A hed bard Bitieiee: seca ny ' tht ayaa aca Hagia ee et Cee iw ats hr m di + wah i ri : s ‘i Bie ae wy Bare 2a x 4; i sy! wet ae a) ape Cieattes oo ee Se ae HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology ie =e os4 me rs) ( ; a ay cmc os ss) : : Ms a i ¥ oe AE Gilbert Dennison Harris (1864 - 1952) Founder of the Bulletins of American Paleontology ( A} SBN 0-87710-4 : SEIS HISS LUME 100, NUMBER 337 Late Triassic cyrtinoid spiriferinacean brachiopods from western North America and their biostratigraphic and biogeographic implications by Peter R. Hoover Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. OCTOBER 16, 1991 | CAS hor PALEONTOLOGICAL RESEARCH INSTITUTIC YN Officers PRESIDENT fhe eo oe eo ee ee Harry A. LEFFINGWELL MIGES PRESIDENTS): can evils it sie Saycicisie nas aketel ore Och aes J. THOMAS DuTRO, JR. SEGRETAR Ys svete ocr eee fi lsya ee Pea eer HENRY W. THEISEN SUREASWRER Oly at As ae en a Ge oe eine JAMES C. 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The Paleontological Research Institution is a non-profit, non-private corporation, and contributions may be U.S. income tax deductible. For more information on P.R.I. programs, memberships, or subscriptions to P.R.I. publications, call or write: Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York 14850 U.S.A. 607-273-6623 OCTOBER 16, 1991 VOLUME 100, NUMBER 337 Late Triassic cyrtinoid spiriferinacean brachiopods from western North America and their biostratigraphic and biogeographic implications by Peter R. Hoover Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. LIST OF ILLUSTRATIONS Text-figure Page 1. Sketch map showing location of North American collecting localities cited in this report .... 2.0.0.0... 0c c cece eee eee eee eee 64, 65 2. Views) of theicooperculum of Spondylospira lewesensis (Lees, 1934) sec. - joc cee coe eee cee aee sees sees snes eee 76 3. Dorsoposterolateral view of ventral valve of Dagyspirifer fascicostata, n. gen. and n. sp., showing one of the two small, paired, ovate pedicle foramena that pierce the apices of both the ridged portion of the interarea and the dental plates ....................... idl 4. Anterodorsolateral (interior) view of ventral valve of Dagyspirifer fascicostata, n. gen. and n. sp., showing the horizontal bar that connects: thedental! plates ‘and lateral'shell wall’ ~...0. se.ace «oc eee one oe ee eo aise cess eke Gert ee ee 78 5. Posterodorsolateral views of portions of a dorsal valve of Phenacozugmayerella mimuncinata, n. gen. and n. sp., showing surface IMICHOOTMAMENUS 15 scree zsepsoc ck eos ech neucien Biers eeocah acess eats ea ves Dadol Oe ee op Te ny Re ee 91 LIST OF TABLES Table Page le Wi pper TnassiciSeries; Stages; substages, and Zones, -52-.4- 50 een sess sere eee sles eee eee ee eee 66 2. Measurements of type specimens of Dagyspirifer fascicostata, n. gen. and N. SP... 2... ee ee ee ce ee cece eee eee eee ee 78 3. Measurements of type specimens of Pseudospondylospira perplexa, n. gen. and N. SP. 2... ee ev vv cece eee eevee eee ee 80 4. Measurements of specimens of Spondylospira lewesensis (Lees, 1934) .... 2.220. ec cece cece ec cc cece ccencccusccvcseresueeeue 82 5. Measurements of the holotype of Spondylospira parmata Hoover, 1983 ............00cccccceccceccueceucccucceuseeceeusceus 84 6: Measurements ofispecimens of Spondylospira tricostaynsisp) ..- 4244s cee so aden eee 85 7. Measurements ofitype’specimens' of Vitimetula parva, n. gen. and\n: sp; ..........+2-os06 sec eee eee eee 87 8. Measurements of type specimens of Zugmayerella americana, N. SP. 2... 2... cece cc nce e ence e ce ceceeceeeuceuseeeeees 88 Ov Measurements of:specimens of. Zugmayerella? spA\ c....-- saaec seen aeac ee neseesee seis te ene nen eee 89 10. Measurements of type specimens of Phenacozugmayerella mimuncinata, n. gen. and N. SP. ..... 2. eee ee ev ee ee eee eee eee eee 90 LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS FROM WESTERN NORTH AMERICA AND THEIR BIOSTRATIGRAPHIC AND BIOGEOGRAPHIC IMPLICATIONS By PETER R. HOOVER Paleontological Research Institution 1259 Trumansburg Road Ithaca, NY 14850-1398, U.S.A. ABSTRACT Cyrtinoid spiriferinacean brachiopods in samples from over fifty Late Triassic marine fossil localities in western North America include examples of three species of Spondylospira Cooper, 1942, one of which (S. tricosta) is new; one new species of Zugmayerella Dagys, 1963; and four new monotypic genera, represented by Dagyspirifer fascicostata, Phenacozugmayerella mimunicinata, Pseudospondylospira perplexa, and Vitimetula parva, all new species. In North America, all of these genera are limited to the Karnian and/or Norian. One species, Spondylospira tricosta, n. sp. appears to be a good indicator of the early Norian. The family Laballidae of earlier workers is divided into the family Laballidae and the new family Spondylospiridae, based on absence or presence, respectively, of functional pedicle foramena; the latter family is divided into two new subfamilies, Spondylospirinae and Dagyspiriferinae, based on the form of those foramena. Two features of cyrtinoid spiriferinacean species biogeography have tectonostratigraphic implications. First, there are no “Tethyan” species among them, and second, the nine species of cyrtinoid spiriferinaceans reported here are predominantly confined to the Americas, including both cratonal and terrane belt localities. Biogeographic evidence from the cyrtinoids, therefore, does not support close links between terranes of the present-day eastern and western Pacific. Cyrtinoid spiriferinaceans provide abundant evidence of biogeographic “‘stitching” of various terranes. For example, Pseu- dospondylospira perplexa, Spondylospira lewesensis, and Spondylospira tricosta occur in localities that are craton-bound as well as in parautochthonous and allochthonous localities. ACKNOWLEDGEMENTS I owe a great debt of gratitude to Dr. Cathryn R. Newton (Syracuse University, Syracuse, NY) for her help with the tectonostratigraphic summaries of the various localities, and for discussions of the paleogeo- graphic implications of this fauna. I am indebted to the National Research Council for awarding me a post- doctoral fellowship at the United States Geological Survey’s Paleontology and Stratigraphy Branch at the United States National Museum of Natural History in Washington, DC, during which this study was initi- ated. I am also indebted to the Trustees of the Pale- ontological Research Institution, of Ithaca, New York for encouraging me to take the necessary time to com- plete the study. Drs. J. Thomas Dutro, Jr. (U. S. Geological Survey, Washington, DC, and my advisor in the post-doctoral study which led to this paper), Norman J. Silberling, (U.S. Geological Survey, Denver, CO), Algirdas S. Dagys (Institute of Ecology, Vilnius, Lithuania), and Derek Ager (Swansea, Wales) reviewed the manuscript and made valuable comments, many of which have been incorporated into the manuscript. Iam grateful to Fred Collier (U. S. National Museum of Natural History, Washington, DC, U. S.A.), Dutro, Silberling, George Stanley (University of Montana, Missoula, MT, U.S. A.), and E. T. Tozer (Geological Survey of Canada, Vancouver, British Columbia, CANADA) for the opportunity to examine collections under their care, and for loans of specimens, encour- agement, constructive criticism, and gracious hospi- tality during the course of this study. The illustrations in this study came from a variety of sources. The photographs on all the plates were produced in 1975 by R. H. McKinney and H. E. Moch- izuki of the U. S. Geological Survey’s Paleontology and Stratigraphy Branch in Washington, DC. I drafted Text- figure 1; the photographs in Text-figure 2A and B were shot by McKinney and Mochizuki, and Jorge Valdes, of S. T. Pees and Associates, Meadville, PA, respec- tively; Text-figures 3 and 4 come from the talented pen of Jude Louviere of Toledo, OH; and the photographs that comprise Text-figure 5 were expertly shot and printed by David Tuttle, of the Department of Geo- logical Sciences at the State University of New York at Binghamton. INTRODUCTION Late Triassic cyrtinoid spiriferinacean brachiopods from western North America historically have been poorly known. This is partially due to their rarity (com- pared with other Late Triassic invertebrate fossils) and partially due to the comparatively greater biostrati- graphic utility of other forms (e.g., ammonites and flat clams, and, more recently, conodonts; see discussion 64 BULLETIN 337 Li AN aN ‘. x Nee ae ny a cae x . eee .. jo ee x va Q : < . U2 . Koq-31 13,17 oe ¢ Se aoe ae cee aU o> oo 6 53 20° fo & Secs ° ; ; TOS = uf ; ost a 0? Text-figure 1.—Sketch map showing location of North American collecting localities cited in this report (arrows point North). below). Although there were sporadic reports under the guise of Cyrtina Davidson, 1858 and Spiriferina d’Orbigny, 1847 during the late nineteenth and early twentieth centuries, not until Cooper’s work of 1942 and 1944, which defined Spondylospira, were these forms studied in appropriate detail. In 1944, in Shimer and Shrock’s Index Fossils of North America, Cooper illustrated a specimen he assigned to Psioidea sp. (p. 359, pl. 140, figs. 52-55). This specimen was differ- ted from the species of Spondylospira illustrated by having a smooth fold and sulcus. The il- shows a very smooth surface, and exami- the specimen (USNM 106250) reveals it to he internal mold of a cyrtinoid spiriferinacean of a size and form similar to Spondylospira lewesensis (Lees, 1934). Because it lacks any remaining vestige of the shell, it cannot confidently be assigned to any known cyrtinoid spiriferinacean genus or species. These stud- ies were followed in short order by the Vokes and Haas (1944) compendium of information on American spe- cies of Spondylospira, which incorporated available information from the Triassic of Peru (KOrner, 1937; Tilmann, 1917). Since then most North American Late Triassic cyrtinoid spiriferinacean brachiopods have been referred to Spondylospira, including some forms that were not even spiriferinaceans (e.g., Spondylospira alia Hall and Whitfield, 1877; see discussion under Spondylospira lewesensis). In recent years, some cyr- tinoid forms that lack a costate fold and sulcus have been assigned to Zugmayerella Dagys, 1963 [see e.g., Zugmayerella uncinata (Schafhautl, 1851), cited in Stanley, 1979, p. 14, pl. 8, figs. 10-13], but none has been formally described. In 1976, as part of a post-doctoral project, I began a study of Late Triassic articulate brachiopods, based primarily on materials deposited in the stratigraphic LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER s N 14 aN a . 4 ‘ 65 ~ 15 > Oi 47 fy a12 , é 28 ’ ; oi a hee ie Sy ~~ 46, ve = sB:49: at ’ 32,41,43 aS ee 18,39, stig a Text-figure 1.—[continued]. collections of the U.S. Geological Survey in Wash- ington, DC, and Menlo Park, CA. This eventually in- volved over 50 collecting localities, spread over an area from central Nevada north to southwestern Alaska (Text-fig. 1). It became immediately apparent that time and equipment were inadequate to do justice to the terebratulids and rhynchonellids, both of which re- quire elaborate preparation of many specimens to ad- equately understand the variation in expression of tax- onomically significant characters. The spiriferoids were commonly better preserved, and their usable taxonom- ic characters more readily accessible. Of these, the cyr- tinoid spiriferinaceans were a convenient group to work on, as they contain a variety of structures that are of great interest in terms of functional interpretations. Indeed, these puzzling functional interpretations are one of several important factors that have delayed pub- lication of this monograph (see Hoover, 1983; 1990). Fourteen years later, in March of 1990, having be- come increasingly aware of the many fine Late Triassic marine fossil localities that had been collected from by Canadian workers, I visited the Geological Survey of Canada in Ottawa, and surveyed its Triassic bio- stratigraphic collections. At the Canadian Survey, as in the U.S., fossils are most commonly collected by workers engaged in mapping, and chiefly interested in those forms that have biostratigraphic significance. Be- cause most biostratigraphically significant Triassic in- vertebrate macrofossils are not brachiopods, but rather ammonites or flat clams, there were few brachiopods in the collections. Indeed, the only Canadian localities represented in the GSC biostratigraphic collections that contained cyrtinoid spiriferinaceans were: (1) Nicola, British Columbia [no further information available], which yielded a single ventral valve questionably re- ferable to Pseudospondylospira, n. gen.; (2) the type 66 BULLETIN 337 Table 1.—Upper Triassic Series, Stages, Substages, and Zones (after Tozer, in House and Senior, 1980). Upper Norian (Sevatian)(UNo) Middle Norian Nomani(Ne) (Alaunian)(MNo) Lower Norian (Lacian)(LNo) Upper Karnian Karina) (Tuvalian)(UC) Lower Karnian (Julian)(LC) area for Spondylospira lewesensis (Lees, 1934) [Lake Laberge region, Yukon Territory]; (3) another area in similar strata about 10 km to the east that also yielded S. lewesensis [Lime Peak, Yukon Territory (Reid, 1985)]; (4) the Karnian of Cameron Island in the Ca- nadian Arctic [GSC loc. 25872]; (5) Trout Lake, Yukon Territory [GSC loc. 86206], which yielded isolated specimens that may be referable to Zugmayerella Da- gys, 1963; and (6) the Halfway River area of British Columbia [GSC loc. 47016], which yielded isolated specimens questionably referable to the genus Spon- dylospira Cooper, 1942. This should in no way be con- strued as a gauge of the rarity of cyrtinoid spiriferi- naceans in Canada, any more than U.S. collections are a gauge of their comparative rarity in the U.S. Rather, they are rarely collected. Because this paper demonstrates the biostratigraphic utility of some of these forms, they may be better sampled in the course of future studies. Here, then, are six genera of spiriferinacean bra- chiopods that may prove useful in future paleoecolog- ical and biostratigraphic studies of the Triassic System. BIOSTRATIGRAPHY All specimens discussed herein were recovered from institutional collections, and their occurrence in the field was documented by workers whose primary in- terest was not necessarily biostratigraphic, or even pa- lecontologic. Thus, much of the information relating to ‘lecting localities themselves is interpretive and was | from evidence recovered subsequent to the ing itself (see Appendix, material in brackets). obable age assignments for the rocks recovered )calities discussed herein vary in their precision; yme are made on the basis of stratigraphic position, and others on the basis of associated fauna. Dagyspirifer fascicostata, n. gen. and n. sp., and Vi- iimetula parva, n. gen. and n. sp., each known from a single locality, occur, respectively, in the earliest No- rian and late Norian [zones LNol, and UNol through Choristoceras crickmayi Zone (UNo3) Cochloceras amoenum Zone (UNo2) Gnomohalorites cordilleranus Zone (UNo1) Himavatites columbianus Zone (MNo2) Drepanites rutherfordi Zone; Cyrtopleurites bicrenatus Zone (MNo1) Juvavites magnus Zone (LNo3) Malayites dawsoni Zone; Malayites paulckei Zone (LNo2) Mojsisovicsites kerri Zone; Guembelites jandianus Zone (LNo1) Klamathites macrolobatus Zone (UC3) Tropites welleri Zone; Tropites subbullatus Zone (UC2) Tropites dilleri Zone (UC1) Sirenites nanseni Zone (LC3) Austrotrachyceras austriacum Zone (LC2) Trachyceras desatoyense Zone; Trachyceras aonoides Zone (LC1) UNo3 of Tozer (1980); see Table 1]. Their full bio- stratigraphic ranges are not known. Pseudospondylospira perplexa, n. gen. and n. sp. oc- curs in beds that range from early late Karnian to latest Norian in age [zones UCI through UNo3 of Tozer (1980)], but most occurrences are in the early Norian. The precision of dating of these localities is, for the most part, insufficiently high to make this form a re- liable biostratigraphic indicator. Spondylospira Cooper, 1942 has commonly been used as an indicator for the Late Triassic, and with one possible exception (Hoover, 1990), that is still true. S. lewesensis (Lees, 1934), the best known species of the genus, has been recovered in this study alone from 22 localities ranging in probable age from earliest Karnian through latest Norian [zones LC] through UNo3 of Tozer (1980)]. The precision of the age assignments on the early end of this range is unknown. Only a single locality (loc. 26, Alexander terrane) includes rocks that may be older than late Karnian [zone UCI of Tozer (1980)]. This date is based on analysis of associated fauna by Silberling (written commun., 1989). Dagys (written commun., 1990) states that the “‘distribution {herein] of S. /ewesensis through all Late Triassic is quite unusual for Triassic cyrtinoid brachiopods. In Tethys, as well as in Boreal region, they are indicators only of part of one stage. Very suspect is distribution of /ewesensis in Carnian.”’ It may be: (1) that the ages presented here are correct; (2) that age assignments of some of the localities included herein are too broad or incorrect; or (3) that some of the material may belong to other taxa. I believe alternative (3) to be unlikely. S. parmata Hoover, 1983 is known from only two localities, but seems such a distinctive form that it could be a reasonable indicator for the middle to late Norian [zones MNol through UNo3 of Tozer (1980)]. S. tricosta, n. sp. qualifies as a very good index fossil. In this study it occurs at six localities in three different terranes, and nowhere is found earlier than the earliest Norian nor later than the late early Norian [zones LNol LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 67 through LNo3 of Tozer (1980)]. Because it is morpho- logically highly distinctive and easily recognizable, it should find great utility as an index for the early No- rian. Zugmayerella Dagys, 1963 as a genus does not ap- pear to occur earlier than the early late Karnian [zone UCI of Tozer (1980)] nor later than the late middle Norian [zone MNo2 of Tozer (1980)], and there are interesting patterns within the species recognized here- in. Z.? sp. A is known from only two specimens, from two localities, one of which is of unknown age, and the other of which is dated as middle late Karnian [zone UC2 of Tozer (1980)] on the basis of associated fauna. Z. sp. is known from a single specimen recovered from beds that are dated from early late Karnian through middle early Norian [zones UC1 through LNo2 of Toz- er (1980)], based on stratigraphic position. Neither can be considered as a good index based on this small information base. Z. americana, n. sp., however, is known from four localities representing three terranes, all dated on the basis of associated fauna and covering the relatively short time range of from middle late Karnian through early early Norian [zones UC2 through LNol of Tozer (1980)]. Phenacozugmayerella mimuncinata, n. gen. and n. sp. is a particularly interesting story. It has a completely disjunct distribution. Four localities, representing the Alexander (loc. 6) and Izee (locs. 48, 49, 51) terranes, give an age spanning the Late Karnian [zones UCI through UC3 of Tozer (1980)], based on associated fauna in the former and on stratigraphic position in the latter, but these ages may not be so well constrained as represented. Eight other localities, representing the Paradise and Jungo terranes, and rocks overlying the Golconda allochthon, give an age mostly in the early Norian, with one locality each from the Paradise and Jungo terranes extending the range up through the mid- dle Norian [through zone MNo2 of Tozer (1980)]. EVOLUTIONARY RELATIONSHIPS OF THE BRACHIOPODS All of the brachiopods described herein are repre- sentatives of the families Spondylospiridae or Labal- lidae of the Superfamily Spiriferinacea. With the pos- sible exception of a single species of Spondylospira Cooper, 1942 from Peru (Hoover, 1990), none is known from strata older than Karnian or younger than No- rian. Where these two structurally bizarre spiriferina- cean families came from is a matter of conjecture. After many years of pondering this problem I am no nearer to a solution. How evolution proceeded within the group is somewhat easier to postulate. The two families are differentiated largely on the basis of the apparent presence in the Spondylospiridae of a large functional pedicle in the adult stage, while this feature is only rarely reported in the Laballidae. Dagys (1974) has reported a small pedicle foramen in Thecocyrtella Bitt- ner, 1892, and in Thecocyrtelloidea Yang and Xu, 1966 (pp. 113, 114, pl. 9, figs. 1-14; pl. 10, figs. 2-3), mul- tiple pedicle tubules that pierce the deltidium have been described. Although Yang and Xu state [p. 113] that Thecocyrtelloidea is typified “by . . . lack of dental plates (hence no spondylium),” Dagys (written com- mun., 1990) feels that a true spondylium is present. Thus it appears that if the descent of mutiple-foramen cyrtinoid spiriferinaceans is monophyletic, this occur- rence in the Ladinian of Gueizhou, China is a logical precursor to Spondylospira and related forms in the later Triassic. Dagys (1974, p. 284) demonstrates that the Spiri- ferinacea, like other groups of Paleozoic brachiopods that survived the great reorganization near the Perm- ian—Triassic boundary, continued to diversify through- out the Triassic. In the Early Triassic (Induan and Olenekian), the superfamily consisted of only a single genus; in the Anisian, it had eight genera; in the Lad- inian, nine; in the Karnian, 19; and in the Norian 12. Studies since 1974 have increased these numbers, but the pattern remains the same. Dagys (1974, Table 3, pp. 287-289) indicated that Zugmayerella had a fossil record covering the entire Norian, while Spondylospira was known only in the Upper Norian. From evidence developed herein, I think it is safe to extend the lower boundaries of both ranges, because good examples of the families Spondylospiridae (typified by Spondylos- pira) and Laballidae (typified herein by Zugmayerella) exist in the Karnian. Dagys (1974, p. 228) cites the obvious homeomorphy between Paleozoic Cyrtinidae (which extend from the Late Silurian through the Early Carboniferous) and the Late Triassic laballids [= La- ballidae + Spondylospiridae herein], but points out that there is no direct phylogenetic connection between the two lineages. Dagys (1974, p. 228) also discounts the possibility that punctae developed more than once in the spiriferids (i.e., that these forms were polyphy- letic), and believes that the Late Triassic spiriferina- ceans (including those discussed herein) evolved from spiriferinacean ancestors, not directly from homeo- morphically-similar