z < n LIBRARIES SMITHSONIAN INSTITUTION NOIiniliSNI* NVINOSHIIIAIS h- 3 NoiiniusNi nvsnoshuws S3 1 uyu an LIBRARIES to SMITHSONIAN SMITHSONIAN INSTITUTION NOIinillSNI co INSTITUTION x to O 2 / NVINOSH11SAJS S3 I dVdS II to yj > 2 LIB 2: co NOlifUllSNI NVINOSHimS S3 I d Vd 8 IT LIBRARIES SMITHSONIAN Z s INSTITUTION NO I o 2 LIBRAR I ES SMITHSONIAN INSTITUTION NOIinillSNI NVINOSHIIIAIS S3 I d Vd 8 SI NOIinillSNI NVIN0SH1IWS S3 I dVd a n to LIBRARIES SMITHSONIAN co : INSTITUTION co S3idvdan NOIinillSNI NVINOSHIIIAIS S3SdVdai1 LIBRARIES SMITHSONIAN .... ' co INSTITUTION LIBRAR ES SMITHSONIAN to INSTITUTION NOIinillSNI CO NVINOSHIIIAIS S3 co dvda ii LU NOIinillSNI NVINOSHIIIAIS LIBRAR I ES SMITHSONIAN INSTITUTION NVIN0SH11IAIS ■ > 2 O 1 LIBRAR s— INSTITUTION N0S1S11I1SNS NV r~ (T\ Number 439 24 February 1994 9^ >v* Contributions in Science Additional Uintan and Duchesnean (Middle and Late Eocene) Mammals from the Sespe Formation, Simi Valley, California Thomas S. Kelly and David P. Whistler Natural History Museum of Los Angeles County Serial Publications of THE Natural History Museum of Los Angeles County Scientific Publications Committee Craig C. Black, Museum Director Daniel M. Cohen Kirk Fitzhugh John M. Harris, Committee Chairman Edward C. Wilson Richard C. Hink, Managing Editor Robin A. Simpson, Head of Publications The scientific publications of the Natural History Mu- seum of Los Angeles County have been issued at irregular intervals in three major series; the issues in each series are numbered individually, and numbers run consecutively, regardless of the subject matter. # Contributions in Science, a miscellaneous series of tech- nical papers describing original research in the life and earth sciences. # Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth sci- ences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now published by the Museum in Contributions in Science. # Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through the Museum Book Shop. A catalog is available on request. The Museum also publishes Technical Reports, a mis- cellaneous series containing information relative to schol- arly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum’s Scientific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Tech- nical Reports may be obtained from the relevant Section of the Museum. Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Printed at Allen Press, Inc., Lawrence, Kansas ISSN 0459-8113 Additional Uintan and Duchesnean (Middle and Late Eocene) Mammals from the Sespe Formation, Simi Valley, California Thomas S. Kelly1 and David P. Whistler1 ABSTRACT. A paleontologic resource impact mitigation program at the Simi Valley Landfill, Ventura County, California, is yielding new taxa and new geologic records of middle and late Eocene mammals from the middle member of the Sespe Formation. New Uintan taxa and new occurrences recorded from Simi Valley are Centetodon sp., cf. C. aztecus; erinaceomorph, gen. and sp. undet.; Uintasorex sp., cf. U. montezumicus; Microparamys woodi n. sp.; Microparamys sp., cf. M. tricus\ Miacis sp. undet.; and Pisectolophid, gen. and sp. undet. New Duchesnean taxa and new occurrences recorded from Simi Valley are Peradectes californicus; Leptotomus sp. undet.; PCamelidae, gen. and sp. undet.; Simimeryx sp., aff. S. hudsoni; and Mammalia, gen. and sp. undet. Sespedectes singulars is now recorded from the Simi Valley Landfill Local Fauna, and this occurrence extends the geologic range of this species upward into the late Duchesnean. Additional specimens of the rare taxa Dyseolemur pacificus, Griphomys alecer, and Protylopus robustus are now available and allow reevaluation of the intraspecific variation in the teeth of these species. INTRODUCTION The middle member of the nonmarine Sespe For- mation, which is exposed along the northern side of Simi Valley, Ventura County, California, has yielded diverse middle and late Eocene land mam- mal assemblages (Stock, 1932;Golz, 1976;Golzand Lillegraven, 1977; Mason, 1988; Kelly, 1990, 1992; Kelly et ah, 1991). Kelly et al. (1991) documented the preliminary results of a paleontologic resource impact mitigation program that is being conducted in the lower and middle members of the Sespe Formation at the Simi Valley Landfill. The program has yielded many new taxa and new geologic and geographic records from the Sespe Formation that were discussed only briefly by Kelly et al. (1991). Many of these new taxa and records are biostrati- graphically significant and have not been adequately described. Kelly (1992) recently described the ro- dents of the families Eomyidae, Heliscomyidae, Simimyidae, and PZapodidae recovered during the program. The report herein describes additional new taxa and new specimens of poorly known taxa that were discovered during the program. 1. Vertebrate Paleontology Section, Natural History Museum of Los Angeles County, 900 Exposition Boul- evard, Los Angeles, California 90007. Contributions in Science, Number 439, pp. 1-29 Natural History Museum of Los Angeles County, 1994 METHODS All specimens were recovered from the middle member of the Sespe Formation by a process described by Kelly et al. (1991) that included wet screening of bulk matrix samples and heavy liquid separation of fossils. All speci- mens described herein are deposited in the Vertebrate Paleontology Collection of the Natural History Museum of Los Angeles County. Measurements of large teeth were made to the nearest 0.1 mm with a vernier caliper, and those of smaller teeth were made with an AO optical micrometer to the nearest 0.01 mm. All teeth were measured at their greatest di- mensions. Metric abbreviations and dental formulae fol- low standard usage. All cladistic analyses were performed using Version 1.5 of the Hennig86 computer program (Farris, 1988) and run on a 386 personal computer. Cladograms were generated using the IE COMMAND that computes the most par- simonious cladograms by implicit enumeration. Weight- ing of characters was accomplished by using the XSTEPS W COMMAND that sets the character weights of the cladograms generated by the IE COMMAND by calcu- lating the best fits of each character based on the product of the character consistency and character retention in- dices. The characters and character states used in the cladistic analyses are presented in Appendix A and the character state matrices for each group of taxa analyzed are presented in Appendices B, C, and D. Institutional acronyms are as follows: CIT — California Institute of Technology LACM — Natural History Museum of Los Angeles County Figure 1. Peradectes calif ornicus (Stock), LMi5 LACM 132422, occlusal view. Scale - 0.5 mm. LACM (C1T) — California Institute of Technology local- ity, numbers and specimens now held by LACM UCMP-— University of California, Berkeley, Museum of Paleontology UCMP-V — University of California, Berkeley, Museum of Paleontology, vertebrate fossil locality UCR — University of California, Riverside, Department of Earth Sciences Abbreviations are as follows: A-P — Anteroposterior ANT — anterior Ar-Ar — argon-argon C.V. — coefficient of variation K-Ar — Potassium-argon L — left Ma — million years before present N — number of specimens O.R. — observed range POST — posterior R — right S.D. — standard deviation TR — transverse SYSTEMATIC PALEONTOLOGY Order Marsupialia Illiger, 1811 Family Didelphidae Gray, 1821 Tribe Peradectini Crochet, 1979 Genus Peradectes Matthew and Granger, 1921 Peradectes californicus (Stock, 1936) Figure 1 REFERRED SPECIMEN. LM„ LACM 132422. LOCALITY. LACM 5876. FAUNA AND AGE. Simi Valley Landfill Local Fauna, late Duchesnean. DISCUSSION. The M, of LACM 132422 ex- hibits the diagnostic characters of Peradectes cal- ifornicus, including a paraconid that is larger than the metaconid, a reduced entoconid, a weak en- toconid notch, a dorsally projecting hypoconulid that is twinned with the entoconid, and a cristid obliqua that extends posteriorly from the trigonid before turning labially. The measurements of LACM 132422 are 1.55 mm A-P, 0.71 mm ANT-TR, and 0.75 mm POST-TR. Peradectes californicus was previously known from the late Uintan Tapo Canyon and Brea Can- yon Local Faunas of the Sespe Formation (Stock, 1936; Golz, 1976; Kelly, 1990; Kelly et al., 1991), the late Uintan Mission Valley and upper PSantiago Formations from the San Diego and Oceanside ar- eas of California (Lillegraven, 1976; Walsh, 1991), the latest Uintan or earliest Duchesnean Camp San Onofre Local Fauna from the PSantiago Formation on the Camp Pendleton Marine Corps Base in Cal- ifornia (Lillegraven, 1976), and the late Uintan to early Duchesnean Badwater Localities 5, 5 Front, 5 Back, 6, and 20 of the PWagon Bed Formation in Wyoming (Krishtalka and Stucky, 1983). The recovery of P. californicus from locality LACM 5876 represents the first Duchesnean record of this species from the Sespe Formation and extends the geologic range of the species into the late Duches- nean. Order Insectivora Illiger, 1811 Family Geolabididae McKenna, 1960 Genus Centetodon Marsh, 1872 Centetodon sp., cf. C. aztecus Lillegraven, McKenna, and Krishtalka, 1981 Figure 2, Table 1 REFERRED SPECIMENS. RM>, LACM 130750; 2LM,s, LACM 130831, 130832 4LM2s, LACM 130830, 130835, 130837, 130838; RM„ LACM 130839; RM„ LACM 130829. LOCALITIES. LACM 5661, 5857, 5866, 5869. FAUNAS AND AGE. Tapo Canyon and Brea Canyon Local Faunas, late Uintan. DESCRIPTION. Only one upper molar referable to Centetodon has been recovered during the im- pact mitigation program. The upper molar has a robust parastyle that is directed anterolabially. The large metastyle is directed posterolabially and po- sitioned much farther labially than the parastyle. The paracone and metacone are conical cusps with sharp-tipped apices and are positioned relatively close together towards the midline of the tooth. The protocone is a well-developed cusp with the preprotocrista and postprotocrista extending to a very small protoconule and metaconule, respec- tively. The anterior cingulum is moderately devel- oped and extends lingually from the middle of the anterior surface of the tooth to the anterolingual base of the protocone. The posterior cingulum is well developed and extends lingually from the pos- terior surface of the tooth, at the level of the meta- conule, to terminate in a distinct small bulge at the posterolingual base of the protocone. Although the roots are broken ofF near the base of the tooth, a single lingual root appears to have been present. The lower molars are typical of those of Cen- tetodon and are characterized by the following: (1) they have a basic tribosphenic pattern; (2) the an- Kelly and Whistler: Sespe Mammals 2 ■ Contributions in Science, Number 439 I Figure 2. Centetodon sp., cf. C. aztecus Lillegraven et al., A. RM1, LACM 130750; B. LM2, LACM 130835. All occlusal views. Scale = 1 mm. terior cingulid is well developed; (3) the paraconid is shelf-like; (4) the protoconid is the tallest cusp; (5) the cristid obliqua is usually medially concave; and (6) the entoconid and hypoconulid are posi- tioned close together on the M,_2. DISCUSSION. The specimens of Centetodon from Simi Valley are very similar morphologically to those of C. aztecus of the Friars and Mission Valley Formations from the San Diego area, Cali- fornia, particularly with regard to the possession of a medially concave cristid obliqua on the lower molars. The mean measurements of the lower mo- lars from Simi Valley are slightly larger than those of the sample of C. aztecus from the San Diego area. However, all of the Simi Valley lower molars fall within the observed range or within less than one standard deviation from the mean of those for C. aztecus. The M1 from Simi Valley is smaller than any of those of C. aztecus from the San Diego area and is about three and one-half standard deviations from the mean of those of the San Diego sample. It is difficult to determine the significance of this size difference because only five M's of C. aztecus are known and such a small sample is not expected Table 1. Measurements (in mm) of dentition of Cen- tetodon sp., cf. C. aztecus. N Tooth/ dimension O.R. Mean 1 M1 A-P 1.49 1 ANT-TR 1.83 1 POST-TR 2.03 2 M, A-P 1.71-1.77 1.74 2 ANT-TR 1.10-1.24 1.17 2 POST-TR 0.99-1.06 1.03 4 M, A-P 1.48-1.75 1.63 4 ANT-TR 1.00-1.15 1.06 5 POST-TR 0.83-0.99 0.90 1 M, A-P 1.67 1 ANT-TR 1.05 1 POST-TR 0.73 Figure 3. Sespedectes singulars Stock, A. RM„ LACM 132459; B. LM2, LACM 132458. All occlusal views. Scale = 1 mm. to include the full range of intraspecific variation. Furthermore, the discrepancies in the comparative upper and lower molar sizes within the Simi Valley sample of Centetodon indicate that the sample ei- ther represents more than one species or a highly variable species. Until a larger sample from the Sespe Formation is available for adequate analysis and comparison with the San Diego sample, the Simi Valley teeth are herein referred to C. sp., cf. C. aztecus , recognizing that this taxon may represent more than one species. Suborder Erinaceomorpha Gregory, 1910 Family Dormaaliidae Quinet, 1964 Subfamily Sespedectinae Novacek, 1985 Genus Sespedectes Stock, 1935c Sespedectes singularis Stock, 1935c Figure 3 REFERRED SPECIMENS. Partial dentary with RM,, LACM 132459; LM„ LACM 132458. LOCALITY. LACM 5876. FAUNA AND AGE. Simi Valley Landfill Local Fauna, late Duchesnean. DISCUSSION. The two lower molars from LACM 5876 are morphologically indistinguishable from those of 5. singularis. The measurements of LACM 132458 are 1.96 mm A-P, 1.35 mm ANT- TR, and 1.32 mm POST-TR, and those of LACM 132459 are 1.85 mm A-P, 1.51 mm ANT-TR, and 1.47 POST-TR. These molars are significant because they rep- resent the highest stratigraphic occurrence of S. singularis in the Sespe Formation and extend the geologic range of this species into the late Du- chesnean. PSespedectinae Novacek, 1985 erinaceomorph, gen. and sp. undet. Figure 4 REFERRED SPECIMENS. RP4, LACM 131088; RM2, LACM 131091. Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 3 A B Figure 4. Erinaceomorph, gen. and sp. undet., A. RP4, LACM 131088; B. RM2, LACM 131091. All occlusal views. Scale - 1 mm. LOCALITY. LACM 5863. FAUNA AND AGE. Brea Canyon Local Fauna, late Uintan. DESCRIPTION. The P4 is subtriangular in oc- clusal outline and the labial surface is straight. The paracone is slightly compressed transversely and taller than the protocone. An incipient metacone is present posterior to the paracone along a sweep- ing posterior crest of the paracone (= metastylar crest). The parastyle is a small distinct cusp. The posterolingual cingulum extends posteriorly from the protocone and joins the posterior cingulum. The measurements of the P4 are 1.86 mm A-P and 1.89 mm ANT-TR. The partial M2 is missing the paracone, meta- cone, and part of the anterior cingulum. The para- conule is smaller than the metaconule and is con- nected to the protocone by the preprotocrista. The metaconule is a prominent, relatively isolated cusp. The protocone is a tall, sharp-crested cusp with the postprotocrista extending posterolabially and ter- minating at the posterior aspect of the metaconule. The hypocone is a well-developed cusp that is con- nected to the posterior and lingual cingulae. Al- though only partially represented, the anterior cin- gulum appears to have been robust. The only measurement that can be estimated for the partial M2 is the A-P, which is 1.9 mm. DISCUSSION. The P4 and M2 from locality LACM 5863 are morphologically most similar to those of Sespedectes, Proterixoides Stock, 1935c, and Crypholestes (Novacek, 1976). The P4 and M2 are smaller than those of Proterixoides, slightly larger than those of Sespedectes, and about the same size as those of Crypholestes. They differ from those of Sespedectes by having the P4 less trans- versely elongated with a smaller metacone on the metastylar crest and more acute cusps (less buno- dont), and the M2 with a less bunodont protocone, a much more robust precingulum, a weaker hy- pocone that is positioned further posteriorly, and a well-developed lingual cingulum between the hy- pocone and protocone. They differ from those of Proterixoides by having the M2 with a lingual cin- gulum present and sharper (less conical) primary A , , B Figure 5. Dyseolemur pacifcus Stock, A. LP4, LACM 131087; B. RM\ LACM 131089. All occlusal views. Scale = 1 mm. cusps. They differ from those of Crypholestes by having the P4 less transversely elongated, the M2 metaconule more isolated, and an M2 lingual cin- gulum present. The P4 and partial M2 from locality LACM 5863 appear to represent a new species of erinaceomorph insectivore most closely related to species of the Sespedectinae. However, due to the lack of ade- quate material, these specimens are referred to er- inaceomorph, gen. and sp. undet. Order Primates Linnaeus, 1758 Family Omomyidae Trouessart, 1879 Genus Dyseolemur Stock, 1934a Dyseolemur pacificus Stock, 1934a Figure 5 REFERRED SPECIMENS. LP4, LACM 131087; RM\ LACM 131089; LP4, LACM 131090; partial RM„ LACM 130749. LOCALITIES. LACM 5616, 5863. FAUNA AND AGE. Brea Canyon Local Fauna, late Uintan. DESCRIPTION. The P4 (LACM 131087), the first upper premolar from the Sespe Formation re- ferred to D. pacificus, is only slightly worn. The paracone is a sharp tall cusp that is slightly com- pressed transversely. The labial margin of the tooth is convex. The posterolabial aspect of the tooth is swollen and larger than the anterolabial aspect. The preparacrista extends anteriorly from the paracone to the posterior aspect of the tooth and then turns labially to join the labial cingulum. The postpara- crista extends posteriorly to the posterior cingulum and then turns labially to join the labial cingulum. A very small metastyle is present at the junction of the postparacrista and the labial cingulum. A small distinct parastyle is present. The protocone is a conical cusp with a sharply pointed apex. A small short preprotocrista extends anterolabially from the 4 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals protocone to the base of the paracone. The post- protocrista is a weak crest that extends posteriorly down the posterior surface of the protocone, wherein it joins the posterior cingulum. A small bulge, which appears to represent an incipient hy- pocone, is present at the junction of the postpro- tocrista and the posterior cingulum. The precin- gulum is a distinct crest that extends labially from the anterior base of the protocone to the parastyle. The posthypocone crista is robust and extends la- bially from the incipient hypoconal bulge to the anterior base of the paracone, near the point where the preparacrista turns labially. The labial cingulum is complete across the labial surface of the tooth but is distinctly thinner along the medial labial as- pect. The measurements of the P4 are 1 .8 1 mm A-P and 2.16 mm TR. The 1VT discovered during the program is well preserved and less worn than the two previously known JVTs of D. pacificus. It differs from these i\Ts by the following characters: (1) the metaconule is better developed and slightly more isolated; (2) a connecting crest from the metaconule to the metacone is lacking; and (3) the hypocone is a dis- tinct cusp on the posterior cingulum. The mea- surements of the NT are 1.88 mm A-P and 2.81 mm TR. The newly discovered P4 and partial M2 are in- distinguishable from previously described P4s and M2s (Stock, 1934a; Szalay, 1976; Kelly, 1990) and, therefore, detailed descriptions of these teeth are not warranted. The measurements of the P4 are 1 .72 mm A-P and 1.44 mm TR, and those of the M> are 2.03 mm A-P and 2.01 mm ANT-TR. DISCUSSION. The impact mitigation program has resulted in the recognition of additional spec- imens of the rare primate D. pacificus, including the first P4 from the Sespe Formation referable to this species. Walsh (1987) assigned a P4 from the Mission Valley Formation in the San Diego area to D. pacificus, and a comparison of the description of this specimen with the newly discovered Simi Valley specimen appears to substantiate his assign- ment. Szalay and Delson (1979) regarded Dyseolemur, Washakius Leidy, 1873, and Shoshonius Granger, 1910, to be related because they share the derived character of having mesostylids present on the low- er molars. Szalay and Delson considered Dyseole- mur more closely related to Washakius than to Shoshonius because the upper molars of Shoshoni- us possess mesostyles whereas in Washakius and Dyseolemur they lack mesostyles. Furthermore, Szalay and Delson noted that the shape of the M,_2 talonid notch of Dyseolemur and Washakius is nearly identical. A close relationship between Dyse- olemur and Washakius is further supported be- cause the Simi Valley P4 of Dyseolemur also ex- hibits greater morphological similarity to those of Washakius than to those of Shoshonius. In Sho- shonius a distinct metacone is present on the P4, resulting in a reduction of the size of the paracone, i i Figure 6. Uintasorex sp., cf. U. montezumicus Lille- graven, LM„ LACM 132461, occlusal view. Scale = 0.5 mm. whereas in Dyseolemur and Washakius the meta- cone is lacking and the paracone is correspondingly larger. However, the P4 of Dyseolemur can be easily distinguished from those of Washakius by having a much weaker hypocone and better developed postprotoconal fold. Order PPrimates Family Microsyopidae Osborn and Wortman, 1892 Subfamily Uintasoricinae Szalay, 1969 Genus Uintasorex Matthew, 1909 Uintasorex sp., cf. U. montezumicus Lillegraven, 1976 Figure 6 REFFERRED SPECIMEN. LM„ LACM 132461. LOCALITY. LACM 5855. FAUNA AND AGE. Tapo Canyon Local Fauna, late Uintan. DISCUSSION. The LM, from locality LACM 5855 is morphologically very similar to those of U. montezumicus from the Mission Valley Formation, San Diego, California, including the following shared characters: (1) a vestigial paraconid that is posi- tioned very close to the metaconid with the apices of these cusps separated by a very small gap; (2) a sharply curved cristid connecting the protoconid with the paraconid and a curved cristid connecting the protoconid with the metaconid; (3) an enclosed trigonid basin; (4) a small hypoconulid positioned close to the entoconid; and (5) a deeply basined and completely rimmed talonid. The measurements of LACM 132461 are 0.88 mm A-P, 0.5 1 mm ANT- TR, and 0.63 mm POST-TR. The LM, from Simi Valley is assigned to U. sp., cf. U. montezumicus because it differs from those of U. montezumicus in its smaller size and by having the widths of the trigonid and talonid slightly small- er relative to the corresponding A-P lengths. The discovery of LACM 132461 represents the first rec- ord of Uintasorex from the Sespe Formation. Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 5 Figure 7. Miacis sp. undet., RM2, LACM 130826, oc- clusal view. Scale = 1 mm. Order Carnivora Bowdich, 1758 Family Miacidae Cope, 1880 Genus Miacis Cope, 1872 Miacis sp. undet. Figure 7 REFERRED SPECIMEN. RM„ LACM 130826. LOCALITY. LACM 5866. FAUNA AND AGE. Brea Canyon Local Fauna, late Uintan. DESCRIPTION. The trigonid of LACM 130826 is moderately elevated above the talonid indicating it is an M2. The trigonid shelf and the talonid basin are deep and the trigonid is open lingually. The cristid connecting the paraconid and protoconid possesses a distinct carnassial notch. A similar notch is present midway along the cristid that joins the metaconid and protoconid. The paraconid, pro- toconid, and metaconid are about equal in height. The entoconid and hypoconid are well developed, whereas the pre-entoconid, hypoconulid, and pre- hypoconid are only represented by very small cus- pules along the talonid cristid. The anterior and posterior labial cingulids are robust. A distinct peri- conid is present on the anterior labial cingulid. The measurements of LACM 130826 are 3.43 mm A-P, 2.74 mm ANT-TR, and 2.14 mm POST-TR. DISCUSSION. The RM2 (LACM 130826) from Simi Valley is most similar to those of Miacis, and it is referred to Miacis sp. undet. It differs from previously known miacids of the Sespe Formation (‘ Tapocyon occidentalis, Stock, 1934b; Procyon- dictis progressus (Stock, 1935b); and Miacis ? hook- wayi Stock, 1934b) by its much smaller size. Order Rodentia Bowdich, 1821 Family Ischyromyidae Alston, 1876 Reithroparamyinae, Wood, 1962 Genus Microparamys Wood, 1959 DISCUSSION. In a very significant paper, Korth (1984) presented a comprehensive review of the early Tertiary evolution and radiation of North American rodents that provided many new insights into rodent phylogenetic relationships. In this re- port, Korth (1984) reviewed the taxonomy of Mi- croparamys wherein he assigned two new species, M. scopaiodon and M. reginensis, to the genus. Korth noted that many species originally referred to Microparamys are now assigned to other genera. He recognized seven North American species of Microparamys: M. minutus (Wilson, 1937), M. tri- cus (Wilson, 1940a), M. dubius (Wood, 1949), M. perfossus Wood, 1974, M. sp. D ( = M. woodi n. sp., this paper), M. scopaiodon, and M. reginensis. In addition to these species, two informal taxa are referred to Microparamys : M. sp., cf. M. minutus from the Friars and Mission Valley Formations of the greater San Diego area of California (Lillegrav- en, 1977), and M. sp., cf. M. tricus from the Sespe Formation (this paper). Wood (1959) designated M. minutus from the Bridger Formation of Wyoming as the type species of Microparamys. The char- acters that define Microparamys have been ex- panded considerably with the addition of M. sco- paiodon and M. reginensis to the genus. Korth noted that M. scopaiodon and M. reginensis also exhibit many similarities to the middle Eocene sci- uravid Pauromys (family Sciuravidae). Korth (1984) revised the characters that define the Reithroparamyinae and included the following genera in this subfamily: Keith r op aramys Matthew, 1920, Acritoparamys Korth, 1984, Apatosciuravus Korth, 1984, Lophiparamys Wood, 1962, and Mi- croparamys. Korth provided evidence that the reithroparamyines were directly derived from an Asian ctenodactyloid rodent ancestor. In order to determine the phylogenetic relations of Micropar- amys to the other reithroparamyines and to deter- mine if Microparamys comprises a monophyletic clade, cladistic analyses were performed at the ge- neric and specific levels using Version 1.5 of the Hennig86 computer program (Farris, 1988) run on a 386 personal computer. The characters and char- acter states used in these analyses are presented in Appendix A. The Asian ctenodactyloid rodent Co- corays lingchaensis Li, Chiu, Yan, and Hsieh, 1979, was selected as the outgroup because it is the most primitive rodent known, making it the best taxon available for determining the plesiomorphic char- acter states used in the analyses. Cocomys is well known from numerous specimens and has been described in detail by Li et al. (1989). Korth (1984) assigned M. scopaiodon and M. reginensis to Microparamys based on the following shared characters of these species with primitive ischyromyids and Microparamys: (1) the M, an- terior cingulid is widely separated from the pro- toconid; (2) the lower molar anterior cingulid arises from the metaconid; (3) the lower molars lack a hypolophid that extends bucally from the ento- conid; and (4) the mandibular masseteric fossa ex- tends forward to a level about equal to those of Microparamys. Korth also used the above char- acters to differentiate M. scopaiodon and M. re- 6 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ! ginensis from Pauromys but recognized that most of these characters are just the plesiomorphic states for primitive ischyromyids. Korth cited the follow- ing similarities of M. scopaiodon and M. reginensis with Pauromys: (1) the lophs on the lower molars are of similar height; (2) the P4 is very reduced; (3) the lower molar mesoconid is anteroposteriorly compressed; (4) the lower molar entoconid is sep- arated from the posterolophid; and (5) the lower molar protoconid is isolated with the long arm of the protoconid extending to the base of the meta- conid and paralleling the anterior cingulid. Of these characters, separation of the entoconid from the posterolophid is the plesiomorphic state observed in ctenodactyloid rodents and cannot be used to establish phylogenetic relationships. In order to determine the phylogenetic relations of M. scopaiodon and M. reginensis to the sciuravid Pauromys and the reithroparamyines, a cladistic analysis including all of these taxa was performed. Character states were based primarily on those of the holotypes and topotypes of each genus. How- ever, in order to avoid using characters that exhibit ! high levels of parallelism or reversals in the analyses, attempts were also made to consider the character states of the least derived species of each genus. The generic character states were determined from the following species: Reithroparamys ctenodac- tylops Korth, 1984, Reithroparamys delicatissimus (Leidy, 1871), Acritoparamys atavus (Jepsen, 1937), Acritoparamys atwateri (Loomis, 1907), Acrito- paramys francesi (Wood, 1962), Lophiparamys murinus (Matthew, 1918), Lophiparamys woodi Guthrie, 1971, Apatosciuravus bifax Korth, 1984, I Microparamys minutus, Microparamys scopaio- don, Microparamys reginensis, Pauromys perditus Troxell, 1923, and Pauromys sp. from Powder Wash, Utah, described by Dawson (1968). Cladistic analysis of M. scopaiodon, M. reginen- sis, Pauromys, and the reithroparamyines was per- formed based on the character state matrix pre- sented in Appendix B. The analysis resulted in four equally parsimonious cladograms with lengths of 60 steps and consistency indices of 66 using un- weighted characters. The analysis was then repeated using successive weighting of characters, a proce- dure that has been shown to avoid the excessive weighting of multistate characters relative to binary characters and a means of basing groupings on more reliable characters without making prior decisions on weighting (Goldman, 1988; Farris, 1988). This procedure resulted in cladogram A of Figure 8 as the most parsimonious cladogram with a consis- tency index of 86. Therefore, cladogram A of Fig- ure 8 is considered to most likely represent the correct relations of the taxa. Based on cladogram A (Fig. 8), M. scopaiodon and M. reginensis are united by the following syn- apomorphies (node 6): (1) the M,_2 anterior cingulid is separated from the protoconid by a distinct groove in early wear that disappears with moderate wear to form a connection between the labial terminus Contributions in Science, Number 439 of the anterior cingulid and the protoconid; (2) the M,_, posterolophid is well developed; and (3) the M | .7 entoconid is isolated and separated from the posterolophid by a narrow distinct groove. In this analysis, Pauromys is the closest sister taxon to M. scopaiodon and M. reginensis, and these taxa are united by the following synapomorphies (Fig. 8, cladogram A, node 5): (1) the M1 2 metaconule is absent or very reduced; (2) the P4 is extremely re- duced relative to the lower molars; (3) a P4 ecto- lophid is absent; (4) the M,_, anterior cingulid is separated from the protoconid by a distinct groove or valley and the labial terminus of the anterior cingulid is not connected to the protoconid; (5) the M,_, mesoconid is anteroposteriorly compressed; and (6) the M,_2 ectolophids are usually absent. These data indicate that M. scopaiodon and M. reginensis are more closely related to the sciuravid Pauromys than to the reithroparamyines and strongly imply that M. scopaiodon and M. regi- nensis are not referable to Microparamys ( sensu stricto). Microparamys scopaiodon and M. regi- nensis are known only from fragmentary material and, until additional material is discovered that can provide a more complete analysis of these taxa, they cannot be confidently assigned to any genus or fam- ily and are herein referred to as “ Microparamys family incertae sedis. The analysis also indicates Apatosciuravus is the closest sister taxon to Pau- romys. Korth (1984) noted that Apatosciuravus shares many characters with Microparamys and sciuravids. Whether Apatosciuravus is a reithro- paramyine or sciuravid cannot be confidently de- termined based on the cladistic analysis presented here. Apatosciuravus is also known from fragmen- tary material and, until additional material of this taxon is discovered that can provide a more com- plete analysis, it is tentatively included in the Reith- roparamyinae. Based on the analysis presented above, the Reith- roparamyinae comprises the following genera: Reithroparamys, Acritoparamys, Microparamys, Lophiparamys, and Apatosciuravus. Cladistic analysis was performed on these genera using the character state matrix presented in Appendix C with the characters unweighted and weighted. This anal- ysis produced a single most parsimonious clado- gram with a length of 43 steps and a consistency index of 83 using unweighted characters and 96 using weighted characters (Fig. 9). The analysis in- dicates that Reithroparamys, Apatosciuravus, and Acritoparamys are successive sister taxa to Micro- paramys and Lophiparamys. The reithropara- myines are united by the following synapomorphies (Fig. 9, node 1): (1) the posterior margin of the anterior root of the zygoma is located in line with the anterior margin of the P4 to the center of the P4; (2) the M1 2 metaconule is lingually positioned; (3) the M1-2 hypocone is well developed; (4) the anterior termination of the masseteric fossa is lo- cated in line between the middle of the posterior half of the M2 to the middle of M2; (5) the P4 is Kelly and Whistler: Sespe Mammals ■ 7 .CO CO c CD .co co c 0) .co co c CD .CO CO c CD Figure 8. A-D. Four equally most parsimonious cladograms (length of 60 steps, consistency index of 66) of genera discussed in text produced using character state matrix presented in Appendix B with characters unweighted. With weighted characters, cladogram A becomes the most parsimonious (consistency index of 86). In cladogram A, Pauromys is closest sister taxon to “ Microparamys ” scopaiodon and “M.” reginensis, united at node 5 by following list of ancestral synapomorphies. Within the parentheses, number to left of colon denotes character number and to right of colon character state of hypothesized ancestor. Node 5: anterior termination of masseteric fossa is located in line between anterior end of M2 to posterior root of M, (2:4); M'~2 metaconule absent or very reduced (11:4-5); P4 extremely reduced in size relative to lower molars (16:3); P4 ectolophid absent (18:0); M,_, mesoconid anteroposteriorly compressed (24:1); M, 2 anterior cingulid separated from protoconid by distinct groove or valley and labial terminus of anterior cingulid not connected to protoconid (26:1); M, 2 ectolophids usually absent (36:1). molariform with two buc'cal cusps (paracone and metacone) and a distinct hypocone; (6) the P4 ec- tolophids are usually developed as complete cristids or almost complete cristids extending from the me- soconid to the protoconid and hypoconid; (7) the P4 hypoconid is well developed; (8) the P4 postero- lophid is well developed; (9) the P4 protoconid is usually absent or vestigial; and (10) the M,_2 pro- toconid is connected or nearly connected to ec- tolophid, and the posterior arm of protoconid is short or if developed is usually directed towards the middle of the metaconid or further forward, and the posterior arm of the protoconid is not parallel with the anterior cingulid. Additional syn- apomorphies noted by Korth (1984) that may unite the reithroparamyines, but not included in the cla- distic analysis because their character states are un- 8 ■ Contributions in Science, Number 439 known for several genera, are: (1) the auditory bul- lae are enlarged and cossified with the skull and (2) the posterior margin of the nasal bones extends further posteriorly than that of the premaxillaries. In this analysis, Microparamys and Lophiparamys are closest sister taxa based on the putative synapo- morphy (Fig. 9, node 4) of crenulated cheek teeth enamel. Based on the cladistic analysis, no autapo- morphies can presently be identified for the reith- roparamyines indicating they may represent a para- phyletic group. However, the lack of identifiable autapomorphies may also be due to the fact that many members of this group are poorly known. Removal of “M.” scopaiodon and “M.” regi- nensis from Microparamys results in the following species assigned to the genus: M. minutus ; M. sp., cf. M. minutus ; M. dubius\ M. tricus\ M. sp., cf. Kelly and Whistler: Sespe Mammals M. tricus ; M. perfossus ; and M. woodi n. sp. Cla- distic analysis of these species was performed using the character state matrix presented in Appendix D with the characters unweighted and weighted. Two equally parsimonious cladograms were produced using unweighted characters with lengths of 53 steps and consistency indices of 77 (Fig. 10A-B). With weighted characters, cladogram A (Fig. 10) becomes the most parsimonious with a consistency index of 92. Based on cladogram A (Fig. 10) the following suite of synapomorphies (node 1) are shared by the species of Microparamys and characterize the ge- nus: (1) the posterior margin of the anterior root of the zygoma is located in line with the anterior margin of the P4; (2) the anterior termination of the masseteric fossa is located in line with the middle of M,; (3) size is small; (4) the cheek teeth enamel is weakly to moderately crenulated; (5) the P4 me- soconid is weakly developed; (6) a P4 ectolophid is present; (7) the P4 hypoconid is well developed; (8) the P4 posterolophid is well developed; (9) the M,_2 anterior cingulid is well developed as a long distinct cristid; (10) the M,_2 anterior cingulid is separated from the protoconid by a distinct groove or valley and the labial terminus of the anterior cingulid is not connected to the protoconid; (11) the M,_2 posterolophid is well developed; (12) the MN2 hy- poconulid is usually present as a moderately well- developed, often elongated, cuspule formed along the posterolophid; and (13) the M,_2 entoconid is isolated and separated from the posterolophid by a narrow distinct groove. The only autapomorphic characters identified that indicate Microparamys is a monophyletic clade are the forward positioning of the zygoma and masseteric fossae and the distinct arrangement of the anterior cingulid and the pro- toconid (numbers 1, 2, and 9, above). Three species groupings are indicated by the analysis: (1) the mi- nutus group, including M. minutus and M. sp., cf. M. minutus ; (2) the woodi group, including M. woodi and M. perfossus ; and (3) the tricus group, including M. tricus, M. sp., cf. M. tricus, and M. dubius. The synapomorphy (node 2) that unites the minutus group is a strongly developed P4 meso- conid. The synapomorphies (node 4) that unite the woodi group are as follows: (1) the occlusal patterns in the Mu2 protoconal and hypoconal regions are U-shaped; (2) an M2 mesolophid is present as a distinct crest originating near the apex of the pro- tocone and extending lingually into the central ba- sin of the tooth; (3) an M,_2 hypoconulid is lacking; and (4) the M,., entoconid is connected to the posterolophid. The synapomorphies (node 5) that unite the tricus group are: (1) the M1 2 postproto- crista is absent or poorly developed and (2) a small distinct cuspule is developed on the labial terminus of the M,_ 2 anterior cingulid. The cladistic analyses presented here are regard- ed as preliminary because many of the taxa are poorly known, the sample sizes are small, and the intraspecific variation of certain characters is not Figure 9. Single most parsimonious cladogram (length of 43 steps, consistency index of 83 with unweighted characters and 96 with weighted characters) produced for genera comprising the Reithroparamyinae using character state matrix presented in Appendix C. The cladogram is supported by the following list of ancestral synapomor- phies. Within the parentheses, number to left of colon denotes character number and to right of colon character state of hypothesized ancestor. Node 1: posterior margin of anterior root of zygoma located in line with center of P4 (1:1); anterior termination of masseteric fossa is located in line between posterior half to middle of M2 (2:1); P4 molariform with two buccal cusps (paracone and meta- cone) and distinct hypocone (7:1); M1 - metaconule po- sitioned lingually and not in close association with me- tacone (12:1); M1 ’ hypocone well-developed cusp (14:1); P4 slightly reduced relative to lower molars (16:1); P4 ec- tolophid present, weakly developed (18:1); P4 hypoconid well developed (19:1); P4 posterolophid present, well de- veloped (20:1); P4 protoconid usually absent or vestigial (21:1); Mi2 protoconid not isolated (30:0); M, 2 ectolo- phids usually present with one or both cristids complete or nearly complete (36:0). Node 2: M1 4 metaconule sin- gle, moderately reduced cusp (1 1:1); M,_, anterior cingulid moderately to well-developed cristid connected to meta- conid (27:1); M,_, hypoconulid present as moderately well developed, often elongated, cuspule formed along pos- terolophid (34:1). Node 3: anterior termination of mas- seteric fossa is located in line between middle of NT to center of anterior half of M, (2:2-3); M, 2 posterolophid well developed (32:1); M, , entoconid isolated and sep- arated from posterolophid by narrow distinct groove (35: 1). Node 4: cheek teeth weakly to moderately crenulated (4:1); M1 - protoconule moderately reduced as elongated cusp (10:1); M,_, anterior cingulid separated from pro- toconid by distinct groove or valley and labial terminus of anterior cingulid not connected to protoconid (26:1). adequately known, especially quantitative charac- ters. However, whenever possible, an attempt was made not to use characters that have been shown by other investigators to be highly variable (e.g., Lillegraven, 1977). The analyses presented here help to clarify the characterization of the genus Micro- paramys and provide a basis for future phyloge- Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 9 Figure 10. A-B. Two equally most parsimonious cladograms (length of 53 steps, consistency index of 77) of species of Microparamys produced using character state matrix presented in Appendix D with characters unweighted. With weighted characters, cladogram A becomes the most parsimonious with a consistency index of 92. Cladogram A is supported by the following list of ancestral synapomorphies. Within the parentheses, number to left of colon denotes character number and to right of colon character state of hypothesized ancestor. Node 1: posterior margin of anterior root of zygoma is located in line with anterior margin of P4 (1:2); anterior termination of masseteric fossa is located in line with middle of M, (2:5); cheek teeth weakly to moderately crenulated (4:1); size small (5:1); P4 molariform with two buccal cusps (paracone and metacone) and distinct hypocone (7:1); M12 metaloph and protoloph incomplete (9: 1); M'~2 single moderately reduced metaconule usually present as relatively distinct cusp (11:1); P4 mesoconid present, weakly developed (17:1); P4 ectolophid usually developed as a complete cristid or almost complete cristid extending from mesoconid to protoconid and hypoconid (18:1); P4 hypoconid well developed (19:1); P4 posterolophid well developed (20:1); M,_, anterior cingulid well developed as long, distinct cristid (25:1); M,_, separated from protoconid by distinct groove or valley and labial terminus of anterior cingulid not connected to protoconid (26:1); M,., posterolo- phid well developed (32:1); M,_, hypoconulid present as moderately well-developed, often elongated, cuspule formed along posterolophid (34:1); M,_, entoconid isolated and separated from posterolophid by narrow distinct groove (35: 1). Node 2: P4 mesoconid strongly developed (17:2). Node 3: M,_2 anterior cingulid separated from protoconid by distinct groove or valley and connected to protoconid by small but relatively persistent thin cristid originating from near labial terminus of anterior cingulid (26:3). Node 4: M'~2 occlusal patterns U-shaped in hypoconal and protoconal regions (8:1); M1-2 metaconule doubled as two small reduced cuspules (11:2); M2 mesolophid present (15:1); M,„2 hypoconulid absent (34:3); M,_, entoconid connected to posterolophids (35:2). Node 5: M1-2 postprotocrista absent or poorly developed (13:0); M,_, labial terminus of anterior cingulid cuspate (28:1). Node 6: M1-2 metaconule doubled as two well-developed cuspules (11:3); P4 slightly reduced relative to lower molars (16:1); P4 protoconid usually absent or vestigial (21:1). netic investigations of this little-known group of rodents. Microparamys woodi new species Figure 11, Tables 2 and 3 Paramys cf. minutus Wilson, 1940a:72, pi. 1, figs. 10-11. Microparamys sp. D Wood, 1962: 165-166, figs. 55G-H. HOLOTYPE. RM2, LACM 130820. TYPE LOCALITY. LACM 5857. HYPODIGM. LdP4, LACM 133988; 3RP4s, LACM 130821, 132415, 132463; LP4, LACM 132414; LM\ LACM 130819; 2RM's, LACM 132422, 133989; 3LM2s, LACM 132416, 132464, 132467; RM2, LACM(CIT) 2155; LP4, LACM 130822; 4RP4s, LACM 130823, 132417, 132466, 133990; 2RM,s, LACM 130818, 133991; 2LM,s, LACM 132471, 132651; 2LM,s, LACM 130824, 132472; RM,, LACM(CIT) 2156; LM„ LACM 132468. DISTRIBUTION, FAUNAS, AND AGE. Lo- calities LACM 5661, 5855, 5857, 5860, 6081, lo- cality LACM(CIT) 180; Tapo Canyon and Brea Canyon Local Faunas; late Uintan. ETYMOLOGY. Named in honor of Albert E. Wood, formerly of the Amherst College, in rec- ognition of his contributions to our understanding of Microparamys. DIAGNOSIS. Microparamys woodi is distin- guished by having the following suite of characters: (1) small ischyromyid; (2) cheek teeth enamel cren- ulated; (3) upper cheek teeth with protolophs and metalophs usually complete, occlusal patterns in the protoconal and hypoconal regions U-shaped, metaconule often doubled as two small cusps on metaloph, and tendency for development of small accessory crest originating from protoloph and ex- tending anteriorly towards the anterior cingulum that occasionally results in division of valley be- tween the anterior cingulum and protoloph into labial and lingual portions; (3) upper molars with mesostyles and from one to two mesolophs; (4) P4 with moderately developed protoconid, weakly de- veloped metaconid, and well-developed postero- lophid; and (5) lower molars with metalophid com- plete, “mesolophids” present, posterolophid well developed and connected to entoconid, valley be- i Kelly and Whistler: Sespe Mammals 10 ■ Contributions in Science, Number 439 G H I i 1 Figure 11. Microparamys woodi new species, A. LdP4, LACM 133988; B. RP4, LACM 132415; C. LM1, LACM 130819; D. holotype, RM-, LACM 130820; E. RP4, LACM 130823; F. RM„ LACM 130818; G. LM,, LACM 130824; H. LM,, LACM 131472; I. LM3, LACM 132468. All occlusal views. Scale = 1 mm. tween anterior cingulid and protoconid moderately developed with small accessory cristid extending across valley connecting protoconid and anterior cingulid, and hypoconulids absent. Differs from Mi- croparamys minutus (Wilson, 1937) and M. sp., cf. M. minutus (Lillegraven, 1977) by having the fol- lowing characters: (1) teeth larger; (2) M1-2 with mesostyles present, mesolophs more strongly de- veloped, and connection of anterior crest of hy- pocone to metaloph stronger; and (3) lower molar hypolophids more developed. Differs from M. tri- cus (Wilson, 1940a) by having the following char- Contributions in Science, Number 439 acters: (1) teeth smaller; (2) M'~2 with relatively smaller mesostyles, less developed and bulbous protoconules and metaconules, less tendency for multiplication of metaconules, greater tendency for multiplication of mesolophs, and greater tendency for development of an accessory crest uniting pro- toloph and anterior cingulum; and (3) lower molars with slightly less separation of protoconid and an- terior cingulid. Differs from M. dubius (Wood, 1949) by having the following characters: (1) P4 with bet- ter developed metaconule, a mesostyle, and larger hypocone; (2) M12 with lingually directed crests Kelly and Whistler: Sespe Mammals ■ 11 Table 2. Measurements (in mm) of upper dentition of Microparamys woodi new species. N Tooth/ dimension O.R. Mean S.D. c.v. 1 dP4 A-P 1.29 1 ANT-TR 1.07 1 POST-TR 1.30 4 P4 A-P 1.15-1.24 1.21 4 ANT-TR 1.38-1.54 1.46 4 POST-TR 1.33-1.45 1.39 3 M1 A-P 1.33-1.42 1.38 3 ANT-TR 1.45-1.55 1.51 3 POST-TR 1.40-1.47 1.44 5 M2 A-P 1.35-1.48 1.42 0.05 3.5 5 ANT-TR 1.58-1.73 1.63 0.06 3.7 4 POST-TR 1.44-1.56 1.49 from mesostyles and U-shaped protocones and hy- pocones; and (3) lower molars with “mesolophids” present and hypolophids that are less developed and posterolabially directed. Differs from M. perfossus Wood, 1974, by having the following characters: (1) P4 and M1 hypocones less separated from pro- tocones; (2) M1-2 with better developed metalophs and greater tendency for multiplication of meta- conules; (3) P4 less anteroposteriorly elongated and protoconid smaller; and (4) lower molars with pos- terolabially directed hypolophids and lacking long cristid from entoconid toward hypoconid. Differs from “M.” reginensis Korth, 1984, by having the following characters: (1) teeth much larger; (2) low- er molar lingual extension of mesoconid lacking and mesoconid not anteroposteriorly compressed; (3) occlusal outline of M3 less rectangular; and (4) enamel more crenulated in basins. Differs from “M.” scnpaiodon Korth, 1984, by having the following characters: (1) P4 much larger, and larger relative to molars; (2) P4 occlusal pattern more complex and protoconid more reduced; (3) lower molars with mesoconids not anteroposteriorly com- pressed, lacking a long posterior arm of protoconid extending to base of metaconid, and entoconids connected to posterolophids; and (4) enamel more crenulated in basins. DESCRIPTION. Only one deciduous P4 of M. woodi has been recovered from the Sespe Forma- tion. The dP4 is subtriangular in occlusal outline and moderately molariform. The protocone is the largest primary cusp, and the metacone and para- cone are about equal in size. A thin crest is present along the labial aspect of the tooth that connects the metacone with the paracone. The preproto- crista extends anterolabially to form a protoloph and the postprotocrista extends posterolabially to form a metaloph. Two small distinct metaconules are present along the crest of the metaloph. The anterior cingulum is robust. A very low accessory crest is present between the protoloph and the an- terior cingulum that divides the valley separating Table 3. Measurements (in mm) of lower dentition of Microparamys woodi new species. N Tooth/ dimension O.R. Mean S.D. C.V. 5 P4 A-P 1.24-1.41 1.34 0.07 5.5 5 ANT-TR 0.97-1.08 1.05 0.05 4.7 5 POST-TR 1.18-1.39 1.30 0.09 6.8 3 M, A-P 1.40-1.46 1.43 4 ANT-TR 1.30-1.51 1.41 3 POST-TR 1.39-1.49 1.44 3 M, A-P 1.44-1.58 1.52 3 ANT-TR 1.42-1.55 1.49 3 POST-TR 1.49-1.58 1.52 1 M, A-P 1.47 1 ANT-TR 1.35 1 POST-TR 1.21 these structures into labial and lingual portions. The posterior cingulum is well developed and extends lingually from the labial aspect of the metacone to the level of the protocone wherein it terminates in a well-defined bulge or hypocone. The P4 is subquadrate in occlusal outline with the protocone and hypocone separated by relatively deep valleys. A small distinct mesostyle is present between the paracone and metacone. One or two metaconules may be present as small cusps or bulg- es on the metaloph. The protoconule is usually a distinct small bulge on the protoloph. The proto- loph is a complete crest except for a small cleft about midway along the protoloph at the point where a small accessory crest projects anteriorly from the protoloph towards the anterior cingulum. The metaloph is a low complete crest. The prepro- tocrista, postprotocrista, and the protocone form a U-shaped wear pattern in the protoconal region. A mesoloph is absent in all known P4s. The anterior cingulum is well developed and a slight enlargement is present at the lingual terminus. The posterior cingulum is well developed and extends lingually from the posterolabial aspect of the metacone to terminate in a well-developed hypocone. A crest from the hypocone extends anterolabially to ter- minate just short of the middle of the metaloph and this crest along with the posterior cingulum and the hypocone results in a U-shaped wear sur- face in the hypoconal region. The enamel is mod- erately crenulated in the basins of the tooth. The first two upper molars are similar in mor- phology to the P4 except for the following differ- ences. The occlusal outline is more quadrate in shape. The mesostyle has a small spur that extends lingually into the central basin of the tooth. The parastylar region is larger or more inflated. Meso- lophs are always present and may vary from a single crest to two separate crests that extend labially from the apex of the protocone, between the prepro- tocrista and postprotocrista, into the central basin to terminate about half way towards the labial as- 12 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals pect of the tooth. The anterior cingulum is more developed, and a lingual terminal bulge of the an- terior cingulum may be present or absent. The ac- cessory crest that originates from the protoloph occasionally extends anteriorly to join with the an- terior cingulum; this results in a division of the valley between the anterior cingulum and the pro- toloph into labial and lingual portions. However, this accessory crest is more commonly absent, or it only forms a partial divider that extends anteriorly from the protoloph a very short distance into the intervening valley. The enamel is more crenulated in the basins of the first two upper molars. The P4 is subrectangular in occlusal outline. The metaconid is the largest and tallest cusp. A small cristid extends posteriorly from the apex of the metaconid towards the entoconid where it termi- nates just anterior to the entoconid. The proto- conid is a low bunodont cusp and the smallest primary cusp. The metalophid is a small low cristid that connects the metaconid and protoconid. The mesoconid is weakly developed as a small rounded bulge or cusp, and a mesolophid is lacking. The entoconid is slightly taller than the hypoconid, and these cusps are connected by a complete, posteri- orly convex posterolophid. The anterior cingulid is a convex cristid extending from the metaconid to the anterolabial aspect of the protoconid. The metalophid is a small cristid originating from the apex of the protoconid and extending anterolin- gually towards the protoconid. The enamel is usu- ally crenulated in the basins of the tooth. The first two lower molars have rectangular oc- clusal outlines. Incipient metastylids may be present as very small cuspules on the lingual cristid con- necting the metaconid and entoconid. The metalo- phid is a complete low cristid that connects the metaconid and protoconid. The mesoconids are well developed. “Mesolophids” are usually present and can vary from a single to as many as three low spurs extending lingually from the mesoconid into the central basin of the tooth. A small short spur is present that extends labial ly from the entoconid into the basin of the tooth. The hypolophid is usu- ally a short posterolabially directed cristid. The an- terior cingulid extends lingually front the base of the metaconid to the level of the protoconid and a moderately developed valley separates the ante- rior cingulid from the protoloph and protoconid. A small accessory cristid extends from the proto- conid across this valley to the anterior cingulid. The posterolophid is well developed and connects the hypoconid with the entoconid. The enamel is cren- ulated in the basins of the tooth. Only two M,s of M. woodi have been recovered during the impact mitigation program. The M?s are morphologically very similar to the first two lower molars except that they have more robust and curved posterolophids. Both M3s lack incipient mesosty- lids, but this may not be a diagnostic character for the M, because the occurrence of mesostylids is variable in the first two lower molars. DISCUSSION. Wilson (1940a) referred two iso- lated teeth, an upper and a lower molar, from the Sespe Formation at locality LACM(CIT) 180 to Paramys cf. minutus. Wood (1962) reevaluated the phylogenetic status of these two teeth and assigned them to an informal taxon, Microparamys sp. D, noting that they represented a very distinct species of Microparamys. Lillegraven (1977), in agreement with Wood, regarded the teeth of M. sp. D a new species but, like Wood, was reluctant to formally name a new taxon until a better sample was avail- able from the Sespe Formation. The new material of M. sp. D discovered during the program at the Simi Valley Landfill has resulted in a better char- acterization of this taxon and allows the assignment of this sample to the new species M. woodi. Because of the larger sample of M. woodi now available, the intraspecific variation of the teeth can be evaluated. The two most variable characters of the teeth of M. woodi are: (1) the degree of the development of an accessory crest extending from the protoloph to the anterior cingulum (= anterior cingulum-protoloph crest) and (2) the development of mesolophs and “mesolophids” in the upper and lower molars, respectively. The anterior cingulum- protoloph crest is usually absent or represented only by a partial crest extending anteriorly from the pro- toloph part way across the transverse valley that separates the anterior cingulum from the proto- loph. Occasionally, the anterior cingulum-proto- loph crest is complete across the intervening valley and connects the anterior cingulum with the pro- toloph, resulting in a division of the valley into labial and lingual portions. Walsh (1987) also found the occurrence of anterior cingulum-protoloph crests to be highly variable in a sample of Micro- paramys teeth from the Mission Valley Formation of the San Diego area in California. In the upper molars, one or two mesolophs may be present, ex- tending labially from the apex of the protocone into the central transverse valley. In the lower mo- lars, from one to three “mesolophids” may be pres- ent as separate low cristids extending lingually from the mesoconid into the central basin of the tooth. Many investigators have discussed in detail the probable relations of M. woodi (= M. sp. D) to other species of Microparamys (Wilson, 1940a; Wood, 1962, 1974; Dawson, 1966, 1974; Lille- graven, 1977; Korth, 1984; Walsh, 1987). Wood (1974) regarded M. woodi as most similar to M. perfossus of the Porvenir Local Fauna, Vieja-Oji- naga area, Texas, wherein both species have U-shaped upper molar occlusal patterns in the pro- toconal and hypoconal regions. Additional shared characters of M. woodi and M. perfossus indicating a close relationship are as follows: (1) the M2 me- soloph is well developed; (2) the M,_2 entoconid is connected to the posterolophid; and (3) the M,_2 hypoconulid is absent or vestigial. Even though the above synapomorphies unite M. woodi and M. per- fossus into a species group, M. woodi can be easily distinguished from M. perfossus by the following Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 13 Figure 12. Microparamys sp., cf. M. tricus (Wilson), A. LM2, LACM 130809, B. LM„ LACM 130810; C LM2, LACM 130816; D. RM3, LACM 130814. All occlusal views. Scale = 1 mm. differences: (1) the P4 and M1 hypocones are less separated from the protocones; (2) the M12 meta- lophs are more developed with a greater tendency for multiplication of the metaconules; (3) the P4 is less anteroposteriorly elongated with a relatively smaller protoconid; and (4) the lower molar hy- polophids are less developed and extend postero- labially. Microparamys sp., cf. M. tricus (Wilson, 1940) Figure 12, Table 4 REFERRED SPECIMENS. RM1, LACM 130812; LM2, LACM 130809; LM„ LACM 130810; RM,, LACM 130815; LM„ LACM 130816; RM3, LACM 130814. LOCALITIES. LACM 5866, 5869. FAUNA AND AGE. Brea Canyon Local Fauna, late Uintan. DISCUSSION. The Microparamys teeth from localities LACM 5866 and 5869 are morphologi- cally very similar to those of M. tricus from the early Duchesnean Pearson Ranch Local Fauna of the Sespe Formation. However, these teeth differ from those of M. tricus by having the following characters: (1) the teeth are smaller; (2) the M2 post- protocrista is much more weakly developed and separated from the metaconal region by a labial continuation of the valley that separates the pro- tocone from the hypocone; (3) the lingually di- rected crest from the mesostyle in the upper molars is slightly less developed; (4) the M2 hypolophid is slightly less robust; (5) the M,„, anterior cingulid has a larger, more prominent cuspule at its labial termination; and (6) the M3 postprotocristid is less developed. The teeth of M. sp., cf. M. tricus are distinguished from those of M. woodi by having the following characters: (1) M'~2 with larger me- sostyles, more bulbous protoconules and metacon- ules, a greater tendency for multiplication of the metaconules, less tendency for multiplication of mesolophs, and usually lacking the low crest that unites the protocone and the anterior cingulum and (2) lower molars with slightly greater separation of the protoconid and the anterior cingulid. The samples of Microparamys teeth from the late Uintan Brea Canyon Local Fauna are slightly less derived, as indicated by their less developed postprotocristae, postprotocristids, mesostylar lin- gual crests, and hypolophids, than those of M. tri- cus and are therefore assigned to M. sp., cf. M. tricus. Lillegraven (1977) assigned 10 isolated teeth from the latest Uintan or earliest Duchesnean Camp San Onofre Local Fauna of the PSantiago Formation, Camp Pendleton Marine Corps Base, California, to M. tricus. Lillegraven noted that even though the Camp San Onofre samples of Microparamys teeth are consistently smaller, they are indistinguishable morphologically from those of M. tricus from the Pearson Ranch Local Fauna. The teeth of M. sp., cf. M. tricus are the same size as those of M. tricus from the Camp San Onofre Local Fauna but have less developed postprotocristae, postprotocristids, mesostylar lingual crests, and hypolophids. These character states indicate that M. sp., cf. M. tricus is less derived than M. tricus from the Camp San Onofre Local Fauna. A single evolutionary lineage is probably represented by the Microparamys sam- ples discussed above, wherein the late Uintan M. sp., cf. M. tricus from the Brea Canyon Local Fauna was probably ancestral to the latest Uintan or ear- liest Duchesnean M. tricus from the Camp San On- ofre Local Fauna, and the latter then gave rise to the larger early Duchesnean M. tricus from the Pearson Ranch Local Fauna. Genus Leptotomus Matthew, 1910 Leptotomus sp. undet. Figure 13 REFERRED SPECIMEN. LM3, LACM 131465. LOCALITY. LACM 5876. FAUNA AND AGE. Simi Valley Landfill Local Fauna, late Duchesnean. DESCRIPTION. The LM3 is well worn, antero- posteriorly elongated, and the occlusal outline is subrectangular. The labial surface of the protocon- id is missing. The anterior cingulid (= anterolophid) is a robust cristid that connects the metaconid with the protoconid. A small metalophid is present and projects lingually from the protoconid into the an- terior basin of the tooth. The hypolophid is a con- tinuous cristid between the entoconid and the hy- poconid. The ectolophid is complete and markedly curved. The posterior cingulid (= posterolophid) is robust and extends lingually from the posterolabial aspect of the tooth to the entoconid, where it is only separated from the entoconid by a small cleft. The measurements of LACM 131465 are 3.06 mm A-P, 2.66 mm ANT-TR (estimated), and 2.67 mm POST-TR. DISCUSSION. The Simi Valley tooth (LACM 131465) appears most similar to those of Lepto- tomus guildayi Black, 1971, and L. sp., near L. 14 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals Table 4. Measurements (in mm) of specimens of Mi- croparamys sp., cf. M. tricus. Specimen Tooth/dimen- sion LACM 130812 RM1 A-P 1.55 (estimated) TR — LACM 130809 LM2 A-P 1.69 TR 1.83 LACM 130810 LM, A-P 1.69 ANT-TR 1.50 POST-TR 1.71 LACM 130815 RM, A-P 1.54 ANT-TR 1.43 POST-TR 1.59 LACM 130816 LM2 A-P 1.75 ANT-TR 1.68 POST-TR 1.77 LACM 130814 RM, A-P 1.89 ANT-TR 1.71 POST-TR 1.50 guildayi (Black, 1971) of the Badwater Localities 5, 6, and 15 from the ? Wagon Bed Formation of Wyoming by having the following shared charac- ters: (1) the metalophid extends from the proto- conid into the central basin of the tooth but does not connect with the metaconid; (2) the hypolophid is complete; (3) the posterolophid is prominent and separated only slightly from the entoconid; and (4) the ectolophid is markedly curved. The Simi Valley tooth differs from those of L. guildayi and L. sp., near L. guildayi by its smaller size and by having a slightly less developed metalophid and a vestigial mesoconid. The tooth from Simi Valley also exhibits some characters that are similar to those of species of Rapamys, including a prominent posterolophid, markedly curved ectolophid, incomplete metalo- phid, and complete hypolophid. However, the Simi Valley tooth differs from those of Rapamys by lacking the wide separation of the posterolophid and the entoconid. The Simi Valley specimen appears to be most closely related to L. guildayi and L. sp., near guil- dayi. However, due to the lack of adequate ma- terial for species identification, it is herein assigned to Leptotomus species undetermined. Family PIschyromyidae Genus Eohaplomys Stock, 1935a Eohaplomys matutinus Stock, 1935a Figure 14 REFERRED SPECIMEN. RM ', and partial RM2, LACM 130827. LOCALITY. LACM 5616. FAUNA AND AGE. Brea Canyon Local Fauna, late Uintan. i 1 Figure 13. Leptotomus sp. undet., LM„ LACM 131465, occlusal view. Scale = 1 mm. DISCUSSION. Kelly et al. (1991) questionably assigned LACM 130827 to an undetermined di- chobunid. Further preparation and examination of this specimen indicates that it is referrable to Eo- haplomys matutinus. The measurements of the RM 1 of LACM 130827 are 4.21 mm A-P and 4.88 mm TR. Superfamily Geomyoidea Weber, 1904 Family PGeomyidae Gill, 1872 Genus Griphomys Wilson, 1940b Griphomys alecer Wilson, 1940b Figure 15, Tables 5 and 6 REFERRED SPECIMENS. In addition to those referred elsewhere (Lillegraven, 1977); 2?RdP4s, LACM 130801, 132427; 2?LdP4s, LACM 130776, 132428; partial left maxilla with P4 and partial M', LACM(CIT) 2525; LP4, LACM 132432; 2RM‘s, LACM 130777, LACM(CIT) 2526; LM1, LACM 130781; 3RM2s, LACM 130778, 132425, 132429; 2LM2s, LACM 130779, 132431; LM3, LACM 132462; holotype, partial right dentary with P4-M2, LACM(CIT) 2522; partial right dentary with P4 and M„ LACM(CIT) 2524; 3RP4s, LACM 130782, 130784, 130789; 2LP4s, LACM 130783, 133993; Figure 14. Eohaplomys matutinus Stock, RM 1 and par- tial RM2, LACM 130827, occlusal views. Scale = 1 mm. Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 15 A M B J C D N O Figure 15. Griphomys alecer Wilson, A. RdP4, LACM 132428; B. LP4, LACM 132432; C. LM1, LACM 130781; D. RM2, LACM 130778; E. LM2, LACM 130779; F. LM2, LACM 132431; G. LM\ 132462; H. RP4, LACM 130782; I. LP4, LACM 130783; J. RP4, LACM 130784; K. RM„ LACM 130785; L. LM,, LACM 130774; M. LM,, LACM 130775; N. RM„ LACM 130780; O. RM„ LACM 130787; P. RM„ LACM 130790. All occlusal views. Scale = 1 mm. RM„ LACM 130785; 2LM,s, LACM 130788, 132470; partial right dentary with M,, LACM(CIT) 2523; 4LM,s, LACM 130774, 130775, 132424, 132426; 2RM,s, LACM 130780, 132430; 3RM,s, LACM 130786, 130787, 130790; LM„ UCMP 79479. LOCALITIES. Sespe Formation, LACM(CIT) 150, 202, 207, LACM 5612, 5616, 5661, 5855, 5857, 5859, 5860, 5866, 5868, 5869; PSantiago Formation, UCMP V-72088. FAUNAS AND AGE. Tapo Canyon, Brea Can- yon, and Pearson Ranch Local Faunas, Simi Valley, and Camp San Onofre Local Fauna, Camp Pendle- ton Marine Corps Base, California; late Uintan to early Duchesnean. DESCRIPTION. Four teeth (LACM 130776, 130801, 132427, 132428), tentatively assigned to dP4s, were recovered during the program from lo- calities LACM 5616, 5661, and 5860. These teeth are morphologically very similar to the permanent upper cheek teeth except for being slightly smaller and relatively narrower transversely, and by having the anterior cingulum usually not extending as far lingually. The permanent cheek teeth of Griphomys alecer have been well described by Wilson (1940b), Lind- say (1968), and Lillegraven (1977). Flowever, due to the small number of teeth in the original samples of G. alecer, additional details on intraspecific vari- ation are described below. All of the upper molars have the central trans- verse valleys open labially and lingually, and all are lacking a preprotocrista. However, one upper molar has a very small mesostyle present between the paracone and metacone, but this cusp is not high enough to block the labial opening of the central transverse valley. A small anterocone is usually pres- ent as a small cusp along and near the lingual ter- mination of the anterior cingulum. The most vari- able character of the upper molars is the development of a protoloph spur, which usually comprises a small crest that originates along the posterior face of the protoloph near its connection with the protocone and extends posteriorly into ^■Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals Table 5. Measurements (in mm) of upper dentition of Griphomys alecer. N Tooth/ dimension O.R. Mean S.D. c.v. 4 dP4 A-P 0.96-1.04 1.03 4 ANT-TR 0.85-1.05 0.96 4 POST-TR 0.83-1.05 0.94 2 P4 A-P 1.14-1.22 1.18 2 ANT-TR 1.08-1.21 1.15 2 POST-TR 1.12-1.16 1.14 4 M' A-P 1.12-1.30 1.20 3 ANT-TR 1.19-1.26 1.24 3 POST-TR 1.15-1.30 1.20 8 M2 A-P 1.11-1.24 1.17 0.05 4.2 8 ANT-TR 1.19-1.42 1.27 0.08 6.0 8 POST-TR 1.13-1.28 1.17 0.05 4.3 2 M’ A-P 1.01-1.05 1.03 2 ANT-TR 1.17-1.18 1.175 2 POST-TR 1.04-1.10 1.07 the central transverse valley of the tooth where it then turns labiad for a short distance. Lillegraven (1977) identified this structure as a low posterior protocrista. Whether this structure is homologous with a postprotocrista or an anterior ectoloph (= anterior mure) and a vestigial mesoloph is uncertain. Nevertheless, the following variations of this spur are exhibited by the Simi Valley upper molars of G. alecer : (1) the spur is completely lacking; (2) the spur is very small and only extends a very short distance posterolabially where it terminates at the posterior base of the protocone and does not ex- tend into the central transverse valley; (3) the spur extends posterolabially from the protoloph about midway into the central transverse valley and re- sembles a vestigial mesoloph; and (4) the spur forms a small cristid that originates at the base of the protoloph, just labial to its connection with the protocone, and then extends posterolabially in a curved fashion into the central transverse valley resembling a small mesoloph. No distinct cusps or swellings are exhibited on any of the spurs that could be regarded as mesocones. In the P4 the most variable character is the de- velopment of the ectolophid and its associated me- soconid. Mesoconids, which may be represented by a distinct isolated cusp in the central transverse valley or by a distinct swelling at the middle of the ectoloph, are usually present. The mesoconid-ec- tolophid structure varies in the Simi Valley P4s as follows: (1) ectolophid incomplete, mesoconid ab- sent, ectolophid represented by a small cristid or loph that projects anteriorly from the hypolophid into the middle of the central transverse valley and terminates about halfway across this valley; (2) ec- tolophid complete, mesoconid absent, ectolophid represented by a small cristid that projects anteri- orly from the hypolophid across the transverse cen- tral valley and connects with the metalophid; (3) Table 6. Measurements (in mm) of lower dentition of Griphomys alecer. N Tooth/ dimension O.R. Mean S.D. C.V. 7 P4 A-P 0.96-1.14 1.08 0.07 6.0 6 ANT-TR 0.71-0.83 0.77 0.04 5.5 6 POST-TR 0.91-1.05 0.99 0.05 5.3 4 M, A-P 1.13-1.19 1.16 4 ANT-TR 0.91-1.04 0.98 4 POST-TR 0.97-1.19 1.05 11 M, A-P 1.18-1.35 1.23 0.05 4.2 11 ANT-TR 0.97-1.25 1.14 0.08 6.9 11 POST-TR 1.00-1.23 1.13 0.07 6.2 4 M, A-P 1.07-1.23 1.15 4 ANT-TR 0.98-1.07 1.01 4 POST-TR 0.86-0.90 0.88 ectolophid complete, mesoconid present, ectolo- phid represented by a complete cristid extending anteriorly from the hypolophid across the central transverse valley where it is connected with the metalophid, and the mesoconid is a distinct bulge or swelling midway along the cristid; and (4) ec- tolophid absent, but an isolated well-developed, triangular-shaped mesoconid is present with one apex of the triangle connected to the middle of the anterior face of the hypolophid. The central trans- verse valley is usually open labially; however, in two premolars a slight bulge, which may represent a vestigial ectostylid, is present midway along a weakly developed cristid at the labial aspect of the tooth between the protoconid and the hypoconid. This bulge does not represent a mesoconid because in both of these premolars a mesoconid is present in the central transverse valley. The lower molars vary in the development of cristids and cusps in the central transverse valleys as follows: (1) cusps or cristids absent; (2) a distinct isolated cusp or mesoconid in the central transverse valley with cristids absent; and (3) an incomplete ectolophid, which may be a distinct cristid or very low indistinct cristid that originates on the anterior face of the hypolophid, lingual to the hypoconid, and extends midway anterolingually or anteriorly into the central transverse valley. In the teeth ex- hibiting an incomplete ectolophid, a distinct me- soconid usually cannot be differentiated from the ectolophid. In one tooth (LACM 130774), two cris- tids originate on the anterior face of the hypolophid and extend anteriorly into the central transverse valley. The most labial cristid appears to represent a partial ectolophid with a small cusp or mesoconid at its termination. The lingual cristid does not ex- hibit a cusp, and this cristid may be a duplication of the ectolophid. Another lower molar (LACM 130775) is also distinctive in that it exhibits a small bulge or cuspule (= Pectostylid) at the posterolabial base of the protoconid and a thin low cristid (= Plabial cingulid) that extends posteriorly from this Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 17 cuspule to join with the anterolabial base of the hypoconid. DISCUSSION. Many investigators have dis- cussed the taxonomic relations of Gripbomys even though the fossil record of this genus was previ- ously very sparse (Wilson, 1940b, 1949; Lindsay, 1968; Emry, 1972; Wood, 1974; Sutton and Black, 1975; Lillegraven, 1977; Black and Sutton, 1984). Wilson (1940b) described Gripbomys alecer and G. sp., near alecer on the basis of five specimens rep- resenting a total of nine teeth from the Sespe For- mation of Simi Valley. Lillegraven (1977) assigned six specimens representing a total of six teeth from the Camp San Onofre Local Fauna, PSantiago For- mation, Camp Pendleton Marine Corps Base, Cal- ifornia, to G. alecer. Lillegraven (1977) also assigned four isolated teeth from the PSantiago Formation to a second species of Gripbomys, G. toltecus. Lind- say (1968) referred an LM3 from the Hartman Ranch Local Fauna of the Sespe Formation, north of the town of Ojai, California, to an undetermined spe- cies of Gripbomys. Golz and Lillegraven (1977) re- ported the occurrence of Gripbomys sp. from the Laguna Riviera Local Fauna, PSantiago Formation, Oceanside area, California. Sutton and Black (1975) questionably assigned an RM3 from the Pilgrim Creek Local Fauna, Colter Formation, Jackson Hole, Wyoming, to Gripbomys. Mary R. Dawson (pers. commun. in Sutton and Black, 1975) reported that Gripbomys may occur in the Badwater faunas from the PWagon Bed Formation of Wyoming. The above records represent the entire previously known occurrences of Gripbomys in the fossil record. Most investigators regard G. sp., near alecer to be syn- onymous with G. alecer, although none have pre- sented new evidence to support this assumption (Lillegraven, 1977; Black and Sutton, 1984; Kelly, 1990; Kelly et al., 1991). The impact mitigation program at the Simi Valley Landfill has resulted in the recovery of 30 additional teeth of G. alecer from 1 1 different stratigraphic levels in the Sespe Formation. The LM3 described by Lindsay (1968) from the Hartman Ranch Local Fauna can now be confidently assigned G. alecer because it is indistinguishable from the newly dis- covered M3s of this species from Simi Valley. A total of 46 teeth are now assigned to G. alecer. The four teeth identified herein as deciduous P4s are only tentatively assigned to G. alecer. These teeth could represent a different species of Gripbo- mys because they are smaller than the permanent P4s of G. alecer. However, these teeth were recov- ered from three different localities that each yielded other teeth that were definitely assignable to G. alecer. Furthermore, these teeth are almost identical in structure to the other upper cheek teeth assigned to G. alecer, including the presence of small pro- toloph spurs and strongly bilophodont occlusal pat- terns. The larger sample of G. alecer teeth allows for reevaluation of intraspecific variation and morpho- logical change through time. Wilson (1940b) divid- ed the specimens of Gripbomys into three taxa: G. alecer from locality LACM(CIT) 207, G. sp., near alecer from locality LACM(CIT) 202, and G. sp., near alecer from locality LACM(CIT) 150. These taxa were separated on the basis of the following characters: (1) slight differences in tooth size; (2) slight differences in the occlusal outlines of the P4s; (3) the presence or absence of vestigial mesoconids on the lower cheek teeth; (4) slight differences in the widths of the lower molar trigonids; and (5) the different stratigraphic occurrences of each taxon. Wilson noted that the teeth of Gripbomys from localities LACM(CIT) 202 and 207 were extremely similar and, considering the amount of individual variation present in other Eocene rodents, were probably conspecific. Wilson considered the sam- ple of Gripbomys from locality LACM(CIT) 150 most likely to represent a different species because the lower cheek teeth lack vestigial mesoconids, whereas those of G. alecer and G. sp., near alecer from localities LACM(CIT) 202 and 207 possess vestigial mesoconids. The development of proto- loph spurs and mesoconids on the teeth of Gri- j pbomys does appear to vary somewhat depending upon the stratigraphic level from which they were collected. In ascending stratigraphic order, these variations are as follows: (1) from the level of lo- cality LACM 5855 to that of locality LACM(CIT) 207, all upper cheek teeth have protoloph spurs and all lower cheek teeth have mesoconids present; (2) from the level of locality LACM(CIT) 202 to that of locality LACM 5866, the presence of pro- toloph spurs on the upper molars and mesoconids in the lower cheek teeth is highly variable; and (3) from the level of locality LACM (C1T) 150, all upper and lower cheek teeth lack protoloph spurs and mesoconids, respectively. Although there is an increase in the loss of protoloph spurs and meso- conids with decreasing geologic age, the cheek teeth that lack protoloph spurs and mesoconids from low in the section are otherwise indistinguishable from those of G. sp., near alecer from locality LACM(CIT) 150. Furthermore, the new teeth recovered during the impact mitigation program at the Simi Valley Landfill and those described by Lillegraven (1977) confirm that all of the characters that Wilson (1940b) used to separate species of Gripbomys are highly variable and not reliable for species identification. Therefore, all the specimens from the Sespe For- mation of Simi Valley and those referred to G. alecer by Lillegraven (1977) from the PSantiago For- mation can confidently be assigned to G. alecer. Gripbomys is generally regarded as belonging to the superfamily Geomyoidea and questionably as- signed to the Geomyidae because of the bilopho- dont structure of the molars (Wilson, 1940b, 1949; Lindsay, 1968; Sutton and Black, 1975; Wood, 1974; Black and Sutton, 1984). Wilson (1949) suggested that Gripbomys may have been derived from an ancestral sciuravine similar to the Bridgerian Tax- ymys Marsh, 1872. Lillegraven (1977) speculated that Gripbomys may be derived from a “ Nama - 18 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals Figure 16. PIsectaiophid, gen and sp. under., right upper cheek tooth, LACM 133987, occlusal view. Scale = 1 mm. tomys fantasma”- like eomyid based on his inter- pretation of the origin of the transverse lophids in Griphomys. “ Narnatomys ” is a distinct genus of early eomyid rodents that differs from Narnatomys Black, 1965 (Chiment, 1977; Lillegraven, 1977; Sto- rer, 1987; Kelly, 1992) and will be assigned to a new genus in a forthcoming report by J.J. Chiment and W.W. Korth (pers. commun., 1992). Storer (1987) considered “ Narnatomys ” to be derived from a sciuravid resembling the Wasatchian Knightomys Gazin, 1961 (Wood, 1965; Korth, 1984). Taxymys is a poorly known taxon and only the upper dentition is confidently assigned to this genus (Marsh, 1872; Wilson, 1938; Bown, 1982). The up- per molars of Griphomys resemble those of Tax- ymys in their bilophodont structure. The major changes in the upper cheek teeth that would be required to derive Griphomys from a Taxymys- like sciuravine are as follows: (1) loss of the P3; (2) development of a hypocone and metaloph on the P4 with subsequent widening of the central trans- verse valley and molarization; and (3) in the upper molars, loss of the metaconules on the metalophs, development of the protoloph spurs, and extreme reduction or loss of the mesostylids. The major changes in the teeth that would be required to derive Griphomys from a “ Narnatomys fantas- raTMike eomyid ancestor are as follows: (1) loss of the crests extending from the anterocone and anteroconid to the protoloph and metalophid, re- spectively; (2) loss of the posterior portion of the ectoloph and ectolophid; and (3) a slight increase in the height and development of the main lophs and lophids (protoloph, metaloph, metalophid, and hypolophid) resulting in greater bilophodonty. The development of the ectoloph, ectolophid, and the crests and cristids extending from the anterocone and anteroconid to the protoloph and metalophid, respectively, are derived characters for “ Namato - mys" (Chiment, 1977) and would not be expected in a basal eomyid ancestor. It appears that Gripho- mys is more easily derived from an ancestral eomyid- like form, as suggested by Lillegraven (1977), than from a Taxymys- like sciuravid. Storer (1987) pre- sented convincing evidence for the derivation of the early Uintan to late Duchesnean “ Narnatomys ” from an ancestral sciuravid resembling the Wa- satchian Knightomys. It is possible that Griphomys was derived from an ancestral “Narnatomys” -like eomyid during the intervening Bridgerian, making Griphomys a sister taxon of the eomyids. However, all of the above proposed evolutionary relation- ships are highly speculative. Details of the cranial and postcranial morphology of Griphomys, which would further clarify the phylogeny of the genus, are unfortunately unknown. Nevertheless, Gripho- mys is a distinctive taxon that exhibits affinities with both the Eomyidae and the Geomyoidea. Order Perissodactyla Owen, 1848 Family Isectolophidae Peterson, 1919 Pisectolophid, gen. and sp. undet. Figure 16 REFERRED SPECIMEN. Right upper cheek tooth, LACM 133987. LOCALITY. LACM 5616. FAUNA AND AGE. Brea Canyon Local Fauna, late Uintan. DESCRIPTION. The position of the upper cheek tooth from locality LACM 5616 within the dental arcade is uncertain; it could be a deciduous P4 or a molar. The parastyle is a distinct cusp that is connected to the ectoloph and the protoloph. The labially positioned paracone is a robust bilobed cusp. The metacone is the tallest and most prominent cusp along the ectoloph. A small metacone rib is present. The posterior crest of the metacone ex- tends posteriorly as a broadly curved crest and joins with the small metastyle. The anterior crest of the metacone extends anteriorly a very short distance where it then bifurcates into two crests, one leading anterolabially to join with the paracone and the other extending anteriorly to join with the para- style. The protocone and hypocone are prominent cusps separated by a central transverse valley that extends from the ectoloph to the lingual aspect of the tooth. The protoloph is a gently curved crest that extends from the protocone to join with the parastyle. The metaloph is slightly less prominent than the protoloph and extends labially from the hypocone to join with the base of the ectoloph at a point about midway between the metacone and the metastyle. The anterior, lingual, and posterior cingulae are well developed, whereas a labial cin- gulum is lacking. The measurements of LACM 133987 are 7.5 mm A-P and 7.6 mm TR. DISCUSSION. Kelly et al. (1991) questionably assigned the upper cheek tooth from locality LACM 5616 to an isectolophid perissodactyl, genus and species undetermined. This tooth exhibits charac- ters that are similar to those of the Isectolophidae, including a prominent ectoloph with a well-devel- oped parastyle and paracone, a reduced metastyle, and a cross lophed occlusal pattern formed by the Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 19 B » — i Figure 17. Protylopus robustus Golz, A. partial RM11, LACM 131448; B. LP4-M3, LACM 131447. All occlusal views. Scale = 1 mm. protoloph and metaloph. However, the tooth is very distinctive and does not compare well with those of any previously described genus or species of isectolophid (Radinsky, 1963; Schoch, 1989). This tooth could represent a previously unrecognized deciduous premolar or a developmental anomaly of a known isectolophid. Until a better sample of this taxon is available and following Kelly et al. (1991), this tooth is very questionably assigned to the Isectolophidae. Order Artiodactyla Owen, 1848 Family Oromerycidae Gazin, 1955 Genus Protylopus Wortman, 1898 Protylopus robustus Golz, 1976 Figure 17, Table 7 REFERRED SPECIMENS. Partial right maxilla with broken M'~3, LACM 131448; partial left den- tary with P4-M3, LACM 131447. LOCALITY. LACM 5616. FAUNA AND AGE. Brea Canyon Local Fauna, late Uintan. DISCUSSION. Golz (1976) described the rare oromerycid Protylopus robustus on the basis of two partial dentaries, the holotype (UCR 12833) from the Laguna Riviera Local Fauna of the ?San- tiago Formation, Carlsbad area, California, and a referred specimen (LACM 26363) from the Brea Canyon Local Fauna of the Sespe Formation of Simi Valley. The holotype is well worn, and the referred specimen from the Sespe Formation is a poorly preserved partial left dentary with the M2 and a broken M3. The impact mitigation program at the Simi Valley Landfill has resulted in the dis- covery of the first upper molars of P. robustus and a well-preserved, slightly worn partial left dentary with P4-M3 (Kelly et al., 1991). The upper molars of LACM 131448 are damaged with the M1 represented by the protocone and a partial paracone and metaconule, the M2 repre- sented by a partial protocone and the metaconule, and the M3 represented by a partial protocone and metaconule. These molars exhibit the following characters: (1) crescentic labial cusps; (2) bifurcated postprotocristae; (3) moderately developed anterior and posterior cingulae; and (4) labial shelves be- tween the protocones and metaconules. The lower molars of P. robustus have been well described by Golz (1976). However, the teeth of the newly discovered dentary are less worn than those of the holotype and exhibit the following differences: (1) the P4 metaconid is slightly more developed; (2) the metaconid in the lower molars 20 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals Table 7. Measurements (in mm) of teeth of Protylopus robustus. Tooth/ dimension LACM 131449 LACM 131448 M1 A-P — TR 8.6 (estimated) M2 A-P — TR — M3 A-P 1 1.5 (estimated) TR 10.6 (estimated) P4 A-P 8.1 TR 4.2 M, A-P 8.1 TR 6.4 M, A-P 9.5 TR 7.5 M, A-P 16.6 TR 8.3 M,-M, 34.3 is not connected to the co-joined cristids from the protoconid and hypoconid; and (3) the M3 hypo- conulid basin has a small cuspule at the anterior labial corner near the termination of the posterior cristid of the metaconid. Family Camelidae Gray, 1821 PCamelidae, gen. and sp. undet. Figure 18 REFERRED SPECIMEN. Partial RM2, LACM 130828. LOCALITY. LACM 5876. FAUNA AND AGE. Simi Valley Landfill Local Fauna, late Duchesnean. DESCRIPTION. The only specimen referable to this taxon is a well-worn, partial RM2 that is missing the mesostylar region, the paracone, the anterola- bial portion of the preprotocrista, and part of the anterior cingulum. The tooth appears to have been transversely elongated with a rectangular occlusal outline. The partial ectoloph, which extends from the metastyle to the anterior aspect of the meta- cone, is relatively straight. The metastyle is very small and the rib on the labial surface of the meta- cone is weakly developed. The protocone is deeply worn. The partial preprotocrista extends antero- labially and, presumably, was connected with the missing parastyle. The postprotocrista is rounded with wear and is separated from the anterior crest of the metaconule by a narrow valley between the protocone and metaconule. The anterior crest of the metaconule extends anterolabially wherein it joins the posterior crest of the paracone and the anterior crest of the metacone. The posterior crest of the metaconule extends posterolabially and joins with the metastyle and the posterior crest of the metacone. The metacone and metaconule are cres- centic and, even in their worn state, exhibit a mod- Figure 18. PCamelidae, gen. and sp. under., partial RM2, LACM 130828, occlusal view. Scale = 1 mm. erate degree of selenodonty. The anterior cingulum is moderately well developed and a small lingual cingulum is present between the protocone and metaconule, whereas labial and posterior cingulae are lacking. The measurements of LACM 130828 are as follows: 5.0 mm (estimated) A-P, 7.6 mm (estimated) ANT-TR, and 6.6 mm POST-TR. DISCUSSION. The partial M2 from Simi Valley differs from those of other middle to late Eocene selenodont artiodactyls, including Protoreodon Scott and Osborn, 1887, Diplobunops Peterson, 1919, Protylopus Wortman, 1898, Oromeryx Marsh, 1894, Eotylopus Matthew, 1910, Mala- quiferous Gazin, 1955, Camelodon Granger, 1910, Leptoreodon Wortman, 1898, Leptotragulus Scott and Osborn, 1887, Poabromylus Peterson, 1931, Hendryomeryx Black, 1978, Simimeryx Stock, 1934c, and Leptomeryx Leidy, 1853, by having a much less developed metastyle and an apparently straighter ectoloph. It further differs from those of species of the Oromerycidae by lacking a bifurcated postprotocrista and having a weaker developed rib on the metacone. The Simi Valley tooth is most similar to those of Poebrodon kayi Gazin, 1955, from the Uinta Formation of Utah. These similarities include a rel- atively straight ectoloph, a very small metastyle, a weakly developed rib on the metacone, a crescentic metaconule, and a tendency for the anterior crest of the metaconule to join with the posterior crest Figure 19. Simimeryx sp., aff. S. hudsoni Stock, A. RM2, LACM 130869; B. RM „ LACM 130868. All oc- clusal views. Scale = 1 mm. Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 21 of the paracorte with wear. The missing paracone, parastyle, and mesostyle in the Simi Valley tooth do not allow a comprehensive comparison with species of Poebrodon. However, it differs from those of Poebrodon by being more transversely expanded and having greater development of the anterior and labial cingulae. The Simi Valley tooth appears to belong to a taxon that is most closely related to the middle Eocene Poebrodon, but, because of the incomplete nature of the Simi Valley specimen, it is herein questionably referred to the Camelidae. Family Hypertragulidae Cope, 1879 Genus Simimeryx Stock, 1934c Simimeryx sp., aff. S. hudsoni Stock, 1934c Figure 19 Simimeryx n. sp. Kelly et al., 1991:8. REFERRED SPECIMENS. Partial RM1 and par- tial RM2, LACM 130867; RM2, LACM 130869; RM„ LACM 130868. LOCALITY. LACM 5876. FAUNA AND AGE. Simi Valley Landfill Local Fauna, late Duchesnean. DESCRIPTION. The upper molars of Simime- ryx sp., aff. S. hudsoni are represented by a partial M1 that is missing the paracone and protocone, a partial M2 that is missing the ectoloph, and a com- plete M2. The paracone is a rounded conical cusp. The parastyle is robust and positioned at the an- terolabial corner of the tooth. The metaconules are well developed and crescentic. The posterior crest of the metaconule extends anterolabially where it joins the posterior cingulum. The anterior crest of the metaconule extends posterolabially to the an- terolabial base of the paracone where it turns lin- gually towards the posterior crest of the protocone. The labial cingulum is reduced on the upper molars so that it is distinct anteriorly, posteriorly, and be- tween the paracone and metacone, but barely dis- cernable along the labial surfaces of the paracone and metacone. The anterior and lingual cingulae are moderately well developed, but the posterior cingulum varies from a moderately distinct crest to absent. The measurements of the only complete upper molar are 5.6 mm A-P, 6.6 mm ANT-TR, and 6.0 mm POST-TR. The M, is anteroposteriorly elongated and mod- erately well worn. The paraconid and entoconid are conical and somewhat bunodont cusps, whereas the protoconid and metaconid are crescentic. The parastylid is a distinct cristid that extends labially from the termination of the preprotocristid wherein it turns posteriorly to join the mediolabial surface of the paraconid. The anterior and lingual cingulids are moderately developed. A robust hypoconulid is present. The measurements of the M, are 8.8 mm A-P and 4.7 mm TR. DISCUSSION. The teeth from locality LACM 5876 are referred to Simimeryx because the upper molars lack mesostyles and have robust parastyles, postprotocristae that are posterolabially directed, and lingual cingulae. Also, the morphology of the M, hypoconulid is more like those of Simimeryx than those of other hypertragulids. Two species of Simimeryx are currently recog- nized: S. hudsoni of the Pearson Ranch Local Fau- na, Sespe Formation, California, and S. minutus Peterson, 1931, of the Lapoint Fauna, Duchenese River Formation, Utah. Fossils of Simimeryx hud- soni are well represented in the collections from the Sespe Formation, whereas S. minutus is known only from fragmentary material. The samples of Simimeryx teeth from locality LACM 5876 differ from those of S. hudsoni by having the following characters: (1) the teeth are slightly larger and slight- ly more selenodont; (2) the parastyle and parastylid are more labially positioned and the parastyle is slightly more reduced; (3) the anterior crest of the metaconule extends further posterolabially to the anterolabial base of the paracone where it turns lingually towards the posterior crest of the proto- cone; (4) the upper molar anterior, posterior, and lingual cingulae are less developed; (5) the upper molar labial cingulum is much less developed; and (6) the M, hypoconulid is proportionately larger. It is difficult to compare the Simimeryx teeth from locality LACM 5876 with those of S. minutus be- cause of the lack of comparative material, but the measurements of the Simi Valley teeth are much larger. The teeth from locality LACM 5876 are more derived than those of S. hudsoni, as indicated by their greater selenodonty, slightly more reduced cingulae and reduced parastyles in the upper molars, and relatively larger M3 hypoconulid, and appear to represent a new species. However, until an ad- equate sample of this taxon is available, these teeth are assigned to S. sp., aff. S. hudsoni. Most investigators regard Simimeryx as a mem- ber of the Hypertragulidae (Stock, 1934c; Gazin, 1955; Golz, 1976). However, Emry (1978) hesitated to include Simimeryx in this family because Sim- imeryx differs from the more typical hypertragulids, Hypertragulus Cope, 1873, and P arvitragulus Emry, 1978, by having (1) the protocones and metaconules of the upper molars in different positions relative to Simimeryx and (2) a very distinctive M, hypo- conulid morphology. The teeth of S. sp., aff. S. hudsoni exhibit some similarities with those of P ar- vitragulus and Hypertragulus, such as the reduced molar cingulae and cingulids. The M3 hypoconulid of S. sp., aff. S. hudsoni is also proportionally larger than those of 5. hudsoni and possesses a deeper central basin that is intermediate to those of S. hudsoni and those of other hypertragulids. The teeth of S. sp., aff. S. hudsoni differ from those of P arvitragulus and Hypertragulus by having the fol- lowing characters: ( 1 ) larger and less hypsodont mo- lars; (2) less transversely compressed upper molars; (3) much less developed paracone and metacone ribs; (4) a greatly reduced metastyle; (5) a more posteriorly directed postprotocrista; and (6) a rel- 22 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals atively smaller M, hypoconulid. The teeth of S. sp., aff. S. hudsoni appear to be morphologically in- termediate to those of S. hudsoni and those of the more typical hypertragulids, suggesting, contrary to Emry (1978), that Simimeryx should be included with the Hypertragulidae. Mammalia gen. and sp. under. Figure 20 REFERRED SPECIMEN. Partial right upper cheek tooth, LACM 130865. LOCALITY. LACM 5876. FAUNA AND AGE. Simi Valley Landfill Local Fauna, late Duchesnean. DESCRIPTION. The partial tooth (LACM 130865) is missing part of the metaconule and pos- terior cingulum, and the metacone. The occlusal outline appears to have been subtriangular in shape. A small metastyle is present. The protocone and metacone are tall conical cusps, with the protocone larger than the metacone. The protocone exhibits a postprotoconular fold that extends posterolabi- ally towards the posterior cingulum and a very ves- tigial postprotocrista that extends between the pro- tocone and the metaconule. The preprotocrista is a complete crest forming a protoloph that extends anterolabially from the protocone to join with the protoconule. The protoconule exhibits an oval wear pattern and appears to be smaller than the meta- conule. Only about half the metaconule is present, and it also appears to have had an oval wear pattern. The anterior cingulum is robust and extends lin- gually from the metastyle to about half way along the anterior face of the protocone. The labial cin- gulum is moderately developed across the labial surface of the metacone. A small distinct cusp (= Ppericone) is present along the anterior cingulum near its lingual termination. The partial posterior cingulum is robust and a small cusp (= Phypocone) is present near its lingual termination. The mea- surements of LACM 130865 can only be estimated as follows: 3.7 mm A-P and 4.6 mm TR. DISCUSSION. The partial upper cheek tooth (LACM 130865) from Simi Valley is perplexing be- cause its ordinal and familial assignment is uncer- tain. Kelly et al. (1991) assigned this tooth to an undetermined omomyine primate based on the presence of a postprotoconular fold. Further prep- aration and examination of this specimen makes this assignment questionable. Because of the in- complete nature of the specimen, it is herein as- signed to Mammalia, gen. and sp. undet. CONCLUSIONS The paleontologic resource impact mitigation pro- gram at the Simi Valley Landfill has yielded large samples of superposed middle to late Eocene small mammal assemblages. New taxa and additional specimens of poorly known taxa were recovered » ) Figure 20. Mammalia, gen. and sp. undet., partial right upper cheek tooth, LACM 130865, occlusal view. Scale = 1 mm. during the program. Prior to this report, many of these taxa were inadequately described because they were represented by fragmentary specimens or very small samples. The discovery of the following Uintan taxa from the middle member of the Sespe Formation is doc- umented: Centetodon sp., cf. C. aztecus ; erinaceo- morph, gen. and sp. undet.; Uintasorex sp., cf. U. montezumicus ; Microparamys woodi n. sp.; Mi- croparamys sp., cf. M. tricus ; Miacis sp. undet.; and Pisectolophid, gen. and sp. undet. Also, the discovery of the following Duchesnean taxa from the middle member of the Sespe Formation is doc- umented: Peradectes californicus ; Leptotomus sp. undet.; PCamelidae, gen. and sp. undet.; Simimeryx sp., aff. S. hudsoni ; and Mammalia, gen. and sp. undet. The impact mitigation program at the Simi Valley Landfill has resulted in larger samples of the teeth of the primate Dyseolemur pacificus, the rodent Griphomys alecer, and the artiodactyl Protylopus rohustus. The new specimens of D. pacificus de- scribed herein include the first P4 and the third M3 of this species recovered from the Sespe Formation. Among the known teeth of D. pacificus, the M3 appears to exhibit the greatest variation in mor- phology. In the teeth of G. alecer, the most variable characters are the development of the upper molar protoloph spurs and the lower molar mesoconids. The teeth of G. alecer exhibit similarities with those of the Eomyidae and the Geomyidae, and Gripho- mys may have originated from a “ Namatomys ”- like eomyid ancestor. The new specimens of P. rohustus described herein include the first upper molars referable to this species and a well-preserved partial left dentary with P4-M3. Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 23 Centetodon sp., cf. C. aztecus and Micropara- mys woodi are now known to have a geologic range from the early late Uintan Tapo Canyon Local Fau- na to the late Uintan Brea Canyon Local Fauna. Sespedectes singularis is now recorded in the Simi Valley Landfill Local Fauna, and this occurrence extends the geologic range of this species upward into the late Duchesnean. The taxa that constitute the late Duchesnean Simi Valley Landfill Local Fauna from locality LACM 5876 (Kelly et ah, 1991; Kelly, 1992) can now be revised to include the following: Peradectes cali- fornicus ; Sespedectes singularis ; Leptotomus sp. undet.; “ Namatomys ” sp.; Paradjidaumo reynold- si Kelly, 1992; Heliscomys sp.; Simiarcitomys whistleri Kelly, 1992; Simimys simplex (Wilson, 1935); Simimys landeri Kelly, 1992; Simimeryx sp., aff. S. hudsoni ; PCamelidae, gen. and sp. undet.; and Mammalia, gen. and sp. undet. Although the taxa that constitute the Simi Valley Landfill Local Fauna do not modify the extensive ecological in- terpretations of the paleoenvironment of the Sespe Formation presented by other investigators (Stock, 1932; Taylor, 1983, 1984; Krishtalka et al., 1987; Mason, 1988; Kelly, 1990), they do add knowledge to the characterization of the Duchesnean Land Mammal Age. The Duchesnean was previously characterized by the first appearances of Leptictis, Protadjidaumo, Ischyromys, P seudocylindrodon, Ardynomys, Jaywilsonomys, Presbymys, Eutypo- mys, Adjidaumo, Aulolithomys, Yoderimys, He- mipsalodon, Hyaenodon, Daphoenus, Duchesneo- dus, Menops, Hyracodon, Amynodontopsis, Mesohippus, Trigonias ?, Subhyracodon ?, Toxo- tberium, Brachyhyops, Archaeotherium, Agrioch- oerus, Poabromylus, Eotylopus, Aclistomycter, Heteromeryx, Hendryomeryx, Leptomeryx, and Hypertragulus and the last appearances of Prote- rixoides, Sespedectes, Simidectes, Chumashius, P areumys, Rapamys, Griphomys, Mytonomys, Is- chyrotomus, Hessolestes, Harpagolestes, Duches- neodus, Amynodontopsis, Triplopus ?, Epihippus, Protoreodon, Leptoreodon, Hyopsodus, Diplobu- nops, and Simimeryx (Wilson, 1986; Kelly, 1990; Kelly et al., 1991; Walsh, 1991; Lucas, 1992). Ad- ditional first appearances that can now be confi- dently recorded in the Duchesnean are Paradji- daumo, Heliscomys, and Simiarcitomys. Additional last appearances recorded in the Duchesnean are “ Namatomys ” and Simimys. Based on the taxa that constitute the Simi Valley Landfill Local Fauna, this fauna can be compared with other key Duchesnean North American land mammal faunas as follows: (1) it is younger than the Pearson Ranch Local Fauna from the Sespe Formation of California, the Skyline Local Fauna from the Agua Fria-Green Valley area of Texas, Lapoint Fauna from the Duchesne River Formation of Utah, and the faunal assemblages of the Bad- water Locality 20 and the Wood and Rodent Lo- calities from the Wagon Bed? Formation of Wyo- ming; (2) it is older than the Porvenir Local Fauna from the Vieja-Ojinaga area of Texas; and (3) it is probably a correlative of the Lac Pelletier Lower Fauna from the Cypress Hills Formation of Sas- katchewan. Prothero et al. (1992) recently reported that the Simi Valley Landfill Local Fauna occurs within the latter part of a reversed magnetozone, which they interpret as Chron C17R. Based on K-Ar radio- metric calibration, Berggren et al. (1985, 1992) re- gard Chron C17R to occur between about 41 and 41.5 Ma. However, revised Ar-Ar radiometric cal- ; ibration presented by Prothero and Swisher (1992) places Chron C17R between about 38.7 and 39 Ma. Based on the data presented by Prothero and Swish- er (1992) and Prothero et al. (1992), the age of the Simi Valley Landfill Local Fauna is about 38.7 to 38.8 Ma. Prothero and Swisher (1992) also provided revised Ar-Ar dates of 39.74 ± 0.07 Ma for the Lapoint tuff that occurs below the Lapoint Fauna between the Lapoint and Dry Gulch Creek Mem- bers of the Duchesne River Formation and 37.8 ± 0.06 Ma for the Buckshot Ignibrite of the Vieja- Ojinaga area that underlies the Porvenir Local Fau- na. In addition, Prothero and Swisher (1992) ques- tioned the validity of a K-Ar date of 42.3 ± 1.4 Ma (Black, 1969) for a biotite-bearing unit over- lying the faunal assemblage from Badwater Locality 20 because it contains older detrital biotite. The age of this biotite-bearing unit has been used by other investigators as a lower age limit for the Du- chesnean (e.g., Krishtalka et al., 1987; Kelly, 1990; Lucas, 1992). Furthermore, Prothero and Swisher (1992) and Prothero et al. (1992) have presented the following additional magnetostratigraphic cor- relations: (1) the Duchesnean Pearson Ranch Local Fauna occurs throughout most of Chron C18N, estimated between about 40.5 and 39 Ma; (2) the Duchesnean Lapoint Fauna occurs in the latter part of Chron C18N, estimated to be younger than about 39.7 Ma; (3) the latest Uintan Strathern Local Fau- na, which underlies the Pearson Ranch Local Fau- j na, occurs in the latter part of Chron C18R, esti- « mated to be about 40.5 Ma; and (4) the late Uintan Tapo Canyon and Brea Canyon Local Faunas occur from Chron C19N to the lower half of Chron C18R, estimated between about 42 and 40.5 Ma. The new Ar-Ar calibration of the magnetic time scale would place the Duchesnean between about 40.5 and 37 Ma (Prothero and Swisher, 1992; Prothero et al., 1992), whereas the K-Ar calibration would place the Duchesnean between about 42 and 39 Ma (Berggren et al., 1985, 1992). Lucas (1992) recently redefined the Duchesnean and regarded Duches- nean faunas as occurring from about 37 Ma to as old as 42 Ma, which he correlated with Chron C18 to part of Chron Cl 6. Irregardless of whether the new Ar-Ar or the K-Ar calibrations of the magnetic time scale are correct, it appears that the early Du- chesnean faunas occur within Chron C18N, the late Duchesnean faunas occur within Chrons C17R to C17N, and the Uintan-Duchesnean boundary oc- curs between Chrons C18N and C18R. All of the 24 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals above data support the biostratigraphic evidence that the Simi Valley Landfill Local Fauna is younger than the Lapoint Fauna, older than the Porvenir Local Fauna, and late Duchesnean in age. | ACKNOWLEDGMENTS We are indebted to the Simi Valley Landfill and Recycling Center (SVLRC), a division of Waste Management of i California, Inc., for providing financial support of the Paleontological Resource Impact Mitigation Program that has led to the recovery of the specimens described in this study. Special thanks is given to E. Bruce Lander and Mark A. Roeder of Paleo Environmental Associates, Inc. and the LACM for their direction of the extensive fieldwork at the Simi Valley Landfill, which resulted in the discovery of the new specimens described herein. We would also like to thank Mike Williams of SVLRC and Kim Uhlich and Scott Ellison of the Ventura County Resource Man- agement Agency Planning Divison for their support and recognition of the scientific significance of the paleon- tological resources of the Simi Valley Landfill. LITERATURE CITED Berggren, W.A., D.V. Kent, J.J. Flynn, and J.A. Van Cou- vering. 1985. Cenozoic geochronology. Bulletin of the Geological Society of America 96:1407-1418. Berggren, W.A., D.V. Kent, J.D. Obradovich, and C.C. Swisher 111. 1992. Toward a revised Paleogene ge- ochronology. In Eocene-Oligocene climatic and bi- otic evolution, ed. D.R. Prothero and W.A. Berg- gren, 29-45. Princeton, New Jersey: Princeton University Press, xiv + 568 pp. Black, C.C. 1965. Fossil mammals from Montana, part 2. Rodents from the early Oligocene Pipestone Springs Local Fauna. Annals of the Carnegie Mu- seum 38( 1 ): 1 —48 . . 1969. 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Preliminary report on a pa- leontologic investigation of the lower and middle members, Sespe Formation, Simi Valley Landfill, Ventura County, California. Paleo Bios 1 3(50): 1 — 13. Korth, W.W. 1984. Earliest Tertiary evolution and ra- diation of the rodents in North America. Bulletin of the Carnegie Museum of Natural History 24:1- 71. Krishtalka, L., and R.K. Stucky. 1983. Paleocene and Eocene marsupials of North America. Annals of the Carnegie Museum 52(10):229-263. Krishtalka, L., R.M. West, C.C. Black, M.R. Dawson, J.J. Flynn, W.D. Turnbull, R.K. Stucky, M.C. McKenna, Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 25 T.M. Bown, D.J. Golz, and J.A. Lillegraven. 1987. Eocene (Wasatchian through Duchesnean) biochro- nology of North America. In Cenozoic mammals of North America, geochronology and biostratig- raphy, ed. M.O. Woodburne, 77-117. Berkeley and Los Angeles: University of California Press, xv + 336 pp. Leidy, J. 1853. 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Bulletin of the American Museum of Natural History 158(4): 221-261. Lillegraven, J.A. , M.C. McKenna, and L. Krishtalka. 1981. Evolutionary relationships of middle Eocene and younger species of Centetodon (Mammalia, Insec- tivora, Geolabididae) with a description of the den- tition of Ankylodon (Adapisoricidae). University of Wyoming Publications 45:1-115. Lindsay, E.H. 1968. Rodents from the Hartman Ranch Local Fauna, California. Paleo Bios 6:1-22. Loomis, F.B. 1907. Wasatch and Wind River rodents. American Journal of Science, series 4 23:123-130. Lucas, S.G., 1992. Redefinition of the Duchesnean Land Mammal “Age”, late Eocene of western North America. In Eocene-Oligocene climatic and biotic evolution, ed. D.R. Prothero and W.A. Berggren, 88-105. New Jersey: Princeton University Press, xvi + 568 pp. Marsh, O.C. 1872. Preliminary description of new Ter- tiary mammals. American Journal of Science and Arts 4:122-128. . 1894. Description of Tertiary artiodactyls. American Journal of Science 48:259-274. Mason, M.A. 1988. Mammalian paleontology and stra- tigraphy of the early to middle Tertiary Sespe and Titus Canyon Formations, southern California. Un- published Ph.D. dissertation, Department of Pale- ontology, University of California, Berkeley, 257 pp. Matthew, W.D. 1910. On the skull of Apternodus and the skeleton of a new artiodactyl. Bulletin of the American Museum of Natural History 28(5):33-42. . 1918. A revision of the Lower Eocene Wasatch and Wind River Faunas. Part 5. Insectivora (contin- ued), Gilres, Edentates. Bulletin of the American Museum of Natural History 38:565-657. . 1920. A new genus of rodents from the middle Eocene. Journal of Mammalogy 1:168-169. Novacek, M.J. 1976. Insectivora and Proteutheria of the later Eocene (Uintan) of San Diego County, Cal- ifornia. Contributions in Science 283:1-51. — . 1985. The Sespedectinae, a new subfamily of Hedgehog-like insectivores. American Museum Novitates 2822:1-24. Peterson, O.A. 1919. Report upon the material discov- ered in the upper Eocene on the Uinta Basin by Earl Douglass in the years 1908-1909, and by O.A. Pe- terson in 1912. Annals of the Carnegie Museum 12(2-4):1-19. . 1931. New species from the Oligocene of the Uinta. Annals of the Carnegie Museum 12:40-168. Prothero, D.R., T.H. Dozier, and J. Howard, 1992. Magnetic stratigraphy and tectonic rotation of the middle Eocene-late Oligocene Sespe Formation, Ventura County, California. Geological Society of America, 1992 Meeting, Abstract No. 17505. Prothero, D.R., and C.C. Swisher III. 1992. Magneto- stratigraphy and geochronology of the terrestrial Eo- cene-Oligocene transition in North America. In Eo- cene-Oligocene climatic and biotic evolution, ed. D.R. Prothero and W.A. Berggren, 46-68. New Jer- sey: Princeton University Press, xvi + 568 pp. Radinsky, L. 1963. Origin and early evolution of North American Tapiroidea. Peabody Museum of Natural History, Yale University Bulletin 17:1-106. Schoch, R.M. 1989. A review of the tapiroids. In The evolution of perissodactyls, ed. D.R. Prothero and R.M. Schoch, 298-320. Oxford Monographs on Ge- ology and Geophysics 15:1-537. Scott, W.B., and H.F. Osborn. 1887. Preliminary report on the vertebrate fossils of the Uinta Formation, collected by Princeton Expedition of 1886. Pro- ceedings of the American Philosophical Society 24: 255-264. Stock, C. 1932. Eocene land mammals on the Pacific Coast. Proceedings of the National Academy of Sci- ences 18(7):518-523. . 1934a. A second Eocene primate from Cali- fornia. Proceedings of the National Academy of Sciences 20:150-154. . 1934b. New Creodonta from the Sespe upper Eocene, California. Proceedings of the National Academy of Sciences 20:423-427. . 1934c. A hypertragulid from the Sespe upper Eocene. Proceedings of the National Academy of Sciences 20:423-427. — . 1935a. New genus of rodent from the Sespe Eocene. Geological Society of America Bulletin 46: 61-68. . 1935b. Pleisomiacis, a new creodont from the Sespe upper Eocene, California. Proceedings of the National Academy of Sciences 21:119-122. — — — . 1935c. Insectivora from the Sespe uppermost Eocene, California. Proceedings of the National Academy of Sciences 21:214-219. — . 1936. Sespe Eocene didelphids. Proceedings of the National Academy of Sciences 22:122-124. Storer,J.E. 1987. Dental evolution and radiation of Eo- cene and early Oligocene Eomyidae (Mammalia, Ro- dentia) of North America, with new material from the Duchesnean of Saskatchewan. Dakoterra 3:108- 117. Sutton, J.F., and C.C. Black. 1975. Paleontology of the earliest Oligocene deposits in Jackson Hole, Wyo- ming. Part 1 . Rodents exclusive of the family Eomyi- dae. Annals of the Carnegie Museum 45(16):299- 315. Szalay, F.S. 1976. Systematics of the Omomyidae (Tar- 26 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals siiformes Primates) taxonomy, phylogeny and ad- aptations. Bulletin of the American Museum of Nat- ural History 156(3):157-450. Taylor, G.E. 1983. Braided-river and flood-related de- posits of the Sespe Formation, northern Simi Valley, California. In Cenozoic geology of the Simi Valley area, southern California, ed. R.L. Squires and M.V. Filewicz, 129-140. Pacific Section, society of Eco- nomic Paleontologists and Mineralogists, Fall Field Trip Volume and Guidebook, 266 pp. . 1984. Depositional environments of the Sespe Formation, northern Simi Valley, Ventura County, California. Unpublished M.S. thesis, California State University, Northridge, 168 pp. Troxell, E.F. 1923. Pauromys perditus, a small rodent. American Journal of Science 5:155-156. Walsh, S.F. 1987. Mammalian paleontology of the southern outcrops of the Mission Valley Formation, San Diego County, California. Unpublished senior thesis, California State University, San Diego, 171 pp. . 1991. Eocene mammal faunas of San Diego County. In Eocene geologic history San Diego re- gion, ed. P.L. Abbott and J.A. May, 161-177. Pacific Section, Society of Economic Paleontologists and Mineralogists Book 68:1-227. Wilson, J.A. 1986. Stratigraphic occurrence and corre- lation of early Tertiary vertebrate faunas, Trans-Pe- cos Texas: Agua Fria-Green Valley areas. Journal of Vertebrate Paleontology 6(4):350-373. Wilson, R.W. 1935. Cricetine-like rodents from the Sespe Eocene of California. Proceedings of the National Academy of Sciences 21:26-32. — . 1937. Two new Eocene rodents from the Green River basin, Wyoming. American Journal of Science, series 5 34:447-456. — — . 1938. Review of some rodent genera from the Bridger Eocene, part III. American Journal of Sci- ence, series 5 35:297-304. ■ . 1940a. California paramyid rodents. Carnegie Institute of Washington Publication 514(5):59-83. — . 1940b. Two new Eocene rodents from Cali- fornia. Carnegie Institute of Washington Publica- tion 514(6):85-95. . 1949. Additional Eocene rodent material from southern California. Carnegie Institute of Washing- ton Publication 584(l):l-25. Wood, A. E. 1949. Small mammals from the uppermost Eocene (Duchesnean) near Badwater, Wyoming. Journal of Paleontology 2 3:556-565. — . 1959. Rodentia. In The geology and paleon- tology of the Elk Mountain and Tabernacle Butte area, Wyoming, ed. P.O. McGrew, J. E. German, M. K. Hecht, G. G. Simpson, and A. E. Wood, 1 57— 169. Bulletin of the American Museum of Natural History 117:117-176. . 1962. The early Tertiary rodents of the family Paramyidae. Transactions of the American Philo- sophical Society 52:1-261. — . 1965. Small rodents from the early Lysite Mem- ber, Wind River Formation of Wyoming. Journal of Paleontology 39(1): 124- 134. — . 1974. Early Tertiary vertebrate faunas, Vieja group, Trans-Pecos Texas: Rodentia. Bulletin of the Texas Memorial Museum 21:1-111. Wortman, J.L. 1898. The extinct Camelidae of North America and some associated forms. Bulletin of the American Museum of Natural History 10(7):93- 142. Received 15 December 1992; accepted 20 May 1993. APPENDIX A CHARACTERS AND CHARACTER STATES USED IN CLADISTIC ANALYSES 1. Position of posterior margin of anterior root of the zygoma. Three states are recognized: 0, located in line between P4 and M1; 1, located in line with center of P4; 2, located in line with anterior margin of P4. 2. Position of anterior termination of masseteric fossa. Six states are recognized: 0, located in line with the trigonid of M,; 1, located in line between the pos- terior half to middle of M,; 2, located in line with the middle of M>; 3, located in line with center of anterior half of M2; 4, located in line between the anterior end of M, to posterior root of M,; 5, located in line with the middle of M,. 3. Number of mental foramina on mandibular ramus. Two states are recognized: 0, single foramen present; 1, two foramina present. 4. Cheek teeth crenulations. Three states are recog- nized: 0, not crenulated; 1, weakly to moderately crenulated; 2, extremely crenulated. 5. Size. Based on A-P length of M,. Four states are recognized; 0, medium 1.40-1.60 mm; 1, small 1.25- 1.40 mm; 2, very small <1.25 mm; 3, large <1.60 mm. 6. P3. Two states are recognized: 0, present; 1, absent. 7. P4. Two states are recognized: 0, non-molariform with single buccal cusp (paracone) and hypocone absent or rudimentary; 1, molariform with two buccal cusps (paracone and metacone) and distinct hypocone. 8. M1-2 occlusal patterns in hypoconal and protoconal regions. Two states are recognized: 0, occlusal pattern not U-shaped; 1, occlusal pattern U-shaped. 9. M1-2 metaloph and protoloph development and oc- clusal shape. Two states are recognized: 0, both lophs well developed and complete forming general V-shaped occlusal pattern; 1, one or both lophs in- complete and do not form a V-shaped occlusal pat- tern. 10. M1-2 protoconule development. Three states are rec- ognized: 0, protoconule well developed as single large conical cusp; 1, protoconule moderately reduced as elongated cusp; 2, absent or vestigial. 11. Mu2 metaconule development. Six states are recog- nized: 0, single unreduced metaconule usually present as large conical cusp; 1, single moderately reduced metaconule usually present as a relatively distinct cusp; 2, two metaconules (doubled) usually present as small reduced cuspules; 3, two metaconules (doubled) usu- ally present as well-developed cuspules; 4, single very reduced metaconule usually present; 5, metaconule absent or vestigial. 