HARVARIT'UNIVERSITY Library of the Museum of Comparative Zoology The Great Basin Naturalist VOLUME 34, 1974 Editor: Stkphen L. Wood Published at Brigham Young University, by Brigham Young University TABLE OF CONTENTS Volume 34 Number 1 — March 31, 1974 Numeric analysis of the lizard genus Sceloporus with spe- cial reference to cranial osteology. Kenneth R. Larsen and Wilmer W. Tanner 1 The generic name of the North American musk turtles. Hobart M. Smith and Kenneth R. Larsen 42 Studies on the tolerance of aquatic insects to low oxygen concentrations. Arden R. Gaufin, Robert Clubb, and Robert Newell 45 Substrate color matching in the grasshopper, Circotettix rabula (Orthoptera: Acrididae). George W. Cox and Darla G. Cox 60 Effects of chemicals on the germination of pollen grains of Torenia asiatica Linn. E. M. V. Nambudiri and M. K. Thomas 71 An unusual spider bite. D. M. Allred 82 Number 2 — June 30, 1974 The species of Calineuria and Doroneuria (Plecoptera: Perlidae). Bill P. Stark and Arden R. Gaufin 83 The name of the Baja California Cape wormsnake. Hobart M. Smith and Kenneth R. Larsen 94 Contributions to the concepts of reproductive cycles and the systematics of the scalaris group of the lizard genus Sceloporus. Hobart M. Smith and William P. Hall 97 Growth and development of the western harvest mouse, Reithrodontomys niegalotis megalotis. Gary H. Rich- ins, H. Duane Smith, and Clive D. Jorgensen 105 Body size, body composition, and behavior of juvenile Belding ground squirrels. Martin L. Morton, Cathe- rine S. Maxwell, and Charles E. Wade 121 New species of American Corthylini (Coleoptera: Scolyti- dae). Stephen L. Wood 135 The anatomy and taxonomy of Vnnclevea ( Asteraceae) . Loran C. Anderson and Phillip S. Weberg 151 A revision of the Echidiocarya section of Plagiobothrys (Boraginaceae) . Larry C. Higgins 161 Number 3 — Sept. 30, 1974 Studies on the biology and ecolog}- of the northern scorpi- C on, Paruroctonus boreus (Girard). Gregor}' Ira Tourtlotte 167 ) Notes on three varieties of Astragalus lentiginosus (Legu- minosae). Carol S. Schoener 180 New species of American Corthylus (Coleoptera: Scolyti- dae). Stephen L. Wood 181 Notes on Viscainopelmatus davewerneri (Orthoptera: Gryllacridadae, vStenopelmatinae) from the Viscaino Desert, Baja CaHfornia Sur, Mexico. Ernest R. Tink- ham, Clark R. Mahrdt, and Benjamin H. Banta 203 The nustrina group of the genus Microvelia (Hemiptera: Veliidae). John T. Polhemus 207 Additional records of Coleoptera collected at the Nevada Test Site, Mercury, Nevada. Vasco M. Tanner and Wilmer W. Tanner 218 A summary of biological investigations concerning the Great ^Salt Lake, Utah (1861-1973). Doyle W. Stephens 221 A new variety of Penstemon thurberi (Scrophulariaceae). James L. Reveal and Janice C. Beatley 230 A new sceloporine lizard from Oaxaca, Mexico. Marlene M. Dasmann and Hobart M. Smith 231 Range extension of the long-nosed snake, Rhinochcilus 1. lecontei, into east-central Utah. William L. Grogan, Jr., and Wilmer W. Tanner 238 Description of a Phyllorhynchus from Cerrovalo Island, Gulf of California, Mexico. Arnold L. Powers and Benjamin H. Banta - 241 Two new varieties of Eriogonum (Polygonaceae) from the Intermountain region. James L. Reveal 245 Number 4 — December 31, 1974 Field behavior and seasonal acti^ itv of the rodent bot fly, Cuterebra tenebrosa^ in central Washington (Diptera: Cuterebridae) . Craig R. Baird 247 Notes on the spotted bat {Euderma maculatum) from southwest Utah. Richard M. Poche and Geoffrey L. Bailie 254 The significance of scale characters in evaluation of the lizard genera Gerrhonotits, Elgaria, and Barisia. James W. Waddick and ITobart M. Smith 257 Revegetation of gopher mounds on aspen range in Utah. W. T. McDonough 267 Observation on voice in the western collared lizard, Crotaphytus collaris bicinctores. Nathan M. Smith 276 New synonymy and records of American bark beetles (Coleoptera: Scolytidae). Stephen L. Wood 277 Description of new species of Milodcres Casey, with com- ments on other species in the genus (Coleoptera: Curculionidae) . Vasco M. Tanner 291 Revision of the plant genus Geranium in Utah. Glen T. Nebeker 297 Utah plant novelties in Astragalus and Yucca. Stanley L. Welsh : 305 A new combination in Penstemon (Scrophulariaceae). James L. Reveal - 311 Dominance relationships of the dark kangaroo mouse {Mi- crodipodops rnegacephalus) and the little pocket mouse {Perognathus longimembris) in captivity. Andrew R. Blaustein and Arthur C. Risser, Jr 312 List of insect type specimens in the entomological collec- tion of the Brigham Young University, Provo, Utah, No. V. Vasco M. Tanner 317 Volume j4, ino. i March 31, 1974 The Great Basin mmwjm Brioham YnrjNf; TJ? GREAT BASIN NATURALIST Editor: Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah Editorial Board: Stanley L. Welsh, Botany, Chairman; Wilmer W. Tanner, Zoology; Joseph R. Murdock, Botany; Vemon J. Tipton, Zoology; Ferron L. Andersen, Zoology Ex officio Editorial Board Members-. A. Lester Allen, Dean, College of Biological and Agricul- tural Sciences Ernest L. Olson, Director, University Press, University Editor The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been continuously published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only original, previously unpublished manuscripts pertain- ing to ^e biological natural history of the Great Basin and western North America will be accepted. Manuscripts are subject to the ap- proval of the editor. SuBscBiPTioN: The annual subscription is $5.00 (outside the United States $5.50). The price for single numbers is $2.00 each. All matters pertaining to toe purchase of subscriptions and back numbers should be directed to Brigham Young University Press, Publication Sales, 205 UPB, Provo, Utah 84601. libraries or other organizations interested in obtaining this journal through a contin- uing exchange of scholarly publications should contact the Brigham Yoimg University Exchange Librarian, The Library, Provo, Utah 84601. Manuscripts: AU manuscripts and other copy for the Great Basin Naturalist should be addressed to the editor. Contributors should consult the instructions printed on the back cover of a recent number. The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University Volume 34 March 31, 1974 No. 1 NUMERIC ANALYSIS OF THE LIZARD GENUS SCELOPORUS WITH SPECIAL REFERENCE TO CRANIAL OSTEOLOGY Kenneth R. Larsen^-^ and Wilmer W. Tanner^ Abstract. — Numerical statistical methods were used to analyze the species in the genus Sceloporus using cranial osteology, external meristic and numeric characters, karyology, display behavior, and geographic distribution. A new classification for the genus is proposed with three major branches or groups. Group I contains 7 species in 3 species groups. Group II contains approxi- mately 19 species in 5 species groups. Group III contains approximately 32 species in 5 species groups. This classification is supported by the cluster analysis of several different sets of data. Cranial osteology, zoogeography, behavior, and karyology are shown to be taxonomically significant as nvuneric characters. Step- wise discriminate analysis shows that this classification of the species of Sceloporus into 3 major groups and 13 species groups is significant at the .999 confidence level: It is concluded that the 3 major groups should be given taxonomic recogni- tion. Cope (1900) stated, "The distinction of many of the species of this genus [Sceloporus] is not accomplished without difficulty. I recommend it as an excellent piece de resistance for those persons who do not believe in the doctrine of derivation of species." This statement was endorsed by Hobart Smith (1938:548-49): Sceloporus is one of the most nearly ideal of living genera of reptiles for the study of speciation and related phenomena. The characteristics which it possesses and which are essential to an ideal genus for such studies are: 1 . A large number of living forms. . . . 2. Prolificity. Where Sceloporus occurs, usually it is the most common of all reptiles, or for that matter, of all vertebrates. 3. A large range, entirely contiguous. The genus occupies practically all of the United States, and occurs as far south as Panama. 4. Great adaptability. Species in this genus have adapted themselves to considerable range of elevation — from below sea level (Death Valley) to about 13,500 feet above sea level. They occur in almost every conceivable terrestrial habitat— deserts, sand dunes, forests, on rocks, trees, or ground in grassy plains or heavy brush, and even on houses, fences and other man-made structures. 5. Lack of obvious distinctive specific characters. Subspecies are numerous and species not so well defined as in many other genera of animals, and for this reason relationships may more definitely be postulated. These characters are indicative of a group of relatively recent development. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 'Present address: E.P.O. Biology, University of Colorado, Boulder, Colorado 84302. GREAT BASIN NATURALIST Vol. 34, No. 1 Smith would probably have added a sixth and seventh character- istic if karyological and behavioral information had been available. It is only proper, in consideration of the foregoing, that Scelo- porus should be considered a suitable candidate for the application of recently developed statistical methods. The study here reported was undertaken with several questions in mind: (1) What is the most natural arrangement of species within the genus? (2) Can satisfactory results be obtained with modern statistical methods? (3) Can significant intrageneric taxo- nomic information be obtained from the cranial osteology of Scelop- orus? (4) Will different sets of characters (scale counts, external morphology, karyotypes, behavior, osteology, etc.) produce similar results? (5) Is Sceloporus a single genus? We extend our deepest gratitude to the following persons and institutions for loans, exchanges, and gifts of specimens and for critical assistance in the preparation of the manuscript: Ticul Al- varez (University of Mexico), Charles C. Carpenter and James Richard Purdue (University of Oklahoma), Charles J. Cole (Ameri- can Museum of Natural History, AMNH), Wilham E. Duellman (University of Kansas), William P. Hall (University of Puerto Rico), Hymen Marx (Field Museum of Natural History, FMNH), Hobart M. Smith (University of Colorado), and Ernest E. Williams (Museum of Comparative Zoology, Harvard). We thank the departments of Zoology, Geology, and Computer Science at Brigham Young University (BYU) for use of their facili- ties and for valuable assistance rendered by numerous staff and faculty members. Fig. 1. Platform for constant angle photography of skulls. March 1974 LARSEN, TANNER: SCELOPORUS This Study was supported by a National Defense Education Act title IV fellowship to the senior author from 1966 to 1970. Review of Literature Taxonomy The term Sceloporus was coined by Weigmann (1828:369-70) from the Greek words scelos (leg) and porus (hole). Our translation of Weigmann's original description follows: Furthermore, there is a Mexican genus with many species which is similar to Tropidurus in body shape, head shape, placement of nostrils and ears, formation of teeth, and form and placement of dorsal and caudal scales. But it differs in that it has femoral pores and the dorsals are enlarged. Both genera have a peculiar dermal pocket on the side of the neck. This dermal pocket is crescent shaped in the new genus. It is formed by a fold in the skin Fig. 2. Characters 41 to 50 on dorsal view of skull. 41=A-B 42 = E-H 43 = H-K 44 = D-F 45 = G-I 46 = I-J R-F 48 = L-N 49 = A-B BE MO N-B 50 = BE C-G P-Q F-G BE BE 47 = B-P BE GREAT BASIN NATURALIST Vol. 34, No. 1 and the inner surface is lined with shagreen-like scales. I usually found a popu- lation of 6-legged orange-colored epizoa in the dermal pocket in which case the scales would be missing. His Highness, the Prince of Neuwied observed the same thing in his description of Tropidurus torquatus (Beitrage zur Naturgeschichte Brasiliens I. p. 148). Hernandez has already mentioned 2 of the species of this genus. He reports that the species which can reasonably be considered typical is a crevice-dweller and eats worms. Because of the large femoral pores, I name this genus Sceloporus. The following is a short provisional description of the species. Weigmann (1828) then gave a short description of the genus and six species: torquatus, spinosus, grammicus, pleurostictus , aeneus, and scalaris. In the heading, he provided the common name Stone Lizard, which name he explained in a footnote: "I have chosen this German name (Stone Lizard) because Hernandez says that the common species of this genus are called Tecoixin in Mexi- co. Tecoixin means Saxorum Lacerta [Saxorum lacerta = stone lizard]." KLM N O P Q R Fig. 3. Characters 51 to 56 on dorsal view of skull. 51 =N^ 52=J-L 53 = M-P 54=K-M 55 = D-C 56 = F-G L-Q L-Q L-Q L-Q D-I EH March 1974 LARSEN, TANNER: SCELOPORUS Hobart Smith (1938:547-48) provided an excellent history of the revisions of this genus which is paraphrased as follows: Weigmann (1834) recognized nine s{>ecies — torquatus, formosus, spinosus, horridus, grammicus, microlepidotus, variabilis, aeneus and scalaris. Dumeril and Bibron (1837) recognized 10 species, adding undulatus Latreille. Bocourt (1834) recognized 22 species. Cope (1885) published a synopsis of Sceloporus, in which he recognized 36 species and subspecies. Boulenger (1885) recognized 33 species and subspecies and Gunther (1890) recognized 30 species and listed 7 other described forms without comment as to validity. Boulenger (1897) presented his conclusions with regard to the species of Sceloporus in his revision of the genus and recognized 36 species and sub- species. . . . In the last monograph of the genus is that of Cope (1900) published in 1900, in "The Crocodilians, Lizards, and Snakes of North America." Forty species and subspecies are recognized. Smith (1939:29) added, "Of the 127 names proposed in the ge- nus, I consider 95 valid. These have been segregated into 15 groups of approximately equivalent morphological value." Fig. 4. Characters 57 to 60 on ventral view of skull. 57 = G-E 59 = C-B 58 = E^ 59 = C-B 60 = F-D A-D A-D A-D AD A-D GREAT BASIN NATURALIST Vol. 34, No. 1 Smith and Taylor (1950) provide the following list of groups and species (15 groups, 54 species; in each, the first species is the group name): (1) formosus, malachiticus, asper, stejnegeri, presy- gous, lunaei; (2) spinosus, lundelli, edwardtaylori, melanorhinus, clarki, orcutti, magister, horridus, olivaceus; (3) undulatus, cautus, occidentalis, woodi; (4) graciosus; (5) grammicus, heterolepsis; (6) megalepidurus, pictus; (7) torquatus, serrifer, mucronatus poinsetti, cyanogenys, bulleri, lineolateralis, ornatus, dugesi, jarrovi; (8) variabilis, cozumelae, teapensis, parvus, couchi; (9) merriami; (10) maculosus; (11) chrysostictus; (12) siniferus, squamosus, carinatus, ochoterenai; (13) utiformis; (14) scalar is, jalapae, aeneus, goldmani; (15) pyrocephalus, gadoviae, nelsoni. Later, Smith and Taylor (1966) added four new species to their checklist: macdougalli, shannonorum, subpictus, and virgatus. Hall (1971) increased the number of species in this genus to 61 by recog- Fig. 5. Characters 61 to 66 on ventral view of skull. 61=A-B 62=B-C 63 = CD 64=GJ£ 65 = E^ 66 = (diagonal) A-E A-E A-E A-E I-J 5 (Tangent P) March 1974 LARSEN, TANNER: SCELOPORUS nizing acanthinus and by adding exsul and insignis. Hall (pers. comm.) has called attention to a new species in Baja California and has suggested the elevation of magister zosteromus and orcutti licki to specific rank. He has also proposed that grammicus contains at least six cryptic species. Hobart Smith (pers. comm.) also has a manuscript species. A second manuscript species described by Smith and Larsen is in press. If these new species are included, the total number in this genus would exceed seventy. Osteology. Avery and Tanner (1971) presented a review of lizard osteology to which the reader is referred. On page 6 they stated: In summary the literature dealing with anterior osteology and myology of lizards is scattered and varied. Descriptions of skulls representing almost all families can be found. With the exception of such papers as Camp (1923), McDowell and Bogert (1954), Savage (1958), Etheridge (1964), and Presch (1969), little has been done, utilizing osteology, to analyze the evolutionary lines within families. Of the above listed papers, only Savage, Etheridge, and Presch considered Sceloporine relationships, and none of these reported on species relationships within the genus Sceloporus. Cope (1900:330-31) described the cranial osteolog}-^ of Sceloporus on the basis of two specimens of undulatus and one specimen of spinosus. He described the following 35 characteristics: [1] Premaxillary bone has a long superior spine and is [2] truncate on the palatal face, and [3] has the button-like process. [4] The nostrils are partially vertical, so that the [5] nasals are a little shortened in front. [6] The latter are rather large and are distinct. [7] The frontal is simple and narrow and is [8] strongly grooved on the middle line below. [9] The parietal is short and wide, and [10] is perforated by a large pineal foramen, [11] which touches the A KB c Fig. 6. Characters 67 to 73 on lateral view of skull. 67 = A;^ 68 = B-C 69 = AD 70 = D-E 71=E-F 72 = HI 73 = J-K AG AG AG AG AG D-E L-M 8 GREAT BASIN NATURALIST Vol. 34, No. 1 coronal suture. [12] Parietoquadrate arch distinct. [13] Supraoccipital broadly but loosely attached [14] confluent with exoccipitals. [15] Prefrontals large, not reaching postfrontals above. [16] Lachrymal small and joining jugal. [17] Postf rental a small splint. [18] Postorbital large, extensively in contact with jugal and supra temporal. [19] Paroccipital small. [20] Vomers short, divaricate, and separated by a deep notch behind. [21] Palatine with the vomerine process longer than maxillary; [22] Palatine foramen large. [23] Palatines and ptery- goids well separated from each other on the middle line; [24] ectopterygoid de- flected at its internal extremity. [25] Basipterygoids developed. [26] Quadrate with two conchs [27] the internal the narrower. [28] Presphenoid rudimental; [29] sphenoid and basioccipital coossified; [30] descending lateral processes of the latter strongly developed. [31] The supraforaminal part of the petrosal is very short; [32] the infraforaminal portion is produced beyond it and is nearly hori- zontal in position. [33] The foramen of the eighth nerve is at the bottom of a fossa. [34] Epipterygoid resting on pterygoid much posterior to ectopterygoid and reaching parietal without touching petrosal. [35] Occipital condyle not sub- divided by grooves. (We disagree with the last characteristic as most of our specimens exhibit a conspicuous pair of grooves that subdivide the occipital condyle.) Lundelius (1957) produced the only computerized statistical analysis of Sceloporus cranial osteology to date. On pages 67 and 68, he listed 32 cranial measurements used in his analysis: (1) midline length of premaxillary, (2) midline length of nasal, (3) midline length of frontal, (4) midline length of pineal, (5) midline length of parietal, (6) total length of skull roof from snout to posterior edge of parietal, (7) inter- narial width, (8) width anterior to orbit, (9) interorbital width, (10) anterior width of parietal, (11) width of pineal, (12) interfenestral width, (13) maximum width of teiniK>ral fenestra (diagonal), (14) distance from basicranial tubera to E .... J K N L O M Fig. 7. Characters 74 to 80 on posterior view of skull. 74 = C^ 75 = D-F 76 = F^ 77 = E^ (along diagonal J) 78 = A-B A-B A-B A-B 79= 5 (Tangent P) 80 = 5 (Tangent Q) K-M N-0 March 1974 LARSEN, TANNER: SCELOPORUS basipterygoid process, (15) length of palatine ramus of pterygoid, (16) length of palatine, (17) length of prevomer, (18) length of quadrate ramus of pterygoid, (19) width across basicranial tubera, (20) width across basipterygoid processes, (21) width across posterior ends of maxillaries, (22) width across descending processes of pterygoid, (23) width across anterior part of palate, (24) tooth row width of premaxillaries, (25) length of maxillary, (26) distance from the pos- terior end of maxillary to posterior edge of quadrate, (27) length of quadrate, (28) total width of skull across exoccipitals, (29) length of exoccipital, (30) medial end of exoccipital to lateral edge of foramen magnum, (31) width of foramen magnum, (32) width of occipital condyle. Our analysis utilitzed all the above measurements, or functions of them, with the exception of numbers 4, 11, 13, 14, and 23. Num- bers 4 and 1 1 were omitted because the thin bone around the parie- tal foramen is easily dissolved in bleach and because we suspect that the dimensions of the parietal foramen may be affected by the time spent in bleach during preparation. Number 13 was omitted because it is a diagonal with no definite points of origin. We measured the width of the temporal fenestra at right angles to the midline. Num- bers 14 and 23 were omitted because they are difficult to define on a photograph (see material and methods below). Karyology The karyology of Sceloporus has attracted much interest because of the high level of intrageneric variation. Gorman, Atkins, and Holzinger (1967) published karyotypic data on 15 genera. On page 287 they reviewed a manuscript presented by William P. Hall: MfCAllPIDURUSr MACUIOSUS ' OCHOTEIIfNAI . IfAPINSIS L. COZUMIIAE / SOUAMOSIS -1 SINtFilUS y CAIINATUS ' CH»TSOSTICTUS- MECAlEPIDURUSv PICTUS V- TiAPtNSIS ■' moCEPNAlUS UAMMICUS N MICtOlEPIDOTUS'" ASPE» \ FCMMOSUS h MElANO«HINUS' OUVACEUS . MAGrSTE* - SPINOSUS - EDWAaOTATlO ACANTHINUS^ SALVINI ' lUNDElLI j lUNAEI ^ Fig. 8. Dendrogram produced by Ward's cluster analysis of Smith's (1939) data for Sceloporus. 10 GREAT BASIN NATURALIST Vol. 34, No. 1 Hall (1965) has summarized all available information on iguanid karyo- types. He listed the formula 12 metacentric Macrochromosomes and 24 micro- chromosomes for the following genera: Anolis, Crotaphytus, Dipsosaurus, and Phrynosoma. Hall characterized the genera termed 'sceloporine' ... as having 12 metacentric Macrochromosomes and a reduced number of microchromosomes, ranging from 10 to 22. Hall's data include members of the genera Holbrookia, Callisaurus, Urosaurus, Uta and Sceloporus. Gorman et al. (1967) established that a formula of 12 meta- centric macrochromosomes and 24 microchromosomes is primitive among many lizards. They concluded: "Chromosome loss would be of a specialized, advanced character, and this correlates with the phylogenetic position of the sceloporines." Lowe, Cole, and Patton (1967) proposed that karyotypical evolu- tion can be a matter of Robertsonian fusion, but they did not al- low for Robertsonian fission. Cole (1970, 1971a, 1971b) published the karyotypes of the spinosus group, the pyrocephalus group, and the five monotypic groups (Smith's groups above). He proposed phylogenies of the two polytypic groups and discussed relationships among the others. CHARACTER Fig. 9. Theory of canonical analysis. March 1974 lausen, tannku: sceloporus See page 14 for caption 11 12 GREAT BASIN NATURALIST See page 14 for caption Vol. 34, No. 1 March 1974 LARSEN, TANNER: SCELOPORUS 13 See page 14 for caption 14 GREAT BASIN NATURALIST Vol. 34, No. 1 Fig. 10. Dorsal, ventral, lateral, and posterior views of 13 species of Sceloporus, representing the major groups within the genus: A, Sceloporus gadoviae; B, S. couchi; C, S. maculosus; D, S. grammicus microlepidotus; E, S. pyrocephalus; F, S. scalaris scalaris; G, S. siniferus cupreus; H, 5. variabilis variabilis; I, S. spinous caeruleopunctatus; J, S. formosus formosus; K, S. undulatus elongatus; L, S. jarrovi jarrovi; M, S. torquatus melanogaster . Hall (1970, 1973) has also attempted to establish a phylogeny of Sceloporus with major emphasis on karyology. With almost no disagreement concerning the karyotypes of different species. Hall and Cole have produced quite different phylogenies. Hall accepts fission as well as fusion. The occurrence of fission was shown in Anolis by Webster, Hall, and Williams (1972). Ethology In 1960 Hunsaker showed that different species of Sceloporus have specific display patterns. He showed that females can distin- guish between the display patterns of closely related forms. In one part of his study, Hunsaker offered females a choice of males of closely related species. His results show that the females seemed to discern which male was most similar to their own species. This pre- ference by females could be a valid systematic tool. For example, Hunsaker mixed female jarrovi with males of jarrovi, dugesii, and ornatus. He found that female jarrovi prefer to associate with male dugesii over ornatus 16 to 11 (61 observations): "These data reflect an apparent tendency of a female jarrovi to associate with male dugesii more frequently than with a male ornatus" (p. 67). Hun- saker also found that female jarrovi preferred male dugesii over male jarrovi 22 to 11 (70 observations) and ornatus over jarrovi 47 to 12 (47 observations). Possibly the females were not receptive and preferred to avoid their own species. As a result, it may be con- cluded that ornatus is closer to jarrovi than is dugesi (the ornatus males looked more like jarrovi to the female jarrovi who chose to avoid males). The results do not agree with Smith (1939), and Hun- saker's work is hardly sufficient for systematic conclusions at this point. However, this method may have future prospects. March 1974 LARSEN, TANNER: SCELOPORUS 15 Ss;?,vj'.'>- MocOougolli Mslonorhinus ^ Mucronarui Cvanog*nyi • ufl«aii - irgotus Afp*r Grocioiuc — M*gol*pidur prison 1 1 - variobiiii' T»op«nsi» Cotum«la SiniUrus. uriformis Carinorus ChryiostK Squ MOCUIOSUS — Ochot*r«na« • jalaptM - !=> 0.5 16 32 Fig. 11. Dendrogram generated by external characters (1 to 40). A more promising aspect of lizard behavior is the analysis of "display action patterns." Carpenter (1962) reported on the display action patterns of Uta, Streptosaurus, and Urosaurus and concluded that Urosaurus is a valid genus because its patterns diverge signifi- cantly from those of Uta. Purdue and Carpenter (1972a) compared one species of Petro- saurus, five species of Uta, and five species of Urosaurus to 22 spe- cies of Sceloporus. In their examination of displaying males, they found that the ratio of hip movement to eye movement is a valuable taxonomic character. They have also shown (1972b) that the ratio of shoulder movement to eye movement is a valid species-specific character. Both ratios have been included in our analysis. Hematology Guttman (1970) analyzed the hemoglobin of 12 species of Scelo- porus using gel electrophoresis. His evidence gives ample support to the proposition that relative movement within the gel is indeed de- termined by genetic factors. His data also support the arrangement of cyanogenys and torquatus in one part and jarrovi in the other part of a distinct group. Although some relationships can be shown with electrophoresis, there are problems that make this method suspect. If a heterozy- gous individual produces two bands, which band is representative of 16 GREAT BASIN NATURALIST Vol. 34, No. 1 HOimiDUS OUTUS — CUTPTUS — lUNAEI — OLIVXCEUS ^— EDW/tROTAriORI MEKNORHINUS SINIFEKUS ^^ CARINATUS ^— UriFORMIS ^— CVtNOGENVS ■ULLERI *SfER NETEROLEPIS ' SPINOSUS ^ lUNDELLI ^ IINIOLAIERALIS • *C«NTHINUS M«GISTER ^ VIRG«TUS ^ COZUMElAt • VARIABILIS - SQUAMOSUS SCALARIS ^— OCHOTERLNAE AENEUS ^^— lALAPAE ^-^— PARVUS •^^— NEISONI — ? ^— h ^=^ ^ 2=h ^J 0.5 16 Fig. 12. Dendrogram generated by skull characters (41 to 80). the position of the species? Sometimes the separation between two bands in a single individual is greater than the distance between single bands of widely divergent species. For example, the total range of relative movement reported by Guttman in the gel is from .11 to .50. S. unclulatus and cyanogenys together cover almost the entire range (.16 to .50). Yet they have a nearly identical band {cy- anogenys .30; undulatus .28 to .33). The relative movement of hemoglobin in an electrophoretic gel is obviously not an indication of degree of relationship. Such a number cannot be used as a nu- meric character, and the interpretation of electrophoresis must re- main subjective and qualitative — which does not rule out its value in systematics. It would be a mistake, however, to consider vari- abilis (.16) and merriami (.17) as more closely related than magi- ster (.20) and orcutti (.41). The members of each pair differ from each other, and further conclusions from electrophoresis may be misleading. Temperature Bogert (1949) computed average body temperatures of 10 forms of Sceloporus (Table 1). Two closely related forms {v. variabilis March 1974 LARSEN, TANNER: SCELOPORUS 17 Fig. 13. Dendrogram generated by external and skull characters (1 to 80). and IK olloporus) are separated by 1.4 degrees. However, a span of 1.3 degrees includes five widely divergent species {magister, undu- latus, poinsctti, grammicus, and merriami) . In fact, grammicus and merriami prefer the same temperature. It is doubtful that these data have any systematic value. Futher studies, however, may show that temperature preference or optimum temperature for enz}TTie systems can be useful. Paleontology Brattstrom (1955) reported some thoracic vertebrae, which he identified as Sceloporus jarrovi, in Late Pleistocene deposits in Zum- pango, Mexico. However, Cole (1970:27) has found that, "The fossil record of Sceloporus is practically nonexistent." Femoral Pore Secretions Hunsaker (1960:72) suggested that lizards can identify femoral pore secretions by olfaction or taste: In poinsetti and cyanogenys there is a marked disposition of the members of each species to separate when put together. The lizards of one species would establish common territories to the exclusion of the other species. When the secretions of each species were transposed, a reversal of the associative patterns 18 GREAT BASIN NATURALIST Vol. 34, No. 1 Fig. 14. Dendrogram generated by external, skull, and distribution characters (1 to 82). occurred, and the members of one species associated with the other and excluded members of their own species. If femoral pore secretions represent a species-specific territorial marker, then perhaps chemical analysis of these secretions will provide another valuable taxonomic character for future workers. Myology Secoy (1971) examined the myolog}' of eight species of Scelop- orus, including an extensive examination of c. clarki. She concluded that intrageneric myological variation is slight and that speciation in this genus is therefore recent. Although myology may be signifi- cant, and even diagnostic, at higher levels or wdth different taxa, its usefulness within the genus Sceloporus must yet be demonstrated. Materials and Methods Specimens for this study were acquired from several museum collections and through extensive field collecting by the authors. Most specimens were collected by noosing or shooting with .22 dust shot. The museum and locality data for specimens from the United States, Mexico, and Central America are as follows: gadoviae, BYU 36148 (skull), 45 March 1974 L.\RSEN, TANNER: SCELOPORUS 19 Clarkii •— Orcutti ^— Magister ^ Undulatus _ Occidenta Torquatus — Mucronatus Ornatus ^_ Dugesir i— Poinsett! Grammicus_ Variabilis Teopensif ^^ Chrysost icus Sini(erus Utiformi i Mer r iami -^^ Maculotus ^^ Ja iapae Py rocephalus- Nelsoni -^-^ h n 0.5 16 Fig. 15. Dendrogram generated by external, skull, distribution, and display characters (1 to 84). km S Neuva Italia, Michoacan; couchi BYU 36418 (skull), 36417, Huestaca Canon, 18 km W Monterrey, Nuevo Leon; merriami merriami BYU 36389 (skull), 13 km S Shumla (Hwy 90 and Pecos River), Val Verde Co., Texas; parvus scutulatus BYU 36125 (skull). 4 km N Zimapan, Hidalgo; parvus parvus BYU 36126. 36127, 7 km W 3 km N Santiago Anaya, Hidalgo; jalapae BYU 36423 (skull) 13 km SE Nochixtlan, Oaxaca. BYU 36422 near Tehuacen (Cacoa- lepam). Puebla; ochoterenae BYU 36004 (skull), 36003, 36005, 36006, Chilpan- cingo. Guerrero; maculosus FMNH 33548 (skull), 32007, 23 km NE Pedricena, Durango; grammicus microlepidotus BYU 36300 (skull), 36015, 36017, 36021, Puebla, Puebla, east side of Orizaba, Veracruz; pictus BYU 36419 (skull), sum- mit Mt. Acultzingo, Veracruz; megalepidurus BYU 36421 (skull), Lake El Chico, Hidalgo. BYU 36094. 36095, 3 km W Limon. Veracruz; cry plus AMNH 65835 (skull), Cerro de Humo, Oaxaca; heterolepis BYU 36420 (skull). Rancho Primarera, near Guadalaiara. Jalisco; asper FMNH 32041 (skull). 32043 Uropan, Michoacan; pyrocephalus BYU 36268 (skull). 36264, 36265, 36266, 24 km N Colima, Colima; nelsoni bar rancor urn BYU 14316 (skull), 14317, 14318, 14319, 14320, Urique, Chihuahua; scalaris scalaris BYU 36132 (skull), Zumpango, Mexico, BYU 36132. Yuridin, Guanajuato, BYU 36133, 2 km S 4 km E Villa Victoria, Mexico; aeneus aeneus BYU 36137 (skull), 3 km S Atlacomulco, Mexico, BYU 36136, 4 km S Mexicaltzingo. Mexico. BYU 36138, Salazar, Mexico, BYU 36139. Lagunas Zempoala. Morelos; siniferus cupreus BYU 36228 (skull), 36225, 26336, 26229. 74-108 km SE Oaxaca. Oaxaca; carinatus BYU 36424 (skull), Rancho Meyapac, Ocozocoautla, Chiapas; utiformis BYU 36400 (skull), 36401, 36402, 36403, 262 km S Guadalajara (Hwy 80). Jalisco; squamosus BYU 36044 (skull), Chinandega Nicaragua; variabilis variabilis BYU 36018 (skull), 36163, 36164, 36172, 39 km E Jalapa, Veracruz; cozumelae BYU 36428 (skull), 36425, 36426. 36427, 8 km W Progreso, Yucatan; teapensis BYU 36121 (skull), 36122, 20 km N Randales, Chiapas. BYU 36123, Montepio, Veracruz, BYU 36124, Catemaco, Veracruz; chrysostictus BYU 36129 (skull). Piste 10 m Yucatan, 20 GREAT BASIN NATURALIST Vol. 34, No. 1 FORMOSUS ■ SPINOSUS - HMRIOUS - OlIVACEUS . CAUTUS EDWARDTAYLODI. ClARKII ORCUTTI OCCIOCNTALIS. GRACIOSUS TOROUATUS CYANOGENYS— MUCRONATUS - POINSETTI JARROvn ORNATUS ASPER . GRAMMICUS MEGAIEPIDURUS. PICTUS VARIABIIIS TEAPENSIS SINIFERuS NELSONI .^-^ SCAIARIS. AENEUS _ COUCHII . 0.5 "P i> & 5=^ ^ ^ 16 Fig. 16. Dendrogram generated by external, skull, distribution, and chromo- some characters (1 to 83). Yucatan, BYU 36128, Isla mujeres, Quintana Roo; spinosus caeruleopunctatus BYU 36213 (skull), 36205, 36212, 36219, 16 km S. Oaxaca, Oaxaca; orcutti orcutti BYU 32321 (skull), mountains S of Cabazoh, Riverside Co., California, BYU 30080, 30081, Canyon Guadalupe, Juarez Mountains, Baja California; clarki clarki BYU 36056 (skull), 36053, 36054, 36055, San Rafael Trail. Arizona; melanorhinus calligaster BYU 14640 (skull), Puerto Vallarta, Jalisco; magister magister BYU 8848 (skull), Panoche. San Benito Co., California. BYU 9850, 26 km W Caliente, Lincoln Co., Nevada, BYU 23666, Leeds. Washington Co., Utah, BYU 12886, Hole in the Rock, Kane Co., Utah; olivaceus BYU 13048 (skull). Camp Bullis, Texas, BYU 36397. 36398, Laredo. Texas; cautus BYU 36250 (skull), 36251, 24 km SE Saltillo, Coahuila; horridus horridus BYU 36387 (skull), 36384, Iguala (185 km S Mexico City), Guerrero, BYU 36024, 36025, Chilpancingo. Guerrero, BYU 36231, 132 km S Mexico Citv Morelos; edwardtaylori BYU 36080 (skull), 8 km NW Salina Citjz, Oaxaca; formosus formosus BYU 36074 (skull), 36075, 36076, Llano de las Flores, Chiapas; lunaei FMNH 64687 (skull), 64691, Santa Clara, Sierra de las Minas, Guatemala; lundelli lundelli FMNH 32123 (skull), 32088, 30261, Balchacaj, Campeche; malachidcus rnalachiticus BYU 36032 (skull), 36029, 36030. 36031, Cerro de la Muerte, 95 km S San Jose, Costa Rica; acanthinus FMNH 20156 (skull). Tiquisata, Guatemala, FMNH 167111, Santa Clara, Sierra de las Minas, Guatemala. FMNH 10991, Hacienda Chileta, Sonsonate, El Salvador; undulatus elongatus BYU 20642 (skull), 20632, 20633, 20635, Yellow Cat Mining District, Grand Co., Utah; virgatus BYU 17031 (skull), 15487, 15488. 17030. 16 km SW San Pedro. Chihuahua; woodi BYU 8370 (skull), Englewood, Florida; occidenlalis biseriatus BYU 30097 (skull). 30094, 23873, 23875, 23878, Rainier Mesa, Nevada Test Site. Mercury-, Nve Co.. Nevada; graciosus graciosus BYU 16700 (skull). 33024, 33049. 21 km NE Provo, Wasatch Co., Utah, BYU 33057, 33058, 5 km E Spanish Fork, Utah Co.. Utah; jarrovi jarrovi BYU 36007 (skull), 36008, 36010. Huachuca Mountains. Arizona, BYU 36072, Saddle Mountain Trail. Arizona; lineolateralis FMNH 100174 (skull), 32030, 10 km NE Pedrecena, Durango; ornatus caeruleus BYU 36262 (skull), 36263, 68 km E Torreon, Coahuila; dugcsi dugesi BYU 36369 (skull), 36342, 36343, 36367, 36370, 165 km S Guadalajara, Jalisco; torquatus melanogaster BYU 36309 (skull), 36302, 36303, 36304, 36306, Morelia. Michoacan; cyanogenys BYU 36011 (skull), Rancho Santa Anna (13 km SE Padilla), Tamaulipas, BYU March 1974 LARSEN, TANNER: SCELOPORUS 21 Clarkii _ Orcutti _ Magister Occidentalis. Torquatus .^ Mucronatus. Poinsetti -^ Ornatus _^ Grammicus— Variabilis^ Teapensis_ Siniferis-^ Nelson! gralar is Merr iami — Maculosus Jalapae 0.5 n ? ^ 8 16 Fig. 17. Dendrogram generated by external, skull, distribution, display, and chromosome characters (1 to 85). 11402, 11404, 11405, Arroyo Vaqueriso, Nuevo Leon; bulleri BYU 40082 (skull), 36381, Autlan (185 km S Guadalajara), Jalisco; macdougalli FMNH 71661, AM 76119, Isthmus of Tehuantepec, Oaxaca; mucronatus omiltemanus BYU 36190 (skull), 36188, 36189. 105 km S Oaxaca, Oaxaca, BYU 36035, Omiltome, Guerrero; serrifer plioporus BYU 36182 (skull), 36183, 36149, 36185, 16 km E Jalapa, Veracruz; poinsetti poinsetti BYU 13812 (skull), 13814, 13815, 13820, 80 km W Chihuahua City, Chihuahua. External Characters The external characters used were chosen because of their suit- abihtv for numerical analysis. Keys and checklists (Smith and Taylor, 1950; Boulenger, 1885; Cope 1900; Van Denburgh, 1922) were examined and all quantitative characteristics were included. Color patterns were omitted because of variations caused by preser- vatives. The forty external characters utilized are: (1) Snout-vent length (mm). (2) Snout-vent/snout-parietal eye. (3) Humerus (from ventral midline to outside of elbow) /snout-vent. (4) Femur (from ventral midline to outside of knee) /snout-vent. (5) Outside length of tibia/snout-parietal eye. (6) Length of fourth toe/femur. (7) Height-to-width ratio of tail at point one head length from vent. (8) Snout-parietal eye (mm). (9) Width of head at parietal eye/snout-parietal -eye. (10) Vertical height of head at parietal eye/snout- parietal eye. (11) Width of head anterior to orbit/snout-parietal eye. (12) Distance between nares/snout-parietal eye. (13) Length of frontal scale (s) /snout-parietal eye. (14) Length of frontal scale (s) /snout-parietal eye. (14) Length of frontal scale (s) /smallest width of frontal. (15) Largest linear measurement on inter- nasal scale/snout-parietal eye. (16) Length of interparietal/width of same (through parietal eye). (17) Width of widest supraocular/snout-parietal eye. (18) Width of widest supraocular/length of same. (19) Parietal eye to posterior edge of interparietal/length of interparietal. (20) Length of median frontonasal/ width of same. (21) Length of median frontonasal/snout-parietal eye. (22) Dorsals from interparietal to posterior margin of thigh. (24) Dorsals equal to 22 GREAT BASIN NATURALIST Vol. 34, No. 1 Clarkii -^ Poinsett i ^[_ Magester I jgeste Orcutti — Ornatus — Nelsoni — Undulatus Occidentalis Grammicus Totquat Mucronat Teapensis Variabilis Siniferus ' Scalaris - Merriam ry cus — ^ us -J atus J Maculosus Jalapae -y 0.5 16 Fig. 18. Dendrogram generated by distribution, display, and chromosome characters (81 to 85). one head length (between points 2 and 3 head lengths posterior to interparietal). (25) Laterals equal to one head length midway between limbs. (26) Ventrals equal to one head length (between points 2 and 3 head lengths posterior of snout). (27) Dorsals equal to V2 head length (counting laterally from midline at a point 2 head lengths from interparietal). (28) Ventrals equal to i/^ head length (counting laterally from midline at a point 3 head lengths from snout). (29) Total femoral pores (both sides). (30) Ventrals between medial limits of femoral pores. (31) Ventrals from vent to a line connecting femoral pore series. (32) Caudals equal to one head length (between points 1 and 2 head lengths from vent). (33) Supra- labials (total both sides and rostral). (34) Infralabials (total both sides and mental). (35) Sublabials (total both sides and mental). (36) Caudals around tail one head length from vent. (37) Dorsals equal to one interparietal (counting posterior from interparietal). (38) Ventrals equal to one interparietal (counting anterior from vent). (39) Head shields in contact with interparietal. (40) Fourth toe lamellae. Skulls Preparation. Skulls were prepared by boiling 15-20 minutes in 50 ml water with a few drops of detergent and NH4OH. After boil- ing, they were allowed to dry until the muscles were easily removed with forceps. This procedure was repeated several times and the last remains of muscle were removed by dipping the skull in Clorox bleach. Whitening of skulls. Kier, Grant, and Yochelson (1965:453-56) described a technique widely used in paleontological preparations but possibly new to investigators of herpetological osteology. The skull was first blackened by dipping in ink. (Shafer's permanent blue-black is excellent because it stains the skulls effectively and is easily removed by dipping the skull in a mild solution of NH4OH.) The blackened skull was then highlighted with a thin layer of NH4CI. The dry NH4CI was placed in the chamber of a March 1974 larsen, tanner: sceloporus 23 24.000 . • 2<000 23.167 . ■ 23.167 22.313 . ■ 22-333 21.500 . • 21.500 20.667 . A • 20.667 19.0)3 . AA 19-833 19.000 . (A "■OOO 18.167 . AAA IB. 167 17.333 . A . 17.333 16.500 . • 16.500 15.667 . . 15.667 14.833 . ■ l<-833 14.000 . 14.000 13.167 . . U-167 12.333 . . 12.333 11.500 . . 11-500 10.667 . 10.667 9.833 . 9 933 9.000 . 9.000 8.167 . 8.167 7.J33 . 7 333 soo . .500 5.667 . 5.667 4.833 . < •33 4.000 . .4.000 3.167 . 3.167 2.333 . 2-333 1.500 . 1-500 0.667 . CC C • 0^" -0.167 . C C • -0-167 -1.000 . CCCaCCCC -1.000 -1.833 . CCC -1 »33 -2.667 . C CC C - -2.667 -3.500 . C • -3.500 -4.333 . - -<-333 -5.167 -6.000 -5.167 -6.000 . 6.833 . Sb -6 "3 -7.667 . B -7.667 -8.500 . B . -8.500 -9.333 . - -9-333 Fig. 19. Canonical display of three groups: 1(A), 11(B), and III(C). 100 ml pipette, and the open end of the pipette was attached with rubber tubing to a squeeze bulb. To vaporize the NH4CI, the pipette was heated over a flame. With careful pressure on the squeeze bulb, the skull was then highlighted with NH4CI vapor. This technique enhances the suture lines in black contrast and facilitates the study of photographs. Skull photography. Several workers have taken measurements directly from skulls with calipers (Weiner and Smith, 1965; Jenkins and Tanner, 1968; Avery and Tanner, 1971). However, the small size of some species makes it virtually impossible to take precise measurements directly from the skulls. Weiner and Smith (1965) made some of their skull measurements with the aid of an ocular micrometer and a microscope. But measurements through a micro- scope or on a photograph are subject to error caused by variation in angle of view. Such measurements would be acceptable, however, if the angle of view were kept constant. Lewis (1944) studied the determination of dress patterns from photographs and found that if the subjects were properly oriented, correct three-dimensional dress patterns could be determined. To minimize the problem of distortion and provide constant orientation of skulls, special equip- ment was constructed. The apparatus (Fig. 1) was constructed to minimize variations in angle of view. This structure consists of a circular outer platform that can be leveled with spirit levels. The skull is placed on a second platform in the center. The inner plat- 24 GREAT BASIN NATURALIST Vol. 34, No. 1 15.6 -11.6 -7.6 -3.6 0. 5.733 . 5.733 5.06 7 . Q .5.06 7 4.400 . 4.400 3.733 . . 3.733 3.067 . #0 0 '•"*' 2.400 . 2.400 1.733 . C '-"^ 1.067 . . 1.067 0.400 . . 0.400 -0.267 . . -0.267 -0.933 . . -0.933 -1.600 . ■ -1.600 -2.267 . . -2.267 -2.933 . . -2.933 -3.600 . f^ -3.600 -4.267 . B ■ -4.267 -4.933 . • ■ -4.933 -5.600 . B ■ -5.600 -6.267 . . -6.267 -6.933 . . -6.933 -7.600 . ■ -7.600 -19.6 -I'.b -11.6 -7.6 -3.6 0.4 4.4 8.4 12.4 16.4 Fig. 20. Canonical display of the three subgroups of group I. form can be tilted along two planes as well as adjusted vertically until specific reference lines on the skull are parallel with the out- side platform. A camera (Nikon FTN with Kodak plus X film) was placed over the skull with spirit levels attached to the camera back so that the reference lines through the skull and the film in the camera were always as nearly parallel as possible. A line through the tip of the premaxilla and the center of the foramen magnum was the first reference for dorsal, ventral, and lateral views. The second reference line for the dorsal view passed through the anterolateral corners of the parietals. The second refer- ence line for the ventral view passed through the lateral tips of the ectoptery golds. The second reference line for the lateral view was the surface of the frontoparietal suture, which was oriented at right angles to the outer platform. The posterior view was arranged so that the surface of the parietal bone was at right angles to the outer platform and a line through the lateral tips of the exoccipitals was parallel to the outer platform. Further to minimize possible error caused by variation in angle of view, all skull measurements were converted to ratios between two distances measured in the same direction on the same photo- graph. Although this technique reduces the effects of distortion, it unfortunately eliminates most of the traditional skull characters (width and length ratios of skull members). Illustrations were prepared by projecting and tracing the photo- graphs with a Saltzman Projector. Detail was added to the tracings with the aid of a binocular microscope (Presch, 1969; Nash and Tanner, 1970). Skull Characters The following 40 characters (numbers 41-80) were computed for each skull: (41) Posterior extent of supraoccipital on midline to anterior border of parietal foramen/parietal foramen to suture between nasals (Fig. 2). (42) Length of suture between nasals/parietal foramen to suture between nasals (Fig. 2). (43) March 1974 LARSEN, TANNER: SCELOPORUS 25 12.0 32.0 S2.0 12.000 S.666 S.333 2.000 -1.334 -4.667 -8.000 12.000 e.666 S.333 2.000 -1.334 -4.667 -e.ooo -11.334 -14.667 12.0 32.0 52.0 Fig. 21. Canonical display of the five subgroups of group II. Length of premaxilla/parietal foramen to suture between nasals (Fig. 2). (44) Posterior tip of suture between frontal and nasal to posterior end of suture be- tween prefrontal and nasal/posterior end of suture between prefrontal and nasal to anterior end of suture between prefrontal and nasal (Fig. 2). (45) Anterior end of suture between prefrontal and nasal to anterior end of suture between maxillary' and nasal/parietal foramen to suture between nasals (Fig. 2). (46) Anterior end of suture between maxillary and nasal to anterior end of suture between maxillary and premaxillary /parietal foramen to suture between nasals (Fig. 2). (47) Pineal foramen to posterior end of suture between prefrontal and lacrimal/parietal foramen to suture between nasals (Fig. 2). (48) Posterior extent of lateral wing of parietal to posterior end of suture between parietal and postorbital/posterior extent of supraoccipital on midline to anterior border of parietal foramen (Fig. 2). (49) Length of postorbital/posterior end of suture between parietal and postorbital to anterior edge of parietal foramen (Fig. 2). (50) Posterior tip of prefrontal to anterior end of suture between prefrontal and nasal/posterior end of suture between prefrontal and lachrymal to anterior tip of prefrontal (Fig. 2). (51) Most narrow width of frontal/anterior width of parietal (along suture with postorbital) (Fig. 3). (52) Lateral side of jugal on transverse line through anterior border of parietal foramen to lateral extent of suture between postorbital and parietal/anterior width of parietal (Fig. 3). (53) Interfenestral width (on line passing through posterior tips of both postorbitals)/ anterior width of parietal (Fig. 3). (54) Lateral edge of parietal on line passing through posterior tips of both postorbitals to posterior tip of postorbital on same side/anterior width of parietal (Fig. 3). (55) Anterior end of suture between prefrontal and nasal to posterior end of suture between prefrontal and lacrimal distance between left and right anterior ends of suture between prefrontal and nasal (Fig. 3.) (56) Internarial width/distance between left and right anterior ends of suture between maxillary and premaxillary (Fig. 3). (57) Posterior tip of occipital condyle to medial comer of tip of basipterygoid process of the basi- sphenoid/lateral tip of ectopterygoid to anterior tip of premaxilla (Fig. 4). (58) Medial comer of tip of basiterygoid process to lateral tip of ectopterygoid/lateral tip of extoptery'goid to anterior tip of premaxilla (Fig. 4). (59) Posterior corner of lateral side of palatine to lateral limit of suture between palatine and maxilla/ lateral tip of ectopterygoid to anterior tip of premaxilla (Fig. 4). (60) Posterior 3S.000 . 30.000 . 2S.000 . 20.000 . li.OOO . 10.000 . S.OOO . -S.OOO . -10.000 . -IS. 000 . 20.000 . • B 25.000 . B 30.000 . 35.000 30.000 25.000 20.000 15.000 10.000 5.000 0.0 -5.000 -10.000 -15.000 -20.000 -25.000 -30.000 Fig. 22. Canonical display of the five subgroups of group III. 26 GREAT BASIN NATURALIST Vol. 34, No. 1 Table 1. Average body temperature of some Sceloporus as reported by Bogert (1949). Temperature C Species 36.9 variabilis variabilis 36.2 ivoodi 35.4 variabilis olloporus 35.3 squamosus 34.9 ■ magister 34.8 , undulatus consobrinus 34.2 poinsetti 33.6 grammicus disparilis 33.6 merriami 32.9 formosus malachiticus tip of quadrate ramus of pterygoid to lateral tip of ectopterygoid/lateral tip of ectopterygoid to anterior tip of premaxilla (Fig. 4). (61) Lateral tip of ecto- pterygoid to medial limit of suture between ma.xilla and ectopterygoid/distance between lateral tips of ectopterygoid (Fig. 5). (62) Medial limit of suture be- tween maxilla and ectopterygoid to posterior corner of lateral side of palatine/ distance between lateral tips of the ectopterygoid (Fig. 5). (63) Posterior corner of lateral side of palatine to medial limit of suture between palatine and pterygoid/ distance between lateral tips of ectopterygoids (Fig. 5). (64) Smallest widtli of basisphenoid/distance between lateral tips of ectopterygoids (Fig. 5). (65) Diagonal distance from lateral tip of ectopterygoid on one side to posterior tip of quadrate ramus of pterygoid on other side/length between the same points on one side (Fig. 5). (66) Five times the tangent of the angle between the midline and the extended line that passes through the midpoint on the tip of the basi- pterygoid process and the midpoint on the most narrow part of the neck of the basipterygoid process (Fig. 5). (67) Tip of premaxilla to most ventral extent of ectopterygoid projected onto a line from the tip of premaxilla to tip of quadrate ramus of pterygoid/tip of premaxilla to posterior tip of postorbital (Fig. 6). (68) Most ventral extent of ectopterygoid to tip of quadrate ramus of pterygoid pro- jected onto a line from the tip of premaxilla to tip of quadrate ramus of pterygoid/ tip of premaxilla to posterior tip of postorbital (Fig. 6). (69) Tip of premaxilla to anterior end of suture between prefrontal and lacrimal (parallel to denomi- nator)/tip of premaxilla to posterior tip of postorbital (Fig. 6). (70) Anterior end of suture between prefrontal and lacrimal to posterior tip of prefrontal (parallel with denominator) /tip of premaxilla to posterior tip of postorbital (Fig. 6). (71) Posterior tip of prefrontal to anterior end of suture between post- frontal and parietal (parallel with denominator) /tip of premaxilla to posterior tip of postorbital (Fig. 6). (72) Anterior end of suture between prefrontal and lacrimal to posterior tip of prefrontal (direct) /same as numerator projected onto the line between the tip of the premaxilla and the posterior tip of postorbital (Fig. 6). (73) Posterior tip of prefrontal to most ventral extent of ecto- pterygoid/anterior end of suture between postfrontal and parietal to tip of quadrate ramus of pterygoid (Fig. 6). (74) Dorsal ridge of supraoccipital to dorsal edge of foramen magnum/top of parietal at midline (passes vertically through medial ridge of supraoccipital and through center of occipital condyle) to ventral edge of parietal at midline (Fig. 7). (75) Height of foramen magnum along midline/top of parietal to ventral edge of parietal (Fig. 7). (76) Ventral edge of foramen magnum on midline to ventral edge of condyle/dorsal-ventral height of parietal (Fig. 7). {77) Dorsal corner of lateral process of exoccipital to ventral corner of lateral process of exoccipital/dorsal-ventral height of parietal (Fig. 7). (78) Distance between right and left dorsal corners of lateral process of exoccipital/distance between right and left ventral corners of basioccipital tubercles (Fig. 7). (79) Five times the tangent of the angle formed by the dorsal comer of the lateral process of the exoccipital and its intersection with the mid line (at right angles) and the ventral comer of the basioccipital tubercle (all points on one side) (Fig. 7). (80) Five times the tangent of the angle formed March 1974 LARSEN, TANNER: SCELOPORUS 27 Table 2. Groups and subgroups in the genus Sceloporus. Group I Group II Group III Subgroup A Subgroup A Subgroup A gadoviae grammicus spinosus picius orcutti megalepidurus clarki cryptus melanorhinus shannonorum* magister heterolepsis olivaceus asper cautus horridus edwardtaylori Subgroup B Subgroup B Subgroup B couchi pyrocephalus formosus merriami nelsoni lunaei malachiticus acanthinus Subgroup C Subgroup C Subgroup C maculosus scalaris undulatus parvus goldmani* virgatus jalapae aeneus woodi ochoterenae occidentalis graciosus Subgroup D Subgroup D siniferus jarrovi carinatus lineolateralis utiformis ornatus squamosus dugesi Subgroup E Subgroup E variabilis torquatus cozumelae cyanogenys teapensis bulleri chrysostictus insignis* macdougalli mucronatus serrifer poinsetti 'Species not examined in this study. by the shortest width of the parietal, its intersection with the midline, and the line from that intersection to the dorsal corner of the lateral process of the exoc- cipital (all points on one side) (Fig. 7). Other Characters Karyology. Extensive karyological data are available for most species of Sceloporus (see Literature Review — Karyolog}')- Al- though many characters can be described for each species, the only karyological character included in this study is the number of micro- chromosomes. A change in microchromosomal number is theoreti- cally a single karyotic event, and such events may indicate rela- tionships. Hopefully, additional characters will soon be available for numeric analysis. Display -action patterns. Although display-action patterns in- volve a complex of activities, only two measurements were con- 28 GREAT BASIN NATURALIST Vol. 34, No. 1 sidered: the ratio of vertical movement of the shoulder to the Aerti- cal movement of the eye and the ratio of vertical movement of the hip to the vertical movement of the eye (Purdue and Carpenter, 1972b). Zoogeography. The approximate latitude and longitude of the center of distribution for each species were included as additional characters. Of course, a simple measure of latitude and longitude does not allow for altitude, climate, habitat preference, or natural barriers such as mountain;s and rivers. However, differences in lati- tude and longitude are a measurement of horizontal distance and represent a crude measure of natural resistance to gene flow. Lati- tude is also somewhat correlated with climatic gradients. Data Analysis Justification. Hennig (1966:74) defined species relationships as follows: "A species 'x' is more closely related to another species 'y' than it is to a third species 'z' if and only if it has at least one stem species in common with species 'y' that is not also a stem species of 'z'." Hennig proposed that classification be based on phylogenetic kinship and not form similarity because frogs and tadpoles should not be different taxa. Bigelow (1958) said that the measure of phylo- genetic relationship is the "relative recency of common ancestry." These definitions of relationship seem to be circular. Phylogenies are based on circumstantial evidence, so it is impossible for Hennig to prove whether or not "x" and "y" have a stem species that is not an ancestor of "z." We believe that phylogenetic relationships are manufactured in the mind of the taxonomist from phenetic data and form similarities. We therefore reject Ilennig's and Bigelow's pro- posals and suggest that the best phenotypic dendrogram (containing representatives of all populations of the group under consideration and including a sufficient number of characters manipulated in the best numeric manner) is also the best source for the most probable phylogeny. Sokal and Sneath (1963) suggested that at least 60 characters are necessary for highly significant results. Our study has Table 4. Means and standard deviations for the eight diagnostic characters in groups I, II, and III. Group IS Group I Group II Group III Character Mean S.D. Mean S.D. Mean S.D. 32 13.97 1.00 9.90 1.55 7.90 0.72 7 10.02 0.88 10.98 1.23 9.45 0.63 27 9.08 1.72 6.59 1.65 4.39 1.03 13 4.05 0.17 3.84 0.22 3.71 0.21 30 4.71 4.55 9.49 5.64 9.04 3.06 6 10.15 3.01 10.93 1.14 9.32 1.00 39 7.19 1.94 7.48 1.57 5.70 1.04 70 2.56 0.19 2.34 0.24 2.55 0.19 March 1974 LARSEN, TANNER: SCELOPORUS 29 Q ^ o> S a £ [x. ^ H C .xo-r-iooocr5 «H'S OOfON.t^t^t^^'O Orrnat^OOODCOCO OOOOOOOOOOOOOOOO 0) ^ o > < a OJ C/D I o5 CO •* vn 00 PC iE < ^ fe o ii Ph [_| 'Ih (J 01-^ *H ^ rl ^'^ o a! f^u2 So 03 03 CO < tovn of of Ol^ *"^ o t^ fl td^' ■ rt 00 tn i/> Si a> o CO ^ -s o crio •jd OOO ^ o c o « Ph 30 GREAT BASIN NATURALIST Vol. 34, No. 1 b Is-I 00 CT) O) 01 'J- roxi -Val stati ro ^ rq 55 00 (£> '< «5 OO OO OO xri b-i C^ ^^6 < ^ ^ 1 PO 05 o ■^^ T-H ^^ T— I 00 t^ be O 0) S 2 t}-" '^'^ '^'^ ^'' '*•" '^'" 'J-" Q cS 4) O tH ^ h fl Ph Ph Ph «« ^ •^ Oi CO CJ) vn ■I C5 CSl O) (M C^ CS) vn vn xn to in vn to cs) PO •<}- \n <£> t^ 3 S bo 3 W U m 00 PO PO to t^ t^ o -ing size. The number of dimensions in the hyperspace is equal to the number of variables measured. This concept of individuals being repre- 32 GREAT BASIN NATURALIST Vol. 34, No. 1 fe 1^ /-N S 31^ X « « < '^ 0) g 4) O M O 01 fa >. ^ ^^ c S3W 3m Ph J^< o oo xn O) ■<*■_ CO ai ^' trj oi oi oi ■^ "}- -^ ^ t}- -^ en i^ xn XT) O") ^ CO (O (£5 iO O T-^ tJ- ^ tJ- ^ csi xn vn u^ xTi xn oo -^ 'i- vn xn XT) CO 00 00 00 00 O^ 05 ro ■; d rc) d cS cS 00 o xn to 00 -r^ rq xn xn -^ C> t}- xri xn t>l d PO to 00 00 O to O PO to xn oj • PO 'ij- March 1974 larsen, tanner: sceloporus 33 Table 12. Stepwise discriminate analysis of group III. Number of pairs separated at .95, .99 and .999 confidence levels for each step. Confidence Level ____^^_ Step ^5 .99 ' .999 16 6 5 2 9 9 8 3 10 10 8 4 10 10 9 5 10 10 8 6 10 10 10 7 10 10 10 8 10 10 9 9 10 10 99 10 10 10 9 sented as points in a p-dimensional space is essential to cluster and discriminant analyses. Ward's method of cluster analysis forms spherical clusters of individuals in tlie hyperspace. New clusters are formed by measuring the distance from each individual in the original cluster to the center of the cluster, called the centroid. These distances are summed to form the error sum of squares for the cluster. The individuals to be added to the cluster are conditionally added, and the new centroid formed. An error sum of squares for the newly formed cluster is calcu- lated. This procedure is done for all possible entries to the original cluster (pos- sible entries include other clusters as well as individuals). The entr>^ that causes the least increase in the error sum of squares is joined to the original cluster. Each new cluster is formed bj^ joining those individuals that move the centroid the smallest distance. In other words, each cluster is composed of those individuals located closest to each other in the hyperspace. Thus, it is seen that this method unites individuals of the highest morphological similarity first (Wishart. 1969). Besides producing a phenetic dendrogram. Wishart's program (1968) provides other useful information, including (1) raw-data listing (numeric and binary), (2) maxima and minima for nu- meric data, (3) standard scores for numeric data, (4) means and standard deviations, (5) product-moment correlation coefficients, (6) principle components eigenvalues, (7) percentage and cumula- tive variances, (8) eigenvectors, (9) binary attribute frequencies, (10) binary attribute percentage occurrences, (11) similarity ma- trix, (12) "normaHzed" classification array, (13) listing of sample numbers for each cluster, (14) cluster means, standard deviations, F-ratios and T-values for continuous variable in each cluster, (15) cluster frequencies for binary attributes, (16) cluster percentage oc- currences for binary attributes, (17) binary attribute percentage ratios. The F-ratios printed for each character in each cluster are com- puted as the variation within that cluster divided by the variation in the total population. It must be remembered that this is not the traditional F-ratio (variance within clusters /variance between clus- ters). A character with a low F-ratio in one cluster is not neces.sarily a diagnostic character. The high total variance may be caused by variance within another cluster rather than variance between clus- 34 GREAT BASIN NATURALIST Vol. 34, No. 1 Table 13. X and Y canonical coordinates of the < species in groups I, II, and III. Species X Y Species X Y Group I Group III gadoviae -8.8 18.4 spinosus 12.5 -1.7 merriaTni -7.8 19.6 edwardtaylori 11.9 -1.8 couchi -^.5 18.1 melanorhinus 10.1 -0.4 parvus -7.7 20.1 clarki 12.7 -1.1 maculosus -8.2 18.7 orcutti 13.7 -0.8 ochoterenae -8.8 17.5 magister 11.4 -1.2 jalapae -8.8 20.9 horridus 11.0 0.6 olivaceus 11.9 -3.3 cautus 11.3 -1.1 Group II formosus 11.3 -0.9 asper -14.9 -6.5 malachiticus 12.9 -1.4 grammicus -15.3 -6.0 lunaei 11.7 -2.7 heterolepis -15.4 -5.7 lundelli 12.7 -1.1 megalepidurus -16.2 -5.2 undulatus 10.1 -2.5 pictus -14.6 -6.6 virgatus 11.3 -1.1 cryptus -15.2 -7.1 occidentalis 12.2 -1.5 pyrocephalus -16.2 -5.1 woodi 12.7 -0.4 nelsoni -15.5 -7.2 graciosus 11.7 -1.2 scalaris -15.8 -5.4 lineolateralis 10.7 1.0 aeneus -13.5 -5.2 ornatus 12.0 -2.4 siniferus -14.1 -6.6 dugesi 9.4 -1.3 squamosus -15.4 -7.8 jarrovi 13.6 -2.7 carinatus -16.1 -8.5 torquatus 11.9 -2.7 utiformis -15.0 -5.7 poinsetti 12.5 -1.0 variabilis -14.3 -5.6 cyanogenys 10.9 -1.5 cozumelae -14.5 -6.4 serrifer 10.4 -1.2 teapensis -17.2 -6.2 mucronatus 12.7 0.3 chrysostictus -13.3 -6.3 bulleri 14.2 -1.3 ters. With this limitation in mind, Wishart's program is extremely valuable. As an independent check on this system, Ward's Cluster Analy- sis was applied to some of the data used by Smith in 1939 (pers. comm.). Because of the nature of Smith's data, only 12 characters were used: (1) snout-vent length (mm), (2) snout-occiput/ snout— vent, (3) snout— ear/snout-vent, (4) length of hind leg/ snout-vent, (5) length of tibia/snout-vent, (6) length of fourth toe/snout-vent, (7) length of fourth toe/length of fifth toe, (8) lamellae on fotirth toe, (9) femoral pores (total both sides), (10) dorsal scales from occiput to back of thigh, (11) ventral scales from front of arm to vent, and (12) scale rows around body. The resulting dendrogram is illustrated in Figure 8. The strong similarity between Figure 8 and Smith's phylogeny (1939) sug- gests that there is a similar mechanism in Ward's Cluster Analysis and the subjective thinking of classical taxonomy. This supports the conclusion that a person capable of considering 60 to 80 characters on 50 to 60 species simultaneously, would arrive subjectively at re- sults similar to those produced by Ward's Cluster Analysis. Stepwise Discriminate Analysis. The purposes of stepwise dis- criminate analysis are (1) to determine the validity of proposed March 1974 larsen, tanner: sceloporus 35 Table 14. X and Y canonical coordinates for the species in the three sub- groups in Group I. Species X Y Species X Y Subgroup A Subgroup C gadoviae -15.3 -3.3 parvus 1.4 3.1 maculosus 1.2 3.3 Subgroup B ochoterenae 0.5 5.1 merriami 12.1 -5.7 jalapae -0.2 2.0 couchi 12.3 -A.2 groups, (2) to determine the relative diagnostic value of each char- acter, and (3) to classify individuals according to the proposed groups. The stepwise discriminate analysis computer program pub- lished by Dixon (1967) considers one character at a time according to F-ratios (variation between groups/variation within groups). Each step "includes" the remaining character with the highest F- ratio. At each step the species are classified according to all the included characters. As more characters are included, the differences between group means become more significant (as indicated by a U-statistic and an approximate F- value), and more individuals are properly classified. Dixon's program produces ( 1 ) means for each group and overall means for each character, (2) standard deviations for each charac- ter in each group, (3) within-group covariance matrix, (4) with- in-group correlation matrix, (5) detailed results for each "step." At each step a new variable is included and the program prints the F-ratios (and degrees of freedom) for all variables included and not included. The program also prints a U-statistic with degrees of freedom for an estimate of the significance of group separation. Since the U-statistic becomes extremely small and exceeds the capa- city of most charts, the program computes an approximate F that, with its degrees of freedom, can show the confidence level for over- all group separations. Dixon's program also produces an F-matrix in which F-ratios are computed for every possible combination of two groups. If the characters included provide good overall separation but fail to dis- tinguish between two groups, this matrix will quickly show which pairs are not separated at .9, .99, and .999 levels of significance. (F tables are not included in the printout.) This program also produces canonical functions for each step. These functions are then used to classify all species according to the information in the included characters. As more characters are in- cluded, the percent of species properly classified increases if the original groupings are valid. At the end of the printout the "dis- tance" is computed from each species to the center (centroid) of each group. Posterior probability for inclusion in each group is also computed for each species. The computer was instructed to "include" characters only if their F-ratio exceeded 1.0 and to "remove" char- acters if their F-ratio fell below .0,5. A summary of variables "in- 36 GREAT BASIN NATURALIST Vol. 34, No. 1 Table 15. X and Y canonical coordinates for the species in the five sub- groups in Group II. Species X Y Species X Y Subgroup A Subgroup D asper -30.0 7.5 siniferus 101.7 -1.2 grammicus -32.2 5.6 squamosus 102.2 -0.1 heterolepis -31.0 7.6 carinatus 101.7 0.1 megalepidurus -32.1 9.0 utiformis 101.1 0.5 pictus -30.0 6.1 cryptus -33.0 6.7 Subgroup E variabilis -57.3 -6.5 Subgroup B cozwnelae -57.3 -6.0 pyrocephalus -9.3 -6.5 teapensis -59.1 -4.8 nelsoni -9.7 Subgroup C -7.4 chrysostictus -56.8 -5.9 scalarus 14.6 -3.3 aeneus 16.1 -1.2 eluded" or "removed" including F-ratios and U-statistics is printed near the end of the printout. Then the program tabulates ( 1 ) eigen- values, (2) cumulative proportion of total dispersion, ( 3) canoni- cal correlations, (4) coefficients for canonical variables, (5) canon- ical variables evaluated at group means, and (6) graph coordi- nates for the first and second canonical variables for each species in each group. The program terminates with a graphic representa- tion of all species on the first two canonical variables. Canonical Analysis. The canonical analysis (a part of Dixon's program) computes a pair of linear coefficients for each character so that the greatest separation of groups can be displayed on a two- dimensional graph. For example. Figure 9 shows groups A and B plotted on characters p and q. To reduce the illustration from two dimensions to one, all points must be projected onto a single line. Line X would show more information about groups A and B than would line Y. In accordance with this idea, the canonical analysis rotates an imaginary plane through a multidimensional hyperspace until the best separation of groups is displayed. The lists of canonical functions are used to classify additional individuals according to the originally proposed classification and the "best" characters. Results Skulls. Skulls of 1 3 species of the genus Sceloporus are illustrated in Figure 10. Cluster Analysis. Ward's cluster analysis was applied to the fol- lowing sets of variables: 1. External characters 1-40 (Fig. 11). 2. Skull characters 41-80 (Fig. 12). 3. External and skull characters 1-80 (Fig. 13). 4. External, skull, and distribution characters 1-82 (Fig. 14). 5. External, skull, distribution, and display characters (Fig, 15). 6. External, skull, distribution, and karyological characters (Fig. 16). March 1974 LARSEN, TANNER: SCELOPORUS 37 Table 16. X and Y canonical coordinates for the species in the five sub- groups of Group III. Species X Y Species X Y Subgroup A Subgroup C spinosus 103.9 -3.9 undulatus -90.4 25.2 edwardtaylori 104.6 -5.3 virgatus -90.7 26.3 melcinorhinus 103.1 -5.8 occidentalis -91.7 27.0 clarki 102.5 -4.2 woodi -92.1 26.9 orcutti 102.4 -5.6 graciosus -91.3 28.1 magister 104.9 -5.8 horridus 104.3 -4.4 Subgroup D olivaceus 102.2 -4.6 jarrovi -22.5 0.32 cautus 104.7 -3.7 ornatus -23.8 2.6 dugesi -23.9 0.2 Subgroup B lineolateralis -22.5 3.1 formosus -158.3 -21.0 malachiticus -158.4 -19.7 Subgroup E lunaei -158.4 -22.5 torquatus 67.5 1.6 lundelli -157.1 -19.5 poinsetti 68.7 1.7 cy^anogenys 67.0 0.8 serrifer 69.9 0.6 mucronatus 66.1 1.0 bulleri 68.0 1.1 7. External, skull, distribution, display, and karyological characters (Fig. 17). 