miles northeast of the junction of highways Idaho 22- 88 along highway Idaho 22 to a point approxi- mately 200 yards north of Birch Creek, thence west 250 yards from paved road. Predominant plants are Artemisia, Chrysothamnus, and Pru- nus ( Chokecherry ) along the creek. This is in McBride's Black Sagebrush-Big Sagebrush- Shadscale type. Area 38. This area is 0.3 mile northeast of the burial grounds which lie southwest of NRTS facility EBR-I. The predominant plants are Agropyron with some Aiiemisia. This is in McBride's Big Sagebrush-Bluebunoh Wheat- grass-Rabbitbrush type. Area 39. This area is 0.8 mile west of NRTS facility GCRF along paved road (Wilson Blvd.), thence O.I mile south of highway in a lava outcrop. The predominant vegetation is Artemisia. This area is in McBride's Big Sage- brush-Rabbitbrush-Foxtail type. Area 40. This area is 0.3 mile west of NRTS facility GCRF along paved road (Wilson BJvd. ), thence 0.1 mile south of highway in a lava outcrop. The predominant vegetation is Artemisia. This area is in McBride's Big Sage- brush-Rabbitbrush-Foxtail type. PROCEDURES Vertebrate Collections Six styles of traps were utilized. The YAW hve-catch trap is made of 3-mesh, galvanized hardware cloth with a galvanized sheetmetal door and reinforcement bands. It is 15 inches long by 4/2 inches square (Fig. 24). The modi- fied Hubbard trap is made of galvanized sheet- metal, and is 8 inches long and 3 inches square ( Fig. 25 ) . The can pit-trap consists of an outer, galvanized metal case, 7 inches in diameter and 14 inches long, with a stainless steel, flanged in- ner can of shghtly smaller size (Fig. 26). Other Fig. 24. YAW live-catch trap. Fig. 25. Modified Hubbard live-catch trap. ! Ticks of Reactor Testing Station 19 Fig. 26. Components of a can pit-trap. traps used were the Museum Special, break-back trap (Fig. 27) and the Oneida-Victor carnivore trap (Fig. 28). The Cahfornia gopher trap was used only to a small extent. The YAW, Museum Special, and Hubbard traps were baited with rolled oats, but bait was not used in the other traps. Rifles and shotguns were used to collect rabbits, carnivores, and birds. Reptiles were captured by hand and in can pit-traps. Visual observations of vertebrates were recorded, and many animals found dead along roads and high- ways were collected. In each of the 12 primary study areas, traps were arranged in a radiating pattern (Fig. 29). Two traps were placed at each station, with stations approximately 30 feet apart. This same procedure was followed in some cases in the secondary areas, although the traps usually were arranged in single-line transects or randomly Fig. 28. Oneida-Victor carnivore trap. placed. Only one can pit-trap was placed at each station, with stations 50 feet apart. Small mammals found alive in the traps were killed by spinal separation, then put into white paper sacks which were sealed by folding and stapling the tops. Other animals captured alive were killed with chloroform, and these as well as those which were shot were similarly put into paper sacks. All animals returned to the labora- tory were refrigerated until further studies could be made. Representative specimens of birds and c o o o ^O o O^ o o O o o o O^ o^ o o o Q o o o o o o o o o o o DnaDDDDDDn • p DnDDDDDDDnH O o O O O O O O o O ^o O o O ^o o o o Y o o Y Fig. 27. Museum-Special break-back trap. Fig. 29. Trap arrangement used in the major study areas. P, metal post; C, can pit-traps; S, Museum- Special traps; H, Hubbard traps; Y, YAW traps. 20 Brigham Voung University Science Bulletin mammals were permanently preserved as muse- um skins, whereas many reptiles were preserved formalin. Rattlesnakes were taken alive in m most cases, examined for ectoparasites, marked for identification purposes, and released. Most of the rabbits, small mammals, and many of the common birds which were damaged were dis- carded after they were examined for ectopara- sites. I)uring the study period, traps were operated for a total of 57,330 trap nights. Relative to each type of trap, this included 14,040 for YAW, 12,000 for Hubbard, 10,800 for Museum Special, and 20,490 for can pit-traps. Invertebrate Collections Most of the ectoparasites were collected by a cooling and warming method. The vertebrate hosts were maintained under normal refrigera- tion (about 40° F) until their bodies had thor- oughly cooled ( a few hours to overnight ) . They were then put into a large, white enamelware pan over which was placed a 100-watt lamp. When the ectoparasites became active and moved away from the cold body of their host, they were retrieved and preserved in vials of 70 percent ethyl alcohol. Additional specimens were obtained by searching through the fur, feathers, or scales of their vertebrate hosts. Fleas, lice, mites, and some ticks were mount- ed on microslides for identification purposes, whereas most ticks were retained in alcohol. Free-hving arthropods were collected prin- cipally in can pit- traps. Others were taken by modified Berlese funnel extraction (Fig. 30), some by hand, and others with aerial and beat- ing nets. Vlany were preserved in 70 percent ethyl alcohol, whereas others were pinned. Fig. 30. Berlese funnel. VERTEBRATES EXAMINED The kinds and numbers of vertebrates ex- amined for ectoparasites are listed below (also see Table 1). Scientific and common names, number examined, months and areas of collec- tion, and additional comments are listed in se- quence, respectively. In most cases the area of collection is listed only when it is one of our 40 specific study sites. Species are listed in alpha- betical sequence without regard to phylogenetic relationship in each major category. Amphibians Rana pipiens hrachycephala (Western Leo- pard Frog). 1; Aug.; area 28. Scaphiopus hammondi intermontanus (Great Basin Spadefoot). 26; June-Aug.; mainly area 6, also in 11, 12. Most taken in July. Reptiles Coluber constrictor mormon (Western Yel- low-bellied Racer). 1; May; area 24. Crotahis viridis hitosus (Great Basin Rattle- snake). 95; May-Oot.; areas 3, 19, 20, 24, 36. Most taken from den at area 19 during May, June. Eumeces skiltonianus utahensis (Western Skink). 5; June, Aug.; area 19. Masticophis taeniatus taeniatus (Desert Striped Whipsnake). 1; May. Ticks of Reactor Testing Station 21 Phrynosoma douglassi douglassi (Pigmy Homed Lizard). 19; April-Sept.; areas 1, 4, 6, 7, 8, 11, 19. Most (11) taken at area 7. Pituophis catenifer deserticola (Great Basin Gopher Snake). 8; May-Oct.; areas 10, 19, and other. Sceloporus graciosus (Spiny Lizard). 314; April-Oct.; areas 1, 2, 3, 6, 7, 8, 10, 11, 12, 18. Most collected in June, and from areas 1, 6, 12. Thamnophis elegans vagrans (Wandering Garter Snake). 3; Oct.; areas 19, 20. Bh-ds Acanthis flammea (Common Redpoll). 1; Nov.; area 24. Actitis macularia (Spotted Sandpiper). 1. July. Aegolius acadicus (Saw Whet Owl). 2; Sept. Aeronautes saxatilis (White-throated Swift). 1; June. Agelaius phoeniceus (Red- winged Black- bird ) . 1 ; June ; Howe ( city ) . Aimophila cassinii (Cassin's Sparrow). 2; May; areas 1, 24. Alectoris graeca (Chuckar). 1; May; area 24. Amphispiza belli (Sage Sparrow). 38; April- Sept.; areas 1, 2, 7, 10, 11, 23, 24, 25, 29. Most collected in April. Amphispiza bilineata (Black-throated Spar- row). 1; Aug.; area 7. Anas discors (Blue-winged Teal). 1; July; area 15. Anas platyrhynchos ( Mallard ) . 2; Jan., May. Anthus spinoletta (Water Pipit). 7; April, Oct.; area 23. Asio flammeus (Short-eared Owl). 4; April, June, Aug., Oct. Asyndesmiis lewis (Lewis' Woodpecker). 2; May. Bomhycilla cedrorum (Cedar Waxwing). 1; Sept.; Atomic City. Bomhycilla garrulus (Bohemian Waxwing). 8; Nov.; area 24. Bubo virginianus (Great Homed Owl). 1; Nov. Buteo lagopus (Rough-legged Hawk). 4; Nov., I>ec., Jan. Buteo regalis (Ferruginous Hawk). 2; Aug.; areas 4, 39. Buteo swainsoni (Swainson's_ Hawk). 3; April, May, Aug.; Atomic City and other. Carpodacus mexicanus (House Finch). 2; Nov.; area 20. Calamospiza melanocorys (Lark Bunting). 4; May, July; areas 7, 24. Centrocercus urophasianus (Sage Grouse). 18; Feb., May, July, Oct., Nov.; area 24 and other. Charadrius vociferus (Killdeer). 4; May, Aug.; area 23 and other. Chordeiles minor (Common Nighthawk). 5; June, Aug.; areas 1, 7, Atomic City and odier. Circus cyaneus (Marsh Hawk). 1; April. Colaptes cafer (Red-shafted Flicker). 5; May, Sept.; area 24, Atomic City. Columha liva (Rock Dove). 2; Feb.; Howe (city). Cyanocitta stelleri (Steller's Jay). 1; Sept.; area 20. Dendrocopos villostis (Hairy Woodpecker). 1; June; area 24. Dendroica petechia (Yellow Warbler). 2; May, June; Atomic City and other. Empidonax difficilis (Western Flycatcher). 3; June. Empidonax hammondii (Hammond's Fly- catcher). 1. Empidonax oberholseri (Dusky Flycatcher). 1; May; area 24. Empidonax traillii (Traill's Flycatcher). 1; June; Atomic City. Eremophila alpestris (Homed Lark). 84; Oct.-Aug.; areas 3, 4, 6, 8, 9, 10, 11, 23, 30, 36. Most birds taken in April, and from area 23. Ereunetes nmuri (Western Sandpiper). 6; May, Aug.; area 19 and other. Euphagus cyanocephalus (Brewer's Black- bird). 9; April, May; area 23, Atomic City and other. Falco sparverius (Sparrow Hawk). 6; April, June, Aug., Sept.; area 39 and other. Hirundo rustica (Bam Swallow). 1; Sept. Catharus guttatus ( Hermit Thrush ) . 2; May; area 24. Catharus ustulatus (Swainson's Thmsh). 1; June. Junco oreganus (Oregon Junco). 30; Mar., April, Sept., Oct.; areas 19, 20, 22, 23, 24, 25 and other. Lanius excubitor (Northern Shrike). 7; Oct., Dec, Jan., Mar.; area 23 and other. Lanius ludovicianus (Loggerhead Shrike). 20; Mar., April; area 2, Atomic City and other. 22 Bricham Young University Science Bulletin Leucosticte tephrocotis (Gray-crowned Rosy Finch). 25; Jan., Feb., Nov; areas 19, 21, 24 and other. Loxia ctirvirostra (Red Crossbill). 3; Nov.; area 2A. Melospiza melodia (Song Sparrow). 1; Mar.; area 3. Mtjadestes townsendt ( Townsend's Solitaire ) . 6; Oct.; area 23, 24. Myiarchus cinerascens (Ash-throated Fly- catcher). 1; May. Nycticorax nycticorax (Black-crowned Night Heron). 1; Aug.; area 15. Niittallornis borealis (Olive-sided Flycatch- er). 2; May, June; area 2, Howe (city). Oreoscoptes montanus (Sage Thrasher). 13; April, May, July, Sept.; areas 1, 7, 10, 11, 36 and other. Parus gambeli (Mountain Chickadee). 2; Sept.; area 20. Passe'rina amoena (Lazuli Bunting). 6; May, June; area 24, Atomic City and other. Perdix perdix (Gray Partridge). 2; Dec. Petrochelidon pyrrhonota (Cliff Swallow). 3; May, June. Phasianus colchicus (Ring-necked Pheasant). 5; Jan., Feb., Dec; Howe (city) and other. Pica pica (Black-billed Magpie) 8; Jan., Feb., June, Oct., Dec. Piranga ludoviciana (Western Tanager). 16; May, June; area 24, Atomic City. Podiceps caspicus (Eared Grebe). 1; May. PoUoptila caerulea (Blue-gray Gnatcatcher). 1; May. Pooecetes gramineus (Vesper Sparrow). 13; April-Aug.; areas 1, 3, 10, 16 and other. Recurvirostra americana ( American Avocet ) . 1; July. Regulus calendula (Ruby-crowned Kinglet). 4; May; area 20 and other. Regulus satrapa (Golden-crowned Kinglet). 1; Oct. Salpinctes ohsoletus (Rock Wren). 17; May, June, July, Sept., Oct.; areas 19, 20, 24 and other. Sayornis saya (Say's Phoebe). 2; May, Sept.; areas 19, 23. Seiurus noveboracensis (Northern Water- thrush). 1; May; area 24. Sehsphorus platycercus (Broad-tailed Hum- mingbird). 1; June; area 24. Sialia currucoides (Mountain Bluebird). 5; Mar., April, Sept., Oct.; areas 9, 25 and other. Spinus pinus (Pine Siskin). 23; Jan., June, Oct., Dec. Spinus tristis ( American Goldfinch ) . 1; June; Atomic City. SpizeUa arborea (Tree Sparrow). 1; Nov.; area 24. SpizeUa brcweri (Brewer's Sparrow). 7; June, Sept.; areas 2, 24 and other. Sturnella neglecta (Western Meadowlark). 7; Mar., April, May, July, Sept., Oct.; areas 3, 24 and otlier. Sturnus vulgaris ( Starling ) . 9; June, Sept., Oct.; Atomic City and odier. Turdus migratorius (Robin). 11; April, June, Sept.; area 24 and other. Tyrannus tyi annus (Eastern Kingbird). 3; May, June; area 24 and other. \ViIs07iia pusiUa (Wilson's Warbler). 3; May, Sept.; area 24 and otlier. Zenaidura macroum (Mourning Dove). 23; April-July; areas 2, 24 and other. Zonotrichia leucophnjs ( White-crowTied Sparrow). 33; May, Sept., Oct.; areas 19, 22, 23 and other. Mammals Antilocapra americana (Antelope). 3; April, May, Nov. Canis latrans (Coyote). 6; Jan., Feb., Aug., Nov., Dec. Dipodomys ordii (Ord Kangaroo Rat). 808; Mar.-Nov.; all areas except 17, 19-21, 24, 25, 29- 32, 34, 35, 38. Mostly in areas 4, 12; seasonally most abundant in Aug. Erethizon dorsatuni (Porcupine). 1; Aug.; area 19. Eutamias minimus (Least Chipmunk). 398; Mar.-Nov.; all areas except 4, 20, 21, 24, 25, 28-35. Most abundant in areas 1 and 36; most common in July. Lepus californicus ( Black -tailed Jackrabbit). 125; every month except Feb.; areas 5, 10, 13 and other. Lynx rufus (Wildcat). 8; Jan., Feb., April, Nov.; area 19 and other. Marmota flaviventris (Yellow-bellied Mar- mot). 6; May-July; areas 4, 36 and other. Microtus montanus (Montane Vole). 25; Jan., Mar., June-Aug., Oct.. Nov.; areas 1, 3, 15, 27, 30, 38, 39. Mus musculus (House Mouse). 1; June; area 32. Mustela frenata (Long-tailed Weasel). 4; July, Aug., Oct.; areas 23, 33, 38 and other. Ticks of Reactor Testing Station 23 Myotis evotis (Long-eared Myotis). 2; Mar.; area 20. Myotis subulatus (Small-footed Myotis). 1; Mar.; area 20. Neotoma cinerea (Bushy-tailed Woodrat). 14; June-Sept., Nov., Dec; areas 5, 17, 19, 20, 24, 29, 32, 33. •Oni/chomys leucogaster (Grasshopper Mouse). 63; Mar.-Etec; areas 1-3, 7, 9-14, 16, 28, 36, 38. Mostly in area 1. Ovis aries. (Sheep). 1; July. Peromyscus maniculatus (White-footed Mouse). 1866; every month; every area. Mostly area 3; most common June-Aug. Perognathus parvus (Great Basin Pocket Mouse). 474; April-Nov.; all areas except 12, 15, 20-22, 24, 25, 27-35. Mostly areas 1, 2, 7, 10, 36; most common May-Aug. Plecotus townsendii (Long-eared Bat). 78; Sept.-April; areas 19, 21, 33 and other. Mostly areas 18, 54. Reithrodontomys megalotis (Harvest Mouse). 39; Mar.-Aug., Oct.; areas 1-3, 7, 10, 11, 13, 15-17, 36, 37. Mostly July, Aug. Sorex merriami ( Merriam's Shrew ) . 9; April, June-Aug., Oct.; areas 1, 6, 7, 11, 12, 19. Spermophilus toicnsendii (Townsend's Ground Squirrel). 60; Mar.-July; areas 1-3, 6-9, II, 13, 36-38 and other. Mostly June; common areas 3, 9. Sylvilagus idahoensis (Pygmy Rabbit). 13; Feb., April, June-Aug., Nov.; areas 1-3, 38 and other. Sylvilagus nuttallii ( Nuttall's Cottontail ) . 28; Feb., Mar., May-Dec; areas 3, 10, 15, 17, 19-21, 32, 36, 38 and other. Taxidea taxus (Badger). 5; April-June, Aug., Nov.; areas 30, 37 and other. Thomomys talpoides (Northern Pocket Gopher). 8; Mar., June, Sept.-Nov.; areas 1, 2, 7, 9, 12, 19, 24. Vulpes fulva (Red Fox). 4; July; Howe (city). Table 1. Total number of maminals examined in each study area. Study No. Study No. area mammals area mammals 1 316 21 59 2 166 22 6 3 561 23 65 4 112 24 5 5 144 25 6 6 165 26 24 7 222 27 9 8 200 28 19 9 139 29 30 10 195 30 14 11 106 31 1 12 165 32 59 13 154 33 24 14 56 34 1 15 88 35 11 16 98 36 349 17 20 37 32 18 53 38 52 19 45 39 35 20 9 40 53 TIGKS Dermacentor andersoni Hosts General infestation. Birds of three species and mammals of 16 species were infested (Tables 2 and 3). Only larvae were found on birds. Rodents generally were infested with larvae and nymphs, whereas adult ticks were found only on rabbits and larger mammals. Exceptions were Lepus which possessed larvae, nymphs, and adults; Sylvilagus which possessed only nymphs; and a single Peromyscus which was host for an adult male tick. Table 2. Total numbers of ticks collected by species and stage of development. Species la ny ad 5 ad 5 Total Dermacentor andersoni 3221 542 16 6 3785 Haemaphysalis leporispalustris 73 94 16 10 193 Ixodes kingi 4806 1972 27 66 6871 Ixodes angustus ( ? ) 1 1 Ixodes marmotae 15 1 16 Ixodes ochotonae 25 6 31 Ixodes sculptus 1 3 4 Ixodes sp. 64 4 68 24 Brigham Young University Science Bulletin Table 3. Hosts, degree of infestation, and general seasonal occurrence for three developmental stages of Dermacentor andersoni. %" of hosts Tick-host Seasonal Host species infested by la, ny, ad index' occurrence Birds Calamospiza melanocortjs la 25 (4)""' 1 July Centrocercus UTophasianus la 6 (18) 1 Pooecetes gramineus la 8 (13) 19 Mammals Dipodomijs ordii la 7 4 June- Aug. ny 6 2 May- Etitamias minimus la 11 5 June- ny 11 2 May- Homo sapiens ad June Lepus califonucus la 3 1 Aug. ny 7 3 July-Aug. ad 8 2 Mar.-April, Aug., Nov. Lynx ruftis ad 1 (8) 1 April Microtus montanus la 8 1 Aug. ny 4 1 July Mustela frenata la 25 (4) 1 July Neotoma cinerea la 7 (14) 10 June ny 29 (14) 5 " -Aug. Onychomys leucogaster la 8 2 July- " ny 13 2 June- Perognathus parvus la 15 9 - ny 14 2 May- Peromyscus manicuUttus la 19 6 June- ny 8 2 May- ad 0.05 1 April Reithrodontomys megalotis la 8 1 July-Aug. ny 3 1 Aug. Spermophilus townsendii la 2 2 June ny 18 3 May-July Sylvilagus idahoensis ny 8 (13) 6 Aug. S. nuttallii ny 11 1 July. Oct. Taxidea taxus ad 20 (5) 1 April °To nearest whole number. "Number in parentheses following percentage in- fested indicates the actual number of hosts examined when less than 20 were collected. Larvae. Birds were not frequently infested. One larva each was taken from Calamospiza and Centrocercus, whereas one Pooecetes pos- sessed 19 larvae. Of the total mammals examined where large numbers were taken ( (30 or more ) , a higher per- centage of Peromyscus (24%) were infested than were other hosts. The tick-host index (ratio of ticks to number of infested hosts) was high- est on Lepus and Perognathus (10 and 9, re- spectively). Highest incidence of infesitation was 39% for Perognathus in August. The highest tick-host index was for Perognathus in June (11.6). Average indices on all hosts varied from a low of 5.2 in June to a high of 6.2 in July. There was no proportional oorrelation between the percentage of animals infested and tick-host index. Relative to an average of all hosts examined, there was no significant difference between the degree of infestation of male and female hosts. However, the males of Perognathus and Pero- myscus were much more heavily infested than were the females (tick-host index 9.3j':6.4 9 and 8.2<:r:4.7$, respectively). The opposite situation occurred with Dipodomijs and Euta- mias (3.1 cf :4.9 ? and 3.7 d' : 5.9 5 , respectively). Nymphs. Nymphs were found on a greater percentage of Spermophilus (26%) than on other hosts. The tick-host index (3) was also highest for this host. The highest incidence of infestation found was for Spermophilus (.50% in- fested in August). Highest tick-host index also was related to Spermophilus — 7.2 in August. Average indices on all hosts varied from a low^ of 1.0 in October to a high of 2.4 in July. There was no proportional correlation between the per- centage of animals infested and the tick-host index. There was no significant difference between the degree of infestation of males and females for any of the host species. Adults. Few adults were taken. Thirteen of the 16 males collected were taken from eight Lepus, and three of the six females from three Lepus. Seasonal Incidence Larvae. Ticks of this stage were found only in June, July, and Augu.sit. Highest percentages of animals were infested in July. Nymphs. Nymphs were found from May into August; two specimens were taken from Sylvilagus ( cottontail ) in October. Highest per- centages of animals were infested in June and August. Adults. Males were taken in March, Aj>ril, June, and November, and females in March, April, June, and August. i Ticks of Reactor Testing Station 25 Haemaphysalis leporispalustris Hosits General infestation. Birds of 10 species and mammals of six species were infested (Tables 2 and 4 ) . Only larvae were foimd on birds witih the exception of three Salpinctes which pos- sessed six nymphs. Pew rodents were infesited, and most mammalian records were for rabbits which possessed larval, nymphal, and adult ticks. Larvae. Of all birds examined where more than a few were taken (10 or more), a higher percentage of Salpinctes (53%) were infested Table 4. Hosts, degree of infestation, and general seasonal occurrence for three developmental stages of Haemaphysalis leporispalustris. X" of hosts Tick-host Seasonal Host species infested by la, ny, ad index" occurrence Birds Amphispiza belli la 5 2 Aug. Centrocercus urophasianus la 6 (18)"° 1 July Junco oreganus la 7 2 Sept.-Oct. Nttttallornis borealis la 50 (2) 2 June Oreoscoptes montanus la 8 (13) 1 July Pica pica la 13 (8) 1 June Polioptila caerulea la 100 (1) 1 May Salpinctes obsoletus la 53 (17) 3 May, Sept.- Oct. ny 18 (17) 2 Sept. Selasphorus platycercus la 100 (1) 1 June Sturnella neglecta la U (7) 1 July Mammals Lepus californicus la 10 6 Mar.-Oct. ny 16 3 April-Nov. ad 23 2 May-Sept. Onychomys leucogaster la 2 1 Sept. Perognathus parvus la 0.2 1 May Peromyscus maniciilatus ny 0.2 1 April-June Sylvilagus idahoensis la 8 (13) 1 July ny 15 (13) 8 June-July ad 8 (13) 1 " S. nuttallii la 18 3 "- Sept. ny 29 9 May- ad 18 4 June-July °To nearest whole number. °'' Number in parentheses following percentage in- fested indicates the actual number of hosts examined when less than 20 were collected. than were other species. The tick-host index for the birds was higihesit for Salpinctes (2.6); for other species it was 2.0 or less. For the mammals, incidence of infestation was highest for Sylvilagus nuttallii (17.8%), al- though Lepus ( which had an infestation rate of 9.6%) had the highest tick-host index, 6.2, as compared with 3.4 for S. nuttallii. The tick-'host index was much higher for male than for female Lepus (14cr:6.4$ ) but was not significantly different for other species of hosts. Nymphs. The only birds possessing nymphs were Salpinctes which had an infes'tation inci- dence of 17.6% and a tick-host index of 2.0. The mammal with the highest incidence of infestation and tick-hosit index was Sijlvilagus nuttallii with 28.6% and 9.4, respectively. One nymph each was found on two Peromyscus; otherwise, most nymphs were found on rabbits. The tick-host index was much higher for male than for female Sylvilagus nuttallii ( 15.3 cT : 6.0 ? ) , but was not significantly different for other species of hosts. Adults. Adult ticks were found only on rabbits. Highest incidence of infestation was for Lepus (23.2%), but the highest tick-host index was for Sylvilagus nuttallii (4.2). Only female Sylvilagus nuttallii were infested with adult ticks, whereas both sexes of other hosts were infesited. Seasonal Incidence Larvae. Larvae were found on birds and mammals from May into October, and one larva was found on a mammal in March. Insufficient numbers were taken to determine the month of highest incidence. Nymphs. Nymphs were found on birds in Septembier, and on mammals from April into November. Highest incidence apparently was June into August. Adults. Males were taken from May into September, and females from May into July. Highest incidence was from June into August. Ixodes kingi Hosts General infestation. Mammals of 11 species were infested (Tables 2 and 5). Principal hosits for these ticks are Onychomys, Dipodoinys, Perognathus, and Peromyscus. Larvae. The highest incidence of infesta- tion occurred with Onychomys. Dipodomys 26 Bricham Young Universitv Science Bulletin Table 5. Hosts, degree of infestation, and general seasonal occurrence for three developmental stages of Ixodes kingi. %° of hosts Host species infested by la, ny, ad Tick-host Seasonal index" occurrence Mammals Dipodomys ordii Eutamias minimus Lepus californicus Microtus montanus Mustela frenata Neotoma cinerea Omjchomijs teucogaster Perognathus parvus Peromyscus maniculatus Spermophilus townsendii Taxidea taxus Thomomys talpoides la Z'J, ny 40 ad 5 la 4 ny 1 ad 0.2 la 2 ny 4 la 0.8 ny 1 ad 2 la 14 (14)° ny 36 (14) la 46 ny 22 ad 2 la 12 ny 14 ad 0.4 la 10 ny 5 ad 0.05 la 5 ny 7 ny 20 (5) ad 60 (5) la 13 (8) ny 13 (8) 13 4 4 2 4 2 1 6 3 1 23 8 1 2 14 12 Mar.-Nov. " - Oct. Mar., June- July, Oct. Mar., June- July June April Aug. July-Oct. June-Aug. " - Sept. Mar., June- Dec. June-Oct. July May-Oct, July Jan. -Dec. "- Oct. March Mar., June Aug. April, Nov. June "To nearest whole number. "Number in parentlieses following percentage in- fested indicates the actual number of hosts examined when less than 20 were collected. were about one-half, and Ferognathiis and Pero- myscus were about one fourth as frequently in- fested. The average tick-host index for Dipodo- mys (11.2) was about twice th'at for other hosts. The tick-host index was not significantly dif- ferent for males and females of host species ex- cept for Oni/chomt/s wherein the index for fe- males was 50% higher than for males (S.lcf : 4.5 9 ). Nymphs. The highest incidence of infesta- tion occurred with Dipodomys. The tick-host index for Onychomys was about two times higher than for Perognathus and Peromyscus, but was only s'hghtly higher than Dipodomys. The tick -host index was not significantly dif- ferent for males and females of host species ex- cept for Onychomys wherein the index for males was almost three times higher than for females {7.5d:2.8 9 ). Adults. Although adults were taken from a variety of hosts, most were infrequent infesta- tions. The average incidence on Dipodomys was about 4%, whereas for Mustela and Taxidea it was almost 100%. The average tick-host in- dex was five on Mustela as compared to about two for other hosts. Seasonal Incidence Larvae. Larvae were found every month except February. Peak populations occurred in March, June, and October. Few specimens were taken in January, September, November, and December. Nymphs. Ticks of this stage were found every month except February and December. Only one nymph was itaken in January, and five in November. Highest incidence occurred in March, June, and October. Adidts. Males were taken from March into October, except in May and September, and predominantly in July and October. Females were taken from March into October, also pre- dominantly in July and October. Other Species Ixodes angustus One larva was taken from a male Spermo- philus in June. Ixodes marmotae Four larvae were taken from a male Mar- mota in May, and 11 larvae from a male Mar- mota in June. One nymph was taken from a female Spermophilus in June. Ixodes ochotonae Twenty-five larvae \vere taken from a female Spermophilus in June. Two nymphs were taken from a male Neotomci in August, and four from a female in September. Ixodes sculptus One male and three female ticks were taken from a male Taxidea in August. Ixodes sp. A number of Ixodes, mostly larvae, unidenti- fiable to species, were taken from the following hosts: Ticks of Reactor Testing Station 27 Agelaius phoeniceus, one larva in June from a male bird. Dendrocopos villosus, 17 larvae in June from a female bird. Lanius ludovicianus, three larvae and one nymph in April. Passerina amoena, one nymph in May from a male bird. Microtus montanus, two nymphs in June. Reithrodontomys megalotis, four larvae in March from a female mouse. Spermophilus townsendii, one larva in March, one in April, and 37 in June. DISCUSSION Trapping Efficiency For the past 17 years during ecological studies in the cold deserts of western North America, my associates and I have used several kinds of animal itraps and bait with various de- grees of success. Live-catch traps have captured more of the small nocturnal rodents than other traps, and in the Idaho study this also was the case. The live-catch traps caught up to four times as many animals as the break-back type. The YAW trap was twice as efficient as the Hubbard trap for harvest mice, white-footed mice, pocket mice, and kangaroo rats (nocturnal species ) . The YAW and Hubbard traps demon- strated about equal efficiency for ground squir- rels and chipmunks (diurnal species). How- ever, three times as many grasshopper mice (nocturnal) were caught in the Hubbard traps compared to the YAW traps. These relative efficiencies likely are related to bait retention, conspicuousness of the trap, and retention of animal odors which may be attractive to some species and repulsive to others. The can pit-trap, efficient for some amphibi- ans, reptiles and ground-dwelling arthropods, occasionally captured rodents. However, the numbers caught are insignificant except for Sorex and Microtus which were frequently caught in the cans. Nuttall's and pygmy cotton- tail rabbits occasionally were found in YAW traps. Ground-dwelling birds, such as the Ves- per Sparrow, Sage Thrasher, Homed Lark, and Sage Sparrow, were caught infrequently in the YAW and break-back traps. Interaction of Tick Species It is assumed that competition occurs be- tween ticks of different species relative to their simultaneous infestation of the same hosit. This was not the situation, however, for Dermacentor and Ixodes (Table 6). In most cases the per- centage of animals infested with ticks of both species simultaneously was higher than expected as demonstrated particularly on Perognathus. Onijchomijs demonstrated the expected pattern, and fewer animals were infested with both ticks than was expected. Other species combinations seldom occurred, although this may have been due to the fewer numbers of ticks collected. No Leptis was taken on which only Dermacentor was found, but 30% of these hosts were infested only with Haemaphysalis; eighteen percent were infested with both Dermacentor and Haemaphysalis. Only four hosts, each of a different species, were infested with different species of Ixodes simul- taneously. Seven other hosts representing four species possessed ticks of different genera simultaneously. Table 6. Interaction relative to occurrence of Dermacentor andersoni and Ixodes kingi on common hosts. Host and sample size D. Actual % of hosts infested with Only Only Both andersoni I. kingi ticks Expected % infestation with both ticks Dipodomys ordii (808) Eutamias minimus (398) Lepus californicus (125) Omjchomys leucogaster (63) Perognathus parvus (474) Peromyscus maniculafus (1866) Spermophilus townsendii (60) 8 63 5 5 2 4 0.5 Q.08 18 2 0.8 0.4 14 64 6 8.9 22 14 13 3 22 11 5 2.4 15 7 5 1 28 Brigham Young University Science Bulletin Ecological Distribution of Ticks Insufficient detailed analyses of ithe habitats were made to correlate the occurrence of ticks with both soil and plant types. However, the frequency of occurrence of Dermacentor and Ixodes ticks was greatest in the sage-grass as- sociations and around reservoirs (Tables 7 and 8). Where grass and sage were present in only small amounts, or where either occurred in abundance with plants of other species, ticks were not as abundant. Ticks were conspicuously absent in the grassy, sinks area of site 15. Al- though mammals occur there in moderate num- bers (Table 1), the periodic, spring flooding of this lowland area may be effective in eliminating the unattached ticks which normally await the passing of a suitable host. In at least six areas where Dermacentor and Ixodes occurred simultaneously, a significant dif- ference between their frequency indices was Table 7. Frequency index of occurrence' of Derma- centor and Ixodes in selected study areas'. Study area Dermacentor Ixodes 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 18 23 29 32 36 38 40 .08 .008 .24 0 .02 .005 .02 .18 .001 .02 .003 .001 .17 .03 .01 .75 1.0 .76 .05 .003 .03 .05 .15 .03 .04 .05 .06 .04 .07 .23 .08 « 0 .08 .04 .91 0 .14 1.0 .34 .17 'Relative to the highest populations found (in area 36 for Ixodes and area 38 for Dermacentor) indicated as 1.0. "Areas where an adequate sample of mammals and the principal hosts were examined during the known seasonal occurrence of the species of ticks. "Not applicable for reasons stated in footnote 2. noted (Table 7). In areas 3, 38 and 40, Derma- centor was greatly predominant, whereas in areas 12, 23 and 36, Ixodes was predominant. In seven of 21 areas Dermacentor was slightly more abundant than Ixodes, whereas in 13 areas Ixodes was the more abundant. Table 8. Frequency inde.\' of ticks in representa- tive plant community types and areas. Predominant plant type or area Frequency index Dermacentor Ixodes Grass .03 .145 Juniper .02 .15 Sage .39 .1 Sage-Grass .87 .67 Sage-Grass-Juniper .001 .23 Sage-Rabbitbrush .029 .058 Sage-Saltbush .18 .05 Rabbitbrush-Grass .003 .07 Rabbitbrush-Horsebrush .005 .03 Goosefoot-Winterfat .001 .06 Sand Dunes ( shifting and semi-permanent ) 0 .04 Reservoirs (semi-permanent) 0 .91 'Average taken between all areas of the specific plant type, so highest index possible will not equal 1.0. Refer to footnote 1 of Table 7. Radiaition Effects Although the evidence relating to the effects of low-level radiation on parasites and itheir hosts is circumstantial and available for only one area, it is sufficient to warrant further detailed studies. In an area where radioactive wastes were buried, fewer ticks and infested mammals were found than in a non-irradiated control area of a similar habitat. At study 13 (radioactive waste burial ground near NRTS facility EBR-I ) the frequency index was lower for Dermacentor and Ixodes than at the non-irradiated control site (area 38) 0.3 mile away (Table 9). Simi- larly, the tick-ho-st index was much lower in the irradiated area than at the control site. Table 9. Relative infestation and abundance ticks in irradiated and non-irradiated control areas. of Irradiated area Non-irradiated area Factor Dermacentor Ixodes Dermacentor Ixodes Frequency index .17 .08 Tick-host index .93 .67 1.0 .34 5.4 2.72 Ticks of Reactor Testing Station 29 SUMMARY During the period from June, 1966, to Sep- tember, 1967, 27 amphibians of two species, 446 reptiles of eight species, 561 birds of 81 species, and 4050 mammals of 26 species were examined for ectoparasites at the National Reactor Testing Station in southern Idaho. Of seven species of ticks collected, the predominant one was Ixodes kingi, found on mammals of 12 species. Next in abundance was De^-nmcentor andersoni found on birds of three species and mammals of 16 species. Haemaphysalis leporispalustris was tak- en from birds of 10 species and six spvecies of mammals. Four other species of Ixodes were taken from birds of four species and seven spe- cies of mammals. Where adequate samples were taken, the highest percentage of hosts infested with Der- macentor andersoni was 8% for birds and 29% for mammals. Haemaphysalis leporispalustris was found on as many as 53% of the birds and 29% of the mammals. Ixodes kingi infested as many as 46% of the mammals. Apparently there was little competi'tion be- tween ticks of different species relative to their simultaneous infestation of the same host. Highest populations of ticks were found in areas where both sagebrush and grass were pre- dominant. In one irradiated area the rate of infestation and population of 'ticks were signifi- cantly lower than in an adjacent, non-irradiated control area. REFERENCES Allred, D. M., D E. Beck, and C. D. Jorgensen. 1963. Biotic communities of the Nevada Test Site. Brigham Young Univ. Sci. Bui., Biol. Ser., 2(2): 1-52. Bates, R. G. 1965. Aeroradioactivity survey and area! geology of the National Reactor Testing Sta- tion area, Idaho. U. S. Atomic Energy Commission, Civil Effects Test Operations, Series CEX-59.4.10. February. McBride, R. NRTS vegetation map. National Re- actor Testing Station. Health and Safety Division, Ecology Branch. Unpublished data. U. S. Atomic Energy Commission. 1965. National Reactor Testing Station. Thumbnail Sketch. U. S. Atomic Energy Comm., Idaho Falls, Idaho. January. INFORMATION FOR CONTRIBUTORS Contributions to the Science Bulletin should be primarily monographic in nature. For the most part only manuscripts of approximately forty or more typewritten pages will be ac- cepted. Papers will be published approximately in the order that they are received, pending avail- ability of funds. Authors must arrange for finan- cing their publications, except that the Univer- sity Press and the hbrary of Brigham Young University will share the publication cost pro- portionately to their needs for copies of the article printed. In the preparation of manuscripts, authors are requested to foUow the Style Manual for Biological Journals, American Institute of Bio- logical Sciences, 2000 P Street, N.W., Washing- ton, D.C., 20006. Manuscripts must be typed on one side of the paper only, double-spaced with ample margins. Footnotes should be avoided. To facihtate review by referees, send to the editor the original manuscript and one carbon copy, together with the illustrations. Copies of the original illustrations may be submitted, but should be of quality equal to the originals. Illustrations should be referred to as figures except for materials requiring inserts of special paper, which may be called plates. Illustrations should be so designed as to fit when reduced into a one-column or fidl-page width. Special care must be taken to allow for proper reduc- tion in lettering (i.e., a 50% reduction of the figiure means also a 50% reduction in the lettering). Photographs should be of a glossy finish, unblurred, and showing sharp con- trast. Line drawings should be made with black ink on heavy white drawdng paper, blue trac- ing cloth, or blue-ruled coordinate paper. Use the same abbreviations on line drawings as in text. Line drawings must be equivalent to a professional draftman's work. Original drawings are preferable to photographs, even if they are large. Illustrations (hne drawings or photo- graphs) should be nimibered consecutively throughout the paper, and the approximate place of insertion should be indicated in the margins of the manuscript pages. Captions for illustrations should be assembled on a separate sheet, and each plate of figures must have its corresponding figm-e niunber pencilled hghtly on the back. Illustrations and cuts will be de- stroyed unless their retimi is requested when proof is returned to tlie editor. A table title should be a short, concise state- ment of what the table purports to show, and should not include information necessary to the interpretation of the table. Every column in the table should carry a head identifying the data in that column; the measm^e in which the data are given should be indicated at the head of each column. Tables should not be used when the same information can be given in a few lines of text, and should not duphcate informa- tion in text, graphs, or charts. Symbols ( asterisk, dagger, etc.) should be used to indicate foot- notes to tables, with footnotes on the same page as the table. An abstiact of less than four percent of the length of the paper should be prepared. This summary should be understandable without ref- erence to the body of the manuscript. The ab- stiact must be on sheets separate from the man- uscript. Proof should be corrected immediately on receipt and returned to the editor. Authors should leave forwarding addresses if they move from the address sent with the manuscript. Reprints should be ordered when the proof is returned. Address all manuscripts to Wilmer W. Tan- ner, Department of Zoology and Entomology, Brigham Yoimg University, Provo, Utah. I j N/)-?r^,ua3 '-'^^ COMP -w,wH 200L' Library ' Brigham Young University ^ Science Bulletin ' ^'^^ ^ NEW RECORDS AND SPECIES OF NEOTROPICAL BARK BEETLES (SCOLYTIDAE: COLEOPTERA) PART IV by STEPHEN L. WOOD ' 1 I BIOLOGICAL SERIES — VOLUME X, NUMBER 2 JUNE 1969 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Wilmer W. Tanner, Department of Zoology and Entomology Brigham Yoimg University, Provo, Utah Associate Editor: Stanley L. Welsh, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: Vernon J. Tipton, Zoology Ferron L. Anderson, Zoology J. V. Beck, Microbiology Joseph R. Muedock, Botany Wilmer W. Tanner, Chairman of the Board Ex officio Members: A. Les' Sci Ernest L. Olson, Chairman, University Publications A. Lester Allen, Dean, College of Biological and Agricultural Sciences The Brigham Young University Science Bulletin, Biological Series, pubhshes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from University Press, Brigham Yoimg University, Provo, Utah. All remittances should be made payable to Brigham Yoimg University. Orders and materials for hbrary exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 84601. Brigham Young University Science Bulletin NEW RECORDS AND SPECIES OF NEOTROPICAL BARK BEETLES (SCOLYTIDAE: COLEOPTERA) PART IV by STEPHEN L. WOOD BIOLOGICAL SERIES — VOLUME X, NUMBER 2 JUNE 1969 I I TABLE OF CON lENl S Xylechinus avarus, n. sp I Phrixosoma clusiae, n. sp 1 Phloeosinus jwlearis. n. sp 2 Chramesus gracilis, n. sp 2 Chraniesus ingen.s, n. sp 3 Cliraniesui acacicolens, n. sp 3 Chrumesus varius, n. sp 4 Chramesus secui, n. sp 4 Chramesiis unicornis, n. sp 4 Chramesus cecropiae, n. sp 5 Chramesus periosus, n. sp 5 Phloeotribus maurus, n. sp 6 Phloeolribus alavus, n. sp 6 Phloeotribus hyslrix, n. sp 7 Phloeotribus hystrix, n. sp 7 Phloeotribus opimus, n. sp 7 Phloeotribus squamatus, n. sp 7 Chaelophloeus phorodendri, n. sp H Liparlhrum americanum, n. sp *< Liparlhriim cracentis, n. sp 9 Cnemonyx nitens, n. sp 9 Cnemonyx insignis, n. sp 9 Cnemonyx opacus, n. sp ' 10 Cnemonyx gracilens, n. sp 10 Cnemonyx recavus, n. sp II Camplocerus infictelis, n. sp 11 Scolytus nodalus, n. sp 11 Scolylus cristatus, n. sp 12 Microborus limatus, n. sp 13 Microborus ambitus, n. sp 13 Scolytodes clusiae. n. sp 14 Scolytodes volcanus, n. sp 14 Scolytodes pan'utus, n. sp 14 Scolytodes acares, n. sp 15 Scolytodes irazuensis, n. ap 15 Scolytodes cecropicolens. n. sp 16 Scolytodes plioebeae, n. sp 16 Scolytodes plumeriae , n. sp 17 Scolytodes cecropiavorus, n. sp 17 Scolytodes cecropiavorus acuminatus, n. siibsp 17 Scolytodes cecropiavorus punctifer, n. siibsp 18 Scolytodes veniistus, n. sp IS Scolytodes pseudopiceus, n. sp 1*^ Scolytodes ochroinae. n. sp 19 Scolytodes cedreUie. ii. sp 19 Scolyttnles immaiiis. n. sp 19 Scolytodes iiliii, n. sp 20 Scolytodes eriucopliilus, n. sp 20 Scolytodes inarginatus, n. sp 20 Scolytoiles iiiarginotiis, n. sp 21 Scolytodes plicalus. n. sp 21 Scolytodes pubescens, n. sp 21 Scolytodes impressus, n. sp 22 Scolytodes exiguus, n. sp 22 Scolytodes pumilus, n. sp 23 Pseudothysanoes acacicolens, n. sp 23 Pseudothysanoes vallatus, n. sp 24 Pseudothysanoes vesculus, n. sp 24 Pseudothysanoes mancus, n. sp 24 Pseudotliysanoes viscivorus, n. sp 25 Pseudothysanoes amassius, n. sp 25 Pseudothysanoes viscicolens, n. sp 25 Pseudothysanoes funnis, n. sp 26 Pseudothysanoes fumalus, n. sp 26 Pseudothysanoes verdicus, n. sp 27 Pseudothysanoes funebris, n. sp 27 Pseudothysanoes peniculus, n. sp 28 Pseudothysanoes bullatus, n. sp 28 Pseudothysanoes plumalis, n. sp 29 Pseudothysanoes coracinus, n. sp 29 Pseudothysanoes crassinis. n. sp 29 Cryplulocleplus quercinus. n. sp 30 Cryptulocleplus arbuti, n. sp 30 Cryptulocleptus carilus, n. sp 30 Cryptulocleplus pumilus, n. sp 31 Cryptulocleplus mendicus, n. sp 31 Cryptulocleplus acares, n. sp 32 Cryptulocleplus aquilus, n. sp 32 Cryplulocleptus mirus, n. sp 32 Cryptulocleplus obesus, n. sp ; 33 Cryptulocleplus excavatus, n. sp 33 Cryptulocleplus spicatus, n. sp 33 Cryptulocleplus subulatus, n. sp 34 Cryplulocleptus mttcronatus, n. sp 34 Thysanoes neotropicalis, n. sp 35 Thysanoes subsulcatus, n. sp 35 Thysanoes epicharis, n. sp 35 Thysanoes adonis, n. sp 36 Micracisella monadis. n. sp 36 Micracisella sciluta. n. sp 37 Micracisella nitidula. n. sp 37 Micracisella nigrellu. n. sp 37 Micracisella si mi lis. n. sp 38 Micracisella vescula. n. sp 38 Micracisella squamatula, n. sp 38 Micracisella divaricalii, ii. sp 39 Micracis costariceiisis, n. sp 39 Micracis carinulus, n. sp 40 Micracis tribulatus, ii. sp 40 Micracis lepidus, n, sp 41 Micracis lignicolus, n. sp. . . . v 41 Micracis detentus, n. sp 42 Micracis unicornis, n. sp 42 Micracis inimicus, n. sp 42 Micracis festivus, n. sp 43 Hylocurus alternus, n. sp 43 Hylocurus aberrans, n. sp 44 Hylocurus incnniptus, n. sp 44 Hylocurus microcornis, n. sp 45 Phloeocleptus plagialus, n. sp 45 Phloeocleptus nanulus, n. sp 45 Phloeocleptus pannis, n. sp 46 LIST OF ILLUSTRATIONS Figure 1. Scolytus nodatus, n. sp 12 2. Scolytus cristalus, n. sp 13 3. Hylocurus incomptus, n. sp 44 NEW RECORDS AND SPECIES OF NEOTROPICAL BARK BEETLES (SCOLYTIDAE: COLEOPTERA) PART IVi BY Stephen L. Wood^ On the following pages 108 species and two subspecies of bark beetles (Scolytidae) are described as new to science. It was intended that most of these would be named in a synopsis of the Scolytidae of Costa Rica; however, the volume of undescribed material proved to be so large and the manuscript treating it so bulky that its completion became impractical. That manuscript, with its keys, host and geographical distribiuions, and biological notes is being expanded into a treatment of the entire Ameri- can continent north of Panama, In order to stabilize new names for that project, they are validated below. The new species represent the following genera: Xylechinus (I), Phrixosoma (1), Phloe- osiniis (1), Chramesiis (8), Phloeolribus (5), Chaetuphloeus (1), Liparihrum (2), Cnejnonyx (5), Camptocenis (1), Scaly Ins (2), Microborus (2), Scolytodes (22 with 2 subspecies), Pseudu- thysanoes (16), Cryptulocleptus (13), Thysanoes (4), Micracisella (8), Micmcis (9), Hylocurus (4), Phloeocleplus (3). Ihey are from the fol- lowing countries: Mexico (49), Guatemala (9), Honduras (7), Costa Rica (49), Panama (5); seven species occurred in two countries, and two species occurred in three countries. Except as noted below, all type series were tentatively deposited in my personal collection presently housed at the Brigham Young Uvi- versity, in order to facilitate preparation of a monograph of the Scolytidae of the western hemisphere. Upon conclusion of that mono- graph, paratypes will be distributed, in so far as possible, to at least two European and two other American collections. The British Museum (Natural History) has already received para- types of many species, Xylechinus avarus, n. sp. The large body size, the stout form, the ele- vated declivital interspaces I and 3 on the elytra, the fine ground vestiture. and the fine, low frontal carina serve to distinguish this species. Of the species known to me it is perhaps more closely allied io full gin osus Blandford, although the relationship is remote. Female. — Length 3.2 mm. (paratypes 2.4-3.3 mm.), 2.1 times as long as wide; body color brown, usually covered by an incustation. Frons convex, with a feeble transverse impression just below upper level of eyes, a low acutely elevated median carina extending from this impression to episto- mal margin; entire surface coarsely reticulate, punctures fine, inconspicuous; vestiture short sparse inconspicuous. Pronotum 0.93 times as long as wide; sides widest and almost parallel on middle half, strongly constricted one-fourth length from anterior margin, rather broadly rounded in front; surface finely grantdate and with rather sparse, round, small vulcanate pinictures; vesti- ture consisting of short, rath-er stout, moderately abun- dant bristles. Elytra 1.2 times as long as wide. 1.8 times as long as pronotum; sides almost straight, very slightly wider at declivital base, narrowly rounded behind; striae im- pressed, the pimctures deeply, not sharply impressed; interstriae about one and one-half times as wide as striae, irregular, the punctures fine, abundant, con- fused, with a median row of larger, granulate punctures, crenulate toward base. Declivity convex, rather steep; striae impressed; interspace 2 impressed, strongly nar- rowed, ending before apex, I. 3, and 9 elevated, about half as high as wide, 3 and 9 join apically. the elevation continued to I. granules not as large as on disc. Vestiture of short, rather abundant, stout, hairlike setae, and interspacial rows of long bristles, spacing between bristles and between rows equal to length of a bristle. Mate. — Similar to female except frons flattened on lower half; and ground vestiture evidently more abundant. Type Locality. — Volcan Irazu, Cartago Prov,, Costa Rica, Hosl. — Oreopanax nubigenus. Type Material. — The male holotype, female allotype, and 205 paratypes were collected at the type locatlity on October 26, 1963, at an eleva- tion of 2,300 m,, by S. L. Wood, from the fallen epiphytic host that was about 10 cm. in diameter. The galleries were similar to those oi fuliginusus. The holotype, allotype antl p;iratype are in my collection. Phrixosoma clusiae, n. sp. This species is very closely related to minor Wood, but it may be distinguished by the larger size, by the much more strongly impressed female vertex, by having the interstrial granules three- ranked almost to elytral base, by the compara- tively wider discal interstriae, and by the less definitely granulate anterior slope of the pro- notum. Female. — Length 2.5 mm. (paratypes 1.9-2.5 nnn.), 1.9 times as long as wide; color black. Frons broadly convex from epistomal margin to upper level of eyes; rather strongly, transversely, arcuate- ly impressed from just above level of eyes to vertex, upper margin of this groove abruptly angled; lateral angles of groove produced laterally causing a pro- 'Part of the Tieki work thai led to the discovery of these insects was sponsored by the National Science Foundation. ^Department of Zoology and Entomology, Brigham Young University, Provo. Utah. Scolytoidea contribution number 38. Brigham Young University Science Bulletin nounced sinuation in line where pubescent and non- pubescenl areas meet; vestitiire hairlike, short, rather abundant, extending to vertex. Pronoliim 0.80 tiines as long as wide; widest at base, sides uniforndy. convergently arcuate to weak constriction just before the rather narrowly rounded anterior margin; anterior third very minutely asperate, asperities on margin somewhat larger; surface rather closely marked by low. subluberculate. short, irregular rugae (remnants of interspaces between obsolete punc- tures), depressed areas reticulate; vestiture consisting of very fine, short hair. Elytra 1.2 times as long as wide. 1.7 times as long as pronotum; outline as in obesa; narrowly impressed, the punctures small, distinct; interstriae almost three tiines as wide as striae, reticulate, granules rather large, rounded, three-ranked almost to base except on 2. the maiginal rows of granules becoming .smaller by middle of disc and obsolete by upper fourth of declivity. Declivity and vestiture as in minor. Male. — Similar to female except frontal carina absent. vertex and anterior margin of pronotum not modified, and asperites on anterior third of pronotum almost absolete. Type Locality. — Zamorano, Moia^aii. Hon- duras. Hos t . — Cliisia flava . Type Material. — The female holotype, male allotype, and ten paratypes were collected at the type locality on April 18, 1964, 700 m. ele- vation. No. 561, in branches and bole of Cliisia flava. This monogamous species constructs biramous, transverse egg galleries in the cam- bium region essentially as in minor. The holotype, allotype, and paratypes are in my collection. Phloeosinus palearis, n. sp. This species is closely allied to serralu.s LeConte, but it may be distinguished by the smaller size, by the numerous minute points and/or fine lines in the interspaces between punctures on tiie pronotal disc, by the less numerous, smaller submarginal crenulations at the elytral bases, and by the less strongly compressed teeth on the male elytral declivity, the anterior profile of each tooth usually evenly arcuate (abrup;'y angled in serratus). Mule. — Length 2.4 nun. (paratypes 1.9-2..') mm.). 1.9.5 limes as long as wide; color very dark brown. Fions feebly convex, almost flat, the lower fointh gradually ascending to epistonial margin; lower hall marked by a fine, low median carina; surface smooth with rather fine, deep, close puuctines, some sub- granulate in area median to upper half of eye; vestiture fine, moderately long, rather abuuckiiu. \'estinne on antennal club short, more abundaut than in allied species, Pronotum 0.82 times as long as wide; widest near ba.se. the sides strongly, convergently arcuate to moderate constriction just before broadly rounded anterior mar gin; surface smooth, with some minute points and lines in interspaces between the modeiately large deep punctines; interspaces one-fourth to equal in width to diameter of a ptuictuie. Vestiture fine, rather short, modeiately abundant. Elytra 1.2 times as long as wide. 1.6 times as long as pronotum; basal margins armed by 12 crenulations on interstriae 2-9; striae narrowly impressed, the small punctures somewhat obscurely indicated; interstriae about three times as wide as striae, feebly convex, the surface smooth but irregular, the punctures modeiately coarse, close, confused, their anterior margins usually elevated, some rather coar.sely crenulate at base. Decli- vity convex, rather steep; striae less strongly impressed; interstriae 1 and 3 moderately, 5, 7, 8 less strongly elevated, these and 6 armed by rather coarse teeth, those on 3 and particularly on 1 larger, 2 bearing two teeth near apex, 2 and 4 flat, finely puncliaeil; inter- striae 1 bearing eight teeth, each only slightly compressed and usually with anterior profile evenly arcuate, % bearing 13 teeth with none of them compressed. Vestiture scanty, but stout, some setae almost scalelike. Female. — Similar to male except frons more stongly convex and more extensively granulate; pronotal lines and points much more abundant; a few elytral crenu- lations extend almost to declivity; declivital teeth smaller with interstriae 2 and 4 armed by at least a few small teeth; declivital vestiture largely of scales each only slightly longer than wide. Type Locality. — Sixteen km. east Chiapa de Corzo, Chiapas, Mexico. Host. — Juniperus sp. Type Material. — The male holotype, iemale allotype, and M paratypes were collected at the type locality on June 17, 1964, at an elevation of about 2.300 m., No. 710. fiom a small Juni- perus branch. Ihe holotype, allotype ami paratypes are in my collection. Chrumesus gracilis, n. sp. This species is in the subgenus Prochramesus and is allied to annectans Wood, but may be distinguished by the larger size, by the much more finely, shallowly ptinctured pronotum, by the smaller, shallow strial punctures, and by other characters. Mate. — Length 2.5 mm. (paratypes 2.2-2.7 mm.), 2.2.') times as long as wide; color dark brown to black, with white vestiture. Frons rather deeply, concavely impressed from upper level of eyes to epistoma. the lateral margins rounded, bearing mesad of margin just above antennal bases, a pair of large squared pro'cesses as in annectayu; surface coarsely reticulate with a few fine, obscure punctures; vestiture fine. long, rather sparse. Antennal club acutely pointed. Pronotum 0,9 times as long as wide; widest at base, sides weakly arcuate and converging slightly on basal two thirds then rather strongly constricted before the broadly rounded anterior margin; surface reticulate, with fine, rather widely separated pimctures. the punc- tures indistinctly vulcanate or suba.sperate in lateral areas; vestiture consisting of stout semirccumbent bristles. Elytra \.b times as long as wide. 1.9 times as long as pronotum; sides straight and parallel on basal two- thirds, rather broadly loimded behind, striae not im- i New Neotropical Scoi.vtidae pressed, the punctures small, round, shallow, distinct; interstriae about three times as wide as striae, almost flat, smooth and shining with rather abundant, minute, setiferous. confused punctures and median rows of fine setiferous gianules. Declivity steep, convex; striae weak- ly impressed, the interspaces feebly convex. Vestiture consisting of rather abinidant, short, subpliimose scales and interstrial rows of somewhat longer erect bristles, each bristle about two-thirds as long as distance between rows. Female. — Similar to male except frons very shallowly impressed, the tubercles somewhat smaller; pronotal tubercles distinctly larger. Type Locality. — University of Costa Rica Campus, San Jose, Costa Rica. Host. — Bambusa vulgaris. Type Material. — The male holotype, female allotype, and 83 paratypes were collected at the type locality on September 12, 1963, at an elevation of about 1,300 m., by S. L. Wood, from dead and dying stuinps of Japanese bamboo which were 10 cm. or more in diameter. The diagonal, biramous, egg galleries were jtist under the surface of the wood. Larval mines were parallel to the grain of the wood. The holotype, allotype, and paratypes are in my collection. Chramesus ingens, n. sp. Among the species known to me this one is iniique in having the antennal bases compara- tively close together and higher than usual: in having the elytral vestiture fine and of uniform length; and in having the male frons not ex- cavated, but the lateral margins elevated with the raised tubercle at a point well above the antennal insertion. Male. — Length 2.7 mm. (paratypes 2.4-2.8 mm.), 1.6 times as long as wide; color dark brown or black. Frons impressed from upper level of eyes to epistoma with a low elevation between antennal bases, lateral margins elevated on middle third with a rounded tubercle just above antennal base; surface reticulate, with very fine punctures at sides and above; antennal bases closer together than eyes, distance from epistomal margin (omitting lobe) to antennal bases 1.6 times greatest width of eye; frons subglabrous. Antennal scape with small tuft of hair. Pronotum 0.8 times as long as wide; base strongly bisinuate, basal angles broadly rounded, sides strongly arcuate, widest just behind middle, moderately con- stricted just behind the broadly rounded anterior mar- gin; entire surface reticulate and with small, sparse, finely subasperate punctures, finer in posteromedian area; glabrous. Elytra 1.03 times as long as wide, 1.5 times as long as pronotum; sides straight and parallel on basal half, broadly rounded behind; striae narrowly impressed, the punctures small, distinctly impressed; interstriae about three times as wide as striae, slightly convex almost smooth with numerous, confused, small rounded seti- ferous gianules. Declivity convex, moderately steep; about as on disc. Vestiture consisting of fine, erect, confused, hairlike setae, all of equal length, each shorter than a distance equal to width of an interspace (about two thirds as long). Female. — Similar to male except frontal margin feebly elevated and not armed; pronotal punctures almost entirely devoid of small asperities. Type Locality. — Tapanti, Cartage Prov., Costa Rica. Host. — Serjania mexicana? (probable error in identification). Type Material. — The male holotype, female allotype, and 270 paratypes were collected at the type locality on October 24. 1963, at an elevation of about 1,300 m., by S. L. Wood, from a vine having three leaflets per leaf. The host was identified as Serjania mexicana, however, the tissues were very moist, soft and non-woody and turned black with age, suggesting an error in identification. This species was also collected at Puerto Viejo, Heredia Prov., and at Guapiles, Limon Prov., in humid lowland areas, but these specimens were not included in the type series. The holotype, allotype, and paratypes are in my collection. Chramesus acacicolens, n. sp. Similar to crenatus Wood, but distinguished by the deeper, wider strial punctures, by the much shorter, wider, erect interstrial scales, by the small tubercles of elytral interspaces, and by other characters. Male. — Length 1.8 mm. (paratypes 1.5-1.8 mm.). 1.65 times as long as wide; color light brown. Frons broadly, concavely excavated from upper level of eyes to epistoma, deepest point near middle, subtoveate; lateral margins acute below, armed by a pair of tubercles just below level of antennal insertion; surface coarsely reticulate, a few miniite punctures scarcely visible; vestiture short, inconspicuous. Pronotum 0.7.3 times as long as wide; median basal area produced slightly into scutellar notch, sides widest near base, rather strongly arcuate, constricted just behind the broadly rounded anterior margin; surface minutely rugulose, dull, rather finely, shallowly punctured, the punctmes becoming finely asperate toward anterior and lateral areas; vestiture consisting of short, stout bristles. Elytra 1.07 times as long as wide. 1.6 times as long as pronotum; sides straight and parallel on basal half, broadly rounded behind; striae impressed, the punc- tures rather laige and deep; interstriae about twice as wide as striae, convex, with small, abundant, squami- ferous punctures and median rows of tubercles that decease in size posteriorly. Dccli\ity convex, moderate- ly steep; as on (lisc except tubercles not evident. Vesti- ture consisting of abundant short scales and interstrial rows of erect scales slightly more than twice as long as ground vestiture. each large scale about three times as long as wide. Female. — Similar to inale except frons weakly con- vex, transversely impressed just above epistoma, minute punctures more evident; pronotal asperities evident only at anterolateral angles. Type Locality. — Finca La Pacifica, 4 km. Bricham Young University Science Bi lletin northwest of Caiias, Guanacaste Prov., Costa Rica. Host. — Acacia sp. Type Material. — The male holotype, female allotype, and 80 paratypes were collected at the type locality on July 13, 1966, at an elevation of 50 m., by S. L. Wood, from the cambium region of branches and twigs of the host tree less than 2 cm. in diameter. The holotype. allotype, and paratypes are in my collection. Chramesus variiis, n. sp. This is the only species of Chramesus known to me in which the frons is concave in both sexes and the epistoma is armed in both sexes by a large median tubercle; the variegated color pattern of the scales is also unique in the genus. Male. — Length 1.5 mm. (paratypes 1.41.7 mm.) 1.7 times as long as wide: color brown with light colored scales on basal fourth and lateral margins of pionotum and on interstriae 1 and lateral areas of elytral disc. dark scales predominate elsewhere. Frons deeply, broadly, concave from eye to eye from well above eyes to epistoma; excavated area reticulate- granulate; lateral margin acute only at level of antennal bases, unarmed; epistomal margin bearing a large, pointed, median tubercle; vestiture consisting of mod- erately abundant, stout setae. Scape bearing a conspicuous tuft of long, yellow, subplumose setae; club rather small for this genus, its apex not pointed. Pionotum 0.82 times as long as wide; widest on basal third, the sides strongly, convergently arcuate to- ward the broadly rounded anterior margin: anterolateral areas with a few fine asperities; surface coarsely reticulate, the punctines rather fine, shallow, moderately close, many of them sub\iilcanate on posterior area: basal margin bisinuate: vestiture stout, short, es.sentially scale- like. Elytra 1.14 times as long as wide, 1.6 times as long as prdnotum: bases armed by 11 marginal crenulations on interstriae 2-9; scutellum not visible in usual sense; striae weakly impressed, stronger toward declivity, the punctures small, impressed; interstriae twice as wide as striae, convex, the surface evidently superficially reticu- late, the punctures very fine, usually obscured by setae. Declivity convex steep; striae slightly deeper, interstriae slightly narrower than on disc. Vestiture consisting of moderately abundant, stout, almost scalelike, erect setae, and rows of erect scales each about twice as long as ground vestiture and about four or five times as long as wide. Female. — Similar to male except frons more narrow- ly, shallowly concave with median tubercle a little smaller: pronotal crenulations larger and more numerous. Type Locality. — Twenty-seven km. north of Ixmiquilpan, Hidalgo, Mexico. Host. — Acacia sp. Type Material. — The male holotype, female allotype, and 12 paratypes were collected at the type locality on July 10, 1967, elevation 1,900 m., No. 188, in an Acacia branch about 4 cm. in diameter, by S. L. Wood. The holotype, allotype, and paratypes are in my collection. Chramesus secus, n. sp. This species is exceedingly similar to i/ariits Wood, described above, but differs by the larger size, by the uniformly light color of the vestiture, by the more finely sculptured pionotum, and by the shorter, more scalelike vestiture. This pos- sibly could be a subspecies oilvariiis. Male. — Length 1.7 mm., 1.7 times as long as wide; color brown with white vestiture on pronotum and interstriae 1. tan elsewhere. Exactly as in varius except as noted above and also setae on all margins of pronotum shorter and stouter: median extension of ba.sal margin of pronotum less pronounced; elytral vestiture appearing more abundant and each seta conspicuously wider. Type Locality. — Laguna Santa Maria, Naya- rit, Mexico. Type Material. — The unique male holotype was taken with a series of the following species on July 6, 1965, near 1,000 m. elevation. No. 201, by S. L. Wood. The host was a large woody vine (liana) having the larger stems square in cross section. The holotype is in my collection. Chramesus unicornis, n. sp. This species is rather closely allied to varius Wood, described above, but it is easily distin- guished by the less extensive male frontal im- pression, by the smaller tuft of setae on the scape, by the subquadrate pronotum, by the more strongly impressed striae and sirial punc- tures, and by the uniformly light colored vesti- tiae. Male. — Length 1.4 mm. (paratypes 1.3-1.6 nrin.), 2.0 times as long as wide, color medium to light brown. Frons deeply excavated from eye to eye from upper level of eyes to epistoma; lateral margins acute from just above level of antennal in.sertion to epistomal mar- gin: surface rather coarsely reticulate above, becoming obscurely reticulate and shining below, armed by a large, pointed, median tubercle immediately above epistomal margin; vestiture sparse, inconspicuous. Anten- nal scape bearing a small tuft of fewer than a dozen long, yellow setae; club small for this genus. Pronotum 0.83 times as long as wiile; widest at base, the sides feebly arcuate and converging slightly on more than basal two-thirds, then rather abruptly constricted just before the broadly rounded anterior margin: surface rather coarsely reticulate, the punc- tures very close, rather shallow, irregular in outline, anterolateral asperities obscure. Vestiture consisting of short, rather abundant, stout, almost scalelike setae. Elytra 1.2 times as long as wide, 1.5 times as long as pronotum: sides straight and subparallel on basal two- thirds to declivital base, then rather broadly rounded behind; basal margins armed by 14 crenidations; striae slightly impressed, the large punctures deeply impressed; interstriae as wide as striae, convex, each bearing a I New Neotropical Scoi.^ tidae row of fine, poiiued tubercles. Declivity moderately steep, convex, essentially as on disc except area at junction of interstriae 3 and 9 slightly elevated. Vestiture con- sisting of moderately abundant, small, broad scales, and ro(vs of erect interstrial scales about three times as long as ground cover, each half as long as distance between rows and almost equal in length to distance between erect scales within a row. about four times as long as wide: not longer on declivity. Female. — .Similar to male except frons coarsely reticulate to epistoma, the median absent and pronotal asperities slightly larger. Type Locality rit, Mexico. convex, tubeicle -Laguna Santa Man'a, Naya- Type Material. — The male holotype, female allotype, and 76 paratypes were collected at the type locality on July 6, 1965, about 1,000 m. elevation, No. 201, from the phloem tissues of the host, by S. L. Wood. The host was a large woody vine (liana) about 7 cm. in diameter, having the cross-section of the larger stems square. Four paratypes were taken 2.5 km. west of Armaria, Colima, Mexico, on June 30, 1965, No. 143, from the same host, by S. L Wood. The holotype, allotype, and paratypes are in my collection. Chramesus cecropiae, n. sp. In this unique species the male frons is not excavated, but more nearly protubrant at a level just above the antennal bases; in addition, the elytral ground vestiture is totally absent, a row of stout bristles marks each interstriae. Ma/f.— Length 1.25 mm. (paratypes 1.3 mm.). 1.7 times as long as wide: color rather dark brown, the elytra somewhat lighter. Frons longitudinally flattened, transversely, weakly convex on upper third, then ascending rather abruptly below to an indefinite, strongly arcuate line between antennal ba.ses, its median point well above (dorsad) its lateral extiemities; lower two-thirds almost flat, the lateral margins below very slightly elevated: surface above coarsely reticulate, lower area transversely strigose; vestiture consisting of a few hairlike setae on lateral areas on lower half. Antennal club moderately large for the genus. Pronotum 0.82 times as long as wide: widest on basal third, sides arcuately converging to a slight con- striction just before the rather bioadly rounded anterior margin; surface shining, subreticulate, with scattered, rather abundant tubercles of variable size on anterior three-fourths, some of those on anterolateral area finely asperate; a few large, obscure punctures evident toward base, \estiture rather sparse, consisting of a few stout setae over entire surface. Elytra 1.1 times as long as wide, 1.4 times as long as pronotum; elytral bases armed by about 12 cienula- tions; striae feebly if at all impiessed, the punctures large, moderately deep, distinct; interstriae very slightly wider than striae, very slightly inegular, brightly shining, each bearing a row of small, pointed, setiferous tubercles. Declivity rather steep, convex, sculptured essentially as on disc. \'estiture consisting of lows of interstrial bristles, each bristle somewhat flattened toward its apex and equal in length to distances between bristles within and between rows: very minute strial hairs also evident. Female. — Similar to male e.xcept frons more evenly convex, coarsely reticulate throughout, with a median fovea on upper half; pronotal asperities larger, the tubercles largely absent but replaced by shallow, rather obscure punctures. Type Locality. — Four km. southwest of Rin- con, Osa Peninsula, Puntarenas Prov., Costa Rica. Host. — Cecropia sp. Type Material. — The male holotype, female allotype, and two paratypes were collected at the type locality between Februai7 21 and 28, 1967, i)y D. D. .Sliwa, from the fallen leaf petiole of a Cecropia tree. The holotype, allotype, and paratypes are in my collection. Chramesus periosits, n. sp. Superficially this species resembles vastus Wood more closely than other species known to me, although its true affinities may lie nearer to crenatus Wood which shared the same host material. From vastus it differs by the more closely, deeply punctured pronotum, by the less strongly iinpressed striae, by the scalelike erect interstrial bristles, and in the male by the frontal impression extending above the eyes. Ma/e.— Length 2.2 mm. (paratypes 2.0-2..') mm.), 1.4 times as long as wide, color brown. Frons broadly concave from epistomal margin to just above upper level of eyes: lateral margins acutely elevated from just below upper level of eyes to epistomal margin, bearing a pointed tubercle at level of antennal insertion; surface reticulate except on siriooth epistomal margin, the punctures small, shallow, obscure: vestiture fine, inconspicuous. Antennae as in related species. Pronotum 0.73 times as long as wide: posterolateral angles one-third pronotum length from base, median area produced posteriorly: sides widest at posterolateral angles, strongly, arcuately convergent to moderate con- striction just before the rather broadly rounded anterior margin; suigace finely reticulate, with fine, moderately deep, rather close punctures, about half of those in central area finely asperate, the asperities larger toward anterolateral angles; vestiture consisting of stout, rather short, recumbent bristles and similar, very fine hair. Elytra 0.90 times as long as wide, 1.4 times as long as pronotum: anterior margins armed by low basally fused crenulalions that merge lateral to striae 4 into a continuous costa; striae moderately impressed, the punctures small, moderately deep, rather widely spaced; interstriae about twice as wide as striae, weakly convex, a median row of fine, nongranulate punctures and con- fused minute punctures on each interstriae. Declivity rather steep, convex; sculpture essentially as on disc. Vestiture consisting of ground covering of small, non- overlapping, interstrial scales each longer than wide, and median rows of erect interstrial bristles, each twice as long as ground vestiture and four to five times as long as wide and spaced within a row by slightly more than length of a bristle. Female. — Similar to male except frons not impressed, almost flat; lateral asperities on pronotum coarsely de- Bricham VoLiNf, University Science Bulletin veloped; each interstriae bearing a iiniseriate row of coarse, narrow tubercles, each ahiiost as high as wide. Type Locality. — Five km. west of Jaltipan, Vera Cruz, Mexico. Type Material. — The male holotype, female allotype, and 56 paratypes were collected at the type locality on June 25, 1967, 50 m.. No. 99, by S. L. Wood, from tunnels in the wood of branches 3-8 cm. in diameter. The host tree, awaiting identification, was also infested by Chrarnesus crenalits and Scolytus propingui Blandford. The holotype, allotype, and paratypes are in my collection. Phloeotribus maitrus, n. sp. This species resembles sulcifrons Chapuis and may be rather closely related; it is easily distin- guished, however, by the large tuft of hair on the male scape, by the much coarser asperities on the anterolateral angles ot the pronotum, by the coarser sculpture of pronotum and elytra, with some interstriae bearing only one row of inter- strial crenulations, and by the median row of coarse setae on each of the interstriae. Male. — Length ,S.O mm. (paratypes 2.3-3.1 mm.). 1.7 times as long as wide, color very dark brown to black. Frons similar to but much narrower than in sulci- frons; sculpture similar; vestiture above antennal bases much coarser than in sulcifrons. Scape bearing a rather large, dense tuft of dark hair. Pronotum outline as in sulcifrons; surface less uni- formly, more deeply punctured, some punctures toward base much larger than in sulcifrons; anterolateral asperi- ties about twice as large as in sulcifrons, the vestiture at lateral margins coarser and longer. Elytra 1.03 times as long as wide; striae strongly impressed, the punctures clearly indicated; interstriae distinctly less than twice as wide as striae, the punc- tures subcrenulate on 2 and 3 to declivity, less distinctly so on 4 to 6, the crenulations uniseriate except somewhat biseriate at base of 2; interstriae narrower and more con vex toward declivity. Declivity convex, moderately steep; interstriae narrower than on disc, each with a median, subserrate row of low tubercles. Vestiture consisting of a row of short, suberect bristles on each interstriae; each bristle not longer than a third the distance between rows; largely abraded on basal half; some fine hair also on sutural interstriae. Female. — Similar to male except frons convex, foveate just above le\el of antennal insertion; tuft of hair on scape absent. Type Locality. — Near Rincon, Osa Penin- sula, Puntarenas Prov., Costa Rica. Host. — Ficus sp. Type Material. — The male holotype, female allotype, and 30 paratypes were collected at the type locality on August 11, 1966, near 30 m. elevation, No. 63, from a Ficus limb 15 cm. in diameter, by .S. L. Wood. The holotype, allotype, and paratypes are in my collection. Phloeotribus atavus, n. sp. The large size, sculpture of the pronotum, and vestiture of pronotum and elytra serve to distinguish this species from all other repre- sentatives of the genus. Female. — Length 3.6 mm. (paratypes 3.3-4.0 mm.), 2.0 times as long as wide; color very dark brown. Frons shallowly, concavely impressed on lower two- thirds, the impre,ssion ending above in a small deep median fovea; lateral margins below antennal bases acute; surface rather coarsely reticluate above fovea, subreticulate to smooth below, the punctures small, deep, close; vestiture inconspicuous. Antennal club with segment 1 three times as wide as long, about as long as scape. Pronotum 0.96 times as long as wide; widest at or just behind middle, sides rather strongly arcuate, con- verging very slightly anteriorly, rather strongly constricted just behind the rather broadly rounded anterior margin; surface completely devoid of asperities, reticulate, the punctures rather small, very close, deep; a faint median line weakly elevated; vestiture consisting of inconspicuous, very fine, short hair. Elytra 1.4 times as long as wide. 1.8 times as long as pronotum; sides straight and parallel on basal two- thirds, broadly rounded and with apex rather narrowly extended on median third behind; striae narrowly im- pressed, the punctures rather obscure; interstriae twice as wide as striae, crenulate on basal half, some crenulations as wide as interspace, the elevations becoming narrower and uniseriate behind, some fine punctures also presenl; shining. Declivity rather sleep, convex; striae impressed: interstriae convex. 1 slightly elevated on lower half, apical costal margin and posterior part of 9 also slightly elevated, each interspace with a row of small tubercles. Vestiture consisting of moderately sparse, small scales on posterior half of disc and on decli\ity. and sparae median interstrial rows of fine long hair, longest hairs equal in length to distance between rows and between adjacent hairs within a row. Male. — Similar to female except frontal impression verv slightly deeper and extending to upper level of eyes, the punctures larger, shallower; interspaces 1 and 2 on elytral declivity finely punctured and almost devoid of tubercles. Type Locality. — Laguna Volcan Poas, Here- dia Prov., Costa Rica. Hosts. — Cliisia sp. (type), and Pudocarpus oleifolius (paratype). Type Material. — The female holotype, male allotype, and 31 paratypes were collected at the type locality on October 6, 1963, at an elevation of about 2,700 m., by .S. L. Wood, from a limb of CAusia about 25 cm in diameter. Three para- types bear the same data but were taken July 14, 1963, from a Podocarpus root buttress. The egg tunnels were biramous, transverse, and almost entirely in the phloem. The species is agrcssive; some specimens were attacking healthy living tissue, and it appeared to have killed several Clusia trees in the area. The holotype, allotype, and paratypes are in my collection. 1 New Neotropical Scolytidae Phloeotribiis hystrix, n. sp. This species evidently is allied to discrepans Blandford, although not closely related. It is distinguished by the more deeply striate elytra, by the more nearly scalelike vestiture, and by the declivital armature. Male. — Length 1.7 mm. (paiatypes 1.7-2.0 mm.), 2.1 times as long as wide; color brown. Frons very shallowly concave below upper level of eyes, sides weakly elevated and armed at level of aniennal insertion by a pair of small tubercles; surface coarsely reticulate, the punctures small, obscure; vestiture sparse, moderately long, coarse. Anlcnual club with segment 1 three times as wide as long. Pronotum 0.9 times as long as wide; widest on ba.sal half, the sides weakly arcuate, almost parallel on basal half, then converging to constriction just behinil broadly rounded anterior margin; surface coarsely. deeply. clo,sely. irregularly punctured, anterior and lateral areas subgranulate or even very finely a.sperate; vestiture short, stout, abundant, almost scalelike. Elytra 1.4 times as long as wide, 1.7 limes as long as pronolum; sides almost straight and parallel on basal three-fourths, obtusely angulate behind, with lateral declivital spines interrupting the profile; striae sharpiv impressed, the punctures large, deep, distinct; interstriae distinctly narrower than striae, irregular, each with a median row of fine setiferous granules and additional subgranulate setiferous punctures on each side. Declivity steep, convex: interspace 1 weakly elevated. 5 with two. 7 with three. 9 with five large pointed tubercles, the posterior most one on each interspace longer, 9 elevated slightly to final spine; costal margin near apex slightly elevated. \'estiture consisting of rather abundant inter- strial subsquamose bristles, short except median low twice as long, erect; equally abundant on disc and declivity. Female. — Similar to male except Irons weakly con vex. unarmed; pronotum more nearly asperate in lateral areas. Type Locality. — Rio Damitas in the Dota Mountains, San Jose Prov., Costa Rica. , Type Material. — The male holotype, female allotype, and 20 paratypes were collected at the type locality on February 18, 1964, at an eleva- tion of about 250 m., by S. L. Wood, from the phloein tissues in a stump of a broken uniden- tified tree 15 an. in diameter. The holotype, allotype, and paratypes arc in my collection. Phloeotribus opimiis, n. sp. As in demessus Blandford, except smaller, the basal margin of the pronotum straight, and the discal interstriae smooth and shining except for the uniseriate granules. Male. — Length 1.4 mm. (paratypes 1.4-I.S mm.). 1.8 times as long as wide; color dark brown, almost black, with light vestiture. Frons deeply excavated from uppei le\el of eyes to epistoma, deepest near center, the lateral margins acute fiom middle to antennal bases; surface rather coarsely reticidate; vestiture largely confined to marginal areas. Antennal scape long, ornamented by a tuft of long yellowish hair. Segment 1 of antennal club three times as wide as long. Pronotum 0.8 times as long as wide; widest one- third length from base, sides rather strongly arcuate, anterior constriction weak, rather broadly rounded in front; surface coarsely, deeply piinctuied, spaces between pimctures almost smooth, subshining, less than half the width of a puncture, with a small rounded, setiferous granule on anterior margin of each puncture; vestiture consisting of short, stout, whitish bristles. Elytra I.l times as long as wide. 1.5 times as long as pronotum; sides straight and [jarallel on basal two- thirds, broadly rounded behind; single row of crenula- tions at elytral base high, coarse; striae slightly impressed, the punctures coarse, deep; interstriae convex, narrower than striae, armed by a row of small setiferous tubercles. Declivity steep, convex; striae somewhat narrower and deeper, interspace 9 weakly elevated, more coarsely serrate. X'estiture consisting of uniseriate rows of suberecl. flat- tened bristles, each bristle separated in its own row by distances equal to its own length, and from adjacent tows by distances about equal to one and one IkHI limes its own length. Female. — Similar to male except frons convex, the lateral margins rounded; and antennal scape without tuft of hair. Type Locality. — Zamorano, Morazan, Hon- Hosts. — Ficus glabrala (type), Ficu.t spp, (paratypes), Celtis iguanae (paratype), and Ser- jania triquetra (paratype). Type Material. — The male holotype, female allotype, and 39 paratypes were taken at the type locality on April 18, 1964, at an elevation of 700 m., by S. L. Wood, from the phloem tissues of twigs of Ficus glabrata; 35 paratypes bear identical data but were taken from Serjania triquetra; 1 bears the same data but is from Celtis iguanae; 16 paratypes were colledted at the lower Rio Teinpisque, Guanacaste Prov,, Costa Rica, on March 25, 1964, at 15 m. elevation from Ficus twigs; 1 paratype is from Rio Damitas in the Dota Mountains, San Jose Prov., Costa Rica, taken on February 18, 1964, 200 m. eleva- tion, from an unknown branch; and 1 paratype is from Palin, Esquintla Piov., Guatemala, taken on May 19, 1964, at 300 m. elevation, from a Ficus twig; all were collected by S. L. Wood. The holotype, allotype, and paratypes are in my collection. Phloeotribus squamatus, n. sp. 1 his unique species is distinguished from other species in the genus by the sinall size, by the unarmed pronotum, by the short, squamose vestiture, by the shallowly impressed male frons, and by other characters. Male. — Length 1.25 mm. (allotype 1.3 mm.). 2.1 times as long as wide; color black, with white squamo.se vestiture. Frons shallowly concave from upper le\el of eyes to epistoma, the lateral margins not acutely elevated 8 Bricham Young University Science Builetin or armed; epistomal margin weakly elevated, and with a median tubercle; surface coarsely reticulate; vestiture inconspicuous, mostly confined to marginal areas. Anlen- nal club small, compact; segment 1 about l.,'> times as wide as long. Pronotum 0.8 tiines as long as wide; widest just behind middle, sides moderately arcuate, moderately constricted just behind the rather narrowly roimded anterior margin; surface rather coarsely reticulate in- cluding the punctures, with close, coarse, shallow punc- tures; vestiture consisting of suberect, short, stout, almost scalelike bristles, most of them about three times as long as wide. Elytra 1.4 times as long as wide. 1.9 times as long as pronotum; crenulations of elytral base coarse, rather high, forming a single row; striae weakly impressed, the punctures large, rather deep; interstriae about half as wide as striae, almost restricted to a narrow row of closely set, subvtilcanate. squamiferous pimctines. De- clivity rather steep, convex; striae somewhat narrower and deeper than on disc. Vestiture consisting of imiseriate rows of erect interstrial scales, each scale less than twice as long as wide; separated from adjacent scales in the same row by distances equal to less than the width of a scale, and between rows by distances equal to aliiK>st twice the length of a .scale. Female. — Similar to male except frons convex, the median epistomal tubercle absent; vestiture of pronotum and elytra very slightly more slentler. Type Locality. — 7apanti, Cartago Prov.. Costa Rica. Type Material. — The male liolotype and female allotype were collected at the type locality on July 8, 1963, at an elevation of 1,300 m., by S. L. Wood, from a dying tree that bore fruiting pods. The holotype and allotype are in my collec- tion. Chaetophloeus phurodendri, n. sp. This species has the same arrangement of crenulations at the base of the elytra as brazilien- sis (Blackman), but it differs by the much larger size, by the reduction of elytral striae, by the very different vestiture and by other characters. Male. — Length 1.8 mm. (paratypes 1.5-2.0 mm.). 1.4 tiines as long as wide; color black, vestiture cinereus. Frons broadly, deeply concave from vertex to the broadly emarginale epistomal margin, and from eye to eye; surface granulose-reticulate; vestiture sparse in con- cavity, a few erect gray setae along sides below eye. margin above eyes ornamented by a row of long reddish hair many of which reach mandibles; mandibles each with an anterior extension half as large as mandible. Antennal club about as in brazitiensis. Pronotum 0.53 times as long as wide; widest at base, strongly arcuately converging to the broadly roimded anterior margin; surface retictdate, with small, rather close setiferous punctures; lateral areas armed by two groups of asperities; vestiture consisting of mixed slender and stout, short bristles and a few bifid hairs in lateral areas. Elytra 1.1 times as long as wide. 2.1 times as long as pronotum; anterior margins armed by six pairs of crenulations. the median pair twice as wide as others. an additional submarginal pair at base of interspace 1; striae reduced, obscure, the punctiues only slightly larger than those of interspaces; interspaces wide, flattened, with numerous punctures variable in size. Declivity steep, convex. Vestiture consisting of short, rather abim- dant, slender scales, and median rows of erect interstrial scales, each scale about three times as long as wide, separated from other scales in .same row by distances equal to length of scale and between rows by one and one-half times length of a scale. Female. — Similar to male except frons flattened, with a small median impression, frontal vestiture short, not specialized; pronotiun armed by three paired groups of asperities. Type Locality. — Zamtjrano, Morazan, Hon- duras. Host. — Plwradendron robustissimum (type), and Plioradendron sp, (paratypes). Type Material. — The male holotype, female allotype, and 40 paratypes were collected at the type locality on April 18, 1964, at an elevation of 700 m., by S. L. Wood, from twigs of the above mistletoe; 4 paratypes were taken at Caiias, Guanacaste Prov., Costa Rica, on April 10, 1966, from Plwradendron twigs; one paratype was taken 20 km. southeast of Liberia, Guanacaste Prov., Costa Rica, on July 10. 1966, from Phora- dendron twigs; all were collected by S. L. Wood. The holotype. allotype, and paratypes are in my collection. Lipartlnum uniericuuuin, n. sp. This species is allied to scjuamomm (Black- man), but is readily di.slinguished by the much smaller, more strongly impressed elytral striae, and by the broadly concave male frons. Male. — Length 0.8 mm., about 2.1 times as long as wide; color rather light brown. Frons shallowly conca\'e almost from eye to eye and from \ertex to episloma; surface reticidate; vestiture largely confined to impressed area, consisting of very fine, minute, abundant, pilose hair of uniform length. Pronotum 0.8 times as long as wide; widest one- third from base, sides moderately arcuate, rather nar- rowly rounded in front; surface obscured by vestiture. evidently minutely subgranidate and with small setiferous granules uniformly distributed; vestiture consisting of stout reciuiibent bristles and erect broad scales in about ecjual numbers. Elytra 1.3 times as long as wide, 1.7 times as long as pronotum; sides straight and parallel on basal two- thirds, rather broadly rounded behind; bases armed by six pairs of crenulations between striae 4; striae not impressed, the punctures rather large, deep; interstriae almost as wide as striae, weakly convex, the moderately large, setiferous punctures imiseriate. feeble gianulate on anterior side. Declivity steep convex; striae and inter- striae somewhat narrower than on disc. Vestiture con- sisting of strial rows of short recumbent, stout hair, and interstrial rows of erect, broad scales, each scale almost as wide as long. Type Locality. — The lower Rio Tcmpisque, Guanacaste Prov., Costa Rica. Host. — Ficus sp. New Neotropical Scoi.'i tidae Type Material. — The unique male holotype was colletted at the type locality on March 25, 1964, at ail elevation of about 20 ni., by S. L. Wood, from small branches of the common nar- row leafed fig that grew near the river. The holotype is in my collection. A second specimen, tentatively assigned to this species, possible the female, is identical to the type except the frons is less deeply, less extensive- ly impressed and the pilose vestiture is not evi- dent. It was taken at Dominical, Puntarenas Prov., Costa Rica, on December 9, 1963, from an iMiidentified branch. Liparthrum cracentis, n. sp. This species is allied to the four previously described American species but it may be dis- tinguished by the small size, by the more slender body form, by the poorly developed female frontal vestiture, and by the narrowei elytral scales. Female. — Length 0.8 nun. (paiatypes 0. 7.5-0. K,5 mm.), 2.2 times as long as wide; color brown. Frons moderately convex, slightly impressed just above epislomal margin; surface rather coarsely reticulate; vestiture sparse, more conspicuous toward episloma, consisting ol stout, siibplnmose setae. Eye anil antenna as in allied species. Pronotmn 0.81 times as long as wide; widest on basal third, sides weakly arcuate from base, converging on an- terior half to the broadly rounded anterior margin; sur- face shining, feebly subreticulate. the obscure, setiferous punctures rather close, their anterior margins very finely asperate. X'esliture consisting of truncate, subtriangular scales, each about one and one-half times as long as wide, and equally abundant stout hairlike setae, each about one and one-half times as long as scales. Elytra l.ti times as long as wide. 1.96 times as long as pronolum; basal margins armed by five pairs of crenulations between interstriae 4; striae not impressed, the punctures coarse, rather deep; interstriae al)oiU half as wide as striae, the punctures fine. unLseriate. evidently not granulate. Declivity rather steep, convex; sculpture about as on disc, \estiture consisting of strial rows of bristles equal in length to scales in interstrial rows; each scale almost as wide as a strial puncture and slightly longer than wide. Male. — Similar to female except pronotum bearing a pair of small, median tubercles on anterior margin and a smaller submarginal pair about one-fourth pronotum length from anterior margin. Type Locality. — Twenty-nine km. west Nilti- pec, Oaxaca, Mexico. Type Material. — The female holotype, male allotype, and 29 paratypes were collected at the type locality on June 23, 1967, at about .30 m. elevation, by S. L. Wood, fiom the pholem of small twigs of a small tree having narrow, thick, waxy, simple leaves. The identification of the host is not yet available. The holotype, allotype, and paratypes are in my collection. Cnernonyx nitetis. n. sp. Evidently allied to atratits (Blandfortl), but distinguished by the more extensive frontal im- pression of the male, by the elongate pronotal punctures and minutely, longitudinally strigose pronotal surface, and by the coarser interstrial punctures of the elytra. Male. — Length 2.5 mm. (paratypes 2.0-2.5 mm.). 2.1 times as long as wide; color almost black. Frons shallowly conca\e on median half from upper level of eyes to epistoma; surface reticulate above, the punctures rather coarse, deep, clo.se. becoming finer and closer below in concave area; vestiture short and rather abundant in concave area, longer at sides of this area, evidently glabrous elsewhere. Antennal club subtriangu- lar. finely, closely covered by hairlike pubescence; sutures not evident. Pronotum 1.0 times as long as wide; widest at base, sides weakly arcuate and converging slightly to the slight constriction just behind the broadly rounded anterior margin; sinlace shining, shallowly etched by minute, longitudinally strigose lines, becoming reticulate anterior- ly, the punctures fine and elongate posteriorly, almost round and larger anteriorly and laterally; glabrous. Elytra 1.2 limes as long as wide, l..*? times as long as pronotum; sides straight and parallel on basal half, then arcualely converging lo the rather narrowly rounded, .sutuially subemarginate, posterior margin; striae 1 weakly. others feebly impressed, the punctures rather small, deep, close; interstriae smooth, shining, about twice as wide as striae, the punctures about half as large as those of striae, uniseriate behind, moderately confused on anterior half of di.sc; elytral bases with fine raised line; base of interspace 1 not depressed; sculelluni small, as wide as long, almost flush with elytral surface. Declivity moderate- ly steep, convex; striae and interstriae narrower than on disc, the striae narrowly impressed; interstriae moderately, broadly convex, shining except sutural half of 1 dull, the punctures iniiseriate. Vestiture confined to declivity, consisting of sparse minuc. short, siiberect scales arising from interstrial punctures. Female. — Similar to male except frons flattened, the \estiture reduced; strigose lines on pronotum rather obscure; interstrial punctures on elytral disc not as con- fused; and surface of declivity less brightly shining. Type Locality. — Puerto Viejo, Heredia Prov., Costa Rica. Type Material. — The male holotype, female allotype, and 34 paratypes were collected at the type locality on March 12, 1964, at an elevation of about 70 m., by ,S. L. Wood, from bark of the limb of a fallen tree. Ihe holotype, allotype, and paratypes are in my collection. Cnernonyx insignis, n. sp. This species evidently is allied to panamensis (Blandford), but is distinguished by the larger size, by the procurved epistomal carina, and by other characters. Male. — Length 3.5 mm. (paratypes 3.5-3.9 mm.). 2.0 times as long as wide; color dark brown. Frons shallowly concave on median two-thirds from upper level of eyes to epistoma; surface reticulate-granu- 10 Bricham Young University Science Bulletin late, and closely, deeply, rather coarsely punctured in concave area, shallowly, less closely punctured above; slightly less than median half of epistoma armed by an elevated, procurved, transverse carina; vestitiire confined to concavity, consisting of short, erect, rather abundant, dark hair. Pronotum 0.9 times as long as wide; widest at base. the sides weakly arcuate and converging toward the rather broadly rounded anterior margin; surface smooth and shining on posterior three-fourths, with minute points and fine punctures, anterior fourth rather coarsely subreticulate, dull, faintly, longitudinally etched in lateral areas, the punctures very slightly larger in anterior area, much larger laterally; glabrous. Elytra 1.1 times as long as wide. 1.2 times as long as pronotum; scutellum as wide as long, not depressed; basal margins of elytra with an irregular raised line; outline as for nitens, above; striae rather strongly im- pressed, the punctures fine, shallow, distinct; interslriae smooth, shining, at least twice as wide as striae, moderate- ly convex at base, becoming rather narrowly costiforin by declivital base, the punctures fine and uniseriate be- hind, confused toward base. Declivity moderately steep, convex; striae broadly impressed, the fine shallow punc- tures evident; interstriae narrowly, acutely, subserralely elevated; striae and interstriae minutely reticulate-granu- late: costal margin rather coarsely serrate. Rows of very minute interstrial scales evident on declivity. Female. — Psesumable ihe female is represented by specimens having the frons less strongly impressed, bul otherwise similar to the male. Type Locality. — Moravia, Cartago Prov., Costa Rica. Type Material. — The male holotype, female allotype, and 5 paratypes were taken at the type locality on March 11, 1964, at an elevation of about 500 m., by S. L. Wood, as dead specimens under loose bark of an unideniified log about 40 cm. in diameter. The holotype, allotype and paratypes are in my collection. Cnemonyx opacus, n. sp. This species, at least superficially, resembles insignis Wood, above, but is easily distinguished by the smaller size, by the narrow frontal carina, by the much more extensively granulate elytra, and by characters mentioned in the key. How- ever, the antennal club has an entirely different shape and has three strongly procurved sutuies clearly marked by rows of setae suggesting that the relationship is superficial at best. Male. — Length 2.2 mm. (paratypes 2.0-2.5 nun.). 2.2 times as long as wide; color dark brown. Frons shallowly concave on median half from upper level of eyes almost to epistoma, the surface closely, deeply, rather coarsely punctured; median fourth of epistoma bearing a strongly elevated, procurved, trans verse carina; vestiture consisting of short, erect, rather abundant, stout hair in concave area. Antennal funicle with inner angle of .segments projecting and bearing tufts of long yellow hair; club obovate, longer than wide, with three strongly procurved sutures conspicuously marked by rows of closely set setae. Pronotum about as wide as long; widest on basal third, the sides rather weakly arcuate and converging to the weak, transverse constriction just behind the rather narrowly rounded anterior margin; surface smooth and shining behind, becoming reticulate anteriorly and to- ward lateral margins, the punctures rather coarse, deep, close, somewhat larger in lateral areas; glabrous. Elytra 1.2 times as long as wide. 1.3 times as long as pronotum; sides straight and parallel on more than basal half, rather narrowly rounded behind; scutellum very small, rounded, not depressed; basal margins of elytra marked by a fine, irregular, raised line; striae impressed, the punctures rather large, deep, clearly impressed in basal area, obscure behind; interstriae convex, finely, completely granulate (including striae) posteriorly, coarse- ly punctured on basal third of disc. Declivity moderately steep, convex, striae deeply impressed, the punctures obscure; interstriae narrow, strongly convex, minutely granulate, finely subserrate; interspace 9 moderately ele- vated, continuing to apex, highest at junction with 3. Vestiture confined to tieclivity, consisting of uniseriate rows of minute interstrial scales. Female. — Similar to male except frons less strongly impressed, almost flat. Type Locality. — Playa del Coco, Guanacaste, Costa Rica. Host. — Hippomane mancinella. Type material. — The male holotype, female allotype, and 64 paratypes were collected at the type locality on October 18, 1963. at sea level, by S. L. Wood, from the bark of the above host tree. This agressive species constructs biramous, transverse, egg tunnels in the bole and branches of the host tree. Adult and the irregular larval tunnels were almost entirely in the phloem tis- sues. The holotype, allotype, and paratypes are in my collection. Cnemonyx gracilens, n. sp. This species is very closely related to exilis Wood, bul it may be distinguished by the smaller size, by the more broadly impressed frons with the epistomal carina higher, Ijy the less numer- ous, more nearly oval pronotal punctures, and by the shorter, finer, less strongly flattened elytral setae. Male. — Length 1.3 mm. (paratypes 1.2-1.4 mm.), 2.6 times as long as wide; color dark brown. Frons as in exilis but more broadly planoconcave, the transverse epistomal carina less procurved. slightly higher, occupying at least two-thirds width of epistomal margin; pubescence perhaps a little more abundant. Pronotum 1.09 times as long as wide; punctures dis- tinctly less numerous than in exilis, each aboul one and one-half times as long as wide. Glabrous. Elvira 1.6 times as long as wide. 1.5 times as long as pronotum; as in exilis interstrial bristles on declivity much shorter, very weakly or ot at all flallened. narrower, not increasing in width apically. Female. — Similar lo male except frons less strongly impressetl. Type Locality. — Fifty -seven km. southeast of New Neotropecal ScoLVTroAE El Cameron. Oaxaca, Mexico. Type Miilenal. — The male holotype, female allotype, and 98 paiatypes were collected at the type locality on June 22, 1967, No. 81, by S. L. Wood. Specimens were taken from phloem tis- sues of twigs of a small tree that had peeling bark, thin simple leaves, and abundant white latex; a sample awaits identification. The holotype, allotype and paratypes are in my collection. Cnemonyx recavus, n. sp. This species is allied to impressus Wood, but it is readily distinguished by the larger size, by the much more strongly impressed frons and by the dull pronotal surface. Male. — Length 1.7 mm. (paratypes \.5-\.H mm.), 2.1 times as long as wide; color dark reddish brown. Frons bioadly, rather deeply concave from just below upper level of eyes to epistomal margin: concavity dense- ly pubescent, the setae moderately, uniformly long; sur- face of maiginal areas and above coarsely reticulate, with fine, close punctures; surface in concavity obscured by pubescence. Scape short; fimicle bearing a dense tuft of long hair; club obovate. with three strongly procur\ed sutures marked by setae. I extending almost two-thirds club length from base. Pronotum 0.92 liines as long as wide; basal margin bisinuate. marked by a fine, raised line; widest at base, arcuately converging to a slight constriction just before the broadly lounded anterior margin; surface didl, minutely, longitudinally etched, the punctures fine, oval, close, nroderately deep. Glabrous. Elytra 1.2 times as long as wide. 1.4 times as long as pronotum; sides almost straight and parallel on slightly moie than basal half, rather narrowly rounded behind; basal margins indistinctly rai.sed along a fine line; striae mipressed, the punctures fine, distinctly impre.s,sed; inter- striae almost twice as wide as striae, convex, dull, each with a uniseriale (confused on 2) row of fine, short, transverse cienulations, the puncture posterior to each crenulation fine, obscure. Declivity convex, moderately steep; about as on declivity, with interstriae 9 weakly elevated and more coarsely tuberculate, 10 with a row of fine tubeicles posteriorly, \estiture confined to decliv- ity, consisting of interstrial rows of very short, flattened bristles each about twice as long as wide. Female. — Similar to male except frons very slightly less strongly impressed. Type Locality. — Forty-eight km. east of Te- huantepec, Oaxaca, Mexico. Type Material.~The male holotype, female allotype, and 87 paratypes were collected at the type locality on June 22, 1967, 120 m., No. 84, by S. L. Wood. The host was s small tree with simple leaves; a sample awaits identification. The holotype, allotype, and paratypes are in my collection. A series taken from "Leche Sillo," 16 km. northwest of Acatlan, Puebla, Mexico, June 15, 1967, 1,700 m.. No. 36, by S. L, Wood, not desig- nated as paratypes, have the posterior half of the pronotum smooth and shining, and the punc- 11 tures behind the interstrial cienulations are more conspicuotis. The series overhip in these charac- ters to such an extent that geographical races can- not be disignated. Camptocerus infidelis, n. sp. This species is sllied to auricomus Blandford, but is not closely related. It differs in the female by the more strongly convex frons above, and narrower below, by the much less closely punc- tured pronotum, by the inore sparsely punctured elytra, by the near absence of elytral scales, by the smaller size, and by other characters. ffma/c— Length 2.7 mm. (paratypes 3.2 mm.). 2.0 times as long as wide; color black. Frons rather strongly convex above, inore nearly llaitened below, with a distinct, transverse impression just abo\e epistomal margin; surface rather coarsely reticulate and with rather abundant, shallow, indefinite punctures; vestiture spaise. inconspicuous, .\ntenna essen- tially as in auricomus. Pronotum 0.98 times as long as wide; widest near base, sides rather weakly arcuate and converging slightly to the rather strongly lOunded anterior angles, broadly rounded in front; surface minutely reticulate anil dull on anterior half, almost smooth and subshining behind; the punctures minute, sharply impressed, sepiirated by distances ec]ual to two or moie diameters of a pimcture; glabrous. Elvtra 1.2 limes as long as wide, 1.3 times as long as pronotum; widest at base, the sides converging pos- teriorly to the narrow, subemarginate posterior niaigin; striae not impressed, the punctures small, shallow, rather widely spaced; interstriae twice as wide as striae, shining, somewhat irregular, very sparsely punctured, the shallow- punctures almost as large as those of striae. Declivity gradual, convex; striae weakly impressed. Vestiture con- sisting of very sparse interstrial scales on sides and de- clivity. Male. — Not represented in the material at hand. Type Locality. — Peralta, Cartago Prov,, Costa Rica. Type Material. — The female holotype and one female paratype were collected at the type locality on March 10, 1964, at an elevation of about 500 m., by S, L. Wood, from a dead tree seedling 1 cm. in diameter. The egg tunnel fol- lowed the central axis of the host for several centimeters, the larval cradles extended into the wood at an acute angle to the central axis. Two adult females and small larvae were in the only tunnel found. Both specimens are in my collection. Scolytus nodatus, n. sp. Fig. 1 In the American Scolytus fauna this species is unique. The female abdominal sterna are un- armed; in the male sternum 2 is armed by a large median tooth, 4 is armed by a large, median, rounded knob, and the lateral margins of 2 and 3 are each armed by a pair of small teeth; the frons is convergently aciculate and pubescent in 12 BrIC.H AM VOUNG UNIVERSITV SCIENCE BULLETIN Fig. 1. Scolytus nodalus abdomen and elytra: male at upper left and below: female at upper right. both sexes, it is devoid of tubercles in both sexes. Male. — Length 3.8 mm. (paralypes 3.1-3.8 mm.). 