12. M12 metaconule position. Two states are recognized: 0, metaconule positioned labially and in close asso- ciation with metacone; 1, metaconule positioned lin- gually and not in close association with metacone. 13. M1'2 postprotocrista development. Two states are recognized: 0, absent or poorly developed; 1, mod- erately well to well developed. 14. M1-2 hypocone development. Two states are recog- Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 27 nized: 0, moderately developed cusp; 1, well-devel- oped cusp. 15. M2 mesolophid. Defined as spur or loph originating near apex of protocone that extends lingually into the central basin of the tooth. Two states are rec- ognized: 0, absent or weakly developed; 1, present as a distinct cristid or loph. 16. P4 size relative to lower molars. Defined as ratio of mean M2 A-P length to mean P4 A-P length. Four states are recognized: 0, moderately reduced (ratio = 0. 75-0.90); 1, slightly reduced (ratio = 0.90-0.98); 2, very reduced (ratio = 0.65-0.75); 3, extremely re- duced (ratio = <0.65). 17. P4 mesoconid. Three states are recognized: 0, absent; 1, present, weakly developed; 2, present, strongly de- veloped. 18. P4 ectolophids. Two states are recognized: 0, absent; 1, usually developed as a complete cristid connecting the protoconid, mesoconid, and hypoconid or two almost complete cristids extending from mesoconid towards protoconid and hypoconid, respectively. 19. P4 hypoconid. Two states are recognized. 0, small and poorly developed; 1, well developed. 20. P4 posterolophid. Two states are recognized: 0, absent or very weakly developed; 1, present, well developed. 21. P4 protoconid. Two states are recognized: 0, usually present as a distinct cusp; 1 , usually absent or vestigial. 22. Complexity of lower molar occlusal patterns. Two states are recognized: 0, simple to moderately simple; 1, very complex with abundant small cristids and lophs present throughout the talonid. 23. M,^2 trigonid size. Two states are recognized: 0, equal or subequal in size to talonids; 1, markedly narrower than talonids. 24. M,_, mesoconid shape. Two states are recognized. 0, not anteroposteriorly compressed; 1, anteroposteri- orly compressed. 25. Mu, anterior cingulid development. Two states are recognized: 0, weakly developed as short low cristid; 1, well developed as long, distinct cristid. 26. M,_, anterior cingulid connection with protoconid. Four states are recognized: 0, anterior cingulid sep- arated from protoconid by small, shallow valley and labial terminus of anterior cingulid connected to pro- toconid; 1, anterior cingulid separated from proto- conid by distinct groove or valley and labial terminus of anterior cingulid not connected to protoconid; 2, anterior cingulid separated from protoconid by nar- row groove in early wear that disappears with mod- erate wear to form connection between labial ter- minus of anterior cingulid and protoconid; 3, anterior cingulid separated from protoconid by distinct groove or valley and connected to protoconid by small but relatively persistent thin cristid originating from near labial terminus of anterior cingulid. 27. M,_, anterior cingulid connection with metaconid. Three states are recognized: 0, anterior cingulid is weakly developed as low, short cristid connected to base of metaconid; 1, anterior cingulid moderately to well-developed cristid connected to the metaco- nid; 2, anterior cingulid moderately developed as a distinct cristid along anterior margin of tooth and separated from metaconid by distinct gap or groove. 28. Mu2 development of cusp on labial terminus of an- terior cingulid. Two states are recognized: 0, not cus- pate; 1, cuspate. 29. M,_, trigonid. Two states are recognized: 0, trigonid usually open posterolingually; 1, trigonid usually closed posterolingually. 30. M,_, protoconid. Two states are recognized: 0, pro- toconid not isolated, but connected or nearly con- nected to ectolophid and posterior arm of protoconid short or if developed usually directed towards middle of metaconid or further forward, and posterior arm is not parallel with anterior cingulid; 1, protoconid relatively isolated cusp with posterior arm of the pro- toconid extending to or nearly to the posterior labial base of metaconid, and posterior arm of protoconid nearly parallel with anterior cingulid. 31. M,., hvpolophid development. Two states are rec- ognized: 0, incomplete, absent or rudimentary cristid is present that originates from the entoconid and is labially directed a short distance into the central basin of tooth; 1, complete cristid present, connecting en- toconid with hypoconid resulting in a distinct en- closed valley between the hypolophid and postero- lophid. 32. Mu2 posterolophid development. Two states are rec- ognized: 0, posterolophid weakly developed; 1, pos- terolophid well developed. 33. M ,_2 “mesolophids.” Defined as small spurs or cristids generally directed lingually from the mesoconid into the central basin of the tooth. Two states are rec- ognized: 0, absent or rudimentary; 1, present as one to three small spurs. 34. M,_, hypoconulid development: Four states are rec- ognized: 0, present as well-developed isolated cusp; 1, present as moderately well-developed, often elon- gated, cuspule formed along posterolophid; 2, incip- ient hypoconulid present as a widening along pos- terolophid; 3, absent. 35. M,_, entoconid isolation. Four states are recognized: 0, M,_2 entoconid isolated and well separated from posterolophid by well-developed wide gap; 1, M,_2 entoconid isolated and separated from posterolophid by narrow distinct groove; 2, M,_2 entoconid con- nected to posterolophids; 3, M, entoconid isolated from posterolophid by narrow groove, whereas M, entoconid connected to posterolophid. 36. ectolophids. Defined as cristids extending from mesoconid to or near protoconid and hypoconid forming anteroposteriorly directed loph along labial aspect of tooth. Two states are recognized: 0, usually present with one or both cristids complete or nearly complete; 1, usually absent. 28 ■ Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals APPENDIX B I; Character state matrix for following group of taxa: Cocomys (outgroup), Reithroparamys, Apatosciuravus, Acrito- paramys, Microparamys, Lophiparamys, “ Microparamys ” reginensis, “ Microparamys ” scopaiodon, and Pauromys. Characters and character states are defined in Appendix A. Characters 1 2 3 4 6 7 10 11 14 16 18 19 20 21 22 23 24 26 27 30 32 34 35 36 Cocomys 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 1 Reithroparamys 1 1 0 0 0 1 0 0 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 Apatosciuravus 2 1 1 0 0 1 2 4 1 2 1 1 1 1 0 0 0 0 1 0 0 1 0 0 Acritoparamys 1 3 1 0 0 1 0 1 1 1 1 1 1 1 0 1 0 0 1 0 1 1 1 0 Microparamys 2 5 0 1 1 1 1 1 1 0 1 1 1 0 0 0 0 1 1 0 1 2 1 0 Lophiparamys ? 2 0 2 0 1 1 1 1 1 1 1 1 1 1 0 0 1 1 0 1 1 1 0 “M.” reginensis ? 4 0 0 ? ? ? ? p ? ? 1 1 ? 0 0 1 2 1 1 1 1 1 1 “M.” scopaiodon p 4 0 0 ? p p p p 3 0 1 1 0 0 0 1 2 1 1 1 1 1 1 Pauromys 2 5 0 0 1 1 2 5 1 3 0 1 1 0 0 0 1 1 2 1 0 2 0 1 APPENDIX C Character state matrix for Cocomys ( outgroup) and reithroparamyine genera. Characters and character states are defined in Appendix A. Characters 1 2 3 4 6 7 10 11 12 14 16 18 19 20 21 22 23 24 26 27 30 32 34 35 36 Cocomys 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 1 Reithroparamys 1 1 0 0 0 1 0 0 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 Apatosciuravus 2 1 1 0 0 1 2 4 1 1 2 1 1 1 1 0 0 0 0 1 0 0 1 0 0 Acritoparamys 1 3 1 0 0 1 0 1 1 1 1 1 1 1 1 0 1 0 0 1 0 1 1 1 0 Microparamys 2 5 0 1 1 1 1 1 1 1 0 1 1 1 0 0 0 0 1 1 0 1 2 1 0 Lophiparamys ? 2 0 2 0 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 0 1 1 1 0 APPENDIX D Character state matrix for Cocomys lingchaensis (outgroup) and species of Microparamys. Characters and character states are defined in i Appendix A. Characters 1 2 4 5 7 8 9 11 13 15 16 17 18 19 20 21 25 26 28 29 31 32 33 34 35 Cocomys lingchaensis 0 0 0 3 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 M. minutus 2 5 1 0 1 0 1 1 1 0 0 2 1 1 1 0 1 1 0 0 0 1 0 2 1 M. sp., cf. M. minutus ? p 1 1 1 0 1 1 1 0 0 2 1 1 1 0 1 1 0 0 0 1 0 2 1 M. duhius ? p 1 1 1 0 1 1 0 0 0 1 1 1 1 0 1 3 1 1 1 1 0 1 1 M. sp., cf. M. tricus ? p 1 3 p 0 1 3 0 0 p ? ? p p p 1 3 1 0 0 1 1 2 1 M. tricus 2 5 1 3 1 0 1 3 0 0 1 1 1 1 1 1 1 3 1 0 0 1 0 2 1 M. woodi ? p 1 0 1 1 0 2 1 1 0 2 1 1 1 0 1 3 0 1 0 1 1 3 2 M. perfossus ? 5 1 0 1 1 1 4 0 1 1 1 1 1 1 0 1 3 p 0 0 1 0 3 2 Contributions in Science, Number 439 Kelly and Whistler: Sespe Mammals ■ 29 Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Number 440 24 February 1994 m m Contributions in Science Late Cenozoic Equids from the Anza-Borrego Desert of California Theodore Downs and George J. Miller Natural History Museum of Los Angeles County Serial Publications of THE Natural History Museum of Los Angeles County Scientific Publications Committee Craig C. Black, Museum Director Daniel M. Cohen Kirk Fitzhugh John M. Harris, Committee Chairman Edward C. Wilson Richard C. Hink, Managing Editor Robin A. Simpson, Head of Publications The scientific publications of the Natural History Mu- seum of Los Angeles County have been issued at irregular intervals in three major series; the issues in each series are numbered individually, and numbers run consecutively, regardless of the subject matter. # Contributions in Science, a miscellaneous series of tech- nical papers describing original research in the life and earth sciences. # Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth sci- ences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now published by the Museum in Contributions in Science. # Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through the Museum Book Shop. A catalog is available on request. The Museum also publishes Technical Reports, a mis- cellaneous series containing information relative to schol- arly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum’s Scientific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Tech- nical Reports may be obtained from the relevant Section of the Museum. Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Printed at Allen Press, Inc., Lawrence, Kansas ISSN 0459-8113 Late Cenozoic Equids from the Anza-Borrego Desert of California Theodore Downs1 and George J. Miller2 CONTENTS ABSTRACT INTRODUCTION METHODS AND MATERIALS SYSTEMATICS cf. Dinohippus sp Equus Characterization of some Equus subgenera . . . Descriptive Terminology Equus ( Dolichohippus ) enormis, new species . . Equus ( Dolichohippus ), cf. E. (D.) simplicidens Equus (Equus) species A Equus (Equus) species B Equus cf. Equus (Equus) species SUMMARY Morphologic Trends Paleoenvironment Paleobiogeography CONCLUSIONS ACKNOWLEDGMENTS LITERATURE CITED APPENDIX A — Abbreviations APPENDIX B — Additional measurements 2 2 2 4 4 9 9 15 19 57 66 69 73 78 78 81 82 84 84 85 89 90 1. Emeritus, Vertebrate Paleontology Section, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. 2. Paleontology Department, Imperial Valley College Museum, 442 Main Street, El Centro, California 92243. De- ceased, December 1989. Contributions in Science, Number 440, pp. 1-90 Natural History Museum of Los Angeles County, 1994 ABSTRACT. Well-preserved fossil Equidae recovered from the earliest Pliocene to middle Pleistocene sequence in the Anza-Borrego Desert of southern California (Downs and White, 1968; Miller, 1985) represent six different species. A small species, cf. Dinohippus sp., occurs in the earliest Pliocene (late Hemphillian) and later Pliocene (early to late Blancan) strata of the Imperial and Palm Spring Formations. A new species of a large dolichohippine, Equus { Dolichohippus ) enormis, was recovered from the Vallecito Creek local fauna, Palm Spring Formation (transitional Blancan to Irvingtonian), the Borrego Badlands area, Palm Spring or Ocotillo Formations (late Irvingtonian), and the lower Coyote Canyon badlands, Ocotillo Formation (late Irvingtonian). Well-preserved crania and dentitions from late Blancan strata (late Arroyo Seco local fauna and early Vallecito Creek local fauna) appear morphologically intermediate between the largest specimens of E. ( Dolichohippus ) simplicidens (Cope, 1892) and E. (D.) enormis and are identified as E. { Dolichohippus ) cf. E. (D.) simplicidens. Three crania and four mandibles are referred to the caballine subgenus Equus {Equus). Equus {Equus) species A is from the Vallecito Creek local fauna, Palm Spring Formation (late Blancan to Irvingtonian). Equus {Equus) species B and E. {Equus) sp. indet. are from the Borrego Badlands, Palm Spring and Ocotillo Formations (late Irvingtonian). Morphological trends link the small cf. Dinohippus sp. to the larger Equus {Dolichohippus) cf. E. (D.) simplicidens and thence to the large E. (D.) enormis n. sp. The dolichohippine and caballine species constitute North American representatives of populations that migrated into Eurasia and Africa during the late Pliocene and early Pleistocene, respectively. INTRODUCTION The equids described in this report were collected from southeastern California, within the Colorado Desert, as first named by Blake in 1858. Specifically, they were retrieved from what is now the Anza- Borrego Desert State Park. This park is at the west- ern margin of the Imperial Valley, San Diego Coun- ty, California. Geologically, the emergent north ex- tension of the Gulf of California, today’s Imperial Valley, is a structural trough filled with a sequence of marine and terrestrial sediments derived from the bordering uplands, the Colorado River delta, and the previous northern extensions of marine deposition in the Gulf of California. Studies of the stratigraphy and geology of the area by Woodard (1974), of the paleomagnetic stratigraphy by Op- dyke et al. (1977), of the fission track data and tectonism by Johnson et al. (1983), and of the bio- stratigraphy by Downs and White (1968) provide background information for this report. The “zones” referred to in this study are lithostrati- graphic entities, not formal or informal biostrati- graphic zones. These and the local faunas referred to in the locality descriptions are described in more detail by Opdyke et al. (1977) and Downs and White (1968). Approximately 20,000 feet of stratified and lat- erally gradational marine and continental sediments were deposited in the type section area. Woodring (1931) named the marine deposits the Imperial For- mation and the overlying predominantly terrestrial sediments the Palm Spring Formation. There are approximately 12,200 feet of the Palm Spring For- mation exposed that preserved vertebrate fossils, but most fossil vertebrates occur in the upper 4,000 feet of the section. Fossils were collected from an approximately 150-square-mile area, extending from the Fish Creek and Vallecito Creek mountains southeast to the Coyote Mountains. Dolichohippine and caballine fossil Equus were also collected in the Borrego Badlands and lower Coyote Canyon of the northwestern Imperial Val- ley. The specimens occurred in the Ocotillo For- mation (Bartholemew, 1970) or Palm Spring For- mation (Dibblee, 1954). Frick (1937:202) published the first record of a vertebrate fossil from the Anza-Borrego area — some cervid elements from “Carrizo Creek, Southern California” collected by Guy E. Hazen in 1936. This area is now recorded on the Arroyo Tapiado Quadrangle (1959) as the Carrizo Valley and Val- lecito Creek. Frick named the fossils Odocoileus cascensis, new species, whose limb proportions “approximate Odocoileus hemionus AM(M) 122667 from Alberta, Canada.” Harley Garbani of San Jacinto, California, brought some vertebrate fossils to the Natural History Mu- seum of Los Angeles County (LACM) in 1954 and sought information from T. Downs. Subsequently, Garbani provided guidance for LACM personnel in the Anza-Borrego Desert which led to further collecting for many more years, with annual col- | lecting permission issued by the California Depart- ment of Parks and Recreation. Specimens now in the collections are recorded as loans from the state to the Natural History Museum of Los Angeles County. The fossil equid materials from Anza-Borrego that are discussed in this report were formerly housed in the Imperial Valley College Museum, El Centro, California, and Natural History Museum of Los Angeles County, Los Angeles, California. The Imperial College Museum Vertebrate Pale- ontology Collection is currently housed in the Stout Laboratory, next to the Anza-Borrego State Park headquarters, Post Office Box 299, Borrego Springs, California 92004. The Anza-Borrego Desert se- quence has yielded additional fossil equids with probable hemionine, asinine, and Hippidion-\ike affinities. These will be described in a future report. METHODS AND MATERIALS In the course of our study of Equus, 119 types of mea- surements or character states were recorded from which 2 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids DENTAL PARTS Metaconid Protoconid Paralophid Metastylid Hypoconid Entoconid Hypostylid DENTAL FOLDS Preflexid Linguaflexid Cement TOOTH COMPONENTS \ X Enamel Postflexid L^r0' Ectoflexid Meta-isthmus b Figure 1. Dental nomenclature: a. Equus ( Dolichohippus ) grevyi (Oustalet, 1882); Holocene, AMNH (M) 54247, partial P3, P4-M3; b. Equus ( Dolichohippus ) simplicidens Cope (1892); early Pleistocene, Nebraska, F:AM 87440, partial P3, P4-M3; a and b reproduced from Skinner (1972:fig. 57). computer printouts of exploratory, two dimensional scat- tergrams and Simpson ratio diagrams were prepared. From these analyses we have selected the most significant fea- tures for the discussions presented in the text, tables, figures, and appendices. Methods of measurement include those used in Gazin (1936), Hibbard (1955), Gromova (1949), and Eisenmann (1979a, b, 1981a, 1983) as well as our own additions. They are recorded in millimeters, using dial or vernier calipers, and were made by either of the authors. Detailed measurements and comparisons were made by the authors on 17 crania and mandibles of E. (D.) simplicidens (Cope, 1892), two crania and mandibles of E. idaboensis Merriam, 1918, eight specimens of E. (D.) grevyi Oustalet, 1882, nine of E. (E.) przewalskii Polia- kov, 1881, and two skeletons of E. {Equus) caballus (Lin- naeus, 1758). Our nomenclature for dental features primarily follows that of Stirton (1940, 1941) and Skinner (1972); see Figures 1 and 2. The depth (or length of penetration of) the ectoflexid in Equus lower cheek teeth was measured from the labial border of the protoconid and hypoconid to the lingual tip of the penetrating ectoflexid (also see Fig. 2E). The paralophid (= parastylid) was measured from the external labial border of the protoconid to the internal- lingual extent of the paralophid tip. We follow Sisson and Grossman (1964) for the skull and postcranial anatomical terminology. Important measurement data were provided to us by Melissa Winans in 1980 following her examination of several collections from United States museums contain- ing samples of E. (D.) simplicidens and E. (Equus) prze- walskii. Abbreviations and symbols used in the text, tables, and figures are presented in Appendix A. Some measurements not presented in the tables are presented in Appendix B. Museum specimens cited and their acronyms are: American Museum of Natural History (AMNH), New York, New York; Field Museum of Natural History (FMNH), Chicago, Illinois; Florida State Museum (FSM), Gainesville, Florida; Idaho State University Museum (ISUM), Pocatello, Idaho; Imperial Valley College Mu- seum (IVCM), El Centro, California; Museum National d’Histoire Naturelle (MNHN), Paris, France; National Museum of Natural History (USNM), Washington, D.C.; National Museums of Kenya (NMK), Nairobi, Kenya; Natural History Museum of Los Angeles County (LACM), California; University of California Museum of Paleon- tology (UCMP), Berkeley, California; and the University of Michigan Museum of Paleontology (UM), Ann Arbor, Michigan. The recent publication Studying Fossil Horses, volume I: Methodology, by Eisenmann et al. (1988), was published after we compiled data for this paper. It is impractical for us to modify our data, and we believe that our methods and nomenclature differ only slightly and are adequately Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 3 illustrated in Figures 1 and 2, or explained in our mea- surement tables and footnotes. Detailed locality information for all specimens de- scribed is on file in the Section of Vertebrate Paleontol- ogy, LACM and IVCM. SYSTEMATICS Class Mammalia Linnaeus, 1758 Order Perissodactyla Owen, 1848 Superfamily Equoidea Hay, 1902 Family Equidae Gray, 1821 Dinohippus Quinn, 1955 cf. Dinohippus sp. Figures 3, 4 MATERIAL. LACM 4356, right M3 from lo- cality LACM 1656; IVCM 2257-1, right M3, IVCM 2257-2, right M2, and IVCM 2257-3, right M1, from locality IVCM 177; IVCM 1873, partial left man- dible with I„ partial L and P2 to M3; a partial right mandible with I,, L, I3, broken canine and partial post-canine diastema; moderate wear on dentition from locality IVCM 537. LOCALITIES. LACM 1656 and IVCM 177 from Fish Creek area, IVCM 537 from Arroyo Seco Wash, Anza-Borrego Desert State Park, San Diego County, California. STRATIGRAPHY AND AGE. LACM 1656 from “zone” 4, Imperial Formation, probable pre-Layer Cake local fauna, late Hemphillian in age (early Pliocene); IVCM 177 from “zones” 7-8, Imperial- Palm Spring transitional Formations, pre- or earliest Layer Cake local fauna, late Hemphillian to early Blancan in age (early Pliocene); IVCM 537 from “zone” 43, middle to upper Palm Springs Forma- tion, late Arroyo Seco local fauna, Blancan age (middle to late Pliocene). The chronostratigraphic range for the Anza-Bor- rego cf. Dinohippus sp. is approximately 3.9 Ma or 4.0 Ma to 2.7 Ma, a total of about 1.3 million years. DESCRIPTION AND COMPARISONS. On the basis of generic characters of equids outlined by Stirton (1940), Quinn (1955), Bennett (1980), and MacFadden (1984), it is now apparent that LACM 4356 and the more recently collected IVCM 2257- 1, 2257-2, and 2257-3 can be distinguished from Equus by their simple enamel pattern, absence of a protoconal heel or definitive protoconal groove, small size, and low height of crown. The molars resemble those of Pliohippus, from which they may be distinguished by their relatively straight crowns, absence of a lingual protoconal groove, and lack of protoconal heel. Straight crowns, lack of heels, and simple enamel pattern are characters shared with Dinohippus, especially with the type of Dino- hippus leidyanus (Osborn, 1918). However, the variation of these features in Dinohippus mexican- us in the LACM collection suggests caution should be exercised in identifying individual upper cheek teeth. For the present, we shall refer to the upper dentition as cf. Dinohippus sp. IVCM 2257-1 is a right M1. The paracone and metacone walls are deeply concave without ribs. The protocone is elongate and bears a suggestion of a lingual groove; there is no protoconal heel. A wide isthmus connects with the protoselene; the post-protoconal valley is deep, wide, and without a plicaballin. There is a very small hypocone and a shallow post-hypoconal groove. The enamel pat- tern is simple with only two small plications in the post-fossette. The parastyle is well developed (5.5 mm long). The mesostyle is 2.0 mm long. There is a minimum of tooth curvature. Measurements of IVCM 2257-1 are: AP 33 mm, TR 25.4 mm, pro- tocone length 12.2 mm, and crown height 41.6 mm. IVCM 2257-3 is also a right M1. The paracone and metacone walls are relatively straight; both the parastyle (4.7 mm long) and mesostyle (3.9 mm long) are deep. The enamel pattern is simple with one pli-protoconule. The protocone is elongate with very slight suggestion of a heel and a lingual groove; the hypocone is small with a minute hypoconal groove. Measurements of IVCM 2257-3 are: AP 26 mm (estimated), TR 30.2 mm, protocone length 10.9 mm, and crown height 39.1 mm. IVCM 2257-2 is a right M2. The parastyle is nar- row (AP 4.7 mm) and wider than the mesostyle (2.8 mm long). The styles are deep, and the walls of the paracone and metacone are concave. There are two small pre-fossette plications, otherwise the tooth has a simple enamel pattern. The protocone is elon- gate, without a lingual groove or protoconal heel. There is a narrow and deep post-protoconal valley, a single wide plicaballin; very slight pre-protoconal groove; small hypocone with a very short hypo- conal groove; and very slight tooth curvature. Mea- surements of IVCM 2257-2 are: AP 28.2 mm, TR 29.0 mm, protocone length 10.9 mm, and crown height 46.1 mm. LACM 4356, a right M3, is somewhat smaller than that of most Equus species. The protocone is elongate without a groove or heel; it connects with the protoselene by a narrow isthmus. The parastyle and mesostyles appear to be narrow; the paracone and metacone walls are concave and relatively deep. There are two small pli-protoconules. There is no apparent hypocone. The enamel pattern is simple, and the tooth has very slight curvature. Measure- ments of LACM 4356 are: AP 29 mm, TR 23 mm, protocone length 9.6 mm, and crown height 30 mm. ; IVCM 1873 comprises partial left and right man- dibles bearing teeth (Fig. 4). The right and left first incisors in IVCM 1873 are complete and well worn, with small circular cups and relatively thin external borders; L is much shorter mesiodistally than in other species compared. The right I2 has an elongate cup, closely adjacent to the lingual border, and the tooth is smaller than in other species compared (see Table 3). Right I3 is slightly worn and has a broad 4 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids Figure 2. A. Equus ( Dolichohippus ) simplicidens s.l. from Coso Mountain local fauna, LACM (CIT) 902, right I,_3 (restored from left side), moderate state of wear, occlusal view. a. infundibulum or cup, b. enamel cingulum (can become infundibulum with wear) also referred to as recessed open “V.” B. Equus ( Dolichohippus ) simplicidens Cope, 1892, LACM (CIT) 2021, from Coso Mountain local fauna California, right I, — 13, well worn, occlusal view. a. enamel, b. nutrient canal, c. basal remnant of recessed open “V.” C. Equus {Equus) caballus (Linnaeus, 1758), Holocene, LACM 51575, right I,-I3, moderate wear, occlusal view. a. enamel, b. enamel bordered infundibulum. D. Equus ( Asinus ), Holocene, LACM 31132, left I,-I3, well worn occlusal view. a. infundibulum, b. recessed open “V.” E. Diagram of lower molars, degrees of penetration of the ectoflexid into the isthmus (estimated) in right M, or M2. a. no penetration, b. slight penetration, c. penetration; d. deep penetration, touching the linguaflexid. F. Diagram linguaflexid shape categories of lower dentition: (1) narrow-deep, (2) broad “V,” (3) deep “U,” (4) broad “U,” (5) irregular. G. Diagram metastylid shape categories of lower dentition: (1) rounded, (2) triangular, indented, (3) triangular, (4) oval. H. Diagram protocone shape categories of upper dentition: (1) short, (2) moderately elongate, (2a) elongate, (3) moderately elongate with narrow lingual groove, (4) elongate with broad lingual groove, (5) elongate with irregular lingual goove. recessed “V” as in Dinobippus and E (D.) simplici- dens. R. H. Tedford (personal communication, 1985) has verified for us the lack of a cup in the I3 of the holotype of D. leidyanus, and he noted this was also true of samples of Dinobippus from the Edson Quarry, Kansas. MacFadden (1984:280) ob- served that in Dinobippus mexicanus from the Ye- pomera local fauna of Mexico the incisors had “cement filled infundibulae.” However, in our sur- vey of the I3’s in the Yepomera collection of D. Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids H 5 Table 1. Characters distinguishing subgenera of Equus from North America (based on characters commonly available in fossil material, initially on information provided by M. F. Skinner). Character states £.(£.)* £.(jD.)* E.{He.)* E.{As.)* E.{Am.) 1. Cranial, rostral proportions: elongate, index1 0.50- 0.55 or less (E). Short or broad, index 0.50-0.55 or more (S). S E S S S2 2. Mandibular rostral proportions: elongate, index3 0.38-0.54 or less (E). Short or broad, index 0.50- 0.68 or more (S). S E S S s 3. Orbit position: posterior to tooth row, index4 0.41- 0.58 or more (P), 0.24-0.40 or less (A). A P A A A 4. Isthmus: P/2 to P/4 strong isthmus; protoconid united directly to metaconid-metastylid column, hypoconid joined to posterior part of protoconid junction.5 + + + + + 5. Isthmus: M/1 to M/3 strong isthmus; protoconid united to metaconid, metastylid more posteriorly placed, lingual to metaconid-protoconid junction, no ectoflexid penetration, index6 0.36-0.49 or less (I). M/1, M/2, and usually M/3 ectoflexid pene- trates and splits isthmus producing antro-, meta-, and post-isthmus, index 0.50-0.78 or more (P). P7 P I I P7 6. Isthmus: M/1 and M/2 usually strong isthmus, pro- toconid to metaconid, M/3 occasionally with penetration of ectoflexid (or hypoconid-metasty- lid union). + 8 7. Linguaflexid: deep “V” to deep, narrow “U” shape, protocone short-to-long with narrow lingual groove (D). Usually moderate-to-broad “U” shape, with broad or irregular lingual groove on elongate protocone (B). B D B D D 8. Incisors: 1/1 to 1/3 with infundibulum or cup, an enamel surrounded pit, cement filled (C). 1/1 and 1/2 with cups; 1/3 usually with a recessed, poste- rior open “V” (O). 1/1 to 1/3 without cups or has a posterior ridge or buttons (R). 1/1 to 1/3 with deep cement filled cups, distinct lateral heel on 1/1 and 1/2, talonid on 1/3 (see subgenera dis- cussion) (El). C8 O9 C9, H10 O9 R9 9. Metapodials: long and slender; estimated indices,11 metacarpal 0.13-0.18, metatarsal 0.12-0.15 (L). Medium to long and stout; estimated indices, metacarpal 0.19-0.22, metatarsal 0.16-0.19 (M). Short and stout; estimated indices, metacarpal 0.23-0.24, metatarsal 0.19-0.21 (S). M M L12 s S E.(E.) = E. {Equus), E.{D.) — E. {Dolichohippus), E.{He.) = E. ( Hemionus ), E.{As.) = E. ( Asinus ), E.{Am.) = E. {Amerhippus). * Subgenera that occur at Anza-Borrego. 1 Transverse width at I3 divided by AP length I1 to P2. 2 Broad in Amerhippus, Hoffstetter (1952) as seen in illustrations and Rancho la Brea specimens. Rancho La Brea specimens narrower than South American species. ’’ Transverse width at I, divided by AP length I, to P,. 4 AP from junction of M' to M2 to anterior edge of orbit divided by length of tooth row. ' This also applies to the possible subgenus E. ( Onohippidion ) but separates the possible subgenus E. ( Hippidion ), which has a deep ectoflexid that penetrates the isthmus producing an antro-, meta-, and post-isthmus on the P2 through M3. See Methods and Figure 2E for an explanation of ectoflexid penetration. E. ( Onohippidion ) has a deep ectoflexid in all the lower deciduous cheek teeth (see MacFadden and Skinner, 1979). 6 Depth of ectoflexid divided by transverse width of tooth. 6 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids mexicanus, we found no cup or infundibula in the I3, and there is some evidence of an open recessed “V.” I3 of IVCM 1873 is relatively smaller than in other species compared; a minute “button” or cap- sule is at the base of the lingual edge of the “V” and is worn to a circular tip. The right canine is well developed but broken lingually and at the tip. The pre- and post-canine diastemata proportions seem to resemble E. (D.) simplicidens and E. (D.) grevyi (Table 2), although IVCM 1873 is smaller. The external enamel walls of the hypoconid in P4 through M3 are somewhat rounded or crescentic in IVCM 1873. Similar roundness occurs in E. (D.) simplicidens (see Gazin, 1936) and in Dinohippus leidyanus and D. mexicanus (see Osborn, 1918; Lance, 1950). The preserved protoconid walls of P4, M2, and M3 of IVCM 1873 are crescentic. The metaconid-metastylid anteroposterior length is somewhat shorter than in E. (D.) simplicidens. The metaconids of M2 and M3 seem to be relatively broad as in E. (D.) enormis, new species (described herein), and thus distinguished from E. (D.) sim- plicidens. The metastylid of P2, M2, and M3 is rounder and smaller than in most other species of Equus compared (see Table 3); Dinohippus mexi- canus resembles IVCM 1873 in this respect. The entoconid, as seen in the M2 and M3 of IVCM 1873, is small as in Dinohippus. The hypostylid, partic- ularly visible in the M3, is within the size ranges of Dinohippus, E. (D.) simplicidens, and E. (D.) grev- yi. The isthmus connecting the hypoconid and en- toconid in P2 is relatively wide, as is somewhat true for P3 through M3. The paralophid is well devel- oped in P3 through M3. The linguaflexids, seen only in P4, M2, and M3, are “V” shaped in IVCM 1873, thereby resembling those of Dinohippus mexican- us, D. leidyanus, and E. (D.) simplicidens (see cat- egories 1-2, Fig. 2F). There is great depth of penetration of the ec- toflexid in M, through M3; it touches the entoflexid in M, and M3 and probably in Mx. The ectoflexid index in M2 (length of ectoflexid divided by trans- verse width of tooth) is 0.69, resembling E. (D.) simplicidens (ectoflexid mean indices 0.70 to 0.71), and contrasts with Dinohippus (probable ectoflexid indices 0.61 in M, and 0.78 in M>, mean 0.70). P2 is relatively short in anteroposterior length compared to other species studied; the width is not measurable in P3 through M3 in IVCM 1873. The lengths of these teeth are shorter than in E. (D.) simplicidens (Table 3). The length of the tooth row is estimated to be 174 mm in IVCM 1873 and thus similar to the smallest E. (D.) simplicidens (173 mm to 211 mm). Dinohippus mexicanus has a mean tooth row length of 150 mm, whereas the type of D. leidyanus measures about 165 mm. The indi- vidual teeth of the young adult D. leidyanus and the mature age IVCM 1873 are generally similar, but the former has larger tooth crowns. IVCM 1873 has an estimated mandibular rostral length (I, to P2) of 98 mm and a transverse width at I3 of 59 mm. These parameters are estimated because necessary supportive matrix adheres to and covers the base of both first incisors where, like a natural cast, it preserves the dimensions and form of the ramus and rostrum. Two methods of mea- surement estimates were attempted, the first from the alveolus of P2, right side, to a point visually estimated as the base of I,, providing an antero- posterior length of 95 mm. The second method, using the same point at P2 but measured to the most posterior edge of the well-preserved occlusal sur- face of I2, which is roughly in line with the base of the Ij in other observed Equus specimens, gave an estimate of 100 mm; the mean estimate is 98 mm. Due to the absence of most of the left I3, width of palate at I3 could only be estimated; by measuring from the ventral surface and by doubling the dis- tance from the symphysis mid-point to the posterior edge of the right alveolus of I3, we obtained an estimated width of palate at I3 of 59 mm. We estimate a rostral index (transverse width di- vided by length) of 0.60 for IVCM 1873. This con- trasts with a more elongate rostrum in E. (D.) sim- plicidens (rostral index 0.41 to 0.58) and in E. (D.) grevyi (rostral index 0.43 to 0.50). IVCM 1873 re- sembles the relatively short rostrum of E. (£.) prze- walskii (rostral index 0.55 to 0.64). The holotype cranium of Dinohippus leidyanus (Osborn, 1918) (placed in Pliohippus by Osborn, 1918:162) is fig- ured in Azzaroli (1982:pl. 1, figs. 1, la). Its cranial rostral index was estimated to be 0.63 based on Osborn’s figure and 0.67 using Azzaroli’s photo- 7 Slight to deep penetration of ectoflexids in E. {Equus), ectoflexid index 0.42-0.52, mean 0.47-0.51, estimates from our data; also see Eisenmann (1981a:fig. 8). See Hoffstetter (1952) in regard to E. ( Amerhippus ), which usually has slight penetration of the ectoflexid. According to Azzaroli (1979), asses are variable; in contrast to this see Eisenmann (1981). 8 See Skinner (1972:120); applies to E. (H.) conversidens only. 4 See Eisenmann (1979a: fig. 2 or 3) and our discussion on “Burchell” zebras in our subgeneric review; variation occurs in E. burchelli and E. zebra; also see Skinner (1972:118). 1,1 E. kiang sometimes lacks cups on I, (see text p. 17). "Proximal TR divided by greatest length. 12 E. (H.) calobatus very long; E. (H.) francisi slender (Lundelius and Stevens, 1970); E. ( H .) conversidens slender (Skinner, 1972). Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 7 Table 2. Measurements of mandibles. Taxon abbreviations used in Table 2 headings and in subsequent tables, where applicable: E. {D.) enor. = E. (D.) enormis ; E. (D.) cf. simp. — E. {D.) cf. simplicidens; E. (£.) sp. A = E. ( Equus ) species A; £. (£.) sp. B = E. {Equus) species B; E. ( E .) cab. = E. {Equus) caballus; E. cf. (£.) = £. cf. {Equus); E. (D.) simp. = E. {D.) simplicidens; E. live. = E. livenzovensis h; E. sanm. = E. sanmeniensis; E. koobi. = E. koobiforensis; E. st. si. = E. cf. Dinohippus E. (D.) enor. E. (£.) sp. A IVCM 1873 IVCM 32 IVCM 1336 LACM 3667 LACM 4335 Length I, post, edge alveolus 556e 600 573e to post, edge of condyle6 (576-587e) Length rostrum, post, edge 95-100e 144e 146 154 I, to P2 (ant.) (148) Length, post, edge I, to post. 274e 362 384 376e M, (alveolus) (374) Length diastema, post, edge 73. 4e 112 124, 138 131 h to P2 (126) Length diastema, I, to 6.1 8.4, 8.4 12.4, 12.5 13.1 canine (alveolus) (ID Length diastema, canine 61 86.1,88.0 95 97 to P2 (92) Length M3 to post, edge 58-60e 175e 150 173 143 of angle (166) Length P2 to condyle 454 386 338 (420) Rostrum, transverse 59e 67 60-67 63e diameter at I3, greatest (64) Rostrum, least TR at 37e 45 45 symphysis (45) Depth mandible below M, 102 113 102 TR diameter at I3/length 0.60e 0.47e 0.39 0.40 from I, to P2, index (0.42) 1 Includes data from Winans (personal communication, 1982). 2 n = number of mandibles, not individuals. 2 Estimates from photos in Teilhard de Chardin and Piveteau (1930:pl. IV). 4 From Teihard de Chardin and Piveteau (1930:35). 5 Estimates from photo plates in Reichenau (1915). 6 A. Azzaroli personally provided us an unpublished photograph of E. livenzovensis, mandible no. 1229, derived from Bajgusheva; from this we estimated a 555 mm length for the mandible and rostral index of 0.43 in E. livenzovensis. graphs, with a mean estimate of 0.65. IVCM 1873 has a similarly short rostrum to E. (£.) przewalskii (rostral index 0.78) and E. ( Asinus ) (rostral index 0.69). Cranial and mandibular indices tend to be sim- ilar, although in general the mandible has a slightly higher index (or a shorter rostrum); therefore, we would judge the mandibular rostral index of D. leidyanus to have been about 0.66, not markedly different from 0.60 of IVCM 1823. The length of the diastema from I3 to P2 in IVCM 1873 is 73 mm and, compared with transverse width at I3 of 59 mm, provides an estimated mandibular diastemal index of 0.81. This contrasts with the much longer diastema in E. (D.) simplicidens (mean mandibular rostral index 0.59) and in E. (D.) grevyi (mandibular rostral index 0.56). The figure of Dinohippus lei- dyanus in Osborn (1918) provides an estimated diastemal index of about 0.83 compared to 0.81 in IVCM 1873. Approximately 47 mm of the ascending ramus is preserved in IVCM 1873. We estimate the ascend- ing ramus arises at an angle of 120° to 125° from the horizontal axis of the tooth row. This compares with about 120° in caballines, 130° in E. (D.) sim- plicidens, and 135° in the type of D. leidyanus as estimated from the drawing in Osborn (1918:pl. 30). The mental foramen is in a position midway be- tween the C and P2. I DISCUSSION. The M1 2 3 4 5 6 (LACM 4356) was re- ported by Downs and White (1968) as }Equus, or Pliohippus, by Opdyke et al. (1977) as cf. Pliohip- pus, by Lindsay et al. (1980) as Pliohippus, and by Lundelius et al. (1987:fig. 7.2) as cf. Dinohippus. 8 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids Table 2. Continued. stenonis sensu lato; E. st. v. = E. stononis vireti; E. = E. ( D .) grevyi; E. (E.) przw. = E. (E.) przewalskii. for a taxon; if followed by an “e,” it is an estimate idaho. = E. idahoensis; E. mosb. A number in parentheses is the i = E. mosbachensis; E. ( D .) grev. mean of the measurements given E. (E.) sp. B E. cf. E. IVCM IVCM 1816 3429 E. ( D .) simp r E. idaho. E. sanmE E. mosb.1 E. ( D .) grev.1 E. (E.) przw.1 414-544' 515e 559e 452-500 399-452 nl96(479) n8(470) n9(426) 115-136' 129 112e 148e 118-124 99-118 nl2(126) n6(123) n8(107) 292-343 325 292e 315-349e 283-302 282-296 n5(322) n4(296) n2(289) 81-117' 110, 111 94e 119, 127e 89-116 78-96 nl3(100) n7(104) n8(85) 5.5-13.0 8.2e 1.0-10.0 2.1-13 n7(7.7) n8(6.6) n4(8) 72 72-98 84e 79, 894 68-92 74-80 n8(82) n7(78) n4(78) 165 165 136-150 156 174e 131-144 124-130 n7(145) n4(136) n2(127) 420 405 341-365 387e 322-340 282-305 n4(351) n4(334) n2(293) 51-67 55-62 nl3(59) n7(57) 37-41 43 58e 35-43 41-44 n4(39) n4(39) n2(43) 118 106 92-104 116e 109e 72-85 75-101 n8(98) n7(82) n2(88) 0.41-0.58 0.51 0.39,0.46 0.43-0.50 0.55-0.64 nl2(0.48) n7(0.47) n8(0.61) No description or comparison was included in these reports. The diversity of published taxonomic ref- erences for LACM 4356 (M3) resulted from the information provided by T. Downs to various au- thors before detailed studies were completed. The teeth preserved in mandible IVCM 1873 display a diversity of characters that resemble either Dinobippus leidyanus or early E. (D.) simplicidens by the deep penetration of the ectoflexid in the IV^ through M3, a definite open recessed “V” in the I3, a “V” shaped linguaflexid, the mental foramen lo- cated midway between the C and P2, the crescentic shape of the external walls of the protoconid and hypoconid, and the well-developed paralophids. Features of IVCM 1873 shared with Dinobippus and E. ( Equus ) comprise the short rostrum, the short diastema, and possibly the more vertical slope (120° to 125°) of the ascending ramus. The relatively short rostrum (0.60 index) and I3 to P2 diastema contrast with the condition typical of the subgenus E. ( Dolicbobippus ), and the “V” or very narrow “U” shaped linguaflexid appears to eliminate affinity with the subgenus E. (Equus). Distinctive features of IVCM 1873 include rel- atively small lower incisors (especially in the 1, and I2) and a relatively wide isthmus between the en- toconid and the hypoconid of the cheek teeth. For the present, it is not deemed desirable to establish a new name for IVCM 1873. Due to a lack of sufficient material, we tentatively refer to it as cf. Dinobippus sp. If it indeed represents a species of Dinobippus, this would be the latest known oc- currence of the genus (about 2.7 Ma, middle to late Blancan). No other specimens referred to Di- nobippus have been recorded later than early Blan- can, or about 4 Ma (see Lundelius et ah, 1987). Equus Linnaeus, 1758 Characterization of Some Equus Subgenera The following comments on subgeneric character- istics are primarily based on expertise, information, and written comments provided by Morris F. Skin- ner in the late 1980s and his 1972 paper, pages 117- 129. The traditional paleontological approach to the identification of fossil Equus specimens was based primarily on cranial and upper dental char- acters, but included some postcranial data. The lower dentition was practically ignored and seldom figured in a useable manner. Recent papers by Ei- Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 9 Table 3. Measurements of lower dentition. cf. Dino- hippus E. (D.) enor. E. (D.) cf. simp. E. (E.) sp. A E. ( E .) sp. B IVCM 1873 IVCM 32 IVCM 1336 LACM 3677 IVCM 2673 LACM 4335 IVCM 1816-2 Length tooth row at P2-M3 174 229 231 222 210e 190 217 alveolus (227) I,, AP at enamel crest 13.4, 13.5 17.8, 18.0 16.0, 15. 3e (16.7) I,, TR at enamel crest 9.6, 10.2 10.8, 11.1 13.9, 15.4e (12.8) I2, AP 14.5 19.7, 19.8 17.8, 16. 7e 16. 7e 18.8 (18) I2, TR 9.2 10.5, 10.8 12.1, 14.0e 11.4 11.1 (11.8) la, AP 16.9 21.3,22.1 18.5, 17.9 17.5 (19.4) I„ TR 8.6 11.0, 11. 7e 9.8, 9.2 11.1 /C, AP at alveolus /C, TR at alveolus 14.8 9.5 20.7, 20.5 16.0, 14.5 (10.6) 17.7, 17.2 (18.4) 12.4, 12.8 (14.4) 15.7 16.5 P2, AP at enamel crest 32.3 42.1,42.6 35.2,36.2 (38) 41.8 37.5, 37.8 35.1 39.0 P2, TR at enamel crest 15.8 14.3, 14.6 13.7, 13.8 (14.7) 16.6 13.9, 12.7 14.2 16.3 P„ AP P3, tr 28.8 38.3 19.2 32.2, 34.0 (35) 19.1, 19.4 (19.6) 37.3 19.4 34.0, 34.2 19.7, 20.3 27.5 18.0 34.4 P4, AP 28.4 37.6 32.0, 34.1 (35) 36.5 32.5, 32.7 29.8 33.8 P4, TR 20.2 19.2, 20.4 (21) 20.6 18.4, 19.0 17.0 23. 5e M„ AP 23.9 32.2 29.9,31.3 (31) 31.8 29.9,31.2 28.0 31.0 M„ TR 18.4 17.8, 17.0 (18) 19.2 15.5, 16.3 15.0 17e M2, AP 24.6 33.4 31.0,31.7 (32) 32.4 29.1,30.7 27.8 30.7 m2, TR 17.7 15.7, 16.3 (17.1) 18.1 17.0, 17.2 12.9 19.3 M3, AP 30.4 39.2 40.7, 42.0 (41) 41.3 36.3,37.3 31.3 45.0 m3, tr 12.9 15.5 14.7, 15.1 (15.9) 16.6 15.4, 15.9 13.0 16.8 P3, depth ectoflexid 7.7 6.0 6.3, 8.0 (7.3) 8.9 9.4, 9.4 5.8 6.7 P4, depth ectoflexid 7.9 6.9 7.6, 8.4 (7.9) 9.0 8.6, 9.2 7.3 9.2 M„ depth ectoflexid 10.2 8.6 8.8, 8.9 (9.4) 11.5 9.2, 10.5 10.0 lO.Oe M2, depth ectoflexid 9.7 8.9 9.1, 9.4 (9.6) 11.0 9.5, 10.7 10.0 11.2 10 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids Table 3. Continued. E. cf. (E.) IVCM 3429 E. (D.) simp.1 E. idaho. E. live. E. sanmS E. koobi.1 E. (D.) grev. E. (E.) przw. 203 173-211 196, 204e 197-219. 62 200e 202e 160-189 175-182 nl8(197) (206.9) n8(179) n9(179) 16.2-16.3 12.4-17.1 17.0-19.0 n7(17.7) n6(14) n4(18) 9.1-11.6 8.8-11.5 8.5-10.0 n7(10.5) n6(10.4) n4(9.2) 9.4-20.8 14.2, 16.5 13.0-17.5 19.0-20.5 n8(18) n6(15.4) n4(19.7) 9.6-11.7 10.0 8.8-11.0 8. 3-9. 6 n8(10.4) n6(10) n4(9.0) 17.0-20.8 14.5, 16.9 13.6-17.7 18.0-20.6 n6(18.5) n7(15.6) n4(19.5) 8. 7-8. 9 10.4 5.3-10.6 9. 3-9. 7 n2(8.8) n7(9.3) n4(9.5) 5.3-17.7 21.2, 17.8e 16.9-22.0 3.6-14.1 n8(10.9) n7(18.4) n3(8.7) 3.0-4. 1 14.6, 13. 3e 12.5-15.3 3.2-11.5 n4(3.6) n8(14.3) n4(7.5) 37.3 36.1-40.0 34.0-30.4 29.1-36.9 30.3-34.4 n8(37.9) n7(34) n4(33) 15.2 16.6-17.5 12.9, 13.4 12.4-19.0 12.1-12.5 nl0(13.7) n7(13.8) n4(12) 35.7 31.0-36.1 30.9,31.0 24.8-31.2 27.7-29.6 nl0(33) n8(30) n4(29) 21.0 15.2-17.1 17.0, 17.2 13.4-17.3 15.7-16.1 nl0(16) n7(15.8) n4(16) 32.3 28.1-34.7 29.2, 29.0 25.5-30.5 27.2-29.7 nl0(32) n8(29) n4(28) 22.3 12.4-18.0 17.2, 17.4 14.4-18.8 14.9-16.6 nl0(15.4) n8(16.3) n4(16) 29.4 26.1-32.2 28.0,27.7 22.0-27 .6 24.6-28.4 nl0(29) n8(25) n4(27) 19.4 13.7-16.2 15.1, 15.2 12.9-15.4 14.2-14.7 nl0(14.7) n8(14.6) n4(14.2) 30.4 28.7-33.9 28.0, 22.8 23.3-28.4 23.2-29.1 n9(31.3) n8(26) n4(27) 18.1 13.4-15.8 15.0, 14.1 12.2-14.7 13.6-15.0 n9(14.4) n8(12.0) n4(14.3) 40.1 32.7-35.7 32.9, 33.0 22.7-32.7 28.3-30.0 n6(34.5) n8(30) n4(29) 17.7 10.2-14.0 14.3, 14.4 11.0-13.6 11.3-13.4 n8(12.5) n8(12.7) n4(12.2) 8.3 5.0— 8.6 7.2, 6.8 5. 5-7.5 4.4-6.4 nl0(7.2) n7(6.7) n4(5.5) 9.9 6.7 -9.5 8.5, 8.0 5. 8-9.2 5. 6-7.4 n9(7.7) n8(7.5) n4(6.4) 10.1 7.8-11.8 9.2, 10.0 8.7e-10.8e4 9.0-9.2e 7.6-8. 5 7.8-9. 6 7.1-7.8 n9(10.4) (9.7) (9.1) (8.1) n8(8.6) n4(7.4) 10.5 8.1-11.1 7.9, 9.2 9.4-19. 8e4 9.2-9.4e 7.6-9. 5 7.9-10.2 5. 8-7.7 n8(10.3) (10.1) (9.3e) (8.8) n8(8.6) n4(6.8) Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 11 Table 3. Continued. cf. Dino- hippus E. (D.) enor. E. (D.) cf. simp. E. ( E .) sp. A E. (E.) sp. B IVCM 1873 IVCM 32 IVCM 1336 LACM 3677 IVCM 2673 LACM 4335 IVCM 1816-2 M3, depth ectoflexid 7.2 7.3 7.9, 9.6 (8.7) 10.2 8.9, 9.9 8.0 10.7 P3, metastylid TR 9.2, 10.2 8.3, 8.6 (9.2) 8.9 9.2, 8.7 6.9 9.8 P4, metastylid TR 8.7 7.7, 8.1 (8.7) 9.2 7.4, 7.5 7.0 8.1 M„ metastylid TR 7.8 7.2, 7.4 (7.3) 6.6 6.2, 6.3 5.5 6.3 M2, metastylid TR 4.8e 7.2 6.2, 6.5 (7) 7.8 6.6, 6.8 4.6 6.0 P3, metaconid TR 8.9, 9.6 8.9, 9.0 (9.0) 9.4 8.2, 8.5 7.7 9.5 P4, metaconid TR 9.3 9.0, 9.0 (9.6) 10.1 7.7, 7.9 7.7 10.3 M„ metaconid TR 8.9 7.8, 8.7 (9.0) 10.1 6.9, 7.1 6.8 7.5e M2, metaconid TR 7.7 7.5, 7.8 (7.9) 8.7 7.9, 6.9 6.4 8.3e P3, linguaflexid shape6 1,1 2,2 2 1,1 1 3 P4, linguaflexid shape 1 1,1 1,2 2 1,3 2 3 M,, linguaflexid shape 1,1 1,2 3 1,3 2 3 M2, linguaflexid shape 1,2 1,2 3 1,3 3 3 P3, metastylid shapeT 2,2 2,2 2 3,3 3 ? P4, metastylid shape 2 2,4 2 3,3 3 ? M„ metastylid shape 2 2,2 1 4,4 3 p M2, metastylid shape P3, depth ectoflexid/TR of P3 2 0.31 2,2 0.33,0.41 (0.38) 1 0.46 4,4 0.46, 0.48 3 0.32 ? P4, depth ectoflexid/TR of P4 0.34 0.37, 0.44 (0.41) 0.44 0.45,0.50 0.43 0.39 M,, depth ectoflexid/TR of M, 0.47 0.50,0.52 (0.52) 0.60 0.59,0.64 0.67 0.59 M,, depth ectoflexid/TR of M2 0.69e 0.50 0.58,0.58 (0.57) 0.61 0.56,0.62 0.78 0.58 P3, TR metastylid/ AP of P3 0.24 0.25, 0.26 (0.25) 0.21 0.26, 0.27 0.25 0.28 P4, TR metastylid/AP of P4 0.23 0.26,0.28 (0.27) 0.25 0.23,0.25 0.24 0.24 Mh TR metastylid/AP of M, 0.24 0.24, 0.24 (0.24) 0.21 0.21,0.21 0.20 0.20 M2, TR metastylid/AP of M2 0.20 0.22 0.20, 0.21 (0.22) 0.24 0.21,0.24 0.17 0.20 P3, TR metaconid/AP of P3 0.23 0.26, 0.28 (0.26) 0.25 0.24, 0.25 0.28 0.28 P4, TR metaconid/AP of P4 0.25 0.26, 0.28 (0.27) 0.28 0.24, 0.24 0.26 0.30 M,, TR metaconid/AP of M, 0.27 0.25,0.29 (0.29) 0.32 0.23, 0.23 0.24 0.24 M,, TR metaconid/AP of M2 0.23 0.24, 0.25 (0.25) 0.27 0.22, 0.27 0.23 0.27 12 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids Table 3. Continued. E. cf. (E.) IVCM E. (E.) 3429 E. (D.) simp.1 E. idaho. E. live. E. sanmS E. koobi 7 E. { D .) grev. przw. 9.5 8.0-9. 7 10.2, 10.2 5. 8-9. 5 4.7 -6.6 n6(8.5) n8(7.8) n4(60) 10.2 6. 0-7.7 7.9, 8.1 5.2-5.9e4 7.6-7.6e 6. 7-7.4 6. 2-7. 5 6.0-7.0 n9(6.8) n3(5.6) (7.6) n3(7.0) n6(6.8) n4(6.5) 10.0 4. 7-7.3 7.1, 7.6 5.5-5.6e4 7.0-7. 4e 6. 1-7.4 5. 8-7.7 5. 1-6.3 n9(6.3) n3(5.5) (7.2) n3(6.9) n6(6.8) n4(5.8) 7.9 5.2-6. 4 5.9-6. 1 4.8-6.2e4 6.2-6.4e 5.0-5. 7 4. 8-6. 5 5.0-6.0 n9(5.8) n3(5.4) (6.3) n3(5.3) n6(5.9) n4(5.5) 7.3 5.0-6.0 6.0, 6.4 4.6-6.2e4 6.0-6.0e 5. 1-5.5 5. 7-6. 3 5. 5-6.4 n8(5.4) n3(5.1) (6.0) n3(5.3) n6(5.9) n4(6.0) 10.0 5. 7-7.7 7.8, 7.8 4.6-6.0e4 7.0-8.2 6. 7-6. 8 6. 5-7.6 7.0-7.3 n9(6.9) n3(5.3) (7.6) n3(6.7) n6(7.0) n4(7.1) 10.7 4.5-8.0 7.2, 7.8 3.2-6.44 7.6-7.7e 7.1-7.2 6. 5-8. 6 5. 6-7.0 n9(6.4) n4(5.6) (7.6) n3(6.9) n6(7.5) n4(59) 9.5 4. 7-7.6 7.0, 7.6 5.3-6.1e4 6.2- 7.6e 6.4-6. 5 6.4-6. 8 4. 6-5. 8 n9(6.3) n3(5.6) (6.9) n3(6.3) n6(6.7) n4(50) 8.0 5.5-6. 1 6.8, 6.9 4.6-6.4e4 6.4-6.4e 5. 6-5. 6 5. 9-6. 7 3. 5-6.4 n8(5.7) n3(5.6) (6.4) n2(5.6) n6(6.3) n4(4.9) 3 l(nl)3(n2)5(nl)8 2, 2 l(n6) 4(n4) 3 I(n6)2(n2) 3, 3 2(n6)3(n2) 4(n4) 1-2 I(n4)3(n4) 3, 3 I(n4)2(nl)3(n6) 4(n4) 2-3 I(n4)2(nl)3(n2) 2, 2 I(n2)2(nl)3(n4)4(nl) 4(n4) 3 I(n4)2(n2)3(n2) 2, 2 I(nl)2(n3)3(n2) 4(n4) 3 I(n7)3(n2) 2, 2 I(nl)2(n3)3(n2)4(n2) 2(n2)4(n2) 4 I(n6)4(n2) 2, 2 I(n2)2(n3)3(n3) 2(n2)4(n2) 3 l(n5)3(nl) 2, 2 I(n3)2(n2)3(n2) 2(n2)4(n2) 0.40 0.32-0.55 0.40, 0.42 0.31-0.48 0.47e 0.37-0.49 0.27-0.41 nl0(0.45) n3(0.40) n7(0.42) n4(0.35) 0.44 0.41-0.61 0.45, 0.46 0.38-0.60 0.54e 0.37-0.63 0.35-0.46 n9(0.49) n3(0.49) n8(0.45) n4(0.41) 0.52 0.55-0.79 0.68,0.68 0.70-0.71 0.58e 0.52-0.68 0.48-0.55 n9(0.70) n4(0.71e) n8(0.59) n4(0.51) 0.58 0.60-0.76 0.65,0.71 0.50-0.75 0.63e 0.55-0.73 0.41-0.53 n8(0.71) n4(0 .66) n8(0.63) n4(0.47) 0.29 0.18-0.27 0.26 0.19 0.22 0.20-0.24 0.21-0.24 n6(0.19) n6(0.23) n4(0.23) 0.31 0.15-0.24 0.25 0.17 0.21 0.23 0.19-0.27 0.19-0.27 n9(20) n6(0.24) n4(20) 0.27 0.16-0.24 0.22 0.18 0.21 0.17 0.19-0.24 0.21-0.21 n9(0.20) n6(0.22) n4(0.21) 0.24 0.15-0.20 0.22 0.18 0.22 0.19 0.20-0.25 0.22-0.23 n8(0.17) n6(0.22) n4(0.23) 0.29 0.16-0.24 0.22e 0.22e 0.21 0.21-0.26 0.25-0.26 n9(0.21) n6(0.24) n4(0.25) 0.33 0.13-0.25 0.25e 0.17 0.22e 0.23 0.21-0.28 0.19-0.27 n9(0.20) n6(0.26) n4(0.23) 0.32 0.16-0.26 0.24e 0.19 0.23e 0.23 0.23-0.27 0.12-0.23 n9(0.21) n6(0.25) n4(0.20) 0.26 0.16-0.20 0.21e 0.19 0.21e 0.20 0.21-0.25 0.12-0.27 n8(0.18) n6(0.24) n4(0.19) Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Liquids H 13 senmann (1975, 1976, 1979a, b, 1981a, 1984, 1986) have, however, greatly increased our knowledge of methods of evaluation of cranial, mandibular, den- tal, and postcranial elements. The natural groupings that represent subgeneric rankings within equid genera from the late Plio- cene, early Pleistocene, and Holocene have re- ceived little attention. Hoffstetter (1952) considered characters of the lower incisors, Azzaroli (1966, 1979, 1982) and Bennett (1980) dealt with skull shapes, lateral views of crania, and mandibles, and Harris and Porter (1980) with postcranial measure- ments. Forsten (1988) separated the genus Equus into two “groups” based on the shape of the “en- toflexids” (= linguaflexid) in lower cheek teeth. Churcher and Richardson (1978) and Eisenmann (1986) have also discussed subgeneric ranking. Comparisons of the subgeneric characters that we interpret to be useful in Equus analyses (Table 1) include proportional length of the cranium and mandibular rostrum, location of the orbit, char- acters of the lower incisors and lower cheek teeth, and relative length and robustness of the metapo- dials. Skeletal and dental parts that are most com- monly available, or better preserved, have been em- phasized. None of these characters can be described in terms of presence or absence, nor are there pre- cise metrics for any one taxon. All show variation and mean trends. Some subgeneric groups appear reasonably distinctive on the basis of character state combinations. Although we are convinced of the usefulness of subgeneric taxa, they may not have become clearly distinguished until middle to late Pleistocene or Holocene time (also see Azzaroli, 1979, 1982). Skinner (1972:118) proposed that the similarities between the living zebra, Equus grevyi, and the North America fossil E. simplicidens justified plac- ing them in the subgenus Dolichohippus, and that the Pliocene E. (D.) simplicidens was ancestral to the African E. (D.) grevyi. Churcher and Richard- son (1978:403) stated, “The occurrence of doli- chohippine equids in Africa by the middle to late Pliocene would appear to preclude the latter as- sertion,” i.e., E. (D.) simplicidens to E. (D.) grevyi descent. Nevertheless, these authors stated that (p. 407) “. . . the largeset E. capensis and E. oldoway- ensis and the more complete skeletal material de- scribed for E. simplicidens (Skinner, 1972) conform to expectations for precursors of the modern E. grevyi .” Winans (1985) emphasized that, because the ho- lotype upper check tooth of E. (D.) simplicidens lacks adequate diagnostic features, this taxon should be construed as a nomen dubium. She stated that the sample representing E. (D.) shoshonensis from Hagerman, Idaho, provides more adequate material to characterize its “zebrine” affinity, such as the deep penetration of the ectoflexid and presence of P* 1 2 3 4 5 6 7 8. Berger and Howe (1987) also noted the possible relationship of E. (D.) shoshonensis to E. (D.) grev- yi. We refer to the Hagerman sample as E. (D.) simplicidens, following Skinner (1972) in consid- ering E. (D.) shoshonensis to be synonymous with E. (D.) simplicidens. Winans (1989:292-294) proposed five “ Equus species groups”; these groups include: Equus sim- plicidens, Equus scotti, Equus laurentius, Equus francesci, and Equus alaskae. On page 292, Winans listed, and seemed to rec- ognize, E. simplicidens and E. shoshonensis as dis- tinct species within her E. simplicidens “group,” contrary to her 1985 position. As noted above we follow Skinner (1972:119) and consider Equus (D.) shoshonensis to be synonymous with E. (D.) sim- plicidens. With reference to her Equus scotti group, Winans indicated, “Site samples referred to this group: All specimens from Rock Creek, Bautista Creek, Grandview, Irvington, Gilliland, Pool Branch, Port Charlotte, Vallecito, American Falls, and In- gleside; large horses from Hay Springs, Arkalon.” Winans may have overlooked the fact that many of the samplings of material referred to Equus (“large and small” or “Equus”) from the Vallecito Creek area, in the Anza-Borrego Desert, are derived from sediments containing the “Vallecito Creek” and “Arroyo Seco” faunas. They were collected within approximately 3,700 feet of stratified sediments, ranging in age from about 2.3 Ma to 1.0 Ma; see Downs and White (1968) and Opdyke et al. (1977: 323). On the basis of M. F. Skinner’s concept of Equus grouping (personal communication and Skinner, 1972:117-125), we characterize five subgenera of Table 3. Continued. 1 Includes data from the authors and Gazin (1936). 2 From Bajgusheva (1798). 3 From Eisenmann (1983), estimate from photo plate 5.9. 4 Estimate from Gromova (1949), line drawings. 5 Estimates from Teilhard de Chardin and Piveteau (1930), photo paltes. 6 See Figure 2F for linguaflexid shape categories. 7 See Figure 2G for metastylid shape categories. 8 l(nl),3(n2) indicates: 1 specimen, category 1; 2 specimens, category 3; etc. 14 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids Equus in North America, including dolichohippine, caballine, hemionine, asinine, and amerhippine (see Table 1). The amerhippine horses are recorded from North America and South America. The other subgenera occur in North America and Eurasia. Equus ( Dol - ichohippus ), Equus ( Equus ), Equus ( Hemionus ), and Equus ( Asinus ) all probably occur in the Anza- Borrego Desert of California. Equus ( Hemionus ) and Equus ( Asinus ) will be described in a forth- coming general account of the Anza-Borrego se- quence. Winans (1989:296) noted that in a relatively short period of time (3.5 million years), the North Amer- ican genus Equus increased from one to up to four contemporary species, suggesting Equus was un- dergoing the first stages of radiation. Our studies also suggest radiation or diversification, although we interpret this on a subgeneric level (see Table 1). DESCRIPTIVE TERMINOLOGY Linguaflexid Shape The metastylid in the lower dentition of the equids was formed by crown fission, or splitting of the metaconid, as stated by Stirton (1941:440) in con- trast to Osborn (1907). The variation in shape of the metaconid-metastylid (the linguaflexid of Skin- ner, 1972) in the Equidae may be viewed as evo- lutionary stages that can be used as temporal in- dicators. Practically all the late Oligocene equids exam- ined (Whitneyan, 30 Ma to 21.6 Ma from the Agate Ash) have a weak cleft on the central crown of the metaconid (see Stirton, 1941:441, fig. 9). Stirton (1940) stated that there is a faint indication of twin- ning or division of the two cusps, apparently de- rived from one in Hyracotherium. This cleft, or twinning, became more prominent as the metastylid expanded posteriorly, with the ultimate production of the metaconid-metastylid column. During Oligocene through Miocene time, the metastylid became a distinct cone by crown fission of the metaconid in early forms such as Mesobippus and Miohippus, and it continued to be more strong- ly expressed in later Miocene equids such as Para- hippus color adensis (see Stirton, 1941:434, fig. 44), Anchitherium Meyer, 1844, and Merycbippus Lei- dy, 1857 (Stirton, 1940:figs. 12, 19). By late Hemingfordian time, specimens of Para- hippus Leidy, 1858, from the Runningwater For- mation of Nebraska (17.5 Ma) show the metaco- nid-metastylid columns as distinct entities in which the protoconid alone united directly with the meta- conid of the premolars and molars. In some spec- imens of Parahippus, the hypoconid united directly to the split, posteriorly expanded part of the meta- conid that had become a distinct entity, the meta- stylid. This resulted in varying degrees of junction and union of these separate parts by structures called “isthmuses” (Skinner, 1972). The presence of isth- Contributions in Science, Number 440 L_= 1 5cm Figure 3. cf. Dinobippus sp.; four upper molars from the Anza-Borrego Desert, Fish Creek area, southern Cal- ifornia, occlusal and lingual views; (a) LACM 4356, right M' “zone” 4, pre-Layer Cake local fauna, late Hemphil- lian; (b) IVCM 2257-3, right M1; (c) IVCM 2257-2, right M-; (d) IVCM 2257-1, right ML IVCM 2257-1-2-3 from “zones” 7-8, early Layer Cake local fauna, transitional late Hemphillian to early Blancan. muses is now an important feature in the suite of characters used to distinguish some Equus subgen- era (see Skinner, 1972; and Table 1, herein). Ectoflexid The deep invagination or valley, called the ecto- flexid by Skinner, 1972 (and see Fig. 2E) separates the hypoconid from the protoconid on the lower cheek teeth in some equids. There may be a cor- relation between an increase in the amount of den- Downs and Miller: Late Cenozoic Equids ■ 15 a i i Q 5 cm Figure 4. a. cf. Dinohippus sp., from Arroyo Seco local fauna, “zones” 44-45, late Blancan, IVCM 1873, mandible, occlusal view, with incisors, b. cf. Dinohippus sp., IVCM 1873, left mandible, lateral view. c. cf. Dinohippus sp., IVCM 1873, left P,-M,, occlusal view. tine in the isthmuses connecting the protoconid and hypoconid to the metaconid and metastylid and increasing hypsodonty and a shortening of the ec- toflexid. There is also a correlation of the length- ening of the postflexid in the molars with shortening of the ectoflexid, especially the Mj and M2 (see Eisenmann, 1983). The molars became more rect- angular, or more “molariform,” in E. ( Asinus ) and E. ( Hemionus ). The apparent increased amount of exposed enamel and dentine served to enhance the grinding surface, probably a useful adaptation for grazing. However, E. (D.) simplicidens, E. sanmen- 16 ■ Contributions in Science, Number 440 iensis, E. (D.) grevyi, and E. ( Equus ) przewalskii tend to retain the deep ectoflexid in M1? M2, and M3. Equus (D.) enormis, new species, and E. (D.) cf. simplicidens of this report tend slightly toward a complete isthmus correlated with a shallow ec- toflexid. Equus ( Hemionus ) and E. ( Asinus ) have molars that are rectangular, with a complete isth- mus, except that occasionally the M3 in E. ( H .) conversidens (Owen, 1869) assumes the caballine type of junction or hypoconid-metastylid junction of Skinner (1972:120). Recently, Forsten (1988:23-24) proposed that I Downs and Miller: Late Cenozoic Equids Equus can be separated into two main groups mor- phologically, characterized by the enamel pattern of their lower cheek teeth, particularly by the shape and form of “divergence” of the metaconid and metastylid. She noted stenonid equids are charac- terized by . . a ‘V’ shape entoflexid between the metaconid and metastylid and caballoid equids by their ‘U’ shape entoflexid.” It is our belief that Equus groupings may require consideration of cranial and mandibular proportions (especially rostral), dental characteristics, and metapodial proportions. Also, we are following Skinner (1972), who referred to the metaconid-metastylid “divergence” as the lin- guaflexid, and not the entoflexid of Forsten (1988). Stirton (1941 :fig. 6) showed the entoflexid to be located between the metastylid and entoconid. Incisor Cups The presence or absence of infundibulae, or cups, on the lower incisors, especially I3, is another char- acter which may be used in differentiating Equus at the subgeneric level. However, Skinner (1972: 118) stated, “The presence or lack of cups in I3, although a useful character for specific separation, is not applicable for subgeneric separation of equids.” Since 1972, additional studies have been made, and it is our opinion that features such as isthmuses, ectoflexids, linguaflexids, rostral pro- portions, orbital and narial notch positions, meta- podial proportions, and lower incisor cup mor- phology may be jointly employed in subgeneric classification. The dominant tendency in the Equidae is for the post-cingulum of the lower incisors to become more hypsodont with time to form enamel-lined pits or infundibulae (Fig. 2A-D). It is by this process that the posterior border of the pit could become sur- rounded by enamel. We avoid the term “commis- sure” (Bennett, 1980:280, fig. 2E) for the open “V” on the posterior side of the lower incisors because this is not a junction of two parts unless the right and left sides of an incisor are considered as such. This is particularly true in I, and I, of the caballinid and ass-like forms. The lack of cups such as found in E. ( Amerhippus ) Hoffstetter, 1950, may also be due to the absence of post-cingula development, a condition observed in some early equids. The pre- dominantly recessed cup, or open “V,” on the I3 of burros, Equus (. Asinus ), and long-headed zebras, E. ( Dolichohippus ), is here interpreted as a failure of the posterior part of the third incisor cingulum to become hypsodont. We do not recognize the occurrence of a “half-infundibulum” (Bennett, 1980: fig. 2E). Azzaroli (1982:92) stated, “There is no proof that the lack of cups is a plesiomorphic feature, on the contrary, paleontological evidence seems to indi- cate that this is a relatively late ‘generalization.’ ” Eisenmann (1979a:55) noted that it is possible to assume that the presence of cups “is a plesiomor- phous character for the genus Equus (though an apomorphous one for the Equidae in general).” With Contributions in Science, Number 440 respect to the loss of cups, she stated, “This loss is the result of a parallel evolution expressing a common evolutive tendency rather than a synapo- morphy.” She recorded a lack of cups in equids from the Americas, Eurasia, and Africa. It is possible, in our estimation, that the presence of a cup is apomorphic, or derived, in early Equus species such as E. simplicidens and E. idahoensis, where a cup is beginning to form as in I3. Our studies indicate that a lack of cups is plesiomorphic, or primitive, in early, less hypsodont Equidae. Most of the hemionids, E. ( Hemionus ) and per- haps E. “(H.) zebra hartmanni ,” have cups on I,, I2, and I3. The Mongolian kiang, E. “(H.) hem- ionus,” also has well-developed cups on all the incisors, but the Indian kiang, or kiang of Kutch, E. “(H.) kiang” lacks cups on the third incisors. Hoffstetter (1950) was one of the first to point out the importance of the total lack of cups on the incisors when he established the genus Amerhippus with the type species Equus andium, now referred to as Equus ( Amerhippus ) andium. We have ob- served numerous examples of this incisor condition in other equids, particularly the Rancho La Brea horse commonly known as Equus “ occidentalis ” (Leidy, 1865), and here considered as E. (Amerhip- pus) “ occidentalis ” (Leidy, 1865), as was also sug- gested by Hofstetter (1950). E. ( Amerhippus ) is perhaps more widespread in North America than previously realized, although rare in most collections. Churcher (1985) reported the “genus” Amerhippus from the Sangamon of Medicine Hat, Alberta, Canada. Lower jaws of a species of E. ( Amerhippus ) have been found in the late Pleistocene Sheridan Deposits of the Hay Springs local faunas of northwest Nebraska, as well as a smaller species from the Bondalier tuff, dated at 1.5 Ma in New Mexico (personal communication from Skinner). Eisenmann (1975) described E. teil- hardi from the early Pleistocene of Nihowan, Chi- na, a species characterized by the absence of lower incisor caps. Winans (1985:293) stated, “All spec- imens from Rancho La Brea . . . are referred to her E. laurentius group.” Other included “types” are E. mexicanus, Hibbard, 1955, and E. midlandensis, Quinn, 1957. Some species of Equus have developed rather prominent talonids on the external posterior por- tion of the lateral incisors such as those on the living E. (EE) hemionus. Another variant was ob- served by Hofstetter (1950:690) when he estab- lished the generic name Pseudoquagga for one of the Burchell zebras in Africa. He pointed out that Pseudoquagga, or the subgenus Equus (Pseudo- quagga) lacks cups on the forward edge of the blade-like lower incisors similar to the condition found in E. (Amerhippus). However, E. (Pseudo- quagga) has a semi-posterior “cingulum” in the form of small, bead-like outgrowths on the pos- terolingual side of the incisors. In 1982, T. Downs inspected samples labeled E. “ burchelli ” (= E. quagga tbahni according to Churcher, personal communication, 1992) from the National Muse- Downs and Miller: Late Cenozoic Equids H 17 ums of Kenya, Nairobi. In all seven specimens, cups were present wherever there was sufficient wear to reveal them. The lack of cups in the lower incisors was also noted by Cope (1892) when he proposed the generic name T omolabis. As emphasized above, these varying conditions of the lower incisors do not permit the use of the presence or absence of the cup as a single diagnostic character. However, in conjunction with the other characters of the lower cheek teeth, skull, and limbs, the nature of the cup apparently provides a useful indication of taxonomic affinity. Dolichocephaly Dolichocephaly appears to have its inception in £. (D.) simplicidens, as noted in the scattergram of the width and length of the rostrum (Fig. 14). How- ever, the snout of E. (D.) simplicidens ranges in variation from a short snout (rostral index 0.60) to a considerably elongated one (rostral index 0.43). The short snout is a trait common in E. { Equus ) and E. { Asinus ) and possibly retained from the pos- tulated ancestral genus, Dinohippus. The elongate versus short condition of the pre- maxillary symphysis, rostrum, and palatal-facial re- gion represent dolichocephalic and brachycephalic tendencies, respectively. Dolichocephaly is ob- served in the long-headed African zebra, E. (D.) grevyi, and in its North American relatives E. (D.) simplicidens and E. (D.) enormis, new species that have been recovered from the Anza-Borrego Desert of California; Hagerman, Idaho; Blanco, Texas; Benson, Arizona; Las Cruces, New Mexico; and other localities. Skinner (1972:118) interpreted a longer, more slender rostral area to accompany lon- ger-headedness as diagnostic characters for the sub- genus Dolichohippus. Bennett (1980:284) noted Equus burchelli, E. andium, E. stenonis Cocchi, 1867, and E. (A.) “ occidentalism of Rancho La Brea also displayed dolichocephaly and possessed long narrow snouts. However, as we interpret them, the Rancho La Brea horse, E. burchelli, and E. andium are not dolichohippine; their rostral indices range from 0.56 to 0.69 for the Rancho La Brea horse, 0.61 to 0.65 for E. andium, and 0.52 to 0.55 for Equus burchelli, as deduced from measurement of figures in Azzaroli (1966). None of these rostral indices suggest dolichocephaly (see Fig. 14 and Ta- ble 1). The position of the orbit relative to the tooth row, particularly the M3, is also of value in deter- mining dolichocephaly. The orbit is generally pos- terior to the M3 in dolichohippines (Azzaroli, 1966; Skinner, 1972). The position of the naso-maxillary notch, or narial notch, relative to the P2 is also of some value in subgeneric character comparisons, being deep in E. {Equus) and E. { Dolichohippus ), above the posterior edge of the P2 or near the P3, compared to being more anterior in other subgen- era. These features, rostrum, palate, orbit, and pos- sibly narial notch positions, are not only useful, but are frequently preserved in the available fossil ma- terials. Thus, they are especially useful in judging the degree of dolichocephaly. Metapodial Proportions Metapodials are relatively common in association with cranial, mandibular, and dental elements in fossil equids. In general, there are three types of metapodials: (1) Medium-to-large size found in the relatively tall dolichohippines such as E. (D.) grevyi, E. (D.) enormis, new species, and E. livenzovensis ; this may also include E. stenonis sd. (see Azzaroli, 1982; Eisenmann and Karchoud, 1982) and E. (E.) caballus. (2) Elongate, or slender, metapodials as in hemionids— - -especially the North American fos- sil E. { Hemionus ) calobatus (Troxell, 1915), the small but equally slim E. (H.) francisi (Hay, 1915), possibly E. (EL) conversidens, and the modern Asi- atic kiang. The metatarsals of the kiang are partic- ularly slender. (3) Short-to-stout metapodials as in E. koobiforensis, the asses, £. (A.) scotti (Gidley, 1901), E. (A.) asinus, the non-dolichohippine ze- bras such as E. burchelli and E. zebra, and the amerhippines such as E. (A.) “ occidentalis ” and E. (A.) andium . There appears to be a trend toward larger skull size and a larger postcranial skeleton in some late Pliocene to middle Pleistocene dolichohippine and caballine equids (see Figs. 13-17). They are larger than modern, or non-domestic, wild Equus. This large size is observable in E. sanmeniensis, E. liv- enzovensis, £. mosbachensis Reichenau, 1915, and £. (D.) enormis, new species, and probably in £. idahoensis, E. {Equus) sp. B, and £., cf. (E.) {Equus) sp. of this report. Our current study proposes to show that E. (D.) simplicidens, £. livenzovensis Bajgusheva, 1968, E. sanmeniensis Teilhard de Chardin and Piveteau, 1930, E. koobiforensis Ei- senmann, 1983, and possibly £. stenonis s.l. could be potential precursors of E. (D.) grevyi. During the Pliocene to Pleistocene, Equus spe- cies display increasing diversity. There seem to be some taxa or populations from North America, Eurasia, and Africa with mixed subgeneric tenden- cies, especially merging dolichohippine and cabal- line characteristics — for example, £. idahoensis, Equus cf. E. caballus (Savage, 1951) of Irvington, and Equus. { Dolichohippus ) cf. £. D. simplicidens of Anza-Borrego in North America have a relatively short or caballine rostral area, or usually a deep- to-moderate penetration of the ectoflexid in the lower molars, and usually no cup or recessed “V” in the I3. In Eurasia, £. mosbachensis has a mod- erately elongate rostrum, the orbit is posterior to the M3, the protocone in the upper molars is broad- ly grooved and elongate, and the ectoflexid in the lower molars is generally deep. However, the lin- 18 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids guaflexid has a broad “U” shape as in the caballines (see Fig. 2E, F for character states). All of these equids occur in the holarctic during Plio-Pleisto- cene time. Hibbard (1955) described a new subgenus and species, Equus ( Hesperohippus ) mexicanus from the Tequixquiac fauna of Mexico. The subgenus is primarily distinguished by a high degree of cranial flexion and anteriorly flattened premaxilla of the holotypic skull. A ramus with P2-M3 is identified as Equus cf. ( Hesperohippus ) mexicanus in Hib- bard (1955:fig. 5B) and as }Equus ( Hesperohippus ) mexicanus in his table IX. The proportions of the rostrum (based on measurements of the cranium in Hibbard, 1 955:pl. IV, fig. 1) indicate a very short rostral index of 0.69 as in E. (Equus), E. ( Asinus ), E. ( Hemionus ), and E. ( Amerbippus ). The referred lower dentition has a broad “U” to “V” shaped linguaflexid, non-penetration of the ectoflexid in M,, M2, and M3, and straight walls of the ectolo- phids. In the upper dentition, the protocones are very elongate and grooved, and the pre- and post- fossettes have very complex enamel patterns. Hib- bard’s material could be either E. ( Hemionus ) or E. ( Amerbippus ) because of the shallow ectoflexid in the molars. Without further examples of E. (H.) mexicanus to suggest the validity of the subgeneric characteristics, we tentatively reject the material as a distinct subgenus of Equus. The subgenus E. (Para sty li dens) parastylidens Mooser and Dalquest, 1975, from central Mexico is based on lower dentition. We believe that with- out the availability of cranial, mandibular, or rostral features for comparison, there is insufficent basis for a reliable subgeneric diagnosis. A key for the subgenera of Equus is presented below. KEY TO SOME SUBGENERA OF EQUUS 1A Cranium and mandible rostra elongate; orbit posterior to tooth row 2 IB Cranium and mandible rostra short; orbit an- terior to tooth row 3 2 Ij_2 with cups; I3 with recessed posterior open “V”; ectoflexid penetrates isthmus on M,_3; deep linguaflexid with narrow lingual groove; stout, medium-to-long metapodials . . Dolichohippus 3A M,_3 with strong isthmus but no ectoflexid pen- etration 4 3B M,_3 ectoflexid penetrates and splits isthmus 5 4A h_3 with cups; broad “U” shaped linguaflexid with broad lingual groove; long and slender metapodials Hemionus 4B I,_2 with cups but I3 with recessed posterior open “V”; deep “V” shaped linguaflexid with narrow lingual groove; short and stout meta- podials Asinus 5 Broad “U” shaped linguaflexid with broad lin- gual groove 6 6 It_3 with cups; medium-to-long and stout meta- podials Equus Equus ( Dolichohippus ) enormis, new species Figures 5-17 DIAGNOSIS. About 7 percent longer cranium and mandible than the largest Equus ( Dolichohip- pus), ranging from 616 mm to 660 mm basilar length and 573 mm to 600 mm mandibular length; with greater elongation of the rostrum and palate (rostral index range 0.40 to 0.45, mean 0.43; palatal index 0.21 to 0.24, mean 0.23); orbital position posterior to M3; adult upper incisors and lower third incisor massive; lower third incisor without infundibulum and with a lingual open recessed “V”; ectoflexid in M„ M2, and M3 penetrates isthmus, although with somewhat less penetration than in E. (D.) simplici- dens and E. (D.) grevyi; metastylid in P3 and Mr M2 broad transversely, triangular to rounded an- teriorly, with or without lingual indentation; meta- conid broad transversely in P3 and M2; radius-ulna with a straight mid-shaft, flattened anterior surface, and fusion of the ulna and radius distal to the fo- ramen separating the two; relatively small meta- carpal 3 and phalanges 1 and 3; metatarsals 3 and 4 relatively larger than in other fossil Equus except perhaps E. livenzovensis and E. mosbachensis. HOLOTYPE. IVCM 32 from locality IVCM 15; partial cranium and right and left mandibles with postcranial elements including: right distal humer- us, nearly complete right radius-ulna, complete left metacarpal 3 with associated articulating unciform, magnum, trapezoid, and a proximal metacarpal 4; phalanges 1, 2, and 3 of the manus; partial pelvis, right femur head and shaft fragment, metatarsal 3 with portions of attached metatarsals 2 and 4 and a phalanx 3 of the pes; mature, five to six year old male. PARATYPE. LACM 4338, partial cranium, with relatively complete ventral palato-rostral area from I1 to posterior of left, post-glenoid process, dorsal aspect from premaxillae to portions of parietals, cast of braincase and original posterior squamosal, mature to old age male, from locality LACM 1528. REFERRED SPECIMENS. LACM 16815, max- illa with partial rostrum, complete maxillary den- tition, associated pelvis, immature, probable male, from locality LACM 6606; LACM 3677, nearly complete left mandible with P2 to M2 and right symphysis, mature to old age male from locality LACM 1253; IVCM 1336, complete right and left mandibles with dentition, mature to old age male from locality IVCM 371. TYPE LOCALITY. IVCM 15, from Vallecito Creek, Anza-Borrego Desert State Park, San Diego County, California, “zone” 54, upper Palm Springs Formation, middle Vallecito Creek local fauna, late Blancan in age (late Pliocene), approximately 2.0 Ma. Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 19 Figure 5. a. Equus ( Dolichohippus ) enormis, new species, holotype, IVCM 32-1 and IVCM 32-2, from Vallecito Creek local fauna, zone 54, late Blancan, palate and rostrum, occlusal view. b. Equus ( Dolichohippus ) enormis , new species, IVCM 32-4, partial mandible, occlusal view. Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 21 Figure 6. a. Equus ( Dolichohippus ) enormis, new species, IVCM 32-1, right upper dentition, occlusal view. b. Equus {Dolichohippus) enormis, new species, IVCM 32-4, right lower dentition, occlusal view. Figure 7. Equus ( Dolichohippus ) enormis, new species, holotype, IVCM 32-1 and IVCM 32-2, maxillae and rostrum, right lateral view. Hatchured areas in this figure, and any succeeding illustrations, indicate restored material. LOCALITIES FOR HYPODIGM. LACM 1528 (paratype) and LACM 6672 from Vallecito Creek area; LACM 1253 from badlands east of Borrego Springs, near “Truckhaven Trail”; IVCM 371 from lower Coyote Canyon badlands north of Borrego Springs; all localities from Anza-Borrego Desert State Park, San Diego County, California. STRATIGRAPHY AND AGE. The holotype, IVCM 32, is from within the type stratigraphic sec- tion of the Palm Spring Formation, late Blancan, “zone” 54. Approximately 2.0 to 2.1 million years old, it predates the Olduvai Gorge sequence of East Africa (see Opdyke et ah, 1977). The paratype, LACM 4338, from locality 1528 is from sediments of fault block K, of the Palm Spring Formation just south of Vallecito Creek Wash and approximately equivalent to LACM locality 1190, “zones” 62-63 of the type stratigraphic section; the sediments and faunal taxa are considered correlative with the late Vallecito Creek local fauna, middle Irvingtonian age. The referred immature palate, LACM 16815, is from “zone” 58 at locality LACM 6606 and is about 250 meters higher in the type section and approximately one half million years younger than the holotype, IVCM 32. Referred mandibles IVCM 1336 and LACM 3677 are geographically separate, being from about 30 miles north of the type section, in sediments of late Irvingtonian age, possibly 0.3 Ma to 0.72 Ma. LACM 1528 (paratype) fault block K and ap- proximately equivalent to “zones” 62-63 of the type section in Upper Palm Springs Formation, Val- lecito Creek local fauna, middle Irvingtonian in age (early-middle Pleistocene), approximately 1.0 Ma; LACM 6606, estimated “zone” 58, Upper Palm Springs Formation, late Vallecito Creek local fauna, early Irvingtonian in age (early Pleistocene), ap- proximately 1.6 Ma. LACM 1253, Palm Springs Formation of Dibblee (1954) or Ocotillo Formation of Bartholemew (1970), undescribed Borrego Badlands local fauna, probable late Irvingtonian (middle Pleistocene), as suggested by presence of Mammuthus Burnett, 1830, Equus (Equus), Megalonyx Harlan, 1825, cf. Nothrotheriops, and Arctodus Leidy, 1854, among other taxa. IVCM 371, Ocotillo Formation, un- described Coyote Canyon badlands local fauna, probable late Irvingtonian (middle Pleistocene), as suggested by presence of Lepus ? Linnaeus, 1758, Equus ( Dolichohippus ), Equus ( Hemionus ) ?Eu- ceratherium Furlong and Sinclair, 1904, Nothroth- eriops, and Tetrameryx Lull, 1921, among other taxa. Miller et al. (1988) presented radiometric dates for the Fryant ash of the Coyote Canyon badlands, ranging from 0.3 Ma to 0.72 Ma. There is about a 2.1 Ma to 0.3 Ma range in time or 1.8 Ma duration for the species; therefore, we cannot assume that all specimens of the new species are synchronous, although they are probably mor- phologically one taxon. ETYMOLOGY. From Latin enormis, of great size, huge, immense. DESCRIPTION AND COMPARISONS. There is no single diagnostic character for the species; the diagnosis is based on a unique combination, or suite, of characters that is not duplicated in other known taxa represented by comparable material. After further collecting and study, it is possible that the specimens referred to E. (D.) enormis might be interpreted to represent more than one species; for the present we interpret observed variation as in- : traspecific — in contrast to the “single tooth” jus- tification of some previously described species. In the following discussion, we use Equus ( Dolicho- hippus) simplicidens as a primary standard for com- parisons. Cranial and mandibulae lengths in E. (D.) enor- mis indicate generally larger size in E. (D.) enormis than in other fossil Equus species (see Fig. 12). Proportions of the cranial rostrum appear sig- 22 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids nificant when comparing E. (D.) enormis with other taxa. About 22 mm of missing bone has been re- stored between the canines and P2 in the holotype, IVCM 32, the estimate of length of rostrum in this specimen is based on extrapolations from mea- surements of length between the upper canine and P2 in more complete specimens of Anza-Borrego dolichohippines. The latter include LACM 4338, the paratype of similar geochronologic age as IVCM 32 (middle Vallecito Creek local fauna); LACM 16815, from the late Vallecito Creek local fauna; and specimens referred to Equus ( Dolichohippus ) cf. E. (D.) simplicidens— IVCM 2673 from the ear- ly Vallecito Creek local fauna and LACM 17614, partial cranium of probable equivalence to the late Arroyo Seco or early Vallecito Creek local faunas. Using these specimens, we compared length from upper canine to P2 with the total length of the tooth row. For LACM 4338 these two figures are 96.7 mm and 203.8 mm, respectively. As IVCM 32 has a longer tooth row length of 217.5 mm, the pro- portional distance for C to P2 should be 103 mm (= 217.5 x 96.7/203). Other extrapolated estimates of C to P2 for IVCM 32 are: 78 mm based on LACM 16815, 106 mm based on LACM 17614, and 76 mm based on IVCM 2673. The mean estimate is 91 mm for IVCM 32 in C to P2. Measurements that are available in the rostrum of IVCM 32 are: I1 to I3, 42 mm; I3 to C, 30 mm; and the basal length of canine, 18 mm. These dimensions plus the mean estimate for C to P2 of 91 mm provide a combined total estimate of 181 mm for the length of the rostrum of IVCM 32. The rostrum is 175 mm long in LACM 4338 and 157 mm in LACM 16815. The estimated mean rostral length is thus 171 mm in E. (D.) enormis. The transverse diameter at I3 is 70 mm in LACM 4338, 70 mm in LACM 16815, and 79 mm in IVCM 32; the mean is 73 mm. These proportions (trans- verse width at I3 divided by length from I1 to P2) indicate a greater elongation of the rostrum than in other species compared (see Fig, 14). Rostral indices of 0.44 in IVCM 32 and 0.40 in LACM 4338 seem particularly significant; the immature specimen LACM 16815, with a rostral index of 0.45, is closer to proportions of E. (D.) simplicidens (mean rostral index 0.50) and E. (D.) grevyi (mean rostral index 0.48) although outside their range of variation (see Table 2). Equus (D.) enormis has an estimated rostral index of 0.40 to 0.45 (mean 0.43). Winans (1985) noted that, because of different rates of growth in different parts, use of ratios to compare proportions can be valid only for speci- mens of the same ontogenetic age. IVCM 32, LACM 4338, LACM 3677, and IVCM 1336 are relatively mature adults, but even the immature LACM 16815 (rostral index 0.45) indicates relatively long rostral length. The position of the anterior edge of the orbit relative to the cheek teeth may be used as an in- direct indication of dolichocephaly and is cited as a subgeneric character by Azzaroli (1966:11). The distance from the posterior edge of the M1 to the anterior edge of the orbit in the paratype LACM 4338 is 100 mm. This compared with the mean tooth row length of 204 mm provides an orbital index of 0.49; in contrast, the orbital index 0.41 to 0.45 (mean 0.43) in E. (D.) simplicidens indicates this species has a more anteriorly located orbit than in E. (D.) enormis. The dorsoventral depths of the cranial pre- and post-canine diastemae of IVCM 32 are relatively shallow compared to those of E. (D.) simplicidens, but those of LACM 4338 and LACM 16815 are deeper than in E. (D.) simplicidens. Comparison of the cranial pre- and post-canine diastema length in IVCM 32 and LACM 4338 indicates the new spe- cies has a relatively long post-canine diastema, al- though immature LACM 16815 is closer to E. (D.) simplicidens and E. (D.) grevyi (see Table 2). The depth of the mandible below M, in IVCM 32 and LACM 3677 is somewhat deeper than other dolichohippine species (see Table 2). The lack of the ventral margin of the mandible in IVCM 1336 precludes comparisons. Basilar length cannot be determined in the new species, but LACM 4338 measures 554 mm from I1 to the temporal condyle. The distance from the temporal condyle to the foramen magnum in our E. (D.) simplicidens sample is about 12% of the basilar length. By using the 12% factor, we can estimate a basilar length of 621 mm for LACM 4338. Mandibular specimens can also assist in obtain- ing estimates of basilar length. The estimates of basilar length of the samples of E. (D.) simplicidens from Hagerman, E. (D.) grevyi, and E. sanmenien- sis are based on excellent photos from Teilhard de Chardin and Piveteau (1930) and data from Eisen- mann (1983). We observed the following differ- ences between mandibular and basilar lengths: the Hagerman sample of E. (D.) simplicidens has a dif- ference of 41 mm to 50 mm, mean 47 mm (n = 3); E. (D.) grevyi has an observed range of 63 mm to 75 mm, mean 68 mm (n = 5); and E. sanmeniensis has a difference of 65 mm. The mean of these differences between basilar and mandibular length is 60 mm. By using the 60 mm factor, the extrap- olated basilar lengths are 660 mm for IVCM 1336 and 633 mm for LACM 3677. Only two-thirds of the holotype mandible IVCM 32 is present. A crude length estimate of 556 mm was obtained from a restoration of the mandible using cranial and mandibular parts that were pre- served. Addition of the 60 mm factor provides an extrapolated 616 mm basilar length for IVCM 32. All samples of E. ( D .) enormis are estimated to range from 616 mm to 660 mm, with a mean of 633 mm in estimated basilar length. If we exclude IVCM 32, the estimate of the mean is 638 mm for basilar length. This compares with 414 mm to 544 mm, mean of 479 mm, in the mandible and 514 mm to 556 mm, mean of 537 mm, in the cranial Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 23 basilar length for E. (D.) simplicidens from Hager- man (Fig. 13 and Table 2). Perhaps the otherwise largest basilar length for Equus species is our es- timated 615 mm for E. livenzovensis or 616 mm for the Ingleside fauna horse referred to Equus complicatus (Lundelius, 1972). The cranial and mandibular lengths for Equus (D.) enormis are es- timated to be about 7 percent greater than for other dolichohippine species (see Fig. 13). Increased size and dolichocephaly is demonstrated by comparison of palatal length with transverse diameter at the juncture of M2 and M3 (see Fig. 15 and Table 2). A ratio of palatal width over length is estimated to be 0.21 to 0.24 (mean 0.23), compared with 0.26 to 0.27 in E. (D.) simplicidens and 0.26 to 0.28 in E. (D.) grevyi. Equus (D.) enormis (LACM 4338) measures 554 mm from I1 to the posterior lateral edge of the temporal condyle compared with the relatively shorter 455 mm to 490 mm (mean 476 mm) in E. (D.) simplicidens. Facial length is estimated to be 475 mm in LACM 4338 compared with a shorter 397 mm to 456 mm in E. (D.) simplicidens. The cranial symphysis is relatively broad and ro- bust, especially in IVCM 32 and moderately so in LACM 4338. The mandibular symphysis in IVCM 1336 seems relatively narrow compared to IVCM 32; this measurement cannot be determined for LACM 3677. The incisive foramen is round and located op- posite the center of the I3 in IVCM 32, LACM 4338, and LACM 16815. The grooves along the palatine process of the premaxillae vary in anterior extension from the center of the I3 in IVCM 32 to behind the I3 in LACM 4338 and LACM 16815. In IVCM 32, the post-maxillary palatal area is mod- erately shallow. The cheek teeth in IVCM 32 and LACM 16815 converge anteriorly. The anterior tips of the posterior narial borders are present, and their lateral borders are wider than in E. (D.) simplicidens. The palate and posterior nares of LACM 4338 appear to be narrower than in IVCM 32, although lateral compression has al- tered their form. The post-palatine foramen is po- sitioned opposite the center of M2 in IVCM 32 and opposite the anterior half of M3 in LACM 4338. The narial notch is relatively shallow in LACM 4338, and the posterior extension is above the metacone of P2 and P3 in E. (D.) simplicidens. The notch is deeper in LACM 4338 than in the living E. (D.) grevyi. The facial crest extends anteriorly as far as the Figure 8. a. Equus { Dolichohippus ) enormis, new spe- cies, paratype, LACM 4338, from fault block K, Vallecito Creek local fauna, approximately “zone” 54, late Blancan, cranium nearly complete, dorsal view. b. Equus ( Doli- chohippus) enormis, new species, paratype, LACM 4338, cranium, left lateral view. c. Equus ( Dolichohippus ) enor- mis, new species, paratype, LACM 4338, occlusal view. 24 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 25 I I 5cm Figure 9. Equus ( Dolichohippus ) enormis, new species, referred, LACM 16815, from Vallecito Creek local fauna, approximately “zone” 58, early Irvingtonian, partial im- mature cranium, occlusal view. center of P4 in IVCM 32 and LACM 4338 and as far as M1 in LACM 16815. The infraorbital fora- men is observable only on the left side of LACM 4338, occurring above ML The conformation of the middle braincase is discernable in LACM 4338, providing an estimated transverse width of 85 mm to 90 mm, compared to the observed range of 99 mm to 113 mm (mean 105 mm) for our sample of E. (D.) simplicidens. The right frontals and nasals in LACM 4338 are narrow throughout their entire length and slope steeply ventrally toward the ex- panding maxilla. Facial fossae appear to be absent from LACM 4338, which has the preorbital area preserved on the right side. The dorsal profile of the frontal and nasal bones (from the right side, lateral view, in LACM 4338) is very slightly concave. The temporal condyle (width in LACM 4338 67 mm, length 23.4 mm) is similar in shape to that of E. (D.) simplici- dens (width 67 mm, length 23 mm). Dimensions of individual cheek teeth, and es- pecially the length of tooth row, provide an esti- mate of overall size if comparisons are made with similar stages of wear. The moderately worn adult premolars and molars of IVCM 32 consistently ap- proach or exceed the upper size range of any one tooth of E. (D.) simplicidens (see Tables 3, 4). LACM 4338, of greater ontogenetic age and thus propor- tionately smaller tooth size because of greater wear, falls within the upper extremes of E. (D.) simplici- dens measurements. Adult E. (D.) enormis individ- ual tooth dimensions are closer in size to those of E. (D.) simplicidens than are the crania, mandibles, and some foot elements. The upper tooth series of £. (D.) enormis, IVCM 32, measures 217 mm com- pared to 181 mm to 207 mm, mean of 195 mm, in E. (D.) simplicidens. LACM 16815 is immature and, with less wear on the teeth, provides the longest upper tooth row measurement of 224 mm. The upper tooth row of LACM 4338, an elderly spec- imen, measures 204 mm and is within the upper size range of E. (D.) simplicidens. The four speci- mens of E. (D.) enormis together average 212 mm, but without the old adult LACM 4338, they average 221 mm and exceed E. (D.) simplicidens (see Table 2) in mean length. The lower cheek tooth row seems similar in length to the upper tooth rows of IVCM 32, IVCM 1336, and LACM 3677, measuring 229 mm, 231 mm, and 222 mm, respectively, with a mean of 227 mm. This contrasts with 173 mm to 210 mm, mean of 194 mm, in E. (D.) simplicidens. Without the old adult LACM 3677, the mean is 230 mm for the new species (see Table 3). The upper incisors are massive and reasonably well represented in LACM 4338, LACM 16815 (immature), and IVCM 32. With wear, proportions of the incisors change, becoming deeper mesiodis- tally, as in LACM 4338. However, regardless of state of wear, the incisors of IVCM 32 and LACM 4338, and the permanent I1 in LACM 16815, are larger than those of other species (see Table 2). The upper incisor infundibulae, or cups, are well de- veloped and broad in IVCM 32 and LACM 16815. Due to wear, LACM 4338 has only a faint sugges- tion of root canals, or nutrient canals, on its inci- sors. Upper canines in IVCM 32 and LACM 4338 are large; the little worn upper canine on the right side of IVCM 32 is broadly based, with a knife-like cutting edge and very slight lingual curvature. P‘ in IVCM 32 is represented by an alveolus 26 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids _Q CO Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 27 Figure 10. a. Equus ( Dolichohippus ) enormis, new species, referred, LACM 3677, from Borrego Badlands east of Borrego Springs probable, late Irvingtonian, mandible, occlusal view, b. Equus ( Dolichohippus ) enormis, new species, referred, LACM 3677, left mandible, lateral view. Table 4. Measurements of upper dentition. E. ( D .) enor. E. (D.) cf. simp. IVCM 32 LACM 4338 LACM 16815 IVCM 2673 LACM 17614 Length tooth row at alveolus, P2-M3 215-220 203-205 (212) 224 198,200 198,201 I1 AP at enamel crest 19.2, 20.5 14.2, 14.2 (19.0) 21.7,21.9 13.2, 13. 7e 18.5, 17.0 P TR at enamel crest 13.5, 15. 2e 15.1, 15.2 (14.0) 12.8, 13.1 11.7, 13.1 11.4 I2 AP 23. 5e, 23.5 15.7, 18.6 (21.0) 20.2, 21. 24 17.5,20.3 18.7,21.3 I2TR 13.2, 16. 3e 13.1, 13.1 (13.0) 19.7, 10.24 13.2 11.6, 13. 2e P AP 26.1,26.2 16.4, 17.7 (20.0) 19.2, 20.8 22.6, 24.9 P TR 11.6, 12.8 11.2, 11.9 (11.0) 8.4, 8.9 10.8, 10.8 C/AP at alveolus 18.2, 19.2 17.6, 17.7 (17) 12.6 17.0, 17.5 P1 present (pr)/ absent (ab) pr, pr ab, ab pr,pr ab, ab pr, pr P2 AP 49.7, 50.1 45e, 45e (47) 49e, 47e 45.5,46.2 43.2, 43.4 P2 TR 31.7, 33.8 27.2, 26.6 (30.4) 31.3,31.1 28.4, 29.2 29.5, 29.1 P; AP 37.7,38.3 35.3,35.0 (37) 37.3,38.0 35.0,35.0 33.0, 32.2 P3 TR 34.5,35.3 31.6,33.4 (34) 34.2, 34.5 32.4, 33.0 28.7, 28.7 P4 AP 35.8,36.8 30.5,31.1 (33) 33.6, 34.3 33.4, 33.6 31.8, 30.6 P4 TR 34.3,34.8 31.2, 32.6 (34) 31.7,36.5 32.1,32.4 27.8, 28.7 M1 AP 29.7,31.5 26.8,27.6 (29.6) 32.9, 33.2 26.7, 27.4 26.6, 26.8 M1 TR 31.7, 32.1 29.7, 29.7 (30.0) 28.4, 28.4 31.0,31.1 26.8,28.5 M2 AP 31.7,31.9 28.9, 28.2 (31) 32.4, 32.9 30.6, 32.0 27.2, 28.2 M2 TR 30.5,31.5 29.8,29.7 (29) 25.3, 24.7 29.0, 30.6 26.3, 28.5 M3 AP 26.2, 27.1 37.3,37.5 (32.5) 30.7,31.7 27.2, 30.6 M3 TR 27.0, 26.2 26.6, 27.2 (27.2) 24.8, 39.0 20.6,21.4 P3 AP protocone at enamel crest, greatest 14.5, 14.5 14.2, 14.2 (15.2) 16.6, 16.6 13.3, 13.1 11.7, 12.0 P3 TR protocone at enamel crest, minimum 5.0, 5.1 6.0, 6.3 4.5, 4.9 (16) 6.4, 7.0 5.7, 6.3 P4 AP protocone 15.1, 16.7 14.7 (16) 15.5 14.6, 15.1 14.2, 14.8 P4 TR protocone 5.0, 5.1 6.5, 7.0 (60) 6.8, 7.2 5.2, 5.7 M1 AP protocone 11.5, 11.6 13.2, 14.3 (13.3) 14.3, 14.7 11.8, 12.3 11.3, 11.8 28 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids Table 4. Continued. £. (£.) sp. A LACM 4335 E. (£.) sp. B IVCM 1816-1 E. cf. (£.) IVCM 3429 £. (D.) simp.' £. idaho. £. (£).) grev. £. (£.) przw. 183 222, 222 203 181-2072 194, 196 167-186 171-180 nl2(195)’ n4(180) n4(176) 16.3, 17.0 15.7-20.5 18.7 12.0-18.5 19.4-20.7 n8(17.7) n8(156) n4(202) 11.2-12.9 11.6 11.5-13.5 11.0-11.2 n5(11.9) n8(12.8) n4(ll.l) 18.6, 19.7 17.4 17.4-20.5 19.0, 18.7 15.2-19.7 19.6-22.3 n6(19.2) n8(17.8) n4(20.4) 10.8 15.6 11.2-12.7 12.6 10.2-13.5 10.7-11.2 n5(11.6) n8(11.9) n3(11.0) 18.1,20.5 16.3-24.0 20.1,20.8 16.2-22.0 19.6-22.3 n6(21.0) n8(19.4) n4(20.4) 9.8, 10.0 10.5-12.9 11.9, 11.7 9.3-12.5 10.7-11.2 n4(11.9) n7(11.3) n4(l 1.0) 6.8 7.1-19.0 14.6, 14.2 15.0-18.8 6.4-11.4 n6(12.3) n8(16.4) n4(8.6) pr, pr pr(6), ab( 1) pr? pr(6), ab(2) ab(4) 39.7,39.6 45.5 45 39.6-44.6 41.4, 42.0 37.2-42.4 39.2-40.5 n 12(42) n8(39.4) n4(40) 23.9, 26.2 29.0e 36.2 25.7-30.7 26.4, 25.5 22.4-29.6 24.5-25.7 nll(29) n8(26.3) n4(25) 30.0, 33.0 35.0, 34.0 35 32.5-38.5 32.0, 30.8 27.7-33.3 28.1-32.0 nl2(35) n8(31) n4(31) 28.0, 28.0 32.0, 32.0 35.6 29.5-32.1 28.6, 29.6 26.8-29.7 25.0-28 nl2(30.8) n8(28.5) n4(27) 30.0,31.0 36.0, 37.0 31.2 30.7-34.5 30.4, 30.7 27.0-30.8 27.5-29.3 nl2(32.5) n8(30) n4(29) 30.1,28.0 30.0, 37.0 35.2 28.4-31.4 30.1,30.8 26.7-29.4 24.8-26.5 nl2(29.5) n8(28.3) n4(25.7) 27.0, 26.0 29.0 27.8 26.2-34.6 25.7, 26.6 22.8-27.1 23.3-28.0 nl2(29) n8(25) n4(26) 29.0, 24.0 32.0 31.5 26.5-29.4 26.1,29.4 26.0-27.5 23.3-26.2 nl2(28.2) n8(27) n4(25) 28.0,29.0 32.0 30.3 27.6-32.8 26.7, 29.0 23.7-27.7 24.5-28.1 nl2(30) n8(26) n4(26) 28.0-29.0 34.0 31.8 27.0-30.8 27.5, 28.4 24.7-27.6 23.0-24.5 nl2(28) n8(26.3) n4(24) 28.0, 25.0 34.6 29.0-32.4 30.6,33.4 24.6-30.3 24.4-27.0 nl0(31) n8(28.3) n4(26) 22.0, 24.0 29.2 21.4-29.0 24.8,25.0 20.7-21.5 19.4-21.4 nl0(24.2) n8(23) n4(20.5) 9.2, 10.1 17.2, 15.7 16.4 7.7-11.7 10.0, 11.3 10.4-14.2 13.5-14.4 n 12(10) n8(13) n4(14) 5.8, 5.0 5.5, 7.0 5A-7.7 5.6 5. 3-6. 6 4.5-5. 6 nl2(6.6) n8(5.8) n4(5.1) 11.9, 11.0 18.8,20.5 17.5 7.8-13.8 11.0, 11.2 11.8-14.7 13.5-13.8 nl2(ll) n8(13.0) n4(13.6) 5. 5,5.0 6.9, 6.5 5.0-6. 5 5.7, 6.2 4. 8-5. 7 4.5-5. 8 nl2(6.0) n8(5.3) n4(5.5) 11.5, 10.7 15.2 14.2 8.9-13.5 8.7, 10.0 4. 7-6. 5 11.4-13.4 nl2(ll) n8(5.6) n4(12.0) Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 29 Table 4. Continued. E. ( D .) enor. E. ( D .) cf. simp. IVCM 32 LACM 4338 LACM 16815 IVCM 2673 LACM 17614 M1 TR protocone 5.0, 5.7 6.8, 7.3 (5-8) 4.7, 5.1 6.7, 6.9 6.1, 6.2 M2 AP protocone 13.9, 14.4 14.2, 14.4 (14.8) 15.0, 15.8 14.7, 14.7 11.7, 12.5 M2 TR protocone 4.3, 4.5 6.0, 6.1 (5.2) 4.8, 4.8 6.4, 6.5 5.5, 5.6 P' protocone shape 3,3 3, 3 3,3 2,2 3, 3 P4 protocone shape 2,2 3A, 3A 2A, 2A 3A, 3A M' protocone shape 2,2 2A, 2A 3,3 2,2 2,2 M2 protocone shape 3A, 3A 2A, 2A 3,3 2,2 3,3 P’ AP protocone/AP tooth 0.38,0.38 0.40,0.41 (0.41) 0.44, 0.45 0.38 0.36 P4 AP protocone/AP tooth 0.41,0.47 0.48,0.50 (0.47) 0.45, 0.44 0.46 M1 AP protocone/AP tooth 0.37,0.39 0.48,0.53 (0.44) 0.46, 0.43 0.42, 0.44 M2 AP protocone/AP tooth 0.44, 0.45 0.49,0.51 (0.47) 0.46,0.48 0.46,0.41 1 Includes data from Gazin (1936) and specimens from IVCM, LACM, and UCMP collections. 2 207 mm in E. livenzovensis. (Not listed in Table 4.) 3 n = number of tooth rows, not individuals. 4 Deciduous teeth. 5 All AP and TR measurements of tooth enamel are at crest or biting surface. b See Figure 2H for shape category. containing the root. P1 was probably present in LACM 16815. P1 is well developed in all five spec- imens of E. (D.) simplicidens compared. P2 has a pseudoparastyle, anterior to the para- style, and appears relatively stronger in IVCM 32 and LACM 16815 than in E. (D.) simplicidens. LACM 4338 lacks the pseudoparastyle. The para- style, which is well developed in E. (D.) simplici- dens, is strong in IVCM 32 but narrow in LACM 4338. The mesostyle is broad and slightly grooved in IVCM 32, moderately broad with slight groove in LACM 4338, and generally broad in E. (D.) sim- plicidens. The metastyle is relatively broad in IVCM 32 but absent in LACM 4338. The protoloph con- nects with the parastyle in IVCM 32 but is not discernable in LACM 4338 or LACM 16815. The following remarks concerning dental char- acters refer to P3 through M3, unless otherwise in- dicated. Where the state of wear in LACM 16815 prevents comparisons, no comment will be made. The pli-protoloph is small and progressively re- duced in P3 through M3 of IVCM 32 and LACM 4338 and in M1 and M2 of LACM 16815. Proto- cones are generally elongate and with narrow lin- gual grooves in E. (D.) enormis (Fig. 2H and Table 2) in comparison to E. (D.) simplicidens. The pro- toconal index (protoconal length compared to an- 30 ■ Contributions in Science, Number 440 teroposterior length) is 0.37 to 0.53 in P2 through M2 of E. (D.) enormis. Equus (D.) simplicidens has mean protoconal indices of 0.28 to 0.44; E. ( D .) grevyi ranges from about 0.43 to 0.52 and is similar to E. (D.) enormis (see Table 2). The generally elon- gate protocone seems to be correlated with the relative increase in length of the “toe” (or posterior extension) of the protocone. P3 and P4 appear to have shorter pre-protoconal grooves in IVCM 32 and LACM 4338 when compared to E. (D.) sim- plicidens. The parastyles are broad and slightly grooved in P3 through P4 but narrow and ungrooved in M!, M2, and M3 of IVCM 32; teeth of LACM 4338 are similar except there is no apparent groove in P3 and P4. M1 and M2 parastyles in LACM 16815 are nar- row without grooves. The mesostyles are similar to the parastyles ex- cept they are narrower in IVCM 32, but wider in LACM 4338, when compared to E. (D.) simplici- dens. Complexity of the enamel pattern, the number of plications in the pre- and post-fossettes including the plicaballin, is determined by actual counts. The pre- and post-fossettes, including the pli-protoloph, tend to be complex in P3 through M2, especially as noted in IVCM 32, with a range of 9 to 13 plica- Downs and Miller: Late Cenozoic Equids Table 4. Continued. E. (£.) sp. A LACM 4335 E. ( E .) sp. B IVCM 1816-1 E. cf. (£.) IVCM 3429 £. (£).) simp.' E. idaho. £. (D.) gre\. £. (£.) przw. 5.6, 5.7 6.3 4. 8-6.4 5.3, 67.1 4.7-6. 5 3 .7-5.3 nl2(5.3) n8(5.6) n4(4.7) 13.0, 13.8 19.4 15.6 11.0-15.7 11.1, 11.5 12.7-14.6 13.2-14.7 nl2(13) o 00 c n4(14) 5.2, 5.6 6.6 4.7-6. 7 5.2, 5.3 4. 1-5. 5 4. 1-5.0 nl2(5.8) n8(5.0) n4(4.5) 3,3 3-4 3 2n8,3n4 2,3 3n8 3n2, 4n2 3,3 4,4 4 2n8,3n4 2,2 3n8 4n2,5n2 5,5 3-4 2 2n8,2An2,3An2 3,3 2n2,3n2,3An2, 4n2 2n2,5n2 2,2 4-5 3 2An8,3n4,3An2 3A, 3A 3n4,3n2,5n2 5n4 0.31,0.31 0.49, 0.46 0.47 0.24-0.33 0.35,0.32 0.38-0.46 0.42-0.51 nl2(0.28) n8(0.42) n4(0.47) 0.41,0.35 0.52, 0.55 0.56 0.23-0.43 0.36, 0.36 0.41-0.48 0.34-0.50 nl2(0.34) n8(0.45) n4(0.45) 0.41,0.42 0.52 0.51 0.34-0.44 0.33,0.39 0.46-0.56 0.45-0.50 nl2(0.38) n8(0.52) n4(0.48) 0.47, 0.49 0.60 0.51 0.32-0.48 0.40, 0.41 0.46-0.57 0.49-0.52 nl2(41) n8(0.52) n4(0.51) tions. Equus (D.) enormis thus tends to be more complex than E. (D.) simplicidens (5 to 11 plica- tions). The plicaballin is well developed in IVCM 32 and LACM 4338, and one to two pli-hypostylids occur in the premolars and molars. The pre-hypoconal groove is present on P2 and P3 of IVCM 32, and incipient on P2-M3 of LACM 4338 and on the M2 of LACM 16815. The post- hypoconal groove is deep and open on P2-M2 of IVCM 32 and LACM 4338 and on M12 of LACM 16815. The M3 of IVCM 32 is not worn sufficiently to show development of a groove; the right M3 in LACM 4338 has the groove in the form of a fos- sette. All the upper and lower teeth have an abundance of cement, both labially and lingually, and are very hypsodont. The crown height of M2 in IVCM 32 is at least 85 mm; right and left P2, 86 mm and 89 mm; right and left P4, 96 mm and 92 mm; and left M2, 99 mm in LACM 16815. The exposed crowns show slight degree of curvature, especially in LACM 16815. The height of crown in E. (D.) simplicidens is 59 mm in P2 of LACM 1520 and 62 mm and 82 mm in P3 of IVCM #H1. The preserved parts of the mandible of E. (D.) enormis mirror the relatively elongate proportions of the cranial rostrum (see Tables 2, 4). Equus (D.) enormis, IVCM 32, IVCM 1336, and LACM 3677, have a mandibular rostral index (length I,-P2/width at I3) of 0.39 to 0.47, mean of 0.42, compared to equivalent indices of 0.43 to 0.58, mean of 0.50, in E. (D.) simplicidens. The mandibular rostral in- dex of IVCM 32 is 0.47, which confirms our esti- mated cranial rostral index of 0.44 for E. (D.) enor- mis. Behind the canines, the symphysis of IVCM 32 is narrow and deeply furrowed dorsally. The preangular notch is broadly concave in IVCM 32 and LACM 3677 but indeterminate in IVCM 1336. The mental foramen is slightly anterior to the P2 in IVCM 32 but slightly posterior to the canine in IVCM 1336. The angle between the slope of the anterior border of the ascending ramus and the horizontal axis of the tooth row is estimated as 130° in IVCM 1336 and 133° in LACM 3677. This compares with 120° to 140° in E. (D.) simplicidens, 135° to 145° in E. (D.) grevyi, and 115° in E. (E.) przewalskii. The lower incisors in IVCM 32, and especially in the I3, are massive and are well preserved; they have very elongate, clearly formed cups on I, and I2. I3 has no cup and possesses a deep to broad recessed “V” with two pointed cusp-like cingulae at the base of the “V.” All the incisors of IVCM 1336 are well worn; due to advanced wear, there is no cup on I15 a minute trace of a cup is lingually placed on L, and a remnant of an open “V” is clearly present on I3 (Fig. 11a), with a small lingual cusp-like cingulum at the mid-point. In LACM 3677, the right I2 is well worn, about 19 mm in height, and lacks a cup, although a nutrient canal may be present. I3 of LACM 3677 is worn, but with a rem- nant of a slightly developed recessed “V” (Fig. 10a). The third incisors of E. (D.) enormis greatly resem- ble those of E. (D.) simplicidens; however, Gazin (1936:302) stated, “I3 is broadly open on the lingual side and in most cases does not exhibit a lingual stylar cup, although in a few instances a small cup is present . . . .” We find no cup on the I3 in our Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids M 31 J' / 1 CO -Q 32 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids Figure 11. a. Equus ( Dolichohippus ) enormis, new species, referred, IVCM 1336, from lower Coyote Canyon badlands, north of Borrego Springs, probable late Irvingtonian, right mandible, occlusal view. b. Equus ( Dolichohippus ) enormis, new species, referred, IVCM 1336, right mandible, lateral view, c, d. Equus ( Dolichohippus ) enormis, new species, composite of (c) LACM 4338 and (d) LACM 3677. CO -Q Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 33 Figure 12. Equus ( Dolichohippus ) enormis, new species, composite of (a) LACM 4338 cranium and (b) LACM 3677 mandible (not associated), left lateral view; from Figures 8b and 10b. E. complicatus CR MR Cp Cp E. (E) przewalski CR MR Pr Pr Pr Pr Pr E. (D) grevyi CR MR GG G G Q G G G G G E. mosbachensis CR MR Mo Mo E. stenonis s.l. CR MR St St E. koobiforensis CR E. sanmeniensis CR MR E. livenzovensis CR MR E. (D) simplicidens CR MR H H HH H H H H H H H H HH HH H H Anza-Borrego Desert E. (Dj enormia E. ( D ) cf simplicidens E. (Dj enormis CR MR _ ®_. 1 e (j> E 4' i " — i t r 600 620 640 660 1 T i r 400 420 440 460 480 500 520 540 560 580 Comparison of cranial and mandibular lengths Figure 13. Cranial and mandibular lengths compared (for abbreviations and symbols, see Appendix A). (1) E. com- plicatus, data from Lundelius (1972). sample of E. (D.) simplicidens, but the above no- tation by Gazin emphasizes the need for caution in noting the presence of the recessed “V.” The vari- ability in this character is documented by Eisen- mann (1979a) and Azzaroli (1979). The lower left canine of IVCM 32 is broad, or rounded, at the basal outline and flattened dorsally with wear and possibly due, in part, to restoration. The canine in IVCM 1336 is unworn, somewhat pointed, and narrower at the base than in IVCM 32. The lower right canine is broken distally in LACM 3677 but preserved in its broad rounded base. Cement covers the unworn surfaces of the incisors and canines in the new species. The pre- and post-canine mandibular diastemae vary in absolute size within the sample of E. (D.) enormis but seem to be similar to E. (D.) simplici- dens in proportions, with the post-canine diastemia being much longer than the pre-canine diastema. The right post-canine diastema is fractured in IVCM 32 and is somewhat shorter than in IVCM 1336 and LACM 3677. However, the fractured margins of the mental foramen openings and sediment-filled inner canal of the foramen are well seen in IVCM 32 and do not distort rostral proportions. A portion of the lingual flange, the juncture of the right and left symphysis, occurs at the fracture. All three spec- imens have appreciably longer pre- and post-canine 34 ■ Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids Anterior posterior diameter, rostrum, I- to P- Transverse diameter, rostrum at I- Figure 14. Rostrum, anterior posterior length for I1 to P2 compared with transverse width at I3 (for abbreviations and symbols, see Appendix A). Contributions in Science, Number 440 Downs and Miller: Late Cenozoic Equids ■ 35 360 — CD ■4— * 03 03 Ql O w ^ ^ e a> 3 H pd i p ON VO OO r. i ■p r PO OO OO •p r- vo r 4 Oi on c OO I OO Tt" VO VO >D VO I PD VO p _ D4 4 o , _ VO VO VO 1 *D J lD >D ' — - •vo -vo 7" c c >D id tp OO 'vO rsj PD ov *D „ PD P 4 ^ VO o I vo iO - • VO vo c *D VO vo ov *D OO OO OO o OO OO o >D I ON PD o OO -3L VO o ^ 4 vo | ON ^ o OO PO D 'd r d i vo 4 P 4-i rt cu W) 4 — • 4-— 03 03 D "O T3 C O -a 24), samples sizes (in parentheses), and minimum and maximum values for external measures for adult specimens of the two forms of common dolphins from California. Snout (= beak) measures are taken from the tip of the upper jaw. Males Females Short-beaked Long-beaked Short-beaked Long-beaked Total length 189.5 ± 7.37 (28) 172-201 219.1 (15) 202-235 180.1 ± 6.5 (37) 164-193 207.7 (10) 193-224 Snout to anus 137.7 (20) 127-148 158.3 (12) 148-174 130.3 ± 4.3 (27) 120-138 145.0 (6) 139-151 Snout to genital slit 121.0 (23) 109-128 138.8 (12) 129-151 123.7 ± 4.4 (26) 116-133 138.0 (6) 132-145 Snout to umbilicus (18) 89.3 83-95 103.1 (10) 100-109 (24) 86.3 80-90 (4) 95.5 94-98 Snout to dorsal fin tip (18) 114.4 105-126 (12) 125.5 119-137 (27) 106.9 ± 4.0 98-115 (7) 122.9 113-133 Snout to anterior dorsal fin (7) r-H o-n . i o oo oo (12) 95.3 88-100 (18) 82.6 76-86 (7) 94.3 88-101 Snout to flipper (17) 44.6 41-49 (10) 49.3 47-52 (28) 41.9 ± 2.1 38-46 (6) 47.5 45-50 Snout to ear (15) 35.9 32.5-40.0 (9) 38.6 37.0-41. (22) 34.2 31.5-37.2 (7) 38.9 36.5-43.0 Snout to eye (20) 30.8 27.3-34.3 (12) 34.4 31.0-38.0 (33) 29.3 ± 1.6 26.5-32.7 (7) 33.6 31.0-37.0 Snout to gape (24) 26.1 21.7-28.0 (12) 29.1 27.0-32.0 (33) 24.8 ± 1.3 22.0-27.5 (7) 28.4 27.0-31.0 Snout to blowhole (19) 31.4 27.7-34.2 (13) 34.8 31.1-40.0 (34) 30.4 ± 1.8 26.4-33.8 (8) 34.7 32.5-38.4 Snout to melon apex (25) 12.3 ± 0.89 10.5-14.0 (14) 15.7 14.0-17.5 (26) 11.6 ± 0.7 10.5-13.0 (8) 15.4 13.3-17.0 Eye to ear (20) 5.3 3. 3-8.0 (9) 5.6 5.0-6. 5 (21) 5.2 4.6-62 (6) 5.6 5.0-6. 1 Eye to gape (20) 5.8 4.6-6. 6 (12) 6.4 5. 5-7.0 (25) 5.8 ± 0.4 4. 7-7.0 (7) 5.8 4. 5-6. 9 Right eye to blowhole (2) 18.3 18.0-18.5 (7) 17.4 16.0-18.5 (10) 17.1 16.3-18.5 (5) 16.3 16.0-1 6.5 Left eye to blowhole (15) 14.3 10.7-18.0 (9) 16.2 15.0-17.5 (19) 14.6 13.2-16.0 (7) 15.0 14.0-16.0 Blowhole length (11) 1.2 0.9-1. 5 (12) 1.3 1. 0-2.0 (21) 1.1 0. 8-1.5 (8) 1.1 0.8-1. 3 Blowhole width (13) 2.4 1. 6-3.0 (12) 2.5 2.0-3.0 (21) 2.1 1. 7-2.9 (8) 2.4 1.9-2. 8 Head diameter at eyes (12) 20.1 19.0-21.1 (11) 20.1 19.0-22.5 (26) 19.9 ± 1.5 17.2-22.8 (7) 18.9 18.0-19.5 Rostral width at melon apex (11) 7.4 6. 1-8.4 (13) 7.2 6. 2-8.0 (19) 7.9 6.0-11.0 (8) 6.5 6.0-7.4 Projection of lower jaw (11) 0.4 0.2-0. 5 (11) 0.7 0.3-1. 5 (16) 0.4 0. 2-1.0 (4) 0.6 0. 5-1.0 Flipper length, anterior (22) 30.0 25.4-34.3 (12) 31.1 28.9-33.0 (27) 27.1 ± 2.1 24.0-31.0 (8) 29.7 28.4-32.0 Flipper length, posterior (22) 21.2 17.2-24.6 (12) 22.7 18.9-25.0 (27) 19.5 ± 1.8 16.2-23.0 (8) 20.4 18.0-23.0 Flipper width (21) 9.9 8.5-11.3 (12) 10.2 8.8-12.2 (25) 8.8 ± 0.8 7.1-11.0 (8) 9.7 9.0-10.7 Length of genital slit (11) 9.6 5.9-12.3 (ID 9.3 6.6-12.5 (11) 12.1 9.5-16.6 (8) 11.5 8.0-13.0 Fluke width (22) 42.9 29.9-52.3 (ID 47.9 40.0-58.0 (25) 38.5 ± 3.9 29.0-46.0 (7) 42.4 38.0-49.0 Fluke depth at lobe (1) 12.5 12.5 (7) 14.7 13.3-16.5 (4) 12.3 11.5-13.0 (4) 13.1 12.0-14.0 12 ■ Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus Delphinus Table 2. Continued. Males Females Short-beaked Long-beaked Short-beaked Long-beaked Fluke depth at notch 12.0 13.0 11.4 11.7 (21) 9.9-14.0 (10) 12.0-14.5 (24) 9.6-13.0 (7) 10.5-13.0 Fluke notch depth 2.4 2.4 2.3 ± 0.6 2.5 (18) 1.8-3. 5 (11) 1. 5-3.0 (25) 1. 0-3.0 (7) 1. 5-3.0 Dorsal fin height 20.1 19.9 16.6 ± 1.4 17.4 (22) 17.0-26.1 (12) 17.0-24.5 (25) 14.0-19.0 (8) 13.0-21.1 Dorsal fin base length 30.1 32.7 27.1 33.0 (8) 24.9-33.0 (10) 30.0-36.0 (10) 19.0-32.0 (8) 27.0-41.1 Girth at eye 70.5 68.6 67.8 67.7 (2) 68-73 (8) 66-73 (6) 66-71 (6) 65-70 Girth at axilla 97.5 100.1 91.7 ± 6.1 95.4 (23) 88-111 (10) 87-110 (26) 73-101 (8) 83-103 Girth, maximum 101.4 112.4 99.2 106.6 (7) 96-108 (10) 93-133 (15) 86-111 (7) 87-116 Girth at anus 63.1 66.9 54.5 ± 4.5 55.9 (21) 49-73 (11) 54-80 (26) 47-66 (7) 50-65 Girth midway anus-fluke notch 32.5 34.8 29.6 31.0 (2) 32-33 (8) 31-38 (5) 28-32 (4) 29-34 Caudal height 15 15.6 13.6 15.8 (2) 15.0 (8) 14-17 (5) 13-14 (4) 14-19 Caudal thickness (1) 3.5 (1) 4.9 3.0 4.4 3.5 4.9 (4) 3-5 (5) 3-6 ■ Long-beaked a Short-beaked Figure 11. Plot of length and mass data for the two forms of common dolphins off California. Note that for the same body length the short-beaked form tends to be heavier. Open squares represent short-beaked specimens, and filled squares long-beaked specimens. Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus DelphinusM 13 Table 3. Means, standard deviations (for sample sizes > 20), samples sizes (in parentheses), and minimum and maximum values for selected external measures as per- centage of total length for adult specimens of the two forms of common dolphins from California. Short-beaked Long-beaked Flipper length anterior Males 15.8 ± 1.1 14.1 (24) 13.8-17.4 (12) 13.5-15.1 Females 15.0 ± 1.2 14.4 (26) 13.3-17.1 (8) 13.9-15.5 Dorsal fin height Males 10.2 ± 2.3 9.0 (24) 8.7-13.6 (12) 7.3-10.4 Females 9.3 ± 0.8 8.4 (24) 7.6-10.6 (8) 6.0-9. 5 Fluke width Males 22.9 ± 2.0 21.7 (22) 17.8-26.5 (11) 18.4-24.7 Females 21.2 ± 2.0 20.5 (25) 17.6-24.7 (7) 19.0-23.2 Snout to melon apex Males 6.4 ± 0.4 7.2 (27) 5. 5-7.1 (14) 6.2-8.0 Females 6.4 ± 0.4 7.6 (25) 5. 6-7.3 (7) 7.3-8. 1 that there are two forms of common dolphins in the eastern North Pacific separated completely by rostral length. To take into account size differences, Banks and Brownell (1969) used the ratio of rostral length to zygomatic width to distinguish the two forms. They found that specimens with a ratio above 1.55 could be assigned to D. bairdii and below 1.53 to D. delphis. In their response to Banks and Brownell’s (1969) study, van Bree and Purves (1972) noted that they found some specimens with intermediate ratios of rostral length to zygomatic width when sampling common dolphins from other ocean basins. How- ever, neither of these two studies separated speci- mens by sex, nor did they include only mature specimens. This confounded sexual dimorphism, ontogenetic variation, and geographical variation with potential subspecific or specific level differ- ences. Evans (1982) re-plotted rostral length on zy- gomatic width, incorporating only sexually mature animals, and obtained more discrete clusters than did Banks and Brownell (1969). We plotted rostrum length on zygomatic width (Figs. 5, 6) using only mature specimens stratified by sex and found that the differences are not just size related; they represent a true shape difference in the skulls of these two forms. We found that adult specimens from California waters were sep- arated by a gap between 1.47 and 1.52 for this ratio (Table 8). Skull measurements are available for specimens of Delphinus sp. from South Africa (Ross, 1984), the type locality of D. capensis. We re-analyzed the ratio of rostrum length to zygomatic width, including only specimens considered to be mature based on either age (over 8 growth layer groups; Ross, 1984, table 90) or known reproductive status (Ross op. cit., tables 92, 93). Adult specimens from this region have an average ratio of 1.68 (n = 12) with a range of values from 1.59 to 1.76. These numbers do not include one specimen lacking ma- turity data with an extremely long beak that has a ratio of 1.87. Based on this small sample, common dolphins from the waters off South Africa have beaks that are modally longer than in the long- beaked form off southern California, with consid- erable overlap in the range of values. These values are even more distinct from those of the short- beaked form off California. The ratio of rostrum length to zygomatic width for 10 specimens of D. tropicalis, excluding two immature skulls, ranges from 1.89 to 2.22 (Pilleri and Gihr, 1973-1974; van Bree and Gallagher, 1978; Mohan, 1985). One specimen from off the Arabian Peninsula with a ratio of 1.72 was considered to represent a specimen of D. delphis by van Bree and Gallagher (1978). This value is well out of the range of the short-beaked form from other regions and within the range of values for the long-beaked form from either California or South Africa. Thus, we believe this specimen does not belong to the short- beaked D. delphis form, but with the long-beaked D. capensis form (see discussion under Taxonomy). In this same paper, van Bree and Gallagher (1978) presented data from a long-beaked specimen from Taiwan with a rostral ratio of 1.75. Kasuya (1973) noted another specimen from Taiwan with a bair- dii-type rostral length. Based on the data in Ross (1984), the long-beaked common dolphins from South Africa have an av- erage upper tooth count of 52 (n = 17) with a range of 47-60 and an average lower tooth count of 51 (n = 13) with a range of 47-57. These values are almost identical to the tooth counts for the long- beaked form from California (Table 7). Tooth counts for 11 D. tropicalis were 57-69/5 4-64 (van Bree and Gallagher, 1978; Mohan, 1985). These values overlap somewhat with the tooth counts for the long-beaked form from South Africa and Cal- ifornia. These counts do not include the specimen from the Indian Ocean with a rostral ratio of 1.72. This specimen has upper tooth counts of 57 and 56, within the region of overlap between D. tropi- calis and the long-beaked form. Perrin et al. (1987) found that vertebral counts were one of two diagnostic characters useful for distinguishing between two species of spotted dol- phins in the Atlantic. In the California sample, we found strong modal differences in several vertebral meristics, but no separation. Thus, some specimens cannot be assigned to a particular form or species based on univariate postcranial characters only. Off South Africa, the range of vertebral formulas in the 14 ■ Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus Delphinus Table 4. Means, standard deviations (for sample sizes > 30), samples sizes (in parentheses), and minimum and maximum values for skull measurements for adult specimens of the two forms of common dolphins from the coast of southern California. Males Females Short-beaked Long-beaked Short-beaked Long-beaked Condylobasal length 421.5 ± 13.04 473.6 406.3 ± 14.06 465.5 (48) 392-445 (19) 446-498 (49) 382-442 (12) 445-486 Rostral length 254.4 ± 10.97 302.0 244.0 ± 10.99 296.2 (48) 227-275 (19) 286-321 (49) 218-264 (12) 281-314 Rostral width At base 86.9 ± 4.03 85.9 84.9 ± 4.10 82.7 (50) 78-98 (19) 77-93 (51) 73-93 (13) 78-88 At 14 length 57.6 ± 2.87 57.4 55.7 ± 2.59 55.3 (48) 52-63 (19) 53-61 (49) 49-63 (12) 52-58 At Vi length 51.3 ± 2.55 50.9 48.7 ± 2.06 48.3 (48) 45-57 (19) 46-55 (49) 43-55 (12) 46-51 At % length 38.1 ± 2.65 38.0 35.4 ± 2.12 35.1 (48) 33-44 (19) 33-47 (49) 32-41 (12) 30-39 Premaxillae width at Vi length 23.6 ± 1.67 23.9 22.7 ± 1.49 23.5 (48) 21-27 (19) 19-28 (49) 20-25 (12) 21-26 Tip of rostrum to external nares 301.9 ± 11.91 359.5 290.3 ± 12.18 343.2 (48) 272-321 (19) 331-375 (48) 263-314 (12) 326-364 Tip of rostrum to internal nares 297.1 ± 11.53 344.5 285.3 ± 12.64 339.5 (47) 270-318 (19) 323-370 (49) 258-316 (12) 322-355 Preorbital width 167.3 ± 4.76 167.4 161.6 ± 5.06 160.3 (50) 158-177 (19) 156-181 (51) 150-175 (13) 150-169 Postorbital width 187.6 ± 4.81 190.4 182.5 ± 4.81 183.1 (49) 176-197 (19) 183-205 (51) 174-192 (13) 173-193 Zygomatic width 184.9 ± 5.27 189.1 179.6 ± 4.91 180.8 (49) 173-195 (19) 181-204 (51) 170-190 (13) 173-191 Width of external nares 45.1 ± 2.26 45.6 44.3 ± 2.35 45.1 (50) 39-50 (19) 41-49 (50) 39-50 (13) 43-50 Greatest width of premaxillae 71.5 ± 2.99 71.1 71.8 ± 3.19 70.8 (50) 66-78 (19) 66-77 (50) 63-79 (13) 68-74 Parietal width 148.9 ± 2.99 153.8 145.4 ± 5.26 149.8 (50) 139-162 (19) 145-162 (51) 132-156 (13) 138-158 Height of braincase 108.0 ± 4.35 112.3 103.5 ± 15.89 108.8 (50) 98-116 (19) 106-126 (50) 94-114 (13) 100-118 Length of braincase 116.1 ± 4.73 119.3 112.0 ± 4.32 116.8 (49) 106-124 (18) 114-129 (49) 104-124 (13) 111-124 Length of temporal fossa 67.6 ± 4.19 73.6 65.8 ± 4.15 69.4 (50) 59-84 (19) 65-81 (51) 54-73 (13) 64-76 Height of temporal fossa 51.0 ± 4.11 57.6 48.9 ± 3.78 55.5 (50) 41-60 (19) 49-63 (51) 42-58 (13) 48-68 Length of orbit 49.3 ± 1.99 49.2 48.7 ± 2.22 48.2 (50) 45-54 (19) 44-55 (51) 45-53 (13) 44-53 Length antorbital process 44.5 ± 2.62 44.9 42.4 ± 2.52 43.8 (50) 39-51 (19) 41-50 (51) 36-48 (13) 38-47 Width of internal nares 54.2 ± 2.15 55.8 52.1 ± 2.16 54.1 (49) 48-59 (19) 53-60 (51) 47-57 (13) 50-60 Length of upper tooth row 214.3 ± 9.90 258.8 205.0 ± 10.09 253.6 (47) 188-232 (19) 241-275 (49) 181-226 (12) 232-271 Length of lower tooth row 208.5 ± 10.77 246.4 200.5 ± 9.84 238.3 (49) 184-232 (20) 228-268 (50) 181-218 (13) 215-255 Length of ramus 354.8 ± 13.34 405.0 343.0 ± 12.69 394.3 (49) 323-379 (20) 384-433 (50) 321-372 (13) 371-419 Height of ramus 64.1 ± 2.52 70.8 62.2 ± 2.89 67.8 (50) 57-70 (20) 66-77 (50) 57-69 (13) 65-71 Tooth width 2.81 ± 0.233 3.71 2.83 ± 0.238 3.57 (41) 2. 