HARVARD UNIVERSITY Library of the Museum of Comparative Zoology (US ISSN 0027-4100) Is ul let in of the seum o Comparative Zoology *«& HH PHP*- . - ■* ■■:•'•■•■ MCZ LIBRARY AUG 1 2 »« HARVARD Volume 1 5f S1TY 1989-1992 HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS 02138 U.S.A. No. 1. The Relationships of the Ai ylopidae (Mammalia v Data and Interpretation. By Richard I ( ifelli ( harh R. Schaff, and Malcolm C. McKenna. Vpril 19£ No. 2. Primary Types of Microlepidoptera in the Museum ol Comparative Zoology (with a Discursion on \ I ( ham bers' Work). By Scott E. Miller and Ronald \\ Hodges April 1990 ,-. No. 3. Early Mississippian Blastoids from Western Montana H\ James Sprinkle and Raymond C. Gutschick Septembei 1990 No. 4. The Neotropical and Mexican Species ol the Orb-Weavei Genera Aranens, Dubiepeira, and Aculepeira ^ran< Araneidae). By Herbert W. Levi. January lcJl)l No. 5. A Review of the South American Lizard Genera I rosti phus and Anisolepis (Squamata: [guania: Polychri- dae). By Richard Etheridge and Ernest E. Williams Ma) 1991 No. 6. The Neotropical Orb- Weaver Genera Edricus and Wag neriana (Araneae: Araneidae). B> Herberts Levi July 1991 No. 7. The Classification of the Naticidae (Mollusca Gastropoda Review and Analysis of the Supraspecific Taxa By Vlan R. Kabat. September 1991 No 8 Amphibians of Southeastern Tanzania, with Special Re! erence to Stephopaedes and Mertensophrym Bufom- dae). By J. C. Poynton. October L991 No. 9. New Flying Lizards and Predictive Biogeography o! I wo Asian Archipelagos. By James Lazell. Max I I 12 (US ISSN 0027-4100) bulletin OF THE Museum of Comparative Zoology The Relationships of the Arctostylopidae (Mammalia): New Data and Interpretation RICHARD L. CIFELLI, CHARLES R. SCHAFF, AND MALCOLM C. McKENNA HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS, U.S.A. VOLUME 152, NUMBER 1 11 APRIL 1989 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952- bulletin 1863- Memoirs 1864-1938 Johnsonia, Department of Mollusks, 1941- Occasional Papers on Mollusks, 1945- SPECIAL PUBLICATIONS. 1. Whittington, H. B., and E. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Tere- dinidae (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4. Eaton, R. J. E., 1974. A Flora of Concord. 236 pp. 5. Rhodin, G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 745 pp. Other Publications. Bigelow, 11. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprint. Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. Creighton, W. S., 1950. The Ants of North America. Reprint. Lyman, C. P., and A. R. Dawe (eds.), 1960. Symposium on Natural Mam- malian 1 [ibernation. Ornithological Gazetteers of the Neotropics (1975-). Peters' Checklist of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1948. (Complete sets onl) Publications <>t the Boston Societ) of Natural History. Price list and catalog <>l MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cam- bridge, Massachusetts, 02138, I S \ This publication has been printed on acid-free permanent paper stock. C The President and Fellows of Harvard College 1989. THE RELATIONSHIPS OF THE ARCTOSTYLOPIDAE (MAMMALIA) NEW DATA AND INTERPRETATION RICHARD L. CIFELLI,' CHARLES R. SCHAFF,- AND MALCOLM C. McKENNA CONTENTS Abstract Introduction Acknowledgments Abbreviations Systematic Paleontology 5 Order Arctostylopida, new 5 Family Arctostylopidae 5 Arctostylops Matthew, 1915 6 A. steini Matthew, 1915 6 Palaeostijlops Matthew and Granger, 1925 1 1 P. iturus Matthew and Granger, 1925 1 5 Gashatostylops new genus 15 G. macrodon (Matthew et al., 1929) .. 15 Sinostylops Tang and Yan, 1976 ... 20 S. promissus Tang and Yan, 1976 ... 20 Bothriostylops Zheng and Huang, 1986 22 B. notios Zheng and Huang, 1986 ... B. progressus (Tang and Yan, 1976) Anatolostylops Zhai, 1978 22 A. dubius Zhai, 1978 23 Asiostylops Zheng, 1979 23 A. spanios Zheng, 1979 Kazachostylops Nesov, 1987 K. occidentalis Nesov, 1987 Arctostylopidae?, incertae sedis Allostylops Zheng, 1979 A. periconatus Zheng, 1979 Arctostylopidae, genus and species indet. Comparative Dental Morphology of the Arcto- stylopidae The Notoungulata of South America Discussion Possible Relationships ^ Remaining Resemblances Distinctness of Arctostylopida 1 Oklahoma Museum of Natural History and De- partment of Zoology, University of Oklahoma, Nor- man, Oklahoma 73019. 2 Museum of Comparative Zoology, Harvard Uni- versity, Cambridge, Massachusetts 02138. 3 American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. Abstract. The dental morpholog steini, hitherto known onl) from pari cheek- tooth series is desi rilx-d on the ba collected specimen including .i near!) per and lower dentition \U arctoMylopid • from North America principally from th Basin, Wyoming, are apparentl) referable l gle species, which therefore ranges from the lati luiiian through the ( larklorkian Other an I genera are restricted to the \si.m I'.ileogi A previously described 5pe< ies "l sty/ops is placed in a new genus Gashal Arctostylopidae and its constituent subordii are diagnosed, and a h\ pothesis "I relationshi| in the family is presented B) comparison witl ungulate morphohpe as represented l>\ / latum, Asiostylops spumes is hypothesized most primitive member ol the family; Both notios and B. progressus retain man) primitivi tures but clearK hear some ol the spec lalizatkx in Palaeostijlops, Arctostylops, and other genera. Of these derived taxa, North \mt tostylops may be the sister taxou to the n genera, all of which are Vsiatfc in distributioi ato/osfy/opi and an as \et unnamed sp< specialized sister taxa that ma) be most dote!) n to Gashatostylops Comparison of morphotypes tor th. daeand for southern Notoungulata sug ivation of one group Irorn within th.- oUm rentlv known, is unlikeb This compai indicates that most notoungulate similai tostylopids were independent!) acquin basis for an exclusive relationship to Notoungulata as sister taxa is a single d. acter. The ankles ol arctostylopids and are divergently specialized and shan not present in a eutherian morpnotype family Arctostvlopulae is. herefon Notoungulata Relatives for th, Holarctic faunas remain unknowi although members of the group r. i er enigmatic .uammaUtrom.h. I Because it is a well-defined mo. out obvious elose relationship I groups, the M,t opidaei Arctostylopida IV. iati Notoungulata removes the i Bull. Mus. Comp Zool 152 ■■ ■ Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 leocene or early Eocene link of Holarctic to Neo- tropical mammal faunas and suggests, in accordance with other evidence now available, that whatever inter American connections of mammal faunas oc- curred must have been earlier in time. The geometry ol li\ pothesized relationships among the Arctostylop- id. te and the fact that the group was most abundant .Hid diverse in Asia suggest an Asian, rather than \.>rth or South American, origin for the family. INTRODUCTION Since the studies of Gaudry (1902, 1904, 1906, 1908) and Scott (1904), it has been widely accepted that South America's fau- na is largely autochthonous, a result of that continent having been isolated by sea barriers from the rest of the world for most of the Tertiary. Endemism at high taxo- nomic levels is particularly conspicuous among the land mammals, which under- \\ ent their great diversification and radia- tions largely within the span of the Ter- tiary. It thus came as a great surprise when, in the first part of this century, apparent members of South America's largest and most characteristic group of hoofed mam- mals, the Notoungulata, were described from specimens recovered in Wyoming Matthew, 1915) and Asia (Matthew and Granger, 1925; Matthew, Granger, and Simpson, 1929). Other possible close rel- atives among Holarctic and Nearctic mammal faunas had been and have con- tinued to be suggested (Ameghino, 1906; Gingerich, 1985; McKenna, 1981). None- theless, none of the proposed relationships seemed so certain, based on characteristic synapomorphies, as in the case of these ungulates, for the Holarctic Arctostylopi- 'l.ir possess a strongly specialized dentition that resembles notoungulates alone among mammals. For this reason, the Arctosty- lopidae have figured prominently in dis- cussions of the origin and early dispersal of South America's native land mammal fauna McKenna, 1981; Simpson, 1951, L978 L980) and of zoogeography in gen- (Colbert, 1973; Darlington, 1957; Simpson. 1965). In addition, because of their presence in North America and Asia, the Arctostylopidae have been integral to the development of correlations of early Tertiary strata (Dashzeveg, 1982; Ginger- ich and Rose, 1977; Matthew and Granger, 1925; Szalay and McKenna, 1971). Arctostylops, represented by the type (and only) species, A. steini, was described by Matthew (1915), based on a partial low- er jaw from the "lower Gray Bull beds, Clark Fork Basin, Wyoming." This local- ity is probably, but not certainly, Clark- forkian in age (Rose, 1981). Matthew re- ferred the genus without question to the Notoungulata, hitherto known only from South America, placing it in the "Entelo- nychia," a mixed assemblage that then contained the most primitive of known no- toungulates. Matthew believed Arctosty- lops to be early Eocene in age, which may well be the case, but is a matter of defi- nition. Further materials of the species were not forthcoming for another 50 years, when a specimen was reported nearby from the Silver Coulee beds of the Polecat Bench Formation near Princeton Quarry. This lo- cality is late Paleocene (late Tiffanian) in age (Jepsen and Woodburne, 1969). Inten- sive collecting by Gingerich, Rose, and as- sociates in Clarkforkian beds of the Clarks Fork Basin has produced four additional specimens, consisting of dentulous lower jaw fragments and isolated teeth (Ginger- ich and Rose, 1977; Rose, 1981). The single report of Arctostylops steini from outside the Clarks Fork Basin is that of McKenna (1980), who recorded the species from beds of Clarkforkian age at Togwotee Pass, northwestern Wyoming. However, related mammals had in the meantime been recovered from Asia. Pa- leontological work at Gashato in Mongolia by the American Museum of Natural His- tory's Central Asiatic Expeditions led to the description of two species, Palaeosty- lops iturus Matthew and Granger, 1925 and "P." macrodon Matthew, Granger, and Simpson, 1929. These species are probably latest Paleocene in age (Szalay and McKenna, 1971). More recent addi- tions to the group have come from slightly • younger deposits at Naran Bulak, Mon- golia (Gradzinski et al., 1969), the Paleo- cene and Eocene (or possibly Oligocene) of China (Tang and Yan, 1976; Zhai, 1978; Zheng, 1979; Zheng and Huang, 1986), and the Paleocene of the USSR (Nesov, 1987), where seven additional described species, placed in six genera, bear witness to a modest radiation of Arctostylopidae in the early Tertiary of Asia.4 Tang and Yan (1976) described Sinostylops, includ- ing two species, from the late Paleocene of Anhui Province, China. S. promissus (from the Dou-mu Formation), the type species, is based on a mandibular ramus with eight teeth; S. progressus (collected in the Shuang-ta-si Group and later trans- ferred to a new genus, Bothriostylops) from six jaw fragments. Anatolostylops dubius was described by Zhai (1978) from the pu- tative early Eocene (but see below) Shi- san-jian-fang Formation of the Turpan Ba- sin, Xin-jiang Province, China. The species is known from a maxillary fragment with well-preserved M2 3. Two additional gen- era and species were published by Zheng (1979). Asiostylops spanios, from the late Paleocene Lan-ni-kong Member of the Chi- jiang Formation, Jiang-xi Province, China, is based on a skull and associated mandible preserving much of the dentition. Because of its primitiveness with respect to other members of the family, Zheng (1979) placed Asiostylops in its own monotypic subfamily. Allostylops periconatus Zheng, 1979, from the late Paleocene Wang-wu Member of the Chi-jiang Formation, Jiang- xi Province, is known from an incomplete rostral part of a skull with poorly preserved P2 to M3. Bothriostylops notios, also from the Wang-wu Member of the Chi-jiang Formation, was described by Zheng and 4 An additional, undescribed genus and species has been reported from the late Paleocene Da-tang Mem- ber of the Nung-shan Formation, Guang-dong Prov- ince, China (Li and Ting, 1983). Dashzeveg (1982) recorded an undescribed species of "Arctostylops from the Bumban Member of the Naran Bulak For- mation, Mongolia, higher in the section than the local occurrence of Palaeostylops iturus. Hua i • Sinostylops pi The most recenl addition to the famil Kazachostylo) Neso\ L981 From the late l tashkent Svita of Kazakhstan i S5 Since the initial descriptii stylops and Palaeostylops Matthew I Matthew and Granger 1 identsl realized that these Holan ti< form some respects, more primitive than known South American Notoungul while in other respects the) are uniquel) specialized. Their primitiveness is refl< ed by placement ol the famil) \n t< lopidae in Simpson's archaic notoungulate suborder Notioprogonia Simpson 1 1945). However, the relationships ol Hoi- arctic to South American forms have n< been considered in detail Foi this reason a variety of opinions exist as to the place of origin of notoungulates and theii sub- sequent dispersal patterns, with authors variously favoring northern Patterson 1958) or. speeitiealb \-i.ui Matthew 1928; Zheng. L979 Smith American (Hoffstetter, 1970: Marshall de Muizon and Sige, 1983; Simpson 1951 and ( en- tral American (Gingerich and I ll»7' centers of origin. Until recently, the data base for making such an assessment has been rather limited The early Tertiar\ South American no- toungulates have received monographs treatment (Simpson. 1948, 1967 rheHol- arctic radiation, the Vrctostylopidae was long represented onl) b) the single lowei dentition originall) described lor \ortl American Arctostylops rteini and b) den- titions of two species referred to Vsiai laeostylops. Although some tonus remain poorly known, Vsian arctostylopid taxa de- scribed in recenl years add substantial!) t knowledge of morphological divers! within the group, offering a dramaticall) improved basis for comparison with N< toungulata Herein we describe the dentition tostylops steini, much of which has • hitherto unknown, based on a new!) 4 Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 BUCCAL ANTERIOR - POSTERIOR LINGUAL Mesostyle Parastyle Fossette Preprotocrista Protocone Precingulum Metastyle Ectoloph Postmetaconule crista Postprotocrista Pseudohypocone Postcingulum Sulcus pseudohypocone Lingual cingulum Ectocingulid cristid obliqua (selenolophid) Paracristid Protoconid Precingulid""^ \yr A \ Talonid basin Hypoconulid Entolophid Entoconid Protocristid Trigonid Metaconid Talonid notch notch Figure 1 . Dental terminology used in describing arctostylopid molars, based on Palaeostylops iturus (after Szalay, 1969). lected and remarkably complete specimen from the late Tiffanian of Wyoming. This specimen (<>rms the basis for a revised di- agnosis of the genus and species and for a comparison w Lth Vsian Axctosty lopidae and South American Notoungulata. Revised diagnoses are presented for previously de- scribed taxa; we refer "Palaeostylops" macrodon to a new genus. Formal descrip- tion of a hitherto unknown species of arc- tostylopid from the Yan-ma-tou Forma- tion, Hunan Province, China, is currently Arctostylopids lia) • ( ifelit ei al in progress; for comparative purposes, we briefly review some of its morphological features. Another new genus and species, from the Da-Tang Member of the Nung- shan Formation, Guang-dong, is being de- scribed by others elsewhere. ACKNOWLEDGMENTS We thank Dr. Donald Baird for loan of a Princeton specimen under his care; Dr. Anne Roe Simpson, Mr. Will Downs, and Mr. Meng Jin for providing us with trans- lations of some important Chinese papers; Mr. Kenneth Sauer for translation of a work in Russian; Ms. Ting Su-yin and Drs. Ken- neth Rose and Louis L. Jacobs for various comments and information; and especially Dr. Philip D. Gingerich for providing us with casts of Asian Arctostylopidae and for facilitating access to areas currently under investigation by the University of Michi- gan Field Program. Dr. Zheng Jia-Jian gra- ciously permitted us access to an unde- scribed specimen under study by him. Part of this research was undertaken while Ci- felli was a Research Associate with the Bi- ological Survey, New York State Museum, and the support of that organization is gratefully acknowledged. Exploration of Silver Coulee sites was supported by Na- tional Geographic Society grant no. 2057 to Schaff. We extend our thanks to the Churchill family of Powell, Wyoming, for their assistance and generous hospitality over the years, in connection with field work by Schaff. Field assistance was pro- vided by Dr. Farish A. Jenkins, Jr., and by Messrs. William Amaral, Mark Goodwin, Charles R. Schaff, Jr., and Rick Schaff. The drawings were prepared by Mr. Laszlo L. Meszoly and Ms. Coral McCallister, and the photographs were taken by Mr. Al- phonso H. Coleman. We thank also Mrs. Lillian W. Maloney for her assistance in preparation of the manuscript. ABBREVIATIONS AMNH, Department of Vertebrate Pa- leontology, American Museum of Natural History; IVPP, Institute of Vertebrate Pa- leontology and Paleoanthropolog jing, People's Republic oi China M( / Museum ol Comparath vard I niversit) I \\ I niversit) -I Mi. I, igan; YPM-Pl . Yale Peabod) Museum Princeton I niversit) ( oil.-, tion Dental terminology is illustrated in I ure 1. SYSTEMATIC PALEONTOLOGY Order Arctostylopida. new Distribution. Extinct; presentl) known only from the Paleocene, 1 ocene .md|H,v sibly the Oligocene <.l \sia; Kit. Palea ene and possibly earl> Kmnic <>1 North Amer- ica. Diagnosis. Small mammals with uppei and lower dentitions forming an evenl) graded series; canines | rl\ or not dif- ferentiated and without diastemal rating them from adjacent teeth Posterioi upper premolarssomewli.it molarized ex- cept in Asiostyl(J))s. I", at least, with a metacone. Upper molars with well-devel- oped centrocrista. becoming a salient, straight ectoloph in advanced g< nera parastyle usually prominent Pn and postprotocristae of upper molars strong, conules lacking; upper lars primitive!) triangular but M1-2 becoming quadrate in advanced forms 1>\ the addition <>t .. |x>s terolingual cusp (pseudohypocone In- terior lower premolars seriall) tricuspid, with strong shearing surfaces; lowei mo- lars primitively biselenodonl with para- cristid lost and various accessor) trigonid structures acquired in advanced taxa Lower molar hypoconid indistinct; ent< conid transverseK expanded and. m ad- vanced forms, developed into an antero- bucally oriented entolophid. Family Arctostylopidae Schlosser. 1923. p. 614 (=Subfamily Arctostylopmae Zheng. 1979. p. 391) Type Genus. Arctostylops Matth< 1915. p. 429 Other Included Genera Anatolostyi Zhai 1978, p 1"'' isiostylopi Z\ 6 Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 1979, p. 388; Palaeostylops Matthew and Granger, 1925, p. 2; Sinostylops Tang and ^ an, 1976, p. 91; Bothriostylops Zheng and Huang, 1986, p. 121; Kazachostylops Ne- sov, 1987, p. 212; Gashatostylops, new; an unnamed genus; and, with some doubt, Allostylops Zheng, 1979, p. 391. Distribution. Paleocene, Eocene, and possibly the Oligocene of Asia; late Paleo- cene and possibly early Eocene of North America. Diagnosis. \s for the order. Zheng (1979) divided the Arctostylopi- dae into two subfamilies: the Arctostylop- inae, which included "typical" genera; and the Asiostylopinae, containing only Asio- stylops itself. While we are in agreement that this last-named genus is the most primitive of known forms, we choose not to recognize a higher taxon (subfamily) on that basis alone. Moreover, the description of species "intermediate" between Asio- stylops spanios and advanced forms (see Zheng and Huang, 1986) largely occludes the morphological hiatus distinguishing the proposed subfamilies, so that they are not even clearly defined grades. Nesov (1987) distinguished two further arctostylopid subfamilies, Sinostylopinae and Kazacho- stylopinae. On the basis of evidence now in hand, we do not believe that such di- vision of the group is warranted. Arctostylops Matthew, 1915, p. 429 Type Species. Arctostylops steini Mat- thew, 1915, p. 429. Included Species. The type only. Distribution. Late Tiffanian to late Clarkforkian, and possibly Wasatchian, North America. Diagnosis. Large arctostylopid differing From Palaeostylops and all other members of the family in having a salient lingual rib on the lower canine, a molarized P., with a low, recurved talonid loph that ex- tends lingually at the posterior margin of the tooth, and a prominent anterolabial cingulum (ectocingulid). Distinct, where known, from primitive genera (Asiosty- lops, Bothriostylops) in having quadrate upper molars with a sulcus separating two lingual cusps on M12. Upper molars differ further from those of Asiostylops in having a strongly developed ectoloph and in lack- ing a paracone fold. Lower molars differ from Asiostylops, Kazachostylops, and Bothriostylops in having paracristid re- duced, prominent ectocingulid with shear surface descending from protoconid, cris- tid obliqua achieving a pronounced labial attachment to the trigonid, and entolophid stronger and more oblique. Metacones on P23 lacking or not so well-developed as in Palaeostylops and Gashatostylops; a lin- gual cingulum is present on P4 and is more salient than in those genera. M1 2 more transverse, less quadrate in occlusal view; M2 sulcus between protocone and pseu- dohypocone not so well-developed as in Palaeostylops or Gashatostylops. Meta- conid of lower molars not forming a dis- tinct column within the talonid basin as in those two genera. Pre- and postprotocris- tae of upper molars high and variably en- closing a very transient fossette, as occa- sionally seen in Palaeostylops and Gashatostylops, but not so strongly devel- oped as in Anatolostylops. Arctostylops steini Matthew, 1915 Figures 2, 8, 9 Arctostylops steini Matthew, 1915, p. 429; Jepsen and Woodbume, 1969, p. 546; Rose, 1981, p. 965 Holotijpe. AMNH 16830, left mandibular ramus with P3 to M3. Referred Material. MCZ 20004, asso- ciated mandible and anterior part of skull with nearly complete upper and lower dentitions; YPM-PU 20397, poorly pre- 5 The listing of this species as "Palaeostylops stei- ni" by Thenius (1985, caption to Fig. 1, p. 151) de- serves mention, although a text explanation is lacking and we are thus uncertain as to whether this is a lapsus or implied synonymy. The figure itself is dia- grammatic but suggestive of Palaeostylops iturus rather than A. steini (for which well-preserved upper molars have not been previously reported otherwise). Vs indicated in the diagnoses, the species are clearly distinct; regardless, Arctostylops is the prior name. ARCTOSTYLOPIDS M wim \i.i \ • ( |/i //, ,., al cm B Figure 2. Stereophotographs of upper (A) and lower (B) dentitions of Arctostylops stein,. MCZ 20004 8 Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 served incomplete skull and mandible; UM is mitten-shaped, with a prominent distal 650^4 left dentarv fragment with worn heel. A cingulum, lacking on the labial side \1 and right dentarv fragment with P3; of the tooth, is well-defined on the lingual I \1 66707, right dentary fragment with portion of the crown. I2 is represented only M, and partial M2; UM 68863, right M2; by a fragmentary part of the crown. As I \1 69280, right P3 (UM specimens are with the preceding teeth, P is single-root- cited from Rose, 1981, p. 96, and have not ed. The crests descending from the single been studied bv us); and AMNH 88141, cusp are sharp; a small heel is present. A trigonid of left M,. weak labial cingulum is present; a lingual Horizons and Localities. The type was cingulum appears to have been well-de- collected in the "Lower Gray Bull beds, veloped, but breakage obscures most of Clark Fork Basin, Wyoming" (Matthew, this side of the tooth. The upper canine is 1915, p. 429), of probable late Clarkfork- similar to the incisors and, unlike those of ian (Rose, 1981) or, possibly, Wasatchian Palaeostylops and Gashatostylops, which (Jepsen and Woodburne, 1969) age. Re- are subequal in size to adjacent teeth, is f erred specimens have been collected from larger than P and P1. The single root is the Willwood Formation at University of round to oval in cross-section and is not Michigan localities SC-19, 116, 188, and well-differentiated from those of the ad- 203 in the Plesiadapis cooki and Phenac- jacent teeth. The crown bears sharp mesial odus-Ectocion zones, Clarkforkian, Clarks and distal crests, is labiolingually corn- Fork Basin, Wyoming (Rose, 1981, p. 96); pressed, and is somewhat inclined poste- in the "lower variegated sequence" (Love, riorly; the labial surface is convex and the 1947) of an unnamed formation, Clark- lingual surface is slightly concave. The dis- forkian, near Togwotee Pass, Wyoming tal coronal crest bears a small, compressed (McKenna, 1980, p. 330); Silver Coulee cusp followed by a faint heel. The cin- beds, Polecat Bench Formation, Plesia- gulum is well-defined both lingually and dapis simonsi zone, Tiff anian (Jepsen and labially; the posterolabial part bears poorly Woodburne, 1969, p. 546), Wyoming. The defined cuspules. There are no diastemata specimen described below, MCZ 20004, adjacent to the canine, was collected by Charles Schaff and Mark P1 is single-rooted and bears a single cusp. Goodwin in 1977, approximately 5 m from The tooth is labiolingually compressed, the Princeton Quarry site (Jepsen, 1930). with a faint lingual bulge, and closely re- The specimen was excavated from a gray- sembles the larger canine. The lingual cin- green siltstone 2.5 m below the Princeton gulum is prominent. Salient crests descend Quarrv level. The localitv (MCZ number from the anterior and posterior ends of the 1/77WYO; SE Va sec. 21, T56N, R100W) tooth to the single cusp. These evidently is about 24 km northwest of Powell, Park were important shearing structures, as a ( .'()., Wyoming, on the west side of Polecat well-defined wear surface is developed on Bench. the lingual side of the tooth, obscuring any Diagnosis. As for the genus. detail that may originally have been pres- ent. P2, also anteroposteriorly elongate, has _ _ two roots and is triangular in coronal view; DESCRIPTION .i . ,, , r . i . j. i. the serial homologue or the protruding hn- The upper and lower dentitions form gual cingulum on P1 is here developed into evenl) graded series, w ithout diastemata a protocone. Labially, the ectoloph is sup- or marked structural gaps between teeth, ported by a single prominent cusp, the is ti.»t preserved in place in MCZ 20004. paracone, from which the loph descends However, two isolated upper incisors, one anteriorly and posteriorly. The anterior o\ which has been lost, were found in as- surface is moderately worn, with the facet iation with the upper dentition and angled sharply with respect to the plane probabl) represenl this tooth. The crown of occlusion. This facet is continuous with AR( lLIA) • < ifelti ei al Table 1. Dental measurements of . AMNH 16830 MCZ 20004 PU 20397 65024* • ' .-.» \. Lower dentition Ci L — — — — — W — — — — — — Pi L — 1.6 — — — W — — — — — P2 L — 3.0 — — — W — 1.8 — — — — P3 L 3.3 3.4 — 3.6 — — W 1.7 2.0 — 2.1 — — 1 7 P4 L 3.7 3.8 — — — — W 1.8 2.0 — — — — MjL 4.0 4.0 — 3.9 — W 1.8 2.0 — 2.3 2.3 — MoL 4.2 4.7 — — — . 1 W 2.1 2.3 — — — 2.2 M3L 3.9 4.5 4.3 — — — W 1.8 1.9 — — — — — MCZ 20004 PU 20397 L w L \\ B. Upper dentition C1 3.1 2.0 — — P1 2.9 1.8 — — p2 3.2 2.5 — — p3 3.5 3.3 — — p4 3.6 4.2 — — M1 3.7 4.9 — — M2 4.5 5.6 3.7 4.1 M3 4.0? 5.1 3.7 4.0 *From Rose (1981, p. 97) another wear surface that extends from the region of the parastyle to the protocone, along the anterior portion of the lingual cingulum. A faint bulge anterior to the paracone suggests that in the unworn con- dition a parastyle was present. The part of the ectoloph distal to the paracone bears a strongly developed wear surface, also steeply angled with respect to the occlusal plane. P3 is larger than P2, with a better developed protoconal region, more salient paracone fold on the labial surface of the ectoloph, and three roots, but is in most other respects similar to P2. P4 bears a well- developed, prominent protocone and is therefore considerably more transverse than P4. The ectoloph is folded at the para- cone. The lingual surface of the ectoloph is considerably worn, but a parastyle and, with less certainty, a metacone maj be distinguished. A well-developed cresl ex- tendsfrom the protoc-nur t.> the parastyle As with P3, wear on this nest is continuous with that on the anterolingual part "t the ectoloph. The remnant ol a small fossette persists in the trigonol theright I' \ well- developed cingulum extends from parastyle around the based the proto and along the posterior border of the I I" is nearl) the same length as \1 \1> 2aremorphologicall) similai toeacn other, the principal difference being that MMs somewhat larger than M' > >< ference in relative size is less in I The ectoloph is anteroposterior!) straigti the only departure from this beingth lient parastyle, which is develop, column on the labial wall of the ectoloph 10 Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 This ectoloph outer wall also has a postero- seems to have been unfused. Small mental inferiorlv developed bulge, probably cor- foramina are located below the right P4 responding to the base of a metacone or and below left I, and I2, respectively, metastvle. The inner face of the ectoloph I, is not preserved in MCZ 20004. I2 is on each molar bears a very well-developed procumbent and spatulate, with a long wear surface, oriented, as on the premo- straight root that is round in cross-section, lars superolingually at a steep angle to the An oblique ridge traverses the lingual sur- plane of jaw occlusion. A lingual sulcus face of the crown. I3 and the lower canine separates the protocone from another cusp much resemble I2, differing in not being distal and somewhat appressed to it; this procumbent. The canine is thus incisiform, latter cusp we believe not to be a true structurally undifferentiated, and not sep- cingulum hvpocone, for reasons developed arated from adjacent teeth by diastemata. below. The crests linking protocone to The crown of C, bears a well-developed parastyle (preprotocrista) and protocone to lingual column; posterior to this, two cus- "pseudohvpocone" (Gregory, 1920; Simp- pules, separated by a notch, are present, son, 1929) to the posterobuccal angle of P! is missing as a result of postmortem the tooth (postprotocrista) maintained a damage in MCZ 20004 and is represented primitive triangular arrangement with re- only by a small remnant of one heel. The spect to the ectoloph and were evidently tooth was single-rooted. P2 is a larger tooth strongly developed, because a small rem- and is double-rooted. It is buccolingually nant of a fossette enclosed by them persists compressed and bears three principal cusps on the left \T and right M2. These heavily that are nearly in line with each other, the worn crests descend6 buccally from the middle of which is the tallest. The ante- protocone to their junction with the de- riormost two cusps are separated by a dis- scending wear surface of the ectoloph de- tinct notch; the third cusp lies on the pos- veloped on the labial face of the trigon terior slope of the middle cusp and has basin, so that the molars appear to be been reduced in this specimen by wear. notched when viewed anteroposteriorly. Behind this the central crest slopes infe- M\ somewhat damaged on both sides of riorly before rising to a sharp heel at the the specimen, is smaller and more trian- distal margin of the tooth. A slight bulge gular in outline than M2. An accessory crest, is present on the inferolabial side of the apparently lacking on M12 but perhaps not tooth, but this is not distinctly formed into seen because of heavy wear on those teeth, a cingulum. P3 to M3 are similar to those sweeps posterolabially from the midpoint of the holotype, AMNH 16830, as figured of the postprotocrista to the base of the by Matthew (1915). P3, like P2, is trenchant metacone (or metastvle). A small accessory and is similar to that tooth except for being crest, the postmetaconule crista, is present larger. P4 is submolariform. The paraconid on the left M" (the right M3 is damaged), is lower than and directly mesial to the As with the more anterior molars, a distinct protoconid; the metaconid is lingualfy lingual cingulum is present and appears to placed. The protoconid and the metaconid be confluent around the base of the pro- are subequal in size. The cristid obliqua tocone attaches to the trigonid somewhat nearer The mandible is shallow and somewhat to the metaconid than to the protoconid I shaped at the symphysis. The symphysis and extends superiorly to a level near the apices of these cusps. The talonid is formed by a simple, crescentic crest that termi- , f „ nates at the posterolingual angle of the I "i uppei teeth we lollow convention in using . .i A n i i • i • i / cend," .l.-cend," "superior," "inferi toofh- A sma11 anterolabial cingulum (ec- I so forth, in a sense relative to the way the) tocingulid) is present. ith reference to orientation in life. The lower molars are morphologically Aw . , , al n similar to each other. This series may differ slightly from that of the holotype, AMNH Palaeostylops Matthew and Granger, 16830, in that M2 is more distinctly the 1925- P- 2 largest of the three. The paraconid and its Type Specit s Palai ostylop Hurt linking crest are altogether lacking, and thew and Granger L92i the protoconid is near the anterolabial Included .Sp, margin of the tooth. From this cusp a crest Distribution Late Paleo descends anterolabially, forming a distinct Eocene (fide Li and Tin ridge (ectocingulid) at that corner of the Diagnosis. Dentall) advi -d arcto tooth; the protoconid is also slightly ex- stylopid generall) similar to in panded into an anterolingually developed but differing in the lack ol a heel ridge. The cristid obliqua has an extremely lack of paracolic Folds on the « tolophs labial attachment to the trigonid; i.e., at P5 '. lack of a lingual rib on ( and in the the protoconid. From this point, at which lesser size differentiation ol the uppei it is nearly as high as the trigonid, it ex- nine from adjacent teeth. Molars \o\ tends distally as a sharp, straight loph, be- crowned than in Anatolostylops; uppei fore curving somewhat lingually to end at molar fossette more rapidK li»s| b) dental the hypoconulid. A hypoconid as such is wear. Differs from Gashatostylops ma lacking. The entoconid is transversely de- don, the most closel) similar form, in de- veloped into a loph (entolophid), which ing a strong sulcus separating tin lingual extends anterolabially to join the principal cusps of M\ three upper incisors and an talonid loph (cristid obliqua and postcris- unconstricted snout, and in lacking ( us- tid) at about its mid-point. Measurements pules on upper molar lingual cingula and are given in Table 1. relative enlargement of the upper and low- Available materials of Arctostylops are er second molars, inadequate to properly assess specific vari- Both Palaeostylops Hunts and Gasha- ability. All specimens in the hypodigm in- tostylops macrodon (herein separated from elude teeth also represented in the type of Palaeostylops) were described from the A. steini (AMNH 16830) and are suffi- type Gashato Formation Matthew and ciently similar to them in known morpho- Granger 1925; Matthew. Crangei and logical features to cause us to consider all Simpson, 1929). Both species, but i specimens to belong to the same species, cially P. iturus, are know n from large sam- P3 and Mj are represented by four speci- pies of rather complete dental materials mens each; M2 and M3 are known by three Further remains ol both spe< ies have beei teeth each. Of these, M2 shows a marked recovered in the Naran Bulak I ormati variability in length (Table 1). P3 seems to by Soviet and Polish-Mongolian ei vary considerably in proportion of length tions (Gradzinski et al., lc>dl). Szala) an to width, but the significance of this cannot McKenna, 1971) to the Nemeg now be determined. about 250 km W'SW ol Gashato; in the As thus recognized, the species A. steini Nomogen Formation, near Nom is known from sediments of late Tiffanian Mongol, by Chinese workers (Plesiadapis simonsi zone) through Clark- 1977; Chow and Qi, 1978); and i forkian (Phenacodus-Ectocion zone) or van Ulan Formation Nei possibly Wasatchian age. This is a wide 1979). Individuals oi these species stratigraphic range for mammalian species sent by far the most abundant of that age; however, several other species, of the Gashato and including the abundant phenacodontids curious fact considering tl Phenacodus primaevus, P. vortmani and close relative, Arctostylops, Ectocion osbornianus, are believed to have temporaneous North Amer • -i /d ioqi ™ 99 9 tin- »i.iui^ similar ranges (nose, 1981, pp. LA-LA). Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 log(M1 area) Figure 3. Plot of log-transformed M, area vs. M2 length for Palaeostylops iturus (dots) and Gashatostylops macrodon (squares). the genus Palaeostylops, its contained species, and the relationships of those species to Arctostylops steini. Simpson (1936a) indicated that the species P. iturus and "P." macrodon might be considered as closely allied but distinct genera; Dash- zeveg (1982) referred both to the North American genus Arctostylops. The super- ficially close similarity of the Asian species (except for size) and the fact that they always co-occur suggested to us, at the out- set of this study, the possibility that a sin- gle, sexually dimorphic, species was rep- resented. Detailed qualitative and quantitative comparisons, presented be- low, together with previously unknown morphology provided by a new specimen, uphold Simpson's view. To explore the dif- Ferences between these superficially sim- ilar speeies, we examined available VMNH) samples of arctostylopid denti- tions from the Gashato and Nomogen lo- calities and performed univariate and multivariate analyses on tooth dimension length, width' data, using the Systat mi- iputer software package. Matthew . ( Granger, and Simpson (1929) distinguished "P." macrodon from P. itu- rus by its larger size and its proportionately larger second upper and lower molars. Comparison of type and referred materials reveal several other consistent morpholog- ical differences, summarized in the diag- noses and description given below. The most obvious difference in specimens as- signed to the two species, other than ab- solute size, is the aforementioned dispro- portionately large upper and lower second molars of "P." macrodon (Table 2). Length measurements of these teeth do not even overlap in range, which would be expected if the difference were due to sexual di- morphism. In most mammals (Gingerich, 1974), M, is the least variable lower molar; in Palaeostylops iturus, the species for which samples are most nearly adequate, variability is comparable between Ml and M3 (Table 2). Because M, is represented by larger samples in both species, this tooth was chosen as a basis for comparison of second molar proportionate size. A plot (Fig. 3) of log-transformed M^ area (length x width) against M2 length indicates that the difference in relative length of the sec- • al 2 -■ 1 -■ *- 0 ■■ o <0 1 -. -2 -■ A -t— • • -+- -1 factor 2 1.0 ■■ 0.5 ■■ o o.o ^ u 10 ■0.5 ■- -1.0 ■- -1.0 ■0.5 B -i- w, w • 1 M3W 0.0 factor 1 2 0.5 .V V_ M L 3 Figure 4. PCA loading plots for six lower molar variables of Palaeostylops and Cash on first two axes (dots = P. iturus: squares = G. macrodon), B. loadings for the six vanables 1.0 Unoi 11 in Museum of Comparative Zoology, Vol. 152, No. 1 TABI 1 7 MEASUREMl NTS KND SI MMAHV STATISTICS OF PALAEOSTYLOPS AND GASHATOSTYLOPS (MM; P. ITURUS, G = G. macrodon). p = A. Upper cr ieek teeth P'L p3W P^L pjw M1 L P G P G P G p G P G \ 2 2 2 2 4 2 3 2 5 5 Minimum 1 880 2.140 1.690 1.850 1.410 2.070 2.210 2.290 2.260 2.670 Maximum 1.990 2.170 1.880 2.000 2.090 2.120 2.320 2.530 2.700 3.220 \li '.in 1.935 2.155 1.785 1.925 1.812 2.095 2.273 2.410 2.532 2.936 si) 0.078 0.021 0.134 0.106 0.304 0.035 0.057 0.170 0.170 0.210 M'W M' ;L M2W M3L M3W P G P G P G p G P G N 5 5 5 5 4 5 4 3 3 3 Minimum 2.340 2.790 2.670 4.320 3.170 3.630 2.030 1.940 2.570 3.020 Maximum 2.960 3.340 3.560 5.190 3.530 4.100 2.340 2.530 3.060 3.100 Mean 2.772 3.026 3.138 4.788 3.398 3.874 2.135 2.323 2.810 3.063 SD 0.250 0.209 0.328 0.321 0.161 0.177 0.139 0.332 0.245 0.040 B. Lower cheek teeth PsI P3w P4L P4W MiL P G p G P G p G P G N i 3 7 3 10 4 9 3 16 7 Minimum 1 760 2.090 0.980 1.230 1.960 2.500 1.040 1.300 2.440 2.890 Maximum 2,310 2.370 1.220 1.280 2.500 2.700 1.280 1.440 2.950 3.220 Mean 2.011 2.240 1.164 1.253 2.303 2.612 1.170 1.393 2.737 3.064 SD 0.208 0.141 0.086 0.025 0.153 0.084 0.081 0.081 0.120 0.141 ( Yar 10 3 — 7,3 — 6.6 — 6.9 — 4.4 — MiW M2L M2W M3L M.3w P (; P G p G p G P G N 16 7 16 12 16 12 11 4 11 4 Minimum 1.270 1.420 3.060 4.150 1.510 1.750 2.370 3.090 1.330 1.330 Maximum 1.630 1 .610 3.850 4.890 2.010 2.550 3.030 3.320 1.520 1.800 Mean 1.425 1,533 3.537 4.592 1.744 2.063 2.694 3.185 1.407 1.505 SI) 0.140 0.077 0.194 0.248 0.145 0.248 0.172 0.097 0.060 0.205 ( Yar 7,3 — 5.5 — 8.3 — 6.4 — 4.3 — ond molar between the two morphs is not a factor of scaling, i.e., an allometric effect attributable to the fact that "P." macrodon is larger than P. Hums. Were this the case, all specimens would have fallen along the same line; in the present situation, two lines, with different Y-intercepts, are apparent. To evaluate the significance of differences in measurement means, independent T-tests were performed on the lower elieek-tooth data. For most variables, means of samples assigned to the two species were si nilieantly different at the 05 level (Table probability of identical means was high( stforP3l i' \\ , M,W,and \1 w and lowesl foi Ml Ml., M3L, and M2W. Principal components analysis, which does not require prior taxonomic sorting, was performed on various com- binations of both untransformed and log- transformed lower molar data (the corre- lation matrix with listwise deletion of miss- ing data and varimax rotation were em- ployed). These analyses consistently separated the specimens into two groups (corresponding to the two species) along the first axis (presumably attributable to size) that, for the untransformed and un- rotated lower molar data, accounted for about 61% of the total variance. Factor loading plots (Fig. 4) indicate that the source of this separation is the three length Arctos ds (Mammalia • i IfeUietal variables (M,L, M2L, M3L), which have very high loadings along the first axis; a result consistent with the univariate anal- ysis. Thus, on a statistical basis, the differ- ences between P. iturus and "P." macro- don are significant and are not attributable to size alone. In addition, dental and cra- nial features indicate greater structural dif- ferences between the species than has hith- erto been appreciated. We consider these differences to be worthy of generic sepa- ration. Palaeostylops iturus Matthew and Gran- ger, 1925, p. 2 Arctostylops iturus Dashzeveg and Rus- sell, 1988, p. 131 Figures 7, 8 Holotype. AMNH 20414, right mandibular ramus with broken I, 2 and with I3 to M3 complete. Referred Specimens. The type, and the following AMNH specimens, consisting of dentulous upper and lower jaws or portions thereof: 20415, 20417, 22143, 101967, 101968 (uppers); 20429, 21723, 101983, 101985; and AMNH 109522 A-J, casts of 10 uncatalogued lower jaw specimens in the IVPP. The AMNH specimens are from Gashato; the IVPP specimens were col- lected at Nomogen. Additional materials referable to the species are housed at the Polish Academy of Sciences, Warsaw, and at the Paleontological Institute, Moscow. These specimens are not listed here be- cause we were not able to compare them directly with the fossils listed above. Horizon and Localities. Late Paleo- cene; Gashato, Bayan Ulan, Naran Bulak, and Nomogen formations, Nei Mongol. Diagnosis. As for the genus. Gashatostylops, new genus Type Species. Palaeostylops macrodon Matthew, Granger, and Simpson, 1929, p. 11. Etymology. Gashato-, for the original locality of the type species; -sty lops (Gr.), pillarlike, a commonly-used suffix for arc- Tabli 3 Homogeneity ,■ M ,.. LOWER ( in i k IMMIM- ./ ' 'S •/■■) / ,,/'S in in s vnd \ ariable >.irlil,.«l P.3 length 1.714 0 1 Pi width 1 7os (1 P4 lenulh 3.768 P, width I l in o.o \li length 5 725 0.000 \li \\ idtli 2 • M2 length 12 '.-,ii 0.0 \lj width 1 21.7 ii 01 \l | length 5 (20 ii 000 M3 width l 501 11 i tostylopid and primitive notoungulate genera. Distribution. Late Paleocene to early Eocene {fide Li and Ting, 1983 . tsia Diagnosis. Advanced arctost) lopids dif- fering from Palaeostylops, the most < losel) similar genus, in having relativel) en- larged upper and lower second molars; in having cuspules, variable in numbei and development, on the lingual cingula oi up- per molars; in the weakness or absem a sulcus separating the lingual cusps ol \l . in the presence of two rathei than three upper incisors; and in having a lateral!) constricted snout, with the dental arcade multiply curved. Differs from Inatolo- stylops, to which it ma) beclosel) related, in having lower-crow ned cheek-teeth and in having upper molars with aivrssor) cusps and plications on the lateral w alls i »l the ectolophs. Gashatostylops macrodon (Matthew. Granger, and Simpson, 1929) Palaeostylops macrodon Matthew. Granger, and Simpson, 1929, p. 11 Figures 5-9 Holotype. AMNH 21725. left mandibulu ramus with P3-M2. Referred Specimens The type, an un catalogued IVPP specimen casts, tMNH 109521) consisting ol the left rostral pari of a skull with roots ,.! two right and lej incisors, left < M and part ol the left mandible w itli M3, pins an astragalus, col- 16 Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 leeted by McKenna; AMNH 21742, two isoalted calcanea; AMNH 21726, isolated right astragalus; and the following AMNH specimens consisting of dentulous upper and lower jaw fragments: 22142, 101967, 101979, 101977, 101963 (maxillary); 101980, 101987, 101984, 101982, 101981, 20416, 21740, 21741, 21723, and 21716 (mandibular). The AMNH specimens were collected at Gashato, the IVPP specimen is from Bayan Ulan. As with Palaeostylops iturus, additional specimens (not listed here) are in the collections of the Polish Academy of Sciences, Warsaw, and the Paleontological Institute, Moscow. Horizon and Localities. Late Paleo- cene; Gashato, Bayan Ulan, Naran Bulak, and Nomogen formations, Nei Mongol. Diagnosis. As for the genus. Although a diagnosis appeared in the original publication (Matthew, Granger, and Simpson, 1929), the morphology of this species has never been described. Im- portant details are provided by the spec- imen represented by AMNH 109521 from Bayan Ulan, which preserves the left side of the rostrum, including the orbit and zygomatic root, left C-M3, and the roots of the incisors on both sides. In addition, further preparation of the original IVPP specimen by one of us (Schaff) revealed the presence of part of the left mandible, including M3 and the condyle, and a right astragalus lodged within the broken cra- nial cavity. These are presumed to be as- sociated with the skull fragment itself. All teeth are in full eruption but wear is light, indicating that the animal was a young adult. The specimen is more or less split sagittally, except that both premaxillae are preserved. The palate, nasal, and frontal regions are crushed, so that the corre- sponding bones are somewhat fragmented. Comparison with the dentition preserved in original paratypesof the species (AMNH 221 I t Ml P3-M2; 22142, right broken M1 and complete M23) and other referred ma- terials from the type locality leave no doubt as to reference of this specimen to Gash- atostylops macrodon. As preserved in AMNH 109521, the snout is short and constricted, flaring broadly at the root of the zygomatic arch, so that in palatal aspect the tooth row as- sumes a double curvature. The form of the dental arcade thus contrasts with that seen in Palaeostylops iturus, which curves gently from front to back. In palatal view, the posterior margin of the maxilla forms a curved process that almost completely encloses a small foramen lingual to the junction of M2 and M3. This foramen in all probability housed the minor palatine branch of the maxillary artery, as it does in many living mammals and in certain notoungulates, such as Notopithecus (see Simpson, 1967, fig. 23). The infraorbital foramen, located above the junction of P3 and P4 about halfway between the base of those teeth and the anteroinferior margin of the orbit, is small. The root of the zy- gomatic arch arises at the base of M2. It is massive and dorsoventrally expanded, flar- ing to an inferior prominence at the squa- mosal suture, suggesting relatively pow- erful development of the masseteric musculature. The nasals are long and nar- row, flaring posteriorly, with the median processes of the f rontals deeply projecting between them. Small, isolated foramina are present in each nasal. The premaxillary- maxillary suture is located in the most an- terior quarter of the snout, just posterior to I2. The maxilla is extensive, incorporat- ing three-quarters of the snout region, and extends to the base of the orbit. The max- illary-jugal suture is oblique and runs above the base of M3. Although upper incisors are not pre- served in the specimen represented by AMNH 109521, roots preserved in the pre- maxillae clearly indicate that only two were itostylops macrodon and associated partial left mandible (uncataloqued IVPP specimen; cast. A iorsal (A), ventral (B), and left lateral (C) views. 17 is Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 present on each side, in contrast to the three known for Palaeostylops iturus and Arctostylops steini. The roots of both in- cisors are subround and approximately equal in size. The base of the upper canine is larger than the roots of the incisors, ap- proximating the base of P1 in size. Whether or not this reflects a notable difference in size between canine and lateral incisor crowns cannot be determined; however, root development is comparable in Pa- laeostylops Hunts, whose anterior teeth nonetheless form an evenly graded series (cf. Matthew, Granger, and Simpson, 1929, p. 12). P1 is single-rooted. Its crown, gen- erally similar to those of corresponding teeth in Palaeostylops and Arctostylops, is buccolingually compressed and bears a sharp mesiodistal crest, which ascends me- dially to the apex of the single cusp. A faint lingual cingulum, not developed into a heel as in Palaeostylops, is present. P2 is double-rooted. It is larger than P1 and structurally similar to it, except that a pro- tocone, smaller than that of Arctostylops and equal to that of Palaeostylops, is de- veloped lingually. Well-defined crests de- scend from this cusp to the anterior and posterior margins of the tooth. The lingual surface of the coronal crest, or ectoloph, is steep and bears well-marked wear facets, as seen in succeeding teeth. P3-4 are suc- cessively larger and more molariform, with more fully developed protocones. As in Pa- laeostylops iturus but in contrast to Arc- tostylops steini, P4 is noticeably smaller than M1. The molars bear sharp, straight ectolophs with well-developed parastyles. On M', the lingual sulcus posterior to the protocone is faint, unlike the condition seen in Palaeostylops. Lingual cingula are well- developed on all upper molars; cusps, vari- able in development, are present on M12. M in tin specimen represented by AMNH L09521 bears two such cusps, one lingual to the protocone and another, larger, pos- terolingual to thai cusp and in a hypoconal position \; is much larger than preceding or succeeding teeth and bears three cusps on the lingua] cingulum. Posterior to the protocone the lingual sulcus is strong, so that the tooth is bilobed. A prominent ac- cessory cusp lies in a median position at the base of the ectoloph, posterior to the parastyle. M3 is generally similar to those of Arctostylops steini and Palaeostylops iturus, except that the lingual cingulum is complete and bears an eminence directly lingual to the protocone. A nearly complete lower dentition is represented in AMNH 21741 from Ga- shato, a left dentary with I^, C, P^, and M^; the last molar bears a moderately damaged talonid. The horizontal ramus is shallow, with a nearly horizontal sym- physis that appears to have been unfused. Small foramina are located below P[ and P4. The three incisors are similar to those of Palaeostylops iturus. \x is spatulate with a rounded point and a median vertical ridge, the crown being less compressed than in Arctostylops steini. I2 is larger and more laterally compressed than Ils with the an- terior part of the crown more expanded and the median vertical ridge better de- veloped. I3 is similar in size and morphol- ogy to I2, except for the presence of an incipient posterior lobe on the median ridge. The canine is subequal in size to I3 and somewhat larger than P,; no diaste- mata separate it from those teeth. The crown of the canine is tricuspid and com- pressed; lingual crests are associated with each cusp. In these respects it generally resembles Palaeostylops iturus rather than Arctostylops steini. The single-rooted P{ is morphologically similar to the canine, although the three coronal cusps are somewhat more distinct. P2 is double-rooted and significantly larger than P,, with the tricuspid pattern clearly defined. P3 is similar to but larger than P2, w ith the protoconid being the tallest cusp. P4 is submolariform, with a serially tricus- pid trigonid and a small, crested heel. The paraconid and metaconid are equal in size, and the protoconid is the tallest cusp. The cristid obliqua attaches somewhat labial to the metaconid. There is no ectocingulid present on any of the lower premolars. ' mmalia • ( tfeUietal cm Figure 6. Stereophotographs of left mandible of Gashatostylops macrodon. AMNH 21 741 The most notable feature of the lower molars is the extremely salient, blade-like, labially-placed cristid obliqua. The molars are morphologically similar to each other, with M2 appearing to be disproportionate- ly larger than preceding and succeeding teeth. The entoconid is expanded into an obliquely-oriented entolophid that con- tacts the cristid obliqua in about the mid- dle of the talonid. On all three molars, the ectocingulid is developed as a distinct an- terolingual ridge at the junction of pro- toconid and cristid obliqua. The protoco- nid is the tallest cusp except on M2, in which the hypoconulid is larger. No directly associated, articulated post- cranial elements are yet known for any of the Arctostylopidae, but proximal ankle bones may now be referred to Palaeosty- lops iturus and Gashatostylops macrodon with little doubt. These species are by a considerable margin the most abundant taxa known from Gashato. Isolated astrag- ali and calcanea, of appropriate size for P. iturus and G. macrodon, occur there in the same relative abundances as dental re- mains of these species. Furthermore, an astragalus was found lodged within the cranial cavity of a specimen from Bayan Ulan referred to G. macrodon (see below). This astragalus, for which association is reasonably inferred, resembles to the point of identity the isolated specimens from Gashato believed on the basis <>l m/<- a\h\ relative abundance to belong to Gasha- tostylops macrodon. In known respects the ankle of Palaeostylops iturus is similai to that of G. macrodon. and it is therefore not described separately. Descriptive ter- minology follows that of Cifelli h VMNH 21726, a right astragalus from Gashato. the astragalai body is mediolateralh compressed, with nearly vertical sides: there is little or no development of a fibular shell on the lat- eral side. The body is more or less i ylin- drical, with the tibial trochlea marked b) a median groove and well-defined raised borders. An astragalar foramen is appar- ently lacking; a pit is present on the tibial trochlea of AMNH 21726 but appears to have been caused !