2.0 times as long as wide; color dark brown, anterior margin of thorax, bases and apices of elytra may be reddish. Frons weakly convex; surface coarsely, somewhat con- vergently strigose toward the shallowly, broadly emar- ginate epistomal margin: punctures not evident: vestiture consisting of fine, rather long, moderately abundant hair. Eye emarginale; very finely granulate. Antennal club 2.0 times as long as wide; sutures not evident, very finely, closely pubescent. Pionotum equal in length and width; widest at base, the sides moderately arcuate and converging to dis- tinct constriction just behind broadly rounded anterior margin; surface smooth, brightly shining, the pinictures rather small, deep. oval, larger in lateral and anterior areas; glabrous on disc, sparse hair at sides and in front. Elytra I.I times as long as wide, l.I times as long as pronotum; sides straight and parallel on basal half, con- verging slightly behinil to the broadly rounded postero- lateral angles, almost straight on median half; posterior margin smooth, striae and interstriae moderately im- pressed, the punctures small, rather deep, those of striae very slightly larger; elytra descending very slightly near posterior margin; vestiture confined /to scutellar impres- sion. Sternum 2 ascending sharply, the margin not pro- duced, armed by a laterally compressed, median spine, its base extending from anterior margin to middle of segment, longitudinal length of base and height of spine about equal; sternum 4 armed by a large, rounded tubercle, 2 and 3 each armed at lateral margins by a pair of small teeth; surface finely, obscurely pinictined, dull except anterior half of 2 shining; pubescence sparse, hairlike. Female. — Similar to male excpet frons more nearly flat; all abdominal sterna unarmed. Type Locality. — Santa Ana, San Jose Prov., Costa Rica. Type Material. — The male holotype, female allotype, and 4 paratypes were taken at the type locality on November 8, 1963, at an elevation of 1,300 m., by S. L. Wood, from the limb of an unidedntified tree. Nineteen other paratypes were collected at Fort Clayton, Panama Canal Zone, on December 22, 1963, at an elevation of 30 m., by S. L. Wood, from a large, woody vine (liana) about 5 cm. in diameter. The holotype, allotype, and paratypes are in my collection. Scolytus cristatus, n. sp. Fig. 2 This species does not appear to be closely related to any known Neoptopical representa- tive of the genus. The frons of both sexes is con- vex, convergently strigose, with a very fine, low, long, median, subcarinate ridge, and in the male moderately abundant, uniformly distributed, long, frontal hair; the second abdominal sternum ascends rather abruptly, not sharply, and is arined by a high, nanow carina on anterior two thirds, its highest point near its posterior ex- tremity. k New Neotropical Scoi.vtidae 13 Fig. 2. Scolytiis crhtalus posterolateral aspect of male abdomen. Male.— Length 2.4 mm. (paratypes 2.3-2.7 mm.), 2.0 times as long as wide; color reddish brown (mature para- types dark reddish-brown). Frons convex, somewhat flattened below; surface convergently strigose, the grooves usually wider than the ridges, the median ridge higher, subcarinate, extending from epistomal maigin almost to upper level of eyes; vestiture fine, long, rather abundant, uniformly distri- buted. Pronotum very slightly wider than long; widest on basal third, sides moderately arcuate and converging slightly to constriction just behind the broadly rounded anterior margin; surface smooth, shining, with .some very minute points and rather widely separated, moderately small, deep, oval punctures, the punctures larger in lateral areas and in front; glabrous. Elytra 1.1 times as long as wide, 1.2 times as long as pronotum; striae and interstriae strongly, equally im- pressed, the punctuies moderately large, shallow, the ridges between rows of punctures smooth, shining, con- vex, almost as wide as grooves; declivity descending slightly; a few scattered, slender scales on declivity. Venter with sterninn 2 ascending abruptly, not sharply, armed by a median, laterally compressed, rather high elevation extending fiom near anterior margin of segment two- thirds of its length, highest point on elevation near its posterior limits; sternum 2 very coarsely, shallowly punc- tured, the smface within each puncture reticulate, the inteispaces subreticulate; sternum 3 with one row of punctures. 4 and 5 not clearly punctured, dull. Female. — Similar to male except frons subglabrous; carinate armature of sternum 2 not as high, occupying middle third of length of segment. Type Locality. — Lower Rio Tempisque, Gua- nacaste Prov., Costa Rica. Type Material. — The male holotype, female allotype, and 29 paratypes were collected at the type locality on March 25, 1964, at an elevation of about 20 m., by S. L. Wood, from the bark of an unidentified broken branch. The transverse, biramous egg tunnels engraved the wood rather deeply; larvae were not present. A series of this species, not included in the type series, was collected 8 km. south of Aten- quique, Jalisco, Mexico, on June 25, 1965, from an unidentified woody vine (liana). The holotype, allotype, and paratypes are in my collection. Microborus limatus, n. sp. This species differs from all previously de- scribed representatives of this genus by the more narrowly rounded (subangulate) posterior mar- gin of the elytra, by the more gradual elytral declivity, and, particularly, fjy the brightly shin- ing subglabious elytral surface with the sparse, declivital, hairlike setae that are separated from one another by one to two or more times the length of a seta. Afa/p.— Length 1.4 mm. (paratypes 1.3 1.5 mm.). 2.7 times as long as wide: body color dark brown, elytra medium brown. Frons narrow and feebly convex between eyes, pro- truding very slightly just above antennal bases; surface smooth and polished, with a few scattered, minute punc- tures; almost glabrous. Eyes very large, almost touching below; anterior maigin very broadly emarginate; very coarsely faceted. Antennal scape short, about twice as long as pedicel; funicle 6-segmenled; club subglobular. sutures obscurely marked by rows of setae. Pronotum 1.25 times as long as wide; sides almost straight and parallel on basal three-fourths, rather broad- ly rounded in front; anterior angles not narrowed; sur- face shining, weakly reticulate, the punctures rather small and shallow, separated from one another by one to two diameters; glabrous. Elytra 1.9 times as long as wide; sides almost straight and parallel on basal three-fourths, narrowly (subangu- lately) rounded behind; striae 1 modeiately. 2 weakly im- pressed, narrow, the punctures small, separated by dis- tances equal to their own diameteis; interstriae distinctly wider than striae, smooth, brightly shining, each with a sparse row of minute punctures. Declivity rather steep, not precipitous; striae and interstriae about as on disc, except inlerstrial punctures very finely granulate; inter- spaces 7 and 9 acutely elevated from declivital ba.se to point of fusion, the acute elevation continuing to apex. Vestiture short, sparse, almost entirely lestricted to de- clivity; each hairlike seta about equal to half the width of an interspace, and separated from one another by about one and one-half to two times the length of a setae. Female. — Similar to male except more finely sculp tured; the declivital gianules scarcely visible. Type Locality. — Finca Gromaco on Rio Goto Brus (34 km. from mouth). Puntarenas Prov., Gosta Rica. Type Material. — The male holotype, female allotype, and 11 paratypes were taken at the type loclaity on July 14, 1963, at an elevation of about 500 m., by S. L. Wood, from the phloem tissues of a large limb of an unknown tree that had been cut at the margin of the jungle. The holotype. allotype, and paratypes are in my collection. Microborus ambitus, n. sp. This species is closely related to liinatus Wood, described above, but it may be distin- guished by the slightly deeper pronotal punc- tures, by the impressed, much more coarsely punctured declivital striae, and by the fine inter- strial granules on the declivity. Female. — Length 1.3 mm. (paratypes 1.3-1.4 inm.). 2.9 times as long as wide; color of head and pronotum very dark brown, posterior areas and legs reddish brown. 14 Br[(;ham Young University Science Bulletin Head, pronotum, and elytral disc as in limalus except. as noted above, the pionotal puctuies slightly deeper. Elytral declivity convex, steep; striae slightly impressed, the punctures coarse, deep; inteistriae slightly narrower than striae, veiy slightly convex, each bearing a iiniseriate row of fine granules; interstriae 7 very acutely, strongly elevated, joining 9 and continuing but declining in height slightly along costal margin to sutural apex, 9 narrowly. not strongly elevated from level of hind coxae to junction with 7. Vestiture confined to declivity, consisting of interstrial rows of fine, short, almost hairlike bristles. Male. — Although both sexes are probably represented in the series sexual differences are not apparent. Type Locality. — Twenty-five km. east of Co- atzacoalcos, Veracruz, Mexico. Type Material. — The female holotype and two paratypes were collected at the type locality on June 26, 1967, 30 m., No. 110, by S. L. Wood. Three paratypes were taken 22 km. west of Car- denas, Tabasco, Mexico on June 26, 1967, 30 m.. No. 106, by S. L. Wood. Both series were taken from phloem tissues of limbs and bole 10-20 cm. in diameter of the same unidentifiable species of cut tree. The holotype and paratypes are in my collec- tion. Scolytodes (s. str.) clusiae, n. sp. Evidently not closely related to previously known species. It has interstriae 10 ending before level of metacoxae, the protibiae armed by a small tooth on posterior face just before tarsal insertion and between terminal mucro and first or second marginal tooth; female frons broadly, shallowly concave. Female. — Length 1.8 mm. (paratypes 1.71.9 mm.). 2.4 times as long as wide; color black. Frons broadly, shallowly concave from upper level of eyes to level of antennal insertion; surface closely, deeply, somwhat coarsely punctured over entire frontal area in and out of concavity; median epistomal lobe con- spicuous, broad, short, continuous with surface of frons; vestiture very fine, not abundant, longer above, not evident along epistoma. Pronotum 1.0 times as long as wide; widest at base, very slightly constricted one-third from base, weakly arcuate on anterior half then broadly rounded in front; surface reticulate, with fine, obscure, rather widely spaced punctures on posterior half, equally fine sparse, minute granules on anterior third; almost glabrous. Elytra 1.4 times as long as wide, 1.7 times as long as pronotum; widest one-third from base, the sides feebly arcuate, abruptly rounded at declivital base, narrowly rounded behind; striae not impressed, the punctures very small, not deep; interstriae about twice as wide as striae, shining, irregular due to obscure surface lines, the punc- tures not evident. Declivity convex, rather steep; striae 1 weakly impressed above, the punctures on all striae obscurely indicated. Vestiture consisting of a few rather short hairs and. on lower half, more numerous, very short, semirecumbent hair. Male. — Similar to female except frons rather strongly convex, finely, obscurely punctined. surface reticulate, vestiture sparse, short, inconspicuous; strial punctures rather obscure. Type Locality. — .Summit of Volcan Poas, He- redia Prov., Costa Rica. Host. — Clusia sp. Type Material. — The female holotype, male allotype, and 16 paratypes were collected at the type locality on August 19, 1966, by S. L. Wood, from green phloem of dying twigs of the common Clusia tree in the area. The holotype, allotype, and paratypes are in my collection. Scolytodes (s. str.) volcanus, n. sp. Closely related to clusiae Wood, described above, but smaller, the small tooth on posterior face of protibiae absent, the strial punctures larger, and the female frons more narrowly im- pressed to vertex. Female. — Length 1.3 mm. (paratypes 1.2-1.4 mm.). 2.3 times as long as wide; color very dark brown, almost black. Frons narrowly, shallowly concave from epistomal margin to above upper level of eyes; surface fineh. shal- lowly but distinctly, densely punctate, epistomal lobe mostly shining; vestiture largely confined to margin of impressed area, fine, short below, somewhat longer above, moderately abundant, minute in concavity. Pronotum 1.1 times as long as wide; sides almost straight and parallel on basal half, then somewhat arcu- ate and rather narrowly lounded in fiont; surface reticu- late, the punctures fine, sparse, rather obscure, devoid of granules; glabrous. Elytra 1.5 times as long as wide. 1.6 times as long as pronotum; sides almost straight and parallel on basal two-thirds, narrowly rounded behind; striae not im- pressed, the punctures moderately large, deep; inter- striae slightly wider than striae, shining, with very few- lines, the punctures obscure, mostly obliterated. Declivity rather steep, convex; an occasional deep, interstrial punc- ture present; shining. Glabrous. Male. — Similar to female except frons convex, reticu- late, obscurely punctuied, the vestiture gieatly reduced, inconspicuous; a few interstrial punctures clearly im- pressed. Type Locality. — Volcan Poas, Heredia Prov., Costa Rica. Type Material. — The female holotype, male allotype, and 24 paratypes were collected at the type locality on July 14, 1963, 100 m. (elevation) below the summit, by S. L. Wood, from a small woody vine and cut tree seedlings. The holotype, allotype, and paratypes are in my collection. Scolytodes (s. str.) paniulus, n. sp. Similar to volcanus Wood, described above, but smaller, the female frons armed by a pair of longitudinal carinae, the strial punctures re- duced: the female frontal vestiture uniforiTily distributed. Female. — Length 1.1 mm. (paratypes 0.9 I.I mm). 2.3 times as long as wide; color rather dark brown. New Neotropical Scolvtidae 15 Fions flattened from upper level of eyes to epistoma, the median half of each side just inside of lateral inargin armed by a shining longitudinal carina; the pair of carinae beginning at level of antennal insertion and end- ing before upper level of eyes; surface finely, rather closely punctured in impressed and lateral areas, reticulate above eyes; vestiture fine, moderately long around margin, somewhat shorter in central area, moderately abundant. Pronotum 1.0 times as long as wide: sides very weakly arcuate and feebly narrowed anteriorly on basal two- thirds, broadly rounded in front; surface reticulate, with sparse, minute punctures extending from base to anterior margin; glabrous. Klvlr;i l.:i limes as long as wide. 1.:") limes as long as pronotum; sides almost straight and parallel on basal half, rather narrowly rounded behind; striae 1 weakly impressed, others not at all impressed, the punctures fine, shallow, rather obscure; interstriae wider than striae, not entiiely smooth, the punctures sparse, fine, obscure. Declivity convex, steep; essentially as on disc but inter- spaces narrower and punctures smaller, the features not sharply defined. Vestiture consisting of less than a dozen moderately long, erect interstrial bristles. Male. — Similar to female except frons convex, with a slight transveise impression just above epistoma, sur- face obscurely reticulate, shining, sparsely punctured. vestiture very sparse, mostly near epistoma; pronotum obscurely reticulate. subshining. Type Locality. — Pandora, Limon Prov., Costa Rica. Host. — Cecropia sp. Type Material. — The female holotype, male allotype, and 12 paratypes were collected at the type locality on August 23, 1963, at an elevation of about 50 m., by S. L. Wood, from Cecropia leaf petioles. The holotype, allotype, and paratypes are in my collection. Scolytodes (s. str.) acares, n. sp. Allied to pamuliis Wood, described above, but the female frontal vestiture longer and much more abundant near vertex, and the pronotal punctures more coarsely, deeply impressed in posterior area. Female. — Length 1.1 mm. (paratypes 0.9-1.1 mm.). 2.4 times as long as wide; color dark brown. Frons narrowly flattened from well above upper level of eyes to epistoma. margined at sides by a pair of shining, longitudinal carinae extending almost from epistoma (half way bewteen level of antennal insertion and episto mal margin) to near upper level of eyes: surface sculpture obscure between carinae. evidently smooth and finely punctured; vestiture largely confined to vertex above upper level of eyes, consisting of a dense, long brush of subplnmose, yellow hair that extends in length to a point below level of antennal bases. Pronotum and elytra as in parvulus (above), except pronotimi mote narrowly rounded in front and its punc- tures somewhat larger. Male. — Similar to female except frons convex with a slight transverse impression just above epistoma. the surface leticulate. sparsely punctured, the vestiture re- duced to a few setae near epistoma. Type Locality. — Rio Damitas in the Mountains, San Jose Prov., Costa Rica. Host. — Cecropia sp. Dota Type Material. — The female holotype, male allotype, and 14 paratypes were collected at the type locality on August 22, 1963, at an elevation of about 250 m., by S. L. Wood, from the petioles of fallen Cecropia leaves. Other paratypes were taken from the same host material, by S. L. Wood, as follows: 17 from Barro Colorado Island, Panama Canal Zone, December 27, 1963, eleva- tion about 70 m.; II from Palin, Esquintla Prov., Guatemala, May 19. 1964, elevation 350 m.; 1 from Rincon de Osa. Puntarenas Prov., Costa Rica, August 1 1 , 1966, elevation 30 m. The holotype, allotype, and paratypes are in my collection. Scolytodes iraziiensis, n. sp. Rather closely related to parvulus Wood, described above, but larger, the female frons only shallowly impressed with setae on vertex ex- tending half the distance to the antennal inser- tion, and the general sculpture coarser. Female. — Length L7 mm. (paratypes I. ,5- 1. 9 mm.). 2.4 times as long as wide; color very dark brown. Frons rather broadly, longitudinally impressed from upper level of eyes to epistoma; the lateral margins above le\el of antennal inscriion siibiarinnleh elev;Llcd almost to upper level of eyes; surface finely, closely, deeply punc- tured: vestiture mostly confined to vertex, rather long, moderately abundnat (largely abraded in type), none of .setae long enough to reach half the distance to level of antennal insertion, only a few setae arise below upper level of eyes. Pronotum 1.0 times as long as wide; sides weakly arcuate anerdiciis except granules in psoterior area reduced and setae finer. Elytra 1.5 times as long as wide. 1.6 times as long as pronotum; outline as In verdicus; striae not impressed, the punctures moderately large, shallow, distinct; inter- striae wider than striae, surface somewhat irregularly marked by lines, the punctures fine, uniseriate. becoming granulate near declivity. Declivity confined to posterior third, rather steep, convex; strial punctures greatly re- duced above, obsolete below; Interstriae 1 devoid of granules on lower three-fourths and 2 on lower half, upper half of 2, all of 3 and 4 and lateral areas armed by moder- ately large, somewhat pointed, isolated, squamiferous granules. Vestiture consisting of erect scales; each scale on disc about four times as long as wide, almost equal in length to distance between rows, more slender, one and one-half times as long and more widely spaced than on dsic; spacing of scales within row on declivity slightly greater than spacing between rows. Female. — Similar to male except frontal concavity slightly deeper and wider; anterior margin of pronotuin 28 unarmed; elytra 1.7 times as long as wide, the surface somewhat more irregular; elytral declivity sculptured to apex as on disc; interstrial scales about six times as long as wide, not longer or more widely spaced on declivity. Type Locality. — Twenty-seven km. west of Durango, Durango, Mexico. Host. — Phoradendron villosum. Type Material. — The male holotype, female allotype, and 41 paratypes were collected on June 4, 1965, about 2,000 m.. No. 12, by S. L. Wood, from tunnels in the phloem of the above named mistletoe. The holotype, allotype, and paratypes are in my collection. Pseudothysanoes peniculus, n. sp. This s])ccics is rather closely allied \api»ebris Wood, described above, but it is readily distin- guished, in the male, by the continuation of all rows of interstrial scales to their normal termina- tion on the declivity, and in the female, by the more extensively concave frons that is orna- mented above by a very long tuft of yellow hair. Male. — Length 1.4 mm. (paratypes 1.4-1.7 mm), 2.3 times as long as wide; color dark brown, part of elytra and summit of pronotum lighter in some specimens. Frons rather strongly, broadly, transversely impressed on lower half, convex above, the epistoma giadually raised to Mungin; surface finely granulale piuulale. liuci in impression; vestiture consisting of moderately abundant, short, subplumose setae. Pronotum 0.96 times al long as wide; outline as in verdicus; surface behind summit rather coarsely reticu- late, the minute, sparse punctures obscure; vestiture of scales and stout hair intermixed. Elytra 1.4 times as long as wide. 1.5 times as long as pronotum; outline as in verdicus: striae not impressed, the punctures small, shallow, obscure; interstriae about as wide as striae, irregularly marked, the punctures fine, obscure, very feebly granulate, squamiferous. Declivity on slightly more than posterior third, convex, moderate- ly steep; sculpture essentially as on disc. Vestiture of rows of erect scales, each about four times as long as wide at base, at least eight times as long as wide on declivity, each slightly longer than distance between rows at base, in- creasing in length toward declivity until about twice as long as distance between rows on declivity, the .spacing within each row equal to or slightly less than length of scale. Female. — Similar to male extepl frons concave from eye to eye, from epistoma to vertex, area above eyes orna- mented by a dense brush of long, yellow hair some of which extends to epistomal luargin; scape with a rather large fringe of long hair; anterior margin of pronotum unarmed; elytra 1.6 limes as long as wide, declivity con- fined to posterior third; elytral surface, particularly on declivity, more nearly minutely rugulose; elytral scales shorter (about two thirds as large) and closer. Type Material. — Thirteen km. west of Tex- melucan, Puebla, Mexico. Host. — Phoradendron sp. Type Material. — The male holotype, female Brk.ham Young Univkrsity Science Bulletin allotype, and nine paratypes were collected at the type locality on June 13, 1967, 2,600 m., No. 26, by S. L. Wood, from tunnels in the phloem of the above mistletoe. The holotype allotype, and paratypes are in my collection. Pseudothysanoes bullatus, n. sp. This species is rather closely allied to peni- culus Wood, described above, but it is easily distinguished, in the male, by the presence of a rather large bulla on the lower half of declivital interstriae 3 and by the broader elytral scales, and, in the female, by the shallowly concave frons with vei7 short hair above. Male. — Length 1.5 mm. (paratypes 1.5-2.0 mm.), 2.3 times as long as wide; color very dark brown, posterior half of elytra usually lighter. Frons moderately, transversely impressed on lower half, convex above; surface rather coarsely punctate- granulate above, finely rugulose on lower third; vestiture consisting of stout, rather sparse, short, subplumose setae. Scape with about a dozen long setae. Pronotum 0.94 times as long as wide; widest at base, the sides arucately converging from base to narrowly lounded anterior margin, with a slight constriction one- third from anterior margin; anterior margin armed by six small teeth; summit behind middle, well developed, broad; posterior area very finely rugulose-reticulale, the minute punctmes obscure; vestiture of scales and hair intermixed. Elytra 1.4 times as long as wide, 1.6 times as long as pronotum; outline as in verdicus; striae not impressed, the punctures rather small distinctly impressed; interstriae wider than striae, minutely, inegularly marked by obscure lines and points, the squamiferous punctures rather fine, usually distinct, not granulate. Declivity occupying slight- ly more than posterior third, convex, rather steep; strial and interstrial punctures reduced, becoming obsolete by iniddle of declivity; points largely replace obscure lines on interstriae. the surface shining; interstriae 3 with a large, low bulla on upper part of lower half of declivity, extending slightly to interstriae 2 and 4. Vestiture con- sisting of erect, interstrial scales, each about two to three times as long as wide, slightly longer toward declivity, those on upper declivity spaced within and between rows by a distance equal to length of a scale; lower two-thirds of declivity glabrous. Female. — Similar to male except frons shallowly con- cave on median three-fourths from epistoma to above upper level of eyes, the upper area bearing rather numer- ous, moderately short, yellow setae; anterior margin of pronotum unarmed and more broadly rounded; declivital interstriae each bearing a uniseriate row of fine granules to apex, the bulla obscure; declivity shorter, steeper; vestiture more abundant, extending to apex, each scale more slender, about five times as long as wide. Type Locality. — Forty-two km. southeast of Nochixtlan, Oaxaca, Mexico. Host . — Phoradendron sp. Type Material. — The male holotype, female allotype, and nine paratypes were collected at the type locality on June 17, 1967, about 2,300 m.. No. 55, by S. L. Wood, from tunnels in the New Neotropical Scoi.ytidae 29 phloem of the above mistletoe. The holotype, allotype, and paratypes are in my collection. Pseudothysanoes plumalis, n. sp. This species evidently belongs to the verdicus Wood, described above, group of species, al- though it is not closely related to any of those treated here. The female frons is feebly concave and not ornamented by long hair; the scape bears a large tuft of hair. The male elytral sculpture is simple and quite unlike related species. Male. — Length 1.4 mm. (paratypes 1.4-1.7 mm.). 2.4 tiines as long as wide; color dark brown. Frons shallowly concave almost to upper level of eyes, convex above; surface rugose-reticulate with a few fine granules in convex area; vestiture sparse, incon- spicuous. Scape with fewer than a dozen setae. Pronotum 0.91 times as long as wide; outline as in nerilicui; posterior area reticulate, the very fine punctures obscure; vestiture of fine and stout hairlike setae. Elytia 1.4 times as long as wide. 1.65 times as long as pronotum; sides straight and parallel on basal two- thirds, rather broadly rounded behind; striae not im- pressed, the punctuies rather coarse, deep; interstriae narrower than striae, almost smooth, the punctures very fine, uniseriate. Declivity largely confined to posterior third, convex, steep; strial punctures greatly reduced, the interstrial punctures very finely granulate. Vestiture consisting of rows of small, erect, interstrial scales of about equal size on disc and declivity; each scale about three times as long as wide, about one-half as long as distance between rows of scales, spaced within rows by distances equal to about three times length of a scale. Female. — Similar to male except frons more broadly, somewhat more deeply concave; scape bearing a large, dense tuft of long, yellow hair; pronotuiii armed by two teeth on anterior margin; declivity slightly shorter and steeper; declivital interstrial granides obsolete. Tyf)(' Locality. — Eighteen km. north of Hua- juapan, Oaxaca, Mexico. Typf Material. — The male holotype, female allotype, and 10 paratypes were collected at the type locality on June 15, 1967, 2,000 m., No. 43, by S. L. Wood, from tunnels in the phloem of an unusual mistletoe that awaits identification. The holotype, allotype, and paratypes are in my collection. Pseudothysanoes coracinus, n. sp. This species is very similar to hopkinsi Black- man, but it is distinguished, in the male, by the more nearly flattened frons, by the more weakly impressed strial punctures, and by the much broader interstrial scales. The female remains unknown. Male. — Length 1.2 mm. (male paratypes 1.2-1.3 mm.). 2.2 times as long as wide; color almost black, pronotal summit reddish. Frons convex above, weakly, transversely impressed just above epistoma, with a narrow impression extending dorsad to center of frons; surface rugulosereticulate; vestiture sparse, consisting of coarse, subplumose setae of moderate length. Pronotum 0.84 times as long as wide; widest a third from base; sides rather strongly arcuate on basal two- thirds then weakly constricted befoie the lather broadly rounded anterior margin; anterior margin armed by four broad, low teeth; summit high, behind middle; posterior area reticulate, the punctures sparse, minute, obscure; vestiture of stout and slender, short, hairlike setae. Elytra 1.3 times as long as wide. 1.6 times as long as pronotimi: sides almost straight and parallel on slightly more than basal half, rather narrowly lounded behind; striae not impressed, the punctures rather small, distinct but not deep; interstriae about as wide as striae, the pimctuies uniseriate, finely granulate. Declivity on slight- ly less than posterior half, convex, rather steep; as on disc except strial punctures slightly smaller, the interstrial granules very slightly higher. Vestiture consisting of minute, fine, strial hair and tows of erect interstrial scales; each scale about twice as long as wide: not longer on declivity. Type Locality. — Thirty-four km. north of Juchitlan, Jalisco, Mexico. Type Material. — The male holotype and three male paratypes were collected at the type locality on July 3, 1965, about 1,300 m.. No. 182a, by S. 1,. Wood, from tunnels in the cambium region in twigs of an unidentified tree. The holotype and paratypes are in my collec- tion. Pseudothysanoes crassinis, n. sp. This species is allied to hopkinsi Blackman, but it is not closely related. It is distinguished, in the male, by the much stouter body, by the longer, more slender interstiral scales, and by the large interstrial granules, and, in the female, by the slightly impressed frons, and by the inuch finer elytral sculpture and smaller scales, Male. — Length 0.9 mm. (paratypes: males 0.8-1.0 mm.; females 1.01.2 mm.). 2.0 times as long as wide; color black. Frons and pronotum as in coracinus (above) except teeth on anterior margin of pronotimi more slender, the summit higher and witler. and the posterior areas obscure- ly reticulate. Elytra 1.1 times as long as wide, 1.25 times as long as pronotum: sides almost straight and parallel on basal two-thirds, broadly rounded behind; striae not impressed, the punctuies obsolete: interstriae each with a uniseriate row of large tubercles, increasing in size toward declivity. Declivity beginning at middle, rather steep, convex; a few minute strial punctuies obscurely indicated; inter- strial gianules greatly reduced on upper half, obsolete on lower half, replaced by very minute, obscure punctures. Vestiture consisting of fine, moderately long, suberect, strial hair and interstrial rows of longer, erect scales; each scale about four times as long as wide on disc, six times as long as wide on declivity; each scale as long as distance between rows on disc, slightly longer on declivity, spaced within rows by a slightly gieater distance. Female. — Similar to male except larger; 2.4 times as long as wide; frontal impression very slightly larger; anterior margin of pronotum unarmed: strial punctures obscurely visible; interstrial granules obsolete: declivity 30 Brigham Young Univfrsity Science Bulletin confined to posterior third of elytra, more narrowly con- vex; interstrial stales slender, at least eight times as long as wide. Type Locality. — Twenty-four km. west of Tehuantepec, Oaxaca, Mexico. Type Material. — The male holotype, female allotype, and 63 paraiypes were collected from the type locality on June 22, 1967, 70 m.. No. 35, by S. L. Wood, from tunnels in the cambium region in the bole of a small, thornless, legumi- nose tree having bipinnately compound leaves. The holotype, allotype, and paratypes are in my collection. Ciyptulocleptus quercinus, n. sp. This if the first species in this genus known to me in which the frons of male and female are equally convex; the scape is also vei7 short and bears a large tuft of hair in the female. Male. — Length 1.0 mm. (paratypes: male 0.8-1.1, female 1.2-1.5 mm.). 2.4 times as long as wide; color very dark brown. Frons convex, narrowly, transversely impressed just above epistoma; rugiilose-reticulate, the piincliires fine; vestitiire moderately abundant, conspicuous, subplumose, moderately short. Scape only slightly longer than pedicle; bearing several long setae. Pronotum 0.81 times as long as wide; widest near middle, subcircular in outline; anterior margin armed by three small teeth (four in paratypes); summit at middle, rather high, anterior slope rather coarsely asper- ate; posterior area smooth, shining, the piuictures sparse, very minute; vestiture consisting of stout erect hair pos- teriorly, of bifid scales in asperate area. Elytra 1.6 times as long as wide, 1.9 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather narrowly rounded behind; striae not impressed, the punctures of moderate size and depth; interstriae almost smooth, as wide as striae, the punctures fine, uniseriate. Declivity rather steep, convex; essentially as on disc but surface more irregular. Vestiture consisting of fine, short, slrial hair and rows of erect interstrial scales; each scale one and one-half times as long as strial hair, about two-thirds as long as distance between rows, similarly spaced within each row, each scale about twice as long as wide. Female. — Similar to male excpet 2.8 times as long as wide; frons as in male; scape very slightly larger, bearing a large, long tuft of hair; anterior margin of pionotum unarmed; strial punctures not as deep, more widely spaced; declivity le.ss clearly punctured; elytral scales more closely spaced within rows, very slightly longer, each scale about four times as long as wide. Type Locality. — Ihirty-seven km. west of Durango, Durango, Mexico. Type Material. — The male holotype, female allotype, and 32 paratypes were collected at the type locality on June 4, 196.'j, about 2,000 m.. No. 10, by S. L. Wood, from phloem tuntiels in a small, shaded-out Qitercus branch 2 cm. in diameter. The holotype, allotype, and paratypes are in my collection. Cryptulocleptus arbnti, n. sp. This species is rather closely allied to quer- cinus Wood, described above, but it is distin- guished by the larger, median, frontal impres- sion, by the greatly reduced tuft of hair on the scape, by the rugulose-reticulate posterior areas on the pronotum, by the larger strial punctures, and by the longer, more slender interstrial scales. Male. — Length 1.2 mm. (paratypes 1.0-1.3 mm., males average slightly smaller than females), 2.7 times as long as wide; color very dark brown. Frons convex above, narrowly, rather strongly, trans- versely impressed just above epistoma, the impression exleucli'il slighily ilorsad al center; vestiture confined to epistomal margin. Scape twice as long as pedicel, bearing about six setae; club oval very slightly longer than wide, Pronotum 0.94 times as long as wide; essentially as in quercinus except posterior area minutely rugulose-reticu- late, most of the punctures minutely granulate. Elytra 1.7 times as long as wide, 1.8 times as long as pronotum; outline about as in quercinus: striae not im- pressed, the punctures rather coarse, close, not sharply impressed; interstriae as wide as striae, minutely irregular, the punctures very fine, uniseriate. Declivity steep, con- vex; strial punctures slightly reduced in size, interstrial punctures minutely granulate. Vestiture consisting of very line, moderately long slrial iKiirand slighllv longci . erect interstrial scales; each scale slender, equal m length to distance between rows, slightly closer within each row, each at least eight times as long as wide. Female. — Similar to male except 3.0 times as long as wide; upper frons bearing rather abundant short, stout setae; scape bearing about a dozen setae; anterior margin of pronotum unarmed; elytral scales a little more slender. Type Locality. — Thirteen km. west of Tex- melucan. Puebla, Mexico. Type Material. — The male holotype. female allotype, and 34 paratypes were collected at the type locality on June 13, 1967, 2,600 m., No. 27, by S. L. Wood, froin twigs of Arbutus sp. The tunnels were immediately below the surface. The holotype, allotype, and paratypes are in my collection. Ciyptulocleptus caritus, n. sp. This species is allied to arbuti Wood, described above, but it is not closely related. It differs from arbuti by the less strongly convex frons which lacks a transverse impression, by the smaller, less abundant, pronotal asperities, and by the very diffeient pronotal and elytral vesti- ture. Male. — Length 1.2 mm. (paratypes 1.2 I. .3 mm.). 2.7 times as long as wide; color dark blown. Fions convex to epistomal margin, a moderately large fovea just below center; surface reticidale in marginal areas, smooth and shining on middle half, with a few scattered, small, punctures: vestiture inconspicuous. Scape elongate; chdj small, oval. New Neotropical Scolytidae 31 Pionotum 1.0 times as long as wide: widest near base, the sides very weakly arcuate, converging slightly to shal- low constriction one-third pronotuni length from the rather narrouly rounded anterior margin; anterior margin armed by four moderately large teeth; summit at middle, moderately high; anterior slope with fine asperities; pos- terior area minutely rugulose. the minute punctures obscure; vestiture slender, hairlike in both asperate and posterior areas. Elytra 1.7 times as long as wide, 1.8 times as long as pronotinn; outline as in quercinus; striae not impressed, the punctures fine; interstriae as wide as striae, with abundant, very minute points, the punctures very fine, obscure. Declivity steep, convex, as on disc except inter- strial punctures finely granulate. Vestitme consisting of very fine long, strial hair and erect interstrial setae; each interstrial seta on disc stout, hairlike, one and one-half times as long as strial hair, slightly longer than distance between rows, more closely spaced within each row; inter- strial setae becoming scalelike on declivity and very slightly longer, each scale at least six times as long as wide. Female. — Similar to male except 2.9 times as long as wide; frons very weakly, transversely impressed just above epistomal margin; anterior margin of pionotum unarmed; interstrial granules on declivity very minute; interstrial setae on declivity stout but essentially hairlike. Type Locality. — Cerro Punta, Chiriqui, Pana- ma. Type Material. — The male holotype, female allotype, and 10 paratypes were collected January 11, 1964, 1,800 m., No. 411, by S. L. Wood, from broken branches of an unidentified tree. The holotype, allotype, and paratypes are in my collection. Cryptiilocleplus pumtlus, n. sp. This species is rather closely allied to siib- pilosus Wood, but it may be distinguished in the male by the more slender body form, and by the much smaller strial punctures, and, in the female, by the more deeply concave frons and by the limitation of vestiture in the frontal concavity to a narrow band on upper margin. Male. — Length 0.8 mm. (paratypes: male 0.8-0.9, female I.O-I.I mm.), 2.3 times as long as wide; color yel- lowish brown. Frons convex, narrowly, transversely impressed just above epistomal margin, more strongly in median aiea; surface shining, rather coarsely reticulate; vestiture con- fined to epistomal margin. Antennal club rather nar- rowly ovate; devoid of sutures. Pionotum 0.90 times as long as wide; widest a third from base, sides evenly arucate to the rather broadly rounded anterior margin; anterior margin arined by four small teeth; summit just behind middle, rather well de- veloped; asperities on anterior slope coaise; posterior area shining, minutely rugulose-reticidate, the fine functures obscure; vestitme largely scalelike. Elytra 1.4 times as long as wide. 1,6 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; striae not impressed, the punctures very small but distinct; inter- striae somewhat irregidar, shining, twice as wide as striae, the squaniiferous punctures equal in size to those of striae but more widely spaced. Declivity rather steep, convex; sculpture as on disc but all punctures very slightly smaller. Vestiture consiting of minute, fine, strial hair and rows of erect, interstrial scales; each scale twice as long as strial hair, almost as long as distance between rows, about twice as long as wide, strongly tapered toward their bases. Female. — Similar to male except 2.5 times as long as wide; frons rather deeply concave on median two-thirds to well above eyes, with a row of stout, subpluinose setae on upper margin; scape bearing a small tuft of long hair; anterior margin of pionotum unarmed; strial punc- tures equal in size but less distinct; elytral scales three times as long as wide. Type Locality. — Fifty-three km. south of Coli- ma, Colima, Mexico. Type Material. — The male holotype, female allotype, and 56 paratypes were collected at the type locality on June 27. 1965, about 700 m., No. 122, by S. L. Wood, from an unidentified desert shrub. The holotype, allotype, and paratypes are in my collection. Ciyptulocleplus mendicus, n. sp. This species is closely allied to pumilus Wood, described above, but it may be distin- guished by the smoother elytral surface, by the much more slender elytral scales, and, in the female, by the less strongly concave frons that is devoid of special setal ornamentation. Male. — Length 0.9 mm. (paratypes: male 0.9-1.0, females 1.1-1.3 mm.), 2.4 times as long as wide; color yel- lowish brown, some specimens darker. Frons as in pumilus: pronotum also as in pumitus except six teeth on anterior margin larger, the posterior area more finely reticulate. Elytra 1.4 times as long as wide, 1.6 times as long as pronotum; sides almost straight and parallel on basal two-thirds, narrowly rounded behind; striae not im- pressed, the fine punctures spaced by two diameters of a puncture; interstriae shining, marked by irregular lines, twice as wide as striae, the punctures distinctly smaller than those of striae. Declivity rather steep, convex; essentially as on disc except punctures much smaller. X'estiture consisting of very minute strial hair and rows of erect interstrial scales; each scale three times as long as strial hair, slightly shorter than distance between rows of scales, each about three times as long as wide. Female. — Similar to male except 2.9 times as long as wide; frons concave on median two-thirds to upper level of eyes, not specially ornamented by setae; scape bearing a small tuft of long hair; anterior margin of pronotum unarmed; elytral punctures much more obscure; elytral scales more slender, each about six times as long as wide. Type Locality. — Eleven km. south of Coli- ma. Colima, Mexico. Type Material. — The male holotype, feinale allotype, and 72 paratypes were collected at the type locality on June 28, 1965, about 1,000 m.. No, 120, by S. L. Wood, from an unidentified thornless, leguininous, desert, shrubby tree. Additional paratypes were taken in Mexico as follows: 19 at 3 km. W. Armeria, Colima, June 32 Bricfiam Young University Science Bulletin 28. 1965. 70 m., No. 137. S. L. Wood; and 17 at 24 km. W. Tehuatepec, Oaxaca. June 22. 1967, 70 m., No. 85, S. L. Wood; presumably all taken from the same host species. The holotype. allotype, and paratypes are in my collection. Ctyptulocleptus acares, n. sp. This species is rather closely related to men- dicus Wood, described above, but it is distin- guished by the larger size, by the more irregular, more deeply punctured elytra, and by the larger, more closely spaced interstrial scales. Male. — Length 1.2 mm. (paratypes: males. 1.1-1.2. females 1.3-1.4 mm.). 2.2 times as long as wide; color yel- lowish brown, the elytra darker. Frons and pronotum as in mendicus, except posterior area of pronotum almost smooth and shining. Elytra 1.3 times as long as wide. 1.5 times as long as pronotum; outline about as in mendicus; striae not impressed, the punctures rather coarse; interstriae as wide as striae, the punctures rather large, uniseriate. Declivity steep, convex; sculpture about as on disc except interstrial punctures closer. Vestiture consisting of minute strial hair and rows of erect interstrial scales; each scale two- thirds as long as distance between rows, similarly spaced within rows; each scale twice as long as wide. Female. — Similar to female of mendicus except 2.7 times as long as wide; frons somewhat more deeply im- pressed, with a few minute granules in upper third of impression; strial and interstrial punctures closer and more distinct, the general surface more irregular; declivity much more coarsely, deeply punctured; interstrial scales much closer, each four to six times as long as wide, very slightly longer. Type Locality. — Thirty km, southeast of El Cameron. Oaxaca, Mexico. Type Material. — The male holotype. female allotype, and 25 paratypes were collected at the type locality on June 21, 1967, near 1,000 m.. No. 80, by S. L. Wood, from the branch of an uniden- tified tree (probably leguminose). The holotype, allotype, and paratypes are in my collection. Cryptulocleptus acjuilus, n. sp. This species is allied to mendicus Wood, described above, but it may be distinguished by the slightly larger size, by the darker color, by the much more boardly rounded anterior margin of the pronotum, by the inuch coarser male strial punctures, by the much more coarsely punctured temale elytral declivity, and by the more exten- sive female front;d impression. Male. — Length 1.1 mm. (female paratypes 1.2-1.4 mm.). 2.3 times as long as wide; color dark brown. Frons and pronotum as in mendicus except anterior margin of pronotum much more broadly rounded, bearing eight teeth, and posterior area more clearly punctured. Elytra 1.4 times as long as wide, 2.1 limes as long as pronotum; outline as in mendicus; striae not impressed, ihe punctures rather coarse, round, deep, interstriae as wide as striae, almost smooth, the punctures minute, uniseriate. Declivity rather steep, convex; strial punc- tures decreasing rapidly in size, almost obsolete at apex; interstrial punctures minute. Vestiture consisting of fine, minute, strial hair and rows of erect interstrial scales; each scale almost as long as distance between rows, each about four times as long as wide. Female. — Similar to male except 2.7 times as long as wide; frons shallowly concave from epistoma to vertex, smooth and shining on median half below, not specially ornamented by setae; anterior margin of pronotum un- armed; elytral surface more irregular, the strial punctures smaller, but distincl; interstrial punctures minute, mostly obscure; declivity more irregular, more coarsely punctured much as in acares: interstrial scales more closely placed. Type Locality. — Twenty-six km. east of Mo- relia, Michoacan, Mexico. Type Material. — The male holotype, female allotype, and 21 female paratypes were collected at the type locality on June 14, 1965, 2,500 m.. No. 56, by S. L. Wood, from phloem tunnels in an Acacia branch. The holotype, allotype, and paratypes are in iny collection. C)-yptulocleptus minis, n. sp. This species is rather closely related to acares Wood, described above, but it may be distin- guished by the more deeply impressed feinale frons above, by the more coarsely sculptured pos- terior areas of the pronotum, and by the much finer declivital punctures. Male. — Length 1.0 nmi. (paratypes: male 1.0-1.1. female 1.2-1.4 mm.). 2.2 times as long as wide; color yel- lowish brown. Frons similar to acares but more strongly convex. Pronotum as in acares except reticulation in posterior areas tending to be very slightly more rugulose. Elytra 1.3 times as long as wide, 1.4 times as long as pronotum; outline as in acares; striae not impressed, the punctures rather coarse, more sharply impressed than in acares; interstriae inegular but much smoother than in acares, as wide as striae, the punctures rather small and more widely spaced than in acares. Declivity steep, con- vex; surface smoother, the punctures very slightly smaller than on disc. Vestiture as in acares except interstrial scales very slightly more widely spaced. Female. — Similar to female of acares except 2.8 times as long as wide; frons more abruptly impressed at upper limits of concavity; posterior areas of pronotum more strongly reticulate; elytral surface less rugulose; declivital punctures reduced to very fine strial and interstrial punc- tures; elytral scales similar, but less abundant. Type Locality. — Thirty-eight km. south of Matias Romero, Oaxaca, Mexico. Type Material. — The male holotype, female allotype, and 40 paratypes were collected at the type locality on June 24, 1967, 250 m., No. 92, by S. L. Wood, from branches 1 to 5 cm. in diameter of a leguminose shrub having a yellow, mimosa- like flower. The holotype, allotype, and paratypes are in my collection. New Neotropical Scolytidae Cryptulocleptus obesus, n. sp. Appareiuly this species is more closely allied to colurnbiuuus Blackmail than to other known species, but it is distinguished by the stouter body, shorter antennal scape; and by the less closely placed interstrial scales. Male. — Length 1.0 mm. (paratypes: male 1.0-1.2, female 1.2-1.4 mm.), 2.0 times as long as wide: color red dish brown, the elytra black (lighter in less mature speci- mens). Frons planoconvex almost to upper level of eyes: minutely punctate above, becoming smoother below; vesti- ture consisting of a few short subphimose setae over flat- tened area. Scape short, little longer than pedicel, bearing a small tuft of hair: club moderately large, oval. Pronotum 0.86 times as long as wide; widest one- third from base, sides rather strongly arcuate converging toward the narrowly rounded anterior margin; anterior inargin armed by four teeth, the median pair larger; sum- mit behind middle, well developed; anterior area coarsely asperate; posterior area very minutely rugulose, the punc- tures obscure; vestitiire largely scalelike. Elytra 1.15 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on slightly more than basal half, rather broadly rounded behind; striae not impressed, the punctures small, obscure, inter- striae wider than striae, the punctures fine, uniseriate. Declivity steep, broadly convex; surface minutely rugulose, as on disc except interstrial punctures closer. Vestiture consisting of interstrial rows of erect scales, each scale less than twice as long as wide. Female. — Similar to male except 2.3 times as long as wide; frons shallowly concave to upper level of eyes; scape larger, broadly triangular, bearing a large tuft of long hair; anterior margin of pronotum unarmed; elytra minutely rugulose, all punctures obscure. Type Locality. — Thirty km. southeast of El Cameron, Oaxaca, Mexico. Type Material. — The male holotype, female allotype, and 26 paratypes were collected at the type locality on June 21. 1967, near 1,000 m.. No. 79, by S. L. Wood, from twigs of a tree awaiting identification. The holotype. allotype, and paratypes are in my collection. Cryplutoc leptus excavatiis.n. sp. This remarkable species is unique in the genus. It is distinguished by the remarkably extensive frontal excavation of both sexes; its larger size and the short, tufted scape also aid in distinguishing it. Male. — Length 1.4 mm. (paratypes: male 1.6, female 1.4-1.6 mm.). 2.1 times as long as wide; color dark brown to black. Frons verv broaillv. deeply, (ouiavciv e\ca\;iic(l from eye to eye. trom epistoma to vertex, the upper margin acute with median fourth projecting cephalad slightlv: surface reticulate, glabrous. Mandibles near midtlle armed by a transverse, strongly developed ridge. Antennal scape slightly longer and twice as wide as pedicel; club twice as long as wide. Pronotum 0.89 times as long as wide; widest two- fifths from base, the sides and anterior margin equally rounded on an almost circular arc; anterior margin armed by two small teeth; summit at middle, broad; anterior slope veiy coarsely asperate; posterior areas almost smooth, obscurely reticulate, shining: small scales almost obsolete except at margins. Elytra 1,2 times as long as wide, 1.4 times as long as pronotum; widest at base, the sides straight on basal two- thirds, very feebly converging posteriorly, broadiv roinided behind; striae not impressed, the punctuies small, shallow, .somewhat obscure; interstriae wider than striae, minutely subrugulose, shining, the punctures fine, obscure. Decliv- ity steep, convex: slrial ptmcttnes greatly reduced, becom- ing obsolete; interstrial punctures finely granulate. \'esti- ture consisting of minute, fine, strial hair and rows of erect interstrial scales; each scale on disc slender, elongate, slightly longer than distance between rows, becoming shorter but not narrower toward declivity, on declivity each scale equal in length to little more than half distance between rows, about twice as long as wide. Posterior margin of sternum 5 narrowly rounded, projecting slightly. Female. — Similar to male except 2. .3 times as long as wide; frontal excavation slightly deeper with median area near epistoma slightly elevated; mandibular ridge absent; scape wider, bearing a small tuft of hair; margin of pro- noliim unarmed; di.scal scales on eyltra slightly shorter; stern tun 5 less strongly produced. Type Locality. — Thirty km. southeast of El Cameron, Oaxaca. Mexico, Type Material. — The male holotype, female allotype, and three paratypes were collected at the type locality on June 21, 1967, near 1,000 m., No. 78, by S. L. Wood, from the twigs of the same tree that harbored the preceeding species, obesus. riie holotype, allotype, and paratypes are in my collection. Cryptulocleptus spicatus, n. sp. This species and the next one form a new species group within the genus. They differ from all representatives of the genus in having the last visible male abdoininal tergum very nar- row, produced medially into a projecting mucro; in the female a similar projecting mucro is formed by the last visible sternum; the female frons is also concave. Male. — Length 1.7 mm. (paratypes 1.5-1.8 mm.). 2.4 times as long as wide; color dark brown, summit of prono- tum reddish brown. Frons convex above, becoming more nearly flattened on lower third; surface rugulose-reticulate and obscurely punctuied; an indeifinite, median, subfoveate impression just below upper level of eyes; vestitiue fine, short, in- conspicuous except long and conspicuous along epistomal margin. Scape elongate. Pronotum 0.90 times as long as wide; widest a third from base, sides arcuate, a weak constriction on anterior third, rather broadly rounded in front; anterior margin armed by about eigth small teeth; siuiimit just behind middle, moderately high; asperities on anterior slope rather large; posterior area rugidose-reticulate, a few fine granules behind summit: vestiture hairlike. 34 Brigham Young University Science Bulletin Elytra 1.5 times as long as wide. 1.6 times as long as pronotum: sides almost straight and parallel on basal two-thirds, rather broadly rounded behind: striae not impressed, the punctures very small, obscure; interstriae almost twice as wide as striae, very finely rugulose. the punctures very fine, their anterior margins very finely gianulate. Declivity convex, steep; sculpture essentially as on disc. Vestiture consisting of fine, moderately long strial hair and rows of slightly longer interstrial bristles; each bristle about two-thirds as long as distance separating rows, separated within rows by distances equal to length of bristle, bristles becoming scalelike on declivity, each scale there at least three times as long as wide. Last visible tergum narrowly produced into a iTuicro-like process that projects beyond elytral apex. Female. — Similar to male except 2.6 times as long as wide; frons shallowly concave to vertex, lower, median half smooth, shining and ascending toward epistomal margin; anterior margin of pronotinn iniarmed; interstrial setae on disc fine, hairlike, bristlelike on declivity; mucro formed by last visible sternum; last tergum rounded on posterior margin. Type Locality. — Cerro Puma, Chiriqui, Pana- ma, Type Material. — The male holotype, female allotype, and 48 paratypes were collected at the type locality on January 11, 1964, 1,800 m.. No. 382, from cambium tunnels in an inidientified log about 1 m. in diameter. The holotype. allotype, and paratypes are in my collection. Cryptulodeplus subulatus, n. sp. This species is rather closely related to spicatus Wood, described above, but it may be distinguished by the much larger and deeper strial punctures, by the longer, broader, elytral scales, and by the much more strongly impressed female frons. Mate. — Length 1.4 mm. (paratypes 1.3-1.7 mm.). 2.3 times as long as wide; color very dark brown to black, pronotal summit reddish brown. Frons and pronotum as in spicalus except part of pronotal vestiture stout. Elytra 1.5 times as long as wide. 1.6 limes as long as pronotum; outline as in spicatus; striae not impressed, the punctures rather large, deep, sharply impressed; inter striae almost smooth, shining, one and one-half times as wide as striae, the uniseriate punctures finely granu- late. Declivity steep, convex; striae very feebly impressed, the punctures more obscure. X'esiiture consisting of shoil, fine strial hair (some flattened on declivity), and rows of longer, interstrial scales; each scale slightly more than half as long as distance between rows, separated within rows by distances equal to length of scale, each about twice as long as wide. Tergal mucro about as in spicatus. Female. — Similar to male excpet 2.6 times as long as wide; frons rather deeply concave on median three- fourths from epistoma to vertex, a small, smooth, median area on lower half, most females ornamented by a small, sparse tuft of hair at upper margin of concavity; anterior margin of pronotum unarmed; strial punctures not as deep, perhaps smaller; interstrial scales very slightly longer and more slender, each scale about four times as long as wide; tergal muao replaced by sternal mucro as in spicatus. Type Locality. — Volcan Pacaya, Esquintla, Guatemala. Type Material. — The male holotype, female allotype, and bl paratypes were collected at the type locality on June I, 1964, 1,300 m.. No. 700, by S. L. Wood, in tunnels in the phloem of branches less than 8 cm. in diameter of a tree named Caldo de Frijol by a local woodcutter. The local host name obviously was derived from the odor of freshly ciu wood. The holotype, allotype, and paratypes are in my collection. Cryptulodeplus miicronatus, n. sp. While most of the generic characters of Cryp- tulocleptus clearly fit this species, it is unique in having the elytral apices acuminately extended to form a short mucro as seen in Micracis or pos- sibly Micracisella. The mucro appears to be a secondary acquisition that should not have a bearing on the generic placement of this species. Male. — Length 1.3 mm. (paratypes 1.2-1.4 mm.), 2.5 times as long as wide; color dark reddish brown. Frons convex, with a feeble impression just above epistoma; surface rather coarsely rugulose above, more finely below, with a small, meidan, smooth area on lower third; vestiture short, sparse. Scape elongate, with fewer than a dozen setae; club broadly oval, unmarked by any indication of sutures. Pronotum 1.0 times as long as wide; widest two-fifths from base, the sides weakly arcuate, moderately constricted a fourth of pronotum length behind the broadly rounded anterior margin; anterior margin armed by four minure teeth; summit at middle, moderately high; anterior slope armed by many small asperities; posterior areas rather coarsely reticulate, with moderately abundant, fine, iso- lated granules behind summit; vestiture of stout, hairlike .setae. Elytra 1.5 times as long as wide. 1.5 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind except for the \ery short, sutural mucro; striae not impressed, the punctures fine, distinct; interstriae minutely subrugulose, wider than striae, the punctures small, obscure, becoming granulate toward declivity. Declivity convex, very steep; strial punctures larger than on disc but very obscure; interstrial punctures replaced by rather large rounded granules. Vestiture consisting of very fine, moderately long strial hair and rows of erect interstrial .scales; the scales slightly shorter and more slender on disc; each scale on declixity spaced within and between rows by distance equal to length of scale, each about four times as long as wide. Apex with a \ery short mucro. Female. — Similar to male except 2.6 times as long as wide; frons less strongly convex, less coarsely rugulose, with median fovea near center; anterior margin of prono- nun unarmed; elytral scales more slender, at least six times as long as wide on declivity. Type Locality. — La Lima, Cortez, Honduras. Type Material. — The male holotype, female allotype, and 44 paratypes were collected on May 5, 1964, 70 m,. No. 572, by S. L. Wood, from tun- nels in the xylem of stems 1 cm. in diameter from an unidentified shrubby, woody vine. New Neotropical Scolvtidae The holotype. allotype, and paratypes are in my collection. Thysanoes neotropicalis, n. sp. This species is more closely related to mexi- canus Wood than to others in the genus, but it is larger, more slender, more finely striate, and it has the female frons more strongly impressed. The is the first species of Thysanoes reported south of Mexico. Female. — Length 2.1 mm. (paratype.s: females 2.0-2.2. males 1.51.9 mm.). 3.2 times as long as wide: color medium brown, pronotuni lighter on type (only). Frons broadly, .somewhat transversely impressed from epistoma to upper level of eyes, the central area more narrowly impresseil. subfoveate; epistomal area smooth and shining to siibfoveale impression, sides anil above finely, obscurely punctured and finely, closely, subacicu- lately granulate; vestitme inconspicuous. Tuft of setae on scape much smaller than in rnexicanus. Pronotum 1.2 times as long as wide; slightly wider near middle, the sides straight and diverging on basal half, converging slightly anteriorly toward (he broadly rounded subserrate anterior margin; summit almost one- third from anterior margin: posterior area reticulate and very finely punctined; vcstilure confined to marginal areas consisting of flattened bristles. Ehlra 2.1 times as long as wide 1.8 times as long as pronotum; sides straight and parallel on basal two-thirds, rather broadly rounded behind; striae not impressed, ihc punctures small, shallow: interstriae about twice as wide as striae, subshining, irregular but not clearly marked b\ lines or punctures. Declivity convex, steep; surface dull, strial punctures smaller than on disc, perhaps deeper: interstriae with occasional fine granules. Vesti- tme confined to declivity, consisting of erect inlerstrial scales with an additional, partial row at suture; each scale three to four times as long as wide. Male. — Similar to female except smaller. 1.8 mm., stouter, 26 times as long as wide; frons less extensively impressed: anterior margin of pronotinn armed by ten serrations; elytra with strial punctures much larger and deeper, the interstrial punctures uniseriately gianulate toward declivity, both strial punctures and moderately large interstrial granules continuing to apex of declivity; decli\ital vestiture longer and extending well onto disc. Type Locality. — San Ignacio de Acosta, San Jose Prov., Costa Rica. Host. — Calliandra confusa. Type Material. — The female holotype, male allotype, and 22 paratypes were collected at the type locality on July 5, 1963, at an elevation of 1,500 m., by S. L. Wood, from dying branches of the above host. The tunnels were in the wood as in other species of the genus. The holotype, allotype, and paratypes are in my collection. Thysanoes subsuhalus, n. sp. This species is allied to fimbraticornis Le- Conte, but it is distinguished by the larger size, by the more deeply impressed female frons, by the rather large interstrial declivital granules. 35 and by the much longer, more slender declivital scales. Female. — Length 2.2 mm. (paratypes 1.9-2.3 nun.), 3.0 times as long as wide; color dark reddish brown. Frons shallowly, broadly concave from epistoma to vertex, with a small, median impression near upper level of eyes; surface strongly reticulate, almost subrugulose, except transversely etched on a rather large area above epistomal margin; vestitme short, stout, .spaise, uniformly distributed. Scape with a small fringe of hair. Pronotum 1.2 times as long as wide; sides straight and parallel on slightly less than basal two-thirds, very broadly rounded in front, anterior margin unarmed; summit well in front of middle, broad; surface strongly reticulate, the punctures minute, very feebly gianulate behind sumniil; vestiture fine, hairlike in posterior areas, stout bristles in asperate areas. Elytra 2.0 times as long as wide, 1.9 times as long as pronotinn; sides straight and parallel on slightly more than basal two-thirds, broadly rounded behind: striae not impressed, the punctures very fine, shallow, distinct; interstriae more than twice as wide as striae, minutely irregular, the punctures fine, obscure. Declivity steep, convex: strial punctures almost obsolete; interstriae 2 slightly elevated, each interstiiae uniseriately. rather finely gianulate, except punctures and granules obsolete on lower three-fourths of I. Vestiture consisting of very minute, fine, strial hair, and uniseriate rows of longer, elect, interstrial scales; each scale on disc about half as long as distance between rows and three times as long as wide, on declivity slightly longer than distance be- tween rows and up to six times as long as wide, obsolete on middle half of ilecli\ ital interstriae I. Male. — Similar to female except frontal concavity deeper, exteniling only to upper level of eyes: anterior margin of pronotum armed by eight small teeth; discal strial punctmes slightly larger, interstrial punctures gianulate on posterior third; declivital granules much larger; declivital scales slightly wider and longer. Type Locality. — Eight km. west of Tulancin- go, Hidalgo, Mexico. Type Material. — The female holotype, male allotype, and eight paratypes were collected at the type locality on June 11, 1967, 2,400 m,, by S. 1.. Wood, fiom Qiiercus branches. The iiolotype, ;illotype, and paratypes are in my lolkiiiou. Thysayioes epicharis, n. sp. This species is allied to fimbraticornis Le- Conte, but it is distinguished by the less deeply impressed, more strongly reticulate female frons, by the more coarsely sculptured elytral disc, by the almost obsolete declivital striae, and by the much broader scales on the elytral declivity. Female. — Length 1.2 mm. (paratypes 1.8-2.2 mm.), 3.0 times as long as wide; color yellowish brown. Frons rather strongly, transversely impressed from epistomal margin to well above eyes, a small median im- pression just below center; entire surface rather strongly rugulose-reticulate; vestiture fine, short, sparse. Scape bearing a rather dense fringe of long hair. Pronotum 1.08 times as long as wide; sides almost straight and parallel on basal half, distinctly constricted on anterior half, bioadlv louniled in front; anterior 36 Brigham Young Univfrsitv Science Bulletin margin unarmed; siimmil in front of middle; posterior area reticulate, the punctures moderately abundant, rather small, shallow; vestitiue of fine hair and stout bristles, rather short. Elytra 1.8 times as long as wide. 1.7 limes as long as pronotum; sides almost straight and parallel on basal three-fourths, bioadly roiuided behind; striae not im- pressed, the punctures small, distinct; interstriae twice as wide as striae, inegularly subriiglo,se, the ptinctures small, obscure. Declivity convex, steep, confined to posterior fourth; striae very weakly impressed, the punctures minute, distinct; interstriae almost smooth, with very fine, uniseriate, squamifeious granules, those on 3 very slightly larger. Vestiture of minute strial hair and uni- seriate rows of erect interstrial scales; each scale on disc half as long as distance between rows, three times as long as wide; declivital scales three-fourths as long as distance between rows, two to three times as long as wide; a partial supplemental low along sutuie. Male. — Similar to female except frontal impression reduced, not reaching upper level of eyes, the central mipression moie pronounced, sculpture of tipper half granulate; anterior margin of pronotum armed by six teeth; declivital granules very slightly larger. Type Locality. — Twenty-four km. south ol Mazaniitla, Jalisco, Mexico, Type Material. — The female holotype, male allotype, and eight paratypes were collected at the type locality on June 22, 1965, 2,500 m., by S, L, Wood, from branches of a leguminose tree. The holotype, allotype, and paratypes are in my collection. Thysanoes adonis, n, sp. This species is allied to mexicanus Wood, but it is distinguished by the larger size, by the sub- foveate frons, by the absence of interstrial gran- ules on basal half of disc, and by longer and much more slender interstrial declivital scales. Female. — Length 2.0 muL (paialypes 1.4-1.9 mm.), 2.6 times as long as wide; color dark brown, sirnmii of pronotum reddish brown. Frons convex above, transversely Impressed on lower half of area below upper level of eyes, a small metlian, stibloveate impression at center, ascending lowaixl episto- mal margin; surface densely, minutely, e\ idently subrugii- losely puncturetl in convex aiea, becoming almost smooth toward epistoma; vestiture of sparse, moderately long, very stout bristles uniformly distributed on convex area. Scape bearing about a dozen long hairs. Pronotum 1.04 times as long as wide; sides almost straight and parallel on basal half, moilerately constricted on anterior half, lather broadly lountled in front; anterior margin siibserrate; summit at middle, rather broad; aspe- rities on anterior slope rather coarse; posterior areas reticulate, the punctures minute, sparse, a few behind summit minutely granulate; vestiture of stout bristles. Elytra 1.6 times as long as wide, 1.6 times as long as proiiolum; sides almost straight and parallel on basal three-lourths. rather broadly loumled behind; striae not impressed, the punctures small, distinct; interstriae about Ihiec times as wide as striae, the surface iircgular, siib- rugulose, the punctures .small, uniseriate, finely gianulate on posterior lialf. Declivity convex, steep, restricted to posterior fourth; strial punctures obsolete, except some of lluiii very finelv granulate; interstriae tiniseriately, finely gi. Ululate; general surface somewhat dull.- Vestiture of \eiv minute strial hair, and uniseriate rows of much longer, interstrial scales; each scale on disc about two- thirds as long as distance between rows, about four times as longer as wide; scales on declivity slightly longer than distance between rows, about six times as long as wide, with a smaller supplemental low at suture. Male. — Similar to female except anterior margin of pronotum more distinctly serrate; elytral granules smaller, llic declivital strial punctures minute but visible; declix ilal stides broader, about four to five times as long as wide. Type Locality. — Twenty-nine km. west of Quiroga, Michoacan, Mexico, Type Material. — The female iioloiype, male allotype, and seven damaged paratypes were col- lected at the type locally on June 17, 1965, 2,200 m,, by .S. L, Wood, from (htercus Ijranches. The holotype, allotype, and paratypes are in my collection, Micracisella monudis, n. sp. This species is perhaps more closely allied to hondurensi.s Wood than to other known species, but it is easily distinguished by the very different frontal sculpture, by the rows of rounded inter- strial granules on the declivity, and by the dis- tinctly elevated declivital interstriae 9. Female. — Length 1.7 mm. (paratypes 1.5-1.7 mm.), 2.9 times as long as wide; color dark brown. Frons planoconvex from epistoma to well above eyes, the epistomal margin slightly elevated toward center; surface very minutely, longitudinally etched, approaching reticulation laterally; glabrous. Scape more than three tiines as long as pedicel, bioad. bearing a large tuft of hair. Pronotum I.I times as long as wide; widest at middle, sides weakly arcuate on posterior two-thirds, weakly con- stricted on anterior third before the rather narrowly rounded anterior margin; anterior margin armed by two basally contiguous, small teeth; summit at middle broad; posterior areas coarsely reticulate with a few minute gianules near and behind summit; a few scales on mar- ginal areas. Elytra 1.7 times as long as wide. 1.5 times as long as pronotum; sides almost straight and parallel on more than basal two-thirds, narrowly rouiuletl behind; striae not impressed, the punctures very small, distinct; interstriae marked by lines, shining, three times as wide as striae, the punctures fine, obscure. Declivity convex, rather sleep; strial punctures larger, deeper, very obscure; interstrial punctures replaced by small rounded granules. Apex of elytra subacuminate. Vestiture consisting of very .short, fine, strial hair and rows of longer, erect, interstrial scales; each scale slightly shorler than distance between rows, similarly spaced within rows, each scale about four times as long as wide, gradually, strongly narrowed to- ward their bases. Male. — Similar to female except frons convex, a narrow, transverse impression just above epistoma, the surface finely rugose-reticulate, a few short, stout setae in lateral areas; tuft of hair on scape smaller; scape more slender; teeth on anterior margin of pronotum larger; declivital scales very slightly wider. Type Locality. — Volcan C.olima, Jalisco, Mexico. Host. — SlrulluDillnis veuelus. New Neotropical Scolytidae Type Material. — The holotype, allotype, and seven paratypes were collected at the type locality on June 23, 1965, 2,500 m.. No. 105, by S. L. Wood, from axial pith tunnels in twigs of the above named mistletoe. The holotype, allotype, and paratypes are in my collection. Micracisellu scilula, n. sp. This species is allied to monadis Wood, de- scribed above, but it is readily distinguished by the strongly convex frons, by the smaller, less numerous declivital interstrial granules, by the wider declivital scales, and by the presence of four teeth on the anterior margin of the pro- notum. Male. — Length 1.(3 mm., 3.0 times a.s long a.s wide; color reddish brown. Frons rather strongly convex, with a narrow, trans- verse impression just above epistomal margin: surface rather coarsely reticulate except almost smooth on small median area on lower half, a few very small granules scattered on upper half; vestiture sparse, very short in convex area, longer below. Pronotum 1.2 times as long as wide; widest near middle, the sides very feebly arcuate, almost parallel on more than basal half, rather narrowly rounded in front; anterior margin armed by four small teeth, median pair slightly larger; summit broad, near middle; posterior area coarsely reticulate, a few shining points evidently represent obsolete punctures; vestiture reduced to a few very stout bristles in marginal areas. Elytra 1.8 times as long as wide. 1.6 times as long as pronotum; sides almost straight and parallel on slightly more than basal two-thirds, rather broadly romided behind except subacuminate at suture; striae not im- pressed, the punctures small, distinct, anterior margins ol some of them very feebly elevated; interstriae shining, subrugulose, the punctures two-thirds as large as those of striae, shallow, some obscmc. Declivity convex, rather steep; strial punctures somewhat obscure; interstrial punctures on upper half replaced by rounded gianules; subreticulale toward suture near apex. Vestiture largelv abraded on disc of type, evidently shorter than on de- clivity; on declivity consisting of rows of erect, interstrial scales, each scale almost as long as distance between rows, more widely spaced within each row except on 1, each scale broad, averaging about twice as long as wide. Apex acuminate, the mucro short; not at all divaricate. Type Locality. — Fourteen km. east of Hua- tusco, Veracruz, Mexico, Type Material. — The male holotype was col- lected at the type locality on July 7, 1967, near 1,300 m., No. 173, by S. L. Wood, from Quenus twigs. The holotype is in my ctjllection. Micracisella tutidiila, n. sp. This species is allied to monadis Wood, described above, but it may be distinguished by the reticulate-granulate frons, by the slender declivital interstrial scales, by the much smaller declivital interstrial granules, and by the 37 presence of four marginal teeth on the pionotum. ffma/f.— Length 1.9 mm.. 3.1 times as long as wide; color black. Frons convex, less strongly on lower half, epistomal margin weakly elevated; surface very finely reticulate- granulate, reduced to minute points on small median area near epistoma; vestiture consisting of rather sparse, con- spicuous, subplumose setae of moderate length to vertex. Scape twice as long as pedicel, bearing a small tuft of hair. Pronotum 1.1 times as long as wide; widest just be- hind middle, sides rather strongly arcuate, weakly con- stricted on anterior third, rather narrowly rouncled in front; anterior margin armed by six small teeth of equal size; summit near middle, rather broad; posterior area minutely reticulate-granulate, with a few minute, almost flat, granules toward summit; vestiture rather stout, moderately abundant, rather long. Elytra 2.0 times as long as wide. 2.0 times as long as pronotum; sides almost straight and parallel on basal three-fourths, narrowly acuminate behind; striae not impressed, the punctures small, shallow; interstriae twice as wide as striae, somewhat irregualar, the punctures very fine, uniserate. Declivity convex, rather steep; strial punctures slightly deeper than on disc; interstrial punc- tures replaced by fine granules; apex subacuminate. mucro not developed, not at all divaricate. \'estiiure consisting of very fine, moderately long strial hair and rows of erect interstrial scales, slightly longer on declivity; each scale on declivity slightlv longer than distance be- tween rows, similarly spaced within e.ich row, each about six times as long as wide. Type Locality. — Ten km. northeast of Tezuit- lan, Puebla, Mexico. Type Material. — The female holotype was collected at the type locality on July 2, 1967, 1,600 m.. No. 142, by S. L. Wood, from an axial pith tunnel in Rubus sp. A second female speci- men, almost certainly this species but not desig- nated as a paratype, was taken 26 km. east of Morelia, Michoacan, Mexico, June 14, 1965, 2,500 m., S, L. Wood, from an Arbutus sp. twig. The holotype is in my collection. Micracisella nigrella, n. sp. This species is allied to nigra Wood, but it is distinguished by the coarser sculpture of strial punctures and declivital granules, by the larger, coarser, more abundant interstrial scales, and by the different arrangement of marginal teeth on the pronotum. Female. — Length 1.7 mm. (paratypes 1.3-1.8 mm.), 3.0 times as long as wide; color almost black, summit of pronotum reddish brown. Frons convex on upper half, transversely impressed on lower half, an indistinct median impression toward center; vestiture sparse, of very short, fine, subplumose setae. Scape twice as long as pedicel, bearing a moderate- ly large tuft of long hair. Pronotum 1.14 times as long as wide; widest just behind midtUe, the sides weakly arcuate on more than basal half, rather narrowly rounded in front anterior margin armed by two rather large teeth; summit at middle, rather broad; posterior area minutely reticulate- Bricham Young University Science Bulletin j granulate, with moderately abundant, isolated, very fine granules; vestiture of rather short, broad scales. Elytra 1.9 times as long as wide. 1.7 times as long as pronotuni; outline as in nitidula; striae not impressed, the punctures fine, distinct, shallow, interstriae with fine lines. twice as wide as striae, the punctures very fine, half as large as those of striae. Declivity convex, rather steep; interstrial punctines about as large as those of striae; apex acuminate, the sutural apex distinctly, shallowly emarginate. Vestiture toiisisiiiig of rows of erect inlevsirial scales; each scale very slightly shorter than distance be- tween rows, similarly spaced within rows, each about two and one-half to three times as long as wide; a supple- mental sutural row of smaller scales on declivity. Male. — Similar to female except frons slightly more convex; scape without tuft of hair; elytral scales very slightly larger. Type Locality. — Volcan de Agua, Esquintla, Guatemala. Type Material. — The female holotype, male allotype, antS 81 paratypes were collected at the type locality on May 19, 1964, 1,000 m.. No. 603, by S. L. Wood, from branches of an unidentified shrub somewhat resembling Rhus. The holotype, allotype, and paratypes are in my collection. Micracisella similis, n. sp. This species is allied to nigra Wood, but it is distinguished by the more slender body form, i)y the less strongly convex frons, with a slight median impression above epistoma, and much greater development of the vestiture, and by the more strongly reticulate pronotal disc. Female. — Length 1.5 mm. (paratypes 1.4 I. .5 mm.). 3.0 times as long as wide; color black, summit of pronolum reddish brown. Frons planoconvex, flattened toward center, giadually ascending to epistomal margin on lower fourth; surface minutely reticulate-gianulate. smoother medially toward epistoma; vestitme moderately long and abundant toward lateral areas, subplumose. Scape twice as long as pedicel, bearing a small tuft of long hair. Pronotum 1.16 times as long as wide; widest at middle, sides on basal half straight, almost parallel, rather narrowly rounded in front; anterior margin armed by two moderately large, basally contiguous teeth; summit at middle, broad; posterior areas minutely reticulate- granulate, with a few. fine scattered gianules; vestiluie short, scalelike. Elytra 1.9 times as long as wide. 1.7 times as long as pronotum; outline as in nitidula: striae not impressed, the punctures small, shallow; interstriae marked by lines, shining, slightly less than twice as wide as striae, the uniseriate punctuies very fine. Declivity convex, rather steep; strial punctures somewhat reduced; interstrial punc- tures replaced by fine gianules on upper half, almost obsolete below; general surface gianulose toward sutural apex; apex acuminate, the sutural apex shallowly emargi- nate. Vestiture consisting of minute, fine, strial hair and uniseriate rows ot longer, erect, interstrial scales; each scale slightly shorter than distance between rows, similarly spaced within each row. each scale on declivity about three times as long as wide. Male. — Similar to female except frons more nearly convex, the frontal vestiture somewhat shorter; reduction of interstrial tubercles on lower half of declivity less pronounced. Type Locality. — Volcan Ceboruco, Nayarit, Mexico. Type Material. — The female holotype, male allotype, and seven paratypes were collected at the type locality on July 5, 1965, 1,200 m., No. 189, by S. L, Wood, from axial pith tunnels in Serjania sp. The holotype, allotype, and paratypes are in my collection. Micracisella vescitla, n. sp. This species is allied to similis Wood, described above, but it may be distinguished by the coarser strial punctures, by the absence of interstrial granules on the declivity, and by the large, broader, interstrial scales. Female. — Length 1.4 mm. (allotype l..'i5 mm.). 3.0 times as long as wide; color ilark brown, summit of pronotum reddish brown. Frons as in similis but less strongly flattened. Scape twice as long as pedicel, bearing a tuft of long hair. Pronotum 1.17 times as long as wide; outline as in similis; anterior margin armed by two moderately large, subcontiguous teeth; posterior area minutely reticulate- granulate, with obscure, subgranulate punctures; vestiture of rather abundant, short scales, each twice as long as wide. Elytra 1.8 times as long as wide. 1.6 times as long as pronotum; outline as in nitidula: striae not impressed, the punctures rather fine, not always clearly defined; inter- striae irregular, almost subrugulose. as wide as striae, the uniseriate punctures fine. Declivity convex, rather steep; strial punctuies deeper than on disc, the interstrial punctures equal in size to those of striae; apex acuminate, shallowly emarginate at sutural apex. Vestiture consist- ing of very fine, short, strial hair and interstrial rows of erect, uniseriate scales, each scale about as long as distance between rows, similarly spaced within each row. each about three times as long as wide; a supplemental row of smaller, sutural scales on declivity. Male. — Similar to female except frons more strongly convex, the frontal vestiture shorter; tuft of hair on scape much smaller; teeth on anterior margin of pronotum slightly larger. Type Locality. — Two km. north of Cerro Gordo. Veracruz, Mexico, Type Material. — The female holotype and male allotype were collected at the type locality on July 6, 1967, 1,000 m.. No. 166. by S. L. Wood, in axial pith timnels in Serjania sp. The holotype and allotype are in my collec- tion. Micracisella squamatida, n. sp. This species is allied to nigra Wood and the three preceeding species, but it is easily distin- guished from these and all other representatives of the genus by the much more widely spaced teeth on the anterior margin of the pronotum, by the scalelike, small, sirial setae in addition to 1 I New Neotropical Scoi.ytidae those of the interstriae, and by the more strongly divaricate elytra at the sutural apex. Female. — Length 1.4 mm. (paratypes 1.3-1.5 mm.). 3.0 times as long as wide; color black, sunnnil of prono- tum reddish brown, vestiture white. Frons convex above, becoming flattened below, then ascending slightly to epislomal margin: surface reticulate- granulate above, becoming almost smooth in central area below except for a few fine points; vestiture rather abundant and long in large dorsolateral areas. Scape about twice as long and slightly wider than pedicel, bear- ing a small tuft of long hair. Pronotum 1.1 times as long as wide; widest at middle, the sides on posterior half almost straight and parallel on basal half, feebly constricted on anterior third, rather narrowly rounded in front; anterior margin armed by two moderately large, rather widely separated teeth; sum- mit at middle, rather broad; posterior area finely reticu- late-granulate, with a few minute granules; vestitiue consisting of short, broad scales. Elytra 1.8 times as long as wide, 1.7 times as long as pronotum; outline as in nitidula; striae not impressed, the punctures small, somewhat indistinct; interstriae inegidarly marked by fine points and lines, one and one- half times as wide as striae, the punctures fine, uniseriate. Declivity convex, rather steep; strial punctures evidently sli!;hil\ deeper; inlerslrial punctures replaced by small rounded granules; sutural apex emarginate. Vestiture consisting of rows of small strial scales, each slightly longer than diameter of a strial puncture, about four times as long as wide, and rows of longer interstrial scales; each interstrial scale one and one-half times as long as strial scales and about equal in length to distance between rows of interstrial scales, each about three times as long as wide. Male. — Similar to female except frons more strongly convex on upper two-thirds, the smooth area much smaller and subreticulate; tuft of hair on scape reduced. Type Locality. — Ten km. south of Huajua- pan, Oaxaca, Mexico. Type Material. — The female holoiype, male allotype, and nine paratypes were collected at the type locality on June 16, 1967, about 2,()()() in., No. 45, by S. L. Wood, from axial pith tunnels in Serjania sp. Five additional paratypes were taken 32 km. south of Huajuapan, Oaxaca, Mexico, on the same date. No. 49, from twigs of an unidentified shrub. The holotype, allotype, and paratypes are in my collection. Micracisella divaricata , n. sp. While several of the tropical species in this genus have the eyltral apices very slightly divari- cate, this character is pronounced in this species and serves to distinguish it. Female. — Length 1.6 mm. (paratypes 1.3 mm.). 2.8 times as long as wide; color black, central area of pro- notum reddish brown. Frons planoconvex from epistoma to upper level of eyes; surface coarsely reticulate above and laterally be- coming smooth toward center, the pimctures fine, moderatelv abinidant; vestiture consisting of moderately long, stout, subplumose setae inoderately abmidant in lateral areas and above. Scape twice as long as pediiel. bearing a small tuft of long hair. 39 Pronotum 1.1 times as long as wide; widest on basal half, the sides alnrost straight and parallel, then aicuately rounded to constriction on anterior third, nanowly rounded in front; anterior margin armed by two. rather large, basally contiguous teeth; stuimiit rather indefinite, at middle; small asperities continue to just behind sum- mit, isolated granules continue to base; surface of posterior area coarsely reticulate; vestiture of small, short scales. Elytra 1.7 times as long as wide. 1.6 times as long as pronotunv, sides almost straight and parallel on more than ba.sal two-thiids. then narrowly rounded to blunt apex; striae not impressed, the punctures very small; interstriae shining, marked by lines, twice as wide as striae, the uniseriate punctures fine, somewhat obscure. Declivity convex, rather steep; strial punctures largely obsolete; interstrial punctures replaced by rounded granules; apex divaricate, the emargination as wide as width of antennal club and half as deep. Vestiture consisting of very minute, very fine strial hair and rows of longer, suberect, inter- strial scales, each scale slightly shorter than distance be- tween rows, similarly spaced within rows except closer on I, each scale about four times as long as wide. Male. — Similar to female except frons more strongly convex, the setae more evenly distributed; elytral scales slightly wider, about three times as long as wide. Type Locality. — Two km. north of Cerro Gordo, Veracruz Mexico. Type Material. — The female holotype, male allotype, and one female paratype were collected at the type locality on July 6, 1967. 1,000 m.. No. 166, by S. L. Wood, froirr axial pith tunnels in Serjania sp. The holotype, allotype, and paratype are in my collection. Micracis costaricensis, n. sp. The only species known to me that is at all similar to this one is cariuulatus Wood. This species may readily be distinguished from cariuu- latus by the dull surfaces, by the more irregularly punctured elytra, by the more delicate, more strongly flattened interstrial scales, and by the slightly impressed second declivital interspace. Female. — Length 2.7 mm. (paratypes 2.2-2.8 mm.). 3.2 times as long as wide; color rather dark reddish brown. Frons strongly, transversely impiessed and slightly concave from epistoma to well above eyes, upper half of impressed area with a fine, low median carina; surface coarsely reticulate except smooth just above epistoma; vestiture consisting of sparse, short setae in reticulate area, with a fringe of long subplinnose hair along margin of impressed aiea above level of eyes. Scape flattened, tri- angular, ornamented by a fringe of long hair; sutures of club strongly procurved, 1 reaching middle. 2 extending three-fourths of club length from its base. Pronotum 1.3 times as long as wide; widest on basal half, sides almost straight and parallel on slightly more than basal half, rather broadly rounded in front; anterior margin armed by 12 rather coarse, isolated teeth; siniimit in front of middle; rather coarsely reticulate behind sum- mit with rather sparse, fine, subgrairulate punctures, Vestitme consisting of rather short, stout, delicate, white setae. Elvira l.tl times as long as wide. 1.6 times as long as pionotum; sides almost straight and parallel on basal three-fourths, rather broadly rounded behind to the 40 Brioham Young University Science Bulletin short, projecting sutural apex: striae not impressed, the punctmes small, shallow, distinct; interstriae about three times as wide as striae, irregular, punctures and other features obscured by irregular surface features. Declivity evenly convex, very steep; striae slightly impressed, the punctures deeper than on disc; interstriae weakly convex, with lower half of 2 somewhat impressed; all interspaces bearing median rows of fine, rounded granules, those in lateral areas a little larger; terminal mucro as in other spceies of this genus. Vestiture consisting of rows of fine strial hair and delicate, very slender interslrial scales of equal length; each scale almost equal in length to distance between rows of scales; easily abraded. Male. — Similar to female except slightly smaller, 2.4 mm., stouter, 2.8 times as long as wide; frons convex with a slight transverse impression just above epistoma. smface finely gianulate, the vestiture greatly reduced; scape not dilated, the tuft of hair almost obsolete; pronotuni and elytra very slightly more coarsely sculptured. Type Locality. — Volcan Poas, Heredia Prov., Costa Rcia. Host. — Calliandra confusa (paratypes). Type Material. — The holotype, allotype, and 41 paratypes were collected at the type locality on June 19, 1966, at an elevation of about 2,000 m., by S. L. Wood, from branches of a leguniin- ose shrub. Additional paratypes were taken in Costa Rica as follows: 18 from Tapanti, Cartago Prov., 1,400 m., Aug. 17, 1963; 25 from Santa Ana, San Jose Prov., 1,400 m., Oct. 4, 1963; and 3 from San Ignacio de Acosta, San Jose Prov., 1,500 m., July 5, 1963, all taken by S. L. Wood, from Calliandra confusa. Eleven specimens not included in the type series are from Zamorano, Morazan, Honduras, April 18, 1964, 700 in., from Salix guatemalensis. The holotype, allotype, and paratypes are in my collection, Micracis carinulus, n. sp. Similar to costaricensus Wood, described above, but smaller, the declivital striae not im- pressed, the impression of the feinale frons not reaching upper level of eyes with the median carina rather sharply elevated. Female. — Length 2.1 mm. (paratypes 1.6-2.1 mm.). 3.1 times as long as wide; color yellowish brown. Frons broadly concave from epistoma to upper level of eyes, upper half divided by a fine, low carina, sinface smooth and shining just above epistoma. then finely punctured and becoming coarsely reticidate on upper half; vestiture consisting of short, stout, close setae in concave area, and long, curved, subplumose hairs along upper margin of concavity. Antenna as in custaricensis. except wider. Pronotuni as in costaricemis except anterior margin unarmed. Elytra as in costarincensis except surface of disc a little more irregular and strial punctures less distinct. Declivity evenly convex except for terminal mucro; strial punctures very slightly impressed; interstriae with small. rather obscure gianules. Vestiture consisting of moderaltly long, rather abundant coarse strial and interslrial setae; those on median row of each interspace slightly longer. Male. — Similar to female except frons convex and finely granulate with a slight transverse impression above epistoma; scape less strongly dilated; anterior margin of pronotiMii armed by about eight small serrations; and median rows of elytral setae more nearly .scalelike. Type Locality. — Playon, Puntarenas Prov., Costa Rica. Host. — Rheedia editlis. Type Material. — The female holotype, male allotype, and 20 paratypes were collected at the type locality on Februai7 22, 1964, at an eleva- tion of about 50 m., by S. L. Wood, fiom branches of the above host. Two additional para- types were taken at Fort Clayton, Canal Zone, Panama, on December 22, 1963 at about 30 m., by S. L. Wood, from a fallen litnb of an uniden- tified tree. The holotype, allotype, and paratypes are in my collection. Micracis tribulatus, n. sp. This species is very colsely related to costari- censis Wood, described above but it may be dis- tinguished by the darker color, by the more slender form, by the armed anterior margin of the female pronotum, by the granulate, discal, interstrial punctures, and by the coarser de- clivital granules. Female. — Length 2.8 mm. (paratypes 2.2-2.8 mm.), 3.6 times as long as wide; color very dark brown. Frons and antenna as in costaricemis. Pronotum 1.2 times as long as wide; sides straight and parallel on basal two-thirds, rather broadly rounded in front; anterior margin armed by six small teeth; summit distinct, in front of middle; posterior areas reticidate- gianulate, with sparse, moderately large, isolated granules to base; vestitme of fine hair and slender scales. Elytra 2.3 times as long as wide, 2.0 times as long as pronotuni; sides straight and parallel on basal three- fourths, narrowly mucionate behind; strial punctures very small, distinct; interstriae at least three times as wide as striae, subshining, marked by lines, the pimclures fine, subgranulate; declivity steep. co\ex. confinetl to posterior fourth; interslrial punctures replaced by moderately large, rounded granides; apex mucionate. \'estiture consisting of fine strial hair and rows of longer, erect, interstrial scales, slightly longer on declivity; each scale on disc about six to eight limes as long as wide, much more slender on declivity. Male. — Similar to female except 2.9 times as long as wide; frons convex above, transversely impressed below, with a small median impression; scape smaller, narrower; teeth on anterior margin of pronotuni larger; declivital scales slightly longer. Type Locality. — Twenty-six km. west of Tex- melucan, Puebla, Mexico. Type Material. — The feinale holotype, male allotype, and 9 paratypes were collected at the type locality on June 6, 1967, 2,900 in.. No. 29, by S. L. Wood, from Salix. The holotype, allotype, and paratypes are in my collection. ( New Neotropical Scolytidae Micracis lepidus, n. sp. This species is not closely related to any species known to me, although there inay be a remote relationship to lignalor Blackman. The size, scalelike elytral setae, armed anterior margin of the female pronotum, and the unexcavated female frons distinguish it. Female. — Length 3.0 mm. (paratypes 2.3-3.0 mm.). 3.2 times as long as wide; color dark brown to black with white scales. Frons convex above, somewhat flattened on lower third: surface leticiilate and finely piinctiiied above, smooth, shining on lower part of flattened area; vestitiire subplumose. erect, rather abundant to vertex except on shining epistomal area, short below, moderately long above. Scape broadly triangidar, ornamented by a inar- ginal fringe of long hair; club 1.6 times a.s long as wide, suture 1 reaching middle. Pronotum 1.3 times as long as wide; sides almost, straight and parallel on basal half, rather broadly rounded in front; anterior margin armed by 12 low teeth; summit in front of middle, distinct; posterior area reticulate- granulate, very minute granules indicated towaid summit in median area; vestiture short, of minute hair and slender scales. Elytra 2.0 times as long as wide, 1.6 times as long as pronotum; sides straight and paralle on basal three- fourths, rather nanowly mucronate behind; striae not impressed, the punctures minute; inteistriae about four times as wide as striae, undulating slightly and with numerous indefinite points, the punctures fine. Declivity steep, convex, confined to posterior fourth; strial and interstrial punctures of equal size, close, confused, rows of minute granules on all inteistriae except 1; apex obtusely mucionate. Vestiture consisting of minute strial hair and erect scales; scales on disc in interstrial rows, more abundant and confused on declivity, each scale three to four times as long as wide; base of declivity also with several pointed bristles longer than scales, sides with some slender, long hair. Male. — Similar to female except averaging lightly smaller; 2.6 times as long as wide; frons more strongly convex, not flattened belou, surface with tnany rounded granules; scape less stongly expanded, the tuft of hair smaller; teeth on anterior maigin of pronotum laiger; elytral scales shorter, broader, mostiv twice as long as wide, the bristles and inteistrial hair also scalelike and slightly larger than other scales. Type Locality. — Cerro Punta. Chiriqui, Pana- Type Material. — The female holotype. male allotype, and 50 paratypes were collected at the type locality on December 19, 1963 and January 11, 1964, 1.800 m., No. 315, 371, 387, 404. by S. L. Wood, from branches of Inga sp. The holotype, allotype, and paratypes are in my collection. Micracis lignicolus, n.sp. This species is closely related to tiguator Blackman, but it is distinguished by the less strongly elevated interstriae 9 at base of male declivity, and by the less strongly tuberculate interstriae at margin of declivity, by the slightly 41 elevated, subserrate, inale declivital interstriae 3 on basal third, by the different arrangement of setae on the female frons and scape, and by the narrower interstrial scales in both sexes. Male. — Length 2.8 mm. (paratypes 2o-3.3 mm.). 2.8 times as long as wide; color reddish brown. Frons convex above, transversely impressed on lower half epistomal margin slightly elevated; surface reticulate- granulate, with sparse, smooth granules on convex area; vestiture sparse, limited to lateral areas. Scape about twice as long as pedicel, distal width equal to length, bearing a small tuft of long hair; club 1.8 times as long as wide, suture 1 extending three-fifths of club length from base. Pionotum I.I times as long as wide; sides almost straight and parallel on basal half weakly constricted just in front of middle, anterior inargin rather broadly rounded and armed by 12 teeth; summit at middle, broad; posterior areas reticulate-granulate, the punctures fine, inconspicuous, some minutely granulate; vestiture consisting of minute hair and slender scales. Elytra 1.7 times as long as wide. 1.6 times as long as peonotum; sides straight and parallel on basal three- fourths, narrowly rounded behind; striae moderately impressed on posterior thiid of disc, the punctured rather coarse and deep; interstriae as wide as striae, shining, somewhat irregular, the punctures uniseriate. rather fine. Declivity rather abrupt, convex, steep; striae not impressed, the punctures smaller and sqiiamifeious; interstriae 1-8 each with one or two rounded nodules at base of declivity as in tiguator, 9 weakly elevated but ending just behind base of declivity, 3 bearing a row of about six pointed tubercles on upper two-thirds. ,5 bearing two similar tubercles near base; apex extended slightly but terminated by ascending costal maigins that form a pseiido-emargina- tion; interstrial punctures coarse, two-thirds as large as those of striae, squamiferous. Vestiture consisting of minute strial hair and interstrial rows of slender, erect scales on disc; on declivity strial setae and some inter- strial setae form moderately short scales each about four times as long as wide, other eiect, interstrial scales in lows, much longer at declivital base, some more than twice as long as ground scales. Female. — Similar to male except 3.2 times as long as wide; frons convex to epistoma. median half glabrous, devoid of granules; scape triangularly extended, bearing a much laiger tuft of hair, but shorter than in lignalor; striae not impressed toward declivity; declivity devoid of tubercles or nodules, 3 weakly convex; vestiture less dense on declivity, the scales more slender, not longer than on disc. Type Locality. — Cerro Peiia Blanca, Hon- duras. Type Material. — The male holotype, female allotype, and 13 paratypes were taken at the type locality on April 23, 1964, 2,000 m., No. 529, in Miconia schlechtendalii, by S. L. Wood; 23 para- types bear identical data except lot 533 taken from branches of Qiiercus williamsi; four para- types were from Zamorano, Morazan Honduras, April 18, 1964, 700 m.. No. 539, Quercus sapotae- folia, S. L. Wood; 15 paratypes were from Volcan de Agua. 1,000 in., No. 598, froin an unidentified broken branch, by S. L. Wood. The holotype, allotype, and paratypes are in my collection. 42 Bricham Young University Science Bulletin Micracis detentiis, n. sp. This species is vei7 colsely related to cari- nulus Wood, described above, but it may be distinguished by the larger size, by the larger granules behind the pronotal summit, by the more deeply impressed female froiis with a more prominent median elevation below, and by the coarser interstrial bristles on the declivity. Female. — Length 2.2 mm. (paratypes 1.8-2.3 mm.). 3.2 times as long as wide; color yellowish brown. Frons transvasely, siibconca\ely impressed from epistoma to upper level of eyes, median third of epistonia moderately elevated, the smooth, impunctate, shining elevation extending about one-third of distance to upper level of eyes; remaining surface reticulate-granulate, the punctures minute, obscure; vestiture of minute sub- plumose hair, becoming longer toward upper margin of impression. Scape nanowlv triangulai, ornamented by a tuft of long hair; club 1.7 times as long as wide, suture 1 extending slightly beyond middle. Pronotum 1.3 times as long as wide; sides straighl and parallel on slightly more than basal half, rather broadly rounded in front; anterior margin unarmed; summit in front of middle; posterior area reticulate-granulate, with rather sparse, fine, isolated gianules; vestiture hairlikc. inconspicuous. Elytra 2.0 times as long as wide. 1.6 times as long as pronotum; sides straight and parallel on basal three- fourths, acuminate behind; striae not impressed, the punctures small, distinct; interstriae at least twice as wide as striae, with lines and points, the punctures uniseriate, very fine. Declivity steep, convex, confined to posterior fourth; minutely gianulose, strial punctures deeper bul somewhat obscure; interstrial punctures replaced by fine granules; apex rather strongly acuminate. Vestiture con- sisting of very fine strial hair and rows of erect, inter- strial bristles, slightly longer and coarser on declivity; each bristle on declivity slightly longer than distance be- tween rows. Male. — Similar to female except 2.7 times as long as wide; frons convex aboxe, a moderate transverse impres- sion on lower half, the median elevation obsolete; scape not as wide as long, bearing a small (lift of shorter hair; anterior margin of pronotum armed by eighl teeth; declivital vestiture slightly coarser. Type Locality. — Twenty-four km. south of Mazamitla, Jalisco, Mexico. Type Material. — The male holotype, female allotype, and five paratypes were collected at the type locality on June 22, 1965, 2,500 m., No. 94, by S. L. Wood, from branches of a leguminose tree; 11 paratypes were taken 8 km. south of Atenquequc, Jalisco, Mexico, on June 25, 1965, 1,000 m.. No. 118, by S. L. Wood, from an un- identified shrub; two paratypes were from Maguarichic, Chihuahua, Mexico. July 13. I960, blacklight, S. L. Wood. The holotype, allotype, and paratypes are in my collection. Micracis unicornis, n. sp. This species evidently is closely related to detenlus Wood, described above, but the male bears a large, pointed median tubercle on the epistoma, the vestiture is finer, and the elytral declivital tubercles and pronotal asperities are larger. Male. — Length 2.1 mm. (male paratype 2.1 mm.), 2.7 times as long as wide; color reddish brown. Frons convex above, tran.sversely impressed on lower half; epistomal margin slightly raised, with a rather indefinite median elevation that bears a conspicuous, moderately large, pointed tubercle; surface reticulate, the punctures obscure, a few fine granules in convex area; vestiture of coarse, subplumose setae of moderate length, rather sparse. Scape twice as long and twice as witle as pedicel, bearing a small tuft of hair; club 1.55 times as long as wide, suture I extending slightly beyond middle of club. Pronotum 1.05 times as long as wide; sides feebly arcuate, subpaiallel on basal half, rather broadly rounded in front; anterior margin armed by 12 coarse teeth; sum- mit at middle; posterior area reticulate-granulate, with a few- minute granules behind summit, the minute punc- tures obsc.-re; vestiture inconspicuous, composed of stout hah li' e setae. Elytra 2.0 times as long as wide, 2.0 times as long as pronotum; sides straight and parallel on basal three- fourths, narrowly mucronate behind; striae not impressed, the punctures small, distinct; interstriae about four times as wide as striae, marked by many lines and a few points, the punctures fine, shallow. Declivity steep, convex, con- fined to posterior fourth; strial punctures larger and im- pressed; interstriae minutely granulose. the punctures replaced by fine granules. Vestiture consisting of minute strial hair and rows of erect, pointed, interstrial bristles, longer and stouter on declivity; each bristle on declivity slightly longer than distance between rows. Type Locality. — Six km. west of Quiroga, Michoacan, Mexico. Type Material. — The male holotype and one male paratype weie collected at the type locality on June 17, 1965, 2,200 m.. No. 72, by S. L. Wood, from a shrubby herbaceous plant. The holotype and paratype are in my collec- tion. Micracis inimicus, n. sp. The scalelike interstrial setae on the elytra and the armed anterior inargin of the pronotum suggest a relationship to evanescens Wood, but this species is smaller and the frontal sculpture of the female is entirely diffeient. Female. — Length 2.3 mm. (paratypes 2.0-2.5 mm.). 