cyrtinids. He further suggests that, although the jugal structures of Late Paleozoic spiri- ferinids have been insufficiently studied, one detail of these forms stands out — the jugum is V-shaped and has its apex directed toward the anterior margin; that is, its structure is intermediate between the jugum of cyrtinids and the jugum of younger spiriferinaceans. Within the Late Triassic spiriferinaceans that have ridges on part or all of the interarea, the course of evolution is clearer (Hoover, 1990). Dagysprifer fas- cicostata, n. gen. and n. sp. (Pl. 9, figs. 1-9), with its globose form and small ridged portion of the interarea, 68 BULLETIN 337 appears a likely precursor morphology to that of Spon- dylospira; it is known only from a single locality in the late Karnian of southeastern Alaska. Forms like Pseu- dospondylospira perplexa, n. gen. and n. sp. (Pl. 10, figs. 14-24) from the late Karnian to late Norian of Nevada, Oregon, and Alaska, have a slightly less glo- bose form and relatively larger ridged portion of the interarea, and they seem likely intermediates in this lineage. This leads through increased relative height of the ventral valve, broadening of the “‘interarea’’, in- creased development of multiple ““honeycomb”’ fora- mena at the ventral valve apex, and decreasing shell thickness, to forms like Spondylospira parmata Hoo- ver, 1983 (Pl. 10, figs. 8-13) in the late Norian. BIOGEOGRAPHY OF THE BRACHIOPODS AND IMPLICATIONS FOR TERRANE COMPARISONS The nine clearly recognizable species of cyrtinoid spiriferinaceans occur in a wide array of different tec- tonic blocks within the western Cordillera. Therefore they provide a new data set for biogeographic com- parisons among various Cordilleran terranes. This is the first such data set published for Early Mesozoic brachiopods of the Cordillera. Of the six spiriferinacean genera in two families of cyrtinoid brachiopods decribed in this paper, only two (Spondylospira Cooper, 1942, and Zugmayerella Da- gys, 1963) appear to be known outside North America, and of these, only Zugmayerella is cosmopolitan. In addition to North American occurrences, Spondylo- spirais also known from South America and the north- easternmost U.S.S.R. (Bychkov and Dagys, 1984; Dagys, written commun., 1990). The two spondylo- spirid genera Pseudospondylospira, n. gen. and Viti- metula, n. gen., are known from but a single locality each, in southeastern Alaska [zones UCI-—UNol of Tozer (1980); Alexander terrane] and Idaho [zones UNol1-UN6o3 of Tozer (1980); Wallowa terrane], re- spectively. Zugmayerella, when first described by Da- gys (1963), was reported from the Norian and Rhaetian of the Alps, the Carpathians, the Crimea, the Caucasus, northwestern Turkey, and the northeastern U.S. S. R. (see Bychkov et a/, 1976). It has since been recognized in Alaska (Wrangellia terrane), California (Eastern ‘amath terrane) and Oregon (Wallowa terrane) [this Chile (Chong D. and Hillebrandt, 1985), Peru unpublished data), Papua New Guinea ‘icoll, and Campbell, 1976), China (Ching ig, 1977), Tibet (Sun, 1981), and New Zealand impbell, 1990). Stanley’s (1979) report of Zugmay- rella in the Late Triassic of Nevada was based on my OWn erroneous preliminary identifications; in the light of the present report, these forms are probably better assigned to Phenacozugmayerella, n. gen. Phenaco- zugmayerella, n. gen., in many respects a homeomorph of Zugmayerella, is known from 12 localities in Alaska (Alexander terrane), Nevada (Paradise terrane, Jungo terrane, and Triassic cover on the Golconda Allo- chthon), and Oregon (Izee terrane). Three features of cyrtinoid spiriferinacean species biogeography are particularly striking and offer insight into terrane comparisons and paleogeography. First, true ““Tethyan” species are absent. Indeed, only one of the six genera reported here (Zugmayerella) includes species that occur in Tethys, but the closest Tethyan relative of an American species is from China, at the eastern end of Tethys. Dagys (written commun., 1990) suggests that “Zugmayerella is [a] cosmopolitan genus, but species are different in remoted [sic] regions — western Tethyan, eastern Tethyan (China), Boreal (Si- beria), eastern Pacific, and perhaps Notal. North American species were connected with Tethyan rather via boreal regions. True Tethyan species of cyrtinoid brachiopods (as well as other characteristic Tethyan species of this group — koninkinids, thecospirids, etc.) were absent in East Pacific region.” Second, some of the forms endemic to North Amer- ica occur in several different types of terranes. Phen- acozugmayerella, n. gen. occurs not only in the al- lochthonous Alexander, Paradise, and Izee terranes, but also in parautochthonous terranes such as the Jun- go, and in rocks associated with blocks such as the Golconda Allochthon, which were linked with North America and ‘‘autochthonous” by the Late Triassic. The distribution of Zugmayerella parallels that of the Late Triassic rhynchonellid brachiopod Halorella Bittner, 1884, whose geographic occurrences have been summarized by Ager (1964). Halorella occurs over a broad geographic range within the Tethyan seaway, and also has been recognized both in allochthonous Cordilleran terranes of eastern Oregon (Ager, 1964), and in rocks of the parautochthonous Jungo terrane (Burke and Silberling, 1973). The nine species of cyrtinoid spiriferinaceans re- ported herein are mostly confined to North and South America, including the terrane belt. The only taxo- nomic overlap with Western Pacific Norian assem- blages is with the species of Spondylospira from the Koryak Range of the northeasternmost U.S.S. R. Cyrtinoid spiriferinacean brachiopods show little bio- geographic evidence for linkages among terranes of the present-day eastern and western Pacific. Abundant evidence does exist for biogeographic “stitching” among the North American terranes — in- cluding biogeographic links among allochthonous, par- autochthonous, and craton-related terranes. Among the spiriferinaceans, several species illustrate these com- plex, overlapping distribution patterns particularly well. (1) Pseudospondylospira perplexa, n. gen. and n. sp. LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 69 is known from the allochthonous Alexander and Wal- lowa blocks. However, it also occurs in the Triassic cover on the Golconda Allochthon, which is thought by most workers to have been tectonically emplaced against the North American craton during Permian to Early Triassic time (Miller et a/., 1984; Brueckner and Snyder, 1985; Silberling et a/., 1987; Babaie and Speed, 1990). New structural and stratigraphic data (Ketner, 1990) indicate that parts of the Golconda Allochthon overthrust Lower Triassic strata as a result of Mid- or Late Mesozoic tectonism. However, the youngest time of initial emplacement is mid- to Late Triassic (Sil- berling et a/., 1987; see discussion under “Triassic cov- er on the Golconda Allochthon’’, p. 71). Thus, occur- rences of P. perplexa in rocks overlying the Golconda Allochthon represent localities whose paleogeographic position was either on or very near the Late Triassic North American continent. Pseudospondylospira perplexa also occurs in the par- autochthonous Jungo terrane of Nevada. Triassic stra- ta of the Jungo terrane have been identified by some workers as basinal equivalents of the Auld Lang Syne Group of the Golconda terrane, which was craton- bound by latest Triassic time (Silberling et a/., 1987). The depositional cyclicity within the Auld Lang Syne Group in turn has been correlated with that of the North American Chinle Formation (Lupe and Silber- ling, 1985). An alternative recent interpretation by Ol- dow, Bartel, and Gelber (1990) is that the basinal Early Mesozoic rocks of the Jungo terrane (““Lovelock as- semblage” of Oldow, Bartel, and Gelber) are alloch- thonous relative to the neighboring craton-related plat- formal sediments of the Golconda terrane. However, Oldow (1984) has interpreted the Jungo—Lovelock Ear- ly Mesozoic sediments as having been thrust eastward during closure of a marginal basin behind the Sierra Nevada arc, and thus, even this “‘allochthonous”’ in- terpretation of Jungo strata does not involve large-scale longitudinal displacement of Jungo rocks. Therefore, the Jungo terrane is probably parautochthonous. Note that the occurrence of P. perplexa in the Jungo litho- tectonic block is significant in bridging the boundary between the craton-related Golconda terrane and the more outboard, truly allochthonous Cordilleran blocks such as the Alexander terrane. In summary, then, P. perplexa is here reported from occurrences that rep- resent craton-bound North American rocks, parauto- chthonous rocks, and allochthonous rocks. (2) Spondylospira lewesensis (Lees, 1934) is known from a broad range of allochthonous terranes within the North American Cordillera, including Eastern Klamath, Alexander, Paradise, Wrangellia, Wallowa, and Chulitna. It also occurs in the parautochthonous Jungo terrane (see discussion above), and is found in rocks overlying the Golconda allochthon, which by Late Triassic time was likely bound to the North Amer- ican craton (see discussion above). In addition, S. /ew- esensis is known from the Stikine terrane (where it was first described), which is inferred to have amalgamated with Cache Creek by Late Triassic time and may have accreted to North America during Jurassic time (Mon- ger, Price, and Templeman-Kluit, 1982; Cordey et al., 1987). Paleomagnetic data from Permian and Triassic strata of the Stikine block initially were thought to indicate significant northward latitudinal displacement of Upper Triassic and younger rocks. Late Triassic reefs described by Reid (1985) further suggest an origin for Stikine far south of the terrane’s present position. However, May and Butler (1986) have shown that when revised North American Upper Triassic poles are used, the Stikine Triassic results are in agreement with ex- pected cratonal inclinations. Comparable results have been obtained for Permian rocks of the Stikine block (Irving and Monger, 1987). Thus, paleomagnetic data show no indication of major post-Permian translati- tudinal motion for Stikine, leading some workers to speculate that Stikine may have ‘evolved near, but seaward of, the margin of North America” (Samson et al., 1989, p. 705). The presence of S. /ewesensis in the Stikine block and in the Golconda terrane shows that this species inhabited North American parauthoch- thonous and cratonal habitats as well as occurring in more distant allochthonous terranes. Thus, S. /ewes- ensis provides yet another biogeographic tie between cratonal North America and the allochthonous ter- ranes. (3) Spondylospira tricosta, n. sp. also occurs in ter- ranes of diverse origins. It is known from the craton- bound Triassic strata overlying the Golconda allo- chthon, but it is also represented in the Paradise terrane of Nevada and the Wallowa terrane of northeastern Oregon and Idaho. The distribution of S. tricosta com- pares with that of Pseudospondylospira perplexa, n. gen. and n. sp. and S. /ewesensis in that this species also occurs in both craton-bound (Golconda) and “‘sus- pect”’ terrane localities. In summary, the absence of ““Tethyan”’ species and the complex, overlapping biogeographic patterns of spiriferinacean brachiopods in this study compare fa- vorably with patterns previously described for Cor- dilleran bivalves by Newton (1987). The most signif- icant pattern documented in this study is the biogeographic “stitching”? of North American craton- bound brachiopod faunas with those of the parauto- chthonous and allochthonous Cordilleran terranes. These shared brachiopod species suggest that either many terranes originated close to the North American continent, or that there was large-scale dispersal of faunas within the proto-Pacific ocean basin. Evidence for the latter is also suggested by apparent species co- 70 BULLETIN 337 occurrences in Spondylospira in North America and in the northeasternmost U.S. S.R., and similarities between North American and Chinese species of Zug- mayerella. TECTONOSTRATIGRAPHIC SETTING OF LOCALITIES The following discussions of North American ter- ranes are ordered geographically, from south to north, and are followed by a brief discussion of the marine sedimentary rocks of the Cerro de Pasco region of Peru, from which Late Triassic cyrtinoid spiriferinacean bra- chiopods have also been recovered. In these discus- sions, a terrane is a descriptive unit having no inherent genetic implications (sensu Dover, 1990). JUNGO TERRANE [Churchill and Pershing counties, Nevada] (locs. 18, 39, 45) The Jungo terrane of northwestern Nevada preserves an Early Mesozoic stratigraphy of deformed Upper Triassic-Lower Jurassic fine clastic strata that repre- sent basinal facies (Silberling, 1990). This terrane, as defined by Silberling et a/. (1987) and Silberling (1990), is the equivalent of the “Lovelock assemblage” dis- cussed by Oldow (1984) and Oldow, Bartel, and Gelber (1990). Lupe and Silberling (1985) and Silberling ef al. (1987) infer that these basinal Jungo strata are the distal equivalents of the Auld Lang Syne Group sed- iments that overlie the Golconda Allochthon. Oldow, Bartel, and Gelber (1990) infer that the Late Mesozoic Fencemaker thrust, which separates the Early Meso- zoic rocks of these two terranes, coincides approxi- mately with the sedimentological transition between equivalent platformal (Golconda) and basinal (Jungo) strata. The Upper Triassic sedimentary units of the Jungo terrane include the lower Norian Hollywood Formation and overlying Antelope Springs Formation, the middle Norian Lori Formation, and the upper No- rian to Lower Jurassic Packard Wash Formation (Ol- dow, Bartel, and Gelber, 1990). PARADISE TERRANE [locs. 8-11, 33-38, 40] (Mineral and Nye counties, Nevada) Nevada terrane was initially assigned k within the Walker Lake terrane of Sil- : (1987), but recently Silberling (1990) el- ted the Paradise to terrane status. This terrane, as scognized by Silberling (1990) and Silberling et al. (1987), is equivalent to the Sand Springs, Pamlico, Luning, and Gold Range lithotectonic assemblages of Oldow (1984). The Paradise terrane consists of Permo- Triassic volcanogenic rocks, overlain by Upper Tri- assic to Lower Jurassic platformal carbonates that also include clastic and volcanogenic components (Silber- ling, 1990). Upper Triassic formations within the Par- adise terrane include the Luning and Gabbs forma- tions, both of which yield cyrtinoid spiriferinacean brachiopods. The stratigraphy of the Luning—Gabbs sequence was first described by Muller and Ferguson (1936; 1939) and, in part, has most recently been re- viewed by Taylor et a/. (1983). The Luning, principally of Norian age, represents a carbonate platform with interspersed clastic sediments; it is overlain by clastic sediments of the upper Norian Gabbs Formation, which is overlain in turn by lowermost Jurassic clastics of the Sunrise Formation (Taylor ef a/., 1983; Silberling et al., 1987). Much biostratigraphic work has been done on rocks of the Gabbs and Sunrise formations, because the Triassic-Jurassic boundary coincides approxi- mately with the formational contact (Guex and Taylor, 1976; Hallam, 1981; Laws, 1982; Taylor et al., 1983; Tozer, 1988). The Paradise terrane is clearly, in the strict sense, allochthonous with respect to cratonal North America, because it is bounded by thrust faults (e.g., Silberling and John, 1989). Exposures of Paradise terrane de- scribed by Silberling and John (1989) occur west of the Sr,;=0.706 isopleth, which is interpreted as the west- ernmost extent of Precambrian continental basement (Levy and Christie-Blick, 1989, and references there- in). However, the magnitude of post-Early Mesozoic displacement relative to the North American continent is not known, and the Mesozoic rocks of the Paradise are often interpreted as parautochthonous rather than allochthonous; for example, Speed, Elison, and Heck (1988) infer that the Paradise rocks were deposited atop an allochthonous Paleozoic basement after su- turing to North America. Some tectonic studies (e.g., Silberling, 1990) have also preferred a fringing-arc in- terpretation of the volcanogenic sediments of this and related terranes, in which these Nevada terranes orig- inated close to the American continental margin but were tectonically reshuffled later in the Mesozoic. One specific variant on this theme was proposed by Oldow (1984), who inferred that the thrusting of the Luning— Fencemaker and other western Nevada allochthons re- flects closure of a back-arc basin behind (eastward of) the Sierra Nevada arc. In this last view, although there may have been hundreds of kilometers of southeast- directed thrusting, the western Nevada terranes would essentially be parautochthous with respect to cratonal North America. More recent work by Oldow, Bartel, and Gelber (1990) has shown that, although there are lithostratigraphic differences among the western Ne- vada terranes (including the Paradise terrane), prov- enance linkages between stratigraphic units do exist — LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 71 for example, between the platformal Triassic sedi- ments of the Golconda terrane and correlative sub- marine fan deposits of the Jungo terrane. Biogeograph- ic evidence from ammonoid cephalopods supports the idea that the Triassic Paradise strata originated near North America (Tozer, 1982). TRIASSIC COVER ON THE GOLCONDA ALLOCHTHON [locs. 32, 41-43] (Lander and Pershing counties, Nevada) The Golconda Allochthon is a Carboniferous to mid- Permian package of sedimentary and volcanic rocks in thrust contact with an older, more eastern Paleozoic allochthon (the Roberts Mountains Allochthon) or its Upper Paleozoic cover. Triassic strata overlying the Golconda Allochthon rest in unconformable contact on a deformed Carboniferous to mid-Permian package of deep-water pelagic rocks, turbidites, and pillow la- vas (Silberling et a/., 1987) variously interpreted as a collapsed back-arc basin or an accretionary prism (see Miller et a/., 1984; and Brueckner and Snyder, 1985, for discussions of each model). Cyrtinoid spiriferinaceans occur within the Upper Triassic Cane Spring Formation of the Star Peak Group and the Osobb, Dun Glen, and (possibly) Winnemucca formations of the Auld Lang Syne Group (as restricted by Oldow, Bartel, and Gelber, 1990). The Cane Spring includes carbonates that represent a range of shallow- water environments (Nichols and Silberling, 1977), whereas the overlying Auld Lang Syne clastic sedi- ments were deposited in partly deltaic, to shallow sed- imentary environments (Burke and Silberling, 1973). Three lines of evidence suggest that the Golconda Allochthon was emplaced against cratonal North America during Late Permian to Early Triassic time. First, stratigraphic evidence presented by Lupe and Silberling (1985) suggests that cratonal, nonmarine sediments of the Chinle Formation in the Great Basin show sedimentary cycles correlative with those of the marine Auld Lang Syne Group, which overlies the Gol- conda Allochthon. Second, the age of the youngest sediments in several areas of the Golconda terrane has been used to con- strain the timing of emplacement of the allochthon, an event generally referred to as the Sonoma orogeny (Sil- berling and Roberts, 1962; Snyder and Brueckner, 1983). In the Schoonover sequence, for instance, the youngest basinal Paleozoic deposits are of Early Perm- ian age, leading Miller et a/. (1984) to conclude that thrust-faulting of the basin occurred in post-Early Permian time. Burke and Silberling (1973) suggest that emplacement occurred during the Late Permian or Ear- ly Triassic, a conclusion that has been supported by numerous other workers (e.g., Speed, 1977; Brueckner and Snyder, 1985; Brueckner, Snyder, and Boudreau, 1987; Speed, Elison, and Heck, 1988). However, there is considerable evidence that deformation of sediments within the Golconda terrane may have been diachron- ous (e.g., Brueckner and Snyder, 1985) and also that the emplacement itself may have occurred at slightly different times in various areas of Nevada. There is also evidence for some post-Triassic movement of rocks in both the Golconda and Roberts Mountains terranes during later orogenic episodes (Ketner, 1984, 1990; Ehman, 1985, p. 145; Little, 1987; Thorman et al., 1990). A third, and more compelling line of evidence for a Triassic or earlier emplacement of the Golconda ter- rane is that the Upper Triassic Auld Lang Syne clastics were derived from an eastern, low-lying continental source — possibly in the Colorado Plateau or Rocky Mountain regions (Compton, 1960; Silberling and Wallace, 1969; Burke and Silberling, 1973). The com- position of these clastics differs from that expected for western Nevada terranes derived from local sources (Burke and Silberling, 1973). Thus, even if the timing of emplacement was diachronous and spanned some- what more than the Permian—Early Triassic range usu- ally assigned to the Sonoma orogeny, an eastern source for Upper Triassic Auld Lang Syne Group clastics (Burke and Silberling, 1973) suggests that by Late Tri- assic time the Golconda terrane adjoined the North American craton (Lupe and Silberling, 1985). EASTERN KLAMATH TERRANE [locs. 1-5, 44] (Shasta county, California) The Eastern Klamath terrane, which represents a Paleozoic—Early Mesozoic island-arc sequence in northern California, preserves a Triassic through Mid- dle Jurassic sedimentary section of carbonate, shale, and coarse volcaniclastic sediments (Sanborn, 1960; Silberling et a/., 1987). Cyrtinoid spiriferinacean bra- chiopods occur in two formations within the Triassic part of the section in Shasta County: the Hosselkus Limestone and the Devil’s Canyon Member of the Modin Formation. The Hosselkus consists of lime- stones and shales and is probably in conformable con- tact with the Pit Shale below, while it is only ques- tionably conformable with the lithologically similar, but more shaly, Brock Shale above. The Brock appears to be in conformable contact with the Modin For- mation above. Sanborn (1960) divided the Modin into three subunits, from oldest to youngest, the Hawkins Creek Member (volcanics and conglomerate), the De- vil’s Canyon Member (tuffaceous limestone, lime- stone, and calcareous sandstone), and the Kosk Mem- ber (argillites, sandstones, and volcanics). 