8. Distribution, display, and karyological characters (Fig. 18). Analysis and Conclusions Species Groups. The minor differences among these eight den- drograms were resolved subjectively to produce three major divisions and 13 species groups (Table 2). Careful examination will reveal the similar patterns generated by different sets of data. Figure 18 shows that distribution, display, and chromosome characters pro- duce results similar to those produced by external and osteological characters. This increases our confidence in Table 2 and demon- strates the taxonomic value of zoogeography, behavior, and kary- ology. Stepwise Discriminate Analysis. Dixon's stepwise discriminate analysis program cannot consider as many as 82 characters in one run. It was therefore used with external characters alone (1-40) and skull characters alone (41-80). For both studies the program was run four times to consider the three groups and their sub- groups. The number of characters in the data set was reduced by eliminating those characters with consistently small F-ratios. The characters removed were"numbers 2, 5, 15, 18, 19, 25, 42, 45, 46, 51, 65, 68, 73, 76, and 78. With the inclusion of latitude and longitude, this program then evaluated 66 characters. Table 3 shows the results of stepwise discriminate analysis of the three groups in the genus Sceloporus. This table shows that when characters are considered according to F-ratios, eight charac- ters are sufficient for correct classification of all species: (1) size of caudal scales, (2) degree of compression of tail, (3) width of 38 GREAT BASIN NATURALIST Vol. 34, No. 1 dorsal scales, (4) length of frontal scale, (5) ventrals between me- dial limits of femoral pores, (6) length of fourth toe, (7) head shields in contact with interparietal, and (8) length of prefrontal bone. The first eight characters contribute variation significant at the .99 level. Of the 39 most diagnostic characters (F-ratio greater than 1.0), 10 are osteological, 1 is geographical (longitude), and 28 are external. The null hypothesis that the three groups are not dif- ferent is rejected at the .999 level after consideration of the first character (number of caudal scales equal to one head length). The same hypothesis is rejected at the .999 level with respect to any com- bination of two groups. The eight diagnostic characters with means and standard deviations are shown in Table 4. Table 5 shows the results of stepwise discriminate analysis for the three subgroups of group I. Two characters are sufficient to classify the species into their subgroups. Both characters contribute variation significant at the .999 level. The first is a measure of the degree of compression of the tail, and the second is the number of ventrals. At every step, the separation of means is significant at the .999 level after inclusion of the second character. Of the six most diagnostic characters (F-ratio greater than 1.0), two are osteological and four are external. Table 6 shows the means and standard devia- tions of the first two characters. Tables 7, 8, and 9 show the results of stepwise discriminate analysis of the five subgroups of group II. Four characters are suffi- cient for 100-percent correct classification of all species: (1) the rel- ative width of the supraoculars, (2) the number of lamellae on the fourth toe, (3) the degree of compression in the tail, and (4) a measure of the relative width of the basisphenoid bone. Of the 13 most diagnostic characters (F-ratio greater than 1.0) 4 are osteologi- cal, 1 is geographical (latitude), and 8 are external. The approxi- mate F (U-statistic) for overall separation of means is significant at the .999 level for every step. Tables 10, 11, and 12 show the results of stepwise discriminate analysis of the five subgroups within group III. Six characters are sufficient for correct classification: (1) the size of ventral scales in the vent region, (2) latitude, (3) the number of dorsals equal to the length of the head, (4) the compression of the tail, (5) the size of dorsals near the interparietal scale, and (6) the shape of the ectopterygoid bone. The separation of the subgroups of group III is as significant as is the separation in group II. Of the 24 most diagnos- tic characters (F-ratio greater than 1.0) 8 are osteological, 1 is geo- graphical (latitude), and 15 are external. Tables 3 to 12 show that the groups and subgroups proposed in Table 2 are distinct at the .999 level of confidence according to the characters used in this study. These tables also show which charac- ters are most diagnostic among the groups. Canonical Analysis. Figure 19 shows the first two canonical di- mensions for the species in each major group. Table 13 gives the x and y coordinates for each species. This canonical separation gives March 1974 larsen, tanner: sceloporus 39 strong support to the conclusion that each of the tliree groups is monoj)hyletic and should be given taxonomic recognition. Figures 20, 21, and 22 and Tables 14, 15, and 16 indicate that the subgroups of each major group are also distinct phenetic units with no overlap. Literature Cited Avery, D. F., and W. W. Tanner. 1971. Evolution of the iguanine lizards (Sauria: Iguanidae) as determined by osteological and myological characters. Brigham Young Univ. Sci. Bull., Biol. Ser. 12(3): 1-79. BiGELOW, R. S. 1958. Classification and phylogeny. Syst. 7 -1. 7:49-50. BocuRT, M. F. 1874. fitudes sur les reptiles. Mission Sciei ' que au Mexique et dans TAnierique Central — Recherches Zoologir ■"■ (q. i.): 3d part. BoGERT, C. M. 1949. Thermoiegulation and ec I ody temperatures in Mexican lizards of the genus Sceloporus. An. In Biol. Univ. Nacion., Mexico 20 ( 1/2): 41 5-426. BouLENGER, G. A. 1885. 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Mus. Nat. Hist. 105(1): 1-142. Nash, D. F., and W. W. Tanner. 1970. A comparative study of the head and thoracic osteology and myology of the skinks Eumeces gilberti Van Den- burgh and Eumeces skiltonianus (Baird and Girard). Brigham Young Univ. Sci. Bull., Biol. Ser. 12(2): 1-32. Presch, W. 1969. Evolutionary osteology and relationships of the horned lizard genus Phrynosoma (family Iguanidae). Copeia 1969(2) : 250-75. Purdue, J. R., and C. C. Carpenter. 1972a. A comparative study of the dis- play motion in the iguanid genera Sceloporus, Uta, and Urosaurus. Herpe- tologica 28(2): 137-41. . 1972b. A comparative study of the body movements of displaying males of the lizard genus Sceloporus (Iguanidae). Behavior 41:68-81. Savage, J. M. 1958. The iguanid lizard genera Urosaurus and Uta, with re- marks on related groups. Zoologica 43(2):41-54. Secoy, D. M. 1971. The myology of Sceloporus c. clarki Baird and Girard (Reptilia: Iguanidae). Brigham Young Univ. Sci. Bull., Biol. Ser. 14(1 ):1- 22, 8 figs. Smith, H. M. 1938. The lizards of the torquatus group of the genus Scelop- orus Weigmann. Univ. Kansas Sci. Bull. 24:539-693. . 1939. The Mexican and Central American lizards of the genus Scelop- orus. Field Mus. Nat. Hist. Zool. Ser. 26:1-397. , and E. H. Taylor. 1950. An annotated check list and key to the rep- tiles of Mexico, exclusive of the snakes. Bull. U.S. Nat. Mus. (199): 1-199. 1966. Herpetology of Mexico: annotated checklists and keys to the amphibians and reptiles. A reprint of Bulletins 187, 194, and 199 of the U.S. National Museum, with a list of subsequent taxonomic innovations. Ashton, Maryland. Eric Lundberg. Smith, N. M., and W. W. Tanner. 1974. A taxonomic study of the western collard lizards, Crotaphytus collaris and Crotaphytus insularis. Brigham Young Univ. Sci. Bull., Biol. Ser. 19(4). In press. SoKAL, R. R., AND P. A. Sneath. 1963. Principles of numerical taxonomy. W. H. Freeman. San Francisco. Van Denburgh, J. 1922. The reptiles of western North America . . . , Vol. 1: Lizards. San Francisco, Calif. Acad. Sci. 611 pp., 57 pis. Webster, T. P., W. P. Hall, III, and E. E. Williams. 1972. Fission in the evolution of a lizard karj'otype. Science 177 (4049) : 61 1-13. Weigmann, A. F. 1828. Beitrage zur amphibienkunde. Isis von oken, v. 21:369- 70. March 1974 larsen, tanner: sceloporus 41 1834. Herpetologia Mexicanna seu descriptio amphibiorum Novae Hispaniae. Pars. Prima. Saurorum species. Berlin, Luderitz. v. i., 54 pp., 10 pis. Weiner, N. J. AND H. M. Smith. 1965. Comparative osteology and classifica- tion of the Crotaphytiform lizards. Amer. Midi. Nat. 73(1) : 170-87. WisHART, D. 1968. A Fortran II programme (CLUSTAN) for numerical clas- sification. Comput. Lab., St. Andrews, Fife, Scotland. 50 pp. THE GENERIC NAME OF THE NORTH AMERICAN MUSK TURTLES Hobart M. Smith^ and Kenneth R. Larsen^ Abstract. — Although several recent workers have used the name Sterno- thaerus for the North American musk turtles, the rules of priority render Ster- notherus the correct name for them. The generic type fixation by Stejneger of 5. odoratus is acceprted. A number of recent regional as well as more general works (e.g., Cochran and Coin, 1970:137-139) use the generic name Sternothae- rus for the North American musk turtles commonly bearing the name Sternotherus. Use of the former generic name in recent years stems from the excellent revision by Tinkle (1958) of the S. carina- tus complex, wherein (p. 51) the generic name Sternotherus is held to be a misspelling that should be replaced by Sternothaerus. As pointed out by Albrecht (1967:82), the latter spelling actually does not supersede Sternotherus. It is true that Bell is credited {''Sternotherus, Bell, Mss.") by Gray (1825:211) as the source of Sternotherus in the first appear- ance oJF either name and that accordingly perhaps Bell should now be regarded as the author of the name. The wording of the descrip- tion is, however, completely different from that of Bell's subsequently published Sternothaerus (Bell, 1826:305), and Gray employs the first person form in commenting on the genus: "Cuvier describes the anterior and posterior lobes of the sternum of these species to be moveable; but the hinder was fixed on the specimens which I have examined, which were all dry." Thus the description appears to be in Gray's words, even though he clearly may have drawn the idea for the genus from Bell's MS. The International Code of Zoological Nomenclature (1964), Ar- ticle 50, specifies, in this context, that "The author ... of a scientific name is . . . the person . . . who first publishes it ... in a way that satisfies the criteria of availability . . . , unless it is clear from the contents of the publication that . . . some other person ... is alone responsible both for the name and the conditions that make it avail- able." Clearly Gray's account is not a copy of Bell's, and accordingly it can only be concluded that Bell did not provide the description in the form in which it appeared in Gray's work, even though he may have provided the name or a basis for the name. Reluctantly, but seemingly incontrovertibly, the conclusion is that, by present rules, Gray must be regarded as the author of Sternotherus. Bell remains author of his name Sternothaerus, based upon a different array of taxa, including one African, one Asian, and two North American nominal species. ^Department of Environmental, Population, and Organismic Biology, University of Colorado, Boulder 80302. 42 March 1974 smith, larsen: turtles 43 The name appearing in 1 825 cannot be rendered as Sternothaerus even if attributed to Bell. Although it is true that incorrect original spellings are to be replaced by correct spellings, as pointed out by Tinkle (1958:51), the code severely limits the concept of "incorrect original spellings." In this case, only an inadvertent error could be called incorrect, whereas there is no evidence whatever that Gray did not use the spelling Sternotherus quite deliberately. Article 32 of the code makes it clear that even if Gray did err in transliteration, such an error is not itself "inadvertent." Again there is no escape from the conclusion that Sternotherus is the correct spelling for the 1825 name. Indeed, Gray repeated this spelling in ihis subsequent works, giving no indication of occurrence of error therein. In turn. Article 56 expressly provides that "even if the difference between two genus-group names is due to only one letter, these two names are not to be considered homonjTns." Accordingly Sterno- thaerus of 1826 must be judged on its own merits, as it were, as a name completely independent of and different from Sternotherus of 1825. Although Gray (1825) included a species of Kinosternon in his Sternotherus ("5. pensylvanica" = K. subrubrum), as well as a spe- cies we now recognize in Sternotherus ("5. odorata"), the name Sternotherus has long been accepted as fixed with its type as S. odor- atus, although the earliest selection of this species as type is question- able. It was not selected either directly or indirectly by Bell, as Stejneger (1902:236) maintained. Gray's name could never be ap- plied properly to an African group, since among the originally in- cluded species (to which type selection is limited) there were no African representatives. Stejneger's (1902:237) restriction of type is explicit and acceptable. The alternative would merely make Sternotherus a sjTion^Tn of Kinosternon Spix, 1824 by fixation with K. subrubrum. The t}TDe of Sternot/iaerus Bell could have been restricted subse- quently to any of the four species included by Bell: trifasciatus sp. n. (^Cuora trifasciata), leachianus sp. n. (=iPelusios subniger Lace- pede, 1789), odoratus (=S. odoratus Latreille, 1801), and boscii (Merrem, 1820) {=S. odoratus). Stejneger (1902:237) explicitly des- ignated .9. odoratus as the type, although by specious reasoning (in modern contexts), since he regarded Bell's Sternothaerus as merely a deviant spelling of Gray's Sternotherus and since he regarded Bell's inclusion in Sternothaerus of but one (S. odoratus) of the species Gray included in Sternotherus as fixing the tynpe of both genera as S. odoratus. This is not admissible under the present code, but at this date no reassignment would be useful. Other workers have long accepted Stejneger's explicit type restrictions, and we recommend that this policy be perpetuated. I^ITERATURE CiTED Albrecht, p. W. 1967. The cranial arteries and cranial arterial foramina of the turtle genera Chrysemys, Sternotherus, and Trionyx: A comparative study 44 GREAT BASIN NATURALIST Vol. 34, No. 1 with analysis of possible evolutionary implications. Tulane Studies Zool. 14(3):81-99, figs. 1-3. Bell, T. 1826. A monograph of the tortoises having a moveable sternum, with remarks on their arrangement and affinities. Zool. J. 2:299-310. (The date in 1825 cited by Stejneger is the date of presentation, not of publication.) Cochran, D. M., and C. J. Goin. 1970. The new field book of reptiles and amphibians. G. P. Putnam's Sons, New York. Gray, J. E. 1825. A synopsis of the genera of reptiles and Amphibia, with a description of some new species. Ann. Philos. 10 (2): 193-21 7. Stejneger, L. 1902. Some generic names of turtles. Proc. Biol. Soc. Washing- ton 15:235-238. Tinkle, D. W. 1958. The systematics and ecology of the Sternothaerus carin- atus complex. Tulane Studies Zool. 6(1): 1-56, figs. 1-57. STUDIES ON THE TOLERANCE OF AQUATIC INSECTS TO LOW OXYGEN CONCENTRATIONS' Arden R. Gaufin-, Robert Clubb^, and Robert NewelH Abstract. — Acute, short-term (96-hour) tests were conducted to determine the relative sensitivity of low oxygen concentrations to 20 species of aquatic insects. In addition, the longer-term effects of low oxygen levels on the survival, molting, growth, and emergence of 21 species were studied. This paper en- compasses work conducted at the University of Montana Biological Station from 1968 to 1970 and at the University of Utah from 1966 to 1972. An evaluation of the average minimum dissolved-oxygen requirements of the different groups of aquatic insects tested indicates that the mayflies are the most sensitive, that the stoneflies are next, and that the caddis flies, freshwater shrimp, true flies, and damselflies follow, in that order. While two species of mayfly could tolerate as low a dissolved-oxygen concentration as 3.3 mg/1 for 10 days, a level of 4.6 mg/1 was required for 50-percent survival at 30 days. Fifty percent of the true flies and damselflies tested were able to survive at levels ranging from 2.2 to 2.8 mg/1 for periods ranging from 20 to 92 days. Oxygen is a basic need of aquatic insects, yet information con- cerning exact oxygen requirements is known for but a very few spe- cies. Gaufin and Tarzwell (1956) pointed out that if the oxygen re- quirements of different species of aquatic insects were better known, it should be possible to estimate in retrospect, with considerable ac- curacy, what oxygen levels have existed in a given aquatic environ- ment during the life history of the organisms. Thus, aquatic insects could be used as an excellent index of water quality. The literature is extensive on oxygen consumption by various ani- mals, 3'et such values are meaningful only for the particular condi- tions of measurement. The conditions under which such measure- ments were made are important because the rate of oxygen consump- tion is influenced by several internal and external variables: activity, temperature, nutrition, body size, stage in life cycle, season, time of day, and previous oxygen experience and genetic background (Pros- ser and Brown, 1961). The highest respirator}' rates usually occur in the small, very active forms, whereas the lowest occur in the large, relatively sedentary forms. Wiggles worth (195(3) and Edwards (1946) summarized much of the work done on respiration rates of insects. The majority of the publications on immature aquatic insects has been on European spe- cies. Extensive work on individual, immature aquatic insects was done by Balke (1957) on European species of the orders Neuroptera, Odonata, Plecoptera, and Trichoptera. The difficulty in selecting a suitable and adequate method for the measurement of the respiratory rate in a particular species of aquatic insect was evaluated by Kam- ^This investigation was supported by Research Contract No. 14-12-438, granted by the National Water Quality Laboratory, Federal Water Pollution Control Administration, to the University oJ Montana Biological Station, Bigfork, Montana. 'Professor of Zoologj-, University of Utah and Montana Biological Station. 'Graduate Assistant, University of Utah, Salt Lake City, Utah. ^Graduate Assistant, University of Montana, Biological Station, Bigfork, Montana. 45 46 GREAT BASIN NATURALIST Vol. 34, No. 1 ler (1969). An analysis of the various factors that influence the oxy- gen requirements and respiratory rates of benthic invertebrates is presented by Hynes (1970). The oxygen consumption of 10 of the most common species of stonefly of the western United States and the factors that modify their metabolic rate are discussed by Knight and Gaufin (1966). The oxygen requirements of immature aquatic insects in relation to their classification as index organisms are thoroughly evaluated by Olson and Rueger (1968). Their statistical analyses of oxygen-consumption rates by 12 representative species of aquatic insects of the upper Great Lakes Region constitute very valu- able data for establishing water-quality criteria for the protection of aquatic life. The principal objectives of the studies presented in this report were to determine the oxygen requirements of representative species of aquatic insects of the Intermountain Region and to determine their relative sensitivity to low oxygen concentrations. Oxygen levels necessary for survival and the long-term effects of low oxygen concentrations on molting, growth rates, times of emergence, and behavior patterns were investigated. This report summarizes the results of acute, short-term (96-hour) tests (TLm^'"') used in screening 20 species of aquatic insects to de- termine their relative sensitivity to low oxygen concentrations. In addition, the longer-term effects of low oxygen levels on the survival, molting, growth, time of emergence, and behavior patterns of 21 species are considered. The 96-hour TLm (Standard Methods, 1965) was used as a measure of survival in the tests. This report en- compasses work conducted at the University of Montana Biological Station from 1968 to 1970 and at the University of Utah from 1966 to 1972. Materials and Methods The organisms used in the tests were all insects, except for one species of Amphipoda. All organisms were collected from streams and ponds in northwestern Montana and northern Utah. The organ- isms for a test were all collected from the same area at the same time. The specimens were kept in well oxygenated holding tanks for three days prior to testing. Only specimens of the same age group were utilized. These were generally of the oldest year class present. Test procedures were those outlined in Standard Methods (1965). Deoxygenated water was obtained from degassing equipment as described by Mount (1964). Modifications included a cooling system and an oxygen "ladder." The ladder is constructed of single pane glass cemented with silicone. The ladder is 5 1/2 feet long, 7 inches wide, and 7 inches deep. It is divided into 15 compartments, each separated by a glass partition 2 inches high. The remainder of the divider is composed of fiberglass screen with a 1 mm mesh opening. The deoxygenated water comes from the degasser through plastic tubing, passes through the cooler and then enters the elevated end of the ladder. As the water flows over the 2-inch compartment dividers toward the lower end, its oxygen content increases. Rates of increase March 1974 gaufin, et al: aquatic insects 47 are dependent upon rate of inflow and angle of inclination of the ladder. At an inclination of 40 degrees from the horizontal and at a flow rate of 1000 cc/min, the oxygen increase per chamber is about 0.5mg/l at IOC. Ten organisms were placed in each of seven test chambers and observed twice daily. Point of death was determined by lack of re- sponse when stimulated. Small rocks were placed in the test chambers to which the organisms could cling. The flow rate was checked weekly and was found to vary ±25 cc/min. The temperature was taken daily with a pocket thermometer and was found to vary ±0.5 C. Oxygen concentration was taken daily using the modified Winkler method, utilizing a 50 ml sample. Variations of plus or minus 0.2 mg/1 occurred. Water used in the tests at the Biological Station was unchlorinat- ed well water with the following chemical composition: pH 7.8; total hardness, 135 mg/1; temperature, 6.4 C; turbidity, 0-5 J. T. U.; car- bon dioxide, 1-2 mg/1. Short-term (Acute) Bioassays Conducted at University OF Montana Biological Station Results Nineteen species of aquatic insect and one species of Amphipoda were studied to determine their 96-hour median tolerance limit (TLm). Eight species of Plecoptera were tested. The mean TLm for this group was 3.04 mg/1 of oxygen. Acroneuria pacifica Banks had the lowest TLm, 1.6 mg/1 at a flow rate of 1000 cc/min (Table 1). The highest TLm was obtained with Pteronarcys californica Newport (3.9 mg/1) at a rate of 500 cc/min. The TLm for this species de- creased to 3.2 mg/1 at a flow of 1000 cc/min. All of the specimens of Arcynopteryx parallela Prison survived at oxygen concentrations of from 2 to 5 mg/1 at a flow of 1000 cc/min. All of the test species were stream forms. Four species of mayfly (Ephemeroptera) were examined. Two species were lotic forms, Hexagenia limbata Guerin and Callibaetis montanus (Eaton). Their TLm's were 1.8 mg/1 and 4.4 mg/1 respec- tively. The lentic forms tested were Ephemerella doddsi Needham and Ephemerella grandis Eaton, with D. O. values of 5.2 mg/1 and 3.0 mg/1 respectively. The mean for the group was 3.6 mg/1. Seven species of Trichoptera were tested, and all were from lentic environments. Several of these organisms could not be identified to the species level. Ninety percent of the specimens of Brachycentrus occidental is Banks survived at oxygen concentrations of 2-4 mg/1 and at a flow rate of 500 jcc/min. Neothremma alicia Banks, a small spe- cies (5 mm), had the lowest TLm of 1.7 mg/1. Neophylax sp. had the highest TLm of 3.8 mg/1. The mean for the entire group was 2.86 mg/1. One Dipteran was tested {Simulium vittatum Zetterstadt) and had a TLm of 3.2 mg/1. One Amphipoda was examined {Gammarus 48 GREAT BASIN NATURALIST Vol. 34, No. 1 Table 1. Test organisms, TLm in mg/1, percent saturation and water flow in cc/min. Organisms TLm Saturation Plow Plecoptera Acroneuria pacifica Banks 1.6 14 1000 Arcynopteryx aurea Smith 3.3 29 1000 Arcynopteryx parallela Prison 100* 2-5 mg/1 1000 Diura knowltoni (Prison) 3.6 32 500 Nemoura cinctipes Banks 3.3 29 1000 Pteronarcys californica Newport 3.9 34 500 Pteronarcys californica Newport 3.2 28 1000 Pteronarcella badia (Hagen) 2.4 21 1000 Ephemeroptera Callibaetis montanus Eaton 4.4 38 500 Ephemerella doddsi Needham 5.2 46 500 Ephemerella grandis Eaton 3.0 27 1000 Hexagenia limbata Guerin 1.8 15 1000 Trichoptera Brachycentrus occidentalis Banks 90* 2-4 mg/1 500 Drusinus sp. 1.8 15 1000 Hydropsyche sp. 3.6 32 500 Lepidostoma sp. 80* 3-4 mg/1 1000 Limnephilus ornatus Banks 3.4 30 500 Neophylax sp. 3.8 33 500 Neothremma alicia Banks 1.7 14 500 Diptera Simulium vittatum Zetterstadt 3.2 28 500 Amphipoda Gammarus limnaeus Smith 80* 3 mg/1 500 * Percentage of survival limnaeus Smith) with a survival of 80 percent at 3 mg/1 of oxygen and at a flow rate of 500 mg/1. The mean TLm for all organisms tested was 3.1 mg/1. The mean for all organisms tested at a flow of 1000 cc/min was 2.55 mg/1 and 3.64 mg/1 at a flow of 500 cc/min. The lowest TLm recorded was 1.6 mg/1 for Acroneuria pacifica, or 14-percent oxygen saturation. The highest TLm was 5.2 mg/1 for Ephemerella doddsi, or 46-per- cent oxygen saturation. Discussion Of the organisms tested, the group most tolerant to low dissolved- oxygen (D.O.) values was the lYichoptera (2.86 mg/1). All of the Trichoptera tested, except Hydropsyche, were cased forms, and all came from lentic environments. All the organisms except Drusinus sp. were tested at a flow rate of 500 cc/min. Higher flow rates would probably reduce the TLm of many of the forms. Acroneuria pacifica, a predacious stonefly, was the most re- sistant form tested with a TLm of 1.6 mg/1 (14-percent saturation). The largest organism tested, Pteronarcys californica, showed a de- crease in TLm as the flow rate increased (3.9 mg/1 to 3.2 mg/1). March 1974 gaufin, et al: aquatic insects 49 The mayfly, Ephcmcrella doddsi, had the highest TLm of 5.2 mg/1 (46-perccnt saturation) at 500 cc/niin. This species is found attached to rocks in fast streams. It has been shown by Knight and Gaufin (1963, 1964) that rate of water flow is very important in determining tolerance limits. And that conclusion is supported by the flow-rate ranges and means for Ptcronarcys californica observed in this study. The TLm range for 11 species tested at 500 cc/min was 1.7 mg/1 to 5.2 mg/1, with a mean of 3.64 mg/1. At 1000 cc/min, the range for 10 species was 1.6 mg/1 to 3.3 mg/1 with a mean of 2.55 mg/1, a substantially lower value. Behavior of organisms during testing was of interest. All of the Plecoptera initiated "push-up" movements upon introduction to the test chambers. Most species ceased this motion after several hours but Pteronarcys californica continued these movements periodically throughout the test. Pteronarcys californica also assumed a position half out of the water in the low oxygen chambers. Nemoura cinctipes assumed a stilted position upon death. Number of gill beats per unit time was indicative of oxygen con- centration. Gill beats in Ephemerella grandis were counted after 12 hours in the test chambers and results are given in Table 2. Each value is the mean number of beats for the ten organisms in each chamber. Except at the lowest D.O. concentration, the gill beat decreased as the oxygen increased. The rhythm of gill beats also became erratic as the oxygen increased. The high TLm of the pond mayfly, Callibaetis montanus, was surprising. It had the second highest TLm of all species tested (4.4 mg/1). Another lotic species Hexagenia limbata had a low TLm of 1.8 mg/1. This could probably be explained by its acclimation to lower oxygen concentrations in its normal environment. In response to low oxygen values, the Trichoptera undulated their abdomens in their cases. Simulium vittatum congregated on the chamber walls where the flow was the greatest. Gammarus limnaeus showed no behavioral response to the low oxygen values. Long-term Bioassays Conducted at the University of Montana Biological Station AND THE University of Utah Results Eight species of aquatic insects from northwestern Montana were studied to determine their tolerance levels and behavior patterns when exposed to low oxygen levels over periods of time longer than 96 hours. Five of these species, and an additional 13 species from northern Utah, were also tested for periods of time ranging from 4 to 104 days to determine their long-term reactions (Tables 3, 4, 5). The results of the longer-term bioassays clearly indicate increased sensitivity and mortality of test specimens with increased length 50 GREAT BASIN NATURALIST Vol. 34, No. 1 Table 2. Gill beats/minute for Ephemerella grandis Eaton. Oxygen cone, (mg/l) Beats Rhythm 2.4 3.0 3.6 4.6 5.0 6.0 176 192 192 184 160 100 steady steady steady erratic erratic erratic Table 3. Long-term dissolved-oxygen bioassays conducted at University of Montana Biological Station. Species Minimum D.O. level (mg/l) Percentage of Survival Survival time (days) Plecoptera Pteronarcella badia (Hagen) Pteronarcys californica Newport Arcynopteryx aurea Smith Acroneuria pacifica Banks 4.4 4.8 4.8 5.8 50 40 30 50 69 97 12 111 Ephemeroptera Ephemerella grandis Eaton 4.6 30 30 Trichoptera Brachycentrus occidentalis Bank; Hydropsyche sp. 3 3.2 4.8 50 30 120 50 DiPTERA Atherix variegata Walker 2.4 90 40 Amphipoda Gammarus limnaeus Smith Flow rate of 1000 cc/min 2.8 50 20 of exposure to low oxygen levels. For example, while 50 percent of the specimens of Acroneuria pacifica in Montana survived an oxygen concentration of 1.6 mg/l for 4 days, the minimal dissolved-oxygen level for 50-percent survival at 111 days was 5.8 mg/l. Similarly, 50 percent of the specimens of Arcynopteryx aurea survived in an oxy- gen concentration of 3.3 mg/l for 4 days, but only 30 percent sur- vived at a dissolved-oxygen level of 4.8 mg/l for 12 days. This in- creased sensitivity can be explained partly on the basis of fungus in- fection and debilitation caused by lack of food. For example, 60 per- cent of the larvae of the crane fly, Holorusia sp., survived for 86 days at a dissolved oxygen level of only 2.0 mg/l. Infection with fun- gus and shrinkage of the larvae bodies owing to starvation caused a rapid die-off after 86 days. Of the eight species of aquatic insects tested at the Biological Sta- tion, the carnivorous stonefly, Acroneuria pacifica, had the highest TLm, with a 50-percent death rate at an oxygen level of 5.8 mg/l for 111 days. The most tolerant species was the Dipteran, Atherix variegata, with 90 percent of the specimens surviving for 40 days at March 1974 GAUFIN, ET AL: AQUATIC INSECTS 51 Table 4. Long-term dissolved-oxygen bioassays conducted at the University of Utah (50-percent-plus survival). i Species Minimum D.O. level (mg/1) Percentage of Survival Survival time (days) Plecoptera Acroneuria pacifica Banks Brachyptera nigripennis (Banks) Isoperla fulva Claassen 3.0 2.3 2.3 50 60 50 24 4 13 Ephemeroptera Ephemerella grandis Eaton Rhithrogena rohusta Dodds Trichoptera 3.3 3.3 3.3 50 50 50 18 7 4 Brachycentrus occidentalis Banks Rhyacophila sp. Arctopsyche grandis (Banks) Parapsyche elsis Milne 2.6 1.4 3.4 5.2 80 50 50 60 91 45 26 30 Diptera Atherix variegata Walker Holorusia sp. 2.4 2.0 90 60 97 86 Odonata Argia vivida Hagen Enallagma anna Williamson Flow rate of 1000 cc/min 3.0 1.4 50 50 56 21 Table 5. Long-term dissolved-oxygen bioassays conducted at the University of Utah (minimum D.O. with survival). Species Minimum D.O. level (mg/l) Percentage of Survival Survival time (days) Plecoptera Acroneuria pacifica Banks Arcynopteryx parallela Prison Brachyptera nigripennis (Banks) Isoperla fulva Claassen Pteronarcella badia (Hagen) Ephem eroptera Baetis hicaudatus Dodds Ephemerella grandis Eaton Trichoptera Parapsyche elsis Milne Diptera Atherix variegata Walker Bibiocephala sp. Odonata Argia vivida Hagen Enallagma anna Williamson Flow rate of 1000 cc/min 3.0 20 41 3.4 10 8 4.2 20 28 3.7 20 9 2.1 10 27 2.0 30 30 3.8 10 3 3.5 50 21 4.8 1.7 3.4 1.7 1.1 40 70 40 10 20 16 90 21 100 35 52 GREAT BASIN NATURALIST Vol. 34, No. 1 Table 6. Average minimum dissolved-oxygen requirements of different groups of aquatic invertebrates*. Montana species Average survival (days) Utah species Average survival (days) Plecoptera Ephemeroptera Trichoptera Diptera Odonata Amphipoda 4.9 mg/1 62 2.8 mg/1 14 4.6 mg/1 30 3.3 mg/1 10 4.0 mg/1 85 3.1 mg/1 48 2.4 mg/1 40 2.2 mg/1 92 2.2 mg/1 39 2.8 mg/1 20 'Averages based on 50-percent-plus survival for time indicated. an oxygen concentration of 2.4 mg/1. This species was also the most tolerant of the Utah forms listed, 90 percent of the specimens surviv- ing at the same oxygen level for 97 days. The higher oxygen re- quirement of Acroneuria pacifica under long-term conditions may be partially owing to its food requirements. Inasmuch as this species is carnivorous, lack of a varied animal diet may have reduced its ability to tolerate low oxygen levels for extended periods of time. A comparison of the long-term median tolerance limits of the same species of aquatic insects from Montana and Utah shows con- siderable variation. Fifty percent of the specimens of the stonefly, Acroneuria pacifica, from Montana died at a dissolved-oxygen level of 4.4 mg/1 in 69 days. The same percentage of Utah specimens sur- vived at a much lower dissolved-oxygen concentration, 3.0 mg/1, but for only 24 days. A mayfly, Ephemerella grandis, was tested from both Montana and Utah with similar results. Thirty percent of the Montana specimens survived at a dissolved-oxygen level of 4.6 mg/1 for 30 days, while 50 percent of the Utah specimens survived at a dissolved-oxygen concentration of 3.3 mg/1, but for only 18 days. The differences in tolerance limits between the same species may have been much less if the tests had been conducted under exactly the same conditions in the two locations. Time did not permit this, so the Utah tests were run at lower oxygen levels to determine maximum surviv- al rates at these much lower oxygen limits. An evaluation of the average minimum dissolved-oxygen re- quirements of the different groups of aquatic invertebrates tested shows that the mayflies are the most sensitive, that the stoneflies are next, and that the caddis flies, freshwater shrimp, true flies, and damselflies follow, in that order. While two species of mayfly could tolerate as low a dissolved-oxygen concentration as 3.3 mg/1 for 10 days, a level of 4.6 mg/1 was required for 50-percent survival at 30 days. Three species of stonefly from Utah survived at a dissolved- oxygen concentration of 2.8 mg/1 for 14 days with 50 percent sur- viving, but an average oxygen concentration of 4.9 mg/1 was re- quired for 30- to 50-percent survival for 62 days. Tests on caddis flies also indicated that higher oxygen levels were necessary with longer exposure, a minimum of 4.() mg/1 being required for 50- percent survival for 84 days. March 1974 GAUFIN, ET AL: AQUATIC INSECTS 53 100^ 80. D.O. mg/I Fig. 1. Acroneuria pacifica: 1000 cc/min, 96-hour TLm re- sults, oxygen. '\/ Fig. 2. Arcynopteryx aurea: 1000 cc/min. 96-hour TLm re- sults, oxygen. T~T T IUU.J 80 60. 1.0 l Fig. 3. Arcynopteryx paral- lela: 1000 cc/min, 96-hour TLm results, oxygen. Fig. 4. Pteronarcella badia: 1000 cc/min, 96-hour TLm re- sults, oxygen. 80 60. 0.0. n>9/' Fig. 5. Pteronarcys calif or - nica: 1000 cc/min. 96-hour TLm results, oxj-gen. 0.0. mg/I / Fig. 6. Nemoura cinctipes: 1000 cc/min, 96-hour TLm re- sults, oxygen. 100.. : 60 J y- 0.0. I mg/l Fig. 7. Hexagenia limbata: 1000 cc/min, 96-hour TLm re- sults, oxygen. Fig. 8. Ephemerella grandis: 1000 cc/min, 96-hour TLm re- sults, oxygen. 54 GREAT BASIN NATURALIST Vol. 34, No. 1 0.0. mg/l 1 I 5" Fig. 9. Lepidostoma sp.: 1000 cc/min, 96-hour TLm results, oxygen. ^\ / o.c. mg/I 1 I i k 5 6 7 Fig. 10. Drusinus sp.: 1000 cc/min, 96-hour TLm results, oxygen. 8o! ^ 60. I liO. / i 20- / D.O. mg/I Fig. 11. Pteronarcys calif or- nica: 500 cc/min, 96-hour TLm results, oxygen. Fig. 12. Diura knowltoni: 500 cc/min, 96-hour TLm re- sults, oxygen. // / 0.0. I mg/I T Fig. 13. Ephemerella doddsi: 500 cc/min, 96-hour TLm re- sults, oxygen. Fig. 14. Callibaetis montan- us: 500 cc/min, 96-hour TLm results, oxygen. D.O. mg/l "1 T Fig. 15. Limnephilus sp.: 500 cc/min, 96-hour TLm results, oxygen. Fig. 16. Hydropsyche sp.: 500 cc/min, 96-hour TLm re- sults, oxygen. March 1974 GAUFIN, ET AL: AQUATIC INSECTS 55 lOO. so- so. I|0 0.0. 1 mq/l Fig. 17. Neothremma sp.: 500 cc/min. 96-hour TLni re- sults, o.xygen. D.O. mg/I Fig. 18. Neophylax sp.: 500 cc/min, 96-hour TLm results, oxygen. 100, 80. 60. dOJ /" 0.0. Fig. 19. Brachycentrus occi- dentalis: 500 cc/min, 96-hour TLm results, oxygen. D.O. I mg/1 T T" Fig. 20. Simulium vittatum: 500 /cc min, 96- hour TLm re- sults, oxygen. 100. 80 6a W 0.0. 1 mg/l Fig. 21. Gammarus limnaeus: 500 cc/min, 96-hour TLm re- sults, oxygen. 56 GREAT BASIN NATURALIST Vol. 34, No. 1 The true flies, freshwater shrimp, and damselflies displayed a much greater tolerance to low oxygen levels than did the previous three groups. Fifty percent of the specimens of these three groups were able to survive for periods ranging from 20 to 92 days at dissolved-oxygen levels ranging from 2.2 to 2.8 mg/1. While the principal objective of this project was to determine the minimal dissolved-oxygen levels required for both short- and long-term exposure, survival without growth and metamorphosis would eventually eliminate a species of aquatic insect. While not all of the species tested molted or emerged during the study, many species did. All of the species on which bioassays were run for over 30 days molted one or more times at the oxygen levels required for 50-percent survival. Species such as the stoneflies Brachyptera nigripennis, Pteronarcys californica, and Pteronarcella badia, the mayfly Ephemerella grandis. and the damselfly Enallagma anna emerged during the tests at oxygen concentrations of 4.8 mg/1 or less. None of the caddis flies or Dipterans emerged, inasmuch as only larvae and not pupae were used for testing purposes. Discussion Aquatic environments in which dissolved oxygen is available in excess at all times are rare. Many aquatic animals possess varied adaptations that facilitate the acquisition of oxygen when it be- comes scarce. In stoneflies, diffusion, along with special ventilation mechanisms, provides extensive absorbing surfaces for the absorption of oxygen from the environment. An adaption utilized by the nymphs of Pteronarcys californica when environmental oxygen be- comes reduced is undulating the body — to destroy the oxygen gradient that develops around the body and gills. Of particular in- terest is the variation in rate of undulation according to year class. In studies conducted at the University of Utah from 1963 to 1965, the undulations of the smaller nymphs of this species (year I, 17-18 mm long) were more rapid than those of the larger (year II, 30 mm long). The respiratory mechanism possessed by different species of aquatic insects greatly influences their ability to withstand low oxygen concentrations. In work conducted by Knight and Gaufin (1966) at the University of Utah, the value of gills in enabling some species to better withstand low dissolved-oxygen levels was demonstrated clearly. The nymphs of Pteronarcella badia, Isoperla fulva, and Acroneuria pacifica were all exposed to an environment of reduced dissolved oxygen of 1.0 cc/1 and water flow of 0.004 feet/second, at 10 C. The forms possessing gills exhibited quite similar mortalities during the exposure period. Pteronarcella badia nymphs exhibited a 13-percent mortality after 24 hours and 48 hours of exposure, and 29-percent at the end of 72 hours, with no further mortality for the remainder of the exposure period. Acro- neuria pacifica showed the same mortality as did Pteronarcella badia after 72 hours of exposure. After 96 hours exposure. March 1974 gaufin, et al: aquatic insects 57 Acroneuria pacifica displayed a 25-percent mortality. No further mortality was noted for the remainder of the experimental period. Eighty percent of the Isoperla fulva nymphs, a species without gills, died within 24 hours. After 144 hours of exposure, all had suc- cumbed. The increased mortality shown by the Isoperla fulva nymphs may have been owing to their smaller size and the fact that they were year class I, as opposed to year class II in the gilled fornis. Isoperla fulva has only a one-year life cycle, so it was im- possible to compare nymphs of similar size. In view of the above, a second evaluation was carried out com- paring nymphs of Acroneuria pacifica (gills) to those of Arcynop- teryx parallela (no thoracic gills). The nymphs were tested at a temperature of 15.6 C with a water flow of 0.25 feet/second and a dissolved-oxygen concentration of 1.0 cc/1. The nymphs of both species were between 25 and 30 mm in length. In general, the results of this test- — like those of the previous one — indicated that forms lacking gills are more sensitive to reduced dissolved oxygen than are forms possessing gills. No mortality of Acroneuria pacifica n\Tnphs occurred during the experimental period, while nymphs of Arcynopteryx parallela showed 82-percent mortality after 10 hours of exposure, 88.5-percent mortality at the end of 24 hours, and 100-percent mortality after 34 hours. The metabolism of poikilotherms rises wdth temperature about two and one-half times per 10 C change in temperature (Prosser and Brown, 1961). This metabolic increase in response to increased environmental temperature produces increased oxygen consumption. The increase in oxygen consumption with increased water tempera- ture would cause an aquatic insect subjected to the higher tempera- ture (15.6 C) to incur an oxygen debt at a higher dissolved-oxygen concentration than one subjected to a situation similar except for a reduced temperature (10 C). Stoneflies, mayflies, and caddis flies do not have an apparent ability to get along without oxygen for an extended period. They do survive for a short period in greatly reduced oxygen by greatly reducing their activity, and they use energy apparently produced by the anaerobic phase of glycolysis. If the oxygen supply is not restored within a certain time, the speci- mens die from asphyxiation. In the work conducted to date by the author and his colleagues, there has been a great difference in the dissolved-oxygen concen- tration at which initial mortality of test organisms was recorded. This difference was greatly influenced by the temperature difference in the experimental environment. In a natural situation in which the dissolved oxygen is gradually reduced for short periods of time (for example, by intermittent discharges of organic oxygen-demand- ing wastes), the onset of stonefly mortality would be influenced by the existing water temperature. Provided that the water flow and other variables remained constant, one could expect the aquatic insects subjected to an environmental temperature of 10 C to with- stand reduced oxygen concentrations about 2.4 times lower than would the same specimens exposed to similarly reduced oxygen 58 GREAT BASIN NATURALIST Vol. 34, No. 1 concentrations at a water temperature of 15.6 C. In a hypothetical situation based on the work of Knight and Gaufin (1966), a stream possessing a temperature of 15.6 C and a dissolved-oxygen con- centration of 0.6 cc/1 would have a stonefly mortality of 18 percent while a stream similar in all respects except that its water tempera- ture was 10 C would exhibit 100-percent survival. Thus, the water temperature of a stream is a very important factor in the survival of aquatic insects when they are subjected to a reduction in dissolved oxygen over a short period of time. The rate of water flow in a stream also is a very important factor in the survival of aquatic insects exposed to low oxygen concentrations. Knight and Gaufin (1966) showed that a gradual reduction of dissolved oxygen with water flow of 0.06 ft/sec pro- duced approximately 50-percent stonefly mortality and that a similar situation provided with a water flow of 0.25 ft/sec resulted in 100-percent survival. In the present study the mean oxygen concentration required for 50-percent survival by 1 1 species of aquatic insects at a flow rate of 500 cc/min was 3.64 mg/1. The mean for 10 species at a flow rate of 1000 cc/min was considerably lower or 2.55 mg/1. Literature Cited American Public Health Association. 1965. Standard methods for the exam- ination of water and wastewater, 12th ed. New York, N.Y. 769 pp. Balke, E. 1957. Der O2 Konsum und die Tracheen-Innenflache bei durch Tracheen Kiemen atmenden Insekten larven in Abhangig Keit von der Korpergrosse. Z. Vergl. Physiol. 40:415-439. Brinck, Per. 1949. Studies on Swedish stoneflies (Plecoptera). Lund Berlingeka Boktryckeiret, Sweden. DoDDS, G. S., AND F. L. Hisaw. 1924. Ecological studies of aquatic insects. II. Size of respiratory organs in relation to environmental conditions. Ecology. 5:262-271. Edwards, G. A. 1946. The influence of temperature upon the oxygen consump- tion of several arthropods. Jour. Cell. Comp. Physiol. 27:53-60. Fair, G. M., and M. C. Whipple. 1948. Revision of the microscopy of drinking water, 4th ed. John Wiley, New York. Fox, H. M. 1936. Oxygen consumption of mayfly nymphs in relation to available oxygen. Nature (London) Dec. 1015. Gaufin, A. R., and C. M. Tarzwell. 1952. Aquatic invertebrates as indica- tors of stream pollution. Pub. Health Rept. 67:57-64. . 1956. Aquatic macroinvertebrate communities as indicators of organic pollution in Lytle Creek. Sewage and Wastes. 28:906-924. Gaufin, R. F., and A. R. Gaufin. 1961. The effects of low oxygen concentra- tions on stoneflies. Proc. Utah Acad. Sci., Arts, Let. 38:57-64. Hynes, H. B. N. 1960. The Biology of polluted waters. Liverpool University Press. . 1970. The ecology of running waters. Univ. of Toronto Press. 555 p. Kamler, E. 1969. A comparison of the closed-bottle and flowing- water methods for measurement of respiration in aquatic invertebrates. Pol. Arch. Hydrobiol. 16(29) :31-49. Knight, A. W., and A. R. Gaufin. 1963. The effect of water flow, tempera- ture, and oxvgen concentration on the Plecoptera nymph Acroneuria pacifica Banks. Proc.'^Utah Acad. Sci., Arts, Let. 40(2): 175-184. . 1964. Relative importance of varying oxygen concentration, tempera- ture, and water flow on the mechanical activity and survival of the Plecoptera March 1974 gaufin, et al: aquatic insects 59 nymph Pteronarcys californica. Newport. Proc. Utah Acad. Sci., Arts, Let. 41 (1): 14-28. . 1965. Function of stonefly gills under reduced dissolved-oxygen con- centration. Proc. Utah Acad. Sci., Arts, Let. 42 (2): 186-190. 1966. Oxygen consumption of several species of stoneflies (Plecoptera). J. Insect Physiol. 12:347-355. KoLKwiTz, R. AND M. Marsson. 1909. Ockologie der tierschen Saprobien. Int. Revue ges. Hydrobiol. Hydrogr. 2:126-152. LiEBMANN, H. 1951. Handbuch der Frischwasser und Abwasserbiologie. vol. 1. R. Oldenbaurg. Munchen. Madsen, B. L. 1968. The distribution of nymphs of Brachyptera risi Mort. and Nemoura flexuosa Aub. (Plecoptera) in relation to oxygen. Oikos 19:304-310. Mount, D. I. 1964. Additional information on a system for controlling the dissolved oxygen content of water. Trans. Am. Fish Soc, 93(1) : 100-103. Olsen, T. E., and M. E. Rueger. 1968. Relationship of oxygen requirements to index-organism classification of immature aquatic insects. J. Water Poll. Control Fed. 40(5) :R188-R202. Petty, W. C. 1967. Studies on the oxygen requirements of two species of stoneflies (Plecoptera). Unpubl. Master's thesis, Dept. of Zool. Entomol., Univ. of Utah, Salt Lake City, 73 pp. Philipson, G. N. 1954. The effect of water flow and oxygen concentration on six species of caddis fly (Trichoptera) larvae. Proc. Zool. Soc. London (Brit.) 124:547-564. Prosser, C. L., and Brown, F. A., Jr. 1961. Comparative animal physiology, 2nd ed. W. B. Saunders, Philadelphia. Reish, D. J., and T. L. Richards. 1966. A technique for studying the effect of varying concentrations of dissolved oxygen on aquatic organisms. Int. J. Air. Wat. Pollut. 10:69-71. Roeder, K. D. 1953. Insect physiology. John Wiley and Sons, Inc., New York, 1100 pp. WiGGLEswoRTH, V. B. 1950. The principles of insect physiology. E. P. Button and Co., New York, N.Y. SUBSTRATE COLOR MATCHING IN THE GRASSHOPPER, CIRCOTETTIX RABULA (ORIHOPTERA: ACRIDIDAE) George W. Coxi and Darla G. Cox^ Abstract. — Mechanisms important in maintaining substrate color matching in the grasshopper, Circotettix rabula, were studied near Aspen, Colorado, during the summers of 1968-70. Studies concentrated on populations on gray shale and red sandstone substrates. In both areas, collections revealed appreciable numbers of mismatched pheno types among all age groups. The possibility of develop- mental homochromy was examined by observation of nymphs held in rearing boxes on matching and contrasting soil colors. The behavioral selection of matching substrate colors was tested by preference experiments. While not negating the possibility of these mechanisms, results suggested that they were of minor importance. Predation experiments, using Sceloporus lizards, demon- strated significant levels of selective predation on mismatched nymphs on both red and gray substrates. Experiments with bird and mammal predators, using adult grasshoppers, gave similar results. Release-recapture experiments with marked adults in areas of red and gray substrates showed markedly higher disappearance rates for mismatched animals. These results are interpreted to indicate that selective predation on mismatched animals is a major factor in maintaining substrate color matching in this species. Over the past 25 years, students of ecological genetics have documented several major cases of rapid evolutionary response by animal populations subjected to strong selective pressures. The most thoroughly investigated of these concern industrial melanism in moths (Kettlewell, 1961), pesticide resistance in a wide variety of animals (Crow, 1966), and the coloration and banding patterns of various land snails (Cain and Sheppard, 1954). For the Peppered Moth, Bistort betularia. an industrial melanic, and for the land snail, Cepaea nemoralis, selective predation by vertebrates has been demonstrated (by experimentation and direct observation) to be an important factor in the observed evolutionary response (Kettle- well, 1955, 1956; Sheppard, 1951; Cain and Sheppard, 1954; Cain and Currey, 1968). The phenomenon of substrate color matching has been recorded for many species of animals of open habitats and is especially common in Acridid grasshoppers (Rowell, 1971). For these animals, the mechanisms involved appear to be complex and varied, includ- ing— at least for different species — predator selection for crypsis, behavioral preference for color-matching substrates, and homo- chromic responses during individual development (ibid.). Acridid species showing spatial or temporal variation in substrate color matching may therefore be of major value in the study of evolu- tionary processes operating on organisms capable of behavioral or developmental modification of genetically based characteristics. In the Roaring Fork Valley near Aspen, Colorado, populations of many animals of open habitats show differences in general body coloration correlated with soil and rock substrate color. The sub- 'Department of Biology, San Diego State University, San Diego, California 92115. 'Biology Department, La Jolla Country Day School, La Jolla, California 92037. 60 March 1974 cox, cox: acrididae 61 strates involved vary strikingly in color as a result of the variety of geological formations exposed in the valley. The most extensive exposures near Aspen consist of red sandstones of the Maroon For- mation, dark gray shales of tlie Mancos Formation, and Precambrian granites on which yellow to brown clay soils are developed. Smaller areas of dark basaltic lavas and white shales also occur in the lower part of the valley. Although sharp changes in substrate color some- times exist at contacts between different parent materials, the presence of a variety of other bedrock formations and the mixing of various parental materials in glacial and alluvial deposits produce an additional variety of substrates of intermediate colors in different parts of the valley. Correlation of general body color with substrate color has been observed in the lizards Sceloporus undulatus and S. graciosus (Cole- man, 1968), and, by the authors, in various species of Acridid grass- hopi)ers, including Arphia conspersa Scudder, Circotettix rabula Rehn and Hebert, Cratypedes neglectus (Thomas), and Trimero- tropis suffusa (Scudder). During the summers of 1968-70, the authors conducted studies of the role of various factors in the maintenance of substrate color matching in populations of the grasshopper Circotettix rabula, an Acridid of the subfamily Oedipodinae, in the vicinity of Aspen. These studies concentrated on populations on exposures of Mancos Shale (gray) and Maroon Sandstone (red). The Mancos Shale population was studied at a location 6 miles west of Aspen on Colorado State Highway 82 at Brush Creek. The Maroon Sandstone population was studied at an area of cliffs on the north side of the Roaring Fork River opposite the community of Snowmass, about 15 miles west of Aspen. Both study areas consisted of relatively open, dry habitats on hilly terrain, with the dominant vegetation being a complex mixture of sagebrush, oak-serviceberry, pinyon- juniper, and riparian elements. Procedure At the outset, three major mechanisms that, separately or in combination, might account for the pattern of substrate color matching were recognized: 1. Homochromy (the adoption, through developmental process in the nymphs, of a body color matching that of the substrate upon which they are living). This mechanism has been demonstrated in many Acridid grasshoppers (Rowell, 1971). 2. Behavioral preference by individuals for substrates of matching color. Such preferences have been recorded for some Oedipodine grasshoppers (Rowell. 1971). This preference could presumably act in conjunction with either 1 (above) or 3 (below) to enhance the degree of color matching achieved by these mechanisms. 3. Predator selection favoring individuals closely matching the substrate in body color. This mechanism might operate in conjunction with 1 or 2 (above) by selection against individuals in which the mechanisms of homochromy or behavioral preference are imperfectly developed. Al- ternatively, selection may relate to matched and mismatched color pheno- 62 GREAT BASIN NATURALIST Vol. 34, No. 1 types resulting directly from genotypic differences in processes of pigment production. The present studies were designed to investigate the relative im- portance of these possible mechanisms. The hypothesis of developmental adoption of environmental color was tested by a combination of field and experimental studies. Nymphs were collected in both study areas during the period 22-25 June 1968. Total body length and color phenotype were recorded for each of these animals.' These animals were visually assigned to four color phenotype classes: gray, yellow brown, brown, and red. These collections were made to determine the presence and general frequency of mismatched phenotypes in the two areas. In addition, during the June and July of 1968, live nymphs from each study area were divided into two groups of equal size, and placed in separate rea'ring boxes with red and gray soils and correspondingly painted walls. Totals of 54 gray and 52 red nymphs were used in this experiment. Food and water were provided in each box. Although it proved impossible to rear many of these nymphs to adulthood, many completed one or more molts and were maintained for periods of up to four weeks. This experiment was performed to determine if changes in body color could be in- duced in nymphs of the size classes for which data on frequency of various color phenotypes had bsen obtained in the field observations. To investigate hypothesis 2, that of possible behavior preferences for matching substrates, four substrate-color preference tests were conducted during the summer of 1968. Preferences were tested separately for nymphs and adults from the two study areas. Tests were conducted outdoors in 4'x8' boxes covered with gray fiberglass screening, in which equal areas of red and gray soils from the study areas were presented. These tests consisted of a series of 1-hour runs, with tne position of the boxes being reversed between suc- cessive runs. Experimental animals were introduced at the boundary of the two substrate types at the start of each run, and their positions were noted at the end of the one-hour period. Several laboratory' and field experiments were conducted to evaluate the intensity of predation on animals of matched and mis- matched body color. The most extensive series was conducted using the lizards Sceloporus undulatus and S. graciosus as predators on nymphs. These experiments were carried out in the covered 4'x8' boxes described above. A layer of red or gray soil was placed in the bottom of the box, and several piles of flat stones of matching color constructed near the center of the box to allow shelter for both liz- ards and grasshopper 'nymphs. This was necessary to minimize the tendency for animals to climb onto the walls and covers of the boxes (which were of different color). Equal numbers of red and gray nymphs were then placed in the box and lizards introduced so that a ratio of 6-8 nymphs per lizard was obtained. These experiments were terminated when approximately one half of the nymphs had been eaten. During the summer of 1968 experiments were conducted on gray substrates. Experiments on red substrates were performed March 1974 cox, cox: acrididae 63 during all three summers to obtain an adequate number of observa- tions. Experiments were carried out in a shaded, outdoor area during 1968, with the duration of each run being 24 hours. In 1969 and 1970, experiments were done indoors under artificial lights. The duration of runs in these experiments varied from 6-12 hours. During the summer of 1969 two series of experiments were con- ducted using the least chipmunk, Eutamias minimus, and the Ameri- can robin, Turdus migratorius, as predators. For these experiments, adult grasshoppers were used. Experiments using the least chipmunk (Hobbs, 1969) were conducted in the 4'x8' box described above. The flight wings and hind legs of the grasshoppers used in these tests were removed to minimize the effects of flight and rapid movement in attracting attention of the predators. Two grasshoppers, one red and one gray, were placed in the box on a given substrate color, and the predator introduced. The grasshopper first captured was then noted. A total of 25 such tests were performed on each sub- strate color. A similar procedure was followed in experiments using the American robin (Coate, 1969), except that tests were performed in a cage 6'x6'x6' in size. Two field experiments were also conducted to compare the rates of disappearance of red and gray adults following their release in an area of one substrate color. The animals used in these experiments were captured in the two study areas at locations about one-half mile from the locations at which releases were subsequently made. Ani- mals were marked with a spot of colored airplane dope on the ven- tral side of the thorax and were released on the same date on which they were captured. During the summer of 1968 two releases were made on a Man- cos Shale (gray) area. On 24-26 July, 37 males and 34 females from the Mancos Shale population and 54 males and 50 females from the Maroon Sandstone population were released. The second release, made on 6 August, consisted of 14 males and 13 females from the Mancos Shale population and 15 males and 30 females from the Maroon Sandstone population. Recapture samples were taken on 1-3 August. 15-16 August, and 21 August. During the summer of 1969, releases were made on an area of Maroon Sandstone (red) substrate. On 16-17 July, 65 males and 53 females from the red sandstone population and 62 males and 55 females from the gray shale population were released. Recapture samples were obtained on 24 July, 29-31 July, and 7-8 August. Results Totals of 48 n}Tnphs from the Mancos Shale population and 70 n\Tnphs from the Maroon Sandstone population were obtained in the field collections made in June 1968 (Table 1). The distribution of these animals among the four color phenotype classes was sum- marized separately for animals less than 12 mm and for those 12 mm or over in total length. These data indicate that an appreciable frequency of mismatched color phenotypes exists, especially among 64 GREAT BASIN NATURALIST Vol. 34, No. 1 Table 1. Frequencies of nymphs of different color in collections from Mancos Shale (gray) and Maroon Sandstone (red) areas, 22-25 June 1968. Color Phenotype Class (Visual Groupings) Size Yellow Light-Dark Substrate Class Gray Brown Brown Red Total Mancos Shale <12 nmi >12 mm Total Maroon Sandstone <12 mm >12 mm Total 18 1 2 5 26 21 0 0 1 22 39 1 2 6 48 0 0 16 12 28 0 2 9 31 42 25 43 70 the smaller nymphs, in both populations. In the Mancos Shale area, for example, 5 of the 25 nymphs less than 12 mm in total length were red. When analyzed by a 2x2 contingency test, the frequencies of matched and totally mismatched phenotypes in the two size groups were found to be significantly different, or nearly so. For the Mancos Shale population the calculated chi square value of 3.80 was nearly significant at the 5 percent level; for the Maroon Sand- stone population, the value of 5.55 was significant at the 5 percent level (5 percent critical chi square, with 1 d.f., equals 3.841). In both cases, the frequency of mismatched individuals was lower among the larger nymphs. The nymphs maintained in rearing boxes on substrates of con- trasting color, however, showed no visually detectable changes in body color, as compared with those kept on matching substrates. In the four experiments designed to test for substrate-color pref- erence by nymphs and adults, the total numbers of preference re- sponses obtained per experiment varied from 58 to 96. However, chi square goodness-of-fit tests, comparing observed results and an ex- pectation of equal numbers selecting each substrate, showed no sig- nificant preferences for substrate color by nymphs or adults from either population. Results of the predation experiments with the lizards Sceloporus undulatus and S. graciosus demonstrated selective predation on nymphs of mismatched color (Table 2). On the gray substrate a 2x2 contingency test of the experimental results showed that the numbers of nymphs eaten and not eaten were significantly influenced by body color (1 percent level); a similar test for the red substrate showed significance at the 5 percent level. Fewer data were obtained for the experiments using the Ameri- can robin and least chipmunk as predators (Table 3). Except for the results of the tests with the least chipmunk on the gray substrate (significant at the 5 percent level) these results do not show signifi- cantly different preferences for matched and mismatched grasshop- pers. However, the pattern shown by these data is similar to that ob- tained for the lizard predation experiments. A common feature of experiments with all of the predators used was a higher intensity of March 1974 cox, COX: ACRIDIDAE 65 (0 ti o 05 « u u Xi CO 3 01 (0 o 2 o w 3 >. c 3 c CT ■^ >> (0 U) -d « 4) u o J3 tfO H O 0.(/) aj cue ^ ^ W J3 to 3 (0 w ^ !« P ^-^ 4) I^ '3'S 3 <0 (C C bi ^- *« <« s_ -p re o Ic s«o H u ^ c « 3 S — 3 0) a 3-3 to 3 ""0 *c -C CO C . to -c « .^^ (/) a, u o J3 (0 ceo H o 66 GREAT BASIN NATURALIST Vol. 34, No. 1 predation on mismatched animals on the gray substrate than on the red. Data from the release and recapture experiments wdth marked adults on both substrates showed rapid decline in the ratio of mis- matched to matched animals with time (Figs. 1, 2). To evaluate the significance of this change, Chi Square tests were performed on the observed frequencies of red and gray animals in the first recap- ture sample obtained after each release, using the release ratio of red and gray animals to provide the expected frequencies. For the 24-26 July release and the 1-3 August recapture samples on the Mancos Shale area (1968), the calculated Chi Square value was 8.74. For the 6 August release and the 15-16 August recapture sample, the calculated Chi Square value was 8.78. Both of these tests indicate a highly significant (1 percent level) change in the ratio of red to gray animals. For the 16-17 July release and the 24 July recapture sample on the Maroon Sandstone area (1969), the calculated Chi Square value was 4.66 (significant at the 5 percent level). The ratio of mismatched to matched animals (Figs. 1, 2) declined exponentially, according to the formula Rt = Roe"' Where: Ro = Ratio on date of release Rt = Ratio t days after release b = Rate of change in mismatched/matched ratio per day. A rough estimate of the average rate of change, b, in this ratio was 1.8- 1.6- 1.4- 54 m 37m 50f R 34f G 15m30fR 14m13fG « R-Red GGray m-Male f- Female ^1.2: ^1.0- Q ^0.8- \ 0.6- 0.4- 0.2- \ 5m3fR N. 7m11f17G \ 3m1f1?R ^, 7m8f1'G ». ~^~-, ImR UR ~~--.^2m3fG --^^_6tG n n ^""~~~'""-~--_''"^ 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1^ T 1 25 30 1 JULY 10 AUGUST 15 20 Fig. 1. Ratios of red (Maroon Sandstone) to gray (Mancos Shale) adults in the release-recapture experiment on the Mancos Shale study area. March 1974 cox, COX: ACRIDIDAE 67 Q H Pi I 62m 55f G 65m 53 f R R-Red G - Gray m - Male f - Female Im 2f G 8m 9f R 3m 4f R I I 10 JULY AUGUST Fig. 2. Ratios of gray (Mancos Shale) to red (Maroon Sandstone) adults in the release-recapture experiment on the Maroon Sandstone study area. obtained by calculating the least square linear regression of In R on t. The slope of this relationship is a measure of b. Values for the rate of change in the red-gray ratio during the experiments initiated in July and August of 1968 on the Mancos Shale area were -0.10 and -0.14, respectively. These values indicate that the disappearance rate of the red animals was of the order of 10 to 14 individuals per 100 individuals per day greater than that of the gray animals. On the Maroon Sandstone area, the differential rate of disappearance of the gray animals, calculated in a similar manner, was -0.09, or 9 individuals per day greater than that of the red animals. Recapture data from the field experiments were also analyzed with respect to frequencies of recapture of individuals of different sex. For the experiment on the Maroon Sandstone area, and for red animals in the Mancos Shale experiment no significant differences were noted between the sex ratios of animals released and those re- captured. However, for gray animals used in the Mancos Shale ex- periment, the sex ratio of the combined groups of released animals (24-26 July and 6 August 1968) was significantly different (Chi Square r= 4.26, 5 percent critical value = 3.841) from that of the combined recapture samples for these releases. Males were less fre- quently recaptured than expected in this case. The males of this species perform a conspicuous aerial display, consisting of a pro- longed dancing flight accompanied by a loud crackling call. This display is attractive to predators, and several observations of actual or attempted predation involved attempts by birds to capture such animals in flight. 68 GREAT BASIN NATURALIST Vol. 34, No. 1 Discussion Although the phenomenon of substrate color matching is well developed in Circotettix rahula, variation in body coloration exists in both nymphal (Table 1) and adult (Trent, 1968) populations in the areas examined in this study. Both of the study areas were adjacent to alluvial terraces of the Roaring Fork River and, thus, to substrates of somewhat different color. Furthermore, the adults are strong fliers, and in such situations it is likely that considerable dispersal occurs between areas differing in substrate color. During the sum- mers of 1969 and 1970, direct observations of such dispersal were made near Aspen at a locality in which a whitish gray shale forma- tion sharply contacts an exposure of Maroon Sandstone. The specific locations from which collections of nymphs (Table 1) were made in 1968, however, were at least 100 m from areas of strongly differing substrate color. The occurrence of strongly mis- matched forms among these nymphs, together with the failure of nymphs of comparable size to show modification of body color in the red and gray rearing boxes, suggests that any homochromic de- velopmental response is relatively weak and that body color is to a large extent under direct genetic control. The possibility that artifi- cial light conditions may have been unfavorable for homochromic response cannot be entirely dismissed, however. Because of the low mobility of the nymphs, it is probable that the existence of mis- matched nymphs in the study areas is the result of dispersal of adults into the areas from other substrate types. No evidence of behavioral selection of matching substrates was obtained in the laboratory preference tests. This evidence is further supported by the direct observations, cited above, of dispersal of adults between substrates of sharply differing color. Nevertheless, it is still possible that behavioral selection of matching substrates does occur to some degree, but was inhibited, especially for adidts, by the artificial conditions of the test boxes. Evidence of selective predation is strong. Field observ'ations indi- cate that a variety of vertebrates prey on this species. Observations of either attempted or successful predation in the field were obtained for the sagebrush lizard, Sceloporus graciosus, the least chipmunk, Eutamins minimus, the western tanager, Piranga ludoviciana, the Steller's jay, Cyanocitta stelleri, the mountain bluebird, Sialia cucur- roides, and the red-winged blackbird, Agelaius phoeniceus. Results of the predator selection experiments (Tables 2, 3) indi- cate that differential predation is of major importance in the main- tenance of substrate color matching. These obsersations correlate with the significantly lower frequency of mismatched nymphs in the larger size classes in the field (Table 1 ) and with the signifi- cantly greater rate of disappearance of mismatched adults in the release-recapture experiments. In the release-recapture experiments, as conducted, it is impos- sible to rule out entirely the possibility of differential dispersal away from the release point by matched and mismatched animals. But, March 1974 cox, cox: acrididae 69 the absence, or at least the weakness, of behavioral preferences for matching substrate color suggests that this is unlikely. The recapture data for males and females are also most consistent with the con- clusion that predation is the primary factor in the differential dis- appearance rates of matched and mismatched adults. Males, which perform frequent and extensive aerial displays, should be both the most vulnerable to aerial predators and the most prone to dispersal from the release site. Mismatched males should then have the great- est opportunity for encountering and selecting more appropriate sub- strates. However, the only instance in which the disappearance rate for males significantly exceeded that for females was for gray adults on the gray substrate. If dispersal, combined with behavioral selec- tion of matching substrates, had been of major importance, differ- ences in male and female disappearance rates should have been greatest for mismatched animals. In addition, the experimental predation studies and the release- recapture experiments share the common feature that mismatched animals are removed, or disappear more rapidly, from the gray shale than from the red sandstone substrate. This observation is difficult to interpret in the absence of detailed measurements of spectral re- flectance of the substrates and animals. However, it may be that gray animals on the red substrate, while sharply contrasted in color, approximate in appearance spots of shadow produced by small objects, or perhaps dead twigs, and are thus not as easily recognized by predators. These observations lead us to conclude that predator selection against mismatched animals is the major mechanism for maintain- ing substrate color matching in this species. Acknowledgments This work was supported by Grants 4753 and 5167 from the Penrose Fund, American Philosophical Society. We wish to thank Frederick G. Astin and Dennis Turner for assistance in the field, and Jerry L. Hobbs and Ralph E. Coate for permission to use un- published data on the results of predation tests with the least chip- munk and the American robin, respectively. We also thank Theo- dore J. Cohn for comments and criticism on the study as a whole. References Cain, A. J., and J. D. Currey. 1968. Ecogenetics of a population of Cepaea nemoralis (L.) subject to strong area effects. Trans. Roy. Soc. Lond. B. 253:447-482. , AND P. M. Sheppard. 1954. Natural selection in Cepaea. Genetics 39:89-116. Coate, R. E. 1969. Avian predation on red and gray populations of the grass- hopper Circotettix rabula. Unpublished manuscript, Aspen Inst. Field Ecol. Coleman, P. R. 1968. Adaptation of thermal balance and concealing coloration by three populations of the sagebrush lizard. Sceloporus graciosus graciosus in the Roaring Fork Valley, Colorado. Unpublished manuscript. Aspen Inst. Field Ecol. 70 GREAT BASIN NATURALIST Vol. 34, No. 1 Crow, J. F. 1966. Evolution of resistance in hosts and pests. Pages 263-275 in Scientific aspects of pest control, Pub. No. 1402, National Acad. Sci. — Na- tional Res. Council, Washington. HoBBS, J. L. 1969. An analysis of differential predation by Eutamias minimus on two populations of the grasshopper Circoiettix rabula. Unpublished manuscript. Aspen Inst. Field Ecol. Kettlewell, H. B. D. 1955. Selection experiments on industrial melanism in the hepidoptera. Heredity 9:323-342. . 1956. Further selection experiments on industrial melanism in the hepidoptera. Heredity 10:287-301. 1961. The phenomenon of industrial melanism in the hepidoptera. Ann. Rev. Ent. 6:245-262. RowELL, C. H. F. 1971. The variable coloration of Acridoid grasshoppers. Adv. Insect Physiol. 