>\ diageneti< corrosion of the fossil (as on the other side ol the same specimen; such pitting is common on fossils from Gashato The neck is ol mod- erate length but is notabl) constricted, so 20 Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 that the head is clearly demarcated. The head itself is subround with, however, the navicular facet developed as a flattened band that does not extend onto its sides. The navicular facet extends far superiorly, onto the dorsal side of the bone, and is developed so that movement between as- tragalus and navicular would have been subparallel rather than highly oblique to that between astragalus and tibia. There is no observable separate facet for the cu- boid. The tarsus might thus tentatively be regarded as "serial" (see discussion by Os- born, 1889), although this cannot be de- finitively ascertained until a well-pre- served, articulated foot is discovered. The sustentacular facet is unremarkable, ex- cept that it is somewhat larger and better developed distally than it is in Protun- gulatum. The ectal facet is very steeply inclined, and the interarticular sulcus sep- arating the two facets is deep. The calcaneus, as represented by AMNH 21742 (complete left calcaneus and right calcaneus lacking the tuber, almost cer- tainly not from the same individual), is notable in having a short neck (that part anterior to the astragalocalcaneal facets) relative to the tuber. The ectal prominence is dominated by a very strongly developed fibular facet, which forms a broad, antero- posteriorly oriented, semicylindrical sur- face. Medial to this lies the ectal facet, which is strongly inclined with respect to the fibular facet. The sustentaculum is un- usual in lying at or near the distal end of the bone; a prominent "beak" is developed on the superior distomedial corner of the bone. The cuboid facet is developed at a moderate angle with respect to the long asix of the calcaneus. Comparisons to other taxa are deferred until the discussion. Sinostylops Tang and Yan, 1976, p. 91 Type Species. Sinostylops promissus Tang and Yan, l()76, p. 92. Included Species. The type only. Distribution. Kate Paleocene (fide Li and Ting, 1983), Asia. Diagnosis. Primitive arctostylopids dif- fering from Asiostylops in having higher- crowned molars and a metaconid on P?3. Distinct from advanced genera such as Arctostylops in retaining a paracristid on the lower molars. Similar to Bothriostylops in having the cristid obliqua attaching to the trigonid of lower molars in a lingual position, but differs from that genus in having higher-crowned molars and a more slender, elongate P3. Sinostylops promissus Tang and Yan, 1976, p. 92 Holotype. IVPP V4263, right mandibular ramus with eight partial or complete teeth. Hypodigm. The type only. Horizon and Locality. IVPP locality 71017, Dou-mu Formation, Anhui Prov- ince, People's Republic of China; late Pa- leocene. Diagnosis. As for the genus. With the removal of referred species "Sinostylops" progressus to Bothriosty- lops, the concept and affinities of Sino- stylops become problematic. The identi- ties of the eight teeth in the holotype and only specimen of Sinostylops promissus are open to doubt. Because the third from the last tooth is remarkably low-crowned and long, unlike either preceding or suc- ceeding teeth, we believe it to be decid- uous. The penultimate tooth, although much smaller than the ultimate, is mor- phologically similar to it; both are badly damaged but apparently were bicrescen- tic, which is not the case for the more anterior teeth. We therefore believe the teeth in this specimen to be I3-P3, dP4, and Mi.2) although other interpretations are possible. Available materials of Sinosty- lops promissus and Bothriostylops pro- gressus suggest further differences be- tween the species beyond those listed in the diagnoses, but because of the uncertain identities of the teeth in IVPP 4263 and because of postmortem damage to that specimen, we are unable to evaluate the significance of these differences. Arctostylopids (1 ,,, .< ifeUtetal fibular facet B sustentaculum fibular facet cuboid facet sustentaculum D sustentacular facet ectal facet navicular facet mm tibial trochlea Figure 7. Right astragalus (unnumbered IVPP specimen associated with rostrum of skull, a cast of which b numbered AMNH 109521) and left calcaneus (AMNH 21742) of Gashatostylops macrodon. A. B. C: calcaneus in distal, dorsal, and planter views. respectively; D, E, F: astragalus in plantar, dorsal, and distal views, respectively. .->.-> Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 Bothriostylops Zheng and Huang, 1986, p. 121 Type Species. Bothriostylops notios Zheng and Huang, 1986, p. 122. Referred Species. The type, and Both- riostylops progressus (Tang and Yan, 1976, p. 92). Distribution. Late Paleocene, Asia. Diagnosis. Primitive arctostylopids with brachydont teeth, differing from Asio- stylops, which they generally resemble, in having a crescentic P4 talonid; and from all known genera in having M3 with an elongate talonid, the hypoconulid forming a distinct lobe. A number of other characters were list- ed in the diagnosis and description of the genus (Zheng and Huang, 1986). Of these, the presence of a deep median labial groove and convex labial wall on lower molars were cited as important similarities to Asiostylops. We see no distinction of arc- tostylopid genera on this basis but, lacking access to the original specimens (especially the type of Bothriostylops notios), we de- fer to Zheng and Huang (1986). Nonethe- less, we observe that a deep median ex- ternal groove is present on lower molars of Sinostylops promissus. Even on the ba- sis of the single, enigmatic specimen avail- able, it is clear that this latter species is rather divergent and not obviously con- generic with other known taxa. Bothriostylops notios Zheng and Huang, 1986, p. 122 Holotype. IVPP V7642, portion of left mandible with P4-H, Referred Specimens. The type only. Horizon and Locality. Wang-wu Mem- ber, Chi-jiang Formation; late Paleocene. North of Zhulin Hill, Dayu County, Jiang- \i Province, People's Republic of China (cited from Zheng and Huang, 1986). Diagnosis. Cheek-teeth lower-crowned than in R. progressus. Entolophid of lower molars not so well-developed as in that sp< • nd, at least on \1 , incomplete; trigonid ol lower molars more open lin- gually, with the paracristid less truncated, than in B. progressus. Bothriostylops progressus (Tang and Yan, 1976) Figures 8, 9 Sinostylops progressus Tang and Yan, 1976, p. 92 Bothriostylops progressus Zheng and Huang, 1986, p. 127 Holotype. IVPP V4264.1, fragment of right mandib- ular ramus with M2. Beferred Specimens. The type, and IVPP 4264.2, right mandible fragment with worn M2_3; 4264.3, right mandible fragment with P3_4; 4264.4, right mandible fragment with P34; 4264.5, right mandible fragment with M2 and with broken M3; 4264.6, right maxillary fragment with bro- ken M1 and with M2-3 well-worn. Horizon and Locality. IVPP locality 71071, Shuang-ta-si Group, Anhui Prov- ince, People's Republic of China; late Pa- leocene (Li and Ting, 1983) or early Eocene (Zheng and Huang, 1986). Diagnosis. Cheek-teeth higher-crowned than in B. notios. Entolophid of lower mo- lars complete and more fully developed than in that species; trigonid of lower mo- lars more compressed, with paracristid more truncated, than in B. notios. Anatolosty lops Zhai, 1978, p. 109 Anatostylops, Schaff, 1985, p. 593 Type Species. Anatolostylops dubius Zhai, 1978, p. 109. Included Species. The type only. Distribution. Late early Eocene or early middle Eocene (fide Li and Ting, 1983) or, perhaps, Oligocene (fide Zhai, personal communication); Asia. Diagnosis. Differs from all other genera, excepting an unnamed form, in having higher-crowned cheek-teeth; ectoloph of upper molars elongate, smooth and fea- tureless, with a large parastyle but no para- stylar fold. Pre- and postprotocristae sa- lient, enclosing a fossette that persists through more advanced wear than in other forms. Sulcus on lingual side of M2 crown ARCTosTYLOPins (Mammalia) • ( :„■//, ei at. not so broad as in Arctostylops or Palaeo- stylops. Differs from a closely similar un- named genus and species in having a lin- gual division of M2, and in lacking the great anteroposterior expansion of the ec- toloph and the strong development of the postcingulum seen on upper molars of that genus. Anatolostylops dubius Zhai, 1978, p. 109 Figure 8 Holotype. IVPP V4357, fragment of left maxilla with Hypodigm. The type only. Horizon and Locality. Shi-san-jian-fang Formation, Turpan Basin, Xin-jiang Prov- ince, People's Republic of China; Eocene or Oligocene (see above). Diagnosis. As for the genus. Asiostylops Zheng, 1979, p. 388 Type Species. Asiostylops spanios Zheng, 1979, p. 388 Included Species. The type only. Distribution. Late Paleocene (fide Li and Ting, 1983), Asia. Diagnosis. Distinct from all other arc- tostylopid genera in the more transverse P2 3, with a lesser development of the pro- toconal region; upper molars lacking a pos- terolingual cusp or other secondary coro- nal complications; metaconid lacking on P3. Lower molars primitive in retaining the paracristid, as in Bothriostylops and Si- nostylops but not other genera; cristid ob- liqua attaching to trigonid in a median position. Entolophid feebly developed and transversely oriented. Asiostylops spanios Zheng, 1979, p. 388 Figures 8, 9 Holotype. IVPP V5042, cranium and associated left mandible. Hypodigm. The type only. Horizon and Locality. IVPP locality 73039, Lan-ni-kong Member, Chi-jiang Formation, Jiang-xi Province, People's Re- public of China. Diagnosis. As for the genus. Kazachostylops Nesov, 1987. p. 212 Type Species. Kazachostylops occid* r». talis Nesov, 1987, p 212 Included Spr.-i, . The type only Distribution. Late Paleocene western Asia. Diagnosis (from Nesov 1987, p. 21 Small arctostylopids with long, tall pai cristid on lower molars; premetacristid and postmetacristid reduced to absent I ■ lophidof VI, ,long,nearl) transverse and joined with the talonid loph. ( Irestsol Hi- lar teeth form practically uninterrupted cutting edges. Lacking access to the two, relatively good specimens of the type and only species ol Kazachostylops, we defer to Neso> s briel diagnosis of the genus, and mint it from the detailed comparisons and discussion presented below. From the figures, Ka- zachostylops appears to be rather similai to Bothriostylops and, perhaps, Sinosty- lops, particularly in the elongate, well-de- veloped M3, the strong, crescentic para- cristid, and in the lingual attachment ol cristid obliqua to trigonid (i.e., at the meta- conid). Kazachostylops occidentalis Nesov. 1987. p. 212 Holot ype. Specimen number Hi 12455 indicated b) Nesov (1987) as being deposited in the Ts.N.1 G.F Museum, Kazakhstan, Dzhilu.i I ssli , onsisting ol a right dentary with C, F, ,. and \1 Hypodigm. The type, and at least one more dentulous jaw fragment figured l>\ Nesov (1987), number 12 L2455, consist- ing of a right maxilla with P1 to M Horizon and Locality. Marginal marine deposits of the Pretashkenl Svita, late Pa- leocene; site TDA-2. Kazakhstan, Dzhilga USSR (fide Nesox. L987, p. 212 Diagnosis. As for the genus Arctostylopidae?, incertae sedis Allostylops Zheng, 1979, p. 391 Type Species Allostylops periconatus Zheng, 1979, p. 391 Included Species.The type onl) 2 I Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 "^TmJllfflmjmrr'Tnr. Figure 8. Comparative series of arctostylopid (A-H) and primitive notoungulate (I) upper dentitions. Teeth standardized to size and reversed where necessary. A, Arctostylops steini (MCZ 20004); B, Palaeostylops iturus (AMNH 22143); C, Gashatostylops macrodon (cast, AMNH 1 09521 ); D, Anatolostylops dibius (IVPP V4357); E, undescribed genus and species (unnumbered IVPP specimen); F, Bothhostylops progressus (IVPP V4264.6); G, Allostylops periconatus (IVPP V5043); H, Asiostylops spanios (IVPP V5042); I. Peripantostylops minutus (AMNH 28494). Distribution. Late Paleocene {fide Li and Ting, 1983), Asia. Diagnosis. Generally primitive ?arcto- stylopids similar to Asiostylops spanios in the low -crowned cheek-teeth, the small size <>f P\ and the presence of a paracone fold on the ectoloph of at least some upper mo- lars, hut differing from that species in hav- ing a hypocone on M2. Differs from ad- vanced arctostylopids (Palaeostylops, Arctostylops, Gashatostylops, Anatolo- stylops) in having lower-crowned cheek- teeth, a smaller P3, smaller upper molar parastyles, and a broadly expanded pos- terior cingulum on M1. Allostylops is dis- tinct from all forms in the family save ;omparative series of arctostylopid (A-E) and primitive notoungulate (F) lower dentitions. Teeth standardized to size j reversed where necessary. A. Arctostylops steini (MCZ 20004); B, Palaeostylops iturus (AMNH 20414); C, Gashatostylops rodon(AMNH 21741); D, Bothhostylops progressus (P^, IVPP V4264.4; M2, IVPP V4264.1; M3 outline, IVPP V4264.2)- E. Asiostylops spanios (IVPP V5042); F, Peripantostylops minutus (AMNH 28494). 25 Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 Figure 1 0. Stereophotographs of P3-M3 of undescribed genus and species of Arctostylopidae (unnumbered IVPP specimen). Gashatostylops in having a prominent pericone anterolingual to the protocone on upper molars. Allostylops periconatus Zheng, 1979, p. 391 Holotype. IVPP V5043, badly preserved rostral por- tion of cranium. Hypodigm. The type only. Horizon and Locality. IVPP locality 73041, Wang-wu Member, Chi-Jiang For- mation, Jiang-xi Province, People's Re- public of China; late Paleocene. Diagnosis. As for the genus. Arctostylopidae, genus and species indet. A In addition to the foregoing previously described species, an unnamed arctosty- lopid is represented by an upper dentition collected from the Yan-ma-tou Formation, Hunan Province, People's Republic of ( !hina. While full description of this species is in progress, we briefly note some of its morphological features here in order to facilitate comparison among other mem- bers of the family and to aid in assessing their relationships. The taxou in question is a small, dentally dvanced arctostylopid similar to Anato- lostylops in having high-crowned cheek- tt i tli and a smooth ectoloph on the upper ii ilars, but it differs from this and all other genera in the great anteroposterior expan- sion of the ectoloph crest on P3-M3 and in the strong development of the postcingu- lum on M1'2. Indeed, the ectolophs of the upper cheek-teeth are so strongly devel- oped that the rest of each tooth appears by comparison to have been constructed as an afterthought. A lingual division of M1-2, seen in all other genera except Asio- stylops, is lacking. The second upper mo- lar is notably larger than the first. The single specimen representing this species was plotted into a measured section (Unit 21 of Zhu-chen, 1986) of beds re- ported to be of Cretaceous age. The basis for this surprising age determination is not entirely clear, but it seems to involve fossil remains believed to be dinosaur eggs (list- ed as Elongatoolithus and other taxa) which, apparently, bracket the arctosty- lopid specimen. Other fossils from this sec- tion are listed merely as "animal bones" or "animal teeth," and are therefore of little help in age determination. However, a mammalian axis vertebra is larger than that of any Chinese Paleocene mammal and would be totally out of place in the Cretaceous, as would a large anterior tooth of some ungulate-sized mammal. We be- lieve on this basis that the locality is much younger than Cretaceous, perhaps even Eocene in age, whether dinosaurs were present or not. Indeed, the advanced mor- ARCTOSH 4AMMAL1 ,,/ :: phology of the small, distinctive arctosty- (strong hiah ,,-„.. lopid from this site is suggestive of the cone nectingpara- Eocene or possibly Oligofene Anatolo- ^^X^S^S^T^1^ stylops dubius. Hut'/ duC€ * stylar prominences; blade-like Arctostylopidae, genus and species serially tricuspidate I' . with notches a indet. B aratmg the cusps; l\ with a trigonid i > Another arctosty lopid, which we have Kfffi^t(nX^ ''IT*11' not seen, occurs in the late Paleocene Da- lower m(lrs i ! ^ ' ' tang Member of the Nung-shan Forma- ZZoni i , , dian S 7^ 'T"" .. tvt . i • /-. i pciidtuiiiu in a media )os turn and i i , turn, Nan-xiong basin Guang-dong, Peo- onid consisting of cristid obli „ land ' , ,' pies Republic of China. The animal is cristid united into a ,„„„ s' , L?aandeTinWl983 fl3) *"*" ^ ^ ^^ and *»** '"' ^ Liand ling (1983, p. 13). tinct. The entoconid of lower molars is iso- lated from the postcristid and developed COMPARATIVE DENTAL MORPHOLOGY int° a f^?tf.trfnS!en^ '0p!' " ^ extends OF THE ARCTOSTYLOPIDAE anterolabially to the tal dcrescenl Man) °r these features are shared b\ pKMiinal.K Review of dental variations among the unrelated groups of mammals, but the se- Arctostylopidae and assessment of the rel- rially tricuspid anterior lower premolars ative primitiveness of various character and the transversely developed entoconid states is based on comparison with an un- (entolophid) of the lower molars are ratlin gulate morphotype as represented by Prot- distinctive characters. ungulatum and various comparable oxy- Arctostylops, Palaeostylops, and Ga- claenine Arctocyonidae (Cifelli, 1983a). shatostylops are distinctively more spe Some of the features that are represented cialized. Advanced characters of th< in available materials of the known species three genera with respect to Asiostylops are summarized in Table 4. The most include higher-crowned posterior premo- primitive arctostylopid for which good lars and molars; an expanded protocone on materials are available is unquestionably P3; upper molars with a high, flat ectoloph Asiostylops spanios Zheng, 1979. Zheng wall including parastylar and metastylar (1979) referred Asiostylops to the Notoun- folds only (Asiostylops has a distinct para- gulata based on the biselenodont lower cone fold); M2, at least, is bifid linguall) molars, with shortened trigonid and loph- with high pre- and postprotoeristae that odont entoconid, and on the upper molar enclose a very transient trigon fossette l>nt ectoloph, with parastyle developed. He which are rapidly reduced l>\ hcaw weai considered Asiostylops to be primitive The lower molars of these three genera are within the order because the cheek-teeth distinctive in a number of respects, such are low-crowned, the premolars are not as: 1) the presence of a salient, pillar-like molarized (in particular, P4 lacks an ento- ectocingulid with a wear surface dea end- conid), the lower molars have a pro- ing along its face; 2) the reduction ol nounced paraconid, and the upper molars trigonid by loss of the paracristid 3) the lack the secondary coronal complications presence of a high, shearing talonid seen in Henricosbornia and more ad- cent (cristid obliqua), which joins the tri- vanced South American notoungulates. gonid labial to position ol the prota onid; Compared to an ungulate morphotype and 4) the strongK developed, oblique en- represented by Protungalatum, Asiosty- tolophid. All three genera have an ante- fops spanios has a greater development of riorly placed P, paraconid, unlike \sio the protocone on P3 \ with a metacone on stylops. The polarities ol some feati those teeth; upper molars with an ectoloph P< are uncertain. Arctos Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 Palaeostylops, Gashatostylops, and most other Arctostylopidae in the presence of a crescentic P., talonid loph, and differs from Gashatostylops macrodon (but not Pa- laeostylops iturus and several other species) in the lingual placement of its P4 meta- conid. Arctostylops steini is probably au- tapomorphous in having a stronger ecto- cingulid on P34, a stronger lingual rib on C,, a prominent heel on I1, and a slightly larger protocone on P2. Palaeostylops and Gashatostylops appear to be derived with respect to Arctostylops in the lesser dif- ferentiation of C1, the lack of paracone folds on the ectolophs of P34, and the pres- ence of a shearing notch on P4. Palaeo- stylops and Gashatostylops differ from Arctostylops also in the more quadrate, less transverse nature of M12 and in the fact that the sulcus between the two in- ternal cusps is better developed, at least on M1. The P4 cusps in Gashatostylops ma- crodon are more or less anteroposteriorly aligned, as with the more anterior pre- molars of all genera; the talonid crest is a straight, bladelike structure. G. macrodon is also distinctive in that the upper and lower second molars are greatly enlarged, in the variable development of one or more cuspules on the lingual cingulum, and in the reduction or absence of a lingual suclus on M1 (further distinctions are given in the diagnoses provided above). Thus, Arcto- stylops, Gashatostylops, and Palaeosty- lops share presumed synapomorphies with respect to Asiostylops. Within this clade of advanced genera, there is some evi- dence to suggest that Gashatostylops and Palaeostylops shared a more recent com- mon ancestor than either did with Arcto- stylops. Because of uncertainty in mor- phocline of several features, the possibility of lineal relationships between any of the included species cannot be evaluated. The remaining species of Arctostylopi- dae are known from less complete mate- ils and there is. accordingly, some un- certainty as to various character states. Although rather primitive, the two species ol Bothriostylops are unique among arc- tostylopids in having an elongate M3 in which the hypoconulid forms a separate lobe. (This also appears to be true of lower molars belonging to Kazachostylops occi- dentalis, which we have not examined firsthand. We are unable to consider the species further here, but note that the above-mentioned feature and several oth- er lower molar characters suggest a close relationship to Bothriostylops spp.) We as- sume, for the purpose of comparison, that these two species form an exclusive unit within the family. Thus conceived, Both- riostylops is, in several respects, interme- diate beteen Asiostylops on the one hand and advanced arctostylopids (Arctosty- lops, Gashatostylops, Palaeostylops) on the other. As in Asiostylops, Palaeostylops, and Arctostylops, the metaconid on P4 is lin- gually placed (we are uncertain of the con- dition in B. notios). The talonid crest of that tooth is curved in Bothriostylops spp, although not so strongly as in Arctostylops. The lower molar trigonids of B. progressus are anteroposteriorly compressed, as in the derived genera, but unlike those forms, part of the paracristid remains, as in Asio- stylops. In B. notios, the trigonid retains a more open arrangement, with the para- cristid little reduced. The cristid obliqua attaches to the trigonid at a lingual posi- tion, near the apex of the metaconid, un- like either Asiostylops on the one hand or Palaeostylops/ Arctostylops on the other. The ectocingulid is feebly developed and not expanded into an occlusal structure. The entolophid varies from well-devel- oped (B. progressus) , as in the advanced forms, to weak and incomplete (B. notios). A partial, very worn, upper molar series is available for Bothriostylops progressus, but it adds little to knowledge of the species. The ectoloph appears to have been high; as far as can be determined, paracone and metastylar folds are lacking although a parastylar fold is well-developed. M1 has a sulcus separating two lingual cusps; this appears not to have been true of M2, which is triangular in outline (as with Asiosty- lops), but excessive wear has obscured de- Arctostylopids (Mammalia) • Cifelli ei al. < D 0 J H C/3 o H U < ac H O z o < en Z o en E < o o cc w H O <: cc < a o j < H Z w Q j CQ < o c wi O 5H O 3 S3 <■> u fi C bC cd O C a c 0-- C»- fL. C "TV ■" — bC ■" — yo "3 cb O - b "_ _ CD "■*- CO CO — ~ CO ft □ -j £. . _2 _2 ._£_:;- Cl. fL. cl. o-. O-. CT-. o_. O, > o3 jj > o3 ~3 B** Sjb > bl 2i = bC c-l. ct-. £V. id .2; o- - . 2 CD ■" bO — 3 uT 3 as tt "> c — — W — £ <^ _ y ' X . _o . o •- • a3 > as f>-. u — > o. '- 2 5 C « -S O __ cd SP cd s _o .5 3= _q fc .5 -C ai « 'S aj Rii i t/> — M-. tii- cS .5 — bC cd c o . i _Q -O co 03 03 C _c a, „ .SP_o Cb_3 03 ID 3 H co S> o3 3 _b 7 £_ C u ''I • o-. f>-. o-' c 03 OJ 3 CO CD bC Lri 3 tt 33 - f- £*-■ CD « *< - a 3 3 3 •£ e CD CD ^2 CD j- crt en ^ co CD O. 03 -3 — ? T3 33 rt -a "O = 3 3 -3 ■- a s 3 O O 03 l-< 03 a 8 © o O a 0- 0- , S;0)3 CD 3 bO tr-3 "K _Q s t>c2 _ -r 2 1) _ o3 i- »> 2r t o 03 -0 o r, O I- CD O o — V SSSS^WUOh 30 Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 tails of crown morphology. M2 may have been slightly larger than the adjacent teeth, but it is not greatly enlarged as in Ga- shatostylops macrodon. As for Sinostylops promissus Tang and Yan, 1976, poor preservation of the type and only known specimen leaves various character states open to question. It cannot be determined if an entolophid was pres- ent on Ml2. It appears that a trigonid cres- cent was retained, as in Asiostylops, and the cristid obliqua attaches to the trigonid at the metaconid, as in Bothriostylops spp. The antemolariform teeth form a graded series and are long, narrow, and bladelike, especially dP4. The premolars are serially tricuspid, with a straight, crested heel. Si- nostylops promissus lacks advanced fea- tures of the lower molars seen in Palaeo- stylops and Arctostylops. The morphology of the premolars would seem to indicate pertinence to the Arctostylopidae; within the family, Sinostylops promissus is sim- ilar only to Bothriostylops spp in the lin- gual attachment of cristid obliqua to tri- gonid. Allostylops periconatus Zheng, 1979, about which little can be said, is repre- sented by the rostral part of a skull with the dentition very poorly preserved. The upper molars resemble those of Asiosty- lops, and are therefore presumably prim- itive, in lacking an enlarged parastyle and in retaining paracone and metacone folds on the ectoloph. There was, apparently, no posterointernal cusp on M1-2; a prominent anterolingual cusp (pericone) is present on the lingual cingulum, as is variably present on upper molars of Gashatostylops mac- rodon. The posterior cingulum of M1-2 is broadly expanded, so that the molars are subquadrate in occlusal aspect. The den- tition as preserved gives little indication of affinity to this group, and the position of Allostylops is therefore indeterminate. Anatolostylops dubius Zhai, 1978, known from M2-\ is clearly a rather spe- i ialized Form and may be significantly younger than the other genera. As in Pa- laeostylops, Arctostylops, Gashatostylops, and Bothriostylops, the ectoloph is high and lacks a paracone fold; unlike those forms, the ectoloph is otherwise feature- less, lacking a parastylar fold or basal bulges in the regions of parastyle and metastyle. The lingual coronal crests (pre- and post- protocristae) are strong and enclose a fos- sette that probably persists into a fairly advanced stage of wear. The sulcus be- tween the lingual cusps on M2 is not so deep as in Palaeostylops or Gashatosty- lops but, as in those genera, it probably persists to advanced wear. A lingual cin- gulum is weak or lacking on M3, as in Both- riostylops progressus; as in Gashatosty- lops macrodon, M2 is considerably larger than M3. Anatolostylops is most closely similar to the unnamed genus and species, with which it shares several derived char- acters not found in other Arctostylopidae. The ectoloph is anteroposteriorly elongate, with labial plications reduced or lost. The lingual division of upper molars is poorly marked in Anatolostylops and absent in the unnamed form; because these genera otherwise appear to be closely related to forms in which it is well-developed (e.g., Palaeostylops), we believe this to repre- sent reduction or loss rather than retention of a primitive condition (as in Asiostylops) . The cheek-teeth of the undescribed genus and Anatolostylops are higher-crowned than in other genera, and the pre- and postprotocristae better developed, enclos- ing a more persistent fossette than in other members of the family. Although Gasha- tostylops is autapomorphous in several re- spects, notably in the development of ac- cessory cuspules on the lingual cingulum and base of the ectoloph of upper molars, it is similar to Anatolostylops and the un- described form in several other respects. These include a reduction of the lingual sulcus on at least the first tooth of the upper molar series and the great size of the sec- ond molar relative to that of adjacent teeth. Among advanced Arctostylopidae, Ana- tolostylops is divergent in having double opposition of upper to lower teeth, as in- dicated by the presence of a distinct wear facet in the mesostylar area of the upper molar (this would correspond to a facet Arctostylopids (Mammalia) • Cifelli ei al. 3] Figure 1 1 . Hypothesized relationships among the Arctostylopidae. Characters at nodes (see Table 4): 1 ) metaconid added to P3, pseudohypocone on at least one upper molar, upper molar paracone fold lost, ectocingulid developed on lower molars, ectocingulid developed on P4, P4 talonid curved, upper molar parastyle enlarged; 2) lower molar cristid obliqua attaches Imgually to rear of trigonid; 3) M3 elongate? (condition unknown in S. promissus), 4) lower molar entolophid well developed, lower molar cristid obliqua attaches labially to rear of trigonid, lower molar ectocingulid strong and pillarlike, P4 paraconid shifted labiaily. lower molar paracristid lost; 5) shearing notch developed on P4 talonid, P3^ paracone fold lost, canines lesser differentiated. 6) second molars enlarged, pseudohypocone lost on M1?; 7) upper molar protocristae salient, M2 pseudohypocone reduced9, parastyle fold on ectoloph of posterior upper cheek teeth lost, ectoloph of upper molars anteropostenorly elongate, cheek teeth very high crowned. anterior to the protoconid on the lower molars, which are not known for Anato- lostylops). This facet is lacking in Arcto- stylops, Palaeostylops, and Gashatosty- lops, which apparently had singly opposing upper and lower cheek teeth. An hypothesis of interrelationships of the Arctostylopidae is given in Figure ll7 (Ka- zachostylops occidentalis, which we have not examined first-hand, and Allostylops 7 The absence of a chronologic dimension is due to uncertainties of relative age, not our lack of appre- ciation for this consideration. periconatus, poorly known and of doubt- ful affinities, have been omitted from this phylogeny). Asiostijlojis spanios is the most primitive taxon known and i^ considered to represent the sister group of all remain- ing taxa. Bothriostylops spp, unique in .it least one character (the presence o\ .i hy- poconulid lobe on \1>. generall) resemble Asiostylops in their retention "t primitive features, but nonetheless appear t" share several derived features with the remain- ing taxa. Among these are the presence ol a P3 metaconid, a curved P< talonid. a lin- gual division of M1, and the loss of the Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 paracone fold and the presence of a large with an ungulate morphotype, of which narastvle on the upper molars. Sinostylops Protungulatum is a good approximation, nromissus (poorly known and lacking all these notoungulates of the earliest fau- much of the most' diagnostic morphology nas share a number of dental speciahza- i„ the type and only specimen) is similar tions (Figs. 81, 9F). The posterior upper onlv to' Bothriostylops spp in its lingual premolars (P34) are somewhat molanzed, cristid obliqua-trigonid attachment; it is with large protocones supporting antenor very tentatively regarded as the sister tax- and posterior lingual cingula and trigonal on of Bothriostylops spp. The remaining crests; the teeth are dominated labially by Vrctostylopidae clearly are united by de- a prominent paracone, which is separated rived morphology not found in Asiostylops from the also well-developed parastyle and or Bothriostylops. These features include metastyle. A metacone, as far as is known mainly specializations of the lower cheek- does not develop on upper premolars of teeth, such as the labial attachment of the notoungulates. Illustrated specimens of cristid obliqua, the presence of a pillar- Henricosbornia lophodonta (Simpson, like ectocingulid, and the loss of the para- 1948, figure 53) and Oldfieldthomasia de- cristid. Among advanced genera, Arcto- bilitata (Simpson, 1967, plate 5) have stylops appears to be the most primitive, metacones on the teeth indicated to be P\ lacking specializations such as a shearing but comparison with other materials be- notch on ?4, found in Palaeostylops and longing to these species indicate that the Gashatostylops. Within the group formed teeth in question are probably deciduous. b\ the remaining genera, the undescribed The upper molars bear a strong ectoloph form and Anatolostylops possess several whose labial wall is marked by sulci sep- svnapomorphies (mainly features related arating parastyle, paracone, and meta- to the hvpertrophied ectoloph of upper cone. M1"2 are quadrate in occlusal view, molars) and both share with Gashatosty- with a posterolingual cusp (hypocone) sep- lops an enlarged second molar. arated from the protocone by a sulcus. M3 does not develop a hypocone, but variants THE NOTOUNGULATA OF SOUTH among even primitive taxa may show AMERICA strong development of the cingulum in this The early Tertiary Notoungulata of region. The crest linking protocone to South America have been fully reviewed paracone (preprotocrista) is strong and is by Simpson (1948, 1967). Additions to developed into a protoloph; on the first two knowledge since publication of these molars, at least, and variably on M\ a monographs have been principally the Ita- metaloph joins hypocone and metacone boraian to Casamayoran notoungulates of (Fig. 12). The metaconule of upper molars Itaborai, Brazil (Paula Couto, 1952, 1954, is expanded anterolabially into the trigon 1978) and of northwestern Argentina basin as a crochet; various other cuspules (Bond, 1981; Pascual, Vucetich, and Fer- and crests characterize this part of most nandez, 1978; Vucetich, 1980). As recog- notoungulate upper molars (see Patterson, nized by Simpson, the major advanced no- 1934; Simpson, 1948). Cingula are present toungulate suborders Toxodonta and anteriorly and posteriorly but not lingual- T\ potheria (including Hegetotheria; see ly. The posterior lower premolars (P3_4) are ( lifelli, 1985a) were differentiated by the molarized (P3 somewhat less than P4): the late Paleocene, \\ ith 5 families collectively trigonid is crescentic, with crests directed represented. Simpson grouped two other anteriorly and posterolingually from the families of the earliest faunas (Riochican protoconid; the talonid is much shorter than ', the Henricosborniidae the trigonid and also bears a crescentic into his paraphyletic crest. The construction of the lower molar . When compared trigonids is extraordinary, and the homol- Arctostylopids (Mammalia) • Cifelli et al. ogies of some parts are open to question. A crest (paracristid?), variable in length, extends anteriorly or anterolingually from the protoconid; a low anterior crest or cin- gulum, on the anterior face of the tooth, may connect with this in heavy wear so that the paracristid (?) appears to run to the lingual margin of the tooth. The great- est variation occurs in the region of the metaconid. That cusp may be anteropos- terior^ expanded (Henricosborniidae, some Oldfieldthomasiidae), bearing an an- terolabial-posterolingually directed crest. Another variant involves the presence of an anterior accessory cusp, which some- times bears the appearance of a paraconid that has lost the paracristid connecting it to the protoconid (most notably in Isotem- nidae but also in some Oldfieldthomasi- idae). Notostylopids are characterized by an accessory cusp on the crest linking pro- toconid to metaconid (protocristid), so that this crest is serially tricuspid. Marshall, de Muizon, and Sige (1983) propose homol- ogies for these trigonid structures, which they argue are variations about a basic no- toungulate pattern that included a pre- and postmetastylid. The talonid consists, in its simplest form (Henricosborniidae), of a crescent (cristid obliqua and postcristid) uniting hypoconid and hypoconulid, which nonetheless are retained as distinct, cusp- like entities. The entoconid is developed transversely (entolophid) and, in advanced forms, joins the postcristid anterior to the hypoconulid. The most primitive condi- tion of this feature is seen in henricosbor- niids such as Henricosbornia itself. The entolophid is incompletely developed, pos- terobucally oriented, and is somewhat more separated from the hypoconulid on the posteriormost molar of Henricosborniidae. On M,, however, this crest runs labially to the hypoconulid or to a point just anterior to that cusp, and it therefore appears that the entolophid is homologous to the crest connecting entoconid and hypoconulid (a portion of the postcristid), and becomes distinct as a separate loph by migrating anteriorly. DISCUSSION In the original description ol thespei ies, Matthew (1915) referred Arctostylops steini to the order "Entelonychia" and. within that group, plated the species w ith some doubt in the Isotemnidae. At thai time, "notoungulate" t<> many students (see, e.g., the influential classifications oi Gregory, 1910; Osborn, L910; and Scotl 1904) was equivalent to "indigenous South American ungulate," and did not explic- itly refer to that group in the sense it is defined today8. "Entelonychia" was a sub- order proposed by Ameghino 1 1 89 i to in- clude the aberrant, clawed Homalodo- therium (a Santacrucian, mid-Miocene form shown by Patterson, L936, to be tox- odont-like in the construction of its eat region and since universally placed in the Toxodonta, a suborder of the Notoungu- lata) within the "Aneylopoda," thus unit- ing it with the similarly clawed chalico- theres of Holarctic faunas. Ameghino had abandoned the use of the term "Entelon- ychia" by the time of his final (1906) clas- sification, but by this time had placed oth- er notoungulate families (Isotemnidae and Leontiniidae, both currently recognized as belonging to the Toxodonta) with the Homalodotheriidae in the "Aneylopoda." The dentition of members of all these fam- ilies are relatively primitive within the No- toungulata. Thus, later workers ignored Ameghino's reference of these and other notoungulates to Holarctic groups, and in- stead resurrected his term "Entelonychia to include generally primitive notoungu- lates. (Scott, 1913, for instance, placed the Notostylopidae under this heading. K\ the time of Matthew's (1915) writing, "Entel- onychia" referred to primitive notoun gulate mammals; then, as now, the Isotem- nidae were considered to be basal meml of the South American notoungulate ra diations (although the henricosborniids are generally acknowledged to be somewhat more primith -The concept of the Notoungulata now current had, however, been made clear b) Roth, I 3 t Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 Matthew and Granger (1925) recog- 1967), was that the Henricosborniidae, then nized that Palaeostylops iturus was strong- known only from the Riochican and early ly specialized in having high-crowned Casamayoran (Cifelli, 1985b), or putative cheek-teeth with well-developed shearing late Paleocene and early Eocene (Marshall, surfaces, and in having reduced lower mo- 1985; Marshall, Hoffstetter, and Pascual, lartrigonids. In this respect, they indicated 1983), represent the most primitive of that (pp. 4-5), "it may be regarded as an- known Notoungulata. By this interpreta- cestral to Arctostylops and through that tion, the order arose in South America from genus to some of the South American the same "ungulate" stock which gave rise Eocene Notoungulata (e.g., Leontinia, also to the other groups of indigenous South Notostylops, etc.) but to the latter only in American ungulates. Migration of a prim- a broad way, as no one of the genera of itive notoungulate to North America and the Deseado fauna can be cited as clearly thence to Asia would thus provide the following the line indicated by Palaeo- source for the Arctostylopidae (Simpson, stylops-Arctostylops." Nonetheless, as im- 1951, 1965, 1978, 1980). Szalay and plied in the foregoing statement, they re- McKenna (1971) followed Simpson in this garded Palaeostylops as more primitive in respect, noting that molars of then known a number of features (for instance, the sim- arctostylopids were more advanced than pie premolars) than the earliest of the South any in the earliest South American no- American notoungulates or Arctostylops. toungulates. Apparent support for a south- They thus believed the Asian genus to be ern origin of the Notoungulata, on both ancestral, at least in a general sense, to all morphological and temporal grounds, is New World forms, and that "the South lent by the proposed referral of Peru- American Tertiary hoofed mammals were therium, from the Late Cretaceous of Peru, originally derived from the north, al- to the order (Marshall, de Muizon, and though undergoing a great secondary evo- Sige, 1983). Placement of this genus, which lution in the Neotropical region" (p. 2). is based largely on two broken molars of Simpson (1934) clearly defined the No- the type and only species, has been a mat- toungulata and its contents. He removed ter of considerable dispute since its initial the Arctostylopidae and Notostylopidae (a description (Grambast et al., 1967), with group of primitive South American no- workers variously suggesting arctocyonid toungulates) from the "Entelonychia" and (Grambast et al., 1967), didolodontid placed them with the Henricosborniidae (Tedford, 1974), periptychid (Van Valen, in a then new paraphyletic suborder, No- 1978), and even marsupial (Hoffstetter, tioprogonia, defined on the basis of prim- 1981) affinities. Marshall, de Muizon, and itiveness of its constituent taxa. This left Sige (1983) suggested that Perutherium the "Entelonychia" as Ameghino had orig- possesses, in common with notoungulates, inally conceived it except that Simpson re- a pre- and postmetastylid in the trigonid moved the Leontiniidae to the Toxodonta. of the lower molars, and that the genus is Thus recognized, the Notoungulata com- a morphologically appropriate antecedent prised four suborders: Notioprogonia, to both the South American notoungulates "Entelonychia," Toxodonta, and Typothe- and the Arctostylopidae. ria. On the basis of further studies (Pat- Patterson (1958; Patterson and Pascual, terson, 1936; Simpson, 1936b), Simpson 1972), on the other hand, followed Mat- later (1945) removed the remaining con- thew (1928; Matthew and Granger, 1925) tents of the "Entelonychia" (Isotemnidae in believing that notoungulates arose in the and I lomalodotheriidae) to the Toxodon- north and, along with several mammalian ta. where they have since remained. companions, colonized South America in aborated in his two the earliest Tertiary, later to radiate and irliest South flourish on that continent. The basis for tunas (Simpson, 1948, this opinion is unclear, but it is likely that Arctostylopids Mammali fellietal Patterson, like Matthew before him, was impressed by the early records of Arcto- stylopidae in North America (then thought to be early Eocene) and Asia (latest Paleo- cene), and by several of the strikingly primitive dental features found in mem- bers of that family. An Asian origin for the Notoungulata was also suggested by Nesov (1987). Gingerich and Rose (1977) pro- posed yet another possibility, that the No- toungulata arose in Central America (where evidence bearing on this issue is lacking) and from there spread both north- ward and southward. Because of the inferred primitiveness of Asiostylops within the Notoungulata (sim- ple premolars, triangular upper molars lacking a hypocone, simple molar lophs, unreduced anterior wing of lower molar trigonids), Zheng (1979) suggested that the order originated in Asia and, more specif- ically, in southern China. Earliest records need not infallibly indicate centers of or- igin, however. Van Valen (1988) consid- ered Asiostylops to be sufficiently primi- tive to be structurally antecedent to trigonostylopids (an archaic group of As- trapotheria, which are endemic to South America). Several recent studies have emphasized the profound differences in dental spe- cializations between the Notoungulata and the Arctostylopidae, and on this basis have tentatively disassociated Holarctic from South American forms (Cifelli, 1983a, .1985a; Schaff, 1985; Thenius, 1985). It is well worth pointing out that it was Simp- son who first flirted with this possibility, before returning to a more traditional view in the same paper: "A possibility that seems not to have been considered but perhaps should be is that Arctostylops, Palaeostylops, and Sinostylops, although quite sure- ly related among themselves, might not after all be true notoungulates. Their dentitions do have derived characters that occur in almost all ear- ly notoungulates with various modi- fications and some marked changes in later, more specialized forms. These apparently diagnostic characters an not known in am other defined ordei of mammals Nevertheless, these arc unlike South American notoungulates in detail and one cannot absolutely exclude the possibility ol conver- gence." (Simpson, L978, p. 325 Possible Relationships Evaluation of these contrasting v iews on the origin and subsequent dispersal ol the Notoungulata, of great interest in both zoogeographical and paleobiologies terms, is dependent on determination ol mor- phocline polarity sequences and the ro- bustness of the phylogenetic framework derived therefrom. The issue of funda- mental interest, one which remains to be examined in detail, is the phylogenetic po- sition of the Arctostylopidae with respect to South American Notoungulata Assum- ing notoungulate monophyly, inclusive of the Arctostylopidae, three possibilities present themselves: 1) arctostylopids took origin from a southern notoungulate as that group is known (southern origin): 2) the southern notoungulates derived from a form that falls within the Arctostylopidae as that group is here conceived (northern origin for the order); and 3) the Arcto- stylopidae and known South American Notoungulata are sister taxa (northern or southern origin). Even without knowledge of the cranial morphology of arctostylopids (a suite ol synapomorphies characterizes tins region in notoungulates; Simpson. 1948), there is rather imposing evidence, in the dentition and proximal ankle, that the southern No- toungulata constitute a monophyletic as- semblage. Derivation of the Arctostylopi- dae from within the order as it iscmreiitK recognized would require man) simplifi- cations (reversals) in the dentition, because Asiostylops in many cases and all arcto- stylopids in some instances are more prim- itive than any known southern not,. un- gulate. The most significant ol th< Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 characters are in the upper molars. All de Muizon, and Sige (1983) suggest that southern notoungulates have secondary the various accessory tngonid structures ot complications, consisting of at least a cro- notoungulates may be homologized with chet (Patterson, 1934; Simpson, 1948) in a pre- and a postmetastylid and that these the trigon basin on all upper molars and a are primitive for the order. Unlike typical liNpocone on M12; all arctostylopids lack South American notoungulates and the the first character and at least Asiostylops Arctostylopidae, Perutherium lacks an en- among that family lacks either a hypocone tolophid on its lower molars. The absence or hypocone-like structure. By analogy of a pre- and postmetastylid in Asiostylops with a series of variants in M3 of Henri- and Bothriostylops would therefore re- cosbornia lophodonta (Fig. 12), which are quire postulation of secondary loss of these not quadritubercular but which illustrate structures in forms which otherwise seem a plausible character state series for the to be rather primitive in the construction addition of the posterolingual cusp on of their lower molars. Thenius (1985) ac- primitive notoungulate anterior upper cepted the lower molar pre- and postmeta- molars, the posterointernal cusp of south- stylid pattern as a synapomorphy of no- em notoungulates appears to be a deriv- toungulates, and excluded arctostylopids ative of the cingulum and therefore a from the order because it was lacking from "true" hypocone (Simpson, 1929). By con- "Palaeostylops steini." trast, in arctostylopids which have quad- The morphotype for the notoungulate ritubercular M12, the posterointernal cusp proximal ankle bones is not strongly spe- is encircled basally by the cingulum and cialized (as compared, for instance, to un- appears to have originated as a transverse, gulate groups such as the Litopterna, Pe- lingual extension of the metacrista from rissodactyla, Artiodactyla, and Hyracoidea, the region of the metaconule9. Thus, the all of which are highly modified at first posterolingual upper molar cusp of south- appearance in the fossil record). Nonethe- ern notoungulates and arctostylopids ap- less, it is characterized by a number of pears to have been acquired indepen- synapomorphies which render it readily dently and in a nonhomologous fashion. recognized (Cifelli, 1983b). These features Even the most primitive of southern No- include a long, constricted astragalar neck, toungulata (Henricosborniidae) have sub- with an oblique dorsal crest; astragalar molariform posterior lower premolars; P4 body with a median (tibial) protuberance; has a complete, curved talonid crescent, astragalar foramen with posterolateral sul- Although the serially multicuspate, blade- cus interrupting continuity of tibial troch- like lower premolars of such forms as Pa- lea and flexor tendon groove; and well- laeostylops may reflect specialization for developed sustentacular-navicular facet shearing (secondary simplification), Asio- contact on the astragalus. stylops lacks the degree of molarization Except for a constricted astragalar neck, seen even in henricosborniids. none of these features is shared with known The proposed addition of Perutherium arctostylopid ankle regions (Gashatosty- (iltiplanense to the Notoungulata (Mar- lops macrodon and Palaeostylops iturus) , shall, de Muizon, and Sige 1983) presents which bear specializations contrasting with further problems for an origin of the Arc- those of notoungulates. The arctostylopid tostylopidae within that group. Marshall, ankle is advanced in having an astragalus with a cylindrical, vertically-walled body, the tibial trochlea extensivelv developed I considered the posteroin- anteroposterior^; lack of a fibular shelf; il i "sl "I arctostylopid upper molars to be a • i r . i i i .1 .1 elj expanded metaconule; it navicular facet developed so that the axis lowever, that the upper molar conules were of movement along the midtarsal joint would have been roughly parallel (rather Arctostylopids (Mammalia) • Cifelli ei al. mm Figure 12. M3 variants in Henhcosbornia lophodonta, AMNH 28964, from the early Casamayoran Canadon Vaca local fauna illustrating hypothesized addition of hypocone through linking of postcingular cusp and metaloph. than oblique) to that at the proximal ankle joint; astragalar cuboid facet lost (?); ectal facet steeply inclined with respect to in- ferior surface of astragalus; calcaneal fib- ular facet strongly developed into a semi- cylindrical surface; and sustentaculum of calcaneus distally located, at or near distal (cuboid) end of the bone. Most of these ankle modifications are usually associated with restriction of lateral and inversion/ eversion movement, with concomitant greater capability for flexion/extension, at the proximal and mid-tarsal joints. Such specializations are commonly found among terrestrial mammals (Cifelli, 1983b). The extreme distal position of the astragalo- calcaneal facets on the calcaneus (a prim- itive condition?), implying poor mechan- ical advantage for rapid flexion of the pes by the gastrocnemius and soleus muscles, is enigmatic in this regard, and contrasts with the condition seen in terrestrial sal- tators or cursors. In any event, regardless of the paleobiological implications of this unusual ankle morphology, it is clear that notoungulates are uniquely derived with respect to arctostylopids, and vice versa. Derivation of southern notoungulates from the Arctostylopidae (Matthew and Granger, 1925; Patterson, 1958; Zheng, 1979) is also contradicted by the available morphological evidence. Neotropical \<>- toungulata have a different style of upper premolar molarization from that of arc- tostylopids and lack a metacone on P The lower molars of henricosborniids show a very primitive state in the development of the typical notoungulate talonid: the major cusps (entoconid, hypoconid. hy- poconulid) remain distinct; the entolophid is weak. The placement and orientation ol the entolophid suggest that it was derived from the entoconid to hypoconulid pari "I the postcristid. Even in primitive arcto- stylopids (e.g.,Asiostylops), the hv poconid is indistinct, having been merged into tin talonid crescent. The entolophid <>1 arc- tostylopids is advanced in being more an- teriorly placed and is oriented anterola- bially (Schaff, 1985). If Perutherium ^ a notoungulate, as argued l>\ Marshall, de Muizon, and Sige (1983), then derivation of South American taxa from arctostylo- pids would require independent acquisi- tion of the entolophid in the Neotropical forms, because that structure is lacking in Perutherium. The final possibility is that known \rc- Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 tostvlopidae and southern Notoungulata (Anatolostylops; unnamed genus and are sister taxa: that they shared an ancestor species), which lacks folds other than those that was exclusive to them and no other for the parastyle and metastyle, is remi- groupof mammals. This hypothesis would niscent of that of notoungulates such as be compatible with all existing scenarios Notostylops (comparison with which was regarding the geographic origin and dis- the basis for the genus and family-group persal of notoungulates. With the addition names of the northern forms) and various to the Arctostylopidae of primitive forms Leontiniidae, but primitive members of such as Asiostylops and Bothriostylops, both the Arctostylopidae and southern No- nearly all of the similarities shared by toungulata have lower, more complexly southern notoungulates and advanced arc- folded ectolophs. Arctostylops and ?Pa- tostylopids would have been acquired in- laeostylops also resemble some southern dependency and therefore represent par- notoungulates, especially Notostylops, in allelisms. These include the reduction of the high talonid crescent, which achieves the lower molar trigonids, the addition of an anterior attachment with the trigonid accessory trigonid structures to those teeth at a very labial position; this, again, is not (the homology of these structures, termed a condition shared by more primitive pre- and postmetastylid by Marshall, de members of either group. Muizon, and Sige, 1983, is open to some question, even among the taxa restricted Remaining Resemblances to South America), the development of a With the dismissal of many arctostylo- talonid on ?4, and the upper molar crown pid-notoungulate similarities as conver- pattern, which is superficially similar but gent acquisitions within each group, it is appears on other grounds to include non- relevant to evaluate the uniqueness of re- homologous features, as discussed above, semblances that remain. The most striking What is known of the ankle region in arc- of these is the transversely developed, lo- tostylopids indicates that they are diver- phate entoconid (entolophid) of the lower gently specialized from notoungulates. One molars. This is an unusual but not excep- specialization of the arctostylopid ankle, tional feature among mammals: it surely the development of the calcaneal fibular developed independently in the Astra- facet into a large, semicylindrical surface, potheria and twice among the Litopterna is found among a group of advanced tox- (Cifelli, 1983a; Cifelli and Soria, 1983). odont Notoungulata (the monophyletic Among Holarctic mammals, an entolophid group including Notohippidae, Leontini- or similar structure developed indepen- idae, and Toxodontidae), but this was dently in numerous rodent lineages (L. L. clearly developed independently by them. Jacobs, personal communication). Without Certain other notoungulate resemblances knowledge of more primitive forms, it is of arctostylopids, which undoubtedly in- not possible to determine if the arctosty- fluenced early workers in their compari- lopid entolophid arose, as in the southern sons and in their speculation regarding re- notoungulates, from part of the postcristid lationships, evidently represent derived or if it is demonstrably non-homologous ( liaracter states within both groups and are (the entolophid of astrapotheres, for in- almost certainly convergent. These in- stances, appears to be a de novo structure), elude the presence of a labial ectocingulid, If, as argued by Marshall, de Muizon, and which is characteristic of most toxodont Sige (1983), Perutherium is a notoungu- lower molars and premolars and of ad- late, then independent acquisition of the vanced forms (e.g., Palaeostylops, Ga- entolophid in the Arctostylopidae is sug- shatostylops, and Arctostylops) among the gested by the fact that they primitively Vrctostylopidae. The smooth ectoloph of lack the accessory trigonid structures pos- y lopid upper molars sibly shared by that genus with Neotrop- Arctostylopids (Mammalia) • Cifelli et al. 39 ical Notoungulata. Other shared dental ambiguously homologous, and in part be- features of arctostylopids and notoungu- cause the evidence of relationship has of ten lates, derived with respect to an ungulate been based on shared primitive features morphotype, might include a crescentic rather than uniquely derived specializa- lower molar trigonid (this condition is tions. The Arctostylopidae have been im- somewhat uncertain in southern notoun- mune to such controversy because, despite gulates, as the trigonid is already reduced some unique aberrancies and retention ol at first appearance), reduced upper molar a few primitive features, the advanced stylar shelf and lobes, and slightly raised genera Arctostylops and Palaeostylops centrocrista between paracone and meta- strikingly resemble notoungulates and no cone on the upper molars ("incipient" ec- other mammals in certain aspects of their toloph). These latter features are not in dental anatomy. Evaluation of the realit) themselves or collectively diagnostic, as of this relationship and its precise nature they represent generalized, almost gradal was long hampered by insufficient knowl- trends in many different groups of Paleo- edge of arctostylopid morphologic diver- cene and Eocene ungulate-like mammals, sity and of the structure and relationships Of the three alternatives of arctostylo- of the most primitive notoungulates of pid-notoungulate relationships discussed South America. With these circumstances above, the most permissive, that they rep- now dramatically improved, considerable resent sister taxa, is the most likely. (This doubt is cast on the close relationship of is true by definition, as the other two pos- the two groups, accepted without question sibilities are more specific and therefore for most of this century. A common no- more susceptible to falsification.) Yet, be- toungulate/arctostylopid ancestor (i.e., a cause most similarities of arctostylopids to morphotype for the two groups, consid- notoungulates must have arisen indepen- ered as sister taxa) might have been suf- dently, whether by parallelism or conver- ficiently primitive to have given rise to gence, the evidence that they collectively many other orders of mammals. In rec- comprise a monophyletic unit with respect ognition of this, and considering the ample to other mammals is slim: it amounts, in evidence for monophyly of the Arctosty- fact, to one possible character (entolophid) lopidae, we have referred the family to its that is known to have developed indepen- own order. Thus recognized, the group dently several times among other, unre- would represent an Asian radiation that lated groups. This is hardly secure docu- managed to disperse to North America, mentation of monophyly. Other evidence, possibly in the late Paleocene. The geo- such as that provided by the ankle region, graphic distribution of arctostylopid taxa, suggests that a common ancestor of the two and the hypothesized immigration to \(>rtn groups would have been exceedingly America, are given in Figure 13. It is in- primitive and, probably, not exclusive. teresting to note that most of the primitive forms are more southerly in distribution. Distinctness Of ArctOStylopida being found in south China, while spe- Since the time of Ameghino, many close cialized taxa are generally northerl) in dis- relationships of South American with Hoi- tribution. arctic forms have been proposed (see, e.g., The broader relationships oi Vrctosty- summaries by Simpson, 1978; McKenna, lopida among the Mammalia are em 1981; and Gingerich, 1985). With the ex- matic. The arctostylopid dental morpho- ception of marsupials, the controversy sur- type bears some similarity to several \muii rounding all ordinal and lower level re- taxa of debatable affinities, such as Lan- ferrals of South American to Holarctic taxa tianius (Cifelli, 1983a) and Petrolemur has been considerable, in part because de- although contrasting specializations rived similarities are incomplete or not un- as loss of premolars in the latter genusj are [Q Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 140 120° 100° T CHI-JIANG BASIN Asiostylops span/os Al lost y lops periconah Bothnostylops notios NAN-XIONG BASIN Arctostylopidae gen.