72 BULLETIN 337 IZEE TERRANE [locs. 48, 49, 51] (Suplee area, east-central Oregon) Triassic strata in the Suplee area of east-central Or- egon are assigned to the Begg and Brisbois formations and the Rail Cabin Argillite (Dickinson and Vigrass, 1965). The Begg Formation, including mudstone, con- glomerate, volcaniclastic rocks, lava, and rare bioclas- tic limestone, rests unconformably on unnamed Pa- leozoic rocks. The Begg contains possibly redeposited mollusks and corals of probable early Karnian age and has a conformable contact with the overlying Brisbois Formation, in part of late Norian age. The Brisbois, lithologically similar to the Begg, has produced all the spondylospirid brachiopods recovered from the area. It is unconformably overlain by the Rail Cabin Argil- lite, of middle to late Norian age, which is overlain concordantly by the Lower Jurassic Graylock For- mation. Hettangian fossils in the basal strata of the Graylock suggest that the uppermost beds of the Rail Cabin could extend upward into the earliest Jurassic. Silberling (written commun., 1990) suggests that ‘‘most of the Late Triassic shelly fossils in the Begg and Bris- bois are reworked,” and that “the Rail Cabin is evi- dently a facies of part of the Brisbois, as interpreted by Dave Taylor.” WALLOWA TERRANE flocs 125152212 28547] (Wallowa County, Oregon, and Nez Perce and Lewis counties, Idaho) The Wallowa terrane, a Late Paleozoic—Early Me- sozoic volcanic arc with overlying Triassic—Jurassic limestones and clastic sediments, contains cyrtinoid spiriferinacean brachiopods of both early and late No- rian ages. Within this terrane, early Norian brachiopod localities occur in Hells Canyon and in the Wallowa Mountains, whereas the only late Norian locality is an isolated limestone quarry, often called the Lewiston locality (Stanley, 1979, p. 25), which is far to the north- east of the other Triassic carbonate outcrops. The var- ious Wallowa localities are discussed in further detail below. Early Norian (locs. 12, 21, 28, 47).—In the Wallowa Mountains of northeastern Oregon, Triassic strata in- clude the Martin Bridge and Hurwal formations, and 's informally designated the Lower Sedimentary Se- [LSS] (Smith and Allen, 1941). The LSS in part rlies, and in part may be equivalent to the largely lover Creek Greenstone. The LSS is com- conformably overlain by the Martin Bridge Formation; and several hundred feet below that con- tact the LSS contains fossils suggestive of the middle upper Karnian Tropites welleri Zone. The Martin Bridge Formation as the name is applied in Hells Canyon and D mM +? rermMian & monly u the northeastern Wallowa Mountains, consists largely of shelf carbonates and includes typical early Norian faunas. It contains the only spondylospirid brachio- pods recovered from the area, and it is apparently con- formably overlain by the Hurwal Formation. The Hur- wal, in the northern Wallowa Mountains, contains several ammonoid faunas of from early to middle No- rian age (Silberling and Tozer, 1968), and it is overlain unconformably by Tertiary volcanics (Reeside et ai., 1957). Late Norian (loc. 15).—An isolated limestone quarry on the Nez Perce Reservation east of Lewiston, Idaho, probably representing a roof pendant in the Wallowa Batholith, contains scleractinian coral patch reefs with an associated fauna of brachiopods, bivalves, and gas- tropods. The coral-associated invertebrates represent an epifauna-dominated assemblage; bivalves from this locality consist of cemented and epibyssate (primarily nestling) species (Newton, in prep.). The paleoecology and structure of the scleractinian reefs has been doc- umented by Stanley (1979). The age of the carbonates at this locality is not well documented. Although no ammonoids have yet been recovered from this site, the reef-associated fauna seems to indicate a late Norian age. Haas (1953) noted, in particular, that the undescribed gastropod fauna at Lewiston closely resembles late Norian gastropods from Peru. Bivalves from the Lewiston site include no age- diagnostic species. A diverse fauna of largely unde- scribed brachiopods has been recovered from this site. Two species of cyrtinoid spiriferinaceans are present, one of which, S. /ewesensis (Lees, 1934), is of some biostratigraphic utility. The other, so far as is known, occurs only at this locality. The spatial isolation of this limestone block makes correlation with units in Hells Canyon (loc. 28) or the Wallowa Mountains (locs. 12, 21, 47) difficult. Indeed, if the late Norian age is correct, there are no time- equivalent rocks of comparable lithology in Hells Can- yon. Stanley (1979) has tentatively compared the Lew- iston sequence with the late Norian Hurwal Formation, though the lithofacies are not identical. Based on its location, most workers have presumed that the Lew- iston locality belongs to the Wallowa terrane, which encompasses nearby parts of the Wallowa Mountains and Hells Canyon areas (Silberling et a/., 1987). STIKINE TERRANE [locs. 29-31] (Lake Laberge region, Yukon Territory, Canada) The Stikine terrane is a Paleozoic through Early Me- sozoic volcanic-are terrane characterized by volcanic and sedimentary strata (Monger and Berg, 1987). The Mesozoic rocks of the Lake Laberge region, Yukon Territory, Canada, are divided into the Lewes River LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 73 Group (Triassic), the Laberge Group (Jurassic), and the Tantalus Formation (Upper Jurassic or Lower Cre- taceous). They are underlain by Permian limestones of the Braeburn Formation. The Lewes River Group, defined in this area by Lees (1934), has been divided into seven formations, from oldest to youngest des- ignated “A” through “G” (Tozer, 1958). These range in age from Karnian through Norian. Formations “C”’, “EE”, and ““G” consist mostly of limestone; ““B”’ and “—D” are composed mostly of clastic rocks; and “A” and ‘“‘F” contain both carbonate and clastic strata. Spondylospirid brachiopods have been recovered only from formations “‘F” and “G”’. ALEXANDER TERRANE [locs. 6, 7, 22-27] (Gravina, Keku, Kuiu, and Screen Islands and Cornwallis Peninsula, southeastern Alaska) The Alexander terrane of southeastern Alaska rep- resents a composite terrane with a complex Paleozoic history that suggests that the three major subterranes — Admiralty, Craig, and Annette — experienced dis- parate tectonic regimes in pre-Late Paleozoic time (Monger and Berg, 1987). Nonetheless, Pennsylvanian plutons common to both the Alexander and Wrangellia terranes indicate that at least part of the Alexander terrane (and perhaps the entire composite terrane) was united with the Wrangellia terrane at least by Late Carboniferous time (Gardner et a/., 1988). A further geographic constraint on the Alexander terrane comes from paleomagnetic evidence that the terrane was lo- cated far to the south in Pennsylvanian time (Van der Voo et al., 1980), but had arrived at its present latitude relative to North America by at least Triassic time (Hillhouse and Grommé, 1980). The Admiralty Subterrane, which contains the cyr- tinoid spiriferinacean brachiopod localities herein dis- cussed, includes Triassic strata of the Hyd Group and the Keku Volcanics. The Hyd Group is divided into the Burnt Island Conglomerate, Cornwallis Limestone, Hamilton Island Limestone, and Hound Island Vol- canics (Muffler, 1967). The Keku Volcanics consist of felsic and mafic volcanic rock and intercalated clastic units, which unconformably overlie the Permian Py- bus Formation. The upper part of the Keku Volcanics includes rare fossiliferous limestones, which contain early to late Karnian faunas. The Burnt Island Con- glomerate may be a lateral equivalent of the upper part of the Keku Volcanics and at one locality contains [in allochthonous pebbles] an early Karnian fauna. The Cornwallis Limestone, of late Karnian to earliest No- rian age, apparently overlies the Keku Volcanics con- formably and the Burnt Island Conglomerate uncon- formably. The Cornwallis is a gray oolitic limestone containing faunas of late Karnian to earliest Nonan age, and it is considered the shallow-water facies equiv- alent of the deep-water Hamilton Island Limestone (Muffler, 1967). Fossils from the Hamilton Island Limestone suggest ages equivalent to that of the Corn- wallis. Both the Cornwallis and Hamilton Island for- mations are apparently conformably overlain by the Hound Island Volcanics. This unit consists largely of basaltic pillow breccia and pillow lava, andesitic vol- canic breccia, and aquagene tuff, but includes some thin-bedded limestone. Fossils from these limestones suggest a late Karnian to late middle Norian age. The Hyd Group is unconformably overlain by unnamed Cretaceous lithic mudstones and sandstones. Silberling (written commun., 1990) suggests that “the complex facies relations implied by all of this probably indicate some substantial tectonic disruption of these rocks.” Cyrtinoid spiriferinacean brachiopods have been re- covered from the Cornwallis Limestone and from the Keku and Hound Island volcanics. WRANGELLIA TERRANE [locs. 13, 20] (Wrangell Mountains, southern Alaska and Nabesna quadrangle, south-central Alaska) Triassic strata in the Wrangell Mountains of south- central Alaska include the Nikolai Greenstone, the Chitistone and Nizina limestones, and the McCarthy Formation (Silberling and Tozer, 1968). Faunas from shaly beds near the base of the Nicolai indicate a late Ladinian to early Karnian age, while conformably overlying the Nicolai, the Chitistone Limestone con- tains 7ropites faunas near its base that are indicative of an early late Karnian age. Fossils from beds tran- sitional between the Chitistone and the overlying Ni- zina Limestone suggest a latest Karnian to earliest No- rian age. Fossils from the highest beds in the Nizina suggest the Juvavites magnus Zone of late early Norian age. Fossils in the lower part of the overlying McCarthy Formation are typical of the Rhabdoceras suessi Zone of late Norian age. Spondylospirid brachiopods have been recovered in this area from beds transitional be- tween the Chitistone and Nizina limestones and are thus early Norian in age. In the Nabesna quadrangle of south-central Alaska, Triassic fossils are reported from three horizons. The ‘‘Upper Triassic limestone” of Richter (1976) contains Monotis subcircularis Gabb, 1864, suggesting an early late Norian age for that unit. Lying unconformably below this unit, the ““Lower Triassic limestone” of Richter has yielded spondylospirid brachiopods refer- able to Spondylospira and has been assigned a late Karnian age. This unit is bounded unconformably be- low by the Nikolai Greenstone, an unfossiliferous unit consisting of volcanics and volcanogenic sediments. 74 BULLETIN 337 The Nikolai is in turn bounded unconformably below by the Eagle Creek Formation. The “Upper Triassic limestone” is unconformably overlain by beds of Late Jurassic to Cretaceous age which are assignable to the “Nutzotin Mountains sequence” of Berg, Jones, and Richter (1972). Wrangellia is demonstrably an allochthonous Cor- dilleran terrane, as attested by its distinctive internal stratigraphy of Paleozoic arc rocks overlain by Triassic basalt and Upper Triassic sedimentary strata (Jones, Silberling, and Hillhouse, 1977; Jones et al., 1987), and by its anomalous paleomagnetic signatures. Pa- leomagnetic data suggest Triassic paleolatitudes of 14— 18°, whereas for the Alaska segment of Wrangellia the expected Triassic paleolatitude would be approxi- mately 35-37° (Hillhouse, 1977; Yole and Irving, 1980; Hillhouse and Grommé, 1984; see compilations in Stone and McWilliams, 1989). Recent paleomagnetic modelling of the Jurassic and later motions of known oceanic plates suggests that if Wrangellia accreted to North America during Cretaceous time, Jurassic and younger longitudinal displacement of Wrangellia with- in the proto-Pacific ocean basin may have been as much as 60° (Debiche, Cox, and Engebretson, 1987). CHULITNA TERRANE [loc. 53] (Alaska Range, south-central Alaska) The Chulitna terrane consists of Upper Devonian ophiolite, overlain by Upper Paleozoic volcanic and sedimentary rocks, Triassic carbonate, terrigenous clastic, and basaltic rocks, and Jurassic clastic sedi- mentary rocks and chert (Jones ef a/., 1980; Jones et al., 1987). Within a complex Upper Triassic unit of interstratified pillow basalt and limestone, Spondylo- spira lewesensis (Lees, 1934) has been recovered in association with large (up to 20 cm) megalodontid bi- valves. Megalodontids are typical of shallow-water platformal facies. CERRO DE PASCO REGION [loc. 16] (Peru) Spiriferina steinmanni Tilmann, 1917 and Spirifer- ina sp. of Tilmann, 1917 (the latter here referred to Spondylospira tricosta, n. sp.), are both referable to the lylospirid genus Spondylospira Cooper, 1942 and -ecovered from beds in the Triassic—Jurassic section of Peru that Tilmann (1917, p. 644) kes and Haas (1944, p. 284) referred to the Szekeley and Grose (1972), in a detailed e-examination of these beds at three localities in the same region (Ninacaca; old trail connecting Carhua- mayo and Pasco; Yanamarca), suggest that most of the Jurassic Aramachay and Condorsinga formations are liddie Lias. missing here. Thus, the fossil-bearing sequence sam- pled by Tilmann is probably best referred to the Cham- bara Formation of the Pucara Group, which, on pa- leontological grounds, is of Norian age. Lower Jurassic (Sinemurian) ammonites recovered by Tilmann from the nearby San Blas area were not found in association with any spondylospirid brachiopods. At one set of localities south of Cerro de Pasco, however, an un- described species of Spondylospira is associated with a sponge fauna that suggests a Jurassic age (Hoover, 1990). Thus, South America may be the only place spondylospirids are not limited to the Upper Triassic. SYSTEMATIC PALEONTOLOGY INTRODUCTION Abbreviations of Repository Institutions The material discussed and described in this report comes from several sources and collections. These bear locality and/or catalogue numbers with acronyms as follows: CASG: California Academy of Sciences, Geology Col- lections, San Francisco, CA. GSC: National Type Collection of Fossil Invertebrates and Plants, Geological Survey of Canada, Ottawa, Ontario, Canada. LSJU: Leland Stanford Junior University, Stanford, CA. USGS: United States Geological Survey, U.S. Na- tional Museum of Natural History, Washington, DC. USGS M: United States Geological Survey, Menlo Park, CA. USNM: U.S. National Museum of Natural History, Smithsonian Institution, Washington, DC. (other): Various collections, identified only by field number, currently housed in the collections of the U. S. Geological Survey, Denver, CO. Measurement Abbreviations and Subscripts In the tables of measurements: a.v. = articulated valves; d.v. = dorsal valve; v.v. = ventral valve. Sev- eral subscripts are used to qualify measurements. The subscript . indicates that the measurement was esti- mated and is probably correct only to the nearest mil- limeter. The subscript . indicates that the specimen was crushed in such a way as to render the measure- ment non-representative; if . is also present, the value given was estimated to correct this deficiency. The sub- script ,, indicates that the specimen is broken and im- plies that the missing portion is necessary to that mea- surement; this may also be accompanied by the subscript . with the same meaning as given above. A measure across a structure with bilateral symmetry LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 75 (e.g., brachiopod hinge width) may be estimated by doubling the half-measure [distance from symmetry plane to distal extremity]. Use of this procedure is indicated by the subscript ,, . Terminology of Types The terminology of types used in this report follows that of Frizzell (1933). Of the types defined therein, only the chorotype is likely to be unfamiliar to the reader. Frizzell (p. 646) defined it as: “‘A fossil speci- men collected from the same stratum as the type, but from a neighboring locality.” Size as a Taxonomic Character Two samples of a continuous species population, taken from temporally or spatially disjunct localities, may exhibit different mean and range in individual size. With such material, there is no way, a priori, to determine whether one is dealing with two genetically- distinct species or with a single size-variable one. In- dividuals that differ on/y in size are here considered conspecific; and that size as a basis for specific dis- crimination, taken alone, is considered invalid. In many monotypic new genera, however, the diagnosis of the type species indicates the mean size of the individuals. This implies that size is of significance in specific dis- crimination. Specific discrimination by size, however unrealistic, is a necessary approximation (as are many paleonto- logical species) in the initial stages of paleontological investigation. The advantage of biological and pale- ontological taxonomy is that it is not static, but is cumulative, and may be emended as more data become available. Thus, in this report, the type species of a monotypic new genus may be diagnosed as “‘small’’, based solely on the subjective opinion that other, larg- er, possibly distinct congeneric species populations re- main to be sampled and described. ““Lumpers” and “Splitters” I do not think of myself as either a “splitter”, a manufacturer of perhaps unnecessary new taxa, nor as a “lumper’’, including new forms within the broad potential morphological variation of a given taxon. Several of the taxa reported herein are new genera represented by single new species, and several of these are represented by relatively few specimens from a small number of localities. The relatively bizarre mor- phological characters of many of these forms suggest to me that they are distinct at the generic level, but the thought remains that I do not have much material on which to base this suggestion. I am left with the task, which is the task of all taxonomists, of deter- mining the closeness of relationship of the forms under consideration to each other and to previously-de- scribed ones, and have reluctantly decided that they are indeed new, and not just on the fringes of variation of existing forms. I am comforted by the belief that taxonomic names, whatever else they are in genetic terms, are “handles” for discussion of a definite group of specimens, and that their correspondence to genetic groups becomes increasingly vague as the temporal distance from the Recent increases. This would not be a problem if all I was doing was describing morphology and making morphological comparisons. I am, however, making paleobiogeo- graphic inferences based on these taxonomic assign- ments, and the taxonomic assignments favor the pa- leogeographic arguments presented. I feel it necessary to state that the taxonomic assignments presented herein were made on morphologic grounds, with no a priori thought to their paleobiogeographic conse- quences, but also that I take full responsibility for those assignments. SYSTEMATICS Order SPIRIFERIDA Waagen, 1883 Suborder SPIRIFERIDINA Waagen, 1883 Superfamily SPIRIFERINACEA Davidson, 1884 Family SPONDYLOSPIRIDAE, new family Diagnosis.—Cyrtiniform spiriferinaceans with large functional pedicle in the adult. Delthyrium open or closed by deltidial structures. Additional posterior covering plates (collectively the cooperculum; see Text- fig. 2) rarely preserved. Dental plates converging an- teriorly to join strong median septum as spondylium; septum continuing posterior to junction of dental plates. Jugal supports attached to valve floor. Spires of brach- idium directed posterolaterally or ventrally. Remarks.—The genera Spondylospira Cooper, 1942 and Vitimetula, Dagyspirifer, and Pseudospondylo- spira, new genera, are included in the new family Spon- dylospiridae. The erection of a new family is justified by evidence that all four genera possessed a compar- atively large functional pedicle in the adult stage, a feature rare in other non-Paleozoic cyrtinoid spirifer- inaceans. The size of the structure is indicated by the area taken up by inferred pedicle foramena. A fora- menate condition has not been reported in the related family Laballidae, except for the two types of foramena cited for the Thecocyrtellinae. Among the genera as- signed to the Thecocyrtellinae (Thecocyrtella Bittner, 1892; Thecocyrtelloidea Yang and Xu, 1966; Klipstei- nella Dagys, 1974), only Thecocyrtelloidea appears to possess other than the normal single pedicle opening. In Thecocyrtelloidea numerous “pedicle tubules” (Yang and Xu, 1966, pl. 9, fig. 14, and pl. 10, fig. 2) appear to pierce the deltidium throughout its length, a scheme different from that seen in the Spondylospiridae. 76 BULLETIN 337 Views of the cooperculum of Spondylospira lewesensis (Lees, 1934). A, thin-section, x12, 11.7 mm dorsad of ventral beak, sural plane, of ventral valve, GSC 69374D (chorotype, from loc. 30), showing ridged interarea, dental plates, and median if rior portion of the spondylium, deltidial plates, and severely distorted cooperculum. Note that the shell structure of all )f the shell is fibrous and laminar, while the cooperculum is clearly punctate externally and laminar internally, as is the (shown only as fragment at extreme upper left of figure), except for the ridged portion of the interarea; B, posterior view, = 14.5, tion of ventral valve, GSC 69373 (chorotype, from loc. 30), showing punctate surface of cooperculum, its irregular mesial suture (ticks and bottom), and the lateral rugae and vertical furrows on its surface. (enlargement, upside down, of Pl. 9, fig. 21]. LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 77 The proximal portions of the pedicle in the Spon- dylospiridae are of two types, which form the basis for definition of two new subfamilies, the Dagyspiriferinae and the Spondylospirinae. Each subfamily is repre- sented by two or more genera and species, which in other respects exhibit considerable homeomorphy. In- dividuals of genera currently assigned to the Laballidae may, on closer examination, prove to be spondylo- spirids rather than laballids. Subfamily DAGYSPIRIFERINAE, new subfamily Included genera.—Dagyspirifer, n. gen.; Pseudo- sponadylospira, n. gen. Diagnosis.—Spondylospirid spiriferinacean bra- chiopods with paired pedicle foramena; foramena ovate, internally rimmed by callus thickening. Shells commonly globose, inflated, rarely pyramidal. Delti- dial or additional posterior covering plates not ob- served. Remarks.—The Dagyspiriferinae may be distin- guished from the Laballidae by their pedicle foramena, from the Thecocyrtellinae by the number and size of these pedicle openings, and the apparent lack of pos- terior covering or deltidial plates in these forms, and from the Spondylospirinae by the paired, rather than multiple pedicle foramena, and the globose, rather than pyramidal shell habit. Text-figure 3.— Dorsoposterolateral view (prepared using a camera lucida), x13, of ventral valve of Dagyspirifer fascicostata, n. gen. and n. sp., USNM 450274 (paratype, from loc. 23), showing one of the two small, paired, ovate pedicle foramena that pierce the apices of both the ridged portion of the interarea and the dental plates. The thin ridge that appears to bisect the foramen is the edge of a very narrow deltidial plate. The dorsal edges of the spondylium are bro- ken. Genus DAGYSPIRIFER, new genus Etymology of name.