8:145-198. Sheppard, p. M. 1951. Fluctuations in the selective value of certain phenotypes in the polymorphic land snail Cepaea nemoralis L. Heredity 5:125-134. Trent, D. G. 1968. An analysis of substrate color-matching in two populations of the grasshopper, Circotettiz rabula. Unpublished manuscript. Aspen Inst. Field Ecol. EFFECTS OF CHEMICALS ON THE GERMINATION OF POLLEN GRAINS OF TORENIA ASIATICA LINN. E. M. V. Nambudirii and M. K. Thomas^ Abstract. — Germination studies of the pollen grains of a Scrophulariaceae plant, Torenia asiatica Linn, have revealed that the maximum percentage of germination and longer tubes were attained in 15 percent sucrose solution. Among the chemicals tried, calcium and boron enhanced germination. Less germination occurred among grains treated with potassium or magnesium salts. Abnormalities such as branched tubes with or without vesicles were found in chemically treated pollen grains. Torenia asiatica Linn., a common garden plant belonging to the tribe Gratioleae of the Scrophulariaceae family is cultivated as an ornamental plant in and around Bombay, India. This small her- baceous plant produces bright violet flowers. Pollen morphology of Scrophulariaceae plants has been described by Risch (1939), Erdt- man (1952), Ikuse (1952), Natarajan (1957), and others. Varghese (1968) described the morphology of pollen grains of Torenia cordi- folia Roxb., along with certain other plants of this family. Successful germination of pollen is a prerequisite for success in fertilization; therefore, many workers have studied the effect of different chemicals on the germination of pollen from various plants. Compounds of boron, calcium, potassium, and magnesium were selected for the study of their effects on the germination of the pollen of T. asiatica. Although the effects of these chemicals on germination of pollen grains of certain other plants have already been studied by previous workers, in this paper the authors are con- cerned wdth the study of various concentrations (20, 40, 60, 80, and too ppm) of boric acid (H3BO3), calcium nitrate (Ca[NO,02'4HoO), potassium nitrate (KNO3) and magnesium sulphate (MgS04»7H20) on the germination and tube growth of T. asiatica pollen. Materials and Methods Pollen grains used in the experiments were collected from flowers just prior to their opening to ensure that the material used was fresh and uncontaminated. Pollen grains of T. asiatica (Fig. 4b) show three colpae. Chemicals employed in the experiments were of AR-BDH. Culture media of sucrose (5, 10, 15, and 20 percent) were prepared in double-distilled water. After the sucrose concentration was standarized at 15 percent, different concentrations of four chemicals were added separately to the 15 percent sucrose solution. Concentrations of chemicals chosen were 20, 40, 60, 80, and 100 ppm. One drop of each of these solutions was then placed on a clean sterilized microslide, and pollen grains were dusted on these media. The microslides were then transferred to a moist filter chamber 'P.O. Box 21563, Dar-es-Salaam, Tanzania. -Botany Department, Wilson College, Bombay 400007, India. 71 72 GREAT BASIN NATURALIST Vol. 34, No. 1 where pollen grains were allowed to germinate and grow for two hours. Experiments were conducted at a pH of 6.5 and a tempera- ture of 26 to 28 C. Percentage germination was calculated by counting 100-200 pollen grains from different fields of the micro- slide. Mean tube length was calculated from 50 pollen tubes selected at random. All experiments were repeated until concurrent results were available. Results During initial experiments, it was noticed that grains did not germinate in water but readily germinated in a sugar solution. Therefore, further germination tests were carried out with different types of sugars; namely, glucose, fructose, and sucrose. Sucrose gave maximum germination percentage and higher mean tube length. Hence, for further experiments with different chemicals, sucrose was selected as the basic medium. Pollen germination in sucrose medium. Pollen grains were allowed to germinate in 5, 10, 15, and 20 percent sucrose solutions. The percentages of germination and tube growth in these media were recorded. Maximum germination percentage and tube growth were observed in 15 percent sucrose solution (Table 1, Fig. 1). A minimum percentage germination (10 percent) along with a lower mean tube length (222. 5/x) were observed in 20 percent sucrose solution. A comparatively low germination percentage and shorter tubes were also found in 5 and 10 percent sucrose solutions (Table 1). It is therefore evident that 15 percent sucrose medium can be used for further experiments with different chemicals. Another benefit in selecting this sucrose concentration is that the bursting of pollen tubes in this medium is lower than in other concentrations. Bursting of pollen tubes before attaining required lengths is undesir- able since such tubes cannot effect fertilization. Pollen germination in different chemicals. Keeping 15 percent sucrose solution as the basic medium, different concentrations of the four chemicals were prepared separately. A comparative ac- count of the results obtained is given in Table 2 and in Figure 2. The major effects of different chemicals on germination are given below: 1. Boric acid (Fig. 3b). A minimum concentration of boric acid enhanced pollen germination and tube length. Eighty-nine percent Table 1. Percentage germination and mean tube length of pollen grains of Torenia asiatica Linn. Percentage concentration of sucrose Mean tube length Percentage in microns after germination two hours 43 290.0 55 351.0 73 443.5 10 222.5 5 10 15 20 March 1974 lOOOr 900 NAMBUDIRI, THOMAS: GERMINATION 73 8o» 7w 0) O) 3 600 *- C Q) 0 0. i«o C 0 Q) £ 400 3'>« 200 sucrose concentration (%) Fig. 1. Torenia asiadca. Graph showing mean tube length for pollen grains in control sucrose medium. of the germinated pollen grains had a mean tube length of 755/(. in 20 ppm of boric acid. Since the present work was confined to study- ing fixed concentrations of different chemicals, and not determining which concentration gave maximum tube length, the concentra- tion of boric acid below 20 ppm was not tried. With an increase in the concentration of the chemical, percentages gradually declined — to a minimum of 70 percent germination and mean tube length to 403/1 — in 100 ppm of boric acid. 74 GREAT BASIN NATURALIST Vol. 34, No. 1 2. Calcium nitrate (Fig. 3a). A condition the reverse of that for boric acid (as described above) was obtained when pollen grains were treated with calcium nitrate solution. A high germination percentage and longer tubes were noticed in the maximum concen- tration of this chemical (100 ppm), as can be observed in Table 2. Mean tube length gradually increased from 418/j, in 20 ppm to 813/.(, in 100 ppm. Similarly, percentage germination progressed upwards from 66 percent in 20 ppm to 91 percent in 100 ppm. 3. Magnesium sulphate. A 40 ppm concentration of magnesium sulphate gave the highest germination percentage (88 percent) and the longest pollen tubes (mean length 556/x) of any of the concen- trations of this chemical used. Higher concentrations, like 100 ppm, were toxic, and they produced minimum germination percentages (69 percent) and shorter pollen tubes (323. S/i). 4. Potassium nitrate. Potassium nitrate produced many abnor- malities. The most favorable concentration of this chemical for pollen germination was 60 ppm. This concentration gave 74 percent germination with pollen tubes of 444/x mean length. At 20 ppm concentration of potassium nitrate, 4 percent of the pollen tubes were branched at their tips. However, rate of branching declined as the chemical concentration increased. Many of the branched pollen tubes had vesicles at the tip of one of their branches. Morphology of pollen tubes: Pollen tubes in different media were studied for an understanding of the morphological variations, Table 2. Percentage germination and mean tube length of Torenia asiatica pollen germinated in 15 percent sucrose solution mixed with different chemicals. Concentration Mean tube length of chemical Percentage in microns Chemical (ppm) germination after 2 hrs. Boric acid 20 89 755.0 (aBO,) 40 86 587.5 60 78 492.7 80 75 429.5 100 70 403.0 Calcium nitrate 20 66 418.0 (CA[N03]2.4H.O) 40 68 484.0 60 80 591.0 80 83 707.5 100 91 813.5 Magnesium sulphate 20 86 439.5 (MgS04 . 7 H2O) 40 88 556.0 60 75 499.5 80 75 379.5 100 69 323.5 Potassium nitrate 20 70 346.0 (KNO3) 40 72 354.5 60 74 444.0 SO 66 302.0 100 62 240.0 March 1974 nambudiri, thomas: germination 75 20 4'> 60 ppm concentration of chemicals ao Fig. 2. Graph to show comparative effects of different chemicals on the mean pollen tube length for Torenia asiatica. if any, shown by the growing tubes. Although pollen grains of T. asiatica consist of three germinal pores, only a single tube resulted. The protoplasm migrated into this tube. In the control sucrose medium and in the four chemicals, the pollen tube developed in either a coiled or a zig-zag manner (Fig. 4c). A combination of these growth characteristics also occurred in the same tube. Occa- sionally, tubes showed callose plugs (Fig. 4a). After its initiation into potassium nitrate, a pollen tube developed normally for a time and then branched. One of these branches had a vesicle at its tip. 76 GREAT BASIN NATURALIST Vol. 34, No. 1 Fig. 3. Torenia asiatica: (a) germination of pollen grains in calcium nitrate- sucrose medium (60 X) ; (b) germinating grains in boric acid-sucrose medium (60 X). March 1974 nambudiri, thomas: germination 11 Discussion Johri and Vasil (1961) noted that germinating media give varied resuhs when used with pollen grains of different species. This study revealed that sucrose is the best sugar for germination of the pollen grains of T . asiatica and that maximum germination occurs in a 15 percent concentration. This conclusion is in accord with that of Goss (1962) who attained maximum germination in 10-15 percent sucrose medium for Ornithogalum caudatum pollen and Nair and Deshpande (1968) for pollen of Luff a cylindrica. Sucrose medium was also used for germination of pollen grains of various plants by Vasil (1961), Biswas and Datta (1964), Cook and Walden (1965), Premnath and Purohit (1969), and others. Linskens (1964) suggested that sucrose was better than any other organic compound as a medium for pollen germination. However, Premnath and Purohit (1969) found no pollen germination in sucrose concen- trations below 40 percent for spinach beet pollen they studied. Biswas and Datta (1964) and Datta and Neogy (1965) observed maximum germination in 4 percent sucrose solution, while Singh (1957) reached a maximum in 10 percent sucrose solution, for Crotalaria pollen. However, as stated above, we determined that a 15 percent concentration of sucrose was the most suitable for pollen germination of T. asiatica. We consider this to be a specific differ- ence depending upon the osmotic pressure of the cell which again may be dependent upon the locality. The role of boron on germinating pollen was described by Stanley and Loewus (1964), Vasil (1964), and others as associated with sugar metabolism, as an inducer of oxygen absorption, and as a requirement for pectin synthesis. The effects of various com- pounds of boron on the germinating pollen have been reported by Stanley and Lichtenberg (1963). In the present experiments a lower concentration of boron (20 ppm) promoted a high percentage germination and longer pollen tubes. In higher concentrations of this chemical, germination was retarded. Higher germination per- centage and longer tubes with boron have been reported by Thomp- son and Batjar (1950), Young (1958), and Bamzai and Randhawa (1969). Young (1958) found that effective germination was possi- ble when pollen grains were allowed to germinate in boric acid concentrations of up to 50 ppm. However, experiments of Bamzai and Randhawa (1969) showed that higher percentage germination and tube length could be achieved with concentrations of up to 20 ppm. The results of the present investigation are in general agree- ment with those of Bamzai and Randhawa (1969) regarding the effect of boric acid. Therefore, we agree that the reduction in the germination percentages and a lower mean tube length in higher concentrations of boric acid may be caused by the toxic effect of this chemical in these higher concentrations. Cook and Walden (1965) reported that the presence of a calcium ion is required for pollen germination. In the present work germination increased with an increase in concentration of calcium 78 GREAT BASIN NATURALIST Vol. 34, No. 1 Fig. 4. Torenia asiatica: (a) branched pollen tube, showing callose plugs (150 X); (b) pollen grain of Torenia asiatica, showing tricolpate condition (250 X);' (c) germinating pollen tube, showing the zig-zag nature (150 X). March 1974 nambudiri, thomas: germination 79 nitrate (Table 2). Thus, in 100 ppm of calcium nitrate maximum germination was noted. The present experiments were intended to demonstrate only the effect of the known range of concentrations of different chemicals on pollen germination. Whether or not a higher concentration of calcium nitrate than 100 ppm would have ennanced pollen germination further was not within the scope of the jiresent study; nevertheless, the study does provide a basis for further research. Cook and Walden (1967) also noted that con- centrations of calcium higher than 100 ppm produced long pollen tubes and that germination does not occur in the absence of this chemical. Importance of calcium to the germinating pollen was given in detail by Faull (1955), Knawck and Brewbaker (1961), and Linskens (1964). Pfahler (1968) noted increased germination when calciiun was added to pollen of some hybrid maize. As in experiments of Cook and Walden (1967), so in the present experi- ment: a relatively higher concentration of calcium gave better re- sults. Most of the resulting pollen tubes did not rupture. This decreased incidence of rupture among the pollen tubes has been attributed to the capacity of calcium to give more rigidity to the cell wall (Brewbaker and Knawck, 1964). Compared with the control medium, potassium and magnesium do not improve germination percentage or mean tube length of pollen grains. However, potassium nitrate induced certain abnormalities, such as branched pollen tubes and vesicles. In the pollen tubes that grew in a coiled or zig-zag manner, the wall was not straight as in many other plants. Such coiled or zig- zag tubes have been obsened by Loo and Hwang (1944) and Vasil (1960). Loo and Hwang (1944) observed that when pollen grains of Antirrhinum ma jus, Then sinensis, Brassica juncea, Triticum vulgarc, and Hordeum vulgare were germinated in the presence of colchicine, the resultant tubes became either zig-zag or coiled. Loo and Hwang (1944) considered such an abnormality to be the direct effect of the treatment of colchicine, not of a change in the osmotic pressure, as had been suggested by earlier workers. Abnormal tubes were recorded by Brink (1924) and Vasil (1960) also. Vasil (1960) found in certain cucurbitas abnormal pollen tubes having branched or swollen tips. Vasil attributed these modifications to hypo- or hypemutrition, high humidity, high temperature and/or stale pollen. However, Nair and Deshpande (1968), working on Luffa cylindrica pollen, noted that apparently there was no signi- ficant correlation between germination results and meteorological data. Because the coiled tubes are found in both the control and the chemically induced pollen, there might be some other hitherto un- explored factor that regulates the morphology of the developing tube of the pollen of T. asiatica. Branched pollen tubes having a vesicle on one of their tips were also found by Loo and Hwang (1944), Vasil (1960), Nair et al. (1964), and Nair and Deshpande (1968). Loo and Hwang sug- gested that these swellings were the result of germination in a medium containing indole-3-acetic acid and that the abnormality 80 GREAT BASIN NATURALIST Vol. 34, No. 1 was the direct effect of this chemical. The branching of the tube and the subsequent swelling of one of these branches in the present experiments is observed only when the pollen grains are germinated in a potassium nitrate-sucrose medium. It is noteworthy that in lower concentrations of the above mixture there was a lower per- centage of branching than in higher concentrations. Whether the abnormalities are a direct result of the addition of KNO3 or an ex- pression of certain other physiological phenomena in the plant cannot be ascertained until more pollen grains belonging to dif- ferent taxa are studied responding to KNO3 in the germinating medium. Acknowledgment The authors are grateful to Dr. D. Kumar, Department of Botany, University of Dar-es-Salaam, Tanzania, for critically re- viewing the manuscript. References Bamzai, R. D., and G. S. Randhawa. 1969. Studies on germination and storage of Mango {Mangifera indica) pollen grains. Jour. Palynol. 5(2) : 104-1 10. Biswas, P. K., and R. M. Datta. 1964. Effects of auxins and antiauxins on the rate of pollen tube growth in Crotalaria juncea L., the Sun hemp. Revist. d. Biol. 4(3-4) :221-228. Brewbaker, J. L., AND B. H. Knawck. 1964. The calcium ion and substances influencing pollen growth. Pages 143-151 in H. F. Linskens, ed. Pollen Physiology and Fertilization. North Holland Publ. Co., Amsterdam. Brink, R. A. 1924. The physiology of pollen 2. Amer. Jour. Bot. 2:283-294. Cook, F. S., and D. B. Walden. 1965. The male gametophyte of Zea mays L. II. In vitro germination. Canad. Jour. Bot. 43:779-786. . 1967. The male gametophyte of Zea mays L. III. The influence of temperature and calcium on pollen germination and tube growth. Canad. Jour. Bot. 45:605-613. Datta, R. M., and A. K. Neogy. 1965. Comparative studies on the rates of growth of pollen tubes of some species of Crotalaria. Acta Biol. Hung. 16(1):35-41. Erdtman, G. 1952. Pollen morphologj^ and plant taxonomy, Angiosperms. Stockholm. Faull, a. F. 1955. Some factors in pollen gennination. Calcium salts, dextrose, drying. Jour. Arnold Arboretum 36:171-188. Goss, J. A. 1962. In vitro germination of Ornithogalum caudatum pollen. Trans. Kansas Acad. Sci. 65(3) : 310-31 7. Ikuse, M. 1956. Pollen grains of Japan, Tokyo. JoHRi, B. M., and I. K. Vasil. 1961. Physiology- of pollen. Bot. Rev. 27:325-381. Knawck, B. H., and J. L. Brewbaker. 1961. The essential role of calcium ion in pollen germination and population effect. Plant Physiol. (Suppl.) 36:16. Linskens, H. F. 1964. Pollen Physiology. Ann. Rev. Plant Physiol. 15:225-270. Loo, T. H., and T. C. Hwang. 1944. Growth stimulation by manganese sulfate, indole-3-acetic acid, and colchicine in pollen germination and pollen tube growth. Amer. Jour. Bot. 31:356-367. Nair, p. K. K., V. R. Balasubramanyam, and H. A. Khan. 1964. Palynological investigation of some guava varieties. Ind. Jour. Hort. 21:29-84. Nair, P. K. K.. and S. U. Deshpande. 1968. Pollen studies in successive flowers of Luffa cylindrica Linn.) M. J. Royen. Jour. Palynol. 5(2):98-106. Natarajan, a. T. 1957. Studies in the morphology of pollen grains. Tubiflorae. Phytom. 8:21-42. March 1974 nambudiri, thomas: germination 81 Pfahler, p. L. 1968. In vitro germinations and pollen tube growth of maize {Zea mays) pollen II. Pollen source, calcium and boron interactions. Canad. Jour. Bot. 46:235-240. Premnath, and S. p. Purohit. 1969. Studies on pollen morphology and physiology in spinach beet {Beta vulgaris var. banghalensis Hort.). Jour. Palynol. 5(2): 111-121. RiscH, C. 1939. Die pollenkomer der in Deutschland wild wachsend on Schrophulariaceen. Ber. Deut. Bot. Ges. 57:108-121. Singh, V. P. 1957. Studies in the artificial germination of pollen grains. I. Germinations of pollen grains of Sesbania aculeata Pers. and Crolalaria juncea L. Agra Univ. Jour. Res. (Sci.) 6(l):63-68. Stanley, R. G., and E. A. Lichtenberg. 1963. The effect of various boron compounds on in vitro germination of pollen. Physiol. Plantarum. 16:337-346. Stanley, R. G., and F. A. Loewus. 1964. Boron and myoinositol in pollen pectin biosj'nthesis. Pages 128-136 in H. F. Linskens, ed. Pollen physiology and fertilization. North Holland Publ. Co., Amsterdam. Thompson, A. H., and L. P. B.atj.^r. 1950. The effect of boron in the germina- tion medium on pollen germination and tube growth for several deciduous tree fruits. Proc. Amer. Soc. Hort. Sci. 56:227-230. Varghese, T. M. 1968. Studies in the family Scrophulariaceae. II. Pollen morphology. Jour. Palynol. 4(2):91-97. Vasil, I. K. 1964. Effect of boron on pollen germination and pollen tube growth. Pages 107-119 in H. F. Linskens, ed. Pollen physiology and fertilization. North Holland Publ. Co., Amsterdam. Young, T. W. 1958. A technique for testing mango pollen viability on artificial medium. Hort. Adv. 2:72-75. AN UNUSUAL SPIDER BITE Dorald M. Allredi Abstract. — The bite of a male blackwidow spider on a Utah woman is described. On 24 October 1973 a woman in Provo, Utah was bitten by a spider which I subsequently identified as a subadult blackwidow, Latrodectus hesperus Chamberlin and Ivie. As she arose in the morning she slipped her bare feet into a pair of solid-toe loafers, and walked into the kitchen. Her right foot immediately began to hurt, and as she removed her shoe she noted a crushed spider on her foot. Pain was localized in her big toe where she noticed a red spot and puncture marks on the top about one-half inch from the base. The pain soon extended to her thigh where it remained for about the first hour. During this time her leg ached and throbbed, occa- sionally with sharp pains. After an hour the pain in her thigh subsided, but remained prominent in her foot. The pain in her foot gradually subsided until the third day after the bite when it com- pletely disappeared. She did not consciously detect other symptoms of arachnidism (rigid abdomen, tightness in the chest, difficulty in breathing, nausea, and sweating, etc.), most likely because she was in her fifth month of pregnancy and was experiencing "morning sickness." The site of the bite remained red during the whole period of pain, although no scab or necrosis occurred. One month after the bite the site remained red, although no pain or tenderness was present. The bedroom of the victim was in a semi-basement apartment. It had two windows, one of which had been open for several days prior to the biting incident. The spider likely moved into the apart- ment through the open window with the advent of cold weather. According to Dr. WilHs J. Gertsch, bites by male black widows are unusual, and such incidences apparently have not been pre- viously recorded. The male likely has venom similar to that of the female, but is not inclined to bite. The entrapment of the spider in the shoe was undoubtedly the stimulus that resulted in the bite. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 82 f iVP^i •W'\ ^J?/ "i ■ ' >• V ■ • V t«- J ^^K^ GREAT BASIN NATURALIST V^H ^^^^ "Special Notic ^^^B Eoimiiein.iJiy wiui vuiLune 35 (Marcn ij/jj iiio vjieat iia jm pralist will be combined with the Brigham Youiig Univei mce Bulletin, Bi< n " ' " the title of The Great Basin mrahst. The foi^ ^ from the present 6x9 inch ■e column page to a 7x10 nicli double column page. The journal Icontinue to be published quarterly, but the number of pages per ioie will increase to approximately 500. Manuscripts up to 100 Bed pages (about 200 manuscript pages) in length wiU be ac- Ibble. Publication and reprint charges will be adjusted to the r format, but will not be significantly different from that indi- rI below. The subscription rate will" be increased to $9.00 per I (outside the United States, $10.00) . [ Notice to Contributors Dngniai manuscripts in EngHsh pertaining to the Biological |ral liistory of western North America and intended for publica- I in the Great Basin Naturalist should be directed to Brigham bg University, StorJi^-n T Wood, Editor, Great Basin Naturalist, io, Utah 84602. *' i wo copies are required, typewritten, double- Ci- ii one side of the paper, with margins of at least pnch on all sides. In general, the style should conform to recom- idations published in the most recent edition of the Council of Io ;ors Style Manual, published by the American Institute Bi^.^^...il Sciences. An abstract, about 3 percent as long as the Kand written in accordance with Biological Abstracts guidelines, Did precede the introductory'' paragraph of each article. Illustrations and Tables: All illustrations should be made I a vriew of having them appear within the limits of tlie printed t. The illustrations that form a part of an article should ac- ipany the manuscript. All illustrations and tables to appear in i'^i'mal are to be made under the supervision of the editor, and of the cuts and labor required to set tables is to be borne by tributor, iNTS: No reprints are furnished free of charj'^. A mi' o list uits and an order form are sent with thf RePMNTS SCHEDI HE GrE.V iNatUIL^LIST Each Additional 2pp. 4pp. 6pp. 8pp, lOpp, 12pp. 2pp. ^copies S12 00 514.00 ,?l(^nn ^i.qnn iODon «?7 nn itin,\ .00 30.00 TABLE OF CONTENTS Numeric analysis of the lizard genus Scelopor us with special reference to cranial osteology. Kenneth R. Larsen and Wilnier W. Tanner 1 The generic name of the North American musk turtles. Hobart M. Smith and Kenneth R. Larsen _ 42 Studies on the tolerance of aquatic insects to low oxygen concentrations. Arden R. Gaufin, Robert Clubb, and Robert Newell 45 Substrate color matching in the grasshopper, Circotettix rabula (Orthoplera: Acrididae). George W. Cox and Darla G. Cox 60 Effects of chemicals on the germination of pollen grains of Torenia asiatica Linn. E, M. V, Nambudiri and M. K. Thomas 7^ An unusual spider bite. D. M. AUred 82 c.,„, -. .1 ,,„v;„^ 4^ ^,,1,^, ,;i,„,.o inside h?s^' '■"■"■-^r Voiiune 6-1', xNo. 2 June 30, 1974 The Great # J]OTiJi]IJ^T PuxiJLioLLED BY Brtoham Young University GREAT BASIN NATURALIST Editor: Stephen L. Wood, Department of Zoology, Brigham Yoi University, Provo, Utah Editorial Board: Stanley L, Welsh, Botany, Chairman; Wihner Tanner, Zoology; Joseph R. Murdock, Botany; Vernon Tipton, Zoology; Ferron L. Andersen, Zoology Ex officio Editorial Board Members: A. Lester AUen, Dean, College of Biological and Agric tural Sciences Ernest L. Olson, Director, University Press, Univers Editor The Great Basin Naturalist was founded in 1939 by Vasco Tamier. It has been continuously published from one to four a year since then by Brigham IJoung University, Provo, Utah, general, oid^ original, previously unpublished manuscripts pei ing to the biological natural history of the Great Basin and west North America will be accepted. Manuscripts are subject to the proval of the editor. Subscription: The annued subscription is $5.00 (outside United States $5.50). The price for single numbers is $2.00 eac. All matters pertaining to me pin*chase of subscriptions and ba< numbers should be directed to Brigham Young University Pre }^ubHcation Sales, 205 UPB, Provo, Utah 84601. Libraries or oth^ organizations interested in obtaining this journal through a conti^ imig exchange of scholarly publications should contact the Brigh£ "Xomig University Exchange Librarian, The Library, Provo, Ut 84601. Manuscripts: All manuscripts and other copy for the Gre Basin Naturalist should be addressed to the editor. Contribut should consult the instructions printed on tlie back cover of a recei number. The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University Volume 34 June 30, 1974 No. 2 THE SPECIES OF CALINEURIA AND DORONEURIA (PLECOPTERA: PERLIDAE)^ Bill P. Stark- and Arclen R. Gaufin- Abstract. — The genus Doroneuria is reviewed, and characters necessary for recognition of its two included species are figured. Doroneuria baumanni, sp. nov., is described for the male, female, and nymph, and a male holotype and female allotype are designated. Calineuria is removed from the synonymy of Doroneuria, and the genus is characterized in detail. Distributional data are pre- sented for species in both genera. Doroneuria was proposed as a subgenus of Acroneuria by Need- ham and Claassen (1922) with the newly described theodora as the type species. This subgenus was omitted from the systematic list in the Needham and Claassen monograph (1925). Kicker (1954), apparently unaware of the former paper, proposed a new subgenus, Calineuria, for the species Acroneuria calif ornica (Banks) and A. theodora Needham and Claassen with calif ornica (Banks) as type species. lilies (1966) gave Doroneuria generic status and considered Calineuria a synonym. A detailed study of these species has indicated clearly that two genera are involved and has led to the discovery of a new species of Doroneuria. Calineuria Ricker Acroneuria {Calineuria) Ricker (1954:39). Type species Perla calif ornica Banks. Doroneuria: lilies (1966:330). In part. Doroneuria: Zwick (1973:274). In part. The genus Calineuria may be characterized in the adult male by ( 1 ) a longitudinally rectangular hammer on abdominal sternum 9, (2) a median, heavily sclerotized tergite on tergum 10 (Fig. 1), and (3) the aedeagus (Figs. 1, 2) bearing on the basal portion 4 prominent patches of long golden-brown spinules (2 lateral, 1 ventral, and 1 apical). Adult females are characterized by (1) the unproduced, shallowly notched posterior margin of sternum 8 with an unsclerotized, U-shaped border around the notch (Fig. 3) and (2) a membranous vagina that extends forward to near the anterior margin of sternum 7 and is not lined internally by spinulae. 'Study supported by EPA Contract, Univ. of Munt.ina #881 1 =Biology Dept., Univ. of Utah, Salt I.akc City, Ulnli 84112 83 84 GREAT BASIN NATURALIST Vol. 34, No. 2 Mature nymphs are characterized by ( 1 ) an incomplete, irregularly spaced row of spines across the occiput, (2) numerous prominent spines on each abdominal tergum in addition to the posterior fringe, (3) a dense fringe of long silky hairs along the mesal surface of the cerci originating in the whorl of spines on each cereal segment (Fig. 14), and (4) usually a complete posterior fringe of spines across Figs. 1-3. Calineuria calif or nica: 1, Male terminalia with aedeagus extruded, dorsal; 2, Aedeagus, lateral; 3, Female abdominal sternum 8. (ap = apical patch, Ip^lateral patch, vp^ventral patch). June 1974 STARK, GAUFIN: PLECOPTERA 85 Fig. 4. Distribution map of C. californica. D. baumanrii, and D. theodora. abdominal sternum 7 (Fig. 14). This monotypic genus is known only from western North America. Calineuria californica (Banks) Perla californica Banks (1905:87). Holotype 9; Claremont, California. (MCZ #11318). 86 GREAT BASIN NATURALIST Vol. 34, No. 2 Perla concolor Banks (1908:255). Syn. Needham and Claassen, 1922. Acroneuria calif or nica: Needliam and Claassen (1925:192). Acroneuria (Calineuria) calijornica: Ricker (1954:39). Doroneuria calif ornica: lilies (1966:330). Male. — Macropterous. Length of forewings 19-21 mm; length of body 18-20 mm. General color yellow brown. Wings hyaline, veins brown. Paraprocts sclerotized, fingerlike, with a small termi- nal acute spine. Aedeagus (Figs. 1,2) as described above. Occasion- al specimens with ventral patch of aedeagal spinules variously reduced in size. Female. — Macropterous. Length of forewings 24-27 mm; length of body 22-23 mm. Subgenital plate (Fig. 3) and vagina as described above. Nymph. — Length of mature male 17-18 mm; mature female 22-23 mm. General color yellow and brown. Head with distinctive large yellow spot covering ocellar area. Abdominal sterna and cerci (Fig. 14) as described above. Calineuria calijornica is a species of the Pacific coast and north- ern Rocky Mountains (Fig. 4). Previous studies (Gaufin et al., 1966; Baumann, 1973; Stark et al., 1973) have recorded the species from Colorado and Utah, but an examination of all this material in the University of Utah collection previously determined as cali- jornica has shown these records are of a presently undetermined species of Acroneuria (possibly depressa Needham and Claassen). Specimens examined in this study: CALIFORNIA: Alameda Co., Oakland, 15-V-lO, E. C. Van Dyke, 1$ (cas); Oakland, 9-V-09, E. C. Van Dyke, \$ (cAs). Alpine Co., Markleeville, 7-VII-49, Allen, 1 9 (cas). Butte Co., Big Chico Creek, 18-XI-71, M. Kainu, nymphs (uu). Eldorado Co., 3 mi S Camino, 26-VI-48, MacNeil, \$ (cas). Humboldt Co., Prairie Creek, 9-VI-36, E. C. Van Dyke, 1^19 (cas); North Fk. Yager Creek, 8-VIII-66, R. E. Leech, 19 (cas). Lake Co., Anderson Springs, 20-III-55, Leuschner, 1^ (inhs); 6 mi S Middletown, 12-V-26, 4 5 5 9 (cas). Los Angeles Co., Arroyo Seco Canyon, 23-VI-18, E. C. Van Dyke, 19 (cas); East Fk. San Gabriel River, 16-Vi-59, nymphs (uu). Marin Co., Muir Woods, 25-IV-64, D. G. Denning, 1^ (rom); Cascade Creek, 13-V-52, R. E. Leech, \$ (cas); Bear Valley, 25-V-19, \$ (cAs); Ross, 25-IV-18, Van Duzee, 19 (cas); Lagunitas, 14-IV-08, Van Duzee, 1 9 (cas); Lagunitas, 30-V-38, Nast, 4 9 (cas). Mendocino Co., Camptche, 7-IV- 36, E. C. Van Dyke, \$ (cas); Yorkville, 30-IV-24, Van Duzee, \$ (cas). Mono Co., Convict Creek, VI-63, 1 $ (uu); Sonora Bridge, 28-VII 62, Rentz and MacNeil, \$ (cas). Placer Co., Truckee River, l-VIII-68, R. W. Baumann, 6 5 29 (usnm). Plumas Co., Mohawk, 12-VII-57, Rentz, \$ (cas); Quincy, 26-VI-73, D. R. Givens and S. D. Smith, 1 $ (cwsc). San Bernardino Co., Lytle Creek, 7-VI-28, E. C. Van Dyke, 7$ 4 9 (cas). Santa Clara Co., San Jose, 13-X-81, Van Duzee, 1 9 (cas); Alum Rock Park, 29-VI-55, Ribble, 1 $ (kan); 29-VI-69. Cobb, 1^ (usnm). Shasta Co., 26-VI-21, J. A. Kuscho. 19 (cas); Burney Falls, 29-VI-47, H. P. Chandler, 1 9 (cas); Castle Crags St. Pk., 20-VI-64, D. G. Denning, 1 $ (rom). Siskiyou Co., Sacramento River. 8-VII-53, H. P. Chandler, 3 5 (cas). Sonoma Co., Guernevillo. 31-V-36. E. C. Van Dyke, \$ (cas); 19-VI-10, J. Kuscho, 1^19 (cas). Trinity Co.. Trinity Center, "6-VI-64, J. Pinto. 2 5 49 (ucr); Coffee Creek, 7-VI-34. E. C. Van Dyke, 15 19 (cas); Butter Creek, 21-VII-68, R. E. Leech, 19 (cas). Tuolume Co., Dana Fk. Tuo- lume River, 15-VII-36, Rayner 15 (inhs). IDAHO: Boise Co., Boise River, 12-VI-31, Ivie, nymphs (uu). Boundary Co., Moyie River, 26-III-65, A. V. Nebe- ker, nymphs (uu). Kootenai Co., Hayden Creek, 7-VI-69, R. W. Baumann, 2 5 June 1974 stark, gaufin: plecoptera 87 2 9 (usnm). Lemhi Co., Salmon River, 28-VI-73, M. and E. Gather, 2$ 29 (uu); Salmon River, 6-VII-64, A. R. Gaufin, 1 $ (uu); Salmon River, ll-VII-67, G. F. Edmunds, 1^ 49 (usnm). Shoshone Co., Coeur d'Alene River, ll-VI-69, A. R. Gaufin, 2^29 (usnm). MONTANA: Camas Co., Glacier Park, 7-VII-67, A. R. Gaufin. \S (uu). Flathead Co., Swan River, 29-VI-65, A. V. Nebeker and R. K. Allen, 415 13 9 (uu) (usnm); Swan River, l-VII-67, P. Milam, 6$ 49 (uu); McGregor Creek, l-VII-67, A. R. Gaufin, \$ 39 (uu); Camas Creek. 7-VII-67, A. R. Gaufin, 1^ 19 (uu). Lake Co., Swan River, 23-VI-67, P. Milam, l\ $ 69 (uu); Swan River, lO-VI-67, A. R. Gaufin, 9$ 29 (uu). Lincoln Co., Yaak River, 21-VI-69, R. W. Baumann, 7$ (uu). Missoula Co., Blackfoot River, 28-VI-67, A. R. Gaufin, 10 5 7 9 (uu). Pondera Co., Dupuyer, 24-VI-65, A. V. Nebeker, 1 $ (uu). Ravalli Co., Bitterroot River, 28-VII-65, J. R. Grierson, 2$ (uu). OREGON: Benton Co., Corvallis, 22-V-34, R. E. Dimick, \S (cAs); Corvallis, 8-V-34, Bali, \$ (inks); Alsea, 27-V-34, J. Schuh, 2$ 7 9 (c.\s) (iNHs). Clatsop Co., Cannon Beach, 15-VI-27, E. C. Van Dyke, 1 $ (cas); Big Creek, 28-V-49, S. G. Jewett, Jr., 5^39 (cas); Nehalem River, ll-VII-64, S. G. Jewett, Jr., 5^19 (sgj). Columbia Co., Clatskanie, 19-V-36, J. Schuh, 1 $ (c.\s); Scappoose, 20-VI-36, Rieder, 3$ 3 9 (inhs). Jackson Co., Rogue River, 17-VI-49, S. G. Jewett, Jr., \$ (cas). Lake Co., Paisley, 28-VI-51, B. Malkin, 2$ 19 (cas) (fmnh). Lane Co., Bob Creek, 13-VIII-32, R. E. Dimick, 1$ (cas). Lincoln Co., Siletz River, 24-V-34, R. E. Dimick, 3$ (cas). Linn Co., Santiam, 26-VII-60, D. G. Denning, 19 (usnm). Marion Co., Breitenbush, ll-VII-68, E. Evans, 1$ (usnm). Multnomah Co., Eagle Creek, 19-V-40, S. G. Jewett. Jr.. \$ (cas). Washington Co., Gales Creek, 14-V-47, S. G. Jewett, Jr., 1 $ (cas). WASHINGTON: Clallam Co., Crescent Lake, 21-VI-40, S. G. Jewett, Jr., 1 $ (cas); Forks. 2-VII-20, Van Duzee, 1 $ (cas); Bogachiel River, 18-VI-67, R. W. Baumann, 2$ 4 9 (usnm); Soleduck River, 23-VII-60, D. G. Denning, 1 9 (usnm). Grays Harbor Co., East Fk. Humptulips River, 2-IX-58, nymphs (uu); Donkey Creek, 18-VI-67. R. W. Baumann, nymphs (usnm). Jefferson Co., Hoh River, 3-IX-58, nymphs (uu). King Co., Rex River, 20-VII-72, C. Baker, 1 $ (uu); Re.x River, 5-X-73, A. R. Gaufin, nymphs (uu). Kittitas Co., Liberty, 27-VII-49, White, 19 (kan). Spokane Co., Little Spokane River, 13-VI-55, nymphs (uu). Thurston Co., Deschutes River, 15-VL67, R. W. Baumann, 10 5 5 9 (usnm). Yakima Co., Toppenish, 9-Vn-35, Beamer, 2 9 (kan); Naches River. 5-IX-58. nymphs (uu). Doroneuria Needham and Claassen Acroneuria {Doroneuria) Needham and Claassen (1922:249). Type species Acroneuria theodora Needham and Claassen by original designation. Acroneuria (Calineuria) Ricker (1954:39). In part. Doroneuria: lilies (1966:330). In part. Doroneuria: Zwick (1973:274). In part. The genus Doroneuria may be characterized in the adult male by (1) a longitudinally rectangular hammer on abdominal sternum 9 (Fig. 11), (2) tergimi 10 without a median tergite, and (3) the aedeagus bearing a spatulate sclerotized plate dorsally and two elongate, narrow, sclerotized lateral bars (Figs. 5, 6). Adult females are characterized by (1) the almost straight (sometimes slightly produced mesally) posterior margin of sternum 8 (Fig. 7) and (2) a membranous vagina that extends forward to the posterior margin of sternum 7 and is lined with numerous small golden brown spinu- lae (Figs. 7, 8). Mature nymphs are characterized by (1) an in- complete, irregularly spaced row of spines across the occiput (Fig. 12), (2) numerous prominent spines on each abdominal tergum in addition to the posterior fringe, (3) a dense fringe of long silky hairs along the me.sal surface of the cerci (Fig. 13), and (4) an 88 GREAT BASIN NATURALIST Vol. 34, No. 2 ^-v (."7!-^ Figs. 5-8. Doroneuria baunianni: 5, Male teniiinalia wiUi acdeagus extrud- ed, dorsal; 6, Aedeagiis, lateral; 7, Female abdominal sterna 7 and 8, vagina shown in outline; 8, Vagina. (pa = paraproct, dp = dorsal plate, lb = lateral bar). incomplete posterior fringe of spines on abdominal sternum 7 (Fig. 13). The genus is presently known only from western North America. Doroneuria baumanni, sp. nov. Acroneuria theodora-. Frison (1942:284-285). In part. Acroneuria {Calineuria) theodora: Jewett (1959:89). In part. Doroneuria theodora: lilies (1966:331). In part. June 1974 STARK, GAUFIN: PLECOPTERA 89 Male. — Brachypterous to macropterous. Length of forewings 18-27 mm; length of body 22-25 mm. General color dark brown. Wings hyaline, veins black. Paraprocts similar to D. theodora, sclerotized and fingerlike. Aedeagus (Figs. 5, 6) with 12-22 promi- nent spines on the lateral sclerotized bars; ventral and lateral sur- faces of aedeagus with numerous minute spinulae. Female. — Macropterous. Length of forewings 31-33 mm; length of body 27-30 mm. Subgenital plate (Fig. 7) unproduced. Vagina (Figs. 7, 8) lined with minute golden brown spinulae. Nymph. — Length of mature male 20-22 mm; mature female 31-34 mm. Similar to D. theodora in coloration and general mor- phology. Occiput with a transverse row of long silky hairs between the postfrontal suture and the row of spines. Frons with a single long seta near each antennal base. Dorsum of thorax and abdomen with a median longitudinal row of long silky hairs. Doroneuria baumanni is a species of the Cascade and Sierra Nevada Mountains of western North America (Fig. 4) with relict populations in Nevada and southeastern Oregon. The lack of records for Doroneuria in Ricker's (1943) study of southwestern British Columbia indicates that the species survived the Pleistocene glacia- 10 Figs. 9-11. Doroneuria theodora: 9, Male terminalia with aedeagus extrud- ed, dorsal; 10, Aedeagus, lateral; 11, Male abdominal sterna 9 and 10. (h = hammer, pa = paraproct, dp = dorsal plate, lb = lateral bar) 90 GREAT BASIN NATURALIST Vol. 34, No. 2 tions in a coastal refuge and has been unable to recolonize severely glaciated areas. Types. — Male holotype, female allotype, and 5 male and 1 female para types from Burney Falls State Park, Shasta Co., Cali- fornia, 8-X-72, D. G. Denning. Holotype male No. 73022 and allo- type female deposited at the United States National Museum, Wash- ington, D.C. Additional paratypes as follows: CALIFORNIA: Sis- kiyou Co., Tate Creek, 8-IX-54, H. P. Chandler, 2d (sgj) (cas); Trinity Co., Grizzly Meadows, 10-15-IX-64, Buxton and Gurney, 3cf 1 ? (usnm); 10 mi W Thompson Peak, 16-IX-64, Buxton, 2 9 (usnm). OREGON: Benton Co., Marys Peak, 3-VI-68, T. Yama- moto and S. D. Smith, 1 ? (rom). Curry Co., Fork Creek, 19-VII-67, E. Evans, 1 d (usnm). Harney Co., Fish Creek, Steens Mts., 20-VII- 69, S. G. Jewett, Jr., 2cf (usnm). Lane Co., McKenzie Bridge, 21-IX-34, R. E. Dimick, 3d" (inks) (uu); McKenzie River, 21-IX- 34, R. E. Dimick, 2cf (osu). Lincoln Co., Tidewater, 6-VI-68, T. Yamamoto and S. D. Smith, 1 cf (rom). Linn Co., Marion Forks, 28-VIII-68, Goeden and Gurney, 1 ? (usnm). Tillamook Co., Nes- kowin, ll-VII-63, G. Wiggins, Id" (rom). Yamhill Co., McMinn- viUe, 4-V-47, K. M. Fender, Id (cas). WASHINGTON: Anderson Creek, Stevens Pass, 29-IX-59, 1 d (inks). Chelan Co., Icicle Creek 9-X-71, L. Whyte, Id (cwsc). Clark Co., Lewis River, 15-IX-57, H. Hacker, Id" (usnm). Additional specimens examined: CALIFORNIA: Butte Co.. Big Chico Creek, 26X11-72, M. Kainu, nymphs (uu). Fresno Co., South Fk. King River, 6-VII-lO, E. C. Van Dyke, 1^ (cas). Modoc Co., 20-VII-22, Lindsey, \$ (inks). Mono Co., Mono Lake, 31-VII-40, Lipovsky, \$ (kan). Tuolume Co., Kennedy Creek, 18-VII-73, D. R. Givens and S. D. Smith, nymphs (cwsc). NEVADA: Elko Co., near Midas, 20-IX-57, G. F. Edmunds and R. K. Allen, nymphs (uu); Franklin River, 19-IX-57, G. F. Edmunds and R. K. Allen, nymphs (uu). OREGON: Benton Co., Philomath, 4-VII-63, G. Wiggins, 19 (rom); Corvallis, 27-VII-62, G. C. Eickwort, 1$ (msu). Hood River Co., Mt. Hood, 24-VI-25, E. C. Van Dyke, \S (cas). Lane Co., McKenzie Bridge, 21-IX-34. R. E. Dimick, \S \9 (iNHs) (cAs); South Fk. McKenzie River, 4-IX-32, R. E. Dimick, \$ (inks); McKenzie River, 21-IX-34, R. E. Dimick, 2$ 2 9 (inks). Multnomah Co., Wahkeenah Falls, 15-VII-63, G. Wiggins, 1 9 2 nymphs (rom). WASHINGTON: King Co., Rex River 5-X-73, A. R. Gaufin, nymphs (uu). Lewis Co., Nickle Creek, l-IX-58, nymphs (uu). Pierce Co., Slippery Creek, 5-IX-58, nymphs (uu). Yakima Co., Chapparal Creek, 2-IX-54, B. Malkin, 1$ (cas). This species is named for Dr. Richard W. Baumann of the Smithsonian Institution who has assisted in obtaining material and by making suggestions for this and other studies in progress. Doroneuria theodora (Needham and Claassen) Acroneuria (Doroneuria) theodora Needham and Claassen (1922:254). Holotype 9, allotype $ ; Yellowstone Nat'l Pk., Wyoming. (Cornell Univ. #1180). Acroneuria theodora-. Frison (1942:284-285). In part. Acroneuria (Calineuria) theodora: Ricker (1954:39). Doroneuria theodora: lilies (1966:331). In part. Male. — Brachypterous to macropterous. Length of forewings 7-17 mm; length of body 19-22 mm. Similar to D. baumanni in June 1974 STARK, GAUFIN: PLECOPTERA 91 jiiiliiljaiJiu .jmulmUmiMM llJJlJjn^ 13 14 Figs. 12-14. Calineuria and Doroneuria nymphal structures: 12, Doronruria, dorsum of head; 13, Doroneuria abdominal sternum 7 (mature $ nymph) and cereal segments basal and apical; 14, Calineuria abdominal sternum 7 (mature $ nymph) and cereal segments basal and ajjical. coloration and general features. Aedeagus (Figs. 9, 10) without spines on the lateral sclerotized bars; ventral and lateral surfaces of aedeagus with numerous minute spinulae. Female. — Brachypterous to macropterous. Length of forewings 15-25 mm; length of body 29-31 mm. Subgenital plate and vagina similar to D. haurnanni. 92 GREAT BASIN NATURALIST Vol. 34, No. 2 Nymph. — Length of mature male 20-22 mm; mature female 31-34 mm. Presently indistinguishable from D. baumanni. Doroneuria theodora is a species of the northern Rocky Moun- tains (Fig. 4). Records are presently from north of the Wind River range in Wyoming into southern Canada and west through Idaho to eastern Oregon. Published records of theodora from Utah and Colo- rado are based on the same species of Acroneuria nymph discussed under Calineuria calijornica above and should be disregarded. Specimens examined in addition to the holotype and allotype: ALBERTA: Banff Nat'l Pk., above Bow Falls, 19-VII-69, C. M. Yarmoloy, \$ (usnm); Spray River, Banff Nat'l Pk., 15-VI-69, C. M. Yai-moloy, nyinphs (uu); Crows Nest River, Crows Nest, 13-IX-75, R. Beck, nymphs (uu). IDAHO: Adams Co., Little Goose Creek, 21-VL64, A. R. Gaufin, nymphs (uu). Bonneville Co., West Fk. Pine Creek, 21-VI-64, J. W. Richardson and S. L. Jensen, nymphs (uu). Boundary Co., Canyon Creek, 12-VIII-63, M. Brusven, 1 $ 4 nymphs (usnm). Idaho Co., Lochsa River, 19-VI-64, A. R. Gaufin, nymphs (uu). Kootenai Co., Beauty Bay Creek, 4-VII-64, A. R. Gaufin, nvmphs (uu). Lemhi Co., Pine Creek, 28-VL64, J. K. Adams, nymphs (uu). MONTANA: Gallatin Co., Hyalite Creek, 17-VIII-52, R. A. Hays, \$ (uu); Hvalite Creek. 16-Vin-52, R. A. Hays, 45 1? (usnm). Granite Co.. Ranch Creek, 23-VII-66, M. L. Miner, \$ (usnm). Ravalli Co., Burnt Fk. Bitterroot River, 16-Vin-65, J. R. Grierson, 2$ 19 (usnm). OREGON: Wallowa Co., French Camp, Lostine Valley, 18- VnL52, G. F. Edmunds. 19 1 nymph (uu). WYOMING: Park Co., Beartooth Creek, 9-VIII-52, R. A. Hays, 1 $ (uu); Beartooth Creek, 31-VIII-51, R. A. Hays, 19 (uu); Beartooth Creek, 31-VI1I-51, R. A. Hays, 5^ (uu); Tower Creek, Yellowstone Nat'l Pk., 17-VIII-59, W. L. Peters and G. F. Edmunds, nymphs (uu). Sheridan Co., Tongue River, 20-VII-40, T. H. Frison, 1 $ (inhs); Tongue River, 28-VII-40, T. H. Frison, \$ (inhs); Big Horn Mts., 30-VII-40, T. H. Frison and T. H. Frison, Jr., 1^ (inhs); South Fk. Camp, Big Horn Nat'l Forest, 4-VIII-52, B. Malkin, \$ (cas). Acknowledgments The authors are grateful to the following individuals and mu- seiuns for the loan or gift of material used in this study: Dr. N. H. Anderson and Ms. J. R. Sawbridge, Oregon State University (OSU), Dr. R. W. Baumann, Smithsonian Institution (USNM), Dr. G. Byers, Kansas University (KAN), Dr. R. L. Fischer, Michigan State University (MSU), Mr. S. Frommer, University of California Riverside (UCR), Mr. S. G. Jewett, Jr., West Linn, Oregon (SGJ), Mr. M. Kainu, Chico State University, Dr. L. L. Pechuman, Cornell University (CU), Dr. M. Prokop, Field Museum of Natural History (FMNH), Dr. R. X. Schick, California Academy of Sciences (CAS), Dr. S. D. Smith, Central Washington State College (CWSC), Mr. D. Webb, Illinois Natural History Survey (INHS), and Dr. G. Wig- gins, Royal Ontario Museum (ROM). We also thank Dr. Baumann and Dr. G. F. Edmunds (University of Utah) for critically reading this manuscript. Other specimens used in this study are deposited in the collection at the University of Utah (UU) . Literature Cited Banks, N. 1905. Plecoptora. In Neuropteroid insects. Invert. Pacif. 1:85-91. . 1908. Neuropteroid insects, notes and descriptions. Trans. Amer. Entomol. Soc. 34:255-267. June 1974 stark, gaufin: plecoptf.ra 93 Baumann, R. W. 1971 Studies on Utah stoneflies (Plecoptera). Great Basin Nat. 33:91-108. Prison, T. H. 1942. Studies of North American Plecoptera with special refer- ence to the fauna of Illinois. Bull. 111. Nat. Hist. Sui-v. 22:235-355. Gaufin, A. R., A. V. Nebeker, and J. Sessions. 1966. The stoneflies (Ple- coptera) of Utah. Univ. Utah Biol. Ser. 14. 93 pp. Illies, J. 1966. Katalog der rezenten Plecoptera. Das Tierreich, 82. Walter de Gruyter and Co., Berlin. 632 pp. Jewett, S. G. 1959. The stoneflies (Plecoptera) of the Pacific Nortliwest. Oregon State Monographs 3. 95 pp. Needham, J. G., AND P. W. Claassen. 1925. A monograph of the Plecoptera or stoneflies of America north of Mexico. Thomas Say Found. Entomol. Soc. Amer. 2. 397 pp. Richer, W. E. 1943. Stoneflies of southwestern British Columbia. Ind. Univ. Pub. Sci. Ser. 12. 145 pp. . 1954. Nomenclatural notes on Plecoptera. Proc. Entomol. Soc. Brit. Columbia 51:37-39. Stark, B. P., B. R. Oblad, and A. R. Gaufin. 1973. An annotated list of the stoneflies (Plecoptera) of Colorado. Entomol. News. In press. ZwicK, P. 1973. Insecta: Plecoptera. Phylogenetisches system und katalog. Das Tierreich, 94. Walter de Gruyter and Co., Berlin. 465 pp. THE NAME OF THE BAJA CALIFORNIA CAPE WORMSNAKE Hobart M. Smith^ and Kenneth R. Larsen^ Abstract. — The type-specimen of Glauconia boettgeri Werner, 1899, is consubspecific with L. h. slevini Klauber, 1931. The valid name of the Baja California Cape wormsnake accordingly is Leptatyphlops humilis boettgeri (Werner) . The allocation of a snake described 75 years ago (Werner, 1899:- 116) as Glauconia boettgeri has long been uncertain, largely because of its unknown type locality. The original description is reasonably good, and the species was stated to be related to Leptotyphlops humi- lis; but no subsequent reviewer has placed it definitively with any known species. Werner (1917:198) later reviewed the whole family but added nothing except the speculation that boettgeri might be grouped with certain African species. He had earlier placed it as a synonym of the African Leptotyphlops labialis, but that species dif- fers in numerous ways (e.g., no preocular supralabial, rostral ex- tending posterior to eye level), as he noted in 1917. He concluded that L. boettgeri might be related to L. latifrons and L. scutifrons, both African species, but is distinct from them. Indeed it is distinct, since L. scutifrons has no preocular labial and L. latifrons has a very large rostral. His final thoughts, seemingly, placed L. boettgeri with African species rather than with L. humilis, his first impression; and perhaps for this reason Klauber (1940) made no attempt to allocate L. boettgeri, although he cited Werner's 1917 monograph. In an attempt to fix the allocation of Werner's name. Dr. Josef Eiselt of the Vienna Museum very kindly loaned us the holotype of Werner's species for more careful examination. Although too faded to reveal the pattern of pigmentation, in other respects the specimen, now No. 15455 in the herpetological collection of the Natural History Museum of Vienna, is a typical representative of the population now known as Leptotyphlops humilis slevini Klauber (1931:338). It has 254 dorsals; 17 subcaudals; 12 scale rows around tail, 14 around body; and median scales on head all about equally wide and little, if any, narrower than the scales in the median row on the neck and trunk. The body length is 203 mm, the tail 10.8 mm. The body- length/diameter ratio is 58, the body-length/tail-length ratio 18.4. For L. humilis slevini, Klauber (1940) records 253 as the mean (range 244-269) for the dorsals; 15 (range 12-18) for the subcau- dals; scalerow counts and median head scales as in L. boettgeri; body- length/diameter ratio moan 49; body-length/tail-length ratio mean These characters clearly place Werner's G. boettgeri with L. humilis slevini. The latter is the only L. humilis subspecies with 'Department of Environmental, Population, and Organismic Biology, University of Colorado, Boulder, 80302. 94 June 1974 smith, larsen: woumsnake 95 equal-sized median head scales, and only one other (L. humilis dugcsi) has as few as 254 dorsals (and its maximum is 257). Only one other (L. humilis cahuilae) has only five pigmented dorsal scalerovvs, but unfortunately this character cannot be determined; however, the general tone ("light brown') is matched. Direct com- parisons of the holotype with all available specimens of L. humilis (unfortunately none of L. humilis slevini) reveal a complete agree- ment in all external features of scutellation, except for the median head scales being equally broad and as large as the median scales of the trunk. This feature Klauber emphasized, however, as distinc- tive of L. humilis slevini. There is a minor deviation of the holotype from the mean body proportions of L. humilis slevini, but the range of variation in these features is considerable. Klauber did not record the variation for L. humilis slevini but noted (1940:99) that in a homogeneous series of 52 L. humilis humilis the range of body- length/diameter ratios varied from "under 45" (1) to "over 70" (1) and that in 54 of the same subspecies from the same area the body- length tail-length ratios varied from "under 16" (1) to "over 24" (3). Thus the ratios of the holotype of G. boettgeri respectively of 58 and 18.4 presumably fall well within the expected range for L. humilis slevini about the means respectively of 49 and 23. Indeed, the slightly shrunken holotype of G. boettgeri was originally mea- sured at 214 mm in body length, tail 11 mm, diameter 3 mm. The latter measurement apparently was an error, for even now the di- ameter differs in various parts of the body, between extremes of 3 and 4 mm, \\ith 3.5 mm being an approximate mean. In a less de- hydrated condition 4 mm would be likely. The length of the body has clearly diminished in the interim. The body-length/diameter ratio of 75 given by Werner (1899:116) is, however, clearly too high; the most reasonable figure (based upon Werner's 214 mm body length and our 4 mm estimate for diameter in the fresh speci- men) is 54, quite in line with that of L. humilis slevini. The name Leptotyphlops boettgeri has not been used frequently, and therefore the possibilit}^ arises of appealing to the International Commission on Zoological Nomenclature for conservation of L. humi- lis slevini. 33 years a junior of G. boettgeri. However, the name L. humilis slevini itself has not been in use for 50 years; nor is it a widely cited name. The case does not justify an appeal, even under the teVms of the 1972 decisions of the ICZN (Corliss, 1972:1120). Accordingly, it is necessary to accept as valid the name Lepto- typhlops humilis boettgeri (Werner, 1899) in replacement of L. humilis slevini Klauber (Klauber, 1931) as a subjective senior syno- n}Tti of the latter name. The Cape region of Baja California was visited by boat so fre- f|uontly before 1900 that it is not at all strange that a specimen of Leptotypfilops from there should have reached a European museum before 1899. The most frequently visited port was La Paz, whence records for L. humilis boettgeri are already available. We accord- ingly here propose that the type locality of G. boettgeri be restricted to I>a Paz, Baja California, Mexico. 96 GREAT BASIN NATURALIST Vol. 34, No. 2 Acknowledgments We are much indebted to Dr. Josef Eiselt for the loan of the holotype of G. boettgeri, and to Dr. T. Paul Maslin for use of com- parative material in the University of Colorado Museum. Literature Cited Corliss, J. O. 1972. Priority and stability in zoological nomenclature: Resolu- tion of the problem of article 23b at the Monaco Congress. Science 178:1120. Klauber, L. M. 1931. Notes on the worm snakes of the southwest, with de- scriptions of two new subspecies. Trans. San Diego Soc. Nat. Hist. 6(23) :333- 352. . 1940. The worm snakes of the genus Leptotyphlops in the United States and northern Mexico. Trans. San Diego Soc. Nat. Hist. 9(18) :87-162, figs. 1-8, maps 1-2, pi. 6. Werner, F. 1899. Beschreibung einiger neuer Schlangen und Batrachier. Zool. Anz. 22:114-117. . 1917. Versuch einer Synopsis der Schlangenfamilie der Glauconiiden. Mitt. Hamburg Zool. Mus. 34:191-208. CONTRIBUTIONS TO THE CONCEPTS OF REPRODUCTIVE CYCLES AND THE SYSTEMATICS OF THE SCALARIS GROUP OF THE LIZARD GENUS SCELOPORUS Hobart M. Smith and William P. HalP Abstr.\ct. — The concept of parity type is proposed as a criterion of relation- ship among members of the Sceloporus aeneus-scalaris complex, with members of 5. arncus being regarded as viviparous and those of S. scalaris, as oviparous. The viviparous S. goldmani is regarded as a relative of S. aerteus, and S. a. subniger is not regarded as intermediate between S. scalaris and S. aeneus. The proposal by Davis and Smith that S. a. aeneus is oviparous and specifically distinct (as S. aeneus) from S. a. bicanthalis (as 5. bicanthalis) is rejected. S. aeneus is hj-potliesized to have a biennial reproductive cycle. S. scalaris slevini auctorum is segregated into two subspecies: S. s. samcolemani subsp. nov. in the Sierra Oriental (northern end), and S. s. slevini sensu stricto in the Sierra Occidental (also northern end) of Mexico and of the adjacent United States. The systematics and phylogenetic history of the scalaris group of Sceloporus has long been viewed as enigmatic, beginning with the statement of the quandary by Smith (1939: 347-348) in explanation of the allocation of the northwestern member, slevini, to the species scalaris despite some contrary evidence. However, Smith and Pog- layen (1958: 13-15) allocated slevini to the species aeneus after another population exhibiting certain features of intermediacy be- tween the two polytypic species scalaris and aeneus was described {S. a. subniger) . Neither of the preceding accounts placed any overt emphasis upon parity types. Smith (1939: 332) made a passing comment that both oviparous and "ovoviviparous" species occur in the group. Evi- dence now available makes it certain that members of the scalaris complex are oviparous, and we here postulate that members of the aeneus complex are viviparous (a broader term, more certainly ap- plicable than ovoviviparity, which has come to imply absence of placental structures, whereas "euviviparity" implies their presence; both conditions exist in viviparous reptiles and only by microscopic techniques can they be determined). Anderson (1962) has demon- strated that slevini is oviparous, and accordingly its allocation with the scalaris complex is correct. The most critical doubt cast upon this hypothesis is the proposi- tion by Davis and Smith (1953: 102) that S. a. aeneus is oviparous, whereas S. a. bicanthalis is incontrovertibly viviparous (Smith, 1939: 356, and personal data). On this basis these taxa were elevated to specific status {S. aeneus, S. bicanthalis) . The data provided in Davis and Smith, however, strongly suggest that in S. a. aeneus viviparity does occur, but on a two-year cycle. Ten of 28 females collected from 25 July to 15 August "contained from 3 to 5 (average 4) large ova, the largest measuring 6 x 12 mm. ... In none . . . was there any recognizable evidence of embryonic development. . . . Many of these 'Deparlment of Environmental, Population, and Orpanismic Biology, University of Colorado, Boulder, 80302; and Departamento de Biologia, Universidad de Puerto Rico, Rio Piedras, Puerto Rico, OOOM. 97 98 GREAT BASIN NATURALIST Vol. 34, No. 2 specimens were taken at an elevation of 10,000 feet. . . ." Obviously these eggs could not be laid, at that elevation, and hatch before win- ter or survdve the winter without hatching. Almost certainly they would be carried in utero through the winter, the embryos develop- ing and the young being born in the spring. Indeed, further observa- tions in Davis and Smith support this contention: "12 of the others appeared to have oviposited and in the others (6) the ova were mi- nute, less than 1 mm. in diameter." The logical inference is that af- ter the young are bom in spring or early summer, ova start to de- velop but do not reach large size until the following year, and the embryos are not ready for birth until the spring or summer after that. We are not aware of any previous record of occurrence of a bien- nial reproductive cycle in lizards, although Fitch (1970) records the occurrence of biennial or triennial cycles (his opinion being that they are sometimes more properly interpreted as irregular cycles, their length being determined by environmental variables) in sever- al species of northern snakes {e.g., Crotalus viridis, Vipera aspis, and V. berus). Drs. Frank N. and Frieda C. Blanchard thought (personal communication) that a biennial cycle occurs in northern Michigan populations of Thamnophis sirtalis (Oliver, 1955: 244), and there is considerable evidence (obtained by Dr. Althea Gerrard of the Laboratory of Comparative Reproduction of the University of Colo- rado, under the direction of Dr. Richard Jones) that a biennial cycle occurs in populations of Thamnophis radix occurring at the extreme periphery of the range of the species in the foothill area near Boul- der, Colorado. Reproductive cycles exceeding one year in length re- quire, as a rule, considerable longevity, which is not ordinarily at- tributed to small lizards such as Sceloporus aeneus. Regardless of the apparent improbability of relative longevity in S. aeneus, a biennial viviparity does appear to occur in both subspecies of the species. De- finitive investigation of this apparently unique phenomenon is of high priority. It should be noted that although small size and probable short life (assumed on the basis of longevity records for related species: no records are available for members of the scalaris group of Scelopo- rus) mitigate against a biennial reproductive cycle (which might not be productive enough to offset the mortality rate), the very short season of activity and the perpetual coolness have the effect of in- creasing longevity. It has been amply documented that in squamatan reptiles and anurans, the members of subpolar or high-altitude popu- lations of any species or group, subjected to cold-induced inactivity much of the year, are longer-lived and have more protracted repro- ductive cycles than do their more temperate or subtropical relatives, which are active most of the year and have shorter reproductive cycles. Indeed, if other lizards with biennial reproductive cycles exist, they are almost certainly high-altitude or subpolar in distribu- tion. Andean lizards would be particularly suspect, since the repro- ductive cycles of the northern temperate lizards are well known and are consistently annual. However, S. aeneus and its viviparous rela- June 1974 smith, hall: sceloporus lizards 99 lives may truly be unique in this respect: whereas most, if not all, other high-altitude lizards are conspicuously heliotherm arboreals or saxicoles, acnrus is secretively terrestrial and graminicolous. S. sca- Inris is equally terrestrial and graminicolous but occurs at lower alti- tudes and is less secretive. S. microlcpidotus occurs at altitudes as great as acncus, but it lengthens its activity period conspicuously by insolation above ground, usually on trees; and insolation on rocks would have the same effect. Thus S. microlcpidotus and other high- altitude species of arboreal or saxicolous habits may, through ex- treme heliophily, so protract their activity that an annual reproduc- tive cycle may be maintained, whereas the terrestrial, seemingly nuich less heliophilous, S. acncus has been forced to adopt a biennial reproductive cycle to survive. Even if it were known to have a lower optimum activity temperature (not yet established) than do other sympatric species, it would not thereby necessarily escape the need for a biennial cjcle; Sphcnodon, with the lowest optimum activity temperature of any living reptile, has a protracted reproductive cycle (±: 13 months), although its freedom from temperatures in the freezing range permits a more rapid development of embryos than would be possible in the strongly seasonal weather to which S. acncus is exposed. Sphcnodon in the habitat of 5. aeneus would undoubtedly also require two years for completion of one reproduc- tive cycle. Although subniger was interpreted when described as occupying an ancestral position relative to the scalaris and aeneus complexes (Smith and Poglayen, 1958: 14), the clear evidence of intergrada- tion between subniger and a. aeneus (and absence of such evidence for subniger and scalaris) leads to the assumption that viviparity oc- curs in subniger, conclusive evidence that it is not an ancestral mem- ber of the scalaris complex. An important link in considerations of relationship and phylog- eny in the scalaris group is goldmani, which possesses a mixture of derived and primitive characters. Its viviparity — indicated by the occurrence of 9 well-developed young in the uteri of the holotype and one paratype of the species (Univ. Michigan Mus. Zool. 80896 and 77266, respectively, both from Charcas, San Luis Potosi) — is critical. Obviously goldmani falls with the acncus complex, not with the scalaris complex, and it seemingly constitutes the northernmost member of its complex. However, the species is known from only the hyjiodigm (Charcas, S. L. P.; Carneros, Coah.) localities and Ojo de Agua, Pablillo, Nuevo Leon (Liner and Olson, 1973: 54). One of us (Hall) has surmised that the species is a grassland inhabitant whoso existence has been critically attenuated by destruction of hab- itat through grazing by goats. A character progression from minimal derived states in northern populations {goldmani) to maximal de- rived states in southern poj)ulations {a. aeneus), through the inter- mediate i)opulations of subniger and bicanf /talis, supports Larsen's (1973) hypothesis of a northern origin for the group. Sympatry of the scalaris and acncus complexes is extensive in central parts of the Mexican plateau, although to the west the sea- 100 GREAT BASIN NATURALIST Vol. 34, No. 2 laris complex occurs without the other. A puzzling area of overlap occurs in the northern part of the Sierra Oriental, where populations referred to slevini are reported from areas occupied also by gold- mani. It appears that the scalaris-com^Xex representatives of that area are isolated from their closest relatives {scalaris slevini) in the western Sierra Occidental, Arizona, and New Mexico (Dixon and Medica, 1965: 73). Re-examination of material from the isolated northeastern populations reveals the existence of constant differences from paratopotypic scalaris slevini that justify recognition of a dis- tinct taxon we here name: Sceloporus scalaris samcolemani, subsp. nov. HoLOTYPE. — University of Michigan Museum of Zoology (UMMZ) 124670, an adult female taken between Providencia and La Paz, Nuevo Leon, Mexico, by P. H. Litchfield, on 16 July 1960. Paratypes. — Seventeen, including the Museum of Comparative Zoology (MCZ) 133167-73, 19 mi. S junction of roads to Doctor Arroyo and San Roberto, Nuevo Leon, 2250 m; and MCZ field nos. Y-25063 - Y-25071, 12 mi. SSE General Cepeda, 2250 m, Coahuila. All paratypes were slit the full length of the abdomen and rather distorted in preservation, but have retained their color and pattern with little change from life. Ilypoparatypes. Seven, including UMMZ 95220(7), from Galeana, Nuevo Leon (all extremely dis- colored, two juveniles 23 and 23.5 mm s-v), taken 12-17 July 1945, by Hellmuth Wagner. Diagnosis. — A short-legged (tibia 76-89 percent of snout-occiput length), oviparous member of the scalaris group of Sceloporus (of Smith, 1939), with essentially parallel rows of lateral scales, and two postrostrals; unique in the combination of usually (90 percent) a single canthal, dorsals 44-50, femoral pores 14-19 (66 percent 16 or more), maximum snout- vent length 51 mm, gular region barred or mottled (prominently in males, variable in females), and a con- sistently multicolored and patterned dorsum. Description of holotype. — Smaller dorsal head scales moder- ately keeled, larger ones smooth or weakly rugose; two postrostrals; four scales between nasals, two between median frontonasal and postrostrals; median and lateral frontonasals broadly in contact with each other, median narrowly separating prefrontals and contacting frontal; frontal divided, broadly contacting interparietal; two fronto- parietals on each side; three enlarged supraoculars on each side, sep- arated from median head scales by one row of small scales, from superciliaries by two rows (except rear supraocular, separated by one row); one large subnasal and one canthal on each side; two loreals; one preocular; one subocular; 2-3 large keeled lower post- oculars, three small upper ones; two rows of lorilabials ventral to loreals, one ventral to anterior half of subnasal, two ventral to poste- rior half; lorilabials reduced to one row over a length of 1-2 scales between subocular and labials; 3^/2 supralabials to below middle of eye. June 1974 smith, hall: scklopouus lizards 101 Fig. 1. Dorsal (left) and ventral (center) views of the holotype of Sceloporus scalaris samcolemani. UMMZ 124670, 51 mm SVL. Right, ventral view of a male paratj-pe of same, MCZ Y-25066, 41 mm SVL. Fivo infralabials; 3-4 pairs of chinshields, scales of anterior pair in medial contact, 2 scales between those of 2nd pair; labiomentals not reaching mental; one large medial auricular lobule; 6 scales be- tween upper enlarged postocular and car opening. Dorsal scales 47 from interparietal to base of tail; 44 ventrals from shoulder level to anus; 44 scales around middle of body; 17-17 femoral f)ores, the two series in medial contact; dorsal scales straight sided, with a strong medial keel, a short medial mucrone, one or no denticule/scale, no lateral mucrones. All ventral scales smooth; lamellar formula for toes 7-11-13-18-12 (7-11-14-18-13). Snout- vent length 51 mm; tail 62 mm; snout to occiput, 10.2 mm; hind leg 25 mm; tibia 8.(5 mm. 102 GREAT BASIN NATURALIST Vol. 34, No. 2 Color a brownish slate gray above, browner on head; a dorso- lateral light line on each side, largely on one scale row, separated by nine scale rows; ten U-shaped markings between shoulders and base of tail, dark-edged and light bordered to the rear, in each of two rows between dorsolateral light lines, separated medially by a space half as wide as markings; a row of similar marks lateral to dorso- lateral light line, bordered laterally by a very faint lateral line; a row of dark dots corresponding in position to the other dark mark- ings, lateral to lateral light line. Ventral surfaces whitish, with scattered gray pigment near chin, on chest, in preanal area and on ventral surfaces of the thigh. Large, well-developed eggs in the uteri, with no evidence of de- veloping young. Variation. — In the entire hypodigm, the dorsals vary from 44 to 50 (22: 44, six; 45, three; 46, four; 47, six; 48, two; 50, one), mean 46.1; ventrals 36-49 (14: 39, one; 40, one; 42, one; 43, two; 44, five; 45, two; 48, one; 49, one), mean 43.9; scales around mid- body 38-49 (17: 38, one; 39, one; 41, one; 42, three; 43, two; 44, five; 45, two; 47, one; 49, one), mean 43.3; femoral pores 14-19 (41: 14, eight; 15, six; 16, seven; 17, sixteen; 18, two; 19, two), mean 16.2. Scales between femoral pore series 0-1 (none, 11; one, 8); scales between second pair of postmentals 1-3 (1, four; 2, nineteen; 3, one); canthals 1-2 (1, forty-three; 2, five); snout-vent lengths 23- 51 mm (23,23.5,36, 37(2), 38(2), 40(2), 41 (2), 42(4), 43, 44, 45, 45.5, 47(3), 49, 51); tibia/snout-occiput ratio 76-88.8, mean (22) 81.25. The dorsal pattern in all paratypes is much like that of the holo- type, but the ventral markings are more prominent in some females and are conspicuous in males. In males the sides of the abdomen are dark blue, with no darker medial border but with scattered scales and white and irregular borders of scales black. No dark streaks tra- verse the blue patches. The gular region is conspicuously black-mot- tled or barred. Some females possess a vestige of the lateral abdomi- nal patches, even with scattered light scales, but the color is gray rather than blue. The gular region is also weakly barred or mottled in some females. Comparisons. — Sccloporus scalaris slevini is clearly the taxon that resembles S. s. colemani most closely — and indeed is the one with which the latter has long been united. S. s. samcolcmani is markedly different, however, in having gular bars, in being consis- tently multicolored above, and in having dorsals 44 or more (100 percent in 22), femoral pores usually (66 percent in 41 ) 16 or more, and s-v length not exceeding 51 mm. In 5. .?. slevini no gular mark- ings are present, the dorsal pattern is frequently unicolor, the dor- sals are usually 43 or fewer (84 percent in 44), the femoral pores usually 15 or fewer (92 percent in 88), and the s-v length frequently exceeds 51 mm (13 in a total of 44 measured), reaching 61 mm. The complete absence of the unicolor phase in S. s. samcolcmani is of course not a certainty, but that phase is so common in S. s. slevini Jime 1974 SMITH, HALL: SCELOPOHUS LIZARDS 103 and S. s. scalar is (no precise figures available) that the frequency in S. s. samcolemani is assuredly of a much lesser order, since 24 do not exhibit it. The single canthal character-state of S. s. sanicolernani is shared with S. s. unicanthalis as well as with S. s. slevini, but S. s. unican- thalis is distinctive in usually having the scales of the second pair of postmen ta Is in contact, whereas they are separated by one to three small scales in the other two subspecies. In addition, S. s. unican- thalis has a longer tibia and larger dorsals and reaches a larger size than either S. s. slevini or S. s. samcolemani. Remarks. — The southern limit of the range of S. s. samcolemani does not reach as far as 12 mi. SW Ahuacatlan, or Ciudad Maiz, both in San Luis Potosi, whence specimens typical of S. s. scalaris (UMMZ 126228, Univ. Illinois Mus. Nat. Hist. 21512, respectively) are available. These have 2-2 canthals; 44 and 35 dorsals, 16-19 and 108 102 96 Fig. 2. Distribution of the subspecies of Sceloporus scalaris. Localities as given in text for S. c. samcolemani, from the literature and museum lists for others: especially Duellnian (1961) for Michoacan, Dixon et al. (1972) for Queretaro, and t)rake (1950) for Durango records of S. s. scalaris, and Grant and Smith (1960: 40) for an Jalisco record of S. s. unicanthalis. 