+ sp. nov ^■\P£\ CHINA gxVhun^n qiaNSHAN BASIN """ Si no sty lops promissus_ _ , - - XUAN- CHENG -" ~=? -anhui u BASIN ^-N /& Bothnostylops propressusi ^ 0,'°*J . ^-Beijing' XINJIANG TURPAN BASIN ^r' Anatolostylops dubius NEMEGT BASIN Palaeostylops iturus Gashatostylops macrodon , Arctostylopids (Mammalia) • Cifelli et al. 41 evident. Both forms were originally re- ferred to the Primates; the ankle of known arctostylopids is completely dissimilar to any belonging to that order. Arctostylopid ankle specializations are shared, as best we are able to determine from published fig- ures (Sulimski, 1968; Szalay, 1977, fig. 16), with the Asian late Paleocene Pseudictops. This taxon has, in turn, been considered to be part of "an endemic Cretaceous and early Tertiary Asian radiation, whose clos- est living relatives are the Lagomorpha" (Szalay and McKenna, 1971, p. 301). Whatever the constituents of this radiation (see also McKenna, 1975; Novacek, 1986; Szalay, 1977), we note that lagomorphs and some of their suspected allies are special- ized for saltatory locomotion (Szalay, 1977; see Bleefeld and McKenna, 1985, for de- scription of some lagomorph ankle spe- cializations); arctostylopids — which may just be primitive in this regard — appar- ently were not, as indicated by the lever mechanics of the calcaneus. The diversity and abundance of arcto- stylopids in early Tertiary Asian faunas, coupled with the proposed close relation- ship of North American Arctostylops to Asian Palaeostylops as rather derived taxa within the family, suggests that dispersal from west to east, rather than the reverse, is the most probable explanation for geo- graphic distribution of the group. Owing to high endemism of Asian faunas older than those of the North American Wa- satchian, correlation of earliest Tertiary mammalian assemblages between the two continents has been problematic (Szalay and McKenna ,1971). The presence of Arc- tostylops in the Tiffanian (late Paleocene) of North America, the geometry of pro- posed relationships among the Arctosty- lopidae, and the fact that more primitive taxa are known from Asia but not North America, suggest a late Paleocene (Dash- zeveg, 1982; Szalay and McKenna, 1971), rather than early Eocene (Gingerich and Rose, 1977) age for Asian faunas, such as Gashato, which include Palaeostylops and Gashatostylops. LITERATURE CITED AMEGHINO, F. 1894. Enumeration synoptique des especes de mammiferes fossiles des formations eocenes de Patagonie. Boletin d<- L Vcademia Nacional de Ciencias de Con 1( il>a 13:259 l~>_: . 1906. Les formations sedimentaires du < tace superieur et du tertiare de Patogonie av« un parallcMc entre leurs faunes mammalogiques et celles de l'ancien continent Anales del Museo Nacional, Buenos Aires, 15(ser. 3. \<>l 8 I 568 Bond, M. 1981. Un nuevo Oldfieldthomasiidai (Mammalia, Notoungulata) del Eoceno inferior (Fm. Lumbrera, GrupoSalta idelNW ^rgentino Anales II Congreso Latino-Americano de Pa- leontologia, Porto Alegre: 521-536. Bleefeld, A. R., and M.C. McKenna 1985. Skel- etal integrity of Mimolagus rodens (Lagomor- pha, Mammalia). American Museum Novitates, 2806: 1-5. Chow Min-chen, Qui Tao. vnd Li Vi \<. 1977 Paleocene stratigraphy and faunal characters of mammalian fossils on Nomogen Commune, Si- zi-wang-qi, Nei Mongol. Vertebrata PalAsiatica. 14: 228-233 [in Chinese; English abstract]. Chow Min-chen, and Qui Tao. 1978. Paleocene mammalian fossils from Nomogen Formation of Inner Mongolia. Vertebrata Pal Asiatica 16: 77- 85 (in Chinese; English abstract). ClFELLI, B. L. 1983a. The origin and affinities of the South American Condylarthra and early Ter- tiary Litopterna (Mammalia). American Mu- seum Novitates, 2772: 1-49. . 1983b. Eutherian tarsals from the late Pa- leocene of Brazil. American Museum Novitates, 2761: 1-31. . 1985a. South American ungulate evolution and extinction, pp. 249-266. In S. D. Webb and F. G. Stehli (eds.), The Great American Iiiotic Interchange. New York: Plenum Pub! ( o itvii + 532 pp. . 1985b. Biostratigraphy of the Casamayor- an, early Eocene, of Patagonia American Mu- seum Novitates, 2820: 1-26. . and M. F. Soria. 1983. Systematic ol the Adianthidae (Litopterna. Mammalia American Museum Novitates, 2771: 1 2 i Colbert, E. H. 1973 Wandering Lands and An imals. New York: E. P. Dutton. 323 pp. Darlington, P. J. 1957. Zoogeograph> The < Figure 13. Lamberts azimuthal equal area projection map of Northern Hemisphere, showing arctostylopid ( hypothesized dispersal route to North America. Bulletin Museum of Comparative Zoology, Vol. 152, No. 1 graphical Distributions of Animals. New York: Wile) and Sons. 675 pp. DASHZEVEG, D. 1982. La faune de Mammiferes du Paleogene inferieur de Naran Bulak (Asie cen- trale) et ses correlations avec l'Europe et l'Ame- rique du Nord. Bulletin de la Societe Geologique de France, 24(2): 275-281. DASHZEVEG, D., AND D. RUSSELL. 1988. Paleocene and Eocene Mixodontia (Mammalia, Glires) of Mongolia and China. Palaeontology, 31: 129- 164. GAUDRY, A. 1902. Recherches paleontologiques de \l Andre Tournouer en Patagonie. Bulletin de la Societe d'Histoire Natural de Autun, 15: 117- 123. . 1904. Fossiles de Patagonie. Dentition de quelques mammiferes. Memoirs de la Societe Geologique de France (Paleontologie), 31: 1-27. . 1906. Fossiles de Patagonie. Etude sur un portion du monde antartique. Annales de Pa- leontologie (Paris), 1: 1-42. 1908. Fossiles de Patagonie. De l'economie dans la nature. Annales de Paleontologie (Paris), 3: 1-28. GlNGERH II. P. D. 1974. Size variability of the teeth in living mammals and the diagnosis of closely related sympatric fossil species. Journal of Pa- leontology, 48: 895-903. . 1985. South American mammals in the Paleocene of North America, pp. 123-137. In F. G. Stehli and S. D. Webb (eds.), The Great Amer- ican Biotic Interchange. New York: Plenum Press, xvii + 532 pp. \\i) K. D. Rose. 1977. Preliminary report on the American Clark Fork mammalian fauna, and its correlation with similar faunas in Europe and Asia. Geobios, Memoire Special, 1: 39-45. Gradzinski, R., J. Kazmiercak, and J. Lefeld. 1969. Geographical and geological data from the Polish-Mongolian paleontological expedi- tions. Palaeontologica Polonica, 19: 33-82. Grambast, L., M. Martinez, M. Mattauer, and L. Thaler. 1967. Perutherium altiplanense nov. gen., nov. sp., premier mammifere meso- zoique d Amerique du Sud. Comptes Rendus de la Academie des Sciences, Paris, ser. D, 264: 707- 710 GREGORY, W. K 1910. The orders of mammals. Bulletin ol the American Museum of Natural History, 27: 1-254. 1920. On the structure and relations of Notharctus, an American Eocene primate. Mem- oirs of the American Museum of Natural History, n.s. 3: 51-243. Hoi i mi i ii it, R. 1970. Radiation initiale de Mam- mileres placentaires et biogeographie. Comptes Rendus i OKI), B. H. 1974. Marsupials and the new pa- leogeography, pp. 109-126. In C. A. Boss (ed.), Paleogeographic Provinces and Provinciality. So- ciety of Economic Paleontologists and Mineral- ogists, special publication 21. Tin NIUS, E. 1985. Zur systematischen stellung der Arctostylopiden (Mammalia, Notoungulata). Eine odontologische analyse. Bazprave IV. Bazreda Sazu (Zbornik Ivana Bakovca), 26: 147-156. V \\ Valen, L. 1978. The beginning of the age of mammals. Evolutionary Theory, 4: 45-80. . 1988. Paleocene dinosaurs or Cretaceous ungulates in South America. Evolutionary The- ory, 10, 79 pp. Vucetich, M. G. 1980. Un nuevo Notostylopidae (Mammalia, Notoungulata) proveniente de la Formacion Lumbrera (Grupo Salta) del noroeste Argentine Ameghiniana, 17: 363-372. Zhai Ben-jie. 1978. More fossil evidences favour- ing an early Eocene connection between Asia and Neoarctic. Memoirs of the Institute of Vertebrate Paleontology and Paleoanthropology, Academ- ica Sinica, 13: 107-115 [in Chinese]. Zheng Jia-jian. 1979. The Paleocene notoungu- lates of Jiang-xi, pp. 387-394. In A Symposium on Cretaceous and Early Tertiary Bed Beds of South China. Beijing: Science Press, Academica Sinica. [in Chinese]. , and Huang Xueshi. 1986. New arcto- stylopids (Notoungulata, Mammalia) from the late Paleocene of Jiangxi. Vertabrata PalAsiatica, 24: 121-128 [in Chinese; English summary]. Zhu-chen. 1986. Late Cretaceous red beds in Hun- an. Journal of Stratigraphy, 10: 54-59 [in Chinese]. (US ISSN 0027-4100) Bulletin of the Museum of Comparative Zoology Y 390 HARVARD Primary Types of MicrolepMbjifeiPaiirvthe Museum of Comparative Zoology (with a Discursion on V. T. Chambers' Work) SCOTT E. MILLER and RONALD W. HODGES HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS, U.S.A. VOLUME 152, NUMBER 2 19 APRIL 1990 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952- Bl LLETIN 1863- Memoirs 1864-1938 Johnsonia, Department of Mollusks, 1941- Occasional Papers on Mollusks, 1945- SPECIAL PUBLICATIONS. 1. Whittington, H. B., and E. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution of Crustacea. 192 pp. 2 Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Tere- dinidae (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4. Eaton, R. J. E., 1974. A Flora of Concord. 236 pp. 5. Rhodin, G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 745 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprint. Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. Creighton, W. S., 1950. The Ants of North America. Reprint. Lyman, C. P., and A. R. Dawe (eds.), 1960. Symposium on Natural Mam- malian 1 libernation. Ornithological Gazetteers of the Neotropics (1975-). Peters" Check-list of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1948. (Complete sets only.) Publications of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publica- tions Office. Museum of Comparative Zoology, Harvard University, Cam- bridge, Massachusetts, 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 1990. PRIMARY TYPES OF MICROLEPIDOPTERA IN THE MUSEUM OF COMPARATIVE ZOOLOGY (WITH A DISCURSION ON V. T. CHAMBERS' WORK) SCOTT E. MILLER1 and RONALD W. HODGES2 ABSTRACT: Primary types (holotypes, lectotypes and syntypes) in the moth superfamilies Erioeranioidea, Hepialoidea, Nepticuloidea, Incurvarioidea, Tineo- idea (except Gracillariidae), Geleehioidea (except Co- leophoridae), Copromorphoidea, Yponomeutoidea, Sesioidea, Cossoidea, Tortricoidea, Zygaenoidea, Pyr- aloidea, and Pterophoroidea are listed. Most of the taxa are Nearctic, several Neotropical. Authors in- cluded are S. E. Cassino, V. T. Chambers, W. G. Dietz, H. Edwards, C. Fish, W. T. M. Forbes, H. Frey and J. Boll, A. R. Grote, T. W. Harris, G. D. Hulst, W. D. Kearfott, A. S. Packard, Lord Walsingham, and P. C. Zeller. Lectotypes are designated herein for Anacampsis quadrimaculella (Chambers) and Is- ophrictis trimaculella (Chambers) (both Gelechi- idae). INTRODUCTION Classification of several groups of North American microlepidoptera is made very- difficult because most early workers on this fauna did not designate type specimens; their descriptions are inadequate to rec- ognize the species; their "type" material was sent to more than one institution; and among them they described several hundred species. The Museum of Com- parative Zoology (MCZ), Harvard Univer- sity, has a significant number of types of these authors, particularly of V. T. Cham- bers. Chambers is notorious for his very brief and inadequate descriptions, the large 1 Museum of Comparative Zoology, Harvard Uni- versity, Cambridge, Massachusetts 02138. Present ad- dress: Bishop Museum, Box 19000-A, Honolulu, Ha- waii 96817. 2 Systematic Entomology Laboratory, Agricultur- al Research Service, USDA, % NHB 168, National Museum of Natural History, Washington, D.C 20560. number of his often very short papers in scattered journals, the large number of species for which no type material exists, and for the fact that his types were dis- persed among the MCZ and contemporar) workers. Because we have studied Cham- bers' papers and believe we have uncov- ered nearly all remaining specimens that can be considered authentic, we have in- cluded a discussion of specimens in other collections (see Appendix) and Chambers' bibliography to apprise other workers of the facts they will need when selecting lectotypes or designating neotypes. Man) species are represented by no extant type material or are not represented by speci- mens in the MCZ; their names do not ap- pear in the catalog. This catalog lists the primary types of 609 taxa of Microlepidoptera located in the MCZ. All holotypes, lectotypes, and syntypes in the families treated are listed along with some paralectotypes and "psen- dotypes." All specimens (except obvious paratypes) with numbered red "MCZ. type" labels are included. These red num- bered labels were placed on the specimens by Nathan Banks and subsequent curators and sometimes are not accurate as noted in this list. Some lectotypes have been des- ignated by previous authors by inference of holotype (Article 74(b), International Code of Zoological Nomenclature, third edition, 1985). For example, a lectotype was designated for Nepticula castaneae- foliella Chambers by inference ol holo- type. We have not designated additional lectotypes here (excepl for two as part oi Bull. Mus. Comp. Zool., 152(2): 45-87, April. 1990 45 46 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 Hodges' research) because this action lection includes the Peabody Academy should be left to specialists as part of the collection (the Peabody Academy still ex- revision process. Each species-group entry ists in Salem, Massachusetts, but no longer has five potential topics: 1) Original com- maintains entomological collections) and lunation, author, and date-page citation; the types from the now defunct Boston 2) Category of type, sex of specimen(s), Society of Natural History. Details on the and MCZ type number; 3) Geographic dis- major type collections discussed here fol- tribution as indicated in the original de- low: scription and/or labels accompanying the Vactor T. Chambers: Most of the re- type specimens; 4) Current valid name (if mains of the Chambers collection are at different from original combination); and the MCZ, with another significant part at 5) Remarks. the USNM. See appendix for discussion of Information presented in brackets [ ] possible Chambers types at the BMNH. represents additions to or corrections of the Chambers deposited types at the MCZ be- original description. The present valid tween 1876 and 1883 (Hagen, 1884); many name of each taxon listed, if it differs from of them are in very poor condition, and the original combination, is also included many probably are not true types but spec- ( following Hodges, et al., 1983 and more imens substituted by Chambers for lost or recent literature). Sex is included only when damaged types. it could be determined readily without Walsingham (1889: 24) wrote, "Cham- damaging the specimen. The locations of bers, in distributing specimens to his var- some other syntypes are indicated if known, ious correspondents, frequently appears to using the following abbreviations: AMNH have attached a wrong name to them. This = American Museum of Natural History, he admits in more than one instance in his New York; ANSP = Academy of Natural writings. The utmost caution is required Sciences, Philadelphia; BMNH = British before accepting a specimen in any col- Museum (Natural History), London; lection as a co-type of any one of his LACM = Natural History Museum of Los species." Angeles County, Los Angeles; and USNM Chambers (1877c: 39) wrote: ". . . But National Museum of Natural History, a few years ago I began to make a collec- Smithsonian Institution, Washington. tion to be preserved as types of all my The following superfamilies (following species. These were all pinned and spread. Hodges, et al., 1983) are included in the Unfortunately, during my absence in Col- list: Eriocranioidea, Hepialoidea, Nepti- orado, the greater part of this collection culoidea, Incurvarioidea, Tineoidea (ex- was destroyed. One or more specimens of cept Gracillariidae), Gelechioidca (except the greater number of species were for- Coleophoridae), Copromorphoidea, Ypon- tunately preserved, and most other species omeutoidca, Sesioidea, Cossoidea, Tortri- can be supplied. This collection is now in coidea. Zygaenoidea, Pyraloidea, and the Cambridge Museum [MCZ]. It contains I'terophoroidea Data For Tineidae and types — pinned and spread — of some- Blastobasidae were provided by D. R. Da- thing over 200 species." (See also Braun, vis and I). Adaniski. respectively. Gracil- 1963: 2; Hagen, 1884; Sattler, 1962.) larudae are under stud\ by D. R. Davis Most Chambers specimens bear only the (I SNM). Coleophoridae are under study data "Kentuckv. /Chambers." (machine b) B. Wright (Nova Scotia Museum) and printed) along with a handwritten deter- were discussed by McDunnough (1944). mination label, and frequently a large All the types listed here are in the main handwritten number (between 12 and 164) MCZ collection, except those in the Harris on a separate label (the meaning of which collection, which is maintained separately remains unknown). Most of these were from the general collection. The MCZ col- presumably collected around his home in MCZ Microlepidoptera Types • Miller and Hodges 47 Covington, Kentucky (Chambers, 1875b: 234). Chambers wrote (1872: 433), "Out of at least one hundred and fifty species of Tineina which I have found here, fully three-fourths have been taken resting upon the leeward side of a board fence not two hundred yards long, at Linden Grove Ce- metary at this place [Covington, Ken- tucky].'' Other major lots of material include those collected by Chambers (and others) in Colorado in 1875 (and other years) and Texas specimens purchased by Chambers from Gustaf W. Belfrage. Belfrage lived near Norse, Bosque County (some 60 km northwest of Waco), Texas, from 1868 to 1882 (Geiser, 1948). Most, but not all, of Belfrage's moth material was probably col- lected there. The USNM has many Chambers types, obtained primarily through acquiring the collections of C. H. Fernald (which in- cluded part of the M. Murtfeldt collec- tion), F. H. Belanger (via Laval University, Quebec), and W. Saunders (Busck, 1903: 768). Type localities for many of Chambers' species are not obvious from the original descriptions. We have followed the label data on the types, unless contradicted by other evidence. Because of the scattered nature of Chambers' publications, we have included all of them on Lepidoptera in our litera- ture section, whether mentioned here or not. William G. Dietz: His types of Blasto- basidae and Gracillariidae are in the MCZ. Heinrich Frey and Jacob Boll: Several types of species described by Frey and Boll (1873) previously assumed to be at the BMNH (e.g., Braun, 1972: 56) are present. These were evidently collected by Boll around Cambridge, Massachusetts, in au- tumn 1871 (Geiser, 1948: 22-23). Thaddens W. Harris: Probably the old- est extant collection of North American insects, most specimens are still in good condition. It is held as a separate unit at the MCZ because the labels are cryptic (Johnson, 1925). Most specimens bear Har- ris numbers, the catalog of which is in the MCZ Archives. Specimens for which fur- ther data are not given probably came from Massachusetts. George D. Hnlst: As discussed by Rindge (1955), the main Hulst collection is at AMNH, but the MCZ has syntypes of some Hulst taxa. William D. Kearfott: The MCZ has a number of Kearfott syntypes (labeled "co- type"), some of which have now become paralectotypes. As discussed by Klots (1942: 392-393), much of Kearfott's collection is in AMNH, but parts are in USNM (via the Barnes collection), and elsewhere (includ- ing MCZ). The best candidates for lecto- type designation for Kearfott taxa will gen- erally be found at AMNH or USNM, not at MCZ. Authorship of lectotype desig- nations in some Kearfott Tortricidae re- mains problematic; Klots (1942) credited many lectotype designations to Heinrich (1923, 1926)'. However, Heinrich did not publish which specimen he considered the type if there was more than one synt\ pe in AMNH. Klots usually did designate in- dividual specimens, and should be consid- ered the designator of most of the lecto- types in question. Alpheus S. Packard, Jr.: Included here are California specimens collected by Henry Edwards. Some of these Edwards specimens bear Edwards' catalog num- bers; data from Edwards' catalog (now in AMNH) are included here. Lord Walsingham [Thomas de Grey]: The MCZ has one Walsingham holotype (Eriocraniidae) and many syntypes (Oec- ophoridae and Plutellidae). The syntypes are duplicates of species described by Wal- singham (1881) from material he collected in California and Oregon in 1871-72 (see Essig, 1941), which were sent to Chambers by Walsingham. Lectotypes for these taxa should be designated from syntypes in the Walsingham collection at the BMNH. Philipp C. Zeller: The MCZ has most of the specimens collected by Jacob Boll around Dallas, Texas in late 1869 and 1870 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 Geiser, 1929. 1948) and described by Zell- er in three papers on North American Mi- crolepidoptera (1872, L873, L875). The Boll specimens were purchased by Louis Agas- si/ tor the \1( r/and sent toZeller for study. The) all bear characteristic labels: ma- chine printed "Dallas/Tex. Boll" and handwritten Zeller determination labels on Hreen paper. The following corrections and additions to recent literature are noted in the list: Several types not located by Wilkinson and coauthors are included (Nepticulidae). The syntypes of 11 1 1 arris species reported lost In Duckworth and Eichlin (1978) are in- cluded (Sesiidae). The following were ac- cidently omitted from Hodges, et al. (1983): "Elachista" texanella Chambers (Scythri- didae), Paralipsa fulminalis (Zeller) (Pyr- alidae) and Marasmarcha pumilio (Zeller) (Pterophoridae). Problems are noted in previous lectotype selections for Pyrausta unt/ascia/w(Packard) (Pyralidae) and Oi- daematophorus grandis (Fish) (Ptero- phoridae). LECTOTYPES are here des- ignated for Anacampsis quadrimaculella (Chambers) and Isophrictis trimaculella (Chambers) (both Gelechiidae). "Scy- thris" albapenella (Chambers) is here transferred from Scythrididae to Blasto- basidae on advice of J. F. Landry and D. Adamski; Adamski will deal with its ge- neric placement in a subsequent publica- tion. ACKNOWLEDGMENTS We thank V. O. Becker, R. L. Brown, C. \ . Covell, D. R. Davis, J. B. Heppner, W. E. Miller, E. G. Munroe, R. W. Poole, J. \ Powell, G. Robinson, K. Sattler,M. Shaf- fer, I C. Thompson, K. Tuck, A. Watson, and P. Whalley for reviewing the manu- script; F. H. Rindge (AMNH) and J. P. Donahue (LACM) for information on types in other collections: and M. D. Bowers, A. 1 Newton, and D. F. Schweitzer for sup- port at MCZ. Development of this list was rtially supported 1>\ National Science lation facilities grant BSR-82 03845 ( » Wilson and \1. D. Bowers of the MCZ. A. G. Wine and T. M. Kuklenski prepared the final manuscript. Publication costs of this study were covered in part by a grant from the Wetmore Colles Fund. SUPERFAMILY ERIOCRANIOIDEA Family Eriocraniidae auricyanea Walsingham, 1882: 204, Micropteryx [sic]; HOLOTYPE male, MCZ 1622; [United States: probably California (see Davis, 1978)]; Dyserio- crania auricyanea (Walsingham). SUPERFAMILY HEPIALOIDEA Family Hepialidae argenteomaculatus Harris, 1841: 295, Hepiolus [sic]; SYNTYPE, MCZ 26378; United States: [Massachu- setts, Cambridge, Harris no. 257]; Sthenopis ar- genteomaculatus (Harris); Often cited as 1842, the description was originally published in 1841, and reprinted in 1842 (page 295 of both works). labradoriensis Packard, 1864c: 394, Hepialus; HO- LOTYPE [?] male, MCZ 160; Canada: Labrador, Straits of Belle Isle, Caribou Island, Salmon Bay, 3 August 1860, A. S. Packard, Jr.; Korscheltellus gra- cilis (Grote) (see Wagner, 1988); Abdomen and wings glued in place. SUPERFAMILY NEPTICULOIDEA Family Nepticulidae apicialbella Chambers, 1873: 127. Neptieula; PAB.A- LECTOTYPES (5), MCZ 1496; United States: Ken- tucky, June, Chambers; Stigmella apicialbella (Chambers); Lectotype (USNM type 523) and 3 paralectotvpes in USNM designated bv Newton and Wilkinson (1982: 367). bosquella Chambers, 1878c: 106, INepticula; SYN- TYPES (2 males), MCZ 14958; United States: Texas, Bosque County; Ectoedemia obrutella (Zeller); A male syntype in USNM (type 524). castaneaefoliella Chambers, 1875a: 117, Neptieula; LECTOTYPE female, MCZ 14956; United States: Kentucky, Chambers: Stigmella castaneaefoliella (Chambers); Wilkinson and Scoble (1979: 46), des- ignated the lectotype (Code, Art. 74(b)). ciliaefuscella Chambers, 1873: 128, Neptieula; SYN- TYPE, MCZ 1301; United States: Kentucky, "at lamp." 23 August [year not stated], Chambers; Stig- mella fuscotibiella (Clemens). clemensella Chambers, 1873: 125, Neptieula; LEC- MCZ Microlepidoptera Types • Miller and Hodges 49 TOTYPE female, MCZ 14955; United States: Ken- tucky, Chambers; Ectoedemia clemensella (Cham- bers); Lectotype and paralectotype in MCZ by Wilkinson and Scoble (1979: 86).' grandisella Chambers, 1880b: 193, Nepticula; HO- LOTYPE male, MCZ 1302; United States: Texas; Ectoedemia grandisella (Chambers). juglandifoliella Chambers, 1878c: 105, INeplicula; SYNTYPE, MCZ 1495; United States: Kentucky, Chambers; Stigmella juglandifoliella (Clemens); Chambers (1878c) used Clemens' (1861) name for "mine and adult, the mine of which was described by Clemens. Despite acknowledgement of Cle- mens' previous use, Chambers called his name a new species. latifasciella Chambers, 1878c: 106, Nepticula; HO- LOTYPE female, MCZ 1497; United States: Ken- tucky, "on . . . chestnut-trees,'' August, Chambers; Stigmella latifasciella (Chambers). maculosella Chambers, 1880b: 193, Nepticula; HO- LOTYPE female, MCZ 1303; United States: Texas; Stigmella nigriverticella (Chambers); Newton and Wilkinson (1982: 425) commented on status of this name. maximella Chambers, 1873: 126, Nepticula; SYN- TYPES (2), MCZ 14951; United States: Kentucky, Chambers; Ectoedemia platanella (Clemens); Not mentioned by Wilkinson and Scoble (1979) or Wil- kinson and Newton (1981). pomivorella Packard, 1870: 237, Micropteryx; SYN- TYPE, MCZ 1499; United States: Massachusetts, Salem, A. S. Packard, Jr; Stigmella pomivorella (Packard); There are two specimens, an adult ("Imra. 12.[18]71") and a cocoon ("Apple June 19"), probably from the same individual as stated by Busck (1901: 52). quericastanella Chambers, 1873: 127, Nepticula; SYNTYPES (3), MCZ 1304; United States: Ken- tucky, Chambers; Stigmella saginella (Clemens). quercipulchella Chambers, 1878c: 105, Nepticula; HOLOTYPE male, MCZ 14957; United States: Kentucky, Chambers; Stigmella quericipulchella (Chambers). resplendensella Chambers, 1875a: 118, Nepticula; LECTOTYPE, MCZ 14954; United States: Ken- tucky, 23 May [year not stated], Chambers; Stig- mella resplendensella (Chambers); Lectotype des- ignated by Newton and Wilkinson (1982: 456) who incorrectly stated it was in ANSP. serotinaeella Chambers, 1873: 126, Nepticula; SYN- TYPE, MCZ 1498; United States: Kentucky, Cham- bers; Stigmella prunifoliella (Clemens); Head and forewings only. thoracealbella Chambers, 1873: 127, Nepticula; LECTOTYPE male, MCZ 14952; United States: Kentucky, June, Chambers; Microc(dij)ttris thora- cealbellus(Chambers); Wilkinson (1979: 70) des- ignated the lectotype (Code, Art. 74(b)). uuifasciella Chambers. L875a: 119, Nepticula; LEC- TOTYPE female, MCZ 1305; United States: Ken- tucky, Chambers; Stigmella uuifasciella (Cham- bers); Lectotype and paralectotype also in MCZ designated by Newton and Wilkinson (1982: 440] Family Tischeriidae aenea Frev and Boll, 1873: 222, Tischeria; S"i \ TYPES (5), MCZ 1349; United States: Massachu- setts, "Cambr.B." [=Cambridge, Boll or Cam- bridge, Boston]; Braun (1972: 56) stated "Type, Texas (probably Dallas)[BM]," but type locality is not specified in original description, and introduc- tion to the paper indicates most of the species were reared by Boll at Cambridge. badiiella Chambers, 1875a: 109, Tischeria; SYN- TYPES (8), MCZ 14941; United States: Kentucky, Chambers; Braun (1972: 21) stated "Type? Ken- tucky (MCZ?); Type [female] Kentucky^?), geni- talia slide 9707 J.F.G.C (USNM)." USNM speci- men is type 516. clemensella Chambers, 1878c: 99, Tischeria; SYN- TYPE, MCZ 14940; United States: Texas; Braun (1972: 34-35) stated type locality is Kentucky as implied by Chambers (1875a: 110, 1878c: 99), but specimen is labelled "Tex." concolor Zeller, 1875: 352, Tischeria; HOLOTYPE female, MCZ 1348; United States: Texas, Dallas, Boll; Braun (1972: 27) stated "Type [female], Texas (MCZ)." fuscomarginella Chambers, 1875a: 110, Tischeria; LECTOTYPE male, MCZ 14938; United States Kentucky, Chambers; Braun (1972: 35) designated the lectotype (Code, Art. 74(b)). heliopsisella Chambers, 1875a: 113, Tischeria; SYN- TYPES (2), MCZ 1503; United States: Kentucky, Chambers. latipenella Chambers, 1878c: 97, Tischeria; HO- LOTYPE male, MCZ 14942; United States: Texas; Tischeria zelleriella Clemens. pulvella Chambers, 1878c: 99, Tischeria; LECTO- TYPE, MCZ 1505; United States: Texas; Braun (1972: 97) designated the lectotype (Code, Art. 74(b)). purinosella Chambers, 1875a: 110, Tischeria; LEC- TOTYPE, MCZ 14939; United States: Kentucky, Chambers; Braun (1972: 29) designated the lecto- type (Code, Art. 74(b)). quercivorella Chambers, 1875a: 109, Tischeria; SYNTYPES (5), MCZ 1506; United States Ken- tucky, Chambers; Tischeria citrinipennella Cle- Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 mens; Braun (1972: 15) stated "Type [male], Ken- tucky (MCZ)," but no specimen was labelled lectotype. roseticola Frev and Boll, 1873: 223, Tischeria; SYN- TYl'l S (2 MCZ 1350; United States: Massachu- setts, "Cambr.B."; See comments under Tischeria aenea regarding type locality. tinctoriella Chambers, 1875a: 108, Tischeria; SYN- T, I'l MCZ 150; United States: Kentucky, Cham- bers; Tischeria quercitella Clemens; Two addi- tional specimens, labelled only "Kentucky./ ( hambers." ma) be syntypes also. SUPERFAMILY INCURVARIOIDEA Family Incurvariidae alba Zeller, 1873: 232, Tegeticula; LECTOTYPE male. MC:Z 2922; United States: Texas, Dallas, Boll; Tegeticula yuccasella (Riley); Lectotype designat- ed b) Davis (1967: 51). aureovireus Dietz. 1905: 39, Incurvaria; HOLO- TYPE, MCZ 2ST4; United States: Pennsylvania, Hazleton, [23] June 1899, Dietz; Phylloporia bis- trigclla (Haworth). bella Chambers. 1873: 73, Adela; SYNTYPE female, MCZ 1 102; United States: Kentucky, May, Cham- bers. Adela caeruleella Walker. chalybeis Zeller. 1873: 226, Adela; HOLOTYPE male, M( IZ 32960; United States: Texas, Dallas, Boll; Ade- la caeruleella Walker. dietziella Kearfott, 1908: 187, fig. 6, Incurvaria?; SYNTYPES (4), MCZ 14236; United States: New Jerse\ . Essex ( iounty. 30 May 1907, W. D. Kearfott; Chalceopla dietziella (Kearfott). paradoxioa Chambers, 1878e: 149, Hyponomeuta [sic]; LECTOTYPE male, MCZ 32959; United States: Colorado, "nine miles north of Colorado Springs and thence 5 miles east of the mountains' ; Pro- doxus quinquepunctellus (Chambers); Lectotype and Imir paralectotypes also in MCZ designated by Davis L967 76). 5-punctella Chambers, L875d: 7, Hyponomeuta [sic]; LECTOTYPE Female, MCZ 1413; United States: Texas, Bosque County; Prodoxus quinquepunctel- lus Chambers); Lectotype designated by Davis 1967 7". rheumapterella Dietz 1905: 37, pi. I: fig. 4. Incur- saria; LECTOTYPE Female, MCZ 2873; United States Colorado, Durango; Prodoxus coloradensis Rile) Lectotype and paralectotype also in MCZ designated b) Davis I L967 83 Family Heliozelidae aesella Chambers 1877a 108, Heliozela; HOLO- TYPE, MCZ L512; l nited states Kentucky, near Covington, 24 April, Chambers; Head and right front wing only. ampelopsifoliella Chambers, 1874a: 168, Antispila; PSEUDOTYPES (3), MCZ 1367; United States: Kentucky, Chambers; Chambers (1874: 168) states "known only in the larval state," so these cannot be types. One of the three specimens is missing from its minuten. gracilis Zeller, 1873: 314, Heliozela; HOLOTYPE male, MCZ 1351; United States: Texas, Dallas, Boll. viticordifoliella Chambers, 1874a: 168, Antispila; SYNTYPES (2), MCZ 1368; United States: Ken- tucky, Chambers. SUPERFAMILY TINEOIDEA Family Tineidae Donald R. Davis apachella Dietz, 1905: 7, Amydria; SYNTYPE fe- male, MCZ 2904; United States: Arizona, Catal[ina] Springs; The type series of apachella is mixed. This specimen (MCZ 2904) is curvistrigella Dietz. One female syntype (Williams, Arizona) of apachella is in USNM. apicisignatella Dietz, 1905: 65, Tinea; SYNTYPES (2 of 3), MCZ 2862; United States: New Hampshire, Hampton and Pennsylvania, Hazleton; Nemapo- gon variatella (Clemens); Both MCZ syntypes lack abdomens and one lacks a forewing. An additional female syntype is in USNM. approximatella Dietz, 1905: 27, Scardia; PARALEC- TOTYPES (8), MCZ 2889 and 1 PSEUDOTYPE, MCZ 2889; United States: New Jersey, Essex Coun- ty (Kearfott); and Pennsylvania, Hazleton, and Mauch Chunk; Scardiella approximatella (Dietz); One Hazleton, Pennsylvania, "syntype" bears a la- bel date of "6/28 06" and therefore may not be a true type. Lectotype male and paralectotype fe- male in USNM, designated by Robinson (1986: 109). argentinotella Chambers, 1876b: 104, Semele; SYN- TYPE female, MCZ 1400; United States: Kentucky, June, Chambers; Homosetia argentinotella (Chambers); No other syntypes are known. arizonella Dietz. 1905: 6, Amydria; SYNTYPE male, MCZ 2903; United States: Arizona, Huachuca; Ab- domen missing. Another male syntype (Phoenix, Arizona) in USNM. auricristatella Chambers, 1873a: 110, Pitys; SY'N- TYPE female, MCZ 1397; United States: Kentucky, Chambers; Homosetia auricristatella (Chambers); No other syntypes are known. auristrigella Chambers, 1873a: 86, Tinea; SYNTYPE male, MCZ 14943; United States: Kentucky, July, Chambers; Isocorypha mcdiostriatella Clemens; In MCZ Microlepidoptera Types • Miller and Hodges 51 poor condition and glued to a paper point. No other syntypes are known. auropulvella Chambers, 1873a: 90, Tinea; SYN- TYPES (1 male, 2 females), MCZ 1391; United States: Kentucky, July; Nemapogon auropulvella (Chambers); Two additional male syntypes in USNM. aurosuffusella Chambers, 1873a: 87, Tinea; SYN- TYPE female, MCZ 1394; United States: Kentucky, Chambers; Hybroma servulella Clemens; No other syntypes are known. behrensella Chambers, 1875b: 249, Tinea; HOLO- TYPE, presumed lost; United States: Californa, San Francisco, J. Behrens. bimaculella Chambers, 1873a: 87, Tinea; SYN- TYPES (3 males), MCZ 1388; United States: Ken- tucky, Chambers; Tinea mandarinella Dietz; Chambers' name is preoccupied by Thunberg, 1794. bipunctella Dietz, 1905: 77, Progona; HOLOTYPE female, [no MCZ number]; United States: Florida, Hastings; Mea bipunctella (Dietz); Specimen most- ly destroyed, only fragments of mesothorax adher- ing to pin. brevipennella Dietz, 1905: 5, Amydria; SYNTYPES (3 males), MCZ 2901; United States: District of Columbia, Washington, A. Busck; Maryland, Plum- mer's [sic] Island, A. Busck; Additional syntypes in USNM (13) and BMNH (3). busckiella Dietz, 1905: 12, Paraplesia; PSEUDO- TYPE female, MCZ 2892; United States: Arizona, Catal[ina] Springs, [E. A. Schwarz]; Hypoplesia busckiella (Dietz); Dietz (1905) states that the only material examined by him was a single male (there- fore the holotype) in the USNM. Consequently the MCZ "cotype" and a male ■'cotype" in LACM are not true types. carbonella Dietz, 1905: 30, Abacobia; SYNTYPES (1 male, 4 females), MCZ 2871; United States: Penn- sylvania, Hazeleton, W. G. Dietz; Elatobia carbo- nella (Dietz); Two additional syntypes (male and female) in USNM. cariosella Dietz, 1905: 17, Epilegis; SYNTYPE male, MCZ 2898; United States: California, Kaweah; Se- tomorpha rutella Zeller; Dietz described this species from two males from Kaweah, California. A female "cotype'' in MCZ and another female "cotype'' in LACM, both of which bear no locality labels, are not types. chrysocomella Dietz, 1905: 43, Isocorypha; HO- LOTYPE, MCZ 2883; United States: Kansas, On- aga, [H. Kahl]. clemensella Chambers, 1873a: 174, Xylesthia; SYN- TYPES (3 males, 3 females, 2 unknown), MCZ 1384; United States: Kentucky, [Chambers]; Xylesthia pruniramiella Clemens; Four MCZ syntypes are badly damaged with abdomens missing from three. Three additional male syntypes in USNM. coloradella Diet/ L905: 6, Amydria; SI Vl'YPES (1 male, 3 females), MCZ 2902; United States: Ari- zona, Santa Rita Mountains; California, Kaweah; and Colorado, Durango; Amydria effrentella Cle- mens; Two additional (male, female) syntypes in USNM. confusella Dietz, 1905: 8, Amydria; SYNTYPES (6 females), MCZ 290(i; United States: California, Pas- adena; Two additional tcinale syntypes in USNM. costotristgella Chambers, 1873a: 87, Tinea; SYN- TYPE male, MCZ 14947; United States: Kentucky, August and September, Chambers; Nemapogon granella (Linnaeus); Specimen in poor condition; right forew ing and abdomen missing. No other syn- types are known. crescentella Kearfott, 1907b: 9, Amydria; SYNTYPE male, MCZ 14235; United States: Arizona, Pima County, Baboquivari Mountains; Acrolophus cres- centella (Kearfott); Three additional male syntypes in USNM. Presumably the fifth remaining syntype is in University of Kansas collection, Lawrence, Kansas. cristatella Chambers, 1875b: 243, Semele; SYNTYPE male, MCZ 1401; United States: Kentucky, June, Chambers; Homosetia cristatella (Chambers); No other syntypes known. croceoverticella Chambers, 1876b: 106, Tinea; SYN- TYPE male, MCZ 1393; United States: Kentucky, Chambers; No other syntypes known. cruciferella Dietz, 1905: 14, Paraneura; SYNTYPES (2 males), MCZ 2895; United States: California, Mountain View; Lindera tessellatella (Blanchard); One male "cotype" each in LACM and USNM are pseudotypes. curviliniella Dietz, 1905: 71, Homostinea; SYN- TYPES (3 males), MCZ 2866; United States: District of Columbia, [Washington, A. Busck]; Kansas, On- aga; Louisiana, Vowells Mill; and, Missouri, St. Louis; Louisiana syntype is lost (only pin and labels re- maining in MCZ). One male syntype (Washington, DC.) is also present in LACM, but a second female "cotype" from Kentucky is a pseudotype. curvistrigella Deitz, 1905: 8, Amydria; SYNTYPE female, MCZ 2905; United States: Arizona, Phoe- nix; Additional male syntype in USNM. dyarella Dietz, 1905: 11, Amydria; HOLOTYPE fe- male, MCZ 14234; United States: Pennsylvania, Hazleton; Two female paratypes also exist, one in MCZ and one in USNM. ehrhornella Dietz, 1905: 13, Paraneura; SYNTYPES (2 males), MCZ 2894; United States: California, Mountain View: Lindera tessellatella (Blanchard). 52 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 eunitariaeella Chambers, L873a 85, Tinea; SYN- rYPES 2 males 2 Females), MCZ 1398; United States Kentucky, Chambers; Eudarcia eunitar- iaeella (< !hambers fasciella Chambers L873a: 111, Pitys; SYNTYPE fe- male, MCZ L347; I nited States Kentucky, Cham- bers; Homosetia fasciella (Chambers); No other syntypes are known Another female in extremely worn condition bearing Chambers' Kentucky label exists in MCZ, but it shows no other indications of being a s> ut\ pe. doridella Diet/. L905: 77. Progona; HOLOTYPE fe- male MCZ 2869; I nited States: Florida. Hastings; Xfea skinnerella (Dietz); Holotype erroneously stat- ed to be a male by Dietz. fractiliniella Dietz, L905; 17 Apotomia; SYNTYPE male. MCZ 2899; I nited States: California, Pasa- dena. Setomorpha rutella Zeller; Additional female s\ lit > pe in I S\\l frigidella Packard. 1867: 62, Oecophora; SYNTYPES 2females ,MCZ 1549; Canada: Labrador, Caribou Mand. Square Island; Niditinea fuscella (Lin- naeus); Both syntypes in ver) poor condition with the abdomen missing from one. fulvisuffusella Dietz, 1901 68, Tinea; HOLOTYPE male MCZ 2864; I nited States: New Hampshire, I [ampton; Nemapogon variatella (Clemens); A male pseudotype from the same locality in USNM. fuscocristatella Chambers, L873a: 111, Pitys; SYN- I "\ I'l S presu d lost; t nited States: Kentucky; Homosetia fuscocristatella (Chambers); Dietz L905 examined the "type" of this species in the MCZ and considered it a synonym of Homosetia miscecristatella Chambers. fu-cofasciella Chambers, 1875b: 257. Euplocamus CO: LECTOTYPE female, MCZ 1385; United States Kentucky, Chambers; Moniescardia fus- cofasciella Chambers); l.ectotype designated by Robinson (1986: 76). fuscomaculella Chambers, lS73a: 88, Tinea; SY'N- n I'l male M< /. 14946; United States: Kentucky, Chambers; Nemapogon granella (Linnaeus); No othei s\ nt\ pes know n. geniculatella Diet/ L905: 62, Tinea; SYNTYPES (2 females), MCZ 2861 I nited States: California, Oc- cidental and Pasadena; \emapogon geniculatella Dietz Deposition ol an additional syntype from Kaweah, California, is unknown griseella Chambers 1873a 88, Tinea; SYNTYPES (2 males MCZ L389 I nited States: Kentucky, Chambers Siditinea Jus, din \ Linnaeus); One syn- type in ver) pooi condition with head, abdomen. i left toiew ing missing hybromella< hambei L874a 51, Oenoe; SYNTYPE m I nited Slates: Kentucky. Cham- bei mi. inamoenella Zeller, 1873: 224, Setomorpha; HO- LOTYPE male, MCZ 2882; United States: Texas, Dallas, Boll; Setomorpha rutella Zeller. interstitiella Dietz, 1905: 68, Tinea; HOLOTYPE male, MCZ 2865; United States: Georgia, Forsyth; Nemapogon interstitiella (Dietz). irrorella Dietz. 1905: 34, Monopis; SYNTYPES (4 males. 3 females), MCZ 2872; United States: Penn- sylvania, Hazleton and Mauch Chunk; Monopis marginistrigella (Chambers); Additional syntypes in LACM (1 male, 1 female) and USNM (2 males). Deposition of syntype(s) from New York unknown. maculatella Dietz, 1905: 84, Homosetia; SYNTYPES (2 males), MCZ 2851; United States: California, Placer County; Pennsylvania, Hazleton; Homose- tia marginimaculella (Chambers); Syntype from Placer County, California, missing abdomen. majorella Dietz, 1905: 15, Setomorpha; SYNTYPES (2 females), MCZ 2896; United States: California, Pasadena; Setomorpha rutella Zeller. marginistrigella Chambers, 1873a: 88, Tinea; SYN- TYPE, MCZ 14950; United States: Kentucky, Chambers; Monopis marginistrigella (Chambers); Specimen in very poor condition, glued to a point with only the head, prothorax, and right forewing. No other syntypes known. margoriella Dietz, 1905: 11, Amydria; SYNTYPES (4 males), MCZ 2891; United States: Florida; Kan- sas, Lawrence; Texas, San Antonio; The type series appears to be mixed, with the eastern records most likely representing Amydria dyarella Dietz. Dietz (1905) mentions syntypic material in USNM, but none has been found. minutipulvella Chambers, 1875e; 212, Tinea; SYN- TYPE female, MCZ 1390; Canada; Nemapogon acapnopennella (Clemens); Another female syn- type in USNM. miriamella Dietz, 1905: 90, Leucomele; SYNTYPES (4 males, 2 females), MCZ 2854; United States: Maryland, Plummers [sic] Island; Pennsylvania, Hazleton and Mauch Chunk; Two additional male syntypes in USNM and one male syntype in BMNH. miscecristatella Chambers, 1873a: 111, Pitys; SY'N- TYPE female, MCZ 1395; United States: Kentucky, Chambers; Homosetia miscecristatella (Cham- bers); No other syntypes known. misceella Chambers, 1873a: 86, Tinea; SYNTYPE(S), presumed lost; United States: Kentucky; Dietz (1905) reported the poorly preserved "type in MCZ. molybdanella Dietz, 1905: 61, Tinea; SYNTYPES (2 females), MCZ 2880; United States: California, Pas- adena; S'emapogoti molybdanella (Dietz). multimaculella Chambers, 1878c: 89, Gelechia?; SYNTYPES (2 males), MCZ 1451; United States: Texas; Setomorpha rutella Zeller. MCZ Microlepidoptera Types • Miller and Hodges 53 multistriatella Dietz, 1905: 59, Tinea; SYNTYPES (2 females), MCZ 2878; [Canada: Toronto]; Nema- pogon multistriatella (Dietz); A female syntype is also present in the LACM (Plummers Island, Mary- land). The MCZ syntypes lack locality labels. nepotella Dietz, 1905: 21, Epichaeta; SYNTYPES (2 females), MCZ 2887; United States: California, Pas- adena; Apreta paradoxella Dietz. nigroatomella Dietz, 1905: 70, Tinea; HOLOTYPE, MCZ 2865; United States: New Jersey, Montclair, W. D. Kearfott; Nemapogon granella (Linnaeus). obliquella Dietz, 1905: 10, Amydria; SYNTYPES (2 females), MCZ 2909; United States: California, Los Angeles County, Pasadena; Additional male syn- type (Los Angeles County) in USNM. obscurella Dietz, 1905: 82, Homosetia; HOLOTYPE male, MCZ 14945; United States: Kansas, Onaga; Homosetia bifasciella (Chambers). oecidentella Dietz, 1905: 9, Amydria onagella; SYN- TYPES, presumed lost; United States: California, Mountain View and Pasadena. oecidentella Chambers, 1880b: 193, Tinea tapetzella; SYNTYPES (1 male, 1 female?), MCZ 1392; United States: California, San Francisco; Tinea oeciden- tella Chambers; According to Chambers, the type series consisted of three specimens. The location of the third syntype is unknown. onagella Dietz, 1905: 9, Amydria; SYNTYPES (2 males), MCZ 2908; United States: Kansas, Onaga; One syntype missing right wings. operosella Zeller, 1873: 223, Setomorpha; HOLO- TYPE male, MCZ 2881; United States: Texas, Dal- las, Boll; Setomorpha rutella Zeller; Abdomen and left wings missing. ophrionella Dietz, 1905: 56, Tinea; HOLOTYPE male, MCZ 2875; United States: New York, Ithaca; Nemapogon ophrionella (Dietz); Left wings miss- ing. orleansella Chambers, 1873a: 85, Tinea; SYNTYPE male, MCZ 14949; United States: Louisiana, New Orleans; Niditinea orleansella (Chambers); Syn- type in poor condition, unspread, and glued to a point. No other syntypes known. pandurella Dietz, 1905: 8, Amydria; SYNTYPES (2 males), MCZ 2907; United States: Arizona, Phoenix; California, Pasadena; Amydria curvistrigella Dietz; Dietz stated that the type series consisted of "two specimens, [male] and [female], in my collection." A second specimen in MCZ is hereby considered a syntype even though it is also a male (i.e., sex mis- determined by Dietz) and bears no type labels. Two "cotypes" in USNM must thereby be regarded as pseudotypes. One of these (from Pasadena) is a female, but it was misidentified and is A. conjusella Dietz. paradoxella Diet/. 1905: 21, Apreta; HOLOTYPE female, MCZ 2186; United States: California, Pas- adena; Dietz ( 1905) inisdetennined the holot) pe as a male. rileyi Dietz, 1905: 59, Tinea; S^i VH PES (3 lemal, MCZ 2879; United States: Pennsylvania, Hazleton; District of Columbia; Florida Hastings; Nemapo- gon rileyi (Dietz); One female syntype collected by Busck in Washington, D.C., and loaned to Dietz has been returned to USNM. The 25 specimens from Hastings, Florida (reared by Rileyi were re- ferred to by Dietz in the original description. roburella Dietz, 1905; 58, Tinea; HOLOTYPE male, MCZ 2877; United States: New Jersey, Essex Coun- ty [Park], VV. D. Kearfott; Nemapogon roburella (Dietz). ruderella Zeller, 1873: 225, Setomorpha; HOLO- TYPE male, MCZ 14233; United States: Texas, Dal- las, Boll; Setomorpha rutella Zeller. scardina Zeller, 1873; 215, Anaphora; SYNTYPES (2 males), MCZ 33318; United States: Texas, Dallas, Boll; Acrolophus popeanella (Clemens); Two ad- ditional male syntypes in BMNH ("Carolina" and Texas). septemstrigella Chambers, 1878c: 79, Tinea; HO- LOTYPE male, MCZ 1386; United States: Texas; Augolychna septemstrigella (Chambers). sepulchrella Dietz, 1905: 74, Tryptodema; SYN- TYPE male, MCZ 2867; United States: Maryland, Plummer's [sic] Island, A. Busck; Three additional syntypes (2 males, 1 female) in USNM. sigmoidella Dietz, 1905: 16, Setomorpha; SYN- TYPES (2 males), MCZ 2897; United States: Col- orado, Glenwood Springs, Pueblo; Setomorpha ru- tella Zeller; One additional male syntype each in USNM and LACM. simulella Dietz. 1905: 13, Paraneura; SYNTYPE fe- male, MCZ 2893; United States: California, Folsom and Los Angeles; Lindera tessellatella Blanchard; A second syntype (Los Angeles) in LACM skinnerella Dietz, 1905: 76, Progona; HOLOTYPE male, MCZ 2868; United States: New Jersey, Cald- well; Mea skinnerella (Dietz). straminiella Chambers, 1873a: 86, Tinea; SYN- TYPE(S), presumed lost; United States: Kentucky, June. texanella Chambers, 1878c: 79, Anaphora; SYN- TYPE male, MCZ 1383; United States; Texas: \. rolophus texanella (Chambers); No other syntypes known. ihoracestrigella Chambers, 1876b: 106, Tinea; SYN- TYPE(S), presumed lost; United States: [type lo- cality not stated]. transversestrigella Dietz. 1905: 20, Semiota; SYN- TYPES (8 males), MCZ 2900; United States: Cali- 54 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 fornia, Pasadena: Setomorpha rutella Zeller; Four additional male syntypes in I SNM. tuscanella Dietz, L905: 53, Tinea; S"V NTi PE female, MCZ 2884; I nited States: Arizona, Tuscan [Tuc- son]. Tinea occidentella Chambers; No other syn- types know n unomaculella Chambers, 1875b: 258, Tinea; SYN- H PE female, M< :Z 1 387; I nited States: Texas; No other S) nt> pes know n. vicinella Dietz, L905 55 Tinea; HOLOTYPE fe- male. MCZ 2885; I nited States Florida, Gotha; Ceratophaga vicinella (Dietz); Sex given as male in original description. visaliella Chambers, L873a: 113, Cyane; SYNTYPE male. MCZ L4974; I nited States: Kentucky, Yisa- lia. Chambers; Choropleca visaliella (Chambers); The single know n s\ ntype is l)adl\ broken with the remaining parts glued to a point. xanthostictella Dietz. 1905 56, Tinea; HOLOTYPE male. MCZ 2S7(i; United States: Georgia, Forsyth. yumaella Kearfott, 1907b: 6, Plutella; SYNTYPE male ] ol L6), M< / 1 I 858; United States: Arizona, Yuma County, Desert; Dyotopasta yumaella (Kearfott); Nme Yuma Count) syntypes in USNM, some of the remaining syntypes (other localities) at Uni- versity ol Kansas. Family Lyonetiidae albella Chambers, 1871a: 23, Cemiostoma; SYN- TYPES 2 ol t . MCZ 1306; United States: Ken- tucks. Chambers; Paraleucoptera albella (Cham- bers); One possible syntype in USNM (type 518). albella Chambers. ISTTd: 140, Eurynome; SYN- TYPES (2), MCZ 1312; United States: Colorado, near Edgerton, elevation about 6,500 feet; Philon- ome albella < Ihambersi: Two possible syntypes in I \( \1 albicapitella Chambers, I875e: 125. Bucculatrix; SYNTYPE (?), M( / I 1962; Canada; Bucculatrix agnclla Chambers; Missing hind wings and abdo- men. Braun L963 79); considered this name "an apparent transposition of s\ I lables" of capitealbella < fiambers. One possible syntype in USNM (type 5776 alniella Chambers, L875c: 303, Lyonetia; S^ V Tl PI S 9 M( Z l.l, l nited States Colorado, "along Grand River, Clear Creek. Fall River and I ontain-qui-Bouille, in all its stages, up to 9,000 feel altitude Vnother syntype in LACM. ambro-.ia<-l«»ii.>lla Chambers, 1875a: 119. Buccula- trix; SYNTYPES 2 MCZ L308; I nited States; Kentucky, bred from leaves ol Ambrosia trifida I innai us < Ihambers apicistrigella Chambers, 1875a: 105, Lyonetia; SYN- T"5 PE, MCZ 1314; United States: Kentucky, Au- gust, Chambers; Lyonetia speculella Clemens; Missing abdomen. canadensisella Chambers, 1875e: 146, Bucculatrix; SYNTYPE, MCZ 1307; Canada; Missing abdomen. According to Braun (1963: 147) there is a female type in USNM (type 5775), but we could not locate it. capitealbella Chambers, 1873a: 150, Bucculatrix; HOLOTYPE, MCZ 14961; United States: Ken- tucky, Chambers; Bucculatrix agnella Clemens; One "syntype" in USNM (type 519). clemensella Chambers, 1874a: 97, Philonome; SYN- TYPES 7 (2 missing from pins), MCZ 1311; United States: Kentucky, Chambers; One possible syntype in USNM (type 522). curvilineatella Packard, 1869: 354, pi. 8, fig. 16, Lith- ocolletis; SYNTYPE, MCZ 1347; [United States]; Bucculatrix pomifoliella Clemens; Broken and missing head. fuscoscapulella Chambers, 1878c: 104, Acanthoc- nemes; SYNTYPE, MCZ 1530; United States: Tex- as, Bosque County; Missing head and forewings. inornatella Chambers, 1880b: 188, Eulyonetia; SYN- TYPE, MCZ 1353; United States: Texas, Belfrage; Missing left wings and abdomen. litigiosella Zeller, 1875: 354, Bucculatrix; SYNTYPE (1 of 2) female (not male as stated by Zeller), MCZ 14959; United States: Texas, Dallas, Boll. luteella Chambers, 1873a: 151, Bucculatrix; SYN- TYPES (7), MCZ 1501; United States: Kentucky, March, Chambers; Braun (1963: 153) stated that some of these syntypes are Bucculatrix packardella, and that there is a female syntype at USNM (type 520) and male syntype at ANSP. luteella Chambers, 1875c: 304, Eurynome; HOLO- TYPE, MCZ 14964; United States: Colorado, Span- ish Bar, 4 July; Philonome luteella (Chambers). magnella Chambers, 1875d: 54, Bucculatrix; SYN- TYPES (6), MCZ 1309; United States: Kentucky, Chambers; Braun (1963: 42-43) listed only one fe- male "type" in MCZ, and two "paratypes in USNM. obscurofasciella Chambers, 1873a: 150, Bucculatrix; S"i NTYPE female, MCZ 1500; United States: Ken- tuck), 23 May, Chambers; Bucculatrix trijasciella Clemens; Braun (1963: 136-137) noted a female syntype in USNM (type 521). packardella Chambers, 1873a: 151, Bucculatrix; s^ \ TYPES" (2), MCZ 14960; United States: Ken- tucks . ( :hambers; Two specimens glued to the same point, labelled only "Kentucky./ Chambers." and MCZ Microlepidoptera Types • Miller and Hodges 55 "Type/ 14960." Braun (1963: 129) stated "Two specimens thus named, presumably by Chambers, in the Museum of Comparative Zoology, but not labeled types, do not represent this species." staintonella Chambers, 1878d: 133, Bucculatrix; LECTOTYPE male, MCZ 1310; United States: Col- orado, Edgerton, elevation 6,000 feet; Originally described by Chambers (1877d: 141) as B. albella, a name preoccupied by Stainton. Lectotype des- ignated by Braun (1963: 74). SUPERFAMILY GELECHIOIDEA Family Oecophoridae [Note: The Walsingham syntypes listed be- low were received by Chambers from Walsingham. Lectotypes should be des- ignated from BMNH specimens.] albaciliaeella Chambers, 1878b: 77, Strobisia; HO- LOTYPE, MCZ 1548; United States: Ohio, Cincin- nati; Menesta tortriciformella Clemens; Head and thorax only. albella Chambers, 1874b: 235, Harpalyce; SYN- TYPES (5), MCZ 1417; United States: Texas, Clif- ton, Belfrage; Durrantia piperatella (Zeller). apioipunctella Chambers, 1875d: 8, Hyponomeuta; LECTOTYPE male, MCZ 1404; United States: Texas, Basque [sic] County, Belfrage; Ethmia ap- icipunctella (Chambers); Lectotype designated by Powell (1973: 88), also paralectotype in MCZ. argillacea Walsingham, 1881: 313, pi. XXXVI, fig. 2, Depressaria;; SYNTYPE, MCZ 14975; United States: California, Tehama County, Newville; Agonopterix argillacea (Walsingham). arnieella Walsingham, 1881: 314, pi. XXXVI, fig. 3, Depressaria;; SYNTYPE, MCZ 14976; United States: California, Mount Shasta; Agonopterix ar- nieella (Walsingham). bicostomaculella Chambers, 1877a: 127, Gelechia; SYNTYPE male, MCZ 1461; United States: Colo- rado, Edgerton, July; Taygete decemmaculella (Chambers); Hodges (1986: 6) transferred decem- maculella to Oecophoridae. boreasella Chambers, 1873a: 189, Oecophora; LEC- TOTYPE male, MCZ 1553; United States: Ken- tucky; Decantha boreasella (Chambers); Lectotype designated by Hodges (1974: 104). canusella Chambers, 1874b: 235, Harpalyce; SYN- TYPE, MCZ 1415; United States: Texas, [13/8]; Antaeotricha humilis (Zeller). chrysurella Dietz, 1905: 42, Breckenridgia [sic]; HO- LOTYPE male, MCZ 33274; United States: [South- west] Colorado, ["6-28-89"]; Ethmia albistrigella (Walsingham). clemensella Chambers, 1876b: 173, Gelechia; SYN- TYPES (5), MCZ 1488; United States: Pennsylva- nia, Easton; Agono])tcrix clemensella (Chambers). coryliella Chambers, 1875b: 242, llyale; SYNTYPE, MCZ 14974; United States: Kentucky, Covington, Chambers; Menesta tortriciformella Clemens. cressonella Chambers, 1878c: 86, Cryptolechia; LECTOTYPE male, MCZ 1420: I nited States: Texas; Psilocorsis cryptolechiella ^Chambers); Lec- totype designated by Hodges (1974: 92), also 2 paralectotypes in MCZ. difficilisella Chambers, 1872a: 66, Evagora: S't V TYPE male, MCZ 1528; United States: Kentucky, Chambers; Taygete attributella (\\ alker); Hodges (1986: 6) transferred attributella to Oecophoridae. discostrigella Chambers, 1877d: 122, Anesychia; LECTOTYPE male, MCZ 1421; United States: Col- orado, Edgerton, 6,500 feet; Ethmia discostrigella (Chambers); Lectotype designated by Powell (1973: 93), also 4 paralectotypes in MCZ. eupatoriiella Chambers, 1878c: 82, Depressaria; SYNTYPE, MCZ 1432; United States: Kentucky, Chambers; Agonopterix eupatoriiella (Chambers). faginella Chambers, 1872a: 131, Hagno; LECTO- TYPE male, MCZ 1419; United States: Kentucky, Chambers; Psilocorsis cryptolechiella (Chambers); Lectotype designated by Hodges (1974: 92), also 4 paralectotypes in MCZ. fernaldella Chambers, 1878c: 82, Depressaria; Pos- sible SYNTYPES (2); United States: Maine; Ma- chimia tentoriferella Clemens; Two specimens with no type labels may be types: one is labelled "Me"; "47" [pencil]; "fernaldella/Chb" [Chambers' pen]; "Machimia/(tentoriferella?)