3.5 times as long as wide; color medium to dark t)rown. Frons broadly concave to uppa le\el of eyes, lower third with unusual sculpture; epistomal margin slightly elevated, smooth on a narrow band on median fourth, then a slight, narrow ridge, then above this a narrow slit-like impression, rather strongly abruptly, transvei.sely elevated above Impression on more than median half, its upper slope marked by two or three minute, transverse ridges in its gradual descent to concavity; vestiture short, sparse, inconspicuous; surface reticulate-granulate. Scape nanowlv triangular, ornamented by a tuft of long hair; club 1.9 times as long as wide, widest on distal half, suture I extending two-thirds of club length from base Pronotum 1.3 times as long as wide; as in detentus except some setae stout, almost scalelike. ■A New Neotropical Scolytidae Elytra 2.1 times as long as wide, 1.75 times as long as pronotiim; outline and disc as in clelentus except strial punctures rather obscure. Declivity steep, convex, confined to posterior fourth; striae weakly impressed, the punctures larger, deeper; interstrial punctures suh- gianulate. except reduced on 2: granules larger in some paratypes. Vestiture consisting of very fine strial hair and rows of erect interstrial scales, slightly longer on declivity; each scale on declivity as long as distance be- tween scales, spaced mote closely within each row. trun cate at their apices, each about five times as long as wide. Male. — Similar to female except 2.7 times as long as wide: frons convex, transversely impressed on lower half, episloma elevated on median third, without elaborate sculpture of female; anterior margin of pronotum armed by six teeth; declivital tubercles more distinct except on interstriae2 and 3. Type Locality. — Volcan de Agua, Esqiiiiula, Guatemala. Type Material. — The female holotype, male allotype, and 21 paratypes were collected at the type locality on May 19, 1964, 1,000 m., No. 603, by S. L. Wood, from branches of an unidentified shrub. llie holotype, allotype, and paratypes are in my collection. Micracisfestivus, n. sp. Apparently the only known species allied to this one is inimicus Wood, described above, but it is easily distinguished by the slender elytral setae, and by the very different frontal sculpture of the female. Female. — Length 2.2 mm. (paratypes 1.7-2.:i mm.). ,"5.4 times as long as wide: color dark brown. Frons concavely impressed from episloma to above upper level of eyes; epistomal margin elevated, median half with a narrow, slit like impression just above margin and a low elevated, transverse, subcarinate ridge immediately above the slit; surface reticulate-giamilate. the punctures fine, obscure; vestiture sparse, fine, short below, loirger above. Scape narrowly triangular, orna- mented by a tuft of long hair; club 1.6 times as long as wide, suture 1 extending beyond middle. Pronotiuii 1.2 times as long as wide; as in detentus except gianules in posterior areas larger and anterior margin anned by six small teeth. Elytra 2.1 times as long as wide, 1.8 times as long as pronotimi: outline and disc as in detentus except strial punctures very slightly deeper. Declivity steep, convex, confined to posterior fourth: striae feebly impressed, the pimctures deeper and slightly larger than on disc; inter- striae each armed by a row of rounded granules similar to but slightly larger than in inimicus: apex stiongly acuminate or mucronate. Vestiture consisting of fine strial hair and rows of longer interstrial bristles, slightly longer on declivity; each bristle up to one and one-half times as long as distance between rows of bristles, more closely spaced within each row. Mate. — Similar to female except 2.7 times as long as wide; frons as in male of inimicus: interstrial punctures on disc as large as those of striae, very feebly granulate: strial punctures smaller on disc and declivity, somewhat obscure on declivity: vestiture with interstrial setae on disc rather narrowly scalelike, truncate, those on declivity longer, more slender and almost pointed at their apices. 43 Type Locality. — Zamorano, Morazan, Hon- duras. Type Material. — The lemale holotype, inale allotype, and 16 paratypes were collected at the type locality on April 18, 1964, 700 m.. No. 506, by .S. L. Wood, from branches of Celt is iguanaea; 14 paratypes bear identical data except they are lot No. 534, from Calliandra houstoniajia. The holotype, allotype, and paratypes are in my collection. Hylocuriis alternus, n. sp. The only species known to me that is closely related to this one is inaequalis Wood; however, it has the male declivital spines much less highly developed. The development of declivital spines is similar in magnitude to medius Wood, but the relatioirship to the latter species is remote. Females of this species lack the abuiulaiu frontal vestiture of inaequalis. Mate. — Length 2.5 mm. (paratypes 1.7-2.5 nmi.). 2.5 times as long as wide; color very dark blown, almost black. Frons concealed in type. In male paralype, convex and gianulate above, transversely impressed and punc- tured below a transverse carina formed at upper level of eyes by four basallv confluent tubercles; vestiture in conspicuous. Pronotum 1.1 limes as long as wide; widest near base, the sides feebly arcuate and converging very slightly on basal two-thirds then rather broadiv lounded in front; anterior margin armed by about 12 coarse serra- tions: sunmiit in front of middle; posterior area sub- shining, rather finely piuictured, the posterior margin of each puncture subasperate, more coarsely sculptureil toward sunmiit. Vestiture consisting of short, incon- spicuous hair. Elytra 1.4 times as long as wide. 1.2 times as long as pronotiun: sides almost straight and parallel to declivital base: piofile of posterior margin interrupted by projection of spines on interspaces 9 and '^ and by sutmal apex; striae not impiessed. the punctures large, deeply impressed; inteistriae narrower than striae, irregularly, sparsely punctured, each ending posteriorlv at declivital margin in a tubercle or spine. Declivity abrupt, very steep; upper and lateral maigins armed by a tow of tubercles, those on interspaces 2. 4 and 6 distinctly shorter, all tubercles abruptly declivous behind. 9 longer and projecting; de- clivital face convex, coarsely puncturet^l with two rather coarse tubeicles on intaspace 3 and about four smaller ones on middle third of 1; apex narrowly acuminate. Vestiture consisting of short, stout, bristles. Female. — Similar to male except frontal carina absent, the frons more finely scidpturetl; anterior margin of pronotiun finely .serrate; elytra not abruptly declivous, without a marginal row of tubercles on declivity; each interspace bearing about three to five small tubercles on upper or lateral half of declivity. Type Locality. — twenty-one km. south- east of Liberia, Guanacaste Prov., Costa Rica. Type Material. — The male holotype, female allotype, and 39 paratypes were collected at the type locality on July 10, 1966, at an elevation of 44 Brigham Young Univf.rsitv Science Bulletin about 50 m., from a branch of an unidentified tree, by S. L. Wood. The holotype, allotype, and paratypes are in my collection. Hylocurus aberrans, n. sp. This species is not allied to any other repre- sentative of the genus known to me. The sutures of the antennal club are more strongly procui-ved than in other Hylocurus, but less so than in Micracis. The posterior face of the anterior tibia is tuberculate as in Hylocurus; the posterior extremity of declivital interspace 9 almost reaches the costal margin thereijy resembling somewhat certain species oi Micracis. Male. — Length 2.2 mm.. 2.6 times as long as wide; color dark reddish brown. Frons broadly convex with a slight transverse impres- sion just above epistomal margin; surface finely, regularly granulate. Antennal club with two prociirved sutiues, 1 extending two-fifths from base. 2 extending four-fifths from base. Pronotum 1.2 times as long as wide; widest on basal half, the sides feebly arcuate, rather broadly rounded in front; anterior margin armed by 16 or more serrations; summit in front of middle; posterior area reticulate, with small, subgianulate punctures of moderate abundance. Vestiture consisting of short, erect scales with a few bristles in asperate area. Elytra 1.5 times as long as wide, 1.6 times as long as pronotum; sides almost straight and parallel to base of declivity then slightly wider at middle of declivity; posterior outline interrupted by declivital spines on interspaces 7. 8, and 9 and by short, broad, sutural apex; striae slightly impressed, the punctures deep, coarse, much smaller toward base; interstriae about half as wide as striae, the punctures on 1 moderatlly large, others mostly minute. Declivity abrupt, very steep, almost entire- ly margined by a row of tubercles or spines; tubercles at base of declivity on interspaces 1 and 2 not pointed, abruptly declivous behind, those on 2 to 8 produced into increasingly large and pointed toothlike spines, 9 similar but lower and extending nearer elytral apex than normal for this genus; apex not strongly produced, its margin subserrate. the small dentitions about half as high as apex of elevation on interspace 9; central area coarsely punc- tured, interspace 3 slightly elevated near center and bearing three small teeth, 1 on lower third gradually elevated to apex and finely .serrate. Vestiture consisting of a few stout bristles or scales at margin and on inter- spaces 1 and 3. Fe?nale. — Similar to male except frons less strongly convex, finely punctured, ornamented by stout, short, rather sparse, setae; scape flattened and bearing a tuft of long, coarse setae; anterior margin of pronotum almost unarmed; elytra more finely sculptured, the declivity convex, without marginal ring of tubercles, the striae regidarly pmictured. odd numbered interspaces bearing several rounded granules; rows of broad scales on upper half of all tleclivital interspaces. Tyl^e Locality. — Dominical, Piuitarenas Prov., Costa Rica. Type Material. — The male holotype and female allotype were taken from the same timnel at the type locality on December 9, 1963, at sea level, by S. L. Wood, from a small branch of a recently cut unidentified tree. The holotype and allotype are in my collec- tion. Hylocurus incomptus, n. sp. Fig. 3 Fig. 3. Hylocurus incomptus dorsolateral aspect of male elytral declivity. This species is allied to femineus Wood, but it is readily distinguished by absence of sexual dimorphism on the frons, by the much more finely sculptured elytral disc and declivity, and by the steeper elytral declivity. Male. — Length 2.3 mm. (paratypes 1.9-2.5 mm.), 2.6 titnes as long as wide; color black. Frons moderately convex, the epistomal margin slightly elevated; a conspicuous, transverse carina occupy- ing median half at upper level of eyes; surface coarsely reticulate, the sparse punctures minute; vestiture incon- spicuous except on epistomal margin. Eye smaller than related species. Pronotum 1.2 times as long as wide; widest at base, sides almost straight, converging anteriorly slightly on posterior two-thirds, broadly rounded in front; anterior margin subserrate; sculptine as in femineus with tubercles behind summit finer, becoming obsolete at base; vestiture inconspicuous, hairlike. Elytra 1.5 times as long as wide, 1.4 times as long as pronotum; sides straight and parallel on basal two-thirds, rather abruptly roinided behind, median fourth extend- ing into a rather well-developed mucio; striae not im- pressed, the punctures fine, shallow but distinct; inter- striae about three times as wide as striae, marked by irregular lines, almost flat, the punctures minute, widely separated, becoming finely granidate on posterior fointh. Declivity convex, very steep; strial punctures obscure; interstriae 3 weakly, 9 moderately elevated. I feebly raised and terminating apically in mucro; all interstriae with a row of tubercles on upper third, minute on 2. rather coarse on 3 and last tubercle on 9, those on 3 extend to middle, on 1 two or three minute gianules scattered on lower half. X'estiture hairlike, consisting of rows of short strial, and slightly longer interstrial setae on declivity. Female. — Similar to male except evidently very slight- ly more finely sculptured; declivital vestiture longer and more abundant. Type Locality. — Volcan de Agua, Esquintla, Guatemala. Type Material. — The male holotype, female allotype, and 63 paratypes were collected at the New Neotropical Scolytidae type locality on May 19, 1964, 1,000 m.. No. 616, by S. L. Wood, from an old fallen limb about 20 cm. in diameter. It appeared that two or more generations had been completed in the same piece of wood. The holotype, allotype, and paratypes are in my collection. Hylocurus microcornis, n. sp. This species evidently is allied to cancellatus Blandford, but it differs in the female frons, in the absence of a tubercle on the male declivital interstriae 5, and evidently in the very small antennal club. Male. — Length 2.7 mm. (paratypes: male 2.7-3.0. females 2.9-3.0 mm.), 2.7 times as long as wide: color black. Frons weakly convex, ascending slightly toward epi- stoma: surface very closely, subgraniilately punctured, with a transverse carina occupying median third at level imme- diately below upper level of eye. Antennal club very slightly longer than distance equal to width of eye, 1.7 tiines as long as wide. Pronotum 1.06 times as long as wide: widest on basal third, the sides weakly arcuate and converging slightly to anterior third, then rather narrowly lounded in front; anterior margin armed by eight teeth; anterior slope asperate, posterior area closely tuberculate. the inter- striae rather coarsely reticulate; vestiture inconspicuous, hairlike. Elytra 1.9 times as long as wide. 2.0 times as long as pronotum: striae slightly impres,sed on posterior third, the punctures deep, small at base, increasing in size to- ward declivity: interstriae as wide as striae, weakly convex except near declivity, terminating abruptly at margin of declivity in a blunt, nonprojecting spine, the punctures rather fine, uniseriate. Declivity abrupt, somewhat convex within ring of subequal marginal tubercles (except those on 1 smaller); costal margin near apex subserrate: slrial punctures impressed, in rows; interstriae shining. 1 bear- ing nine small pointed tubercles. 3 with three small granules, lateral interstriae unarmed, mucro well de- veloped, reticidate. Vestiture confined to declivity, con- sisting of minute strial hair and rather long, slender bristles. Female. — Similar to male except frons above more nearly flattened, below slightly more protuberant, orna- mented on median two-thirds by a conspicuous tuft of rather long, yellow hair from near epistoma to vertex, median area immediately above epistoma smooth and shining, remaining area rather finely reticulate-granulate, punctures obscure, carina absent; anterior margin of pro- notum unarmed; tubercles on declivital margin absent: declivital striae as on disc, interstriae except 2 armed by fine tubercles, those on I. 3. 7 and 9 extending to middle of declivity, others ending on upper fourth; elytral vesti- ture finer, shorter, extending to disc. Type Locality. — Twenty-six km. east of Tex- melucan, Puebla, Mexico. Host. — Salix sp. Type Material. — The male holotype, female allotype, and 6 paratypes were collected at the type locality on June 13, 1967, 2,900 m,. No. 29, by S. L. Wood, from branches of willow. The holotype, allotype, and paratypes are in my collection. 45 Phloeocleptus plagiatus, n. sp. This species is closely allied to obscurus Wood, but may be distinguished by the much larger size, by the transverse impression of the lower frons and presence of a low median eleva- tion above, by the absence of teeth on the anterior margin of the male pronotum, and by the somewhat more strongly elevated declivital interstriae 9. Male. — Length 1.5 mm. (paratypes 1.41.8 mm.), 1.4 times as long as wide; color very dark brown, almost black in mature specimens. Frons convex above, a moderate, transverse impres- sion at level of antennal insertion; a short, longitudiani, median carina at upper level of eyes; surface smooth and shining at center below, becoming coarsely reticulate elsewhere, finely punctured toward smooth area, a few fine gianules laterally and above; vestiture inconspicuous except near epistoma. ."Antennal scape short, about two and one-half times as long as wide, bearing a small tuft of hair; club small, two straight sutures indicated by setae only at lateral margins. Pronotum 1.0 times as long as wide; sides almost straight on posterior half, converging to a feeble constric- tion just anterior to middle, rather broadly rounded on the unarmed anterior margin; anterior slope asperate to summit, summit at middle: posterior area rather coarsely reticulate, with small, isolated granules decreasing in abundance toward basal margin, granules replaced by minute punctures in lateral areas. Vestiture moderately abundant, consisting of stout hairlike setae, longer in median area behind summit. Elytra 1.4 times as long as wide, 1.4 times as long as pronotum: sides almost straight and parallel on basal two-thirds, subaciuelv mucronate behind: striae feebly if at all impressed, the punctures of moderate size and depth: interstriae shining, as wide as striae, the punctures fine, uniseriate. Declivity convex, steep: strial punctures reduced, obscured by subgranulate reticulation of entire declivital surface; interstriae each armed on upper half by a series of small tubercles, I and 3 feebly elevated. 9 more strongly raised and subcarinate on liasal half: apex gradually extended towaid poorly developed mucro. Vestiture on disc and declivity consisting of fine, erect strial hair, and slightly longer, moderately stout inter- strial bristles: more conspicuous on declivitv. Female. — Similar to male except antennal scape tri- angular and bearing a larger tuft of hair; elytral vestiture finer and somewhat shorter. Type Locality. — Seven km. west of Tepic, Nayarit, Mexico. Type Material. — The male holotype, female allotype, and 33 paratypes were collected at the type locality on July 13, 1965, about 1,000 m.. No, 240, by S. L. Wood, from cut branches of a roadside shade tree. The tunnels were in the phloem and were similar to those of caudatus Wood. The holotype, allotype, and paratypes are in my collection, Phloeocleptus nanuliis, n. sp. This species is allied to plagiatus Wood, described above, but it is easily distinguished 46 Brigham Young University Science Bulletin by the uniformly convex frons, by the armed anterior margin of the pronotum in both sexes, by the interstrial granules on the male disc, by the greater development of tubercles at the base of the elytral declivity and by the more strongly elevated interstriae 9. Mate. — Length 1.5 mm. (paratype 1.6 mm.) about 2.5 times as long as wide; color almost black. Frons convex except epistomal margin weakly ele- vated; surface subreticulate. with small isolated granules except almost smooth in median epistomal area; vestiture inconspicuous. Antennal scape short, about twice as long as wide, bearing a small tuft of hair. Pronotum 1.0 times as long as wide; outline as in plagialus but anterior margin armed by four teeth, the median pair larger; sculpture as in plagiatus, vestiture coarser. Elytra about 1.6 times as long as wide (elytra spread slightly), 1.6 times as long as pronotum; outline as in plagialus; striae perhaps feebly impressed, the punctures rather small, deep, larger posteriorly; interstriae almost as wide as striae, the punctures finely granulate (sub- vulcanate), the granules slightly larger near declivity. Declivity steep, convex; as in plagialus except all tubercles slightly larger particularly those at base of declivity in lateral areas; interstriae 9 iiuuli more strongly elevated. Interstrial bristles on declivity longer and coarser than in plagialus. Female. — Similar to male except antennal scape longer, wider, and bearing a larger tuft of hair; anterior margin of pronotum armed by two small teeth; discal interstriae not granulate; lateral declivital tubercles smaller, interstri.te 9 not as strongly elevated; declivital vestiture not as coarse. Type Locality. — Seven km. west of Tepic, Nayarit, Mexico. Type Material. — The male holotype, female allotype, and one female paratype were collected at the type locality on June 13, 1965, about 1,000 m., No. 240, by S. L. Wood, in series with plagi- atus which was described above. The holotype, allotype, and paratype are in my collection. Phloeocleplus paiuus, n. sp. This species is more closely allied to plagi- atus, described above, than to others in the genus, but it is easily distinguished by the more slender antennal scape, by the presence of several rather large punctures on the base of the pronotum, and by the reduced number of tubercles on the elytral declivity. Male. — Length 1.3 mm.. 2.6 times as long as wide; color dark brown (probably not fully mature). Frons not visible. Scape little if any wider than pedicel, about three times as long as wide. Pronotum 1.16 times as long as wide; widest behind middle, sides feebly arcuate, almost straight and parallel on basal two thirds, broadly rounded in front; anterior margin unarmed; summit in front of middle; posterior area reticulate, a few small, isolated granules near summit. several moderately large, deep punctures toward basal margin and in entire lateral area; vestiture fine, laigely abraded. Elytra 1.5 times as long as wide, 1.3 times as long as pronotum; outline and disc as in plagialus. Declivity steep, convex, shining to apex; strial punctures slightly confused; interstriae I and 2 unarmed, 2 greatly narrowed near middle. 3 armed by a tubercle at middle and another at junction with 7, 7 also bearing a small tubercle at declivital base, 9 more strongly elevated than in plagialus. Vestiture confined to declivity, consisting of very short strial and moderately long, stout, interstrial bristles. Type Locality. — Los Corchos, Nayarit, Mexi- co. Type Material. — The male holotype was col- lected 2 km. south of highway 15 on the Los Corchos road on July 10, 1965, No. 206, by S. L. Wood, from a broken branch of an unidentified tree. The holotype is in my collection. INFORMATION FOR CONTRIBUTORS Contributions to the Science Bulletin should primarily monographic in nature. For the )st part only manuscripts of approximately ty or more typewritten pages will be ac- pted. Papers will be published approximately in } order that they are received, pending avail- Uity of funds. Authors must arrange for finan- ig their publications, except that the Univer- / Press and the Hbrary of Brigham Young iversity will share the pubhcation cost pro- irtionately to thedr needs for copies of the icle printed. In the preparation of manuscripts, authors ; requested to follow the Style Manual for )logical Journals, American Institute of Bio- ;ical Sciences, 2000 P Sti-eet, N.W., Washing- i, D.C., 20006. Manuscripts must be typed on 3 side of the paper only, double-spaced with pie margins. Footnotes should be avoided. To facilitate review by referees, send to the tor the original manuscript and one carbon )y, together with the illustrations. Copies of : original illustrations may be submitted, but 1)uld be of quahty equal to the originals. Illustrations should be referred to as figiues *;ept for materials requiring inserts of special 3er, which may be called plates. Illustrations )uld be so designed as to fit when reduced 0 a one-colimin or full-page width. Special e must be taken to allow for proper reduc- Q in lettering (i.e., a 50% reduction of the ure means also a 50% reduction in the tering). Photographs should be of a glossy ish, unblurred, and showing sharp con- Ist. Line drawings should be made with black t on heavy white drawing paper, blue trac- i^ cloth, or blue-ruled coordinate paper. Use 1^ same abbreviations on line drawings as in It. Line drawings must be equivalent to a (pfessional draftman's work. Original drawings i; are preferable to photographs, even if they are large. Illustrations (Hne dravvrings or photo- graphs) should be numbered consecutively throughout the paper, and the approximate place of insertion should be indicated in the margins of the manuscript pages. Captions for illustrations should be assembled on a separate sheet, and each plate of figures must have its corresponding figure number pencilled lightly on the back. Illustrations and cuts will be de- stroyed unless their retmm is requested when proof is returned to the editor. A table title should be a short, concise state- ment of what the table pm^jorts to show, and should not include information necessary to the interpretation of the table. Every colvram in the table should carry a head identifying the data in that column; the measiue in which the data are given should be indicated at the head of each column. Tables should not be used when the same information can be given in a few lines of text, and should not duplicate informa- tion in text, graphs, or charts. Symbols (asterisk, dagger, etc.) should be used to indicate foot- notes to tables, with footnotes on the same page as the table. An abstract of less than foiu: percent of the length of the paper should be prepared. This siunmary should be imderstandable vdthout ref- erence to the body of the manuscript. The ab- stract must be on sheets separate from the man- uscript Proof should be corrected immediately on receipt and retiu-ned to the editor. Authors should leave forwarding addresses if they move from tlie address sent with the manuscript. Reprints should be ordered when the proof is retiuned. Address all manuscripts to Wilmer W. Tan- ner, Department of Zoology and Entomology, Brigham Young University, Provo, Utah. 5 • r ro V 0 J Brigham Young University Science Bulletin '■''JS. COMP. L/BRARy 200L. SEP 5 ^969 Harvard A STUDY OF THE WEEVIL TRIBE CELEUTHETINI OF THE SOLOMON ISLANDS (COLEOPTERA: CURCULIONIDAE) by VASCO M. TANNER BIOLOGICAL SERIES — VOLUME X, NUMBER 3 JUNE, 1969 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Wdlmer W. Tanner, Department of Zoology and Entomology Brigham Young University, Provo, Utah Associate Editor: Stanley L. Welsh, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: Vernon J. Tipton, Zoology Ferron L. Anderson, Zoology J. V. Beck, Microbiology Joseph R. Murdock, Botany Wilmek W. Tanner, Chairman of the Board Ex officio Members: A. Les: Sci Ernest L. Olson, Chairman, University Pubhcations A. Lester Allen, Dean, College of Biological and Agricultural Sciences The Brigham Young University Science Bulletin, Biological Series, pubHshes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from University Press, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for Hbrary exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 84601. Brigham Young University Science Bulletin A STUDY OF THE WEEVIL TRIBE CELEUTHETINI OF THE SOLOMON ISLANDS (COLEOPTERA: CURCULIONIDAE) by VASCO M. TANNER BIOLOGICAL SERIES — VOLUME X, NUMBER 3 JUNE, 1969 GlllADALCANAL I SLAND -'i.^t/Q^^r^ffAh-. . SAN C ( hrM SnUn It) ^•:,«5--. "^^i?"" BOUGAINVILLE ISLAND .^TL , ^^^*'.. t^ "■rti.rtjrtttL ', , , Rot Roir I c ■ 'J*^ ■■.'5*«o^^ - Island -^^ KolombangiTa ^ N Island 3^ J toi%. NEW GEORGIA GROUP Rendova Island, >* .U; \ New Georgia lova Islands- 'i^W' ^ ''i^'" JLiW *'4"-"'f' S \^"- Fig. 1. Map cf the Solomon Islands. By permission of the United States A-my Map Service TABLE OF CONTENTS Page INTRODUCTION 1 SYSTEMATIC CONSIDERATIONS 2 TAXONOMIC KEYS Key to the Genera of Celeuthetini Found on the Solomon Islands ; 5 Key to the species of Kietana 7 Key to the species of Platyacus 11 Key to the species of Trigonops 21 GENERA AND SPECIES DESCRIPTIONS 46 LITERATURE CITED 46 INDEX LIST OF ILLUSTRATIONS Figure Page 1. Map of the Solomon Islands II 2. Zeugorrhinus granulatus Mshl 2 3. Kietana episomoides Hllr 8 4. Kietana gressitti n. sp 8 5. Celeuthetes paganus GyU 9 6. Atactophijsis cordata Hllr 10 7. Platyacus suhalatus Hllr 13 8. Platyacus laticollis Hllr 14 9. Platyacus kolomhangarae Mshl 14 10. Platyacus malachiticus Hllr 15 11. Platyacus nigrocristatus Hllr 16 12. Platyacus decoratus Fst 17 13. Platyacus wehsteri Fst 18 14. Platyacus leveri M.shl 18 15. Female ventrites of species of Platyacus 19 I. P. laticollis; 2, P. kolomhangarae; 3. P. decoratus; 4. P. minor Mshl. 16. Hypotactus ruralis Fst 19 17. Hypotactus papillatus Fst 20 18. Hypotactus novohritannicus var. suturalis Hllr 20 19. Paratastus carbunculus Hllr 21 20. Trigonops rugosa Guer 24 21. Trigonops planicollis HUr 25 22. Trigonops paravicinii Hllr 26 23. Trigonops minuta n. sp 26 24. Trigonops irrorata Hllr 27 25. Trigonops froggutti Hllr 28 26. Trigonops solomonis Hllr 29 27. Trigonops coerulescens Blanch 30 28. Trigonops marshalli n. sp 30 29. Trigonops exophlhalmus Hllr 31 30. Trigonops becki n. sp 32 31. Trigonops becki 9 genitalia 33 32. Trigonops platessa Hllr 34 33. Trigonops platessa $ genitaha 34 34. Trigonops platessa 5 genitalia and spermatheca 35 35. Trigonops granulosa n. sp 35 36. Trigonops isabellae n. sp 36 37. Trigonops carinitlwrax Hllr 37 38. Trigonops bouguinvillensis n. sp 38 39. Trigonops guadalcanalensis n. sp 39 40. Trigonops hclleri n. .sp 40 41. Trigonops seriatopunctata Hllr 41 42. Trigonops dilaticoUis Gnth 42 43. Trigonops notaticollis Hllr 43 44. Trigonops forticornis Hllr 43 45. Trigonops gressetti n. sp 44 46. Trigonops gloriosa n. sp 45 A STUDY OF THE WEEVIL TRIBE CELEUTHETINI OF THE SOLOMON ISLANDS (COLEOPTERA: CURCULIONIDAE) by Vasco M. Tanner' INTRODUCTION This study is concerned with the genera and species of the tribe Celeuthetini known to occur in the Solomon Islands. The tribe is confined, in the main, to New Guinea and neighboring Islands. With the exception of Celeuthetes, the genera and species of Celeuthetini on which this paper is based are endemic to the Solomon Is- lands. On the basis of collections, Celeuthetes, has a wide distribution from Australia to the Moluccas. The Solomon Islands consist of eleven large islands which form two chains. The northern row of islands are Buka and Bougainville, just south of New Britain, followed by Choiseul, Santa Isabel, and Malaita. The southern group, which essentially parallels the northern row, con- sists of Vella Lavella, Kolombangara, New Geor- gia, Russell, Guadalcanal, and San Cristobal. Bouganville, the largest of the Solomon Is- lands, has an area of 3,900 square miles, while Gualacanal, the second largest, is 2,500 square miles in area. The main topographical features of the islands are the 10,000-foot active volcano. Mount Bally, on Bouganville and an 8,000-foot peak on Guadalcanal ( map. Fig. 1 ) . The climate is tropical; the average temperature is about 82 degrees Fahrenheit. During the rainy season, from January to March, the temperature and humidity are high. The weather from April to November is fairly cool. In general the climate, flora, and fauna of the Solomon Islands are simi- lar to that of Guadalcanal. The topography, di- rection of the mountain ranges, and size of the islands have influenced the floral patterns, den- sity of the plant growth, and, to some extent, determined the insect life on each of the islands (Gressitt 1961). The following observations made on the climate and plant life of Guadal- canal may be applicable to the other islands of the Solomon group. Mr. Robert C. Pendleton, who spent twenty- two months on the Solomon Islands in 1944-45, published his findings on "The Rain Shadow Effect on the Plant Formations of Guadalcanal" (1949). The following excerpts are from this study. Guadalcanal is well within the true tropic belt and the work of many plant geographers indicates that a rain forest t^pe of vegetation should be e.vpected. However, this island differs in having the major portion of the North Coast covered with coarse grass whUe a tnie rain forest vegetation occurs in the south portion ;md on the moiuitains. The main mountain ranges on the island are approximately parallel and occupy a central position. Comprehensive ground studies were made only on the north coastal plain, lxx:aiise travel to other portions was exteremeU' diffi- cult. Guadalcixnal is the only island of the group having a mountain chain at right angles to the prevailing wind direction. It is character- ized by having a rain forest on the south side and grasslands on the north side of the island. The ecological factors responsible for the grass- lands of the island are both climatic and topo- graphic. In the rain shadow insufficient rain falls during several months to support a forest. This great ecological diversity which pre- vails in the Solomon Islands, no doubt, accounts for the many species of insects found on the Islands. Although knowledge of the Celeuthetini has been accumulating since the pioneer contribu- tions of Guerin-Meneville (1841); Wallace (18.54-1862); Lacordaire (1861); Pasco (1885); Cheveolat (1885); Faust (1897); Heller (1910, 1934); Gunther (1937); E. G. Zimmerman ( 1942); and J. L. Gressitt ( 1966 a & b); the rich insect fauna of the Solomon Islands is still poor- ly known. Collecting throughout the year on all areas of the islands will, no doubt, produce many ad- 'Deparlnient of Zoology and Entomology. Brigham Young University. Provo. Utah. Bricham Young University Science Bulletin ditions to the fauna as well as contribute infor- mation concerning the economic importance and life history of the insects. The specimens that have come under my observation have been collected mainlv at a few coastal centers on the Solomon Islands. The monumental study by Sir Guy Marshall (1956) brought order out of a previously dis- organized classification of the Celeuthetini. It represents a critical enlargement of the revisional work of Faust (1897). Faust had recognized 31 genera as belonging to the tribe but had failed to discern the generic characters and groupings of the tribe. Marshall pointed out the restricted range and high endemicity of several of the genera as evidenced by the separation of the 36 species of the old wide-spread genus Trigonops. These he considered as confined to the Solomon Islands area. Several species known to occur on the islands have not been described, since they are repre- sented in collections only by a single specimen. The type material of all new species described in this study have been deposited in the collec- tions from which the specimens came to me by loan. Tvpe specimens are designated and their disposition indicated as a part of each descrip- tion. SYSTEMATIC CONSIDERATIONS The subfamily Otiorrhvnchinae," the broad- nose weevils, as now constituted, is separated into 15 tribes, genera and species manv of which are world-wide in their distribution. The tribe Celeuthetini, however, is restricted to the Pacific area, extending from Northern Australia, west- ward to Java, northward from the Celebes to the Philippine Islands, southeastward to Hal- mahera. Admiralty, Bismarck, New Britain, New Ireland, Solomon, and New Hebrides Islands. The Philippines and New Guinea seem to be the two focal areas of the tribe since of the 72 known genera 23 are endemic to New Guinea, 10 to the Philippines, and 5 to the Solomon Is- lands. The important tribal characteristics of Celeu- thetini are rostrum as long or longer than head, robust, bent downward at apex, narrowed in the middle; scrobes angular, well developed, an- tennae long, scape extending to the anterior por- tion of the prothorax, funicular club elongate; transverse sulcus separating rostrum from head; eyltra not wider than prothorax at base, with shoulders rounded or rectangular; mesepimera much reduced, remote from the base of the elytra; metepisternal suture much abbreviated front coxae more or less separated, except in some very small species; corbels of the hind tibiae placed in a more dorsal position, so that the tarsal cavity is largely, or even entirely con- cealed when the tibia is viewed on its inner surface, level of the corbel with its inner edge entirely bare of setae. The keys to the genera and species are based upon the morphological characters as revealed by careful study of specimen of each species. Since R. M. Heller described many of the species included in this study, it was necessary that specimens from the type series be made avail- able. Through the kindness of Dr. Gotz cotype specimens of all of Heller's species were loaned to me and proved to be invaluable for making comparisons and drawings. As a result specimens of 43 of the species have been studied, 23 of which were designated as Ti/pus specimens. A Typus (sic) specimen of the distinctive species Atactophtjsis cordata Hllr., rare in collections and unkno\vn to Sir Guy Marshall, was made avail- able to me. The Typus specimen is labeled Solomo Ins. T. Bodecker. No other specimens have been observed. A drawing of the Typus specimen is included in this study. Great care has been taken to have the draw- ings represent the correct proportions of the insect structures as well as the vesture. A draw- ing of the insect in toto illustrating the rostral shape and sculpturing, along with that of the eyes, antennae, prothorax, and elytra will supple- ment and clarify the accompanying descriptions. A description is a subjective appraisal of the objective material on the part of the author, which is oft-times brief and minus important at- -Recently it has been proposed, because of priority, that this subfanulv be known as Brai )ivrhininae. See Coleoptcronim Cataioeus pars 100:290-315 for a listing of the tribes. Weevil of the Tribe Celeuthetini tributes of the object. A good drawing, therefore, is more objective and thus is an important sup- plement to an insect description. Similarity of genitalia was noted after many species had been studied. Therefore, illustrations of female and male genitalia of only Trigonops becki and T. platessa are provided as represen- tative of the species of this genus. A widely de- bated subject today among entomologists is that of species classification. In order to justify the ef- forts put forth in this study I wish to present the following point of view as it relates to the pro- cedures and proposals of this study. Systematic workers acquainted with the theory and practice of present-day systematic zoology will admit that to continue to describe new species, as some have done in the past, is only to add to the great number of partially known "species" with which taxonomic literature is encumbered. The solution, however, would seem not to go to the extreme and abandon the "species level" activity as suggested by Oldroyd, ( 1966 ) . Man's knowl- edge of the animate world is largely due to the results of past practices of segregating, describ- ing, and naming of species. As a result much progress has been made in the categorization of plants and animals. It is granted that some past species taxonomy is inadequate since it is based upon a meager description of a single type-speci- men. Presently the refinement of naming a new species is based upon an analysis of a number of specimens, as to internal as well as external morphology, along with data on habitat, life history, distribution, and food habits. A descrip- tion based upon the above, and accompanied by drawings, conveys information on an objective basis. It makes possible the recording of minute details, which are free from those difficulties which are present in the verbal description. Thi.s has been the attempted approach in dealing with the species of this study. The Celeuthetini are restricted to the oceanic Pacific Islands, as pointed out above, and since so little is known about the insect fauna of the New Guinea-Solomon Islands area (Gressitt, 1961), studies following Marshall (1956), should aid in pointing up the endemicity and distribu- tion of this tribe. A study of the origin of the insect fauna of the Solomon Islands presents many interesting and difficult problems. The origin of the fauna is closely associated with the origin of the islands. Recently much interest has been manifest in the different theories of continental drift. Darlington (1965) is of the opinion "that the southern con- tinents have drifted." Additional research, how- ever, must be carried on before the picture of the past relationship of the Solomon Islands with the surrounding island complex is clarified. Along with additional knowledge as to the age and origin of the islands must be added information dealing with the geology, geography, climate, ecology, and a more complete sampling of tlie islands insect fauna. More intensive collecting, especially in the interior of the larger islands, will, no doubt, extend the range of presently known species, as well as result in the discovery of many new ones. As an example we may cite Gressitt's (1966) paper dealing with the Papuan weevil genus Gi/mnopholus ( Leplopitnae ) which points up the possibilities of intensive collecting on the mountainous areas of the islands. Prior to Gressitt's study only 14 species were consigned to Gijmnopholus. Collecting at high altitudes on the northeast interior of New Guinea resulted in his adding 32 new species to the genus. The faunal relationships of insects, amphibi- ans and reptiles of the Solomon Islands and of New Guinea suggest that at an early geological period these islands were connected. It has also been pointed out by Marshall (1956) and stu- dents of the reptiles that there is a faunal rela- tionship among species found in the Solomons, the northern islands, and Asian mainland. The accumulated information on the origin and phy- logeny of the fauna of these oceanic islands indicates that it is predominately Oriental. The following recent report on the "Pale- ogeography of the Tropical Pacific" by Menard and Hamilton ( 1961 ) supports the belief that many of the islands are oceanic: The oldest known fossil fauna from the Pacific Basin is the Middle Cretaceous reef coral-rudistid fauna from the flat tops of sea- mounts in the Mid-Pacific Mountains. A num- ber of other locaHties have been dated as early and late Eocene. There are enough of these dates to indicate that the present topography or ridge, seamounts, and islands probably origi nated during and after the CretaceoiLs. The evi- dence indicates that there were never any simken continents in the area, but the ideas concerning animal migration by island step- ping-stones (between present and former is- lands) appear to be more and more valid. The fact that many of the islands are con- tinental while others are oceanic, that some are geologically much older, also, that some have been separated or isolated and surrounded by deep water for longer periods of time, and that climatic and ecological conditions are different on many of the islands, present conditions that Bricham Young University Science Bulletin have influenced the rate of evolution, survival, and dispersal of the species of the respective islands. Too little is now known about the rich insect fauna of the Solomon Islands. In this paper 25 species of Trigonops are discussed, and of this number 7 are known to occur on Bougain- ville, and 6 are known from Guadalcanal. This study has revealed that much more col- lecting and association of the specimens with their host plants along with the gathering of eco- logical infoniiation is necessary before conclu- sions as to the extent of the fauna and its eco- nomic importance will be available. A program of this nature will necessitate segregation of the specimens into groups or species and the designa- tion by some symbol or name of each group if we are to deal intelligently with the specimens of this tribe. ACKNOWLEDGMENT This study was possible only because of the suggestions, cooperation, and specimen material made available by the late Sir Guy Marshall of the British Museum of Natural History, Herm Dr. Wilhelm Gotz of the Dresden Museum, and Dr. J. L. Gressitt of the Bishop Museum, Hono- lulu. Marshall made available specimens of Trigonops from the British Museum collection and also determined and made comments on 21 species belonging to the tribe Celeuthetini which I sent to him from the Brigham Young University Collection. My association with him at the British Museum in 1957 was most rewarding. I also wish to thank E. China, B. T. Thompson, and J. P. Doncaster, persomiel of the British Museum, who have been patient with the return of mu- seum specimens. Dr. Gotz loaned cotypes of species of Trigonops, Kietana, Atactophijsis, Col- posternum and Hoplotrigonops described by Dr. K. M. Keller, and cotypes of four species of PJatijacus described by J. Faust. I am grateful to these entomologists and staff members for their aid rendered in this project. I also extend my thanks to J. L. Gressitt, Chairman of the Entomological Staff, Bishop Museum, for a loan of specimens which he col- lected in the Solomon Islands. Thanks is also due Mr. Hugh B. Leech, Curator of Insects, Cali- fornia Academy of Sciences, San Francisco, and Mrs. Bose Ella Spilman, U. S. National Museum for a loan of specimens of the genus Trigonops. I wish to thank Dr. Carl H. Lindroth of the Department of Entomology, Zoological Institute, University of Lund, Sweden, who interceded in my behalf for a loan of specimens of Celeuthe- tini from Herm Dr. Wilhelm Gotz. The aid and help that came through the loan of type speci- mens and the cooperation of the personnel of the Dresden Museum is greatly appreciated. To the late Dr. D Elden Beck, Professor Ernest Reimschiissel, the late Harry P. Chandler, and Dorald Taylor, who were stationed on Guadalcanal and Morotai Islands in 1944 and 1945, as personnel of the United States armed forces, I express my appreciation and thanks for the results of their collecting efforts. They sent to tlie Brigham Young University, their alma mater, thousands of insect specimens. I must thank Professor J. Beuben Clark III, Miss Alberta Jacobs of the Classical Language Department, Mr. Kenneth Larson and Gundi Alt of the Zoology and Entomology Department of the Brigham Young University, for their assis- tance in the translation of some foreign litera- ture. Finally, I greatly appreciate the ability and cooperation of Professor L. Douglas Hill of the English Department of the Brigham Young Uni- versity, my artist, who has been so careful and accurate in the execution of the drawings that illustrate the species of this paper. The illustra- tions accompanving the descriptions of the 45 species dealt with in this study should add to further understanding of the Celeuthetini fauna of the Solomon Islands up to this date. Thank.s is also due Dr. H. Tracy Hall and Dr. Lane Compton of the research division of the Univer- sity for their cooperation in supplying funds for some of the art work of this project. I also extend my thanks to the members of tlie Editorial Board for the cooperation and financial aid in the pub- lication of this study Weevil of the Tribe Celeuthetini KEY TO THE GENERA OF CELEUTHETINI FOUND ON THE SOLOMON ISLANDS 1. (2 2. (3 3. (4 4. (22 5. (7 6. (5 10 18 21 20 19 18 (4 28 24. (25) Rostrum abruptly declivious dorsally at the apex. Elytra often granulate but without conical tubercles. Tibiae not carinate dorsally. Elytra with more than ten striae, sometimes confused. Rostrum with the basal sulcus broadly interrupted in the middle the ros- trum being there continuous with the frons. Head constricted behind the eyes; mentum sessile, prothorax granulate; ely- tra indistinctly striate, rather densely granulate (Bougainville Is.) . ... Zeugorrhinus Marshall Rostrum with the basal sulcus not interrupted. Front tibiae not denticulate. Joint 2 of the funicle, not or but slightly longer than 1. Meostemal process longer than the shortest distance beween the coxae. Rostrum dilated apically. Funicle stout, widest at the base narrowing distally; eyes nearly flat, not pro- jecting beyond the temples, mentum with only two setae; stria 12 on the elytra abbreviated; front coxae in the middle of the prester- num (Solomon. Isl. ) Kietana Heller Funicle slender, not narrowing distally; eyes convex, projecting. Mentum with four or more setae; front coxae in the middle of the proster- num; stria 12 abbreviated. Rostrum with the dorsal area shallowly excised, the narrowest part two to three times as wide as the apex of the scape, the declivity without scales or a dorsal elevation (New Guinea, Solomon Islands) Celeuthetes Schonh Mesostemal process as broad as long or broader than long. Front coxae nearer to the front margin of the prostemum; metastemum shorter than a median coxa. Rostrum with the basal sulcus not pro-duced dovniward in front of the eyes. Prothorax with the basal margin elevated; elytra cordate flattened; metas- stemum with the deep rugose depression on each side. ( Solomon Isl. ) Atactophtjsis Heller Prothorax with the base not elevated; elytra ovate, convex, metastemum without depressions. Rostrum with the basal sulcus produced downward in front of the eyes. Elytra with ten regular striae, but the tenth often abbreviated. All of the femora with one or more small teeth; mesostemal process transverse. Front margin of prostemum sinuate; rostrum distinctly longer than broad; frons very steep, nearly vertical, its width less than the length of an eye. 6 Brigham Young University Science Bulletin eyes directed obliquely forward; with only four visible ventrites. ( Sol- omon Isl. ) Platyacus Faust 25. (23) Femora without teeth. 26. (27) Mesostemal process much broader than long. 27. (28) Declivity rostrum squamose. 28. (29) Funicle with joint 2 not longer tlian 1. 29. (33) Eyes more or less conical, highest behind the middle mostly very strongly produced. Elytra with stria 10 approaching very closely to 9, the punctures comparatively larger in the basal, third, much smaller be- hind; five visible ventrites, ventrites 3 and 4 narrow in width (Solo- mons) Trigonops Guerin GENUS ZEUGORRHINUS MSHL. (1956) Marshall, The Otiorrhynchine Curculionidae of the tribe Celeuthetini ( Col. ) , 1956, p. 28, British Museum, London Fig. 2 Marshall based the genus Zeugorrhinus on a species collected at Kieta, Bougainville Islands, in 1937 ( J. L. Fraggott ) . As far as the writer is aware, granuJattis is the only species now as- signed to the genus. No specimens of this species have been available for study. Marshall indicates that this taxon is most nearly allied to Elytrochei- his Fst., consisting of some 20 species which are mainly found on New Guinea, Louisiade Archi- pelago and Woodlark Islands. An examination of several specimens of Elytrocheihts confinis Fst. from Oro Bay, New Guinea collected by Harry P. Chandler, 1944, confinns Marshall's state- ment as to the characteristics of the genus Elytro- clieiliis; "the basal sulcus of the rostrum is com- plete; the head is not constricted behind the eyes; the prothorax is vertically truncate at the apex; the elytra are strongly carinate at the base; and the mesostemal process is somewhat trans- verse and almost parallel-sided." Zeugorrliinus is characterized as follows: "the head constricted behind the eyes, separated from the rostrum by a deep sulcus which is rather broadly interrupted in the middle but is continuous downward later- ally at apex, the basal margin finely carinate. Mesothoracic process narrowly triangular much longer than broad." A good illustration of Z. granulatus Mshl. contained in Marshall's paper is here reproduced for the convenience of those interested in the Celeuthetini of the Solomon Islands. Fig. 2. Zeugorrhinus granulatus Mshl. - 9 Weevil of the Thibe Celeuthetini GENUS KIETANA HELLER (1910) Heller, Wien. Ent. Zeit, XXIX, 1910, p. 195 Marshall, The tribe Celeuthetini, 1956, p. .38 This genus may be characterized as follows: Head level with rostrum, separated from it by a deep curved sulcus; rostrum longer than broad, dilated at genae, apical declivity devoid of scales, dorum with two carinae and an eleva- tion at the top of declivity. Antennae scape stout, rugosely punctuate and parallel sides; funicle stout at base and narrowing distally, joints 1 and 2 equal. Prothorax about as long as broad, granu- late, not construicted at apex. Elytra broadly ovate, with 12 striae, twelfth abbreviated. Legs, with femora moderately clavate, hind pair not reaching apex of elytra. Prostemum with coxae in middle and contiguous; ventrite 2 longer than 3 and 4, 1 and 2 granulated laterally. Key to Species of KIETANA HELLER Marshal, 1956, p. 38 1. (7) Funicle with joints 4-6 transverse or not longer than broad. 2. (3) Setae on the elytra and declivity spatulate and recumbent; prothorax with kidney-shaped granules (Bougainville Is.) size 9.8 mm in length-4 mm in breadth. Fig. 3 episomoides Hllr. 3. (2) Setae on declivity of the elytra not spatulate, longer, stouter erect or suberect. 4. (5) Prothorax without punctures, the granules dense and simply convex; elytra with the indistinct granules in the striae of about the same size as those on the intervals ( Russel Is. ) size 5.5-8.0 mm., in length-2.5-4.0 mm in breadth tessellata Mshl. 5. (4) Prothorax long, feebly rounded laterally, disc flat longitudinally, granules lu- nate due to a lateral puncture from which a small brown setae issues, gran- ules in the striae about the same size as those on the intervals. (Gua- dalcanal Is.) size 8.9-9.1 mm., in length-4.0 mm in breadth. Fig. 4 gressitti n. sp. 6. (5) Prothorax distinctly rounded laterally, disc elevated near base slopping to- ward the apex; discal granules large, lunate owing to a large puncture on the inner side of each; granules in the striae much smaller than those on the intervals (Isabel Is.) size 8.0 mm in length-3.8 mm in breadth isabellae Mshl. 7. ( 1 ) Funicle with all joints distinctly longer than broad; discal granules on pro- natum lunate; setae on the elytra subrecumbent (Shortland Is.) size 8- 10 mm in length-3.7^.8 mm in breadth aluensis Mshl. Kietana episomoides Heller Wien. Ent. Zeit. 39: 195, 1910. Fig. 3 The following description was made from one cotype specimen kindly loaned by Dr. Wilhelm Gotz of the Dresden Museum. Derm black, with white scales; pronotum with narrow median stripe of denser scales; elytra with scales grouped between large granules on intervals. Head slightly punctuate up to vertex, frons convex, eyes obovate, slightly convex. Ros- trum stout and not so roughly punctuate as in some other species of Kietana; lateral and angu- lar area with long white setae. Antennae scape parallel-sided, widening slightly from base to apex; funicle joints 1-2-.3-7 equal in length; joints 4-5-6 smaller and equal, with long brown setae. Prothorax slightly longer than broad, gent- ly rounded, laterally, widest at middle; dorsum feebly convex longitudinally, granules on disc lu- nate due to a puncture on the inner side of each one; each bearing a spatulate recumbent white seta. Eh/tra ovate, longer than broad, widest at apical third, declivity abrupt, with long flat setae, disc flat with large granules irregularly placed along intervals making wavy arrange- ment, seta issuing from side of granule, small granules along stria, and with scales betvveen shining granules. Legs, femora and tibia red- Bhicham Young University Science Bulletin Fig. 3. Kietana episomoides HUr. brown clothed with white scales and setae, tarsi black; ventral area and ventrites with white scales and setae. Length 9.S mm; breadth 4 mm. Type locality: Bougainville, Kieta (Dr. Schlaginhaufen) 1908-5 tijpus, 1908-5 Staatl, Mu- seum fiir Tierkunde, Dresden. Kietana gressitti n. sp. Fig. 4 Demi black with small pale greenish scales, pronatum with a narrow median strip of scales; elytra, legs, and ventral surface with small groups of green scales. Head rugose with small punctures on apical portions of frons, diminishing toward base; eyes large, slightly convex, lateral and in contact with rostral suture; rostrum longer than head, dilated at genae, dorsum and apical declivity punctuate, with a slight elevation and fovea at junction of two carinae and declivity. Antennae scape widen- ing gradually from base to apex, punctures deep, clothed with long recumbent brownish setae and a few green scales; scape reaching the anterior two-fifths of prothorax; funicle slightly longer than scape, all segments pyrifomi, 1 a little larger than 2. Prothorax longer than head and ros- trum combined; widest at middle, sides practical- ly parallel; truncate at base and apex; disc con- vex, highest in middle, sloping to apex and base, closely set with rounded granules, from one side of which issues a small brown seta, giving gran- ules a lunate appearance, green scales sparsely placed between granules, median area more heavily covered with scales. Elytra broadly ovate, widest behind middle, dorsum convex highest before middle, punctures in striae, granules in main, along intervals, some small ones in striae paths; small brown setae arise from side of gran- ules; some scales between granules of intervals; setae long, brownish and recumbent on precipi- tous declivity. Underside, prosternum, mesoster- num, metastemum, ventrites 1 and 2 sparsely clothed with green scales and short, white setae. Fig. 4. Kietana gressitti n. sp. Weevil of the Tribe Celeuthetini Legs, femora, tibia, and coxa reddish-brown, tar- sus black with white recumbent setae. Length S.0-8.1 mm.; breadth 4.0 mm. Type locality: Solomon Islands: Holotype Guadalcanal, Gold Ridge, 500 M. June 24, 1956, (J. L. Gressitt). One paratype, same data as the Holotype; one paratype Guadalcanal Is. Gold Ridge, March 21, 1955, E. S. Brown. Holotype, deposited in the Entomological Collection Bishop Museum, Honolulu, Hawaii; one paratype in the Entomological Collection, British Museum of Natural History, London; and one paratype in the Entomological Collection, Brigham Young University, Provo, Utah. Remarks: The species gressitti is black with a narrow prothorax, similar in shape to that of tesseUata, but is larger, with green scales, and more deep punctures on the dorsal area of the rostral declivity. K. ahiensis in contrast to gressitti has a more rugose and rounded prothorax, and is larger with conspicuous white scales on body and legs. The rostral carina and elevation at top of the declivity is distinctive in each of these species. GENUS CELEUTHETES SCHONH. Schbenheer. Gen. Spec. Cure. VII, 1, 1843, p. 250. Rostrum as long as the head, robust, abruptly declivious dorsally at apex; declivity without scales or a dorsal elevation, scrobes deep, triangu- lar, dorsal area shallowly excised, narrowest part two to three times as wide as apex of scape. An- tennae submedium, rather long and slightly ro- bust, scaled and with setae; scape subcylindrical, straight except for slight arch at base; funicle segments 1-2 elongate, 1 longer than 2, 3-5 sub- pyrifonn, 6-7 subglobular, club oval. Eijes rather large, round, protruding. Prothorax subcylindri- cal, slightly rounded at sides; tnmcate at apex and base. Elytra short and moderately rounded; concave dorsally in cT or regularly convex in 2 . Elytra not wider than prothorax at base, with shoulders rounded or rectangular; often granular, but without conical tubercles. Mesostemal pro- cess longer than shortest distance between coxae. Corbel of posterior legs cavernous and claws on tarsus free. Celeuthetes paganus Gyll Gyllenhal in Schonh. Gen. Spec. Circ. II, 2, 1834, p. 539. Fig. 5 Denn reddish black, with small white scales and setae which arise from small polished gran- ules. Head and rostrum as long as prothorax; ros- trum with a furrow, narrowest part three times as wide as apex of scape, declivity without scales or a dorsal elevation; a small fovea at base of rostral groove; rostral sulcus extending laterally and forward to anterior margin of eyes; rostrum two-thirds as long as head. Antennae scape slight- ly bowed, same diameter throughout, extending to anterior one-third of prothorax; segments 1-2 of funicle as long as 3-6; segments 3-7 globoid, eyes large, round, flattish and placed well down on lateral portion of head. Prothorax as long as wide; apex slightly concave, base truncate, trans- Fig. 5. Celeuthetes paganus Gyll. 10 Brigham Young University Science Bulletin versely convex, widest at middle, disc and lateral portions granulate and punctate. Eh/tra widest at middle, one-fourth longer than wide, stria 12 abbreviated; intervals on disc twice as wide as striae, interval 6 units just back of humerus with 5 and 7 fomiing a low obscure carina, (there is also a union of these intervals at the declivity which results in fonning a low keel which ex- tends to tlie apex) shallow closely set punctures in striae, small shining granules with decumbent setae on intervals, scales and setae abundant on declivity down to apex. Legs reddish, thickly covered with white scales and setae. Prothoracic coxae large and narrowly separated, ventrites 1-2 rugose laterally free from punctures and setae in middle; 3-5 free from scales and setae. Length 7.8 mm; breadth 3.2 mm. Type locality: Santa Cruz Islands, Vanikora Island. Specimen studied: One from the Solomon Islands, Guadalcanal. Teneru River, (D E. Beck), 1944. Bougainville Is. 2 specimens, June 5, 1944, (A. B. Gumey); two specimens, June 30, 1956, Simba Mission (E. J. Ford, Jr.). The species assigned to Celeiithetes are wide- ly distributed from Australia to New Hebrides, New Guinea, Bismarck, and Solomon Islands. The species paganus is the only member of this genus thus far reported from the Solomon Is- lands. GENUS ATACTOPHYSIS HELLER Heller, Verb. Naturf. Ges. Basel XLV, 1934, p. 9. Body depressed, with few scales, rostrum as long as wide, dorsal without keel, basal sulcus deep and crescent shaped, eyes not in contact with rostrum, due to deep sulcus. Antennae scape cylindrical, straight, apically knot shaped; not extending to middle of prothorax, segments 1-2 of funicle elongate; equal in length to segments 3-5 combined; club as long as segments 5-7 com- bined. Frothorax as wide as long, base and apex truncate, convex transversely; with median longi- tudinal elevation and shallow close set punctures. Elytra with eleven striae, depressed, heart- shaped, humerus ec|ual to base of prothorax, sloping from base to obtuse point, in contact with coxa of metathoracic leg, apex acuminate. Femora extending slightly beyond apex of elytra. ATACTOPHYSIS CORDATA HELLER Heller, Verb. Naturf. Ges. Basel XLV, 1934, p. 9. Fig. 6 Derm black with bluish-white scales, sparse white setae, prothorax finely punctuate, elytra with rows of granules between interspaces. Head as long as rostrum, vertex rugose; ros- tral sulcus deep, crescent shape; antennae scape narrow, except at apex, practically devoid of scales and setae, funicle segments 1-2 as long as 3-5, club as long as 5-7; funicle devoid of any vesture; rostral base broad with deep punctures, no carina, but with a slight elevation at rostral declivity, scrobes large, separated dorsally by about width of apex of scape. Eyes lateral, round, flattish and contiguous with rostral sulcus. Pro- thorax greater in breadth than length; base and apex truncate, disc convex transversely punc- tures shallow and irregular in placement; low median carina, a few white scales on lateral mar- gin, body scales denuded, except for a few scattered bluish and white ones. Elytra 4 mm in width at base, 7 mm in greatest width and 7.6 mm in length; disc flat, heart-shaped; striae con- sisting of shallow punctures and rows of small polished granules. The 11 intervals smooth, on discal area practically free of scales; posterior margin emaginate, gentle slope at declivity, cov- ered with more scales and deeper punctures; an- terior lateral margin of metathorax impressed and with deep punctures, lateral margin of elytra in contact with coxa of metathorax leg. Legs elon- gate, prothoracic very narrowly separated; meta- thoracic coxa widely separated. Ventrites 1-2 ru- gose laterally with fine wavy lines in medial area, 5 greater in width than 3-4 combined. Fig. 6. Atactophysis cordata Hllr Weevil of the Tribe Celeuthetini 11 Length 13.5 mm; breadth 7 mm. Type locality: Solomon Islands (T. Bodeck- er) Tijplius, Stattl. Museum fiir Tierkunde, Dres- den. The genus Atatophijsis is characterized by its elytra which are flattened and heart-shaped. It is widest where the lateral margin of the elytra comes in contact with the coxa of the metathor- acic leg. The 11 striae, the rows of small, pol- ished granules, the smooth intervals, shallow punctures and acuminate apex of the elytra, along with the deep rostral sulcus, deep punc- tures on the base of the rostrum, slender scape of the antennae, and the flat eyes which come in contact with the lateral extension of the rostral suture are all outstanding characters. I have been unable to compare this genus with Atactus be- cause of the lack of correctly determined speci- mens. GENUS PLATYACUS FAUST (1897) Platyacus Faust, 1897, Stett, Ent., Zeit. 58: 236, 270 Colposternum Heller, 1910, Wien., Ent. Zeit. 29: 191 Hoplotrigonops Heller, 1934, Verb. Naturf. Ges. Basel 45:21 This genus, peculiar to the Solomon Islands, is characterized by the presence of several small teeth on the femora, a distinct modification of the fourth ventrite, ( Figs. 1-4), a sinuation of the front margin of the prosternum, the steep frontal part of the head, with the eyes closely approxi- mate, and a long slender rostrum with an ele- vated straight keel. Through the kindness of Dr. Wilhelm Gotz of the Dresden Museum I have seen cotype specimens of Platijacus siibalatus ( Hllr. ) ; P. lati- collis Hllr.; P. malachiticus Hllr.; P. nigrocristcitus Hllr.; P. decoratus Fst.; P. ivebsteri Fst.; Hijpo- tactus ruraJis (Fst.); H. papillatus (Fst.); H. novobritannicus var. suturalis Hllr.; and Paratac- tus carbtincuhis Hllr. Drawings have been made of all the species of Platyacus as listed above, also of Hijpotactes ntralis, papillatus, and novobritannicus var. su- turalis Hllr., and Paratactus carbunculus Hllr. Both Faust and Heller were not sure as to the generic characters of some of the species of this complex. Marshall's analysis of the genera Platya- cus, Colposternum, and Hoplotrigonojis resulted in his concluding that the two latter ones were synonyms of Platyacus. He also observed that the two Faust species Platijacus ruralis and papillattis were not congen- eric with the genotype of Platyacus {tvebsteri Fst). "They differ from Platyacus in the much shorter rostrum, the truncate (not sinuate) front margin of the prosternum, the gently sloping frons, the width of which is as great as, or greater than, the length of an eye, and the more lateral position of the eyes. The female has only four visible ventrites." Marshall thus erected the genus Hypotactus for these two species. I have con- cluded, from my study that P. novobritannicus var. suturalis Hllr. also belongs in Hypotactus. The species of Platyacus are, at this stage of our knowledge, recorded only from the Solomon Islands, while species of Hypotactus have been reported only from the Bismarck Archipelago. The cotype specimen Hypotactus novobritanni- cus var. .suturalis Hllr. bears a locality label "N. Pommem" now known as New Britain. The other tv\'o species ruralis and papillatus are also from New Britain. This leaves the species carbunculus Hllr. which does not belong in Platyacus, but has been assigned to Paratactus by Marshall. This study is concerned only with the celeu- thetini of the Solomon Islands; but since Hypo- tactus ruralis, papillatus and var. .suturalis and Paratactus carbunculus have been dealt with as species of Platyacus or in genera now considered as synonyms of Platyacus and since they are not well represented in most entomological collec- tions, drawings of cotypes from the Dresden Mu- seum are included. It is my belief that illustra- tions of rare species will be an aid in future stud- ies of the weevils of the Solomon Islands. 1. 2. (4) (3) KEY TO THE SPECIES OF PLATYACUS Sir Guy Marshall's Key to the species of Platyacus. Marshall notes that the species subalatus and laticollis of this Key were known to him from the description only. Prosternum with a projecting tooth on each side of the apical sinuation. Elytra with the dorse-lateral margin raised into a comb-like ridge in the middle two-thirds, the intervals without granules; funicle longer than the scape (Bougainville I.) Fig. 7 subalatus Hllr. 12 Brigham Young University Science Bulletin 3. (2) Elytra without such a ridge, the intervals granulate; funicle shorter than the scape (Bougainville I.) Figs. 8, 14 laticollis Hllr. 4. ( 1 ) Prostemum without teeth on the front margin. 5. (20) Rostrum slightly widening apically, or at least parallel-sided, the scrobes comparatively large, ill-defined behind, the narrowest space between them not wider than a scape. 6. (7) Elytra with a distinct caUus on intervals 4 at the top of the declivity, the apical lateral margin deeply excised in 5 ( Kolombangara I.) Figs. 9, 14 kolombangarae Mshl. 7. (6) Elytra with no posterior callus in interval 4, the apical lateral margin straight or shallowly sinuate. 8. (11) Elytra with a callas or short ridge at the top of the declivity on interval 5. 9. (10) Prothorax rounded laterally, flattened on the disk; tibiae with a fine dorsal carina on the basal half; tarsi with joint 2 transverse; elevations on interval 5 of the elytra without suberect black setae; length 6.0-7.5 mm. (Bougainville I.) Fig. 10 malachiticus Hllr. 10. (9) Prothorax parallel-sided in the basal half, transversely convex on the disk; tibiae without any dorsal carinae, tarsi with point 2 somewhat longer than broad; elevations on the elytra witli short black suberect setae; length 4.5-5.0 mm. (Bougainville I.) Fig. 11 nigrocristatus Hllr. 11. (8) Elytra without any trace of a callus or ridge on interval 5. 12. (19) Scape rapidly widened close to the base and then parallel-sided to the apex, compressed, with tlie lower edge sharply carinate. 13. (14) Elytra with bands of yellowish-green scales; a complete, basal band which emits three backward elongations along the suture and the sixth in- tervals, a much shorter band behind the middle extending to stria 4 and broadly interrupted at the suture, and a band covering the whole apical area except the suture; scape narrower and dilated apically (Isabel I.) Figs. 12, 14 decoratus Est. 14. (13) Elytra without bands, the green or yellowish scales more generally distrib- uted; scape thicker, almost parallel-sided from near the base to the apex. 15. (16) Elytra with the scales sharply confined to the intervals, the bare striae (as broad as the intervals) appearing like black stripes; ventrite 4 of 9 with the processes at the external apical angles only half as long as the median process (Alul.) Fig. 13 websteri Fst. 16. (15) Elytra with the green scales covering the septa between the punctures in the striae, the bare punctures producing a tessellated effect, with no sign of stripes; ventrite 4 of 9 with the external apical processes nearly as long as the median process. 17. (18) Prothorax rounded laterally (Florida I., Giza I.) Fig. 15 leveri Mshl. 18. (17) Prothorax parallel-sided in the basal half (Fauro I.) fauronus Mshl. 19. ( 12) Scape more slender, gradually widening from base to apex, not compressed, nor carinate on the lower edge (Bougainville I.) Fig. 14 minor Mshl. 20. (5) Rostrum gradually narrowing from base to apex, the scrobes small, ovate, sharply delimited behind, the space between them much wider than a scape (Bougainville I.) narinus Mshl. Weevil of the Tribe Celeuthetini 13 Plattjaciis suhalatus Hllr. Heller, Wien. Ent. Zeit. XXIX, 1910, p. 194 Fig. 7 Derm black, except the proximal portions of the femora which are red, with small green and white scales, irregularly and sparsely placed on dorsum of body, antennae and femora. Rostrum much longer than head, parallel- sided dorsum narrow, carina with small eleva- tion at top of declivity; separated from head by sulcus which extends lateral and anterially to scrobes; antennae long, funicle reaching middle of prothorax bowed slightly, same diameter throughout, except for rounded distal end, funi- Fig. 7. Plahjaciis suhahitus HUr. cle segments all elongate. Prothorax, prostemum with a projecting tooth on each side of apical sinuation, wider than long, widest posterior of middle, truncate at base and apex, slight con- striction at apex; disc flat, rugose, with varying- sized granules, no punctures or setae, no marked median area, and few small scales. Elytra with dorso-lateral margin raised into a comblike ridge in middle two-thirds, intervals without granules, striae punctured, short white setae at apex. Legs, femora red in color on proximal and club areas, reaching just beyond apex of elytra and with small tooth on each femora. Tibia straight with long white setae. Ventral body and ventrites devoid of sailes, ventrites with short suberect setae, ventrite 5 longer than .3-4. Length 9.8 mm; breadth 5.2 mm. Type locality: Bougainville Island-Kieta. The above description was made from the following specimen: Bougainville - Kieta (Dr. Schlagin- haufen), 1908-5; Typus, Staatl. Museum fiir Tier- kunde, Dresden. Other specimens studied: Solo- mon Islands: Bougainville Islands, Kokure, Nr. Crown Prince Ra. 900 M. VI-8 1956 (J. L. Gressitt ) . Plattjacus laticoUis Hllr. Heller, Wien. Ent. Zeit. XXIX, 1910, p. 193 Fig. 8, 14 Derm black with white scales, some irides- cent. Rostrum twice as long as head, narrow in middle, expanded at apex, carina ending in U- shaped elevation at declivity which is finely punctuate. Sulcus separating rostrum from head circular in shape. Antenna scape enlarged at distal end, reachmg middle of prothorax, funicle shorter than scape, segments elongate, club as long as segments 5-6-7 combined. Prothorax, pro- stemum with projecting tooth on each side of apical sinuation; more than one-fourth wider than long, widest back of middle, base truncate, apex concave, disc flat, slightly concave along medial area, with granules. Elytra widest near base, broadly ovate and acuminate behind, inter- vals 3-4 slightly elevated at declivity, punctures along striae and granules on interval, granules on intervals 3-4 at declivity prominent, with short setae, white scales in irregular transverse rows across elytra. Legs, femora not reaching apex of elytra, one large and several smaller teeth on each of femora, no scales, but few setae on un- derside of body; fourth ventrite of female deeply bimarginate forming a median lobe as shown in Fig. 14, segment 3 is sparsely covered with setae. 14 Brigham Young Univebsity Science Bulletin Platyacus kolombangarae Mshl. Marshall, Otiorrhynchine Curculionidae, Tribe Celeuthetini, 19.56, pp. 68-70, Fig. 25, British Fig. 8. Platyacus laticollis HUr. Length 10.2 mm; breadth 5.0 mm. Type locahty: Bougainville Lsland. The above description from one specimen from Salamo In- seln-Kieta 1910, 12-typus-Staatl. Museum fiir Tierkunde, Dresden. Remarks: The fourth ven trite of kolomban- garae Mshl. is very different to that of laticollis. This is a good 9 character which may prove to be of aid in separating species of this genus. Fig. 26: p. 70 of Marshall's "Otiorrhynchine, Curcul- ionidae of the tribe Celeuthetini" is mislabeled, it should be venter of Platyacus kolombangarae Mshl. Mus., Nat. Hist. Fig. 9 Denn black with dense green scales, except in the middle of the venter. Rostrum narrow, carina ending at declivity without an evident elevation, sulcus V-shaped; antennae wide from origin to apex, slightly bowed, with dense green scales and decumbent black setae, scape reaching slighdy beyond mid- dle of prothorax; funicle a little longer than scape, segments elongate, especially 1-2. Prothor- ax wider than long, rounded laterally, slightly constricted near apex, widest behind middle, base and apex truncate, disc convex with small black granules and dense green scales, median area more densely set with scales, punctuate on pleurae. Elytra of female ovate, acuminate be- hind with lateral margin sinuate near apex, a distinct sub-apical elevation near apex of inter- val 4; male without callus, or margin sinuation, shortly acuminate behind and with apex pro- duced downward; dorsum convex, striae with deep naked punctures, intervals between striae covered with dense green scales which obscure Fig. 9. Platyacus Kolombangarae Mshl. Weevil of the Tribe Celeuthetini 15 small granules with short setae. Legs, black and reddish-brown, all femora with a small tooth, tibia straight. Covered dorsally with green scales. Ventrites without scales, except 1 and 2 with scales laterally and along posterior margin of 2. Fourth ventrite of female modified as shown by Sir Guy Marshall. Length 5-6 mm; breadth 3.0-3.6 mm. Type locality: Solomon Islands: Kolomban- gara Island. Locality of specimens studied: New Georgia Group, Kolombangara, Island Kukundu, S. W. Coast 1-12 M, 10 VII, 1959, (J. L. Gres- sitt). New Georgia Group: Nr. Egolo, 1-2.5 m. July 16, 1959, (J. L. Gressitt) Freijcinetia. Solo- mon Islands: Kolombangara-Iri-iri; 27 VII, 1958, (P. G. Fenemore); Vella Lavella, Tiopari, 26 VII, 1958, (P. G. Fenemore); Choisaul, LuH, 1 VIII, 1958, (P. G. Fenemore). Remarks: Dr. Gressitt collected specimens of kolomhangarae on the following plants; Calo- fhyllum, Freijcinetia, Flagellaria, and the palm. Platyacus malachiticus Hllr. Heller, Wien. Ent. Zeit. XXIX, 1910, p. 192 Fig. 10 Derm black with green and some iridescent scales. Rostrum short, expanded some at apex, car- ina acute, small elevations at declivity, declivity scaleless, and punctate; antennae scape slightly bowed, same in diameter throughout, reaching beyond the middle of prothorax, punctate and covered with scales and decumbent setae; fun- icle segments elongate. Two-longer than 1. Head shorter than rostrum, rugose and punctate to the apex; eyes round, slightly convex, frons, between eyes as wide as diameter of eye; sulcus separat- ing head from rostrum circular. Prothorax 3.2 mm in breadth, 2.3 mm long, constricted near apex, widest a little behind middle, truncate at base, slightly concave at apex, punctate on pleu- rae, disc flat, with black shiny granules and punc- tures obscured by green scales. Prostemum has a slight sinuation and small projection which resembles a tooth. Elytra broadly ovate, widest before middle, elevation on interval 5, without subrect black setae, sharply acuminate behind narrow declivity, dorsum only slightly convex transversely; shallow striae with punctures, in- tervals with cluster of scales and small black granules, small recumbent white setae. Legs proximal portion of femora reddish; anterior femora with one large and several small teeth. Ventral surface with setae devoid of scales; ex- cept lateral portion of metastemum; ventrites scaleless, 4-3 times as long as 3, nonnal in shape. Length 9 mm; breadth 5.0 mm. Type locality: Solomon Islands, Bougainville Islands. The above description from one speci- men from Bougainville Island-Kieta ( Kapt. Kurtz). 1908-3-Typus Staatl. Museum fiir Tier- kunde, Dresden. Other specimens studied: Bou- gainville Is.-(S.) Kieta, XI. 30 1959, (T. C. Maa), Bishop Museum Bougainville Is. Pukpuk, Nr. Kieta VI 26, 1956, (E. J. Ford, Jr.). Bougain- ville Is.-Kaure 690 m. VI -8-18-1956 (J. L. Gressitt). Kokure, Nr. Crown Prince Ra. 900 M. VI-10-1956 (J. L. Gressitt). Remarks: The specimen of malachiticus from Dresden has a small projection on the proster- Fig. 10. Platt/acus muUicliiticus Hllr. 16 Brigham Younc University Science Bulletin nal apical sinuation, likewise, some specimens collected by Dr. Gressitt. This should be con- sidered when using the above key by Sir Guy Marshall. Platyaciis nigrocristatus Hllr. Heller, Verb. Naturf. Ver. Basel XLV, 1934, p.22 Fig. 11 Derm brownish, black, with dense white scales. Rostrum a little longer than head, paral- lel sides, carina distinct, scales dense with small black granules showing through, slight eleva- tion at declivity which is reddish-brown with sparse white setae, scrobes small separated by a greater width than diameter of antennae scape. Antennae slightly bowed, gradually increasing in diameter to apex, densely covered with white scales and decumbent black setae, extending be- yond middle of prothorax, funicle a little longer than scape; segments 1-2-3 as long as segments 4-5-6-7 combined. Head densely covered with scales, a frontal carina which meets rostral ca- rina at curved sulcus; punctate extending to apex of head, granules interspersed among scales; eyes large, ovate, space between them less than diameter of eye. Prothorax 1.6 mm. long, 1.8 mm. in breadth, parallel sided in basal half, transver- sely convex on disc with granules and punctures, scales on septa around punctures (except in medial area which is sparsely covered); base and apex truncate, pleurae punctate and densely cov- ered with scales; a short white setae issuing from each puncture. Elytra, greatest width just before middle, shallow striae with punctures similar to those on prothorax; white scales on septa between punctures and on intervals; slightly convex at middle becoming concave between black elevations on fifth interval, shortly acumin- ate behind with apex produced dovmward. Legs brownish covered with white scales and setae; all femora with a large brown-colored tooth; venter and ventrites 3-4-5, with white decumbent setae, ventrite 4 of 9 sculptured similarly to that of webesteri Faust. Length 5.0-6.S mm; breadth 3.0-4.0 mm. Type locality: Solomon Islands, Bougainville Islands. This description was made from one specimen from Bougainville, 30-5-60, 1926-5 Plat- tjacus (Hoplotrigonops) nigrocristata L'N H typus 33 (the genus and species is in Heller's hand writing) Staad. Museum fiir Tierkunde, Dresden. Other specimens studied: Solomon Islands: Bougainville Is. (5). Kieta, XI. 27, 1959, (T. C. Maa). Bishop and Simba Mission, June 29, 1956, (E.J. Ford, Jr.). Fig. 11. Plati/acus nigrocristatus Hllr. Platyacus decoratus Fst. Faust, Stett. Ent. Zeit. LVIII, 1897, p. 272 Fig. 12 Derm with black small blue-green and yel- lowish scales. Rostrum, sides parallel, one and one-half time as long as head, carina with small elevation at declivity; antennae scape narrow at origin, en- « Weevil of the Tribe Celeuthetini 17 Fig. 12. Platyucus decoratus Fst. covering entire apical except suture, striae with larged at apex, reacliing antenior half of prothor- ax, funicular segments 1-4 pyriform, 5-6-7 diam- eter equal to lengthy club slightly longer than preceding three funicular segments. Head short, with small blue scales and irregular wavy ele- vated lines from between eyes to apex, frons be- tween eyes narrower than width of eye. Eyes frontal and moderatley convex. ProtJwrax wider than long, base and apex truncate, slightly con- stricted near apex, widest before middle, trans- versely convex, disc rather rugose, granules showing through scales which obscure punc- tures. Medial line with dense green scales. Elytra ovate, sharply acuminate especially in 9 with bands of yellowish green scales, a complete basal band from which extends three backward elongations along the suture and 5-6-7 intervals, a shorter band behind middle extending to stria 4 and broadly interrupted on suture and a band punctures, septa and intervals with small blue and green scales. 9 with tuft of setae on suture of declivity. Small granules with short setae on intervals. Legs black with blue and green scales on femora with a large tooth and several smaller ones. Tibia slender, straight, and long. Venter covered with scales, except middle area of ventrites 1-2 and all of 3-4-5. Ventrites 4 with modified fourth segment ( Fig. 14 ) . Length 5.1-6.5 mm; breadth 2.9-3.2 mni. Type locality: Solomon Islands: Isabel Is- land. The above description was made from one specimen with the following notation on the small gold-colored square label on the pin - Isa- bel I. (Webster). Decoratus Faust; Coll. J. Faust, Ankauf 1900 - Typhus - Staatl. Museum fiir Tierkunde, Dresden. Other specimens stu- died: Solomon Islands - New Georgia Group, N. Georgia Is. - Munda 1 - 30 M. VII - 15 - 1959 (J. L. Gressitt) acalypha. Remarks: The fourth ventrite of the 9 is distinctive. Fig. 14. Platyacus loebsteri Faust Faust, Stett. Ent. Zeit. LVIII, 1897, p. 270 Fig. 13 Derm black, clothed with a mixture of green- white and blue scales or entirely all blue scales. Rostrum parallel-sided and almost twice as long as wide; dorsum narrow with a central carina, bordered on each side with a small ca- rina, very small elevation at top of declivity; separated from head by a sulcus. Carina of rostrum continuous with carina of frons. Anten- nae long, scape reaches beyond three-fifths of prothorax; slightly curved, as wide at base as apex, funicle segments elongate, segments 1-2 as long as 3-4-5 combined, club long and slender; scales on dorsal portion of scape. Prothorax wider than long, widest at middle, truncate at base and apex, slightly constricted at apex; dorsum flat toward base, closely set with small granules and punctures, median line densely covered with scales. Prostemum without teeth on front mar- gin. Eh/tra widest at anterior third, gently slop- ing to apex, 10 striae with deep punctures, dis- tinctly separated from intervals which are cov- ered with rows of scales. Declivity and apex practically devoid of setae. Legs, femora reach- ing tip of elytra, basal portion of femora narrow, club not robust, teeth on all femora, tibia straight, ventral surface of body covered by scales except median portion of ventrites 1-2 and all of 3-4-5 with densely covered fine setae. 18 Brigham Young University Science Bulletin ? websteri Faust; Coll. J. Faust, Ankauf, 1900, Typus, Staatl. Museum fvir Tierkunde, Dresden. Other specimens studies: Solomon Islands; Buka Islands: Gagon, 40 M. VI - 15 - 1956 (J. L. Gres- sitt). Remarks: The statement in the key, "ventrite 4 of 9 with the processes at the external apical angles only half as long as the median process," could not be observed on specimens listed above. The one specimen labeled "Solomonus ?" is well marked with blue scales in contrast to the speci- men labeled "Alu I - Shortland Is." which is cov- ered with scales colored green-white and with some blue ones. Fig. 13. Phiti/dcus iieb.itcri Faust. Length 6.0-6.2 mm; breadth 2.9-3.0 mm. Type locality: Solomon Islands: Alu Island (Webster). The above description was made from two specimens; one with a small gold-col- ored square, Alu I - Shortland Is. (Webster); websteri Faust; Coll. J. Faust, Ankauf 1900; Typus, Staatl. Museum fiir Tierkunde, Dresden, one with small gold-colored square; Solomonus Fig. 14. Female ventrites of species of Platyacus. ( 1 ) P. laticollis Hllr.; (2) P. Kohmhangasae Mshl.; (3) P. dccoratus Fst.; (4) P. minor Mshl. The fourth ventrite is structurally distinctive. GENUS HYPOTACTUS MARSHALL Marshall, the Otiorrhynchine Curculionidae of the tribe Celeutlhetini '( Col. ) , 1956; p. 66, British Museum, London. The genus Hi/potacttis was erected for two species, Platyacus ruralis and papillatus Fst. As noted above I have referred P. novobritannicus var. suturalis Hllr. to this genus. These species are recorded only from the Bismarck Islands. Species of Platyacus are known only from the Solomon Islands. Weevil of the Tribe Celeuthetini 19 Fig. 15. Phtt/acus leveri Mshl. Fig. 16. Hypotactus ruralis Fst. 20 Bricham Young Univebsitv Science Bulletin Fig. 17. Hi/pntacttis papilliitus Fst. Fig. 18. Hypotactus novohritamictis var. suturalis HUr. Weevil of the Thibe Celeuthetini 21 Fig. 19. Paratustus carbunculus Hllr. GENUS TRIGONOPS GUERIN (1841) Guer. Rev. Zool. 1841, 10. 128. The genus Trigonops, so far as now known, is confined, in the main, to the Solomon and New Hebrides Islands. It was first recognized and described by Guerin in 1841, the genotype being nigosa Guerin. The saliant characteristics of the genus are: Body and prothorax globular; rostrum abruptly declivous dorsally at apex, declivity of rostrum squamose; scape similar in both sexes, not com- pressed, subcyltndrical, very gradually widening from base to apex; funicle with joint 2 not longer than 1, eyes highest behind middle, mostly very strongly produced; elytra often granulate but without conical tubercles, with ten regular striae, but tenth often abbreviated and approaching very closely to 9, punctures comparativelv large in basal third, much smaller behind, female in many species with tuft of setae on posterior de- clivity of elytra. Tibiae not carinate dorsally; metastemal suture, visible only near its base; femora without teeth; mesostemal process much broader than long; legs rather long with femora swollen toward extremity. Sir Guy A. K. Marshall in his study of the tribe Celeuthetini divided the thirty-six species of the old widespread genus Trigonops as listed in the Coleopterorum Catalogus, pars 160, 1937, pp. .31.3-315, into six genera thus leaving only fifteen species as typical Trigonops, these he considered as confined to the Solomon Islands area. Since K. M. Heller of the Dresden Museum had described eleven of the fifteen species it was most necessary that specimens from the type series be made available for study. Through the kindness of Herm Dr. Wilhelm Gotz, Haupt- direktor, Stattliches Museum fiir Tierkunde, Dresden, a loan of cotype specimens of all of the Heller species was made available. Drawings and comparisons with specimens from the assembled collections of Trigonops from the Solomon Is- lands were made. I have studied specimens of all twenty-five species included in this report, with the exception of vitticoUis Fairm, which is unknown to me. Since some species are rare and not in most collections, much care has been taken to provide a carefully prepared illustration of each species dealt with in this genus. KEY TO SPECIES OF GENUS TRIGONOPS GUERIN 1. Elytra punctate with longitudinal series of strongly elevated tubercles, close to each other, fomiing lines or ribs between which are a series of smaller tubercles; prothorax rugose; size 5.5-6.5 mm in length-3.5-3.6 mm in width. Solomon Islands, genotype. Fig. 20 rugosa Guerin - Elytra without strongly elevated tubercles, close to each other, forming lines or ribs 2 22 Brigham Young University Science Bulletin 2. Humeral margin of elytra, on both sides, with an outward projecting tooth 3 — Humeral margin of elytra without an outward projecting tooth 4 3. Prothorax flattened on the disc; sides strongly rounded, wider than long. Size 5.5 mm in length-3 mm in width. Kolombangara, Western Group. Fig. 21 . phnicoUis Hllr. — Prothorax not flattened, sides moderately rounded; rugulae; wider than long. Size 5.0-6.5 mm in length-2..5-3.0 in width. New Georgia Island. Fig. 22 paravicinii Hllr. — Prothorax longer than wide, reduced at apex; with large gray and greenish scales; small species, size 3.0 mm in length-1.5 mm in width. Shortland Is- land. Fig. 23 minuta n. sp. 4. Latemal margin of prothorax in basal half not angular; longer than wide, taper- ing from base to apex; with deep punctures; elytra with deep punctures; small iridescent scales. Size 5.0 mm in length-2.9 mm in width, Bougainville Is- land. Fig. 24 irrorata Hllr. — Lateral margin of prothorax in basal half angular; as long as wide; coarsely and deeply punctured; lateral border with a sharp edge; scales green; elytra ro- timdate, punctures regular and shallow. Size 5.0 mm in length-2.5 mm in width. Bougainville Island. Fig. 25 salomonis Hllr. 5. Eyes blunt and conical, the vertex shifted toward the back 6 — Eyes round, less conical 8 6. Prothorax as wide as long, punctures not as large as the interspaces; elytral punc- tures deep, small keel on fourth interval at declivity on the d" . Size 5.0-^.6 mm in length-2.7-2.9 mm in breadth. Russel Island. Fig. 26 froggatti Hllr. — Prothorax wider than long, sparsely punctate, elytral punctures shallow; deeply emarginate at apex. Size 7.2 in length-3.5 mm in breadth. Solomon Islands. Fig. 27 coerulescens Blanch. 7. Prothorax wider than long; surface closely tuberculate, tips of tubercles black, shining; scales sparsely placed on the interspaces; elytra longer than wide; punctures deep with clusters of scales and small black tubercles showing through from the intervals between the punctures. Size 4.5-5.2 mm in length-2.5- 2.0 mm in breadth. Santa Isabel Island. Fig. 28 marshalU n. sp. 8. Rostrum narrow, as long as head; sulcus deep and more crescent shaped; eyes prominent; prothorax wider than long, convex; keel on 4-5 intervals of elytra at declivity; size 6.0-7.1 mm in length-3.0-3.8 mm in breadth. Malaita Island. Fig. 29 exophthahnus Hlli. — Rostrum narrow, as long as head; sulcus shallow V shaped; eyes conical; pro- thorax as wide as long; clothed with closely set green scales; body compact, punctures prominent and deep on elytra; keel on declivity less prominent. Size 4.5-5.2 mm in length-3.0-3.4 mm in breadth. Guadalcanal Island. Figs. 30-31 becki n. sp. 9. Eyes round, large, convex; rostrum wider than long; scrobes large, hollow, sulcus deep, straight, prothorax with large lateral punctures; keel extending from middle of elytra to declivity; body robust. Size 7.1-9.2 mm in length-3..5-4.2 mm in breadth. Guadalcanal Island. Figs. 32-.34 platessa Hllr. 10. Eyes round, placed well down on side of head; antennae scape slender, scrobes open posteriorly; carina with prominent elevation at rostral declivity, de- clivity scaleless; prothorax surface granular, with a fine black medium carina; Weevil of the Tribe Celeuthetini 23 elytra with small anterior elevation on intervals 2-3; and on intervals 4-5 at declivity. Derm black with small, round, brown scales. Size 4.(S-8.0 mm in length-2.5-3.1 mm in breadth. Florida and Guadalcanal Islands. Fig. 35. granulosa n. sp. — Eye round, small, slightly pointed; rostrum wider than long, scrobes prominent, sulcus crescent-like; scape slender; prothorax with distinctive scale pattern; elytra with deep punctures, small species. Size 4.0-4.3 mm in length-2. 0-2.3 mm in breadth. Isabel, Florida, and Guadalcanal Islands. Fig. 36 isabellae n. sp 11. Prothorax much wider than long; median carina extending the full length of pro- thorax, elytra emarginate at apex; interval 9 enlarged and thickly covered by gray scales and short setae; size 8.5-8.6 mm in length-3. 1—3.3 mm in breadth. Bougainville Island. Fig. 37 . carinithorax Hllr. — Prothorax wider than long; median carina obscure; elytra less emarginate at apex, apical enlargement of interval 9 more pronounced; scales green with distinc- tive pattern; size 7.th active. The divide between the two nests was 125 feet higher than A-.3b and 225 feet high- er than A-4. A third nest active in 1968 (A-5) completed a triangle whose sides measured: A-.3b to A-4, 0.7 miles; A-4 to A-5, 1.5 miles; and A-5 to A-3b, 1.5 miles. The area of this triangle was 0.5 square miles. Home range- Only in area A was an attempt made to measure home range. The direction of all eagles going to or from their nests was re- corded and only those nests with four or more sightings appear in Fig. 4. Recordings for both years were included when the same nest or known alternate nests were involved. The birds of nest A-15a and b and A-8a and b had their activity rather evenly distributed in four directions. Those of nest A-15a and b had 33.3 percent of their activity to both the north and south and 16.6 percent to both the east and the west; while A-8a and b had 18.2 percent to the east and the west, .36 percent to the south, and 27 percent to the north. Pairs A-5, A-4, A-3, and A-2a and b had their activity somewhat restricted to various direc- tions. The birds of nest A-2 had no sightings to the north while 37.5 percent were to both south and east, with 25 percent to the west. Pairs A-.3, A-4, and A-5 had their activity pri- marily in opposite directions Fifty-four percent of the activity of A-.3a and b was to the north, while 45.5 percent and 31.8 percent of the ac- tivity of A-5 was to the east and south respec- tively. All of the activity of pair A-4 was evenly divided between both the south and west. Similarly, pairs A-9, A-10, and A-11 had the greater part of their activity in directions op- posite to each other. Sixty-one percent of the activity of pair A-10 was to the south, while pair A-11 had .50 percent to the north and .33.3 per- cent to the east. Pair A-9 had 66.6 percent to the south and .33.3 percent to the cast and no reported activity to the north and west. Alternate nests. Many of the pairs of eagles had one or more alternate nests Eleven of the 21 p;iirs considered had known alternate nests. Of these, 8 pairs had three nests and 3 pairs had two. Three of the 11 pairs with alternates are known to have used different nests the same year. The distance from active to alternate nests raneed from slightly less than 25 meters at A-4 to 1.3 miles at A-8. Nesting AcrrviT^' Nest preparation. Preparation of the nests varied greatly, with several being almost doubled in size prior to nestine, while others remained virtually unchanged. Nest A-10 was increased in height nearly 0.4 meter for the Biological Series, Vol. 10, No. 4. Ecology of Raptorial Birds o c 11 S o bC \- \/^- 10 BniGH.\M Young Univer.sitv Science Bulletin 1968 .season; in contrast, nest A-5 was enlarged veiy little during the same period. Of all the nests studied for the two years, 54.5 percent had an increase in height of 0.1 meter or more, while the remaining 45.5 percent had less than 0.1 meter of new material added. Egg Im/ing. Egg laying dates varied from year to year as well as within a given year. The first egg was noted in nest A-3b on Febru- ary 25, in A-10 on February 26, and in A-14 on March 1, 196S. Two eggs were seen in nests B-5 and B-6 on the second of March, 1968, and in nest A-15 on the fourth of March, 1968. This is in contrast to 1967 when, based on a 42-day incubation period, the first eggs were laid on March 6 at nest A-10 and March 9 at Nest .'\-14. Two eggs were found in nest A-3a on March 11, 1967, but unfortunately no laying date could be detennined, as the nest was de- stroyed before hatching. The last two eggs laid in 1968 were found on April 11 at nest B-5b, and were abandoned about May 16, with evidence that incubation had lasted nearly 35 days. This is believed to be the second nesting effort for this pair, their first having ended with two abandoned eggs about March 17, 1968 at nest B-5a. Another late nesting date occurred at nest A-7 where one young hatched about May 5. Based on a 42-day incubation period, the egg would have been laid about March 24. Temperature data from the two years indi- cate no appreciable differences and provide no obvious correlation between the laying dates for both years Incubation periods. Fairly accurate incuba- tion periods were obtained from three nests. Nest A- 15b had an egg present on March 4 and the young eaglet was cstiinated to have hatched between April 12 and 14. This would indicate a minimum of 39 to 41 days of incubation. However, it is believed that this egg was laid March 2 when the bird was observed for 35 minutes to be sitting on the nest. This would increase the incubation period to a probable 41 to 43 days. Similarly, nests A-14 and B-6 held two eggs each on March 1 and 3, respectively. The estimated dates of hatching are April 13 to 15 for the eggs of nest A-14 and April 14 to 16 for the eggs of nest B-6. This results in a mini- mum 41- to 43-day incubation period for both clutches. Fledging. The length of time required for the young to fledge varied from 8.5 weeks to 10 weeks. Pair A-8b hatched two birds about April 7, shortly after which one died Fifty-nine days later, on June 5, the nest was empty, and what is believed to be the remaining bird was seen 1.5 miles east of the nest resting on a low ledge. It allowed our vehicle to approach within 100 meters, and then flew poorly for about 0.25 mile, and landed roughly on a hillside. One young at nest A-15b fledged in approxi- mately ten weeks in 1968, and two young fledged in 9 weeks in 1967. Three other pairs (A-5, A-12, and A-14) each fledged one bird in nine weeks in 1968, and pair A-4 fledged two birds in this same period in 1968 Productivity In 1967 and 1968, 23 nesting pairs of Golden Eagles produced eggs for an average of 1.91 eggs per nest. Three pairs produced one egg each, 19 produced two eggs each, and the re- maining pair produced three eggs (Table 3). A total of 31 active pairs in 1967 and 1968 hatched 35 birds for an average of 1.13 young per pair. Twenty-six or 74 percent of these subsequently fledged for an average of 0.84 fledglings per pair for both years (Table 4). Unsuccessftd nests- Six of the 13 nesting at- tempts that failed are believed to have been interfered with by humans. Nest A-3a and b were destroyed during incubation in both years of study. In 1967 the female was shot off the nest by a small caliber bullet and both eggs destroyed, and in 1968 all three eggs were found broken in the same area below and to the side of the nest. Nests A-6, A-10, and A-11 had the eggs removed, and no remains could be found in or around the sites. Four people with a rope were seen near nest A-11 on the last day the eggs were reported present. Human interference is also blamed for the abandonment of nest A-8a in 1967 before eggs were laid. It is known that at least one man attempted to approacli this nest with a rope on the last day it was reported active. One nest (A-2) was abandoned in 1967 after a fire of unknown origin destroved a wood rat den 12 meters to the side of the nest. Although the nest and egg were undamaged they were TABLE 3 Clutch size for various Golden Eagle nests for Areas A and B CIntrb Size N.I. I'lii" N,.. l, MUS. COMP. ZOOL- 5.C0f^--yt,rov;oT LIB,RARY ^ AUG 1 8 1970 Brigham Young UniversWy Science Bulletin HARVARD UISUVERSITYi OSTEOLOGICAL AND MYLOGICAL COMPARISONS OF THE HEAD AND THORAX REGIONS OF CNEM/DOPHORUS TIGRIS SEPTENTRIONALIS BURGER AND AMEIVA UNDULATA PARVA BARBOUR AND NOBLE (FAMILY TEIIDAE) by Don Lowell Fisher '^ and Wilmer W. Tanner ^ BIOLOGICAL SERIES — VOLUME XI, NUMBER 1 JUNE 1970 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Stanley L. Welsh, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: Stanley L. Welsh, Chairman of the Board Veknon J. Tipton, Zoology Febbon L. Anderson, Zoology J. V. Beck, Microbiology Joseph R. Muedock, Botany Wilmer W. Tanner, Zoology Ex officio Members: A. Les' Sci Ernest L. Olson, Chairman, University Pubhcations A. Lester Allen, Dean, College of Biological and Agricultural Sciences The Brigham Yoimg University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from University Press, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for library exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 84601. Brigham Young University Science Bulletin OSTEOLOGICAL AND MYLOGICAL COMPARISONS OF THE HEAD AND THORAX REGIONS OF CNEMIDOPHORUS TIGRIS SEPTENTRIONALIS BURGER AND AMEIVA UNDULATA PARVA BARBOUR AND NOBLE (FAMILY TEIIDAE) by Don Lowell Fisher and Wilmer W. Tanner BIOLOGICAL SERIES — VOLUME XI, NUMBER 1 JUNE 1970 TABLE OF CONTENTS Page INTRODUCTION 1 METHODS AND MATERIALS 2 OSTEOLOGY 3 General Description of the Sia, A. it. hartwegi, and A. u. sinistra. It is either absent or questionably present in C. t. tigris, C. t. septentrionalis, C. t. gracilis, C. t. caniis, C. t. aethiops, C. gularis, C. exsanguis, C. d. deppei, C. d. lineatissisimus, C. h. hyperythrus, C. o. lemniscatus, and A. auberi. It is evident from the above distribution that the presence or absence of the interme- dium may not be of paleotelic significance at the generic level, but may be at the specific level. It may also be intraspecifically variable. 7. The musculature. The greatest myological variation appears to be centered around the anterior segment of the ventral musculature. The mylohyoideus com- plex interdigitates frequently with the geniohy- oideus in A. u. parva. Nine separate bundles were found in the species studied. Camp (1923) stated that above the family level in suarians, eight or more bundles was a valid indicator of primitiveness. The other muscles of generic var- iation include the m. cervicomandibularis, m. constructor colli, m. mandibulohyoideus II, m. omphyoideus, m. styloglossus and the associ- ated basal tongue sheath. The anatomical differences existing in the sheath associated with the tongue of^. u. parva is of special significance. A basal sheath is connected to the tongue sheath and extends for attachment to cranial and mandibular bones. This establishes a real anatom- ical basis for the tongue sheath character now used by some to separate these genera. Further exploration of species in these genera should be made to determine the stability of the character. This is particularly true in view of the comments made by Burt (1931b) con- cerning this structure, which, based on his findings, may be an inadequate character to separate these gen- era. It is evident from the findings that specialization has occurred not only intergenerically but intragener- ically, and care must be taken to distinguish the two levels of variation. Clearly the presence or absence of the intermedium is of little significance intergener- ically, but may be of importance intragenerically. The presence or absence of the pterygoid teeth may also fit this category, but with our present understanding of its paleotelic significance within this family it is impractical to draw a conclusion. Barbour and Noble (1915) and Burt (1931b) concluded from their study of external morphology that Ameiva is a more primi- tive genus than Cnemid phonis. Our observations of osteological and myological structures which are seemingly of paleotelic significance also support this conclusion. It is to be noted that the terminology of this muscle has not been determined in the literature. The name of a similarly placed mammalian muscle is temporarily adopted until complete homologies can be determined. 39 LITERATURE CITED Adams, W. E. 1953. The carotid arch in lizards with partic- ular reference to the origin of the internal carotid artery. Journal of Morphology, 92: 1 15-155. Avery, D. F. and W. W. Tanner. 1964. The osteology and myology of the head and thorax regions of the obesus group of the genus Sauromalus Dumeril (Iguanidae). Brigham Young University Science Bulletin, Biological series, 5(31:1-30. Barbour, T. and G. K. Noble. 1915. A revision of the lizards of the genus Ameiva. Cambridge Mass. Bulletin of the Museum of Comparative Zoology, 59(6):417-479. Beargie, K. and C. J. McCoy, 1964. Variation and relation- ships of the teiid lizard Cnemidophonis an^isticeps. Copeia, 1964 (3):561-570. Beebe, W. 1945. Field notes on the lizards of Kartobo Brit- ish Guinea and Caripito Venezuela. Ill Teiidae. Zoolog- ica. New York, 30:7-31. Boulenger, G. A. 1884. Synopsis of the families of e.xisting lacertilia. Annals and Magazine of Natural History, Ser. 5, 14(80);1 17-122. (reprint of the Ohio Herpetological Society, 1966:86-112.) . 1891. Notes on the osteology oi Helodenna honi- dum and H. suspectuin. with remarks on the systematic position of Helodermatidae and on the vertebrae of the lacertilia. Proceedings of the Zoological Society of Lon- don, 1891:109-118. 1899. Description of a new lizard of the genus Ameiva from Equador. Proceedings of the Zoological Society of London, 1899:517. Bradley, O. C. 1903. The muscles of mastication and the movements of the skull in lacertilia. Biologische Jahr- bucher-abtheilung fur Anatomic und Ontogenie der Tiere, 18:475-488. Brattstrom, B. H. 1954. Amphibians and reptiles from Gypsum Cave. Nevada. Bulletin Southern California Academy of Sciences, 53(1):8-12. Broom, R. 1903. On the development of the pterygoquad- rate arch in lacertilia. Journal of Anatomy and Physio- logic Normal and Pathological, 17(2): 107-1 1 1. . 1925. On the origin of lizards. Proceedings of the Zoological Society of London, 1925(1): 1-16. . 1935. On the structure of the temporal region in liz- ard skulls. Annals of the Transvaal Museum, 18:13-22. Burger, W. L. 1950. New, revived, and reallocated names of the North American whiptailed lizards, genus Cnemi- dophonis. Chicago Academy of Science Natural History Misc., ^¥65: 1-9. Burt.C. E. 1929. The genus of teiid lizards, Kerr/fanfl Cope, 1869, considered as a synonym of Cnemidophonis V^^g- ler, 1830, with a key to the primitive genera of the teiidae. Proceedings of the Biological Society of Washing- ton, 42: 153-156. — . 1931a. The status of the spotted race-runner, fnem- idophoms sexlineatus gularis (Baird and Girard). Pro- ceedings of the Biological Society of Washington, 44:73-78. — . 