—Honoring A. S. Dagys, Lith- uanian student of Triassic brachiopods. Description.—Globose shells with subpentagonal outline. Surface macro-ornament pauciplicate. Micro- ornament of co-marginal growth lamellae and fine pus- tules. Ventral valve deep, swollen in lateral view, with distinct sulcus bounded by largest costae. Sulcus pro- duced anterodorsally as tongue producing W-shaped commissure in anterior view. Beak produced consid- erably posterior to hingeline, commonly incurved strongly, reaching to commissural plane. Interarea low, small, narrow, vertically ridged, bordered anteriorly by row of hinge denticles. Delthyrium wide for the subfamily, open, rimmed by narrow apparent deltidial plates. Spondylium as for subfamily. Paired pedicle foramena at delthyrial apex piercing apical portions of both ridged interarea and dental plates (i.e., spondy- lium) and opening into shell interior cavity on either side of median septum (see Text-fig. 3). Ventral interior surface radially plicate, reflecting ex- terior ornament. Muscle impressions indistinct. Dorsal valve unknown. Type Species.— Dagyspirifer fascicostata, n. sp. Diagnosis.—Globose Dagyspiriferinae with pauci- costate radial macro-ornament, distinct ventral sulcus floored by costae, and narrow ridged portion of inter- area, and small, oval, paired pedicle foramena. Occurrence.—As for species. Comparison.— Dagyspirifer, n. gen. is superficially most similar to Orientospira Dagys, 1965, from which it differs in its costate, rather than smooth ventral sul- cus, its delicate internal structures and its paired apical pedicle foramena. It differs from other cyrtinoid spi- riferinaceans in its markedly globose, rather than py- ramidal form. It differs from the other known dagys- piriferine, Pseudospondylospira, n. gen., in this respect, in the much narrower ridged portion of its interarea, and in its coarser macroornament. Dagyspirifer fascicostata, new species Plate 9, figures 1-9 new genus and new species of Early Norian brachiopod. Hoover, 1991, p. 394, fig. 2. Etymology of name.—L. fascis = bundle + L. costata = ribbed. Description.—Medium-sized globose shells; outline subpentagonal in dorsal view. Surface macro-orna- ment of four to six coarse costae on each flank, ante- riorly broadening from apex to commissure; low broad smaller costae arising from flanks of larger costae, broadening anteriorly. Micro-ornament of co-marginal growth lamellae and fine pustules. 78 BULLETIN 337 Table 2.— Measurements (in mm) of type specimens of Dagyspirifer fascicostata, n. gen. and n. sp. See p. 74 for explanations of abbreviations and subscripts. hinge width maximum width length depth specimen type locality 23 USNM 450274 (paratype) 93 18.4 14.7, 8.6 V.V. USNM 450275 (paratype) 10.6 24.2 19.1,. 10.7 v.V. USNM 450276 (holotype) 13.3 25.0 25.0 1322 V.V. Ventral valve deep, swollen in lateral view; having prominent flat-bottomed sulcus, floored by several very weakly rounded broad costae, bordered by two most prominent costae, these running distinctly from apex to anterior commissure, there produced dorsally as tongue. Commissure in anterior view W-shaped. Beak produced considerably posterior to hingeline, com- monly incurved to commissural plane. Interarea low, small, about one-half maximum shell width, vertically ridged, marginally rimmed by low, anteromesially-di- rected hinge denticles. Delthryium wide, open, rimmed by narrow apparent deltidial plates. Small spondylium formed by ventral convergence of dental plates; plates ridged on mesial surfaces, ridges reflecting previous positions of anterior edges of dental plates during on- togeny. Low median septum emergent from bottom 4.—Anterodorsolateral (interior) view (prepared using x 13%, of ventral valve of Dagyspirifer fascicostata, gen. and n. sp., USNM 450275 (paratype, from loc. 23), showing » horizontal bar (at right) that connects the dental plates and lateral vall. The opening above the bar connects to one of the two I -d pedicle foramena that pierce both the ridged portion of the interarea and the dental plate (see Text-fig. 3). The Y-shaped struc- ture at lower left is the posterior portion of the spondylium. mera 1ucica), furrow of spondylium, produced a short distance an- terodorsal of dental plates. Paired oval pedicle fora- mena at apex of delthyrium opening into shell interior. Ventral interior with thin median septum extending about one-half valve length. Surface radially costate, reflecting exterior macro-ornament. Thin, small, hor- izontal plate joining apical dental plate to lateral shell wall (Text-fig. 4). Muscle impressions indistinct. Dorsal valve unknown. For measurements, see Table 2. Occurrence.—Dagyspirifer fascicostata, n. sp., has been recovered only from locality 23 (Alaska, Corn- wallis Limestone, Alexander terrane), from beds as- signed to zone LNol of Tozer (1980), based on asso- ciated fauna [Silberling, written commun., 1989]. Diagnosis.—Fascicostate Dagyspirifer. Types.—Holotype, USNM 450276; paratypes, USNM 4590274, USNM 450275. Comparison.— Dagyspirifer is at present monotypic. The fasciculate macro-ornament of D. fascicostata is assumed to be of significance only at the species level. D. fascicostata is probably more closely related to spe- cies of Pseudospondylospira, n. gen. than to those of Orientospira Dagys, 1965, even though it is superfi- cially homeomorphic with the latter. It differs from species of Orientospira in its paired pedicle foramena, its more complex macro-ornament and its more del- icate internal structures. Genus PSEUDOSPONDYLOSPIRA, new genus Etymology of name.—L. pseudo = false + Spon- dylospira. Description.—Subglobose, unequally biconvex, cos- tate shells with modestly developed dorsal fold and ventral sulcus. Ornament weaker on mesial than on lateral shell surfaces. Outline subsemicircular in dorsal view, with rounded dorsal umbo and pointed ventral beak. Valves meeting at acute angles on flanks, at right angles in mesial regions. Ventral valve high. Costae broadening distally. In- terarea duplex: laterally smooth, punctate, mesially vertically ridged, impunctate; inner ridged portion lat- erally rimmed by low serrate ridge in some specimens, anteriorly bordered by hinge denticles that terminate vertical ridges. Delthyrium high, narrow, open, with narrow, marginal, disjunct apparent deltidial plates. Paired ovate apical pedicle foramena piercing both LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 719 ridged interarea and apical portions of dental plates. Low extensions of median septum protruding poste- riorly from floor of spondylium. Dorsal valve ornamented as for ventral. Umbo low, broad. Ventral interior surface reflecting exterior ornament. Spondylial elements very thin, delicate. Paired ovate apical pedicle foramena bearing raised callus rims. Dorsal interior with bilobate anteroposteriorly stri- ate cardinal process, flanked by anteriorly produced articulatory shelf. Broad inner socket ridges flanking wide, open notothyrium. Descending branches of spire arising from dorsal margins of socket ridges, joined to valve floor by narrow ridge. Smooth ovate paired ad- ductor muscle scars. Surface reflecting external orna- ment. Type species.— Pseudospondylospira perplexa, ni. sp. Diagnosis.—Subglobose Dagyspiriferinae with cos- tate radial ornament, and large duplex interarea, and relatively large, paired, oval apical pedicle foramena. Occurrence.—As for species. Comparison.—Pseudospondylospira, n. gen., out- wardly resembles Spondylospira Cooper, 1942, very closely. In specimens that preserve the apex of the ventral valve, however, it may easily be distinguished from Spondylospira, by its paired, ovate, rather than multiple, slit-like pedicle foramena. It is otherwise dis- tinguished from Spondylospira by its more delicate shells, more globose, less pyramidal shape, and by the limitation of the ridged portions of its interarea to a smaller portion of the posterior face of the ventral valve. Some specimens of two species of Spondylospira [S. lewesensis (Lees, 1934) and S. parmata Hoover, 1983] preserve both deltidial plates and cooperculum, and the shell structure of the ridged portion of the interarea in Pseudospondylospira leads me to suggest that a cooperculum was present in the latter genus as well. Pseudospondylospira is distinguished from Da- gyspirifer, n. gen., by its less globose form, its costate, rather than paucicostate macro-ornament, its relative- ly large, paired, oval pedicle foramena, and by its less pronounced fold and sulcus development. Its differs from laballid spiriferinaceans by having pedicle fora- mena, and from thecocyrtelline spiriferinaceans by these foramena being paired, rather than single or mul- tiple, and by having an open delthyrium. Pseudospondylospira perplexa, new species Plate 10, figures 14-24 new genus and new species of Late Karnian to Late Norian bra- chiopod. Hoover, 1991, p. 394, fig. 3. Etymology of name.—L. perplexus = obscure, am- biguous. Description.—Subglobose, inequibiconvex, costate shells with modest development of dorsal fold and ventral sulcus. Costae weaker, narrower, more closely spaced on fold and sulcus than on flanks. Shell outline subsemicircular in dorsal view, with rounded dorsal umbo and pointed ventral beak. Commissural angle acute on flanks, about 90° mesially. Ventral valve transverse, wider than long; high, rounded, with tongue-like semicircular dorsal exten- sion of mesial sulcus. Costae low, rounded, gradually broadening distally. Small, pointed beak hooked pos- terior to hingeline. Interarea duplex; laterally smooth, mesially vertically ridged; in some specimens rimmed by low serrate ridge on lateral margins of mesial ridged portion; anteriorly bordered by articulatory hinge den- ticles. Delthyrium high, narrow, open, with narrow, marginal, disjunct apparent deltidial plates. Paired elongate oval pedicle foramena piercing ridged inter- area and apical portions of dental plates. Spondylium floored mesially by thin posterior extension of median septum; septum projecting somewhat dorsad of edges of dental plates. Dorsal valve bearing moderately strong to weak macro-ornament of radial costae; costae more narrow, weak, and closely spaced on fold than on flanks, in- creasing in number anteriorly by bifurcation, median costa rarely broader or higher than adjacent costae. Anterior commissure in dorsal view mesially resected by sulcal tongue of opposite valve. Umbo low, broad, produced slightly posterior to hinge; incurved only slightly ventrad to hingeline. Ventral interior surface reflecting exterior ornament of costae, smooth distal interarea and ridged mesial interarea. Elements of spondylium (dental plates, me- dian septum) thin, delicate. Pedicle foramena bearing raised callus rims; muscle impressions otherwise in- distinct. Dorsal interior with bilobate, anteroposteriorly stri- ate cardinal process, flanked by anteriorly-produced articulatory shelf. Shelf bearing denticulate dorsad step at about mid-length; narrow lengthwise furrows pos- terior to step; deep, oval articulatory fossae just an- terior to step. Broad inner socket ridges flanking wide, open notothyrium. Descending branches of spire aris- ing from dorsal margins of socket ridges, joined to valve floor in narrow ridge at about one-half valve length. Median myophragm separating smooth, ovate, paired adductor scars; myophragm variably devel- oped, low; may extend anteriorly about one-fifth valve length. Interior surface radially striate, reflecting ex- terior ornament. For measurements, see Table 3. Occurrence.— Pseudospondylospira perplexa, Nn. sp., has been recovered in the present collections from lo- calities 7 [Alaska, Cornwallis Limestone, Alexander terrane, zones UCI-LNol(?) of Tozer (1980), based on associated fauna], 18 [Nevada, Jungo terrane, zone 80 BULLETIN 337 Table 3.—Measurements (in mm) of type specimens of Pseudospondylospira perplexa, n. gen. and n. sp. See p. 74 for explanations of abbreviations and subscripts. eS owoo.._“ striate hinge maximum width interarea width width length thickness specimen type ee ee SSS SS SSS Ss locality 22 USNM 450277 (paratype) — Ips 20 24 15:3 a.v. USNM 450278 (holotype) 11.0 15.6 28, 24 22.5 a.v. LNo3 of Tozer (1980), based on associated fauna], 22 [Alaska, Cornwallis Limestone, Alexander terrane, zones UCI-—UNol of Tozer (1980), based on associ- ated fauna], 23 [Alaska, Cornwallis Limestone, Alex- ander terrane, zone LNol of Tozer (1980), based on associated fauna], 24 [Alaska, Cornwallis Limestone, Alexander terrane, zones UC1-—LNo2 of Tozer (1980), based on stratigraphic position], 25 [Alaska, limestone bed in Hound Island Volcanics, Alexander terrane, zones UC2-LNol of Tozer (1980), based on associated fauna], 27 [Alaska, Cornwallis Limestone, Alexander terrane, zones UCI-UNol of Tozer (1980), based on stratigraphic position], 28 [Oregon, Martin Bridge For- mation, Wallowa terrane, zone LNol of Tozer (1980), based on associated fauna], and 41 [Nevada, Osobb Formation, Triassic cover on the Golconda Alloch- thon, zones LNo2—LNo3 of Tozer (1980), based on stratigraphic position]. See Table 1 for Tozer’s (1980) zonal scheme. Diagnosis.—Transverse Pseudospondylospira with low dorsal fold and shallow ventral sulcus. Types.—Holotype, USNM 450278; paratypes, USNM 450277, USNM 450279, USNM 450280, USNM 450281, USNM 450282. Comparison.— Pseudospondylospira perplexa, n. sp., is at present the only species in the genus. The as- sumption is made that the outline and strength of fold and sulcus development are specific rather than generic characters. Mean size may also prove to be significant, but no conclusions can be made in the absence of large population samples of P. perplexa or specimens of oth- er congeneric species. Several specimens from a single locality in Oregon have been tentatively assigned to this species (see below). Their stronger macro-orna- ment and more pronounced fold and sulcus develop- “nt are near the limit of what I would consider nor- pecific variation. Pseudospondylospira perplexa, n. sp., ified from silicified material, which com- nly preserves the subfamilially diagnostic apex of ventral valve. Calcareous “‘crackout’”’ specimens, however, commonly lack this portion of the shell, and levelopment of such specimens from matrix involves ngthy and complex preparation. I have not identified specimens as P. perplexa unless, in addition to other criteria, the diforamenate condition could be ascer- tained. Hence, many individuals that may be P. per- plexa have been assigned to less restrictive taxonomic categories, and the cited occurrence of the species may not be representative of its true geographic and strati- graphic range. Pseudospondylospira aff. P. perplexa, new species Plate 10, figures 25-27 Remarks.—A small partial ventral valve and a larger complete dorsal valve of a form closely related to P. perplexa were recovered from locality 28 (Oregon, Martin Bridge Formation, Wallowa terrane) in beds assigned, on the basis of associated fauna, to zone LNol of Tozer (1980) by Silberling (written commun., 1989). The paired pedicle foramena clearly place these spec- imens within the Dagyspiriferinae, and the subglobose form and comparatively subdued macro-ornament suggest Pseudospondylospira. The macro-ornament is, however, stronger than that in P. perplexa, and is ac- companied by more pronounced development of the fold and sulcus. The range of variation in macro-or- nament observed in P. perplexa (see Pl. 10, figs. 20, 21) suggests this form may also lie within P. perplexa, but too few specimens are present to permit more con- clusive identification than the above. Types.— Figured specimens, USNM 450283, USNM 450284. Subfamily SPONDYLOSPIRINAE, new subfamily Included genera.—Spondylospira Cooper, 1942; Vi- timetula, n. gen. Diagnosis.—Spondylospirid spiriferinacean bra- chiopods with more than two pedicle openings. Del- tidial plates and coopercula may be present. Shells commonly subpyramidal, apically hooked, rarely glo- bose or inflated. Remarks.—The Spondylospirinae may be distin- guished from the Laballidae by their pedicle opening, from the Thecocyrtellinae by the size of that opening, from the Dagyspiriferinae by their multiforamenate rather than diforamenate pedicle opening, and in some species of Spondylospira from all others by the pres- ervation of two distinct kinds of coopercula (see Hoo- ver, 1983; and Text-fig. 2 and Plate 10, figs. 10-13 herein). LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 81 Genus SPONDYLOSPIRA Cooper, 1942 Type species.— Spondylospira reesidei Cooper, 1942, p. 232; Cooper, in Shimer and Shrock, 1944, p. 359, pl. 140, figs. 43-47. Diagnosis.—Spondylospirine spondylospirids with weak to strong dorsal fold and ventral sulcus, bearing ornament of strong costae on flanks and weaker costae on fold and sulcus. Interarea vertically ridged; del- thyrium commonly preserved as open, in some spec- imens covered by deltidial plates or deltidium; inter- area in some specimens shielded by multipartite shield (cooperculum). Hinge denticulate. Ventral interior with spondylium basally bisected by median septum; dorsal interior with descending branches of spire jugate an- teriorly, supported dorsally by meshwork connected to valve floor. Occurrence.—Spondylospira occurs in beds of from Karnian to Norian age in North and South America. Spondylospira cf. S. lewesensis (Lees, 1934) is the only species referred to the genus reported from outside the Western Hemisphere [by Dagys (written commun., 1990) from the Koryak mountains of the northeast- ernmost U.S.S.R.]. This species was originally re- ferred to Spondylospira alia (Hall and Whitfield, 1877) by Bychkov and Dagys (1984, p. 13). Comparison.—Well-preserved specimens of Spon- dylospira may be distinguished from other non-spon- dylospirine cyrtinoid spiriferinaceans by their multiple apical pedicle foramena. Less well-preserved speci- mens may be distinguished from Zugmayerella Dagys, 1963, Laballa Moisseiev in Dagys, 1962, Pseudola- balla Dagys, 1974, Spinolepismatina Dagys, 1974, and Orientospira Dagys, 1965, by their costate fold and sulcus. The latter characters also distinguish Spondy- lospira from the other known spondylospirine, Viti- metula, n. gen. Discussion.— Vokes and Haas (1944) referred the following species to Spondylospira: Cyrtina lewesensis Lees, 1934, Spiriferina acrotamboensis Korner, 1937, Spirifera (Spiriferina) alia Hall and Whitfield, 1877, Spiriferina canavarica Tommasi var. robusta Korner, 1937, Spiriferina koessenensis Zugmayer, 1882, Spi- riferina steinmanni Tilmann, 1917, Spiriferina sp. of Tilmann, 1917, Spiriferina(?) sp. indet. of K6rner, 1937, and Spondylospira reesidei Cooper, 1942. Cyrtina lewesensis Lees, 1934 is the name first ap- plied to the form which typifies Spondylospira. Widely variable in size, strength of costation, and number of costae, this species is the senior synonym of both Spi- riferina acrotamboensis [Pl. 9, figs. 27-29], and Spon- dylospira reesidei [Pl. 9, figs. 22-26]. Hoover (1983) reported that preparation of the ventral valve which is the holotype of S. (S.) alia Hall and Whitfield, 1877 [USNM 12671; Pl. 10, figs. 4-7] reveals a shell that may not even be punctate, a low, non-ridged interarea, non-denticulate hinge, and absence of the “‘cyrtinoid”’ spondylium that is the uniting characteristic of the laballids and spondylospirids. S. (S.) alia cannot be placed within either family, and only questionably fits in the Spiriferinacea. The specimen figured by Cooper (in Shimer and Shrock, 1944, pl. 141, figs. 48-51) as Spondylospira alia (Hall and Whitfield, 1877) is here placed in synonymy with S. /ewesensis (Lees, 1934). Specimens of S. canavarica robusta, S. steinmanni, and S. (?) sp. indet. of K6rner, were not available for comparison, but from the published illustrations sug- gest assignment to Spondylospira. Spiriferina sp. of Tilmann (1917) is herein referred to Spondylospira tri- costa, n. sp. Spiriferina koessenensis was removed from Spondylospira by Dagys (1963), who defined it as the type species of Zugmayerella. Spondylospira lewesensis (Lees, 1934) Plate 9, figures 10-38; Plate 10, figures 1-3 Cyrtina lewesensis Lees, 1934, p. 35, pl. 1, figs. 14-16. Spiriferina acrotamboensis Korner, 1937, p. 168, pl. 11, figs. 5-8. Spondylospira reesidei Cooper, 1942, p. 232; Cooper, in Shimer and Shrock, 1944 [part], p. 359, pl. 140, figs. 43-47 [USNM 103468a- c, e-g only; not USNM 103468d (= Vitimetula parva, n. gen. and n. sp.)]; Dagys, 1974, pl. 54, figs. 5-8; Hoover, 1983, figs. 3G-3K. Spondylospira alia (Hall and Whitfield). Cooper, in Shimer and Shrock, 1944, p. 359, pl. 140, figs. 48-51 [not Spirifera (Spiriferina) alia Hall and Whitfield, 1877, p. 281, pl. 6, fig. 17]. Spondylospira aff. S. alia Hall and Whitfield [sic]. Rangel Z., 1978, p. 25, pl. 1, figs. 3-5 [not Spirifera (Spiriferina) alia Hall and Whitfield, 1877, p. 281, pl. 6, fig. 17]. Spondylospira lewesensis Lees. Tozer, 1962, p. 27, pl. 12, figs. 11- 13; Dagys, 1974, pl. 51, figs. 9a—b; Hoover, 1983, pp. 1026-1028, figs. 3L-3S, 4E, 4F, 5. Spondylospira aff. S. acrotamboensis Korner [sic]. Rangel Z., 1978, p. 25, pl. 1, fig. 6, pl. 2, figs. 1-4. Spondylospira cf. S. acrotamboensis Korner [sic]. Rangel Z., 1978, p. 24, pl. 1, figs. 1, 2. Description.—Spondylospira lewesensis (Lees) has been amply described in the above accounts. Its dif- ferences from related species are detailed under Com- parison below. For measurements, see Table 4. Occurrence.—Spondylospira lewesensis (Lees) has been recovered from deposits of from early Karnian to late Norian age. In the present collections, it is rec- ognized from localities 5 [California, 7Modin Forma- tion, Eastern Klamath terrane, zones LNol1—UNo3 of Tozer (1980), based on associated fauna], 6 [Alaska, Alexander terrane, zones UC1—UC3 of Tozer (1980), based on associated fauna], 10 [Nevada, middle mem- ber of Gabbs Formation, Paradise terrane, zones UNo2-UNo3 of Tozer (1980), based on associated fauna], 13 (Alaska, Wrangellia terrane, age unknown), 15 [Idaho, Wallowa terrane, zones (?) UNol-—UNo3 of Tozer (1980), based on associated fauna (see Appendix for discussion of age uncertainty at this locality)], 16 [Peru, zones UCI-—UNo3 of Tozer (1980), based on associated fauna], 21 [Oregon, Martin Bridge Forma- 82 BULLETIN 337 Table 4.—Measurements (in mm) of specimens of Spondylospira lewesensis (Lees, 1934). See p. 74 for explanations of abbreviations and subscripts. dorsal costae on hinge width maximum width length thickness fold/flank specimen type locality 15 USNM 103468a' 9.1 13.7 8.4. 