104 GREAT BASIN NATURALIST Vol. 34, No. 2 14-15 femoral pores, respectively; and s-v measurements of 58 cT and 53 9 mm, respectively. The female is unicolor above; the male has faint dark gular bars. The female has the smallest dorsal count of any recorded specimen; Smith (1939:347) records counts no lower than 37. The type series of 24 specimens includes 16 adult females, all with large eggs in the uteri, none showing evidence of embryonic development. In view of the lower altitude and longer season for activity than in S. aeneus, the early-middle July dates suggest ample time for egg deposition and hatching later the same year. Apparent- ly, therefore, the population is oviparous and on that basis as well as pattern seems properly allocated with S. scalaris rather than with S. aeneus. Localities other than those represented by the hypodigm include Hda. Pablillo, above Galeana, Nuevo Leon (Smith, 1939:349), the Gomez Farias region, Tamaulipas (Martin, 1955: 173); and Cedri- tos, Coah. (Amer. Mus. Nat. Hist. 77245). The subspecies is named for Dr. Sam Coleman, who wrote the programs for processing the enormous quantity of data pertaining to the herpetology of Mexico and who thereby has contributed far more to a synopsis of Mexican herpetology than he realizes. Acknowledgments. — We are much indebted to Dr. Arnold Kluge, Dr. D. F. Hoffmeister, and Dr. Ernest E. Williams for the privilege of studying material under their care in their respective institutions. Literature Cited Anderson, J. D. 1962. Egg laying and nesting in Sceloporus scalaris slevini. Herpetologica 18(3) : 162-164. Davis, W. B., and H. M. Smith. 1953. Lizards and turtles of the Mexican state of Morelos. Herpetologica 9:100-108. Dixon, J. R., C. A. Ketchersid, and C. S. Lieb. 1972. The herpetofauna of Queretaro, Mexico, with remarks on taxonomic problems. Southwestern Naturalist 16:225-237, fig. 1. , AND P. A. Medica. 1965. Noteworthy records of reptiles from New Mexico. Herpetologica 21:72-73, fig. 1. Drake, J. J. 1958. The brush mouse Peromyscus boylii in southern Durango. Publ. Mus. Michigan St. Univ., Biol. Ser. 1(3): 97- 132, pis. 1-3, fig. 1-8. DuELLMAN, W. E. 1961. The amphibians and reptiles of Michoacan, Mexico. Univ. Kansas Publ. Mus. Nat. Hist, 15(1): 1-148, pis. 1-6, figs. 1-11. Grant, C, and H. M. Smith. 1960. Herpetozoa from Jalisco. Herpetologica 16:39-43. Larsen, K. R. 1973. Speciation in the genus Sceloporus, as determined by cranial osteology and other characters. Provo, Utah, Brigham Young Univ. Ph.D. Dissertation, xii, 272 pp., 64 figs. Liner, E. A., and R. E. Olson. 1973. Adults of the lizard Sceloporus torquatus binocularis Dunn. Herpetologica 29(l):53-55, fig. 1. Martin. P. S. 1955. Herpetological records from tlie Gomez Farias region of southwestern Tamaulipas, Mexico. Copeia 1955(3):173-180. Oliver, J. A. 1955. The natural history of North American amphibians and reptiles. New York, Van Nostrand. xi, 359 pp., 12 pis., 74 figs. Smith, H. M. 1939. The Mexican and Central American lizards of the genus Sceloporus. Field Mus. Nat. Hist., Zool. Ser. 26:1-397, figs. 1-59, pis. 1-31. , AND L PoGLAYEN. 1958. Noteworthy herptiles from Mexico. Herpeto- logica 14: 11-15, pi. 1. GROWTH AND DEVELOPMENl^ OF THE WESTERN HARVEST MOUSE, REITHRODONTOMYS MEGALOTIS MEGALOTIS Gary- H. Richins^-^, H. Duane Smith', and Clive D. Jorgensen' Abstract.- - Growth rates, gestation period, litter size, reproductive age, sex ratios, and development were studied on 198 litters of Reithrodontomys megalotis megalolis (Baird). Growth was characterized by several standard skull and body measurements and was partitioned into four phases of 1-3, 4-12, 13-22, and 23-70 days. Growth was best described by measurements of tail length, ear length, and dried eye-lens weight. Reproductive activity began as early as 38 days for females and 59 days for males. The gestation period was 22 days, and the mean litter size was 3.83 (range 1-7). Sex ratio was 53.49 percent males to 46.51 percent females. Repro- ductive efficiency was 53.53 percent. The development of R. m. megalotis was very similar to tliat of R. m. dychei. Early breeding, postpartum estrous, year-round breeding, high reproductive efficiency-, and a short gestation period contribute to a high reproductive potential in /?. m. megalotis. Small mammals occupy vital positions in the bioenergetics of North American deserts because they are important primary con- sumers, but man}- of their interactions within the ecosystem are not understood. Productivity estimates of rodent populations seldom take into account the biomass and biotic potential of young animals, especially when these estimates are made solely on the basis of trap- pable animals with disregard for such dynamic variables as litter size, gestation period, reproductive age, sex ratios, and biomass. Because this type of data is difficult to obtain in the field, it is usually provided by laboratory studies. This study will define the growth and development of Reithrodontomys megalotis megalotis (Baird), a small cricetine found throughout much of North America. The objectives of this study are (1) to characterize growth from birth to maturity by measurements of various body and skull parameters, (2) to determine gestation period, litter size, reproduc- tive age, sex ratios, and development of behavioral sequences, and (3) to correlate these data, where possible, with similar studies on related species. Literature Review Growth rates have been studied for several small mammals (e.g., Meyer and Meyer, 1944; Pournelle, 1952; Chew and Butterworth, 1959; Layne, 1960; Goertz, 1965; Ilayden and Gambino, 1966; Jones, 1967; Horner and Taylor, 1968), but Reithrodontomys, par- ticularly R. megalotis, remains basically unstudied. Life histories of R. humulis (Holding and Royal, 1952; Layne, 1959; Kaye, 1961), R. montanus (Leraas, 1939), R. fulvescens (Svihla, 1950), and 'Departn\cnt of Znolopy, Brighani Young University, Prove, Utah 84f)02 ^Currently Wildlife Biologist, Danics and Moore. Cincinnati . Ohio 105 106 GREAT BASIN NATURALIST Vol. 34, No. 2 R. megalotis (Svihla, 1931; Smith, 1939; Brummel, 1961) have been partially studied. Bancroft (1966) completed an intensive study of reproduction, development, and behavior of R. megalotis dychei from Kansas, but no correlation of age with growth was attempted, nor has Bancroft's (1966) information been published. Materials and Methods Animals used in this study were reared from 20 pairs of R. m. megalotis captured approximately 19 km SE of Benmore Guard Station, Tooele Co., Utah, from 1-9 September 1971 and subsequently housed in a laboratory at Brigham Young University. The animals were caged in standard small-mammal cages with wood shavings pro- vided as substrate and cotton as nesting material. Free water and Purina mouse breeder chow were available continuously. Laboratory temperature was held at 22 =t 2 C with a photoperiod of 12 hr. light and 12 hr. dark; varying intensities simulated dawn and dusk. Ani- mals brought into the laboratory were sexed, paired, and checked daily for food, water, and presence of a litter. When a litter was born, each litter member was marked by sequential toe clipping. The following measurements were taken daily from days 1-22 and then weekly for 7 additional weeks on a sample of 100 animals: (1) body weight, (2) total length, (3) tail length, (4) ear length from notch, and (5) hind-foot length. Daily observations on development and behavior were recorded for each litter. Body weight was determined to the nearest 0.05 g; total and tail lengths, to the nearest 0.5 mm; and ear and hind-foot lengths, to the nearest 0.01 mm. After the eyes had opened and the indi- viduals had become more active (about 10 davs of age), they were anesthetized with Penthrane.^ Ten individuals were killed daily from 1 to 22 days and weekly from 4 to 10 weeks for additional studies on skull measurements and eye lens weights to correlate age with the external measure- ments. On the day that an animal was to be killed, it was removed from the nest and administered a lethal dose of Penthrane. Standard body measurements were taken and the carcass placed in a 10- percent formalin solution to fix the eye lenses. After four days the carcass was taken from the formalin solution; the head was removed and skinned; and the lenses were extracted with a curved forceps after a slit had been made in the cornea. The lenses were stored in vials of 10-percent formalin, after which they were dried at 100 C for one week before weighing to the nearest .0001 g. After the lenses had been extracted, the skull was stained and the (1) total length, (2) zygomatic breadth, (3) foramen magnum height, (4) mastoidal breadth, (5) nasal length, and (6) cranium width were measured to the nearest 0.01 mm with dial calipers. Staining followed the methods described by Humason (1967), except that the concentration of alizarine stock solution used was increased 10 times. ^Abbott laboratories, North Chicago, Illinois. June 1974 richins, et al: harvest mouse 107 The instantaneous relative growth rate (IGR) described by Brody (1945) and Lackey (1967) was calculated to express growth as a rate between times of measurement and percentage of adult size. The IGR is expressed as dW/dt/W, where W is the parameter measured at the instant the rate of change of dW/dt is measured. Because it is not entirely possible to develop the "instantaneous" rate of growth, it was necessary to integrate the infinite number of growth rates to derive W = Ae'^K This is conveniently written as InW = hiA + kt where: InW is the natural logarithm of the variable measured (W) at ^ = 0, In A is the natural logarithm of a theoretical constant based on the value oi W at t = 0, k represents the instantaneous relative growth rate (when multiplied by 100, k = percentage growth rate), and t is time (age in days). For comparative purposes, the IGR (k) is determined with lnW„ - lnW„_, kn - — *n ~ *n-i Thus k is definite and can be used to compare differences in rates of growth. Reproductive efficiency, defined as the weight of the litter multi- plied by 100 and divided by the weight of the mother (Kaye, 1961) was determined for 19 adult female R. m. megalotis. In all instances both measurements were taken immediately after birth. Results and Discussion Growth Rates The instantaneous relative growth rates k for body weight, total length, tail length, ear length, and hind-foot length that were de- termined each day for 100 R. m. megalotis from 1 to 22 days and then weekly for the next 7 weeks are reported in Table 1. Growth rates {k) were also calculated for dried eye-lens weight, skull total length, zygomatic breadth, foramen magnum height, mastoidal breadth, nasal length, and cranium width on 10 animals each day from 1 to 22 days and then weekly for the next 7 weeks. This re- duction in sample size from 100 to 10 per day was necessary, owing to the time required to raise animals to the proper age and to prepare the skulls and lenses. Previous studies (Smith and Jorgensen, 1972) involving com- puter analyses of growth data using linear, quadratic, cubic, com- bined linear-(iuadratic, and combined linear-quadratic-cubic distribu- tions with different time intervals were conducted to characterize species growth curves. Since the growth curves turned out to be curvilinear, none of the time intervals used with the above distribu- tions aTid combinations of distributions adequately described the entire growth curve for any parameter. Brody's (1945) logarithmic approach adopted in this paper and used by Smith and Jorgensen (1973), however, does accurately describe the growth of R. m. 108 GREAT BASIN NATURALIST Vol. 34, No. 2 be fo m vo (N ■^ (O IT) o^ 00 CT) in o O CS) Ol Ol vn (£5 00 CT) vn 00 o CM po lo vn (N t^ vn o -^ (M ^o -^ o -^vnoopo CM 00 Cs) ro « CM o t^ ooiovn (Y^ CN oi O) (o -"i- o a> CT) c^) m T^ T^t^\r)•>-' ■ co •rt O 00 t^ t^ C^ O ro ■.- -^ -^ tN. tJ- t}- fO T-. WTO t}- oi CO 00 (^ o) vn 00 t^ (Nt^ rOOOOloo int^co-^ ■ •r-< \n ■^ t^ t^ t^ po •^ vn xn -"^ 00 t^ Tj- ^ vn •»}■ oo o ■^ vn ■^ -^ "^ CM 00 tJ- ^ (OxnroO oc^-^o — c o o o OOOO -hOOO int^oo t^oo->J-oi rorornrn 00 oo O) O oo 00 vn o 00 CO -rt o C^OOO OOOO o 00 t>. ■>}■ 05 ■<}- cT> o cr> T-H t^ vo --H 00 CT) o oi t>. ■.ot^ai O'^oooo cO'T-cNcsi cvn-^0 ooccro-^ •^OOO OOOO -r-^OO (TiOOO OOOO ■"J'-^OO (!>5 0i COfO 00 CM t^ vn t}- O CO O CM ■}■ ro ro ■r-. CM ^CS) CM •^CSIO CMtNO CMCMO CMCMO rO'^CMt^ rO'^CMt^ rO-^CMt^ rO'^CMt^ ■^•^roro -r-crj-roro •^■"J-coro T-c-? -^CN ■rtCM ^CM t W ^•5 c a 0) June 1974 RICHINS, ET AL: HARVEST MOUSE 109 COOivnO lOO-<}-ro t^-*»r)CO t^ m tJ- o 1 00 Oi CO -"HO t^ ■«}• xr> fo 00 -^ -^ y-> •+ O xn CO Q (O ■+ o ro o CM (o — (N •rHxn-r-io o -^ o o o vr> o o "^(^(o-^ csi -—i o PO -rH (>o t^ tOtO^^O O t^ O 00 to (O fO CSI -"J- cq to ro -^ 00 '^ -r- 00 t^ T-. .,-1 O) to O O t^ Ci CT) O CM ■. — ro tO tO -^ O to fO •^ O •^ Cs) •^ O vn CM O O ro ro O O vn (M O ■* CSI o o oooo oooo oooo oooo oooo oooo — cMoo -- — to vnro •«}• to to t^ o) po tooo OOlCSl CM 00 O) — CTi in fo vn (OOOOCM CMCQCMCM «-".-CMCM «^ CM ■* O CMOOCOO 00 O t^ (T) rocji-^xT! PO(75tOO oouicrito POCMtOCTi -H^xntM iflTi-'rf-c^ t>.ooocM xnt^O'^ 00 o -^ 00 Oo^"^ ^^fvjCS o^^^ t^ -r-toxn 00 00 t^ -^ t^ CM oooo J^O) ■^ CM CMCMO (?4CMO C^CMO -' CM ^- PO -^ CM t^ PO -^ CM t^ •^ ■^ PO PO -ri T^ PO fO ■•-htJ-POPO ■<-h rf CO pO ■^ tJ- PO pO ii •^ PO pO •"-■CM -iCM —"CM ■>-'C>l •^CM -^CM -2 3 C C Hj3 C/Ji-J NCQ 2-£ CO bc ^•5 to to CO tu CO 'p^ 110 GREAT BASIN NATURALIST Vol. 34, No. 2 megalotis for the time intervals (growth phases) 1-3, 4-12, 13-22, and 23-70 days. Not only is growth characterized by these four time intervals, but developmental sequences also tend to correspond to them. From 1 to 3 days the young harvest mouse is adjusting to the rigors of life outside the uterus. Because there is a rapid elongation of the ex- tremities and a constant gain in weight (Figures 1-4), but little visible external development, there is probably considerable internal change taking place. During the 4-12-day growth phase, however, a great deal of external change is manifest. The extremities and body continue to elongate, and the animal increases in weight — although at a slower rate (Figures 1-4). The ears unfold, and the pinnae begin to develop. The toes separate and become functional. Pelage develops, and the teeth erupt from the gums. The eyes open, and there is increased activity about the nest. The external auditory meatus opens, and the sense of hearing develops. Little external change other than a continued elongation of the extremities (Figures 1-4) characterizes the 13-22-day growth phase. Near the beginning of this period the young animals begin to take solid food, and by 22 days of age they are completely weaned. From 23 to 70 days the growth of extremities all but ceases, and body weight is stabilized (Figures 1-4). Sexual development proceeds, and by 70 days both sexes have reached maturity. The R^ values (Table 1 ) indicate the amount of variation ac- counted for by growth of the parameter during a particular time interval. When R^ values are converted to r, they indicate statistical significance. When time is partitioned into growth phases, a signifi- cant r indicates correlation between hiW and age and is used to establish confidence in k. Tail length, ear length, and dried eye-lens weight provided the best correlations of relative growth with age (Table 1), r being significant during all four growth phases (n = 10). Although dried eye-lens weight was the best indicator of growth — and, ultimately, of age — in the laboratory, tail length or ear length would be the best parameter to measure in the field because these measurements are easily taken and do not involve killing the animal. Significance was spotty among the other parameters for the various growth phases, but all of the parameters were significantly correlated with age dur- ing the 4-12 day interval when the majority of growth and develop- ment occurred. Graphic representations of means, standard errors, and instan- taneous relative rates {k) were prepared for the three most signifi- cant growth parameters — tail length (Fig. 1), ear length (Fig. 2), and dried eye-lens weight (Fig. 3) as well as for body weight (Fig. 4). The close correlation of k with the actual data means indicates that the k values for each of the four growth phases characterize the growth of R. m. megalotis. If a detailed growth analysis as is pre- sented here is unfeasible, a workable growth curve could be presented using the means and standard errors; but it is imperative that these be -plotted on a logarithmic scale for the curve to be meaningful. June 1974 RICHINS, RT AL: HARVEST MOUSE 111 /^=.0084 / ^=.1300 lA 21 28 35 42 Age (days) 49 56 63 70 Fig. 1. Means, standard errors (p = .95), and instantaneous relative growth rates for tail lengtli of Reithrodontomys megalotis megalolis. The presence of four growth j)hases (1-3, 4-12, 13-22, and 23-70 days) in R. m. megalotis differs from the results of other workers. Layne (1959) found two growth phases in R. humulis: 0-3 weeks and 3-7 weeks. Bancroft (1966) found three growth phases in R. m. dychei: 0-3 weeks, i-7 weeks, and 7-10 weeks. 112 GREAT BASIN NATURALIST Vol. 34, No. 2 100 90 80 70 60 50 AG 30 20 10 ?;=.0015 /c=.0467 fe=.1051 /i=.1329 lA 21 28 35 42 A9 Age (days) 56 63 70 Fig. 2. Means, standard errors (/? = .95), and instantaneous relative growth rates for ear length of Reithrodontomys megalotis megalotis. Age prediction Because the correlation of some growth parameters with age is significant, one might consider using these parameters to predict age. Although the process seems evident at first (it would involve simply reading the predicted age from a graph), the results cannot June 1974 RICHINS, ET AL: HARVEST MOUSE 113 A:=.0010 ^=.0539 K=.0888 ^=.3401 14 21 28 35 42 Age (days) 49 56 63 70 Fig. 3. Means, standard errors {p = .95), and instantaneous relative growth rates for dried eye-lens weight of Reithrodontomys megalotis megalotis. be interpreted with statistical confidence because of the lack of variation among days. Calculation of confidence limits about the regression line presents special problems, because the X axis (age) is a non-random variable, selected arbitrarily by the investigator; thus, only the regression of Y on X can be estimated with confidence. 114 GREAT BASIN NATURALIST Vol. 34, No. 2 1000 900 800 700 600 500 400 300 200 100 90 80 70 60 50 40 30 20 10 9 8 7 6 5 4 W^ /c=.0337 /c=.0159 fe=.1820 ^=.4867 14 21 28 35 42 Age (days) 49 56 63 70 Fig. 4. Means, standard errors (p = .95), and instantaneous relative growth rates for body weight of Reithrodontomys megalotis megalotis. When a least squares regression equation is used to predict ages of unknown animals, age becomes a random variable (Dapson and Irland, 1972; Smith and Jorgensen, 1972); consequently, the analy- ses of these data to provide age determinations await nonparametric procedures. June 1974 richins, et al: harvest mouse 115 Gestation Period The minimum gestation period of 22 days, as determined by the minimum interval between successive litters, was shorter than pre- viously reported: Svihla (1931) reported 23 days and EcoDynamics (1971) reported 25-28 days as the gestation period for R. m. mega- lotis. One litter was born after only 20 days, but because all four young died within a few hours, the litter was assumed to be pre- mature. Our results are similar to those of other workers. Bancroft (1967) reported 23 days as the gestation period for R. m. dychei, while Layne (1950) listed 24 days for R. humulis, although later Kaye (1961) reported 21 days. Leraas (1938) reported 21 days as the gestation period for R. niontanus. Litter Size Litter size had a mean of 3.83 and ranged from 1 to 7 (Table 2), which is comparable to Bancroft's (1966) reporting of 3.84 and a range of 1-7 for R. m. dychei, although litter size had been stated earlier by Long (1962) to include up to nine. Svihla (1931) listed an average litter size of 2.60, ranging from 1 to 7, for R. m. ni- grescens and R. m. megalotis. Other findings on litter size of R. megolotis vary somewhat, although always within the ranges pre- viously stated: 4-6 in Oregon (Bailey, 1936); 1-4 and a mean of 2.0 in South Dakota (Zukerman, 1935); 1-7 for captive bred, with a mean of 4.27, and up to eight for laboratory bred (Egoscue, Bitt- menn and Petrovich, 1970; Egoscue, 1972). Egoscue (1972) further reported an average litter size of 5.64 for wild caught females, rang- ing from 3 to 6, based on embryo counts of 56 placenta examinations. Litter size varied with the number of litters females had already produced. The mean litter size (3.29) of 69 first litters of R. m. megalotis increased to 4.71 for 17 fourth litters before decreasing to 3.60 for five ninth litters (Table 2). Bancroft (1966) stated that mean litter size of three R. m. dychei was 4.30 for first litters, in- creased to a maximum of 6.00 for fifth litters, and decreased to a low of 3.30 for eleventh litters. This would indicate that female R. m. megalotis reach their maximum reproductive capabilities from 20-25 weeks of age — assuming that most are bred by 10 weeks and produce a new litter approximately every 3-3.5 weeks. While this reproduc- tive rate is possible under laboratory conditions, it is likely to proceed more slowl}- in a field situation in which successful male-female en- counters are logistically reduced. Sex Ratio The sex ratios of newly born R. m. megalotis resulted in 53.5 percent males and 46.5 percent females (Table 2). This sex ratio varies from those previously re})ortod. Egoscue (1972) lists a sex ratio of 50.99 percent males to 49.01 })ercent females for 901 R. m. megalotis. whereas Bancroft (1966) ref)f)rts a sex ratio at birth of 51 percent mal(>s to 49 })erceiit females for ^^H R. m. dychei. Layne 116 GREAT BASIN NATURALIST Vol. 34, No. 2 Table 2. Mean Number of Young Reithrodontomys megalotis megalotis Born in the Laboratory per Successive Litter. Successive Number of Average Percentage Litter Litter Number Litters Size Males Females Males Females 1 69 3.29 1.90 1.39 52.7 42.3 2 37 3.86 2.24 1.62 58.0 42.0 3 30 4.23 1.90 2.33 44.9 55.1 4 17 4.71 2.76 1.94 58.8 41.2 5 12 4.25 2.17 2.08 50.9 49.1 6 10 4.10 1.80 2.30 43.9 56.1 7 9 4.00 2.00 2.00 50.0 50.0 8 8 3.88 2.00 1.88 51.6 48.4 9 5 3.60 1.60 2.00 44.4 55.6 10 1 5.00 2.00 3.00 40.0 60.0 1-10 198 3.83 2.05 1.78 53.49 46.51 (1959) found an even sex ratio at birth in R. humulis, but Kaye (1961) reported a ratio of 33.4 percent males to 66.6 percent females for 27 R. humulis. Reproductive Age Svihla (1931) reported 128 days as the youngest age at which a male or female R. m. megalotis had been found to breed. Leraas (1938) found that female R. montanus albescens gave birth as early as 90 days. Layne (1959) reported that one female R. humulis humulis became pregnant at 77 days and that males had enlarged testes as early as 49-50 days. Bancroft (1966) listed one female R. megalotis dychei as giving birth at 72 days and males as reaching sexual maturity at from 56 to 70 days. During this study of R. m. megalotis, a female 29 days old was paired with a male 50 days old. This pairing resulted in a litter of four young born when the female and male parents were 62 and 83 days of age respectively. Ciranting a 22-day gestation period, R. m. megalotis can become reproductively active as early as 38 days and 59 days for females and males respectively, which supports the 35 days reported but unpublished by EcoDynamics (1971) but which supplants previously published data. Reproductive Efficiency R. megalotis has the highest reproductive efficiency reported for any mammal (Bancroft, 1966). Reproductive efficiency was calcu- lated for 19 adult female R. m. megalotis resulting in a high of 53.33 percent for a litter of six. The mean reproductive efficiency of the 19 animals was 43.44 ± 2.91 percent (31.5-53.3 percent). Frank (1956) listed Microtus arvalis as having the highest reproductive efficiency of any mammal, the litter weight amounting to 53.2 per- cent of the mother's weight, but Bancroft (1966) reported 62.5 per- cent for a litter of five R. m. dychei. Kaye (1961) listed a high 50.5 percent for R. humulis. Dunaway (1962) listed 77.7 percent as the June 1974 richins, et al: harvest mouse 117 reproductive efficiency of a litter of eight R. humulis, but because the measurements were taken some 10-15 hours after birth, it is probably erroneous. Reproductive Potential Several of the foregoing factors contribute to the high reproduc- tive potential of R. mcgalotis: early breeding, postpartum estrous, year-round breeding, high reproductive efficiency, and a short gesta- tion period. Early breeding allows for a longer reproductive life. Postpartum estrous which was evidenced during this study enables R. m. mega- lotis to maximize the number of litters born during a single season. In Utah the breeding season runs from mid-May to early September. Therefore, a harvest mouse born in early June could have as many as two litters its first year. A mature harvest mouse could have 4-5 litters during this time and could contribute 15-20 individuals to the population in a single year. A high reproductive efficiency allows a greater percentage of energy intake to go for reproduction, thus maximizing reproductive potential. A short gestation period in concert with postpartum estrous is also advantageous because it tends to minimize the time between successive litters. Reproductive potential for any species is clearly a function of litter rapidity and the length of time reproductive activity occupies in the life span of females. Under laboratory conditions, these fac- tors can be maximized, although field conditions obviously impose yet another constraint- — environmental limits, including population interactions. Bancroft (1966) reported postpartum estrous in R. m. dychei; and, although they can breed year-round in the laboratory, he doubts this would persist in nature, except perhaps in the southern part of their range. Development Condition at hirth. Neonatal R. m. megalotis are similar in ap- pearance to R. m. dychei (Bancroft, 1966) in that they are smooth and pink and are naked except for natal vibrissae. Eyes are closed and covered with a transparent membrane, including the crease marking the persisting fusion of upper and lower eyelids. Pupils are clear, with the exception of coloration in the iris. Ear pinnae are folded and appressed against the head. Toes are still fused, and nails are not yet present. The young vocalized faintly when handled. One day. There is little morphological change, although the ear pinnae are sometimes unfolded. A gray dorsal stripe is visible, and the skin has lost its newborn transparency. Hair is not evident, although Bancroft (1966) reported fine hair visible upon close ex- amination of the dorsum in R. m. dychei. Layne (1959) also noted abundant short and long hair on the dorsum of day-old R. humulis. Two days. Ear pinnae have unfolded, and the dorsal pigmenta- tion is darker, with dorsal hairs visible when held to the light. Toes 118 GREAT BASIN NATURALIST Vol. 34, No. 2 are about one-third separated and claws are visible. Young move their feet rapidly and vocalize sharply when handled. Three days. The young cling to the female's nipples and are often dragged around the cage several times before falling off when disturbed. Vocalization is a high-pitched squeak, 2-8/second. The toes are about one-half separated. Eyelid slits are visible. The dorsal pigmentation is almost black, and a white, scaly epidermal material observed by Bancroft (1966) in R. m. dychei and by Layne (1959) in R. humulis is present on the dorsum. Four days. The toes are almost separated. The hair on the head is thicker and colored a pale gray brown. Bancroft (1966) stated that at this age R. m. dychei had claws approximately 5 mm long and that the ears were only fleshy lobes. Five days. Dorsal furring extends about one-third of the way along the back, and the upper and lower incisors have erupted in some. Bancroft (1966) stated that the lower incisors had erupted in R. m. dychei. Layne (1959) also reported that the lower incisors were visible in R. m. megalotis on the fifth day. Six days. There is little external change evident, although Ban- croft (1966) reported that hair had appeared on the dorsal side of the tail in R. m. dychei. Seven days. The dorsum is completely furred, and ventral hair is more pronounced. Mammae are visible in females. Eight days. The external auditory meatus is open, and the young are very active and vocal. Opening of the external auditory meatus almost always precedes the opening of the eyes by one day. Nine days. The venter, feet, and pinnae are completely furred, and the eyes are beginning to open. Smith (1936) reported that the eyes first opened at six days but were of little use for 48 hours. Bancroft (1966) found that the eyes were beginning to open in R. m. dychei at seven days but were not yet fully functional. Svihla (1931) found that the eyes opened on days 11-12 in R. m. megalotis, as did Brummel (1961). Leraas (1938) observed that the eyes opened on the eighth day in R. montanus. Layne (1959) reported that the eyes opened at from 7 to 10 days in R. humulis. In the present study, the eyes of R. megalotis megalotis opened at from 9 to 13 days. Ten days. The eyes are open and fully functional in the ma- jority of the young. Fourteen days. Face washing is observed. There is very little vocalization. Observations of the feces suggest that the young have started eating solid food. Sixteen days. Although nursing is observed until the sixteenth day, the majority of the young are completely weaned. There is no detectable change in appearance or behavior on days 11, 12, 13, and 15. Summary and Conclusions The growth date of R. m. megalotis can be divided into four distinct phases: 1-3, 4-12, 13-22, and 23-70 days; whereas R. m. June 1974 richins, et al: harvest mouse 119 dychei and R. humulis hurnulis have three and two growth phases respectively (Bancroft, 1966; Layne, 1959). Growth of R. m. mega- lotis through all four phases is most accurately characterized by tail length, ear length, and dried eye-lens weight measurements. R. m. megalotis had a 22-day gestation period, one day shorter than previously reported. Litter size varied from 1 to 7, and sex ratios at birth were approximately even. Western harvest mice became reproductively active earlier than previously reported. One R. m. megalotis became pregnant at 38 days of age. A high reproductive efficiency, 53.33 percent in this study and 62.5 percent (Bancroft, 1966), has been evidenced in R. m. megalotis. R. m. megalotis has a high reproductive potential because of early breeding, postpartum estrous, year-round breeding, high re- productive efficiency, and a short gestation period. The development of R. m. megalotis closely followed that of R. m. dychei as reported by Bancroft (1966), but differed slightly in that the eyes opened at from 9 to 13 days in R. m. megalotis while in R. m. dychei the eyes opened at from 7 to 11 days. In the laboratory R. m. megalotis grows faster, has a shorter gestation period, has larger litters, breeds earlier, and has a higher reproductive efficiency than previously reported for this genus. Literature Cited Bailey, V. 1936. The mammals and life zones of Oregon. N. Amer. Fauna 55:1-416. Bancroft, W. L. 1966. Reproduction, development and behavior of the western harvest mouse, Reithrodontomys megalotis. Unpub. Thesis, U. of Kansas. 80 pp. . 1967. Record fecundity for Reithrodontomys megalotis. J. Mammal. 48:306-308. Brody, S. 1945. Time relations of growth of individuals and poulations. In: Bioenergetics and Growth. Reinhold, N. Y. 1023 pp. Brummel, C. N. 1961. Some aspects of the life history and ecology of the western harvest mouse in southeastern South Dakota. Proc. South Dakota Acad. Sci. 40:85-92. Chew, R. M., and B. B. Butterworth. 1959. Growth and development of Merriam's kangaroo rat, Dipodomys merriami. Growth 23:75-96. Dapson, R. W., and J. M. Iri.and. 1972. An accurate method of determining age in small mammals. J. Mammal. 53(1): 100-106. Dunaway, p. B. 1962. Litter size record of eastern harvest mouse. J. Mam- mal. 43(3): 428-429. EcoDynamics. 1971. Ecological studies in western Utah, Annual Report. EcoDynamics Inc., Salt Lake City, Utah. 116 pp. Egoscue, H. J., J. G. BiTTMENN, AND J. A. Petrovich. 1970. Some fecundity and longevity records for captive small mammals. J. Mammal. 51(3):622- 623. Frank, F. 1957. The causalty of microtinc cycles in Germany. J. Wild). Mgmt. 21 (2): 113-1 21. GoERTZ. J. W. 1965. Sex. age, and weight variation in cotton rats. J. Mam- mal. 46:471-477. Hayden, p., and J. J. Gambino. 1966. Growth and development of the little pocket mouse, Perognathus longimembris. Growth 30:187 197. 120 GREAT BASIN NATURALIST Vol. 34, No. 2 Holding, B. F., and 0. L. Royal. 1952. The development of a young harvest mouse, Reithrodontomys. J. Mammal. 33:388. Horner, B. E., and J. M. Taylor. 1968. Growth and reproductive behavior in the southern grasshopper mouse. J. Mammal. 49:644-660. HuMASoN, G. L. 1967. Animal Tissue Techniques. 2nd. ed., W. H. Freeman and Co., San Francisco, pp. 179-181. Jones, C. 1967. Growth, development, and wing loading in the evening bat, Nycticeius humeralis (Rafinesque). J. Mammal. 48:1-19. Kaye, S. V. 1961. Laboratory life history of the eastern harvest mouse. Am. Midi. Nat. 66:439-451. Lackey, J. A. 1967. Growth and development of Dipodomys stephensi. J. Mammal. 48:624-632. Layne, J. N. 1959. Growth and development of the eastern harvest mouse, Reithrodontomys humulis. Bull. Florida State Mus., Biol. Sci. 4(2):61-82. Leraas, H. J. 1938. Observations on the growth and behavior of harvest mice. J. Mammal. 19:441-444. Long, C. A. 1962. Records of the reproduction for harvest mice. J. Mammal. 43:103-104. Meyer, B. J., and R. K. Meyer. 1944. Growth and reproduction of the cotton rat, Sigmodon hispidus hispidus, under laboratory conditions. J. Mammal. 25:107-129. Pournelle, G. H. 1952. Reproduction and early postnatal development of the cotton mouse, Peromyscus gossypinus gossypinus. J. Mammal. 33:1-20. Smith, C. F. 1936. Notes on the habits of the long-tailed harvest mouse. J. Mammal. 17(3) : 276-278. Smith, H. D., and C. D. Jorgensen. 1972. Demographic and individual growth studies for Dipodomys ordii and Peromyscus maniculatus. Desert Biome IBP. Progress Rpt. December 31, 1972. Smith, H. D., and C. D. Jorgensen. 1973. Demographic and individual growth studies for Dipodomys ordii and Peromyscus maniculatus. Desert Biome IBP. Final Rpt. December 31, 1973. Svihla, R. D. 1930. Notes on the golden harvest mouse. J. Mammal. 11(1): 53-54. Svihla, R. D. 1930. Notes on the desert and dusky harvest mouse {Reithro- dontomys megalotis megalotis and R. m. nigrescens). J. Mammal. 12:363- 365. Zukerman, S. 1953. The breeding seasons of mammals in captivity. Proc. Zool. Soc. London. 122:827-950. BODY SIZE, BODY COMPOSITION, AND BEHAVIOR OF JUVENILE BELDING GROUND SQUIRRELS Martin L. Morton^, Catherine S. Maxwell', and Charles E. Wade' -2 Abstract.— Juvenile Belding ground squirrels were studied in the Sierra Nevada. Females were more trappable, had smaller home ranges, and tended to enter hiboination earlier than males. The primary sex ratio was 1:1. Individuals first emerged from the natal burrow at tliree to four weeks of age and a body weight of 35 g. Body weight and linear dimensions increased thereafter until hibernation began. Maxinuim prehibernatory weight of 200 to 260 g was at- tained at about 12 weeks of age. Prehibernatory fattening began at about six weeks of age. Maximum lipid stores attained weighed about 80 percent of the lean, dry body compartment. Seasonal changes occurred in weight of white and brown fat depots, adrenal glands, spleen, heart, kidneys, liver, and testes. Annual variations in snowpack and emergence schedule caused the reproductive period, and thus phenology of juveniles, to vary by as much as three weeks. The last animals to innnerge were unusually small, being from late litters. Nonetheless, they may have had lipid stores sufficient for sui-viving hibernation. In natural history studies of ground squirrels (Spermophilus sp.), juvenile members of the population are often ignored or examined only casually. Yet these young, inexperienced animals are logical subjects if one is trying to uncover fundamental adaptations of species to environments, such as high altitude, that are only seasonal- ly accessible. Spermophiles survive in such environments through hibernation, and there is evidence that mechanisms favoring the success of juvenile hibernators, such as an accelerated ontogeny, are particularly important to the species as a whole (Mayer and Roche, 1954; Clark, 1970; Morton and Tung, 1971a). In this article we present the biology of juvenile Belding ground squirrels, Spermophilus beldingi beldingi, from a population residing at high altitude in the Sierra Nevada of central California. Parame- ters of growth, behavior, prehibernatory fattening, and effects of annual variations in snowpack on the schedule of juveniles are discussed. Methods Data were gathered during a five-year period (1969-1973) on 5. h. beldingi living on the eastern slope of the Sierra at the upper edge of the Mono Basin near Tioga Pass, Mono County, California. In this area there are subalpine meadows at elevations of 2926 to 3048 m (9600 to 10,000 ft) that are occui)ied by substantial numbers of S. h. beldingi. Squirrels were live-trapped in Tomahawk wire mesh traps baited with peanut butter. Those to be released were toe-clipped for later identification, and those kept for dissection were etherized. Linear moasuromonts were taken with calipers. Body weights were measured to the nearest 0.1 g on a Welch triple-beam pan balance. Fat pads, 'Deparlnicnt of Biology, Occidental College, Los Angeles, C^TJifornia 90041 'Department of Physical Education, University of California, Davis, California 95GI() 121 122 GREAT BASIN NATURALIST Vol. 34, No. 2 kidneys, heart, spleen, and liver were weighed to the nearest 0.01 g on an Ohaus analytical pan balance. Gonads and adrenals were weighed to the nearest 0.01 mg on a Federal Pacific torsion balance. Carcasses, including dissected portions, were kept frozen for later laboratory analysis of water and lipid content. Water content was determined by drying carcasses to a constant weight in a vacuum oven at 55C. They were then homogenized and extracted for lipid with petroleum ether in a soxhlet apparatus (Morton and Tung 1971b). Data on snow depth at Tioga Pass and predictions of snow- melt runoff from the Mono Basin were taken from Bulletins 120 and 129 of the California Department of Water Resources. Results and Discussion Seasonal Schedule of Events and Behavior. — Juvenile S. b. beldingi usually appeared above ground in mid-July. During the rest of the summer they gained in body size, deposited fat, and began entering hibernation in September at about three months of age. The last animals disappeared from the surface in the final ten days of September (1970) and the first ten days in October (1969, 1971, 1972, and 1973). Differences were noted in the behavior of males and females in terms of trappability, size of home range, and onset of hibernation. In 1971 a grid of 64 traps with 20 m spacing was set and centered on a burrow system containing a group of newly born litters. Fifty- five juveniles (30 females, 25 males) born in this burrow system were captured. During the course of the summer, females were captured an average of 9.53 times (range, 1-30), significantly (P<0.05) more often than males, 5.32 times (range, 1-21). This difference in number of recaptures may be due to sex-specific dif- ferences in home range and dispersal. Thirteen juveniles were always trapped within the outer rows of traps. Six were males trapped an average of 14.2 times, and seven were females trapped an average of 22.3 times. The mean home range of these animals, calcu- lated by the "halfway to the next trap" method (Hayne, 1949), was 6408 m- for males and 4434 m- for females. The home range of males was significantly larger (P<0.05). There are indications that males tended to remain active later in the season than did females. In 1973 juveniles were consistently trapped and released at burrow systems on a section of meadow roughly 300 m x 300 m. Each trapping session was conducted for about four hours with about 40 traps. Traps were checked once per hour. The daily catch increased from late July through August (Fig. 1). During that time, males and females were captured in about equal numbers during each trapping session, and 93 different individuals were handled: 47 males and 46 females. Beyond the first week in September, however, more males than females were captured during every trapping session except the final one, on 5 October. During September 10 individuals were trapped that had not been handled previously; nine were males, and only one was a June 1974 MORTON, ET AL: GROUND SQUIRRELS 123 18 - Pig. 1. Number of juvenile Spermophilus beldingi beldingi captured per trapping session during the 1973 season. female. The four males captured on 28 September and the two females ca{)tured on 5 October were the last squirrels trapped or observed in the 1973 season, and all six were kept as specimens. The larger home ranges of males, their lower rate of recapture, and their tendency to enter hibernation later than females are probably all in(hcators of greater dispersal of males from the natal area. Sex-specific emigration is common among young mammals and has been noted many times in ground squirrels (Quanstrom, 1971; Turner, 1972; Yeaton, 1972). One might predict that the hazards associated with dispersal would lead to greater mortality among juvenile males (Murray, 1967) and that an unbalanced sex ratio in favor of females would exist among those members of the population exceeding one year of age. These data raise a question concerning the sex ratio in young squirrels. During the five years of this study, juveniles were cap- tured or shot throughout the habitat at Tioga Pass known to harbor 124 GREAT BASIN NATURALIST Vol. 34, No. 2 S. b. beldingi. In all, 672 different individuals were handled: 336 males and 336 females. Clearly the sex ratio in juveniles was 1:1. Seasonal Changes in Body Size. — Growth of young squirrels was evident in terms of their linear dimensions and body weights, as the season progressed. Both body length and hind-foot length increased throughout the summer (Fig. 2). There was considerable variance in foot length, however, and it would seem to be a poor indicator of age. Body length was a more reliable indicator and has been used to show that maximum body size is not achieved in S. b. beldingi until the end of the second season of life or beyond (Morton and Tung, 1971a). In mid-September the oldest, largest juveniles began to enter hibernation, whereas the smallest, most immature individuals tended to remain active. This resulted in a decreased mean body size in late September samples. E|7 4.0 39 - 3.8 - 3.7 36 20 30 July 20 30 20 30 Aug Sept Fig. 2. Seasonal changes in mean body length and hind-foot length in juvenile Spermophilus beldingi beldingi at 10-day intervals. Data were pooled from five seasons, 1969-1973. Vertical lines indicate two S.E. Sample size shown in parentheses. June 1974 morton, et al: ground squirrels 125 When they first emerged from the natal burrow, juveniles weighetl about 35 g. For examj)le, on 15 July 1970, members of one litter of six that were probably venturing out for the first time were observed for 40 minutes, then hand-caj)tured and weighed. They paid no attention to an observer seated one m from them, and they cHd not go more than 1 m from the burrow entrance. Nevertheless, they sat erect, trilled, nibbled on grass, and when handled everted their anal glands. Coupling this information with that from captives of known age (Morton and Tung, 1971a), we estimate that these juveniles emerged at an age of three to four weeks. Their mean body weight was 36.4 g (range, 35.5-41.2 g), only one-half of what captives weigh at the same age (Morton and Tung, 1971a). Clearly, captives are precocious in comparison to wild juveniles, but the de- gree of precocity is difficult to determine. Turner (1972) has ob- sers'ed that S. b. oregonus juveniles attempt to nurse for the first few days after emerging from the natal burrow. Apparently the same behavior occurs in S. b. beldingi. Morhardt (1971) suggests that juvenile S. b. beldingi experience maternal care for about the first week following emergence. Mean body weight increased steadily each season of the study, from about 100 g in late July to about 215 g in mid-September, where it leveled off as the fatter individuals began hibernating and were no longer sampled by us (Fig. 3). Because of the inability of the live traps to retain small juveniles and because of our preoccupa- tion with other projects, no useful sample sizes were obtained until late July each year. As the season progressed thereafter, trapping success increased as the juveniles became larger and more active. During the final portion of September, trapping success decreased as the number of active squirrels decreased. There was considerable variation in mean body weight of the juvenile population from year to year. For example, 15 individuals weighed between 20 and 30 July 1973 were twice as heavy as 10 that were weighed during the same interval in 1969. There was a disparity of as much as three weeks in the dates at which specific mean body weights were reached. Obviously, young squirrels were not born within a certain very limited time year after year. These annual differences in growth schedules indicate that there is considerable variation in the timing of reproduction. Within a given season, variation occurred because females, even on the same meadow, emerged over a span of about a month, with adults tending to emerge ahead of yearlings. Furthermore, small yearling females often did not come into estrous until several weeks after emergence. Their offspring tended to be the ones that were active at the very end of the season. There was also variation in the schedule of reproduction that could be attributed to annual differences in snowpack. Snow depth at the study area was measured every year in February and April, and in May 1969, by the State of California (Fig. 4, upper panel). Snowmelt runoff for the watershed of our study area (Mono Basin) was also forecast by them each month from February through May (Fig. 4, lower). Both of these measurements show that 1969 was GREAT BASIN NATURALIST Vol. 34, No. 2 1970 1971 12 1972 18 1973 ]5 Total 65 42 109 95^ 246 17 50 102 52 131 352 Sept 39 51 21 58 21 0 87 78 43 26 30 9 103 Z2 67 313 252 140 30 Fig. 3. Seasonal changes in mean body weight of juvenile Spermophilus beldingi beldingi at 10-day intervals. Heavy line shows means when data were pooled for all five seasons. Sample sizes are below each class interval. the heaviest snow year during the study and that 1972 was the lightest. In fact, snowpack in 1969 was the heaviest recorded during the 44 years that the surveys have been taken, and snow was present on the study area for about a month longer than usual. In 1972 runoff was unusually early because of lack of precipitation and hot weather in March. These extremes in snow conditions strongly affected the timing of reproduction. As a result, juveniles reached prehibernatory weights several weeks earlier than usual in 1972, and they were larger early in the season than at compar- able dates of other years. It is interesting that, despite their late start, the 1969 juveniles attained prehibernatory weight levels at about the same calendar time as in other years (Fig. 3). The basis for their increased growth rate is not known, but the wet, green phase of the meadow environment persisted much later than usual in 1969. This could have provided a highly favorable nutritional situation for young squirrels. The data on mean body weights tend to obscure some ratlier important facets of growth and fattening as it actually occurred in individuals. Our records show that frequently captured, toe-clipped June 1974 MORTON, ET AL: CUOUND SQUIRRELS 127 E 300 o Q. o Q o c CO 200 00 0 ro E 250 ^ o c <4— o 200 *•> c 0) o ^ 0) 150 Q. <4- >4— o c 100 D q: •<-> s ^ r/ 'j8;aiuBip aSBJSAy rf 'j8}9uiBip isapi^ + + + I + n. + + + '^ (O CM t~~ ^ to ^ fO Csl O ^ !>0 ■* * 05 o - 00 o ro ro § (M CM ■^ ■^ ■^ ■^ "^ O) ^O t^ o t^ ro -^ c^ s § 05 in CM CM CM CM eg c5 ^ ^o 00 CO CM ro ■^ t^ CM CM -i- t^ »o xn o> o O OO Oi cq 'J- § CM CM vn § vr> t^ CSl t^ O) xn CM to o 00 CO ? l^ ^ 5^ 00 00 p o (O ■^' CT) vn to CTi 00 00 C a S On ^ e) ej e) !i: c u June 1974 anderson, weberg: vanclevea 155 Aspects of Vanclevea floral morphology are listed in Table 2. The heads are always discoid. Flower numbers (from single head counts) in our other taxa include: Acamptopappus schockleyi, 12 ray and 54 disc flowers; A. sphaerocephalus, 22 disc; Eastwoodia elegans, 50 disc; Grindelia decumbens, 12 ray and 59 disc; G. squar- rosa, 38 ray and 270 disc; G. squarrosa var. nuda, 222 disc; Haplo- pappus salicinus, 12 disc; and H. scopulorum, 9 disc in ours and 24 in Eastwood & Howell 7111 (rsa) . Differences in pappus are frequently considered characteristics for distinguishing genera in the Asteraceae. Among species under study, extremes are seen between Grindelia and Haplopappus. The former has two to eight deciduous, paleacous awns, whereas the latter has numerous persistent, capillary bristles. Pappus of Van- clevea is somewhat intermediate, though more like the Grindelia in having 15-18 tardily deciduous, paleacous awns. However, one Van- clevea flower from Holmgren & Hansen 3801 was found with a small gall (chalcid-fly induced) arising from the top of the achene beside the corolla. (The senior author has found similar galls oc- casionally in Chrysothamnus and Haplopappus.) The pappus ad- jacent to the gall is composed of numerous capillary bristles 6 mm long, whereas the pappus adjacent to the corolla is typical, consisting of eight paleacous awns each about 2.5 mm long. Perhaps too much significance is attached to pappus differences in the Asteraceae. Shinners (1949) certainly thought so. Data on floral anatomy are presented in Table 3; the format follows that in Anderson (1970a). Frequency classes are: -| — \-, abundant; -}-, frequent; -, rare; and 0, absent. Zones I and II are the proximal and distal areas of the achene, respectively; zones III - V are from the corolla; and VI and VII, from the style. Trichome types d, n, and g are duplex (the nonglandular twin-hairs character- istic of achenes), uniseriate nonglandular, and biseriate glandular, respectively. Corolla thickness (cell number) was determined at Table 2. Features of floral morphology in Vanclevea stylosa % i c y G ^ u ^ ^^ ^ sir" £ > > o '^ o ^ t? '^ ^ Collection « >B >S, E EO^c^ "^ B Anderson 3156 55.6 12.0 6.9 44.2 7.8 1.0 13.5 42.2 Anderson 3337 51.0 10.5 7.5 43.6 7.3 1.0 11.6 41.8 Eastwood 6 Howell 6660 46.5 12.4 6.9 34.0 8.2 0.9 13.6 43.5 Cutler 3/55 66.0 10.0 7.5 36.5 8.0 1.2 13.2 45.8 Holmgren & Hansen 3801 45.5 11.9 7.2 32.0 7.4 1.2 11.4 35.9 156 GREAT BASIN NATURALIST Vol. 34, No. 2 t/ 'qiSuaq Xouanbajjj 0) 1 adAx •n t/ 'i{:i3u37 H jtouanbaj^ >1 (0 Q 2 C u < •rH T3 4a > S 3 adAx IIA lA A AI III II I aSejaAy XO ;£) (O^O^OCOCOtO xn (O «o 00 o ooooCTioooa) -hc),-i o o o o T-i CM + + M U) oooooo ooo ooooo ooooo O O -^ CN ro +++++ ++++ o oo ooo •^ oo poo t^ + + +' ' + ''O H '^ CT3 0 bCO '^ 'W 'WO 'W o o +++ ooo o o +++ ooo o o °+++++ +++ +++ + ++ o o °+++ ' + +++ +++ + + o o O 1++0+ + o-|-o + + 0000_|_^ +++ + + + + + oooo_^o 1 1 o + + o o iocr> tN.00 t>. 00 00 xnoqoJcM^j^ xn^if) p O p p (M rq 05 05 00 xrioicMoivriod ^vovr! -^ 3 C O ft) a 3 o-o-g 3 C 1/5 3 -^ ^J^ ^~( «^ C C C |^« p Si vj Oj Vi v. C 1^ • •■^,^- "^ i^^ c c June 1974 anderson, weberg: vanclevea 157 three levels; those levels (A-C) and the seven zones studied for secretory canals are diagrammatically shown in Fig. 24 of Anderson (1970a). Data relating to ray flowers are not included in Table 3 because several of the taxa are eradiate. Ray flowers of Grindelia decumbens have short (90/(,), biseriate glandular trichomes, though none are found on the disc flowers. Trichomes in Acamptopappus are distinc- tive. Long isotropic, nonglandular trichomes (shag hairs) are found abundantly with the anisotropic duplex hairs on the achene walls. Duplex hairs were found only distally on Vanclevea achenes, and in Grindelia squarrosa the rare glandular hairs are restricted to the very top of the otherwise glabrous achenes. Ovarian vascular bundle number and the pattern of secretory canal distribution (present in achenes and corollas but absent in styles) are similar in the Haplopappus and Vanclevea. At least a few flowers of all samples from those two genera have additional corolla vasculature (midveins) ; the other taxa had only five veins in their disc corollas. Grindelia is set apart from the other taxa by its highly reduced vasculature in the achenes. Xeromorphy in Vanclevea flowers is evidenced in the thickness of its corollas and the massive sclerenchyma sheaths that surround the veins in the achenes. Taxonomy Although Vanclevea stylosa was first described as a Grindelia, that relationship apj)arently is not close. In addition to having fea- tures of habit, morphology, and anatomy presented here, the two groups are distinguishable chromosomally. In Grindelia, x — Q, whereas the single documented count for Vanclevea is /2 = 9 {Anderson 1976 in Solbrig et al., 1964). The remaining taxa studied here are also n — 9 or chromosomally unknown, as in H. salicinus. The purported relationships of Vanclevea to Acamptopappus and Chrysothamnus (Styermark, 1937) now do not appear close. The genus has many features in common with Haplopappus salicinus and //. scopulorum. It is with that section of Haplopappus (Hespero- doria) that Vanclevea is most closely allied. Although pappus in H . ciliatus (section Prionopsis) more than in any other Haplopappus resembles that of Vanclevea, H. ciliatus differs from Vanclevea in most other features of morphology, anatomy, and cytology. Haplo- pappus {sens, lat.) is badly in need of revision, but we believe that after such a study, Vanclevea will still be considered a separate genus in the Astereae. Vanclevea Greene, Pittonia 4:50. 1899. E. L. Greene dedicated this monotypic genus to a Mr. J. W. Van Cleve of Dayton, Ohio. Vanclevea stylosa (Eastw.) Greene, Pittonia 4:51. 1899. Basionym: Grindelia stylosa Eastw., Proc. Calif. Acad. Sci. II 6:293. 1896. 158 GREAT BASIN NATURALIST Vol. 34, No. 2 Type: [Epsom Creek] Barton's Range, San Juan Co., Utah, 13 July 1895, A. Eastwood 36. CAS (holotype) US!, photo - KSC! Slender branching shrubs (4) 6-7 (10) dm tall, older stems with shedding epidermis or dull white bark, often with axillary fascicles of small leaves, younger stems greenish white, glutinous, glabrous; leaves alternate, linear lanceolate, rigid, spreading to falcate and recurved, occasionally conduplicate, entire, (2.5) 3-3.5 (4.5) cm long and (2.5) 3-4 (5) mm wide; inflorescence cymose or a solitary head, discoid, viscous; involucres broadly turbinate, (9.5) 10-12 (12.5) mm tall, bracts graduated, subulate to broadly linear, acute to acuminate, often squarrose; flowers (31) 35-45 (48); corollas yellow, 7-8.5 mm long, lobes usually 1 mm long; styles well exerted, appendages longer than stigmatic lines; pappus of (12) 15-18 pa- leacous awns, stramineous, 2.5-3 mm long, often exceeding the in- volucre at anthesis; achenes narrowly cylindric, 4-5 mm long, nearly glabrous, n = 9. Variation within the species is not great, and no subspecific taxa are recognized. In most plants the bracts are squarrose and acu- minate to apiculate, but some are ascending and only acute. The degree to which the pappus is visible at anthesis also varies. Distribution: Sandy washes and sand hills; from Emery and Grand counties south through the Canyonlands Section of Utah to Coconino and Navajo counties in Arizona (Fig. 1). Phenology: Primarily blooming July through September, but occasionally later in the autumn and in March. Fig. 1. Flowering twig of Vanclevea slylosa and the known distribution of the species. June 1974 Anderson, wkbeug: vanclevea 159 ExsiccATAE: ARIZONA. Coconino Co.: Page, elev. 4300 ft, L. C. Anderson 1916 (ksc); 11 mi SSW Page, elev. 5100 ft, L. C. Anderson 1715 (uc, ny, rsa), 2619 (ksc); 9.3 mi SSW Page, A'. D. Atwood, S. L. Welsh, <& B. Wood 3359 (bry) ; 6 mi SW Page, elev. 4500 ft, /. Daney in 1971 (asc), R. H. Hevly & J. States in 1972 (asc); 1.5 mi below Lee's Ferry, H. C. Cutler 3/55 (mo, ny, us, SMu); Kaibito Plateau, elev. 6000 ft, L. D. Love in 1934 (ariz). Navaja Co.: Monument Valley, A. Eastwood & J. T. Howell 6660 (cAs, uc, us), J. T. Howell & G. True -14882 (cas), E. McClintock in 1963 (cAs), M. Bias 38 (cas); 7 mi SW Utah line toward Kayen- ta, elev. 5400 ft, J. T. Howell <& G. True 45018 (cas, ny); Tsegi Canyon, L. C. Whitehead in 1916 (ariz). UTAH. Emery Co.: Temple Wash, T25S, RUE NWy^. Sec 12, L. C. Anderson 3337 (ksc); 2 mi E Gilson Butte, Green River Desert, elev. 4800 ft. A. Baker 5 (us); Andy Moore North Spring [T26S, R15E], San Rafael Desert, elev. 5000 ft, Bryan & Read in 1938 (utc); 5 mi E Jeffrey Well, Green River Desert, elev. 4500 ft, W. P. Cottam 17761 (colo, ut); W Robber's Roost, L. A. Stoddart in 1943 (utc); Dry Lake Wash, ca. 15 mi S Green River, S. L. Welsh, N. D. Atwood, & G. Moore 10833a (bry, ny). Garfield Co.: 18 mi E Boulder vie. Circle Cliffs, B. Maquire in 1940 (ny); Baker's Ranch [T37S, R7E], B. Markham in 1940 (utc); Ticebo Mesa, T36S, R12E, J. C. Pederson 26 (bry). Grand Co.: S Double Arch, Arches Nat'l Monument, L. C. Anderson 66 (utc); Courthouse Towers, Arches Nat'l Monu- ment, S. L. Welsh, B. F. Harrison, & G. Moore 2268 (bry) ; S Turn- bow Cabin, Salt Wash, Arches Nat'l Monument, S. L. Welsh & G. Moore 2717 (bry, ny); Little Sand Flats, E Moab, elev. 4500 ft, K. Goodspeed in 1968 (utc). Kane Co.: 2.5 mi W Glen Canyon City, L. C. Anderson 3156 (ksc); 2 mi E Glen Canyon City, A'. D. Atwood 3095 (bry) ; 6 mi E Glen Canyon City, A^. D. Atwood & D. Kaneko (ut); Fiftymile spring S Escalante rT40S, R8E], J. R. Murdock 375 (bry); 55 mi E Kanab, S. L. Welsh 9418 (bry). San Juan Co.: Forbidding Canyon, Rainbow Bridge area, elev. 3600 ft, R. A. Darrow 2806 (ariz) ; Monument Valley, A. Eastwood & J. T. Howell 6675 (cas, k, ny, uc); S. Needle Rock, Monument Valley, A. H. Holmgren & S. Hansen 3801 (ny, uc, us, utc); Lower Beef Basin [T32S, R8E], NW Monticello, W. A. Shands 140 (us); N Mexican Water, elev. 4900 ft, B. Smith in 1966 (utc). Wayne Co.: 20 mi N Hanksville. R. Jensen in 1941 (utc); Burr Point [T30S, R13E], B. Markham in 1940 (utc); SSE Hanksville near county line, W. A. Shands 124 (us); 2 mi E Wadlerman Home, S Hanks- ville, W. A. Shands 140 (ut); San Rafael Desert, elev. 4500 ft, W. D. Stanton 1068 (ut); Barrier (Horseshoe) Canyon, 5. L. Welsh, N. D. Atwood. d^ G. Moore 10867 (bry, ny); 5 mi N Hanksville, 5. L. Welsh & G. Moore 3614 (bry). Literature Cited Anderson. L. C. 1963. Studies on Petradoria (Compositae) : anatomy, cytology, taxonomy. Trans. Kans. Acad. Sci. 66:632-684. 160 GREAT BASIN NATURALIST Vol. 34, No. 2 1964. Taxonomic notes on the Chrysothamnus viscidiflorus complex (Astereae, Compositae). Madrono 17:222-227. — . 1970a. Floral anatomy of Chrysothamnus (Astereae, Compositae). Sida 3:466-503. — . 1970b. Embryology of Chrysothamnus (Astereae, Compositae). Ma- drono 20:337-342. 1972. Studies on Bigelowia (Asteraceae). II. Xylary comparisons, woodiness, and paedomorphosis. J. Arnold Arb. 53:499-514. Carlquist, S. 1960. Wood anatomy of Astereae (Compositae). Trop. Woods 113:54-84. Dalbey, N. E. 1914. On the anatomy of Grindelia squarrosa. Kans. Univ. Sci. Bull. 9:31-41. Eastwood, A. 1896. Report on a collection of plants from San Juan County, in southern Utah. Proc. Calif. Acad. Sci. II 6:270-328. GiROUX, J., AND J. SusPLUGAS. 1935. £tude anatomique du Grindelia robusta. Bull. Sci. Pharm. 42:89-102. Greene, E. L. 1899. Neglected generic types. Pittonia 4:45-51. Howe, T. A. 1926. Development of embryo sac in Grindelia squarrosa. Bot. Gaz. 81:280-296. Martin, G. W. 1892. Development of the flower and embryo sac in Aster and Solidago. Bot. Gaz. 17:353-358. Shinners, L. H. 1949. Notes on Texas Compositae. III. Field and Lab. 17: 170-176. SoLBRiG, O. T., L. C. Anderson, D. W. Kyhos, P. H. Raven, and L. Rudenburg. 1964. Chromosome numbers in Compositae. V. Astereae II. Amer. J. Bot. 51:513-519. Steyermark, J. A. 1937. Studies in Grindelia, III. Ann. Mo. Bot. Gard. 24: 225-262. A REVISION OF THE ECHIDIOCARYA SECTION OF PLAGIOBOTHRYS (BORAGINACEAE) Larry C. Higgins* Abstr.\ct. — This revision discusses two closely related species of Plagio- bothrys (Boraginaceae) found in the southwestern United States and central Chile. One of the two species, P. collinus, is composed of five varieties that range from southern California and western Arizona to central Chile in South America. The following new combinations are proposed: P. collinus var. cali- fornicus, P. collinus var. gracilis, P. collinus var. ursinus, and P. collinus var. fulvescens. Detailed keys, synonymy, and descriptions are given for each entity. The section Echidiocarya, as here defined, includes those species of the genus that are annuals \^dth opposite or alternate lower leaves and that have the nutlet scar elevated on the end of a conspicuous stipe. The section Allocarya (to be treated in a later paper) differs mainly in that the nutlets lack the cylindrical stipe and the leaves are nearly ahvays oppositely arranged. The section Echidiocarya (Gray) Johnston is composed of two species and five varieties that are confined to the western half of the American continent, ranging from central California to central Ari- zona and southward to northern Sonora and northern Baja Californ- ia, Mexico, with an outl3'ing population in central Chile. The habi- tats of the various entities occur from sea level to steep mountain slopes and are characterized by scanty rainfall, warm temperatures, and high evaporation. In 1857 R. A. Phillipi described the first species in the section when he published Eritrichium coUinum. In 1908 K. Reiche trans- ferred the species to Cryptantha in his treatment of the Boraginaceae published serially in the Anales de la Universidad de Chile and in 1910 as part of his Flora de Chile. In 1895 Phillipi described E. inconspicuum, which falls into synonymy with E. collinum. Asa Gray (1876) proposed the genus Echidiocarya for Echidio- carya arizonica (Plagiohothrys pringlei Greene) . It was characterized by the long-stiped nutlets. Gray (1877) added E. calif ornica and (1885) E. ursina to the genus. Gray (1885), commenting on the very close relationship of his genus to Plagiohothrys, said, "The com- paratively recent discover}' of the preceding species [P. ursina'] of this section has made it clear that both of them should fall in Plagio- hothrys.'' He then transferred E. californica and ursina to Plagio- hothrys. leaving the original species by iteslf. Greene (1887) disposed of Echidiocarya, stating that it had "every aspect and every character of Plagiohothrys, except that there is a stipe between the scar, or point of attachment to the gynobase, and the body of the nutlet." Johnston (1923) placed Echidiocarya as one of his sections under Plagiohothrys. He also described two additional varieties of P. call- 'Department of Biology and Killgoie Research Center, West Texas Slate University, Canyon, Texas "OOf;. 161 162 GREAT BASIN NATURALIST Vol. 34, No. 2 fornicus. Since Johnston's treatment only Brand (1931) and Abrams (1951) have accepted Echidiocarya as a distinct genus. During my research — which led ultimately to a complete revision of Plagiobothrys- — a realignment of the members of this section be- came apparent because the South American P. collinus is conspecific with P. californicus of the southwestern United States and Mexico. Johnston (1927) was aware of the close relationship even though he never made any nomenclatural changes. He said, "the relationship between the Californian and Chilean plants is very clear and strong, so incontrovertible in fact, that some changes in the classification of the North American forms will probably be made." Those changes are the primary basis of this paper. Special thanks are extended to the curators of the following herbaria for their loan of material for this study: BRY, CAS, GH, ND, ND-G, NY, UC, US. I also wish to thank West Texas State University and the Killgore Research Center for their support both physical and monetary. Taxonomy Plagiobothrys Fisch. and Mey. Sect. Echidiocarya (Gray) Johnston, in Contr. Gray Herb. 68:65, 1923. Echidiocarya Gray, in Benth. and Hook. Genera PI. 2:854, 1876; Proc. Amer. Acad. Arts 11:89, 1876. Species Holotypus: (£. arizonica) P. pringlei Greene. Annual, usually diffusely branched, herb with the lowest leaves opposite or sometimes alternate; flowers in slender spikes, usually bracteate, calyx lobes linear-lanceolate, corolla white, the throad not crested; nutlets 4, incurved, conspicuously rugulose dorsally, con- spicuously keeled ventrally; scar on a slightly to prominently ele- vated stipe. 1. Stipe of nutlet elongate, approximating the body in length; nutlets commonly united in pairs; distribution in southern Arizona and northern Sonora 1. P. pringlei V. Stipe of nutlet short, 0.5 mm long or less, much shorter than the body of the nutlet 2. P. collinus 2. Corolla 4-7 mm broad; pubescence fine, appressed, usually somewhat silky; California, Arizona, and northern Mexi- co 2A. var. californicus 2'. Corolla 1-3 mm broad; pubescence usually spreading; coarse, setulose 3 3. Leaves narrowly linear, 2-2.5 mm broad 4 3'. Leaves oblanceolate, 3-5 mm broad 5 4. Pubescence fine, nutlets rounded-rugulose; California and Baja California, Mexico 2B. var. gracilis 4'. Pubescence coarse, nutlets sharply rugose; Chile. 2C. var. collinus 5. Racemes dense, hidden among leaves; San Bernardino Mountains, San Jacinto Mountains 2D. var. ursinus June 1974 higgins: plagiobothrys 163 5'. Racemes elongate, projecting from among the leaves and evident; southern California and Baja California, Mexico 2E. var. fulvescens 1 . Plagiobothrys pringlei Greene Echidiocarya arizonica Gray, in Benth. and Hook. Genera PI. 2:854, 1876; Proc. Amer. Acad. Arts 11:89, 1876. Not P. arizonicus (Gray) Greene; Plagio- bothrys pringlei Greene, in Pittonia 1:21, 1887. Mesa Verde, Arizona, Dr. Smart sn. Holotype GH! Stems several to many branched from near the base, prostrate or decumbent to nearly erect, slender, 1-4 dm long, spreading setose; leaves numerous below, less so above, narrowly oblanceolate, 2-4 cm long, 2-5 mm broad, rounded or obtuse to nearly acute at apex, approssed strigose or canescent to conspicuously setose; floral bracts lanceolate to narrowly oblong, 1-2 cm long; spikes elongating in age, bractless near the apices; calyx segments lance-linear, about 3 mm long in fruit, hirsute; corolla 2-3 mm broad, inconspicuous; nutlets 4, the lowermost commonly joined in pairs, the upper separate, ovate, the apex acute, 1.8-2 mm long, dorsal keel evident above fad- ing to rather distinct tuberculations below, dorsal surface rugulose with short ridges also conspicuously tuberculate; scar elevated on a j>rominent stipe at least 1 .3 mm long and usually as long as the nut- let, ventral keel evident; style shorter than the stipe of nutlet.— Representative collections: J. J. Thornber sn. (cas, ny); F. R. Fos- berg 10663 (cas, gh); V. L. Cory 3398 (gh); Pultz and Phillips 1572 (gh, ny); I. L. Wiggins 8689, 8190 (gh); L. C. Higgins 2822 (bry, wtsu). Distribution Common in Cochise, Maricopa, Pima, and Pinal counties of Ari- zona and in northern Sonora, Mexico. Growing on sandy desert flats and mesas. The most remarkable characteristic of Pringle's Plagiobothrys is the long-stiped nutlets, unique among the borages. This species is very closely related to P. collinus var. fulvescens, which it re- sembles in most details, except that the former has stalked nutlets and slightly longer calyx segments. 2a. Plagiobothrys collinus var. californicus (Gray) Higgins Plagiobothrys collinus (Ph.) Johnston var. californicus (Gray) Higgins comb, nov., based on Echidiocarya californica Gray, in Proc. Amer. Acad. Arts 12:164, 1877. Plagiobothrys californicus (Gray) Greene, in Bull. Calif. Acad. Sci. 2:407, 1887. Plagiobothrys coopcri Gray, in Proc. Amer. Acad. Arts 20:285, 1885. Allocaryastrum californicum (Gray) Brand, in Pflanzenr. 4"": 100, 1931. Southeastern California in San Bernardino County, 1876, Parry and Lemmon 278. Holotype GH! Leaves numerous below, oblanceolate, 1-3 cm long, 2-5 mm broad, rounded or obtuse at apex, thinly hirsute with ascending hairs or sometimes canescent with appressed pubescence; spikes slender, at maturity elongate and remotely flowered; calyx segments 3 mm long in fruit, linear-lanceolate, hirsute to hispid; corolla 4-6 mm 164 GREAT BASIN NATURALIST Vol. 34, No. 2 broad; nutlets about 1.5 mm long, scar with a short stipe near base. — Representative collections: J. T. Howell 3935 (gh, cas); T. S. Brandegee 1637 (gh); L. Abrams 3276 (gh); I. M. Johnston 1839 (gh). Distribution San Benito County, California, southward to northern Baja Cali- fornia, Mexico. P. collinus var. californicus is most easily recognized by its large corollas, 4-7 mm broad and tlie more canescent pubescence; even at this, it intergrades with varieties gracilis and fulvescens. 2b. Plagiobothrys collinus var. gracilis (Johnston) Higgins Plagiobothrys collinus (Ph.) Johnston var. gracilis (Johnston) Higgins comb, nov., based on Plagiobothrys californicus var. gracilis Johnston, in Contr. Gray Herb. 68:73, 1923. Allocaryastrum gracile Brand, in Pflanzenr. 4"-: 100, 1931. Echidiocarya calif ornica subsp. gracilis (Brand) Abrams, in 111. Fl. Pacific Stat. 3:571, 1951. San Diego, San Diego County, California, on the mesas. 10 April 1902. T. S. Brandegee 1658. Holotype GH! Stems slender, with spreading hispid pubescence; leaves linear- lanceolate, sparsely pubescent, with spreading setose hairs, 2-2.5 mm broad, acute or acutish; calyx segments linear, ca 2 mm long, very slender, hirsute; corolla 1.5-2 mm broad; nutlets 1-1.5 mm long, inconspicuously rugulose. — Representative collections: F. R. Fosberg 7681 (gas, gh); I. L. Wiggins 4462 (gh); F. F. Gander 4955 (gh) ; Raven & Mathias 12520 (gh). Distribution. San Diego County, California, southward to northern Baja Cali- fornia, Mexico, and on the islands of Santa Cruz, Santa Catalina, and San Clemen te. The variety gracilis is differentiated from the other varieties by the sparse spreading hirsute pubescence, narrower leaves, and small- er corollas. Its closest relatives are to be found in the varieties ful- vescens and collinus. Intermediates are not uncommon with variety fulvescens and are very difficult to separate from true gracilis. 2c. Plagiobothrys collinus var. collinus Eritrichium collinum Ph. Linnaea 29:17, 1857. Cryptantha collina Reiche, in Anales Univ. Chile 121:828, 1908; Fl. Chile 5:233, 1910. Plagiobothrys collinus (Ph.) Johnston, in Contr. Gray Herb. 78:81-82, 1927. Hills, Huanta, August 1836, Gay 1623. Holotype MS! Isotype fragment and photograph GH! Eritrichium inconspicum Ph. Anales Univ. Chile 90:534, 1895. Cryptantha inconspicua ReicJie, in Anales Uiuv. Chile 121:820, 1908; Fl. Chile 5:225, 1910. Coquimbo: La Serena, October 1878. Philippi s.n. Holotype MS! Fragment and photograph GH! Stems several, mostl}' erect, 3-15 cm long, hispidulous-villous; leaves linear to oblance-linear, obtusish, conspicuously hispidulous, 2-4 mm broad; calyx segments 1.5-2.5 mm long, fulvescent; corolla 1-1.5 mm broad, inconspicuous; nutlets 1.2-1.5 mm long, reticulately rugose with sharp ridges; style approximating nutlets. — Represen- tative collections: C. O. Skottsberg 1371, 723 (ny, gh); G. Montero June 1974 higgins: plagiobothrys 165 1895 (gh); E. Barros 6316, 6361 (gh); Worth & Morrison 16340 (gh). Distribution Chile, in the providence of Coquimbo. Apparently restricted to the area around Coquimbo. This variety is most closely related to the varieties gracilis and fulri'scens. It differs from the former in being coarser and in having broader obtusish more densely pubescent leaves and sharper and more prominently rugulose nutlets. From fulvescens it differs in hav- ing narrower leaves that are not so harshly pubescent, a more erect habit, and more prominently rugulose nutlets. 