/AB May 1900"; the other "L.I." [?, pencil, illegible]; "Ken- tucky.[crossed out]/Chambers."; "22" [Chambers' hand]. fernaldella Riley, 1889: 155, Setiostoma; SYNTYPES (2 of 12), MCZ 11907; United States: California. Los Angeles County; Other syntypes in USNM; Lectotype should be designated from USNM spec- imen. hagenella Chambers, 1878c: 80, Anesychia; LEC- TOTYPE male, MCZ 1422; United States: Texas, Bosque County; Ethmia hagenella (Chambers); Lectotype designated by Powell (1973: 110), also paralectotype in MCZ. lithosina Zeller, 1873: 244, Cryptolechia; HOLO- TYPE male, MCZ 1717; United States: Texas, Dal- las, Boll; Antaeotricha unipunctclla (Clemens). longimaculella Chambers, 1872a: 43, Hyponomeuta; LECTOTYPE male, MCZ 1403; United States: Kentucky, Chambers; Ethmia longimaculella (Chambers); Lectotype designated by Powell (1973: 178), also 4 paralectotypes in MCZ. 56 i Museum of Comparative Zoology, Vol. 152, No. 2 mirusella Chambers, L874b; 233, Anesychia; LEC- TOTYP1 male, MCZ L423; I nited States: Texas, [frage; Ethmia mirusella (Chambers); Lectotype designated l>\ Powell (1973: 193), also 4 paralec- tot) pes in \K / maltipunctella Chambers, L874b: 233, Anesychia; LECTOTYPE male. MCZ 1425; United States: Texas Wan.. Belfrage; Ethmia semilugens (Zell- ei . Lectotype designated In Powell (1973: 86), also L3 paralectot) pes in NH :/.. nebeculosa Zeller. L873: 245. fig. 12, Cryptolechia; SYNTYPES (4), MCZ 1720; United States: Texas, Dallas. Boll; \ntaeotricha humilis (Zeller). uovi-mundi Walsingham, 1881: 318, Depressaria; SYNTYPE, MCZ 14971; liiited States: California and Oregon; Exaeretia thoracenigraeella (Cham- bers). nubilerella Walsingham, 1881: 316, pi. XXXVI, fig. i- Depressaria; SYNTYPE, MCZ 14977; United States: Oregon, Rogue River; Agonopterix nubi- ferella (Walsingham). obscuromaculella Chambers, 1878c: 86, Cryptole- chia?; SYNTYPE, MCZ 1418; United States: Texas, Bosque County, [11/8.]; Inga obscuromaculella Chambers;. Another syntype in USNM (Hodg 197 1 L02 es. piperatella Zeller. 1873: 239, Cryptolechia; HO- LOTYPE male. MCZ 1719; United States: Texas, Dallas. Boll; Durrantia piperatella (Zeller). posticella Walsingham, 1881: 315, pi. XXXVI, fig. 5, Depressaria; SYNTYPE, MCZ 14972; United States: ( alifornia and Oregon; Agonopterix posticella \\ alsingham). quinqueferella Walsingham, 1881: 322. Glyphipte- r>x [sic]; PARALECTOTYPE (?), MCZ [no num- bei | I nited States: [California]; Fabiola quinque- ferella (Walsingham); Designated a paralectotype b> Heppner ll)TS 19), but, as noted by Heppner L984 338), it might not be one of the original s\ nt\ pes. shaleriella Chambers 1875a: 1 14, Oecophora; SYN- I 5 PI \l( ./. 1512: I nited State's: Kentucky, Cham- bers Fabiola shaleriella (Chambers); Head and thorax onl) texanella Chambers, 1880b 180, Hyponomeuta; I I < ion PI MCZ I 106; I nited States: Texas; Ethmia zelleriella Chambers); Lectotype desig- nated b) Powell PC ill, ihoracefasciella Chambers, 1875b: 24(i. Gelechia; si MAPI \l( / I 165; I nited States: California; / taeretia ihoracefasciella (Chambers thoracenigraeella < hambers, L875b: 246, Gelechia; si MAPI MCZ L466; I ...led States California; Exaeretia thoracenigraeella (hambers!. I'rag- ments in capsule tortricella Chambers, 1874b: 235, Harpalyce; SYN- TYPES (3), MCZ 1416; United States: Texas; An- taeotricha unipunctella (Clemens). trifurcella Chambers, 1873: 12, Anesychia; LEC- TOTYPE female, MCZ 1426; United States: Ken- tucky, Chambers; Ethmia trifurcella (Chambers); Lectotype designated by Powell (1973: 194). umbraticostella Walsingham, 1881: 318, pi. XXXVI, fig. 8,; Depressaria; SYNTYPE, MCZ 14973; United States: California and Oregon; Exaeretia umbra- ticostella (Walsingham). vestalis Zeller, 1873: 247, Cryptolechia; HOLOTYPE female, MCZ 1718; United States: Texas, Dallas, Boll; Antaeotricha vestalis (Zeller). xanthobasis Zeller, 1875: 324, Setiostoma; HOLO- TYPE male, MCZ 33256; United States: Texas, Dal- las, Boll; Rectiostoma xanthobasis (Zeller). zelleriella Chambers, 1878c: 80, Hyponomeuta; LECTOTYPE female, MCZ 1405; United States: Texas, Bosque County; Ethmia zelleriella (Cham- bers); Lectotype designated by Powell (1973: 113). Family Elachistidae parvipulvella Chambers, 1875d: 56, Elachista; PSEUDOTYPES (4), MCZ 1513; United States: Texas; Braun (1948: 51) stated that none of the 4 MCZ specimens is an Elachista and that the real type is in USNM. staintonella Chambers, 1878c: 96, Elachista; SYN- TYPE male, MCZ 1514; United States: Texas. Family Blastobasidae David Adamski ampla Dietz, 1900a: 103. pi. VI, fig. 1, Ploiophora; SYNTYPES (4 males), MCZ 2923; Adamski gen. slide nos. 2159, 2160, 2161; United States: Penn- sylvania, Hazleton; 1 syntype missing abdomen; 1 male erroneously labeled as "type" (no. 6135) in USNM (Adamski gen. slide no. 2471). angustipennella Dietz, 1900a: 108, Pigritia; SYN- TYPES (5), MCZ 2930; Adamski gen. slide nos. 2166 (male), 2167; (male), 2168 (female); 1 syntype missing abdomen; 1 syntype not dissected; United States: Pennsylvania, Hazleton; 1 male erroneously labeled as "type" (no. 6157) in USNM (Adamski yen. slide no. 2473). annectella Dietz, 1910: 63, pi. IV, fig. 34a, Holcocera zelleriella var.; SYNTYPES (2 females of 3), MCZ 2932; Adamski gen. slide nos. 2150, 2151; United States: Iowa, Iowa City; Louisiana, Vowell's Mill; 1 syntype lost. annulipes Dietz, 1910: 58, Holcocera crescentella x ai ; 1 IOLOTYPE female, MCZ 2950; United States: MCZ Microlepidoptera Types • Miller and Hodges 57 Arizona, Bahoyquivaria [sic] Mountains; Abdomen missing. argyreella Dietz, 1900a: 113, Pseudopigritia; HO- LOTYPE male, MCZ 2920; Adamski gen. slide no. 2201; United States: Pennsylvania, Hazleton. argyrosplendella Dietz, 1910: 22, pi. II, figs. 13, 13a, 13b, Calosima; SYNTYPES (3 of 3), MCZ 2968, 1 male, 1 female; Adamski gen. slide nos. 2059, 2060; United States: Pennsylvania, Hazleton; Florida, Hastings; Louisiana, Vowells Mill; Syntype from Louisiana missing abdomen. arizonella Dietz, 1900a: 109, Pigritia; SYNTYPES (2 of 2 males), MCZ 2912; Adamski gen. slide nos. 2169, 2170; United States: Arizona, Huachuca, and Nochales [sic]. aufugella Zeller, 1873: 301, Blastobasis; HOLO- TYPE, MCZ 14978; United States: Texas, [Dallas], Boll; Pigritia laticapitella Clemens; Abdomen missing. basilarella Dietz, 1900a: 105, pi. VI, fig. 6, Pigritia; SYNTYPES (3 of 3), MCZ 2928; Adamski gen. slide no. 2171 (female); United States: Pennsylvania, Ha- zleton; Kansas, Lawrence; Iowa; 2 syntypes missing abdomens. basipallidella Dietz, 1910: 26, Holcocera dives var.; SYNTYPES (2 of 3), MCZ 2951, 1 male and 1 female; Adamski gen. slide nos. 2082, 2085; United States: Pennsylvania, Hazleton; New Hampshire, Hampton; 1 syntype from Cohasset, Massachusetts in USNM (Adamski gen. slide no. 2434 — male). boreasella Dietz, 1910: 47, pi. Ill, fig. 22, Holcocera; SYNTYPES (4 of 5), MCZ 2960, females; Adamski gen. slide nos. 2064, 2065, 2066; R. B. Selander gen. slide no. 701; United States: New Hampshire, Web- ster; Canada: Montreal; 1 syntype lost. busckiella Dietz, 1910: 36, pi. II, fig. 19, Holcocera; SYNTYPES (2 males of 7), MCZ 2956; Adamski gen. slide nos. 2067, 2068; United States: Maryland, Plummer's [sic] Island, July and August 1903, A. Busck; 5 syntypes in USNM; 1 male dissected (Adamski gen. slide no. 2424), 4 without abdomens. canariella Dietz, 1900a: 118, pi. VII, fig. 17, Dryope; HOLOTYPE, male, MCZ 14226; Adamski gen. slide no. 2212; United States: California, Sonoma Coun- ty; Dryoperia canariella (Dietz). confectella Zeller, 1873: 303, Hypatima; HOLO- TYPE female, MCZ 2083; United States: Texas, [Dallas], Boll; Valentinia confectella (Zeller); Ab- domen missing. confluentella Dietz, 1910: 36, pi. II, fig. 18, Holco- cera; SYNTYPES (2 of 4), MCZ 2955, female; Adamski gen. slide no. 2156; 1 syntype missing abdomen; United States: Pennsylvania, Hazleton; Central New York; Holcocerina confluentella (Dietz); 1 syntype from Cohasset, Massachusetts, in USNM (Adamski gen. slide no. 2129 — female). 1 syntype lost. confusella Dietz. 1900a: 104, pi. VI, fig. 4. Pigritia; SYNTYPES (4), MCZ 2925; Adamski gen. slide nos. 2172 (female), 2173 (male), 2174 (female); 1 syn- type not dissected; United States: Pennsylvania, [Hazleton]; 1 synt> pe Erom [Montclair], New Jersej . in USNM (type no. 6131) (Adamski gen. slide no. 2476 — female). crescentella Dietz, 1910: 57. pi. 1\ . fig. 31. Holco- cera; HOLOTYPE female, MCZ 29 19 Adamski gen. slide no. 2081; United States: Utah, Stockton, T. Spalding. dianella Dietz, 1910: 22, Calosima; HOLOTYPE male, MCZ 2969; Adamski gen. slide no. 2061; United States: Georgia, Forsyth, 1895. discopunctella Dietz, 1900a: 118, pi. VII, fig. 16, Dryope; HOLOTYPE female, MCZ 14223; Adam- ski gen. slide no. 2213; United States: Pennsylvania, Hazleton; Dryoperia discopunctella (Dietz) dives Dietz, 1910: 26, pi. II, fig. 14 (male), Holcocera; SYNTYPES (2 of 7), MCZ 2951, females; Adamski gen. slide nos. 2083, 2084; United States: Pennsyl- vania, Hazleton; Maryland, Plummer's [sic] Island; Canada: Toronto; 3 syntypes from Charleroi, Penn- sylvania in USNM (Adamski gen. slide nos. 2431, 2433 (females) and 2432 (male)). 1 syntype lost. dorsomaculella Dietz, 1900a: 112, pi. VII, fig. 10, Pseudopigritia; SYNTYPES (2 of 2), MCZ 2917, male and female, both specimens missing abdo- mens; United States: Pennsylvania, Hazleton. elyella Dietz, 1910: 49, pi. Ill, fig. 25, Holcocera; SYNTYPES (7 of 17), MCZ 2941; Adamski gen. slide nos. 2086 (female), 2087 (male), 2088 (male), 2089 (female); United States: Connecticut, East River, C R. Ely; Maryland, Plummer's [sic] Island, A. Busck, and Frederick; New Jersey, Essex Coun- ty, W. D. Kearfott; Specimens comprising remain- der of original syntype series cannot be determined. equitella Dietz. 1900a: 112, Pseudopigritia; SYN- TYPES (3 females), MCZ 2918; Adamski gen. slide nos. 2204, 2205; 1 syntype missing adbomen; United States: Pennsylvania, Hazleton. estriatella Dietz, 1910: 28, pi. II, fig. 15, Holcocera; HOLOTYPE male, MCZ 2970; Adamski gen. slide no. 2090; United States: Massachusetts, [Cohasset]. fenyesella Dietz, 1900a: 119. Dryope; HOLOTYPE male, MCZ 14224; Adamski gen. slide no. 2214; United States: California, Pomona, A. Fenyes; Dry- operia fenyesella (Dietz). fidella Dietz, 1900a: 103, pi. VI, figs. 2, 2a (male 2b (female), 2c (male), Ploiophora; SYNTYPES (5), MCZ 2924; Adamski gen. slide nos. 2162 (male), Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 2163 (female), 2164 (female); two syntypes not dis- sected; I nited States: Pennsylvania, Hazleton; 1 syntype and 1 specimen erroneously labeled as "type" (type no. 6126) in USNM (Adamski gen. slide ik's 2472. 2473 — males). Iloridella Dietz. 1910: 17 p] I. fig. 10, Valentinia; S^ NT"} PES (1 ol I MCZ 2966, female (no ab- domen : I nited States: Florida, Cresent City, [male] cone of '/Mima integrifolia; 3 syntypes (male and 2 females) in I SNM; I dissected (Adamski gen. slide no 2115 — male); 1 male erroneously labeled as "cotype," MCZ 2966, from Florida, G. D. Hulst ( lollection. fluxella Zeller. 1873: 301, Blastobasis; HOLOTYPE male, MCZ 1711; Adamski gen. slide no. 2091; I nited States Texas, [Dallas], Boll; Holcocera flux- ella Zeller). fratcrnella Dietz, 1900a: 113, pi. VII, fig. 11, Pseu- dopigritia; HOLOTYPE male, MCZ 2919; Adam- ski gen. slide no. 2207; United States: Pennsylvania, Hazleton. fumerella Dietz. 1910: 35, Holcocera chalcofron- tella var.; SYNTYPES (6), MCZ 2954, females; adamski gen. slide nos. 2074, 2075, 2076; 3 spec- imens not dissected.; I nited States: [Pennsylvania, Hazleton], bred from fruit racemes of sumach. funebra Dietz, 1910 44. Holcocera; SYNTYPES (2 of5 MCZ2092, adamski gen. slide no. 2092 (male); I nited States: Maryland, Plummer's [sic] Island, 1903, A Busck; Pennsylvania, Hazleton; 3 syntypes from Plummer's [sic] Island, Maryland in USNM not dissected ln-<-opurpurella Dietz, 1910: 9, Blastobasis plum- merella var.; HOLOTYPE female, MCZ 2962; Vdamski gen. slide no. 2962; United States: Mary- land, Plummer's [sic] Island: Blastobasis plummer- ella Diet/ fascosuffusella Dietz, 1900a: 117, pi. VII, fig. 13, Dry.pe: SI N I \ PES (2 of 2), MCZ 14222, male .Hid female; Adamski gen. slide nos. 2215, 2216; I nited Slates Missouri, St. Louis; Dryoperia fus- ((/sujjtisctla i Diet/ giganteUa Chambers L876b: 219, Blastobasis; LEC- TOR I'l male MCZ 1551; J. A. Powell gen. slide no 1008 1 nited states: Colorado; Holcocera gi- gantella Chambers . Lectotype and 3 paralecto- types in \l( / designated l>\ Powell (1976). One other spei mien 1 1 ni 1 1 ( iolorado is at MCZ. may not In' |).ut ol original series. gruella Diet/ 1900a 116 |>l VII, figs. 12, 12a, 12b male 12. (female) Dryope; SYNTYPE male, M( / I t22~ Vdamski yen slide no. 221 S: United Pennsylvania, Hazleton; Missouri, Saint Louis; Dryoperia grisella (Dietz); \t least I syntype lost. I syntype from collecti I ( :. V. Riley in I s\M no 71 heidemannella Dietz, 1900a: 111, pi. VII, fig. 9, Epi- gritia; SYNTYPES (2 of 7), MCZ 2916, females; Adamski gen. slide no. 2208; 1 syntype missing left hindwing, right forewing, and abdomen; United States: Pennsylvania, Bedford County, Sulfur Springs, on Abies excelsa, O. Heidemann; Epigritia ochrocomella (Clemens); 1 syntype in USNM (type no. 6133) (Adamski gen. slide no. 2483 — male); 4 syntypes lost. hulstella Dietz, 1910: 7, pi. I, fig. 2, Blastobasis; HOLOTYPE female, MCZ 2977; Adamski gen. slide no. 2042; United States: Texas, G. Hulst. iceryaeella Riley, 1887: 485, Blastobasis; SYNTYPES (4?), possible syntypes in MCZ; Adamski gen. slide nos. 2097, 2098 (1 male, 1 female); United States: California, Los Angeles County, Pasadena; Hol- cocera iceryaeella (Riley); At least 1 syntype from Alameda Co. and Los Angeles Co., California in USNM. illibella Dietz, 1910: 57, pi. Ill, fig. 30, Holcocera; SYNTYPES (3 of 3), MCZ 2948, 2 males; Adamski gen. slide nos. 2101, 2102; United States: Maryland, Frederick; 1 syntype missing abdomen. inclusa Dietz, 1910: 51, pi. Ill, fig. 27, Holcocera; HOLOTYPE male, MCZ 2943; Adamski gen. slide no. 2103; United States: Pennsylvania, Hazleton, 20 July 1904. insulatella Dietz, 1910: 50, pi. Ill, fig. 26, Holcocera; SYNTYPES (2 of 2 males), MCZ 2942; Adamski gen. slide nos. 2104, 2105; United States: Colorado, Glenwood Springs, August. interpunctella Dietz, 1910: 67, pi. IV, fig. 39, Hol- cocera; SYNTYPES (3 of 3), MCZ 2938, 2 males, 1 female; Adamski gen. slide nos. 2107, 2108, 2909; United States: Utah, Stockton, T. Spalding. livorella Zeller, 1873: 299, Blastobasis; HOLOTYPE male, MCZ 2082; Adamski gen. slide no. 2110; United States: Texas, [Dallas], Boll; Holcocera li- vorella (Zeller). luteopulvella Chambers, 1875d; 73, Dryope; SYN- TYPE female, MCZ 1438; Adamski gen. slide no. 2178; United States: Kentucky, Chambers; Pigritia laticapitella Clemens. mediofasciella Dietz, 1900a: 107, pi VI, fig. 5, Pi- gritia; HOLOTYPE female, MCZ 2927; Adamski gen. slide no. 2179; United States: [Pennsylvania, Hazleton], July. melanostriatella Dietz, 1910: 66, pi. IV, fig. 38, Hol- cocera; SYNTYPES (7 of 9), MCZ 2937, 2 males; Vdamski gen. slide nos. 2111, 2112; 5 syntypes miss- ing abdomens; United States: Pennsylvania, Hazle- ton; Connecticut, East River, C. R. Ely; Maryland, Frederick; 2 syntypes from Frederick, Maryland, and Fast River, Connecticut, are lost. messelinella Dietz, 1910: 52, pi. Ill, fig. 29, Holco- MCZ Microlepidoptera Types • Miller and Hodges 59 cera; SYNTYPES (2 of 2 males), MCZ 2945; Adam- ski gen. slide nos. 2115, 2116; United States: Mary- land, Frederick; Florida, Hastings. minnicella Dietz, 1900a: 116, pi. VII, fig. 15, Dryope; SYNTYPES (5 of 5), MCZ 14221; Adamski male gen. slide nos. 2229, 2230 (3 syntypes not dissected); United States: Georgia, Forsyth; Drijoperia min- nicella (Dietz); 1 syntype without "cotype" label, or lost. minorella Dietz, 1910: 34, pi. II, fig. 17e, Holcocera chalcofrontella var.; SYNTYPES (4 of 6), MCZ 2953, female; Adamski gen. slide no. 2077; 3 syn- types not dissected; United States: Pennsylvania, Lewisburg, Chestnut burrs; 2 syntypes lost. murtfeldtella Chambers, 1874a: 50, Dryope; SYN- TYPES (14), MCZ 1437; Adamski gen. slide nos. 2234 (male), 2235 (female); 8 syntypes with dam- aged or missing abdomens; 4 syntypes not dissected; United States: Kentucky; Drijoperia murtfeldtella (Chambers). nubilella Zeller, 1873: 297, fig. 36. Blastobasis; HO- LOTYPE female, MCZ 1714; Adamski gen. slide no. 2054; United States: Texas, [Dallas]; Valentinia glandulella (Riley). obscurella Dietz, 1900a: 110, Pigritia; SYNTYPES (2 of 2), MCZ 2914; Adamski gen. slide no. 2180 (male); 1 syntype missing abdomen; United States: Pennsylvania, Hazleton. occidentella Dietz, 1900a: 115, pi. VII, fig. 14, Dryope; HOLOTYPE female, MCZ 11300; United States: California, Kaweah; Drijoperia occidentella (Dietz); Missing left forewing and abdomen. ochrocephala Dietz, 1910: 31, Holcocera; POSSI- BLE SYNTYPE male, MCZ 2952; Adamski gen. slide no. 2123; United States: Maryland, [Plummers Island]; Male syntype from Aurora, West Virginia in USNM (Adamski gen. slide no. 2455). ornatella Dietz, 1900a: 107, pi. VI, fig. 7, Pigritia; SYNTYPES (7 females, 1 male), MCZ 2929; Adam- ski gen. slide nos. 2182, 2183, 2184, 2186, 2187 females, 2185 male; 2 syntypes not dissected; United States: Pennsylvania, Hazleton; 1 specimen erro- neously labeled as "ornatella" in USNM (Adamski gen. slide no. 2480 — female); 1 erroneously sexed syntype in MCZ. pallidotinctella Dietz, 1900a: 111, pi. VII, fig. 8, 8a (male), 8b (female), Epigritia; SYNTYPES (2 of 3), MCZ 2915; both specimens missing abdomens; United States: Pennsylvania, Hazleton; Epigritia ochromella (Clemens); Male erroneously labeled as cotype in USNM (Adamski gen. slide no. 2482); 1 syntype lost. plagiatella Dietz, 1910: 40, pi. Ill, fig. 20, Holcocera; HOLOTYPE male, MCZ 2957; Adamski gen. slide no. 2124; United States: Arizona, [Williams]. plummerella Diet/.. 1910: 8, pi I, fig. 4, Blastobasis; SYNTYPES (2 males of 6), MCZ 2961; Adamski gen. slide no. 2043 (one specimen missing abdo- men); United States: Maryland, Plummers [sic] Is- land; 3 syntypes in USNM (not dissected); 1 syntype lost. pulchella Dietz, 1910: 20. pi. II, figs. 12, 12a, Euresia; HOLOTYPE male, MCZ 2967; Adamski gen. slide no. 2058; United States: District of Columbia, Washington. purpurella Dietz, 1900a: 105, Pigritia; S^i NTYPES (3 of 3 females), MCZ 2926; Adamski gen. slide nos. 2188, 2189, 2190; United States: Pennsylvania, Hazleton. pusilla Dietz, 1910: 65, pi. IV, fig. 37, Holcocera; HOLOTYPE male, MCZ 2935; Adamski gen. slide no. 2139; United States: Texas, Brownsville. quaintancella Dietz, 1910: 15, pi. I, fig. 9, Valentinia; SYNTYPES (2 females of 6), MCZ 2965; Adamski gen. slide nos. 2055, 2056; United States: locality unknown, bred from apple, Quaintance; 4 syntypes in USNM; 1 dissected (Adamski gen. slide no. 2416 — female), 2 missing abdomens, 1 with broken ovipositor. quisquiliella Zeller, 1873: 298, Blastobasis; SYN- TYPES (3 of 5), MCZ 1712, 1 male and 2 females; Adamski gen. slide nos. 2072, 2073; United States: Texas, [Dallas], Boll; Holcocera chalcofrontella quisquiliella (Zeller); 1 female is missing abdomen. reductella Dietz, 1910: 45, Holcocera funebra var.; SYNTYPES (1 of 2), MCZ 2959, Adamski gen. slide no. 2095 (male); Canada: Manitoba, Aweme, Crid- dle; 1 syntype in USNM (missing abdomen). rufopunctella Dietz, 1910. 65, Holcocera; HOLO- TYPE female, MCZ 2936; Adamski gen. slide no. 2140; United States: Colorado, Denver, Oslar. sagitella Dietz, 1910: 9, pi. I, fig. 5, Blastobasis; HO- LOTYPE male, MCZ 2964; Adamski gen. slide no. 2046; United States: Pennsylvania, Hazleton [Aug. 2, 1898]. sciaphilella Zeller, 1873: 295, fig. 34. Blastobasis; HOLOTYPE male, MCZ 1713; Adamski gen. slide no. 2141; United States: Texas, [Dallas], Boll; Hol- cocera sciaphilella (Zeller). simplicella Dietz, 1910: 9, Blastobasis plummerella var.; HOLOTYPE male, MCZ 2963; Adamski gen slide no. 2045; United States: Iowa, Iowa City; Spec- imen originally misidentified as female. simulella Dietz, 1910: 52, pi. Ill, fig. 28, Holcocera; SYNTYPES (2 of 4), MCZ 2944, female; Adamski gen. slide no. 2143; 1 syntype missing abdomen; United States: Arizona, Williams; Texas Fedora; 1 syntype from Williams, Arizona in USNM (Adam- ski gen. slide no. 2459 — female); syntype from Stockton, Utah lost. 60 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 spoliatella Dietz, 1900: L10, Pigritia; SYNTYPES 2 ol 2 males), M< / 2913; Adamski gen. slide nos. 21lJl. 2192: 1 syntype missing left forewing and abdomen; United States: Pennsylvania, Hazleton; \n additional damaged MCZ specimen is probabl) n.it a syntype. 1 specimen erroneously labeled co- t> pe in L SW1 spoliatella Dietz, 1910: 53, Holcocera messelinella var.; SYNTYPES (3 of 6), MCZ 2946, 3 males; Adamski gen. slide nos. 2117. 2118, 2119; United States Pennsylvania, Hazleton; Holcocera messe- linella Diet/: 3 s\nt\[>es lost including those from New Terse) spretella Dietz. 1910: 58, pi. IV, fig. 32, Holcocera; s\ \ n PE 1 1 ot 4), MCZ 2931; Adamski gen. slide no. 2145 — female: L nited States: Arizona, Phoe- nix: 3 syntypes from Williams, Arizona in USNM \n I 0: 7. pi. I. fie,. 3, Blastobasis; SYNTYP1 S male and female: ^dai slid,-,,.. 2iil7 2048; United States: Texas, bred from Yucca baccata, emerged 18 April 1897; 3 syntypes in USNM (not dissected). zelleriella Dietz, 1910: 62, pi. IV, fig. 34, Holcocera; SYNTYPES (2 of 2 males), MCZ 2081; Adamski gen. slide nos. 2148, 2149; United States: Texas, Dallas; Only 1 male has locality label. Family Momphidae albocapitella Chambers, 1875d: 33, Laverna; SYN- TYPE, MCZ 1374; United States: Texas, Basque [sic] County, September; Mompha murtfeldtella (Chambers). bicristatella Chambers, 1880b: 187, Elachista; SYN- TYPE, MCZ 1355; United States: Texas, Belfrage; Mompha bicristatella (Chambers). cephalonthiella Chambers, 1871b: 221, Laverna; SYNTYPES (6), MCZ 1372; United States: Ken- tucky, Chambers; Mompha cephalonthiella (Chambers). circumscriptella Zeller, 1873: 312, fig. 42, Laverna; PSEUDOTYPE, MCZ 1380; United States: Texas, Dallas, Boll; Mompha circumscriptella (Zeller). coloradella Chambers, 1877d: 136, Laverna(?); SYNTYPE, MCZ 1379; United States: Colorado, Edgerton, July; Mompha coloradella (Chambers). definitella Zeller, 1873: 111, fig. 41, Laverna; SYN- TYPE, MCZ 1377; United States: Texas, Dallas, Boll; Mompha definitella (Zeller). grisseella Chambers, 1875c: 295, Laverna; PSEL - DOTYPES (2), MCZ 14970; Mompha murtfeld- tella (Chambers); MCZ has 2 specimens from Ken- tuck), but description was based on 3 specimens from Spanish Bar, Colorado. ignobilisella Chambers, 1875d: 33, 51, Laverna; SYNTYPE, MCZ 1519; United States: Texas, Bas- que [sic] County, September; Mompha ignobili- sella (Chambers). murtfeldtella Chambers, 1875b: 237, Laverna; HO- LOTYPE, MCZ 1375; United States: [Missouri, 15 September 1874, Murtfeldt]; Mompha murtfeld- tella (Chambers). obscurusella Chambers, 1875d: 53, Laverna; HO- LOTYPE, MCZ 1371; United States: Texas, Basque [sic] County; Mompha murtfeldtella (Chambers). ornotheraesemenella Chambers, 1876b: 138, Lav- erna; SYNTYPE, MCZ 14969; United States: Mis- souri; Mompha brevivittella (Clemens). tricristatella Chambers, 1875e: 211, Leucophryne; SYNTYPE, MCZ 1381; Canada; Mompha tricris- tatella (Chambers). unifasciella Chambers. lS76b: 159, Laverna; SYN- TYPE, MCZ 1378; United States: California; Mom- MCZ Microlepidoptera Types • Miller and Hodges 61 pha unifasciella (Chambers); Missing hindwings and abdomen. Family Agonoxenidae bicristatella Chambers, 1875e: 210, Gelechia; SYN- TYPES (2), MCZ 1459; Canada; Blastodacna bi- cristatella (Chambers). bipunctella Chambers, 1880: 187, Aelia; SYNTYPE, MCZ 1360; United States: Texas. Family Cosmopterigidae albalineella Chambers, 1878c: 95, Eriphia?; HO- LOTYPE female, MCZ 14965; United States: Tex- as, Bosque County; Eralea albalineella (Chambers). eoncolorella Chambers, 1875d: 55, Elachista?; HO- LOTYPE female, MCZ 1352; United States: Texas, Bosque County; Periploca orichalcella (Clemens). eoncolorella Chambers, 1875d: 55, Eriphia; LEC- TOTYPE male, MCZ 1356; United States: Texas; Ithome eoncolorella (Chambers); Leetotype des- ignated by Hodges (1961b: 87), also 3 paraleeto- types in MCZ. determinatella Zeller, 1873: 289, Oeeophora; HO- LOTYPE male, MCZ 1710; United States: Texas; Triclonella determinatella (Zeller). erransella Chambers, 1874a: 52, Perimede; HO- LOTYPE, MCZ 1521; United States: Kentucky, Covington, Chambers; Description based on one specimen, but 4 specimens present in MCZ. gleditschiaeella Chambers, 1876b: 135, Laverna(?) (Anybia?); LECTOTYPE male, MCZ 1373; United States: Kentucky, Chambers; Periploca gledit- schiaeella (Chambers); Leetotype designated by Hodges (1962a: 88), also paralectotype in MCZ. miscecolorella Chambers, 1875d: 51, Laverna; LEC- TOTYPE male, MCZ 1370; United States: Texas, Bosque Count); Walshia miscecolorella (Cham- bers); Leetotype designated by Hodges (1961a: 70). montisella Chambers, 1875c: 297, Cosmopteryx [sic]; HOLOTYPE, MCZ 14967; United States: Colo- rado, Spanish Bar; Cosmopterix montisella Cham- bers. nigrilineella Chambers, 1878c: 96, Eriphia?; HO- LOTYPE male, MCZ 1357; United States: Texas, Bosque County; Melanocinclis nigrilineella (Chambers). ostryaeella Chambers, 1874a: 74, Aeaea; HOLO- TYPE male, MCZ 14966; United States: Kentucky; Stilbosis ostryaeella (Chambers). purpuriella Chambers, 1874a: 73, Chrysopeleia; SYNTYPES (3), MCZ 1358; United States: Ken- tucky. quadricustatella Chambers, 1880b; 186, Aeaea; HO- LOTYPE male, MCZ 1359: I nited States: Texas, Waco; Stilbosis quadricustatella (Chambers). l-lineella Chambers, 1878c: 95, Cosmopteryx [sic]; HOLOTYPE female, MCZ 1518; United' States: Texas, Bosque County; Cosmopterix quadrilineella Chambers. sexnotella Chambers, 1878c: 88, Gelechia; HOLO- TYPE, MCZ 1542; United States: Texas, Bosque County; Stagmatophora sexnotella (Chambers). unimaculella Chambers, 1875d: 94, Ithome; LEC- TOTYPE male, MCZ 1520; United States: Texas; Ithome eoncolorella (Chambers); Leetotype des- ignated by Hodges (1961b: 87), and paralectotype in MCZ. Family Scythrididae albapenella Chambers, 1875d: 11, Butalis; S^ V TYPE male, MCZ 1516; United States: Texas, Bas- que [sic] County, October; "Scythris" albapenella (Chambers) [misplaced]; Specimen in poor condi- tion, missing left wings, and part of thorax. arizoniella Kearfott, 1907: 8, Holcocera; SYNTYPE, MCZ 15022; United States: Arizona, Phoenix, Oc- tober, Kunze; Scythris eburnea (Walsingham). immaculatella Chambers, 1875d: 10, Butalis; SYN- TYPE, MCZ 1515; United States: Texas, Basque [sic] Count\, April; Scythris impositella (Zeller). plausipennella Chambers, 1875d: 10, Butalis; SYN- TYPE, MCZ 1517; United States: Texas, Basque [sic] County; Scythris plausipennella (Chambers); Missing left wings. Family Gelechiidae albimarginella Chambers, 1875c: 291, Gelechia; SYNTYPE male, MCZ 2992; United States: Colo- rado, Grand River; Gnorimoschcma albimargi- nella (Chambers); Abdomen and left wings miss- ing. Specimen (MCZ 2992) probably is incorrectly labelled Kentucky. It matches a specimen from West Fork of Oak Creek, Coconino Count), Ari- zona; and the type locality is Grand River, Colo- rado. albistrigella Chambers, 1872a: 171, Gelechia; SYN- TYPES" (2), MCZ 1522; United States: Kentucky, June, Chambers; Untomia albistrigella (Cham- bers); Untomia albistrigella was described from a single specimen; one of the specimens is a false type. amorphaeel la Chambers, lS77a: 124. Gelechia; SYNTYPES (3 of 4), MCZ 1480; United States: Colorado, Edgerton, Chambers; Filatima ornati- fimbriella (Clemens). 'ulletin Museum of Comparative Zoology, Vol. 152, No. 2 anarsiella Chambers, L877a: 126, Gelechia; SYN- ryp] S 8 M< Z 1 184; United States: Colorado, I dgerton, ( Ihambers. apicistrigella Chambers, L872a: 66, Parasia; HO- LOTYPE Female, MCZ 1523; United States: Ken- tucky, Chambers; Battaristis nigratomella (Cle- mens Specimen lacks abdomen and right hindwing. atrupictella Dietz. 1900 350, Kucordylea; HOLO- hl'l male, \1C:Z 2986; United States: Pennsyl- vania, Hazleton; Coleotechnites atrupictella (Dietz). basifasciella Zeller, L873: 269, Gelechia (Poecilia?); HOLOTYPE male, MCZ 2988; United States: Tex- as. Boll; Pseudotelphusa basifasciella (Zeller). basistrigella Zeller. L873: 270, Gelechia; HOLO- TYPE female. MCZ 2981; United States: Texas, Boll; Xenolechia basistrigella (Zeller). bifasciella Chambers, 1874a: 76, Gelechia; SYN- TYPE male, MCZ 1362; United States: Kentucky, Chambers; Theisoa constrictella (Zeller). bifidella Dietz, 1900: 351, Nealyda; SYNTYPE male, MCZ 2990: United States: Colorado, Glenvvood Springs. bimaculella Chambers, 1877d: 122, INothris?; HO- I OTYPE male. MC:Z 1557; United States: Colo- rado. Edgerton; Dichomeris georgiella (Walker). bimaculella Chambers, 1872a: 108, Depressaria; st \ n I'l'. female, MCZ 1524; United States: Ken- tucky Chambers; Fascista bimaculella (Cham- bers). biminimaculella Chambers, 1880b: 183, Gelechia; SYNTYPES 2 MCZ 1525; United States: Texas, \\ aco, Belfrage; Filatima biminimaculella (Cham- bers). biscolorella Chambers, lS72a: 195. Agnippe; SYN- \\ I'l female, MCZ 1281; I nited States: Kentucky, ( Ihambers. bosqueella Chambers. L875d: 92, Oecophora; SYN- TYPE female, MCZ L443; United States: Texas; Stegasta bosqueella (Chambers); bight hindwing missing. canopulvella Chambers L878c: 91, Gelechia; HO- I I > n I'l : female, M( Z I 152; I nited States: Texas, Bosque County; Filatima obscurosuffusella ( hambers). cercerisella Chambers L872a: 108, Depressaria; SYNTYPES (2 MCZ 1427; United States: Ken- tu< k\ ( hambers. Fascista cercerisella (Chambers); Specimens are double mounted on one block of pith. Male lacks head, temale lacks left forewing 1 abdomi cilialineella I S7 lb 242, Gelechia; S'l \- TYP1 nited States: Texas; Isophrictis collinusella Chambers, 1877d: 128, Gelechia; HO- LOTYPE male, MCZ 1526; United States: Colo- rado, Edgerton; Gnorimoschema collinusella (Chambers); Right wings only. concinnusella Chambers, 1875b: 253, Gelechia; SYNTYPES (2), MCZ 1460; United States: Texas; Battaristis concinnusella (Chambers). consonella Zeller, 1873: 251, Gelechia; SYNTYPE male, MCZ 2982; United States: Texas, Boll; An- acampsis rhoifructella (Clemens). costarufoella Chambers, 1874b: 240, Gelechia; LEC- TOTYPE, MCZ 1527; United States: Texas, Cham- bers; Dichomeris costarufoella (Chambers); Lec- totype designated by Hodges (1986: 114). crescentifasciella Chambers, 1874b: 237, Gelechia; SYNTYPES (7), MCZ 1477; United States: Texas; Compsolechia crescentifasciella (Chambers). cristatella Chambers, 1875b: 241, Gelechia; SYN- TYPES (2), MCZ 1472; United States: Kentucky, Chambers; Coleotechnites cristatella (Chambers). cristifasciella Chambers, 1878c: 87, Gelechia; SYN- TYPE female, MCZ 1474; United States: Kentucky, 1 1 May [no year stated], Chambers; Arogalea cris- tifasciella (Chambers); Worn specimen. Described from two syntypes. curvistrigella Chambers, 1872a: 133, Telphusa; SYN- TYPE female, MCZ 2939; United States: Kentucky, Chambers; Telphusa longifasciella (Clemens); Ab- domen missing. depressostrigella Chambers, 1874b: 236, Gelechia; SYNTYPES (2), MCZ 1434; United States: Texas; Filatima ochreosuffusella (Chambers). disconotella Chambers, 1878c: 86, Gelechia; SYN- TYPE, MCZ 1475; United States: Kentucky, Cham- bers; Monochroa disconotella (Chambers); Left wings missing. discoocellella Chambers, 1872a: 194, Gelechia; SYN- TYPES (4), MCZ 1439; United States: Kentucky, September, Chambers; Chionodes discoocellella (Chambers). dorsivittella Zeller, 1873: 265, fig. 20, Gelechia (Tel- eia?); HOLOTYPE male, MCZ 1707; United States: Texas, Dallas, Boll; Coleotechnites vagatioella (Chambers); Haustellum and abdomen missing. • I ii l>i irl la Chambers, 1872a: 92, Depressaria?; LEC- TOTYPE male, MCZ 1529; United States: Ken- tucky, Chambers; Dichomeris juncidella (Cle- mens); Lectotype designated by Hodges (1986: 111). elegantella Chambers, 1874b: 239, Gelechia; SYN- TYPES (8), MCZ 1494; United States: Texas, Chambers; Aristotelia elegantella (Chambers). eupatoriella Chambers, 1872a: 221, Ypsolophus; LECTOTYPE male, MCZ 1531; United States: [Kentucky], Chambers; Dichomeris setosella (Cle- MCZ Microlepidoptera Types • Miller and Hodges 63 mens); The original description indicates that one specimen was reared; it should be the holotype. None of the specimens bearing MCZ 1531 could be recognized as the one that Chambers described. Hodges (1986: 82) designated one of the four as lectotype. fragmentella Zeller, 1873: 271, Gelechia (Poecilia?); HOLOTYPE female, MCZ 1706; United States: Texas, Dallas, Boll; Pseudotelphusa quercinigra- cella (Chambers); Specimen lacks right wings and abdomen. fuscocristatella Chambers, 1875d: 9, Naera; SYN- TYPES (4), MCZ 1382; United States: Texas. fuscoluteella Chambers, 1872a: 106, Depressaria; HOLOTYPE male, MCZ 1462; United States: Ken- tucky, Chambers; Gelechia albisparsella (Cham- bers); Abdomen missing. fuscomaculella Chambers, 1872a: 170, Gelechia; HOLOTYPE male, MCZ 1469; United States: Ken- tucky, Chambers; Chionodes fuscomaculella (Chambers). fuscoochrella Chambers, 1872a: 106, Gelechia; SYN- TYPE female, MCZ 1532; United States: Kentucky, Chambers; Chionodes mediofuscella (Clemens). fuscopalidella Chambers, 1875b: 231, Sinoe; SYN- TYPES (3), MCZ 1533; United States: Kentucky, Chambers; Sinoe robiniella (Fitch); One specimen has the left forewing; the other specimen has the right forewing. fuscopulvella Chambers, 1872a: 195, Agnippe; HO- LOTYPE male, MCZ 1282; United States: Ken- tucky, April, Chambers. fuscopulvella Chambers, 1872a: 170, Gelechia; HO- LOTYPE female, MCZ 1492; United States: Ken- tucky, June, Chambers; Chionodes obscurusella (Chambers). fuscoslrigella Chambers, 1876b: 30, Polyhymno; HOLOTYPE male, MCZ 1547; United States: Tex- as, Belfrage; Polyhymno luteostrigella Chambers; Abdomen missing; labial palpus nearly devoid of scales ventrally. fuscotaeniaella Chambers, 1878c: 89, Gelechia; HO- LOTYPE male, MCZ 1448; United States: Texas; Rifseria fuscotaeniaella (Chambers); Head, thorax, and left wings only. gilviscopella Zeller, 1873: 266, Gelechia; SYNTYPES (2), MCZ 1708; United States: Texas, Dallas, Boll; Coleotechnites quercivorella (Chambers). glandiferella Zeller, 1873: 275, Gelechia; PARA- LECTOTYPE female, MCZ 1709; United States: Texas; Deltophora glandiferella (Zeller); Lecto- type in BMNH designated by Sattler (1979: 297). glycyrhizaeella Chambers, 1877a: 124, Gelechia; HOLOTYPE female, MCZ 1483; United States: Colorado, Edgerton; Filatima glycyrhizaeella (Chambers); Abdomen missing. innocuella Zeller, 1873: 249, Gelechia (Tachvptilia); SYNTYPE male, MCZ 1721; United States: Texas, Boll; Anacampsis innocuella (Zeller). intermediella Chambers, 1879a: 89, Gelechia; HO- LOTYPE female, MCZ 1447; United States: Texas, Bosque County; Aristotelia intermediella (Cham- bers); Labial palpi, metathorax, and abdomen miss- ing. inversella Zeller, 1873: 248, Epicorihylis; HOLO- TYPE male, MCZ 1715; United States: Texas, Boll; Dichomeris inversella (Zeller). latifasciella Chambers, 1875b: 251, Gelechia; PSEI DOTYPE female, MCZ 2940; United States: Mis- souri, St. Louis; Telphusa latifasciella (Chambers); The original description indicates that the moth is light. The specimen (MCZ 2940) is dark and rep- resents the dark form of latifasciella. lavernella Chambers, 1874b: 242, Gelechia; HO- LOTYPE female, MCZ 1473; United States: Texas; Symmetrischema lavernella (Chambers). leuconota Zeller, 1873: 268, Gelechia; HOLOTYPE male, MCZ 2973; United States: Texas, Boll; Evippe leuconota (Zeller). liturosella Zeller, 1873: 265, Gelechia (Lite); HO- LOTYPE female, MCZ 1704; United States: Texas, Dallas, Boll; Chionodes mediofuscella (Clemens). luteostrigella Chambers, 1874b: 247, Polyhymno; PSEUDOTYPE female, MCZ 1546; United States: Texas; The specimen (MCZ 1546) is a Coleophora. The original description applies to the current con- cept of the gelechiid species Polyhymno luteostri- gella . maculimarginella Chambers, 1874b: 241, Gelechia; SYNTYPES (2), MCZ 1446; United States: Texas; Chionodes fuscomaculella (Chambers). marmorella Chambers, 1875b: 239, Gelechia; HO- LOTYPE, MCZ 1471; United States: Kentucky, Chambers; Gnorimoschema emancipatum (Meyr- ick), REVISED STATUS; Gelechia marmorella Chambers, 1875 is a junior primary homonym of Gelechia marmorella Doubleday, 1859. Phthori- maea emancipata Meyrick, 1925 is an objective replacement name for Gelechia marmorella Chambers, 1875. minimaculella Chambers, 1874b: 235, Gelechia; HOLOTYPE male, MCZ 1485; United States: Tex- as; Aroga thoracealhclla (Chambers). minimella Chambers, 1S741> 24 5, Gelechia; SYN- TYPES (2), MCZ 1487; United States: Texas; Ar- istotelia pullusella (Chambers). monumentella Chambers, I877d: 125, Gelechia; HO- LOTYPE male, MCZ 1482; United States: Colo- 64 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 rado, Monument Park; Scrobipalpa monumentella ( hambers). multifasciella Chambers, L875d: 93, Theisoa; SYN- nil MCZ L363; I nited States: Texas. nigrella Chambers, L875b: 250. Gelechia; PSEU- DOTYPE MCZ L489; I nited States: Texas; Syn- copacma nigrella (Chambers); The single, abdo- menless specimen does not match the original description. aoostrigella Chambers, 1878c: 92, Dasycera; HO- LOTYPE male. MCZ 1544; United States: Ken- tucky ( lhambers; Dichomeris nonstrigella (Cham- hcrs). obliquifasciella Chambers, 1880b: 182, Gelechia; HOLOTYPE male, MCZ 1170; United States: Tex- as. Telphusa longifasciella (Clemens). obliquistrigella Chambers, 1872a; 65, Anarsia; SYN- TYP1 S 2 MCZ 1535; United States: Kentucky, Chambers; Coleotechnites obliquistrigella (Cham- bers). obscurosuffusella Chambers, L878c: 90, Gelechia; s^ NT1 PE Female MCZ 1453; United States: Texas, Bosque County; Filatima obscurosuffusella ( hambersi; Abdomen and left forewing missing. obscurusella Chambers, 1872a; 106, Depressaria; SYNTYPE male MCZ 1632; United States: Ken- tucky Chambers; Chionodes obscurusella (Cham- bers). occidentella Chambers, 1875b; 24b, Gelechia; SYN- TYPE Female, MCZ 1464; tinted States: Califor- nia Behrens; Chionodes occidentella (Chambers). ocellella Chambers, L877a; 126, Gelechia?; HO- LOTYPE male. MCZ 1440; I nited States: Colo- rado. Edgerton; Gelechia hianulella (Chambers); The holotype is a male, not a female as indicated b\ Chambers. ochreocostella Chambers, ISTSc: 91, Gelechia; HO- LD H PE female, MCZ I 155; United States: Texas, Bosque County; Anacampsis rhoifructella (Cle- mens ochreostrigella Chambers, L875b: 247, Gelechia; HOLOTYPE male MCZ L463; United States: Cal- ifornia, Behrens; Chionodes ochreostrigella ( hambers); Abdomen missing. ochreostrigella Chambers, L877a; 120, Gelechia; SI Vn PES 1 M< ./. 2995; I nited States: Colo- rado Edgerton; Scrobipalpula henshawiella (Busck); The syntypes represent two species olympiadella Z. II. i L873 259, Gelechia; PSEU- DOTYPES 2 males M< :/. 2021. I nited States: Tex- rcerisella (Chambers); These speci- S weir nol seen b\ Zeller when he described th< pallidastrigella Chambers, 1874b: 244, Cleodora; SYNTYPE, MCZ 1561; United States: Texas; Iso- phrictis pallidastrigella (Chambers). pallidella Chambers, 1874b: 245, Gleodora; SYN- TYPES (2), MCZ 1562; United States: Texas; Iso- phrictis pallidella (Chambers); The specimens are very poor and probably represent two species. pallidochrella Chambers, 1872a: 126, Depressaria;; HOLOTYPE female, MCZ 1536; United States: Kentucky, May, Chambers; Symmetrischema pal- lidochrella (Chambers). pallidochrella Chambers, 1873a: 188, Helice; SYN- TYPES (4), MCZ 1620; (13), MCZ 1534; United States: Kentucky, Chambers; Theisoa pallidochrel- la (Chambers); Two MCZ type numbers with as- sociated specimens represent the same species. palpiannulella Chambers, 1872a: 68, Gelechia; SYN- TYPES (4), MCZ 1493; United States: Kentucky, Chambers; Monochroa absconditella (Walker). palpilineella Chambers, 1875b: 252, Gelechia; SYN- TYPES (3), MCZ 1486; United States: Kentucky, Chambers; Syncopacma palpilineella (Chambers). pedmontella Chambers, 1877a: 123, Gelechia; HO- LOTYPE male, MCZ 1478; United States: Colo- rado, Edgerton; Gnorimoschema pedmontella (Chambers). pennsylvanica Dietz, 1900b: 353, Pseudochelaria; HOLOTYPE female, MCZ 2974; United States: Pennsylvania, Hazleton. physaliella Chambers, 1872a: 173, Gelechia; SYN- TYPE female, MCZ 33255; United States: Ken- tucky, Chambers; Aristotelia physaliella (Cham- bers); Chambers described physaliella from two specimens reared from Physalis viscosa L. The specimen in the MCZ was received from the Pea- body Academy collection. A second syntype is in USNM. platanella Chambers, 1872a: 146, Cirrha; PSEU- DOTYPE male, MCZ 1430; United States: Ken- tucky, Chambers; Gelechia albisparsella (Cham- bers); Cirrha platanella Chambers is a replacement name for Depressaria albisparsella Chambers and as such does not have a separate type from the name it replaces. However, no type material of albisparsella exists. This specimen could be consid- ered for designation as neotype of albisparsella. It has only the head, thorax, and right forewing. plutella Chambers, 1874b: 238, Gelechia; PSEU- DOTYPES (2), MCZ 1429; United States: Ken- tucky; Dichomeris serrativittella (Zeller); These specimens are from Kentucky, not Texas as stated in the original description, and the color pattern on the forewings is reversed from that of the orig- in, il description. MCZ Microlepidoptera Types • Miller and Hodges 65 plutella Chambers, 1874b: 244, Neda; SYNTYPES (2), MCZ 1369; United States: Texas; Megacras- pedus plutella (Chambers); Another svntvpe in USNM. plutella Chambers, 1875d: 106, Phaetusa; PSEU- DOTYPES (2), MCZ 1429; United States: Ken- tucky; Evippe leuconota (Zeller); These specimens are from Kentucky, not Texas as stated in the orig- inal description. One female syntype in USNM. prunifoliella Chambers, 1873a: 186, Evippe; SYN- TYPE male, MCZ 1537; United States: Kentucky, Chambers. pseudacaciella Chambers, 1872a: 107, Depressaria; SYNTYPES (11), MCZ 1490; United States: Ken- tuck), Chambers; Filatima pseudacaciella (Cham- bers). pudibundella Zeller, 1873: 273, Aristotelia; LEC- TOTYPE male, MCZ 1441; United States: Texas, Dallas, Boll; Lectotvpe designated bv Forbes (1932: 429). c| u;i< 1 1 i mac ii I < I la Chambers, 1874b: 237, Gelechia; LECTOTYPE male, MCZ 1436; United States: Texas; Anacampsis rhoifructella (Clemens); The lectotype, present designation, bears the following labels: l)"Type 1436"; 2)"Gelechia quadrimaculel- la Cham. Texas"; 3)" Anacampsis rhoifructella Clem. =quadritnaculclla Cham. AB 1902"; 4)"Lectotype R. W. Hodges. It is selected to ensure that the name continues to be associated with An- acampsis rhoifructella (Clemens). The second syn- type is a species of Neodactylota. querciella Chambers, 1872a: 127, Depressaria; SYN- TYPES (3), MCZ 1538; United States: Kentucky, Chambers; Neotelphusa querciella (Chambers). querciella Chambers, 1872a: 223, Ypsolophus; PSEUDOTYPE female, MCZ 1560; United States: Kentucky, Chambers; Dichomeris ventrella (Fitch); The identification label reads "Ypsolophus quer- cicoella var. pomatella 482/1", and the specimen is Dichomeris ligulella Huebner. The original de- scription of querciella clearly states that the wings are broad and thus does not apply to this specimen. quercinigracella Chambers, 1872a: 170, Gelechia; HOLOTYPE male, MCZ 1701; United States: Ken- tucky, Chambers; Pseudotelphusa quercinigracella (Chambers); Abdomen and right wings missing. quercipominella Chambers, 1872a: 222, Ypsolo- phus; HOLOTYPE female, MCZ 1560; United States: Kentucky, Chambers; Dichomeris ligulella Huebner. quercivorella Chambers, 1872a: 173, Gelechia; HO- LOTYPE female, MCZ 1539; United States: Ken- tucky, Chambers; Coleotechnites quercivorella (Chambers). quinqueannulella Chambers. 1872a: 191, Gelechia; HOLOTYPE female, MCZ 1467; United States: Kentucky, Chambers; Trypanisma prudens Cle- mens. reedella Chambers. 1872a: 222, Ypsolophus; S1! V TYPES (3), MCZ 1559; United States: Kentucky, Chambers; Dichomeris ligulella Huebner; The three specimens are the same species ribesella Chambers, 1875c: 290, Gelechia; HOLO- TYPE female, MCZ 2984; United States ( Jolorado, Spanish Bar, Clear Creek. rileyella Chambers, 1872a: 106, Depressaria; S1! V TYPE male, MCZ 1431; United States: Kentucky, Chambers; Gelechia rileyella (Chambers). rubensella Chambers, 1872a: 193, Gelechia; PSEU- DOTYPES (3), MCZ 1449; United States: Ken- tucky, Chambers; Aristotelia rubidella (Clemens); The status of these specimens as types is question- able. Chambers described rubensella from a single specimen that had its fringes singed by a gas light. None of these is singed. rufusella Chambers, 1874b: 240, Gelechia; SIN- TYPES (3), MCZ 1444; United States: Texas; An- acampsis fullonella (Zeller). saphirinella Chambers, 1875b: 250, Gelechia; HO- LOTYPE, MCZ 1468; United States: [Kentucky, Chambers]; Gnorimoschema saphirinella (Cham- bers); Labial palpi, right hindwing, and abdomen missing. The right forewing is glued on a block. Although the original description implies Texas as the type locality, the label states "Kentucky./ Chambers." saundersella Chambers, 1876b: 173, Gelechia; SYN- TYPES (2), MCZ 1540; United States: Kentucky. Chambers; Taijgete saundersella (Chambers). scutellariaeela Chambers, 1873a: 175, Gelechia; SYNTYPE male, MCZ 1541; United States: Ken- tucky, Boone County, Verona; Scrobipalpa .scutel- lariaeela (Chambers). sella Chambers, 1874b: 238, Gelechia; LECTOTYPE male, MCZ 1481; United States: Texas; Deltophora sella (Chambers); Lectotvpe designated by Sattler (1979: 294). serratipalpella Chambers, 1877a: 123. Gelechia; HOLOTYPE male, MCZ 1479; I nited States: Col- orado, Edgerton; Gnorimoschema serratipalpella (Chambers); The abdomen is glued to the rest of the specimen. serrativittella Zeller, 1873: 280, Gelechia; PARA- LECTOTYPE female, MCZ 1705; United States: Texas, Dallas, Boll; Dichomeris serrativittella (Zell- er); Lectotype female in BMNH designated by Hodges (1986: 101). 66 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 ..■x-irigella Chambers, 1874b: 248, Polyhymno?; SYNTYPES J), M< Z L545; I nited States: Texas; Calliprora sexstrigella (Chambers), solaniiella Chambers, L873a: 176, Gelechia; PSEU- D( >T"i PE female, \l( !Z 2453; hophrictis similiella ( lumbers); Gelechia solaniiella Chambers is a re- placement name for Gelechia similiella Chambers, 1872 straminiella Chambers, 1872a: 224, Ypsolophus; LECTOTYPE female, MCZ 1558; United States: Kentucky, Chambers; Dichomeris punctidiscella ( lemens); Lectotype designated by Hodges (1986: 54 ternariella Zeller, 1873: 264, Gelechia (Lita); HO- LOTYPE female, MCZ 1702; United States: Texas, Boll; Fascista bimaculella (Chambers). texanella Chambers, 1880b: 179, Anesychia; LEC- TOTYPE male, MCZ 1424; United States: Texas; Lita texanella (Chambers); Lectotype designated b) Hodges (1966: 30). thoracealbella Chambers, 1874b: 235, Gelechia; S"i NTYPE male. MCZ 1435; United States: Texas, Waco, Belfrage; Aroga thoracealbella (Chambers). trialbamaculella Chambers, 1875b: 250, Gelechia; HOLOTYPE female, MCZ 1456; United States, Texas. Waco. Belfrage; Aroga trialbamaculella ( hambers ; Head and right wings missing. Uifasciella Chambers, 1875b: 252, Gelechia; SYN- TYP1 S 2), MCZ 1445; United States: Texas, Waco, Belfrage; Filatima albilorella (Zeller). trilineella Chambers, 1877a: 125, Gelechia; SYN- rYPES (6), MCZ 1454; United States: Colorado, Edgerton; Aroga trilineella (Chambers); One syn- type is not conspecific with the other five. trimaculella Chambers, 1874b: 243, Anarsia; LEC- TOTYPE mal. MCZ 1556; United States: Texas, Waco, Belfrage; Isophrictis trimaculella (Cham- bers); The lectotype, present designation, bears the following labels: 1 l"Type L556"; 2)"Texas"; 3)"72"; l '996": 5 "trimaculella"; 6)"Lectotype R W Hodges"; 7)"RWH genitalia slide 3314." It is se- lected to ensure continued use of the name in Is- ophrictis. Three species are represented among the s\ nt) pes trimaculella Chambers, lS7l!>: 238, Gelechia; PAR \l I ( ion PES - MCZ 1476; I nited States: Texas, Waco, Belfrage; Helcystogramma melan- ocarpum Meyrick); Lectotype in USNM desig- nated b) Hodges 1986: L30 trimaculella Packard L867: 61, Gelechia; SYN- TYP1 S 2 \K / L563; Canada: Labrador, Straw- berry Harbor: Chionodes continuella (Zelli triocellella ( hambers L877a 127 Gelechia; SYN- TYPES 6 MCZ L453; I nited states: Colorado, Edgerton; Gnorimoschema triocellella (Cham- bers). unctulella Zeller. 1873: 257, Gelechia; HOLOTYPE male, MCZ 1703; United States: Texas, Boll; Fila- tima ornatifimbriella (Clemens). variella Chambers, 1872a: 174, Gelechia; SYN- TYPES (2 of several), MCZ 1544; United States: Kentucky, Chambers; Coleotechnites variella (Chambers); One female syntype is in USNM. versutella Zeller, 1873: 253, Gelechia; HOLOTYPE female, MCZ 1457; United States: Texas, Boll. violaceofusca Zeller, 1873: 258, Gelechia; HOLO- TYPE male, MCZ 2981; United States: Texas, Dal- las, Boll; Chionodes discoocellella (Chambers). walsinghami Dietz, 1900b: 352, Pseudochelaria; SYNTYPE male, MCZ 3266; United States: Penn- sylvania, Hazleton. SUPERFAMILY COPROMORPHOIDEA Family Glyphipterigidae exoptatella Chambers, 1875b: 234, Glyphipteryx [sic]; HOLOTYPE, MCZ 1564; United States: Kentucky, Linden Grove Cemetery, [June], Chambers; Di- ploschizia impigritella (Clemens); Heppner (1981: 322) gives type locality as '"[Covington?, Kenton Co.]." montisella Chambers, 1875c: 292, Glyphipteryx [sic]; LECTOTYPE male, MCZ 32887; United States: Colorado, Denver, South Park, 10,000 feet; Gly- phipterix montisella Chambers; Lectotype desig- nated by Heppner (1985: 124). SUPERFAMILY YPONOMEUTOIDEA Family Plutellidae NOTE: The Walsingham syntypes in this family were sent to Chambers by Wal- singham. We suggest that lectotypes should be selected from syntypes at the BMNH because Walsingham's personal collection is there. He most likely sent syntypes (a term not used in the 1880s) to Chambers as examples of his species. Types (holo- types and lectotypes in current terms) he would have retained. canariella Walsingham, 1881: 309, pi. XXXV, fig. 11, Cerostoma; SYNTYPES (2), MCZ 14982; United States California, Lake County, Scott's Valley, 18 May 1871; Ypsolopha canariella (Walsingham). castella Walsingham, 1881: 310, pi. XXXV, fig. 13, Euceratia; SYNTYPES (2), MCZ 14984; United States. California, San Francisco. MCZ Microlepidoptera Types • Miller and Hodges 67 deniiferella Walsingham, 1881: 308, pi. XXXV, fig. 10, Cerostoma; SYNTYPES (2), MCZ 14985; United States: California, Mount Shasta, August 1871; Yp- solopha deniiferella (Walsingham). falciferella Walsingham, 1881: 307, pi. XXXV, fig. 7, Cerostoma; SYNTYPE (1 of 18), MCZ 14988; United States: California and Oregon; Ypsolopha falciferella (Walsingham). frustella Walsingham, 1881: 309, pi. XXXV, fig. 12, Cerostoma; SYNTYPES (2 of 26), MCZ 14981; United States: California, Shasta County, 28 July 1871; Ypsolopha frustella (Walsingham). ochrella Chambers, 1880b: 181, Pluteloptera; SYN- TYPE, MCZ 1414; United States: Texas, Belfrage; Ypsolopha ustella (Clemens); Missing left wings. polita Walsingham, 1881: 302, pi. XXXV, fig. 2, Cal- antica; SYNTYPES (2), MCZ 14992; United States: California; Eucalantica polita (Walsingham). securella Walsingham, 1881: 311, pi. XXXV, fig. 14, Euceratia; SYNTYPE (1 of 17), MCZ 14983; United States: California, Sonoma County, May 1871. subfasciella Walsingham, 1881: 303, pi. XXXV, fig. 3, Araeolepia; PARALECTOTYPE (1 of 17), MCZ 14986; United States: Oregon, Currant Creek (an affluent of John Day River), 16 April 1872; Lee- totvpe in BMNH designated by Heppner (1982: 278). vanella Walsingham, 1881: 305, pi. XXXV, fig. 6, Plutella; SYNTYPE, MCZ 14987; United States: California, San Francisco. Family Yponomeutidae celastrusella Kearfott, 1903: 150, Zelleria; SYN- TYPES (2), MCZ 14228; United States: New Jersey, Essex County, "Pk." and Caldwell. crassivenella Zeller, 1872: 563, fig. 27, Enaemia; HO- LOTYPE female, MCZ 15001; United States: Tex- as, Dallas, Boll; Lactura papula (Huebner). Family Argyresthiidae altissimella Chambers, 1877d: 130, 147, Argyresthia; SYNTYPE, MCZ 1412; United States: Colorado, Mount Elbert, 11,000 feet, July. austerella Zeller, 1873: 305, fig. 38, Argyresthia; SYNTYPES (3 of 6), MCZ 14254; United States: Texas, Dallas, Boll. belangerella Chambers, 1875e: 145, Argyresthia; SYNTYPE, MCZ 1407; Canada; Missing right wings. deletella Zeller, 1873: 305, Argyresthia; SYNTYPES (3), MCZ 14254; United States: Texas, Dallas, Boll. TYPE, MCZ 1411; United States: Colorado, Ed- gerton, July. pedmontella Chambers, 1877d: 131, Argyresthia; SYNTYPES (2), MCZ 1409; United States: Colo- rado, Edgerton, July. quadristrigella Zeller, 1873: 304, Argyresthia; IIO- LOTYPE male, MCZ 14252; United States: Texas, Dallas, Boll; Missing abdomen. quercicolella Chambers, 1877d: 130, Argyresthia; SYNTYPES (2), MCZ 1410; United States: Colo- rado, Edgerton, June. thuiella Packard, 1871: 24, pi. 1, fig. 6, Bucculatrix; SYNTYPES, MCZ 14963; United States: Maine, Brunswick, July, on cedar tree; Argyresthia thuiel- la (Packard); Fragments of several specimens in vial. ii mlu Intel la Chambers, 1874a: 10, Argyresthia; SYN- TYPES (6), MCZ 1408; United States: Kentucky, Chambers. Family Heliodinidae abroniaeella Chambers, 1876b: 217, Lithariapteryx; SYNTYPES (9), MCZ 1565; United States: Colo- rado, Edgerton, over 6000 feet, July. bella Chambers, 1875d: 73, Aetole; SYNTYPE, MCZ 1364; United States: Texas, August, Belfrage; He- liodities bella (Chambers). SUPERFAMILY SESIOIDEA Family Sesiidae aureopurpura H. Edwards, 1880: 72, Aegeria;; HO- LOTYPE female, MCZ 928; United States: Texas, Dallas, Boll; Carmenta bassiformis (Walker). caudata Harris, 1839: 311, Aegeria;; SYNTYPES (1 male, 1 female), MCZ 26354; United States: [Mas- sachusetts, Neponset River, 30 August 1823, Harris no. 87]; Alcathoe caudata (Harris). cucurbitae Harris, 1828: 33, Aegeria;; SYNTYPES (5 adults and 5 pupal cases), MCZ 33258; United States: [Harris no. 249]; Mclittia cucurbitae (Harris). denudatum Harris, 1839: 310. Troehilium; SYN- TYPE, MCZ 26359; United States: [Harris no. 31 1 ]; Sesia asilipennis (Boisduval). fulvipes Harris, 1839: 312, Aegeria; SYNTYPE, M( Z 26361; United States: [Harris no. 17]; Synanthedon fulvipes (Harris). maculipes Grote and Robinson, 1868: 184, Zenodox- us; PSEUDOTYPE female, MCZ 929; United States: Texas, Dallas, Boll; Not a syntype according to Duckworth and Eichlin (1978: 16). montella Chambers, 1877d: 130, Argyresthia; SYN- marginatum Harris, 1839: 309, Troehilium; SYN- 68 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 TYPE Female, MCZ 26356; United States: New Hampshire, [Dublin], L. W. Leonard [Harris no. 38]; Pennisetia marginata I Harris). aebraskae II Edwards, 188 1 I SI, Kuhagena; HO- I ( ) ] "i I'l male, M< / 930; I nited States; Nebraska, Mr. Austin; Broken and in poor condition. polisiiformi* H.n is L854; 216, Aegeria; SYNTYPES males. 2 it-males. 3 pupal cases), MCZ 26358; I nited Malts North Carolina. Albermarle, F. J. Kron [Harris no. 791]; Vitacea polistiformis (Har- ris). pyri Harris. L830; 2. \egeria; SI \ TYPE, MCZ 26363; I nited States; [Massachusetts. Dorchester and Cambridge, Harris no. 702]; Synanthedon pyri (Harris); Missing abdomen *citula Harris. 1839: 313, Aegeria; SYNTYPES (2 adults. 