5-3.4 (15) 3. 3-4.0 (33) 2.3-3.3 (11) 3.2-4.0 Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus DelphinusM 15 Table 5. Medians, sample size (in parentheses), and min- imum and maximum values for tooth row counts for the two forms of common dolphins from the coast of south- ern California. Short-beaked Long-beaked Upper teeth, left 49 53 (136) 42-54 (45) 48-59 Upper teeth, right 49 53 (138) 42-54 (45) 47-58 Lower teeth, left 47 51 (143) 41-53 (46) 47-55 Lower teeth, right 47 51 (172) 41-53 (47) 47-55 capensis) in Japanese waters based on the length of the rostrum and tooth count. For D. delphis, he found the range of total vertebrae was 74-78 (n = 6) and the ranges of the tooth counts were 44-49/ 44-50 (n = 8). The pigmentation pattern in Ogawa’s (1936) photograph of D. delphis agrees with our definition of the short-beaked form. The tooth count ranges listed by Ogawa for D. capensis are 54-55/51-55 (n = 3). Based on a small sample, Kasuya (1973) found that the sigmoid process of the tympano-periotic complex was larger in the short-beaked form (D. delphis, n = 6) than in the long-beaked form (D. bairdii, n = 2). long-beaked form is C 7, T 14-17, L 18-23, Ca 30-34, for a total of 72-76 (n = 11; Ross, 1984). The total counts are below the range of counts for the long-beaked form off California and even below the average vertebral counts for the short-beaked form off California (Table 8). This is due primarily to fewer caudal vertebrae. In summary, common dolphins from off South Africa are very similar to the long-beaked form in coloration, rostral length, and tooth counts but differ in average total vertebral count. Casinos (1984) examined specimens of Delphinus from the Atlantic Coast of South America. Based on his data, only 6 of the 10 specimens are assign- able to form. These all represent the long-beaked form based on rostral ratios (1.55-1.77, n = 5) or tooth counts (60-59/53-55, n = 1). The locality for these specimens ranged from Venezuela to the La Plata region of Argentina. Ogawa (1936) recognized both the short-beaked form (D. delphis ) and the long-beaked form (D. ECOLOGY To occur sympatrically, these two forms must be exploiting the environment in at least subtly dif- ferent ways. Banks and Brownell (1969) suggested that until the end of the 1800s the long-beaked form was more common, and that in this century the short-beaked form was more prevalent in southern California waters. These authors also sug- gested that the long-beaked form is more abundant off California during periods of relatively warm wa- ter. We analyzed stranding data from 1970 to 1990 for common dolphins stranded along southern Cal- ifornia shores (Fig. 15) to evaluate any potential temporal patterns. A gradual shift in relative abun- dance of the two forms is evident. From 1970 to 1982 inclusive, 84.4% (n = 109) of the common dolphins strandings were of the short-beaked form. During the years 1983 to 1988, 88.2% (n = 34) of the stranded common dolphins were of the long- beaked form. This shift roughly coincides with the onset of the warm water El Nino event of 1982/ 1983, which also apparently shifted the distribution « c 24), samples sizes (in parentheses), and minimum and maximum values for postcranial meristics for the two forms of common dolphins from southern California. Short-beaked Long-beaked Vertebral count 76 ± 1.48 78 ± 0.64 (80) 74-80 (25) 77-80 No. of lumbar vertebrae 20 ± 1.12 20 ± 0.96 (76) 18-24 (25) 19-22 No. of caudal vertebrae 34 ± 1.30 35 ± 1.04 (75) 30-36 (25) 33-37 First vertebra with 54 ± 1.70 55 ± 1.46 vertical foramen (76) 49-59 (26) 53-57 Last vertebra with 58 ± 1.26 59 ± 1.22 transverse process (75) 54-61 (26) 55-60 Last vertebra with 63 ± 1.45 65 ± 1.03 neural arch (75) 60-68 (25) 63-67 First vertebra with 43 ± 1.19 44 ± 0.83 chevron (75) 41-46 (25) 42-45 No. of chevrons 25 ± 2.32 27 ± 1.78 (71) 18-29 (25) 23-30 No. of vertebral ribs 15 ± 0.63 15 ± 0.56 (78) 13-16 (25) 14-16 No. of sternal ribs 8 ± 0.76 8 ± 0.72 (77) 6-10 (26) 7-9 Number of carpals 10 10 and metacarpals (33) 10 (15) 10 Phalanges Digit 1 2 2 (33) 1-2 (10) 1-3 Digit 2 8 ± 0.62 8 (32) 7-9 (9) 8-9 Digit 3 6 ± 0.45 6 (33) 5-7 (9) 6-7 Digit 4 2 ± 0.5 2 (31) 1-3 (10) 1-3 Digit 5 1 1 (29) 1 (9) 1 Number of cervicals 2 ± 0.42 2 fused (75) 2-4 (25) 2 Last vertebra with 66 ± 1.56 68 facet for chevron (65) 61-70 (24) 59-70 Widest vertebra 23 ± 1.42 24 ± 1.46 (70) 21-26 (25) 22-27 of the long-beaked form. Long-beaked specimens have been collected from coastal central Peru (Banks and Brownell, 1969). The “Guerrero common dol- phin” that occurs along the west coast of Mexico could be either the long-beaked form or a large- bodied population of the short-beaked form (Perrin et al., 1985). In the eastern North Pacific, all sightings of long- beaked animals have been within about 100 nau- tical miles of shore (Perrin et al., 1985; Hill and Barlow, 1992). The long-beaked form off South Africa also seems to have a nearshore distribution (Ross, 1984). Nishiwaki (1967), based primarily on the work of Ogawa (1936), described and mapped the dis- tribution of common dolphins in the western Pa- cific. He stated that the long-beaked form, D. ca- pensis, favors warm waters and is only found in the more southern regions of Japan such as the waters off Kyushu during the summer. The short-beaked form, D. delpbis, avoids cooler waters but during summer may be found offshore north of Honshu in the warm Kuroshio current. TAXONOMY For all species of delphinids studied to date, mod- erate to strong patterns of morphological variation over sometimes short geographical distances seem to be typical (Perrin, 1984). For most species, only small samples are available from restricted locali- ties. The strong geographical variation and small sample sizes have hampered studies regarding al- pha-level taxonomy, because samples collected over wide areas are then needed to distinguish between intraspecific (gamma-level taxonomy) and interspe- cific differences (e.g., Perrin et al., 1981, 1987). The current study is unique in that the study area is quite small and thus theoretically free from the effects of geographical variation. Also, the samples of both forms are large enough to support statistical analysis, even when stratified by sex and age. One interpretation is that the above evidence of complete morphological separation of these sym- patrically occurring forms suggests that the forms represent two species. Wiley (1981:62) considered such cases of completely distinct phenotypic pop- ulations in a region of sympatry prima facie evi- dence for two species. In addition to our morpho- logical data, initial research utilizing molecular data also suggests two species. Based on analyses of mi- tochondrial DNA sequence data, Rosel et al. (in press) found that the California short-beaked form was more closely related to common dolphins from the Black Sea than to the sympatrically occurring long-beaked form. Based on the ratio of rostrum length to zygomatic width, Banks and Brownell (1969) also found that Black Sea animals clustered with the short-beaked form from the eastern North Pacific. An alternative hypothesis is that various demes of common dolphins form a Rassenkreis and the two forms we describe are examples of the ex- tremes in a single species that do not interbreed. Although the Rassenkreis phenomenon is consid- ered rare for large vagile vertebrates (Mayr, 1969), one potential delphinid example has been pro- posed. Ross (1977) examined a small series of bot- tlenose dolphins from South Africa and distin- guished two species, Tursiops truncatus and T. aduncus, based on differences in size, morphology, and coloration. Subsequently, in evaluating speci- mens of Tursiops sp. from Australia, Ross and Cockcroft (1990) found that the two forms off South Africa appear to represent the extremes of a dine 18 ■ Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus Delphinus Table 7. Means, standard deviations (for sample size > 24), sample sizes (in parentheses), and minimum and maximum values for postcranial measures for adult specimens of the two forms of common dolphins from California. Males Females Short-beaked Long-beaked Short-beaked Long-beaked Width of atlas 126.5 ± 8.1 133.3 120.0 ± 5.6 127.4 (26) 110-143 (6) 125-142 (25) 108-129 (5) 121-135 Height of atlas 51.7 ± 2.7 54.5 48.1 53.8 (26) 46-58 (6) 53-56 (24) 42-52 (5) 50-58 Length of atlas lateral process 24.7 ± 2.4 27.8 23.4 25.4 (26) 19-30 (6) 27-30 (24) 20-28 (5) 24-28 Length of atlas dorsal spine 45.6 ± 4.7 50.2 44.5 ± 4.2 50.2 (26) 35-56 (6) 46-54 (25) 38-55 (5) 47-54 Height of first thoracic vertebra 47.5 51.0 45.1 51.0 (11) 42-52 (4) 50-52 (13) 41-49 (2) 47-55 Width of first thoracic vertebra 88.0 ± 5.8 90.3 82.6 88.6 (26) 77-100 (6) 85-95 (23) 74-94 (5) 79-99 Length of first thoracic spine 33.2 39.0 28.7 30.8 (23) 23-41 (6) 34-45 (23) 20-48 (5) 21-35 Height of first lumbar vertebra 51.0 57.4 48.3 54.2 (23) 48-56 (5) 55-59 (23) 41-53 (5) 50-58 Width of first lumbar vertebra 191.6 ± 9.0 204.5 184.3 200.2 (25) 177-208 (6) 192-211 (24) 165-200 (5) 187-216 Length of first lumbar spine 74.2 80.2 68.4 75.8 (24) 66-84 (6) 75-84 (24) 56-78 (5) 71-82 Length of first vertebral rib 121.5 ± 6.0 132.0 112.2 ± 4.1 121.0 (25) 112-131 (6) 121-142 (25) 105-121 (4) 115-130 Length of longest vertebral rib 269.6 298.8 256.2 ± 9.0 275.5 (24) 249-287 (6) 287-322 (25) 241-278 (4) 268-285 Maximum width of manubrium 102.2 ± 5.7 114.8 96.6 ± 4.6 108.0 (26) 91-112 (5) 105-121 (25) 87-109 (4) 99-122 Height of scapula 126.1 ± 6.1 139.0 118.2 ± 6.8 128.5 (25) 117-139 (5) 136-144 (25) 104-130 (4) 118-143 Length of scapula 122.0 ± 6.6 135.0 112.6 ± 6.5 199.0 (25) 112-132 (5) 133-137 (25) 101-123 (4) 110-129 Maximum length of humerus 56.6 62.0 52.9 ± 3.3 58.3 (24) 50-62 (5) 57-65 (25) 47-61 (4) 57-60 Maximum length of radius 76.7 83.8 70.6 ± 5.0 76.8 (23) 70-84 (5) 81-88 (25) 63-82 (4) 65-86 Maximum length of ulna 66.6 73.6 61.4 ± 5.1 70.0 (23) 59-73 (5) 70-78 (25) 54-73 (4) 68-72 Maximum width of humerus 40.4 42.8 36.8 41.5 (24) 37-43 (5) 41-44 (24) 33-40 (4) 38-46 Length of longest chevron 46.3 ± 4.1 51.3 41.8 ± 3.8 48.0 (26) 37-53 (6) 48-59 (25) 34-50 (5) 41-54 Length of longest pelvic 92.7 93.4 62.9 73.8 (20) 64-111 (5) 79-101 (17) 47-76 (5) 58-84 Width of widest vertebra 193.9 ± 9.6 207.4 187.3 202.8 (25) 174-214 (5) 200-211 (23) 169-205 (5) 187-220 Length of centrum of first lumbar 23.5 ± 1.8 25.5 21.8 24.2 (26) 20-27 (4) 23-30 (23) 20-24 (5) 22-27 found along the coasts of Australia. This cline, par- ticularly relating to body size, is strongly correlated with water temperature. We believe our study does not represent such a pattern and that their study does not represent a Rassenkreis for the following reasons. First, the two forms of Tursiops occurring off the coast of South Africa are essentially allo- patric, associating with different bodies of waters or currents (Ross, 1977, 1984; Ross and Cockcroft, 1990), and thus are potentially physically isolated from each other. The two forms of common dol- phins off the coast of southern California often are seen within minutes and kilometers of each other, so no physical barrier can be invoked as a genetic Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus DelphinusM 19 25 73 74 75 76 77 78 79 80 81 VERTEBRAL COUNT ■■ short-beaked long-beaked Figure 14. Total vertebral counts for common dolphins from waters off southern California. Filled bars are short- beaked specimens and hatched bars long-beaked specimens. isolating mechanism. Second, the differences be- tween the Southern African forms of T ursiops were based on extremely small samples (n = 5 for adult T. truncatus; Ross, 1977, table 1). Intermediates between the two forms may be found when a larger sample is examined. Finally, the two forms of com- mon dolphins are found sympatrically in several distinct regions (e.g., western Pacific, eastern Pacific, eastern Atlantic). A Rassenkreis would not explain this type of pattern. We conclude that the two forms of common dolphins represent two distinct species. Below, we review the nomenclature for Delphinus, provide re- descriptions of the two species, and suggest further research. REVIEW OF DELPHINUS NOMENCLATURE The annotated list of the nominal species of com- mon dolphins provided here is based largely on Hershkovitz (1966). We converted measurements in the original descriptions to metric units (centi- meters for external morphometries and millimeters for cranial morphometries). Based on the type ma- terial, we attempted to assign each nominal species Table 8. Means, standard deviations (for sample size > 30), sample sizes (in parentheses), and minimum and max- imum values for the ratio of rostrum length to zy- gomatic width for adult specimens from the coast of California. Short-beaked Long-beaked Males 1.37 ± 0.046 1.60 (47) 1.21-1.46 (19) 1.52-1.67 Females 1.36 ± 0.055 1.64 (48) 1.23-1.47 (12) 1.55-1.77 to either the long-beaked or short-beaked form of Delphinus as recognized and diagnosed herein. There are numerous problems inherent in such an exercise. Foremost is that most of the type spec- imens lack data relating to maturity, either sexual or physical. Thus, many type specimens may be unsuitable for comparative studies because they may be juveniles. For the majority of type specimens, not even the sex is recorded. Published tooth counts are also a potential problem. In a previous study of tooth counts in spotted dolphins ( Stenella ), Per- rin et al. (1987) found that their tooth counts were typically higher than those listed in previous works. They believed that previous workers overlooked the anteriormost teeth because they are very small and/or buried in remnants of tissue. We found this to be true for specimens of Delphinus as well. For example, the published tooth counts for the type of D. major (Gray, 1866a) are listed as 46/47, whereas our counts for this specimen are 52/49. Therefore, based upon high tooth counts, we are confident in referring that type to the long-beaked form; the same is true for referring very low-tooth- count specimens to the short-beaked form. For specimens at the upper range of the short-beaked form, we are less confident regarding assignments based on this character. Lastly, description of col- oration is often not useful as the pigmentation of cetaceans darkens quickly postmortem (Norris and Prescott, 1961; Mitchell, 1970) and many descrip- tions and illustrations are obviously not from fresh specimens. We consulted with the original description for each species and note in the text if we were able to examine any type material. Several of the type specimens are housed in the Natural Flistory Mu- seum, formerly the British Museum (Natural Flis- tory). These specimens are listed here with the orig- inal museum acronym (BM) as found in the literature. 20 ■ Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus Delphinus YEARS D. del phis D. capensis Figure 15. Strandings by year of common dolphins classified to species from southern California. The differences of absolute abundance of strandings reflects, in part, differences in response effort to strandings. Filled bars are short- beaked specimens and hatched bars long-beaked specimens. We did not include the “varieties” of common dolphins ( variegatus , balteatus, moschatus, and marginatus ) listed by Fischer (1881), because they were used to describe color variation. According to the International Code for Zoological Nomen- clature (Article 45g), such designated varieties pub- lished prior to 1961 are considered subspecies. It is beyond the scope of this paper to discuss the validity of subspecies. Delphinus delphis Linnaeus, 1758 No type specimen was designated. This species is based on description of an animal from “ Oceano europaeo ” by Artedi (1738). Delphinus vulgaris Lacepede, 1804 This species is represented by an illustration of the whole animal and skull. The illustration is an ex- tremely crude rendering of a dolphin that we be- lieve is not referable even to genus. The published tooth count of 42-38/47-47 is well within the exclusive range of the short-beaked form and the type description indicates that Lacepede (1804) in- tended this species to supplant D. delphis. Delphinus capensis Gray, 1828 The type specimen (BM 41.1734) was a stuffed whole mount with skull inside collected from the Cape of Good Hope housed in the Natural History Mu- seum (London). The published external measure- ments are as follows: total length, 206; maximum girth, 107; snout to blowhole, 33; snout to melon apex, 18 cm; snout to dorsal fin, 97 cm; dorsal fin height, 25; fluke width, 46 cm. The tooth counts are reported as 50/50. Gray (1828) noted that this species is distinguished by its relatively short beak. This comment appears to be made in comparison to Stenella longirostris, then regarded to be in the genus Delphinus. The skull was later removed from the mount. We measured the type and found the rostral ratio to be 1.71 and the tooth counts 54- 53/51-52. The maxillae and premaxillae are fused along the entire length of the rostrum, indicating the specimen is an adult animal. The skull mea- surements and tooth counts of the type specimen associate it with the long-beaked form from Cali- fornia (Banks and Brownell, 1969; van Bree and Purves, 1972). Delphinus longirostris G. Cuvier, 1829 This name is preoccupied by D. longirostris Gray, 1828, now placed in the genus Stenella (Hershko- vitz, 1966). Delphinus novae-zelandice Quoy and Gaimard, 1830 The following external measurements for the type specimen have been published (Quoy and Gaimard, 1830; Gray, 1850): total length, 188; snout to blow- hole, 33; snout to eye, 30; snout to flipper, 43; snout to dorsal fin, 83. The tooth counts are reported as 43/47, and the skull measurements are as follows: skull length, 356 mm; nose, 203; length of mandible 305. Gray (1850) also noted that the skull was very much like that of D. janira. The low tooth counts associate this nominal species with the short-beaked form. Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus DelphinusM 21 Delpbinus loriger Wiegmann, 1846 This name was published in 1846 in Schreber’s Saugtbiere (Wagner, 1846). Hershkovitz (1966) considered this species to be a junior synonym of Stenella dubia (G. Cuvier, 1812). Perrin et al. (1987) stated that based on the published pigmentation pattern this specimen was a common dolphin. This species is only known from a description and il- lustration of a whole animal (Wagner, 1846, PI. CCCLXII). The animal had a flipper stripe like that of the short-beaked form. Thus, we consider this species to be a junior synonym of D. delpbis. The year this species was named is problematic. Hershkovitz (1966) listed the author and date as “Wiegmann, 1841 (?)” and secondarily listed the 1846 edition of Schreber’s Saugtbiere edited by J. Wagner. This listing has been followed by Hall (1981) and Perrin et al. (1987). This ambiguity may be due to the fact that Wagner (1846) mentioned that the species was based on the papers of A. Wiegmann. We examined the volumes of Arcbiv fur Naturgescbichte for the years 1836 through 1847. This series was edited by A. Wiegmann in the late 1830s and early 1840s and includes a sup- plement that summarized the published biological work of the previous year. There is no mention of D. loriger in any volume. Sherborn (1935) consid- ered that Wiegmann had published this species name in the 1846 edition of Saugtbiere. According to the Catalogue of the Books, Manuscripts, Maps and Drawings in the British Museum (Natural History) (1903:1861), A. Wiegmann began editing the 1846 edition of Schreber’s Saugtbiere, but the volume was completed by Wagner; this may explain Wag- ner’s mention of the “papers” of Wiegmann. Delpbinus janira Gray, 1846 The type specimen is a skull collected from New- foundland and housed in the Museum of the Bristol Institution. The published diagnostic features and measurements are listed as the following: skull length, 453; skull width at orbit, 195; length of mandible, 318; tooth counts 43/42. The very low tooth counts associate this nominal species with the short-beaked form. Delpbinus sao Gray, 1846 The type of this species is a skull from Madagascar that Gray stated was located in the Paris Museum. However, Robineau (1990) does not list this spec- imen in his list of cetacean types in the Museum national d’Histoire naturelle, Paris. Measurements and characters published by Gray are as follows: tooth counts, 55 / 55-60 /60; skull length, 430; beak length, 267; tooth row length (upper?), 222. Hersh- 22 ■ Contributions in Science, Number 442 kovitz (1966) erroneously listed this species as au- thored by Gray (1850). The tooth count is above the range of the short-beaked form and, thus, this nominal species groups with the long-beaked form. Delpbinus forsteri Gray, 1846 This species is based on a description and unpub- lished illustration of an animal from near New Cal- edonia by Forster (1844). Gray (1846) published this illustration and noted that there were white patches on the dorsal fin and flippers of this specimen. The tooth count was listed as 44/44. Based on the low tooth count and the distinct light patches on the flippers and dorsal fin, we tentatively refer this spe- cies to the short-beaked form. Delpbinus fulvifasciatus Wagner, 1846 The type specimen was collected off Hobart Town, Tasmania and housed in the Museum national d’Histoire naturelle, Paris (CAC: A. 3025). Robi- neau (1990) published the following skull mea- surements: condylobasal length, 432; rostrum length, 270; rostral width at base, 90; preorbital width, 173; postorbital width, 191; zygomatic width, 184; length of mandible, 383; height of ramus, 68. The tooth counts are listed as 42 + , 45/44, 45. Based on the published skull measurements, the ratio of rostral length to zygomatic width of 1.47 associates this specimen with the short-beaked form. Delpbinus albimanus Peale, 1848 The type originally consisted of a mounted skin with partial mandibles and manus bones within, collected off the coast of Chile. A rather crude illustration of the whole type specimen was pub- lished by Cassin (1858). True (1889) examined the type mounted skin, dissected out the partial man- dibles, and commented that he believed that this species was like D. forsteri, a variety of D. delpbis with white patches on the flipper. The mounted skin of the type specimen is now lost (Poole and Schantz, 1942), but the mandibles are still available in the National Museum of Natural History. Based on the distinct white patches on the flipper and the comments of True (1889), we tentatively refer this nominal species to the short-beaked form. Delpbinus fritbii Blyth, 1859 The type specimen was collected on a voyage from England to India and is represented by a skull in the Calcutta Museum. The published tooth counts are 55/50; the upper count is within the exclusive range of the long-beaked form. Heyning and Perrin: Common Dolphins, Genus Delphinus Delphinus algeriensis Loche, 1860 The type consists of a skin with skull and possibly a complete skeleton collected from the coast of Algiers housed in the Natural History Museum of Algiers. The illustration of the type specimen de- picts an animal with a mixture of Stenella coeru- leoalba and Delphinus spp. coloration. However, the description of the skull (Loche, 1860:477) clear- ly mentions the long grooves on the palate, a char- acter unique to the genus Delphinus. The published tooth counts of 41-41 /41-39 would associate this nominal species with the short-beaked form. Delphinus major Gray, 1866 The type specimen is a skull in the Natural History Museum (BM 1852.10.5.2) from an unknown lo- cality. The original published measurements (Gray, 1866a) of the type skull are the following: condy- lobasal length, 533; rostrum length, 318; rostrum width at notches, 106; mandible length, 445; tooth counts, 46/47. We examined the type skull. The published ratio of rostrum length to skull width is 1.73 (van Bree and Purves, 1972), whereas our mea- surements yield a slightly different ratio of 1.70. We examined the type and report the tooth counts as 52/49. We found almost complete fusion of the maxillae to the premaxillae along the rostrum, sug- gesting adulthood. Both tooth counts and the ros- tral ratio place this species with the long-beaked form from the eastern North Pacific. This is in agreement with the conclusions of Banks and Brownell (1969) and Evans (1975). Delphinus moorei Gray, 1866 The type specimen is represented by a skull in the Liverpool Free Museum, from a 192-cm female collected southwest of the Cape of Good Hope. The original published measurements of the type skull are as follows: skull length, 445; rostrum length, 279; mandible length, 368; rostral width at notch, 91; rostrum width midlength, 49; tooth counts, 44/48. External measurements (in cm) are as follows: 192; beak length, 14.6; snout to eye, 33; snout to blowhole, 33; snout to flipper, 47; snout to anus, 141; snout to genital slit, 138. The published illustration of a whole specimen (Gray, 1866b:736, fig. 1) depicts a nondescript, three-tone dolphin. However, the lack of white below the eye is characteristic of the long-beaked form. For a female of this size, the rostrum tip to eye and ros- trum to melon apex measures are above the range of the short-beaked form. Also, the condylobasal length and rostrum length are at the lower ends of ranges of the long-beaked form and above the rang- es of the short-beaked form. The lower tooth count is in the region of overlap, but the upper count is in the short-beaked range. However, the published tooth counts are likely to be low. We consider this species referable to the long-beaked form. Delphinus walkeri Gray, 1866 The type specimen is a skull in the Liverpool Free Museum, from a 202-cm female collected south- west of the Cape of Good Hope. We have not examined it. This species was described by Gray (1866b) as being extremely similar to D. moorei, with the following features: skull length, 419; ros- trum length, 279; rostrum width at notches, 85; rostrum width midlength, 47; tooth count, 47/49. The external measurements were described as being very similar to those of D. moorei. The published illustration (Gray, 1866b:737, fig. 2) does not pro- vide enough details to ascertain which form this species represents. Delphinus pomeegra Owen, 1866 The type specimen is a skull collected from the Madras coast of India now housed in the Natural History Museum (BM 1478a-66.2.5.5). The tooth counts given in the original description are listed as 42-41/45-46. In Owen’s (1866, PI. 8) illustra- tion, the tip of the rostrum appears somewhat worn. True (1889) remarked that the tip is damaged and estimated that about 7 cm are missing. Based on our examination of the skull, we believe this may be an overestimate of the amount of the rostrum that is missing. Comparing the overall skull mor- phology to other specimens, we estimate that 2 cm or less are missing. There is no fusion of the max- illae to the premaxillae along the rostrum, indicat- ing that the specimen was not yet sexually mature and the tip of the rostrum was still growing (Perrin and Heyning, 1993). This specimen is from the northern Indian Ocean, where so far only animals with very long rostra have been documented, such as the type specimen and referred specimens of D. tropicalis. The published coloration of the type specimen as very dark, almost black above and lighter below, appears to represent postmortem darkening of the coloration, as no form of Del- phinus that we know of has such a simple color pattern. Because the type specimen is immature and the rostrum is damaged, we regard this species as a nomen dubium. Eudelphinus tasmaniensis Van Beneden and Gervais, 1880 This species is based on a skull in the Museum national d’Histoire naturelle, Paris (CAC: A. 3071) collected in Tasmania. Robineau (1990) was unable to locate this skull in the collection. No type de- scription is provided, only an illustration of the palate and pterygoid region (Van Beneden and Ger- vais, 1880, PI. 39, fig. 9). True (1889) reported a Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus DelphinusM 23 condylobasal length of 445 mm and an orbital width of 168 mm. The upper tooth counts (53-50) of the type obtained from the illustration cannot be con- sidered accurate as the counts widely differ bilat- erally in other specimens on the same plate. We consider this species to be a nomen dubium. Delphinus dussumieri Blandford, 1891 This name was proposed for D. longirostris G. Cu- vier, 1829, which is preoccupied by Stenella lon- girostris (Gray, 1828). However, D. dussumieri is preoccupied by D. dussumieri Fischer, 1829 as a new name for D. capensis F. Cuvier, 1829, which in turn is preoccupied by D. capensis Gray, 1828. Both D. dussumieri Fischer, 1829 and D. capensis F. Cuvier, 1829 are considered junior synonyms of Cephalorhynchus heavisidii (Hershkovitz, 1966). Delphinus tropicalis van Bree, 1971 was proposed as a new name for the species to which the names D. longirostris G. Cuvier and D. dussumieri Bland- ford, 1891 had been applied. Delphinus microps Burmeister, 1866 (not Gray) This species is based on a short description of three skulls collected from the coast of Brazil. The skulls are all about 460 mm long and have a zygomatic width of about 190 mm with a tooth count range of 47-49/44-48. Burmeister’s original description of the types noted that this species appears almost identical to D. walkeri. The skull length and width are within the range of the long-beaked form and outside the range for the short-beaked form. The tooth counts are low for the long-beaked form, but, as mentioned previously, the true counts may be higher than given in some of the original de- scriptions. We therefore consider this type attrib- utable to the long-beaked form. Delphinus marginatus Lafont, 1868 (not Pucheran, 1868) Hershkovitz (1966) considered this species to be based on a misidentification and therefore not a new species name. Delphinus bairdii Dali, 1873 This species was based on two specimens collected by Charles Scammon off Point Arguello, California. The type specimens are lost (Poole and Schantz, 1942). See discussion below. Delphinus tropicalis van Bree, 1971 This is a new name proposed for D. longirostris and D. dussumieri, as both names are unavailable. 24 ■ Contributions in Science, Number 442 The type specimen is a skull collected from the Malabar coast housed in the Museum national d’Histoire naturelle in Paris (specimen numbers CAC: A. 3065 and CAG: B 11/64; Robineau, 1990). We did not examine the skull. The tooth counts of the type are reported as 65-65/5 7-58 and the ratio of rostral length to zygomatic width as 2.06 (van Bree and Gallagher, 1978). Both the tooth counts and the ratio of rostral length to zygomatic width are above the range of values for the long- beaked form off California. In reviewing the nomenclature, we find that the senior synonym for the short-beaked form is Del- phinus delphis Linnaeus, 1758 and for the long- beaked form Delphinus capensis Gray, 1828. The species Delphinus delphis is based on the description by Linnaeus (1758) of “ corpore oblongo subtereti, rostro antenuato acuto .” This diagnosis was used to differentiate the common dolphin from Orcinus orca and Phocoena phocoena, then con- sidered to be congeneric. No type was designated, and the type locality was listed as “ Oceano Euro- paeo.,'> The lack of a type specimen and detailed diagnosis raises several potential problems of no- menclature. The first relatively complete description of a common dolphin from European waters is provid- ed by Lilljeborg (1866). This specimen of unknown sex was collected from Kristiania Bay, Norway and housed in the Zoological Museum of the University of Kristiania (Oslo). The total length of the speci- men was 181 cm with a snout to melon apex length of 10.5 cm. The ratio of rostral length to zygomatic width is 1.48. Both the external measurement of the snout relative to total length and the rostral ratio of the skull are within the exclusive ranges of values for the short-beaked form. The next rather detailed description of a com- mon dolphin from European waters is that of Flow- er (1880). The specimen he described was an im- mature female 156 cm in length caught off the Cornwall coast of the United Kingdom in 1878. The complete skeleton was collected and housed originally in the Museum of the Royal College of Surgeons, then later transferred to the Natural His- tory Museum, London. This animal was illustrated (Flower, 1880) by a watercolor of moderately good quality that depicts some strong accessory stripes, such as the primary and secondary abdominal stripes and the eye-to-anus stripe of Mitchell (1970). The flipper stripe intersects with the lower jaw about one-third the distance anterior to the hind-end of the gape. Based on the illustrated color pattern, it is not possible to assign this specimen to one or the other of the forms we describe from off California. The tooth counts of Flower’s specimen were listed as 46-44/48-47. This is near the mean number of teeth we found for the short-beaked form and just at or below the lower limit for the long-beaked form off California. Both of the above early described specimens of common dolphins from European waters were of Heyning and Perrin: Common Dolphins, Genus Delphinus the short-beaked form. Additionally, the range of values for zygomatic width to rostrum length (1.37- 1.54) for 31 specimens (albeit not stratified by age or sex) from European waters (van Bree and Purves, 1972) clustered with the short-beaked form. As we are unable to find specimens of the long-beaked species from European waters, we refer the short- beaked form to the species Delpbinus delphis. The type of Delpbinus bairdii was described by W. H. Dali (1873) based on two females caught off Point Arguello, California by Captain Charles Scammon. In the original description, Dali (1873) mentioned that one entire skeleton was sent to the U.S. National Museum. Poole and Schantz (1942) noted that there are no records of the specimen arriving at the museum, nor are there catalog num- bers for it. The type specimen has not been found subsequently (J. Mead, National Museum of Nat- ural History, pers. comm.). What is available for the syntypes are the total lengths, a few skull measure- ments, and tooth counts published in the original description. In addition, Scammon (1874) illustrat- ed the pigmentation of one of the syntypes, which we reproduce here (Fig. 16). As the type specimens have been lost, it is im- portant to firmly establish which of the two forms present off California was considered D. bairdii in order to stabilize the nomenclature. The tooth counts for the type specimen were reported as 53/ 47 (Dali, 1873). The upper tooth count is at the upper limit of the tooth count we found for the short-beaked form and well within the range of the long-beaked form. The lower tooth count is at the minimal range of tooth counts for the long-beaked form but well within the range of the short-beaked form. However, Dali (1974) mentioned that there was room for four to five additional teeth in each mandibular tooth row. Tooth counts as originally listed are often lower than those obtained by to- day’s methods (see Perrin et al., 1987). Thus, it is possible that the tooth count of the type specimen of D. bairdii would be higher and within the ex- clusive range of the long-beaked form if re-exam- ined today. The total lengths of the two females mentioned in the type description (Dali, 1873) were 201 and 206 cm. These lengths fall well outside the range of total length for the short-beaked form and are very close to the average length (207.7 cm) of phys- ically mature female iong-beaked common dol- phins in this region. The condylobasal length and the rostrum length of the type skull are significantly above the range of these measures for female short- beaked specimens and even above the mean con- dylobasal and rostrum length for the long-beaked specimens we examined. The illustration provided by Scammon (1874) depicts an animal clearly as- signable to the long-beaked form based on the cri- teria we used in our analysis of color pattern, es- pecially the flipper stripe angling toward the corner of the mouth. Therefore, based on total length, skull measures, and pigmentation, the types of D. bairdii belong to the long-beaked form. In order to provide taxonomic stability, we pro- pose a neotype for Delpbinus bairdii, LACM 84240. This specimen consists of a complete skeleton of a physically mature male. The external measure- ments and skull and postcranial data are listed in Tables 9-11. A photograph of the whole specimen is found in Figure 8 and a photograph of the skull in Figure 18. In their analysis of rostral length vs. zygomatic width, Banks and Brownell (1969) found that the type specimens of D. bairdii Dali, 1873; D. major Gray, 1866; and D. capensis Gray, 1828 were in- cluded in a cluster with the long-beaked form. In their review of the previous study, van Bree and Purves (1972) noted that there was no reason to call the long-beaked form D. bairdii, because there are no described differences between D. capensis, D. major, and D. bairdii, and D. capensis is the senior synonym. The published values (van Bree and Purves, 1972) for the ratios of zygomatic width to rostral length for D. capensis (1.74) and D. major (1.73) clearly group these two nominal species with the long-beaked form from California (see Table 10). The long-beaked forms from South Africa and California are similar in overall skull shape, includ- ing relative rostral length, in general coloration pat- tern, and in tooth count. The two populations differ significantly in total vertebral count. We believe that this difference alone does not warrant assigning species status to these populations. If they are con- sidered one species, the range of vertebral count is nine (72-80). Perrin et al. (1987) found that the Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus DelphinusM 25 Figure 17. Typical skull of adult Delphinus delphis from the waters off southern California (LACM 84225, male, total length 188 cm). 26 ■ Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus Delphinus Figure 18. Typical skull of adult Delphinus capensis from the waters off southern California (LACM 84240, male, total length 235 cm). This specimen is also designated as the neotype of Delphinus bairdii, considered to be a junior synonym. Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus DelphinusM 27 Table 9. External morphometries (in centimeters) for the neotype of Delphinus bairdii (LACM 84240). Total length 234.5 Snout to anus 170.0 Snout to genital slit 145.0 Snout to umbilicus 107.0 Snout to dorsal fin tip 133.0 Snout to anterior dorsal fin 99.0 Snout to flipper 46.5 Snout to ear 37.5 Snout to eye 33.0 Snout to gape 27.5 Snout to blowhole 33.5 Snout to melon apex 14.5 Eye to ear 5.5 Eye to gape 5.5 Right eye to blowhole 18.5 Left eye to blowhole 16.5 Blowhole length 1.7 Blowhole width 2.0 Head diameter at eyes 19.5 Rostral width at melon apex 7.5 Projection of lower jaw 1.0 Flipper length, anterior 33.0 Flipper length, posterior 24.0 Flipper width 11.0 Length of genital slit 12.0 Fluke width 58.0 Fluke depth at lobe 16.0 Fluke depth at notch 13.5 Fluke notch depth 3.0 Dorsal fin height 24.5 Dorsal fin base length 33.0 Girth at eye 71.0 Girth at axilla 105.0 Girth, maximum 118.0 Girth at anus 78.0 Girth midway anus-fluke notch 38.0 Caudal height 16.5 Caudal thickness 4.2 pan-tropical spotted dolphin ( Stenella atennuata) has a range of nine in total number of vertebrae. Thus, the variation noted in the long-beaked com- mon dolphin is comparable to the range of variation found in another widely distributed delphinid. Evans (1982) found that sexually mature animals from the eastern Pacific formed two discrete clus- ters; the few available specimens of D. tropicalis formed another cluster, with rostral lengths even greater than in the long-beaked form from the east- ern Pacific. The sample size of Delphinus sp. spec- imens from the Indian Ocean is still small. One specimen from this region has a rostrum-to-zygo- matic width ratio of 1.72, within the range of D. capensis. Another specimen from South African waters has a ratio of 1.87, near the lower range of values for D. tropicalis. If D. tropicalis is a valid species, then by using our criteria D. capensis also occurs in the Indian Ocean. The alternative hy- Table 10. Skull measurements (in millimeters) and tooth counts for the neotype of Delphinus bairdii (LACM 84240). Condylobasal length 469 Rostral length 295 Rostral width At base 93 At 14 length 60 At Vi length 55 At % length 46 Premaxillae width at V2 length 27 Tip of rostrum to external nares 346 Tip of rostrum to internal nares 339 Preorbital width 167 Postorbital width 190 Zygomatic width 185 Width of external nares 45 Greatest width of premaxillae 70 Parietal width 156 Height of braincase 109 Length of braincase 114 Length of temporal fossa 72 Height of temporal fossa 53 Length of orbit 47 Length of antorbital process 42 Width of internal nares 55 Length of upper tooth row 246 Length of lower tooth row 237 Length of ramus 339 Height of ramus 74 Tooth width 3.7 Upper teeth, left 58 Upper teeth, right 55 Lower teeth, left 51 Lower teeth, right 52 pothesis is that rostral length increases clinally in the Indian Ocean. The coloration of specimens of the nominal D. tropicalis is very similar to that of D. capensis from the west coast of Africa, and thus on this evidence D. tropicalis may be a junior syn- onym of D. capensis. RE-DESCRIPTION OF DELPHINUS DELPHIS Delphinus delphis Linnaeus, 1758 Delphinus vulgaris Lacepede, 1804 Delphinus nova-zealandice Quoy and Gaimard, 1830 Delphinus zelandce [sic] Gray in Dieffenbach, 1843 D[elphinus] novae zeelandice [sic] Wagner 1846 Delphinus novae zealandice [sic] Gray 1850 Delphinus longer Wiegmann, 1846 Delphinus fulvifasciatus Wagner, 1846 Delphinus janira Gray, 1846 Delphinus foresteri Gray, 1846 Delphinus albimanus Peale, 1848 Delphinus algeriensis Loche, 1860 28 ■ Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus Delphinus HOLOTYPE. None. TYPE LOCALITY. Oceano Europaeo. DIAGNOSIS. The coloration consists of the ba- sic criss-cross pattern with the thoracic patch rel- atively light gray to a medium golden-yellow in color, contrasting sharply with the very dark gray to black spinal field. The flipper-to-anus stripe is weakly formed or absent in most animals. One or more abdominal accessory stripes may be present. The flipper stripe does not angle toward the corner of the gape and fuses with the lip patch one-third to one-half of the gape length anterior to the corner of the mouth. The flipper stripe narrows distinctly anterior to the eye. There is always a wide, lightly pigmented region between the flipper stripe and the eye which extends anteriorly to the gape region. The eye patch and bridle are black, contrasting sharply with the adjacent thoracic patch. The white of the abdominal field extends above the flipper stripe to at least under the eye. Light gray to white patches with diffuse edges are found on the dorsal fin and flippers of many adults. This species of common dolphin is relatively heavier and deeper-bodied anteriorly, with a more rounded melon that inserts onto the beak at a sharp angle when viewed in profile. The flippers and dor- sal fin are larger than in D. capensis. The total length ranges from 172 to at least 223 cm in mature males and from 164 to 215 cm for mature females. The tooth counts range from 42/41 to 54/53. The vertebral formula range is as follows: total, 74-80; thoracic, 13-16; lumbar, 18-24; caudal, 30- 36. The ratio of rostral length to zygomatic width for adults ranges from 1.21 to 1.47 (Fig. 17). DISTRIBUTION. This species is found in tem- perate and tropical waters of all major oceans and some seas (Fig. 19). In the North Pacific, D. delphis has been found from British Columbia south to Chile and out to 135°W. There are few records from the Gulf of California. In the western Pacific this species is documented from New Caledonia, New Zealand, and Japanese waters. There are rec- ords from north of Hawaii. Thus, the range may extend entirely across the tropical and temperate North Pacific. Records from the western North Atlantic range from at least Florida to Newfound- land and in the eastern Atlantic from northern Eu- rope south to the west coast of Africa. This species is found in the Mediterranean and Black Seas. GEOGRAPHICAL VARIATION. Accessory stripes are more common on animals from the North Atlantic. The distinct flank blaze found on animals from the North Atlantic is less conspicuous or ab- sent on animals from the eastern North Pacific. RE-DESCRIPTION OF DELPHINUS CAPENSIS Delphinus capensis Gray, 1828 Delphinus sao Gray, 1846 Delphinus frithii Blyth, 1859 Delphinus major Gray, 1866 Table 11. Postcranial measurements (in millimeters) and meristics for the neotype of Delphinus bairdii. Vertebral count 79 No. of lumbar vertebrae 22 No. of caudal vertebrae 36 First vertebra with vertical foramen 55 Last vertebra with transverse process 59 Last vertebra with neural arch 66 First vertebra with chevron 44 No. of chevrons 29 No. of vertebra! ribs 15 No. of sternal ribs 8 Number of carpals and metacarpals 10 Number of cervicals fused 2 Last vertebra with facet 69 Widest vertebra 22 Width of atlas 138 Height of atlas 54 Length of atlas lateral process 30 Length of atlas dorsal spine 54 Height of first thoracic vertebra 52 Width of first thoracic vertebra 95 Length of first thoracic spine 34 Height of first lumbar vertebra 59 Width of first lumbar vertebra 211 Length of first lumbar spine 82 Length of first vertebral rib 142 Length of longest vertebral rib 294 Maximum width of manubrium 118 Height of scapula 140 Length of scapula 133 Maximum length of humerus 65 Maximum length of radius 84 Maximum length of ulna 73 Maximum width of humerus 44 Length of longest chevron 59 Length of longest pelvic 99 Width of widest vertebra 211 Length of centrum of first lumbar 30 Delphinus moorei Gray, 1866 Delphinus microps Burmeister, 1866 Delphinus bairdii Dali, 1873 Delphinus bairdi [sic] Norris and Prescott, 1961 HOLOTYPE. Whole mount with skull (BM 41.1734) in the Natural History Museum (London). TYPE LOCALITY. Cape of Good Hope. DIAGNOSIS. The overall pigmentation pattern is more muted than that of D. delphis. The ground coloration consists of a criss-cross pattern with the thoracic patch relatively darker, not contrasting as sharply with the dark gray spinal field as in D. del- phis. The flipper-to-anus stripe is weakly to strongly formed. The flipper stripe angles toward the corner of the mouth and fusees with the lip patch at the corner of the gape to one-third anterior along the gape or closely parallels the gape. The flipper stripe narrows moderately anterior to the eye. The eye patch does not contrast strongly with the adjacent Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus DelphinusM 29 -a > X) T3 a/ u=: c cu "O a, C/5 C o -a Qh 03 .ij P c & W> < ■§ 30 ■ Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus Delphinus 100° 120° 140° 160° 180° 160° 140° 120° 100 Figure 20. Approximate known distribution of the long-beaked common dolphin, Delphinus capensis, based on specimen records or sightings at sea identified by g characters listed in this paper. Records of the nominal species D. tropicalis are represented by x s. thoracic patch, and the white of the abdominal field rarely extends above the flipper stripe to below the eye. There may be a slight lightening of the flippers and dorsal fin of some adults. This species is slightly more slender than D. del- phis, with a flatter melon that inserts onto the beak at a more gradual angle when viewed in profile. Total length for mature specimens ranges from 193 to at least 222 cm for females and from 202 to 254 cm for males. The tooth counts are 47-60/47-57. The verte- bral formula is as follows: cervical, 7: thoracic, 14- 17; lumbar, 18-23; caudal, 30-37, for a total of 72-80. The range of ratio of rostral length to zy- gomatic width is 1.52-1.77 (Fig. 18). DISTRIBUTION. Restricted to nearshore trop- ical to temperate waters of some oceans (Fig. 20). In the eastern North Pacific, this species is recorded from Point Conception south to Peru, including the Gulf of California. In the western North Pacific, specimens have been recorded from the coasts of Korea, southern Japan, and Taiwan. In the Atlantic, records are available from coastal Venezuela south to the La Plata region of Argentina. This species is found along the west coast of Africa and in South African and Madagascan waters. GEOGRAPHICAL VARIATION. Animals from the coast of southern Africa have lower vertebral counts (72-76) than animals from the eastern North Pacific (77-80). The animals from southern Africa also may have relatively longer beaks and an overall larger body size than animals from the eastern North Pacific. SPECIATION. These two similar species cur- rently have a parapatric distribution pattern. The narrow regions of sympatry seem to be limited to nearshore waters in several ocean basins. One ques- tion posed by such a pattern of distribution is whether speciation occurred sympatrically or al- lopatrically. Allopatric speciation still seems to be the most favored model (Mayr, 1963; Coyne, 1992). For large, vagile marine species, the most likely geographical isolation would involve entire ocean basins. Both the species of common dolphins are found primarily in tropical to moderate temperate waters, avoiding cold temperate zones. The Amer- icas form a significant barrier to marine organisms between the Atlantic and Pacific Oceans, except for cold temperate forms that range around the tip of South America. Common dolphins are not record- ed from this region (Goodall, 1978). The tip of southern Africa does not presently seem to be a barrier to temperate species between the Indian and Atlantic Oceans. The long-beaked common dol- phin is found in the mixed waters of the Agulhas Current off the southeast coast of southern Africa (Ross, 1984). The cooling of waters during the Pleistocene could have resulted in southern Africa becoming a barrier to tropical and warm temperate species (Davies, 1963). This potential isolation of the Atlantic from the Indo-Pacific waters provides a geographical isolating mechanism for the speci- 32 ■ Contributions in Science, Number 442 ation in common dolphins based on the allopatric model. Unfortunately, our data are not appropriate to test this hypothesis. COMMON NAMES. Because of the nearshore sympatry, the vernacular names “offshore” to de- scribe the short-beaked species and “neritic” or “Baja-neritic” for the long-beaked species are per- haps misleading. We suggest the common names “short-beaked common dolphin” and “long-beaked common dolphin” for the two species. FUTURE RESEARCH Common dolphins are a highly variable, widely dis- tributed group consisting of two or possibly three species. Due to the complexity of this problem, we outline several areas for further research on this genus. First, more molecular-genetic (isozymes and DNA) evidence should be examined over a wide geographical range. Morphological analyses of specimens from other regions could address both interspecific and intraspecific variation of the two species of Delphinus. These studies will better de- fine the species characteristics and population struc- ture. Ecological and distributional studies need to be performed to elucidate non-morphological dif- ferences between the two species. Such ecological studies should focus on regions where the two spe- cies occur sympatrically and compare these to regions where only one species predominates, such as D. capensis in the Gulf of California. The remaining existing type specimens for nom- inal species not here referred to either of the two species should be examined for maturity, relative rostrum length, and accurate tooth counts so their taxonomic status can be resolved. Further sampling and a review of D. tropicalis is needed to determine whether it is an extremely long-beaked form along a cline of D. capensis found in the Indian Ocean or represents a third species of common dolphin. I ACKNOWLEDGMENTS We thank S. Smith and J. Tutak for assistance in gathering data. We thank A. Rea for access to the SDMNH col- lection. Pat Pinkard produced Figure 1. T. Kasuya assisted with references. S. 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Review of geographical stocks of tropical dolphins ( Stenella spp. and Delphinus delphis) in the eastern Pacific. NOAA Technical Report NMFS 28, iv + 28 pp. Pilleri, G., and M. Gihr. 1972a. A rare species of dolphin Delphinus tropicalis van Bree 1971 ( =dussumieri Blanford, 1891) from the coast of Pakistan. Mam- malia 36:406-413. Pilleri, G., and M. Gihr. 1972b. Contribution to the knowledge of the cetaceans of Pakistan with a par- ticular reference to the genera Neomeris, Sousa, Delphinus, and T ursiops and description of a new Chinese porpoise ( Neomeris asiaeorientalis). Inves- tigations on Cetacea 4:107-162. Pilleri, G., and M. Gihr. 1973-1974. Contribution to the knowledge of the cetaceans of the southwest and monsoon Asia (Persian Gulf, Indus Delta, Mal- abar, Andaman Sea and Gulf of Siam). Investigations on Cetacea 5:95-149. Poole, A.J., and V.S. Schantz. 1942. Catalog of the type specimens of mammals in the United States National 34 ■ Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus Delphinus Museum, including the Biological Survey specimens. United States National Museum Bulletin 178, xiii + 703 pp. Quoy, J.R.C., and P. Gaimard. 1830. Voyage de decon- vertes de l’ Astrolabe execute par ordre du Roi, pen- dant les annees 1826-1827-1828-1829, sous le Commandemant de M.J. Dumont D'Urville, Zool- ogie. Paris: J. Tastu, i-1 + 268 pp. + atlas. Robineau, D. 1990. Les types de cetaces actuels du Museum national d’Histoire naturelle II. Delphini- dae, Phocoenidae. Bulletin du Museum National d’Historie Naturelle, Paris 4 (Series 12, Section A, l):197-238. Robson, F. 1976. Thinking dolphins, talking whales. Wellington, New Zealand: A.H. &c A.W. Reed Ltd., vii + 199 pp. Rosel, P.E., A.E. Dizon, and J.E. Heyning. (in press) Ge- netic analysis of sympatric morphotypes of common dolphins (Genus Delphinus ). Marine Biology. Ross, G.J.B. 1977. The taxonomy of bottlenose dol- phins Tursiops species in South African waters, with notes on their biology. Annals of The Cape Provin- cial Museums (Natural History) 1 1(9):135— 194. . 1984. The smaller cetaceans of the south east coast of southern Africa. Annals of The Cape Pro- vincial Museums (Natural History) 15(2):173-410. Ross, G.J.B. , and V.G. Cockcroft. 1990. Comments on Australian bottlenose dolphins and the taxonomic status of Tursiops aduncus (Ehrenberg, 1832). In The bottlenose dolphin, ed. S. Leatherwood and R. R. Reeves, 101-128. San Diego: Academic Press. Scammon, C.M. 1874. The marine mammals of the north-west coast of North America, described and illustrated: Together with an account of the Amer- ican whale-fishery. San Francisco: John H. Carmany and Co., v + 319 pp. Schnell, G.D., M.E. Douglas, and D.J. Hough. 1986. Geographical variation in offshore spotted dolphins (Stenella attenuata) of the eastern tropical Pacific Ocean. Marine Mammal Science 2:186-213. Schwartz, M., A. Hohn, H. Bernard, S. Chivers, and K. Peltier. 1992. Stomach contents of beach cast ce- taceans collected along the San Diego County coast of California, 1972-1991. Southwest Fisheries Sci- ence Center Administrative Report LJ-92-18, 33 pp. Sherborn, C.D. 1935. Index Animalium, Sectio Secun- da, H, I, J, K, L. London: Oxford University Press. Smithers, R.H.N. 1983. The mammals of the Southern African Subregion. Pretoria, South Africa: Univer- sity of Pretoria, xxii + 736 pp. Sullivan, R.M., and W.J. Houck. 1979. Sightings and strandings of cetaceans from northern California. Journal of Mammalogy 60:828-833. True, F.W. 1889. Contributions to the natural history of the cetaceans, a review of the family Delphinidae. Bulletin of the United States National Museum 36: 191 pp + 45 pis. Van Beneden, P.J., and P. Gervais. 1880. Osteographie des Cetaces vivant et fossil comprenant la descrip- tion et I’iconographie du squelette et du systeme dentaire de ces animaux ainsi que des documents relatifs a leur histoire naturelle. Paris: Arthus Ber- trand, viii + 634 pp. + atlas. Wagner, J. A. 1846. {Revision of} J.C.D. von Schreber’s Die Sdugthiere, Vol. 7. Leipzig: Die Ruderfuesser und Fischzigthiere, vii + 427 pp. Wells, R.S., L.J. Hansen, A. Baldridge, T.P. Dohl, D.L. Kelly, and R. H. Defran. 1990. Northward exten- sion of the range of bottlenose dolphins along the California coast. In The bottlenose dolphin, ed. S. Leatherwood and R. R. Reeves, 421-431. San Diego: Academic Press, xviii + 653 pp. Wiley, E.O. 1981. Phylogenetics: The theory and prac- tice of phylogenetic systematics. New York: John Wiley and Sons, xv + 439 pp. Received 14 September 1992; accepted 3 December 1993. APPENDIX List of specimens examined by catalog number. Museum acronyms are as follows: LACM = Natural History Mu- seum of Los Angeles County, SDMNH = San Diego Mu- seum of Natural History, SWFC = Southwest Fisheries Science Center. Three specimens are not identifiable to species due to a lack of sufficient accompanying data, specimen damage, or immaturity. Delphinus delphis (n = 263) LACM: 27088, 27090, 27099, 27100, 27401, 30860, 31491, 40127, 43471, 43474, 47147, 52320, 54065, 54066, 54068, 54070, 54071, 54073, 54074, 54075, 54179, 54180, 54454, 54461, 54558, 54565, 54568, 54619, 54621, 54622, 54625, 54626, 54627, 54628, 54629, 54631, 54632, 54633, 54634, 54635, 54636, 54640, 54641, 54642, 54643, 54644, 54732, 54736, 54738, 54741, 54743, 54744, 54745, 54746, 54747, 54748, 72181, 72278, 72279, 72280, 72281, 72282, 72283, 72287, 72288, 72293, 72299, 72333, 72334, 72335, 72336, 72337, 72338, 72339, 72340, 72341, 72342, 72343, 72344, 72345, 72346, 72347, 72348, 72349, 72350, 72351, 72352, 72353, 72354, 72355, 72356, 72357, 72358, 72359, 72360, 72361, 72362, 72363, 72364, 72365, 72366, 72367, 72368, 72369, 72370, 72371, 72372, 72373, 72374, 723 75, 72376, 72377, 72378, 72379, 72380, 72381, 72382, 72383, 72384, 72385, 72386, 72387, 72388, 72389, 72390, 72391, 72392, 72393, 72394, 72395, 72396, 72397, 72398, 72399, 72400, 72401, 72402, 72403, 72404, 72405, 72406, 72407, 72408, 72409, 72410, 72411, 72412, 72413, 72414, 72415, 72416, 72417, 72418, 72419, 72420, 72421, 72422, 72423, 72425, 72426, 72428, 72454, 72468, 72495, 72496, 72497, 72498, 72503, 72505, 72543, 72587, 84007, 84039, 84041, 84042, 84045, 84046, 84050, 84054, 84067, 84074, 84078, 84090, 84094, 84096, 84105, 84106, 84108, 84109, 84118, 84129, 84131, 84132, 84134, 84136, 84137, 84138, 84139, 84140, 84143, 84155, 84170, 84172, 84173, 84177, 84178, 84181, 84196, 84197, 84199, 84207, 84208, 84209, 84216, 84225, 84226, 84227, 84229, 84230, 84231, 84232, 84255, 84257, 84261, 84279, 84280, 84282, 84283, 88904; SDMNH: 961, 2591, 19144, 19145, 20140, 20141, 20142, 21204, 21205, 21206, 21207, 21208, 21209, 21210, 21211, 21266, 21269, 22841, 22865, 23017, 23018, 23024, 23582, 23741, 23743, 23744, 23745, 23810; SWFC: 39, 40, 43, 44, 45, 48. Delphinus capensis (n = 54) LACM: 54067, 54463, 54618, 54735, 72284, 72285, 72286, 72289, 72424, 72427, 72429, 72430, 72469, 72494, 72499, 72500, 72502, 72544, 72593, 72595, 84009, 84021, 84040, 84071, 84077, 84083, 84091, 84092, 84100, 84121, 84125, 84127, 84130, 84135, 84163, 84183, 84184, 84185, 84220, 84221, 84223, 84228, 84233, 84236, 84239, 84240, 84241, 84254, 84256, 84258, 84278, 84281; SWFC: 39, 43. Delphinus sp. LACM: 72501, 84038, 84171. Contributions in Science, Number 442 Heyning and Perrin: Common Dolphins, Genus DelphinusM 35 >fu & Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Number 443 13 June 1994 t Contributions in Science Revision of Phanaeus Macleay, a New World Genus of Scarabaeine Dung Beetles (CoLEOPTERA: SCARAB AEIDAE, SCARABAEINAE) Natural History Museum of Los Angeles County Serial Publications of THE Natural History Museum of Los Angeles County Scientific Publications Committee Craig C. Black, Museum Director Daniel M. Cohen Kirk Fitzhugh John M. Harris, Committee Chairman Charles L. Hogue Edward C. Wilson Richard C. Hink, Managing Editor Robin A. Simpson, Head of Publications The scientific publications of the Natural History Mu- seum of Los Angeles County have been issued at irregular intervals in three major series; the issues in each series are numbered individually, and numbers run consecutively, regardless of the subject matter. • Contributions in Science, a miscellaneous series of tech- nical papers describing original research in the life and earth sciences. • Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth sci- ences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now published by the Museum in Contributions in Science. • Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through the Museum Book Shop. A catalog is available on request. The Museum also publishes Technical Reports, a mis- cellaneous series containing information relative to schol- arly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum’s Scientific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Tech- nical Reports may be obtained from the relevant Section of the Museum. Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Printed at Allen Press, Inc., Lawrence, Kansas ISSN 0459-8113 Revision of Phanaeus Macleay, a New World Genus of Scarabaeine Dung Beetles (CoLEOPTERA: SCARABAEIDAE, SCARABAEINAE) W. D. Edmonds1 CONTENTS ABSTRACT 3 RESUMEN 3 INTRODUCTION 3 HISTORY OF THE GENUS PHANAEUS 3 PRESENT CLASSIFICATION 4 Taxonomic Characters 4 Primary Type Material 5 Taxonomic Summary 8 Phanaeus Macleay, 1819 8 Invalid Names Assigned to Phanaeus Macleay 8 BIOLOGY, GEOGRAPHY, AND EVOLUTION OF PHANAEUS 9 South American Patterns 12 Middle American Patterns 13 North American Patterns 13 KEY TO THE PHANAEINE GENERA 17 SYSTEM ATICS OF THE GENUS PHANAEUS MACLEAY 17 Key to the Subgenera of Phanaeus 18 Notiophanaeus, New Subgenus 18 Key to the Species Groups of Notiophanaeus 19 The Splendidulus Group 19 Key to the Species of the Splendidulus Group 19 Phanaeus ( Notiophanaeus ) splendidulus (Fabricius) 22 Phanaeus ( Notiophanaeus ) dejeani Harold 23 Phanaeus ( Notiophanaeus ) haroldi Kirsch 24 Phanaeus ( Notiophanaeus ) melihaeus Blanchard 24 The Chalcomelas Group 25 Key to the Species of the Chalcomelas Group 27 Phanaeus ( Notiophanaeus ) chalcomelas (Perty) 28 Phanaeus ( Notiophanaeus ) achilles Boheman 30 Phanaeus ( Notiophanaeus ) camheforti Arnaud 31 Phanaeus ( Notiophanaeus ) meleagris Blanchard 31 The Bispinus Group 33 Key to the Species of the Bispinus Group 33 Phanaeus ( Notiophanaeus ) alvarengai Arnaud 34 Phanaeus ( Notiophanaeus ) bispinus Bates 35 The Palaeno Group 35 Key to the Species of the Palaeno Group 36 Phanaeus ( Notiophanaeus ) kirbyi Vigors 37 Phanaeus ( Notiophanaeus ) palaeno Blanchard 38 90007. Investigador Invitado, Instituto de Ecologia, Xalapa, Veracruz. Contributions in Science, Number 443, pp. 1-105 Natural Flistory Museum of Los Angeles County, 1994 The Endymion Group 39 Key to the Species of the Endymion Group 39 Phanaeus ( Notiophanaeus ) halffterorum Edmonds 41 Phanaeus ( Notiophanaeus ) endymion Harold 43 Phanaeus ( Notiophanaeus ) pyrois Bates 44 Subgenus Phanaeus Macleay . 46 Key to the Species Groups of Phanaeus, sen. str 46 The Hermes Group 47 Key to the Species of the Hermes Group 48 Phanaeus ( Phanaeus ) hermes Harold 48 Phanaeus ( Phanaeus ) prasinus Harold 49 The Tridens Group 49 Key to the Species and Subspecies of the Tridens Group 50 Phanaeus ( Phanaeus ) nimrod Harold 52 Phanaeus ( Phanaeus ) tridens Laporte-Castelnau 53 Phanaeus ( Phanaeus ) eximius Bates 54 Phanaeus (Phanaeus) furiosus Bates 54 Phanaeus ( Phanaeus ) daphnis Harold 55 The Triangularis Group 56 Key to the Species and Subspecies of the Triangularis Group 56 Phanaeus (Phanaeus) adonis Harold 57 Phanaeus (Phanaeus) triangularis (Say) 58 The Mexicanus Group 60 Key to the Species and Subspecies of the Mexicanus Group 60 Phanaeus (Phanaeus) flohri Nevinson 63 Phanaeus (Phanaeus) demon Laporte-Castelnau 65 Phanaeus (Phanaeus) scutifer Bates 67 Phanaeus ( Phanaeus ) lunaris Taschenberg 67 Phanaeus (Phanaeus) wagneri Harold 68 Phanaeus (Phanaeus) mexicanus Harold 69 Phanaeus (Phanaeus) amithaon Harold 69 The Beltianus Group 71 Key to the Species of the Beltianus Group 71 Phanaeus (Phanaeus) beltianus Bates 72 Phanaeus (Phanaeus) howdeni Arnaud 73 Phanaeus (Phanaeus) sallei Harold 73 The Amethystinus Group 74 Key to the Species and Subspecies of the Amethystinus Group 74 Phanaeus (Phanaeus) melampus Harold 74 Phanaeus (Phanaeus) amethystinus Harold 77 The Quadridens Group 79 Key to the Species of the Quadridens Group 79 Phanaeus (Phanaeus) palliatus Sturm . 79 Phanaeus (Phanaeus) damocles Harold 81 Phanaeus (Phanaeus) quadridens (Say) 82 The V index Group 83 Key to the Species of the V index Group 83 Phanaeus (Phanaeus) igneus Macleay 85 Phanaeus (Phanaeus) difformis LeConte 88 Phanaeus (Phanaeus) vindex Macleay 89 Incertae Sedis 91 Phanaeus labreae (Pierce) 91 Phanaeus (?) antiquus Horn 92 ACKNOWLEDGMENTS 92 LITERATURE CITED 93 APPENDIX 99 2 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ABSTRACT. This study is a revision of the genus Phanaeus as defined by Edmonds (1972). The intro- duction includes a history of the genus, a key separating it from related genera, a discussion of taxonomic characters and primary type material, a summary of the biology and geography of the genus, and con- clusions about its evolution. The systematic section treats 48 species and subspecies in two subgenera, one of which, Notiophanaeus, is new. Notiophanaeus comprises five species groups embracing 15 mostly South American species. Phanaeus, sen. str., includes 31 species and subspecies arranged in eight species groups. Keys separate subgenera, species groups, species, and subspecies. Phanaeus labreae (Pierce) and P. antiquus Horn, known from fossil deposits, are considered incertae sedis. One new subspecies is described, P. (P.) triangularis texensis. The following new combinations result from reductions in rank from species to subspecies: P. (P.) amethystinus guatemalensis Harold, P. (P.) tridens pseudofurcosus Balthasar, and P. (P.) wagneri pilatei Harold. Neotypes are designated for the following taxa: Copris triangularis Say, C. quadridens Say, Phanaeus tridens Laporte-Castelnau, P. demon Laporte-Castelnau, P. laevicollis LaPorte-Castelnau (= P. kirbyi Vigors), P. difformis LeConte, P. igneus Macleay, and P. vindex Macleay. Lectotypes are designated for the following (valid name, where appro- priate, appears parenthetically preceded by =): Scarabaeus carnifex Linnaeus 1767, not 1758 (= P. vindex Macleay); P. minos Erichson (= P. meleagris Blanchard); P. bogotensis Kirsch (= P. hermes Harold); P. torrens LeConte (= P. triangularis [Say]); P. pyrois Bates; P. furiosus Bates; P. eximius Bates; P. beltianus Bates; P. scutifer Bates; P. excelsus Bates (= P. demon Laporte-Castelnau); P. obliquans Bates (= P. demon Laporte-Castelnau); P. scintillans Bates (= P. mexicanus Harold); and P. tepanensis Bates (= P. amethystinus guatemalensis Harold). The following new synonymy is proposed in Notiophanaeus (synonym precedes valid name): P. kirbyi truncaticornis Olsoufieff = P. kirbyi Vigors, P. blanchardi Olsoufieff = P. pyrois Bates, and P. funereus Balthasar = P. pyrois Bates. The following are new synonyms in Phanaeus, sen. str.: P. bogotensis Kirsch = P. hermes Harold, P. lugens Nevinson = P. prasinus Harold, P. furcosus Lelsche = P. furiosus Bates, P. herbeus Bates = P. daphnis Harold, P. coerulus Bates = P. daphnis Harold, P. tricornis Olsoufieff = P. daphnis Harold, P. niger Olsoufieff = P. triangularis (Say), P. obliquans Bates = P. demon Laporte- Castelnau, P. obliquans Bates = P. demon Laporte-Castelnau, P. excelsus Bates = P. demon Laporte- Castelnau, P. scintillans Bates = P. mexicanus Harold, P. martinezi Halffter = P. amethystinus Harold, P. quadridens borealis Olsoufieff = P. quadridens (Say), P. vindex rubervirens Robinson = P. vindex Macleay, P. floridanus Olsoufieff = P. igneus Macleay, and P. difformis magnificens Robinson = P. difformis LeConte. A list of all known available names and a previously established synonymy are included in the intro- duction. Geographical distribution data for all species and subspecies are contained in the appendix. RESUMEN. Este trabajo es una revision del genero Phanaeus Macleay, segiin Edmonds (1972). La parte introductoria consiste en una relacion de la historia taxonomica del genero, una clave que lo separa de generos vecinos, un resumen de la biologia y geografia del grupo, y una serie de conclusiones acerca de su evolucion. La seccion sistematica trata de 48 especies y subespecies colocadas en dos subgeneros, de los cuales Notiophanaeus es nuevo. Notiophanaeus comprende cinco grupos de especies, que, en su mayoria, son sudamericanas. Phanaeus, sen. str., consta de 31 especies y subespecies, principalmente mexicanas y estadounidenses, clasificadas en ocho grupos. Se presentan claves para la identificacion de subgeneros, grupos de especies, especies y subespecies. Phanaeus labreae (Pierce) y P. antiquus Horn son especies supuestamente extintas de incertae sedis. Los datos de distribucion geografica comprenden el apendice. Se describe una nueva subespecie, P. (P.) triangularis texensis. El nivel taxonomico de tres taxa, anteriormente consideradas como especies, se baja a subespecie. Se designan neotipos por ocho especies y lectotipos por otras trece. Se sehalan 19 casos de nueva sinonimia. INTRODUCTION The present paper is a continuation of my previous work on the phanaeine dung beetles (Edmonds, 1972). One of the more interesting problems I left open at that time was the systematics of the genus Phanaeus. These are very well known beetles, but until now the only comprehensive study of species has been that of Gregor d’OlsoufiefF (1924). The systematic portion of the present study is based on the examination of more than 17,000 specimens, including most primary types, from over 75 insti- tutional and private collections. It is preceded by a brief historical account, a summary of the taxon- omy of the genus, and a discussion of its biology, geography, and evolution. HISTORY OF THE GENUS PHANAEUS William Sharp Macleay was the first serious student of what today is generally known as the family Scarabaeidae. In 1819 he published the first part of Horae Entomologicae, an elaborate reexamination of Linnaeus’ genus Scarabaeus (as defined in the twelfth edition of Sy sterna Naturae, 1767) within the somewhat broader taxonomic context of P. A. Latreille’s “Lamellicornes” (1825, and elsewhere). While Macleay was primarily concerned with the Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 3 “higher” classification of Lamellicornes by estab- lishing five-part “circular relationships” among families he recognized, he was a strong believer in the “naturalness” of the genus. Macleay (1819) thus included the original descriptions of a number of new scarabaeid genera, one of which is Phanaeus. Macleay defined Phanaeus primarily on the basis of the shape of the antennal lamellae and arranged the ten species he recognized according to five “types.” His “Typus 5,” which included carnifex, sensu Linnaeus, 1767, vindex as a variation of car- nifex, and igneus as species inquirenda, is the least inclusive direct antecedent of Phanaeus as defined here. Macleay’s concept of Phanaeus endured almost intact until d’OlsoufiefPs revision of the “Tribu Phanaeides” in 1924. The only significant change before then was Laporte-Castelnau’s (1840) crea- tion of Oxysternon, which received, among others, those species comprising Macleay’s “Typus 4.” During the latter part of the nineteenth century, however, many new species were described. Of the 46 species and subspecies recognized here, 34 were named between 1830 and 1890 as well as many others later placed in synonymy. Especially impor- tant in expanding the size of Phanaeus and related genera {sensu Edmonds, 1972) were the works of Harold (1863, 1868a, b, 1871b, 1875a, b) and Bates (1887, 1889). The first key to species in a modern format appeared in Harold’s (1863) review of the Mexican species of Phanaeus. The genus was cat- aloged by Gemmingerand Harold (1869), Nevinson (1892a), Gillet (1911b), and, most recently, Black- welder (1944). Regional reviews postdating d’Ol- soufieff’s revision include Pessoa (1934; Brazil), Islas (1942; Mexico), and Robinson (1948; United States). Gregor d’Olsoufieff’s (1924) revision largely con- served the taxonomic limits of Phanaeus (excluding Oxysternon) originally established by Macleay, al- though the number of species had increased more than nine-fold in the preceding 105 years. However, he reorganized what had become a highly diverse group into five subgenera: Sulcophanaeus, Mega- phanaeus, Coprophanaeus, Metallophanaeus, and Phanaeus, sen. str. {Scarahaeus mimas L., included in Macleay’s “Typus 2,” was moved to the new genus T aurocopris, now known as Diabroctis. ) The first of these subgenera was the descendent of Mac- leay’s “Typus 3” and remainder of “Typus 2”; the second and third descended from “Typus 1” (nei- ther species assigned to Metallophanaeus was known to Macleay). Phanaeus, sen. str., preserved Macleay’s “Typus 5” and was the initial step in restricting the taxonomic scope of the name Phan- aeus. My revision (Edmonds, 1972) of the supraspecific taxonomy of the phanaeines resulted in the splitting of Phanaeus into three genera based largely upon the subgenera created by Olsoufieff (1924). Ac- cordingly, Phanaeus became one of nine genera (and eight subgenera) comprising the phanaeine ge- nus group (not recognized by me as a tribe or sub- tribe). The sense of Phanaeus used here is the same as that I defined in 1972. PRESENT CLASSIFICATION TAXONOMIC CHARACTERS The characters used in the keys, descriptions, and elsewhere employ the terminology established by Edmonds (1972). Easily assessed morphological characters separate most species of the genus. How- ever, the observation I made in 1972 still holds true in spite of the introduction of many new characters here. The incidence of irregular distribution of character states, and the continuous variation of characters whose extremes are easily definable, make the taxonomy of some groups difficult. Most of these difficulties occur in Phanaeus, sen. str., and are discussed in the taxonomic section. Pronotal sculpturing is widely used here to de- scribe supraspecific taxa. Sculpturing in the genus varies from a smooth condition virtually devoid of punctures or other relief (Fig. 166) to extreme ru- gosity (Fig. 339). In 1972 I recognized only two, very generalized states of pronotal sculpturing: “smooth-punctate” and “roughened” (or “ru- gose”). Present purposes, however, require that these descriptors be refined. The smooth-punctate con- dition is characteristic of the subgenus Notiophan- aeus. However, in certain instances described in the introduction to that subgenus, it can be ob- scured by superposition of various kinds of rough- ening. In Phanaeus, sen. str., the pronotum is al- ways rugose to some degree, often entirely so. The kind of roughening in this subgenus falls into two general categories: granulation and granulorugosi- ty. In the former case (Figs. 224, 236, 253), sculp- turing consists of discrete granules, the size and density of which can vary among species and species groups. An area described as granulate for a given taxon will bear granules of usually uniform size that only infrequently coalesce into larger entities. Granulorugosity, on the other hand, consists of a heterogenous mixture of asperities of various shapes and sizes producing a highly fractured surface (Figs. 274, 310, 319, 320). The overall texture produced by granulorugosity varies greatly and continuously such as to preclude any objective definition of sub- states. Thus, granulorugose is an imprecise term describing a roughened texture that is not clearly granulate. Phanaeus species are very efficient diggers, and those structures used directly for digging are highly subject to wear. The effects of wear can lead to errors in assessing the shapes of the anterior margin of the clypeus and front tibiae. In very extreme cases, wear can alter the nature of surface sculp- turing (especially shagreening) and coloration. Most Phanaeus are brilliantly colored (Figs. 1- 107). Intraspecific variation in coloration and color pattern is the rule and can be very striking. More- over, individuals themselves are seldom uniformly 4 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus colored. While its taxonomic usefulness is limited, coloration is an important aspect of the morphol- ogy of these beetles and is addressed in all species descriptions. Assessment of coloration should be made under low magnification ( x 5-10) with a high- intensity, broad-spectrum light source, such as halogen. A summary of known distributional information (geographical, ecological, altitudinal, and temporal) is included in the treatment of each species. De- tailed information is contained in the appendix. Geographical data can be critical in identifying cer- tain species, especially those in which females and smaller males are indistinguishable morphological- ly- Traditionally, much taxonomic emphasis has been placed on the secondary sexual features of Phan- aeus, especially the shape of the cephalic horn and pronotum of the male. Male armature, however, is subject to often extreme allometric intrapopulation variation (as in Fig. 214) as well as to interpopu- lational differences in shape (as in Figs. 275-280). In general, taxonomically reliable male sexual fea- tures are characteristic only of large, well-devel- oped (“major”) individuals, although there can be some striking exceptions (see, e.g., the case of Phanaeus demon). Secondary sexual features, par- ticularly the shape of the pronotum, are much more useful among females. A potential source of morphological characters not explored exhaustively during this study is the structure of the genitalia. Zunino (1971, 1978, 1979, 1983) relied successfully on the bursa copulatrix of the female for reliable taxonomic characters in Onthophagus. I attempted, without success, to find similar characters applicable to the definition of species only in the particularly difficult cases of the mexicanus and tridens species groups of Phanaeus, sen. str. Proper assessment of many taxonomic characters used here requires viewing under magnification. Where appropriate, the minimum augmentation recommended is noted. PRIMARY TYPE MATERIAL I have examined primary type material for the great majority of species group names considered in this study. The only valid name used for which no type was examined or designated is Phanaeus lunaris Taschenberg. In several cases I have found it necessary to des- ignate neotypes and lectotypes; particulars con- cerning these specimens appear in the comments section of the species in question. Those names for which neotypes are designated are as follows (de- scribed as Phanaeus unless indicated otherwise; valid name, where appropriate, indicated parenthetical- ly): Copris triangularis Say, Copris quadridens Say, tridens Laporte-Castelnau, demon Laporte-Castel- nau, laevicollis Laporte-Castelnau (= kirbyi Vig- ors), difformis Leconte, igneus Macleay, and vin- dex Macleay. Lectotypes are designated in the following cases: Scarabaeus carnifex Linnaeus, 1767; minos Erichson (= meleagris Blanchard); bo- gotensis Kirsch (= hermes Harold); pyrois Bates; furiosus Bates; eximius Bates; beltianus Bates; scu- tifer Bates; excelsus Bates (= demon Laporte-Cas- telnau); obliquans Bates (= demon Laporte-Cas- telnau); scintillans Bates (= mexicanus Harold); tepanensis Bates (= amethystinus guatemalensis Harold); and torrens LeConte (= triangularis Say). The Thomas Say Collection has been long pre- sumed to have been destroyed in the 1830s. Since Say is known to have worked with the collections of John E. LeConte (which became the nucleus of the collection of his son, John L. LeConte) and T. W. Harris (A. Newton, pers. comm.), it has been customary to select necessary neotypes of Say spe- cies from the Harris or LeConte Collection, both of which are housed in the Museum of Compar- ative Zoology, Harvard University. In neither case (■ Copris triangularis nor C. quadridens) for which a neotype of a Say species was necessary did ac- ceptable specimens exist in either of these two col- lections. Over the period of roughly 1800-1890, the Mac- leays of Sydney, Australia (Alexander; his son, Wil- liam Sharp; and nephew, William John), together amassed an enormous insect collection of great his- torical importance (Horning, 1984, and references therein). Their combined collections form the Mac- leay Museum at the University of Sydney. I was unable to locate types of either vindex or igneus in material from the Macleay Collection, including that part transferred to the Australian National In- sect Collection in Canberra in 1969 (Britton and Stanbury, 1981). The whereabouts of type material for species described by Francois L. N. de C. Laporte (who later in life received the French royal title, Comte de Castelnau) has long been a mystery (see, e.g., Paulian, 1976). Many phanaeines ( Phanaeus and related genera) exist in the Castelnau Collection at the National Museum of Victoria in Melbourne, Australia, where Laporte served as Counsel General of France. Specimens bearing labels in Laporte-Cas- telnau’s hand also appear in the Godfrey Howitt Collection, also in Melbourne (A. Neboiss, pers. comm.). Neither collection includes designated type material, nor does either include specimens that can reasonably be taken as typical. Laporte wrote under his surname as well as his title, Comte de Castelnau. As a result, authorship of Laporte species appears in the literature alternately as Laporte or Castelnau. For bibliographic purposes, I have elected to refer to him as Laporte-Castelnau. Arnaud (1982a) is responsible for the designation of many phanaeine lectotypes in the collections of the Museum National d’Histoire Naturelle in Paris. In the introduction to his paper, he states, in the case of species described by H. W. Bates (1887, 1889), “En effet, les seuls specimens portant la de- signation manuscrite de l’espece par la main de l’au- Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 5 [1] Phanaeus splendidulus d [2] Phanaeus splendidulus d [3] Phanaeus splendidulus ? [4] Phanaeus dejeani d [5] Phanaeus dejeani ? [6] Phanaeus melibaeus [7] Phanaeus melibaeus [8] Phanaeus haroldi [9] Phanaeus haroldi [10] Phanaeus palaeno d [11] Phanaeus palaeno d gp® mmm [12] Phanaeus palaeno [13] Phanaeus ^ kirbyi [14] Phanaeus kirbyi [15] Phanaeus bispinus d [ 1 9] Phanaeus pyrois d [16] Phanaeus bispinus ? [20] Phanaeus pyrois ? [17] Phanaeus endymion [2 1 ] Phanaeus halffterorum [18] Phanaeus endymion [22] Phanaeus halffterorum ? Figures 1 - 22. Shown natural size. 6 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus [27] Phanaeus meleagris [32] Phanaeus hermes [28] Phanaeus meleagris 9 [33] Phanaeus hermes 9 [29] Phanaeus meleagris d [34] Phanaeus prasinus d [35] Phanaeus prasinus 1000 m) forests of the coastal mountains of ex- treme southeastern Brazil in Sao Paulo, Rio de Ja- neiro, Minas Gerais, and Espirito Santo. Presumed coprophagous. Collection dates: November-Feb- ruary. COMMENTS. The coloration of dejeani is strik- ing and unlike that of any other Phanaeus. The intense, deep blue of the elytra, in particular, is unapproached in other species. Not pointed out elsewhere is the fact that, unlike the female of splendidulus, which bears a single, large black spot on the pronotal disk, the female of dejeani has a pair of small, black spots immediately behind the anteromedian concavity. The distribution of dejeani appears to be over- lapped completely by the northern part of that of splendidulus. Reliable data, while scarce, suggest that it usually occurs at higher elevations (>1000 m). Phanaeus ( Notiophanaeus ) haroldi Kirsch Figures 8, 9, 111, 112, 118, 120, 122, 124, 128, 131, 135, 136 Phanaeus haroldi Kirsch, 1871:358 Type: Female holotype (“Bogota”), Staatliches Museum fiir Tierkunde, Dresden. OTHER REFERENCES. Nevinson, 1892a; Gil- let, 1911b; Olsoufieff, 1924; Paulian, 1935; Pessoa, 1934; Blackwelder, 1944; Gacharna, 1951; Vulcano and Pereira, 1967; Barrera, 1969; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 8, 9). — Pronotum, posterior portion of head weakly to moderately shining dark green; pronotal spines of both sexes black. Elytra dull, dark green or, more often, almost black with weak greenish or bluish luster. Pygidium dark, weakly shining green; venter somber, with weak green or blue-green reflections. Pronotum. — Basal fossae conspicuous. Anterolateral angle drawn out into laterally projecting tooth, posterior to which cir- cumnotal ridge strongly sinuous (Fig. 122); anterior portion of ridge (that behind paraocular areas) straight and approximately perpendicular to lon- gitudinal axis of body. Puncturing (x30) widely spaced and often visible only posterolaterally in male; more evenly distributed in female (Fig. 118). Metasternum. — -Anterior prominence (seen in pro- file; Fig. 128) rounded, sometimes almost lobate, (seen from below) compressed laterally, keeled lon- gitudinally. Front Tibiae (Fig. 124). — Quadriden- tate; basal (fourth) tooth sometimes poorly devel- oped; first three teeth carinate longitudinally. Apex of spur truncated, inner angle acute, strongly pro- duced mesally. Secondary Sexual Characters. — Male: In larger individuals (Fig. Ill), cephalic horn slender, erect, not curving posteriorly; reduced to small, triangular tubercle in smallest individuals. In large individuals, roughly triangular portion of pronotal disk concave; posterolateral angles pro- duced as pair of apically convergent, long, antero- dorsally projecting spines; in smaller individuals, disk convex, spines reduced to pair of acute, lat- erally compressed denticles near anterior margin. Female: Cephalic carina bowed anteriorly, strongly trituberculate (Fig. 131), less so in smaller individ- uals. Anterior one-half of disk (Figs. 112, 120) bear- ing concavity flanked by pair of elongate, antero- dorsally directed, acute spines, reduced to acute denticles in small individuals. Specimens Exam- ined.— 73 males, 66 females (length 14-20 mm; width 9-13 mm). DIAGNOSTIC REMARKS. The secondary sex- ual features of this species distinguish it from all other Phanaeus. DISTRIBUTION (Fig. 136; appendix). Forests of the eastern slopes and foothill valleys of the Andes (“Yungas”) from Colombia to northern Peru, and isolated localities in Venezuela. 500-1000 m (estimated). Coprophagous. Collection dates: all year. COMMENTS. It is surprising to me that this large, striking Phanaeus is so poorly known. It ap- pears to be a resident of the higher forests of the northern portion of the eastern Andean slopes (“Yungas”) from which it can descend to lowland areas. I have seen two specimens collected in the Orinoco River basin (solid squares in Fig. 136) near Ciudad Bolivar, where they probably inhabit gallery forest. This lowland population is very far removed from the main range and may be a relictual. Several specimens examined were labeled, most likely in error, “Panama” (one, “Chiriqui”); Howden and Young (1981) do not report haroldi from Panama. Its closest relative, melibaeus, occurs far to the southeast. Phanaeus ( Notiophanaeus ) melibaeus Blanchard Figures 6, 7, 113, 114, 119, 123, 125, 132, 136 Phanaeus melibaeus Blanchard, 1843:176 Type: Male holotype (“Chiquitos”), Museum National d’Histoire Naturelle, Paris. Phanaeus sculpturatus Olsoufieff, 1924:87 (Pereira and Martinez, 1956:236) 24 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Type: Female holotype (“Chiquitos”), Museum National d’Histoire Naturelle, Paris. NOMENCLATURAL REMARKS. Blanchard s description was based on a single, well-developed male. Olsoufieff based sculpturatus on a single spec- imen which he rightly suspected was the female of melibaeus. OTHER REFERENCES. Lacordaire, 1856; Gemmingerand Harold, 1869; Harold, 1869; Nev- inson, 1892a; Gillet, 1911b; Olsoufieff, 1924; Pessoa, 1934; Pessoa and Lane, 1937; Blackwelder, 1944; Pereira and Martinez, 1960; Edmonds, 1972; Arnaud, 1982a. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 6, 7). — Pronotum, posterior portion of head shining green with weak to strong yellow or copper reflections, particularly on posterome- dian portion of notum. Elytra shining green to blue- green, interstriae strongly shining midlongitudinally and only weakly so along striae. Pygidium shining green to yellow-green. Venter dark, with shining green on abdominal sterna and legs. Pronotum. — Basal fossae conspicuous. Anterolateral angles den- tate, only weakly so in female; circumnotal ridge sinuous behind anterolateral angle in male (Fig. 123), almost straight in female; anterolateral portion of ridge (behind paraocular areas) slightly sinuous. Metasternum. — Anterior prominence (seen in pro- file) rounded, (seen from below) compressed lat- erally, keeled longitudinally. Front Tibiae (Fig. 125).—— Strongly quadridentate, fourth (basal) tooth distinct even after severe wear; first three teeth car- inate longitudinally. Spur truncated apically, inner angle acute, strongly produced mesally. Secondary Sexual Characters.— Male: In large individuals (Fig. 113), cephalic horn, seen laterally, slender, except for slight posterior flexion near base, almost erect; that of smallest specimens reduced to small, conical tubercle. Pronotal disk bearing pair of prominent, round concavities in front of raised area along pos- terior margin; sides of disk drawn out as pair of tapering, mesally curved, dorsally directed spines; in smaller individuals, pronotal spines reduced to slightly upturned, blade-like processes. Female: Ce- phalic carina trituberculate; seen from above, al- most straight; in larger individuals, carina strongly raised with highly accentuated, acute tubercles, its outline resembling scalloped margin of crown (Fig. 132). Pronotum (Figs. 114, 119) convex, with round, anteromedian concavity preceded by three, strong, anteriorly directed denticles, middle one of which quadrate, lateral of which acute; surface in front of denticles almost vertical, seen anteriorly, shal- lowly concave between two weak ridges directed outwardly from points on the notum above the eyes. Specimens Examined. — 19 males, 14 females (length 13-18 mm; width 8-11 mm). DIAGNOSTIC REMARKS. This species is sim- ilar to baroldi; the two species are easily distin- guished by coloration and secondary sexual char- acters. DISTRIBUTION (Fig. 136; appendix). Southern Amazon Basin in Brazil. Coprophagous. Collection dates: October-February. COMMENTS. This species is uncommon, even in large collections. Known collection localities are all along the upper reaches of major tributaries of the Amazon River and doubtlessly reflect the routes followed by collectors. P. melibaeus is unknown from central, “open” Amazonia and may be a spe- cies that prefers intermediate habitats rather than fully developed hylea. The distribution of this spe- cies is separated from those of splendidulus and dejeani to the east by the extensive “campos cerra- dos” of the Brazilian Highlands and from that of haroldi to the northwest by uninterrupted Ama- zonian forest. I have not seen specimens from Bo- livia other than the holotypes, both of which are labeled “Chiquitos.” The known distribution of melibaeus and its scarcity suggest peculiar ecolog- ical requirements, but I have no data that indicate what these could be. The Chalcomelas Group DIAGNOSIS. [1] Anterior margin of clypeus weakly bidentate; [2] clypeal process (seen from front) rounded, narrow, almost toothlike (Fig. 156); [3] cephalic carina of female at most only barely trituberculate (Fig. 149), usually appearing simple (except achilles, Fig. 148); [4] anterolateral portions of pronotum punctate, punctures either [a] (Fig. 147) very broad, shallow, often confluent, each with minute, central shining spot (punctures sometimes almost effaced and visible only from a low angle) or [b] (in achilles, Fig. 146) smaller, deeper, and most distinct opposite eyes and around lateral fos- sae; [5] pronotal disk (and sometimes also sides) with few to many irregularly shaped, often conflu- ent, flat, blister-like black rugosities (most strongly developed in achilles) giving surface a peppered or mottled appearance (Figs. 137-145); [6] pronotum of male flattened dorsally, very evenly so except in achilles, with distinct posterolateral angles (Figs. 137, 139, 141, 143, 144); [7] pronotum of female evenly convex (Figs. 138, 140, 142, 145); [8] basal pronotal fossae present or absent; [9] front tibiae (Figs. 152, 153) tridentate (fourth [basal] tooth sug- gested in some females); [10] elytral striae fine ( x 30), each bearing 15-20 conspicuous, variable-sized, dark, shallow punctures easily visible to unaided eye (Figs. 150, 151); interstriae flat; [11] dorsal col- oration usually somber (Figs. 23-29) (bright red or green in achilles, Figs. 30, 31); Amazonian rain for- ests (except achilles) (Fig. 162). The chalcomelas group brings together four South American species distinguished from other Notio- phanaeus by their pronotal sculpturing (4 and 5 above) and conspicuous, fossa-like punctures of the elytral striae (Figs. 150, 151). Three species, chal- comelas, meleagris, and cambeforti, inhabit rain forests; their somber coloration camouflages them against a background of forest floor litter. The fourth Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 25 Figures 137-145. Phanaeus ( Notiophanaeus ) chalcomelas group, oblique lateral views (137, P. chalcomelas, male; 138, same, female; 139, P. cambeforti, male; 140, same, female; 141, P. achilles, male; 142, same, female; 143, 144, P. meleagris, male; 145, same, female. species, achilles, inhabits the desert scrub region of southwestern Ecuador and northwestern Peru along the Pacific coast; it is isolated taxonomically within the group (see “Comments” under achilles). The pronotal sculpturing of this group is remi- niscent of that of most Phanaeus, sen. str., which is most closely approached in achilles. In this con- text, the chalcomelas group is an annectant taxon between the two subgenera. There exists a contin- uum of variation among the species of this group in the degree of roughness of the pronotal surface, particularly the disk. In chalcomelas and cambe- forti (Figs. 137-140), the rugosities are finer and less dense. In meleagris (Figs. 143-145), they are coarser and more densely placed, but, as in the two former species, they retain a flat, blister-like appearance. In achilles, however (Figs. 141, 142), the rugosities are more clearly defined (“crisper”), denser, and more widely distributed on the disk and sides such that the sculpturing of the pronotum closely resem- bles that of certain Phanaeus, sen. str. In all four species, however, the sides of the pronotum, which may bear rugosities like those on the disk, retain at least some distinct puncturing. Moreover, except in some male achilles and meleagris, there is at least limited puncturing also on the central area of the disk. The large, fossate punctures of the elytral striae are seen otherwise only in bispinus (Fig. 169). The striae themselves are extremely fine, superficial lines; they are not carinulate, as I wrongly stated in 1972. P. chalcomelas and cambeforti bear a strong re- semblance to each other, particularly in size, shape, and color. Indeed, cambeforti escaped detection until 1982, and the two species are often commin- gled in collections. Both occur in the Amazon Basin and, as far as I know, cambeforti is always found along with chalcomelas (the converse, however, is not true). Certain color forms of meleagris resemble these species, and all three can occur sympatrically (e.g., the environs of Villavicencio, Colombia). How these three species coexist ecologically would be the subject of an interesting field study. P. achilles differs morphologically, geographi- cally, and ecologically from the other members of the group; it is the only xeric adapted species of 26 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Figures 146-151. Phanaeus ( Notiophanaeus ) chalcomelas group. (146, anterolateral view pronotum of P. achilles, female; 147, same, P. chalcomelas, female; 148, dorsal view head of P. achilles, female; 149, same, P. chalcomelas, female; 150, dorsal view left elytron of P. achilles, female; 151, same, P. chalcomelas, female). Notiophanaeus. At first glance, it strongly resem- bles members of the mexicanus group of Phan- aeus, sen. str., none of which possesses the large, fossate punctures of the elytral striae nor an acu- minate anterior angle of the metasternum. KEY TO THE SPECIES OF THE CHALCOMELAS GROUP la. Anteromedian angle of metasternum seen from below (Fig. 154) capped by raised arrowhead or V-shaped swelling; sides of swelling visible laterally as distinct ridges. Paraocular ridge al- most always distinct and extending from pos- terior margin of paraocular area to point even with middle of lateral margin of eye. Male: Cephalic horn of large individuals as in Figures 159, 160, abruptly narrowed at about mid- length. Female: Pronotum (Fig. 138) with shal- low, anteromedian concavity surrounded by four small tubercles, anterior one of which is largest, occasionally toothlike. Dorsum dull, dark, usually olive brown or olive green (Figs. 23, 24). Length 12-18 mm. Tropical rain for- ests of South America from Guiana to Bolivia (Fig. 162) Phanaeus (N.) chalcomelas (Perty) b. Anteromedian angle of metasternum not cap- ped; this angle smooth and drawn out anteri- orly as small, slightly upturned acute tooth best seen in profile (Fig. 155). Paraocular ridge pres- ent or absent. Male: Cephalic horn tapering evenly (Fig. 157); if widened basally (some meleagris. Fig. 158), narrowing not as abrupt as above. Female: Pronotum evenly convex, lacking distinct anteromedian concavity, bear- ing three small, rounded tubercles (almost ef- faced in achilles ) near anterior margin (Figs. 140, 142, 145). Color and distribution variable 2 Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 27 159 156 ' I Figures 152-161. Phanaeus ( Notiophanaeus ) cbalcomelas group. (152, P. chalcomelas, male, left tibia; 153, same, female; 154, P. chalcomelas, anterior end of metasternum [A = lateral view; B = ventral view]; 155, P. meleagris, same; 156, P. chalcomelas, anterior view clypeal process; 157, P. cambeforti, male, anterior view left side of head; 158, P. meleagris, same; 159-161, P. chalcomelas, same). 2a. Elytral interstriae distinctly, but sparsely punc- tured (xlO), irregularly wrinkled (Fig. 150). Pronotal punctures small, deep, intermingled with well-defined, black rugosities (Fig. 146). Pronotal disk of large males flattened, but not evenly so (Fig. 141); posterolateral angles, seen from above, rounded laterally such that disk appears nearly heart-shaped. Middle of ce- phalic carina of female thickened and raised, forming a distinct, isolated, anteriorly bowed ridge (Fig. 148). Paraocular ridge distinct. Dor- sum shining, coppery red or dark green (Figs. 30, 31). Desert scrub region of southwestern Ecuador and northern Peru Phanaeus (N.) achilles Boheman b. Elytral interstriae completely smooth, bearing no punctures or wrinkling (Fig. 151). Pronotal punctures (Fig. 147) large, very shallow, often confluent, each with a shining, central micro- spot (x40; pronotal puncturing often visible only when viewed from a low angle). Pronotal disk of large males evenly flattened, sides of posterolateral angles more or less straight such that disk, seen from above, appears almost tri- angular (Figs. 137, 139, 143, 144). Cephalic carina of female simple or only weakly tritu- berculate (Fig. 149). Paraocular ridge absent or indicated only by a slight swelling adjacent to eye. Dorsum dull, usually somber. Evergreen forests east of Andes 3 3a. Pronotal puncturing, while never strong, usu- ally distinct both on sides and disk; central microspots often very difficult to detect ( x 40), sometimes lacking. Basal pronotal fossae lack- ing or represented only by small punctures ( x 10). Dorsum olive green or olive brown; me- tallic highlights on pronotum and pygidium al- ways green. Smaller-sized, length rarely ex- ceeding 15 mm. Amazon Basin from Guiana to Colombia and Peru Phanaeus (N.) cambeforti Arnaud b. Pronotal puncturing very weakly defined and often appearing almost effaced; central shining microspots almost always distinct ( x 40) even if associated punctures are obsolete. Basal pronotal fossae always distinct. Dorsal color- ation variable: (a) including reddish or pinkish reflections, or green-brown tones infused with reddish highlights at least along circumnotal ridge, but often also on sides of disk and pro- notum (Figs. 27, 28); or (b) uniformly weakly lustrous, dark blue with an “oily” sheen (Fig. 29); or (c) uniformly dull, dark green or blue; if general dorsal coloration approaches olive green or olive brown, always associated with red- dish or pinkish highlights on pronotum and py- gidium. Farger-sized, length seldom less than 15 mm. Rain forests (“Yungas”) along eastern slopes of Andes from Venezuela to Bolivia Phanaeus (N.) meleagris Blanchard Phanaeus ( Notiophanaeus ) chalcomelas (Perty) Figures 23, 24, 137, 138, 147, 149, 151-154, 156, 159-162 Onitis chalcomelas Perty, 1830:40 28 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Type: Female holotype? (“Brasilia”), Zoolo- gische Staatssammlung, Munich (see “Com- ments”). Phanaeus chalcomelas (Perty) (Harold, 1859:198) OTHER REFERENCES. Bates, 1868; Gemmin- ger and Harold, 1869; Nevinson, 1892a; Gillet, 1911b; Olsoufieff, 1924; Pessoa, 1934; Paulian, 1935; Halffter and Matthews, 1966; Vulcano and Pereira, 1967; Barrera, 1969; Edmonds, 1972; Scherer, 1983. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 23, 24).— Dorsum dull olive green to olive brown with green highlights, occasionally coppery brown, rarely with strong green reflections on pronotum. Pronotal coloring broken by black rugosities, which produce mottled or peppered ap- pearance; extent of mottling varies from very little to covering most of disk. Pygidium green; venter with shining, dark green reflections on legs, pleura, and abdominal sterna. Head.— Paraocular ridge distinct. Pronotal Sculpturing .- — Low, blister-like rugosities usually fairly dense, covering most of disk (Figs. 137, 138) (very sparse to almost absent in male Guiana specimens); sides, anterolateral angles with few, if any, rugosities, densely punctate (Fig. 147); punctures broad, shallow, each with incon- spicuous central microspot. Pronotum. — Basal pronotal fossae present, punctiform. Metaster- num.— Anterior prominence capped by a raised swelling which, seen from below (Fig. 154B), shaped as an arrowhead or narrow V; sides of swelling, seen laterally (Fig. 154A), visible as distinct ridges. Secondary Sexual Characters.— -Male: Basal one- half of cephalic horn of large individuals (seen from front, Figs. 159, 160) greatly expanded laterally; apical one-half abruptly narrowed, tapering; basal expansion of horn progressively less pronounced in smaller specimens; horn reduced to small, conical tubercle in smallest individuals (Fig. 161). Pronotal disk of large individuals (Fig. 137) evenly flattened, sides more-or-less straight, posterolateral angles (seen from above) projecting beyond lateral margin of pronotum such that disk almost triangular; in smallest individuals, posterolateral angles reduced to small, elongate tubercles near middle of disk, flattened area restricted to area between and in front of these tubercles. Female: Pronotum (Fig. 138) with small, oval, anteromedian concavity surround- ed by four tubercles; anterior tubercle largest, toothlike in largest specimens; in smallest individ- uals, tubercles and concavity virtually effaced. Spec- imens Examined. — 136 males, 131 females (length 12-18 mm; width 9-12 mm). DIAGNOSTIC REMARKS. The coloration and overall size and shape of chalcomelas are very sim- ilar to those of cambeforti; the two species are easily confused. They differ in secondary sexual characters (shape of male cephalic horn and female pronotum) and in the shape of the anterior prom- Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 29 inence of the metasternum. Certain meleagris also strongly resemble chalcomelas, but they also differ in the aforementioned characters. Smaller individ- uals, particularly males, must be examined carefully to avoid confusing these three species. DISTRIBUTION (Fig. 162; appendix). Ever- green forests of Guiana and the Amazon Basin of Brazil, Colombia, Ecuador, Peru, and Bolivia. 0- 500 m. Coprophagous. Collection dates: all year. COMMENTS. The known distribution of chal- comelas coincides with practically all of the Am- azonian hylea and covers the largest area of that of any Phanaeus. In spite of its wide distribution, there is remarkably little geographic variation in the char- acters that I have studied. Guianan males are more coppery brown and have fewer, smaller pronotal rugosities than specimens from elsewhere. This spe- cies is sympatric with cambeforti, and also with meleagris in those peripheral areas where meleagris descends to lower elevations (e.g., Buena Vista, Bo- livia; Pucallpa, Peru; Villavicencio, Colombia). Although I have listed it above, I am not sure that the Munich specimen is indeed the holotype of chalcomelas. It bears the indentification label “Phanaeus chalcomelas,” but the writing does not match the sample of Perty’s handwriting in Horn and Kahle (1937, Plate 34, Fig. 16). Moreover, Perty placed this species in Onitis, not Phanaeus. Phanaeus ( Notiophanaeus ) achilles Boheman Figures 30, 31, 141, 142, 146, 148, 150, 162 Phanaeus achilles Boheman, 1858:42 Type: Male holotype (“Puna”), Naturhistoriska Riksmuseet, Stockholm. Phanaeus foveolatus Harold, 1880a:152 (Gillet, 1911a:319) Type: Unknown to me. OTHER REFERENCES. Nevinson, 1892a; Gil- let, 1911b; Olsoufieff, 1924; Vulcano and Pereira, 1967; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 30, 31). — Dorsum brightly shining red, green or red with weak green or yellow high- lights except for black pronotal rugosities. Pygidi- um weakly shining dark green, yellow-green, or nearly black. Venter weakly shining black, some- times with weak green highlights. Head. — Para- ocular carina present, extending to lateral margin of eye. Pronotal Sculpturing. — Male (Fig. 141): Disk rugose; rugosities irregularly shaped, becoming weaker posteromedially, replaced by strong, rough puncturing; sides granulorugose with intermingled punctures anteriorly, strong punctures posteriorly. Female (Figs. 142, 146, 150): Anterior two-thirds of surface covered by melange of granules and rough, confluent punctures producing distinct black ru- gosities in some areas (especially on disk); posterior one-third punctate, becoming more weakly so pos- teriorly. Both sexes with distinct puncturing, at least on sides, sometimes obscured by surrounding, rough sculpturing in some specimens, particularly males. Elytra. — Interstriae distinctly, but finely, sparsely punctured ( x 10), with some irregular wrinkling (Fig. 150). Metasternum. — Anterior prominence acu- minate, tip produced as small, acute, slightly up- turned tubercle best seen in profile (as in Fig. 155). Secondary Sexual Characters. — Male: In large in- dividuals (Fig. 141), anterior three-fourths pronotal disk flat, posterior one-fourth weakly, transversely concave; concave posterior area bounded antero- medially by weakly defined, transverse carina or by small, raised, V-shaped ridge; sides of disk rounded, angulate posterolaterally, each interrupted near an- terior margin of pronotum by strong, conical tu- bercle. (No very small males observed.) Female: Cephalic carina (Figs. 142, 148) thickened, raised medially as weakly, anteriorly bowed ridge. Pro- notum (Fig. 142) evenly convex, with three weak, closely set, round tubercles near anterior margin; these tubercles often coalesce to form a thick, flat, trilobed ridge. Specimens Examined.— 15 males, 17 females (length 15-20 mm; width 9-13 mm). DIAGNOSTIC REMARKS. This species is easily distinguished from other members of the group by pronotal sculpturing, color, and distribution. The male, in particular, superficially resembles certain Phanaeus, sen. str. (e.g., wagneri ), from which it differs by having distinct puncturing of the elytral striae and an acuminate anterior prominence of the metasternum. DISTRIBUTION (Fig. 162; appendix). Fowland desert scrub region of southwestern Ecuador and northern Peru. Coprophagous. Collection dates: October, January-February. COMMENTS. P. achilles is a unique species that I have assigned to the chalcomelas group on the following basis: conspicuous, foveolate puncturing of the elytral striae; black pronotal rugosities mixed with distinct puncturing (particularly in the female), acuminate anterior prominence of the metasternum (as in meleagris and cambeforti ), and female sec- ondary sexual characters (similar to those of mele- agris and cambeforti). In 1972 I included it in the “ mexicanus complex” on the basis of a resem- blance of the male (the female was unknown to me at the time) to such species as wagneri. The female, however, is quite different from those of the mexicanus group (q.v.); it most closely resem- bles those of meleagris and cambeforti. Neverthe- less, achilles is a taxonomic isolate within the chal- comelas group not only morphologically (form of the male pronotum, pronotal sculpturing, colora- tion) but also ecologically and geographically. Whereas other members of the group inhabit the rain forests of the Amazon Basin, achilles is re- stricted to the desert scrub region of southern Ec- uador and adjacent northern Peru. Bruce Gill (pers. comm.) has collected this species from feces in the region of dry scrub vegetation 45 km west of Gua- 30 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus yaquil as well as from “wet forests” on the outskirts of the same city. Pbanaeus ( Notiophanaeus ) cambeforti Arnaud Figures 25, 26, 139, 140, 157, 162 Pbanaeus cambeforti Arnaud, 1982b:122 Type: Male holotype (“French Guiana, Saul; IV- 1977”), Museum National d’Histoire Natu- relle, Paris. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 25, 26). — Dorsum dull olive brown to olive green with speckling on pronotum pro- duced by black rugosities (more extensive on fe- male). Pygidium dark green or olive brown; venter with dark, shining green reflections. Head. — Para- ocular ridge lacking. Pronotal Sculpturing. — Disk sparsely (in male) to densely (in female) covered with low, black, blister-like rugosities; entire sur- face of pronotum with large, shallow punctures (best seen from low angle); each puncture with cen- tral, shining microspot, which can be difficult to detect (x40). Pronotum. — Basal fossae usually ab- sent; if present, small and partially effaced. Meta- sternum.— Anterior prominence acuminate, not capped by a swelling. Secondary Sexual Charac- ters.— Male: Cephalic horn (seen from front, Fig. 157) gradually tapering from base to apex; prono- tum as in chalcomelas. Female: Pronotum (Fig. 140) evenly convex, with three small, round, smooth tubercles near anterior margin. Specimens Exam- ined.— 16 males, 19 females (length 12-15 mm; width 7-10 mm). DIAGNOSTIC REMARKS. While this species can closely resemble meleagris, its more distinct pronotal puncturing, much weaker and less exten- sive pronotal rugosity, absent (or very small) basal pronotal fossae, smaller size and olive coloration lacking red or pink highlights will distinguish it from its closest relative. (See also “Diagnostic Re- marks” under chalcomelas.) DISTRIBUTION (Fig. 162; appendix). Known from scattered localities in Guiana and Amazon Basin of Brazil, Colombia, and northern Peru. 0- 50 m. Coprophagous. Collection dates: November- April (but probably all year). COMMENTS. All examined specimens of cam- beforti bearing precise data were collected from localities where chalcomelas also occurs (the con- verse, however, is not true). On superficial exami- nation these two species closely resemble each oth- er; cambeforti was not described until 1982. Pbanaeus ( Notiophanaeus ) meleagris Blanchard Figures 27-29, 143-145, 155, 158, 162 Pbanaeus meleagris Blanchard, 1843:176 Type: Male lectotype (“Yungas”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:114). Pbanaeus minos Erichson, 1847:106 (Harold, 1870: 105) Type: Male lectotype; 2 male, 1 female paralec- totypes (“mountains of Peru”), Museum fiir Naturkunde, Berlin, PRESENT DESIGNA- TION. OTHER REFERENCES. G uerin-Meneville, 1855; Lacordaire, 1856; Gemminger and Harold, 1869; Kirsch, 1873 (1874) (as minos ); Nevinson, 1892a; Gillet, 1911b; Olsoufieff, 1924; Pessoa, 1934; Blackwelder, 1944; Gacharna, 1951; Halffter and Matthews, 1966; Vulcano and Pereira, 1967; Ed- monds, 1972; Arnaud, 1982a. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 27-29). — Variable, three classes as follows: (a) dorsum dull black except for shining red to yellow-red (rarely yellow-green) highlights on pronotum and posterior portion of head; prono- tal coloring usually restricted to sides and to narrow line along posterior margin, but sometimes extend- ing onto disk; elytra with at most extremely weak, bluish sheen (x20); pygidium weakly shining red, red-green, or yellow-green; venter with strong high- lights colored like pygidium; (b) dorsum evenly dark blue or green, dull to weakly shining; color of pro- notum broken only by black rugosities; venter with strong blue (or green) reflections; (c) dorsum dull, usually mostly black; elytra, colored areas of pro- notum black-brown or mahogany brown, rarely olive green; pronotum with weak pink, yellow-red, yellow-green, or coppery highlights at least antero- laterally and along posterior margin; pygidium weakly shining pink to red or red-green; venter, particularly femora, with strong pink highlights. Head. — Paraocular ridge absent or only weakly in- dicated. Pronotal Sculpturing. — Rugosity dense on disk and often also on sides; shallow punctures covering most of surface except where obliterated by rugosities, but often very difficult to see even at low angle; each puncture bearing small, central, shining microspot which persists even if associated puncture effaced. Pronotum. — Basal fossae pres- ent, sometimes punctiform, rarely almost effaced. Metasternum. — Anterior prominence acuminate, not capped by swelling (Fig. 155). Secondary Sexual Characters. — Male: Cephalic horn of large individ- uals (seen from front, Fig. 158) swollen basally and tapering apically; basal swelling not abruptly nar- rowed so as to produce angular prominences; horn reduced to conical tubercle in smallest individuals. Pronotum as described for cambeforti. Female: Pronotum (Fig. 145) evenly convex, with three, smooth, round tubercles near anterior margin; sur- face sometimes distinctly flattened in front of these tubercles. Specimens Examined. — 67 males, 81 fe- males (length 14-21 mm; width 8-14 mm). Contributions in Science, Number 443 Edmonds: Revision of PhanaeusM 31 Figures 163-170. Phanaeus ( Notiophanaeus ) bispinus group (163, P. bispinus, male; 164, same; 165, same, female; 166, same, anterolateral portion of female pronotum; 167, P. alvarengai, holotype male; 168, same, allotype female; 169, P. bispinus, left elytron; 170, P. alvarengai, same). DIAGNOSTIC REMARKS. See under cbalco- melas and cambeforti. DISTRIBUTION (Fig. 162; appendix). Eastern slopes of Andes (“Yungas”) from vicinity of Co- chabamba, Bolivia, to southwestern Venezuela. 