2d. Plagiobothrys collinus var. ur sinus (Gray) Higgins Plagiobothrys collinus (Ph.) Johnston var. ursinus (Gray) Higgins comb, nov., based ort Echidiocarya ursina Gray, in Proc. Amer. Acad. Arts 19:90, 1883. Plagiobothrys ursinus Gray, in Proc. Amer. Acad. Arts 20:285, 1885. Plagio- bothrys californicus var. ursinus (Gray) Johnston, in Contr. Gray Herb. 68:74, 1923. Echidiocarya californica var. ursina (Gray) Jepson, in Fl. Cahf. 3:370, 1943. California, San Bernardino Mountains, S. B. Parish 927. Lecto- type GH! Stems much branched, dense, compact, 2-8 cm long; spikes short; flowers concealed by the hispidulous leaves and bracts which are 1- 2.5 cm long; corolla 1-2 mm broad; nutlets weakly rugulose. — Rep- resentative collections: S. B. Parish 3247 (ny); P. A. Munz 5725 (gh); Munz & Johnston 4550 (gh); C. R. Orcutt 908 (gh). Distribution California, in the San Bernardino and the San Jacinto Moun- tains, growing in sandy to gravelly soils. The caespitose, compact habit of variety ursinus serves best to distinguish it from all the other varieties. It also grows at a higher, more montane elevation. 2e. Plagiobothrys collinus var. fulvescens (Johnston) Higgins Plagiobothrys collinus (Ph.) Johnston var. fulvescens (Johnston) Higgins comb, nov., based on Plagiobothrys californicus var. fulvescens Johnston, in Contr. Gray Herb. 68:74, 1923. Allocaryastrum ursinum var. fulvescens (Johnston) Brand, in Pflanzenr. 4'": 101, 1931. Echidiocarya californica subsp. fulves- cens (Brand) Abrams, in 111. Fl. Pacific Stat. 3:571-572, 1951. California, Santa Barbara. 1888, T. S. Brandegee s.n. Holotype GH! Plagiobothrys micranthus A. Nels., in Amer. J. Bot. 25:115, 1938. Arizona, Pres- cott, moist creek banks, 28 April 1925. A. Nelson 10232, Holotype RM! Iso- type GH! Stems slender, elongate, prostrate, hispid-pubescent, leaves ob- lancoolate, obtusish to acutish, hispidulous 3-5 mm broad; spikes \ery slender at maturity, remotely flowered; corolla about 2 mm broad. — Representative collections: Eastwood and Howell 3914, 4156 (c.AS, gh); F. R. Fosberg 10706 (cas, gh); J. T. Howell 31075 (gas); L. Abrams 3315 (ny); T. H. Kearney s.n. (cas); C. B. Hardham 3049,5(502, 10048 (cas). 166 GREAT BASIN NATURALIST Vol. 34, No. 2 Distribution Santa Barbara County, California, southward to northern Baja CaHfornia, Mexico, eastward to central Arizona. Also on the islands of Santa Rosa, Santa Catalina, and Anacapa. Most closely related to the varieties collinus and gracilis as dis- cussed under collinus. Variety fulvescens also introgresses consider- ably with varieties californicus and gracilis. Literature Cited Abrams, L. 1951. Illustrated flora of the Pacific states. 3. Stanford Univ., Calif.: Stanford Univ. Press. Bentham, G., and J. D. Hooker. 1876. Genera Plantarum. 1. London: Reeve and Co. Brand, A. 1931. Borraginaceae-Borraginoideae-Cryptantheae. Pflanzenr. IV. 252:100-101. Gray, A. 1876. Contributions in North American botany, IV. Miscellaneous botanical contributions. Proc. Amer. Acad. Arts 11:89. 1877. Contributions in Norlli American botany, XIV. Characters of some little-known or new genera of plants. Proc. Amer. Acad. Arts 12:163- 164. . 1883. Contributions in North American botany, I. Characters of new Compositae, with revisions of certain genera and critical notes. Proc. Amer. Acad. Arts 19:90. 1885. Contributions in North American botany. A revision of some Boragineous genera. Proc. Amer. Acad. Arts 20:285. Greene, E. L. 1887. Some west American Asperifoliae. Pittonia 1:9-21. Johnston, I. 1923. Studies in the Boraginaceae, IV. A synopsis and redefini- tion of Plagiobothrys. Contr. Gray Herb. 68:57-80. . 1927. Studies in the Boraginaceae, VI. A revision of the South Ameri- can Boraginoideae. Contr. Gray Herb. 78:81-82. Nelson, A. 1938. Rocky Mountain herbarium studies. Amer. J. Bot. 25:115. Philippi, R. 1857. Linnaea 29:17. . 1895. Plantas nuevas Chilenas. Anales Univ. Chile 90:534-535. Reiche, K. 1908. Estudios criticos sobre la flora de Chile. Anales Univ. Chile. 121:820-829. . 1910. Flora de Chile. 5:225-233. GREAT BASIN NATURALIS Si CoiiuTieiicmg with volume 35 (March 1975) The i< ^ "in \'ii;Mi!ist will be coml^inod witli the Brigham Youno iiv Bulletm, Bioi >eries under the title of 7 \ iuactust. The foniiat wni be changed from the prt-o.-nt v^.v./ m. .., Miigle column page to a 7x10 inch double colimm page. The journal will continue to be publi' ' .... ■• ,ges per No'iime will increase to . to 100 I pages (about 200 mdiiuscij, ic- . ^ .\ Publication and reprint cl ^, ^ I ho new format, but will not be significantly different from that indi- cated below. The subscription rate ^'^""^' ^'^ ^r^ry•n'.iM.H|<,.|^r...| (.,.|., ,.|.Mr'r,^-*r..|.HMi IIIMTIMI |.||| r| I'|'II'|M.|' 31 10 20 30 10 20 30| 10 20 30, 10 20 3p HAV JUNE ILLY AUGUST SEPTEMBER Fig. 5. Number of male P. boreus observed per female, between 31 May and 25 September 1972. 176 GREAT BASIN NATURALIST Vol. 34, No. 3 birth during the late suininer and early fall, and that females usually remain in their burrows or other shelters while carrying young on their dorsum. The tendency of vaejovids to remain in seclusion fol- lowing parturition would probably account for the paucity of females observed beyond 27 July. Environmental influence. — Despite the fact that a multitude of factors probably interact to regulate the surface activity of ter- restrial organisms, some interesting relationships between surface activity of P. boreus and temperature and rainfall were identified. Surface activity was not observed when the air temperature was below 10 C. As temperature increased, an increase in surface ac- tivity was noted (Fig. 6), except for the period between 29 June and 21 July, when there was no measurable precipitation (Fig. 6). The dry conditions during this period may have been responsible for the decrease in scorpion surface activity. Following the rainfall of 19 and 21 July, the original relationship of activity versus tem- perature was resumed. Any measurable precipitation was immediately followed by a decrease in surface activity (Fig. 6). This decreased activity was followed within three to five days by a period of increased activity. Surface activity was not observed during periods of rainfall. Reproduction. — No courtship activity was observed during this study. However, several gravid individuals were observed from 29 June to 27 July. During this period five gravid females were transported to the laboratory to determine fertility. Only two of the scorpions gave birth. A third died prior to parturition and was dis- sected. A female with only one pedipalp and a normal female each gave birth on 17 August 1972. The female with one pedipalp de- livered 13 young, 10 of which she consumed before their first molt. The normal female delivered 39 young, all of which lived beyond their first instar. The dissected female contained 51 nearly mature embryos. :^l 10 20 30 MAY JUNE Fig. 6. Relationship between scorpion activity, air temperature, and rainfall, between 31 May and 25 September 1972. Sept. 1974 TOURTLOTTE: SCORPIONS 177 Both females observed assumed a stilting position during parturi- tion. The young, completel}^ encased in a transparent birth mem- brane, passed through the genital aperture one by one. Within 10 to 20 minutes the young had freed themselves from the birth mem- brane and had chmbed up the mother's walking legs to occupy a nonrandom position on her dorsmn. The young scorpions first mohed 12 days following birth, and within the next week they began to move from the mother's dorsum to assume an independent existence. The birth process and subsequent development of the young were similar to those described by Williams (1969). The mean litter size for the three observations was 34 ± 11. Miscellaneous observ.\tions. — With few exceptions, all scor- pions observed by blacklight were stationary. Similar behavior was reported by Hadley and Williams (1968) for other vaejovids. Most scorpions were observed sitting directly on the ground surface; how- ever, several were seen on vegetation, as high as 50 cm above the ground. Although scorpions were not observ-ed in the act of capturing prey, feeding behavior was observed seven times during the study. Prev organisms included one grasshopper, two spiders, one scorpion (in a pitfall trap), two pentatomids, and one unidentified arthropod. One adult female was uncovered by excavation on 10 March 1973. She was in a cavity approximately 30 mm long, 15 mm wide, and 10 mm high, at a depth of 25 cm. The air temperature at the time of capture was -1 C, and the ground temperature was approxi- mately 8 C. No organisms were observed to prey upon P. horeus. although several scorpions were killed in pitfall traps by Jerusalem crickets, Stenopelmatus sp. Cannibalism was observed only once in the field, but again in a pitfall. Discussion and Conclusions P. boreus was active on the surface only during the nocturnal hours. Nocturnal surface activity was not equally distributed throughout the night but was more intense shortly after sunset, diminishing until dawn. Hadley and Williams (1968) reported similar activity patterns for V . confusis and P. mesaensis. Decreased surface activity throughout the night might be attributed to de- creasing air and soil temperatures during this period. Surface activity of P. boreus w^as not evidenced until 22 March 1972, after which time it seemed to be contingent upon a minimum air temperature of 10 C. Surface activity was observed throughout the summer but diminished in the fall and was not evidenced after 13 October 1972. Causes for the variation in surface activity were not determined, but a relationship between activity, temperature, and rainfall seemed apparent. Many other factors, such as photo- period, relative humidity, and soil moisture probably influence sur- face activity as well as temperature and rainfall, but these were not considered in this investigation. 178 GREAT BASIN NATURALIST Vol. 34, No. 3 Frequency of surface activity varied considerably among indi- viduals. This variability might be explained by examination of such biologically necessary activities as feeding, mating, and population- dispersal. Nocturnal surface activity could provide a means of satis- fying these necessities, after which the scorpion could remain in its burrow. Hadley and Williams (1968) suspected that nocturnal surface activity might be a means of maintaining water balance in a dry environment. The high frequency of surface activity for females would permit a higher probability of mating encounters with the more mobile males. Once she has located a suitable shelter, the female appears to limit her surface activity to this vicinity. Williams (1966) sug- gested that the differentiation of movement patterns in male and female A. phaeodactylus might relate to the mate-seeking activity of males. Sex ratios for P. boreus varied greatly, depending on time of year and sampling method. Allred (1973) assumed that males wander more extensively than females and are therefore more apt to fall into the pitfall traps. Because of the differentiation in mo- bility between males and females, data from pitfall studies were misleading in sex-ratio determinations. And because female vae- jovids become less active on the surface in late summer and early fall, even blacklight data from this period tended to underestimate the number of females in the population. Probably the best method to determine sex ratio would involve comparison of the total number of males observed to the total number of females observed over the entire study period. On the basis of the three litters examined, the variation in litter size for P. boreus is considerable. The smallest litter recorded (13) may have been a result of the female's handicapped condition, while the largest litter (51) may represent only the potential fecundity, since parturition had not yet occurred. The observed range in litter size is well within the range described by Baerg (1954) for several species of Scorpionida. Literature Cited Allred, D. M. 1973. Scorpions of the National Reactor Testing Station, Idaho. Great Basin Nat. 33:251-254. Anderson, R. C. 1972. Personal communication. Idaho State University, Pocatello, Idaho. Baerg, W. J. 1954. Regarding the biology of the common Jamaican scorpion. Ann. Entomol. Soc. Amer. 47:272-276. EwiNG, H. E. 1928. The scorpions of the western part of the United States, with notes on those occurring in northern Me.xico. Proc. U.S. Nat. Mus. 73:1-24. Gertsch. W. J., AND D. M. Allred. 1965. Scorpions of the Nevada test site. Brigham Young Univ. Sci. Bull., Biol. Ser. 6(4): 1-15. Gertsch, W. J., and M. Soleglad. 1966. The scorpions of the Vejovis boreus group (subgenus Paruroctonus) in North America (Scorpionida, Vejovidae). Amer. Mus. Novitates 1966 (2278): 1-54. Hadley, N. F., 1970. Micrometeorology and energy- exchange in two desert arthropods. Ecology 51:434-444. Sept. 1974 TOURTLOTTE: SCORPIONS 179 Hadley, N. F., and S. C. Williams. 1968. 'Surface activities of some North American scorpions in relation to feeding. Ecology 49:726-734. HoNETscHL.^GER, L. D. 1965. A new method for hunting scorpions. Turtox News 43:69. McDaniel, M. M. 1968. Notes on the biology of California scorpions. Entomol. News 79:278-284. Stahnke, H. L. 1966. Some aspects of scorpion behavior. Bull. So. Calif. Acad. Sci. 65:65-80. TouRTLOTTE, G. I. 1973. A preliminary investigation of the biology and ecology of the Northern Scorpion, Vaejovis boreus (Girard). Masters Thesis, Dept. Biol., Idaho State University. Pocatello, Idaho. 30 p. Williams, S. C. 1966. Burrowing activities of the scorpion Anuroctonus phaeodactylus (Wood) (Scorpionida: Vejovidae). Proc. Calif. Acad. Sci. 34:419-428. . 1967. Birth activities of some North American scorpions. Proc. Calif. Acad. Sci. 37:1-24. . 1968. Metliods of sampling scorpion populations. Proc. Calif. Acad. Sci. 36:221-230. . 1970. Coexistence of desert scorpions by differential habitat preference. Pan-Pacific Entomol. 46:254-267. 1972. Four new scorpion species belonging to the genus Paruroctonus (Scorpionida: Vaejovidae). Proc. Calif. Acad. Sci. 94:1-16. NOTES ON THREE VARIETIES OF ASTRAGALUS LENTIGINOSUS (LEGUMINOSAE) Carol S. Schoener^ Abstract. — A taxonomic review of the Astragalus lentiginosus complex in Utah indicates that the epithet var. albiflorus, antecedes and therefore should replace var. diphysus. The range of var. vitreus has been extended to include Kane and Garfield cos., Utah, and var. frernontii has now been identified from the state. In a taxonomic review of the Astragalus lentiginosus complex in Utah, it was found that the name, var. diphysus, is anteceded by the name A. diphysus var. albiflorus Gray. The varietal name diphysus is more commonly used due to its previous status as a species. It also appears to be a misnomer to refer to this bright purple flowered plant by the epithet albiflorus. Barneby (1964, Mem. N. Y. Bot. Gard. 13:941) discusses this problem. According to the rules of botanical nomenclature {International Code of Botani- cal Nomenclature, Article 11, 1972), the older name in the same rank has precedence. Because the oldest name at varietal rank is albiflorus, I propose the new combination, to wit: Astragalus lenti- ginosus var. albiflorus (Gray) Schoener stat. nov. based on Astraga- lus diphysus var. albiflorus Gray in Memoirs of the American Academy II, 4 (PI. Fendl.):34, 1894. Herbarium specimens of Astragalus lentiginosus var. frernontii were collected in the vicinity of Terrv's Ranch, at Beaver Dam Wash in Washington Co., Utah: Barnum 997 (BRY); Higgins 411 (BRY, NY) and 878 (BRY); Welsh 5265 (BRY, NY). This estabhshes a new state record: previously its distribution had been thought to be limited to the desert areas of southern Nevada and the northern and eastern Mohave Desert in California. The var. frernontii is quite distinctive with its small purple flowers (banner 9.4-11.5 [13.8] mm long, keel 6.3-7.9 mm long) and loose racemes which elongate as the flowers mature. Astragalus lentiginosus var. vitreus has now been identified in Utah from Kane and Garfield cos. Kane Co.: Four Mile Bench, Atwood 4064 (BRY) ; Nipple Springs Road, Atwood and Allen 2820 (BRY); north end of Brigham Plain, Atwood and Allen 02791 A (BRY); northeast slope of Smoky Mt., Cronquist 10022 (BRY, NY, RM, UTC). Garfield Co.: Star Spring, Henry Mts., Welsh 3942 (BRY, NY). Previous collection localities were in northern Mohave Co. and northwestern Coconino Co., Arizona, and in Washington Co., Utah. The variety can be characterized by its lax racemes (3.4) 4.0-9.0 (13.4) cm long and its inflated, papery-membranous, glabrous pods which become lustrous with maturity. Pods on two Kane Co. collections (Atwood and Allen, 02791 A; Cronquist, 10022) are atypically mottled. 'Department of Botany and Range Science, Brigham Young University, Prove, Utah 84602. 180 NEW SPECIES OF AMERICAN CORTHYLUS (COLEOPTERA: SCOLYTIDAE)' Stephen L. Wood- Abstr.\ct. — Tlio following species are described as new to science: Corthylus nolenae, C. spinosus (Mexico), C. diligens (Guatemala), C. minimus (Honduras), C. granulifer, C. reiusus, C. retusifer. C. oculatus, C. calamarius, C. cannularius, C. simplex, C. comosus, C. brunneus, C. calmicolens, C. zelus, C. trucis, C. py- gamaeus, C. sobrinus, C. subserratus (Costa Rica), C. pwnilus (Panama), C. donaticus. C. additus, C. cirritus (Venezuela), C. insignis, (Colombia), C. con- similis (Mexico and Guatemala), C. concisus (Mexico, Costa Rica, and Panama), C. sentus (Guatemala and Costa Rica), C. villifer, C. strigilis, C. irunculus, C. serratus (Costa Rica and Panama). A lectotype is designated for C. compressi- cornis (Fabricius). The 31 species of ambrosia beetles of the genus Corthylus de- scribed below as new to science were discovered during the prepara- tion of my monograph of the Scolytidae of North and Central America. They are named here in order to avoid nomenclatural confusion and to facilitate identification in the interim period before publication of the monograph. The species described here are from Mexico (2), Guatemala (1), Honduras (1), Costa Rica (15), Pana- ma (1), Colombia (1), Venezuela (3), Mexico and Guatemala (1), Mexico, Costa Rica, and Panama (1), Guatemala and Costa Rica (1), Costa Rica and Panama (4). Corthylus compressicornis (Fa- bricius) is redescribed, and a lectotype is designated. Corthylus granulifer, n. sp. This species is distinguished from retusus Wood by the smaller size, by much smaller tubercles on the elytral declivity, by the uni- formly convex declivity lateral from striae 1, and by the more slen- der body form. Male. — Length 2.0 mm, 2.5 times as long as wide; color dark bro\vn. Frons broadly convex, ascending slightly toward epistomal mar- gin; surface shining at epistoma, reticulate above, reticulate area with sparse, rather fine punctures; vestiture sparse, inconspicuous. Antennal club as in retusus. Pronotum 1.0 times as long as wide; anterior margin with median pair of serrations much larger, otherwise as in retusus female. Elytra 1.5 times as long as wide, 1.5 times as long as pronotum; similar to retusus, except elytral surface smoother, strial punctures near declivity almost in rows, declivital interstriae 2 not impressed, 3 not elevated, tubercles much smaller, declivital vestiture apparent- ly longer and slightly more abundant. 'The researcli that resulted in the discoverj- of these insects was sponsored bj' the National Science Foundation. -Department of Zoology, Brigham Young University, Provo, Utah 84602. Scolytidae contribution No. 53. 181 182 GREAT BASIN NATURALIST Vol. 34, No. 3 Type Locality. — Volcan Poas, Heredia, Costa Rica. Type Material. — The male holotype was taken at the type locality on 9-XI-63, at 2500 m, No. 261, from a tree branch, by me. One male paratype is from 15 km SE Cartago, 24-IX-63, 1800 m, No. 200, Myrica pubescens, S. L. Wood. The holotype and paratype are in my collection. Corthylus retusus, n. sp. This species is distinguished from granulifer Wood by characters summarized in the diagnosis of that species. Female. — Length 2.3 mm (paratypes 2.2-2.4 mm), 3.3 times as long as wide; color very dark brown. Frons deeply concave from eye to eye, from epistoma to vertex, lateral margin carinate from eye to lateral part of vertex; surface of lower half reticulate, upper half shining, slightly irregular, minutely, rather closely punctured, glabrous; a pair of widely spaced, slender, hornlike denticles near upper margin of concave area. Antenna! club asymmetrically subquadrate, 1.4 times as long as wide; anterior surface minutely pubescent; suture 1 indicated by an obscure sep- tum, 2 marked by an obscure groove; posterior face with a small tuft of hair extending slightly be3'ond apex of club. Pronotum 0.93 times as long as wide; sides almost straight and parallel on basal half, abruptly converging to rather broadly rounded anterior margin; anterior margin with about 10 low serrations (viewed from cephalic aspect) ; summit indefinite, slightly anterior to middle; anterior slope coarsely asperate; posterior areas subshin- ing, weakly reticulate, punctures minute, moderately abundant. Glabrous. Elytra 1.3 times as long as wide, 1.5 times as long as pronotum; sides very feebly arcuate and subparallel on basal two-thirds, rather broadly rounded behind; disc smooth, shining, with obscure, irregu- lar lines indicated, punctures small, confused, moderatel}'^ abundant. Declivity convex, steep; interstriae 1 uniformly abruptly elevated from near base to near apex, about as high as wide on middle half, its summit smooth, shining, with sparse punctures; interstriae 2 moderately, broadly impressed, 3 convex, slightly elevated, 2-5 each bearing a row of conspicuous, closely spaced, pointed tubercles. Vestiture confined to declivity, of fine, long, interstrial hair. Protibia with posterior face subrugulose, unarmed. Male. — Similar to female, except frons and anterior margin of pronotum as in male granulifer; antennal club smaller, less strongly asymmetrical, tuft of hair absent. Type Locality. — Tapanti, Cartago, Costa Rica. Type Material. — The female holotype and one paratype were taken at the type locality on 26-XI-63, at 1300 m, No. 265, from Phoebe mexicana branches. The male allotype and one paratype are from the same locality, taken 2-VII-63, No. 9, Miconia. Eight para- Sept. 1974 WOOD: American scolytidae 183 types are from 15 km SE Cartago, Cartago, Costa Rica, 24-IX-63, 1800 m. No. 198, Solanuni torviim; one para type bears this same data except it is from No. 195, tree seedling. All specimens were taken by me. The holotj'^pe, allotype, and paratypes are in my collection. Corthylus retusifer, n. sp. This species is distinguished from retusus Wood by the characters of the antennal club and elytral declivity as cited below. Female. — Length 2.0 mm (paratypes 1.7-2.0 mm), 2.3 times as long as wide; color very dark brown (holotype not fully darkened). Frons as in retusus; antennal club more nearly subcircular, su- tures 1 and 2 entire!}- obsolete except for a very small piece of septum of 1 on lateral margin. Pronotum and elytra as in retusus, except declivital interstriae 2 impressed, very strongly narrowed, almost obsolete, unarmed, 3 weakly convex, 3-5 armed by small granules about as in granulifer Wood. Male. — Similar to female, except frons, antenna, and anterior margin of pronotum as in male granulifer. Type Locality. — Tapanti, Cartago, Costa Rica. Type Material. — The female holotype and two paratypes were taken on 26-XI-63, at 1300 m. No. 265, Phoebe mexicana. The male allotype and two paratypes are from the same localit}^ 24-X-63, No. 266, Cecropia branch. One paratype bears data identical to the type except it is No. 271, Werklia insignis; two paratypes are from the same locality, one taken 2-VII-63, No. 7, wood}- vine, and one 17-IX-63, No. 176, tree branch. All were taken by me. The holotype, allotype, and paratypes are in my collection. Corthylus villi fer, n. sp. This unique species is distinguished from all others in the genus by the small, slender form, by the abundant, minute setae on the posterior half of the elytra, by the female antennal club and frons, and by other characters. Female. — Length 1.5 mm (paratypes 1.4-1.7 mm), 2.5 times as long as wide; color brown. Frons rather shallowly concave from eye to eye from epistoma to upper level of eyes, a median, shining callus on epistoma; surface shining, punctures coarse, close, deep; vestiture of fine, long, uni- formly distributed hair. Antennal scape rather strongly flattened; club large, asynii7ietrical, 1.3 times as long as wide, sutures obsolete except 1 at mesal margin, uniformly, finely pubescent. Pronotum 1.05 times as long as wide; sides almost straight and parallel on basal half, rather broadly rounded in front; anterior margin unarmed; summit indefinite, slightly in front of middle; rather finely asperate on anterior slope; posterior areas reticulate, 184 GREAT BASIN NATURALIST Vol. 34, No. 3 punctures small, obscure, moderately abundant; lateral margins rounded, not marked by a raised line. Subglabrous. Elytra 1.6 times as long as wide, 1.6 times as long as pronotum; sides straight and parallel on more than basal two-thirds, broadly rounded behind; disc smooth and shining at base, becoming strongly reticulate toward declivity, punctures sparse, small, confused. De- clivity steep, shallowly, subconcavely impressed on median half, lateral and basal margins broadly rounded; surface minutely, closely punctured; impressed area extending from suture to position of inter- striae 3. Vestiture largely confined to posterior fourth, of dense, minute hair, a few longer setae on marginal areas. Protibiae subinflated on posterior face, a few very minute gran- ules indicated. Male. — Similar to female, except frons convex, reticulate, punc- tures sparse, obscure, subglabrous; anterior margin armed by one median pair of large, slender teeth. Type Locality. — Fifteen km SE Cartago, Cartago, Costa Rica. Type Material. — The female holotype, male allotype, and 37 paratypes were taken at the type locality on 24-IX-63, 1800 m, Nos. 187, 199 (type), and 248, Siparuna nicaraguensis. Fourteen para- types bear identical data except No. 201, woody vine. Three para- types are from Tapanti, Cartago, Costa Rica, 2-VII-63, 1300 m, No. 177, wood)^ vine. Twenty-six paratypes are from Cerro Punta (labeled Volcan Chiriqui), Chiriqui, Panama, 11-1-64, 1800 m. No. 385, woody vine. All were taken by me. The holotype, allotype, and paratypes are in my collection. Corthylus oculatus Wood This species is distinguished from villifer Wood by the smaller size, by the ^'ery large eyes, by the narrower, less strongly impressed female frons, and by the strongly elevated declivital interstriae 1. Female. — Length 1.2 mm (paratypes 1.0-1.2 mm), 2.3 times as long as wide; color brown. Frons narrow, shallowly concave from e3"e to eye, from epistoma to slightly below upper level of eyes; surface smooth and shining on upper half, reticulate below, punctures sparse, fine; vestiture rather sparse, moderately long. Eye very large, coarsely faceted. Antennal club similar to villifer but slightly smaller. Pronotum and elytra about as in villifer^ except elytral disc strongly reticulate to base, declivity more nearly convex, declivital interstriae 1 strongly, acutely elevated from base to near apex, punctures not evident, a few minute granules on interstriae 3; vesti- ture confined to declivity, consisting of very short, moderately abun- dant hair, a few long, slender bristles on declivital interstriae 1, 3, and lateral areas. Male. — Similar to female, except frons convex, reticulate, punc- Sept. 1974 WOOD: American scolytidae 185 tures obscure; anterior margin of j^roiiotuin amied by a median pair of slender teeth. Type Locality. — Pandora, Limon, Costa Rica. Type Material. — The female holotype, male allotype, and eight paratypes were taken at the type locality on 23-VIII-63, at 50 m, No. 142, in a tree branch, b}' me. The holotype, allotype, and paratypes are in my collection. Corthylus calamarius, n. sp. This species is distinguished from cannularius Wood by the distinct strial punctures that are in rows, by the larger, regularly placed interstrial granules on the declivity, and by the more coarsely punctured female frons. Female. — Length 2.9 mm (paratypes 2.5-2.9 mm), 2.6 times as long as wide; color yellowish brown, with anterior third of j)ronotum and elytral declivity dark brown to black. Frons deeply concave from eye to eye, from epistoma to vertex; surface reticulate, punctures rather close, moderately coarse; vesti- ture of moderatel}' abundant, fine, rather long hair, uniformly distributed except absent on a small median area at epistoma. An- tennal club subreniform, 1.2 times as long as wide; suture 1 repre- sented by septum on lateral third, 2 and 3 represented by arcuate grooves (secondary strengthening features, not true sutures); a small tuft of hair on posterior face not reaching apex. Pronotum 1.02 times as long as wide; widest near base, sides weakly arcuate and converging ver}' slightl}' on basal half, rather narrowly rounded in front; anterior margin serrate; summit in- definite, slightly in front of middle; anterior slope moderately steep, coarsely asperate; posterior areas finely reticulate, punctures very small, rather sparse; lateral margin rounded. Glabrous. Elytra 1.6 times as long as wide, 1.6 times as long as pronotum; sides straight and parallel on basal two-thirds, rather broadly round- ed behind; disc reticulate, punctures near base minute, obscure, con- fused, those on posterior third distinct and in definite strial rows. Declivity convex, steep; striae 1 and 2 with punctures minute; inter- striae 2 broadly, very weakly impressed, 1-4 each with a row of small, regularly spaced granules. Vestiture confined to declivity, consisting of interstrial rows of moderately long, slender bristles. Male. — Similar to female, except frons convex, reticulate sparsely punctured, subglabrous, antennal club smaller, less irregular in shape; median pair of serrations on anterior margin of pronotum much larger than others. Type Locality. — Volcan Poas, Heredia, Costa Rica. Type Material. — The female holotype, male allotype, and nine paratypes were taken at the type locality on 19-XT63, near 2000 m elevation, from recently cut native bamboo, by me. The holotype, allotype, and paratypes are in my collection. 186 GREAT BASIN NATURALIST Vol. 34, No. 3 Corthylus cannularius, n. sp. This species is distinguished from calamarius Wood by the con- fused, obscure to obsolete punctures near the elytral declivity, by the very minute to obsolete, irregularly placed interstrial granules on the declivity, and by the more finel}" punctured female frons. Female. — Length 2.5 mm (paratypes 2.5-2.8 mm), 2.5 times as long as wide; color yellowish brown, anterior third of pronotum and elytral declivity darker. Frons as in calamarius, except surface almost smooth, punctures minute. Pronotum as in calamarius, except anterior margin more coarsely serrate (somewhat variable). Elytra as in calamarius, ex- cept punctures near declivity confused or obsolete, strial punctures on declivity obsolete, interstrial granules on declivity much smaller and more widely, less regularly spaced, declivital vestiture finer and less abundant. Male. — Similar to female, except frons convex, reticulate, sparse- ly punctured, subglabrous; antennal club smaller, less irregular in shape; anterior margin of pronotum armed by only two coarse teeth. Type Locality. — Cerro de la Muerte, San Jose, Costa Rica. Type Material. — The female holotype, male allotype, and 38 paratypes were taken at the type locality on 6-VIII-63, at 3200 m. No. 46, in native bamboo, by me. The holotype, allotype, and paratypes are in my collection. Corthylus comosus, n. sp. This species is distinguished from the very closely related villosus Eggers by the smaller pale area on the lower fifth of the female frons, by the smaller punctures on the elytral disc, by the more closely punctured elytral declivity, by the much more abundant declivital vestiture, and by the larger size {villosus 2.0-2.5 mm). Female. — Length 2.6 mm (paratypes 2.5-2.8 mm), 2.1 times as long as wide; color dark brown. Frons deeply, uniformly concave from eye to eye, from epistoina, to vertex; almost half of concave area above eyes; surface reticulate, finely, closely, uniformly punctured, lower one-sixth yellow and with punctures finer and much closer; vestiture of rather abundant, uniformly distributed, short, fine hair, slightl}^ longer on upper margin. Antennal club large, reniforni; tuft of hair on posterior face ending before apex of club. Pronotum 0.90 times as long as wide; widest near bases, sides weakly arcuate and subparallel on basal third, broadly rounded in front; anterior margin armed by about 14 low serrations; summit indefinite, anterior slope moderately steep; asperities commence slightly behind middle and continue^ to anterior margin; posterior Sept. 1974 WOOD: AMERICAN SCOLYTIDAE 187 area reticulate, punctures minute. Minute, hairlike pubescence moderately abundant, but usually abraded. Elytra 1.2 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal half, broadly rounded behind; disc smooth, a few ver}' obscure irregular lines evident, punctures very fine, moderately abundant, confused. Declivity oc- cui)ying jDosterior third, steep, evenly, broadly convex; surface smooth, shining, punctures very fine, close, confused, much closer than in villosus. Vestiture sparse on disc, abundant on entire de- clivity, of fine, moderately long hair, more abundant than in villosus. Male. — Similar to female, except frons convex, reticulate, rather finely punctured, vestiture sparse; anterior margin of pronotum armed by two median teeth. Type Locality. — Tapanti, Cartago, Costa Rica. Type Material. — I'he female holotype, male allotype, and 30 para types were taken on 26-XI-63, at 1300 m, No. 264, from Piper, by me. The holotype, allotype, and para types are in my collection. Corthylus simplex, n. sp. This species is distinguished from rubricollis Blandford by the characters of the elytral declivity and of the female frons as indi- cated below. Female. — Length 2.9 mm (paratypes 2.7-3.0 mm), 2.4 times as long as wide; color black. Frons as in rubricollis, except punctures larger, vestiture coarser, slightly less abundant, slightly longer, broad median line on upper third impunctate and glabrous. Antennal club 1.3 times as long as wide; sutures 1 and 2 slightly procurved, parallel, clearly marked; tuft of hair on posterior face extending about half club length be- yond apex. Pronotum and elytra as in rubricollis except upper two thirds of declivity more gradual, declivital interstriae 2 less strongly im- pressed, strial punctures much smaller, declivital bristles (when present) about twice as long. Male. — Similar to female, except frons convex, reticulate, punc- tures fine, sparse, subglabrous. Type Locality. — Cerro de la Muerte, San Jose, Costa Rica. Type Material. — The female holotype and male allotype were taken at the type locahty on 6-VIII-63,' at 3100 m. No. Ill, from a tree seedling. Six paratypes are from Volcan Poas, Heredia, Costa Rica, three on 14-VII-63, 2500 m. No. 46, Podocarpus oleifolius, and three 19-XI-63, 2500 m. No. 262, tree branch. All were taken by me. The holotype, allotype, and paratypes are in my collection. 188 GREAT BASIN NATURALIST Vol. 34, No. 3 Corthylus consimilis, n. sp. This species is distinguished from panamensis Blandford by the larger elytral punctures, by the feebly impressed interstriae 2, and by the larger elevated callus on the female vertex. Female. — Length 2.8 mm (paratypes 2.5-2.8 mm), 2.2 times as long as wide; color -very dark brown. As in panamensis, except frontal callus larger, extending from vertex to slightly below upper level of eyes; elytral punctures dis- tinctly larger, particularly on declivity; declivital interstriae 2 feebly impressed; declivital tubercles slightly larger and more widely distributed. Male. — Similar to female, except differing as in panamensis. Type Locality. — Nine km NE Teziutlan, Puebla, Mexico. Type Material. — The female holotype, male allotype, and 22 paratypes were taken at the type locality on 2-VII-67, 1600 m, Nos. 136, 140 (type), from tree boles, or an unnumbered sample from Alnus. One paratype is from the same locality, taken 27-VI-53. Five paratypes are from Volcan Pacaya, Esquintla, Guatemala, l-VI-64, 1300 m. No. 668, tree branch. Four paratypes are from Volcan de Agua, Guatemala, 19-V-64, 1000 m, No. 615, Acacia. All were taken by me. The holotype, allotype, and paratypes are in my collection. Corthylus brunneus, n. sp. This species is distinguished from calmicolens Wood by the smaller size, by the more slender body form, by the much more coarsely serrate anterior margin of the pronotum, and by the setal characters indicated below. Female. — Length 3.2 mm (paratypes 3.0-3.3 mm), 2.4 times as long as wide; color dark brown, discal area of elytra lighter. Frons as in punctatissimus (Zimmermann), except vestiture more abundant and slightly longer on margins of lower half. An- tennal club very large, strongly subreniform, suture 1 weakly, 2 moderately procurved; tuft of hair on posterior face large, its tip reaching apex. Pronotum as in punctatissimus, except anterior margin very coarsely serrate, median serrations larger. Elytra as in punctatissimus. except punctures slightly smaller, declivity not as steep, more broadly convex, punctures much less numerous, very fine, confused except on striae 1, about four minute granules on interstriae 1; ventrolateral margin of declivity with a low, short, subacute carina extending from costal margin toward interstriae 8. Vestiture fine, hairlike, sparse, not confined to declivity, mostly confined to odd-numbered interstriae except on sides, rather sparse on interstriae 8 and 9 and not longer than elsewhere. Sept. 1974 WOOD: American scolytidae 189 Male. — Similar to female, except frons convex, subshining, sparsely, coarsely ])uiictured; median j)air of serrations on anterior margin of pronotum larger. Type Locality. — Volcan Barba, Heredia, Costa Rica. Type Material. — The female holotype, male allotype, and 13 paratypes \vere taken on 19-XI-63, at 2500 m. No. 258, from native bamboo, by me. The holotype, allotype, and paratypes are in my collection. Corthylus ralmicolens. n. sp. This species is distinguished from hrunneus Wood by the larger size, by the stouter body form, b}' the much more finely serrate anterior margin of the pronotum, and by the setal characters indi- cated below. Female. — Length 4.0 mm (paratypes 3.7-4.0 mm), 2.3 times as long as wide; as in hrunneus.. except anterior margin of pronotum finely serrate, elytral declivity more broadly convex, and vestiture along sides of elytra much more abundant and longer. Male. — Similar to female, except differing as in male hrunneus. Type Locality. — South slope Volcan Barba, Heredia, Costa Rica. Type Material. — The female holotype, male allotype, and 11 paratypes were taken at the type locality at Finca Vota Steinworth, on 8-II-65, from native bamboo, by J. B. Karren. One paratype is from Volcan Poas, 19-XI-63, 2500 m, No. 258, native bamboo, by me. The holotype, allotype, and paratypes are in my collection. Corthylus strigilis, n. sp. This species is distinguished from comatus Blandford by the more deeply impressed declivital interstriae 2, by the much less closely, more coarsely punctured female frons, by the very small tuft of hair on the posterior face of the female antennal club, and by the finely serrate apical margin of the female antennal club. Female. — Length 2.7 mm (paratypes 2.7-3.1 mm), 2.3 times as long as wide; color dark brown. Frons as in comatus. except less deeply concave, sparsely, less uniformly punctured, an indistinct median callus on upper third, vestiture much more sparse, very fine, absent on callus. Antennal club 2.8 times as long as wide, large, rather strongly asymmetrical; sutures 1 and 2 indicated by grooves, weakl}- procurved; tuft of hair on posterior face very small, ending remote from apex. Pronotum and elytral disc as in comatus. Elytral declivity as in comatus, except interstriae 2 more strongly impressed, 1 more strongly elevated, half as high as wide, lateral convexities more nar- rowly rounded. 190 GREAT BASIN NATURALIST Vol. 34, No. 3 Male. — Similar to female, except differing as in comatus. Type Locality. — Sixteen km SE Cartago, Cartago, Costa Rica. Type Material. — The female holotype, male allotype, and three paratypes were taken at the type locality on 24-IX-63, at 1800 m. No. 205, in Myrica pubescens, by me; five paratypes bear identical data except No. 204 from Miconia globuliflora. Thirty-nine para- t3'pes are from Tapanti, Cartago, Costa Rica, 2-VII-63, No. 9, Miconia, 17-IX-63, No. 176, tree limb, 24-X-63, No. 241, Phoebe mexicana, 26-XI-63, No. 265 on Phoebe mexicana and 270 on Werklia insignis. Two paratypes are from Escasu, San Jose, Costa Rica, 2-X-63, 1300 m. No. 217, Ficus. Fifteen paratypes are from Cerro Punta (labeled Volcan Chiriqui), Chiriqui, Panama, 11-1-64. 1800 m. No. 393 in a tree branch. No. 407 in a tree seedling. All were taken by me. The holotype, allotype, and paratypes are in my collection. Corthylus diligens, n. sp. This species is distinguished from comatus Blandford by the much more deeply impressed declivital interstriae 2 and by char- acters of the female frons and antennal club as indicated below. Female. — Length 2.5 mm (paratypes 2.5-2.6 mm), 2.4 times as long as wide; color brown. Frons as in comatus, except concave area more broadly oval, 1.2 times as long as wide (1.4 times in comatus); tuft of hair on pos- terior face of antennal club with tip ending remote from apex. Pro- notum as in comatus., except anterior margin more coarsely serrate; punctures on disc minutely, obscurely granulate. Elytra as in co- matus, except declivity more strongly impressed on interstriae 2, su- ture not as high, lateral convexities higher and more narrowly rounded. Male. — Similar to female, except sexual differences as in comatus. Type Locality. — Volcan de Agua, Esquintla, Guatemala. Type Material. — The female holotype and five paratypes were taken at the type locality on 19-V-64, 1000 m, Nos. 593 in Clusia, 600 (type) in Alnus. The male allotype and one paratype are from Palin, Esquintla, Guatemala, 19-V-64, 300 m, No. 687, Ficus. Two paratypes are from Lago Amatitlan, lO-VI-64, 700 m, tree branch. All were taken by me. The holotype, allotype, and paratypes are in my collection. Corthylus zelus, n. sp. This species is distinguished from diligens Wood b}' the larger size and by characters of the female frons and antennal club as indi- cated below. Sept. 1974 WOOD: American scolytidae 191 Female. — Length 3.5 mm (paratypes 3.4-3.5 mirai), 2.5 times as long as wide; color very dark brown. Frons similar to diligens, except concave area less strongly im- pressed, upper margin not clearly delimited; frontal vestiture longer. Tuft of hair on antennal club with its tip almost attaining apex. Anterior margin of pronotum less coarsely serrate. Elytral declivity with punctures on striae 1 in a row, shallow, of moderate size; lateral j)unctures nuich smaller. Male.— Similar to female, except sexual differences as in comatus. Type Locality. — Cerro de la Muerte, San Jose, Costa Rica. Type Material. — The female holotype, male allotype, and one male paratype were taken at the type locality on 6-VIII-63, at 2500 m, No. 114, in Brunellia costaricensis. by me. The holotype, allotype, and paratype are in my collection. Corthylus trunculus, n. sp. This unique species is distinguished from all other representatives of the genus by the small size, by the truncate elytral declivity that is armed at its base by a pair of denticles on interstriae 3, by the con- vex female frons, and by the strongly procurved sutures on the female antennal club. Female. — Length 1.4 mm (paratypes 1.3-1.4 mm), 2.2 times as long as wide; color brown. Frons weakly convex, except somewhat flattened on lower fifth; surface subreticulate, punctures small, obscure, moderately abundant; vestiture of uniformly distributed moderately long hair from epi- stoma almost to vertex. Antennal club subtriangular, more nearly symmetrical than normal for this genus, 1.1 times as long as wide; sutures 1 and 2 strongly procurved, 1 apparently partly septate but not showing on posterior face; posterior face with tip of small tuft of hair reaching apex. Pronotum 1.1 times as long as wide; basically as in comatus Bland ford, except sculpture much finer, posterior areas strongly reticulate. Elytra 1.2 times as long as wide; 1.1 times as long as pronotum; sides almost straight on basal five-sixths, very slightly narrower behind, abruptly, very broadly rounded behind; disc almost smooth, subshining. points, lines, and punctures small, confused, somewhat obscure. Declivity truncate, very steep, margin from base to apex abruptly rounded, slightly elevated (not acutely margined); basal margin armed by a pair of small pointed spines at interstriae 3; sutural interstriae uniformly, subacutely, rather weakly elevated; surface strongly reticulate, punctures abundant, confused, shallow, very obscure. Vestiture sparse, short, confined to sides and margins of declivity. Posterior margin of protibia subinflated and armed by four to six confused, minute denticles. 192 GREAT BASIN NATURALIST Vol. 34, No. 3 Male. — Similar to female, except frons more strongly convex, subglabrous; antennal club normal, sutures weakly procurved, tuft of hair absent; anterior margin of pronotum armed by two or more coarse teeth. Type Locality. — Thirteen km or 8 miles S El Hato del Volcan, Chiriqui, Panama. Type Material. — The female holotype, male allotype, and six para types were taken b}' me at the type locality on 7-1-64, 1000 m. No. 371, from a tree seedling, by me. One paratype is from Bever- ley, Limon, Costa Rica, 26-VIII-63, 7 m. No. 154, woody vine, by me. The holotype, allotype, and paratypes are in my collection. Corthylus minimus, n. sp. This species is distinguished from minutissimus Schedl by the characters of the head and elytral declivity as indicated below. Female. — Length 1.3 mm (paratypes 1.3-1.4 mm), 2.3 times as long as wide; color brown. Frons very narrow, very shallowly concave on central half, im- pressed area not attaining upper level of eyes; surface shining, ob- scurely reticulate, a few punctures in marginal areas; subglabrous. Eyes very large, coarsely faceted. Antennal club 1.5 times as long as wide, moderately asymmetrical; sutures almost straight, 1 partly septate, 2 very obscure; posterior face without a tuft of hair. Pronotum and elytra essentially as in minutissim.us, except elytral declivity with interstriae 2 narrowed, moderately impressed, 1 carinate, more strongly elevated, 3 distinctly elevated and armed by about four minute tubercles, 2 without setae. Male.- — Similar to female, except frons convex, rugose-reticulate, punctures more uniformly distributed; eyes of normal size; antennal club smaller, less elongate; anterior margin of pronotum armed by a pair of coarse, median teeth. Type Locality.— La Ceiba, Atlantida, Honduras. Type Material. — The female holotype, male allotype, and 15 paratypes were taken at the tvpe locality from 20-V to 26-Vin-49 (type 1-VL49), at light, by E. C. Becker. The holotype, allotype, and paratypes are in my collection. Corthylus pumiius, n. sp. This species is distinguished from minutissimus Schedl by char- acters of the frons and elytral declivity as indicated below. Female. — Length 1.7 mm (paratypes 1.5-1.8 mm), 2.2 times as long as wide; color brown. Frons broadly, deeply concave from epistoma to vertex; surface evidently minutely, densely punctured and finely, densely, uniform- Sept. 1974 WOOD: American scolytidae 193 ly pubescent except smooth and glabrous on narrow median area on upper fourth; vestiture rather short, of uniform length. Antennal club 1.4 times as long as wide; strongly reniform, with grooved sutures, and with a small tuft of hair- on i)osterior face as in minutissimus. Pronotum and elytra as in minutissimus, except elytral declivity with interstriae 1 more strongly elevated, 2 weakly impressed, en- tirely unarmed, and devoid of setae, 4 feebly elevated and armed by al)OUt four minute tubercles. Vestiture as in minutissimus, except more slender, slightly longer, absent on interstria-e 2. Male. — Similar to female, except Irons convex, reticulate, sub- glabrous; antennal club normal, anterior margin of pronotum armed by a pair of coarse, median teeth. Type Locality. — Madden Forest, Canal Zone, Panama. l^pe Material. — The female holotype, male allotype, and eight paratypes were taken at the type locality on 2-1-64, at 70 m. No. 366, in tree branch, by me. Eight paratypes are from Ft. Clayton, Canal Zone, Panama, 22-XII-63, 30 m, No. 363, tree branch, by me. The holotype, allotype, and paratypes are in my collection. Corthylus trucis, n. sp. This species is distinguished from uniseptis Schedl by the larger size, by the armed declivial interstriae 2, and apparently by other characteristics. Female. — Length 3.0 mm (parat3^pes 2.9-3.1 mm), 2.3 times as long as wide; color dark brown, almost black. Frons deeply concave from eye to eye, from epistoma to vertex; surface smooth, shining, finely, closely jjunctured, except narrowly impunctate on entire median line; vestiture of fine, erect mioderate- ly long hair of uniform length, except a few longer setae on lateral margins. Antennal club large, strongly asymmetrical, 1.25 times as long as wide, with three rather strongly procurved sutures, median half of 1 septate, cirrus extending slightly beyond apex. Pronotum and elytral disc essentially as in comatus Blandford. Elytral decli^■ity broadly convex, steep; posterolateral margin sub- acutely elevated from costal margin to interstriae 8; surface shining, impressed points rather numerous, strial punctures only slightly larger than points, mostly in rows; interstriae 1 distinctly elevated, subcostate, 2 distinctly, rather shallowly impressed, 1-3 each armed by about four to six moderately large tubercles, usuall}- less numer- ous on 2. Vestiture mostly confined to declivity, of fine, long, inter- strial hair of moderate abundance. Male. — Similar to female, except frons convex, reticulate, sparsely punctured, subglabrous; antennal club much smaller, more nearly symmetrical, wdth only two sutures; anterior margin of pronotum more strongly serrate, median pair larger. 194 GREAT BASIN NATURALIST Vol. 34, No. 3 Type Locality. — Sixteen km SE Cartago, Cartago, Costa Rica. Type Material. — The female holotype, male allotype, and three paratypes were taken at the type locality on 24-IX-63, at 1800 m, No. 204, in Miconia glohuliflora. by me. The holotype, allotype, and paratypes are in my collection. Corthylus nolenae, n. sp. This species is distinguished from detrimentosus Schedl by the smaller size, by the female frons, and by the elytral declivity as noted below. Female. — Length 2.3 mm (paratypes 2.0-2.3 mm), 2.3 times as long as wide; color reddish brown, elytra black. Frons as in detrimentosus, except spongy area smaller, ending well below upper level of e3^es, dorsal half of excavated area with a strongly elevated median carina, marginal setae slightly longer, cirrus on antennal club smaller, shorter. Pronotum much as in detrimentosus. except pronotal disc with several broad subasperate crenulations, and elytral disc with strial punctures much larger and mostly in definite rows. Elytral declivity similar to detrimentosus but not as steej) and more narrowly convex; strial punctures rather coarse, deep; interstriae 1 distinctly elevated, 3 feebly elevated on basal half, 1 and 3 each with about four to six small tubercles; ventrolateral margin as in detrimentosus. Male. — Similar to female, except sexual differences as in detri- mentosus. Type Locality.- — Huajuapan, 21 km or 13 miles SE Oaxaca, Oaxaca, Mexico. Type Material. — The female holotype, male allotype, and 49 paratypes were taken at the type locality on 4-VII-53, in a Nolena fruiting stalk, by me. The holot3qDe, allotype, and paratypes are in my collection. Corythylus spinosus, n. sp. This species is distinguished in the male b}' the stouter bod}' and by the very different elytral declivity. Male. — Length 2.0 mm, about 2.0 times as long as wide; color very dark brown. Frons, antenna, ])ronotmn, and elytral disc as in uniseptis Schedl, except pronotum with summit nearer base, its disc with rather numerous low, rather broad, transverse crenulations. Elytral de- clivity much as in uniseptis, except wider, strial punctures slightly larger, interstriae 2 narrower and slightly deeper, 1 and 3 each bearing about four very coarse, sharply pointed tubercles, each high- er than wide (unusually large for this genus), several smaller tubercles in lateral areas. Vestituro slightly longer and more abun- dant than in uniseptis. Sept. 1974 WOOD: American sc;olytidae 195 Type Locality. — Fortin de las Flores-Sumidero, Veracruz, Mexico. Type Material. — The male holot'V'pe was taken at the type locality on 27-IV-65, 900 m, at blacklight, by H. V. Weems, Jr. The holotype is in my collection. Corthylus sentus, n. sp. This species is distinguished from minutus Bright by the larger size and b}' the characters of the female frons. Female. — Length 2.0 nun (paratypes 2.0-2.4 mm), 2.3 times as long as wide (2.4 times in Guatemalan specimens); color brown. Frons strongly concave from eye to eye, from epistoma to vertex; surface of upper half almost smooth, evidently with a few minute punctures, lower half from epistoma to upper level of eyes entirely occupied by a pair of oval, yellow, spongy areas, separated by a fine, low, acute, median carina, margins of spongy areas without a row of hair in most specimens, an irregular row in others. Antennal club strongly asymmetrical, apex subacute, sutures 1 and 2 moderately procurved, both partly septate at median end, a feeble suture 3 sug- gested; cirrus entirely absent. Pronotvmi as in uniseptis Schedl, except discal areas with numer- ous weak, broad crenulations. Elytral disc as in uniseptis, except punctures more obscure; declivity about as in spinosus, except punc- tures obsolete, interstriae 1 weakly elevated, unarmed, 2 impressed, strongly narrowed, essentially obsolete, 3 and lateral areas convex, 3 armed by about three coarse spines (larger than in spinosus), usually with one to two smaller spines at base, lateral areas with about four to eight pointed tubercles. Vestiture confined to declivity, of sparse hair. Male. — Similar to female, except frons convex, reticulate, sparse punctures obscure, antennal club smaller, less strongly asymmetrical; anterior margin of pronotum armed by one pair of median teeth. Type Locality. — Two km SE Cartage, Cartago, Costa Rica. Type Material. — The female holotype, male allotype, and 39 paratypes were taken at the type locality on 2-VIII-63, at 1300 m, No. 98, from a tree branch. Ten paratypes are from Cerro Pena Blanca, Honduras, 23-IV-64, 2000 m, No. 529, Miconia schlechten- dalii. Five paratypes are from Guatemala City, Guatemala, 30- V- 64, 1300 m, No. 644 in leguminous vine, or 643 in a shrub branch. One j)aratyj)e is from Quezaltenango, Guatemala, 26-V-64, No. 623, tree branch. All were taken by me. The holotype, allotype, and paratypes are in my collection. Corthylus pygmaeus, n. sp. This species is distinguished from spinifcr Schwarz by the smaller size and by the numerous minor features indicated below. 196 GREAT BASIN NATURALIST Vol. 34, No. 3 Female. — Length 1.8 mm (paratypes 1.5-1.8 mm), 2.0 times as long as wide; color brown. Frons as in spinifer, except more shallowly impressed (trans- versely flat at upper level of eyes), punctures averaging smaller, longest setae on upper third less numerous. Cirrus on antennal club shorter, its length from insertion of last hair to its apex less than length of club. Pronotum and elytra as in spinifer, except declivital sulcus not quite as deep, punctures slightly finer, impressed points largely obso- lete, posterolateral margin more abruptly, less strongly elevated. Male. — Similar to female, except sexual differences as in spinifer. Type Locality. — Finca La Lola, Limon, Costa Rica. Type Material. — The female holotype, male allotype, and four paratypes were taken at the type locality on 17-1-63, from Theo- bronia cacao branches, by J. L. Saunders. Eight paratypes are from Pandora, Limon, Costa Rica, 23-VIII-63, 50 m. No. 149, tree branch, by me. The holotype, allotype, and paratypes are in my collection. Corthylus sobrinus, n. sp. This species is distinguished from flagellifer Rlandford by the larger size and by characters of the frons and elytral declivity men- tioned below. Female. — Length 3.0 mm (paratypes 3.0-3.4 mm), 2.1 times as long as wide; color ver}^ dark brown. Frons as in flagellifer, except much more strongly impressed, particularly on upper half; lower fifth with a pair of rather widely separated, oval, yellow spongy areas (vestiture not interrupted by their presence) ; vestiture similar to flagellifer.. except most setae strongly flattened, less abundant at upper margin. Antennal cirrus much more poorly developed but only slightly shorter. Pronotum and el3'tra as in flagellifer, except punctures on el}'tral disc larger but very shallow, and elytral declivity somewhat more strongly, more broadly impressed. Male. — Siniiln?- to female, except sexual differeiues a'^ in spinifer. Type Locality. — Turrialba, Cartago, Costa Rica. Type Material. — The female holotype, male allotype, and 14 paratyi)es were taken at the type locality on 7-L63, in Theobroma cacao, by J. L. Saunders. Two paratypes are from the same locality taken 9-III-64, 700 m. No. 468B, from a fence post, by me. The holotype, allotype, and paratype are in my collection. Sept. 1974 WOOD: American scolytidae 197 Corthylus subserratus, n. sp. This species is distinguished from scrrulalus Eggers (from BoHvia and Peru) by the larger size and by conspicuous differences on the head and elytral declivity as indicated below. Female. — Length 2.5 mm (parat3pes 2.3-2.5 mm), 2.4 times as long as wide; color black, with one-third to two-thirds of elytra yel- lowish brown (area and color intensity of pale portion variable). Frons deeply concave from eye to eye, from epistoma to vertex, much deeper than in serrulatus; entire lower third occupied b}' yel- low spong}- area, upper areas dull, coarsely punctured; vestiture absent at center of concavity, increasing in size and density toward margins above eyes; margins above eyes ornamented by a dense brush of long, yellow^, strongly plumose setae (as in serrulatus) . Anteimal club as in serrulatus, with suture 1 partly septate, 2 and 3 weakly impressed; cirrus entirely absent. Pronotuni as in serrulatus, except posterior areas more strongly reticulate, weak transverse crenulations on disc much narrower. Elytra as in serrulatus, except surface of disc less shining; declivity less convex, posterolateral margin ending slightly below middle of declivity, surface strongl}^ reticulate, pmictures mostly obscure, inter- striae 1 slightly less strongly elevated, its summit conspicuously undulating, forming about six rounded serrations (summit an almost uniform costa in serrulatus) . Vestiture confined to declivity, of rows of erect interstrial hair on interstriae 1 and 3 and base of 2, a few setae in lateral areas. Male. — Similar to female, except frons convex, reticulate, sparsely punctured, vestiture sparse, inconspicuous; antennal club smaller, more nearly symmetrical; anterior margin of pronotum armed by one median pair of teeth. Type Locality. — Volcan Poas, Heredia, Costa Rica. Type Material. — The female holotype, male allotype, and 65 paratypes were taken at the type locality on 19-XI-63, 2500 m, Nos. 260, 261 (type), from tree branches. Twenty-eight paratypes were taken 14-VII-63, No. 44B, 12 paratypes were taken 6-IX-63, No. 169 from Vaccinium consanguineum, and four paratypes were taken 19-VII-66, at 1300 m, from the same locality. Three paratypes are from Guapiles, Limon, Costa Rica, 22-VIli-66, 100 m. All were taken by me. The holotype, allotype, and paratypes are in my collection. Corthylus serratus, n. sp. This species is distinguished from serrulatus Eggers and subser- ratus Wood by the smaller size and by conspicuous differences on the head and elytral declivity as noted below. Female. — Length 2.0 mm (paratypes 2.0-2.3 mm), 2.5 times as long as wide; color dark brown. 198 GREAT BASIN NATURALIST Vol. 34, No. 3 Frons basically shallowl}- concave as in scrrulatus, and with oval area on median half from near epistoma to near vertex abruptly, much more strongl}- concave; central concavit}' smooth, with a few {)unctures and hairlike setae; lateral areas minutely, densely punc- tured, becoining spongy and yellowish above on median margins, this area very minutely, densely pubescent; plumose vestiture on upper margin as in subserratus. Antenna as in serrulatus. except cirrus smaller, slightly shorter. Pronotum and elytral disc as in serrulatus, except punctures on elytral disc slightly larger. Elytral declivity with margins abruptly angled, except rounded on basal fifth, face much more weakly con- vex than serrulatus. its surface shining and punctured about as in serrulatus^ interstraie 1 more strongly elevated than in serrulatus, its crest with about four to six coarse crenulations, some of them acutely pointed. Interstriae 1, 3, and 4 with very long hairlike setae. Male. — Similar to female, except sexual differences as in sub- serratus. Type Locality. — Tapanti, Cartago, Costa Rica. Type Material. — The female holotype was taken at the type locality on 24-X-63, 1300 m, No. 241, in Phoebe mexicana; two paratypes are from the same locality, one taken 2-VII-63, No. 9, Miconia, and one 24-XI-63, No. 271, Werklia insignis. Three para- types are from 16 km SE Cartago, Cartago, Costa Rica, 24-IX-63, 1800 m. No. 204, Miconia globuli flora. The male allotype and 11 paratypes were taken from Cerro Punta (labeled Volcan Chiriqui), Chiriqui, Panama, 11-1-64, 1800 m. No. 371 in Inga (allotype), Nos. 388, 399, in tree branches; four paratypes from the same locality were taken 19-XII-63, No. 315, Inga. All were taken by me. The holotype, allotype, and paratypes are in my collection. Corthylus donaticus, n. sp. This species is distinguished from serratus Wood by the larger size, by the less strongly elevated declivital interstriae 1, by the abrupt, angulate margins being restricted to the lower half of the elytral declivit}, and by the different female frons as described below. Female. — Length 2.4 mm (paratypes 2.4-2.6 mm), 2.2 times as long as wide; color very dark brown. Frons rather deeply concave from eye to eye from epistoma to vertex, a feeble median carina on floor of upper half of concave area; lateral area just mesad of lateral margin with an elongate, elevated, yellow spongy area from up])er level of eyes to vertex, dorsal ends of these spongy areas separated from one another by a distance slightly less than half cHstance between eyes; spongy areas with minute [)ilo, their lateral margins marked by a dense row of long yellow hair very similar to but shorter than in serratus. Pronotum and elytral disc as in serratus. Elytral declivity as in Sept. 1974 WOOD: American scolytidae 199 serratus. except lateral margin subacutely angulate only on lower half and interstriae 1 less strongly elevated, crest almost as high as wide at highest point; setae on interstriae 1 short to obsolete. Male. — Similar to female, except sexual differences as in ser- ratus. Type Locality. — Colonia Tovar, Aragua, Venezuela. Type Material.- — The female holotype, male allotype, and 28 paratypes were taken at the type locality on 4-V-70, 1700 m, Nos. 478 (1), 484 (8), 487 (1), 493 (2), 496 (holotype and 3 paratypes), 497 (allotype and 9 paratypes), 502 (2), 509 (2), from various hosts, by me. The holotype, allotype, and paratypes are in my collection. Corthylus additus, n. sp. This species is distinguished from serratus Wood by the larger size, by the more slender body form, and by the female frons as described below. Female. — Length 2.7 mm (paratypes 2.3-2.7 mm), 2.8 times as long as wide; color very dark brown. Frons similar to serratus, except lower half of excavated area as strongly impressed as dorsal half; lateral margins of impressed area narrowly, more strongly elevated, spongy and with Aery short pile- like pubescence from upper level of eye to upper level of ocular emargination, these spongy areas of yellow color; pubescence about as in serratus except slightly more abundant, a row of longer hair along lateral margins of spongy area to epistoma. Antenna as in serratus, with sutures 1 and 2 clearly marked, 1 septate. Pronotum and elj^tra as in serratus. Male. — Similar to female, except sexes differing as in serratus. Type Locality. — La Carbonera experimental forest, about 50 km (airline) NW Merida, Merida, Venezuela. Type Material. — The female holotype, male allotype, and 19 paratypes were taken at the type locality on 14-X-69, 2500 m. No. 50 (12) and 51 (1) in Clusia, 53 (2) and 61 (holotype, allotype, 2 parat3'pes), and 62 (1) in Nectandra, 66 (1). Other paratypes from the same locality were taken lO-X-69, No. 121 (1) in Rubus, 122 (2) and 123 (1) in a woody vine, 125 (2) in Nectandra. 128 (10) in a tree seedling; 9-Xn-69, No. 171 (2) in a small log, 174 (2) in Nectandra. Three paratypes are from La Mucuv, Merida, Vene- zuela, 20-X-69, 2500 m. No. 74 (1) in Nectandra. and 12-XL69. No. 131 (1) and 132 (1) in tree branches. Two ])aratypes are from Merida, Merida, Venezuela, 22-IX-69, 1700 m. No. 8 in Ficus and 12 in Vismia. All were taken by me. The holotype, allotype, and paratypes are in my collection. 200 GREAT BASIN NATURALIST Vol. 34, No. 3 Corthylus cirritus, n. sp. This species is distinguished from additus Wood by the stouter body form, by the subangulate margin of the elytral declivity ex- tending from the apex to the base, and by the conspicuous differences in the female frons and antenna as described below. Female.- — Length 2.3 mm (paratypes 2.2-2.4 mm), 2.3 times as long as wide; color very dark brown. Frons similar to additus, except dorsal half more broadly im- pressed, spongy areas less strongh' elevated, slightly wider; hair on lateral margins more abundant, extending to level of antennal inser- tion, a separate tuft of plumose hair on lateral areas of epistoma. Antennal club similar in shape to additus but sutures replaced by three arcuate calluses; cirrus slender, exceedingly long, more than four times length of club. Pronotum and elytral disc as in additus. Declivity similar to additus, except margin subacutely elevated on a complete circle, interstriae 1 less strongly elevated, tubercles on interstriae 3 almost obsolete. Male. — Similar to female, except sexual differences as in ser- ratus Wood. Type Locality. — La Carbonera experimental forest, about 50 km (airline) NW Merida, Merida, Venezuela. Type Material. — The female holotype, male allotype, and 23 paratypes were taken at the type locality on 9-IV-70, 1100 m, Nos. 404 (1), 412 (11), 413 (holotype, allotype, 6 paratypes), 419 from Piper (1), 431 from Nectandra (2), 433 (1), 445 (1), by me. The holotype, allotype, and paratypes are in my collection. Corthylus insignis, n. sp. This species is distinguished from schaufussi Schedl by the smaller size and by characters of the frons and elytra as described below. Eggers erroneousl}' considered this species to be cornpressi- cornis (Fabricius), as indicated by his specimens "mit type ver- gleichen." Female. — Length 1.8 mm (paratypes 1.8-2.1 mm), 2.2 times as long as wide; color very dark brown. Frons as in .schaufussi, except short, pale setae in central area more strongly flattened, and more strongly plumose, marginal red- dish, long setae much more densely placed, particularly on vertex and toward bases of mandibles. Antennal club as in schaufussi. Pronotum as in schaufussi, except disc more strongly reticulate and with minute, transverse crenulations much more obscure, almost obsolete. Elytral declivity as in schaufussi except steeper, inter- striae 1 less strongly elevated, with its crest smooth, wdthout indica- tions of serration; lateral areas more evenly convex, denticles on interstriae 3 much smaller. Sept. 1974 WOOD: American scolytidae 201 Type Locality. — Twenty-four km E Barbosa, Antioquia, Co- lombia. Type Material. — The female holotype, male allotype, and 26 paratypes were taken at the type locality on 18-VII-70, at 1200 m, No. 699, in Inga, by me. The holotype, allotyj^e, and paratypes are in my collection. This species was reported by Schedl from Brazil and Peru as praeustus Schedl; praeustus apparently is restricted in its distribu- tion to Central America. Corthylus concisus, n. sp. This species is distinguished from praeustus Schedl by the larger size and by the characters on the head mentioned below. Female. — Length 2.2 mm (paratypes 2.0-2.3 mm), 2.1 times as long as wide; color dark brown to almost black. Frons flat, a weak transverse callus at level of antennal insertion; large central area smooth, glabrous, impunctate, marginal areas with fine punctures; lateral, submarginal areas with a tuft of hair from level of antennal insertion to upper level of eyes, margin above upper level of eyes ornamented by a dense row of very long plumose setae. Antennal club very broad, 1.4 times wider than long, cirrus at least three times longer than club (club longer than wide and cirrus shorter than club in praeustus) . Pronotum and elytral disc essentially as in petilus Wood, except pronotal disc \\ath minute, transverse crenulations and punctures on elytral disc minute, confused. Elytral declivity truncate, subvertical, margin abruptly, subacuteh' elevated on a complete circle from apex to base; face slightly convex, strongly reticulate, punctures small, confused; interstriae 1 strongly costate, as high as wide on middle half, middle area of 2 distinctly impressed, 3 weakly elevated on upper half and armed by about two to four small, pointed tubercles. Glabrous except for a few setae on declivital interstriae 1 and 3. Male. — Similar to female, except sexual differences as in petilus. Type Locality. — Tapanti, Cartago, Costa Rica. Type Material. — The female holotype, male allotype, and 15 paratypes were taken at the type locality on 17-VIII-63, 1300 m. No. 106, from a woody vine; two paratypes are from the same locality taken 24-X-63, "Nos. 224, 245, and one paratype 2-VII-63, No. 10, Conostegia oersted iana. Other paratypes include 20 from Turrialba, Cartago, Costa Rica, 7-1-63, Theobroma cacao, J. L. Saunders; 1 from 16 km SE Cartago, Cartago, Costa Rica, 3-VII-63, No. 11, woody vine; 6 from Peralta, Cartago, Costa Rica, 10-III-64, 500 m, Nos. 461, 462, woody vines; 2 from Puerto Viejo, Heredia, Costa Rica, 12-III-63, 70 m, No. 480, woody vine; 2 from Volcan, Puntarenas, Costa Rica, ll-XII-63, 1000 m. No. 304, tree branch; 4 from Escasu, San Jose, Costa Rica, 2-X-63, No. 215 in Guazuma ulmifolia, No. 216 in Spondias purpurea. No. 218 in a tree seedling; 202 GREAT BASIN NATURALIST Vol. 34, No. 3 10 from Santa Ana, San Jose 4-X-63, 1300 m. No. 223, tree branch, and 5 on 9-X-63, No. 229, tree branch; 2 from Rio Damitas, Dota Mts., San Jose, 18-11-64, 250 m. No. 436, tree seedling; 4 from 13 km or 8 miles S El Ha to del Volcan, Panama, 7-1-64, 1000 m. No. 371, tree seedling; 3 from Cerro Campana, Panama, 26-VII-66, 1000 m, No. 32, tree branch. All were taken by me except as noted. The holotype, allotype, and paratypes are in m}^ collection. This species was treated as Corthylus compressicornis by Bland- ford (1904, Biol. Centr. Amer. Coleopt. 