1931b. A study of the teiid lizards of the genus Cnemidophonis with special reference to their phylo- genetic relationships. Bulletin of the United States National Museum, 154:1-280. . 1931c. South American lizards in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural History, New York, 61:227-395. Camp, C. L. 1923. Classification of the lizards. Bulletin of the American Museum of Natural History. New York, 48(11):289-481. Cope, E. D. 1892a. On the homologies of the posterior cra- nial arches in the reptilia. Transactions of the American Philosophical Society, 27:1 1-26. 1892b. A synopsis of the species of the teiid genus Cnemidophonis. Transactions of the American Philo- sophical Society, 27:27-52. Davis, D. D. 1936. The terminology of reptilian muscula- ture. Herpetologica, 1:12-17. Dubois, E. P. 1942. Technique for sectioning small skulls. Copeia, 1942(l):55-56. . 1943. Osteology of the skull of Cnemidophorus. American Midland Naturahst, 30(2):5 10-517. Duellman, W. E. and J. Wellman. 1960. A systematic study of the lizards of the deppei group (genus Cnemidoph- onis) in Mexico and Guatemala. Misc. Publications of the Museum of Zoology, University of Michigan, 111: 1-80 Duellman, W. E. and R. G. Zweifel, 1962. A synopsis of the lizards of the sexlineatus group (genus Cnemidophonis). Bulletin of the American museum of Natural History, 123(3):158-210. Edgeworth, F. H. 1931. On the muscles used in shutting and opening the mouth. Proceedings of the Zoological Soci- ety of London, 1931:817-818. Edmund, A. G. 1960. Tooth replacement phenomena in the lower vertebrates. Royal Ontario Museum, Life Sciences Division, Toronto, Canada, 5 2 : 1 - 1 90. Etheridge, R. 1960. Additional notes on the Cragin Quarry fauna. Papers of the Michigan Academy of Science, Arts, and Letters, 45:113-117. Gadow, H. 1906. A contribution to the study of evolution based upon the Mexican species o{ Cnemidophonis. Pro- ceedings of the Zoological Societv of London, 1906(11:277-375. Gehlbach, F. R. 1965. Amphibians and reptiles from the pliocene and pleistocene of North America; A chronolog- ical summary and selected bibliography. Texas Journal of Science, 17(l):56-70. George, J, C. 1948. The muscular system of Uromastix hardwickii (Gray). Journal of the University of Bombay, 17(3): 1-23. Gnanamuthu. C. P. 1937. Comparative study of the hyoid and tongue of some typical genera of reptiles. Proceed- ings of the Zoological Society of London, Ser. B., 107:1-63. Gray, J. E. 1825. A synopsis of the genera of reptiles and amphibia, with a description of some new species. Annals of Philosophy, Set. 2, 10:193-217. (Reprint by the Ohio Herpetological Society, 1966) Hofer, H. 1960. Vergeleichende Untersuchungen am Schadel von Titpinambis und Varanus mit besonderer Beruchsichtigung ihrer Kinetik. Gegenbaurs Jahrbuch, 100:706-746. 40 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Howell, A. B. 1936. Morphogenesis of the shoulder architec- ture part [V. Reptilia. Quarterly Review of Biology, ll(2):183-208. Howes, G. B. 1902. The morphological method and prog- ress. Nature, 66:522-530. Huntington, G. S. 1903. Present problems of myological research and the significance and classification of muscu- lar variations. American Journal of Anatomy, 2(2):157-175. Jollie, M. T. 1960. The head skeleton of the lizard. Acta Zoologica, 41:1-64. Kesteven, H. L. 1919. The pterygoids in amphibia and rep- tiles and the parasphenoid. Journal of Anatomy, 53:223-238. Kingsley, J. S. 1905. The bones of the reptilian lower jaw. The American Naturalist, 39:59-64. Licht, P. 1956. Reproduction in lizards: Influence of tem- perature on photoperiodism in testicular recrudescence. Science, 154:1668-1670. Lowe, C. H. and S. R. Goldberg. 1966. Variation in the cir- cumtesticular leydig cell tunic of teiid lizards (Cnemi- dophorus and Ameiva). Journal of Morphology, 119(3):277-282. Maslin, T. P. 1959a. The nature of amphibian and reptilian species. Journal of the Arizona Academy of Science, 1(1):8-17. — . 1959b. An annotated check list of the amphibians and reptiles of Colorado. University of Colorado Studies, Biology, 6:1-98. . 1961. All-female species of the lizard genus Cnemi- dophorus, Teiidae. Science, 135(3499):212-213. — . 1963. Notes on a collection of herpetozoa from the Yucatan peninsula of Mexico. University of Colorado Studies, Biology, 9:1-20. . 1966. The sex hatchlingsof five apparently unisexual species of whiptail lizards (Cnemidophorus. Teiidae). The American Midland Naturalist, 76(2):369-378. Maslin, T. P., R. G. Beidleman, and C. H. Lowe, Jr. 1958. The status of the lizards Cnemidophorus perplexus Baird and Girard (Teiidae). Proceedings of the U. S. National Museum, 108( 3406):331-345. McCoy, C. J. 1966. Geographic variation in ovarian cycles and clutch size in Cnemidophorus tigris (teiidae). Science, 154:1671. Meyer, F. A. A. 1795. Syn. Rept. 27-Rept. Nopcsa, F. 1928. The genera of reptiles. Paleobiologica, 1(1):163-188. Oelrich, T. M. 1956. The anatomy of the head of Cteno- saura pectinata (Iguanidae). Misc. Publications of the Museum of Zoology, University of Michigan, 94:1-1 18, figs. 1-57. Patten, B. M. 1951. The embryology of the chick. Blakiston and McGray-Hill, New York. p. 85. Pennock, L. A. 1965. Triploidy in parthenogenetic species of the teiid lizard, genus Cnemidophorus. Science, 149(3683):539-540. Peters, J. A. 1964. The lizard genus Ameiva in Equador. Bulletin of the Southern California Academy of Sciences, 63:113-127. Poglayen-Neuwall, I. 1954. Die Kiefermuskalature der Ei- dechsen und ihre Innervation, Zeitschrift fur Wissen- schaftliche Zoologie, 158:79-132. Reese, A. M. 1923. The osteology of the Tegu Tupinambis nigropunctatus. Journal of Morphology, Philadelphia, 38:1-17. Robison, G. W. and W. W. Tanner. 1962. A comparative study of the species of the genus Crotaphytus Holbrook (Iguanidae). Brigham Young University Science Bulletin, Biological Series, 2(1): 1-31. Romer, A. S. 1924. Pectoral limb musculature and shoulder girdle structures in fish and tetrapods. Anatomical Rec- ord, 27(2):1 19-143. . 1956. Osteology of the reptiles. University of Chicago, Chicago, Illinois, p. 1-772. — . 1964. The vertebrate body. Saunders, Philadelphia. p. 1-569. 1966. Vertebrate paleontology. University of Chicago, Chicago, Illinois, p. 1-396. Sinitsin. D. T. 1928. The types of skull in the family Teiidae. Zoologische Anzeiger, 76:232-236. Smith, H. M. 1960. Evolution of chordate structure. Holt, Rinehart and Winston, New York. p. 1-492. Smith, H. M. and W. L. Burger. 1949. The identity of Ameiva tesselata Say. Bulletin of the Chicago Academy of Science, 8(13) :277-284. Smith, H. M. and L. E. Laufe. 1946. A summary of the Mexican lizards of the genus Ameiva. University of Kan- sas Science Bulletin, 3I(l):7-73. Smith, H. M. and E. H. Taylor. 1950. An annotated check- list and key to the reptiles of Mexico exclusive of the snakes. Bulletin of the U. S. National Museum, 199:170-192. Snyder, R. C. 1954. The anatomy and function of the pelvic girdle and hindlimb in lizard locomotion. The American Journal of Anatomy, 95( I): 1-45. Stebbins, R. C. 1966. A field guide to western reptiles and amphibians. Riverside, Cambridge, Mass. p. 125-133. Stokely, P. S. 1950. The occurrence of an intermedium in lizards. American Midland Naturalist, 43(11:179-182. Stuart, L. C. 1942. Comments on the undulata group of Ameiva (Sauria). Proceedings of the Biological Society of Washington, 55:143-149. Tanner, W. W. 1952. A comparative study of the throat musculature in the Plethodontidae of Mexico and Cen- tral America. University of Kansas Science Bulletin, 34:583-677. Taylor, E. H. 1940. Palatal sesomoid bones and palatal teeth in Cnemidophorus. with notes on these teeth in other saurain genera. Proc. Biol. Soc. Washington, 53:119-124. Taylor, H. L. and P. A. Medica. 1966. Natural hybridization of the bisexual teiid lizard Cnemidophorus inorantus and the unisexual Cnemidophorus perplexus in southern New Mexico. University of Colorado Studies Series in Biol- ogy, 22:1-9. Uzzell, T. M., Jr. 1959. Teiid lizards of the genus /'/ocosoma. Occ. papers of the Museum of Zoology, University of Michigan, 606:1-16. Wagler, J. 1830. I-VI:l-354. Naturlieher system der amphibien. 41 Wake. D. B. 1966. Comparative osteology and evolution of letin of the American Museum of Natural History, the lungless salamanders, family Plethodontidae. Mem- 117:60-116. 4"m04 "'' ^°""'"" ^''"'°'"'' Academy of Science. jg^, ^^^,y^^^ ^^ hybridization between two sub- species of the desert whiptail lizard, Cnemidophonts Zweifel, F. W. 1965. A handbook of Biological Illustration. ''^''- C°P^'^- 1962(4):749-766. University of Chicago, Chicago, Illinois, p. 1-131. . 1965. Variation in and distribution of the unisexual Zweifel, R. G. 1959. Variation in and distribution of lizards lizard, Cnemidophonis tesselatus. American Museum of western Mexico related to Cnemidophonis sacki. Bui- Novitates. 2235 : 1-49. INFORMATION FOR CONTRIBUTORS Contributions to the Science Bulletin should be primarily monographic in nature. For the most part only manuscripts of approximately forty or more typewritten pages will be ac- cepted. Papers will be published approximately in the order that they are received, pending avail- ability of funds. Authors must arrange for finan- cing their pubhcations, except that the Univer- sity Press and the hbrary of Brigham Yoimg University vsdll share the pubhcation cost pro- portionately to their needs for copies of the article printed. In the preparation of manuscripts, authors are requested to follow the Style Manual for Biological Journals, American Institute of Bio- logical Sciences, 2000 P Street, N.W., Washing- ton, D.C., 20006. Manuscripts must be typed on one side of the paper only, double-spaced vdth ample margins. Footnotes should be avoided. To fadhtate review by referees, send to the editor the original manuscript and one carbon copy, together vdth the illustrations. Copies of the original illustrations may be submitted, but should be of quahty equal to the originals. Illustrations should be referred to as figvires except for materials requiring inserts of special paper, which may be called plates. Illustrations should be so designed as to fit when reduced into a one-column or full-page vddth. Special care must be taken to allow for proper reduc- tion in lettering (i.e., a 50% reduction of the figure means also a 50% reduction in the lettering). Photographs should be of a glossy finish, xmblurred, and showing sharp con- trast. Line dravidngs should be made with black ink on heavy white drawdng paper, blue trac- ing cloth, or blue-ruled coordinate paper. Use the same abbreviations on line drawings as in text. Line drawings must be equivalent to a professional draftsman's work. Original dravvdngs are preferable to photographs, even if they are large. Illustrations (hne drawings or photo- graphs) should be munbered consecutively throughout the paper, and the approximate place of insertion should be indicated in the margins of the manuscript pages. Captions for illustrations should be assembled on a separate sheet, and each plate of figures must have its corresponding figure nvimber pencilled hghtly on the back. Illustrations and cuts will be de- stroyed unless their retvun is requested when proof is returned to the editor. A table title should be a short, concise state- ment of what the table piu^orts to show, and should not include information necessary to the interpretation of the table. Every column in the table should carry a head identifying the data in that column; the measure in which the data are given should be indicated at the head of each colvimn. Tables should not be used when the same information can be given in a few lines of text, and should not duplicate informa- tion in text, graphs, or charts. Symbols ( asterisk, dagger, etc.) should be used to indicate foot- notes to tables, with footnotes on the same page as the table. An abstract of less than foiu: percent of the length of tlie paper shoidd be prepared. This simimary should be understandable vdthout ref- erence to the body of the manuscript. The ab- stract must be on sheets separate from the man- uscript. Proof should be corrected immediately on receipt and returned to the editor. Authors should leave forwarding addresses if they move from the address sent with the manuscript. Reprints should be ordered when the proof is returned. Address all manuscripts to Wilmer W. Tan- ner, Department of Zoology and Entomology, Brigham Young University, Provo, Utah. -^r^ 5-/^/1- P/rovoJ Brigham Young Universifys comp zooi_. LIBRARY Science Bulletin ,^pp ^ ^ ,^,„ HARVARO UNIVERSITY PINYON-JUNIPER SUCCESSION AFTER NATURAL FIRES ON RESIDUAL SOILS OF MESA VERDE, COLORADO by < James A. Erdman BIOLOGICAL SERIES— VOLUME XI, NUMBER 2 JUNE 1970 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Stanley L. Welsh, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: Vernon J. Tipton, Zoology Ferbon L. Anderson, Zoology Joseph R. Murdock, Botany Wdlmer W. Tanner, Zoology Ex officio Members A. Les Sc Ernest L. Olson, Chairman, University Publications A. Lester Allen, Dean, College of Biological and Agricultiu-al Sciences The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from Publication Sales, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for hbrary exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 84601. n u... Frontispiece. The Morfield Burn of 1959 several days after the natural fire was suppressed. Ash and mud were temporarily impounded behind this road across East Canyon. Brigham Young University Science Bulletin PINYON-JUNIPER SUCCESSION AFTER NATURAL FIRES ON RESIDUAL SOILS OF MESA VERDE, COLORADO by James A. Erdman BIOLOGICAL SERIES— VOLUME XI, NUMBER 2 JUNE, 1970 CONTENTS Page Introduction 1 Methods 3 Results 4 The Morfield Burn of 1959 4 The Wetherill Burn of 1934 6 The Long Mesa Burn of 1873 10 The Long Mesa climax site 13 Additional evidence on climax 15 Dendrochronology: a new application 18 Discussion and conclusions 18 Pinyon-juniper succession after natural fires on residual soils of the Mesa Verde . . .18 The status of the present forest stands 21 Acknowledgments 24 Literature cited 25 ILLUSTRATIONS Figure Page 1 . Map of the eastern part of Mesa Verde with locations of the study sites 2 2. Aspect of the Morfield Burn in July 1963 6 3. Aspect of the Wetherill Burn in July 1963 8 4. Soil profile underlying the Wetherill Burn near the macroplot sample 9 5. Dating the Long Mesa Burn; skeleton plot of the wedge of a burned Douglas-fir cross-dated with the Mesa Verde master chronology 11 6. Dating the Long Mesa Burn: part of the wedge of a burned Douglas-fir from which the skeleton plot shown in figure 5 was made 11 7. Dating the Long Mesa Burn: increment core of Douglas-fir cross-dated with the Mesa Verde master chronology • • ■ • 1 1 8. Aspect of the Long Mesa Burn in July 1963 12 9. Soil profile underlying the Long Mesa Burn 13 10. Aspect of the Long Mesa climax stand 14 1 1 . Soil profile underlying the Long Mesa climax stand 15 12. View of the pinyon-juniper/mountain brush element that characterizes the higher parts of Mesa Verde in the Ute Mountain Indian Reservation \(, 13. A pinyon pine growing in the burned-out base of a juniper on Big Mesa 18 14. Suggested successional stages after fire on residual soil of Mesa Verde 18 15. Climographs comparing temperature and precipitation in Mesa Verde (Park Point) and at Pagosa Springs, Colorado 23 TABLES Page 1. Summary of Mesa Verde climate 3 2. Cover and frequency of the vegetation in the Morfield Burn sample 5 3. Cover and frequency of the vegetation in the Wetherill Burn sample 7 4. Description of the soil profile underlying the Wetherill Burn near the macroplot sample 9 5. Outside ring dates of pinyon pines killed in the Long Mesa Burn of 1873 10 6. Dates of pinyon-juniper establishment following the Long Mesa Burn of 1873 . . 10 7. Cover and frequency of the vegetation in the Long Mesa Burn sample 13 8. Cover and frequency of the vegetation in the Long Mesa climax forest sample ..15 9. Point quarter analysis of the pinyon-juniper element in the Long Mesa climax forest 16 10. Tree density analysis of the Long Mesa climax forest 16 1 1 . Description of the soil profile underlying the Long Mesa climax forest 17 12. Point quarter analysis of the pinyon-juniper element in the Ute Mountain Indian Reservation climax forest 17 13. Species listing by community according to successional status 19 14. Comparison of soil moisture between tree canopies and adjacent openings, Chapin Mesa 20 15. Ages of stands surveyed subsequent to the timber cruises of Spencer (1964) .... 21 16. Comparison of the Mesa Verde climatic data with similar data characteristic of the interior ponderosa pine forests of the Southwest 23 17. Comparison of temperature and precipitation at Mesa Verde (Park Point) and at Pagosa Springs, Colorado 23 PINYON-JUNIPER SUCCESSION AFTER NATURAL FIRES ON RESIDUAL SOILS OF MESA VERDE, COLORADO by James A. Erdman* ABSTRACT A mountain brush vegetation typified by Gambel oak (Quercus gambelii) and serviceberry (Amelan- chier ittahensis) dominates the uplands of Mesa Verde National Park. There is evidence that this brush ele- ment is a successional stage that has been maintained by repeated natural fires in a large part of the Mesa Verde landscape. This study was conducted to deter- mine the role of fire in the region's ecology, the nature of the climax pinyon pine (Piiius edulis) and Utah juniper (Junipenis osteosperma) vegetation, and the major successional stages that lead to the climax condition. Three postburn plant communities and two climax stands overlying residual soils at elevations of about 7,500 feet were studied. The successional stands had been burned through natural causes in 1873, 1934, and l^S^. The earliest date was determined by cross- dating trees affected by the fire with the Mesa Verde master chronology, since the fire antedated any writ- ten records of the region. Pith dates from pinyon pines in the climax forest stands indicated an age of about four centuries. Permanent sample plots representative of each stand were established, and the vegetation was ana- lyzed using a modification of the line intercept and macro plot methods. The vascular species that occurred in each sample were noted, and their cover value and frequency were summarized. Plant succession proceeds rapidly from a pioneer stage of the shade and competition intolerant weeds. Helianthus annuus and Chenopodium pratericola, to a meadow stage dominated by the native grasses, Ory- zopsis hymenoides, Sitanion hystrLx. and Poa fendler- iana. In the most recent burn, several species of per- ennial exotic grasses have been introduced to check erosion. After about 25 years, a crown-sprouting brush ele- ment, consisting primarily oi Quercus gambelii, Ame- lanchier utahensis, Cercocarpus montanus, and Pur- sfiia tridentata becomes the dominant vegetation. At this time, seedlings of Pimis edulis and Juniperus osteosperma are also established. This open shrub stage gradually becomes more dense, and in about 100 years forms a thicket stage. Young trees come up through the shrubs, eventually overtopping them. As the fire sere proceeds toward a climax condition, the brush species are gradually suppressed by the matur- ing forest. After several centuries, the understory is composed mainly of a sparse shrub component, a grass (Poa feudleriaiia), prickly-pear (Opuntia poly- acantha), and several forbs. The data tend to support the theory that recurrent fires throughout previous centuries have permitted a chaparral-like, floristically rich shrub vegetation to persist as a fire climax along the uplands of Mesa Verde National Park. Under the present management policies of fire suppression, however, a pinyon-juniper forest is slowly replacing the former extensive shrub vegetation. INTRODUCTION Mesa Verde, a highly eroded plateau in southwest- ern Colorado (Fig. 1), is on the eastern edge of the Colorado Plateau physiographic province. A pinyon- juniper forest dominates most of the mesa, but dense brush vegetation is common at higher elevations, especially within the boundaries of Mesa Verde National Park. The ecological status of the brush has been a puzzle because, while it appears relatively stable, other sites on the mesa with an apparently similar environment are occupied by stands of pinyon-juniper. Moreover, evidences of fire predating the recent human occupation of the area suggest that the brush may be a product of fire (Anderson, 1961). As a member of the staff of the park's Wetherill Mesa Project (Osborne, 1964), I was charged with helping to determine the environmental conditions that existed during the prehistoric Indian occupation of the area, which extended up to the close of the thirteenth century. It was apparent that an under- standing of succession following fires would be help- ful in the paleoenvironment researches. A fire in Morfield Canyon, in July 1959, provided me with an opportunity to study early stages in the sere, and shortly thereafter I began an intensive investigation of "Present address; U.S. Geological Survey, Denver, Colorado. BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Fig. 1. Map of the eastern part of Mesa Verde. Residual soils 7,500 feet. The study sites are as follows: 1) Morfield Burn, Climax. An additional climax site that was studied is located the Wetherill Burn. the mountain brush, or chaparral (Bader, MS.; Ander- son, 1961). The Mesa Verde sandstones and shales of Upper Cretaceous age (Wanek, 1959) rise 1,500 feet above the valleys to the north which reach a maximum ele- vation of 8,500 feet. These beds dip southward towards the Mancos River. Here the canyons attain depths of 1,000 feet. Travel across the mesa, there- fore, is generally along the upper reaches, where the relief consists of rolling swales and ridges. MESA VERDE NATIONAL PARK AND VICINITY MONTEZUMA COUNTY COLORADO cover the upper third of the mesa, at elevations generally above 2) Wetherill Burn. 3) Long Mesa Burn, and 4) the Long Mesa on the Ute Mountain Indian Reservation about 2 miles west of Residual soils are developed from the underlying sandstone on the narrow, more easily eroded ridges of the higher parts of Mesa Verde— the area under con- sideration. Most of the soils on the remainder of the plateau, however, appear to be eolian in origin (Arr- henius and Bonatti, 1965), with accumulation contin- uing even today though at a minimal rate. The cal- cium carbonate content is higlily variable; in some places it forms a thick caliche layer, while in others free lime is absent. I BIOLOGICAL SERIES. VOL. 11, NO. 2 PINYON JUNIPER SUCCESSION AFTER NATURAL FIRES The climate of Mesa Verde is mild, for in this semi- arid region fair weather prevails throughout the year. Temperatures on the mesa (Table!) are suprisingly moderate, considering the altitude. This is a result of both generally continuous winds from the southwest and lack of cold-air drainage. Extreme temperature fluctuations, rare on the mesa tops, are pronounced along the canyon floors, however (Erdman, Douglas, and Marr, 1969). TABLE 1 SUMMARY OF MESA VERDE CLIMATE, U.S. WEATHER BUREAU STATION AT HEADQUARTERS, CHAPIN MESA. ELEVATION 7,070 FEET. RECORD PERIOD, 1923-1963* Montli Air Temperature = F Precipitation Maximum Minimum Mean in inches January 62 -20 29.8 1 .93 February 68 -15 33.0 1.95 March 72 4 38.6 1.76 April 84 9 48.1 1.32 May 90 23 57.0 1.05 June 101 32 67.4 .70 July 102 43 72.9 1.68 August 101 41 70.8 2.01 September 94 28 64.4 1.53 October 85 13 52.7 1.66 November 75 -3 39.5 .98 December 67 -6 32.1 1.71 Annual 102 -20 50.5 18.28 *U.S. Weather Bureau, 1964, and Mesa Verde National Park weather records. Precipitation is irregular. A proportionately large amount comes as snow during the late winter months. Normal snowfall is 78 inches. The typically dry months of May and June are followed by a period of intense and often highly localized thunderstorms. Rainfall during the late summer and early autumn may be heavy, but because of rapid runoff and higli evaporation it is much less effective than winter mois- ture in recharging ground water supplies. Except for the Mancos River, which skirts the mesa on the south, , the only natural water supplies available to wildlife are occasional springs, seeps, and potholes that have been filled by rains or melted snow. Pollen studies by Martin and Byers (1965) and simultaneous dendroclimatic investigations by Fritts, Smith, and Stokes (1965) indicate that the climate has been relatively stable in the Mesa Verde region during the past millennium. This period would seem to be adequate for plant species adapted to the cli- matic conditions here to establish themselves and achieve stable communities. Mesa Verde lies within the pinyon-juniper climax region, an area of 60 million acres lying mostly in Nevada, Utah, Colorado, Arizona, New Mexico, and western Texas (Arnold, Jameson, and Reid, 1964). These forests have been described by Howell (1941), Woodbury (1947), Merkle (1952), and Woodin and Lindsey(1954). I have not used Merriam's terms. Upper Sonoran and Transition zones, since I agree with Johnsen (1964) that "The terms Upper and Lower Sonoran are occasionally used in generalized, limited area studies by nonbotanists to avoid detailed descriptions of flora and fauna; and, as such, the terms have a gen- eral but limited descriptive value." Most of the plateau is mantled with stands of pin- yon pine (Pinus ediilis Engelm.) and Utah juniper (Juniperus osteospenna (Torr.) Little). The areas underlain by loess have an understory dominated by muttongrass (Poa fendleriana (Steud.) Vasey), and those underlain by residual soils have an understory dominated by shrubs. Dominance, as used here, is based upon "an approximation of the area over which a plant exerts its influence upon other components of the ecosystem" (Daubenmire, 1959). Elements of the montane tlora are common at the more mesic sites in the Mesa Verde area. These sites are dominated by aspen {Popiilus tremuloides Michx.), ponderosa pine {Pinus ponderosa Dougl.), and Dougl as- fir ( Pseud otsiiga menziesii (Mirb.) Franco), or a combination of all three. As an exten- sion of the Great Basin flora to the west, big sage- brush {Artemisia tridentata Nutt.) assumes a climax status on the deep, friable, alluvial terraces in the can- yons. The flora of Mesa Verde is characterized by Madro-Tertiary species (Weber, 1965). In addition to the pinyon-juniper element, the mountain-brush thickets so extensive along the North Rim contain several species belonging to this assemblage, the prin- cipal one of which is Gambel oak (Quercus gambelii Nutt.). Some of the Madro-Tertiary representatives, however, are exceedingly rare relicts in the Mesa Verde. Among these are single-leaf ash (Fraxinus anomala Torr. ex S. Wats.), big-toothed maple {Acer grandidentatum Nutt. ex 1. & G.), and Ajo oak {Quercus ajoensis Muller), a recent addition to the Colorado flora (Erdman, Weber, and Tucker, 1962). This study is concerned with the mountain-brush element associated with the pinyon-juniper forest on residual soils. METHODS I have modeled this study on Viereck's research (1966), which followed the methods of Jenny (1941), Major (1951), and Crocker (1952). Jenny proposed that soil is the result of five independent variables-climate, organisms, topography, parent BRIGHAM YOUNG UNIVERSITY SCIENCE BULLEIIN material, and time-all acting together to produce a characteristic type of soil. He emphasized the value of finding examples in nature where all the variables but one are relatively constant. In his terminology, for example, a chronosequence develops with all factors except time held constant. Major (1951) pointed out that these variables are just as valuable in vegetational studies, and illustrated a series of vegetation sequences in which all variables except one were kept constant. Crocker (1952) suggested that the ecosystem would be a more logical unit to use than either soil or vegetation, for it is the ecosystem that is the function of the five independent variables. The present study is based on this concept. In addition, 1 have employed the principle of succession, which states that given an adequate period of time and barring disturbance, unstable ecosystems with the same climate, potential organisms, relief, and parent material will converge towards a single climax type. Through reconnaissance studies and inspection of aerial photos, a selection was made of several young postburn stands (i.e., concrete ecosystems of Marr, 1961) of different ages and of two much older, rela- tively stable ecosystems. A permanent sample area, visually determined to be representative of each stand, was established, and a species list for it was compiled. The line intercept method (Canfield, 1941) and the macroplot method (Poulton and Tisdale, 1961) were used. Both methods are based on the line transect, although the latter adds small rectangular quadrats, called observation plots, for estimating her- baceous cover. Ten parallel, 50-foot lines, each subdivided into 1 0-foot units, were laid out in each sample area at 25-foot intervals along a base line. Tree and shrub canopies projected onto these lines provided a meas- ure of the relative importance or dominance of each species in the sample. According to Lindsey (1956), this canopy coverage is "the most important single parameter of a species in its community relations." Earlier studies of the pinyon-juniper type have also used the intercept technique (Cable, 1957; Jameson, Williams, and Wilton, 1962; and Arnold, Jameson, and Reid, 1964). Frequency, expressed as the per- centage of subsamples in which a species occurs, was used to measure the relative distribution throughout the stand. Understory was studied within lx2-foot observa- tion plots placed at 10-foot intervals along each line. Cover within these quadrats was estimated by using the Domin scale (Cain and Castro, 1959). The abundance and reproduction of pinyon pine and juniper were compared by counting those individ- uals lound within ten 4x50-foot belt transects adja- cent to each line. The following size categories were used: seedlings, less than 1 foot in height; saplings. stem diameters less than 2 inches; and trees, greater than 2 inches in diameter. In the climax-forest samples, trees were selected for importance values and associated data, using the point quarter method of Cottam and Curtis (1956). Ten points were located at 50-foot intervals. Tree-ring samples were dated according to the Douglass system (Clock, 1937). Increment cores pro- vided ages of the recovering forest, while cross sec- tions from charred snags were collected from the Long Mesa Burn to establish the date of that fire. Robert F. Nichols, archeologist and dendrochronolo- gist with the Wetherill Mesa Project, helped collect samples from both the Long Mesa Burn and the Ute area climax forest. David G. Smith, dendrochronolo- gist-climatologist formerly with the Laboratory of Tree-Ring Research, provided most of the dates. Dating mature junipers was not attempted because this species often has an extremely erratic growth and iobing habit, and shows frequent doubling of rings. Up-to-date discussions of the principles and appli- cations of dendrochronology have been published by Bannister (1963), McGinnies (1963), and Fritts (1965a). A good treatment of "signature groups" and key years that occur in the Mesa Verde chronology is to be found in an article by Spencer (1964). Soil profiles were described and additional infor- mation was provided by Orville Parsons, of the Soil Conservation Service. Where possible, photographs of these profiles were made by Fred Mang, Jr. RESULTS The Morfield Burn of 1959: Pioneer Weeds and Meadow Serai Stages Between July 15 and August 7, 1959, a lightning- sparked fire burned 2,043 acres extending over sec- tions of both Mesa Verde National Park and Ute Res- ervation lands of the east end of Mesa Verde. The flanks of the burn generally paralleled Morfield Can- yon on the west and Whites Mesa to the east (Fig. 1). Razing of the vegetation was complete (see frontis- piece). After spending several days on a fire line at Whites Mesa, the fire crew was relieved from duty, and as we drove through the moonlit pall of ash and smoke in Waters Canyon the tree snags glowed like a myriad of campfires. Some of the burned area had not been completely forested prior to the fire. Much of the eastern part of the park is covered with dense brush and scattered young pinyon and juniper trees. This vegetation has developed since the mid-1800's, when a large fire swept over the terrain. The western part of the Morfield Burn had been a pinyon-juniper forest with trees up to 30 feet in height. On an isolated ridge of this portion of the burn (Fig. 1) a permanent study BIOLOGICAL SERIES. VOL. 11, NO. 2 PINYON JUNIPER SUCCESSION AFTER NATURAL FIRES plot was established to monitor the vegetation change. Vegetation. According to park files, a mixture of commercial seed was air-dropped over most of the burned area in early October 1959, to check erosion. Equal amounts of intermediate wheatgrass {Agropyron intemie- diiim), crested wheatgrass (A. desertonim), and smooth brome (Bromus inennis) were applied at the rate of 6 pounds per acre. The small ridge in Figure 1 was not seeded intentionally so that natural succes- sion could be studied. But, because of seed drift, the exotic grasses soon became well established in the sample plot also. In order to promote tree recovery, 30 pounds of pinyon seeds were planted in a 1-acre plot on Whites Mesa. However, germination and growth have been poor. The vegetation was leveled by the fire except for the snags of the mature trees. A layer of ash was evi- dent over most of the landscape. But by autumn, many of the fire-adapted shrubs were sprouting from root crowns, and these were developing vigorously the following year. Gambel oak had crown-sprouted and was in leaf within a few months after the fire- impressive evidence of its adaptability to fire. The pioneer vegetation during the first two years of recovery was dominated by two annuals, sunflower (Helianthiis anmius) and pigweed {Chenopodium pratericola). Other advent ives found at the site in 1960 were wheat (Tritiaim aestivum) and the brome grasses, Bromus carinatiis and B. japonicus. Further evidence suggests that the Helianthus- Chenopodium community-type constitutes an early serai stage throughout Mesa Verde. In the summer of 1960, a small plot on Chapin Mesa was intentionally burned in a fire-school demonstration. Sunflower and pigweed appeared the following year. By 1962, the third year after the Morfield burn, the introduced and native grasses began supplanting the pioneer annuals. In August 1963 the grass com- munity in the study plot was analyzed for cover and frequency, using the modified line intercept-macro- plot method (Table 2). The grasses by then were well established, with scattered clumps of resprouted shrubs assuming a minor role (Fig. 2). The composite, Machaemnthem bigelovii, and Russian thistle (Salsola kali) were the most abundant forbs, but the original pioneer annuals were evident only as litter. Of the herbs listed in Table 2, only lupine (Lupinus cauda- tiis) and Penstemon liiuirioides are components of the climax forest. The following perennial herbs, also common in the climax forest, were observed in the study area by 1963 but did not occur in the sample plot: Poa fendleriana. Sitaiiion hystrix. Yucca bac- cata, Petradoria piimila. Opiintia polyacantha, Com- andra iimbellata, Arabis selbyi. and Eriogonum race- mosum. A summary of all the species occurring in the stands sampled is given in Table 13. TABLE 2 COVER AND FREQUENCY OF THE VEGETATION IN THE MORFIELD BURN SAMPLE ELEVATION 7,550 FEET. AUGUST, 1963 Species % Cover % Frequency* Shrubs Amelanchier utahensis** -)-*** 2 Fendlera rupicola + 2 Qiierciis gambelii + 2 Herbs Machaeranthera bigelovii <4 74 Agropyron desertonim <4 60 Salsola kali <4 46 Bromus tectontm <4 42 Agropyron intermedium <4 34 Viguiera multiflora <4 18 Bromus inermis <4 10 Oryzopsis hymenoides <4 6 Descwainia pinnata + 44 Lactuca scariola + 14 Lappula redowskii + 8 Lupinus caudatus + 8 Chenopodium fremontii + 6 Penstemon linarioides + 6 Sphaeralcea coccinea + 6 Camelina microcarpa -1- 4 Moldavica paiviflora + 2 Ground soil 25-33 92 litter 4-10 100 rock <4 46 *Based on 50 subplots. **Species authorities are given in the synoptical list ofTable 13. ***Indicates presence but no significant cover value. Soils. The soils on the study ridge are higlily variable. Parsons considered the soil just north of the sample plot to be Witt loam developing on a thin mantle of loess sediments over Cliffliouse sandstone. This pocket of relatively deep soil (one auger hole reached 60 inches to bedrock) is more like the eolian soils common at lower elevations to the south, where the mesa tops are broad and subject to less erosion. The shallower soil south of the plot is Montvale stony fine sandy loam. No description was made of the profile, but the following data were obtained from auger holes at the corners of the macroplot: southeast corner 16 inches to bedrock, B^ at 6 inches, + test for CaCO-, at 12 inches depthfsouth- west corner 12 inches to bedrock, rock slabs at 8 inches, CaCO^ test -; northwest corner 17 inches to bedrock, slight + CaCOj test at bottom, B2 among BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN rock fragments; northeast corner 12 inches to bed- rock, CaCOj test + at bottom. Rock outcrops occur along the edges of the ridge and in parts of the sample plot. In general, the soil within the macroplot is prob- ably less than two feet in depth. Some CaCOj occurs at the bedrock-soil interface. The Wetherill Burn of 1934; the Mountain Brush Serai Stage Biesen (1931) prefaced his description of the coni- fers of Mesa Verde with these observations: While the trees of the park completely cover the mesa areas, they do not grow densely together. . . For this reason as well as for the fact that usually heavy downpours accompany each severe elec- trical storm, the fire hazard of the park is not considered to be very great. It is interesting to note that Mesa Verde has not been troubled with severe fires in recent years. During the present season Fig, 2. Aspect of the Morfieid Burn in July 1963. The vegetation at this stage consists of exotic and native grasses, some forbs, and scattered fire-adapted shrubs, such as this small clone of Gambel oak in the foreground. Soils on this ridge are variable. Elevation, 7,500 feet. I BIOLOGICAL SERIES. VOL. 11. NO. 2 PIN YON JUNIPER SUCCESSION AFTER NATURAL FIRES only two small fires have been reported, and they were quickly extinguished by the Park's efficient protective organiza- tion. This was the feeling that prevailed in the park prior to 1934. Fire was never considered a serious threat to the mantle of pinyon-juniper, often referred to as "the asbestos forest." That view soon changed when the largest fire in park history taxed the efforts of over 1,000 men attempting to control it. Don Watson (1934), then ranger-historian of the park, gave this account of the event: On the ninth of July a small cloud, wandering across the Mesa Verde, caused an electrical storm that sent a few streaks of lightning down into our forests. One of these lightning streaks, striking a tree, started a fire that was to prove the most disastrous Mesa Verde has ever known. For two weeks it raged, ultimately con- suming 4,540 acres of timber and brush lands. The Wetherill Burn was in reality the result of two fires that tlared up consecutively, one on the Ute Mountain Indian Reservation on July 9, and the other in the park on July 11, 1934. During a flare-up on July 14, they merged into a fire that was not declared out until July 26. This burn, too, left a desolate land- scape in its wake. The following incident provides further evidence of the fire potential of lightning. An extremely abbre- viated electrical storm that resulted in a "smoke" occurred one summer afternoon in 1964 while I was visiting Long House, a cliff dwelling on Wetherill Mesa. The weather was fair with no sign of shower activity. Yet while I was in the recesses of the large rock shelter I heard what later proved to have been a lightning bolt strike a large juniper tree about 1 mile up the mesa. The fire was quickly suppressed before it spread, but this event underscores the major role of fire in the dynamics of the regional vegetation prior to modern fire-fighting technology. Contrary to the statement of Biesen (1931), elec- trical storms are a consequential factor in determining the nature of the Mesa Verde vegetation. The Wether- ill Burn, and, more recently, the Morfield fire were liglitning-caused. Vegetation. As with the Morfield Burn, attempts to reforest the area artificially have met with but limited success. There were 42,000 trees planted on Wetherill Mesa in 1942 (unpublished park records). Several years ear- lier, the Civilian Conservation Corps planted 200,000 seedlings. Only a few isolated stands of these pinyon pines and Rocky Mountain junipers (Jiinipems scopu- lonim Sarg.) persist today, in most cases as rows across shallow washes off the main ridges. The present open brush vegetation was sampled in 1963 in an area heavily forested before the 1934 fire (Fig. 3). A macroplot was established on the main ridge of Wetherill Mesa at an elevation of 7,650 feet (Fig. 1). The data given in Table 3 are for only those plants intercepted by the sample lines or included in TABLE 3 COVER AND FREQUENCY OF THE VEGETATION IN THE WETHERILL BURN SAMPLE ELEVATION 7,650 FEET. AUGUST, 1963 Species 7' Cover % Frequency* Trees Piiuis echilis +** 2 Shrubs Amelanchier iitahensis 10 20 Yucca baccata 6 16 Chrysolhamnus nauseosus 2 10 Cercocarpus montanus 1 6 Fendlera nipicola I 6 Svmphoricarpos oreophilits + 4 Herbs Poa fendleriana 4-10 88 Sitanion hystrix <4 24 Sphaeralcea coccinea <4 22 Heterotheca villosa <4 18 Bromus tectonim + 22 Penstemon linarioides + 16 Schoenocrambe linifoliwn -1- 16 Lappula redowskii + 10 Viguiera multiflora ■1- 8 Polygonum racemosum + 6 Zygadenus elegam + 6 Petradoria pumila + 6 Oiyzopsis liymenoides -1- 6 Moldavica pamflora -1- 4 Comandra umbellata + 4 Machaeranthera bigelovii + 4 Stipa comata + 2 Lupinus caudatus -1- 2 Calochortus nuttalUi + 2 Achillea lanulosa + 2 Arabis selbyi + 2 Ground soil 25-33 88 litter 4-10 96 rock <4 20 *Based on 50 subplots. **Indicates presence but no significant cover value. 5 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Fig. 3. Aspect of the Wetherill Burn in July 1963. Dead snags of the former pinyon-juniper forest show little weathering. The mountain-brush species, released from competition with the trees, are becoming more dominant. Elevation, 7,650 feet. the observation plots. Gambel oak, for example, was not sampled but was abundant along the edges of the ridge. Bitterbrush (Piirshia tridentata), too, was com- mon in the burn but not in the immediate area. A list- ing of those species occurring in the macroplot area, however, is given in Table 13. The stand was characterized by serviceberry (Ame- lanchier utahemis). Yucca baccata, Poa fendleriana, and Heterotheca villosa. One pinyon pine sapling occurred in the macroplot and was included in the line intercept. Since the pith date by ring count was 1953, this tree was not part of the earlier planting program. Such a delay in natural establishment of the tree component is consistent with the postfire recov- ery on Mesa Verde. Regardless of the extent of the area burned, 20 to 30 years usually lapse before the pinyon-juniper element begins to return. The few nonvascular plants encountered are, for the most part, the moss Grimmia, found on rocks scattered over the ridge. A small complex of lichens. principally CoUema tenax, covers parts of the soil sur- face. Soil. The soil profile (Fig. 4 and Table 4) was studied at a pit located a short distance down the ridge from the macroplot. According to Parsons (personal communi- cation), the soil representative of this site is tenta- tively identified as Roubideau very fine sandy loam. It has developed primarily from weathered products of the underlying sandstone, although eolian soils have modified it to some extent. Soil depth around the macroplot, however, is shallower. Holes augered at the corners of the sample revealed the following depths to bedrock: southeast corner, 14 inches; southwest corner, 10 inches; northwest corner, 6 inches; and northeast corner, 6 inches. Tests for CaCOj were negative. Several additional test holes inside the macroplot ranged from 20 to 43 inches in depth. One of these had 10 inches of caliche (CaCO-,) in the lower part of the profile. BIOLOGICAL SERIES. VOL. U, NO. 2 PINYON JUNIPER SUCCESSION AFTER NATURAL FIRES V\^' Fig. 4. Soil profile underlying the Wetherill Burn near the macroplot sample. The soil in this pit typifies the Roubideau very fine sandy loam. Note the deep, highly structured Bq horizon and the absence of CaCOq, TABLE 4 DESCRIPTION OF THE SOIL PROFILE UNDERLYING THE WETHERILL BURN NEAR THE MACROPLOT SAMPLE Horizon Depth (inches) Description 'l\ 0-4 Brown (7.5YR 5/2, dry) to dark brown (7.5YR 3/2, moist) very fine sandy loam; moderate to strong very fine granular structure; consist- ence soft dry, very friable moist; noncalcareous; weakly platy structure in the upper inch or two; lower boundary clear and smooth. 4-7 Reddish-brown (5.0YR 5/3, dry) to reddish-brown (5.0YR 4/3, moist) very fine sandy loam; weak medium structure breaking to weak to TABLE 4 (Continued) moderate medium subangular blocks; consistence slightly hard dry, and very friable moist; many fine pores, numerous fine worm casts; noncalcareous; lower bound- ary clear and smooth. Bj 7-12 Reddish-brown (5.0YR 5/4, dry) to dark reddish-brown (5.0YR 3/4, moist) heavy loam; moderate medium prismatic structure breaking to moderate medium subangular blocks; consistence hard dry, and very friable moist; a few very thin and patchy clay skins on soil peds, noncalcareous; lower boundary clear and smooth B-,, 12-19 Reddish-brown (5.0YR 5/4, dry) to dark reddish-brown (5.0YR 3/4, moist) clay loam; moderate medium pris- matic structure breaking to moderate to strong medium angular blocks; consistence very hard dry to firm moist; many thin patchy clay skins on ped surfaces with some slickenside surfaces; a few weathered sandstone frag- ments; noncalcareous; lower boundary clear and smooth. B-n 19-32 Reddish-brown (slightly red- der than 5.0YR 5/4, dry, and 5.0YR 4/4, moist) heavy clay loam; strong medium prismat- ic structure breaking to strong medium angular blocks; consistence very hard dry, and firm moist; thin almost continuous clay skins on soil peds; slickenside numerous; noncalcareous; lower boundary clear and smooth. R 32 Relatively unweathered sand- stone with extensions of B-,-, into cracks. 10 The Long Mesa Burn of 1 873 : the Mountain Brush-Incipient Forest Serai Stage Long Mesa lies due east of Wetherill Mesa (Fig. 1). Its northern ridges are within the area of the 1934 Wetherill Burn. Much of the northern part of Long Mesa and a large segment of Long Canyon show evi- dence of a fire that occurred before the park was established. To my knowledge, this burn antedates the settling of the region by early ranchers. A site was selected on a narrow ridge where the fire, originating in Long Canyon, had spread over part of the mesa top (Fig. 1). This small section of the Long Mesa Burn was bordered on the north and south by pinyon-juniper forest. Dating the Burn. Cross sections were cut from pinyon pines that had been killed by the fire. The results given in Table 5 indicate that the event must have occurred after 1870. This was not a bark date since the terminal rings had weathered away, and the date presumably represents the latest date of the outermost persisting rings. TABLE 5 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN TABLE 6 DATES FOR PINYON-JUNIPER ESTABLISHMENT FOLLOWING THE LONG MESA BURN OF 1873 Mesa Verde Species Specimen No. 3047 Utah juniper 3048 Utah juniper 3055 pinyon pine 3056 pinyon pine 3057 pinyon pine 3058 pinyon pine 3059 pinyon pine 3060 pinyon pine 3063 pinyon pine 3064 pinyon pine 3065 pinyon pine ^ Based on increment cores -nc indicates "near center" -'p indicates "pith" Inside Ring Date 7 1896±nc^ 1908 nc 1927 p3 1901 p 1912 nc 1907 nc 1907 nc 1900 p 1905 p 1900±nc 1912 nc OUTSIDE RING DATES OF PINYON PINES KILLED IN THE LONG MESA BURN OF 1873 Mesa Verde Specimen No. Outside Ring Date 3002 3003 3004 3005 3009 301 1 3015 3016 3019 3020 1870 1810± C.1846 no date 1868 no date no date no date 1869 C.I860 The most recent year in which the fire miglit have occurred is 1896. This is the pith date of the oldest living tree within the burn. This second line of evi- dence—increment cores from trees that have come in subsequent to the fire (Table 6)-provided the latest dates possible for the burn. More precision was made possible by studying two Douglas-fir trees that were caught in the upward sweep of the fire as it moved out of Long Canyon. A wedge (Mesa Verde specimen No. 3066) was cut from one of the trees destroyed by the fire. The ring series from this section was plotted and matched with the master chronology developed for Mesa Verde by den- drochronologists of the Laboratory of Tree-Ring Research (Fig. 5). The pith date was 1663; the extremely eroded outermost ring was 1871 (Fig. 6), a date that was in accord with those from the mesa-top samples. At the same time the wedge was collected, an increment core (Mesa Verde specimen No. 3067) was taken from a Douglas- fir tree that was living but had probably been affected by the fire. David Smith studied the core and arrived at a probable fire date of 1874 based on a suppression wood sequence from 1874 through 1889 (Fig. 7). Apparently no xylem increment occurred in the position from which the increment core was taken during the initial years, for only ten rings are evident in the suppression series. In light of radial growth studies reported by Doug- las and Erdman (1967) and a closer analysis of the increment core in Figure 7, the year 1873 may be a more accurate date of the Long Mesa fire. Radial growth in Douglas-fir terminates by late spring, cer- tainly by early June. Therefore, the increment laid down the year of the fire should be normal, not locally absent or suppressed; 1873 is the last normal ring before the sequence, 1874 through 1889. BIOLOGICAL SERIES. VOL. 1 1, NO. 2 FIN YON JUNIPER SUCCESSION AFTER NATURAL FIRES 11 Hd= Hii ■2il^Mt ci n li mnn nr ilil iHl mm fi^it iill Ml Fig. 5. Dating the Long Mesa Burn: Cross-dating the skeleton plot derived from the wedge of a burned Douglas-fir with the Mesa Verde master chronology. A diagnostic grouping, or "signature" occurs in the 1818-24 sequence. Other key years are 1847, 1851, and 1861. The annual increments for 1847 and 1861 are missing from the specimen (indicated by the letter m). The extremely eroded outermost ring of 1871 is indicated by the symbol vv. "^K^.-'^^vX^v'^VxV,' -^ 1850 1871vv Fig. 6. Dating the Lons Mcs.i Bum: I'.nl .'I tin- wedge of a burned Douglas-fir (Mesa Verde specimi-ii No. 3066) from which the skeleton plot shown in Figure 5 was made. The cross-dating provided an outermost ring date of 1871. The single pinholes indicate decades; the double pinhole, the 1850 increment. Vegetation. The present vegetation in the Long Mesa Burn is a dense entanglement of shrubs and young trees (Fig. 8). The dead, charred snags of the former forest are generally screened by this new vegetation. The pinyon-juniper component of the stand has been increasing since the turn of the century, when the first seedlings were established (Table 6). Although the older trees are 15 to 20 feet in height, the mountain brush species continue to characterize the stand (Table 7). The shrubs with the maximum cover and greatest frequency were bitterbrush {Purshia tridentata) and Fendlera nipicola, both almost tree-like in their growth habits here, in con- trast to their small size in the climax forest. *f - Fig. 7. Dating the Long Mesa Burn; Increment core of Douglas-fir cross-dated with the Mesa Verde master chronology. The length of each vertical bar on the graph is inversely proportional to the relative width of the rmg; average width rings are not recorded. The suppression wood between the years 1874 and 1889 is atypical and reflects the effect of the fire. Micro-rings occurred in 1847, 1851, 1861, 1902, and 1904. 12 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN I Fig. 8. Aspect of tin I, out; Mesa Burn in July 1963. The mountain brush species dominate the stand although the cone-bear- ing pinyon-juniper canopy is overtopping them. The charred snag in the center was a mature Utah juniper. Elevation, 7,400 feet. Most of the herbs sampled were perennials, the only exceptions being Polygonum sawatcheiise, found in the pinyon-juniper forest throughout Mesa Verde, the widespread cheatgrass (Bronius tectonim), and the almost equally common v/eed,Lappula redowskii. Soil. The soi: underlying the burn is identified as a shai-' low phase of Montvale fine sandy loam. It is very stony and higiily calcareous, as is evident in Figure 9. Eolian sediments are possibly a minor influence in this solum, developing residually from weathered Cliffliouse sandstone. Plant growth may be enhanced by the underlying fractured sandstone, which serves as a reservoir for additional soil moisture, and by some lateral flow along the bedrock surface. BIOLOGICAL SERIES. VOL. 1 1 , NO. 2 PIN YON JUNIPER SUCCESSION AFTER NATURAL FIRES TABLE 7 COVER AND FREQUENCY OF THE VEGETATION IN THE LONG MESA BURN SAMPLE ELEVATION 7,400 FEET. SEPTEMBER, 1963 13 Species % Cover % Frequency* Trees Pinus eduUs 19 46 Jiinipents osteosperma 6 12 Shrubs Piirshia tridentata 26 82 Fendlera nipkola 14 48 Yucca baccata 13 40 Cercocarpos montamis 1 10 Quercus gambelii + ** 4 Amelanchier utahensis + 2 Herbs Poa fendleriana <4 56 Lappula redowskii <4 32 Oryzopsis hymenoides <4 16 Opiintia polyacantha <4 6 Eriugomtm umbeUatum <4 6 Bromits tectonim + 22 Penstemon linarioides + 16 Sitanion hystrix + 8 Senecio multilobatus + 6 Boil teloua gracilis + 4 Cryptantha bakeri + 4 Polygonum sawatchense + 4 Artemisia liidoviciana + 2 Schoeiiocrambe linifoliuin r + 2 Gutierrezia sarothrae + 2 Sphaeralcea coccinea + 2 Petradoria pumila + 2 Penstemon strictis + 2 Ground litter 25-33 100 soil 10-25 46 rock <4 36 *Based on 50 subplots. **Indicates presence but no significant cover value. This profile is similar to the profile described in the climax forest nearby (Table 11), although the former is four inches less in total depth and has a slightly coarser-textured B2 horizon. The Long Mesa Climax Site; the Pinyon- Juniper/Mountain Brush Stage One mile north of the Long Mesa Burn lies a small stand of pinyon-juniper that was missed by both the Fig. 9. SoU profile underlying the Long Mesa Burn. This soil is transitional between Montvale fine sandy loam and shal- lower series. The highly structured, blocky B2 is developing between fragments of Cliffhouse sandstone which overly a 1873 and 1934 fires (Fig. 1). This stand was studied for two reasons. First, as a remnant of the forest that was destroyed in the 1873 burn, it served as a con- trol. Secondly, and of more significance, it is con- sidered to be representative of the type of vegetation that can develop on relatively shallow soils in the Mesa Verde only after centuries of little disturb- ance~the climax forest-type under the present cli- matic regime. Vegetation. This 400-year-old stand consists of but a few char- acteristic species. Those apparent in Figure 10 are Yucca baccata, Poa fendleriana, the prickly-pear, Opuntia polyacantha, and some scraggy examples of the mountain brush element. On the basis of the macroplot analysis (Table 8), the pinyon pine and Utah juniper are clearly codomi- nants, with similar cover values and frequencies. But from the results given in Table 9, juniper is more dominant than pinyon pine, and is of considerably more importance in the stand. These differences are due to differences in the criterion used to determine dominance. The macroplot method stresses cover from the canopy; the point quarter method summar- ized in Table 9 emphasizes board feet. Cover was 14 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN based on canopy projection irrespective of tree size; the point quarter study excluded trees less than four inches in diameter. Pinyon pine had a relatively small trunk compared with juniper, averaging seven inches in diameter. Trunks of juniper averaged 20 inches. The density transects reported in Table 10 indicate that pinyon pine was more numerous when seedlings and saplings are considered. Thus, although juniper was the aspect dominant, pinyon pine was more abundant. Both species attained crown heights of 35 feet, about the maximum reported for other parts of the Mesa Verde. >5^^ Fis. 10. Aspect of the Long Mesa clima.x stand in July 1963. The sparse understory consists mostly of Poa fendleriana. Yucca baccara, and generally suppressed shrub species such as the mountain mahogany in the foreground. Elevation, 7,600 feet. BIOLOGICAL SERIES. VOL. 1 1, NO. 2 PINYON JUNIPER SUCCESSION AFTER NATURAL FIRES 15 The age of the stand was based upon increment cores taken from the older trees in the stand. The old- est pith ring date, 1576, came from a 21-inch diame- ter pinyon. The considerably greater diameters of jun- iper trunks, however, suggest that this species may be the older of the tree components. Unfortunately, mature juniper trees can rarely be dated. TABLE 8 COVER AND FREQUENCY OF THE VEGETATION IN THE LONG MESA CLIMAX FOREST ELEVATION 7,600 FEET. AUGUST, 1963 Species % Cover % F requency* Trees Jiinipents osteosperma 31 50 Piniis edulis 27 48 Shrubs Cercocarpos inoiuaims 14 28 Yucca baccala T 14 Symphoricarpus oreophiliis 2 12 Fendlera mpicola 2 8 Amelanchier ittahensis +** 2 Purshia tridentata -H 2 Herbs Poa fendleriana 4-10 80 Peiistemon linarioides <4 24 Opuntia polyacantha <4 10 Comandra umbellata -H 14 Eriogonum umbellatiim -1- 8 Petradoria pumila -1- 8 Polygonum sawatchense -H 6 Schoenocrambe linifolium -1- 6 Lupinus ammophilus -H 4 Oryzopsis hymenoides + 4 Lappula redowskii + 4 Bronius tectonim + 4 Sitanion liystrix ■h 2 Pedicularis centranthera -^ 2 Cryptantha bakeri + 2 Chenopodium fremontii -1- 2 Balsamonliiza sagittata -1- 2 Ground litter 33-50 94 soil 4-10 48 rock <4 20 *Based on 50 subplots. **Indicates presence but no significant cover value. Fig. 11. Soil profile underlying the Long Mesa climax stand. The Montvale fine sandy loam has a finer-textured B horizon than is typical, but otherwise it is representative. Soil. This soil, too, has been tentatively identified as Montvale fine sandy loam. It has a finer-textured B horizon than is typical of most of the Montvale soils mapped in the area by Parsons (MS), but it is other- wise representative. The CaC03 component is less extensive than that in the burn sample. The profile (Fig. 1 1) is described in Table 1 1. Additional Evidence on Climax: Ute Pinyon-Juniper/Mountain Brush Forest An answer to the problem of determining climax vegetation is especially difficult in the park proper because prehistoric burns cover the entire northern third of its area. On the other hand, the Ute Indian holdings of the Mesa Verde to the west remain rela- tively unaffected by fire. For whatever reason, a mature pinyon-juniper forest covers that section of the mesa. The mountain brush or chaparral "zone" so familiar in the park is not evident on the Reservation. In 1963, a reconnaissance flight over Mesa Verde prompted a brief excursion into the reservation forest. On July 1 1 , 1964, a small field party ascended the North Rim in the vicinity of the Mesa Verde Mine, about four miles west of Wetherill Mesa. A 16 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN TABLE 9 POINT QUARTER ANALYSIS OF THE PINYON-JUNIPER ELEMENT IN THE LONG MESA CLIMAX FOREST* Species Number Mean Basal of Distance Area Trees (feet) (inches-) Relative Importance Density ^^^^^ p^.^^^ j^^^^^^ y^j^^ No./Acre -req. (%) (%) (%) (D+F+Do) Juniperus osteosperma 24 Pinus edulis 16 7,309 155 60 53 92 205 647 103 40 47 8 95 all trees 40 13 7,956 258 100 100 100 300 *Sampling method included only those trees greater than four inches in diameter. TABLE 10 TREE DENSITY ANALYSIS OF THE LONG MESA CLIMAX FOREST BASED ON TEN 4x50-FOOT BELT TRANSECTS Species Seedlings* Saplings**Trees*** Total Pinus edulis 6 10 Juniperus osteosperma 5 0 24 14 *Individuals less than one foot in height. **Individuals taller than one foot but with the main stem less than two inches in diameter. ***Individuals with main stems greater than two inches in diameter. pinyon-juniper stand with a mountain-brush under- story similar to the one studied on Long Mesa was selected for observation (Fig. 12). The elevation of this ridge-top site was 7,900 feet. The species composition was generally similar to that in the Long Mesa climax stand, except for some herbs along the North Rim in the park. The tree stratum was very similar to that on Long Mesa, although the Ute stand tended to be more open (Table 12). Maximum height was 35 feet and the approximate age was 400 years. One of the largest pinyon pines observed in the Mesa Verde was cored at this site; the diameter was 32 inches, and the date of the center ring was 1539. Several other pinyons dated around 1600. A Utah juniper, 20 feet tall and 15 inches in diameter was also cored. The center ring Fig. 12. Detail of the pinyon-juniper/mountain bnjsh element that characterizes the higher parts of the Mesa Verde in the Ute Mountain Indian Reservation. Elevation at this ridge-top site is 7,900 feet. date was 1570, nearly as old as the pinyon of twice the size. The largest juniper in the point quarter anal- ysis was 36 inches in diameter. These data, along with similar results obtained elsewhere, suggest that the juniper component in much of the forested sections of the Mesa Verde is considerably older than the pin- yon pine codominant. BIOLOGICAL SERIES. VOL. 1 1, NO. 2 PIN YON JUNIPER SUCCESSION AFTER NATURAL FIRES TABLE 1 1 Table 1 1 (Continued) 17 DESCRIPTION OF THE SOIL PROFILE UNDERLYING THE LONG MESA CLIMAX FOREST Horizon Deptii (inches) Description 0-4 Brown (7.5YR 5/3, dry) to dark brown (7. SYR 3/3, moist) fine sandy loam; moderate to strong very fine granular structure; con- sistence soft dry, and very friable moist; considerable dark organic flakes and very many fine almost matted roots; non- calcareous; lower boundary clear and smooth. 4-7 Brown (slightly redder than 7.5YR 5/4, dry) to dark brown (slightly redder than 7.5YR 3/4, moist) loam; weak medium sub- angular structure; consistence slightly hard dry, and very fri- able moist; very many fine roots and pores; noncalcareous; lower boundary clear and smooth. 7-10 Brown (slightly redder than 7.5YR 5/4, di7) to dark brown (slightly redder than 7.5YR 3/4, moist) heavy loam; weak medium prismatic structure breaking to moderate medium angular blocky structure; con- sistence very hard dry, and very friable moist; thin patchy clay skins; many fine roots. B3c R numerous cicada casts, 5 percent weathered sandstone chips; non-calcareous; lower boundary clear and smooth. 10-14 Reddish-brown (5.0YR 5/4, dry) to reddish-brown (5.0YR 3/4, moist) clay loam; moderate medium prismatic structure breaking to strong medium and fine angular blocky structure; consistence very hard dry, and firm to slightly friable moist; almost continuous clay skins with some slickensides; 25 per- cent weathered sandchips; non- calcareous; lower boundary clear and smooth. 14-16 Brown (7. SYR 5/4. dry) to dark brown (7. SYR 4/4, moist) heavy loam or light clay loam; weak medium subangular blocky structure; consistence hard dry, and friable moist; white chalky lime concretions, 25 percent sandstone fragments; very thin patchy clay skins; strongly cal- careous; lower boundary clear and smooth with large roots fol- lowing along underlying sand- stone surface. 16 Relatively indurated sandstone. TABLE 12 POINT QUARTER ANALYSIS OF THE PINYON-JUNIPER ELEMENT IN THE UTE MOUNTAIN INDIAN RESERVATION CLIMAX FOREST* Species Number of Trees all trees 40 Mean )istanc- (feet) Basal Area Density (inches-^) No./Acre Relative Dens. Freq. Dom. (%) (%) (%) Juniperus osteosperma 27 Pimis ediilis 1 3 22 11,131 90 100 100 100 Importance Value (D-HF+Do) 9,020 61 68 56 81 205 - 2,111 29 32 44 19 95 300 "Sampling method included only those trees greater than four inches in diameter. 18 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Dendrochronology; A New Application The counting of rings from actively growing trees has been used for years to determine the age of forest stands and even some geomorphological events. For instance, ring counts have provided some information on the effects of fire and other disturbances on the pinyon-juniper type in Arizona (Arnold, Jameson, and Reid, 1964). This simple dating technique is not usable, however, in dating fires because we do not know how soon living trees become established after burns (Fig. 13). On the other hand, the method of cross-dating specimens directly affected by a fire with a master chronology for the region can provide rather accurate dating information. Fig. 13. A pinyon pine began growing in the burned-out base of a juniper about 1880, and thereby established a baseline from which to work back in time to determine the date of the fire. This site is on Big Mesa in the eastern part of the Mesa Verde. An 18-inch increment borer provides a scale. In the Mesa Verde, for example, the northern ridge of Moccasin Mesa is covered with a scattering of young pinyon-junipers, most of which were estab- lished around the turn of the present century. All the charred specimens collected for dating purposes, how- ever, placed the burn in the 18th century. Perhaps recurrent flash fires prevented the return of pinyon- junipers for almost a century. In any event, this infor- mation would not have been available from simple ring counts. In summary, where master chronologies are avail- able or can be built, cross-dating trees destroyed or otherwise affected by fire provides reliable data con- cerning the time of the disturbance. This technique might well be applied to other ecological work of this sort. DISCUSSION AND CONCLUSIONS Pinyon-Juniper Succession after Fires on Residual Soils of Mesa Verde In Mesa Verde sites where dense stands of pin- yon-juniper on shallow, residual soils have been burned, recovery begins with the establishment of annuals, which reach a maximum density in two to three years. Helianthus and Chenopodium are the initial dominant pioneers, followed by Russian thistle (Salsola kali) and Machaeranthera bigelovii. By the fourth year perennial grasses dominate the burn. Under natural conditions, these consist of Indian rice- grass (Oryzopsis hymenoides), muttongrass {Poa fen- dleriana). and squirreltail (Sitanion hystrix). The grass and forb stage eventually gives way to a persistent mountain-brush vegetation type. This postfire sere is generally consistent with the one reported by Arnold, Jameson, and Reid (1964) in Arizona. When fires recurred periodically in prehistoric times, a chaparral fire climax persisted along the upper reaches of Mesa Verde. But under the present protective regime, a young, open stand of trees is gradually suppressing the shrub element, and even- tually a mature forest may again be established. These successional stages following a fire are given in Figure 14. . b I m A4 yv pinyon-juniper forest \ FIRE skeleton forest & bare soil i annual stage perennial grass/forb stage shrub stage shrub/open tree stage i CLIMAX pinyon-juniper forest in years O 2 4 25 lOO 300 ♦ Fig. 14. Suggested successional stages after fire on residual soils of the Mesa Verde. Details of species change in the postfire sere, sum- marized in Table 13, lead to the following con- clusions. First, in the change toward a climax status there is a definite trend towards impoverishment in the fioristic composition. In the young Morfield Burn community there were 47 species, compared to 40 in the older Wetherill Burn, 28 in the Long Mesa Burn, and 26 in the climax forest. This trend is due pri- marily to the reduction of weedy annuals and bien- nials. BIOLOGICAL SERIES. VOL. II. NO. 2 PINYON JUNIPER SUCCESSION AFTER NATURAL FIRES 19 TABLE 13 SPECIES LISTING BY COMMUNITY ACCORDING TO SUCCESSIONAL STATUS* Species* 1959 Burn 1934 Burn 1873 Burn Climax Forest Trees Pinus edulis Engelm. Juniperus osteosperma (Torr.) Little Shrubs Chrysotbaminis nauseosiis (Pall.) Britt. Quercus gambelii Nutt. Amelanchier utahensis Koehne Cercocarpiis montanus Raf. Fendlera rupicola A. Gray Yucca baccata Torr. Purshia tridentata (Pursh) DC. Symphoricarpos oreophUus A. Gray Grasses Agropyron desertorum (Fisch.) Schult. Agropyron intermedium (Host.) Beauv. Bromus cariiiatus Hook. & Arn. Bromus iucrmis Leyss. Bromus japonicus Thunb. Triticum aeslivum L. Bromus tectorum L. Oryzopsis hymenoides (R. & S.) Ricker Poa fendieriana (Steud.) Vasey Sitanion hystrix (Nutt.) J. G. Smith Koeleria gracilis Pers. Stipa comala Trin. & Rupr. Bouteloua gracilis (H.B.K.) Lag. Sporobolus cryplandrus (Torr.) A. Gray Subshrubs and Forbs Helianlbus anmms L. Chenopodium pratericola Rydb. Salsola kali L. Camelina microcarpa Andrz. Descurainia pinnata (Walt.) Britt. Descuraiiiia sopliia (L.) Webb, ex E. & P. Melilotus officinalis (L.) Lamb Arabis selbyi Rydb. Astragalus scopulontm T. C. Porter Lithospermum ruderale Dougl. Cirsium cf. undulatum (Nutt.) Sprcng. Lactuca scariola L. Machaeranthera bigelovii (Gray) Greene Moldavica parviflora (Nutt.) Britt. Tragopogon dubius Scop. Viguiera multiflora (Nutt.) Blake Erigeron flagellaris A. Gray Eriogonum racemosum Nutt. Heterotheca villosa ( Pursh) Shinners Lupinus caudatus Kell. Zygadenus paniculatus (Nutt.) Wats. Sphaeralcea coccinea (Pursh) Rydb. Lappula redowskii (Hornem.) Greene Polygonum sawatchense Small Schoenocrainbe linifolium (Nutt.) Greene Balsamorrhiza sagittata (Pursh) Nutt. Comandra umbellata (L.) Nutt. Opuntia polyacantha Haw Penstemon linarioides A. Gray Petradoria pumila (Nutt.) Greene Senecio multilobatus T. & G. e.\ A. Gray Senecio mutabilis Greene Artemisia ludoviciana Nutt. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + 20 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN TABLE 13 (Continued) Gutierrezia sarothrae (Pursh) B. & R. Crepis sp. Cymopleris bulhosus A. Nels. Penstemon strictis Benth. ex DC. Cryptantha bakeri (Greene) Payson Eriogonum umbellatum Torr. Lupimis ammophilus Greene Pedicularis centranthera A. Gray ex Torr. *Within groups pioneer plants are listed first while those more restricted to the climax forest are last. **Nomenclature follows that of Welsh and Erdman (1964); more recent nomenclatural changes were provided by William A. Weber, University of Colorado Museum. Inasmuch as several of the grasses listed in Table 13 are adventives, the permanency of those artifi- cially seeded will require further study. Poa fen- dlehana, on the other hand, is the most characteristic grass in the Mesa Verde area and in much of the pin- yon-juniper climax region of the Southwest. All the shrubs listed, except rabbitbrush (Chry- sothammis nauseosus), are common in the pinyon-ju- niper/mountain-brush forests of the Southwest; and probably all of these, except rabbitbrush, reproduce in varying degrees by crown-sprouting. Resprouting of the bitterbrush (Purshia tridentata) was not ob- served, although it is known to sprout or stem layer in California (Nord, 1959). Snowberry (Symphori- carpos oreophilus) may propogate similarly, for it occurs in the fairly recent burn at Wetherill Mesa. Another species of snowberry, S. rivulahs, crown-sprouts along coastal California (Wells, 1962). The rapid recovery of the subshrubs after the Mor- field fire suggests that they, too, must resprout. Crown-sprouting shrubs have a decided advantage over Utah juniper and pinyon pine during at least the first 100 years of succession because reestablishment of these trees depends entirely on seed production. The chief agents for dispersal of juniper seed are animals, as reported in a scat analysis by Arnold, Jameson, and Reid (1964). Cheek pouches of a chip- munk trapped in Mesa Verde were full of such seeds (C. L. Douglas, personal communication). The means by which pinyon seeds are spread are not as well known, but rodent caches are probably a major factor. Woodbury (1947) stated that Utah juniper may precede pinyon pine in succession towards climax, although I found no conclusive evidence for this oc- curring in the Mesa Verde area. In the sample on Wetherill Mesa, pinyon was the first tree in forest es- tablishment; but on Long Mesa, Utah juniper pre- ceded pine by several years (Table 6). Juniper may have a small recovery advantage in that it needs no nurse plant, whereas pinyon invariably does (Phillips, 1909; Meagher, 1943; Erdman, MS.). Shade has traditionally been considered the factor required to produce sufficiently moist conditions for pinyon pine germination. But data collected in 1964 (Table 14) indicate that tree canopies may intercept a considerable amount of precipitation, with the result that soil moisture regimes under the litter beneath the trees are poorer than are those in the more insolated openings. Or, the lack of moisture under the canopies may be the result of depletion due to roots in that area. TABLE 14 COMPARISON OF SOIL MOISTURE (% DRY WEIGHT) BETWEEN TREE CANOPIES AND ADJACENT OPENINGS, CHAPIN MESA* Date Canopy Opening 1 May 1964 11.7 20.0 1 June 19f-,4 8.9 11.0 1 July 1964 7.4 6.5 1 August 1964 9.5 14.3 1 September 1964 7.9 8.1 *Based on gravimetric samples from six inches below the soil surface; about four inches of litter covered the soil under the canopy. An alternative explanation is the relatively low optimum temperature required for germination. I have found that 70°F is the optimum germinating temperature for pinyon pine seeds from Mesa Verde and this agrees with earlier studies by Kintigh (1949). Kintigh not only found that germination was most rapid and complete at this temperature, but that it was also most favorable for the growth of the seed- lings. 1 discovered that the lethal threshold occurred between 85° and 100°F. In a recent study of the regional and local environments (Erdman, Douglas, and Marr, 1969), it was found that temperatures two inches below the ground surface in unshaded sites during summer and early fall months were commonly above 100°F, albeit for only a few hours at a time. Even at 6-inch depths, temperatures exceeded 90°. ^ BIOLOGICAL SERIES. VOL. 1 1, NO. 2 PINYON JUNIPER SUCCESSION AFTER NATURAL FIRES 21 By comparison, soil temperatures under the tree canopies were found to be moderate. Such conditions are believed to be due more to the insulating effects of tree litter than to the shade afforded by the trees themselves (Oosting, 1956). The Status of the Present Forest Stands The pinyon-juniper stands on Mesa Verde are ap- proaching climax status, the condition in which no appreciable change is taking place under prevailing conditions (Hanson and Churchill, 1961). These stands have been largely free of disturbance for several centuries, yet the structure of age-classes in the pinyon population does not fit the typical struc- ture of a climax stand. The oldest age-class, 400 to 500 years (Table 15), is younger than the age this species is capable of attaining in this environment (Smiley, 1958). Furthermore, there is a dearth of dead and fallen pinyon trees. At the same time, there TABLE 15 AGES OF STANDS SURVEYED SUBSEQUENT TO THE TIMBER CRUISES OF SPENCER (1964)* Stand Location Inside Age RingDate Asof 1965 Ute climax, Mesa Verde Mine** Long Mesa climax*** Glades climax, Chapin*** Knife-Edge, North Rim*** Ml station, Chapin M2 station, Chapin*** CCC Camp, Chapin Spur, Chapin Research Laboratory, Chapin Cedar Tree Tower, Chapin Rock Springs, Wetherill Mug House, Wetherill South end, Wetherill Park-Ute boundary, Wetherill Double House*** Step House Rock Canyon, west talus Rock Canyon, east talus Navajo Canyon, west talus Navajo Canyon, east talus 1539 426 1576 389 1711 254 1581 384 1529 436 1647 318 1538 382 1561 404 1601 364 1531 434 1457 508 1606 359 1581 384 1586 379 1500 465 1669 296 1594 371 1644 321 1698 267 1723 242 *The majority of these dates are from an unpub- lished catalog of collections made the summer of 1961 by Thomas P. Harlan, Laboratory of Tree-Ring Research. **Dated by James A. Erdman. ***Dated by David G. Smith. is an abundance of old and of dead juniper trees (Erdman, MS.). I have been able to determine the age of some of the smaller juniper trees by cross-dating cores with the master tree-ring chronology for Mesa Verde. Extrapolating from these data, it is reasonable to conclude that the oldest junipers are several hun- dred years older than the most ancient pinyons. These observations suggest the possibility that some factor in the environment has affected adversely the pinyon component of an otherwise relatively stable climax forest. Several possible factors such as porcupine feeding pressure, blight, and drought are discussed below. Spencer, (1964), after making a thorough timber survey of Mesa Verde, stated that the scattered, older pinyons are from 300 to 500 years of age, indicating a relatively young pinyon element on the mesa. Dates in support of this finding are given herein (see Table 15). Spencer contended that porcupines have con- tributed to the maintenance of this condition, but the evidence he presents does not entirely bear out his conclusion. While feeding pressures were high in the canyons and along the perimeters of the mesa, the incidence of porcupine feeding on the mesa tops ranged from zero to only 7 percent. This can hardly account for the relatively young stands of pinyons extant on the mesa tops. A more serious toll on pinyon pine has been exacted by the root fungus, Leptographium, as re- ported by Mieike (MS.), and by Wagener and Mielke (1961). This disease was first discovered in Mesa Verde in 1942, but field evidence indicates the in- festation is of long standing. Losses of mature trees seem to have been heavy, but there are no indications that the fungus ever threatened to eliminate pinyon altogether. Wagener and Mielke observed; Most of the escapes [sic] are small trees with limited root systems that do not come in close contact with diseased roots. These escapes provide the nucleus for a new stand on the areas over which the disease has passed. In Mesa Verde National Park all stages of residual and replacement stands of pinyon in old disease areas can be found. Such blights also occur in many areas of the mesa beyond the park boundaries, and it may well be that these diseased stands, with a spread rate of about 40 feet each 6 years, could account for a youthful pinyon forest, especially where the incidence is highest in the deeper-soil mesa centers. Bark beetles of the genus Ips are known to invade Leptographium-iniected trees. Broods originating in such trees sometimes attack and kill neighboring pinyons, but losses from primary Ips attack are apparently low (Wagener and Mielke, 1961). Pressures by porcupine and Leptographium may have been minor compared to the effects of wide- 22 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN spread drought the Southwest experienced several centuries ago, causing considerable die-out of the less resistant species, pinyon included (Smiley, 1958). According to Malde (1964), this drought occurred about A.D. 1600, and in the following two centuries the "forests expanded again around small stands of the older trees (those which flourished between 1300 and 1500) that had survived in sheltered habitats." Malde concluded: "It is a remarkable fact that most conifers in the Southwest are not older than 350 years, but that a few are nearly twice that age." These data resulted from research done under the Navajo Land Claim Project (Smiley, 1958). Smiley suggested that with the advent of a more moderate climate after the drought, the forests spread from the "islands" of older trees that acted as seed sources. In a dendroclimatic study of western North America, Fritts (1965b) mapped regional variations in climate, based upon 10-year departures in tree-ring indices, which tend to support Smiley's theory. Of the period A.D. 1566-85, Fritts writes, "Dry con- ditions intensify in the Southwest and northern Rockies; a major drought develops until it finally ex- tends throughout the entire West." In the Mesa Verde area, a center of low growth persisted from 1571-95. The 1600's ushered in an extremely good growth regime, for, as Fritts relates for the 1601-25 period, "An extensive, extremely moist climate expands throughout the central portions of the West." And of the period between 1641 and 1650, Fritts says, "Moist conditions intensify in the Rio Grande and Upper Colorado River Basins, expand into the southern and north-central portions of the West, but do not reach their previous maximum extent." Though the maps represent only a tentative recon- struction of otherwise undocumented climatic patterns, they do provide a plausible answer to the problem of age relations in the pinyon-juniper forests of the Mesa Verde. Utah juniper, the more xerophytic species, apparently was able to withstand the drought conditions. A final problem in regard to climax at Mesa Verde involves the role of ponderosa pine. Getty (1935) stated, "Only rarely does yellow pine (Pinus pon- derosa) occur in the [prehistoric] ruins, indicating that then, as now, this tree grew only in the northern and higher parts of the Mesa Verde, remote from most of the ruins." This misconception that pon- derosa pine grows only along the upper reaches of the mesa seems quite prevalent. 1 have found pockets of this species, however, throughout the plateau in es- pecially mesic sites, from Morfield Canyon on the east to Horse Springs and beyond in the Ute Indian Reservation; from the rimrock above The Knife-Edge at the northern escarpment to an extensive stand in the Pine Canyon area overlooking the Mancos Canyon to the south. Some of these stands admittedly are no more than single fire- and lightning-scarred sentinels, looming above the surrounding brush and pinyon-ju- niper-possibly relicts of formerly more extensive stands. Yet in Bobcat Canyon on Wetherill Mesa, trees approaching 100 feet in height tower over saplings and seedlings of the same species. But in this instance the exceptional growth and probably the success of the stand are products of good ground-wa- ter supplies from local seeps and springs. Rand (1966) discussed factors related to distribution of ponderosa and pinyon pines at the Grand Canyon, Arizona. Some of the factors discussed are relevant to the Mesa Verde. There is little indication that ponderosa pine has changed appreciably during the last millenium from its present pattern of distribution. As Getty (1935) said, ponderosa pine is rarely found in the prehistoric ruins, which predate the fourteenth century. And in the Wetherill Mesa excavations, those sites yielding some ponderosa artifacts are near the present stand of this species in Bobcat Canyon. During numerous surveys along the North Rim I found many large stumps charred from undated fires; but the trees were pinyon pine and Rocky Mountain juniper {Junipenis scopiilonim), both common along the North Rim today. It is true that with elevations generally above 7,000 feet and yearly precipitation averaging 18 inches. Mesa Verde should support a better stand of ponderosa pine. But because of its physiography, temperatures over most of the plateau exceed those normally expected at such an altitude. Based on the findings of Pearson (1931) and of Curtis and Lynch (1957) Parsons (MS.) attributed the status of pon- derosa pine in Mesa Verde to an unfavorable tem- perature regime. Pearson (1950) summarized the climate of the forest interior: The geographic range of interior pon- derosa pine in the Southwest is associated with certain climatic characteristics which vary little from one locality to another. In general terms, the climate may be de- scribed as cold and dry, mean annual tem- perature varying from 42° to 48°F and growing season temperatures (June through September) from 58° to 65°. Maximum temperatures seldom exceed 95° and frost may occur in any month of the year. Precipitation ranges generally between 18 and 22 inches, though occa- sionally it may be higher or lower. A comparison of these values with data from the weather station near park headquarters (Table 16) would suggest that Parsons' contention is valid. But similar measurements from Park Point, given in the same table, cast doubt on the temperature argument, BIOLOGICAL SERIES. VOL. 11, NO. 2 PINYON JUNIPER SUCCESSION AFTER NATURAL FIRES TABLE 16 23 COMPARISON OF THE MESA VERDE CLIMATIC DATA WITH SIMILAR DATA CHARACTERISTIC OF THE INTERIOR PONDEROSA PINE FORESTS OF THE SOUTHWEST Station Mean Temperature °F Annual June-September Precipitation in inches Mesa Verde, park headquarters* Mesa Verde, Park Point** Forest Interior*** 50.5 45.4 42-48 68.9 62.5 58-65 18.28 18.84 18-22 *U.S. Weather Bureau, 1964. Data are from a Weather Bureau station at 7,070 feet elevation for the period 1923-1963. **Erdman, Douglas, and Marr, 1969. Data are from Park Point on the North Rim at 8,575 feet elevation for the period 1962-1963. ***Pearson, 1950. TABLE 17 COMPARISON OF TEMPERATURE AND PRECIPITATION AT MESA VERDE (PARK POINT)* AND AT PAGOSA SPRINGS, COLORADO** RECORD PERIOD, 1962-1963 Mean Temperature, °F Precip. in Park inches Month Park Pagosa Pagosa Point Springs Point Springs January 23.0 17.2 1.02 1.84 February 30.9 28.5 2.00 1.99 March 28.5 29.8 1.25 1.12 April 41.6 43.2 .47 .69 May 52.6 50.5 .54 .20 June 59.6 57.1 .55 .21 July 66.9 64.3 1.99 .64 August 64.1 62.6 4.79 2.70 September 59.2 57.4 1.29 2.28 October 50.7 48.5 2.82 2.56 November 37.9 35.5 1.27 1.58 December 29.8 26.1 .82 .74 Annual 45.4 43.4 18.84 16.20 *Erdman, Douglas, and Marr, 1969. Data are from a temporary weather station on the North Rim at 8,575 feet elevation. **U.S. Weather Bureau, 1963 and 1964. Data are from a standard Weather Bureau station at 7,238 feet elevation. for they fall well within the ranges of the ponderosa forest interior. A comparison of weather data (U.S. Weather Bureau 1963, 1964) from Pagosa Springs, Colorado, located in the heart of a ponderosa pine climax region 70 u. 60 o 3 o 50 • a E » 40 c o 30 c o S 20 10 — 1 1 r Summer Winter _L Pagosa Springs Mesa Verde - _L _L -L 0 1.0 2.0 3.0 4.0 5.0 Monthly Mean Precipitation, inches Fig. 15. Climographs comparing temperature and precipita- tion in the Mesa Verde (Park Point) and at Pagosa Springs, Colorado. Monthly temperature means and total precipita- tion have been plotted against each other and the points con- nected by lines. Except tor the marked difference in August rainfall, the polygons are quite similar. 100 miles east of Mesa Verde, with data from Park Point (Table 17) reveals a striking similarity of monthly and annual values. Park Point is warmer but has more precipitation than Pagosa Springs. Climo- graphs based on these data (Fig. 15) only serve to underscore the enigma concerning the scarcity of this species of pine in the Mesa Verde. Pinyon seedlings and saplings are abundant today even in stands wherejuniper is dominant in the older age-classes. Although Spencer (1964) asserted that pinyon mortality is high during the first 100 years, I 24 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN found that seedlings, at least, survive very well. In five 1 -meter quadrats established in 1962 under various microhabitats, 42 pinyon and 3 juniper seedlings were marked and their progress was followed. The latest observations in 1966 showed a loss of only four pinyon pines; the remaining seedlings appeared firmly established. In light of the greater number of pinyon seedlings than juniper seedlings and their good sur- vival rate, some change in the present forest can be expected. It is reasonable to conclude that the per- centage of pinyon tree cover will increase in the older age-classes during the next few hundred years in those stands left undisturbed. ACKNOWLEDGMENTS My sincere gratitude is extended to John W. Marr of the University of Colorado, who advised me through my ecological studies at Mesa Verde. To William A. Weber I owe my familiarity with the flora of the area, primarily through his determinations of most of my collections, but also as a result of my assisting him for several years in the University of Colorado Museum's Herbarium. Many other persons have contributed significantly to the collection of the data that has culminated in this paper. In particular, I acknowledge the help of Marilyn Colyer who assisted me in the field; of Orville A. Parsons, Soil Conservation Service, Fort Collins, Colorado, who provided the soils information; of Fred E. Mang, Jr., Project photographer; and of George A. King, architect of Durango, Colorado, for drafting the base map. Dendrochronological corrobo- ration was provided by Harold C. Fritts, Robert F. Nichols, David G. Smith, and Thomas P. Harlan, of the Laboratory of Tree-ring Research, University of Arizona, Tucson. To my wife, Margaret Ann, I give deepest thanks for her patient encouragement througli the many stages involved in the completion of this study. This paper is a revision of the author's doctoral dissertation. The research was supported by the National Geographic Society and the National Park Service througli the Wetherill Mesa Project. The man- uscript was reviewed by Richard P. Wheeler of the National Park Service, Washington, D.C. This is Contribution 49 of the Wetherill Mesa Proj- ect. LITERATURE CITED Anderson, S. 1961. Mammals of Mesa Verde National Park, Colorado. Univ. Kansas Pub!., Mus. Nat. Hist. 14: 31-67. Arnold, J. F.. D. J. Jameson, and E. H. Reid. 1964. The pin- yon-juniper type of Arizona: effects of grazing, fire, and tree control. U.S. Dep. Agr. Forest Service Production Res. Rep. No. 84. Arrhenius, G., and E. Bonatti. 1965. The Mesa Verde loess, p. 92-100. In Contributions of the Wetherill Mesa Archcological Project. Memoirs Soc. Amer. Archaeol. 19, Amer. Antiquity 31. Bader, E. H. MS. The vegetation of the Mesa Verde National Park, Colorado-a preliminary study. Unpublished M.A. thesis, Univ. Colo., 1932. Bannister, B. 1963. Dendrochronology, p. 161-176. In D. Brothwell and E. Higgs |eds.|. Science in Archaeology. Basic Books Inc., New York. Biesen, C. 1931. Evergreens of Mesa Verde. Mesa Verde Notes 2: 12-16. Cable, D. R. 1957. Recovery of chaparral following burning and seeding in central Arizona. Res. Notes, No. 28, For- est Service, Rocky Mountain Forest and Range Exp. Sta. Cain, S. A., and G. M. Castro. 1959. Manual of vegetation analysis. Harper & Bros., New York. Canfield, R. H. 1941. Application of the line interception method in sampling range vegetation. J. Forest. 39: 388-394. Cottam, G., and J. T. Curtis. 1956. The use of distance measures in phvtosociological sampling. Ecology 37: 451-460. Crocker, R. 1952. Soil genesis and the pedogenic factors. Quart. Rev. Biol. 27: 139-168. Curtis, J. D., and D. W. Lynch. 1957. Sylvics of ponderosa pine. U.S. Forest Service, Intermountain Forest and Range Exp. Sta. Misc. Publ. No. 2. Ogden. Utah. Daubenmire, R. F. 1959. A canopy-coverage method of veg- etational analysis. Northwest Sci. 33: 43-64. Douglas, C. L., and J. A. Erdman. 1967. Development of terminal buds in pinyon pine and Douglas-fir trees. The Pearce-Sellards Ser. No. 8. Texas Memorial Museum, Austin. Erdman, J. A. MS. Ecology of the pinyon-juniper woodland of Wetherill Mesa, Mesa Verde National Park, Colorado. Unpublished M.A. thesis. Univ. Colo., 1962. Erdman, J. A., C. L. Douglas, and J. W. Marr. 1969. The environment of Mesa Verde. Colorado. Archeol. Res. Ser. No. 7-B. National Park Service. Washington. D.C. Erdman, J. A., W. A. Weber, and J. M. Tucker. 1962. Quercus ajoensis in Colorado. Southwestern Natur. 7: 269-270. Fritts, H.C. 1965a. Dendrochronology, p. 871-879. /» H. E. Wright, Jr., and D. G. Frey |eds.]. The Quaternary of the United States. Princeton Univ. Press, Princeton, N. J. . 1965b. Tree-ring evidence for climatic changes in western North America. Monthly Weather Rev. 93: 421-443. Fritts, H. C. D. G. Smith, and M. A. Stokes. 1965. The bio- logical model for paleoclimatic interpretation of Mesa Verde tree-ring series. In Contributions of the Wetherill Mesa Archeological Project. Memoirs Soc. Amer. Archaeol. 19, Amer. Antiquity 31. Getty, H. T. 1935. New dates from Mesa Verde. Tree-Ring Bull. 1: 21-23. Glock, W. S. 1937. Principles and methods of tree-ring anal- ysis. Carnegie Institution of Washington, Publ. No. 486. Hanson, H. C, and E. D. Churchill. 1961. The plant community. Reinhold Publ. Corp., N. Y. 218 p. Historic Preservation, National Park Service, Washington, D.C. Howell, J. 1941. Pinon and juniper woodlands of the South- west. J. Forest. 39: 542-545. Jameson, D. A., J. A. Williams, and E. W. Wilton. 1962. Vegetation and soils of Fishtail Mesa, Arizona. Ecology 43: 403-410. Jenny. H. 1941. Factors of soil formation. McGraw-Hill. New York. 281 p. Johnsen, T. N. 1964. Merriam's pinyon zone. Plateau 37: 65-66. Kintigh, R. G. 1949. Effects of temperature on germination of pinon. J. Forest. 47: 622-626. Lindsey. A. A. 1956. Sampling methods and community attributes in forest ecology. Forest Sci. 2: 287-296. Major, J. 1951. A functional, factorial approach to plant ecology. Ecology 32: 392-412. Malde. H. E. 1964. Environment and man in arid America. Science 145: 123-129. 1950. Management of Ponderosa Pine in the South- west. U. S. Forest Service, Agr. Managr. No. 6. 218 p. Marr, J. W. 1961. Ecosystems of the east slope of the Front Range in Colorado. Univ. Colo. Studies, Ser. Biol. No. 8. 134 p. Martin, P. S., and W. Byers. 1965. Pollen analyses from the Wetherill excavations. In Contributions of the Wetherill Mesa Archeological Project. Memoirs Soc. Amer. Archaeol. 19, Amer. Antiquity 31. McGinnies, W. G. 1963. Dendrochronology. J. Forest. 61: 5-11. Meagher, G. S. 1943. Reaction of pinon and juniper seed- lings to artificial shade and supplemental watering. J. Forest. 41: 480-482. Merkle. J. 1952. An analysis of a pinyon-juniper community at Grand Canyon, Arizona. Ecology 33: 375-384. Mielke, J. L. MS. Leptographium root disease of pinyon pine. Mesa Verde National Park, Colorado. Unpublished report from the Division of Forest Pathology, 1948. 4 p. Nord, E. C. 1959. Bitterbrush ecology-some recent find- ings. U.S. Forest Service, Pacific Southwest Forest and Range Exp. Sta. Res. Note 148. Costing, H. J. 1956. The study of plant communities. W. H. Freeman, San Francisco. 440 p. Osborne, D. 1964. A prologue to the project, p. 1-17. /n A. C. Hayes, The archeological survey of Wetherill Mesa, Mesa Verde National Park, Colorado. Archeol. Res. Ser. No. 7-A. National Park Service, Washington, D.C. Parsons, O. A. MS. Soil survey of Wetherill Mesa, Mesa Verde National Park, Colorado. Manuscript, 1966, on file with the Division of Archeology, Office of Archeology and Pearson, G. A. 1931. Forest types in the Southwest as deter- mined by climate and soil. U.S. Dep. Agr. Tech. Bull. 247. -. 1964. Climatological data, Colorado. Annual Sum- Phillips, F. J. 216-223. 1909. A study of pinon pine. Bot. Gaz. 48: Poulton, C. E., and E. W. Tisdale. 1961. A quantitative method for the description and classification of range vegetation. J. Range Manage. 14: 13-21. Rand, P. J. 1966. Factors related to the distribution of pon- derosa and pinyon pines of Grand Canyon, Arizona. Diss. Abstr. 26(10):5679-5680. Smiley, T. L. 1958. Years, centuries, and millennia, p. 11-18. In T. L. Smiley (ed.j, Climate and man in the Southwest. Univ. Ariz., Tucson. Spencer, D. A. 1964. Porcupine population tluctuations in past centuries revealed by dendrochronology. J. Appl. Ecol. 1: 127-149. U.S. Weather Bureau. 1963. Climatological data, Colorado. Annual Summary, 1962, Vol. 67: 204-214. mary, 1963, Vol. 68: 204-214. Viereck, L. A. 1966. Plant succession and soil development on gravel outwash of the Muldrow Glacier, Alaska. Ecol. Monogr. 36:181-199. Wagener, W. W., and J. L. Mielke. 1961. A staining-fungus root disease of ponderosa, Jeffrey and pinyon pines. Plant Dis. Rep. 45: 831-835. Wanek. A. A. 1959. Geology and fuel resources of the Mesa Verde area, Montezuma and La Plata Counties, Colo- rado. U.S. Geol. Survey Bull. 1072-M: 667-721. Watson, D. 1934. The Navajos as fire fighters. Mesa Verde Notes 5: 28-31. Weber, W. A. 1965. Plant geography in the Southern Rocky Mountains, p. 453-468. In H. E. Wright, Jr., and D. G. Frey [eds.] , The Quaternary of the United States. Princeton Univ. Press, Princeton, N. J. Wells, P. V. 1962. Vegetation in relation to geological substratum and fire in the San Luis Obispo Quadrangle, California. EcoL Monogr. 32: 79-103. Welsh, S. L., and J. A. Erdman. 1964. Annotated checklist of the plants of Mesa Verde, Colorado. Brigham Young Univ. Sci. Bull., Biol. Ser., Vol. 4 (2). 32 p. Provo, Utah. Woodbury, A. M. 1947. Distribution of pigmy conifers in Utah and northeastern Arizona. Ecology 28: 113-126. Woodin, H. E., and A. A. Lindsey. 1954. Juniper-pinon east of the Continental Divide, as analyzed by the line-strip method. Ecology 35: 473-489. INFORMATION FOR CONTRIBUTORS Contributions to the Science Bulletin should be piimarily monographic in nature. For the most part only manuscripts of approximately forty or more typewritten pages will be ac- cepted. Papers will be published approximately in the order that they are received, pending avail- ability of funds. Authors must anange for finan- cing their publications, except that the Univer- sity Press and the hbrary of Brigham Young University will share the publication cost pro- portionately to their needs for copies of the article printed. In the preparation of manuscripts, authors are requested to follow the Style Manual for Biological Journals, American Institute of Bio- logical Sciences, 2000 P Street, N.W., Washing- ton, D.C., 20006. Manuscripts must be typed on one side of the paper only, double-spaced with ample margins. Footnotes should be avoided. To facilitate review by referees, send to the editor the original manuscript and one carbon copy, together with the illustrations. Copies of the original illustrations may be submitted, but should be of quality equal to the origiaals. Illustrations should be referred to as figures except for materials requiring inserts of special paper, which may be called plates. Illustrations shouJd be so designed as to fit when reduced into a one-column or fuU-page width. Special care must be taken to allow for proper reduc- tion in lettering (i.e., a 50% reduction of the figure means also a 50% reduction in the lettering). Photographs should be of a glossy finish, unblurred, and showing sharp con- trast. Line drawings should be made with black ink on heavy white drawing paper, blue trac- ing cloth, or blue-ruled coordinate paper. Use the same abbreviations on hne drawings as in text. Line dravdngs must be equivalent to a professional draftsman's work. Original drawings are preferable to photographs, even if they are large. Illustrations (line drawings or photo- graphs) should be numbered consecutively throughout the paper, and the approximate place of insertion should be indicated in the margins of the manuscript pages. Captions for illustrations should be assembled on a separate sheet, and each plate of figures must have its coiTesponding figure number pencilled hghtly on the back. Illustrations and cuts will be de- stroyed unless their retiuii is requested when proof is returned to the editor. A table title shouJd be a short, concise state- ment of what the table purports to show, and should not include information necessary to the intei-pretation of the table. Every column in the table should carry a head identifying the data in that column; the measure in which the data are given should be indicated at the head of each column. Tables should not be used when the same information can be given in a few lines of text, and should not duphcate informa- tion in text, graphs, or charts. Symbols (asterisk, dagger, etc.) should be used to indicate foot- notes to tables, with footnotes on the same page as the table. An abstract of less than four percent of the length of the paper should be prepared. This summary should be understandable without ref- erence to the body of the manuscript. The ab- stract must be on sheets separate from the man- uscript. Proof should be corrected immediately on receipt and returned to the editor. Authors should leave forwarding addresses if they move from the address sent with the manuscript. Reprints should be ordered when the proof is returaed. Address all manuscripts to Stanley L. Welsh, Department of Botany, Brigham Young Univer- sity, Provo, Utah 84601. I Brigham Young University Science Bulletin MU3. CCy.P. ZOOL. LIBRARY DEC 41970 HARVARD UNIVERSITY) DISTRIBUTION OF THE NATIVE TREES OF UTAH by Kimball S. Erdman BIOLOGICAL SERIES— VOLUME XI, NUMBER 3 SEPTEMBER 1970 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Stanley L. Welsh, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: Vernon J. Tipton, Zoology Feeeon L. Anderson, Zoology Joseph R. Murdock, Botany WiLMER W. Tanner, Zoology Ex officio Members: A. Lester Allen, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Chairman, University Publications The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from Pubhcation Sales, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for hbrary exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 84601. Brigham Young University Science Bulletin DISTRIBUTION OF THE NATIVE TREES OF UTAH by Kimball S. Erdman BIOLOGICAL SERIES— VOLUME XI, NUMBER 3 SEPTEMBER 1970 * TABLE OF CONTENTS Page ACKNOWLEDGMENTS 1 INTRODUCTION 1 METHODS 2 SPECIES DISCUSSION 4 DISCUSSION 26 TABLES 27 BIBLIOGRAPHY 33 DISTRIBUTION OF THE NATIVE TREES OF UTAH by Kimball S. Erdman* ABSTRACT Although botanists have been actively collecting in Utah for well over a century, the distribution of many plants are as yet imperfectly known. This is, in part, due to the scattered nature of Utah herbarium material. Also the terrain of the state has impeded collection in more rugged and remote areas. The present study gathers together the data from the major collections of native Utah trees and clarifies our present understanding of tree distributions in the state. The older collection records have been exten- sively supplemented by the field work of the author and other botanists. The great diversity of environments due to geog- raphy, physiography, and climatic features of Utah has resulted in an interesting tree flora of fifty species and at least three naturally occurring, recognized hybrids. Although a majority of Utah's trees are mountain species, a number are strictly desert plants restricted to the Virgin or Colorado River drainages. ACKNOWLEDGMENTS No distribution study of this scope is completed without the assistance of many people. I am grateful to Dr. Bertrand F. Harrison and Dr. Kent H. McKnight for their counsel and advice when this pro- ject was initiated as a master's thesis. Dr. Stanley L. Welsh and Dr. Earl M. Christensen have given enthu- siastic support to the continuation of this research and have added much to our present revised distri- bution maps. They have also been most helpful in the criticism of the manuscript. I would like to thank Dr. Albert E. Little, chief dendrologist of the United States Forest Service for his help and advice. I am also grateful to the curators of herbaria who assisted me: Dr. Walter P. Cottam, formerly of the University of Utah; Professor Arthur H. Holmgren of Utah State University; Mr. Stephen Clark at Weber State College; Dr. LeRoy K. Henry of the Carnegie Museum; Dr. Dan Nicholson of the U. S. National Herbarium; Dr. Charles Feddema of the U. S. Forest Service Herbarium; and to Dr. Howard S. Irwin and Dr. Patricia Kern of the New York Botanical Garden. Finally. I would like to acknowledge the contri- butions of many botanists during the past hundred years. The time and effort expended in searching out the plants of Utah's rugged terrain have been monu- mental. INTRODUCTION Althougli botanists have been active in Utah for well over a hundred years, there is still a real need for accurate surveys and studies of plant distribution. This paper is an attempt to summarize the distri- bution data for all the native trees of Utah. The earliest distribution maps of Utah tree species are those of Sudworth (1915, 1916, 1917, 1918, 1934) on the gymnosperms and Salicaceae of the Rocky Mountains. His maps were based on collec- tions, reportings, and generalizations based on terrain. He did not show sites of specimens except on his working maps. Sudworth's maps were later revised and republished by Munns (1938). There have been a number of works since then such as Preston's Trees of the Rocky Mountains (1940), Harlow and Harrar's treatment of economic trees (1958) and Fowell's work on the silvics of forest trees (1965) which in- cluded generalized maps of tree distribution. Some work on tree distribution is included in Choate's Forests in Utah (1965). Much more detailed maps of the genus Pimis were published by Critchfield and Little (1966) and Mirov (1967). Christensen pub- lished distributions of Qiterciis gambeUi and on certain naturalized species. Tamarix, Ulmiis, and Elaeagmis not covered in this paper (1949, 1950, 1955, 1962, 1963, 1964, 1965). Recently Youngberg (1966) treated the willows of Utah. Unfortunately, however, liis distribution maps are not particularly informative inasmuch as they record only the pres- ence of a species in a given county. •"Department of Biology, Slippery Rock State College, Slippery Rock, Pennsylvania. BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN There have been a number of studies of the veg- etation of portions of the state both pubUshed and unpubHshed. Among these are Graham's study of the Uinta Basin (1937), Stanton's thesis on the ecology and floristics of the Henry Mountains (1931), Wood- bury's paper (1933) on the biotic communities of Zion National Park, Burke's thesis on the plants of the Wellsville Range (1934), MacMilian's study of the floristics and ecology of the Deep Creek Mountains (1948), Preece's work on the plant communities of the Raft River Mountains (1950), Welsh's thesis on plant ecology in Dinosaur National Monument (1957), Buchanan's Plant Ecology of Bryce Canyon National Park (1960), Ellison's paper (1954) on the subalpine vegetation of the Wasatch Plateau, and Foster's thesis on major plant communities of the Great Basin in Utah (1966). Flowers (1959, 1960), Lindsay (1959), Hall (1960), and Woodbury and others (1958, 1959, 1960), published reports on the plants of major reservoir sites in Flaming Gorge and Glen Canyon. Harrison, Welsh, and Moore have pub- lished on the plants of Arches National Monument (1964). The selection of species to be included in this paper was difficult. In Utah thousands of acres are covered with pigmy forest where trees rarely reach 30 feet in height. One, therefore, is inclined to be liberal in his definition of a tree. In general, however, if a species consistently or at least commonly exceeded 10-15 feet in heiglit, was at least two or three inches in diameter, and had a fairly well defined trunk and crown, it was included. Plants such as Ribes aurem and Cormis stolonifera, which may be this large but definitely many stemmed and shrubby, are excluded. Also, plants which are commonly shrubs and rarely reach a height of 10 or 15 feet, such as Cowania stansbuhana, are not included. METHODS It has been the general practice of botanists when mapping distribution of plants to use base maps showing political subdivisions such as states and coun- ties and a few major lakes and rivers. The size of Utah's counties and the remarkable variation of topography within them necessitates a more detailed base map. I have therefore included rivers and also major mountain ranges (See Fig. 1 and the accompa- nying key for the features included). The distribution of many of Utah's trees are corre- lated with mountain ranges with elevations over 7,000 feet. Generally such areas will have sufficient precipitation to support a tree flora more varied than that of the pigmy forests typical of lower elevations. Only ranges with several square miles above 7,000 feet are shown on the map. The location of species as determined from herbar- ium specimens are indicated by dots on the maps. Certain of these specimens are cited in the species dis- cussion. There the herbaria are listed according to their standard abbreviations; U. S. National Herbar- ium (US), U. S. Forest Service Herbarium (USES), New York Botanical Gardens (NY), Carnegie Museum (CM), Utah State University (UTC), University of Utah (UT), Brigham Young University (BRY), Weber State College (no abbreviation.) Sightings and recorded occurrences are indicated by an "X." In most cases these represent species dis- cussed in theses or in published works, but are not validated by specimens in the herbaria consulted. Sightings represent communications from various bot- anists to the author. All of these sightings and recorded occurrences should eventually be verified by collections. MOUNTAIN RANGES AND PLATEAUS (highest elevation included); 1. Raft River Mountains 9,892 ft. 2. Grouse Creek Mountains 9,042 ft. 3. Goose Creek Mountains 8,690 ft. 4. Pilot Range ca. 8,000 ft. in Utah. 5. Cedar Mountains 7,039 ft. 6. Stansbury Mountains 11,031 ft. 7. Oquirrh Mountains 10.626 ft. 8. East Tintic Mountains 8,218 ft. 9. West Tintic and Sheeprock Mountains 9,154 ft. 10. Onaqui Mountains 9,067 ft. II. Simpson Mountains 8,450 ft. 12. Fish Springs Range 7,680 ft. 13. Deep Creek Mountains 12,101 ft. 14. Canyon Mountains 9,717 ft. 15. House Range 9,678 ft. 16. Confusion Range (southern part) 8,300 ft. 17. Needle Range 9,785 ft. 18. Wah Wah Mountains ca. 9,200 ft. 19. San Francisco Mountains 9,669 ft. 20. Mineral Mountains 9,619 ft. 21. Pine Valley Mountains 10.238 ft. 22. Beaver Dam Mountains 7,746 ft. 23. Markagunt Pla- teau 11,307 ft. 24. Paunsaugunt Plateau 9,630 ft. 25. Aquarius Plateau 11,328 ft. 26. Sevier Plateau 11,307 ft. 27. Tushar Mountains 12,173 ft. 28. Pavant Range 10,279 ft. 29. Wasatch Plateau 11,300 ft. 30. San Pitch Mountains 9,487 ft. 31. Wasatch Range 11,877 ft. 32. Wellsville Mountains 9,356 ft. 33. Bear River Range 9,980 ft. 34. Uinta Mountains 13,498 ft. 35. Roan Plateau (or West Tavaputs Plateau) 10,235 ft. 36. Roan Plateau (or East Tavaputs Plateau) 9,510 ft. 37. San Rafael Swell 7,921 ft. 38. La Sal Mountains 12,721 ft. 39. Abajo Mountains (or Blue Mountains) 1 1 ,360 ft. 40. Elk Ridge ca. 9,200 ft. 41. Henry Mountains 11,615 ft. 42. Kaiparowits Plateau 7,610 ft. 43. Navajo Mountain 10,388 ft. BIOLOGICAL SERIES. VOL. 11, NO. 3 NATIVE TREES OF UTAH ,->*• „ nwn UTAH OUTLINE MAP County linoe and physiographic features Fig. 1. Key to Map ptiysiograpliic Features RIVERS and LAKES: „ r^ ,, u . , A Bear Lake B. Bear River C. Great Salt Lake D. Weber River E. Jordan River F. Utah Lake G. Provo River H. Strawberry Reservoir I. Strawberry River J. Indian Creek K. Duchesne River L. Lake Fork Creek M. Uinta River N. Green River O. White River P. San Rafael River Q Col- orado River R. Dirty Devil and Fremont Rivers S. Escalante River T. San Juan River U. Pana River V. Virgin River W. Santa Clara River X. Sevier River Y. San Pitch River BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN SPECIES DISCUSSION UTAH OUTLINE MAP Fig. 2. Abies concolor (Gord. & Glend.) Lindl. Abies concolor (GovA. & Glend.) Lindl. (Fig. 2) White fir is one of the most widespread of Utah's conifers. It is commonly associated with Douglas-fir, blue spruce, and aspen throughout the state. With Douglas-fir it forms the very common conifer com- munity typical of the cooler north slopes in mountain canyons from 5,000 to 9,000 feet. White fir is conspicuously absent from Graham's (1937) treatment of the flora of the Uinta Basin. However, I found in looking over his collections at the Carnegie Museum that at least two of his collec- tions, numbers 9641 and 9940 had been identified as Abies lasiocarpa but were actually /I. concolor. There have been several additional collections in the Uinta Basin since then. Representative Specimens: Beaver Co., Wah Wall Mts., W.P. Cottam 8080, June 10, 1940 (UT); Box Klder Co., Raft River Mts., K.S. Erdman 15.