9.0. — v.v. USNM 103468b? 14.0, 15.0, 9.0, 3.0. 6/6 d.v. USNM 103468c? 14.1 14.1 6.1 3.0 4/6 d.v. USNM 103468d? 10.1 12.4 8.8 6. — V.V. locality 29 GSC 9619 (lectotype) 17.9 22.6 18.5 20.7 4/6 a.v. GSC 9619a (paralectotype) 17.0 2257 20.7 21 6/6 a.v. locality 30 GSC 69373 (chorotype) 14.9 22.4 23.8 17.8 6/6 a.v. locality 39 CASG 66253.02 6.8 7.9 6.4 6.3 4/6 a.v. CASG 66253.03 8.3 11.0 8.5 9.0. 4/6 a.v. CASG 66253.04 10.4, 12.3 9.1 9.2 4/6 a.v. CASG 66253.01 1S 14.0 10.0 9.5 6/7 a.v. locality 45 CASG 66258.05 6.8 8.3 etl 5.8 4/4 a.v. CASG 66258.01 7.6 8.6 5.4 6.2 4/4 a.v. CASG 66258.06 6.8 8.4 5.6 5.0 4/4 a.v. CASG 66258.07 8.0 9.2 7.5 6.3 4/7 a.v. CASG 66258.08 7.9 9.7 6.8 8.3 = a.v. CASG 66258.03 8.6 10.2 8.0 ed 4/6 a.v. CASG 66258.02 10.4 10.5 6.5 6.9 4/6 a.v. CASG 66258.09 10.4 11.5 8.2 8.0 4/6 a.v. CASG 66258.10 10.8 12.8 8.9 8.0 4/8 a.v. CASG 66258.04 13.2 13.8 8.5 9.8 6/6 a.v. ' Holotype of Spondylospira reesidei Cooper, 1942. ? Paratype of Spondylospira reesidei Cooper, 1942. tion, Wallowa terrane, zone UC3 of Tozer (1980), based on associated fauna], 26 [Alaska, limestone bed in Keku Volcanics, Alexander terrane, zones LCI-UNo?2 of Tozer (1980), based on associated fauna], 27 [Alaska, Cornwallis Limestone, Alexander terrane, zones UC 1—- UNol of Tozer (1980), based on stratigraphic posi- tion], 29 [Yukon, Formation “‘F” of Lewes River Group, Stikine terrane, zone UNo2 of Tozer (1980), based on associated fauna], 30 [Yukon, Formation “F” of Lewes River Group, Stikine terrane, zone UNo2 of Tozer (1980), based on associated fauna], 31 [Yukon, Formation “F” of Lewes River Group, Stikine terrane, ‘one UNo2 of Tozer (1980), based on associated fau- 3 | Nevada, lower member of Luning Formation, terrane, zone LNol of Tozer (1980), based it this stratigraphic level], 35 [Nevada, Lun- ag Formation, Paradise terrane, zone LNo3 of Tozer 980), based on associated fauna], 36 [Nevada, Lun- ing Formation, Paradise terrane, zone LNol1 of Tozer (1980), based on fauna at this stratigraphic level], 37 [Nevada, lower Luning Formation, Paradise terrane, zones LNol—LNo2 of Tozer (1980), based on fauna at this stratigraphic level], 39 [Nevada, Jungo terrane, zone LNo3 of Tozer (1980), based on associated fau- na], 40 [Nevada, Luning Formation, Paradise terrane, zone LNo3 of Tozer (1980), based on stratigraphic position], 41 [Nevada, Osobb Formation, Triassic cov- er on the Goldconda allochthon, zones LNo2—LNo3 of Tozer (1980), based on stratigraphic position], 45 [Nevada, Jungo terrane, zone MNo2 of Tozer (1980), based on associated fauna], 47 [Oregon, Martin Bridge Formation, Wallowa terrane, zones LNol—-MNo2 of Tozer (1980), based on associated fauna], and 53 [Alas- ka, Chulitna terrane, zone UC3 of Tozer (1980), based on associated fauna]. This indicates a range of zones LC1 through UNo3 of Tozer (1980), with ages in the older end of the range most common in the Alexander terrane. The early and middle Karnian extension of the range is supported by only a single locality (loc. 26), and Dagys (written commun., 1990) doubts that the species really ranges down into the Karnian at all. Occurrences that may be Karnian are limited to the Alexander and Chulitna terranes of Alaska. Diagnosis.—Spondylospira with strong dorsal fold costae that increase initially by equal bifurcation, sub- sequently by equal or unequal bifurcation; widest at or LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 83 slightly anterior to hingeline; fold and sulcus moder- ately to strongly developed; bears posterior cooper- culum on interarea, but cooperculum rarely preserved. Types.—Lectotype, GSC 9619; paralectotype, GSC 9619a; chorotype, GSC 69373; chorotype of S. acro- tamboensis Korner, 1937, USNM 450285; figured specimens, CASG 66248.01, CASG 66248.02, CASG 66251.01, CASG 66253.01, CASG 66258.01—CASG 66258.04, USNM 103468a, USNM 103468b, USNM 103468f, USNM 106521, USNM 450286, USNM 450287; measured specimens, CASG 66253.01-—CASG 66253.04, CASG 66258.01-CASG 66258.10, GSC 9619, GSC 9619a, GSC 69373, USNM_ 103468a- USNM 103468d. Comparison.— Spondylospira lewesensis (Lees, 1934) includes forms that vary considerably in most of the characters which define the species. It may most easily be distinguished from S. tricosta, n. sp., by the manner of initial increase in number of costae on the dorsal fold. S. tricosta adds a single subordinate costa on ei- ther side of a single initial costa, so that bilateral tri- furcation takes place. S. /ewesensis simply bifurcates the initial costa, and at least in the early ontogenetic stages, bears an even number of dorsal fold costae, as new ones are added symmetrically. S. parmata Hoo- ver, 1983 may fall within the range of S. /ewesensis. It was identified as a distinct species primarily because (1) it is close to the limits of variation of S. /ewesensis and (2) it exhibits features of particular interest that are not well shown in S. /ewesensis, although they may be present in some individuals of that species. S. par- mata may be distinguished by its weak development of the median fold and sulcus, and by its very numer- ous, thin, closely spaced costae. It bears a cooperculum that appears to be tripartite, as opposed to the bipartite set of elements seen in some specimens of S. /ewesensis. S. lewesensis includes multicostate forms referred by Cooper to S. alia (Hall and Whitfield, 1877). Hoover (1983) recognized that the type of Hall and Whitfield’s species was not a spondylospirid. In Spondylospira the dorsal interior spire support meshwork memorialized by the generic name is not always well-preserved. The beautiful silicification seen in the types of S. acrotamboensis (KOrner, 1937) and S. reesidei Cooper, 1942 is the exception rather than the rule. Thus specific characters have tended to be external. The types of S. reesidei (USNM 103468a-c, e-g) however, suggest how variable such external char- acters may be. In addition, it seems subjectively rea- sonable that individuals of this elusive genus from such geographically widely-separated regions as Peru, Ida- ho, and the Yukon Territory of Canada might, in the absence of other collections, belong to distinct species. Thus, S. /ewesensis (Lees) from the Yukon Territory of Canada was considered distinct from the Peruvian species S. acrotamboensis (K6rner, 1937), and both of those were considered distinct from S. reesidei Cooper. These three species can only be differentiated as fol- lows: S. acrotamboensis and S. lewesensis are larger, and have more lateral costae, than does S. reesidei. This distinction, as suggested above, is insufficient ba- sis for specific discrimination when a large database is available. S. /ewesensis and S. acrotamboensis have more (4—6 vs. 4—5) lateral costae than does S. reesidei. The latter phenomenon can be related directly to size. Discounting these differences, the three species above must be considered to form a single, externally vari- able, hemispherically cosmopolitan species, Spondy- lospira lewesensis (Lees). Discussion.—The type material of S. reesidei in- cludes individuals with two distinct types of ornament. Most bear strong, commonly sinuous costae, that bi- furcate anteriorly. A single ventral valve [USNM 103468d] bears only simple, very weak costae, and is comparatively smooth. Examination of topotypic ma- terial [loc. 15] has produced further specimens of this smooth form. As no transitional forms occur, the smooth form is here assigned to a distinct taxon. As it lacks a costate fold or sulcus, it cannot be assigned to Spondylospira [sensu stricto], and is therefore here placed in the new genus Vitimetula. Ventral valves of both forms show the multiple apical foramena char- acteristic of the new subfamily Spondylospirinae. Spondylospira parmata Hoover, 1983 Plate 10, figures 8-13 Spiriferina cf. S. suessi Winkler. Sanborn, 1960, pp. 20-21, pl. 2, figs. 7, 8 [not Spiriferina suessi Winkler, 1859 = Laballa Moisseiev in Dagys, 1962]. Spondylospira parmata Hoover, 1983, pp. 1025, 1026, text-figs. 3A— 3F; Hoover, 1991, p. 395, fig. 4. Description.— Pyramidal, inequibiconvex costate shell, with weak dorsal fold and ventral sulcus. Costae increasing in number distally by bifurcation. Interarea about half-covered by multipartite cooperculum, ex- posed apically and basally. Ventral valve about equal in length, width, and depth. Outline triangular in lateral, anterior, posterior, or ven- tral aspects. Surface macro-ornament of closely spaced fine costae; at 10 mm from beak, about 15 on each flank, and six narrower ones in shallow sulcus. Inter- area mostly planar, but beak strongly hooked poste- riorly at apex. Interarea surface vertically finely ridged; about 17 ridges in a 5 mm distance at base. Hingeline denticulate; denticles serving articulatory function. Delthyrium high, open, narrow. Cooperculum of single basal and two lateral plates, together covering entire width of basal two-thirds of interarea, exposing only oval apical mesial and low, transverse slitlike basal areas. Surface rugose, with ap- 84 BULLETIN 337 Table 5.—Measurements (in mm) of the holotype of Spondylospira parmata Hoover, 1983. See p. 74 for explanations of abbreviations and subscripts. maximum hinge width width interarea height length thickness specimen type locality 44 CASG 60975.01 (holotype) 21 be 2 ye parent suture-lines demarcating basal one or two plates, and disjunct right and left lateral plates. Basal plate relatively smooth, lateral plates co-marginally rugose around pedicle opening, bearing coarse rounded ver- tical grooves on lateral plates. Lateral plates thickest (ca. 2 mm) around pedicle opening, thinning distally and toward the hingeline. Dorsal valve with costate ornament in number and character like that of opposite valve. Low mesial fold developed. Hinge area bearing longitudinally-ridged articulatory shelf and striate cardinal process. Ventral interior with spondylium of dental plates, supported and divided longitudinally by median sep- tum. Interiors of either valve not otherwise observed. For measurements, see Table 5. Occurrence.—Only a single specimen confidently identifiable as S. parmata Hoover, 1983, is known. It was recovered from locality 44 (California, Devil’s Canyon Member of the Modin Formation, Eastern Klamath terrane), in beds assigned to zones UNol- UNo3 of Tozer (1980) on the basis of associated fauna (Silberling, written commun., 1989). Several other specimens which appear conspecific but have not pre- served the cooperculum characteristic of S. parmata have been recovered from locality 44, and from locality 2 (California, 7Modin Formation, Eastern Klamath terrane). Beds at the latter locality may be assigned to zones MNol through UNo2 of Tozer (1980) based on associated fauna (Silberling, written commun., 1989). Diagnosis.—Equidimensional Spondylospira with numerous fine costae, weak fold/sulcus development, and a tri-partite cooperculum covering portions of the ventral interarea. Types.— Holotype: CASG 60975.01. Comparison.—S. parmata may fall within the ex- tremely broad range of variation of S. /ewesensis (Lees), as discussed above. It is, however, near the limits of yee of that variation, and its numerous fine costae fold and sulcus development distinguish it ; of S. acrotamboensis, S. lewesensis, and S. {t is more similar in exterior ornament to spondylospira perplexa, n. sp., but differs from form in having an interarea that covers the max- num width of the shell, and in having multiple rather han paired pedicle foramena. Its tripartite coopercu- jum distinguishes it from all previously described spe- cies of the genus. Topotypic specimens of S. /ewesensis (Lees) also bear a cooperculum, which in that species 20.3 23. 29.2 ay. appears to be comprised of only two paired lateral plates. Spondylospira tricosta, new species Plate 11, figures 1-15 Spiriferina sp. Tilmann, 1917, p. 689, pl. 25, figs. 12a, 12b. Etymology of name.—L. tri = three + L. costa = rib. Description.—Small, inequibiconvex, paucicostate shells with distinct, costate dorsal fold and ventral sul- cus; commonly high, thicker than long; transverse, wider than long. Ventral valve outline subsemicircular in ventral as- pect, dorsally tongued in anterior aspect; pointed beak commonly projecting posteriorly. Distinct median sul- cus commonly bearing two weak radial costae; flanks bearing four to five stronger costae; mesial costae only originating at beak; others originating further dorsad on flanks of posterior margin. Interarea planar to con- cave posteriorly, bounded laterally by raised rim; ver- tically finely ribbed, ribs terminating dorsally in fine hinge denticles. Delthyrium narrow, open. Micro-or- nament of fine co-marginal growth lines and radial striae. Dorsal valve anteriorly truncate in lateral aspect; hinge width commonly slightly less than maximum width, subequal to length. Broad median fold com- monly bearing three costae on crest; mesial costa strongest, all arising about four mm anterior to beak by trifurcation. Flanks commonly bearing fine costae. Ventral interior with spondylium and posteriorly- produced median septum. Hinge teeth low, feeble, ar- ticulation chiefly by hinge denticles. Dorsal interior unknown. For measurements, see Table 6. Occurrence.—Spondylospira tricosta, n. sp., is known only from the early Norian [zones LNol through LNo3 of Tozer (1980)]. It has been recognized in the present study at localities 8 [Nevada, lower member of Luning Formation, Paradise terrane, zone LNol of Tozer (1980), based on associated fauna], 28 [Oregon, Martin Bridge Formation, Wallowa terrane, zone LNol of Tozer (1980), based on associated fauna], 32 [Nevada, Osobb Formation, Triassic cover on the Golconda AI- lochthon, zones LNol1-LNo3 of Tozer (1980), based on stratigraphic position], 40 [Nevada, Luning For- mation, Paradise terrane, zone LNo3 of Tozer (1980), based on stratigraphic position], 41 [Nevada, Osobb LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 85 Table 6.— Measurements (in mm) of specimens of Spondylospira tricosta, n. sp. See p. 74 for explanations of abbreviations and subscripts. costae on dorsal hinge width maximum width length thickness fold/flank specimen type locality 8 USNM 450288 17.0 2273. 17.6, Zee 5/5 a.v. locality 32 CASG 66247.01 (holotype) Wey 13.8 11.4 11.5 3/5 a.v. CASG 66247.02 (paratype) 10.6 15.4 11.5, 12.6 3/5 a.v. CASG 66247.03 (paratype) 10.2 17.3 13:0; 13.8 3/5 a.v. CASG 66247.04 (paratype) 14, 18.5 13.9 15.8 4/5 a.v. locality 40 CASG 66254.01 11.1 14.8 12.0 15.0 5/5 a.v. locality 41 CASG 66255.01 ale 16.0 132 15.6 3/5 a.v. CASG 66255.03 LES 16.5 16.6 14.0, 3/5 a.v. CASG 66255.04 14.1 16.5, 14.8 14.3 3/5 a.v. CASG 66255.05 14.2 18.1 14.5 16.8, 3/5 ay. CASG 66255.06 19.7 21.6 15.0 18. 3/5 a.v. CASG 66255.02 18.3. 18.8. 19.7, 24.0. 3/5 a.v. locality 42 CASG 66256.01 Ud 10.5, Sie 8.4 3/4 a.v. locality 43 CASG 66257.01 13.8 16.9 14.0 14.0 3/5 a.v. CASG 66257.02 14.8 17.6 17.0 16.6 3/5 a.v. Formation, Triassic cover on the Golconda Allo- chthon, zones LNo2-LNo3 of Tozer (1980), based on stratigraphic position], 42 [Nevada, Dun Glen For- mation, Triassic cover on the Golconda Allochthon, zone LNo3 of Tozer (1980), based on associated fau- na], and 43 [Nevada, ?Winnemucca Formation, Tri- assic cover on the Golconda Allochthon, zones LNo2- LNo3 of Tozer (1980), based on stratigraphic position]. It also appears to be present in rocks assigned to the ““Middle Triassic’ of Peru by Tilmann (1917), where it was reported as Spiriferina sp. The dorsal valve ex- terior figured by Tilmann (1917, pl. 25, fig. 12a), shows the trifurcate costae on the fold that are characteristic of this species. Diagnosis.—Paucicostate Spondylospira bearing an odd number of costae on the dorsal fold, which arise from bilateral trifurcation of a single initial mesial cos- ta. Types.—Holotype, CASG 66247.01; paratypes, CASG 66247.02—CASG 66247.04; figured specimens, CASG 66255.01, CASG 66255.02, CASG 66256.01, USNM 450289; measured specimens, CASG 66255.01-CASG 66255.06, CASG 66256.01, CASG 66257.01, CASG 66257.02, USNM 450288. Comparison.—S. tricosta, n. sp. may be distin- guished from most species of Spondylospira by its pau- cicostate macro-ornament. Among paucicostate cyr- tinoid spiriferinacean species, it may be distinguished from SS. /ewesensis (Lees) by the odd number of costae borne on its dorsal fold. It may be distinguished from Vitimetula parva, n. sp. by its more pronounced or- nament, and presence of a dorsal fold. It may be dis- tinguished from species of Zugmayerella Dagys, 1963, which it resembles in form and outline, by its costate fold and sulcus. Discussion.—S. tricosta, n. sp. is probably closest in overall appearance to S. /ewesensis, although no tran- sitional forms have been observed. As is the case in S. lewesensis, sporadic bifurcation of mesial and lateral costae results in both paucicostate and multicostate phenotypes, although the transition from one to the other is not so well-represented by samples as is the case with S. /ewesensis. Samples of S. tricosta from Nevada most resemble species of Zugmayerella Dagys, 1963, differing most obviously in bearing costae on the fold and sulcus. Genus VITIMETULA, new genus new genus aff. Spondylospira. Hoover, 1983, p. 1026. Etymology of name.—L. vietus = wrinkled + L. me- tula = conical or pyramidal figure. Description.—Strongly inequibiconvex shells with high spire-like ventral and shallow bowl-like dorsal valves. Both valves may bear faint simple costae; cos- tae more pronounced distally. Distinct, shallow, very narrow ventral sulcus, extending from apex to anterior commissure; no corresponding dorsal fold. Ventral valve outline variable in ventral aspect. In- terarea margins laterally rimmed; interarea surface pla- nar, bearing coarse, sinuous vertical ridges, which may 86 BULLETIN 337 bifurcate commissurally, ridges terminating at hinge in prominent simple denticles. Delthyrium high, nar- row, open. Apical pedicle foramena multiple, “hon- eycombed” as typical of subfamily. Dorsal valve quite shallow, commonly widest at hinge. Surface smooth or distally faintly costate. Ventral interior with shallow spondylium. Articu- lation by hinge denticle and by anterior portions of dental plates that apparently match hinge sockets of opposite valve. Surface smooth or faintly reflecting exterior ornament. Muscle scars indistinct. Dorsal interior posteriorly bearing short articulatory shelf; shelf bearing anteroposteriorly elongate fossae that accomodate articulatory hinge denticles of op- posite valve. Shallow hinge sockets present. Median septum absent, but low median myophragm separating muscle scars. Umbo poorly developed, not produced posterior to hinge line. Descending branches of spire produced anteriorly from anterior margins of hinge sockets, anteriorly forming a jugum. All elements joined to valve floor by complex meshwork. Type species.— Vitimetula parva, n. sp. Diagnosis.—Faintly costate Spondylospirinae with high ventral and shallow dorsal valves, narrow shallow ventral sulcus but no dorsal fold. Interior as for Spon- dylospira Cooper. Occurrence. — Vitimetula, n. gen. is at present known only from locality 15 (Idaho, Wallowa terrane), where it co-occurs with Spondylospira lewesensis (Lees, 1934). The fauna at this locality is probably of late Norian age (Stanley, 1979) [zones UNol-UNo3 of Tozer (1980). See Appendix for discussion of problems with the age of this locality.]. Comparison.—Vitimetula, n. gen. is easily distin- guished from Spondylospira by the absence of a dorsal fold and the absence of costae on the mesial portions of either valve. It is distinguished from other laballids and spondylospirids that have smooth mesial portions by its narrow sulcus and comparatively smooth surface overall. It is distinguished from Psioidea Hector, 1879 by the absence of a dorsal fold and its relatively short hingeline and non-transverse outline. Discussion.—A single specimen of the type lot of Spondylospira reesidei Cooper [USNM 103468d] is here issigned to Vitimetula, n. gen. Vitimetula parva, new species Plate 11, figures 16-32 reesidei Cooper, 1942, p. 232 [part: unfigured para- fi 1M 103468d)]; Cooper, in Shimer and Shrock, 1944, p. > [part: unfigured paratype (USNM 1034684d)]. Description.—Small, strongly inequibiconvex shells with high conical ventral and shallow bowl-like dorsal valves. Both valves bearing faint simple costae; costae more pronounced distally. Distinct, very narrow, shal- low ventral sulcus, but no corresponding dorsal fold. Ventral valve commonly four times as deep as dor- sal; outline in ventral aspect variable. Margins of in- terarea commonly produced posteriorly as low mim; posterodorsal hinge margins commonly flared distally. Interarea bearing coarse, sinuous, dorsally bifurcate vertical ridges; ridges terminating at hinge in promi- nent simple denticles. Apical portion of area pierced by several posteroventrally oriented slitlike openings (see Pl. 11, fig. 18). Delthyrium long, narrow, open. Dorsal valve quite shallow, commonly widest at hinge; surface smooth or marginally faintly costate. Ventral interior with shallow spondylium formed by junction of short dental plates and median septum; septum length variable in terms of spondylium sup- port, invariably projecting posteriorly into spondy- lium. No functional paired hinge teeth, but anterior portions of dental plates apparently articulating with hinge sockets of opposite valve. Articulation mostly by hingeline denticles. Muscle scars not evident. In- terior surface smooth or faintly reflecting subdued ex- terior costation. Dorsal interior posteriorly bearing short articulatory shelf. Shelf bearing anteroposteriorly elongate fossae which accomodate articulatory hinge denticles of op- posite valve. Shallow hinge sockets present. No median septum, but low rounded myophragm separating stri- ate muscle scars. Umbo poorly developed, not pro- duced posterior to hinge line. Descending branches of spire produced anteriorly from anterior margins of hinge sockets, anteriorly forming a jugum, all elements joined to valve floor by complex meshwork. For measurements, see Table 7. Occurrence.— Vitimetula parva, n. sp. is at present known only at locality 15 (Idaho: Wallowa terrane), where it co-occurs with Spondylospira lewesensis (Lees, 1934). The fauna there suggests a late Norian age [Stan- ley (1979); zones UNol-UNo3 of Tozer (1980), based on associated fauna, but see Appendix for discussion of problems with this age]. Diagnosis.—Small Vitimetula. Types.—Holotype, USNM 450290; paratypes, USNM 103468d, USNM 450291-USNM 450298. Comparison.