4(6):255). Corthylus compressicornis (Fabricius) Bostrichus compressicornis Fabricius, 1801, Systema Eleutheratorum 2:388 (Lecto- type, female; Essequibo, British Guiana; Copenhagen Mus., present designa- tion) Two female type specimens of this species are in the Fabricius collection. The pin of each specimen bears a small green label without writing, and a red label with the printed word "Type". In addition, the first specimen bears a handwritten label, "Essequibo. Schmidt. Mus. J. Lund. Bostrichus compressicornis Fabr." The first specimen is here designated as the lectotype of Bostrichus com- pressicornis Fabricius and now bears my red lectotype label. Female lectotype. — Length 2.2 mm, 2.1 times as long as wide; color dark brown. Frons visible to upper level of eyes; flat to feebly concave, lateral margins along inner margin of eye subacutely elevated, ending ven- trally in a small denticle just below upper limits of ocular emargina- tion; epistomal margin weakly elevated; surface of subconcave area smooth and shining except for two large, contiguous, longitudinally oval, spongy areas occupying middle half from level of antennal in- sertion to a point well below upper level of eye; spongy areas yellow, almost smooth, their margins acutely elevated well above surfaces of frons or spongy areas; surface of spongy areas elevated slightly above surface of remainder of frons; areas of frons dorsad from spongy areas bearing fine, long, yellow hair over entire surface. Antennal club strongly asymmetrical much as in concisus Wood, and bearing a cirrus twice as long as width of club. Pronotum as in schaufussi Schedl, except less strongly reticulate and transverse discal rugosites much more obscure. Elytra as in schaufussi, except disc less shining, declivity dull. Pronotum 1.2 times as long as elytral disc. Notes. — This species has been reported from A^arious parts of Central and South America; however, except for the two females in the Fabricius collection, it is unknown to me. At least two Central American species, concisus Wood and praeustus Schedl, have been reported as this species. The confusion of South American forms with this species is much more complex. NOTES ON VISCAINOPELMATUS DAVEWERNERI (ORTHOPTERA: GRYLLACRIDIDAE; STENOPELMATINAE) FROM THE VISCAINO DESERT, BAJA CALIFORNIA SUR, MEXICO Ernest R. Tinkham,i Clark R. Mahrdt,- and Benjamin H. Banta^ Abstract. — A male nympkh in stadium 3 of Viscainopelmatus davewerneri Tinkham is reported and illustrated. The specimen was taken near Laguna Scam- mon, Baja California Sur, Mexico. Tinkham (1970) described Viscainopelmatus davewerneri on the basis of a single specimen of undetermined sex (lacking an abdo- men) from the coastal dunes at Laguna Manuela, approximately 25 km NE of Guerrero Negro, Baja California Norte, Mexico. The holotype was found while excavating a rodent burrow on 5 July 1965. We report a second specimen obtained from a rodent burrow by Banta on 30 December 1971. The specimen, deposited at Cali- fornia Academy of Sciences (CAS), was taken on the southwest side of Laguna Scammon, approximately 4 km east of the lagoon entrance, Baja California Sur, Mexico. The site is located at 114° 20' W long., 27° 45' N lat, near sea level. The dominant vegetation in this area of predominantly coastal sand dunes includes Abronia maritima, Lycium sp., Seauvium verrucosum. Despite numerous visits in this area by scientists of diverse disci- pUnes, only two specimens have been obtained. Tinkham and Mahrdt visited the area on two separate occasions (June 1968 and January 1971, respectively) but failed to obtain additional stenopel- matids. The second specimen, a male, is apparently in the third stadium of nymphal development, and is significantly smaller than the type (Table 1). Size variation cannot be attributed to sexual Table 1 neri. I. Morphometric data (in millimeters) of Viscainopelmatus davewer- Source Body Length Head + Thorax Pronotum Caudal Femur Caudal Tibia Type (Tinkham, 1970) CAS ca. SO- BS mm 26.8 11.3 9.5 7.2 X 8.8 4.2 x 5.4 10.0 X 4.1 6.7 X 2.4 10.8 X 3.5 7.2 X 2.0 dimorphism, because male and female stenopelmatids are usually consistent in their morphological features (Tinkham, 1970). The main difference between the two specimens, other than size, is that in the nymph the dorsal outline of the pronotum is rather quadrate ^81-441 Date Palm, Indio, California 92201. =San Diego Natural History Museum, P.O. Box 1390, San Diego, California 92112. 'Department of Biology, United Stales International University, San Diego, California 94131. 203 204 GREAT BASIN NATURALIST Vol. 34, No. 3 Figs. 1-12. Viscainopelmatus davewerneri: comparison of the holotype to the male stadium 3 nymph taken near Scammon Lagoon. 1, Apical chaetotaxy of right foretibia, ventral aspect, nymph; 2, same for holotype; 3, ventral aspect of right mesotibia of nymph; 4, same for holotype; 5, apical chaetotaxy of meta- tibia of nymph, dorsal aspect; 6, left metathoracic tarsus, showing plantar surface from ventromesal aspect, nymph; 7, same from right tibia of holotype; 8, outline of pronotum of nymph from dorsal aspect; 9, apical chaetotaxy of left meta- thoracic tibia of holotype, dorsal aspect; 10, apical terminalia of nymph, dorsal aspect; 11, outline of pronotum of holotype; 12, same, ventral aspect. AH drawn with the aid of a stereoscopic microscope at 15 X magnification. (Fig. 8), whereas in the type the pronotum is ampliate forward with all angles well rounded (Fig. 9). The supra-anal and sub- genital plates of the nymph are roundly triangular and rather similar (Figs. 10, 11). Leg Spination: Figures 1 and 2 show that the foretibiae, exclu- sive of size, are similar. Figures 3 and 4 depict identical chaetotaxy in the calcars, but the two dorsal subapical teeth are the merest black dots in the nymph. The dorsal subapical teeth are of impor- tant taxonomic value in Stenopelmatidae. Figures 5, 6, and 7 show relative sizes of the caudal tibia and tarsus and reveal that the pe- culiar feature of having a laterally flattened and twisted caudal tarsus, characteristic of Viscainopelmatus, is evident in the third Sept. 1974 TINKHAM ET AL: DESERT ORTIIOPTERA 205 Stadium nymph. The nymph differs from the t}'pe in possessing only two minute, subapical, vestigial, external dorsal teeth instead of three. In Tinkham's (1970:174) key the sixth line should be amend- ed to read, "only 2 or 3 very small, vestigial, external, dorsal, sub- apical teeth." The coloration of the head, thorax, and abdomen in alcohol is brown pink to tan, 91 -L in Maerz and Paul (1950). The abdomen is unicolor without alternating black and white bands. V iscainopclmatus davcwcrncri, pre\'iously known only from the type locality, occupies a coastal sand dune habitat. Our specimen >- *% \ ' < 4. Fig. 13. Viscainopelmatus davewerneri male nymph in stadium 3: 1, dorsal aspect; 2, ventral aspect; 3, ventral aspect showing black-edged spathulate calcars of meta thoracic tibiae and twisted tarsi; 4, ventral aspect of protibial calcars. 206 GREAT BASIN NATURALIST Vol. 34, No. 3 was taken in ecological conditions like those of the type; its occur- rence 60 km southwest on the south side of Scammon Lagoon further indicates a distribution confined to coastal sand dunes of the Viscaino Desert. Indeed, the salient features show an adaptation to such an arenicolous environment. However, there is still a need for more intensive sampling in the Viscaino Desert to discern the distribu- tional parameters and extent of variation in this species. Acknowledgments: We are grateful to Dr. Theodore Cohn for suggestions and criticisms of the manuscript, and to Mr. Randall Nieman for photographs of figures 1-4. Literature Cited Maerz, a. J., AND M. R. Paul. 1950. A dictionary of color. McGraw-Hill, New York. 208 p. TiNKHAM, E. R. 1970. Studies in Nearctic Desert sand dune Orthoptera. Part 12. A remarkable new genus and species of stenopelmatine crickets from the Viscaino Desert, Baja California, Mexico, with key. Great Basin Nat. 30:173-179. THE AUSTRINA GROUP OF THE GENUS MICROVELIA (HEMIPTERA; VELIIDAE) John T. Polhemus^ AssTRfVCT. — Six new species of Microuelia ai-e described from North and Central America: depressus, glabrosulcata, leavipleura, paura, psilonota, and reflexus. These species are compared to Microuelia austrina Bueno, and the austrina group is designated to hold them. The habitats and phoresy of the group are discussed. Microuelia austrina Bueno is a widespread species, occurring from the southeastern United States into Central Mexico. No closely related species have been previously described. Through extensive collecting from the United States to Panama, I have been able to secure series of Microuelia containing a complex of closely related species which I here designate as the austrina group. In the austrina group the males offer little in the way of diag- nostic characters, but the apterous females exhibit various modifica- tions to facilitate the males' riding astride or "piggyback," and these features provide excellent specific characters. Many of the small microvelias exhibit such modifications in the female, but they have rarely been used as ke}" characters (Polhemus, 1970). The late Carl Drake described many Microuelia, including some with obvious female modification, but did not use the character in discrimination, which may explain why several of the species described below stand in his collection under M. austrina. (Loans from the Drake collec- tion are not permitted, so those specimens are not included in this study). All of the species of the austrina group are found in cryptic habitats during daylight hours as they are apparently strongly nega- tively phototropic. Only once were members of this group observed running on open water; this was in the early morning in canopy jungle, where M. paura n. sp. could be seen in the dim light as dark specks moving slowly over a pool in the small stream. The most successful method of collecting is violent sweeping and splashing in dark caverns or under overhanging banks adjoining slow clear water streams. Many specimens collected were paired even after being dashed about, but no mating pairs were observed; Esaki (1937) noted a similar phoresy in M. notophora Esaki. The modifications to facilitate the male's riding "piggyback" would seem to serve two purposes. First, a male would be readily available at the time of mating; second, the specialized modification exhibited by each species would be a prcmating reproductive iso- lating mechanism. At this time it is not known whether or not there are strong postmating reproductive isolating mechanisms in the group, but several species are sympatric, and as Littlejohn (1969) has pointed out, premating isolation would permit the most efficient use of gametes. >3115 S. York, Englewood, Colorado 80110 207 208 GREAT BASIN NATURALIST Vol. 34, No. 3 The austrina group will be described first, giving the generalized characters; the species descriptions will follow, providing additional detail and key characters. For all measurements 60 units equals 1 mm. Females are intentionally named as holotypes: those types being retained in the Polhemus collection are irrevocably committed to later placement in a designated type repository. Much of the material reported here is in the Polhemus collection (JTP). Most of the remaining specimens belong to the University of Colorado Museum (CU), and I am indebted to Dr. Peter Robinson for the opportunity to study them. The latter were collected by Martin S. Polhemus and myself during a Mexican Expedition partly financed by a grant from the University of Colorado. I am indebted to the late Robert L. Usinger for permitting me to study specimens from the University of California at Rerkeley (UCB), to the late Jean L. Lafoon for the exchange specimens from the Iowa State Uni- versity Collection, to Harold C. Chapman for the gift of specimens from his collection, and to Peter D. Ashlock for the loan of recently collected specimens from the University of Kansas (KU). Microvelia austrina Group Description. — Small (1.2 to 2.2 mm), stout (see Figs. 1, 2). Head short, broad, recessed into anterior margin of pronotum; ground color blackish brown with fine greyish pubescence. Prono- tum long, covering mesonotum. Legs short, stout. Antenna short, stout, fourth segment fusiform. Hemelytral markings and veins in- conspicuous. Body of male somewhat depressed dorsoventrally. Male parameres small, inconspicuous; male genital segment not prominent (Fig. IE, F). Male foretibial comb prominent, extending beyond distal end of tibia. Key to the Microvelia austrina Group 1. Humeri depressed in both sexes depressus n. sp. Humeri not depressed 2 2. Anterior lobe of pronotum with transverse orange brown band extending laterally onto episternum; body length at least 1.95 mm in male, 2.15 mm in female .... rcflexus n. sp. Anterior lobe of pronotum with shorter transverse orange brown band, usually not extending past lateral margins; if transverse band extends onto episternum {paura), then body length not more than 1.3 mm in male, 1.75 mm in female 3 3. Distal segment of middle tarsi either shorter or longer than proximal segment by about 20 percent; coxae yellowish or dark brown 4 Distal segment of middle tarsi subequal to proximal segment; coxae yellowish 5 Sept. 1974 POLHEMUS: AQUATIC HEMIPTERA 209 4. Distal segment of middle tarsi longer than proximal segment (7:9); coxae dark brown glabrosulcuta n. sp. Distal segment of middle tarsi shorter than proximal segment (11:9); coxae yellowish paura n. sp. 5. Proepisternum yellowish or light orange brown with very obvious but tiny black conical setae psilonota n. sp. Venter unicolorous or proepisternum dark orange brown; pro- episternum not light colored and black conical setae not obvious 6 6. Posterior acetabula with glabrous area laevepleura n. sp. Posterior acetabula without glabrous area .... austrina Bueno Microvelia austrina Bueno 1924 Microvelia austrina Bueno 1924. Bull. Brooklyn Ent. Soc. 19(5): 191 (North Carolina) Drake and Hussey (1955) gave the distribution of this species as Indiana, Maryland, Mississippi, North Carolina, Tennessee, Virginia, and Mexico. Description. — Female body shape elongate, quite similar to Fig. 1. Microvelia leavipleura n. sp.: a, Apterous male; b, Apterous female; c, Macropterous female; d. Pronotum and pleura, apterous female, side view; e. Genital segments, male; f. Male paramere. 210 GREAT BASIN NATURALIST Vol. 34, No. 3 Fig. 2. Microvelia spp.: a, paura n. sp., apterous female; b-g. Pronotum and pleura, apterous females, side view; b, paura n. sp.; c, depressus n. sp.; d, psilonola n. sp.; e, glabrosulcala n. sp.. f, reflexus n. sp., g, austrina Bueno. Figure IB; male body shape elongate, tapering caudad, very similar to Figure lA. Anterior lobe of pronotum with yellowish to dark orange brown band, continuous or weakly interrupted on midline, extending almost to lateral margins in some females, shorter and darker in males. Anterior lobe of pronotum, humeri, and pleura of apterous female with long stiff setae (Fig. 2G) ; alate females with a patch of stiff setae on anterolateral margins of pronotum; apterous and alate males with shorter scattered setae on pronotum and elytra. Apterous fe- male without glabrous areas on pronotum or pleura, but with semi- glabrous depressed area on propleura and mesopleura. Connexiva slightly raised in male, almost flat to vertical in ap- terous females. Dorsum of abdomen covered with short inconspicu- ous pubescence. Venter uniforml}' dark, except dark orange brown next to eyes on proepisternum, anterior and middle acetabula; venter of abdomen with semilong decumbent setae in female, shorter in male. Rostrum basally, coxae, trochanters, femora basally and legs beneath yellow- ish to testaceous, remainder of extremities brown. Legs with the following proportions (apterous female plesiotype) : Tarsal 2 13 12 Anterior Middle Posterior Femur 28 32 33 Tibia 20 26 36 Tarsal 1 13 13 Antenna! proportions I; II; III; IV: 9; 10; 12; 14. Body measurements: Mean length of 10 apterous $ $ : 1.59 mm (max. 1.70, min. 1.47). Mean width of 10 apterous $ $ : 0.60 mm (max. 0.68, min. 0.55). Sept. 1974 POLHEMUS: AQUATIC HEMIPTERA 211 Mean length of 10 apterous $ $ : 1.87 nun (max. 2.00, min. 1.80). Mean widtli of 10 apterous $ 9 : 0.74 mm (max. 0.80, min. 0.70). Plesiotype: Length 1.88 mm, width 0.72 mm (apterous 9 ). Material examined. — MEXICO: San Luis Potosi,- 1 $ apterous, 25 S $ , 23 $ $ macropterous, Ciudad del Maiz, CL528, 9-1-1971. J. T. & M.S. Polhemus (JTP); Ver.\cruz; \ $. { 9 apterous, 8 $ $, 2 9 9 macropterous, Tantoyuca, CL523, 8-1-1971. J. T. & M. S. Polhemus (JTP). UNITED STATES; Georgia,- Calhoun, 8-VIII-1958, H. C. Chapman, 22 $$, 17 9 9, all apterous (JTP). Tennessee,- Kno.xville, 27-VI-1891 to 2-\TM891, H. E. Summers Collection, 13,4 9 9, all macropterous (JTP). Discussion. — The description of austrina by Torre-Bueno (1924) is quite adequate but is given here for consistency. Curiously, Torre- Bueno chose a female for the type of austrina, but undoubtedly with a different motivation than mine in describing additional members of the group. Austrina can be separated from the other group species by the key characters and by the small degree of modification in the apterous female. Microvelia depressus, n. sp. Female bod}- shape elongate oval, widest across metapleura; male body shape similar to female but slightly narrower. Anterior lobe of pronotum with orange brown band reaching inner margin of eyes, often weakly interrupted medially. Female pronotiun with two groups of stiff but slender, long setae, most noticeable from side (Fig. 2C) ; male with shorter scattered setae; humeri depressed in both sexes, pronotum narrowed laterally, and entire dorsum depressed below level of first visible abdominal tergite; female wdth glabrous area on lateral margin. Connexiva moderately raised in males, usually vertical in females or reflexed over last three tergites. Venter uniformly dark except small brown spot on proepisternum cephalad of anterior coxae; jugum of head and entire venter covered ^^'ith minute black conical setae, most noticeable on brown area of proepisternum. Legs, rostrum, and antennae brown; basal part of femora and trochanters yellowish. Posterior acetabula with glabrous spot. Legs with following proportions: Anterior Middle Posterior Antennal proportions I; II; III; IV: 10; 8; 7; 13. Body measurements: Mean lengtli of 9 apterous $ $ : 1.74 mm (max. 1.78. min. 1.53). Mean width of 9 apterous $ $ : 0.75 mm (max. 0.77, min. 0.72). Mean length of 10 apterous 9 9 : 167 mm (max. 1.72, min. 1.60). Mean width of 10 apterous 9 9 : 0.78 mm (max. 0.80, min. 0.73). Holotype: Length 1.67 mm, width 0.78 mm. Material examined. — Holotvpe, apterous 9, 10 mi. N Chapala. Jalisco, Mexico, 22-IV-1964, J. T. & M." S. Polhemus (CU). Paratypes as follows; Femur Tibia Tarsal 1 Tarsal 2 21 17 10 . . 27 22 8 11 30 30 5 8 212 GREAT BASIN NATURALIST Vol. 34, No. 3 MEXICO: Jalisco,- 9 $ $ , 18 9 9 apterous, 10 mi. N Chapala, 22-IV-1964, J. T. & M. S. Polhemus (CU, JTP). Discussion. — The strongly depressed humeri in both sexes are characteristic of this species and set it off from the remainder of the group. Microvelia glabrosulcata, n. sp. Female body shape elongate, abdomen slightly wider than thorax, not as robust as psilonota; male body shape elongate, tapering cau- dad, similar to Figure lA. Anterior lobe of pronotum with orange brown uninterrupted band reaching inner eye margin, rarely beyond. Anterior lobe of alate and apterous females with long stiff setae, apterous females also with stiff setae on humeri directed cephalad (Fig. 2E), thinner long setae on propleura of females; males with shorter scattered setae, not prominent. Apterous and alate females with a shallow glabrous sulcus on lateral margin of pronotum cephalad of humeri. Connexiva almost flat in males and some females, strongly raised in most females, rarely almost vertical. Venter uniformly dark, except proepisternum dark grey brown; entire venter set with minute black conical setae, most noticeable on episternum. Legs, rostrum, and antennae brown, anterior femora lighter beneath. Acetabula with prominent glabrous areas. Legs with following proportions: Femora Tibia Tarsal 1 Tarsal 2 Anterior 24 17 10 — Middle 25 22 7 9 Posterior 26 26 6 9 Antennal proportions I; II; III; IV: 8; 7; 6; 10. Body measurements: Mean length of 10 apterous $ $ : 1.34 mm (max. 1.38, min. 1.28). Mean width of 10 apterous $ $ : 0.58 mm (max. 0.60, min. 0.55). Mean length of 10 apterous $9 : 1.66 nun (max. 1.72, min. 1.58). Mean width of 10 apterous $ 9 : 0.79 mm (max. 0.82, min. 0.72). Holotype: Length 1.62 mm, width 0.73 mm. Material examined. — Holotype, apterous 9 , Ciudad del Maiz, San Luis Potosi, Mexico, 9-1-1971, CL528, J. T. & M. S. Polhemus (in Polhemus Collec- tion). Paratypes as follows; MEXICO: San Luis Potosi,- 10 5 5 , 22 9 9 apterous, 14 $5, 21 9 9 macropterous, Ciudad del Maiz, 9-1-1971, CL528, J. T. & M. S. Polhemus (JTP). Jalisco,- 1 $, 1 9 apterous, 10 mi. N Chapala, 22-IV-1964, CL1031, J. T. & M. S. Polhemus (CU). UNITED STATES: Arizona; 2 $ 5, 11 9 9 apterous, 1 9 macropterous, Superior, 7-X-1964, CL313, J. T. Polhemus (JTP); 3 $$,7 9 9 apterous, Sabino Canyon, 27-XI-1970, CL497, J. T. Polhemus (JTP). Discussion. — This species is widely distributed, but its known range is restricted to northern Mexico and Arizona. At Ciudad del Maiz, San Luis Potosi, it was taken with austrina. The glabrous sulcus on the female pronotum, the dark coxae, and the long distal segment of the middle tarsi are diagnostic. The body shape is compact and somewhat resembles depressus. Sept. 1974 POLHEM US: AQUATIC HEMIPTERA 213 Microvelia leavipleura, n. sp. Female body shape elongate (Fig. IB) ; male body shape elongate, tapering caudad (Fig. lA). Anterior lobe of pronotum with yellowish to dark orange brown band, continuous or weakly interrupted on mid-line, not extending to lateral margins. Anterior lobe of pronotum, humeri, and pleura of apterous female with long stiff setae (Fig. ID); anterior lobe of pronotimi and corium of hemelytra in alate female with long stiff setae; apterous and alate males with shorter scattered setae on pro- notum and elytra. Apterous female with a glabrous sulcus on the metapleura (Fig. ID); alate female with shallow sulcus in same position, not glabrous. Connexiva slightly raised in male, reflexed over abdomen in female. Dorsum of abdomen covered \Adth semilong decumbent setae, shorter in male. Venter uniformly dark, covered with semilong decumbent setae in female, much shorter in male. Femora yellowish basally and beneath, remainder of femora and tibia deep brown. Coxae, tro- chanters, and beak yellowish, antennae deep brown. Legs with the following proportions (holotype): Femur Tibia Tarsal 1 Tarsal 2 Anterior 39 27 14 — Middle 40 33 13 13 Posterior 42 42 10 13 Antennal proportions I; II; III; IV: 12; 15; 14; 17. Body measurements: Mean length of 10 apterous $ $ : 1.77 mm (max. 1.80, min. 1.72). Mean width of 10 apterous $ $ : 0.68 mm (max. 0.70, min. 0.67). Mean length of 10 apterous 9 $ : 2.17 mm (max. 2.38, min. 2.05). Mean width of 10 apterous 9 9 : 0.78 mm (max. 0.83, min. 0.72). Holotype: Length 2.20 mm, width 0.83 mm. Genital segment and paramere of male as in Figure IE and F respectively. MATERIAL EXAMINED. — Holotype, apterous 9 , Rincon Chamula. Chis., Mexico, 6-1-1974, CL671, J. T. Polhemus (in Polhemus Collection). Paratypes as follows; MEXICO: Chiapas,- 21 $ $ , 17 9 9 apterous, Rincon Chamula, CL671, 6-1-1974, J. T. Polhemus (JTP); 13 $$, 13 9 9 apterous, 2 $$.3 9 9 macropterous, 1 nymph, 12 mi. N Bochil. CL1091, 4 May 1964, J. T. and M. S. Polhemus (CU). Discussion. — This is the only member of the group in which the apterous female has an obvious glabrous sulcus on the meta- pleura. In general body shape, leavipleura resembles austrina, but the glabrous area on the posterior acetabula, lacking in austrina, will separate them. The collections of this species were made over a rather restricted area in northern Chiapas. Microvelia paura, n. sp. Female body shape robust (Fig. 2A) ; male body shape similar to figure lA. Anterior lobe of pronotum with orange brown band extending 214 GREAT BASIN NATURALIST Vol. 34, No. 3 to lateral margins in females, evanescent laterally in males; epister- num orange brown. Dense area of bristly setae on each humerus of apterous females (Figs. 2A, B), absent in alate female; bristly setae sparsely set laterally on anterior pronotal lobe; males without long hairs on pronotum. Small glabrous spot, sometimes absent, on lateral margin of pronotum of apterous female (Figs. 2A, B) ; metapleura with a shallow transverse, faintly shining depression. Elongate glabrous area on pronotum in a shallow transverse sulcus cephalad of humeri, reaching lateral margin, and irregular glabrous spot on mesopleura of alate female. Connexiva slightly raised, sometimes almost vertical in female. Dorsum of abdomen covered with semilong decumbent setae, shorter in male. Venter uniformly dark except episternum; thickly set with minute black conical setae, sparser posteriorly, very noticeable on episternum. Femora yellowish, tibia yellow brown, knees em- browned in females; males slightly darker. Legs with following proportions (holotype) : Femur Tibia Tarsal 1 Tarsal 2 Anterior 25 19 10 — Middle 26 21 11 9 Posterior 30 30 7 9 Antennal proportions I; II; III; IV: 9; 10; 9; 12. Body measurements: Mean length of 7 apterous $ $ : 1.11 nun (max. 1.14, min. 1.07). Mean width of 7 apterous $ $ : 0.44 mm (max. 0.44, min. 0.43). Mean length of 10 apterous $ 9 : 1.63 mm (max. 1.72, min. 1.57). Mean width of 10 apterous 9 9 : 0.74 mm (max. 0.83, min. 0.68). Holotype: Length 1.58 mm, width 0.68 mm. Material examined. — Holotype, apterous 9 , Palenque, Chis., Mexico, 13-XII-1973, CL608. J. T. Polhemus (in Polhemus Collection). Paratvpes as follows; EL SALVADOR: 3 5 5,13 99 apterous, 11 5 5. 33 99 macropter- ous. La Majadita, CL1256, 12-XII-1970, J. T. Polhemus (JTP). MEXICO: Chia- pas- 5 $ $ , 10 9 9 apterous, 4 $ $, 2 9 9 macropterous, Palenque. CL608, 13- XII-1973, J. T. Polhemus (JTP); 1 9 apterous. Rincon Chamula. CL671. 6-1-1974, J. T. Polhemus (JTP); 1 9 apterous, 2 $ $, 2 9 9 macropterous. Rizo de Oro, CL1331, 14-M970, J. T. Polhemus (JTP); 2 9 9 apterous, Simojovel 22-VII- to 19-VIII-1958, J. A. Chermak (UCB). PANAMA: 3 $ $, 2 9 9 macropterous, Cerro Campana, 7-V-1973, P. D. Ashlock (JTP, KU). Discussion. — The female body shape in paura is quite robust, and the transverse orange brown band on the anterior lobe of the pronotum reaches the lateral margins or extends onto the propleura; these characteristics give the females a rather distinctive appearance. The males, on the other hand, are very difficult to separate from the other species. The dense field of minute black conical setae on the venter, the orange brown episterntim, and the long proximal segment of the middle tarsi are helpful. Paura occurs throughout Central America. Microvelia psilonota, n. sp. Female body shape robust, similar to paura (Fig. 2A); male body shape elongate, tapering caudad, similar to Figure 1 A. Sept. 1974 POLHEMUS: AQUATIC HEMIPTERA 215 Anterior lobe of proiiotum with uninterrupted light to dark orange brovvn band not reaching lateral margins, rarely extending laterally to behind middle of eye. Anterior lobe of pronotum and propleura of alate and apterous females with long stiff setae (Fig. 2D); apterous and alate males with shorter scattered setae, not prominent. Apterous and alate females with a shallow glabrous sulcus of somewhat variable shape on lateral margin of })ronotum (Fig. 2D) . Pronotal disc of apterous female broadly feebly depressed. Connexiva almost flat to feebl}' raised in males and most females, strongly raised and almost vertical in some females. Venter uniformly dark, except proepisternum light orange brown; jugum of head, inner portion of anterior coxae, proepister- num behind eyes, lateral margins of metapleura and abdominal ventrites above spiracles with a field of minute black conical setae. Legs yellowish, embrowned dorsally on tarsi, tibia, and distal part of femora. Acetabula brown, posterior acetabula with glabrous area. Trochanters, coxae, rostrum and ventral part of first three antennal segments yellowish to yellowish brown; remainder of antennae brown. Legs with following proportions (holotype) : Femora Tibia Tarsal 1 Tarsal 2 Anterior 23 18 11 — Middle 24 21 9 9 Posterior 25 27 5 8 Antennal proportions I; II; III; IV: 8; 10; 7; 12. Body measurements: Mean length of 10 apterous $ $ : 1.28 mm (max. 1.32, min. 1.21). Mean width of 10 apterous $ $ : 0.53 mm (max. 0.55, min. 0.52). Mean length of 10 apterous 9 ? : 1.70 mm (max. 1.77, min. 1.62). Mean width of 10 apterous 9 2 : 0.89 mm (max. 0.95. min. 0.80). Mean length of 10 alate $ $ : 1.55 mm (max. 1.62, min. 1.50). Mean width of 10 alate $ $ : 0.72 mm (max. 0.75. min. 0.70). Mean length of 10 alate 5 9 : 1.85 mm (max. 1.95, min. 1.72). Mean width of 10 alate ? 9 : 0.87 mm (max. 0.95, min. 0.83). Holotype: Length 1.62 mm, width 0.83 mm. Material examined.— Holotype, apterous 9 . 20 mi. SE Acatlan, Puebla, Mexico. 16-XII-1969. CL1243. J. T. Polhemus (in Polhemus Collection). Para- tvpes as follows; MEXICO: Puebla; 6 $ $ , 15 9 9 apterous. 13 $ $ . \5 9 9 macropterous, 20 mi. SE Acatlan. 16-XII-1969, CL1243, J. T. Polhemus (JTP); 27 $ $ . 3i 9 9.3 n\'mphs apterous. 4 $ $ , 2 99 macropterous. same place, 28-IV-1964, J. T. & M. S. Polhemus (CU, JTP); 1 $ macropterous, Izucar de Matamoros. 27-IV-1964. CL1050, J. T. & M. S. Polhemus (CU). Durango; 1 $ macropterous. La Palmita, 20-IV-1964; CL1018. J. T. & M. S. Polhemus (CU). Sinaloa; 2 9 9 macropterous, Santa Lucia. 20-IV-1964, CL1019, J. T. & M. S. Polhemus (CU, JTP). Discussion. — The robust body shape, nature of the glabrous pronotal sulcus, and setae ])attern make separation of psilonota females a simple matter, but the males are very difficult to separate. J he combination of body size, yellowish proepisternum with the field of tiny conical black setae, and subequal middle tarsal segments are helpful, but for certain identification, associated females are desirable. 216 GREAT BASIN NATURALIST Vol. 34, No. 3 Microvelia reflexus, n. sp. Female body shape elongate, similar to Figure IB. Male body shape elongate, similar to Figure lA, but with pronotum narrower than metanotum and lateral margins sinuate. Anterior lobe of pronotum with orange brown band extending laterally onto episternum in both sexes. Sparsely set areas of long setae on anterior lobe of pronotum, humeri, and metanotum of ajiterous female (Fig. 2F) ; anterior lobe of pronotum and corium of hemelytra with similar setae in alate female; apterous and alate males with much shorter pubescence. Lateral pronotal margin of apterous female with a depressed area, feebly shining but not gla- brous (Fig. IF), absent in alate female. Connexiva raised in male, but not vertical; vertical in female, abruptly reflexed over abdominal tergite 4 then diverging posteriorly and less strongly reflexed on segments 5 and 6, strongl}' hirsute on segment 6. Abdominal dorsum clothed with very fine pubescence; female with long decumbent setae on last abdominal tergite. Venter uniformly dark, except episternum and genital segments orange brown; episternum thickly set with minute conical black setae and a few long setae. Male with last ventrite depressed, form- ing a broad transverse sulcus. Legs yellowish, embrowned dorsally on tibia, tarsi, and distal part of femora. Coxae, trochanters, beak, and ventral part of first antennal segment yellowish; remainder of antennae brown. Legs with following proportions (holotype): Anterior Middle Posterior Antennal proportions I; 11; III; IV: 10; 10; 10; 14. Body measurements: Apterous male, length 1.95 mm, width 0.78 mm. Macropterous male, length 2.14 mm, width 0.88 mm. Apterous female (holotype), length 2.15 mm, width 0.88 mm. Macropterous female, length 2.22 mm, width 0.97 mm. Material examined. — Holotype, apterous 9 , San Vito de Java, Costa Rica, CL1287, 30-XII-1968, J. T. Polhemus (in Polhemus Collection). Paratypes, same data as holotype, 2 $ $ apterous, 4 5 $, 2 ? $ macropterous. Discussion. — The relatively larger size and the transverse orange brown band on the anterior lobe of the pronotum, extending onto the episternum, set reflexus apart from other members of the group. Literature Cited Drake, C. J., and R. H. Hussey. 1955. Concerning the genus Microvelia Westwood, with descriptions of two new species and a checklist of the American forms (Hemiptera: Veliidae). Florida Ent. 28(3) :95-l 15. EsAKi, T. 1937. Die Gcrroidea Mikronesiens (Hemiptera-Heteroptera). Ten- thredo 1:351-362, 3 pis. ?mur Tibia Tarsal 1 Tarsal 2 29 20 12 — 32 27 12 12 35 38 7 10 Sept. 1974 POLHEMUS: AQUATIC HEMIPTERA 217 LiTTLEJOHN, M. J. 1969. The systematic significance of isolating mechanisms. Proc. Int. Conf. Syst. Biol.. Univ. Michigan, Nat. Acad. Sci. Pub. 1692: 459-482. PoLHEMUS, J. T., AND J. L. Herring. 1970. Ergebnisse der Osterreichischen Neukaledonien Expedition. Aquatic and Semiaquatic Hemiptera. Proc. Ent. Soc. Wash. 72 (2): 179- 187. ToRRE-BuENo, J. R. DE LA. 1924. A preliminary survey of the species of Microvelia Westwood (Veliidae. Heteroptera) of the Western World, with description of a new species from the southern United States. Bull. Brooklyn Ent. Soc. 19(5): 187-194. ADDITIONAL RECORDS OF COLEOPTERA COLLECTED AT THE NEVADA TEST SITE, MERCURY, NEVADA Vasco M. Tanner^ and Wilmer W. Tanner^ Abstract. — Thirty-nine species of Coleoptera not previously reported for the Nevada Test Site are listed. Several reports dealing with the Coleoptera collected at the Nevada Test Site were based on collections taken from can pit-traps placed in various sites throughout the area (Allred, Beck, and Jor- gensen, 1963; Allred and Beck, 1965; Tanner and Packham, 1965; and Tanner, 1963, 1966). Very few specimens have been reported other than those taken from traps. This has resulted in the sampling of insects that move about on the ground. There are many forms, however, that rarely would be taken by such methods. From 1965 to 1969 the following list of Coleoptera were taken by means of insect nets and from the vegetation of the Test Site area. Only species not reported previously are contained in this list. From the experience gained in our collecting, we are quite certain that many interesting species from this desert area have not yet been reported. Since the area is still being studied, this list may be of interest to workers in the area. The present study was a part of the ecological research done under the Atomic Energy Commission Contract No. At( 11-1) 1946 awarded to the junior author and Brigham Young University. Most of the specimens were collected by the authors; however, J. M. Hopkin and R. Walker aided in the field work. Family Carabidae Rhadine jejuna Lee. 20 specimens, Mercury, N.T. Site, July 1967. An abundant species around Cane Springs. Rhadine myrmecodes Horn 1 specimen, Pahute Mesa, 27-VII-67. Family Melyridae Collops punctulatus utahensis Schffr. 2 specimens, Pahute Mesa, 27-VII-67. Trichochrous varius Csy. 5 specimens. Rainier Mesa. July 1967. Eutrichopleurus concinnus Blais. 1 specimen, Nevada Test Site, 1969. Attalus futilis Fall 1 specimen, 3 miles W area 12, Nevada Test Site, 29-VII-67. Family Buprestidae Acmaeodera sp. 1 specimen. Frenchman Flat, on Mesquite bush. 29-Vni-67. 'Department of Zoology, Brigham Voumr University, Pinvo. Utah 84602. 218 Sept. 1974 TANNER & TANNER: COLEOPTERA 219 Family Cucujidae Oryzaephilus surinamensis L. 3 specimens, lO-X-66. Family Phalacridae Phalacrus sp. 25 specimens, Paliute Mesa, 27-VII-67. 5 specimens, Rainier Mesa, 29-VII-67. Familj' Coccinellidae Hippndamia apicalis Csy. 3 specimens. Pahute Mesa, 29-VII-67. Hippodamia convergens Guer. 4 specimens. Pahute Mesa, 27-VII-67. Hippodamia ambigua Lee. 1 specimen, Pahute Mesa. 27-VII-67. Hippodamia convergens uteana Csy. 1 specimen. Rainier Mesa. 28-VII-67. Hyperaspis quadrivittata Lee. 1 specimen, Pahute Mesa. 27-Vn-67. 1 specimen Frenchman Flat. 29-Vn-67. Hyperaspis taeniata tacniata Lee. 2 $ 1 Rainier Mesa and 13 miles W area 12, 27-VII-67. Hyperaspis taeniata nevadica Csy. 1 specimen, Frenchman Flat, 29-Vn-67. Hyperaspis pleuralis Csy. 1 specimen, 3 miles W area 12, 27-Vn-67. Scymnus pollens Lee. 1 specimen. Rainier Mesa, 28-Vn-67. Scymnus aridus Csy. 3 specimens. Pahute Mesa, 27-Vn-67. Family Alleculidae Hymenorus prolixus Csy. 3 specimens, Nevada Test Site. 22-Vn-65. Family Tenebrionidae Zopherodes uteanus Csy. 6 specimens. Rainier Mesa, lO-IX-69. Eleodes cognata Hald. 2 specimens, Nevada Test Site. 1965. Eleodes concinna Blais. 7 specimens. Rainier Mesa, 29-Vn-69. Eleodes nevadensis Blais. 2 specimens. Nevada Test Site. 27-VI-65. Eleodes pimelioides subsp. pairuelis Blais. 2 specimens. Rainier Mesa. Aug. 1968. Eleodes carbonaria subsp. interstitialis Blais. 5 specimens, Nevada Test Site, 3 IX-65. Eleodes omissa subsp. pygmaea Blais. 1 specimen. Pahute Mesa. 27-Vn-69. Trogloderus tuberculatus Blais. 2 specimens. Rainier Mesa. 8-IX-67. Eusattus oblongulus Csy. 4 specimens. Nevada Test Site, collected in the Grayia-Lycium community, 5-V-63. 220 GREAT BASIN NATURALIST Vol. 34, No. 3 Family Cerainbycidae Prionus californicus subsp. 1 specimen, Rainier Mesa, 31-VII-67. Moneilema gigas Lee. 1 specimen. Rainier Mesa, 31-VII-67. Family Chrysomelidae Chlamys memnonia Lac. 22 specimens. Frenchman Flat, 29-VII-67, abundant on Larrea divaricata. Family Curculionidae Crocidema californica Van Dyke 23 specimens, can trap, Rainier Mesa, 10 to 15-IX-67. 1 specimen. Rainier Mesa, 1969. 1 specimen. Rainier Mesa, 1970. Anthonomus ochreopilosus Dietz 5 specimens, area 12, 27 to 29-VII-67. Ceutorhynchus tescorum Fall 2 specimens, near Pahute Mesa, 1969. Anthonomus near juniperilius Dietz Pahute Mesa, 1967. Anthonomus inermis Boh. 2 specimens, Nevada Test Site, 1969. Tychius (Sibinia) setosus Lee. 1 specimen, on mesquite, Nevada Test Site, 1969. Cleonus quadrilineatus (Chev.) Nevada Test Site, 1971. Literature Cited Allred, D. M., D E. Beck, and C. D. Jorgensen. 1963. Biotic communities of the Nevada Test Site. Brigham Young Univ. Sci. Bull. BioL Ser. 2 (2): 1-52. Allred, D. M., and Beck, D E. 1965. A list of Scarabaeidae beetles of the Nevada Test Site. Great Basin Nat. 25:77-79. Tanner, V. M. 1963. A new species of Craniotus (Coleoptera: Temebrionidae) Great Basin Nat. 23:167-170. Tanner. V. M. 1966. Rhynchophora beetles of the Nevada Test Site. Brig- ham Young Univ. Sci. Bull., Biol. Ser. 8(2): 1-34. Tanner, V. M., and W. A. Packam. 1965. Tenebrionidae beetles of the Nevada Test Site. Brigham Young Univ. Sci. Bull. Biol. Ser. 6(1): 1-44. A SUMMARY OF BIOLOGICAL INVESTIGATIONS CONCERNING THE GREAT SALT LAKE, UTAH (1861-1973) Doyle W. Stephens^ Abstract. — The early stages in the historj- of biological investigation of the Great Salt Lake involved the identification and establishment of taxonomic relationships of the indigenous flora and fauna. A result of advancements in systematic biology is that many of the earlier names of organisms have been placed in synonomy. Recent interest in the lake has centered on biological pro- ductivity and interactions of components of the ecosystem. The creation of two ecologically distinct lakes bv the construction of a railroad causeway has further enhanced the biological complexity' of what was originally believed to be a lifeless bodj' of brine. Because of the late settlement of the Bonneville Basin, scientific investigation into the biological composition and biotic mechanisms of the Great Salt Lake was nonexistent before the latter half of the nineteenth century. An 1861 issue of Scientific American reported, "No living thing of any kind exists in the lake," ignoring Captain B. L. E. Bonneville's note of small animals in the water during his 1831-1833 explorations. By 1889 three species of algae (Farlow, 1879, cited in Kirkpatrick, 1934), a brine fly (Packard, 1871), and brine shrimp (Verrill, 1869) had been named from the lake, yet Jordan (1889) stated that no life could exist in the lake with the exception of brine shrimp. Schwarz (1891) investigated various forms of insect life adjacent to the lake and concluded that the brine fly, Ephydra cinerea Jones (as Ephydra gracilis Packard), was the only insect inhabitant of the lake. He made note of the adult flies' habits regarding oviposition and feeding in the water. Tilden (1898) reported five species of algae from the lake: Aphanothece Utahensis Tilden, Polycystis packardii Farlow, Dichothrix utahensis Tilden, Enteromorpha tuhulosa (Kiitzing) Reinbold, and Chara contraria Braun. It was now evident that the Great Salt Lake could support life and that additional biological inquiry was needed. Considerable in- terest and speculation centered around the introduction of marine organisms to the estuaries formed where fresh water entered the lake. Moore (1899) examined the chemical and physical character- istics of the lake and concluded that even with dilution, the waters would not support anv introduced crustaceans or fish. The possi- bilitv of introducing oysters into the estuaries was considered, but he concluded that a self-replenishing colony could not exist from year to year, and commercial exploitation was not feasible. Aldrich (1912) reported on the morphology and ecology of the brine flies E^phydra cinerea and Hydropyrus (as E.) hians (Say) from the lake, stating that a pulpy alga of the Nostoc group was the probable food of the Ephydra larvae. In his collection of notes on 'Department of Biology-, University of Utah. Salt Lake City, Utah 84112. 221 222 GREAT BASIN NATURALIST Vol. 34, No. 3 fauna in the lake, Vorhies (1917) stated that this Nostoc form was probably the alga Alphanothece packardii Setchell. He also com- mented on the viability of Artemia and Ephydra in various densi- ties of lake water. Vorhies noted several protozoans in his culture flasks including an amoeba (similar to Amoeba flowersi Jones), a ciliate protozoan (similar to Uroleptus)^ and a species of Euglena. The alga Chlamydomonas (reported in Daines, 1917) appeared regularly and in great numbers in his cultures. He noted that the brine shrimp was never collected from the lake when the water temperature was below 9C Complete absence of predators was suggested as an explanation for the great abundance of shrimp and brine flies. In repudiation of Vorhies's (1917) comment that brine shrimp and brine flies were abundant due to lack of predators, Wetmore (1917) noted that a wide variety of waterfowl fed heavily on the animals in the lake. He stated massive production of offspring to be the most likely explanation for the abundance of Artemia and Ephydra. A companion paper on the flora by Daines (1917, cited in error as Daniels) appeared with the observations of Vorhies on the fauna. Daines briefly mentioned Tilden's (1898) description of six algae and added a new one, Chlamydomonas sp. to the list. Two genera of diatoms {Navicula and Cymbella) were observed in the estuaries around the lake, and Daines concluded that they were adapted to dilute brines. He noted five bacteria, three being chromogenic, but offered no identifications. Daines noted a considerable size variation between the Chlamydomonas cells but through his experimentation concluded that it was not induced by differences in salinity. The brine shrimp, Artemia gracilis Verrill was further observed and its general morphology described in some detail by Jensen (1918). He reported the optimum density for hatching and growth of Artemia to be between specific gravities of 1.044 and 1.089. The eggs would not hatch in a saturated brine solution. Pack (1919) described a new species of protozoan, Prorodon utahensis Pack and studied the effects of brine dilution upon this species and another ciliate, Uroleptus packii Calkins. In less dense media, the animals increased in size, became more active, and de- veloped more flexible and contractile bodies. Pack also believed that by "slo\\ing down the rate of dilution, some of these Great Salt Lake forms may be transformed into fresh water animals." Seville Flowers (1934), in his monograph on the vegetation of the Salt Lake area, reported the following algae as endogenous to the lake: Aphanothece utahensis Tilden Microcystis packardii Farlow (Tilden) Oscillatoria tenuis var. tergestina (Kiitzing) Oscillatoria tenuis var. natans (Kiitzing) Chlamydomonas sp. Tetraspora lubrica var. lacunosa Chauv. The work of Flow'ers was followed by that of Kirkpatrick (1934) on the algal forms within the lake. Her conclusions touched upon Sept. 1974 STEPHENS: GREAT SALT LAKE BIOLOGY 223 the variety of organisms reported by earlier workers and are repro- duced with added comments by this author: 1 . There are five colonial forms of blue-green algae of the Great Salt Lake. Most of these forms did not thrive under laboratory conditions. 2. There are two species of Chlamydomonas in the Great Salt Lake. This form thrived at all densities (1.0145 to saturation). 3. Two species of diatoms, resembling Navicula, are present. They did not thrive in the weakest (sp. grav. 1.0145) nor the strongest (saturated) con- centrations present. 4. A species of Chroococcus and one of Gleocapsa developed in the lower con- centrations (sp. grav. 1.0225). They cannot be considered native lake forms. 5. A filament of Oscillatoria was seen twice. It is possible that this foiTn exists in small quantities in the lake. It is very abundant in the hot springs along the shore and could readily be washed into the main body of water. 6. A great number of the species listed by former workers did not develop in the cultures, nor were they observed in examination of fresh material. It is possible that many of these fonns came from extraneous sources, or were not able to survive the increased density of the lake water at its present low level. 7. The fauna observed in this experiment consists of three ciliates {Uroleptus packii Calkins, Prorodon utahensis Pack, and an unidentified species), one amoeba, one crustacean (Artemia) and one fly larvae (Ephydra). The reported occurrence of diatoms by Daines (1917) and Kirk- patrick (1934) was further investigated by Ruth Patrick (1936). She found a variety of diatoms in the sediments of the lake, presum- ably originating from the Lake Bonneville era, but did not find any evidence of their currently living in the Great Salt Lake. The bacterial composition within the lake was first investigated by Frederick (1924). Through the use of colonial morphology and several media, she isolated eleven forms, which she identified as the following: Serratia salinaria (Harrison and Kennedy) Bergey Cellulomonas subcreta (McBeth and Scales) Bergey Bacillus freudenreichii (Miguel) Chester Achromobacter solitarium (Ravenel) Bergey Bacillus cohaerens Meyer and Gottheil Flavobacterium arborescens (Frankland and Fiankland) Bergey Micrococcus sulflavus Chester Achromobacter hartlebii (Jensen) Bergey Bacteriodes rigidus (Dista) Bergey Bacillus mycoides Fliigge Achromobacter album (Pagliani) Bergey A brief mention of the brine shrimp, Artemia fertilis Verrill {Artemia salina Leach), larval Ephldra. the blue-green alga, Apha- nothecc utahensis as Aphanothicd packardii. diatoms, and the green alga Chlamydomonas is made by AUee (1926). An observation of water bugs of the family Corixidae is also noted, probably referring to a similar observation by Schwarz (1891). Woodbury (1936) provided the first comprehensive analysis of the lake ecosystem. His description of the aquatic system is brief, mentioning the apparent interspecific competition between two closely related species of Ephydra and Artemia. Several historical notes made by Fremont and Stansbury during early surveys were cited. 224 GREAT BASIN NATURALIST Vol. 34, No. 3 The bacterial composition within the lake was further investi- gated by Smith (1936). The majority of his results and conclusions are published in Smith and ZoBell (1937). Their conclusions, while not definitive, are of interest and are reproduced here: The attachment of bacteria to sterile glass slides submerged in Great Salt Lake indicates the presence of abundant and varied bacterial flora. Controlled experiments demonstrate that only living bacteria attach themselves to slides in appreciable numbers. This, together with the fact that micro-colonies develop on slides in the lake, indicates that the bacteria are multiplying in the lake and are not merely passive inhabitants. The inability of soil, sewage or marine bacteria to attach to slides in lake water supplies further proof for the latter contention. Most of the lake bacteria are small gram-negative rods besides other morphological varieties which do not fit into any conventional classification. The direct micro- scopic procedure offers possibilities for studying tlie seasonal and geographic distribution of bacteria in the lake. A summary of research on the brine shrimp, Artemia, prior to 1936 is given in Relyea (1937), but no new data are presented. An excellent compilation of research on the physiology of Artemia salina Leach appeared in the work of Quinn (1940). Magnesium ion concentrations of twice that in the lake did not appreciably change the time of nauplius emergence from the egg but did inversely affect the time of egg hatching. The time of emer- gence was found to vary inversely with the temperature, and effects of magnesium ion concentration on nauplii were restricted to the earliest developmental stages. A complete bibliography including many European articles is given. A short paper presented by Rees (1942) presents a popular view of animal life within the lake. It is followed by an article by Behle (1942) listing four species of colonial nesting birds found on islands in the Great Salt Lake. Distribution and general ecology is presented for: American white pelican {Pelecanus erythrorhynchos Gmelin), double-crested cormorant {Phalacrocorax auritus auritus Lesson), Treganza great blue heron (Ardea herodias Treganza), and the California gull {Larus calif or nicus Lawrence) . The amoeba noted by previous workers (Vorhies, 1917; Kirk- patrick, 1934; Woodbury, 1936) was described as Amoeba flower si Jones by Jones (1944). He also described the Euglena seen by Vorhies (1917) and Kirkpatrick (1934), naming it Euglena cham- berlini Jones. Woodbury (1948) briefly mentioned the work of Quinn (1940) concerning salinity effects on Artemia. No new data are given. The Ephydridae of Utah (Jorgensen, 1956) lists Ephydra cinerea Jones as the most common brine fly from the lake, with E. auripes Aldrich, E. riparia Fallen (as E. subopaca) and Hydro- pyrus (as E.) hians (Say) also being reported by various other col- lectors. Evans and Thompson (1964) list a new genus of ciliate proto- zoan, Pseudocohnilembus, occurring in the lake. Only one species, P. persalinus Evans and Thompson, was collected. Further work by Professor Evans and his students has resulted in the isolation of Sept. 1974 STEPHENS: GREAT SALT LAKE BIOLOGY 225 an additional ciliate, Euplotes parsalinus Reddy from the lake (Reddy, 1971). Population pressures and its problems were becoming evident when McDonald (1956) investigated the effects of pollution upon lake organisms. Dissolved oxygen concentrations were found to be below 3.3 mg/liter with localized decomposition of brine organisms being primarily responsible for the oxygen demand. He reported that an experimental mixture of 2 percent commercial oil and gaso- line added to lake water containing the alga Chlamydomonas re- sulted in complete elimination of cells within 45 hours. A comprehensive study of the bird life associated with the lake was authored by Behle (1958). As the lake lies within the Pacific Flywa}', there are numerous local and transient birds associated with its marshes, constituting the major predatory source for the macrofauna. Evans (1960) listed five new genera of protozoa and three un- identified types from the lake. Crystigera, Cyclidium, Euplotes, and Oikomonas were believed to be bacterial feeders. Podophyra was predareous upon Euplotes. An amoeba and two unidentified ciliates were also observed. Cristigera exhibited optimum growth in salt concentrations of 1 to 18 percent; its growth completely declined at 23 percent salt. Cysts of Cristigera, however, could survive long periods in a saturated salt solution. Preliminary tests on other pro- toza indicated that growth is inhibited at 15-18 percent salt con- centration. Evans concluded that Cristigera and the amoeba were specialized halophilic protozoa and that certain of the other species of protozoa may be salt-tolerant, freshwater forms. The occurrence of algal biostromes or tufa precipitated from the brine as a result of the action of blue-green algae was mentioned by Flowers (1934). Carozzi (1962) reported Aphanothece packardii to be the most predominant blue-green, forming the biostromes in distinct morphological zones. He concluded that the algae have no characteristic grow'th pattern of their own, but have developed on raised areas separating a system of erosional channels extending at right angles to the shoreline. A fairly complete summar}^ of plant and animal species found in and around the Great Salt Lake appeared in Flowers and Evans (1966). Their work lists two species of blue-green algae, Coccoch- loris elahens Drouet and Daily and Entophysalis rivularis (Kiitzing) Drouet, and two undescribed species of green algae, Chlamydomonas, as inhabiting the lake proper. The listing of bacteria follows that of Frederick (1924) with several forms listed in synonomy. The brine shrimp, Artemia salina is mentioned as the most conspicuous animal. The brine flies Ephydra cinerea Jones and E. hians Say are the only insects reported within the lake. The list of protozoa appearing in Evans (1960) was revised and expanded, listing the following ciliates: Urolepius packii Calkins Chilophyra utahensis (Pack) Podophyra sp. 226 GREAT BASIN NATURALIST Vol. 34. No 3 Euplotes sp. Pseudocohnilembus sp. Colhurnia sp. Two unidentified amoeba were noted as common, and several species of flagellates including Tetramitus, Oikomonas and at least two others were seen in large numbers from the lake and nearby salt ponds. Mention is made concerning the deposition of carbonate tufa by blue-greens, but the exact mechanism is unknown. Vegetation surrounding the lake is well described and its distribution noted. Gaskill (1970) reported on waterfowl commonly associated with the southeastern shore of the Great Salt Lake concluding that coots were the most prevalent of nesting birds (39 percent of total), with cinnamon teal, redhead, mallard, and pintails of considerable im- portance. The report of a National Science Foundation student-originated studies program (Carter, 1971) considered ecological relationships within the Farmington Bay Estuary of the Great Salt Lake, and the general terrestrial ecology of Antelope Island State Park. Por- tions of the aquatic study are relevant to the lake biology and are presented here. The estuary is less polluted now (1971) by coliform bacteria than it was in 1965. The coliforms are more heavily distributed on the estuary bottom than in the upper layers of water. Most coliforms are killed or fail to multiply in NaCl concentrations greater than 5.5 percent, with some of tlie bacteria being sensitive to concentrations of as little as 1.8 percent. The freshening of Farming- ton Bay could cause a definite increase in the coliform population. There are large numbers and many species of protozoans living in the estuary resulting from freshening of the lake due to construction of the causeway from Syracuse to Antelope Island. Because of the increase in the protozoan population, it is reasonable to e.xpect an increase in the overall biological pro- ductivity as protozoans are an important food and energy source. The distribution, number, and species diversity- of zooplankton and phyto- plankton were established. Through comparison with the water chemistry of samples taken at the same locations, it was found that the distributions of Arte- mia salina, Diaptomus, sp., a Corixid, Daphnia .sp.. and Nodularia sp. are depen- dent on the salinity. A predator-prey relationship between the Corixid and Arte- mia salnia was suggested, and it is concluded that the introduction of marine game fish or fresh water fish to the area for sport fishing is not feasible. The construction of a rock-filled railroad causeway between Little Mountain and Lakeside in 1957 resulted in the creation of two ecologically distinct lakes due to salinity imbalances. Its effect on the biota was reported by Gillespie, Wirick, and Stephens (1971). They concluded that the saline waters of the Great Salt Lake pro- vided an extremely rigorous, and therefore relatively simple eco- system. The northern basin contains saturated brine with a depau- perate biota consisting of Dunaliella salina Teodoresco plus unidenti- fied protozoa and bacteria. In the southern basin, two major energy- flow sequences dominate the system: a planktonic sequence con- sisting of {Dunaliella) -> (Artemia) and a benthic sequence con- sisting of (blue-green algae -f- detritus) -> {Ephydra) . There is some crossover in that much of the detritus consists of dead Artemia, and Artemia will feed on benthic algae and detritus when Duna- liella are scarce. Sept. 1974 STEPHENS: GREAT SALT LAKE BIOLOGY 227 Further work by Wirick (1972) demonstrated that the main phytoplanktor. DuiwlicUa viridis Teodoresco (previously reported as a Chlamydomonas) exhibited one bloom per year in April. The zooplankter, Artcmia salina is present and grazing DunalicUa only when the water temi)erature is above 6C. Construction of a mathematical simulation model suggested that the growth rate of th(^ DunalicUa population is light limited and density dependent at high algal concentrations. Ponella and Holnian (1972") concluded that inorganic nitrogen is apparently the limiting factor for growth of phytoplankton in the Great Salt Lake water. Carbon may also be limiting. Phosphorus, iron, and other trace elements seem to be in abundant supply. Their observations were confirmed by algal bioassays. Growth and repro- duction of the brine shrimp on Dunaliella alone was superior to yeast alone as a food source. The optimum utilization by the brine shrimp was about 1,000 algal cells per brine shrimp per day. Dif- ferent concentrations and ages of added algae had no apparent effect on whether the mature brine shrimp produced live young (nauplii) or resistant cysts. It was their belief that a feasible aquacmture based ouDunaliella sp. and Artemia sp. could be developed for brine shrimp isolated from the Great Salt Lake. Production of algae and brine shrimp in lake enclosures may be increased by addition of specific nutrients. Basic schemes for energy flow within the north and south lake basins were presented by Stephens and Gillespie (1972). They found that the northern basin supports a depauperate biota consisting primarily of an alga, Dunaliella salina, several protozoa, and bac- teria. The southern basin exhibits two energy- flow systems with only minor interactions: the planktonic system with a dominant phytoplanktor, {Dunaliella viridis), and a single zooplankter, (Arte- mia salina) ; and a benthic svstem of blue-green alga {Coccochloris elahens), detritis, and brine fly larvae (Ephydra) . The only outflow from either system occurs when birds feed upon the shrimp or fly larvae. The Dunaliella population seems to be limited early in the calendar year by temperature and light. Dunaliella viridis reaches its peak population density (24 x lO'Vliter) in April and its decline to less than 1 x 10'^ cells/liter) occurs in May and June as a conse- quence of the rapidly expanding Artemia salina population. The availability of the nutrients nitrogen and phosphorous does not seem to be a limiting factor for Dunaliella. The apparent conflict of the Porcella-Holman study (1972) and that of Stephens-Gillespie (1972) regarding limiting factors to phy- toplankton growth is currently under investigation by Stephens (1975). Initial conclusions indicate that Dunaliella is (1) light limited during the April-May bloom and (2) nitrogen, carbon, and possibly vitamin limited later in the year. Grazing by Artemia could prevent additional algal blooms even if necessary nutrients were available. Most recently. Van Auken and McNulty (1973) published on the factors limiting growth in laboratory cultures of Dunaliella sp. 228 GREAT BASIN NATURALIST Vol. 34, No. 3 isolated from the Great Salt Lake. Optimum growth was obtained under the following conditions: (1) temperature 32 C, (2) NaCl 19.2 percent (w/v), (3) CO^ 1-2 percent at a rate of 2.2 ml/min/ml of culture media, (4) light intensity of 25-35 klux, (5) pH 5.8-6.5. The K^/Na"^ ratio should not be more than 0.1. The specific growth constant for this halophyte under the above conditions was 0.069 hrs~\ which is equal to a doubling time of 10 hours. Chemical control of the massive swarms of Ephydra in the beach areas was reported by Nabrotzky, Rosay, and Sadler (1973). Control lasting several hours to several da3^s was obtained using both mala- thion and Dowco 214 insecticides. At the concentrations applied, no damage to Artemia or water bugs (Corixidae) was evident. An indigenous wasp parasite of Ephydra larvae collected near the lake indicates biological control of the brine flies may be possible. Literature Cited Aldrich, J. M. 1912. The biology of some western species of the dipterous genus Ephydra. J. N.Y. Ent. Soc. 20:77-102. Allee, W. C. 1926. Some interesting animal communities of northern Utah. Sci. Mon. 23:481-495. Behle, W. H. 1942. The colonial nesting birds of Great Salt Lake. Min. Soc. Ut. News Bull. 3:59-63. . 1958. The Bird Life of the Great Salt Lake. Univ. Utah Press, Salt Lake City. 230 pp. Carozzi, a. V. 1962. Observations of algal biostromes in the Great Salt Lake. J. Geol. 70:246-252. Carter, C. K. 1971. Some ecological considerations of Farmington Bay and adjacent Great Salt Lake State Park. Report NSF Grant #GY-9120. Univ. Utah Press, Salt Lake City. 247 pp. Daines, L. L. 1917. [reported in error as Daniels] On the flora of Great Salt Lake. Am. Nat. 51:499-506. Evans, F. R. 1958. Culture of protozoa from Great Salt Lake. J. Protozool. 5:13. . 1960. Studies on growth of protozoa from the Great Salt Lake with special reference to Cristigera sp. J. Protozool. 7:14-15. Evans, F. R., and J. C. Thompson, Jr. 1964. Pseudocohnilembidae n. fam., a h3^menostome ciliate family containing one genus, Pseudocohnilembus n.g., with three new species. J. Protozool. 11:344-352. Flowers, S. 1934. Vegetation of the Great Salt Lake Region. Bot. Gaz. 95: 353-418. Flowers, S., and F. R. Evans. 1966. The flora and fauna of the Great Salt Lake region, Utah. Pages 367-393 in Hugo Boyko (ed.) Salinity and aridity: new approaches to old problems. Junk, the Hague. Frederick, E. 1924. On the bacterial flora of Great Salt Lake and the via- bility of other microorganisms in Great Salt Lake water. M.S. thesis, Univ. Utah. 65 pp. Gaskill, J. H. 1970. Waterfowl studies on the southeastern Great Salt Lake. M.S. thesis, Univ. Utah. 51 pp. Gillespie, D. M., C. D. Wirick, and D. W. Stephens. 1971. The Great Salt Lake: plankton ecology in a rigorous environment. Bull Ecol. So. Am. 52:31. Jensen, A. C. 1918. Some observations on Artemia gracilis, the brine shrimp of Great Salt Lake. Biol. Bull. 34:18-25. Jones. D. T. 1944. Two protozoans from Great Salt Lake. Bull. Univ. Utah. 35:1-11. Jordan, D. S. 1889. Report of exploration of Colorado and Utah during the summer of 1889. U.S. Comm. Fish. 9:31-68. Jorgensen. E. C. 1956. The Ephydridae of Utah. M.S. thesis, Univ. Utah. 62 pp. Soj)t. 1974 STEPHENS: GREAT SALT LAKE BIOLOGY 229 KiRKPATRiCK, R. 1934. The life of Great Salt Lake, with special reference to tlie algae. M.S. thesis. Univ. Utah. 30 pp. McDoN.\LD. D. B. 1956. The effects of pollution upon Great Salt Lake, Utah. M.S. thesis. Univ. Utah. 42 pp. Moore, H. F. 1899. An inquirj^ into the feasibility of introducing useful marine animals into the waters of Great Salt Lake. U.S. Comm. Fish. H. of Rep. Doc. 692, part 25:231-250. Nabrotzky. F. W., B. Ros.\y. and T. Sadler. 1973. Further studies on the ecology and control of brine flies of the Great Salt Lake. Proc. Ut. Mosq. Abate. Assoc. (Oct. 1-2) Salt Lake City. P.\ckard, a. S.. Jr. 1871. On insects inhabiting salt water. Am. J. Sci. 1:100-110. . 1879. The seaweeds of Great Salt Lake. Am. Nat. 13:701-703. P.ATRicK, R. 1936. Some diatoms of Great Salt Lake. Torrey Bot. Club Bull. 63:157-166. PoRCELLA, D. B.. AND J. A. HoLMAN. 1972. Nutrients, algal growth, and cul- ture of brine shrimp in the southern Great Salt Lake. Pages 142-155 in J. P. Riley (ed.) The Great Salt Lake and Utah's water resources. Proc. First Ann. Confer, of Utah Sec. Am. Water Res. Assoc. Utah Water Res. Lab.. Utah State Univ. Reddy, Y. J. R. 1971. A description of a new species of Euplotes from Great Salt Lake, Utah. M.S. thesis. Univ. Utah. 31 pp. Rees, D. M. 1942. Animals living in the Great Salt Lake. Min. Soc. Ut. News Bull. 3:57-58. Relye.'v, G. M. 1937. The brine shrimp of Great Salt Lake. Am. Nat. 71: 612-616. Scientific American. 1861. Great Salt Lake. 9:131-132. ScHWARz, E. A. 1891. Preliminary remarks on the insect fauna of the Great Salt Lake. Utah. Can. Ent 23:235-241. Smith, W. W. 1936. Evidence of a bacterial flora indigenous to the Great Salt Lake. M.S. thesis. Univ. Utah. 101 pp. Smith. W. W., and C. E. ZoBeel. 1937. Direct microscopic evidence of an autochthonous bacterial flora in Great Salt Lake. Ecology 18:453-458. Stephens, D. W. 1973. Primary production and the influence of limiting factors in the Great Salt Lake, Utah. Ph.D. dissertation, Univ. Utah (in preparation). Stephens, D. W. and D. M. Gillespie. 1972. Community structure and ecosystem analysis of the Great Salt Lake. Pages 66-72 in J. P. Riley (ed.) The Great Salt Lake and Utah's water resources. Proc. First Ann. Confer. of Utah Sec. Am. Water Res. Assoc. Utah Water Res. Lab., Utah State Univ. TiLDEN. J. 1898. American Algae. Cent. TIL No. 298. Van Auken, O. W.. and L B. McNulty. 1973. The effect nf environmental factors on the growth of a halophylic species of algae. Biol. Bull. 145:210-222. Verrill, a. E. 1869. Twelfth Annual Report of U.S. Geological & Geographi- cal Survey the Territories of Wyoming and Idaho, Part I. 1878. U.S. Govt. Print. Office, Washington. Vorhies, C. T. 1917. Notes on the fauna of the Great Salt Lake. Am. Nat. 61:494-499. Wet.more, a. 1917. On the fauna of Great Salt Lake. Am. Nat. 51:753-755. WiNOET, R. N.. D. M. Rees, and G. C. Collett. 1969. Preliminary investiga- tions of the brine flies in the Great Salt Lake. Utah. Proc. Utah Mosquito Abate. Assoc. 22:16-18. WiRicK, C. D. 1972. Dunaliella-Artemia plankton community of the Great Salt Lake. Utah. M.S. thesis, Univ. Utah. 44 pp. Woodbury. A. M. 1936. Animal relationships of Great Salt Lake. Ecology 17:1-8. . 1948. Animals and salinity in the Great Basin. Am. Nat. 82:171-187. A NEW VARIETY OF PENSTEMON THURBERI (SCROPHULARIACEAE) James L. Reveal' and Janice C. Beatley- The Thurber Beard-tongue, Penstemon thurberi Torr. is cur- rently known from the sandy desert regions of southwestern United States and northern Mexico ranging from the Sierra San Pedro Martir in Baja Cahfornia northward to the Providence Mountains of San Bernardino County, Cahfornia, hence eastward into southern Mohave and Yavapai counties, Arizona, and across the southern tier of Arizona counties into western New Mexico. The recent dis- covery of a northern, disjunct population of P. thurberi from ex- treme northwestern Clark County, Nevada, in the near vicinity of the Nevada Test Site, has now been studied in detail and has proved to represent a distinct variant of the species which we hereby name. Penstemon thurberi Torr. var. anestius Reveal & Beatley, var. nov. A var. thurberi floribus 8-9 mm longis (nee 10-15 mm longis) cum inferior labiis (1) 2-3 (3.5) mm longis (non 4-6 mm longis) et tubis (4) 6-7 mm longis (non 7-10 mm longis), capsulis 4-5 mm longis differt. Typus: NEVADA: Clark Co.: In deep volcanic sands on the upper bajada below the southwest end of the Buried Hills, east of Frenchman Flat dry lake, associated with Larrea and Am- brosia, at about 3800 feet elevation, 20 June 1973, Beatley <& Acker- man 13460. Holotypus, US! Isotypi, 30 duplicates will be distributed from US. Other Specimens Examined: All from the type area: 22 June 1971, Beatley & Bamberg 12843 (MARY, NTS, US) . The var. anestius (from the Greek anestios, meaning homeless, alluding to its disjunct distribution) differs from var. thurberi primarily in the features of the flowers, which are smaller and less obvious than the flowers of the typical form. In var. anestius the flowers are 8-9 mm in length; those of var. thurberi are (10) 12-15 mm. The lower lip of var. thurberi is up to twice the length of that of var. anestius, whereas the tube of the Nevada plants is shorter than that of var. thurberi. The mature capsules of var. anestius are 4-5 mm long; those of var. thurberi are 6-9 (10) mm. Both forms occur in similar habitats and have the same general habit of growth, although the Nevada plants rarely exceed 5 dm in height. These features, coupled with the disjunct distribution pattern, have led us to recognize the Nevada variant as distinct. 'Department of Botany, University of Maryland, College Park 20742, and National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Research supported by National Science Foundation Cirant GB-22645. -Department of Biological Sciences, University of Cincinnati, Cincinnati, Ohio 45221. The work reporlcd hero was (oriducted in part under U.S. Atomic Energy Commission contracts No. AT (04-1) Gen-12 and AT (11 -I) 2307. 230 A NEW SCELOPORINE LIZARD FROM OAXACA, MEXICO Marlene M. Dasmann' and Hobart M. Smiths Abstract. — Sceloporus megalepidurus halli, subsp. nov., is described from San Jose Lachiguiri, Oaxaca, Mexico, from one adult male. Among specimens recently collected for the University of Colo- rado Museum by the late Thomas B. MacDougall in a remote sector of the Sierra Madre del Sur of Oaxaca, Mexico, is a single specimen of a distinct, unnamed race of Sceloporus megalepidurus. Despite an appeal for search for more material before his death, he was un- able to secure it. The senior author attempted to reach the locality with directions provided by Mr. MacDougall only a week before his death but was unable to do so. Since additional material is not now likely to be acquired for at least several years, we here describe and name Sceloporus megalepidurus halli, subsp. nov. HoLOTYPE. — University of Colorado Museum 41137, adult male, collected by Thomas MacDougall in October 1967 at San Jose Lachiguiri, Oaxaca, Mexico. Diagnosis. — A Sceloporus megalepidurus of small size, maxi- mum snout-vent measurement 47 mm; dorsal scales 46 from occiput to base of tail; no ventral coloration. Description of holotype. — Head scales above (Fig. 1) smooth, weakly pitted on frontonasals and prefrontals; interparietal about 3 mm wide, broader posteriorly than anteriorly, posterior edge fairly straight; parietals divided in two, each about one-sixth size of inter- parietal; a moderately large scale posterior to parietal, considered to be a secondary parietal; a pair of moderate-sized rectangular fronto- parietals, separated medially by broad contact of frontal and inter- parietal; frontal transversely divided, posterior section three-fourths as large as anterior section; prefrontals moderate in size, narrowly contacting each other medially; median frontonasal somewhat larger than either lateral frontonasal; a pair of square scales in front of median frontonasals, preceded by another, similar pair of scales; four postrostrals; four enlarged supraoculars on each side, separated from median head scales by a complete row of small scales; one complete and another incomplete row of scales separating supraocu- lars from superciliaries; six superciliaries on each side, normal; one canthal on each side; subnasal present, about same size as loreal; preocular not divided; subocular long, single, followed posteriorly around margin of orbit by two postoculars; two incomplete rows of lorilabials (Fig. 2), reduced to one row below subocular; one row of 'Department of Environmental, Population and Organismic Biology. University of Colorado, Boulder, Colorado 80302. 