200- 2000 m (estimated). Reportedly necrophagous (Halffter and Matthews, 1966); inhabiting forests, probably mostly cloud forests (“montanas”). Col- lection dates: March-January. COMMENTS. The coloration of meleagris is more variable than that of any other species in this group. The lectotype of meleagris exhibits pattern (a) above, black with red markings along the prono- tal margin; that of minos is dark, black-brown (c). Except for two labeled “Columbia” [s/c], all spec- imens (about 45) with color pattern (a) that I have seen are from the cloud forests (“Yungas”) of Bo- livia. These specimens are also slightly, but consis- tently, smaller than meleagris from other places, and they may later prove to belong to a distinct taxon. Most meleagris examined conform to color pat- tern (c), a highly variable combination of colors best described collectively as “dirty brown” or, sim- ply, “brownish.” The common feature of the (c) pattern is pink or reddish highlights on the pro- notum and pygidium, which are never seen in (a) and (b). Certain of these specimens, because they can assume olive brown tones, resemble cambeforti and chalcomelas; this is the only form of meleagris that I have seen from lower (<300 m) localities. There is not enough precise locality data to clear- ly define the geographic distribution of the species. While I suspect that it is most common in cloud forests, it nevertheless clearly enters lowland rain forest where it occurs along with chalcomelas and cambeforti. Thanks to Patrick Arnaud, I have seen nine un- usual specimens that I have tentatively assigned to meleagris. Three are dark, weakly shining blue and labeled “Peru,” “Bolivie-Coroico” (in the “Yun- gas” region northeast of La Paz), and “Bolivie-Son- go” (not located); six are dark olive green from Pucallpa, Peru (Amazonian hylea). All nine are large, the largest meleagris I have seen, and in the males, the cephalic horn is more strongly swollen basally 32 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus 183 Figures 171-183. Phanaeus ( Notiophanaeus ) bispinas group (171, P. bispinus, small male, dorsal view of head; 172, same, lateral view; 173, P. alvarengai, male holotype, dorsal view of head; 174, same, lateral view; 175, frontal view cephalic carina, P. alvarengai, male holotype; 176, same, P. bispinus, small male; 177, same, P. bispinus, female; 178, P. bispinus, female, anteromedian portion of pronotum [A = lateral view; B - frontal view]; 179, P. alvarengai, allotype female, same; 180, P. alvarengai, holotype male, front tibia [stippling = reconstruction of tibial teeth]; 181, P. bispinus, female, front tibia; 182, same, female; 183, same, frontal view clypeal process). than in typical meleagris ; otherwise, they differ in no significant way from other specimens. It may well be that these specimens will later prove to represent a distinct taxon, but until more material and more precise geographical data are available, it will be difficult to consider them more than un- usual variants of meleagris. The lectotype of minos designated here is from among four specimens labeled in Erichson’s hand from the Berlin Museum. It agrees with the original description in all respects. The Bispinus Group DIAGNOSIS. [1] Anterior margin of clypeus (Figs. 171, 173) bidentate medially, strongly so in male alvarengai ; [2] clypeal process (seen from front) rounded (Fig. 183); [3] cephalic process of female a trituberculate carina; [4] anterolateral portions of pronotum smooth, virtually devoid of sculpturing (Fig. 166); [5] pronotal disk smooth; [6] pronotum of male rounded, lacking distinct posterolateral an- gles, bearing anteromedian concavity from whose upper margin project pair of slender, anteriorly pro- jecting spines (Figs. 163, 167); [7] pronotum of fe- male as in Figures 178, 179; [8] basal pronotal fossae present, conspicuous; [9] front tibiae tridentate (Figs. 180-182); [10] elytral striae fine, carinulate, punc- tate (Figs. 169, 170); interstriae weakly convex; [11] dorsum dark, somber (Figs. 15, 16); [12] Amazonian rain forest (Fig. 184). This group comprises two species, bispinus and alvarengai. The pronotal armature of both sexes is unique to the genus and approached only re- motely in baroldi. Besides those characters men- tioned above, these species differ from other Phan- aeus by possessing distinct notching of the hind wing (not observed in alvarengai) and proportion- ately larger eyes. Their coloration, like that of chal- comelas and cambeforti, suggests camouflage. KEY TO THE SPECIES OF THE BISPINUS GROUP la. Clypeus of male (Figs. 173, 174) drawn out anteriorly, sides almost straight; median teeth prominent, strongly upturned. Pronotum of fe- male bearing flat, quadrate ridge (Figs. 168, 179). Punctures of elytral striae not conspicuous to unaided eye (Fig. 170). Front tibiae tridentate; all three teeth distinctly carinate on outer sur- face (Fig. 180). Amazonia Phanaeus ( N .) alvarengai Arnaud b. Clypeus of both sexes rounded, median teeth inconspicuous and not strongly upturned (Figs. 171, 172). Pronotum of female (Figs. 165, 178) with strong, circular, anteromedian concavity, upper margin of which bears a strong, laterally flattened spine or (in small individuals) vertical, keel-like tubercle. Punctures of elytral striae large, easily seen with unaided eye (Fig. 169). Front tibiae tridentate; only apical two teeth distinctly carinate on outer surface (Figs. 181, 182). Amazonia (Fig. 184) Phanaeus (N.) bispinus Bates Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 33 Phanaeus ( Notiophanaeus ) alvarengai Arnaud Figures 167, 168, 170, 173-175, 179, 180 Phanaeus alvarengai Arnaud, 1984:60 Type: Male holotype (“Bresil, Matto Grosso”), private collection of Patrick Arnaud, Saintry, France. DESCRIPTIVE REMARKS. Color and Color Pattern. — Dorsum dull, almost black except for metallic green pronotum, posterior portion of head of male, lateral parts of female pronotum. Pygidium green; venter almost black. Pronotal Sculptur- ing. — Appearing smooth to unaided eye; with mod- erate magnification (xlO) weak puncturing visible along sides, most conspicuous in female. Elytra (Fig. 170). — Striae fine, carinulate ( x 10), each bear- ing 30-35 small punctures not at all conspicuous to unaided eye; interstriae weakly convex. Front Tibiae. — Tridentate in both sexes, all three teeth distinctly carinate longitudinally (Fig. 180). Second- ary Sexual Characters. — Male : Clypeus (Figs. 173, 174) drawn out anteriorly, sides almost straight, median teeth prominent and strongly upturned; length along midline about twice that of flat frontal area between eyes, behind transverse cephalic Ca- rina. Cephalic process (Fig. 175) transverse carina weakly bituberculate medially. Pronotum (Fig. 167) with anteromedian concavity from top margin of which projects pair of nearly straight, slender, par- allel spines. Female: Clypeus rounded; median teeth inconspicuous (worn in allotype); length about equal to that of frontal area. Cephalic process transverse carina (worn in allotype, presumably weakly tritu- berculate). Pronotum (Figs. 168, 179) convex, with weak, quadrate ridge above anterior margin; this ridge (seen from front) rounded laterally, (seen from above) minutely beaded in the middle; surface weakly concave beneath ridge. Specimens Exam- ined.— 1 male (holotype), 1 female (allotype) (length 16-22 mm; width 8-9 mm). DIAGNOSTIC REMARKS. See bispinus, be- low. DISTRIBUTION. The allotype female is labeled “Brazil, Rondonia, Rio Japunda (September)”; nei- ther this nor the stated type locality is precise enough to locate on a map. COMMENTS. Arnaud described this species from two specimens. While each is definitely dif- ferent from other known Phanaeus, only addition- al evidence will support the conclusion that they are, indeed, conspecific. The allotype is in the col- lection of the Rijksmuseum van Natuurlijke His- toric in Leiden. 34 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Phanaeus ( Notiophanaeus ) bispinus Bates Figures 15, 16, 163-166, 169, 171, 172, 176-178, 181-184 Phanaeus bispinus Bates, 1868:89 Type: Male holotype (“Canelos, Ecuador”), Mu- seum National d’Histoire Naturelle, Paris. Phanaeus digitalis Olsoufieff, 1924:34 (Pereira and Martinez, 1956:237) Type: Female holotype (“Guyana”), Museum National d’Histoire Naturelle, Paris. OTHER REFERENCES. Gemminger and Har- old, 1869; Nevinson, 1892a; Gillet, 1911b; Black- welder, 1944; Pereira and Martinez, 1960; Halffter and Matthews, 1966; Edmonds, 1972; Arnaud, 1982a. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 15, 16). — Dorsum dull, dark olive green to olive brown, with narrow line of shining green along circumnotal ridge, lateral margins of elytra. Pygidium usually bright moderately shining green, sometimes almost black; venter weakly shin- ing, dark green. Head. — Clypeus of both sexes weakly bidentate medially, evenly rounded laterally (Fig. 171); length along midline equal to or less than that of frontal area between eyes, behind clypeal process. Pronotal Sculpturing (Fig. 166). — Virtually smooth, unsculptured, at most with largely effaced, minute puncturing visible only under high magni- fication (x40), appearing smooth even under mod- erate (x20) magnification. Metasternum. — Ante- rior prominence compressed laterally; seen from side, rounded, longitudinally keeled. Elytra (Fig. 169). — Striae fine, carinulate (x30), each bearing 15-20 punctures conspicuous to unaided eye; in- terstriae weakly convex, smooth except for weak puncturing (x30). Front Tibiae (Figs. 181, 182). — Tridentate in both sexes, slightly narrower in fe- male; first two teeth carinate longitudinally, third (basal) tooth sometimes with weak carina. Second- ary Sexual Characters. — Male: Cephalic horn of larger individuals (Fig. 163) slender, suberect, slight- ly bowed posteriorly; apex (seen from front) slightly expanded laterally, compressed anteroposteriorly; that of smallest individuals (Figs. 164, 176) a carina, middle of which strongly raised, bituberculate. Pro- notum of large individuals (Fig. 163) with antero- median concavity from middle of posterior edge of which arises pair of closely set, long, slender an- teriorly directed horns; that of smallest individuals (Fig. 164) more-or-less evenly convex, bearing two prominent acute tubercles near anterior margin. Fe- male: Cephalic carina (Fig. 177) trituberculate, in larger individuals distinctly elevated. Pronotum of larger individuals (Figs. 165, 178) with deep, round- ed anteromedian concavity, sides of which carinate, posterior margin of which bears single, strong, fin- ger-like, anteriorly projecting, ventrally curved spine; that of smallest individuals lacking concavity and bearing two short longitudinal ridges, with prom- inent blunt tubercle near anterior margin. Speci- mens Examined. — 16 males, 15 females (length 13- 17 mm; width 7-11 mm). DIAGNOSTIC REMARKS. This species differs from alvarengai by its secondary sexual characters, the shape of the male clypeus, non-carinate third tibial tooth, and puncturing of the elytral striae. DISTRIBUTION. (Fig. 184; appendix). Known from scattered Amazon Basin localities in Brazil, Venezuela, Colombia, Ecuador, Peru, and Bolivia. 0-500 m (estimated). Necrophagous. Collection dates: all months except March, November, and December; probably all year. COMMENTS. Although widely distributed geo- graphically, this curious species is rather rare in collections. Although he suspected conspecificity, Olsoufieff (1924) described females available to him as digitalis. Of interest is the fact that the hind wing of bispi- nus, like Coprophanaeus, is notched basally. This species is one of very few in the genus that does not prefer excrement. The Palaeno Group DIAGNOSIS. [1] Clypeus strongly bidentate me- dially (Fig. 196); [2] clypeal process spiniform (Fig. 195); [3] cephalic carina of female distinctly tritu- berculate; [4] cephalic process of male dimorphic, either slender, cylindrical, posteriorly curved, ta- pering horn (in larger individuals, Fig. 185) or carina with strongly raised, medially bidentate process (smallest individuals, Fig. 187); [5] pronotum smooth except for very fine, sparse puncturing ( x 30); [6] shape of male pronotum variable (Figs. 185, 187); in female, disk distinctly swollen posterolaterally on each side of midlongitudinal depression and (seen in profile) distinctly raised above level of elytra; [7] basal pronotal fossae absent, basal angle prominent, acute; [8] elytral striae carinulate, not punctate; in- terstriae convex; [9] anterior prominence of meta- sternum (seen in profile, Fig. 194B) lobate, (seen from below, Fig. 194A) compressed laterally and longitudinally keeled; [10] pteropleura and sides of metasternum clothed by dense pile of long white or yellowish hair (indistinct in worn or very dirty specimens); [11] front tibiae (Figs. 197, 198) quad- ridentate, apical three teeth longitudinally carinate; [12] dorsum (Figs. 10-14) highly shining, bright me- tallic green (rarely dark blue or blue-green); female with pair of black, often confluent areas behind anteromedian pronotal carina; [13] inhabiting sa- vanna (“campos cerrados”) of Brazilian Highlands and adjacent areas to south (Fig. 199). This group comprises two species, palaeno Blan- chard and kirbyi Vigors, that share several char- acters not found in other Phanaeus: spinate clypeal process; strongly bidentate cephalic carina of cer- tain males; and thick, whitish pile on underside of Contributions in Science, Number 443 Edmonds: Revision of Phanaeus £1 35 Figures 185-191. Phanaeus ( Notiophanaeus ) palaeno group (185, P. kirbyi, male; 186, same, female; 187, same, frontal view small [arrow indicates pronotal tubercles]; 188, same, frontal view female; 189, P. palaeno, male; 190, same, female; 191, same, frontal view female). pterothorax. All of these features, however, are shared with Oxysternon palaemo Laporte-Castle- nau ( palaemon of authors), which is easily confused with F. palaeno, in particular. The two species are sympatric and ecologically very similar. In this species group, males have either a strongly bidentate cephalic carina or a clearly horndike pro- tuberance; there is no intermediate state, and males of the same (always intermediate) size may possess either option. Dimorphism of the male cephalic process is also seen in F. bispinus and endymion as well as in the phanaeine genus Oxysternon. This is the only species group of Phanaeus (but not of phanaeines) that inhabits the “campos cerra- dos” of the Brazilian Highlands. As savanna species, their recent distribution has probably expanded considerably since pre-European times as forests in southern Brazil have been cut to produce pasture- lands. KEY TO THE SPECIES OF THE PALAENO GROUP la. Paraocular areas flat above. Anterior portion of the circumnotal ridge raised behind and nar- rowly excised medially, excision separating two small, dentiform tubercles which are rarely ef- faced even in smallest individuals (Fig. 187, arrow). Elytral interstriae weakly convex. Male : Pronotum of large specimens (Fig. 185) flat- tened above and expanded laterally as round- ed, flat lobes. Female: Pronotum of all but smallest individuals bearing almost straight, transverse anteromedian carina (Fig. 188) . . . Phanaeus (N.) kirbyi Vigors b. Upper surface of paraocular areas swollen ad- jacent to eyes, flattened laterally. Anterior por- tion of circumnotal ridge not noticeably raised and neither excised nor bidentate medially. 36 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ Elytral interstriae moderately convex. Male: Pronotum of larger individuals (Fig. 189) shal- lowly concave above, posterior angles pro- duced upward as pair of apically convergent, widely separated horns. Female: Pronotum of all but smallest individuals (Fig. 191) bearing a wide, U-shaped anteromedian carina ......... Fbanaeus (N.) palaeno Blanchard Phanaeus ( Notiophanaeus ) kirbyi Vigors Figures 13, 14, 185-188, 196-199 Phanaeus kirbyi Vigors, 1825:539 (as kirbii ) Type: Male holotype (“Brazil”), British Museum (Natural History), Fondon. Phanaeus planicollis Perty, 1830:40 (Nevinson, 1892a:5) Type: Male holotype (no data), Zooligsche Staatssammlung, Munich. Phanaeus subtricornis Perty, 1830:41 (Harold, 1859: 198) Type: Male holotype (no data), Zooligsche Staatssammlung, Munich. Phanaeus laevicollis Laporte-Castelnau, 1840:82 (Nevinson, 1892:5) Type: Male neotype (“Bresil”), Castelnau Col- lection, National Museum of Victoria, Mel- bourne, PRESENT DESIGNATION. Phanaeus kirbyi truncaticornis Olsoufieff, 1924: 91, NEW SYNONYMY Type: Unknown to me. NOMENCLATURAL REMARKS. Although Harold (1869:65) synonymized planicollis and laev- icollis, Nevinson (1892a) was the first to syn- onymize them with kirbyi. OlsoufiefPs (1924) trun- caticornis is a small male described as a variation of kirbyi. OTHER REFERENCES. Klug, 1841; Guerin- Meneviile, 1855 (as planicollis ); Lucas, 1857; Har- old, 1871a; Preudhomme de Borre, 1886; Gahan and Arrow, 1903; Gillet, 1911b; Pessoa, 1934; Pes- soa and Lane, 1941; Blackwelder, 1944; Lange, 1947; Halffter and Matthews, 1966; Barrera, 1969; Ed- monds, 1972; Scherer, 1983. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 13, 14). — Dorsum shining green to yellow-green; rarely dark, shining blue-green. Ven- ter mostly shining green or blue-green, otherwise dark brown. Paraocular Areas.-— Upper surface flat, not swollen adjacent to eyes. Pronotum. — Anterior portion of circumnotal ridge strongly raised, almost vertical; posterior margin of ridge bearing pair of Figures 192-198. Phanaeus ( Notiophanaeus ) palaeno group (192, P. palaeno, female [Corrientes, Argentina; A = lateral view cephalic carina; B = dorsal view of same; C = dorsal view pronotal carina]; 194, P. palaeno, anterior prominence of metasternum [A = ventral view; B = lateral view]; 195, same, subclypeal process [A = frontal view; B = lateral view]; 196, P. kirbyi, dorsal view clypeal mar- gin; 197, same, front tibia of male; 198, same, front tibia female). closely set, small median denticles (Fig. 187, arrow) separated by shallow, rounded notch; denticles and notch only rarely almost effaced. Secondary Sexual Characters. — Male: Pronotum of larger individuals (Fig. 185) unevenly flattened, posterolateral angles (seen from above) produced laterally as broadly rounded flanges, outer margins of which narrowly upturned; these angles reduced to low, angular tu- mosities in smaller specimens. Female: Pronotum (Figs. 186, 188) bearing sharp, almost straight trans- verse carina near anterior margin; carina followed by narrow, transverse depression; carina and de- pression almost effaced in some small specimens. Specimens Examined. — 38 males, 31 females (length 15-23 mm; width 9-13 mm). DIAGNOSTIC REMARKS. See palaeno, below. DISTRIBUTION (Fig. 199; appendix). Brazilian Highlands and extensions thereof into eastern Bo- livia and eastern Paraguay. 300-1000 m (estimated). Coprophagous (one record from carrion); inhabit- ing “campos cerrados” and similar savanna-like for- mations resulting from forest clearing for pasture- land. Collection dates: September-June (most November-March). COMMENTS. As discussed above, kirbyi is sympatric with palaeno, although its known range is somewhat smaller. This is the larger of the two species, and it exhibits less color variation than palaeno. The elytra are always completely green (or Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 37 palaeno group palaeno kirbyi "Corrientes" palaeno Figure 199. Distribution of the Phanaeus ( Notiophanaeus ) palaeno group. blue-green); they never have the yellow-brown markings seen in most palaeno. There are two specimens of laevicollis in the Castlenau Collection in the National Museum of Victoria (Melbourne). One is a female, the other a small male with a bidentate cephalic carina. I hereby designate the latter as neotype. Laporte-Castelnau’s (1840) original description included a large and small male, the latter of which he mistakenly assumed was a female (“. . . tete . . . avec . . . un petit tuber- cule bifide sur la [femelle] . . .”). Phanaeus ( Notiophanaeus ) palaeno Blanchard Figures 10-12, 189-195, 199 Phanaeus palaeno Blanchard, 1843:176 Type: Male holotype (“Bolivie, Concepcion de Chiquitos”), Museum National d’Histoire Na- turelle, Paris. Phanaeus blanchardi Harold, 1871a:114 (not Ol- soufieff, 1924:92), new name for palaeno Blan- chard Phanaeus palaeno exisicornis Balthasar, 1939:247 (emended to excisicornis, Balthasar, 1940:39) (Arnaud, 1982c:125) Type: Male holotype (“Brazil, Matto Grosso”), Narodni Muzeum Prirodovedecke Muzeum, Prague. NOMENCLATURAL REMARKS. Harold s “ blanch ardr was based on the mistaken notion that Blanchard’s species was the same as Oxyster- non palaemo Laporte-Castelnau (see Gemminger and Harold, 1869:1016). The error was corrected by Nevinson (1892a). OTHER REFERENCES. Lacordaire, 1856; Lu- cas, 1857; Gahan and Arrow, 1903; Bruch, 1911; Pessoa, 1934; Pessoa and Lane, 1941; Blackwelder, 1944; Martinez, 1959; Halffter and Matthews, 1966; Barrera, 1969; Edmonds, 1972; Arnaud, 1982a. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 10-12). — Dorsum usually shining green to yellow-green, often with golden to cop- pery highlights on head, anterior portions of pro- notum; rarely evenly blue-green or dark blue, much less shining; elytra either evenly colored, or with yellow-brown lateral markings with green high- lights, or (very rarely) almost completely yellow- brown with green highlights. Venter dark brown with green to blue-green highlights. Paraocular Ar- eas.— Upper surface usually distinctly swollen ad- jacent to eyes, flattened laterally. Pronotum. — An- terior portion of circumnotal ridge simple, neither 38 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus raised nor bituberculate medially. Secondary Sex- ual Characters. — Male: Pronotum of larger indi- viduals (Fig. 189) broadly, shallowly concave dor- sally, each posterolateral angle produced upward as wide process tip of which (seen laterally) acute, directed forward; these processes lower, quadrate in smaller individuals; in smallest specimens, re- duced to pair of anteriorly converging, low, round- ed ridges extending from near anterior margin to middle of disk. Female: Pronotum of all but small- est individuals with fine, broadly U-shaped antero- median carina (Fig. 191), middle of which may bear small tubercle; this carina followed by concavity interrupted by low, truncated or round swelling or tubercle, flanked on each side by elongate concavity extending to anterior margin of pronotum behind eye. In smallest specimens, carina and adjacent fea- tures less pronounced to virtually effaced (see also “Comments,” below). Specimens Examined. — -96 males, 82 females (length 10-19 mm; width 7-12 mm). DIAGNOSTIC REMARKS. This species is easily distinguished from kirbyi by the shape of the an- terior portion of the circumnotal ridge and by sec- ondary sexual features; only the very smallest in- dividuals of the two species are likely to be confused. DISTRIBUTION (Fig. 199; appendix). Brazilian Highlands and extensions thereof into eastern Par- aguay, eastern Bolivia, and extreme northeastern Argentina. 300-1000 m (estimated). Coprophagous, inhabiting “campos cerrados” and similar savanna- like formations resulting from forest clearing for pastureland. Collection dates: October-July (most October-March). COMMENTS. As I pointed out in 1972, this species bears a striking resemblance to Oxysternon palaemo. Well-developed males of the two species differ by the shape of the pronotum, which in O. palaemo is much more strongly concave. Females, on the other hand, are so similar that they can be confused even on fairly close inspection if the shape of the metasternum (drawn out anteriorly into a sharp, upturned spine projecting between the front coxae in Oxysternon) is not examined. P. palaeno is widely distributed in the “campos cerrados” and similar areas of the Brazilian High- lands, where it occurs along with kirbyi and O. palaemo. I do not know if these species are sepa- rated ecologically in some obvious way (foraging time, preferred ecotones, etc.); they would make interesting subjects for a serious community study. Balthasar’s subspecies exisicornis was based upon a single small male; it is not a valid taxon. I have seen a series of eight specimens tentatively assigned here to palaeno collected by Antonio Martinez in Corrientes Province, Argentina, that has the fol- lowing unusual female secondary sexual features: the cephalic carina is strongly raised and the middle tubercle, which is much larger than the lateral two, is abruptly offset posteriorly (Fig. 192); the pronotal carina (Fig. 193) is brace-shaped ({), more elevated than its counterpart in other specimens, and not associated with a concavity. This series may later prove to represent a distinct taxon. The Endymion Group DIAGNOSIS. [1] Anterior margin of clypeus strongly bidentate medially; [2] clypeal process (seen from front) transverse; [3] pronotum usually ap- pearing completely smooth to unaided eye (Figs. 200-208); sculpturing ( x 10-30) restricted to simple puncturing except on disk of male, which can bear minute granulations and similar formations (Figs. 209-211); [4] cephalic horn of male (Figs. 200, 203, 206) strongly curved posteriorly over pronotum, thickened basally, strongly tapered to apex; re- duced to feebly bituberculate carina in smallest in- dividuals; [5] pronotum of larger males (Figs. 200, 203, 206, 214) flat above, disk assuming a triangular shape with prominent, acute posterolateral angles; flattened, triangular disk of smaller males propor- tionately smaller, located anteriorly, with only weak, tuberculate posterolateral angles; [6] pronotum of female (Figs. 201, 204) convex, with three antero- median tubercles of varying shape and configura- tion; [7] basal pronotal fossae punctiform; [8] front tibiae (Figs. 219, 220) quadridentate, narrower in male; fourth (basal) tooth separated from third by narrow, slit-like notch; [9] apical spur of front tibia, viewed anteriorly, more-or-less straight, apex bent posteriorly; [10] elytral striae fine, not carinulate, simple to (rarely) punctate (x5) (Figs. 212, 213); [11] dorsal coloration (Figs. 17-22) almost always at least partially shining green, red, or dark blue; pronotum of female with irregularly shaped, central black area of variable size; [12] coprophagous, co- pronecrophagous (one species mycetophagous); [13] inhabiting neotropical forests from southern Mex- ico to northwestern South America. This group brings together three (or four) closely related Mesoamerican species that closely resemble each other in general form: endymion Harold, py- rois Bates, and halffterorum Edmonds (a possible fourth species is known from two Oaxacan speci- mens here included in endymion). Member species are distinguished primarily on the basis of color and color pattern, relief and sculpturing of the pro- notum and elytra, secondary female sexual char- acters of the pronotum, and geographical distri- bution. Both pyrois and endymion are common, widely distributed species that vary considerably, but in ways too inconsistent to warrant recognition of subspecies. Because of their variability, each is difficult to define taxonomically, but most speci- mens are quickly identifiable by color and place of origin. P. halffterorum is a rare, fungus-feeding spe- cies known only from a few isolated localities in southern Mexico. KEY TO THE SPECIES OF THE ENDYMION GROUP la. Inner margin of each elytron upturned to form sharp ridge progressively more elevated pos- Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 39 Figures 200-208. Phanaeus ( Notiopbanaeus ) endymion group (200, P. endymion, male; 201, same, female; 202, same, dorsal view anteromedian portion of prothorax; 203, P. pyrois, male; 204, same, female; 205, same, dorsal view anteromedian portion of pronotum; 206, P. halffterorum, male; 207, same, anterolateral portion of pronotum of female; 208, same, dorsal view base of pronotum of female). teriorly, forming small tooth at the apical angle of elytron (Figs. 217, 218); margin of elytron slightly excised adjacent to this tooth. Pygidi- um shallowly impressed on each side. Male: Disk of pronotum (Fig. 211) distinctly but very finely granulorugose, more strongly so laterally ( x 10); middle of anterior margin of pronotum of large individuals sometimes bearing small, acute tooth; posterior angles of disk of large specimens directed posteriorly. Female: Pro- notum (Fig. 208) bearing a midlongitudinal de- pression extending from near base to near mid- dle of disk, and three isolated anteromedian tubercles followed by shallow concavity. Dor- sum shining green (brighter in male) or dark blue (Figs. 21, 22); pronotum of female shining black except for green (or blue) areas along margins. Mycetophagous; south central Mex- ico (Fig. 221) Phanaeus (N.) halffterorum Edmonds b. Inner margins of elytra (Figs. 215, 216) not upturned to form ridge terminating in an acute tooth; apical margin of elytra straight. Pygidi- um evenly convex, lateral impressions lacking or only very weakly indicated. Male: Disk of pronotum distinctly granulorugose at most only laterally, central area velvety smooth, only rarely sparsely granulate (x15; except Oaxaca pop- ulation of endymion, which is evenly and mi- nutely granulate); anteromedian pronotal tooth lacking; posterior angles of pronotal disk di- rected laterally. Female: Pronotum lacking dis- tinct midlongitudinal impression (except Oaxa- can endymion ); otherwise variable. Dorsum dull black, dull black with shining red, shining green, or dull blue. Coprophagous, copronecropha- gous; southern Mexico to northwest South America 2 2a. Elytral interstriae (Fig. 213) evenly convex and shining midlongitudinally; striae impressed ba- sally as distinct fossae. Male: Pronotal disk (Fig. 209) ( x 20) velvety smooth medially, asperate laterally and sometimes also medially (distinct- ly evenly granulate [ x 10] in Oaxaca specimens; see below). Female: Pronotum (Fig. 201) with anteromedian concavity bounded anteriorly by raised U- or V-shaped ridge; this ridge pro- duced medially as an acute or rounded tuber- 40 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Figures 209-213. Phanaeus ( Notiophanaeus ) endymion group (209, dorsal view left side pronotum of male P. endymion; 210, same, P. pyrois; 211, same, P. halffterorum; 212, dorsal view left elytron P. pyrois; 213, same, P. balffterorum). cle, and at each end as rounded or elliptical tubercle; in some individuals, concavity inter- rupted posteriorly by small, rounded bump; in some, usually small individuals concavity ob- solete, ridge reduced to three isolated, round tubercles. Dorsum (Figs. 17, 18) shining green or dark blue (Oaxaca population shining black); rarely shining green with strong yellow reflec- tions. Southwestern Mexico and southeastern Mexico to Honduras (Fig. 221) Phanaeus (N.) endymion Harold b. Elytral interstriae (Fig. 212) distinctly flattened and uniformly dull (more convex and shining in some Central American populations); striae not strongly impressed basally, anterior ends usually bearing deep punctures rather than large fossae. Male: Pronotal disk (Fig. 210) dull, vel- vety smooth medially, finely asperate, brighter laterally. Female: Pronotum (Fig. 204) evenly convex, lacking anteromedian concavity even in largest specimens, bearing three round, smooth tubercles in transverse line near ante- rior margin. Head and pronotum (Figs. 19, 20) largely highly shining metallic red to nearly completely dull black with metallic red re- stricted to ridges and isolated areas on anterior part of pronotum; elytra dull to weakly shining black; pygidium usually metallic red medially, green peripherially, sometimes completely red or green. Southern Nicaragua through Central America into western Colombia and Ecuador (Fig. 221) Phanaeus (N.) pyrois Bates Phanaeus ( Notiophanaeus ) halffterorum Edmonds Figures 21, 22, 206-208, 211, 217, 218, 221 Phanaeus halffterorum Edmonds, 1979:99 Type: Male holotype (“Mexico, State of Mexico, 8km west Temascaltepec, 2360m; 11 July 1976; fungus in pine-oak forest”), California Acad- emy of Sciences, San Francisco (Ento. Type No. 13184). OTHER REFERENCES. Hinton, 1935 (as en- dymion)', Edmonds, 1972 (as incertae sedis ); Ed- monds and Halffter, 1978; Moron, 1984; Delgado- Castillo, 1989 (as “ Phanaeus sp.”). DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 21, 22). — Except for central black area on female pronotum (difficult to see in dark specimens), dorsum either shining green or dark blue, female may be entirely black; in green males, color highly iridescent on head, pronotum, duller on elytra; in green females, blue individuals of both sexes, iridescence lacking; pronotum of male evenly colored, coloration of female pronotum restricted to sides, anterolateral angles. Venter red-brown to chocolate brown with green (or blue) reflections on pteropleura, legs, abdominal sterna. Pronotal Sculpturing. — Disk of male pronotum (Fig. 211) unevenly covered with variable-sized granules which become coarser laterally, progressively weaker ba- sally ( x 15) until replaced by minute punctures; sides, Contributions in Science, Number 443 Edmonds: Revision of PhanaeusM 41 Figures 214-220. Phanaeus ( Notiophanaeus ) endymion group (214, P. pyrois, allometric variation of head and prothorax of male [right side of pronotum, left side of head, lateral view cephalic process]; 215, tip of elytron and half of pygidium, P. endymion; 216, tip of elytra, P. endymion; 217, same as 215, P. halffterorum; 218, same as 216, P. halffterorum ; 219, P. pyrois, female, front tibia [arrow indicates notch]; 220, same, male). anterolateral angles weakly punctate. Female pro- notum punctate, punctures sometimes effaced me- dially (x 15), coalescing laterally to produce weak rugosity. Except for granulated disk of male (visible to unaided eye as fine texturing), pronotum ap- pearing glassy smooth without magnification. Ely- tra.— Striae fine, interstriae evenly but weakly con- vex, evenly shagreened such that no area appears smoother or brighter than any other. Inner margin of each elytron (Figs. 217, 218) with ridge progres- sively more raised, keel-like posteriorly extending slightly beyond apical angle as small, rounded tooth; apical margin (viewed posteriorly) shallowly excised adjacent to this tooth. Pygidium.— Usually bearing shallow, elongate concavity on each side (often vis- ible only from angle) (Fig. 218). Secondary Sexual Characters. — Male (Fig. 206): Head, pronotum as described for endymion group except that well- developed individuals with posterior angles of pronotal disk more rounded laterally, proportion- ately larger than in other species, anterior margin of pronotum bearing small, acute, median tooth (except in certain Guerrero specimens; see “Com- ments,” below). Female: Cephalic carina trituber- culate. Pronotum bearing weakly raised, anterior transverse ridge with three isolated, equal-sized tu- bercles followed by shallow concavity and midlon- gitudinal linear depression extending from near middle of disk to posterior margin (Fig. 208). Spec- imens Examined. — 19 males, 10 females (length 12-19 mm; width 8-12 mm). DIAGNOSTIC REMARKS. This species differs from endymion, its closest relative, by the follow- ing: pronotal shape and sculpturing of male, shape of inner margin of elytra, shape of female prono- tum, relief of pygidium, and distribution. DISTRIBUTION (Fig. 221; appendix). Known only from the environs of Temascaltepec, state of Mexico, and from the Sierra Madre del Sur from Chilpancingo to Acahuizotla. 750-2400 m. Pine/ oak forests; crepuscular/nocturnal. Mycetopha- gous. Collection dates: June-November. 42 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus COMMENTS. The unique ecological and geo- graphical characteristics of halffterorum were dis- cussed by me in 1979. Since describing this species, I have discovered four of the ten missing specimens of the original Hinton series (see Edmonds, 1979, footnote p. 105); one is in the Howden collection, three in the British Museum (Natural History). The original type series of this species included a large male from the Sierra Madre del Sur near Chilpancingo, Guerrero. I have since examined ad- ditional species from the environs of Acahuizotla, about 40 km west of the Chilpancingo site. These specimens resolve the doubt I had in 1979; the Guerrero populations are, indeed, halffterorum as defined here. They were collected in carrion-baited traps (which attract other essentially mycetopha- gous species) and at lights. Guerrero males are dark green and lack an anteromedian pronotal tooth; females are virtually all black and lack distinct im- pressions on the base of the pygidium. Guerrero populations of halffterorum are isolat- ed from those in the Temascaltepec area by the arid valley of the Balsas River and occur at much lower elevations (ca. 750 m vs. >2000 m). This species is evidently much more ecologically variable than I suspected in 1979. The intervention of the arid conditions of the Balsas Valley, which has un- dergone xerification only recently (Pleistocene), suggests that the origin of halffterorum was earlier than I previously thought. Phanaeus ( Notiophanaeus ) endymion Harold Figures 17, 18, 200-202, 209, 213,215,216,221 Phanaeus endymion Harold, 1863:163 Type: Male lectotype (“Mexico”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:114). OTHER REFERENCES. Bates, 1887; Nevinson, 1892a; Gillet, 1911b; Olsoufieff, 1924; Islas, 1942; Blackwelder, 1944; Halffter and Matthews, 1966; Barrera, 1969; Edmonds, 1972; Moron, 1979, 1984, 1987; Moron et al., 1986; Moron and Fopez-Men- dez, 1985; Moron et al., 1985; Palacios-Rios et ah, 1990. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 17, 18). — Except for irregularly shaped, shining black area on disk of female pro- Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 43 notum (difficult to see in dark specimens), dorsum uniformly dark green or dark blue-green, rarely completely black (Oaxaca specimens); dull to mod- erately shining, brightness often more subdued on elytra, more accentuated on male pronotum; oc- casional females with highly shining, yellow-green pronotum. Color extending ventrally to abdominal sterna, ventral surfaces of femora, tibiae, ptero- pleura, metasternum. Pronotal Sculpturing. — Ap- pearing smooth in both sexes to unaided eye (Figs. 200-203). In male (Fig. 209) ( x 20), disk shagreened, bearing minute asperities that become larger, dens- er, more apparent to naked eye laterally; sides, an- terolateral angles more weakly shagreened than disk, with feeble punctures that become larger, some- what denser anterolaterally; shagreening mutes brightness of surface such that disk appears velvety to unaided eye and under low magnification (x5). Disk of Oaxaca specimen with evenly distributed, minute granulations. In female ( x 20), entire surface punctate, punctures dorsally extremely minute, widely spaced, becoming larger, denser anterolat- erally; to unaided eye, appearing glassy smooth. Elytra. — Striae simple, bearing (xl5) punctures varying from barely perceptible to distinct. Inter- striae distinctly convex (Fig. 213), shagreened; shin- ier (“polished”) medially than adjacent to striae; longitudinal “polishing” most distinct in green in- dividuals, then in first four or five interstriae; in worn specimens, shininess effaced, entire interstria uniformly dull. Elytral margin simple (Figs. 215, 216). Secondary Sexual Characters. — Male: Head, pronotum as described for endymion group, above. Female: Except in worn, small individuals, cephalic carina almost always distinctly trituberculate, only rarely simple. Except in small individuals, prono- tum with small, rounded anteromedian concavity bounded anteriorly by a U- or V-shaped ridge, in few large specimens, posteriorly by small, rounded convexity; anterior ridge bearing median tubercle (dentiform in large specimens), terminating at each end as low rounded or oval tubercle; in some small individuals, concavity only weakly indicated or ab- sent, ridge replaced by three isolated, rounded tu- bercles. Specimens Examined. —137 males, 123 fe- males (length 11-20 mm; width 7-12 mm). DIAGNOSTIC REMARKS. This species most closely resembles F. halffterorum, from which it differs by those features discussed under the latter (see above). Males bear a superficial resemblance to F. adonis (q.v.). DISTRIBUTION (Fig. 221; appendix). Central Honduras, Guatemala, El Salvador, Belize, south- ern Mexico east of the Sierra Madre Oriental from Veracruz to Oaxaca, and southwestern Mexico. Sea level to 2200 m (most records 0-150 m). Coproph- agous; also collected in carrion, fungi, and decom- posing fruits and leaves; diurnal. Collection dates: Mexico — all year; Central America — May-Decem- ber. COMMENTS. The range of endymion is the northern half of Mesoamerica. Its center of distri- bution is southern Mexico where it occurs widely in tropical evergreen and semideciduous forests. There is, however, a disjunct area in southwestern Mexico (western Jalisco and adjacent areas of bor- dering states) where the species is not uncommon; specimens from this area do not differ significantly from those of eastern populations. The intervening region (from Jalisco along the Pacific coast to south- ern Oaxaca) is a drier area where it probably sur- vives in isolated pockets of suitable habitat. The close similarity of eastern and western populations of endymion suggests that this species, until re- cently, also inhabited south-central Mexico. The same factors that resulted in its extinction there (drier climate and associated vegetational changes) may have been also those which promoted speci- ation in relict populations. Such a series of events could explain the existence of halfferorum and of what appears to be a unique Oaxaca population (see below) in areas now peripheral to the distri- bution of endymion. The Oaxaca population mentioned above and referred to in “Descriptive Remarks” is known from two specimens collected by Henry and Anne How- den from the mountains (2150 m) south of the city of Oaxaca. These specimens, a well-developed male and female, differ from typical endymion as fol- lows: they are completely shining black except for subtle remnants of blue coloration along the prono- tal margin (x20); the disk of the male pronotum ( x 15) is evenly, but not densely, covered with small granules, and the pronotum of the female bears a distinct, midlongitudinal impression (like that of halffterorum ). The Howden specimens were col- lected from wild mushrooms in a montane forest, suggesting an ecological distribution similar to that of halffterorum. My guess is that more specimens will support recognizing this Oaxaca population as a distinct species, whose origin is attributable to the same set of circumstances as that of halfftero- rum. While the female pronotum of endymion almost always bears a trituberculate ridge, it sometimes resembles that of pyrois by having three isolated tubercles. Specimens with this atypical pronotal shape almost always also possess another uncom- mon feature: distinctly, broadly punctate elytral striae (xO-5). Neither of these unusual characters, nor the combination of both, varies in any consis- tent geographical way. The lectotype of endymion carries Harold’s label with the notation “Mexico”; his original descrip- tion designates Cordova, Veracruz, as type locality. I have not seen the female cotype described by Harold. Phanaeus (Notiophanaeus) pyrois Bates Figures 19, 20, 203-205, 210, 212, 214, 219-221 Phanaeus pyrois Bates, 1887:58 44 H Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Type: Male lectotype (“Chontales, Nicaragua, T. Belt, B.C.A. p.58 sp.8, sp. figured”), British Museum (Natural History), PRESENT DES- IGNATION. Phanaeus blanchardi Olsoufieff 1924:92 (not Har- old, 1871a:114), NEW SYNONYMY Type: Male lectotype (“Colombia, Cauca River Valley”), Museum National d’Histoire Natu- relle, Paris (Arnaud, 1982a:116). Phanaeus funereus Balthasar, 1939:241, NEW SYNONYMY. Type: Male holotype (“Ecuador”), Narodni Mu- zeum, Prague Phanaeus olsoufieffi Balthasar, 1939:242 (new name for P. blanchardi Olsoufieff, 1924) (Martinez and Pereira, 1967:68) Phanaeus hothrus Blackwelder, 1944:209 (new name for P. blanchardi Olsoufieff, 1924) (Mar- tinez and Pereira, 1967:68) NOMENCLATURAL REMARKS. Olsoufieff s “blanchardi” was first recognized as a junior pri- mary homonym of Phanaeus blanchardi Harold by Balthasar (1939), who proposed the replacement name Phanaeus olsoufieffi. The homonomy was also recognized independently by Blackwelder (1944), whose replacement name Phanaeus bothrus becomes a junior objective synonym of P. olsou- fieffi Balthasar. Thus, the valid name of Olsoufieff’s taxon is Phanaeus olsoufieffi Balthasar. P. olsou- fieffi (as blanchardi Olsoufieff) was first synony- mized with P. funereus by Martinez and Pereira (1967). The name “apollinaris Gillet” has been used for pyrois; as far as I know, it is a nomen nudum. OTHER REFERENCES. Nevinson, 1889, 1892a; Pittier and Biolley, 1895; Gillet, 1911b; Gacharna, 1951; Edmonds, 1972, 1978; Young, 1978; How- den and Young, 1981; Peck and Forsyth, 1982 (as “apollinaris”). DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 19, 20). — Usually as follows: Poste- rior two-thirds of clypeus, paraocular areas, if not all at least sides and anterolateral angles of pro- notum bright metallic red; all non-colored areas dull or weakly shining black; elytra entirely dull black with at most very weak bluish luster (usually visible only under magnification with strong light); pygidium bright red medially, green around margin. Following variations known: (a) certain specimens from western Panama, Costa Rica, and Colombia, all specimens examined from Ecuador (“funereus”) almost entirely black; colored areas very incon- spicuous, restricted to pronotal, pygidial margins and often visible only under magnification ( x 10); (b) certain specimens from central Costa Rica (Tur- rialba) completely dull, dark metallic blue or weak- ly shining green; elytra distinctly duller than pro- notum (most Turrialba specimens exhibit usual coloration described above); and (c) any dorsal col- ored areas, regardless of size or location, in occa- sional specimens can be red-yellow or red-green. Venter black to black-brown, usually with distinct, weak greenish or bluish reflections on legs, abdom- inal sterna, other isolated areas. Pronotal Sculp- turing.— To unaided eye pronotum of both sexes appears smooth (Figs. 203, 204); magnification ( x 20) reveals following: In male (Fig. 210) flattened, tri- angular disk shagreened, bearing minute asperities which become larger, denser laterally; rounded sides, anterolateral angles with feeble punctures becom- ing somewhat larger, denser anterolaterally; sha- greening tends to mute brightness of surface such that disk appears almost velvety to unaided eye or under low magnification ( x 5). In female, entire sur- face punctate, punctures extremely minute, widely separated dorsally, becoming larger, denser later- ally, anteriorly. Pronotum. — Basal fossae reduced to punctiform pits in both sexes. Elytra (Fig. 212). — Striae fine but apparent width enhanced somewhat by usually weakly convex, dull interstriae; in some specimens (particularly large ones) interstriae vir- tually flat (frequency of specimens with flat, uni- formly dull interstriae higher in South American populations than in Central American ones). Inter- striae usually extremely minutely, evenly sha- greened (x80) with no difference in reflectivity be- tween median areas, those adjacent to striae. In some specimens viewed with unaided eye, median portions of interstriae 1-4 appear somewhat shinier such that their apparent convexness is accentuated; any variability in reflectivity visible to unaided eye disappears under higher magnifications ( x 15 + ), in- terstriae appearing uniformly dull. All interstriae smooth except for lateral-most, which bears few, scattered, small basal punctures. Secondary Sexual Characters. — Male: Head, pronotum as described for endymion group above. Female: Cephalic carina at most only weakly trituberculate, often simple carina. Pronotum (Fig. 205) evenly convex, always bearing three low, round tubercles arranged in transverse line, middle one often lying before or behind imaginary line connecting outer two. Spec- imens Examined. — 127 males, 1 16 females (length 11-20 mm; width 7-12 mm). DIAGNOSTIC REMARKS. This species most closely resembles endymion, from which it is dis- tinguished by the following combination of char- acters: color and color pattern, elytral sculpturing, female secondary sexual characters of the prono- tum, and distribution. DISTRIBUTION (Fig. 221; appendix). Southern Nicaragua (north of Lake Nicaragua), Costa Rica, Panama, Colombia west of and along east side of Andes, and Ecuador east of the Andes. Sea level to 2200 m (most records 0-1500 m). Apparently re- stricted to forest habitats; diurnal. Coprophagous but sometimes collected from carrion and decom- posing fruit. Collection dates: Panama — all year; Costa Rica — May-August; Colombia — May-Oc- tober; Ecuador — March-September. Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 45 COMMENTS. As here defined, pyrois includes two previously recognized species based primarily on differences in color pattern, P. olsoufieffi and funereus. Were it not for the fact that specimens virtually identical to funereus occur in Colombian and Central American populations of pyrois, I would be inclined to maintain funereus as a subspecies. The type of olsoufieffi, as well as similar specimens from Central America, fall within the normal range of variation of pyrois and likewise do not merit separate taxonomic status. The dark blue and dull green specimens of pyrois from Turrialba, Costa Rica, resemble endymion closely enough to suggest that they are intermedi- ates between the two otherwise distinct species. I have not, however, seen any endymion from Nic- aragua; moreover, the specimens in question were collected along with perfectly “good” pyrois. Nev- ertheless, the question of the relationship between the two species is somewhat clouded by the Tur- rialba specimens. The British Museum (Natural History) collection includes the two specimens upon which Olsoufieff (1924) based the “aberration viridicollis ” of his P. blanchardi. Both are labeled “Balzar Mountains], Ecuador.” The male is small; its head and pronotum are bright, shining green, the elytra dark, almost black with some greenish tinges, and pygidium shin- ing green. The female is like the male except that the head and pronotum are blue-green. The spec- imens appear to be composites resulting from mis- matching head-prothorax sections and pterotho- rax-abdomen sections (which remain on pins when the former break free). I cannot, however, clearly detect any repair, although there is glue beneath the right elytron of the male (but none visible be- tween the pro- and metathorax). These specimens are unique, and, if it can later be determined that they are members of a morphologically and geo- graphically distinct population, OlsoufiefPs “viri- dicollis” may prove to be a valid taxon. The distribution of pyrois covers the southern half of Mesoamerica. I consider it the sister group of endymion and its relatives, which occupy the northern half. Subgenus Phanaeus Macleay Phanaeus Macleay, 1819:124. Type-species: ( Scar - abaeus carnifex L., 1767) = Phanaeus vindex Macleay, 1819; subsequent designation by Ol- soufieff, 1924. Lonchopborus Germar, 1824:106. Type-species: (. Scarabaeus carnifex L., 1767) = Phanaeus vin- dex Macleay, 1819; subsequent designation by Edmonds, 1972. Onthurgus Gistel, 1857:602. Type-species: ( Scar- abaeus carnifex L., 1767) = Phanaeus vindex Macleay, 1819; original designation. Palaeocopris Pierce, 1946:130. Type-species: Pa- laeocopris labreae Pierce, 1946; original desig- nation by monotypy. Phanaeus, sen. str., differs from Notiophanaeus primarily on the basis of pronotal sculpturing, which in this subgenus is granulate or granulorugose an- terolaterally (see discussion under Notiophanaeus). This subgenus comprises 31 species and subspecies arranged in eight species groups. Except for lunaris, hermes, and prasinus, all species and subspecies are endemic to either Central or North America. The primary characters separating species groups relate to pronotal sculpturing, shape of the longer me- sotibial spur (especially in females), shape of the female pronotum, and geographic distribution. In general, separation of species on morpholog- ical grounds in this subgenus is more difficult than in Notiophanaeus. This is especially true for the mexicanus and tridens groups, females and small males of which can be virtually impossible to dis- tinguish. KEY TO THE SPECIES GROUPS OF PHANAEUS , SEN. STR. la. Pronotum granulate anterolaterally (Figs. 224, 236, 253). Longer mesotibial spur not dilated subapically (Fig. 245). Basal segment middle tarsus of female not strongly dilated 2 b. Pronotum granulorugose or rugose anterolat- erally (Figs. 274, 310, 319, 320). Longer meso- tibial spur dilated subapically (Fig. 289), es- pecially in females (except flohri and some scutifer). Female mesobasitarsus variable . . 4 2a. Front tibiae tridentate. Anterolateral pronotal granules of variable sizes. Elytral interstriae finely, but distinctly (x 10) punctate (Figs. 254, 255). United States and northeastern Mexico (Fig. 256) triangularis group b. Front tibiae quadridentate (Figs. 243, 244), bas- al tooth small (can be indistinct in worn spec- imens). Anterolateral pronotal granules more- or-less uniform in size, usually dense. Elytral interstriae only minutely punctured ( x 40) (Figs. 225, 226). Northwest Mexico to northwestern South America 3 3a. Pronotal disk of well-developed males (Figs. 228-235) concave at least anteriorly, usually coarsely rugose, bordered laterally by raised, usually jagged ridges, and bearing a central or posteromedian process of some type (Figs. 237- 242). Northwestern Mexico to Costa Rica (Fig. 246) tridens group b. Pronotal disk of well-developed males flat and uniformly granulate (Fig. 222). Northwestern South America to Panama (Fig. 227) hermes group 4a. Pronotum of female with wide anteromedian prominence flanked on each side by elongate, vertical depression extending from disk to an- terior pronotal margin behind eye (Figs. 273, 274). Extreme southwestern United States, Mexico, Central America, and northwestern South America (Fig. 290) . . mexicanus group b. Pronotum of female more-or-less evenly con- vex anteriorly, with or without small antero- 46 ^ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Figures 222-226. Pbanaeus ( Phanaeus ) hermes group (222, P. prasinus, male; 223, same, female; 224, same, oblique view female pronotum; 225, P. hermes, dorsal view elytron; 226, P. prasinus, same). median concavity and tubercles, and lacking vertical depressions 5 5a. Midventral carina of front femora angulate near base; base of posterior surface with large, coarse punctures (Fig. 297). Pronotum of large males (Figs. 291-293) with smooth posteromedian area bounded anteriorly by transverse tubercle or carina. Pronotum of female (Fig. 294) with narrow, transverse anteromedian ridge fol- lowed by oval concavity whose posterior mar- gin interrupted by low, rounded gibbosity. Southeastern Mexico to Panama (Fig. 301) . . heltianus group b. Front femora not angulate at base (Fig. 298); basal puncturing same size and texture as that extending to apex of femora. Male pronotum variable, but not as above 6 6a. Pronotum finely granulorugose anterolaterally. Pronotum of large males with smooth raised area along posterior margin which is often toothed medially (Figs. 307-309). Cephalic ca- rina of female (seen from above) even with anterior margin of eyes. Eastern and southern Mexico to Guatemala (Fig. 312) amethystinus group b. Pronotum (Figs. 319, 320, 336-340) very densely and coarsely granulorugose anterolat- erally, and usually also over entire pronotal surface (sculpturing effaced in igneus). Male pronotum variable, but not as above. Cephalic process of female (seen from above) positioned in front of eyes. Central and western Mexico, United States 7 7a. Elytral interstriae smooth or minutely punctate (x30) (Figs. 321, 322). Outer margin of head notched between clypeus and paraocular areas (Fig. 325). Central and western Mexico and extreme southwestern United States (Fig. 329) quadridens group b. Elytral interstriae densely and coarsely punc- tatorugose (Figs. 341-346). Outer margin of head not distinctly notched. Extreme northern Mexico, United States (Fig. 360) vindex group The Hermes Group DIAGNOSIS. [1] Anterior margin of clypeus at most only weakly bidentate, often rounded; [2] clypeal process (seen from front) a rounded ridge; [3] cephalic carina of female at most only weakly tridentate; [4] pronotal disk of large males (Fig. 222) flattened dorsally, posterior angles curved laterally and sharply upturned, anterior angles each bearing conical tubercle; [5] pronotum of female evenly convex, with bowed transverse carina near anterior margin, sometimes weakly toothed medially and followed by a weak concavity; [6] anterolateral por- tions of pronotum evenly covered by flattened scale- like granules (Fig. 224); [7] basal pronotal fossae present; [8] front tibiae quadridentate, basal (fourth) tooth small, apical two teeth longitudinally cari- nate; [9] longer mesotibial spur slender, not dilated subapically; [10] elytral striae punctate (Figs. 225, 226); [11] forests of Panama and northwest South America (Fig. 227). This group consists of two closely related species, hermes Harold and prasinus Harold (see also dis- cussion under the triangularis group). Contributions in Science, Number 443 Edmonds: Revision of Phanaeus B 47 hermes group hermes prasinus Figure 227. Distribution of the Phanaeus ( Pbanaeus ) hermes group. KEY TO THE SPECIES OF THE HERMES GROUP la. Elytral interstriae dull, appearing almost flat (Fig. 225); median area not noticeably shinier than that adjacent to striae; punctures of striae visible to unaided eye. Dorsum weakly shining to bright coppery brown or coppery green (Figs. 32, 33). Northwestern Colombia to Costa Rica (Fig. 227) .... Phanaeus (P.) hermes Harold b. At least first three, but usually all interstriae evenly shining or shining only midlongitudi- nally and appearing convex (Fig. 226). Punc- tures of elytral striae very small, sometimes effaced, clearly visible only under magnification (xlO). Dorsum weakly shining dark green, black, or, rarely, coppery green (Figs. 34-36). Northwestern Colombia and western and northern Venezuela (Fig. 227) Phanaeus (P.) prasinus Harold Phanaeus (Phanaeus) hermes Harold Figures 32, 33, 225, 227 Phanaeus hermes Harold, 1868b:82 Type: Male lectotype (“Columbia” [sfc]), Mu- seum National d’Histoire Naturelle, Paris (Ar- naud, 1982a:115). Phanaeus bogotensis Kirsch, 1871:359, NEW SYNONYMY Type: Male lectotype (“Bogota”), Staatsliches Museum fur Tierkunde, Dresden, PRESENT DESIGNATION. NOMENCLATURAL REMARKS. Harold (1880b) synonymized bogotensis and prasinus; Kirsch’s species, however, is hermes. OTHER REFERENCES. Gemminger and Har- old, 1859; Bates, 1887; Nevinson, 1892a; Pittierand Biolley, 1895; Gillet, 1911b; Olsoufieff, 1924; Blackwelder, 1944; Gacharna, 1951 (as “oliva- ceus”); Vulcano and Pereira, 1967; Edmonds, 1972; Howden and Young, 1981. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 32, 33).- — Dorsum weakly shining to bright coppery brown, usually with green or yellow- green highlights on head, pronotum, margins of elytra; pronotum of female with central black spot that can cover most of disk; females usually darker, duller than males. Pygidium colored as dorsum. Venter dark with strong green reflections on legs, sterna. Pronotal Sculpturing. — In large males, gran- ulate except for smoother, strongly punctate tri- angular area adjacent to posterior margin, for punc- tate areas beneath posterolateral angles; granulation strongest, best defined on disk, becoming weaker, scale-like laterally, where it forms some ridging. In females, small males, anterior one-half to two-thirds of surface granulate, granules becoming scale-like anterolaterally, in females, coalescing dorsally to form distinct rugosities; basal one-third to one-half strongly punctate, most weakly so posteromedially. Elytra (Fig. 225). -—Striae punctate, punctures small but clearly visible to unaided eye. Interstriae almost flat, minutely punctured (xlO), dull. Secondary Sexual Characters. — As described for the species group. Specimens Examined. — 57 males, 40 fe- males (length 12-20 mm; width 7-14 mm). DIAGNOSTIC REMARKS. This species differs from prasinus in color and elytral relief and texture. Small individuals can resemble chalcomelas, which 48 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus has very large strial punctures and tridentate front tibiae. DISTRIBUTION (Fig. 227; appendix). Moun- tain forests (400-1000 m) of northwestern Colom- bia (including interandean valleys), Panama, and Costa Rica. Coprophagous. Collection dates: May- August, December. COMMENTS. The eastern distributional limit of hermes appears to be the Cordillera Oriental of the Andes, where it overlaps prasinus. Locality rec- ords are scattered, but I suspect that this species inhabits montane forests throughout northwestern Colombia. Phanaeus ( Phanaeus ) prasinus Harold Figures 34-36, 222-224, 226, 227 Phanaeus prasinus Harold, 1868a:83 Type: Male lectotype (“Venezuela”), Museum National d’Histoire Naturelle, Paris (Arnaud, 1982a:115). Phanaeus lugens Nevinson, 1889:179, NEW SYN- ONYMY Type: Male lectotype (“Venezuela, Merida”), Museum National d’Histoire Naturelle, Paris (Arnaud, 1982a:116). OTHER REFERENCES. Harold, 1868b; Can- deze, 1891; Nevinson, 1892a; Gillet, 1911b; Sharp and Muir, 1912 ( lugens ); Olsoufieff, 1924; Black- welder, 1944; Gacharna, 1951; Roze, 1955; Vul- cano and Pereira, 1967; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 34-36). — Dorsum either dull to weakly shining black (“lugens”) or dark to bright green to blue-green rarely with coppery highlights; pronotal disk of green females with central black spot that can virtually cover disk. Pygidium colored as dorsum. Venter black or with dark green reflec- tions on legs, sterna. Pronotal Sculpturing. — As de- scribed for hermes except that in black females, puncturing less pronounced. Elytra (Fig. 226). — Striae punctate ( x 10), punctures scarcely if at all visible to unaided eye. Interstriae weakly convex, minutely punctate; surfaces evenly shining or shin- ing at least midlongitudinally. Secondary Sexual Characters. — As described for the species group. Specimens Examined. — 106 males, 91 females (length 12-20 mm; width 7-13 mm). DIAGNOSTIC REMARKS. The black females of prasinus can superficially resemble dark females of triangularis while dark green individuals can resemble similarly colored females of igneus (see also “Comments” under hermes). DISTRIBUTION (Fig. 227; appendix). Forests of northern Colombia, western and northern Ven- ezuela, and isolated localities in the Gran Sabana of Venezuela. 0-2000 m. Coprophagous. Collec- tion dates: January-October. COMMENTS. Nevinson’s species differs from prasinus only in color, and I have concluded for now that the two are best considered synonyms. The synonymy is open to some question, an answer to which will depend upon a greater quantity of more precise locality data than I have now. Most available “lugens” bear imprecise locality data, and there are few series or even pairs collected from the same place. I know of no locality where black and green forms occur together; moreover, there are few specimens intermediate between the two. The green form is more common in collections. Most specimens are from mid-elevation localities in the Cordillera Oriental of the Colombian Andes. Some are from the coastal mountains and lowlands of northern Venezuela, and a few from scattered localities in the Gran Sabana of central Venezuela. These latter localities are on rivers and suggest that prasinus follows the gallery forests into that region. I know the black form from the Cordillera de Me- rida (Venezuela), some specimens labeled “Bogota” (see below), and from Trinidad. Trinidad individ- uals can be black, but more often they have a dis- tinct blue cast on the upper surface and are shinier and smaller than black specimens from the main- land. I have seen specimens of both forms of prasinus as well as hermes labeled “Bogota.” Their occur- rence there needs confirmation. Among a series of specimens from gallery forest near Ciudad Bolivar (Venezuela) are green individuals with strong cop- pery reflections that make them resemble hermes in color. The Tridens Group DIAGNOSIS. [1] Outer margin of head not dis- tinctly notched laterally; [2] shape of clypeal pro- cess variable; [3] cephalic carina of female weakly tridentate, set in front of eyes; [4] anterolateral por- tions of pronotum (Fig. 236) granulate, granules small, uniformly distributed, fairly dense, only oc- casionally confluent (around lateral fossa); [5] basal pronotal fossae present; [6] disk of pronotum of large males concave at least anteriorly, usually coarsely rugose, bearing a median or posteromedian process whose shape and position varies by species (Figs. 237-242) (trace of this process persists even in smallest individuals); sides usually raised as jagged ridges; posterolateral angles (seen from behind) rounded, upturned; [7] pronotum of female con- vex, bearing thick, black, transverse anteromedian ridge usually followed by shallow concavity (ridge and concavity obliterated in small individuals); [8] front tibiae quadridentate (Figs. 243, 244); [9] fe- male mesobasitarsus narrow (Fig. 245); [10] longer metatibial spur not dilated subapically (Fig. 245); [11] dorsum usually completely green or greenish, occasionally dark blue or coppery red (Figs. 86- 95); [12] smaller-sized, length rarely exceeding 15 mm; [13] central Sonora southward along Pacific coast of Mexico, southern Mexico, and Central America from Guatemala to Costa Rica (Fig. 246). Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 49 Figures 228-236. Phanaeus ( Phanaeus ) tridens group (228, P. tridens, male; 229, same; 230, P. furiosus; 231, P. daphnis, male; 232, same; 233, P. nimrod, male [mouthparts removed]; 234, same, female; 235, P. eximius, male; 236, P. furiosus, female, lateral view pronotum). The tridens group embraces five closely related species, tridens, eximius, nimrod, daphnis, and fu- riosus. Of those characters listed above, the sculp- turing of the pronotum and form of the pronotum of large males are the most distinctive. The species themselves, however, are difficult to separate mor- phologically. The most consistent characters are details of the shape of the pronotum of large males and geographic distribution. Females and small males can be virtually impossible to identify if un- accompanied by precise locality data. In general, the combination of distribution, male pronotal fea- tures, and some other morphological character or combination of characters will distinguish 90% of all specimens. For each species, there is a combi- nation of features that, while “usual,” is subject to intraspecific variation; this fact should be taken into account when using the key presented below. The observation that the members of the tridens group are not separated by clean morphological gaps indicates that the group is likely the product of rapid speciation. Of special interest is the Colima population of tridens because it suggests a close phylogenetic connection among itself, daphnis, and furiosus. This group is equivalent to “cluster 8” of Phan- aeus that I recognized in 1972 with the addition of eximus; at that time, this latter species was assigned to “cluster 5” (now the quadridens group). Avail- able distributional data suggest that all members of the tridens group are allopatric. But two areas, in particular, must be more thoroughly sampled be- fore some zones of sympatry can be ruled out: central Michoacan (where daphnis and furiosus may overlap) and the Chiapas-Guatemala border region (where tridens may contact eximius). KEY TO THE SPECIES AND SUBSPECIES OF THE TRIDENS GROUP la. Clypeal process (seen from front) very narrow, acute, distinctly longer (usually much longer) than width at base. Posteromedian process of male pronotum (Fig. 240) very large, laterally flattened, triangular ridge (Fig. 233). Female al- most always with small, round black spot be- neath lateral pronotal fossa. Elytral interstriae usually distinctly shinier midlongitudinally 50 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Figures 237-245. Phanaeus ( Phanaeus ) tridens group (237, dorsal view posterior portion male pronotum, P. furiosus; 238, same, P. tridens ; 239, same, P. daphnis ; 240, same, P. nimrod; 241, same, P. eximius ; 242, same, P. daphnis [“tricornis” form]; 243, P. tridens, male front tibia; 244, P. eximius, female front tibia; 245, P. tridens, tip of mesotibia and basitarsus). (uniformly dull in worn specimens). Higher val- leys of central Oaxaca above 1400 m, and Pa- cific coast of Oaxaca (Fig. 246) Phanaeus (P.) nimrod Harold b. Shape of clypeal process variable, but almost always broader than long, never appearing con- ical. Not occurring in central or coastal Oa- xaca. Other characters variable 2 2a. Clypeal process (seen from front) triangular, or weakly rounded with small median tooth. Pos- teromedian process of male pronotum usually elongate, apically bifurcate, spine-like projec- tion (Fig. 238). Female lacking black spot be- neath lateral pronotal fossa. Distribution as in Figure 246 Phanaeus (P.) tridens Laporte-Castelnau 3 b. Clypeal process (seen from front) usually rounded; if more-or-less triangular (some daphnis ), then male pronotum as in Figures 231, 232. Other characters variable 4 3a. Elytral interstriae uniformly shining. Southern Chiapas, extreme southwestern Guatemala . . . Phanaeus (P.) tridens pseudofurcosus Balthasar b. Elytral interstriae distinctly shinier rnidlongitu- dinally. Lowlands of central Veracruz and Co- lima Phanaeus (P.) tridens tridens Laporte-Castelnau 4a. Elytral interstriae ( x 30) usually minutely ru- gose or punctatorugose adjacent to striae (sometimes visible only apically), smooth me- dially, appearing strongly convex to unaided eye. Posteromedian process of male pronotum a short, conical tooth (Fig. 241). Pronotal disk of male with pair of conical tubercles near anterior margin. Southern Guatemala to cen- tral Costa Rica (Fig. 246) Phanaeus (P.) eximius Bates b. Elytral interstriae uniformly shiny or dull; oc- casionally shagreened adjacent to striae, shinier medially, but never minutely rugose-puncta- torugose adjacent to striae. Male pronotum not as described above. Not occurring in Cen- tral America 5 5a. Posteromedian process of male pronotum broad, apically emarginate projection (Fig. 237); raised sides of disk lacking strong conical tooth; granulations of sides of pronotum reaching posterior margin. Clypeal process rounded. Elytral interstriae convex. West-central Mex- Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 51 ico, extending along Pacific coastal region northward into central Sonora (Fig. 246) . . . Phanaeus ( P .) furiosus Bates b. Posteromedian process of male pronotum a short conical tooth (Fig. 239), or long, some- times weakly apically bifurcate, spine-like pro- jection (Fig. 242); raised sides of disk each with strong, conical tooth (Figs. 231, 232). Clypeal process usually rounded, sometimes triangular. Elytral interstriae flat to weakly convex. Balsas River valley and peripheral areas along escarp- ment of Transverse Volcanic Range of south- ern Mexico (Fig. 246) Phanaeus (P.) daphnis Flarold Phanaeus ( Phanaeus ) nimrod Harold Figures 90, 233, 234, 240, 246 Phanaeus nimrod Harold, 1863:167 Type: Male lectotype (“Mexico”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:115). Phanaeus bahori Balthasar, 1939:246 (Arnaud, 1982c:125) Type: Male holotype (“Mexiko: Oaxaca”), Na- rodni Muzeum, Prague. OTHER REFERENCES. Gemminger and Har- old, 1869; Bates, 1887; Nevinson, 1892a; Villada, 1901; Gillet, 1911b; Olsoufieff, 1924; Islas, 1942; Blackwelder, 1944; Edmonds, 1972 (references to nimrod in Halffter and Matthews, 1966, should read daphnis, q.v.). DESCRIPTIVE REMARKS. Color and Color Pattern (Fig. 90). — Dorsum dark blue, blue-green, green, or yellow-green; yellow-green sometimes combined with coppery highlights. Head.— Clyp- eal process long, acute, spine-like, longer, usually much longer, than width at base. Pronotum. — Bas- al fossae present. Base of female disk impressed midlongitudinally. Pronotal Sculpturing.— Male: In large individuals, sides granulate to finely gran- ulorugose; disk smooth to finely punctate anteri- orly, with sparse but coarse rugosities posteriorly. Smaller individuals, as above except disk granulate to granulorugose anteriorly, punctate posteriorly. Female: Sides granulate; disk granulorugose ante- riorly, granules progressively weaker posteriorly un- til replaced by sparse puncturing near posterior margin. Elytra. — Striae fine, impressed, minutely punctured (x30). Interstriae convex, minutely 52 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus punctate (x30), shagreened; shagreening usually denser adjacent to striae such that interstriae (es- pecially 1-3) appearing smoother, shinier midlon- gitudinally under weak magnification (xlO). Sec- ondary Sexual Characters. — Male: Pronotum as in Figures 233, 240. Female: As described for species group. Specimens Examined. — 64 males, 49 fe- males (length 11-16 mm; width 7-10 mm). DIAGNOSTIC REMARKS. The shape of the clypeal process and male pronotum along with dis- tribution will separate nimrod from other members of the group. DISTRIBUTION (Fig. 246; appendix). Central valley of Oaxaca and adjacent areas to south and southwest, and Pacific coast of Oaxaca. 0-1800 m. Coprophagous. Collection dates: May-September. COMMENTS. The known distribution of nim- rod is the smallest among members of the species group. Other than records from El Ocotito in cen- tral Guerrero and from Puerto Angel on the Pacific coast of Oaxaca, it is restricted to the central valleys of Oaxaca. Its presence in the coastal scrub forest could be the result of recent introduction from the interior. In spite of its limited distribution, color varies considerably. As in eximius, no one color predominates, and small series from single localities can include all variations. This species resembles tridens tridens in the microsculpture of the inter- striae. Phanaeus ( Phanaeus ) tridens Laporte-Castelnau Figures 91-93, 228, 229, 238, 243, 245, 246 Phanaeus tridens Laporte-Castelnau, 1840:81 Type: Male neotype (erroneously labeled “Chi- li”), Castelnau Collection, National Museum of Victoria, Melbourne, PRESENT DESIG- NATION. Phanaeus frankenbergeri Balthasar, 1939:245 (Ar- naud, 1982c:125) Type: Male holotype (“Mexico”), Narodni Mu- zeum, Prague. Phanaeus pseudofurcosus Balthasar, 1939:244 Type: Male holotype (“Mexico”), Narodni Mu- zeum, Prague. Phanaeus tridens tridens Laporte-Castelnau, NEW STATUS Phanaeus tridens pseudofurcosus Balthasar, NEW COMBINATION NOMENCLATURAL REMARKS. Hope’s (1837:53) suspicion that Scarabaeus tridens Fabri- cius, 1775 ( Copris tridens Fabricius, 1801), be- longed in Phanaeus created the possibility that La- porte-Castelnau’s name was a junior homonym. Fa- bricius’ species, however, is an Onthophagus. OTHER REFERENCES. Klug, 1841 (as “nep- tunus”); Lacordaire, 1856; Harold, 1863; Gemmin- gerand Harold, 1869; Bates, 1887; Nevinson, 1892a; Villada, 1901; Gillet, 1911b; Olsoufieff, 1924; Islas, 1942; Blackwelder, 1944; Halffter and Matthews, 1 966; Barrera, 1 969; Edmonds, 1 972; Edmonds and HalfFter, 1978; Halffter and Edmonds, 1982. DESCRIPTIVE REMARKS. Color and Color Pattern. — (a) tridens (Figs. 91-93): Posterior por- tion of head, pronotum green, yellow-green, cop- pery yellow, coppery red, dark black-red; sides of pronotum (especially males) usually contrasting with that of disk, sometimes tricolored; elytra green, yellow-green or coppery red with green highlights, usually less brightly colored than pronotum. More common color combinations: (i) pronotal disk bright coppery red, sides yellow-green, elytra yellow-green; (ii) pronotal disk yellow-green, elytra, sides of pro- notum deeper green, (b) pseudofurcosus: Much less variable, contrasting than above, usually as in com- bination (ii) above. Head. — Clypeal process (seen from front) triangular, or weakly rounded with small median tooth. Pronotal Sculpturing. — Male: In larger individuals, sides finely granulate becoming granulorugose on raised outer margin of disk; disk weakly but coarsely rugose, more sparsely so pos- teriorly. In smaller individuals, remains of central part of disk sparsely, irregularly granulate, becom- ing punctate posterolaterally. Female: Sides finely granulate, granules becoming larger, flattened on disk, progressively more effaced posteromedially where replaced by asperate punctures. Elytra. — Striae simple, impressed, minutely punctured ( x 30). In tridens, interstriae 1-4 (or 5) shinier medially than along striae, appearing polished midlongitu- dinally. In pseudofurcosus, interstriae uniformly shining. Secondary Sexual Characters. — Male: Pronotum as in Figures 228, 229, 238; postero- median process of disk usually elongate, apically bifurcate, spine-like projection (except Colima pop- ulation of tridens; see “Comments,” below). Fe- male: As described for species groups. Specimens Examined. — tridens — 237 males, 196 females; pseudofurcosus — 67 males, 74 females (length 10- 17 mm; width 7-11 mm). DIAGNOSTIC REMARKS. The shape of the clypeal process will separate this species from all but a few daphnis. Certain males of Colima pop- ulations superficially resemble “tricornis” males of daphnis. DISTRIBUTION (Fig. 246; appendix), tri- dens— tropical lowlands of central Veracruz and Colima. 0-1500 m. pseudofurcosus — southern Chiapas and extreme southwestern Guatemala. 0- 1800 m. Both subspecies — Coprophagous. Collec- tion dates: May-October (most June-August). COMMENTS. Specimens of tridens tridens from the northern limit of its range in Veracruz usually have yellow-green pronota and green elytra. Else- where, this subspecies usually has a coppery red pronotal disk and light green elytra with strong yellow or coppery highlights. Most pseudofurcosus are uniformly dark green. The Colima population of tridens tridens exhib- Contributions in Science, Number 443 Edmonds: Revision of Phanaeus M 53 its a surprising amount of variability in the shape of the posteromedian process of the male prono- tum. Certain males resemble those of nimrod, and others look like the “tricornis” males of daphnis. I have seen only 20 specimens from Colima; of these, four are large males, and none has a typical tridens pronotum. This population may later prove to be a distinct taxon, but its relationship with tridens from eastern Mexico is clear. The Colima population of tridens, along with disjunct south- west Mexico populations of endymion, are prob- ably survivors of broadly distributed tropical forest populations that antedated the onset of desert con- ditions in the Balsas River valley and southern coastal regions of Mexico. I have seen one specimen of this subspecies collected in Tequesquitengo, Mo- relos, far from the Colima and Veracruz popula- tions. In order to rule out an accidental introduc- tion in modern times, more specimens must be collected. The original type material of tridens is apparently lost (see the introduction). I have designated a spec- imen of tridens from the Laporte-Castelnau col- lection in Melbourne as neotype. It is mounted with a small male of scutifer and both are labeled “tri- dens Chili.” Phanaeus (Pbanaeus) eximius Bates Figures 86, 235, 241, 244, 246 Pbanaeus eximius Bates, 1887:62 Type: Male lectotype (“Guatemala, Salle, B.C.A. p.62 sp. 22, sp. figured”), British Museum (Nat- ural History), London, PRESENT DESIG- NATION. OTHER REFERENCES. Bates, 1889; Nevinson, 1892a; Gillet, 1911b; Olsoufieff, 1924; Blackwelder, 1944; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Fig. 86). — Dorsum dark blue, blue-green, green, yellow-green, or coppery red. Pygidium same color as elytra. Venter dark, colored reflections on legs, sterna. Head.— Clypeal process semicircular or broadly rounded, never angulate. Pronotum. — Base of disk impressed in female. Basal fossae pres- ent. Pronotal Sculpturing. — Male: In large individ- uals, sides granulate, disk more or less evenly, coarsely rugose except posteromedially where coarsely punctatorugose. Small individuals as above except disk granulorugose anteriorly finely punc- tatorugose to punctate posteriorly. Females: Gran- ulate laterally; granules becoming flat on disk, co- alescing anteriorly as transverse ridges, becoming progressively weaker posteriorly where they may become asperate punctures; posterior portion of pronotum never simply punctured. Elytra.— Striae simple, impressed, minutely punctured (x30). In- terstriae convex, distinctly but minutely ( x 30) ru- gose-puntatorugose adjacent to striae, smooth me- dially; interstrial rugosity sometimes distinct only apically. Secondary Sexual Characters.— Males: Pronotum as in Figures 235, 241. Female: As de- scribed for species group. Specimens Examined. — 35 males, 27 females (length 12-17 mm; width 7- 11 mm). DIAGNOSTIC REMARKS. This species most closely resembles tridens, from which it differs by shape of the clypeal process, sculpturing of the elytral interstriae and of the male pronotum, and distribution. DISTRIBUTION (Fig. 246; appendix). Central America south of the central cordilleras from Gua- temala to Costa Rica. 300-1800 m. Coprophagous. Collection dates: May-August. COMMENTS. The elytral sculpturing charac- teristic of eximius is sometimes indistinct; in these cases, the interstriae might be described as “coarsely shagreened.” Phanaeus eximius resembles pallia- tus, and I grouped the two species in 1972. The two differ by pronotal sculpturing. The color vari- ation of eximius is great among the few specimens examined. The distribution of eximius is poorly known; well over one-half of the specimens examined lack precise locality data. Pbanaeus ( Pbanaeus ) furiosus Bates Figures 87-89, 230, 236, 237, 246 Phanaeus furiosus Bates, 1887:61 Type: Male lectotype (“Ventanas, Mexico, 2000ft, Forrer, Hoege, B.C.A. p.61 sp.19, sp. figured”), British Museum (Natural History), PRESENT DESIGNATION. Pbanaeus furcosus Felsche, 1901:155 (new name for furiosus ), NEW SYNONYMY NOMENCLATURAL REMARKS. In his orig- inal description, Bates said, “I have adopted the name ‘P. furiosus Duges,’ given to this species in the Salle collection.” Felsche (1901), who was in- formed by Salle that the name in his collection was furcosus, not furiosus, emended the name accord- ingly. Gillet (1911b) and Olsoufieff (1924) accepted Felsche’s emendation, and the name furcosus has been used widely in collections for the last 60 years. It is my opinion that Felsche’s action was an un- justified emendation (Article 33[a] [ii] of the Code) and, therefore, that Phanaeus furiosus Bates is the valid name of this species. Accordingly, P. furcosus becomes a junior objective synonym of P. furiosus Bates. OTHER REFERENCES. Nevinson, 1892a; Blackwelder, 1944; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 87-89).— Dorsum dark blue, shining blue-green, or green; rarely shining yellow-green with coppery highlights. Pygidium colored as elytra. 54 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Venter dark with colored reflections on legs, sterna. Head. — Clypeal process rounded. Pronotum. — Basal fossae present. Pronotal Sculpturing. — Male: Sides granulate; granulation extending to posterior margin, near which it can become somewhat coars- er. Disk of larger individuals becoming coarsely rugose in front of posteromedian process; disk of smaller individuals granulate to granulorugose an- teriorly, weakly punctatorugose posteriorly. Female (Fig. 236): Sides granulate; disk granulate anteriorly, granules becoming larger, scale-like medially, weakly punctatorugose along posterior margin. Elytra. — Striae fine, minutely punctate ( x 30). Interstriae con- vex, with variable sculpturing (xlO): very finely roughened, weakly shining, to almost entirely smooth, brightly shining. Secondary Sexual Char- acters.—Male: Pronotum as in Figures 230, 237. Female: As described for species group. Specimens Examined. — 415 males, 403 females (length 11-18 mm; width 7-11 mm). DIAGNOSTIC REMARKS. Large males differ from other members of the group by pronotal shape; females are virtually identical to daphnis (the only other group member with which it can be sympat- ric). DISTRIBUTION (Fig. 246; appendix). — South- western Mexico from northern Michoacan, south- ern Guanajuato, and Zacatecas westward through Jalisco and northward along the Pacific coast from Nayarit to central Sonora. (Selander and Vaurie [1962] identify the type locality as Villa Corona, 115 km WSW of the city of Durango, Durango, Mexico.) 0-1500 m. Coprophagous; common in pasturelands. Collection dates: June-December (most June-August). COMMENTS. This species is most closely re- lated to daphnis. The females of the two are vir- tually indistinguishable morphologically, and pos- itive identification requires a sample including associated large males. I have chosen to maintain furiosus as a distinct species because of a lack of well-developed males intermediate between it and daphnis and because it displays less intraspecific variation than does daphnis. The coloration of furiosus varies geographically. In Sonora, Sinaloa, Nayarit, and western Jalisco, individuals are green and often highly shining; dark blue individuals are rare. In northern Jalisco, dark blue, blue-green, and dark green individuals are common; bright green is unusual. In Michoacan, practically all specimens examined are either dark blue or blue-green. I conclude that the range of the species has most recently expanded northward and, during the expansion, green coloration has become virtually fixed. The sculpturing of the elytral inter- striae also varies geographically. Along the western coast of Mexico (Nayarit, Sinaloa, and Sonora), they are commonly uniformly microrugose ( x 30); to the southeast, they are commonly uniformly mi- crorugose ( x 30); to the southeast, they are usually smooth or roughened only along the striae, rarely completely roughened. Phanaeus ( Phanaeus ) daphnis Harold Figures 94, 95, 231, 232, 239, 242, 246 Phanaeus daphnis Harold, 1863:166 Type: Male lectotype (“Mexico”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:114). Phanaeus herbeus Bates, 1887:61, NEW SYN- ONYMY Type: Male holotype (“Mexico”), British Mu- seum (Natural History), London. Phanaeus coerulus Bates, 1887:61, NEW SYN- ONYMY Type: Male holotype (“Mexico”), British Mu- seum (Natural History), London. Phanaeus tricornis OlsoufiefF, 1924:105, NEW SYNONYMY Type: Male lectotype (no data), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:l 16). Phanaeus substriolatus Balthasar, 1939:245 (Ar- naud, 1982c:125) Type: Male holotype (“Mexiko”), Narodni Mu- zeum, Prague. NOMENCLATURAL NOTES. Bates described coerulus and herbeus as varieties of daphnis ; the former name is sometimes attributed to Harold (1863), who proposed no name for his variety of daphnis. Arnaud (1982a:116) synonymized herbeus (as “herbaeus”) and tricornis. OTHER REFERENCES: Gemminger and Har- old, 1869; Bates, 1889; Nevinson, 1892a; Villada, 1901; Gillet, 1911b; Hinton, 1935; Blackwelder, 1944; Islas, 1942; Halffter and Matthews, 1966 (as nimrod ); Barrera, 1969; Edmonds, 1972; Edmonds and Halffter, 1972, 1978; Kohlmann and Sanchez- Colon, 1984. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 94, 95). — Dorsum dark blue, blue- green, green, or yellow-green; rarely with coppery reflections on pronotum. Head. — Clypeal process usually rounded, sometimes triangular. Prono- tum.— Basal fossae present; base of disk only weak- ly impressed in female. Pronotal Sculpturing. — Male: In large individuals, sides granulate anteri- orly, granules becoming flattened posteriorly, re- placed by effaced granules and punctures beneath posterolateral angles of pronotum; disk coarsely rugose, usually more densely so posteriorly. In smaller individuals, sides evenly granulate to pos- terior margin; disk finely granulate to granulorugose Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 55 anteriorly, punctatorugose to punctate posteriorly. Female: Sides granulate; disk granulate to granu- lorugose anteriorly, granules becoming scale-like posteriorly until replaced by progressively more punctate area near posterior margin. Elytra: Striae fine, minutely punctate (x30). Interstriae flat to weakly convex, sometimes completely shagreened, dull, sometimes smooth, brightly shining. Second- ary Sexual Characters.- — Male: Shape of pronotum variable, as in Figures 231, 232, 239, 242. Female: As described for species group. Specimens Exam- ined.— 314 males, 239 females (length 10-17 mm; width 6-10 mm). DIAGNOSTIC REMARKS. The spine-like pro- jections from the sides of the pronotal disk distin- guish large males of daphnis from other species in the group. Females and small males, however, can be virtually impossible to separate morphologically from those of furiosus. DISTRIBUTION (Fig. 246; appendix). Balsas River valley of southern Mexico from Puebla to Michoacan, extending into transitional areas along southern limit of Transverse Volcanic Range. 1000- 1600 m. Coprophagous. Collection dates: June- October (most June-September). COMMENTS. This species is the most variable of the group and, as defined here, may include sev- eral yet cryptic taxa I have been unable to distin- guish. Characters usually fixed in other members of the group (elytral relief and sculpturing, shape of clypeal process, shape of posteromedian process of pronotum of large males) vary in daphnis. In- deed, the only consistent features of the species are ecogeographic distribution and the acute lateral processes of the male pronotum. This species, like its Balsas valley companion demon, ascends the southern slopes of the Transverse Volcanic Range. The posteromedian process of the pronotum of large males varies in shape and size. In typical daphnis (Figs. 231, 232, 239) it is acute and con- siderably shorter than the lateral processes of the disk. In other individuals, it is much longer and either acute or weakly bituberculate apically (Fig. 242). These variants were called “var. coerulus ” by Bates (1887) in reference to Harold (1863) and later accorded species status by Olsoufieff (1924) under the name tricornis. I consider the “tricornis” con- dition just one aspect of the intraspecific variation of daphnis. It is common in Michoacan popula- tions and appears also in populations in Guerrero. I have not observed the “tricornis” variant in spec- imens collected elsewhere although Olsoufieff cites Cholala, Puebla, as the source of one of his cotypes. Color, on the contrary, is less variable in daphnis than in all other members of the group except fu- riosus. Most (>95%) observed specimens are green or yellow-green. Rare individuals are dark blue (“var. herbeus ” of Bates, 1887); rarer still are yellow-green specimens with coppery highlights on the prono- tum. These rare color forms are most frequent in populations from Tuxpan, in eastern Michoacan. Field data from Cuernavaca, Morelos, suggest that daphnis, unlike most Phanaeus, prefers equine dung (J. Blackaller, pers. comm.). The Triangularis Group DIAGNOSIS. [1] Outer margin of head not notched laterally; [2] clypeal process a transverse, rounded ridge; [3] cephalic process of female a tri- tuberculate carina in front of eyes; [4] anterolateral portions of pronotum finely granulate, more dense- ly so in females, small males (Fig. 253); [5] basal pronotal fossae present, sometimes almost effaced; [6] disk of male pronotum (Figs. 247, 249, 251) flat, posterolateral angles not strongly rounded laterally; [7] pronotum of female (Figs. 250, 252) with nar- row, medially tuberculate anteromedian transverse ridge followed by weak concavity and feeble tu- mosity; [8] elytral interstriae (Figs. 254, 255) con- spicuously ( x 10) punctured, usually convex (flat in triangularis texensis)\ puncturing fine to coarse; [9] front tibiae tridentate (fourth [basal] tooth some- times very weakly indicated in unworn specimens); [10] basal segment of female middle tarsus not wid- ened apically; [11] longer mesotibial spur not dis- tinctly dilated subapically; [12] Mesa Central and Sierra Madre Oriental of Mexico and southeastern United States (Fig. 256). In 1972 I overlooked the close relationship be- tween triangularis and adonis, which are brought together in this species group. The sculpturing of the elytra of adonis and triangularis, sen. str., shape of the mesotibial spur and mesobasitarsus, second- ary sexual characters, and pronotal sculpturing are the salient group characters; they suggest a closer relationship to the hermes group than to the tridens group. The case of triangularis is interesting in that one subspecies, texensis, approaches the vindex group in the relief and sculpturing of the elytral interstriae; it is the only taxon outside the vindex group that has a regular interstrial pattern of ru- gosity, albeit very fine. KEY TO THE SPECIES AND SUBSPECIES OF THE TRIANGULARIS GROUP la. Dorsum uniformly dark blue, blue-green, or bright green (Figs. 63-65). Pronotum rather densely granulate anterolaterally. Mesa Central and Sierra Madre Oriental of Mexico (Fig. 256) Phanaeus (P.) adonis Harold b. Dorsum somber, black to weakly shining cop- pery red (Figs. 58-62); if weakly to strongly shining green, then elytral interstriae flat. Pro- notum usually sparsely and finely granulate, more so in male. United States 2 2a. Elytral interstriae shining, distinctly convex (rarely dull, almost flat), with distinct, simple puncturing (Fig. 255); first interstria not dis- tinctly shinier than others. Dorsum uniformly either black or muted coppery red (Figs. 58, 59). Southeastern United States from Kansas to Louisiana-Texas border eastward to Carolinas 56 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Figures 247-255. Phanaeus ( Phanaeus ) triangularis group (247, P. triangularis texensis, male; 248, same, female; 249, P. t. triangularis, male; 250, same, female; 251, P. adonis, male; 252, same, female; 253, P. triangularis texensis, anterolateral portion female pronotum; 254, same, dorsal view elytron; 255, P. t. triangularis, dorsal view elytron). 247 249 252 Phanaeus (P.) triangularis triangularis (Say) b. Elytral interstriae 2-8 dull, flat, evenly and densely covered by large, shallow punctures that coalesce to form reticulate pattern of ridg- ing ( x 20) (Fig. 254); first interstria convex, sim- ply punctate, distinctly shinier than other in- terstriae. Dorsum uniformly black to muted coppery red, rarely dull to brightly shining green (Figs. 60-62). Texas . .Phanaeus (P.) triangularis texensis, NEW SUBSPECIES Phanaeus ( Phanaeus ) adonis Harold Figures 63-65, 251, 252, 256 Phanaeus adonis Harold, 1863:169 Type: Male lectotype (“Mexico”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:114). OTHER REFERENCES. Harold, 1868a; Gem- miner and Harold, 1869; Bates, 1887; Nevinson, 1892a; Villada, 1901; Gillet, 1911b; Olsoufieff, 1924; Blackwelder, 1944; Halffter and Matthews, 1966; Barrera, 1969; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 63-65). — Dorsum uniformly dark blue, blue-green, dark green, or bright yellow-green. Pronotum of male completely colored; that of fe- male with black anteromedian carina often fol- lowed by small, transverse black spot. Pygidium colored as elytra. Venter dark with colored reflec- tions on legs, sterna. Pronotal Sculpturing. — Male: In larger individuals, sides densely, finely granulo- rugose, rugosities becoming flatter, larger toward sides of disk, grading into coarse puncturing mixed with flat rugosities posteriorly. Disk more coarsely, evenly granulorugose, rugosities not forming ap- preciable ridging; texture of disk sculpturing chang- ing (sometimes abruptly) posteriorly to dense, coarse granulorugosity followed by conspicuous smooth, Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 57 punctate area. Smaller individuals as above except that remnant of flattened portion of disk finely gran- ulorugose to almost granulate, posterior portion of disk punctatorugose medially, simply punctate lat- erally. Female: Sides as in male; rugosities becoming flat on disk, grading posteriorly first into asperate punctures followed by simple punctures near pos- terior margin. Elytra. — Striae simple to minutely punctate, shagreened (x30). Interstriae convex, shining, bearing sparse, simple puncturing ( x 10); sculpturing coarser, denser on interstriae 6-8. All interstriae finely roughened at least along striae, occasionally over entire surface. Secondary Sexual Characters. — Male (Fig. 251): Pronotal disk flat, strongly triangular. Female (Fig. 252): As described for species group. Specimens Examined. — 92 males, 76 females (length 12-17 mm; width 7-11 mm). DIAGNOSTIC REMARKS. The most obvious differences between adonis and triangularis are size, color, and distribution. DISTRIBUTION (Fig. 256; appendix). Mid- to high-elevation open forests of the Sierra Madre Ori- ental from Nuevo Leon south to Hidalgo, and the eastern Mesa Central, Mexico. 350-2100 m. Co- prophagous. Collection dates: June-September. COMMENTS. In 1972 I overlooked the close relationship between this species and triangularis (see “Comments” under that species). Phanaeus ( Phanaeus ) triangularis (Say) Figures 58-62, 247-250, 253-256 Copris triangularis Say, 1823:206 Type: Male neotype (“Arkansas, Little River County, 2mi S Foreman”), Museum of Com- parative Zoology (MCZ Neotype No. 32895), Harvard University, PRESENT" DESIGNA- TION. Phanaeus triangularis (Say), LeConte, 1847:85 Phanaeus torrens LeConte, 1847:85 (LeConte, 1854: 217) Type: Male lectotype (no data), Museum of Comparative Zoology (MCZ Type No. 3710), Harvard University, PRESENT DESIGNA- TION. Phanaeus niger Olsoufieff, 1924:95, NEW SYN- ONYMY Type: Male lectotype (“Mexico”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:116). 58 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Phanaeus goidanichi Balthasar, 1939:243 (Arnaud, 1982c:125) Type: Male holotype (“North America-PMexi- co”), Narodni Muzeum, Prague. Phanaeus triangularis triangularis (Say), NEW STATUS Phanaeus triangularis texensis, NEW SUBSPE- CIES Type: Male holotype (“Texas, Dallas Co., 2mi N. Cedar Hill”), Museum of Comparative Zo- ology (MCZ Type No. 33405), Harvard Uni- versity (314 paratypes deposited in all major collections cited in the acknowledgments; no allotype designated; label data appear in the appendix). NOMENCLATURAL REMARKS. There has been considerable confusion attending use of the names “triangularis,” “torrens,” and “niger.” Re- cent usage follows that established by Robinson (1948) based on Olsoufieff (1924). In my opinion, both misidentified “triangularis” in the sense of Say (1823) in applying the name only to certain Texas populations of this species. While Say’s type ma- terial is assumed to have been destroyed, his original description of triangularis is clear enough to ex- clude these Texas populations and to include tor- rens, niger, and goidanichi. LeConte (1854) himself synonymized torrens and triangularis. All Texas populations examined (except those from far east- ern portions of the state) comprise a distinct taxon for which, in the absence of any available name, I propose texensis (see “Comments,” below). OTHER REFERENCES. Lacordaire, 1856; LeConte, 1859, 1863; Gemmingerand Harold, 1869; Blanchard, 1885; Nevinson, 1892a; Henshaw, 1885; Gillet, 1911b; Leng, 1920; Brown, 1927; Lindquist, 1935; Cartwright, 1939 (as niger); Blackwelder and Blackwelder, 1948; Loding, 1945; Lengerken, 1954; Halffter and Matthews, 1966; Fincher et al., 1969; Barrera, 1969; Edmonds, 1972; Fincher, 1973a, 1981 (as torrens ). DESCRIPTIVE REMARKS. Color and Color Pattern.— (a) triangularis (Figs. 58, 59): Dorsum moderately shining black (“niger”) or moderately shining to dull coppery red (“torrens”); interme- diates almost black with weak coppery sheen visible especially on pronotum. Pygidium venter black, sometimes with weak reddish reflections, (b) tex- ensis (Figs. 60-62): Dorsum dull black or coppery red, occasionally weakly shining; rarely dull to moderately shining green with or without coppery red highlights on pronotum. Pygidium, venter black, sometimes with weak-colored reflections. Pronotal Sculpturing. — Male: In large individuals, sides fine- ly granulate, granules sparse to moderately dense and becoming scale-like, forming some ridging be- hind lateral fossae. Disk granulate, somewhat more coarsely so than sides, appearing finely roughened to unaided eye; granules replaced posteromedially by ill-defined area of coarse, rugose puncturing. Smaller individuals as above except posterior por- tion of disk rather densely punctate, each puncture bearing minute tubercle anteriorly, occasionally also (especially in texensis) minute, shining central spot. Female: As in smaller male except disk more dense- ly granulate, granules distinctly more flattened, scale- like. Elytra.— Striae fine, minutely punctate (x30). (a) triangularis (Fig. 255): Interstriae distinctly con- vex, evenly shining; rarely appearing both dull and flattened; bearing simple punctures (x20), some- times almost effaced; first interstria not usually shin- ier than other interstriae. (b) texensis (Fig. 254): Interstriae 2-8 dull, flat, densely covered by large, flat punctures which coalesce to form reticulate, often shining ridging (x 10); each puncture usually bearing minute, shining, central spot (x30). First interstria convex, rather densely punctate, almost always much shinier than other interstriae. Second- ary Sexual Characters. — Male: Pronotum as in Fig- ures 247, 249. Female: As described for species group. Specimens Examined. — triangularis: 163 males, 151 females; texensis: 171 males, 175 fe- males (length 12-21 mm; width 7-12 mm). DIAGNOSTIC REMARKS. This species differs from adonis most obviously by color and distri- bution; pronotal sculpturing will distinguish it from other United States species. Small individuals can resemble the dark phase of prasinus, from which they differ in elytral sculpturing. The new subspe- cies, texensis, differs from the nominate subspecies by elytral sculpturing. DISTRIBUTION (Fig. 256; appendix). trian- gularis— Forested and semiforested areas of south- eastern United States from eastern Kansas, Okla- homa, and Texas to near the Atlantic Coast (unknown from Florida). 0-1000 m. Copropha- gous. Collection dates: April-November (mostly June-October in northern part of range), texen- sis— Forested and semiforested areas of Texas ex- cept far western and Panhandle areas. Copropha- gous; preferring areas with rocky soils. Collection dates: March-October except July (apparently bi- voltine). COMMENTS. In 1972 I included triangularis in the “ vindex complex” along with the present members of the vindex group (q.v.). This species, along with adonis, forms a distinct group more closely related to the tridens and hermes groups than to the other United States species. Some recent workers (e.g., Fincher, 1973a, 1981) have considered torrens to be a distinct species with two subspecies, torrens and niger, and have used the name triangularis almost exclusively in refer- ence to the Texas form. The nomenclatural changes proposed above may cause some confusion since torrens LeConte becomes a subjective junior syn- onym of triangularis Say. The distinction between the two subspecies is clear if the elytral and pronotal sculpturing of typ- ical individuals are compared; the variation of ely- Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 59 tral relief and sculpturing is similar to that observed in igneus (q.v.). Intermediates between the two sub- species occur among populations in extreme eastern Texas (Sabine and Shelby Counties) and adjacent areas of Louisiana across the Sabine River, and in Arkansas (Carroll County); moreover, most popu- lation samples of triangularis, sen. str., include in- dividuals with distinctly roughened interstriae bear- ing obscure puncturing. All texensis specimens examined, however, have a reticulate pattern of fine interstrial ridges produced by large, shallow, con- fluent punctures on a flat surface. The color of triangularis, sen. str., varies geo- graphically. Specimens from the southern part of its range are shining black or nearly black with weak coppery highlights; specimens from north of about 33 degrees N latitude always have a distinct, dull coppery red cast. Observed populations of texensis usually include both black and dull coppery red individuals; I have seen green specimens from only three, widely separated counties (Dallas, Menard, and Harris). Regardless of color, triangularis, sen. str., is almost always moderately shining while tex- ensis is ordinarily dull. Information available to me suggests that tex- ensis most commonly occurs in, but is not restricted to, areas with rocky or sandy soils and patchy ground cover consisting of a mixture of clumped grasses and low trees such as mesquite and postoak. The type locality of this subspecies is an area of lime- stone outcroppings in southwest Dallas County, Texas, covered by a mixed oak-juniper woodland with some mesquite and patchy pasturelands. On the other hand, triangularis, sen. str., appears to be a resident of moister forests and semiforested areas. The type of triangularis, sen. str., is among those specimens of the Say collection presumed lost in the 1830s. The usual practice has been to select needed neotypes for Say species from the ]. L. LeConte or T. W. Harris Collection in the Museum of Comparative Zoology, Harvard University. These collections were presumably seen by Say, or com- pared to material in Say’s collection before it was destroyed. The LeConte Collection includes 15 specimens labeled triangularis, of which 12 are tex- ensis. None of the remaining three specimens agrees fully with Say’s original description. I have chosen, therefore, a large male from modern material col- lected in Arkansas (Say’s type locality) that agrees fully with the original description for present des- ignation as neotype. The Mexicanus Group DIAGNOSIS. [1] Outer margin of head scarcely to strongly notched laterally (Figs. 286, 287); [2] clypeal process rounded to quadrate; [3] cephalic carina of female transverse trituberculate carina (Fig. 283) or narrow corniform process in front of eyes (Fig. 282); [4] anterolateral portions of pronotum densely granulorugose (Fig. 274); [5] basal pronotal fossae present or absent; [6] disk of male pronotum flat (Figs. 262, 263) or convex (Figs. 264, 265) (con- cave in demon, Fig. 257), posterolateral angles rounded, upturned (except demon ); [7] pronotum of female (Figs. 273, 274) with broad, usually salient anteromedian prominence usually carinate anteri- orly and flanked on each side by elongate depres- sion extending toward eye (depressions weak in pilatei) ; [8] front tibiae tridentate (weak fourth tooth in demon), wider in female than male; [9] basal segment of female middle tarsus broad apically (Figs. 288, 289); [10] longer mesotibial spur dilated sub- apically (Fig. 289) (except in flohri and scutifer, Fig. 288), dilation stronger in female; [11] color variable (Figs. 66-85); [12] larger size, length usually ex- ceeding 20 mm; [13] southeastern Arizona, Mexico (except northcentral and northeastern portions), Central America as far south as Costa Rica, and southern Ecuador (Fig. 290). The mexicanus group comprises seven species: flohri, mexicanus, amithaon, wagneri (with two subspecies), lunaris, scutifer, and demon. The most conspicuous feature of the group is the form of the female pronotum — an anteromedian prominence flanked by trough-like depressions that descend to- ward the eyes. For reasons discussed fully below, lunaris and flohri are taxonomic isolates in the group. The taxonomic and geographic picture of the mexicanus group is very similar to that of the tri- dens group and suggests a common history. In both groups, females and small males are similar and often indistinguishable morphologically; moreover, the distribution patterns are such that, except for lunaris (which occurs in Ecuador), each species is paired ecogeographically with a member of the oth- er group (although not on a strictly one-to-one, nor a completely sympatric, basis) as follows: • mexicanus with daphnis or nimrod • amithaon with furiosus or tridens tridens • scutifer with tridens tridens • demon with tridens pseudofurcosus, daphnis, or nimrod • wagneri wagneri with eximius The intragroup relationships here present an in- teresting picture. Three species, amithaon, mexi- canus, and wagneri, form a subgroup whose dis- tribution depicts linear differentiation (Fig. 290). The close morphological similarity of females and small males of these species suggests recent diver- gence. Of the remaining species, demon and scutifer form a species pair, while lunaris and flohri are taxonomic isolates. KEY TO THE SPECIES AND SUBSPECIES OF THE MEXICANUS GROUP la. Elytral interstriae (Fig. 272) smooth, shining medially, dulled laterally by shagreening that extends into striae; interstriae uniformly dull 60 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Figures 257-269. Phanaeus ( Phanaeus ) mexicanus group (257, P. demon, male; 258, same; 259, same {“excelsus”}; 260, same [“mirabilis”]; 261, same, female; 262, P. scutifer, male; 263, P. flohri, male; 264, P. amithaon, male; 265, P. mexicanus, male; 266, same, female; 267, P. lunaris, male; 268, same, female; 269, P. w. wagneri, male). in highly worn specimens). Sides of pronotum densely, coarsely punctatorugose around, be- hind lateral fossae. Pronotal disk of large males (Fig. 263) flat, posterolateral angles strongly rounded, entire disk (seen from above) almost heart-shaped. Longer mesotibial spur not di- lated subapically (as in Fig. 288). Clypeal pro- cess usually almost quadrate. Dark blue-black (Fig. 66); sometimes weakly shining green. Known from isolated localities in Veracruz, Guerrero, Morelos, Puebla Jalisco, Sonora (Fig. 290B) Phanaeus (P.) flohri Nevinson b. Elytral interstriae evenly shining (or dulled by severe wear); shagreening, if any, restricted to striae. Sides of pronotum granulorugose around and at least some distance behind lateral fossae. Male pronotum variable. Longer metatibial spur dilated subapically (except scutifer ), especially in female. Other characters variable 2 2a. Outer margin of head deeply notched between clypeus, paraocular areas (Fig. 286, arrow), notch usually more-or-less right-angled such that paraocular area appears to project laterally beyond margin of clypeus. Raised outer margin Contributions in Science, Number 443 Edmonds: Revision of Phanaeus M 61 Figures 270-274. Phanaeus ( Phanaeus ) mexicanus group (270, P. demon, dorsal view elytron; 271, P. w. wagneri, same; 272, P. flohri, same; 273, P. mexicanus, female, frontal view; 274, same, anterolateral view). of mesocoxal cavity always abruptly widened posteriorly resulting in posterior width about double anterior width (Fig. 285, arrow). Shape of male pronotum as in Figures 257-260. Ce- phalic carina of female raised medially, (seen from front) quadrate or weakly emarginate (Fig. 281). Anteromedian prominence of female pronotum lacking distinct transverse carina. Dorsal surface, especially pronotal disk and el- ytra, highly shining, appearing polished; color variable (Figs. 69-73). Southwestern Mexico, Balsas River valley, southern Oaxaca, Chiapas, Pacific coast of Central America to central Cos- ta Rica (Fig. 290B) . . . Phanaeus (P.) demon Laporte-Castelnau b. Outer margin of head at most only weakly notched (Fig. 287). Raised outer margin of me- socoxa not usually widened posteriorly (Fig. 284). Male pronotum not as above; disk flattened or convex medially. Female cephalic carina either trituberculate carina or narrow, bituberculate process (Fig. 282). Anteromedian pronotal prominence usually with distinct transverse ca- rina. Other characters variable 3 3a. Cephalic process of female narrow, almost conical bituberculate projection (Fig. 282). Longer of mesotibial spurs not usually distinct- ly dilated subapically. Pronotal disk of large males flattened dorsally (Fig. 262). Dorsum green to yellow-green, rarely coppery (Figs. 77, 78). Coastal lowlands of central Veracruz (Fig. 290B) Phanaeus (P.) scutifer Bates b. Cephalic process of female trituberculate ca- rina. Longer mesotibial spur distinctly dilated subapically, more strongly so in female. Prono- tal disk of large males convex medially, usually bearing central tubercles or ridges of some sort. Color and distribution variable 4 4a. Pronotum of large males (Fig. 267) with distinct posteromedian, bowed transverse carina pos- terior to which the pronotal surface is abruptly much smoother than anteriorly. Transverse an- teromedian ridge of female pronotum fol- lowed by fairly deep, transverse concavity. Deep shining green. Western and southern Ecuador (Fig. 290A) Phanaeus (P.) lunaris Taschenberg b. Male pronotal disk lacking any well-defined smooth basal area set off by transverse carina (posterior area can, however, be smoother than rest of disk). Pronotum of female at most only weakly concave behind anteromedian carina. Color variable. Central America and Mexi- co 5 (Note: The following taxa can be difficult or im- possible to separate if reliable locality data are not available. Females and smaller males are indistin- guishable morphologically.) 5a. Basal pronotal fossae distinct in female and usually also in male; occasionally small, punc- tiform, rarely lacking in male. Disk of male pronotum usually flat. Southern Mexico to Costa Rica (Fig. 290A) Phanaeus (P.) wagneri Harold 6 b. Basal pronotal fossae usually absent, repre- sented at most by small, punctiform pits. Disk of male pronotum usually either weakly con- 62 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Figures 275-289. Phanaeus ( Phanaeus ) mexicanus group (275-279, P. demon, variation in shape of posterior portion of pronotum [lateral views] among well-developed males; 280, P. fiohri, lateral view head and pronotum of Sonoran male; 281, P. demon, frontal view cephalic process; 282, P. scutifer, same; 283, P. mexicanus, same; 284, P. mexicanus, mesocoxa [crosshatchedj and edge of coxal depression; 285, P. demon, same [arrow indicates widening of raised margin]; 286, dorsal view head, P. demon; 287, same, P. mexicanus; 288, tip of female mesotibia and basitarsus, P. fiohri; 289, same, P. mexicanus). vex or concave. Southern Mexico west to Pa- cific coast, north to Arizona 7 6a. Pronotum with conspicuous basal fossae sep- arated by distance usually no more than three times diameter of single fossa. Elongate an- terolateral concavities of female pronotum weak. Pronotum of large males with transverse, almost ridge-like tumosity between center, posterior margin of disk. Dark green-blue, blue- green, or green (Figs. 75, 76). Yucatan Penin- sula south to northern Chiapas and Guatemala (Fig. 290A) Phanaeus (P.) wagneri pilatei Harold b. Pronotum with small basal fossae separated by distance greater than three times diameter of single fossa; sometimes lacking in male. An- terolateral concavities of female pronotum conspicuous. Pronotal disk of large males lack- ing transverse tumosity, often with distinct smooth area adjacent to middle of posterior margin. Usually coppery red (Fig. 74); occa- sionally green, yellow-green, or dark blue. Guatemala to Costa Rica (Fig. 290A) .... Phanaeus (P.) wagneri wagneri Harold 7a. Pronotal disk of large males (Fig. 265) convex, bearing central triad of conspicuous tubercles, posterior two of which may be fused as single ridge. Balsas River valley, central Oaxaca, Fos Tuxtlas region of Veracruz Phanaeus (P.) mexicanus Harold b. Pronotal disk of large males (Fig. 264) slightly concave, lacking central tubercles. Central Mesa of Mexico from Tlaxcala to Jalisco, northward along Pacific coast to southern Sonora, interior of Sonora north into southern Arizona and New Mexico Phanaeus (P.) amithaon Harold Phanaeus ( Phanaeus ) fiohri Nevinson Figures 66, 263, 272, 280, 290B Phanaeus fiohri Nevinson, 1892b:33 Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 63 Figure 290. Distribution of the Phanaeus ( Phanaeus ) mexicanus group (letters following species name indicates map). Type: Male lectotype (“Jalapa”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:l 16). OTHER REFERENCES. Nevinson, 1892a; Gil- let, 1911b; Olsoufieff, 1924; Rlackwelder, 1944; Ed- monds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Fig. 66). — Dorsum dark, weakly shining blue, often almost black, occasionally weakly shin- ing green. Coloration on pronotum restricted large- ly to anterolateral areas, to disk of male, to mid- longitudinal depression of disk of female. Elytra with weak bluish cast. Head. — Clypeal process al- most quadrate. Outer margin not notched. Pronotal Sculpturing.— -Male: In large individuals, sides gran- ulorugose anteriorly, punctatorugose around, be- hind lateral fossae; sculpturing grading posteriorly 64 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus to simple puncturing beneath posterolateral prom- inences of disk. Disk rather finely, sparsely granu- lorugose, becoming punctate on raised, smooth, posterior transverse area extending between pos- terolateral prominences. Smaller individuals with posterior two-thirds of pronotum rather coarsely, densely punctate to punctatorugose; granulorugose areas restricted to anterolateral portions of pro- notum, small, round, concave remnant of disk near anterior margin. Female: As in smaller males except puncturing on posterior portion of disk weaker, usually sparser, sometimes becoming almost effaced near posterior margin. Pronotum. — Basal fossae present in both sexes, although sometimes only weakly indicated. Mesotibiae. — Longer spur not di- lated subapically (Fig. 288). Elytra (Fig. 272). — Stri- ae simple, impressed. Interstriae subconvex, smooth, brightly shining medially, dulled laterally by sha- greening such that each interstria appears broadly polished midlongitudinally (interstriae uniformly dull in highly worn specimens). Secondary Sexual Characters. — Male: Pronotum as in Figure 263; disk of large individuals flat, seen from above almost heart-shaped; raised posterior area impressed mid- longitudinally. Female: Cephalic carina tritubercu- late, in front of eyes; pronotal prominence carinate anteriorly in large individuals. Specimens Exam- ined.— 17 males, 10 females (length 16-20 mm; width 9-12 mm). DIAGNOSTIC REMARKS. This species differs from other members of the group by the following combination of characters: microsculpture of ely- tral interstriae, shape of longer mesotibial spur, shape of clypeal process, shape of large male pronotum, and sculpturing of sides of pronotum. DISTRIBUTION (Fig. 290B; appendix). Known only from isolated localities in the Mexican states of Guerrero, Mexico, Jalisco, Puebla, Sonora, and Veracruz. 600-1800 m. Habits unknown. Collec- tion dates: June-October. COMMENTS. Except for one series, I have seen only a few specimens of this interesting species col- lected from scattered localities. The series in ques- tion numbers 16 individuals collected by Leonardo Delgado-Castillo and Julian Blackaller in the vicin- ity of Acahuizotla, Guerrero, 650-1400 m, in the Sierra Madre del Sur (see Delgado-Castillo, 1989). Most of the series was collected from traps baited with feces or carrion placed in protected areas cov- ered by very large rocks where exposed soil is re- stricted to cracks and crevices populated by scrubby vegetation— the kind of habitat attractive to bur- rowing animals. That this species has been collected almost exclusively under such unusual conditions may very well portend that flohri will be the first Phanaeus shown to be an inquiline in mammal nests. Four large, unusual males are known from the three Sonoran localities listed in the appendix. These specimens, collected and very kindly brought to my attention by Scott McCleve, are tentatively assigned to flohri pending discovery of associated females. They differ from other male flohri I have seen in two striking respects: the cephalic horn is strongly curved and almost hook-shaped (Fig. 280), and the pronotum (seen laterally, Fig. 280) is distinctly shal- lower and (seen from above) wider anteriorly than posteriorly. Otherwise, these specimens display the salient features of this species given above. This population could later prove to be a distinct taxon. The general facies of this species is reminiscent of the hermes group, particularly the black phase of prasinus (“lugens”). It occupies an isolated tax- onomic position within the mexicanus group (see “Diagnostic Remarks”). The rather extensive punc- tation of the sides of the pronotum behind the lateral fossae is observable elsewhere in the sub- genus only in the hermes group, with which flohri also shares narrow mesotibial spurs. While in many respects flohri can be regarded as an annectant be- tween the hermes and mexicanus groups, the shape of the female pronotum clearly places it in the latter. Phanaeus ( Phanaeus ) demon Laporte-Castelnau Figures 69-73, 257-261, 270, 275-279, 281, 285, 286, 290B Phanaeus demon Laporte-Castelnau, 1840:81 Type: Male neotype (“Mexico: Michoacan, La Huacana”), Castelnau Collection, University of Victoria, Melbourne, PRESENT DESIG- NATION. Phanaeus pegasus Sturm, 1843:331 (Harold, 1863: 169, as “damon”) Type: Unknown to me. Phanaeus obliquans Bates, 1889:389 (new name for mirabilis Bates, 1887:65, junior primary homonym of P. mirabilis Harold 1877:98), NEW SYNONYMY Type: Male lectotype (“Mazatlan, Mexico, Salle, B.C.A. p.65 sp.31, sp. figured”), British Mu- seum (Natural History), London, PRESENT DESIGNATION. Phanaeus excelsus Bates, 1889:389, NEW SYN- ONYMY Type: Male lectotype (“Chontales, Nicaragua, Janson, B.C.A. p.389 sp.32, sp. figured”), Brit- ish Museum (Natural History), London, PRESENT DESIGNATION. NOMENCLATURAL REMARKS. The name demon often appears incorrectly as “damon” in collections and in the literature. At the time Bates described mirabilis, the name was preoccupied by Harold’s species. Bates corrected his error by re- naming his species obliquans. Harold’s taxon was Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 65 moved to Taurocopris (now Diabroctis ) by Olsou- fieff (1924), where it remains. Because Bates’ mi- rabilis is a primary junior homonym and perma- nently unavailable, the valid name of his species is obliquans. OTHER REFERENCES. Imhoff, 1856 (as pega- sus ); Lacordaire, 1856; Nevinson, 1892a; Villada, 1901; Gillet, 1911b; Olsoufieff, 1924; Paulian, 1935; Blackwelder, 1944; Islas, 1942; Halffter and Mat- thews, 1966; Barrera, 1969; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 69-73). — Dorsum variable, but al- ways highly shining except in highly worn speci- mens. Color varies geographically as follows: Mex- ico from Isthmus of Tehuantepec westward, usually clear, brilliant green, often with yellow reflections, to golden green except for anterior portion of clyp- eus, tips of male pronotal prominences, areas of variable size on female pronotal disk which are black. Populations in southern Puebla (occasional specimens from other localities) often include in- dividuals of brilliant copper-red color with green reflections. From Chiapas eastward into Central America, color varies from dark green through sil- ver-green, yellow-green to silver-blue, or dark vi- olet-blue except for those black areas noted above. In all populations, disk of female pronotum may be nearly covered by irregularly shaped black area, or marked by anteromedian black spot, two smaller posterolateral ones. Pygidium, venter shining, with areas on sterna, femora colored like dorsum. Head. — Outer margin, at junction of clypeus, para- ocular areas (Fig. 286, arrow), with distinct notch resulting from lateral expansion of paraocular ar- eas, lateral margin of which can be almost straight; notch larger, more distinct in larger individuals, especially males. Underside of posterior portion of paraocular area of males with deep, semicircular concavity (more prominent in larger specimens). Prothorax. — Underside of anterior angles with dis- tinct, round concavity (more prominent in females), beneath which rounded tooth extends beneath paraocular area of head. Pronotum lacking basal fossae. Sides of pronotum (seen from above) round- ed in females, small males, distinctly bowed in large males between prominent anterolateral, lateral an- gles. Pronotal Sculpturing. — Larger males: densely granulorugose laterally; disk with large transverse rugosities except posteromedially and on pronotal horns where smooth. Smaller males, females: Densely granulorugose except posteriorly where smooth to weakly punctate; rugosities finer, denser anterolaterally. Front Tibiae.