^7, June 8„ 1965 (Weber State College); Cache Co., Logan Canyon, S. Flowers 1374, July 12, 1924 (UT); Grand Co., Roan Plateau (east),£.//. Graham 9940, July 31. 1935 (CM); Kane Co., Bryce Cmyon. Bitdiaih an 307, August 16, 1957 (UT); Piute Co., Tusliar Mts., K.S. Erdman 129, May 13, 1960 (BRY); Salt Lake Co., Oquirrli Mts., J.F. Page s.n.. May 13, 1965 (Weber State College); San Juan Co., Elk Ridge, K.S. Erdman 277, July 29, 1960 (BRY); Sevier Co., Gunnison, I..F. Uto-rf658. August 29, 1875 (US); Summit Co., China Meadows. D. Stephens s.n., July 9. 1966 (Weber State College); Tooele Co., Stanshury Range, B. Maguire 21789, June 25, 1943 (UTC); Utah Co.. Hobble Creek, g.F. //flmxON 8319, May 10, 1938 (BR Y); Washington Co., Beaver Dam Mts., L.S. Higgins 705. June 18, 1966 (BRY). Abies lasiocarpa (Hook.) Nutt. (Fig. 3) The finest and most extensive stands of subalpine fir in Utah occur in the high basins of the Uinta Mountains. This typical spireform habit of subalpine fir is a symbol of the high Uintas and other timberline areas throughout the state. From its upper altitudinal limits in southern Utah of about 11,500 feet, it descends as low as 8,000 feet on the cooler north slopes of the Raft River Mountains and other ranges of northern Utah. At higher elevations it is found most commonly with Englenian spruce and occasion- ally with lodgepole pine, limber pine, and aspen. Like other species which grow at high altitudes, its present known distribution retlects incomplete sampling of Utah's upper mountain slopes. UTAH .Abies lasiocarpa (Hook.) Nutt. Representative Specimens: Box Elder Co., Raft River Mts., K.S. Erdman 1545, June 8, 1965 (Weber State College); Cache Co., Garden City Summit, K.S. Erdman 2954, Septem- ber 11, 1965 (Weber State College); Carbon Co., Roan Pla- teau (west), S.L. Welsh & E.M. Christensen 6542, August 10, 1967 (BRY); Duchesne Co., Uinta Mts., E.H. Graham 6633, July 3, 1931 (CM); Garfield Co., Boulder Mt., K.S. Erdman 2825, September 5, 1965 (Weber State College); Garfield Co., Henry WH., R.McVaiigh 14584, June 19-20, 1953 (NY); Grand Co., La Sal Mts., A'.5. Erdman 216, July 28, 1960 (BRY); Iron Co., Cedar Breaks. S.L. Welsh & E.M. Christen- sen 2654, September 11, 1963 (BRY); Salt Lake Co., Lamb's Canyon, W.P. Coltam 14373. July 5. 1956 (UT); Sevier Co., Monroe Mt., K.S. Erdman 2711, September 3, 1965 (Weber State College); Summit Co., Bald Mt., K.S. Erdman 314, August 25, 1960 (BRY); Tooele Co., Stansbury Mis., E.M. Christensen s.n., July 29, 1963 (BRY). Junipents osteosperma (Torr.) Little (Fig. 4) Utah juniper is probably the most widespread and BIOLOGICAL SERIES. VOL. II, NO. 3 NATIVE TREES OF UTAH UTAH OUTLINE MAP Counly linoB and Fig. 4. Jiinipenis osteospenna (Torr.) Little common conifer in Utah. It is the characteristic tree of lower and desert mountain slopes from 4,000 to 7,000 feet elevation, and often forms pure stands of considerable size, particularly in regions of recent invasion. Together with the two pinyon pine species in Utah it forms the pigmy forests of the desert ranges and plateaus. At higlier elevations Utah juniper is associated with Ganibel oak, ponderosa pine and other species. Sargent (1922) and Preston (1940) report a closely related species, J. monosperma, as common through- out Utah. However, the collection records and field work do not reveal its presence in Utah. Representative Specimens: Beaver Co., Wall Wah Mts., W.P. Cotlam 8045, June 8, 1940 (UT); Box Elder Co., Raft River Mts., K.S. Erdman 1519, June 8, 1965 (Weber State College); Millard Co., Canyon Mts., K.S. Erdman 133, May 13, 1960 (BRY); Millard Co., House Rock Mt,s.,/F. Brenckle 51027, March 24, 1951 (NY); Piute Co., Marysvale, P.A. Rvdberg & E.C Carlton 7008, July 21, 1905 (US); San Juan Co., Island in the Sky, G. Moore 243, July 9, 1964 (BRY); San Juan Co., Monument Valley, //.C Culler 3000, August 28, 1939 (NY); Summit Co., Echo, M.E. Jones s.n.. May 7, 1890 (US); Tooele Co., Stansbury Island, W.P. Collam 7607, April 13, 1940 (LT); Uintah Co., Dinosaur Nat'l. Monu., E.H. Graham 9162, June 10, 1935 (CM); Washington Co., Toquerville, O.A. Olsen 399, December 22, 1930 (USPS). Although /. scopiilonim may be a foothill species, it is common up to 9,000 feet on mountain slopes and is a characteristic tree of high rocky crags. The oldest known specimen of this species, commonly known as the Jardine Juniper, is so situated on the cliffs of Logan Canyon in northern Utah. This tree, probably the largest of its kind, is 27 feet in circum- ference and 44 feet in height and has been estimated to be over 3,000 years old. (Pomeroy & Dixon 1966.) UTAH Fig. 5. Juniperus seopulonim Sarg. Representative Specimens: Beaver Co., Beaver Canyon, fi. Slahmam 8113a, December 15, 1934 (US); Box Elder Co., Raft River Mts., K.S. Erdman 1515, June 8, 1965 (Weber State College); Daggett Co., Flaming Gorge, H.C. Cutler 3445, June 21, 1940 (NY); Duchesne Co., Roan Plateau (west), E.H. Graham 9463, June 29, 1935 (CM); Garfield Co., Henry Mts., R. McVau^h 14488, June 7, 1953 (NY); Millard Co., Canyon Mts., B.L. Robins 90, August 16, 1913 (USPS): Millard Co., Wah Wah Mts., B. Maguire & ,4. Holm- gren 25100, May 19, 1945 (UTC); San Juan Co., Elk Ridge, Twiss s.n., July 16, 1913 (UT); San Pete Co., Six Mile Can- yon, J.A. Willey 356, June 19, 1913 (USPS); Summit Co., Holiday Park, Mrs. J. Clemens s.n., August 14, 1911 (NY); Tooele Co., Stansbury Mts., E.M. Christensen s.n., July 29, 1963 (BRY); Tooele Co., Wendover, W.P. Cottam 7085, May 3, 1937 (UT); Washington Co., Beaver Dam Mts., L.C. Hig- gins 11 A, July 9, 1966 (BRY); Weber Co., Ogden Canyon, l.H. Pommel & RE. Blackwood 3722, July 17, 1902 (US). Junipents seopulonim Sarg. (Fig. 5) Rocky Mountain Juniper is probably second only to Utah juniper as the most widespread tree species in Utah. In view of the fact that it is found on almost every major mountain range in Utah, further work should reveal its presence in the La Sal and Pine Valley Mountains. Picea engelmannii Parry (Fig. 6) The high rugged summits of Utah's many moun- tain ranges are not very accessible. Our knowledge of the distribution of subalpine species such as Engle- man spruce will reinain incomplete until all such areas are carefully searched. Apparently this is a wide- spread species found not only in the main mountain BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN UTAH OUTLINE MAP Fig. 10. Pimis ediilis Enge\m. Representative Specimens: Beaver Co., Beaver, S.5. Hutch- ings 8444, May 2, 1938 (NY): Carbon Co., Price, R. Hardv s.n.. May 20, 1937 (UT); Duchesne Co., Red Crecl<, J. Brotherson 480, July 20, 1965 (BRY): Emery Co., Emery,/. Tidestrom 1422, July 8, 1908 (US): Garfield Co.. Aquarius Plateau, Vickerv 615, May 19, 1956 (UT): Grand Co., Arches N.M.. K.S. Erdman 188, July 26, 1960 (BRY): Kane Co., Kanab, Knudscn 39, December 29, 1950 BRY): San Juan Co., Elk Ridge, K.S. Erdman 279, July 29, 1960 (BRY); Tooele Co., Lofgreen, B.F. Harrison 7236, April 15, 1934 (BRY): Uintah Co., Dinosaur Nat'l. Monu., E.H. Graham 9155, June 10, 1935 (CM). Pimis flex His James. (Fig. 1 1) Limber pine is a characteristic tree of high exposed ridges, where it is often in pure stands. However, it inay also be associated with Cercocarpus ledifoliits, Juniperus scopulorum and other high mountain con- ifers. Occasionally it may occur as low as 7,500 feet. A tree near Alta in the Wasatch National Forest 28 feet in circumference and 43 feet in heiglit is the larg- est known limber pine. (Communication from the U.S.F.S.) A closely related species, Pimis alliicaulis, has re- peatedly been reported to occur in Utah (Mirov 1967, Critchfield & Little 1966, Choate 1965). However, extensive field and herbarium investigations have fail- ed to reveal any evidence of this species occurring in Utah. Both Little and Choate (personal communic- ation) have indicated that the claims of distribution in Utah may be based on old unverified Forest Ser- vice reports. Representative Specimens: Box Elder Co., Pilot Mt.. W.P. Cottam 4543, July 6, 1929 (BRY): Duchesne Co., Roan Fig. 1 1 . Pimis fle.xilis James Plateau (west), K.S. Erdman 2565, August 18, 1965 (Weber Slate College): Garfield Co., Bryce Canyon, B.F. Harrison 12039, May 10, 1953 (BRY): Garfield Co., Henry Mts., R. McVaugh 14478, June 7, 1953 (NY): Grand Co., Roan Plateau (east), C Vickerv & D. Wiens 1804, July 13-15, 1956 (UT): San Pete Co., Manti, J. Willey 358, June 19, 1913 (irrC): Summit Co., near Kamas, K.S. Erdman 300, August 25, 1960 (BRY): Tooele Co., Stansbury Mts., E.M. Chrislensen s.n., }u\y 29, 1963 (BRY): Tooele Co., Stansbury Mts., J.A. Harris C20702, Julv 23. 1920 (CM): Uintah Co., Brush Creek, EH. Graham 8271, June 23, 1933 (CM); Wash- ington Co., Pine Valley Mts., W.P. Cottam 8924, June 26, 1941 (UT): Wayne Co., Thousand Lake Mt., G.K. Gilbert 121, July 23, 1875 (US); Weber Co., Ben Lomond Peak, W. Call 109, August 30, 1940 (UT). Finns monophylla Torr. & Frem. (Fig. I 2) Singleleaf pinyon is a common member of the pigmy forest throughout the basin and range province of Utah and Nevada. Although this species is in the canyon and East Tintic Mountains, it is inexplicably absent from the main mountain system in Utah ex- cept in the extreme northern and southern portions. Growing with P. edulis in southern Utah, it extends as far east as the Paria River. There are a few isolated stands in the northern Wasatch Mountains and it is re- ported to be present in the Crawford Range. Rich County on the Utah-Wyoming border. Representative Specimens: Beaver Co.. Wah Wah Mts., B. Magidre 20965, June 20, 1941 (UTC); Box Elder Co.. Raft River Mts., W.P Cottam 2953. June 6, 1928 (BRY): Cache Co., Blacksmith lork. Flowers 17038, May 12, 1961 (BRY); Kane Co., Paria, B.F. Harrison 12055, May I I, 1953 (BRY); Millard Co., Canyon Mts., K.S. Erdman 2642, September 1, 1965 (Weber State College); Tooele Co., Deep Creek Mts.. J.A. Harris C28613, August 5, 1928 (CM); Tooele Co., West Tintic Mts., B. Magiiirc 20730, June 15, 1941 (NY); Wash- ington Co., Rockville, M.E. Jones 5220, May 15, 1894 (US). BIOLOGICAL SERIES. VOL. 1 I, NO. 3 NATIVE TREES OF UTAH UTAH OUTLINE MAP Fig. I 2. Pinus iiionophyHa Torr. & Frem. Pimts ponderosa Laws. (Fig. 13) Ponderosa pine is one of the most common con- ifers at elevations between 6,500 and 9.500 feet in the mountains and plateaus of southern Utah. Al- thougli occasionally occurring in open park-like stands, it is commonly associated with other species such as Gambel oak, aspen, juniper, white fir, etc. Ponderosa pine is not present in the mountains of the basin and range province of western Utah except for possibly two exceptions. It has been collected in the Wah Wah Mountains, and there is a specimen (USPS) collected in 1913 in Oak Creek Canyon, Canyon Mountains, by Bert L. Robins. This canyon is the site of a ponderosa pine plantation, but I did observe a few larger more isolated individual trees about one-fourth mile up the canyon. It is undetermined whether the older trees are natives to the canyon or remnants of earlier experimental planting. Ponderosa pine occurs only sporadically in the Wasatch Plateau of Sevier County and only in a few sites in the Wasatch Mountains of Utah County. How- ever, it is common at moderate elevations throughout the Uinta Basin. It has been collected on the north slope of the Uintas only in Daggett County. Why this species is absent from the northern Wasatch Range when it grows far to the north in Idaho and Montana is a most interesting problem that reflects the general distribution of this species in the West. Ponderosa pine is common from Arizona to Canada and from the Pacific coast to the Black Hills of South Dakota and Nebraska. It is absent from the central part of its range, namely in northern portions of Nevada and Utah and in southern regions of Idaho and Wyoming. No satisfactory answer has been found, although Baker and Korstian (1931 ) suggested that moisture deficiency early in the growing season might be a possible explanation. ^0 [( 1 ii ■•. i [ ' • ;1U= •;■:■' ! ' / UTAH OUTLINE MAP County linae nnd Fig. 13. Pimts i^onderosa Laws. Representative Specimens: Beaver Co.. Wah Wah Mts., S.L. Welsh & EM. Christensen 2687, September 13. 1963 (BRY); Daggett Co., Eagle Creek, B.F. Harrison & Larsen 1%19, June 19, 1934 (BRY); Duchesne Co., Moon Lake, B.F. Harrison & Larsen 7599, May 13, 1934 (BRY): Garfield Co., Henrv Mts,,/?. McVaugh 14479, June 7, 1953 (NY);Garfield Co., Panguitch Lake, A/.£. Jones 6025, September 10, 1894 (US): Grand Co., Roan Plateau (east), E.H. Graham 9933, July 29, 1935 (CM); San Juan Co., Abajo Mts., K.S. Erdman 248, July 28, 1960 (BRY); San Juan Co., La Sal Mts., H.C. Cutler 2655, July 9, 1939 (NY); San Pete Co., Ephraim Can- yon, /. Tidestrom 1144, June 11, 1908 (US); Sevier Co., Tushar Mts., K.S. Erdman 2702, September 2, 1965 (Weber State College); Utah Co.. Hobble Creek, B.F. Harrison 8319, May 10, 1938 (BRY); Washington Co., Pine Valley Mts.. W.P. Cottam 6851, May 10. 1936 (UT). Pseudotsuga menziesii (Mub .) Franco (Fig. 14) In nearly every mountain range in Utah over 7,000 feet in elevation there is an appropriate habitat for Douglas-fir, namely a canyon with a moist, north- facing slope. This species is almost universally present throughout Utah's mountains, and its absence from a range on the distribution map probably reflects lack of collection rather than absence of the species. Douglas-fir is adapted to a variety of habitats and may be associated with nearly all of the other coni- fers of the state. Although Douglas-fir is logged to BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN systems of Utah, such as the Wasatch and the Uintas where it is especially common, but also on isolated ranges such as the Deep Creek, La Sal, and Abajo Mountains. UTAH UTAH OUTLINE MAP Fig. 6. Picea engelinannii Parry. Representative Specimens: Box Elder Co., Raft River Mts.. W.M. Stephensen s.n.. May 18. 1965 (Weber State Col- lege); Cache Co.. near Monte Cristo. K.S. Erdinan 2600, August 28. 1965 (Weber State College); Duchesne Co.. Moon Lake, EH. Graham 6523, July 1, 1931 (CM); Grand Co.. Roan Plateau (east), Vkkerv 1802, July 13-15, 1956 (UT); Iron Co., Cedar Breaks. 5./.. Welsh & E.M. Christemen 265 3, September 11. 1963 (BRY); San Juan Co.. Abajo Mts.. P Rvdberg & A.O. Ganetl 9718. August 17-20. 1911 (NY); Sevier Co., Mt. Hilgard, H'. li'. Eggleston 10350, August 1. 1914 (US); Tooele Co., Stansbury Mts., E.M Chnstemen s.n., July 29, 1963 (BRY). Picea pungens Engelm. (Fig. 7) Througliout various canyons in Utah's mountain backbone from Cedar City to the Uintas, there are small groves of spruce growing alone or in association with Douglas-fir, white fir, and aspen. The traits of symmetry and blue or silver gray cast so common in horticultural plantings, are rarely seen in moun- tain-grown blue spruce. Rarely is it abundant; and of- ten, while common in one canyon, it will be absent from similar canyons just over the ridge. Its presence in the Deep Creek Mountains and on Navajo Moun- tain certainly indicates that deserts need not prohibit its migration. Careful searches on other isolated ranges such as the La Sal may yet reveal its oc- currence. Representative Specimens: Beaver Co.. Beaver Canyon, S.S. Hiilchmgs 8443, May 2, 1948 (US); Duchesne Co., Moon Fig. 7. Picea pungens Engelm. Lake, Harrison di Larson 7729. June 16, 1934 (BRY); Gar- field Co.. Escalante Mts.. F. W. Pennell & R.L. Schaeffer 21943. June 19. 1938 (NY); Iron Co., Cedar Creek, S.L. Welsh & E.M. Christensen 2616. September 1 1, 1963 (BRY); Piute Co., Monroe Mt., A'..S. Erdman llil. September 3, 1965 (Weber State College); San Pete Co., East of Mayfield, K.S. Erdman 2940. September 7. 1965 (Weber State Col- lege); Utah Co.. Tucker, S.L. Welsh & G. Moore 3503, August 14, 1964 (BRY). Pimis aristata Engelm. (Fig. 8) Slow growing and often exceptionally long-lived, the bristlecone pine is one of the most interesting trees native to Utah. Its most characteristic habitat is on the wind swept ridges near timberline. Some in- dividual trees in the White Mountains of California and Nevada have survived for over 4,000 years. (Fer- guson 1968;Schulman 1958.) Some of the best stands of bristlecone pine in Utah are in Cedar Breaks National Monument and in the adjoining Fishlake National Forest. Age studies during the past several years have been conducted on trees in these groves. So far the oldest tree sampled is in the Blowhard Point area of Cedar Breaks. It is esti- mated to be 1,630 years in age. A 1,560-year-old tree was found in Bryce Canyon National Park (personal communication. Dixie National Forest officials). The distribution of bristlecone pine in Utah is very spotty. Its tendency to grow in higli rugged mountain areas has been a hinderance to collection. Stanton (19.^1 ) described the vegetation of the Henry Moun- tains. His list of plants lacks a reference to bristlecone pine. However, in 1953 a botanist collected speci- BIOLOGICAL SERIKS. VOL. 1 1 , NO. 3 NATIVE TREES OF UTAH mens from a grove on the southern slopes of Mt. Mill- ers in the Henry Mountains. Likewise, groves have re- cently been discovered in the remote Roan Plateau of the Uinta Basin. As yet, the reports of bristlecone pine in the Uinta Mountains are unconfirmed. It has been found in three Great Basin ranges in Utah: Wah Wah, Needle, and Deep Creek Mountains. There are unconfirmed reports of it in the San Francisco Moun- tains. UTAH OUTLINE MAP County lineB find Likewise the Critchfield and Little report (1966) of lodgepole pine in the Raft River Mountains of north- western Utah appears to be in error. UTAH Fig. 8. Pimis anslata Engelm. Representative Speeiincns: Beaver Co., Needle Range, S.Z,. Welsh & G. Moore 3188. July 18, 1964 (BRY); Beaver Co.. Wall Wah Mts., B. Magiiire & AM. Holmgren 25102, May 19, 1945 (UTC); Carbon Co., Price Canyon, ,S. Flowers s.n., Jan- uary 12. 1934 (UT); Duchesne Co., Roan Plateau (west), D. AtwooJ 748, September 2(1, 1965 (BRY); Garfield Co., Escalante Mts., K.S. Erdman 35, May 10, 1960 (BRY); Gar- field Co., Henry Mts., R. McVaugh 14481, June 7, 1953 (NY); Garfield Co., Panguitch Lake, M.E. Jones 6003, Sep- tember 7, 1894 (US); Iron Co., Cedar Breaks, B.F. Harrison 9864, August 24, 1940 (BRY); Washington Co., Pine Valley Mts..F. H'. Gould 1876, July 9, 1942 (NY). Pinus coiUurta Dougl. (Fig. 9) Lodgepole pine is abundant in the Uinta Moun- tains where it grows from 7,000 to 10,000 feet, and in the extreme northern Wasatch or Bear River Range. Contrary to published reports, the southern most station of this species yet found in Utah is Daniels Canyon near the summit. Although Critch- field and Little in Geographic Distribution of the Pines of tlie World ( 1966) show this species through- out the Wasatch Plateau region south of Daniels Can- yon, this is apparently based on unconfirmed reports. Kig. 9. Ptnus contorts Dougl. Representative Specimens: Cache Co., Garden City Sum- mit. K.S. Erdman 2955, September 11, 1965 (Weber State College); Duchesne Co.. Mt. Hmmons, B.F. Harrison & Larsen 11A4. June 16. 1934 (BRY); Summit Co., Soapstone, A'.5. Erdman 305, August 25, 1960 (BRY); Uintah Co., Taylor Mt., E.H. Graham 6382, June 24, 1931 (CM); Wasatch Co., Daniels Canyon, K.S. Erdman 2448, August 16, 1965 (Weber State College). Pinus edulis Engelm. (Fig. 1 0) Pinyon pine is widespread and abundant on lower mountain slopes and plateaus of eastern Utah. It does not extend into the ranges of the basin and range pro- vince except in the Mineral, East Tintic, and Canyon Mountains. Although present in the Wasatch Moun- tains it does not occur north of Spanish Fork Can- yon. Pinyon pine is common on the southern slopes of the Uintas but only in the Flaming Gorge area does it grow on the north slope. Althougli the ranges of Utah's two pinyons are generally separate, there is overlap in the East Tintic Mountains and Canyon Mountains as well as in ex- treme southwestern Utah. In these localities there is evidence of hybridization between Pinus edulis and P. monophylla (Cole & Hiner 1963). The largest known specimen of Pinus edulis, 33 feet in height and 1 1 feet in circumference, is in the Manti-La Sal National Forest of southeastern Utah. (Pomeroy & Dixon 1966.) 10 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN some extent, it is most important in Utah as a watershed species. UTAH OUTLINE MAP County linoB and Fig. 14. Psctidotsiiga menziesii {Miib.) Franco Representative Specimens: Box Elder Co., Raft River Mts., K.S. Erdimn 1533, June 8, 1965 (Weber State College): Carbon Co., Roan Plateau (west), E.H. Graham 9642, July 13, 1935 (CM); Daggett Co., Flaming Gorge, L. Williams 494, June 2, 1932 (NY); Duchesne Co., Moon Lake, EH. Graham 9386, June 25, 1935 (CM); Garfield Co., Henry Mts., A/.£ Jones 5695bc, July 27, 1894 (US); Grand Co., La Sal Mts., K.S. Erdman 212, July 28, 1960 (BRY); Millard Co., Chalk Creek, E.M. Christensen s.n., August 3, 1966 (BRY); San Juan Co., Abajo Mts., K.S. Erdimn 247, July 28, 1960 (BRY); San Juan Co., Navajo Mt.,//.C. Cutler 2817, July 30, 1939 (NY); Tooele Co., Stanshury Island, W.P. Cottam 7606, April 13, 1940 (UT); Utah Co,, Rock Canyon, B.F. Harrison 6181, May 12, 1933 (BRY); Washington Co., Pine Valley Mts., L.N. Goodding 875, May 17. 1902 (US); Weber Co., Ben Lomond Peak, IV. Call s.n., June 28, 1940 (UT). Acacia greggii A. Gray (Fig. 1 5) The Beaver Dam Wash of Washington County, re- presents the northern limits of the catclaw acacia, named for its sharp spines, and is the only recorded collecting site in Utah. There is one stand near Terry's ranch which numbers about 40 individual trees. Representative Specimen: Washington Co., Beaver Dam Wash, ,?./.. Welsh 1480, April 8, 1961 (BRY). Acer glabntmJorr. (Fig. 16) This maple is widespread in Utah mountains and generally grows at higher elevations than /leer ^ra«c//- deiitatum. Rocky Mountain maple occurs along streams and in conifer and aspen forests. The leaves are variable in shape and may even be palmateiy compound. A common fungus, almost un- Flg. 1 5. A cacia greggii A. Gray iversal in this area, results in striking briglit red patches on the leaves. Representative Specimens: Box Elder Co.. Raft River Mts., C McMillan 1504, June 18, 1948 (UT); Cache Co., Dry UTAH OUTLINE MAP i Fig. 16. Acer glabriim Ton. BIOLOGICAL SERIES. VOL. 11. NO. 3 NATIVE TREES OF UTAH 11 Canyon. B. Maguire 13731. May 9, 1936 (NY); Carbon Co.. Castlegatc. A.H. Holmgren & S. Hansen 3508. June 29, 1944 (NY): Garfield Co.. Bryce Canyon, K.E. Weight B-31-90, June 29. 1931 (US); Garfield Co.. Henry Mts., W.D. Stanton 43, June 20, 1930 (UT); Grand Co., Roan Plateau (east). E.H. Graham 9953. August 1, 1935 (CM): Juab Co.. Deep Creek Mts., W.P. Cottam 3202, June 16, 1928 (UT); Millard Co., Wall Wall Mts., B. Maguire & A. H. Hohneren 25096, May 19, 1945 (UTC); Piute Co., Box Creek, H.J. Skidmore 20, August 16, 1940 (USPS); Salt Lake Co., Big Cottonwood Canyon, P.A. Rvdherg 6535, June 30, 1905 (US): San Juan Co., hdSiiyH^., P.A. Rvdberg & A.O. Garrett 8&12, iuly 12, 1911 (NY): Tooele Co.. Sheeprock Mts.. E.M. Cliristensen s.n., July 28, 1963 (BRY): Uintah Co., Diamond Mt.. E.H. Graham 8125, June 7, 1933 (CM): Washington Co., Pine Val- ley MU..F.W. Gould 1912, July 22, 1942 (NY). UTAH OUTLINE MAP County ImoB old phy9iograpKic teatui Fig. 17. Acer grandidentattim Nutt. Acer grandidentatum Nutt. (Fig. 17) The bigtooth maple is common along the nortli- south mountain axis in Utah and is one of the major contributors to Utah's autumn pageant. Wliile primar- ily a riverbottom species associated with willows, birch, alders and aspen, it is also frequently found throughout the mountain brush zone along with oak, chokeberry, and other species. Althougli A. grandi- dentatum is abundant from the Wasatch Mountains south to the Pine Valley and Beaver Dam Mountains, it is uncommon in both the more eastern and western portions of the state. It has been collected in Wah Wah Canyon, East Tintic Mountains and reported to be in the Stansbury and Oquirrh Mountains in the basin and range province. Bigtooth maple has been collected numerous times in the Abajo Mountains but it has not yet been found in the La Sals or Henrys. It has not been collected east of Rock Creek in the Uinta Mountains even though it occurs sporadically far to the east along the southern rim of the Uinta Basin. Representative Specimens: Beaver Co., Beaver Canyon, K.S. Erdman 109, May 13. 1960 (BRY): Beaver Co., Wah Wah Mts,, W.P. Cottam 8051, June 8, 1940 (UT): Cache Co., Logan Canyon, K.S. Erdman 2942, September 11, 1965 (Weber State College): Duchesne Co., Red Creek, E.H. Gra- ham 9411, June 26, 1935 (CM): Garfield Co., Bryce Canyon, H Buchanan 371, June 28, 1958 (UT); Grand Co., Desola- tion Canyon, £".//. Graham 9958, August 1, 1935 (CM): Kane Co.. Kaiparowits Plateau. J.R. Miirdock 399. May 3, 1962 (BRY): Millard Co., Canyon Mts.. A'.5. Erdman 2637, Sep- tember 1, 1965 (Weber State College); Salt Lake Co.. Red Butte Canvon, A.O. Garrett \1S7. July 12, 1906 (UT): San Juan Co.. Abajo Mts., P.A. Rvdherg & A.O. Garrett 9264, July 28-29, 1911 (NY); San Pete Co., Ephraim Canyon, /. Tidestrom 1003, May 16, 1908 (US); Washington Co., Pine Valley Mts. N. Goodding 859, May 17, 1902 (US). UTAH Fig. 18. Acer negundo h. Acer negiindo L. (Fig. 18) Well adapted to a variety of climates, box elder is widely distributed throughout the United States. In Utah it is restricted to streamsides and washes where it is often one of the most common trees. Box elder's distribution in Utah is spotty and may reflect in part an extension of its range due to its frequent use as an ornamental tree. Representative Specimens: Beaver Co.. Beaver Canyon, /. Tidestrom 2872, September 2, 1909 (US); Duchesne Co., Roan Plateau (west), S.L. Welsh & E.M. Christensen 6618, August 12, 1967 (BRY); Garfield Co., Henry Mts., R. McVaugh 14490. June 7, 1953 (NY); Salt Lake Co.. Lamb's Canyon, Vickerv 1505, October 8, 1957 (UT): San Juan Co., La Sal Mts., H.C. Cutler 2643, (NY): San Juan Co,, Natural Bridges Nat'l. Monu., S.L. Welsh & G. Moore 3587, August BRICHAM YOUNG UNIVERSITY SCIENCE BULLETIN 15, 1964 (BRYl; Sevier Co., Salina Canyon, A'.S. Enlman 2919, September 7, 1965 (Weber State College): Tooele Co., Dugway Range, J.A. Harris C286:4, Angiist 6, 1928 (CM): Uintah Co., Dinosaur Nat'l. Monu., E.H. Graham 1114. May 18, 1933 (CM): Utah Co., Thistle, A'.5. ErJman 2589, August 18, 1965 (Weber State College); Washington Co., Beaver Dam Mts., i.e. Hi^giiis 688, May 27, 1966 (BRY); Weber Co., Ogden Canyon, A.E. Hilchcock 1499, August 19. 1913 (US). UTAH UTAH Fig. 19. Aliiiis Icmdfolia Nutt. Alnus temiifolia Nutt. (Fig. 19) Thin-leaf alder forms large shrubby thickets in high mountain regions, althougli it is more common along streams at lower elevations. It is like bristlecone pine and blue spruce in that it has a very spotty distri- bution. While very abundant in one canyon, it may be absent from drainages on either side. For exainple, alder is very common in the drainages of the Fremont River and Salina Creek, but is absent from Twelve- mile Creek ten miles to the north. It has been found at only one site along the Wasatch Plateau but is common in certain canyons of the Wasatch and Uinta Mountains. It is present in only three of the isolated ranges of the state, the Raft River, La Sal and Abajo Mountains. Representative Specimens: Box Elder Co., Raft River Mts., Preece 775, June 29, 1947 (UT); Salt Lake Co., Red Butte Canyon, 5. G'. ^roi-ess.n.. May 15, 1900 (US): San Juan Co., Abajo Mts., P.A. RvJhcrg & A.O. Garrett 9252, July 28-29. 191 1 (NY): Sevier (\x, Saline Canyon, K.S. Enlman 2918, September 7, 1965 (Weber State College): Uintah Co., Ashley Creek, EH. Graham 6259, June 23, 1931 (CM): Wasatch Co., Strawberry Reservoir, /?.//. Foster 268, October 13, 1964 (BRY): Wayne Co., Fremont River, K.S. Enlman 2807, September 4, 1965 (Weber State College). Fig. 20. Bcliila occideiilahs Hook. Betitla occicleiitalis Hook. (Fig. 20) River birch is common throughout the state and is usually found near running water, springs, and seeps. In the basin and range province, it has been collected only in the Deep Creek Mountains. Representative Specimens: Box Elder Co., Raft River Mts., Porter 75, July 2, 1949 (UT): Davis Co., Bountiful, Mrs. J. Clemens s.n., September 22, 1909 (NY); Duchesne Co., Roan Plateau (west), K.S. Erdman 2517, August 17, 1965 (Weber State College); Garfield Co., Bryce Canyon, K.E. Weigh! G-31-114, July 10, 1931 (US); Grand Co., La Sal Mts., A'.5. Erdman 221, iu\v 28. 1960 (BRY): Iron Co., Paro- wan, W.W. Eggleston 14902, September 17, 1918 (US); Millard Co., Pavant Mts., yifrs. F.M. Stone 223, May 9, 1934 (NY); San Juan Co., Elk Ridge, /'..4. Rvdherg& A.O. Garrett 9321, July 31. 1911 lNY);San Pete (To., Ephraini Canyon, Jorgensen 42, August 5, 1950 (UT); Uintah Co., Dinosaur Nat'l. Monu., E.H. Graham 7542, April 28, 1933 (CM); Utah Co., Provo Canyon, S.L. Welsh 3813, October 20, 1964 (BRY). Celtis reticulata Torr. (Fig. 2 1 ) In Utah the netleaf hackberry is very often over- looked. Indeed, there is little to make this tree at- tractive. Its gnarled and twisted branches, often in- fected with a fungus causing a witches broom effect, bear leaves equally deforined by insect galls. The distributional pattern of this tree type in Utah is certainly distinctive. Generally it is a tree of the Colorado River and Virgin River drainages, but it is found in a few other isolated areas. Its presence in the Uinta Basin along the Green River can be explained by its migration up that stream valley. However, it is quite common on the foothills of the Wasatch Moun- BIOLOGICAL SERIES. VOL. 1 1, NO. 3 NATIVE TREES OF UTAH 13 tains and on low mountain ranges immediately around the Great Salt Laiemont poplar may have had a greater range. Other stands could be hybrids between narrow-leaf cottonwood and other species such as P. sargentii. (Bennion, Vickery & Cottam 1961.) Representative Specimens: Popiilus fremontii: Carbon Co., Castle Dale, / Boiins s.n.. May 19, 1963 (NY); Grand Co., Arches Nat'l. Menu., B F. Harrison 11162, April 27, 1947 (BRY); Kane Co., Kanab, W.P Cottam 4319, June 13, 1929 (UT); San Juan Co., Natural Bridges Nat'l. Monu., R Rvdberg & A.O. Garrett 9484, August 4-6, 1911 (NY); Sevier Co., Pavant Range, K.S. Erdman 2705, September 2, 1965 (Weber State College); Uintah Co., Jensen, EH. Graham 7541 & 7540, April 27, 1933 (CM): Utah Co., Utah Lake, E.M. Christensen s.n., June 24, 1964 (BRY): Washington Co., St. George, ME. Jones s.n., March 30, 1880 (US). Representative Specimens; Populus angustifolia x P. fremontii (previously P. acuminata): Box Elder Co., Raft River Mts., K.S. Erdman 1491, June 8, 1965 (Weber State College): Garfield Co., Henry Mts., Stanton 474, July 15, 1930 (BRY); Iron Co., Cedar Canyon, W.P. Cottam 6736, May 9, 1936 ( UT); Salt Lake Co., S. Flowers 5000, August 8, 1933 (UT): San Juan Co., Natural Bridges Nat'l. Monu.,5.Z,. Welsh & G. Moore 2864, August 12, 1963 (BRY); Sevier Co., BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Salina Canyon, K.S. Erdman 2928, September 7, (Weber State College). 1965 Populus tremuloides Michx. (Fig. 34) Apparently the present climate of Utah is unfavor- able for the sexual reproduction of aspen (Moss 1938, Baker 1925). Althougli some seeds may germinate, the main method of reproduction appears to be root suckering. An aspen stand therefore is usually a clone originating many hundreds of years ago. The vast interconnected root systems lie a few feet under the surface of the ground and at intervals produce sprouts which eventually become independent of the old roots (Gifford 1966, Barnes 1966). Aspen is very intolerant of shade and therefore yields readily to invading conifers such as white fir, Douglas fir, and ponderosa pine. Although aspen communities may be easily replaced by such conifer stands, continued growth of aspen is favored as long as fire is a common component of its environment. Forest fires kill the conifers but the aspen roots remain alive and suckering will produce a new grove after the fire. Because of the effective fire protection measures used today, many aspen communities throughout the state are now being replaced by coni- fers. This species is widespread in Utah as it is through- out temperate North America. Some of the largest specimens of aspen in the United States are located in the Grove of the Aspen Giants in the Manti-La Sal National Forest. There are approximately 20 huge aspen trees with diameters up to three feet. A large UTAH OUTLINE MAP tree of this species, near Cedar City, is 1 1 feet in cir- cumference and 75 feet in height (Littlecott 1969, USDA 1969). Representative Specimens: Beaver Co., Needle Range, S.L. Welsh & G. Moore 3480, August 14, 1964 (BRY); Box Elder Co., Raft River Mts., Preece 903. May 18, 1948 (UT); Cache Co., near Monte Cristo, K.S. Erdman 2599, August 28, 1965 (Weber State College); Garfield Co., Escalante Mts., Beck & Tanner s.n., June 12, 1936 (BRY); Grand Co., Roan Plateau (east) E.H. Graham 9252, June 18. 1935 (CM); Millard Co., Canyon Mts., B.L. Robins 94, August 16, 1913 (USFS);Salt Lake Co., Mill Creek, ,4.0. Garrett 6098, May 8, 1932 (UT); San Juan Co., Abajo Mts., K.S. Erdman 259, July 29, 1960 (BRY); Sevier Co.. Monroe Mt.. D.G. Harrington 16, August 10, 1940 (USPS); Uintah Co., Uinta Mts., L.N. Goodding 1215, June 30, 1902 (US); Washington Co., Silver Reef, Af.f. yo/!« 5169, May 5, 1894 (US). UTAH OUTLINE MAP Fig. 34. Populus tremuloides Michx. Fig. 35. Prosopis glandulosa Tott. Prasopis glandulosa Torr. (Fig. 35) Mesquite is a very common plant in the southwest, and is an aggressive invader of disturbed land. In Texas it has spread over millions of acres of grazing land. In Utah it is restricted to the valley of the Virgin River. Representative Specimens: Washington Co., St. George, W.W. Eggleston 14799, September 7-1 2, 1918 (US); Washing- ton Co., Rockville, K.S. Erdman 63, May 11, 1960 (BRY). Prosopis pubesceiis Benth. (Fig. 36) The Fremont screwbean, so named because of the tiglitly coiled nature of its fruit, is a small shrubby tree of the Virgin River basin in Washington County. It is less common than/", glandulosa. Representative Specimens: Washington Co., Beaver Dam Mts., L.C. Higgins 818, August 19, 1966 (BRY); Washmgton Co., Santa Clara, W.P Cottam 3396, June 21, 1928 (BRY). BIOLOGICAL SERIES. VOL. 11, NO. 3 NATIVE TREES OF UTAH 19 UTAH OUTLINE MAP Fig. 36. Prosopis pubescens Benth. ^mj\ UTAH OUTLINE MAP County linoB find phySiOfjraphiG tk^uturvo Fig. 37. h'liinis lirginiana L. var. mclanocarpa (A. Ncls.) Moldenke Pniniis virginiana L. var. melanocarpa (A. Nels.) Moldenke (Fig. 37) Chokecherry is found throughout Utah's moun- tains in a variety of habitats. Under favorable condi- tions it may form large streamside groves where the trunks are many inches in diameter. However, it is also an early invader of talus slopes where plants, a foot or two in height, may be many years old. Representative Specimens: Beaver Co., Wah Wah Mts., S. Welsh & E.M. Christensen 2692, September 13, 1963 (BRY); Box Elder Co., Raft River Mts., K.S. Erdman \S11 . June 8. 1965 (Weber State College); Duchesne Co., Rock Creek,/ Brotherson 904, June 19, 1965 (BRY); Garfield Co., Bryce Canyon,//. Buchanan 165, July 3, 1957 (UT); Garfield Co., Henry Mts., R. McVaugh 14491, June 7, 1953 (NY); Grand Co., Roan Plateau (east), EM. Graham 9862, July 28, 1935 (CM); Juab Co., Deep Creek Mts., W.P. Cottam 8189, June 16, 1940 (LIT); Piute Co., Tushar Mts.. K.S. Erdman 26, May 9, 1960 (BRY); Rich Co., Bear Lake, L.K. Henrv s.n., June 21, 1956 (CM); San Juan Co., Abajo W.H..P. Rvdherg & .A.O. Garrett 9668, August 1911 (NY): San Pete Co.. Ephraim, /. Tidestrom lA^l. July 14, 1909 (US); Tooele Co., Sheeprock Mts..^ FrischkneduM, iune 1, 1961 (USPS); Wasatch Co., Deer Creek, R.H. Foster 224, October 10, 1964 (BRY); Weber Co., Ogden Canyon, /I. f. Hitchcock 1479, August 19, 1913 (US). UTAH OUTLINE MAP Fig. 38. Quercus gambelii Suti. Quercus ganibelii Nutt. (Fig. 38) Gambel oak forms great belts which extend for hundreds of miles along the foot hills of mountain ranges in Utah. It is a common member of the moun- tain brush zone where it is usually associated with maples, chokecherry, mountain mahogany, and other species. It is generally a small tree, and often little more than a shrub, but under favorable conditions the stems are often over a foot in diameter. The most common method of reproduction in Gambel oak is through root suckering, but there is some reproduction by acorn. Thus large oak clones are common on the mountainsides and may be several 20 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN hundred years in age. The expansion of the clone is slow, perhaps as little as a foot is added to the margin per year. There is some evidence that Gambel oak is far more widespread now than it was a hundred years ago. (Christensen 1950.) The northern-most stands of Gambel oak are in Parry Canyon and near Mantua in southern Box Elder County, and Little Bear Canyon, Cache County. However, it appears that these may represent consid- erable extension of the range in the past hundred years. Ferris, a very careful reporter, indicated that oak was not present in Ogden Hole (Huntsville Valley). Weber County in 1830 but today it is abun- dant on the foothills. (Christensen 1950.) Q. gambelii is generally not a species of the basin and range province in Utah, but it is present in the Mineral, Canyon, Sheeprock, and Oquirrh Mountains. Some of these colonies may be of great age, but Christensen (1950) suggests that others may have resulted from recent introduction of acorns by squir- rels or other animals. Gambel oak has long presented problems to the taxonomist. It is high variable species, which has been alternately considered as one species or treated as several species or subspecific taxa. (Rydberg 1922, Tucker 1961a). Today it is generally regarded as a highly polymorphic species which has hybridized readily with other oaks throughout the southeast. The discovery in 1954(Cottam& Drobnick 1955) of an unusual oak clone in the Oquirrh Mountains of UTAH OUTLINE MAP northern Utah led to the study of its relationship to the Gambel oak with which it was associated to Q. tiirbinella of southern Utah which it resembled. It was determined that the unusual clone was a hybrid between these two oaks, a conclusion full of interest- ing implications inasmuch as one parental species, Q. litrbinella is 300 miles to the south (Cottam & Tucker 1956). The name proposed for this hybrid was Qiiercus x paiiciloba (Tucker 1961a). Further studies revealed similar hybrid clones scat- tered from Ogden south to St. George and Arizona. It was postulated that during the altithermal period fol- lowing the most recent glaciation, temperatures may have been such that Q. tiirbinella grew far north of its present range and hybridized with Gambel oak. If so, some hybrid clones may be several thousand years in age. The distribution map (Fig. 39). for Q. x paiici- loba is based on specimens cited by Tucker (I96lb). U /• ;;;> ;Wi UTAH r^ /; «. ! ^,"'' / OUTLINE MAP : IS C-nD -M County linoB and physiocrat f^ic fcaturvn 1 ^ U A ,-J '-■■■'i'!^ l^ .'•; V JV, -V'"""" \ ^ / i,! ^''=T-pA~S/ , \ c f ^..A^. - Fig. 39. Qucrcus \ pauciloha Rydb. Fig. 40. Qucrcus uudulala Torr. This particular study helped focus renewed atten- tion on the taxonomic problems of a common shrubby oak of southeastern Utah, long identified as Q. iindiilata (Fig. 40). As Tucker pointed out (I'^'hla). the taxonomic history of this species is very confused and troublesome. He proposed that this highly polymorphic group represents hybrids between Q. gambelii and a number of other oaks throughout the southwest. Q. x paiiciloba represented hybrids be- tween Q. gambelii and Q. tiirbinella and these were most common along the western limits of Gambel oak's range. On the other hand, there was little evi- BIOLOGICAL SERIES. VOL. 1 1 , NO. 3 NATIVE TREES OF UTAH 21 dence that Q. turbinella was a parent of the sprawling oak of southeastern Utah. Tucker suggested that Q. liavardii, a low, rhizomatous shrub of sandy areas in Texas and New Mexico, had crossed with Gambel oak to produce what has long been called Q. undulata in Utah. (Tucker 1961a, 1961b.) Representative Specimens; Qiiercus gambelii: Bo.\ Elder Co., Parry Canyon, E.M. Clirislensen s.n., July 20, 1948 (LIT); Cache Co., Little Bear Canyon, Bancroft 'l031, June, 1928 (UT); Carbon Co., Book Cliffs, E.H. Graham 83.^9a, June 28, 1933 (CM); Duchesne Co.. Red Creek,/. Brotherson 507, June 21, 1965 (BRY); Garfield Co., Aquarius Plateau, Beck & Tanner s.n., June 23, 1936 (BRY);Grand Co., La Sal Mts., K.S. Erdnian 197, July 27, 1960 (BRY); Kane Co., Kanab, B.F. Harrison 11087, August 27, 1946 (BRY); Salt Lake Co., Emigration Canyon. Vickerv 2516, August 30, 1959 (Utah); San Juan Co., Abajo Mts.,' W.P. Cottam 14392, July 11, 1956 (UT); San Juan Co., Aztec Creek, /.(//t/sffv 154, July 25. 1958 (UT): San Pete Co., Mt. Pleasant,/. Tidcstrom 1867, September 2, 1908 (US); Sevier Co., Salma Canyon, M.E. Jones 5447, June 16, 1894 (US); Tooele Co., Shecprock Mts., W.P. Cottam 14340, July 1, 1956 (UT); Uintah Co.. Roan Plateau (east), E.H. Graham 9825, July 26, 1935 (CM); Washington Co., Beaver Dam Mts., L C. Higgins 663, May 27. 1966 (BRY). Representative Specimens: Quercus x pauciloba (taken from Tucker 1961b): Beaver Co., Mineral MU.^Drohnick 86 (UT); Iron Co., four mi. NW of Kanarravillc, 6,000 ft.. Cot- ram . Youngberg eU, October 2, 1965 (BRY). Salix lasiolepis Benth. (Fig. 48) This willow grows along streams and in wet areas in Utah, where it is of infrequent occurrence. Representative Specimens: Iron Co., Parowan, W.W. Eggle- ston 14903, September 17, 1918 (US); Millard Co., Canyon Mts., K.S. Erdman 134, May 13, 1960 (BRY); Sevier Co., Monroe Mt., K.S. Erdman 5, May 9. 1960 (BRY); Washing- ton Co.. Pine Valley, F.W. Gould 1781, May 26, 1942 (NY). Salix nigra Marsh (Fig. 49) Western black willow is restricted to streamsides in Washington County and the Colorado River drainage in eastern Utah. It extends as far north as Moab in Grand County. Representative Specimens: Garfield Co., Bullfrog Creek, 5. Welsh 3969, May 6, 1965 (BRY); Grand Co., Arches Nat'l. Monu., S. Welsh 2820, Mav 30, 1965 (BRY); Washington Co., St. George, W.W. Eggleston 14773, September 7-12, 1918 (US). 24 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN iferous or aspen forests. Its altitudinal range is from 5,000 to 1 1,000 feet elevation. UTAH OUTLINE MAP Fig. 48. Salix lasiolcpis Benth. UTAH OUTLINE MAP County liUBB and physiogrciphic featurcB Fig. 49. Salix nigra Marsh Salix scouleriana Barratt (Fig. 50) Scouler or fire willow, like Bebb willow, is wide- spread througliout Utah and not limited to stream- sides. It is a common understory tree or shrub in con- Fig. 50. Salix sioiilcnaiw Barratt Representative Specimens; Box Elder Co.. Raft River Mts., W.P. Cottam 2949, June 5, 1928 (BRY); Cache Co., Lo- gan Canyon, C.R. Ball 1866, August 18, 1914 (US); Daggett Co.. Red Canyon, L. Williams 593, June 10, 1932 (NY): Davis Co., Farmington Canyon, A.D. Youngbcrg 527, Sep- tember 3, 1965 (BRY); Garfield Co., Henry Mts., Stanton 188, June 29, 1930 (UT); Grand Co., La Sal Mts., K.S. Erd- man 211, July 27, 1960 (BRY); Juab Co., Deep Creek Mts., B. Maguire 2533a, June 20, 1933 (UTC); San Juan Co., Abajo Mts., B. Magiitre 1722a, July 1, 1932 (UTC); San Pete Co., Mt. Pleasant, /. Tidestrom 1912, September 4, 1908 (US); Tooele Co., Stansbury Mts., E.M. Cliristensen s.n., July 29, 1963 (BRY); Utah Co., Provo Canyon, S. Welsh 3385, August 5, 1964 (BRY): Waslungton Co., Pine Valley Mts.,B. Maguire 20184, August 1 , 1934 (UTC). Sambiicus caenilca Raf. (Fig. 5 1 ) This elderberry is common throughout most of Utah's mountains. This shrubby plant is common among stands of ponderosa pine, Douglas fir, and as- pen, especially in more open areas near streams. Representative Specuiiens: Beaver Co., Needle Range, S.L. Welsh & G. Moore 3202a, July 18, 1964 (BRY); Carbon Co., Roan Plateau (west), S.L. Welsh & E.M. Christcnsen 6564, August 11, 1967 (BRY); Duchesne Co., Moon Lake, E.H. Graham 6410, June 30, 1931 (CM):Garfield Co., Bryce Can- yon, // Buchanan 183, July 9, 1957 (UT): Grand Co., B. Maguire 21716, July 15, 1933 (NY): Salt Lake Co., Emigra- tion Canyon, \iekery 2514, August 30, 1959 (UT);San Pete Co., Ephraim Canyon, /. Tidestrom 138, August 12, 1907 (US); Sevier Co., Tushar Mts., R.G. Warnock s.n., July 7, 1961 (BRY); Tooele Co., Stansbury Island, W.P. Cottam 9174, July 7, 1942 (UT): Uintah Co., Dinosaur Nat'l. Monu., AH. Holme^ren 14248, June 29, 1962 (NY): Utah Co., Provo Canyon, St.. Welsh 3243, June 1964 (BRY); Washington Co., Bellvue, \. Tidestrom 9412, May 10, 1919 (USl-S): We- ber Co., North Fork, Call 89, July 26, 1940 (UT). BIOLOGICAL SERIES. VOL. 1 I , NO. 3 NATIVE TREES OF UTAH 25 UTAH OUTLINE MAP Fig. 51. Sambuciis cacruica Rat. Fig. 53. Sorbus scopulina Greene UTAH OUTLINE MAP County linoB find physiograp'iic taaturve i 10 »0 .„...-.i,:-^^ ' .., ^Ili§i0W^^'^' ^''^i0^^m^^^ t'^k^.^rw^ - — r ^ ^M Fig. 1. Typical sagebrush-grass community comprized of Artemisia tridentata, Stipa comata. and Oryzopsis hyinenoides lig. 2. Sinks area luukiiig east. BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION ~ \R31E I R32E LEGEND Paved Roods oS»ateHwy. vUSHwy Unsurfoced Roads * NRTS Instalations NRTS Boundry ■ Spring or well • Towns T 4 N tra \3\ ^/ r-- i"' ,&/ 1 ---^-/s ■ <$y 1 0/ ~/ ' -?' / ^^ , / / V--- /"- Vn ~ / v ~., ~~^-% T3' ■-/ ^ I •P' 01, o ,' D \ /'""~~/- Atq9e__ _|^r_d. - -n ^-- " , ebr-1 middle ---,-'' butte (:r}\ ' "east butte R28E frenchman well - l'R29E X j:f / "= ^ Springs ^ big" ■"-' southern butte IR30E Atomi cI^ieX'r R32E T 2 N T 3 N Fig. 3. Principal roads used to make plant collections. Modified from Allied (1968). BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Fig. 4. Big Lost River during spring run off. across the northeastern end of the Snake River Plain (presently the northeastern portion of the NRTS), and camped on the Camas River (June 20-23). From this camp Wyeth penned a number of letters, one to his friend Nuttall in Cambridge, mentioning a package of plants being shipped which, however, never reached its destination. It is interesting that this is the earhest account of botanical collecting in the intermountain region. The next collections were those of Nuttall and Townsend in their travels to the Columbia River with Wyeth's second transcontinental expedition. They were in the area of the Lost River and Camas prairie from August 7th to August 12, 1834. To my knowledge. Palmer was the next botanist to collect plants in the vicinity of the site and was there around 1893 (Davis, 1968 personal communication and Reveal, 1969 personal communication). Since 1900 the most important collections have been made by R. J. Davis who collected in this area periodically from the late 20's untill about 1952. The Atomic Energy Commission supported workers who made a substantial contribution in 19.50 when they collected a series of plants (about 180) during the spring of that year. Many of these are deposited in the herbarium of Idaho State College. Some of these plants are also present in a small collection in the ecology branch of the NRTS. Geographical description of the National Reactor Testing Station The National Reactor Testing Station is situated along the western edge of the upper Snake River Plain in southeastern Idaho. Most of the test site lies in the eastern portion of Butte County, at the base of the Lost River, Lehmi, and Beaverhead Mountains; the remainder of the site occupies portions of Bingham, Jefferson, Clark, and Bonneville Counties. The 894 square miles which comprise the station is more or less rolling to broken land, typical of this portion of the Columbia Plateaus Province. The plain is broken by three large buttes, all volcanic in nature. Middle and East Buttes, which are located in the south- eastern portion of the test site, rise from the plain at an elevation of 5,200 feet, to elevations of 6,394 feet and 6,605 feet respectively. Big Butte is located just south of the NRTS and towers to an elevation of 7,576 feet. Little Lost River and Birch and Camas Creeks enter the station from the north and drain into the sinks in the northwestern portion of the site. Geology The Snake River Plain was formed by the inter- bedding of volcanic rocks and of lake and alluvial deposits. The alluvial deposits are quaternary in age BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION BLAINE y^*Blackfoot S^f Fig. 5. Location of National Reactor Testing Station in relationsiiip to some cities in southeastern Idaho. Modified from Allred(1968). BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN i y^-mt* '•M. ■u:.^ -.»t,.- t :B lig. 6. Lemhi Mountains in northwestern portion of National Reactor Testing Station. ¥:\ I'lg. 7. Middle and ha.sl liiau ,>. iih wild ijc paiLli ui kacgiuuiid. BIOLOGICAL SERIES, VOL. I I NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION NRTS VEGETATION TYPE MAP LEGEND o NRTS Boundry • Highway Intermittent Streams Instclotions Big Sagebrush- Bluebunch Wheotgross-Rabbitbrush Big Sagebrush - Rabbitbrush - Foxtail Big Sagebrush -ThickspikeVi'heatgrass-Needlegrass Big Sagebrush -Winterfat- Rabbitbrush Big Sagebrush -Winterfat -Saltbush Big Sagebrush -Shadscale-Rabbitbrush Big Sagebrush - Ricegrass-Needlegrass Black Sagebrush-Big Sagebrush -Shadscale Small Sagebrush -Shadscale -Foxta Crested Wheatgross (seeded) Bluebunch Wheatgross -Threetip Sage- Rabbitbrush Bluestem Wheatgross- Wiregross -Iva Ricegross - Needlegross -Rabbitbrush -Cactus Canada Wild Rye- Rabbitbrush -Big Sagebrush Juniper - Big Sogebrush - Bluebunch Wheatgross Horsebrush -Rabbitbrush -Big Sagebrush Rabbitbrush -Big Sagebrush- Gross Saltbush- Winterfat- Indian Ricegross Mixed Shrub Sond Dunes °Ar, Idaho Falls ■A. IE -Oc^^ R32E Fig. 8. Major plant communities as designated by McBride. Modified from McBride (1968). BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN and are present along the streams flowing onto the site. The northern part of the station, south and west of Mud Lake, is covered' by lake and eolian deposits; these are mostly of unconsolidated clay, silt, and sand and are represented as far west as the sinks. Ex- posures of basalt are less common within this area. East, Middle, and Big Butte are probably sources of much of the lava flow. Middle Butte is composed entirely of basalt. Big Butte is mostly rhyolite with some basalt, and East Butte consists of trachyte and rhyolite with some basalt, and East Butte consists of trachyte and rhyolitic rocks. All three Buttes are middle teritary in age, somewhat older than the quat- ernary basalt flows surrounding them. Many of the basalt flows are in excess of 1 ,500 feet in thickness. Ecology The NRTS is located at the eastern end of the Snake River Plain in a broad, more or less tlat valley. The prevailing air masses pass over the mountain ranges to the west and north before entering the site, and in doing so lose the major portion of their mois- ture. Annual precipitation at the station is between seven and eight inches. Temperatures vary from 102 degrees F as a high and minus 43 degrees F as a low. The average annual temperature is 42 degrees F. The intense cold common in most of that region east of the Rocky Mountains does not generally prevail in the Snake River Plain, due to the effective barrier of the Beaverhead and Centennial Mountains. The vegetation of this semidesert belongs to the northern desert shrub biome. According to French, et al {\965). Artemisia tridentata occupies about 80 per- cent of the site. McBride (1968), in his vegetation map (Fig. 1 ) of the NRTS, recognized 19 major plant communities. Allred (1968), in his ecological inves- tigations, established study areas in the major vegeta- tion types as outlined by McBride. Detailed plant analysis within Allred's study areas demonstrated some variation from McBride's designations. Poorly drained, saline soil depressions support vari- ous members of the family Chenopodiaceae, such as Sarcobatiis vermiculatus. Grayia spinosa, Eurotia lan- ata, and several Atriplex species. Jiinipenis osteosper- ma is dominant on rocky ridges extending onto the site. Sandy areas, particularly south of Mud Lake, support stands of grasses of wliich Oryzopsis hymen- oides. Stipa comata, and Aristida fendleriana are dominant. Species of Chrysothamnus are also an important part of the vegetation in sandy areas. Among the grasses, Agropyron spicatum appears to be the most important single species, althougli Poa secunda, Koeleria cristata, Stipa spp., Agropyron dasystachyitm, Oryzopsis hymenoides, and others, are common. The streams which enter the site support various species of trees, shrubs, grasses and forbs. . iini.iii iniiKilii^nfiimfniiii'ii iiiirii-iia&rrrirT-'i-^'ii ^K I ig. ''. I )r\zi>p\i\ hvniciii)ult\-Slipa coniaUi gi.iss mini lui nil \ in I ijctoi 1 Lit on noillieast portion ot site. BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION ^^M Fig. 10. Stand of juniper along Idaho liiglnvay 21. live miles north of junction with Idaho highway '< fe -■-^ iSfe-- •^'fi^^ »(-F ^!*K' Fig. 11. Artemisia-Atriplex community three miles north of Pole Line Road. Lemhi Mountains in background. ■■i-f^ BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 1 W^'Mr Fig. 12. Mixed comiiuinity oi Chrysothamnus. grass, Tetradymia. and Opuntia. jiA-^ !."'■ i^?':; 'i. rJj ..' .-^^ :m^^' -M Fig. 13. Sand dunes witli Russian tlustie in loreground. Located just norllnvcst ot junction witli Idaho iiigliway X8- Lincoln Blvd. Arrangement of Plant Families, Genera, and Species All taxa from division to species are arranged in made for all families, genera, and species. In addition, alphabetical order. The primary purpose of this work distribution, phenology, and a voucher specimen cita- is to provide a means of identifying the plants of the tion is given for each of the species. National Reactor Testing Station. Keys have been BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION TAXONOMIC TREATMENT 11 Key to the Higher Taxa 1. Plants reproducing by seeds; cone or flower bearing plants DIVISION SPERMATOPHYTA (2) 1. Plants without seeds or tlowers, reproducing by spores in sporangia; fernlike or rushlike plants .... DIVISION PTERIDOPHYTA, p. 44 2. Seeds borne in a closed carpel of a flower; stigmas present; plants mostly deciduous . . . CLASS ANGIOSPERMAE (3) 2. Seeds borne on the surface of scales or bracts, arranged in cones; stigmas none; plants evergreen trees CLASS GYMNOSPERMAE, p. 43 3. Embryo usually with 1 cotyledon; floral parts usually in threes; leaves mostly parallel veined . . SUBCLASS MONOCOTYLEDONEAE, p. 39 3. Embryo usually with 2 cotyledons; tloral parts commonly in fours or fives; leaves mostly net-veined SUBCLASS DICOTYLEDONEAE (4) 4. Corolla lacking; calyx, when present, with segments much alike, sometimes petaloid . . . SERIES APETALAE, p. 1 1 4. Corolla and caly.x present, commonly differ- ent in texture and color 5 5. Petals more or less united SERIES GAMOPETALAE, p. 12 5. Petals distinct, at least at the base SERIES POLYPETALAE. p. 1 2 CLASS ANGIOSPERMAE SUBCLASS DICOTYLEDONEAE SERIES APETALAE 1 . Plants woody, trees or shrubs 2 1 . Plants herbaceous, sometimes woody only at the base 8 2. Flowers borne in catkins SALICACEAE, p. 35 2. Flowers not in catkins, perfect or unisexual . 3 3. Flowers several to many, in a head surround- ed by an involucre of bracts; calyx wanting; ovary inferior COMPOSITAE, p. 17 3. Flowers 1 to many; calyx usually present . .4 4. Ovary inferior; flowers in corymbose or ter- minal cymes, perfect; stamens usually 5 . . . . CORNACEAE, p. 23 4. Ovary superior; trees or shrubs 5 5. Ovary of I carpel, with 1 locule, 1 style, 1 stigma ROSACEAE, p. 34 5. Ovary of 2 or more carpels, several loculed . 6 6. Leaves opposite, lobed to compound; fruit a double samara ACERACEAE, p. 14 6. Leaves alternate or opposite, entire to tooth- ed, but not compound; fruit not a double samara, various 7 7. Flowers subtended by a gamophyllous invol- ucre, perfect; stamens mostly 6 to '^; fruit an achene POLYGONACEAE, p. 32 7. Flowers perfect or unisexual, but not sub- tended by a gamophyllous involucre; sta- mens I to 5, fruit a utricle CHENOPODIACEAE, p. 16 8. Ovary inferior, adnate to the perianth or hypanthium, only appearing fused in Nycta- ginaceae 9 8. Ovary superior 12 9. Ovary 2-loculed, 1 ovule in each locule; fruit 2-seeded 10 9. Ovary 1 -loculed, 1-2 ovules per locule; fruit 1 -seeded II 10. Perianth parts united; flowers in cymes, not umbellate; leaves opposite or whorled RUBIACEAE, p. 35 10. Perianth parts separate; flowers in umbels; leaves basal or alternate UMBELLIFERAE, p. 38 1 1 . Plants parasitic on the roots of shrubs; per- ennial herbs with woody base; leaves alter- nate; flowers greenish white SANTALACEAE, p. 36 11. Plants not parasitic; leaves opposite; inflor- escence rarely corymbose; ovules I to an ovary NYCTAGINACEAE, p. 30 12. Pistils numerous in a single flower; stamens commonly many RANUNCULACEAE, p. 33 12. Pistils I to a tlower; stamens 1 to many, usu- ally 1 0 or less 13 13. Ovary 2 or more loculed; milky juice present EUPHORBIACEAE, p. 26 13. Ovaiy 1-loculed; plants without milky juice . 14 14. Ovary with several to many ovules; fruit a several- to many-seeded capsule; styles 2 to 5; calyx of distinct segments CARYOPHYLLACEAE, p. 15 14. Ovary with a single ovule; fruit a 1 -seeded achene or utricle 15 15. Leaves with stipules united to form a sheath around the stem, above the nodes POLYGONACEAE, p. 32 15. Stipules lacking; leaves alternate or opposite 16 i; BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 16. Perianth large, over 5 mm long; style 1, stig- ma 1 ; leaves opposite and entire NYCTAGINACEAE, p. 30 16. Perianth mostly less than 5 mm long; styles or stigmas more than I (usually) ...... .17 17. Fruit an achene; plants with stinging hairs .. URTICACEAE, p. 38 17. Fruit a utricle; plants without stinging hairs . 18 18. Bracts and perianth more or less hyaline or scarious; leaves alternate; plants weedy an- nuals AMARANTHACEAE, p. 14 18. Bracts and perianth herbaceous-tleshy; plants annual or perennial CHENOPODIACEAE, p. 16 SERIES GAMOPETALAE Ovary superior 2 Ovary inferior 15 Stamens free from the corolla 3 Stamens at least partially adnate to the cor- olla 5 3. Corolla regular; stamens many, united into a tube (monodelphous); leaves alternate, simple MALVACEAE, p. 29 3. Corolla irregular; stamens 4 to 10, diadelphous 4 4. Sepals 2, scalelike; petals 4, in 2 sets; leaves finely dissected FUMARIACEAE, p. 26 4. Sepals 4 to 5, not scalelike; petals 5, papilio- naceous LEGUMINOSAE, p. 27 5. Plants devoid of chlorophyll; parasitic on other plants OROBANCHACEAE, p. 30 5. Plants with green chlorophyll; plants not parasitic 6 6. Ovaries 2, entirely separate below and with a common stigma above; plants with milky juice ASCLEPIADACEAE, p. 14 6. Ovary 1; plants without milky juice 7 7. Corolla regular or nearly so; stamens mostly 5 (2 or 4) 8 7. Corolla irregular 13 8. Ovary 4-loculed, 4-lobed, developing into nutlets; inflorescence a scorpioid cyme .... BORAGINACEAE, p. 14 8. OVARY not developing into nutlets; fruit a capsule or berry; I to 3-loculed 9 9. Ovary 1-loculed; leaves usually alternate . . . 10 9. Ovary with 2 or more locules 11 10. Corolla dry -scarious, veinless; scapose herbs; sepals and petals 4; stamens 2 or 4; flowers in bracteate spikes or heads PLANT AGINACEAE, p. 31 10. Corolla not as above; plants commonly leafy stemmed; perianth 5-merous (in ours); sta- mens 5 HYDROPHYLLACEAE, p. 26 1 1. Style 3-cleft; ovary 3-loculed POLEMONIACEAE, p. 31 1 1. Style 2-cleft; ovary commonly 2-loculed. .12 1 2. Stems trailing or twining CONVOVULACEAE, p. 23 12. Stems not trailing or twining; style I ; stigma 1 , entire or 2-lobed; fruit a capsule or berry . SOLANACEAE, p. 38 13. Fruit of 2 to 4 nutlets; leaves opposite . . .14 13. Fruit a capsule; ovary 2-loculed; leaves most- ly opposite; corolla sometimes almost regular SCROPHULARIACEAE, p. 36 14. Ovary 4-lobed; style arising in the center between the lobes LABIATAE, p. 27 14. Ovary not 4-lobed; style apical VERBENACEAE, p. 38 15. Stamens numerous; anther sacs opening by slits; petals numerous; plants spiny and suc- culent CACTACEAE, p. j 5 15. Stamens 5 or less and not with the above combination of characters 16 Stamens distinct; leaves whorled or opposite 16 17 16. Stamens united by their anthers into a ring or tube around the style; flowers in invol- ucral heads COMPOSITAE, p. 17 17. Fruit 2-lobed, 2-loculed, separating into 2 one-seeded nutlets; leaves more or less whorled RUBIACEAE, p. 35 Ovary 2 to 5-loculed; fruit a berrylike drupe; ours shrubs CAPRIFOLIACEAE, p. 15 SERIES POLYPETALAE 1 . Ovary inferior 2 1 . Ovary completely superior 13 2. Petals many; stems thick and succulent, spiny CACTACEAE, p. 15 2. Petals less than 10; stems not succulent or only slightly so, not spiny (see also Rosa- ceae) 3 3. Stamens as many as the petals and opposite them; plants shrubby with thick, leathery leaves RHAMNACEAE, p. 34 3. Stamens fewer or more than the petals, or if the same number than alternate with them . 4 4. Plants woody, trees and shrubs 5 4. Plants herbaceous or woody at the base . . .8 5. Fruit a 1- to 2-seeded drupe; flowers many in a flat-topped cyme . . . .CORNACEAE, p. 23 Fruit not a drupe, various, 3 or more seeded; flowers in racemes or corymbs 6 Leaves opposite; fruit a capsule SAXIFRAGACEAE, p. 36 Leaves alternate; fruit a pome or berry ... .7 Fruit a pome; leaves pinnately veined, lobed or compound; petals commonly much 5 BIOLOGICAL SERIES, VOL. I 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 13 larger than the petals ROSACEAE, p. 34 7. Fruit a berry; leaves palmately veined; sim- ple; calyx larger and usually more showy than the petals . . GROSSULARIACEAE, p. 26 8. Stamens 10, 11 or more; plants rough-hairy; fruit a capsule LOASACEAE, p. 29 8. Stamens 10 or fewer; plants not rough-hairy; fruit various 9 9. Ovules and seeds more than 1 in each locule; ovary 1 to 4-loculed 10 9. Ovules and seeds 1 to each locule; ovary 2 to 6-loculed 11 10. Style 1 ; stamens 4 or 8 ONAGRACEAE, p. 30 10. Styles 2 or more; stamens as many as the petals, twice as many or numerous SAXIFRAGACEAE, p. 36 11. Leaves verticillate, entire; ovary commonly 2-loculed; style 1 CORNACEAE, p. 23 11. Leaves never verticillate; ovary 4 or 2-loc- uled and then with 2 styles present 12 12. Stamens 2, 4, or 8; petals 2 or 4-merous; style 1 ; ovary 4-loculed ONAGRACEAE, p. 30 12. Stamens 5; petals 5-merous (rarely 4); ovary with 2 or more styles, 2-loculed; fruit dry, 2-seeded UMBELLIFERAE, p.38 13. Corolla definitely irregular 14 13. Corolla regular or nearly so 20 14. Leaves variously compound 15 14. Leaves simple, entire to lobed or pinnatified, not compound 18 15. Petals smaller than the sepals; stamens num- erous; ovaries commonly 2 or more or if car- pels these united at the base RANUNCULACEAE, p. 33 15. Stamens less than 14; petals as large or larger than the sepals; ovary 1, if merely carpelate then these completely united 16 16. Stamens 6; calyx segments 2 or 4; corolla not papilionaceous 17 16. Stamens commonly 10; calyx of 5 usually united sepals; corolla mostly papilionaceous LEGUMINOSAE, p. 27 17. Sepals 2; corolla spurred FUMARIACEAE, p 17. Sepals 4; corolla not spurred CAPPARIDACEAE, p. 18. Stamens many; leaves palmately cleft or parted; carpels commonly more than 1; fruit a follicle RANUNCULACEAE, p. 33 18. Stamens 10 or less; fruit a capsule or legume; leaves simple to compound, but not palmately cleft 19 19. Stamens 5; ovary 1-loculed with 3 placentae; lower petal saccate at the base or spurred . . . V10LACEAE,p. 39 19. Stamens 10; petals longer than the sepals; ■>■ 26 15 20. 20. 21. 21. 22. 22. 23. 23. 24. 24. 25. 25. 26. 26. 27. 27. 28. 29. 29. 30. 30. 31. 31. 32. 32. fruit a legume LEGUMINOSAE, p. 27 Stamens the same number as the petals and opposite them 21 Stamens fewer or more than the petals, or if the same number then alternate with them . 22 Leaves simple and pinnately veined, not lobed, alternate; petals 5, hooded and clawed RHAMNACEAE, p. 34 Leaves palmately veined and lobed, oppo- site; petals 4 or 5, not hooded and clawed . . ACERACEAE, p. 14 Ovary 1, 1-loculed, at least above 23 Ovary more than 1, or if 1 then 2 or more loculed 25 Ovary simple; style I ; stigma 1 ; placenta 1 . . 24 Ovary compound with more than 1 placenta, style or stigma; plants woody shrubs; leaves alternate ANACARDIACEAE, p. 14 Stamens and petals inserted on the calyx tube; leaves various, but not bipinnate ROSACEAE, p. Stamens and petals not inserted on the calyx tube; leaves commonly bipinnate LEGUMINOSAE, p. Stamens numerous, more than 12 26 Stamens not numerous, 1 2 or less 28 Stamens monodelphous MALVACEAE, p. Stamens distinct or in several sets 27 Stamens and petals inserted on the calyx tube; flowers perigynous ROSACEAE, p. Stamens and petals inserted on the recep- tacle; flowers hypogynous RANUNCULACEAE, p. 33 Ovaries 2, united at the tips by a discoid stig- ma head; plants with milky juice ASCLEPIADACEAE, p. 14 Ovaries 1 to many, if united then only at the base; juice rarely milky, if so then not with the foregoing characters 29 Ovaries more than 1, separate or united at the base, each with a style 30 Ovary 1 ; style 1 per ovary 31 Leaves fleshy, entire; carpels 3 to 5 CRASSULACEAE, p. 23 Leaves not fleshy; carpels 2 SAXIFRAGACEAE, p. 36 Ovules 1 to 2 in each cell 32 Ovules more than 2 in each cell 35 Flowers imperfect, borne in calyx-like invol- ucres; plants with milky juice EUPHORBIACEAE, p. 26 Flowers perfect, not borne in calyx-like involucres; plants without milky juice . . .33 34 27 29 34 14 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 33. Perianth 4-merous; stamens 6 (4+2) CRUCIFERAE, p. 23 33. Perianth 5-merous; stamens 5 or 10 34 34. Stamens 5: leaves alternate . .LINACEAE, p. 29 34. Stamens 10; leaves commonly opposite .... GERANIACEAE, p. 26 35. Styles 2, distinct; ovary 2-celled; fruit a cap- sule or berry SAXIFRAGACEAE, p. 36 35. Style 1, distinct; ovary 2-celled; fruit a silicle or silique CRUCIFERAE, p. 23 ACERACEAE. MAPLE FAMILY 1. Acer L. Maple L Acer glabmm Torr. Commonly of the moun- tains, April to June. Idaho, west from Alaska to Cal- ifornia, east from Alberta to New Mexico and Ne- braska. Webb Springs, Butte Co., 26 May 1967,874. AMARANTHACEAE. AMARANTH FAMILY 1. Amaranthus L. Pigweed; Amaranth 1. Amaranthus hybridus L. A weed of waste places and cultivated ground. Widely distributed throughout most of North America, June to Novem- ber. T.6N., R.3 IE., Butte Co., 29 July 1967, 1182. ANACARDIACEAE. SUMAC FAMILY I. Rhus L. Sumac 1. Rhus trilobata Nutt. Common along streams and dry hills. May to July. North Dakota to Washing- ton, south to Texas and California. Big Lost River, Butte Co., 1 July 1967, 1093. ASCLEPIADACEAE. MILKWEED FAMILY 1. Asclepias L. Milkweed 1. Asclepias speciosa Torr. Common over much of the western and central states, June to August. South of junction 88-Lincoln Blvd., Butte Co., 24 August 1967, 1209. BETULACEAE. BIRCH FAMILY 1. Betula L. Birch 1. Betula occidentalis Hook. Along streams of the plains and mountains. Utah, west to northern Cal- ifornia, less frequent to southern British Columbia and south central Idaho. Birch Creek, Clark Co., 25 June 1967, 1053. BORGINACEAE. BORAGE FAMILY 1. Prostrate annuals; style deeply 2-cleft3. Coldenia 1. Plants mostly erect or the style entire or shallowly lobed 2 2. Nutlets with prickles on the margins, attached at or below the middle 3 2. Nutlets without prickles 4 3. Annual plants; pedicles erect; style longer than the nutlets 6. Lappula 3. Plants perennial; pedicles reflexed; styles com- monly shorter than the nutlets 5. Hackelia 4. Calyx enlarged in fruit, veiny, folded, and flat- tened; stems procumbent 2. Asperugo 4. Plants not as above 5 5. Flowers blue or reddish in bud; nutlets obliquely attached 8. Mertensia 5. Flowers other than blue 6 6. Stigmas 2; nutlets broadly attached at the base to a flat receptacle 7. Lithospermum 6. Stigmas solitary; nutlets attached a little above the base 7 7. Corolla white (ours); fornices present in the throat and the throat almost closed by them 4. Cryptantha 7. Corolla yellow or orange; fornices wanting and the throat open 1 . Amsinckia 1 . Amsinckia Lehm. Fiddle-neck 1. Amsinckia nienziesii (Lehm.) N. & M. In culti- vated fields in dry or moist places. May to September. Idaho to British Columbia, south to California, and Nevada. R.29E., T.5N., Butte Co., 25 June 1967, 957. Birch Creek, Clark Co., 25 June 1967; 1049. 2. Asperugo L. Madwort 1. Asperugo procumbens L. A weedy plant of fields and waste places. May to July. Over much of North America; Eurasia. R.29E., T.5N., Butte Co., 12 June 1967,958. 3. Coldenia L. 1. Coldenia nuttallii Hook. Sandy soil of the plains. May to August. Idaho, west from Washington to California, east to Wyoming, south to Arizona. Tractor Flat; Jefferson Co., 22 July 1967, 1199. R.31E., T.6N.. Butte Co., 18 July, 1967, 1 166. 4. Cryptantha Lehm. 1 . Plants annual 3 1 . Plants perennial 2 2. Leaves strongly ciliate margined, setose; nutlets ovate, muricate or short rugose, the scar open and triangular 3. C. humilis 2. Leaves not strongly ciliate margined, setose and subtoinentose or tomentose; nutlets lanceolate, rugose, the scar closed or subulate 5 . C spicuUfera 3. Calyx circumscissile, plants low mat forming; flowers solitary in the upper axils I . C circumscissa 3. Calyx not circumscissile; plants erect, not form- ing mats; inflorescence spikelike 4 4. Nutlets heteromorphous, three of them tuberu- late, the others smooth, 1 .8-2.3 mm long 4. C. kelseyana 4. Nutlets homomorphous, all smooth, lanceolate, 1 .5-2 mm long 5 5. Margins of the nutlets prominent and sharply an- gled, at least above; fruiting calyx 2-4.5 mm long; BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 15 corolla 1 to 2 mm wide 6. C. watsonii 5. Margins of the nutlets obtuse or rounded; fruit- ing calyx 4 to 6 mm long; corolla about 1 mm wide 2. C. fendleri 1. Cryptantha circumscissa (H. & A.) Johnst. Dry, sandy soil, April to July. Washington to California, east through the Snake River Plains of Idaho, south to Utah, Arizona; Chile and Argentina. R.34E., T.5N., Jefferson Co., 20 June 1967, 1005. 2. Cryptantha fendleri (Gray) Greene. Mostly in sandy soil, May to July. Saskatchewan to Washington, east and south to Idaho to Nebraska, Nevada and New Mexico. Tractor Flat, Jefferson Co., 22 July 1967, 1195. 3. Cryptantha humilus (Greene) Pays. var. shantzii (Tidestr.) Higgins. Idaho, south to Colorado, west to Utah to California. T.4N., R.28E., Butte Co., 798. 4. Cryptantha kelseyana Greene Dry, sandy soil of the plains, June to July. Saskatchewan and Montana, south to Idaho and Utali. Tractor Flat, Jefferson Co., 20 June 1967,991. 5. Cryptantha spiculifera Pays. Foothills to the plains, May to July. Idaho, west to Washington and Oregon, north to Montana and Saskatchewan. East of Arco, Butte Co., 17 May 1936, Twitchell and Shaw, s.n. (IDS). East of Arco, 22 June 1939, Davis, 937 (IDS). 6. Cryptantha watsonii (Gray) Greene. Foothills and lower valleys. May to August. Western Montana to central Washington, south to Nevada and Colo- rado. Big Butte, Butte Co., 22 June 1950, AEC, 91 (IDS). 1 . Lappula echinata Gilib. Mostly weedy plants of waste places, roadsides, and cultivated ground, June to August. Birch Creek, Clark Co., 25 June 1967, 1046. R.29E., T.6N., Butte Co., 10 June 1967, 941. R.34E., T.5N., Jefferson Co., 20 June 1967, 990. 2. Lappula redowskii (Hornem.) Greene. Weedy plants. May to July. Common in western North America; Eurasia. T.4N., R.31E., Butte Co., 20 May 1967,782. 7. Lithospermum L. Gromwell 1. Lithospermum niderale Lehm. Dry or quite moist areas in the plains and foothills, April to June. Alberta and British Columbia to California and Colo- rado. Webb Springs, Butte Co., 24 May 1967, 843. 8. Mertensia Roth. Bluebells 1. Mertensia oblongifolia (Nutt.) G. Don Sage- brush plains to open slopes, April to July. Montana, south to Utah and Nevada. East Butte, Bingham Co., 18 May 1967,772. CACTACEAE. CACTUS FAMILY I. Opuntia Mill. Prickly-pear; Cactus I. Opuntia polyacantha Haw. In sagebrush flats, May to June. British Columbia to Alberta, south to Oregon, Arizona to Texas. Webb Springs; Butte Co., 1 July 1967, 1091a. CAPPARIDACEAE. CAPER FAMILY 1. Cleome L. Bee Plant 1. Cleome lutea Hook. In desert plains, usually in sandy soil. May to July. Washington to California, 5. Hackeha Opiz. Stickseed east to Montana, Nebraska and Texas. T.6N., R.30E., 1. Corolla limb blue; stems spreading-hirsute at least Butte Co., 6 June 1967,898. above; cauline leaves well developed; fornices mi- nutely papillate 1. //. jessicae CAPRIFOLIACEAE. HONEYSUCKLE FAMILY 1. Corolla limb white to ochroleucous, usually 1. Symphoricarpus Duhamel. Snowberry marked with light blue; stems with appressed 1- Corolla 7 to 10 mm long, long and narrow; fruits pubescence above; cauline leaves evidently 7 to 10 mm long 2. S. oreophilus smaller; fornices villous-puberulent 1- Corolla 5 to 7 mm long, short and broad; fruits 2. H. patens 1 0 to 15 mm long \. S. albus 1. Hackelia jessicae (McGregor) Brand From open 1. Symphoricarpus albus (L.) Blake. Open slopes slopes to meadowlands, June to August. Alberta to of the plains to wooded areas in the mountains. May California, south througli Montana to Idaho, Wyo- to August. Alaska to Quebec, south to California, ming and Utah. Webb Springs, Butte Co., 1 July central Idaho, and Nebraska. East Butte, Bingham 1967, 1082. Co., 22 June 1967, 1022. 2. Hackelia patens (Nutt.) Johnst. Sagebrush 2. Symphoricarpus orephilus Gray. Open ground plains to higher elevations in the mountains, June to in the foothills to well up into the mountains, June to August. Montana, west to Idaho, south to Wyoming, August. British Columbia to California, New Mexico. Utah and Nevada. Northeast of Howe, Butte Co., 10 R.30E., T.6N., Butte Co., 1 1 June 1967, 961. June 1067,927. CARYOPHYLLACEAE. PINK FAMILY 6. Lappula Gilib. Stickseed 1 . Arenaria L. Sandwort 1. Nutlets with the marginal prickles in 1 row; co- 1. Inflorescence congested to glomerate; glandular rolla limb up to 4 mm wide .... 2. L. redowskii pubescent, at least above 2 1. Nutlets with the marginal prickles in 2 rows;co- 1. Inflorescence not congested; plants not glandular rolla limb up to 4 mm wide 1 . L. echinata 3. A. kingii 16 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 2. Flowering stems usually less than 10 cm tall, leafy; sepals 5 to 12 mm long •. . .2. A. franklinii 2. Flowering stems taller, from 15 to 50 cm high, nearly leafless; sepals 3 to 6 mm long 1 . A. congesta 1. Arenaria congesta Nutt. var. lithophila Rydb. Plains and mountains, June to August. Montana to Washington, Colorado, and California. Big Butte, Butte Co., 1 July 1967, 1107. 2. Arenaria franklinii Dougl. war. franklinii. Sandy plains and sagebrush slopes. May to June. Central Idaho to central Washington and eastern Oregon and northern Nevada. R.28E., T.4N., Butte Co., 21 May 1967,807. 3. Arenaria kingii (Wats.) Jones var. glabrescens (Wats.) Maguire. Dry ridges to alpine slopes, June to August. East central California, southeastern Oregon, east to central Idaho in Custer, Lemlii and Butte Counties. R.31 E., T.6N., 9 June 1967, 918. CHENOPODIACEAE. GOOSEFOOT FAMILY I. Leaves and bracts spinulose at the tips; fruiting ca- lyx with a broad, transverse wing 8. Salsola 1. Leaves and bracts not spinulose 2 2. Spiny shrubs; leaves linear, semiterete; endosperm lacking; embryo spirally coiled ... .9. Sarcobatus 2. Plants herbs or nonspiny shrubs, if spinose then with flattened leaves and the embryo annular; endosperm present 3 3. Flowers imperfect; perianth lacking in the pistil- late flowers; plants more or less shrubby (in ours) 4 3. Flowers perfect or polygamous; perianth present; on all flowers, often small; plants mostly herbs 6 4. Bracts dorsally compressed; pubescence mainly of inflated hairs which become scurfy when dry 1 . Atriplex 4. Bracts laterally compressed; pubescence of branched hairs not inflated 5 5. Fruiting bracts densely long-villous; leaves narrow with revolute margins 4. Eurotia 5. Fruiting bracts glabrous; leaves entire, not revolute 5 . Grayia 6. Stamens usually over 3; perianth enclosing the fruit and more or less concealing it 7 6. Stamens 1 ; perianth small, fruit exposed 7. Monolepis 7. Flowers solitary in the axils of leaves or bracts; each calyx segment with a hooked spine when mature 2. Bassia 1. Flowers variously disposed in glomerules; calyx segments without hooked spines 8 8. Leaves tipped with a bristlelike hair, fleshy; flow- ers in small axillary glomerules .... 6. Halogeton 8. Leaves without a bristlelike hair, not fleshy; flow- ers usually borne in terminal or axillary glome- rules 3. Chenopodium 1. Atriplex L. Saltbrush; Shadscale 1. Plants annual; monoecious 4. A. rosea 1. Plants perennial shrubs; dioecious 2 2. Fruiting bracts 4-winged; leaves sessile or nearly so \. A. canescens 2. Fruiting bracts not winged; leaves petioled ....3 3. Plants woody througliout, sharply spinose; most- ly 4 to 8 dm tall 2. A. confertifolia 3. Plants woody only at the base, not spinose; low spreading plants, mostly I to 5 dm tall i. A. falcata 1. Atriplex canescens (Pursh) Nutt. var. cfl«e.sce«5 Mostly on saline soils, June to August. Eastern Wash- ington and Oregon, to adjacent California, east to Al- berta, South Dakota, Kansas, Texas and Mexico. Big Lost River Bridge, Butte Co., 6 June 1967, 895. 2. Atriplex confertifolia (Torr.) Wats. Dry fairly saline soil, June to July. North Dakota to eastern Ore- gon, south to northern Arizona and California. R.31E.,T.6N.,ButteCo.,8 June 1967, 1118. 3. Atriplex falcata (M.E. Jones) Standi. Usually on concentrated saline soil, June to August. Saskatch- ewan to Alberta, south to Colorado, Utah, New Mex- ico, California, north to Washington. Pole Line Road, Butte Co., 11 July 1967, 1133. Snake River Plains, Butte Co., 9 August 1950, R. J. Davis; 63 (IDS). 4. Atriplex rosea L. Along roadsides and waste areas, July to September. Much of temperate North America; Old World. Five mi west of Big Butte, Butte Co., 27 July 1950; Davis, 145 (IDS). 2. Bassia Allion 1. Bassia hvssopifolia (Pall.) Kuntze. Saline soil and irrigated land, July to September. British Colum- bia to California, east to Nevada, Idaho, and Mon- tana; Asia; sporadic in the eastern United States. Snake River Plains, Butte Co., 9 August 1950, Davis s.n.(lDS). 3. Chenopodium L. Lamb's Quarters; Pigweed 1. Leaves narrowly linear to linear, entire, 1 -nerved; plants densely farinose .... 3. C. leptophylhim 1. Leaves broader; plants only sparsely farinose to grayish-farinose and greenish above 2 2. Leaves 2 to 4 times longer than wide; pericarp usually adherent to the seed I. C. album 2. Leaves wider, nearly as broad as long; pericarp not tightly adherent to the seed .2. C. fremontii 1. Chenopodium album L. Plants weedy, mostly in waste places, June to September. Eurasia; over much of North America. Big Butte, Butte Co., 27 June 1950; Davis 130 (IDS). 2. Chenopodium fremontii Wats. From saline soil depressions to sagebrush tlats and pine woodlands, June to September. 1. Leaves lanceolate, mostly entire; sparsely fari- nose 1. var. atrovirens BIOLOGICAL SERIES, VOL, 1 I NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 17 1, Leaves mostly triangular and hastately-lobed; grayish-farinose at least below, greenish above , . ,• • • • 2. var. fremontii 2a. Chenopodiiim fremontii Wats. var. atrovirem (Rydb.) Fosberg. British Columbia to Alberta, south to Iowa and Colorado, west to Oregon, Nevada, and California. By EBR. II Reactor, Butte Co., 3 July 1967, 1112. 2b. Chenopodiiim fremontii Wats. var. fr-emontii. British Columbia, Alberta, south to Texas, Mexico and southern California. Big Lost River, Butte Co., 13 July 1967, 1140. 3. Chenopodium leptophyllum Nutt. var. siibgla- bnim Wats. Dry soils of the plains and foothills. May to August, Manitoba, Alberta, and Oregon, south to Texas and Cahfornia. South of Big Butte, Butte Co., 27 June 1950, Davis 150 (IDS). 4. Eurotia Adans. Winter Fat I. Eurotia laiiata (Pursh) Moq. In saline or alka- line soil of the plains and foothills, May to July in our area. Saskatchewan to Washington, south to Texas and California. T.6N., R.32E., Jefferson Co., 28 July 1967, 1177. Butte Co.. 1950, AEC 174 (IDS). 5. Grayia H. & A. Hopsage 1. Grayia spinosa (Hook.) Moq. Saline soil and dry plains and slopes, April to June, Wyoming to Washington, south to Arizona and California. Cinder Butte, Butte Co., 18 May 1967, 762. Base of Big Butte; Butte Co., 22 June 1950; AEC 96 (IDS). 6. Halogeton C. A. Meyer I. Halogeton glomeratus C. A. Meyer. Roadsides and waste ground, July to September. Idaho, Utah, Nevada and California, spreading rapidly. Junction of Higliway 88-Lincoln Blvd., Butte Co., 20 June 1967, 1185. 7. Monolepis Schrad. Poverty weed 1. Monolepis nuttalliana (Schultes) Greene Dry or moist, often saline soil, deserts to middle elevations. May to July. Alberta to Washington, south to Texas and California. Big Butte, Butte Co., 20 June 1950, AEC 14 (IDS). T.2N., R.30E., Butte Co., 13 July 1967, 1135. 8. Salsola L. Russian thistle 1 . Salsola kali L. In moist arid regions of the world, a noxious weed, June to August. Eurasia; througliout most of the western United States, occa- sionally eastward. T.6N., R.31E., Butte Co.. 29 July 1967, 1183. 9. Sarcobatus Nees. Greasewood 1 . Sarcobatus vermiculatus (Hook.) Torr. On saline or alkaline soil in dry regions. May to July. Alberta to Washington, south to New Mexico and California. Pole Line Road, Butte Co., 26 July 1967, 1176. COMPOSITAE. COMPOSITE FAMILY 1. Flowers all ligulate and perfect; plants with milky juice GROUP A 1. Flowers not all ligulate; ray flowers, when pres- ent, marginal and ligulate; juice watery 2 2. Heads radiate 3 2. Heads discoid or disciform 5 3. Rays white to pink, purple or blue . . GROUP D 3. Rays yellow or orange, sometimes purple or red- dish brown at the base 4 4. Pappus chaffy, or of firm awns, or none; recep- tacle chaffy, bristly or naked GROUP B 4. Pappus partly or wholly of capillary bristles, sometimes plumose; receptacle naked GROUP C 5. Pappus of many capillary bristles, sometimes plumose GROUP E 5. Pappus of scales, or awns, or very short chaffy bristles or a mere crown, or lacking, not plumose GROUP F Group A. Flowers all ligulate and perfect; juice milky 1. Pappus of plumose bristles, at least in part ... .2 1. Pappus none, of scales or simple bristles, not plu- mose 3 2. Achenes beaked at the tip; involucres 2 cm long or more; leaves entire, large . . . .34. Tragopogon 2. Achenes not beaked, involucres less than 2 cm long; leaves small and often scalelike 30. Stephanomeria 3. Achenes more or less flattened; stems leafy; heads in panicles or umbels; achenes beaked .... 23. Lactiica 3. Achenes not tlattened; stems leafy or scapose; heads solitary to many 4 4. Flowers pink or purple; leaves linear, subulate or reduced 24. Lygodesmia 4. Rays yellow, sometimes drying to pink; leaves larger, entire to pinnatifid 5 5. Heads several or numerous, rarely solitary 13. Crepis 5. Heads 1 per scape or stem 6 6. Achenes 8 to 10-ribbed or nerved, not spinulose- mucronate; involucre bracts imbricated in several series 2. Agoseris 6. Achenes 4 to 5-ribbed, spinulose-mucronate, principal bracts of the involucre in 1 series 31. Taraxacum Group B. Rays yellow; pappus chaffy, or of firm awns, or none. Receptacle chaffy or bristly throughout . 18 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 1 . Receptacle naked 3 2. Rays neutral; style branches flattened; pappus of 2 readily deciduous awns; leaves mostly basal forming rosettes, cauline leaves reduced 7. Balsamorhiza 2. Rays pistillate and fertile; style branches slender; pappus wanting; leaves mostly cauline 20. Helianthus 3. Rays short and inconspicuous. 1 to 5 mm long; heads smaller, less than 5 mm wide 18. Giitierrezia 3. Rays well developed, 5 to 30 mm long; heads 5 mm or more wide 4 4. Pappus of firm deciduous awns; involucre strong- ly resinous 17. Grindelia 4. Pappus of chaffy scales; involucre tomentose . . . 15. Eriophylhtm Group C. Rays yellow; pappus partly or wholly of capillary bristles. 1. Plants compact dwarf shrubs . . 19. Haplopappus 1 . Herbaceous plants 2 2. Rays white, pink, purple, or violet 3 2. Rays yellow or orange 4 3. Style tips deltoid, obtuse or rounded; phyllaries usually equal or little graded; rays narrow 14. Erigeron 3. Style tips lanceolate or narrower; phyllaries either subequal and the outer leafy, or usually imbricate; leaves alternate and spinulose-toothed 25. Machaeranthera 4. Leaves opposite at least below 5. Arnica 4. Leaves alternate 5 5. Phyllaries in a single series, equal; style tips trun- cate 28. Senecio 5. Phyllaries in 2 or more series, more or less un- equal and graduated 6 6. Heads numerous, small, in panicles or cymes, in- volucre usually 2 to 7 mm high; fibrous rooted plants from a rhizome 29. Solidago 6. Heads few, and relatively large, involucre more than 7 mm high; plants from a taproot 19. Haplopappus Group D. Rays white to pink, purple, red, or blue. 1. Receptacle chaffy or bristly throughout; pappus of graduated bristles, narrow scales, or none . . .2 1 . Receptacle naked; pappus various 3 2. Receptacle chaffy throughout; pappus none .... 1 . Achillea 2. Receptacle densely bristly; pappus of graduated bristles, narrow scales, or reduced 8. Centaurea 3. Pappus of the disk flowers partly or wholly of capillary bristles 4 3. Pappus a short crown, reduced, or none 5 4. Rays numerous, filiform, about equal or slightly longer than the disk flowers; annual plants 12. Conyza 4. Rays larger than the disk flowers . . .14. Erigeron 5. Receptacle conic or hemispheric; leaves pinnately dissected; pappus a minute crown 26. Matricaria 5. Receptacle flat or nearly so; leaves entire 33. Townsendia Group E. Heads or disciform; pappus capillary I . Receptacle densely bristly or leaves more or less spiny and thistlelike 2 1. Receptacle naked; leaves not at all thistlelike, (spiny in species of Tetradymia) 3 2. Plants thistlelike with spiny-margined leaves; in- volucres all spiny 11. Circium 2. Plants not tliistlelike, leaves not spiny-rnargined; involucres not spiny (ours with one species lacer- ate-margined) 8. Centaurea 3. Shrubs 4 3. Herbs 5 4. Involucral bracts 4 to 6, equal 32. Tetradymia 4. Involucral bracts over 6, numerous, imbricate . . . 10. Chrysothamnus 5. Herbage densely white-woolly; leaves simple, entire, alternate and basal, generally tufted 3. Antennaria 5. Herbage not at all white-woolly; stems leafy, without basal tuft of leaves 12. Conyza Group F. Heads discoid or disciform; pappus chaffy, or of awns or none 1. Involucres, at least in part, covered with hooked prickles, or nutlike or burrlike and armed with tubercles or straight spines 2 1. Involucres not as above, all similar 4 2. Heads all alike; involucral bracts hooked at the tip; receptacle bristly 4. Arctium 2. Heads unisexual; involucre of the pistillate heads nutlike or burrlike, armed with hooked prickles, spines or tubercles; staminate involucral heads unarmed; receptacle chaffy 3 Pistillate involucre with hooked prickles 35. Xanthium 3. Pistillate involucre with one to several rows of tuberucles or spines 16. Franseria 4. Receptacle chaffy throughout; heads very small, the involucre only 1 .5 to 4 mm high . . . .22. Iva 4. Receptacle naked, or beset with long hairs in Artemisia 5 5. Involucral bracts herbaceous, not scarious or hya- 3. J BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 19 line; pappus of well developed scales 9. Chaenactis 5. Involucral bracts definitely scarious or hyaline, at least at the apex; pappus a very short crown, scales, or none 6 6. Involucral bracts nearly equal, pale yellowish, with more or less petaloid tips 21. Hymenopappus 6. Involucral bracts not at all petal-like nor yellow- ish, more imbricate, scarious marginal as well as tips 6. Artemisia 1. Achillea L. Yarrow 1. Achillea millefolium L. Circumboreal, April to October. Webb Springs. Butte Co., I July 1967, 1086. 2. Agoseris Raf. False Dandelion 1. Agoseris glaciia (Pursh) Raf. Moist to dry, open places, May to September. British Columbia to Man- itoba, south to California, Arizona, and Minnesota. Big Butte, Butte Co., 22 June 1950, AEC 84 (IDS). 3. Antennaria Gaertn. Everlasting; Pussy-toes 1. Perennial stoloniferous plant; heads several or many 2. A. rosea 1. Perennial plant, without stolons; heads solitary, terminal 1 . A. dimorpha 1. Antennaria dimorpha (Nutt.) T. & G. Foothills and plains, April to May. British Columbia and Montana, south to California, Nebraska and Colo- rado. Middle Butte, Bingham Co., 3 May 1956; French 45 (NRTS). 2. Antennaria rosea Greene. Dry places on the plains to middle elevations. May to August. Alaska to Ontario, south to California and New Mexico. East Butte, Bingham Co., 22 June 1967, 1019. R.29E., T.7N., Butte Co., 10 June 1967, 934. Birch Creek, Clark Co., 25 June 1967, 1043. t. Arctium L. tsuraocK 1. Arctium minus (Hill) Bernh. A weedy plant of waste places, July to October. Over much of the United States and southern Canada. Birch Creek, Clark Co., 5 September 1967, 1214. 5. Arnica L. Arnica 1. Arnica cordifolia Hook. Foothills and moun- tain slopes and woods, April to June. Alaska, south to California and New Mexico; Michigan. Webb Springs, Butte Co., 15 July 1967, 1160. 6. Artemisia L. Sagebrush; Wormwood 1 . Plants all shrubs; flowers all perfect 2 1 . Plants shrubs or herbs; marginal flowers pistillate 5 2. Heads sessile in the axils and surpassed by the subtending leaves; leaves deciduous, 3 to 5-cleft 5. A. rigida 2. Heads not sessile in the axils, upper heads with the subtending leaves not surpassing them; leaves persistent through winter, deeply cleft to merely tridentate 3 3. Flowers mostly 5 to 8 in each head; leaves com- monly deeply cleft into narrow segments, these often 3-cleft 8. A. tripartata 3. Leaves mostly merely toothed at the apex ... .4 4. Plants mostly 1 to 4 dm tall; leaves mostly 1.5 to 5 cm long or less ]. A. arbuscula 4. Plants taller 4 to 20 dm tall; leaves mostly 1.5 to 5 cm long 7. A. tridentata 5. Disk flowers sterile, ovary abortive 6 5. Disk flowers fertile, ovary normal 7 6. Plant spinescent subshrub; achenes and cor- ollas cobwebby with long hairs 6. A. spinescens 6. Unarmed herbs; achenes and corolla glabrous . .-. 3. A. dracunculus 1 . Inodorous (more or less), glabrous, taprooted an- nual or biennial plant 2. A. biennis 7. Aromatic perennial from a rhizome; pubescence white-tomentose 4. A. ludoviciana 1. Artemisia arbuscula Nutt. var. nova (Nels.) Cronq. Dry plains, July to September. Washington to Montana, south to California and New Mexico. South of Junction 88-Lincoln Blvd., Butte Co., 6 Sept- ember 1967. 1224. 2. Artemisia biennis Willd. Along waterways and in sandy soil, August to October. Widely distributed, especially in northwestern United States. Birch Creek, Clark Co., 5 September 1967, 1216. 3. Artemisia dracunculus L. Plains to middle ele- vations, July to October. Yukon to Manitoba, south to California, Texas, and New Mexico; Eurasia. Big Lost River, Butte Co., 13 July 1967, 1139. 4. Artemisia ludoviciana Nutt. var. latiloba Nutt. Dry, open places, June to October. British Columbia to Ontario, south to California, New Mexico, and Arkansas. T.3N., R.29E., Butte Co., 11 July 1967, 1129. 5. Artemisia rigida (Nutt.) Gray. Mostly on dry, icky foothills and plains, September to October. Montana to Washington and Oregon. East Butte; Bingliam Co., 22 June, 1967, 1013. Big Butte, Butte Co., 26 May 1967,866. 6. Artemisia spinescens Eaton. Plains and foot- hills, April to June. Montana to Oregon, south to California and New Mexico. T.6N., R.30E., Butte Co., 6 June 1967,902. 7. Artemisia tridentata Nutt. Mostly of lower ele- vations, but sometimes to timberline, July to Sept- ember. Widespread in the western United States. Birch Creek, Clark Co., 5 September 1 967, 1 2 1 2. 20 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 8. Artemisia tripartita Rydb. Dry plains to moist areas in the mountains, July to September. British Columbia to Montana, south to California and Colo- rado. T.IN., R.30E., Butte Co., August 1965, Harniss, s.n., (UTC). 11. Cirsium Mill. Thistle 1. Cirsium vulgare (Sav.) Airy-Shaw. A weedy plant, July to September. Over much of North America; native of Eurasia. Birch Creek, Clark Co., 5 September 1967, 1213. 7. Balsamorhiza Nutt. Balsamroot 1. Balsamorhiza sagittata (Pursh) Nutt. Valleys and open slopes, April to July. British Columbia to Montana, south to California, South Dakota, and Colorado. Webb Springs, Butte Co., 1 July 1967, 1078. 1 2. Conyza Less. 1. Conyza canadensis (L.) Cronq. A weedy plant, July to September, Southern Canada; Tropical America; over much of the United States. Big Lost River, Butte Co., 1 3 July 1 967, 1 1 46. 13. Crepis L. Hawk's-beard 8. Centaurea L. Star-thistle; Knapweed j involucres, or lower part of stem, or both con- L Perenmal from creepmg roots; involucre greenish- spicuously setose, not glandular ... .2 strammeous, middle and outer bracts broad, with , involucres and stem slightly or not at all setose, subentire hyahne tips 2. C. repens jf ,^,^56 then with gland-tipped setae 3 1. Biennial or short-lived perennial from a taproot; 2. Plants mostly less than 3 dm tall; heads 1 to 9, involucre greenish, the middle and outer each with 10 to 60 flowers ...3. C modocensis bracts with short, dark pectinate tips 2. Plants usually 3 to 8 dm tall; heads 6 to 70, each 1 • C- maculosa vvith 8 to 25 flowers 2. C. barbigera 1. Centaurea maculosa Lam. A weedy plant of 3 ^gi, developed plants 2 to 7 dm tall, with 20 to waste places, June to October. Common in the north- ,00 heads; herbage gray-tomentose to glabrate eastern United States, less common westward; above; heads 5 to 10 flowered .. 1. C acumwato Europe. North^ot Junction 88-22, Butte Co., 5 Sept- 3^ p,^,^^^ ^^^H^^ 0 5 j^ 4 ^^ j^n^ ^^^^ ,_ j^ 25 ember 1967, l^^l. heads; often glandular-hirsute above; heads 10 to 2. Centaurea repens L. A noxious weed, June to 4Q lowered 4. C. occidentalis September. Over much ot the arid portions of the , ^ ■ ■ xi i- 1 n . j . . , . c * u T £M n -M c D ** /-- 1- Crepis acununata Nutt. roothills to moderate west, June to September. T.6N., R.31E., Butte Co., , „. , , .. „, , ?8 I 1 1967 1 ''01 elevations. May to July. Montana to Washington, ' " ' south to California and New Mexico. Sec. 29, T.3N., R.28E., Butte Co., 18 June 1956, McBride, s.n., 9. Chaenactis DC. False yarrow; (NRTS). Dusty Maiden 2. Crepis barbigera Leib. ex Coville. Dry plains 1. C/zae«acr/s(^oi/^/fls// (Hook.) H.& A. Rocky or and foothills. May to July. Oregon to Washington, sandy soil of the plains and foothills. May to Septem- east to central and northern Idaho. R.29E., T.7N., ber. British Columbia to Montana, south to Califor- Butte Co., 9 June 1967,921. nia, Colorado and Arizona. R.3 IE., T.4N., Butte Co., 3. Crepis modocensis Greene ssp. modocensis. 8 June 1967, 1008. Plains and foothills. May to July. British Columbia to Montana, south to Wyoming, and Colorado, and California. Big Butte, Butte Co., 20 June 1950, AEC, 10. Chrysothamnus Nutt. (IDS) Rabbit brush 4 Crepis occidentalis Nutt. ssp. occidentalis. 1. Twigs covered by a feltlike tomentum pi^j^j ^^j foothills, May to July. British Columbia to ' • ^- t^auseosus California, east to South Dakota, and New Mexico. 1. Twigs glabrous to minutely puberulent, not R.3 1 £., T.6N., Butte Co., 8 June 1967, 909. tomentose 2. C vicidiflorus 1. Chrysothamnus nauseosus (Pall.) Britt. Plains and foothills, August to October. British Columbia to Saskatchewan, south to Texas, New Mexico and Calif- ornia. CFA, Butte Co., 9 September 1955, French, 2, (UTC). South of Junction 88-Lincoln Blvd., 24 August 1967, 1210. 2. Chrysothamnus viscidiflonis (Hook.) Nutt. Plains and foothills, July to September. British Columbia to California, east to North Dakota, and New Mexico. South of Junction 88-Lincoln Blvd., Butte Co., 24 August 1967, 1203. 14. Erigeron L. Daisy; Fleabane 1. Leaves trifid to lernately dissected; rays reduced or wanting 1. E. compositus 1. Leaves entire to toothed; rays well developed ..2 2. Rays well over 100; plants fibrous rooted 3 . E. glabellus 2. Rays 1 00 or less; plants from a taproot 3 3. Lower leaves with 3 more or less prominent veins; rays 35 to 65 2. E. corymbosus BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 21 3. Leaves not trinei'ved; rays 50 to 100 4. E. pumihis 1. Erigeron compositus fursh. Ury sana or rocky ground, May to August. Greenland to California, Arizona, South Dakota, and Quebec. T.4N., R.28E., Butte Co., 21 May 1967, 804A. 2. Erigeron corymbosiis Nutt. Sagebrush plains and slopes, June to July. British Columbia to Montana, south to Oregon and Wyoming. Big Butte, Butte Co., 15 July 1967, 1156. 3. Erigeron glabellus Nutt. Along rivers and other moist places, June to July. Alaska to Utah, South Dakota, and Wisconsin. Big Lost River, Butte Co., 13 July 1967, 1152. 4. Erigeron pumihis Nutt. Plains and foothills, May to July. British Columbia to Saskatchewan, south to California and New Mexico. Big Lost River, Butte Co., 13 July 1967, 1148. Middle Butte, Bingham Co., 22 June 1967, 1031. ewan to Colorado, west to Montana, central Idaho, Oregon and California. R.38E., T.4N., Butte Co., 21 May 1967,805. 2. Haplopappus nanus (Nutt.) Eaton Gravelly soil and rocky ledges. May to August. Idaho, west to Washington and California, south to Utah. Big Butte, Butte Co., 28 May 1938, Davis, 161, (IDS). 20. Helianthus L. Sunflower 1. Helianthus petiolaris Nutt. Plains, usually in waste places. June to September. Widespread in the United States. R.31E., T.6N.. Butte Co., 2 July 1967, nil. 21. Hymenopappus L'Her 1 . Hymenopappus filifolius Hook. Gravelly or sandy soil of the plains. May to July. Washington to California, east to North Dakota and Texas. R.30E., T.6N., Butte Co., 1 1 June 1967, 944. 15. Eriophyllum Lag. 1. Eriophyllum lanatum (Pursh) Forbes var. inte- grifolium (Hook.) Smiley. Dry plains to moderate ele- vations in the mountains. May to August. Widespread east of the Cascades. South slope of East Butte, Bing- ham Co., 22 June 1967, 1007A. 16. Franseria Cav. 1. Franseria acanthicarpa (Hook.) Cov. Mostly along streams in sandy soil, July to October. Sas- katchewan to Washington, south to California and Texas. R.31E., T.6N., Butte Co., 18 July 1967, 1 165A. Payne (1964) has placed Franseria as synon- omy nndiQx Ambrosia. 17. GrindeliaWilld. Gum Plant; Resinweed 1. Grindelia squarrosa (Pursh) Dunal. var. serni- lata (Rydb.) Steyerm. Dry, open places, July to Sep- tember. British Columbia to Minnesota, south to Cali- fornia and Texas. T.3N., R.28E., Butte Co., 28 July 1967,1181. 22.1vaL. 1. Annual; leaves long-petiolate ... 1. /. xanthifolia 1. Perennial from a creeping rootstock; leaves sessile or subpetiolate 1 . /. axillaris 1 . Iva axillaris Pursh. Plains and foothills, often in alkaline soil. May to September. Manitoba to British Columbia, south to California and Oklahoma. Big Lost River, Butte Co., 13 July 1967, 1137. 2. Iva xanthifolia Nutt. Along streams and in bottomlands, August to October. Alberta to Wash- ington, south to New Mexico and Texas. Sec. 17, T.3N., R.30E., Butte Co., 2 August 1956, McBride, 79,(NRTS). 23. Lactuca L. Lettuce 1. Biennial or winter annual; leaves prickly on the midrib beneath and on the margins; heads rela- tively small 2. L. scariola 1. Perennial; leaves glabrous or glabrate, not prick- ly heads large and showy 1 . L. pulchella 1. Lactuca pulchella (Pursh) DC. Moist places, June to September. Alaska to California, east to Minnesota and Missouri. T.2N., R.30E., Bingham Co., 23 July 1967, 1173. 2. Lactuca scariola L. A weedy plant of fields and waste places, July to September. Europe; over much of the United States. North of Highway 20, east side, Bingham Co., 29 September 1955, French, 9, (NRTS). 18. Gutierrezia Lag. Matchbrush; Snakeweed 1. Gutierrezia sarothrae (Pursh) Britt. & Rusby. Plains and foothills, July to September. Sas- katchewan, south to California, Mexico, and Kansas. Tractor Flat, Jefferson Co., 22 July 1967, 1203A. 24. Lygodesmia D. Don 19. Halopappus Cass. Rush Pink; Skeleton-weed 1. Low branching shrubs, not caespitose 1. Branches spine-tipped, divaricate, rigid; involucre 2. H nanus imbricate, 7 to 1 3 mm long 2. L. spinosa 1. Caespitose plant, with herbaceous flowering 1. Branches not spine-tipped; involucre merely cal- stems 1 . H. acaulis yculate, 1 8 to 22 mm long ... \. L. grandiflora 1. Haplopappus acaulis (Nutt.) Gray Foothills to 1. Lygodesmia grandiflora (Nutt.) T. & G. Foot- well up in the mountains. May to August. Saskatch- hills and valleys May to July. Idaho and Wyoming to 22 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 30. Stephanomeria Nutt. Rush Pink; Skeletonweed 1. Annual or biennial; pappus bristles plumose on the upper 1/2 to 2/3; achenes rugose-tuberculate and pitted 1 . S. exigua 1. Perennial; pappus bristles plumose to the base; achenes smooth 2. S. temiifolia 1. Stephanomeria exigiia Nutt. Plains and foothills in dry, often sandy soil. June to August. Central Oregon to Wyoming, south to California and New Mexico. T.3N., R.29E., Butte Co., 18 July 1967, 1169. 2. Stephanomeria temdfolia (Torr.) Hall. Plains and foothills to middle elevations, June to Sept- ember. British Columbia and Washington to Montana, south to California and Texas. Junction of Higliway 22-Atomic City, Butte Co., 14 July 1967, 1153. 31. Taraxacum Hall Dandelion 1. Taraxacum officianale Webber in Wiggers. A weedy plant, March to December. Cosmopolitan; native of Europe and adjacent Asia. Sec. 10, T.2N., R.29E., Butte Co., 3 August 1956, McBride, 36, (NRTS). California and New Mexico. T.3N., R.38E., Butte Co., 11 July 1967, 1128. 2. Lygodesmia spinosa Nutt. Dry, rocky or sandy places, July to August. British Columbia to Montana, south to California and Arizona. Tractor Flat, Jeff- erson Co., 22 July 1967, 1200. 25. Machaeranthera Eaton 1 . Machaeranthera canescens Gray. Dry, open places of the plains and foothills, July to October. Saskatchewan to British Columbia, south to Calif- ornia and Colorado. Big Butte, Butte Co., 7 September 1967, 1232. 26. Matricaria L. 1 . Matricaria maritima L. A weed of waste places and roadsides, July to September. Europe; widely established in North America. R. 29E., T.3N., Butte Co., 11 July 1967, 1132. 27. Microseris D. Don 1. Microseris troximoides Gray. Dry, open places in the foothills and plains, April to June. British Columbia, south to California, Montana and northern Utah. R.29E., T.7N., Butte Co., 10 June 1967, 929. 32. Tetradymia DC. Horse Brush 28. Senecio L. Groundsel; 1. Primary leaves forming spines; fiowers 5 to 9 ... Butterweed 2. T. spinosa 1. Cauline leaves well developed, with well devel- 1- Primary leaves not forming spines; flowers 4 oped tuft of basal leaves; glabrous or puberulent 1 • T. canescens near the base 3. S. serra 1- Tetradymia canescens DC. Foothills and plains, 1. Basal leaves well developed and the cauline leaves June to September. British Columbia to Montana, gradually reduced upwards; crisp-villous to arach- south to California and New Mexico. R.31E., T.4N., noid-villous or densely tomentose 2 Butte Co., 18 July 1967, 1171. 2. Pubescence loosely crisp-villous or arachnoid- -■ Tetradymia spinosa H. & A. Foothills and villous, sometimes sparse at flowering time; stems P'ains, May to August. Southwestern Montana and solitary 2. S. inteserrimus central Idaho to Oregon, south to Colorado, and 2. Pubescence strongly white-tomentose; stems usu- Utah. T.6N., R.30E., Butte Co., 6 June 1967, 903. R. ally several 1 S caniis 34E., T.5N., Jefferson Co., 20 June 1967, 1004. 1. Senecio canus Hook. Plains and foothills. May to August. Saskatchewan to British Columbia, south 33. Townsendia Hook. Daisy to Cahfornia, Colorado, and Nebraska. R.29E.,T.3N., 1- Townsendia florifer (Hook.) Gray. Plains and Butte Co. 1 July 1967, 1092. foothills. May to July. Washington to Idaho, south to 2. Senecio integerrimus Nutt. Plains to high in the Nevada, Utah and Wyoming. T.4N., R.28E., 21 May mountains, May to July. British Columbia to Calif- 1967,804. ornia, east to Saskatchewan and Minnesota. R.31E., T.6N., Butte Co., 1 1 June 1967, 942. -''4. Tragopogon L. Goatsbeard 3. Senecio serra Hook. Open places in the foot- 1 • Tragopogon dubiiis Scop. Dry ground, es- hills to middle elevations in the mountains, south to pecially along roadsides. May to July. Over much of California and Colorado. Webb Springs, Butte Co., 15 'he United States; Europe. T.6N., R.31E., Butte Co., July 1967 1158 29 July 1967. 1 184. R.34E., T.5N., Jefferson Co., 20 June 1967,988. 