— Vitimetula, n. gen. is monotypic. V. parva, n. sp. may be distinguished from species of Spondylospira by its smooth narrow ventral sulcus and its lack ofa dorsal fold, and from other spondylospirids and laballids by its high ventral valve, smooth exterior, and narrow ventral sulcus. Its internal details indicate that it is probably most closely related to Spondylos- pira. Its ventral valve somewhat resembles that of Zug- mayerella americana, n. sp., but consistently bears a narrower sulcus, while the dorsal valves of the two forms are markedly dissimilar. LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 87 Table 7.—Measurements (in mm) of type specimens of Vitimetula parva, n. gen. and n. sp. See p. 74 for explanations of abbreviations and subscripts. height/depth or hinge width maximum width length thickness specimen type locality 15 . USNM 450290 (holotype) 4.8 6.3 6.0 10.2 a.v. USNM 450291 (paratype) 5:5 5:5 8.2 6.2 v.V. USNM 450292 (paratype) 5.4 5.4 3.8 4.5 V.V. USNM 450293 (paratype) 6.04. 6.05. SiS 6.3 v.V. USNM 450294 (paratype) BiOne 3/6 3.7 4.2 V.v. USNM 450295 (paratype) 5:5 5:5 5.0 1.0 d.v. USNM 450296 (paratype) 7.0 7.0 4.8 135. d.v. USNM 450297 (paratype) 6.0, Ue. 4.5. Dis d.v. USNM 450298 (paratype) 9.0, 9.0, => 1.6 d.v. USNM 103468d (paratype) 8.0 8.0 6.0. 4.8 V.V. Family LABALLIDAE Dagys, 1962 Subfamily LABALLINAE Dagys, 1962 Genus ZUGMAYERELLA Dagys, 1963 Type species.—Spiriferina koessenensis Zugmayer, 1882, p. 28, pl. 3, figs. 2, 3, 5, 13. Diagnosis.—Small to medium-sized laballines with high ventral valve, smooth, sharply differentiated fold and sulcus, and paucicostate flanks. Interior as for Spondylospira Cooper, 1942. Microornament of densely packed spines. Occurrence.— Zugmayerella has been recovered from Norian strata in the Alps, the Carpathians, the Balkans, the Dinarides, Crimea, the Caucasus, northeastern U.S.S.R., Papua New Guinea, China, Tibet, New Zealand, and South America. In the present collec- tions, Zugmayerella and a species questionably re- ferred to that genus have been recovered from seven localities in Alaska, California, and Oregon, ranging in age from late Karnian to early Norian. Comparison.— Zugmayerella may be distinguished externally from Spondylospira Cooper, 1942, by its smooth, rather than costate fold and sulcus. It differs from Vitimetula, n. gen. in its more transverse outline, broader sulcus, and presence of a dorsal fold. It differs from Spinolepismatina Dagys, 1974, and Pseudola- balla Dagys, 1974, in having a longitudinally ridged, rather than smooth interarea. It differs from K/ipstei- nella Dagys, 1974, in its denticulate hinge, and absence of a deltidium, and from Laballa Moisseiev in Dagys, 1962, by its denticulate hinge and by the strong pli- cation of both valves. It differs from Orientospira Da- gys, 1965, in its subpyramidal rather than globose shell form, and its comparatively high, narrow interarea. It differs from Phenacozugmayerella, n. gen. in its spi- nose, rather than capillate—cancellate microornament. Discussion.— Placement of the genus Zugmayerella in the Laballidae rather than the Spondylospiridae is tentative. It was so assigned by Dagys (1962), and the critical characters of the ventral apex necessary to dis- tinguish between the two families are not well-shown in any of the specimens of Zugmayerella available for this study. It is, however, possible that better preser- vation in specimens recovered in the future may dictate that this genus be placed in the Spondylospiridae with the other genera described in this paper. Zugmayerella was originally distinguished from Lepismatina Wang, 1955 (Dagys, 1963, pp. 99-100) on the basis of its ridged interarea and resultant denticulate hinge, fea- tures which Dagys believed were not shared by Lep- ismatina. Later examination by Sun (1981) of topo- typic material of MLepismatina revealed that Lepismatina does not belong in the Laballidae, but rather in the Spiriferinidae. One character common to known species of Zug- mayerella is spinose microornament (Dagys, written commun., 1990). This character, though mentioned in the original diagnosis of the genus (Dagys, 1963), was not there accorded the importance brought out by later observations (Dagys, written commun., 1990). The spines in Zugmayerella are small and easily broken, and a pustulose surface microornament is commonly the result. Pearson (1977, p. 23), in one of the few comprehensive discussions of the genus in English, states that the shell surface is “‘pustulose, possibly spi- nose.” It is this character that, among others, sets this genus apart from the new genus Phenacozugmayerella, in which the cancellate microornament is formed by beaded capillae. Zugmayerella americana, new species Plate 11, figures 33-50; Plate 12, figures 1—5 Description.—Small, unequally biconvex shells hav- ing high, pyramidal ventral and shallow bowl-like dor- sal valves. Ventral sulcus and dorsal fold only slightly developed. Microornament of densely-packed spines. Ventral valve semicircular in ventral aspect, trian- gular in anterior aspect. Depth commonly subequal to shell width, slightly less than height. Median sulcus shallow, rounded, bounded laterally by large plicae. 88 BULLETIN 337 Table 8.—Measurements (in mm) of type specimens of Zumayerella americana, Nn. sp. See p. 74 for explanations of abbreviations and subscripts. ee hinge width maximum width length thickness specimen type locality 3 USNM 450305 (paratype) 5.8), 5.8), 4.1 4.0 V.V. locality 20 USNM 450299 (paratype) 4.8, 6.0, 4.1 3.9 V.V. USNM 450300 (paratype) 6.2 6.8 4.4 4.54. V.V. USNM 450301 (paratype) 11.8 13.2 10.0 9.1 V.V. USNM 450302 (paratype) 10.5 14.1 10.3 3.4 d.v. USNM 450303 (paratype) 8.8 13.3 1253 3.2 dv. locality 21 USNM 450304 (paratype) Wes) 8.4 6.5 4.3 V.V. locality 28 USNM 450306 (holotype) 5.4 5.4 51 4.4 a.v. USNM 450307 (paratype) 7.9, TES 4.6 4.7 V.V. USNM 450308 (paratype) 4.6 7.6 6.1 4.6 V.V. USNM 450309 (paratype) 9.6, 12.0 10.0 9.8 V.V. USNM 450310 (paratype 122) 14.1 10.0 9.7 V.v. USNM 450311 (paratype) 10.0 11.3 7.6 3.2 dv. USNM 450312 (paratype) 7.9. 11.6 8.8 2.3 d.v. USNM 450313 (paratype) 10.6 11.6 10.0 2.6 d.v. Flanks bearing four to six prominent plications. In- terarea sharply recurved posteriorly at apex, otherwise flat; entire surface vertically ridged, ridges terminating as denticles at hinge. Delthyrium open, narrow. Hinge denticulate; primary teeth absent, anterior portions of dental plates may be marginally functional in articu- lation. Dorsal valve quadrate to transverse or semicircular in dorsal aspect, broadly rounded in anterior or lateral aspects. Umbo tiny, not produced posterior to hinge- line. Median fold low, square-shouldered, medially de- pressed; bordered laterally by four to six prominent rounded plications; fold commonly lower than adja- cent plicae. Plicae originating from central portion of hingeline, but not from mesially-located “point-source” beak. Ventral interior generally reflecting exterior macro- ornament. Thin median septum extending one-half to two-thirds valve length, may be asymmetric with re- spect to sulcus. Dental plates united with median sep- tum just below its crest, forming small spondylium. Muscle scars indistinct. Dorsal interior with simple, anteroposteriorly striate | process. Fossae accomodating denticles of ngeline anteroposteriorly elongate, may be 1esially. Hinge sockets present, probably nally functional in articulation. Low median my- phragm present or absent, if present extending about me-half valve length. Descending branches of spire joined to valve floor at about mid-length of valve, united anteriorly to that point by simple jugum. For measurements, see Table 8. Diagnosis.—Small Zugmayerella with pyramidal, comparatively high ventral valve, shallow, flat-bot- tomed ventral sulcus and low, mesially depressed dor- sal fold, and four to six lateral plications. Dorsal umbo not produced posterior to hingeline. Occurrence.— Z. americana, n. sp. has been recov- ered in this study from rocks ranging in age from mid- dle late Karnian [zone UC2 of Tozer (1980)] to earliest Norian [zone LNol of Tozer (1980)], at localities 3 (California, 7Hosselkus Formation, Eastern Klamath terrane, zone UC2 of Tozer (1980), based on asso- ciated fauna], 13 (Alaska, Wrangellia terrane, age un- known), 20 (Alaska, beds transitional between Chiti- stone and Nizina formations, Wrangellia terrane, zone LNol of Tozer (1980), based on associated fauna], 21 [Oregon, Martin Bridge Formation, Wallowa terrane, zone UC3 of Tozer (1980), based on associated fauna], and 28 (Oregon, Martin Bridge Formation, Wallowa terrane, zone LNol of Tozer (1980), based on asso- ciated fauna]. All specimens recovered are silicifed. The fine preservation of this form wherever it is found and its comparatively wide distribution, coupled with its relatively short stratigraphic range, combine to make it a fine potential biostratigraphic index. Etching of material in suitable matrix should provide usable spec- imens of this small but important form. Types.—Holotype, USNM 450306; figured para- types, USNM 450300-450302, USNM 450305, USNM 450308, USNM 450309, USNM 450311, USNM 450313; measured paratypes, USNM 450299- USNM 450313. Comparison.— Zugmayerella americana, n. sp. may be distinguished from most known species of the genus by its lower fold and shallower sulcus, and by lacking LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 89 a dorsal umbonal bulge that projects posterior to the hingeline. It is smaller and less globose than Z. eurea Dagys, 1965, Z. inaequiplicata Dagys, 1965, or Z. koessenensis Zugmayer, 1882. It is smaller and has more lateral plications than does Z. hemipyramida Sun, 1981, and is smaller, and has plications that are less angular than those of Z. uncinata (Schafhautl, 1851). It is smaller, narrower, and lacks the characteristic rib in the sulcus and furrow on the fold that are charac- teristic of Z. osmana (Bittner, 1902). Among the other North American cyrtinoid spiri- ferinaceans, it is easily distinguished from specimens of Phenacozugmayerella mimuncinata, n. sp., by its smaller size and less acute plications and by the lack of spinose ornament in the latter form. From all the species of Spondylospira Cooper, 1942, and from Pseu- dospondylospira perplexa, n. gen. and n. sp., Dagys- pirifer fascicostata, n. gen. and n. sp., and Vitimetula parva, n. gen. and n. sp., it is distinguished by its ap- parent lack of pedicle foramena in the ventral valve. The small shell of Zugmayerella americana, n. sp. is most similar to Z. yueliangpingica Ching and Feng, 1977 from the Upper Triassic of Gueizhou, China, from which it differs in size. The latter form, described on the basis of only four specimens, two ventral ex- ternal molds, and one dorsal external mold, and one dorsal internal mold, ranges from 15 to 20 mm wide, while the largest known specimen of Z. americana is just over 14 mm wide. Discussion.— Zugmayerella americana, n. sp. be- longs to the only cosmopolitan cyrtinoid genus among those described in this report. Its close similarity to the Chinese Z. yueliangpingica Ching and Feng, 1977 may suggest origin via trans-Pacific dispersal. Zugmayerella species Plate 12, figure 34 Discussion.—A single silicified dorsal valve [USNM 450317] assignable to Zugmayerella, but not confi- dently associated with any species, shows the sup- porting structures of the brachidium in best detail. The descending branches of the spire extend anterodorsally from the hingeline and contact the valve floor at about midlength. A simple jugum unites the branches ante- riorly. It was recovered from locality 1 [California, ?Hosselkus Formation, Eastern Klamath terrane, zones UCI-LNo2 of Tozer (1980), based on stratigraphic position]. Measurements (in mm)*.— Hinge width, 10.6,,.; valve width, 10.6,. ; valve length, 8.8,..; valve height, 5.0. Type.— Figured specimen, USNM 450317. *see p. 74 for explanations of abbreviations and subscripts. Table 9.— Measurements (in mm) of specimens of Zugmayerella? sp. A. See p. 74 for explanations of abbreviations and subscripts. maxi- speci- hinge mum men width width length height type locality 3 USNM 450315 122 17.6, 15. — V.V. locality 12 USNM 450314 10.5 14.2 12.7 11.6 V.V. USNM 450316 7529 24315 13.7, IWS 2 V.V. Zugmayerella? species A Plate 12, figures 25-33 Discussion.— Three partial ventral valves (one silic- ified, two calcareous) of a new species here question- ably assigned to Zugmayerella were recovered from localities 3 [California, 7Hosselkus Formation, Eastern Klamath terrane, zone UC2 of Tozer (1980), based on associated fauna] and 12 (Oregon, ?Martin Bridge Formation, Wallowa terrane, age unknown). All ap- pear to have been somewhat decorticated, but repre- sent a pauciplicate form with a smooth broad sulcus and very faint lateral plications. The rarity and the poor preservation of this form make formal species designation unwarranted here, and the lack of un- equivocally preserved microornament make generic identification questionable. For measurements, see Table 9. Types.— Figured specimens, USNM 450314, USNM 450315; measured specimens, USNM 450314-USNM 450316. Genus PHENACOZUGMAYERELLA, new genus Etymology of name.—Gr. phenax = cheat, impostor, + Zugmayerella. Type species.— Phenacozugmayerella mimuncinata, Nn. sp. Diagnosis.— Medium-sized laballines with high ven- tral valve, smooth, sharply differentiated fold and sul- cus, narrow denticulate hinge, and paucicostate flanks. Surface microornament cancellate, with capillae bro- ken into transverse bars by growth increments (Text- fig. 5). Occurrence.—As for species. Comparison. — Phenacozugmayerella is perhaps most easily confused with Zugmayerella Dagys, 1963, from which it differs in its microornament, which is capil- late-cancellate rather than densely spinose. In this re- spect is also differs from most other laballids, except aspinose forms like Yanospira Dagys, 1977, which dif- fers in having a duplex interarea, with the central por- tion ridged, and the outer part smooth, and by being more transverse. It may be distinguished from Spon- dylospira Cooper, 1942, by its smooth, rather than 90 BULLETIN 337 Table 10.—Measurements (in mm) of type specimens of Phenacozugmayerella mimuncinata, n. gen. and n. sp. See p. 74 for explanations of abbreviations and subscripts. eS maximum — dorsal valve hinge width width locality 31 CASG 66248.03 (paratype) 14.5 19 CASG 66248.04 (paratype) 19 21.5 CASG 66248.06 (paratype) 13 15 locality 32 CASG 66249.01 (paratype) 9.5 9.5 CASG 66249.02 (paratype) 11.5 13.5 locality 34 CASG 66250.01 (paratype) 15 17 CASG 66250.02 (holotype) 18.5 20 locality 35 CASG 66251.02 (paratype) 19 19 CASG 66251.03 (paratype) 17 20 locality 36 CASG 66252.01 (paratype) 20 21 interarea specimen length total length __ thickness height type 15 19.5 18.5 11.5 a.v. 15 20 18 12 a.v. 10 12.5 13. 10.5. a.v. 5.5 Te 6.5 6.0. a.v. 10.5 10.5 11.0, 10.0. a.v. 16 21 17 9 a.v. 12.5 DAES 17 12.5 a.v. 14 17 15.5 10 a.v 18.5 20. 19 12. d.v 15 18.5 16.5 13 a.v Se ee eee eee costate fold and sulcus, and from Vitimetula, n. gen., Dagyspirifer, n. gen., and Pseudospondylospira, n. gen., by its equidimensionally pyramidal plicate shell, and from all of these by its apparent lack of a functional pedicle. Discussion.— Assuming (with Dagys, 1974) that the nature of the microornament in cyrtinoid spiriferina- ceans is relatively important in distinguishing among the various genera, Phenacozugmayerella is probably most closely related to Yanospira Dagys, 1977, from the Norian of Okhotsk, U.S. S. R. Phenacozugmayerella mimuncinata, new species Plate 12, figures 6-24 Zugmayerella uncinata (Schafhautl). Stanley, 1979, p. 14 [as Z. in- cinata], pl. 8, figs. 10-12] (non Spirifer uncinata Schafhautl, 1851, p. 135, pl. 24, fig. 33). Etymology of name.—Gr. mimos = actor + uncin- ata. Description.—Shell small- to medium-sized, un- equally biconvex, with pronounced dorsal fold and ventral sulcus. Fold height about two to four, com- monly two, times the height of adjacent plicae. Flanks f both valves corrugated by radial plicae that expand plicae of less magnitude laterally. Fold and to subrounded in section; commonly iced, rarely interrupted mesially by one or ) faint radial plicae. Valve outline in ventral aspect juadrate, with straight lateral margins that converge slightly anteriorly, and mesially incurved anterior mar- zin. Outline in dorsal aspect transverse with rounded anterior margin in small specimens, becoming quad- rate in larger, apparently more mature individuals. Outer shell layer commonly not preserved; where pre- served, surface micro-ornament of radial capillae, composed of obliquely-disposed transverse bars formed by the intersection of capillae with growth lines, the two elements of microornament together presenting a finely cancellate surface. Three to five, most commonly three plicae on either side of fold on dorsal valve, one or more on ventral. Hinge width subequal to maximum shell width. Ventral valve deep, varying from about one-half to four-fifths of hinge width in height. Valve length and width, and shell thickness all subequal, but variable. Interarea high, triangular, with narrow to moderately broad triangular delthryium; interarea surface longi- tudinally ridged, ridges fine, extending dorsal to hinge as fine denticles; denticles fitting into articulatory re- cesses along hinge of opposite valve. Interarea flat, catacline to apsacline, angle to commissural plane mea- sured through shell 90° to about 110°, most commonly about 100°. Ventral apex of ventral beak commonly hooked posteriorly in last few mm. Dorsal valve comparatively shallow, with fold pro- viding curved quadrate section in lateral view. No in- terarea exposed, but short low beak extending about | mm beyond the hinge line in well-preserved speci- mens. Lateral plicae not straight, but gently curved and concave distally. Ventral interior with strong median septum extend- ing along valve floor about one-half valve length, rising anteriorly as a thin blade. Apparent dental plates join- ing inner margins of delthyrium to about mid-height of median septum, to form spondylium; median sep- tum continuing posterior to spondylium as thin blade not quite reaching plane of interarea surface. Muscle scars indistinct. Interior surface plicate, reflecting ex- terior macro-ornament. LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 91 Dorsal interior with minute notothyrium and hinge- line bearing row of anteroposteriorly elongate fossae that accommodate ventral valve hinge denticles. For measurements, see Table 10. Occurrence.—Phenacozugmayerella mimuncinata, Nn. sp. 1s apparently endemic to North America. In the present collections it has been recovered from rocks ranging in age from late Karnian to middle Norian [zones UCI through MNo2 of Tozer (1980)], at local- ities 6 [Alaska, Alexander terrane, zones UC1—UC3 of Tozer (1980), based on associated fauna], 32 [Nevada, Osobb Formation, Triassic cover on the Golconda Al- lochthon, zones LNol-LNo3 of Tozer (1980), based on stratigraphic position], 33 [Nevada, lower member of Luning Formation, Paradise terrane, zone LNol of Tozer (1980), based on fauna at this stratigraphic level], 34 [Nevada, upper member of Luning Formation, Par- adise terrane, zone LNo3 of Tozer (1980), based on stratigraphic position], 35 [Nevada, Luning Forma- tion, Paradise terrane, zone LNo3 of Tozer (1980), based on associated fauna], 36 [Nevada, Luning For- mation, Paradise terrane, zone LNol1 of Tozer (1980), Text-figure 5.—Posterodorsolateral views of portions of a dorsal valve of Phenacozugmayerella mimuncinata, n. gen. and n. sp., CASG 66250.02 (holotype, from loc. 36), showing (a) the five folds on one flank, <8; (b) the surface microornament of beaded capillae on the next- to-the-top fold, x 17; and (c) the capillate and capillate—cancellate (x) surface microornament on the top fold. 92 BULLETIN 337 based on fauna at this stratigraphic level], 37 [Nevada, lower Luning Formation, Paradise terrane, zones LNol-LNo2 of Tozer (1980), based on fauna at this stratigraphic level], 38 [Nevada, Luning Formation, Paradise terrane, zones LNol-MNo2 of Tozer (1980), based on stratigraphic position], 45 [Nevada, Jungo terrane, zone MNo2 of Tozer (1980), based on asso- ciated fauna], 48 [Oregon, Brisbois Formation, Izee terrane, zones UC1—UC3 of Tozer (1980), based on stratigraphic position], 49 [Oregon, Brisbois Forma- tion, Izee terrane, zones UC1I—UC3 of Tozer (1980), based on stratigraphic position], and 51 [Oregon, Bris- bois Formation, Izee terrane, zones UC1—UC3 of Toz- er (1980), based on stratigraphic position]. Diagnosis.—Small to medium-sized pauciplicate Phenacozugmayerella with acute-angled plications and distinct, comparatively large fold and sulcus. Types.—Holotype, CASG 66250.02; figured para- types, CASG 66248.03—CASG 66248.05, CASG 66249.01, CASG 66249.02, CASG 66250.01, CASG 66251.02, CASG 66252.01; measured paratypes, CASG 66248.03, CASG 66248.04, CASG 66248.06, CASG 66249.01, CASG 66249.02, CASG 66250.01, CASG 66251.02, CASG 66251.03, CASG 66252.01. Comparison. — Phenacozugmayerella mimuncinata, n. sp. is the only species of the genus, and is most easily confused with Zugmayerella uncinata (Schafhautl, 1851), from which it differs in its capillate-cancellate surface micrornament, and in lacking the duplex in- terarea that is characteristic of that species. APPENDIX LOCALITIES CITED IN THIS REPORT Locality information from specimen labels (not enclosed in square brackets) in many cases has been enhanced by comparison of original register entries, field labels, maps, notebooks, more recent topographic maps, and geologic maps and reports. Such additions appear in brackets. Zonal abbreviations refer to the zonation for the Triassic presented by Tozer (1980, Table II; see Table 1 herein) and have been assigned by N. J. Silberling (written commun., 1989). Terrane assignments were made by comparison of map position with those in the “Folio of the Lithotectonic Terrane Maps of the North American Cordillera” [U. S. Geological Survey, Miscellaneous Field Studies Maps, MF-1874-A (Jones et al., 1987), MF-1874-B (Monger and Berg, 1987), and MF-1874-C (Silberling et al., 1987)], and by consultation with N. J. Silberling (written commun., 1989). Locality numbers as used in this report (see also Text-fig. 1) are followed by the field number, in parentheses. Locality 1 (USGS loc. 2309).— California, [Shasta Co., Bollibokka Mtn. 15’ quad.;] E. line of sec. 30, T. 34 N., R. 1 W.; on both sides of Little Cow Creek [= Little Cedar Ck.], just below mouth of Bear Gulch; Collectors: [J.] Storrs and [C.] Washburne, August 8, 1901. [formation not given, probably Hosselkus Fm.; zones UCI-—LNo2 of Tozer (1980), based on stratigraphic position; Eastern Klamath terrane]. Locality 2 (USGS loc. 2314).—California, Shasta Co., [Burney 30’ quad.] near ctr., sec. 20, T. 34 N., R. 1 W.; 5 mi NE of Furnaceville; Collectors: [J.] Storrs and [C.] Washburne, August 8, 1901. [for- mation not given, probably Modin Fm.; zones MNol-UNo2 of Tozer (1980), based on associated fauna; Eastern Klamath terrane] Locality 3 (USGS loc. 2446).