1 pupal shell), MCZ 26362; United States: [Harris no. 333]: Synanthedon scitula (.Harris). syringae Harris, 1839: 311, Aegeria; LECTOTYPE male. MCZ 26360; I nited States: [Harris no. 464]; Podosesia syringae (Harris); One male and one female paralectotype, MCZ 26360. Lectotype des- ignated by Purrington and Nielsen (1987: 551). tibiale Harris. 1839: 309, Trochilium; SYNTYPE fe- male. MCZ 26355: United States: New Hampshire, [Dublin], L. W. Leonard [Harris no. 387]; Sesia tibialis (Harris). tricincta Harris. 1839: 310, Aegeria; SYNTYPES (1 male, 1 female!. MCZ 26357; United States: [Harris mi 322]; Paranthrene tabaniformis (Rottemburg). SUPERFAMILY COSSOIDEA Family Cossidae "crepera Harris. 1833: 591, Cossus"; "SYNTYPE" male MCZ 26393; I nited States: New Hampshire, [Dublin, L. W. Leonard. Harris no. 582]; Prion- oxystus robiniae (Peck); The original description of crepera is generally cited as Harris (1935a: 592; 1835b: 72), but it is only listed there (as Cossus crepera), not described or illustrated. The first de- scription seems to be that ol Packard (1864c: 388) as Xyleutes crepera SUPERFAMILY TORTRICOIDEA Family Tortricidae (including Cochylidae) allutana Zellei 1875 295, fig. 27, Crapholitha; S'i N n PES 2 MCZ I 1325 i nited States New York, ind Texas, Dallas, Boll; Episimus argutanus < ilemens). amphorana Walsingham, 1879: 63, pi. LXXIV, fig. 9, Semasia; SYNTYPES (2), MCZ 15006; United States: Oregon, John Day River, Camp Watson, April, Walsingham; Phaneta amphorana (Wal- singham). angleseana Kearfott, 1907a: 64, Enarmonia; PARA- LECTOTYPE, MCZ 15014; United States: New Jersey, Anglesea, V-00 (?), W. D. Kearfott; Gra- pholita angleseana (Kearfott); Lectotype in AMNH designated by Klots (1942: 398); see also Heinrich (1926: 31). bobana Kearfott, 1907a: 26, Eucosma; PARALEC- TOTYPE, MCZ 14322; United States: Texas, Harris County, 7-5-1899; Described from three syntypes. Heinrich (1923: 103) designated the Salida, Col- orado, specimen as lectotype; it is in USNM, not AMNH as he indicated. The third specimen, from Southwest Colorado, is in AMNH (Klots, 1942: 399). See also Powell (1968:10). bolanderana Walsingham, 1879: 42, pi. LXIX, fig. 10, Paedisca; SYNTYPE, MCZ 15003; United States: California, Mount Shasta, August 1871, Walsingh- am; Eucosma bolanderana (Walsingham); Missing abdomen. cockerellana Kearfott, 1907a: 71. Tortrix; PARA- LECTOTYPE, MCZ 14320; United States: Colo- rado, Glenwood Springs, September 1899; Argy- rotaenia cockerellana (Kearfott); Lectotype male in AMNH designated by Powell (1964: 225). constrictana Zeller, 1875: 305, fig. 36, Paedisca; HO- LOTYPE male, MCZ 14335; United States: Texas, Dallas, Boll; Sonia constrictana (Zeller). desertana Zeller, 1875: 306, fig. 37, Paedisca; LEC- TOTYPE male, MCZ 14338; United States: Texas, Dallas, Boll; Epiblema desertana (Zeller); Lecto- type designated by Miller (1976: 50). dietziana Kearfott, 1907a: 92, Epinotia; PARALEC- TOTYPES (2), MCZ 14302; United States: Penn- sylvania, Hazleton, W. G. Dietz; Rhopobota diet- ziana (Kearfott); Lectotype in AMNH designated by Klots (1942: 401); see'also Heinrich (1923: 191). dudana Kearfott, 1907a: 27, Eucosma; PARALEC- TOTYPE, MCZ 15019; United States: SW Colo- rado, 7-13-1889, W. G. Dietz; Missing abdomen. Lectotype in AMNH designated by Klots (1942: 401); see also Heinrich (1923: 106).' dodecana Zeller, 1875: 311, fig. 40, Paedisca; SYN- TYPES (5), MCZ 14341; United States: Texas, Dal- las, Boll; Pelochrista scintillana (Clemens). eclipsana Zeller, 1875: 298, fig. 29, Crapholitha [sic]; HOLOTYPE male, MCZ 14328; United States: Texas, Dallas, Boll; Missing abdomen. ednana Kearfott, 1907c: 161, pi. VIII: fig. 13, Phal- onia; PARALECTOTYPES (2), MCZ 15017; United MCZ Microlepidoptera Types • Miller and Hodges 69 States: Pennsylvania, Hazleton; Anopina ednana (Kearfott); Leetotvpe in AMNH designated bv Klots (1942: 417). exasperatana Zeller, 1875: 238, Torlrix; SYNTYPES (2), MCZ 14309; United States: Texas, Dallas, Boll; Platynota exasperatana (Zeller). fagigemmaeana Chambers, 1878b: 74, Exartema; SYNTYPE, MCZ 15007; United States: Kentucky, Chambers; Olethreutes fagigemmaeana (Cham- bers); Wings only, plus pupal case. Chambers refers to a single specimen in the MCZ, but then describes both sexes. fragariana Packard, 1869: 335, Lozotaenia; SYN- TYPE, MCZ 14315; United States: Maine; Clepsis persicana (Fitch); Missing abdomen. frigidana Packard, 1856: 57, Penthina; LECTO- TYPE male, MCZ 14306; Canada: Labrador; Apo- tomis frigidana (Packard); This species was de- scribed from two syntypes. Adamski and Peters (1986: 664) designated the lectotype (Code, Art. 74(b)). The male paralectotype is also in MCZ. frustrana Comstock, 1880: 236, Retinia; PSEUDO- TYPES (23 + ), MCZ 30298; United States: Massa- chusetts, Nantucket Island, S. H. Scudder; Rhy- acionia frustrana (Comstock); The name Retinia frustrana was published by both Comstock (1880) and Scudder (1883). These specimens are Scudder 's invalid type series. The lectotype in USNM des- ignated by Miller (1967: 591). ' fulvifrontana Packard, 1866: 59, Penthina; SYN- TYPES (2), MCZ 14305; Canada: Labrador; Oleth- reutes septentrionana (Curtis). glauoofuseana Zeller, 1875: 245, Conchylis; HO- LOTYPE female, MCZ 14303; United States: Tex- as, Dallas, Boll. gomonana Kearfott, 1907b: 78, Eucosma; PARA- LECTOTYPE, MCZ 15021; United States: New Jersey, Essex County, 5-11-1900, W. D. Kearfott; Lectotype in AMNH designated bv Klots (1942: 403); see also Heinrich (1923: 119). haimbachiana Kearfott, 1907a: 51, Epinotia; PARA- LECTOTYPE, MCZ 14300; United States: Penn- sylvania, Philadelphia, VL23-1904, F. Haimbach; Gypsonoma haimbachiana (Kearfott); Lectotype in AMNH designated by Klots (1942: 403); see also Heinrich (1923: 163). inimicella Zeller, 1872: 559, fig. 20, Galleria; HO- LOTYPE male, MCZ 14275; United States: Texas, Dallas, Boll; Pseudogalleria inimicella (Zeller). leucophaleratana Packard, 1866: 56, Pandemis; SYNTYPE, MCZ 14331; Canada: Labrador; An- cylis tineana (Huebner). longipalpana Powell, 1985: 67, Syllonoma; HOLO- TYPE male; United States: South Carolina, Horry County, Myrtle Beach, 9 July 1943, C. T. Parsons. marcidana Zeller, 1875: 260, Phoxopteris; SYN- TYPE male, MCZ 14330; United States: Texas, Dal- las, Boll; Ancylis platanana (Clemens). merrickanum Kearfott, 1907c: 156, pi. VIII, fig. 1, Exartema; PARALECTOTYPL, MCZ 15016; United States: Pennsylvania, New Brighton, VI 1-5- 1904, H. D. Merrick: Olethreutes merrickana (Kearfott); Lectotype in AMNH designated !>v Klots (1942: 405); see also Heinrich (1926: 149). miscana Kearfott, 1907a: 91, Eucosma; LECTO- PARATYPE, MCZ 15020; United States: Califor- nia, Placer County, Cisco; Epinotia miscana (Kear- fott); Lectotype in AMNH designated bv Klots (1942). murina Packard, 1867:60, Penthina; LECTOTYPE male, MCZ 14307; Canada: Labrador, Caribou Is- land; Olethreutes metallicana (Huebner); Lecto- type designated by Miller (1985: 410); also para- lectotype male. nebulosana Packard, 1866: 61, Grapholitha; SYN- TYPE male, MCZ 14312; Canada: Labrador; Gyp- sonoma nebulosana (Packard); An additional MCZ specimen may be a syntype. Heinrich (1923: 261) cites a male "type in the Fernald collection. niveosana Packard, 1866: 55, Sciaphila; SYNTYPES (6), MCZ 14313; Canada: Labrador; Eana niveo- sana (Packard). numerosana Zeller, 1876: 317, Paedisca; HOLO- TYPE male, MCZ 14339; United States: Texas, Dal- las, Boll; Epiblema numerosana (Zeller). ochromediana Kearfott, 1907a: 11, Olethreutes; PARALECTOTYPE, MCZ 14332; United States: Pennsylvania, Hazleton, 6/21/1902; Olethreutes osmundana (Fernald); Lectotype in AMNH des- ignated by Heinrich (1926: 170). olivaeeana Fernald, 1882: 71, Eccopsis; PARALEC- TOTYPES (2), MCZ 14334; United States: [no data]; Olethreutes olivaeeana (Fernald); Lectotype in USNM designated by Miller (1970: 292). osmundana Fernald, 1879: 156, Penthina; PARA- LECTOTYPE, MCZ 15010; United States: Maine. Orono, feeding on Osmunda regalis, emerged ] July 1879, A. Allen; Olethreutes osmundana (Fer- nald); Lectotype in USNM designated by Miller (1970: 292). packardi Zeller, 1875: 300, fig. 31. Grapholitha [sic]: HOLOTYPE, MCZ 14329; United States: Texas, Dallas, Boll; Missing abdomen. peculiana Zeller, 1875: 210, fig. 1, Teras; HOLO- TYPE female, MCZ 14316; United States: Texas, Dallas, Boll; Acleris subnivana (Walker). Udletin Museum of Comparative Zoology, Vol. 152, No. 2 perfluana Zeller. I 875 299, fig. 30, Grapholitha [sic]; HOLOTYPE female, MCZ L4327; United States: rexas, Dallas. Boll; Sereda tautana (Clemens). pinatubana Kearfott, 1905: 9, Eulia; PARALEC- rOTYPE, MCZ 143-44; United States: Massachu- setts. Winchendon, V-26-1902; Argyrotaenia pin- atubana (Kearfott); Lectotype in AMNH designated b> Klots (1942: 415). plumholineana Kearfott, 1907a: 53, Epinotia; PAR VLECTOTYPE, M( Z 14301; Canada: British ( olumbia, Wellington; Lectotype in AMNH des- ignated by Klots (1942i primulana Walsingham, 1S79: 45, pi. LXX, fig. 7, Paedisca; SYNTYPE, MCZ 15004; United States: ( alifornia, Mendocino County, 10 June 1871, Wal- Miiidiain. Eucosma primulana (Walsingham). prosperana Kearfott, 19071): 128, Enarmonia; PARALECTOTYPE, MCZ 15013; United States: ( alifornia, San Luis Obispo, HI-[no year], A. H. Vachell; Cydia prosperana (Kearfott); Lectotype in AMNH designated by Klots (1942: 408); see also Heinrich (1926: 57 ptychogrammos Zeller, 1875: 213, Teras hastiana va- riety ;HOLOTYPE male, MCZ 14317; United States: Texas, Dallas, Boll; Acleris ptychogrammos (Zell- er); Missing abdomen and hindwings. quintana Zeller, 1875: 304, figs. 34, 35, Paedisca; S^i NTYPES (2), MCZ 14340; United States: Texas, Dallas. Boll; Eucosma robinsonana (Grote). raracana Kearfott 1907a: 44, Thiodia; PARALEC- TOTYPE, MCZ 14320; United States: [no locality], ["8-12-99"]; Phaneta raracana (Kearfott); Lecto- type in \M\H designated by Heinrich (1923: 41). geriatana Zeller, 1875 244, Conchylis; HOLOTYPE male. MCZ 11301; United States: Texas, Dallas, Boll; Aethes seriatana (Zeller). sescuplana Zeller. 1 S75 220, Torlrix; SYNTYPES I \K IZ 14314; I nited States: Texas, Dallas, Boll; Clepsis virescana (Clemens). Bpaldingana Kearfott, 1907a: 19, Eucosma; PARA- LECTOTYPE, MCZ 14321; United States: Utah, Stockton, "VII-26-3", T Spaulding; Lectotype in \M\II designated by Klots (1942: 410); see also Heinrich 192 - M spiculana Zeller, L875 2s>) fig. 23, Grapholitha; HO- LOTYP1 female, M( :Z 1 132 l l nited States: Tex- Dallas, Boll; Phaneta spiculana (Zeller); Missing abdomen and tiulit forew ing. terracoctana Walsingham, 1879: 39, Paedisca; S1! N- TYP1 \K / r 5 I nit. d States California. Mount Shasta; Epinotia terracoctana (Walsingham); This species was described from 7 syntypes from Mi unit Shasta. Six syntypes and an additional 24 specimens from Mount Shasta and Mendocino City are now in the BMNH. Like other Walsingham California syntypes in the MCZ, this specimen does not bear labels in Walsingham's own handwriting, but was presumably received by Chambers. tessellana Packard, 1866: 58, Penthina; SYNTYPES (6), MCZ 14308; Canada: Labrador; Olethreutes intermistana (Clemens). testulana Zeller, 1875: 241, Cenopis; HOLOTYPE male, MCZ 14311; United States: Texas, Dallas, Boll; Sparganothis directana (Walker). trifurculana Zeller, 1875: 226, Tortrix; SYNTYPE, MCZ 14319; United States: Texas, Dallas, Boll; Ar- gyrotaenia quercifoliana (Fitch); Missing left fore- wing. tripartitana Zeller, 1875: 308, fig. 39, Paedisca; HO- LOTYPE female, MCZ 14337 United States: Texas, Dallas, Boll; Epiblema tripartitana (Zeller). tristriata Kearfott, 1907a: 67, Sparganothis; SYN- TYPE, MCZ 14310; United States: Minnesota, Du- luth, ["15. Fern/new"]; Missing right forewing, 1 cotype in AMNH (Klots, 1942: 416). variolana Zeller, 1875: 212, Teras; HOLOTYPE male, MCZ 14318; United States: Texas, Dallas, Boll; Acleris minuta (Robinson). vertumnana Zeller, 1875: 310, Paedisca; LECTO- TYPE female, MCZ 14336; United States: Texas, Dallas, Boll; Epinotia vertumnana (Zeller); Lec- totype designated by Brown (1987: 343). verutana Zeller, 1875: 247, Bactra lanceolana vari- ety; SYNTYPES (2), MCZ 14333; United States: Texas, Dallas, Boll; Bactra verutana Zeller. vestaliana Zeller, 1875: 286, fig. 21, Grapholitha [sic]; HOLOTYPE male, MCZ 14323; United States: Texas, Dallas, Boll; Hystrichophora vestaliana (Zeller); The holotype is a male, not a female as stated by Zeller. vitivorana Packard, 1869: 336, pi. 8, fig. 22, Penthina; SYNTYPE, MCZ 15008; United States: Ohio, Hud- son, M. C. Reed; Endopiza viteana Clemens; Miss- ing abdomen and right hindwing. worthingtoniana Fernald, 1878: 83, Paedisca; SYN- TYPE, MCZ 14342; United States: "North Illinois", [June], C. E. Worthington; Eucosma bipunctella (Walker). /ana Kearfott, 1907a: 61, Enarmonia; PARALEC- TOTYPE, MCZ 15015; Canada: British Columbia, Wellington, VI-[no year], T. Bryant; Grapholita caeruleana Walsingham; Lectotype in AMNH des- ignated In Klots ( 1 942: 412); see also Heinrich (1926: 31). MCZ Microlepidoptera Types • Miller and Hodges 71 SUPERFAMILY ZYGAENOIDEA Family Zygaenidae sanborni Packard, 1864a: 32, Harrisina; HOLO- TYPE [?], MCZ 27316; United States: Virginia, Al- exandria, J. O. Treat; Acoloithus falsarius Clemens; This may not be the holotype, which was "loaned ... by Mr. Sanborn." Family Limacodidae bifida Packard, 1864c: 338, Euclea; SYNTYPES [?] (2 males), MCZ 16028; United States: Maine, Bruns- wick, "at light", August; Euclea delphinii (Bois- duval). biguttata Packard, 1864c: 341, Limacodes; SYN- TYPE, MCZ 26382; United States: Pennsylvania [Harris no. 753]; Apoda biguttata (Packard); Two additional Harris specimens may be syntypes. cinereum Forbes, 1942: 389, Palaeophobetron; HO- LOTYPE male, MCZ 26268; Panama: Canal Zone, Barro Colorado Island, 31 December 1934, [A. Friedman]; Label states collector was A. Friedman, not M. Bates as stated by Forbes. ephippiatus Harris, 1869: 301, pi. I, fig. 7, pi. II, fig. 10, Limacodes; SYNTYPES (2 males, 2 females, 3 pupal cases), MCZ 33259; United States; "probably from Pennsylvania", T. Hill [Harris no. 775]; Acharia stimulea (Clemens) (see Becker and Miller, 1989). ferruginea Packard, 1864c: 338, Euclea; HOLO- TYPE female, MCZ 16027; United States: "St. Ca- tharines, C. W. (Coll. Scudder)"; Euclea delphinii (Boisduval); Specimen is a female, not a male as stated by Packard. geminata Packard, 1864c: 343, Cyrtosia; SYNTYPE male, MCZ 16031; United States: Maryland, Janes- ville, R. Stratton; Packardia geminata (Packard); Packard (1964) and Tietz ([1952]: 151) record syn- type(s) from Philadelphia in ANSP. monitor Packard, 1864c: 337, Euclea; SYNTYPES (2 males), MCZ 16029; United States: Massachusetts, Cambridge and Boston; Euclea delphinii (Bois- duval); Additional 2 male, 2 female syntypes in Harris Collection. testacea Packard, 1864c: 348, Tortricidia; SYN- TYPES (2), MCZ 26384; United States: [Harris no. 315]. y-inversa Packard, 1864c: 341, Limacodes; HOLO- TYPE, MCZ 26383; United States: Pennsylvania [Harris no. 781]; Apoda ij-inversa (Packard); A pseudotype in main collection, MCZ 16030. Family Megalopygidae crispata Packard, 1864c: 335, Lagoa; SYNTYPES (1 male, 1 female), MCZ 27315; United States: Mas- sachusetts, Brookline, C. A. Shurtleff; Megalopyge crispata (Packard); Packard notes "thirteen speci- mens reared from the blackberry bush by Mr. Shur- tleff." immaculaia Cassino, 1928. 91, Lagoa; HOLOTYPE male, MCZ 32865; United States: Texas, Davis Mountains, 1-7 February 1926, O. C. Poling; Me- galopyge immaculaia (Cassino); Probably a syn- onym of Megalopyge laycei (Barnes and Mc- Dunnough). pulla Forbes, 1942: 401, Trosia; HOLOTYPE fe- male, MCZ 26269; Panama: Canal Zone, Barro Col- orado Island, 10 October 1934, M. Bates. SUPERFAMILY PYRALOIDEA Family Pyralidae albiplagiatella Packard, 1873b: 269, Myelois; SYN- TYPES (2 males of 2), MCZ 14291; United States: New Hampshire, May and June, C. A. Walker; Pima albiplagiatella (Packard). anticostalis Grote, 1871: 104, Botys; SYNTYPES (1 male, 1 female), MCZ 14263; United States: Ala- bama, "about Demopolis"; Eulepte anticostalis (Grote). argillaceellus Packard, 1867: 54, Crambus; SYN- TYPES (2), MCZ 14295; Canada: Labrador, Square Island, 14 July 1864; Raphiptera argillaceella (Packard). betulella Hulst, 1890: 125, Acrobasis; PARALEC- TOTYPE female, MCZ 14284; United States: New Y'ork, July; Lectotype in AMNH designated by Neunzig (1986: 59). borealis Packard, 1867: 53, Pyrausta; HOLOTYPE female, MCZ 14270; Canada: Labrador, Square Is- land, 14 July [1864]; Pyrausta subsequalis borealis Packard. caeculalis Zeller, 1875: 333, pi. 10: fig. 46, Perispasta; HOLOTYPE male, MCZ 14259; United States: Texas, Dallas, Boll. californicalis Packard, 1873a: 264, Eromene; LEC- TOTYPE male, MCZ 14297; United States: Cali- fornia, H. Edwards; Euchromius californicalis (Packard); Capps (1966: 5) saw two of the three males now in the collection, and designated one of the two as lectotype, without so labelling it. Since there are three specimens, not two. it is not clear which one is the lectotype. One of the three may not be a syntype because it was collected by Beh- rens, not Edwards. Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 carpenterellus Packard, L874: 348, fig. 1, Crambus; SYNTYP1 S I ol 8), MCZ 14299; United States: ( olorado, "Mountains of Colorado", 19 July, 12 Vugust, and 8 September 1873, W. L. Carpenter; ( 'rambus hamellus carpenterellus Packard; Brown L972: 217) restricted the type locality to Weston lass. Park County, Colorado. comptoniella Hulst, 1890; 125. Acrobasis; LECTO- H PE male, MCZ 14283; United States: New York, Long Island, July; Lectotype designated by Neun- zig (1986 56] consobrinella Zeller L872: 548, Nephopteryx; HO- LOTYPE male, MCZ 14287; United States: Texas, Dallas, Boll; Glyptocera consobrinella (Zeller). cuprina Zeller. L872: 497, Aglossa; SYNTYPES (1 male. 1 female), MCZ 14257; United States: Texas, Dallas, Boll. decinierella Hulst, 1SSS: 1 17, Lipographis; Probable S . \ n PE female, MCZ 14288; United States: Tex- as, Blanco County, August; Stylopalpia scobiella Grote). edmandsii Packard. 1864b: 120, Nephopteryx; SYN- TYPE, MCZ 14278; United States: Massachusetts; Vitula edmandsii (Packard). electella Hulst. 1887: 137, Anerastia; Probable SYN- TYPE, MCZ 14279; United States: Texas, Blanco County; Homoeosoma electella (Hulst). fenestrella Packard, 1873a: 259, Pempelia; SYN- TYPES (2 of 6), MCZ 14292; United States: Cali- fornia, H. Edwards; Lipographis fenestrella (Pack- ard . Edwards number 711 indicates Angel Island, \pril. feriella Hulst. 1888: 115, Tacoma; Probable SYN- n PE, MCZ 14286; United States: Texas, [Blanco ( ount) . August]. feudalis Grote 1875: 231. Botis; SYNTYPE, MCZ L6095; I nited States New York and Massachusetts; Herpetogramma theseusalis (Walker). frigidella Packard. 1867: 53, Eudorea; SYNTYPE, MCZ 14296; Canada: Labrador, Caribou Island; Pyla fusca (Haworth); Abdomen missing. fulminalis Zeller I S72 560, fig. 19, Melissoblaptes; HOLOTYPE male. MCZ 11276; United States: Texas, Dallas. Hoik Paralipsa fulminalis (Zeller). glacialis Packard L867: 52, Botys; HOLOTYPE fe- male MCZ L5354 Canada: Labrador, fiopedale, - \uuust ishi ( dea inquinatalis (Zeller); Packard 1867 indicates one sex onl) and no range of size. hospitella Zeller 1875 338, Ephestia; PARALEC- rOTYPES 6), MCZ 1 1277. I nited States: Texas. rythmia hospitella (Zeller); Lec- in PM\1I designated by Heinrich (1956: integra Zeller, 1873: 328, fig. 44, Scoptonoma; SYN- TYPES (3), MCZ 16094; United States: Texas, Dal- las, Boll; Lineodes integra (Zeller); Although la- belled as types by Hagen, these specimens were probably not seen by Zeller, who described the species from "Ein paar in meiner Sammlung, meh- rere nach Hagens Angabe im Cambridger Mu- seum." interrupta Zeller, 1873: 329, Scoptonoma; SYN- TYPES (4), MCZ 16093; United States: Texas, Dal- las, Boll; Lineodes interrupta (Zeller); As with Scop- tonoma integra above, Zeller probably did not see these specimens. latifaseiatella Packard, 1873b: 269, Nephopteryx; HOLOTYPE female, MCZ 14290; United States: Maine, A. S. Packard; Telethusia ovalis (Packard). leoninella Packard, 1873a: 259, Pempelia; SYN- TYPES (2 of 3), MCZ 14293; United States: Cali- fornia, H. Edwards; Lipographis leoninella (Pack- ard); Edwards number 706 indicates San Mateo County, pastures, May. lentiflualis Zeller, 1872: 525, Homophysa; SYN- TYPE, MCZ 14274; United States: Texas, Dallas, Boll; Aethiophysa lentiflualis (Zeller). melanogrammos Zeller, 1872: 546, fig. 24, Tetralo- pha; HOLOTYPE male, MCZ 14256; United States: Texas, Dallas, Boll; Specimen illustrated by Hol- land and Schaus (1925: 65). metalliferalis Packard, 1873a: 265, Calaclysta; SYN- TYPES (2 of 4), MCZ 33257; United States: Cali- fornia, H. Edwards; Dicymolomia metalliferalis (Packard); Edwards number 208 indicates San Ma- teo County, marshy places. May. mustelinalis Packard, 1873a: 262, Botys; SYNTYPES (2 of 2), MCZ 14269; United States: California, H. Edwards; Mecyna mustelinalis (Packard); One specimen has Edwards number 773 indicating An- gel Island, May. occidentalis Packard, 1873a: 260, Scopula; LEC- TOTYPE female, MCZ 14264; United States: Cal- ifornia, H. Edwards; Achyra occidentalis (Pack- ard); Lectotype designated by Capps (1967: 51). Edwards number 716 indicates Point Lobos, pas- tures, May. ochrifrontella Zeller, 1875: 337, Ephestia; HOLO- TYPE male, MCZ 14280; United States: Texas, Dal- las, Boll; Eulogia ochrifrontella (Zeller). octonalis Zeller, 1873: 211, Orobena; HOLOTYPE male, MCZ 14273; United States: Texas, Dallas, Boll; "Lygropia" octonalis (Zeller) (needs new ge- nus according to Munroe, pers. comm.). ovalis Packard, 1873b: 269, Pempelia; HOLOTYPE, MCZ I42N9; t nited States: Maine, A. S. Packard; Telethusia ovalis (Packard); Missing abdomen. MCZ Microlepidoptera Types • Miller and Hodges 73 perrubralis Packard, 1873a: 264, Botys; SYNTYPES (2 of 3), MCZ 14267; United States: California; Pyrausta perrubralis (Packard). plumbicostalis Grote, 1871: 103, Botys; HOLOTYPE male, MCZ 32943; United States: Alabama, "about Demopolis"; Lygropia plumbicostalis (Grote); Pro- thorax, left forewing, and right wings only. profundalis Packard, 1873a: 261, Botys; SYNTYPES (2 of 5), MCZ 14271; United States: California, H. Edwards; Udea profundalis (Packard); Edwards number 705 indicates San Mateo Countv, pastures, May. reniculalis Zeller, 1872: 526, Homophysa; SYNTYPE male, MCZ 14261; United States: Texas, Dallas, Boll; Nephrogramma reniculalis (Zeller). roseatella Packard, 1873b: 270, Nephopteryx; LEC- TOTYPE male, MCZ 14282; United States: Mas- sachusetts, Dorchester, F. G. Sanborn; Peoria ap- proximella (Walker); Lectotype, and paralectotype also in MCZ, designated by Shaffer (1968: 30). rubrifasciella Packard, 1873b: 267, Acrobasis; SYN- TYPES (4 of 15), MCZ 14285; United States: Maine, Orono, A. S. Packard; See Neunzig (1986: 57) for discussion of identity of types. semirubralis Packard, 1873a: 263, Botys; HOLO- TYPE male, MCZ 14268; United States: California, [Sausalito, 6 May 1872], H. Edwards, Pyrausta se- mirubralis (Packard); A second male from "S. Ne- vada" (Sierra Nevada Mountains of California) is also labelled "type" in Packard's handwriting, but is probably not the holotype, since there are other specimens in the MCZ with the same data but with- out type labels. serratissimalis Zeller, 1872: 521, Crocidophora; PARALECTOTYPE female, MCZ 14262; United States: Texas, Dallas, Boll; Lectotype in BMNH designated by Monroe (1976: 22). sesquialteralis Zeller, 1873: 209, fig. 5, Botis; HO- LOTYPE male, MCZ 32924; United States: Texas, Dallas, Boll; Microtheoris o. ophionalis (Walker). sincera Zeller, 1875: 332, Oectoperia; SYNTYPES (2 of 2), MCZ 14258; United States: Texas, Dallas, Boll; Salobrena sincera (Zeller). subdivisalis Grote, 1871: 126, Desmia; HOLOTYPE [?], MCZ 14260; United States: Alabama, "about Demopolis"; Specimen consists of right wings only. subolivalis Packard, 1873a: 261, Botys; SYNTYPES (2), MCZ 14266; United States: Maine, Brunswick, in grass uplands, and Orono, Maine, A. S. Packard, Jr.; Pyrausta unifascialis subolivalis (Packard). tetradella Zeller, 1872: 552, Anerastia; PARALEC- TOTYPES (2), MCZ 14281; United States: Texas, Dallas, Boll; Peoria tetradella (Zeller); Lectotype in BMNH designated by Shaffer (1968: 17). unifascialis Packard, 1873a: 261, Botys; SYNTYPES (2 of 2), MCZ 14265; United States: California, H. Edwards; Pyrausta unifascialis (Packard); Klots (1942: 422) designated a lectotype and paralecto- type in AMNH. However, the MCZ specimens are probably the real types, since Packard kept the other types of California Pyralidae from Edwards. Beutenmueller(1892: 196) listed one type in Wl\ll unistriatellus Packard, 1867: 54, Crambus; SYN- TYPE male, MCZ 22641; Canada: Labrador, Car ibou Island. vibicalis Zeller, 1873: 208, fig. 4, Botis; HOLOTYPE male, MCZ 14272; United States: Texas, Dallas. Boll; Microtheoris vibicalis (Zeller) (see Munroe, 1972: 147, about misspelling of name as ribicalis). Family Thyrididae dimidiata Forbes. 1942: 345, Bhodoneura; HOLO- TYPE female, MCZ 26263; Panama: Barro Colo- rado Island, 2-XII-1934, M. Bates; Missing abdo- men. longalis Forbes, 1942: 340, Dysodia; HOLOTYPE male, MCZ 26264; Panama: Barro Colorado Island, 29-XI-1934, M. Bates. maculata Harris 1839: 313, Thyris; SYNTYPES (4), MCZ 26346; United States: [New Hampshire, Har- ris no. 219]; All missing abdomens. mesogramma Forbes, 1942: 341, Ochrothyris; HO- LOTYPE male, MCZ 26265; Panama: Barro Col- orado Island, 10-1-1935, A. Friedman. SUPERFAMILY PTEROPHOROIDEA Family Pterophoridae cervinidactylus Packard, 1873a: 266, Pterophorus; HOLOTYPE, MCZ 1785; United States: Califor- nia, H. Edwards; Platyptilia pallidactyla (Ha- worth); Missing right wings and abdomen. cineraceus Fish, 1881: 73, Oidaematophorus; SYN- TYPE male, MCZ 1781; United States: Washington Territory, H. K. Morrison; Barnes and Lindsey (1921: 392) consider this specimen a paratype. grandis Fish, 1881: 141, Lioptilus; "LECTOTYPE" male, MCZ 1782; United States: California; Oidae- matophorus grandis (Fish); Cashatt (1972: 5) des- ignated this specimen as lectotype, unaware of a previous lectotype designation by Klots (1942: 423) of a female in the AMNH. lacteodactylus Chambers, 1873: 72, Pterophorus; HOLOTYPE male, MCZ 1783; I nited States: Ken- tucky, Chambers; Oidaematophorus lacteodacty- lus (Chambers); Missing abdomen. paleaceus Zeller, 1873: 326, LeJoptilus; SYNTYPES Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 M< / L787; I nited States: Texas, Dallas, Boll; Oidaematophorus paleaceus (Zeller). pergracilidactylus Packard, 1873a: 265. Pteropho- rus; HOLOTYPE, MCZ L786; United States: Cal- ifornia, H. Edwards; Emmelina monodactyla (Lin- naeus); Missing left wings and abdomen. pumilio Zeller. L873: 324, Mimesoptilus; HOLO- n I'l. male. MCZ 1788; United States: Texas, Dal- las, Boll: Marasmarcha pumilio (Zeller). x-micostatus Zeller. 1873: 323. Mimeseoptilus; SYN- TYPES (2 of 2), MCZ 1789; United States: Texas, Dallas, Boll; Stenoptilia zophodactyla (Dupon- chel); One male and one female, although Zeller stated two males. sulphureodactylus Packard, 1873a: 266, Pteropho- rus; SYNTYPES (6 of 8), MCZ 1784; United States: California, Siskiyou County, Goose Lake, 26-27 July, J. Holleman; Oidaematophorus sulphureo- dactylus (Packard). APPENDIX Chambers "types" in the British Museum (Natural History) The collection of the BMNH contains specimens sent by Chambers to Stainton in the 1870s. Some of them probably are syntypes of Chambers' species, others are not syntypes but are topotyes, and others are taxa described by other authors. The topotypes might prove useful to document Chambers' species concepts, or as neotype candidates. There are three lots of Chambers spec- imens now in the Stainton collection. The \ngust 1877 shipment to Stainton was ac- companied by the following letter from Chambers: "Wishing to preserve types of species of Tineina described by me, and likewise to make some acknowledgement ... it has been my intention to send you as complete a collection of the species of this region [Covington, Kentucky] as 1 could make. 1 nfortunately during my absence in Col- orado more than half the collection that I had made before I went there was de- stroyed, and the remainder with a small collection that I made in Colorado, was senl to the . [MCZ]. This spring and sum- mer I have attempted to supply a few cab- in this country and your own. Un- fortunately the continued ill-health of my son again calls me to Colorado and un- willing to risk the destruction of my col- lection while I am absent I distribute it — so far as I have renewed it — now. I enclose here with 67 species of Tineina and one of Tortricina." The specimens are listed here as iden- tified by Chambers. We have made no attempt to verify identifications. The lo- calities listed here are those indicated by Chambers in correspondence or on labels, and are not necessarily the type localities. The first lot, sent December 1876, includes Lithariapteryx abroniaeella Chamber "from Colorado where the larva mines the leaves of Abronia fragrans up to an alti- tude of about 7,000 feet above sea level" and Batrachedra praeangusta Chambers from Colorado. The second lot, sent in January 1877, now includes 5 species, but originally in- cluded 6. 1. Lithariapteryx abroniaeella Chambers Colorado 2. Lithocolletis salicifoliella Chambers 3. Lithocolletis amphicarpeaeella Cham- bers Kentucky 4. Laverna magnatella Chambers = La- verna oenotheriella Chambers 5. Gracilaria [sic] salicifoliella Chambers 6. Eurynome albella Chambers Colorado [must have been damaged in transit, not at BMNH] The largest lot was sent in August 1877. All bear Chambers' handwritten numbers and Stainton's name labels. The numbers not cited in the following list refer to species described by authors other than Cham- bers. 1 . Xylesthia clemensella Chambers Ken- tucky 2. Tinea bimaculella Chambers Ken- tucky 3. Tinea caemetariiella [sic] Chambers Kentucky 5. Semele cristatella Chambers Ken- tucky MCZ Microlepidoptera Types • Miller and Hodges 75 6. Agnippe biscolorella Chambers Ken- tucky 7. Hyponomeuta longimaculella Cham- bers Kentucky 8. Depressaria eupatoriiella Chambers Kentucky 9. Gelechia querciella Chambers Ken- tucky 10. Gelechia cristatella Chambers Ken- tucky 1 1 . Gelechia rubensella Chambers Ken- tucky 13. Helice pallidochrella Chambers Ken- tucky 17. Argyresthia undulatella Chambers Kentucky 18. Gracilaria packardella [sic] Chambers Kentucky 19. Corisium albanotella Chambers Ken- tucky 20. Gracilaria [sic] fasciella Chambers Kentucky 21. Gracilaria [sic] purpuriella Chambers Kentucky 23. Coleophora ochrella Chambers Ken- tucky 24. Coleophora caryaefoliella Chambers Kentucky 25. Laverna cephalanthiella Chambers Kentucky 26. Chrysopelia purpuriella Chambers Kentucky 27. Laverna? gleditschiaeella Chambers Kentucky 28. Perimede erransella Chambers Ken- tucky 30. Dryope murtfeldtella Chambers Ken- tucky 31. Tischeria quercivorella Chambers Kentucky 32. Tischeria heliopsisella Chambers Kentucky 33. Bucculatrix luteella Chambers Ken- tucky 35. Philonome clemensella Chambers Kentucky 36. Phyllocnistis vitifoliella Chambers Kentucky 37. Phyllocnistis ampelopsiella Cham- bers Kentucky 40. Leucanthiza amphicarpeaefoliella Chambers Kentucky 41. Lithocolletis clemensella Chambers Kentucky 43. Lithocolletis fuscocostella Chambers Kentucky 44. Lithocolletis celtisella Chambers Kentucky 45. Lithocolletis cincinnatiella Chambers Kentucky 47. Lithocolletis ulmella Chambers Ken- tucky 49. Lithocolletis tiliaeella Chambers Kentucky 50. Lithocolletis ornatella Chambers Kentucky 51. Lithocolletis corylisella Chambers Kentucky 52. Lithocolletis ambrosiaeella Chambers Kentucky 53. Polyhymno sexstrigella Chambers Texas 54. Perimede unomaculella Chambers Texas 55. Elachista parvipulvella Chambers Texas 58. Ornix prunivorella Chambers Ken- tucky 59. Gelechia fuscoochrella Chambers Kentucky 60. Lithocolletis tritaeniaella Chambers Kentucky 61. Nepticula apicialbella Chambers Kentucky 63. Gracilaria 12-lineella Chambers Ken- tucky 64. Gelechia bimaculella Chambers Ken- tucky 65. Lithocolletis corylisella Chambers Kentucky 66. Gelechia variiella [sic] Chambers Ken- tucky 68. Microaethia amphicarpeaeana Chambers Kentucky The correspondence also indicates a shipment in November 1872 which in- cluded 41 Chambers species. However, these specimens could not be located at the BMNH. Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 LITERATURE CITED \DAMSKI, D. VND T. M Pi ii RS. 1986. Review of Nearctic ipotomis Huebner (Lepidoptera: Tor- tricidae: Olethreutini) Canadian Entomologist, UK: 649 689 Becker, V. O., and S. E. Miller. 1989. The iden- tic of Sphinx brunnus Cramer and the taxonom- ic position of Acharia Huebner (Lepidoptera: Li- macodidae). Journal of Research on the Lepidoptera 26: 219-224 Bl i ii wii ii i ik. W. 1889. Chambers' corrections to his paper on the illustrations of the neuration of the wings of American Tineidae. Entomolo- gica Americana, 5: 37-38. . 1892. List of types of Lepidoptera in the Edwards collection of insects. Bulletin of the American Museum of Natural History, 4: 167- 198. BRAUN, A. F. 1948. Elachistidae of North America Microlepidoptera). Memoirs of the American Entomological Society, 13: 110 + ii pp., pi. I- \\\ I . 1963. The genus Bucculatrix in America north of Mexico (Microlepidoptera). Memoirs of the American Entomological Society, 18: iii + 208 + ii pp., pi. I-XLV. . 1972. Tischeriidae of America north of Mexico i Microlepidoptera). Memoirs of the American Entomological Society, 28: i + 148 pp. BROWN.F.M. 1972. The type locality for two moths (Pyralididae [sic], Saturniidae) collected by Lt. W L. Carpenter, U.S.A., in Colorado, 1873. Jour- nal of the Lepidopterists' Society, 26: 245-247. BROWN, R. L. 1987. Resurrection of Catastega Cle- mens and revision of the Epinotia vertumnana (Zeller) species-group (Tortricidae: Olethreutin- ae). Journal of the Lepidopterists' Society, 40: 327-346. ("1986"] Bl S< k V 1903 A revision of the American moths nl the- family Gelechiidae, with descriptions of new species. Proceedings of the United States National Museum, 25: 767-938. Cuts H W. 1966. Review of New World moths of the genus Euchromius Guenee with descrip- tions oi two new species (Lepidoptera: Crambi- dae). Proceedings ol the I nited States National Museum, I 19(3551): 1-9. . 1967 Review of some species of Loxostege Huebner and descriptions ol new species (Lep- idoptera, Pyraustidae: Pyraustinae). Proceedings of the United States National Museum. 120(3561): 1-75. HAT1 E I) 1972 Notes on the halanotes Meyrick) group ol Oidaematophorus Wallen- gren with description ol a new species (Ptero- phoridae [ournal of the Lepidopterists' Societ) 26: I INO S ! iiieneu Lasio* ainpidae. The iterist, 4: 89 96 IsTUa \ singular case. Canadian Entomologist. 2: 43-44. [saturniid larvae and their parasites] — . 1870b. Harmless parasites on the larva of the luna moth. American Entomologist, 2: 125. . 1870c. Time of the appearance of the poly- phemus moth in Louisiana and Kentucky. Amer- ican Entomologist, 2: 156. . 1870d. An anomalous grape sphinx moth. American Entomologist, 2: 210. . 1870e. A rare capture. American Ento- mologist, 2: 242. [on Phymaphora pulchella] . 1870f. Salt marsh caterpillar. American Entomologist, 336. . 1871a. A new species of Cemiostoma (Mi- cro-Lepidoptera, Tineina). Canadian Entomol- ogist, 3: 23-25. . 1871b. Micro-Lepidoptera. Canadian Entomologist, 3: 54-58, 84-88, 108-112, 127- 130, 146-149, 161-166, 182-185, 205-209, 221- 224. 1872a. Micro-Lepidoptera. Canadian Entomologist, 4: 7-12, 25-29, 41-44, 65-69, 92, 106-108, 126-133, 146-150, 169-175, 191- 195, 206-209, 222-226. . 1872b. Microlepidoptera. American Nat- uralist, 6: 432-433. [notes on how to collect] . 1872c. Notes on Cemiostoma. American Naturalist, 6: 489-490. . 1873a. Micro-Lepidoptera. Canadian Entomologist, 5: 12-15, 44-50, 72-75, 85-91, 110-115, 124-128, 147-152, 173-176, 185-190, 229-232. . 1873b. Cemiostoma again. American Nat- uralist, 7: 47-49. . 1874a. Micro-Lepidoptera. Canadian Entomologist, 6: 8-11, 49-52, 72-77, 96-97, 128- 130, 149-153, 166-170, 197-198, 217-220. . 1874b. Tineina from Texas. Canadian Entomologist, 6: 229-249. . 1874c. Prof. Frey, of Zurich, and some American Teneina [sic]. Cincinnati Quarterly Journal of Science, 1: 193-211. . 1874d. Notes and errata on a former paper, by V.T. Chambers, on Prof. Frey, and some American Teneina [sic]. Cincinnati Quarterly Journal of Science, 1: 338-340. . 1875a. Tineina of the Central United States. Cincinnati Quarterly Journal of Science, 2: 97- 121. . 1875b. Teneina [sic] of the United States. Cincinnati Quarterly Journal of Science, 2: 226- 259. 1875c. Teneina [sic] of Colorado. Cincin- nati Quarterly Journal of Science, 2: 289-305. — . 1875d. Tineina from Texas. Canadian Ento- mologist, 7: 7-12, 30-35, 51-56, 73-75, 92-95, 105-108. . 1875e. Tineina from Canada. Canadian Entomologist, 7: 124-128, 144-147, 209-213. — . 1875f. On some European 'micros' away from home. Entomologist's Monthly Magazine, 11: 279-280. MCZ Microlepidoptera Types • Miller and Hodges 11 . 1876a. Micro-Lepidoptera. Canadian Entomologist, 8: 18-19. . 1876b. Tineina. Canadian Entomologist, 8: 30-35, 103-106, 135-138, 158-160, 171-173, 217-220. . 1876c. Correpsondence [sic]. Canadian Entomologist, 8: 39-40. [on denuding wings of Lepidoptera] . 1877a. Tineina. Canadian Entomologist, 9: 13-15, 108-110, 123-127, 145-147, 194-196, 206-208. . 1877b. Tineina from Texas. Canadian Entomologist, 9: 22-26, 71-74. . 1877c. Correspondence. Canadian Ento- mologist, 9: 38-40. . 1877d. The Tineina of Colorado. Bulletin of the United States Geological and Geographical Survey of the Territories, 3: 121-142. . 1877e. Notes on a collection of tineid moths made in Colorado in 1875 by A. S. Packard, Jr., M. D. Bulletin of the United States Geological and Geographical Survey of the Territories, 3: 143-145. . 1877f. On the distribution of Tineina in Colorado. Bulletin of the United States Geolog- ical and Geographical Survey of the Territories, 3: 147-150. . 1877g. Insects feeding on Gleditschia. Ca- nadian Entomologist, 9: 231-235. . 1877h. Notes upon the American species of Lithocolletis . Psyche, 2: 81-87. . 1878a. Tineina. Canadian Entomologist, 10: 50-54. . 1878b. Micro-lepidoptera. Canadian Ento- mologist, 10: 74-78, 109-114, 238-239. . 1878c. Descriptions of new Tineina from Texas, and others from more northern localities. Bulletin of the United States Geological and Geo- graphical Survey of the Territories, 4:79-106. . 1878d. Index to the described Tineina of the United States and Canada. Bulletin of the United States Geological and Geographical Sur- vey of the Territories, 4: 125-167. . 1878e. On Pronuba yuccasella (Riley), and the habits of some Tineina. Journal of the Cin- cinnati Society of Natural History, 1: 141-154. . 1878f. Tineina and their food-plants. Bul- letin of the United States Geological and Geo- graphical Survey of the Territories, 4: 107-123. . 1878g. On larvae of Tineina, especially of Lithocolletis. Psyche, 2: 137-153. . 1879a. Micro-lepidoptera. Canadian Ento- mologist, 11: 5-10, 72-75, 89-93, 118-119, 125- 127. . 1879b. Tineina. Canadian Entomologist, 11: 143-146. . 1879c. Annual address of V.T. Chambers, Esq., President Cincinnati Society of Natural History. Journal of the Cincinnati Society of Nat- ural History, 2: 71-92, 1 pi. . 1879d. [corrections to Chambers, 1878g]. Psyche, 2: 227. . 1880a. Some new species of Tineina from North America. Canadian Entomologist, 12: 225- 226. . 1880b. Descriptions of some new Tineina, with notes on a few old species. Journal of the Cincinnati Society of Natural History, 2: 179- 194. . 1880c. Illustrations of the neuration of the wings of American Tineina. Journal of the Cin- cinnati Society of Natural Historv, 2: 194-199, 4 pis. . 1880d. On the changes that take place in the mouth-parts and legs of some leaf-mining lepidopterous larvae. American Entomologist, 3: 255-262. . 1880e. Notes upon some tineid larvae. Psy- che, 3: 63-68. . 1880f. Further notes on some tineid larvae. Psyche, 3: 135-137, 147-149. . 1880g. Wandering habit of larvae belong- ing to the genus Bucculatrix. American Ento- mologist, 3: 50. 1880h. Insects injuring the black locust (Ro- binia pseudacacia). American Entomologist, 3: 59-61. . 1880L Pronuba vs. Prodoxus. American Entomologist, 3: 177. . 1881a. On the early stages of Gracilaria [sic] stigmatella, Fabr. Canadian Entomologist, 13: 25-28. . 1881b. On the larva of Mordella. Canadian Entomologist, 13: 173-175. . 1881c. On some North American Tineidae, bv Thomas, Lord Walsingham, F.Z.S. Canadian Entomologist, 13: 191-194. . 1881d. New species of Tineina. Journal of the Cincinnati Society of Natural History, 3: 289- 296. . 1881e. Nepticula pteliaeella, n.sp. Psyche, 3: 276. . 1882a. Notes on the larva of Bucculatrix ambrosiaefoliella. Canadian Entomologist, 14: 153-160. . 1882b. Burrowing larvae. Nature, 25: 529. . 1882c. "Tineidae" or "Tineina." Papilio, 2: 115-119. . 1882d. On the antennae and tropin of Lep- idopterous larvae. Journal of the Cincinnati So- ciety of Natural History, 5: 5-24. . 1883. The classification of the Tineidae. Psyche, 4: 71-74. COMSTOCK, J. H. 1880. Report of the entomologist, pp. 185-348 In Annual Report of the I nited States Commissioner of Agriculture for 1879. Washington, D.C [not seen] Davis, D. R. 1967. A revision of the moths of th( subfamily Prodoxinae (Lepidoptera: Incurvari- idae). Bulletin of the United States National Mu- seum, 255: 1-170. . 1978. A revision of the North American moths of the Superfamily Eriocranioidea with the proposal of a new family, Acanthopterocte- Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 tidae (Lepidoptera) Sinithsonian Contributions t\\ l I I . J. A. 1964. Biological and taxonomic stud- it's on tortricine moths, with reference to the species in California. Llniversity of California Publications in Entomology, 32: iv + 317 pp. . 1968. Host associations and taxonomy of Nearctic conifer cone moths in the genus Eu- cosma (Lepidoptera: Tortricidae). Hilgardia, 39: 1-36. . 1973. A systematic monograph of New World ethmiid moths (Lepidoptera: Gelechioi- dea). Smithsonian Contributions to Zoology, 120: iv + 302 pp. 1985. Discovery of two new species and genera of shaggy tortricids related to Synnoma and NiOSOma (Tortricidae: Sparganothini). Jour- nal ol Research on the Lepidoptera, 24: 61-71. Pi RRINGTON, F. F. \\i)D. G. NIELSEN. 1987. Dis- covery ot the T. W. Harris collection at Harvard I niversity and designation of a lectotype for Po- dosesia syringae Harris (Lepidoptera: Sesiidae). Proceedings of the Entomological Society of Washington, 89: 549-551. RlDOl I. B V. 1977 Two new synonymies of Mi- i rolepidoptera (Gelechiidae: Gnorimosehemini and Oecophoridae: Deprcssariinae). Entomolo- gists Gazette, 28: 38 12 Bui-, C. V 1887. Report of the Entomologist, pp. \~>') 592, pis I \l in Report of the Commis- sioner nt Viniculture L886. Washington, DC. 719 PP L889 Two brilliant and interesting micro- lepidoptera new to our fauna. Proceedings of the Entomological Societ) of Washington, 1: 155- 1 59 RlNDGE, F H L955 The type material in the J. B. Smith and G D Hulst collections o! Lepidoptera iii the American Museum ol Natural History. Bulletin ol the American Museum of Natural lli-tmv, (06: 91 172. Hoiunson (. s L986 Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae). Bulletin of the British Museum (Natural History). Ento- mology, 52: 37-181. Sutler, K. 1962. Ein Beitrag zur Geschichte der Mikrolepidopterologie. Entomologische Zeit- schrift, 72(13): 1-3. . 1979. A taxonomic revision of the genus Deltophora Janse, 1950 (Lepidoptera: Gelechi- idae). Bulletin of the British Museum (Natural History), Entomology, 38: 263-322. ScLDDER, S. H. 1883. The pine moth of Nantucket, Retinia frustrana. Publication of the Massachu- setts Society for the Promotion of Agriculture, [not seen] Shaffer, J. C. 1968. A revision of the Peoriinae and Anerastiinae (Auctorum) of America North of Mexico (Lepidoptera: Pyralidae). Bulletin of the United States National Museum, 280: 1-124. TlETZ, H. M. [1952]. The Lepidoptera of Pennsyl- vania. Pennsylvania State College, State College, Penn. xii + 194 pp. WAGNER, D. L. 1988. Taxonomic status of Korschel- tellus Borner in North America (Lepidoptera: Hepialidae). Journal of the New York Entomo- logical Society 96: 345-354. Walsingham, Lord. 1879. Illustrations of typical specimens of Lepidoptera Heterocera in the col- lection of the British Museum. Part IV. - North- American Tortricidae. British Museum (Natural History), London, xi + 84 pp., pi. LXI-LXXVII. . 1881. On some North-American Tineidae. Proceedings of the Zoological Societv of London, 1881: 301-325, pi. XXXV, XXXVI. . 1882. Notes on Tineidae of North America. Transactions of the American Entomological So- ciety, 10: 165-204. 1889. Steps toward a revision of Chambers index, with notes and descriptions of new species. Insect Life, 2: 23-26. Wilkinson, C. 1979. A taxonomic study of the micro-lepidopteran genera Microcalyptris Braun and Fomoria Beirne occurring in the United States of America (Lepidoptera, Nepticulidae). Tijdschrift voor Entomologie, 122: 59-90. Wilkinson, C, and M. J. Scoble. 1979. The Nep- ticulidae (Lepidoptera) of Canada. Memoirs of the Entomological Society of Canada, 107: 1- 129. ZELLER, P. C. 1872. Beitraege zur Kenntniss der nordamericanischen Nachtfalter, besonders der Microlepidopteren. Erste Abtheilung. Verhand- lungen der Zoologisch-Botanischen Gesellschaft in Wien, 22: 447-566, pi. II— III. — . 1873. Beitraege zur Kenntniss der norda- mericanischen Nachtfalter, besonders der Mi- crolepidopteren. Zweite Abtheilung. Verhand- lungen der Zoologisch-Botanischen Gesellschaft in Wien, 23: 201-334, pi. III-1V. 1875. Beitraege zur Kenntniss der norda- mericanischen Nachtfalter, besonders der Mi- crolepidopteren. Verhandlungen der Zoologisch- Botanischen Gesellschaft in Wien, 25: 207-360, pi. VIII-X. MCZ Microlepidoptera Types • Miller and Hodges 81 INDEX Original names of all taxa cited are indexed, along with present names if the present species name differs (e.g., present names for which only the generic name differs from the original are not indexed). Family names and significant people cited in the introduction are also indexed. abroniaeella, Lithariapteryx, 67, 74 absconditella, Monochroa, 64 acapnopennella, Nemapogon, 52 Adamski, D., 56 aenea, Tischeria, 49 aesella, Heliozela, 50 agnella, Bucculatrix, 54 Agonoxenidae, 61 alba, Tegeticula, 50 albaciliaeella, Strobisia, 55 albalineella, Eriphia, 61 albanotella, Corisium 75 albapenella, Butalis, 61 albella, Cemiostoma, 54 albella, Eurynome, 54, 74 albella, Harpalyce, 55 albicapitella, Bucculatrix, 54 albilorella, Filatima, 66 albimarginella, Gelechia, 61 albiplagiatella, Myelois, 71 albisparsella, Gelechia, 63, 64 albistrigella, Ethmia, 55 albistrigella, Gelechia, 61 albocapitella, Laverna, 60 allutana, Grapholitha, 68 alniella, Lyonetia, 54 altissimella, Argyresthia, 67 ambrosiaeella, Lithocolletis, 75 ambrosiaefoliella, Bucculatrix, 54 amorphaeella, Gelechia, 61 ampelopsiella, Phyllocnistis, 75 ampelopsifoliella, Antispila, 50 amphicarpeaeana, Microaethia, 75 amphicarpeaeella, Lithocolletis, 74 amphicarpeaefoliella, Leucanthiza, 75 amphorana, Semasia, 68 ampla, Ploiophora, 56 anarsiella, Gelechia, 62 angleseana, Enarmonia, 68 angustipennella, Pigritia, 56 annectella, Holcocera zelleriella, 56 annulipes, Holcocera crescentella, 56 anticostalis, Botys, 71 apachella, Amydria, 50 apicialbella, Nepticula, 48, 75 apicipunctella, Hyponomeuta, 55 apicisignatella, Tinea, 50 apicistrigella, Lyonetia, 54 apicistrigella, Parasia, 62 approximated, Scardia, 50 approximella, Peoria, 73 argenteomaculatus, Hepiolus, 48 argentinotella, Semele, 50 argillacea, Depressaria, 55 argillaceellus, Crambus, 71 argutanus, Episimus, 68 argyreella, Pseudopigritia, 57 Argyresthiidae, 67 argyrosplendella, Calosima, 57 arizonella, Amydria, 50 arizonella, Pigritia, 57 arizoniella, Holcocera, 61 arnicella, Depressaria, 55 asilipennis, Sesia, 67 atrupictella, Eucordylea, 62 attributella, Taygete, 55 aufugella, Blastobasis, 57 aureopurpura, Aegeria, 67 aureovireus, Incurvaria, 50 auricristatella, Pitys, 50 auricyanea, Micropteryx, 48 auristrigella. Tinea, 50 auropulvella, Tinea, 51 aurosuffusella, Tinea, 51 austerella, Argyresthia, 67 badiiella, Tischeria, 49 basifasciella, Gelechia, 62 basilarella, Pigritia, 57 basipallidella, Hococera dives, 57 basistrigella, Gelechia, 62 bassiformis, Carmenta, 67 behrensella, Tinea, 51 Belanger, F.H., 47 belangerella, Argvresthia, 67 Belfrage, G.W., 47 bella, Adela, 50 bella, Aetole, 67 betulella, Acrobasis, 71 bicostomaculella, Gelechia, 55 bicristatella, Elachista, 60 bicristatella, Gelechia, 61 bifasciella, Gelechia, 62 bifasciella, Homosetia, 53 bifida, Euclea, 71 bifidella, Nealyda, 62 biguttata, Limacodes, 71 bimaculella, Depressaria, 62 bimaculella, Fascista, 66 bimaculella, Gelechia, 75 bimaculella, Nothris, 62 bimaculella, Tinea, 51, 74 biminimaculella, Gelechia, 62 bipunctella, Aetia, 61 bipunctella, Eucosma, 70 bipunctella, Progona, 51 biscolorella, Agnippe, 62, 75 tin Museum of Comparative Zoology, Vol. 152, No. 2 bistrigella, Phylloporia. 50 Blastobasidae, 56 bobana, Eucosma, 68 bolanderana, Paedisca, 68 Boll. J.. 47 borealis, Pyrausta. 71 boreasella, Holcocera, 57 boreasella, Oecophora, 55 bosqueella, Oecophora, 62 bosquella, Nepticula, 48 Boston Society of Natural History, 46 brevipennella, Amydria, 51 brevivittella, Mompha, 60 busckiella, Holcocera, 57 busckiella, Paraplesia, 51 caeculalis, Perispasta, 71 caemetariiella, Tinea, 74 caeruleana, Grapholita, 70 caeruleella, Adela, 50 californicalis, Eromene, 71 canadensisella, Buceulatrix, 54 canariella, Cerostoma. 66 canariella, Dryope, 57 canopulvella, Gelechia, 62 canusella, Harpalyce, 55 capitealbella, Buceulatrix, 54 carbonella, Abacobia, 51 cariosella, Epilegis, 51 carpenterellus, Crambus, 72 caryaeloliella, Coleophora, 75 castaneaefoliella, Nepticula, 45, 48 castella, Euceratia, 66 caudata, Aegeria, 67 celastrusella, Zelleria, 67 celtisella, Lithocolletis, 75 cephalonthiella, Laverna, 60, 75 cercerisella, Depressaria, 62 cercerisella, Faseista, 64 cervinidactylus, Pterophorus, 73 chalcofrontella, Holcocera, 59 chah l>i'is. Adela, 50 ( chambers, V.T., 46 chrysocomella, Isocorypha, 51 chrysurella, Breckenridgia, 55 ciliaefuscella. Nepticula, 48 cilialineella, Gelechia, 62 cincinnatiella, Lithocolletis, 75 i meraceus, Oidaematophorus, 73 cinereum, Palaeophobetron, 71 circumscriptella, Laverna, 60 citrinipennella, Tischeria, 49 clemensella ( lelechia, 55 clemensella, Lithocolletis, 75 clemensella, Nepticula. is clemensella Philonome, 54 75 clemensella, Tischeria. 49 clemensella, \\ lesthia, 51. 7 1 'i\ lidae, 68 cockerellana, Tortrix, 68 ( loleophoridae, 46 collinusella, ( lelechia <->-2 coloradella, Amydria, 51 coloradella, Laverna, 60 coloradensis, Prodoxus, 50 comptoniella, Aerobasis, 72 concinnusella, Gelechia, 62 concolor, Tischeria, 49 concolorella, Elachista, 61 concolorella, Eriphia, 61 confectella, Hypatima, 57 confluentella, Holcocera, 57 confusella, Amydria, 51 confusella, Pigritia, 57 consobrinella, Nephopteryx, 72 consonella, Gelechia, 62 constrictana, Paedisca, 68 constrictella, Theisoa, 62 continuella, Chionodes, 66 coryliella, Hyale, 55 corylisella, Lithocolletis, 75 Cosmopterigidae, 61 Cossidae, 68 costarufoella, Gelechia, 62 costotristgella, Tinea, 51 crassivenella, Enaemia, 67 crepera, Cossus, 68 crescentella, Amydria, 51 crescentella, Holcocera, 57 crescentifasciella, Gelechia, 62 cressonella, Cryptolechia, 55 crispata, Lagoa, 71 cristatella, Gelechia, 62, 75 cristatella, Semele, 51, 74 cristifasciella, Gelechia, 62 croceoverticella, Tinea, 51 cruciferella, Paraneura, 51 cryptolechiella, Psilocorsis, 55 cucurbitae, Aegeria, 67 cuprina, Aglossa, 72 curvilineatella, Lithocolletis, 54 curviliniella, Homostinea, 51 curvistrigella, Amydria, 51, 53 curvistrigella, Telphusa, 62 Davis, D.B., 50 decemmaculella, Taygete, 55 decimerella, Lipographis, 72 dehnitella, Laverna, 60 deletella, Argyresthia, 67 delphinii, Euclea, 71 dentiferella, Cerostoma, 67 denudatum, Trochilium, 67 depressostrigella, Gelechia, 62 desertana, Paedisca, 68 determinatella, Oecophora, 61 dianella, Calosima, 57 Dietz, W.G., 47 dietziana, Epinotia, 68 dietziella, Incurvaria, 50 difficilisella, Evagora, 55 dimidiata, Bhodoneura, 73 directana, Sparganothis, 70 disconotella, Gelechia, 62 MCZ Microlepidoptera Types • Miller and Hodges 83 discoocellella, Chionodes, 66 discoocellella, Gelechia, 62 discopunctella, Dryope, 57 discostrigella, Anesychia, 55 dives, Holcocera, 57 dodana, Eucosma, 68 dodecana, Paedisca, 68 dorsivittella, Gelechia, 62 dorsomaculella, Pseudopigritia, 57 dubitella, Depressaria, 62 duodecemlineella, Gracilaria, 75 dyarella, Amydria, 51 dyarella, Amydria, 52 eburnea, Scythris, 61 eclipsana, Grapholitha, 68 edmandsii, Nephopteryx, 72 ednana, Phalonia, 68 Edwards, H., 47 effrentella, Amydria, 51 ehrhornella, Paraneura, 51 Elachistidae, 56 electella, Anerastia, 72 elegantella, Gelechia, 62 elyella, Holcocera, 57 emancipatum, Gnorimoschema, 63 ephippiatus, Limacodes, 71 equitella, Pseudopigritia, 57 Eriocraniidae, 48 erransella, Perimede, 61, 75 estriatella, Holcocera, 57 eunitariaeella. Tinea, 52 eupatoriella, Ypsolophus, 62 eupatoriiella, Depressaria, 55, 75 exasperatana, Tortrix, 69 exoptatella, Glyphipteryx, 66 fagigemmaeana, Exartema, 69 faginella, Hagno, 55 falciferella, Cerostoma, 67 falsarius, Acoloithus, 71 fasciella, Gracilaria, 75 fasciella, Pitys, 52 fenestrella, Pempelia, 71 fenyesella, Dryope, 57 feriella, Tacoma, 72 Fernald, C.H., 47 fernaldella, Depressaria, 55 fernaldella, Setiostoma, 55 ferruginea, Euclea, 71 feudalis, Botis, 72 fidella, Ploiophora, 57 floridella, Progona, 52 floridella, Valentinia, 58 fluxella, Blastobasis, 58 fractiliniella, Apotomia, 52 fragariana, Lozotaenia, 69 fragmentella, Gelechia, 63 fraternella, Pseudopigritia, 58 Frey, H., 47 frigidana, Penthina, 69 frigidella, Eudorea, 72 frigidella, Oecophora, 52 frustella, Cerostoma, 67 frustrana, Retinia, 69 fullonella, Anacampsis, 65 fulminalis, Melissoblaptes, 72 fulminalis, Paralispa, 48 fulvifrontana, Penthina, 69 fulvipes, Aegeria, 67 fulvisuffusella, Tinea, 52 fumerella, Holcocera chalcofrontella, 58 funebra, Holcocera, 58 fusca, Pyla, 72 fuscella, Niditinea, 52 fuscocostella, Lithocolletis, 75 fuscocristatella, Naera, 63 fuscocristatella, Pitys, 52 fuscofasciella, Euplocamus, 52 fuscoluteella, Depressaria, 63 fuscomaculella, Gelechia, 63 fuseomaculella, Tinea, 52 fuscomarginella, Tischeria, 49 fuscoochrella, Gelechia, 63, 75 fuscopalidella, Sinoe, 63 fuscopulvella, Agnippe, 63 fuscopulvella, Gelechia, 63 fuscopurpurella, Blastobasis plummerella, 58 fuscoscapulella, Acanthocnemes, 54 fuscostrigella, Plyhymno, 63 fuscosuffusella, Dryope, 58 fuscotaeniaella, Gelechia, 63 fuscotibiella, Stigmella, 48 Gelechiidae, 61 geminata, Cyrtosia, 71 geniculatella, Tinea, 52 georgiella, Dichomeris, 62 gigantella, Blastobasis, 58 gilviscopella, Gelechia, 63 glacialis, Botys, 72 glandiferella, Gelechia, 63 glandulella, Valentinia, 59 glaucofuscana, Conchylis, 69 gleditschiaeella, Laverna, 61, 75 glycyrhizaeella, gelechia, 63 Glyphipterigidae, 66 gomonana, Eucosma, 69 gracilis, Heliozela, 50 gracilis, Korescheltellus, 48 Gracillariidae, 46 grandis, Lioptilus, 73 grandis, Oidaematophorus, 48 grandisella, Nepticula, 49 granella, Nemapogon, 51, 52, 53 griseella, Tinea, 52 grisella, Dryope, 58 grisseella, Laverna, 60 hagenella, Anesychia, 55 haimbachiana, Epinotia, 69 hamellus, Crambus, 72 Harris, T.W., 47 heidemannella, Epigritia, 58 Heliodinidae, 67 heliopsisella, Tischeria, 49, 75 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 Heliozelidae, 50 henshawiella, Scrobipalpula, 64 Hepialidae, 18 hospitella, Ephestia, 72 Hulst, G.D., 47 hulstella, Blastobasis, 58 humilis, Antaeotricha, 55, 56 hybromella, Oenoe, 52 iceryaeella, Blastobasis, 58 ignobilisella, Laverna, 60 illibella, Holcocera, 58 imamoenella, Setomorpha, 52 immaculata, Lagoa, 71 immaculatella, Butalis, 61 impigritella, Diploschizia. 66 impositella, Scythris, 61 inclusa, Holcocera, 58 Incurvariidae, 50 inimicella, Galleria, 69 innocuella, Gelechia, 63 inornatella, Eulyonetia, 54 inquinatalis, Udea, 72 insulatella, Holcocera, 58 integra, Scoptonoma, 72 intermediella, Gelechia, 63 intermistana, Olethreutes, 70 interpunctella, Holcocera, 58 interrupta, Scoptonoma, 72 interstitiella, Tinea, 52 inversella, Epicorthylis, 63 irrorella, Monopis, 52 juglandifoliella, Nepticula, 49 juncidella, Dichomeris, 62 Kearfott, W.D., 47 labradoriensis, Hepialus, 48 lacteodactylus, Pterophorus, 73 laticapitella, Pigritia, 57, 58 latitasciatella, Xephopteryx, 72 latifasciella, Gelechia, 63 latifasciella, Nepticula, 49 latipenella, Tischeria, 49 lavernella, Gelechia, 63 la> t ti Megalopyge, 71 lentiflualis, Homophysa, 72 Iconincllu Pempelia, 7.2 leuconota, Evippe, 65 leuconota, Gelechia, 63 leucophaleratana, Pandemis, 69 ligulella, Dichomeris, 65 Limacodidae, 7 1 lithosina, Cryptolechia, 55 litigiosella, Bucculatrix, 54 liturosella, Gelechia, 63 livorella, Blastobasis, 58 longalis, Dysodia, 73 longifasciella, Telphusa, 62, 64 longimaculella, Hyponomeuta, 55, 75 longipalpana S) llonoma, 69 luteella, Bucculatrix, 54. 