29. Solidago L. Goldenrod 1. Solidago occidentalis (Nutt.) T. & G. Stream- banks and other moist areas, July to October. Alberta to British Columbia, south to California, New Mexico, and Nebraska. Big Lost River, Butte Co., 13 July 1967, 1149. ! I I i 35. Xanthium L. Cocklebur 1. Xanthium strumarium L. var. canadense (Mil[.) T. & G. Disturbed ground, April to October. Cosmo- politan. Sec. 33, T.5N., R.30E., Butte Co., 26 July 1956; French, 75, (NRTS). BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 23 CONVOLVULACEAE. MORNING-GLORY FAMILY \. Convolvulus L. Bindweed; Morning-glory 1. Convolvulus arvensis L. A very troublesome weed introduced from Europe and now well estab- lished in North America. May to August. T.2N.. R.30E., Butte Co., 3 July 1967, 1116. CORNACEAE. DOGWOOD FAMILY 1 . Cornus L. Dogwood 1 . Cornus stolonifera Michx. Common along streams and in moist woods. May to July. Widespread in North America. Webb Springs, Butte Co., 1 July 1967, 1087. Birch Creek; Clark Co., 25 June 1967, 1052. CRASSULACEAE. STONECROP 1. Sedum L. Stonecrop 1. Sedum stenopetalum Pursh. Variable in habitat from the sagebrush plains to subalpine ridges. May to July. Western temperate North America. Big Butte, Butte Co., 1 July 1967, 1106. CRUCIFERAE. MUSTARD FAMILY 1. Pods stipitate; stipe 1 mm long or more 1. Pods not stipitate or stipe less than I mm long .3 2. Stipe 5 mm long or less; petals white or purple . . 15. Tlielvpodium 2. Stipe over 5 mm long, mostly 1 to 3 cm long; petals yellow 14. Stanleya 3. Silique composed of two distinct segments, upper segment commonly sterile; beak stout, over 5 mm long 2. Brassica 3. Pods composed of one segment, seed bearing; beak not stout 4 4. Fruit a silique 5 4. Fruit a silicic 17 5. Petals yellow 6 5. Petals never yellow, white to purple 10 6. Plants with appressed, mostly 2-forked hairs; stig- mas deeply 2-lobed 5. Erysimum 6. Plants glabrous or if pubescent not as above . . .7 7. Petals about 2 mm long; seeds biseriate, some- times unlseriate in Descurainia 8 7. Petals over 2 mm long; seeds uniseriate 9 8. Plants aquatic or semiaquatic; fruit 1 -nerved; pubescence not stellate 11. Rorippa 8. Plants terrestrial; fruit several-nerved; pubescence stellate 4. Descurainia 9. Valves of the silique with a prominent midnerve; plants not rhizomatous; annual or biennial 13. Sisymbrium 9. Valves indistinctly nerved; plants rhizomatous perennials 12. Schoenocrambe 10. Perennial, caespitose plants from a branched cau- 13, 14. 14. 15. 16. 16. dex; leaves entire, mostly basal; pubescence gray- ish with branched hairs; petals 10 to 15 mm long 10. Phoenicaulis Plants erect, not as above; leaves toothed to lobed or smaller with the petals less than 10 mm long 11 Seeds biseriate; pods terete 6. Halimilobos Seeds uniseriate 12 Pubescence not branched or lacking 13 Pubescence branched to stellate 14 Stamens tetradynamous; pods strongly flat- tened parallel to the partition; style lacking 1. Arabis Stamens 6; pods terete or 4-sided, not flattened; style short 15. Thelypodium Leaves 1 to 3 times pinnately compound 4. Descurainia Leaves mostly simple, at least the cauline so . .15 Pods strongly flattened parallel to the partition; leaves simple l . Arabis Pods terete or 4-sided (or nearly so); leaves sim- ple to bipinnatifid 16 Annuals; flowers bracteate 9. Malcolmia Biennial or perennials; flowers ebraceate 6. Halimilobos Silicles strongly flattened, not turgid 18 Silicles not strongly flattened, if so weakly, dis- tinctly turgid 19 Fruit 1-loculed and 1 -seeded; silicle broadly winged 17. Thysanocarpus Fruit 2-loculed, several seeded; silicle only slight- ly winged 8. Lesquerella Pods inflated, nearly globose to oblong-rotund 8. Lesquerella Pods more or less obcompressed 20 Seeds 1 per locule; pubescence simple, when present 7. Lepidium Seeds 2 to many per locule; pubescence stellate, when present 21 Fruit triangular in shape; plants hirsute and stel- late; silicles not wing-margined 3. Capsella Fruit not triangular; plants glabrous; silicles wing- margined all around 16. Thlaspi 1 . Arabis L. Rock Cress 1 . Siliques erect, not over 2 mm broad; seeds with a wing 0.3 mm broad or less, wingless; stems simple 5. A. nuttallii 1 . Siliques erect to reflexed; seeds winged, the wing 0.3 mm broad or more 2 2. Plants usually less than 2 dm tall; silique various but not reflexed; stems 1 mm thick or less 4. A. microphylla 2. Plants mostly over 3 dm tall; siliques frequently reflexed 3 3. Leaves mostly densely pubescent, never ciliate; siliques spreading to pendulous, not sharply re- flexed; cauline leaves auriculate 4 19. 19. 20. 20. 21. 24 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 3. Leaves not densely pubescent, greenish to gray- ish, ciliate or the siliques sharply reflexed, or the stem leaves not auriculate 5 4. Petals 4 to 6 mm long; seeds with a wing at least 0.5 mm broad; style short 1 . A. cobremis 4. Petals 6 to 9 mm long; wing of the seeds less than 0.5 mm broad; style lacking .... 3. A. ligiiifera 5. Lower cauline leaves usually petiolate; siliques 3 to 7 cm long; 1 -nerved to about midlength 2. A. holboellii 5. Lower cauline leaves usually auriculate; siliques 4 to 12 cm long, 1 -nerved for nearly the full length 6. A. sparsiflora 1 . Arabis cobrensis Jones. Sagebrush hills and plains, May to July, Wyoming, east across south central Idaho, to central Oregon and northern Nevada. Near Big Butte, Butte Co., s.n., Davis, 167, (IDS). R.29E., T.3N., Butte Co., 6 June 1967, 882. 2. Arabis holboellii Hornem. var. retrofacta (Grab.) Rydb. Sagebrush plains to higher elevations in the mountains. May to August. Montana, north to British Columbia, south to California, east to Colo- rado. R.28E., T.4N., 20 May 1967, 797. 3. Arabis Hgnifera A. Nels. In sagebrush areas. May to June. Wyoming, Idaho to Arizona and Nevada. N.W. sec. 4, T.3N., R.28E., Butte Co., 18 May 1954, Fineman, 152, (UTC). 4. Arabis microphylla Nutt. in T. & G. var. sax- imontana Rollins. Commonly in the lower montane areas and higher mountains, April to July. Montana and Wyoming, to Idaho, and Utah. Webb Springs, Butte Co., 22 May 1967, 829. East Butte, Bingham Co., 18 May 1967,774. 5. Arabis nitttallii Robin. Low valleys to higher in the mountains, April to August. Alberta to eastern Washington and Nevada, south to Wyoming, Idaho and Utah. R.29E., T.7N., Butte Co., 10 June 1967 928. 6. Arabis sparsiflora Nutt. var. sparsiflora. Sage- brush plains, April to June. Idaho, Utah to north- eastern California. Dry plains on Little Lost River, Butte Co., 30 May 1938, probably Davis, 162, (IDS). This plant should be looked for on the NRTS, partic- ularly on the N.W. portion of the site. 2. Brassica L. Mustard I. Brassica juncea (L.) Coss. Commonly a weed along roadsides and in waste places, May to August. Common througliout North America; Native of Europe, T.2N., R.30E., Bingham Co., 23 July 1967, 1174. 3. Capsella Medic. Shepherds purse I. Capsella bursa-pastoris (L.) Medic. A weed, March to July. Common througliout most of North America. Atomic City, Bingham Co., 1 July 1967, 1089. 4. Descurainia Webb. & Berth. Tansy mustard 1 . Pods subclavate to clavate 2. D. pinnata 1 . Pods linear, not enlarged at the apex 2 2. Leaves 2 to 3 times pinnate; pods over 20-seeded A. D. Sophia 1. Leaves only pinnate; pods less than 20-seeded . . . 3 3. Siliques fusiform; style commonly over 0.5 mm long 1 . D. californica 3. Siliques not fusiform; style less than 0.5 mm long 3. D. richardsonii 1 . Descurainia californica (Gray) Schulz. Wyo- ming to Oregon, south througli the Great Basin. Big Butte, Butte Co., 27 June 1950, AEC, 1 19, (IDS). 2. Descurainia pinnata (Walt.) Britt. var. filipes (Gray) Peck. A weedy and common species, April to July. Throughout most of southern Canada and the United States. Big Butte, Butte Co., 26 June 1950, AEC, 111, (IDS). 3. Descurainia richardsonii (Sweet) Schulz. Plains to Middle elevations, June to August. Throughout western North America. Big Butte, Butte Co., 22 June 1950, AEC, 82, (IDS). 4. Descurainia sophia (L.) Webb. A weedy plant along roadsides and in cultivated areas, March to July. Europe; Most of North America. T.4N., R.31E., Butte Co., 20 May 1967,783. 5. Erysimum L. Walltlower 1. Pedicels 4 to 8 mm long; petals pale yellow. 7 to 1 1 mm long 2. E. inconspicuum 1. Pedicels 8 to 13 mm long; petals yellow to deep orange or somewhat reddish, 15 to 25 mm long . 1 . E. asperum 1. Erysimum asperum (Nutt.) DC. In our area commonly in sagebrush. May to July. In most of the western states, to Minnesota, Kansas, and Oklahoma. Arco, Butte Co., 17 May 1936, Qirst, s.n., (IDS). Near Big Butte, Butte Co., 29 May 1938, Davis, 164, (IDS). 2. Erysimum inconspicuum (Wats.) MacMillan. Plains, often in alkaline soil, June to July. Ontario to Alaska, south to southeastern Oregon, and Montana, to Colorado; central United States. Near Big Butte, Butte Co., 29 May 1938, Davis, 164, (IDS). 6. Halimolobos Tausch. 1. Seeds biseriate; siliques glabrous; cauline leaves auriculate 2. H. virgata 1 . Seeds uniseriate; siliques hairy; cauline leaves ses- sile but not auriculated 2 2. Siliques not constricted between the seeds, con- sistently stellate; leaves mostly entire; stigma not lobed, rounded 3. H. whitcdii 2. Siliques strongly torulose, sometimes glabrate; leaves commonly lyrate; stigma often lobed .... 1 . H. perplexa BIOLOGICAL SERIES, VOL. r 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 25 Eastern Montana to the Dakotas, Minnesota and Illi- nois, south to central Idaho, Utah, and Colorado. Six miles S.W. of Atomic City, Butte Co., 26 May 1967, 861. 9. Malcolmia (L.) R. Br. 1. Malcolmia africana (Willd.) R. Br. Becoming a weedy plant in Colorado, south and central Idaho, Nevada and Arizona; native of Africa, April to June. T.2N., R.3IE., Bingliam Co., 22 June 1967, 1026. Junction of Higliway 28-22, Clark Co., 25 May 1967, 852. 1. Halimolobos perplexa (Henderson) Rollins. Dry soil and loose rocky areas. May to June. Central and west-central Idaho. Big Butte, Butte Co., 20 June 1950, AEC. (IDS). 2. Halimolobos virgata (Nutt.) Schulz. Open plains and foothills, June to July. Yukon to Saskatch- ewan, to central Idaho to Wyoming, Utah and Colo- rado. Big Creek, Butte Co., 30 May 1 938, Davis, 1 78, (IDS). Should be looked for on the site. 3. Halimolobos whitedii (Piper) Rollins. Sage- brush and desert plains. May to June. Central Wash- ington and Bingliam County, Idaho. East Butte, Bingham Co., 22 Jime 1967, 1024. 10. Phoenicaulis Nutt. 1. Phoenicaulis cheriranthoides Nutt. in T. & G. 7. Lepidium L. Pepper grass Sagebrush plains and hills, April to June. Idaho to 1. Style present; cauline leaves auriculate to perfoli- central Washington, south to Nevada and California, ate 2. L. perfoliatum North slope of Big Butte, 26 May 1967, 878. 1. Style lacking; cauline leaves not auriculate or per- foliate 2 11. Rorippa Scop. Yellow water-cress 2. Petals usually lacking or rudimentary; pods ob- 1. Rorippa msturtiiim-aqiiaticiim (L.) Schinz. & long to obovate 1 . L. densiflonim Thell. Along streams and other fresh water bodies, in 2. Petals sometimes present; pods elliptic to oval shallow water. March to October. Native of Europe, 3. L. virginicum now well established in North America. Birch Creek, \ . Lepidium densiflonim Schrad. var. macroc- Clark Co., 25 June 1967, 1021. arpum Mulligan. Dry soil, April to June. Eurasia; now widespread in North America. This variety occurs 12. Schoenocrambe Greene from Alaska to California, and Arizona, east to \ . Schoenocrambe linifolia {^\M.) Greene. }Aosi\y Montana and Colorado. R.30E., T.6N., Butte Co., 1 1 "i sagebrush and juniper plains. May to June. British June 1967,948. Columbia, south to Nevada, east to Montana, south 2. Lepidium perfoliatum L. Dry, waste ground, to Idaho and Utah. Arco Desert, Butte Co., 16 May March to June. Native of Europe and now well estab- 1936, Davis, s.n., (IDS). R.29E., T.4N., Butte Co., 22 lished in the western and central part of the United May 1967, 819. States. Sec. 12, T.6N., R.33E., Jefferson Co., 6 June , , „. , . inc<; \A D -A zi. /MDTo '-^- Sisymbrium 1956, McBride, 56, (NRTS). , o- j, ■ ;,■ • i m .i i j -,.,.... , ,. ,^ 1. Sisymbrium allissimum L. Mostly along road- 3. Lepidium virginicum L. var. medium (Greene) ■, ; . , , j m » c . u „,,,';,„*. , , .. , sides and in waste places, weedy. May to September. C. L. Hitchc. Becoming a weedy plant, March to ^^^^^^ ^^ ^^ ^^„ established in the United June. Norl^hern Call lornia to eastern Washington, east ^^^^^^ ^^^^ ^^^ _ g^^^^ ^^ j2 June 1967, 959. through Idaho to Wyoming, south to Arizona, New Mexico, and Texas. Butte Co., June 1950, AEC, 142, (IDS). 14. Stanleya Nutt. Prince's plume 1 . Stanleya viridiflora Nutt. Dry plains and valleys, June to August. Wyoming, west througli cen- 8. Lesquerella Wats. Bladder-pod tral Idaho to central Oregon and Montana, south to 1. Pedicles uniformly recurved, not straight or sig- Utah and Nevada. T.4N., R.31E., Butte Co., 4 July moid; fruit globose; leaves mostly narrowly ob- 1967, 1117. lanceolate 2. L. ludoviciana 1. Pedicels sigmoid or straight; fruit ovate to ob- 15. Thelypodium Endl. long; leaves broader than oblanceolate 1 ■ Cauline leaves sagittate or clasping at base; stipe 1 . /,. alpiiia less than 2 mm long 2. T. sagittatum 1. Lesquerella alpina (Nutt.) Wats. var. laevis 1- Cauline leaves not as above; stipe 2 to 4 mm long (Pays.) C. L. Hitchc. Dry plains and foothills and 1- T. laciniatum mountain ridges. May to July. Alberta, south to Mon- I. Thelypodium laciniatum (Hook.) Lindl. var. tana, and western South and North Dakota, west to milleflorum (Nels.) Pays. Deserts and plains, April to Colorado and central and southeastern Idaho. R.30E., July. Wide spread in western North America. Near Big T.6N., Butte Co., 11 June 1967, 945. Should be Butte, Butte Co., 29 May 1938, (IDS). looked for on the site. 2. Thelypodium sagittatum (Nutt.) Endl. Com- 2. Lesquerella ludoviciana (Nutt.) Wats. Probably mon in valleys, deserts, and lower mountain areas, yai. ludoviciana. Mostly of the plains. May to June, usually somewhat moist ground. June to July. Mon- 26 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN tana to Idaho and Wyoming, west to Washington, Oregon and Cahfornia. Birch Creek, Clark Co., 25 June 1967, 1045. Arco, Butte Co., 16 May 1956, no author, s.n., (IDS). 16. Thlaspi L. Penny cress 1 . Thlaspi arvense L. A common weed in North America, April to August. Webb Springs, Butte Co., 26 May 1967, 871. Birch Creek, Clark Co., 25 June 1967, 1048. 17. Thysanocarpus Hook. Fringe Pod 1. Thysanocarpus curvipes Hook. Moist, often open hillsides; April to June. British Columbia to Cal- ifornia, east to Idaho. Arco Desert, Butte Co., 20 May 1956, Quinn, s.n., (IDS). EUPHORBIACEAE. SPURGE FAMILY 1 . Euphorbia L. Spurge 1. Plants prostrate; stipules present; seeds with transverse ridges 2. E. glyptosperma 1 . Plants erect ; stipules lacking; seeds smooth 1 . E. esiila 1. Euphorbia esula L. Mostly weedy. May to July. Eurasia; over much of the United States. Big Lost River, Butte Co., 11 July 1967, 1130. 2. Euphorbia glyptosperma Englm. Dry, sandy soil of the plains. June to September. Idaho, west from British Columbia to California, east to New York, south to Texas. T.6N., R.31E., Butte Co., 29 July 1967, 1186. FUMARIACEAE. FUMITORY FAMILY 1. Corydalis Medic. 1. Corydalis aurea WiUd. In moist and dry habi- tats, variable. May to July. Nova Scotia, west to Alaska, south to Pennsylvania, New Mexico and Cali- fornia. Arco Desert, Butte Co., June 1950, AEC Pro- ject, 6, (IDS), (IDS). GERANIACEAE. GERANIUM FAMILY 1. Erodium L. 'Her. Stark's-bill 1 . Erodium cicutarium (L.) L.'Her. Native of Eurasia and Africa, April to July. Widespread in the United States. Lost River, Butte Co., 11 July 1930, Davis, s.n., (IDS). GROSSULARIACEAE. CURRENT AND GOOSEBERRY FAMILY 1. Ribes L. Currents and Gooseberries 1. Flowers glabrous, briglit yellow ... 1. R. aureum 1. Flowers pubescent glandular, or both, not bright yellow 2 2. Hypanthium twice as long as the sepals; berries 5 to 7 mm in diameter, red 2. R. cereum 2. Hypanthium about equal to the sepals; berries 10 to 12 mm in diameter, blue to black 3. R. viscosissimum 1. Ribes aureum Pursh. In moist wooded areas in the mountains to stream banks at lower elevations, April to May. Idaho, west to north central Washing- ton to California, east to the Rocky Mountains from Saskatchewan and South Dakota to New Mexico. Webb Springs, Butte Co., 22 May 1967, 824. Big Lost River, Butte Co., 13 July 1967, 1136. 2. Ribes cereum Dougl. var. inebrians (Lindl.) Hitchc. Variable in habitat from dry rocky areas to moist shaded areas, April to June. Central Idaho to central Montana, east to Nebraska, south to Utah and Nevada. Webb Springs, Butte Co., 26 May 1967, 875. 3. Ribes viscosissimum Pursh. Along creeks, open areas, from moist areas to dry plains. May to June. Idaho, west to the Cascade Mountains, from British Columbia to northwestern Colorado. T.6N., R.30E., Butte Co., 18 May 1967,760. HYDROPHYLLACEAE. WATERLEAF FAMILY 1. Flowers mostly solitary; leaves entire 3 1 . Flowers in scorpioid or subcapitate cymes; leaves entire to pinnately lobed or dissected 2 2. Flowers in scorpioid cymes; placentae narrow . . . 4. Phacelia 2. Flowers in subcapitate clusters; placentae broad . 2. Hydrophyllum 3. Plants acaulescent, perennial mesophytes 1 . Hesperochiron 3. Plants caulescent; annual xerophytes .. .3. Noma 1 . Hesperochiron Wats. 1. Hesperochiron pumilus (Griseb.) Porter. In the lower foothills and valleys to meadow lands in the mountains, April to June. Idaho, west from Washing- ton to California, east and south from Montana to Colorado, Utah, and Arizona. Five miles south of Atomic City, Butte Co., 22 May 1967, 820. 2. Hydrophyllum L. Waterleaf 1 . Hydrophyllum occidentale Gray. Mostly in moist places in the woods, April to July. Elmore and Jefferson County, Idaho, east to Oregon and Califor- nia, south to Utah and Arizona. R.34E., T.5N., Jef- ferson Co., 20 June 1967, 1001. 3. Nama L. 1. Nama densum Lemmon var. pan'iflonim (Greene) Hitchc. Sandy soil of the plains. May to July. Washington to California, east to Idaho, Nevada, Utah, and Wyoming. Sec, 24, T.3N., R.30E.. Butte Co., 25 June 1957, 134, Goodwin, (NRTS). 4. Phacelia Juss I . Leaves mostly entire 2 1 . Leaves pinnatifid or pinnately compound 3 2. Plants perennial; stamens strongly exserted 3. /*. hastata BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 27 2. Plants annual; stamens barely exserted or included 4. P. humiliis 3. Filaments long exserted; ovules and seeds 4 or less; style cleft to near the base 2. P. glandulosa 3. Filaments included; ovules and seeds more than 4, style cleft Va of its length . . . 1 . P. glandulifera 1 . Phacelia glandulifera Piper. Commonly in sage- brush or sandy soil of the plains, May to June. Wash- ington to California, east througli central Idaho to Wyoming and to Nevada. Shell Road, Butte Co., 23 May 1967,837. 2. Phacelia glandulosa Nutt. Rocky ridges of the plains and mountains, June to August. Centra! Idaho to Montana, Colorado and Utah. Plot 4, Snake River Plains, Butte Co., 17 July 1950, AEC, 82, (IDS). 3. Phacelia hastata Dougl. ex Lehm. In a wide va- riety of habitats. May to August. British Columbia, east to Colorado and Nebraska. East Butte, Bingham Co., 22 June 1967, 1018. Shell Road, Butte Co., 8 June 1967,923b. 4. Phacelia humihis T. & G. Mostly in fairly moist habitats. May to July. Washington to California, east to Nevada, Utah, and Idaho. Webb Springs, Butte Co., 13 June 1967,969. LABIATAE. MINT FAMILY 1. Agastache Clayton 1. Agastache urticifolia (Benth.) Kuntze. Foothills and mountain slopes, June to August. British Colum- bia, south to Colorado. Webb Springs, Butte Co., 1 July 1967. 1081. LEGUMINOSAE. PEA FAMILY 1. Fertile stamens 5, flowers small in dense spikes, corolla irregular 8. Petalostemon 1. Fertile stamens usually 10; flowers papilionaceous 2. Stamens separate; corolla yellow, racemose 10. nennopsis 2. Flowers not yellow, or if so then the stamens united (stamens mostly united) 3 3. Leaves glandular-punctate and 3-foliate or the fruit with hooked spines 4 3. Leaves not glandular-punctate or if so then the leaves not 3-foliate or with hooked spines on the fruit 5 4. Plants 1.5 to 5 dm tall; leaves mostly 3-foIiate, fruit not spiny 9. Psoralea 4. Plants usually larger, 3 to 1 2 dm tall; leaflets 7 to 15; fruit spiny 2. Glycyrrhiza 5. Leaves even-pinnate, terminal portion modified into tendrils 11. Vicia 5. Leaves with a terminal leaflet, odd-pinnate or tri- foliate 6 6. Leaves palmately compound or trifoliate; flowers in elongate racemes or heads 9 6. Leaves not palmately compound 7 7. Fruit a loment; keel longer than the wings 3. Hedysarum 1 . Fruit not a loment; keel shorter than the wings . . 8 8. Plant mostly without leafy stems; keel beaked . . 7. Oxytropis 8. Plant usually with leafy stems; keel not beaked . . 1 . Astragalus 9. Leaves palmately compound 4 Lupinus 9. Leaves trifoliate 10 10. Pods falcate to coiled, heavily veined 5. Medicago 10. Pod straight or nearly so, lightly veined 6. Melilotus 1. Astragalus L. Locoweed; Milk vetch 1. Flowers ascending to erect, in a subcapitate raceme, purplish; legumes erect, villous 1 . A. agrestis 1 . Flowers spreading to descending or less com- monly erect; the raceme subcapitate to elongate, ochroleucous to bluish or purplish; legumes vari- ous but if erect not villous 2 2. Leaflets 3 to 7, the petioles persistent as a thatch at the base of plant; legumes commonly curved, bilocular and sessile 2. A. calycosus 2. Leaflets various in number, commonly more than 7; petioles usually not persistent; legumes vari- ous, but if bilocular than bladdery inflated or woody to leathery and straight 3 3. Plants acaulescent; leaves and peduncles arising from the apex of the caudex branches; legumes woolly -villous; calyx tubular-cylindric 10. A. purshii 3. Plants with well developed stems; leaves and ped- uncles arising from well separated nodes; legumes strigose-glabrate or glabrous 4 4. Plants from elongate, subrhizomatous branches; terminal leaflet confluent with the rachis; leg- umes 1-loculed, bladdery inflated, purplish mot- tled 4. A. ceramicus 4. Plants from caudices, the caudex branches not or seldom (except in A. miser) subrhizomatous; ter- minal leaflet jointed to the rachis; legumes var- ious, but if bladdery inflated then 2-loculed . . .5 5. Calyx tube cylindric, over 5 mm long 6 5. Calyx tube campanulate, less than 5 mm long . .9 6. Ovaries and legumes stipitate, the stipe more than 2.5 mm long, the body of the legume curved or straight at maturity 7 6. Ovaries and legumes sessile, the body of the leg- ume straight to slightly curved at maturity ... .8 7. Body of the legume curved at maturity; calyx gibbous-saccate dorsally behind the pedicel 6. A. curvicarpus 7. Body of the pod nearly or quite straight; calyx not gibbous-saccate 7. A. filipes 28 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 8. Flowers bluish to purplish, the wing tips whitish; legumes unilocular; plants flowering in early spring 5. A. cibarius 8. Flowers ochroleucous, the keel tip purplish; leg- umes bilocular; plants flowering in late spring or early summer 3. A. canadensis 9. Flowers with distinctly purplish veins; keel tip produced into an elongate point at riglit angles to the keel; legumes unilocular, spreading to de- scending; leaflets more than three times longer than broad 9. A. miser 9. Flowers lacking purplish veins; keel tip various, but not as above; legume unilocular or bilocular, ascending or spreading; leaflets less than three times longer than broad 10 10. Flowers commonly less than 12 mm long; pods bilocular, ovoid-inflated, membranous, spreading 8. A. lentigiuosiis 10. Flowers more than 12 mm long; pods unilocular, oblong-subcylindric, leathery, erect-ascending . . . 11. A. terminalis 1. Astragalus agrestis Dougl. ex Hook. Commonly on clay soil of the plains and up in the mountains. Idaho, north to Manitoba and the Yukon, south to California and New Mexico. Birch Creek, Clark Co., 25 June 1967, 1127. 2. Astragalus calycosus Torr. ex Wats. Sagebrush to dry open slopes. May to July. Central Idaho to Wyoming, south to California, Arizona and New Mex- ico. Six miles southwest of Atomic City, Bingham Co., 26 May 1967. 864. R.28E., T.4N., Butte Co., 1 2 June 1967,953. 3. Astragalus canadensis L. var. hrevidens (Grandg.) Barn. From moist areas to alkaline flat, June to July. Idaho, west to California, north and east to British Columbia to the Atlantic Coast, and from the Great Plains to Texas. T.6N., R.31E., Butte Co., 8 June 1967,906. 4. Astragalus ceramiciis Sheld. Mostly in sandy areas. May to July. Central Idaho to eastern Montana, the Dakotas and Nebraska, south to Arizona. T.6N., R.31E., Butte Co., 8 June 1967,911. R.34E., T.5N., Jefferson Co., 20 June 1967, 993. 5. Astragalus cibarius Sheld. Sagebrush plains. May to June. Central Idaho, north to Montana, south to Nevada, Utah, Colorado, and New Mexico. R.28E., T.4N., Butte Co., 20 May 1967, 792. 6. Astragalus cun'icarpus (A. Hell.) Macbr. var. cun'icarpus. Sagebrush plains. May to July. Idaho, west to Oregon, south to Nevada and California. Mid- dle Butte, Bingham Co., 6 May 1967, 884. 7. Astragalus filipes Torr. Sagebrush slopes. May to June. Idaho, north and west from British Columbia to California and Nevada. Big Lost River Bridge, Butte Co., 6 June 1967,885. 8. Astragalus lentiginosus Dougl. ex Hook. var. salinus (Howell) Barn. Desert alkaline tlats to middle elevations in the mountains. T.6N., R.31E., Butte Co., 8 June 1967,910. 9. Astragalus miser Dougl. ex Hook. var. praeter- itus Barn. From dry ground at lower elevations to moist meadows in the mountains. May to July. Cen- tral Idaho, south to Wyoming, west to Montana. East Butte, Bingham Co., 22 June 1967, 1016. T.4N., R.28E., Butte Co., 1 3 June 1 967, 980. 10. Astragalus purshii Dougl. ex Hook. var. purshii. Sagebrush plains to the lower foothills in the mountains, April to June. Idaho, north and west from British Columbia to California, east to Alberta, the Dakotas, and Colorado. Big Lost River Bridge, Butte Co., 6 June 1967,897. lOa. Asgragalus purshii Dougl. ex Hook. var. concinnus Barn. On the plains, from central Idaho to west central Montana. Six miles east of Arco, Butte Co., 13 May 1933, Davis s.n., (IDS). 11. Astragalus terminalis Wats. Common in sage- brush. May to June. Central Idaho to Montana, east to Wyoming. T.4N., R.28E., Butte Co., 12 June 1967,956. 2. Glycyrrhiza L. Licorice 1 . Glycyrrhiza lepidota Pursh. Common along streams and in waste places. May to August. British Columbia to Ontario, south to New York, New Mexico, and California. Big Lost River, Butte Co., 29 July 1967, 1188. 3. Hedysarum L. 1. Hedysarum boreale Nutt. Dry plains. May to August. Saskatchewan to central Idaho, south to New Mexico and Arizona. R.31E., T.6N., Butte Co., 8 June 1967,914. 4. Lupinus L. Lupine 1 . Plants annual 4. L. pusillus 1 . Plants perennial 2 2. Leaves mostly cauline; plants erect with a well branched crown 1 . L. argenteus 2. Leaves basal and cauline 3 3. Plants low and spreading to matted 2. L. lepidus 3. Plants erect, cauline leaves with petioles to 20 cm long 4 4. Leaflets pubescent on the upper surface, acumi- nate to apiculate; flowers bluish to violet or white in our area 3. L pninophilus 4. Leaflets glabrous on the upper surface, usually rounded or acute; flowers deep violet to purple . 5. L. wyethii 1. Lupinus argenteus Pursh subsp. argenteus var. tenellus (Dougl. in G. Don) Dunn. Central Idaho to central Montana, Wyoming and Utah, May to July. Big Butte, Butte Co., 26 May 1967, 869. 2. Lupinus lepidus Dougl. ex Lindl. Dry plains and slopes, June to August. California to British Columbia, east to Montana, Wyoming and Colorado. BIOLOGICAL SERIES, VOL. 11 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 29 Five miles west of Arco, Butte Co.. 14 May 1939, O. Mays, s.n., (IDS). 3. Liipimis X pninophilus M. E. Jones. Plains and foothills. May to June. Central and eastern Washing- ton, northeastern Oregon, southcential Idaho, south to Nevada, Utah, and Colorado. Webb Springs, Butte Co., 24 May 1967.840. 4. Lupimis piisiUus Pursh. Mostly in sandy soil. May to June. Idaho, west to Washington, south to California, east to Alberta to Nebraska. Webb Springs. Butte Co., 24 May 1967,838. 5. Lupiniis wyethii Wats. Open ridges and forests to sagebrush plains. May to July. Idaho, west from British Columbia to California, east from Alberta to Colorado. Big Lost River Bridge, Butte Co., 6 June 1967,886. 5. Medicago L. I. Medicago sativa L. Native to the Old World. Often escaping from cultivation, June to October. Camas Creek, Jefferson Co., I September 1955. French, 23, (NRTS). 6. Melilotus Mill. Sweet clover 1. Melilotus officinalis (L.) Lam. Native of the Mediterranean area, a weedy plant in our area. May to September. Sec. 3. T.6N.. R.33E., Jefferson Co.. 7 June 1956, McBride, 1 20. (NRTS). 7. O.xytropis DC. Stemless loco 1 . Corolla purple to reddish-purple; pubescence seri- ceous-silky; leaves 3 to 7 cm long; leaflets 7 to 1 5 mm long; pods 1 .5 cm long 1 . O. lagopiis I. Corolla white to yellow; pubescence densely grayish villous to strigose; leaves 5 to 30 cm long; leaflets 1-3.5 cm long; pods 1.5-2.5 cm long 2. O. sericea 1. Oxytropis lagopiis Nutt. var. lagopus. Plains and lower mountains. May to June. Idaho and Mon- tana. R.28E., T.4N., Butte Co., 12 June 1967, 950. 2. Oxytropis sericea Nutt. var. sericea. Middle ele- vations to the plains. May to July. Western Montana to northeastern Nevada, New Mexico, and Texas. R.31 E., T.3N., Bingham Co.. 24 May 1967, 856. Co.. 20 June 1967, 996. R.31E.. T.6N., Butte Co 18 July 1967, 1167. 10. Thermopsis R. Br. Buck-bean 1 . Thermopsis moiuana Nutt. Mostly along streams or moist places. May to August. Idaho, west from British Columbia to California, east from Mon- tana to Colorado and Utah. Birch Creek, Clark Co., 25 June 1967, 1044. 1 1 . Vicia L. Vetch I. Vicia sativa L. Introduced from Europe and now widespread in the United States, May to July. R.3IE.. T.6N.. Butte Co., 8 June 1967, 907. LINACEAE, FLAX FAMILY I. Linum L. Flax 1. Linum perenne L. Mostly in dry areas. May to July. Over much of western North America; Eurasia. North of junction 22-28, Clark Co., 24 June 1967 1060. LOASACEAE. BLAZING-STAR FAMILY 1 . Mentzelia L. Blazing star 1. Annual plants 1 to 4 dm tall; petals 1.5 cm long . I . M. alhicaulis 1. Biennial plants 3 to 10 dm tall; petals 1.5-8 cm long 2. M. laevicaulis 1. Mentzelia albicaulis Dougl. ex Hook. Desert plains, in sandy soil. May to July. Idaho, west from British Columbia to California, east to Montana to New Mexico. Cave by EBR.II Reactor. Butte Co., 18 May 1967.769. 2. Mentzelia laevicaulis (Dougl.) T. & G. Desert and mountains. July to September. Idaho, west from British Columbia to California, east to Wyoming. Arco desert. Butte Co.. June 1950. Davis, 183, (IDS). MALVACEAE. MALLOW FAMILY I. Sphaeralcea St. Hil. 8. Petalostemon Michx. Prairie clover 1. Petalostemon ornatum Dougl. ex Hook. Com- monly in sagebrush, sandy or rocky soil. May to July. South and central Idaho, west to Washington and Oregon. Sec. 9. T.3N.. R.30E., Butte Co., 26 June 1957, Goodwin, 131, (NRTS). 9. Psoralea L. I. Psoralea lanceolata Pursh. Mostly in sandy soil. May to September. Idaho, west from Washington to Nevada, east to Nebraska. R.34E., T.5N., Jefferson 1. 1. Leaves crenate or if lobed less than halfway to the midrib; calyx bracteolate 2. S. munroana Leaves divided almost to the midrib; calyx tri- bracteolate 1 . S. grossulariaefolia 1. Sphaeralcea grossulariaefolia (H. & A.) Rydb. Mostly on dry soil, June to July. Central Idaho, west from Washington to Nevada and Utah. Big Lost River, Butte Co., 1 July 1967, 1094. 2. Sphaeralcea munroana (Dougl.) Spach. ex Gray. Dry plains and mountains. May to August. Idaho, west and south from British Columbia, to Call- 30 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN fornia and Utah. T.4N.. R.30E.. Butte Co., 1 June 1953, McBride, s.n.,(UTC). NYCTAGINACEAE. FOUR-O'CLOCK FAMILY Abronia Juss. Sand verbena I. 2. 3. 4. Oenothera L. Evening Primose Plants with leafy flowering stems 2 Plants without leafy flowering stems 4 Petals large, over 1 cm long, wliite or pink 3. O. pallida Petals smaller, less than I cm long or yellow . . .3 Plants annual, 1 to 5 dm tall; stigmatic lobes less ] 1. Abronia mellifera Dougl. ex Hook. Dunes and than 5 mm long 4. O. scapoidea sandy soil, May to July. Columbia River Gorge, east 3. Plants biennial or perennial, mostly over 5 dm i to central and southeastern Washington and adjacent tall (up to 5 m); stigmatic lobes about 5 mm long ! Oregon, east to eastern Idaho. R.31E., T.6N., Butte 2. O. hookeri I Co., 9 June 1967, 916. 4. Leaf margins ciliate; capsules short-pedicellate .. I ONAGRACEAE. EVENING PRIMOSE FAMILY '^- ^ ^aespitosa var. 1 . Seeds with a tuft of hairs (coma) at the tip; , ' ■ ! .• ; •.; "^'"'Si"^^'' \ I II ■, T z7 ■; 1. 4. Leai margins not ciliate; capsules sessile leaves usually opposite 2. Epdobium '^ ,, ^ . ' 1 . Seeds without coma; leaves mostly alternate . . .2 "'■ ^- ^^^/^'^^^^ ^ar. montana 2. Ovary 2-loculed 3. Gayophytum ^^- Oenothera caespitosa Nutt. var. marginata j 2 Ovary 4-loculed 3 (Nutt.) Munz. Commonly ot dry, rocky soil in the j 3. Stigma 4-lobed '.'.'.'.'.'.'.'.'.'.'.'.'.'.'.4. Oenothera P'^'"^- ^^V ^o July. Idaho, west to California, south , 3. Stigma capitate 1. Camissonia ^^ ^^^'^^^^^^ ^"'l, ^''^: ^^st ot Big Lost River Bridge, i ^ ^ Butte Co., 13 July 1967, 880. lb. Oenothera caespitosa Nutt. var. montana 1. Camissonia Link. (Nutt.) Durand. Washington, Oregon and Idaho. East I. Petals yellow; capsule sessile, straight, fusiform, Butte, Bingham Co., 22 June 1967, 1007. T.4N., - crowded \ . C. andina R.31E., Butte Co., 20 May 1967, 784. 1. Petals white, often reddish; capsule sessile, often 2. Oenothera hookeri T. & G. In moist areas of \ contorted 2. C. minor the plains and lower mountain streams, June to Sep- j 1. Camissonia andina (Nutt.) Raven. Dry soil. May tember. Central Idaho, west to Washington, Oregon, 1 to June. Desert near Big Butte, Butte Co., 17 May and California, south to Utah, New Mexico, east to j 1938, Davis, 139, (IDS). Te.xas. Big Lost River, Butte Co., 13 July 1967, 1142. j 2. CaH!/MO«/fl m/«or (Nels.) Raven. Dry soil. May 3. Oenothera pallida Lindl. Mostly in hot, dry ; to June. Near Big Butte, Butte Co., 28 June 1950, sandy soil. May to July. Idaho, west to Washington AEC Project, 164 (IDS). and Oregon, south to Arizona and New Mexico. EBR. II Reactor, Butte Co., 3 July 1967, 1115. 2. Epilobium L. Willow herb 4. Oenothera scapoidea Nutt. in T. & G. Com- i 1. Epilobium watsonii Barn. var. parishii (Trel.) monly in sagebrush and sandy soil. May to July. Cen- j Hitchc. Commonly in moist places, June to Septem- tral Idaho, west to Oregon, south to Wyoming, Colo- ber. Spreading throughout the United States. Birch rado, and Utah. Shell Road, Butte Co., 8 June 1967, Creek, Clark Co., 5 September 1967, 1219. 922. 3. Gayophytum Juss 1. Plants usually branched from the base; capsules OROBANCHACEAE. BROOMRAPE FAMILY nearly sessile; petals I mm long or less 2. G. racemosum \ . Orobanche L. Broomrape; Cancerroot I. Plants usually branched from above; capsules j pedicelled; petals 1 to 5 mm long 1 . Flowers long-pedicellate, without bractlets; co- 1 1 ■ G. mittallii roUa equally 5-cleft 1 . O. fasiculata 1. Gayophytum mittallii T. & G. Dry sandy and 1. Flowers sessile or pedicels less than 3 mm long gravelly soil of the plains and foothills, June to Au- with a pair of bractlets beneath the calyx; calyx gust. Common in the western states. Shell road, Butte unequally 5-cleft 2. O. grayana Co., 23 May 1967, 836. Our plants belong to variety 1. Orobanche fasiculata Nutt. Foothills and diffusum and villosum and are quite distinctive in this plains. May to July. Michigan to British Columbia, ; area. south to Texas and California. Tractor Flat, Jefferson j 2. Gayophytum racemosum T. & G. Lower desert Co., 20 June 1967, 995. I plains to well up in the mountains. May to July. 2. Orobanche grayana Beck. British Columbia, Washington to California, east from Montana to Colo- south to California and Nevada, June to September, rado and Utah. T.4N., R.31E., Butte Co., 20 May Six miles southwest of Atomic City, Butte Co., 18 1967,781. July 1967, 1162. BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 31 PLANTAGINACEAE. PLAINTAIN FAMILY 1 . Plantago L. Plaintain 1. Small annual plants; leaves small, linear to nar- rowly lanceolate; seeds 2 per capsule 2. P. piirshii 1. Perennial plants; leaves broad, large; seeds 6 to 30 per capsule 1 . p, major 1. Plantago major L. A weedy species. May to September. Cosmopolitan. Birch Creek, Clark Co., 5 September 1%7, 1220. 2. Plantago piirshii R. & S. Dry soil. Ontario to British Columbia, south to Texas and California. R.29E.,T.3N., Butte Co., 1 July 1967, 1095. POLEMONIACEAE. PHLOX FAMILY 1. Cauline leaves lacking, represented only by the persistent cotyledons and basally conate bracts; small annuals 4. Gymnosteris 1. Cauline leaves more or less well developed; an- nual or perennial plants 2 2. Calyx-tube tending to remain unbroken to ma- turity of the capsule; leaves prevailingly alter- nate, at least above 3 2. Calyx-tube tending to split along intercoastal membranes 4 3. Stamens unequally inserted on the corolla tube; calyx enlarging in fruit 1 . CoUomia 3. Stamens more or less equally inserted close to the sinuses of the corolla; calyx not enlarging in fruit 7. Navarrctia 4. Leaves entire, mostly opposite; filaments very unequally inserted 8. Phlox 4. Leaves dissected or lobed; filaments more or less equally inserted 5 5. Annuals with the tlowers borne in leafy- bracteate, tomentose heads; anthers deeply sagit- tate 2. Eriastnim 5. Perennials or if annual the flowers not in heads . . 6 6. Leaves sessile, palmatifid or pinnatifid; shrubby pungent plants 6. Leptodactylon 6. Leaves not sessile and palmatifid, principal leaves alternate, pinnately lobed to entire 7 7. Leaf lobes setose (spine-tipped); sepals unequal . 5. Langloisia 7. Leaf lobes usually not spine-tipped; sepals equal or subequal 3. Qilia 1. Collomia Nutt. 1. Plants 1 to 6 dm tall, unbranched; filaments un- equally inserted 1 . C. linearis 1. Plants 1 to 2 dm tall, freely branched; filaments equally inserted 2. C. tenella 1. Collomia linearis Nutt. Dry to moist habitats, ' to August. Common in western North America 'oving eastward. Webb Springs, Butte Co., 1 '070. 2. CoUomia tenella Gray. Common in sagebrush and open places, June to July. Central Idaho, west to Washington and Oregon, east to Wyoming, south to Utah and Nevada. Webb Springs, Butte Co., 1 July 1967, 1072. 2. Eriastrum Woot. & Standi. 1. Eriastrum sparsiflorum (Eastw.) Mason var. wilcoxii (A. Nels.) Cronq. Desert plains, often in sand. May to August. Idaho, west to California, south to Utah. EBR. II Reactor, Bingham Co., 3 July 1967, 1113. 3. Cilia R. &P. 1 . Plants biennial or perennial 2 1 . Plants annual 3 2. Corolla red; stems mostly over 2 dm tall I . G. aggregata 2. Corolla white, stems mostly 2 dm or less tall .... 2. G. congesta 3. Leaves usually entire, basal and stem leaves pres- ent 4 3. Leaves toothed to pinnatifid, mostly basal; co- rolla 1 .5-4 mm long 5. G. tenerrima 4. Leaves toothed or lobed; seeds mostly 8 to 12 per locule, less than 1 mm long 3. G. leptomeria 4. Leaves usually pinnatifid; seeds 2 to 8 per locule, 1 .5 mm long 4. G. simtata 1. Gilia aggregata (Pursh) Spreng. Rocky, open ground of the plains to middle elevations. May to August. British Columbia to Idaho, east to Colorado, south to Mexico. Southwest of Howe, Butte Co., 12 June 1967,954. 2. Gilia congesta Hook. Dry rocky ground at lower elevations to higlier in the mountains, June to August. Idaho, west to Oregon and California, east to North Dakota, Nebraska and Colorado. R.3 1 E., T.6N, Butte Co., 8 June 1967,905. 3. Gilia leptomeria Gray. Mostly in sandy soil of the plains. May to July. Idaho, west from Washington to California, south to Colorado to New Mexico. Webb Springs, Butte Co., 24 May 1967,839. R.34E., T.5N., Jefferson Co., 20 June 1967, 989. 4. Gilia sinuata Dougl. ex Benth. Mostly of the plains. May to July. Idaho, west from Washington to California, east to Wyoming, south to New Mexico. T.6N., R.30E., Butte Co., 18 May 1967, 759. 5. Gilia tenerrima Gray. Plains to middle ele- vations, June to August. Central Idaho and Montana, west to Oregon, south to Wyoming, Colorado, and Utah. Six miles west of Atomic City, Butte Co., F July 1967, 1164. 4. Gymnosteris Greene 1. Gymnosteris nudicaule (H. & A.) Greer sandy places, April to May. Baker County and the snake Rivpr Plains cnuth to Nev 32 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN toward Arco, Butte Co. (IDS). 13 May 1933, Davis, s.n.. 5. Langloisia Greene 1. Langloisia setosissinia (T. & G.) Greene. Com- monly in sandy soil of the plains, May to July. Oregon, south to California, east and south from Idaho to Mexico. T.6N., R.30E., Butte Co., 6 June 1967,899. 6. Leptodactylon H. & A. 1. Plants diffusely branched, up to 30 cm tall; leaves predominately alternate 1 . L. pungens 1. Plants only moderately branched, up to 15 cm tall; leaves predominately opposite 2. L. watsonii 1. Leptodactylon pungens (Torr.) Rydb. Rocky or sandy soil of the plains and desert. May to July. British Columbia to California, east to Montana, Nebraska, south to New Mexico. Shell Road, Butte Co., 23 May 1967,835. 2. Leptodactylon watsonii (Gray) Rydb. Common on rocky ledges and gravelly soil, June to July. Idalio, Colorado, and Utah. R.28E., T.4N, Butte Co., 22 7. Navanetia R. & P. 1. Navanetia breweri (Gray) Greene. Dry or moist ground, commonly as understory. May to July. Central Idaho, west to Washington to California, east from Wyoming to Arizona. Big Butte Canyon, Butte Co., 1 July 1967, 1108. 8. Phlox L. Phlox 1. Caespitose, glandular to hirsute plants not espe- cially mat forming; leaves firm, mostly over 1 cm long; style over 5 mm long 2 1. Mat torming, arachnoid plants; leaves firm and pungent, mostly 0.5-lcm long; flowers mostly sessile; style 2 to 5 mm long 2. P. hoodii 2. Flowers with long pedicels; style 6 to 15 mm long; plant with well developed internodes 3. P. longifolia 2. Flowers short-pedicellate; style 5 to 10 mm long; plant with short internodes 1 . P. aculeata 1. Phlox aciilaeata A. Nels. Commonly in sage- brush, alkaline, or sandy clay soil, April to May. Snake River Plains, west to Oregon. East Butte, Bingham Co., 18 May 1967, 773. T.4N., R.28E., Butte Co., 20 May 1 967, 695. 2. Phlox hoodii Rich. Dry, rocky plains and foothills, April to June. Common througli the western states and southern Canada. R.28+., T.4N., 20 May 1967,789. 3. Phlox longifolia Nult. Dry, rocky places, April to July. British Columbia to Montana, south to Calif- ornia, Wyoming, Colorado, and New Mexico. Bit Butte, Butte Co., 20 June 1950, AEC, 15, (IDS). POLYGONACEAE. BUCKWHEAT FAMILY 1 . Leaves without sheathing stipules; stamens mostly 9 2 1. Leaves with sheathing stipules; stamens 4 to 8 .. 3 2. Involucre with 3 to 6 spine-tipped teeth or lobes 2. Oxytlieca 2. Incolucre with 3 to 10 nonspinose teeth or lobes 1. Eriogonum 3. Perianth segments mostly 5 (sometimes 4 or 6), never with grainlike callosities, stigma capitate . . 3. Polygonum 3. Perianth segments usually 6 (sometimes 4), 1 or more often with a grainlike callosity; stigmas tufted 4. Rumex 1. Eriogonum Michx. 1 . Annual plants 2 1 . Perennial plants 3 2. Involucres sessile, spicate, mostly single at the di- chotomies and along the unbranched stem tips, not retlexed 7. E. vimineum 2. Involucres with retlexed peduncles; leaves all basal 2. E. cermium 3. Perianth narrowed to a slender stipelike base which is jointed to the pedicel 4 3. Perianth rounded to acute at base, narrowed to the point of attachment (joint) with the pedicel but without a slender stikelike base 5 4. Flowering stems with a whorl of leaves (bracts) about midlength; plants erect and well over 10 cm tall (1-4 dm tall) 3. E. heracleoides 4. Flowering stems without a whorl of leaves (bracts) about midlength; plants low and spread- ing, 1 0 cm or less tall 1 . E. caespitosum 5. Plants matforming; involucres often capitate or umbellate 6 5. Plants not matforming; involucres borne in broad, compound, cymose inflorescences 5. E. rnicrothecum 6. Perianth segmented for 2/3 to 3/4 or its length; pale grayish-lanate throughout 4. E. maiKum 6. Perianth segmented nearly to the swollen basal joint; from pannose-lanate and almost white on both surfaces to less tomentose, sometimes greenish above 6. E. ovaUfolium 1. Eriogonum caespitosium Nutt. Sagebrush flat, grassy hills and pinyon-juniper lands, April to July. Eastern California and Oregon, eastward to Montana, Idaho, Wyoming, and Colorado. Lava beds, Bingliam Co., 27 May 1 969, E.C. Moran, s.n., ( BRY). 2. Eriogonum cermium Nutt. var. cernuum. In the hills and valleys of the plains, June to August. South central Oregon to southeastern California, east across southern Idaho to eastern Montana, Nebraska, Colo- rado, and New Mexico. By ERB. II Reactor, Bingham Co., 3 July 1967, 1114. BIOLOGICAL SERIES, VOL. 1 I NO. 4 ELORA OF THE NATIONAL REACTOR TESTING STATION 33 3. Eriogonum lieracleoides Nutt. var. Hera- cleoides. Gravelly to loamy soil in the desert and' rocky ridges to about 6,000 feet. May to August. Utah, Nevada, Wyoming, east to western Montana, northeastern California, and British Columbia. East Butte, Bingliam Co., 22 June 1967, 1021. 4. Eriogonum mancum Rydb. Sagebrush flats and grassy hills, June to July. Granite and Powell Coun- ties Montana, south to Lehmi and Clark Co., Idaho. Birch Creek, Clark Co., 25 June 1967, 1057. 5. Eriogonum microthecum Nutt. var. laxiflonim Hook. Sandy deserts and low mountain slopes, June to August. Washington and California, east to Mon- tana, Nebraska, Arizona, New Mexico and Colorado. South of Junction 88-Lincoln Blvd., Butte Co., 24 August 1967, 1202. 6. Eriogonum ovalifolium Nutt. var. ovalifolium. From sagebrush slopes to above timberline. May to August. British Columbia, south through Washington, western Oregon, and California, on the eastern side of the Rocky Mountains. T.4N., R.28E., Butte Co., 21 May 1967, 802, T.6N., R.30E., Butte Co., 6 June 1967,900. 7. Eriogonum vimineurn Dougl. in Benth. Pine woodlands, to sagebrush and desert lands in sandy or rocky soil, June to September. Washington and Ida- ho, southward througli Oregon into northwestern Nevada and central California. Should be looked for on the site. 2. Oxytheca Nutt. 1 . Oxytheca dendroidea Nutt. var. dendroidea. Common in sagebrush, June to August. Central Wash- ington, Oregon, south to California, east to Idaho and Nevada; Chile and Argentina. T.3N., R.30E., Butte Co., 25 June 1956, McBride, 105, (NRTS). 3. Polygonum L. Kjiotweed 1. Perianth undivided except for a short limb; plants fragile 2. P. polycenoeoides 1 . Perianth divided at least to the middle; plants not fragile 2 2. Calyx lobes yellowish; plants erect, 1 m tall .... 3. P. ramisissinnim 2. Calyx lobes white or pinkish margined; plants prostrate to ascending, 1 to 4 dm tall 1. P. achoreum 1. Polygonum achoreum Blake. Dry waste ground, July to September. Eastern Oregon and Montana, south to Idaho and Colorado, east to Canada and nor- thern United States to Quebec and New York. S.W. of Big Butte, Butte Co., 27 Jtme 1950, AEC, 144, (IDS). 2. Polygonum polyceneoides Juab. & Spach. Growing in sandy areas. New York and south central Idaho; Asia, June to August. East of Jet. 88-Lincoln Blvd., 24 August 1967, 1207. 3. Polygonum ramosissinuim Michx. Moist to dry wasteground, July to September. In most of the United States, except the S.E.; over most of southern Canada. Webb Springs, Butte Co., 13 June 1967, 970. 4. Rumex L. Dock 1 . Flowers mostly imperfect; leaves hastate 1 . R. acetosella 1 . Flowers perfect; leaves not hastate 2 2. Plant rhizomatous, leaves leathery 4. R. venosus 2. Plant with a strong taproot; leaves various 3 3. Leaf margins crisped; petioles usually papillose- puberulent 2. R. crispus 3. Leaf margins plane to undulate, but not crisped; petioles glabrous 3. R. salicifolius 1. Rumex acetosella L. A very weedy plant. May to August. Native of Europe and now over most of the world. Desert, Butte Co., 1950, AEC, 37A,(IDS). 2. Rumex crispus L. A weed of fields and waste places, generally in moist areas, June to September. Over most of the United States; Europe. Webb Springs, Butte Co., I July 1967, 1066. Flood Control area, Butte Co., 1 July 1967, 1096. 3. Rumex salicifolius Weinm. Sandy and moist areas in the plains and mountains, June to September. Alaska to California, east to Nebraska and Texas, and in Canada and northern U.S. to Quebec, south to New York; Europe. Big Butte, Butte Co., June 1950; AEC. 128. (IDS). 4. Rumex venosus Pursh. Dry sandy soil, April to June. Alberta to Washington, California, east to Texas. T.4N., R.28E., Butte Co., 21 May 1967,883. RANUNCULACEAE. BUTTERCUP FAMILY Flowers irregular; upper sepal spurred 2. Delphinium Flowers more or less regular; upper sepal not spurred, petals may be spurred 2 Petals spurred; fruit of follicles .... 1. Aquilegia Petals not spurred; fruit of achenes 3. Ranunculus 1. Aquilegia L. Columbine I. Aquilegia formosa Fisch. Moist mountains, meadows, and slopes. May to August. Widely dis- tributed in western North America. Webb springs, 1 July. 1080. Birch Creek, Clark Co., 25 June 1967, 1041. 2. Delphinium L. Larkspur Stems I -several, commonly over 4 dm tall, from a very fibrous root system 1. D. andersonii Stems usually single, usually less than 4 dm tall, from a tuberous root system ... .2. D. mcnziesii 1 . Delphinium andersonii Gray. Commonly in sagebrush and juniper of desert ranges, April to June. 1. 1. 34 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN California, Nevada and south central Idaho. South slope of Middle Butte, Bingliam Co., 22 June 1%7, 1029. 2. Delphinium menziesii DC. subsp. iitahense (Wats.) D. Sutherland. Prairies, canyons, and lower montane meadows. British Columbia, south to Calif- ornia and Utah, April to June. R.28+., T.4N., Butte Co., 21 May 1967,812. 3. Ranunculus L. Buttercup; Crowfoot 1. Plants not both scapose and perennial; achenes strongly compressed 2 1. Perennial scapose plants; achenes not strongly compressed 1 . R. audersouii 2. Low annual plants, 7 mm, or less high, appearing early in the spring; achenes woolly, usually densely so 4. R. testiculatus 2. Mostly perennial plants (in ours), over 7 cm tall; achenes not woolly 3 3. Receptacle globose; achenes each with a broad thin wing or stipe at base 2. R. glaberrimus 3. Receptacle ellipsoid; achenes not stipitate or winged at base 3. R. macounii 1 . Raminciilus andersonii Gray. Sagebrush and juniper-pinyon hills, April to June. California, south to Nevada, north to eastern Oregon and central Idaho. North of Middle Butte, Bingliam Co., 21 April 1967,752. 2. Ranancidiis glabemimis Hook. var. ellipticits Greene. Sagebrush and grassland valleys, March to June. California, to British Columbia, east to North and South Dakota, Nebraska, and New Mexico. T.IN., R.30E., Butte Co., 22 May 1967; 823. 3. Ranunculus macounii Britt. var. macounii. Usually in damp places at low elevations. May to July. Northern California, north to British Columbia and Alberta, south to Arizona and New Mexico, east to Labrador, Michigan to Nebraska. Birch Creek, Clark Co., 25 June 1967, 1050. 4. Ranunculus testiculatus Crantz. Commonly in sagebrush areas, March to May. Oregon and eastern Washington, east to Idaho and Colorado, south to Nevada. North base of East Butte, Bingham Co., 18 May 1967,775. RHAMNACEAE. BUCKTHORN FAMILY 1. Ceanothus L. 1. Ceanothus velutinus Dougl. ex. Hook. var. ve- lutinus. Moist areas in the mountains, June to August. Idaho, west from British Columbia to Nevada, east to Montana, South Dakota and Colorado. Big Butte, Butte Co., 1 July 1967, 1101. ROSACEAE. ROSE FAMILY 1 . Plants bearing prickles; leaves pinnate; carpels several 9. Rosa 1. Plants not with the above combination of charac- ters 2 2. Ovary more or less inferior, compound, 2 to 5-lo- cular; fruit a pome 1 . Amelanchier 2. Ovary superior, with a single carpel; fruit of achenes, follicles, drupes, or druplets 3 3. Fruit a fleshy, 1-seeded drupe; pistil 1; leaves simple 7. Pninus 3. Pistils 1 to several, if 1 then not drupaceous; leaves simple, compound or persistent 4 4. Petals lacking; pistil 1 , developing into an achene; style plumose; leaves simple ....2. Cercocarpus 4. Petals present; pistils usually more than 1, if 1 then without a plumose style; leaves simple or compound 5 5. Leaves compound with 3 or more leaflets 6 5. Leaves simple or deeply lobed, but not compound 8 6. Petals bright yellow; fruit of dry achenes 6. Potentilla 6. Petals white; fruit of follices or druplets 7 7. Plants with bristles or prickles; fruit fleshy, of druplets 10. Rubus 1. Plants without bristles or prickles; fruit of follicles 3. Chamaebatiaria 8. Leaves entire, not lobed 4. Holodiscus 8. Leaves toothed to shallowly lobed 9 9. Leaves deeply 3-toothed at the tip only, small, revolute-margined; pistils solitary; fruit an achene 8. Purshia 9. Leaves usually 3 to 5-palmately lobed, larger, not revolute-margined; pistils 1 to 5; fruit a follicle . . 5. Physocarpus I. Amelancier Medic. Serviceberry 1. Amelanchier utahensis Koehne. Commonly in sagebrush deserts to well up in the mountains. May to June. Central Idaho, west to California and Oregon, east to southern Montana, Wyoming, Colorado, New Mexico and Texas. East Butte, Bingliam Co., 22 June 1967, 1023. Webb Springs, Butte Co., 26 May 1967, 870. 2. Cercocarpus HBK Mountain Mohogany 1. Cercocarpus ledifolius Nutt. in T. & G. Lower foothills to rocky mountain ridges, April to June. Montana, southwest to Washington, Oregon, Calif- ornia and Arizona, west to Colorado and Utah. R.28E., T.4N., Butte Co., 21 May 1967, 803. 3. Chamaebatiaria (Porter) Maxim. Fern Bush 1 . Chamaebatiaria millefolium (Torr.) Maxim. Desert plains and foothills, commonly in lava in our area. June to August. Oregon, south to Nevada and California, Arizona, east througli central Idaho to Utah. R.29E., T.7N., Butte Co., 10 June 1967, 1165b. 4. Holodiscus Maxim. Mountain Spray 1. Holodiscus dumosus (Hook.) Heller. Rocky soil BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 35 of the plains to moist areas in the mountains, June to August. Oregon, south to Cahfornia, east and south to central Idaho, Wyoming, Colorado, New Mexico, Utah, Arizona, and Nevada. East Butte, Bingham Co., 22 June 1967, 1014. Webb Springs, Butte Co., 13 June 1967,973. 5. Physocarpus Maxim. Nine Bark 1 . Physocarpus alternans (Jones) Howell. Dry rocky slopes at low elevations. Central Idaho, south to Utah and Nevada. R.29E., T.7N., Butte Co., 10 June 1967, 926. Should be looked for at the NRTS. 6. Potentella L. Cinquefoil 1 . Stamens 1 0 to 20; style terminal 2 1 . Stamens more than 20; style lateral 3 2. Lower portion of the stems hirsute; stamens mostly over 15 4. P. norvegica 2. Lower portions of the stems soft pubescent; stamens 10 or 1 5 2. P. biennis 3. Stems creeping (stoloniferous) and rooting; flowers solitary; pubescence grayish, silky- tomentose I . P. anserina 3. Stems erect, not rooting; flowers in terminal cyiTies; pubescence glandular-puberulent to pi- lose-glandular 3. P. glandulosa 1. Potentella anserina L. Moist areas of the plains and meadowlands. May to August. Atlantic coast, west to California. Flood Control Area, Butte Co., 1 July 1967, 1098. 2. Potentella biennis Greene. Moist shady areas at lower elevations to higher up in the mountains. May to August. Idaho, west to Washington, north and south from Saskatchewan to Colorado. Big Lost River, Butte Co., 13 July 1967, 1 145. 3. Potentella glandulosa Lindl. var. glandulosa. Moist soil in the plains to middle elevations. May to July. Washington, Idaho, and Oregon. Big Butte Canyon, Butte Co., 1 July 1967, 1 100. 4. Potentella non'egica L. A weed in the eastern United States, now over much of North America. May to August. Flood Control Area, Butte Co., 1 July 1967, 1097. 7. Prunus L. Cherry 1. Prunus virginiana L. Variable in habitat from the lower grassland and sagebrush plains to well up in the mountains. May to July. Webb Springs, Butte Co., 1 July 1967, 1064. 8. Purshia DC. ex Poir Bitter Brush 1 . Purshia tridentata (Pursh) DC. Commonly from sagebrush to wooded areas, April to June. Idaho, north and west from British Columbia to Nevada and California, east to Montana, Wyoming, Utah, and New Mexico. Six miles south west of Atomic City, Bingham Co., 22 May 1967, 865. Big Butte Co., 20 June 1950, AEC, 16, (IDS). 9. Rosa L. Rose I. Rosa woodsii Lindl. Commonly along stream- banks or moist areas. May to July. Idaho, west to Oregon, Washington, and California, east to Minne- sota and to Texas. Big Butte, Butte Co., 22 June 1950, AEC, 89, (IDS). 10. Rubus L. Blackberry; Raspberry 1. Rubus ideaus L. Streambanks to moist woods. May to July. Over much of North America; Asia. Big Butte, Butte Co., 1 July 1967, 1101. RUBIACEAE. MADDER FAMILY 1. Galium L. Bedslraw, Cleavers 1. Plants perennial from creeping rhizomes; leaves mostly in whorles of 4; fruit without ucinate hairs (may be flexuous) 2. G. multiflonim 1. Plants annual trom a taproot; leaves 2 to 4 in a whorle; fruit with ucinate hairs . . I. G. bifolium 1. Galium bifolium Wats. Often as understory in dry to moist places. May to August. British Columbia and Montana, south to California and Colorado. Big Butte, Butte Co., 27 June 1950, AEC, 120,(IDS). 2. Galium multiflonim Kell. Desert plains and rocky slopes. May to August. Central Idaho, west from Washington to California, south to Utah, and Arizona. Webb Springs, Butte Co., 13 June 1967, 968. SALIACEAE. WILLOW FAMILY 1. Buds with several scales; stamens numerous; flowers borne on a cuplike disk 1 . Populus 1. Buds with a single scale; stamens commonly 1 to 2 or many; flowers without disks 2. Salix I. Populus L. Cottonwood 1. Petioles strongly flattened laterally; bark of the trunk mostly smooth and white 2. P. tremuloides 1 . Petioles terete or nearly so; bark of the trunk not as above, generally ashy gray . . \. P. angustifolia 1 . Populus angustifolia James. Common along streams. Saskatchewan and Alberta, south to Ne- braska, New Mexico and Arizona. Big Lost River, Butte Co., 4 September 1967, 1028. 2. Populus tremuloides Michx. Common in sandy or gravelly soil of mountains and hillsides. Southern Labrador to Alaska, south to New Jersey, New Mexico and Arizona. Webb Springs, Butte Co., 1 July 1967, 1090. 2. Salix L. Willow 1. Stamens 3 to 8; leaves narrow, finely serrulate, more than three times as long as broad; filaments hairy at base 3. S. lasiandra 1 . Stamens mostly 2 or less; other characters various 2 36 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 4. 5 Scales of the aments early deciduous; leaves 5 to 15 times as long as wide, narrow, stigmas lobed nearly to the base 2. S. exigiia Scales of the aments persistent, mostly brown to blackish; leaves usually broader 3 Ovaries and capsules glabrous 4 Ovaries and capsules more or less densely short- hairy to sparsely hairy with age 5 Aments appearing before the leaves (precocious); fruits 3 to 5 mm long; leaves oblanceolate 4. S. lasiolepis Aments coetaneous or serotimous; leaves loosely long-woolly-villous when young, less so with age, entire to toothed 1. S. commutata Style mostly 0.7-1.7 mm long; leaves glabrous when fully expanded 5. S. phylicifoUa Style 0.5-0.8 mm long; leaves reddish-strigose be- neath, sometimes glabrous .... 6. S. scouleriana 1. Salix commutata Bebb. Wet places at middle to high elevations, July to September. Western Wyoming to northern Utah into northern California, north through central and southeastern Idaho to southern Yukon and southern Alaska. Birch Creek, Clark Co., 25 June 1967, 1055. 2. Salix exigiia Nutt. Moist situations, commonly along streams, Washington and Oregon to Wyoming and Colorado. Big Lost River, Butte Co., 29 July 1967, 1192. Birch Creek, Clark Co., 25 May 1967, 854B. 3. Salix lasiandra Benth. Stream banks. Upper Sonoran and Transition Zones. California and New Mexico, north to British Columbia and Alberta, sparingly to Alaska and the Yukon. T.2N., R.29E., Butte Co., 4 July 1967, 1123. 4. Salix lasiolepis Benth. Along streams at lower elevations. British Columbia to Baja California, east to northern and southwestern Idaho, Nevada, southern Utah, western Texas and adjacent Mexico. Webb Springs, Butte Co., 1 July 1967, 1084. 5. Salix phylicifoUa L. var. monica (Bebb) Jeps. Diverse in habitat, usually in moist places. West central Montana and central Idaho to New Mexico, Utah, Nevada, and eastern California. North of Junction 28-22, Clark Co., 25 May 1967, 854A. 6. Salix scouleriana Barratt. Transition and Canadian Zones. Alaska and Yukon to California, Arizona, and New Mexico, east to Manitoba and South Dakota. Near Big Butte, Butte Co., 19 June 1950, AECs.n., (IDS). SAXIFRAGACEAE. SAXIFRAGE FAMILY 1. Plants usually bulblet-bearing above or below ground level; styles or branches 3 2. Lithophragma 1. Plants not bearing bulblets; styles or branches us- ually 2 1 . Heuchera 1 . Heuchera L. Alumroot 1. Heuchera paniflora Nutt. ex T. & G. Gravelly to talus slopes, June to August. Utah and New Mexico, and Colorado, north to Wyoming, central Idaho and Nevada. Big Butte, Butte Co., 1 July 1967. 1102. 2. Lithophragma Nutt. Woodland Star I. Lithophragma tenella Nutt. Dry sagebrusl plains. May to June. Washington to southern Oregon east through central and southern Idaho to Montan and Wyoming. S.W. of East Butte, Bingham Co., I May 1967,770. SrROPHULARIACEAE. FIGWORT FAMILY Anther-bearing stamens 5; corolla more or le; regular 7. Verbascur Anther-bearing stamens 2 or 4; corolla mostly ii regular Anther-bearing stamens mostly 2; corolla nearl regular 3. Veronic, Anther-bearing stamens 4; corolla irregular . . . .; Corolla with a slender spur at the base 4. Linarii Corolla without a spur '■ Corolla with a fifth sterile stamen present, some times reduced; leaves opposite or whorled 5 Corolla without a fifth sterile stamen; leaves al ternate or opposite i P'a-ts annual, small, lax and weak; sterile stamer reduced and glandlike 2. Collinsia Plants perennial, large and usually stout; sterile stamen slender and elongate 6. Penstemon Flowers borne in heads or spikes; corolla glaeate. forming a hood or beak 7 6. Flowers axillary; corolla bilabiate, but not form- ing a hood or beak 5. Mimulus 1 . Galea surpassing the lower lip; plants perennial . . 1 . Castilleja 1. Galea about equal with the lower lip; plants an- nual 3. Cordylantlius 5. 6 SANTALACEAE. SANDALWOOD FAMILY 1. Comandra Nutt. False Toad-Flax 1. Commandra umbellata (L.) Nutt. var. pallida 'DC.) Jones. Commonly in sandy to well drained soil, April to August. Western United States and southern Canada. R.3IE., T.6N., Butte Co., 8 June 1967,915. I. Castilleja Mutis. Indian Paintbrush 1. Bracts yellow to yellowish; palea short, rarely over '/: the length of the corolla tube 2. C. longispica 1. Bracts red or purple; galea usually longer, some- times as long or longer than the length of the co- rolla tube 1. C. august ifolia BIOLOGICAL SERIES, VOL. 1 I NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 37 1. Castilleja angiistifolia (Nutt.) G. Don. Com- monly in sagebrush of lower elevations. May to July. Wyoming, Idaho and eastern Oregon. Webb Springs, Butte Co., 22 May 1967, 825. R.32E., T.3N., Bing- ham Co., 18 May 1967,765. 2. Castilleja longispica A. Nels. Sagebrush plains and lower meadows. May to July. Idaho, west to Ore- gon and California, east to Wyoming and Montana. Webb Springs, Butte Co., 22 May 1967, 826. South- west of Atomic City, Butte Co., 26 May 1967, 863. 2. CoUinsia Nutt. Blue-eyes Mary 1. CoUinsia parviflora Dougi. Variable in habitat from low elevations to higli in the mountains, March to July. Ontario to British Columbia, south to Michi- gan, Colorado to Arizona and California. Southwest of East Butte, Bingham Co., 18 May 1967, 771. T.IN., R.30E., 22May 1967,822. 3. Cordylanthus Nutt. Bird's-beak 1 . Cordylanthus ramosus Nutt. ex Benth. Usually growing in sagebrush or open places, June to August. Oregon and California, east to Wyoming and Colo- rado. NRTS, Goodwin, 139, (NRTS). 4. Linaria Mill. Toad flax; Butter and Eggs 1. Linaria vulgaris Hill. Scattered, mostly in waste places, June to September. Over much of temperate North America; Eurasia. Birch Creek, Clark Co., 5 September 1967, 1211. 5. Mimulus L. Monkey Flower 1. Mimulus nanus H. & A. Sandy or gravelly soil of the plains. May to April. Idaho, west to Washing- ton, Oregon and California, east and south from Mon- tana to Nevada. R.29E., T.3N., 21 May 1967, Butte Co., 818. 5. Basal rosette well developed; calyx 2.5 to 6 mm long; corolla 10 to 17 mm long, not whitish ven- trally; capsule 4 to 6 mm long .... 4. P. humilis 5. Leaves mostly cauline; calyx 5 to 9 mm long; co- rolla 16 to 25 mm long, whitish ventrally; cap- sule 5 to 8 mm long 6. P. radicosus 1. Penstemon cyaneus Pennell. Often in sagebrush and open places on the plains to middle elevations in the mountains, June to August. Idaho, east to Wyo- ming and Montana. Flood Control Area, Butte Co., 1 July 1967, 1099. 2. Penstemon deustus Dougl. Rocky, dry ground to middle elevations. May to July. Idaho, west from Washington to California, east to Montana and Wyo- ming, south to Utah and Nevada. T.IN., R.30E., Butte Co., 1 3 June 1967, 963. 3. Penstemon erianthenis Pursh. var. redactus Pen- nell & Keck. Mostly on dry ground in the plains and foothills. May to July. Idaho, west from British Co- lumbia to Oregon, east to North Dakota, Nebraska and Colorado. R.29E., T.7N., 8 June 1967, 920. 4. Penstemon humilis Nutt. ex Gray. Frequently dry, open places on the plains and foothills to well up in the mountains. May to July. Central Idaho, west to Washington, Oregon and California, south to Utah, Nevada, Wyoming and Colorado. R.29E., T.7N., Butte Co., 10 June 1967,932. 5. Penstemon pumilus Nutt. Commonly in sage- brush at lower elevations. May to July. Salmon River to the Snake River Plains, Idaho. R.31E., T.6N., Butte Co., 8 June 1967,908. 6. Penstemon radicosus A. Nels. Associated with sagebrush or in open places, from the plains to middle elevations in the mountains. May to July. Southwest- ern Montana, to central and eastern Idaho, Wyoming, Utah, Nevada and Colorado. East Butte, Bingliam Co., 24 May 1967,841. 6. Penstemon Mitch. Beardtongue 7. Verbascum L. Mullein 1. Leaves sharply toothed (sometimes entire); flow- 1. Verbascum thapsus L. A weedy plant, June to ers white or whitish; plants distinctly woody at September. Over much of temperate North America; the base 2. P. deustus Eurasia. Junction of Highway 20-26, Butte Co., 4 1. Leaves mostly entire; flowers blue to bluish- September 1967, 1205. white; plants not distinctly woody 2 2. Corolla large 20 to 40 mm long; palate glabrous . 8. Veronica L. Speedwell 1- P cyaneus 1. Plants from rhizomes; leaves petiolate; corolla 5 2. Corolla smaller, usually less than 25 mm long; to 10 mm wide; fruiting pedicels divaricate palate bearded 3 1 . y, americana 3. Calyx elongate, 7 to 1 1 mm long; corolla 18 to 1. Plants fibrous-rooted; leaves sessile and clasping; 30 mm long, inflated; staminode in at least some corolla about 5 mm wide; fruiting pedicels flowers exserted 3. P. erianthenis strongly ascending 2. V. anagallis-aquatica 3. Calyx 7 mm long or less; corolla mostly less than 1. Veronica americana Sctiwein. Moist places at 18 mm, not inflated, narrow; staminode included lower elevations to higlier in the mountains. May to 4 July. Over most of temperate North America. Birch 4. Plants small less than 2 dm tall; corolla glabrous Creek, Clark Co., 25 June 1967, 1056. Webb Springs, within 5. P. pumilus Butte Co., 1 July 1967, 1088. 4. Plants mostly over 2 dm tall; corolla bearded on 2. Veronica anagallis-aquatica L. Along streams or the palate 5 other moist habitats, June to September. Over much 38 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN of the United States; Europe. Big Lost River, Butte Co., 13 July 1967, 1144. SOLANACEAE. POTATO OR NIGHTSHADE FAMILY 1 . Corolla over 2 cm long, funnelform or salver- form; fruit a capsule 1 . Nicotiana 1. Corolla less than 2 cm long, rotate to campanu- late; fruit a berry 2. Solamim 1 . Nicotiana L. Tobacco 1. Nicotiana attenuata Torr. Commonly in sandy soil, moist or dry rocky slopes and hills, June to Sep- tember. Idaho, west from British Columbia to Califor- nia, east to Texas. Big Lost River, Butte Co., 13 July 1967, 1151. 2. Solanum L. Nightshade 1. Solanum dulcamera L. Along streams or in thickets. May to September. Common in the United States and southern Canada; Eurasia. Birch Creek, Clark Co., 5 September 1967, 1218. Poisonous. UMBELLIFERAE. PARSLEY FAMILY 1. Fruit bristly -hispid; leaves with well defined leaf- lets, not dissected 3. Osmorhiza 1. Fruit not bristly-hispid; leaves more or less dis- sected, without broad leaflets 2 2. Mature fruit with both lateral and dorsal wings, ovoid to oblong 1 , Cymoptems 2. Mature fruit with lateral wings only, linear to obricular 2. Lomatium fined vaneties foenicitlacais (T. & G.) Cronq. and cal- careus (T. & G.) Cronq. 2. Lomatium Raf. Desert Parsely 1. Ultimate leaf segments (at least some) over 1 cm long 2 1. Ultimate leaf segments less than 1 cm long 2. L. foeniculaceum 2. Some of the ultimate leaf segments 3 mm wide and over 1 cm long; taproot elongate and seldom thickened 3. L. tri tenia titni 2. Ultimate leaf segments less than 3 mm wide and barely over 1 cm long; taproot very large and woody 1 . L. dissectum 1. Lomatium dissectum (Nutt.) Math. & Const, var. multifidam (Nutt.) Math. & Const. Rocky plains and slopes, April to June. British Columbia and Al- berta to Wyoming, west through central Idaho to Ore- gon and Nevada. Webb Springs, Butte Co., 24 May 1967,842. 2. Lomatium foeniculaceum (Nutt.) Coult. & Rose var. macdougalii (Coult. & Rose) Cronq. Valleys and plains to higlier in the mountains, April to Au- gust. Central and southern Idaho, west to Oregon to Arizona and Nevada, east to Montana to the southern Great Plains. T.4N., 4.28E., Butte Co., 21 May 1967, 810. 3. Lomatium triternatum (Pursh) Coult. & Rose From the dry plains to higlier, moist habitats. May to July. Alberta and British Columbia to Colorado, Utah and California. T.4N., R.28E., Butte Co., 21 May 1967,809. 1 . Cymopterus Raf. 1. Plants mostly with 1 to 2 pseudoscapes; leaves forming a flat rosette 1. C. acaulis 1. Plants caespitose, with several leafy stems 2 2. Flowers white; calyx teeth wanting; plants acau- lescent 2. C. bipinnatus 2. Flowers yellow; calyx teeth evident; plants cau- lescent 3. C. terebinthinus 1 . Cymopterus acaulis (Pursh) Raf. Dry plains and valleys, April to May. Central Saskatchewan and west- ern Minnesota to Colorado, Montana and central Ida- ho to Oregon. Southwest to Howe, Butte Co., 1 2 April 1939, Davis, s.n., (IDS). 2. Cymopterus bipinnatus Wats. Open rocky places, from the foothills to above timberline. May to July. Montana and Wyoming, to central Idaho, Ore- gon, Nevada and Utah. Near Howe, Butte Co., July 1939,0. Mays, s.n., (IDS). 3. Cymopterus terebinthinus (Hook) T. & G. Dry, sandy or rocky ground on the plains and foot- hills, April to June. Central Idaho, east to Montana, west to Oregon and Washington. R.29E., T.3N., 6 June 1967, 881. Our plants belong to the poorly de- 3. Osmorhiza Raf. Sweet Cicely 1. Osmorhiza chilensis Hook & Arm. Mainly in moist wooded areas, April to June. Idaho, north and west from Alaska to California, east from Alberta, South Dakota, Colorado to Arizona; Chile and Argen- tina. Webb Springs, Butte Co., 1 July 1967, 1083. URTICACEAE. NETTLE FAMILY 1. Urtica L. Nettle 1. Urtica dioica L. ssp. gracilis var. holosericea (Nutt.) Hitchc. Sagebrush plains to mountain slopes. May to September. Widespread. Big Butte, Butte Co., 27 June 1950, AEC, 137, (IDS). I VERBENACEAE. VERBENA FAMILY I. Verbena L. Verbena , 1. Verbena bracteata Lag. & Rodr. A weedy plant 4 along roads, waterways, and disturbed areas. May to j September. Througliout most of temperate North ,j America. Big Lost River, Butte Co., 13 July 1967, 1143. BIOLOGICAL SERIES, VOL. I 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 39 VIOLACEAE. VIOLET FAMILY 1. Viola L. Violet 1. Viola mittalUi Pursh Commonly in sagebrush, from the plains to the mountains, April to July. Brit- ish Columbia to California, east to the central United States. Webb Springs, Butte Co., 22 May 1967, 828. CLASS ANGIOSPERMAE SUBCLASS MONOCOTYLEDONEAE Key to the Families I. 3 Perianth lacking or inconspicuous, of bristles or scales, not petallike (see also Junca- ginaceae and Juncaceae) 2 Perianth commonly of 2 series, the inner us- usually petaloid (except in Juncaceae and Juncaginaceae) 4 Flowers sessile in the axils of chaffy, imbri- cate bracts; plants with jointed stems and sheathing leaves, grasslike; fruit 1 -seeded. . .3 Flowers not sessile in the axils of chaffy bracts; fruit of 3 to 6 carpels JUNCAGINACEAE, p. 42 Stems round or flat and mostly hollow; leaves 2-ranked; anthers attached at the mid- dle GRAMINEAE, p. 39 Stems mostly triangular and solid; leaves 3-ranked; anthers attached at the base CYPERACEAE, p. 39 4. Perianth segments greenish or brownish, not showy; plants somewhat grasslike 5 4. Inner perianth divisions showy and petallike; plants not grasslike 6 5. Carpels 3 to 6, separating at maturity; flow- ers in racemes or spikes JUNCAGINACEAE, p. 42 5. Carpels 1 to 3, united; inflorescence panicu- late or corymbose JUNCACEAE, p. 6. Pistils several to many in a head or ring, each ovary with 1 -carpel ALISMACEAE,p. 6. Flowers with a single pistil, of several united carpels 7 7. Stamens 3 or fewer; ovary inferior 8 7. Stamens commonly 6; ovary superior LILIACEAE,p. 8. Perianth irregular; stamens 1 or 2 ORCHIDACEAE,p. 8. Perianth usually regular; stamens commonly 3 IRIDACEAE,p. 42 42 39 43 ALISMACEAE. WATER PLAINTAIN FAMILY 1. Alisma L. Waterplantain 1. Alisnm plant ago-aqiiatica L. In water and wet places, June to September. Found over most of the United States and Canada. Sinks, Butte Co., 5 Sep- tember 1967, 1222. CYPERACEAE. SEDGE FAMILY 1. Achene enclosed by an inner scale (perigynium); flowers imperfect; perianth lacking ... .1. Carex 1. Achenes subtended by 1 outer flat scale, not en- closing the achene; some flowers of each spikelet perfect; perianth of 1 to 12 bristles 2. Eleocharis 1. Carex (Rupp.) L. Sedge I. Carex douglasii Boott. Open more or less dry ground, April to August. Manitoba to British Colum- bia, south to California and New Mexico. Shell Road, Butte Co., 25 May 1967, 848. There are probably more species in this genus and in the family which occur on the site particularly in the sinks area. This area, during the period of this work, received an un- usual amount of runoff and was quite inaccessible. 2. Eleocharis R. Br. Spikerush 1. Eleocharis palustris (L.) R. & S. Moist or wet ground. May to August. Alaska and British Columbia, south to California and Mexico. Five miles S.E. of Big Butte, 24 June 1950, AEC, 172, (IDS). GRAMINEAE. GRASS FAMILY 1 . Spikelets with one perfect terminal floret and a sterile or staminate floret below; articulating be- low the spikelet 5. Echinochloa 1. Spikelets I to many flowered, reduced florets, when present, above the perfect tTorets; articula- tion above the glumes 2 2. Spikelets sessile or subsessile in spikes or spike- like racemes 3 2. Spikelets pedicellate in open or contracted pan- icles 6 3. Spikelets solitary at each node of the rachis .... 1 . Agropyron 3. Spikelets more than one at each node of the ra- chis (Elymus with part of the spike with solitary spikelets) 4 4. Spikelets 3 at each node of the rachis, usually 1 -flowered, the lateral ones pedicelled, usually re- duced to awns 8. Hordeum 4. Spikelets 2 at each node of the rachis, 2 to 6- flowered 5 5. Rachis readily disarticulating, glumes subulate and produced into long divergent awns 14. Sitanion 5. Rachis continuous, glumes broad or narrow, acute to aristate, entire 6. Elymus 6. Spikelets 1-flowered 7 6. Spikelets 2 to many flowered 11 7. Fruit indurate, terete, awned; callus well devel- oped, bearded 8 7. Fruit thin or Arm; callus not well developed. . .10 8. Awn trifid, the lateral ones sometimes short .... 3. Aristida 40 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 8. Awn simple 9 9. Awn not twisted (early deciduous), usually less than 3 to 4 times as long as the fruit; callus mostly obtuse 11. Oryzopsis 9. Awn twisted, persistent, and bent, more than 4 times longer than the fruit; callus sharp pointed . 16. Stipa 10. Glumes longer than the lemma; palea small or wanting 2. Agrostis 1 0. Glumes not longer than the lemma, mostly shorter; palea well developed ... 15. Spurobohis 1 1. Glumes at least as long as the lowest floret; lem- mas awnless or awned from the back 9. Koeleria 1 1. Glumes shorter than the first floret, lemmas awn- less or awned from the tip or a bifid apex ... 1 2 12. Plants large, culms 2 m or more tall; panicles large, plumelike 12. Phragmites 12. Plants smaller, culms less than 1 .5 m tall 13 13. Lemmas awnless, mostly rounded on the back 14 13. Lemmas mucronate or awned, keeled at least to- ward the summit 15 14. Glumes papery; upper florets reduced 10. Melica 14. Glumes not papery; upper florets similar to the lower ones; blades with boat shaped tips \i. Poa 15. Lemmas awned from between the teeth of the minutely bifid apex 4. Bromiis 15. Lemmas awnless; plants dioecious; stigmas sub- plumose 7. Hesperochloa I . Agropyron Gaertn. Wheatgrass 1 . Plants with creeping rhizomes 2 1. Plants without creeping rhizomes 5 2. Lemmas awned, the awn divergent at maturity; lemmas pubescent 1. A. albicans 2. Lemmas awnless or with a short straight awn . . .3 3. Glumes rigid, tapering into a short awn 5. A. smithii 3. Glumes not rigid, acute or abruptly awn pointed 4 4. Joints of the rachilla villous; lemmas villous .... 2. A. dasystachyum 4. Joints of the rachilla glabrous or scaberlous .... 4. A. riparium 5. Spikelets much compressed, crowded on the rachis 3. A. desertorum 5. Spikelets not much compressed, not crowded on the rachis 6 6. Spikelets awnless or nearly so; lemmas glabrous . 8. A. trachycaulum 6. Spikelets awned 7 7. Awn straight or nearly so, blades 3 to 8 mm wide; glumes 4 to 7-nerved 7. A. subsecundum 7. Awn divergent when dry; blades 1 to 2 mm wide; glumes about 4-nerved 6. A. spicatum 1. Agropyron albicans Scribn. & Smith. Plains and dry hills. South Dakota to Alberta and Idaho, Colora- do and Utah. R.30E., T.5N., Butte Co., 13 June 1967,986. 2. Agropyron dasystachyum (Hook.) Scribn. Plains and sandy shores. May to July. Michigan to British Columbia, south to Illinois, Nebraska, Colora- do, Nevada and Oregon. By cave near EBR. II Reac- tor, Bingham Co., 30 June 1967, 1061. T.6N., R.32E., Jefferson Co., 28 July 1967, 1178. T.4N., R.31E., Butte Co., June 1965, Harniss, (UTC). 3. Agropyron desertorum (Fish.) Schult. Russia; widespread in the western United States, June to August. R.30E., T.5N., Butte Co., 13 June 1967, 985. 4. Agropyron riparium Scribn. & Smith Dry or moist ineadows and hills. May to July. Western United States and southern Canada. EBR. I, Butte Co., 4 July 1967, 1124. 5. Agropyron smithii Rydb. Moist, usually alka- line soil, June to August. Widespread in east and west- ern North America and southern Canada. R.30E., T.5N., Butte Co., 13 June 1967,987. 6. Agropyron spicatum (Pursh) Scribn. & Smith Plains, canyons, and slopes, June to August. Alaska, south to California, east to New Mexico, South Dako- ta and Michigan. Webb Springs, Butte Co., 13 June 1967,974. 7. Agropyron subsecundum (Link) Hitchc. var. andimim (Scribn. & Smith) Hitchc. Mountain mea- dows and slopes. Montana to Washington, south to Colorado and Nevada. East Butte, Bingham Co., 22 June 1967, 1011. 8. Agropyron trachycaulum Malte. Alaska to Lab- rador, south to California, Mexico, West Virginia and Missouri. T.2N., R.29E., Butte Co., 4 July 1967, 1124. T.6N., R.32E., Jefferson Co., 28 July 1967, 1179. 2. Agrostis L. Bentgrass 1. Agrostis alba L. Cultivated extensively and es- caping, June to September. Mostly in the cooler parts of the United States, Eurasia. Big Lost River, Butte Co., 29 July 1967, 1191. 3. Aristida L. Three-awn 1. Aristida fendleriana Steud. Plains and hills, June to July. North Dakota to Montana, south to Texas, Nebraska and California; Mexico. Tractor Flat, Jefferson Co., 22 July 1967, 1 197. \ B. inermis 9 4. Bromus L. Bromegrass Plants perennial Plants annual Lemmas narrow, with a sharp callus, gradually acuminate, bifid; awns usually 1.5 cm long or more 2. B. tectorum J i BIOLOGICAL SERIES, VOL. 1 I NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 41 2. Lemmas broad, rounded above, not acuminate; sheaths glabrous 3. B. secalinus 1. Bromus inermis Leyss. Cultivated, escaping, June to August. Europe; northern half of the United States. R.30E., T.5N., Butte Co., 13 June 1967,984. 2. Bromus secalinus L. A weed, throughout most of the United States, June to July. Birch Creek, Clark Co., 25 June 1967, 1054. 3. Bromus tectonim L. Along roadsides and waste places, April to June. Sporadic in the United States as far as South Carolina and Texas. R.28E., T.4N.. Butte Co., 21 May 1967,817. 5. Echinochloa Beuv. 1. Echinochloa cmsgalli (L.) Beauv. Moist places, June to October. Widespread. Sec. 33, T.5N., R.30E., Butte Co., 26 July 1956, McBride, 74, (NRTS). 6. Elymus L. Wildrye I . Plants with slender, creeping rhizomes 3. E. flavescens 1. Rhizomes wanting or short and stout in E. cin- ereus; plants tufted 2 2. Spike 1 to 2 cm thick; spikelets usually 2 to 5 at a node; culms many 2. E. cinereus 2. Spike more slender; some or most of the spike- lets solitary at the nodes; culms few 1 . E. ambiguus 1. Elymus ambiguus Vasey & Scribn. Open slopes. May to July. Montana, Colorado, Idaho and Utah. R.30E., T.6N., Butte Co., 1 1 June 1967, 946. 2. Elymus cinereus Scribn. & Merr. Moist river banks or dry slopes and plains, June to July. Minne- sota to British Columbia, south to California and Colorado. Flood Control Area, Butte Co., 1 July 1967, 1110. 3. Elymus flavescens Scribn. & Smith Sand dunes, June to July. Eastern Washington and Oregon, Idaho, South Dakota. R.31E., T.6N., Butte Co., 28 July 1967, 1180. 7. Hesperochloa (Piper) Rydb. 1. Hesperochloa kingii (S. Wats.) Rydb. Mountains and hills, June to August. Oregon to California, east to Montana. Nebraska and Colorado. Webb Springs, Butte Co., 13 June 1967,971. 8. Hordeum L. Barley 1. Hordeum jabatum L. Moist soil, especially in saline areas, June to August. Newfoundland to Alas- ka, south to Texas. California, and Mexico. Junction 88-22, Butte Co., 13 June 1967, 983. 9. Koeleria Pers. Junegrass 1. Koeleria cristata (L.) Pers. Prairie, open woods, and sandy soil. May to July. Ontario to British Columbia, south to Delaware, Louisiana, Mexico and California. East Butte, Bingham Co., 22 June 1967, 1015. Webb Springs, Butte Co., 15 July 1967, 1155. 10. Melica L. Melicgrass; Oniongrass 1. Melica bulbosa Geyer. Woods and hills. May to July. British Columbia, south to California, Colorado and Texas. Big Butte, Butte Co., 13 June 1967,982. 11. Oryzopsis Michx. Ricegrass 1. Oryzopsis hymenoides (R. & S.) Richer. Des- erts and plains. May to June. Manitoba to British Columbia, south to California, Texas and New Mexico. Two miles west of Big Lost River Bridge, Butte Co., 6 June 1967,887. 12. Phragmites Trin. Common reed 1. Phragmites communis Trin. Marshes, banks, and other moist places, August to September. Cosmo- politan. Should be looked for on the Big Lost River and in the Sinks area. 13. Poa L. Bluegrass 1. Spikelets distinctly compressed, the glumes and lemmas keeled 2 1. Spikelets little compressed, narrow, much longer than wide; lemmas convex on the back 3 2. Creeping rhizomes present; lemmas webbed at the base 5. P. pratense 2. Creeping rhizomes wanting; lemmas not webbed at the base 2. P. fendleriana 3. Lemmas crisp— puberulent on the back toward the base 4 3. Lemmas glabrous or minutely scabrous, but not crisp-puberulent 5 4. Culms slender mostly less than 30 cm tall; num- erous short innovations at base; blades usually folded 6. P. secunda 4. Culms stouter, usually more than 30 cm tall; in- novations usually not numerous ... 1. P. canbyi 5. Sheaths scaberulous; ligules long, decurrent .... 4. P. nevadensis 5. Sheaths glabrous; blades involute 3. P. juncifolia 1. Poa canbyi (Scribn.) Piper Dry soil. Yukon to Michigan, south to Nebraska, Arizona and California. T.2N., R.29E., Butte Co., 4 July 1967, 1125. Eight miles east of Howe, Butte Co., 27 June 1941, Hull 265, (IDS). 2. Poa fendleriana (Steud). Vasey Rocky slopes and mesas. May to August. British Columbia to Manitoba, south to South Dakota, Nebraska, and Idaho to Texas and California; New Mexico. R.29E., T.3N.. Butte Co., 23 May 1967, 833. 3. Poa juncifolia Scribn. Saline soil. May to July. British Columbia to Montana, south to California and Colorado. Desert, Butte Co., June 1950, AEC, 41, (IDS). 4. Poa nevadensis Vasey ex Scribn. Moist places, June to July. Montana to eastern Washington and the Yukon, south to Colorado, Arizona, and California; Maine. East Butte, Bingham Co., 3 July 1950, AEC 42 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Project, 170 (IDS). T.3N., R.29E., Butte Co., 7 June 1967,892. 5. Poa prateuse L. Open woods, meadows and plains, May to October. Widespread throughout the United States and northward; Europe. R.29E., T.3N., 21 May 1967,833. 6. Poa seciuida Presl. Plains, dry woods and rocky slopes. North Dakota to the Yukon, south to Cali- fornia, New Mexico, Nebraska; Chile. T.2N., R.29E., Butte Co., 4 July 1967, 1125. 14. Sitanion Raf. Squirreltail 1. Sitanion hystrix (Nutt.) J. G. Smith. Dry hills, plains and woods. May to July. South Dakota to British Columbia, south to California, Texas and Missouri. T.6N., R.31E., Butte Co., June 1957, McBride, s.n., (UTC). 15. Sporobolus R. Br. Dropseed 1. Sporobolus cryptandms (Torr.) Gray Sandy open ground, June to July. Alberta to Ontario, south to Washington, North Carolina, Indiana, Louisiana, southern California, and New Mexico; Maine. R.31E., T.4N., Butte C, 4 July 1 967, 1 1 20. 16. Stipa L. Needlegrass 1. Awn plumose below, the hairs ascending or spreading 2 1 . Awn scabrous or nearly glabrous, rarely appress- ed-hispid, not plumose 3 2. Ligules 3 to 6 mm long, hyaline 4. S. thurberiana 2. Ligules minute, mostly hairy. . . 3. S. occidentalis 3. Lemmas mostly more than 1 cm long, glabrous or sparsely pubescent above the callus 2. S. coma t a 3. Lemmas 7 mm long or less; distinctly pubescent on the upper part at least 1 . S. columbiana 1 . Stipa columbiana Macoun. Dry plains and woods. May to August. South Dakota to Yukon Territory, south to California and Texas. Webb Springs, Butte Co., 1 July 1967, 1073. 2. Stipa comata Trin. & Rupr. Plains and hills. May to July. Indiana to Yukon Territory, south to California and Texas. R.28E., T.4N., Butte Co., 21 May 1967,816. 3. Stipa occidentalis Thurb. Plains, hills, and woods. May to August. Wyoming to Washington, Arizona, and California. East Butte, Bingham Co., 22 June 1967, 1010 Webb Springs, Butte Co., 1 July 1967,1073. 4. Stipa thurberiana Pi[>er Rocky slopes and plains. May to June. Idaho to Washington and Calif- ornia. South of Middle Butte, Bingham Co., 22 June 1967, 1028. IRIDACEAE. IRIS FAMILY 1. IrisL. 1 . Iris missouriensis Nutt. Moist meadows, marshes and along streams. New Mexico and Calif- ornia, north to British Columbia and North Dakota, 5,000 to 10,000 feet. May to July. Birch Creek, Clark Co., 25 June 1967, 1040. JUNCACEAE. RUSH FAMILY 1 . Juncus L. Rush 1. Juncus balticus Willd. Moist places, June to August. Montana to Alaska, south to Kansas, New Mexico and California. Birch Creek, Clark Co., 25 June 1967, 1038. I JUNCAGINACEAE. ARROWGRASS FAMILY j 1. Triglochin L. Arrowgrass j 1. Triglochin palustris L. Greenland to Alaska, south to most of the northeastern United States and scattered in the western states, June to August. Birch Creek, Clark Co., 25 June 1967, 1039. > LILIACEAE. LILY FAMILY 1 . Perianth segments all alike or nearly so 2 1, Perianth segments unlike, 3 outer small and se- r^ paloid, 3 inner large and petaloid '■'■■. 2. Calochortus , 2. Flowers borne on scapose peduncles, in umbels; j when fresh, emitting an onionlike odor 1 . Allium 2. Flowers not in umbels, on scapose peduncles; on- ' ion odor not present 3 3. Plants from bulbs or corms; fruit a capsule; leaves linear-lanceolate 4 I 3. Plants from rootstocks; fruit a berry, leaves Ian- ' ceolate 4. Smilacina \ 4. Flowers small, usually 1 cm long or less; styles 3, i distinct; flowers whitish 5. Zygadenus * 4, Flowers usually over 1 cm long; style 1 , entire or ' divided; tlowers purple, brown, or yellow \ 3. Fritillaria , { 1. Allium L. Onion ■ 1 . Outer bulb coats of coarse anastomosing fibers; ovary crested with 6 low knobs 2 \ 1 . Outer bulb coats without fibers, never fibrous- reticulate; ovary inconspicuously 3 crested I . .4. acuminatum ; 2. Leaves commonly 2 per scape; tips of the inner '■ perianth segments spreading; flowers usually whit( 3. A. textile 2. Leaves usually 3 or more per scape; tips of the inner perianth segments erect; flowers commonly | pink 2. A. geyeri 1. Allium acuminatum Hook. Dry plains and hills, May to July. British Columbia south to Wyoming, Colorado and California. South of Middle Butte, BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 43 Bingliam Co., 22 June 1967, 1052. Arco Desert, Butte Co., June 1 950, AEC, 67, (IDS). 2. Allium geyeri S. Wats. Plains, meadows and slopes. May to June. Alberta, south to Texas, Arizona and Washington. South of Middle Butte, Bingham Co., 22 June 1967, 1025. R.29E., T.7N., Butte Co., 10 June 1967,935. 3. Allium textile Nels. & Macbr. Plains and dry hills. May to June. Saskatchewan to Alberta, south to New Mexico and Arizona. R.28E., T.4N., Butte Co., 20 May 1967,794. 2. Calochortus Pursh. Sego lily 1 . Sepals usually exceeding the petals; anthers linear 2. C. macrocarpus 1. Sepals shorter than the petals; anthers oblong to lanceolate 1 . C. bnineaunis 1 . Calochortus bivneauim Nels. & Macbr. Dry soil, June to July. Nevada and adjacent Oregon in the southeastern part, southwestern Idaho and south- western Montana. R.34E., T.5N., Jefferson Co., 20 June 1967, 1006. 2. Calochortus macrocarpus Dougl. Montana to British Columbia, south to Nevada and northeastern California, May to July. East Butte, Bingham Co., 23 July 1967, 1175. 3. Fritillaria L. 1 . Flowers dull purple, mottled with greenish- yellow; capsule sharply angled 1 . F. atropurpurea 1. Flowers bright yellow-orange, when young; cap- sule obtusely lobed 2. F. pudica 1. Fritillaria atropurpurea Nutt. Dry plains and slopes. May to June. North Dakota across central and southern Idaho, to Oregon, south to California and New Mexico, April to June. Six miles S.W. of Atomic City, Butte Co., 22 May 1967, 821. Webb Springs, Butte Co., 26 May 1967,876. 2. Fritillaria pudica (Pursh) Spreng. Plains and hillsides, March to June. Montana to British Colum- bia, south to Utah and California, April to June. Webb Spring, Butte Co.. 2 1 April 1 967, 75 1 . 4. Smilacina Desf. False Solomon's seal 1 . Smilacina stellata (L.) Desf. Moist, usually shaded ground, May to June. Throughout most of temperate North America; Europe, May to June. Webb Springs, Butte Co., 15 July 1967, 1159. Big Lost River, Butte Co., 1 3 July 1 967, 1 1 50. 5. Zygadenus Michx. Death camas 1. Flowers mostly paniculate; perianth-parts acute or acuminate; plants of dry areas in the foothills . 1 . Z. paniculatus 1. Flowers commonly racemose; perianth-parts ob- tuse or rounded at apex; plants of moist areas in the mountains 2. Z. venenosus 1. Zigademis paniculatus (Nutt.) Wats. Montana to Washington, south to California and New Mexico, 5,500 to 8,000 feet. May to June. R.28E., T.4N., Butte Co., 21 May 1967,811. 2. Zigadenus venenosus S. Wats. Montana to British Columbia, south to Utah and California, June to July. Big Butte, Butte Co., 1 July 1967, 1 103. ORCHIDACEAE. ORCHID FAMILY 1 . Corallorhiza (Hall.) Chat. Coral root 1 . Corallorhiza maculata Raf. Commonly in shaded woods, on rich humus soil. Alaska to Nova Scotia, south to Florida, New Mexico and California, at 6,000 to 9,000 feet. June to July. Webb Springs, Butte Co., 1 July 1967, 1062. CLASS GYMNOSPERMAE Key to the Families Leaves scalelike, opposite or in whorls, im- bricate; cones berrylike, the scales becoming fleshy at maturity 5. CUPRESSACEAE, p. 43 Leaves in fasciles or solitary, needle-shaped or narrowly linear; cones elongate (more or less), dry and woody at maturity 4. PINACEAE,p. 43 CUPRESSACEAE. CYPRESS FAMILY 1. Juniperus L. Juniper 1. Fruit reddish-brown or bluish; seeds 1 to 2; branchlets not flattened; leaves obtuse to acute 1 . J. osteosperma 1. Fruit briglit blue; seeds I to 3; branchlets flat- tened; leaves acute to acuminate 2. / scopulorum 1 . Juniperus osteosperma (Torr.) Little. Dry plains and foothills. Southwestern Wyoming, south- western Idaho, western Colorado, Utah, and western New Mexico to northern Arizona and the south- eastern part of Cahfornia. R.28E., T.4N., Butte Co., 21 May 1967,815. 2. Juniperus scopulorum Sarg. Commonly on rocky dry ridges at altitudes of 5,000 to 6,000 feet. May to June. Alberta and British Columbia, south to Arizona and Nevada. Webb Springs, Butte Co., 1 July 1967, 1068. PINACEAE. PINE FAMILY Leaves in clusters of 2 to 5, sheathed at the base, at least when young; cone scales very thick and woody I. Pinus I 44 1. Leaves single on the twigs, not sheathed at the base; cone scales not thick and woody 2. Pseudotsuga 1. Pinus L. Pine 1. Leaves 2 in a fasicle, each with 2 fibro-vascular bundles; cones 3 to 5 cm long \. P. contorta L Leaves usually 5 in a fasicle, each with 1 fibro- vascular bundle; cones 1 to 2.5 dm long 2. P. monticola \. Pinus contorta Dougl. var. munayana Engel. Hills and mountains, commonly in dry situations, April to June. Saskatchewan to Alaska, south to California and Colorado. Big Butte, Butte Co., 7 September 1967, 1227. 2. Pinus monticola Dougl. Moist areas at middle altitudes in the mountains. May to June. California, north to Idaho, western Montana and British Colum- bia. Big Butte, Butte Co., 7 September 1967, 1228. 2. Pseudotsuga Carr. Douglas fir 1. Pseudotsuga menziesii (Mirb.) Franco. Hills and mountains, commonly on deep soils of northern slopes, April to May. Alberta to British Columbia, south to California, Arizona and Western Texas. Webb Springs, Butte Co., 1 July 1967, 1079. BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 1 . Plants with leaves reduced to scales, sheath- ing the internodes at the joints; leaves sessile EQUISETACEAE, p. 44 2. Sporangia borne in hard sporocarps; leaves palmately 4-foliate; plants rooting in mud . . MARSILEACEA, p. 44 2. Sporangia borne on the veins of the pin- nules, leaves bi- or tri-pinnate; plants com- monly in crevices of cliffs and ledges POLYPODIACEAE, p. 44 EQUISETACEAE. HORSETAIL FAMILY 1 . Equisetum an'cnse L. In damp soil, along streams, woods and in sterile soil of roadsides. New- foundland to Alaska, south throughout most of the United States; Greenland; Eurasia. Birch Creek, Butte Co., 25 June 1967, 1042. MARSILEACEAE. PEPPERWORT FAMILY 1 . Marsilea vestita Hook. & Grey. Edges of ponds, ditches and rivers. Saskatchewan, British Columbia, south to California, Texas, Arkansas and South Dakota. Sinks, Butte Co., 5 September 1967, 1223. J I I i DIVISION PTERIDOPHYTA Key to the Families Plants with broad leaves, usually exceeding 2 cm in length; leaves petioled ..2 POLYPODIACEAE. FERN FAMILY 1. Woodsia oregana Eaton. Common in moist or dry shaded cliffs, on talus slopes and under rocks. Quebec to British Columbia, south to California, Arizona, New Mexico and Oklahoma. Webb Springs, Butte Co., 1 July 1967, 1076. % BIOLOGICAL SERIES, VOL. 1 1 NO. 4 FLORA OF THE NATIONAL REACTOR TESTING STATION 45 LIST OF REFERENCES Abrams, L. 1923. Illustrated Flora of the Pacific State.s. 1: Ferns to Birthworts. Stanford University, California: Stanford University Press. 1944. Illustrated Flora of the Pacific States. 2: Buckwheats to Kramerias. Stanford University, California: Stanford University Press. 1951. Illustrated Flora of the Pacific States. 3: Geraniums to Figworts; Stanford University, California: Stanford University Press. and R. S. Ferris. 1960. Illustrated Flora of the Pacific States. 4: Bigonia to Conipositae. Stan- ford University, California: Stanford University Press. Allrcd, D. M. 1968. Ticks of the National Reactor Testing Station. Brigham Young University Science Bulletin. 10(1): 1-29. Atwood, N. D. 1969. Flora of the National Reactor Testing Station. Idaho. Unpublished master's thesis. Department of Botany, Brigham Young Uni- versity. Provo, Utah. Bailey, L. H. 1949. Manual of Cultivated Plants. New York: The Macmillan Company. Barneby, R. C. 1952. A Revision of the North American Species of Oxvtropis DC. Proc. Cal. Acad. Sci. 27:117-312. Bates, R. G. 1963. Aeroradioactivity Survey and Arcal Geol- ogy of the N.R.T.S. Area. Idaho (ARSM-I). U.S. Geological Survey and Division of Biology and Medicine, U.S.A.E.C. Britton, N. L. 1905. The Flora of the Northern United States and Canada. New York: Herery Holt and Company. Clements, F. E. and E. S. Clements. 1920. Rocky Mountain Flowers. New York: H. W. Wilson Company. Cronquist, A. and D. D. Keck. 1957. A reconstitution of the Gsnus MachaeraiUhera. Britt. 9:231-239. Davis. R. J. 1952. Flora of Idaho. Wm. C. Brown Company, Dubuque, Iowa. __. 1968. Idaho State College. Personal Communication, Eastwood, Alice. 1934. A Revision of Arctostaphylos. Leatl. West. Bot. 1:105-127. Ewan, J. 1945. A Synopsis of the North American Species of Delphinium. Univ. of Colo. Phys. & Biol. Studies 2(2):55-244. French, N. R., R. McBride, and J. Detnier. 1965. FertiUty and Population Density of the Black-tailed Jack- rabbit. J. Wildl. Mgmt. 29(I):14-26. Hanson. C. A. 1962. Perennial Atriplex of Utah and the Northern Deserts. Unpublished master's thesis, Brigham Young University, Provo, Utah. Harniss, R. O. 1968. Vegetational Changes Following Live- stock Exclusion on the National Reactor Testing Station, Southeastern Idaho. Unpublished master thesis in Plant Ecology, Utah State University, Logan, Utah. Harrington, H. D. 1954. Manual of the Plants of Colorado. Sage Books, Denver. Hitchcock, A. S. 1950. Manual of the Grasses of the United States. U. S. Dept. of Agriculture Misc. Publ. 200. Hitchcock. C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1955-1964. Vascular Plants of the Pacific Northwest. Parts 2-5. Univ. of Washington Press, Seattle. Holmgren, A. H. and J. L. Reveal. 1967. Checklist of the Vascular Plants of the Intermounlain Region. U. S. Forest Service Res. Paper Int. 32. Ogden, Utah. Jackson, B. D. 1881. Guide to the Literature of Botany. London: Longmans, Greene and Company. . 1895. Index Kewensis. Oxford: The Clarendon Press. Johnston, I. M. 1924. Studies in the Boraginaceae. A Synopsis of the American Native and Immigrant Borages of the Subfamily Boranginoideae. Contr. Gray Herb. 70:46. Kearney. T. H. and R. H. Peebles. 1960. Arizona Flora. Uni- versity of California Press: Berekeley and Los Angeles. Lanjouw, J. and F. A. Stafleu. 1964. Index Herbariorum. Part 1. 5th ed. Utrecht, Netherlands: Kemink en Zoon. Ley, A. 1943. A Taxonomic Revision of the Genus Hoi- odiscus. Bull. Torrey Bot. Club 70:275-288. Maguire, B. 1951. Arenaria in America North of Mexico- A Conspectus. Amer. Midi. Natur. 46:493-511. McBride, R. 1968. NRTS Vegetation Type Map. In: AUred, D. M. Ticks of the National Reactor Testing Station. Brigham Young University Sci. Bull. Bio. Ser. 10:1-29. McKelvey, S. D. 1955. Botanical Exploration of the Trans- Mississippi West, 1790-1850. Arnold Arboretum of Harvard University. Janiacia Plain, Mass. Munz, P. and D. D. Keck. 1959. A Cahfornia Flora.University of California Press: Berkeley and Los Angeles. Nace, R. L., M, Deutsch, and P. T. Voegeli. 1956. Geography, Geology and Water Resources of the National Reactor Testing Station, Idalio: Part 2, Geography and Geology. U.S. Geological Survey, Admin. Rept. IDU-22033-USGS. Nelson, A. 1912. Contributions from the Rocky Mountain Herbarium. XI. New Plants from Idaho. Bot. Gaz. 54:136-151. Ornduff, R. 1968. Index to Chromosome Numbers for 1966. Regnum Vegetaile Vol. 55. Over, W. H. 1932. Flora of South Dakota. Vermillion: Uni- versity of South Dakota. Ownbey, M. 1940. A Monograph of the Genus Calochortus. Ann Mo. Bot. Card. 27:371-560. ^ 1950. The Genus Allium in Idaho. Research Studies of the State College of Washington. 18:3-39. Payson, E. B. 1927. A Monographic Study of Thelypodium and its Immediate Allies. Ann. Mo. Bot. Gard. 9. 46 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Pennell, F. W. 1936. Travels and Scientific Collections of Thomas Nuttall. Bartonia 18:10-61. Porter, C. L. 1959. Taxonomy of Flowering Plants. Freeman and Company, Inc. San Francisco. Pretzel, G. A. 1872. Thesaurus Literature Botanicae. Lipsiae; F. A. Brockhuas. Raven, P. H. 1962. The Systematics of Oenothera subgenus Chylismia University of California Press: Berkeley and Los Angeles. 1964. The Generic Subdivisions of Onagraceae, Tribe Onagreae. Britt. 16(2): 276-288. Reveal, J. L. 1968. Brigham Young University. Personal Communication. - 1969. A revision of the genus Eriogonum (Polygonaceae). Unpublished doctoral dis- sertation. Department of Botany, Brigham Young University, Provo, Utah. Rickett, H. W. 1944. The Classification of Inflorescences. Bot. Rev. 10:187-231. Rollins, R. C. 1941. Monographic Study of A ra bis in Western North America. Rhodora 43:289-325, 348-411, 411-481. Ross, C. P. and J. D. Forrester. 1958. Outline of the Geology of Idaho: Idaho Bur. Mines GeoL Bull. No. 15. Rossbach, G. B. 1958. The Genus Erysimum (Cruciferae) in North America North of Mexico -A Key to the Species and Varieties. Madrono 14:261-267. Rydberg, P. A. 1922. Flora of the Rocky Mountains. Intelligencer Printing Company: Lancaster, Pa. Sargent, C. S. 1955. Manual of the Trees of North America. Vol. 1-2. Dover Publications, Inc.: New York. Schwarten, L. and H. W. Rickett. 1958. Abbreviations of Titles of Serials Cited by Botanists. Bull. Torrey Club 85:277-300. Stearns, H. T., L. Crandall, and W. G. Steward. 1938. Geology and Ground-Water Resources of the Snake River Plain in Southeastern Idaho. U. S. Geol. Survey Water-Supply Paper 774. . L. L. Bryan, and L. Crandall. 1939. Geology and Water Resources of the Mud Lake Region, Idaho. U. S. Geol. Survey: Water-Supply Paper 818. Tidestrom, I, 1925. Flora of Utah and Nevada. Contrib. U. S. Natl. Herbarium, Vol. 25. Washington, D. C. Turner. B. L. 1956. A Cytotaxonomic Study of the Genus Hvmenopappus. Rhodora 58:163-186,208-242, 253-269, 295-308. Welsh, S. L., M. Trcshow, and G. Moore. 1964. Guide to Common Utah Plants. Brigham Young University Press, Utah. Wherry, E. T. 1955. The Genus Phlox. Morris Arboretum Monogr. Ill, 174 pp. Yanskey, G. R., E. H. Markee Jr., and A. P. Richter. 1966. Climatography of the National Reactor Testing Station. Air Res. Field Res. Office, Idaho Falls, Idaho. INFORMATION FOR CONTRIBUTORS Contributions to the Science Bulletin should be pri- Tiarily monographic in nature. For the most part only Tianuscripts of approximately forty or more typewritten Dages will be accepted. Papers will be published approximately in the order hat they are received, pending availability of funds. \uthors must arrange for financing their publications, ;xcept that the University Press and the library of 3riglram Young University will share the publication ;ost proportionately to their needs for copies of the irticle printed. In the preparation of manuscripts, authors are re- }uested to follow the Style Manual for Biological Jour- lals, American Institute of Biological Sciences, 2000 P Street, N.W., Washington, D.C., 20006. Manuscripts nust be typed on one side of the paper only, double- paced with ample margins. Footnotes should be voided. To facilitate review by referees, send to the editor the )riginal manuscript and one carbon copy, together with he illustrations. Copies of the original illustrations may le submitted, but should be of quality equal to the orig- nals. Illustrations should be referred to as figures except or materials requiring inserts of special paper, which nay be called plates. Illustrations should be so designed s to fit when reduced into a one-column or full-page vidth. Special care must be taken to allow for proper eduction in lettering (i.e., a 50% reduction of the figure neans also a 50% reduction in the lettering.) Photo- iraphs should be of a glossy finish, unblurred, and show- 'ng sharp contrast. Line drawings should be made with ilack ink on heavy white drawing paper, blue tracing loth, or blue-ruled coordinate paper. Use the same bbreviations on line drawings as in text. Line drawings nust be equivalent to a professional draftsman's work. Original drawings are preferable to photographs, even if they are large. Illustrations (line drawings or photo- graphs) should be numbered consecutively throughout the paper, and the approximate place of insertion should be indicated in the margins of the manuscript pages. Captions for illustrations should be assembled on a sep- arate sheet, and each plate of figures must have its corre- sponding figure number pencilled lightly on the back. Illustrations and cuts will be destroyed unless their re- turn is requested when proof is returned to the editor. A table title should be a short, concise statement of what the table purports to show, and should not include information necessary to the interpretation of the table. Every column in the table should carry a head identify- ing the data in that column; the measure in which the data are given should be indicated at the head of each column. Tables should not be used when the same infor- mation can be given in a few lines of text, and should not duplicate information in text, graphs, or charts. Symbols (asterisk, dagger, etc.) should be used to indi- cate footnotes to tables, with footnotes on the same page as the table. An abstract of less than four percent of the length of the paper should be prepared. This summary should be understandable without reference to the body of the manuscript. The abstract must be on sheets separate from the manuscript. Proof should be corrected immediately on receipt and returned to the editor. Authors should leave forwarding addresses if they move from the address sent with the manuscript. Reprints should be ordered when the proof is re- turned. Address all manuscripts to Stanley L. Welsh, Depart- ment of Botany, Brigham Young University, Provo, Utah 84601. ''\ 1 i e)"^"" nV^^ Brigham Young University Science Bulletin DEC 1 ' UMlVEHSiTY, MITES AND LICE OF THE NATIONAL REACTOR TESTING STATION by Dorald M. Allred BIOLOGICAL SERIES— VOLUME XII, NUMBER 1 OCTOBER 1970 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIt BIOLOGICAL SERIES Editor: Stanley L. Welsh, Department of Botany, Brigham Young University, Prove, Utah Members of the Editorial Board: Vernon J. Tipton, Zoology Ferron L. Anderson, Zoology Joseph R. Murdock, Botany Wilmer W. Tanner, Zoology Ex officio Members: A. Lester Allen, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Chairman, University Publications The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from PubUcation Sales, Brigliam Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for library exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 8460 1. Brigham Young University Science Bulletin MITES AND LICE OF THE NATIONAL REACTOR TESTING STATION by Dora Id M. All red BIOLOGICAL SERIES— VOLUME XII, NUMBER 1 OCTOBER 1970 I MITES AND LICE OF THE NATIONAL REACTOR TESTING STATION 1 Dorald M. Allred* INTRODUCTION This is the fourth in a series of reports on arthro- pods of the National Reactor Testing Station in Idaho (Allred, 1968 a & b, 1969). A fifth paper by Atwood (1970) deals with the plants of the sites where the arthropods were studied. The initial publication of the series (Allred, 1968a) discusses the location and physical characteristics of the NRTS, the study areas and procedures, and lists the kinds of vertebrates ex- amined. The reader is referred to that report for de- tails not included herein. The National Reactor Testing Station is situated approximately 30 miles west of Idaho Falls in south- eastern Idaho. The central and southern parts are typified by basalt flows which are exposed in some areas. The northern section is primarily lake and eolian deposits, and exposed basalt flows are less evi- dent. Annual precipitation averages less than 10 inches, and the vegetation is ciiaracteristic of the cool, northern desert, shrub-type biome. Twelve principal study sites established on the basis of their predominant vegetation were studied periodically: (1) C h r y so t lia m uu s-A r t e misia-gtasses; (2) Artemisia- Chiysollmmmis-grasses; (3) Elynnis: (4) Oryzopsis- Sripa: (5) Junipems; (6) Chn'sothammis-Tetra- dymia-Artemisia; (7) Chrysothammis-Artemisia- Eurotia: (8) Artemisia- A triplex: (9) Cheiiopodiiim- Eiirotia: (10) Artemisia-Opuntia; (11) Chrysotham- nits-graises-Tetraclymia; (12) Jimiperus-Chrysotham- inis-Eurotia-Artemisia. Twenty-eight other sites simi- lar to the major ones but with minor variations of plant associations occupying smaller geographic areas were studied less frequentlv. Most of the mammals were captured with live- catch or break-back traps. Rabbits, carnivores, and birds were shot, and reptiles were captured by hand or in can pit-traps. Ectoparasites were retrieved from their hosts by the cooling and warming method de- scribed by Allred ( 1968a). Financial support for these studies was provided by U.S. Atomic Energy Commission Contract AT(1 1-1)-1559 with Brigham Young University. Lo- gistics (in part) were provided through the AEC Oper- ations Office at Idaho Falls, Idaho. The chigger mites and many of the mesostigmatids were identified by Mr. Morris Goates. and the lice by Dr. W. L. Jellison. PARASITE-HOST ASSOCIATIONS Entries in the listings below may be interpreted by using the first two lines of the listing of mite-host as- sociations as an example (each specific entry is indi- cated in boldface type); Androlaelaps leviculus ( Mar-Aug) 1 3 dny Id 17 9: Androlaelaps leviculus = the species of parasite collected. (Mar-Aug) = the inclusive period of time when the parasites were found. 