— Quadridentate in both sexes; basal tooth small, often worn away. Meta- sternum.— Raised outer margin of mesocoxal de- pression (Fig. 285, arrow) almost always more-or- less abruptly widened posteriorly such that posterior width almost double anterior width. Mesotarsus. — Basal segment in female broadly triangular, width at apex about equal length. Elytra (Fig. 270). — Striae fine, sometimes minutely punctured (xlO); striae 5-7 fainter than 1-4; striae 2-5 impressed basally. Interstriae 5-6 distinctly flatter than 2-4. Elytral suture sometimes bearing exposed row of erect, black setae than can be brush-like. Secondary Sex- ual Characters. — Male: In larger specimens, pro- notum with posterolateral angles varying in shape (as seen laterally) from recumbent conical processes (“obliquans,” Figs. 260, 279) to erect, massive quad- rate processes (“demon”/“excelsus,” Figs. 257-259). Female: Cephalic carina (Fig. 281) isolated, raised, seen from front more-or-less quadrate. Antero- median prominence of pronotum lacking transverse carina bordering anterior margin. Specimens Ex- amined.— 737 males, 562 females (length 10-21 mm; width 7-14 mm). DIAGNOSTIC REMARKS. The shape of the head margin, brilliant coloration, and secondary sexual features make this is a distinctive species not easily confused with other members of the group or genus. DISTRIBUTION (Fig. 290B; appendix). Pacific coastal areas of Mexico and Central America from southern Sinaloa to northern Costa Rica. Copro- phagous. Semidesert scrub and open deciduous for- ests south of major mountain systems. 0-1900 m. Collection dates: April-November (most June-Au- gust). COMMENTS. Phanaeus demon is a highly vari- able species which, as here defined, includes two variants heretofore regarded as separate species: ob- liquans and excelsus. As discussed below, these forms are best regarded as extremes in the variation of a single, widely distributed polymorphic species. The most striking aspects of the variation of de- mon are in color, fringing of the elytral suture, and shape of the pronotum of large males. Of these, only the latter varies in a way suggestive of sub- speciation. Through most of the range of the spe- cies, the projections from the pronotum of large males are massive, vertical processes that widen apically (Figs. 257, 258, 275). In southwestern Mex- ico populations (Colima, Jalisco, Nayarit, and Sina- loa), these projections are attenuated anteriorly to various degrees and appear in profile as narrow, posteriorly inclined processes (Figs. 276-279). In Central America, the projections are narrower, more erect, and more parallel-sided (Fig. 259); their shape, along with a pair of long, acute denticles near the anterior pronotal margin lacking in Mexican pop- ulations, give the pronotum a markedly different shape. The pronotal shape of females and small males is essentially the same throughout the range. The most frequent coloration of demon is bril- liant green usually combined with yellow reflections (Figs. 69, 70, 72). Two color variants are notewor- thy. Populations in southern Puebla show high fre- quencies of coppery green and coppery red indi- viduals (Fig. 71); reddish individuals are rare in populations from Morelos and Guerrero. In Cen- tral America, populations often include silver-blue (Fig. 73) or dark blue specimens, although light silver-green individuals are not uncommon. The “greens” of populations from Chiapas and Central 66 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus America tend to be less brilliant and darker than those from the remainder of Mexico. Most populations of this species lack a conspic- uous fringe of setae projecting from the elytral su- ture. A weak fringe is common in populations from El Salvador and Nicaragua. Populations from Mi- choacan and southwestern Mexico (state), how- ever, have a prominent, brush-like fringe. Other than the shape of the pronotum of large males, I have discovered no consistent characters which would support formal recognition of sub- species of demon. The distribution of the species appears to be continuous from southwestern Mex- ico to Costa Rica, but two intermediate areas need to be better sampled before ruling out distribu- tional gaps: the coasts of Oaxaca and southern Gua- temala. The occurrence of demon in Guanajuato, although not completely unreasonable, needs con- firmation. This is a common species apparently re- stricted to regions with hot, wet summers and warm, dry winters. The original type material of demon is unknown to me. There is no specimen assignable to demon in the Castelnau Collection at the Victoria Museum in Melbourne, nor is there any record to suggest that one existed. The neotype designated here agrees with Laporte-Castelnau’s original description in all respects except that the pronotum lacks any trace of pinkish or reddish reflections (“corselet ru- gueux”). Phanaeus ( Phanaeus ) scutifer Bates Figures 77, 78, 262, 282, 290B Phanaeus scutifer Bates, 1887:60 Type: Male lectotype (“Misantla, Mexico, Hoege, B.C.A. p.60 sp.14, sp. figured”), British Mu- seum (Natural History), London, PRESENT DESIGNATION. OTHER REFERENCES. Bates, 1889; Nevinson, 1892a; Gillet, 1911b; Olsoufieff, 1924; Blackwelder, 1944; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 77, 78). — Dorsum dark, highly shin- ing green, often with yellowish (rarely coppery) highlights on pronotum. Pygidium green to yellow- green. Venter with green reflections on sterna, legs. Head. — Outer margin with distinct notch at junc- tion of clypeus, paraocular area (as in Fig. 286). Pronotum. — Basal fossae absent or reduced to small punctures; present more often in females. Pronotal Sculpturing. — Male: In large individuals, disk coarsely, but not densely granulorugose except along posterior margin where smoother, rather finely punctate to punctatorugose; smoother basal area longest medially, strongly shortened laterally; sides densely granulorugose. Smaller individuals as above except smoother basal area less well demarcated, more clearly punctate, disk rugosities finer. Fe- males: Sides densely granulorugose, disk coarsely granulorugose to punctatorugose except posteri- orly where punctate. Mesotarsus. — Basal segment widened in female. Metatibia. — Longer spur not dilated subapically. Secondary Sexual Charac- ters.— Male (Fig. 262): Pronotal disk flat, with small acute tubercle near middle of anterior margin; pos- terolateral angles of disk rounded laterally, up- turned in larger individuals, reduced to rounded ridges near middle of disk in smaller individuals. Females: Cephalic process an apically bituberculate, conical process (Fig. 282) set forward of imaginary line connecting eyes; anteromedian process of pro- notum distinctly carinate anteriorly. Specimens Ex- amined.— 45 males, 38 females (length 15-20 mm; width 9-13 mm). DIAGNOSTIC REMARKS. The conical ce- phalic process is unique to females of the mexi- canus group. Large males differ from those of other species in the group by having a flat pronotal disk with a distinct, raised basal area along the posterior margin. DISTRIBUTION (Fig. 290B; appendix). Trop- ical lowlands of central Veracruz, Mexico. Copro- phagous. 0-1000 m. Collection dates: June-July. COMMENTS. I erroneously considered this spe- cies to be a close relative of the amethystinus group in 1972 because of the similarity in structure of the pronotum of large males (flattened with a smooth, raised basal area). The female, unknown to me then, as well as distinct similarities to demon (character- istic luster, notching of head margin) place scutifer in the mexicanus group. 1 have seen two long series (20+ specimens) of scutifer, one from Conejos (between the cities of Jalapa and Veracruz), the other from Palma Sola (on the coast north of Cardel), in addition to in- dividuals from scattered localities in central Vera- cruz. The Palma Sola population is unusual in that, despite several years of intensive collection in the area, no large, well-developed males have been found. Phanaeus ( Phanaeus ) lunaris Taschenberg Figures 67, 68, 267, 268, 290A Phanaeus lunaris Taschenberg, 1870:183 Type: Unknown to me. Phanaeus charon Harold, 1880a:151 (Gillet, 1911a: 319) Type: Unknown to me. OTHER REFERENCES. Nevinson, 1892a; Ohaus, 1909; Gillet, 1911b; Olsoufieff, 1924; Black- welder, 1944; Lengerken, 1954; Vulcano and Pe- reira, 1967; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 67, 68). — Dorsum dark, usually mod- erately shining green; occasionally with yellowish reflections that result in a dark bronze sheen. Py- gidium green. Venter black with green reflections Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 67 on legs, sterna. Head. — Outer margin with shallow notches separating clypeus from paraocular areas. Pronotal Sculpturing. — Male: In large individuals, sides densely granulorugose except adjacent to eyes where almost smooth, punctate beneath postero- lateral angles of disk; disk coarsely rugose, rugos- ities forming strong transverse ridges; posterome- dially, disk abruptly smoother, with asperate punctures that can coalesce as fine ridges; smoother posterior portion separated from remainder of disk by anteriorly bowed, transverse carina. Smaller in- dividuals as above except rugose area of disk re- duced, not separated from smoother posterior area by transverse carina. Female: Disk, behind antero- median ridge, with flattened scale-like granules that become progressively weaker posteriorly until re- placed by asperate punctures in concave postero- median area; sides as in male. Pronotum:- — Basal fossae punctiform. Mesotibia. — Longer spur dilat- ed subapically, more weakly so in male. Elytra.— Striae minutely punctate ( x 30), distinctly duller than interstriae. Interstriae convex, minutely, sparsely punctured (x30), appearing lacquered, distinctly shinier than striae. Secondary Sexual Charac- ters.— Male: Pronotum as in Figure 267; disk with smooth posteromedian area separated from re- mainder of disk by transverse ridge. Female (Fig. 268): Cephalic carina strongly tridentate, set in front of eyes. Anteromedian ridge of pronotum followed by tranverse concavity. Specimens Examined. — 64 males, 49 females (length 16-25 mm; width 10-15 mm). DIAGNOSTIC REMARKS.— This species dif- fers from other members of the group by distri- bution and form of the pronotum. DISTRIBUTION (Fig. 290A; appendix). West- ern and southern Ecuador (probably also extreme northwestern Peru). 0-2300 m. Coprophagous. Collection dates: probably all year (no records from April, June, and August). COMMENTS. The coloration of lunaris is con- siderably less variable than that of other members of the group. It occurs in arid and semiarid habitats, as does achilles. Although I have seen no specimens from there, I assume its range extends into the arid region of extreme northwestern Peru. The form of the pronotum of both sexes is reminiscent of that of the beltianus group, to which I presently con- sider lunaris an annectant. Its placement in the mexicanus group is based primarily upon the fe- male pronotum, which possesses the lateral de- pressions characteristic of the group. I have not been able to locate the type specimens of either lunaris or charon. Phanaeus { Phanaeus ) wagneri Harold Figures 74-76, 269, 271, 290A Phanaeus wagneri Flarold, 1863:170 Type: Male lectotype (“Nicaragua”), Museum National d’Histoire Naturelle, Paris (Arnaud, 1982a:115). Phanaeus pilatei Harold, 1863:170 Type: Male lectotype (“Yucatan”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:115). Phanaeus wagneri wagneri Harold, NEW STA- TUS Phanaeus wagneri pilatei Harold, NEW COM- BINATION NOMENCLATURAL REMARKS. Both the names wagneri and pilatei were proposed simul- taneously by Harold on the same page. The choice of the former name as nominate subspecies is by precedence of position. OTHER REFERENCES (to both subspecies un- less indicated otherwise). Bates, 1887, 1889 ( pila- tei); Nevinson, 1892a; Pittier and Biolley, 1895 (; wagneri ); Villada, 1901 ( wagneri ); Gillet, 1911b; Olsoufieff, 1924; Blackwelder, 1944; Barrera, 1969; Edmonds, 1972; Moron et al., 1986 {pilatei); Pa - lacios-Rios et al., 1990 {pilatei). DESCRIPTIVE REMARKS. Color and Color Pattern. — (a) wagneri (Fig. 74): Dorsal coloration highly variable; dark blue, blue-green, green, yel- low-green, or coppery red. Pronotum of male com- pletely colored except for black tips, sides of pos- terolateral angles; that of female with black anteromedian spot. Pygidium colored as elytra; ven- ter with colored reflections on legs, sterna, (b) pi- latei (Figs. 75, 76): Dorsum deep blue-green or dark green; otherwise as above. Pronotum. — Basal fos- sae almost always present; in wagneri, small, punc- tiform, sometimes absent in male; always present, usually conspicuous in female; in pilatei, conspic- uous in both sexes. Pronotal Sculpturing. — Male: In large individuals, sides densely granulorugose, disk coarsely rugose except posteromedially where usually distinctly smoother, punctate to punctato- rugose (in pilatei, disk raised transversely as ridge- like tumosity in front of smooth posteromedian area). Small individuals as above except disk less coarsely rugose, rugosities becoming scale-like me- dially, progressively weaker posteriorly. Female: Sides densely granulorugose, rugosities becoming scale-like on disk, progressively weaker posteriorly where replaced by asperate punctures. Elytra. — Striae minutely punctate (x40). Interstriae evenly convex, smooth to minutely roughened, punctate ( x 30). Secondary Sexual Characters .- — Male: Pro- notum as in Figure 269; Female: Cephalic carina trituberculate. Convexities flanking anteromedian pronotal prominence weaker, less shining in pilatei. Specimens Examined. — wagneri: 180 males, 146 females; pilatei: 88 males, 85 females (length 14- 20 mm; width 8-13 mm). DIAGNOSTIC REMARKS. This species most closely resembles mexicanus, from which it differs 68 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus by having basal pronotal fossae, in shape of the male pronotum, finer, more asperate puncturing of the female pronotum, and distribution. Small males also closely resemble sallei, which possesses several unique species group characters (q.v.) that distin- guish it from all members of the mexicanus group. DISTRIBUTION (Fig. 290A; appendix), wag- neri — -Transitional habitats from Chiapas to Costa Rica (500-2200 m). Coprophagous. Collection dates: May-December. pilatei—- Drier deciduous and ev- ergreen forests of the Yucatan Peninsula (0-1000 m). Coprophagous. Collection dates: May-Decem- ber. COMMENTS. It could be argued that the two subspecies recognized here should continue to be regarded as distinct species. I have elected to make them conspecific in recognition of their close sys- tematic relationship. The chief characters separat- ing them (basal pronotal fossae and shape of the male pronotum) intergrade especially among spec- imens from Chiapas, where the two subspecies are sympatric. Most of the specimens of wagneri that I have seen were collected long ago and bear imprecise locality data. Thus, the distribution dipicted in Fig- ure 290A is, at best, a rough approximation. Phanaeus ( Phanaeus ) mexicanus Harold Figures 79-82, 265, 266, 273, 274, 290A Phanaeus mexicanus Harold, 1863:171 Type: Male lectotype (“Mexico, Veracruz”), Museum National d’Histoire Naturelle, Paris (Arnaud, 1982a:115). Phanaeus divisus Harold, 1863:171 (Bates, 1887: 65) Type: Male lectotype (“Mexico”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:l 14). Phanaeus scintillans Bates, 1887:64, NEW SYN- ONYMY Type: Male lectotype (“Mexico, Oaxaca, Salle, B.C.A. p.64 sp.27, sp. figured”), British Mu- seum (Natural History), London, PRESENT DESIGNATION. NOMENCLATURAL REMARKS. The name mexicanus takes precedence over divisus by po- sition on page. OTHER REFERENCES. Blanchard, 1885; Nev- inson, 1892a; Villada, 1901; Gillet, 1911b; Leng, 1920; Olsoufieff, 1924; Blackwelder, 1944; Black- welder and Blackwelder, 1948; Robinson, 1948; Halffter and Matthews, 1966; Edmonds, 1972; Ed- monds and Halffter, 1972, 1978; Halffter and Ed- monds, 1982; Moron, 1984. DECRIPTIVE REMARKS. Color and Color Pattern (Figs. 79-82). — Dorsal coloration highly variable — dark blue, green, yellow-green, coppery red; most often yellow-green or coppery red with green highlights; dark blue phase rare. Pronotal disk completely colored except for following black ar- eas: posterolateral angles of male; posterolateral angles, irregular anteromedian area of female. Py- gidium colored as elytra. Venter dark with colored highlights on legs, sterna. Pronotum. — Basal fossae usually absent; rarely indicated in female by small punctiform pits. Pronotal Sculpturing. — Male: In large individuals, sides densely, finely granuloru- gose; disk rugose, rugosities forming transverse ridging medially; posteromedially sculpturing varies from coarsely punctatorugose to smooth with scat- tered punctures; posterolateral angles of disk punc- tate behind. In smallest individuals, granulorugosity extends to middle of disk, grades into dense, coarse puncturing near posterior margin. Female: Granu- lorugose except posteromedially where disk smooth, sparsely punctate to punctatorugose. Elytra. — Stri- ae simple to minutely punctate (x40), impressed. Interstriae evenly convex, moderately shining, mi- nutely punctate ( x 40), appearing smooth to unaid- ed eye. Interstriae 6-8 (rarely 1-8) often unevenly roughened by minute puncturing ( x 20). Secondary Sexual Characters.— Male: Pronotum of large specimens as in Figure 265; disk bearing three acute tubercles posterior two of which may fuse as trans- verse ridge. Female (Figs. 273, 274): Cephalic carina trituberculate. Transverse carina bordering antero- median prominence of pronotum slightly bowed anteriorly, weakly toothed medially; occasionally sinuous, distinctly, but not strongly tuberculate me- dially. Specimens Examined. — 179 males, 133 fe- males (length 14-23 mm; width 9-15 mm). DIAGNOSTIC REMARKS. As mentioned above, only large, well-developed males of this spe- cies are distinguishable from amithaon on mor- phological grounds. Reliable locality information is necessary to distinguish females and small males of the two species. DISTRIBUTION (Fig. 290A; appendix). Appar- ently disjunct as follows: Balsas River valley, central Oaxaca, escarpment of Mesa Central in east-central Veracruz, and the Los Tuxtlas region of Veracruz. Coprophagous. 0-1500 m. Collection dates: May- October (May-January in Los Tuxtlas). COMMENTS. The apparently disjunct distri- bution of mexicanus may be an artifact of col- lecting, at least in all cases except Los Tuxtlas. This species appears to be allopatric with its closest rel- ative, amithaon, although I have seen some inter- mediate specimens that suggest the two hybridize in limited areas of eastern Michoacan (e.g., Tuxpan; see also “Comments” under amithaon). Phanaeus ( Phanaeus ) amithaon Harold Figures 83-85, 264, 290A Phanaeus amithaon Harold, 1875b:88 Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 69 Figures 291-298. Phanaeus ( Phanaeus ) beltianus group (291, P. beltianus, male; 292, P. sallei, male; 293, P. howdeni, male; 294, P. beltianus, female; 295, same, dorsal view elytron; 296, P. sallei, same; 297, P. sallei, posterior view front femur and tibia [upper arrow indicates field of large punctures; lower arrow indicates expanded margin]; 298, P. quadridens, same, cf. 297). Type: Male lectotype (“Mexico, Guanajuato”), Museum National d’Histoire Naturelle, Paris (Arnaud, 1982a:114). NOMENCLATURAL REMARKS. For unstated reasons, Bates (1887) emended Harold’s name to amythaon, and his spelling has received wide sub- sequent use. The emendation, however, was un- justified. Nevinson (1892a) synonymized scintillans Bates and amithaon; Bates’ taxon is a synonym of mexicanus (q.v.). OTHER REFERENCES. Blanchard, 1885 (as mexicanus ); Villada, 1901; Gillet, 1911b; Leng, 1920; Olsoufieff, 1924; Islas, 1942; Blackwelderand Blackwelder, 1948; Robinson, 1948; Halffter and Matthews, 1966; Barrera, 1969; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 83-85). — Dorsal coloration highly variable— green, yellow-green (often with coppery reflections), copper-red, deep red; moderately shin- ing to dull. Pronotum of male completely colored except for black posterolateral angles; that of fe- male with irregular black anteromedian spot, elon- gate black areas posterolaterally. Pygidium colored as elytra. Venter dark with colored reflections on legs, sterna. Pronotum. — Basal fossae absent ex- cept in females from Pacific coast. Pronotal Sculp- turing.— Male: In large individuals, sides finely, densely granulorugose; disk coarsely rugose, ru- gosities often forming transverse ridging, except for somewhat smoother punctatorugose area poster- omedially; smoother posteromedian area of disk 70 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus sometimes bounded anteriorly by incomplete, bowed ridge; posterolateral angles of disk punctate behind. Smaller specimens as above except sculp- turing finer, denser. Female: Sides densely granu- lorugose, sculpturing becoming coarser on disk; disk punctate to punctatorugose posteromedially where somewhat smoother than remainder of disk. Ely- tra.-—Striae fine, simple to minutely punctate (x40). Interstriae evenly convex, appearing smooth to unaided eye; interstriae 6-8 (rarely 2-8) often roughened by minute punctures (x40). Secondary Sexual Characters. — Male: Pronotum as in Figure 264. Female: Cephalic carina tridentate. Pronotum as in Figures 273, 274; transverse anteromedian ca- rina straight or sinuate medially. Specimens Ex- amined.— 627 males, 492 females (length 13-25 mm; width 9-16 mm). DIAGNOSTIC REMARKS. As mentioned in the key to species, amithaon is similar to mexicanus (see “Comments,” below). DISTRIBUTION (Fig. 290A; appendix). Mesa Central of Mexico westward to Colima and Jalisco, northward along the Pacific coast into southeastern Arizona. 0-1800 m. Coprophagous, common in pastures. Collection dates: June- December (most June-August). COMMENTS. This is a widely distributed spe- cies replaced to the south and east by mexicanus, to which it is closely related. Without reliable geo- graphic data, females and small males of the two are inseparable. There is little question that the two species are vicariants. As mentioned in the introduction to the mexi- canus group, amithaon is often collected along with furiosus. Both species occur in large numbers in cow dung. I have been able to discover no dif- ference between Arizona populations and those in Mexico. I assume that its spread north is recent and associated with the introduction of cattle into the southwestern United States. Most of the variation in coloration of amithaon is observable only in the central portions of Mex- ico. Coastal populations in Nayarit and Sinaloa are of a fairly uniform green, to which are added cop- pery highlights in populations in Sonora and Ari- zona. I have seen specimens labeled Torreon, Monclo- va, and Saltillo (all in Coahuila). These localities need confirmation. The Beltianus Group DIAGNOSIS. [1] Outer margin of head at most only scarcely notched; [2] clypeal process rounded; [3] cephalic carina of female trituberculate, set in front of eyes; [4] anterolateral portions of prono- tum densely granulorugose; [5] basal pronotal fos- sae present or almost effaced; [6] disk of male pro- notum (Figs. 291-293) flat, posterolateral angles rounded, upturned; base with smooth area sepa- rated from rest of disk by transverse carina or tu- bercle; each anterolateral angle of disk bearing con- Figures 299, 300. Phanaeus ( Phanaeus ) beltianus group (299, P. beltianus, anterolateral margin of pronotum; 300, P. sallei, same). ical tubercle; [7] pronotum of female (Fig. 294) with narrow, medially tuberculate carina followed by oval concavity posterior margin of which inter- rupted by rounded gibbosity; [8] front tibiae tri- dentate; [9] midventral carina of front femora an- gulate near base; base with field of large, coarse, setose punctures (Fig. 297); [10] basal segment of middle tarsus of female broad apically; [11] longer mesotibial spur dilated apically, more strongly so in female; [12] evergreen forests from central Ve- racruz to eastern Panama (Fig. 301). This group comprises three closely related Mid- dle America species, all of which inhabit moist trop- ical evergreen forests: beltianus Bates, sallei Har- old, and howdeni Arnaud. The principal group characters are the form of the male and female pronota and the unique shape and puncturing of the front femora. The pronota, especially that of the male, resemble that of lunaris, which I regard as an annectant between the beltianus and mexi- canus groups. The distributions of these three species form a chain extending most of the length of Middle America. The northern and widest range is that of sallei, a species replaced by beltianus in southern Nicaragua and Costa Rica; howdeni is known only from Panama. The distributions of all three species, but particularly beltianus, are not well known. Un- corroborated data suggest that sallei reaches Pan- ama and, therefore, may be sympatric with both other species. KEY TO THE SPECIES OF THE BELTIANUS GROUP la. Elytral interstriae shining medially, dull adja- cent striae such that interstriae appear polished midlongitudinally (Fig. 295). Sculpturing of sides of pronotum extending to anterolateral mar- gins, where considerably weaker but nonethe- less visible ( x 30). Basal pronotal fossae prom- inent in female; very small, but usually distinct in male. Pronotum of female not distinctly im- pressed midlongitudinally. Lateral margin of pronotum (seen from above) usually distinctly curved inward between anterolateral and lat- eral angles (Fig. 299). Lowland forests of Nic- aragua and Costa Rica (Fig. 301) Phanaeus (P.) beltianus Bates b. Elytral interstriae evenly shining or dull, not distinctly shinier medially. Sculpturing of sides Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 71 of pronotum effaced adjacent to anterior mar- gin. Basai pronotal fossae usually reduced. Pro- notum of female usually distinctly impressed midlongitudinally. Lateral margin of pronotum more or less straight between anterolateral and lateral angles (Fig. 300) 2 2a. Elytral interstriae evenly dulled by dense sha- greening ( x 20). Smooth posterior area of pro- notal disk of male extending at least one-half distance to anterior margin of disk (best judged in large individuals) (Fig. 293). Disk of female pronotum more coarsely sculptured along mid- longitudinal impression than areas to each side. Lowland forests of Panama (Fig. 301) Phanaeus (P.) howdeni Arnaud b. Elytral interstriae evenly smooth and shining, not dulled by shagreening. Smooth posterior area of male pronotum extending less than one- half (usually about one-third) distance to an- terior margin of disk (Fig. 292). Texture of sculpturing of disk of female pronotum more or less uniform, not distinctly coarser or denser in median depression. Forests of east central Veracruz and of southern portion of Yucatan and adjacent areas of northern Chiapas and Guatemala (Fig. 301) Phanaeus (P.) sallei Harold Phanaeus (Phanaeus) beltianus Bates Figures 37, 38, 291, 294, 295, 299, 301 Phanaeus beltianus Bates, 1887:63 Type: Male lectotype (“Chontales, Nicaragua, B.C.A. p.63 sp.24, sp. figured”), British Mu- seum (Natural History), London, PRESENT DESIGNATION. OTHER REFERENCES. Nevinson, 1892a; Gil- let, 1911b; Olsoufieff, 1924; Blackwelder, 1944; Ed- monds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 37, 38). — Posterior portion of head, pronotum weakly shining yellow-green with strong coppery red reflections; elytra coppery red, some- times with strong greenish highlights. Pronotal disk of male completely colored except for black pos- terolateral angles; that of female mostly dull black. Venter with strong yellow-green reflections on legs, sterna. Pronotum. — Lateral margin bowed inward between anterolateral and lateral angles (Fig. 299). Basal fossae prominent in female, minute or effaced in male. Disk of female not inmpressed midlongi- tudinally. Pronotal Sculpturing. — Male: In large in- dividuals, sides finely granulorugose, becoming somewhat more coarsely rugose near disk, punctate beneath posterolateral angles. Disk coarsely gran- ulorugose except posteromedially where abruptly smoother and bears minute asperities. Smooth basal area partially separated from remainder of disk by straight, transverse carina. Smaller individuals as above except disk reduced to small, circular, con- cave area near anterior margin, sculpturing of sides extending onto disk as flat scale-like granules re- placed by puncturing near posterior margin. Fe- male: Sides finely granulorugose, rugosities becom- ing scale-like near disk; disk with asperate punctures 72 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus grading to weak, simple punctures posteriorly, ap- pearing smooth to unaided eye. Elytra (Fig. 295). — Striae simple or minutely punctate ( x 40). Interstriae convex, highly shining medially, dulled laterally by dense shagreening. Secondary Sexual Charac- ters.— Male: Pronotum as in Figure 291. Female: Pronotum as in Figure 294; disk not impressed mid- longitudinally. Specimens Examined. — 6 males, 6 females (length 16-20 mm; width 10-13 mm). DIAGNOSTIC REMARKS. The salient features distinguishing this species from other members of the group are microsculpturing of the elytra and form of the male pronotum. DISTRIBUTION (Fig. 301; appendix). Known only from scattered localities in Costa Rica and Chontales district of Nicaragua. Lowland evergreen forest below 500 m. Collection dates: July- August. COMMENTS. This is one of the rarer species of the genus. Howden and Young’s (1981) refer- ences to beltianus apply to howdeni. Phanaeus (Pbanaeus) howdeni Arnaud Figures 41, 42, 293, 301 Phanaeus howdeni Arnaud, 1984:61 Type: Male holotype (“Panama, Gatun Lake, Lion Flill Island”), Anne and Henry Howden Col- lection, in Canadian National Collection, Ot- tawa. OTHER REFERENCES. Howden and Young, 1981 (as beltianus ); Howden and Gill, 1987. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 41, 42). — Dorsum dark blue-green, yellow-green with weak coppery highlights, or green. Disk of female pronotum with anteromedian, two posterolateral black spots, all three of which may coalesce. Pygidium colored like elytra. Venter with colored reflections on legs, sterna. Pronotum .— Lateral margin weakly sinuate between anterolat- eral, lateral angles. Basal fossae almost effaced. Disk of female usually distinctly impressed midlongitu- dinally. Pronotal Sculpturmg. — Male: In large in- dividuals, sides granulorugose, sculpturing becom- ing weak along anterolateral margin, grading into weak puncturing (or smooth areas) beneath, behind posterolateral angles of disk. Disk coarsely rugose, with transverse ridging except for much smoother ovoid posteromedian area, which bears weak, as- perate punctures; smoother area extending at least one-half distance to anterior margin, bearing a crest- like tubercle in middle of its anterior edge. Smaller individuals as above except disk less coarsely ru- gose, smooth posteromedian area bounded ante- riorly by wide V-shaped ridge distinctly carinate only medially. Female: Sides granulorugose; sculp- turing almost effaced along anterolateral margin, becoming scale-like toward disk, grading into punc- tures posteriorly. Midlongitudinal impression of disk usually more coarsely sculptured than adjacent ar- eas to its sides. Elytra. — Striae simple to minutely punctate (x40). Interstriae convex, evenly dulled by shagreening. Secondary Sexual Characters. — Male: Pronotum as in Figure 293. Female: Prono- tum as in Figure 294. Specimens Examined. — 17 males, 15 females (length 14-19 mm; width 9-12 mm). DIAGNOSTIC REMARKS. See under sallei. DISTRIBUTION (Fig. 301; appendix). Known only from tropical evergreen forests in central Pan- ama. 0-500 m. Collection dates: May-December. COMMENTS. This species is fairly common on Barro Colorado Island, Panama Canal Zone, and nearby regions to the east and west. Howden and Young (1981) referred to it as beltianus, a much rarer species. Phanaeus ( Phanaeus ) sallei Harold Figures 39, 40, 292, 296, 297, 300, 301 Phanaeus sallei Harold, 1863:168 Type: Male lectotype (“Mexico”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:115). OTHER REFERENCES. Gemminger and Har- old, 1869; Bates, 1887; Nevinson, 1892a; Villada, 1901; Gillet, 1911b; Olsoufieff, 1924; Blackwelder, 1944; Barrera, 1969; Edmonds, 1972; Moron, 1979; Moron et ah, 1985; Palacios-Rios et ah, 1990. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 39, 40). — Dorsum shining coppery red (often combined with yellow-green highlights), yellow-green, green, or dark blue-green. Disk of female with black anteromedian spot, black pos- terolateral angles. Pygidium colored as elytra. Ven- ter dark with colored reflections on legs, sterna. Pronotum. — Lateral margin more or less straight between anterolateral, lateral angles (Fig. 300). Bas- al fossae almost effaced. Disk of female weakly impressed midlongitudinally. Pronotal Sculptur- mg.— Male: In large individuals, sides granuloru- gose, becoming smooth near anterolateral margin, punctate to smooth beneath posterolateral angles of disk. Anterior portion of disk coarsely rugose, with isolated transverse ridging. Posterior portion of disk smooth, bearing small asperate punctures; this smooth basal area separated from anterior area by transverse carina. Smaller individuals as above except disk less coarsely rugose, transverse carina separating anterior, posterior portions of disk re- placed by a weak tumosity. Female: Sides densely granulorugose, becoming smooth along anterolat- eral margin, coarser, scale-like posteriorly; middle of disk with flat, scale-like granules grading to small, simple punctures posteriorly. Midlongitudinal im- pression (if distinct) not more coarsely sculptured than areas of disk to either side. Elytra (Fig. 296). — Striae simple to minutely punctured (x40). Inter- striae evenly convex, shining; smooth to minutely punctured, occasionally minutely roughened ( x 40). Secondary Sexual Characters. — Male: Pronotum Contributions in Science, Number 443 Edmonds: Revision of PhanaeusM7 3 Figures 302, 303. Phanaeus ( Phanaeus ) amethystinus group (302, P. melampus, hind basitarsus; 303, P. ame- thystinus, same). as in Figure 292; in large specimens, smooth pos- terior area of disk extending no more than one- third (usually less) distance to anterior margin. Female: Pronotum as in Figure 294. Specimens Examined.-— 97 males, 81 females (length 14-20 mm; width 8-12 mm). DIAGNOSTIC REMARKS. This species most closely resembles howdeni, from which it is distin- guished by features of the pronota of both sexes. It also superficially resembles wagneri wagneri, from which it can be separated most easily by the struc- ture of the front femora. DISTRIBUTION (Fig. 301; appendix). Tropical forests of east-central Veracruz, southern Yucatan Peninsula and adjacent areas in northern Chiapas and Guatemala (and probably also Pionduras). 0- 1500 m. Coprophagous. Collection dates: May- October. COMMENTS. The most frequent color of sallei is coppery red, especially in populations from Ve- racruz. Examined populations from Chiapas are al- ways green. The distribution of this species is poorly known; of the over 150 specimens examined, only about 50 carry precise locality data. Since its dis- tribution is limited ecologically to lowland ever- green forests, it is likely that the geographical dis- tribution of sallei, like that of endymion, is now far more restricted and fragmented that it was in pre-European times. The Amethystinus Group DIAGNOSIS. [1] Outer margin of clypeus at most only weakly toothed medially; [2] clypeal process (seen from front) transverse rounded ridge; [3] ce- phalic process of female narrow, trituberculate Ca- rina (seen from above) on line even with anterior margins of eyes; [4] pronotal disk of large males flattened and coarsely granulorugose and (seen from above) triangular; posterolateral angles rounded, upturned; bearing smooth, raised transverse trian- gular area immediately in front of posterior margin (Figs. 307-309); [5] pronotum of female convex, granulorugose anterolaterally, with weak transverse anteromedian ridge followed, in large individuals, by small concavity; [6] basal pronotal fossae pres- ent; [7] front tibiae tridentate; [8] longer mesotibial spur of female dilated subapically; [9] mountains of Guatemala and southern Mexico (Fig. 312). This group brings together two montane species, melampus and amethystinus. Leonardo Delgado- Castillo of the Instituto de Ecologia in Xalapa, Ve- racruz, has kindly brought to my attention a new species assignable to this group being described by him (see “Comments” under amethystinus). KEY TO THE SPECIES AND SUBSPECIES OF THE AMETHYSTINUS GROUP la. Posteromedian portion of female pronotum appearing almost smooth to unaided eye; weak magnification (xlO) reveals field of shallow punctures against smooth background of dull, velvet-like shagreening (Fig. 311). Elytra dull dark brown, rarely with bluish cast (Figs. 51, 52); interstriae appearing flat or very weakly convex to unaided eye (Fig. 307); striae super- ficial. Hind basitarsus of female narrow api- cally, ridges on plantar surface more or less straight (Fig. 302). Isolated localities in Chiapas and Veracruz (Fig. 312) Phanaeus (P.) melampus Harold b. Posteromedian portion of female pronotum distinctly roughened to unaided eye (Fig. 310), bearing (xlO) coarse punctures against a roughened, shining background; if punctures almost effaced, surface shining. Elytra blue or blue-green (Figs. 53-56), rarely green or green- brown (Fig. 57), always at least partially shin- ing; interstriae appearing convex to unaided eye, striae impressed. Hind basitarsus of female widened apically; outer ridge of plantar surface usually distinctly more sinuous than inner ridge (Fig. 303). Mountainous areas of Chiapas and western Guatemala, eastern Oaxaca northward to southern San Luis Potosi Phanaeus (P.) amethystinus Harold 2 2a. At least interstriae 2 and 3, but usually 2-5, highly shining medially, dull adjacent to striae and appearing costate to unaided eye (Fig. 309). Sides of pronotum densely granulate-granulo- rugose. Highlands of Guatemala (Fig. 312) . . . . . Phanaeus (P.) amethystinus guatemalensis Harold, NEW STATUS b. All elytral interstriae evenly and usually bright- ly shining, evenly convex (Fig. 308). Sides of pronotum only moderately densely granulo- rugose. Highlands of Chiapas, mountains of Guerrero, eastern Oaxaca, western Veracruz, northern Hidalgo, and southern San Luis Po- tosi (Fig. 312) . .Phanaeus (P.) amethystinus amethystinus Harold, NEW STATUS Phanaeus {Phanaeus) melampus Harold Figures 51, 52, 302, 304, 307,311,312 Phanaeus melampus Harold, 1863:165 Type: Male lectotype (“Mexico”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:l 15). 74 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Figures 304-311. Phanaeus ( Phanaeus ) amethystinus group (304, P. melampus, male; 305, P. a. amethystinus, male; 306, same, female; 307, P. melampus, dorsal view elytron; 308, P. a. amethystinus, same; 309, P. a. guatemalensis, same; 310, P. a. amethystinus, dorsal view female pronotum; 311, P. melampus, same). OTHER REFERENCES. Gemminger and Har- old, 1869; Bates, 1887; Nevinson, 1892a; Villada, 1901; Gillet, 1911b; Olsoufieff, 1924; Blackwelder, 1944; HalfFter, 1955; Halffter and Matthews, 1966; Barrera, 1969; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 51, 52). — Dorsum somber, often ap- pearing black or dark brown without intense illu- mination, which can reveal bluish or greenish cast on pronotum or elytra or both. Pronotum. — Basal fossae present. Pronotal Sculpturing. — Male: In large individuals (Fig. 304), disk coarsely rugose; sides more densely but more finely granulorugose except along lateral margins and on anterolateral angles where almost smooth; base of disk with raised, transverse area that is smooth except for some coarse puncturing in front of fossae. In small- er individuals, disk as above except smooth basal area lacking, posterior portion of disk shallowly punctate. Female: Posteromedian portion of disk bearing shallow punctures against dull, velvety background; punctures becoming progressively Contributions in Science, Number 443 Edmonds: Revision of Phanaeus M7 5 Figure 312. Distribution of the Phanaeus ( Phanaeus ) amethystinus group (open circle = sp. nov. near amethystinus; see text). stronger near middle of disk where they coalesce to form fine transverse ridging; ridging becoming progressively stronger anteriorly. Elytra (Fig. 307). — Striae fine, shining. Interstriae weakly convex, ap- pearing almost flat to unaided eye, uniformly dull. Hind Tarsus. — Basal segment of female (Fig. 302) narrow apically, apical width <0.75 length along inner margin; both plantar ridges more or less straight. Secondary Sexual Characters.— Raised smooth area at base of male pronotum distinct only in larger individuals; otherwise as described for spe- cies group. Specimens Examined. — 45 males, 25 females (length 17-25 mm; width 11-15 mm). DIAGNOSTIC REMARKS. This species can be confused easily with amethystinus, especially smaller specimens lacking any conspicuous blue color. The most dependable features separating the two spe- cies are relief and luster of the interstriae and sculp- turing of the pronotum, particularly that of the female. Small female melampus resemble the dark phase of prasinus, from which it differs by the form of the cephalic carina. DISTRIBUTION (Fig. 312; appendix). Poorly known (see “Comments,” below). Assumed to be at lower elevations (<1400 m) along eastern slopes of Sierra Madre Oriental and northern slopes of Chiapan Highlands. COMMENTS. Almost all the specimens of me- lampus I have examined bear incomplete or im- precise locality data. Moreover, all but 4 of the 65 specimens available were, or appear to have been, collected many decades ago, and their labels, if any, bear such entries as “Mexico,” “Veracruz,” or, somewhat more precisely, “Jalapa,” “Cordova,” and other Mexican sites popular with nineteenth cen- tury collectors. In spite of the fact that these regions have been well collected in the last 25 years, I have seen no melampus from them. The lack of modern 76 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus material suggests that melampus is now extinct in many areas it once inhabited, that it occurs in some highly restricted habitat, or both. The four specimens examined collected in recent years are from two localities in Chiapas on the northern slopes of the central Highlands (appen- dix). This area supports wet tropical evergreen for- est, although it is now quite heavily disturbed by agricultural activities. Both localities are at middle elevations (ca. 1500 m). It is not unreasonable to suppose that melampus also occurs in similar, iso- lated areas in the mountainous regions from Oaxaca to northern Puebla. I suspect that its distribution parallels that of amethystinus but at somewhat lower elevations in more restricted habitats. Phanaeus ( Phanaeus ) amethystinus Harold Figures 53-57, 303, 305, 306, 308-310, 312 Phanaeus amethystinus Harold, 1863:169 Type: Male lectotype (“Orizaba”), Museum Na- tional d’Histoire Naturelle, Paris (Arnaud, 1982a:l 14). Phanaeus martinezi Halffter, 1955:75, NEW SYN- ONYMY Type: Male holotype (“Mexico: Veracruz, Pre- sidio”), personal collection of Gonzalo Halff- ter, Xalapa, Veracruz. Phanaeus guatemalensis Harold, 1871 b:l 14 Type: Male lectotype (“Guatemala”), Museum National d’Histoire Naturelle, Paris (Arnaud, 1982a:114). Phanaeus tepanensis Bates, 1889:388 (Nevinson, 1892a:4) Type: Male lectotype (“Guatemala, Tepan, Con- radt, B.C.A p.388 sp.22a, sp. figured”), British Museum (Natural History), London, PRES- ENT DESIGNATION. Phanaeus amethystinus amethystinus Harold, NEW STATUS Phanaeus amethystinus guatemalensis Harold, NEW COMBINATION OTHER REFERENCES. To amethystinus — Gemminger and Harold, 1869; Halffter and Mat- thews, 1966. To both subspecies — Gillet, 1911b; Olsoufieff, 1924; Blackwelder, 1944; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern. — (a) amethystinus (Figs. 53, 54): Dorsum shining deep blue or blue-green (elytra dull in some specimens from San Luis Potosi; see “Comments,” below), (b) guatemalensis (Figs. 55, 56): As above except some specimens completely bright green, or brown-green (“bronze”) with green highlights on sides of pronotum, coppery brown area on pronotal disk of male (Fig. 57). Pronotum. — Basal fossae present in female, sometimes effaced in male. Pro- notal Sculpturing (as follows for both subspecies but generally denser in guatemalensis). — Male : In large individuals (Fig. 305), disk coarsely rugose; sides granulorugose to granulate; base of disk (Figs. 308, 309) with raised transverse area bearing scat- tered, coarse punctures. Small individuals as above except raised basal area often replaced by coarsely punctured area. Female (Fig. 310): Posteromedian area of disk punctate to punctatorugose, rarely al- most smooth. Elytra. — Striae fine, often minutely punctate (xlO). (a) amethystinus (Fig. 308). Inter- striae more or less evenly convex, uniformly shin- ing. (b) guatemalensis (Fig. 309): Interstriae 2-3 (usually also 4-5 or 6) shining medially, dull adja- cent to striae such that interstriae appear polished midlongitudinally. Hind Tarsus. — Female basitar- sus (Fig. 303) wide, apical width >0.75 length along inner margin; outer ridge on plantar surface usually distinctly more sinuous than inner ridge. Secondary Sexual Characters. — As described for species group; raised area along base of pronotum distinct in all but smallest individuals. Specimens Examined. — amethystinus: 82 males, 59 females; guatemalensis: 138 males, 109 females (length 13-25 mm; width 8-16 mm). DIAGNOSTIC REMARKS. This species differs from melampus by the luster of the elytral inter- striae and sculpturing of the posteromedian portion of the pronotal disk of the female. DISTRIBUTION (Fig. 312; appendix). (a) ame- thystinus.— Forests and adjacent open habitats in mountains of Chiapas, Sierra Madre del Sur of Guerrero, eastern Oaxaca, and the Sierra Madre Oriental of Veracruz, northern Hidalgo and south- eastern San Luis Potosi. 1000-2200 m (usually >1500 m). (b) guatemalensis. — Forests and adja- cent open habitats in Guatemalan Highlands. 1500- 2400 m. Coprophagous; both subspecies common in meadows and clearings grazed by livestock. Col- lection dates: May-October. COMMENTS. The two subspecies recognized here have heretofore been regarded as separate spe- cies. The primary difference between the two is the microsculpturing of the elytral interstriae, which appear costate in guatemalensis. Specimens exhib- iting intermediate states in the luster of the inter- striae occur in Chiapas and, I suspect, also in far western Guatemala. Males of guatemalenis rarely lack at least some trace of the smooth basal area of the pronotum; this area is often lacking in ame- thystinus, even in large males. In addition to those characters mentioned in the descriptions above, some members of this species possess shallow fos- sae near the posterior margin of the median portion of the metasternum. These fossae are largest in guatemalensis and lacking from populations in San Luis Potosi. Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 77 Figures 313-322. Phanaeus ( Phanaeus ) quadridens group (313, P. quadridens, male; 314, same, female; 315, F. damocles, male; 316, same, female; 317, F. palliatus, male; 318, same, female; 319, F. quadridens, dorsal view female pronotum; 320, F. damocles, same; 321, F. quadridens, dorsal view elytron; 322, F. palliatus, same). Specimens from southwestern San Luis Potosi and adjacent area of Hidalgo (along Highway 85 descending toward Tamazunchale) differ from typ- ical amethystinus by being smaller, by having a distinctly duller coloration (especially on the ely- tra), and by lacking metasternal fossae. These in- dividuals are reminiscent of melampus. This pop- ulation is separated from more typical amethystinus in northeastern Hidalgo by an intervening xeric re- gion in north-central Hidalgo (the Hidalgan Des- ert), and further study may prove it worthy of sub- specific status. I have only one specimen of amethystinus from Oaxaca. Only further collection in the mountains of northern Oaxaca will settle the question of whether or not the distribution of the subspecies extends continuously from Veracruz to the south. Leonardo Delgado-Castillo (1991) of the Insti- tuto de Ecologia in Xalapa, Veracruz, has described a new species (under the name Phanaeus hlackal- leri) closely related to amthystinus from the Sierra Madre del Sur of Guerrero and Oaxaca. Specimens from the Sierra de Alquitran (1400-1670 m; open circle in Fig. 312) in central Guerrero and from the environs of Candelaria Loxicha in extreme south- ern Oaxaca are smaller (length 16-20 mm; width 8-12 mm) and very dark blue, nearly black in color. Moreover, the female pronotum bears an almost 78 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus straight, transverse carina that is weakly tuberculate medially and not followed by any trace of a con- cavity. These populations inhabit montane forests and are coprophagous. The Quadridens Group DIAGNOSIS. [1] Outer margin of clypeus at most only weakly toothed medially; [2] clypeal process (seen from front) transverse, rounded ridge; [3] ce- phalic process of female trituberculate carina in front of eyes; [4] outer margin of head distinctly notched between clypeus, paraocular areas (Fig. 325, arrow); [5] pronotum densely, coarsely granuloru- gose (Figs. 313-320; [6] pronotal disk of large males (Figs. 313, 315, 317) triangular, concave medially, with prominent posterolateral angles; [7] pronotum of female more or less evenly convex, with weak ridge shaped almost like the Greek letter omega (co) near anterior margin; [8] basal pronotal fossae pres- ent; [9] front tibiae tridentate; [10] longer mesoti- bial spur usually dilated subapically in female; [11] mountains and plateaus above 1500 m in south- central and western Mexico northward to the southwestern United States (Fig. 329). This group comprises three montane species: damocles Harold, palliatus Sturm, and quadridens (Say). The former is the rarest of the three and more closely related to palliatus, with which it is allo- patric. All three species are coprophagous. The combined range of the group is extensive, including most of the mountainous regions of Mexico west of the Isthmus of Tehauntepec. The degree of coarseness of pronotal sculpturing is greater in this group than in any other Phanaeus. Females, es- pecially, are distinguished from others in the sub- genus by the extreme coarseness of the postero- median portion of the pronotum (Figs. 319, 320). The smooth elytra (Fig. 321) with weak, almost effaced striae of damocles and quadridens, are not approached in other Phanaeus, sen. str. KEY TO THE SPECIES OF THE QUADRIDENS GROUP la. Interstriae convex; striae fine, clearly impressed (Fig. 322). Raised outer margin of pygidium usually effaced apically (Fig. 324). Dorsum bright coppery red, dark blue, or bright green (Figs. 48-50). Transverse Volcanic Range of Mexico from northern Puebla to Jalisco northward to southern Durango (Fig. 329) Phanaeus (P.) palliatus Sturm b. Interstriae flat; striae superficial, very fine, ap- pearing almost effaced to unaided eye (Fig. 321). Raised outer margin of pygidium effaced api- cally or not. Dorsal color variable, never cop- pery red. Distribution variable 2 2a. Pronotum of male as in Figure 315. Disk of female pronotum extremely coarsely puncta- torugose posteromedially (Fig. 320). Raised outer margin of pygidium often effaced or near- ly so apically (as in Fig. 324). Higher elevations Figures 323-328. Phanaeus ( Phanaeus ) quadridens group (323, P. quadridens, pygidium; 324, P. palliatus, same; 325, P. palliatus, clypeal margin {arrow indicates notch}; 326, P. quadridens, female mesotibial spurs; 327, P. damocles, longer female mesotibial spur; 328, P. pal- liatus, same). (1800-2900 m) of the Sierra Madre del Sur of Guerrero and Oaxaca, Mexico Phanaeus (P.) damocles Harold b. Pronotum of male as in Figure 313. Disk of female pronotum less coarsely punctatorugose posteromedially (Fig. 319). Raised outer mar- gin of pygidium always complete (Fig. 323). Transverse Volcanic Range from Veracruz to Jalisco, Sierra Madre Occidental from Durango to southwestern United States, Sierra Madre Oriental from Hidalgo to San Luis Potosi (Fig. 329) Phanaeus (P.) quadridens (Say) Phanaeus ( Phanaeus ) palliatus Sturm Figures 48-50, 317, 318, 322, 324, 325, 328, 329 Phanaeus palliatus Sturm, 1843:332 Type: Male holotype (“Mexico”), Zoologische Staatssammlung, Munich. OTHER REFERENCES. Harold, 1859, 1863; Gemminger and Harold, 1869; Bates, 1887; Nev- inson, 1892a; Gillet, 1911b; Olsoufieff, 1924; Black- welder, 1944; Halffter and Matthews, 1966; Bar- rera, 1969; Edmonds, 1972; Edmonds and Halffter, 1972, 1978; Moron, 1984; Moron and Zaragoza, 1976. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 48-50). — Dorsum exhibiting three Contributions in Science, Number 443 Edmonds: Revision of PhanaeusM79 Figure 329. Distribution of the Phanaeus ( Phanaeus ) quadridens group. distinct color phases: (a) bright coppery red, often with isolated greenish reflections, (b) dark blue, (c) bright green; intermediates exist among all three. Red phase predominates in most populations; blue- green most common intermediate (see “Com- ments”). Pygidium colored as dorsum, shining. Venter dark, with strong colored reflections. Pro- notal Sculpturing. — Male: Disk of larger individ- uals coarsely rugose except for raised area along posterior margin, which is smooth except for me- dian field of coarse punctures; smooth basal area of disk longest medially, its anterior angle bearing strong tooth; sides of disk rather densely, coarsely granulorugose. Disk of smaller individuals as above except smooth basal area replaced by coarsely but sparsely punctatorugose area. Female: Disk coarse- ly, densely punctatorugose; sides coarsely, densely granulorugose. Mesotibia. — Longer spur dilated subapically (Fig. 328). Elytra (Fig. 322). — Striae fine, impressed, minutely punctate (x20); interstriae convex, evenly, sparsely, finely punctate ( x 20). Py- gidium.— Raised outer margin usually effaced api- cally. Secondary Sexual Characters. —Male (Fig. 317): Pronotal disk of larger individuals concave, posterior angles rounded laterally, upturned; smooth basal area strongly toothed anteromedially. Pro- notum of smaller individuals with less well defined disk, but almost always retaining at least trace of basal tooth. Female (Fig. 318): Cephalic carina nar- row, trituberculate, well in front of eyes. Pronotum weakly concave in front of basal fossae, otherwise evenly convex, with low, smooth transverse area 80 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus near anterior margin; this transverse area bounded posteriorly by doubly bowed crease. Specimens Ex- amined.— 270 males, 193 females (length 13-19 mm; width 8-13 mm). DIAGNOSTIC REMARKS. This species is easily distinguished from other members of the group by the relief of the elytra. Superficially it resembles eximius, from which it differs in pronotal sculp- turing and distribution. DISTRIBUTION (Fig. 329; appendix). Trans- verse Volcanic Range of southern Mexico from northern Puebla to eastern Jalisco, and southern Sierra Madre Occidental of Durango. Forest mar- gins at middle to high elevations (1800-2800 m). Coprophagous. Collection dates: June-September. COMMENTS. This species appears to prefer dis- turbed habitats where forest has been cleared or severely thinned for pastureland. Such a preference suggests to me that in pre-livestock raising times, it was restricted to transition zones between forest and meadowlands. Indeed, before the careful field work of Gonzalo Halffter revealed localities where it is relatively common, palliatus was considered a rare species. I and others have collected this species in the Sierra Madre Occidental of southern Duran- go in habitat similar to that it occupies farther south. Barring its introduction into Durango along with cattle, it seems reasonable to suppose that its range between the Tranverse Volcanic Range of eastern Jalisco and the sierras of southern Durango is con- tinuous. Although there is an absence of collection records for northern Jalisco and southwestern Za- catecas, I suspect that palliatus occurs in these areas also. Phanaeus palliatus is one of the few species of the genus that exhibits distinct, almost mutually exclusive color phases. Intermediate colors do oc- cur (see “Descriptive Remarks”), but they account for less than about 5% of the observed variation in local populations. The approximate frequencies of phases among all material studied were 3% green, 27% blue, and 70% red, distributed equally be- tween the sexes. In most populations, red is by far the predominant color; a series examined from Tu- lancingo, Hidalgo, and nearby Huauchinango, Puebla, comprise about 80% blue and blue-green individuals and 20% red. This species is closely related to damocles, and their relative distributions suggest that they are vi- cariants. Contrary to my 1972 conclusion, palliatus is not a close relative of eximius, a member of the tridens group (q.v.). Phanaeus ( Phanaeus ) damocles Harold Figures 46, 47, 315, 316, 320, 327, 329 Phanaeus damocles Harold, 1863:165 Type: Male lectotype (“Mexico”), Museum Na- tional d’Histoire Naturelle, Paris (the desig- nated type locality is “Oaxaca, Capulalpam”) (Arnaud, 1982a:114). OTHER REFERENCES. Gemminger and Har- old, 1869; Bates, 1887; Nevinson, 1892a; Gillet, 1911b; Olsoufieff, 1924; Blackwelder, 1944; Bar- rera, 1969; Edmonds, 1972. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 46, 47). — Pronotum, posterior por- tion of head completely black, dark blue, or green with golden to coppery reflections on sides. Elytra weaking shining, sometimes completely black or dark blue, but usually with distinct green reflections at least along inner margins, on apices. Pronotum of male, if not entirely black, completely colored except along posterior margin; that of female sel- dom completely colored, usually mostly black on disk. Pygidium black, or colored as dorsum. Venter black, sometimes with blue or green tinges of color. Pronotal sculpturing. — Male: Disk of larger indi- viduals coarsely rugose except for raised area along posterior margin, which is smooth except for few coarse median punctures; this raised area longest medially, its anteromedian angle bearing strong, dentiform tubercle. Sides densely, coarsely granu- lorugose. Small males as above except generally less coarsely rugose, raised basal area replaced field of coarse punctures between which surface is smooth. Female (Figs. 316, 320): Disk punctatorugose, more coarsely so posteriorly, laterally; sides granuloru- gose. Mesotihia. — Longer spur dilated subapically (Fig. 327). Elytra (as in Fig. 321). — Appearing very smooth to unaided eye. Striae amost effaced; first, usually also second striae coarsely punctured; re- maining striae only weakly punctured (xlO). In- terstriae flat except for convex apical one-half of first interstria. Pygidium. — Raised outer margin usually effaced apically (as in Fig. 324). Secondary Sexual Characters. — Male (Fig. 315): In larger in- dividuals, disk concave, posterior angles rounded laterally, upturned; raised basal area strongly toothed medially. In smaller individuals, features of disk less demarcated but almost always retaining at least trace of posteromedian tooth. Female (Fig. 316): Ce- phalic carina trituberculate, set well in front of eyes. Pronotum weakly concave in front of basal fossae, otherwise evenly convex with low, smooth trans- verse area near anterior margin; this transverse smooth area bounded posteriorly by double arched crease. Specimens Examined. — 39 males, 31 fe- males (length 12-19 mm; width 8-12 mm). DIAGNOSTIC REMARKS. The form of the pronotum of both sexes strongly resembles their counterparts in palliatus, which has distinctly con- vex elytral interstriae. Females are easily confused with those of quadridens, which have a less coarse- ly sculptured pronotum and entire outer margin of the pygidium. DISTRIBUTION (Fig. 329; appendix). Pine/oak and oak forests at higher elevations (1800-2900 m) of the Sierra Madre del Sur of Guerrero and Oa- xaca. Coprophagous. Collection dates: June-Sep- tember. Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 81 COMMENTS. This uncommon species appears to be restricted to the montane forests of Guerrero and Oaxaca. It can occur above 2500 m, well above the upper altitudinal limit of most other North American Phanaeus. Phanaeus ( Fhanaeus ) quadridens (Say) Figures 43-45, 313, 314, 319, 321, 323, 326, 329 Copris quadridens Say, 1835:176 Type: Male neotype (“Mexico, D. F.”), Museum of Comparative Zoology, Harvard University (MCZ Type No. 32877), PRESENT DESIG- NATION. Phanaeus quadridens (Say), Harold, 1859:198 Phanaeus violaceus Laporte-Castelnau, 1840:81 (Harold, 1859:198) Type: Unknown to me. Phanaeus laevipennis Sturm, 1843:333 (Harold, 1859:198) Type: Female holotype (“Mexico”), Zoologische Staatssammlung, Munich. Phanaeus quadridens borealis Olsoufieff, 1924:100, NEW SYNONYMY Type: Unknown to me. OTHER REFERENCES. Lacordaire, 1856; LeConte, 1859; Harold, 1863; Blanchard, 1885; Bates, 1887; Nevinson, 1892a; Henshaw, 1885; Vil- lada, 1901; Kolbe, 1905; Gillet, 1911b; Leng, 1920; Blackwelder, 1939; Blackwelder and Blackwelder, 1948; Robinson, 1948; Halffter, 1952, 1955, 1961, 1964, 1976; Halffter and Matthews, 1966; Barrera, 1969; Edmonds, 1972; Halffter and Edmonds, 1982; Moron, 1984; Moron and Zaragoza, 1976. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 43-45). — Dorsal coloration varying between two extremes: (a) dark blue to blue-violet (sometimes almost black), often with light blue re- flections on anterolateral angles of pronotum; and (b) bright, deep green to yellow-green. Color pres- ent on posterior portion of head, sides of prono- tum, elytra; disk of pronotum usually completely colored in male and mostly black in female; elytra of blue individuals evenly colored; those of green individuals bright along periphery, dark, sometimes almost black, centrally. Pygidium colored as dor- sum. Venter almost black, with moderately shining colored highlights in blue individuals, dark with strong green reflections in green specimens. Prono- tal Sculpturing. — Male: Sides densely granuloru- gose, disk coarsely rugose except posteromedially, where it is coarsely punctatorugose. Female (Fig. 319): Sides densely granulorugose, disk punctate to punctatorugose. Mesotibia.— Longer spur not di- lated subapically (Fig. 326). Elytra (Fig. 321). — Ap- pearing smooth to unaided eye. Striae fine, often almost effaced; minutely punctured (x40), punc- tures strongest at bases of striae 1 and 2. Interstriae flat, minutely, sparsely punctured (x40), most strongly so laterally. Elytral suture sometimes bear- ing fringe of black setae. Pygidium.— Raised outer margin complete (Fig. 323). Secondary Sexual Characters.— Male (Fig. 313): Disk of pronotum of larger individuals flattened, with distinct postero- median convex area, four strong, acute tubercles; posterolateral angles strongly produced laterally, only weakly upturned. Female (Fig. 314): Cephalic carina trituberculate, narrow, set in front of eyes, often strongly raised in larger specimens. Pronotum evenly convex, distinctly concave in front of basal fossae; anteromedian prominence only weakly pro- duced, with straight or double-curved transverse carina. Specimens Examined. — 858 males, 720 fe- males (length 13-23 mm; width 8-14 mm). DIAGNOSTIC REMARKS. Females can closely resemble those of damocles, from which they differ by less coarse pronotal sculpturing, completely margined pygidium, and distribution. DISTRIBUTION (Fig. 329; appendix). Trans- verse Volcanic Range from extreme west-central Veracruz to northern Michoacan; the Altiplano of Queretaro and Guanajuato (and probably also Za- catecas); the southern Sierra Madre Oriental; and the Sierra Madre Occidental from northwest Jalisco to extreme southeastern Arizona (and adjacent ar- eas in Nayarit, Sinaloa, Sonora, and southwestern New Mexico). Grassland, mixed forest-grassland area at mid-elevations (1500-2000 m). Copropha- gous, common in places grazed by livestock. Col- lection dates: June-November (most July-Septem- ber in Mexico, August-September in Arizona). COMMENTS. The distribution of quadridens corresponds rather closely to the mountain and plateau regions which border the central, xeric in- terior of Mexico. It rarely occurs at elevations be- low 1500 m (5000 ft). Halffter (1964, and elsewhere) refers to this pattern of distribution as “dispersion en el Altiplano.” Throughout most of the southern portion of its range, it is sympatric with palliatus. Olsoufieff (1924:100) introduced the name “bo- realis” as a variety of quadridens including green specimens he observed from northwestern Mexico and the southwestern United States: “. . . il se peut que tous les 4-dens de la Sonora du Nord, Cali- fornie Sud, Arizona et N.-Mexique soient verts aus- si, en constituant une race locale (var. borealis n.), ce qui serait interessant a etablir.” Although it has never been proposed formally as such, “borealis” has often been cited as a subspecies of quadridens, a status consistent with the International Code. I have not located the type, which one would expect to find in Paris, and doubt that Olsoufieff designated one. I have chosen not to recognize borealis as a subspecies. While rare individuals in southern pop- ulations have a distinctly blue-green color, greenish 82 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus individuals do not appear regularly until central Chihuahua. Here, more than 50% of the specimens studied are blue-green or green. Farther north in Arizona, more than 85% of observed individuals are greenish and lack any trace of bluish highlights. Insofar as coloration is concerned, central Chihua- hua is a transition zone between blue populations to the south and largely green populations to the north. Other than frequency of color phases, I can find no differences between northern and southern populations of quadridens. I have seen one blue specimen of this species labeled “near Hot Springs, Las Vegas, New Mex- ico”; this record needs corroboration. Contrary to early records (e.g., Olsoufieff, 1924), quadridens does not occur in southern California. The type of quadridens was evidently among those destroyed in the 1830s (A. Newton, pers. comm.). Moreover, no specimens occur in either of the collections of T. W. Harris and the LeContes at Harvard University. The designated type locality is “Mexico,” and I assume Say himself collected his material during a sojourn there in 1827-1828 in the company of William McClure. During this time he traveled from the port city of Veracruz to Mexico City, presumably collecting along the way (Weiss and Ziegler, 1931). Since Mexico City, or more generally the Distrito Federal, is well within the known range of quadridens, and since it is reasonable to assume that Say collected there, I have chosen as neotype a male from this area. The neotype is a large violet-blue specimen with an in- conspicuous fringe of setae along the elytral suture. The vindex Group DIAGNOSIS. [1] Outer margin of head not notched laterally; [2] clypeal process a rounded, transverse ridge; [3] cephalic process of female sim- ple carina (Fig. 357) or strongly raised, apically trun- cate or emarginate, corniform process (Fig. 356); [4] anterolateral portion of pronotum strongly gran- ulorugose (Figs. 339, 340); [5] basal pronotal fossae usually distinct; [6] disk of male pronotum flat, with prominent posterolateral angles (Figs. 347-348) (re- duced in igneus, Fig. 349); [7] pronotum of female more-or-less evenly convex ( igneus , Fig. 335) or with transverse anteromedian prominence anterior to which surface abruptly declivitous (Figs. 331, 333); [8] elytral interstriae (Figs. 341-346) densely, heavily punctured; punctures often coalescing to form reticulate or longitudinal ridging; [9] front tibiae tridentate (Figs. 352, 353) (small fourth tooth in difformis. Figs. 352, 353); [10] female mesobas- itarsus not widened apically; [11] longer mesotibial spur dilated subapically; [12] pygidium densely punctatorugose; [13] eastern two-thirds of the Unit- ed States and extreme northern Mexico (Fig. 360). This group consists of three U.S. species, igneus Macleay, vindex Macleay, and difformis LeConte, distinguished from closely related groups by the sculpturing of the elytra and pronotum. The elytral sculpturing of the vindex group is most closely approached in triangularis texensis (q. v.), on the strength of which I included triangularis in the “ vindex complex” in 1972. KEY TO THE SPECIES OF THE VINDEX GROUP la. Sculpturing of disk of pronotum (Fig. 340) con- sisting of large, flattened rugosities with ill-de- fined margins such that distinct ridging at most only weakly defined; rugosities often mixed with sparse micropuncturing ( x 40) most clearly vis- ible posteriorly. Front apical tibial spur abrupt- ly bent mesally (Fig. 355). Ventral surfaces of middle, hind tibiae smooth. Elytral interstriae flat, punctatorugose (Fig. 341) to weakly con- vex, strongly, but simply punctured (Fig. 342). Male: Cephalic horn of large individuals (Figs. 334, 358) never reaching posterior margin of pronotum, inclined but only slightly curved posteriorly and compressed apically; postero- lateral prominences of pronotum broadly rounded (Figs. 334, 349), disk never presenting flat, triangular dorsal surface. Female: Anterior portion of circumnotal ridge more or less straight, not distinctly angulate medially (Fig. 338); cephalic carina low, thickened mesally and at most only barely trituberculate (Fig. 357); pronotum in profile evenly rounded from pos- terior to anterior margins; anteromedian pro- notal prominence almost always either effaced or consisting of two low, rounded tubercles (Fig. 338), rarely in form of weak transverse carina. Florida, adjacent coastal plains from eastern Louisiana to North Carolina (Fig. 360) Phanaeus ( P .) igneus Macleay b. Sculpturing of disk of pronotum consisting of well-defined, ridge-like rugosities with distinct margins (Fig. 339); micropuncturing absent. Front apical tibial spur evenly curved mesally or nearly straight (Fig. 354). Ventral surfaces of apices of four posterior tibiae finely, but distinctly rugose. Elytral interstriae flat, densely punctate to punctatorugose (Figs. 343-346). Male: Cephalic horn of large individuals usu- ally reaching or surpassing posterior margin of pronotum, evenly curved posteriorly, conical apically (Figs. 330, 332, 359); posterolateral prominences of pronotum acute, disk always distinctly flattened dorsally, more-or-less tri- angular (Figs. 347, 348). Female: Anterior por- tion of circumnotal ridge strongly angulate mesally (Figs. 336, 337); cephalic carina strong- ly raised medially as apically truncate or emar- ginate, corniform process (Fig. 356); pronotum of large individuals, seen in profile, abruptly vertical anterior to distinct transverse ridge (Figs. 331, 333). Distribution variable 2 2a. Bases of elytral interstriae 2-3 raised, smooth- er, shinier, less densely punctate than adjacent areas, never bearing distinct midlongitudinal Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 83 Figures 330-340. Phanaeus ( Phanaeus ) vindex group (330, P. vindex, male; 331, same, female; 332, P. difformis, male; 333, same, female; 334, P. igneus, male; 335, same, female; 336, P. vindex, dorsal view anterior portion female pronotum; 337, P. difformis, same; 338, P. igneus, same; 339, P. vindex, anterolateral portion female pronotum; 340, P. igneus, same). costae (Fig. 346); interstriae otherwise densely punctatorugose, punctures coalescing to pro- duce irregular, usually reticulate (as opposed to longitudinal) ridging (Figs. 344, 346). Elytral striae 1-2 narrow, widths near base usually less than one-fifth widths of interstriae 2-3, re- spectively. Circumnotal ridge irregularly ser- rate behind anterolateral angle (Fig. 348B). Front tibiae quadridentate, fourth (basal) tooth small but distinct in all but worn specimens (Figs. 352, 353). Male (Figs. 332, 348): In large individuals, sides of disk of pronotum curved; posterolateral angles of disk elongate, their tips (seen from above) not extending beyond lateral margin of pronotum. Female: In large individ- uals, transverse, anteromedian ridge of pro- notum effaced medially, from above (Fig. 337) usually bowed posteriorly, not bounded pos- teriorly by complete, carina-like crease; in small specimens, anteromedian pronotal prominence reduced to two isolated, rounded tubercles. South-central United States, extreme north- 84 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Figures 341-346. Phanaeus ( Phanaeus ) vindex group (341, 342, P. igneus, dorsal view elytron; 343, P. vindex, same; 344, P. difformis, same; 345, P. vindex, same; 346, P. difformis, same). eastern Mexico (Fig. 360) Phanaeus (P.) difformis LeConte b. Elytral interstriae 2-3, sometimes also 4-5, each with single, strong midlongitudinal costa (Figs. 343, 345); costae of interstriae 2-3 often ex- tending length of elytra; those of interstriae 3- 4, if present, usually less than one-half length of elytra. Sculpturing of interstriae 2-3 con- sisting otherwise of ridge-like rugosities often joining to form fine ridges which parallel costae (Fig. 345); remainder of interstriae densely punctatorugose to granulorugose. Elytral striae broad, flat (xl5), widths near base usually greater than one-fifth those of interstriae 2-3, respectively. Circumnotal ridge simple behind anterolateral angle (Fig. 347B); rarely with one or two small teeth or (in some Texas and Ar- izona specimens) with weak serration. Front tibiae tridentate (Figs. 350, 351), fourth (basal) tooth sometimes suggested by indistinct an- gulation of tibial margin or rarely distinct. Male: In large individuals, sides of pronotal disk straight, longitudinal axes of posterolateral an- gles diverging posteriorly such that angles (seen from above, Fig. 347A) project to or slightly beyond lateral margin of pronotum. Female: Transverse anteromedian prominence of pro- notum complete, seen from above (Fig. 336), straight, bounded posteriorly by uninterrupted, carina-like crease; in very small individuals, prominence reduced to two isolated, flat, rounded tubercles. Widely distributed in the eastern half and southwestern portions of the United States and in extreme northern Mexico (Fig. 360) . . . Phanaeus (P.) vindex Macleay Phanaeus ( Phanaeus ) igneus Macleay Figures 104-107, 334, 335, 338, 340-342, 349, 355, 357, 358, 360 Phanaeus igneus Macleay, 1819:133 Type: Male neotype (“Georgia”), Macleay Mu- Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 85 Figures 347-359. Phanaeus ( Phanaeus ) vindex group (347, P. vindex, male pronotum [A = dorsal view; B = anterolateral margin]; 348, P. difformis, same; 349, P. igneus, same; 350, P. vindex, front tibia, male; 351, same, female; 352, P. difformis, front tibia, male; 353, same, female; 354, P. vindex, front tibial spur; 355, P. igneus, same; 356, P. vindex, female cephalic process [A = lateral view; B = frontal view]; 357, P. igneus, same; 358, P. igneus, lateral view male cephalic process; 359, P. vindex, same). seum, University of Sydney, Sydney (Type No. MMINV 8), PRESENT DESIGNATION. Phanaeus floridanus Olsoufieff, 1924:94, NEW SYNONYMY Type: Male lectotype (“St. John’s Bluff, E. Flor- ida”), Museum National d’Histoire Naturelle, Paris (Arnaud, 1982a:116). OTHER REFERENCES. Blanchard, 1885; Nev- inson, 1892a; Henshaw, 1885; Gillet, 1911b; Leng, 1920; Hayden, 1925; Cartwright, 1934, 1939; Brim- ley, 1938; Blackwelder, 1939; Loding, 1945; Robinson, 1948; Blackwelder and Blackwelder, 1948; Miller, 1961; Miller et al., 1961; Howden, 1955; Halffter and Matthews, 1966; Fincher et al., 1969, 1970, 1971; Edmonds, 1972; Fincher, 1973a, b, 1975a, b, 1979, 1981; Woodruff, 1973. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 104-107).— Dorsum brightly shining to dull; color highly variable: (a) uniformly black, dark blue to blue-green, color occasionally more pronounced on pronotum; to (b) pronotum cop- pery red, elytra bright green to coppery red. Many intermediates occur, particularly in shades of col- oration, combinations of pronotal and elytral col- ors. Pygidium colored as elytra. Venter dark, with few muted colored reflections to brighter with strongly shining colored areas on legs, sterna. Pro- notum.— Anterolateral margin not serrate. Basal fossae usually at least indicated. Pronotal Sculp- turing.— Male: In large individuals, sides rather fine- ly, sparsely to more coarsely, densely granuloru- gose; sculpturing coarsest anterolaterally, clearly finer below lateral fossae. Central area of disk coarsely rugose, rugosities flattened, sometimes with ill-defined margins so as to appear partially effaced; widely scattered micropunctures usually visible (x30) in spaces separating rugosities; micropunc- tures usually most distinct in shiniest specimens. Sculpturing of posterolateral portions of disk weak- er than in center of disk, often grading into widely scattered, simple punctures. Disk somewhat raised posteromedially; this raised area impressed midlon- gitudinally, distinctly more coarsely sculptured (punctatorugose) than adjacent areas. In smaller in- dividuals, sculpturing as above except that disk not raised posteromedially, area of disk behind line connecting remnants of posterolateral angles rather coarsely, densely punctatorugose medially, punc- tate to granulorugose laterally. Female (Figs. 338, 340): As in male except posterior two-thirds of disk, in an approximately triangular area whose apices Edmonds: Revision of Phanaeus 86 ■ Contributions in Science, Number 443 are anteromedian prominence, posterolateral an- gles, coarsely, rather densely punctate to granulo- rugose; sculpturing denser in, near middle of disk, sometimes partially effaced in shiniest specimens. Disk weakly, but usually perceptibly impressed midlongitudinally from posterior margin to middle. Tibiae. — Front tibiae tridentate; fourth (basal) tooth at most only weakly indicated; apical spur abruptly bent mesally (Fig. 355). Ventral surface of apices of middle, hind tibiae smooth. Elytra. — Striae sim- ple to minutely punctured ( x 30). Interstriae varying between two extremes: (a) (Fig. 342) all interstriae appearing uniformly shining, convex to unaided eye; first interstria smooth, basal one-third to one-half of second and third interstriae with few, widely scattered, small simple punctures or largely smooth; remaining interstriae, distal portions of second and third distinctly, uniformly densely punctured, these punctures round, separated by shining raised areas, rarely confluent; (b) (Fig. 341) first interstria smooth, convex; other interstriae appearing uniformly dull, flat to unaided eye but under magnification (xl5) appearing uniformly, densely punctatorugose, punctures large, flat, coalescing frequently to pro- duce reticulate pattern of fine, shining rugosities. Secondary Sexual Characters. — -Male (Figs. 334, 349): In largest individuals, pronotal disk flattened medially, posterolateral angles produced as broad, slightly upturned rounded lobes; in smallest indi- viduals, pronotum almost evenly convex, postero- lateral angles reduced to round tubercles. Cephalic horn, even in largest individuals, extending little beyond middle of disk; in smallest individuals, re- duced to small, weakly emarginate tubercle. Fe- male: Cephalic carina narrow, thickened mesally, usually simple (faintly trituberculate in some spec- imens showing no wear). Anteromedian promi- nence of pronotum (Fig. 338) usually weak, often appearing as two separated tubercles or altogether absent; at most (rarely so even in largest individuals) weak, thick, transverse ridge; pronotum in profile more or less evenly rounded from posterior to an- terior margins. Specimens Examined.— 896 males, 722 females (length 13-22 mm; width 7-14 mm). DIAGNOSTIC REMARKS. This species is easily distinguished from its U.S. relatives by its pronotal and elytral sculpturing and secondary sexual char- acters. DISTRIBUTION (Fig. 360; appendix). Coastal plains of the southeastern United States from ex- treme southeastern Louisiana through extreme southern portions of Mississippi and Alabama, southern and southwestern Georgia, eastern South Carolina, southeastern North Carolina, and all of Florida except the keys. 0-100 m. Usually associ- ated with sandy soils and pine or mixed pine forests. Coprophagous (rarely necrophagous). Collection dates: March-December (most March-September). COMMENTS. Almost everything about the gen- eral appearance of igneus suggests a deemphasis of Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 87 what are the salient features of the vindex group — that is, an igneus looks like a vindex which has been subjected to some kind of evolutionary eraser! The most obvious “erasures” have affected sec- ondary sexual characters and sculpturing. Male ig- neus are never as well developed as large (“major”) individuals of other species; nor are female char- acters ever strongly developed. Moreover, dorsal sculpturing, while similar to that of vindex and difformis, is never sharply defined and, therefore, never appears coarse or dense. This species is restricted to the coastal plains of the United States. It probably originated in the Flo- ridian refugium created during Pleistocene episodes of glaciation. The northern limit of its distribution coincides with the presumed coastline of the south- eastern United States during maximum interglacials (Howden, 1963). The Florida peninsula and adja- cent areas of neighboring states are characterized by predominently sandy soils supporting pine and mixed pine forests. P. igneus is sympatric with vin- dex throughout its range, although this latter spe- cies prefers soils with a significant clay content. It has been collected from many types of excrement and is attracted in large numbers to pitfall traps baited with propionic acid. Several features of igneus that vary geographi- cally prompted Woodruff (1973) to recognize two subspecies: igneus igneus and igneus floridanus. The extremes in variation are, indeed, distinct; but in my opinion, the variation of this species is not predictable enough geographically to warrant rec- ognition of subspecies. Of particular interest in this context is the variability of the elytra in color and sculpturing. Specimens from Louisiana, Mississippi, Alabama, and the panhandle of Florida west of the Chatahoochee River are always subdued in color (ordinarily dull green or blue-black) and have flat elytral interstriae rather densely covered by flat, confluent punctures. The elytral sculpturing of these specimens is reminiscent of that of triangularis tex- ensis. Specimens from peninsular Florida are almost always brightly colored and have weakly convex interstriae with simple punctation (certain speci- mens from Miami resemble specimens from west of the Chatahoochee River). Individuals from Georgia northward are essentially intermediate, al- though they generally are more somber colored and have more densely sculptured elytral interstriae than peninsular specimens. Regardless of whether or not subspecies are recognized, it is evident that igneus has undergone regional differentiation, perhaps in response to climatic and geographical oscillations associated with cycles of Pleistocene glaciation. Differences in the pronotal sculpturing between igneus and the two other members of the group, while consistent, are subtle. In general, the rugos- ities of igneus lack clear definition when compared to the much more sharply defined granules and ridges found in vindex and difformis. The whereabouts of Macleay’s type material of igneus is uncertain. I have examined the purported types of this species on deposit at the Australia National Insect Collection (Commonwealth Sci- entific and Industrial Research Organization, Divi- sion of Entomology, Canberra) and cited by Britton and Stanbury (1981) as “ igneus Macleay 1833.” These two specimens are labeled “syntypes” and include a male and female transferred to Australia National Insect Collection from the Macleay Col- lection in Sydney. The female of igneus was un- known to Macleay; the male conforms to the orig- inal description but no better so than certain other specimens that remain in the Macleay Collection. The Macleay Collection, graciously loaned to me for study by Dr. D. S. Horning, Jr., its curator, includes 11 specimens bearing labels in Macleay’s handwriting. It is certain that not all these speci- mens were at hand when Macleay described igneus. His original description treats only the male, and nothing about it suggests that he observed more than a single, small specimen. It is my opinion, therefore, that the series in the Macleay Museum along with the pair in Canberra cannot be consid- ered syntypic, and that the only reasonable course of action is to choose a neotype from among them. From among the specimens in the Macleay Col- lection, I have chosen a small male labeled “car- nifex Georgia” as the neotype. It conforms to the original description in all respects and is the only specimen labeled Georgia, the designated type lo- cality. Phanaeus ( Phanaeus ) difformis LeConte Figures 101-103, 332, 333, 337, 344, 346, 348, 352, 353, 360 Phanaeus difformis LeConte, 1847:86 Type: Male neotype (no data), Museum of Com- parative Zoology, Harvard University (MCZ Type No. 32896), PRESENT DESIGNA- TION. Phanaeus difformis magnificens Robinson, 1948: 302, NEW SYNONYMY Type: Male holotype (“Romeo, Florida”), Na- tional Museum of Natural History, Washing- ton, D.C. (Type No. 65634). NOMENCLATURAL REMARKS. Woodruff (1973:62) synonymized magnificens with vindex. Robinson’s holotype, however, is a well-developed male difformis that was very likely mislabeled; it is probably from Texas. OTHER REFERENCES. Lacordaire, 1856; LeConte, 1863; Gemminger and Harold, 1869; Blanchard, 1885; Nevinson, 1892a; Henshaw, 1885; Gillet, 1911b; Leng, 1920; Olsoufieff, 1924; Brown, 1927; Lindquist, 1933; Blackwelder and Black- welder, 1948; Dillon and Dillon, 1961; Halffterand Matthews, 1966; Barrera, 1969; Edmonds, 1972; Blume and Aga, 1976, 1978; Nealis, 1977; Fincher, 88 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus 1981; Fincher and Marti, 1982; Howden and Scholtz, 1986. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 101-103. — Two dorsal color phases: (a) posterior portion of head, sides of prothorax, elytra, pygidium weakly shining dark blue; disk of pronotum weakly shining dark green with feeble yellow highlights; and (b) posterior portion of head, sides of prothorax, elytra, pygidium strongly shin- ing green with conspicuous yellow highlights; disk of pronotum coppery red. Intermediates dark green to blue-green on head, elytra, coppery red with strong yellow highlights on pronotum. Pronotum of male completely colored; that of female black and adjacent to anteromedian prominence. Venter dark with strongly colored (blue or green) reflec- tions on posterior four femora. Clypeus.— Weakly, but perceptibly broadly bidentate. Pronotum.— Anterolateral margin serrate (Fig. 348B). Anterior margin angulate medially (Fig. 337). Basal fossae usually present; rarely absent in large males. Prono- tal Sculpturing.— Male: In large individuals, sides densely, fairly coarsely granulorugose; disk more coarsely but less densely granulorugose except for nearly semicircular, coarsely, densely punctatoru- gose posteromedian area bounded anteriorly by dis- tinct, transverse semicircular ridge. Smaller individ- uals as above except posteromedian ridge absent or distinct only medially, punctatorugose area rel- atively larger. Female: Sides granulorugose; disk coarsely, densely punctatorugose, most strongly so medially where surface impressed longitudinally. Tibiae. — Front tibiae quadridentate in both sexes (Figs. 352, 353), most obviously so in male; fourth (basal) tooth small, but distinct in all but most worn specimens. Ventral surface of apices of posterior four tibiae finely rugose (xl5). Elytra (Figs. 344, 346). — -Striae simple to minutely punctate ( x 40); narrow, basal widths of second and third striae less than one-fifth width of second and third interstriae, respectively. First interstria narrow, evenly, strongly convex, smooth; remaining interstriae flat, coarsely, densely punctatorugose except for bases of inter- striae 2-3, which are widely raised, usually much less densely punctured than elsewhere. Basal one- half to one-third of interstriae 2-3 appearing smooth, shining to naked eye; these raised, smooth areas rarely extending more than one-half length of interstria; width of these raised areas at least one-half, usually three-fourths or more of total width of respective interstriae, only rarely narrow enough to produce distinct costa. Interstriae 4-5 always densely granulorugose. Secondary Sexual Features.- — Male: Pronotum of large individuals as in Figures 332, 348. Female: Cephalic carina raised medially as apically truncate or emarginate corni- form process (as in Fig. 356). Anteromedian prom- inence of pronotum medially, lacking complete ca- riniform crease behind anterior margin, seen from above (Fig. 337), usually distinctly bowed posteri- orly; in smallest individuals, anterior prominence reduced to two round flat tubercles. Specimens Ex- amined.— 402 males, 285 females (length 13-23 mm; width 7-14 mm). DIAGNOSTIC REMARKS. Red-green individ- uals often closely resemble vindex, from which they differ by elytral sculpturing, shape of the front tibiae, and, in large individuals, configuration of the pronotum. DISTRIBUTION (Fig. 360; appendix). South- central United States and extreme northeastern Mexico. 0-1200 m. Coprophagous; open habitats with predominantly sandy soils. Collection dates: March-December (most April-October). COMMENTS. LeConte’s description of diffor- mis was based on a single small male. The LeConte Collection in the Museum of Comparative Zool- ogy, Flarvard University, includes 12 specimens of difformis, one of which is labeled “Type 3709.” This latter specimen does not conform with Le- Conte’s description and cannot be regarded as the holotype. Moreover, since the description was based on a single specimen, the series cannot be regarded as syntypic. Among these specimens, however, is one that conforms well with the original descrip- tion and that I hereby designate as neotype. This species is especially common in the sandier areas of the coastal plains of Texas and adjacant area of Louisiana and of the eastern Edwards Pla- teau. It is sympatric with vindex in all but the cen- tral Texas portion of its distribution (see “Com- ments” under vindex.) Its range extends northward and westward along major river drainage systems, via which it reaches Arkansas, Kansas, and Colo- rado (Arkansas River), the Texas Panhandle (Red River), and southeastern New Mexico (Pecos River). I have seen one record from El Paso, which if not an error, suggests that it has also followed the Rio Grande westward through the Big Bend region of Texas and Mexico; I have not, however, seen spec- imens recorded from the Rio Grande valley be- tween El Paso and the mouth of the Pecos River. Locality records about which I have serious doubts include Saltillo, Coahuila (Mexico), and Romeo, Florida (the designated type locality of magnifi- cens). The blue-elytra phase of difformis appears to have a simple, intrapopulational genetic basis (Blume and Aga, 1976, 1978); it occurs only in the coastal plains. Aspects of the ecological and genetic rela- tionships between difformis and vindex are dis- cussed below. Phanaeus ( Phanaeus ) vindex Macleay Figures 96-100, 330, 331, 336, 339, 343, 345, 347, 350, 351, 354, 356, 359, 360 Phanaeus vindex Macleay, 1819:133 Type: Male neotype (“North America”), Mac- leay Museum, University of Sydney, Sydney Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 89 (Type No. MMINV 7), PRESENT DESIG- NATION. Scarabaeus carnifex Linnaeus, 1767:546 (not 1758, vol. 1:346) (Barber, 1928:383) (permanently unavailable name; see “Nomenclatural Re- marks”) Type: Male lectotype; 3 male, 1 female paralec- totypes (no data), Linnean Society, London, PRESENT DESIGNATION. Phanaeus vindex cyanellus Robinson, 1938:107 (Woodruff, 1973:62) Type: Male holotype (“Chokolostra, Florida”), National Museum of Natural History, Wash- ington, D.C. (Type No. 65636). Phanaeus vindex rubervirens Robinson, 1948:301, NEW SYNONYMY Type: Male holotype (“Chiricahua Mts., Cochise Co., Arizona”), the Academy of Natural Sci- ences of Philadelphia (Type No. 10700). NOMENCLATURAL REMARKS. The great majority of references to this species prior to about 1965 are under the name carnifex L. in spite of the fact that Barber pointed out the confusion attend- ing the name almost 40 years earlier (see also Leng, 1927). The name carnifex is now properly applied only to the Jamaican species, Sulcophanaeus car- nifex (L.) (Matthews, 1966), and, in accordance with Article 52(b) of the International Code, Scar- abaeus carnifex Linnaeus, 1767, is a permanently unavailable name. In 1972 I suggested that recourse to the International Commission on Zoological Nomenclature was in order to fix usage of the names carnifex and vindex. I did not follow through be- cause it was not necessary. Matthew’s (1966) action of designating a lectotype for carnifex L., 1758, fixed the identity of the Jamaican species with the result that vindex remained as the only available name for this North American species. Robinson’s (1948) magnificens, which Woodruff (1973:62) syn- onymized with vindex, is in fact difformis (q.v.). OTHER REFERENCES. To carnifex: Drury, 1770; Say, 1823; Laporte-Castelnau, 1840; Le- Conte, 1847, 1859, 1863; Lacordaire, 1856; Gem- minger and Harold, 1869; Blanchard, 1885; Nev- inson, 1892a; Henshaw, 1885; Blatchley, 1910; Gil- let, 1811b; Leng, 1920; Olsoufieff, 1924; Hay- den, 1925; Blatchley, 1910; Brown, 1927; Lind- quist, 1933; Cartwright, 1934, 1939; Brimley, 1938; Loding, 1945; Robinson, 1948. To vindex: Becton, 1930; Leng and Mutchler, 1933; Cartwright, 1934; Blackwelder, 1939; Ritcher, 1945, 1966; Black- welder and Blackwelder, 1948; Virkki, 1959; Dillon and Dillon, 1961; Miller et ah, 1961; Stewart and Kent 1963; Matthews, 1966; Halffter and Mat- thews, 1966; Stewart, 1967; Stewart and Davis, 1967; 90 ■ Contributions in Science, Number 443 Barrera, 1969; Fincher et ah, 1969, 1970, 1971, 1981; Edmonds, 1972; Edmonds and Halffter, 1972, 1978; Fincher, 1972, 1973a, b, 1975a, b, 1979, 1981; Blume and Aga, 1976, 1978; Fincher and Marti, 1982; Moron, 1984; Howden and Scholtz, 1986; Ratcliffe, 1991. DESCRIPTIVE REMARKS. Color and Color Pattern (Figs. 96-100). — Dorsum brightly shining, usually tricolored as follows: posterior parts of head, sides of pronotum golden yellow to yellow-green, disk of pronotum coppery red, elytra golden green to dark green; rarely (a) almost completely dark blue or nearly black or (b) head, pronotum com- pletely golden green, elytra dark blue with green reflections. Variations in shades, distribution of col- ors together produce many intermediate combi- nations. Pygidium colored like elytra. Venter with strong colored reflections on legs, often also on sterna. Clypeus. — Tvenly rounded, rarely with two distinct median teeth. Pronotum. — Anterolateral margin simple or weakly serrate (Fig. 347B). An- terior margin in female angulate medially, forming posteriorly directed angulation (Fig. 336). Basal fos- sae usually at least indicated in female, usually ef- faced in male. Pronotal Sculpturing. — Male: In larger individuals, sides coarsely granulorugose; disk somewhat more rugose than sides with frequent transverse ridging; raised posteromedian area sep- arated from remainder of disk by usually ill-defined bowed ridge, strongly punctato- to granulorugose. Smaller individuals as above except bowed ridge reduced or absent. Female (Figs. 336, 339): Sides granulorugose, disk more coarsely rugose than sides with frequent ridging of irregular orientation. Tib- iae.— Front tibiae (Figs. 350, 351) tridentate in both sexes; fourth (basal) tooth sometimes indicated, but rarely distinct; apical spur approximately straight, apex not strongly bent mesally. Apices of four pos- terior tibiae minutely rugose (x 15) on ventral sur- faces. Elytra (Figs. 343, 345).- — Striae minutely punctate ( x 30); widths at base of second and third striae more than one-fifth width (usually one-third to one-fourth) of second interstria. First interstria smooth, narrow, evenly, strongly convex; remain- ing interstriae flat, coarsely sculptured as follows: interstriae 2-3 costate medially for at least one- half, often entire length; basal width of second interstrial costa seldom exceeding one-half width of interstria itself; basal width of third interstrial costa never exceeding width of interstria itself; sec- ond and third interstrial costae each flanked by several fine ridges; interstriae 3-4 either densely punctatorugose, or puncturing obsolete such that relief of interstria appears fragmented into series of irregular ridges, granules; bases of interstriae 3-4 sometimes costate, but these costae usually shorter, less well defined than those on interstriae 2 and 3. Secondary Sexual Characters.- — Male: Pronotum as in Figures 330, 347. Female: Cephalic process raised, apically truncate or weakly emarginate cor- niform projection (Figs. 333, 356); anteromedian process of pronotum (seen from above) straight, Edmonds: Revision of Phanaeus bounded by complete, cariniform crease (Fig. 336) or (in small individuals) reduced to two small, round, flat tubercles. Specimens Examined. — 1504 males, 1624 females (length 11-22 mm; width 7-13 mm). DIAGNOSTIC REMARKS. This species, es- pecially females and smaller males, can closely re- semble F. difformis. Features of the elytral striae, sculpturing of the interstriae and the front tibiae will, in combination, separate practically all “look- alikes”; secondary sexual characters will distinguish large individuals. DISTRIBUTION (Fig. 360; appendix). United States east of the Continental Divide and south of about 43 degrees N latitude except the Edwards Plateau and far western regions of Texas; occurring west of the divide only in New Mexico, Arizona, and northern Chihuahua, Mexico. 0-1400 m. Co- prophagous, occasionally collected from carrion. Highly eurytopic. Collection dates: March-Octo- ber. COMMENTS. The distribution of vindex is both geographically and ecologically very broad, from the Sonoran Desert region of Arizona, to subtrop- ical Florida, to New England. Only rarely has it been collected north of about 43 degrees N lati- tude. Blume and Aga (1978) recorded vindex from southeastern Montana, and it is reasonable to as- sume it also occurs in South Dakota. I have seen one specimen labeled “Ottawa, Ont[ario],” but its occurrence in Canada needs confirmation (Henry Howden, pers. comm.). This species is sympatric with difformis except in the Edwards Plateau and far west regions of Texas, and its distribution com- pletely overlaps that of igneus in the southeastern United States. Many characters of vindex vary geographically, but none in ways predictable enough to support, in my opinion, formal recognition of subspecies. Individuals collected east of the Mississippi and south of the Ohio rivers tend to be darker and less shining than those collected to the west and north; those collected east of about 90 degrees W longi- tude tend to have more sharply defined costae on interstriae 2-3 and obscure to no distinct punctur- ing on interstriae 4-5. Other variations, such as those upon which Robinson based cyanellus and rubervirens, appear to be highly localized or re- peated in widely disjunct areas. Blume and Aga (1976, 1978) reported successful interbreeding between vindex and difformis in the laboratory. Both they and I have observed what appear to be intermediates among field collections, and it could very well be that these two species hybridize under natural conditions. Careful field studies to test the hypothesis have not been done, but one excellent place to conduct a study of the interactions between these two species would be in the area of Medora, Reno County, Kansas, where both species and intermediates occur fairly com- monly. All Medora “intermediates” I have seen, however, are vindex which resemble difformis, never vice versa. While they always strongly resem- ble difformis in color, size, and general appearance, and are easily confused on first (and often also on second) glance, Medora vindex are always clearly identifiable as such. I have not observed interme- diates between vindex and either of the other spe- cies with which it is sympatric, igneus and trian- gularis. Macleay (1819) based his description of vindex on at least two specimens, a female and a small male, that he regarded as a variety of Scarabaeus carnifex L., 1867. The Macleay Collection in Syd- ney contains eight vindex, none of which is iden- tifiable as a type. From these eight I have selected a small male labeled “N. America” conforming to the original description that I hereby designate as neotype. Incertae Sedis Phanaeus labreae (Pierce) Palaeocopris labreae Pierce, 1946:130 Type: Holotype male (“Rancho La Brea Tar Pits, Los Angeles, California”), Natural History Museum of Los Angeles County, Los Angeles (Type No. LACMIP 3059 = C116a). Phanaeus labreae (Pierce), Miller et al., 1981:627 OTHER REFERENCES. Halffter, 1959; Mat- thews, 1961; Matthews and Halffter, 1968; Miller, 1983. COMMENTS. The holotype of Phanaeus la- breae, the anterior portion of the head of a small male, is among the numerous insect fragments col- lected by W. D. Pierce from deposits in Pit 81 of the Rancho La Brea Tar Pits in Los Angeles, Cal- ifornia. Miller et al. (1981), in their reevaluation of Pierce’s species of Scarabaeidae, assigned labreae to Phanaeus while pointing out that the leg frag- ments included in the original description belong to other genera ( Copris and, perhaps, Deltochilum and Onthophagus). I agree with their conclusion. Lurther, I believe it highly likely that labreae is assignable to an extant species, most probably vin- dex. The latter species now occurs in Arizona. Pos- itive identification of the La Brean species can be made if well-preserved elytra, prothoraces, or both are later discovered among the many, yet unex- amined remains. Rancho La Brea, the type locality of the Ran- cholabrean Land Mammal Age, comprises perhaps the world’s best known late Pleisotcene deposits. The site records roughly 30,000 years of the history of a rich vertebrate fauna that became extinct some 10,000 years ago (Kurten and Anderson, 1980). Bone collagen of Equus from Pit 81 (not associated with labreae ) has yielded as radiocarbon date of ca. 11,000 yr BP. Most La Brean insect species described by Pierce have been synonymized with extant taxa (Miller, 1983). However, two other dung beetles described by Pierce (1946), Copris pristinus and Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 91 Onthopbagus everestae, are regarded as extinct spe- cies by Miller (op. cit.). Miller et al. (1981) pointed out that the paratype legs of everestae are assignable to Canthon. Collectively, the conclusions of Miller and Miller et al. suggest that the Los Angeles dung beetle fauna of La Brean times was very similar to that we now see in southeastern Arizona. The de- mise of the La Brean dung beetle fauna was no doubt provoked by both the climatic changes and the extinction of the contemporaneous fauna of large land mammals. Phanaeus^) antiquus Horn Phanaeus antiquus Horn, 1876:245 Type: Unknown to me. COMMENTS. Horn based his description of antiquus on portions of elytra and abdominal ster- na from “. . . some masses of clay from which were obtained many fragments of Coleoptera” from Port Kennedy Cave, Montgomery County, Pennsylva- nia. Port Kennedy Cave is a classic Pleistocene site studied by E. D. Cope, whom Horn accompanied when he collected this species as well as another dung beetle from the same deposit described as Choeridium (now Ateuchus ) ebeninum. Kurten and Anderson (1980) refer Port Kennedy Cave deposits to the Aftonian or early Kansan intervals of the late Irvingtonian Land Mammal Age (mid-Pleistocene; ca. 900,000 yr BP). I have not seen Horn’s specimens and, therefore, cannot judge if antiquus is indeed referrable to Phanaeus. His description is based on the elytra, which suggested to him “[a] species . . . somewhat larger than [P.] carnifex [= vindex ]” whose elytral sculpture more closely resembled that of [P.] pluto (now placed in Coprophanaeus). Copropbanaeus pluto is a Mexican species of neotropical origin that, in my opinion, has penetrated only very re- cently (Holocene) into northern Mexico and ex- treme southern Texas (see biogeographical com- ments in the introduction). If antiquus is a Phanaeus, Horn’s description suggests it is more likely to be triangularis, sen. str., or its relative. ACKNOWLEDGMENTS This study would have been impossible without the con- siderable cooperation of many colleagues, whose kind- nesses I deeply appreciate. I have received continuous support from my university in many ways. I also acknowl- edge with great appreciation the support of the Instituto de Ecologia in Xalapa, Veracruz, where I have a long association. This project has received direct support from funding provided to the Instituto by the Secretaria de Educacion Publica under the program Ecoetologia Ani- mal en el Estado de Veracruz en Ambientes Naturales y Perturbados (subproject: Ecoetologia de Escarabajos de Estiercol (Coleoptera: Scarabaeinae) en Selvas y Pastizales Aledanos). I have profited much from the careful review of the manuscript by Gonzalo Halffter, and Miguel Angel Mo- ron. Their efforts have considerably improved the final product. The color photographs (Figs. 1-107) are the work of J. Mark Rowland, whose assistance I gratefully ac- knowledge. The following individuals have graciously provided the material upon which this revision is based from their personal collections and those collections under their care: Kenneth Ahlstrom, North Carolina Insect Survey, Ra- leigh, North Carolina Patrick Arnaud, Saintry-Seine, France Donald Azuma, The Academy of Natural Sciences of Philadelphia, Philadelphia, Pennsylvania M. E. Bacchus, British Museum (Natural History), Lon- don Julian Blackaller, Mexico, D.F. H. D. Blocker, Kansas State University, Manhattan, Kan- sas Richard R. Blume, USDA, College Station, Texas M. Brancucci, Museum d’FIistoire Naturelle, Basel, Swit- zerland Horace R. Burke, Texas A&M University, College Sta- tion, Texas George W. Byers, University of Kansas, Lawrence, Kansas Yves Cambefort, Museum National d’Histoire Naturelle, Paris Joan B. Chapin, Louisiana State University, Baton Rouge, Louisiana Jose A. Clavijo A., Univerdidad Central de Venezuela, Maracay Cleide Costa, Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo, Brazil Charles V. Coveil, University of Louisville, Louisville, Kentucky Roger Damoiseau, Insitut Royal des Sciences Naturelles de Belgique, Brussels Robert L. Davidson, Carnegie Museum of Natural His- tory, Pittsburgh, Pennsylvania Leonardo Delgado C., Instituto de Ecologia, Xalapa, Ve- racruz, Mexico R. M. Dobson, University of Glasgow, Glasgow, Scotland W. A. Drew, Oklahoma State University, Stillwater, Okla- homa S. Endrodi, Hungarian Natural History Museum, Buda- pest G. Truman Fincher, USDA, College Station, Texas Eric R. Fisher, California Department of Food and Ag- riculture, Sacramento, California G. S. Forbes, New Mexico State University, Las Cruces, New Mexico Saul Frommer, University of California, Riverside, Cali- fornia Bruce Gill, Carleton University, Ottawa, Canada James E. Gillaspy, Texas A&I University, Kingsville, Texas John D. Glaser, Baltimore, Maryland Robert D. Gordon, National Museum of Natural History, Washington, D.C. Gonzalo Halffter, Instituto de Ecologia, Xalapa, Veracruz, Mexico Alan R. Hardy, California Department of Food and Ag- riculture, Sacramento, California Lee H. Herman, Jr., American Museum of Natural His- tory, New York Charles Hogue, Los Angeles County Museum of Natural History, Los Angeles, California D. S. (Woody) Horning, Jr., The Macleay Museum, Syd- ney, Australia Henry F. Howden, Carleton University, Ottawa, Canada 92 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus James H. Hunter III, Texas A&M University, College Station, Texas C. D. Johnson, Northern Arizona University, Flagstaff, Arizona David H. Kavanaugh, California Academy of Sciences, San Francisco, California Bert Kohlmann, Coacalco, Mexico Michael Kosztarab, Virginia Polytechnic Institute, Blacks- burg, Virginia R. Krause, Staatsliches Museum fur Tierkunde, Dresden W. L. Krinsky, Peabody Museum, Yale University, New Haven, Connecticut Robert J. Lavigne, University of Wyoming, Laramie, Wy- oming Robert E. Lewis, Iowa State University, Ames, Iowa Per Lindskog, Naturhistoriska Riksmuseet, Stockholm Ivan Lobl, Museum d’Histoire Naturelle, Geneva Ole Martin, Zoologisk Museum, Copenhagen Antonio Martinez, Rosario de Lerma, Salta, Argentina Scott McCleve, Douglas, Arizona Jean McNamara, Biosystematics Research Institute, Ot- tawa, Canada Miguel Angel Moron R., Instituto de Ecologia, Xalapa, Veracruz, Mexico Arturs Neboiss, National Museum of Victoria, Sydney, Australia Alfred F. Newton, Jr., Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts Mark F. O’Brien, University of Michigan, Ann Arbor, Michigan W. L. Overal, Museu Paraense Emilio Goeldi, Belem, Brazil L. L. Pechuman, Cornell University, Ithaca, New York Gary L. Peters, Oregon State University, Corvallis, Oregon Arwin Provonsha, Purdue University, West Lafayette, In- diana Brett C. Ratcliffe, University of Nebraska State Museum, Lincoln, Nebraska Pedro Reyes C., Instituto de Ecologia, Xalapa, Veracruz, Mexico E. G. Riley, University of Missouri, Columbia, Missouri Gerhard Scherer, Zoologische Staatssammlung, Munich Danny Shpeley, University of Alberta, Edmondton, Can- ada Cecil L. Smith, University of Georgia, Athens, Georgia R. zur Strassen, Forschungsinstitut Senckenberg, Frank- furt a. M. Terry W. Taylor, Ft. Davis, Texas Charles A. Triplehorn, Ohio State University, Columbus, Ohio Robert H. Turnbow, Texas A&M University, College Station, Texas Manfred Uhlig, Museum fur Naturkunde, Berlin Kenneth Vick, USDA, Gainesville, Florida William Warner, Chandler, Arizona Larry E. Watrous, Field Museum of Natural History, Chi- cago, Illinois Richard L. Westcott, Salem, Oregon Robert Woodruff, Bureau of Entomology, Gainesville, Florida Richard S. 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Essai preliminaire sur revolution des armures genitales des Scarabaeinae, par rapport a la taxonomie du groupe et a revolution du comporte- ment de nidification (Col. Scarabaeidae). Bulletin de la Societe Entomologique de France 88:532-542. Received 16 October 1991; accepted 15 June 1992. 98 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus APPENDIX: DISTRIBUTION DATA The appendix comprises geographical and temporal label data from specimens examined during this study. No data are included solely from published sources. They are ar- ranged by species and subspecies, which are listed alpha- betically. For each taxon, information is listed alphabet- ically by country and, for each country, alphabetically by first-order political subdivision (state, department, or province; political subdivisions for Belize, Guyana, Suri- name, French Guiana, and Trinidad and Tobago are not employed). Map points within political subdivisions are listed randomly, and each is followed, as available, by elevation and by month(s) of collection in roman nu- merals. For the United States, country and parish records for which there are no known map points appear par- enthetically at the end of the state listing. Relevant ex- planatory information appears parenthetically where nec- essary. Phanaeus achilles Boheman ECUADOR: Guayas — 40 km SW Guayaquil, 50 m, II; 45 km W Guayaquil; Guayaquil I; Bucay; Posorja, 0 m; Loja — Catamayo, X; Loja, IX, X. PERU: Lambayeque — Olmos, III; Tumbes — Cazaderos. Phanaeus adonis Harold MEXICO: Guanajuato — -Guanajuato, VIII; Hidalgo — 8 mi NE Jacala, 5200 ft, VI— VII; Agua Fria, X; Durango, IX; Fiuasca, IX; Barranca de Meztitlan, VI; 3 mi E Tepeji del Rio, VII; 4 mi S Tepeji del Rio; Ajacuba, VI; Puerto La Zorra, X; La Placita, VI; Mexico — Ixtapantongo, VIII; Ciudad Satelite, VII; Santa Cruz Acatlan, VIII; Nuevo Leon — • 7 mi W El Cercado, 2800 ft, VI; Villa Santiago, 1500 ft, VI; Chipinque Mesa (near Monterrey) 5400 ft, VII; Monterrey, VI; Queretaro — 3 mi S Queretaro, 6200 ft; San Luis Potosi— Microwave Station “Tortugas,” km 82 on Hwy 70, VIII; Valles, 300 ft, VI; Tamaulipas — 9 mi N Ciudad Victoria, VI; Gomez Farias, VI; 4 mi W Antiguo Morelos, 1200 ft, VI; 5 mi NE Llera, VII; Mi- quihauana, VIII; Canon de Caballeros, VI. Phanaeus alvarengai Arnaud No precise locality data available; see systematic section of text. Phanaeus amethystinus amethystinus Harold MEXICO: Chiapas — San Cristobal de las Casas, 2150 m, V-IX; 16 km W San Cristobal de las Casas, VII; Ocosingo, 905 m; 20 mi W Bochil, 5600 ft, VIII; Lagunas de Mon- tebello, V; jitotol, 6700 ft, VI; 3 mi W Jitotol, VI; Santa Rosa, V, VIII— IX; NW slope of Cerro Baul, 21 km W of Rizo de Oro, 1770 m, X; 11 km N Tzontehuitz, 1980 m, X; Hidalgo — 3 mi N Tlanchinol, 5100 ft, VII; 4 mi SW Chapalhuacan, 3500 ft, VII; 10 mi NE Jacala, VIII; Laguna Atezca, IX; Zacualtipan, VIII; 3 km S Zacualtipan, 2060 m, VII; Oaxaca— -Juquila Mixes, VI; San Luis Potosi— 7 mi W El Naranjo, 2400 ft, VI; 3 mi W Xilitla, 4800 ft, VI; Ciudad de Maiz, VII; Tamaulipas — Gomez Farias (Rancho “El Cielo”), 1100 m, IV; Veracruz — 15 mi NW Jalapa, VII-VII; Citlaltepetl, 7500 ft, VI; Las Vigas; Pre- sidio, VII; Huayacocotla, 2400 m, V; Teapan, 1770 m; Acajete, 2000 m. Phanaeus amethystinus guatemalensis Harold GUATEMALA: Cbimaltenango — Chimaltenango, 1750 m, VII-IX; Zaragoza, 2000 m, VIII-X; Quetzaltenango — San Juan Ostuncalco, 2400 m, V; Zunil, 2000 m, VII; Santa Maria de Jesus, 1550 m, VI; Quiche — Chupol, 2300 m, X; Chichicastenango, 6000 ft, VIII; Chuvexa, 2300 m, VII; Sacatepequez — Antigua Guatemala, VI; San Mar- cos— San Marcos, 2500 m, V; Solola — 12 km SSE Na- huala, 2275 m, VIII; Santa Lucia Utatlan, 2450 m, V; Chuchexik, 2200 m, X; Solola, VII; Xajaxac, 2325 m, V. Phanaeus amithaon Harold MEXICO: Aguascalientes — Aguascalientes; Colima — Manzanillo, XII; Colima, VI; Jalisco state line, Hwy 110; Guanajuato — Irapuato, VI, XI; 10 mi W Irapuato, VII; Valle de Santiago, VII; Dolores Hidalgo, VIII, IX; 22 mi E Penjamo, 5200 ft, IX; 6 mi W Penjamo, VIII; Celaya, VII; Hidalgo — 15 mi SW Huichapan, IX; Actopan, VIII; Jalisco — Cuzalapa, 660 m, VIII; El Tigre, 700 m, VII; Plantanarillo, 620 m, X; 2 mi S Tamazula de Gordiano, 3800 ft, XI; 22 km NE Zapotlanejo, VII; 3.4 km SE Jocotepec, 1493 m, IX; 36 mi N Cuidad Guzman, IX; 18 mi E Cuidad Guzman, VI; 10 mi E Union de Tula, IX; 15 mi SE Tequila, VIII; Lagos de Moreno, VI; 6 mi E Lagos de Moreno, VII; Etzatlan, VIII; Cocula, VII, XI; 13 mi SW Cocula, 5600 ft, IX; Chapala, VIII— IX; 6 mi W Chapala, VI; Santa Cruz del Astillero, 1417 m, IX; 22 mi NW Mascota, IX; Guadalajara, VI-VIII; 40 mi NW Gua- dalajara, VI; 11 mi W Guadalajara, IX; Magdalena, 4600 ft, VI-VIII; Ajijic, 1550 m, VII-VIII; La Barca, VII; 3 mi SW Autlan, VII; 17 mi NE Tecolotlan, VII; Juan Acatlan, IX; Micboacan — Maratavio, VIII; Los Reyes, VII; Eron- garicuaro, VI; 6 mi W Sahuayo, IX; Carapan, VII, IX; Morelia, IX; 4 mi E Morelia, VII; 33 mi E Morelia, IX; 17.4 km E Morelia, 2134 m, IX; 8 km E Zacapu, VII; Cotija, IX, XII; 5 km W Quiroga, VII; 13 km Quiroga, 2134 m, X; Patzcuaro, 2150 m, VI-IX; 7.7 km NE Patz- cuaro, 2088 m, IX; Zamora, VI; Tuxpan, 1900 m, VI- VIII; 7 mi SE Tuxpan, IX; 1 mi N Irimbo, 8400 ft, VII; 18 mi W Sahuayo, 4700 ft, VI; Janitzio, VII; 4 mi W Jiquilpan, 6100 ft, VIII, IX; Uruapan; Nayarit — Tepic, VI, XI; 24 mi SE Tepic, 4000 ft, VI- VII; 9.4 km SE Tepic, 975 m, IX; 17 mi NW Tepic, XI; San Bias, IX; 25.7 km E San Bias, IX, XII; 3 mi NW Santa Maria del Oro, VI; 8 mi N Ahuacatlan, XI; Ixtlan del Rio, VI; 4 mi E Ixtlan del Rio, IX; 4 mi S Sinaloa state line, Hwy 200, XII; 40.2 km SE Penitas, 366 m, IX; Ojo de Aguila, VII; Las Varas, XII; Compostela, 3000 ft; VIII, X; Acaponeta, VIII; 26 mi E Acaponeta, IX; 34 mi S Acaponeta, VIII; Quereta- ro— Tequisquiapan, IX; Queretaro, VI, IX; 35 mi SE Que- retaro, VIII; Sinaloa — Escuinapa; Culiacan; 12 mi SE Villa Union, VI; 13 mi E Concordia, 800 ft, VIII; 5 mi N Mazatlan, VII; 3 km E Mazatlan, VIII; 8 mi SW Con- cordia, 300 ft, IX; 8 mi NE Concordia, VII; Los Mochis, 14 m, IX-XI; Sonora — Guirocoba, VIII; Alamos, VIII; Hermosillo, VII; 17 mi SW Moctezuma, 944 m, VII; Tlax- cala — Tlaxcala; UNITED STATES: Arizona — Tucson, X; Tumacacori National Monument, VIII-X; Bisbee, X; Phoenix, X; 6.4 mi N Amado (“Canoa Ranch’’), VIII; 11 mi S Green Valley, VIII; Continental, VIII. Phanaeus{ ?) antiquus Horn UNITED STATES: Pennsylvania — Port Kennedy Cave. Phanaeus beltianus Bates COSTA RICA: Puntarenas — Osa Peninsula (“La Selva’’ Biological Station); San Jose — 16 km SW Sabanilla, V; NICARAGUA: “Chontales.” Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 99 Phanaeus bispinus Bates BOLIVIA: Cochabamba — Puerto Chimore, 25 m, I; Santa Cruz — Buena Vista; BRAZIL: Amazonas — Manaus, X; Para - — Belem; Cachimbo, VIII, X; COLOMBIA: Ama- zonas— Leticia, 700 ft, II; ECUADOR: Napo — Tena, 400 m, X; Pastaza— Canelos, X; FRENCH GUIANA: Beli- zon, V; Cayenne, VII; GUAYANA: Kartabo, VI; Mora- balli Creek, X; PERU: Huanuco— Tingo Maria, 700 m, VII, IX-X; Junin — Satipo, IV; Sani Beni, IX; Loreto — Ucayali, I; Panguana, 260 m, III; Pucallpa, 200 m, VI; TRINIDAD: Maracas Bay, II, VI, VIII; VENEZUELA: Amazonas — San Carlos de Rio Negro, 65 m, III. juapan de Leon, IX; Puebla — 13 mi SE Acatlan, VIII; 9 mi SE Izucar de Matamoros, 4800 ft, VIII; 12 mi SE Izucar de Matamoros, 3700 ft, VII; 30 mi S Atlixco, VIII; 7 mi NW Tehuitzingo, VII; 13.3 mi NE Tehuitzingo, VII; 9 mi N Amatitlan, XII. Phanaeus dejeani Harold BRAZIL: Espirito Santo— Fazenda Jerusalem, I; Minas Gerais— Serra Cara^a, 1380 m, XI; Fazenda dos Campos, Virginia, 1500 m, I; Rio de Janeiro — Petropolis, I— II, X- XII; Teresopolis; Itatiaya, 1100 m, XI; Nova Friburgo; Sao Paulo— Campos do Jordao, II, XII. Phanaeus cambeforti Arnaud BRAZIL: Amazonas — 15 km E Manaus, VII; Tefe, I, III; COLOMBIA: Amazonas— Leticia, 700 ft, II— III; Meta — Villavicencio, II— III; FRENCH GUIANA: Bas Moroni; PERU: Huanuco — Tingo Maria, XI— XII; Loreto — Ata- laya, II; Boqueron Abad, 550 m, IX. Phanaeus chalcomelas (Perty) BOLIVIA: Cochabamba — Puerto Chimore, 250 m, I; Rio Coni, 400 m; La Paz — Tumupasa, IX; Ixiamas, XII; Santa Cruz — Buena Vista, 1700 ft, I— IV; El Espejo, II; BRAZIL: Amazonas — Tefe; Benjamin Constant; 15 km E Manaus, VII; Para ■ — Cachimbo, X; Obidos, XII \Rond6nia — Porto Velho; COLOMBIA: Amazonas— Leticia, II; Huila — Gi- gante, III-IV; Vegas de Caqueta, I; Meta — Villavicencio, 500 m; 33 km E Villavicencio, III; Putumayo — Santa Rosa; Mocoa, 530 m, II; ECUADOR: Morona Santiago — Ma- cuma, VII; Napo — Lago Agrio, 250 m, VI; Limoncocha, 250 m, VI— VII; FRENCH GUIANA: Cayenne; St. Jean du Maroni; St. Laurent du Maroni; Massikiri (Oyapock River), XI; PERU: Cuzco — Marcapata; Huanuco — Tingo Maria, XI— XII; Loreto — Pucallpa, V; Boqueron Abad, VII; SURINAME: Anapaike, XL Phanaeus damocles Harold MEXICO: Guerrero — Omilteme, 8000 ft, VII, IX; Amula, 6000 ft, VIII; Oaxaca — -Municipio Suchixtepec, Rio Gua- jolote, 1975 m, VII; 4 km W Calpulalpan, 2000 m, VI; 5 mi N Oaxaca, 1700 m, VI; km 154 on Hwy 175 S Oaxaca, VI; 36 km NE Oaxaca (El Cerezal), 2300 m, VI; 3 mi N Suchixtepec, 9500 ft (km 144 on Hwy 175 S Oaxaca), VI; Nochistlan, VI; Monte Alban, VIII. Phanaeus daphnis Harold MEXICO: Guerrero — Taxco, IX; Coacoyula, VI; Telo- loapan; Iguala, VI; 13 km NNW Iguala, 1035 m, VIII; 4 mi S Chilpancingo, VIII; Agua Bendita, VII; Tierra Co- lorada; Mexcala, 4500 ft, VII; 1.5 mi N El Mogote, 1500 m, VIII; Amula, 6000 ft, VIII; Hueyecantenango (Chilapa), VIII; Mexico — Temascaltepec, 6000 ft, VII; Tejupilco, VII; Tonatico, VII, IX; Nuevo Santo Tomas de los Pla- tanos, IX; Ixtapan de la Sal, VII; Chalma, 2600 ft, VI; Michoacdn — Tuxpan, 1450 m, VII-IX; La Huacana, VII; 19 mi S Uruapan, 3000 ft, VII; 2.5 mi N Huetamo de Nunez, 3100 ft, VII; Apatzingan, 1200 ft, VIII; Morelos — Puente de Ixtla, VII; Progreso, VI; Tepoztlan, 5000 ft, VI— IX; Cuernavaca, 1650 m, V-IX; 2 mi SE Cuatla, 1372 m, IX; 11.5 mi W Cuatla, 4500 ft, VIII; 3 mi W Tlayecac, VI; San Vicente, VI; 3 mi W Moyotepec, VI; 17 mi NE Amacuzac, VI; Oaxtepec, VI— VII; Xochitepec, VI; Cau- tela, VII; 7 km S Alpuyeca, 3200 ft, VI— VII; Xochicalco, 4000 ft, VII; Tequesquitengo, VI; Oaxaca — 8 mi N Hua- Phanaeus demon Laporte-Castelnau COSTA RICA: Guanacaste — 6 km S La Cruz, VI; Las Canas; 5 km NW Las Canas, VII; 25 km N Las Canas, V; 15 km SW Bagaces; Palmira, V; Santa Rosa National Park, VI; San Jose — San Jose; EL SALVADOR: La Liber- tad— 6 mi W Quetzaltepeque, 500 m, VIII; La Union — Volcan Conchagua, V; Playa El Icacal, VII— VIII; Santa Ana — San Diego, VI; Usulutan — Usulutan, VII; 15 km E Usulutan, VIII; GUATEMALA: Izabal — Los Amates; HONDURAS: Choluteca — Choluteca, IX; 2 mi E Cho- luteca, IX; MEXICO: Chiapas — 18 km S La Trinitaria, 914 m, XII; Tuxtla Gutierrez, VI, IX; 65 km S Tuxtla Gutierrez, 823 m, IX; Ocozocoautla, VI, IX; 2 mi N Oco- zocoautla, VIII; Santa Rosa, IX; 10 mi NW Arriaga, 122 m, VIII; 18 mi N Arriaga, VIII; Cintalapa, VIII; Las De- licias, X; Colima— Colima, VI; 7 mi W Colima, VIII; 20 mi S Colima, VI; 4 mi E Colima, 1500 ft, VIII; Guerrero — Colotlipa, VII; Chilpancingo, VIII— IX; 22 mi N Chilpan- cingo, VIII; 24 mi N Chilpancingo (Canon del Zopilote), VII; 5 mi S Chilpancingo, VIII; 30 mi S Chilpancingo, 4000 ft, VIII; Iguala, V-VII; 8 mi SW Iguala, VII; 3 mi S Iguala, VIII; 13 mi NNW Iguala, 1035 m, VIII; Taxco, VII; 20 mi E Taxco, IX; Mexcala, VII; 2 mi N Mexcala, VIII; Zumpango, VII; Huitzuco, XI; Coacoyula, IX; 75 km E Acapulco, X\ Jalisco— Autlan, IX; 3 mi SW Autlan, VII; Barra de Navidad, VI; Juchitlan, IX; Manzanillo, VI; Chamela (Estacion Biologica UNAM), VII; Michoacdn — Tocambaro, 1300 m, VI; Playa Azul, 0 m, VII; 3 mi W Tangamandapio, 5250 ft, VI; Apatzingan, 1200 ft, VIII; La Huacana, VII; 10 mi S Tzitzio, VII; 5 mi SW Tiquicheo, VII; 20 mi E, 23 mi S Morelia, 2950 ft, VII; Morelos — 2 mi W Moyotepec, VI; Tepoztlan, IX; Tequesquitengo, VII; Palo Bolero, VIII; Zacatepec, VI; Cuautla, VIII— IX; 21 km SE Cuautla, 1372 m, IX; Huajintlan, IX; Tlalti- zapan, 1100 m, VIII; Xochitepec, VI; 15 km W Xochi- tepec, 1250 m, X; Villa de Ayala; Alpuyeca, VI— VII; Cuer- navaca, VI- VIII; 10 mi S Cuernavaca, VII; 16 mi S Cuernavaca, VIII; Puente de Ixtla, VI; Yautepec, VI; 7 mi SW Yautepec, 3500 ft, VII; Tetecala, VI; Tepalcingo, VIII; Alpuyeca, VII; 7 km S Alpuyeca, 3200 ft, VI— VII; Jojutla, V; Xochicalco, VIII; Nayarit — Campostela, IV; 6 mi SW Sayulita, VII; Oaxaca — 4 km E Tequisistlan, 700 ft, VIII; 6 mi N Juchitan, VII; 15 km E Juchitan, VII; 17 mi E La Ventosa, VII; 5 mi N La Ventosa, VII; Tehuantepec, XII; 8 mi NE Tehuantepec, VII; 14 mi W Tehuantepec; 6 mi S Tehuantepec, VII; Puerto Angel, 40 m, VI; 38 mi N Puerto Angel; 3 mi N Totolapan, 3400 ft, VI; Presa Benito Juarez, VII; 7 mi W Tehauntepec, 90 m, VII; Huajuapan de Leon, VI; 1.5 mi E Zopilote, VI; Puebla — 22 mi SE Izucar de Matamoros, VIII; 12 mi N Izucar de Mata- moros, 5000 ft, VI; 17 km S Izucar de Matamoros, 4200 ft, VIII; 13 km NE Tehuitzingo, VII; 6 mi NW Tehuit- zingo, 3600 ft, VII; 10 km SE Tehuitzingo, 1150 m, VII; Acatlan, VII; 13 mi SE Acatlan, VIII; 45 mi N Acatlan; 7 100 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus mi S Atlixco, 4900 ft, VII; NICARAGUA: Chinandega — 20 km NW Leon (Depto. Leon) Managua — Managua, VI; 3 mi SW Managua, VI. Phanaeus difformis Leconte MEXICO: Tamaulipas — Tampico; 16 mi SSW San Fer- nando, 100 m, VIII; UNITED STATES: Arkansas — 2 mi E Ozark, VIII; Pine Bluff; Colorado — Lamar, VIII; Kan- sas— Medora (Seward Co.; Clark Co.; Ford Co.; Coman- che Co.); Louisiana — Cameron, IV, VI; Johnsons Bayou, V (Cherokee Parish); New Mexico — Eunice, VII; Los Me- danos, VIII; Mescalero Sands, VII; Oklahoma — Little Sa- hara, VI; Grand, VIII; Red River at Interstate Hwy 35 (Love Co.; Cherokee Co.); Texas — Sarita, X; Kingsville, X; Lyford, V; Riviera, VIII; Rio Grande City, IX; Sinton, VI, VIII; Duval City, VI; Edinburg, VI; Raymondville, V- VI; Santa Ana National Wildlife Refuge, VII; Aransas National Wildlife Reserve, IV; 18 mi SE Laredo, IX; Padre Island National Seashore, IX; Dallas, VI; San Antonio, XII; 7 mi S Seagoville, VI; Pearsall, VI; Cypress Mills; Round Mountain; Galveston; Sabinal, XII; Fedor; Coyote Lake, VIII; Proctor, VI; New Braunfels; Brownsville; Ca- nadian; Brownwood, X; Gainesville; Columbus, V; Vic- toria; 24 mi N Falcon Dam; Port Isabel, IX; Corpus Chris- ti, VIII; San Diego, XI; Mustang Island; Hockley; McAllen, VI; College Station, IV; Waco, VII; 8 mi S Cuero, IX; 6 mi N Cuero, IX; 4 mi W Cookes Point, IX; Camp Creek Lake, IX; 4 mi S Buffalo, IX; 2 mi E Bastrop, V; Freer, IV; Sheffield, VII; Fredericksburg, X; 3 mi E Goliad, IX; 5 mi N Industry, V; Robstown, VI; Waskom, IV; Heb- bronville, VIII; Uvalde, X; Streeter, V; San Jacinto River at Interstate Hwy 45, Montgomery Co.; Winter Haven; 8 mi S Falfurrias, VII; Frelsburg, VI; Crane; Austwell; Higgins, VI; 8 mi E Tyler, IX; 10 mi SE Tyler, IX; Mule- shoe, VIII; Marfa (Karnes Co.; Anderson Co.; Dimit Co.; Grimes Co.). Phanaeus endymion Harold BELIZE: Caves Branch, VIII; Augustine, 1500 ft, VIII; EL SALVADOR: La Libertad — 4 km S Nueva San Salvador (= Santa Tecla), V; GUATEMALA: Alta Verapaz — Lan- quin, 350 m, VIII; Coban; 35 km SW Sebol, 915 m, VII; Escuintla — 10 km NE Escuintla, 700 m, VII; Huehuete- nango — 16 km NW Huehuetenango, VIII; Izabal — Ca- yuga, VIII; Peten — Tikal, VIII; Retalhuleu — San Sebas- tian, VIII; Santa Rosa — 6 km E Oratorio, 670 m, VII; HONDURAS: Atlantida — Tela, V; Francisco Mora- zdn — Cedros, XI; MEXICO: Chiapas — Palenque, V; 7 mi S Palenque, 350 m, VIII; 48 km NE Las Margaritas, 2100 m, X; 1 mi SW Rizo de Oro, 2700 ft, VIII; 6.6 mi W El Bosque, 4800 ft, VIII; 11 mi E La Trinitaria, 5200 ft, VIII; 2 km S La Trinitaria, X; 11 mi NW Ocozocoautla, 3400 ft, VIII; 15 mi NW Ocozocoautla, 2800 ft, VIII; 22 mi N Bochil, 5600 ft, VIII; Santa Rosa, VIII; Simojovel, V; Bonampak, 300 m, V, VIII; 9 mi N Arriaga, VIII; Lagos de Montebello, 4900 ft, VIII; Lacanja-Chansayab, 300 m, II, IV, VIII; 4.1 mi S Tuxtla Gutierrez (“Parque Cerro Hueco”), X; Rosario Izapa, V; El Escopetazo (km 40 on hwy between Tuxtla Gutierrez and San Cristobal de las Casas), VIII; Jalisco— 13 mi SW Cocula, 5600 ft, IX; Te- calitlan; 10 mi SW Autlan, 4200 ft, IX; Michoacan — 12 km NW Calcoman, VII; Nayarit — 24 mi SE Tepic, 4100 ft, VII; Oaxaca — 6 mi S Valle Nacional, 2000 ft, VII; 13 mi S Valle Nacional, 3700 ft, VIII; 15 mi S Valle Nacional, 5600 ft, VII; 9 mi E El Camaron, 4300 ft, IX; Juquila Mixes, VIII, X; Niltepec, IX; Loma Bonita, VIII; Nueva Esperanza, IX; Portillo del Rayo (Finca “Dos Angeles”), VII; Quintana Roo — X-Can, VI; Carrillo Puerto, XII; Ta- basco— 3 mi NE Teapa; 8.2 mi W Teapa; Veracruz — 22 km W Palma Sola, 800 m, VIII; 12 km N Sontecomapan, 125 m, VII; 10 km NE Catemaco, 500 m, VII; Dos Amates, V; Cordova, VII; Huatusco, VIII; Tiro de Hayas (Ejido “El Sumidero”), 1360 m, VIII— IX; Fortin de las Flores, VII; Yucatan — 3 mi N Muna, 100 ft. Phanaeus eximius Bates COSTA RICA: Cart ago — Turrialba, VII; Guanacaste — 6 km S La Cruz, VI; 25 km N Las Canas, V-VI; EL SALVADOR: Chalatenango — Chalatenango, VII; La Libertad — 2 mi E Quetzaltepeque, VII; 6 mi W Quetzalte- peque; Santa Ana — Santa Ana, VI; GUATEMALA: Ju- tiapa — 10 km W Jalpatagua, 600 m, VII; Zacapa — La Union, X; HONDURAS: Cortes — San Pedro Sula; La Lima, 2000 ft, V; Francisco Morazdn — 25 km E Tegucigalpa; NICARAGUA: Managua — Managua. Phanaeus flohri Nevinson MEXICO: Chiapas — Tapachula*; Guerrero — Acahuizo- tla, 650 m, VIII, X; 9 km S Palo Blanco (Sierra Alquitlan), 1400 m, VII; Jalisco — Ajijic; 5.5 mi NE Autlan; More- los— San Jose de los Laureles, V-VI; Santo Domingo, VI; Puebla — 7 mi S Izucar de Matamoros; Sonora — 10 mi NW Yecora, 4320 ft, VII; 20.1 mi E Rio Yaqui, 3010 ft, VII; 3.2 mi NW Huicache, 5170 ft, VII; Veracruz — Ja- lapa*; Las Vigas*. (*Localities are not confirmed by mod- ern records.) Phanaeus furiosus Bates MEXICO: Guanajuato — Irapuato; Valle de Santiago, VI; Silao, VII; Jalisco — Guadalajara, VIII; 30 mi S Guadala- jara; 5 mi S Guadalajara, VI; 28 mi E Guadalajara; 10.8 mi S Talpa de Allende, 1900 ft; Etzatlan, VIII; Ajijic, 1500 m, VI— VIII; Magdalena, VIII; 22 km NE Zapotlanejo, VII; 22 mi NW Mascota, IX; 13 mi SW Cocula, IX; Cha- pala; Tala, 5200 ft, VIII; Tequila, VII; La Venta; Tecolo- tlan, VII; Michoacan — Patzcuaro, VII— VIII; Cojumatlan; 6 mi W Sahuayo, IX; 18 mi W Sahuayo, VI; Cotija, IX; Jacona, VIII; Coalcoman, 4000 ft, VII; Nayarit — San Bias, VI; 9 mi E San Bias, VIII; Jesus Maria, VII; Compostela, 2500 ft, IX-X; 17 mi NW Tepic, XI; 34 mi S Acaponeta, IX; 4 mi E Ixtlan del Rio, IX; San Juan Peyotan, VIII; Sinaloa — 28 mi NE Concordia, VIII; 2 mi E Concordia, VI; 13 mi E Concordia, VIII; Santa Lucia, 4000 ft, VIII; Escuinapa; 27 mi E Villa Union, VII; Copala, VII— VIII; 7 mi S Culiacan; La Concha, VIII; 1.5 mi SW Panuco, VII; Mazatlan, XI; Sonora — Rio Mayo, VIII; 55 km SW Moc- tezuma, 1066 m, VII; 17 km SW Moctezuma, 944 m, VII. Phanaeus halffterorum Edmonds MEXICO: Guerrero — Acahuizotla, 750 m, X-XI; 9 km S Palo Blanco (Sierra de Alquitlan), 1450 m, VI-VII; 22 mi S Chilpancingo, 2800 ft, VIII; Mexico — 8 km W Temas- caltepec, 2360 m, VII; 5 km E Temascaltepec (“Real de Arriba”), 2200 m, VII. Phanaeus haroldi Kirsch COLOMBIA: Caqueta — Vegas del Caqueta, 1; Fluila — Gigante, III— IV; Campoalegre, V; Meta — Villavicencio, VIII-X; ECUADOR: Napo — Lago Agrio, 250 m, VI; Lo- reto, XII; Pastaza — Puyo, II; Canelos, XII; Tena, 400 m, II; PERU: Huanuco — Tingo Maria, IV, VI; San Martin — Moyobamba, VIII; San Martin, 1500 ft, XII; VENEZUE- Contributions in Science, Number 443 Edmonds: Revision of Phanaeus M 101 LA: Amazonas — San Fernando de Atabapo, VI; Boli- var— El Playon (Rio Cuara), XI; Maripa, VI; Tachira — Rio Negro, 500 m, VIII. Phanaeus hermes Harold COLOMBIA: Magdalena — Aracataea, VIII; Rio Frio, 2000 ft, VI; Sierra de Perija, 650 m, VII; Norte de Santander — La Playa, VI; Ocana, VI; Santander — Bucaramanga, V; COSTA RICA: Puntarenas — 11.6 mi S Buenos Aires, VI; 15 km NE Potrero Grande, IX; San Jose — San Isidro del General, V; PANAMA: Chiriqui — David; Tole; Pama- ma — Cerro Campana, 850 m, V; La Chorrera, V; Chepo, XII; Las Cumbres, XII; VENEZUELA: Zulia — km 40 on hwy between Machiques and Colon (Hacienda “San Ma- rino”); Kasmera (Sierra de Perija), 250 m, IX. Phanaeus howdeni Arnaud PANAMA: Canal Zone — Barro Colorado Island, V-XII; Lion Hill Island; Panama — Maje (8 km SE Bayano Ridge). Phanaeus igneus Macleay UNITED STATES: Alabama— Mobile, III; Springhill, V; Hartford, III; Claireborne (Baldwin Co.); Florida — Mi- ami, IV— VII; Gainesville, VI-X; Jacksonville, V, XI; Gulf- port, III; Lake Worth; Ft. Myers, VI; Fruitland Park; Tar- pon, VI; 3.5 mi N Salt Springs, IV; Koreshan State Park, VI; Mariana, IV; De Funiak Springs, III; 12 mi S Chatta- hoochee, VII; Brooksville, VI; Enterprise; Clarksville, III; 1.4 mi SW Archer, III; Ocala National Forest, III; 14.3 mi E Ocala; Largo, VII; Sarasota; Inverness, VII; Lake Placid, III; 8 mi SE Interlachen, III; Daytona, IV; Edge- water; Crescent City; Winterpark, III; Ormond, III; New- man’s Lake, XII; Walaca, IV; La Belle, VII; Kissimmee, V; Homosa Springs, IV; Sanford, X; Tampa, III; 3.7 mi N Old Town, V; Lake Alfred; 6 mi E Quincy, III; North Smyrna, XII; Clearwater, VI; 8 mi E Bronson, III; Talla- hasee, IV; Pensacola; Lutz; St. Petersburg, V; Torreya State Park, IV; Glen St. Mary, XII; Plant City; Romeo, IV; Georgia — Jekyll Island; Tifton, V; Vidalia; 10 mi SE Waycross, VI; Thomasville, III— IV; Darien, IV; Lumber City, IX; St. Marys; Little Ocmulgee State Park, IX; St. Catherines Island, VI; Brunswick, VII; Augusta; Swains- boro, VIII; Blackbeard Island, V-VII (Camden Co.; Glynn Co.; Echols Co.); Louisiana — Covington, V; Mississip- pi— Hattiesburg, VIII; Ocean Springs, VI; Lucedale, VII; North Carolina — Southern Pines, II— X; West End, VII; New Bern; Southport, X; Wilmington, V; Faison; South Carolina — Myrtle Beach, VII; Aiken; Beaufort, VI; Flor- ence, IV-VI; Hilton Head Island, VII; Walterboro, IX; Blackville, VIII; Isle of Palms, VII; White Pond, III; Mt. Pleasant, VIII; Summerton, VII; Dillon, X; Darlington, VIII; Barnwell, VIII; Bishopville, IX; Charleston, IV. Phanaeus kirbyi Vigors BOLIVIA: Santa Cruz — Buena Vista, 1700 ft, I— IV, XI- XII; Chiquitos, XI; BRAZIL: Goias — 20 km N Sao Joao da Alianga, V; 40 km S Peixe, VI; Mato Grosso — Cha- pada, 2600 ft, II, IX-XI; Tres Lagoas, VI; Rio Verde, XI; Corumba; Uitiariti (Rio Papagaio), XI; Alto Gargas, XII; Murtinho, XI; Minas Gerais — Sertao de Diamantina, XI; Parana — Villa Velha, II, XI; PARAGUAY: Alto Para- na— Belle Vista, I; Boqueron — Puerto Casado; Caagua- zu — Caaguazu, XII. Phanaeus labreae (Pierce) UNITED STATES: California — Los Angeles (Rancho La Brea Tar Pits). Phanaeus lunaris Taschenberg ECUADOR: Azuay — Huigra, 1300 m; Bolwar — Balza- pamba; El Oro — Zaruma, II; Esmeraldas — San Mateo, IX; Telimbelo; Guayas — Guayaquil; Los Rios — Queve- do, 75 m, I; Babahoyo, I; 45 km N Babahoyo, 700 ft, II; Loja — -Loja, XII; Cariamanga, VII. Phanaeus melampus Harold MEXICO: Chiapas— 6.6 mi W El Bosque, 4800 ft, VIII; Mahosik’ Tenejapa, 4800 ft, VII; Veracruz — Barranca de Metlac, VII; Orizaba, I; Fortin de las Flores, VIII. Phanaeus meleagris Blanchard BOLIVIA: Cochabamba — Yungas del Palmar, 1600 m, II, IX; Crista Mayu, 600 m, IX-XI; La Paz — Coroico; Calisaya, V; Chuani; Puente Llorosa, I; Santa Cruz — Buena Vista, 450 m; Parapet! River Valley, near Lagunillas; COLOMBIA: Boyaca — Muzo; Meta — Villavicencio, V; Tolima — Honda; ECUADOR: Pastaza — Mera; Canelos; San Francisco del Rio Pastaza, 1200 m, X; Zamora Chin- chipe — Sabanilla, IX; PERU: Cuzco — Marcapata; Hua- nuco — Pozuzo; Tingo Maria, VII; Junin — La Merced; Loreto - — Pucallpa, 200 m, XI; Pasco — Chuchurras; VEN- EZUELA: Tachira— San Cristobal, 1200 m, V, VIII-IX. Phanaeus melibaeus Blanchard BRAZIL: Goias — Aragar^as, I, X; Mato Grosso — Rosa- rio Oeste, II; confluence of Tapirape and Araguaia Rivers, XI-XI; Para — 45 km E Caninde, X; Rondonia — Porto Velho, X. Phanaeus mexicanus Harold MEXICO: Guerrero — Acahuizotla, 750 m, XI; 11 mi E El Ocotito, 3200 ft, VIII; Tierra Colorada, VII; Chilapa, VIII; Mexico — Temascaltepec, VIII; 5 km E Temascal- tepec (Real de Arriba); Tejupilco; Ixtapan de la Sal, 1975 m, VII; Valle de Bravo, VIII; Tenancingo, V, IX; Villa Guerrero, VII; Malinalco, IX; Oaxtepec, X; Morelos — Cuernavaca, 6500 ft, VI- VIII, XI; Cuautla, VIII-IX; Jo- jutla, VI; Puente de Ixtla, VII; Yautepec, VII; Tepoztlan, VI— VIII; Atonyo, 1630 m, VI; Oaxaca — Oaxaca, 1600 m, V-IX; 4 mi NW Oaxaca, IX; Juquila Mixes, IX; Puebla — 6 mi E Atlixco, IX; 7 mi S Izucar de Matamoros, 4500 ft, VI; Acatlan, VII; Veracruz — Salinas, VI; Presidio, VII; Catemaco (Dos Amates), I, IV— XII; Fortin de las Flores, VII; San Andres Tuxtla, VII— VIII; Orizaba, IX; Tezonapa, IX; Cotaxtla, VIII; Cerro Azul, XII. Phanaeus nimrod Harold MEXICO: Oaxaca — 2.7 mi NW El Camaron, VII; 9 mi E El Camaron, IX; Ocotlan; 20 mi S Ocotlan; Oaxaca, 5050 ft, V, VIII; 15 mi S Oaxaca, VIII; 4 mi NW Oaxaca, IX; Monte Alban, VII-VIII; 48 mi W Tehuantepec, VIII; 1.5 mi E El Zopilote, VI; Puerto Angel, 40 m, VI. 102 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus Phanaeus palaeno Laporte-Castelnau ARGENTINA: Corrientes — Villa Olivari, XII; BOLIVIA: Beni — Magdalena, I— III; Santa Cruz — Chiquitos, XI; BRAZIL: Goias — Jatai (Fazenda Nova Orlandia), I; Goia- tuba, I; Mato Grosso — Tres Lagoas, V-VI; Rosario Oeste, XI— XII; Santa Teresinha, XI; Utiariti, X; Murtinho, XII; Rio Verde, XI; Corumba; Chapada (near Cuiaba), IV— VII, X-XI; Aragar^as; Rancho Grande, XI; Nobres, I— III; Ron- donia — Vilhena, II; Santa Catarina — Joinville; Sao Pau- lo— Campinas, II, XII; Batatais; Agudos; Epitacio, XII; Boa Esperan^a do Sul, I; PARAGUAY: Boqueron — Puer- to Casado; Concepcion — Horqueta, III; Guaira — Villari- ca; Borja, II; Presidente Hayes — Asuncion. Phanaeus palliatus Sturm MEXICO: Distrito Federal — Atzcapotzalco, VII; Con- treras, 2700 m, VII; Durango — 145 km S Durango (Sierra de Michis, “La Michilia” Biosphere Reserve), 2300 m, VIII; Hidalgo — 4 mi W Tulancingo, 7600 ft, VII; Huau- chinango, VIII; 12 mi W Huauchinango, 6700 ft, V\; Jalis- co— Sierra Manantlan (“Las Joyas” Biological Station), 1900 m, VII; Ajijic, 1950 m, VIII; 3 mi WSW Mazamitla, VII; 8 mi S Autlan, VII; Mexico — Tepotzotlan, VI; Te- mascaltepec; Amecameca, VI; Toluca, 2640 m, VI— VIII; Almoloya, VII— VIII; Ocoyacac, VII— VIII; Villa del Car- bon, VI; Tejupilca; San Bartolo Tenayuca, VI; Chapingo, 2250 m, VIII; Morelos — Cuernavaca, 1500 m, Michoa- can — 5 mi E Quiroga, 7600 ft, VII; Tancitaro, 6300 ft, VI; Matujeo, VII; Carapan, 6200 ft, VI; Patzcuaro, 2150 m, VI— VII; 7.7 km NE Patzcuaro, 2080 m, IX; Puerto El Tigre, VII; Paricutin, VII. Phanaeus prasinus Harold COLOMBIA: Norte de Santander — 3 km N Chinacota, 3000 ft, V; 30 km S Cucuta, 700 m, V; TRINIDAD AND TOBAGO: Port of Spain; Maracas Bay, VI— VIII; Narva; Balandra Bay, X; Arima Valley, V; Diego Martin Valley, VIII; VENEZUELA: Apure — San Fernando, X; Aragua — Rancho Grande (Maracay), VII; Pozo del Diablo (Mara- cay), 500 m, IV; Limon, 450 m, IX; Barinas — Las Maravi- llas, IV; Barinitas, 500 m, IV-VI; Bolivar — 48 km ESE Cuidad Bolivar (Rancho “Santa Rita”), VII; Carabobo — San Esteban, 100 m, II, VII; Valencia, 500 m, IV; Distrito Federal — Chichiriviche, 0 m, X; Caracas, VI, XI; Falcon — Yaracal (Hato “Corralito”), 60 m, I; Gudrico — Las Mer- cedes (Hato “El Saman”), V; Merida — Merida; La Pedre- goza, 2000 m; Tabay, IV; Miranda — Birongo, I; Parque Nacional Guatopo, III; Monagas — Caripito, VIII; Yara- cuy — Yumare, VI. Phanaeus pyrois Bates COLOMBIA: Antioquia — Puerto Berrio, VIII; Boyacd — Muzo, IX; Choco — Quibdo, IX; Valle del Cauca — 70 km E Buenaventura (Anchicaya Dam), 500 m, V, VIII; COSTA RICA: Cart ago — Turrialba, 600 m, II; Heredia — Puerto Viejo, 90 m, IX; Limon — Llanuras de Santa Clara, VI; Puntarenas — 6 km S San Vito, 1000 m, V, VIII; Rincon, 100 m, IX; ECUADOR: Esmeraldas — 11 km SE San Lo- renzo, VI; Los Rios — Quevedo, 45 m, V, VIII; Manabi — 78 km NE Chone, 450 m, VI; Pichincha — 4 km SE Santo Domingo de los Colorados, 500 m; PANAMA: Canal Zone — Barro Colorado Island, I— II, VII— XII; Chiriqui — 2 mi N Santa Clara (“Hartmann’s Finca”), 1200 m, VI- VO; Panama — Cerro Campana, 900 m, V. Phanaeus quadridens (Say) MEXICO: Chihuahua — Matachic, VII; 27 mi W Parrita, VII; Madera, 7200 ft, VII; 9.7 mi W Cuauhtemoc, VII; Cusihuiriachic; 13.2 mi S Villa Matamoros, 6000 ft, VIII; 8 mi W Matachic, 7200 ft, VII; San Francisco del Oro Mesa, 7300 ft, IX; Charcos, 6000 ft, VII; Santa Barbara, 7500 ft, VII; 63 mi W Santa Barbara, 5500 ft, VII; 26 mi S Hidalgo del Parral, VIII; Santa Clara; Mesa de Huracan, 7400 ft, VII; Valle de Olivos, 5500 ft, VII; 11 mi E Hue- jotitlan, 5900 ft, VII; 6 mi N-3 mi E El Sueco, X; Distrito Federal — Mexico City, VII; Villa Madero, VII; Guada- lupe, IX; Durango — 3 mi E El Salto, 2400 m, VIII; 6 mi NE El Salto, 8500 ft, VIII; 45 mi E El Salto, VIII; Otinapa, 8200 ft, VIII; Durango, VIII; 5 mi W Durango, 6500 ft, VI; 30 mi W Durango, VI; 43 mi W Durango, 1900 m, VII— VIII; 145 km S Durango (Sierra de Michis, “La Michi- lia” Biosphere Reserve), 2300 m, VI— VIII; 32 mi N Du- rango, VII; 7 mi SW Buenos Aires, 8800 ft, VIII; Gua- najuato— Yuriria, VII; Leon, VI; Irapuato, VII; Hidalgo — 12 mi W Huauchinango, Puebla, 6700 ft, VI; Tulancingo, 6750 ft, VI-VII; 4 mi E Tulancingo, VIII; Pachuca, VIII; 15 mi SW Huichapan, 7800 ft, IX; Huasca, IX; Jalisco — 3 mi W Magdalena, VI; Zacualtipan, VII— IX; Mexico — Toluca, VI; Temascaltepec, VII; 25 mi E Mexico City, VII, XI; Michoacdn — 3 km E Zacapu, VII; 7.7 km NE Patzcuaro, 2100 m, IX; Morelia, VIII; 33 mi E Morelia, IX; 5 km W Quiroga, VII; 6 mi W Sahuayo; Nayarit — Santa Teresa, 2070 m, X; Puebla — 2.5 mi S Huauchinan- go, VIII; 11 mi SW Puebla; Sinaloa — 6.5 mi E Potrerillos, VIII; El Palmito, VI; 8 mi W El Palmito, 6100 ft, VI; 38 mi NE Concordia, 6200 ft, VI; Puerto Loberas, 2000 m, VII; Sonora — Yecora, VI; Veracruz — Acutzingo, XI; 8 mi N Cuidad Mendoza, VII; Pie del Cumbres (km 295 on Hwy 140), 8000 ft, VI; Zacatecas — Sombrerete, VII; UNITED STATES: Arizona — Pinery Canyon (Chiricahua Mts.), VIII— IX; Portal, 4800 ft, IX; 15 mi W Portal, VIII; Patagonia, VII; 4.3 mi SW Patagonia, IX; Madeira Canyon (Santa Rita Mts.), IX; Miller Canyon (Huachuca Mts.), 5200 ft, IX; 28 mi SE Sonoita (Lyle Canyon), IX; Pena Blanca Canyon (Pajarito Mts.), VII— VIII; 65 mi N Willcox (Galuro Mts., Deer Creek Ranch), VII; Ft. Huachuca, VII; Duquense, IX; New Mexico — Tyrone, VIII; Animas Mts. (Birch Spring), IX. Phanaeus sallei Harold BELIZE: Indian Church, VII; GUATEMALA: Alta Vera- paz — 9 km E Panacajche; Zacapa— La Union, 850 m, IX-X; MEXICO: Chiapas — Boca del Chajul, X; Bonam- pak, 300 m, VII-IX; Santa Rosa, VIII; Lagunas de Mon- tebello, 4900 ft, VII; Palenque, VI-VII; Veracruz — Pre- sidio, VII; Naolinco, IIX; Fortin de las Flores, IX; Banderilla; Jalapa, X; Catemaco, 100 m, V; Cosautlan, 900 m. Phanaeus scutifer Bates MEXICO: Veracruz — Cotaxtla, VII; 14 mi W Conejos, VI; 24 mi N Huatusco, 650 m, VII; 14 mi SE Jalapa, VII; 7 km W Palma Sola, 100 m, VI-VII. Phanaeus splendidulus (Fabricius) ARGENTINA: Misiones — Dos de Mayo, XI; Aristabulo del Valle, XII; San Antonio, X; BRAZIL: Bahia — Rio Una, X; Espirito Santo — Vitoria, IX; Sao Leopoldina; Mi- nas Gerais — Ipatinga, XII; Mesquita, XI; Itacolmi, III; Rio de Janeiro — Rio de Janeiro, X-XII, III; Nova Friburgo; Teresopolis; Itatiaya, 700 m, III— IV; Santa Catarina — Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 103 i! Corupa, XI— XII; Sao Paulo — Itapetininga, II; Santos (Ilha Santo Amaro), IV; Mogi das Cruzes (Serra do Mar, Bora- ceia Biological Station), X. Phanaeus triangularis texensis Edmonds MEXICO: Coahuila — Sierra La Encantada, VII; UNIT- ED STATES: Texas — Houston; Austin, X; San Antonio, IX; Sheffield, VI— VII; Macdona, VII; Abilene, VII; Kerr- ville, IX; Mountain Home, VII; Leakey, VII; Sonora, X; Tyler, IV; Rio Frio, V; Alvin, IX; Brownsville, V-VI; Com- fort; Liberty; Columbus, VI; New Braunfels; San Angelo, VI; 5 mi W Cross, IX; Hockley; Plano, VII; Frio State Park, IX; Victoria; Dallas, V-VI; Roosevelt, X; Kingsville, X; Uvalde, V; Garner State Park, V; Eagle Pass, V; Men- ard, IV-VI; 15 mi S Marathon, VI; 33 mi S Marathon, VII; Cedar Hill, III-VI, X; Clay, IV; d’Hanis, VI; 5 mi NW Taylor, V; Ft. Worth; Dead Mans Canyon (Davis Mts.), VII (Burnett Co.; Lavaca Co.). Phanaeus triangularis triangularis (Say) UNITED STATES: Alabama — Monte Sano, VI; Arkan- sas— 2 mi S Foreman, VI (Carroll Co., VI, IX); Georgia — (Dougherty, VII— VIII; Rabun Co., VIII); Florida — Tor- reya State Park, V; Kansas — Osage (Riley Co.; Davis Co.; Douglas Co.); Kentucky — (Pulaski Co., VI); Louisiana — 12 mi SW Alexandria, IV; Kisatchie National Forest (Red Bluff Campground), IX (St. Charles Parish, V; Sabine Par- ish, IV); Oklahoma — 10 mi SSW Heavener, VI; 3.8 mi S Eagle Fork Creek bridge, Hwy 259; Mississippi — Santa Rosa, VI; Lucedale, VII— VIII; Missouri — Mark Twain National Forest, VII (St. Louis Co., VIII); North Caroli- na— Elizabethtown, IV; South Carolina — 3 mi S Wal- terboro, IX; Tennessee — Memphis (T. O. Fuller State Park), VI; Burrville, V-VII; Deer Lodge, VII; Reelfoot Lake, V; Texas — (Shelby Co., IV-VI, XI; Sabine Co., V-VI); Vir- ginia— Jones Creek. Phanaeus tridens pseudofurcosus Balthasar GUATEMALA: Huehuetenango — 16 km NW Huehue- tenango, VIII; MEXICO: Chiapas — Tuxtla Gutierrez, VI; 96 km S Tuxtla Gutierrez, 730 m, X; 65 km S Tuxtla Gutierrez, 820 m, XI; 3 km W Soyalo, 1220 m, IX; Corni- tan, V; 32.5 mi E Comitan, 2200 ft, IX; Las Delicias, IX- X; 18 km S La Trinitaria, 900 m, XII; 17 mi SE Teopisca, VI; 10 mi S Teopisca, VII; Santa Rosa, VII; junction Hwys 190 and 195, VII; Arriaga, VIII; 18 mi W Arriaga, VIII; 20 km NW Ocozocoautla, VIII; El Porvenir, 2300 m, IX; Mazapa, 1220 m, X; El Pozuelo, 1060 m, XI; 45 km SW Cintalapa, 2500 ft, VIII. Phanaeus tridens tridens Laporte-Castelnau MEXICO: Colima— Colima, VI; 20 km S Colima; Coli- ma-Jalisco state line at Hwy 110, VI; Jalisco — El Tuito, IX; El Tigre, 700 m, VII; Morelos— Tequesquitengo, VI (introduced?); Oaxaca — 15 mi N La Ventosa, VII; 5 mi NE Tehuantepec, 200 m, VIII; 13 mi SE Tehuantepec, 35 m, VIII; Veracruz — Palma Sola, 0 m, VI— VII; 12 km W Palma Sola, 300 m, VI; 6 mi SE Jalapa, VII; 14 mi SE Jalapa; 21 mi SE Jalapa, VII; 8 mi N Cuidad Mendoza, VII; 26 mi E Cuidad Mendoza, VII; Acultzingo, X; Salinas, VI; Conejos, 200 m, VIII; Presidio; 14 mi W Conejos, VI; Catemaco, V-VII; 21.7 mi NW Alvarado; 10 mi W Rin- conada; Tapalapa, X; 24 mi N Huatusco, 4800 ft, VII; Puente Nacional, VIII; Cotaxtla, IX; Trapiche del Cami- no, 550 m, I; 13 mi E Cuitlahuac. Phanaeus vindex Macleay MEXICO: Chihuahua — 15 mi S Camargo, IX; Gallego; Estacion Conchos, 4000 ft; UNITED STATES: Ala- bama— Mobile, VI; 6 mi S Jackson, V; Alabaster, IX; Enterprise, VI; Selma, VIII; Pinckard, III; Seddon, VI; Delchamps, VIII; Blount Springs, VIII; Birmingham, IX; Argo, IX; Tuskegee Army Air Field, VIII; Geneva, IV; Arizona — 3 mi S Willcox, VII; Willcox Dry Lake, VII; 12 mi NW Willcox; Ft. Huachuca, VII; Carr Canyon (Huachuca Mts.); 2 mi SE Apache, VIII; Portal, 4700 ft, IX; 10 mi E Portal, VIII; Tucson, IX; Fairbank, VIII; 8 mi E Drake, VIII; Prescott, VIII; Nogales, VIII; 12 mi E Sonoita; Phoenix; San Simon Valley, 4170 ft; 10 mi SE Patagonia, VIII; Tombstone, VIII; Ramsey Canyon (Hua- chuca Mts.), 6000 ft, VI; 1 mi E Douglas; Double Adobe, VII; 5 mi SW McNeal, VII; Sierra Vista, VIII; Webb, VII; Arkansas — Mena; Batesville, V; Imboden, III— VII, IX; Rich Mountain, IV; Pine Bluff, VI; Ozone, VII (Wash- ington Co.; LaFayette Co.); Colorado — Nunn; Eckley; Trinidad, VI; Limon, VI; Connecticut — Pomfret; Cole- brook, VII; Woodbridge, VII; Clinton, VIII; Putnam, IX; Florida — Clarkesville, VII; Monticello, III, IX, XI; Lutz; 13 mi N O’Brien, III; LaBelle, IV; Ocala, IV; Orlando; Ft. Drum; St. Petersburg; Jacksonville, VI; Lochloosa, I; Tam- pa, III; Miami, V, XII; Sanford; 6 mi E Greenville; Gains- 1 ville, V; Paradise, III; Bronson, IX; Pensacola; Tallahasee, IV; Ft. Myers, XI; Blountstown, III; Zolfo Springs, V; Brooksville, VI; Elfers, IV; Torreya State Park, VI; Lake Miccosukee, IV; Englewood, II; Orange, III; Anthony; Enterprize; Captivity Key; 3 mi E Rodman, VII; Brighton, VI; St Augustine; Lake Placid, III; Sarasota; Ormond, IV; Pass-a-grille, IV; 4 mi E Tarpon Springs; Lloyd, V; Moore Haven, VIII; 13 mi W Perry, VII; Newport, IX; Georgia— Tifton, VII; Swainsboro, VIII; Vidalia, X; Clayton, VI; Ft. Valley, VI; Camilla, IX; Reynolds; Nahunta, VI; Perry, IV; Rochelle, VIII; Atlanta, IV; Thomasville, VIII; States- boro; Dublin, VI; Putney, IV; Suches; Quitman, IX; 6 mi 5 Cordele; Athens, IX; Boston, IV; Ochlochnee, V; Blake- ly, VII; Butler, III; Fargo, V; Ellaville, IV; 12 mi WSW Faceville, VI; Acworth; Rutledge, VII; Moultrie, XI; Bain- bridge, VIII; Forsyth, IV; Okeefenokee National Wildlife Refuge, IX; Ft. Benning, VI; Brunswick, VI; Wadley, X; Savannah, X; Preston, V; Illinois — Glencoe; Chicago; Waggoner, VII; Mahomet, VI; Rockport, VI; Mossville, V; Indiana — Pekin, VII; State Forest, VII (Vigo Co.; Spen- cer Co.; Posey Co.; Laporte Co.; Marion Co.; Whitley Co.; Dubois Co.); Iowa — Knoxville, VII; Shenandoah, VII; Ames, VIII; Des Moines; Leon, VIII; Bloomfield, VI; Iowa City, V; 2 mi WSW Volga; Kansas — Elmdale, VI; Manhattan, V-VI, IX; Mt. Hope, IX; Onoga, IV; Lakin, 3000 ft, IX; Kansas City; Leavenworth, VI; Little Gobi Desert, VII; Medora; Abilene; Benedict; Ottawa, IV, IX; Baldwin, IX; Belvidere; Sylvia; Marquette, IX; Wathena; 6 mi N St. Mary’s, III; Ellis; Wellsville, VII; Wheeler; Osage; St. Francis; Dodge City (Geary Co.; Meade Co.; Commanche Co.; Hodgeman Co.; Clay Co.; Hamilton Co.; Montgomery Co.); Kentucky — Henderson, V; Tay- lorsville; Lexington, VIII; Rough River State Park, IV; Louisville, VII; Trenton, IX; Hickman, VII; Mammoth Cave National Park (Flint Ridge), VII; Water Valley, VI; Louisiana — 16 mi S Baton Rouge, XII; 7.7 mi S Ringgold, VIII; 3 mi W Winfield, VIII; 2 mi S De Ridder, VIII; Clayton, VII; 2 mi N Noble, VII; 7.9 mi W Kelly, VII; Shreveport, IX-XI; Alexandria, IV, VII; Lake Iatt, VI; New Orleans, IV; Covington; Vinton, IX; Frierson, V; Magnolia, IX; Kisatchie National Forest (Red Dirt Ref- uge), IX; Pearl River; Bogalusa, IV; Cameron, IV; Dry Prong, V; Simsboro, VIII; Leesville, III, VII; Maryland — 104 ■ Contributions in Science, Number 443 Edmonds: Revision of Phanaeus La Plata, V; Upper Marlboro, VI; Port Tobacco, VII; Bowie, VI; Towson, IX; Plum Point, VII; Massachu- setts— Northfield, VII; Sherborn, VII; Martha’s Vineyard, VIII; Woods Hole; Nantucket, IX; Springfield, VI; Wil- braham, VII; Fall River, VI; North Truro, IV; Amherst; Michigan — Ann Arbor, VIII; Wakeley, VII; Hartland, V, VII, X; Mississippi — State College, IV-VI; Lucedale, IV, VII, IX; Bay St. Louis, IX; Gainesville, XII; Richton, V; Leland; Cannonsburg, VI; Falkner, V; Missouri — Kansas City, VI; Columbia, V; Rock Port, V; Boonville, V; St. Louis, V; Louisiana, V-VI; Williamsville, VI, X; Butler, VIII; Allenton, V; Peevely, VI; Sedalia, V; Naylor, IX; 5 mi E Nevada, VI; Wentzville, VI; Albany, VII; Fayette, VII; Willard, VI; Charleston; Hayti, VII; St. Charles, VIII; Poplar Bluff, VII; Warrensburg, VIII; Alton, VIII; Halls, VII; Archie, VII; Nebraska — Benkelman; Omaha, VI; 3 mi S Hyannis, VII; Orleans, VII; Ogallala, VIII; Blair; Emerald, V; South Bend; Rulo; Oshkosh, IX; Lincoln, V; Halsey, VII; Eustis, V; Beaver City, VII (Franklin Co.); New Jersey — Phillipsburg; Egg Harbor; 3 mi NE Chats- worth, VIII; Ocean City, VIII; Barnegut Pinebarrens, VIII; Elizabeth; New Mexico — 2 mi N Rodeo, VIII; Albu- querque, VI; Koehler; Tucumcari, VI; Santa Fe, VII; Aco- ma, VII; Las Cruces, VIII; 15 mi N Las Cruces, VII; Las Lunas, VII— VIII; Hondo, VIII; Clayton, V; 10 mi S, 5 mi E Sofia, VIII; Deming, VII; Bernardo, VII; 8 mi E Las Vegas, VI (Eddy Co.; Grant Co.); New York — Fallsburg, IX; Shelter Island, IX; Orient; Somers, VIII; Rye, V; Brooklyn, VII; North Carolina — Beaufort, IX; Black Mountains, VIII; Southern Pines, IV; Chapel Hill, X; Car- thage, VII; Balsam, IX; Elizabeth City, IV; Ashville, VII; Cranberry; New Bern; Andrews, VIII; Graham, V; Greens- boro, VIII; Raleigh, XI; Reidsville, V; Faison, V, IX; Creedmore, IV; Willard, V; Lake Waccamaw, IX; Bryson City, 2000 ft, VIII; Grimesland, V; Highlands, IX; Ohio — Glen Este, VIII; Logan, V; Wooster, VII; Millport, V; Canton; Pickerington, VIII; Oklahoma — Stillwater, IX; Grainola, VII; Kenton, VI; Optima, VII; 5 mi S Sulphur, VI; Catoosa, VII— IX; Wichita Mountains Refuge, VI; Hin- ton, VII; 8 mi SE Guymon, VI; Spring Creek, VI; String- town, IV; Black Mesa State Park, VII; Geary, VI; Sayre, VIII; Boise City, VII; Bluejacket, VI; Davis, IV; Norman, IV; Woodward, V, VII; Clinton, VI; Bridgeport, VI (Cot- ton Co.; Marshal Co.; Payne Co.; Osage Co.; Choctaw Co.; Oklahoma Co.; McCurtain Co.; Latimer Co.); Penn- sylvania— Jeanette, V; Reading, VI; Downing Town, VII; Bethlehem; Philadelphia; Charleroi; Media, IX; Broomall, V; Slippery Rock, VI; Pittsburgh, VIII; Ambridge, VII; Rhode Island — Warwick, VII; Johnston, VIII; Barring- ton, VI; South Carolina — Aiken; Columbia, III; Charles- ton, IX; Florence, III; Clemson College, IX; Walterboro, IX; Camden, VI; White Pond, III; Bishopville, IX; Barn- well, IV; Rocky Bottom, V; Manning, V; Bluffton, V; Myrtle Beach, IV; Tennessee — Allardt, 1650 ft, VIII; Knoxville, IV; Kingston; Deer Lodge, VII; Roan Moun- tain, 6300 ft, VIII; Oak Ridge, X; Tullahoma, V; Burrville, VII— VIII; South Pittsburgh; Tusculum, VI; Newport; De- cherd; Nashville, VIII; Natchez Trace State Park, VI; Tex- as— Elkhart, V; Wascon, IV; Fedor, V-VI; Palo Duro State Park, VI— VII; Dallas, IV; New Braunfels; Stanton, VII; Abilene, X; Kingsville; Mt. Pleasant, IX; Higgins; Forestburg, V; College Station, V, VIII; New Boston, V; Wills Point, III— IV; Columbus; Big Spring; Amarillo, IV; Gainesville, V; Wichita Falls, III; Fred, V; Longview; 10 mi W Palestine, VII; Denton, V; Petrolia, IV; 4.5 mi N St Jo; Canadian, VIII; Angleton, VI; Perryton, V; Calvert, IV; Refugio, IV; Victoria, VIII; Nancy, IV; Weatherford; Stanton, VII; Sherman; Pampa, V; Nacagdoches, X; Bas- trop, III; 4 mi W Cooke’s Point, IX; New Ulm, IV; Mo- rales, IX; Frelsburg, IV; Brenham, IV; 9 mi E Yoakum, IX; Navasota, III; 7 mi S Seagoville, X; Pittsburg, IX; 1 mi N Centerville, VI; Paris, III; 10 mi SE Tyler, IX (Crosby Co.; Fayette Co.; Lavaca Co.; Eastland Co.; Shelby Co.) Vermont — Bennington; Virginia — Port Royal, IX; Wil- liamsville, IX; McKenney, IX; Holland, III; 5 mi S Waynes- boro, IX; Bedford Springs, VI— VII; Blacksburg; Virginia Beach, VII; Charlottesville, VIII; Farmville, IX; Falls Church, VI; Bristol, VI; Blue Ridge Parkway, mile 210, 2800 ft, V; West Virginia — Patterson Creek, VII; Glen Dale, VI; Wisconsin — Sauk City; Madison, VI; Wyo- ming— 7 mi NE Douglas, VII; Glendo, V. Phanaeus wagneri pilatei Harold GUATEMALA: Peten— Tikal, VII; MEXICO: Quintana Roo — Solferino, VII; Nuevo X-Can, V; 3 km E Tres Reyes, XI; 20 km N Felipe Carillo Puerto, VI; Yucatan — X-Can, VIII— IX; Tekax; Colonia Tucatan, VIII; Chichen Itza, VI; Uxmal, VI. Phanaeus wagneri wagneri Harold COSTA RICA: Cartago — Turrialba; Cachi, 2000 ft; Car- tago, XI; Guanacaste — 5 mi N Las Canas, VII; 20 km N Las Canas, VII; Sabana Grande, XI— XII; Santa Rosa Na- tional Park; San Jose — Aserri, 1300 m, VIII; San Jose, 1100 m, II, V— VIII; EL SALVADOR: Chalatenango — La Palma, VI; La Libertad — Quetzaltepeque, 500 m, VI-VII; San Salvador — San Salvador, VI; GUATEMALA: Chi- maltenango — Chimaltenango, 2000 m, VI, X; Chiqui- mula — Belen, IX; Guatemala — Guatemala, VII, X; Quetzaltenango — 17 km S Quetzaltenango, 4000 ft; Za- capa — La Union, X; HONDURAS: Comayagua — Minas de Oro, 4000 ft, V; Cortes — San Pedro Sula; Francisco Morazan — 5 km SW Suyapa, 5400 ft, VIII; Cedros, XI; MEXICO: Chiapas — 2 mi S La Trinitaria, 5200 ft, VIII; Bochil, VII; 9 mi SE Teopisca, VI; Aguacatenango, 1770 m, VIII; Cerro Huitepec, 2280 m, XI; El Sumidero, VIII; 12 mi S San Cristobal de las Casas, VIII. Contributions in Science, Number 443 Edmonds: Revision of Phanaeus ■ 105 Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 A/ Number 444 12 August 1994 CONTRIBUTIONS in Science Macropaleontology of the Eocene Crescent Formation in the Little River Area, Southern Olympic Peninsula, Washington Richard L. Squires and James L. Goedert Natural History Museum of Los Angeles County Serial Publications of THE Natural History Museum of Los Angeles County Scientific Publications Committee James L. Powell, Museum President Brian V. Brown Kenneth E. Campbell Daniel M. Cohen, Committee Chairman Kirk Fitzhugh Edward C. Wilson Robin A. Simpson, Managing Editor The scientific publications of the Natural History Mu- seum of Los Angeles County have been issued at irregular intervals in three major series; the issues in each series are numbered individually, and numbers run consecutively, regardless of the subject matter. # Contributions in Science, a miscellaneous series of tech- nical papers describing original research in the life and earth sciences. # Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth sci- ences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now published by the Museum in Contributions in Science. # Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through ij the Museum Book Shop. A catalog is available on request. The Museum also publishes Technical Reports, a mis- cellaneous series containing information relative to schol- arly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum’s Scientific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Tech- nical Reports may be obtained from the relevant Section i of the Museum. Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Printed at Allen Press, Inc., Lawrence, Kansas ISSN 0459-8113 Macropaleontology of the Eocene Crescent Formation in the Little River Area, Southern Olympic Peninsula, Washington Richard L. Squires1 and James L. Goedert2 ABSTRACT. In the upper part of the Crescent Formation in the Little River area, southern Olympic Mountains, Grays Harbor County, Washington, basalt units are intercalated with marine sedimentary units as much as 100 m thick. Ten macrofossil localities were found and 42 macrofossil taxa were identified to genus and/or species. These include 1 hexactinellid sponge, 1 colonial coral, 4 solitary corals, 1 inarticulate brachiopod, 2 articulate brachiopods, 1 polychaete worm, 1 scaphopod, 15 gastropods, 13 bivalves, 1 “gooseneck” barnacle, and 2 sharks. One new species is described and named: an inarticulate brachiopod, Craniscus edwilsoni, new species. The macrofauna is dominated by bivalves, limpets, and brachiopods that lived on a hard substrate produced by the accumulation of volcanic-island or pillow basalt in shallow marine waters. After a short distance of postmortem transport into adjacent subtidal waters, the macrofauna and eroded basalt debris accumulated as pebbly deposits. Parts of the animals that were cemented to the basalt did not survive the erosion/transport processes. Locally, the pebbly deposits grade into overlying muddy deposits where soft-bottom-dwelling solitary corals, mobile gastropods, shallow-burrowing bivalves, and hexactinellid sponges lived. Macrofossils in the upper part of the Crescent Formation in the Little River area indicate a middle early Eocene age equivalent to the molluscan “Capay Stage.” Three of the genera previously known from only the Atlantic Ocean are reported for the first time from the Paleogene of the Pacific coast of North America. They are the brachiopod Craniscus, the “gooseneck” barnacle Aporolepas, and the shark Palaeohypotodus. Eight genera, three tentatively identified genera, and 16 species (mostly mollusks) previously known from the Pacific coast are reported for the first time from the upper Crescent Formation. Of these, the gastropods Diodora and Erginus and the bivalve Spondylus carlosensis have their earliest record in the Little River area. A few of these previously known taxa may have immigrated into the Little River area from Alaska or Russia. INTRODUCTION The basement rock in the Olympic Peninsula of western Washington is the upper Paleocene to low- er middle Eocene Crescent Formation (Snavely, 1987). The formation consists predominantly of oceanic tholeiite basalt flows. Several models pro- posed for the origin of the formation were reviewed by Snavely (1987). Most envisage accretion of sea- mounts. The upper third of the Crescent Formation ranges from a deep-to-shallow marine environment to one that is locally terrestrial with marine sedi- mentary rocks interbedded with the basalt flows (Cady, 1975). Locally, these marine sedimentary rocks contain fossils. There have been several de- tailed studies of the benthic foraminiferal assem- 1. Department of Geological Sciences, California State University, Northridge, California 91330, and Research Associate in Invertebrate Paleontology, Natural History Museum of Los Angeles County. 2. 15207 84th Avenue Ct. NW, Gig Harbor, Wash- ington 98329, and Museum Associate in Vertebrate Pa- leontology, Natural History Museum of Los Angeles County. Contributions in Science, Number 444, pp. 1-32 Natural History Museum of Los Angeles County, 1994 blages (Rau, 1964, 1966, 1967), but, until recently, there were no detailed studies of the macrofossil assemblages. Prior to 1992, the previous macro- fossil studies dealt with a few new species of mol- lusks or corals mainly from the type section of the Crescent Formation at Crescent Bay (Fig. 1) along the south shore of the Strait of Juan de Fuca. Squires et al. (1992) studied the shallow marine upper part of the Crescent Formation at Pulali Point (Fig. 1), west of Seattle, and made the first detailed analysis of a macrofossil assemblage in the formation. Al- though Weaver (1943) did a monograph on Tertiary marine macrofossils from Washington and Oregon, his coverage of species from the Crescent Forma- tion was very limited. The present study is a continuation of work by the authors on the paleontology of little-studied early Eocene macrofaunal assemblages in the Olympic Peninsula. It summarizes the macrofossil assemblages of the upper Crescent Formation in the Little River area, southern part of the Olympic Mountains, Grays Harbor County, Washington (Fig. 1), and discusses the paleoenvironmental, biostrati- graphic, and paleobiogeographic aspects of the as- semblage. Figure 1. Index map showing the location of the Little River area and outcrops (shown as parallel inclined lines) of the Crescent Formation (compiled from Tabor and Cady, 1978; Walsh et ah, 1987) in the Olympic Peninsula, Washington. The most detailed geologic mapping of the study area and vicinity was done by Rau (1966), whose work was used by Tabor and Cady (1978) in their compiled geologic map of the Olympic Peninsula. Rau (1966) did a detailed study of the benthic fo- raminiferal assemblage from the upper Crescent Formation in the study area and noted the presence of mollusk shell fragments in some of the beds. Armentrout (1973) identified the genera of a few of the more abundant mollusks. Our study is the first species-level study of the macrofauna in the Little River area. Grain and clast (> 2 mm) size terms used in the present study are from Wentworth (1922). Litho- logic and grain roundness terms follow those used in Compton (1985). The molluscan stage termi- nology used in this report stems from Clark and Vokes (1936), who proposed five mollusk-based provincial Eocene stages: “Meganos,” “Capay,” “Domengine,” “Transition,” and “Tejon.” The stage names are in quotes because they are informal terms. Givens (1974) modified the use of the “Capay Stage,” and it is in this modified sense that “Capay Stage” is used herein. Saul (1983) correlated the stages to the calcareous nannofossil zonation of Okada and Bukry (1980). 2 ■ Contributions in Science, Number 444 STRATIGRAPHY Outcrops of the upper Crescent Formation in the Little River area are part of a monoclinal structure in the foothills along the southern side of the Olym- pic Mountains. The outcrops strike generally north- west and dip about 40° southwest. Although partial sections were measured at all the places where mac- rofossils were collected, structural and facies com- plexities, as well as dense forest cover, make it extremely difficult to correlate these sections. As a result, the stratigraphy of the upper Crescent For- mation in the study area can be depicted only in an idealized manner (Fig. 2), with the stratigraphic positions of the macrofossil localities only shown approximately. California State University, North- ridge (CSUN) localities 1553, 1553a, and 1554 are especially tenuous due to fault problems. Only the upper part of the Crescent Formation was studied. Most of the non-sedimentary rocks are weathered pillow basalt. Some individual flows, especially in the lower part of the section, are in- tensely fractured and commonly appear to be brec- ciated or sheared. Rau (1966) observed these frac- ture zones and attributed them to autobrecciation, in some cases, and to structural deformation, in others. Squires and Goedert: Crescent Formation Eocene Fossils The main macrofossiliferous sedimentary unit is approximately 100 m thick and consists mostly of dark green, very fine to medium basaltic sandstone with rounded to subrounded grains. Near the base of this sedimentary unit, there are pebble conglom- erates (clasts up to 13 mm in length) that grade rapidly upward into coarse sandstone. These pebble conglomerates and associated sandstones locally contain many macrofossils. At CSUN locality 1553 (Fig. 3) there is a 1-m-thick sequence of these rocks that is the most fossiliferous deposit in the study area. Macrofossils make up as much as 30 percent of the clasts and form a coquina. Thirty-five species were found associated with a diverse microfossil assemblage. Intermixed with the macrofossils are subrounded basalt clasts that range in size from granule to medium (up to 11 mm in length). There are also some large pebble-sized (up to 20 mm in length) lumpy calcareous nodules. The matrix of the pebble conglomerate is mudstone and the amount is variable. In the upper part of the 1-m- thick sequence, the matrix makes up 60 percent of the rock, and the rock type is a sandy mudstone with shell debris and some scattered small pebbles of basalt. Locality 1553 is similar to localities 1554, 1555, and 1557 in rock type and abundance and diversity of macrofauna. At locality 1557, there is a thin lens of Glycymeris sp. cf. G. crescentensis coquina. The stratigraphic position of CSUN locality 1555, which is known with more certainty than for CSUN lo- calities 1553, 1554, and 1557, is in the lower part of the main sedimentary unit. Localities 1553, 1554, and 1557, therefore, also are placed in the lower part of the section (Fig. 2). Approximately 2 m upsection at locality 1553, there is a 3-m-thick, pyritic argillaceous black mudstone with sparse macrofossils, including some hexactinellid sponges. Locality 1553a is in this black mudstone, which is not present elsewhere in the study area. Near the middle of the main sedimentary unit, macrofossils were found at only one locality (CSUN loc. 1560). A few species were recovered from silty very fine sandstone to argillaceous fine sandstone. At the top of the main sedimentary unit, there are pebble sandstones that do not have muddy matrix and have only a few macrofossils. The basalt unit overlying the main sedimentary unit is approximately 90 m thick. It consists mostly of numerous, thick-bedded pillow basalts, but near the top of the unit, in the vicinity of locality 1556, there is a flow that shows columnar jointing along its lower and upper contacts. The middle of the flow is massive but fractured. There are minor de- posits of siltstone interbedded with the basalt unit, but these siltstones contain no macrofossils. Unconformably overlying the Crescent Forma- tion in the Little River area is a sequence of gray micaceous thin-bedded mudstone, siltstone, and silty sandstone. Rau (1984) indicated on a geologic sketch map that these strata should be mapped as part of the Humptulips formation that he informally named Figure 2. Idealized stratigraphy of the upper Crescent Formation in the Little River area and approximate strati- graphic positions of CSUN macrofossil localities. and recognized just west of the study area. These strata were originally mapped as “sedimentary rocks of late Eocene age” (Rau, 1966; Tabor and Cady, 1978). Rau (1966) mapped a nearly continuous band of upper Crescent Formation sedimentary rocks above the highest volcanic rocks throughout the study area. This interpretation does not correspond to the general structure of the Crescent Formation within the study area as we interpret it, and this band of sedimentary rocks could not be found at all of the areas indicated by Rau (1966). In addition, basalt is exposed in roadcuts in the south half of SW14 of the SEVi of section 12, in an area previously mapped (Rau, 1966) as “sedimentary rocks of the late Eocene age” (= Humptulips formation). Basalt and sedimentary rocks of the Crescent Formation are also present in the NEVt of the NW14 of section 15, again in an area mapped (Rau, 1966) as Hump- tulips formation. Detailed work (beyond the scope Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 3 Figure 3. Index map showing location of CSUN macrofossil localities in the Little River area. Contour interval is 400 ft. Base map is U.S. Geological Survey, 7.5 minute, Grisdale, Washington, quadrangle (provisional edition, 1990). of this paper) on the structure of this part of the Crescent Formation is needed to accurately deter- mine the stratigraphic relationships of the macro- fossiliferous strata and the basalt units. Rau (1966) also believed that the sedimentary deposits in the vicinity of locality 1553 are above the stratigraphically highest volcanic rock because there is no overlying basalt there. Our interpretation is that stratigraphically higher volcanic units have been eroded or faulted out in the vicinity of locality 1553 (as well as loc. 1554). At locality 1553, the contact with underlying pillow basalt is a vertical fault of unknown displacement. FAUNAS A total of 776 macrofossils were collected from 10 localities (Fig. 3). Preservation ranges from poor to good, but many of the fossils are in poor condition. In addition, fossils in mudtone matrix are easily destroyed if not carefully removed from the brittle matrix. Limpet gastropods are mostly preserved as internal molds. Most of the macrofossils are about 10-15 mm in maximum dimension. Forty-two of the macrofossil taxa could be iden- tified to genus and / or species. These include 1 hex- actinelled sponge, 1 colonial coral, 4 solitary corals, 1 inarticulate brachiopod, 2 articulate brachiopods, 1 polychaete worm, 1 scaphopod, 15 gastropods, 13 bivalves, 1 “gooseneck” barnacle, and 2 sharks. Other taxa too poorly preserved for generic iden- tification include calcareous sponges?, bryozoans, minute serpulid worms, a few gastropods and bi- valves, an anomurid (thalassinoid) shrimp claw, a 4 ■ Contributions in Science, Number 444 xanthid? crab claw, sea urchin spines and plates, | and two fragments of irregular echinoids. On the basis of comparison with Crescent For- mation macrofaunal data in Squires et al. (1992), ,j eight genera, three tentatively identified genera, and 16 species (Table 1) are reported for the first time from the Crescent Formation. Two of the species, Calyptraea diegoana and Bracbidontes (B.) cow- litzensis, have been reported previously from the Crescent Formation, but the previous records were questionable (Squires et al, 1992). One new species is reported from the Crescent Formation in the Little River area. It is the inarticulate brachiopod Craniscus edwilsoni, new species. Rau (1966) made a detailed study of benthic for- aminifers from the upper part of the Crescent For- mation in the study area and found 55 species. His localities F-36 and F-37 are in the immediate vicin- ity of locality 1553. At locality 1553, abundant large benthic foraminifers, as well as some fish otoliths, are detectable with a hand lens. A few radiolarians were found at localities 1553 and 1560. A microfossil sample was collected at locality 1553 and analyzed for benthic foraminifers and calcareous nannofossils. The benthic foraminiferal assemblage is dominated by a robust species of Amphistegina. There are also fragments of prob- able discocyclinids, along with rare specimens of Quinqueloculina sp. and Nodosaria latejugata Giimbel, 1868 (Heitman, pers. comm., 1992). Only very rare, poorly preserved specimens of the cal- careous nannofossils Dictyococcites sp., Coccolith- us pelagicus (Wallich, 1877), and Toweius (?) sp. were found (M.V. Filewicz, pers. comm., 1992). Squires and Goedert: Crescent Formation Eocene Fossils DEPOSITIONAL ENVIRONMENT the grains and clasts in the upper Crescent For- mation sedimentary deposits were derived by high- energy erosion of pillow basalts that were extruded into shallow water. These basalt buildups provided a hard-substrate habitat for marine invertebrates, and macrofossils in the sedimentary deposits are indicative of shallow water and were predomi- nantly hard-substrate-dwelling species. All of these animals were transported after death into adjacent j subtidal waters where there were soft-subtrate sands and muds. The distance of postmortem transport j was not far because the shells are unabraded and | mostly complete. Four types of hard-substrate-dwelling animals are | present that would have been able to live on sur- faces of the pillow basalts. They are bivalves at- tached by byssal threads ( Barbatia , Brachidontes, and Nayadina ( Exputens )); bivalves, brachiopods, and polychaete worms attached by cementation (, Spondylus , Ostrea, Craniscus, and Rotularia ); limpets and turbinid gastropods ( Erginus , Patelloi- da ?, Diodora, and Homalopoma ); and brachio- pods attached by a pedicle ( Gryphus ? and Terebra- tulina). In addition, two specimens of an encrusting colonial coral ( Leptastrea ?) are present. Modern species of the byssate bivalves Barbatia and Brachidontes commonly live wedged in among rocks or colonial corals in intertidal or shallow subtidal waters (Stanley, 1970). Nayadina ( Expu- tens) batequensis was interpreted to have been bys- sate and to have lived in shallow marine waters associated with reef corals (Squires, 1990a; Squires and Demetrion, 1992). This bivalve is the most common macrofossil in the upper Crescent For- mation and 120 specimens were found at locality 1553. All specimens are disarticulated, but there are equal numbers of right and left valves indicating that the distance of postmortem transport was short. At locality 1554, an articulated specimen of this species was found. Modern species of the bivalves Spondylus and Ostrea are commonly cemented to rocks, shells, and colonial corals in the intertidal zone and off- shore in shallow subtidal depths (Keen, 1971; Ab- bott and Dance, 1982). Spondylus carlosensis is present at nearly every locality in the study area. At locality 1553, it is the second-most abundant macrofossil, and 60 specimens were found. All are free (non-cemented) valves, except for a single spec- imen of a left valve cemented to another S. car- losensis. Like Spondylus, the specimens of Ostrea are nearly all free valves. Only a single specimen of a cemented valve was found and it was attached to the brachiopod Craniscus. The near absence of the cemented valves of these types of bivalves indicates that they only rarely survived the erosion of the rocky substrate. A modern species of a similar crani- acean brachiopod, Neocrania californica Berry, 1921, can be found cemented to rocky walls of glacially formed fjords in depths as shallow as 35 Table 1. Little River area macrofossil taxa reported for the first time from the Crescent Formation. Hexactinellid sponge Apbrocallistes ? sp. Colonial coral }Leptastrea hertleini Durham Solitary corals Balanophyllia variabilis Nomland Trocbocyathus crooki Merriam and Turner Turbinolia dickersoni Nomland Inarticulate brachiopod Craniscus edwilsoni, new species Articulate brachiopod Gryphus ? sp. Gastropods Calyptraea diegoana (Conrad) Colwellia sp. Conus sp. Dentimitra cretacea (Gabb) Diodora sp. Erginus vaderensis (Lindberg) Homalopoma sp. cf. H. umpquaensis domenginensis Vokes Homalopoma sp. aff. H. wattsi (Dickerson) Lyria andersoni Waring Patelloida ? sp. cf. P. vokesi (Hickman) Patelloida} sp. Surculites mathewsonii (Gabb) Tenagodus bajaensis Squires Turritella andersoni Dickerson Bivalves Acila ( Truncacila ) decisa (Conrad) Barbatia sp. aff. B. (B.) landesi (Weaver & Palmer) Brachidontes (B.) cowlitzensis (Weaver &c Palmer) Corbula ( Caryocorbula ) dickersoni Weaver & Palmer Glycymeris ( Glycmerita ) sagittata (Gabb) Nayadina ( Exputens ) batequensis Squires Spondylus carlosensis Anderson Barnacle Aporolepas sp. Shark Isurus ? sp. Palaeohypodotus sp. m in northern British Columbia (Tunnicliffe and Wilson, 1988). This modern species, which was formerly assigned to genus Crania Retzius, 1781, was recently reassigned by Lee and Brunton (1986) to genus Neocrania Lee and Brunton, 1986. Spec- imens of Craniscus were found only at locality 1553, and like the Spondylus and Ostrea speci- mens, nearly all are free (non-cemented) brachial valves. The polychaete worm Rotularia ( Rotularia ) is extinct but is a common component in shallow Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 5 marine Eocene deposits on the Pacific coast of North America (Squires, 1984, 1987, 1988a). Evidence for attachment for Rotularia (R.) tejonense is a speci- men found attached to a pebble at locality 1553, and this specimen (showing the attachment scar) is illustrated in Figures 19 and 20. Modern species of the limpets Patelloida and Diodora typically inhabit wave-swept rocky coasts (Keen, 1971; McLean, 1978). Modern species of Erginus live intertidally or subtidally (Lindberg, 1988). Most of the limpets found in the upper Cres- cent section are complete and those that possess radial ribbing are unabraded; therefore, the distance of postmortem transport had to have been short. Reports of limpet faunas are uncommon for the Paleogene of the Pacific coast of North America because shells of taxa from this high-energy envi- ronment rarely survive. As mentioned by Lindberg and Squires (1990), subtidal environments juxta- posed to rocky intertidal habitats provide a good preservational site for the limpets. Other gastropods in the upper Crescent Formation that lived on the pillow basalts are two species of the turbinid Hom- alopoma. Modern Homalopoma lives on hard sub- strates in intertidal and shallow subtidal depths (McLean, 1978). The brachiopod T erebratulina unguicula is an extant species found from Kamchatka to San Diego on rocky bottoms in depths as shallow as 10 m (Hertlein and Grant, 1944). The species is common in glacially formed fjords in northern British Co- lumbia and lives attached to rocky walls in depths less than 100 m where turbulent tidal action is present (Tunnicliffe and Wilson, 1988). Numerous single valves of T erebratulina unguicula weaveri were found at locality 1553, but only one articu- lated specimen was found. Little is known about the ecology of modern Gryphus, but species are known to inhabit deep waters near volcanic islands like the Galapagos Islands (Hertlein and Grant, 1944). An entire 8-cm-long corallum of the colonial coral }Leptastrea hertleini was found at locality 1553. This colonial coral may have also lived at- tached to the substrate because living species of Leptastrea commonly encrust in reef habitats in shallow tropical seas in the Indo-Pacific region and Red Sea area (Durham, 1942; Wood, 1983). Lep- tastrea is a hermatypic faviid colonial coral (Wells, 1956). Hermatypic corals are restricted to shallow waters by the light requirements of their symbiotic algae and flourish in depths less than 50 m (Britton and Morton, 1989). There are some macrofossils in the Crescent For- mation that could have lived among the rocky and shelly rubble deposited adjacent to the rocky sur- face of the basalt flows. They are sessile gastropods ( Calyptraea ), a cowrie-like gastropod ( Eratotrivia ), a small bivalve ( Corbula ), and sea urchins. Modern species of Calyptraea live attached to other shells and rocks in the intertidal zone and shallow subtidal depths (Keen, 1971). Trivid gastropods, similar to 6 ■ Contributions in Science, Number 444 the extinct Eratotrivia, are commonly found among rocky rubble intertidally and subtidally in warm waters (Abbott and Dance, 1982). Although most modern species of Corbula live offshore on sandy bottoms in relatively shallow waters, some live in- tertidally under rocks or among gravel and cling by means of a byssus (Keen, 1971). Modern sea urchins (cidaroids) prefer a hard bottom, such as rocks or shell debris (Fell, 1966). Calyptraea specimens are moderately common in the Crescent beds, but only a few specimens of Eratotrivia and Corbula were found. The sea urchin remains are quite common, consisting of only the spines and basal plates of spines, and the spines possess delicate spinelets that show no signs of abrasion. Two capitular plates of the “gooseneck” bar- nacle Aporolepas were found at localities 1553 and 1554. This extinct genus is indicative of very shal- low marine waters where it lived attached to hard substrate (V.A. Zullo, pers. comm., 1993). Several specimens of the sessile gastropod Tena- godus were found at localities 1553 and 1553a. They may have lived embedded in sponges like certain modern species of Tenagodus that have been reported in the western Atlantic Ocean (Morton, 1955). Some of the macrofossils in the sedimentary de- posits were soft-substrate-dwelling mollusks, in- I eluding the scaphopod ( Dentalium ), mobile gastro- pods ( Turritella , Lyria, Dentimitra, Surculites, and Conus), and shallow-burrowing bivalves (Acila, Glycymeris, Venericardia, and Nemocardium). Nearly all of these genera are extant and have been reported from relatively shallow, offshore sandy to muddy bottoms (Abbott and Dance, 1982; Squires, 1984). Additional soft-substrate-dwelling taxa in the Crescent Formation are the ahermatypic solitary corals Flabellum, T rochocyathus, and possibly Bal- anophyllia. Little is known about the living habits of most of the modern ahermatypic solitary corals, but Flabellum is usually found on muddy sub- strates, probably with the sharp base pushed deeper into the mud as the coral increases in size (Vaughan and Wells, 1943). Cairns (1989) reported a depth range of 22-2,300 m for modern species of Fla- bellum s.s. from the Philippine Islands and adjacent waters. Modern species of T rochocyathus have a similar depth range of 32-1,573 m (Vaughan and Wells, 1943) and probably live like Flabellum. Sol- itary corals are common at locality 1553, where they are mostly complete and have partial size se- ries. The muddy matrix of the sandstone and the mudstones at this locality could have provided suit- able soft substrate for these corals. The most abun- dant solitary coral at locality 1553, however, is Bal- anopbyllia. Modern species can live on or under rocks in protected niches in the low intertidal zone where waves beat the hardest (Durham, 1947; Mor- ris et al., 1980) or on soft substrates in depths to approximately 1,000 m (Vaughan and Wells, 1943). The Balanophyllia specimens at locality 1553 were Squires and Goedert: Crescent Formation Eocene Fossils most likely transported from the nearby intertidal zone into the adjacent deeper, muddier waters. Armentrout (1973) concluded that the macro- fossils he found at his locality LR-1 (see also Ar- mentrout, 1975), which is equivalent to CSUN 1553, are suggestive of a very shallow nearshore environ- ment of perhaps less than 18 m. He based his in- terpretation on the presence of Terebratulina, Ac- maea, Turritella, Calyptraea, Barbatia, and Glycymeris in the coarse sediment. Benthic foraminifers from the microfossil sample at locality 1553 yielded an inner to middle neritic assemblage indicative of warm, clear water (H.L. Heitman, pers. comm., 1992). There are probable discocyclinid fragments in this sample, and these would indicate shallow marine conditions. Else- where in lower to middle Eocene strata along the Pacific coast of North America, specimens of dis- cocyclinids have been found in nearshore to off- shore, normal-salinity environments (Squires, 1984; Squires and Demetrion, 1992). Vaughan (1945) re- ported that discocyclinids are indicative of depths less than 100 m. Rau (1966) found the benthic foraminifers Amphistegina, Quinqueloculina, and certain Cibicides from locality 1553 and reported that they indicate possible reef conditions, perhaps in places less than 30 m in depth. Locality 1553a represents the muddiest macro- fossil deposit. It is a very thin deposit and not lat- erally extensive. Macrofossils are sparse and rep- resent some of the substrate-attached species found farther downsection and a few small-sized, thin- shelled nuculanid? bivalve species found nowhere else in the section. According to Keen and Coan (1974), nuculanids are usually deep-water bivalves. The hexactinellid sponge Aphrocallistes ? becomes more plentiful at locality 1553a. According to Rig- by and Jenkins (1983), this type of sponge is common in deep waters. The macrofaunal assemblage at this locality is a mixture of storm-derived shallow ma- rine species deposited with indigenous deep-water offshore species. The depositional environment of the upper Cres- cent Formation in the Little River area is very sim- ilar to that in the upper Crescent Formation at Pulali Point in the eastern Olympic Peninsula, Washing- ton. The Pulali Point area is approximately 70 km northeast of the Little River area (Fig. 1). At Pulali Point, the upper Crescent Formation contains pil- low basalts interbedded with macrofossiliferous sedimentary rocks (Squires et al., 1992). Like the study area, the extrusion of each basalt unit at Pulali Point caused shoaling of the marine waters. Eroded rubble was transported a short distance into shal- low subtidal depths. Living mostly among the basalt rubble were substrate-attaching species of bivalves (. Pododesmus [see Squires, 1993] and Ostrea ), rhyn- chonellid brachiopods ( Hemithiris ), and colonial corals. The solitary coral identified by Squires et al. (1992) as tTrochocyathus nomlandi Bentson, 1943, from the upper Crescent Formation at Pulali Point is actually Flabellum californicum Vaughan, 1900, which is also present in the upper Crescent For- mation in the Little River area. The depositional environment of the upper Cres- cent Formation in the Little River area also is similar to that in the lower Hamlet formation (informal name of Niem and Niem, 1985) in northwestern Oregon. The upper middle Eocene Roy Creek member of this formation at Rocky Point quarry consists of bouldery rubble derived from basaltic headlands and deposited in pocket beaches along a storm-dominated coastline (Mumford and Niem, 1992). Macrofossils are common and are domi- nated by substrate-attaching animals, including bi- valves attached by byssal threads ( Barbatia , Podo- desmus, Mytilus, and Septifer ), bivalves attached by cementation {Ostrea), limpets ( Acmaea ), sessile gastropods ( Calyptraea ), articulate brachiopods (. Rhynchonella and “ Terebratula ”), and colonial corals (Warren and Norbisrath, 1946; Steere, 1957; Niem and Van Atta, 1973). AGE Sedimentary units in the upper Crescent Formation in the Little River area are middle early Eocene on the basis of macrofossils and benthic foraminifers. The most age-diagnostic macrofossils are Turritella andersoni, Tenagodus bajaensis, Glycymeris ( Tu - cetona) fresnoensis, and Nayadina ( Exputens ) ba- tequensis. They are known elsewhere from the Pa- cific coast of North America in strata that have been well documented as correlative with the pro- vincial molluscan “Capay Stage” of middle early Eocene age (Vokes, 1939; Squires, 1984, 1987, 1988a, 1988b, 1990a, 1990b; Squires and Deme- trion, 1992). Additional age-diagnostic macrofossils are ILeptastrea bertleini, T rochocyatbus crooki, and Eratotrivia crescentensis. They are not as well doc- umented in their geologic ranges, but they have been reported elsewhere from the Pacific coast of North America in strata that would be considered by most workers to be correlative with the “Capay Stage” (Weaver and Palmer, 1922; Merriam and Turner, 1937; Durham, 1942; Bentson, 1943; Weaver, 1943). On the basis of benthic foraminifers, Rau (1966, 1981) reported that the upper Crescent Formation in the Little River area is correlative to the Ulatisian Stage (as used by Mallory, 1959, and applied by Rau, 1981). Almgren et al. (1988) emended the age of this stage and showed that it is middle early Eocene through late early Eocene. Squires et al. (1992) illustrated how the Ulatisian Stage (in this emended sense) is equivalent to the upper part of the molluscan “Capay Stage” and to the lower part of the “Domengine Stage.” The calcareous nan- nofossil assemblage from CSUN locality 1553 yielded long-ranging species that are only indicative of Eocene undifferentiated (M.V. Filewicz, pers. comm., 1992). The microfossil and macrofossil age determinations, therefore, indicate a middle early Eocene age (equivalent to the interval in which the Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 7 foraminiferal Ulatisian Stage, in the emended sense, and the molluscan “Capay Stage” overlap). PALEOCLIMATE AND PALEOBIOGEOGRAPHY Early Eocene time was the warmest interval of the Cenozoic, and tropical to subtropical conditions were widespread. Addicott (1970) reported that warm-water conditions existed at that time as far north as the Gulf of Alaska. There was no con- nection between the Arctic Ocean and the north- eastern Pacific Ocean (Marincovich, 1992), and tropical macrofaunas inhabited areas as far north as the northern shore of the Olympic Peninsula (Durham, 1942, 1950). The geographic dispersal of the Little River Cres- cent Formation’s inarticulate brachiopod Cranis- cus, the bivalve Nayadina ( Exputens ) batequensis, and the shark Paleohypotodus was dramatically re- lated to these early Eocene events. Previously, North American fossil craniacean brachiopods have been found only rarely in lower Tertiary rocks on the Atlantic and Gulf coasts of the United States, and their generic assignments have been tentative (Toul- min, 1940; Cooper, 1988). The presence of Cranis- cus in the upper Crescent Formation in the Little River area, therefore, is very significant. This is the first report of this genus from North America. Pre- viously, it has been reported only from Europe, Japan, Australia, and the Indo-Pacific region (Lee and Brunton, 1986). It is likely that Craniscus reached the Pacific coast of North America by way of tropical currents that flowed westwardly from southern Europe and the Mediterranean region to- ward the New World. Squires (1987) discussed that the dispersal route of Old World (Tethyan) mac- roinvertebrate taxa was probably via a seaway in the southern Central American region. Nayadina ( Exputens ) is a warm-water malleid bivalve with Old World Tethyan affinities, and N. (£.) batequensis is the earliest known species. Ap- parently, during the early Eocene it migrated west- erly from Jamaica (Squires, 1992) to Baja California Sur, Mexico (Squires, 1990a, 1992; Squires and De- metrion, 1992), and then northerly into the Little River area. Additional evidence of migration of taxa from the Atlantic Ocean region into the Pacific coast of North America is the limpet Diodora, the “goose- neck” barnacle Aporolepas, and the shark Palaeo- hypotodus. The Little River specimens of Diodora are the earliest record of this genus from the Pacific coast of North America. Paleocene and other early Eocene records, however, are known from the southern and eastern United States (Palmer and Brann, 1966). Aporolepas is primarily known from western Europe and the Gulf coast of North Amer- ica (Newman et al., 1969; V.A. Zullo, pers. comm., 1993). The shark P alaeohypotodus is a monotypic genus whose single species P. rutoti (Winkler, 1874) has been reported from Paleocene rocks in Green- 8 ■ Contributions in Science, Number 444 land, western Europe, Maryland, and North and South Dakota, as well as from lower Eocene rocks in western Europe (Cappetta, 1987; Ward and Wiest, 1990; Cvancara and Hoganson, 1993). Only a single tooth of this genus was found in the upper Crescent Formation, but it is well preserved and closely re- sembles P. rutoti Winkler. Of the identified species reported for the first time from the Crescent Formation (Table 1), migration into the Little River area was mostly from the south based on their previously known geographic dis- tribution (Nomland, 1916; Merriam and Turner, 1937; Vokes, 1939; Squires, 1984, 1987; Squires and Demetrion, 1992). The upper Crescent Formation in the Little River area is now the known northern limit of geographic distribution of most of these species. The exceptions are the bivalves Acila (' Truncacila ) decisa, Brachidontes (B.) cowlitzen- sis, and Glycymeris ( Glycymerita ) sagittata, whose northern limit of geographic distribution is in either Alaska or Russia (Devyatilova and Volobueva, 1981; Marincovich, 1988). These species may have emi- grated southerly from Alaska or Russia into the Pacific coast region of North America. The limpet gastropod Erginus vaderensis has not been found farther south than the upper Eocene “Gries Ranch beds” of the Lincoln Creek Forma- tion in southwestern Washington, and the modern distribution of the genus is throughout the boreal Pacific Ocean, Arctic Ocean, and North Atlantic Ocean (Lindberg, 1988). Erginus vaderensis is an- other one of the Little River Crescent Formation species that may have originated in northern lati- tudes and then migrated to the south. The Little River area specimens of this gastropod are the ear- liest record of this genus. SYSTEMATIC MATERIALS AND METHODS Synonymies, primary type material, molluscan stage range, geographic distribution, local occurrence, and remarks are listed for species/subspecies. Terms used to denote spec- imen abundance are defined as follows (number or spec- imens in parentheses): rare (1-4), uncommon (5-9), com- mon (10-29), and abundant (30 or more). Abbreviations for catalog and/or locality numbers are as follows: ANSP: Academy of Natural Sciences, Philadelphia. CAS: California Academy of Sciences, San Francisco. CSUN: California State University, Northridge. IGM: Instituto de Geologia, Universidad Nacional Au- tonoma de Mexico, Mexico City. LACMIP: Natural History Museum of Los Angeles Coun- ty, Invertebrate Paleontology Section. LACMVP: Natural History Museum of Los Angeles County, Vertebrate Paleontology Section. SU: Stanford University (collections now housed at the California Academy of Sciences). UCMP: University of California, Museum of Paleontol- ogy, Berkeley. USNM: National Museum of Natural History, Washing- ton, D.C. UW: University of Washington, Seattle. Squires and Goedert: Crescent Formation Eocene Fossils The bulk of the Little River area macrofossil collection is housed at CSUN. New species primary type material and hypotypes of the invertebrates used for illustrations in this report are deposited at LACMIP. The shark-teeth hypotypes are deposited at LACMVP. SYSTEMATICS Phylum Porifera Grant, 1861 Class Hexactinellida Schmidt, 1870 Family Aphrocallistidae Gray, 1867 Genus Aphrocallistes Gray, 1858 Aphrocallistes ? sp. Figure 4 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12220. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1553a. REMARKS. Specimens are uncommon at local- ity 1553a and rare at locality 1553. Preservation of the sheet-like or knobby fragments is as internal molds. Walls of the specimens, which are less than 7 mm in length, show a honeycomb-like, polygonal pattern like the morphology of Aphrocallistes. Preservation is too poor to allow positive identifi- cation to genus. If the Little River area specimens of Aphrocal- listes} eventually do prove to be Aphrocallistes, they would be the earliest record of this genus on the Pacific coast of North America. Previously, Aphrocallistes has been reported by Rigby and Jen- kins (1983) from upper Eocene to Miocene rocks of southwestern Washington. It has not been pre- viously reported from the Crescent Formation. Phylum Cnidaria Flatschek, 1888 Class Anthozoa Ehrenberg, 1834 Order Scleractinia Bourne, 1900 Family Faviidae Gregory, 1900 Genus Leptastrea Milne Edwards and Haime, 1848 }Leptastrea hertleini Durham, 1942 Figure 5 Leptastrea hertleini Durham, 1942:507, pi. 44, figs. 1, 4, 5. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12201. LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. Two specimens were found. The largest one, which is figured, is an entire colony whose diameter is 80 mm. The massive and flat- tened corallum resembles L. hertleini, known only from lower Eocene rocks in Media Agua Creek, Temblor Range, southwestern San Joaquin Valley, south central California. Family Caryophyllidae Gray, 1847a Genus Turbinolia Lamarck, 1816 Turbinolia dickersoni Nomland, 1916 Figures 6, 7 Turbinolia dickersoni Nomland, 1916:61, pi. 3, figs. 5-8. Palmer, 1923:30 6, pi. 56, figs. 2, 4. Quayle, 1932:98-99, pi. 6, figs. 1-5. Squires, 1984:15, figs. 5d, e; 1 991 :pl. 1, figs. 2-3. Squires and Deme- trion, 1992:24, figs. 45, 46. PRIMARY TYPE MATERIAL. UCMP holo- type 12020 (missing since 1932), Cerros Shale Mem- ber of the Lodo Formation, UCMP loc. 1817. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12202. MOLLUSCAN STAGE RANGE. Upper Paleo- cene through “Domengine.” GEOGRAPHIC RANGE. Eastern Laguna San Ig- nacio area, Baja California Sur, Mexico, to Little River area, Washington. LOCAL OCCURRENCE. CSUN loc. 1557. REMARKS. Only a single specimen was found and it is the first record of this minute species from the Crescent Formation and from Washington. Pre- viously, the northern limit of this species was Vaca- ville, northern California (Palmer, 1923). Genus Trochocyathus Milne-Edwards and Fiaime, 1848 Trochocyathus crooki Merriam and Turner, 1937 Figure 8 Trochocyathus crooki Merriam and Turner, 1937: 106, pi. 5, figs. 7-9. Bentson, 1943:293, pi. 50, figs. 12-14. PRIMARY TYPE MATERIAL. UCMP holo- type 33697; UCMP paratypes 3396, 3398; all from UCMP loc. A-1315. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12203. MOLLUSCAN STAGE RANGE. “Capay.” GEOGRAPHIC RANGE. West side of Sacra- mento Valley, northern California to Little River area, Washington. LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. Specimens are common, mostly complete, and range in height from 6 to 16 mm. This is the first report of this solitary coral from the Crescent Formation and from Washington. Pre- viously, the northern limit of this species was the Guinda area, northern California (Merriam and Turner, 1937; Bentson, 1943). Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 9 Family Flabellidae Bourne, 1905 Genus Flabellum Lesson, 1831 Flabellum californicum Vaughan, 1900 Figures 9, 10 Flabellum californicum Vaughan, 1900:69, pi. 4, figs. 11-12. Nomland, 1916:60, pi. 4, figs. 14- 16. Durham, 1943:196. ^Trochocyathus nomlandi Bentson. Squires et al., 1992:pl. 1, figs. 1-2. Not Trochocyathus nom- landi Bentson, 1943. PRIMARY TYPE MATERIAL. Specimens at USNM, type locality is 4 km (2.5 mi.) northeast of Clayton, Contra Costa County, northern California. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12204. MOLLUSCAN STAGE RANGE. “Capay” through “Domengine.” GEOGRAPHIC RANGE. Contra Costa County, northern California to Pulali Point, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1555. REMARKS. This distinctly cuneiform species is common at locality 1553 and rare at locality 1555. Specimens are mostly complete and approximately 9 mm in height. Durham (1943) assigned the rocks at the type locality of this species to the “Domen- gine Stage.” Family Dendrophylliidae Gray, 1847a Genus Balanophyllia Wood, 1844 Balanophyllia variabilis Nomland, 1916 Figure 11 Balanophyllia variabilis Nomland, 1916:66, pi. 4, figs. 2-3, 4, 13. Balanophyllia variabilis Nomland var. Merriam and Turner, 1937:101, 106, pi. 5, figs. 12-13. PRIMARY TYPE MATERIAL. Specimens at UCMP, Mount Diablo region, Contra Costa Coun- ty, northern California, UCMP loc. 714. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12205. MOLLUSCAN STAGE RANGE. “Capay” through “Domengine.” GEOGRAPHIC RANGE. Northern California to Little River area, Washington. LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. Specimens are abundant and many are complete. There is a partial growth size series with specimens ranging from 13 to 23 mm in height. This is the first report of this species from the Crescent Formation and from Washington. Previ- ously, the northern limit of this species was the Guinda area, northern California (Nomland, 1916; Merriam and Turner, 1937). Merriam and Turner (1937) assigned the rocks at the type locality of this species to the “Domengine Stage.” Phylum Brachiopoda Dumeril, 1806 Class Inarticulata Huxley, 1869 Order Craniidina Waagen, 1885 Family Craniidae Menke, 1828 Genus Craniscus Dali, 1871 Craniscus edwilsoni, new species Figures 12, 13, 14 DIAGNOSIS. Brachial valve with widely spaced radial ribs, apex slightly posterior of center, a well- delinated tripartite interior with a minutely pustu- lose flattened margin. COMPARISON. Compared to other brachio- Figures 4-27. Little River area upper Crescent Formation, sponge, colonial coral, solitary corals, inarticulate bra- chiopod, articulate brachiopods, polychaete worm, scaphopod, and gastropods. CSUN loc. 1553 unless otherwise noted. All specimens coated with ammonium chloride. 4. Sponge. Aphrocallistes ? sp., x3.5, LACMIP hypotype 12200. 5. Colonial coral. }Leptastrea hertleini Durham, 1942, basal view of corallum, x0.9, LACMIP hypotype 12201. 6-11. Solitary corals. 6, 7. Turbinolia dickersoni Nomland, 1916, x7, LACMIP hypotype 12202, CSUN loc. 1557. 6. Dorsal view. 7. Lateral view. 8. Trochocyathus crooki Merriam and Turner, 1937, lateral view, x3.3, LACMIP hypotype 12203. 9, 10. Flabellum californicum Vaughan, 1900, x3.3, LACMIP hypotype 12204. 9. Lateral view. 10. End view. 11. Balanophyllia variabilisNomland, 1916, lateral view, x2, LACMIP hypotype 12205. 12-14. Inarticulate brachiopod. Craniscus edwilsoni, new species, x4.3, LACMIP holotype 12206. 12. Exterior view. 13. Interior view. 14. Side view. 15-17. Articulate brachiopods. 15, 16. Gryphus ? sp., pedicle valve, x 1.1, LACMIP hypotype 12212. 15. Interior view. 16. Exterior view. 17. Terebratulina unguicula weaveri Hertlein and Grant, 1944, dorsal view, x2.8, LACMIP hypotype 12213. 18-20. Polychaete worm. Rotularia ( Rotularia ) tejonense (Arnold, 1910). 18, 19. x5, LACMIP hypotype 12214. 18. Umbilical view (showing attachment scar). 19. Lateral view. 20. End view, x4, LACMIP hypotype 12215. 21. Scaphopod. Dentalium sp. lateral view, partial specimen, x7.3, LACMIP hypotype 12216. 22-27. Gastropods. 22, 23. Erginus vaderensis (Lindberg, 1979), internal mold, x4, LACMIP hypotype 12217. 22. Dorsal view. 23. Left-profile view. 24, 25. Patelloida ? sp. cf. P. vokesi (Hickman, 1980), internal mold, x2.9, LACMIP hypotype 12218. 24. Dorsal view. 25. Left-profile view. 26, 27. Patelloida ? sp., CSUN loc. 1555. 26. Rubber cast of external mold, dorsal view, x2.9, LACMIP hypotype 12219. 27. Internal mold, left-profile view, x2.8, LACMIP hypotype 12220. 10 ■ Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 11 pod genera, craniacean genera have undergone few major revisions in the past two centuries. Lee and Brunton (1986) recently reviewed and updated the genera in family Craniidae. Using their new diag- noses and descriptions of the genera, we can assign the Little River area craniacean species to genus Craniscus Dali, 1871. The type species of Cra- niscus, the Late Jurassic Crania tripartita Munster (in Goldfuss, 1840) from Germany, is unusual in that it has thin, wall-like septa in the interior of the brachial valve that divide the interior into three chambers. Other species of Craniscus have an in- ternal tripartite subdivision without the thin, wall- like septa. Craniscus edwilsoni, new species, was compared to the few known species of Craniscus and is most similar to certain specimens of C. skeatsi (Allan, 1940) illustrated by Archbold (1990:figs. 21, 2L, 2N). Craniscus skeatsi is known from upper Oligocene to lower Miocene rocks in southeastern Australia (Archbold, 1990). The new species differs from C. skeatsi in having a brachial valve with radial ribbing, a more distinct tripartite interior, more elevated and more diverging anterior adductor pedestals, and a pustulose limbus. Only a few craniacean species have been reported from the Paleogene of North America, and they are all unnamed species tentatively assigned to gen- era. Toulmin (1940) reported a single unnamed spe- cies of Crania ? from the lower Eocene Salt Moun- tain Limestone of Alabama, and Cooper (1988) reported two unnamed species of Crania ? from the predominantly middle Eocene Castle Hayne For- mation of North Carolina. Cooper (1988) also re- ported two unnamed species of Isocrania ? from the middle Eocene Santee Formation of South Car- olina and another unnamed species of Isocrania ? from the Oligocene Cooper Formation of South Carolina. The new species has close affinity to Crania ? sp. 1 Cooper (1988:5, figs. 4-6), from the Castle Hayne Formation of North Carolina, but differs in having a brachial valve with a more centrally located apex, fewer and much coarser radial ribs, and a more elongate anteromedian ridge. The new species also has close affinity to Isocrania ? sp. 2 Cooper (1988: 5, pi. 3, figs. 9, 10; pi. 6, figs. 2, 3), from the Oli- gocene Cooper Formation of South Carolina, but differs in having a less pointed apex, fewer radial ribs, a more prominent anteromedian ridge, and a better developed pustulose limbus. The new species differs from the other unnamed species of Cooper (1988) in having much more swollen anterior adductor pedestals, the presence of an anteromedian ridge, and a less ribbed exterior. Craniscus edwilsoni resembles Toulmin’s (1940: 228, pi. 28, figs. 1-3; Stenzel, 1942:card no. 122, figs. 1-3) Crania ? sp., from the lower Eocene Salt Mountain Limestone of Alabama, but differs in having a brachial valve with non-spinose radial ribs, a more prominent ridge around the interior of the valve, more pronounced posterior adductor ped- 12 ■ Contributions in Science, Number 444 estals, and apparently a more elongate anterome- dian ridge. The anteromedian ridge region in the Salt Mountain species is incomplete, but it appears to be less elongate and less raised than the new species. Lee and Brunton (1986) reported that genus Cra- nia Retzius, 1781, is restricted to the Upper Cre- taceous (Campanian-Maastrichtian) and that Iso- crania Jaekel, 1902, is restricted to the Upper Cretaceous to lower Tertiary (Campanian-Danian). The unnamed Eocene species of Crania ? discussed by Toulmin (1940) and the unnamed Paleogene species of Crania ? and Isocrania ? discussed by Cooper (1988), therefore, should be reevaluated systematically. Toulmin’s (1940) Crania ? sp. and Cooper’s (1988) Crania ? sp. 1 and Isocrania ? sp. 2 closely resemble Craniscus edwilsoni, new species, and should be assigned to genus Craniscus. Their species, along with Craniscus edwilsoni, however, differ from other species of Craniscus by possessing a pustu- lose limbus like that seen in the genera Crania and Isocrania. With more study, these particular spe- cies of Toulmin and Cooper and the new species may prove to be a distinct subgenus of Craniscus. DESCRIPTION. Brachial valve small (up to 12 mm wide), circular, flattened to moderately inflated. Raised apex slightly posterior of center. Surface marked by 10-20 thin, radial riblets separated by wide spaces. Radial ribs tend to be much finer along posterior margin. Interior marked on posterior mar- gin by moderately raised posterior adductor ped- estals. Anterior adductor pedestals near center of valve consisting of two short oblique elevations diverging posteriorly with V-shaped depression be- tween them. Anteromedian ridge [= median myo- phore] moderately raised and extending anteriorly from the anterior adductor pedestals to slightly an- terior of midvalve. Anterior adductor pedestals and anteromedian ridge divide valve into three sections or chambers. Inner margins of brachial valve flat- tened [= limbus] and pustulose. Pedicle valve not found. HOLOTYPE. 7 mm wide, 6.5 mm long, 1 mm high. PRIMARY TYPE MATERIAL. LACMIP ho- lotype, 12206 (illustrated), LACMIP paratypes 12207 to 12211; all from CSUN loc. 1553. TYPE LOCALITY. CSUN loc. 1553. MOLLUSCAN STAGE RANGE. “Capay.” GEOGRAPHIC RANGE. Larch Mountain and Rock Candy Mountain in the Black Hills area, and Little River area, Washington. LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. Specimens are common at the type locality, and most are unbroken single valves. A few brachial valves are fairly thick and smooth, apparently because of abrasion. Ongoing work by us has revealed rare specimens of the new species from the upper Crescent For- mation in the vicinity of Larch Mountain (CSUN loc. 1563) and in the vicinity of Rock Candy Moun- Squires and Goedert: Crescent Formation Eocene Fossils tain (CSUN loc. 1564), Black Hills area (Fig. 1), about 50 km southeast of the Little River area. According to Rowell (1965), Craniscus is a cos- mopolitan genus whose geologic range is Jurassic to Recent. Lee and Brunton (1986), however, re- stricted the geographic distribution of the genus to Europe, Australia, Japan, and the Indo-Pacific area. The specimens of Craniscus edwilsoni in the upper Crescent Formation in the Little River and Black Hills areas, Washington, are the first record of Craniscus from the Pacific coast of North Amer- ica and the first named species of this genus in the New World. ETYMOLOGY. The new species is named after Edward C. Wilson, Curator, Invertebrate Paleon- tology Section, Natural History Museum of Los Angeles County, who has made valuable contri- butions to the study of invertebrate paleontology. Class Articulata Huxley, 1869 Family Terebratulidae Gray, 1840 Genus Grypbus Megerle von Miihlfeld, 1811 Grypbus ? sp. Figures 15, 16 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12212. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554. REMARKS. Specimens are common at locality 1553 but are rare at locality 1554. All specimens are single valves and at locality 1553 they are up to 40 mm in length. Preservation is too poor to allow generic identification. Family Cancellothyrididae Thomson, 1926 Genus Terebratulina d’Orbigny, 1847 Terebratulina unguicula weaveri Hertlein and Grant, 1944 Figure 17 Terebratulina unguicula Carpenter var. B. Weaver, 1943:15, pi. 1, fig. 11. Terebratulina unguicula weaveri Hertlein and Grant, 1944:81-83, pi. 6, figs. 4-5, 8-9, text fig. 20. Squires et al., 1992:pl. 1, figs. 6-8. PRIMARY TYPE MATERIAL. CAS holotype 7303, upper Crescent Formation, Jefferson County, Washington, UW loc. 353. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12213. MOLLUSCAN STAGE RANGE. “Capay” to ap- proximately the “Capay”/“Domengine” boundary. GEOGRAPHIC RANGE. Black Hills area, Little River area, Pulali Point, and Quilcene, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554, 1555, 1559. REMARKS. Specimens are abundant at locality 1553 where they range in size from 6 to 20 mm in length. Only one specimen from this locality is ar- ticulated. A few fragmental specimens were found at locality 1554 and specimens are rare at the other localities where fragments were found also. Squires (1992) discussed the age of the type lo- cality of this species and assigned it to near the “Capay Stage”/“Domengine Stage” boundary. Phylum Annelida Lamarck, 1809 Class Polychaeta Grube, 1850 Order Sabellida Malmgren, 1867 Family Serpulidae Johnston, 1865 Genus Rotularia Defrance, 1827 Subgenus Rotularia s.s. Rotularia ( Rotularia ) tejonense (Arnold, 1910) Figures 18, 19, 20 Spiroglyphus ? tejonense Arnold, 1910:51, pi. 4, fig. 18. Dickerson, 1916:pl. 37, figs. 5a-b. Vokes, 1939:162-1 63, pi. 20, figs. 20-22. Stewart, 1946: pi. 11, fig. 21. Spiroglyphus ? capayensis Merriam and Turner, 1937:106, pi. 5, figs. 10, 11. Sprioglyphus tejonensis Arnold. Hanna and Hert- lein, 1941:figs. 62-4, 62-20. ?T ubulostium capayense (Merriam and Turner). Keen and Bentson, 1944:195. ?T ubulostium tejonense (Arnold). Keen and Bent- son, 1944:195. Rotularia tejonense (Arnold). Nilsen, 1973:table 1. Squires, 1977:table 1; 1984:15, fig. 5g. Rotularia ( Rotularia ) tejonense (Arnold). Squires, 1987:19, 21, figs. 7, 8; 1988a:8, figs. 5, 6. Squires et al., 1992:pl. 1, figs. 12, 13. PRIMARY TYPE MATERIAL. USNM holo- type 165658 of Sprioglyphus ? tejonense, Avenal Sandstone, Kings County, California, USGS loc. 4617. UCMP holotype 33700 and UCMP paratype 33699 of Sprioglyphus ? capayensis, Capay For- mation, Capay Valley, California, UCMP loc. A-1313. ILLUSTRATED SPECIMENS. LACMIP hypo- types 12214-12215. MOLLUSCAN STAGE RANGE. “Capay” through “Transition.” GEOGRAPHIC RANGE. Simi Valley, southern California to Pulali Point, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1553a, 1554, 1556, 1560. REMARKS. A few specimens were found at lo- cality 1553. At the other localities, specimens are rare. Most specimens are coiled, but a few are fairly long, straight fragments. One specimen, which is Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 13 figured, was attached to a small pebble, and the attachment scar is very prominent. As noted by Vokes (1939), the holotype of R. (R.) tejonense is not representative of the ordinary forms in this very variable species. We agree with Voices’ (1939) comment that “the ordinary forms are inclined to be hexagonal in section, with a prominent sulcation on either side of a prominent median ridge on both top and bottom of the whorl, and with two minor sulcations on the outer edge.” Merriam and Turner (1937) reported Spriogly- phus ? capayensis Merriam and Turner from the lower Eocene of northern California and south- western Oregon, as well as from the middle Eocene of central California and southwestern Oregon. Their species is herein regarded as conspecific with R. {R.) tejonense. Phylum Mollusca Linne, 1758 Class Scaphopoda Broun (in Bronn and Keferstein, 1862) Order Dentalioida Palmer, 1974 Family Dentaliidae Gray, 1834 Genus Dentalium Linne, 1758 Dentalium sp. Figure 21 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12216. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554, 1559. REMARKS. A few specimens were found at lo- cality 1553. At the other localities specimens are rare. Specimens are small-sized fragments. More complete specimens are needed to identify the spe- cies. Class Gastropoda Cuvier, 1797 Order Patellogastropoda Lindberg, 1986 Family Lottiidae Gray, 1840 Genus Erginus Jeffreys, 1877 Erginus vaderensis (Lindberg, 1979) Figures 22, 23 Acmaea simplex Dickerson, 1917:181, pi. 31, figs, lla-b. [Not Acmaea cubensis simplex Pilsbry, 1891:42.] Acmaea clarki Van Winkle. Effinger, 1938:374. Weaver, 1943:282. [Not Acmaea clarki Van Winkle, 1918:85, pi. 6, fig. 4.] Acmaea vaderensis Lindberg, 1979:223 [nom. nov.]. Erginus vaderensis (Lindberg). Lindberg, 1988:5, pi. 1E-G. PRIMARY TYPE MATERIAL. CAS holotype 14 ■ Contributions in Science, Number 444 438, “Gries Ranch beds” of the Lincoln Creek For- mation, Lewis County, Washington, CAS loc. 181. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12217. MOLLUSCAN STAGE RANGE. “Capay” through “Tejon.” GEOGRAPHIC RANGE. Vader, Porter Creek (Grays Harbor County), Discovery Bay (Jefferson County), and Little River area, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554, 1555, 1557. REMARKS. A few specimens were found at lo- calities 1553 and 1555. At the other localities, spec- imens are rare. Preservation is mostly as internal molds, but a few specimens have some shell. The shell profile is high with the apex at one-third of length from the anterior end. The shell is smooth except for minute growth lines. The Little River specimens are late juveniles about 5 mm in length. According to Lindberg (1988), shell length in this species can be up to about 9 mm. The Little River area specimens of Erginus vade- rensis are the earliest record of both this genus and species and the first record of them from the Cres- cent Formation. Previously, E. vaderensis has been reported only from Washington and only from the upper Eocene part of the Lincoln Creek Formation (including the “Gries Ranch beds”) and the Quim- per Sandstone (Lindberg, 1988). Genus Patelloida Quoy and Gaimard, 1834 Patelloida ? sp. cf. P. vokesi (Hickman, 1980) Figures 24, 25 Acmaea vokesi Hickman, 1980:14-15, pi. 1, figs. 1-4. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12218. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554, 1555, 1560. REMARKS. A few specimens were found at lo- cality 1553. At the other localities, specimens are rare. Specimens are mostly molds. They closely re- semble Patelloida vokesi (Hickman, 1980) from the upper Eocene Keasey Formation in northwestern Oregon. Preservation is too poor to allow positive identification of the Little River specimens. If these eventually do prove to be Patelloida, they would be the earliest record of this genus from the Pacific coast of North America. The earliest undoubted report of this genus from this area are two species from middle Eocene rocks in the Tehachapi Moun- tains, southern California (Lindberg and Squires, 1990). Patelloida ? sp. Figures 26, 27 ILLUSTRATED SPECIMENS. LACMIP hypo- type 12219-12220. Squires and Goedert: Crescent Formation Eocene Fossils LOCAL OCCURRENCE. CSUN Iocs. 1555, 1557, 1560. REMARKS. Specimens are rare and are molds. The low profile and radial sculpture evidence sug- gest assignment to the genus Patelloida, but pres- ervation is too poor to allow positive identification. Patelloida ? sp. has many more and finer ribs than Patelloida ? sp. cf. P. vokesi, also from the Little River area. As mentioned above, the earliest un- doubted report of Patelloida from the Pacific coast of North America is middle Eocene from southern California. Order Vetigastropoda Salvini-Plawen, 1980 Family Fissurellidae Fleming, 1822 Genus Diodora Gray, 1821 Dio dor a sp. Figure 28 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12221. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554, 1555. REMARKS. A few specimens were found at lo- cality 1553. At the other localities, specimens are rare. Specimens are either internal molds or inte- riors of shells. They show evidence of a truncate internal callus, which is the main diagnostic feature of genus Diodora. Preservation is too poor to allow identification as to species. A few specimens show evidence of reticulate ornamentation that resem- bles Diodora stillwaterensis (Weaver and Palmer, 1922:27, pi. 11, figs. 3, 6) known from the upper middle Eocene Cowlitz Formation, southwestern Washington (Weaver, 1943), and possibly from the middle Eocene (“Transition Stage”) Friars Forma- tion, San Diego County, southern California (Squires and Demere, 1991). Diodora stillwaterensis is the only previously reported Diodora from the Eocene of the Pacific coast of North America. The Little River area specimens of Diodora are the earliest occurrence of this genus on the Pacific coast of North America and the first occurrence of this ge- nus from the Crescent Formation. Paleocene and early Eocene reports of this genus from elsewhere in the United States are confined to Maryland, Al- abama, and Mississippi (Palmer and Brann, 1966). Family Turbinidae Rafinesque, 1815 Genus Homalopoma Carpenter, 1864 Homalopoma sp. cf. H. umpquaensis domenginensis Vokes, 1939 Figures 29, 30 Homalopoma umpquaensis domenginensis Vokes, 1939:179-180, pi. 22, fig. 27. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12222. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554. REMARKS. Specimens are abundant at locality 1553 and rare at locality 1554. Nearly all the spec- imens are fragments. Preservation is too poor to allow positive identification as to species. Although the illustrated specimen is worn, the nodes on the body whorl spirals are fairly prominent. This is the first report of genus Homalopoma from the Crescent Formation. If the Little River area specimens eventually prove to be H. umpquaensis domenginensis Vokes, 1939, they would be the first occurrence of this subspecies in Washington and in the Crescent Formation. The northern limit of this subspecies is presently central California (Squires, 1988a). Homalopoma sp. afF. H. wattsi (Dickerson, 1916) Figures 31, 32 Monodonta wattsi Dickerson, 1916:494, pi. 40, figs. 3a, 3b. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12223. LOCAL OCCURRENCE. CSUN loc. 1555. REMARKS. Specimens are rare. They resemble Homalopoma wattsi (Dickerson, 1916), reported (Weaver, 1943) from lower Eocene strata in Cali- fornia and southwestern Washington. The Crescent Formation specimen, like H. wattsi, has many spiral ribs and two nodes on the inner lip. The Crescent Formation specimens differ from H. wattsi in the following features: somewhat finer spiral ribs and no tendency for those in the middle part of the body whorl to be coarser than the others. The Crescent Formation specimens of Homa- lopoma sp. aff. H. wattsi are very similar to spec- imens of H. aff. H. wattsi reported (Squires, 1991: pi. 1, fig. 9) from “Capay Stage” strata in the Oro- copia Mountains, southern California. Order Caenogastropoda Cox, 1960 Family Turritellidae Woodward, 1851 Genus Turrit ell a Lamarck, 1799 Turritella andersoni s.s. Dickerson, 1916 Figure 33 Turritella andersoni Dickerson, 1916:501-502, pi. 42, fig. 9a, b. Turner, 1938:83, pi. 22, figs. 4-6. Merriam, 1941:76-77, pi. 9, figs. 1-2; pi. 10, figs. I, 3-5, 8; pi. 12, figs. 1-3. Hanna and Hertlein, 1941:figs. 62-6, 62-7, 62-16. Givens, 1974:62, pi. 5, figs. 7-10. Squires, 1984:16-17, fig. 6e; 1987: 24-25, figs. 20-21; 1988a:10, fig. 14; 1988b:205, fig. 2; 1988c:pl. 1, fig. 2; 1 991 :pl. 1, fig. 12. Squires and Demetrion, 1992:26, fig. 58. Turritella andersoni susanae Merriam, 1941:79, pi. II, fig. 6. Saul, 1983:pl. 2, fig. 5. Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 15 PRIMARY TYPE MATERIAL. UCMP holo- type 12131, UCMP paratype 12132, both missing since 1959, Cerros Shale Member of Lodo For- mation, Fresno County, California, UCMP loc. 1817; UCMP neotype 38193 (designated by Squires, 1988c), UCMP loc. 1817. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12224. MOLLUSCAN STAGE RANGE. “Capay.” GEOGRAPHIC RANGE. Eastern Laguna San Ig- nacio area, Baja California Sur, Mexico, to Little River area, Washington. LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. Specimens are abundant, well-pre- served small fragments showing the diagnostic whorl profile that is broadly concave medially between a pair of broadly spaced primary spiral ribs. This is the first report of this species from Wash- ington. Previously, T. andersoni had been reported as far north as southwestern Oregon (Turner, 1938; Squires and Demetrion, 1992). Weaver (1937) re- ported this species from the Metchosin Volcanics near Victoria, British Columbia, in rocks that have been correlated with the Crescent Formation (Mul- ler, 1980), but the specimens belong to a different species (Squires, 1988b). Family Siliquariidae Anton, 1838 Genus T enagodus Guettard, 1770 Tenagodus bajaensis Squires, 1990b Figures 34, 35 Tenagodus bajaensis Squires, 1990b:298, figs. 6-8. Squires and Demetrion, 1992:27-28, fig. 64. PRIMARY TYPE MATERIAL. IGM holotype 5102, IGM paratype 5103, Bateque Formation, eastern Laguna San Ignacio area, Baja California Sur, Mexico, CSUN loc. 1291a. ILLUSTRATED SPECIMENS. LACMIP hypo- type 12225-12226. MOLLUSCAN STAGE RANGE. “Capay.” GEOGRAPHIC RANGE. Eastern Laguna San Ig- nacio area, Baja California Sur, Mexico, to Little River area, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1553a. REMARKS. Specimens are rare and the shell ma- terial easily falls off leaving only internal molds. Most specimens show the slit that is so character- istic of this genus. This is the first report of this species from the Crescent Formation and from Washington. Previ- ously, it had only been found in the “Capay Stage” part of the Bateque Formation, eastern Laguna San Ignacio area, Baja California Sur, Mexico (Squires, 1990b: Squires and Demetrion, 1992). Family Calyptraeidae Blainville, 1824 Genus Calyptraea Lamarck, 1799 Calyptraea diegoana (Conrad, 1855) Figure 36 Trochita diegoana Conrad, 1855:7, 17; 1857:327, pi. 5, fig. 42. Galerus excentricus Gabb, 1964:136, pi. 20, fig. 95; pi. 29, fig. 232a. Dickerson, 1913:264. Calyptraea calabasaensis Nelson, 1925:419, pi. 54, figs. 8a-b. Calyptraea ( Galerus ) calabasaensis Nelson. Clark and Woodford, 1927:120, pi. 21, figs. 10-13. Calyptraea diegoana (Conrad). Stewart, 1927:340- 341, pi. 27, fig. 15. Turner, 1938:89-90, pi. 20, figs. 1-2. Effinger, 1938:378. Weaver, 1943:351- 352, pi. 71, figs. 16, 20; pi. 103, fig. 3; 1953:29. Stewart, 1946:pl. 11, fig. 5. Kleinpell and Weaver, 1963:186, pi. 24, fig. 7. Hickman, 1969:79, 82, pi. 11, figs. 7-8; 1980:33-34, pi. 2, figs. 18-21. Smith, 1975:469, table 2. Givens and Kennedy, Figures 28-48. Little River area upper Crescent Formation, gastropods and bivalves. CSUN loc. 1553 unless otherwise noted. All specimens coated with ammonium chloride. 28-43. Gastropods. 28. Diodora sp., internal mold, dorsal view, x5.9, LACMIP hypotype 12221. 29, 30. Homalopoma sp. cf. H. umpquaensis domenginensis Vokes, 1939, x4.4, LACMIP hypotype 12222, CSUN loc. 1555. 29. Apertural view. 30. Abapertural view. 31, 32. Homalopoma sp. aff. H. wattsi (Dickerson, 1916), x4.9, LACMIP hypotype 12223. 31. Apertural view. 32. Oblique apertural view. 33. Turritella andersoni Dickerson, 1916, partial specimen, apertural view, x8.5, LACMIP hypotype 12224. 34, 35. Ten- agodus bajaensis Squires, 1990, internal molds. 34. Apertural view, x2, LACMIP hypotype 12225. 35. Oblique view of longitudinal slit on body whorl, x4, LACMIP hypotype 12226. 36. Calyptraea diegoana (Conrad, 1855), lateral view, x4.2, LACMIP hypotype 12227, CSUN loc. 1555. 37, 38. Eratotrivia crescentensis (Weaver and Palmer, 1922), x6.9, LACMIP hypotype 12228. 37. Abapertural view. 38. Abapertural view. 39. Colwellia sp., apertural view, x5, LACMIP hypotype 12229. 40. Dentimitra cretacea (Gabb, 1864), apertural view, anterior end missing, x5, LACMIP hypotype 12230, CSUN loc. 1555. 41. Lyria andersoni Waring, 1917, apertural view, x5.7, LACMIP hypotype 12231. 42. Surculites mathewsonii (Gabb, 1864), lateral view, x4, LACMIP hypotype 12232, CSUN loc. 1560. 43. Conus sp., abapertural view, x3.4, LACMIP hypotype 12233. 44-48. Bivalves. 44. Acila ( Truncacila ) decisa (Conrad, 1855), left valve, x3.7, LACMIP hypotype 12234. 45. Barbatia sp. aff. B. (B.) landesi (Weaver and Palmer, 1922), internal mold, left valve, x2.5, LACMIP hypotype 12235. 46, 47. Glycymeris sp. cf. G. crescentensis Weaver and Palmer, 1922. 46. Left? valve, x5, LACMIP hypotype 12236. 47. Left-valve interior, x2.1, LACMIP hypotype 12237. 48. Glycymeris (< Glycymerita ) sagittata (Gabb, 1864), right? valve, xl.8, LACMIP hypotype 12238. 16 ■ Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 17 1979:table 2. Demere, Sundberg, and Schram, 1979:pl. 2, fig. 7. Squires, 1984:21, fig. 6q; 1987: 32, fig. 4; 1988a:ll, fig. 19; 1991:pl. 1, fig. 20. PRIMARY TYPE MATERIAL. USNM holo- type 1856, Tejon? horizon, San Diego, California. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12227. MOLLUSCAN STAGE RANGE. “Martinez” (upper Paleocene) through lower Oligocene. GEOGRAPHIC RANGE. San Diego, California, to Little River area, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1555. REMARKS. Specimens are uncommon at both localities. This is the first confirmed report of this species in the Crescent Formation. Although Ar- nold and Hannibal (1913:572) listed this species in a faunal list of macrofossils from the Crescent For- mation in the Crescent Bay area, Washington, their specimens were not available for study. Family Triviidae Troschel, 1863 Genus Eratotrivia Sacco, 1894 Eratotrivia crescentensis (Weaver and Palmer, 1922) Figures 37, 38 Eratopsis crescentensis Weaver and Palmer, 1922: 36-37, pi. 11, figs. 10, 22. Weaver, 1943:393, pi. 76, fig! 21. Eratotrivia crescentensis (Weaver and Palmer). Schilder and Schilder, 1971:12. PRIMARY TYPE MATERIAL. UW holotype 201, Crescent Formation at Crescent Bay, Clallam County, Washington, UW loc. 358. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12228. MOLLUSCAN STAGE RANGE. “Capay.” GEOGRAPHIC RANGE. Little River area and Crescent Bay, Washington. LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. Specimens are uncommon at local- ity 1553. Preservation is good, and the surface or- namentation is well delineated. Previously, this spe- cies was known only from the Crescent Bay area, Washington (Weaver, 1943). Family Nassariidae Iredale, 1916 Genus Colwellia Nuttall and Cooper, 1973 Colwellia sp. Figure 39 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12229. ' LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. Only a single, poorly preserved specimen was found. Preservation is too poor to allow identification to species. The specimen re- sembles Colwellia antiquata (Gabb, 1864:97, pi. 18 ■ Contributions in Science, Number 444 18, fig. 50) known from lower to upper middle Eocene strata in southern California and south- western Oregon (Allmon, 1990). The Little River area specimen of Colwellia is the first record of this genus from the Crescent Formation. Family Mitridae Swainson, 1831 Genus Dentimitra Koenen, 1890 Dentimitra cretacea (Gabb, 1864) Figure 40 Mitra cretacea Gabb, 1864:103, pi. 28, fig. 215. Stewart, 1927:406, pi. 27, figs. 9-10. Mitra cretacea Gabb var. Turner, 1938:74, pi. 15, fig. 9. Weaver, 1943:497, pi. 95, figs. 5, 6. Uromitra (?) cretacea (Gabb). Vokes, 1939:134-135, pi. 18, fig. 19. Proximitra ? cretacea (Gabb). Givens, 1974:87. Squires, 1977:table 1; 1984:34, fig. 8p. Dentimitra cretacea (Gabb). Cernohorsky, 1976: 510. PRIMARY TYPE MATERIAL. ANSP holotype 4302, Tejon Formation s.L, Martinez, California. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12230. MOLLUSCAN STAGE RANGE. “Capay” through “Domengine.” GEOGRAPHIC RANGE. Simi Valley, southern California, to Little River area, Washington. LOCAL OCCURRENCE. CSUN loc. 1555. REMARKS. Only a single specimen was found. It is well preserved except for missing the anterior end of the shell. This specimen is unworn and shows the sculpture better than any other illustrated spec- imen of this species. This is the first report of this species from the Crescent Formation and from Washington. Previously, the northern limit of this species was southwestern Oregon (Turner, 1938). Family Volutidae Rafinesque, 1815 Genus Lyria Gray, 1847b Lyria andersoni Waring, 1917 Figure 41 Cancellaria irelaniana Cooper. Arnold, 1910:52, pi. 4, fig. 22. Not Cancellaria irelaniana Cooper, 1894. Lyria andersoni Waring, 1917:97, pi. 15, fig. 12. Clark, 1929:pl. 9, figs. 7-8. Clark and Vokes, 1936:876, pi. 1, fig. 17. Turner, 1938:73, pi. 18, fig. 5. Vokes, 1939:136, pi. 18, figs. 22, 24. Hanna and Hertlein, 1941:170, fig. 62-21. Squires, 1984: 35, pi. 9d; 1987:47, fig. 69; 1988a:16, fig. 36; 1988c:pl. 1, fig. 16; 1 991 :pl. 2, fig. 8. Squires and Demetrion, 1992:33, fig. 90. PRIMARY TYPE MATERIAL. SU holotype 195, SU paratype 196, Llajas Formation, Simi Valley, Ventura County, California, SU loc. 2696. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12231. MOLLUSCAN STAGE RANGE. “Capay” through “Domengine.” Squires and Goedert: Crescent Formation Eocene Fossils GEOGRAPHIC RANGE. Eastern Laguna San Ig- nacio area, Baja California Sur, Mexico, to Little River area, Washington. LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. Specimens are rare. This is the first report of this species from the Crescent Formation and from Washington. Previously, the northern limit of this species was southwestern Oregon (Turner, 1938). Family Turridae Swainson, 1840 Genus Surculites Conrad, 1865 Genus Surculites matbewsonii (Gabb, 1864) Figure 42 Fusus matbewsonii Gabb, 1864:83, pi. 18, fig. 33. Dickerson, 1914:pl. 16, fig. 2. Bela clathra Gabb, 1869:152, pi. 26, fig. 31. }Pleurotoma decipiens Cooper, 1894:40, pi. 2, fig. 32. Potamides ? davisiana Cooper, 1894:44, pi. 1, fig. 13. Surcula davisiana (Cooper). Dickerson, 1913:279, pi. 12, figs. 6a-b. SurculaQ) sp. Waring, 1 91 7:pl. 15, fig. 16. Surcula decipiens (Cooper). Hanna, 1927:324, pi. 54, figs. 6, 8. Surculites matbewsonii (Gabb). Stewart, 1927:420- 421, pi. 26, figs. 12-14. Clark, 1929:pl. 9, figs. 3- 4. Turner, 1938:69-70, pi. 17, figs. 6, 10. Vokes, 1939:123, pi. 17, figs. 8, 19. Weaver, 1943:526, pi. 97, figs. 24, 29; pi. 98, figs. 1, 5; 1953:29. Givens, 1974:90, pi. 11, figs. 5, 7. Zinsmeister, 1974:164, pi. 17, fig. 6; 1983:table 1. Squires, 1977:table 1; 1984:37, fig. 9j; 1987:48, fig. 73; 1 991 :pl. 2, fig. 11. Givens and Kennedy, 1979: 87, tables 1, 3. PRIMARY TYPE MATERIAL. ANSP lectotype 4180, designated by Stewart (1927), Tejon For- mation s.L, near Martinez, northern California. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12232. MOLLUSCAN STAGE RANGE. “Martinez” (upper Paleocene) through “Transition.” GEOGRAPHIC RANGE. San Diego, California, to Little River area, Washington. LOCAL OCCURRENCE. CSUN loc. 1560. REMARKS. Specimens are rare. This is the first report of this species from the Crescent Formation and from Washington. Previously, the northern limit of this species was southwestern Oregon (Turner, 1938). Family Conidae Rafinesque, 1815 Genus Conus Linne, 1758 Conus sp. Figure 43 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12233. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1555. REMARKS. Specimens are uncommon and too poorly preserved to allow identification as to spe- cies. The specimens are prominently noded. This is the first report of this genus from the Crescent Formation. Class Bivalvia Linne, 1758 Order Nuculoida Dali, 1889 Family Nuculidae Gray, 1824 Genus Acila H. and A. Adams, 1858 Subgenus Truncacila Schenck, in Grant and Gale, 1931 Acila ( Truncacila ) decisa (Conrad, 1855) Figure 44 Nucula decisa Conrad, 1855:11-12; 1857:pl. 3, fig. 19. Acila gabbiana Dickerson, 1916:481, pi. 36, fig. 1. Anderson and Hanna, 1925:176, pi. 9, fig. 12. Nucula {Acila) stillwaterensis Weaver and Palmer, 1922:6, pi. 8, fig. 8. Acila lajollaensis Hanna, 1927:270, pi. 25, figs. 1, 3, 5, 7-8, 12, 15. Acila ( Truncacila ) decisa (Conrad). Schenck, 1936: 53-56, pi. 3, figs. 1-9, 11-15; pi. 4, figs. 1-2; text fig. 7 (22, 23, 25). Turner, 1938:41-42, pi. 5, figs. 2-3. Vokes, 1939:41, pi. 1, figs. 7-8. Weaver, 1943:22-23, pi. 6, figs. 1, 4, 8; pi. 7, figs. 8-9. Moore, 1968:30, pi. 13a; 1983:10, pi. 1, fig. 14. Givens, 1974:38, pi. 1, fig. 1. Zinsmeister, 1974: 67-68, pi. 6, fig. 3; 1983:table 1. Squires, 1977: table 1; 1984:41, fig. 10a; 1987:54, fig. 86; 1988a: 17, fig. 44; 1 991 :pl. 2, fig. 15. Throckmorton, 1988:pl. 1, fig. 19. PRIMARY TYPE MATERIAL. UCMP neotype 31132, designated by Schenck (1936), Ardath Shale, UCMP loc. 5062. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12234. MOLLUSCAN STAGE RANGE. “Martinez” through upper Eocene ( Turritella scbencki dela- guerrae Zone of Kleinpell and Weaver, 1963). GEOGRAPHIC DISTRIBUTION. San Diego, California, through Kamchatka, Russia. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554. REMARKS. Specimens are rare at both localities and consist of broken single valves. This is the first report of this species from the Crescent Formation. Previously, it had been reported from the Cowlitz Formation in Washington (Weaver, 1943). Order Arcoida Stoliczka, 1871 Family Arcidae Lamarck, 1809 Genus Barbatia Gray, 1842 Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 19 Subgenus Barbatia s.s. Barbatia sp. aff. B. (B.) landesi (Weaver and Palmer, 1922) Figure 45 Area ( Barbatia ) landesi Weaver and Palmer, 1922: 9-10, pi. 8, fig. 4. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12235. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1553a, 1555, 1557, 1558. REMARKS. Specimens are common at locality 1553 and rare at the other localities. Specimens are fragments of single valves resembling Barbatia ( B .) landesi (Weaver and Palmer, 1922) from the upper middle Eocene Cowlitz Formation, southwestern Washington. The Crescent Formation specimens, like B. (B.) landesi, have ribs that are not coarsely beaded and that do not dichotomize. The Crescent Formation specimens differ from B. (B.) landesi in the following features: dorsal margin slopes down just anterior to the umbones (rather than projecting a moderate distance) and no tendency for second- ary ribs in the interspaces. This is the first report of genus Barbatia from the Crescent Formation. Family Glycymerididae Newton, 1922 Genus Glycymeris da Costa, 1778 Glycymeris sp. cf. G. crescentensis Weaver and Palmer, 1922 Figures 46, 47 Glycymeris crescentensis Weaver and Palmer, 1922: 11, pi. 8, figs. 10, 12. ILLUSTRATED SPECIMENS. LACMIP hypo- type 12236-12237. LOCAL OCCURRENCE. CSUN iocs. 1553, 1557. REMARKS. Specimens are abundant at locality 1557 and common at locality 1553. Preservation is too poor to allow positive species identification. The specimens are assigned tentatively to G. cres- centensis Weaver and Palmer, 1922, known from lower Eocene strata in southwestern Oregon and from the Crescent Formation at Crescent Bay, Washington (Turner, 1938; Weaver, 1943). Subgenus Glycymerita Finlay and Marwick, 1937 Glycymeris ( Glycymerita ) sagittata (Gabb, 1864) Figure 48 Axinaea ( Limopsis ?) sagittata Gabb, 1864:197-198, pi. 31, figs. 267-267a. 20 ■ Contributions in Science, Number 444 Glycimeris hannibali Dickerson, 1916:483, pi. 36, figs. 8a-b. Glycymeris sagittatus (Gabb). Dickerson, 1 91 6:pl. 36, figs. 5a-b. Glycymeris sagittata (Gabb). Anderson and Hanna, 1925:181-182, pi. 1, fig. 6. Stewart, 1930:71-73, pi. 12, fig. 10; 1946:pl. 12, fig. 3. Turner, 1938: 43-44, pi. 6, figs. 1-3 [as “ Glycimeris"]. Vokes, 1939:45-46, pi. 1, figs. 18-20. Weaver, 1943:54- 55, pi. 9, figs. 17-18; pi. 11, fig. 15. Kleinpell and Weaver, 1963:196-197, pi. 28, fig. 10; pi. 29, figs. 1-2. Demere, Sundberg, and Schram, 1979:pl. 1, figs. 1-2. Glycymeris ( Glycymerita ) sagittata (Gabb). Givens, 1974:42-43. Squires, 1977:table 1; 1984:42, fig. lOd; 1987:56, fig. 91; 1988a:17-18, fig. 46. Dev- yatilova and Volobueva, 1981:37, pi. 11, fig. 4. Moore, 1983:54-55, pi. 12, fig. 17. Marincovich, 1988:269, pi. 1, fig. 3. Squires and Demetrion, 1992:36-37, figs. 100, 101. PRIMARY TYPE MATERIAL. ANSP lectotype 4422, designated by Stewart (1930:72). Tejon For- mation near Ft. Tejon (NVi of section 29, T 10 N, R 19 W, Kern County, California. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12238. MOLLUSCAN STAGE RANGE. “Capay” through “Tejon,” Oligocene?. GEOGRAPHIC RANGE. Eastern Laguna San Ig- nacio area, Baja California Sur, Mexico, through far eastern Russia. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554. REMARKS. Specimens are rare. This is the first report of this species from the Crescent Formation. Subgenus Tucetona Iredale, 1931 Glycymeris [Tucetona) fresnoensis Dickerson, 1916 Figure 49 Glycimeris fresnoensis Dickerson, 1916:483, pi. 36, fig. 7. Turner, 1938:44, pi. 5, fig. 9. Glycymeris fresnoensis Dickerson. Vokes, 1939:48, pi. 1, fig. 17. Weaver, 1943:pl. 11, fig. 11. Glycymeris ( Tucetona ) fresnoensis Dickerson. Moore, 1983:55, pi. 12, figs. 2, 3. PRIMARY TYPE MATERIAL. UCMP holo- type 11795, Cerros Shale Member of the Lodo Formation, Fresno County, California, UCMP loc. 1817. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12239. MOLLUSCAN STAGE RANGE. “Capay.” GEOGRAPHIC RANGE. Central California through Crescent Bay, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1557. REMARKS. Specimens are rare. This small spe- cies is characterized by 16 sharp radial ribs. Squires Squires and Goedert: Crescent Formation Eocene Fossils Figures 49-62. Little River area upper Crescent Formation, bivalves, “gooseneck” barnacle, and shark teeth. CSUN loc. 1553 unless otherwise noted. All specimens coated with ammonium chloride. 49-59. Bivalves. 49. Glycymeris (' Tucetona ) fresnoensis Dickerson, 1916, left? valve, x5.5, LACMIP hypotype 12239. 50. Brachidontes ( Brachidontes ) cowlitzensis (Weaver and Palmer, 1922), right valve, dorsal view, xl.9, LACMIP hypotype 12240. 51-53. Nayadina (. Exputens ) batequensis Squires, 1990. 51. Right valve, x3, LACMIP hypotype 12241. 52. Left valve, x2.1, hypotype LACMIP 12242. 53. Left-valve interior, x2.4, LACMIP hypotype 12243. 54. Parvamussium sp., left? valve, xl.5, LACMIP hypotype 12244, CSUN loc. 1555. 55. Spondylus carlosensis Anderson, 1905, left valve, x2.7, LACMIP hypotype 12245, CSUN loc. 1555. 56. “ Ostrea ” sp., left-valve interior, x 1.7, LACMIP hypotype 12246. 57. Venericardia sp., internal mold, right valve, xl.6, LACMIP hypotype 12247. 58. Nemocardium linteum (Conrad, 1855), internal mold, left? valve, x4.4, LACMIP hypotype 12248. 59. Corbula ( Caryocorbula ) dickersoni Weaver and Palmer, 1922, left valve, x4, LACMIP hypotype 12249. 60. Aporolepas sp., tergum, x3.6, LACMIP hypotype 12250, CSUN loc. 1554. 61. Paleohypotodus sp., anterior tooth, labial view, x2.7, LACMVP 134133, CSUN loc. 1553 = LACMVP loc. 6117. 62. Isurus ? sp., upper lateral? tooth, lingual view, x5.4, LACMVP 134434, CSUN loc. 1560a = LACMVP loc. 6123. (1988b) allocated UCMP locality 1817 to the “Ca- Order Mytiloida Ferussac, 1822 pay Stage.” Although Weaver (1943) listed this spe- cies from the Domengine Formation of central Cal- Family Mytilidae Rafinesque, 1815 ifornia, Vokes (1939) clearly stated that it is not found in that formation. Genus Brachidontes Swainson, 1840 Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 21 Subgenus Brachidontes s.s. Brachidontes ( Brachidontes ) cowlitzensis Weaver and Palmer, 1922 Figure 50 Modiola ornata Gabb, 1864:184-185, pi. 24, fig. 166. Not Mytilus ornatus d’Orbigny, 1843:283, pi. 342, figs. 10-12. Modiolus ornatus Gabb. Arnold, 1907:pl. 38, fig. 4. Modiolus ( Brachydontes ) cowlitzensis Weaver and Palmer, 1922:16-17, pi. 9, fig. 19 [ nom . nov.]. Brachydontes ornatus (Gabb). Anderson and Han- na, 1925:188, pi. 3, fig. 4. Modiolus ( Brachydontes ) ornatus Gabb. Clark and Woodford, 1927:89, pi. 14, fig. 10. Clark, 1929: pi. 3, fig. 6. Brachidontes cowlitzensis (Weaver and Palmer). Turner, 1938:45-46, pi. 6, figs. 7-8. Kleinpell and Weaver, 1963:197, pi. 29, fig. 3. Wolfe, 1977:3. Givens and Kennedy, 1979:table 2. Volsella ( Brachidontes ) cowlitzensis (Weaver and Palmer). Weaver, 1943:113-114, pi. 26, fig. 4. Brachidontes ( Brachidontes ) cowlitzensis (Weaver and Palmer). Givens, 1974:43. Squires, 1977:table 1; 1984:42, fig. lOe; 1987:56, fig. 92; 1988a:18, fig. 47. Moore, 1983:66-67, pi. 17, fig. 1. Throck- morton, 1988:pl. 1, fig. 7. Marincovich, 1988: 269-270, pi. 1, fig. 4. PRIMARY TYPE MATERIAL. ANSP lectotype 4450 of Modiola ornata Gabb, designated by Stew- art (1930), Domengine? Formation, Martinez, Cal- ifornia. CAS holotype 7406 of Modiolus ( Brach- ydontes:) cowlitzensis Weaver and Palmer, Cowlitz Formation, Lewis County, Washington, UW loc. 329. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12240. MOLLUSCAN STAGE RANGE. “Meganos” through lower Oligocene ( Turritella variata loren- zana Zone of Kleinpell and Weaver, 1963). GEOGRAPHIC RANGE. San Diego, California, through Gulf of Alaska. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1557. REMARKS. Specimens are rare. This is the first confirmed report of this species from the Crescent Formation. Arnold and Hannibal (1913) listed this species in a faunal list of macrofossils from the Crescent Formation in the Crescent Bay area, Washington. Order Pteroida Newell, 1965 Family Malleidae Lamarck, 1819 Genus Nayadina Munier-Chalmas, 1864 Subgenus Exputens Clark, 1934 22 ■ Contributions in Science, Number 444 Nayadina ( Exputens ) batequensis Squires, 1990a Figures 51, 52, 53 Nayadina ( Exputens ) batequensis Squires, 1990a: 308-309, figs. 3-25; 1992:133-134, figs. 1-6. Squires and Demetrion, 1992:37, fig. 103. PRIMARY TYPE MATERIAL. IGM holotype 5108, IGM paratypes 5109-5110, Bateque For- mation, eastern Laguna San Ignacio area, Baja Cal- ifornia Sur, Mexico, CSUN loc. 1220b. ILLUSTRATED SPECIMENS. LACMIP hypo- type 12241-12242. MOLLUSCAN STAGE RANGE. “Capay.” GEOGRAPHIC RANGE. Northwestern Jamai- ca; Baja California Sur, Mexico; and Little River area, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554, 1555, 1557. REMARKS. Specimens are abundant at localities 1553 and 1554 but are rare at the other two lo- calities. This is the first report of this species from the Crescent Formation and from Washington. Pre- viously, it had only been found in the lower to middle Eocene Chapelton Formation, northwest- ern Jamaica and in the middle lower Eocene (“Ca- pay Stage”) parts of the Tepetate and Bateque For- mations, Baja California Sur, Mexico (Squires, 1990a, 1992; Squires and Demetrion, 1992). Order Pterioida Newell, 1965 Family Propeamussiidae Abbott, 1954 Genus Parvamussium Sacco, 1897 Parvamussium sp. Figure 54 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12243. LOCAL OCCURRENCE. CSUN Iocs. 1553a, 1554, 1555. REMARKS. Specimens are uncommon at local- ity 1553a and rare at the other localities. The spec- imens are small (2-3 mm height) and show only the right-valve interior. There are 10 ribs that extend almost to the ventral margin. Parvamussium is characterized by internal ribs that extend to, or almost to, the ventral margin (Moore, 1984). There are three known Paleogene species of Parvamus- sium from the Pacific coast of North America and they all have 10-12 interior ribs (Moore, 1984). Due to the absence of any data on the exterior sculpture, identification to species is not possible for the Little River area specimens. Parvamussium sp. was previously reported from the Crescent For- mation at Pulali Point (Squires et al., 1992). Family Spondylidae Gray, 1826 Genus Spondylus Linne, 1758 Squires and Goedert: Crescent Formation Eocene Fossils Spondylus carlosensis Anderson, 1905 Figure 55 Spondylus carlosensis Anderson, 1905:194, pi. 13, fig. 1. Arnold, 1910:pl. 2, figs. 6-7. Dickerson, 1915:pl. 1, fig. 7. Anderson and Hanna, 1925: 189-190, text fig. 10. Vokes, 1939:57, pi. 3, figs. 10, 13. Kleinpell and Weaver, 1963:199, pi. 31, fig. 6. Squires, 1984:43, fig. lOj. PRIMARY TYPE MATERIAL. CAS holotype 56, Domengine Formation, west and north of Coa- linga, NW14 of section 35, T 20 S, R 14 E, Fresno County, California. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12245. MOLLUSCAN STAGE RANGE. “Capay” through “Domengine.” GEOGRAPHIC RANGE. Simi Valley, southern California, through Little River area, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1553a, 1554, 1555, 1556, 1557. REMARKS. Specimens are abundant at locality 1553, uncommon at 1554 and 1556, and rare else- where. All specimens are single valves and many of these are preserved as molds. The Little River area specimens of Spondylus carlosensis Anderson, 1905, are the earliest record of this species from the Pacific coast of North America and the first record from the Crescent For- mation and from Washington. Previously, it had been reported only from the “Domengine Stage” (Squires, 1984) and had been found as far north as northwestern Oregon (Baldwin, 1964). Family Ostreidae Rafinesque, 1815 Genus Ostrea Linne, 1758 Ostrea sp. Figure 56 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12246. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554, 1555, 1556. REMARKS. Specimens are abundant at locality 1553 and rare at the other localities. Most speci- mens are fragments. The illustrated specimen is a complete single valve that shows only the interior. Due to the absence of any data on the exterior sculpture, species identification is not possible. The specimen strongly resembles Ostrea sp. from the Crescent Formation at Pulali Point (Squires et al., 1992:pl. 1, figs. 30-31). Order Veneroida H. and A. Adams, 1856 Family Carditidae Fleming, 1828 Genus Venericardia Lamarck, 1801 Venericardia sp. Figure 57 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12247. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554. REMARKS. Specimens are common at locality 1553 and rare at 1554. Specimens are small frag- ments of single valves. Preservation is too poor to allow positive identification to species. The speci- mens, which have approximately 20 radial ribs, are quadrate and resemble the illustrated specimen (Squires et al., 1992:pl. 1, fig. 32) of Venericardia hornii s.s. (Gabb, 1864) from the Crescent For- mation at Pulali Point, Washington. Family Cardiidae Lamarck, 1809 Nemocardium linteum (Conrad, 1855) Figure 58 Cardium linteum Conrad, 1855:3, 9; 1857:pl. 2, fig. 1. Anderson and Hanna, 1925:166-167, pi. 3, fig. 3. Cardium cooperii Gabb, 1864:172, pi. 24, figs. 154- 154a. Arnold, 1907:pl. 38, figs. 2-2a. Waring, 1 91 7:pl. 13, fig. 3. Hanna, 1927:285, pi. 41, figs. 6,7. Cardium dalli Dickerson, 1913:289, pi. 14, figs. 4a- c. Not Cardium dalli Heilprin, 1887:131, pi. 16a, fig. 70. Cardium marysvillensis Dickerson, 1916:482 [ nom . nov.]. Cardium {Protocar dium) marysvillensis Dicker- son. Clark and Woodford, 1927:94, pi. 15, fig. 12. Nemocardium linteum (Conrad). Stewart, 1930: 275-277, pi. 8, fig. 6. Turner, 1938:52, pi. 10, fig. 10. Vokes, 1939:76-77, pi. 11, figs. 6, 9. Weaver, 1943:159-160, pi. 38, fig. 3; 1953:28. Stewart, 1946:pl. 11, fig. 19. Moore, 1968:30, pi. 13d. Zinsmeister, 1974:97-98, pi. 9, figs. 7-9; 1983: pi. 2, fig. 7. Givens and Kennedy, 1979:table 4. Squires, 1984:49-50, fig. 12c; 1987:65, 67, fig. 113; 1988a:19, fig. 51. Squires et al., 1992:pl.l, fig. 33. Squires and Demetrion, 1992:42, fig. 121. Cardium ( Nemocardium ) linteum Conrad. Klein- pell and Weaver, 1963:202, pi. 34, fig 4. Nemocardium ( Nemocardium ) linteum (Conrad). Givens, 1974:49. Squires, 1977:table 1. PRIMARY TYPE MATERIAL. USNM holo- type 1834, Domengine Formation near Martinez, California. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12248. MOLLUSCAN STAGE RANGE. “Martinez” through “Tejon.” GEOGRAPHIC RANGE. Eastern Laguna San Ig- nacio area, Baja California Sur, Mexico, through Pulali Point, Washington. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1553a, 1554, 1555. REMARKS. Specimens are uncommon at local- ity 1553 and rare at the other localities. Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 23 Order Myoida Stoliczka, 1870 Family Corbulidae Lamarck, 1818 Genus Corbula Bruguiere, 1797 Subgenus Caryocorbula Gardner, 1926 Corbula ( Caryocorbula ) dickersoni Weaver and Palmer, 1922 Figure 59 Corbula dickersoni Weaver and Palmer, 1922:24- 25, pi. 9, figs. 9-10. Clark, 1938:700, pi. 1, fig. 17. Weaver, 1943:257-258, pi. 61, figs. 13, 16- 17, 20. Demere, Sundberg, and Schram, 1979:pl. 2, fig. 11. Corbula ( Caryocorbula ) dickersoni Weaver and Palmer. Vokes, 1939:98, pi. 16, figs. 1, 5, 9. Giv- ens, 1974:57, pi. 4, fig. 7. Squires, 1984:53, fig. 12m; 1987:70-71, fig. 124. PRIMARY TYPE MATERIAL. CAS holotype 7452, CAS paratypes 7452A-B, Cowlitz Formation, Lewis County, Washington, UW loc 329. ILLUSTRATED SPECIMEN. LACMIP hypo- type 12249. MOLLUSCAN STAGE RANGE. “Capay” through “Tejon.” GEOGRAPHIC RANGE. San Diego, California, through Little River area, Washington. LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. Specimens are rare. This is the first confirmed report of this species from the Crescent Formation. Squires et al. (1992) tentatively reported it from the Crescent Formation at Pulali Point, Washington. Phylum Arthropoda Siebold and Stannius, 1848 Class Cirripedia Burmeister, 1834 Order Thoracica Darwin, 1854 Family Scalpellidae Pilsbry, 1916 Genus Aporolepas Withers, 1953 Aporolepas sp. Figure 60 ILLUSTRATED SPECIMEN. LACMIP hypo- type 12250. LOCAL OCCURRENCE. CSUN Iocs. 1553, 1554. REMARKS. A single capitular plate was found at each locality. The Little River area specimens of this genus constitute the first published record of Aporolepas from the Pacific coast of North Amer- ica. One other locality is in the middle Eocene 24 ■ Contributions in Science, Number 444 Torrey Sandstone in San Diego County, southern California (V.A. Zullo, pers. comm., 1993). Phylum Chordata Balfour, 1880 Class Chondrichthyes Huxley, 1880 Order Lamniformes Berg, 1958 Family Odontaspididae Muller and Henle, 1837 Genus Palaeohypotodus Gliickman, 1964 Palaeohypotodus sp. Figure 61 ILLUSTRATED SPECIMEN. LACMVP 134133. LOCAL OCCURRENCE. CSUN loc. 1553. REMARKS. A single specimen was found. It closely resembles P. rutoti (Winkler, 1874) from the Paleocene of western Europe, Greenland, Maryland, and North and South Dakota, as well as from the lower Eocene (Ypresian Stage) of west- ern Europe (Cappetta, 1987; Ward and Wiest, 1990; Cvancara and Hoganson, 1993). Cappetta (1987:88, figs. 27A-C, 79E-J) and Cvancara and Hoganson (1993:figs. 3MM-3NN) illustrated P. rutoti. The Little River area specimen is the first record of Palaeohypotodus from the Pacific coast of North America. Family Lamnidae Muller and Henle, 1838 Genus Isurus Rafinesque, 1810 Isurus ? sp. Figure 62 ILLUSTRATED SPECIMEN. LACMVP 134434. LOCAL OCCURRENCE. CSUN loc. 1560a. REMARKS. A single specimen was found. It is not diagnostic enough to be positively identified as Isurus. Previously, Isurus has not been reported from the Crescent Formation. LOCALITIES CSUN MACROFOSSIL LOCALITIES 1220b. Along a prominent ridge, north side of a minor canyon on the west side of Mesa La Salina, 84-130 m above the bottom of the exposures of the Bateque For- mation in this area, approximately 1.25 km southeast of the intersection of 113°00'W and 26°45'N, Mexican gov- ernment topographic quadrangle map (scale 1:50,000) of San Jose de Gracia (number G12A64), Baja California Sur, Mexico, 1982. 1291a. South side of a minor canyon near the southern end of Mesa La Salina at 120 m elevation, at 112°56'13"W and 26°40fN, Mexican government topographic quadran- gle map (scale 1:50,000) of San Jose de Gracia (number G12A64), Baja California Sur, Mexico, 1982. 1553. At elevation of 870 ft., pebbly sandstone with Squires and Goedert: Crescent Formation Eocene Fossils mud matrix, on east side of Little River and logging road, just south of bridge over river, 60 m (200 ft.) south and 570 m (1,900 ft.) west of northeast corner of section 14, T 21 N, R 7 W, USGS topographic quadrangle, Grisdale, Washington, 7.5 minute, provisional edition 1990, 1:24,000. 1553a. Argillaceous siltstone 2 m stratigraphically above CSUN loc. 1553. 1554. At elevation of 760 ft., steeply dipping silty very fine-grained sandstone, on both sides of small stream on north side of road, on line between sections 10 and 15 (and nearly in center of south border of section 10), 720 m (2,400 ft.) west of southeast corner of section 10, T 21 N, R 7 W, USGS topographic quadrangle, Grisdale, Wash- ington, 7.5 minute, provisional edition 1990, 1:24,000. 1555. At elevation of 1,100 ft., pebbly sandstone in roadcut on west side of abandoned logging road, 1,110 m (3,700 ft.) and 510 m (1,700 ft.) west of northeast corner j of section 12, T 21 N, R 7 W, USGS topographic quad- rangle, Grisdale, Washington, 7.5 minute, provisional edi- tion 1990, 1:24,000. 1556. At elevation of 1,640 ft., pebbly sandstone near ridge top, on west side of logging road, 1,050 m (3,500 ft.) south and 810 m (2,700 ft.) west of northeast corner of section 12, T 21 N, R 7 W, USGS topographic quad- rangle, Grisdale, Washington, 7.5 minute, provisional edi- tion 1990, 1:24,000. 1557. At elevation of 1,660 ft., pebbly sandstone, 750 m (2,500 ft.) south and 780 m (2,600 ft.) west of northeast corner of section 12, T 21 N, R 7 W, USGS topographic quadrangle, Grisdale, Washington, 7.5 minute, provision- al edition 1990, 1:24,000. 1558. At elevation of 1,470 ft., poorly exposed sand- I stone on east side of an abandoned logging road on crest of low ridge, 1,170 m (3,900 ft.) south and 660 m (2,200 ft.) west of northeast corner of section 12, T 21 N, R 7 W, USGS topographic quadrangle, Grisdale, Washington, 7.5 minute, provisional edition 1990, 1:24,000. 1559. At elevation of 1,940 ft., pebbly sandstone ex- posed along a logging road on northwest side of ridge, 480 m (1,600 ft.) south and 960 m (3,200 ft.) west of northeast corner of section 12, T 21 N, R 7 W, USGS topographic quadrangle, Grisdale, Washington, 7.5 min- ute, provisional edition 1990, 1:24,000. 1560. At elevation of 1,960 ft., sandstone on east side of ridge, 510 m (1,700 ft.) south and 810 m (2,700 ft.) west of northeast corner of section 12, T 21 N, R 7 W, USGS topographic quadrangle, Grisdale, Washington, 7.5 minute, provisional edition 1990, 1:24,000. 1560a. At elevation of 1,960 ft., pebbly sandstone along east side of hill, 600 m (2,000 ft.) south and 870 m (2,900 ft.) west of northeast corner of section 12, T 21 N, R 7 W, USGS topographic quadrangle, Grisdale, Washington, 7.5 minute, provisional edition 1990, 1:24,000. 1563. At elevation of 2,230 ft., roadcut exposure, 300 m (984 ft.) north and 50 m (164 ft.) east of southeast corner of section 1, T 17 N, R 4 W, USGS topographic quadrangle, Capitol Peak, Washington, 7.5 minute, pro- visional edition 1986, 1:24,000. 1564. At elevation of 530 m, roadcut exposure, 800 m (2,624 ft.) north and 50 m (164 ft.) west of southeast corner of section 25, T 18 N, R 4 W, USGS topographic quadrangle, Summit Lake, Washington, 7.5 minute, 1981, 1:24,000. CAS LOCALITIES 181. Just back of the Gries Ranch house, about 6.5 km (4 mi.) east of Vader, Lewis County, Washington. SU LOCALITY 2696. Chivo Canyon, 5 km (3 mi.) N20°E of Bench Mark 961 at Santa Susana, USGS topographic quadrangle, Santa Susana, California, 7.5 minute, 1951 (photorevised 1969), 1:24,000. UCMP LOCALITIES 714. NE14 of SEV4 of section 21, T 1 S, R 1 E, Black Hills, Mt. Diablo State Park area, USGS topographic quadrangle, Tassarjara, California, 7.5 minute, 1968, 1:24,000. 1817. Opposite the place where Urruttia Canyon enters Salt Creek, 30 m (98 ft.) up fourth small draw from west end of ridge, SEV4 of the NWV4 of section 15, T 18 S, R 14 E, USGS topographic quadrangle, Joaquin Rocks, Cal- ifornia, 7.5 minute, 1969, 1:24,000. 5062. In sea cliff south of mouth of Soledad Valley, due west of midpoint between “P” and “u” of “Pueblo,” USGS topographic quadrangle, Del Mar, California, 7.5 minute, 1967, 1:24,000. A-1313. In Smith Canyon on west side of Capay Valley and west of Tancred, in the south central part of section 28, T 11 N, R 3 W, USGS topographic quadrangle, Guin- da, California, 7.5 minute, 1959 (photorevised 1980), 1:24,000. A-1315. About 3.2 km (2 mi.) north of Putah Creek and about 7.65 km (4.7 mi.) west-northwest of Winters, near center of section 14, T 8 N, R 2 W, USGS topo- graphic quandrangle, Monticello Dam, California, 7.5 minute, 1959, 1:24,000. USGS LOCALITY 4617. On southwest flank of Reef Ridge, north of McLure Valley, 3.6 km (2.2 mi.) south-southeast of El Cerrito oil well, section 27, T 23 S, R 17 E, USGS to- pographic quadrangle Cholame, California, 7.5 minute, 1961, 1:24,000. UW LOCALITIES 329. On north bank of the Cowlitz River at bend 1.5- 2.5 km (0.9-1. 5 mi.) east of Vader, section 28, T 1 N, R 2 W, USGS topographic quadrangle, Castle Rock, Wash- ington, 15 minute, 1953, 1:62,500. 353. From boulder-sized rocks not in place but in a modern landslide block, 2 km (1.2 mi.) south of Quilcene on west shore of Quilcene Bay just south of latitude 47°47'30"N, NEV4, section 36, T 27 N, R 2 W, USGS topographic quadrangle, Quilcene, Washington, 7.5 min- ute, 1953, 1:24,000. 358. At Joyce Station in Crescent Bay area, 0.4 km (0.25 mi.) east of Tongue Point, section 22, T 31 N, R 8 W, USGS topographic quadrangle, Joyce, Washington, 7.5 minute, 1978, 1:24,000. ARMENTROUT’S LOCALITY LR-1. Equivalent to CSUN loc. 1553 (see above). ACKNOWLEDGMENTS Gail H. Goedert assisted with fieldwork and collecting. Edward C. Wilson (Natural History Museum of Los An- geles County) provided geologic literature on corals. James H. McLean and LouElla Saul (Natural History Museum of Los Angeles County) helped with the identification of some of the gastropods. David R. Lindberg (University of Contributions in Science, Number 444 Squires and Goedert: Crescent Formation Eocene Fossils ■ 25 California, Museum of Paleontology) helped with the identification of the patellogastropods. Victor A. Zullo (University of North Carolina, Wilmington) identified the barnacle and shared his knowledge about Aporolepas. Ross E. Berglund (Bainbridge Island, Washington) iden- tified the shrimp and crab remains and provided some geologic literature. J.D. Stewart (Natural History Museum of Los Angeles County) helped identify the shark teeth. James H. McLean and David R. Lindberg shared their knowledge regarding the status of gastropod systematics. Lindsey T. Groves (Natural History Museum of Los An- geles County) and Charlotte A. Oyer (Oviatt Library, Cal- ifornia State University, Northridge) expertly helped in finding many of the more obscure taxonomic references. Hal L. Heitman and M.V. Filewicz (Unocal Corpora- tion, Houston, Texas) processed a microfossil sample for benthic foraminifers and calcareous nannofossils, respec- tively. William F. Kamin (Simpson Timber Company, Shelton, Washington) issued a special permit allowing fieldwork to be completed during the dry summer of 1 992. LITERATURE CITED Abbott, R.T. 1954. American seashells. New York: Van Nostrand Reinhold Company, 541 pp. Abbott, R.T., and S.P. Dance. 1982. Compendium of seashells. New York: E.P. Dutton, 411 pp. Adams, H., and A. Adams. 1853-1858. The genera of Recent Mollusca; arranged according to their or- ganization, 2 vols. London: John van Vorst, 660 pp. Addicott, W.O. 1970. Latitudinal gradients in Tertiary molluscan faunas of the Pacific coast. Palaeogeog- raphy, Palaeoclimatology, Palaeoecology 8(4):287- 312. Allan, R.S. 1940. Studies of the Recent and Tertiary Brachiopoda of Australia and New Zealand: Part 2. Records of the Canterbury Museum 4:277-297. Allmon, W.D. 1990. 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Los An- geles: Pacific Section, Society of Economic Paleon- tologists and Mineralogists, Volume and Guide- book. Received 13 April 1993; accepted 18 February 1994. Squires and Goedert: Crescent Formation Eocene Fossils Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Number 445 12 August 1994 IX I Contributions, in Science A Review of the Rissoiform Gastropods of Southwestern South America (Mollusca, Gastropoda) W. F. Ponder and T. M. Worsfold Natural History Museum of Los Angeles County Serial Publications ol THE Natural History Museum oe Los Angeles County Scientific Publications Committee James L. Powell, Museum President Daniel M. Cohen, Committee Chairman Brian V. Brown Kenneth E. Campbell Kirk Fitzhugh Edward C. Wilson Robin A. Simpson, Managing Editor The scientific publications of the Natural History Mu- seum of Los Angeles County have been issued at irregular intervals in three major series; the issues in each series are numbered individually, and numbers run consecutively, regardless of the subject matter. • Contributions in Science, a miscellaneous series of tech- nical papers describing original research in the life and earth sciences. • Science Bulletin, a miscellaneous series of monographs ]' describing original research in the life and earth sci- ii ences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now/!' published by the Museum in Contributions in Science. • Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through I the Museum Book Shop. A catalog is available on request.! The Museum also publishes Technical Reports, a mis- cellaneous series containing information relative to schol- arly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum’s Scientific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Tech- nical Reports may be obtained from the relevant Section of the Museum. H Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Printed at Allen Press, Inc., Lawrence, Kansas ISSN 0459-8113 A Review of the Rissoiform Gastropods of Southwestern South America (Mollusc a, Gastropoda) W. F. Ponder1 and T. M. Worsfold2 CONTENTS ABSTRACT 3 RESUMEN 3 INTRODUCTION 3 MATERIALS AND METHODS 4 ABBREVIATIONS 4 TAXONOMY 5 PART 1. Species from Peru to Tierra del Fuego, South America 5 Superfamily Cingulopsoidea 6 Family EATONIELLIDAE 6 Genus Eatoniella Dali, 1876 6 Subgenus Eatoniella 6 Eatoniella ( Eatoniella ) turricula n. sp 6 Eatoniella ( Eatoniella ) denticula n. sp 7 Eatoniella ( Eatoniella ) cf. denticula 11 Eatoniella ( Eatoniella ) cf. cana Ponder, 1983 12 Eatoniella ( Eatoniella ) ebenina n. sp 12 Eatoniella ( Eatoniella ) picea n. sp 12 Eatoniella ( Eatoniella ) nigra (Orbigny, 1840) 14 Eatoniella ( Eatoniella ) castanea n. sp 15 Eatoniella ( Eatoniella ) glomerosa n. sp 15 Subgenus Albosabula Ponder, 1965 19 Eatoniella ( Albosabula ) mcleani n. sp . 19 Genus Pupatonia Ponder, 1965 19 Pupatonia magellanica n. sp 19 Family CINGULOPSIDAE 21 Genus Skenella Pfeffer, 1886 21 Skenella hallae n. sp 21 Genus Eatonina Thiele, 1912 21 Subgenus Mistostigma Berry, 1947 21 Eatonina ( Mistostigma ) fusca (Orbigny, 1840) 21 Superfamily Rissooidea 23 Family RISSOIDAE 23 Genus Pusillina Monterosato, 1884 23 Subgenus Haurakia Iredale, 1915 23 Pusillina ( Haurakia ) averni n. sp 23 Pusillina ( Haurakia ) cf. averni 25 Genus Manzonia Brusina, 1870 25 Subgenus Alvinia Monterosato, 1884 25 Manzonia ( Alvinia ) limensis n. sp 25 Genus Powellisetia Ponder, 1965 25 Powellisetia microlirata n. sp 26 1. Australian Museum, 6-8 College Street, Sydney, New South Wales, 2000, Australia. 2. % Unicomarine, 7a, Diamond Centre, Letchworth, Hertfordshire SG61LW, U.K. Contributions in Science, Number 445, pp. 1-63 Natural History Museum of Los Angeles County, 1994 Genus Onoba H. and A. Adams, 1852 26 Onoba protofimbriata n. sp 26 Onoba subincisa n. sp 31 Onoba striola n. sp 36 Onoba subaedonis n. sp 36 Onoba sulcula n. sp 39 Onoba scytbei (Philippi, 1868) 39 Onoba fuegoensis (Strebel, 1908) 43 Onoba georgiana (Pfeffer, 1886) 45 Onoba erugata n. sp 45 Onoba amissa nom. nov 46 Onoba (?) algida n. sp. 47 Onoba (?) lacuniformis n. sp 47 Family BARLEEIDAE 48 Genus Barleeia Clark, 1853 48 Barleeia meridionalis n. sp 48 Family Anabathridae 49 Genus Amphithalamus Carpenter, 1864 49 Amphithalamus cf. inclusus Carpenter, 1864 49 Superfamily RISSOELLOIDEA 49 Family RISSOELLIDAE 49 Genus Rissoella J.E. Gray, 1847 49 Subgenus Rissoella 49 Rissoella ( Rissoella ) peruviana n. sp . 49 PART 2. Additional species and records for South Georgia and Falkland Islands 51 Family Eatoniellidae 51 Eatoniella ( Eatoniella ) strebeli n. sp 51 Eatoniella ( Eatoniella ) cf. cana Ponder, 1983 52 Eatoniella ( Eatoniella ) occulta Ponder, 1983 52 Eatoniella ( Eatoniella ) bennetti (Preston, 1912) 52 Family Rissoidae 54 Onoba georgiana (Pfeffer, 1886) 54 Onoba subaedonis n. sp 54 Onoba scytbei (Philippi, 1868) 54 Onoba cf. protofimbriata 54 Onoba anderssoni (Strebel, 1908) 54 Onoba filostria (Melvill and Standen, 1912) 54 Onoba turqueti (Lamy, 1905) 54 Onoba cf. gelida (E.A. Smith, 1907) 54 Powellisetia australis (Watson, 1886) 54 Family Cingulopsidae 54 Skenella georgiana Pfeffer (in Martens and Pfeffer), 1886 54 Skenella wareni n. sp 55 Family Rissoellidae 55 Rissoella (Jeffrey siella) cf. powelli Ponder, 1983 55 PART 3. Species from Juan Fernandez Islands 55 Family Eatoniellidae 55 Eatoniella ( Eatoniella ) zigzag n. sp 55 Family Rissoidae 56 Onoba (?) isolata n. sp 56 Onoba (?) protopustulata n. sp 56 DISCUSSION 57 ACKNOWLEDGMENTS 58 LITERATURE CITED 58 APPENDIX: LIST OF LOCALITIES 59 Natural History Museum of Los Angeles County (LACM) Localities 59 National Museum of Natural History, Washington, D.C. (USNM) Material 61 Natural History Museum (London) (BMNH) Material 62 National Museum of Wales (NMW) Material 62 Swedish Museum of Natural History (SMNH) Material 62 T. Worsfold Stations 62 Locality Numbers 62 2 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ABSTRACT. Rissoiform gastropods from southwestern South America (Peru to Tierra del Fuego) and Juan Fernandez Islands are described and additional records are given. Taxa are also described for South Georgia and the Falkland Islands and new locality records of previously described taxa are listed from those areas. The southwestern South American fauna described consists of 10 species of Eatoniellidae comprising nine species of Eatoniella, seven of these new and one of which is in the subgenus Albosabula, and one new species of Pupatonia, the first record of the latter genus outside New Zealand. There are two species of Cingulopsidae; one, a species of Skenella, is new. The 15 taxa included in the Rissoidae (subfamily Rissoinae only) comprise 1 new species each of Pusillina { Haurakia ), Powellisetia, and Man- zonia ( Alvinia ) and 12 species of Onoba, 8 of which are new. A single new species of Barleeia (Barleeidae) and Rissoella (Rissoellidae) is also described and an Amphitbalamus (Anabathridae), close to A. inclusus Carpenter, is recorded. Three new taxa are described from Juan Fernandez Islands, one Eatoniella and two rissoids tentatively included in Onoba. New species of Eatoniella and Skenella are described from South Georgia. The biogeography of the fauna is briefly discussed. The South American taxa as far north as latitude 42°S are similar to those encountered on sub-Antarctic islands, whereas north of latitude 42°S the few species present are like those of southern California and Mexico. A replacement name, Eatoniella afronigra, is provided for the South African eatoniellid Rissoa nigra Krauss, 1948, a secondary homonym of P aludestrina nigra Orbigny, 1840, also an Eatoniella. RESUMEN. Se describen los gastropodos rissoiformes del sureste de Sudamerica (Peru a Tierra del Fuego) e islas Juan Fernandez y se entregan registros adicionales. Se describen ademas taxa para las islas Georgia del Sur y Malvinas y se entrega una lista con nuevos registros de localidades de especies descritas previamente de estas areas. La fauna sudamericana descrita consiste de diez especies de Eatoniellidae, comprendiendo nueve especies de Eatoniella, siete son nuevas y una esta en el subgenero Albosabula, y una especie de Pupatonia, el primer registro de este ultimo genero fuera de Nueva Zelandia. Existen dos especies de Cingulopsidae; una de ellas, del genero Skenella, es nueva. Los 15 taxa que se incluyen en Rissoidae (solamente la subfamilia Rissoinae) comprenden una sola especie de los generos Pusillina {Haurakia), Powellisetia y Manzolia {Alvinia), y 12 especies de Onoba, 8 de las cuales son nuevas. Se describe tambien una nueva especie de Barleeia (Barleeidae) y de Rissoella (Rissoellidae) y se registra una de Amphitbalamus (Anabathridae), cercana a A. inclusus Carpenter. Se describen tres nuevos taxa de las islas Juan Fernandez, uno de Eatoniella y dos rissoideos, tentativamente incluidos en Onoba. Se describe una nueva especie de Eatoniella y de Skenella de las islas Georgia del Sur. La biogeografia de la fauna se discute brevemente. Los taxa sudamericanos hasta la latitud 42°S son similares a aquellos encontrados en islas subantarticas, mientras que al norte de la latitud 42°S las pocas especies presentes se parecen a aquellas del sur de California y Mexico. Se proporciona un nombre de reemplazo, Eatoniella afronigra, para el eatoniellido Rissoa nigra Krauss, 1948, un homonimo secundario de P aludestrina nigra Orbigny, 1840, tambien un Eatoniella. INTRODUCTION Small gastropods similar to members of the Ris- soidae have proved to be difficult to classify on shell characters alone. It is only in the last three decades that studies on radulae and anatomy have provided a solidly based classification for these an- imals. Because many workers have difficulty in sep- arating the families involved, they are treated to- gether in this work. An attempt is made in this paper to describe the majority of rissoid and rissoid-like taxa occurring in southwestern South America from Peru to Tierra del Fuego. Three taxa from a sample from Juan Fernandez Islands are also described. The taxa reviewed in this work include members of three superfamilies, two of which are closely related. These are the caenogastropod Rissooidea (Rissoidae) and Cingulopsoidea (Eatoniellidae and Cingulopsidae) (see Ponder, 1988, for discussion on relationships) and the superficially similar but very different Rissoelloidea (Rissoellidae), which be- longs in the subclass Heterobranchia. All of these groups closely resemble rissoids and have, in the past, been included in, or closely associated with, that family. This paper is intended to supplement and com- plement a similar work on the Antarctic and sub- Antarctic fauna (Ponder, 1983a). The only family not dealt with herein that was included in Ponder (1983a) is the Orbitestellidae, of which taxa are present in the region (Ponder, 1990). The oppor- tunity is also taken to report on some additional collections from South Georgia and the Falkland Islands. There are very few studies on small marine gas- tropods from South America. Of the 30 species recognized below, only seven have been described previously, with two additional available names proving to be synonyms of earlier named taxa. A few species described from Argentina in the last 20 years (Castellanos and Fernandez, 1972a, b, 1974) have not been available for study but, as far as can be determined from their descriptions and figures, are different from any of the taxa dealt with in this report. Dell (1990), in his major review of Antarctic Mollusca, dealt with some material from South America but did not include records of rissoiform species from that area. Similarly, other faunistic Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 3 Figure 1. Map of South America showing location of outlying localities and insets indicating location of maps shown in Figures 2 and 3. studies on the Mollusca of the Pacific Coast of South America (e.g., Dali, 1909; Dell, 1971) have mainly dealt with the larger species, the microm- olluscan fauna remaining largely unknown. Members of the genus Rissoina (Rissoidae, Ris- soininae) are not discussed here, as they are being investigated by W. Sleurs. MATERIALS AND METHODS Shells were measured using an eyepiece micrometer in a Wild M5 stereomicroscope. The measurements presented are given as the raw data rather than means because the specimens measured were not extracted from the avail- able material randomly, but rather well-preserved, intact adult specimens were selected. The full locality data are given in the appendix. These data are only repeated in the material examined section for each taxon for the primary type localities of new taxa, additional material being referred to by station numbers and the institution in which it is deposited. The number of specimens in each lot that can confidently be assigned to the species is given in square brackets. Where the entire sample consists of empty shells, this is indicated by the letter “d” (=dead) in square brackets. Some juveniles, fragments, and other specimens in the lots that cannot be reliably identified are not included in the number of spec- imens given. The localities are listed in the appendix by institution and then numerically by station number. The location of each station is given in Figures 1-4. Because 4 ■ Contributions in Science, Number 445 of the large number of stations involved, these were grouped into tight geographic clusters and each cluster numbered. These numbers are given for each station in the material examined section of each species and also in the appendix. This was done to facilitate the geographic location of any particular station by reference to Figures 1-4. In addition, the locality numbers are also listed nu- merically with a list of all of the station numbers referrable to each of these numbers. Radulae, shells, and opercula were examined using the scanning electron microscope (SEM) after standard prep- aration. The orientation of the opercula (anterior and posterior ends, inner and outer edges) are given as though the operculum is retracted in the shell aperture. In each case the number of radulae examined is indicated. In the relatively few cases where ethanol-preserved material is available, the pigmentation of the head-foot and visceral coil is noted. ABBREVIATIONS INSTITUTIONS AMS — Australian Museum, Sydney BMNH — Natural History Museum, London LACM — Los Angeles County Museum of Natural His- tory MCZ — Museum of Comparative Zoology, Harvard Uni- versity, Cambridge NMNHP — Museum National d’Histoire Naturelle, Paris NMNZ — National Museum of New Zealand, Welling- ton NMW — National Museum of Wales, Cardiff SMNH — Naturhistoriska Riksmuseet, Stockholm USNM — National Museum of Natural History, Smith- sonian Inst., Washington, D.C. ABBREVIATIONS USED IN LOCALITY DETAILS DE — Discovery Expedition Station E — Eltanin Station H — Hero Cruise 712 Station R/V — Research Vessel SNAE — Scottish National Antarctic Expedition Station SSPE — Swedish Southpolar Expedition Station Sta. — Station TW — T. Worsfold Station V — Verna Station ABBREVIATIONS USED IN SHELL MEASUREMENTS AL — aperture length BA — number of axials on last whorl BS — number of spirals on last whorl PA — number of axials on penultimate whorl PD — protoconch diameter PS — number of spirals on penultimate whorl PW — number of protoconch whorls SL — shell length SW — shell width TW — number of teleoconch whorls Ponder and Worsfold: Review of Rissoiform Gastropods Figure 2. A, Localities in northern Peru. B, Localities in southern Peru and northern Chile. C, Localities in central Chile. TAXONOMY This section deals with species grouped according to three geographic areas: 1. Species from Peru to Tierra del Fuego, South America. 2. Species and records for South Georgia and the Falkland Islands additional to those recorded by Ponder (1983a). 3. Species from Juan Fernandez Islands. Part 1 Species from Peru to Tierra del Fuego, South America Subclass PROSOBRANCHIA This grouping is paraphyletic according to recent analyses of gastropod phylogeny (e.g., Haszprunar, 1988). It is used here in the traditional way pending an alternative ranked classification. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 5 Figure 3. Localities in southern Chile and Tierra del Fue- go. The inset indicates location of Figure 4. Order Caenogastropoda Indo-west Pacific. None are known from northern South America, Central America, or North America where it is replaced by a family with very similar shell features, the Barleeidae. The Argentinean Ea- toniella rubrooperculata Castellanos and Delicia, 1971, is a member of the Barleeidae. The eaton- iellids of New Zealand (Ponder, 1965a), Australia (Ponder and Yoo, 1977a), and the Antarctic-sub- Antarctic (Ponder, 1983a) have been reviewed. The family is distinguished by having a well-developed peg on the operculum, a “littorinid”-like radula, no true penis, and open pallial genital ducts. Genus Eatoniella Dali, 1876 Species of this genus have a depressed-ovate to con- ic shell with a markedly prosocline outer lip. The operculum usually has an opaque insertion area and lacks a distinct internal ridge. See Ponder and Yoo (1977a) for a formal synonymy and diagnosis. Subgenus Eatoniella The typical subgenus is differentiated from the next by having the midbasal part of the central teeth of the radula not markedly produced ventrally. Eatoniella (Eatoniella) turricula n. sp. Figures 5A, 6D, 7A, 8A Superfamily Cingulopsoidea Family EATONIELLIDAE This family is distributed around the southern con- tinents and a few taxa are found in the northern ETYMOLOGY. Turricula — Latin. A little tow- er. Refers to the shell shape. MATERIAL EXAMINED. Types. Holotype, LACM 2656, 205 paratypes, LACM 2657; 8 paratypes, AMS C. 167414. 33R 71-328. 6.4 km N Cabo San Juan, Isla de 33 J Figure 4. Map showing the localities around Isla de los Estados. 6 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods los Estados, Tierra del Fuego, Argentina. 54°39.1'S, 63°50.1'W, 135-137 m, Sta. 874, USARP-SOSC-R/V Hero Cr.715, 26 Oct. 1971. Additional Material Examined. Southern Chile: 22A USNM E 960 [1]. 22B USNM E 958 [6]. Tierra del Fuego: 27B USNM V 17-48 [1(d)]. 27C USNM E 219 [3(d)]. 28 71-271 [3(d)], 29B 71-305 [17(d)], 30A 71-342 [4(d)], 30H 71-332 [2]; 30H USNM H 664 [many(d)]. 301 71-329 [many]. 33A USNM H 656 [9(d)], 33B 71-347 [many]; 71- 267 [1(d)]. 33C 71-348 [many], 33D 71-352 [20(d)]. 33E 71-351 [many]. 33H 71-313 [4(d)]. 33K 71-315 [3(d)], 33L 71-316 [10(d)]. 33P 71-319 [1(d)]. 33Q 71-327 [10 ( + many d)]. 35 37B USNM E 967 [2(d)]. 37A BMNH DE 88 [1(d)]. (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Figs. 5A, 6D). Small (maxi- mum length 2.8 mm), conical, thin, translucent when fresh, with 3.2-4. 1 teleoconch whorls. Spire with straight outlines, whorls very lightly convex; pe- riphery of last whorl lightly angled. Sutures im- pressed, simple. Teleoconch smooth and rather glossy with faint prosocline growth lines. Proto- conch (Fig. 6D) smooth except for fine spiral grooves, of 1.2-1. 7 whorls. Aperture moderately large, circular, with sharp peristome, lacking exter- nal varix. Inner lip narrow, outer lip moderately to strongly prosocline. Umbilical chink minute. Peri- ostracum very thin, transparent. Color yellowish- white, sometimes pale grey, fading to white. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 2.75 1.46 1.87 0.87 3.13 4.1 1.2 0.41 Paratypes 2.47 1.47 1.68 0.85 2.91 3.3 1.7 0.48 2.75 1.50 1.83 0.94 2.91 3.8 1.3 0.46 2.66 1.57 1.69 0.93 2.85 3.2 1.7 0.53 2.75 1.53 1.80 0.92 2.99 4.1 1.5 0.41 2.66 1.43 1.84 0.91 2.91 3.8 1.4 0.45 2.78 1.53 1.80 0.95 2.90 3.4 1.6 0.48 2.48 1.36 1.82 0.81 3.04 3.2 1.7 0.53 2.62 1.52 1.72 0.93 2.80 3.4 1.6 0.48 2.49 1.42 1.74 0.82 3.01 3.3 1.5 0.44 Operculum (Fig. 7A). Pale yellow, oval, weakly angled posteriorly and anteriorly; inner and outer edges equally convex. Peg curved, well developed. Radula (Fig. 8A). Central teeth with cusp formula 3 + 1+3, median cusp moderately long, narrow. Lateral teeth with cusp formula 2+1+3, primary cusp narrow, pointed. Inner marginal teeth with cusp formula 5 + 1 + 5, primary cusp small, outer- most cusps very small. Outer marginal teeth with about 6 small cusps (based on 2 radulae). Animal. Unpigmented. REMARKS. The shell of this species is distin- guishable from the next (E. denticula n. sp.) by its slightly larger size and more conical shape, the whorls being flatter. The shell surface is also smoother and glossier when fresh. The two species also differ in radular details (see below) and occur at different depths. Eatoniella turricula is similar to E. cana Ponder, 1983, but the shell has a broader base, is more conical, is glossier, and differs in color, E. cana having a dark spire and a non-pigmented last whorl. Eatoniella turricula is also similar in shell and rad- ular features to E. kerguelenensis (E.A. Smith, 1875) but has a more evenly conical shell because of the flatter whorl outlines. The shell of E. turricula is about the same size as that of E. kerguelenensis kerguelenensis, but smaller than that of E. kergue- lenensis regularis (E.A. Smith, 1915), these latter taxa also usually having a more darkly pigmented shell. Most of the records of E. turricula are based on empty, faded shells. A shallow-water species, E. ebenina n. sp., has a darkly pigmented shell but faded specimens of the two species closely resemble one another and it is possible that some of our identifications based on dead shells may be incor- rect. Eatoniella turricula and E. ebenina are con- trasted in the remarks under the latter species. DISTRIBUTION. Southern Chile and Tierra del Fuego in 40-900 m. Common. Eatoniella ( Eatoniella ) denticula n. sp. Figures 5B, C, 6C, 7B, C, 8B, F, G ETYMOLOGY. Denticula — Latin. A little tooth. Refers to the shell shape. MATERIAL EXAMINED. Types. Holotype, LACM 2658, 9 paratypes, LACM 2659; 5 paratypes, AMS C. 167415. 32H 71-287. Puerto Cook, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.25'S, 64°2.3'W, inter- tidal, Sta. 71-2-37, USARP-SOSC-R/V Hero Cr.712, 17 May 1971. Additional Material Examined. Southern Chile: 17 73- 75 [8(d)]. 21 73-71 [many]. 24 75-48 [26]. T ierra del Fuego: 25 73-69 [21(d)]. 28 71-270 [13( + 19d)]; 71-271 [4( + 14d)]. 30A USNM H 654 [1(d)]. 30E 71-339 [2(d)]. 31A 71-274 [6]. 32B 71-295 [4]. 32D 71-293 [2], 32G 71-311 [7]. 32H 71-287 [3], 33A 71-258 [2(d)]; 71-308 [9(d)]; USNM H 656 [many(d)]. 33G 71-266 [9]; 73-66 [many(d)]. 331 71- 317 [17(d)]. 33J 71-264 [many(d)]. 330 71-310 [many(d)]. (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Figs. 5B, C, 6C). Small (max- imum length 2.0 mm), ovate-conical, moderately thick, translucent when fresh, with 2. 5-3. 5 teleo- conch whorls. Spire with lightly convex to straight outlines, whorls lightly to moderately convex; pe- riphery of last whorl rounded. Sutures impressed, simple. Teleoconch moderately smooth with faint growth lines. Protoconch (Fig. 6C) smooth, of 1.2- 1.7 whorls. Aperture subcircular, weakly angled posteriorly, with moderately sharp peristome, lack- ing external varix. Inner lip narrow, outer lip mod- erately prosocline. Umbilical chink very small. Peri- ostracum very thin, transparent. Color pale grey to pale yellowish-white, fading to white. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 7 Figure 5. Shells of species of Eatoniella. A, Eatoniella turricula n. sp., holotype, length 2.75 mm. B, C, Eatoniella denticula n. sp.; B, holotype, length 1.87 mm; C, Sta. 71-270, shell, length 1.94 mm. D, Eatoniella picea n. sp., holotype, length 2.44 mm. E, Eatoniella ebenina n. sp., holotype, length 3.25 mm. F, Eatoniella nigra (Orbigny), Sta. 75-20, length 1.69 mm. Scale bars: 500 jum. Dimensions. Sta. 71-270 Fig. 5C 1.94 1.08 1.79 0.63 3.07 3.4 1.3 0.36 SL/ SL/ 1.63 0.98 1.65 0.61 2.67 3.1 1.2 0.32 SL SW SW AL AL TW PW PD 1.62 0.98 1.64 0.56 2.90 3.2 1.2 0.33 Holotype 1.87 1.13 1.65 0.68 2.73 3.1 1.4 0.40 1.77 1.03 1.72 0.64 2.75 3.4 1.3 0.32 Paratypes 1.83 1.14 1.60 0.68 2.67 2.7 1.6 0.43 1.71 1.02 1.68 0.63 2.71 3.2 1.2 0.33 1.74 1.08 1.61 0.65 2.67 2.8 1.5 0.42 Sta. 71-311 1.90 1.16 1.65 0.63 3.02 3.4 1.3 0.32 1.67 1.04 1.61 0.64 2.60 2.9 1.4 0.38 2.04 1.21 1.69 0.63 2.77 3.5 1.2 0.33 1.71 0.98 1.74 0.63 2.71 2.8 1.5 0.43 1.99 1.17 1.71 0.66 3.00 3.2 1.3 0.36 1.64 0.98 1.66 0.62 2.64 2.6 1.6 0.42 1.81 1.07 1.69 0.64 2.82 3.0 1.4 0.41 1.74 1.03 1.70 0.63 2.76 2.7 1.6 0.43 1.71 1.04 1.65 0.57 3.02 3.1 1.2 0.34 1.72 1.06 1.63 0.66 2.60 2.6 1.7 0.43 1.67 0.98 1.70 0.63 2.64 2.5 1.7 0.42 Operculum (Fig. 7B, C). Pale yello’ w, oval, more 1.69 0.96 1.76 0.62 2.72 2.6 1.6 0.43 strongly angled posteriorly than anteriorly, inner 8 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods Figure 6. Protoconchs of species of Eatoniella. A, B, Eatoniella castanea n. sp., paratype, apical view (B) and teleoconch microsculpture (A). C, Eatoniella denticula n. sp., Sta. 71-270, lateral view. D, Eatoniella turricula n. sp., paratype, protoconch microsculpture. E, Eatoniella glomerosa n. sp., Sta. 71-283, lateral view of protoconch. Scale bars: A, D, 20 yum; B, 100 jam; C, E, 50 /mi. and outer edges approximately equally convex. Peg curved, well developed. Radula (Fig. 8B, F, G). Central teeth with cusp formula 2-3 + 1+2-3 median cusp moderately long, narrow. Lateral teeth with cusp formula 2+1+2 ( + 1 outer denticle in some), primary cusp narrow, pointed. Inner marginals apparently with cusp for- mula 3 + 1+2 (somewhat obscured in mounts). Outer marginals also obscured but with at least 7- 8 small cusps (based on 4 radulae). Animal. Unpigmented or with a slight grey tinge on visceral coil. REMARKS. The shell of this species is contrast- ed with E. turricula above and differs from that species in its smaller shell, which has more convex Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 9 Figure 7. Opercula of species of Eatoniella. All views inner side. A, Eatoniella turricula n. sp., paratype. B, C, Eatoniella denticula n. sp.; B, Sta. 71-270; C, paratype. D, Eatoniella picea n. sp., paratype. E, Eatoniella nigra (Orbigny), Sta. 75- 20. F, H, I, Eatoniella glomerosa n. sp.; F, Sta. 71-270; H, paratype; I, Sta. 71-283. G, Eatoniella ebenina n. sp., Sta. 73-69. Scale bars: A-F, H, I, 100 jum; G, 200 ixm. 10 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods Figure 8. Radulae of Eatoniella species. A, Eatoniella turricula n. sp., paratype. B, F, G, Eatoniella denticula n. sp.; B, Sta. 71-270; F, G, paratypes. C-E, Eatoniella ebenina n. sp., Sta. 73-69; C, E, detail of central teeth. Scale bars: A-C, E-G, 10 jum; D, 20 /urn. whorls. The radula in the two species are similar but in E. denticula the primary cusps of the inner marginal, lateral, and central teeth are blunter and the number of cusps on the inner marginal teeth apparently smaller. Eatoniella denticula is rather similar to the Antarctic E. demissa (E.A. Smith, 1915), but the shell of the latter species is broader and the radula has more cusps on the central teeth. The shell of E. denticula is somewhat similar to the southern Australian E. victoriae Ponder and Yoo, 1978, but the shell of that species is thinner, slightly broader, and smaller. DISTRIBUTION. Southern Chile and Tierra del Fuego in algae and among rocks; mainly intertidal with occasional specimens to 15 m. Empty shells have been found down to about 30 m. Common. Empty shells and a few live specimens found down to 124 m may be a different species and are listed separately; see below. Eatoniella ( Eatoniella ) cf. denticula MATERIAL EXAMINED. Southern Chile: 22B USNM E 958 [6(d)]. Tierra del Fuego: 27B USNM V 17-48 [2(d)]. 27C USNM E 219 [1(d)]. 27E BMNH DE 388 [many(d)]. 30H USNM H 664 [28(d). 33A USNM H 656 [1(d)]. 33B 71-267 [1(d)]. 33C 71-348 [4(d)], 33E 71-351 [6(d)], 33L 71-316 [5(d)], 330 71-310 [2(d)]. 33Q 71-327 [10]. 33R 71-328 [2(d)]. 37B USNM E 967 [3]. (All material LACM unless otherwise indicated.) REMARKS. Some lots of empty shells from deeper water, often occurring together with E. tur- ricula, are generally similar to E. denticula but are Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods! 11 larger and more elongate and mostly (bleached?) white. Although some intermediate shell forms are seen, it is quite possible that this material represents another taxon, and we do not feel confident in unreservably assigning it to E. denticula. DISTRIBUTION. Southern Chile and Tierra del Fuego in 18-124 m. Eatoniella ( Eatoniella ) cf. cana Ponder, 1983 Eatoniella cana Ponder, 1983a: 6 (in part). MATERIAL EXAMINED. Southern Chile: 17 LACM 73-75 [12(d)]. Tierra del Fuego: 27 A NMNHP “Baie Or- ange,” Mission du Cap Horn [11]. BMNH 37A DE 88 [1(d)]. Falkland Islands: SNMH SSPE 48 [10]. REMARKS. A few empty shells from LACM and Discovery stations and a single lot found among the old “spirit” collections in the National Muse- um, Paris, are very similar to E. cana but differ in having a slightly smaller shell and more convex whorls. They agree closely with specimens from the Falkland Islands, and it is possible that the ma- terial from these two areas is referable to a separate species-group taxon. The opportunity is taken to list an additional locality for this species from the Falkland Islands. DISTRIBUTION. Falkland Islands, southern Chile, and Tierra del Fuego. Typical form from South Georgia and South Orkney Islands. Eatoniella ( Eatoniella ) ebenina n. sp. Figures 5E, 7G, 8C-E ETYMOLOGY. Ebenina-— Latin. Black. Refers to the shell color. MATERIAL EXAMINED. Types. Holotype, LACM 2660, 12 paratypes, LACM 2661; 3 paratypes, AMS C. 167416. 20 73-72. Bahia Tom, Magallanes Prov., Chile. 50°11.3'S, 74°47.9'W, 14 m, P. Dayton (R/V Hero), 21 May 1973. Additional material examined. Southern Chile: 21 73- 71 [15(d)]. 23 73-70 [28(d)]. 24 75-49 [13(d)]. Tierra del Fuego: 25 73-69 [10( + 21d)]. 26 73-68 [1(d)]. 28 71-296 [4(d)], 32F 71-326 [1(d)]. 33G 73-66 [21(d)], 330 71-310 [12(d)]. (All material LACM.) DIAGNOSIS. Shell (Fig. 5E). Small (maximum length 3.3 mm), elongate-conic, moderately thin, opaque, with 3. 5-4.1 teleoconch whorls. Spire with straight outlines, whorls very lightly convex; pe- riphery of last whorl sometimes slightly angled. Su- tures impressed, simple. Teleoconch smooth with faint growth lines and rather dull surface. Proto- conch smooth, of 1.5-1. 8 whorls. Aperture ovoid, weakly angled posteriorly, with sharp peristome, lacking external varix. Inner lip narrow, outer lip moderately prosocline. Umbilical chink very small or absent. Periostracum very thin, transparent. Col- 12 ■ Contributions in Science, Number 445 or uniform dark grey to black except for yellow- white to white patch on lower base. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD Holotype 3.25 1.65 1.97 1.05 3.10 4.0 1.8 0.54 Paratypes 3.19 1.66 1.92 1.10 2.89 4.1 1.7 0.56 3.05 1.60 1.90 1.03 2.97 4.0 1.8 0.49 2.86 1.55 1.84 0.86 3.10 3.9 1.8 0.46 3.03 1.73 1.75 1.07 2.83 3.8 1.5 0.49 3.14 1.71 1.83 1.06 2.96 4.4 1.7 0.42 2.95 1.63 1.82 1.07 2.76 3.5 1.8 0.53 2.83 1.59 1.78 0.98 2.88 3.5 1.7 0.50 3.04 1.62 1.88 1.00 3.05 3.8 1.8 0.52 3.16 1.64 1.93 1.05 3.01 3.7 1.8 0.54 Operculum (Fig. 7G). Yellow, oval, with post- erior end more strongly angled than anterior. Inner and outer edges of equal convexity. Peg curved, well developed. Radula (Fig. 8C-E). Central teeth with cusp for- mula 3 + 1+3, median cusp long, bluntly pointed to spatulate, outermost cusps very small. Lateral teeth with cusp formula 2 + 1 + 3. Inner marginal teeth with cusp formula 4-5 + 1 + 3-5, outermost cusps minute. Outer marginal teeth with 10+ small, | sharp cusps (based on 3 radulae). Animal. Unknown. REMARKS. This species can be distinguished from the other dark-colored South American spe- cies by its larger size and relatively flatter whorls. Eatoniella turricula is similar in shell shape but can be separated on shell color and its larger size, gloss- ier surface, and flatter whorls. The radulae are also similar, but the primary cusps of the central and lateral teeth are blunt in E. ebenina and sharp in E. turricula. As noted under E. turricula, empty, faded shells of these two species are difficult to distinguish, and it is possible that some records are incorrectly assigned. There is also some similarity with E. cana, but E. ebenina has a thinner, larger shell and uniform dark grey or black color. Other somewhat similar dark-colored eatoniellids include the thicker- shelled, broader sub-Antarctic E. kerguelenensis, which has more convex whorls, and the smaller and less angled New Zealand E. olivacea (Hutton, 1882). DISTRIBUTION. Southern Chile and Tierra del Fuego from intertidal to 15 m. Live-collected spec- imens all subtidal. Not common. Eatoniella ( Eatoniella ) picea n. sp. Figures 5D, 7D, 9A, B ETYMOLOGY. Picea— Latin. Pitch black. Re- fers to the shell color. MATERIAL EXAMINED. Types. Holotype, LACM 2662, 100 paratypes, LACM 2663; 8 paratypes, AMS Ponder and Worsfold: Review of Rissoiform Gastropods Figure 9. Radulae of Eatoniella species. A, B, Eatoniella picea n. sp., paratypes. C, Eatoniella nigra, Sta. 75-20. D, Eatoniella castanea n. sp., paratype. Scale bars: 10 n m. C. 167417. 32G 71-311. Observatorio, Isla de los Estados, Tierra del Fuego, Argentina. 54°39.5'S, 64°08'W, intertid- al rocks. Sta. 699, USARP-SOSC-R/V Hero Cr.715, 19 Oct. 1971. Additional Material Examined. Southern Chile: 24 75- 48 [1]. Tierra del Fuego : 28 71-270 [many]; 71-271 [many]. 31A 71-274 [1(d)]. 31B 71-276 [1(d)]. 31D 71-283 [13]. 32A 71-273 [many]. 32A 71-309 [16]. 32C 71-294 [1(d)]. 32D 71-291 [2(d)]; 71-293 [1(d)]. 32E 71-289 [many]; 71- 290 [4(d)], 32F 71-326 [2], 33G 73-66 [21(d)]. 32H 71-287 [many]. 33F 71-265 [1(d)]. 330 71-310 [1(d)]. 38 75-51 [1], 39 71-268 [11(d)]. (All material LACM.) DIAGNOSIS. Shell (Fig. 5D). Small (maximum length 2.4 mm), elongate-conic, moderately thin, with 3. 5-4.0 teleoconch whorls. Spire with very lightly convex outlines, whorls moderately convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth and rather glossy, with moderately prominent prosocline growth lines. Protoconch smooth, of 1.1-1. 3 whorls. Aperture oval, weakly angled posteriorly, with sharp peri- stome, lacking external varix. Inner lip narrow, out- er lip slightly to moderately prosocline. Umbilical chink very small or absent. Periostracum very thin, transparent. Color black or grey, often with reddish tinge, occasionally yellowish, paler near growing edge. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 2.44 1.23 1.98 0.85 2.89 3.9 1.3 0.35 Paratypes 2.44 1.25 1.95 0.82 2.96 — — — 2.22 1.16 1.92 0.79 2.81 3.8 1.3 0.36 2.21 1.14 1.93 0.75 2.96 3.6 1.2 0.33 2.25 1.26 1.78 0.83 2.69 4.0 1.2 0.30 2.40 1.25 1.91 0.80 2.99 4.0 1.2 0.35 2.14 1.19 1.80 0.76 2.82 — — — 2.30 1.28 1.79 0.85 2.83 3.5 1.3 0.38 2.10 1.11 1.88 0.73 2.88 — — — 2.28 1.22 1.87 0.83 2.73 3.8 1.1 0.35 Operculum (Fig. 7D). Pale yellow with brown markings to yellow-brown or almost black, oval with almost equally angled anterior and posterior ends, inner edge slightly angled. Peg rather narrow, curved. Radula (Fig. 9A, B). Central teeth with cusp for- mula 3 + 1 + 3, median cusp large, spatulate, with finely denticulate end. Lateral teeth with cusp for- mula 3 + 1+3, primary cusp narrow, sharp. Inner marginal teeth with cusp formula 5-6 + 1+2, pri- mary cusp large and rather blunt. Outer marginal teeth with about 7 small cusps, outermost largest (based on 2 radulae). Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 13 Figure 10. A, Eatoniella nigra (Orbigny), syntype, length 1.77 mm. B, Onoba amissa nom. nov. pro Paludestrina striata (Orbigny), lectotype, length 2.18 mm. C, Eatonina fusca (Orbigny), syntype, length 1.28 mm. D, Eatoniella bennetti (Preston), lectotype, length 1.67 mm. Scale bar: 500 ixm. Animal. Head and foot grey, mantle and viscera mottled dark grey. REMARKS. The shell of this species is similar in color to that of E. ebenina and E. nigra but is much smaller with more convex whorls than the former and has a higher spire and thinner, more conical shell than E. nigra. It also has a shell similar to the New Zealand species E. stewartiana Ponder, 1965, but is darker and slightly larger. The sym- patric E. denticula has a smaller, broader shell that is, at most, pale grey in color. Eatoniella picea dif- fers from other similar taxa, except E. nigra, in having a brown operculum and the radula is also unusual in having a blunt median cusp on the cen- tral teeth, whereas the equivalent cusp on the lateral teeth is sharp. It also shares this latter character with E. glomerosa n. sp., but that species differs markedly in shell characters. DISTRIBUTION. Southern Chile and Tierra del Fuego in algae and among rocks. Mainly intertidal with occasional specimens to 15 m. Empty shells to 50 m. Common. Eatoniella ( Eatoniella ) nigra (Orbigny, 1840) Figures 5F, 7E, 9C, 10A Paludestrina nigra Orbigny, 1840: 387, pi. 75, figs. 16-18 (36 syntypes, BMNH 1854.12.4.351, in- 14 ■ Contributions in Science, Number 445 eluding one very worn specimen of “ Potamoli - thus" sp.; Arica, Tarapaca Prov., Chile). Orbigny, 1854: 31. Eatoniella ( Eatoniella ) latina Marincovich, 1973: 26, figs. 51, 57, 58 (Holotype, Iquique, Tarapaca Prov., Chile, intertidal; LACM 1587, see Marin- covich (1973) for details of paratypes). MATERIAL EXAMINED. Types. Syntypes of P. ni- gra. 6 paratypes of E. latina, AMS C. 167464. Iquique, Tarapaca Prov., Chile. Additional Material Examined. Northern Chile: 6A 64-16 [many(d)]; 70-66 [1(d)]; 75-12 [many(d)]; AMS C. 167467 [20(d)]. 6B 75-10 [many]; AMS C.167461 [17]. 7 75-21 [2(d)]. 8A 75-17 [many]; AMS C.167463 [l( + many d)]. 8B 75-19 [many]; AMS C.167465 [20], 8C 75-15 [many]; 75-20 [many]; AMS C.167460 [19]. 9 75-25 [1]. 10 75-28 [6]; AMS C. 167466 [1]. 12 75-33 [4(d)]; AMS C.167462 [5(d)], 13 75-37 [7]. Southern Chile: 15 75-41 [7(d)]. 17 73-75 [many(d)]. 18 73-74 [Id]. 19 73-73 [many]. 20 73- 72 [many(d)]. 23 73-70 [5]. 24 75-48 [many]; 75-49 [many]. Tierra del Fuego: 25 73-69 [many(d)]. USNM 30A H 654 [1(d)]. 31A 71-274 [1(d)]. 33G 73-66 [many(d)]. 330 71- 310 [20(d)]. (All material LACM unless otherwise indi- cated.) DIAGNOSIS. Shell (Figs. 5F, 10A). Minute (max- j imum length 1.8 mm), ovoid, solid, with about 3.3- 3.5 teleoconch whorls. Spire and whorls with very lightly convex outlines; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth with faint prosocline growth lines. Proto- conch smooth of 1.2-1. 4 whorls. Aperture oval, very weakly angled posteriorly, with sharp peri- stome, lacking external varix. Inner lip moderately narrow, outer lip strongly prosocline. Umbilical chink absent. Periostracum very thin, transparent. Color black or grey, often with reddish tinge, paler near growing edge. Dimensions. 1 SL/ SL/ SL SW SW AL AL TW PW PD Figured 1.77 1.04 1.70 — — — — — syntype (Fig. 10 A) Paratypes of 1.78 1.11 1.61 0.65 2.74 3.3 1.3 0.28 E. latina 1.58 0.96 1.65 0.56 2.58 3.4 1.2 0.26 1.58 0.99 1.60 0.61 2.56 3.5 1.3 0.26 1.71 1.10 1.61 0.67 2.57 3.4 1.3 0.29 1.76 1.10 1.66 0.65 2.72 3.4 1.2 0.27 Sta. 75-20 Fig. 5F 1.69 0.99 1.70 0.63 2.68 3.5 1.3 0.25 1.71 1.04 1.65 0.66 2.58 — — — 1.59 0.94 1.69 0.60 2.66 3.4 1.3 0.27 1.77 1.06 1.67 0.68 2.58 3.5 1.2 0.24 1.79 1.05 1.70 0.66 2.69 3.3 1.3 0.27 1.81 1.05 1.72 0.68 2.64 3.5 1.4 0.25 1.62 0.96 1.68 0.62 2.60 3.5 1.2 0.20 1.72 1.03 1.64 0.62 2.77 3.4 1.2 0.26 1.69 1.03 1.64 0.62 2.72 3.4 1.2 0.21 1.66 1.02 1.63 0.59 2.82 3.5 1.2 0.24 Operculum (Fig. 7E). Color yellow with brown markings or brown; oval, angled posteriorly, outer edge more strongly convex than inner edge. Peg very stout, curved. Radula (Fig. 9C). Central teeth with cusp formula I Ponder and Worsfold: Review of Rissoiform Gastropods 3 + 1 + 3, median cusp elongate, sharp (somewhat worn in figured radula, typically more than 2x length of adjacent cusps). Lateral teeth with cusp formula 2-3 + 1 + 2-3, primary cusp elongate, sharp. Inner marginal teeth with cusp formula 3-4 + 1+? cusps, outermost obscured in mounts. Outer mar- ginal teeth with about 6 small, sharp cusps, out- ermost largest (based on 2 radulae). Animal. Unknown. REMARKS. Comparison of paratypes of Eaton- iella latina with the types of Ealudestrina nigra show them to have identical shells and we regard them as conspecific. Eatoniella nigra is shorter and more ovoid in shape than other dark-colored South American species and has a thicker shell. The shell of the most similar South American species, E. pic- ea, is contrasted above. Eatoniella nigra has an operculum with similar coloration to that of E. picea, but in the latter species the opercular peg is much narrower. A somewhat similar species from New Zealand, E. olivacea, has a more elongate, larger shell. The southern Australian E. melanochroma (Tate, 1899) has a very similar shell to that of E. nigra, although it is slightly thinner, with more convex whorls, and the radula and operculum in the two species are also similar. Da Silva and Davis (1983) did not include this species in their review of Orbigny’s species of Palu- destrina. The South African Rissoa ( =Eatoniella ) nigra Krauss (1848) is a secondary homonym of this spe- cies. A replacement name, Eatoniella afronigra n. sp. is provided, as none of the other South African taxa appear to be synonymous. This species also has a shell very similar to that of E. nigra but is slightly broader and usually has more convex whorls. DISTRIBUTION. Northern Chile to Tierra del Fuego in algae and among rocks. Mainly intertidal; some specimens to 15 m. Empty shells to 50 m. Common. Eatoniella ( Eatoniella ) castanea n. sp. Figures 9D, 1 1 F, H, 12A ETYMOLOGY. Castanea — Latin. Chestnut col- or (brown). Refers to the shell color. MATERIAL EXAMINED. Types. Holotype, LACM 2664, 34 -paratypes, LACM 2665; 5 paratypes, AMS C. 167418. 31B 71-277. NW arm Bahia York, Isla de los Estados, Tierra del Fuego, Argentina. 54°47.15'S, 64°17.9'W, intertidal rocks, Sta. 71-2-22, USARP-SOSC- R/V Hero Cr.712, 5 May 1971. DIAGNOSIS. Shell (Fig. 1 1 F, H). Minute (max- imum length 1.7 mm), ovoid, moderately thin, with 2. 2-3.0 teleoconch whorls. Spire with lightly con- vex outlines, whorls lightly convex; periphery of last whorl rounded. Sutures impressed, simple. Tel- eoconch smooth, with faint axial growth lines and faint spirals. Protoconch smooth, of about 1.2-1. 5 whorls. Aperture oval, with sharp peristome, lack- ing external varix. Inner lip narrow, outer lip mod- erately prosocline. Umbilical chink minute. Peri- ostracum very thin, transparent. Color reddish- brown, whitish near growing edge. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 1.31 0.85 1.53 0.54 2.44 3.0 1.2 0.27 Paratypes Fig. 1 1 F 1.66 1.00 1.65 0.63 2.63 2.5 1.4 0.37 1.42 0.93 1.53 0.60 2.37 2.4 1.2 0.35 1.37 0.91 1.51 0.60 2.33 2.4 1.3 0.35 1.44 0.92 1.57 0.62 2.33 2.4 1.4 0.35 1.50 0.96 1.55 0.60 2.55 2.4 1.5 0.41 1.36 0.91 1.49 0.57 2.40 2.2 1.5 0.39 1.18 0.96 1.43 0.56 2.12 2.2 1.3 0.33 1.36 0.92 1.48 0.56 2.44 2.3 1.4 0.35 1.36 0.90 1.51 0.55 2.49 2.3 1.5 0.35 1.25 0.86 1.46 0.55 2.29 2.3 1.5 0.38 Operculum (Fig. 12A). Yellow, oval, slightly more strongly angled anteriorly than posteriorly. Peg moderately stout, curved. Radula (Fig. 9D). Central teeth with cusp formula 2+1+2, median cusp moderately large, trowel- shaped. Lateral teeth with cusp formula 2+1 + 3, primary cusp rather narrow, sharply pointed. Inner marginal teeth with cusp formula 2(?) + 1 + 1 , pri- mary cusp sharp, triangular. Outer marginal teeth with at least 5 small cusps, outermost largest (based on 2 radulae). Animal. Unknown. REMARKS. Eatoniella castanea is distinguished from other South American species by its small, oval, red-brown shell. Faded shells of E. nigra are sometimes reddish in color but are larger and broader and have more flattened whorls. Eatoniella nigra also differ in having a broader opercular peg and a brown operculum and in radular details. Ea- toniella subrufescens (E.A. Smith, 1875) from Ker- guelen Island is similar to E. castanea but has a smaller, narrower shell. Eatoniella argentinense Castellanos and Fernandez, 1972, from Punta Loma, Argentina, is somewhat similar as far as can be determined from the description and figures (Cas- tellanos and Fernandez, 1972a) but is white in color. DISTRIBUTION. Tierra del Fuego on intertidal algae. Uncommon. Eatoniella ( Eatoniella ) glomerosa n. sp. Figures 6E, 7F, H, I, 11A-E, 13A-D ETYMOLOGY. Glomerosa — Latin. Like a ball, round. Refers to the shell shape. MATERIAL EXAMINED. Types. Holotype, LACM 2666, 230 paratypes, LACM 2667; 8 paratypes, AMS C. 167419. 32H 71-287. Puerto Cook, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.25'S, 64°02.3'W, in- tertidal rocks, Sta. 71-2-37, USARP-SOSC-R/V Hero Cr.712, 17 May 1971. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods® 15 Figure 11. Shells of species of Eatoniella. A-E, Eatoniella glomerosa n. sp.; A, holotype, length 1.26 mm; B, C, E, Sta. 71-283, showing variation; B, length 1.04 mm; C, length 1.03 mm; E, length 1.48 mm; D, Sta. 73-73, length 1.16 mm. F, H, Eatoniella castanea n. sp.; F, paratype, length 1.66 mm; FI, holotype, length 1.31 mm. G, Eatoniella ( Albosabula ) mcleani n. sp., holotype, length 0.96 mm. I, Pupatonia magellanica n. sp., holotype, length 1.01 mm. J, Pupatonia cf. atoma Ponder, NMNZ, 67004, 60 m, off Taiere, Dunedin, New Zealand, length 0.76 mm. Scale bars: A-D, G, I, J, 200 |im; E, 400 jam; F, H, 500 /xm. 16 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods Figure 12. Opercula of Eatoniellidae and Rissoidae. All views inner side. A, Eatoniella castanea n. sp., paratype. B, Eatoniella ( Albosabula ) mcleani n. sp., paratype. C, Pupatonia magellanica n. sp., paratype. D, Powellisetia microlirata n. sp., paratype. E, Pupatonia cf. atoma Ponder, 60 m, off Taiere, Dunedin, New Zealand. F, “Owobtf” lacuniformis n. sp., paratype. Scale bars: A-C, E, 100 jum; D, F, 200 jum. Additional Material Examined. Northern Chile: 9 75- 25 [3(d)], 10 75-28 [11(d)]; AMS C.167459 [1(d)]. 11 75- 30 [1(d)]. 13 75-37 [4(d)]. Southern Chile: 15 75-41 [8(d)]. 16 75-46 [1(d)]. 17 73-75 [5(d)]. 19 73-73 [many(d)]. 24 75-49 [1]. Tierra del Fuego: 28 71-270 [5(+2d)]; 71-271 [7]. 31A 71-345 [1(d)]. 31B 71-276 [6]; 71-277 [30]. 31C 71-281 [28], 31D 71-283 [many], 32A 71-273 [many]. 32B 71-295 [3(d)]. 32D 71-293 [7]. 32E 71-289 [many]; 71-290 [4(d)]. 32F 71-326 [1(d)]. 32G 71-311 [3]. 32H 71-286 [1]; 71-323 [1], 33G 73-66 [13]. 39 71-268 [19]. (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Figs. 6E, 11A-E). Minute (maximum length 1.5 mm), globular, thin, trans- lucent, with about 1. 3-3.0 teleoconch whorls. Spire with convex outlines, whorls strongly convex; pe- riphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth and rather glossy, with faint prosocline growth lines. Protoconch (Fig. 6E) smooth, of 1.2-1. 7 whorls. Aperture ovoid, angled posteriorly, with sharp peristome, lacking external varix. Inner lip moderately broad, upper part de- tached from last whorl; outer lip strongly proso- cline, more strongly convex than inner lip. Umbil- icus moderate to distinct. Periostracum very thin, transparent. Color white. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 1.26 0.98 1.29 0.65 1.96 2.3 1.5 0.35 Paratypes 1.16 0.87 1.33 0.61 1.90 3.0 1.5 0.23 1.13 0.77 1.47 0.50 2.26 2.9 1.4 0.23 1.10 0.81 1.36 0.58 1.90 2.5 1.4 0.27 1.10 0.80 1.38 0.52 2.12 2.5 1.4 0.26 1.19 0.87 1.37 0.62 1.92 2.5 1.3 0.27 1.23 0.99 1.24 0.62 1.98 2.5 — — Sta. 71-283 Fig. 11B 1.04 0.83 1.24 0.56 1.87 1.3 1.6 0.44 Fig. 11C 1.03 0.82 1.25 0.58 1.78 1.3 1.6 0.41 Fig. 11E 1.48 1.18 1.25 0.74 2.00 — — — 1.02 0.92 1.10 0.58 1.76 1.7 1.3 0.37 1.03 0.80 1.28 0.57 1.81 1.4 1.6 0.41 1.16 0.93 1.24 0.57 2.04 1.6 1.6 0.42 1.03 0.84 1.21 0.57 1.81 1.4 1.4 0.38 1.11 0.87 1.28 0.55 2.03 1.4 1.7 0.41 1.17 0.95 1.22 0.55 2.14 1.5 1.6 0.41 1.16 0.82 1.40 0.53 2.16 1.6 1.5 0.43 Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 17 Figure 13. Radulae of Eatoniella species. A-D, Eatoniella glomerosa n. sp.; A, Sta. 71-283; B, paratype; C, Sta. 71- 270; D, Sta. 71-283. E, F, Eatoniella ( Albosabula ) mcleani n. sp., paratypes. Scale bars: A-D, 10 /am; E, F, 5 /am. 0.98 0.81 1.21 0.53 1.84 1.4 1.5 0.39 1.48 1.08 1.37 0.65 2.26 — — — 1.41 1.04 1.36 0.67 2.09 1.5 1.6 0.44 Sta. 73-73 Fig. 11D 1.16 0.90 1.29 0.56 2.08 2.2 1.2 0.31 1.07 0.86 1.23 0.53 2.00 2.2 1.2 0.27 1.14 0.87 1.32 0.54 2.10 2.2 1.2 0.31 1.19 0.89 1.34 0.54 2.18 2.3 1.2 0.31 1.21 0.88 1.38 0.44 2.22 2.3 1.2 0.27 1.20 0.93 1.29 0.56 2.15 2.3 1.3 0.30 1.32 0.96 1.37 0.58 2.28 — — — 1.23 0.92 1.37 0.59 2.09 2.3 1.3 0.31 1.26 0.96 1.31 0.57 2.23 2.3 1.2 0.28 1.36 1.02 1.34 0.60 2.27 2.3 1.2 0.28 Operculum (Fig. 7F, Fi, I). Pale yellow, oval, weakly angled posteriorly, outer edge much more convex than inner. Peg moderately narrow, curved. Radula (Fig. 13A-D). Central teeth with cusp formula 3 + 1+3, median cusp large, spatulate. Lat- eral teeth with cusp formula 3 + 1 + 3, primary cusp small, narrow, sharp. Inner marginal teeth with cusp formula 3 + 1 + 1 (—2), primary cusp large, sharp. Outer marginal teeth with about 6 small cusps, out- ermost largest (based on 6 radulae). Animal. Usually unpigmented, sometimes with darker visceral coil. REMARKS. The shell of this species is variable (see measurements) in size, shape, number of whorls, and umbilical size and may possibly represent a species complex. Specimens from the north of its range generally have a larger umbilicus. There are no other globular species of Eatoniella known from South America, although several species from other southern areas have similar shells. The New Zea- land E. pullmitra Ponder, 1965, has a larger shell aperture, and the Australian E. sbepherdi Ponder and Yoo, 1977, has a more elongate shell. The Kerguelen Island E. hyalina Thiele, 1912, is similar, but smaller. A few specimens, including some in the paratype series, have narrower shells somewhat similar in shape to the New Zealand E. aterviseralis Ponder, 1965. This shell form, however, intergrades with the broader, typical form and agrees with it in all other respects. DISTRIBUTION. Central Chile to Tierra del Fuego among rocks and algae. Mainly intertidal; some specimens and dead shells to 15 m. Common. 18 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods Subgenus Albosabula Ponder, 1965 Species in this subgenus were previously known only from New Zealand (Ponder, 1965a) and Tas- mania (Ponder and Yoo, 1977a). They are distin- guished by their small, conical shells that are never colored and in having the midbasal margin of the central teeth of the radula markedly produced ven- trally. Ponder and Yoo (1977a) suggested that Ris- soa georgiana Martens and Pfeffer, 1886, was pos- sibly a member of this group, but subsequent examination of that species showed that it is a mem- ber of the rissoid genus Onoba (Ponder, 1983a). Eatoniella ( Albosabula ) mcleani n. sp. Figures 11G, 12B, 13E, F ETYMOLOGY. Named for Dr. James McLean of the Natural History Museum of Los Angeles County as a small recognition of his efforts in col- lecting the material on which this report is largely based. MATERIAL EXAMINED. Types. Holotype, LACM 2668, 295 paratypes, LACM 2669; 11 paratypes, AMS C. 167420. 10 75-28. Los Molles, Aconcagua Prov., Chile. 32°14'S, 71°32'W, intertidal, Sta. 19, J.H. McLean, 16- 18 Oct. 1975. Additional Material Examined. Northern Chile: 9 75- 25 [23(d)]; AMS C.167474 [3]. 11 75-30 [3(d)]; AMS C. 167473 [1]. Southern Chile : 15 75-41 [19]. 16 75-46 [3(d)]. 19 73-73 [17(d)]. (All material LACM unless oth- erwise indicated.) DIAGNOSIS. Shell (Fig. 11G). Minute (maxi- mum length 1.0 mm), ovate, thin, translucent, with 2. 5-2. 9 teleoconch whorls. Spire with lightly con- vex outlines, whorls lightly convex; periphery of last whorl rounded. Sutures impressed, simple. Tel- eoconch smooth and glossy with moderately prom- inent prosocline growth lines. Protoconch smooth of 1. 1-1.3 whorls. Aperture oval, angled posteri- orly, with sharp peristome, lacking external varix. Inner lip moderately broad, outer lip slightly pro- socline, slightly more convex than inner. Umbilical chink minute. Periostracum very thin, transparent. Color pale brown when fresh, empty shells often white. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 0.96 0.60 1.61 0.42 2.31 2.5 1.3 0.20 Paratypes '0.93 0.56 1.67 0.39 2.35 2.8 1.2 0.16 0.95 0.56 1.71 0.39 2.40 2.8 1.2 0.17 0.91 0.55 1.67 0.41 2.24 2.9 1.2 0.19 0.99 0.61 1.63 0.41 2.45 2.9 1.2 0.18 0.95 0.57 1.70 0.39 2.41 2.9 1.1 0.17 0.86 0.57 1.60 0.35 2.42 2.7 1.3 0.19 0.96 0.59 1.64 0.41 2.37 2.5 1.3 0.21 0.99 0.62 1.60 0.43 2.33 2.5 1.3 0.21 0.97 0.59 1.65 0.42 2.33 2.5 1.2 0.19 Operculum (Fig. 12B). Pale yellow, oval, more strongly angled posteriorly than anteriorly, and more strongly convex on outer edge. Peg stout, slightly curved, and flared at extremity. Radula (Fig. 13E, F). Central teeth with cusp formula 2 + 1 + 2, median cusp long, narrow, point- ed; with midbasal tongue. Lateral teeth with cusp formula 2 + 1+2-3, all cusps pointed. Inner mar- ginal teeth with cusp formula 4 + 1+2, all cusps pointed, primary cusp slightly larger than adjacent inner cusp, outer adjacent cusps much smaller. Out- er marginal teeth with 5-6 small cusps. (Note: Cusps somewhat worn in illustrated specimen; the second specimen examined has sharper cusps but is a very poor mount.) Animal. Unpigmented. REMARKS. This is the only known species re- ferrable to the subgenus Albosabula outside Aus- tralasia. It can be distinguished from other South American species of Eatoniella by its small, oval, translucent shell. There are three species assigned to Albosabula in New Zealand (Ponder, 1965a). Eatoniella (A.) lampra (Suter, 1908) and E. (A.) poutama (E. Smith, 1962) both have larger shells, whereas the shell of E. (A.) rakiura Ponder, 1965, is broader. The only Australian species, E. (A.) pellucida (Tate and May, 1900), has a slightly larger, broader shell. DISTRIBUTION. Central and southern Chile from intertidal to 17 m. Moderately common. Genus Pupatonia Ponder, 1965 This genus has previously only been recorded from southern New Zealand and the New Zealand sub- Antarctic islands (Ponder, 1965a; Powell, 1979). It was distinguished from other genera in the family by the pupiform shell. The familial position of this genus has never previously been confirmed by the examination of the radula and operculum because all known specimens were empty shells. However, recently, a single dried animal of a New Zealand species, P. cf. atoma Ponder, 1965, was found in the NMNZ collections. The radula and operculum were extracted from this specimen and are figured here for comparison with the South American spe- cies (Figs. 1 1J, 12E, 14D, F). The operculum (Fig. 1 2C, E) in both taxa is very similar to that of species of Eatoniella. The radula, for which only poor mounts were available for both species, is also gen- erally similar to that of species of Eatoniella but differs in having markedly less prominent basal pegs on the central teeth as well as having smaller, more equal-sized cusps on the inner marginal teeth than seen in most species of Eatoniella. Pupatonia magellanica n. sp. Figures 111, 12C, 14A-C ETYMOLOGY. Named after the Magellanic Province of Chile. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 19 Figure 14. Radulae of Pupatonia species. A-C, Pupatonia magellanica n. sp., paratype, detail of central teeth. D-F, Pupatonia cf. atoma Ponder, 60 m, off Taiere, Dunedin, New Zealand. Scale bars: A-D, F, 2 /urn ; E, 5 nm. MATERIAL EXAMINED. Types. Holotype, LACM 2670, 3 paratypes, LACM 2671. 32 71-273. Bahia Cross- ley, Isla de los Estados, Tierra del Fuego, Argentina. 54°46.2'S, 64°42.7'W, intertidal, rocks and mussel beds, Sta. 71-2-16, USARP-SOSC-R/V Hero Cr.712, 27 Apr. 1971. 5 paratypes, LACM, 2, AMS C.167421. 17 73-75. Isla Westhoff, Chiloe Prov., Chile. 43°54'S, 73°43.5'W, 23 m, P. Dayton (R/V Hero), 25 May 1973. Additional Material Examined. Northern Chile: 9 75- 25 [1(d)]. Southern Chile: 14 75-43 [6(d)]. 19 73-73 [2(d)]. 22B USNM E 958 [5(d)], Tierra del Fuego: 28 71-270 [1(d)]. 33D 71-352 [3(d)]. (All material LACM unless oth- erwise indicated.) DIAGNOSIS. Shell (Fig. 111). Minute (maximum length 1.1 mm), cylindrical-pupoid, thin, translu- cent, with 2. 6-3.1 teleoconch whorls. Whorls strongly convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth and glossy with faint growth lines. Protoconch of 1.3- 1.5 whorls, apparently smooth (eroded in available material). Aperture oval, protruding, with sharp peristome, lacking external varix. Inner lip mod- erately broad and separated from last whorl, outer lip orthocline. Umbilical chink absent. Periostra- cum very thin, transparent. Color white. Dimensions. 20 ■ Contributions in Science, Number 445 SL SW SL/ SW AL SL/ AL TW PW PD Holotype 1.01 0.53 1.91 0.36 2.84 2.8 1.5 0.17 Paratypes 0.99 0.51 1.94 0.35 2.84 2.7 1.3 0.17 0.91 0.49 1.86 0.33 2.76 2.8 1.4 0.18 Sta. 75-43 0.98 0.51 1.92 0.36 2.69 3.0 1.4 0.16 0.91 0.48 1.88 0.33 2.77 3.1 1.5 0.17 1.05 0.57 1.85 0.37 2.83 3.0 1.4 0.17 0.91 0.50 1.83 0.32 2.83 2.6 1.4 0.17 Operculum (Fig. 12C). Oval, angled posteriorly, and rounded anteriorly. Inner and outer edges equally convex. Peg stout, curved. Radula (Fig. 14A-C). Central teeth with cusp formula 2 + 1+2, median cusp pointed. Lateral teeth with cusp formula 2 + 1+3, primary cusp pointed. Inner marginal teeth with 5-6 cusps about equal in size. Outer marginal teeth with about 4 cusps. Teeth apparently rather soft, causing cusps to fold (based on 1 specimen). Animal. Unpigmented. REMARKS. The shape of the shell of species of Pupatonia is distinctive. Only four species have been assigned to this genus, all from New Zealand. Of these, P. magellanica is most similar in shape Ponder and Worsfold: Review of Rissoiform Gastropods to P. minutula (Powell, 1933), which differs in its slightly larger shell and in having weak spiral sculp- ture. Pupatonia gracilispira (Powell, 1933) is about the same size as the new species but is smooth and has a narrower spire. DISTRIBUTION. Central Chile to Tierra del Fuego; live-collected specimens all intertidal. Shells found to 100 m. Uncommon. Family CINGULOPSIDAE Members of this family have trochiform to elon- gate-conic shells that, like the other members of the superfamily, have an inner “chitinous” layer (Ponder, 1988). Like eatoniellids, they have a pegged operculum and males are aphallate but differ in radular characters and, markedly, in anatomical de- tails. Notable differences include the lack of a style sac in the stomach and closed pallial genital ducts. The operculum of cingulopsids never has an opaque muscle insertion area. The cingulopsid genera have been reviewed by Ponder and Yoo (1980), with some modifications by Ponder (1983a). Genus Skenella Pfeffer, 1886 This genus was included in the Eatoniellidae by Ponder (1965a) and Ponder and Yoo (1977a) but was transferred to the Cingulopsidae by Ponder (1983a) following examination of topotype mate- rial of the type species. It is distinguished from the other genera included in the family by the radula having simple central teeth, the operculum having a small peg and lacking a ridge on the inner surface, and the columella of the shell having a weak bulge. Skenella hallae n. sp. Figure 15A, F ETYMOLOGY. Named for Jane Hall, who was initially involved with this project, taking many of the SEM photographs and measuring many of the specimens. MATERIAL EXAMINED. Types. Holotype, LACM 2672, 8 paratypes, LACM 2673, 1 paratype, AMS C. 167422. 25 73-69. Punta Valparaiso, Canal Cockburn, Magellanes Prov., Chile. 54°22.2'S, 71°21.7'W, 17 m, P. Dayton (R/V Hero), 17 May 1973. Additional Material Examined. Tierra del Fuego: LACM 33M 71-263 [16(d)], DIAGNOSIS. Shell (Fig. 15A). Minute (maxi- mum length 1.2 mm), ovate, moderately thick, with 2. 3-3.0 teleoconch whorls. Spire with convex out- lines, whorls moderately convex; periphery of last whorl rounded. Sutures impressed, simple. Teleo- conch smooth except for faint growth lines. Proto- conch apparently smooth, of 1.0-1. 2 whorls. Aper- ture oval, weakly angled posteriorly, with sharp peristome. Inner lip moderately broad with an in- distinct swelling on columella, outer lip moderately to strongly prosocline. Umbilical chink minute. Color reddish-brown. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 0.88 0.64 1.37 0.39 3.28 2.3 1.2 0.15 Paratypes 0.96 0.73 1.31 0.43 2.22 2.5 1.0 0.14 0.98 0.71 1.38 0.43 2.27 2.5 1.0 0.14 0.97 0.71 1.36 0.44 2.19 2.8 1.1 0.13 0.96 0.72 1.33 0.43 2.22 2.5 1.1 0.13 1.24 0.81 1.53 0.49 2.50 3.0 1.0 0.17 0.97 0.73 1.32 0.43 2.25 2.7 1.2 0.14 0.97 0.73 1.32 0.43 2.25 2.8 1.2 0.14 1.01 0.74 1.36 0.45 2.24 2.3 1.2 0.15 1.02 0.73 1.40 0.43 2.37 2.8 1.2 0.15 Operculum (Fig. 15F). Pale yellow, oval, anterior and posterior ends moderately angled, inner edge weakly angled. Peg short, narrow, almost straight. Radula. Present but too poorly prepared to de- scribe from available material. Animal. Unknown. REMARKS. This is the only known South Amer- ican Skenella and can be distinguished from other rissoiform taxa with similar shells by the small swelling on the columella. The shell is most similar to Eatoniella castanea but is smaller and narrower. DISTRIBUTION. Southern Chile and Tierra del Fuego; living specimens collected from 17 m; empty shells from 30 m. Uncommon. Genus Eatonina Thiele, 1912 In this genus the columella of the shell is simple, the operculum has a large peg and a heavy ridge on its inner surface, and the central radular teeth usu- ally have cusps. Subgenus Mistostigma Berry, 1947 Only two species are known from this subgenus, the type species from California (E. (M.) albida Carpenter, 1864) and the one described below. Ponder and Yoo (1980) described the radula and operculum of the type species. The subgenus is distinguished by members of the typical subgenus on radular details (Ponder and Yoo, 1980). Eatonina ( Mistostigma ) fusca (Orbigny, 1840) Figures 10C, 15B, E Paludestrina fusca Orbigny, 1840: 387, pi. 75, figs. 13-15 (2 syntypes, BMNH 1854.12.4.348; Arica, Tarapaca Prov., Chile). Orbigny, 1854: 31. Eatonina ( Saginofusca ) atacamae Marincovich, 1973: 27, figs. 52, 59, 60 (Holotype, LACM 1589, many paratypes, LACM 1590; 20 paratypes, AMS C. 162624). Eatonina ( Mistostigma ) atacamae : Ponder and Yoo, 1980: 32, fig. 14h, i. MATERIAL EXAMINED. Types. Syntypes of P. fus- ca; paratypes of E. atacamae [20]. Additional Material Examined. Northern Chile: 6A 64-16 [many SEM]; 75-12 [4(d)]; AMS C.167472 [1(d)]. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 21 Figure 15. Shells, radulae, and opercula of Cingulopsidae. A, F, Skenella hallae n. sp.; A, holotype, shell, length, 0.88 mm; F, paratype, operculum, inner side. B, E, Eatonina fusca (Orbigny) Sta. 64-16; B, shell, length, 1.45 mm; E, paratype, Iquique, Chile, lateral view of protoconch. C, D, G, H, Skenella wareni n. sp.; C, holotype, shell, length 1.64 mm; D, G, H, paratypes; D, protoconch microsculpture; G, radula; H, operculum, inner side. Scale bars: A, 200 /urn; B, C, 500 /urn; D, E, 50 /urn; F, H, 100 /u m. 22 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods 6B 75-10 [many]; AMS C.167469 [3(+lld)]. 7 75-21 [9]. 8A 75-17 [many]; AMS C.167468 [3( + 13d)]. 8B 75-19 [3(d)]; AMS C.167471 [1]. 8C 75-15 [3(d)]; 75-20 [8]; AMS C. 167470 [1]. 9 75-25 [1(d)]. 10 75-28 [2], (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Figs. 10C, 15B, E). Minute (maximum length 1.8 mm), ovate-conic, moderate- ly thin, opaque, with 2. 8-3.3 teleoconch whorls. Spire with lightly convex outlines, whorls moder- ately to strongly convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth with faint prosocline growth lines and ex- tremely fine spiral striae. Protoconch (Fig. 15E) with close, fine spiral grooves, of about 1.3- 1.4 whorls. Periostracum thin, transparent. Aperture oval to almost round, very weakly angled posteriorly, with sharp peristome. Inner lip narrow, detached from parietal wall; outer lip strongly prosocline. Umbil- icus distinct. Color typically reddish-brown, occa- sionally pale or white. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Figured syntype (Fig. 10C) 1.25 1.10 1.14 — — — — — Paratypes of 1.52 1.21 1.25 0.73 2.07 3.3 1.4 0.24 E. atacamae 1.38 1.07 1.28 0.68 2.04 3.1 1.4 0.22 1.17 1.02 1.15 0.66 1.77 2.8 1.3 0.25 Sta. 64-16 Fig. 15B 1.45 1.13 1.29 0.73 1.98 3.0 1.3 0.21 Sta. 75-10* 1.57 1.20 1.30 0.72 2.18 — — — 1.61 1.24 1.30 0.76 2.11 — — — 1.48 1.26 1.18 0.70 2.12 — — — 1.54 1.13 1.36 0.70 2.20 — — — 1.75 1.21 1.44 0.74 2.36 — — — 1.52 1.19 1.27 0.69 2.20 — — — 1.56 1.23 1.27 0.74 2.10 — — — 1.54 1.16 1.32 0.77 1.99 — — — 1.48 1.11 1.34 0.71 2.09 — — — 1.62 1.19 1.36 0.75 2.16 - — — _ * Protoconch and upper whorls of shells from this station badly eroded. Operculum. Yellow, oval to rectangular. Peg broad and short with thickened ridge running from it to opposite end of operculum (figured by Marin- covich, 1973: fig. 60). Radula. Central teeth with dorsal edge with 3 rudimentary cusps; face of teeth with 4 prominent cusps. Lateral teeth with 4 large, triangular cusps, outermost smallest. Inner marginal teeth with 4 cusps, third largest, forth (outermost) smallest. Out- er marginal teeth very short, with 3 cusps (figured by Marincovich, 1973: fig. 59, and Ponder and Yoo, 1980: fig. 14h, i). Animal. Unknown. REMARKS. This species is much larger and more globular than Skenella hallae and lacks the bulge on the columella. Shell shape is somewhat similar to that of Eatoniella glomerosa but is immediately distinguished by its red-brown color. DISTRIBUTION. Northern and central Chile; mainly intertidal, occasional animals and empty shells to 30 m. Common. Superfamily RISSOOIDEA (=Truncatelloidea, ICZN Opin. 1664) Family RISSOIDAE This family is distinguished from the foregoing by the shell lacking an inner “chitinous” layer, the males are phallate, and there are also many other anatomical differences (see Ponder, 1985a, 1988). Some rissoid genera have a peg on the inner side of the operculum, but this is not the case in any of the genera covered below. The generic classification adopted here follows Ponder (1985a). Genus Pusillina Monterosato, 1884 This genus is similar to Rissoa in anatomy and shell morphology, differing mainly in details of anatomy, notably the possession of a pallial prostate gland. Subgenus Haurakia Iredale, 1915 Members of this subgenus usually have an anterior excavation of the anterior part of the aperture and the radula has a tongue-like midventral edge on the central teeth. Species of Pusillina ( Haurakia ) were known from all of the southern continents and New Zealand except South America (Ponder, 1985a). The species described below is tentatively assigned to Pusillina ( Haurakia ) pending exami- nation of the animal. No species in this group were recorded from the Antarctic-sub-Antarctic by Pon- der (1983a), although some are known from the New Zealand sub-Antarctic islands (Powell, 1979). Pusillina ( Haurakia ) averni n. sp. Figure 16A, B ETYMOLOGY. Named for Geoff Avern, who prepared, mounted, and photographed with the SEM some of the material used in this paper. MATERIAL EXAMINED. Types. Holotype, LACM 2674, 5 paratypes, LACM 2675; 1 paratype, AMS C. 167455. 30E 71-339. NW arm Bahia York, Isla de los Estados, Tierra del Fuego, Argentina. 54°47.2'S, 64°18.4'W, 38 m, Sta. 891, USARP-SOSC-R/V Hero Cr.71 5, 1 Nov. 1971. Additional Material Examined. Southern Chile: 25 73- 69 [1(d)]. Tierra del Fuego: 30H 71-332 [1(d)]. 33 A USNM H 656 [2(d)]. 33E 71-351 [2(d)]. (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Fig. 16 A, B). Small (maxi- mum length 2.3 mm), elongate-conic, thin, trans- lucent when fresh; with 2. 7-3. 2 teleoconch whorls. Spire with lightly convex to straight outlines, whorls moderately convex; periphery of last whorl round- ed. Sutures impressed, simple. Teleoconch with about 6-20 fine, spiral striae on penultimate whorl Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 23 Figure 16. Shells and protoconchs of species of Manzonia and Pusillina. A, B, Pusillina ( Haurakia ) averni n. sp., holotype; A, shell, length 2.05 mm; B, lateral view of protoconch. C, Pusillina ( Haurakia ) cf. averni, Sta. 71-344, shell, length, 2.13 mm. D, E, Manzonia ( Alvinia ) limensis n. sp., holotype; E, shell, length, 2.03 mm; D, lateral view of protoconch. Scale bars: A-C, 250 yum; D, E, 100 yum ; F, G, 25 yu m. and 16-30 on last whorl and base. Protoconch (Fig. 16B) of 1.1-1. 3 whorls, sculptured with irregularly shaped pits arranged more or less spirally. Aperture oval, weakly angled posteriorly. Inner lip narrow, attached to parietal wall in upper portion; outer lip orthocline with small apertural varix. Umbilical chink small to moderate. Periostracum very thin, transparent. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD PS BS Holotype 2.05 1.19 1.72 0.80 2.58 3.2 1.1 0.34 20 30 Paratypes 2.29 1.41 1.63 0.80 2.88 3.2 1.2 0.35 2.12 1.29 1.64 0.80 2.66 2.9 1.2 0.40 Sta. 71-351 1.98 1.25 1.59 0.82 2.42 2.7 1.2 0.48 24 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods 2.17 1.34 1.62 0.89 2.43 2.7 1.3 0.50 Sta. 71=332 2.11 1.31 1.61 0.89 2.36 2.8 1.2 0.44 11 16 Operculum, radula, and animal unknown. REMARKS. This species is tentatively included here, pending examination of the radula. It most closely resembles Pusillina ( Haurakia ) infecta from New Zealand but differs in having a white (not brown) protoconch, more convex teleoconch whorls, stronger axials, and stronger spirals on the base. This species resembles a few taxa included in Onoba but has finer sculpturing than any similarly sized South American species. DISTRIBUTION. Tierra del Fuego. Known only from empty shells; uncommon in 40-900 m. Pusillina ( Haurakia ) cf. averni Figure 16C MATERIAL EXAMINED. Tierra del Fuego: 25 73-69 [1(d)]. 27C USNM E 219 [1(d)], 29B 71-305. 30C 71-344 [1(d)], 301 71-329 [5(d)], 33D 71-352 [1(d)], 33E 71-351 [3(d)], 35 USNM E 1596 [1(d)], (All material LACM unless otherwise indicated.) DIAGNOSIS. See Remarks. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD PS BS Sta. 71-344 Fig. 16C 2.13 1.28 1.66 0.88 2.41 2.8 1.3 0.47 8 21 Sta. 71-351 2.20 1.50 1.46 0.97 2.28 2.3 1.2 0.54 10 18 1.94 1.35 1.44 0.92 2.10 2.0 1.2 0.46 1.93 1.30 1.49 0.89 2.17 2.0 1.2 0.58 9 17 Sta 71-329 2.05 1.35 1.52 0.89 2.30 2.8 1.1 0.39 2.17 1.49 1.45 0.93 2.32 2.3 1.2 0.49 Sta. 71-305 2.12 1.26 1.68 0.90 2.34 3.0 — — 6 17 REMARKS. Specimens from the above localities differ from the types of P. averni in shell dimensions and/or strength of sculpturing (e.g., Fig. 16C) and may represent variations of one species or closely similar, separate species. We cannot make a deci- sion on the available material. Genus Manzonia Brusina, 1870 This genus is distinguished by its shell having a duplicated peristome and strong, smooth basal spi- rals as well as anatomical and other details (see Ponder, 1985a). Subgenus Alvinia Monterosato, 1884 The species included here is related to a small group of North and Central American species that were assigned to Manzonia ( Alvinia ) by Ponder (1985a). Manzonia ( Alvinia ) limensis n. sp. Figure 16D, E ETYMOLOGY. Named after the Province of Lima. MATERIAL EXAMINED. Types. Holotype, LACM 2676, 2 paratypes, LACM 2677. 4 72-78. Isla Chincha Norte, lea Prov., Peru. 13°38'S, 76°25'W, 6-12 m, ex- posed side, J.H. McLean, V. Alamo, 1 Apr. 1972. Additional Material Examined. Peru: 1 74-6 [many(d)]; AMS C. 167456 [10(d)]. 2A 38-208 [1(d)]. 4 35-159 [1(d)], (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Fig. 16D, E). Small (maxi- mum length 2 mm), ovate-conic, solid, opaque, with about 3 teleoconch whorls. Spire with lightly con- vex outlines, whorls moderately convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch with strong, sharp spiral ridges with wider interspaces and equally prominent axial ribs; 3 spiral ridges on penultimate and last whorl and an additional 3-5 on base; interspaces almost smooth with only very indistinct growth lines. Axial ribs reach suture, but do not extend onto base, and form low, peaked nodules at intersections with spi- ral ridges. Protoconch (Fig. 16D) of about 1.2 whorls, with 5-6 low spiral ridges bearing scattered granules, interspaces between spiral ridges smooth. Aperture oval, angled posteriorly. Inner lip mod- erately broad, attached to parietal wall in upper portion; outer lip orthocline with prominent ap- ertural varix immediately behind lip; peristome weakly duplicated. Umbilical chink represented by narrow groove. Periostracum not observed. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD PS BS PA BA Holotype 2.03 1.20 1.69 0.77 2.62 2.9 1.2 0.41 3 8 7 8 Paratype 1.99 1.18 1.68 0.77 2.57 2.9 1.2 0.40 3 7 7 8 Sta. AHF 534 1.97 1.16 1.69 0.74 2.66 3.0 1.2 0.38 3 6 7 8 Operculum, radula, and animal unknown. REMARKS. This is the only known species of Rissoinae in Peru with coarse axial sculpture. It resembles some of the other eastern Pacific species assigned to Manzonia ( Alvinia ) by Ponder (1985a) but differs in having a broader shell than any of the known species except M. (A.) almo (Bartsch, 191 1), which is much smaller. DISTRIBUTION. Peru (lea and Lima Provs.); known only from empty shells from 7 to 100 m. Uncommon. Genus Powellisetia Ponder, 1965 This genus was introduced for a group of New Zealand species with diverse shell morphology by Ponder (1965b) and has subsequently been redi- agnosed (Ponder, 1985a). The majority of species assigned to this genus are found in New Zealand, one from southern Australia (Ponder, 1985a), a few from the sub-Antarctic islands, and one from Ant- arctica (Ponder, 1983a). Members of the genus are Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 25 similar to some species included in Onoba but differ in usually having a more ovate shell, which is either smooth or finely spirally striate. The radula also differs in having more numerous (>10) cusps on the outer side of the lateral teeth. Powellisetia microlirata n. sp. Figures 12D, 17A-G, 18A ETYMOLOGY. Micro — Greek. Minute. Lira- ta — Latin. Ridge. Refers to the teleoconch sculp- ture. MATERIAL EXAMINED. Types. Holotype, LACM 2678, 175 paratypes, LACM 2679; 8 paratypes, AMS C. 167454. 20 73-72. Bahia Tom, Magallanes Prov., Chile. 50°11.3'S, 74°47.9'W, 14 m, P. Dayton (R/V Hero), 21 May 1973. Additional Material Examined. Southern Chile: 16 75- 46 [1(d)]. 17 73-75 [13]. 21 73-71 [2(d)], 23 73-70 [3], Tierra del Fuego: 25 73-69 [many(d)]. 30A 71-342 [1(d)]. 30F 71-334 [4(d)]. 301 71-329 [1(d)]. 30J 71-260 [9(d)]. 33A USNM H 656 [1(d)]. 33B 71-347 [10(d)]. 33D 71- 352 [20(d)]. 33F 71-357 [5(d)]. 33M 71-263 [1(d)]. 33R 71-328 [2], 36 USNM E 974 [1(d)]. 39 71-268 [2]. (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Fig. 17). Minute (maximum length 1.6 mm, often smaller), conical to elongate- conic, thin, translucent when fresh, with 2. 4-2. 9 teleoconch whorls. Spire with lightly convex to straight outlines, whorls moderately convex; pe- riphery of last whorl rounded. Sutures impressed, simple. Teleoconch with about 20-25 fine, rounded spiral cords on penultimate whorl and 30-40 on last whorl and base but cords too indistinct to be counted precisely in available material. Protoconch (Fig. 17D-F) of 1 .0—1.3 whorls, sculptured with rather scattered, spirally aligned, elongate granules and irregular low wrinkles. Aperture oval to almost round, weakly angled posteriorly. Inner lip narrow, attached to parietal wall in upper portion only; outer lip orthocline. Umbilical chink moderate. Periostracum very thin, transparent. Color white. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 1.00 0.67 1.49 0.43 2.32 2.6 1.3 0.25 Paratypes Fig. 17A 1.23 0.88 1.40 0.44 2.45 2.6 1.3 0.25 1.47 0.99 1.49 0.62 2.36 2.8 1.1 0.32 1.50 1.04 1.44 0.61 2.46 2.7 1.2 0.32 1.48 0.96 1.55 0.61 2.42 2.5 1.1 0.34 1.52 0.97 1.57 0.60 2.52 2.6 1.1 0.35 1.49 0.98 1.53 0.58 2.57 2.5 1.2 0.34 1.48 0.98 1.52 0.60 2.46 2.8 1.2 0.33 1.64 1.01 1.63 0.61 2.68 2.9 1.1 0.35 1.53 1.00 1.53 0.63 2.41 2.8 1.1 0.34 1.52 1.00 1.52 0.62 2.43 2.8 1.0 0.33 1.49 0.92 1.62 0.58 2.57 2.7 1.2 0.36 Sta. 71-328 Fig. 17C 1.30 0.80 1.64 0.51 2.57 2.5 1.2 0.35 Sta. 73-71 1.36 0.92 1.48 0.56 2.44 2.5 1.2 0.35 1.30 0.86 1.51 0.52 2.52 2.4 1.2 0.34 26 ■ Contributions in Science, Number 445 Operculum (Fig. 12D). Thin, horny, paucispiral. Oval but with strongly convex outer edge and weakly convex inner edge. Weakly angled posteriorly, rounded anteriorly. Radula (Fig. 18 A). Central teeth with cusp for- mula 7-8 + 1 +7-8, median cusp narrowly trian- gular; cutting edge triangular; single pair of well- developed basal denticles. Lateral teeth with cusp formula 7-8 + 1 + c. 15-19, primary cusp long, tri- angular. Inner marginal teeth with about 15 cusps on outer edge, inner edge not visible in mounts. Cusps on inner side of outer marginal obscured in mounts, outer side simple (3 radulae from 3 local- ities examined). Animal. Visceral coil darkly colored; head, foot, and mantle apparently unpigmented. REMARKS. This species has a shell similar to the New Zealand P. subtenuis (Powell, 1937) but differs in having a slightly different shell outline due to looser coiling and a less inflated last whorl (also resulting in a smaller aperture). It also lacks a varix on the outer lip. Powellisetia microlirata is smaller than most of the South American species of Onoba and has much finer teleoconch sculpture. Specimens from deeper water have more elongate shells with more convex whorls than the type, but specimens of intermediate shape have been observed. DISTRIBUTION. Valparaiso, Chile, to Tierra del Fuego; mainly intertidal. Empty shells from 30 to 900 m. Common. Genus Onoba FT and A. Adams, 1852 This genus, as recognized and diagnosed by Ponder (1985a), has a worldwide distribution, with major radiations in the northeastern Pacific and the south- ern oceans, including the Antarctic-sub-Antarctic. The genus is distinguished from other rissoids found in South America by the elongate-conic shell, which is smooth or has predominantly spiral sculpture. The radula has fewer than 10 cusps on the outer part of the lateral teeth. Onoba protofimbriata n. sp. Figures 19F, 20B, D ETYMOLOGY. Protos-— Greek. First. Fimbria- ta-— Latin. Fringed. Refers to the fimbriate proto- conch microsculpture. MATERIAL EXAMINED. Types. Holotype, LACM 2680, 11 paratypes, LACM 2681; 1 paratype, AMS C. 167423. 19 73-73. Bahia San Andres, N of Golfo de Penas, Aisen Prov., Chile. 46°35.3'S, 75°30.6'W, subtidal, P. Dayton (R/V Hero), 23 May 1973. Additional Material Examined. Southern Chile: 17 73- 75 [2(d)]. Tierra del Fuego: 27D USNM E 740 [1(d)]. 30G 71-259 [1(d)]. 32D 71-293 [1]. 330 71-310 [2(d)]. Falkland Islands: AMS C.167498, TW4 [1(d)]; AMS C.167497, TW5 [1(d)]. (All material LACM unless otherwise indi- cated.) DIAGNOSIS. Shell (Figs. 19F, 20A, B). Small (maximum length 2.2 mm), elongate-conic, mod- Ponder and Worsfold: Review of Rissoiform Gastropods rS-'N Figure 17. Details of shells of Powellisetia microlirata n. sp. A, Paratype, shell, length 1.23 mm. B, Holotype, shell, length 1.00 mm. C, Sta. 71-328, shell (deep water form), length 1.3 mm. D-G, Paratypes; D, lateral view of protoconch; E, apical view of protoconch; F, protoconch microsculpture; G, teleoconch microsculpture. Scale bars: A-C, 250 n m; D, E, 100 /xr n; F, G, 25 /urn. erately solid, possibly translucent when fresh, with 2. 7-3.1 teleoconch whorls. Spire with lightly con- vex outlines, whorls moderately convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch with moderately flat spiral cords, in- terspaces of approximately equal width; 8-9 (usu- ally 8) on penultimate whorl, 13-14 (usually 13) on last whorl and base. Microsculpture in interspaces of fine but distinct axial growth lamellae and fine spiral striae. Protoconch (Fig. 20A, B) of about 1.1- Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 27 Figure 18. Radulae of species of Rissoidae. A, Powellisetia microlirata n. sp,, paratype. B, Detail of central teeth. C, Onoba striola n. sp., paratype. D, Onoba georgiana (Pfeffer), Sta. 75-49. E, Onoba erugata n. sp., paratype. F, Detail of central teeth. Scale bars: A, C-E, 10 jam; B, F, 5 jam. 28 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods Figure 19. Shells of species of Onoba. A, G, Onoba erugata n. sp.; A, paratype, length 1.52 mm; G, holotype, length 1.49 mm. B-D, Onoba georgiana (Pfeffer); B, Sta. 75-49, length 1.79 mm; C, Sta. 75-49, length 1.86 mm; D, Sta. 75- 48, length 1.86 mm. E, Onoba subincisa n. sp., holotype, length 1.79 mm. F, Onoba protofimbriata n. sp., holotype, length 1.90 mm. H, Onoba cf. macra (Watson), DE 399, length 1.75 mm. Scale bars: 500 yum. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 29 Figure 20. Detail of shells of species of Onoba. A, C, Onoba ? lacuniformis n. sp., holotype; A, lateral view of protoconch; C, protoconch microsculpture. B, D, Onoba protofimbriata n. sp., holotype; B, lateral view of protoconch; D, protoconch microsculpture. E, F, Onoba sulcula n. sp., holotype; E, lateral view of protoconch; F, protoconch microsculpture. G, “ Onoba ” algida n. sp., holotype, protoconch microsculpture. Scale bars: A, B, E, 100 /urn; C, G, 50 /urn; D, F, 20 /urn. 1.2 whorls, sculptured with very fine, sharp, irreg- ular, curved axial lamellae interlocking to give a tessellated pattern showing basically spiral pattern. Aperture oval, angled posteriorly. Inner lip narrow, 30 ■ Contributions in Science, Number 445 detached from parietal wall; outer lip orthocline with moderate apertural varix immediately behind lip. Umbilical chink narrow groove. Periostracum not observed. Color white. Ponder and Worsfold: Review of Rissoiform Gastropods Dimensions. SL/ SL/ SL SW SW AL AL TWPW PD PS BS Holotype 1.90 0.99 1.91 0.72 2.66 2.8 1.2 0.35 8 13 Paratypes 1.98 0.99 1.99 0.74 2.68 2.8 1.2 0.37 9 14 1.85 1.03 1.80 0.78 2.37 2.7 1.1 0.35 8 13 2.01 1.04 1.94 0.77 2.61 2.9 1.1 0.38 8 13 2.18 1.05 2.09 0.77 2.83 3.1 1.1 0.37 8 13 2.11 0.91 2.12 0.76 2.35 3.1 1.1 0.39 8 13 Operculum, radula and animal unknown. REMARKS. This species is somewhat similar to O. subincisa n. sp. described below, but the shell has stronger sculpture and sharper ribs with wider interspaces. The spiral ribs on the shell are much finer in O. fimbriata than in O. fuegoensis (Strebel, 1908), although some forms of the latter species are otherwise somewhat similar in shell shape. The distinctive protoconch microsculpture, from which this species obtains its name, is not seen in any other South American species, although it it is similar, but more weakly developed, in O. georgiana (Fig. 21 B, D, F). Onoba protofimbriata resembles two New Zealand taxa in teleoconch characters: O. fallal (Powell, 1955) from the New Zealand sub- Antarctic is similar in size and in the number of spiral cords but differs in having linear interspaces between the cords (not wider than the cords); O. moriora (Powell, 1933) from the Chatham Islands is very similar to O. fallal but has the interspaces about equal to the cords. DISTRIBUTION. Southern Chile, Tierra del Fuego, and (possibly) Falkland Islands (see Part 2). Known mainly from dead shells from intertidal to 50 m. One similar shell from E. 740, 340-490 m, may be a different species. Uncommon. Onoba subincisa n. sp. Figures 19E, 22A-D, 23B, D, 241 ETYMOLOGY. Sub-Latin. Somewhat, less than. Incisa— Latin. Cut. Refers to the teleoconch microsculpture. MATERIAL EXAMINED. Types. Holotype, LACM 2682, 37 paratypes, LACM 2683; 4 paratypes, AMS C. 167424. 24 75-48. Punta Santa Ana, Fuerte Bulnes, Brunswick Peninsula, Strait of Magellan, Chile. 53°38'S, 70°54.5'W, intertidal, Sta. 38, 1. 1 i. McLean, 16 Nov. 1975. Additional Material Examined. Southern Chile: 24 75-48[2]; 75-49 [6(d)], T terra del Fuego: 28 71-270 [2(d)]. 31B 71-276 [1(d)]; 71-277 [1(d)]. 32A 71-273 [5(d)], 32G 71-311 [2(d)]. 33G 73-66 [10(d)]. 33] 71-264 [1(d)]. (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Figs. 19E, 22A-D). Minute (maximum length 2 mm), elongate-conic, thin, translucent, with 2. 2-3.1 teleoconch whorls. Spire with lightly convex outlines, whorls lightly convex; periphery of last whorl rounded. Sutures impressed, simple. Teleconch with broad, flat spiral cords sep- ; arated by narrow grooves; 7-9 on penultimate | whorl, 11-14 on last whorl and base; spiral cords may be very' faint, especially on upper whorls. Fine, well-spaced spiral threads over whole surface, es- pecially in interspaces. Protoconch (Fig. 22A-C) of 1 .0-1.2 whorls, sculptured with fine, close-set spiral wrinkles linked with irregular axial threads creating impression of irregular, spirally arranged shallow pits. Aperture oval, rather strongly angled posteri- orly. Inner lip narrow, separated from parietal wall in lower half; outer lip orthocline with weak ap- ertural varix immediately behind lip. Umbilical chink minute to absent. Periostracum very thin, trans- parent. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD PS BS Holotype 1.79 0.83 2.15 0.75 2.40 2.2 1.2 0.33 9 14 Paratypes 1.69 0.90 1.87 0.66 2.57 2.6 1.1 0.33 9 12 1.71 0.86 1.99 0.67 2.56 2.7 1.1 0.32 9 13 1.85 0.69 2.07 0.69 2.69 2.9 1.1 0.37 8 13 1.86 0.95 1.95 0.69 2.70 2.7 1.2 0.39 9 14 1.73 0.88 1.96 0.67 2.58 2.7 1.2 0.38 9 13 1.72 0.95 1.82 0.77 2.24 2.5 1.1 0.35 9 14 1.85 0.97 1.91 0.71 2.61 2.4 1.2 0.39 — — 1.73 0.96 1.81 0.64 2.40 2.7 1.1 0.35 9 14 1.62 0.87 1.S2 0.69 2.36 2.4 1.2 0.38 7 12 1.77 0.87 2.04 0.69 2.58 2.2 1.2 0.34 8 13 Sta. 73-66 1.78 0.88 2.02 0.65 2.72 2.9 1.1 0.37 8 12 1.68 0.88 1.91 0.66 2.53 2.7 1.1 0.35 8 11 1.73 0.86 2.02 0.67 2.57 2.7 1.1 0.35 9 13 Sta. 75-49 2.00 0.95 2.10 0.72 2.79 3.1 1.0 0.38 8 13 2.00 0.97 2.05 0.78 2.56 3.0 1.1 0.35 9 13 1.77 0.94 1.87 0.72 2.46 2.6 1.2 0.38 8 13 Operculum (Fig. 241). Yellow, thin, horny, pau- cispiral, oval with strongly convex outer edge and moderately convex inner edge. Weakly angled pos- teriorly, rounded anteriorly. Radula (Fig. 23B, D). Central teeth with cusp formula 4-5 + 1+4-5, median cusp narrow, sharp; cutting edge triangular; single pair of well-devel- oped basal denticles. Lateral teeth with cusp for- mula 4-5 + 1 +4-7, primary' cusp narrowly trian- gular, sharp. Inner marginal teeth with about 16- 20 shaip cusps on outer side, inner side obscured in mounts. Outer marginal teeth with 9-10 cusps on inner side, outer side simple (based on 3 radulae). Animal. Unpigmented. REMARKS. The characters that distinguish O. subincisa from O. protofimbriata, O. striola n. sp., and O. sulcula n. sp. are discussed in the remarks under those species. There are no other known South American elongate-conic Onoba species with finely spirally sculptured shells. The protoconch mi- crosculpture is similar to that of O. georgiana, but the shell of that species has only subobsolete spirals and is broader. Onoba macra (Watson, 1886) from Tristan da Cunha is similar to O. subincisa but differs in being larger with more and wider whorls. A few shells of a smaller undescribed species very similar to O. macra were available for SEM examination. This undescribed species, which was obtained from off Gough Island, near Tristan da Cunha, is figured here Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 31 Figure 21. Details of shells of Onoba erugata and O. georgiana. A, C, E, Onoba erugata n. sp., paratypes; A, lateral view of protoconch; C, apical view of protoconch; E, protoconch microsculpture. B, D, F, Onoba georgiana (Pfeffer), Sta. 75-48; B, lateral view of protoconch; D, apical view of protoconch; F, protoconch microsculpture. Scale bars: A- D, 100 yu.m; E, 20 yum; F, 50 yum. Figure 22. Details of shells of Onoba subincisa and O. cf. macra. A-D, Onoba subincisa n. sp., paratypes; A, lateral view of protoconch; B, protoconch microsculpture; C, apical view of protoconch; D, teleoconch microsculpture. E H, Onoba cf. macra Discovery Expedition Sta. 399; E, lateral view of protoconch; F, protoconch microsculpture; G, apical view of protoconch; H, teleoconch microsculpture. Scale bars: A, C, E, G, 100 yum; B, D, 50 yum; F, 20 yum; H, 200 yum. 32 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 33 Figure 23. Radulae of species of Onoba. A, C, Onoba scythei (Philippi); A, Sta. TW3; C, Sta. 75-49. B, D, Onoba subincisa n. sp., paratypes. E, F, Onoba fuegoensis (Strebel); E, Sta. 71-262; F, Sta. 71-344. Scale bars: A, 20 /am; B, E, 10 /am; C, 25 /am; D, F, 5 /am. for comparison with O. subincisa (Figs. 19H, 22E- H) but is not described because the small amount of available material is poor. Onoba subincisa has a shell somewhat similar to that of the New Zealand O. fumata (Suter, 1898), but the aperture extends much more markedly for- ward (i.e., more opisthocline outer lip) in the New Zealand species. In addition, there are more spiral 34 ■ Contributions in Science, Number 445 cords (9-11 on the penultimate whorl of O. fu- mata), and the cords are narrower and flatter with narrower interspaces in O. subincisa. The New Zealand sub-Antarctic species O. fallai is usually slightly larger, with stronger spiral cords, and has a distinct apertural varix. Another New Zealand species, Onoba inornata (Powell, 1933), is some- what similar in sculptural details but has a similar Ponder and Worsfold: Review of Rissoiform Gastropods Figure 24. Opercula of species of Rissoidae. All views inner side unless otherwise indicated. A-C, Onoba fuegoensis (Strebel); A, Sta. 71-342; B, Sta. 71-344; C, Sta. 71-262. D, H, Onoba scythei (Phillippi); D, Sta. 75-45; H, Sta. TW3, outer side. E, Onoba sfriola n. sp., paratype. F, Onoba erugata n. sp., paratype, G, Onoba georgiana (Pfeffer), Sta. 75- 49. I, Onoba subincisa n. sp., paratype. Scale bars: 200 um. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 35 aperture to that in O. fumata. The sub-Antarctic species O. steineni (Strebel, 1908) and O. turqueti (Lamy, 1905) have much larger and broader shells, and O. steineni has stronger sculpture. DISTRIBUTION. Southern Chile and Tierra del Fuego; intertidal; empty shells to 18 m. Moderately common. Onoba striola n. sp. Figures 18C, 24E, 25 ETYMOLOGY. Striola — Latin. A little furrow or line. Refers to the teleoconch microsculpture. MATERIAL EXAMINED. Types. Hoiotype, LACM 2684, 4 paratypes, LACM 2685; 1 paratype, AMS C. 167426. 32H 71-287. Puerto Cook, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.25'S, 64°2.3'W, inter- tidal, Sta. 71-2-37, USARP-SOSC-R/V Hero Cr.712, 17 May 1971. DIAGNOSIS. Shell (Fig. 25). Minute (maximum length 2 mm), elongate-conic, thin, translucent, with 2. 5-3.4 teleoconch whorls. Spire with straight to lightly convex outlines, whorls moderately convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch glossy with faint, slightly op- isthocline growth lines and weak, flat-topped spiral cords separated by much narrower grooves; about 8-9 cords on penultimate whorl, 18-20 on last whorl and base, typically very faint and sometimes almost obsolete. Protoconch (Fig. 25B-D) of 1.1- 1.3 whorls, with weak, irregular but mostly spirally aligned spiral wrinkles with interconnecting trans- verse wrinkles giving it finely reticulated appear- ance. Aperture oval, weakly angled posteriorly, with sharp peristome. Inner lip narrow, separated from parietal wall; outer lip slightly opisthocline with weak apertural varix immediately behind lip. Um- bilical chink very narrow to absent. Periostracum very thin, transparent. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TWPW PD PS BS Hoiotype 2.00 1.01 1.97 0.73 2.73 3.4 1.3 0.35 — 21 Paratypes 1.64 0.89 1.84 0.64 2.54 2.5 1.3 0.37 1.90 0.98 1.94 0.73 2.62 2.7 1.2 0.41 2.02 1.00 2.01 0.71 2.85 3.0 1.3 0.43 9 — 1.74 0.90 1.93 0.67 2.61 2.7 1.2 0.42 2.03 1.00 2.03 0.73 2.80 3.1 1.1 0.39 — — Operculum (Fig. 24E). Thin, horny, paucispiral. Elongate-oval with fairly strongly convex outer edge and moderately convex inner edge. Weakly angled posteriorly, rounded anteriorly. Radula (Fig. 18C). Central teeth with cusp for- mula 5-6 + 1 +5-6, median cusp long, narrow, pointed; cutting edge triangular; single pair of prominent basal denticles and weak pair of “pseu- dodenticles” arise from thickened lateral edges. Lateral teeth with cusp formula 7-8 + 1 + 7-8, pri- mary cusp narrowly triangular, outer cusps larger and less crowded than inner cusps, inner cusps rath- er irregular. Inner marginal teeth with about 16 36 ■ Contributions in Science, Number 445 small, rather irregular cusps on outer edge, inner edge obscured in mount. Outer marginal teeth sim- ple on outer side, inner side obscured in mount (single radula examined). Animal. Unpigmented. REMARKS. The only species in South America with a similar shell is O. subincisa, which has stron- ger and coarser teleoconch sculpture. The oper- culum is narrower in O. striola and the radula has a distinct second pair of basal cusps developed on the lateral edges of the central teeth, and the cutting edge of these teeth is more narrowly triangular than in O. subincisa. The protoconch microscupture is similar to that of O. subincisa and O. georgiana but is more weakly developed. DISTRIBUTION. Tierra del Fuego; intertidal. Uncommon. Onoba subaedonis n. sp. Figure 26B, C ETYMOLOGY. Idas a similar shell to O. ae- donis (Watson). MATERIAL EXAMINED. Types. Hoiotype, LACM 2686, 33 paratypes, LACM 2687; 5 paratypes, AMS C. 167427. 19 73-73. Bahia San Andres, N of Golfo de Penas, Aisen Prov., Chile. 46°35.3'S, 75°30.6'W, subtidal, P. Dayton (R/V Hero), 23 May 1973. Additional Material Examined. Southern Chile: 17 73- 75 [4(d)]. Tierra del Fuego: 27E BMNH DE 388 [6(d)]. Falkland Islands: AMS C.167499 TW5 [2(d)]. DIAGNOSIS. Shell (Fig. 26B, C). Minute (max- imum length 2 mm), ovate-conic, solid, opaque, with 2. 7-3.0 teleoconch whorls. Spire with straight outlines, whorls moderately convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch with moderately strong, rounded spiral cords with interspaces of equal width; 6-10 cords on penultimate whorl, 12-18 on last whorl and base. Interspaces have extremely fine axial growth lamellae and spiral striae. Very fine irregular axial i sculpture on upper spire whorls sometimes present. Protoconch (Fig. 26C) of 1.0-1. 3 whorls with close- ly spaced, fine, irregular spirally aligned wrinkles. Aperture broadly oval, weakly angled posteriorly. Inner lip narrow to moderately broad, attached to parietal wall in upper part only; outer lip orthocline, with very small posterior notch and moderate aper- tural varix immediately behind lip. Umbilical chink moderate. Periostracum not observed. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TWPW PD PS BS Hoiotype 1.80 1.14 1.60 0.71 2.56 3.0 1.3 0.42 Paratypes 1.92 1.18 1.63 0.73 2.63 3.0 1.1 0.41 7 15 2.02 1.19 1.69 0.76 2.65 3.0 1.0 0.42 6 12 1.90 1.16 1.64 0.75 2.53 3.0 1.1 0.41 7 13 1.83 1.12 1.63 0.73 2.50 2.8 1.2 0.42 10 18 1.85 1.12 1.65 0.74 2.49 2.8 1.0 0.41 7 13 1.90 1.12 1.70 0.68 2.81 2.9 1.2 0.41 6 — 1.76 1.11 1.59 0.69 2.56 2.7 1.2 0.43 7 14 1.85 1.16 1.59 0.75 2.46 2.8 1.2 0.41 Ponder and Worsfold: Review of Rissoiform Gastropods Figure 25. Details of shells of Onoba striola n. sp. A, Holotype shell, length 2.00 mm. B, Apical view of protoconch. C, F, Protoconch microsculpture; C, apex; F, last half whorl. D, Lateral view of protoconch. E, Teleoconch micro- sculpture. Scale bars: A, 500 jum; B, D, 100 /xm; C, E, F, 25 jum. 1.90 1.29 1.69 0.70 2.72 2.7 1.2 0.44 6 — 1.82 1.12 1.62 0.72 2.52 2.7 1.2 0.42 6 14 REMARKS. This species is very similar to Rissoa {Onoba) aedonis Watson, 1886, from Tristan da Cunha. Comparison of the South American mate- rial with the types of O. aedonis shows that the former material is slightly smaller and has more compact coiling, less convex whorls, and a slightly shorter spire. Unfortunately, no material of O. ae- Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 37 Figure 26. Detail of shells of Onoba subaedonis and O. glaphyra. A, D-F, Onoba glaphyra (Watson), Discovery Expedition Sta. 399; A, shell, length 1.75 mm; D, lateral view of protoconch; E, apical view of protoconch; F, protoconch microsculpture. B, C, Onoba subaedonis n. sp., holotype; Scale bars: A, B, 500 nm; C, F, 20 jam; D, E, 100 jam. donis was available for examination with the SEM. Rissoa ( Ceratia ) qlaphyra Watson, 1886, another species of Onoba from Tristan da Cunha, has a thinner shell that is more finely sculptured. A spec- imen from nearby Gough Island is figured for com- parison (Fig. 26A, D-F). B, shell, length 1.80 mm; C, protoconch microsculpture. Compared with other similar South American species, O. subaedonis has a smaller shell than O. scythei (Philippi) and more numerous spiral cords, always with the interspaces narrower than the ribs (unlike O. scythei). The shell of O. subaedonis has a shorter spire than any other South American Ono - 38 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ha. There is also some resemblance to Powellisetia microlirata n. sp., but that species has a smaller, more finely sculptured shell. Although somewhat similar to O. beta (Powell, 1955) from the New Zealand sub-Antartic, the shell of O. subaedonis is much smaller, broader, and with relatively much coarser spiral cords. It also resembles the New Zealand O. foveauxana (Surer, 1898), but the latter species is larger and often has heavier spirals. The New Zealand species of Onoba tend to have the outer edge of the shell aperture inclined forward (i.e., the lip is opisthocline) where- as it is more nearly orthocline in O. subaedonis. A few shells from a beach collection at the Falk- land Islands are essentially indistinguishable from the South American material and are tentatively recorded as this species. DISTRIBUTION. Southern Chile and Tierra del Fuego. Known only from empty shells from sub- tidal to 80 m. Uncommon. Beach specimens, prob- ably this species, from Falkland Islands. Onoba sulcula n. sp. Figures 20E, F, 27C ETYMOLOGY. Sulculus- — Latin. Little furrow. Refers to the teleoconch microsculpture. MATERIAL EXAMINED. Types. Holotype, LACM 2688, 1 paratype, LACM 2689; 301 71-329. 8 km S Punta Ventana, S side Isla de los Estados, Tierra del Fuego, Argentina. 54°54.5'S, 63°56'W, 771-903 m, Sta. 875, USARP-SOSC-R/V Hero Cr.715, 27 Oct. 1971. 10 Para- types, LACM; 1, AMS C. 167672. BMNH 27E DE 388. Off Cape Horn, Chile. 56°19.3'S, 67°09.45'W, 121 m. Discovery Expedition, 16 Apr. 1930. Additional Material Examined. Tierra del Fuego: 35 NMW, SNAE 346 [2(d)]?. DIAGNOSIS. Shell (Figs. 20E, F, 27C). Small (maximum length 2.5 mm), elongate-conic, mod- erately solid, opaque, with 2. 