75 luteella, Eur) nome, 5 I luteopulvella Dryope, 58 luteostrigella, Polyhymno, 63 Lyonetiidae, 54 maculata, Thyris, 73 maculatella, Homosetia, 52 maculimarginella, Gelechia, 63 maculipes, Zenodoxus, 67 maculosella, Nepticula, 49 magnatella, Laverna, 74 magnella, Bucculatrix, 54 majorella, Setomorpha, 52 mandarinella, Tinea, 51 marcidana, Phoxopteris, 69 marginatum, Trochilium, 67 marginimaculella, Homosetia, 52 marginistrigella, Tinea, 52 margoriella, Amydria, 52 marmorella, Gelechia, 63 maximella, Nepticula, 49 mediofasciella, Pigritia, 58 mediofuscella, Chionodes, 63 mediostriatella, Isocorypha, 50 Megalopygidae, 71 melanocarpum, Helcystogramma, 66 melanogrammos, Tetralopha, 72 melanostriatella, Holcocera, 58 merrickanum, Exartema, 69 mesogramma, Ochrothyris, 73 messelinella, Holcocera, 58, 60 metallicana, Olethreutes, 69 metalliferalis, Calaclysta, 72 minimaculella, Gelechia, 63 minimella, Gelechia, 63 minnicella, Dryope, 59 minorella, Holcocera chalcofrontella, 59 minuta, Acleris, 70 minutipulvella, Tinea, 52 miriamella, Leucomele, 52 mirusella, Anesychia, 55 miscana, Eucosma, 69 misceeolorella, Laverna, 61 miscecristatella, Pitys, 52 misceella, Tinea, 52 molybdanella, Tinea, 52 Momphidae, 60 monitor, Euclea, 71 monodactyla, Emmelina, 74 montella, Argyresthia, 67 montisella, Cosmopteryx, 61 montisella, Glyphipteryx, 66 monumentella, Gelechia, 63 multifasciella, Theisoa, 64 multimaculella, Gelechia, 52 multipunctella, Anesychia, 56 multistriatella, Tinea, 53 murina, Penthina, 69 Murtfeldt, M., 47 murtfeldtella, Dryope, 59, 75 murtfeldtella, Laverna, 60 mustelinalis, Botys, 72 nebeculosa, Cryptolechia, 56 nebraskae, Euhagena, 68 MCZ Microlepidoptera Types • Miller and Hodges 85 nebulosana, Grapholitha, 69 nepotella, Epichaeta, 53 Nepticulidae, 48 nigratomella, Battaristis, 62 nigrella, Gelechia, 64 nigrilineella, Eriphia, 61 nigriverticella, Stigmella, 49 nigroatomella, Tinea, 53 niveosana, Sciaphila, 69 nonstrigella, Dasycera, 64 novi-mundi, Depressaria, 56 nubiferella, Depressaria, 56 nubilella, Blastobasis, 59 numerosana, Paedisca, 69 obliquella, Amydria, 53 obliquifasciella, Gelechia, 64 obliquistrigella, Anarsia, 64 obrutella, Eetoedemia, 48 obscurella, Homosetia, 53 obscurella, Pigritia, 59 obseuroiasciella, Buceulatrix, 54 obseuromaeulella, Cryptoleehia, 56 obscurosuffusella, Filatima, 62 obscurosuffusella, Gelechia, 64 obscurusella, Chionodes, 63 obscurusella, Depressaria, 64 obscurusella, Laverna, 60 occidentalis, Scopula, 72 occidentella, Amydria onagella, 53 occidentella, Dryope, 59 occidentella, Gelechia, 64 occidentella, Tinea, 53, 54 ocellella, Gelechia, 64 ochrella, Coleophora, 75 ochrella, Pluteloptera, 67 ochreocostella, Gelechia, 64 ochreostrigella, Gelechia, 64 ochreosuffusella, Filatima, 62 ochrifrontella, Ephestia, 72 ochrocephala, Holcocera, 59 ochromediana, Olethreutes, 69 ochromella, Epigritia, 59 octonalis, Orobena, 72 Oecophoridae, 55 oenotheraesemenella, Laverna, 60 oenotheriella, Laverna, 74 olivaceana, Eccopsis, 69 olympiadella, Gelechia, 64 onagella, Amydria, 53 operosella, Setomorpha, 53 ophionalis, Microtheoris, 73 ophrionella, Tinea, 53 orichalcella, Periploca, 61 orleansella, Tinea, 53 ornatella, Litliocolletis, 75 ornatella, Pigritia, 59 ornatihmbriella, Filatima, 61, 66 osmundana, Olethreutes, 69 osmundana, Penthina, 69 ostryaeella, Aeaea, 61 ovalis, Pempelia, 72 ovalis, Telethusia, 72 Packard, AS., Jr., 47 packardella, Buceulatrix, 54 packardella, Gracilaria, 75 packardi, Grapholitha, 69 paleaceus, Leioptilus, 73 pallidactyla, Platyptilia, 73 pallidastrigella, Cleodora, 64 pallidella, Cleodora, 64 pallidochrella, Depressaria, 64 pallidochrella, Helice, 64, 75 pallidotinctella, Epigritia, 59 palpiannulella, Gelechia, 64 palpilineella, Gelechia, 64 pandurella, Amydria, 53 paradoxella, Apreta, 53 paradoxica, Hyponomeuta, 50 parvipulvella, Elachista, 56, 75 Peabody Academy, 46 peculiana, Teras, 69 pedmontella, Argyresthia, 67 pedmontella, Gelechia, 64 pennsylvanica, Pseudochelaria, 64 perfluana, Grapholitha, 70 pergracilidactylus, Pterophorus, 74 perrubralis, Botys, 73 persicana, Clepsis, 69 physaliella, Gelechia, 64 pinatubana, Eulia, 70 piperatella, Cryptoleehia, 56 piperatella, Durrantia, 55 plagiatella, Holcocera, 59 platanana, Ancylis, 69 platanella, Cirrha, 64 platanella, Eetoedemia, 49 plausipennella, Butalis, 61 plumbicostalis, Botys, 73 plumbolineana, Epinotia, 70 plummerella, Blastobasis, 58, 59 plutella, Gelechia, 64 plutella, Neda, 65 plutella, Phaetusa, 65 Plutellidae, 66 polistiformis, Aegeria, 68 polita, Calantica, 67 pomifoliella, Buceulatrix, 54 pomivorella, Micropteryx, 49 popeanella, Acrolophus, 53 posticella, Depressaria, 56 praeangusta, Batrachedra, 74 primulana, Paedisca, 70 profundalis, Botys, 73 prosperana, Enarmonia, 70 prudens, Trypanisma, 65 prunifoliella, Evippe, 65 prunifoliella, Stigmella, 49 pruniramiella, Xylesthia, 51 prunivorella, Ornix, 75 pseudacaciella, Depressaria, 65 Pterophoridae, 73 ptyehogrammos, Teras hastiana, 70 Bulletin Museum of Comparative Zoology, Vol. 152, No. 2 pudibundella, Aristotelia, 65 pulchella, Euresia, 59 pulla, Trosia, 71 pullusella, Aristotelia, 63 pulvella, Tischeria, 49 pumilio, Marasmarcha, 48 pumilio, Mimesoptilus, 74 punctidiscella, Dichomeris, 66 punctiferella, Holcocera, 60 pupula, Lactura, 67 purinosella, Tischeria, 49 purpurella, Pigritia, 59 purpuriella, Chrysopeleia, 61 purpuriella, Chrysopelia, 75 purpuriella, Gracilaria, 75 pusilla, Holcocera, 59 Pyralidae, 71 pyri, Aegeria, 68 quadricustatella, Aeaea, 61 quadrilineella, Cosmopterix, 61 quadrimaculella, Anacampsis, 48 quadrimaculella, Gelechia, 65 quadristrigella, Argyresthia, 67 quaintancella, Valentinia, 59 quercicolella, Argyresthia, 67 querciella, Depressaria, 65 querciella, Gelechia, 75 querciella, Ypsolophus, 65 quercifoliana, Argyrotaenia, 70 quercinigracella, Gelechia, 65 quercinigracella, Pseudotelphusa, 63 quercipominella, Ypsolophus, 65 quercipulchella, Nepticula, 49 quercitella, Tischeria, 50 quercivorella, Coleotechnites, 63 quercivorella, Gelechia, 65 quercivorella, Tischeria, 49, 75 quericastanella, Nepticula, 49 quinqueannulella, Gelechia, 65 quinqueferella, Glyphipteryx, 56 quinquepunctellus, Prodoxus, 50 quintana, Paedisca, 70 quisquiliella, Blastobasis, 59 raracana, Thiodia, 70 reductella, Holcocera I'unebra, 59 reedella, Ypsolophus, 65 reniculalis, Homophysa, 73 resplendensella, Nepticula, 49 rheumapterella, Incurvaria, 50 rhoifructella, \nacampsis, 62, 64, 65 ribesella, Gelechia, 65 rileyella, Depressai ia, 65 n l<\ i, Tinea. 5 I robiniae, Prionoxystus, 68 robiniella, Sinoe, 63 robinsonana, Eucosma, 70 roburella, Tinea, 53 rnseatclla. \ephopter\ \. 73 roseticola, Tischeria, 50 rubensella, Gelechia, 65, 75 rubidella, Aristotelia. 65 rubrifasciella, Acrobasis, 73 ruderella, Setomorpha, 53 rufopunctella, Holcocera, 59 rufusella, Gelechia, 65 rutella, Setomorpha, 51, 52, 53, 54, saginella, Stigmella, 49 sagitella, Blastobasis, 59 salicifoliella, Gracilaria, 74 salicifoliella, Lithocolletis, 74 sanborni, Harrisina, 71 saphirinella, Gelechia, 65 Saunders, W., 47 saundersella, Gelechia, 65 scardina, Anaphora, 53 sciaphilella, Blastobasis, 59 sciaphilella, Holcocera, 60 scintillana, Pelochrista, 68 scitula, Aegeria, 68 scobiella, Stylopalpia, 72 scutellariaeela, Gelechia, 65 Scythrididae, 61 securella, Euceratia, 67 sella, Gelechia, 65 semicostatus, Mimeseoptilus, 74 semilugens, Ethmia, 56 semirubralis, Botys, 73 septemstrigella, Tinea, 53 septentrionana, Olethreutes, 69 sepulchrella, Tryptodema, 53 seriatana, Conchylis, 70 serotinaeella, Nepticula, 49 serratipalpella, Gelechia, 65 serratissimalis, Crocidophora, 73 serrativittella, Dichomeris, 64 serrativittella, Gelechia, 65 servulella, Hybroma, 51 sescuplana, Tortrix, 70 Sesiidae, 67 sesquialteralis, Botis, 73 setosella, Dichomeris, 62 sexnotella, Gelechia, 61 sexstrigella, Polyhymno, 66, 75 shaleriella, Oecophora, 56 sigmoidella, Setomorpha, 53 similiella, Isophrictis, 66 simplicella, Blastobasis plummerella, 59 simulella, Holcocera, 59 simulella, Paraneura, 53 sincera, Oectoperia, 73 skinnerella, Mea, 52, 53 skinnerella, Progona, 53 solaniiella, Gelechia, 66 spaldingana, Eucosma, 70 speculella, Lyonetia, 54 spiculana, Grapholitha, 70 spoliatella, Holcocera messelinella, 60 spoliatella, Pigritia, 60 spretella, Holcocera, 60 staintonella, Bucculatrix, 55 staintonella, Elachista, 56 stimulea, Acharia, 71 MCZ Microlepidoptera Types • Miller and Hodges 87 straminiella, Tinea, 53 straminiella, Ypsolophus, 66 subdivisalis, Desmia, 73 subfasciella, Araeolepia, 67 subnivana, Acleris, 69 subolivalis, Botys, 73 subsenella, Hypatima punctiferella, 60 subsequalis, Pyrausta, 71 sulphureodactylus, Pterophorus, 74 syringae, Aegeria, 68 tabaniformis, Paranthrene, 68 tartarella, Holcocera, 60 tautana, Sereda, 70 tenebrella, Dryope, 60 ternariella, Gelechia, 66 terracoctana, Paedisca, 70 tessellana, Penthina, 70 tessellatella, Lindera, 51, 53 testacea, Tortricidia, 71 testulana, Cenopis, 70 tetradella, Anerastia, 73 texanella, Anaphora, 53 texanella, Anesychia, 66 texanella, Elachista, 48 texanella, Hyponomeuta, 56 theseusalis, Herpetogramma, 72 thoracealbella, Aroga, 63 thoraeealbella, Gelechia, 66 thoracealbella, Nepticula, 49 thoracefasciella, Gelechia, 56 thoracenigraeella, Gelechia, 56 thoracestrigella, Tinea, 53 thuiella, Bucculatrix, 67 Thyrididae, 73 tibiale, Troehilium, 68 tiliaeella, Lithocolletis, 75 tinctoriella, Tischeria, 50 tineana, Ancylis, 69 Tineidae, 50 Tischeriidae, 49 tortieiformella, Menesta, 55 tortricella, Harpalyce, 56 Tortricidae, 68 transversestrigella, Semiota, 53 trialbamaculella, Gelechia, 66 triangularisella, Holcocera, 60 tricincta, Aegeria, 68 tricristatella, Leucophryne, 60 trifasciella, Bucculatrix, 54 trifasciella, Gelechia, 66 trifurcella, Anesychia, 56 trifurculana, Tortrix, 70 trilineella, Gelechia, 66 trimaculella, Anarsia, 66 trimaculella, Gelechia, 66 trimaculella, Isophrictis, 48 triocellella, Gelechia, 66 tripartitana, Paedisca, 70 tristella, Pigritia, 60 tristriata, Sparganothis, 70 tritaeniaella, Lithocolletis, 75 tuscanella, Tinea, 54 ulmella, Lithocolletis, 75 umbraticostella, Depressaria, 56 unctulella, Gelechia, 66 undulatella, Argyresthia, 67, 75 unifascialis, Botys, 73 unifascialis, Pyrausta, 48, 73 unifasciella, Laverna, 60 unifasciella, Nepticula, 49 unimaeulella, Ithome, 61 unipunctella, Antaeotricha, 55, 56 unistriatellus, Crambus, 73 unomaculella, Perimede, 75 unomaculella, Tinea, 54 ustella, Ypsolopha, 67 vagatioella, Coleotechnites, 62 vanella, Plutella, 67 variatella, Nemapogon, 50, 52 variella, Gelechia, 66, 75 variolana, Teras, 70 ventrella, Dichomeris, 65 versutella, Gelechia, 66 vertumnana, Paedisca, 70 verutana, Bactra lanceolana, 70 vestaliana, Grapholitha, 70 vestaliella, Holcocera, 60 vestalis, Cryptolechia, 56 vibicalis, Botis, 73 vicinella, Tinea, 54 violaceofusca, Gelechia, 66 virescana, Clepsis, 70 visaliella, Cyane, 54 viteana, Endopiza, 70 viticordifoliella, Antispila, 50 vitifoliella, Phyllocnistis, 75 vitivorana, Penthina, 70 Walsingham, Lord, 47 walsinghami, Pseudochelaria, 66 worthingtoniana, Paedisca, 70 xanthobasis, Setiostoma, 56 xanthostictella, Tinea, 54 Yponomeutidae, 67 yuccaecolella, Blastobasis, 60 yuccasella, Tegeticula, 50 yumaella, Plutella, 54 y-inversa, Limacodes, 71 zana, Enarmonia, 70 Zeller, PC., 47 zelleriella, Holcocera, 60 zelleriella, Hyponomeuta, 56 zelleriella, Tischeria, 49 zophodactyla, Stenoptilia, 74 Zygaenidae, 71 (US ISSN 0027-4100) Bui Latin of the Museum of Comparative Zoology MCZ LIBRARY OCT 2 4 1990 HARVARD Early Mis®te$f5$aff Blastoids from Western Montana JAMES SPRINKLE and RAYMOND C. GUTSCHICK HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS, U.S.A. VOLUME 152, NUMBER 3 25 SEPTEMBER 1990 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952- BULLETIN 1863- Memoirs 1865-1938 Johnsonia, Department of Mollusks, 1941- Occasional Papers on Mollusks, 1945- SPECIAL PUBLICATIONS. 1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Tere- dinidae (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4. Eaton, R. J., 1974. A Flora of Concord. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 725 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprint. Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. Creighton, W. S., 1950. The Ants of North America. Reprint. Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First International Symposium on Natural Mammalian Hibernation. Ornithological Gazetteers of the Neotropics (1975-). Peters' Check-list of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1947. (Complete sets only.) Proceedings of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publications Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 1 990. EARLY MISSISSIPPIAN BLASTOIDS FROM WESTERN MONTANA JAMES SPRINKLE1 AND RAYMOND C. GUTSCHICK2 ABSTRACT. Several faunas of Early Mississippian blastoids occur in the Lodgepole and Allan Mountain Limestones of western Montana. More than 1,400 complete specimens representing at least nine genera and 16 species have been collected from three dif- ferent zones, making this one of the largest blastoid collections known from western North America. The largest and most diverse blastoid fauna occurs just above the base of the Lodgepole (lower Paine Mem- ber) and Allan Mountain Limestones at 31 localities and consists of nearly 1,200 specimens belonging to four blastoid genera (Tanaoblastus, Strongyloblas- tus, Orophocrinus, and Metablastus). A second fauna occurs in the middle Lodgepole Limestone (upper Paine Member) at four localities where about 195 specimens and five blastoid genera occur (Koryschis- ma, n. gen., Montanablastus, n. gen., Strongyloblas- tus, Cryptoblastus?, and Hadroblastus). The highest fauna occurs near the top of the Lodgepole Limestone (upper Woodhurst Member) at three localities in the Bridger Range and is represented by 19 specimens and three blastoid genera (Cryptoblastus?, Oropho- crinus, and Phaenoschisma). Most of the blastoids in these faunas are fairly well silicified, and, when extracted with heated acetic acid, a few show excellent preservation of plate ornament and ambulacral structures. One blastoid occurrence in the middle Lodgepole has calcitic specimens with complete brachioles and attached stem segments that lack distal attachment structures. The Lodgepole blastoid faunas appear to be middle Kinderhookian to early Osagean (early to middle Tournaisian) in age, and are most similar to other Early Mississippian (or earliest Carboniferous) blastoid faunas in Missouri, Alberta, New Mexico, and Belgium. The diverse low- er Lodgepole fauna is dominated by a small globular spiraculate (Tanaoblastus) at nearly all sections, whereas other pyramidal, elongate, and club-shaped spiraculate or fissiculate blastoids are much less com- mon. Most of these blastoids were apparently at- tached, medium-level, suspension feeders living on a lime mud bottom in a carbonate ramp setting near 1 Department of Geological Sciences, University of Texas, Austin, Texas 78713-7909. 2 Department of Earth Sciences, University of Notre Dame, Notre Dame, Indiana 46556-1020 (Present ad- dress: 2901 Leonard, Medford, Oregon 97504). or well below normal wave base. At several localities, members of the lower and middle Lodgepole blastoid faunas are found adjacent to or just below Waulsor- tian-type bioherms. New taxa include the fissiculates Koryschisma ele- gans, n. gen., n. sp., and Orophocrinus macurdai, n. sp., and the spiraculates Metablastus milliganensis, n. sp., Strongyloblastus breimeri, n. sp., S. laudoni, n. sp., Montanablastus baldyensis, n. gen., n. sp., and Tanaoblastus allanensis, n. sp. INTRODUCTION Blastoids are usually considered a rela- tively rare element in the Early Mississip- pian faunas of the northern Rocky Moun- tain region in the western United States. Only four blastoid species from the Early Mississippian of this region have been de- scribed in the 120 years between 1865 and 1985. However, several authors have re- ported the presence of unidentified blas- toids in faunal lists during this period. Be- tween 1963 and 1968, we made an extensive collection of blastoids from the Lodgepole and Allan Mountain Lime- stones of Early Mississippian age in west- ern Montana and adjacent states. This new material and a restudy of previously de- scribed specimens form the basis for this paper. The present authors independently dis- covered blastoids in the Early Mississip- pian of western Montana during the sum- mers of 1962 and 1963. During the following three summers (1964-66), we re- turned to Montana to work together on the biostratigraphy and paleontology of the Sappington Member of the Three Forks Formation and the Lodgepole Limestone under NSF-sponsored grants (see Ac- knowledgments; Sprinkle, 1965; Sprinkle Bull. Mus. Comp. Zool., 152(3): 89-166, September, 1990 89 90 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 Table 1. List of collecting localities where we collected blastoids from the Lodgepole and Allw Mountain Limestones in western Montana and southeastern Idaho. Localities for borrowed USGS specimens (Sola w Creek, Gallatin Range; Old Baldy, Gravelly Range; and Brazer Locality Code Range Land-grid location Antelope Valley AV SW, NE, NE, sec. 2, T1S, R2W Ant Park AP Little Belt Mtns. NW, SW, sec. 35, T12N, R9E Bacon Rind Creek BA Gallatin Range SE, SE, sec. 22, T10S, R5E Bakly Mountain BY Bridger Range NW, SE, sec. 11, T1S, R6E Bandbox Mountain BD Little Belt Mtns. NE, NW, sec. 20, T14N, R10E Bridger Mountain BG Bridger Range SW, NE, sec. 11, T1S, R6E Brow nback Gulch BB Tobacco Root Mtns. NE, NW, SE, sec. 20, T1S, R3W Cowboy Canyon CB Madison Range SE, NW, sec. 27, T4S, R2E ( tow ii Mountain CR Lewis & Clark Mtns. C, SW, sec. 28, T19N, R9W Dry Hollow DH SW, NE, sec. 3, TIN, R1W Dudley Creek DC Madison Range NW, SE, sec. 32, T6S, R4E Ellis Mountain (Ellis Peak) EM Gallatin Range NW, SE, sec. 14, T3S, R6E Gallop Creek GP Bridger Range SE, SW, sec. 11, T3N, R5E Grendah Mountain GH Little Belt Mtns. SE, NE, NW, sec. 31, T13N, R9E Jordan Creek JC Madison Range W/2, sec. 23, T5S, R1E Little Antelope Creek LA Tobacco Root Mtns. S'/z, sec. 27, T1S, R2W London Hills LH NV2, NW, sec. 4, T1S, R2W Milligan Canyon MC NE, SW, sec. 36, T2N, R1W Milligan Canyon East MC(E) NE, SW, sec. 31, T2N, R1E North Frazier Lake FR Bridger Bange NW, SE, sec. 9, T2N, R6E North Sawtooth Mountain NS Lewis & Clark Mtns. SW, NE, sec. 13, T21N, R9W Northeast Baldy Mountain NB Bridger Range SE, SE, SE, sec. 2, T1S, R6E Pole Canyon PC Tobacco Root Mtns. SW, sec. 8, T1S, R3W Roy Gulch RG Horseshoe Hills NW, NW, sec. 28, T4N, R4E Sacagawea Peak SA Bridger Range SE, NW, sec. 27, T2N, R6E Saddle Peak SP Bridger Range EV2, NE, sec. 35, T1S, R6E Sand Creek sc SW, NE, sec. 4, T1S, R1W Sixteen Mile Creek SX Horseshoe Hills SE, SW, sec. 4, T4N, R3E South Boulder SB Tobacco Root Mtns. SE, SW, sec. 20, T1S, R2W Squaw Creek Ranger Station so Gallatin Range NW, NE, sec. 28, T4S, R4E Standard Creek ST Gravelly Range NE, SE, sec. 6, T11S, R1W Targhee Peak TG Henrys Lake Mtns. NE, SW, SW, sec. 9, T16N, R43E Timber Butte TB Gallatin Range NE, SE, sec. 11, T5S, R5E and Gutschick, 1967; Gutschick, McLane, and Rodriguez, 1976; Sprinkle and Gut- schick, 1983). This research resulted in the discovery that blastoids are relatively com- mon and diverse in the basal part of the Lodgepole and Allan Mountain Lime- stones over much of western Montana and that they are also present at two higher levels in the Lodgepole. Blastoids have now been found at 33 sections (Table 1) in southwestern, west-central, and north- western Montana and in southeastern Idaho (Text-Fig. 1), and are undoubtedly present at many other localities in western Montana and adjacent states. During these three summers of field work and shorter visits in 1967 and 1968, we collected more than 1,400 blastoid specimens, belonging to nine genera and 16 species. PREVIOUS STUDIES OF MADISON GROUP STRATIGRAPHY The Madison Group in western Mon- tana is made up of the thinner-bedded Lodgepole Limestone below (Text-Fig. 2) and the overlying more massive-bedded Mission Canyon Limestone. Together these two units represent 750 ft (229 m) to more than 2,000 ft (610 m) of Early and Middle Mississippian tropical-shelf carbonates. Lodgepole thicknesses range from about Mississippian Blastoids from Montana • Sprinkle and Gutschick 91 (continued) Canyon, northern Utah) are not included because we did not visit or collect BLASTOIDS THERE. ABBREVIATIONS FOR BLASTOID GENERA INCLUDE: C = CRYPTOBLASTUS?, H = HADRO- BLASTUS, K = KORYSCHISMA, M = METABLASTUS, Mo = MONTANABLASTUS, O = OROPHOCRINVS, P = Phaenoschisma?, S = Strongyloblastus, and T = Tasaoblastus. Topographic map County State Fauna Blastoid composition Jefferson Island Madison Mont. Lower 4 T, 1 M Sand Point Meagher Mont. Middle ? 1 S, 1 K? Tepee Creek Gallatin Mont. Lower 1 T Sedan Gallatin Mont. Lower, Upper 3 T; 1 O, 1 C Bandbox Mountain Judith Basin Mont. Lower, Middle 8 T; 27 K, 4 C Sedan Gallatin Mont. Lower 8T Waterloo Madison Mont. Lower 1 T Ancenney Madison Mont. Lower 14 T Choteau Lewis & Clark Mont. Lower 174 T Three Forks Jefferson Mont. Lower 21 O, 1 S, 1 T Spanish Peaks Gallatin Mont. Lower 1 T Mystic Lake Gallatin Mont. Lower 14 T Maudlow Gallatin Mont. Lower 10 T King's Hill Judith Basin Mont. Lower 1 T Ennis Madison Mont. Lower 10 T Harrison Madison Mont. Lower 25 T, 2 O Jefferson Island Madison Mont. Lower 220 T, 1 M Three Forks Jefferson Mont. Lower 17 S, 2 O, 1 T, 1 M Three Forks Broadwater Mont. Lower 19 S, 15 O, 5 T, 2 M Sedan Gallatin Mont. Lower 25 T Sawtooth Ridge Lewis & Clark Mont. Lower 2T Sedan Gallatin Mont. Lower, Middle 4 T, 2 M; 29 Mo, 10 S, 1 C Whitehall Madison Mont. Upper? 1 C Maudlow Gallatin Mont. Lower 2 T Sedan Gallatin Mont. Lower, Upper 2 T; 11 C, 1 K? Sedan Gallatin Mont. Lower, Upper 1 T, 1 S; 1 P Three Forks Gallatin Mont. Lower 1 O Toston Broadwater Mont. Lower 7 T Harrison Madison Mont. Lower 2 T, 2S Garnet Mountain Gallatin Mont. Lower 18 T Monument Ridge Madison Mont. Lower, Middle 437 T, 2 O, 1 S; 1 H Targhee Peak Fremont Idaho Lower 51 T, 1 S Garnet Mountain Gallatin Mont. Lower 8 T 500 ft (152 m) to more than 1,000 ft (305 m) (Gutschick, McLane, and Rodriguez, 1976, pp. 107-108, figs. 8-12, 8-13). These competent structurally-deformed rocks commonly form the backbone and crest of many mountain ranges (Text-Figs. 3-6). The enormous volume of carbonate rock and the physiographic obstacles make ac- cess, observation, and collection difficult for depositional and paleontological field studies. Nevertheless, many local and re- gional contributions have been published for Montana. Some useful studies of the Lodgepole Limestone which contains the blastoid fau- nas described here include the following: Sloss and Hamblin, 1942; Laudon and Sev- erson, 1953; Andrichuk, 1955; Roberts, 1966; Wilson, 1969; Sando, Mamet, and Dutro, 1969; Craig, 1972; Smith, 1972, 1977; Sando and Dutro, 1974; Sando, 1976; Rose, 1976; Gutschick, McLane, and Ro- driguez, 1976; Roberts, 1979; Gutschick, Sandberg, and Sando, 1980; and Sandberg and Gutschick, 1983, 1984. Studies of the Allan Mountain Lime- stone in the Sun River area include: Sloss and Laird, 1945; Mudge, Sando, and Du- tro, 1962; and Haines, 1977. Several oc- currences of Waulsortian-like bioherms are known from the lower and middle Lodge- pole Limestone and are discussed in 92 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 North Sawtooth Mountain \ ^ * T Augusta \ * V L \Crown Mountain ,«>^ ^ \ \ 1 . 5 . 1 III c K N A 0 10 20 30 40 50 0 10 20 30 40 50 Kilometers J? ■*° MISSISSIPPIAN LODGEPOLE LIMESTONE BLASTOID LOCALITIES, WESTERN MONTANA DeeperWater MONTANA '\ U \ \ / I s HELENA s s y MONTANA • fc'^^" /1r/V .' nt& e°U r>\° sx Sixteen Mile Creek RG, / / /u X L MC Boy Gulch GP L •Gallop Creek wuitcuii i MI|H9an Canyon L X MC(E) _ X 9i L London Hills LH .L Canyon "U Pole C .„,o„PCa . .v3CaLdEc..e,_ Brownback Gul?h l-*»L AMalop* Valley South Boulder LA Llt,'» *n««'0P« i North Frailer Lake L ?,Siciga»8i Peak U BRIDGER L SP RANGE (J NB aSaddle Peak £ - T. 1 N BgJne Baldy Mountain M By Brldger Mountain I Baldy T. 1 S Deeper Water "n,,,»« o Baldy u Or..* BOZEMA* Mountain L LIV>|NGST0N EM , Ellis • L Mountain Cowboy CB \ Canyon'*" 3quaw3«P T(J LJ.° „ graene,k,rL 'Timber Butt. O DILLON i ennia i«- ■ - Hangoi \ O Jordan V, Station 7. x glnla X / •s. \ U Upper blaaloid fauna M Middle blaatold fauna L-Lower blaatold fauna ■z- Standard Creek M L era** Ta City U> V L O Dudley Creek e°u o\e X^ •v°v / ^1 BAf— -- Bacon RlndlCreek I 4 5- WYOMING i\) ^^-J 113* CENTENNIAL RANGE IDAHO 112' Targhee West; J Peak i> v \\ 1 K I \i N \ B VSED PARTLY ON A FIELD CENSUS TAKEN FROM TALUS BLOCKS AT STANDARD CREEK IN 1966; MIDDLE FAUNA BASED PARTLY ON ACID RESIDUES FROM BANDBOX MOUNTAIN (IDENTIFIED BY FRANCIS ZlMMEB ) WD ANT P\Kk. WD A CENSUS OF SLAB SURFACES COLLECTED FROM NORTHEAST BALDY MOUNTAIN Fossil group Lower blastoid fauna Middle blastoid fauna Protozoa Foraminiferida Textulariina Fusulinina Porifera Coelenterata Anthozoa Rugosa Tabulata Br\ozoa ( sstoporata ( rj ptostomata I ■■ ■ i it-strata Hyperammina rockfordensis Pseudastrorhiza digitata P. 2 species Trepeilopsis glomospiroides Ammobaculites leptos Septglomospiranella sp. Septabrunsiina sp. Latiendothyra sp. Siliceous spicules Amplexus sp. Amplexizaphrentis sp. Ample xocarinia sp. Cyathaxonia tantilla Cleistopora placenta Metriophyllum deminutivum Neaxon? sp. Palaeacis sp. Aulupora sp. Cladochonus sp. Several Fenestellids Hyperammina rockfordensis-H. kentuckyensis transit. Rheophax calathus R. raymoorei Tolypammina sp. Chernyshinella sp. Paleospiroplectammina sp. Rectoseptaglomospiranella sp. Siliceous spicules Globular form with spicules Amplexus sp. Amplexizaphrentis sp. Amplexocarinia sp. Cyathaxonia tantilla Sychnoelasma subcrassum Stelechophyllum microstylum? Aulopora sp. Syringopora sp. Cystodictya sp. Fistulipora sp. Sulcoretepora? sp. Unidentified Cystodictyonid Nicklesopora sp. Rhombopora or Rhabdomeson sp. 6 genera of Fenestellids Hemitrypa sp. Penniretepora sp. Ptylopora sp. Unidentified Acanthocladid Septopora sp. Mississippian Blastoids from Montana • Sprinkle and Gutschick 111 (continued) in 1966 and 1984-85. Published sources for information on particular groups listed below include: gutschick (1964), gutschick, weiner, and young (1961), and mamet and skipp (1970), forams; sando (1983) and sando and bamber (1985), corals; mcklnney (personal communic- ation, 1987), middle fauna bryozoans; rodriguez and gutschick (1968, 1969), lower fauna bra- chiopods; Gordon (1986), lower fauna ammonoids; Rodriguez and Gutschick (1970), trace fossils; Laudon and Severson (1953), crinoids; Gutschick, Canis, and Brill (1967), holothurians; and Sandberg et al. (1978), conodonts. Fossil group Lower blastoid fauna Middle blastoid fauna Brachiopoda Inarticulata Articulata Orthida Strophomenida Rhynchonellida Spiriferida Terebratulida Mollusca Gastropoda Bivalvia Cephalopoda Nautiloidea Ammonoidea 'Worms' Crania sp. cf. C. blairi Rhipidomella sp. Caenanoplia logani? Productina lodgepolensis Leptagonia analoga Camarotoechia sp. Axiodeaneia platypleura Cleiothyridina sp. Crurithyris parva? Cyrtina burlingtonensis Hustedia texana Nucleospira obesa Plectospira? problematica Spirifer sp. Dielasma? sp. cf. D. utah Platyceras sp. Several other genera Unidentified small bivalves Triboloceras digonum Imitoceras sp. Gattendorfia costata Pericyclus rockymontanus Rotopericyclus sp. Spirorbis sp. Caenanoplia logani? Buxtonia? sp. Leptagonia analoga Camarotoechia metallica C. tuta C. inaequa? Cleiothyridina obmaxima C. glenparkensis C. sp. cf. C. incrassata Eumetria osagensis? Hustedia texana Prospira greenockensis ? Punctospirifer solidirostris Reticulata cooperensis? Spirifer missouriensis S. albapinensis Dielasma sp. cf. D. utah Platyceras paralius P. 3 sp. Goniospira sp. Bellerophon sp. At least 8 other genera Palaeoneilo missouriensis Allorisma? sp. Leptodesma sp. 1 orthoconic genus 1 or more goniatites Spirorbis nodulosus S. sp. Tentaculites sp. Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 Table 3. Continued. Fossil group Lower blastoid fauna Middle blastoid fauna Trace Fossils Arthropoda Trilobita Ostracoda Echinodermata Crinoidea Inadunata Camerata Blastoidea Fissiculata Spiraculata Vsteroidea Ophiuroidea Edrioasteroidea Echinoidea Holothuroidea Conodonta Vertebra 1. 1 Osteichthyes Totals Cosmorhaphe sp. Scalarituba missouriensis Horizontal burrows Several genera Amphelecrinus madisonensis Linocrinus walsallensis Unidentified microerinoids Abactinocrinus rossei Actinocrinites sp. Platycrinites bozemanensis Orophocrinus macurdai O. sp. cf. O. gracilis Tanaoblastus haynesi T. allanensis Strongyloblastus breimeri S. sp. Metablastus milliganensis Achistrum coloculum A. gamma Eocaudina columcanthus E. subhexagona E. marginata Microantyx botoni M. mudgei Rota campbelli R. martini Siphonodella crenulata Several other genera & species from Lower crenulata Zone 67+ Genera Cosmorhaphe sp. Scalarituba missouriensis Zoophycos sp. Horizontal burrows Richterella snakedenensis? Several genera Amphelecrinus madisonensis Cactocrinus arnoldi Platycrinites bozemanensis Rhodocrinites douglassi Koryschisma elegans Hadroblastus sp. Cryptoblastus? sp. B C? sp. C Strongyloblastus laudoni Montanablastus baldyensis Starfish arm 2 unidentified genera 2 specimens of 1 genus Archaeocidaris aliquantula Siphonodella crenulata Siphonodella isosticha Several other genera & species from Up- per crenulata-isosticha Zone Brachyodont crushing tooth 83+ Genera Mississippian Blastoids from Montana • Sprinkle and Gutschick 113 hypodeltoid, pointed or hooded, having wide growth front on thecal surface; reg- ular deltoids fairly small, either barely ap- pearing on thecal surface with tiny exter- nal DR growth sector, or confined to ambulacral sinuses, radials strongly over- lap deltoids near thecal surface but overlap gradually reverses deeper into sinuses; am- bulacra moderately long, linear to lanceo- late, extending out from mouth in shallow sinuses or down theca in relatively deep sinuses, lancet slightly exposed, side plates usually conceal about two-thirds of slits in sinus walls; brachioles small, ridged on sides; stem made up of flanged columnals, cirri and rootlets present distally for at- tachment of stem to substrate using re- cumbent rhizoid holdfast. Occurrence. Early Mississippian (Late Kinderhookian = Tournaisian) to latest Early Carboniferous (Late Visean and Early Namurian). Montana, New Mexico, Algeria. Etymology. The generic name is de- rived from korys, korystos (Greek), crest- ed, and schisma (Greek), slit, referring to the strongly raised deltoid crests bearing hydrospire slits in this genus. Discussion. Koryschisma is represented by an excellent collection of silicified ma- terial from the middle Lodgepole Lime- stone at Bandbox Mountain in west-central Montana, including about 145 complete or partial thecae, several hundred separate plates and ambulacral fragments, several hundred stem segments and individual columnals, and even a few brachiole frag- ments. Quality of the silicification is gen- erally very good, making it easy to study the morphology, and the numerous sepa- rate plates have yielded additional infor- mation about internal features. Koryschisma differs from other Devo- nian and Mississippian phaenoschismatids by having medium to high deltoid crests, two anal deltoids with the hypodeltoid oc- curring on the thecal surface, and medi- um-length ambulacra with the lancet part- ly exposed. It most closely resembles Leptoschisma and Pleuroschisma from the Devonian and Phaenoschisma and Had- roblastus from the Mississippian. It ap- pears to be intermediate between these Devonian and Mississippian genera, as not- ed by Breimer and Macurda (1972, p. 219, textfig. 104). Koryschisma differs from Leptoschis- ma by having only two anal deltoids, larg- er deltoid crests with more hydrospire slits exposed, somewhat wider ambulacra with the lancet partly exposed, and no BA axis in the basals. Koryschisma differs from Pleuroschisma by having only two anal deltoids, wider and less depressed ambu- lacra that conceal more hydrospire slits and have the lancet partly exposed, usually lower deltoid crests with fewer hydrospire slits, and other minor differences. Kory- schisma differs from Phaenoschisma by having a prominent hypodeltoid on the thecal surface, usually narrower ambula- cra having less of the lancet exposed and covering fewer of the hydrospire slits, and (in the type species) regular deltoids that barely appear on the thecal surface. Ko- ryschisma differs from Hadroblastus by having a more elongate thecal shape, higher deltoid crests with depressed am- bulacra, less exposure of the lancet, usually fewer hydrospire slits, some of which are concealed, and other differences. Breimer and Macurda (1972, pp. 18-20, 217-221) and Macurda (1983, pp. 60-65) described some of the morphologic and growth features of Koryschisma elegans (then unnamed), and informally assigned two other phaenoschismatid species to this genus. We agree with their assignments, and have briefly diagnosed and compared these other two species (Koryschisma sa- harae and K. parvum) with the type species described here in detail. KORYSCHISMA ELEGANS Sprinkle and Gutschick, new species Plate 2, Figures 1-59; Text-Figures 9 and 11-12 "New Lower Mississippian genus from Montana; Phaenoschismatid n. gen.; 'undescribed phaeno- schismatid B' (UB); Phaenoschismatid, new genus, new species. Miss., Lodgepole Fm., Bandbox Moun- tain, Cascade Co., Montana, USA.; undescribed 1 14 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 phaenoschismatid genus; undescribed genus; X' . . . to 1 .53 and averages 1 .39, V/P ratio ranges undescribed phaenoschismatid from Montana (UB); from 0.48 to 0.80, averaging 0.67, and pel- phaenoschismatid (UB) . from the Mississippian yic je ranges from 42° to 58° and av- i2&Vi?5tt^£SSi -ages 52'. Summit nearly flat with sharp 348, 362, and 366, textflgures 72, 100, and 104, and fluted adoral deltoid edges. table 2; Macurda, 1983, pp. 61-62 and 189, table Basals three, medium-sized forming 20. about 40% of pelvis, normally arranged, two larger and one smaller (azygous), azy- Diagnosis. Theca large, obconical, L/W gous basal elongate pentagonal, larger bas- averages 1.39, pelvis somewhat longer than als hexagonal; in large basal set (Plate 2, vault, V/P averages 0.67, pelvic angle av- Figs. 46-48), azygous basal 5.7 mm long, erages 52°, crests moderately high with 4.7 mm wide, larger basal about same sharp raised edges, adoral edge of deltoids length and 5.7 mm wide; stem facet formed serrated, even with summit; ambulacra by prominent secondary deposits bridging long, lanceolate, lancet about one-fourth over triangular tip of basals to form large, of ambulacral width; 3-9 hydrospire slits nearly circular platform bearing stem fac- per group, number slightly reduced on anal et with small central lumen. Oblique deep side; hypodeltoid prominent, hooded, oth- depression about 0.8-1.0 mm long near er deltoids barely appearing on thecal sur- middle (C ray) of BD basal about 1.0 mm face, heavy secondary deposits at tip of from stem lumen (Plate 2, Fig. 42), ap- basals; brachioles ridged; stem long with parent site of internal organ near thecal alternating flanged columnals, apparently base. attached distally using a recumbent rhi- Radials five, large, forming most of the- zoid holdfast. cal surface and 60% of pelvis, RD axis Description. About 145 partial and greater than RB axis at all sizes; each radial complete specimens plus about 500 sepa- roughly rectangular with deep ambulacral rate plates, ambulacral pieces, stem seg- sinus in adoral end, sides convex, profile ments and columnals, and brachiole seg- convex with large radial lip at origin con- ments available for study. Type specimens tinuing pelvis profile; each ambulacrum include holotype MCZ 915, 30 paratype strongly depressed below edge of radial thecae, including the 16-specimen growth sinus, which has sharp raised ridge about series studied by Breimer and Macurda 1 mm higher than plate surface (Plate 2, (1972), and 31 paratype fragments or Figs. 14-15). plates. Regular deltoids four, small, crested, Theca obconical, pelvis longer than barely reaching thecal surface (tiny vault, maximum width at tips of ambu- V-shaped external DR growth sector just lacra above midheight (Text-Fig. 11A), aboral to end of crest), crests horizontal on pelvis conical with nearly straight sides summit with wavy, serrated, or "cocks- (basal profile very slightly convex, radial comb" edge, forming incipient paired spi- profile very slightly concave), stem facet racles adorally behind small deltoid lip relatively large with prominent secondary (Plate 2, Fig. 11), small spine often on lip deposits, interambulacra nearly straight between spiracles, mouth rounded pentag- ignoring large radial lips, slightly concave onal, about 1.2 mm in diameter in large with lips (Plate 2, Fig. 11). Holotype (larg- specimen, radials strongly overlap deltoids est theca in available collections; Plate 2, at top of sinus but overlap slightly reversed Figs. 1, 12-13) 19.0 mm long, 12.4 mm at and below edges of ambulacra (Plate 2, wide, with a vault 7.9 mm long and pelvis Fig. 32). 11.2 mm long; smallest theca (Breimer and Anal deltoids two, medium-sized epi- , 1972, textfig. 72.1) about 5.8 mm deltoid with long depressed limbs and small nd 4.8 mm wide. In eight complete diamond-shaped hypodeltoid on thecal MCZ thecae, L/W ratio ranges from 1.27 surface. Epideltoid inverted V-shaped, lip Mississippian Blastoids from Montana • Sprinkle and Gutschick 115 BR -HS BCP OSP 1 1 1 ; i i -R, .-. B D G Text-Figure 11. Morphology of Koryschisma elegans, n. gen., n. sp. A-B, side and summit views of a large theca based on holotype MCZ 91 5 and paratype MCZ 91 6 showing greatest width (short lines) above midheight, enlarged hypodeltoid appearing on thecal surface, and heavy secondary deposits forming stem facet. C, much-enlarged epideltoid based mostly on paratype MCZ 949; note location of anus (A), adoral ends of hydrospire slits (HS) on limbs, and ridge on right limb extending further adoral than left limb. D, enlarged view of ambulacrum in paratype MCZ 942 showing central lancet (L) and inner and outer side plates (ISP and OSP) bearing brachiole facets (BF) at the edge; small arrow points toward mouth (M). E, enlarged cross section of ambulacrum and adjacent radial (R) drawn mostly from paratypes MCZ 956, MCZ 942 (inside of ambulacrum), and MCZ 924 (hydrospires); note side plates (SP) wrapping around edge of central ridged lancet (L) with slits in the ambulacral sinus (AS) leading to internal hydrospire folds (HF). F, much-enlarged side view and cross section of isolated brachiole fragment MCZ 959 showing ridge (Rl) on biserial brachiolar plates (BP) and tiny brachiolar cover plates (BCP). G, much-enlarged side view and columnal face of proximal stem segment MCZ 931 ; note equatorial flange (FL) on alternating columnals and smaller central lumen (LU). slightly wider than other deltoid lips, limbs infolded into hydrospires with folds ex- tending from limbs into space for hindgut, adorally ridge from "C" limb slightly higher than that for "D" limb (Text-Fig. 11C; Plate 2, Figs. 17, 30). Hypodeltoid greatly enlarged over other deltoid bodies with large external HDR sector, slightly to moderately hooded, adoral edge project- ing slightly above other deltoid crests and summit (Plate 2, Figs. 14, 21), forms aboral side of elliptical anus slightly larger than mouth (1.8 mm long in 18 mm long theca), forms strongly convex sutures with radials, grows aborally from near tip of hood. Ambulacra five, relatively long, mod- erately wide, lanceolate, moderately convex in cross section, slightly curved in profile, lancet slightly exposed in center, one-third to one-half of its width and one- fourth of ambulacral width, side plates curve around lateral edges of lancet (Text- Fig. HE; Plate 2, Figs. 18, 34, 37), lancet grooved at bottom with adoral keel, inner side plates large, constricted abmedially, outer side plates small, rounded triangular, notch abmedial aboral edge of inner side plate, inner and outer side plates form bra- chiole facet at abmedial edge of ambula- crum, brachiolar pit small, at end of side food groove on side plate suture (Text-Fig. 11D), brachiolar facets together hemiel- liptical, about 0.25 mm long, canted to- ward each other so that deepest part be- tween them on side plate suture. Side food grooves enter main food groove at 45-70° angle, four lobes per side plate along main food groove, 3-5 adorally and two small lobes aborally along side food grove. Oral cover plates present on summit of one paratype (UMMP 60694; Plate 2, Fig. 38), form five domed covers about 0.4 mm 116 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 high and 0 6 mm wide made up of tiny Disarticulated stem material abundant plates about 0.15 mm in size over adoral (Text-Fig. 9); distinctively flanged and ambulacra converging at mouth. Sym- somewhat heteromorphic proximally, de- metry appears pentagonal over mouth with veloping cirri and rootlets distally. Longest no apparent "2-1-2" arrangement in cov- stem segment 22 mm long with 33 colum- ers. Several paratvpes (especially UMMP nals (Plate 2, Fig. 56); one short stem seg- 65893) show remnants of small spines about ment attached to small basal set (Plate 2, 0.6 mm long and 0.15 mm in diameter Fig. 59); one theca had single columnal near mouth and anus (Plate 2, Fig. 38), attached but lost it during etching. Prox- apparentlv to protect these summit struc- imal columnals thin, wide, with a large tUres. flange (Plate 2, Figs. 52, 54, and 56); typ- Hydrospires in 10 groups, slits partly ex- ical proximal columnal 0.4 mm long, 1.6 posed in sinus and crest walls, mostly hid- mm wide, having a circular equatorial den beneath ambulacra, 3-9 folds per nor- flange 2.5-3.0 mm in diameter; columnal mal group, number slightly reduced on faces round with 49-50 small crenulae anal side to 2-7 folds; lower folds hang around margin and small, nearly circular down into thecal cavity, deepest at radio- lumen 0.1 mm in size in center (Text-Fig. deltoid suture (Plate 2, Figs. 16, 33), upper 11G). Two or three sizes of flanged colum- folds extend laterally in from sinus edges, nals alternating in proximal stem, either short slit and fold at top of sinus (Plate 2, in sequence "lg.-sm.-med.-sm., lg.-sm.- Fig. 15) probably added late in growth; in med.-sm., . . ." or as "lg.-sm., lg.-sm., . . ." internal view folds bend abmedially at ra- (Plate 2, Figs. 54, 56). Distal columnals diodeltoid suture, folds pinched together longer (0.5-0.7 mm long), narrower (1.2- at aboral end near radial origin (Plate 2, 1.3 mm wide), with smaller flanges (1.3- Fig. 16); adoral edges of ambulacra form 1.5 mm in diameter) that alternate some- incipient spiracles at adoral edge of deltoid what in size and appear partly covered by crests. subsequent lateral growth of columnals Ornament consists of fine to medium- (Plate 2, Fig. 55). Cirri (rootlets?) attached strength growth lines parallel to margins to distal columnals on flanges or sutures on basals and radials, stronger growth lines (Plate 2, Fig. 58), most cirrals about 0.25 on RHD front (Plate 2, Figs. 15, 17), very mm long, 0.5-0.8 mm in diameter, with fine growth lines on hypodeltoid and sides faces bearing 11-14 small crenulae. Cirri of deltoids. Several heavy layers of sec- apparently concentrated on one side of best ondary deposits over origins of basals to distal stem segment (Plate 2, Fig. 58), im- produce large circular platform for stem plying a recumbent rhizoid holdfast (Brett, attachment from smaller triangular tip of 1981, pp. 348, 351). Total length of stem basal cone (Plate 2, Figs. 40-42, 46-47, unknown, but hardly any gradation in size and 49); secondary deposits also forming or morphology noted in longest preserved large, pointed, radial lip up to 1 .5 mm long segments. that continues pelvic profile, lip covers few Brachiole segments up to 6 mm long also growth lines at origin of each radial, and preserved in acid residues (Plate 2, Figs, bears median raised ridge adorally to sep- 23-24); brachioles ridged, roughly pen- arate brachiole groups (Plate 2, Figs. 12, tagonal in cross section, biserially plated 17). Thin 1 mm high ridge of secondary (Text-Fig. 1 IF); brachiolar plates about 0.3 calcite along edge of each ambulacral sinus mm long, 0.33 mm wide, and 0.15 mm above plate surface (Plate 2, Figs. 15, 27); deep (across food groove), possibly one bi- several lateral-pointing spines of apparent serial set of slightly domed, triangular, bra- secondary calcite on adoral-most side plates chiolar cover plates over shallow, V-shaped (Plate 2, Fig. 34); and small spine of sec- food groove (Text-Fig. 11F), about three ondary calcite on some deltoid and epi- brachiolar cover plates per brachiolar plate deltoid lips. on each side. MISSISSIPPIAN Blastoids from Montana • Sprinkle and Gutschick 117 20 16 E E 12 g 8 20 16 12 rt 8 > 4 8 12 Width (mm) • % •+-% 16 0 4 8 12 Pelvis (mm) 15 12 E E I 6 < / 10 20 30 No. Side Plates 40 15 I" • CO 9 DC o CC 6 DC a Q 3 CC / / o°o 8P 2.9 -2.0 Hadroblastus sp. MCZ 748 16.2 -12.0 6.0 10.1 — 1.1 -6.1 -4.4 Orophocrinus cf. MCZ 840 MCZ 836 MCZ 838 MCZ 839 O. gracilis 6.8 16.1 6.5 -15.1 2.6 8.7 -10.0 4.2 7.5 -65° -95° -70° -1.2 1.6 2.3 -2.4 4.6 4.6 -2.0 >4.3 -6.5 >6.0 Orophocrinus sp. MCZ 884 14.0 13.1 4.8 9.2 75° 2.8 4.0 5.5 Mctablastus milliganensis MCZ 804 MCZ 803 (Holotvpe) MCZ 805 -11.0 11.0 -13.7 6.4 6.6 8.2 5.7 4.6 6.0 5.2 6.4 -7.7 -60° -55° -50° 0.8 -3.5 3.7 2.9 3.2 Montanablastus baldyensis MCZ 889 MCZ 893 MCZ 886 (Holotvpe) MCZ 892 6.4 7.7 8.6 10.2 5.0 -5.6 6.5 7.3 3.0 3.8 5.1 5.7 3.3 3.9 -3.6 5.1 -75° -65° 80° 75° -0.6 0.7 -0.9 0.7 1.8 1.9 1.9 2.5 1.6 2.0 2.3 2.8 Koryschisma elegans differs from K. sa- harae by being less elongate (lower L/W ratio), having a higher V/P ratio, higher crests with wider ambulacra, a longer, more pointed hypodeltoid, and by being consid- erably older (Late Tournaisian equivalent vs. Late Visean to Early Namurian). KORYSCHiSMA SAHARAE (Breimer and Macurda), 1972 Pentremites sp., Pareyn, 1961, pp. 223-224. Phaenoschisma? saharae, "Phaenoschisma" saha- rae, Breimer and Macurda, 1972, pp. 18-20, 387, plate II, figures 4-5 and 10; Macurda, 1983, pp. 61-65, plate 14, figures 1-13, table 21. Diagnosis. Theca large, elongate coni- cal, L/W averages 1.71, pelvis much long- er than vault, V/P averages 0.23, pelvic angle averages 38°; deltoid crests low to medium, hypodeltoid large, other deltoids appear confined to ambulacral sinuses; era nearly linear, lancet making up about one-fourth of width; 5-9 hydrospire slits per group, number reduced by about one-third on anal side; subdued secondary deposits at tip of basals. Discussion. This species, from the Late Visean and Early Namurian of Algeria, is larger and more elongate than K. elegans and K. parvum, with a shorter vault, nar- rower ambulacra, lower crests, and more subdued secondary deposits around the large stem facet. KORYSCHISMA PARVUM (Macurda), 1 983 "UA (undescribed phaenoschismatid A . . .); phae- noschismatid (UA)," Breimer and Macurda, 1972, pp. 217-219, plate IV, figures 17, 20, plate V, fig- ures 1-2, textfigure 71. Phaenoschisma? parvum, Macurda, 1983, pp. 60-61, plate 13, figures 9-10, 14-17, 19-20, 23-24, table 19. Diagnosis. Theca small, widely biconi- cal, L/W averages about 1.1, pelvis longer than vault, V/P averages about 0.57, pel- Mississippian Blastoids from Montana • Sprinkle and Gutschich 121 (continued) Text-Figure 12). A - preceding a number indicates that this measurement was esti- mated IN A DAMAGED, INCOMPLETE, OR CRUSHED SPECIMEN; SPECIMENS WITH A — WERE TOO INCOMPLETE OR DAMAGED TO MEASURE. RB front (mm) RR axis (mm) RR front (mm) RD axis (mm) RD front (mm) Del len. (mm) Del. wid. (mm) Amb. len (mm) Amb. wid (mm) No. of side pis -1.1 0.9 2.0 -1.0 -1.5 ■1.0 -4.6 3.3 5.6 4.7 2.3 -4.5 -4.3 -6.5 -1.2 >18 -4.5 2.0 — 3.0 0.4 1.4 0.9 4.2 1.0 15 4.3 — 7.2 1.2 3.0 2.1 >9.2 >1.3 -19 5.0 -7.0 7.1 1.3 — -2.2 >9.0 -2.0 >20 5.5 >9.2 9.2 1.2 4.9 2.4 11.2 1.7 28 -4.5 4.1 -8.0 -5.0 1.6 3.5 2.0 7.1 1.7 -29 2.5 1.8 7.3 5.5 <0.1 <0.1 <0.2 2.5 1.9 6.9 5.7 <0.1 <0.1 <0.2 2.5 — — -6.9 <0.1 <0.1 <0.2 1.5 1.1 3.7 3.0 -0.2 0.7 -0.4 1.8 1.6 -5.0 3.9 — -0.7 — 2.2 1.7 5.7 4.7 -0,3 0.9 0.6 2.8 2.2 -7.0 5.6 0.8 -1.9 -0.9 5.4 0.8 -19 5.1 0.7 20 6.5 0.7 -23 2.9 0.8 13 4.3 1.0 15 4.6 1.0 16 6.2 >0.7 -15 vie angle averages 67°; deltoid crests high, slope down to mouth, hypodeltoid rela- tively large, on thecal surface, other del- toids confined to ambulacral sinuses; am- bulacra lanceolate, lancet only slightly exposed in center; 4-6 hydrospire slits per group, number slightly reduced on anal side. Discussion. This species, from the Osa- gean of New Mexico, is smaller and much wider than K. elegans and K. saharae, with a shorter vault, higher deltoid crests, and fewer hydrospire slits. Because of its small size, it may be a paedomorphic derivation of the slightly older K. elegans. Genus PHAENOSCHISMA Etheridge and Carpenter, 1886 Type Species. Pentatrematites acutum Sowerby, 1834. Diagnosis. Fissiculate blastoids with a conical to obconical theca; 10 hydrospire groups having slits partly exposed, number of slits slightly reduced on anal side; two anal deltoids, hypodeltoid small, not on thecal surface; ambulacra lanceolate, lan- cet widely exposed. Occurrence . Early to Middle Mississip- pian, central and western U.S.A., Early Carboniferous, England and Ireland. PHAENOSCHISMA? SP. Plate 3, Figure 28; Text-Figure 13B; Table 4 A single small specimen of an apparent phaenoschismatid was found in the float near the top of the Lodgepole Limestone at Saddle Peak in the Bridger Range. It was preserved as a crushed but nearly complete calcitic theca on a slab (Plate 3, Fig. 28); the theca was extracted using an air abrasive unit, but unfortunately it proved to be incomplete with only the bas- als and radials still preserved. Theca about Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 AMB H A Text-Figure 13. A, reconstructed side view of Hadroblastus sp. based on MCZ 748; note thecal shape (greatest width at short lines), wide ambulacral sinuses with hydrospire slits (HS) exposed, curved raised ambulacra (AMB), and size of proximal stem (PS), x 1.8. B, slightly reconstructed side view of Phae- noschisma? sp. (MCZ 885) showing shape (greatest width at short lines), long basals and radials, and inferred shape of missing vault with short ambulacra, *3.8. 5.0 mm long with incomplete base and missing deltoids and ambulacra, original length at least 5.2 mm, maximum width (crushed) 3.9 mm, original width estimat- ed at 3.2 mm; pelvis about 4.6 mm long and incomplete vault at least 0.6 mm long. L/W ratio probably near 1.5-1.6 origi- nally, V/P ratio about 0.13 originally, and pelvic angle now 40° on crushed side. Pelvis conical, straight to slightly con- cave in profile; basals three, appear nor- mally arranged with two larger, one small- er, pointed at top, occupy 50-60% of preserved pelvis, about 2.9 mm long with small amount added for missing stem fac- et; stem facet slightly triangular, second- ary deposits not observed. Radials five, pentagonal, occupy slightly less of pelvis than basals, notched at top for ambulacra and deltoids, notch on apparent posterior side larger than others implying external hypodeltoid possibly present (Text-Fig. 13B), regular deltoids small. No deltoids or ambulacra preserved, only ends of hydrospire slits on adoral edges of radials, apparently at least four hydrospire slits per ambulacral side. Other summit structures unknown. The only theca is MCZ 885 which was found in the float about 100-200 ft (30- 61 m) below the top of the Lodgepole Limestone at Saddle Peak, southern Brid- ger Range, southwestern Montana. Discussion. This single specimen is too incomplete to tell whether it belongs to the genus Phaenoschisma, but this is consid- ered the most likely possibility based on its preserved morphology and Early Mis- sissippian age. It is rather similar in side view to P. laevisculum and to P. gracilli- mum (see Breimer and Macurda, 1972, plate 3, figures 14, 19, and 26-27), both from the similar-aged Burlington Lime- stone. These species are somewhat larger, differ slightly in their thecal proportions, and do not have an enlarged hypodeltoid contributing to the posterior thecal sur- face. This is the only relatively complete theca of a possible Phaenoschisma known from the Lodgepole Limestone, but a few elongate basals perhaps belonging to a sim- ilar blastoid are also known from the lower fauna in the Allan Mountain Limestone at the North Sawtooth Mountain Section in northwestern Montana. Family NEOSCHISMATIDAE Wanner, 1 940 Genus HADROBLASTUS Fay, 1962c Type Species. Hadroblastus convexus Fay, 1962c. Diagnosis. Fissiculate blastoids with bi- convex theca, vault usually shorter than pelvis, deltoid crests low to medium; 10 hydrospire groups, slits almost completely exposed in wide shallow sinuses alongside ambulacra, slits usually reduced on anal side; two anal deltoids, epideltoid forms anal hydrospires, hypodeltoid forms part of theca wall; ambulacra lanceolate, often raised, lancet exposed throughout length. Occurrence. Early to Middle Mississip- pian, central and western U.S.A.; Early Carboniferous, Ireland? and Scotland? HADROBLASTUS SP. Plate 3, Figure 29; Text-Figure 13A; Table 4 Hadroblastus sp., Breimer and Maeurda, 1972, pp. 30, 382, plate 18, figure 1. A single specimen from an unknown po- sition in the middle Lodgepole Limestone was found at Standard Creek, southwest- Mississippian Blastoids from Montana • Sprinkle and Gutschich 123 BRF BRF OSP BRF ISP D BRF I Text-Figure 14. Morphology of Orophocrinus macurdai, n. gen., n. sp., (A-B, E-F), Orophocrinus cf. O. gracilis (Meek and Worthen) (C-D), and O. sp. (G-l). A-B, side and summit views of a large theca (based on MCZ 821 and holotype MCZ 811) showing shape, maximum width (short lines), fairly wide ambulacra with spiracular slits alongside, slightly concave interambulacra, and size and shape of hypodeltoid. C, reconstructed side view based on MCZ 836 showing shape, longer ambulacra, and location of greatest width (short lines) near midheight. D, enlarged plan view of ambulacrum in MCZ 836 showing lancet (L) slightly exposed in center (small arrow points toward mouth [M]), large inner and small outer side plates (ISP and OSP) occupying most of width and supporting a brachiole facet (BRF) near edge, *9.5. E-F, enlarged plan view and cross section of ambulacrum in paratype MCZ 823; note lancet (L) exposed in center, large inner and small elongate outer side plates (ISP and OSP) together supporting an elliptical brachiole facet (BRF), well-developed cover plate lobes and sockets, and convex cross-sectional shape with depressed, outward-slanting facets, x 1 1 .8. G-H, side and summit views of MCZ 884 showing similarity in thecal shape to A and B except for more concave interambulacra. I, cross section of E ambulacrum in MCZ 884 showing concave surface with inward-slanted brachiole facets (BRF) quite different from F, x7.1. ern Montana. This specimen was figured by Breimer and Macurda (1972) using a pre-preparation photograph supplied by Sprinkle in 1966. Subsequently, the spec- imen was partly uncovered using an air abrasive unit although the matrix proved too hard and deep to uncover an entire side (Plate 3, Fig. 29). Part of the proximal stem was also found still attached to the theca. Description. Only known specimen partly buried and crushed on slab with exposed plates silicified. Theca fairly large, apparently biconvex, 16.2 mm long, at least 12 mm wide (incomplete but crushed), vault 6.0 mm long, pelvis 10.1 mm long, L/W ratio approximately 1.35 based on exposed width, V/P ratio 0.59. Pelvic an- gle difficult to measure, perhaps 70-80° originally (Text-Fig. 13A). Basals fairly long, occupying 50-60% of pelvis, at least 7 mm long, azygous basal quadrate in shape, about 3.5 mm wide. Radials fairly large, about 6.5 mm long, perhaps as much as 6.5 mm wide, radial body about 4.5 mm long, shallow ambu- lacral sinuses about 2 mm long. Deltoids difficult to see, occupying broad ambula- cral sinuses, approximately 3.5 mm long and about 3.5 mm wide, little or no deltoid crest present. Ambulacra occupying cen- ters of broad ambulacral sinuses, at least 6.5 mm long and about 1.2 mm wide, ap- pear to be flat to slightly convex in cross section, strongly convex in lateral view and considerably raised above surrounding si- nuses (Text-Fig. 13A), too highly silicified to distinguish lancet or side plates. Hydro- spires fully exposed, apparently 7-8 per ambulacral side, longest slits extending nearly full length of adjacent ambulacra, slits converge at center-line of deltoid which is not raised into crest above sinuses. Few brachioles attached to left ambu- 124 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 lacrum in this specimen, brachioles incom- plete, about 10-11 mm long, approxi- mately 0.3 mm wide and deep, poorly preserved because of partial silicification (Plate 3, Fig. 29). Proximal stem attached to facet on bas- als, preserved stem about 12.5 mm long extending off edge of slab (Plate 3, Fig. 29), about 1 mm in diameter both proxi- mally and distally, made up of at least 47 columnals varying from about 0.17 mm long proximaliy to about 0.33 mm long distally. Ornament on thecal plates difficult to see because of silicification and abrasion of plates during preparation, no trace of coarse ornament or growth lines. Material and Occurrence. Only known specimen is MCZ 748 from an unknown height in the middle Lodgepole Lime- stone, slab found in the float above the lower cliffs containing Tanaoblastus at Standard Creek, Gravelly Range, south- western Montana. Discussion. This blastoid from the mid- dle Lodgepole Limestone may represent a new species of Hadroblastus, but is not named here because the only known spec- imen is not well preserved or exposed. This form is larger and more elongate than the type species H. convexus Fay (1962c), which has a small squat theca with a low vault and moderate deltoid crests. It has a higher vault with lower deltoid crests than H. breimeri Ausich and Meyer (1988). It differs from H. whitei by having longer hydrospire fields without deltoid crests and perhaps fewer slits (7-8 vs. 9-10). It may have been similar to H. blairi, especially in ambulacral height and curvature, but was less squat and had almost no deltoid crests. Family OROPHOCRINIDAE Jaekel, 1918 Genus OROPHOCRINUS von Seebach, 1864 Type Species. Pentremites stelliformis Owen and Shumard, 1850. Diagnosis. Fissiculate blastoids having a conical, conoidal, or parachute-shaped theca, ten long spiracular slits and hydro- spire groups alongside ambulacra; 4-11 hydrospire folds per group; two anal del- toids present, relatively small epideltoid with long aboral limbs and relatively small hypodeltoid visible on thecal surface; am- bulacra relatively wide, usually raised, lan- cet narrowly exposed along much of length. Occurrence. Early to Middle Mississip- pian, central, southwestern, and north- western United States; Early Carbonifer- ous (Tournaisian and Visean), Belgium, Great Britain, and Ireland. Discussion. Two species of Orophocri- nus occur in the lower Lodgepole Lime- stone, and an additional specimen of a third species occurs in the upper Lodgepole Limestone of southwestern Montana. These occurrences extend the geographic range of this genus into the northwestern United States. Orophocrinus is a very wide-rang- ing genus in the Mississippian (Early Car- boniferous) known from both North America and Europe. It differs from sim- ilar genera in the Orophocrinidae such as Brachyschisma by having a full set of anal hydrospires and only two anal deltoids, from Katoblastus by having the hydro- spire slits completely hidden and only two anal deltoids, and from Pentablastus and Acentrotremites by having a different the- cal shape with ambulacra that do not usu- ally extend down the theca. OROPHOCRINUS MACURDA! Sprinkle and Gutschick, new species Plate 1, Figure 2; Plate 3, Figures 1-16; Text-Figures 14A-B, E-F, and 15 Diagnosis. Theca conical, L/ W ratio av- eraging 1.07, V/P ratio averaging 0.29, pelvic angle averaging 64°, interambula- cra flat to slightly concave, RD axis less than RB axis at all sizes, hypodeltoid widely borders spiracular slits, ambulacra strongly convex, brachiolar facets abmedial, usu- ally five hydrospire folds per ambulacral side. Description. Forty-one specimens and fragments available for study; description based on holotype MCZ 811, 11 additional Mississippian Blastoids from Montana • Sprinkle and Gutschick 125 15 i« CO 9 CC o S 6 £ 3 4 8 12 Width (mm) / ..