13 dny Id 17 9 = the total number of parasites of each develop- mental stage and sex that were collected (la = larva, pny = protonymph, dny = deuto- 'bYU-AEC Report No. COO-1559-5. *Department of Zoology, Brigham Young University, Provo, Utah. nymph, im = immature, d = adult male, 9 = adult female). Dipodomys ordii 2 (808) 1.5-9: jun Jul: Dipodomys ordii = a specific host on which parasites of the species listed above it were found. 2 = number of hosts infested with parasites of that particular species. (808) = total number of hosts ex- amined. 1.5 = the parasite-host index (total number of parasites of that BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN species found on that partic- ular host, divided by the number of hosts of the species infested with those parasites). 9 = the specific stage of develop- ment and/or sex of parasite found on that host. Jun Jul = the specific months when these parasites were found on that host. Asterisks preceding the names of the hosts indi- cate that of all of those listed, they are the ones most commonly infested. Except where indicated other- wise, the hosts are mammals. Mite-Host Associations Androlaelaps leviculiis (Mar-AuR) 13 dny 1 d 17 9 Dipodomys ordii 2 (808) 1.5-9: Jun Jul Eutamias minimus 1 (398) 1 .0 - 9: Aug * Onychomys leiicogaster 2 (63) - 4.5 - dny 9: Mar Aug * Peromyscus manicidatus 7 (1866) 2.6 - dny 6 9:Mar June-Aug Androlaelaps sp. (Jul-Sep) 3 dny 1 d 7 9 Dipodomys ordii 3 (808) 1 .0 - 9: Jul-Aug Onychomys leiicogaster 1 (63) 4.0 - dny 9: Aug Perognatlius panms 1 (434) 1.0 -d: Aug Peromyscus manicidatus 2 (1866) 1.5 - 9: Aug Sep Bernia niarita { Aug-Nov) 2 la Eremophila a Ipestris ihird) 1 (84) 1.0: Nov Peromyscus manicidatus 1 (1866) 1.0: Aug Brevisterna sp. (Aug) 2 dny Neotoma cinerea 1(14) 2.0 Chatia ochotona (Aug-Oct) 21 la Neotoma cinerea 1(14) 7.0: Aug Plecotus townseiidii 1 (78) 14.0: Oct Dermaiiyssus gallinae (Feh-iun) 1 d 14 9 Asyndesmus lewis (bird) 1 (2) 1.0-9: May Dendrocopos villosiis (bird) 1(1) 4.0 - 9: Jun * Eremophila alpestris (hnd) 5 (84) 1 .6 - d 9: Feb Apr Tardus migratorius (bird) 1 ( 1 1 ) 1 .0 - 9: Jun Dermanyssiis sp. (Apr-Jun) 2 pny 2 9 1? Eremophila alpestris (bird) 3 (84) 1.0 - pny 9: Apr Piranga ludoviciana (bird) 1 (16) 1 .0 - ?: Jun Dipodomys ordii 1 (808) 1 .0 - 9: Apr Eubrachylaelaps circularis (Mar-Jun) 4 9 Peromyscus maniculatus 3 (18t)6) 1.3: Mar Jun Eubrachylaelaps crowei (Mar-Oct) 202 9 Dipodomys ordii 2 (808) .5: Sep * Onychomys leucogaster 22 (63) 8.8: Mar Jun-AugOct Peromyscus maniculatus 5 (1866) 1.6: Mar Aug Oct Eiibrachvlaelaps debilis (Jan-Dec) 1 pny 12 dny 1 d 998 9 Crotalus viridis (snake) 1 (95) 1 .0 - 9: Jun Centrocercus urophasianus {hud) 1 (18) 1.0-9: Feb Dipodomys ordii 1 (808) 5.0 - 9: Mar Jul Aug Onychomys leucogaster 3 (63) 5.3-9: Mar Perognatlius parx'us 5 (474) 1.2 - 9: May Jun Sep * Peromyscus maniculatus 222 (1866) 4.6 - pny dny d 9: Jan-Sep Nov Dec Eubrachylaelaps sp.{Peb-Uov) 1 pny 10 dny Dipodomys ordii 1 (808) 1 .0 - dny: Apr Peromyscus maniculatus 1 (1866) 9.0 - dny: Feb May Jun Nov ? host - pny; Jul Euschoengastia cordiremus (Jul-Oct) 19 la Dipodomys ordii 1 (808) 7.0: Jul Aug Peromyscus maniculatus 2 ( 1866) 6.0: Oct Euschoengastia criceticola (Oct) 1 la Peromyscus maniculatus 1 (1866) 1.0 Euschoengastia decipiens (Mar-Dec) 488 la Zonotrichia leucophrys (bird) 1 (33) 3.0: Sep * Dipodomys ordii 13' (808) 7.0: Mar-May Jul Aug Oct Eutamias minimus 4 (398) 2.8: Oct * Lepiis californicus 2 ( 1 25) 1 1 .0: Dec Neotoma cinerea 1 (14) 3.0: Sep * Perognatlius pannis 12 (474) 13.6: May Aug- Oct * Peromyscus maniculatus 14 (1866) 5.9: Apr May Jul-Nov Sylvilagus idahoensis 1(13) 56.0: Nov * Sylvilagus niiitallii 3 (28) 14.0: Oct-Dec Thomomys talpoides 1 (8) 15.0: Sep Euschoengastia fasolla (Oct) 8 la Eutamias minimus 2 ( 398) 4.0 Euschoengastia land (Jul) 8 la Peromyscus maniculatus 1 ( 1866) 8.0 Euschoengastia luteodema (Dec) 5 la Lepus californicus I ( 1 25) 5.0 BIOLOGICAL SERIES, VOL. 12 NO. 1 MITES AND LICE Euschoengastia oregonensis (iu\) 1 1 la Salpinctes obsoletus (bird) 1 (17) 1 1 .0 Euschoengastia pomerantzi (Oct) 1 la Eutamias minimus 1 (398) 1 .0 Eusclwengastia radforcli (Apt-Dec) 80 la * Ampliispiza belli (bird) 4 (38) 2.3: Apr Centwcercus uwphasiamis (bird) 1 (18) 4.0: Nov Churdeiles minor (bird) 1(5) 2.0: Aug Eremophila alpesths (bird) 2 (84) 2.5: Apr Nov Junco oregamis (bud) 1 (30) 1 1.0: Oct Lanius ludovicianus (bird) 1 (20) 1.0: Apr Leucosticte tepliwcotis (bird) 1 (25) 3.0: Nov Dipodomys ordii 1 (808) 5.0: Oct * Lepus californicus 5 (125) 4.6: Dec Pewgnathus parvus 1 (474) 1.0: May Peromyscus maniculatus 1 (1866) 6.0: Aug Sylrilagus mittallii 2 (28) 3.0: Oct Dec Euschoengastia sciuricola (May -Oct) 7 la Eutamias minimus 2 (398) 2.0: Oct Marmota flaviventris 1 (6) 3.0: May Eusclwengastia sp. ( Aug-Oct) 19 1a Dipodomys ordii 3 (808) 1 .0: Aug Oct Microtus montanus 1 (25) 8.0: Aug Perognathus pairus 2(474) 2.5: Aug Peromyscus maniculatus 2 (1866) 1.5: Sep Oct Haemogamasus ambulans (Mar-Oct) 8 dny 1 d 80 9 * Dipodomys ordii 1 0 (808) 1 .4 - 9: Jun Jul Eutamias minimus 1 (398) 1 ,0 - dny: Mar * Onychomys leucogaster 6 (63) 5.7 - dny 6 9: Jun-Aug Oct Perognathus pamis 2 (474) 1 .0 - 9: Jun Aug * Peromyscus maniculatus 31 ( 1866) 1.2 -9: Mar Jun-Aug Oct Reithrodontomys megalotis 1 (39) 1.0-9: Aug Thomomys talpoides 1 (8) 1 .0 - dny 9: ? Haemogamasus longi tarsus (Jun) 7 9 * Onychomys leucogaster 3 (63) 1.0 Peromyscus maniculatus 4 ( 1 866) 1 .0 Haemolaelaps casalis (iun-Oct) 19 1? Perognathus pan'us 1 (474) 1.0: Oct Perotnyscus maniculatus 1 (1866) 1.0; Jun Haemolaelaps glasgowi (Jan-Nov) 26 pny 64 dny 57 6 1062 9 Eremophila alpestris (bird) 1 (84) 1.0 - 9: Mar * Dipodomys ordii 11 (808) 2.1 - pny dny d 9: Mar-Aug Oct Nov * Eutamias minimus 24 (398) 4.9 - pny 9: Mar Jun-Oct Microtus montanus 1 (25) .1 - d 9: Jan Jun-Aug Nov * Onychomys leucogaster 36 (63) 5.4 - pny dny d 9: Mar Apr Jun-Oct * Perognathus pannis 33 (474) 2.8 - pny dny d 9: Apr-Oct * Peromyscus maniculatus 281 (1866) 2.1 -pny dny d 9: Jan-Nov Plecotus townsendii I ( 78) 5.0-9: Apr Reithrodontomys megalotis 1 (39) 1.0-9: Aug Spermophilus townsendii 9 (60) .3-9: Apr-Jul Thomomys talpoides 1 (8) 1 .0 - 9 : Jun Haemolaelaps sp. (Mar-Jun) 1 pny 1 d Dipodomys ordii 1 (808) 1.0 -d: Mar Peromyscus maniculatus 1 (1866) 1.0 - pny: Jul Hirstionyssus bisetosus (Sep) 2 9 Neotoma cinerea 1 (14) 2.0 Hirstionyssus hilli (Mar-Aug) 1 7 9 Eutamias minimus 1 (398) 1.0: Jul Onychomys leucogaster 1 (63) 1.0: Mar Jun * Perognathus pan'us 5 (474) 2.6: May Jul Aug Peromyscus maniculatus 2 (1866) 1.0: Jul Aug Hirstionyssus incomptus (Mar-Nov) 91 9 * Dipodomys ordii 37 (808) 21.0: Mar Jun-Oct Eutamias minimus 1 (398) 1.0: Jun Perognathus parvus 2 (474) 2.0: Jun Jul Peromyscus maniculatus 6(1866) 1.2: Jun-Aug Nov Hirstionyssus isabellinus (Nov) 1 9 Microtus montanus 1 (25) 1.0 Hirstionyssus longichelae (Jun-Oct) 6 9 Dipodomys ordii 1 (808) 1.0: Jun Peromyscus nianiculatus 1 (1866) 1.0: Jun * Thomomys talpoides 2 (8) 2.0: Jun Oct Hirstionyssus neotomae (Sep-Oct) 48 9 Eutamias minimus 1 (398) 4.0; Oct Neotoma cinerea 1(14) 44.0; Sep Hirstionyssus thomomys (Mar-Oct) 7 9 Onychomys leucogaster 1 (63) 1.0: Jul Peromyscus maniculatus 1 ( 1866) 1.0: Sep * Thomomys talpoides 3 (8) 1 .7: Mar Jun Oct Hirstionyssus triacanthus (Apr-Oct) 1 85 9 Sceloponts graciosus (lizard) 1 (314) 1.0: Sep Chordeiles minor (bird) 2 (5) 1 .5; Aug Dipodomys ordii 1 1 1 (808) 2.4: Apr-Oct Eutamias minimus 5 (398) 1 .0; Jul Oct Neotoma cinerea 1 (14) 1.0: Sep Perognathus parvus 2 (474) 2.0: Jun Jul Peromyscus maniculatus 5 (1866) .8: Jun-Aug Spermophilus townsendii 2 (60) 1 .0: Jun BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Hirstionyssus utahensis (Mai-Nov) 54 9 Dipodomys ordii 2 (808) 1.5: Jun Aug * Eutamias minimus 9 (398) 2.8: Jul Aug Onychomys leucogaster 1 (63) 1.0: Jun Perognatlius pan'iis 5 (474) 1.0: Jun Jul * Peromyscus maniciilatus 74 (1866) .3: Mar Jun-Sep Nov Spermoptiilus townsendii 1 (78) 1.0: Jul Hirstionyssus sp. (Mar-Oct) 71 dny 36 d 14 9 Dendrocopos villosus (bird) 1 ( 1 ) 1 .0 - d: '? Dipodomvs ordii 18 (808) 2.9 - dny d 9: Jun- Oct Eutamias minimus 5 (389) 1.4 - dny d: Jun Jul Sep Oct Microtus montanus 1 (25) 1 .0 - dny: ? Neotoma cinerea 2 ( 14) 4.0 - dny d 9: ? Onycliomys leucogaster 1 (63) 1.0 - dny 9: Mar Apr Perognathus pamis 6 (474) .7 - dny d 9: May- Aug Peromyscus maniculatus 23 (1866) .8 - dny d 9: Jun-Aug Tliomomys talpoides 3 (8) 4.0 - dny d: ? Hypoaspis lubrica (Aug) 1 9 Perognathus pan'us 1 (474) .0 9: Ischyropoda armatus (May-Oct) 3 dny 25 d 186 9 Crotalus virdis (snake) 1 (95) 1 .0 - 9.- Sep * Dipodomys ordii 33 (808) 1 . 1 - d 9: Jun-Sep Lepus californicus 1 ( 1 25) 1 .0 - dny: Oct Microtus montanus 3 (25) .7-9: Jul Aug * Onychomys leucogaster 23 (63) 3.0 - dny d Jun-Oct * Perognathus pan'us 14 (474) 1 .0 - d 9: May-Sep * Peromyscus maniculatus 61 (1866) 1.3 - dny d 9: May-Oct Sorex merriami 1 (9) 7.0 - 9: Aug Thomomys talpoides 1 (8) 1 .0 - 9: Oct Ischyropoda furmani ( Jan-Aug) 3 d 25 9 * Dipodomys ordii 9 (808) 1.2-9: Jun-Aug Onychomys leucogaster 2 (63) 1 .0 - 9: Aug Perognathus pan'us 5 (474) 2.6 - pny dny d 9: Jun-Aug Peromyscus maniculatus 9 ( 1866) 1 .0 - d 9: Jan Jun-Aug Ischyropoda sp. ( Apr-Sep) 5 pny 55 dny 14 d 23 9 Dipodomys ordii 7 (808) 2.3 - pny dny d 9: Jun-Sep Eutamias minimus 3 (398) 1 .0 - dny 9: Aug Oct Neotoma cinerea 1(14) 2.0 - dny d 9: Jun Aug Onychomys leucogaster 7 (63) 5.6 - dny d 9: Jun Aug Perognathus pamis 5 (474) 2.6 - pny dny d 9: Jun-Aug Peromyscus maniculatus 15 (1866) 1.4 - pny dny d 9: Jan Jun-Aug Sorex merriami 1 (9) 3.0 - dny: Aug Spermophilus townsendii 1 (60) 1.0 -dny: Apr Kleemania sp. (Apr-Sep) 154 9 Chordeiles minor (bird) 1 (5) 1.0: Aug * Dipodomys ordii 8 (808) 6.5: Jun-Sep Eutamias minimus 1 (398) 1.0: Jul Microtus montanus 1 (25) 12.0: Aug Neotoma cinerea 1 (14) 1.0: Sep * Onychomys leucogaster 5 (63) 3.8: Jun-Aug * Perognathus pannis 1 (474) 4.3: Apr Jun-Sep * Peromyscus maniculatus 32 (1866) 1.2: Jun- Sep Sorex merriami 1 (9) 1.0: Aug Leeuwenhoekia americana (Jul) 6 la Dipodomys ordii 1 (808) 6.0 Listrophorus sp. (Mar-Nov) 960 Chordeiles minor (hhd) 1 (5) 1.0: Aug * Dipodomys ordii 53 (808) 18.1 : Mar-May Aug- Nov Peromyscus maniculatus 1 (1866) 2.0: Jun Macronyssus unidens ( Feb-Mar) 1 I a 1 pny 1 dny 1 d + others Plecotus townsendii 35 ( 78) ? Odontacanis liiisdalei (Jul) 6 la Dipodomys ordii 1 (808) 6.0 Odontacanis micheneri {Ju\) 10 la Dipodomys ordii 1 (808) 10.0 I Ornithonyssus bacoti (Oct) 1 9 Dipodomys ordii 1 (808) .0 Ornithonvssus svlvianim (Jul-Sep) 16 9 Amphispiza beUi (hud) 2 (38) 2.0: Jul Oreoscoptes montanus (bird) 1(13) 2.0: Jul * Poocetes gramineus (bird) 3(13) 1.3: Jul Zonotrichia leucophrys (bird) 1 (33) 4.0: Sep Radfordia bachai [Aug) 1 la Dipodomys ordii 1 (808) .0 Trombicula arenicola (Jul-Oct) 325 la Dipodomys ordii 26 (808) 1 2.5 Trombicula bakeri (Jul-Aug) 211a Dipodomys ordii 1 (808) 21.0 Trombicula belkini {}u\-Aug) 167 la Phrynosoma douglassi (lizard) 1 (19) 7.0: Aug * Sceloporus graciosus (lizard) 19(314) 8.4: Jul Aug BIOLOGICAL SERIES. VOL. I 2 NO. 1 MITES AND LICE Tnnnbiciila doremi (iu\-Aug) 98 la * Dipodomys ordii 24 (808) 3.6: Jul Aug Onychomys leucogaster 1 (63) 1.0: Aug Perognathus pan'us 5 (474) 2.0: Aug Twmbicula sp. (Aug-Oct) 12 1a Sceloporus graciosits (lizard) 1 (314) 1.0: Aug Dipodomys ordii 3 (808) 3.0: Aug Oct Onychomys leucogaster 1 (63) 1.0: Aug Peromyscus tiwniculatus 1 (1866) 1.0: Aug Louse-Host Associations Enderleinelliis sp. (prob. sutiiralis) (Apr-Oct) 1 d 3 9 Perognathus parvus 1 (434) 1.0-9: Oct Peromyscus nwniculatus 1 (1866) 1.0 - d: Aug * Spermophihis townsendii 2 (60) 1.0 - 9: Apr Jul Fahrenholzia pinnata (Mar-Nov) 63 d 137 9 Sceloporus graciosus (lizard) 1 (314) 1.0 - 9: Aug * Dipodomys ordii 86 (808) 1.8 - d 9: Mar Apr Jun-Nov Eutamias minimus 4 (398) 1.5 - d 9: Jun Jul Oct * Perognathus pan'us 15 (474) 1.6 - d 9: May- Aug Oct Peromyscus maniculatus 11 (1866) 1.4 - d 9: Apr Jun-Sep Sylvilagus idahoensis 1 (13) 1 .0 - d: Nov Fahrenholzia sp. (prob. pinnata) ( Jun-Oct) 1 d 4 9 47 im 1 ? * Dipodomys ordii 27 (808) 1.0 - 9 im: Jun-Aug Eutamias minimus 1 (398) 1 .0 - im: Jul Onychomys leucogaster 1 (63) 1.0 - im: Jun Perognathus parvus 6 (474) 2.2 - d 9: im: May- Aug Oct Peromyscus maniculatus 2 (1866) 2.0 - im '?: Jun-Jul Oct Geomydoecus sp. (Oct) 4 d 9 9 2 im Thomomys talpoides 1 (8) 15.0 Haemodipsus serow; (Feb-Jul) 21 d 28 9 18 im Lepus californicus 2 ( 125) 12.0 - d 9 im: May Jul Peromyscus maniculatus 1 (1866) 1.0 -9: Mar * Sylvilagus nuttalii 2 ( 28) 21 .0 - d 9 im: Feb Hoplopleura acanthopus (Mar-Aug) 8 d 20 9 Micro tus mon tonus 6 (25) 4.7 Hoplopleura arboricola (Mar-Oct) 136 d 257 9 1 im Sceloporus graciosus (lizard) 1 (314) 2.0 - 9: Sep Ereunetes mauri (bird) 1 (6) 1 .0 - 9: Aug Dipodomys ordii 2 (808) 3.5 - d 9 : Mar Jul * Eutamias minimus 83 (398) 4.0 - d 9 : Mar May- Aug Oct Marmota flaviventris 2(6) 1 .0 - d: Jun Perognathus parvus 1 (474) 5.0 - d 9: Aug Peromyscus nwniculatus 16 (1866) 2.1 - d 9: Mar Jun-Aug Spermophilus townsendii 2 (60) 4.0 - d 9 im: Jun-Jul Hoplopleura erratica (Jul-Oct) 1 d 4 9 * Eutamias minimus 4 (398) 1.0 - d 9: Jul-Aug Oct Perognathus parxnis 1 (474) 1.0-9: Jul Hoplopleura hesperomydis (Jan-Dec) 1 6 d 440 9 Ereunetes mauri (bird) 1 (6) 8.0 -6 9: Mar Jun Aug Dipodomys ordii 1 (808) 1 .3 - d 9 : Jun Aug Eutamias minimus 7 (398) 1.1 - d 9: Mar Jun Aug Microtus montanus 2 (25) 1 .0 - d 9: Jun Aug Onychomys leucogaster 1 (63) 1.0-9: Jul Perognathus parvus 3 (474) 2.0 -d 9: Aug * Peromyscus maniculatus 167 (1866) 3.4 - d 9: Jan-Dec Reithrodontomys megalotis 1 (39) 2.0 - d 9: May Neohaematopimis inornatus (Aug) 2 d 2 9 Neotoma cinerea 2(14) 2.0 Neohaematopinus laeviusculus (Apr-Aug) 68 d 107 9 60 im Eutamias minimus 2 (398) 1 .0 - d: Jul-Aug Marmota flaviventris 2(6) 1.0-9: May Perognathus pan'us 1 (474) 2.0 - d 9: Jul Peromyscus maniculatus 5 (1866) 2.8 - d 9: Mar-Jun Aug * Spermophilus townsendii 26 (60) 8.5 - d 9 im: Apr-Jul Neohaematopinus marmota (Apr-Aug) 25 d 26 9 13 im * Marmota flaviventris 4 (6) 14.0 - d 9 im: May- Jun Peromyscus maniculatus 4 (1866) 2.3 - d 9: Apr Aug Neohaematopinus pacificus (May-Nov) 2 Id 46 9 * Eutamias minimus 27 (398) 2.1 -d9: May-Aug Oct Nov Perognathus parvus 2 (474) 1.5 - d: Jul Peromyscus maniculatus 4 (1866) 1.8 - d 9: Feb Mar Aug Reithrodontomys megalotis 1 (39) 1.0-9: Aug Spermophilus townsendii 1 (60) 2.0-9: Jul Neohaematopinus sp. (Aug) 1 ? Dipodomys ordii 1 (808) 1 .0 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Neotrichodectes intermptofasciatiis ( Apr-Nov) 15 9 23im Taxidea taxiis 2(5) 28.5 19 6 .0 -6: Polyplax aimcularis (Jan-Dec) 331 6 669 9 Sceloponis graciosus (reptile) 2 (314) Sep Ereunetes inaiiri (hnd) 1 (6) 1 .0 - 9: Aug Dipodomys ordii 4 ( 808 ) 1 .0 - d 9 : Jul-Aug Nov Eutamias minimus 7 (398) 1,0 - d 9: Mar Jun- Aug Neotoma cinerea 1 (14) 1.0-9: Jun Perognatlnis pannis 2 (474) 1 .0 - 9: Jun Aug * Peromyscus maniculatus 224 (1866) 4.3 - 6 9: Jan-Dec * Spermophilus townsendii 5 (60) 1.0 - c5 9: Apr Jun-Jul Sylvilagus nuttallii 1 (28) 1.0 - 9: Aug Polyplax spiimlosa (Jul) 1 6 Microtus montamts I (25) .0 Polyplax sp. (Jun-Jul) 2 9 Microtus moiitamis 2 (25) 1 .0 Mallophaga (Jan-Oct) 35 c5 70 9 68 im 6 ? Buteo regalis (bird) 1 (4) 24.0 - d 9 im: Aug Centrocercus urophasianus (bird) 2 (18) 2.5 -9 im: Jul Circus cyaneus (bird) 1(1) 1 .0 - im: Apr Didodomys ordii 3 (808) 1.0 - ?: Apr Sep-Oct Ercmophila atpestris (bird) 1 (84) 4.0 - d 9: Jul Ereunetes mauri (bird) 4 (6) 2.5 - 9 im: Mar Jun-Aug Eutamias minimus 6 (398) 3.2 - 9 im: Mar Jun- Aug Falco span'erius (bird) 1 (6) 1 .0 - d: Jun Junco oreganus (bird) I (30) 1.0 - im: Lepus californicus 1 (125) 9.0 - d 9: Lynx nifus 1 (8) 2.0 - im: Jan Oreoscoptes montamts (bird) I (13) 1.0-9: Apr Apr Jul Perognatlnis pan'us 5 (474) 3.4 - d 9 im; ? : Apr-May Jul-Aug Peromyscus maniculatus 9 (1866) 1.9 -d 9 im: Mar-May Jul-Aug Pica pica (bird) 1 (8) 1.0 - im: Feb Plecotus townsendii 1 (78) 4.0 - im: Apr Reithrodontomys megalotis 1 (39) 15.0 - d 9 im: Apr Sceloponis graciosus (lizard) 1 (314) 3.0 - 9 im: Jun Spermophilus townsendii 5 (60) 7.6 - d 9 im: Apr-Jun Spinus pimts (bird) 1 (23) 1 .0 - ?: Jun Sternella neglecta (bird) 1(7) 1.0 - im: Jul Zenaidura macroura (bird) 1 (23) 2.0 - d im: Jul HOST-PARASITE RELATIONSHIPS ( * = the mite and/or louse which occurred most commonly on that host; H = new host record based on replications— other associations listed may re- present new records, but are not so indicated because of infrequent occurrence considered accidental infes- tations or contaminations). Reptiles Crotalus viridis Eubrachylaelaps dehilis Ischyropoda armatus Phrynosoma douglassi Trombicula belkini Sceloponis graciosus Hirstionyssus triacanthus * Trombicula belkini Fahrenholzia piniwta Hoplopleura arboricola Polyplax auricularis Mallophaga Birds Amphispiza belli * Euschoengastia radfordi Ornitlionyssus sylviarum Asyndesmus lewis Dermanyssus gallinae Buteo regalis Mallophaga Centrocercus urophasianus Eubrachylaelaps debilis * Euschoengastia radfordi Mallophaga Circus cyaneus Mallophaga Chordeiles minor Euschoengastia radfordi Hirstionyssus triacanthus Kleemania sp. Listrophonis sp. Dendrocopos villosus * Dermanyssus gallinae Hirstionyssus sp. Eremopliila alpestris Bernia marita * Dermanyssus gallinae Euschoengastia radfordi Haemolaelaps glasgowi Mallophaga Ereunetes mauri Hoplopleura arboricola H. hesperomydis Polyplax auricularis Mallophaga BIOLOGICAL SERIES, VOL. 1 2 NO. 1 MITES AND LICE Falco sparverius Mallophaga Junco oreganus Euschoengastia radfordi Mallophaga Laniiis ludovkianus Euschoengastia radfordi Leucosticte teplirocotis Euschoengastia radfordi Oreoscoptes muntamis Ornithonyssus sylviarum Mallophaga Pica pica Mallophaga Piranga ludoviciana Dermanyssits sp. Pooecetes gramineus Ornithonyssus sylviarum Salpinctes obsoletus Euschoengastia oregonensis Spinus pinus Mallophaga Sternella ncglecta Mallophaga Turdus migratorius Dcrnmnyssus gallinae Zenaidura macroura Mallophaga Zonotrichia leucoplirys Euschoengastia decipiens * Ornithonyssus sylviarum Mammals Dipodomys ordii Androlaelaps leviculus Derrnanyssus sp. Eubrachylaelaps crowei E. debilis Euschoengastia cordiretnus * E. decipiens E. radfordi Hacmogamasus ambulans * Haemolaelaps glasgowi * Hirstionyssus incomptus H. longichelae H. triacanthus H. utahensis * Iscliyropoda armatus I. furmani * Kleemania sp. Leeuwenhoekia americana * Listrophorus sp. Odontacarus linsdalei O. micheneri Ornithonyssus bacoti Radfordia bacliai * Trombicula arenicola T. bakeri * T. doremi * Fahrenholzia pinnata Fahrenholzia sp. Hoplopleura arboricola H. hesperomydis ■ H Neohaemotopinus sp. Polyplax auricularis Mallophaga Eutamias minimus Androlaelaps leviculus Euschoengastia decipiens - H E. fasolla E. pomerantzi E. schuricola Hacmogamasus ambulans * Haemolaelaps glasgowi Hirstionyssus hilli H. incomptus H. neotomae H. triacanthus - H H. utahensis Ischyropoda sp. - H Kleemania sp. Fahrenholzia pinnata Fahrenholzia sp. * Hoplopleura arboricola H. erratic a H. hesperomydis - H Neohaematopinus laeviusculus * N. pacificus Polyplax auricularis - H Mallophaga Lepus californicus Euschoengastia decipiens E. luteodema * E. radfordi Ischyropoda armatus Haemodipsus setoni Mallophaga Lynx nifus Mallophaga Marmota flaviventris Euschoengastia sciuricola Hoplopleura arboricola Neohaematopinus laeviusculus * N. marmotae Microtus montanus Euschoengastia sp. * Haemolaelaps glasgowi Hirstionyssus isabellinus Ischyropoda armatus ■ H Kleemania sp. * Hoplopleura acanthopus - H H. hesperomydis Polyplax spinulosa Polyplax sp. Neotoma cinerea 8 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Brevistema sp. Chatia ochotona Eiischoeiigastia decipiens Hirstionyssits bisetosus * H. neotomae H. triacanthus Ischyropoda sp. Kleemania sp. * Neohaematopimis inurnatus - H Polyplax aiirkularis Onychomys leucogaster Androlaelaps leviculus * Eubradniaelaps crowd E. debilis - H Haemogamasus ambulans H. I o 11 gi tarsus - H * Haemolaelaps glasguwi Hirstionyssits liilli H. thomomys H. iitahensis * Ischyropoda annatiis I. furmani Kleemania sp. Trombicula doremi Fahreiiholzia sp. Hoploplenra liesperoniydis Perognathus paints Androlaelaps sp. Dennaiiyssus galliiiae Eitbrachylaelaps debilis - H * Eiischoeiigastia decipiens E. radfordi Haemogamasus ambulans Haemolaelaps casalis * H. glasgowi * Hirstioiiyssus hilli H. incomptus H. triacanthus H. utahensis - H Hypoaspis lubrica * Ischyropoda armatus I. furmani - H * Kleemania sp. Trombicula doremi - H Enderleinellus sp. * Fahreiiholzia piiinata Fahreiiholzia sp. Hoploplenra arboricola H. erratica H. hesperomydis Neohaeinatopinus laeviusculus N. pacificus Polyplax auricularis Mallophaga Peromyscus maniculatus Androlaelaps leviculus Beniia niarita Eubrachylaelaps circularis E. crowei * E. debilis Euschoengastia cordiremus E. criceticola * E. decipens E. laiiei E. radfordi * Haemogamasus ambulans H. longitarsus Haemolaelaps casalis * H. glasgowi Hirstionyssus hilli H. incomptus H. longichelae H. thomomys H. triacanthus * H. utahensis * Ischyropoda armatus I. furmani * Kleemania sp. Listrophorus sp. Trombicula sp. Enderleinellus sp. Fahrenholzia pinnata - H Fahreiiholzia sp. Haeniodipsus setoni Hoploplenra arboricola - H * H. hesperomydis Neohaeinatopinus laeviusculus N. marmotae N. pacificus * Polyplax auricularis Mallophaga Plecotus townsendii Chatia ochotona Haemolaelaps glasgowi * Macronyssus unidens Mallophaga R eithrod( nitomys megalo tis Haemogamasus ambulans Haemolaelaps glasgowi Hoploplenra hesperomydis Neohaeinatopinus pacificus Mallophaga Sorex merriami Ischyropoda armatus Kleemania sp. Spcrmophilus to wnsendii "•^ Haemolaelaps glasgowi Hirstionyssus triacanthus H. utahensis Ischyropoda sp. Enderleinellus sp. Hoploplenra arboricola * Neohaeinatopinus laeviusculus N. pacificus Polyplax auricularis Mallophaga I BIOLOGICAL SERIES, VOL. 12 NO. 1 MITES AND LICE Sylvilagus idahoensis Eiischoengastia decipiem Fahrenholzia pinnata Sylvilagiis nuttallii * Eiischoengastia decipiens E. radfordi * Haemodipsus setoni Polyplax aiiricidaris Taxidea taxiis Neotrichodectes intemiptofasciatus Thomomys talpoides Eiischoengastia decipiens Haemogamasus ambulans Haemolaelaps glasgowi Hirstionyssiis longichelae * H. thomomys Ischyropoda armatus Geomydoecus sp. ECOLOGICAL CONSIDERATIONS Degree of Host Infestation The degree of infestation of mammals differed rel- ative to their geographic distribution (Table 1 ). A greater percentage of Peromyscus maniculatiis was infested with mites in Area 12 than in other areas, with the lowest percentages in Areas 6, 10 and 11. Pcrognatlnts pannis in Areas 1 and 5 were the most heavily infested, whereas those in Area 7 were the least. In Area 5 Eiitamias minimus were heavily infested, whereas in Area 3 none were infested. The greatest infestation rate for Dipodomys ordii was in Area 8 and the lowest in Area 4. Except in Area 5, no two species of mammals in the same area had a higli rate of infestation with mites. The same was true for the lowest rates of infestation. A greater percentage of Dipodomys ordii was in- fested with lice in Areas 6 and 8, and fewer animals in Areas 5 and 10 were infested than in other areas. The greatest percentage of infested Eutamias minimus was in Areas 2 and 10, and fewest in Area 7. No signifi- cant differences were noted for Perognathiis parvus except in Areas 7, 9 and 10, where no infested ani- mals were found even though 108 were examined. A greater percentage of Peromyscus maniculatus was in- fested in Area 4, and fewer infested animals were found in Area 1 1 than in other areas. On the basis of frequency and degree of infestation, the areas where louse infestation was optimum for Dipodomys ordii Table 1 . Percentage of hosts' infested with mites and lice in each of 1 2 major study areas. Host %^ of hosts infested in each study area-* 10 11 12 MITES Dipodomys ordii 71 50 45 58 87 97 77 71 89 53 Eutamias minimus 10 6 0 24 6 4 Perognatlius pan'us 35 16 21 38 15 24 20 Peromyscus maniculatus 42 32 60 45 14 50 26 45 18 18 79 LICE Dipodomys ordii 17 17 12 6 21 23 8 6 16 14 Eutamias minimus 20 32 24 24 6 40 Perognatlius parvus 8 8 3 8 0 3 0 Peromyscus maniculatus 27 29 23 30 12 18 22 11 12 8 21 'Only those hosts that were widely distributed geographically are included. ^To nearest whole percent. ^Data not included when less than 10 hosts from the area were examined. 10 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN are 6, 8 and 11, for Eiitamias minimus 2 and 3, and for Peromyscus maniculatiis 2 and 5. Considering lice of all species, hosts in Area 2, 5 and 9 were more heavily infested than those in other areas. In each of tiie 1 2 study areas, a greater percentage of the hosts belonging to Dipodomys ordii, Perog- nathus parvus, and Peromyscus maniculatus were in- fested with mites than with lice. The reverse situation occurred with Eutamias minimus. Four exceptions to these conditions wherein about equal percentages of hosts were infested with mites and lice were Eutamias minimus in Areas 5 and 7, and Peromyscus manicula- tus in Areas 2 and 6. Host Abundance and Species Variety The number of species 'parasites which occurred on a particular kind of host generally was in direct proportion to the number of hosts examined (Table 2). Table 2. Number of mammals examined and number of species of mites and lice found on each kind. Host No. species Species No. examined Mites Lice Peromyscus maniculatus 1 866 Dipodomys ordii 808 Perognathus parvus 474 Eutamias minimus 398 Lepus californicus 125 Plecotus townsendii 78 Onychomys leucogaster 63 Spermophilus townsendii 60 Reithrodontomys megalotis 39 Sylvilagus nuttalli 28 Microtus montanus 25 Neotoma cinerea 14 Sylvilagus idahoensis 1 3 Sorex merriami 9 Thomomys talpoides 8 Marmota flaviventris 6 25 25 17 14 4 3 13 4 2 2 5 8 1 2 6 1 Mammals of three species did not fit the expected ratio of direct proportion with reference to mites- Peromyscus tnaniculatus and Sylvilagus idahoensis had fewer species of mites than would be expected, and Onychomys leucogaster more species than ex- pected. I assume that the numbers oi' Peromyscus ex- amined approached the upper end of the "numbers examined— species present" ratio, whereas the unex- pected ratios for Sylvilagus and Onyclumiys may be related to their behavior patterns and/or habitat. Lice are more host specific than mites, fleas or ticks. Consequently the sucking lice in this study were more restricted in host distribution than fleas or mites, but followed the similar trend of number of species found in direct proportion to number of hosts examined. Exceptions were Plecotus townsendii on which no lice were found, and Spermophilus towns- endii. Microtus montanus and Marmota flaviven- tris, which possessed more species of lice than expected. Degree of Infestation by Sex of Host Some significant differences in the rate of infesta- tion on mammals of different sexes were noted for parasites of some species (Table 3). Table 3. Comparative rates of infestation by mites and lice on the different sexes of mammals of some species. Parasite-host index Parasite and host 6 hosts 9 hosts Mites Eubracliylaelaps crowei Onychomys leucogaster 5.4 5.0 Eu brachylaelaps debilis Peromyscus maniculatus 2.2 3.3 Eiischoengastia decipiens Dipodomys ordii Perognathus parvus Pennnyscus maniculatus Haemogamasus ambulans Dipodomys ordii Onychomys leucogaster Peromyscus maniculatus Haemolaclaps glasgowi Dipodomys ordii Eutamias minimus Microtus tncmtanus Onychomys leucogaster Perognathus parvus Peromyscus maniculatus Spermophilus townsendii Hirstionyssus incomptus Dipodomys ordii Hirstionyssus triacan thus Dipodomys ordii Hirs tioi lyssus Utah ensis Eutamias minimus Peromyscus tnaniculatus Ischyropoda armatus Dipodomys ordii Onychomys leucogaster Perognathus pan'us Peromyscus maniculatus Ischyropoda furmani Dipodomys ordii 1.3 1 .0 10.4 3.0 19.9 4.0 6.0 5.2 1.3 1.2 3.5 2.4 1.2 1.3 1.6 1.6 1.8 7.3 1.6 1.1 6.0 2.4 1.8 2.9 1.8 1.6 2.8 2.5 1.4 3.2 2.6 2.5 4.7 1.5 2.0 5.3 2.0 1.6 2.6 2.5 1.0 1.1 1.4 1.2 BIOLOGICAL SERIES, VOL. 1 2 NO. 1 MITES AND LICE Table 3. (Continued) 11 Seasonal Occurrence Parasite-host index Parasite and host 6 hosts 9 hosts Peromyscus maniculatus 1.0 1.0 Kleemania sp. Dipodomys ordii 2.1 2.5 Peromyscus maniculatus 1.6 1.0 Listrophonis sp. Dipodomys ordii 20.0 17.6 Trombicula arenicola Dipodomys ordii 9.0 1.2 Trombicula belkini Sceloporus graciosus 6.7 8.3 Trombicula doremi Dipodomys ordii 3.0 4.0 Lice Fahrenhohia pimwta Dipodomys ordii 2.0 1.6 Hoplopkura arboricola Eutamias minimus 4.7 4.0 Hoplopkura hesperomydis Peromyscus maniculatus 4.1 4.1 Neohaematopinus laeviuscuhis Spermophilus townsendii 12.6 5.1 Neohaematopinus pacificus Eutamias minimus 2.4 1.4 Polyplax auricularis Peromyscus nmniculatus 3.6 2.4 Complete seasonal occurrences of the mites and lice are given in the listings in the sections "Mite-Host Associations" and "Louse-Host Associations." Their occurrence on commonly collected hosts is summar- ized in Tables 4 and 5, I assume that a direct correlation exists between the number and kinds of hosts examined and the number of kinds of parasites found. This was true ex- cept for October when proportionately more species of mites and lice were found than would be expected from the number of hosts examined. The number of species of parasites taken during July and August were equal, although more hosts were examined in August than in July. The number of kinds of parasites in relationship to the number of kinds of hosts ex- amined was in direct correlation for other months ex- cept for April, July and November when fewer kinds of mites, and in June when fewer lice were found than expected from the number of kinds of hosts ex- amined. In February more kinds of lice were found than expected. The absence of Mallophaga on so many birds was unusual, particularly on Amphispiza belli.. Lanius ludovicianus. Leucosticte tephrocotis, and Zono- trichia leucophrys. For the sucking lice an unusual situation was the apparent absence of these parasites during specific months. Lice were found on Peromyscus maniculatus every month, yet were absent on other of their com- mon hosts at times when one would expect them to be present. Significant examples of absence are in May for Dipodomys ordii, August and December for Lepus californicus. and September for Perognathus parvus. Males of Dipodomys ordii dnd Perognathus parvus were more heavily infested with mites of Euschoen- gastia decipiens than were females. Relative to mites of Haemolaelaps glasgowi, females of Eutamias mini- mus were more heavily infested, whereas the males of Onychomys leucogaster were more heavily infested. For Hirstionyssus utahensis. male Eutamias minimus were more heavily infested, whereas female Pero?nys- cus maniculatus were more heavily infested. Males of Dipodomys ordii were more heavily infested with mites of Trombicula arenicola than were females. For the most part, a greater percentage of the male hosts were more heavily infested with lice than were the females. This was most evident with Polyplax aur- icularis on Peromyscus maniculatus. In every case but one (Hoplopleura hesperomydis on Peromyscus nmni- culatus) the louse-host index was higher for males than for females, although the difference was not sig- nificant except for Neohaematopinus laeviuscuhis on Spermophilus townsendii. Reproduction In Mites An egg or larva was observed within the idiosoma of mites of seven species. Females oi Eubrachylaelaps crowei were gravid with eggs during June, July, August and October, and with larvae from June to October, inclusive. Females of E. debilis contained eggs from February througli December (except in September and October), and larvae during the same period except October. Mites of Haemolaelaps glas- gowi were gravid with eggs from March through October, and with larvae for the same period except in May and September. Mites of Hirstionyssus hilli contained eggs in August, those of H. incomptus in June and July, those of//, neotomae in September, and those of//, thomomys in October. The cosmopolitan species Haemolaepaps glasgowi, which occurs on such a variety of hosts, was common on mammals of five species at the NRTS, but those taken from Dipodomys ordii contained the greatest 12 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Table 4. Seasonal infestation of some common hosts^ with mites^ and lice.'^ jjQjt No. vertebrates examined and parasites- present Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Reptiles Crotalus viridis 47 Sceloponis graciosus 1 14 Birds Amphispiza belli Eremophila alpestris 6 3 M 4 M 15 29 M Junco oreganus 1 12 L Laniiis ludovicianus Leucosticte tephrocotis Spinus pinus 5 3 5 1 19 33 2 1 6 M M 02 53 65 77 M M M 5 2 8 7 7 L 3 7 2 3 M 16 2 1 17 L Zenaidura inacroura 2 14 5 2 L Zonotrichia leucophrys 6 26 Mammals Dipodomys ordii Eiitamias minimus Lepus californicus Microtus montamis Onychomys leucogaster Perognathus parvus Peromyscus maniculatus Reithrodontomys megalotis Spermophilus townsendii Sylvilagus nuttalii 50 20 26 183 175 245 60 64 13 LM LM M LM LM LM LM LM LM 51 4 7 53 137 103 7 25 6 LM L LM LM LM M LM L 6 8 13 LM 3 L 10 13 L 27 11 7 LM 7 20 M 8 1 L 1 L 5 L 7 L 2 1 7 1 15 13 14 5 6 1 1 M M LM LM M M LM 14 66 117 122 130 19 6 1 LM LM LM LM LM M LM 5 43 114 40 13 439 514 475 69 82 41 40 LM LM LM LM LM LM LM LM LM LM LM LM 6 1 L 3 L 1 9 17 L 4 5 5 LM 8 LM 33 LM 8 LM 7 1 1 4 4 4 2 2 1 3 L ^Only those are listed wherein a total of 20 or more animals were examined of those species which are common hosts for mites or lice. ^AU species. -^L = lice, M = mites. Table 5. Numbers of species of mites and lice collected each month relative to numbers and Icinds of mammals. Item Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec No. hosts examined 19 50 243 98 121 856 1001 1022 173 198 71 64 Kinds hosts examined 3 2 9 8 8 10 10 9 7 9 8 4 Kinds parasites found: Mites 3 4 13 8 10 19 25 25 16 21 8 4 Lice 2 4 8 7 8 8 11 11 4 8 5 2 1 BIOLOGICAL SERIES, VOL. 12 NO. 1 MITES AND LICE 13 percentage of gravid females (13%) when compared with those on Peromyscus mamculatiis {17o), Onv- chomys leiicogaster (5%), Perognathiis parvus (4%), and Eutamias minimus (3%). Consortism Where sufficient numbers of parasites were found to make comparisons, different degrees of species consortism were noted (Table 6). Mites of Euschoengastia decipens. although found on a variety of hosts, were seldom in association with mites of other genera. All of the common species ex- cept Trombicula arenicola occurred as the only species of mite on their hosts in more than half the collections. Euschoengastia radfordi. E. decipiens and Hirstionyssus incomptus frequently were associated with other species of the same genus. Consortism between lice of different species was not as common as with other ectoparasites. Lice of the species Polyplax auricularis occurred as the only Table 6. Frequency of intrageneric and species asso- ciations for some commonly collected mites and lice. %^ of times found: Parasite As only species- With species of on host same genus Mites Trombicula belkini 100 0 Euschoengastia radfordi 94 17 Hirstionvssus incomptus 84 11 Kleemania sp. 83 0 Eubrachylaelaps debilis 80 1 Haemogamasus ambulans 79 0 Euschoengastia decipiens 79 11 Listrophorus sp. 77 0 Hirstionyssus utahensis 77 5 Trombicula doremi 75 6 Ischyropoda armatus 67 2 I. furmani 67 8 Hirstionyssus triacanthus 66 4 Hacmolaelaps glasgowi 55 1 Trombicula arenicola 39 9 Lice Polyplax auricularis 73 0 Fahrenholzia pinnata 67 0 Hoploplcura arboricola 64 9 Hoplopleura hesperomydis 60 18 Neohaematopinus laeviusculus 53 6 Neohaematopinus pacificus 34 6 'Nearest whole percent. ■^Of mites or of lice, respectively. ones on their host a greater percentage of the time than did other lice. All of the commonly collected species except Neohaematopinus pacificus occurred as the only lice on their hosts in more than half of the collections. A significant correlation occurred with A/; pacificus which was associated with Hoplopleura ar- boricola in 43 percent of its collections. A similar cor- relation was noted between Hoplopleura hesperomy- dis and Polyplax auricularis. Significant intrageneric associations were noted for Hoplopleura and Neo- haematopinus. Geographic Distribution The distribution of parasites of most species was directly correlated with the distribution of the host on which the parasites were most commonly found. Of the mites, Eubrachylaelaps debilis, Hirstionyssus incomptus. Listrophorus sp. and Trombicula areni- cola were more widely distributed than expected, and Euschoengastia radfordi, Bernia marita and Ornitho- nyssus sylviarum were more geographically restricted than were the hosts on which they were found. Lice of the species Neohaematopinus pacificus, and especially those of Hoplopleura erratica. were more geographically restricted than the hosts on which they were most commonly found. The numbers of species of mites and lice found in each study area are shown in Table 7. Although fewer species than expected were found in every area, the greatest deviations from the average numbers of species present were Areas 5 and 7 for the mites and Areas 4 and 8 for the lice. Radiation Influence Animals differed in their degree of infestation with ectoparasites in an irradiated area when compared with an ecologically similar nonirradiated one (Table 8). Animals of Eutamias minimus and Peromyscus maniculatus were less frequently infested with mites in the irradiated area than in the nonirradiated one. The mite-host index was about equal for Eutamias minimus in the two areas, but was higher for Pero- myscus maniculatus in the irradiated one. Fewer species of lice were found, a smaller per- centage of mammals were infested, and a lower louse- host index for Eutamias minimus occurred in the ir- radiated area. Although the louse-host index was lower for Peromyscus maniculatus, a higher per- centage of the mammals were infested and more species of lice were found in the irradiated area. Unusual Host Records Routine processing of many animals for ectopara- sites creates a potential for error and contamination 14 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN even though the greatest care is exercised. Conse- quently, many of the host records in the list of host- parasite relationships must be considered as tentative, especially when they represent only one or two col- lections. On the other hand, some of the records re- present sufficient replications to be valid, and con- sequently must be considered as new. These are Euschoengastia decipiens, Hirstionvssus triacanthus and Ischywpoda sp. on Eutamias minimus: Ischy- wpoda armatus on Microtus montamis: Eubrachylae- laps debilis and Haemogamasiis longitarsus on Ony- chomvs leucogaster; and E. debilis. Hirstionyssus iitaliensis, Ischywpoda furmani and Trombicula dorciiii on Perogimtlnts pamis. Table 7. Numbers of species of mites and lice found in the major study areas in proportion to the kinds and numbers of hosts examined. Study Area No. hosts examined^ No. species mites No. species lice Individuals Species Expected- Found Expected- Found 1 169 6 2 155 4 3 311 7 4 40 3 5 117 4 6 51 3 7 57 4 8 148 5 9 93 3 10 39 4 11 39 4 12 49 3 45 18 12 6 29 13 10 5 25 12 12 7 25 9 9 3 41 12 10 5 41 13 10 6 25 14 10 5 41 14 10 7 41 13 9 5 45 17 10 5 41 13 10 4 41 15 10 5 ^Only those are included that are known to be common hosts of mites or lice. ■^Based on number and kinds of hosts examined in relationship to parasites found on these hosts in at least one other study area. Table 8. Differences in degree of infestation by mites and lice on mammals of two species in irradiated and nonirradiated areas. Area No. hosts examined % hosts infested with; No. species present Mites Lice Mites Lice Parasite-host index Mites Lice Eutamias minimus 38 (control) 13 (irradiated) 18 20 67 39 4 30 30 4 7 4 1.2 1.5 13.1 1.7 Peromysciis mauiculatus 37 (control) 13 (irradiated) 22 80 95 14 5 50 26 9 2 4 .6 26.0 11.0 4.3 BIOLOGICAL SERIES. VOL. 12 NO. 1 MITES AND LICE 15 New records for lice, which likely are not contami- nations, are Falircniuilzia piiinata on Peromvscus maniciilatus. Hoplopleiira acanthopus on Micwtiis montanus, H. arboricola on P. maniculatus, H. hesper- omydis on Dipodomys ordii and Eiitamias minimus. and Polyplax auricularis on E. minimus. Taxonomic Considerations of the Lice I have taken the liberty to include some applicable comments made by Dr. William T. Jellison relative to his identifications of the lice. Enderleinellus sp.— These probably were of groimd squirrel origin, and likely are E. suturalis. Fahrenholzia pinnata. -This is a characteristic par- asite of Dipodomys. The specimens from Perognathus were only tentatively relegated to this species by Jellison. On the Idaho specimens the upper left-hand pleural plate is consistently longer than on typical F. pinnata. Haemodipsus setoni.— Lice relegated to this species were taken from Lepus and Sylvilagus. A different species likely occurs on each of these hosts, but so far no distinction has been recognized. Hoplopleura erraf/ca. —Western chipmunks have two louse parasites—//, erratica arboricola and Neo- haematopimis pacificus. The Idaho specimens are closer to the subspecies H. e. erratica, typical of Tamias, than to the western// e. arboricola. Polyplax auricularis.— This typically infests Pero- myscus and mice of several other genera, and Jellison seriously questions the records from Dipodomys, Per- ognathus and Spermophilus. DISCUSSION Community Relationships In the 12 major study areas where collections were made during all seasons, the degrees of infestation of each species of hosts were not consistent between dif- ferent areas. However, in Areas 7 and 10 the degree of infestation was lower for more species than for the other areas. The same predominant plants were pres- ent in Areas 7 and 1 0 and in some of the other areas, but total composition and edaphic differences likely exist which affect parasite infestation of the host as well as its nest. Species Variety For those ectoparasites that are not considered host specific, the number of species of ectoparasites found on a particular species of host was proportion- ate to the number of hosts examined. The fewer kinds of mites than expected to be found on Peromv- scus maniculatus is unusual in consideration of the abundance, distribution and habits of these rodents. The greater number of species of mites than expected on Onychomys leucogaster is not unusual in light of its carnivorous habits. The unusual number of species of lice on Spermophilus. Microtus and Marmota is un- expected because of the apparent host specificity of these ectoparasites and the habitat and behavior of their hosts. Sexual Differences Where degree of infestation according to sex of host differed for a given species, the males were more often and more heavily infested than the females, al- though this varied somewhat with the species of para- site. This may be related to the reproductive, nesting and food-getting activities of the different hosts. The greater degree of infestation of males is contrary to an assumption that females are potentially apt to be more heavily infested because they spend more time associated with the nest because of their reproductive activities. The nest is assumed to be the principle res- ervoir of many ectoparasites of the nest-building ro- dents. On the other hand, the wandering activities of the males may provide for greater potential contact with ectoparasites seeking a host. Furthermore, the nest itself may contain plant materials and dusts which act as pesticides against the ectoparasites. Seasonal Occurrence The summer months (July and August) are ex- pected to represent the optimum period for the great- est number of species of ectoparasites on the hosts in- asmuch as this is the time when populations of hosts are attaining their peak, and environmental condi- tions should be optimum for ectoparasite reproduc- tion. The decline in September and subsequent in- crease in October likely is related to the maturation of immatures produced by the mid-summer adults. The expected decline in winter months occurred for the mites, but populations of lice in February were higlier than expected. This latter situation may be due to the optimum environmental conditions of the nest as a result of animal hibernation, or decreased amounts of activity outside of the nest. Consortism Whether the degree of consortism is directly re- lated to the species of ectoparasite involved or to eda- phic or other environmental factors is not known. For the lice, considered as being more host specific than the mites, individual species seldom occurred with other lice, especially with those of the same genus. Although mites of different species frequently BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN were associated together, those of two species repre- sented the extremes of consortism. Chigger mites of Euschoengastia decipiens seldom were found in asso- ciation with other mites, whereas the mesostigmatid Eubrachylaelaps crowei was almost always found in association with other species. The occurrence of mites of two species of the same genus on the same host was not considered common in proportion to the number of times each species was found, but was much more common in the mites than with the lice. Geographic Distribution One may assume that the distribution of an ecto- parasite should be in direct relationship to the distri- bution of its common hosts, especially with the lice where host specificity is more evident than with the mites. Such was the case with most of those mites and lice studied. Based on host relationships, how- ever, mites of three species were more widely distrib- uted than expected, whereas those of three other species were more restricted than expected. Un- doubtedly environmental factors other than the host are influential on these mites. No correlation with a predominant type of vegetation was evident, and higliest and lowest populations were found in two communities which contained the same species of predominant plants. On the other hand, other species which occurred in greatest numbers were associated with plant associations wherein greater cover and or- ganic debris were present. Radiation Effects The effects of radiation, per se. on the rate of ectoparasite infestation are not known. However, in j one disturbed area fewer ectoparasites occurred than > in an ecologically similar undisturbed area. This situa- i tion occurred for lice and mites on Eutamias mini- 1 imts. but was different for ectoparasites on Peromy- j sens maiiiculatus. Althougli fewer mice were infested j with mites in the irradiated area than in the undis- j turbed area, the mite-host index was higher. Exactly j the reverse situation occurred with lice on P. mauicu- \ latus. I believe that the differences in rates of infes- tation are not due to the effects of radionuclides, but rather to the physical disturbance of the habitat, i.e., destruction of plants and soil manipulation. SUMMARY Mites of 47 species and lice of 16 species were taken from reptiles of 3 species, birds of 23 species, and mammals of 18 species at the National Reactor Testing Station between June, 1966, and September, 1967. Mammals differed in degree of infestation rela- tive to their geographic distribution at the station. The number of species of ectoparasites which oc- curred on a particular kind of host generally was in direct proportion to the number of hosts examined, but exceptions occurred in some instances. In some cases the rate of infestation differed relative to the sex of the host. The greatest numbers of species of mites and lice were found during July and August, fewest species of mites from December through Feb- ruary, and fewest kinds of lice during December and January. Host preferences for mites of some species were evident during their reproductive and nonrepro- ductive periods. Consortism among the ectoparasites differed in degree by species. Except for mites of seven species and lice of two species, the geographic distribution of the ectoparasites was in direct propor- tion to the distribution of the host on which they were commonly found. Mammals of two species dif- fered in their frequency of infestation with ectopara- sites in an irradiated area when compared to an eco- logically similar nonirradiated control area. Ten new host records for mites and six for lice were estab- lished. 17 REFERENCES Allred, D. M.. and D E. Beck. 1966. Mites of Utah mammals. BYUSci. Bull., Biol. Ser., 8(1):1-123. Allred, D. M. 1968a. Ticks of the National Reactor Testing Station. BYUSci. Bull., Biol. Ser., 10(1): 1-29. . 1968b. Fleas of the National Reactor Testing Station. Great Basin Natl., 28(2):73-87. . 1969. Spiders of the National Reactor Testing Station. Great Basin Natl., 29(2): 105-108. Atwood, N. D. 1970. Flora of the National Reactor Testing Station. BYU Sci. Bull., Biol. Ser., 1 1(4) (in press) Ferris, G. F. 1919-1935. Contributions Toward a Monograph of the Sucking Lice. 2(1-8), Stanford Univ. Press; Stanford, Calif. . 1951. The Sucking Lice. Pacific Coast Entomol. Soc, San Francisco. Ignoffo, C. M. 1956. Notes on louse-host associations of the Great Salt Lake Desert with keys to the hce. Great Ba.sin Natl., 16(l-4):9-17. J INFORMATION FOR CONTRIBUTORS Contributions to the Science Bulletin should be pri- marily monographic in nature. For the most part only manuscripts of approximately forty or more typewritten pages will be accepted. Papers will be published approximately in the order that they are received, pending availability of funds. Authors must arrange for financing their publications, except that the University Press and the library of Brigham Young University will share the publication cost proportionately to their needs for copies of the article printed. In the preparation of manuscripts, authors are re- quested to follow the Style Manual for Biological Jour- nals, American Institute of Biological Sciences, 2000 P Street, N.W., Washington, D.C., 20006. Manuscripts must be typed on one side of the paper only, double- spaced with ample margins. Footnotes should be avoided . To facilitate review by referees, send to the editor the original manuscript and one carbon copy, together with the illustrations. Copies of the original illustrations may DC submitted, but should be of quality equal to the orig- nals. Illustrations should be referred to as figures except for materials requiring inserts of special paper, which ■nay be called plates. Illustrations should be so designed is to fit when reduced into a one-column or full-page A'idth. Special care must be taken to allow for proper eduction in lettering (i.e., a 50% reduction of the figure Tieans also a 50% reduction in the lettering.) Photo- graphs should be of a glossy finish, unblurred, and show- ng sharp contrast. Line drawings should be made with ilack ink on heavy white drawing paper, blue tracing ;loth, or blue-ruled coordinate paper. Use the same ibbreviations on line drawings as in text. Line drawings Tiust be equivalent to a professional draftsman's work. Original drawings are preferable to photographs, even if they are large. Illustrations (line drawings or photo- graphs) should be numbered consecutively throughout the paper, and the approximate place of insertion should be indicated in the margins of the manuscript pages. Captions for illustrations should be assembled on a sep- arate sheet, and each plate of figures must have its corre- sponding figure number pencilled lightly on the back. Illustrations and cuts will be destroyed unless their re- turn is requested when proof is returned to the editor. A table title should be a short, concise statement of what the table purports to show, and should not include information necessary to the interpretation of the table. Every column in the table should carry a head identify- ing the data in that column; the measure in which the data are given should be indicated at the head of each column. Tables should not be used when the same infor- mation can be given in a few lines of text, and should not duplicate information in text, graphs, or charts. Symbols (asterisk, dagger, etc.) should be used to indi- cate footnotes to tables, with footnotes on the same page as the table. An abstract of less than four percent of the length of the paper should be prepared. This summary should be understandable without reference to the body of the manuscript. The abstract must be on sheets separate from the manuscript. Proof should be corrected immediately on receipt and returned to the editor. Authors should leave forwarding addresses if they move from the address sent with the manuscript. Reprints should be ordered when the proof is re- turned. Address all manuscripts to Stanley L. Welsh, Depart- ment of Botany, Brigham Young University, Provo, Utah 84601. MU& COMR ZOOL. LIBRARY Brigham Young University FEB 19 1971 Science Bulletin harvard UNIVK«Olt/ A COMPARATIVE STUDY OF THE HEAD AND fHORACIC OSTEOLOGY AND MYOLOGY OF THE KINKS EUMECES GILBERTI VAN DENBURGH AND EUMECES SKILTONIANUS (BAIRD AND GIRARD) by David F. Nash and Wilmer W. Tanner BIOLOGICAL SERIES— VOLUME XII, NUMBER 2 DECEMBER 1970 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Stanley L. Welsh, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: Vernon J. Tipton, Zoology Ferron L. Anderson, Zoology Joseph R. Murdock, Botany Wilmer W. Tanner, Zoology Ex officio Members: A. Lester Allen, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Chairman, University Publications I The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from Publication Sales, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for library exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 84601. Brigham Young University Science Bulletin a comparative study of the head and thoracic osteology and myology of the jKinks eumeces gilberti van denburgh and eumeces skiljonianus (baird and girard) by David F. Nash and Wilmer W. Tanner BIOLOGICAL SERIES— VOLUME XII, NUMBER 2 DECEMBER 1970 t TABLE OF CONTENTS Page LIST OF FIGURES i INTRODUCTION 1 MATERIALS AND METHODS 1 OSTEOLOGY 2 General Description of the Skull 2 Description of the Skull Elements 2 General Description of the Mandible 5 Description of the Mandible Elements 6 Description of the Hyoid Apparatus 6 MYOLOGY 7 General Description of the Myology 7 Description of the Myology 7 DISCUSSION 30 Osteology 30 Myology 30 SUMMARY AND CONCLUSIONS 31 ACKNOWLEDGEMENTS 31 LITERATURE CITED 32 LIST OF FIGURES Figure Page 1 . Dorsal view of skulls. A. Eumeces gilbert! (9.2X) (SSC 303). B. Eumeces skilton- iainis (9.3X) ( BYU 3 1 960) 8 2. Lateral view of skulls. A. Eumeces gilberti (8.9X) (SSC 303). B. Eumeces skil- toniamis (9.3X) ( BYU 3 1 960) 9 3. Ventral view of skulls. A. Eumeces gilberti (9,2X) (SSC 303). B. Eumeces skil- r(wwnMs(9.6X) (BYU 31961) 10 4. Mandible. A. Medial view. B. Lateral view. C. Dorsal view, fwmecex x/:;7ro/Ha/;ws (9.3X) ( BYU 1 2468) 11 5. Dorsal view of hyoid apparatus. Eumeces gilberti (5. OX) (BYU 31956) 12 6. Musculature dorsal view. A. Superficial depth. B. Fust depth. Eumeces gilberti (3.7X)(BYU3I956) 18 7. Musculature dorsal view. A. Second depth. B. Third depth. Eumeces gilberti (3.7X) (BYU 31956) 19 8. Musculature dorsal view. A. Fourth depth. B. Fifth depth. Eumeces gilberti (3.7X) (BYU 31956) 20 1 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN 9. Musculature lateral view. Superficial depth, liumeces gilbert! {?<.1\) (BYU 31956) 21 10. Musculature lateral view. First depth. A. Ewneces gilberti {3,.1X) (W^\J 31956). B. Eumeces sklltonianiis (5.7.X) (BYU 12474). Showing anterior and posterior bundles of the depressor mandibularis 22 11. Musculature lateral view. Second depth. £';/;He(7eig;7Z7e/-//(3.7X) (BYU 13956) . 23 12. Musculature lateral view. Third depth. Eumeces gilberti {3, JX) {BY\J 31956) ... 24 13. Musculature lateral view. Fourth depth. Eumeces gilberti (3.7X) (BYU 31956) . . 25 14. Musculature lateral view. Filth depth. Eumeces gilbert! {3.7X) {BYU 31956) ... 26 15. Musculature ventral view. A. Superficial depth. B. First deplh. Eumeces gilbert! (3.7X) (BYU 31956) 27 16. Musculature ventral view. A. Second depth. B. Third depth. Eumeces gilbert! (3.7X) (BYU 31956) 28 17. Musculature ventral view. A. Fourth depth. B. Fifth depth. Eumeces gilbert! (3.7X) (BYU 31956) 29 A COMPARATIVE STUDY OF THE HEAD AND THORACIC OSTEOLOGY AND MYOLOGY OF THE SKINKS, EUMECES GILBERT/ VAN DENBURGH, AND EUMECES SKJLTONIANUS (BAIRD AND GIRARD> by David P. Nash and Wilmer W. Tanner* INTRODUCTION A. F. Weigmann (1834) described the genus Eumeces in Herpetologica mexicana and E. pavimeu- tatiis (Geoffroy-St. Hillaire) was designated genotype. The two species involved in this study belong to the skiltomanus group which are characterized by the presence of two dorsolateral and two lateral light lines on the body. Eumeces skiltunianus is found in British Columbia, Idaho, Montana, Utah, Nevada, northern Arizona, California, and Baja California. Baird and Girard (1852) described E. skiltonianiis and designated Oregon as the type locality. Van Denburgli (1896) described a new species from southern California which he called Eumeces gil- berti. Camp, (1916) disagreed with the distinctness of this new species and considered all western skinks to belong to the same species, skiltonianus. Since the publication of Camp's paper the distinctness of these two species has been verified by more thorough inves- tigations, which have been based on geographical distribution, size, color pattern, scale counts, and other external morphological characters. Taylor (1935) was the first to do a complete study of the genus Eumeces and he recognized these two species; Smith (1946) further characterized them in his Hand- book of Lizards; and Rodgers and Fitch (1947) set forth the distribution of gilberti and the distinguish- ing characters of that species. Tanner (1957) charac- terized skiltonianus, designated its type population, and described two new subspecies. The papers of Rodgers and Fitch, and Tanner set forth the external anatomical characteristics of these species. They also recognized that all subspecies of skilton- ianus exhibit a striped pattern; whereas, only the immature forms o( gilberti have the striped pattern. Adults of gilberti are patternless and much larger than the adults of skiltonianus: the average snout-vent length for gilberti is about 88 mm and for skilton- ianus about 64 mm. The studies of external laxo- nomic characters of these two species seem to be complete; therefore, it is the purpose of this study to further clarify their taxonomic and phylogenetic relationships by comparing and examining the oste- ology and myology of their head and thorax regions. Extensive literature reviews on general and specific osteology and myology have been done by other authors: Robison and Tanner (1962), Oelrich (1956), Avery and Tanner (1964), Jenkins and Tanner (1968), and Fisher and Tanner (1970). Only those references which are relevant to this report are given. The skulls of ten species within the genus Eumeces including gilberti, were compared by Kingman ( 1932). To date the only myological work within this genus available to the author is the publication by Edgeworth (1935), in which he describes E. schnidcri. Materials and Methods The subspecies used in this study are E. gilberti placerencis (5), E. g. nibricaudatus (3), E. skilton- ianus skilonianus ( 10), and E. s. utahensis (5). The specimens of E. gilberti have the following numbers: Brigham Young University (31956-58 in- clusive, 32622, and 32623); Sacramento State College (303, 437, and 528). The specimens of E. skilton- ianus have the following numbers: Brigham Young University (8721, 12467, 12468, 12472, 12474, 13750, 13756, 21939, and 31959-65 inclusive). Osteological material was prepared by skinning the specimens, removing the superficial muscles using forceps, placing the specimens in a 50% liquid bleach solution for 30 to 60 seconds and rinsing them after- wards in clear water. The last three procedures were alternated until the desired results were obtained. Care was taken not to destroy the osteological ele- ments when removing the muscles with forceps nor to leave the specimens in the bleach solution for an extended length of time. Illustrations were produced I ''Department of Zoology, Brigham Young University, Provo, Utah. 1 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN by photographing the skulls througli a binocular scope using 400 ASA, 35 mm black and white tllm. The pictures were then enlarged to ten inches and traced onto mylar tracing sheets using a binocular scope for detail. A full description of each skeletal element of E. gilberti is given under "G." If differences in skilton- ianus are noted, they are given under "S." A mini- mum of five skulls of each species is used for the description and comparative differences. All elements are discussed in alphabetical order. Preserved specimens were carefully skinned for myological comparisons. Illustrations were obtained by taking measurements from the preserved speci- mens, enlarging them onto tracing paper, and then transferring them onto mylar tracing sheets. A full description of each muscle for E. gilberti is given under "G." If differences in skiltonianus are noted, they are given under "S." A minimum of four specimens of each species is used for the description and comparative differences. All elements are dis- cussed in alphabetical order. OSTEOLOGY General Description of the Skull Generally the skull is delicate and for the most part completely ossified. The extremities are bounded by the rostrum, the occipital condyle, and the quad- rates laterally. The rostrum is rounded and contains large narial openings which are bounded by the premaxillae, the nasals, and the prefrontals. The artic- ulation between the nasals and the frontals marks an area of abrupt change in elevation with the frontals being higher. Anterodorsally the frontals contact the maxillae thus preventing the nasals from articulating with the prefrontals. The frontals form the medial margin of the orbits; whereas, the prefrontals and postfrontals form the anterior and posterior margins of the orbits respectively. The parietal is slightly rugose and contains a centrally located parietal fora- men. Its posterior margin is dorsoventrally slanted and has two lateral and two medial processes. The postorbitals articulate with the jugais either separ- ately or in conjunction with the postfrontals. In some specimens the postorbitals are reduced to such an extent that only a small slender bone remains. The squamosals, however, do not come in contact with the jugais. A supratemporal fontanelle located pos- teriorly is bordered by the parietal, postfrontal, postorbital and squamosal elements. The occipital and otic regions are posteroventral to the parietal and are generally highly fused. The fusion of elements does not always correspond directly with the size of the skull. The supraoccipital forms an ascendens process with the parietal. The basioccipital is gen- erally fused with the other occipital elements, with the basisphenoid and also with the prootics. The para- sphenoid is a slender pointed element attached to the ventral border of the interorbital septum and extends anteriorly. The anteromedial limb of the pterygoids contains a number of pleurodont teeth. The palatines contact only the maxillae, vomers, and pterygoids. Two pairs of Jacobson's organs are housed in the vomers. The jugais have small spines extending poster- iorly on their lateroventral margins. The maxillae and premaxillae contain cylindrical pleurodont teeth. The right premaxilla is larger and contains one more tooth than the left premaxilla. Two orbital fenestrae, two infraorbital fenestrae, two infratemporal fenestrae, and two posttemporal fenestrae are located in the skull. Description of the Skull Elements Basioccipital. (Figs. 2 and 3) G. The basioccipital contributes to the cranium floor, is roughly hexagonal in shape, and forms the basal portion if not all of the occipital condyle. Kingman (1932) reported that the exoccipitals formed the lateral portions of the occipital condyle. In the specimens studied, however, fusion obscures the possible presence of sutures with the exoccipitals. Articulation with the basisphenoid anteriorly and the prootics laterally are generally by fusion. If sutures are present, the basioccipital-basisphenoid suture is irregular and the basioccipital-prootic suture extends posteriorly and then dorsally, separating the prootic and opisthotic portions of the otic capsule and termi- nating on the ventral margin of the fenestrae ovali. The prominent basioccipital processes project ventro- laterally and in some cases have an epiphysis. These processes appear to be formed in part by the exocci- pital and the paroccipital elements. Anterior to the above processes smaller protuberances are found which make up part of the otic capsules. Basisphenoid. (Fig. 3) G. The basisphenoid contributes to the anterior portion of the cranial tloor. Anteriorly it contacts the parasphenoid. Anterolaterally the basipterygoid processes emerge and enlarge distally into tlangelike processes which articulate with the pterygoids by a cartilaginous pad. Lateral contact is generally made with the prootics by fusion of the two elements. The sella turcica is posterior to the basipterygoid pro- cesses. A small foramen located on the lateral surface of the basisphenoid forms a canal with the prootics. BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS Ectopterygoid. (Figs. 1 and 3) G. The ectopterygoids have dorsal and ventral processes which enclose the lateral processes of the pterygoids. Contact is also made with the maxillae and jugals. These bones do not articulate with the palatines. Epiotic. (Not figured) G. The epiotics are part of the otic capsule and enclose the dorsal parts of the two vertical semicir- cular canals (JoUie, 1960). They appear as large rounded protuberances when viewed from the fora- men magnum and extend medially and dorsally. They fuse inside the braincase with the supraoccipital, the opisthotics, t; ". paroccipital processes, and the exoc- cipitals. Epipterygoid. (Fig. 2) G. The epipterygoids are relatively straight ossified rods extending from a well developed socket on the dorsal margin of the pterygoids to the anterior descending processes of the parietal and posterior descending processes of the prootics. They have a slight lateral cant ventrodorsally. Exoccipital. (Fig. 2) G. The exoccipitals form the ventrolateral margins of the foramen magnum. They are fused with the supraoccipital dorsally and the basioccipital ventrally. Fusion in this area makes it difficult to determine if these bones contribute to the occipital condyle. Laterally they fuse with the paroccipital processes. Anteriorly they form the posterior margin of the fenestrae rotunda. One, two, or three hypoglossal nerve foramina are located in the exoccipitals near the junction with the paroccipital processes. The jugular formina are bordered by these latter two bones. Frontal. (Figs. 1 , 2, and 3) G. The frontal element forms the cranial roof between the orbits. The anterior lateral processes of the frontal always contact the maxillae. Posteriorly articulation is with the postfrontals and parietal. The suture with the parietal is slightly convex. The des- cending processes in the orbits are slender and con- tact the dorsal palatine limbs medial to the prefront- als. S. Anteriorly this bone articulates with the nasals, prefrontals, and may contact the maxillae depending upon the extent of the anterior lateral processes of the frontal. Interorbital Ossification. (Fig. 2) This is an ossified element found in both species, and which is connected anteriorly to the internasal septum. It extends posterior and slightly ventral, and is found within the interorbital septum. In one speci- men of E. gilherti this element extends posteriorly as far as the braincase. In both species it varies from short to long and from triangular to almost rectangu- lar in shape. Jugal. (Figs. 1, 2, and 3) G. The jugals form the lateroventral margin of the orbits and the anterior margin of the infratemporal fenestrae. Their dorsoposterior margin generally artic- ulates with the postfrontals and always with the post- orbitals. They do not contact the squamosals. These bones articulate with the ectopterygoids ventrally. On their lateroposteroventral surface is a short posterior spine. Kingman (1932) states that this projection suggests "the place of continuance of some element in the past." Anteroventrally the jugals are sand- wiched between the lateral and medial processes of the maxillae. Lacrimal. (Figs. 1 and 2) G. The lacrimals are small rectangular splints of bone forming a portion of the ventroanterior margin of the orbits between the prefrontals and the max- illae. They are not always fused with the prefrontals. On their posteroventral surface there is an arch of this element forming one to two foramina which lead into the large lacrimal duct anteriorly. Maxilla. (Figs. 1 , 2, and 3) G. The maxillae are the tooth-bearing bones of skull with from 15 to 20 teeth per side, and an aver- age of 19. Replacement teeth are present. The teeth are pleurodont. The medial surface is level with the palatine. The teeth are cylindrical, homodont, rounded at the tips, and extend slightly laterally. Dor- sally the maxillae articulate with the nasals and with the frontals posteriorly. Anteriorly in the narial open- ings the maxillae send a slender process inward to meet the anterior process of the septomaxillae. The maxillae articulate with the premaxillae in the floor of the narial openings, and on the ventroanterior sur- face of the skull they overlap the lateral processes of the premaxillae. Posteriorly, articulation is with the prefrontals, the lacrimals, and the jugals. Ventrally, the maxillae articulate with the premaxillae and the vomers anteriorly, the palatines medially by the posterior palatine processes, the jugals posteriorly, the ectopterygoids medially, and the prefrontals in the anterolateral portion of the orbits. The maxillae are not always fused with the lacrimals. The lateral surfaces of the maxillae contain six to eight supra- labial foramina of varying sizes and shapes. S. The maxillae may or may not contact the frontal posteriorly. They have a range of 17 to 20 teeth per side, with an average of 18 and contain four to seven supralabial foramina. BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Nasal. (Figs. 1 and 2) G. The nasals cover most of the anterior dorsal surface of the skull. They contact the premaxillae and the narial openings anteriorly, the maxillae laterally, the frontal posteriorly, and do not articulate with the prefrontals. Three to five nasal foramina are present, the posterior one being located near the junction of the prefrontals, nasals, and frontal elements. Sutures with the frontal are higlily irregular and lobed. S. The nasals may or may not contact the pre- frontals. Two to four nasal foramina are generally present. The suture with the frontal is not as irregular as that of gjiberti unless articulation of the prefrontal and nasal elements is absent. Opisthotic. (Fig. 1 ) G. The opisthotics form the posterior portion of the otic capsule; the anterior portion being formed by the prootics (Smith, I960). A suture separating these two bones terminates at the ventral margin of the fenestrae ovali. On the dorsal surface the opisthotics are fused with the epiotics and the prootics. The posterior portion of the opisthotics or the paroccip- ital processes connect the occipital region to the quadrate region. These processes are wide and gener- ally fuse with the supratemporals and the quadrates and posteroventrally they contact the prootics. OsPalpabra. (Fig. 1) G. The OS palpabrae are irregular tear-shaped bones which are attached to the dorsal posterior margin of the prefrontals by connective tissue. Palatine. (Figs. 1, 2, and 3) G. The palatines have a ventral and a dorsal limb. Posteriorly these two limbs join each other and artic- ulate with the medial limb of the pterygoids. The Ventral ■ hmbs articulate with the maxillae by the posterior palatine processes. The dorsal limbs are arched upward forming the wall of the internal nares and contact the descending processes of the tVontals and the prefrontals. Anteriorly this limb^ articulates with the vomers. The dorsal limbs contact each other for about one-fourth their length before separating posteriorly to jdin the ventral limbs. The separated halves form a sliglit ventral ridge. The palatine canals are located in the ventral limbs just medial to the articulation with the maxillae. Parasphenoid. (Figs. 1 and 2) G. This element is a cartilaginous tapered rod extending in an anterior direction from the center of the basisphenoid and forming the ventral border of the interorbital septum. postfrontals and the postorbitals laterally. The latter articulation is not present if the supratemporal fon- tanelle separates these two elements. The postero- lateral processes contact the squamosals, the supra- temporals, the paroccipital processes, the opisthotics, and the prootics. Medially two shorter processes extend posterior and lateral to the ascendens process of the supraoccipital forming a metakinetic type of articulation (Romer, 1956). The posterior edge of the parietal including the medial and lateral processes slant ventrally with the result that the main body of the parietal is higher than the dorsal occipital region. The lateral borders of the parietal are slightly curved inward. Ventrally the descending wall laminae are drawn out and articulate with the anterodorsal border of the epipterygoids. A parietal foramen lies on the median line near the center of the bone. Articulation with the quadrate is absent. Postfrontal. (Figs. 1 and 2) G. The postfrontals contribute to the posterior margin of the orbits. Articulation is with the parietal medially and frontals anteriorly. Laterally contact is made with either the postorbitals or the postorbitals and the jugals. The suture with the postorbitals may be fused in some places. Postorbital. (Figs. 1, 2, and 3) G. The postorbitals vary in width and length, and frequently show a reduction of relative size when compared to other adjacent bones. They are found lateral to the postfrontals and medial to the squa- mosals. Anteriorly they articulate with the jugals and the postfrontals. The suture with the postfrontals may be fused in some places. Posteriorly these ele- ments may articulate with the parietal depending upon the size of the supratemporal fontanelle. They do not articulate with the supratemporals. S. The postorbitals vary in size, but are only occa- sionally reduced to the same degree as mgilbeni. Prefrontal. (Figs. 1 and 2) G. The prefrontals articulate with the frontals medially and with the maxillae laterally. These ele- ments have a large anterior orbital process which descends to contact the palatines, the frontals, and the lacrimals. In the anterior ventral portion of the orbit, articulation is made with the maxillae dorsally. A small foramen is either located in the descending process of the prefrontals or between the prefrontals and frontals. S. The prefrontals articulate with the frontals medially, but may or may not articulate with the nasals. Parietal. (Figs. I, 2, and 3) Premaxilla. (Figs. 1,2, and 3) G. The parietal is a large posterior dorsal element G. The right premaxilla is the larger and contains which articulates with the frontal anteriorly and the four pleurodont teeth as compared with three teeth BIOLOGICAL SERIES. VOL. 12. NO. 2 OSTEOLOGY AND MYOLOGY Ol- THE SKINKS on the left element. This unequalness is produced by the extension of the midline suture into the left ele- ment. These elements may be fused in places. The premaxillae border the narial openings medially and the maxillae and vomers ventrally. Prootic. ( Figs. 1 , 2, and 3) G. The prootics are very complex elements form- ing a major portion of the cranium and the otic capsules. Ventrally they articulate, sometimes by fusion, with the basisphenoid and the basioccipital. If these elements are not fused, the suture continues posterior to the ventral margin of the fenestrae ovali. The lateroventral portion of the prootics is thin and deeply concave. The lateral wall of the prootics contain a canal which comes from the basisphenoid and continues to the otic capsule. Posteroventrally fusion is made with the paroccipital processes. The anterior descending processes contact the postero- dorsal margin of the epipterygoids. Pterygoid. (Figs. 1,2, and 3) G. The pterygoids are "Y" shaped bones sliglitly less than one-half the length of the skull. The anterior limbs are large, heavy, and articulate with the pala- tines by an overlapping suture. These limbs have a depression in the center containing four to ten pleur- odont teeth. The teeth are attached to the lateral side, face somewhat medially, may be in a single row, or in irregular groups; and they are more sharply pointed and smaller than those found elsewhere. The lateral limbs are sandwiched between the ventral and dorsal processes of the ectopterygoids. At the junc- tion of these two hones the lateral surface of the pterygoids is flanged into the coronoid process. The posterior limbs curve inward anteriorly and then outward posteriorly to articulate with the quadrate, and then extend sliglitly beyond the quadrate articu- lation. In the middle of the pterygoids the basiptery- goid processes articulate by a cartilaginous pad. Dorsal to the basipterygoid processes there is a socket into which the epipterygoids fit. Anterior to these processes, the pterygoids are highly concave medially and posterior to the above processes the pterygoids are highly convex medially. Quadrate. (Figs. 1 , 2, and 3) G. The quadrates are large units articulating with the squamosal, supratemporal, and paroccipital elements posteriorly; and with the pterygoids anter- iorly. The articulating surface with the pterygoids is saddle shaped. Septomaxilla, (Fig I) G. The septomaxillae are inverted cup-shaped structures lying dorsal to Jacobson's organ and ex- tending anteriorly. They contact the premaxillae and the maxillae in the tloor of the narial openings. There is a thin ridge of bone extending upward and outward from the anterior portion of the septomaxillae which is joined by an inward process from the maxillae. Stapes. (Fig. 3) G. The stapes are thin cylindrical bones with flared bases which fit into the fenestrae ovali. The lateral ends are somewhat expanded. These bones are directed posterolaterally and extend into the quad- rate region. Supraoccipital. (Fig. I) G. The supraoccipital forms the superior margin of the foramen magnum and generally fuses with the exoccipitals ventrally. Laterally they fuse with the opisthotics and with the epiotics inside the braincase. A slight midsagittal ridge is formed and the ascendens process extends posteriorly under the parietal and forms a metakinetic articulation with the medial posterior processes of the parietal. Supratemporal. (Figs. 1, 2, and 3) G. The supratemporals are small slender bones wedged between the squamosals and the parietal dorsally. Ventrally they contact the quadrates and the paroccipital processes. Squamosal. (Figs. I, 2, and 3) G. The squamosals are larger than the postorbitals and have a medial expansion of variable width. They are separated from the jugals by the postorbitals, and posteriorly they curve laterally and downward to articulate into a small depression of the quadrate. Posterior articulation is with the supratemporals and the parietal. Vomer. (Fig. 3) G. The vomers form the anteroventral surface of the skull. They are rolled laterally with the medial area being the more ventral. They are separated from each other by a deep medial longitudinal groove centrally. Anteriorly they become flat, constricted, and articulate with the premaxillae and the maxillae. They articulate posteriorly with the palatines. An- terior to the latter articulation two ventrally pro- jected toothlike structures are present. The right and left halves are for the most part fused along their entire length. Each half contains one pair of Jacob- son's organs. General Description of the Mandible Since the mandibles of these two species are simi- lar except for the differences described below, figures of both species are not given. E. skiltonianus (Fig. 4) was chosen because of the superior illustration, and is typical for either species. Each ramus is slightly bowed laterally. The rami BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN are united anteriorly by the mental symphysis and articulate posteriorly with the quadrates. An open sulcus of Meckel's cartilage is located on their ventro- medial surface. Each ramus consists of seven ele- ments; the angular, the articular, the coronoid, the dentary, the prearticular, and the surangular. The surfaces are smooth. The dentary bears a single row of pleurodont teeth. The retroarticular processes are thin and spoonshaped. The prearticulars are generally fused with the articulars. Description of the Mandible Elements Angular. (Fig. 4) G. The angular is located on the ventral surface of the mandibular ramus. It is projected dorsally in an anteroposterior direction. On the lateral surface of the ramus it contacts the surangular dorsally, the dentary anteriorly, and the prearticular posteriorly. On the medial surface of the ramus the angular articu- lates by a forward splint of bone between the dentary and the splenial elements. Dorsally, articulation is with the prearticular. This element contains a small foramen anteriorly. Articular. (Fig. 4) G. The articular forms the articulating surface of the mandible for the quadrate. This element is an ossification of Meckel's cartilage and is bordered by dermal elements. The articular process extends medi- ally. The articular is generally fused with the surangu- lar dorsally and with the prearticular ventrally. It does not contact the angular. Coronoid. (Fig. 4) G. The coronoid is an inverted "V" shaped ele- ment which is centrally located and projects dorsally. It has one lateral and two medial processes. The lat- eral process contacts the surangular ventrally and the dentary anteriorly. The anterior medial process lies along the posterior portion of the dentary. The pos- terior medial process contacts the surangular and the prearticular. Between the two medial processes a well developed coronoid fossa is found which contains anterior portions of the surangular and prearticular. These two latter elements are sometimes fused. Dentary. (Fig. 4) G. The dentary is the tooth-bearing bone of the mandible. This element extends posteriorly to a dis- tance equal to the posterior medial process of the coronoid. Nineteen to 23 pleurodont teeth, with an average of 21, are ioeated on the dorsal surface of the dentary. Meckel's cartilage is open ventrally and is equal in length to no more than one-half that of the dentary. The dentary contacts the splenial at the posterior end of Meckel's cartilage. Anterior articu- lation is with the anterior process of the coronoid. the surangular, and the angular. The lateral surface contains 3 to 6 mental foramina of various sizes and shapes. S. The dentary contains 18 to 22 teeth, with an average of 21, and contains 3 to 7 mental foramina. Prearticular. (Fig. 4) G. The prearticular is a major component of the medial surface of the ramus. It is generally fused with the articular posteriorly and sometimes fused with the surangular dorsally. Ventrally it comes in contact with the angular. This element forms the retroartic- ular process of the ramus. The mandibular fossa is located posteriorly and is formed by the surangular dorsally and the prearticular ventrally. Splenial. (Fig. 4) G. The splenial lies on the medial anterior portion of the ramus, ventral to the dentary. It sends forward two small bones which enclose the posterior portion of the open sulcus of Meckel's cartilage prior to artic- ulation with the dentary. This element extends posterior to the posterior margin of the posterior medial process of the coronoid. The splenial contains a foramen just posterior to the open portion of Meckel's cartilage. Surangular. ( Fig. 4) G. Laterally the surangular covers the dorsal por- tion of the ramus between the dentary and the artic- ular. It is generally fused with the articular poster- iorly and forms the anterior portion of the articular process. It contacts the coronoid dorsally, the den- tary anteroventrally, and the angular ventrally. On the medial surface it is sometimes fused with the pre- articular. Posteriorly it contributes to the mandibular fossa with the prearticular. On the lateral surface two foramina are located. Description of the Hyoid Apparatus The general morphology of the hyoid apparatus in both species is similar. Variations, however, were noted in the average ratios of the length of the ento- glossal process to either the length of the proximal segment of the hyoid cornu or the length of the ceratobranchial 2, with gilbeni being greater in both cases. A sufficient number of specimens was not avail- able for statistical analyses. The hyoid apparatus is represented by H. gilbeni in Figure 5. Hyoid Apparatus. (Fig. 5) G. Unless stated differently, all elements are carti- laginous and higlily calcified. The main body or the corpus is prolonged anteriorly into the entoglossal process. The latter is imbedded in the tongue muscu- lature. The remainder of the corpus has two postero- lateral projections which are connected to the other BIOLOGICAL SERIES. VOL. 12, NO. OSTEOLOGY AND MYOLOGY OF THE SKINKS elements. The hyoid cornu consists of two segments. The posterior portion of the proximal segment is almost completely ossified and articulates between the corpus and the ceratobranchial 1. The remainder of this segment is tubular and connects to the distal segment of the hyoid cornu. The distal segment is initially flat and wide; it then becomes thicker and somewhat tubular, and distally it again becomes flatter. This segment has a calcified core throughout its length. The ceratobranchial fied except near its articulation Medially it articulates with the the hyoid cornu. Initially it is center it becomes wider, and again and has a flared articulat chial. The epibranchial is a element. The ceratobranchial 2 indistinguishably to the corpus. 1 is completely ossi- with its epibranchial. proximal segment of tubular, toward the distally it is tubular ion with its epibran- slender and pointed is tubular and fused MYOLOGY General Description of the Myology This description reflects the general myological condition of both species. Differences are discussed later. Ventrally the intermandibularis anterior is sub- divided, interdigitates with the geniohyoideus, and is continuous with the intermandibularis posterior. The sternohyoideus is separable into a superficial and a profundus layer; the medial border is partially separ- ated from its partner by an elliptical space. This space is covered by fascia; but when uncovered, portions of the trachea, thyroid gland, clavodeltoidus, and the clavicle can be seen. The pectoralis is a large muscle covering the sternum and the scapula; posteriorly this muscle interdigitates with the rectus abdominus. Ventrolaterally the pectoralis is covered by an an- terior slip of the rectus abdominus which extends almost to the insertion of the pectoralis. A ligamen- tous fascia covers a large portion of the pectoralis and is connected to the anterior slip of the obliquus abdominus rectus posteriorly, to the lateral process of the interclavicle, and to the muscles immediately anterior to this process. There is only one mandibulo- hyoideus present. Both the clavodeltoideus and the coracobrachialis longus have a dorsal and a ventral origin on the clavicle and the scapula respectively. In the adductor muscle complex, the adductor mandib- ularis externus medius, the adductor mandibularis externus profundus, and the adductor mandibularis posterior are at equal depth. The second of these muscles extends through the posttemporal fenestrae to partially originate from the opisthotic and prootic elements. No levator anguli oris is present. The depressor mandibularis inserts on the retroarticular process and intermandibularis posterior, and in some specimens is separable into two bundles. The tra- pezius varies in thickness and in some specimens can also be separated into two bundles. The trapezius and the sternocleidomastoideus interdigitate ventrally to insert along with the sternohyoideus onto the inter- clavicle lateral process. There is both an anterior and a posterior scapulohumeralis. The obliquus abdom- inus externis muscle is divided into a superficial and a profundus layer. Dorsally the rectus capitis posterior is separated into an anterior and a posterior bundle; the former has a medial and a lateral slip. The obliquus capitis magnus and the longissimus cervicis are joined to the spinalis capitis by a myosepta. No longissimus capitis is present. Description of the Myology M. Adductor Mandibularis Externis Medius. (Figs. 6, 8, and 10) G. This muscle is deep to the adductor mandib- ularis externus superficialis from which it is separated with difficulty. It lies dorsal to the adductor mandib- ularis externis profundus and is separated anteriorly from this muscle by a tendon from the bodenapo- neurosis. These two muscles interdigitate posteriorly. The muscle originates from the prootic and opisthotic elements deep to the spinalis capitis and lies between the medial and lateral slips of the rectus capitis posterior. The muscle fibers extend in a lateral direc- tion and as they enter the posttemporal fenestrae turn anteroventrally, and have other origins from the beveled surface of the parietal, ventral surfaces of squamosals, postorbitals, jugals, and the dorsal surface of the quadrates. The fibers converge anter- iorly to insert onto the coronoid by way of the bodenaponeurosis. M. Adductor Mandibularis Externis Profundus. (Fig. 10) G. This muscle lies ventral to the adductor mandibularis externis medius, and dorsal to the adductor mandibularis posterior. It is separated anter- iorly from the latter two muscles by a fascia. They interweave posteriorly. The muscle originates from the anterior and anterodorsal surfaces of the quad- rate, and from the lateral surface of the prootic. Insertion is onto the bodenaponeurosis and also onto the medial base of the coronoid. M. Adductor Mandibularis Externis Superficialis. (Figs. 6, 9, and 15) G. This thin superficial muscle is scarcely distin- guishable from the deeper muscles. The muscle is separated by the bodenaponeurosis that extends from URIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN CO — — o 2 "O a. •J c o = ■ - o !* ^ = ■-00 ■- o. a. o. i rl n £ I r3 73 73 i Q. n. Q. G. i_ u.^ Oj:;, CTfU 3 33 iU - J5 _2 rj o>- rn aa OOOD.o.a.a.a.BL. Ju ^ ■^ .i a. .il 3 -5 = H - > 5 3 o Q a. ■ x-4 • 0000.0.0.0.0.^ 0 0 ffl n 0 73 _ £ _3 U. "q. 0 '*- at ra ^. >", :> J- ^ -J szzo w w t. S=i -, J s z ; .0 ;o BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS OQ — 3 'o O _ -.-J - '^ Li o O"^ 73 TZ ' ■ ^ "^ !/! ^ D. c ir. a '= ~ ^ ;; .^ t; t: £ ii o .. — ::! D-ti- a-p-=3 a.££al£Ov;in'J5 d> 23-Jii;^x-,-jSZOa.a.a.o.a.B.o-a.buO'iy5c«c«. BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN O 60 -± o ri 73 — ',V: " o o — a a. ■f I S5-S i 2 2 ^ a. a. D. Ct/i C/5C/1 > 2 o t o [jL, o ."^ -^ r: to c T3 ■— — Um cd ■ — O H _ *- c >>Z ,■:: °- — - *-- ^ i— ') -1 o-a.c o - 2 - o 5 2 2 5"^ SO O c/l O 7 f-^ ^ .^ ■^- BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS 11 D A Angular AF Angular Foramen AR Articular C Coronoid D Dentary MC Meckel's Cartilage MEF .... Mental Foramen MF Mandibular Foramen PA Prearticular S Splenial SA Surangular SAF .... Surangular Foramen SF Splenial Foramen MC AF A SA MF AR PA B A. Medial view B. Lateral view C. Dorsal view Figure 4. Mandible. Eumeces skiltoniamis (9.3X) (BYU 12468) BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN C Corpus CBj .... Ccratobranchial 1 CBf .... Ceratobranchial 2 EPT Entoglossal Process H| Proximal Segment pt tli Hyoid Cornu . Hi. ..... Distal Segment of tlie Hvoid Cornu ' Figure 5. Dorsal view of hyoid apparatus. Eiimeces gilbciti (5. OX) (BYU 31956) the coronoid and almost to the posterior margin of the muscle. It arises from the ventral surface of the squamosal, the dorsoanterior surface of the quadrate, and the anterior margin of the ju^al. Insertion is into the bodenaponeurosis and on the lateral surface of the surangular. M. Adductor Mandibularis Posterior. (Fig. 10) G. This inuscle is deep to the adductor mandibu- laris'CXternis superficialis and ventral to the adductor mandibularis exteinis profundus. It is separated anter- iorly from the latter muscle by a tendon; posteriorly the fibers of these two muscles interweave. Origin is on the anterior surface of the quadrate with insertion from the anterior surface of the quadrate with inser- tion on the medial dorsal surface of the surangular, the rt^andibular fossa, and the most anterior surface of the articular. M. Anconaeus Coracoideus. (Fig. 6) G. The anconaeus coracoideus is a small dorsal muscle arising by a tendon, posterior to the insertion of the latissimus dorsi, from the sternoscapular ligament and uniting with the anconaeus scapularis prior to insertion onto the olecranon process of the ulna. This muscle is found posterior to the anconaeus scapularis and anterodorsal to the anconaeus medialis humeralis. M. Anconaeus Humeralis Lateralis. (Fig. 15) G'. This muscle originates medial to the insertion of the scapulodeltoideus and the clavodeltoideus from the proximal head and the anterior surface of the humerus. It then unites with the anconaeus scapu- laris to insert onto the olecranon process of the ulna. Some fibers will interdigitate distally with the brachi- alis inferior. This muscle is located anterior to the anconaeus scapularis and dorsal to the brachialis in- ferior. M. Anconaeus Humeralis Medialis. (Fig. 6) G. This muscle originates from the proximal end of the humerus, below the sternoscapular ligament, and from the posterodorsal surface of the shaft of the humerus. It then unites with the other anconaeus muscles for a common insertion onto the olecranon process of the ulna. This muscle lies posterior to the anconaeus coracoideus and anterior to the coraco- brachialis longus. M. Anconaeus Scapularis. (Figs. 6 and 1 5) G: This large upper-arm muscle originates from the sternoscapular ligament dorsal to the scapulodel- toideus and clavodeltoideus. It then follows the humerus to join tendonously with the other ancon- aeus muscles. The inuscle is divided into two bundles. The anterior bundle fibers extend first lateroven- BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS 13 trally, then parallel to the humerus, and then ventro- medially. This muscle is posterodorsal to the ancon- aeus humeralis lateralis and anterior to the anconaeus coracoideus. fascia to the trapezius, the sternocleidomastoideus, and the pectoralis. In some specimens fat tissue is found in abundance on the ventral and lateral sur- faces. M. Biceps. (Figs. 15 and 17) G. The biceps is a large ventral muscle that inserts tendonously onto the proximal head of the ulna and radius. The muscle continues along the humerus and forms a broad tendon which is connected to the head of this muscle. The origin is from the medial ventral border of the scapula. On the humerus the muscle lies between the brachialis inferior and the coracobrach- ialis longus. Just lateral to the deltopectoral process an anterior slip is attached tendonously to this pro- cess. On the scapula a thin layer of the supracora- coideus covers the anterior portion of this muscle. M. Brachialis Inferior. (Fig. 17) G. This ventral muscle originates friim the antero- lateral surface of the deltopectoral process and the anteroventral surface of the shaft of the humerus. Insertion is primarily onto the radius; however, some fibers interdigitate distally with the biceps to also insert onto the ulna. This muscle is located anterior to the biceps and posterior to the anconaeus humer- alis lateralis. M. Branchiohyoideus. (Fig. 17) G. This straplike muscle originates from the lateral three-fourths of the ceratobrachial I and inserts onto the posterior and ventral surlaces of the central por- tion of the hyoid cornu. This muscle is superficial to the sternocleidomastoideus and trapezius laterally, and is strongly attached to these two muscles by con- nective tissue. M. Clavodeltoideus. (Figs. 6, 9, 12, 13, 15, 16, and 17) G. This complex muscle inserts by a tendon onto the deltopectoral process near the origin of the brach- ialis inferior and the insertion of the scapulodel- toideus. Some of the fibers interdigitate with the scapulodeltoideus prior to insertion. From the inser- tion 'the muscle fibers extend in two directions to originate from the clavicle. One bundle of fibers extends anteromedially under the clavicle bar, rotates ventrally and laterally to originate from the medio- ventral surface of the clavicle. The other bundle of fibers extends anteromedially under the clavicle- interclavicle bar to originate from the dorsomedial surface of the clavicle. M. Constrictor Colli. ( Figs. 6, 9, and I 5) G. This is a wide cervical muscle originating from the middorsal fascia and the tympanic fascia. Inser- tion is with its partner in the median fascia. This mus- cle is strongly attached laterally and ventrally by M. Coracobrachialis Brevis. (Fig. 17) G. This posterior arm muscle originates from the medioventral surface of the scapula with the fibers converging laterally to insert onto the deltopectoral process and onto almost the entire length of the posteroventral surface of the humerus. This muscle is located anterior to the coracobrachialis longus and is deep to the biceps. M. Coracobrachialis Longus. (Figs. 6 and 15) G. This muscle has a dorsal and a ventral slip which have a common insertion onto the ulnar pro- cess of the humerus. The ventral slip is considerably larger and originates from the posterolateroventral tip of the scapula. The dorsal slip originates from the dorsal surface of the scapula just posterior to the origin of the subscapularis 2. It leaves the scapula dorsal to the origin of the ventral slip and shortly terminates in a long slender tendon to insert onto the ulnar process. M. Costocoracoideus. (Not figured) G. This broad, thin muscle originates from the first sternal rib and extends anteriorly to insert onto the sternum and the sternoscapular ligament. M. Depressor Mandibularis. (Figs. 6, 9, and 15) G. This muscle originates from the middorsal fascia, the posterodorsal surface of the squamosal, and the most lateral surface of the parietal. The anterior muscle fibers converge centrally, attach to the posterodorsal edge of the quadrate, the posterior margin of the tympanic fascia and then insert onto the retroarticular process by a wide tendon. The more posterior fibers converge anteroventrally to insert lateroventrally onto the superficial fascia of the inter- mandibularis posterior. This muscle is not easily distinguishable nor easily separable into anterior and posterior bundles. It is deep to the constrictor colli and superficial to the geniohyoideus, the sterno- cleidomastoideus, the trapezius, and the most pos- terior edge of the adductor mandibularis externus superficialis. S. This muscle is easily separable into anterior and posterior bundles. The anterior bundle is small and originates from the posterodorsal surface of the squamosal. It terminates in a wide tendon, and then joins with the posterior bundle to insert onto the retroarticular process. It is deep to the posterior bundle and superficial to the sternocleidomastoideus. The posterior fibers of the posterior bundle originate and insert similarly to gilberti. 14 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN M. External Sternocoracoideus. (Not figured) G. The external sternocoracoideus originates from the anterolateral border of the sternum with fibers extending anteriorly to fuse with the internal sterno- coracoideus for a tendonous insertion onto the scapula, medial to the origin of the subscapularis muscles. M. Genioglossus. (Figs. 10 and 15) G. The genioglossus originates from the antero- medial one-fifth of the mandible and inserts poster- iorly by medial and lateral slips onto the hyoglossus and tongue fascia. It is strongly attached dorsally to the oral membrane. There is a small slip of this muscle which attaches anteromedially to the skin fascia. The muscle is lateral to the entoglossal process and the trachea, and medial to the mandibulo- hyoideus 1. M. Geniohyoideus. (Fig. 15) G. The origin of the geniohyoideus is along the anterior margin of the ceratobranchial 1 and the posteroventral one-third of the entoglossal process of the hyoid apparatus. The muscle fibers continue anterolaterally, interdigitate with six or seven slips from the intermandibularis anterior, and insert along the medial surfaces of the dentary and angular. It is deep to the intermandibularis anterior, the constric- tor colli, and the depressor mandibularis. The body of the muscle is superficial to the pterygomandibularis, the mandibulohyoideus 1, the hyoglossus, the genio- glossus, and the trachea. S. The only difference noted is that this muscle is interdigitated with five to seven slips from the inter- mandibularis anterior. (See M. Intermandibularis Anterior) M. Hyoglossus. (Fig. 16) G. The hyoglossus is a thin sheet muscle that origi- nates from the anterior side of the ceratobranchial 1 and inserts anteriorly with the tongue fascia and with interdigitation by medial and lateral slips of the genioglossus. This muscle is deep to the genio- hyoideus and superficial to the oral membrane, the mandibulohyoideus 1 , and portions of the hyoid apparatus, the branchiohyoideus, and the pterygio- mandibularis. M. Intercostalis Externis. (Fig. 14) G. This muscle originates from the posterolateral edge of the ribs and inserts onto the anterolateral edge of the following posterior rib. This muscle is deep to the obliquus abdominus profundus and to the sacrolumbalis and superficial to the intercostalis internis. It does not cover the intercostalis internis ventrally and the muscle become thicker dorsally. Dorsally and anteriorly, fibers of this muscle join with those of the sacrolumbalis and obliquus abdo- minus profundus to insert onto the ventrolateral sur- face of the axis. M. Intercostalis Internis. (Figs. 14 and 17) G. The fibers of this muscle originate from the ribs and insert onto the following posterior rib. On the ventral surface the muscle is nearly parallel to the bilateral axis of the body, and as it extends dorsally the fibers change to a vertical and slightly posterior direction. It is deep to the intercostalis externis and superficial to the obliquus abdominus internis. M. Intermandibularis Anterior. ( Fig. 1 5 ) G. This ventral muscle originates from the medial surface of the mandible and inserts with the opposite muscle in the median fascia. Posteriorly the muscle becomes thinner and is continuous with the mandib- ularis posterior. This muscle is divided into six or seven medially fused slips which are irregular in size and which interdigitate with the geniohyoideus. In the specimens studied the number of slips was approximately equally divided between six and seven. S. This muscle is subdivided into five to seven medially fus,ed slips which are irregular in size and which interdigitate with the geniohyoideus. A sub- specific difference is observed. The number of slips in skiltonianus ranges from five to six; whereas, in utah- ensis the number of slips ranges from six to seven. M. Intermandibularis Posterior. (Figs. 9 and 15) G. This thin sheet muscle originates ventral to the adductor mandibularis externis superficialis by fascia from the ventrolateral surface of the angular and the prearticular, and also from the fascia covering the quadrate. Insertion is ventrally with its partner in the median fascia. The muscle begins just posterior to the last slip of the intermandibularis anterior and term- inates slightly posterior to the tympanic cavity. It is deep to the depressor mandibularis posteriorly. This muscle is superficial to the anterolateral portion of the pterygomandibularis. M. Internal Sternocoracoideus. (Not figured) G. This thin rectangular muscle arises from the posterolaterodorsal surface of the sternum and unites anteriorly with the external sternocoracoideus for a common tendonous insertion onto the scapula, medial to the origin of the subscapularis muscles. M. Latissimus Dorsi. (Figs. 6, 7, 9, and 15) G. The fibers of the latissimus dorsi originate from the ligamentum nuchae, converge ventrally, and pass most medially between the anconaeus coracoideus and anconaeus scapularis to insert tendonously onto the shaft of the humerus. These muscle fibers are deep to the skin and are superficial to the posterior portion of the scapulodeltoideus. M. Levator Plerygoideus. (Fig. 12) BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS 15 G. This muscle is deep to the pseudotemporaiis profundus. It originates from the lateral surfaces of the prootic and parietal descending processes. Fibers extend ventrally to insert onto the lateral and medial surfaces of the pterygoid, just posterior to the fossa columella. M. Levator Scapulae Profundus. (Figs. 12 and 16) G. This narrow somewhat triangular muscle origi- nates in common with the levator scapulae superfi- cialis from the atlas and inserts posteriorly onto the clavicle bar, ventral to the levator scapulae superfi- cialis. It is superficial to the esophageal membrane and deep to the trapezius. M. Levator Scapulae Superficialis. (Figs. 7 and 12). G. This muscle arises from the atlas with the leva- tor scapulae profundus and inserts onto the antero- lateral surface of the suprascapula. This muscle is thick anteriorly but becomes thinner and fan-shaped posteriorly. It is deep to the lateral trunk muscles and is lateral to the rectus capitis anterior. M. Longissimus Cervicis. (Figs. 8, 12, 13, and 17) G. This muscle is located ventral to the obliquus capitis magnus. Its origin is by fusion with the myo- septa between the obHquus capitis magnus and the lateral bundle of the spinalis capitis. The fibers extend in a ventroanterior direction to insert tendon- ously onto the basioccipital process. M. Mandibulohyoideus l.