—California, Shasta Co., [?Millville 15’ quad.] on Toll Road by Cedar Creek, 10'2 mi [?] below [west of] Round Mtn.; Collectors: J. S. Diller and J. Storrs, June, 1892. [for- mation not given, probably Hosselkus Fm.; about zone UC2 of Tozer (1980), based on associated fauna; Eastern Klamath terrane] Locality 5 (USGS loc. 2769).— California, Shasta Co., [?Big Bend 15’ quad.] 5 mi. E. of Grizzly Peak, W. face of Table; bed of s. s. d cong.; Collector: J. Storrs, June 19, 1903. [Formation not given, lin Formation (based on lithology); zones LNol1-UNo3 of Toz- based on associated fauna; Eastern Klamath terrane] SGS loc. 10548).— Alaska, Southeast, Screen Islands, 3} 1:63,360 quad.;] Clarence Strait, west coast of Etolin 35’ of dark gray limestone 400’ thick; Collector: E. M. e, 1905. [Formation not given; zones UC1—UC3 of Tozer (1980), n associated fauna; Alexander terrane] ’ (USGS loc. 11406).—Alaska, Southeast, [Port Alex- ander D—1 1:63,360 quad.]; west side of Keku Straits, 7 mi NW of Pup Island, Kuiu Islands; Collector: A. F. Buddington, 1922. [Corn- wallis Limestone; zones UC1—LNol(?) of Tozer (1980), based on associated fauna (Muffler, 1967, p. C34); Alexander terrane] Locality 8 (USGS loc. 11814).—Nevada, [Mineral Co.,] Haw- thorne [1°] quad., T. 7 N., R. 35 E., N. 65° E. from Mina, first lime- stone exposure in Dunlap Gulch; Collectors: [H. G.] Ferguson, [S. H.] Cathcart, Heiges and [T. W.] Stanton, July 13, 1923. [Lower member of the Luning Formation, zone LNol of Tozer (1980), based on associated fauna (sponges); Paradise terrane] Locality 10 (USGS loc. 13354).—Nevada, [Mineral Co., Haw- thorne 1° quad.;] Gabb’s Valley Range, S. of Mayflower Ball Mill E. of Road, 100'—200' above top of blue limestone with small amount of brown sandy material; Collector: H. G. Ferguson, September 2, 1925. [Middle member of Gabbs Formation, zones UNo2-UNo3 of Tozer (1980), based on associated fauna; Paradise terrane] Locality 12 (USGS loc. 15550).—Oregon, [Wallowa Co.,] Imnaha River, [Harl Butte 15’ quad.;] NE, sec. 15, T. 2 S., R. 48 E., isolated outcrop over more or less 2 square mile area; Collectors: [R. W.] Richards and [B. N.] Moore, August 14, 1930. [formation not given, ??Martin Bridge Fm.; no information on stratigraphic position; Wal- lowa terrane] Locality 13 (USGS loc. 16266).—Alaska, South Central, Nutzotin Mtns., [Nabesna C-5 1:63,360 quad.;] Long. 143°10' W.; Lat. 62°35’ N.; head of tributary of Jack Creek that lies directly S. of Bear Ck. glacier, Copper River Region; Collector: F. H. Moffit, August 30, 1931. [formation not given; Wrangellia terrane] Locality 15 (USGS loc. 17424).—Idaho, [Nez Perce-Lewis Co. line, Culdesac 15’ quad.;] about 6 mi up Mission Ck. from Jaques R. R. Station, past St. Joseph’s Mission and Slickpoo Settlement, at first narrows of Mission Creek, collection from black limestone abt. 70' above water, E. side of ck.; Collectors: J. Reed and J. S. Williams, June 17, 1936. [formation not assigned, zones UNol-UN0o3 of Tozer (1980), based on associated fauna [the brachiopods (Cooper, 1942) and gastropods (Haas, 1953) suggest a Norian—Rhaetian age. Stanley (1979) notes that many of the coral species are not known elsewhere LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 93 in North America, but have been reported from the Zlambach beds (Norian) of Fischerweise, Austria. He states (p. 27) that he has “‘cho- sen tentatively to regard the Lewiston locality as Upper Norian,” but that “no diagnostic ammonites have been discovered, and the age of the known fauna is equivocal.”; Wallowa terrane] Locality 16 (USGS loc. 24387).—Peru, Atacocha Dist.; Dept. of Pasco; Machican Mine area; Collectors: R. F. Johnson and R. W. Lewis, 1952. [formation not given, = Chambara Fm. of Pucara Gp.; zones UC1-—UNo3 of Tozer (1980), based on associated fauna; ter- rane unknown] Locality 18 (USGS loc. M96).—Nevada, Pershing Co., [Buffalo Mtn. 15’ quad.;] NE", sec. 5, T. 26 N., R. 34 E, and sec. 32, T. 27 N., R. 34 E.; along strike of beds at base of limestone cliffs, approx. 74 miS. to 2 mi W. of Nevada Quicksilver [= Juniper] mine; Collectors: N. J. Silberling, R. E. Wallace, W. P. Irwin, 1956. [formation not given, but = Antelope Springs Fm. of Oldow, Bartel, and Gelber (1990); zone LNo3 of Tozer (1980), based on associated fauna; Jungo terrane] Locality 20 (USGS loc. M1708).—Alaska, Wrangell Mtns., Mc- Carthy B—5 1:63,360 quad.; on west side of East Fork Valley just E. of largest patch of Cretaceous capping Green Butte Ridge; Collector: N. J. Silberling, July 7, 1962. Transitional beds between Chitistone and Nizina formations. [zone LNol of Tozer (1980), based on as- sociated fauna; Wrangellia terrane] Locality 21 (USGS loc. M1749).—Oregon, [Wallowa Co.,] Wal- lowa Mtns., Enterprise 15’ quad.; ctr. of W'2 of boundary line bet. sec. 3, T. 3 S.,R. 44 E., and sec. 34, T. 2 S., R. 44 E.; elev. ca. 6240’ on W. side of Hurricane Ck., about 1 mi N. of Little Granite Ck.; Collector: B. Nolf [Oregon State Univ.], 1962. Martin Bridge For- mation. [zone UC3 of Tozer (1980), based on associated fauna; Wallowa terrane] Locality 22 (USGS loc. M1906).— Alaska, Southeast, Keku Strait, Port Alexander [D-1 1:63,360] quad.; NE side of Cornwallis Pen- insula ca. 1 mi E. of Point 5600’ S. 85° E. from triangulation station Corn; silicified fauna in medium to thick-bedded limestone; Collec- tor: L. J. P. Muffler, June, 1963. Cornwallis Limestone. [Silberling, written commun., 1963 (cited by Muffler, 1967, p. C30, table 2) dates this locality as of “‘uncertain age in the Late Triassic,” based on “‘scleractinian corals and (or) the spiriferid brachiopod Spondy- lospira; arcestid or clydoniacid ammonites.” He later (written com- mun., 1989) assigned it to zones UC1—UNo of Tozer (1980), based on associated fauna; Alexander terrane] Locality 23 (USGS loc. M1911).— Alaska, Southeast, [Keku Strait, Port Alexandria D-1 1:63,360 quad.]; S. side of small cove of E. shore, Kuiu Island, 1.5 mi SSE, or 8300’ S. 19° E. from triangulation station ‘“‘Low”; silicified fauna from 185’ above base of limestone; Collector: N. J. Silberling, June, 1963. Cornwallis Limestone. [Sil- berling, written commun., 1963 (cited by Muffler, 1967, p. C30, table 2) dated this locality as “latest Karnian or earliest Norian,” based on the occurrence of species of Halobia and Mojsisovicsites. He later (written commun., 1989) assigned it to zone LNo1 of Tozer (1980), based on associated fauna; Alexander terrane] Locality 24 (USGS loc. M1917).—Alaska, Southeast, Keku Strait; Port Alexander D-1 1:63,360 quad.; NE shore of Kuiu Island ca. 3% mi SE of Pt. Cornwallis; 17,950’ S. 69° E. from triangulation station CORN; impure limestone 20-30’ above base of limestone; Collectors: N. J. Silberling, L. J. P. Muffler, June 27, 1963. [Corn- wallis Limestone; Silberling (written commun., 1963, cited by Muf- fler, 1967, p. C28, table 1) dated this locality as of “‘uncertain age within the Late Triassic,” based on “‘scleractinian corals and (or) the spiriferid brachiopod Spondylospira, [and] arcestid or clydonitacid ammonites’. He later (written commun., 1989) assigned it to zones UCI-LNo2 of Tozer (1980), based on stratigraphic position; Al- exander terrane] Locality 25 (USGS loc. M1919).— Alaska, Southeast, Keku Strait, Port Alexander D-1 1:63,360 quad.; E. shore Kuiu Island 0.5 mi SSE, or 2500’ S. 23° E. from triangulation station LOW; impure limestone interstratified with mafic tuff, but below main body of Tmiassic volcanics; Collector: N. J. Silberling, June 29, 1963. Hound Island Volcanics. [Silberling (written commun., 1963, cited by Muf- fler, 1967, p. C30, table 2) dated this locality as “latest Karnian or earliest Norian,”’ based on species of Halobia or Mojsisovicsites. He later (written commun., 1989) assigned it to zones UC2-LNol of Tozer (1980), based on associated fauna; Alexander terrane] Locality 26 (USGS loc. M2135).— Alaska, Southeast, Keku Strait, Port Alexander D-1 1:63,360 quad.; easternmost tip of islet in NW Keku Islets, 4.41 mi S. 78.5° E. from triangulation station CORN; dense thick-bedded limestone at tidal level; Collector: L. J. P. Muf- fler, 1963. Keku Volcanics; Silberling (written commun., 1963, cited by Muffler, 1967, p. C28, table 1) dated this locality as of “uncertain age within the Late Triassic,” based on scleractinian corals and (or) the spiriferid brachiopod Spondylospira, and arcestid or clydonitacid ammonites. He later (written commun., 1989) assigned it to zones LCI-—UN0o2 of Tozer (1980), based on associated fauna; Alexander terrane] Locality 27 (USGS loc. M2136).— Alaska, Southeast, Keku Strait, Port Alexander D—-1 1:63,360 quad.; on S. side of islet in SE Keku Straits, | mi SE of triangulation station THUM, 1.03 mi N. 34° E. from triangulation station LOW; Collector: [L. J. P.] Muffler, 1963. Cornwallis Limestone. [Silberling (written commun., 1963, cited by Muffler, 1967, p. C30, table 2) dated this locality as of “uncertain age in the Late Tnassic,” based on “‘scleractinian corals and (or) the Spiriferid brachiopod Spondylospira; [and] arcestid or clydonitacid ammonites.” He later (written commun., 1989) assigned it to zones UCI1-UNo of Tozer (1980), based on stratigraphic position; Alex- ander terrane] Locality 28 (USGS loc. M2672).— Oregon, [Wallowa Co.,] Home- stead 1:62,500 quad.; Oregon side of Snake River Canyon, elev. 2970’ on N. side McGraw Ck., abt. 0.85 mi NW from its mouth, abt. 200’ about base of limestone unit; Collector: T. L. Vallier, June 25, 1964. [Martin Bridge Fm.; Silberling (cited in Vallier, 1967, pp. 246-247) assigned this locality to zone LNo1 of Tozer (1980), based on the occurrence of the ammonoid Tropiceltites cf. T. columbianus; Wallowa terrane] Locality 29 (GSC loc. 10229).—Canada, Yukon Territory, Lake Laberge area ca. 61° N. lat.; 135° W. long.]; High ridge of banded appearance 4 mi NE of Braeburn road house; from interbedded coral reef; Collector: E. J. Lees, 1930. [probably Formation “F”, Lewes River Gp.; see note under loc. 31, below; Stikine terrane] Locality 30 (GSC loc. 23418).—Canada, Yukon Territory, Lake Laberge area. East side of Lake Laberge (Loc. 11 of G. S.C. Bull. 43) [61°15'25” N. lat.; 135°12'9” W. long.]; Formation “F’’, Lewes River group; Collector: E. T. Tozer, 1953. [See note under loc. 31, below; Stikine terrane] Locality 31 (GSC loc. 23462).—Canada, Yukon Territory, Lake Laberge area; East side of Lake Laberge (Loc. 10 of G. S.C. Bull. 43) [61°14'55” N. lat., 135°11'26” N. long.]; Formation ““F”, Lewes River group; Collector: E. T. Tozer, 1953. [Tozer (1958, p. 19) as- signs this and other localities in Formation “‘F” of the Lewes River group in the Lake Laberge area to “late Norian (mid-Upper Triassic) [sic]” and cites the occurrence there of several species of pelecypods in association with Spondylospira lewesensis (Lees, 1934). This age call is apparently based on general similarity of the fauna to an assemblage “widely distributed in southern Yukon and in British Columbia west of the Rocky Mountain Trench.” He specifically cites similarities to the fauna of the Tyaughton group of southern British Columbia, which occurs on the west coast of Vancouver Island in “beds overlying shales with Monotis subcircularis.” Silberling (writ- ten commun., 1989) assigns this locality to zone UNo2 of Tozer (1980), based on associated fauna; Stikine terrane] Locality 32 (LSJU loc. 720-C).—Nevada, Lander Co., [Cain Mountain 15’ quad.; Augusta Mountains] “W. slope of S. end of 94 BULLETIN 337 Lone Peak [Cane or Boundary Peak of 40th Parallel Survey], immed. beyond the divide of the first deep canyon S. of Jenkins Ranch; beds lie conformably on Carboniferous? limestone [= Upper Triassic Cane Spring Formation]; fossils from top of 500’ thick zone, beginning 60’ above ?Carb. [Upper Triassic] Ist.’’; Collector: S. W. Muller, September, 1928. [Osobb Formation; zones LNol-LNo3 of Tozer (1980), based on stratigraphic position; Triassic cover on the Gol- conda Allochthon] Locality 33 (LSJU loc. 724).—Nevada, Mineral Co., Hawthorne [1°] quad., Pilot Mtns.; Dunlap Canyon, 3 mi up from the mouth of the canyon on the W. side of the canyon; Collector: S. Muller, April, 1928; September, 1934; September, 1935. [lower member of Luning Fm.; zone LNol of Tozer (1980), based on fauna at this stratigraphic level; Paradise terrane] Locality 34 (LSJU loc. 764).—Nevada, Mineral Co., Hawthorne [1°] quad.; Pilot Mtns.; E. of Mina, abt. 1 mi S. of the mouth of Dunlap Canyon, abt. 1750’ due S. of a rhyolite knob; impure lime- stones faulted against slates; Collector: S. Muller, May, 1932. Upper member of Luning Formation. [zone LNo3 of Tozer (1980), based on stratigraphic position; Paradise terrane] Locality 35 (LSJU loc. 800-B).— Nevada, Nye Co., [Ione 15’ quad., T. 12 N., R. 39 E. (unsurveyed)]; about 1 mi from the mouth of Union Canyon, E. side; Collector: S. Muller, September, 1930. Lun- ing Formation. [zone LNo3 of Tozer (1980), based on associated fauna; Paradise terrane] Locality 36 (LSJU loc. 844).—Nevada, Mineral Co., Hawthorne and Tonopah [1°] quads.; Pilot Mtns., Dunlap Canyon; lower lime- stone near coral bed; Collector: S. Muller, September 12, 1934. Lun- ing Formation. [zone LNo1 of Tozer (1980), based on fauna at this stratigraphic level; Paradise terrane] Locality 37 (LSJU loc. 872).—Nevada, Mineral Co., Hawthorne and Tonopah [1°] quads.; Pilot Mtns., between Cinnabar and Dunlap Canyons at the boundary of the two quads.; low hills between the two canyons N. of the prominent, andesite, rhyolitic hill; Collector: S. Muller, September, 1934. [lower Luning Formation; zones LNol- LNo2 of Tozer (1980), based on fauna at this stratigraphic level; Paradise terrane] Locality 38 (LSJU loc. 1014).—Nevada, Mineral Co., Tonopah [1°] quad., Pilot Mtns.; Cinnabar Canyon, near mouth of first trib- utary from the east, above the “Slate Gorge”’; Collector: S. Muller, 1936. Luning Formation. [zones LNol—-MNo2 of Tozer (1980), based on stratigraphic position; Paradise terrane] Locality 39 (LSJU loc. 1377).—Nevada, Pershing Co., “Lovelock [1°] quad. [Buffalo Mountain 15’ quad.]; abt. 16 mi E. of Lovelock, abt. | mi SW of Nevada Quicksilver Mine; brownish thinly lami- nated shales and gray limestones”; Collectors: Muller, Bush, John- ston, [E. T.] Schenk and [H. E.] Wheeler, June-July, 1930. [same as loc. 18, formation not given, but = uppermost Hollywood Fm. of Oldow, Bartel, and Gelber (1990); zone LNo3 of Tozer (1980), based on associated fauna; Jungo terrane] Locality 40 (LSJU loc. 1517).—Nevada, Mineral Co., Hawthorne and Tonopah [1°] quads.; Pilot Mtns., Cinnabar Gulch, abt. 1'2 mi above the Slate and Cg. Gorge, last exposure of Upper Triassic | stone before getting back into Slate and Congl.; Collector: S. W. 934. Luning Formation. [zone LNo3 of Tozer (1980), based ic position; Paradise terrane] LSJU loc. 2336-A).—Nevada, Lander Co., Sonoma juad. [Cain Mountain 15’ quad.]; Augusta Mtns., 2!2 mi Jenkins Ranch, T. 25 N., R. 39 E., 400-500’ above base of unit; Collector: S. W. Muller, date unknown. [Osobb Formation. zones LNo2-LNo3 of Tozer (1980), based on stratigraphic position; Tniassic cover on the Golconda Allochthon] Locality 42 (LSJU loc. 2348).—Nevada, Pershing Co., Sonoma Range [1°] quad. [Rose Creek 15’ quad.]; East Range, N. end, at the head of Rose Ck., elev. 100’ above base of limestone; Collector: S. W. Muller, August, 1940. [Dun Glen Formation; zone LNo3 of Tozer (1980), based on associated fauna; Triassic cover on the Gol- conda Allochthon] Locality 43 (LSJU loc. 2546).—Nevada, Pershing Co., Sonoma Range [1°] quad. [Cain Mountain 15’ quad.]; Augusta Mtns., 2'2 mi SE of Jenkins Ranch, abt. 400-500’ above base of the unit; Collector: S. W. Muller, date unknown. ?Winnemucca Formation [on strike with loc. 41; zones LNo2—LNo3 of Tozer (1980), based on strati- graphic position; Triassic cover on the Golconda Allochthon] Locality 44 (LSJU loc. 2970).—California, Shasta Co., Big Bend 15’ quad.; [NEY4sNE“%SW'4, sec. 25, T. 38 N., R. 1 W.; Alder Creek Trail, abt. 2 mi N. of confluence with Devil’s Canyon] Collector: A. F. Sanborn, 1950. Modin Formation, Devil’s Canyon Member. [zones UNol-UNo3 of Tozer (1980), based on associated fauna; Eastern Klamath terrane] Locality 45 (LSJU loc. 3148).— Nevada, Churchill Co., [Shoshone Meadows 15’ quad.;] N. end of Clan Alpine Range, Shoshone Ck., Ys mi N. of the road at a point 1.8 mi down the creek from Shoshone Springs; coral bed abt. 200’ below the limestone ledge; Collectors: [E.] Blackwelder, Sahni, [N. J.] Silberling, August 8, 1952. [formation not given; zone MNo2?2 of Tozer (1980), based on associated fauna; Jungo terrane] Locality 47 (Field No. HC341).— Oregon, Wallowa Co., Jim Creek Butte [15’] quad.; NE cor., sec. | [unsurveyed], T. 5 N., R. 47 E.; 45°56'38" N., 116°52'55” W.; crest of ridge on N. side of Coon Creek; limestone lens in Seven Devils volcanics; Collector: G. C. Simmons, 1974. [Martin Bridge Formation; zones LNol—-MNo2? of Tozer (1980), based on associated fauna; Wallowa terrane] Locality 48 (Field No. 56-133).—Oregon, [Crook Co.,] Suplee area, NENE', sec. 2, T. 18 S., R. 25 E.; isolated limestone knob 750' WNW of road cut 2700’ N. of Camp Ck. crossing; limestone con- glomerate stratigraphically below point estimated 1800’ above base of Brisbois Formation; Collectors: S. W. Muller, [W. R.] Dickinson, {L. H.] Vigrass, 1956. Brisbois Formation. [zones UC1—UC3 of Toz- er (1980), based on stratigraphic position; Izee terrane] Locality 49 (Field No. 56-135).—Oregon, [Crook Co.,] Suplee area; NW cor., NE", sec. 1, T. 18 S., R. 25 E.; limestone outcrop 1000’ E. of Camp Creek on summit of short arcuate E/W ridge; prob. same horizon as 56-133 [loc. 48]; Collectors: [S. W.] Muller, [W. R.] Dick- inson, [L. W.] Vigrass, 1956. Brisbois Formation. [zones UC1-UC3 of Tozer (1980), based on stratigraphic position; Izee terrane] Locality 51 (Field No. 57-417).—Oregon, [Crook Co.,] Suplee area; W boundary, SE’4sNE',, sec. 23, T. 17 S., R. 25 E., 100’ SE of ““Pau- lina telephone” located at junction of Paulina—Suplee road with road to A. Bernard Ranch house; about 700’ above base of unit; Collector: L. W. Vigrass, 1957. Brisbois Formation. [zones UC1-UC3 of Tozer (1980), based on stratigraphic position; Izee terrane] Locality 53 (Field No. 77-S-111).—Alaska, South Central, Alaska Range, Upper Chulitna district, Healy (A-6) quad.; limestone and pillow basalt unit on crest of secondary spur on SW side of valley of the W. fork of the Chulitna River, 3.80 km S. 89° E. from VABM Joyce (6363); Collector: N. J. Silberling, 1977. [zone UC3 of Tozer (1980), based on associated fauna; Chulitna terrane] REFERENCES CITED Ager, D. V. 1964. The supposedly ubiquitous Tethyan brachiopod Halorella and its relations. Palaeontological Society of India, Jour- nal, vols. 5—9 (1960-1964), pp. 54-70. Babaie, H. A., and Speed, R. C. 1990. Origin of kink bands in the Golconda Allochthon, Toiyabe Range, Nevada. Geological Society of America, Bulletin, vol. 102, pp. 315-321. LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER 95 Berg, H. C., Jones, D. L., and Richter, D. H. 1972. Gravina—Nutzotin belt — Tectonic significance of an upper Mesozoic sedimentary and volcanic sequence in southern and southeastern Alaska. United States Geological Survey Professional Paper 800-D, 24 pp. Bittner, A. 1884. Aus den Salzburger Kalkhochgebirgen. Zur Stellung der ' Halstdtter Kalke. Kaiserlich-Koniglichen Reichsanstalt, Verhandlungen, No. 6, pp. 99-113. 1892. Brachiopoden der Alpinen Trias. Nachtrag der Kaiserlich- en-K6niglichen Geologischen Reichsanstalt, Jahrbuch, vol. 17, No. 2, 40 pp., 4 pls. 1902. Brachiopoden und Lamellibranchiaten aus der Trias von Bosnien, Dalmatien und Venetien. Kaiserlichen—K6nig- lichen Geologischen Reichsanstalt, Jahrbuch, Jahrgang 1902, Band 52, Heft 3, 4, pp. 495-642, pls. 18-27. Brueckner, H. W., and Snyder, W. S. 1985. Structure of the Havallah sequence, Golconda allochthon, Nevada: evidence for prolonged evolution in an accretionary prism. Geological Society of America, Bulletin, vol. 96, pp. 1113-1130. Brueckner, H. K., Snyder, W. S., and Boudreau, M. 1987. Diagenetic controls on the structural evolution of siliceous sediments in the Golconda allochthon, Nevada, U.S. A. Journal of Structural Geology, vol. 9, pp. 403-417. Burke, D. B., and Silberling, N. J. 1973. The Auld Lang Syne Group of Late Triassic and (?) Jurassic age, north-central Nevada. U.S. Geological Survey, Bul- letin 1394-E, 14 pp. Bychkov, Yu. M., and Dagys, A. S. 1984. /The Late Triassic fauna of the Koryakian mountain sys- tem and its importance for paleogeographic and paleotec- tonic reconstruction.], pp. 8-18 in Stratigrafiya, Fauna i Flora Triasa Sibiri [Dagys, A. S., ed.]. Akademiya Nauk S.S.S.R., Sibirskoye Otdelenie, Institut Geologii 1 Geo- fiziki, Trudy 600, 80 pp., 16 pls. Bychkov, Yu. M., Dagys, A. S., Efimova, A. F., and Polubotko, I. V. [eds.] 1976. Atlas Triasovoi Fauny i Flory Severo-vostoka SSSR. Min- isterstvo Geologii R. S. F.S. R, Severo-vostoknoye Or- dena Trudovogo Krasnogo Zhnameni Territorialnoye Geologicheskoye Upravlaniye. Moscow. Campbell, H. 1990. Permian, Triassic, and Jurassic brachiopod faunas in Southland and south Otago, New Zealand. Guidebook for Field Excursion NZ4, Second International Brachiopod Congress, Dunedin, New Zealand (Geological Society of New Zealand Miscellaneous Publication 46A), 53 pp. Ching Y., and Feng B. 1977. Upper Triassic brachiopod fauna from the area in the east of the Hengduan mountains, western Yunnan. Mesozoic fossils from Yunnan, China, fasc. 2, pp. 39-69, 6 pls. Chong D., G., and Hillebrandt, A. 1985. El Triasico preandino de Chile entre los 23°30’ y 26°00’ lat. sur. 1V Congréso Geologico Chileno (Universidad del Norte, Antofagasta: August, 1985), Proceedings, vol. 1, pp. 162-210. Compton, R. R. 1960. Contact metamorphism in Santa Rosa Range, Nevada. Geological Society of America, Bulletin, vol. 71, pp. 1383- 1416. Cooper, G. A. 1942. New genera of North American brachiopods. Washington Academy of Sciences, Journal, vol. 32, No. 8, pp. 228- 235. 1944. Phylum Brachiopoda, pp. 277-365 in Shimer, H. W., and Shrock, R. R. [eds.], Index Fossils of North America. Tech- nology Press, Massachusetts Institute of Technology. Cordey, F., Mortimer, N., DeWever, P., and Monger, J. W. H. 1987. Significance of Jurassic radiolarians from the Cache Creek terrane, British Columbia. Geology, vol. 15, pp. 1151- 1154. Dagys, A.S. 1962. [New Late Triassic Spiriferinacea from the northwestern Caucasus}. Paleontologicheskii Zhurnal, vol. for 1962, No. 2, pp. 47-56 [in Russian]. 1963. Verkhnetriasovye Brakhiopody Yuga S. S. S. R. Akade- miya Nauk S. S. S. R, Sibirskoye Otdelenie. 