# V d7 16 ..*' 4 8 12 Pelvis (mm) ~5 E ^4 _j 3 °2 1 / 15 12 £9 < 4 / / 5 Al- io 20 30 40 No. Side Plates + ,4 E 3 E 00 -• 0 o • ■// 3 6 9 12 15 0 1 2 3 4 0 3 6 9 12 Growth Front (mm) Max. Del. W. (mm) RDo RB» (mm) 15 12 E q . h Q DC 6 3 0 C / • y/ - • 1 2 4 6 8 RRo RB« (mm) 8 • E rx cc / 3 6 RB (mm) E E • cc CO o Q CC 15 12 9 6 ■ =/ 3 n I2 2 4 6 Del. L. (mm) Text-Figure 15. Growth plots for the 12 measured specimens (MCZ 811-822) of Orophocrinus macurdai, n. sp. Best-fit lines in all plots were hand fit, and short lines with central tick mark represent estimates for large incomplete holotype MCZ 81 1 that lacks basals. 126 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 complete paratypes in growth series, and 12 other paratype specimens and frag- ments. Theca conical, made up mostly of conical pelvis with straight to slightly con- vex sides, capped by convex vault (Text- Fig. 14A); growth series specimens rang- ing from 4.5 mm long to incomplete ho- lotype 13.2 mm long (no basals; original complete length estimated at 18.2 mm). In 12-specimen growth series, L/W ratio ranging from 0.88 to 1.18 and averaging 1.07, decreasing slightly during growth; V/P ratio ranging from 0.29 to 0.65 and averaging 0.41 for same specimens, in- creasing slightly late in ontogeny; pelvic angle ranging from 50° to 85° and aver- aging 64°, increasing slightly during growth. Greatest width at tips of ambu- lacra, well above midheight; cross section here pentagonal, interambulacral areas flat to slightly concave (Plate 3, Figs. 8-12). Basals three, normally arranged, two regular and one small (azygous), azygous basal quadrate, 3.0 mm long, 3.1 mm wide in medium-sized specimen, regular basals hexagonal, about same length and 4.3 mm wide, basals making up about 40% of pelvis (Plate 3, Figs. 5-7); some secondary de- posits extending short distance up inter- basal sutures from large round to some- what triangular stem facet 1.7 mm in diameter, with 0.2 mm lumen in center. Radials five, relatively long, RD axis less than RB axis at all sizes (Text-Fig. 15), 2 mm less in very large holotype (Plate 3, Fig. 13), RD front nearly straight except on posterior side where distinctly concave against hypodeltoid, large lip at radial or- igin pointing obliquely adoral. Regular deltoids four, relatively narrow, elongate hexagonal. Adoral part bulbous, with several thick overlayerings of second- ary calcite, middle part constricted, strongly concave in profile with raised ridge alongside adoral end of each spiracular slit, aboral part slightly concave in profile (Plate 1, Fig. 2), ornamented with medium, reg- ularly spaced, growth lines. DR sutures nearly straight, forming 150-160° angle, radials slightly overlap deltoids. Mouth pentagonal to star-shaped, surrounded by regular deltoid and epideltoid lips. Anal deltoids two, hypodeltoid fairly small, squat pentagonal, easily lost (Plate 3, Figs. 8-11); extends further down theca than adjacent regular deltoids, entire lat- eral margins border spiracular slits, sutures with radials often moderately curved, adoral edge usually raised in center form- ing hood over anus (Plate 3, Fig. 16). Epi- deltoid having small, pentagonal, adoral part bordering mouth and anus on oppo- site sides, and two long aboral limbs ex- tending down alongside anus and under hypodeltoid, epideltoid limbs infolded to form hydrospire folds below spiracular slit. Anus elliptical with hypodeltoid in place (Plate 3, Fig. 12), about same size as mouth. Ambulacra five, relatively short and wide, 7.5 mm long and 1.9 mm wide in very large holotype, in shape changing from petaloid to lanceolate during growth, strongly convex in cross section, even with or slightly raised above adjacent plate mar- gins (Plate 3, Figs. 1-7), lancet exposed, making up central 20% in adoral two-thirds of ambulacrum (Text-Fig. 14E). Inner and outer side plates supported by lancet, inner side plates grow laterally as they move up ambulacrum, forming raised abmedial lip around outside of large elliptical brachio- lar facets which are abmedial (Plate 3, Fig. 14), 8-9 side plate sets per 3 mm length of ambulacrum, tiny brachiolar pit at end of each food groove near highest point on each side of ambulacrum (Text-Fig. 14F). For each side plate, 3-4 lobes along main food groove plus 3-4 lobes adorally and 2-3 lobes aborally along side food groove (Text-Fig. 14E). Spiracular slits 10, slightly arcuate, moderately long, extending about two- thirds of ambulacral length, few milli- meters of aboral end closed off internally by radial growth beneath lancet, adoral end near narrowest point on deltoid, slits do not quite reach adoral edge of anus in CD interray (Text-Fig. 14B). Hydrospires usually five per ambulacral side (10 mea- surements), possibly four in few cases, pos- Mississippian Blastoids from Montana • Sprinkle and Gutschick 127 sibly six in one case, top slit about 0.6 mm width, vault much greater than pelvis, a deep below deltoid edge adorally, aboral moderate pelvic angle, RD less than RB at end of this slit sometimes visible near ra- all sizes, hypodeltoid widely bordering the dial lip when ambulacrum damaged or spiracular slits, convex ambulacra with ab- side plates missing, enlarged tube appar- medial brachiole facets, and usually five ently present at inner end of each hydro- hydrospires per ambulacral side. Oropho- spire fold. crinus macurdai is probably most closely Ornament consists of medium-strength, related to O. orbignyanus of Belgium and widely spaced, growth lines (Plate 1, Fig. perhaps to O. conicus from the Late Kin- 2; Plate 3, Fig. 13) on basals, deltoids, and derhook of the Mississippi Valley; all of most of radials; RD front of radials consists these species are nearly the same age. of coarse, widely spaced, growth lines Orophocrinus macurdai has only been (Plate 3, Figs. 7 and 14). Secondary de- found in a thin east-west strip of sections posits present around stem facet, at radial near the center of the study area in south- origin and along edges of ambulacra, and western Montana (see Text-Fig. 8). over adoral parts of deltoids (probably fill- Five additional poorly preserved spec- ing in adoral ends of spiracular slits). imens of Orophocrinus also from the lower Measurements of specimens in growth Lodgepole have a different thecal shape series graphed in Text-Figure 15. with much longer ambulacra than O. mac- Stem, brachioles, and cover plates un- urdai and apparently belong to a species known. very similar to O. gracilis from the Late Studied Specimens. Holotype MCZ 811, Kinderhook and Osage of the Mississippi paratypes MCZ 812-834 (23 specimens and Valley, fragments), and MCZ 835 (17 additional ^cTrrence. Known from the lower SROPHOCRINUS cf. O GRACILIS Lodgepole Limestone at five localities in (^ and Worthen) 1870 southwestern Montana; 20 specimens and !Ilatf *' higur?^ ' ' . . , fragments from Dry Hollow 20-50 ft (6- Text-F.gures 14C-D; Table 4 15 m) above the base of the Paine Member, Diagnosis. Theca conoidal, L/W ratio the holotype and 14 other specimens and about 1.1, V/P ratio about 1.3, pelvic angle fragments from Milligan Canyon East 12- about 83°, RD axis much greater than RB 20 ft (3.7-6 m) above the base, four spec- axis, hypodeltoid borders spiracular slits, imens and fragments from Milligan Can- ambulacra long, convex in cross section, yon 15-20 ft (4.5-6 m) above the base, and raised above thecal plates, brachiolar fac- single specimens from Sand Creek 23 ft (7 ets abmedial to central, 4-5 hydrospires m) above the base and from Little Ante- per ambulacral side. lope Creek in the float 20-50 ft (6-15 m) Description. Five poorly preserved and above the base. fragmentary specimens available closely Etymology. Named for D. Bradford resembling this species. Theca conoidal, Macurda, Jr., of The Energists, Houston, pelvis broadly conical, sides of pelvis near- who revised this genus and its species in ly straight, vault parabolic with long am- the 1960s. bulacra extending down theca (Text-Fig. Discussion. Orophocrinus macurdai is 14C; Plate 3, Fig. 22). Smallest apparent a fairly distinctive species and represents specimen 6.8 mm long, largest approxi- one of the earliest occurrences of the ge- mately 19 mm long (basals missing). L/W nus. It differs from other similar species, ratio 1.0 and 1.2 in two nearly complete such as O. orbignyanus and O. conicus, specimens, V/P ratio ranges from 0.6 to by having a conical shape throughout its 1.46, pelvic angle averages 83° in three growth with length slightly greater than incomplete and crushed specimens. Great- 128 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 est width at tips of ambulacra well below five lobes along main food groove and 1- midheight, cross section here pentagonal 2 more along adoral edge of side food with slightlv concave interambulacra. grooves (Text-Fig. 14D). Basals three, normally arranged, two Spiracular slits 10, long, nearly linear, larger and one smaller (azygous) make up appear to extend most of ambulacral length nearly 50% of pelvis; in large specimen but aboral 2-3 mm closed off by radial azygous basal quadrate 5.2 mm long, ap- growth beneath lancet, adoral end at nar- proximately 4.5 mm wide, larger basals rowest point on deltoids. Either four or five same length, about 6.2 mm wide. Stem folds per ambulacral side (two observa- facet large, 2.4 mm in diameter with small tions), folds thin with enlarged tube ap- central lumen 0.1 mm wide; small second- parently present at bottom (Plate 3, Fig. ary deposits up interbasal sutures to pro- 22). duce circular facet. Ornament consists of medium-strength Radials five, long, RD much greater than growth lines parallel to plate margins (Plate RB in all but smallest specimen, nearly 3 3, Fig. 24). Measurements for few known mm longer in largest theca, RD front near- specimens listed in Table 4. ly straight, fairly large lip at radial origin Studied Specimens. MCZ 836-840 (five pointing laterally (Plate 3, Fig. 22). partial specimens). Regular deltoids four, relatively narrow, Occurrence. Known from the lower elongate hexagonal. Adoral part with 1-2 Lodgepole Limestone at three localities in concentric growth lines, middle part con- southwestern Montana: two specimens stricted, concave, aboral part slightly con- from the talus piles at Standard Creek from cave in profile, growth lines subdued (Plate beds 15-55 ft (4.5-17 m) above the base 3, Fig. 24). DR sutures nearly straight, form of the Paine Member, two specimens from 160° angle, radials appear to overlap del- Dry Hollow 20-30 ft (6-9 m) above the toids. base, and a single specimen from Little Anal deltoids apparently two, missing Antelope Creek 26-35 ft (8-11 m) above or poorly preserved on all specimens ex- the base. cept smallest where epideltoid present Discussion. These five poorly preserved (Plate 3, Fig. 18). Hypodeltoid not seen specimens look somewhat different from but probably reaches spiracular slits be- specimens of Orophocrinus macurdai, with cause epideltoid limbs depressed below which they occur at two localities in Mon- thecal surface. Epideltoid has small pen- tana. Instead they closely resemble speci- tagonal part bordering mouth and anus mens of the distinctive Kinderhook and plus two depressed limbs extending ab- Osage form O. gracilis from the Mississip- orally and infolded to form hydrospires. pi Valley (see Macurda, 1965, pp. 1073- Anus probably elliptical in shape with hy- 1077). The thecal shape and long ambu- podeltoid present. lacra extending down the theca are very Ambulacra five, long and fairly narrow, similar (compare Plate 3, Figs. 21-22). No raised above adjacent thecal plates, linear hypodeltoid was seen, but it apparently to lanceolate, moderately convex, lancet borders the spiracular slits on both sides exposed in center along much of length because the epideltoid limbs are de- flate 3, Figs. 23-24). Inner and outer side pressed. The basal angle is similar, RD is plates supported by lancet, side plates ap- much greater than RB in all except the parently do not grow laterally, brachiole smallest specimen (Table 4), the number facets appear to be abmedial or perhaps of hydrospires is similar (four or five vs. central (Text-Fig. 14D). Longest ambu- four), and the brachiolar facets are in a lacra 1 1 .5 mm long and 2.0 mm wide with similar position on the ambulacra. In ad- 25 side plate sets, over much of am- dition, some specimens from the Mississip- lm six side plates per 3 mm length; pi Valley (O. cf . O. gracilis from the Mississippian Blastoids from Montana • Sprinkle and Gutschick 129 B D Text-Figure 16. Morphology of Metablastus milliganensis, n. sp. A-B, side and summit views of reconstructed theca based on holotype MCZ 803 and paratype MCZ 804 showing greatest width (short lines) just above midheight and slightly concave interambulacra. C, enlarged cross-sectional view of broken ambulacrum in paratype MCZ 806; note that lancet (L) is "keeled" on the interior, covered externally by side plates (SP), and lacks hydrospires beneath it or the adjacent radials (R). D, much- enlarged plan view of ambulacrum in holotype MCZ 803 showing inner and outer side plates (ISP and OSP) covering lancet, pores (P) at edge of ambulacrum between brachiole facets (BF), and main food groove (MFG) and short side food grooves (SFG) bearing cover plate lobes and sockets. Small arrow points in direction of mouth (M). Northview Shale of southwest Missouri; see Macurda, 1965, pp. 1075 and 1077) are nearly the same age as the Lodgepole ma- terial. OROPHOCRINUS SP. Plate 3, Figures 25-27; Text-Figures 14G-I; Table 4 A single coarsely silicified specimen of Orophocrinus was found near the top of the Lodgepole Limestone in the southern Bridger Range. The specimen was etched from the slab on which it was collected, and was found to differ from the two Orophocrinus species from the lower Lodgepole. However, it is not well enough preserved to establish a new species name for it, but is briefly described and figured here. Description. Theca conical in shape with rounded vault and conical pelvis; fairly large theca 14.0 mm long, 13.2 mm wide, giving L/W ratio of 1.1; vault 5.1 mm long, pelvis 8.9 mm long, giving V/P ratio of 0.57; pelvic angle about 75° (Plate 3, Fig. 26). Theca pentagonal in summit view with slightly to moderately concave inter- ambulacra, stem facet relatively large. Basals apparently three, fairly large, slightly convex in profile, occupy about 50% of pelvis. Radials five, long, occupy 50% of pelvis and most of vault, RB axis appears greater than RD axis, body slightly concave in profile, little or no radial lip at tip of ambulacra. Regular deltoids four, relatively short, form spiracular slits on margins with ambulacra, moderately con- cave in cross section. Anal deltoids two, not well preserved, hypodeltoid partly missing but appears to reach spiracular slit on each side, hypodeltoid slightly larger than other deltoid bodies. Ambulacra fair- ly long, only preserved in two or three rays, lanceolate, appear concave in cross section with raised margins against adjacent ra- dials and deltoids and moderately de- pressed centers (Plate 3, Figs. 25-26), giving a wide V-shaped cross section (Text- Fig. 141); lancet partly exposed in center, side plates numerous but not well pre- served. Spiracular slits alongside ambula- cra, apparently sealed aborally, probably extend half of ambulacral length or less, posterior spiracular slits do not quite reach adoral edge of anus. Summit structures poorly preserved. 130 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 Studied Specimen and Occurrence. Discussion. The discovery of a new MCZ 884 from a sequence of thick light Metablastus species in Montana extends beds near the top of the Woodhurst Mem- the range of this genus down to the Kin- ber of the upper Lodgepole Limestone in derhookian. About half the described the pass just south of Baldy Mountain, species have slightly to moderately flared southern Bridger Range, southwestern basals with a triangular stem facet, appar- Montana. ently ancestral to the strongly flared basals Discussion. This specimen with its con- of the genus Tricoelocrinus. The new cave ambulacra appears quite distinct from species of Metablastus from Montana has species of Orophocrinus found in the low- non-flared basals with a round stem facet, er Lodgepole and from other previously and was probably ancestral to other Osa- described Orophocrinus species, none of gean and Meramecian species with the which have flat or slightly concave am- same feature. Metablastus differs from bulacra (see Macurda, 1965, table 2). The other closely related genera such as Troos- thecal shape indicates that this specimen ticrinus, Tricoelocrinus, and Costatoblas- is probably related to species such as tus (see Sprinkle and Gutschick, 1967, p. Orophocrinus orbignyanus, O. conicus, 391) by having four anal deltoids, regular and O. macurdai, n. sp. The concave am- deltoids not visible in side view, lancet bulacra, fairly large stem facet, short spi- completely covered by the side plates, and racular slits, and anal deltoid morphology a steeply conical to biconical theca without separate it from all of these species. If other a strongly inflated base, plus its occurrence specimens are found and confirm the de- in the Mississippian. scribed morphology, this form will even- ..—-.-,. AO-rno un ■ i^amftmoio tually need to be described as a separate METABLASTUS Mil JGANENSIS species. Sprinkle and Gutschick, new species Plate 4, Figures 1-13; Order SPIRACULATA Jaekel, 1918 Text-Figure 16; Table 4 Family TROOSTRICRINIDAE Bather, 1899 Diagnosis. Theca biconical, pelvis Genus METABLASTUS Ethendge and somewhat longer than vault, basals non- Carpenter, 1886 flaring with round stem facet, 2?-3 hy- Type Species. Pentremites lineatus drospires per ambulacral side. Shumard, 1858. Description. Six partly complete speci- Diagnosis. Spiraculate blastoids having mens and two separate radial plates avail- an elongate theca (usually biconical); four able for study; specimens thin-plated and paired spiracles and a paired anispiracle, most damaged during acid extraction; de- anus surrounded by an enlarged hypodel- scription primarily taken from holotype toid, a smaller adoral superdeltoid, and two MCZ 803. Thecal shape nearly biconical hidden cryptodeltoids; 2?-5 hydrospire with slightly expanding conical pelvis folds per ambulacral side; ambulacra nar- and rounded conical vault (Text-Fig. 16A). row, lancet completely covered by side Holotype 11 mm long, 6.6 mm maximum plates, one pore per side plate along both width (crushed), original width estimated radial and deltoid margins; hydrospire to be 5.5-6.0 mm. L/W ratio now 1.7 plate lacking; deltoids small, strongly over- (crushed), original L/W ratio probably 1.8- lapped by radials, not appearing on thecal 2.0. Vault of holotype 4.6 mm long, pelvis plate surface except for enlarged hypo- 6.4 mm long, V/P ratio 0.69, probably un- deltoid; basals sometimes flared with tri- affected by crushing. In two best para- angular stem facet. types, L/W and V/P ratios 1.76 and 0.91, Occurrence. Early to Middle Mississip- plus 1.82 and 0.76, respectively. Basal an- uri, Illinois, Iowa, Indiana, gle in holotype now 55-60°, probably clos- Kentucky, and Montana. er to 50° in original uncrushed specimen. Mississippian Blastoids from Montana • Sprinkle and Gutschich 131 Maximum width at base of ambulacra above midheight. Interradial areas slightly concave; ambulacra slightly convex in cross section, only slightly depressed below sur- face of radials (Text-Fig. 16C). Stem at- tachment round without flaring basals. Basals three, normally arranged in a me- dium-sized cone, making up slightly more than half of pelvis, slightly concave in pro- file. Two larger and one smaller (azygous) basal; latter in AB interray, 4.0 mm long and 2.5 mm wide; larger basals about same length and about 3.5 mm wide in holotype. Stem facet at tip of cone, nearly round, about 0.8 mm in diameter with a tiny cen- tral lumen, secondary deposits very minor around stem facet. Radials five, elongate, making up most of thecal surface. Each radial has nearly parallel lateral sutures, and most limbs ex- tend nearly to a point at their adoral end. In holotype, radials 7.2 mm long, 2.7 mm maximum width, with body 2.8 mm long and limbs 4.6 mm long along each am- bulacrum. Radial body nearly straight in profile, radial limbs slightly curved in pro- file. Regular deltoids four, small, not visible on thecal plate surface, strongly over- lapped by radials. In holotype, deltoids 0.9 mm long in ambulacral sinus, about 0.2 mm wide, with low crest on summit slop- ing down to spiracles and deltoid lip. Ra- diodeltoid suture only slightly raised over ambulacral surface. Spiracles apparently paired, with thin depressed deltoid septum not completely separating spiracles from adjacent ambulacra (Plate 4, Fig. 13). Anal deltoids poorly exposed or missing from all specimens, should be four in num- ber. Enlarged aboral hypodeltoid exposed on the thecal surface, small adoral super- deltoid, and two hidden cryptodeltoids be- neath hypodeltoid. Anispiracle apparently paired also and anus not completely sep- arated from posterior spiracles. Ambulacra five, narrow, elongate. In holotype, ambulacra about 5.5 mm long, about 0.7-0.8 mm wide along much of length. In separate radials, ambulacral si- nus up to 8.0 mm long. Lancet thick, keeled on interior (Text-Fig. 16D), completely covered by side plates, about 10 side plate sets per 3 mm length. Holotype has about 19 side plate sets per ambulacral side, one brachiole facet per side plate set. Sutures between opposing sets of inner side plates in main food groove; small outer side plate notched aboral-abmedial edge of inner side plate (Text-Fig. 16D). One pore per side plate set along radial and short deltoid margins (pores alternate with brachiole facets), short pore furrows indistinct. Side plate sets become smaller (and less oblique) adorally, especially alongside deltoids; side food grooves enter main food groove at 30° angle aborally, nearly 60° angle adorally. Hydrospires poorly known, apparently at least two and more likely three hydro- spires per ambulacral side in paratype MCZ 807. No hydrospire plate present. Ornament on basals and radials consists of closely spaced growth lines paralleling sutures; best exposed on holotype and non- silicified separate radials (Plate 4, Figs. 1- 2 and 10-11). Secondary deposits nearly lacking from stem facet, only small lip at tip of each ambulacrum at radial origin. Growth features poorly known because many specimens incompletely preserved; smallest specimen (holotype MCZ 803) 11 mm long, largest specimen (paratype MCZ 806) an estimated 17 mm long (pelvis mostly missing). Measurements for three most complete specimens in Table 4. Stem and brachioles unknown in present ma- terial. Studied Specimens. Holotype MCZ 803, paratypes MCZ 804-810 (five partial spec- imens and two radial plates). Occurrence. Known from the lower Lodgepole Limestone at five localities in southwestern Montana: holotype from An- telope Valley 39-45 ft (12-14 m) above the base of the Paine Member, three para- types from Milligan Canyon East 12-20 ft (3.7-6 m) above the base, one paratype from Milligan Canyon 18-20 ft (5.5-6 m) above the base, one paratype from London Hills about 40 ft (12 m) above the base, Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 and two radial plates in the float from Northeast Baldy Mountain 20-50 ft (6-15 m) above the base. Etymology. The species is named for Milligan Canyon, southwestern Montana, where four of the six partial specimens were found. Discussion. Metablastus milliganensis is the seventh species to be described for this genus (see Fay, 1961, pp. 77-82). It can be differentiated from several Meta- blastus species because it does not have flared basals and a triangular stem facet. It differs from the type species M. lineatus by being less elongate (much lower L/W ratio), and from M. bipyramidatus and M. varsouviensis by having the pelvis longer than the vault. At present, M. milliganen- sis is the earliest described species and may have been ancestral to several later species of Metablastus, especially M. lineatus which occurs in the Burlington Limestone and is the next oldest species. Family PENTREMITIDAE Orbigny, 1851 Genus MONTANABLASTUS Sprinkle and Gutschick, new genus Type Species. Montanablastus bal- dyensis Sprinkle and Gutschick, new species. Diagnosis. Spiraculate blastoids with an obconical theca, vault usually equal to or slightly longer than pelvis; four spiracles and an anispiracle; two or three hydro- spires per ambulacral side, two anal del- toids, hypodeltoid enlarged; regular del- toids smaller but appearing on side of theca, form low crests above depressed summit, normal V-shaped radiodeltoid sutures with radials abutting deltoids (no overlap); am- bulacra moderately long but fairly narrow, lancet exposed toward adoral end, one pore per side plate along radial and deltoid mar- gins; hydrospire plate apparently absent; plates ornamented with fine growth lines; brachioles about two and a half times the- PLATE4 Figures 1-13. Metablastus milliganensis Sprinkle and Gutschick, n. sp., lower Paine Member, lower Lodgepole Limestone, 1-3 and 12-13 from Antelope Valley, 4-7 from Milligan Canyon, 8 from London Hills, 9 from Milligan Canyon East, and 10-11 from Northeast Baldy Mountain, southwestern Montana. 1-3, 12-13, A-side, CD-side, top, and E- and B-ambulacral views, respec- tively, of relatively small holotype MCZ 803 showing shape of nearly complete but crushed theca and morphology of two well- preserved narrow ambulacra, x2.6 and *6.5; 4-7, B-side, D-side, top, and bottom views of medium-sized paratype MCZ 804; note shape, missing tip of basals, and concave interambulacra, x2.6; 8, side view of large paratype MCZ 805 showing large holes in theca and serpulid (left) attached to radials, x2.6; 9, side view of very large incomplete paratype MCZ 806 still partly in matrix, x2.6; 10-11, paratype radials MCZ 809 and 810 showing elongate but relatively narrow ambulacral sinuses, x2.6. Figures 14-28. Cryptoblastus? sp. A, Woodhurst Member, upper Lodgepole Limestone, Sacagawea Peak, Bridger Range, southwestern Montana. 14-15, C-side and top views of relatively small theca MCZ 1049 showing globular shape and eight closely-set spiracles on summit, x2.7; 16-17, C-side and bottom views of medium-sized theca MCZ 1045; note long ambulacra and concave basal cavity, x 2.7; 18, side view of medium-sized theca USNM 20670 showing elongate shape and rather coarse silicification, *2.7; 19, partial radial and ambulacrum MCZ 1051; note side plates and radial ornament, x2.7; 20, large radial, deltoid, and ambulacrum USNM 20670; note short deltoid at upper right, x2.7; 21-22, B-side and top views of relatively large globular theca MCZ 1 046 showing eight closely-set spiracles on summit, x 2.7; 23-24, C-side and bottom views of large elongate theca MCZ 1047; note long ambulacra and small concave basals, x2.7; 25, interradial side view of large broken theca MCZ 1050 with fine growth lines, *2.7; 26, side view of large broken theca MCZ 1055 in etched slab showing disrupted plates at top of theca from horizontal worm burrow, x2.7; 27, side view of very large broken theca MCZ 1048; note growth lines and long ambulacral sinus, x 2.7 ; 28, top view of eroded theca MCZ 1 053 showing summit features and trace of hydrospires beneath C-ambulacrum, x2.7. Figures 29-30. Cryptoblastus? sp. C, upper Paine Member, middle Lodgepole Limestone, Northeast Baldy Mountain, Bridger Range, southwestern Montana. Side and bottom views of weathered globular theca MCZ 1040 in slab showing eroded slightly convex base, growth lines on radial (30, upper right), and brachioles radiating from all five long ambulacra, x 2.8. Figures 31-34. Cryptoblastus? sp. B, upper Paine Member, middle Lodgepole Limestone, Bandbox Mountain, west-central Montana. 31, surface view of small deltoid MCZ 1041 ; note closely-set spiracles, two large spines in center, and growth lines aborally, x6; 32, edge view of small deltoid MCZ 1042 showing three large spines and trace of hydrospire folds, x6; 33, radial fragment MCZ 1044 with long ambulacral sinus, x6; 34, basal set MCZ 1043 with parts of D and E radials; note stem facet on slightly convex basals, growth lines, and large lip at each radial origin, x6. Mississippian Blastoids from Montana • Sprinkle and Gutschick 133 y-vv ,v ^ m- 6 .■■■■■■■ *r- A-J M on W T5 ^4 134 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 cal length; small-diameter stem having slightly flanged columnals. Occurrence. Early Mississippian (Late Kinderhookian) (=Earliest Carboniferous- Tournaisian), Montana. Etymology. Named for the state of Montana, where this new genus was dis- covered. Discussion. Most of the available spec- imens of this genus are exceptionally well preserved with attached brachioles and stem; unfortunately, as in most blastoid occurrences preserved like this, it is very difficult to identify these specimens and study their thecal morphology. Several specimens with buried appendages were partly silicified, and the appendages were sacrificed to uncover the theca by acid etching. However, this was only partly suc- cessful because of incomplete silicihcation and small size of the specimens. Montana- blastus resembles several other genera in the Familv Pentremitidae, but cannot eas- ily be assigned to any of them. It differs from Hyperoblastus, Conuloblastus, De- vonoblastus, and Eleutheroblastus (all De- vonian genera, see Fay and Wanner, 1968) by apparently having only two anal del- toids plus other differences in thecal shape, deltoids, ambulacra, and the later age. It differs from Early Mississippian genera such as Petaloblastus (see Fay, 1962a) by having much narrower ambulacra with less lancet exposure and V-shaped radiodeltoid sutures, from Pentremoblastus by having narrower ambulacra and only two anal deltoids, and from early species of Pen- tremites by having narrower ambulacra and smaller but crested deltoids. Speci- mens of Montanablastus show consider- able resemblance to Metablastus and Cos- tatoblastus (see Sprinkle and Gutschick, 1967, table 1), but these have paired spi- racles, thus belonging in a different family, plus differently shaped deltoids and thecal ornament. MONTANABLASTUS BALDYENSIS Sprinkle and Gutschick, new species Plate 5, Figures 1-8; Plate 6, Figures 33-43; Text-Figure 17; Table 4 Diagnosis. Theca obconical, vault usu- ally slightly greater than pelvis; fairly nar- row ambulacra, lancet making up only about 25% of ambulacral width; hypodel- toid about one and a half times as long as other deltoid bodies; ornament consists of fine growth lines, large radial prong at tip of ambulacra; columnals slightly flanged. Description. At least 30 specimens avail- able for study, including holotype MCZ 886, 26 paratypes either etched out from the matrix or on slabs with attached ap- pendages, and several other possible spec- imens. Theca obconical, pelvis conical, vault truncated conical to parabolic, sum- PLATE5 Figures 1-8. Montanablastus baldyensis Sprinkle and Gutschick, n. gen., n. sp., slab specimens from middle Paine Member, middle Lodgepole Limestone, Northeast Baldy Mountain, Bridger Range, southwestern Montana. 1, paratype MCZ 898 showing many long, complete, recurved brachioles, long stem with segment missing, and theca tangled up with Y-shaped ramose bryozoan, x2; 2, paratype MCZ 903 with crushed theca, few recurved brachioles, and long stem, x2; 3, paratype MCZ 882 showing many brachioles and deeply-buried stem, x2; 4, paratype MCZ 900 with tightly-recurved long brachioles hiding most of theca, '2; 5, paratype MCZ 899 showing long recurved brachioles attached to edges of ambulacra, x2; 6, paratype MCZ 905 with splayed-out brachioles and deeply-buried stem, x2; 7, paratype MCZ 901 showing short broken brachioles and long stem with recurved tip, x2; 8, paratype MCZ 902 with long brachioles and long, deeply-buried, kinked stem emerging from edge of slab (arrow), x2. Figures 9-11. Strongyloblastus laudoni Sprinkle and Gutschick, n. sp., slab specimens from middle Paine Member, middle odgepole Limestone, 9 from Ant Park, Little Belt Mountains, west-central Montana, 10-11 from Northeast Baldy Mountain, 3ridger Range, southwestern Montana. 9, paratype MORI 001 (Welch Collection) immersed in water showing badly-crushed :overed by long brachioles and long, fairly large stem incomplete distally, x2; 10, paratype MCZ 872 with thecal growth long brachioles attached to edges of ambulacra, and short stem segment, x2; 11, paratype MCZ 873 showing broken thecal base, small but visible deltoids, and brachioles, x 2. Mississippian Blastoids from Montana • Sprinkle and Gutschick 135 ^r / - "N % fa u (ffl ■ r / / j ft .-3- ' • / 9 ■■*?■: 136 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 mit depressed with deltoids projecting above peristome (Text-Fig. 17A). Most specimens fairly small; smallest theca 2.2 mm long, largest theca 10 mm long. L/W ratio ranges from 1.13 to 1.45 and averages 1.32 (10 measurements), gradually in- creasing with size; V/P ratio ranges from 0.7 to 1.36 and averages 1.02 (eight mea- surements), showing considerable varia- tion but no obvious trends; and pelvic an- gle varies from 55° to 75° and averages 69° (eight measurements), gradually increas- ing with size. Maximum width at large radial lips usually just below midheight; interambulacral areas flat to slightly con- vex ignoring radial lips, but somewhat con- cave if radial lips included (Text-Fig. 17B). Basals three, normally arranged, rep- resent about 50% of pelvis, flat to slightly concave in profile, two larger and one smaller (azygous), azygous basal quadrate, larger basals hexagonal. Stem facet slightly triangular with only small secondary de- posits forming platform. Radials five, large, forming about 50% of pelvis and most of vault, RD axis ap- parently greater than RB axis at all sizes, large lip up to 0.9 mm long near origin of radials pointing obliquely outward and continuing pelvis profile, lip probably PLATE 6 Figures 1-20. Strongyloblastus breimeri Sprinkle and Gutschick, n. sp., lower Paine Member, lower Lodgepole Limestone, 1, 3-5, 9-12, 16, and 19 from Milligan Canyon East, 2, 6, 7, 13, and 17 from Milligan Canyon, 8 and 14-15 from South Boulder, 18 from Dry Hollow, and 20 from Saddle Peak, southwestern Montana. 1,10, E-side and top views of very small paratype MCZ 843 showing elongate shape, short ambulacra, and separate spiracles, x2.3; 2, B-side view of small paratype MCZ 844; note vault now longer than pelvis, x2.3; 3, E-side view of small paratype MCZ 847 showing longer ambulacra and growth lines on radials, *2.3; 4, 11, D-side and top views of medium paratype MCZ 849; note separate spiracles and damaged base, x2.3; 5, 12, E-side and bottom views of medium paratype MCZ 851 showing elongate shape and stem facet, x2.3; 6, B-side view of crushed paratype MCZ 853; note deltoid length and growth lines on radial, x2.1 ; 7, C-side view of large paratype MCZ 855 still partly enclosed in matrix, x2.1 ; 8, 14-15, D-side, top, and bottom views of large holotype MCZ 841 showing elongate theca with vault much longer than pelvis, separate spiracles on summit, and secondary deposits around stem facet, 2.1; 9, B-side view of very large paratype MCZ 857; note very long, wide ambulacra and deltoid bodies ending well below summit, x2.1 ; 13, top view of medium paratype MCZ 860 showing separate spiracles and C-spiracle partly cut off from rest of anispiracle, x2.3; 16, AB-side view of medium paratype MCZ 862 with rounded summit, x2.1; 17, C-side view of very large broken paratype MCZ 861 showing enlarged hypodeltoid (top left) and separate spiracles, x2.1; 18, oblique EA-side view of medium paratype MCZ 854 still partly in matrix, note well-preserved ambulacra showing brachiole facets plus lobes and sockets, and epideltoid (top rear) somewhat larger than other deltoid lips, x3; 19, side view of large crushed paratype MCZ 866 missing most of the ambulacra but having well-preserved growth lines on the radials, x2.3; 20, side view of very large crushed paratype MCZ 859 in slab showing long wide ambulacra, visible deltoids, and separate spiracles, x2.1. Figures 21-23. Strongyloblastus sp., lower Paine Member, lower Lodgepole Limestone, Targhee Peak, southeastern Idaho; A-side, top, and bottom views of small but well-preserved theca MCZ 870 showing different shape from 1-3 above, separate spiracles with C-spiracle cut off from rest of anispiracle (22), and growth lines on radials, x2.5. Figures 24-32. Strongyloblastus laudoni Sprinkle and Gutschick, n. sp., upper Paine Member, middle Lodgepole Limestone, Northeast Baldy Mountain, Bridger Range, southwestern Montana. 24-25, oblique EA-side and top views of medium paratype MCZ 874 partly etched from matrix; note wide ambulacra, separate spiracles, silicified brachioles from back ambulacra, and stem emerging from matrix (24, bottom), x3; 26, side view of medium paratype MCZ 875 showing short but visible deltoids and lancet in center of ambulacrum, x2.9; 27, side view of medium paratype MCZ 878; note well-preserved ambulacra with brachiole facets and pore furrows, x3; 28-29, top and D-side views of large silicified but broken holotype MCZ 871 showing separate spiracles, thin septum barely cutting off C spiracle from rest of anispiracle, wide ambulacra with brachiole bases still attached to some facets, and cover plates over proximal ambulacra and mouth, x3; 30, two wide lancet plates (paratype MCZ 881) on a slab, x3; 31, radial plate (paratype MCZ 879) with wide sinus and well-developed growth lines, x3; 32, lancet with partial side plates (paratype MCZ 880) showing lancet width and brachiole facets (left), x3. Figures 33-43. Montanablastus baldyensis Sprinkle and Gutschick, n. gen., n. sp., upper Paine Member, middle Lodgepole Limestone, Northeast Baldy Mountain, Bridger Range, southwestern Montana. 33, very small weathered paratype MCZ 909 in slab; note attached stem segment and two adjacent brachioles, x3; 34, side view of small silicified paratype MCZ 891 showing exposed plate sutures, x3; 35, 37, top and C-side views of medium silicified paratype MCZ 889; note four spiracles plus anispiracle (35), plate sutures, and large radial lips, x3, 36, 39, top and A?-side views of medium paratype MCZ 888 showing deltoids, traces of oral cover plates, and fairly narrow ambulacra, x3; 38, top view of medium silicified paratype MCZ 887; note ■ spiracles and epideltoid bordering anispiracle, x 3; 40-41 , top and side views of medium silicified paratype MCZ 893 in slab showing thecal shape and many brachiole segments, x3; 42-43, top and E-side views of fairly large silicified holotype MCZ 886 showing thecal shape, relatively narrow ambulacra, and large radial lips, x3. Mississippian Blastoids from Montana • Sprinkle and Gutschick 137 -v i j\] T f:\ tp >' ivJ 6 ""<#/ wH Ml r, &3 9 '4/5 i x 21 0. W t^rffe '^ ****** '^J gM %f 34 SfiS 35 V' 138 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 SPL BRF -FL BCP -fl G * H Text-Figure 17. Morphology of Montanablastus baldyensis, n. gen., n. sp. A-B, side and summit views of large theca based on paratype MCZ 889 and holotype MCZ 886; note shape, location of maximum width just below midheight (short lines), and summit features. C, much-enlarged plan view of ambulacrum in paratype MCZ 896 showing lancet (L) exposed in center, large inner and small wedge-shaped outer side plates (ISP and OSP) together supporting a hemi-elliptical brachiolar facet (BRF), pore furrow (PF) curving around lower edge of facet, and cover plate lobes and sockets. D-E, much-enlarged plan and side views of spiracle (SP) and mouth (M) on depressed summit with adjacent raised deltoid (D) and adoral ambulacra; based mostly on paratypes MCZ 886 and 887. F, enlarged cross section through adoral theca in paratype MCZ 895 showing ambulacrum made up of lancet (L) and side plates (SPL), adjacent radial limbs (R) with raised edges, and apparently three, poorly preserved, silicified hydrospire folds (HF) on each side, x8.6. G, proximal stem in paratype MCZ 898 showing columnal shape and thin flange (FL) around center; arrow points to attachment at base of theca, x17.2. H, side view and cross section of brachiole in paratypes MCZ 898 and 905; note striations on side and low biserial set of cover plates (BCP) over relatively deep, V-shaped, food groove; arrow points to attachment on ambulacrum, x21.5. formed from secondary deposits, but one theca has lip with closely spaced growth lines. Regular deltoids four, small, body tri- angular, extends only short distance down thecal surface but visible in side view, V-shaped suture between radials and del- toid forming 80-85° angle, deltoids slightly concave in profile, radials abut deltoids without obvious overlap. In several thecae, deltoids 0.8-0.9 mm long, projecting above nearly flat summit (Plate 6, Figs. 36-39), sharp adoral edge of deltoids dropping away rapidly to below summit level; four spiracles formed in front of projecting del- toid bodies by edges of ambulacra and curved deltoid lips, spiracles teardrop- shaped, small, with depressed deltoid sep- tum not reaching surface (Text-Figs. 17D- E). Anal deltoids apparently two, somewhat enlarged hypodeltoid aborally, 1.2-1.5 mm long (about one and a half times length of regular deltoids), slightly ridged with stronger growth lines; epideltoid small, separates mouth from anispiracle, de- pressed aboral side has three troughs for central anus and two lateral hydrospire groups (Plate 6, Fig. 38), septa separating these troughs depressed to form true ani- spiracle, no evidence of cryptodeltoids. Ambulacra five, moderately long, fairly narrow, slightly convex in cross section, lancet partly exposed in center of adoral half, forming about 25% of ambulacral width, side plates on bevelled abmedial edges of lancet, appear to be normally ar- ranged but not well preserved or exposed in most specimens, 14 side plate sets pres- ent in one ambulacrum 4.0 mm long, side food grooves meet main food groove at 45-60° angle, inner side plates apparently large, rectangular, outer side plates not ob- vious but probably small triangular wedges underlying half of brachiole facets, bra- chiole facets large, at slight angle to side Mississippian Blastoids from Montana • Sprinkle and Gutschick 139 food grooves, occupy about half of am- Etymology. Named for the Northeast bulacral width (Text-Fig. 17C). Main food Baldy Mountain locality where all the groove has 4-5 lobes between each pair of studied specimens were collected, side food grooves, which have about three Discussion. Montanablastus baldyensis smaller lobes on the adoral and aboral sides, is an unusual blastoid for the Early Mis- Hydrospires in 10 groups, apparently 2- sissippian. The fairly narrow ambulacra 3 folds per ambulacral side based on two are not particularly similar to other forms poorly preserved silicified specimens and in the Family Pentremitidae; even species a sectioned slab specimen (Text-Fig. 17F). such as Pentremites conoideus have wider Ornament consists of fine growth lines ambulacra with more lancet exposure. The on basals and radials, somewhat coarser radial lips are very large for a blastoid, growth lines on deltoids, hypodeltoid and continue the profile from the pelvis, and radial-hypodeltoid growth front. the lip on one specimen shows apparent Nearly two-thirds of specimens have growth lines, indicating that the lips were brachioles preserved and nearly half have produced by periodic small increments of proximal stem attached; complete bra- growth and not by secondary deposits, chioles about 15-16 mm long in 6 mm long Many specimens with appendages are ex- theca (Plate 5, Fig. 1) with rounded tri- cellently preserved and were cleaned with angular cross section, biserial brachiolar an air abrasive unit, but are almost useless plates (BP), and one low set of tiny biserial for trying to work out the thecal mor- eover plates (BCP) often pyritized along phology of this species. More and better with wide V-shaped brachiolar food groove silicified materials will be necessary before (Text-Fig. 17H). Brachiolar plates about more complete information can be ob- 0.2 mm long, 0.25 mm wide, with a very tained. small central ridge and fine striations extending down length (Text-Fig. 17H); Fami|y PENTREMITIDAE? Orbigny, 1851 about two BCP/BP where observable. Genus STRONGYLOBLASTUS Fay, 1962b Stems incomplete, ranging up to 29 mm Type Species. Strongyloblastus petalus long (attached to theca 5.5 mm long; Plate Fay, 1962b. 5, Fig. 7); this stem has about 190 colum- Diagnosis. Spiraculate blastoids with an nals in this length. Columnals having ovoid, ellipsoidal, or obconical theca; eight rounded edges and small equatorial flange, divided spiracles plus variable arrange- a tiny central lumen, and averaging 0.14- ment on anal side (anispiracle, "C" spi- 0.15 mm long, 0.5 mm wide proximally, racle plus half-paired anispiracle, or "C" and 0.3 mm (or 0.35 mm with flanges) and "D" spiracles plus anus); 3-5 hydro- wide distally (Text-Fig. 17G). spire folds per ambulacral side; two anal Growth information for the few mea- deltoids, prominent epideltoid often larger surable specimens summarized in Table 4. and higher than other deltoid lips, and hy- Studied Specimens. Holotype MCZ 886, podeltoid with slightly enlarged body and paratypes MCZ 882, 887-913, additional adorally projecting septum more promi- specimens MCZ 914. nent than those of other deltoids; regular Occurrence. All described specimens deltoids appear on side of theca, deltoid come from the Northeast Baldy Mountain body short to moderately long, sometimes locality in the southern Bridger Range of heavily ridged, radials overlap deltoids; southwestern Montana. Specimens occur ambulacra medium to long, very wide, in series of beds with 6-8 in. (0.15-0.20 lancet completely exposed, occupying 40- m) of limestone interbedded with shaly 60% of ambulacral width, one pore per dolomite from the middle Paine Member side plate along radials, pores either pres- between 150 and 175 ft (46-53 m) above ent along deltoids or closed off short dis- the base of this member. tance above radiodeltoid suture, no hy- 140 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 drospire plate present; ornament consisting of fine to coarse growth lines. Occurrence. Early to Middle Mississip- pian (Early Carboniferous), southern Ca- nadian Rockies, northern U.S. Rockies, Mississippi Valley, and Alaska. Discussion. Two new species of Stron- gyloblastus occur in the Lodgepole Lime- stone in southwestern Montana, one in the lower Paine Member and the second in the upper Paine Member. A single additional specimen from the lower Paine Member is unassigned at present. A third new species belonging either to Strongyloblastus or to Pentremites (as presently defined) occurs in the younger Castle Reef Dolomite in the Sun River Canyon area of northwest- ern Montana; this form will be described in a separate paper. Strongyloblastus was described by Fay (1962b) as a Devonian blastoid from west- ern New York State, based on the label that accompanied the holotype specimen. Macurda and Breimer (1977, p. 693) re- ported that "Strongyloblastus was com- pletely anomalous when compared with other Devonian Mastoids," and that similar specimens occur in the Banff Formation of Early Mississippian (probably Late Kin- derhookian) age in the southern Canadian Rockies, and concluded that the label with the holotype was incorrect. Occurrences of similar blastoids belonging to different species are known from the northern U.S. Rockies (see following) and from Alaska. Fay (1964) assigned Strongyloblastus to the Family Schizoblastidae, but Macurda and Breimer (1977, p. 694) assigned it to the Family Pentremitidae after some dis- cussion because of its overall resemblance to several early species of Pentremites, al- though this family then cannot be char- acterized alone by having four spiracles and an undivided anispiracle. Strongyloblastus is most closely related to several early species of Pentremites with divided spiracles, such as P. elongatus and P. kirki (Macurda, 1975; Horowitz, Waters, and Macurda, 1981; Horowitz, Macurda, and Waters, 1986). Strongyloblastus dif- fers only slightly from Pentremites species having divided spiracles (see Macurda and Breimer, 1977, p. 696) by having non- functional ambulacral pores along most of the deltoid margin and higher septa sep- arating the anus from one or both of the posterior spiracles. These differences are very minor and intermediate stages occur in some of the new species described here, in contrast to the difference between di- vided and undivided spiracles, a differ- ence that previously would have placed these blastoids in different families (see Fay, 1964; Fay and Wanner, 1968). Instead of having these two genera sep- arated by such minor differences at pres- ent, we propose that all species with di- vided spiracles now assigned to or inferred to belong to Pentremites (P. elongatus, P. kirki, and one or more undescribed species from Montana, western Canada, and Alas- ka) be assigned to the genus Strongylob- lastus Fay (1962b). This proposed change would restrict Pentremites to species hav- ing undivided spiracles (like its type species P. godoni), remove two named and de- scribed species from the genus, and restrict its range to Middle Mississippian (Mer- amecian) to Early Pennsylvanian (Mor- rowan). Strongyloblastus would range throughout much of the Early Mississip- pian from the late Kinderhookian in the Rocky Mountains (Lodgepole Limestone, Banff Formation) to at least the late Osa- gean in the Midwest (Burlington Lime- stone) and perhaps higher in the Rocky Mountains. Strongyloblastus may have been ancestral to Pentremites by suppres- sion of the deltoid septa in the regular spi- racles and anispiracle, retention of func- tional pores along the deltoid margins, and perhaps enlargement of the deltoid body. STRONGYLOBLASTUS BREIMERI Sprinkle and Gutschick, new species Plate 1, Figures 4-5; Plate 6, Figures 1-20; Text-Figures 18A-E and 19 Diagnosis. Theca changing from obcon- ical to elongate ellipsoidal during growth, L/W ratio averages 1.61, V/P ratio av- Mississippian Blastoids from Montana • Sprinkle and Gutschick 141 Text-Figure 18. Morphology of Strongyloblastus breimeri, n. sp. (A-E), S. laudoni, n. sp. (F-l), and S. sp. (J-K). A-C, enlarged side views of large paratype theca MCZ 857 and small paratype theca MCZ 843 and summit view of large theca showing considerable change in shape between biconical juvenile and ellipsoidal adult, very long and wide ambulacra, separate spiracles and half-paired anispiracle on summit, and short lines at maximum width. D, much-enlarged deltoid body (D), thin deltoid septum (DS) separating elliptical spiracles (SP), and deltoid lip (DL) separating spiracles from mouth in paratype MCZ 854, x8.6. E, plan view of ambulacrum in MCZ 854 showing wide exposure of lancet (L) in center, large inner and small triangular outer side plates (ISP and OSP) supporting a brachiole facet (BRF) at the ambulacral edge, curved pore furrow (PF) extending from pore (P) toward center of ambulacrum, and numerous cover plate lobes and sockets, x14.6. F, side view of paratype theca MCZ 875 showing different shape and shorter ambulacra from A above (maximum width at short lines), x 2.1 . G, proximal and medial stem in MORI 001 ; note enlarged proximal stem with thin columnals just below attachment and lack of flanges on columnals, x 3.4. H, plan view of ambulacrum in MCZ 878 showing lancet (L) exposed in center, inner and outer side plates (ISP and OSP), brachiole facets (BRF) at ends of side food grooves, and somewhat longer pore furrows (PF) just reaching lancet, x3.9. 1, much- enlarged side view and cross section of brachiole in MCZ 872; note smooth brachiolar plates and cover plates over shallow food groove, x12. J, side view of small theca MCZ 870 showing difference in shape from B above (short lines at maximum width), x 3.2. K, enlargement of anal side in MCZ 870 showing epideltoid (ED) septum cutting off C spiracle from rest of anispiracle and reaching hypodeltoid (HD), x7.6. erages 2.29, pelvic angle averages 82°, in- terambulacra slightly concave; ambulacra long, moderately to strongly convex, lancet fully exposed, occupying about 50% of am- bulacra! width; one pore per side plate set along radials, pores absent just above ra- diodeltoid suture because of ridges on edge of deltoid; eight spiracles and a half-paired anispiracle on summit, one epideltoid limb partly separates "C" spiracle, "D" spiracle and anus not separated; deltoids fairly long but body on thecal surface short except for enlarged hypodeltoid, surface of deltoids not ridged; three hydrospire folds per am- bulacral side. Description. Forty-seven specimens and fragments available for study; description based on holotype MCZ 841, 17 additional nearly complete paratypes in growth se- ries, and 10 other specimens and frag- ments. Theca obconical in small specimens changing to ellipsoidal in medium to large 142 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 ones (Text-Figs. 18A-B); in adults, theca spiracles aborally; septum long, about 0.4 made up mostly of long parabolic vault mm wide, relatively sharp, separates spi- with a short conical pelvis. Growth series racles and adjacent ambulacra, grooved on specimens range from 3.9 mm long to 22.9 edges facing spiracles; deltoid body tri- mm long, holotype a large, slightly com- angular, flat to slightly concave, nearly pressed theca 18.6 mm long (Plate 6, Figs, smooth on surface, edges grooved at each 8, 14-15). In 18-specimen growth series, brachiole facet on ambulacrum (Text-Fig. L/W ratio ranges from 1.30 to 2.24 and 18D), with a slightly M-shaped DR suture averages 1.61, increasing gradually above making an angle ranging from 60° to 140° a length of 15 mm; V/P ratio ranges from in different specimens; radials strongly 0.96 to 3.88 and averages 2.29, increasing overlap deltoids at surface but suture be- dramatically throughout growth; and pel- comes nearly vertical at level of ambula- vic angle ranges from 65° to 95° and av- era. In paratype MCZ 854 (Plate 6, Fig. erages 82°, increasing in small specimens 18), total deltoid length 4.2 mm with body up to about 9 mm long. Greatest width 1.6 mm long and 0.7 mm wide, septum near midheight in large specimens, at or about 1.8 mm long and 0.35 mm wide just above tips of ambulacra in small between spiracles, and lip 0.8 mm long and thecae (Text-Figs. 18A-B). Pelvis profile 1.0 mm wide. slightly concave, interambulacra slightly Anal deltoids two, slightly enlarged epi- concave near midheight, ambulacra mod- deltoid adorally and enlarged hypodeltoid erately to strongly convex all along length, aborally. Epideltoid triangular to inverted edges slightly depressed below adjacent U-shaped, slightly wider than other del- thecal plate surfaces. toid lips, extends higher above mouth, Basals three, normally arranged, form- notched aborally by "C" spiracle (cut off ing about half of pelvis; two larger, one by low epideltoid septum) and half-paired smaller (azygous), azygous basal quadrate, anispiracle (Plate 1, Fig. 5). Hypodeltoid 2.4 mm long and 1.9 mm wide in medium- enlarged over other deltoids, body extends sized specimen; larger basals hexagonal, about 1 mm further down theca in several about same length, approximately 2.9 mm large specimens, nearly 1.6 times size of wide (somewhat distorted); stem facet rel- other deltoids in one fragment (4.2 vs. 2.6 atively large, up to 1.7 mm in diameter mm; Plate 6, Fig. 17), hypodeltoid septum with slight ridge around periphery and wider and higher than other deltoid septa, small central lumen about 0.2 mm in di- extending up to form raised hood over ab- ameter, facet on raised platform of sec- oral edge of half-paired anispiracle (Plate ondary deposits covering origin of basals. 