(Fig. 16) G. The origin of this muscle is from the ventro- medial surface of the dentary and the angular, and inserts onto the most anteroventral surface of the distal segment of the hyoid cornu. One to two slips of the intermandibularis anterior interdigitate with this muscle. It is superficial to the oral membrane and the medial portion of the pterygomandibularis, and lies lateral to the geniohyoideus. M. Obliquus Abdominus Externis. (Figs. 6, 9, and 15) G. This sheet muscle originates anterolateral^ from the ribs of the sixth and seventh vertebrae and laterally from the sacrolumbalis. Insertion is with the pectoralis and rectus abdominus ventrolaterally. M. Obliquus Abdominus Internis. (Fig. 14) G. This thin, loosely arranged sheet muscle origi- nates from the inner surface of the ribs and inserts anteriorly onto the esophageal membrane, the ster- num, and the ventral heads of the thoracicolumbar ribs. The fibers are superficial to the transversalis and deep to the intercostalis internis. They extend in a dorsal and slightly posterior direction. M. Obliquus Abdominus Profundus. (Figs. 13 and 17) G. This muscle has the same origin as the externis muscle except that some fibers will continue antero- dorsally and become almost parallel with the sacro- lumbalis to interdigitate with that muscle and the intercostalis externis to insert onto the axis. Insertion is also by individual slips onto the sternal ribs with the most lateral fibers being attached to the most posterior rib. This muscle is generally heavier than the externis muscle. M. Obliquus Capitis Magnus. (Figs. 8, 12, and 13) G. This muscle originates from a myosepta with the lateral slip of the spinalis capitis. The myosepta extends lateroventrally from the fifth vertebrae to the atlas. Insertion is onto the supratemporal, the paroc- cipital process, and the parietal. M. Pectoralis. (Figs. 1 5 and 16) G. The pectoralis is a thick muscle having its origin from the lateral process of the interclavicle, the sternum, the mesosternum and the ribs which are attached to the mesosternum, and the first two thora- cicolumbar ribs. Its origin along the ribs is in a steplike fashion laterally, with the most lateral slip fusing with the rectus abdominus. All fibers converge to insert tendonously onto the deltopectoral process of the humerus. This muscle is deep to the skin and to a lateral slip of the rectus abdominus. It overlies a broad ligament tVom the interclavicle lateral process to the sternum and the posterior portion of the scapula with its attached muscles. M. Protractor Pterygoideus. (Fig. 12) G. The anterior portion of this muscle is deep to the pseudotemporaiis profundus. Origin is from the anterolateral surface of the prootic. Fibers extend lateroventrally and posteriorly to insert onto the pterygoid and the quadrate process of the pterygoid. M. Pseudotemporaiis Profundus. (Fig. 1 1) G. This muscle is deep to the adductor mandib- ularis externis profundus and the adductor mandib- ularis externis medius. It originates from the descend- ing processes of the prootic and the parietal; and also from the lateral, posterior and anterior surfaces of the epipterygoid. Some of the dorsal fibers originate from the ventral surface of the parietal. The muscle con- tinues in a nearly anteroventral direction to insert onto the base of the coronoid and posteriorly onto the surangular and the mandibular fossa. M. Pseudotemporaiis Superficialis. (Fig. 1 1) G. This muscle is located medial to the adductor mandibularis externis medius and dorsolateral to the adductor mandibularis externis profundus. Origin is from the ventral surface of the parietal medial to the supratemporal fontanelle. Fibers extend ventrally to insert tendonously onto the dorsal surface of the coronoid. 16 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN M. Pterygomandibularis. (Figs. 7, 10, 13, 15 and 17) G. The pterygomandibularis originates tendon- ously from the pterygoid and the epipterygoid. The muscle continues posteriorly along the pterygoid, attaches to the pterygoid, the basipterygoid process, and then covers the ventrolateral surfaces of the angular, the articular, the medial surface of the surangular, and the medial dorsal surface of the retro- articular process. The muscle then extends dorsally around the posterior end of the retroarticular process to insert onto the condyle of the quadrate and the angular process of the articular. It is deep to all ven- tral and lateral muscles and is covered by an extensive fascia. M. Rectus Capitis Anterior. (Figs. 14 and 17) G. This is a thick straplike muscle originating by two slips from the ventral surface of the eighth and seventh vertebrae and their rib capitulae. Anteriorly the two slips fuse. The combined muscle interdigi- tates with its opposite partner and secondarily arises from the ventral spinous processes of the remaining anterior vertebrae. Insertion is medial to the longis- simus cervicis onto the basioccipital and its process. The body of the muscle lies dorsal to the esophagus and ventral to the longissimus cervicis. M. Rectus Capitis Posterior. (Figs. 8, 13, and 17) G. Anteriorly this muscle is deep to the spinalis capitis and medial to the obliquus capitis magnus. In the vicinity of the seventh vertebrae the muscle is deep only to the dorsal fascia, is medial to the lateral portion of the spinalis capitis, and lateral to the ligamentum nuchae. It is separated into anterior and posterior bundles by a myosepta with some interdigi- tation in the vicinity of the fourth vertebrae. The anterior bundle is divided into a lateral and a medial slip which are interdigitated but separable. The lateral slip makes the myosepta connection with the poster- ior bundle and originates from the second, third, and fourth vertebrae with insertion onto the paroccipital process, the parietal, and the supratemporal. The medial slip originates from the atlas and inserts onto the prootic and the supraoccipital elements. The most anterior portion of the posterior bundle lies deep to the anterior bundle and inserts tendonously onto the axis. M. Sacrolumbalis. (Figs. 6 and 9) G. This superficial epaxial muscle originates from the crest of the ilium and inserts tendonously onto the ribs. Anteriorly, fibers of this muscle will join those of the intercostalis externis and the obliquus abdominus profundus to insert onto the ventrolateral process of the axis. This muscle interdigitates laterally with the obliquus abdominus muscles. This muscle is superficial to the intercostalis externis dorsally. M. Scapulodeltoideus. (Figs. 6, 9, 1 2, and 1 3) G. This triangular shaped muscle originates primar- ily from the lateral surface of the suprascapula with some fibers originating from the clavicle bar. Inser- tion is tendonously onto the deltopectoral process of the humerus. The muscle is ventral to the levator scapulae muscles and superficial to the scapulo- humeralis muscles. M. Scapulohumeralis Anterior. (Figs. 14 and 17) G. This muscle has two slips. The dorsal slip inserts onto the anterior edge of the humerus next to the glenoid fossa beneath the sternoscapular ligament. The fibers continue anterodorsally to originate from the most dorsoanterior surface of the scapular bar and the most anteroventral surface of the supra- scapula. The second slip, which is the longer, has the same origin but inserts onto the ventrolateral border of the scapula. Centrally the two slips are joined by a myosepta. M. Scapulohumeralis Posterior. (Figs. 8, 14, and 17) G. This rectangular shaped muscle inserts onto the proximal end of the humerus next to the glenoid fossa, posterior to the insertion of the scapulo- humeralis anterior and medial to the insertion of the latissimus dorsi. The fibers continue anterodorsally covering some of the fibers of the serratus ventralis to originate from the posterolateral surface of the supra- scapula and the scapular bar. M. Serratus Dorsalis. (Fig. 8) G. This muscle arises from the lateral surface of the first three cervical ribs by three slips and inserts onto the dorsal surface of the suprascapula. They sliglitly overlap one another by the posterior suc- ceeding one. M. Serratus Ventralis. (Figs. 8 and 13) G. This is a wide straplike muscle which originates from the ribs of the sixth and seventh vertebrae, dorsal to the origin of the obliquus abdominus muscles. Insertion is onto the posteromedial border of the suprascapula and the dorsolateral portion of the scapular bar. This muscle is deep to the latissimus dorsi and the trapezius. M.SpinalisCapitis.(Figs. 7, 8, 12, and 13) G. This muscle originates along the cervical and thoracicolumbar vertebrae and inserts along the entire posterior portion of the parietal. Anteromedially, fibers insert tendonously onto the side of the supra- occipital ridge. The muscle has a medial and a lateral portion which are unseparable. The medial portion is superficial to the rectus capitis posterior and thins posteriorly to terminate near the seventh vertebrae. The lateral portion participates in a myosepta with some of the deeper muscles and continues poster- BIOLOGICAL SERIES. VOL. 12, NO. OSTEOLOGY AND MYOLOGY OF THE SKINKS iorly, lateral to the rectus capitis posterior. M. Subscapularis 1 . (Not figured) G. Tiiis muscle has three slips which converge posteriorly to insert by a common tendon onto the humeral head. The most anterior slip originates from the anterodorsal border of the scapula. The most lateral slip of the two posterior slips originates from the scapular bar and the suprascapula; whereas, the medial slip originates from the scapular bar. They both converge posteriorly to join the anterior slip. M. Subscapularis 2. (Not figured) G. The fibers of this muscle originate from the majority of the dorsal surface of the scapula and converge to insert onto the humeral head dorsal to the insertion of the subscapularis 1. This muscle is located between the anterior slip of the subscapularis I and the dorsal slip of the coracobrachialis longus, and lies lateral to the origin of the internal sterno- coracoideus. M. Supracoracojdeus. (Fig. 17) G. This ventral muscle has a wide origin from the anteromedial surface of the scapula with the fibers converging to insert tendonously onto the medial surface of the del topect oral crest of the humerus. This muscle is located posterior to the clavodel- toideus and anterior to the biceps. On the scapula a thin layer of fibers will cover the anteroventral surface of the biceps. It is thicker anteriorly. M. Steriiocleidomastoideus. (Figs. 6, 9. 11, and 15) G. This wide, thick ribbonlike muscle originates from the posterior surfaces of the parietal, the squamosal, and the quadrate. It extends ventropos- teriorly to insert onto the lateral process of the interclavicle. Near its insertion this muscle is fused with the trapezius. It is deep to the depressor mandib- ularis, the ceratobranchial 1, the branchiohyoideus, and the lateral portion of the trapezius. It is super- ficial to the neck muscles and to the insertion of the trapezius and the sternohyoideus. M. Sternohyoideus. (Figs. 10, 12, 15, and 16) G. The sternohyoideus is a thick sheet muscle having a superficial and a profundus layer. Some of the fibers slightly intermix medially and tend to become parallel to the bilateral axis; however, the muscle is easily separable into two layers. The ventral layer is sliglitly larger and originates from the pos- terior and ventral surfaces of the ceratobranchial 1 and medially from the corpus. Its fibers extend posterolaterally to insert onto the lateral process of the interclavicle and the clavicle-suprascapular bar, adjacent to the attachment of the trapezius. At its insertion this muscle joins with the sternocleido- mastoideus and the trapezius. It is deep to portions of the sternocleidomastoideus, trapezius, depressor mandibularis, and the constrictor colli. The dorsal layer extends further laterally than the superficial layer and has its origin from the posterior surface of the ceratobranchial 1 and 2. Fibers continue postero- medially to insert onto the lateral process of the interclavicle. This layer is superficial to the pharyn- geal membrane, the thyroid gland, and portions of the clavodeltoideus and the scapulodeltoideus. Medially, this muscle borders an elliptical space as previously mentioned in the section, General Descrip- tion of the Myology. M. Transversalis. (Fig. 14) G. This loosely arranged muscle arises from a fascia which extends from the pelvis to the neck region and inserts ventrally onto the ribs, the lateral surface of the sternum, and the dorsal surface of the rectus abdominus. The fibers extend in a dorsoanter- ior direction, are superficial to the peritoneal mem- brane and are deep to the obliquus abdominus in- ternis. M. Trapezius. (Figs. 6, 7, 9, II, 15, and 16) G. The trapezius originates from the middorsal fascia and covers a majority of the lateral muscula- ture. It narrows laterally to insert ventrally onto the clavicle bar and the lateral process of the interclavicle which in turn is attached to the sternum by a broad ligament. On the ventral surface the muscle fuses with the sternocleidomastoideus. In some specimens this muscle is attached to the pectoralis by a tendonous fascia. The trapezius is variable in thickness with the anterior fibers usually being thicker. Toward the center they become thinner and in some specimens can be separated into two bundles. In this case the anterior bundle inserts onto the lateral process of the interclavicle and the most ventral portion of the inter- clavicle-clavicle bar. The posterior bundle inserts onto the posterior border of the clavicle bar, dorsal to the sternocleidomastoideus. Unnamed Muscle. (Not figured) The following muscle was found in E. gilberti (BYU 31956). It is a thin horizontal straplike muscle originating by fascia from the ribs of the twelfth and thirteenth vertebrae and inserting onto the anterior edge of the insertion of the serratus ventralis, and also onto the suprascapula. This muscle is deep to the latissimus dorsi. It was not present in any other speci- men o^ gilberti ox skiltonianus studied. BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN ADD MAND EXT SUPER CONST COLLI TRAPEZIUS ANC SCAP ANC CORAC LAT DORSI AND EXT MED RESSOR MAND NOCLEIDOMAST CL A VODELTOID APULODELTOID NC HUM MED COR BRA LON OBL ABD EXT SACROIUMB , s A. Superficial depth B. I-irst depth Figure 6. MusLiilalure dorsal view. Eumcces gilhcrli (3.7X) (I3YU 31956). BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS 19 PTERYGOMAND TRAPEZIUS LAT DORSI SPINALIS CAPITIS LEV SCAP SUP A. Second depth B. Third depth Figure 7. Musculature dorsal view. Eumeces gilherti (3.7X) (B^■U 31956). 20 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN j ADD MAND EXT MED OBL CAP MAGNUS SERRATUS OORSALIS SERRATUS VENT SPINALIS CAPITIS RECTUS CAPITIS POST LONG CERVICIS SCAPULOHUM POST RECTUS CAPITIS POST ] A. Fourth dcplli B. fifth depth I'lgure 8. Musculature dorsal view. i'»mfccsi;(//)m/( 3.7 X) (BYU 31956). BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS 21 INTERMAND POST STERNOCLEIDOMAST SCAPULODELTOID CLAVODELTOID RECTUS ABD ADD MANO EXT SUPER CONST COLLI DEPRESSOR MAND TRAPEZIUS LAT DORSI OBL ABD EXT SACROLUMB l-igure 9. Musculature lateral view. Superficial depth. Eumeces gilherti (3.7X) (BYU 31956). 22 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN GENIOGLOSSUS- PTERYGOMAND- STERNOHYOID DEPRESSOR MAND- POST ANT- B ADD MAND EXT MED ADD MAND EXT PRO ADD MAND POST ligure 10. Musculature lateral view. Mrs! depth. A. Eiimeces gi/herli (3.7X)(BYU 31956). H. t'timeces skilloiiiainis (5 JX) i\iYV 12474), Showing anterior and posterior bundles ot' the depressor mandibularis. BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS 23 STERNOCLEIDOMAST PSEUDOTEMP SUP PSEUDOTEMP PRO •TRAPEZIUS F'igure 1 1. Musculature lateral view. Second depth. Eumeces gilberti (3.7X) (BVU 31956). 24 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN STERNOHYOID PRO LEV SCAP PRO CLAVODEITOID LEV PTERYGOID PROT PTERYGOID SPINALIS CAPITIS OBL CAP MAGNUS LONG CERVICIS LEV SCAP SUP SCAPULODELTOID Figure' 1 2. Musculature lateral depth. Third depth. Eumeces gilberri (3.7X) (BYU 31956). BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS 25 PTERYGOMAND ClAVODELTOID SCAPULODELTOID SERRATUS VENT OBI ABD PRO RECTUS CAPITIS POST OBL CAP MAGNUS LONG CERVICIS SPINALIS CAPITIS I Figure 13. Musculature lateral depth. Fourth depth. Eumeces gilberti (3.7X) (BYU 31956). 26 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN SCAPULOHUM ANT SCAPULOHUM POST INTERCOST iNT INTERCOST EXT RECTUS CAPITIS ANT TRANS OBL ABO INf I'iguri; 14. Musculature lateral view. Iifth depth. Eumeces gilherli (3.7X) (BYU 31956). BIOLOGICAL SERIES. VOL. 12. NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS 27 GENIOGLOSSUS INTERMAND ANT ADD MAND EXT SUPER INTERMAND POST- CONST COLLI SPACE FASCIA- ANC SCAP- BICEPS LAT DORSI- OBL ABD EXT- •GENIOHYOID PTERYGOMAND •DEPRESSOR MAND •STERNOHYOID -STERNOCLEIDOM AST TRAPEZIUS CLAVODEL TOID — ANC HUM LAT -COR BRA LON RECTUS ABD -PECTORALIS A. Superficial depth B. F-irst depth I'igure 15. Musculature ventral view. Humeces gilberti (3.7X) (BYU 31956). 28 BRIGHAM YOUNG UNIVERSITY BULLETIN HYOGIOSSUS STERNOHYOID SUPER TRAPEZIUS MANDIBULOHYOID I STERNOHYOID PRO LEV SCAP PRO CLAVODELTOID PECTORALIS A. Second deplli B. Third depth ligiire 16. Musculature ventral view. Eumeces gilherti iiJX) IBYLI 31956). BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY .\ND MYOLOGY OF THE SKINKS TONGUE FASCIA BRANCHIOHYOID CLAVGDELTOID SUPRACORACOID BICEPS OBL ABD PRO PTERYGOMAND TUS CAPITIS ANT US CAPITIS POST LONG CERVICIS SCAPULOHUM ANT APULOHUM POST COR BRA BRE BRA INF INTERCOST INT A. Fourth depth B. Fifth depth Fiaure 17. Musculature ventral view. Fifwiffes^^/Z/Kv// ( 3. 7X) (BYU 31956). 30 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN DISCUSSION Osteology Upon comparing the lizard skull with other groups of the class Reptilia one tends to agree with Jollie (1960) "that all are specialized to some extent and none is strikingly more primitive in the totality of its structure than the others." He further states that the variation of structures can be explained largely by adaptive modification and that the Reptilia show "radiation rather than an anatomical heirarchy of groups." To determine what the ancestral lizard or the first lizard may have looked like one must rely on the fossil record and the anatomy of living lizards. Broom (1935) is of the opinion that lizards arose from Eosuchian stock in the Upper Triassic and then later developed osteoderms over the skull and the body regions. In the Middle Mesozoic some primitive groups retained the osteoderms while others lost them; thus giving rise to the living forms which have a variety of skull variations. All of the families within the infraorder Leptoglossa (Sincomorpha) have osteoderms except the family Teiidae. The genus Ameiva within this family has os palpabre elements (Fisher and Tanner, 1970) which may be suggestive of cranial osteoderms sometime in the past. In fam- ilies that still retain the osteoderms, Laurtidae and Scincidae, the supratemporal fenestrae are extremely reduced or lost in contrast with the teiids which have large fenestrae. The postfrontals are expanded in the former while in the latter the postfrontals are re- duced, lost, or fused with the postorbitals. Following is a discussion of the osteological comparisons between £". gilberti and E. skiltonianus. The variations noted are; 1 . In gilberti the anterior lateral processes of the frontals prevent articulation of the prefrontals with the nasals; whereas, in skiltonianus articulation with the nasals and prefrontals can either be present or absent. 2. In gilberti the maxillae contain 6 to 8 supra- labial foramina and 15 to 20 teeth, average 19; whereas, in skiltonianus the maxillae contain 4 to 7 supralabial foramina and 17 to 20 teeth, average 18. 3. In gilberti there are 3 to 5 nasal foramina compared with 2 to 4 in skiltonianus. 4. In gilberti the dentary has 3 to 6 mental for- amina and 19 to 23 teeth, average 2 1 ; compared with skiltonianus which has 4 to 7 mental foramina and 18 to 22 teeth, average 21. 5. In gilberti the postorbitals frequently show a reduction of the relative size when compared to other adjacent bones while in skiltonianus the same degree of reduction is only occasionally observed. Condens- ing these variations it is found that: 1 . the number of foramina in the maxillae, nasals, and dentary overlap and are variable within species with neither species showing a definite trend; 2. the same applies to the number of teeth in the maxillae and mandible; 3. in gilberti the nasal-prefrontal articulation is constant; whereas, in skiltonianus it is variable; and 4. in gil- berti the postorbitals frequently show a reduction of the relative size when compared to other adjacent bones, whereas, in skiltonianus this same degree of reduction is only occasionally observed. Myology In contrast with osteology, the homologies of muscles, and thus phylogenies, are difficult to trace when comparing different taxonomic categories. If significant changes are to be found, they are in the cranial and throat myology. These areas seem to be the most easily traceable among groups and in turn are phylogenetically the most important. Brock (1938) working with the genus Gecko found the throat musculature to be stable enough for generic discriminatien. This conclusion has been supported by other papers by Tanner (1952) with Plethoden- tidae: Robison and Tanner (1962) with Crotophytus: Avery and Tanner (1964) with Sauromalus; Jenkins and Tanner (1968) with Phrynosyma: and Fisher and Tanner ( 1970) with Cnemidophorus and Ameiva. Smith (1960) lists a number of trends which have taken place in the musculature of the more primitive to the more advanced vertebrates. These are: I. the loss of myosepta and the fusion of myotomes; 2. the separation of muscles into layers; 3. the expansion of muscles from a limited size; 4. the shifting of location of muscles; 5. the multiplication or fusion of muscle attachments; and 6. the subdivision of muscles. In general these two species show both primitive and advanced characters. The presence of myosepta in the axial musculature would certainly indicate a primitive condition. The subdivision of the interman- dibularis anterior, the rectus capitis posterior, the possible trend toward a complete separation of the trapezius and the depressor mandibularis and the distinct layers of the obliquus abdominus externis and sternohy oideus muscles would indicate an advanced condition. When comparing the myology of these two species little difference is observed. The intermandibularis shows a subspecific difference in /:'. skiltonianus. In two specimens observed of the E. skiltonianus sub- species, a range of five to seven subdivisions of the intermandibularis anterior was found. In E. skilton- ianus the number of subdivisions tends to be toward the lower limit; whereas, in /:'. utahensis the number of subdivisions is toward the higher limit. In contrast both subspecies of E. gilberti tend to orient them- selves equally with six to seven subdivisions Rir this BIOLOGICAL SERIES. VOL. 12, NO. 2 OSTEOLOGY AND MYOLOGY OF THE SKINKS 31 muscle. The only myological character found in this study which differentiates these two species consist- ently is the depressor mandibularis. In E. skiltonianus this muscle is easily separated into an anterior and a posterior bundle; whereas, in E. gilberti this same muscle is somewhat intcrdigitated and not* easily separable. All other muscles mentioned are similar in both species. SUMMARY AND CONCLUSIONS The differences observed from the osteology and myology are few and subtle. Neither species is consistent m having all of the anatomical characters equal. E. skiltonianus, however, is more variable than gilberti. This species: 1. exhibits a wider range of variation in the number of slips of the intermandibularis anterior; 2. has two distinct bundles of the depressor mandibularis; and 3. has variation in the anterior suture pattern of the frcmtal, nasal, prefrontal and maxilla elements. E. gilberti on the other hand has: 1. only one distinct bundle of the depressor mandibularis; 2. a frequent reduction of the relative size of the postorbitals when compared to other adjacent bones; and 3. a more limited suture pattern in the anterior portion of the skull. The major anatomical difference between these two species seems to be size. From the specimens used in this study the average snout-vent length of gilberti is approximately 30mm greater than that of skilton- ianus. We believe that skiltonianus shows more anatom- ical plasticity than gilberti. Plasticity in this case would indicate a more recent gene flow between peripheral and central individuals, and genetic incon- stancy which would provide more variation and more flexibility for adaptation and selection. When the geographical distribution of these two species is considered it also appears that skiltonianus has more flexibility. Taylor ( I')35) has suggested that these species may have arisen simultaneously from a common ancestral stock. On the basis of this anatomical study there is little reason to doubt their close relationship. How- ever, it seems logical to propose an alternate sugges- tion; namely, that the anatomical variations of £". skiltonianus. when compared with similar characters in E. gilberti, indicate that E. gilberti may have arisen from E. skiltonianus. ACKNOWLEDGEMENTS We wish to express our appreciation to Drs. Herbert H. Frost, Ferron L. Andersen and H. Duane Smith of the Department of Zoology, Dr. Morris S. Petersen of the Department of Geology and David F. Avery for their criticisms and review of the manu- script. Gratitude is extended to Dr. R. L. Livezey of the Sacramento State College and to Dr. John Wright of the Los Angeles County Museum for providing speci- mens which were used in the preparation of this thesis. Other specimens were obtained from the Ver- tebrate Natural History Museum collections of Brigham Young University. Financial assistance was provided by the Depart- ments of Zoology and Geology. We also wish to extend our thanks to others who have assisted in the preparation of specimens, field work, photography, and illustrations. These include Gary L. Burkholder. Nathan M. Smith, and Kenneth R. Larsen of the Department of Zoology; Samuel R. Rushforth and Dr. William D. Tidwell of the Depart- ment of Botany; and William L. Chesser of the Department of Geology. 32 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN LITERATURE CITED Avery, D. F. and W. W. Tanner. 1964. The osteology and myology of the head and thorax regions of the ohesiis group of the genus sauromalus Dumeril (Iguanidae). Brigham Young University Seienee Bulletin, Biological Series. 5(3): 1-30. Baird, S. F. and C. Girard. 1852. Characteristics of some new reptiles in the museum of the Smithsonian Institution. Proceedings of the Academy of Natural Science, Phila- delphia. 1852:68-70. Brock, G. T. 1938. The cranial muscles of the Gecko, a general account, with a comparison of the muscles in other Gnathostomes. Proceedings of the Zoological Society of London. Series B, 108:735-761. Broom, R. 1935. On the structure of the temporal region in lizard skulls. Annals of the Transvaal Museum. 18:13-22. Camp, Charles Lewis. 1916. Notes on the local distribution and habits of the amphibians and reptiles of southeastern California in the vicinity of the Turtle Mountains. Univ. Calif. Pub. Zool.. 12( l7):502-544, pis. 19-22. Edgeworth. F. H. 1935. The cranial muscles of vertebrates. Cambridge University Press, London. Fisher, D. L. and W. W. Tanner. 1970. Osteological and myological comparisons of the head and thorax regions of Cnemidophonis tigris scptcntrionalis Burger and Ameiva tindulala parva Barbour and Noble (Family Teiidae). Brigham Young Liniversity Science Bulletin. Biological Series. I 1(1): 1-41. Jenkins, R. L. and W. VV. Tanner. 1968. Osteology and myology of Phrynosoma r. platrrhiitos Girard and PInynosoma d. heniandesi (iirard. Brigham Young University Science Bulletin. Biological Series, 9(4): 1-34. Jollie, M. T. I960. The head skeleton of the lizard. Acta Zoologica, 41: 1-64. Kingman, R. H. 1932. A comparative study of the skull in the genus t'umeces of the Scincidac (a preliminary pa- per). The University of Kansas Science Bulletin. 20(15): 273-295. Oelrich, T. M. 1956. The anatomy of the head of Ctenosaura pectinata (Iguanidae). Miscellaneous Publications of the Museum of Zoology, University of Michigan. (94): I-1 18. Robison, G. W. and W. W. Tanner. 1962. A comparative study of the species of the genus Crotophyliis Holbrook (Iguanidae). Brigham Young Liniversity Science Bulletin, Biological Series, 2(1): 1-31. Rodgers, T. L. and H. S. Fitch. 1947. Variation in the skinks (Reptilia: Lacertilia) of the skiltoiiianiis group. Univer- sity of California Publications in Zoology, 48:169-209. Romer, A. S. 1956. Osteology of the Reptiles. University of Chicago Press, Chicago. Smith, H. M. 1946. Handbook of lizards. Comstock Publish- ing Company, Ithaca, Nev\ York. 1960. Fvolution of Chordate Structure. Holt, Rinehart and Winston, New York. Tanner, W. W. 195 2. A comparative study of the throat musculature in the Plethodontidae of Mexico and Cen- tral America. University of Kansas Science Bulletin, 34:583-677. 1957. A taxonomic and ecological study of the west- ern skink it'Dim'ces skillotiiamis). The Great Basin Naturalist. 17( .3-4):59-94. Taylor. [■'. H. 1935. A taxonomic study of the cosmopolitan scincoid lizards of the genus t'tinieces with an account of the distribution and relationships of its species. The University of Kansas Science Bulletin, 36( 14): 1-643. Van Denbufgh, J. 1896. Description of a new lizard ih'u- meces gilbcni) from the Sierra Nevada of California. Proceedings of the California Academy of Science, 2(6):350-352. Weigmann A. f. 1834. Herpetologia Mexicana 1834. pp i-vi 1-54. pIsI-X. INFORMATION FOR CONTRIBUTORS Contributions to the Science Bulletin should be pri- marily monographic in nature. For the most part only manuscripts of approximately forty or more typewritten pages will be accepted. Papers will be published approximately in the order that they are received, pending availability of funds. Authors must arrange for financing their publications, except that the University Press and the library of Brigliam Young University will share the publication cost proportionately to their needs for copies of the article printed. In the preparation of manuscripts, authors are re- quested to follow the Style Manual for Biological Jour- nals, American Institute of Biological Sciences, 2000 P Street, N.W., Washington, D.C., 20006. Manuscripts must be typed on one side of the paper only, double- spaced with ample margins. Footnotes should be avoided. To facilitate review by referees, send to the editor the original manuscript and one carbon copy, together with the illustrations. Copies of the original illustrations may be submitted, but should be of quality equal to the orig- inals. Illustrations should be referred to as figures except for materials requiring inserts of special paper, which may be called plates. Illustrations should be so designed as to fit when reduced into a one-column or full-page width. Special care must be taken to allow for proper reduction in lettering (i.e., a 50% reduction of the figure means also a 50% reduction in the lettering.) Photo- graphs should be of a glossy finish, unblurred, and show- ing sharp contrast. Line drawings should be made with black ink on heavy white drawing paper, blue tracing cloth, or blue-ruled coordinate paper. Use the same abbreviations on line drawings as in text. Line drawings must be equivalent to a professional draftsman's work. Original drawings are preferable to photographs, even if they are large. Illustrations (line drawings or photo- graphs) should be numbered consecutively throughout the paper, and the approximate place of insertion should be indicated in the margins of the manuscript pages. Captions for illustrations should be assembled on a sep- arate sheet, and each plate of figures must have its corre- sponding figure number pencilled lightly on the back. Illustrations and cuts will be destroyed unless their re- turn is requested when proof is returned to the editor. A table title should be a short, concise statement of what the table purports to show, and should not include information necessary to the interpretation of the table. Every column in the table should carry a head identify- ing the data in that column; the measure in which the data are given should be indicated at the head of each column. Tables should not be used when the same infor- mation can be given in a few lines of text, and should not duplicate information in text, graphs, or charts. Symbols (asterisk, dagger, etc.) should be used to indi- cate footnotes to tables, with footnotes on the same page as the table. An abstract of less than four percent of the length of the paper should be prepared. This summary should be understandable without reference to the body of the manuscript. The abstract must be on sheets separate from the manuscript. Proof should be corrected immediately on receipt and returned to the editor. Authors should leave forwarding addresses if they move from the address sent with the manuscript. Reprints should be ordered when the proof is re- turned. Address all manuscripts to Stanley L. Welsh, Depart- ment of Botany, Brigham Young University, Provo, Utah 84601. i i S-^' Brigham Young University f?!AR12j97d Science Bulletin \ EVOLUTION OF THE IGUANINE LIZARDS (SAURIA, IGUANIDAE) AS DETERMINED BY OSTEOLOGICAL AND MYOLOGICAL CHARACTERS by David F. Avery and Wilmer W. Tanner BIOLOGICAL SERIES — VOLUME Xil, NUMBER 3 JANUARY 1971 Brigham Young University Science Bulletin EVOLUTION OF THE IGUANINE LIZARDS (SAURIA, IGUANIDAE) AS DETERMINED BY OSTEOLOGICAL AND MYOLOGICAL CHARACTERS by David F. Avery and Wilmer W. Tanner BIOLOGICAL SERIES — VOLUME XII, NUMBER 3 JANUARY 1971 TABLE OF CONTENTS Page LIST OF TABLES LIST OF ILLUSTRATIONS INTRODUCTION 1 LITERATURE 1 MATERIALS AND METHODS 8 OSTEOLOGY 9 Skull and Jaws 9 Teeth 22 Hyoid Elements 23 Sterna and Ribs 23 MYOLOGY 34 Throat Musculature 34 Neck Musculature 36 Temporal Musculature 38 OTHER CHARACTERS 40 Tongues 40 Hemipenes 67 DISCUSSION 67 Osteology 67 Myology 69 Tongues 70 Hemipenes 70 Iguanine Distribution 70 ACKNOWLEDGMENTS 73 CONCLUSIONS AND SUMMARY 73 LITERATURE CITED 75 LIST OF TABLES Table Page 1 . Skull Length and Width 9 2. Skull Length and Heiglit 9 3. Basisphenoid Bones 10 4. Basioccipital Bones 10 5. Exoccipital Bones 11 6. Supraoccipital Bones 11 7. Pterygoid Bones 12 8. Ectopterygoid Bones j 2 9. Vomer Bones j 2 10. Palatine Bones 13 1 1 . Premaxillary Bones 13 1 2. Maxillary Bones ] 4 13. Nasal Bones I4 14. Prefrontal Bones 14 1 5 . Lacrimal Bones 15 16. Frontal Bones 15 17. Postfrontal Bones I5 1 8. Jugal Bones 16 1 9. Parietal Bones 17 20. Parietal Wings 17 2 1 . Postorbital Bones 17 22. Squamosal Bones 18 23. Quadrate Bones 18 24. Supratemporal Fossa 19 25. Orbits I9 26. Fenestra Exonarina I9 27. Dentary Bones 20 28. Articular Bones 20 29. Angular Process 20 30. Surangular Bones . . , 21 31. Splenial Bones 21 32. Angular Bones 21 33. Coronoid Bones 22 34. Teeth 23 35. Summary of Important Myological Differences 39 36. Tongue Measurements 40 37. The Number of Osteological Similarities between Genera 57 LIST OF ILLUSTRATIONS Figure Page 1 . Ventral view of skull 24 2. Ventral view of skull 25 3. Dorsal view of skull 26 4. Dorsal view of skull 27 5. Lateral view of skull 28 6. Lateral view of skull 29 7. Medial view of mandibles 30 8. Ventral view of hyoid bones 31 9. Ventral view of sternum 32 10. Ventral view of sternum 33 11. Ventral view of throat musculature; superficial layer shown at left and first depth at right _^ I 12. Ventral view of throat musculature; superficial layer shown at left and first depth at right 42 13. Ventral view of throat musculature; second depth at left and third depth at right 43 14. Ventral view of throat musculature; second depth at left and third depth at right 44 15. Ventral view of throat musculature; fourth depth at left and fifth depth at right . 45 16. Ventral view of throat musculature; fourth depth at left and fifth depth at right . 46 17. Dorsal view of throat and neck musculature; superficial depth at left and first depth at right 47 18. Dorsal view of throat and neck musculature; superficial depth at left and first depth at right 4g 19. Dorsal view of head and neck musculature; second depth at left and third depth at right 49 20. Dorsal view of head and neck musculature; second depth at left and thiid depth yl riglit 50 21. Dorsal view of head and neck musculature; fourth depth at left and fifth depth at right 5] 22. Dorsal view of head and neck musculature; fourth depth at left and fifth depth at right 52 23. Lateral view of head and neck musculature; superficial depth 5^ 24. Lateral view of head and neck musculature; superficial depth 54 25. Lateral view of the head and neck musculature; first depth 55 26. Lateral view of the head and neck musculature; first depth 55 27. Lateral view of head and neck musculature; second depth 57 28. Lateral view of head and neck musculature; second depth 58 29. Lateral view of head and neck musculature; third depth 59 30. Lateral view of head and neck musculature; third depth (,0 31 . Lateral view of head and neck musculature; tourth depth 61 32. Lateral view of head and neck musculature; fourth depth 62 33. Lateral view of head and neck musculature; fifth depth 53 34. Lateral view of head and neck musculature; fit~th depth 64 35. Dorsal view of the tongue 65 36. Hemipenes 66 37. Phylogenetic relationships of the Madagascar Iguanidae and the genera of iguanine lizards 71 Ctenosiiura l^ecliihil.i (Wiegmann) taken 50 miles S.W. of Guadalajara (Hwy. 80) by Kenneth R. Larsen, 18 July 1970. EVOLUTION OF THE IGUANINE LIZARDS (SAURIA, IGUANIDAE) AS DETERMINED BY OSTEOLOGICAL AND MYOLOGICAL CHARACTERS by David F. Avery and Wilmer W. Tanner INTRODUCTION The family Iguanidae is almost completely re- stricted to the Western Hemisphere with its main radiations occurring in North and South America. There are also representatives on Fiji, Tonga, and the Galapagos Islands in the Pacific Ocean. Two distinctly related iguanid genera are also found on Madagascar. These genera, Chalarodon and Opiums . possess abdominal ribs and are therefore considered to be the most primitive members of the family. Although the iguanid lizards are familiar to most scientists inter- ested in the tropics, their anatomy and evolution are still poorly understood. Because the family Iguanidae is a large and diverse group of lizards, several distinct phylogenetic lines have been recognized. In this study we are concerned with that group of genera belonging to the iguanine line, which includes the following genera: Ambly- rhynchm and Conolophus from the Galapagos Is- lands, Brachylophits from Fiji and Tonga Islands, Enyaliosaurus from Central America, Ctenosaura and Iguana from Central and South America, Cyclura from the West Indies, and Dipsosaunis and Sauro- malus from North America. Those iguanid lizards which have a discontinuous distribution all belong to the iguanine line, or are the most primitive members of the family. Explaining the discontinuous distribution pattern between the West- ern Hemisphere mainland iguanines, the Pacific Island forms, and their Madagascar relatives has proven to be an enigma for zoogeographers and herpetologists. The purpose of this study is to establish the degree of relationship between the iguanines of the Galapa- gos, Fiji, and Tonga Islands with the mainland genera. We will also attempt to define more completely the relationships between the Madagascar genera and the iguanine line. In order to ascertain these relationships, the anterior osteology and myology of each genus has been investigated along with such specialized features as the tongue, hyoid bones, sterna and hemipenes. Hopefully the morphological relationships between the ten genera can be clarified by the use of these relationships, and the evolution and distribution of the iguanine iguanids can be explained. Of all the genera listed above, only Enyaliosaurus has not been studied in detail as only two skulls and one complete specimen were available for examination. LITERATURE Literature concerning the anatomy of lizards is varied, widely scattered and incomplete. Because of the large amount of material dealing with this subject, this discussion will be limited, with some exceptions, to that literature which pertains to those anatomical features treated in this paper; namely the anterior osteology and myology, hyoid bones, sternum, the tongue, and the hemipenes. One of the earliest discussions of the head-osteol- ogy or myology of lizards is that of Mivart (1867) who published a detailed account of the myology of Iguana tuberculala (Iguanidae). This work was fol- lowed by Mivart's (1870) paper on the myology of Chamaeleon parsonii (Chamaeleonidae). The latter is detailed and when used with his paper on Iguana con- stitutes two of the most complete discussions of liz- ard myology in the literature. Sanders (1870) published an account of the my- ology of Platydactylus japonicus (Gekkonidae) which is a comprehensive presentation but lacks adequate illustrations. Sanders (1872) again published a lizard myology, with an account on the musculature of Lio- lepis belli (Agamidae). As with the earlier papers of Mivert, the paper is well illustrated. Gervais (1873) published a brief note on the skull and teeth of the Australian agamid Molock. Notes and illustrations dealing with the myology of Phrynosoma coronatum (Iguanidae) were related by Sanders (1874). Parker (1880) described the skull of Lace rta agilis, L. vihclis and Zootoca vivipara (Lacertidae). That Department of Biology, Soutticrn Connecticut State College, New Haven, Connecticut. "Department of Zoology. Brigham Young University, Provo, Utah. work WHS followed by De Vis's (1883) paper on the myology of Chlamydosaiims kingii (Agamidae). Unfortunately, his paper was poorly illustrated. Boulenger (1885 to 1887) published his monu- mental catalogue of lizards in the British Museum in which are scattered his observations on the osteology of lizards, including a discussion of the distinctive cranial features of Amblyrhynclms, Bnichyloplnis. Conolophus, Ctenosaura, Cyclura and /^;w;w (Iguani- dae). Gill (1886) reviewed Boulenger's classification system for lizards and summarized the important osteological differences between the families. Boulen- ger ( 1890) further summarized his osteological obser- vations on the distinctive cranial characters of the iguanid lizards related to Iguana. Even at this early stage of investigation, the iguanine line of evolution was recognized in the family Iguanidae as a natural group. All seven genera listed by Boulenger are today still considered to be iguanines. Boulenger (1891) published a series of remarks concerning the osteol- ogy o'i Hcloderma and presented a conclusion for the systematic position of the family Helodermatidae. E. D. Cope was also actively publishing on lizard anatomy during this period. Cope (1892a) com- mented on the homologies of the posterior cranial arches in reptiles, and his conclusions in this matter have laid the foundation for understanding the com- ponents of the posterior skull of lizards by later workers. During the same year. Cope's ( 1892b) classic work on lizard osteology was published. Not only does Cope provide a comparison of the cranial osse- ous elements, but he describes in detail osteological features of the iguanines, Dipsosaiinis and Saiiro- maliis. This material was also incorporated into Cope's ( 1900) comprehensive taxonomic work. The German worker Siebenrock, during the close of the 19th century, made several contributions to our knowledge of the anatomy of lizards. He pub- lished a brief paper on the skeleton of Uroplatiis fiiii- briatus (Gekkonidae) (1892a) and a more lengthy dis- cussion on the skulls of skinks, anguids and Geirho- saurus (Cordylidae) (1892b). These papers were fol- lowed by Siebenrock's ( 1893) discussion of the skel- eton of Braukesia superciliam (Chamaeleonidae); an account of the skeleton of Lacerta simonyi (Lacert- idae) (1894); and a comprehensive discussion on the skeleton of the agamid lizards ( 1895). Bradley (1903) discussed the muscles of mastica- tion and the movement of the skull in lizards. Broom (1903) named Paligiiana wliitei (Eosuchia) from the Triassic beds of South Africa. This find is of consider- able importance as it may represent an animal ances- tral to lizards. The presence of this fossil also estab- lishes the great geologic age of lizards in general. He also studied (1903b) the development of pterygo- quadrate arch in lizards. Following these investiga- tions, Bcddard ( 1905) published notes on the skull of Uroiuaslix (Agamidae), and in a separate paper dis- BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN cussed some aspects of Chlaniydosaunis kingi and other agamids. Kingsley (1905) examined the reptile jaw bones and figured the medial stnface of the Iguana (Iguanidae) mandible. Beddard (1907) exam- ined the internal anatomy of several genera of lizards and described the uniqueness of various characters to particular families. Bryant (1911) revised the iguanid genus Phryno- sonia and its synonum Anota. In this paper he pre- sented some osteological observations on the species and genera treated in the study. A most useful paper on the phylogeny of jaw mus- cles in vertebrates was published by Adams (1919). Although the paper is concerned with reptiles in gen- eral it describes the jaw musculature of Iguana (Iguan- idae) and Varanus (Varanidae) in particular. Kesteven (1917) analyzed the pterygoids and parasphenoids of reptiles and amphibians. Rice (1920) described the development of the skull in the skink Eiimeces quingialineatus. Camp ( 1923) published his classic work on the classification of lizards, based on their anatomy, in this account, Camp figured the throat musculature of Sphenodon ( Rliynchocephalia), Amphisbaenia ( Amphisbaenidae), Cok'ony.x (Eublepharidae), Umplalus (Gekkonidae), Typhliips. ( Typhlopidae), Tupinainbis (Teiidae), Varanus (Varanidae), Genhosaurus. Zononis, Cha- macsaura (Cordylidae), Lialis (Pygopodidae), Brachyl- opluis. Plirynosoma (Iguanidae), Calmes (Agamidae), Clianiacleon (Chamaeleonidae), Xantusia (Xantusi- dae), Trachysawus (Scincidae). Lacerta ( Lacertidae), Heloderma (Helodermatidae), Gerrlionotus (Anguin- idae), Xenosaurus (Xemisam'idae). A nniiila (Anniell- idae), and Gekko (Gekkonidae). Reese (1923) ana- lyzed the osteology of Tupinanihis nigropunctalus (Teiidae). Broom (l'-)24) discussed the origin of lizards by tracing the cranial elements of the fossil forms Youngina. Mesosuchus. and Paliguana (eosuchia). These genera were compared with modern skinks, chamaeleonids, varanids and agamids. Broom indi- cated the closeness of Paliguana to the modern lizards and suggested ways whereby Paliguana could have evolved into recent forms. Dubecq ( 1925) discussed the elevating muscles of the lower jaws in reptiles, and Williston published his treatise on the osteology of reptiles. This latter work is of interest as Williston figured skulls of Conolophus (Iquanidae), Varanus (Varanidae), Amphisbacna (Amphisbaenidae), and a chamaeleon. He also classi- fied the Squamata in the Subclass Parapsida with the lizardlike fossil Araeoscclls. Gilmore (1928) summarized the fossil lizards of Nt)rth America and discussed the osteology of many forms as well as establishing the existence of some families of lizards in North America as early as the flpper Cretaceous. Nopcsa ( 1928) presented a synop- sis of the genera of reptiles. For each family he ciled \ BIOLOGICAL SERIKS. VOL. 12, NO. 3 LVOLUTION OF THE [GU.^NINF. LIZARDS osteological cliuractcristics and summarized the fossil and recent genera found in each. Lastly, Sinitsin (1^)28) analyzed skulls in the family Teiidae and separated the family into two divisions based on cran- ial osteology. Goodrich ( 1930) published his major work on the structure and development of the vertebrates. In it he figured and described the skulls of Varamis (Varani- dae) and Lacerta (Lacertidae). Edgeworth (1931a, 1931b) presented two papers on reptile anatomy in which he discussed the development of the eye, mas- ticatory and hyoid muscles of SpheiuKlon (Rhyncho- cephalia) and an account of muscles used in opening and shutting the mouth of vertebrates. His remarks in the second paper were restricted to the lizard genera Lacerta (Lacertidae), Platydactyhis (Gekkonidae), and Calotes (Agamidae). Brock (1932) continued early investigations on lizard anatomy and the devel- opmental stages in the skulls of the geckos Lyguduc- tyliis capeiisis and PacliyJactyhis maculosa. Kingman (1932) studied the skull of the -iV-mk. Hwneces nhsD- letus. Davis (1934) published a laboratory manual for Cwtaphytus (Iguanidae) which was one of the most complete studies on lizard anatomy, in the year 1935 important papers on lizard anatomy were published by Brock, Broom, and Edgeworth. Brock's discussion dealt with the problem of temporal bones in lizards, birds, and mammals. Most of Brock's comments were relegated to skinks and geckos. Broom's work also dealt with the temporal bones and correlated the information known for the fossil Paligiiana and )' Genus Mm Mean Mas Min. Mean Max. Mm, Mean Ma\ Min, Mean Max Min. Mean Max. Mm, Mean Max, Amhlvrhvinhii^ 41.4-47.7-53 2 30.3-36,7-43,1 .732--789-.809 Amhlvrhvnchus 41,4-47,7-53,2 20.8-23.2-26,0 .402-.460-.489 Braihytophui 27.7.30 7-34.2 16,8.19.8.24,9 .535-644-726 Braihvlophus 27.7-30,7-34 2 11.2-12.0-13.8 .372-390-404 Chatarodon 12.5.13.1-14.1 8,8- 9,2-10 0 .666-705-792 Chalarodon 12.5-13,1- 4,1 4.4- 4.8- 5.1 .335-368-400 Conolophus 85 0-79.1-86.0 37.0-54.1-64.3 .618-.679-,747 Conolophus 85.079 1-86,0 20.6-30.5-34,3 .344-.379-.398 Ctenosaura 30.1-40 5-54.1 17.6-24.0-32.1 .584-.593-,605 Ctenosaura 30,1-40 5-54-1 9.7-12.8-16.4 .303-316.331 Cvclura 45, 773 3-97.1 26.7-43,6-56.6 .582-593-623 Cvclura 45 7-73 3-97,1 15.2-23.9-24.6 ,306-,326-,334 Dipsosaums 22.2-22,3-23.9 14,0-16,6-18,3 .700-.745-7S1 Dipsosaums 22,2-22,3-23.9 7.3- 8.6- 9.4 .365,386-400 Iguana 49.3-60,0-70,3 28.1-35.4-36.5 .5 19-549-590 Iguana 49,3-60.0-70.3 17.6-20.6-22.6 .318-.344-.356 Opiums 18-1.21.6-30,6 11.2-14.5-20.0 .608-.670--729 Opiums 18 1-21,6-30.6 6.4- 7.7-10.3 .336-357-386 Sauromalus 23,4-34.4-48.1 14.3-22.6-32.8 .6 11-65 3- 684 Sauromalus 23,4-34,4-48,1 6.8- 9.9-14.1 .273-.286--308 10 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN For the sake of convenience the skull has been divided into a posterior occipital segment and an an- terior maxillary segment. The occipital segment forms a median axis for the attachment of the neck and articulation of the re- mainder of the skull. It consists of two parts, (a) the braincase (basisphenoid, basioccipital, prootic, exoc- cipital. supraoccipital. and the associated semicircular canals), and (b) the foramen magnum (enclosed by the basioccipital, exoccipital and supraoccipital). A tripartate occipital condyle is located on the posterior end of the basioccipital and the lateral exoccipital in all genera of iguanine lizards. Basisphenoid Basisphenoid (Figures 1 and 2) forms a portion of the floor of the braincase, is bordered posteriorly by the basioccipital, and is attached dorsally to the pro- otic bone. Anteriorly the bone is expanded into two anterolateral basipterygoid processes which articulate laterally, with the pterygoid bones. Anteromedially the basisphenoid is extended forward as the parasphe- noid process. The basisphenoid forms points of origin for the inferior part of the protractor pterygoideus muscle. Measurements of the length of the basisphenoid were made from the suture between basisphenoid and basioccipital, to the beginning of the parasphenoid process. Width was computed as the distance between the widest extension of the basipterygoid processes. An examination of the ratio means in Table 3 reveals that the lowest ratio is possessed by Chalanidon (.360) while the higliest is that of Opiums (.755). Among the New World genera, Dipsosaums (.469) has the lowest ratio and Igiiana (.652) has the highest. A low ratio indicates that the bone is much longer than wide, whereas the higlier ratios indicate bones that have lengths and widths almost equal. Observations of the bone's position in the skull indicates some variability in the articulation between basipterygoid process and the pterygoid bone. This articulation occurs medial and posterior to an expan- sion of the pterygoid bone just posterior to the ptery- TABLE .^ BASISPHENOID BONES Lcngrh Widdi Widrh-Longih Ratio Min. Ml-jd Mj\. Mm, Mean Mjs. Mm. Mcjn Md\ Amhiyrfniuitin Brachvlnpliiis Chalarudt'it Comitophu^ Crfiiosoiira Cvcliira Dipsusannis Opiiiris Saiirumaliii .SI- 6 l> 7.0 4 1- 4 s. 5 n 1 II- II- 1-1 7 5- 1 11.5- 1 J- 1 -1.9- 5..1- 7 4 d 2-lll(H4 4 :,5- 21- }j 1-1- 7.9. S.7 2 5- 3,4- 5,1 3.»- 5.7- «.l '),I-I2,«.|6.7 6.9- 7.9- 9,5 2 K- 2.9- .1.2 112-19 9-2.1.9 h 7- 8.1-in 7 III J-I6.I-22.3 5.4- 6-3- 6.9 HI.VI 2.2-14.5 3.4- 4.5- 6.6 6.7- 9.6-12.2 .419-.4S2-.56n .526-.570-.594 .333-.360-.406 . 506-53.1-567 .549 .633-.69I .546-.619-685 .412-.469-536 .6110-65 2- 699 .717-755-780 .500-.5H9-.669 gold's midpoint. This point of articulation is almost completely obscured ventrally by the pterygoid bone m Amblyrhynchus, Chalawdun, and Opiums. In Dip- sosaums the articulation is visible ventrally but occurs dorsal to the ventral border of the pterygoid. In Bracliylophus. Conolophus, Ctenosaura, Cyclura, Iguana and Sauromalus the anterior articulating por- tion of the basipterygoid process is partially obscured and two thirds of the articulation occurs on the ven- tromedial border of the pterygoid bones, rather than on the medial face. Basioccipital Basioccipital (Figures 1 & 2) forms part of the occipital condyle and the posterior floor of the braincase. It is the main point of attachment for ven- tral axial musculature. Anteriorly the basioccipital joins the basisphenoid and dorsolaterally it attaches to the exoccipitals and the prootic bones. The third bundle of the longissimus dorsi muscle attaches here. Length and width measurements were made of the basioccipital with the length being taken from the suture between the basisphenoid and basioccipital to the posterior tip of the occipital condyle. The width of the bone was considered as the distance between the tips of the lateral extensions of the spheno- occipital tubercles. An examination of the ratio means in Table 4 indicates that the highest ratios for the basioccipital are possessed by Brachylophus (.VC-)), Ctenosaura (.664), Amblyrhynchus (.663), and Sauromalus (.649) while Dipsosaurus (.369) and Conolophus (.477) possess the lowest ratios. High ratios denote that lengths and widths are nearly equal for the bone. TABLE 4 BASIOCCIPITAL BONES Length Width Width-Length Ratio Mm. .Mean Ma Min Mean Max. Min. Mean Max. ■Xmhh'rhvnihus BrOihvlophns Chaliiriiduii Conoltipfnn Ctenosaura Cyclura Dipwsaimis Ifniona Oplitnii Saiironialiis 8.7- 9.81 1.6 5.0- 6.2- 7.4 2.0- 2.1- 2.2 8.5- 9.4-10.7 4.2- 5 7- 6.7 6.2- 94-12.1 2.1- 2.2- 2 5 6.6- 8.1. 9.8 3.1- 3.3- 4.11 3.4- 5.8- 7.8 12.3-15.0-17.6 3.5- 4.2- 5,0 3.2- 3.4- 3.6 15.0-20.0-23.4 6.2- 86-10.5 I 1.615. 1-24.2 5.9- 6.1- 6.7 I I 5-16.0-19.6 5.4- 5.7- 6.4 5.1- 9.8-13.0 .573-.663.739 .648-.709-.781 .600-,625-.656 ,420-,477-,5S0 .650-.644-.677 50n-.534-.596 . 350-369-389 .5O0-.513-.573 .534-.583-.625 .600-.649-.696 Prootic Prootic bones form the anterolateral wall of the braincase and contain the elements of the ear. Poster- iorly, the prootic is bordered by the supraoccipital, basisphenoid, basioccipital. and the exoccipital. The anterior portion of the bone is surrounded by mem- branes that contain the optic nerve. The pseudotem- poralis and protractor pterygoideus muscles originate BIOLOGICAL SERIES. VOL. 1 2. NO. 3 EVOLUTION OE THE IGUANINE LIZARDS II on the prootic bone. Because of difficulties in mea- suring, the prootic was not studied in detail. Exoccipital Exoccipitals bones form the posterolateral wall of the braincase and the lateral parts of the occipital condyle. Mediolateral articulations form with the par- ietal, supratemporal and quadrate bones. The exoc- cipital also articulates at its most lateral projection with the prootic bone. The longissimus dorsi and epi- sternocleidomastoideus muscles insert on the para- occipital process of the bone. The length of the exoccipital bone was measured from the lateral wall of the foramen magnum to the point of articulation by the paraoccipital process with the squamosal and quadrate bones. Width is repre- sented as the distance between the exoccipital articu- lation with the supraoccipital bone and the union with the basioccipital at the occipital condyle. As Table 5 indicates, the lowest ratio means for exoccip- itals are possessed by Dipsosaums (.594) and Conolo- phus (.626). The largest ratios are found in Brachylo- phus (.858), Amblyrhynchm (.830), and Chalarodon (.813). As with the other bones, near equal relation- ships between length and width are expressed as higli ratios. TABLE 6 SUPRAOCCIPITAL BONES TABLE 5 EXOCCIPITAL BONES Length Width Width-Length Ratio Genus Min. Mean Max Min, Mean Max. Min. Mean Max. Amblyrhynchus 8.9-11.0-13.0 8 5-11.6-16.0 .762-.830-,861 Brathylophus 5.0- 5.5- 6.3 4.5- 4.7- 5.0 .793-.858-,900 Chalarodon 2,2- 2.2- 2.4 1.8- 1-8- 1.9 .782-.8 13-863 Conotophus 8.5-12.4-14.2 13.0-20.0-23.8 .571-.626-,6S3 Ctenosaura 3.0- 4.9- 6.9 4.5- 7.8-12.5 .552-.649-.724 Cyclura 6,2.|1. 3-15.8 8.5-17.1-25.8 .605-.661-,720 Dipsosaums 2.8- 3.3- 3.6 5,4- 5,6- 5.8 .509..594-,648 Iguana 7.0- 7.7. 8-2 9,4-11,2-11,9 .606-.671.,744 Opiums 3.3- 3.9- 5.0 4,2- 5,1- 6,8 .735-.774-,809 Saiironialus 4 0- 5,6- 8.2 5.1- 8,2-12,4 ,602-,692. 759 Supraoccipital Supraoccipital (Figures 3 & 4) forms the roof of the posterior part of the braincase and the dorsal rim of the foramen magnum. It articulates with the pro- otic bone anterolaterally, the exoccipital posteriorly, and the parietal at its anterior extreme. The supraoccipital lengths were measured as the distance between its posterolateral sutures with the exoccipital bones in the area of the foramen magnum. The width was measured as the distance between the suture with the parietal bone anteriorly, and the dor- sal lip of the foramen magnum posteriorly. Table 6 indicates that the genera possessing supraoccipitals with the lowest ratios include Chalarodon (.583), Opiums (.675), and Sauromalus (.680), wliile the highest ratios are found in Brachylophus (.982) and Iguana (.919). Lenplh Width WTdth-Lenglh Ratio Genus Min, Mean Max, Mm. Mean Max, Mm. Mean Max. Anihlvrhviithtt)i 6.4- 6.8- 7-8 9.0- 9.8-11.9 .655-.700-.755 Brachylophus 5-5- 5.9- 6,1 5.2- 5.4- 5,6 .852-.906-.982 Chalarodon 1.2- 1.5- 2,1 2,4- 3.0- 3,7 .413-.524-.583 Coiiolophiii 11.4.14.7.21,1 7,4-13.5-13.9 .648-.706-.793 Chnosoiira 3 3- 4.7- 6,7 5.1- 6.9- 9.2 .632-.677-.739 Cvchira 5.8-10.9-14.6 8, 1-14, 4-18,1 .700-.75 1-806 Dipwsaiiriis 3.7- 4.0- 4,2 4,6- 4.8- 5,2 .795-.820-,874 Iptana 7 9. 8.6- 9,6 7,6- 8.8- 9.4 .900-.949-.979 Ophinii 2,4- 3,2- 5,0 4.0- 5.2- 7,4 .521-.596-.675 Sattromahn 3.3- 4.6- 6.8 5,1- 7. 2-10.0 .538-.632-.680 Orbitosphenoid Orbitosphenoid is a vertical element surrounding the optic foramen. The anterior border forms the pos- terior margin of the optic foramen and the inferior process forms an area of origin for the superior rectus muscles at the eye. The orbitosphenoid also has con- nections with the prootic bone and the alar process of the basisphenoid. It was found to be absent in most of the cleaned skulls in the collections examined. The combination of bones in the anterior region of the skull is referred to as the maxillary segment. It consists of four parts, (a) the palate (pterygoid, ectopterygoid, vomer, palatine, premaxilla, and max- illa), (b) orbits (frontal, postfrontal. and jugal), (c) nasal capsule (nasal, prefrontal, lacrimal, and septo- maxilla). and (d) temporal fenestra (parietal, supra- temporal, postorbital, squamosal, quadrate and epi- pterygoid). These bones are discussed as listed above. Pterygoid Pterygoid (Figures 1 , 2, 3, 4, 5 and 6) extends pos- teriorly as the posterior part of the palate. The ptery- goids are paired bones sutured at the anterior end of the palatines, anterolaterally to the ectopterygoids and posteromedially to the basisphenoids. They com- prise a major area of motion between the occipital and maxillary regions as the posterolateral portions articulate with the quadrate which is movable. Articu- lating with the pterygoid on the dorsal border of the quadrate projection is the columella. At the ventral border, the pterygoid forms the posterior limit of the oral cavity and contributes an area for the origin of the pterygomandibularis muscle. On the dorsal ridge, medial to the columellar fossa, is an area of insertion for the levator pterygoideus muscle. The insertion of most of the protractor pterygoideus muscle is located on the medial surface of the quadrate process of the pterygoid bone. The posterior fibers of the pterygo- mandibularis muscles arise along the ventrolateral border of the lateral side of the quadrate process. Measurements taken of the pterygoid bone include length; represented as the distance between the anter- ior portion of the pterygoid where it sutures with the 12 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN palatine and the most posterior tip of the quadrate process; and the width as the distance between the articulation with the basipterygoid process of the basisphenoid bone and the suture with the ectoptery- goid bone. Table 7 summarizes these measurements and a survey of the ratio means indicates that the lowest pterygoid ratio (long, narrow bones) are possessed by Cyclura (.283), Sauromalus (.293), and Iguana (.309). The highest ratios (short, wide bones) are found in Bmchylophiis (.458) and Chalawdon (.435). The unique relationships of the pterygoid to the basipterygoid process of the basisphenoid bone have already been reviewed. The shape of the medial border of the pterygoid also controls the shape of the pyriform recess (Figs. 1 and 2) of the palate. This shape varies from a gradually widening slit as seen in Brachylophus, Chalawdon and Opiums to a more severe and rapid change in width of the recess as seen in Amblyrhynchus. Conolophus. and Cyclura. The remaining genera are intermediate between the above conditions. TABLE 7 PTERYGOID BONES Lt-nglh Widlli Width-Lcnglh Ralio Min. Mejn Ma\. Min. Mean Max. Mm. Mean MaV- Amblyrhynchus 23.9-28.8-32.6 8.4-10.3-13.5 309-.346-4I4 Brachylophus 14.0-15.5-18.4 6.1- 7.0- 7.5 380 ,458-535 Chatarodoti 5 6- 5.7- 5.7 2.4- 2.4- 2.6 ,421 435 464 Conolophus 34.2-51,0-59.5 11.8-18.8-22.3 ,345 ,367 .382 Ctenosaura 18.7-24.5-33.5 5.8- 7.5-10.8 ,287 311 355 Cyclura 29.1-39.0-63.1 7.3-13.5-20.6 ,250 283 ,326 Dipsosaurus 10.1-12.0-13.1 4.1- 4.2- 5.3 .333 ,353 404 Iguana 26.5-31.5-35.6 7.0- 9.5-11.1 ,264 ,309 ,364 Oplurus 8.4-11.6-19.6 2.9- 3.9- 6.2 ,316 ,347 ,377 Sauromalus 13.0-19.8-30.0 3.9- 5.8- 9.1 ,252 ,293 .343 Ectopterygoid Ectopterygoid (Figs. 1, 2, 3, 4, 5 and 6) forms a brace between the palate and the external roofing bones. Laterally the ectopterygoid sutures with the jugal and the posterior process of the maxilla, and mesially with the pterygoid. Dorsally it forms the posterolateral part of the floor of the orbit. Measurements made on the ectopterygoid include the length, taken as the distance between the suture with the pterygoid and the suture with the jugal and maxilla. The width was measured as the greatest diameter of the bone at its point of union with the jugal and maxilla. These measurements are summar- ized in Table 8 and an examination of the ratio means shows that the lowest ratio (long, narrow bones) is found in Amblyrhynchus (.275) and the form with the highest ratio (short, wide bones) is Dipsosaurus (.600). Vomers Vomers (Figs. 1 and 2) are paired bones forming the most anterior part of the palate, the medial bor- ders of the fenestra vomeronasalis externus and the medial borders of the fenestra exochoanalis. Poster- iorly the vomer is attached to the palatines and dor- sally to the medial surface of the maxilla. Medially the vomers support the nasal septum and the cartilage of Jacobson's organ. At the extreme anterior end the vomer sutures with the premaxilla. The length of the vomer was measured from the anterior suture with the premaxilla to the most pos- terior point of the suture with the palatine bone. Width of the vomer was the distance between the medial border of the vomer at the ventral midline and the most lateral border where it attached to the max- illa. Table 9 summarizes the measurements of vomer bones and the ratio means show that the shortest, widest, vomers (highest ratios) are possessed by Chal- arodon (.711) and the narrowest longest vomers (low- est ratios) are possessed by Amblyrhynchus (.253), Cyclura (.274), and Ctenosaura (.278). The vomers possess a small blunt projection that protrudes from the lateral border of the bone into the opening of the fenestra exochoanalis and fenestra vomeronasalis externa and separates the opening. The possession of this anterolateral projection is seen in all ten genera. Some genera have a secondary projec- tion which extends into the lumen of the fenestra vomeronasalis externa. This projection can be seen in Brachylophus. Ctenosaura, Cyclura, Iguana, and in Sauromalus. TABLE S TABLE 9 ECTOPTERYGOID BONES VOMER BONES Length Width Width-Length Raliu Genus Length Width Width-Length Ralio Mm, Mean Max Min. Me n Max. Min, Mean Max. Mm. Mean Max Min. Mean Max. Min. Mean Max. Amblyrhynchus 7 2- 8 9-10 4 1,8. 2.4 3.1 250-275- .103 A mbhrhynchus 1 1.2-13.0-15.0 2.8- 3.2- 4.0 ,200-253-294 Brachylophus 3,0- 4 1)- 5 II 1,0- 1,5 2.2 .333-371.440 Brachylophus 6.5. 7.0- 7.6 2.4- 2.5- 2.7 .315-.359-.380 Chalarodon 14. 14. IS 0,6- 0.6 0.7 ,40O-.431--466 Chalarodon 1.8- 1,9- 2,0 1.3- 1..3- 1.4 .70O-.711-.722 Conolophus 10-9. 15 ,1-17 ^ 5.0- 6.5 7.5 .405.429-458 Conolophus 12,2-13.6-17,1 4.1- 5.5- 7.0 .335-,372-.409 Ctenosaura 3 3. 5 6. K '1 1 4- 2.3 3.6 404-.421-.444 Ctenosaura 7 2. 9,7.11.1 2 0- 2.9- 4.2 .270-.278-.328 Cyclura 6.1. 108-15 : 2 2- 4,1 6.3 356-377. .414 Cvclura 1 1.6-18 2-25-8 3.1- 5.5- 7,7 .235-.274-.335 Dipsosaurus 2.-1. 2,6. 2,8 14- 1,5 1.8 55I-.600-.642 Dipsosaurus 3.9- 4.2- 4.8 2.0- 2.|. 2.3 .479-.497-.512 Iguana 6.1- 7.9- 9.3 3,0- 3,5 4,0 -41 5.-448-49 1 Iguana 13.5-17,4-21,1 4,8. 5,5- 6.9 .265-.322-.389 Oplurus 2 2- 2.6- 3,6 0 9- 1.1 15 .409-.4 19-434 Oplurus 3.1. 4.2- 6.0 14- 1,6- 2 2 .365-.399-.45I Sauromalus 2.4- 5,4- 8,5 1.0- 2,0 3,2 .323-392.457 Sauromalus 5.7. 7.8-11.8 14. 2 6- 3.7 .245-.329-.390 BIOLOGICAL SERIES. VOL. 1 2, NO. 3 EVOLUTION OF THE IGUANINE LIZARDS Palatine Palatine (Figs. 1 , 2,3, and 4) bones form the main part of the palate, the floor of the orbit and nasal capsule. This bone has three processes; the anterior or vomerine, forms the posterior floor of the olfactory capsule; the pterygoid process, which attaches dor- sally to the pterygoid, forms the medial rim of the inferior orbital fossa and the floor of the orbit; and the maxillary process attaches dorsally to the pre- frontal and ventrally to the jugal and maxillary bones. The length of the palatine was taken as the dis- tance from the anterior suture with the vomer bone at the midline to the most posterior extension of the suture with the pterygoid bone. The width of the palatine bone was considered to be the distance from the palatine medial border at the skull's midline to the lateral suture between the palatine and the max- illa. Table 10 summarizes these measurements for the ten genera under discussion. The ratio means column indicates that the shortest and widest bones (highest ratios) are possessed by Chalarodon (.846) while the longest and narrowest bones (lowest ratios) are found in Cyclura (466). TABLE 10 PALATINE BONES Length Width Width-Length Ratio Mm. Mean Max. Min. Mean Max, Min, Mean Max. Ambtyrhyntinis 12, 0-13. 2-14.4 7.7- 8.6- 9.2 .604--653- 721 Brachylophus 8.0- 8.2- 8.4 4.5- 4.9- 5.4 .560-.601-.658 Chalarodon 2.8- 2.8- 2.9 2.3- 2.4- 2.5 .821-.846-.862 Conotophus 14.2-23. 1-28.4 9.1-14.4-17.7 .S77-.627-.648 Ctenosaura 8.5-10.8-15,1 5.0- 6.0- 8.2 .543-.565-.588 Cvchtra 12, 1.20.9.30.0 6.0- 9.9-13.9 ,406-.466-,S02 Dipsosatinn 5,6- 6.4. 7,9 3.7- 4.1- 4.6 .582-644-678 Iguana 11.2-15.9-20.1 7.4- 9.1-10.6 .432-.598-.671 Opiums 5.9-7.5-12.0 3.4- 3.6- 5.3 .44I-.532-.576 Sauromalus 5.8- 9.7-12.9 3.6- 5.3- 7.0 .542-.557-.620 Premaxillae Premaxillae (Figs. 1, 2, 3, 4, 5, and 6) is the most anterior bone of the skull, and joins the maxilla later- ally and the nasal bones dorsally. It forms the ros- trum of the skull. The length of the maxillae was considered to be the distance from the anteroventral tip of the bone to its dorsal union with the nasal bones at the dorsal midline of the skull. The distances between the lateral sutures shared by the premaxilla with the maxilla on the ventral surface of the bones were considered to be the width of the premaxillae. The length and width values of the premaxilla are expressed in Table 11. The ratio mean column indicates the highest ratios (shortest, widest bones) is found in Conolophus (.•514) while the lowest ratio (longest, narrowest bone) are found in Cyclura {A9\), Ctenosaura (.5 \ 2) And [guana (.521). The suture between the posterodorsal portion of the premaxilla and the nasal bones differs from genus to genus, in the degree of penetration the premaxilla makes between the two nasal bones. \n Amblyrhyn- chus. Conolophus, and Iguana, the penetration of the premaxilla is very shallow with that of Conolophus forming a shallow curve and penetrating the nasals very little. Deep penetrations of the premaxilla are found in Brachylophus. Chalarodon and Oplurus. Moderate penetrations are found in the remaining genera. y TABLE 1 1 PREMAXILLARY BONES Length Width Width-Length Ratio GenuN Mm. Mean Max Mm, Mean Max, Min. Mean Max. Amblyrhynchui; 10. 1-12.6-14 4 6.3- 7,3- 8.2 .566-,585-.623 Brachylophus 7,5- 8.5- 9,8 4.2- 4.7- 5,6 ,442- 558-629 Chalarodon 2 5- 2 6- 2 9 1.4- 1.6- 2.0 ,560-61 1-689 Conolophus 15 8-17.919,3 13.6-18 4-20.5 .860-,914-.963 Ctenosaura 8,0-11.0-15.0 4.1- 5-7- 8.4 .464.5 12-560 Cvclura 11.7-21.6-29.1 5.1-10.7-15.0 .43S-.491-.566 Dipwsauriis 4,0- 4.9- 6.5 2.5- 3.1- 4.3 .607-.629-.661 Iguana 13,5-16.2-18 9 6.7- 8.4- 9,9 .48S-.S21-,S81 Opiums 4-4- 5 7- 9,1 2.5- 3.1- 5.1 .500-.542-.568 Sauromalus 4.7. 7.3-12.3 2.6- 4.3- 6.3 ,509-.603-.676 Maxillae Maxillae (Figs. I, 2, 3, 4, 5 and 6) forms the major lateral surface of the snout. The ventral margin bears a single row of pleurodont teeth. There are three pro- trusions from the main region of the maxilla. The first protrusion is the premaxillary process which overlaps and is attached to the maxillary process of the premaxilla. The medial part of the maxillae is attached to the vomer whereas the dorsal extension forms the inferior rim of the fenestra exonarina. The posterior process of the maxillae is attached to the jugal and larcinial bone dorsally, and to the ectopterygoid medially. This process forms the lateral part of the rim of the inferior orbital foramen. The third process of the maxillae extends dorsally to form the lateral wall of the nasal capsules, the pos- terior rim of the fenestra exonarina anteriorly, and dorsally attaches to the nasal and prefrontal bones. The length of the maxillae was taken from the anterior most extension of the premaxillary process where it formed a suture with the premaxillae to the posterior most extension of the maxillae where it joined the jugal and ectopterygoid bones. The width of the maxillae was considered to be the vertical dis- tance from the ventral border of the maxillae to the dorsal most extension of the bone at the point where it sutured with the nasals and prefrontals. Table 12 summarizes the maxillary measurements. The ratio mean column indicates the longest and lowest maxilla (lowest ratios) are found in Chalarodon (.334), Oplurus (.358), Ctenosaura (.371), Brachylophus (.373) and Sauromalus (311). The shortest and 14 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN widest bones (highest ratio) are found in Amblyrhyn- chus {.bl9). TABLE 12 MAXILLARY BONES Length Width Width-Length Ratio Mm, Mean Ma\- Min. Mean Max. Min, Mean Max. Amhhrhvnctms Bracliyluphua Chalarudon Conolophus Ctcnusatira Cychira Dtpsosatini'. Opli,ni\ Satirunwlm 2\ 7-25.3-28.8 lS.0-17.4-18.9 6.3- 6.7- 7.2 30.0-4L0-45.6 16.1-21.3-29.3 23.4-40.6-54.3 111,1-10.8-1 1.9 27,6-35.0-40.6 9 4-1 1.4-15,9 15 1-18,9-27.1 13. 5-15.4-17. 5 6.0- 6.6- 7.5 2.2- 2.2- 2.3 14.3-19.3-21.5 5.5- 8.0-11.7 8.9-15.5-21,4 5.0- 5.5- 6.1 11.9-13.9-15,5 .1 5- 4.1- 5.7 4.3- 7,2-10,8 596-619-.642 ,346, 373-400 , 319-334-349 432-466-483 341-371-399 .359- 383-402 495- 5 1 3- 564 IKI- 199-,431 ,346- 358-,372 ,355-,377-,398 Nasal Nasal (Figs. 3, 4, 5 and 6) forms the sloped top of the snout and partially covers the nasal capsule. The nasals attach posteriorly to the frontals, anteriorly to the premaxillae, and laterally to the prefrontals. Part of the anterior border of the nasal bone forms the dorsal border of the fenestra exonarina. The measurement of length of the nasal bone was taken from the tip of the ventral border as it formed the fenestra exonarina to the posterodorsal extension that sutured with the prefrontal. Width was defined as the widest portion of the bone from its medial suture with its opposite member to the most lateral extension of the bone where it sutured with the max- illa and prefrontals. These measurements are ex- pressed in Table 13 where the ratio mean column shows the nasals with the greatest ratio of length to width (short, wide bones) are found in Ctenosaura (.555) and Brachylophus (.522), while those with the lowest ratio (long, narrow bones) are found in Amblyrhynchus (.375). The basic shape of the nasal bones differs from genus to genus. The major ditTerences include the amount of nasal bone that borders the premaxilla, the shape of the posterior border that sutures with the frontal bone, and the shape and position of the lateral border that sutures with the maxilla and prefrontals. The nasals border a large portion of premaxilla in Brachylophus. Chalarodon and Ophtrus. A short bor- der with premaxilla is seen in Amblyrhynchus, Cono- lophus and Iguana. The posterior border of the nasal forms an interfingering suture with the tVontal bone m Amblyrhynchus. Conolophus. Dipsosaurus. Iguana, and Sauromalus. The posterior projection forms a smooth suture in the remaining genera. The shape of the posterior border of the nasal bone may be roughly straight as in Amblyrhynchus. Conolophus and Iguana or it may form a posteriorly projecting triangle as in Brachylophus. Chalarodon, Ctenosaura. Cyclura. Dipsosaurus. Opiums, and Suuromalus. The lateral borders of the nasals form a shallow curve in Brachylophus. Chalarodon. Conolophus. Ctenosaura, Cyclura, Dipsosaurus, Iguana and Oplurus. In Amblyrhynchus and Sauromalus this curvature is disrupted at its anterior end by an inden- tion for the dorsal projection of the maxilla. TABLE B NASAL BONES Length Width-Length Ratio Mm. Mean Max, Min, Mean Max, Min, Mean Max, Amhiyrhyiuhiii Brachylophus Chalarodon Conolophus Ctenosaura Cyclura Dipscsaunis Iguana Ophmis Sauromalus 17 2-20.1-24 1 7 2- 8 7-12.4 2,3- 2,5- 2.7 15 6-22.4-26.1 8,1-11.5-14.8 12,6-21.4-27.0 6.8- 7,2- 7,5 14.5-19,7-22,5 4.5- 5 3- 7,7 5 1- 8,8-13,0 6.3- 7,5- 8,4 3-9- 4,4- 5.3 1.1- 1.1- 13 8.6-10.6-11.9 4.2- 6.4- 8.6 5.8-10.2-15.1 2.9- 3.6- 3.9 6.4- 8.9-10.5 1.9- 2.4- 3.7 2.5- 4,3- 6.1 .348-375 .427-.522 .423-.455 .409-.480 .507-.555 .428-.472 .426-.500- .443-449- .422-448 ,434-496 -.416 -564 -481 |-,551 -618 -.559 -541 -466 -480 -583 Prefrontal Prefrontal (Figs. 3, 4, 5 and 6) forms the anterior angle of the orbit. Medially it attaches to the frontal and nasal bones, ventrally to the maxillae and poster- iorly to the lacrimal. Length measurements were taken from the suture between the prefrontal and lacrimal bones at the anterior lip of the orbit, to the suture between the prefrontal and frontal bones on the dorsal lip of the orbit. The width of the prefrontal bone was consid- ered to be from the suture between the prefrontal and lacrimals to the medial point where the frontal, nasal, and prefrontal bones suture together as seen in Table 14. The prefrontals with the greatest ratio of length to width (shortest, widest bones) are possessed by Amblyrhynchus (.776). Those genera with pre- frontals having the lowest ratio (long, narrow bones) include Chalarodon (.512) Sauromalus (.553). and Brachylophus (.571 ) (Table 14). TABLE 14 PREFRONTAL BONES Genus Length Width Width-length Ratio Min. Mean Max. Min. Mean Max, Mm, Mean Max, Amhlvrhvnchus 13.514.9-15 9 10,0-13,3-18,9 ,718-.776-.849 Brachvlophus 8,4- 9.0-10.1 4 5- 5,1- 5,8 .535-.571-.619 Chalarodon 2.9- 2.9- 3.1 1-5- 1,5- 1.6 .500-5 12-.5 17 Conolophus 17.5-22.7-25.7 10.9-14.6-17.3 .595-.636-,676 Cfi'uosaura 8.0-10.5-13.4 5.0- 6.8- 9.1 .625-.645-679 Cvclura 10.5-18.3-23.8 5.7-11.3-14.9 .542-.603-,668 Dipsosaurus 6.3- 6.6- 6.8 4.2- 4.3- 4.5 .656.662, 666 Iguana 12,4-16.0-20.3 8,2-10.7-13.9 .621.672-723 Ophmis 4 4- 5 4- 8 0 2.8- 3.4- 5.1 6I2-.636-.676 Sauromalus 5,3- 8.1-1 1 II 2.7- 4.6- 7.1 .500-553-645 Lacrimal Lacrimal (Figs. 5 and 6) is a small bone on the BIOLOGICAL SERIES. VOL. 1 2, NO. 3 EVOLUTION OF THE IGU.ANINE LIZARDS 15 anteroventral rim of the orbit. Dorsaily it is attached to the prefrontal, anteriorly to the maxillae, ventrally to the jugal. and ventromedially to the prefrontal. Measurements taken on the lacrimal include length as the greatest diagonal distance from the anterodor- sal border as it sutures with the prefrontal and max- illa to the posterior border on the rim of the orbit as it sutures with the jugal. Width was considered as the vertical distance between the dorsal border of the lacrimal at the rim of the orbit to the ventral border of the lacrimal at its suture with the maxilla. Those measurements summarized in Table 15 show the low- est ratio (long, narrow bones) for the lacrimal bone is found in Chalarodon (.293). The highest ratio (short, wide bones) is that for Conolophus (.542), Cteno- saura (.532), Cvclura (.526), and Brachvlophiis (.523). In shape the lacrimal differs from genus to genus. The most common form of the bone is that of a slightly curved rhomboid. This rhomboid shape is most perfectly reproduced in Conolophus. Cteno- saura. Cvclura and Iguana. In Amblyrliynchus the bone is reduced to a splinterlike structure while in Brachylophus, Chalarodon, Dipsosaurus and Ophirus the rhomboid shape is distorted by the curvature of the bone to fit the rim of the orbit. In Sauromalus the bone has its dorsal part reduced so as to form a rougli trapezoid shape. TABLE 15 LACRIMAL BONES Genus Length Width Width-Length Ratio MIn Mean Md\. Min Mean Ma\, Min. Mean Max. Amblvrhvn(.hiis 3.2- 4,0- 5,0 1.5- 1,9- 2.7 .405-.471..540 Brachylophus 2.4- 3,3- 4,3 1.4- 1.6- 1,9 .428-523-640 Chalarodon 1.0- 1,0- 1,1 0.3- 0.3- 0,3 .272-.293-.300 Conolophus .14- 6 1- 7,8 2.0- 3.3- 4.2 .507-.542-.588 Clcnosaiira 2,4- 3,«- dO 1.2- 2.0- 3.3 .500-.532-.555 Cyclura .1.5- 8,3-13.4 2.0- 4.2- 5.9 .506-526- 588 Dipsosaiinn -) ■). 2.6- 3.2 1.0- 1.0- 1.0 .312-.392-4>4 Iguana 7.7- 9,4-10,8 3.4- 4.2- 5.0 .441-451-462 Opiums 17- 1,7- 1,8 0.6- 0.6- 0.7 .352-.370-,38K Sauromalus 2.3- 3.9- 5,7 1,0- 1,5- 1,9 .333-.385-,4l.l Septomaxilla Septomaxilla is found within the nasal capsule where it covers Jacobson's organ and houses the anterior part of the nasal capsule. Because of the dif- ficultness in taking measurements of this bone, it has not been studied in detail. Frontal Frontal (Figs. 3, 4, 5 and 6) forms the dorsal bor- der of the orbits and the anterior roof of the brain- case. At its posterior extreme the frontal is attached to the parietal and postorbital bones. Anteriorly, it is sutured to the nasal and the prefrontal. The pineal foramen penetrates the posteromedial portions of this bone or the suture it shares with the parietal. Table 16 represents length and width measure- ments of the frontal bone. Length was considered to be the distance from the most anteromedial suture shared with the nasals to the most posteromedial suture shared with the parietal bone. The width of the frontal was measured as the distance between the most lateral posterior projections as they sutured with the parietal and postfrontal bones. The greatest width to length ratio (short, wide bones) is to be found in Ctenosawa (.936) and Brachylophus (.907). The smallest ratio (long, narrow bones) is that of Amblyrliynchus (.629). The shape of the frontal is basically the same for all genera but differs in some small features between genera. The greatest difference in shape occurs at the anterior end where the frontal sutures with the nasals and prefrontals. In Conolophus the anterior border interfingers with the nasals but is essentially straight. In Brachylophus. Ctenosaura. Cyclura, Dipsosaurus. Iguana. Opiums, and Sauromalus. the anteromedial portion is triangle shaped and forms a wedge between the paired nasals. This triangle in Dipsosaurus is still more unique by being bifurcated at its tip by a secondary triangular projection of the nasal bones. In Ctenosaura and Sauromalus the lateral sides of the frontafs triangular projection is further bifurcated by secondary triangles. The lateral sutures of this process in Brachylophus interfinger with the prefrontals and nasals but is essentially straight as it is in Cyclura. Amblyrliynchus and Chalarodon lack this anterior traingular projection and in its place possess a depres- sion which fits around a triangular projection formed by the posterior borders of the nasals. Thus in the latter two genera the anterior portion of the frontal bone is bifurcated and sends a projection anteriorly which serves to separate the prefrontals and nasals for a portion of their length. The frontals of Amblyrliynchus, Brachylophus. Conolophus and Cyclura are wider than long. In the other genera the frontals are longer than wide. The placeiTient of the pineal foramen in the frontal bone or the suture between the frontal and parietal bones is perplexing. The foramen was found to be in TABLE 16 FRONTAL BONES Length Width Width-Length Ratio Genus Min. Mean Max. Mm. Mean Max. Mm, Mean Max. Amblyrhynchus 12.1-15.5-18.1 18.9-24.7-30.3 .597-.629-.653 Brachylophus 11,0-12,6-14.2 12.6-13.8-14.8 ,808-,907-.985 Chalarodon 5,9- 6,0- 6.2 4.8- 4,9- 5,0 806-,840-,847 Conolophus 16.6-21,3-24,4 22,4-31.2-36,3 ,63l-,689-.746 Ctenosaura 1 1,6-14,2-16.6 11,1-14.7-18,7 ,887-,936-,963 Cvclura 16,3-22,5-29,1 17,3-25,8-34,1 ,839-,886-,942 Dipsosaurus 8,4- 9,5-10.3 7,5- 8.3- 9,2 ,838-.873-,910 Iguana 18.3-21. 8- 25. 9 15,4-22,1-24,1 ,806-,894-,946 Opiums 7.3- 8.6-12,1 6,0- 7,0-10.0 ,800-,815-,835 Sauromalus 11.0-13.7-18,1 9.6-12.0-17.4 ,800-.873-.966 16 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN the suture between the two bones in all specimens examined of Brachylophus, Chalarodon, Ctenosaitra. and Igitana. The foramen appears completely em- bedded in the frontal bones in one specimen each of Ainblyrhynchus, Conolophus, and Opiums, whereas other specimens of these genera possessed a foramen in the suture. In Cychira the pineal foramen is found in the frontal bone in three of four specimens examined while it occurred in the frontal bone in all four specimens oi Dipsosaurus and in five of six speci- mens of Sauromahts. Post frontal Postt'rontal (Figs. 3, 4. 5 and 6) forms a small part of the posterodorsal margin of the orbit. Posteriorly this bone is sutured to the frontal, and laterally to the postorbital and the parietal. The length of the postfrontal was measured as the distance between the extremities of its longest axis. The width was the distance between the parallel bor- ders on the axis at right angles to the length. The values for these measurements are presented in Table 17 and it can be seen that the genus with the smallest ratio (longest, narrowest bone) is Chalarodon (.200), while Oplurus (.625) has the largest ratio (shortest, widest bones). The postfrontal is usually splinterlike in shape as it is in all genera except Cyclura. Iguana and Opiums. In Cyclura the anterolateral portion of the bone forms a short projection out over the posterodorsal part of the orbit in some individuals. This condition is espe- cially well developed in Cyclura cornuta. In Iguana the lateral portions of the postfrontal is developed into a prominant knob on the anterodorsal face of the postorbital bone. In Oplurus the postfrontal is small, almost spherical in shape, and in at least one skull (MCZ 37191) this bone could not be located. TABLE 17 POSTFRONTAL BONES Length Width Widlh-Length Ratio Genus Min. Mean Max. Min Mean Max. Mm. Mean Max. Amhlyrh\-nthu<. 6.9- 8.7.10.3 3.8- 4.2- 4.6 .436-.489. .550 Brathylophtis 4.0. 4.7. 6.2 1.1- 13- 1.8 .275-.286- .300 Chalarodon 0.5- 0.5- 0.5 0.1- 0.1- 0.1 .200-.200- 200 Conolophus 10.0-13.6-16.7 2.3- 3.5- 4.4 .230-256- ,269 Cwnosaiira 39- 4.8- 5.8 0.8. l.j. 1.7 .207-.234- .293 Cyclura 6.5- 9.4-12.4 1.9- 2.7- 3.9 .219-.287- 326 Dtpsosaurus 2.4- 2.7- 3.0 0.5- 0.6- 0.8 ,204-235- 266 Iguana 4.6- 7.0- 8.4 2.0- 2.7- 3.2 .360-.394- .434 Oplurus 0.2- 0.2- 0.3 0 1- 0.1- 0.3 ,500-625-1.000 Sauromalus 2.1- 3.8. 5.6 0.7- L3- 1.9 ,3IO-,380- ,388 to the ectopterygoid. The dorsal surface forms the anteroventral wall of the orbit. The length of the jugal is considered as the hori- zontal distance between the most anterior projections of the bone as it sutured to the lacrimal and maxillae, to the most posterior projection which sutured to the anteroventral border of the postorbital. The width was the widest distance between the two parallel bor- ders at riglit angles to the length. Measurements of the jugal bones are presented in Table 18. The smallest ratio (longest, narrowest bone) of length to width is found in Ctenosaura (.136), Sauromalus (.141), and Oplurus (.148). The largest ratios (short- est, widest bones) are found in Amblyrhynchus (.207), Conolophus (.195), and Cyclura (.191)' The posterior extension of the jugal may extend so far posteriorly as to touch or overlap the squamosal, thereby completely covering the ventral border of the postorbital bone. Such a condition exists in Conolo- phus, DIpsosaurus, Iguana, and Oplurus. In the other genera the jugals and squamosals do not touch. TABLE 18 JUGAL BONES Length Width Width-Length Ratio Genus Mm. Mean Max. Mm Mean Max, Min. Mean Max. Amhlvrhynthus I8.I-2I.O-24.7 3,4. 4,3- 5.2 .169-,207.,236 Brachylophus 12.5-13.7-15.9 2,0- 2,7- 3.5 .J60-.n2-,220 Chalarodon 6.1- 6.4- 6.8 1,0- 1,1- 1,2 ,163-.I69-,176 Conolophus ^ 26.2-34. 2-38.2 5.4- 6,6- 7,2 188-195-, 206 Cwnosatira 14.0-19.2-27.3 1.6. 2.6- 3.7 ,1]4-,I36.,155 Cvihira 24,0-38.5-50.8 3.1- 7.7- 1.8 .129.,l9l-,265 Dtpsosaums n. 6-12. 1-12.8 1.7- 1.9. 2.2 ,141..16|.,I80 Iguana 20,0-26.8-31.9 3.2- 4,2- 5.0 ,14|.,I56-,I67 Oplurus 9,4-10.7-13.9 1.3- 1.6- 2.1 .139-.148-,159 Sauromalus 11,9-15,9-24.1 1.3- 2.4- 3.4 ,109-.141- 196 Jugal Jugal (Figs. 1, 2, 3, 4, 5 and 6) forms the ventral border of the orbit and a small part of the supratem- poral arch. Anteriorly it attaches to the maxillae, ven- trally to the lacrimal and palatine bones, and medially Parietal Parietal (Figs. 3, 4, 5 and 6) forms the roof of the posterior part of the skull. It articulates with the frontal and postfrontal anteriorly and with the post- orbital posteriorly and ventrally. Posteriorly the par- ietal overlays the supratemporal and articulates with the exoccipitals and supraoccipital. The anterior two thirds of the dorsal surface of the parietal gives origin to the pseudotemporalis superfi- ciatus muscle, and the posterior one-third gives origin to the adductor externus medius. The posterior tip of the supratemporal process supplies the origin for a part of the adductor externus profundus muscle. Laterally the parietal bone gives origin to the levator pterygoideus. The posterior border of the parietal provides attachment to the origin of the depressor mandibularis and the insertion of the spinus dorsi, longissimus dorsi, and episternocleidomastoideus muscles. Measurements were taken of two areas of the par- ietal bone. The anterior two-thirds of the bone was BIOLOGICAL SERIES. VOL. 12, NO. 3 EVOLUTION OF THE IGUANINE LIZARDS 17 subjected to length-width measurements, with the length being the distance along the midline, from the anterior suture with the frontal to the suture between the parietal and the supraoccipital. The width of the parietal was considered as the distance between the two most anterolateral projections of the bone where they sutured with the postorbital and postfrontals. The measurements are presented in Table 19. The greatest length width ratio (shortest, widest bone) is found in Conolophus (.751) while Dipsosaurus (.431), and Brachylopfms (.448) possess the smallest ratio (longest, narrowest bones). The second portion of the parietal to be measured was the wings or posterior dorsolateral projections of the bone that sutured with the supratemporal, squa- mosal, and articulated with the quadrate. Tiie length of the parietal wings is the diagonal distance from the anterolateral portion of the parietal bone to the opposite posterior tip of the parietal wing. The width is the distance between the most posterolateral sur- face of the two wings. The parietal wing ratios are summarized in Table 20 and show the greatest length width ratios (shortest, widest bones) to be possessed by Dipsosaurus (.945) and Saurumalus (.926). The lowest ratios (longest, narrowest bones) are those of Brachylophus (.765) and Ctenosaura (.781). Supratemporal Supratemporal provides support for the postero- TABLE 19 PARIETAL BONES Genus Length Width Width-Length Ratio Mm. Meun Mj\ Min. Mean Max Min. Mean Ma\. Amhlvrhv'ulms 9.9.12.2-16.1 19.4-24, 2-29,2 .444-.502-,S51 Brachylophus 5.9- 6.5- 7.4 12.8-14 7-18.7 .395-,448-,496 Chalarodon 2.6- 2.8- i.] 4.9- 5.2- 5.5 ,S30-,550-,563 Conolophus 16.3-24.9-29.3 24.7-32,5-37,1 ,659-,7Sl-,789 Ctenosaura 7.|. 9.0-11,0 11,6-13,4-21,0 ,522-552-612 Cvclura 10.3-19.2-32.9 18.1-28.8-41.7 .515-.640-,788 Dipsosaurus 3,4- 3,9- 4.3 8.1- 9,1- 9.7 ,4l9-,431-.443 Iguana 10.9-12.8-14.7 18,4-21.8-24,5 566-,585-.60O OpUirus 3.9- 4.4- 5.5 7,8- 8,7-11.3 .486-5 10-.555 Sauromalus 5.0- 7.6-10.0 10.1-13.2-19.3 .495-.566-.646 TABLE 20 PARIETAL WINGS lateral angle of the parietal bone. Posteriorly, it articulates with the exoccipital, squamosal, and the quadrate. On the medial border of the anterior part is a par- tial origin for the adductor mandibularis externus profundus muscle and on the lateral border is the origin of the adductor mandibularis externus medius muscle. Because of the difficulties in measuring, this bone has not been studied in detail. Postorbital Postorbital (Figs. 1, 2, 3, 4, 5 and 6) forms part of the posterior rim of the orbit. The dorsal part is sutured anteriorly to the postfrontal and posteriorly to the parietal bone. The anteroventral border articu- lates with the jugal and the posteroventral border of the squamosal. The ventral free border of the postorbital gives origin to the anterior half of the levator angularis oris muscle, whereas the medial surface of the posterior half gives origin to the adductor mandibularis ex- ternus superficialis muscle. The length of the postorbital was measured as the greatest distance between anteroventral and post- eroventral projections. The width was considered as the distance from the ventral border to the tip of the dorsal projection where it sutured with the parietal and postfrontal bones. Table 21 shows that the great- est length width ratio (shortest, widest bones) is found in Iguana (.886), Amblyrhynchus (.882), and Conolophus (.876). The lowest ratio (longest, narrow- est bones) is possessed by Sauromalus (.556) and Cyclura (.585). The postorbital bone is roughly triangular in shape in all genera. Its ventral border has been discussed in the description of the jugal. This border is completely covered by the squamosal and jugal bones in Con- olophus, Dipsosaurus, Iguana, and Oplurus. In the other genera the central part of the ventral border is free. In Amblyrhynchus the anterior face of the dorsal process is enlarged and contributes to the knoblike structure of the postfrontal. The postorbital of Cyclura appears normal and does not contribute to the postfrontal knob over the orbit. TABLE 21 POSTORBITAL BONES Length Width Width-Length Ratio Length Width Width-Length Ratio Genus Genus Min. Mean Max Mm. Mean Max, Min, Mean Max, Min. Mean Max Mm. Mean Max Mm. Mean Max. Amblyrhynchus 28.1-33,1-38,1 24.4-29,1-33.5 .868-.878-,891 Amblyrhynchus 9.1-13.5-17.5 8.5-11.8-15.3 .850-.882-.934 Brachylophus 13.4-14.3-16,1 17,3-18,3-20,5 .734-.765-785 Brachylophus 5.5- 6.4- 7.8 4,|. 4,9- 6,0 ,745-.771-,80O Chalarodon 7,0- 7.4- 7,7 6.3- 6,5- 6.7 ,840-.880-.957 Chalarodon 3.1- 3.3- 3.7 2,4- 2,4- 2,6 ,702-.734-.774 Conolophus .14. 1-50,8-58,5 27,4-40.6-48.4 .771-.802-.827 Conolophus 16.0-25.7-30.4 14.3-23. 1-27-4 ,824-,876-,901 Ctenosaura 15 7-25.4-30,9 12.5-18.9-25.8 .747-.781-.802 Ctenosaura 8.0-11,3-15.0 4.2- 6.9- 9.1 ,525-,607.,690 Cvclura 25,0-42,4-58,1 l9..1-35,0-48,8 .772-.817-,839 Cvclura 14.0-22.1-35.3 7.5-15.2-21.6 ,535-,585-,6l4 Dipsosaimis 12,6-13,6-14 4 12-4.12 8-13,2 .916-.945-,984 Dipsosaurus 5.4- 6.4- 7.1 4.0- 4.9- 5.9 ,740-.789-835 Iguana 23,2-29.3-33.4 19.6-25, S-28. 7 .814..S70-.925 Iguana 9,8-14.1-16.7 9.0-12.4-15,4 ,842-,886-922 Oplurus 11.0-12.7-18.0 9.0-10.5-15,1 .810-824-833 Oplurus 5.0- 5.8- 8.2 3.6- 4.3- 5,9 ,704-.719-,760 Sauromalus 12.9-19.7-29.0 11.9-19,1-28,0 .897-.926-,980 Sauromalus 7.3-10.0-15.5 3.7- 5.7- 9.2 ,500-,556-,605 n BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Squamosal Squamosal (Figs. 1, 2, 3, 4, 5 and 6) is attached to the postorbital bone on the posterolateral border of the skull. The expanded posterior part of the squ- amosal is attached to the dorsal surface of the supra- temporal and the quadrate. The lateral surface of the squamosal provides an area of origin for the adductor mandibularis externus superficialis and part of the levator angularis oris muscle. The medial surface gives origin to the ad- ductor mandibularis externus medius muscle. The length of the squamosal was measured as the distance between the most anterior and the posterior extremities of the bone. The width was the greatest distance between the parallel borders on an axis at right angles to the length. These measurements are presented in table 22 and show the greatest ratio (shortest, widest bones) to be found in Ambly- rhynchus (.736). The smallest ratio (longest, narrow- est bones) occurs in Chalarodon (.063). The shape of the squamosal bone differs not only in size but in shape as well. The posterior projection of the bone has a dorsal and ventral hooklike pro- jection in Chalarodon and Opiums. Those of Opiums are not as pronounced as those in Chalarodon. The posterior portion of the bone in other genera is swollen but the projections are in the forms of small triangular processes rather than curving hooks as in Chalarodon and Opiums. The greatest development of these triangular projections is found in Ambly- rhyiiclnis. Conolophus. Ctcnosaura. Cyciura. Iguana, and Sauromalus. The squamosals take the form of a long split in Dipsosaums and Brachylophus. TABLE 22 SQUAMOSAL BONES Length Width Wttith-Length Ratio Mm. Mean Max Mm, Mean Max. Mm. Moan Max Amhlvrhynchiis 7,1.10.2-12.9 5,7- 7.4- 9.5 .606-.736-.802 Brachylophus 6.1- 7.1- 8.5 2.0- 2.2- 2.6 ,280-,304-,327 Chalarodon 3.0- 3.1- 3.S 0.2- 0.2- 0.2 ,057-06 3-, 066 Conohphu s I3.0.20.6-24.3 5,2-10,0-13.4 ,400-,476-,55 1 Ctenosaura 5.9- 9.1-13.0 1.3- 2,6- 5,2 ,202-,234-,268 Cyciura 9.8-18,1-23.1 2,4- 4,9- 8,0 ,244-,297.,346 Dipwsaimis 60- 6,4- 7-1 1,5- 1,5. 1.7 ,245-,268-,-309 Iguana 9,8-1 1,2-12,2 3.0- 4,1- 5,0 , 306-369-434 Opiums 4.1- 5.4- 8.6 1.3. 1,5- 2.2 ,244-,296-,369 Sauromalus 4.6- 7.9-10.1 1.0- 2,4- 4,3 .217-299-373 Quadrate Quadrate (Figs. 1, 2, 3, 4, 5 and 6) is found at the posterolateral angle of the skull where it provides the articulating area between skull and lower jaw. It also forms the seal of the middle ear and attachment for the adductor musculature and tympanic membrane. Mesially the quadrate is attached to the parietal, dorsally to the supratemporal, and laterally lo the squamosal bone. Other parts of its dorsal surface provide for the origin of the adductor mandibularis superficialis muscles. The medial half and its crest serve as origin for the adductor mandibularis poste- rior muscle. The posterior end of this area attaches to the prootic bone and the quadrate process of the pterygoid. The tympanic crest serves as a place of attachment for the tympanic membrane and the origin of the adductor mandibularis externus superficialis muscle. The length of the quadrate is the distance from the dorsal border of the bone where it attaches to the squamosal and the ventral extremity of the condyle of articulation with the articular bone of the lower jaw. Width is the greatest distance between medial and lateral borders of the bone. Table 23 summarizes the quadrate measurements. Quadrates with the great- est length-width ratios (shortest, widest bones) are found in Sauromalus (.737). The smallest quadrate ratio (longest, narrowest bone) is that oi Dipsosaums (.5 31), Iguana (.537), and Brachylophus (.546). TABLE 13, QUADRATE BONES Genus Length Width Width-Length Ratio Min. Mean Max Mm, Mean Max, Mm. Mean Max. Amhlvrh\'nclntl 9,8-11,6-13.5 7,0- 8,0- 9,3 .614-.690-,718 Brachylophus 7,0- 7.4- 8.0 4,1- 4.3- 4.5 .537-.549-,642 ClmlaroUori 2.7. 2.9- 3.2 1.8- 1.8- 1.9 ,593-,628-,666 Conolophus 13.4-19.7-24.9 8,3-13.7-17.0 .619-.693-.774 ClenosauKa 7.0- 8.9-11.0 4.4- 5.8- 8.1 .625-.6S3-.736 Cvilura 10.0-17.9-24.2 5,0- 8.9-15.0 .500-.557-.615 Dipsosaums 6.0- 6.2- 6.8 3,1- 3,3- 3,7 ,5l6-.53l-.544 Iguana 10.5-12.7-13.6 4.9- 6.9- 7.9 .457-.537-.580 Opiums 4.0- 4.7- 6.7 2.4- 3.1- 4,1 .600-,662-,731 Sauromalus 5.0- 7.0-10.0 3.2- 5,2- 7.1 ,640-.737-.887 Epipterygoid Epiterygoid extends between the parietal and the pterygoid. Its dorsal tip appears to be held in place by the origin of the pseudotemporalis superficialis mus- cle. The upper one-third of this bone serves as the origin for the pseudotemporalis superficialis muscle. The ventral two-thirds serves as origin for the pseudotemporalis profundus muscle. This bone was not studied in detail. Supratemporal fossa Supratemporal fossa (Figs. 3 and 4) is the large cavity that dominates each half of the dorsal posterior third of the skull. It is bordered medially, posteromedially, and anteromedially by the parietal bone, anterolaterally by the postorbital, and postero- laterally by the squamosal. Length and width measurements were taken of the supratemporal fossa, with length being the inside dis- tance on the longest axis and width being the inside distance on the longest axis at right angles to the length. Table 24 shows the largest length-width ratios BIOLOGICAL SERIES. VOL. 12, NO. 3 EVOLUTION OF THE IGUANINE LIZARDS (shortest, widest opening) are possessed by Dipso- saurus (.647), Sauronwlus (.620), Amblyrhynchm (.616), and Conolophus (.609). The smallest ratio (longest, narrowest opening) is found in ChalaroJon (.443). TABLE 24 SUPRATEMPORAL FOSSA Length Width Width-Length Ratio Min. Meiin Max. Min. Mean Max. Min. Mean Max. Amblvrhynihu>i Brai hylophus Chalarodon Conolophus Ctcnosaura Cychtra Dtpsosaurus Iguana Opiums Sauromalus 12.5-15.2-18.3 8.0- 9.4-12.1 3.4- 3.6- 4.3 18.0-28.8-34.0 8.1-1LO.I3.9 12.0-21.2-28.0 4.2- 5.0- 5.7 10.9-14.2-16.8 4.2- 5.3- 8.0 5.0. 8.0.12.6 6.3- 9.4-1 1.9 4.3- S.4- 6.9 1.5- 1.6- 1.9 12.0-17.4-21.2 4.9- 8.8- 9.1 5.1-11.3-19.1 2.6- 3.2- 3.7 4.6- 7.1- 8.3 2.1- 2.8- 4.1 2.4- 5.0- 8.1 504-6 16--7S1 514-.577..637 .428-44 3-462 ,560-.609-.666 .S89-,615-.654 .425-,515-.6«2 .619-.647-.687 .422..499.,567 .500-.524-.586 .48O-.620-.672 Orbit Orbit (Figs. 3 and 4) is the dominate lateral cavity of the skull and in life is the area where the eye is located. The orbit is basically circular and is bordered dorsally by the frontal; anteriorly by the prefrontals, lacrimals and jugals. ventrally by the jugal and poste- riorly by the postorbital and the postfrontal. The length of the orbit was measured as the great- est distance between lacrimal and postorbital. The width was the greatest distance between jugal and frontal bones. These relationships are expressed in Table 25 which shows the greatest length-width ratios (most circular opening) to be found in Coiioloplius (.969) and the smallest ratio (most eliptical opening) in Chalarodon ( .682). TABLE 25 ORBIT Length Width Width-Length Ratio Genus Min. Mean Max. Min. Mean Max. Min. Mean Max. Amblvrhvnchus 12.4-14.3-16.3 10.9-15,8.19.5 .802-.827-.866 Brachylophus 11.3-11.9-13.2 9.0- 9.7-10.9 .796-.811..825 Chalarodon 5.2- 6.2- 7.3 3.9- 4.2- 4.5 .61 1-682-769 Conolophus 17.8-23.9-26.6 17.2-23.3-26.9 .958-969-988 Ctenosaura 12.1-15.3-18.3 7.6-12.0-14.1 .770-782-793 Cyclura 16.8-25.6-33.1 12.8-20.7-27.5 .760-803-830 Dtpsosaurus 8.6- 9.5-10I 7.0- 7,5. 7.9 .770-.790..813 Iguana 19.0-21.3-24.9 15.3-17.5-19.3 .775..825..867 Oplurus 7.6- 8.3-12.1 5.9- 7.0- 9.2 .760-.795-.839 Sauromalus 9.6.13,1-17.9 7.1- 9.9-13.5 .711-.741-.776 Fenestra exonarina Fenestra exonarina (Figs. 3, 4, 5 and 6) is the paired prominent opening on the dorsal rostrum of the skull. It is bordered medially and anteroventrally by the premaxilla, laterally by the maxilla, and posterodorsally by the nasals. In life the fenestra exonarina provides the opening for the external nares. The length of the fenestra exonarina was con- sidered to be the internal distance between the suture of the lateral projection of the premaxilla and the maxilla and the suture between the nasal and maxilla. The width was considered to be the greatest inside distance between the lateral border of the premaxilla and the anterior border of the maxilla. Table 26 re- presents a tabulation of these values and shows the greatest ratio (most circular opening) to exist in Brachylophus (.872) and the lowest ratio (most elip- tical opening) to be found in Cyclura (.443). The lower jaw consists of two paired rami which unite anteriorly in a mental symphysis and each artic- ulates with the quadrate of the skull posteriorly. The dentary of each ramus bears a single row of pleuro- dont teeth, whereas the remaining bones (articular, surangular, angular, splenial and coronoid) are eden- tate. TABLE 26 FENESTRA EXONARINA Genus Length Width Width-Lensilh Ratio Min. Mean Max. Min, Mean Max. Min, Mean Max, Amblvrhvnchus 8.1- 9.1-10.3 5.8- 7.0- 8.0 .716..764..817 Brach vlophus 4.0- 4.2- 4.8 3.3- 3,7- 4,0 .804-.872-.951 Chalarodon 1.4- 1.5- 1.8 1,0- 1,0- 1,1 .611-.664-.714 Conolophus 11.7-15.4-17.5 8,5-11,9-13,6 .726-.767-.803 Ctenosaura 4,1. 5.5- 6.8 3,0- 4.1- 5.0 .700-.743-.806 Cvclura 7,3-20,4-32,1 5.3- 9.0-14.5 .428-.443-.452 Dipstisaunis 3,2- 3,8- 4,2 2.0- 2,3- 2,5 .571..S90-.606 Iguana 8.2-11.7-14.3 5,5- 7.8- 8.7 .608-.669-.714 Opiums 1.9- 2.5- 4,2 1.4- 1.8- 3.1 .700-7 18-.738 Saurotnatus 3,0- 4.6- 6,8 2.1- 4.1- 5.0 .600-.75I-.833 Dentary Dentary (Figs. 5, 6 and 7) is the largest bone of the lower jaw and bears teeth on its dorsomedial border. At its posterior margins the dentary inter- digitates with the coronoid, splenial, angular, artic- ular, and surangular. The ventral border provides the origin for the anterior fibers of the mandibulo- hyoideus I and HI and the genioglossus muscles. Measurements taken on the dentary included the length as the distance from the anterior tip of the bone to the posterior most projection on the lateral surface of the mandible. The width of the dentary was that vertical distance between the top and the bottom of the mandible immediately in front of the coronoid. An examination of Table 27 reveals that the largest length-width ratio (smallest, widest bone) for the dentary is possessed by Aniblyrhynchus (.297) and Conolophus (.282). The smallest ratio (longest, narrowest bone) is that of Oplurus (.147) and Chalarodon (.174). The relationship between the posterolateral border of the dentary, the coronoid and the surangular differs from genus to genus. In Chalarodon and 20 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN Opiums the dentary is not overlapped by the coro- noid on its posterodorsal surface. In Brachylophus, Ctenosaura, Dipsosaunis, Iguana and Sauromalus, the dentary is overlapped dorsally by the coronoid and the ventral border of the coronoid and its suture with the dentary is parallel to the ventral border of the mandible. The posterior suture of the dentary in Brachylophus, Ctenosaura. Iguana and Sauromalus is concave in nature. In Dipsosaurus this suture is con- vex, Amblyrhynchus. Conolophus and Cyclura have complex rounded or slanting suture between the ventral border of the overlapping coronoid and its suture with the dentary. The posterolateral suture in these genera is complex with two posteriorly pointing triangular projections being present in Ambly- rhynchus and Conolophus. Cyclura possesses a smooth gently curving concave suture. TABLE 27 DENTARY BONES Length Widlh Widlh-Length Ralio Min. Mean Max Mm. Mean Max. Mm. Mean Max, Amblyrhynchus 20.3-28. 3-34. 3 7,5- 8.3-10.2 ,271-,297-.320 Brachylophus 19.8-21.4-23.1 4.0- 4.5- 5.0 .202-.209-,216 Chalarodon 8.1- 9.2-10.0 1.3- 1.5- 2,0 .l42-.n4-.209 Conolophus 40.4-51.2-55.6 8.7-14.7-17.4 .215-.282-.319 Ctenosaura 19.2-26.7-35.6 3.4- 5.2- 7.5 .171-.190-.210 Cyclura 29.5-51.3-72.3 6.5-10,9-15.0 .202-.214-.228 Dipsosaurus 12.9-14.7-15.9 3,1- 3,6- 3.9 .240-.243-.246 Iguana 30.0-40.3-47.5 6.7- 9.3-10.8 .213-.22S-.239 Oplurus 13.2-17.1-24.0 2.1. 2.4- 3.1 .129-.147-.159 Sauromalus 13.6-20.0-28.4 2.5- 3.6- 5.5 .143-.181-.203 Articular Articular (Figs, 5, 6 and 7) is that part of the man- dibles which articulates with the quadrate of the skull. This bone with its retroarticular process serves as the point for insertion for the depressor mandib- ularis, intermandibularis posterior, fibers of the ptery- gomandibularis, and adductor mandibularis externus muscles. The medial border is sutured to the splenial bone. The length of the articular bone was measured from its most anterior projection on the medial surface where it sutured to the coronoid and splenial to the most posterior tip of the retroarticular process. The width was taken from the most ventromedial projection of the angular process to the opposite border of the articular bone where it sutured with the surangular on the mandible's lateral surface. These measurements are summarized in Table 28 which shows the greatest length-width ratio (shortest, widest bones) to be found in Conolophus (.390) and the lowest ratio is possessed by Chalarodon (.200). The angular process of the articular bone differs in shape and size from genus to genus. This projection was also subjected to length-width measurements. The length was taken as the greatest length of the mandible and was contrasted with the width of the articular which in part is a result of the size of the angular process. The ratio between those measure- ments is expressed in Table 29 which shows Con- olophus (.219) to have the greatest ratio (shortest, widest bones), while the smallest ratios (longest, narrowest bones) are those of Chalarodon (,105) and Oplurus {.[20). As previously indicated, the shape of the angular process (Figure 7) differs from genus to genus. The projection which extends medially from the posterior ventromedial surface of the articular bone is either a small rounded bump as in Amblyrhynchus and Iguana, or a pointed spinelike anteriorly pointing pro- jection as in the remaining genera. The smallest pro- jection is that of Amblyrhynchus. Of the pointed pro- jections, that of Cyclura is the most massive and pro- nounced, whereas the projection in Conolophus is nearly as large. The angular process of Ctenosaura and Sauromalus is less massive and projecting than that of Cyclura and Conolophus. It is similar in shape but less pointed in the last two genera. In Brachylophus, Chalarodon, Dipsosaurus and Oplurus, the angular projection is more triangular shaped than hooklike. In all four genera the anterior face terminates as a sharp spine. The process of Brachylophus and Oplurus is more sharply curved and hooklike than that of Chala- rodon and Dipsosaurus. TABLE 28 ARTICULAR BONES Length Width-Length Ratio Min. Mean Max. Min. Mean Max. ilin. Mean Max. Amblvrhvnchus 18.4-25 (1-30,0 5 9- 8.1-10.8 258-320-360 Brachylophus 16.0-17,4-18,4 4,3. 5.3- 6.4 240-,309.400 Chalarodon 6.1- 6.3- 6.7 1.2- 1.2- 1.4 193-200-208 Conolophus 34.2-52.8-61.5 11,7.21.1-27.2 342-.394-.457 Ctenosaura 14,1.2I.4-21.2 4,6- 6,4-10,6 238-.297-.340 Cvclura 24.2-42.4-57.8 4.7-12.0-20.3 182-,264-.351 Dipsosaurus 8.9-1 1.3-12.6 3.0- 3.6- 4.1 298-320-337 Iguana 23.4-31.9-38.7 7.4- 8.6- 9.1 227-.27|.,316 Oplurus 8,3-11.3-15.8 1.9- 3.0- 4,9 228-.257-.310 Sauromalus 14.8-20.7-25.5 2.8- 5.5- 9.3 185-.253-,325 TABLE 29 ANGULAR PROCESS Length Width Width-Length Ralio Mm, Mean Max. Min. Mean Max. Min. Mean Max. Amblvrhvnchus 35.0-48.4- 57.1 6.4- 9.5-11.5 182-.196-,203 Brachylophus 32.2-33.6- 35.6 4.2- 5.4- 6.6 130-1 5 7, 185 Chalarodon 12.0-I2.6- 13.4 1.0- 1.3- 1.8 08 1-1 05. 134 Conoluplui s 66.6-95. 1-107.6 11.6-21.3-27.2 174-219-. 256 Ctenosaura 30,2-43. 1- 60.4 3.9- 6.3-10.6 12.1-.141-.174 Cvclura 48.7.84.7.117.3 7.9-14.5-21.1 162-.169-.179 Dip\nsauni\ 19,9-23.2- 25.4 3.5- 4.1- 4.4 169-1 74-. 175 Iguana 50.0-65.4- 77.2 7.5- 8.6- 9.2 112-134-150 Oplur^is 18.2-24.3-34.7 1.9- 3.0- 4.9 104-. 120-141 Sauromalus 22.4-33.4-43.1 2.8- 5,5- 9.3 113- 1 38-. 167 BIOLOGICAL SERIES. VOL. 1 2, NO. 3 EVOLUTION OF THE IGUANINE LIZARDS 21 Surangular Surangular (Figs. 5, 6 and 7) forms the lateral wall of the posterior third of mandible. The dorsal border serves as the area of insertion for the adductor man- dibularis externus muscle, and the intermandibularis posterior muscle inserts on its lateral surface. The surangular's length is taken as the longest anterior-posterior axis on the lateral surface of the mandible. The width is considered to be the longest dorsal-ventral axis in the area of the anterior sutures with the dentary and coronoid on the mandible's lateral surface. Table 30 indicates the largest length- width ratio (shortest, widest bone) is found in Amblyrhynchiis (.425) and the smallest ratio (longest, narrowest bone) in Sauromalus (.270) and Iguana {,.21^). TABLE 30 SURANGULAR BONES TABLE 31 SPLENIAL BONES U'litilli Width-Lcnglh Ratio Min. MiMri Mj\. Min. Mean Ma\. Min. Mean Max. AmhtrrlniK lni\ ILW. II, 1,-11.6 5.8- 6.9- 8.9 .365-.425-.487 Bracliyln[}hii\