248 pp. 1965. Triasovye Brakhiopody Sibiri. Akademiya Nauk S. S. S. R, Sibirskoye Otdelenie, Instituta Geologii i Geofiziki, 186 pp. 1974. Triasovye Brakhiopody (Morfologiya, Sistema, Filogeniya, Stratigraficheskoe Znachenie i Biogeografiya). Akademiya Nauk S.S. S. R, Sibirskoye Otdelenie. Instituta Geologii i Geofiziki Trudy, No. 214, 386 pp. 1977. [New Triassic brachiopods from the north-east of U. S. S.R.), pp. 5-22, pl. 1 in Stratigrafiya i fauna borealnogo Triasa. Akademiya Nauk S. S. S. R., Sibirskoye Otdelenie, Insti- tuta Geologii i Geofiziki, Trudy No. 344. Davidson, T. 1851-1886. .4 Monograph of the British fossil Brachiopoda. Pa- laeontographical Society Monographs, 6 vols. Debiche, M., Cox, A., and Engebretson, D. C. 1987. The motion of allochthonous terranes across the North Pa- cific basin. Geological Society of America, Special Paper 207, 49 pp. Dickinson, W. R., and Vigrass, L. W. 1965. Geology of the Suplee-Izee area, Crook and Harney coun- ties, Oregon. Oregon, State Department of Geology and Mineral Industries, Bulletin 58, viii + 109 pp., 3 pls., 30 figs., 12 tables. Dover, J. H. 1990. Problems of terrane terminology — Causes and effects. Geology, vol. 18, No. 6, pp. 487-488. Ehman, K. D. 1985. Paleozoic stratigraphy and tectonics of the Bull Run Moun- tains, Elko County, northern Nevada. unpublished Ph. D. dissertation, University of California, Davis. Frizzell, D. L. 1933. Terminology of types. American Midland Naturalist, vol. 14, No. 6, pp. 637-668. Gabb, W. M. 1864. Description of the Triassic fossils of California and adjacent territories. Geological Survey of California, Paleontology, vol. 1, No. 2, pp. 19-35. Gardner, M. C., Bergman, S. C., Cushing, G. W., MacKevett, E. M., Jr., Plafker, G., Campbell, R. B., Dodds, C. J., McClelland, W. C., and Mueller, P. A. 1988. Pennsylvanian pluton stitching of Wrangellia and the Al- exander terrane, Wrangell Mountains, Alaska. Geology, vol. 16, No. 11, pp. 967-971, 6 figs., 2 tables. Guex, J., and Taylor, D. G. 1976. La limite Hettangien—Sinémurien, des Préalpes romandes au Nevada. Eclogae Geologicae Helvetiae, vol. 69, pp. 521-526. Haas, O. 1953. Mesozoic invertebrate faunas of Peru. American Museum of Natural History, Bulletin, vol. 101, 328 pp. Hall, J., and Whitfield, R. P. 1877. Paleontology, part II, vol. 4, in Geological Exploration of the Fortieth Parallel. United States Army Engineering De- partment Professional Paper No. 18, 650 pp. 96 BULLETIN 337 Hallam, A. 1981. The end-Triassic bivalve extinction event. Palaeogeogra- phy, Palaeoclimatology, Palaeoecology, vol. 35, pp. 1-44. Hector, J. 1879. On the fossil Brachiopoda of New Zealand. New Zealand Institute, Transactions and Proceedings, vol. 11, pp. 537- 539. Hillhouse, J. W. 1977. Paleomagnetism of the Triassic Nikolai greenstone, Mc- Carthy quadrangle, Alaska. Canadian Journal of Earth Science, vol. 14, pp. 2578-2592. Hillhouse, J. W., and Gromme, C. S. 1980. Paleomagnetism of the Triassic Hound Island volcanics, Alexander terrane, southeastern Alaska. Journal of Geo- physical Research, vol. 85, pp. 2594-2602. 1984. Northward displacement and accretions of Wrangellia: new paleomagnetic evidence from Alaska. Journal of Geo- physical Research, vol. 89, pp. 4461-4477. Hoover, P. R. 1983. The Cooperculum: a new structure in the Phylum Brach- iopoda, and its functional significance. Journal of Pale- ontology, vol. 57, No. 5, pp. 1017-1029. 1990. Occurrence of Spondylospira in the marine lower Jurassic of Central Peru, and its biostratigraphic significance. Ab- stracts of the Second International Brachiopod Congress, Dunedin, New Zealand, February, 1990. [Abstract] p. 48. 1991. Occurrence of Spondylospira in the marine Early Jurassic of central Peru, and its biostratigraphic significance, pp. 393-396, incl. 5 figs., in Brachiopods through time (MacKinnon, D. I., Lee, D. E., and Campbell, J. D., eds.], Proceedings of the Second International Brachiopod Con- gress, Dunedin, New Zealand, 5—9 February, 1990. A. A. Balkema, Rotterdam, xii + 447 pp. House, M. R., and Senior, J. R. [eds.] 1980. The Ammonoidea. Systematics Association Special Vol- ume 18. Academic Press, London and New York. Irving, E., and Monger, J. W. H. 1987. Preliminary paleomagnetic results from the Permian As- itka group, British Columbia. Canadian Journal of Earth Science, vol. 24, pp. 1490-1497. Jones, D. L., Silberling, N. J., Coney, P. J., and Plafker, G. 1987. Lithotectonic terrane map of Alaska (west of the 141st me- ridian. U. S. Geological Survey, Miscellanous Field Stud- ies Map MF-1874-A. Jones, D. L., Silberling, N. J., Csejtey, B., Jr., Nelson, W. H., and Blome, C. D. 1980. Age and structural significance of ophiolite and adjoining rocks in the upper Chulitna district, south-central Alaska. U.S. Geological Survey, Professional Paper 1121-A, 21 pp. Jones, D. L., Silberling, N. J., and Hillhouse, J. 1977. Wrangellia — a displaced terrane in northwestern North 4merica. Canadian Journal of Earth Science, vol. 14, pp. 2565-2577. oe ctatersreiel = ehey lay stois © evs) sre ievev lst w= «ee s/o) sevete) yee steer Holotype, partial ventral valve, (4) ventral, (5) dorsal, and (6) lateral views, x 1, and (7) dorsolateral view, x 2, showing low beak, wide delthyrium, shallow valve profile, and lack of spondylospirid spondylium; USNM 12671; collected 1.5 mi south of Dun Glen Pass, Pah-Ute Range, Nevada, by A. Hague; figured for comparison with PI. 9, figs. 36, 37 [Spondylospira alia (Hall and Whitfield, 1877) of Cooper (in Shimer and Shrock, 1944) = Spondylospira lewesensis (Lees, 1934)]. WE SPONGY lOSpiraparmata TIGOVET MOSS) se cvs crs ceye MNCL oe heer 505 SHED AH. eS SS NEP AE, Sle SAPS NER Site SEE Seis a settistcfeuecetesa elaushateieassea 13 Holotype, articulated valves, (8) dorsal, (9) ventral, (10) anterior, and (11) lateral views, x 1, showing form, outline, fine ornament and subdued dorsal fold; (12) posterior and (13) posterolateral views, x 1 (stereo), showing denticulate hinge, high ridged interarea, narrow delthyrium, and fragmentary tripartite cooperculum of one basal and two lateral plates; CASG 60975.01, from loc. 44. Pseudospondylospira perplexa new genus and New 'SpeCieS: . 2... . ce ee ee ee eee Sure eee ceesenyn eiete weiey eis eee sie ese ene here 14-18. Holotype, articulated valves, (14) anterior, (15) posterior, (16) dorsal, (17) ventral, and (18) lateral views, x 1, showing globose profile, narrow interarea, and hooked ventral apex; USNM 450278, from loc. 22. 19, 20. Paratype, dorsal valve, (19) ventral (interior) and (20) dorsal views, 1.5, showing circular outline, narrow denticulate hinge, and surface of a strongly ornamented variant; USNM 450279, from loc. 22. 21. Paratype, dorsal valve, dorsal view, x 1.5, showing surface of a weakly ornamented variant, USNM 450280, from loc. DD: 22. Paratype, partial articulated valves, anteroventrolateral view, x2, in ventral valve showing anterior protrusion of dorsal portion of median septum, and spondylospirid spondylium; in dorsal valve showing hinge sockets, crural bases, and descending branch attachment ridges on valve floor; USNM 450281, from loc. 22. 23, 24. Paratype, partial ventral valve, (23) posteroventrolateral (interior) view, x 2, and (24) anterodorsolateral (interior) view, x 3, showing exterior and interior configurations of paired apical pedicle foramena; USNM 450282, from loc. 22. pe Pseudosponaylospira alt. P. perplexa> New Senus and NEw, SPECIES) pieyeyevsyev-are atest ie ate hete eatee etsterer eae 2 ee ected ovetere aia) stetarat ata 25, 26. Figured specimen, dorsal valve, (25) dorsal and (26) ventral (interior) views, <1, showing straight, angular costae and short denticulate hinge of a large specimen; USNM 450283, from loc. 28. 27. Figured specimen, partial ventral valve, anterolateral (interior) view, x 2, showing denticulate hinge and spondylospirid spondylium of a smaller specimen; USNM 450284, from loc. 28. 101 81 83 79 80 Figure BULLETIN 337 EXPLANATION OF PLATE 11 1215 Spondylospira tricosta\ newispecies) 4. --.aa:2 = 202s ee 2" 3-6. 14, 15. Figured specimen, articulated valves, (1) dorsal and (2) posterior views, x 1, showing size and form of a small individual; CASG 66256.01, from loc. 42. Figured specimens, articulated valves, (3, 5) dorsal and (4, 6) lateral views, x 1, showing size, form, and characteristic trifurcate costa on dorsal folds of two specimens, one well-preserved, the other largely decorticated; (3, 4) CASG 66255.01, (5, 6) CASG 66255.02, both from loc. 41. . Holotype, partially decorticated articulated valves, (7) dorsal view, <1, for scale; (8) dorsal, (9) ventral, (10) posterior, (11) lateral, and (12) anterior views, = 1.5, showing form, outline, ridged interarea, spondylospirid spondylium, and characteristic costal trifurcation on dorsal fold; (13) anterior view, * 3, showing finely cancellate surface microornament and contrast between shell and internal mold surfaces; CASG 66247.01, from loc. 32. Figured specimen, dorsal valve, (14) ventral (interior) and (15) dorsal views, x 2, showing denticulate hingeline, attachment ridges for descending branches of spire, and characteristic costal trifurcation on dorsal fold; USNM 450289, from loc. 28. 16-32. Vitimetula parva, new genus and NEW SPECIES... 6 ee jes on en cere ta niche ae sie esis = a cls oe a 16-21. 26-29. 30. 3532. Holotype, articulated valves, (16) anterior view, x 1, for scale; (17) anterior, (18) posterior, (19) dorsal, (20) ventral, and (21) lateral views, <3, in narrow, deep, faintly costate ventral valve showing high, narrow, ridged interarea; in low, rounded caplike dorsal valve showing absence of radial ornament, USNM 450290, from loc. 15. . Paratype, ventral valve, (22) ventral and (23) posterior views, <3, showing rough, irregularly ridged, laterally-rimmed interarea with very narrow delthyrium and multiple apical pedicle foramena; USNM 450293, from loc. 15. . Paratype, dorsal valve, (24) ventral (interior) and (25) dorsal views, x 3, showing weak radial surface ornament, denticulate hingeline, and anteriorly jugate descending branches of spire joined to valve floor by meshwork; USNM 450296, from loc. 15. Paratype, ventral valve, (26) anterior, (27) dorsal (interior), (28) lateral, and (29) posterior views, <3, showing narrow, non-costate sulcus and weakly costate flanks, high, ridged, laterally-rimmed interarea, with narrow delthyrium, spon- dylospirid spondylium and denticulate hingeline; USNM 450291, from loc. 15. Paratype, dorsal valve, ventral view, <3, showing denticulate hingeline, descending branches of spire, and supportive meshwork; USNM 450298, from loc. 15. Paratype, dorsal valve, (31) dorsal view, = 3, showing valve outline and absence of radial ornament; (32) ventral (interior) view, <3 (stereo), showing bowl-like valve form and curvature of descending branches of spire, USNM 450297, from loc. 15. [mesial ridge is silicified crack filling]. 33-50: Zugmayerella americana, NEW SPECIES! <0). =< 2 <2 oe oie oie cosine ering aah ha Sica hss ae Ache ea 38-42. 43-50. 3: (34, 37) from loc. 20; (35, 36) from loc. 28. Paratype, ventral valve, (38) lateral, (39) dorsal (interior), (40) ventral, (41) posterior, and (42) anterior views, x 2, showing triangular profile and outline, weakly paucicostate flanks, flat ridged interarea, broad, flat, non-costate mesial sulcus, spondylospirid spondylium and denticulate hingeline; USNM 450301 [same as Pl. 11, fig. 37], from loc. 20. Holotype, articulated valves, (43) anterior view, 1, for scale; (44) anterior, (45) posterior, and (46) lateral views, x3, showing form, outline, and narrower ventral sulcus of small individual: ventral valve, (47) dorsal (interior) and (48) ventral views, x 3, showing spondylospirid spondylium and denticulate hingeline; dorsal valve, (49) dorsal and (50) ventral (interior) views, x3, showing denticulate hingeline and paucicostate surface ornament; USNM 450306, from loc. 28. 86 87 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 100 PLATE 11 ‘MUA iiny” oe “ee ry i, 4 i | % y a BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 100 PLATE 12 Figure LATE TRIASSIC CYRTINOID SPIRIFERINACEAN BRACHIOPODS: HOOVER EXPLANATION OF PLATE 12 1=5. Zugmayerella.americana, NEW SPECIES) 2... ee he ees oie ene eo mince ts seine et te ae aisle le wae elie) e 0104 oo ve ee eieicieit ie eiieisie 25-33. 34. Me 2-5. 6-8. Paratype, dorsal valve, dorsal view, x 2, showing valve outline and flattened median fold, USNM 450311, from loc. 28. Paratypes, dorsal valves, (2, 4) dorsal and (3, 5) ventral (interior) views, x 2, showing faint incipient costae on flattened dorsal fold, denticulate hingeline, hinge sockets, median septum and ridges marking point of attachment of descending branches of spire to valve floor; (2, 3) USNM 450313, from loc. 28; (4, 5) USNM 450302, from loc. 20. . Phenacozugmayerella mimuncinata, new genus and new species .... 2... ee eee eee Paratypes, partially decorticated articulated valves, x 1, showing size, form, and outline of a partial simulated ontogenetic series; (6) CASG 66249.01 and (7) CASG 66249.02, from loc. 34; (8) CASG 66251.02, from loc. 37. 2. Paratype, partially decorticated articulated valves, (9) posterior, (10) dorsal, (11) ventral, and (12) lateral views, x1, showing denticulate hingeline, ridged interarea, and pauciplicate ornament; CASG 66251.02 [same as PI. 12, fig. 8], from loc237: . Paratype, partially decorticated articulated valves, (13) anterior, (14) posterior, (15) dorsal, (16) ventral, and (17) lateral views, <1, showing pauciplicate ornament, somewhat globose form and ridged interarea; CASG 66250.01, from loc. 36. . Paratype, partially decorticated articulated valves, (18) dorsal and (19) lateral views, x1, showing form and outline; CASG 66248.03, from loc. 33. . Holotype, partially decorticated articulated valves, (20) anteroventral view, 1, for scale, (21) anteroventral view, x3, showing finely capillate-cancellate surface micro-ornament in ventral mesial sulcus, CASG 66250.02, from loc. 36. . Paratype, fragment of articulated valves, anterolateral view, x 1, showing relation of mesial to lateral plications in a large individual; CASG 66248.04, from loc. 33. . Paratype, partially decorticated articulated valves, anterior view, x2, showing incipient secondary plication in mesial sulcus; CASG 66252.01, from loc. 38. . Paratype, decorticated articulated valves, posterior view, x2 [photographed under glycerin], showing spondylospirid spondylium and posteriorly-produced ventral median septum; CASG 66248.05, from loc. 33. LZ UGINGY TELA TASPECIES Atartsrerarsescre) crete cnet tence ieted een eey ee ohatoone ote earn cna r= See este case cht a ev suenereLelepens} side seis chats (erehavat= lafolegeKedelorote feels 25-29. 30-33. Figured specimen, ventral valve, (25) posterior view, x1, for scale, (26) posterior, (27) anterior, (28) lateral, and (29) ventral views, x 1.5, showing laterally rimmed ridged interarea, globose form and very faint lateral plications, USNM 450314, from loc. 12. Figured specimen, partial ventral valve, (30) posterior view, x1, for scale, (31) posterior, (32) anteroventrolateral and (33) dorsal (interior) views, x 1.5, showing spondylospirid spondylium, very weak lateral plications, and globose form; USNM 450315, from loc. 3. PANG CCL IG ISS EM nog na pba bic cocopdncd oom Gtioanracds pAdonbdnc HouULSe ro OsuonoSoaonOnonee CAndooun nooo Dy abpuDaDo dT Oc Figured specimen, partial dorsal valve, ventral (interior) view, x 2 [stereo], showing mesially recurved, anteriorly jugate descending branches of spire attached to valve floor at midlength, USNM 450317, from loc. 1. 103 90 89 89 104 BULLETIN 337 INDEX Note: Page numbers are in light face; plate numbers are in bold face type; the page numbers on which principal discussions occur are in italics. @yaldefined|| \seess soe eeee ssc ose tos eee otras anes salen ears 74 acrotamboensis, SDINULCHUM Cl metas ere a ae eee e ee aN ace 81,99 SpOndylOspirae ee ee ELI 83,84 AERO GA ire cot onece con econce: Gate tnns Snemstendousectecewde nose rasteuenet es 68 A BORSA SAVE ws te di aace va soe aw ae ae salsa a wate saaosndseeasewaslvewsde cities sa sacaa'es 63 Alaskan sees sin eens sci d.72 se eee 68,78-82,87,88,91 south-central, Alaska; Ranges. ance erncseth- cosas tenet eee tne ans Saes sess Steenes 74 Upper Chulitna district, Chulitna River ..................... 94 INabesnaliquadran gle; recesccsescnsescnss-cacese -a2scse-cneeeateceeeee 733 INutzotinelountalnsipecs.csceeces se seccerseceeeece ec eeceen on eceeeeeee 92 J BSE AG) al) a Se P epee CEE ep ERB ERE ES SHOaEE PEC CaF eR Ce Aue ne ceaede se 92 Copper River Region ..... OD NACKIG LEEK Eee wes ences cer ecesceceres cdnenncs oncescunes Sotvenertcs too 92 SOltheastemnrss eee eee eee eee ere 68,73,92 GComnwallisfheninsulayec.cssessesce terre eee ease ee eer ee eee 73,93 Gravinarisland! yo: tics coos se ten seeme oe oat oem a cs ysensheweers 73 eekuiislan dl Pr vct cece cassenrseess anceeeeetissedsctesachecactssscnecers 73 IekutS traits | ooic. codec soscesencs cssecetes soe teect sotseesctsrscteuveuve 92,93 INGKUSISI Ot SBeernscee ssucs econo ee ars sesaccnecsetees er enscercscaete 93 thiangulationistationiGORIN) wicsecesceeescerece-secscecreecueeres 93 triangulation station LOW 52,9 93 tang wlation’station MEWIMUeicccresc-.-c-cceccecesscoseccers 93 Kine Slan ds eens. eee es reece eee rane eects crc mee 73,92,93 Pups sland yrete. te sscteecccsesce cots centerenenc saccecsecncvcwscrsescesees 92 ScreenylslandSyescrcnes cnceces crores sere eee ease nee 73,92 Glarence|StraitEtolinul slandiseeccs.ssei cee sete eas see seas eaee tee 92 southern, Wrangell Mountains East Fork Valley ....... Green Butte Ridge .... Alexandenterrame sn. .-ctesccancsresneneteseeeee ss Admiral tysubterrane tecccec. esse vec teerteree seer core ee eee Annettesubterrane mete fe see atone ee tec ere ee ree eee Graipisubtenane pessssscccicace sc sence cota caiorecetes stueeednoceeie sre tee alia, Spirifera (Spiriferina) .... 81,100 SSDONGYIOSDING....00.+.500500-< ... 64,81,83,99,100 ILA (alta) eS DONAVIOSDIV are recone eee terete meron seo eee 81 LDS qaeere cerca eee en een ea Tee Cnn oRe TE SHOR eee eo raese rate roee eres 68,87 americana, Zugmayerella ... 11,12 ...... 67,86,87,88,89,101,102 Antelope; Sprines#ROnmMatlOn!:..cscscseecesencastarsatteceer ese access 70,93 Aramachay, ROrmationy sn sctes-c.scevesosseenesenetiods cotenatee reoneerene: 74 Auldiicang:Syne!Groupieseesssesees soaks eee enero reece 69-71 Dun Glen Formation ... 71,85,94 Osobbiormatione.. aamcccesecessoteneeeeens 71,80,82,84,85,91,94 WANNneMuUuCccasEOLMAatlOnimrcssseersee teens eee 71,85,94 Austria; PisGhenwelsersesw esc. ase see aeeesee ae acenae aero eeaae 93 fustr hyceras austriacum Zone [Zone LC2 of Tozer (1980)] ... se eee decasvecsescnseteeee erate ers COOLS 2199 and Speed (1990) Baikans 3egg Formation serg, Jones, and Richter (1972) Big Bend 15’ quadrangle Bittner (1884) Bittner (1892) Bittner (1902) Blackwelder; Es ecsese-s-ecoeccsseesoscee= 71 RockysMountainviirenchits.ccc seeesa- noes eee seen ween ses heen = 93 Rosei@reekol'S quadrangle feec. -o. -- sec cans tore caanccece-c-aserevanee-ese 94 Seta rn errs re area eo oa eco oee costes noc seesssencaecens cacstcsctecsce-seees 94 Samisonrenala (U9 89) secacccerascoscaeoce vancetecence secs esos roe eee cee 69 San bormi(i9 GO) he sresrore nace coco cce coe ones Serer oa ee 71,83 Sanborns Acc b yo ccsssnceaasens canes onope Coenen soe oaeen dee rewe tacts anticsars 94 Sand Springs lithotectonic assemblage of Oldow (1984) ......... 70 Schathautli(8sil))c. ceae aden scossaceesocevees coarse veeee oeneeeoceeea 89,90,92 Schenk ESM cc. oic conc ase ones cies odoasacssateeessucteoceencessecedeoessweacsces 94 SchoonovermSequence ya. .isccaea-cancosnces tes cnes seco note oe sas aa seee econo 71 Seven Devils volcanics 94 Shimer/and'Shrocki(l944)i o-oo csecsmecenecs 64,81,86,99,100 Shoshone Meadows 15’ quadrangle .....................ss.ceeeeeeeeeee 94 Sierrav Neva Ga arGrncceecocesaseceacssacccsesscaedneasessdaceudiceeeeeesees Silberlingi(US90) eo ses cssane cooneca=ceece- 5: sce ecna snot coe Ceon soe ee eee Silberling and John (1989) Silberlingverjaly (9 8i)lns.-cescceecaseacea-cseses=-eeses neste eee Silberling and Roberts (1962) Silberling and Tozer (1968) ............ Silberling and Wallace (1969) SilberlingsJNG J), cc sccescasessenasecscetoeess 63,66,72,73,78,80,84,92-94 Sim MONS GAG ecw ecces cons cost evaene eaeeee oa eee enc ae te tee ene eee 94 Sirenites nanseni Zone [Zone LC3 of Tozer (1980)] ...... 66,82,93 Skwarko, Nicoll, and Campbell (1976) ...................ceeeseeeeeees 68 SmithrandFAeni(lS4)iiscscc. accewccccroncanncaseasessaancoeesase soseeeese 72 Snyder/and! Brueckner (983) <......2.c0s.c-cessersesesecuceatess eee -eaees 71 SOnOmasorOgenly, eee see eee es eee eee a ee Re 71 Sonoma Range lciquadrangles 220. 2s2e. 2.2 sseeee nce eceeeceoetg-ese-a <0 94 SOUtHVA Meri Can: recor ee reeecn: see ceeeee ce eeeececene ... 68,74,81,87 (GH iar rsre Panes Svcd ace eet oe cease cwaansus besneaescsesteneeoeese ee ntes 68 Perulyeseecdec sua sicewessascasset ie csacaessacseans 64,67,68,72,74,82,83,85 Atacocha District, Department of Pasco, Machican Mine area Be Ri Fe Go ne ea aa gu a ace RUNGE TE RCP ATS COREE EES ROR 93 Carhuamayo 74 @erroydesPasco region y-cs-..c-scnccaccs-teawewncecceasettersesce os 70,74 Nimacaat ics: acciscesacseonscawcas soetedeudocnse tanec tecnesteeoeccesuoaes 74 Pasco 74 San Blas area 74 Yanamarca 74 dea: b.8. be ccc 64 fe 74,81,84,85 iyerella : 67,89,102 p. A; Zugmayerellat. ee 12-2 67,89,102 p. indet., Spiriferina(?) peed (1977) <2 See heaves s- cones ooetoce eae eee eee Speed, Elison, and Heck (1988) SpinolepismatinaiDagys, 1974: x ..2 2. c..- =) Ld. ie) & 455 & _ a - _ Ma -_ -_ ate = = 7 = ptnst "Eee s 1 a ol -_ ae) ey tee eed (eee 0.44 e006 Gees! 2106 14 = Pe ie ee ens reo Seal hae oe Or Otis Gms 4 PF * bbe 7 we mw, 1. tints c — oO : a en 7 - 7 —— ANON MAAS MPS : r=: © —p* == - =—28- & 2d -G of Ais. Es ee a “Sas . ap Ce Ne aS a ee ey ee es Ow ~~ =) we Ghai tal 22>” ictal ee 7 Sas ny ag > Gein ates.) (eww Grea See eel... OtE wes it AS ipsa =) Pay » Oe = fe —— a - Seed, see iy) Meanie 9? a = <2 = a _ =e Same) ees = o — —- _ : -— tee ee = ee ee <_—_* GO st= et Se ~~ = —_, ae: sr) ee ee 2 Qa ay ee See oe td Sea eae ea Sh, oe , aba vO een ee Ge ae e= an a f= 2 i) ee Se a i ee ae SyGet.e8 .°. 7.2 oO =e Se a) ieee cee et os 9S SE ee et ee ee ee hj fee S > i G12: Fle @ eee GaP te ae et eee 5 oe Ee OC - See ant f& @«) eG es ae 2 tt oa ee ee caer 3 se PA = LS lL Se ee, Ee ae re oor a > eS a ee sy fie ie pene ae sew = a, Sew He ihe 2 le - a ad a aa ee & S01! (ep fe pe (ens patas seme oo + a Fh i ee > «Wa: ©... 4 Ga eee “= 1 a _ a eet a iol ial 7 . : - ’ That - er PREPARATION OF MANUSCRIPTS Bulletins of American Paleontology usually comprises two or more sep- arate monographs in two volumes each year. This series is a publication outlet for significant longer paleontological monographs for which high quality photo- graphic illustrations and the large quarto format are a requisite. 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