231 232 GREAT BASIN NATURALIST Vol. 34, No. 3 Fig. 1 . Dorsal head scales of the holotype of S. m. halli. lorilabials continuous around end of snout; five supralabials to a point at posterior margin of eye. Mental pentagonal, with a labial border about half that of rostral; outer row of labiomentals separated from mental by narrow contact of first postmental and first infralabial; four pairs of well-differen- tiated postmentals, followed by several scales not well differentiated from adjoining gular scales; first pair of postmentals in contact Sept. 1974 DASMANN & SMITH: MEXICAN LIZARDS 233 Fig. 2. Lateral head scales of the holotype of S. m. halli. medially; gular scales all nearly equal in size, somewhat smaller than scales on chest, smallest below ear and between postmentals. Auricular lobules three-four; about six scales between auricular lobules and postoculars; temporal scales keeled, larger than scales between ear and lateral nuchal fold, smaller than largest auricular lobule; scales between ear and lateral nuchal fold keeled, mucronate. Dorsal scales 46 from occiput to base of tail, not reduced in size on nape, weakly keeled, weakly mucronate; lateral scales about one- third smaller than dorsals, not abruptly differentiated from them; lateral scales in oblique rows converging posterodorsally; most ven- tral scales with a single apical notch; scales in axilla and groin im- bricate, some notched; scales on chest slightly larger than mid ventral scales; preanal scales somewhat smaller than lateral abdominal scales, subequal in size to smallest midventral scales; dorsal scales on rump somewhat reduced; dorsal caudal scales at least twice as large as scales on rump; scales around middle of body 54. Dorsal scales of upper foreleg keeled, mucronate, equal in size to dorsal scales on body, slightly larger than largest dorsals on lower foreleg; scales on ventral surface of upper foreleg very small, keeled, weakly mucronate, those of lower foreleg larger, keeled, mucronate; lamellar formula for fingers 8-12-16-19-12 (9-11-16-18-12). Dorsal scales of shank keeled, mucronate, equal in size to median dorsal scales, those of thigh slightly smaller; scales on anterodorsal surface of shank near tibiometatarsal joint greatly reduced in size; ventral scales of shank smooth, smaller than dorsal scales of same member, decreasing in size on ventral surface near femoral pores; scales preceding femoral pores subequal to preanal scales; median scales on posterior surface of thigh keeled, mucronate, subequal to scales in preanal region, decreasing toward series of femoral pores; 234 GREAT BASIN NATURALIST Vol. 34, No. 3 femoral pores 13-14, the two series separated by four scales; no postfemoral dermal pocket; enlarged postanals present, broader than long, separated narrowly by two small scales; lamellar formula for toes 9-13-17-20-13 (10-14-16-20-13). Color and Pattern (Fig. 3). Dorsal surface brown gray; a broad, clove brown band from posterior margin of orbit to rump, bordered on each edge by a light line; below this, irregular spots of clove brown, absent toward ventral surface; black area on shoulder; limbs with narrow clove brown bands; posterior surface of thigh irregu- larly reticulated with clove brown; tail with narrow clove brown bands. Male with immaculate ventral surface; gular scales brown gray, appearing banded toward throat. Discussion. — The present specimen is placed in the megalepi- durus group of Smith (1939). Its only major physical distinction in comparison with Sceloporus pictus is its lack of ventral color. This characteristic is also found in Sceloporus megalepidurus, a close relative of S. pictus. However, the specimen has fewer scales from occiput to base of tail (46) than does S. megalepidurus (52-63). Indeed the new taxon probably will prove to have a lower mean dorsal scale count than even pictus, perhaps even diagnostically lower. Comparisons are given in Table 1, from our own counts. The differences between the present type and both megalepidurus and pictus are such that we believe its accorded rank should be subspecific despite the extensive isolation presently indicated. There is no particular resemblance to either S. subpictus or S. cryptus, the only other taxa of the megalepidurus group; each has distinct lateral belly patches, dorsals no more than 37 from occiput to base of tail, and other distinctions. If S. pictus were regarded as specifically distinct from S. mega- lepidurus, the taxon here described would most reasonably be re- garded as a subspecies of the former {S. pictus halli). However, Fig. 3. Dorsolateral view of the holotype of 5. m. halli. Sept. 1974 DASMANN & SMITH: MEXICAN LIZARDS 235 Table 1. Selected scale counts (means and ranges) in S. megalepidurus. Scales Scales between Sample Dorsal Ventral around Femoral femoral Name size scales scales midbody pores pores Sceloporus m. pictus 13 50.6 49.4 56.9 15.0 4.8 46-54 45-54 53-61 12-18 3-7 S. m. megalepidurus 45 56.7 56.2 57.4 14.7 5.7 52-63 45-69 46-68 13-17 4-7 Hybrids (pictus megalepidurus) 51 57.8 53.6 55.2 14.5 5.9 52-63 46-70 45-65 12-18 4-8 S. m. halli 1 46 50 54 13-14 4 recently Dr. William P. Hall discovered an apparent intergradation zone between the ranges of S. pictus and S. megalepidurus. The specimens collected from this area are quite similar to megalepidurus in nmnbers of dorsal scales, ventral scales, scales around the mid- belly, femoral pores, and scales between the femoral pore series. However, the males have distinct blue belly patches, much as in pictus.. although they are not as clearly defined as is typical of the latter race. Some of the hybrid males have thin black lines outlining these blue belly patches. We thus conclude that indeed megalepi- durus and pictus do intergrade and should be ranked as conspecific subspecies; accordingly our new taxon must fall as a subspecies of S. megalepidurus. Sceloporus m. pictus and S. m. megalepidurus probably evolved from the same ancestral race, and, due to geographic isolation, became phenotypically (and presumably genetically) distinct. The present zone of intergradation appears to be secondary, with reunion of the populations occurring after a number of differences between them had evolved. Because the specimens from this intergradation zone have the S. m. pictus ventral coloration and S. m. megalepi- durus scale counts, we assume that S. m. halli was not of a similar origin (i.e., not a result of interbreeding between pictus and megalepi- durus). It appears more likely that the new taxon is an offshoot of a common ancestral population. 5. m. halli occurs in southern Oaxaca far removed from the present ranges of 5. m. pictus or S. m. mega- lepidurus and far from the intergradation zone (Fig. 4). A founder population could conceivably have been displaced this far from its natural range by human agency, but it seems unlikely. Probably the range of the common ancestral population once maintained continuity from northern Oaxaca into this area, and has since con- tracted, leaving this population isolated to evolve on its own. Since all of the present subspecies of megalepidurus are so closely related, thev probably carry many of the same genes. It would appear that S. m. halli has paralleled some of the mutations or cornbinations phenotvpically expressed in 5. m. megalepidurus. becoming like it in being imrnaculate ventrally but otherwise remaining similar to S. m. pictus, which presumably is more like the ancestral population than is either of the peripheral subspecies. Unfortunately, only one 236 GREAT BASIN NATURALIST Vol. 34, No. 3 • m. megalepldurut • m. pictus :)e IT., megalepidurus x picfus O m. halll ..>v' >^ Fig. 4. Distribution of the subspecies of S. megalepidurus. specimen of S. m. halli is available; it is hoped that future collecting efforts will concentrate on the area where the holotype was found. The area around San Jose Lachiguiri, Distrito Miahuatlan, Oaxaca, is unusual in several ways other than serving as the habitat of S. m. halli. Unexpected taxa of the genera Phrynosoma and Barisia are also found here, all far removed from their close relatives. At one time the ranges of these and other species could have ex- tended in continuity to this area. Due to climatic change, or some other limiting factor, their ranges contracted, leaving relic popula- tions isolated in this area, where they became to some degree dif- ferentiated from their parental stock. Why this area appears to contain so many relictual populations is not yet understood. What- ever the reason, it constitutes an unusually distinctive faunal district. Specimens Examined. — Specimens have been examined in the Museum of Comparative Zoology (mcz). University of Colorado Sept. 1974 DASMANN & SMITH: MEXICAN LIZARDS 237 Museum (cum), the private collection of Edward H. Taylor (eht) (now in part in the University of Illinois Museum of Natural His- tory) and the private collection of Earl Olson (eo), as follows: S. m. megalepidurus - Veracruz: Mt. Orizaba (mcz 14157); Mai Paiz (cum 50383). Puebla: Lago Alchichica (cum 29111-21, 29123-39);4 mi. NE Entronque Zacatepec (mcz 133158-65); 32 km SE Perote (mcz 122162-3); 6 mi ESE San Salvador el Seco (mcz 133155-7). Mexico: Teotihuacan Valley (mcz 133166). S. m. pictus - Veracruz: Acultzingo (eht 7623, 7629 A) ; Cumbres de Acultzingo (cum 48372-6; EO 764, 1191). Puebla: near Alseseca (eht 7620A, 7620-2); 20 km N Tehuacan (eht 7624-5, 7625A, 7626-9); Tehuacan (mcz 42140-1); 15 km SSE Amozoc de Mota (mcz 121885-8). S. m. megalepidurus X S. m. pictus - Puebla: 5 km SE Ciudad Serdan (mcz 133123-54); 8 km SE Ciudad Serdan (mcz Y-25591, 122164-81). S. m. halli - Oaxaca: San Jose Lachiguiri (cum 41137). Acknowledgments. — We are especially indebted to Dr. Wil- liam P. Hall for calling our attention to the intergrade specimens and for permission to study them: the new taxon bears his name in recognition of these courtesies and of his numerous contributions to the systematics of this complex genus. In addition, we are grateful for the loan of comparative material by Dr. T. Paul Maslin, Dr. Ernest E. Williams, Dr. D. F. Hoffmeister, and Earl Olson, from CUM, MCZ, the University of Illinois Museum of Natural History, and the Earl Olson private collection, respectively. Finally, w^e are very grateful to Neal Burstein for the photographs. Literature Cited Smith, H. M. 1939. The Mexican and Central American lizards of the genus Sceloporus. Zool. Ser. Field Mus. Nat. Hist. 26:1-397, figs. 1-59, pis. 1-31. RANGE EXTENSION OF THE LONG-NOSED SNAKE, RHINOCHEILUS L. LECONTEl, IN 10 EAST-CENTRAL UTAH William L. Grogan, Jr.i and Wilmer W. Tanner- Abstract. — A review of the distribution of Rhinocheilus I. lecontei, in Utah, with a northward extension of its range in the Upper Colorado River Basin. The long-nosed snake, Rhinocheilus I. lecontei, was previously thought to inhabit only the southwestern part of Utah (Tanner, 1935. Its range in Utah included only the Lower Sonoran life zone in the Virgin River Valley of Washington County. Tanner (1941) reported it for the Great Rasin (Millard Co.), and Tanner and Heinrichs (1964) extended its range into southeastern Kane County. Stebbins (1966) reported a specimen from San Juan Coun- ty, Utah (northeast of Page, Arizona), extending its range into the Colorado Plateau of southeastern Utah. On 29 May 1970 at Dragerton, Carbon County, Utah, a specimen of Rhinocheilus I. lecontei was collected by William Ingram from under a rock. This specimen extends the range of Rhinocheilus well Fig. 1. Dorsal view of Rhinocheilus I. lecontei from Carbon County, Utah. 'Department of Entomology, University of Maryland, College Park, Maryland 20742. ^Department of Zoology, Brigham Young University, Prove, Utah 84602. 238 Sept. 1974 GROGAN & TANNER: UTAH SNAKE 239 into the Upper Colorado River Basin and is the first specimen taken in Utah east of the Wasatch mountains. It represents a range exten- sion of approximately 1 75 miles north of previous records for south- eastern Utah. Known locality records for Utah are plotted in Fig. 2. The above specimen, a female, from Carbon County (Fig. 1), is now at Brigham Young University, Provo, Utah (BYU 33306). It is an example of the lecontei phase of this species as defined by Shannon and Humphery (1963) and Tanner and Jorgensen (1963). The pattern in alcohol is 35 dark slate-gray saddles and 35 cream- Fig. 2. Locality records of Rhinocheilus I. lecontei in Utah The Carbon County specimen is represented by the sohd triangle. : solid circles). 240 GREAT BASIN NATURALIST Vol. 34, No. 3 colored saddles. The following scale counts were noted: ventrals 205; caudals 34; scales in 25-23-19 rows; upper labials 8; lower labials 9; preoculars 1; postoculars 2; loreals 1; prefrontals 1; inter- nasals 1; right temporals l-)-3+2; left temporals 2+3+2. The caudal number of 34 is apparently the lowest ever recorded for this species: Wright and Wright (1957) record a low of 41, and the lowest number reported by Tanner (1941) in specimens from Utah was 43. Additional specimens are needed from eastern Utah to determine the significance of this low ventral count. The senior author visited the Dragerton locality during August 1972 in an attempt to obtain additional specimens. Four days were spent in the area driving roads at night and turning rocks, without success. Few other reptiles were seen, perhaps because of the ex- treme dryness of the area and the time of the year. The significance of the Dragerton specimen is to again indicate the importance of the Upper Colorado Basin as a distribution route for many species. Western species {Sauromalus obesus, Phrynosoma platyrhinos, Xantusia vigilis, Lampropeltis getulus californiae, Tan- tilla planiceps, and others) and eastern species {Eumeces multivir- gatus gaigei, Lampropeltis triangulum taylori, and Elaple guttata) have entered the basin from the south and moved northward. Be- cause of the isolation factor for populations established in the Upper Basin, many distinct subspecies have evolved. Unfortunately, many populations are widespread and individuals are difficult to obtain, which leaves us with many unanswered systematic problems. Literature Cited Shannon, F. A., and F. L. Humphery. 1963. Analysis of color-pattern poly- morphism in the snake, Rhinocheilus lecontei. Herpetologica 19:153-160. Stebbins, R. C. 1966. A field guide to western reptiles and amphibians. Houghton Mifflin Co., Boston, 279 p. Tanner, V. M. 1935. Western Worm-Snake, Siagonodon humilis (B and G) found in Utah. Utah Acad. Sci., Arts and Letters 12:267-270. Tanner, W. W. 1941. A study of the variation in the less common snakes in Utah. Great Basin Nat. 2(1): 16-28. . 1964. An extension of Arizona e. philipi and Rhinocheilus I. lecontei into south-central Utah. Southwest Nat. 9(l):45-46. Tanner, W. W., and C. D. Jorgensen. 1963. Reptiles of the Nevada Test Site. Brigham Young Univ. Sci. Bull.. Biol. Ser. 3(3): 1-31.. Wright, A. H., and A. A. Wright. 1957. Handbook of snakes. Comstock Publ. Co., Ithaca, New York, 1105 p. DESCRIPTION OF A PHYLLORHYNCHUS FROM CERRALVO ISLAND, GULF OF CALIFORNIA, MEXICO Arnold L. Powers^ and Benjamin H. Banta^ Abstract. — Phyllorhynchus decurtatus porelli ssp. nov. is described from a single specimen from Cerralvo Island, Gulf of California, Mexico, and is regarded as different from P. d. decuratus on the basis of an increased number of dorsal blotches (41 as compared to 18-33) and an increased number of supralabials (7 as compared to 6), combined with geographical isolation and the high degree of endemism found in other reptiles on Cerralvo Island. Among the herpetological specimens secured from Cerralvo Island, Gulf of California, Mexico, in January 1973 is an example of Phyllorhynchus decurtatus (Cope). It constitutes the first record of this species for Cerralvo Island. The specimen resembles a speci- men of P. decurtatus (sdsnh 44682) reported by Soule and Sloan (1966) from San Jose Island, located approximately 90 kilometers to the northwest (Fig. 1). When compared with the published meristic and morphological data provided by Klauber (1940) and Smith and Langebartel (1951), a number of distinctive features were found (Table 1). These data combined with the geographical isolation from the most similar populations on the Baja California peninsula and the high degree of endemism found in other reptiles occurring on Cerralvo Island merit the recognition of a distinct geographic race. Table 1. Summary of selected morphometric and meristic characteristics of nominal forms of Phyllorhynchus adapted from Klauber (1935, 1940); Soule and Sloan (1966); Savage and Cliff (1954); Smith and Langebartel (1951). Sub- Supra- Dorsal Species Ventrals caudals labials blotches decurtatus 157-167 33-36 6 18-33 decurtatus* 153 33 6 28 porelli 160 33 7 41 arenicola 164 39 6 30-32 norrisi 151-156 31-34 6 28-48 perkinsi 168-182 32-41 6 26-57 •from San Jose Island Phyllorhynchus decurtatus porelli, ssp. nov. HoLOTYPE.^ — Subadult male; Natural History Society of Mary- land number R1800 nhsm collected by Ismael Aviles from south- west Cerralvo Island, Gulf of California, 15 January 1973. Diagnosis. — A new geographic race of Phyllorhynchus decur- tatus closely related to peninsula populations of P. decurtatus in 'Department of Environmental, Population, and Organismic Biology, University of Colorado, Boulder, Colorado 80302. -Department of Biology, United States International University, San Diego, California 92131. 241 242 GREAT BASIN NATURALIST Vol. 34, No. 3 LOCALITY RECORDS PHYLLORHYNCHUS P d porelli Pa decurtatus Pd perkinsi Pd nubilus Perkinsi-nubilus infergrades Pb browni Pbljcidus SCALE OF MILES ^eerr Cerrolvo I NOTE - THE SMALL SCALE PREVENTS SHOWING ALL P d perkinsi records IN CALIFORNIA AND ARIZONA Adopted from Klouber ( 1940 1 Fig. 1. Distribution of the genus Phyllorhyncus. having an increased number of dorsal blotches (41 as compared to 18-33) and an increased number of supralabials (7 as compared to 6, Table 1). Description.— Rostral large, truncate in dorsal profile, edges protruding and striated; intemasals triangular, separated by rostral, in contact with upper loreal and both sections of nasal. Body mod- erately stout, slightly flattened below but cylindrical. Tail relatively Sept. 1974 POWERS & BANTA: MEXICAN SNAKE 243 **■■ a~ f :t? ^ftK ^ Fig. 2. Head of P. d. porelli, dorsal (a), ventral (b), and anterior (c) views. Fig. 3. Head views of P. d. porelli from Cerralvo Island (top) and P. d. decurtatus from San Jose Island (bottom). short, decreasing rapidly in girth posteriorly. Head only slightly distinct from body, short with a blunt snout; not as broad at trunk as at mid-body. Scale rows 21-19-16; apical scale pits single; ven- trals 160; anal single; subcaudals 33, all paired. The primary dorsal pattern is a series of 41 irregular brown blotches with a cream-white "S. . ' .. -ft' 1* <> * /A i*. ■ Fig. 4. Dorsal views of P. d. decurtatus from San Jose Island and P. d. porelli from Cerralvo Island. 244 GREAT BASIN NATURALIST Vol. 34, No. 3 background. On the sides are auxiliary series of spots, irregular in form (Figs. 1-3). Remarks. — Geographically, porelli is most closely related to decurtatus^ which occurs on the adjacent Cape Region to midcentral Raja California. A relationship of porelli, as with the San Jose specimen of decurtatus, can also be deduced with the western Mexico mainland norrisi, found approximately 500 kilometers northward across the Gulf of California in Sonora. The Monserrate Island species arenicola, can also be readily related to porelli based on com- parable ventral counts. A better appraisal of the systematic status of this insular form can be obtained only if and when additional material becomes available for study. Acknowledgments The expedition was partially sponsored by the Natural History Society of Maryland, Inc., through the generosities of Mr. Herbert S. Harris, Jr., and Dr. Robert S. Simmons. Mr. Clark R. Mahrdt, San Diego Society of Natural History (sdsnh) provided access to the society's extensive collections. Dr. Hobart M. Smith, EPO Riology Department, University of Colorado, provided editorial sug- gestions. Photographs were provided by Mr. Roy Porello, for whom the new race is named in appreciation of his many photographic contributions to the authors. Literature Cited Klauber, L. M. 1940. Two new subspecies of Phyllorhynchus, the leaf-nosed snake, with notes on the genus. Trans. San Diego Soc. Nat. Hist. 9 (20): 195-214, plates 8, map. . 1935. Phyllorhynchus, the leaf -nosed snake. Bull. Zool. Soc. San Diego 12:4-31, figs. 1-4, map. Savage, .1. M., and F. S. Cliff. 1954. A new snake, Phyllorhynchus arenicola from the Gulf of California, Mexico. Proc. Biol. Soc. Washington 67:69-76, figs. 1-2. Smith, H. M., and D. A. Langebartel. 1951. A new geographic race of leaf- nosed snake from Sonora, Mexico. Herpetologica 7(4) : 181-184. SouLE, M.. AND A. J. Sloan. 1966. Biogeographv and distribution of the rep- tiles and amphibians on islands in the Gulf of California, Mexico. Trans. San Diego Soc. Nat. Hist. 14(11):137-156, figs. 1-4. TWO NEW VARIETIES OF ERIOGONUM (POLYCJONACEAE) FROM THE INIERMOUNTAIN REGION James L. Reveal^ Abstract. — Two new variants of Eriogonum are proposed: E. brevicaule var atwoodii from near Fredonia. Mohave Co., Arizona and E. pharnaceoides var. cervinum from southwestern Utah and adjacent northern Arizona and south- eastern Nevada. After a detailed review of the genus Eriogonum (Polygonaceae) for the state of Utah was pubHshed (Reveal, 1973a, b), material representing two undescribed varieties was discovered, and these are hereby proposed as new. Eriogonum thompsonae S. Wats. var. atwoodii Reveal, var. nov. A var. thompsonae floribus albis et a var. albifloro foliis linearis differt. Typus: Arizona: Mohave Co.: Along Arizona Highway 389, 4.3 miles west of the junction of U.S. Highway 89A at Fredonia, on rolling reddish clay hills, associated with Atriplex, Ephedra, Stan- leya. and Eriogonum mortonianum. at about 4700 feet elevation, 15 August 1973, Reveal & Reveal 5211. Holotypus, US! Isotypi, 15 duplicates to be distributed from US. Low spreading herbaceous perennials (1) 1.5-3 dm high and 2-5 dm across; leaves sheathing up the stems 1-3 cm, linear, 3-8 (10) cm long including the petiole, 2-4 (6) mm wide, thinly to moderate- ly tomentose below, usually glabrous above, infrequently glabrous on both surfaces, the margin inrolled or at least thickened, the nearly indistinguishable petiole about 1/5 the length of the leaf-blade; stems erect, 5-10 (12) cm long, glabrous; inflorescences open, cy- mose, (5) 8-15 cm long, glabrous; involucres turbinate, 2.5-4 mm long, 1.5-2 mm wide, glabrous, the 5 acute teeth 0.3-0.5 mm long; flowers white, 3-3.5 mm long, glabrous, the tepals oblong; achenes light brown, 2.5-3 mm long. Other Specimens Examined: From the type area: 21 Jul 1973, Atwood 5584 (bry, us). This variety differs from typical var. thompsonae in having white flowers. The var. thompsonae occurs nearby on the steep slopes near Kanab westward to beyond Pipe Springs, but neither Atwood nor I found var. thompsonae in the vicinity of var. atwoodii. The var. alhiflorum Reveal is similar in that it too has white flowers, but var. atwoodii differs from it (and var. thompsonae as well) in having long, narrowly linear leaves. The discovery of var. atwoodii confirms the seemingly close association I have assumed Eriogonum thomp- sonae has had wdth the more northerly E. brevicaule Nutt. and the more easterly E. lonchophyllum Torr. & Gray, both of which gen- erally have narrow leaf-blades. The variant is named for N. Duane Atwood, a former fellow 'Department (if Rotany. University of Maryland, Cdllege Park 20742. and National Museum of Natural Ilistory. Smithsonian Institution. Washington. DC. 20560, Research supported by National Science Foiindaton Grant GB-22fi45. 245 246 GREAT BASIN NATURALIST Vol. 34, No. 3 graduate student at Brigham Young University, and an outstanding plant collector. Eriogonum pharnaceoides Torr. in Sitgr. var. cervinum Reveal, var. nov. A var. pharnaceoides floribus f la vis differt. Typus: Utah: Washington Co.: Foothills south of Pinto on the north slope of the Pine Valley Mountains, associated with pinyon- juniper and other shrubs, 18 August 1973, Atwood & Higgins 58 95. Holotypus, US! Isotypi, 20 duplicates to be distributed by US. Erect herbaceous annuals 1-3 dm high; leaves basal and cauline, the basal ones linear-lanceolate to linear-oblanceolate, 1-3 (4) cm long, 1-2.5 (4) mm wide, lanate below, villous and greenish above, the petiole 1-3 (5) mm long, the cauline ones linear, 0.5-2 (2.5) cm long, 0.5-2 (3) mm wide, tomentose to lanate below, thinly villous to infrequently glabrous above, sessile or on petioles less than 2 mm long; stems erect, villous, leafy; inflorescences open, 0.5-2.5 dm long and up to 2 dm wide, the branches villous throughout; peduncles slender, erect or nearly so, (1 ) 2-5 cm long, villous to glabrous; invo- lucres campanulate, the tube 1-2 mm long with 5 ± erect, lanceolate lobes 1-2 (3) mm long, villous; flowers yellow, 1-3 mm long, gla- brous, the tepals dimorphic, those of the outer whorl oblong-ovate with a large saccate base on each side of the truncate to cordate base, those of the inner whorl linear-oblong and erect, usually far exceed- ing the length of the outer tepals; achenes brown, 1.8-2.3 mm long. Other Specimens Examined: Arizona: Mohave Co.: Upper Trumbull Mountain, 6500 ft elev, 8 Sep 1953, Merkle & Merkle 950 (GCNP). Nev.ada: Lincoln Co.: Deer Lodge, 15 Aug 1935, Hall s.n. (bry); Deer Lodge, 7200 ft elev, 1 Sep 1935, Hall s.n. (bry). Utah: Millard Co.: Cane Springs, 6000-7000 ft elev, 1898, Purpus 6229 (k, uc, us). Washington Co.: Pine Valley Mountains, 22 Jul 1973, Atwood 5418 (bry, mary, us); near Grass Valley Reservoir, 22 Jul 1973, Higgins 7921 (us); Mountain Meadows, 15 Aug 1935, Stanton 6583 (so). The var. cervinum differs from var. pharnaceoides in having yellow instead of white flowers and occupying a geographical area north and west of the typical variant. The flowers of var. cervinum are also a bit smaller, but the measurements overlap. As I indicated before (Reveal, 1973a), the problem with this variety has been the paucity of material for critical evaluation and use as a type. In 1973, Atwood and Larry C. Higgins obtained specimens of this variant for me, and they succeed rather well. In addition, a visit to the Grand Canvon National Park resulted in the discovery of var. cervinum in northern Arizona, and thus the known range can be expanded to include this state for the variety. Literature Cited Reveal, J. L. 1973a. Eriogonum (Polvgonaceae) of Utah. Phvtologia 25: 169-217. . 1973b. "Eriogonum." In: Welsh, S. L., and G. Moore, Utah plants, tracheophyta. Brigham Young University Press. Provo, Utah. pp. 254-275. 1974. Two shrubby novelties in Eriogonum (Polygonareae) from the deserts of Utah and Arizona. Brittonia 26:90-94. GREAT BASIN NATURALIST Special Notice Commencing with volume 35 (March 1975) The Great Basin list will be combined with i' " ' ' ^' ■: University Bulletin, Biological Series ui Great Basin list. The format will be changed irom liie { ,._ column page to a 7x10 inch double colunm pa^ ^ i il continue to be published quarterlj^ but the number of pages per LLune will increase to approximately 500. Manuscripts up to 100 iited pages (about 200 manuscript pages) in lengtli will be ac- I table. Publication and reprint charges will be adjusted to the ;v format, but will not be significantly different from that indi- ed below. The subscription rate will be increased to $9.00 per .r (outside the United States, $10.00) . Notice to Coin.JiiiJi.vi.ors Original manuscripts in Enghsh pertaining to the Biological rural history of western North America and intended for publica- II in the Great Basin Naturalist should be directed to Brigham nmg University, Stephen L. Wood, Editor, Great Basin Naturalist, >vo, Utah 84602. Manuscripts: Two copies are required, typewritten, double- iced tliroughout on one side of the paper, with margins of at least -^ inch on all sides. 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Reprints: No reprints are furnished free of charge. r reprints and an nrrlor form are '^'^^^t \\\\h tlip Tn-ncif, Repj EUNTS SCI lEDULE OF THE L tREuVI DASIN iNATUR^SLlilST Each Additional 0 copies ^M copies ^m copies ^K copies 2pp. $12.00 15.00 18.00 20.00 4pp, 6pp. $14.00 $16.00 17.00 19.00 20.00 22.00 22.00 24.00 8pp. 10pp. 12pp. 2pp. $18,00 $20,00 $22.00 $3.00 21.00 23.00 25.00 24.00 26.00 28.00 26 00 23.00 30.00 wL TABLE OP CONTENTS Studies on the biology and ecology of the northern scorpii Paruroctoiius boreus (Girard). Gregory Ira Tourtlotte .... oji Notes on three varieties of Astragalus lentiginosus (Legumi- nosae). Carol S. Schoener 180 New species of American Corthylus (Coleoptera: Scolytidae). Stephen L. Wood 181 Notes on Viscawopelmntus davewerneri (Orthoptera; Gryl- inae) from the Viscaino Desert, Aico. Ernest R. TinMiam, Clark ri. j.>iiuirnx, invl xjeiJjHinin H. Banta 203 The austrina group of the genus Microvelia (Henxiptera: Veliidae). John T. Polhemus 207 Additional records of Coleoptera collected at the Nevada Test Site. Mercury, Nevada. Vasco M. Tanner and Wihiier W. Tanner 218 A summary of biological investigations concerning the Great Salt Lake, Utah (1861-1973). Doyle W. Stephens .......... 221 A new variety of Penstemon thurberi (Scrophulariaceae). James L. Reveal and Janice C. Beatley 230 A new sceloporine lizard from Oaxaca, Mexico. Marlene M. Dasmann and Hobart M. Smith 231 Rartge extension of the long-nosed snake, Rhinocheilus I. lecontei, into east-central Utah. William L. Grogan, Jr. and Wihner W. Tanner 238 Description of a Phyllorhynchus from Cerrovalo Island, Gulf if California, Mexico. Arnold L. Powers and Benjamin 1. Banta 241 Two new varieties of Eriogonum (Polygonaceae) from the Xntermountain region. James L. Reveal 245 Volume 34, No. ^' December 31, 1974 The Grea ]]j] j u ji jjij^ J PUBUSHED BY Brigham Young UNivEHSiT^f GREAT BASIN NATURALIST Editor: Stephen L. Wood, Department of Zioology, Brigham Young University, Provo, Utah Editorial Board: Stanley L. Welsh, Botany, Chairman; Wilmer W. Taimer, Zoology; Joseph R. Murdock, Botany; Veinon J. Tipton, Zoology; Ferron L. Andersen, Zoology Ex officio Editoricd Board Members: A. Lester Allen, Dean, College of Biological and Agricul- tural Sciences Ernest L. OIsod, Director, University Press, University Editor The Great Basin Naturalist was foimded in 1939 by Vasco M. Tannei\ It has been continuously published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only original, previously impublished manuscripts pertain- ing to the biological natural histor}'' of the Great Basin and western North America will be accepted. Manuscripts cire subject to the ap- proval of the editor. SuBscBiPTiON: The annual subscription is $5.00 (outside the United States $5.50). The price for single numbers is $2.00 each. All matters pertaining to me purchase of subscriptions and back numbers should be directed to Brigham Young University Press, Publication Sales, 205 UPB, Provo, Utah 84601. Libraries or other organizations interested in obtaining this journal through a contin- uing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, The Library, Provo, Utah 84601. Manuscripts: All manuscripts and other copy for the Great Basin Naturalist should be addressed to the editor. Contributors ' ' ' lilt the instructions printed on the back cover of a recent The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University Volume 34 December 31, 1974 No. 4 FIELD BEHAVIOR AND SEASONAL ACTIVITY OF THE RODENT EOT FLY, CUTEREBRA TENEBROSA, IN CENTRAL WASHINGTON (DIPTERA: CUTEREBRIDAE) ' Craig R. Baircl- Abstract. — Behavior and activity of the rodent hot fly, Cuterebra tenebrosa Coquillett, was studied at a natural aggregation site in central Washington. Daily flight activity extended from 1700 hr to 2030 hr (PDT) beginning in mid-July of 1970 and 1971. The peak of flight activity, in August, was followed by a decline in activity through September. Male flies aggregated on the vertical faces of basalt cliffs, where they awaited females. Daily activity reached a peak at temperatures of 30-35 C. Eggs were laid in rock crevices witliout regard to the pro.ximity of Neotoma wood rat hosts. Developing fly pupae were found in litter in wood rat tunnels in basalt cliffs. Adult bot flies are rarely encountered in nature, and their general behavior was little known until recent years. Most early information was gathered from scattered accounts of one or two flies captured during the warm months, and specific identification was often im- possible due to lack of adequate keys. The first in-depth study of adult Cuterebra activity and behavior was by Catts (1967), who worked with C. latifrons Coquillett in California. Adult C. polito Coquillett activity at aggregation sites in Utah was studied by Gra- ham and Capelle (1970) and Capelle (1970). Recently, Hunter and Webster (1973) reported behavioral studies of Cuterebra grisea Coquillett and C. tenebrosa Coquillett in British Columbia. Very little additional information is available on the activity of Cuterebra tenebrosa, a large black cuterebrid whose larvae parasitize Neotoma wood rats in the western United States and western Canada. Parker and Wells (1919) obtained a female in September in Montana, and Moilliet (1950) reported the capture of a female in British Columbia in August. During 1970 and 1971, adult C. tenebrosa were studied at natural aggregation sites in central Washington. The purpose of this paper is to describe: (1) seasonal and diurnal occurrence of adult C. tene- brosa at aggregation sites, (2) flight activity and mating behavior, and (3) oviposition activity and sites. 'Sricnlific Paper No. 419+. College of Agriculture Hcscarrli Ot'LCr. Washington Stale Universi- IV Work londiKted under Projetl 9043, suppurlcd in part l)y a National Defense F.ducation Act Tille IV Fellowship. -Department of Enloniologv. Washington State University. Pullman. Washington 99!()3 Ciinpnt address University of Idaho Extension Service, P. O. Bo.x 1()'>S. Caldwell, Idaho 83005 247 248 GREAT BASIN NATURALIST Vol. 34, No. 4 Materials and Methods Field observations were conducted at the Columbia National Wildlife Refuge, Othello, Washington. The area is largely composed of rock slides and 15-35 m high basalt cliffs (Fig. lA) with sage- brush and dry land grasses the main vegetation. Despite the overall dryness, a number of small streams and lakes exist in the lower areas (Fig. IB). Observations were made intermittently during all months of the year; however, the main attempts to study fly activity were between April and October of 1970 and 1971. Searches for aggregation sites Ni fr *• ■:ak- B iig. I. Cjeiit'idl \ it'w ol til til will] I- LiiUitbid /rnrh/ Oiu i> , Direction K ) ' ^x / 1 \^ Dropped X x"^^ to Ground V ^ i '^-- ^ I ^\.^ •^ V WARNER ^^^^ VALLEY ^ ^ ^ "^"^^^^% "^-^ ^— ^ 0 .5 1km Ruins -^^ D ^^ \ / .'« ^°'V^^^ '"o^"^ ° ^.^-^ "^ \y vo'< ''"' y^ ,-,-* ,,'' UTAH \ X- --' ARIZONA 1 Fig. 1. The Washington Co., UtaJi, study site where spotted bat flight and behavior was observed. 256 GREAT BASIN NATURALIST Vol. 34, No. 4 spotted bat indicated that the diet was highly selective, consisting only of moths [Ross, 1961; Easterla, 1965; and Barbour and Davis, 1969a]. Easterla [1965] went so far as to speculate that the bat ate only moths of the family Noctuidae.) The descents to the ground were preceded by split-second hovering. After four minutes of cruising and feeding over the low-lying vegetation, the spotted bat disappeared near a crevice only 300 m from the point of release. We were unable to approach the apparent roost before nightfall due to the rough terrain. More detailed studies are currently under way. We feel that the population in the region is high as compared with estimates in pre- vious reports on the spotted bats. Indications are that density may be higher in Mohave County, Arizona, where a similar habitat exists in close proximity to Fort Pierce Wash. We would like to thank the Nevada Power Company, of Las Vegas, Nevada, for funding this project and for the continuation of natural history studies of the bat. Our appreciation is extended to Clyde Jones for critically reviewing this note. Literature Cited Barbour, R. W. and W. H. D.avis. 1969. Euderma maculatum from Bats of America. Univ. Kentucky Press. Lexington, pp. 161-163. Benson, S. B. 1954. Records of the spotted bat {Euderma maculata) from California and Utah. J. Mammal. 35:117. CoNSTANTiNE, D. G. 1961. Spotted bat and big free-tailed bat in northern New Mexico. Southwest Nat. 6:92-97. DuRANT. S. D. 1935. Occurrence of the spotted bat in Utah. J. Mammal. 16:226. Easterla, D. 1965. The spotted bat in Utah. J. Mammal. 46:665-668. . 1970. First records of the spotted bat in Texas and notes on its natural history. Am. Mid. Nat. 83:306-308. FiNDLEY, J. S., AND C. JoNES. 1965. Comments on spotted bats. J. Mammal. 46:679-680. Hall, E. R. 1935. Occurrence of the spotted bat at Reno, Nevada. J. Mammal. 16:148. Hardy, R. 1941. Some notes on Utah bats. J. Mammal. 22:289-295. Jones, C. J. 1961. Additional i-ecords of bats in New Mexico. J. Mammal. 42:538-539. Rodeck, H. G. 1961. Another spotted bat from New Mexico. J. Mammal. 42:401. Ross, A. 1961. Notes on food habits of bats. J. Mammal. 42:66-71. VoRHiEs, C. T. 1935. The Arizona specimen of Euderma maculatum. J. Mammal. 16:224-226. THE SIGNIFICANCE OF SCALE CHARACTERS IN EVALUATION OF THE LIZARD GENERA GERRHONOTUS, ELGARIA, AND BARISIA James W. Waddick' and Hobart M. Smith= Abstr.'\ct. — Data taken on external scale characters of 1003 specimens rep- resenting nine of the sixteen species of Gerrhonotus sensu Stebbins, 1958, strong- ly indicate that Tihen's 1949 arrangement of those species in three genera {Gerrhonotus, Elgaria, Barisia) is valid. Misinterpretation of the identity of the head scales in various species of this group has led erroneously to disregard of them as indicators of relationships. Actually the scales are as constant as in most other lizards and seemingly provide finn clues to natural associations. The proper generic allocation of species of "gerrhonotine" lizards, defined as those appropriately referred to Gerrhonotus Wiegmann {sensu lato) as understood before 1942 (Smith, 1942) has remained enigmatic despite the documentation provided by the most recent review of the group by Tihen (1949), based upon osteology and external scutellation. The primary doubt was cast upon the validity of Tihen's groupings by Stebbins (1958), who proposed an alternative grouping based upon reproductive habits, color patterns, and habitat. Haunted by the impression that external scutellation provides more reliable clues to relationships in this group than was thought by Stebbins, we initiated a re-examination of this particular aspect, utilizing materials in the University of Illinois Museum of Natural History (uimnh), University of California Museum of Vertebrate Zoolog}^ (mvz). University of Kansas Museum of Natural History (kumnh), University of Michigan Museum of Zoology (ummz). United States National Museum (usnm), Brigham Young University Museum of Natural History (byu). University of Colorado Museum (cum) and University of Texas Natural History Collection (tnhc). We are much indebted to authorities at these institutions for the privilege of borrowing material from them; particularly instrumen- tal were Dr. Donald F. Hoffmeister, Dr. Robert C. Stebbins, Dr. E. Raymond Hall, the late Dr. Norman Hartweg, the late Dr. Doris Cochran, Dr. Wilmer W. Tanner, Dr. T. Paul Maslin, and Dr. W. F. Blair. As is apparent from this list, the work here reported was completed more than a decade ago. Its results remain valid and of current interest. Descriptive Terminology The definition of the genera of gerrhonotine lizards requires a definitive identification of the head and body scales involved. Uni- formity of terminology' has not existed in the past. Indeed, misidenti- fication of scales was important in Stebbins' (1958) rejection of scutellation as a reliable indicator of relationship. The nomenclature 'Curator of Education, New York Zoological Society, 185lli & Southern Blvd., Bronx, New York, 104T-iir),- ? o o o 05 o o o o VO »f5 CSl CO o o o O) o 05 rr> ? •^ •^ ■^ xr> OO »« ^ O O) o O) o o o o o o o O O " ^ O o o o o o o o O O ro CM O _. __ ^ c^ O O o o O 05 o 00 o o o o o OO vf) o ? ooorooooo O -H ^ ^ O) t^ (5 vo v_- : -~y ; o^ W3 o^ '-N -sP ; ^-^ o^ v_^ v^o o^ ^' .o C u -^ nj CO 5 e^ (U oj aj ci QJ ^ fC CO CO J° ^ g CO *3 • 4J *J Ou^coCCoo 3 3 H O 266 OREAT BASIN NATURALIST Vol. 34, No. 4 Table 3. Contrasts between the genera Gerrhonotus, Elgario, and Barisia. Genera GERRHONOTUS ELGARIA BARISIA 1 . Postrostral absent 2. Nasal-rostral contact 3. Cantholoreal present 4. Ant. intern, present 5. Ant. intern, cont. 6. Supranasals cont. 7. Two ant. gulars Seldom Rarely not Usually Rarely Rarely not Rarely Seldom Rarely not Usually Always Rarely Rarely not Never Never Usually Never Rarely not Rarely Usually Rarely Rarely to question. The differential characters, however (even though each separately overlaps at least slightly), collectively indicate a long history of independent evolution that we regard as being consistent with separate generic status. We anticipate that ethological, sero- logical, osteological, and karyological work in the future will sub- stantiate these groupings and their generic rank. It is quite evident that Barisia is the more variable and plastic of the three more closely related gerrhonotine genera, and it is pre- sumably the most primitive of them. Elgaria and Gerrhonotus ap- pear to be almost equally specialized derivatives from ancestral forms of Barisia^ although the imbricata series of the latter genus is almost as specialized as the genera Elgaria and Gerrhonotus. Intermediates link all members of Barisia, however, whereas the members of both Elgaria and Gerrhonotus are trenchantly distinctive. Literature Cited Criley, B. B. 1968. The cranial osteology of gerrhonotiform lizards. Am. Midi. Nat. 80(1): 199-219, 3 figs. Smith, H. M. 1942. Mexican herpetological miscellany. 3. A tentative re- arrangement and key to Mexican Gerrhonotus, with the description of a new race. Proc. U. S. Nat. Mus. 92:363-369. Stebbins, R. C. 1958. A new alligator lizard from the Panamint Mountains, Inyo County, California. Am. Mus. Novit. (1883): 1-27, figs. 1-6. Tanner, Wilmer W. 1959. The status of Gerrhonotus in Utah. Herpetologica 15:178-180, fig. 1. Tihen, J. S. 1949. The genera of gerrhonotine lizards. Am. Midi. Nat. 41 (3): 580-601. Wermuth, H. 1969. Liste der rezenten Amphibien und Reptilien. Anguidae, Anniellidae, Xenosauridae. Tierreich 90:i-xii, 1-41. Woodbury, A. M. 1945. A new Gerrhonotus lizard from Utah. Proc. Biol. Soc. Washington 58:5-10, figs. 1-2. REVEGETATION OF GOPHER MOUNDS ON ASPEN RANGE IN UTAH W. T. McDonoughi Abstract. — The colonization of the pocket-gopher {Thomomys talpoides) mounds b}- annual and perennial species of the understory of aspen woodland was observed over a four-year period. New and old gopher mounds exist as a mosaic of sites in one of three surface conditions: bare, dominated by annuals with a few seedlings of perennials, and dominated by perennials with annuals in peripheral areas. The regular creation of these new sites for plant colonization appears to favor the maintenance of aggressive perennials at high densities in the understory vegetation. On mountain rangeland in the western United States, foraging and soil-displacement activities of pocket gophers {Thomomys tal- poides) have important effects on vegetation (Inlander et al., 1969; Turner, 1969). The mounds and casts created by runway and cavity excavation present a new surface for plant colonization — seedbed conditions that may favor the establishment of particular species (Laycock, 1958). The purpose of this study was to describe the colonization of gopher mounds over a four-year period ( 1 969 through 1972) on aspen {Populus tremuloides Michx.) range in Utah. The aspen woodland of the study area (Fig. 1) occupies approxi- mately 2 ha on a 19-degree south-facing slope at an elevation of 1900 m in Tony Grove Canyon on the Cache National Forest in northern Utah. Density of aspen (DBH greater than 5 cm) averages 2900/ha, and there is a dense understory vegetation of shrubs and annual and perennial forbs and grasses. Florez (1971) described the soil, climate, and vegetation of the site. Methods In early July 1968, 80 new gopher mounds were identified and numbered. In July 1969, the 50 mounds with the lowest assigned numbers and no previously established perennials emerging from them were designated for stud}'. Percentage canopy cover of indi- vidual colonizing species was estimated visually in early Jidy and early September of each year. To determine the density, frequency, and distribution of new mounds each year, two belt transects, each 128 m long and consisting of 128 quadrats (each 1 m-), were established 30 m apart near midslope where mound counts had indicated high mound-building activity. The transects were examined in early July of each year for new mounds. A mound was counted if any part of it was found within a quadrat. The distribution of mounds was estimated by use of variance-mean ratios (Greig-Smith 1964). In mid-June and mid- July of 1970, plant species in each of the quadrats of the transects 'Plant physiologist, Intennountain Forest and Hangc Experiment Station, ORden. Utah 84401; stationed in Logan, Utah, at the Forestry Sciences laboratory in cooperation with Utah State University. 267 268 GREAT BASIN NATURALIST Vol. 34, No. 4 » r *^M:^' . " Fig. 1. The aspen woodland study area. were noted. Dimensions and other characteristics of mounds were taken from 43 new mounds found in the transects in 1969. Fre- quency percentages of species on mounds over the four years of colonization were taken from the 50 mounds selected in 1969; quadrat frequencies were based on the transect data collected in 1970. Soil samples of equal volume were taken from five new mounds and adjacent, undisturbed topsoil in late September of each year for standard fertility analysis — texture, pH, conductivity (total salts), organic matter, P, K, Ca, and NO3-N. Subsamples from these collec- tions were used to determine seed content by sieving and flotation on water. Additional collections of mound and adjacent topsoil were used in pot culture of two common annual {Collomia linearis, Veronica biloba) and two perennial {Rudheckia occidentalism Senecio serra) species with high densities in the study area. The purpose was to determine whether the soils affected plant growth. There were six 20-liter containers per species and four plants per container. After two months' growth in the greenhouse, shoots were harvested for oven-dry weight determinations. The statistical significance of differences was evaluated at the 5 percent level. Results Plants. — Based on quadrat frequency percentages, the most prevalent species were Ncmophila brevi flora and Polygonum doug- lasii among the annuals and Brornus cnrinatus and Rudbeckia occi- dentalis among the perennials (Table 1). Seed dispersal occurs Dec. 1974 mcdonough: gopher mounds 269 Table 1. Species frequencies in the quadrats of belt transects in 1970 and on gopher mounds during four years of colonization. Frequency (%) Species Quadrat Mound Aruiual Chenopodium album L 40 26 Collomia linearis Nutt 43 38 Galium bifolium S. Wats 34 46 Nemophila breviflora A. Gray 87 82 Polygonum, douglasii Greene 68 56 Veronica biloba L 57 64 Perennial Achillea millefolium L 15 12 Agastache urticifolia (Benth.) Kuntze 13 18 Agropyron trachycaulum (Link) Malte 42 52 Bromus carinatus Hook. & Am 83 74 Delphinium occidentale (S. Wats.) S. Wats 19 20 Elymus glaucus Buckl 21 34 Phacelia utahensis J. Voss 30 22 Rudbeckia occidentalis Nutt 95 82 Senecio serra Hook 40 56 Stellaria jamesiana Torr 18 34 among the annuals from late June and early July (A^. breviflora, V. biloba) through September {P. douglasii, Chenopodium alburn)^ and toward the end of summer in most of the perennials. Seeds of the species listed in Table 1 are gravity dispersed, except for S. serra (wind dispersed) and Galium bifolium (adhesive). Gopher mounds. — The outline of mounds varied from nearly circular to elliptic, with mean dimensions of 46 by 38 by 9 cm depth. Density of mounds per m- ranged from 0.06 to 0.22 and frequency percentages from 5 to 17 in the transects over the four-year period (Table 2). Variance-mean ratios were significantly greater than unity in four samples, indicating clustered distributions, and did not differ significantly in three samples, indicating random distributions (Greig-Smith 1964). Mound clusters were observed in some loca- tions (Fig. 2), usually in midsummer. New mounds occurred in Table 2. Density, frequency, and variance-mean distribution ratios of new mounds in the quadrats of belt transects. Year Transect Density (m2) Frequency (%) Variance- mean ratio 1969 1 2 0.17 .16 13 12 1.41* 1.44* 1970 1 2 .06 .15 5 12 1.50* 1.13 1971 1 2 .12 .22 10 17 1.08 1.27* 1972 1 2 .09 .19 8 16 1.11 1.05 •Differs significantly from unity. 270 GREAT BASIN NATURALIST Vol. 34, No. 4 Fig. 2. Clustered mounds were observed midsummer (30 cm ruler in foreground). in some locations, usually in Dec. 1974 MCDONOUGH: GOPHER MOUNDS 271 different quadrats from year to year, and 40 percent of the quadrats were occupied one or more times over the four-year period. Soil. — The soil of new mounds was higher in conductivity and lower in NO3-N than was adjacent topsoil (Table 3). Since the soil of the mounds had been reworked by gopher action, it was uncom- pacted and friable as compared with the undisturbed topsoil. How- ever, these chemical and physical differences did not affect the dry weight of shoots of annuals (C. linearis, V . hiloba) or perennials {ft. occidentalis, S. serrd) after two months' growth in the greenhouse. Seeds. — No seeds were found in new mounds constructed before seed dispersal began; equivalent volumes of adjacent topsoil con- tained a mean of 36 seeds from nine species. Their weatnered ap- pearance suggested that most of these seeds were ungerminated from previous years and probably were not viable. By the end of Sep- tember when seed dispersal had ceased, the mean count per mound was 37, significantly higher than the 23-seed mean for equivalent columns of adjacent topsoil. Mounds appeared to act as catchment areas; dispersed seeds were often buried in the loose mound soil. Four species (A^. hreviflora, P. douglasii, B. carinatus, and R. occi- dentalis) accounted for 64 percent of all seeds recovered. Colonization. — By October of the first year, 8 percent of the new mounds had been colonized by seedlings of three species: Aga- stache urticifolia, R. occidentalis. and V . biloba. By early summer of the second year, 94 percent of the mounds had been colonized by seedlings; and 100 percent colonization was attained by late spring of the third year. For species occurring on both quadrats and mounds, quadrat frequencies (greater than 10 percent) in transects in the summer of 1970 and frequencies on 50 mounds over the four-year period are given in Table 1. Photographs of representative mounds in various stages of colonization are shown in Figures 3 through 6. Of the 8 annual and 14 perennial species that colonized mounds, A^. brevijlora was the best represented annual and R. occidentalis the most prevalent perennial. 1 he correlation coefficients between mound and quadrat frequencies were significant for both annuals Table 3. ResuUs of soil analysis from gopher mounds and adjacent un- disturbed topsoil. Factor Mound soil Undisturbed topsoil Texture Silt loam Silt loam pH 6.2 6.3 Conductivity (rrmihos/cm)* .33 .25 Organic matter (%) 4.7 4.9 P (ppm) 56 65 K (ppm) 466 480 Ca 0 0 NO3-N (ppm)* 7.2 8.0 •Results differ significantly. 272 (iREAT BASIN NATURALIST Vol. 34, No. 4 Fig. 3. First-year colonization by C. linearis, N. breviflora, and S. jamesiana. Fig. 4. Second-year colonization by A', breviflora. Doc. 1974 MCDONOUGH: GOPHER MOUNDS 273 t^V-^-' .^1 Fig. 5. Third-year colonization by R. nccidentalis. Fig. 6. Fourth-year colonization hy .S. serra. 274 GREAT BASIN NATURALIST Vol. 34, No. 4 80 - 60 o " 40 20 - PERENNIALS ANNUALS xD 1969 1970 Year 1971 1972 Fig. 7. Percentage of annual and perennial species cover during four years of colonization. (r = 0.85) and perennials (r = 0.94). This reflects gravity dis- persal of seeds to the mounds from adjacent plants, with higher probabilities for colonization by the more widely distributed species in the area. The revegetation of mounds progressed from an initial dominance by annuals to dominance by perennials^ — a displacement largely completed within the four-year period (Fig. 7). The annuals reached maximum cover values in the second year and declined thereafter, while cover by perennials rose steadily through the period. Conclusions Soil disturbance by gophers creates small and constantly shifting new sites for colonization by plant species in the vicinity. Because new mounds appeared in 40 percent of the quadrats in the sample transects over the four-year period, it seems probable that a large portion of the woodland soil is overturned and laid bare over longer periods. Gopher mounds thus represent a mosaic of sites, constantly shifting in time and space, that individually exist in one of three soil-surface conditions — bare, invaded and dominated by annuals with a few seedlings of perennials, and dominated by perennials with a few persistent annuals in peripheral areas. In aspen vegetation in Wyoming, Laycock (1958) found that 95 percent of all individual plants growing on gopher mounds during one summer were annuals. He concluded that the persistence and spread of annuals in the communit}" were related to the level of mound-building activity. The constant renewal of bare surfaces keeps the aspen understory open to annuals that otherwise might be eliminated or persist only in low densities. Dec. 1974 mcdonoijgh: gopher mounds 275 The constant renewal of bare surfaces also appears to work to the advantage of aggressive perennials, such as R. occidentalism which is known to be an increaser s})ecies on aspen range. Rudbcckia pro- duces an abundance of seed of high and ra})id germinability over a wide temperature range, and there is vigorous seedling growtli with relatively low mortality (Florez 1971; Florez and McDonough 197S). Rudbcckia appears, along with B. carinatus, to take advan- tage of gopher mounds to increase its importance in the aspen under- story vegetation. However, the presence of aggressive species does not negate the effect of gopher activity in maintaining diversity in the species composition of tlio understory vegetation. Literature Cited I'r.oRF.z, J. A. 1971. Ecological life history of Rudbcckia occidentalis Nutt. Ph.D. dissertation. Utah State Univ., Logan. 165 p. Florez, A.. .\nd W. T. McDonough. 1973. Seed germination, and growth and development of Rudbcckia occidentalis Nutt. (western coneflower) on aspen range in Utah. Am. Midi. Nat. 91:160-169. Greig-Smith, p. 1964. Quantitative Plant Ecology. 2nd ed. Academic Press, N.Y. Jui-.^NDER, O., J. B. Low, .\ND O. W. MoRRis. 1969. Pocket gophers on seeded Utah mountain range. J. Range Manage. 22:325-329. T>.\YCOCK, W. A. 1958. The initial pnttem of revegetation of pocket gopher mounds. Ecology 39:346-351 . Turner. G. T. 1969. Responses of mountain grassland vegetation to gopher control, reduced grazing, and herbicide. J. Range Manage. 22:377-383. OBSERVATION OF VOICE IN THE WESTERN COLLARED LIZARD CROTAPHYTUS COLLARIS BICINCTORES Nathan M. Smith^ Abstract. — A noosed female Crotophytus collaris bicinclores Smith and Tanner voiced an alarm similar to that of C. wislizeni. Lizards, with the exception of the geckos, are considered voiceless (Smith, 1946; Stebbins, 1954). The gecko voice, which can be heard up to one hundred meters, is believed to serve to threaten or warn (Goin & Goin, 1971). Apparently some members of Iguanidae pos- sess a weak voice that also serves to threaten when provoked. Jor- gensen, Orton, and Tanner (1963) noted under laboratory conditions that the leopard lizard Crotaphytus wislizeni Baird and Girard changes its usual hissing to a high-pitched squeal before a vicious attack of its agitator. They concluded that the voice was characteris- tic of the Nevada Test-Site population. In late July 1970, a noosed male lizard and a noosed female wislizeni squealed and attempted to bite as they were removed from the noose. They were captured ap- proximately six miles north of Wendell, Gooding County, Idaho, just off state highway 46. On the same collecting trip, a few days later, a captured young female western collared lizard, Crotaphytus collaris bicinctores Smith and Tanner (BYU 3113), made a squealing sound while being re- leased from the noose. The squeal resembled that of the C. wislizeni. To my knowledge this is the first record of a distinct voice for this species. The capture was made on a hot afternoon about eight miles south of Bruneau, Owyhee County, Idaho, on the east side of the Bruneau River. At this time a comparative study of several members of the family Iguanidae and Coleonyx variegatus Baird is underway to describe anatomical voice mechanisms in these taxa. References Goin, C. J., and O. B. Goin. 1971. Introduction to herpetology. W. H. Free- man and Co., San Francisco, p. 119. JoRGENSEN. C. D., A. M. Orton, AND W. W. Tanner. 1963. Voice of the leopard lizard Crotaphytus wislizeni Baird and Girard. Proc. Utah Acad. Sci., Arts, Let. 40(1 ): 1 15-116. Smith, H. M. 1946. Handbook of lizards: lizards of the United States and Canada. Comstock Publishing Co., Ithaca, New York. p. 47. Stebbins, R. C. 1954. Amphibians and reptiles of western North America. McGraw-Hill Book Co., Inc., New York. p. 201 . 'Gradunic Department of Library and Infonnation Sciem cs. BriKliam Yniinp TTniversilv, Provo, Utah 84602. 276 NEW SYNONYMY AND RECORDS OF AMERICAN BARK BEETLES (COLEOPTERA: SCOLYTIDAE) Stephen L. Wood^ Abstr.-\ct. — New synonymy afforting American Scolytidae is proposed as follows: Acanthotomicus granulatus (Ferrari), n. comb. {^Miinips uncinatus Wood). Araptus dcyrollei (Blandford). new comb. ( -^Araplus insinuatus Wood), Cnesinus adiistus Schedl { = Cnesinus atrodeclivis Wood). Corlhylocurus mexi- canus (Schedl). n. comb. {=Corthylus cylindricus Schodl). Corthylus collaris Blandford {=Corthylus splendens Wood). Corthylus coma/us Blandford (= Cor- thylus spendidus Bright). Corthylus uniseptis Schedl {= Corthylus reburrus Bright.) Dendrocranulus carbonarius (Ferrari), n. comb. {^--Xylocleptes flori- densis Hopkins, Xylocleples anonne Hopkins). Dendrocranulus maurus (Bland- ford), n. comb. {^Dendrocranulus huehuetanus Schedl). Dendroctonus valens LeConte ( = Dendroctonus rhizophagus Thomas and Bright). Gymnochilus conso- cius (Blandford), n. comb. { = Problechilus trimaculatus Schedl), Gymnochilus minor (Blandford), n. comb. ( = Problechilus varius Schedl), Gymnochilus zona- tus Eichhoff ( = Problechilus freyi Schedl). Hypothenemus areccae (Hornung) {=Hypothenemus vafer Blandford). Hypothenemus erudiius W^estwood {=Cry- phalus obscurus Ferrari). Micracis grandis Schedl ( ^Micracis costaricensis Wood). Micracis swainei Blackman (=Micracis robustus Schedl. Micracis pyg- rnaeus Schedl). Microcorthylus parvulus Ferrari {=Pterocyclon exile Eichhoff), Monarthrum bicolor (Ferrari), n. comb. ( = Corthylus signatus Ferrari. Phthorius edentatus Hagedorn). Alonarthruni consimile (Blandford). n. comb. (^Ptero- cyclon pseudosulcatum Schedl). Monarthrum egenum (Blandford). n. comb. {=--Brachyspartus hisetosus Schedl). Monarthrum fimbraticorne (Blandford) {=Pterocyclon turbinatum Schedl). Monarthrum gnarum (Schedl), n. comb. { — Amphicranus spinalus Bright). Monarthrum ralidum (Ferrari), n. comb. {=Amphicranus mexicanus Eggers. Pterocyclon jalapae Schedl). Phloeoborus scaber Erichson ( = Phloeoborus opacithorax Schedl). Phloeosinus serratus (Le- Conte) ( ^Phloeosinus rugosus Swaine). Phloenlribus setulosus Eichhoff i^Phloeotribus rudis Eichhoff. Phloentribus sodalis Blandford. Phloeotribus bolivianus Eggers). Pycnarthrutn reticulatum Schedl { ^Pycnarthrum fici Wood), Scolytodes rugicollis (Schedl). n. comb. ( =Scolytodes plicatus Wood). Xylosan- drus morigerus (Blandford) ( = Xyleborus luzonicus Eggers). New names are proposed as follows: Araptus decorulus for Araptus decorus Wood. Cnemonyx nigrellus for Loganius niger W^ood, Monarthrum bicoloratum for Monarthrum bicolor Wood. Araptus sobrinus, n. sp. (Mexico), and Dendroctonus vitei, n. sp. (Guatemala), are named as new to science. Dendroctonus mexicanus Hopkins is removed from synonymy. During the preparation of a ta.xonomic monograph of the Scolyt- idae of North and Central America, the unpublished synonymy summarized in the above abstract and treated on the following pages was encountered. In order to stabilize nomenclature and fix estab- lished names, several lectotypes are designated. Specimens or series referred to in the discussion are in my collec- tion unless a definite statement indicates otherwise. The species are arranged in alphabetical order except that those described as new to science are presented at the end of the article. Acanthomicus granulatus (Ferrari), n. comb. Xylocleptes granulatus Ferrari. 1867, Die Forst- und Baumzuchtschadlichen 'Department of ZooIorv, Brighain YmiriK Univcrsitv. Provo, Utah 84ri02. Scnlvtidac rnnlrib\itif>ii No. .54. 277 278 GREAT BASIN NATURALIST Vol. 34, No. 4 Borkenkafer, p. 40 (Lectotype, male; Venezuela, probably at Colonia Tovar; Vienna Mus., present designation) . Mimips uncinatus Wood, 1971, Brigham Young Univ. Sci. Bull., Biol. Ser. 15(3) :41 (Holotype, male; Colonia Tovar, Aragua, Venezuela; Wood Coll.). New synonymy The type series of four specimens of Xylocleptes granulatus Ferrari collected by Moritz in 1858 was compared directly to the type series of Mimips uncinatus Wood. They are identical in every respect. It is a common species in the vicinity of the Moritz home at Colonia Tovar; it was not found in the Caracas area where Moritz lived at an earlier date. Araptus decor ulus, n. n. Araptus decorus Wood, 1974 (nee. Bright, 1972), Brigham Young Univ. Sci. Bull., Biol. Ser. 19(1 ):47 (Holotype, male; Rincon de Osa, Puntarenas, Costa Rica; Wood Coll.). Araptus decorus Wood is a junior homonym of Neodryocoetes decorus Bright (1972, Bull. Inst. Jamaica, Sci. Ser. 21:96), because the genera to which they belong have been synonymized. The new name Araptus decorulus is proposed as a replacement for decorus Wood. Araptus deyrollei (Blandford), n. comb. Pityophthorus deyrollei Blandford. 1904, Biol. Centr. Amer., Coleopt. 4(6):245 (Lectotype, male; Mexico; British Mus. Nat. Hist., present designation). Araptus insinuatus Wood, 1974, Brigham Young Univ. Sci. Bull., Biol. Ser. 19(1) :43 (Holotype, male; Guatemala; Wood Coll.). New synonymy Blandford named Pityophthorus deyrollei from two syntypic specimens taken in Mexico. Unfortunately these specimens were not available for loan at the time my study of this genus was made for the monograph, and I depended on specimens identified by other authors for my concept of Blandford's species. When the types were studied it was found that deyrollei is identical to Araptus insinuatus Wood. The first specimen in Blandford's series, a male, is here designated as the lectotype of deyrollei. Cnemonyx nigrellus, n. n. Loganius niger Wood, 1961 (nee. Eggers, 1933), Great Basin Nat. 21:95 (Holo- type, female; 26 km W Tehuantepec, Oaxaca, Mexico; Wood Coll.). Both Ceratolepsis niger Eggers (1933, Trav. Lab. Ent. Paris, Mem. 1:13) and Loganius niger Wood have been transferred to the genus Cnemonyx (Wood, 1972, Bull. Ent. Res. 62:344), thus creating homon}TTiy. The new name nigrellus is proposed to replace the junior name, niger (Wood). Schedl (1962, Beitr. Ent. 12:486) previously transferred Cera- tolepsis niger Eggers to the genus Camptocerus and thus created secondary homonjiny with Camptocerus niger (Fabricius). He then renamed Eggers's species Camptocerus nigricans Schedl. However, Dec. 1974 wood: American scolytidae 279 the type of the Eggers species is not congeneric wdth the type of the Fabricius species; consequently, the replacement name nigricans must be rejected and Eggers name restored to its original position. Cnesinus adustus Schedl Cnesinus adustus Schedl. 1949, Rev. Brasil. Biol. 9:266 (Lectotype, female; Turri- alba, Costa Rica; Schedl Coll.. present designation). Cnesinus atrodeclivis Wood, 1968, Great Basin Nat. 28:108 (Holotype, female; Zamorano, Morazan, Honduras; Wood Coll.). New synonymy Schedl named adustus from a syntypic series of females. The specimen in his collection labeled "Holotype" is here designated as the lectotype of the species. This lectotype was compared directly to the holot}'pe of atrodeclivis Wood; only one species is represented by this material. Corthylocurus mexicanus (Schedl), n. comb. Brachyspartus mexicanus Schedl, 1950, Dusenia 1:163 (Holotype, male; Comitan, Chiapas, Me.xico; Schedl Coll.). Corthylus cylindricus Schedl, 1963, Ent. Arb. Mus. Frey 14:164 (Holotype, female; Jalapa, Veracruz, Mexico; Schedl Coll.). New synonymy Several long series of this species were taken in Mexico, Guate- mala, Costa Rica, and Panama. A male and a female in my collec- tion taken with a series from the same branch were compared direct- ly to the holotypes of Brachyspartus mexicanus Schedl and Corthylus cylindricus Schedl and were identical in all respects. Since these two names apply only to the opposite sexes of the same species, the name cylindricus must be placed in synonymy. Corthylus collaris Blandford Corthylus collaris Blandford, 1904, Biol. Centr. Amer. Coleopt. 4(6): 261 (Holo- type, male; Cerro Zunil. Guatemala; British Mus. Nat. Hist.). Corthylus splendens Wood, 1967. Great Basin Nat. 27:138 (Holotype, female; 16 km SE Cartago, Cartago. Costa Rica; Wood Coll.). New synonymy The holotypes of collaris Blandford and splendens Wood were examined. In addition, 43 specimens from Guatemala and 109 specimens from Costa Rica were studied. The northern and southern material can be distinguished by the small differences outlined in the description of splendens. However, in the absence of material from intermediate areas, it appears advisable to place splendens in sy- non>Tny initil the existence of geographical races can be more con- clusively demonstrated. Corthylus coniatus Blandford Corthylus coniatus Blandford. 1904, Biol. Centr. Amer.. Coleopt. 4(6):258 (Holo- type, female; Cerro Zunil. Guatemala; British Mt s. Nat. Hist.). Corthylus splendidus Bright, 1972. Canadian Ent. 104:1371 (Holotype, female; 5.6 km or 3.5 miles S Suchixtepec. Oaxaca, Mexico; Canadian Nat. Coll.). New synonymy 280 GREAT BASIN NATURALIST Vol. 34, No. 4 The holotypes of comatus Blandforcl and splendidus Bright were compared to my material from Mexico, Guatemala, Costa Rica, and Panama. I am unable to find any character that provides a means of subdividing this common species; consequently, the name splen- didus is placed in synonymy. Corthylus uniseptis Schedl Corthylus uniseptis Schedl, 1961, Pan Pacific Ent. 34:229 (Holotype, male; Cordoba. Veracruz, Mexico; California Acad. Sci.). Corthylus reburrus Bright, 1972, Canadian Ent. 104:1375 (Holotype, female; Palenque Ruins, Chiapas, Mexico; Canadian Nat. Coll.). New synonymy The male holotype of uniseptis Schedl was compared directly to a topotypic male paratype of reburrus Bright. Except for the slightly lighter color of the dark brown uniseptis type I see no difference between the two specimens. Since the slight degree of color differ- ence has no taxonomic value in this section of the genus, only one species is represented by these names. Dendrocranulus carhonarius (Ferrari), n. comb. Xylocleptes carhonarius Ferrari, 1867, Die Forst- und Baumzuchtschadlichen Borkenkater, p. 41 (Holotype, male; Cuba; Vienna Mus.). Xylocleptes floridensis Hopkins, 1915, U.S. Dept. Agric. Rept. 99:43 (Holotype, female; Biscayne Bay. Florida; U.S. Nat. Mus.). New synonytny Xylocleptes anonae Hopkins, 1915, U.S. Dept. Agric. Rept. 99:43 (Holotype, female; Florida; U.S. Nat. Mus.). New synonymy The holotypes of carhonarius (Ferrari), floridensis (Hopkins), and anonae Hopkins were compared directly to my series from Cuba. Only one species is represented by these names. Dendrocranulus maurus (Blandford) Dryocoetes maurus Blandford, 1898, Biol. Centr. Amer., Coleopt. 4(6): 191 (Holotype. female; El Tumbador, San Marcos. Guatemala; British Mus. Nat. Hist.). Dendrocranulus huehuetanus Schedl, 1940, An. Esc. Nac. Cienc. Biol.. Mexico 1:344 (Lectotype, male; Huehuetan, Chiapas, Mexico; Schedl Coll., present designation). New synonymy The male syntype from Huehuetan that was labeled by Schedl as the male "type" is here designated as the lectotype of Dendro- cranulus huehuetanus Schedl. That lectotype was compared directly to my series of maurus (Blandford) which was compared directly to the holotype of this species in 1965. Since they represent the same species, Schedl's name must be placed in synonymy. Dendroctonus valens LeConte Dendroctonus valens LeConte. 1860, Pacific R. R. Expl. 5(2) :59 (Holotype. male; San Francisco, California; Mus. Comp. Zool.). Dendroctonus rhizophagus Thomas and Bright. 1970, Canadian Ent. 102:479 (Holotype, male; 16 km or 10 miles SW El Salto, Durango, Mexico; Canadian Nat. Coll.). New synonymy Dec. 1974 wood: American scolytidae 281 The name rhizophagus Thomas and Bright was proposed for a long series of small specimens taken from the roots of pine seedlings during a severe drought near El Salto, Durango, in 1964. In 1965, I was unable to find specimens in seedlings at the same locality. All of the Dendroctonus taken in the area by me were valens LeConte of normal or near normal size. As with other Mexican valens, the El Salto specimens may construct parental tunnels that are linear, of the cave type, or a combination of these, and the frontal area of the head may be quite different from that seen in specimens from the United States and Canada. After examining the holotype and several paratypes of rhizophagus, the holotype and several thousand other specimens of valens. including several hundred from Mexico, Guate- mala, and Honduras, I am able to recognize only one species. There is a certain amount of variability in the frons in all areas, but these intergrade to such a degree that geographical races are difficult if not impossible to characterize. Gymnochilus consocius (Blandford), n. comb. Problechilus consocius Blandford, 1897. Biol. Centr. Amer., Coleopt. 4(6): 171 (Holotype. male; Cerro Zunil. Guatemala; British Mus. Nat. Hist.). Problechilus trimaculatus Schedl, 1935, Arch. Instit. Biol. Veget. 2:91 (Holotype, female; Venezuela, probably Colonia Tovar; Schedl Coll. ). New synonymy The holotypes of consocius Blandford and trimaculatus Schedl were compared to 102 specimens of this species from Venezuela. Approximately 90 percent of the specimens match the characters of consocius very well. In the type of trimaculatus the discal striae are not at all impressed and the strial punctures are much smaller and less strongly impressed than expected and the color is more nearly reddish brown. About 10 percent of my Venezuelan material ex- hibits variations that completely bridge the gap in characters between the extremes. For this reason, Schedl's name must be placed in synonymy. Gymnochilus minor (Blandford), n. comb. Problechilus minor Blandford. 1897, Biol. Centr. Amer. Coleopt. 4(6): 172 (Syntypes; Guatemala City. Guatemala; British Mus. Nat. Hist.). Problechilus varius Schedl, 1951. Dusenia 2:86 (Holotype, female; Mexico; Schedl Coll.). New synonymy Eight syntypes of minor (Blandford) and the holotype of varius Schedl were examined and compared to several series in my collec- tion. The holotype of varius is a typical female of this species. Gynmochilus zonatus Eichhoff Gymnochilus zonatus Eichhoff. 1867. Berliner Ent. Zeitschr. 11:399 (Holotype. sex?; Colombia; lost with Hamburg Mus.). Meringopalpus fallax Hagedorn. 1905. Bull. Mus. Paris 10:547 (Holotype. male; Venezuela, by Moritz, presumably Caracus or Colonia Tovar; Paris Mus.); Eggers, 1932, Wiener Ent. Zeit. 49i:226. Synonymy 282 GREAT BASIN NATURALIST Vol. 34, No. 4 Problechilus freyi Schedl, 1966, Ent. Arb. Mus. Frey 17:103 (Holotype, female; Rancho Grande near Maracay, Aragua, Venezuela; Frey Mus.)- I^ew synonymy Eggers compared the holotypes of Gymnochilus zonatus Eichhoff and Meringopalpus fallax Hagedorn and concluded that they were synonymous. Since the type of zonatus (3.3 mm) was subsequently lost in the destruction of the Hamburg Museum, Eggers's observa- tion is accepted as correct. The male holotype of fallax (3.2 mm) and the female holotype of freyi (Schedl) (2.8 mm) were examined and were found to represent the same species. This species is almost identical to consocius (Blandford), but it is distinguished by the slightly larger size, by the less strongly impressed male frons, by the more strongly convex female frons, by the reticulate surface among the asperities on the anterior half of the pronotum, by the more coarsely punctured basal fourth of the pronotum, and by the slightly longer elytral vestiture. Hypothenernus areccac (Hornung) Bostrichus areccae Hornung. 