1, Fig. 5), right edge meets raised septum Radials five, very long, making up most from epideltoid cutting off "C" spiracle, of thecal surface, RD axis slightly greater Half-paired anispiracle slightly asymmet- than RB axis in smallest specimens, many ric, elliptical, slightly larger than mouth; times greater in largest ones (Text-Fig. 19), "C" spiracle elongate, slightly smaller than RD and RHD fronts nearly straight, radials other spiracles. overlap deltoids along short suture, radial Ambulacra five, long, very wide, strong- limbs raised along ambulacral margins ly convex aborally to moderately convex (slight secondary deposits on surface) but adorally, about 15 mm long in holotype, edges not grooved here, small wide lip at about 2.5 mm wide at widest point; lancet radial origin pointing laterally. completely exposed in center, occupies Regular deltoids four, body short and about 50% of ambulacral width, fairly thin narrow, above level of adjacent ambula- in cross section; side plates on bevelled edge era, septum and lip long and narrow, at of lancet, inner side plates medium-sized, or just above level of ambulacra (Text-Fig. wide, nearly rectangular, outer side plate 18D). Deltoid lip triangular, notched by small, triangular, on aboral-abmedial edge Mississippian Blastoids from Montana • Sprinkle and Gutschick 143 20 16 E E 12 ? 8 Q) / ft 4 8 12 Width (mm) 20 r 16 12 re 8 16 • % 0» / 0 4 8 12 Pelvis (mm) 15 12 £9 t 6 < / / / • • ST. / 10 20 30 40 No. Side Plates 15 I" • CO g CC o E 6 D £ 3 a HXJ ^r.v *- * 7r 6 ~ 5 E £4 _: 3 °2 (9 — o' 3 6 9 12 15 Growth Front (mm) 4 E 3 E DC 2 CO « ■J 0 / » o °° -<3»o O •o / O O 012340 3 6 9 12 Max. Del. W. (mm) RDo RB* (mm) 15 12 !9 Q DC 6 o* cm I & II E E. cc CC / E E CC CO o Q CC 15 /. 12 0 0 0 9 9 / 0 0 6 f 3 7 , , , 2 4 6 8 RRo RB« (mm) 3 6 RB (mm) 0 2 4 6 Del. L. (mm) Text-Figure 19. Growth plots for 1 9 measured specimens (MCZ 841-859) of Strongyloblastus breimeri, n. sp., plus single small specimen (MCZ 870) of S. sp. (white or clotted diamond). Note large size range and nearly equal RR and RB growth vs. much faster RD growth. Best-fit lines in all plots were hand fit. 144 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 of inner side plates, each side plate set forms one brachiole facet at edge of ambulacrum (Text-Fig. 18E). Main food groove extending down center of each am- bulacrum, between side food grooves hav- ing about four lobes on each side formed by lancet; side food grooves long, empty into main food groove at 50-85° angle, each has 5-6 lobes on adoral side, 3-4 cryp- tic lobes aborally (formed by lancet and inner side plate), large brachiolar pit at end of side food groove; brachiole facets well developed, elliptical, tilted about 30- 40° to side food groove, closely spaced, made up of two shallow depressions. One pore per side plate set along radials, alter- nating with brachiole facets along smooth radial edge, short to medium-length pore furrow extending in from pore between brachiole facets almost to lancet; pores ap- parently absent from deltoid margin just above radiodeltoid suture because deltoid edge grooved (at each brachiole facet) and vertical deltoid ridge between grooves ex- tends into apparent pore position at edge of ambulacrum. Hydrospire groups 10, extend short dis- tance into coelomic cavity from ambula- crum sides, three hydrospires per group (poorly preserved in three specimens), short slit and enlarged tube at bottom of each hydrospire, no hydrospire plate present. Ornament consists of medium to strong growth lines parallel to plate sutures on basals and radials, fine growth lines on del- toids (Plate 6, Figs. 19-20), chevrons on radial limbs below enlarged hypodeltoid somewhat coarser than other growth lines, deltoids not coarsely ridged so far as known. Measurements of specimens in growth series plotted in Text-Figure 19. Stem, brachioles, and cover plates un- known. Studied Specimens. Holotype MCZ 841, paratypes MCZ 842-868 (27 specimens and fragments), and MCZ 869 (19 additional partial specimens and fragments). Occurrence. Known from the lower Lodgepole Limestone at six localities in southwestern Montana; 21 specimens from Milligan Canyon East 12-20 ft (3.7-6 m) above the base of the Paine Member, 20 specimens and fragments from Milligan Canyon 20-25 ft (6-7.5 m) above the base, the holotype and one other specimen from South Boulder Canyon in the float from beds about 40 ft (12 m) above the base, and single specimens from Dry Hollow 30- 35 ft (9-11 m) above the base, from the talus piles at Standard Creek from beds 15-54 ft (4.5-16.5 m) above the base, and from Saddle Peak 55-60 ft (17-18 m) above the base. Etymology. Named for Albert Breimer, State Museum of Geology and Mineralogy, Leiden, Netherlands, one of the authors who restudied the type species Strongy- loblastus petalus and corrected its age and occurrence and re-evaluated its phyloge- netic position. Discussion. Strongyloblastus breimeri is somewhat intermediate in its morphol- ogy between S. petalus, the type species, and "Pentremites" elongatus, perhaps an argument for assigning all three of these species to the same genus. It resembles S. petalus by having a half-paired anispiracle (see below), the epideltoid and hypodel- toid raised and somewhat enlarged, by having the pores closed off along much of the deltoid margin, by having smooth ra- dial edges vs. ridged deltoid edges, and by being almost the same age (Late Kinder- hookian). It resembles "P." elongatus in its general theca shape, by having three hydrospire folds per ambulacral side, by having nearly smooth deltoid bodies, and by having the hypodeltoid somewhat en- larged over the regular deltoids. It differs from both of these species by having a somewhat different thecal shape, much shorter deltoid bodies, and the "C" spi- racle just barely separated from the half- paired anispiracle. An excellently preserved vault of S. pet- alus from the Spreng Collection, Univer- sity of Missouri, Rolla (UMR 6967; Plate 1, Figs. 6-7), shows the anispiracle and summit better than any of the specimens figured by Macurda and Breimer (1977, Mississippian Blastoids from Montana • Sprinkle and Gutschick 145 plate 1). It definitely has a half-paired ani- spiracle similar to but more strongly de- veloped than in S. breimeri, with the epi- deltoid limb separating the "C" spiracle much wider and somewhat higher (Plate 1, Fig. 7). Apparently the epideltoid limbs have variable development in S. petalus, with some specimens having a half-paired anispiracle and others having an anus and two separate spiracles on the posterior side. In the Spreng Collection specimen of S. petalus, the epideltoid is horseshoe-shaped, much larger and higher than the regular deltoid lips, and produces a mouth that is much wider than high, plus four ambu- lacra ("B"-"E") that are curved on the summit (Plate 1, Fig. 7), both rather un- usual features for a blastoid. The critical specimen of S. breimeri for showing the half-paired anispiracle was taken out of the acetic acid bath at the Indiana University Field Station in south- western Montana just after the summit and adoral ambulacra were exposed (see Plate 1, Fig. 5), and brought back to Harvard University to be photographed. After being coated with Glyptal, it was put in acid again to extract the rest of the specimen from the matrix; unfortunately, nearly all the earlier summit detail was destroyed by later acid etching (see Plate 6, Fig. 11). Later specimens from the Milligan Can- yon localities showing excellent detail dur- ing acid etching were removed perma- nently, even if incompletely exposed. STRONGYLOBLASTUS LAUDONI Sprinkle and Gutschick, new species Plate 1 , Figure 3; Plate 5, Figures 9-1 1 ; Plate 6, Figures 24-29; Text-Figures 1 8F-I and 20 Diagnosis. Theca obconical, L/W ratio averages 1.46, V/P ratio averages 1.25, pelvic angle averages 73°, interambulacra flat to slightly convex; ambulacra moder- ately long, very wide, slightly to moder- ately convex, lancet fully exposed, occu- pying 40-50% of ambulacral width; one pore per side plate along radials, pores ap- parently filled in just above radiodeltoid suture; deltoid body relatively short, crest long, usually depressed below edges of ad- jacent ambulacra, eight spiracles and a half- paired anispiracle on summit, "C" spira- cles barely split off from anispiracle, hy- podeltoid not enlarged; three? hydrospire folds per ambulacral side. Description. Approximately nine spec- imens and 14 plates available for study, including partly complete, etched holotype MCZ 871, four paratypes in slabs with bra- chioles and (in three cases) stems pre- served, four paratypes in slabs without ap- pendages, and three separate plate paratypes. Theca obconical, changing only slightly in shape during growth through preserved size range, made up of moder- ately long parabolic vault and moderately short conical pelvis (Text-Fig. 18F). Com- plete specimens range from about 5.0 mm long to about 14.5 mm long. Holotype a large, well-preserved, incomplete theca etched from slab; as preserved, about 12.0 mm long with complete vault 9.8 mm long and upper part of broken pelvis; original length estimated at 15-16 mm (Plate 6, Fig. 29). Greatest width 9.5 mm just above radial lips, just below apparent midpoint of theca. Interambulacra here flat to slight- ly convex. In six measurable specimens, L/W ratio ranges from 1.09 to 1.86, av- eraging 1.46; V/P ratio ranges from 1.25 to 2.11 and averages 1.56; and pelvic angle ranges from 60° to 80°, averaging 73°. Pel- vis profile slightly concave, ambulacra slightly to moderately convex. Basals three, normally arranged, incom- pletely exposed (or missing) in most spec- imens, forming about 50% of pelvis, two larger, one smaller (azygous), regular bas- als hexagonal, about 3.6 mm long and at least 3.5 mm wide in a large specimen; stem facet in this specimen about 1.4 mm in diameter with secondary deposits form- ing rounded to slightly triangular platform for stem attachment. Radials five, long, making up most of thecal surface, RD axis greater than RB axis in all available specimens, many times greater in large specimens (Text-Fig. 20), 146 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 RD front nearly straight, relatively short; plate set forming one brachiole facet at small wide lip at radial origin pointing edge of ambulacrum (Text-Fig. 18H). laterally. Main food groove extending down cen- Regular deltoids four, body short to me- ter of each ambulacrum, between side food dium in length, fairly narrow in most spec- grooves having about 3-5 lobes on each imens, septum and lip long and narrow, side formed by lancet; side food grooves in some specimens septum below level of long, empty into main food groove at 45° ambulacra so that side plates from adja- angle aborally to 80° angle adorally, each cent ambulacra in contact (Plate 6, Fig. having 8-11 lobes on adoral side, 7-9 28). Deltoid body 2.5-3.3 mm long, 1.0- smaller lobes on aboral side, both formed 1.8 mm wide in large specimens, much by lancet and inner side plates, each side smaller in small specimens; septum and lip food groove leading to large brachiolar pit at least 2.5 mm long, just barely splitting about 0.3 mm from edge of ambulacrum spiracles at surface of summit, septum and two shallow depressions making up sharp-crested, part above surface of am- brachiole facet about 0.4 mm long and bulacrum somewhat ridged, radiodeltoid 0.25 mm wide, turned at 20-30° angle to suture makes angle between 90° and 120° side food groove, and slanted abmedially in different specimens, radials moderately and adorally (Text-Fig. 18H; Plate 1, Fig. overlap deltoids with about 60° angle from 3); edge of radial smooth, edge of deltoid plate surface. body somewhat ridged, one pore per side Anal deltoids two, hypodeltoid appar- plate along radials, alternating with bra- ently not enlarged over other deltoids. Epi- chiole facets; pores apparently absent along deltoid triangular, slightly wider than oth- deltoid margin just above radiodeltoid su- er deltoid lips, sends thin septum aborally ture, because deltoid edge grooved at each to cut off "C" spiracle from rest of half- brachiole facet and ridges between grooves paired anispiracle, anus and "D" spiracle extend into apparent pore positions at edge apparently not separated (Plate 6, Fig. 28). of ambulacrum. Pore furrows well devel- Hypodeltoid relatively long, body similar oped, arcuate, extending around lower to other deltoids but septum higher and edge of braciole facet, then laterally along not depressed below adjacent ambulacra, raised center of inner side plate and often may project slightly at aboral edge of ani- reaching lancet, pore furrows present along spiracle, which is elliptical except for both radial and deltoid margins, slightly flattened side along "C" spiracle Hydrospire groups 10, poorly known, septum. "C" spiracle similar in size and apparently three hydrospires per group shape to other spiracles. below each ambulacral side. Ambulacra five, moderately long, very Measurements for few complete speci- wide, slightly to moderately convex in cross mens without appendages plotted in Text- section, in holotype, about 10.5 mm long, Figure 20. 3.3. mm maximum width; lancet com- Ornament consists of fine to medium pletely exposed in center, occupies be- growth lines on basals and radials (Plate 1, tween 40 and 50% of ambulacral width, Fig. 3), somewhat coarser growth lines on about 0.3-0.5 mm thick in several ambu- deltoids and along RHD growth front, lacral fragments, side plates abutting edge Brachioles preserved in at least four of lancet, suture nearly vertical every- specimens, at least 21 mm long in largest where except near aboral end of ambu- example, tightly packed along edge of ara- lacrum where lancet somewhat bevelled, bulacrum where attached (Plate 5, Figs, inner side plates medium-sized, wide, 10-11), apparently expanding in size away nearly rectangular, outer side plates small, from theca for some distance before be- triangular, located on aboral-ab- coming smaller again; in theca about 15 of inner side plates, each side mm long, brachioles from middle of am- Mississippi an Blastoids from Montana • Sprinkle and Gutschick 147 20 16 E E 12 ? 8 CD / / 20 16 12 co 8 4 8 12 16 Width (mm) 0 4 8 12 Pelvis (mm) 15 12 E E t 6 < 3- .* 10 20 30 No. Side Plates "40 15 (lulu) • 1 CO 9 CC ■1 o □ Q 3 a d? / 3 J • / • • / o^. __ o 0 c / • _- — oo ) 3 6 9 12 15 7r 6 ^5 E £4 Q • • E 3 E CO / / Growth Front (mm) 0 1 2 3 4 0 Max. Del. W. (mm) 3 6 9 RDo RB« (mm) 12 15 • 12 E q . / b / / *«■— ■* / y/ 6 3 °c O °o • • ) 2 4 6 8 RRo RB« (mm) 6 - E E cr CC 3 6 RB (mm) 15 12 E E cc m o Q CC 9 - •♦ 2 4 6 Del. L. (mm) Text-Figure 20. Growth plots for six measured specimens (MCZ 871-876) of Strongyloblastus laudoni, n. sp. Because many specimens were incomplete, only 3^4 specimens could be plotted in some cases; short lines through some measurements indicate estimated values in broken specimens. Best-fit lines in all plots were hand fit. 148 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 bulacrum biserially-plated, about 18-19 Discussion. Strongyloblastus laudoni mm long, 0.25 mm wide, and 0.3 mm deep, retains its juvenile obconical shape with a apparently a single set of distally slightly- shorter vault and ambulacra and a lower imbricate brachiolar cover plates present, pelvic angle into the adult stage instead of about 3.3 cover plates per brachiolar plate becoming elongate ellipsoidal as S. brei- on each side, cover plates slightly arched meri does. Other minor differences in- over brachiolar food groove (Text-Fig. 181). elude the hypodeltoid apparently not being In holotype, tiny cover plates scattered over enlarged in S. laudoni, and the lancet oc- ambulacra and still organized into domed cupying somewhat less of the ambulacral structure over adoral food grooves and width; however, other features of the ara- central mouth (Plate 6, Figs. 28-29); many bulacra, pore development, half-paired basal brachiolar plates still attached to anispiracle, and anal deltoids are very sim- edges of ambulacra in this specimen. Stem ilar. This species is also similar to S. kirki preserved in three paratypes; in theca about (see Strongyloblastus Discussion) but is less 15 mm long, 4.5 mm of proximal stem elongate with a lower L/W ratio and a preserved, decreasing from 1.5 mm at the- higher pelvic angle, has a smaller stem ca to 0.8 mm at preserved distal tip, prox- facet, no pores along most of the deltoid imal columnals thin, expanding to basal because the deltoid edge is ridged, and attachment, about 5 per mm, distal colum- may show other differences in the anispi- nals thicker, about 3 per mm (Text-Fig. racle and anal deltoids. Studied Specimens. Holotype MCZ 871, STP(JN?Yi0Bl^SIi,S SP> paratvpes MCZ 872-881 (10 specimens and l}atfX' Fl9s-ZJr 77' plates), and MORI 001 (Welch Collection, Text-Figures 18J-K and 19 Museum of the Rockies, Montana State A single small specimen from the lower University, Bozeman); 11 additional plates Lodgepole Limestone apparently does not in MCZ 883. belong to either of the named species of Occurrence. Known from two localities Strongyloblastus known from this for- in the Bridger Range, southwestern Mon- mation. This specimen (MCZ 870) is im- tana, and one locality in the Little Belt mature (5.5 mm long), but is well pre- Mountains, west-central Montana: all ex- served and not like similar-sized specimens cept one MCZ specimen from Northeast of S. breimeri or S. laudoni in its shape. Baldy Mountain in the southern Bridger It is briefly described here but not named. Range (21 specimens and plates, including Description. Specimen godoniform in the holotype) from two 6-8 in. (0.15-0.20 shape (Text-Fig. 18J), relatively wide, in- m) beds in the middle Paine Member about terambulacra flat to slightly concave; theca 150-175 ft (46-53 m) above the base of 5.5 mm long, 5.2 mm wide, L/W ratio = this member, single ambulacrum from the 1.1; vault widely parabolic, slightly re- float about 115 ft (35 m) above the base curved, 3.5 mm long, pelvis low conical, at Baldy Mountain just to the south, and slightly concave in profile, 2.0 mm long, a single complete specimen from an un- V/P ratio = 1.8; pelvic angle 100-105°, known Lodgepole horizon at Ant Park in much higher than either of the other the Little Belt Mountains (Welch Collec- Lodgepole species at this size. Basals three, tion). relatively large, make up 50-60% of pelvis, Etymology. Named for Lowell R. Lau- slightly bulbous with small stem facet; ra- don, formerly at the University of Wis- dials five, large, fairly long, raised above consin, Madison, who first discovered com- ambulacra; regular deltoids four, very plete blastoids in the Bridger Range and short, but just appearing on side of theca; directed us to the rich Northeast Baldy ambulacra moderately long but without Mountain locality in 1966. many side plate sets, slightly convex, am- Mississippian Blastoids from Montana • Sprinkle and Gutschick 149 bulacral pores appear to die out just above radiodeltoid suture; spiracles well divided by deltoid septa, "C" spiracle separated from rest of half-paired anispiracle (Text- Fig. 18K); ornament consists of relatively coarse growth lines with several of these raised (Plate 6, Fig. 21). Growth features included with S. brei- meri for comparison (Text-Fig. 19). Studied Specimen. MCZ 870. Occurrence. Known from the lower Paine Member, Lodgepole Limestone, from beds 22-33 ft (7-10 m) above the base at Targhee Peak, just inside south- eastern Idaho. Discussion. This specimen occurs with Tanaoblastus in the same beds of the lower Lodgepole Limestone as S. breimeri nor- mally does, but unless it is an abnormal growth variant, it does not appear to be- long to this species. It also occurs in a dif- ferent part of the field area from the nar- row east-west strip where most specimens of S. breimeri have been found. This spec- imen is much wider with a lower L/W ratio and a larger pelvic angle than small specimens of S. breimeri (or S. laudoni); the raised growth lines are also different than most specimens of these species. Per- haps larger specimens of this form would also have been godoniform in thecal shape, but additional specimens will be necessary to determine this. Family GRANATOCRINIDAE Fay, 1961 Subfamily CRYPTOBLASTINAE Fay, 1964 Genus TANAOBLASTUS Fay, 1961 Type Species. Pentremites roemeri Shumard, 1855. Diagnosis. Spiraculate blastoids with a globular theca, ambulacra extending to base or nearly so, base moderately convex to flat, interradial sutures not depressed; eight widely separated spiracles and an anispiracle, two hydrospire folds beneath each side of ambulacrum; four anal del- toids, adoral superdeltoid, aboral hypo- deltoid, and two hidden cryptodeltoids lying beneath hypodeltoid; regular del- toids of moderate size usually visible in side view, occupying from one-sixth to nearly one-half of thecal length, hypodel- toid usually slightly enlarged over regular deltoids, radials usually overlapping del- toids; ambulacra long and relatively nar- row, lancet partly exposed toward adoral end, hydrospire plate present along radials bearing between 1.0 to 1.6 pores per side plate, pores usually absent along deltoids except for 1-3 pores located just above radiodeltoid suture in several species. Occurrence. Early Mississippian (Late Kinderhookian = Late? Tournaisian), cen- tral and western U.S.A. (Missouri, Mon- tana, Idaho, Utah, Wyoming?, Nevada?, Arizona?). Discussion. Tanaoblastus differs from other closely related genera, such as Cryp- toblastus, by having a slightly different thecal shape without depressed interradial sutures and by having a flat to moderately convex base. It differs from Mesoblastus by having only two hydrospire folds per side, a different number of pores along the radials, and larger but less ornamented deltoids. Two species of Tanaoblastus are especially abundant in western Montana and adjacent areas such as southeastern Idaho and northernmost Utah. Very sim- ilar species apparently occur in the early Mississippian of Missouri, especially in the Chouteau Limestone. None of the Mon- tana material shows the hidden crypto- deltoids on the anal side, and it is still un- certain whether Tanaoblastus has four anal deltoids as Fay (1961) reported or whether it might be more advanced and have only two anal deltoids, an epideltoid, and a hy- podeltoid. The Montana material studied here probably makes up the largest col- lections of Tanaoblastus that have ever been assembled. TANAOBLASTUS HAYNESI (Clark), 1917 Plate 1, Figure 1; Plate 7, Figures 1-30; Text-Figures 21A-B, E-F, H, and 22 Schizoblastus haynesi, Clark, 1917, pp. 371-373, plate 1, figures 15-20. Mesoblastus haynesi, Fritz and Cline, 1937, pp. 308- 150 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 309, plate 17, figures 10-12 (but not plate 17, figures 1-9). Tanaoblastus haijnesi, Fay, 1961, pp. 102 and 104, plate 37, figures 7-9. Diagnosis. Theca globular, L/W ratio varies from about 0.9-1.2; ambulacra al- most reaching base in small individuals, reaching base and protruding slightly in large ones; basals flat to moderately con- vex, fairly small, deltoids occupying one- third of theca length in small individuals, about one-sixth of thecal length in large ones, interambulacra flat to slightly con- cave; spiracles wide apart on summit, last pore at radiodeltoid suture; ornament con- sists of fine growth lines on radials, coarser bands along RD fronts, low ridges on bas- als. Description. About 925 specimens of this species available for study from about 30 localities in western Montana (Table 1), ranging throughout the lower 75 ft (23 m) of the Lodgepole Limestone. The follow- ing description is based on the holotype specimen MCZ 347 described by Clark (1917) and about 40 other newly-collected specimens. Theca globular, varying from elongate to squat, small thecae nearly spherical, larger ones more variable; vault rounded, pelvis slightly convex, basals moderately convex to flat, interambulacra usually slightly concave, but may be flat in some specimens, greatest width near midheight; smallest theca about 2.8 mm long, largest theca about 10 mm long and 9 mm wide (Plate 7, Figs. 1-25). PLATE 7 Figures 1-30. Tanaoblastus haynesi (Clark), lower Paine Member, lower Lodgepole Limestone, 1-3 from Brazer Canyon, northeastern Utah, 4-6 and 8-13 from Targhee Peak, southeastern Idaho, 7 from White Peak, 14 and 17-19 from Squaw Creek, 1 5 from North Sawtooth Mountain, northwestern Montana, 1 6 from Northeast Baldy Mountain, 20-22 from Old Baldy Mountain, 23-25 and 29-30 from London Hills, 26 from Cowboy Canyon, 27 from Timber Butte, and 28 from Sixteen Mile Creek, all except 1-6, 8-13, and 15 from southwestern Montana. 1-3, D-side, top, and bottom views of very small theca USNM 16515 showing raised radial edges alongside ambulacra, x 2.7; 4-6, E-side, top, and bottom views of very small theca MCZ 1 030; note relatively short ambulacra and nearly round cross section, x2.7; 7, top view of medium-sized theca USNM 20163 showing eight spiracles and anispiracle, x 2.7; 8-1 0, B-side, top, and bottom views of medium-sized theca MCZ 1 027; note globular shape and pentagonal cross section, x2.7; 11-13, C-side, top, and bottom views of medium-sized elongate theca MCZ 1031 showing ornament, enlarged and slightly raised hypodeltoid, and partly-exposed lancet, x2.7; 14, B-side view of medium-sized squat theca USNM 20602 having rather coarse silicification, x2.7; 15, C-side view of large elongate theca MCZ 1026; note fine ornament and flat interambulacra, x2.7; 16, C-side view of medium-sized elongate theca MCZ 1067 showing plate ornament, ambulacral pores ending at radiodeltoid suture, and slight lancet exposure, x4; 17-19, A-side, top, and bottom views of large squat theca USNM 20603; note fine ornament, raised basals, and slightly concave interambulacra, x2.7; 20-22, E-side, top, and bottom views of large squat holotype MCZ 347 showing ornament, short but visible deltoids, and relatively large flat basals (A ray at bottom), '3.0; 23-25, D-side, top, and bottom views of very large elongate theca MCZ 1028; note coarse silicification, small raised basals, and missing side plates, x2.7; 26, cluster of seven small to medium-sized thecae on partly-etched slab MCZ 1038, x3.2; 27, cross section MCZ 1025 showing two well-preserved silicified hydrospires beneath each ambulacral edge, x5; 28, enlarged top view of theca MCZ 1 035; note eight spiracles and anispiracle plus well-preserved ambulacra, x 4; 29-30, top views of abnormal thecae MCZ 1036 and 1037, both of which lack the A ambulacrum, x3.2. Figures 31-56. Tanaoblastus allanensis Sprinkle and Gutschick, n. sp., lower Allan Mountain Limestone, four localities around Crown Mountain, northwestern Montana. 31-33, E-side, top, and bottom views of small angular paratype MCZ 964 showing ambulacra not reaching base of theca, x2.8; 34-36, C-side, top, and bottom views of small rounded paratype MCZ 965; note well-preserved spiracles and anispiracle, x2.8; 37-39, C-side, top, and bottom views of medium-sized very angular paratype MCZ 968 showing pentagonal cross section and depressed ambulacra, x 2.8; 40-42, A-side, top, and bottom views of medium- sized angular paratype MCZ 970; note long deltoids and abnormal basals (three azygous, one zygous), x2.8; 43-45, B-side, top. and bottom views of large rounded holotype MCZ 963 showing ornament on large deltoids and convex base with secondary deposits around small stem facet, x 2.8; 46-48, C-side, top, and bottom views of large rounded paratype MCZ 972; note large deltoids with low central ridge and rounded cross section, x2.8; 49, AB? interray view of large crushed paratype MCZ 975 showing plate sutures and ornament, x 2.7; 50, B-side view of large crushed paratype MCZ 977; note well-preserved ornament and few ambulacral pores above radiodeltoid suture, x2.7; 51, oblique E-side view of rounded paratype MCZ 967 showing few pores alongside lower deltoids, x2.7; 52, oblique E-side view of angular paratype MCZ 976; note ambulacrum, spiracles, and few ambulacral pores above radiodeltoid suture, x2.7; 53, oblique E-side view of abnormal paratype MCZ 969; E ambulacrum and its lancet missing and surrounding thecal plates in contact across sinus, x4; 54, cluster of nine or more small to medium- sized thecae on paratype slab MCZ 978, x2.5; 55, paratype cross section MCZ 974 showing holes for hydrospires in chert- filled interior, x 4.3; 56, oblique top view of very large crushed paratype MCZ 973 in slab; note large deltoids and broken thecal plates, x2.5. Mississippian Blastoids from Montana • Sprinkle and Gutschick 151 Ki V s ill mi m , a! II * 4 40 -*fe£/V ^> 43 X'32 ^v &fe * U % - ^ 38% ?*V 28 -MtS> 7 -H V m l rvV" ftli 49 «/ ^R. 2§ 49 ../ l v -: : • 4 lit Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 B /d i l r/ •tz /HP •1 •1 — P r.B Text-Figure 21. Morphologic features of Tanaoblastus haynesi (Clark) (A-B, E-F, H) and 7. allanensis, n. sp. (C-D, G, I). A- D, side and summit views of large elongate thecae showing maximum width (short lines) near midheight, longer deltoids and shorter ambulacra in T. allanensis (C-D), and difference in interambulacral shape (flat to slightly concave in T. haynesi vs. convex to humped in T. allanensis). E, enlarged ambulacrum in figured T. haynesi specimen MCZ 1024; note central lancet (L) and inner and outer side plates (ISP and OSP) supporting large brachiole facets (BF) at edge of ambulacrum with pore furrow (PF) wrapping around lower edge; cross section is along side food groove and facet, and small arrow points toward mouth (M). F-G, comparison between T. haynesi (F) and T. allanensis (G) showing that pores (P) through hydrospire plate (HP) either stop at suture between radials (R) and deltoid (D) when side plates missing from lancet (L), or extend short distance further along deltoid edge; F drawn from specimen MCZ 1024, G from paratypes MCZ 965 and 967. H-l, cross sections of ambulacra in T. haynesi specimen MCZ 1 025 (H) and T. allanensis paratype MCZ 974 (I); note lancet (L) overlapped by side plates (SP) in ambulacra and shape of two hydrospire folds (HF) and hydrospire plate (HP) beneath each side of ambulacrum and adjacent radial (R) or deltoid (D). Basals three, normally arranged, two larger and one smaller (azygous), usually slightly convex in profile except in small thecae where moderately convex, relative- ly small, occupying 50 to 60% of pelvis (Plate 7, Figs. 19, 22, and 25). In medium- sized theca, azygous basal 1.6 mm long, 1.6 mm wide, larger basals about same length, and 2.4 mm wide. Center of basal set covered with thin secondary deposits obscuring growth lines and bearing central stem facet about 0.7-0.9 mm in diameter, with about 25-26 crenulae extending one- fourth of distance in from margin, and small round central lumen about 0.07-0.08 mm in size surrounded by a slightly de- pressed region (Plate 7, Fig. 19). Radials five, large, making up most of thecal surface (greater than 60%), re- curved at base with short body occupying 40-50% of pelvis, and long moderately curved limbs enclosing long ambulacral sinus. Radiodeltoid suture nearly straight, RD axis dominates growth (Text-Fig. 22), small radial lip near origin of radials point- ing outward or slightly downward. Regular deltoids four, relatively small, make up 16-20% of length in large thecae, DR suture makes angle of about 135°; del- toid lips small, form edges of mouth, two elliptical spiracles notched in aboral cor- ners of lip with small ridge around inside edge of spiracles; deltoid body larger, dia- mond-shaped, moderately concave adoral- ly, slightly concave to flat aborally, radials slightly overlap deltoids at radiodeltoid su- ture. Anal deltoids apparently four, adoral su- perdeltoid (=lip of regular deltoids), two deep, hidden cryptodeltoids (forming septa separating anus from posterior hydrospires beneath hypodeltoid), and large aboral hy- podeltoid (=regular deltoid bodies), hy- podeltoid slightly enlarged over other del- Mississippian Blastoids from Montana • Sprinkle and Gutschick 153 20 16 E E 12 g 8 • •/• 4 8 12 Width (mm) 20 r 16- 12 « 8 > 16 4 ■ » 4 8 Pelvis (mm) 12 15 12 £9 1 6 < 10 20 30 No. Side Plates 40 15 !« CO 9 CC o S 6 9 3 B X^°o *o 3 6 9 12 15 Growth Front (mm) ^5 E £4 — 3 CD °2 / 5 - 4 - E 3 E CD % • CD o°-° .o°o 012340 3 6 9 12 Max. Del. W. (mm) RDo RB« (mm) 15 • 12 I9 ■II O CC 6 • 0 •O • O 3 ■ ip t? 0 ( il . ) 2 4 6 8 RRo RB« (mm) 6 - E E cc CC / 3 6 RB (mm) 15 12 E E CC CD o Q CC o o 0 2 4 6 Del. L. (mm) Text-Figure 22. Growth plots for 11 measured specimens (MCZ 1026-1028, 1030-1031, 1066-1067, holotype MCZ 347, USGS 1 681 5 and 20602-20603) of Tanaoblastus haynes/ (Clark). Note that the pelvis shows no apparent growth in its contribution to thecal length (top center) and very slow growth in RB and RR. Best-fit lines in all plots were hand fit. 154 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 toid bodies. Anispiracle nearly circular, Hydrospires in 10 groups, two folds per formed by superdeltoid adorally, hypo- ambulacral side, folds hang down into the- deltoid aborally, and small segment of side cal cavity (Text-Fig. 21H; Plate 7, Fig. 27), plates from adjacent adoral ambulacra lat- folds have thin parallel walls and an en- erally (Plate 7, Figs. 18 and 21). larged tube at bottom. Mouth central on summit, formed by Ornament consists of fine growth lines regular deltoid lips plus superdeltoid, pen- along RR front with slightly pustular pen- tagonal to slightly star-shaped, slightly odic markings, basals have closely spaced larger than anispiracle, has lobes and sock- pustular periodic markings, relatively ets on margins. coarse growth lines on RD fronts expand- Ambulacra five, long and fairly narrow, ing towards deltoids, DR growth front usually extending to or near base, mod- variable ranging from fine growth lines to erately curved along length, moderately to fairly coarse pustule-bearing bands (Plate strongly convex (actually biconvex) in cross 1, Fig. 1). Radials and deltoids have raised section (Text-Fig. 21E), highest points at nodes along ambulacra, moderate second- or slightly above level of adjacent thecal ary deposits forming stem facet, slight sec- plates, edges slightly depressed below the- ondary deposits forming radial lips, edges cal plates, widest at radiodeltoid suture, of ambulacra, and slightly raised adoral lancet slightly exposed along most of edge of hypodeltoid. length, about one-third to one-half of its Measurements of growth series speci- width exposed, occupying about one-fourth mens plotted in Text-Figure 22. Small to one-third of ambulacral width, side plate specimens have a fairly large convex base sets on bevelled lancet edge, inner side with short ambulacra, a relatively small plates modified rectangular, outer side V/P ratio, and fairly large deltoids. Large plates small and triangular, occupy ab- specimens tend to become more elongate medial aboral margin of inner side plates, or more squat with relatively small base, each set of inner and outer side plates bears ambulacra usually reaching base, higher a fairly large, nearly circular, brachiole V/P ratio, and deltoids occupying less of facet at edge of ambulacrum, small round thecal length. brachiolar pit near highest point of am- No stems or brachioles known for this bulacrum at end of side food groove, two species. Three abnormal specimens in depressed facets just abmedial to this, about 925 examined (0.3%), all of these where brachiolar plates attached (Text-Fig. four-sided with an ambulacrum that re- 21 E). Large pore furrow curving around mained very small or never developed. Ra- aboral side of brachiole facet; 3-4 lobes dials and deltoids appear normal, but ra- along main food groove usually in lancet dial sinus closed and deltoids in lateral material, 3-4 lobes adorally and usually contact; two abnormalities affect "A" am- two lobes aborally along short side food bulacrum, one affects "D" ambulacrum, grooves mostly on inner side plates. Hy- very small spiracles possibly present in ab- drospire plate present along radials be- normal ray (see Plate 7, Figs. 29-30). neath edge of ambulacra, formed by radial Studied Specimens. Holotype MCZ 347, material, side plate impressions on lancet paratypes MCZ 341; other studied or mea- and adjacent deltoid edge but not on ad- sured specimens MCZ 1024-1038, USGS jacent radial edge or hydrospire plate Collections 16815, 20163, and 20602-3; (Text-Fig. 2 IF), one row of pores in hy- other specimens MCZ 1039. drospire plate, between 1.5-1.6 pores per Occurrence. Known from between 5 and side plate set, pores slightly elongate along 75 ft (1.5-23 m) above the base of the ambulacral length, last pore at radiodel- Paine Member of the Lodgepole Lime- uture round to very elongate, appar- stone at 30 or more localities in south- ; where new pores inserted. western and west-central Montana, south- Mississippian Blastoids from Montana • Sprinkle and Gutschich 155 eastern Idaho, and northeastern Utah; also found in the lower Allan Mountain Lime- stone at North Sawtooth Mountain in northwestern Montana (see Table 1). Discussion. Tanaoblastus haynesi char- acterizes the lower Paine Member of the Lodgepole Limestone in much of western Montana, southeastern Idaho, and north- eastern Utah. This blastoid seems to be present at most sections of the Montana Facies of Sando (1976). The preservation of these silicihed specimens ranges from only fair (see Plate 7, Figs. 23-25), at lo- calities such as London Hills, to excellent when extracted with acetic acid at local- ities such as Standard Creek (see Plate 1, Fig. 1) and Targhee Peak (Plate 7, Figs. 8-13). This species is similar to several species from the Mississippi Valley, such as T. missouriensis and T. tenuis; it differs from these species by usually being less elongate, by having flat to slightly concave interambulacra, by having somewhat stronger growth bands on the basals and periodically on the radials, by having a somewhat different number of pores per side plate set along the radials, and by having different length (usually shorter) deltoids. Tanaoblastus haynesi differs from T. allanensis, n. sp., by having a shorter base, flat to slightly concave interambu- lacra, the last pore at the radiodeltoid su- ture, and shorter deltoids. TANAOBLASTUS ALLANENSIS Sprinkle and Gutschick, new species Plate 7, Figures 31-56; Text-Figures 21C-D, G, I, and 23 Diagnosis. Theca globular, length near- ly equal to width, V/P ratio averaging 3.1, interambulacra moderately convex to strongly angular; deltoids long, occupying between one-third and one-half of thecal length, radials abut deltoids; ambulacra relatively long, not reaching base of theca, 1-3 pores just above radiodeltoid suture; ornamented with moderately coarse growth lines. Description. At least 174 specimens available for study, all from four localities in the lower Allan Mountain Limestone at Crown Mountain in northwestern Mon- tana. Holotype MCZ 963, and 16 other paratypes used for the following descrip- tion. Theca globular to elongate, sometimes squat, rounded to flaring pentagonal in cross section, vault rounded, pelvis slightly to moderately convex, maximum width at or considerably above midheight, ambu- lacra flush with adjacent thecal plates; smallest theca 2.9 mm long, largest free theca about 8.3 mm long and 6.8 mm wide, very large obliquely crushed theca in slab at least 8.5 mm long and 11.5 mm wide; L/W ratio ranging from 0.85 to 1.21, av- eraging 1.0, V/P ratio ranging from 1.95 to 4.38, and averaging 3.1, pelvic angle ranging from 90° to 120° and averaging 112°, based on 10 specimens in growth se- ries. Basals three, normally arranged, two larger and one smaller (azygous), fairly small, make up 50-60% of pelvis, usually convex in side view with small stem facet (Plate 7, Fig. 42), in large specimen azy- gous basal 2.1 mm long and wide, larger basals about same length and about 2.8 mm wide. Radials five, fairly large, make up much of thecal surface (about 50%), have small lip at radial origin, body fairly short, mak- ing up about 40% of pelvis, limbs fairly long, making up half or more of ambu- lacral sinuses, adoral end of radials raised, producing convex to angular cross section. Regular deltoids four, fairly large, oc- cupying one-third or more of curved ara- bulacral sinuses, between one-third and one-half of thecal length (Text-Fig. 21C), rhombic-shaped, aboral end strongly con- vex or crested to meet raised adoral radials, radiodeltoid suture forms angle between 90° and 130°, radials appear to abut del- toids without any overlap. Spiracles eight, slightly elliptical, at lateral margins of del- toid lips, about 0.6 mm apart across del- toids and across ambulacra, slight ridges just inside spiracles on edges of deltoids; mouth central on summit, about 0.6-0.7 156 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 mm in diameter, slightly wider than long, into thecal cavity with thin lamellae and star-shaped to pentagonal, margins formed enlarged tubes at bottom, entrance to hy- by regular deltoid and superdeltoid lips. drospires through pores in hydrospire plate Anal deltoids apparently four, small su- (Text-Fig. 21H; Plate 7, Fig. 55). perdeltoid adorally (=lips of other del- Ornament consists of moderate to fairly toids), two crvptodeltoids apparently hid- coarse growth lines on basals, radials, del- den below hypodeltoid, and aboral, fairly toids, and hypodeltoid, ornament along RD large hypodeltoid (=body of other del- and DR fronts especially coarse (Plate 7, toids); hypodeltoid either same size or Figs. 44 and 50). Secondary deposits mi- slightly enlarged over other deltoids, raised nor, forming stem facet and small radial and slightly hooded adorally over anispira- lips. cle, which is surrounded by superdeltoid Measurements of 10 specimens in growth adorally, few side plates of ambulacra lat- series plotted in Text-Figure 23. Relatively erally, and hooded hypodeltoid aborally, little change in thecal shape with increas- anispiracle rounded to pentagonal, slightly ing size. smaller than mouth, about 0.6 mm in di- No stems or brachioles known for this ameter. Cryptodeltoids difficult to see, may species. Two abnormal thecae found in 174 form septa separating posterior hydro- examined (1.1%); one has no "E" ambu- spires from anus deep within anispiracle. lacrum although "E" radial appears nor- Ambulacra five, relatively long but not mal-sized with limbs in contact and DE reaching base of theca, rather narrow, flush and EA deltoids abut each other appar- with or slightly depressed below adjacent ently with two tiny spiracles near normal thecal plates, slightly to moderately con- positions (Plate 7, Fig. 53). Other theca vex; lancet slightly exposed along most of has four basals, one larger and three small- length, occupying one-fourth to one-third er; one of larger basals (apparently DA) of ambulacral width, mostly forming main split into two smaller plates, resembling food groove and adjacent lobes in center azygous AB basal. of ambulacrum, side food grooves enter Studied Specimens. Holotype MCZ 963, main food groove at 40-70° angle. Side paratypes MCZ 964-979, other additional plates normally developed, inner side plates specimens MCZ 980. rectangular, small triangular outer side Occurrence. All studied specimens come plates notch aboral abmedial edge of inner from the area around Crown Mountain, ones, together supporting relatively small between 11 and 35 ft (3.5-11 m) above brachiole facet at edge of ambulacrum, the base of the Allan Mountain Limestone Between side food grooves, 4-5 lobes along in northwestern Montana, main food groove, usually 3 adoral lobes Etymology. Named for the Allan Moun- and 2 small aboral lobes along each side tain Limestone, where this species occurs, food groove, which leads to small brachio- Discussion. Tanaoblastus allanensis ap- lar pit and two small hemispherical pears to be a paedomorphic derivative of depressions where brachiole attached, some other Tanaoblastus species, perhaps Pores developed in hydrospire plate along- T. haynesi which also occurs in the Early side ambulacra, side plates do not com- Mississippian of western Montana. It shows pletely cover hydrospire plate and pores, considerable resemblance to the juvenile average of about 1.2 pores per side plate specimens of this species and some simi- set along radials, pores absent along much larity to species known from the Chouteau of deltoid except at aboral end where 1- Limestone in Missouri. The convex base, 3 extra pores located just above radiodel- long deltoids, relatively long ambulacra toid suture (Text-Fig. 21G). that do not reach the base, and convex to Hydrospires in 10 groups, two hydro- angular shape in cross section are all fea- spires per ambulacral side, folds hang down tures similar to juveniles of T. haynesi, Mississippian Blastoids from Montana • Sprinkle and Gutschick 157 E E 20 16 ■ 12 • 8- .•; 4 8 12 Width (mm) 20 16 ■ | 12 3 8 / > i 16 4 8 12 Pelvis (mm) 15 12 E £9 1 6 < 3 ■ / / 10 20 30 No. Side Plates 40 15 E E 12 CO cc o cc cc a O CC L D a $£&> 3 6 9 12 15 Growth Front (mm) £4 _• 3 0) °2 5 " E 3 E CO •• 00 012340 3 6 9 12 Max. Del. W. (mm) RDo RB» (mm) 15 12 I9 Q a: 6 / 6 - E E, tr cc 15 12 E E, • CO o 0 cc 00 o 00 ^:< 00 2 4 6 8 RRo RB» (mm) 3 6 RB (mm) 0 2 4 6 Del. L. (mm) Text-Figure 23. Growth plots for 1 0 measured specimens (MCZ 863-872) of Tanaoblastus allanensis, n. sp. Note slow growth in pelvis because of slow growth in RB and BR. Best-fit lines in all plots were hand fit. 158 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 although the cross section is not so angular in that form. The small size of most adult specimens of T. allanensis may also agree with this possible derivation. Tanaoblastus allanensis shows some resemblance to Mis- sissippi Valley forms such as T. tenuis (Hambach) and T. roemeri (Shumard) in its globular cross section and protruding base, but the deltoids are not so angular in either of these Missouri forms and the base is much less pronounced than in T. alla- nensis. These Chouteau species do not dif- fer much among themselves, and we ques- tion whether all of these are really distinct species. The cross-sectional profile of T. allanen- sis is probably the most variable feature of this species. Rounded forms and highly angular forms are fairly easy end members to pick out of the available material, but many intermediates exist and some forms cannot be assigned to one or the other with any certainty. We considered the possi- bility that two separate species might be present in this Crown Mountain material, but decided that these are probably highly variable individuals in a single species be- cause of the many intermediates and the occurrence of both forms at all four of the Crown Mountain sections. The angularity in cross section probably represents a growth feature that was not highly con- trolled genetically. Genus CRYPTOBLASTUS Etheridge and Carpenter, 1886 Type species. Pentremites melo Owen and Shumard, 1850. Diagnosis. Spiraculate blastoids with an ellipsoidal or ovoid theca, base fairly small, usually with small, depressed basals; eight spiracles plus an anispiracle; four anal del- toids present, small adoral superdeltoid, two deep, hidden cryptodeltoids, and ab- oral hypodeltoid that is not enlarged; am- bulacra long, slightly depressed, extend to base of theca, lancet slightly exposed along most of length, hydrospire plate present with about 1.5 pores per side plate set along radials, pores absent along deltoids; inter- radial sutures often depressed, radials overlap deltoids. Occurrence. Early to Middle Mississip- pian (Kinderhookian to Osagean), Missis- sippi Valley and northwestern Rockies, plus southern Canadian Rockies. Discussion. At least five species of glob- ular blastoids occur in the U.S. and Ca- nadian Rockies that may belong to Crypto- blastus; three of these occur in the Lodgepole Limestone in western Mon- tana. Two other species, including the form called Mesoblastus haynesi described by Fritz and Cline (1937) and a form that occurs in the Banff Formation near Lake Minnewanka in southern Alberta, also ap- pear to belong to Cryptoblastus. The three Lodgepole species occur in the middle and upper parts of this unit at different local- ities; unfortunately, none of them is par- ticularly common, complete, or well pre- served. For this reason they are not formally named in this paper, but are only briefly described and illustrated. CRYPTOBLASTUS? species A Plate 4, Figures 14-28; Text-Figures 24A-D and 25 About 11-12 silicified specimens and several fragments and plates from two lo- calities near the top of the Lodgepole Limestone appear to belong to one species of Cryptoblastus. This species has an el- lipsoidal theca with a rounded vault and a medium-sized base having a concave basal cavity, a L/W ratio ranging from 1.04 to 1.32, averaging 1.15 in the five nearly complete specimens, relatively short deltoids, and nearly paired, closely set spi- racles. Theca ellipsoidal, base medium-sized, depressed in center, profile in oral view pentagonal with flat to slightly convex in- terambulacra. Large, nearly complete the- ca 8.0 mm long, 7.6 mm wide, vault oc- cupying entire length, pelvis depressed, basals inset about 1.0 mm above tips of ambulacra. Basals three, small, make up shallow basal cavity, deepest part occupied by fairly small stem facet about 1.0 mm Mississippian Blastoids from Montana • Sprinkle and Gutschick 159 in diameter; radials five, very large, oc- cupy about 80% of thecal surface, strongly recurved at base with short body about 1.5 mm long and long limbs about 7.0 mm long; deltoids four, short, barely appearing on side of theca, body about 1.2 mm long, with short lip adorally; hypodeltoid also small, not enlarged, apparent superdeltoid slightly wider than other deltoid lips, cryp- todeltoids not seen but thin septa internally separate anus from posterior spiracles; eight closely spaced spiracles on summit, thin septum connects deltoid body and lip, and barely separates spiracles at surface, spi- racles widely separated across adjacent ambulacra. Ambulacra five, long, extend- ing to base of theca, where small radial lips present, ambulacra convex and an- gular, nearly flush with adjacent thecal plates, lancet appears slightly exposed along most of length, side plates numerous, hy- drospire plate with 1.5-1.6 pores per side plate set along radials, pores apparently absent along short deltoids, thin raised ridges on edges of radials and deltoids. Ornament consists of fine growth lines with small pustules on radials, coarser growth lines on RD fronts and deltoids, radial body and basals nearly smooth. Studied Specimens. MCZ 1045-1061 plus one theca and several plates in USGS Collection 20670 (Sando Collection). Occurrence. Upper Lodgepole Lime- stone, most specimens from about 655 ft (200 m) above the base in the upper half of Woodhurst Member at Sacagawea Peak, northern Bridger Range, southwestern Montana; single specimen (MCZ 1059) with similar features in a float slab from an un- known footage in the upper Lodgepole? Limestone at Pole Canyon, northern To- bacco Root Mountains, southwestern Mon- tana. Discussion. This form has many fea- tures similar to Cryptoblastus melo, but some that are different. The medium-sized, slightly concave base is somewhat differ- ent, and this form lacks the depressed su- tures that characterize C. melo. The num- ber of anal deltoids is unknown; if it is four as suspected, this form would be most closely related to Cryptoblastus and has been questionably assigned to that genus here. More and better-preserved speci- mens will be necessary to confirm this as- signment. CRYPTOBLASTUS? species B Plate 4, Figures 31-34; Text-Figure 24E A small number of distinctive plates be- longing to a globular blastoid were re- covered from acid residues from the large Koryschisma block found at Bandbox Mountain in west-central Montana. This form is known from only four plates or fragments, but may also belong to Cryp- toblastus. Available material includes a partial base with all three basals and parts of two radials, a partial radial with one nearly complete limb and ambulacral margin, and two small spine-bearing del- toids. Base nearly flat, fairly wide, basals three, slightly convex, occupy 55-60% of short pelvis, stem facet protrudes slightly, about 1.2 mm in diameter, azygous basal about 1.9 mm long, 2.1 mm wide, larger basals about same length, about 2.5 mm wide, both basals and radial bodies ornamented with fine growth lines. Radial bodies short, limbs long, radials at least 6.5 mm long in largest fragment, ambulacra absent, but some evidence for pores along radial mar- gin (in hydrospire plate?), radials have me- dium-sized lips pointing outward at tips of ambulacra, possibly two hydrospire folds per ambulacral side, ambulacra apparent- ly narrow. Deltoids short, body orna- mented with fairly coarse growth lines, adoral tip of deltoid body bears either two or three large spines (Plate 4, Fig. 31-32), thin septum leading from body to lip im- plies spiracles closely spaced or possibly paired, possibly two hydrospire folds be- neath deltoid body leading to spiracles, ra- dials appear to overlap deltoids, and ra- diodeltoid suture forms angle near 125°. Studied Specimens and Occurrence. MCZ 1041-1044 from a block of limestone 160 Bulletin Museum of Comparative Zoology, Vol. 152, No. 3 BRF B Text-Figure 24. Morphology of Cryptoblastus? sp. A (A-D), Cryptoblastus? sp. B (E), and Cryptoblastus? sp. C (F). A, side view of large theca based on MCZ 1047 showing ellipsoidal shape (short lines at maximum width) with wide concave base, long fairly narrow ambulacra, and short deltoids just visible in side view, x3.3. B, enlarged summit view in MCZ 1046; note closely set spiracles, small deltoids (D), and hypodeltoid (HD), x5. C, plan view of ambulacrum based on MCZ 1050 and 1056 showing lancet (L) slightly exposed in center, large inner and small wedge-shaped outer side plates (ISP and OSP), and small brachiole facet (BRF) at edge of ambulacrum alongside hydrospire plate with pores (P), x6.6. D, cross section of ambulacrum in MCZ 1054; note lancet (L) mostly covered by side plates (SP) and two hydrospire folds (HF) beneath each side of the ambulacrum, x 5. E, reconstructed thecal shape based on fragments MCZ 1 041 -1 044 showing slightly convex base and spiny deltoids, x 2.5. F, reconstructed thecal shape in MCZ 1 040 which is mostly buried in a slab and draped with brachioles (BR), x 3.3. about 170-175 ft (52-53 m) above the base of the Paine Member, Lodgepole Lime- stone, at Bandbox Mountain, Little Belt Mountains, west-central Montana. This species is apparently a rare spiraculate oc- curring with the fissiculate Koryschisma elegans. Discussion This species has a flat to slightly convex base and spiny deltoids un- like the other species of Cryptoblastus? and unlike the type species C. melo. The species is probably new but is not named here because of the fragmentary speci- mens. CRYPTOBLASTUS? species C Plate 4, Figures 29-30; Text-Figure 24F This form is known from a single spec- imen from the middle Lodgepole Lime- stone at Northeast Baldy Mountain, south- ern Bridger Range, southwestern Montana. It occurs with numerous specimens of Montanablastus baldyensis and less com- mon specimens of Strongyloblastus lau- doni. The specimen is sitting vertically in a slab of limestone with the base exposed, weathered, and partly silicihed; brachioles are splayed out on the slab surface from all five ambulacra (Plate 4, Figs. 29-30). Ambulacra long and recurved, appar- ently reaching base of theca; using growth lines and internal calcite, base of theca (now eroded) apparently slightly convex and basals small to medium in size. Radials long with fine growth lines, ambulacra long, fairly narrow, with many side plate sets, brachioles still preserved attached to all five ambulacra, at least 18 mm long and about 0.2 mm wide, fairly well pre- served, brachiolar plates 0.25 mm long and deep, food groove not seen except in cross sections where filled with pyrite specks. Back of slab ground down perpendicular to thecal axis to intersect summit (Text- Mississippian Blastoids from Montana • Sprinkle and Gutschich 161 20 16 (lulu) jz *-* g 8 _l • 4 - / 0 ( ) 4 8 12 16 20 16 12 rt 8 > Width (mm) 4 8 12 Pelvis (mm) 15 12 £9 I 6^ < / • • 10 20 30 No. Side Plates "40 15 t'2f CO 9 - CC o CC 6 CC 9 3- cc •• 00 o o 3 6 9 12 Growth Front (mm) CD Q —1 15 E 3 E CO 0 12 3 4 Max. Del. W. (mm) OO