1842, Stett. Ent. Zeit. (Lectotype, female; in Betel palm nuts presumably of East Indian origin; Berlin Zool. Mus., present designation). Hypothenernus vafer: Wood, 1972. Great Basin Nat. 32:52. As indicated in the above reference to my recent partial review of the synonymy of this species, there has been much confusion con- cerning the identity of areccae Hornung. The confusion was created by Eggers when he labeled specimens of both areccae and eruditus Westwood as having been compared to the "type." Two syntypes of of areccae in the Berlin Zoologisches Museum are of the species I have referred to as areccae ( = vafer Blandford, fungicola Eggers, hispidus Eggers, etc.). The first of those syntypes has the impressed frons partly obscured by frass; therefore, the second syntype is here designated as the lectotype of areccae Hornung, as indicated above. This species is common in seeds and fruits, although it is also able to breed in twigs, bark, and other materials. Apparently it is native to the Indo-Malayan region; eruditus is less common in seeds and fruits and was not introduced into the Indo-Malayan region until about a half century after areccae was described. Hypothenernus eruditus Westwood Hypothenernus eruditus Westwood, 1836, Trans. Ent. Soc. London 1(1):34 (Syntypes; England). Cryphalus obscurus Ferrari, 1867. Die Forst- und Baumzuchtschiidlichen Borken- kafer. p. 17 (Holotype, female; Cuba; Vienna Mus.). New synonymy Although Cryphalus obscurus Ferrari has been considered a junior homonym of a Fabricius species, the species has never been clarified. The female holotype was examined. It has the entire frontal area immersed in glue, but other features clearly indicate that it is a normal specimen of eruditus V^estwood. Dec. 1974 wood: American scolytidae 283 Micracis grandis Schedl Micracis grandis Schedl, 1948, Rev. de Ent. 19:575 (Holotype. female; San Jose de Ixtepec, Chiapas, Mexico; Schedl Coll.). Micracis coslaricensis Wood, 1969, Brighani Young Univ. Sci. Bull., Biol. Ser. 10(2) :39 (Holotype, female; Volcan Poas, Heredia, Costa Rica; Wood Coll.). New synonymy The female holotypes of grandis Schedl and costaricensis Wood were compared du'ectly to one another and to series from Honduras and Costa Rica. Only one species is represented by this material. Micracis swainei Blackman Micracis swainei Blackman, 1920, Mississippi Agric. Expt. Sta. Tech. Bull. 9:32 (Lectotype, female; luka, Mississippi; U.S. Nat. Mus., designated by Wood, 1963, Great Basin Nat. 33:178). Micracis robustus Schedl. 1948. Rev. de Ent. 19:576 (Holotype, male; Esmeralda, Chiapas, Mexico; Schedl Coll.). New synonymy Micracis pygamaeus Schedl, 1948, Rev. de Ent. 19:577 (Holotype, male; Hue- huetan, Chiapas, Mexico; Schedl Coll.). New synonymy The male holotypes of robustus Schedl and pygamaeus Schedl were compared directly to male specimens from the same series as my homotypes of swainei Blackman; they are of the same species. More than 500 specimens of this species from New York to Hon- duras were examined. The holotype of robustus is of normal size (1.8 mm) and is badly rubbed; the holotype of pygamaeus is near the lower limits of size (1.4 nnn) and has the vestiture in good con- dition. The "second specimen" of pygamaeus, mentioned in the original description, is a female with the epistomal brush of setae that is characteristic of this species. Microcorthylus parvulus Ferrari Microcorthylus parvulus Ferrari, 1867. Die Forst- und Baumzuchtschadlichen Borkenkafer. p. 58 (Holotype, male; Venezuela, probably Colonia Tovar; Vienna Mus.). Pterocyclon exile Eichhoff, 1878, Mem. Soc. Roy. Sci. Liege (2) 8:451 (Holotype, male; Nova Grenada; Brussels Mus.). New synonymy The male holotypes of parvulus Ferrari and exile (Eichhoff) were examined and compared directly to my material. In all, 159 specimens from Costa Rica to Colombia and Venezuela were ex- amined. Following a detailed study of the genus from these and neighboring area, it was concluded that only one species was repre- sented by this material. Minute variation occurs on the frons; other characters are constant. Monarthrum bicolor (Ferrari), n. comb. Corthylus bicolor Ferrari, 1867, Die Forst- und Baumzuchtschiidlichen Borken- kafer, p. 56 (Holotype. female; Venezuela, probably Colonia Tovar; Vienna Mus.). Corthylus signalus Ferrari. 1867. Die Forst- und Baumzuchtschiidlichen Borken- kafer. p. 56 (Syntypes, male; Venezuela, probably Colonia Tovar; Vienna Mus.). New synonymy 284 GREAT BASIN NATURALIST Vol. 34, No. 4 Pthorius edentatus Hagedorn, 1905, Bull. Mus. Paris 10:549 (Holotype, male; Colonia Tovar, Venezuela; Paris Mus.). New synonymy The holotypes of bicolor (Ferrari) and edentatus (Hagedorn) and the two male syntypes of signatus Ferrari were examined and compared directly to my material. The female frons is very distinc- tive, thus making this species easily identified. It is common at high elevations in Colombia and Venezuela. Since bicolor and signatus were taken in Venezuela in 1858 by Moritz, the type localities ap- parently are at or near the Moritz home at Colonia Tovar, Aragua, Venezuela. The male declivity varies slightly in certain series in its steepness and in the abruptness of the basal and lateral margins. Blandford's label on the syntypes of signatus erroneously identified them as Monarthrum chapuisii Kirsch. Monarthrum bicoloratum, n. n. Monarthrum bicolor Wood, 1968 (nee. Ferrari, 1867), Great Basin Nat. 28:4 (Holotype, male; Mile 10 on the Bartica-Potaro Road, British Guiana; British Mus. Nat. Hist.). The transfer of Corthylus bicolor Ferrari to Monarthrum (above) made a junior homonym of Wlonarthrum bicolor Wood. The new name bicoloratum is proposed as a replacement name for bicolor Wood. Monarthrum consimile (Blandford), n. comb. Pterocyclon consimile Blandford, 1904, Biol. Centr. Amer., Coleopt. 4(6):275 (Holotype, female; Volcan de Chiriqui, Panama; British Mus. Nat. Hist.). Pterocyclon pseudosulcatum Sc.hedl, 1935, Rev. de Ent. 5:348 (Holotype, male; Vara Blanca, Heredia, Costa Rica; Schedl Coll.). New synonymy Series of specimens containing both males and females from the type localities of both consimile (Blandford) and pseudosulcatum (Schedl) were collected and compared to the holotypes of these species. The holotype of consimile is the female, and pseudosulcatum is the male of the same biological species; consequently, Schedl's name is placed in synonymy. Monarthrum egenum (Blandford), n. comb. Pterocyclon egenum Blandford, 1904. Biol. Central Amer.. Coleopt. 4(6):280 (Holotype, female; San Juan, Verapaz, Guatemala; British Mus. Nat. Hist.). Brachyspartus bisetosus Schedl, 1954, Dusenia 5:38 (Syntypes, female; Rio Caraguata, Matto Grosso, Brazil; Schedl Coll., etc.). New synonymy This species is common in Inga and other leguminous trees from Guatemala to Brazil. Females from Costa Rica were compared to the holotype of egenum (Blandford), to two topotypic female syntypes of bisetosus (Schedl), and to several series from Costa Rica, Colombia, and Brazil. Only one species is represented by this material. Schedl's males of bisetosus actually are females with frontal setae; his females lack part or all of these setae. Dec. 1974 wood: American scolytidae 285 Monarthrum fimbraticorne (Blaiidford) Pterocyclon fimbraticorne Blandford, 1904, Biol. Centr. Amer. Coleopt. 4(6):285 (Holotj-pe, female; Purula, Verapaz, Guatemala; British Mus. Nat. Hist.). Pterocyclon turbinatum Schedl, 1961, Pan Pacific Ent. 37:230 (Holotype, male; Cordoba, Veracruz, Mexico; California Acad. Sci.). New synonymy The female holotype of fimbraticorne (Blandford) was compared to representatives of several series of this species from Costa Rica. Additional long series have been taken in Venezuela and one addi- tional specimen was seen from Guatemala. Males of these series were compared directly to the male holotype of turbinatum (Schedl). The type of turbinatum is rather small for this species, but males of similar small size occur throughout the range of this species. Monarthrum gnarum (Schedl), n. comb. Pterocyclon gnarum Schedl, 1950. Dusenia 1:169 (Holotype, female; Mexico; Schedl Coll.). Amphicranus spinatus Bright, 1972, Canadian Ent. 104:1383 (Holotype. male; 15 km or 32 miles S Valle Nacional. Oaxaca, Mexico; Canadian Nat. Coll.). New synonymy The holotype of Pterocyclon gnarum Schedl is a female, not a male as stated in the original description. This holotype was com- pared to a topotypic female paratype of Amphicranus spinatus Bright and to another female from Hidalgo. The holotype, allotype, and other para types of spinatus were also examined. It is now clear that all represent one species that is very closely related to praerup- tum (Blandford). Monarthrum validum (Ferrari), n. comb. Corthylus validus Ferrari, 1867, Die Forst- und Baumzuchtschadlichen Borken- kafer, p. 55 (Lectotype, male; Mexico; Vienna Mus.. present designation). Amphicranus mexicanus Eggers. 1931, Ent. Blatt. 27:18 (Holotype, male; Mexico; Berlin Zool. Mus.). New synonymy Pterocyclon jalapae Schedl, 1939, Mitt. Miinchn. Ent. Ges. 29:584 (Holotype, male; Jalapa, Veraci-uz, Mexico; Schedl Coll.). New synonymy The name Corthylus validus Ferrari was based on a male from Mexico, and a female from Venezuela. Blandford (1904, Biol. Centr. Amer., Coleopt. 4(6): 271) transferred the female to another species. The male is here designated as the lectotype of validus. This lecto- type and the male holotype of Amphicranus mexicanus Eggers were compared directly to my series from Costa Rica. Only one species is represented by this material; it occurs from Mexico to Panama. Later, my Costa Rican specimens were compared directly to the holotype of Pterocyclon jalapae Schedl and were also found to be identical. Phloeoborus scaber Erichson Phloeoborus scaber Erichson, 1836, Archiv. Naturgesch. 2(1):55 (Syntypes; Brazil; Berlin Zool. Mus.). Phloeoborus opacithoraz Schedl. 1940. Arb. Morph. Taxon. Ent. Berlin-Dahlem 286 GREAT BASIN NATURALIST Vol. 34, No. 4 7:205 (Syntypes; Panzos, Guatemala; Institut fiir Pflanzenschutzforschung Kleinmachnow). New synonymy The female syntypes of opacithorax Schedl have the eyes slightly more narrowly spaced and the elytral vestiture stouter than do the types of scaber Erichson. The spacing of the eyes varies throughout the range of this species. The elytral vestiture becomes longer and stouter over a gradual cline from northern South America to southern Mexico, although there is considerable variation within series; and the elytral crenulations also become narrower, more numerous, and confused over the same cline. While Brazilian and Guatemalan specimens may be rather strikingly different, intergradation is such that distinctive races cannot be separated. Phloeosinus serratus (LeConte) Hylesinus serratus LeConte, 1868, Trans. Amer. Ent. Soc. 2:170 (Holotype, male; Middle States; Mus. Comp. Zool.). Phloeosinus rugosus Swaine, 1917, Dom. Canada Dept. Agric. Ent. Br. Tech. Bull. 14(1 ):9 (Lectotype, female; Scaffold Meadow, Sequoia N.F., California; Canadian Nat. Coll., 9259, designated by Bright, 1967, Canadian Ent. 99:677). New synonymy The female lectotype of rugosus Swaine lacks tubercles on de- clivital interstriae 2; it is also rather small (2.6 mm). The two fe- male paratypes in the Canadian National Collection both have tubercles on declivital interstriae 2. It is clearly evident that all three specimens fall within the range of variation of serratus (LeConte); consequently, rugosus must be placed in synonymy under the older name. Phloeotribus setulosus Eichhoff Phloeotribus setulosus Eichhoff, 1868. Berliner Ent. Zeitschr. 12:149 (Lectotype, male; Colombia; Brussels Mus., designated by Wood, 1973, Great Basin Nat. 33:182). Phloeetribus rudis Eichhoff, 1868, Berliner Ent. Zeitschr. 12:149 (Syntypes?: male; Brazil; lost with Hamburg Mus.?). New synonymy Phloeotribus sodalis Blandford, 1897, Biol. Centr. Amer.. Coleopt. 4(6): 168 (Lectotype, male; Cerro Zunil, Guatemala; British Mus. Nat. Hist., present designation) . New synonymy Phloeotribus bolivianus Eggers, 1933, Trav. Lab. d'Ent. Mus. Nat. d'Hist. Nat. Paris, Mem. 1:5 (Holotype, male; Cochabamba, Bolivia; U.S. Nat. Mus.). New synonymy The holotypes of setulosus Eichhoff and bolivianus Eggers, the lectotype of sodalis Blandford, and two specimens of rudis Eichhoff that were identified b}- Eichhoff and deposited in the Chapuis collec- tion were all compared directly to series of this species in my collec- tion and were found to represent one species. The two specimens of rudis probably are unmarked syntypes and evidently represent the only reasonably authentic representatives of rudis in existence. The first specimen in Blandford's series, a male from Cerro Zunil, is here designated as the lectotype of sodalis Blandford. This specimen was previously labeled as the type, but it was never so designated. Dec. 1974 wood: American scolytidae 287 Pycnarthrum reticulatum Schedl Pycnarthrum reticulatus Schedl. 1940, An. Esc. Nac. Cienc. Biol., Me.xico 1:355 (Lectotype, female; Tonala. Chiapas, Mexico; Schedl Coll., present desig- nation). Pycnarthrum fici Wood. 1971. Brigham Young Univ. Sci. Bull., Biol. Ser. 15(3): 11 (Holotype, male; 5 km W El Pino, Merida, Venezuela; Wood Coll.). New synonymy A female of reticulatus [sic] Schedl in the Schedl collection, from Tonala, Chiapas, has been labeled as the "Type" of this species by its author, although it has never been so designated. I here desig- nate that female as the lectotype of reticulatus. This lectotype was compared directly to the holotype of fici Wood. Only one species is represented by this material. Scolytodes rugicollis (Schedl), n. comb. Hexacolus rugicollis Schedl, 1940, Arb. Morph. Taxon. Ent. Berlin-Dahlem 7:205 (Lectotype, female; Hamburgfarm on Rio Reventazon, Limon, Costa Rica; Schedl Coll., present designation). Scolytodes plicatus Wood, 1969, Brigham Young Univ. Sci. Bull., Biol. Ser. 10 (2): 21 (Holotype, female; 25 km SE Guapiles, Limon, Costa Rica; Wood Coll.). New synonymy Schedl named Hexacolus rugicollis from a syntypic male and a female mounted together on the same microcard. In the original description the sexes were reversed. I designate the female synt5'pe, marked on the microcard by a male symbol, as the lectotype of Schedl's species. This lectotype was compared directly to the holo- type of Scolytodes plicatus Wood and was found to represent the same species. Xylosandrus morigerus (Blandford) Xylehorus morigerus Blandford. 1894, Insect Life 6:264 (Syntypes; intercepted in England from Dendrobium orchids imported from New Guinea; British Mus. Nat. Hist.). Xyleborus luzonicus Eggers, 1923, Zool. Meded. 7:174 (Lectotype, female; Mt. Makiling, Insel Luzon, Philippinen; U.S. Nat. Mus.; designated by Anderson and Anderson, 1971, Smithsonian Contrib. Zool. 94:18). New synonymy Six syntypes of morigerus (Blandford) were compared directly to my specimens; these were later compared directly to the lectotype of luzonicus Eggers. Only one species is represented by this material. Araptus sobrinus, n. sp. This species is distinguished from schwarzi (Blackman) by the weakly convex, more sparsely punctured female frons, by the dif- ferent elytral vestiture, and by the more deeply, more broadly sul- cate declivity. Female. — Length 2.3 mm (i)aratyj:)es 1.9-2.3 mm), 2.5 times as long as wifle; color reddish brown. Frons weakly convex; surface almost smooth, punctures fine, moderately abundant; vestiture of fine, short, moderately abundant 288 GREAT BASIN NATURALIST Vol. 34, No. 4 hair; frons about as in male schwarzi except for epistomal area. Pronotuni about as in schwarzi, except disc with more numerous impressed points and punctures slightly larger and more nearly circular (punctures in schwarzi oval to crescent-shaped) . Elytra as in schwarzi except with abundant impressed points (usually absent in schwarzi)^ declivity more deeply, more broadly impressed, and vestiture greatly reduced. Strial setae entirely absent, interstrial setae rare on disc, sparse on declivity except at sides. (In schwarzi interstrial rows of erect setae usually extend to elytral base and small strial hairs occur on disc and declivity.) Male. — Similar to female, except frons more strongly convex above, slightly impressed on lower half; epistoma broadly, shallowly emarginate about as in male schwarzi. Type Locality. — Siguatepec, Honduras. Type Material. — The female holotype, male allotype, and 10 para types were taken at the type locality on 25-V-1972, in rust cones on Pinus oocarpa^ by R. Billings. The holotype, allotype, and paratypes are in my collection. Dendroctonus mexicanus Hopkins This species was placed in synonymy (Wood, 1963, Great Basin Nat. 23:41) under Dendroctonus frontalis Zimmermann, because series of this species from the southwestern United States could not be distinguished from variable series from Honduras. When it was recently noticed that two separate emergences took place in Hon- duras from the same logs, it became apparent that two distinct bio- logical species occur in Honduras and that both were responsible for the epidemic of 1964. Once this event was apparent, and pure sam- ples from each emergence were studied, the variable Honduras ma- terial was easily sorted into two groups, one having poorly developed to obsolete interstrial crenulations (frontalis) and a slightly larger, darker form with moderate to rather coarse interstrial crenulations. The former, frontalis, tends to occur at low elevations, less than 700 m, the latter at elevations above 700 m. Thev may occur in the same trees in the area of altitudinal overlap. The latter form appears to be of the same species as occurs throughout Mexico; consequently, the name mexicanus is removed from synonymy to designate it. Speci- mens from Arizona and New Mexico appear to be intermediate but are tentatively grouped with frontalis. Ultimately it may be neces- sary to reduce mexicanus to subspecific rank, due to intergradation in the northern area, even though the two populations behave as valid species in Honduras. In 1970 I received specimens of an additional Guatemalan species in the frontalis complex, from E. W. Clark. Since then Dr. .1. P. Vite and his associates have investigated Mexican and Central American Dendroctonus more thoroughly. Their results are being published elsewhere. It appears that mexicanus originally extended Dec. 1974 wood: American scolytidae 289 only as far south as Chiapas and was replaced in (juatemala b}- intei, described below. Both mcxicanus and frontalis appear to have been introduced into Honduras at a conii)aratively recent date, where they jointly caused ei)idt^niic losses of pine a decade ago. Apparently neither species has penetrated the range of vitei. Dendroctonus vitei, n. sp. This species is distinguished from me.ricanus Hopkins by the slightly larger average size, by the much darker color, and by other differences cited below. The most reliable characters for distinguish- ing it are the pronotal granulation, the declivital characters, the frons, and the seminal rod. Male. — Length 3.4 mm (paratypes 2.6-4.1 mm) (exclusive of head), 2.2 times as long as wide; color almost black. Frons similar to mcxicanus, except epistomal process wider (67 percent of epistomal width, 58 percent in mexicanus) , its lateral arms much more strongly elevated; area from epistomal process to lateral summits on frons more strongly, more broadly impressed (often subconcave), more finely sculptured; largest tubercles at and near lateral summits less numerous, smaller than in mcxicanus. Pronotum similar to that of mcxicanus, except punctures averag- ing much smaller, closer, interspaces averaging less than half diameter of a puncture; interspaces over most of surface minutely granular, subshining, smooth shining areas usually restricted to less than one-fifth of total surface (in mcxicanus interspaces average more than half diameter of a puncture, their surface smooth, brightly shining over almost entire surface, granulation rare and restricted; small tubercles sometimes occur in lateral areas). Vestiture more uniformly distributed, slightly coarser and longer than in mcxicanus. Elytra similar to those of mcxicanus. except discal striae less strongly impressed, punctures averaging smaller, obscurely impressed to obsolete in most specimens (larger and rather distinctly impressed in most specimens of mcxicanus) ; interstrial crenulations narrower, lower, more numerous, rarely as much as one-third width of an inter- striae (in mcxicanus about one-third of crenulations at least half as wide as an interstriae, a few extend entire width of an inter- striae). Declivity with striae feebly if at all impressed, punctures usually obsolete (striae and pimctures rather strongly imjirossed in mcxicanus): interstriae feebly if at all convex, crenulations less numerous and much smaller than in mcxicanus. Vestiture slightly more abundant and coarser than in mcxicanus. particularly at sides. Female. — Similar to male, except epistomal process narrower, not as high, frontal summits more poorlv developed, usually not tuberculate; pronotal callus more poorly developed than in female mexicanus. Type Locality. — Patzun. Guatemala. Type Material. — The male holotype. female allotvpo. and 26 paratypes were collected at the type locality on 19-IX-1974. from 290 GREAT BASIN NATURALIST Vol. 34, No. 4 Pinus tenuifoUa, by J. P. Vite. Two hundred and sixty paratvpes all with their abdomens dissected bear the same data except they were taken 22-V-1974; 50 paratvpes are from the same locality taken 9-X-1974 from "P. maxirninoi" by R. Liihl. Ten parat^'pes are from the same locality, taken on 12-III-1974, and four paratypes are from the same locality, taken on 14-X-1973, by .1. P. Vite fall dis- sected). Three paratypes are from Puente Tzantzir, Sololn. Guate- mala, 2-II-1972, from Pinus mnntezuma, by E. W. Clark. Vite and his associates are studying the biology and liehavior of the frontalis complex and adding significant information about these species (Vite, Islas, Renwdck, Hughes, and Kliefoth, 1974. Zeit. Angew. Ent. 75:422-435). Thev will report additional biological and biochemical characters of this species. Their illustration of the male seminal rod (Fig. 2E on p. 426) is significant. In June 1974 it was discovered that both Vite and I had inde- pendently prepared a description of this species. He kindly con- sented to withdraw his manuscript. This species is named in recog- nition of his numerous contributions to our understanding of these important insects and for the independent discovery of this species. DESCRIPTION OF NEW SPECIES OF MILODERES CASEY, WITH COMMENTS ON OTHER SPECIES OF FHE GENUS (COLEOPTERA: CURCULIONIDAE) Vasco M. Tanner' Abstract. — Miloderes allredi, from Utah, and M. tingi, from California, are described as new to science. Miloderes allredi, n. sp. Figs. 1-5 Derm black, clothed with bluish-green, iridescent, densely placed scales; side of prothorax and elytra with long brownish setae, disc of prothorax and elytra with sparse shorter setae. Rostrum continuous, with no transverse impression; apex one-half width base of head. Origin of scrobes near apex of rostrum, well developed and extending to lower base of eye. Antenna brown, with setae and scales; scape reaching to middle of eye; segments 1 and 2 of funicle elongate, as as long as segments 3-6 combined. Segment 7 asymmetrical, cuplike widest on outer margin, clavate at apex. Mentum large, flat, wider than long, filling entire gular cavity. Eyes small, vertical; row of scales between eye and vibrissae of prothorax. Prothorax widest at anterior third, sides arcuate, strongly convergent toward base; base well separated from elytra; postocular lobes small, with developed vibrissae; sides and disc punctate, scales compact, setae short and sparse on disc; apex slightly constricted. Scutellum obscure. Elytra widest at basal fourth; sides feebly arcuate, rounded behind, pos- terior declivity perpendicular; disc punctate, with sparse short brown setae, a mixture of compact blue, green, and iridescent scales. Abdomen, ventrites and legs clothed with scales and setae similar to those on dorsal areas of body; ventrite 1 at midline as long as 2 and 3 combined. Metathoracic tibia wdth corbel open, margin with amber-colored row of short spines. Prothoracic tibia corbel open and with an outward projection of the distal portion of tibia. Male spermatheca and female genitalia distinctive; related to setosus Ting, 1940. Length. — 5.0-5.8 mm; breadth: 2.5-3.0 mm. Type locality. — Cotton Bench, Glen Canyon City, Kane Coun- ty, Utah. Type specimens. — Male holotype and female allotype in the entomological type collection at Brigham Young University; one paratype in the entomological collection, U.S. Natural Museum. Specimens of this species were collected by Dr. Dorald M. Allred and assistants in May and .Tune 1973, while collecting plant and animal species of the Lake Powell area in connection with the En- 'Departmetit of Zoology, Brigham Young University, Provo, lUah 84(i02. 291 292 GREAT BASIN NATURALIST Vol. 34, No. 4 vironmental Monitoring Project in relationship to Navajo Power Plant Project. Specimens dealt with above were collected on the sand dunes, in an association of Ephedra viridis Coville and Yucca bailey i Woot. Standi. I am pleased to name this striking species in honor of Dr. Dorald M. AUred, Professor of Zoology at Brigham Young University. Dr. Allred is a capable field worker who has devised many means of collecting arthropods and is a leader in environmental studies. Figs. 1-5. Miloderes allredi: 1, dorsal view of adult female; 2-3, ? and 5 genitalia; 4, spermatheca; 5, front and hind tibia. Dec. 1974 tanner: new curculionidae 293 In making this study, the genitalia of several of the species of Miloderes have been compared. Drawings of tibia, male, female genitalia, and spermatheca of five species are included in this study. Miloderes tingi, n. sp. Fig. 6-9 Derm dark chestimt brown to black, with small, irregular- shaped, rather compact, ash-grey scales that clothe legs, head, pro- thorax, and dorsal and ventral parts of body, each scale with central puncture; long grey setae on sides of prothorax, sides and posterior dorsal sides of elytra. Rostrum continuous with head, with slight transverse impression, apex one-half width base of head. Scrobes shallow, extending to well below base of eye. Antennae reddish brown, scape slender, first segment of funicle enlarged, as long as segments 2 and 3 combined, club large and setiferous; prementum wider than long, with few short setae; eye ovate. Prothorax slightly wider than long; widest at middle, sides arcuate, weakly convergent toward base, moderately convex; sides covered with dense whitish scales and long setae; dorsally tuberculate, punctate, with few scales. Scute! I um and postocular well developed; postocular lobes small, with well-developed vibrissae, apex slightly constricted. Elytra widest at anterior fourth, sides feebly arcuate, rounded behind, pos- terior declivity perpendicular; disc punctate, scales covering surface, with scattered long setae on ])Osterior half of disc, scales with a cen- ter puncture. Ventrites covered with scales and setae; ventrite 1 as long as 2 and 3 combined, 2 as long as 3 and 4 combined. Metathor- racic tibia open, with a row of eleven amber-colored spines. Size, shape, and number of tibial spines and tibia distinctive. Prothoracic tibia with distal portion spatulate, 10 spines (Fig. 7). Spermatheca and genitalia distinctive (Fig. 8-9). Length.- — 7 mm; breadth: 3.6 mm. HoLOTYPE. — Female, deposited in type collection, Brigham Young University. Type locality. — Essex, San Bernardino Co., California. 29-IV- 1937, Collectors: P. T. Ting, M. Cazier. M. tingi is uniformly a dark brownish species, clothed with small dense greyish scales; head and prothorax tuberculate, no traces of striation on elytra, and devoid of setae on the central areas of pro- thorax and elytra. The female genitalia and spermatheca differ from similar structures of other sj)ecies of this genus. This species is closely related to M. setosa Csy; however, it is more robust with smaller scale punctation on prothorax, and the female genitalia and spermatheca are different (Figs. 8-9). I am pleased to name this species in honor of Peter T. Ting who contributed much to our knowledge of this group of Brachyrhininae weevils (1940). Edwin C. Van Dyke and David G. Kissinger have done much to bring about an orderly arrangement and understand- ing of this interesting western-American weevil fauna. 294 GREAT BASIN NATURALIST Vol. 34, No. 4 The keys to the genera and species of this group as contained in Ting's paper (1940) will be useful, in combination wdth the descrip- tion and illustrations of this paper, in separating the species of Figs. 6-9. Miloderes tingi: 6, dorsal view adult female; 7, front and hind tibia; 8, 9 genitalia; 9, spermatheca. Dec. 1974 TANNER: NEW CURCULIONIDAE 295 Fig. 10: Miloderes nelsoni Kissinger, front and hind tibia. Fig. 11: M. setosus Casey. Fig. 12: M. nelsoni. $ genitalia. Fig. 13: M. mercuryensis Tan- ner, 5 genitalia. Fig. 14: M. setosus Csy., 9 genitalia. Fig. 15: mercuryensis Tanner, front and hind tibia. Fig. 16: M. setosus Csy. $ genitalia. 296 GREAT BASIN NATURALIST Vol. 34, No. 4 Miloderes. I have had the privilege of examining specimens of all six of the species now included in Miloderes. Unfortunately, I did not make drawings of the tibia of M. viridis Pierce. Dr. Elbert L. Sleeper has made a study of some species of Milo- deres, but I do not have access to his writings dealing with the species of this genus. He kindly contributed specimens of M. nelsoni Kissin- ger which I have reported on. I wish to express my thanks to Dr. Rose Ella Warner for her aid in this study and for loan of specimens of M. viridis and M. setosus from the U.S. National Museum and courtesies extended while I studied at the museum. Literature Cited Casey, T. L. 1888. On some North American Rhynchophora I. Ann. N.Y. Ac. Sci. pp. 252-254. Kissinger, D. G. 1960. Description of a new species of Miloderes Casey with notes on some broad-nosed weevils (Curculionidae). Coleopterists Bull. 14: 25-28. Pierce, W. D. 1909. Studies of North American weevils. Proc. U.S. Nat. Mus. 37:348. Tanner, V. M. 1966. Rhynchophora beetles of the Nevada Test Site. Brig- ham Young Univ. Sci. Bull., Biol. Ser. 8(2): 16. Ting, P. C. 1940. Revisional notes concerned with Cimbocera and related genera. Bull. So. Cal. Ac. Sci. 39:128-157. REVISION OF THE PLANT GENUS GERANIUM IN UTAH Glen T. Nebekci' Abstract.- - Within the state of Utah are seven species of Geranium, two of which are annual. The nature of the caudcx and the growth habit have been used to separate the perennial species. These characters are inadequate for separating the species. Better morphological characters, keys, and descriptions are presented. Two characters used to separate various species of Geranium are not effective. One of these characters is "plants somewhat cae- spitose." Use of this character in keys would lead to plants which are sometimes three feet tall and with the same growth habit of plants considered not caespitose. The other character is the branching or simple nature of the caudex. In herbarium specimens it is usually impossible to see the caudex. In addition, authors vary in the de- scription of the caudex; for example, G. richardsonii is described as: "caudex often slightly branched" (Hanks and Small, 1907), "the usually simple caudex" (Jones and Jones, 1943), "plants . . . erect from a simple caudex" (Harrington, 1959). The caudex actually varies from simple to very branched. (VanCott, 1969). These char- acters illustrate a need for a revision of the genus. Hanks and Small (1907) treated the known species of North America Geranium. In Knuth's (1912) worldwide monograph of the genus the treatment of the North American species was more or less copied from Hanks and Small's work. Jones and Jones (1943) treated the perennial species north of Mexico, and Moore (1943) included one Utah species in his coverage of the Mexican species. Taxonomic Characters As mentioned above, the nature of the caudex and the growth habit are poor taxonomic characters. Six taxonomically significant characters are described below. The variations of each of them were measured, assigned numbers, then averaged for each character and plotted on polygonal graphs. (Fig. 1) All observations were made under a dissecting microscope. Stylodia: These are the branches at the tip of the style column which bear the stigmatic surface. These were first soaked with Pohl- stoffe then measured with a standard mm ruler. Only mature flowers were measured. For each species the length was averaged. Petal pilosity. This is the pubescence on the upper surface of the petals. Three degrees of pilosity were recognized and assigned num- bers for comparison. The degree and corresponding numbers are: 1/4 of the petal covered = 1; V3 of the petal covered = 2; and i/^ of the petal covered = 3. 'Department of Botany and Range Science, Brigliam Younn University, Provo, Ulali 84602. 297 298 GREAT BASIN NATURALIST Vol. 34, No. 4 Fig. 1: 1, stylodia; 2, petal pilosity; 3, pubescence of pedicels; 4, pubescence of stems; 5, petals recurving or not; 6, petal color; A, G. atropurpurcum; B, G. fremontii; C, G. marginale; D, G. parryi; E. G. richardsonii. Pubescence of the pedicels-. The pedicels of some species are glandular pubescent, while others are pubescent but not glandidar. The color (purple or not) of the glands was also noted. The num- bers given these characters are: glandular = 1; purple glands = 2; and nonglandular = 3. Dec. 1974 nebeker: geranium revision 299 Pubescence of the stems: Pubesceiuo on the stems varied from very glandular to glabrous. The only significant difference in pube- scence was glandular = 1 ; nonglandular = 2. Petals recurving or not: The petals of most species extend out flat or curve upward slightly. One species has petals that curve downward abruptly. Numbers assigned to these are: petals not recurving ■= 1 ; petals recurving = 2. Petal color: White petals = 1 ; varying shades of pink to purple = 2. The above characters have been applied to the five perennial species that occur in the state. The characteristics used to separate the annual species are the number of fertile stamens and awned or nonawTied sepals. Shaw (1952) has worked out the cytology of four species that occur in Utah. The chromosome numbers that he reported are as follows: G. carolinianum 2n = 52; G. pusillum 2n = 26; G. richard- sonii n = 26; G. fremontii (which he called G. nervosum) n = 26. Other etiological work should be done because there are indications of hybridization betweeen G. richardsonii and G. fremontii. The probable phylogenetic relationships of the perennial species within the state are outlined on Figure 2 and their distributions are plotted on Figure 3. Taxonomy Geranium L. Sp. PI. 676 (1753) Annual or perennial herbs, often with a woody caudex; stems glabrous to glandular-villous; leaves palmately lobed, cleft or parted, basal leaves generally larger than the cauline ones; inflorescence compact to spreading; flowers complete, actinomorphic; sepals 5 usually awn tipped; petals 5, deciduous, purple to white, usually pubescent toward the base; ovary 5 lobed, 5 loculed with 2 ovules per locule becoming 1 seeded, elastically recoiling at maturity but not twisting. Key to the Species 1. Plants annual, petals less than 1 cm long 2 Plants perennial, petals more than 1 cm long 3 2(1). Sepals awnless, fertile stamens 5 1. G. pusillum Sepals awned, fertile stamens 10 2. G. carolinianum 3(1). Plants nonglandular (sometimes nonglandular in G. richardsonii) 4 Portions of the plant glandular 5 4(3). Petals reflexing at maturity, pilose on the petals ^3-/4 their length 3. G. atropurpureum Petals not reflexing at maturity, pilose on the petals 14 their length 4. G. marginale 300 GREAT BASIN NATURALIST Vol. 34, No. 4 5(3). Pedicels and lower portions of the plant glandular 5. G. parryi Lower portions of the plant nonglandular, pedicels glandular 6 6(5). Petals white, petals pilose Vs-V^ their length 6. G. richardsonii Petals purple, petals pilose 1/4 their length .... 7. G. fremontii 1 . G. pusillum Burm. f . Sp. Bot. Geran. 27 ( 1 759) Annual; stems 10-60 cm long, decumbent or prostrate, puberu- lent; leaves reniform to orbicular, 1-6 cm broad, 3-7 parted; sepals 2.5-5 mm long, awnless; petals purple to violet; 5 fertile stamens; style column 6-9 mm long, glandular puberulent; carpel bodies 2 mm long; seeds smooth. A weed of fields and waste places. Type locality, England and France. Provo Bench near Pleasantview, Utah Co., G. atropurpureum G. marginale Fig. 2. The probable phylogenetic relationships of the perennial species of Utah Geranium. Dec. 1974 NEBEKER: GERANIUM REVISION 301 C. atropurpureum ^S G. carol inianum /\ Fig. 3. Distribution of Geranium in Utah. Harrison 7543 (bry). Univ. St. and 1st. So., Salt Lake Co., WSF 729 (ut). Along Provo River, Utah Co., Sanders sn (ut). Near Salt Lake City, Salt Lake Co., Garrett 1766 (ut). Above Pelican Pond, Cache Co., Thieret 149 (utc). Green Canyon, Cache Co., Shaw 36 (uTc). Logan, Cache Co., Smith 17669 (utc). 2. G. carolinianum L. Sp. PI. 682 (1753) G. bicknellii Britt. var. longipes (Wats.) Fern. Annual; stems 17-40 cm long, erect or branching at the base, short pubescence; leaf blades 2.5-7 cm wide, orbicular to reniform, 5-7 302 GREAT BASIN NATURALIST Vol. 34, No. 4 palmately parted; inflorescence very compact; sepals 6-8 mm long, tipped with awn 1-2 mm long; petals about as long as sepals, pink to wnitish; 10 fertile stamens; style column 12-18 mm long with glan- dular hairs; carpel bodies 3-3.5 mm long; seeds reticulate. Open places and fields throughout North America. Type locality, Carolina. Antelope Island, Howard sn. (ut) Fern Hollow near Ogden, Weber Co., Cardon 313 (utc) "Geranium carolinianum has been confused with G. bicknellii Britt., the latter having been included in the Wasatch region by at least two authors. Inclusion of G. bicknellii in the flora might pos- sibly have been based upon a collection from Logan Canyon in 1910 (C. P. Smith 2164). The stage of the plant's development makes it impossible to determine its true identity." (Shaw, 1952) 3. G. atropurpureum Heller Bull. Torr. Bot. Club 25:195 (1898) G. caespitosum James ex Torr. apud Gray Perennial; stems 10-90 cm long, erect sometimes becoming de- cumbent with age, retrorsely pubescent; basal leaves 2-6 cm wide, reniform to orbiculate, divided nearly to the base into 3-5 lobes, generally larger than the cauline leaves; inflorescence open and spreading; pedicels retrorsely pubescent but not glandular; sepals 9-11 mm long, awned; petals 10-15 mm long, pink to purple, pilose on petals V3-V2 the petals' length, petals recurving; stylodia average 6.55 mm long. Often fovnid growing in association with oak. Type from Santa Fe Creek, New Mexico (Heller 2723). (34 seen) Pine Valley, Wash. Co., Cottam 8905 (ut). Beaver Canyon, Beaver Co., Warnock sn (ut). Devils Canyon Campground, San Juan Co., Cottam 9523 (ut). Long Canyon above Orderville, Kane Co., Cot- tam 4279 (ut). 4 miles north of Glendale, Kane Co., VanCott 1002 (bry). Sheba mine, Millard Co., Cottam and Biddulph 3200 (bry). "It was to this species that Gray assigned the name G. caespito- sum (Gray 1849) believing it to be the species described by .lames. Material in the Gray Herbarium shows G. atropurpureum extending northward into the southern and southwestern counties of Colorado but not into the northeastern region where James is supposed to have collected his "caespitose" Geranium as pointed out by Heller (1898). Lacking collections from the general region or actual specimen col- lected by James, it seems advisable to follow Heller in calling this distinctly southern species G. atropurpureum and to consider G. caespitosum James ex Torr. a nomen dubium." (Moore, 1943) 4. G. marginale Rydb. ex Hanks and Small. N. Am. Fl. xxv 16 (1907) Perennial; stems 10-30 cm long, retrorsely pubescent, slightly exceeding the basal leaves; leaves 2-3.5 cm wide, 5 parted; in- florescence not compact; pedicels retrorsely pubescent, not glandular; sepals 7-9 mm long, puberulent to nearly glabrous; petals purple, pilose 14 the petal length; style column 1.5-2 cm long, stylodia aver- age 4.33 mm long; carpel bodies 4 mm long; seeds 3 mm long, faintlv reticulate. Type ]oca^it^• is the Aquarius Plateau at the head Dec. 1974 nebeker: geranium revision 303 of Poison Creek, Utah, Rydberg and E. C. Carlton 7401. (Hanks and Small, 1907). This is probably the smallest of the perennial species found in Utah. Aquarius Plateau, 35 miles north of Escalante in Wayne Co. Holmgren, Reveal, and LaFrance 2080 (bry). Aquarius Plateau, Garfield Co., VanCott 1196a (bry). 2 miles north of Fishlake, Sevier Co., VanCott 933 (bry). Bryce Canyon, Weight B-31/26 (ut). Wildcat Ranger Station, Garfield Co., Cottam 14130 (ut). Aquarius Plateau, Garfield Co., Cottam 9113 (ut). 5. G. parryi (Engelm.) Heller. Cat. N. Amer. PL ed. 2. 7 (1900) G. pattersonii Rydb., G. fremontii var. parryi Engelm. Perennial; stems 10-45 cm tall, glandular-pubescent throughout; petioles glandular; leaves 2-7 cm wide, deeply 3-5 parted; inflor- escence open and spreading; pedicels glandular-pubescent; sepals 6-10 mm long; petals 12-15 mm long, purple, pilose on petals I/4-V3 their length; style column 1.5-3 cm long, glandular, stylodia average 5.23 mm long; carpels 4-5 mm long; seeds 3-3.5 mm long, reticulate. No specimens of this plant have been seen from the state, but it should be looked for in the Uinta Mountains and in other parts of eastern Utah. Another completely glandular species with which this species may be confused is G. viscosissimum. The distinctive characteristic separating them is the compact inflorescence of G. viscosissimum. 6. G. richardsonii Fisch. & Trautv. Ind. Sem. Hort. Petrop. 4:37 (1837) G. gracilentum Greene, G. albiflorum sensu Hooker, G. pentagynum Engelm., G. loloense St. John. Perennial; stems 30-90 cm tall, glabrous to pubescent; petioles long; leaves 3-15 cm wide, deeply 3-5 parted; inflorescence open; pedicels glandular-pubescent, glands usually purple; sepals 6-12 mm long; petals 1.5-2 cm long, white sometimes bluish, pilose I/3-V2 the petals' length; style column 2-2.5 cm long, glandular, st3dodia aver- age 4.06 mm long; carpel bodies 2.5-4.5 mm long. Found in partial shade or in rather moist ground. (36 seen) 3 miles north of Roosevelt, Duchesne Co., Hardy 113 (bry). Pine Valley Campground, Pine Valley Mts., Washington Co., Higgins 3454 (bry). Cedar Mountain near Navajo Lake, Iron Co., Higgins 4666 (bry). East of Kamas 1 mile above Upper Falls, Summit Co., VanCott 967 (bry). % mile west of Puffer Lake, Beaver Co., VanCott 904 (bry). 10 miles west of Monte Cristo, Cache Co., VanCott 898 (bry). Aspen Grove, Utah Co., Hardy 87 (bry). Vernon, Tooele Co., Frischkmecht 106 (bry). Skyline drive east of Sterling, Sanpete Co., VanCott 943 (bry). Kigalia Ranger Station, San .luan Co., WDS sn (ut). Deep Creek Mts., Juab Co., Lindsay 265 (ut). 7. G. fremontii Torr. ex Gray. PI. Fendl. 26 (1849) G. caespitosum sensu Rydb., G. furcatum sensu Hanks & Small Perennial; stems 20-70 cm long, glabrous to pubescent, not glan- 304 GREAT BASIN NATURALIST Vol. 34, No. 4 dular; petioles long on northern plants shorter in southern speci- mens; leaves 3-10 cm wide, 5-7 parted, cauline leaves much smaller; inflorescence open, spreading; pedicels glandular-pubescent; sepals 7-12 mm long; petals 1-1.5 cm long, purple, pilose 1/4-/^3 their length; style column 2.5-3 cm long, stylodia average 5.23 mm long; carpel bodies 4-5 mm long. Open areas of the foothills and mountains. (49 seen) 4 miles east of Logan, Cache Co., VanCott 886 (bry). Alpine Loop, Wasatch Co., Mullins 17 (bry). South shore of Pine- view, Weber Co., VanCott 896 (bry) . 4 miles west of Garden City, Rich Co., VanCott 885 (bry). Near Cedar Breaks, Iron Co., Higgins 4643 (bry). Pole Canyon, Utah Co., Hartman 127 (bry). Tanners Flat, Little Cottonwood Canyon, Salt Lake Co., Day 30 (bry). 13 air miles south of Wah Wah summit, Beaver Co., Holmgren and Bethers 3824 (bry). Devils Canyon Campground, San Juan Co., Cottam 9523 (ut). Pine Valley, Washington Co., Cottam 8905 (ut). Beaver Canyon, Beaver Co., Warnock sn (ut). 15 miles NW of Orderville, Kane Co., Maguire 18821 (utc). This species is often confused with G. nervosum, which grows north of Utah. The difference between the two is in the inflor- escence: G. nervosum has a compact inflorescence, and G. fremontii has a spreading inflorescence. Bibliography Fernald, M. L. 1935. Geranium carolinianum and allies of northeastern North America. Rhodora 37:295-301. Gray, A. 1849. Geraniaceae. Plantae Fendlerianae Novi-Me.\icanae. Mem. Am. Acad. 4:1-116. Hanks, L. T.. and J. K. Small. 1907. Geraniaceae. N. Amer. Fl. 25(1): 3-24. Harrington, H. D. 1954. Manual of the plants of Colorado. Denver, Sage Books. 666 p. Heller, A. A. 1898. New plants from western North Ainerica. 1. Bull. Torr. Bot. Club 25:193-201. Jones, G. M.. and F. F. Jones. 1943. A revision of the perennial species of Geranium of North America and Canada. Rhodora 45:5-25, 32-53. James, E. P. 1825. Catalogue of plants collected dining a journej' to and from the Rocky Mountains, during the summer of 1820. Trans. Am. Phil. Soc. (new ser.) 2:172-190. Knuth, R. 1912. Geraniaceae. Das Pflanzenreich. Hefte 4, 129(53) :43-221. Moore, H. E. 1943. A revision of the genus Geranium in Mexico and Central America. Contr. Gray Herb. 144:1-108. MuNz, P. A. 1959. A California flora. Univ. Calif. Press. Berkelcv. 1681 p. Osterhout, G. E. 1923. What is Geranium caespitosum James? Bull. Torr. Bot. Club 50:81-84. Shaw, R. J. 1952. A cytotaxonomic study of the genus Geranium in the Wasatch Region of Idaho and Utah. Madrono 11:297-304. VanCott, J. W. 1969. A note on Geranium richardsonii Fisch. 8z Trautv. Great Basin Nat. 29:25-27. UTAH PLANT NOVELTIES IN ASTRAGALUS AND YUCCA Stanley L. Welshi Abstract. — Astragalus iselyi and A. stocksii are described as new species. The former is from the La Sal Mountains of Grand and San Juan counties in eastern Utah, and the latter is from the Henry Mountains in eastern Garfield County, Utah. Yucca toftiac is described as new from materials collected along Glen Canyon in San Juan and Kane counties, Utah. Studies of plant collections taken from previously unexplored or little-known regions of Utah have demonstrated the presence of taxa not included in the known descriptions of plant species of the state. Two species of Astragalus and one of Yucca are included in the new entities. The La Sal Mountains and the Henry Mountains yielded the Astragalus species. The Yucca is from the margin of Glen Canyon. Astragalus iselyi Welsh sp. nov. Fig. 1 A Astragalo sabuloso Jones differt floribus parvioribus et petalis albidioribus carina immaculata et leguminibus parvioribus. Herba perennis radice palari forti et caudice ramificanti; caules 8-25 cm alti recti aut ascendentes; stipulae 3-9 mm longae firmae purpurascentes amplectentes sed non connatae; folia 3.2-8.5 cm lon- ga; foliola (3) 5-11 (13), 7-23 mm longa, 3-9 mm lata elliptica ad rhombica strigosa pilis simplicibus utrinque glabrescens; pedimculi 1.7-10 cm longi; racemi 1.2-3 cm longi, floribus 7 ad multus; brac- teolae nullae; calyx strigulosus tubo cvclindrico 5.5-6.3 mm longi dentibus 1.8-3.1 mm longis; subulatis; flores 17-18 mm longi, petalis ochroleucis, carina apice immaculato; legimiina 25-32 (38) mm longa stricta subcylindrica unilocularia coriacea strigosa. Utah: San Juan Co., La Sal Mts., Brumley Ridge, ca. 1.5 miles north of Pack Creek Ranch, on Morrison formation, in pinyon- juniper community, S. L. Welsh 10970, 5 May 1971 (Holotypus BRY; Isotypi ISC, and many others) ; same locality, J. Pederson 23, 29 March 1967 (bry); S. Daines 39, 5 May 1971 (bry); C. Schoener 75, 11 June 1971 (bry). Grand Co., Paradox formation, gypsiferous clay. Onion Creek, Fisher Valley, S. L. Welsh 11929, 11929a. 30 May 1973 (bry). Astragalus iselyi is a near congener of A. sahulosus Jones. It is similar in habit, leaflet shape and number, and pod features. The flowers are consistently smaller in all parts. Even the average pod size seems smaller than in A. sabulosus. The flowers of A. iselyi are only 17-18 mm long, whereas those of A. sahulosus are 28-31 mm long. Both of these entities comploto flowering dviring earh' spring- time, and it seems probable that the existence of A. iselyi has been 'Department of Botany and Range Science, Brighani VounK University. Prnvo, Utah 84602. 305 306 GREAT BASIN NATURALIST Vol. 34, No. 4 Fig. 1. Astragalus iselyi. A, Habit sketch; B, Detail of pods. Dec. 1974 welsh: utah plants 307 obscured by the fact that most previous specimens were collected in fruit only. It seems certain that the phase of the sabulosus-iselyi complex with massive flowers is that known as A. sabulosus. The measure- ments set forth in the original description (Jones, 1891) and in the Revision of North American Species of Astragalus (Jones, 1923:156- 157) are those of the phase including the type of A. sabulosus. Ma- terials collected on the La Sal Mountains by Rydberg and Garrett (9088, 19 July 1911 ut) and by Maguire et al. (16536, 14 July 1933 UTc) are almost assuredly A. iselyi. From the present informa- tion, it seems that A. sabulosus is a plant of bajadas and drainages over Mancos Shale in the Thompson-Cisco region of Grand County. A. iselyi is also a selenophyto, but grows on the Paradox and Morri- son formations on the foothills of the La Sal Mountains. This species is named in honor of Duane Isely, legume specialist, teacher, colleague, and philosopher. Astragalus stocksii Welsh sp. nov. Fig. 2 Species habitu cum Astragalo musiniensi Jones differt foliolis plus nimierosis floribus parvioribus ochroleucis et leguminibus strigosis. Herba perennis acaulis, caudice ligneo ramificanti, ferenti petiolis marcescentibus; stipulae lanceolatae 3-5 mm longae strigosae librae; folia 2.7-10.6 cm longa; foliola 7-13 mm longa obovata oblong- lanceolata vel elliptica, plumbea aut argentea, supra viridia saepe, strigosa utrinque. truncata ad mucronata aut acuta ad apicem; pe- dunculus 1.1-4 cm longus; racemus (2) 3-8 floribus. compactus; bracteae lanci-subulatae, strigulosae; flores ascendentes. 15-16 mm longi; calyx strigulosus, tubo cylindrico, 9-9.2 mm longo. dentibus lanci-subulatis 2-2.8 mm longis; corolla ochroleuca, venis purpures- centibus, carina-apice maculata; alis apicibus purpurescentibus; le- gumina unilocularia, compressa lateralle, lanci-ovoidea, rostro elongato tenui contracto, stricta aut arcuata, strigosa. Utah: Garfield Co., Henry Mountains, Penellen Pass, S. L. Welsh 11740, 30 May 1972 (TTolotypus bry; Isotvpi us. lsc). Near same locality, S. L. Welsh 9817, 16 Mav 1970 (bry, distributed as A. newberryi Gray); S. L. Welsh 11739, 11751. 30 May 1972 (bry. Para types). Astragalus stocksii is compared to the distinctive A. wusinirnsis Jones in the above description. It is indeed similar to that entity both in having a persistent thatch of petioles and in shape and texture of the pods. Habitually it is apparently nearer to A. iinrberryi Gray and to A. eurckensis .lones. From the former, A. stocksii differs in the smaller, paler flowers and merely strigose pods, and from the latter it differs in having smaller flowers and merely strigose pods. The species is named in honor of the late Dayna L. vStocks, botanist, teacher, and extraordinary human being. 308 GREAT BASIN NATURALIST Vol. 34, No. 4 Fig. 2. Astragalus siocksii. A, Habit sketch; B, Detail of pod. Yucca toftiae Welsh sp. nov. Fig. 3 Differt haec species a Yucca angustissima in uterque amplitudine et habitu. Planta acaulescens vel brevicaulescens, solitaria vel caespitosa; caules 0-7 (10) dm alti; folia 2-7.5 dm longa, 0.4-1.7 cm lata linearia Dec. 1974 WELSH: UTAH PLANTS 309 Fig. 3. Yucca toftiae. A. Habit sketch; B, Detail of leaf; C, Detail of flower; D, Detail of capsule and seeds. 310 GREAT BASIN NATURALIST Vol. 34, No. 4 plano-convexa vel concavo-convexa ad plaiio-carinata effusa rigide, viridia vel flavo-viridia, margo filifer; inflorescentiae 1.2-3.4 m altae, scapi folia 2-vel 6-plo longior; paniculae lineares vel ovoidae, 2.5-20 dm longae, ramulis 0-12 (23); flores numerosi globosi vel campanu- lati albi, cremei, vel viriduli plerumque exti rubro-purpurei suffusi, segmenta elliptica vel lanc-ovata, acuta; sepala 2.5-4.6 cm longa; petala 31-52 mm longa; filamenti staminale 7-24 mm longa; pistila 15-32 mm longa; styli (3) 7-11 mm longi; capsulae erectae, 3.5-5.5 (6) cm longae, cylindricae, plurumque constrictae, ligneae et per- sistentes. Utah: San Juan Co., Three Gardens, ca. 1 mile north of conflu- ence of San Juan and Colorado rivers. Lake Powell, S. L. Welsh 11935a, 4 June 1973 (Holotypus bry; Isotypi us, ny). Ribbon Canyon garden, east side of Lake Powell at ca. mile 66, N. D. At- wood 4112, 10 June 1972 (bry). Kane Co., tributary of Drv Rock Creek, Lake Powell, S. L. Welsh and G. Moore 11779, 4 June 1972 (bry). This entity grows on sandy alluvium and on sandstone outcrops along the shores of Lake Powell in Glen Canyon of eastern Kane and western San Juan counties, Utah. The affinities of Y. toftiae are with Y. angustissima Englem ex Trel, from which it differs in both size and habit. Habitually, Y. toftiae is similar to Y. utahensis Mckelvy and Y . verdiensis Mckelvy. From the former it differs in the inflorescence which branches only near the base (or not at all), and from the latter in the short pistils and longer woody capsules. This striking taxon is named in honor of Catherine Ann Toft, botanist. References Barneby. R. C. 1964. Atlas of North American Astragalus. Mem. N. Y. Bot. Gard. 13:1-1188. Tones, M. E. 1891. New species and notes of Utah Plants. Zoe 2:236-252. — ■ . 1923. Revision of North American species of Astragalus. Salt Lake City, Utah: by the author. 288 pp. Mckelvy, S. D. 1938. Yuccas of the soutlawesteni United States. Part 1. Arnold Arboretum. Harvard Univ. 150 pp. . 1947. Yuccas of the southwestern United States. Part 2. Arnold Arboretum, Harvard Univ. 197 pp. A NEW COMBINATION IN PENSTEMON (SCROPHULARIACEAE) James L. ReveaP One of the more distinctive beard-tongues found in the sandy portions of northern Arizona, southern and western Utah, and south- eastern Nevada is Penstemon angustifolius ssp. venosus. The pinkish- lavender to pinkish flowers of ssp. venosus mark it as one of the more readily recognized of the penstemons found in this part of the western United States. The typical subspecies is found far to the east on the high plains of the central United States. The ssp. angusti- folius generally has narrower, more sharply acute foliage, typical blue flowers that are shorter, and mature capsules that are shorter and narrower than those found in ssp. venosus. The bracts in the inflorescences of ssp. angustifolius are not venose on both sides as in ssp. venosus. and the former has the unfortunate feature of blacken- ing upon drying. As a result of its green, glaucous foliage, bright pinkish to lavender flowers, and general rareness, the ssp. venosus is often collected preferentially by even the most seasoned collector. Many collectors have noted their dissatisfaction with Keek's place- ment of this plant with Penstemon angustifolius. but as yet no one has proposed a specific name for the taxon. So that a specific name might be available to Janice C. Beatley for her floristic work on the Nevada Test Site, the following combination is proposed: Penstemon venosus (Keck) Reveal, stat. et comb, nov., based on P. angustifolius Nutt. ex Pursh ssp. venosus Keck in Kearne}- and Peebles, J. Wash. Acad. Sci. 29:490. 1939. As a distinct species, Penstemon venosus may be distinguished by its flower color, leaf size and shape, the nature of the floral bracts, and its disjunct distribution. 'Department of Botany, University of Maryland. College Park 20742, and National Museum of Natural History, Smithsonian Institution, Washington. D.C. 20560. Research supported by National Science Foundation Grant GB-22645. 311 DOMINANCE RELATIONSHIPS OF THE DARK KANGAROO MOUSE (MICRODIPODOPS MEGACEPHALUS) AND THE LITTLE POCKET MOUSE {PEROGNATHUS LONGIMEMBRIS) IN CAPTIVITY Andrew R. Blaustein' and Arthur C. Risser, Jr.^ Abstract. — Interspecific interactions between the little pocket mouse (Perog- nathus longimembris) and the dark kangaroo mouse {Microdipodops megaceph- alus) were tested in the laboratory. P. longimembris was statistically dominant over M. megacephalus. The dominant-subordinate relationships shown by our laboratory results indicate that interspecific aggression may be one mechanism involved in keeping these sympatric species ecologically separated. In certain areas of northern Nevada, the dark kangaroo mouse (Microdipodops megacephalus) and the little pocket mouse (Perog- nathus longimembris) are sympatric (Hall and Kelson, 1959). These species are similar in size and, being primarily granivorous (Hall, 1946), probably have similar feeding habits. As compared with what is known about other members of their family (Heteromyidae), little is known about the ecology of these species. This study investi- gates the possibility that interspecific agonistic behavior may be a mechanism by which ecological isolation occurs between these two species in the field. To assess this possibility we observed inter- specific interactions between pairs of captive M. megacephalus and P. longimembris. The possible role of interspecific agonistic behavior in the habitat segregation of small mammals has been reviewed by Grant (1972). Three Microdipodops (two females and one male) and four Perognathus (two females and two males) were trapped in Warm Springs Valley, Washoe Co., Nevada. The animals were caged individually in steel cages measuring 34 x 24 x 24 cm. The front of each cage was covered with 1 x 1.25-inch hardware mesh. Sand one centimeter deep was placed in each cage. The cages were cleaned periodically. A mixture of sunflower seeds, rolled oats, and millet was given to the animals daily. Lettuce was provided once a week. No water was provided. The housing cages were placed under a 12-hour light — 12-hour dark controlled photoperiod. The light and dark periods were reversed, allowing these normally nocturnal ani- mals to be observed during convenient daytime sessions. The light phase was illuminated by two incandescent 60-watt white light bulbs, and the dark phase was slightly illuminated by two incandescent 25-watt red light bulbs. Encounters took place in a cage measuring 34.5 x 45.7 x 122 cm. Three sides of the cage were aluminum, and the front was plexi- glas. The top was covered with .25-inch hardware cloth, and an aluminum partition divided the cage into two equal sections, each 'Andrew R. Blaustein and Arthur C. Risser, Jr., Department of Biology, University of Nevada, Reno 89507. Present address of Blaustein: Department of Biological Sciences, University of California. Santa Barbara 93106. 312 Dec. 1974 blaustein, risser: rodent behavior 313 containing a nesting area and a food dish. Two cm of sand covered the floor of the cage and was replaced after each interaction. To begin an interaction, one rodent was placed on each side of the central partition. The cage was subjected to the same reversed photoperiod as the housing cages. The animals were left alone for at least 24 hours as an adjustment period. Then the partition was removed, and the behavior and interrelationships of the rodents were recorded for 15 minutes. Interactions with the same two individuals were not repeated. At least two days were allowed to elapse before an individual was used again. All trials were conducted between 1000 hr and 1500 hr. Any overt interaction between two animals was called an en- counter. Agonistic encounters usually ended when one animal took the dominant role by attacking and chasing its opponent and the other became subordinate by escaping or trying to defend its nest area from attack. The behavioral patterns and postures observed between the mice were classified into five major categories. These have been modified from Eisenberg (1963). Upright posture. — The animal assumes a posture with the long axis of the body at an angle greater than 45 degrees to the substrate. Rushing. — The animal usually has an elongated posture. The ears are erect and the eyes are wide open. When the opponent is reached, the animal may strike at it with its forepaws. This is the major aggressive movement. Chase. — The animal runs after the opponent and tries to bite its rump. Nest defense. — This usually involves an upright posture in de- fending a nest area from an opponent. It may involve short elongate rushes whereby the defender rushes in short spurts, not straying far from its nest area. Escape leap. — Wild erratic jumps are used to escape a rush or subsequent chase. Fighting usually began within the first two minutes after the partition was lifted and was initiated by P. longimembris in 11 of 12 interactions. The predominant patterns during interactions were rushing by Perognathus and escape leaping by Microdipodops. Perognathus was usually deliberate in rushing Microdipodops and when moving away it assumed a slow quadrupedal gait. Micro- dipodops defended its nest area vigorously during interactions but was usually driven away by Perognathus. Rushing by Perognathus also induced Microdipodops to defend its nest area via an upright stance or via a partial rush; that is, Microdipodops started to rush Perognathus but stopped short, not advancing far from its nest area. If Microdipodops escape leaped, Perognathus usually moved away or, rarely, chased Microdipodops. If Microdipodops defended its nest area, Perognathus would usually move away only to return several 314 GREAT BASIN NATURALIST Vol. 34, No. 4 seconds later, rush Microdipodops and take over its nest area. Micro- dipodops only entered the Perognathus half of the cage after they were rushed or chased. Although Eisenberg (1963) noted chasing and locking fight (two animals meet and lock together by gripping with all four limbs) to be quite common between pocket mice intra- specifically, chasing was not frequent and locking fight did not occur in the present study. During interactions both species squealed in high-pitched tones. However, Microdipodops vocalized more often and this usually occurred when defending a nest area. During 12 interactions of all combinations of sex pairings be- tween Microdipodops and Perognathus, P. longimembris was domi- nant 11 times (Table 1). In one interaction involving a female of each species, neither individual was clearly dominant over the other. In all other cases, Perognathus was clearly dominant regardless of sex. Perognathus was equally dominant in both halves of the cage. Since each individual was used in more than one trial, the Mann- Whitney U test (Wilcoxon's two-sample test) was utilized to deter- mine whether P. longimembris is significantly dominant over M. megacephalus. The Mann- Whitney U test in this case can be com- puted as the number of times a P. longimembris was dominant over M. megacephalus out of the 12 possible combinations (see Sokal and Rohlf, 1969). In this case, the "U" of the Mann-Whitney U test is either 0 or 1 ; the probability of this occurring if there is no difference between the species is either 1 or 2 ( 1 < P < 2). Thus, P. longi- 35 35 35 35" membris is significantly dominant over M. megacephalus. How does this compare with Grant's (1972) generalization that larger species are usually dominant over smaller species? We decided to compare differences in weights between the two species by utilizing Student's t test for the difference between mean weights and found that P. longimembris^, the dominant species, was significantly lighter (P < 0.05; N = 4, X = 7.38g, SE=t:0.69) than Microdipodops (N = 3, X=10.5g, SE±0.88). The dominant-subordinate relationships between these two species may be a mechanism by which these species are ecologically separat- ed in the field. O'Farrell's (1973) population study of the desert rodents in the same area where we collected our experimental ani- Table 1. Dominance relationships in Microdipodops megacephalus (Mm) — Perognathus longimembris (PI) interactions. Percentages are in parentheses, dom = dominant Number of Combination interactions Mm-dom Pl-dom None-dom Male Mm-Female PI 2 0 2 0 Male Mm-Male PI 2 0 2 0 Female Mm-Male PI 4 0 4 0 Female Mm-Female PI .... __4 0 3 1 Totals 12 0 11 (91.7) 1 (8.3) Dec. 1974 blaustein, risser: rodent behavior 315 nials lends credence to this idea. His data show tliat on a 2.7 hectare grid M. megacephalus emerged in early March and steadily increased until the middle of April. Towards the end of April, a steady decline of Microdipodops corresponded with the emergence and increase in numbers of P. longimembris. During the summer very few Microdipodops were captured and Perognathus covered the entire grid. In September Perognathus activity declined and Micro- dipodops activity again increased. Throughout the spring and fall, the centers of activity of the two species did not overlap. Thus, they were spatially isolated. O'Farrell (1973) believes that P. longimembris is the more general species and occupies a broader niche than Microdipodops because it was found on all habitat types on the grid while Microdipodops was generally restricted to areas of fine, loose sand. Furthermore, O'Farrell (1973) believes that M. megacephalus probably occupies an included niche within the funda- mental niche of P. longimembris (see Miller, 1967, for terminology"). An interesting situation may exist in Warm Springs Valley because the more general species (P. longimembris) appears to be competi- tively superior to tne specialized species {M. megacephalus). This is not common, according to Miller (1967). He stated that if there are two s}TTipatric species one of which occupies a smaller included niche within the broader niche of the other, then for coexistence to continue, the species occupying the smaller niche must be the su- perior competitor. How, then, has Microdipodops avoided extinction? The answer to this question lies in the temporal aspects of the niches of the two species. As stated above, O'Farrell (1973) found that P. longimem- bris is active primarily during the summer while M. megacephalus is active primarily during the spring and the fall. Thus, temporally, P. longimembris, the superior competitor, has the narrower niche; and if a time axis were included in Microdipodops' s fundamental niche, this species niche would not be totally within the fundamental niche of P. longimembris. Presumably, M. megacephalus would not show decreased summer activity if P. longimembris were absent. When the temporal components of the niche are taken into account, it is seen that this system may conform to Miller's (1967) generali- zation. The dominant-subordinate relationships shown by our labora- tor)^ results and the fact that M. megacephalus apparently becomes rare as Perognathus increases in summer may indicate that inter- specific aggression is one mechanism by which ecological isolation is maintained between P. longimembris and M. megacephalus in the field. We would like to thank Drs. Fred A Ryser, .Jr., and Beatrice Gardner (University of Nevada, Reno) for critically reviewing early drafts of the manuscript. Dr. Michael J. O'Farrell (Savannah River Ecology Laboratory) critically reviewed an earlier draft of the manuscript and allowed us access to some of his unpublished data. For this we wish to express our appreciation. Dr. Allan Oaten (Uni- versity of California, Santa Barbara) helped with the statistical analysis of the data. Special thanks go to Dr. Stephen I. Rothstein 316 GREAT BASIN NATURALIST Vol. 34, No. 4 (University of California, Santa Barbara) for critically reviewing the manuscript and offering many helpful suggestions. Literature Cited EiSENBERG, J. F. 1963. The behavior of heteromyid rodents. Univ. Calif. Publ. Zool. 69:1-114. Grant, P. R. 1972. Interspecific competition among rodents. Ann. Rev. Ecol. Syst. 3:79-106. Hall, E. R. 1946. Mammals of Nevada. Univ. Calif. Press. Berkeley, 710 pp. Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. Ronald Press, New York, l:xxx + 1-546 + 79 pp. Miller, R. S. 1967. Patterns and process in competition. Adv. Ecol. Res. 4:1-74. O'Farrell, M. J. 1973. Seasonal activity patterns, population dynamics, and spatial relationships of rodents in a sagebrush community. Unpublished Ph.D. dissertation, Univ. Nevada, Reno. 158 pp. SoKAL, R. R., AND F. J. RoHLF. 1969. Biometry: the principles and practice of statistics in biological research. W. H. Freeman and Co., San Francisco, xxi + 776 pp. LIST OF INSECT TYPE SPECIMENS IN THE ENTOMOLOGICAL COLLECTION OF BRIGHAM YOUNG UNIVERSITY, PROVO, UTAH, NO. V. Vasco M. Tanner^ Abstract. — Type material added to the Brigham Young University Life Sciences Museum insect collection includes 6 holotypes. 5 allotypes, 106 para- types. 78 cotypes, 48 homotypes, 1 metatype. and 1 androtype. Since the previous lists (I-IV)- were published, the following type specimens have been added to the University Type Collections. Thousands of specimens have also been added to the research col- lection. The pinned collection now consists of more than 900,000 specimens. These are arranged in cabinets, catalogued as to orders, families, genera and species. Fortunately, rather intensive collecting was conducted in many areas of this region prior to the changes that have been made in the lands, watersheds, and drainage of the Great Basin. The biota of one hundred jears ago is now greatly changed. Introduced plants and animals, along with killing and poisoning activities, have reduced and may have exterminated some species. In 1974, Mrs. Lucile Maughan Johnson, wife of the late D. El- mer Johnson, contributed their bee-fly (Bombyliidae) collection to the Life Sciences Museum at Brigham Young University. This col- lection consisted of 6,400 pinned specimens. Included in it are many type specimens. Order Hemiptera Family Belostomatidae Abedus herberti utahensis Menke Univ. Calif. Publ. Ent. 16(8) : 423-424. 1960. 3 paratj-pes: St. George. Utah; V-1917. V. M. Tanner. Order Homoptera Family Hydrometridae Hrdrometra becki Drake Bull. So. Calif. Acad. Sci. 50(2): 103. 1951. 3 paratopes: Solomon Islands; Guadalcanal: VI-30-1944. D E. Beck. Order Coleoptera Family Cantharidae Malthodes ihedae Fender Northwest Sci. 42(3) : 108-1 10. Holotype (male):Hat Creek. Lassen National Poorest. California; VL24-1961, S. L. Wood, J. B. Karren. and D. E. Bright. Family FLlateridae Megopenthes apacheorum Becker Canadian Ent. 103(2) : 157-160. 1971. ^Depailnient of Zoology. Briglu-ini Yoimt; Univcisit\-. I'mvii. Uliili .SK)OJ. =I.ist I. IVoc. L't. .\cad. Sri.. .\ils fin. I r.Ptlcrs IJISl I'M. I<1i5. I.isI II. I'm.. Ll. .\<,ul. S- g to the Biological natural history of western North Ameriv iitended for publica- tion in the Great Basin Naturalist should be du-ected to Brigham Young University, Stephen L. Wood, Editor, Gre?>t R.t-Iti Naturalist, Provo, Utah 84602. Manuscripts: Two copies are required, typewritten, double- spaced throughout on one side of the paper, with margins of at least one inch on all sides. In general, the style should confonn to recom- mendations published in tlie most recent edition of the Council of Biology Editors Style Manual, published by the American Institute of Biological Sciences. An abstract, about 3 percent as long as the text and written in accordance with Biological Abstracts guidehnes, should precede the introductory paragraph of each article. Illustrations and Tables: All illustrations should be made with a view of having them appear within the limits of the printed page. The illustrations that form a part of an article should ac- company the manuscript. All illustrations and tables to appear in this journal are to be made under the supervision of the editor, and the cost of the cuts and labor required to set tables is to be borne by the Contributor. Reprints: No reprints are L -j of charge. A price list for reprints and an order form are sent with the proof. Reprints Schedule of the Great Basin Naturalist Each Additional 2pp. 4pp. 6pp. 8pp. 10pp. 12pp. 2pp. 50 copies $12.00 114.00 $16.00 $18.00 $20.00 $22.00 $3.00 100 copies 15.00 17.00 19.00 21.00 23.00 25.00 200 copies 18.00 20.00 22.00 24.00 26.00 28.00 300 copies 20.00 22.00 24.00 26.00 28.00 30.00 TABLE OF CONTENTS Field behavior and seasonal activity of the rodent bot fly. Cuterebra tenebrosa, in central Washington (Diptera: Cuterebridae). Craig R. Baird 247 Notes on the spotted bat (Euderma maculatum) from southwest Utah. Richai'd M. Poche and Geoffrey L, BaiUe 254 The significance of scale characters in evaluation of the lizard genera Gerrhonotus, Elgaria, and Barisia. Jc-mies W. Waddick and Hobart M. Smith 267 Revegetation of gopher mounds on aspen range in Utah. W. T. McDonough 267 Oljserv^ation on voice in the western coUarde lizard, Crotaphytus collaris bicinctores. Nathan M. Smith 276 iWw synonymy and records of American bark beetles (Coleoptera: Scolytidae). Stephen L. Wood 277 Description of new species of Miloderes Casey, with com- ments on otiier species in the genus (Coleoptera: Curcurlionidae) . Vasco M. Tamier 291 czrtcme Bookbinding Co., Inc. 300 Sur.uncr Street Boston, Mass. 02210 3 2044 072 231 061 Date Due