9-3.4 teleoconch whorls. Spire with lightly convex outlines, whorls moderately convex; periphery of last whorl round- ed. Sutures impressed, simple. Teleoconch with broad, flat spiral cords with slightly narrower inter- spaces, 5-7 on penultimate whorl and 10-11 on last whorl and base. Interspaces have crisp axial growth lamellae and very fine spiral striae. Proto- conch (Fig. 20E, F) of 1.2-1. 5 whorls, smooth ex- cept for numerous scattered, minute punctures (Fig. 20F). Aperture oval, angled posteriorly. Inner lip moderately broad, attached to parietal wall except for extreme lower portion; outer lip orthocline, with small posterior sinus and prominent apertural varix immediately behind lip. Each whorl usually bears weak but distinct varix. Umbilical chink ab- sent. Periostracum not observed. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TWPW PD PS BS Holotype 2.27 1.31 1.74 0.88 2.58 2.9 1.4 0.48 7 11 Paratype 2.43 1.26 1.92 0.87 2.80 2.9 1.2 0.52 7 10 Sta. DE 388 2.50 1.24 2.00 0.83 3.02 — — — 6 11 2.33 1.19 1.97 0.80 2.91 3.4 1.5 5 10 Operculum, radula, and animal unknown. REMARKS. This species has a larger and broader shell than the other South American species of Onoba with elongate-conic shells and has a very distinctive protoconch microsculpture. Of the Ant- arctic species, O. sulcula most resembles O. steineni in general shell features but is smaller and more conical in shape. It is also somewhat similar to O. protofmbriata but is larger and differs in proto- conch microsculpture. Compared with New Zea- land taxa, the shell of O. sulcula resembles that of O. sorensoni (Powell, 1955), but the latter species has more numerous, narrower ribs (up to 12 on the penultimate whorl) and flatter whorls. The shell of O. sulcula is about the same size as that of O. beta but has flatter spirals with narrower interspaces. The protoconch microsculpture suggests that O. sulcula might be a member of the genus Lironoba, although the teleoconch sculpture and shape are typical of Onoba. DISTRIBUTION. Tierra del Fuego; uncommon, 120-900 m. Onoba scythei (Philippi, 1868) Figures 23A, C, 24D, H, 28, 29 Rissoa scythei Philippi, 1868: 225 [Holotype (lo- cation unknown, presumed lost), ? Strait of Ma- gellan]; Cared les, 1950: 55; Carcelles and Wil- liamson, 1951: 272. MATERIAL EXAMINED. Southern Chile: 17 73-75 [4(d)]. 20 73-72 [many(d)]. 23 73-70 [8(d)]. 24 75-48 [3(+2d)]; 75-49 [many]; USNM 212271, St.2778 [27(d)], Tierra del Fuego: 25 73-69 [20]. BMNH 27E DE 388 [1]. 28 71-270 [1(d)]; 71-271 [1(d)]. 32H 71-287 [1(d)]. 33B 71-267[many(d)]; 71-347 [many(d)]. 34 USNM E 363 [1(d)]. 37B USNM E 967 [3]. Falkland 'islands: USNM 368419, [3 (d)]; SMNH SSPE 39 [6]; AMS C.167496, TW1 [many]; AMS C. 167495, TW2 [3]; AMS C. 167494, TW4 [2(d)]; AMS C. 167493, TW5 [5(d)], (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Figs. 28, 29). Small (maxi- mum length 2.7 mm), ovate-conic, solid, opaque, with 2. 7-3. 5 teleoconch whorls. Spire with lightly convex to straight outlines, whorls moderately con- vex; periphery of last whorl rounded. Sutures im- pressed, simple. Teleoconch with strong, spiral cords that may be flat with interspaces of equal width to cords or sharp with wider interspaces; number of cords very variable, 3-7 on penultimate whorl, 7- 14 on last whorl and base. Interspaces with rather coarse, distinct axial growth lamellae and very fine spiral striae. Irregular axial sculpture may be present on upper whorls. Protoconch (Figs. 28D, E, G, 29A, D, E, G) of 1 .0-1.5 whorls, with numerous scattered granules. Aperture oval, angled posteriorly. Inner Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 39 Figure 27. A, Onoba ? algida n. sp., holotype, shell, length 2.72 mm. B, E, Onoba ? lacuniformis n. sp.; B, holotype, shell, length 2.02 mm; E, paratype, radula. C, Onoba sulcula n. sp., holotype, shell, length 2.27 mm. D, Amphithalamus cf. inclusus Carpenter, Sta. 75-15, shell, length 1.24 mm. F, G, Barleeia meridionalis n. sp., holotype; F, shell, length 1.75 mm; G, protoconch microsculpture. Scale bars: A-C, 500 yum; D, F, 250 yu m; E, 10 yum; G, 20 jum. 40 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods Figure 28. Detail of shells of Onoba scythei (Philippi). A, Sta. 75-48, shell, length 2.26 mm. B, Sta. 75-49, shell, length 2.06 mm. C, Sta. 73-69, shell, length 2.26 mm. D, Sta. 75-48, lateral view of protoconch. E, Sta. 75-48, apical view of protoconch. F, Sta. 75-48, teleoconch microsculpture. G, Sta. 75-48, protoconch microsculpture. Scale bars: A-C, 500 jum; D, E, 100 /Lim; F, G, 25 ;um. lip narrow to moderately broad, attached to parietal wall in upper portion; outer lip orthocline, with weak posterior indentation and moderate apertural varix immediately behind lip. Umbilical chink very narrow to moderate. Periostracum well developed, yellowish-brown. Color white or pale yellow. Dimensions. SL/ SL/ SL SW SW AL AL TWPW PD PS BS Sta. 71-267 Fig. 29C 2.47 1.43 1.73 0.89 2.61 3.2 1.3 0.41 5 8 Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 41 Figure 29. Detail of shells of Onoba scythei (Philippi). A, B, D, F, Sta. TW1; A, apical view of protoconch; B, shell, length 2.01 mm; D, protoconch microsculpture; F, teleoconch microsculpture. C, E, G, Sta. 71-267; C, shell, length 2.47 mm; E, lateral view of protoconch; G, protoconch microsculpture. Scale bars: A, E, 100 yum; B, C, 500 /urn; D, F, 50 (um; G, 25 yum. 2.26 1.37 1.65 0.90 2.50 2.9 1.2 0.41 4 8 2.24 1.30 1.72 0.91 2.45 2.7 1.3 0.42 6 12 2.47 1.40 1.77 0.87 2.84 3.2 1.2 0.42 5 9 2.34 1.31 1.79 1.06 2.22 2.8 1.1 0.41 6 13 2.50 1.44 1.74 0.97 2.50 3.4 1.2 0.38 5 9 2.20 1.33 1.65 0.91 2.41 3.0 1.2 0.35 5 12 2.47 1.44 1.69 0.92 2.63 3.3 1.2 0.42 4 9 2.18 1.29 1.69 0.94 2.33 2.8 1.1 0.39 7 13 2.47 1.48 1.67 0.95 2.61 3.3 1.2 0.34 4 8 2.24 1.38 1.62 0.88 2.54 2.8 1.2 0.43 6 12 2.62 1.45 1.81 0.95 2.77 3.3 1.3 0.41 4 9 2.18 1.20 1.81 0.86 2.54 2.8 1.2 0.43 6 12 2.26 1.45 1.80 0.94 2.79 3.5 1.1 0.34 4 8 2.28 1.40 1.63 0.98 2.33 2.9 1.0 0.38 7 13 2.33 1.57 1.71 0.98 2.75 3.5 1.2 0.41 5 9 2.20 1.27 1.74 0.94 2.36 2.8 1.0 0.41 6 12 2.68 1.53 1.75 0.97 2.77 3.5 1.2 0.41 3 7 Sta. TW1 Sta. 75-48 Fig. 29B 2.01 1.28 1.57 0.87 2.32 2.9 1.3 0.37 5 12 Fig. 28A 2.26 1.41 1.60 0.99 2.29 2.8 1.4 0.41 5 13 - Sta. 73-69 Fig. 28C 2.26 1.31 1.72 0.90 2.50 3.0 1.5 0.41 5 14 Sta. 75-49 Operculum (Fig. 24D, H). Yellow, thin, horny, Fig. 28B 2.06 1.26 1.62 0.91 2.26 2.8 1.3 0.36 5 14 paucispiral, oval, with strongly convex outer edge 2.27 1.36 1.66 0.92 2.45 2.9 1.2 0.41 6 12 and weakly convex inner edge. Weakly angled pos- 2.24 1.31 1.70 0.86 2.60 2.9 1.1 0.42 7 12 teriorly, rounded anteriorly. 42 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods Radula (Fig. 23A, C). Central teeth with cusp formula 3-4 + 1 +3-4, median cusp blunt, short, markedly larger than adjacent cusps; cutting edge almost straight; single pair of basal denticles. Lat- eral teeth with cusp formula 5-7+ 1+4-5, primary cusp broadly triangular, pointed. Inner marginal teeth with 8-9 sharp cusps, all near distal end. Outer marginal teeth with about 9 sharp cusps on inner edge, outermost largest, about equal in size with those on inner marginal teeth; outer edge simple (single radula examined). Animal. Apparently unpigmented (from dry ma- terial). REMARKS. The type material of this species cannot be located, and we assume that it is lost. Philippi’s description is sufficiently detailed to iden- tify this species with reasonable confidence, even though it was not figured. This species, as we have interpreted it, is very variable in the shape and sculp- ture of the shell. There appear to be two main forms, although they sometimes intergrade. One is a shallow water form (Fig. 28A, B), which has a shorter, broader shell with more numerous and more flattened spiral cords than the other, an offshore form (Figs. 28C, 29C). Both forms, and the speci- mens from the Falkland Islands (Fig. 29A, B, D, F), all have the distinctive granules on the protoconch (Figs. 28D, E, G, 29A, D, E, G), which are also shared with O. fuegoensis. We have only been able to examine a radula from a shallow-water speci- men, and this is unusual in the central teeth having a simple, almost straight cutting edge rather than the triangular cutting edge seen in most species of Onoba. The shallow-water form is similar in shell char- acters to O. foveauxiana from New Zealand, but that species is larger and more elongate-conic in shape. The shell of another New Zealand species, O. alpha, is similar in size but is slightly narrower and has six to eight spiral cords on the penultimate whorl. In addition, there is a stronger posterior apertural sinus in the New Zealand species. The extreme deep water form is similar to O. delecta Ponder, 1983, from off South Georgia but has a larger shell with less distinct interstitial axial sculp- ture. DISTRIBUTION. Southern Chile, Tierra del Fuego, and Falkland Islands. Fine sculptured form mainly intertidal; common. Coarse sculptured form known only from empty shells found from 15 to 36 m; less common. Onoba fuegoensis (Strebel, 1908) Figures 23E, F, 24A-C, 30 Rissoia ( Cingula ?) fuegoensis Strebel, 1908: 56, pi. 6, fig. 90a, b (Tierra del Fuego, 36 m). Rissoa [Onoba) fuegoensis: Melvill and Standen, 1912: 350. Cingula fuegoensis: Carcelles, 1950: 55, pi. 2, fig. 22; Carcelles and Williamson, 1951: 271. Onoba fuegoensis: Ponder, 1983a: 20. MATERIAL EXAMINED. Types. Lectotype (here chosen) and damaged paralectotype, SMNH SSPE 3, Ti- erra del Fuego. 54°43'S, 64°08'W, 36 m, 6 Jan. 1902. Additional Material Examined. Tierra del Fuego: 27C USNM E 219 [3(d)], 27E BMNH DE 388 [27(d)], 29B 71- 305 [2(d)]. 30A 71-342 [2], 30B 71-346 [9(d)]. 30C 71-344 [many]; USNM H 659 [14(d)], 30D 71-340 [1(d)], 301 71- 329 [13], 33B 71-267 [15]; 71-347 [11(d)]. 33D 71-352 [3(d)]. 33E 71-351 [3]. 33F 71-265 [3(d)]; 71-357 [4], 33N 71-262 [4(d). 33N 71-262 [1]. 33Q 71-327 [2(d)]. 33R 71- 328 [3]. 37 A BMNH DE 88 [1(d)]. Burdwood Bank: 35 NMW, SNAE 346 [2(d)]. (All material LACM unless oth- erwise indicated.) DIAGNOSIS. Shell (Fig. 30). Small (maximum length 3.1 mm), elongate-conic, solid, opaque, with 3. 0-4. 3 teleoconch whorls. Spire with lightly con- vex outlines, whorls moderately convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch with strong, sharp spiral cords with wider interspaces; 4-6 (usually 5) cords on penul- timate whorl, 8-12 on last whorl and base; inter- spaces with rather coarse, distinct axial growth la- mellae and very fine spiral striae. Irregular axial sculpture may be present on upper whorls. Proto- conch (Fig. 30A, E, F) of 1.1-1. 3 whorls, with nu- merous scattered granules, sometimes aligned more or less spirally. Aperture oval, angled posteriorly. Inner lip narrow to moderately broad, attached to parietal wall; outer lip slightly opisthocline with a moderate posterior notch and prominent apertural varix immediately behind lip. Umbilical chink ab- sent or narrow groove. Periostracum thin, trans- parent. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD PS BS Lectotype 2.58 1.21 2.12 0.90 2.88 4.0 1.3 0.38 4 9 Sta. 71-342 Fig. 30C 2.68 1.19 2.26 0.79 3.40 4.2 1.2 0.36 4 9 Sta. 71-346 2.73 1.19 2.29 0.81 3.39 4.3 1.2 0.37 5 10 2.96 1.30 2.27 0.89 3.31 4.2 1.1 0.46 6 12 2.84 1.34 2.11 0.88 3.22 3.9 1.2 0.46 5 10 2.77 1.34 2.06 0.89 3.11 3.6 1.2 0.46 5 10 2.88 1.37 2.11 0.92 3.12 3.8 1.2 0.47 5 10 3.01 1.40 2.15 0.95 3.18 4.0 1.2 0.46 5 10 Sta. 71-262 3.05 1.42 2.15 0.94 3.26 4.1 1.2 0.44 5 10 2.42 1.27 1.91 0.87 2.78 3.2 1.3 0.47 4 8 2.99 1.34 2.22 0.91 3.27 4.2 1.2 0.41 5 10 2.83 1.32 2.14 0.91 3.09 3.8 1.2 0.44 5 11 Sta. 73-344 Fig. 30B 2.73 1.34 2.00 0.87 3.13 4.2 1.2 0.41 5 9 Fig. 30D 2.73 1.39 1.96 0.95 2.87 4.1 1.3 0.37 4 9 2.61 1.26 2.08 0.83 3.16 3.7 1.2 0.47 4 10 3.04 1.45 2.10 0.95 3.22 4.2 1.2 0.45 6 11 2.92 1.31 2.23 0.88 3.32 4.1 1.2 0.44 5 10 2.91 1.39 2.10 0.91 3.19 4.1 1.1 0.41 5 10 3.00 1.38 2.18 0.91 3.29 4.2 1.1 0.41 5 9 2.69 1.24 2.13 0.87 3.02 3.8 1.2 0.43 5 9 2.96 1.29 2.29 0.91 3.23 4.2 1.2 0.42 5 9 2.93 1.38 2.13 0.92 3.17 4.2 1.2 0.44 5 9 2.96 1.40 2.11 0.97 3.06 4.0 1.2 0.44 5 11 Sta. 71-329 Fig. 30G 1.97 1.00 1.97 0.69 2.86 3.0 1.2 0.43 5 10 2.29 1.13 2.03 0.75 3.04 3.9 1.2 0.32 5 8 Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 43 44 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods 2.37 1.18 2.01 0.78 3.03 3.6 1.2 0.40 5 10 1.99 1.00 1.99 0.69 2.89 3.1 1.2 0.42 5 8 Sta. 71-262 2.77 1.27 2.15 0.96 2.90 3.8 1.2 0.42 5 8 Operculum (Fig. 24A-C). Yellow, thin, horny, paucispiral, oval, with strongly convex outer edge and weakly convex inner edge. Weakly angled pos- teriorly, rounded anteriorly. Radula (Fig. 23E, F). Central teeth with cusp formula 4-5 + 1+4-5, median cusp small, sharp; cutting edge triangular; single pair of well-devel- oped basal denticles. Lateral teeth with cusp for- mula 6-8 + 1 + 6-8, primary cusp triangular, outer cusps larger and less crowded than small, irregular inner cusps. Inner marginal teeth with 16-17 sharp cusps on outer edge, inner edge obscured in mount. Outer marginal teeth with c. 8 cusps on inner side, outer side simple, cusps about equal in size to those on inner marginal teeth (4 radulae from 3 localities examined). Animal. Unpigmented. REMARKS. There is some variation in size and shape, particularly with respect to relative width. Extremes of this variation are shown in Figure 30B- D, G. The types are closest to the elongate form shown in Figure 30C. Some specimens from deeper water are much smaller and shorter (Fig. 30G) than the typical form, although intermediates are found (Fig. 30B, D). Some forms of O. scythei approach the shorter varieties of O. fuegoensis, but this latter species is always narrower. Both species have a sim- ilar granulate protoconch microsculpture, but the radula of O. fuegoensis has a triangular cutting edge, not almost straight as in O. scythei. This latter char- acter needs to be confirmed, however, as only a single radula of O. scythei has been examined. DISTRIBUTION. Tierra del Fuego in 50-900 m; shells found as shallow as 36 m. Common. Onoba georgiana (Pfeffer, 1886) Figures 18D, 19B-D, 21B, D, F, 24G Rissoa georgiana Pfeffer (in Martens and Pfeffer), 1886: 92, pi. 2, fig. 3 (SMNH 910; Swedish Ant- arctic Expedition, Sta. 28, South Georgia, 12-15 m); Carcelles and Williamson, 1951: 272. Onoba georgiana: Ponder, 1983a: 15, figs. 9f-h, lOa-h (gives full synonymy). MATERIAL EXAMINED. Southern Chile : 17 73-75 [7(d)]. 18 73-74 [7(d)]. 19 73-73 [many(d)]. 24 75-48 [22]; 75-49 [7(d)]. Tierra del Fuego : 28 71-271 [1(d)]. 29A 71- 302 [1(d)] (doubtful identification). 30H USNM H 664 [1(d)]. 39 71-268 [4], Falkland Islands: AMS C. 167651, TW1 [2(d)]. (All material LACM unless otherwise indi- cated.) DIAGNOSIS. See Ponder, 1983a. Shell (Figs. 19B-D, 21 B, D, F). Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Sta. 75-48 Fig. 19D 1.86 1.03 1.81 0.75 2.49 2.8 1.5 0.41 1.89 1.07 1.76 0.78 2.44 2.8 1.3 0.39 2.30 1.20 1.92 0.87 2.63 2.9 1.4 0.42 2.33 1.20 1.92 0.89 2.62 2.9 1.4 0.42 Sta. 75-49 Fig. 19B 1.79 1.15 1.56 0.77 2.32 2.5 1.4 0.41 Fig. 19C 1.86 1.12 1.66 0.78 2.37 2.7 1.3 0.41 1.69 1.09 1.55 0.75 2.24 2.7 1.2 0.39 1.83 1.12 1.63 0.76 2.39 2.8 1.1 0.38 Sta. 73-74 2.00 1.20 1.66 0.81 2.48 2.8 1.3 0.43 1.90 1.19 1.59 0.80 2.39 2.5 1.3 0.47 1.87 1.17 1.60 0.76 2.45 2.8 1.3 0.42 1.84 1.12 1.64 0.77 2.37 2.7 1.3 0.42 2.09 1.25 1.67 0.82 2.55 2.7 1.3 0.46 1.88 1.09 1.73 0.75 2.50 2.7 1.3 0.43 Sta. 73-73 1.96 1.10 1.78 0.76 2.56 2.9 1.3 0.39 1.96 1.12 1.75 0.76 2.56 2.9 1.2 0.42 1.97 1.13 1.74 0.77 2.54 2.8 1.2 0.42 1.98 1.17 1.63 0.80 2.41 2.5 1.3 0.42 1.90 1.14 1.67 0.78 2.42 2.8 1.2 0.40 2.00 1.10 1.81 0.78 2.53 2.8 1.3 0.43 1.93 1.10 1.76 0.73 2.65 2.8 1.3 0.42 1.82 1.07 1.69 0.69 2.64 2.6 1.3 0.43 2.00 1.13 1.79 0.77 2.58 2.8 1.3 0.43 1.91 1.09 1.76 0.75 2.54 2.7 1.4 0.43 Operculum (Fig. 24G). Radula (Fig. 18D). REMARKS. This species is not redescribed here because it is fully described by Ponder (1983a), the South American material being figured here for comparison. Of the other South American species, O. georgiana is most similar to O. erugata n. sp. in shell and radular characters and is contrasted with that species below. The records from southern Chile, Tierra del Fuego, and the Falkland Islands represent a significant range extension. DISTRIBUTION. South Georgia, South Orkney Islands, Falkland Islands, Tierra del Fuego, and southern Chile. Intertidal and shallow subtidal. Of- ten common. One specimen, doubtfully this spe- cies, from 270 m. Onoba erugata n. sp. Figures 18E, 19A, G, 21A, C, E, 24F ETYMOLOGY. E — Latin. Out of, from. Ru- Figure 30. Detail of shells of Onoba fuegoensis (Strebel). A, E, Sta. 71-344; A, lateral view of protoconch; E, protoconch microsculpture. B, D, Sta. 71-344, shells, both length 2.73 mm. C, Sta. 71-342, shell, length 2.68 mm. F, G, Sta. 71- 329; F, protoconch microsculpture; G, shell, length 1.97 mm. Scale bars: A, 100 jam; B-D, G, 500 yum; E, F, 50 /am. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 45 gata — Latin. Crease, wrinkle. Refers to protoconch microsculpture. MATERIAL EXAMINED. Types. Holotype, LACM 2690, 94 paratypes, LACM 2691; 6 paratypes, AMS C. 167425. 28 71-270. W side Bahia Buen Suceso, Tierra del Fuego, Argentina. 54°47.8'S, 65°16'W, intertidal, sand, rocks, and mussels, Sta. 71-2-8, USARP-SOSC-R/V Hero Cr.71 2, 23 Apr. 1971. Additional Material Examined. T ierra del Fuego: 30A USNM H 654 [1]. 32A 71-273 [3]. 32B 71-295 [1(d)]. 32H 71-287 [3]. 33G 73-66 [2(d)]. (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Figs. 19A, G, 21A, C, E). Small to minute (maximum length 1.6 mm), ovate- conic, thin, translucent when fresh, with 2. 5-2. 7 teleoconch whorls. Spire with straight to lightly convex outlines, whorls moderately to strongly convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth and some- times glossy but may have moderately strong or- thocline growth lines. Protoconch (Fig. 21A, C, E) of 1. 1-1.4 whorls, almost smooth but with about 14 faint, spiral rows of short, curved, axial wrinkles separated by narrow, spiral, smooth spaces (Fig. 21 E). Aperture oval, weakly angled posteriorly, with sharp peristome. Inner lip narrow, attached to pa- rietal wall in posterior part; outer lip orthocline, with moderate posterior notch and weak apertural varix immediately behind lip. Umbilical chink mi- nute to absent. Periostracum thin, transparent. Col- or white to pale yellow. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 1.49 0.85 1.77 0.63 2.35 2.6 1.2 0.37 Paratypes Fig. 19A 1.52 0.96 1.69 0.63 2.39 2.6 1.3 0.31 1.64 0.99 1.66 0.64 2.54 2.7 1.1 0.40 1.48 0.93 1.59 0.64 2.30 2.6 1.2 0.35 1.59 0.94 1.70 0.66 2.43 2.6 1.3 0.39 1.53 0.97 1.58 0.66 2.32 2.6 1.1 0.33 1.60 0.10 1.60 0.68 2.37 2.5 1.4 0.35 1.57 0.98 1.60 0.67 2.35 2.7 1.3 0.38 1.55 0.95 1.64 0.64 2.41 2.5 1.3 0.39 1.59 1.00 1.59 0.64 2.46 2.6 1.2 0.39 1.54 0.95 1.54 0.65 2.37 2.6 1.1 0.32 1.47 0.92 1.59 0.62 2.36 2.6 1.2 0.35 Operculum (Fig. 24F). Thin, horny, paucispiral, oval with strongly convex outer edge and moder- ately convex inner edge. Moderately angled pos- teriorly, rounded anteriorly. Radula (Fig. 18E). Central teeth with cusp for- mula 4+1+4, median cusp moderately long, nar- row, sharp; cutting edge triangular; single pair of well-developed basal denticles. Lateral teeth with cusp formula 4-5 + 1 + 6-8, primary cusp triangular, sharp, inner cusps smaller than outer cusps. Inner marginal teeth with about 20 small cusps on outer edge and distal end, inner edge obscured in mounts. Outer marginal teeth with simple outer edge, inner edge obscured in mounts (2 radulae examined). Animal. Unpigmented. REMARKS. This species is very similar to O. georgiana but has a smaller shell (compare dimen- sions with those of O. georgiana above), typically with a relatively smaller aperture, and more convex whorls. Also, unlike O. georgiana, there is no spiral sculpture or obvious periostracum on the teleo- conch and the protoconch microsculpture, al- though somewhat similar, is more weakly devel- oped and a different pattern (compare Figs. 21A, C, E and 21D, F). In addition, fresh (“live-collect- ed”) shells are translucent, whereas those of O. georgiana are opaque. Onoba melvilli (Hedley, 1916) from Macquarie Island SW of New Zealand is very similar to O. erugata in teleoconch shape and the lack of spiral sculpture but resembles O. georgiana in other characters, including proto- conch microsculpture (Ponder, 1983a). DISTRIBUTION. Tierra del Fuego. Mainly in- tertidal. Locally common. Onoba amissa nom. nov. Figure 10B Paludestrina striata Orbigny, 1840: 386, pi. 75, figs. 7-9 (Lectotype, BMNFI 1854.12.4.349 ( + 3 spec- imens of Powellisetia sp.); southern coast of Pa- tagonia, near Puerto San Julian (Argentina) and Falkland Islands); Da Silva and Davis, 1983: 144, fig. 17 (also as Onoba). Not Turbo striatus J. Adams, 1797 = Onoba semicostata (Montagu, 1803). Hydrobia striata: Carcelles, 1950: 55. Hydrobia ? striata : Carcelles and Williamson, 1951: 270. ETYMOLOGY. Amissa — Latin. Lost. Refers to the unknown locality of this taxon. DIAGNOSIS. A shell description and dimensions are given by Da Silva and Davis (1983). Dimensions. SL/ SL/ TW SL SW SW AL AL + PW Lectotype (from Da Silva and Davis, 1983) 2.18 1.4 1.56 1.04 2.10 4.25 REMARKS. The type of this species was appar- ently obtained on the southeastern coast of Pata- gonia or at the Falkland Islands and does not appear to have been recollected. The type is figured for comparison with the fauna described herein. It dif- fers from all of the other South American species of Onoba in its ovate shape and in having more numerous, closely spaced spiral cords. Da Silva and Davis (1983) figured this species and noted that there are two species in the syntype series. The largest specimen was chosen as the lec- totype (Fig. 10B) by Da Silva and Davis (1983), and they noted that the three smaller shells (Da Silva and Davis, 1983: figs. 18, 19) are a species of Pow- ellisetia. They stated (p. 144) that “[t]he largest specimen is Onoba striata, a rissoid species.” Un- 46 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods fortunately there is another Onoba striata in the literature, Turbo stnatus J. Adams, 1797, which is a synonym of Onoba semicostata (Montagu, 1803) because it is a primary homonym of the prior T urbo striatus da Costa, 1778. Onoba striata (Orbigny, 1840) is thus invalid (ICZN, Art. 57(c)(i) and a re- placement name is proposed above. Onoba (?) algida n. sp. Figures 20G, 27A ETYMOLOGY. Algida — Latin. Cold. Refers to sub-Antarctic location. MATERIAL EXAMINED. Types. Holotype, LACM 2692, 14 paratypes, LACM 2693; 3 paratypes, AMS C. 167457. 30C 71-344. Bahia Capitan Canepa, Isla de los Estados, Tierra del Fuego, Argentina. 54°50.2'S, 64°29.4'W, 67-71 m, Sta. 897, USARP-SOSC-R/V Hero Cr.715, 4 Nov. 1971. Additional Material Examined. Tierra del Fuego: 30B 71-346 [3(d)]. 30J 71-260 [2(d)], 33B 71-267 [many(d)]. 33F 71-265 [1(d)]; 71-357 [8(d)], (All material LACM.) DIAGNOSIS. Shell (Figs. 20G, 27A). Small (max- imum length 3.1 mm), ovate-conic, solid, opaque, with 2. 8-3. 5 teleoconch whorls. Spire with lightly convex to straight outlines, whorls moderately con- vex; periphery of last whorl rounded. Sutures im- pressed, simple. Teleoconch with strong, rounded spiral cords with wider interspaces and 5-9 prom- inent, broad axial folds that reach suture but not extending onto base. Five to 9 (usually 6 or 7) spiral cords on penultimate whorl, 11-13 on last whorl and base; interspaces with fine axial growth lamellae and very fine spiral striae. Spiral cords generally pass over tops of folds and do not form gemmules. Protoconch (Fig. 20G) of 1.2-1. 3 whorls, micro- sculpture of short to long ridges arranged as some- what irregular spiral lines. Aperture oval, angled posteriorly. Inner lip narrow to moderately broad, attached to parietal wall in upper portion; outer lip very slightly opisthocline with prominent apertural varix immediately behind lip. Umbilicus very small. Periostracum not observed. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD PS BS PA BA Holotype 2.72 1.56 1.74 0.99 2.75 3.2 1.2 0.50 6 11 5 6 Paratypes 3.01 1.71 1.76 1.06 2.83 3.3 1.2 0.49 9 12 6 8 2.87 1.70 1.69 1.04 2.75 3.3 1.2 0.48 6 12 6 9 2.96 1.66 1.79 1.04 2.84 3.2 1.2 0.54 — — — — 2.70 1.55 1.77 1.02 2.64 3.1 1.2 0.54 7 13 6 7 2.68 1.56 1.72 1.02 2.62 3.1 1.2 0.52 6 12 5 7 3.13 1.78 1.75 1.11 2.83 3.5 1.3 0.55 7 12 5 6 Sta. 75-357 2.71 1.37 1.71 0.85 2.77 3.3 1.2 0.50 7 12 5 7 2.75 1.48 1.86 0.95 2.91 3.4 1.2 0.48 7 11 5 6 2.39 1.40 1.71 0.97 2.47 2.8 1.2 0.57 5 12 6 7 Operculum, radula, and animal unknown. REMARKS. This is the only known species in the area with coarse axial sculpture on the shell, giving it a superficial resemblance to Rissoa ( Al - vania) lusciniae Watson, 1886, from Tristan da Cunha, but is broader, with stronger axials that persist onto the last whorl. Rissoa lusciniae was placed in Onoba ( Subestea ) by Ponder (1985a: 59). The types have about seven spiral ridges on the protoconch, this character being one of the factors influencing the subgeneric placement. Onoba al- gida has weak spiral sculpture on the protoconch made up of roughly aligned, short, rather irregular ridges. Partly because we have not been able to examine O. lusciniae using the SEM, it is by no means clear whether or not there is any likely re- lationship between these two species but our in- terpretation of the gross teleoconch and proto- conch characters favors the idea that the resemblance is superficial. Although we tentatively include the new species in Onoba, we have no firm basis for doing this. The protoconch microsculp- ture (Fig. 20G) is not distinctive, somewhat similar microsculpture being seen in species of Onoba and Alvania. The general shell morphology is also somewhat similar to species included in Alvania and Onoba and, although the strong axial sculpture on the teleoconch suggests a relationship with Al- vania, we judge this to be due to convergence. Other eastern Pacific species included in Onoba by Ponder (1985a) develop rather strong axial sculp- ture— for example O. carpenteri (Weinkauff, 1885) — although it is more prominently developed in O. algida than in other taxa included in Onoba s.s. to date. Final determination of the generic po- sition of this species must await anatomical ex- amination. DISTRIBUTION. Tierra del Fuego; known only from empty shells, moderately common in 35-70 m. Onoba (?) lacuniformis n. sp. Figures 12F, 20A, C, 27B, E ETYMOLOGY. Similar in shape to the gastro- pod genus Lacuna. MATERIAL EXAMINED. Types. Holotype, LACM 2694, 2 paratypes, LACM 2695; 1 paratype, AMS C. 167458. 32H 71-287. Puerto Cook, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.25'S, 64°2.3'W, inter- tidal, Sta. 71-2-37, USARP-SOSC-R/V Hero Cr.712, 17 May 1971. Additional Material Examined. Southern Chile: 24 75- 48 [1(d)]; 75-49 [3(d)]. Tierra del Fuego: 32G 71-311 [1(d)]. Falkland Islands: AMS C.l 67486, TW1 [2(d)], (All ma- terial LACM unless otherwise indicated.) DIAGNOSIS. Shell (Figs. 20A, C, 27B). Small (maximum length 2.2 mm), ovoid, moderately thick, opaque, with 1. 5-2.2 teleoconch whorls. Spire with moderately convex outlines, whorls strongly con- vex and slightly shouldered; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth. Protoconch (Fig. 20 A, C) of 1. 1-1.3 whorls Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 47 with fine, irregular reticulation so that surface ap- pears to be covered in small, irregular, shallow pits. Aperture oval, weakly angled posteriorly, with slightly prosocline, simple outer lip. Umbilical chink extremely narrow or absent. Periostracum not ob- served. Color pale yellowish grey to white. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD Holotype 2.02 1.27 1.59 0.93 2,17 2.2 1.3 0.49 Paratype 2.05 1.21 1.69 0.90 2.25 1.8 1.2 0.60 Sta. 75-49 2.13 1.34 1.57 0.97 2.18 1.6 1.2 0.64 2.15 1.38 1.56 1.03 2.08 1.5 1.1 0.62 Sta. 75-48 2.02 1.38 1.47 1.02 1.98 1.9 1.1 0.53 Sta. 71-311 2.13 1.31 1.62 1.00 2.13 2.0 1.1 0.53 Note: Limits of protoconch difficult to determine. Operculum (Fig. 12F). Simple, paucispiral with weakly angled posterior end and almost equally convex inner and outer edges. Radula (Fig. 27E). Central teeth with cusp for- mula 4-5 + 1 +4-5, median cusp minute, smaller than adjacent cusps; lateral cusps sharp, small, and somewhat irregular; cutting edge convex; single pair of short basal denticles; outer edges of tooth weak- ly thickened, straight. Lateral teeth with cusp for- mula 4 + 1+5, primary cusp long, triangular, sharp, inner cusps smaller than outer. Inner marginal teeth with 15-16 small, sharp cusps on inner side and distal end, outer edge obscured in mount. Outer marginal teeth with simple outer edge, inner side obscured in the single specimen available. Animal. Unknown. REMARKS. This species is only tentatively as- signed to Onoba, and the Rissoidae, because, on the basis of shell and radular characters, it may represent a new genus. The shell has a somewhat larger, differently shaped aperture compared with species of Onoba, although the protoconch mi- crosculpture is similar to that seen in O. georgiana. The radula (based on the single specimen available) differs considerably in not having the lateral edges of the central teeth strongly thickened and arched and the pair of basal cusps is smaller than in species of Onoba. The convex cutting edge of the central teeth bears minute cusps and the median cusp is smaller, not markedly larger as in species of Onoba and in all other rissoids. The shell has some simi- larity to members of the Elachisinidae (Ponder, 1985b), but the radula of that group differs consid- erably from O. lacuniformis. A new genus-group name is not provided pending anatomical infor- mation to establish the relationships of this species to other taxa. DISTRIBUTION. Strait of Magellan, Tierra del Fuego, and Falkland Islands; intertidal to shallow subtidal. Uncommon. Family BARLEEIDAE The genera of this family have been reviewed and diagnosed by Ponder (1983b). The family is now 48 ■ Contributions in Science, Number 445 (Ponder, 1988) restricted to what was previously the subfamily Barleeinae (Ponder, 1983b). Barleeid shells closely resemble those of eatoniellids in shape and size and in possessing an inner “chitinous” lay- er, but barleeids have a straight (not curved) oper- cular peg and the operculum has a heavy internal ridge. In addition, barleeids are very different an- atomically and are phallate. Genus Barleeia Clark, 1853 This genus has been diagnosed by Ponder (1983b), who also listed many of the named species. A char- acteristic feature is the dark wine-red operculum and the protoconch having pitted microsculpture. This genus has a wide distribution but is unknown from New Zealand. Barleeia meridionalis n. sp. Figure 27F, G ETYMOLOGY. Meridionalis— Latin. Southern. MATERIAL EXAMINED. Types. Holotype, LACM 2696, 4 paratypes, LACM 2697. 2B 74-24. Isla San Loren- zo, midway on NE side, rocky point N of naval base, Lima Prov., Peru. 12°5.75'S, 77°12.9'W, 0-4.5 m, J.H. McLean, J.A. Coyer, and J.M. Engle, 29 Jan. 1974. 9 paratypes, LACM 2754; 3, AMS C.167453. 3 72-76. Pu- cusana (Chilca), Lima Prov., Peru. 12°30'S, 76°49'W, 0-6 m, J.H. McLean, Mar.-Apr. 1972. Additional Material Examined. Peru: 2A 38-208 [1(d)]. 2C 35-147 [1(d)]. 4 72-78 [many(d)]; 35-159 [8(d)]. 5 35- 156 [1(d)]. (All material LACM unless otherwise indicat- ed.) DIAGNOSIS. Shell (Fig. 27F, G). Small (maxi- mum length 1.75 mm), ovoid-conic, moderately thick, opaque, with 2. 5-2. 8 teleoconch whorls. Spire with lightly convex outlines, whorls lightly convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth except for very faint axial growth lines and almost imperceptable traces of spirals on base. Protoconch with 12-14 spiral rows of punctures, of 1.2-1. 3 whorls. Aperture oval, weakly angled posteriorly, with sharp peristome. Inner lip narrow, outer lip moderately prosocline. Umbilical chink absent. Periostracum not observed. Color uniform reddish-brown, some shells bleached to white. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 1.75 1.05 1.66 0.68 2.59 2.8 1.2 0.39 Paratypes 1.64 1.01 1.63 0.57 2.89 2.7 1.2 0.35 1.59 1.11 1.63 0.66 2.75 2.7 1.3 0.42 Sta. 72-78 1.68 0.98 1.71 0.63 2.64 2.5 1.2 0.40 1.66 0.99 1.67 0.63 2.61 2.6 1.2 0.34 1.67 1.05 1.58 0.66 2.54 2.6 1.2 0.39 1.52 1.02 1.59 0.59 2.75 2.5 1.2 0.41 1.64 1.02 1.61 0.63 2.59 2.5 1.2 0.39 1.71 1.00 1.71 0.63 2.69 2.7 1.2 0.35 1.67 1.00 1.67 0.66 2.54 2.7 1.2 0.33 Operculum, radula, and animal unknown. Ponder and Worsfold: Review of Rissoiform Gastropods REMARKS. Because the shell (including the di- agnostic protoconch microsculpture) is typical of the genus, we have no hesitation in placing this species in Barleeia despite the lack of confirmatory evidence from the operculum and radula. Com- pared with other eastern Pacific species, the shell of this species is similar in size and shape to three taxa: B. alderi (Carpenter, 1856) differs in its yellow color with narrow color bands; B. californica Bartsch, 1920, differs in its subangled periphery and is white with wide pale brown bands; and B. car- penteri Bartsch, 1920, is white. Barleeia rubrooperculata (Castellanos and Fer- nandez, 1972) from the Golfo de San Manas, Ar- gentina, has a much more elongate shell. This spe- cies was described as an Eatoniella, but the radula and operculum (Castellanos and Fernandez, 1972b: 231, 233, figs 5, 6) indicate its placement in Barleeia (Ponder, 1983b). DISTRIBUTION. Peru; empty shells from in- tertidal to 90 m. Uncommon. Family Anabathridae This family was separated from the Barleeidae by Ponder (1988) on the basis of anatomical differ- ences but, like that family, the shell has an inner “chitinous” layer. The genera it contains were re- viewed and diagnosed by Ponder (1983b). Genus Amphithalamus Carpenter, 1864 This genus has been diagnosed by Ponder (1983b). It has a wide distribution in temperate to tropical waters, although no Recent species are known from the eastern Atlantic/Mediterranean Sea. Species of Amphithalamus are distinguished by their small, thick shells that have a deep groove separating the inner lip from the parietal wall. Amphithalamus cf. inclusus Carpenter, 1864 Figure 27D MATERIAL EXAMINED. Northern Chile: 8A 75-17 [5(d)], 8CB 75-15 [2(d)]. 8B 75-19 [1(d)]. (All material LACM.) DIAGNOSIS. See remarks. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD Sta. 75-15 Fig. 27D 1.24 0.83 1.49 0.54 2.32 1.8 1.4 0.34 1.35 0.89 1.52 0.62 2.18 2.2 1.4 0.38 REMARKS. The only available material of this species is in rather poor condition. The shells are very similar to the Panamic A. inclusus (see Ponder, 1983b, for synonymy) and also to its Caribbean analogue A. vallei Aguayo and Jaume, 1947. DISTRIBUTION. Northern Chile. Intertidal; only empty shells. Subclass HETEROBRANCHIA Superfamily RISSOELLOIDEA Family RISSOELLIDAE The genus-group taxa in this family were reviewed by Ponder and Yoo (1977b), who recognized only one genus and four subgenera. Comments by Hasz- prunar (1988) suggest that some of these are very distinct anatomically and will ultimately be raised to generic rank. Members of the family are readily distinguished by their simple, thin, translucent shells and distinctive operculum bearing a short, blunt peg in the middle of the columellar edge. Genus Rissoella J.E. Gray, 1847 Subgenus Rissoella Members of this subgenus have a distinctive radula with large central and lateral teeth that bear many small cusps. Rissoella ( Rissoella ) peruviana n. sp. Figure 31 ETYMOLOGY. Named after Peru. MATERIAL EXAMINED. Types. Holotype, LACM 2698, 21 paratypes, LACM 2699; 4 paratypes, AMS C. 167452. 1 74-6. Isla Lobos de Afuera (NW and NE of isthmus), Peru. 6°57.1'S, 80°42.3'W, 2-10 m, rocks and sand, J.H. McLean, J.A. Coyer, and J.M. Engle, 19-20 Jan. 1974. Additional Material Examined. Peru: 2B 74-24 [1(d)]. Northern Chile: 6A 64-16 [13]; AMS C. 167475 [2]. 8A 75-17 [10(d)], 8B 75-19 [5(d)], (All material LACM unless otherwise indicated.) DIAGNOSIS. Shell (Fig. 31A, C, D). Minute (maximum length 1.7 mm), ovate-conic, thin, semi- transparent, with about 3 teleoconch whorls. Spire with straight to lightly convex outlines, whorls moderately to strongly convex; periphery of last whorl rounded. Sutures impressed, simple. Teleo- conch smooth with faint growth lines. Protoconch (Fig. 31D) smooth of about 1.1-1. 2 whorls. Ap- erture oval, angled posteriorly, with sharp, simple peristome. Inner lip thin, outer lip slightly proso- cline. Umbilical chink moderate. Colorless when fresh, dead shells white. (Description only based on type series.) Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Holotype 1.54 0.97 1.60 0.68 2.29 3.1 1.2 0.21 Paratypes 1.60 0.97 1.66 0.70 2.27 3.1 1.1 0.22 1.64 0.99 1.65 0.73 2.24 3.1 1.2 0.27 1.74 1.06 1.64 0.76 2.29 3.3 1.1 0.24 1.38 0.86 1.61 0.61 2.27 2.8 1.2 0.25 1.49 0.92 1.61 0.69 2.16 2.9 1.2 0.25 Sta. 64-16 Fig. 31A 1.17 0.76 1.53 0.54 2.18 2.0 1.2 0.33 0.95 0.65 1.47 0.48 1.96 2.0 1.2 0.25 0.98 0.70 1.40 0.48 2.02 1.9 1.3 0.27 Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 49 Figure 31. Rissoella spp. A, Rissoella cf. peruviana, Sta. 64-16, shell, length 1.17 mm. B-G, Rissoella peruviana n. sp.; B, D-F, paratypes; B, operculum, inner side; D, protoconch; E-G, radula; E, detail of marginal teeth; F, detail of central teeth. C, Holotype, shell, length 1.54 mm. Scale bars: A, C, 250 jum; B, 100 /urn; D, 50 /*m; E, 10 jam; F, 5 fim; G, 20 nm. 50 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods 1.13 0.73 1.54 0.53 2.14 2.1 1.0 0.25 1.00 0.69 1.45 0.49 2.02 1.7 1.2 0.32 1.01 0.67 1.52 0.48 2.09 1.7 1.2 0.32 1.12 0.74 1.51 0.55 2.04 2.0 1.2 0.32 1.09 0.72 1.51 0.45 2.40 2.0 1.1 0.30 1.14 0.77 1.47 0.54 2.12 2.0 1.1 0.31 1.06 0.68 1.57 0.49 2.15 1.8 1.2 0.30 Operculum (Fig. 31B). Typical of genus, inner surface with small twisted peg, median transverse ridge and two ridges along inner edge. Nucleus marginal, at middle of inner edge. Radula (Fig. 31E-G). Symmetrical; central teeth large, subrectangular, with about 7-8 sharp, main cusps alternating with small, sharp cusps about Vi- V3 size of main cusps; no clearly differentiated me- dian cusp. Lateral teeth elongate, with cusp for- mula, 9-12 + 1 + 6-8, primary cusp triangular, sharp, markedly larger than adjacent cusps, with 1-2 cusps on either side larger than other lateral cusps, all cusps sharp. Marginal teeth smaller than lateral teeth (outer marginal teeth absent), 7-9 + 1 + 7-8, cusps sharp, primary cusp triangular, markedly larger than adjacent cusps (based on 2 radulae). Animal. Dried, not obviously pigmented. REMARKS. The only other rissoellid species rec- ognized from the eastern Atlantic are Rissoeila tu- mens (Carpenter, 1856), R. asteriaphila (Carpenter, 1864), and R. hertleini A.G. Smith and Gordon, 1948. The two former taxa are more ovate, the first having a ridge on the base. The last species is more similar in shape but is larger (length 2.2 mm com- pared to less than 1.8 mm) and is yellowish-brown rather than colorless. None of the other eastern Pacific species have had their radula examined to enable more detailed comparison of these other- wise rather featureless taxa. Empty shells, which agree rather well with the type series, have been found well south of the type locality of Rissoeila peruviana (see additional ma- terial examined). One lot (LACM 64-16) contained a specimen with a dried animal, but the radula was not successfully mounted, although the operculum is typical of the family and like that illustrated for R. peruviana. Until radulae are examined, the spe- cific status of these specimens must remain in some doubt. They are usually smaller in size than the type series of R. peruviana and, consequently, have fewer whorls (see the dimensions section) but they agree well in most other shell characters. DISTRIBUTION. Peru to northern Chile; inter- tidal to 15 m. Uncommon. Part 2 Additional species and records for South Georgia and Falkland Islands Family Eatoniellidae Eatoniella ( Eatoniella ) strebeli n. sp. Eatoniella kerguelenensis forma contusa Strebel, 1908: 57 (in part). Eatoniella aff. caliginosa Ponder, 1983a: 6, figs. 4c- e, 5h, i. ETYMOLOGY. Named for the German mala- cologist H. Strebel (1834-1915) in recognition of his early work on sub-Antarctic faunas. MATERIAL EXAMINED. Types. Holotype (speci- men figured by Ponder, 1983a, fig. 4c) and 50+ paratypes, SMNH 5 paratypes, AMS C. 302328. SSPE 25, South Georgia. 54°22'S, 36°27'W, 24-25 m. Additional Material Examined. South Georgia: SMNH SSPE 28 [2]. Burdwood Bank: SMNH SSPE 59 [many], DIAGNOSIS. Shell. Minute (maximum length 1.7 mm), ovate-conic, moderately thick, with 3.1- 3.5 teleoconch whorls. Spire with lightly convex outlines, whorls moderately convex; periphery of last whorl rounded. Sutures impressed, simple. Tel- eoconch smooth with faint growth lines. Proto- conch smooth, of about 1.2-1. 4 whorls. Aperture oval to near circular, very weakly angled posteri- orly, slightly reflexed, with moderately sharp peri- stome, lacking external varix. Inner lip moderately broad, detached from parietal wall, outer lip mod- erately prosocline. Umbilical chink absent or very narrow. Periostracum very thin, transparent. Color white to pale cream. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD Holotype 1.54 1.12 1.38 0.74 2.07 3.1 1.3 0.26 Paratypes 1.72 1.13 1.52 0.74 2.31 3.5 1.4 0.30 1.34 0.92 1.45 0.59 2.26 3.2 — — 1.70 1.10 1.54 0.72 2.37 3.3 1.2 0.29 1.56 1.10 1.41 0.72 2.17 3.2 1.3 0.27 1.64 1.02 1.61 0.69 2.38 3.2 1.2 0.24 Operculum. Pale yellow, oval. Peg stout, curved. Radula. Central teeth with cusp formula 3 + 1+3, median cusp with rounded end, narrow. Lateral teeth with cusp formula 2 + 1+3, primary cusp tri- angular. Inner marginals with cusp formula 4 + 1 + 1- 2, primary cusp triangular. Outer marginals with 6- 7 small, sharp cusps (based on 2 radulae). Animal. Unpigmented or pale yellowish, with distinct orange-brown spot on visceral coil. REMARKS. This species was recognized as a probable undescribed taxon by Ponder (1983a), who figured the shell, radula, and operculum. Exami- nation of the additional material now available shows that the shell form, radula, and operculum are consistent, and we are now confident in rec- ognizing these specimens as a new species. Of the southern species, E. strebeli is similar to E. caligi- nosa from which it differs in its yellowish-white color (not grey or black) and smaller size. It also appears to be similar to E. argentinensis, being of similar size and shape, but direct comparison of specimens has not been possible. However, the rad- ula of E. argentinensis has two cusps on either side of the median cusp of the central teeth, the oper- cular peg is straighter, and the shell aperture is not as flared. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 51 I Figure 32. Species of Eatoniella from Falkland Islands and Juan Fernandez Islands. A-C, Eatoniella bennetti (Preston), Sta. TW2; A, shell, length 2.01 mm; B, operculum, inner side; C, radula. D, Eatoniella zigzag n. sp., holotype, shell, length 2.14 mm. Scale bars: A, D, 500 /urn; B, 200 /um; C, 20 /urn. DISTRIBUTION. South Georgia, Burdwood Bank, and (possibly) South Orkney Islands. Eatoniella (Eatoniella) cf. cana Ponder, 1983 See Part 1 for details. Eatoniella ( Eatoniella ) occulta Ponder, 1983 Eatoniella occulta Ponder, 1983a: 12, figs. 2f, 8a-c. MATERIAL EXAMINED. Falkland Islands: SMNH SSPE 43 [7]; SMNH SSPE 46 [many]; AMS C.167492, TW2 [3]; AMS C.167491, TW3 [2]. Burdwood Bank: SMNH SSPE 59 [many]. REMARKS. This species was described from a single sample, but the above records indicate that it is rather common at the Falkland Islands and its range is now extended to the Burdwood Bank. Eatoniella ( Eatoniella ) bennetti (Preston, 1912) Figures 10D, 32A-C Laevilitorina bennetti Preston, 1912: 636, pi. 21, fig. 1. Ponder and Worsfold: Review of Rissoiform Gastropods 52 ■ Contributions in Science, Number 445 Figure 33. Species of Rissoidae from Juan Fernandez Islands. A, C, Onoba ? isolata n. sp., holotype; A, lateral view of protoconch; C, shell, length 1.91 mm. B, D, Onoba ? protopustulata n. sp., holotype; B, shell, length 1.77 mm; D, lateral view of protoconch. Scale bars: A, D, 100 /urn; B, C, 500 jam. MATERIAL EXAMINED. Type. Holotype, BMNH, 1913.7.31.186. Port Stanley, Falkland Islands, associated with Tonicia, A.G. Bennett. Additional Material Examined. Falkland Islands: AMS, C. 167484, C. 167485 [many], BMNH [6], LACM [6], all TW2. DIAGNOSIS. Shell (Figs. 10D, 32A). Small (max- imum length 2.1 mm), elongate-conic, rather thick, opaque, with about 4 teleoconch whorls. Spire with straight outlines, whorls very lightly convex; pe- riphery of last whorl rounded or very slightly an- gled. Sutures impressed, simple. Teleoconch smooth, sometimes glossy, with regular, faint growth lines. Protoconch smooth of 1.3-1. 5 whorls. Aperture ovoid to almost circular, weakly angled posteriorly, with sharp peristome, lacking external varix. Inner lip narrow, outer lip moderately to strongly proso- cline. Umbilicus absent. Periostracum very thin, transparent. Color uniform dark grey to black, oc- casionally pale grey. Dimensions. SL SW SL/ SW AL SL/ AL TW PW PD Lectotype 1.67 1.05 1.59 0.58 2.88 3.5 1.4 0.30 Sta. TW2 Fig. 32A 2.01 1.12 1.80 0.69 2.92 3.8 1.4 0.29 2.14 1.13 1.89 0.76 2.82 4.1 1.4 0.29 2.03 1.19 1.71 0.68 3.00 4.0 1.5 0.28 1.96 1.12 1.75 0.69 2.84 4.1 1.4 0.28 1.99 1.20 1.66 0.69 2.88 3.9 1.3 0.30 2.10 1.16 1.81 0.72 2.92 4.1 1.5 0.29 2.04 1.13 1.80 0.70 2.90 4.0 1.4 0.27 Operculum (Fig. 32B). Externally black with yel- lowish mottling; oval. Peg rather narrow, curved. Radula (Fig. 32C). Central teeth with cusp for- mula 3 + 1+3, median cusp short, spade-like, with truncated to rounded end; cusps very small. Lateral teeth with cusp formula 2 + 1 + 2, primary cusp tri- angular. Inner marginal teeth with cusp 3-4 + 1 +2- Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 53 3, primary cusp triangular, outermost cusps very small. Outer marginal teeth with 8-10 small, sharp cusps (based on 2 radulae). Animal. Unknown. REMARKS. The lectotype is slightly smaller than our virtually topotypic series but agrees in all other characters. Preston’s measurements (2.25 x 1.25) are slightly larger than any of our specimens. Be- cause of this discrepency between the specimen labeled “holotype” in the BMNH and the pub- lished dimensions, and in the absence of any evi- dence to suggest that there was only one specimen before Preston when he described the species, we have designated the BMNH specimen as the lec- totype. Specimens of this species with pale-colored shells are somewhat similar to E. cana but are smaller, with fewer, slightly more convex whorls. Similarly colored sub-Antarctic species include E. kerguele- nensis, which is much larger, and E. caliginosa, which is shorter and broader. Dark-colored South American species are thinner-shelled with the ex- ception of E. nigra, which is smaller and relatively shorter. DISTRIBUTION. Falkland Islands; intertidal. Locally common. Family Rissoidae Onoba georgiana (Pfeffer, 1886) See Part 1 for details. Onoba subaedonis n. sp. Described above; see Part 1 for details. Onoba scythei (Philippi, 1868) Described above; see Part 1 for details. Onoba cf. protofimbriata n. sp. Described above; see Part 1 for details. REMARKS. Small quantities of a species of Ono- ba were examined from beach sand from the Falk- land Islands (TW4, TW5). They resemble O. proto- fimbriata n. sp. in size and general shell form but were not well enough preserved to allow exami- nation of protoconch microsculpture for confir- mation of the specific identity. Onoba anderssoni (Strebel, 1908) Rissoia anderssoni Strebel, 1908: 55, pi. 4, fig. 54a-c (South Georgia). Onoba anderssoni: Ponder, 1983a: 20, fig. 18c. MATERIAL EXAMINED. South Georgia: SMNH, 28 [11]. REMARKS. The additional specimens are from residues from the same station as the holotype, the species being described from a single specimen. 54 ■ Contributions in Science, Number 445 Onoba filostria (Melvill and Standen, 1912) Rissoa (Onoba) flostria Melvill and Standen, 1912: 349, fig. 9. Onoba filostria: Ponder, 1983a: 14, figs. 9a-e, 13g. MATERIAL EXAMINED. South Georgia: SMNH SSPE, Grytviken [3] (no other data). REMARKS. These four specimens agree rather well with typical material and, if identified cor- rectly, represent the only record of this species from outside the South Orkney Islands. Onoba turqueti (Lamy, 1905) Onoba turqueti: Ponder, 1983a: 16, figs. Ilf, g, 12a-e, 14c-e (gives full synonymy and descrip- tion). MATERIAL EXAMINED. South Georgia: SMNH SSPE 28 [many]. REMARKS. This is the first record of this species from South Georgia. It was previously known from Burdwood Bank, South Orkney Islands, South Shetland Islands, Antarctic Peninsula, and Terre Adelie. Onoba cf. gelida (E.A. Smith, 1907) Onoba gelida: Ponder, 1983a: 20, figs. 13a, b, 16a-d (gives full synonymy and description). MATERIAL EXAMINED. South Georgia: SMNH SSPE 34 [1], REMARKS. A single specimen from the above station represents the second record of this taxon from South Georgia, although well-preserved spec- imens are needed for examination to confirm the species identification. Powellisetia australis (Watson, 1886) Powellisetia australis: Ponder, 1983a: 25-26, figs. 18b, 19d-f (gives full synonymy and description). MATERIAL EXAMINED. Falkland Islands: AMS C. 167489, TW4 [1(d)]; AMS C.167490, TW5 [5(d)]. REMARKS. This is the only record from the Falkland Islands, the species being previously known from Kerguelen and Macquarie Islands. Family Cingulopsidae Skenella georgiana Pfeffer (in Martens and Pfeffer), 1886 Skenella georgiana Pfeffer (in Martens and Pfeffer), 1866: 97, pi. 2, fig. 6a, b. Skenella ( Skenella ) georgiana: Ponder, 1983a: 29, figs. 23a-e, 26a-c (gives full synonymy). MATERIAL EXAMINED. Falkland Islands: SMNH SSPE 40 [3]. Ponder and Worsfold: Review of Rissoiform Gastropods ' H . I JH REMARKS. This species was described from South Georgia and, previously, only two specimens were recorded from the Falkland Islands by Ponder (1983a). Skenella wareni n. sp. Figure 15C, D, G, H ETYMOLOGY. Named after Dr. Anders Wa- ren, as a small mark of recognition for his contri- butions to the study of small gastropods. MATERIAL EXAMINED. Types. Holotype and 4 paratypes (2 damaged), SMNH. SSPE 28. 54°22'S, 36°28' W, South Georgia, 12-15 m, sand and algae, 24 May 1902. 4 Paratypes, SMNH; (1 paratype) AMS C. 167673. SSPE 25. 54°22'S, 36°28'W, South Georgia, 24-52 m, algae, 12 May 1902. DIAGNOSIS. Shell (Fig. 15C, D). Minute (max- imum length 1.6 mm), ovate-conic, moderately thick, with 2. 2-2. 5 teleoconch whorls. Spire with very lightly convex outlines, whorls moderately convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth, with faint, irregular growth lines. Protoconch (Fig. 15D) of about 1.4-1. 6 whorls with very fine spiral grooves on last half whorl. Aperture oval to near circular, weakly angled posteriorly, with sharp peristome and lacking varix (1 from SSPE 25 has strong varix on penultimate whorl). Inner lip moderately broad with indistinct swelling on columella; outer lip moder- ately prosocline. Umbilicus small to moderate. Col- or pale cream, with reddish tinge; reddish-brown when wet. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD Holotype 1.64 1.27 1.29 0.76 2.16 2.5 1.6 0.42 Paratypes 1.38 1.16 1.19 0.72 1.92 2.3 1.5 0.42 1.17 0.86 1.05 0.51 2.30 2.5 1.6 0.35 Paratypes Sta. SSPE 25 1.16 0.88 1.31 0.51 2.27 2.2 1.5 0.36 1.16 0.91 1.27 0.55 2.10 2.4 1.4 0.38 0.98 0.76 1.29 0.44 2.22 2.3 1.4 0.38 1.17 0.84 1.39 0.57 2.07 2.4 1.4 0.36 Operculum (Fig. 15H). Oval, with posterior end angled; peg slightly curved, narrow, extends beyond edge of operculum. Radula (Fig. 15G). Central teeth square, with 2 small, square cusps and moderate lateral thicken- ings. Lateral teeth with cusp formula 3-4 + 1+3- 7, median cusp large, spatulate, lateral cusps slen- der, sharp. Inner marginal teeth with cusp formula 1 + 1 + 1, small pointed outer cusp, and 2 large shov- el-shaped to spatulate inner cusps, median cusp be- ing largest. Outer marginal teeth with 2 large, tri- angular cusps, outermost largest (based on 2 radulae). Animal. Unknown. REMARKS. This species is somewhat interme- diate between S. umbilicata and S. paludinoides in shell shape, being more depressed and convex than S. paludinoides and narrower than S. umbilicata. The reddish-brown color of S. wareni is absent in S. paludinoides and is more pronounced in S. um- bilicata. Both S. paludinoides and S. umbilicata are larger than S. wareni and have broader oper- cular pegs. Skenella wareni is similar to S. palu- dinoides in radular details but more like S. umbil- icata in shell morphology. Some of the paratypes from Sta. SSPE 25 have slightly narrower shells but appear to be identical in other respects. DISTRIBUTION. South Georgia; sublittoral; uncommon. Family Rissoellidae Rissoella (Jeffrey siella) cf. powelli Ponder, 1983 Rissoella [Jeffrey siella) powelli Ponder, 1983a; 32, figs. 26i, 28a-d (Signy Island, South Orkney Is- lands). MATERIAL EXAMINED. South Georgia: SNMH SSPE 25 [1]. REMARKS. A single specimen of a species of Rissoella resembling R. (J.) powelli is recorded from South Georgia. There is not enough material for confident identification. Part 3 Species from Juan Fernandez Islands The opportunity is taken to describe three species from Juan Fernandez Islands that were obtained during the Indian Ocean Expedition in 1965. Family Eatoniellidae Eatoniella ( Eatoniella ) zigzag n. sp. Figure 32D ETYMOLOGY. Zigzag — French (and English). Alternatively changing direction by sharp angles. Refers to the color pattern. MATERIAL EXAMINED. Types. Holotype and 6 paratypes, MCZ 293707, 1 paratype, LACM 2700; 1 para- type, AMS C. 167476. Juan Fernandez Is., R/V Anton Bruun, Cr.12, SE 65 256, 9-12 m. 33°42'S, 78°55'W, S. Earle, 15 Dec. 1965. DIAGNOSIS. Shell (Fig. 32D). Small (maximum length 2.1 mm), conic, moderately thick, with 2.8- 3.4 teleoconch whorls. Spire and whorls lightly convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch smooth and glossy. Protoconch of 1.2- 1.5 whorls. Aperture oval; inner lip narrow, outer lip strongly prosocline. Umbilicus absent. Periostracum very thin, transparent. Color white, usually with pink or purple transverse zigzag lines. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods! 55 Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD Holotype 2.14 1.17 1.83 0.76 2.81 3.4 1.4 0.34 Paratypes 1.87 1.13 1.65 0.73 2.57 3.3 1.3 0.31 1.60 1.07 1.50 0.64 2.50 2.8 1.4 0.34 1.73 1.15 1.50 0.67 2.57 3.2 1.3 0.31 1.68 1.08 1.55 0.65 2.57 3.0 1.2 0.32 1.74 1.10 1.58 0.71 2.47 3.1 1.2 0.31 1.59 1.08 1.47 0.66 2.40 2.9 1.2 0.32 Operculum, radula, and animal unknown. REMARKS. The zigzag color pattern and the thick, conical shell allow this species to be distin- guished from all other described eatoniellids. The only other described species with a somewhat sim- ilar color pattern, E. limbata (Hutton, 1883) from New Zealand, has a larger, broader shell. The shell characters allow little doubt that this species is a member of the Eatoniellidae, although confirma- tion with radular and opercular data is necessary before this can be stated with certainty. DISTRIBUTION. Juan Fernandez Islands. Family Rissoidae Onoba (?) isolata n. sp. Figure 33A, C ETYMOLOGY. Isolata — Latin. Detached, sep- arate. Refers to the isolated location of this species. MATERIAL EXAMINED. Holotype, MCZ 293708, R/V Anton Bruun, Cr.12, SE 65 256, 9-12 m, 33°42'S, 78°55'W, S. Earle, 15 Dec. 1965. DIAGNOSIS. Shell (Fig. 33A, C). Minute (length 1.9 mm), elongate-conic to cylindrical, solid, opaque, with 3.5 teleoconch whorls. Spire with lightly con- vex outlines, whorls lightly convex; periphery of last whorl rounded. Sutures impressed, simple. Teleoconch with strong, rounded spiral cords with wider interspaces and slightly stronger opisthocline axial ribs; 7 spiral cords on penultimate whorl, 15 on last whorl and base. Interspaces smooth. Eleven axial ribs on penultimate whorl; 13 on last whorl; ribs reach suture but do not extend onto base; bluntly tuberculate at point of intersection with spirals. Protoconch (Fig. 33A) of about 1.3 whorls; microsculpture of small granules, some on upper 2/3 of last whorl run together forming broken, al- most zigzag spirals. Aperture oval, rather strongly angled posteriorly. Inner lip moderately broad, at- tached to parietal wall; outer lip opisthocline with moderately distinct posterior notch and prominent apertural varix immediately behind lip. Umbilical chink absent. Periostracum not observed. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD PS BS PA BA Holotype 1.91 0.95 2.02 0.71 2.7 3.5 1.3 0.38 7 15 11 13 Operculum, radula, and animal unknown. REMARKS. Although known from a single spec- imen, this species is readily recognizable by its elon- gate-conic outline, regularly reticulated sculpture, and unusually sculptured protoconch. This distinc- tive species is tentatively included in Onoba on the basis of shell characters, although confirmation is needed. DISTRIBUTION. Juan Fernandez Islands. Onoba (?) protopustulata n. sp. Figure 33B, D ETYMOLOGY. Proto— -Latin. First (whorl of shell). Pustulata — Latin. Bubble, pimple. Refers to the protoconch microsculpture. MATERIAL EXAMINED. Types. Holotype and 1 paratype, MCZ 293707a, R/V Anton Bruun, Cr.12, SE 65 256, 9-12 m, 33°42'S, 78°55'W, S. Earle, 15 Dec. 1965. DIAGNOSIS. Shell (Fig. 33B, D). Small (maxi- mum length 2.0 mm), elongate-conic, solid, opaque, with 3. 6-3. 8 teleoconch whorls. Spire with lightly convex outlines, whorls convex; periphery of last whorl rounded. Sutures impressed, simple. Teleo- conch with sharp spiral cords with wider inter- spaces and prominent axial ribs, 1 spiral cord on lower part of penultimate whorl, 8 on last whorl and base; interspaces smooth; 8 axial ribs on pen- ultimate whorl, 10 on last whorl; axial ribs reach suture but end abruptly at intersection of spiral cord immediately below periphery of last whorl, below which, on base, spirals most significant sculpture. Axials and spirals bluntly tuberculate at intersec- tions. Protoconch (Fig. 33D) of 1.3-1. 5 whorls, with numerous scattered granules loosely arranged in irregular spirals, especially near apex. Aperture oval, angled posteriorly. Inner lip moderately broad, attached to parietal wall; outer lip orthocline with small posterior notch and prominent apertural varix immediately behind lip. Umbilical chink absent. Periostracum not observed. Color white. Dimensions. SL/ SL/ SL SW SW AL AL TW PW PD PS BS PA BA Holotype 1.77 0.96 1.85 0.61 2.89 3.6 1.3 0.29 1 8 8 10 Paratype 2.00 0.99 2.02 0.67 3.00 3.8 1.5 0.32 1 8 8 10 Operculum, radula, and animal unknown. REMARKS. This species has a teleoconch sculp- ture superficially similar to species often included in Alvania, with fewer, much stronger axial ribs than O. isolata and markedly more convex whorls. The protoconch is similar to the last species except that the granules are not fused into irregular lines. We tentatively include this species in Onoba, along with the previous species, because of the likely re- lationship of these two taxa, judging from the teleo- conch and, especially, the protoconch morphology. DISTRIBUTION. Juan Fernandez Islands. 56 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods DISCUSSION The biogeography of the Antarctic and sub-Ant- arctic fauna has produced considerable interest. The area from which most of the material in this report was obtained is the Magellanic Province, which includes Patagonia (from Isla Chiloe on the west coast and Cabo Blanco on the east coast), Tierra del Fuego and the Falkland Islands, and Burdwood Bank (Powell, 1960). Excluding Tierra del Fuego, Chile comprises the majority of this region, and the South American material dealt with in this report is mainly from that country, although a few samples were available from Peru. As can be seen from the data in Table 1, the majority of the 30 South American species-group taxa dealt with in this report are, within South America, found only in southern Chile and Tierra del Fuego or only in one of these areas (22 taxa, 73%), most (15 taxa, 55%) with distributions through southern Chile and Tierra del Fuego, with 7 taxa (23%) known only from Tierra del Fuego. Of the other South American species, one taxon (3%) is found in southern and northern Chile and two (7%) (. Eatoniella nigra and E. glomerosa ) are found from northern Chile to Tierra del Fuego. Of the southern species-group taxa, only six (20%) are also found at the Falkland Islands and only one (O. georgiana) has a wider sub-Antarctic distribution (E. cf. cana being regarded as a separate taxon from E. cana). Five species-group taxa (17%) are only found in northern Chile and/or Peru, and three of these represent families not found farther south (Bar- leeidae, Anabathridae, and Rissoellidae), the other two species belonging to genera ( Manzonia and Eatonina) found farther north on the Pacific Coast of the Americas. The Barleeidae is well represented in western Central and North America (Ponder, 1983b) and A. inclusus is a Panamic species. The Anabathridae, however, as a group, is not well rep- resented in North and Central America but is pri- marily a southern family, with numerous taxa in southern Australasia and southern Africa. The vir- tual absence of this group from South America and its presence in Central and North America, as well as the Mediterranean and eastern Atlantic, suggests that its distribution is not Gondwanan but Pangean. The most detailed biogeographic analysis of the marine fauna of Chile is that of Brattstrom and Johanssen (1983). They confirmed the results of earlier investigations in showing that there are two temperate regions along the Chilean coast, a north- ern warm temperate and southern cold temperate region with the border at about 42°. We have used this same cutoff point to distinguish northern and southern Chile. These authors found that northern species dominate in the littoral and in shallow water and the southern species in deeper water, although they observed that in general the fauna has a “northern stamp” with northern species having wider distributions than southern ones. This is not the case with the families investigated here, the Table 1. South American species listed according to their geographic distributions. IT = intertidal, shelf = on con- tinental shelf, SL = sublittoral. Northern Chile = N of 42°S; southern Chile = S of 42°S. Tierra del Fuego only Eatoniella castanea (IT) Onoba algida (shelf) O. striola (IT) O. sulcula (shelf) O. erugata (IT-SL) O. fuegoensis (shelf) O. lacuniformis (IT-SL) (also Falklands) Southern Chile and Tierra del Fuego Eatoniella turricula (40-900 m) E. denticula (IT-SL) E. cf. denticula (shelf) E. ebenina (IT-SL) E. picea (IT-SL) E. cf. cana (IT-SL) (also Falklands; typical form from South Georgia and South Orkney Islands) Pupatonia magellanica (IT-SL) Powellisetia microlirata (IT-shelf) Pusillina averni and P. cf. averni (shelf) Onoba subincisa (IT-SL) O. scythei (IT-SL) (also Falklands) O. subaedonis (SL) (also Falklands?) O. protofimbriata (IT-SL) (also Falklands?) O. georgiana (IT) (also Falklands and sub-Antarctic) Skenella hallae (SL) Southern and northern Chile Eatoniella ( Albosabula ) mcleani (IT-SL) Northern Chile to Tierra del Fuego Eatoniella nigra (IT-SL) E. glomerosa (IT) Northern Chile and/or Peru Eastonina fusca (IT-SL) Rissoella peruviana (IT-SL) Alvinia limensis (shelf) Barleeia meridionalis (IT-SL) Amphithalamus cf. inclusus (IT-SL?) southern elements dominating the fauna with the few northern elements being limited to northern Chile and Peru. The main elements of the southern fauna are the Eatoniellidae and the genus Onoba (Rissoidae). The eatoniellids are reasonably diverse in the Antarctic and sub-Antarctic but have their greatest diversity in Australasia (Ponder, 1965a; Ponder and Yoo, 1 977a) and southern Africa (Ponder and Y oo, 1 977a; Ponder, unpublished). There are fewer eatoniellids in South America (11 species-group taxa) than in the Antarctic-sub-Antarctic and southern Australia (19 and 18 species-group taxa, respectively) and New Zealand (43 species-group taxa). Onoba, as at pres- ent recognized, is a diverse genus found in cool temperate parts of the northeastern Pacific and north Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 57 Atlantic as well as in the temperate parts of the southern continents (Ponder, 1985a) and Antarctica and the sub-Antarctic islands (Ponder, 1983a). There are 10 taxa from South America, half the number known from the Antarctic and sub-Antarctic. None of the species-group taxa dealt with in this report have protoconchs, suggesting that they pos- sess planktonic larvae, and yet several have consid- erable distributions. Some sub-Antarctic and Ant- arctic species-group taxa also exhibit large ranges, possibly maintaining genetic continuity through dis- persal on drifting algae. Only six of the South Amer- ican species appear to be conspecific with Falkland Islands taxa, although available material from the Falklands is very limited. The poor state of the knowledge of the Falklands fauna is evidenced by the collections made by one of us (TMW), just in the vicinity of Stanley, which revealed five new rec- ords. As might be expected, the three species from the isolated Juan Fernandez Islands are apparently restricted to those islands. The one small sample available does not, however, enable a more defin- itive comment about this fauna. ACKNOWLEDGMENTS We are grateful to Jane Hall, who was involved in the preliminary phase of this project and who did many of the measurements and some of the SEM work, the re- mainder of the scanning being done by Geoff Avern. This work would not have been possible without the considerable amount of time and effort spent by Dr. J.H. McLean of LACM and his volunteers in accumulating and sorting the material on which this paper is based. We thank Dr. McLean for his encouragement and generosity in making available not only the bulk of the material for this report but also a considerable amount of comparative material for this and related projects. We thank Ms. K. Way of the BMNH for the loan of type specimens and Dr. R. Kilias for searching for Philippi material. We thank Dr. R.K. Dell and Mr. B.A. Marshall for the loan and use of collections in the National Museum of New Zealand and Dr. Dell for sending South American material that he had on loan from other institutions. Ma- terial has also been borrowed from the NMNHP, MCZ, NMW, SMNH, and USNM, and we thank the staff of those museums for making it available. Ms. A. Miller and Mr. P. Colman helped check the manuscript and A. Miller prepared the plates and maps. The Spanish translation of the abstract was provided by Mr. Guillermo Herrera. LITERATURE CITED Brattstrom, H., and A. Johanssen. 1983. Ecological and regional zoogeography of the marine benthic fauna of Chile. Report no. 49 of the Lund University Chile Expedition 1948-49. Sarsia 68:233-339. Carcelles, A.R. 1950. Catalogo de los Moluscos Mari- nos de Patagonia. Anales del Museo Nahuel Huapi Perito Dr. Francisco P. Moreno 2:41-100. Carcelles, A.R. , and S.I. Williamson. 1951. Catalogo de los Moluscos marinos de la Provincia Magellanica. Revista del Instituto Nacional de investigacion de las ciencias naturales. Ciencias Zoologicas 2:225- 383. 58 ■ Contributions in Science, Number 445 Castellanos, Z.J. A., and D. Fernandez. 1972a. Unanueva specie de Eatoniella (Mollusca, Rissoacea). Neotro- pica 18:6-8. . 1972b. Resultados de la Campana Exploratoria S.A.O.I. — 1971. Nuevos Moluscos para Golfo San Matias. 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Transactions of the Royal Society of Ed- inburgh 48:33-366. Orbigny, A.d’ 1835-1846. Voyage dans L’ Amerique Meridionale. Tome Cinquieme. 3rd part: Mol- lusques. Paris, 377-408 (1840). . 1854. List of the shells of South America in the collection of the British Museum. London: British Museum, 89 pp. Philippi, R.A. 1868. Conchylia nova potissimum ma- gellanica. Malakozoologische Blatter 1868:222-226. Ponder, W.F. 1965a. The family Eatoniellidae in New Zealand. Records of the Auckland Institute and Mu- seum 6:47-99. — . 1965b. A revision of the New Zealand Recent species previously known as Notosetia Iredale, 1915 (Rissoidae, Gastropoda). Records of the Auckland Institute and Museum 6:101-131. — . 1983a. Rissoaform gastropods from the Ant- arctic and sub-Antarctic. 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Annals and Magazine of Natural History, series 8, 11:218-223. Smith, E.A. 1907. Mollusca and Brachiopoda. National Antarctic Expedition (S.S. Discovery,) 1901-04, Natural History, 2:1-12. Strebel, H. 1908. Die Gastropoden. Wissenschaftliche Ergehnisse der schwedischen Siidpolar Expedition. 1901-1903 6:1-112. Watson, R.B. 1886. Report on Scaphopoda and Gas- teropoda collected by H.M.S. ‘Challenger’ during the years 1873-76. Challenger Report 15:1-756. Received 4 March 1992; accepted 11 March 1993. APPENDIX: LIST OF LOCALITIES NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY (LACM) LOCALITIES The localities are listed in order according to station num- ber. The number in parentheses following the station number are the locality reference points used in Figures !-4- 64-16 (6A). Iquique, near end of Ave Baquedana, Tara- 1 paca Prov., Chile. 20°13'S, 70°10'W, intertidal, L. Marin- covich, June-Aug. 1964. 70-66 (6A). end of Ave Baquedana, Iquique, Tarapaca Prov., Chile. 20°13'S, 70°10'W, intertidal rocks, L. Mar- incovich, July 1970. 71-258 (33A). Bahia Crossley, Isla de los Estados, Tierra del Fuego, Argentina. 54°47.1'S, 64°42.1'W, 13-37 m,Sta. 658 and 5257, USARP, R/V Hero Cr.712, 28-29 Apr. 1971. 71-259 (30G). Puerto Vancouver, Isla de los Estados, Tierra del Fuego, Argentina. 54°46.8'S, 64°04'W, 31 m, Sta. 663, USARP, R/V Hero Cr.712, 9 May 1971. 71-260 (30J). Puerto San Juan del Salvamento, Isla de los Estados, Tierra del Fuego, Argentina. 54°44.85'S, 63°52.9'W, 44 m, Sta. 665, USARP, R/V Hero Cr.712, 11 May 1971. 71-262 (33N). W side Puerto Ano Nuevo, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.1'S, 64°7.3' W, 50 m, Sta. 672, USARP, R/V Hero Cr.712, 19 May 1971. 71-263 (33M). Puerto Basil Hall, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.5'S, 64°9.8'W, Sta. 673,674, USARP, R/V Hero Cr.712, 20 May 1971. 71-264 (33J). 6.6 km E Cabo Colnett, Isla de los Es- tados, Tierra del Fuego, Argentina. 54°43.7'S, 64°14.2'W, 18 m, Sta. 676, USARP, R/V Hero Cr.712, 22 May 1971. 71-265 (33F). E arm Bahia San Antonio, Isla de los Estados, Tierra del Fuego, Argentina. 54°46.5'S, 64°23.5' W, 51 m, Sta. 677, USARP, R/V Hero Cr.712, 24 May 1971. 71-266 (33G). 1.5 km E Cabo Colnett, Isla de los Es- tados, Tierra del Fuego, Argentina. 54°43.3'S, 64°19.8'W, 14 m, Sta. 678, USARP, R/V Hero Cr.712, 25 May 1971. 71-267 (33B). W arm Bahia San Antonio, Isla de los Estados, Tierra del Fuego, Argentina. 54°46.1'S, 64°25.1'W, 36 m, Sta. 679, USARP, R/V Hero Cr.712, 26 May 1971. 71-268 (39). Punta Arenas, Bahia de San Sebastian, Ti- erra del Fuego, Argentina. 53°9.2'S, 68°14.6'W, intertidal, rocks and mussel beds, Sta. 71-2-3, USARP, R/V Hero Cr.712, 20 Apr. 1971. 71-270 (28). W side Bahia Buen Suceso, Tierra del Fuego, Argentina. 54°47.8'S, 65T6'W, intertidal, sand, rock, and mussels, Sta. 71-2-8, USARP, R/V Hero Cr.712, 23 Apr. 1971. 71-271 (28). N side Bahia Buen Suceso, Tierra del Fue- go, Argentina. 54°48.2'S, 65°14.7'W, intertidal rocks, Sta. 71-2-14, USARP, R/V Hero Cr.712, 25 Apr. 1971. 71-273 (32A). Bahia Crossley, Isla de los Estados, Tierra del Fuego, Argentina. 54°46.2'S, 64°42.7'W, intertidal, rocks and mussel beds, Sta. 71-2-16, USARP, R/V Hero Cr.712, 27 Apr. 1971. 71-274 (31A). E arm Bahia Capitan Canepa, Isla de los Estados, Tierra del Fuego, Argentina. 54°50'S, 64°26.8' W, intertidal rocks and mussel beds, Sta. 71-2-19, USARP, R/V Hero Cr.712, 3 May 1971. 71-275 (31A). N arm Bahia Capitan Canepa, Isla de los Estados, Tierra del Fuego, Argentina. 54°49.2'S, 64°27.8' W, intertidal rocks and mussel beds, Sta. 71-2-19, USARP, R/V Hero Cr.712, 3 May 1971. 71-276 (31B). NW arm Bahia York, Isla de los Estados, Tierra del Fuego, Argentina. 54°47.3'S, 64°18.7'W, inter- tidal, mussels and rocks. Sta. 71-2-21,71-2-25, USARP, R/V Hero Cr.712, 4,6 May 1971. 71-277 (31B). NW arm Bahia York, Isla de los Estados, Tierra del Fuego, Argentina. 54°47.15'S, 64°17.9'W, in- tertidal rocks, Sta. 71-2-22, USARP, R/V Hero Cr.712, 5 May 1971. 71-281 (31C). N end Bahia Blossom Isla de los Estados, Tierra del Fuego, Argentina. 54°46.4'S, 63°57.7'W, inter- tidal rocks, Sta. 71-2-30, USARP, R/V Hero Cr.712, 10 May 1971. 71-283 (31D). NE part Puerto San Juan del Salvamento, Isla de los Estados, Tierra del Fuego, Argentina. 54°44.25'S, Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 59 63°51.25'W, intertidal rocks, Sta. 71-2-32, USARP, R/V Hero Cr.712, 12 May 1971. 71-286 (32H). Puerto Cook, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.4'S, 64°2.5'W, intertidal rocks and mussel beds, Sta. 71-2-36, USARP, R/V Hero Cr.712, 16 May 1971. 71-287 (32H). Puerto Cook, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.25'S, 64°2.3'W, intertidal, Sta. 71-2-37, USARP, R/V Hero Cr.712, 17 May 1971. 71-289 (32E). Puerto Basil Hall, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.45'S, 64°10.1'W, in- tertidal rocks, Sta. 71-2-39, USARP, R/V Hero Cr.712, 20 May 1971. 71-290 (32E). Puerto Basil Hall, Isla de los Estados, Tierra del Fuego, Argentina. 54°45.45'S, 64°9.55'W, in- tertidal rocks, Sta. 71-2-40, USARP, R/V Hero Cr.712, 21 May 1971. 71-291 (32D). 3.3 km W Puerto Basil Hall, Isla de los Estados, Tierra del Fuego, Argentina. 54°43.85'S, 64°13.6'W, intertidal rocks, Sta. 71-2-41, USARP, R/V Hero Cr.712, 22 May 1971. 71-293 (32D). 3.3 km W Puerto Basil Hall, Isla de los Estados, Tierra del Fuego, Argentina. 54°43.3'S, 64°14'W, intertidal rocks, Sta. 71-2-43, USARP, R/V Hero Cr.712, 23 May 1971. 71-294 (32C). S end of E arm Bahia San Antonio, Isla de los Estados, Tierra del Fuego, Argentina. 54°47.6'S, 64°22.35'W, intertidal rocks, Sta. 71-2-44, USARP, R/V Hero Cr.712, 24 May 1971. 71-295 (32B). SW arm Bahia San Antonio, Isla de los Estados, Tierra del Fuego, Argentina. 54°46.2'S, 64°24.7' W, intertidal rocks, Sta. 71-2-46, USARP, R/V Hero Cr.712, 26 May 1971. 71-296 (28). Bahia Buen Suceso, Tierra del Fuego, Ar- gentina. 54°47.9'S, 65°14.7'W, 10 m, Sta. 680, USARP, R/V Hero Cr.715, 13 Oct. 1971. 71-302 (29A). 8 km E Caleta San Mauricio, Tierra del Fuego, Argentina. 54°45'S, 65°4.6'W, 75 m, Sta. 687, USARP, R/V Hero Cr.715, 14 Oct. 1971. 71-305 (29B). 12.5 km E Ensenada Patagones, Tierra del Fuego, Argentina. 54°52'S, 65°05'W, 144 m, Sta. 690, USARP, R/V Hero Cr.715, 16 Oct. 1971. 71-308 (33A). Bahia Crossley, Isla de los Estados, Tierra del Fuego, Argentina. 54°46.6'S, 64°41.4'W, 9 m, Sta. 694, USARP, R/V Hero Cr.715, 17 Oct. 1971. 71-309 (32A). Bahia Crossley, Isla de los Estados, Tierra del Fuego, Argentina. 54°47.6'S, 64°40.7'W, intertidal, mussels and rocks. Sta. 695, USARP, R/V Hero Cr.715, 17 Oct. 1971. 71-310 (330). Isla Observatorio, Isla de los Estados, Tierra del Fuego, Argentina. 54°39.5'S, 64°7.1'W, subtid- al, Sta. 698, USARP, R/V Hero Cr.715, 19 Oct. 1971. 71-311 (32G). Observatorio, Isla de los Estados, Tierra del Fuego, Argentina. 54°39.5'S, 64°08'W, intertidal rocks, Sta. 699, USARP, R/V Hero Cr.715, 19 Oct. 1971. 71-312 (33G). 6.4 km N Cabo Colnett, Isla de los Estados, Tierra del Fuego, Argentina. 54°39'S, 64°20'W, 48 m, Sta. 852, USARP, R/V Hero Cr.715, 20 Oct. 1971. 71-313 (33H). 14.4 km N Cabo Colnett, Isla de los Estados, Tierra del Fuego, Argentina. 54°34'S, 64°20'W, 91 m, Sta. 853, USARP, R/V Hero Cr.715, 20 Oct. 1971. 71-315 (33K). 16 km N Isla Observatorio, Isla de los Estados, Tierra del Fuego, Argentina. 54°29'S, 64°10'W, 1 10 m, Sta. 855, USARP, R/V Hero Cr.715, 20 Oct. 1971, 71-316 (33L). 8 km N Isla Observatorio, Isla de los Estados, Tierra del Fuego, Argentina. 54°34'S, 64°10'W, 73 m, Sta. 856, USARP, R/V Hero Cr.715, 20 Oct. 1971. 71-317 (331). 3.3 km W Puerto Basil Hall, Isla de los 60 ■ Contributions in Science, Number 445 Estados, Tierra del Fuego, Argentina. 54°43.9'S, 64°14.1'W, 10 m, Sta. 861, USARP, R/V Hero Cr.715, 21 Oct. 1971. 71-319 (33P). 9.6 km NE Isla Observatorio, Isla de los Estados, Tierra del Fuego, Argentina. 54°35.5'S, 63°58.7'W, 87 m, Sta. 864, USARP, R/V Hero Cr.715, 22 Oct. 1971. 71-323 (32H). Puerto Cook, Isla de los Estados, Tierra de! Fuego, Argentina. 54°45.6'S, 64°2.6'W, intertidal rocks and mussel beds, Sta. 869, USARP, R/V Hero Cr.715, 23 Oct. 1971. 71-326 (32F). Puerto Aho Nuevo, Isla de los Estados, Tierra del Fuego, Argentina. 54°44.9'S, 64°6.6'W, inter- tidal rocks and mussel beds, Sta. 872, USARP, R/V Hero Cr.715, 25 Oct. 1971. 71-327 (33Q). 14.4 km N Cabo San Juan, E end Isla de los Estados, Tierra del Fuego, Argentina. 54°34'S, 63°50'W, 118 m, Sta. 873, USARP, R/V Hero Cr.715, 26 Oct. 1971. 71-328 (33R). 6.4 km N C. San Juan, Isla de los Estados, Tierra del Fuego, Argentina. 54°39.1'S, 63°50.TW, 135— 137 m, Sta. 874, USARP, R/V Hero Cr.715, 26 Oct. 1971. 71-329 (301). 8 km S Pta. Ventana, S side Isla de los Estados, Tierra del Fuego, Argentina. 54°54.5'S, 63°56'W, 771-903 m, Sta. 875, USARP, R/V Hero Cr.715, 27 Oct. 1971. 71-332 (30H). 3.3 km S Bahia Blossom, Isla de los Estados, Tierra del Fuego, Argentina. 54°50'S, 63°59.7' W, 205-208 m, Sta. 880, USARP, R/V Hero Cr.715, 28 Oct. 1971. 71-334 (30F). 3.3 km W Cabo Kendall, Isla de los Estados, Tierra del Fuego, Argentina. 54°50'S, 64°10'W, 76 m, Sta. 882, USARP, R/V Hero Cr.715, 29 Oct. 1971. 71-339 (30E). NW arm Bahia York, Isla de los Estados, Tierra del Fuego, Argentina. 54°47.2'S, 64°18.4'W, 38 m, Sta. 891, USARP, R/V Hero Cr.715, 1 Nov. 1971. 71-340 (30D). 8 km SE Cabo Kempe, Isla de los Es- tados, Tierra del Fuego, Argentina. 54°55.2'S, 64°20.4'W, 303-358 m, Sta. 893, USARP, R/V Hero Cr.715, 2 Nov. 1971. 71-342 (30A). 11.2 km SW Cabo San Bartolome, Isla de los Estados, Tierra del Fuego, Argentina. 55°00'S, 64°48.7'W, 438-548 m, Sta. 895, USARP, R/V Hero Cr.715, 3 Nov. 1971. 71-344 (30C). Bahia Capitan Canepa, Isla de los Esta- dos, Tierra del Fuego, Argentina. 54°50.2'S, 64°29.4'W, 67-71 m, Sta. 897, USARP, R/V Hero Cr.715, 4 Nov. 1971. 71-345 (31A). S side Bahia Capitan Canepa, Isla de los Estados, Tierra del Fuego, Argentina. 54°50.8'S, 64°29'W, intertidal rocks and mussel beds, Sta. 898, USARP, R/V Hero Cr.715, 4 Nov. 1971. 71-346 (30B). 2.4 km E Cabo San Bartolome, Isla de los Estados, Tierra del Fuego, Argentina. 54°55'S, 64°40'W, Grab smpl, Sta. 899, USARP, R/V Hero Cr.715, 5 Nov. 1971. 71-347 (33B). 28.8 km N Bahia Crossley, Isla de los Estados, Tierra del Fuego, Argentina. 54°29.3'S, 64°40.4'W, 116-120 m, Sta. 902, USARP, R/V Hero Cr.715, 6 Nov. 1971. 71-348 (33C). 20.8 km N Bahia Crossley, Isla de los Estados, Tierra del Fuego, Argentina. 54°34.3'S, 64°40'W, 84-85 m, Sta. 903, USARP, R/V Hero Cr.715, 6 Nov. 1971. 71-351 (33E). 22.6 km N Cabo San Antonio, Isla de los Estados, Tierra del Fuego, Argentina. 54°29'S, 64°29.2'W, 122-124 m, Sta. 906, USARP, R/V Hero Cr.715, 7 Nov. 1971. 71-352 (33D). 14.4 km N Cabo San Antonio, Isla de los Estados, Tierra del Fuego, Argentina. 54°34'S, 64°30'W, Ponder and Worsfold: Review of Rissoiform Gastropods 73-76 m, Sta. 907, USARP, R/V Hero Cr.715, 7 Nov. 1971. 71- 357 (33F). E arm Bahia San Antonio, Isla de los Estados, Tierra del Fuego, Argentina. 54°46.5'S, 64°23.3'W, Grab smpl, Sta. 916, USARP, R/V Hero Cr.715, 10 Nov. 1971. 72- 76 (3). Pucusana (Chilca), Lima Prov., Peru. 12°30'S, 76°49'W, 0-6 m, J.H. McLean, 30 Mar. and 10 Apr. 1972. 72- 78 (4). Isla Chincha Norte, lea Prov., Peru. 13°38'S, 76°25'W, 6-12 m, on exposed side, J.H. McLean, V. Alamo, 1 Apr. 1972. 73- 66 (33G). Isla Colnett, off N side Isla de los Estados, Tierra del Fuego, Argentina. 54°43.45'S, 64°14.3'W, 15 m, black sand, P. Dayton, (R/V Hero), 4 May 1973. 73-68 (26). Beagle Canal, Caleta Awaiakirrh, Tierra del Fuego, Chile. 55°00'S, 69°2.2'W, 17 m, P. Dayton, (R/V Hero), 16 May 1973. 73-69 (25). Punta Valparaiso, Canal Cockburn, Ma- gallanes Prov., Chile. 54°22.2'S, 71°21.7'W, 15 m, P. Day- ton, (R/V Hero), 17 May 1973. 73-70 (23). Isla Carlos III, Strait of Magellan, Magalla- nes Prov., Chile. 53°39.4'S, 72°14.8'W, 11-12 m, P. Day- ton, (R/V Hero), 18 May 1973. 73-71 (21). Punta Dashwood, Canal Smyth, Magallanes Prov., Chile. 52° 24'S, 73°39.7'W, 12 m, P. Dayton, (R/V Hero), 19 May 1973. 73-72 (20). Bahia Tom, Magallanes Prov., Chile. 50°11.3'S, 74°47.9'W, 14 m, P. Dayton, (R/V Hero) 21 May 1973. 73-73 (19). Bahia San Andres, N of Golfo de Penas, Aisen Prov., Chile. 46°35.3'S, 75°30.6'W, subtidal, P. Day- ton, (R/V Hero), 23 May 1973. 73-74 (18). Canal Darwin, Aisen Prov., Chile. 45°27.8'S, 74°24.8'W, 8 m, P. Dayton, (R/V Hero), 24 May 1973. 73- 75 (17). Isla Westhoff, Chiloe Prov., Chile. 43°54'S, 73°43.5'W, 23 m, P. Dayton, (R/V Hero), 25 May 1973. 74- 6 (1). Isla Lobos de Afuera (NW and NE of isthmus), Peru. 6°57.1'S, 80°42.3'W, 2-10 m, rocks and sand, J.H. Mclean, J.A. Coyer, and J.M. Engle, 19-20 Jan. 1974. 74- 24 (2B). Isla San Lorenzo, midway on NE side, rocky point N of naval base, Lima Prov., Peru. 12°5.75'S, 77°12.9'W, 0-4.5 m, J.H. McLean, J.A. Coyer, and J.M. Engle, 29 Jan. 1974. 75- 10 (6B). Pozo Toyo, S of Iquique, Tarapaca Prov., Chile. 20°25'S, 70°10.5'W, intertidal, Sta. 1, J.H. McLean, 29 Sept, and 1 Oct. 1975. 75-12 (6A). Iquique, marine lab. of Universidad del Norte, Tarapaca Prov., Chile. 20°15.5'S, 70°08'W, inter- tidal, Sta. 3, J.H. McLean, 30 Sept, and 2 Oct. 1975. 75-15 (8C). S end of Antofagasta, Antofagasta Prov., Chile. 23°42'S, 70°27'W, intertidal Piura beds, Sta. 6, J.H. McLean, J. Tomicic, 5,6 Oct. 1975. 75-17 (8A). Mainland E of Isla Santa Maria, N of An- tofagasta, Antofagasta Prov., Chile. 23°25'S, 70°36'W, in- tertidal, Sta. 8, J.H. McLean and J. Tomicic, 7 Oct. 1975. 75-19 (8B). Los Colorados, rocky headland N of Bahia Antofagasta, Antofagasta Prov., Chile. 23°29'S, 70°22'W, 0-6 m, Sta. 10, J.H. McLean, 9 Oct. 1975. 75-20 (8C). Antofagasta, S end of city, Antofagasta Prov., Chile. 23°42'S, 70°27'W, 2-3 m, out from Piura beds, Sta. 11. J.H. McLean, 10 Oct. 1975. LACM. 75-21 (7). NW of Mejillones, N of Antofagasta, An- tofagasta Prov., Chile. 23°02'S, 70°31'W, 8-23 m, Sta. 12, in Aulacomya beds, J.H. McLean et al., 11 Oct. 1975. 75-25 (9). S side of Bahia Herradura, S of Coquimbo, Coquimbo Prov., Chile. 29°59'S, 71°22'W, 6-17 m, Sta. 16, J.H. McLean, 14 Oct. 1975. 75-28 (10). Los Molles, Aconcagua Prov., Chile. 32°14'S, 71°32'W, intertidal, Sta. 19, J.H. McLean, 16-18 Oct. 1975. 75-30 (11). Estacion de Biol. Marina, Montemar, Val- paraiso Prov., Chile. 32°57'S, 71°32'W, intertidal, Sta. 20, J.H. McLean, 19,20 Oct. 1975. 75-33 (12). Algarrobo, Valparaiso Prov., Chile. 33°22'S, 71°42'W, 3-8 m, Sta. 23, T. Suchanek, K. Sebens, 23 Oct. 1975. 75-37 (13). Mehuin, small offshore island out from Rio Lingue, Valdivia Prov., Chile. 39°26'S, 73°16'W, intertidal, Sta. 27, J.H. McLean, 1 Nov. 1975. 75-41 (15). Pumalin, W of Isla Talcan, Golfo de Cor- covado, Chiloe Prov., Chile. 42°42'S, 72°52'W, intertidal, Sta. 31, J.H. McLean, 4-6 Nov. 1975. 75-43 (14). Islote Nihuel, Golfo de Corcovado, Chiloe Prov., Chile. 42°38'S, 72°57'W, 3-14 m, Sta. 33, J.H. Mc- Lean, 7 Nov. 1975. 75-46 (16). E side Isla Laitec, off SE end of Isla de Chiloe, Chiloe Prov., Chile. 43°14'S, 73° 36'W, 3-6 m, Sta. 36, J.H. McLean, 9 Nov. 1975. 75-48 (24). Punta Santa Ana, Fuerte Bulnes, Peninsula Brunswick, Strait of Magellan, Chile. 53°38'S, 70°54.5'W, intertidal, Sta. 38, J.H. McLean, 16 Nov. 1975. 75-49 (24). Puerto el Hambre, cove S of Punta Askew, Penin. Brunswick, Strait of Magellan, Chile, 53°37'S, 70°56'W, intertidal, J.H. McLean, 16,19 Nov. 1975. 75-51 (38). S of Rio Grande, Atlantic Coast, Tierra del Fuego, Argentina. 53°02'S, 70°49'W, intertidal, Sta. 41, J.H. McLean, 22 Nov. 1975. 35-147 (2C). Near rocks off Isla San Lorenzo, Calloa, Lima Prov., Peru. 12°09'S, 77°15'W, 46 m, R/V Velero III, Sta. BS-522, 11 Jan. 1935. 35-156 (5). E of Isla Viejas, Bahia Independencia, lea Prov., Peru. 14°16'S, 76°10'W, 37 m, R/V Velero III, Sta. BS-531, 13 Jan. 1935. 35-159 (4). N of Isla Medio, Islas de Chincha, lea Prov., Peru. 13°39'S, 76°22'W, 33 m, R/V Velero III, Sta. BS- 534, 15 Jan. 1935. 38-208 (2A). Off Hormigas de Afuera, Lima Prov., Peru. 11°57'S, 77°47'W, 82 m, R/V Velero III, Sta. BS-569, 12 Feb. 1938. NATIONAL MUSEUM OF NATURAL HISTORY, WASHINGTON, D.C. (USNM) MATERIAL The localities are listed in order according to station num- ber. The number in parentheses following the station number are the locality reference points used in Figures 1-4. E = Eltain station, H = Hero cr 712 station, V = Vema station. E 219 (27C). 55°47'S, 66°17'W, 115 m, off Cape Horn. E 363 (34). 57°09'S, 58°58'W, east of Tierra del Fuego. E 740 (27D). 56°06'S, 66° 19'W, 384-494 m, off Cape Horn. E 958 (22B). 5 2°56'S, 75°00'W, 92-101 m, west of Tierra del Fuego. E 960 (22A). 52°40'S, 74°58'W, 64 m, west of Tierra del Fuego. E 967 (37B). 53°42'S, 66°19'W, 81 m, east of Tierra del Fuego. E 974 (36). 53°42'S, 64°57'W, 119-124 m, east of Tierra del Fuego. E 1596 (35). 54°39'S, 57°09'W, 124 m, Burdwood Bank. V 17-48 (27B). 55° 10'S, 66°23'W, 42 m, off eastern Tierra del Fuego. H 654 (30A). 54°40'S, 65°14'W, 12 m, off eastern Tierra del Fuego. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 61 H 656 (33A). 54°48'S, 64°42'W, 18 m, off eastern Tierra del Fuego. H 659 (30C). 54°51.5'S, 64°27.1'W, ? m, off eastern Tierra del Fuego. H 664 (30H). 54°46.1'S, 63°57.9'W, 29 m, off eastern Tierra del Fuego. Sta. 2778 (24), 212271, Strait of Magellan, 53°01.00'S, 70°42.25'W, 111 m, 23 Jan. 1887. 368419, Above battle monument, Port Stanley, Falk- land Islands, boat dredge through kelp, W.L. Schmitt. 23 Feb. 1927. NATURAL HISTORY MUSEUM (LONDON) (BMNH) MATERIAL The localities are listed in order according to station num- ber. The number in parentheses following each station number is the locality reference point used in Figures 1- 4. DE = Discovery Expedition. DE 88 (37A). 54°00'S, 65°00'W, east of Tierra del Fue- go, 118 m. DE Sta. 388 (27E). 56°19.3'S, 67°09.45'W, off Cape Horn, 121 m, 16 Apr. 1930. DE Sta. 399. 1 mi southeast of southwest point of Gough I., Tristan da Cunha, 141-102 m, 18 Apr. 1930. NATIONAL MUSEUM OF WALES (NMW) MATERIAL Scottish National Antarctic Expedition (SNAE) Station. SNAE Sta. 346 (35). 54°25'S, 57°32'W, Burdwood Bank, 102 m, 1 Dec. 1903. SWEDISH MUSEUM OF NATURAL HISTORY (SMNH) MATERIAL Swedish Southpolar Expedition (SSPE) 1901-1903 Sta- tions. Full list in Strebel, 1908: 1-6. SSPE 3, 54°43'S, 64°08'W, Tierra del Fuego, 36 m, 6 Jan. 1902. SSPE 25, 54°22'S, 36°27'W, South Georgia, 24-52 m, some algae, 12 May 1902. SSPE 28, 54°22'S, 36°28'W, South Georgia, 12-15 m, sand and algae, 24 May 1902. SSPE 34, 54°1TS, 36°18'W, center of Cumberland Bay, South Georgia, 252-310 m, a few stones, 5 June 1902. SSPE 39, Port William, Falkland Islands, 51°40'S, 57°4TW, 40 m, sand, stones, algae, 4 July 1902. SSPE 40, 51°33'S, 58°W, Berkeley Sound, Falkland Is- lands, 16 m, gravel and shells with algae, 19 July 1902. SSPE 43, 5 1°33'S, 58°09'W, Greenpatch, Port Louis, near bridge, Falkland Islands, shallow subtidal, stony bot- tom with algae amongst Macrocystis, 28 July 1902. SSPE 46, 51°32'S, 58°07'W, Carenage Creek, Port Lou- is, Falkland Islands, 1 m, sand with Codium, 9 Aug. 1902. SSPE 48, 51°34'S, 57°55'W, Berkely Sound, Falkland Islands, 25 m, sand and stones, 10 Aug. 1902. SSPE 59, 53°45'S, 61°10'W, Burdwood Bank, 137-150 m, broken shells and stones, 12 Sep. 1902. T. WORSFOLD STATIONS The localities are listed in order according to station num- ber. Voucher material is lodged in the AMS. TW1: Off Sparrow Pt., Port William, East Falkland, 51°39.95'S, 57°48.04'W, 3 m, coarse sand, moderately sheltered, 24 June, 1990. TW2: 2 km east of Stanley, East Falkland, 51°41.73'S, 57°48.83'W, intertidal, filamentous algae, sheltered inlet, 24 Feb. 1990. TW3: 1 km west of Stanley, East Falkland, 51°41.43'S, 57°52.52'W, intertidal, filamentous algae, sheltered inlet, 25 Feb. 1990. TW4: Gypsy Cove, East Falkland, 51°40.57'S, 57°48.22'W, intertidal shell sand, sheltered bay, 8 Apr. 1990. TW5: Surf Bay, East Falkland, 51°41.90'S, 57°46.35'W, intertidal shell sand, exposed coast, 24 Feb. 1990. LOCALITY NUMBERS Numbers 1 see Figure 2A; 2-8, see Figure 2B; 9-17, see Figure 2C; 18-27, 36-39, see Figure 3; 28-33, see Figure 4; 34-35, see Figure 1. Peru (Fig. 2A, B) 1. 74-6. 2A. AHF 569. 38-208. 2B. 74-24. 2C. AHF 522. 35-147. 3. 72-76. 4. 72-78, AHF 534. 35-159. 5. AHF 531. 35-156. Northern Chile (Fig. 2B, C) 6A. 64-16, 70-66, 8C. 75-15, 75-20 75-12. 9. 75-25. 6B. 75-10. 10. 75-28. 7. 75-21. 11. 75-30. 8A. 75-17. 12. 75-33. 8B. 75-19. 13. 75-37. Southern Chile (Figs. 2C, 3) 14. 75-43. 21. 73-71. 15. 75-41. 22 A. E 960. 16. 75-46. 22B. E 958. 17. 73-75. 23. 73-70. 18. 73-74. 24. 75-48, 75-49, 19. 73-73. USNM Sta. 2778 20. 73-72. Tierra del Fuego (Figs. 1, 3, 4) 25. 73-69. 31A. 71-274, 71-275. 26. 73-68. 71-345. 27A. “Baie Orange,” 31B. 71-276, 71-277, Mission du Cape 31C. 71-281. Horn (NMNHP). 31D. 71-283. 27B. V 17-48. 32A. 71-273, 71-309. 27C. E 219. 32B. 71-267, 71-295. 27D. E 740. 32C. 71-294. 27E. DE 388. 32D. 71-291, 71-293. 28. 71-270, 71-271, 32E. 71-289, 71-290. 71-296. 32F. 71-326. 29A. 71-302. 32G. 71-311. 29B. 71-305. 32H. 71-286, 71-287. 30A. 71-342, H 654. 71-323. 30B. 71-346. 33A. 71-258, 71-308. 30C. 71-344, H 659. H 656. 30D. 71-340. 33B. 71-267, 71-347. 30E. 71-339, 71-359. 33C. 71-348. 30F. 71-334. 33D. 71-352. 30G. 71-259. 33E. 71-351. 30H. 71-332, H 664. 33F. 71-265, 71-357. 301. 71-329. 33G. 71-266, 71-312, 30J. 71-260. 73-66. 62 ■ Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods 33 H. 331. 33 J. 33K. 33L. 33M. 33N. 330. 33P. 71-313. 33Q. 71-327. 71-317. 33R. 71-328. 71-264. 34. E 363. 71-315. 35. E 1596, SNAE 346 71-316. 36. E 974. 71-263. 37A. DE 88. 71-262. 37B. E 967. 71-310. 38. 75-51. 71-319. 39. 71-268. Contributions in Science, Number 445 Ponder and Worsfold: Review of Rissoiform Gastropods ■ 63 Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Number 446 12 August 1994 si/ Contributions in Science CORYNEXOCHIDA AND PTYCHOPARIIDA (Trilobita, Arthropoda) of the Ehmaniella Biozone (Middle Cambrian), Utah and Nevada Frederick A. Sundberg Natural History Museum of Los Angeles County Serial Publications of THE Natural History Museum of Los Angeles County Scientific Publications Committee James L. Powell, Museum President Daniel M. Cohen, Committee Chairman Brian V. Brown Kenneth E. Campbell Kirk Fitzhugh Edward C. Wilson Robin A. Simpson, Managing Editor The scientific publications of the Natural History Mu- seum of Los Angeles County have been issued at irregular intervals in three major series; the issues in each series are numbered individually, and numbers run consecutively, regardless of the subject matter. • Contribu tions in Science, a miscellaneous series of tech- nical papers describing original research in the life and earth sciences. • Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth sci- ences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now published by the Museum in Contributions in Science. • Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through the Museum Book Shop. A catalog is available on request. The Museum also publishes Technical Reports, a mis- cellaneous series containing information relative to schol- arly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum’s Scientific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Tech- nical Reports may be obtained from the relevant Section of the Museum. Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Printed at Allen Press, Inc., Lawrence, Kansas ISSN 0459-8113 CORYNEXOCHIDA AND PTYCHOPARIIDA (Trilobita, Arthropod a) of the Ehmaniella Biozone (Middle Cambrian), Utah and Nevada Frederick A. Sundberg1 CONTENTS ABSTRACT 4 INTRODUCTION . 4 STRATIGRAPHY AND DEPOSITIONAL SETTING 6 BIOSTRATIGRAPHY 7 Proehmaniella Subzone 9 Elrathiella Subzone 9 Ehmaniella Subzone 10 Altiocculus Subzone 11 Chronostratigraphic Correlations 13 SYSTEMATIC PALEONTOLOGY 14 Species and Subspecies 14 Measurements, Ratios, and Terminology 15 Phylum Arthropoda Siebold and Stannius, 1845 16 Class Trilobita Walch, 1771 16 Order Corynexochida Kobayashi, 1935 17 Family Dorypygidae Kobayashi, 1935 17 Genus Dorypyge Dames, 1883 17 Dorypyge swasii, new species 17 Genus Kootenia Walcott, 1889 19 Kootenia dawsoni (Walcott, 1889) 21 Kootenia cf. K. dawsoni (Walcott, 1889) 24 Kootenia acicularis Resser, 1939b 24 Kootenia cf. K. havasuensis Resser, 1945 26 Kootenia ? margoindistincta, new species 27 Kootenia quadriceps (Hall and Whitfield, 1877) 29 Genus Olenoides Meek, 1877 31 Olenoides serratus (Rominger, 1887) 31 Family Oryctocephalidae Beecher, 1897 31 Subfamily Oryctocephalinae Beecher, 1897 31 Genus Opsiosoryctocephalus , new genus 33 Opsiosoryctocephalus ophis, new species 33 Subfamily Tonkinellinae Reed, 1934 35 Genus Tonkinella Mansuy, 1916 35 Tonkinella breviceps Kobayashi, 1934 35 Tonkinella valida Tchernysheva, 1962 37 Family Dolichometopidae Walcott, 1916 40 Genus Wenkchemnia Rasetti, 1951 40 Wenkchemnia ? housensis, new species 40 Wenkchemnia swasensis, new species 41 Family Zacanthoididae Swinnerton, 1915 43 Genus Parkaspis Rasetti, 1951 43 1. Research Associate, Invertebrate Paleontology Section, Natural History Museum of Los Angeles County. Contributions in Science, Number 446, pp. 1-137 Natural History Museum of Los Angeles County, 1994 Parkaspis drumensis, new species 43 Genus Zacanthoides Walcott, 1888 46 Zacanthoides divergensis Rasetti, 1951 (s.l.) 46 Zacanthoides divergensis divergensis Rasetti, 1951 48 Zacanthoides divergensis limbus, new subspecies 48 Zacanthoides divergensis, new subspecies A 48 Zacanthoides fronslicinus, new species 49 Zacanthoides walapai Resser, 1945 51 Order Ptychopariida Swinnerton, 1915 53 Suborder Ptychopariina Richter, 1933 53 Family Ehmaniellidae, new family 53 Subfamily Ehmaniellinae, new subfamily 53 Genus Ehmaniella Resser, 1937 53 Ehmaniella angustigena, new species 54 Ehmaniella fronsplanata, new species 55 Ehmaniella fronsplanata fronsplanata, new subspecies 58 Ehmaniella fronsplanata convexa, new subspecies 58 Ehmaniella fronsplanata concava, new subspecies 58 Ehmaniella waptaensis Rasetti, 1951 60 Genus Elrathiella Poulsen, 1927 61 Elrathiella decora Resser, 1945 61 Elrathiella domensa, new species 63 Elrathiella dugwayensa, new species 65 Elrathiella euthyopsis, new species 68 Elrathiella} intermedia, new species 69 Elrathiella} cf. £.? intermedia, new species 71 Elrathiella rectangularia, new species 71 Genus Proehmaniella, new genus 73 Proehmaniella basilica (Resser, 1945) 73 Proehmaniella eldoradensa, new species 75 Proehmaniella} granulosa, new species 76 Proehmaniella hebes} (Resser, 1945) 77 Genus Pseudomexicella, new genus 78 Pseudomexicella hestia, new species 78 Genus Trachycheilus Resser, 1945 80 Trachycheilus granulosus, new species 80 Trachycheilus whirlwindensis, new species 82 Trachycheilus whirlwindensis whirlwindensis, new subspecies 84 Trachycheilus whirlwindensis drumensis, new subspecies 84 Trachycheilus} sp. A 85 Trachycheilus} sp. B 86 Genus Tympanuella, new genus 87 Tympanuella transversa, new species 87 Genus Uncertain 90 Ehmaniellinae sp. A 90 Ehmaniellinae sp. B 90 Subfamily Altiocculinae, new subfamily 91 Genus Altiocculus, new genus 91 Altiocculus drumensis, new species 93 Altiocculus concavus, new species 94 Altiocculus cf. A. harrisi (Robison, 1971) 96 Genus Pseudoalokistocare, new genus 96 Pseudoalokistocare swasii, new species 97 Pseudoalokistocare cf. P. paranotatum (Rasetti, 1951) 98 Pseudoalokistocare ?, sp. unidet 98 Family Asaphiscidae Raymond, 1924 99 Genus Glyphaspis Poulsen, 1927 99 Glyphaspis concavus, new species 100 Glyphaspis cf. G. concavus, new species 101 Family Uncertain 102 Genus Bathyocos, new genus 102 2 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites Bathyocos housensis, new species 102 Genus Chanciaopsis, new genus 103 Chanciaopsis heteromorphos, new species 105 Genus Deltina, new genus 107 Deltina limb a, new species 107 Deltina angustigena, new species 108 Genus Elrathina Resser, 1937 109 Elratbina wheeler a, new species 109 Genus Spencella Rasetti, 1963 Ill Spencella ? buttsi (Resser, 1938a) Ill Spencella ? utahensa, new species 112 Genus Trymataspis Robison, 1964a 114 Trymataspis cf. T. depressa Robison, 1964a 114 Genus Yuknessaspis Rasetti, 1951 114 Yuknessaspis benningtonis, new species 114 Yuknessaspis benningtonis benningtonis, new subspecies 117 Yuknessaspis benningtonis convexus, new subspecies 117 Yuknessaspis benningtonis planus, new subspecies 117 Genus Uncertain 118 Ptychopariida sp. A 118 ACKNOWLEDGMENTS 119 LITERATURE CITED 119 APPENDIX: Locality Register 122 Drum Mountains 122 House Range 123 Dugway Range 129 Snake Range 131 Patterson Pass (Southern Schell Creek Range) 133 Northern Egan Range 134 Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites ■ 3 ABSTRACT. Trilobites from the Ehmaniella Biozone (Middle Cambrian) of the North American craton have received relatively little attention, especially the assemblages of the Great Basin. Described here are several of the corynexochid and ptychopariid trilobites of the Ehmaniella Biozone of the Great Basin. Several ptychopariid trilobites from the Ehmaniella Biozone are assigned to the new family Ehmaniellidae based on their close stratigraphic occurrence and their morphological similarity in possessing: (1) thin exoskeletons; (2) relatively elongate glabellae with bluntly rounded to rounded frontal lobes, bifurcated FI lateral glabellar furrows, and converging, relatively straight axial furrows; and (3) micropygous pygidia with poorly defined borders, no or very shallow border furrows, anterior and posterior pleural bands that are moderately convex in exsagittal profile, and pleural and interpleural furrows that join at the border. Ehmaniellidae is further subdivided into two subfamilies, Ehmaniellinae and Altiocculinae. Genera in the Ehmaniellinae include: Ehmania Resser; Ehmaniella Resser; Elrathia Walcott; Elrathiella Poulsen; Pa- rehmania Deiss; Proehmaniella, new genus; Pseudomexicella, new genus; Trachycheilus Resser; and T ympanuella, new genus. Genera in the Altiocculinae include: Altiocculus, new genus; Pseudoalokistocare, new genus; and Schopfaspis Palmer and Gatehouse. The stratigraphic ranges of ptychopariid trilobites are used to subdivide the Ehmaniella Biozone into four local subzones, which are (in ascending order): Proehmaniella, Elrathiella, Ehmaniella, and Altioc- culus subzones. These subzones, which are based on shallow-water taxa, are chronostratigraphically correlated with the deep-water faunules of the “ Bathyuriscus-Elrathina ” Biozone of the Stephens For- mation, British Columbia, Canada, and with other regions in North America. New genera described here are the ptychopariids Altiocculus, Bathyocos, Chanciaopsis, Deltina, Proehmaniella, Pseudoalokistocare, Pseudomexicella, and T ympanuella and the corynexochid genus Opsiosoryctocephalus. New species and subspecies include the ptychopariids Altiocculus concavus, Al- tiocculus drumensis, Bathyocos housensis, Chanciaopsis heteromorphos, Deltina angustigena, Deltina limba, Ehmaniella angustigena, Ehmaniella fronsplanata fronsplanata, Ehmaniella fronsplanata con- cava, Ehmaniella fronsplanata convexa, Elrathiella domensa, Elrathiella dugwayensa, Elrathiella euthy- opsis, Elrathiella intermedia, Elrathiella rectangularia, Elrathina wheelera, Glyphaspis concavus, Proeh- maniella eldoradensa, Proehmaniella ? granulosa, Pseudoalokistocare swasii, Pseudomexicella hestia, Spencella ? utahensa, Trachycheilus whirlwindensis drumensis, Trachycheilus whirlwindensis whirlwind- ensis, Trachycheilus granulosus, T ympanuella transversa, Yuknessaspis benningtonis benningtonis, Yuk- nessaspis benningtonis convexus, and Yuknessaspis benningtonis planus and the corynexochids Dorypyge swasii, Kootenia ? margoin distinct a, Opsiosoryctocephalus ophis, Parkaspis drumensis, Wenkchemnia ? housensis, Wenkchemnia swasensis, and Zacanthoides fronslicinus. A new subspecies of Zacanthoides divergensis Rasetti is named Z. divergensis limbus. INTRODUCTION Trilobites of the Ehmaniella Biozone are wide- spread in the Great Basin and have been extensively Figure 1. Index map showing location of measured sec- tions. used for the correlation of Middle Cambrian strata both among mountain ranges within this region (Robison, 1964b, 1976; Palmer, 1971a; Kepper, 1972, 1976; Hintze and Robison, 1975) and with other regions in the United States and Canada (Ro- bison, 1964b; Palmer, 1971b). Despite the bio- stratigraphic importance of this trilobite assem- blage, no taxonomic study of this fauna from the Great Basin has been published. Assemblages from the Ehmaniella Biozone are known from Arizona (Resser, 1945), Montana (Deiss, 1938), British Co- lumbia (Rasetti, 1951), Tennessee (Resser, 1938a; Sundberg, 1989a), Alabama (Schwimmer, 1989), and Greenland (Poulsen, 1927), but none of the species from these assemblages have been reported from the Great Basin. This paper describes the corynexo- chid and ptychopariid assemblages of the Ehma- niella Biozone from six stratigraphic sections in the central Great Basin, west-central Utah and east- central Nevada (Fig. 1). In addition, the Ehmaniella Biozone is subdivided into four local subzones, the Proehmaniella, Elrathiella, Ehmaniella, and Al- tiocculus subzones, and is correlated to other regions within North America. An important evolutionary aspect of the trilobite faunas of the Ehmaniella Biozone is their occur- rence at the bottom of the Marjumiid biomere. The lower boundary of the Ehmaniella Biozone is an extinction event that marks the end of the cory- nexochid-dominated Corynexochid biomere 4 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites Biozone Glossopleura Ehmaniella Subzone Proehmaniella Elrathiella Ehmaniella Altiocculus Fossil Horizons mill _LL ilium mi Proehmaniella eldoradensian. sp. Kootenia cf. K. havasuensis Resser Proehmaniella basilica (Resser) ' Zacanthoides walapai Resser ~ " Trachycheilus ? sp. A Proehmaniella hebe ? (Resser) '' Deltina angustigena n. sp. Proehmaniella ? granulosa n. sp. - Pseudomexicella hestia n. sp. - Wenkchemnia ? housensis n. sp. - - Spencella? buttsi (Resser) Kootenia dawsoni (Walcott) Elrathiella domensa n. sp. Elrathiella decora Resser Elrathiella euthyopsis n. sp. Trachycheilus granulosus n. sp. Ehmaniellinae sp. A - - Tympanuella transversa n. sp. “ “ Elrathiella ? intermedia n. sp. - Trachycheilus whirlwindensis n. sp. - ■ Elrathiella dugwayensa n. sp. Ehmaniella waptaensis Rasseti - Kootenia ? margoindistincta n. sp. - - - Ehmaniella fronsplanata n. sp. Ehmaniellinae sp. B - Pseudoalokistocare cf. P. paranotatum (Rasetti) Elrathiella rectangularia n. sp. Olenoides serratus (Rominger) Wenkchemnia swasensis n. sp. - Kootenia cf. K. dawsoni (Walcott) - - Ehmaniella angustigena n. sp Pseudoalokistocare swasii n. sp. Altiocculus drumensis n. sp. Dorypyge swasii n. sp. Deltina limba n. sp. - - - - Zacanthoides fronslicinus n. sp Tonkinella breviceps Kobayashi Kootenia quadriceps (Hall and Whitfield) Zacanthoides divergensis Rasetti - Ptychopariida sp. A - - - Glyphaspis concavus n. sp. Parkaspis drumensis n. sp. Tonkinella valida Tchernysheva Chanciaopsis heteromorphos n. sp. Kootenia aciculahs Resser _ - _ Spencella ? utahensa n. sp. Trachycheilus ? sp. B Trymataspis cf. T. depressa Robison - Yuknessaspis benningtonis n. sp. Altiocculus concavus n. sp. Bathyocos housensis n. sp. Altiocculus cf. A. harrisi (Robison) Glyphaspis cf. G. concavus n. sp Elrathina wheelers n. sp. Opsiosoryctocephalus ophis n. sp. - T- I. -V i \ .1- ■}. -t Drum Mountains Stratigraphic Chisholm Formation Dome Limestone Whirlwind Formation Swasey Limestone C «5,2 O 4- c/> « ■g E (0 i. x: o 0) ll —15536-15548 _1 5534,1 5535 imm “1 5526,1 5527 -15525 -15523 "15522 -15521 “15520 I .«•#§•#•••• • I mil Altiocullus Subzone Ehmaniella Subzone • • Elrathiella Subzone Figure 5. Species range chart for the Dugway Range section. (See Fig. 79 for lithologic symbols.) studied from the Great Basin, Ehmaniella is rec- ognized only from the upper half of the interval that is normally referred to as the Ehmaniella Bio- zone. Taxa previously referred to Ehmaniella in the lower half of the biozone (Resser, 1945; Oldroyd, 1973; Campbell, 1974; Hintze and Robison, 1975; Robison, 1976) are either Elrathiella Poulsen, 1927, or Proehmaniella. Ehmaniella waptaensis Rasetti, 1951, is the lowest-occurring species of the genus. This species occurs approximately 35 m above the Glossopleura Biozone in the deep-water Stephen Formation, British Columbia (Fritz, 1971), and ap- proximately 95 m above the Glossopleura Biozone in the shallow-water Whirlwind Formation, Utah. These stratigraphic positions indicate that Ehma- niella does not extend down to the top of the Glossopleura Biozone and that it may be limited to only the upper portion of the Ehmaniella Bio- zone. The base of the Ehmaniella Biozone has not been raised to the lowest occurrence of the genus, because further analysis of additional sections cov- ering a broader geographic area and spectrum of depositional environments is needed to confirm the limited stratigraphic occurrence of Ehmaniella pri- or to changing the biozonal boundary. Genera common at various levels in the Ehmani- ella Biozone of the study area are: Altiocculus, new genus; Dorypyge Dames, 1883; Ehmaniella; Elra- thia Walcott, 1924; Elrathiella; Kootenia Walcott, 1889; Glyphaspis Poulsen, 1927; Pagetia Walcott, 1916; Parkaspis Rasetti, 1951; Peronopsis Hawle and Corda, 1847; Proehmaniella; Ptychagnostus Jaekel, 1909; Spencella Rasetti, 1963; Tonkinella Mansuy, 1916; Trachycheilus Resser, 1945; Tym- panuella, new genus; Wenkchemnia Rasetti, 1951; and Zacanthoides Walcott, 1888 (Oldroyd, 1973; Randolph, 1973; White, 1973; Robison, 1982; Ko- paska-Merkel, 1983; this study). Based on the first occurrences of key genera, the Ehmaniella Biozone is divided into four local subzones, which are (in ascending order): Proehmaniella, Elrathiella, Eh- maniella, and Altiocculus subzones. Each subzone is named for its diagnostic genus and is discussed below. The composite stratigraphic ranges of the taxa described in this paper are shown in Figure 2. The taxa ranges within the six stratigraphic sections are shown in Figures 3-8. The Ehmaniella Biozone correlates to the Bathy- uriscus-Elrathina Biozone defined by Rasetti (1951). Robison (1976) stated that both Bathyuriscus Ra- setti, 1948, and Elrathina Resser, 1937, are not restricted to the biozone, and he suggested the abandonment of Rasetti’s biozonal name. In turn, Robison proposed the use of the Ehmaniella Bio- zone for intracratonic-basin deposits and the Or- yctocephalus Biozone for open-ocean deposits. The Oryctocephalus Biozone spans a large stratigraphic interval that is equal to most of the Middle Cam- brian including the Plagiura-Poliella, Albertella, Glossopleura, and Ehmaniella biozones (Robison, 1976). However, Glossopleura Poulsen, 1927, and Ehmaniella are also known from open-ocean de- posits (Resser, 1938a; Rasetti, 1951; Fritz, 1971; Palmer, 1971a; Campbell, 1974; Briggs and Collins, 8 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites 1988), indicating that the Glossopleura and Eh- maniella biozones can also be recognized in open- ocean deposits. The Ehmaniella Biozone and its subzones cor- relate with the Bathyuriscus-Elrathina Biozone and its faunules from the Stephen Formation (Fig. 9). Fritz (1971) recognized five faunules, which are (in ascending order): Kootenia sp. 1, Ogygopsis klotzi, Pagetia bootes, Ehmaniella burgessensis, and Ba- thyuriscus adaeus faunules. The correlation of the Proehmaniella and Elrathiella subzones with the Kootenia sp. 1 and Ogygopsis klotzi faunules is based on their stratigraphic position between the Glossopleura Biozone and the first occurrence of Ehmaniella. Other than their general assignment to the Ehmaniella Biozone, these two subzones and two faunules do not have common taxa that can be used for correlation. The correlation of the Eh- maniella and Altiocculus subzones with the Pagetia bootes, Ehmaniella burgessensis, and Bathyuriscus adaeus faunules is discussed below. The correlation of the lower three subzones of the Ehmaniella Biozone in the study area, the Proehmaniella, Elrathiella, and Ehmaniella sub- zones, with the Ehmaniella to lower Bolaspis-Gly- phaspis biozones of Schwimmer (1973, 1975) from Montana (Fig. 9) is based also on the stratigraphic position of the subzones and Schwimmer’s biozone between the Glossopleura Biozone and the first oc- currences of Altiocculus, Peronopsis intermedius (Tullberg, 1880), and Peronopsis gaspensis Rasetti, 1948. Schwimmer (1973, 1975) recognized the oc- currence of Ehmaniella specimens slightly above the Glossopleura Biozone in his Montana sections. However, these specimens resemble Elrathiella and not Ehmaniella. As a result, Schwimmer’s Ehmani- ella Biozone correlates to a portion of the Elra- thiella Subzone in the study area. Other than the genus Elrathiella, the three subzones in the study area and the faunas of the Ehmaniella to lower Bolaspis-Glyphaspis biozones in Montana have no taxa in common that are used for correlation. The correlation with the Altiocculus Subzone to the upper Bolaspis-Glyphaspis, Ehmania, and Pareh- mania Biozones is discussed below. PROEHMANIELLA SUBZONE This subzone is the lowermost subzone of the Eh- maniella Biozone. The lower boundary is defined by the first occurrence of Proehmaniella, which is well constrained with several fossil collections bracketing the boundary (Fig. 2) in four of the Great Basin sections. Common species in this subzone are Proehmaniella basilica (Resser, 1945); Zacan- thoides walapai Resser, 1945; Wenkchemnia ? hou- sensis, new species; and Kootenia dawsoni (Wal- cott, 1889). Only one species from the underlying Glossopleura Biozone ranges into this subzone; Glossopleura boccar (Walcott, 1916) occurs in the lower 10 cm of the Proehmaniella Subzone in the Drum Mountains (Sundberg, 1990). Figure 6. Species range chart for the northern Egan Range section. (See Fig. 79 for lithologic symbols.) ELRATHIELLA SUBZONE This subzone occurs above the Proehmaniella Sub- zone. The lower boundary is defined by the first occurrence of Elrathiella, which is poorly con- strained since only a few fossil collections are lo- cated near the boundary in the studied sections (Fig. 2). Common species in this subzone are Elrathiella decora Resser, 1945; Elrathiella dugwayensa, new species; Trachycheilus granulosus, new species; Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites II 9 Figure 7. Species range chart for the Snake Range section. (See Fig. 79 for lithologic symbols.) Tracbycheilus wbirlwindensis, new species; and Tympanuella transversa, new species. EHMANIELLA SUBZONE This subzone occurs above the Elrathiella Subzone. The lower boundary is defined by the first occur- rence of Ehmaniella, which is well constrained with several fossil collections bracketing the boundary (Fig. 2). Common species in this subzone are Eh- maniella waptaensis, Ehmaniella fronsplanata, new species, Wenkcbemnia swasensis, new species, and T ympanuella transversa . The Ehmaniella Subzone correlates to the upper Ogygopsis klotzi, Pagetia bootes, and Ehmaniella burgessensis faunules of Fritz (1971; Fig. 9). The stratigraphic position of Ehmaniella waptaensis in both regions provides the correlation of the sub- zone to the three faunules. It is not known how much of the E. burgessensis faunule correlates to either the Ehmaniella Subzone or the overlying Al- tiocculus Subzone because the boundary between the two subzones is poorly constrained and because of the low diversity of the £. burgessensis faunule (Rasetti, 1951; Fritz, 1971). In the Stephen Formation, Ehmaniella waptaen- sis occurs with Ptychagnostus praecurrens (Wes- tergard, 1936) ('= Ptychagnostus burgessensis [Ra- 10 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites = 15580, 15581 Altiocullus Subzone Ehmaniella Subzone Elrathiella Subzone Figure 8. Species range chart for the Patterson Pass section. (See Fig. 79 for lithologic symbols.) setti, 1951] of Fritz [1971]; Robison [1982, 1984]). This indicates that the Ehmaniella Subzone cor- relates to part of the Ptychagnostus praecurrens Biozone (Fig. 9). ALTIOCCULUS SUBZONE This subzone occurs above the Ehmaniella Sub- zone and is the uppermost subzone in the Ehmani- ella Biozone. The lower boundary is defined by the first occurrence of Altiocculus, which is poorly con- strained because fossil collections containing as- semblages diagnostic of either of the two subzones are separated by a relatively thick, unfossiliferous interval of the Swasey Limestone (Fig. 2). The upper boundary of the Altiocculus Subzone is defined by the first occurrence of Bolaspidella Resser, 1937, which marks the lower boundary of the Bolaspi- della Biozone. In the sections studied, the common species in this subzone are Altiocculus drumensis. This Paper Agnostoid Robison, 1982, 1984 Schwimmer, 1973, 1975 Fritz, 1971 Young and Ludvigsen, 1989 Bolaspidella P. atavus Bolaspidella Zone 3 V) P. gibbus Parehmania ~$atHyurlscus~ Zone 2 | Ehmania adaeus Faunule Zacanthoides gilbeti ““| Fauna J” “ Bolaspis- Giyphaspis Ehmaniella burgessensis _ ^Faunule Ehmaniella iella P. praecurrens Pagetia g bootes £ Faunule iu " Ogygopsis £ klotzi J. Faunule tg CQ i £ Ehmaniella Elrathiella Kootenia sp. 1 Faunule 1 C 1 Gbssopleura Gbssopleura Gbssopleura Figure 9. Correlation chart of polymeroid biostrati- graphic units from the study area to agnostid and other polymeroid sequences. new species, Altiocculus concavus, new species, Dorypyge swasii, new species, Glyphaspis conca- vus, new species, Parkaspis drumensis, new spe- cies, Peronopsis cf. P. fallax (Linnarsson, 1869), and Tonkinella breviceps Kobayashi, 1934. In the Wheeler Shale, the trilobite faunas are dominated by the agnostoids Peronopsis amplaxis Robison, 1982, Ptychagnostus gibbus (Linnarsson, 1869), and Ptychagnostus intermedius (see Robison, 1982; Rowell et ah, 1982). As defined here, the Altioc- culus Subzone also incorporates the upper part of the Ptychagnostus praecurrens agnostoid biozone and the majority of the P. gibbus agnostoid biozone (Fig. 9). The Glyphaspis fauna of Hintze and Robison (1975) and Rees (1986) is equivalent to the faunas of the Altiocculus Subzone in the uppermost por- tion of the Swasey Limestone. Robison (1964b) correlated the Parkaspis endecamera faunule of Rasetti (1951) from British Columbia to the faunas in the uppermost portion of the Swasey Limestone. The strong similarity between the two faunas is shown by the occurrence of similar or identical species of the genera Altiocculus; Glyphaspis; Par- kaspis; Peronopsis; Tonkinella; Yuknessaspis Ra- setti, 1951; and Zacanthoides in both assemblages. Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobitesl 11 a* c rt u 00 ON c o C/5 15 o - a C vs rt _r .a »o t3 r- C On 3 ro O t^ Q ON 3 «- •r w '*h ^ O ON c/s ' — • 5 N .2 -s g>£ ^ «N ii ns 0/ T-( -c S£ 1 & * £ 8 « £ ■f? 3 Q\, C/5 T-H u, X c* | £ > o i/s ,H C o ns • - > £ ~o o l-l-N 3 w M-n C/5 O 0/ -C t: IN « (U -C -o u c tO '3 4-> 3 O eg Ui o X S-H u. ns O 4-N u ns Q d o _o t-h 10 mm) of Chanciaopsis have posterior area of fixigenae 110- 120% glabella length, whereas Chancia has pos- terior area of fixigenae 80-100% glabella length. Ratios for Chancia are based on the illustrated specimens of C. ebdome, C. agusta, C. evax, and C. vestita (Resser, 1939a) in Resser (1939a, b), of which the first three species are compressed in shales, which would increase the lengths of the fixigenae and posterior area of fixigenae. Chanciaopsis heteromorphos has a broad range of morphologies; one extreme morphology (Fig. 67.4) suggests that Chanciaopsis may be closely related to Altiocculus. This specimen has very shal- low lateral glabellar furrows; relatively short, mod- erately strongly tapered, and broad glabella, convex axial furrows; and very faint secondary basal lobes. These features are very similar to larger specimens of Altiocculus drumensis. Furthermore, if C.? pal- liseri is included in the genus, then Chanciaopsis also has a pygidium similar to Altiocculus. These similarities suggest that Chanciaopsis may belong in Altiocculinae. Chanciaopsis occurs in the Altiocculus Subzone of the Ehmaniella Biozone. ETYMOLOGY. Greek, -opis = like, for the like- ness to Chancia. Chanciaopsis heteromorphos , new species Figure 67 DIAGNOSIS. Chanciaopsis with two cranidial types. Specimens without prominent medial swell- ing with convergent, concave facial suture anterior of palpebral lobes; flat, level anterior border, very slight median swelling in anterior border furrow; moderate to large palpebral lobes, moderately pos- terior directed ocular ridges; anterior area of fixi- gena and preglabellar field slightly downsloping to very strongly downsloping; and moderately narrow posterior area of fixigenae (30-40% glabellar length). Specimens with prominent medial swelling with di- vergent convex facial sutures anterior of palpebral lobes; flat, level anterior border relatively wide; moderate to large palpebral lobes, moderately pos- terior directed ocular ridges; anterior area of fixi- gena and preglabellar field moderately downsloping 15517, partially exfoliated, x4.1. 5, Stereo pair of small paratype cranidium (LACMIP 11950), LACMIP locality 15541, exfoliated, x?.8. 6, Stereo pair of larger paratype cranidium (LACMIP 11951), LACMIP locality 15543, partially exfoliated, x2.8. Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites ■ 105 to very strongly downsloping; and moderately nar- row posterior area of fixigenae. DESCRIPTION. Cranidium moderate size, length 4.2-12.1 mm; subpentagonal, length 50-55% width; low to strong sagittal and moderate transverse con- vexity, height 20-30% width; anterior margin slightly to moderately, evenly curved, anterior margin with 55-70% cranidial width; posterior margin, exclud- ing occipital ring, slightly backswept. Facial sutures slightly convergent and concave to slightly diver- gent and convex from ends of palpebral lobes to anterior border furrow; strongly convergent across border; strongly divergent posterior of palpebral lobes. Glabella moderately elongate, length 65-70% and width 50-60% cranidial length; glabella width 25-35% cranidial width; strongly to very strongly tapered, width at anterior end 60-70% glabellar width; moderate convexity (sag. and trans.); frontal lobe moderately rounded, 3 lateral glabellar lobes, very poorly developed secondary basal lobe. Axial furrows moderately deep to deep, deeper posteriorly, slightly convex in larger specimens, very slightly concave in smaller specimens; preglabellar furrow shallow to moderately deep, narrow, moderately curved; lateral glabellar furrows very shallow to deep, FI furrow bifurcated, directed posteriorly, F2 furrow directed slightly posteriorly, F3 directed slightly anteriorly. Occipital ring sagittal length 20~ 25% glabellar length, slightly elevated above gla- bella, low convexity; small occipital node; furrow curved anteriorly, moderately deep laterally, shal- low medially; posterior margin slightly curved to straight. Frontal area length 30-35% cranidial length, subequally to unequally divided. Preglabellar field may possess median swelling, circular or subtrian- gular, moderately convex, area around swelling de- pressed; slightly to strongly downsloping, length 50-65% frontal area length. Anterior border flat, level, moderately tapered laterally, evenly curved, sagittal length 35-50% frontal area length. Anterior border furrow evenly curved, moderately narrow, shallow, shallower than axial furrows, very low transversely elongate medial swelling. Fixigena width, exclusive of palpebral lobe, 60-75% gla- bellar width, moderate convexity, upsloping, an- terior area moderately to strongly downsloping. Palpebral lobe narrow to moderate width, width 25-40% lobe length; moderate length to long, length 30-45% glabellar length; anterior margin located adjacent to 20-30% glabellar length from anterior margin of frontal lobe; upturned, furrow shallow. Ocular ridge weak to moderate strength, moder- ately thin, directed moderately posterolaterally from glabella at 65-70° to axial trend. Posterior area of fixigena width 30-40% glabellar length; length 90- 120% glabella length; downsloping and backswept laterally, roundly terminated. Posterior border high, strongly convex, wider laterally; border furrow nar- row, deep, curved posteriorly, uniform width. Librigena large, length 12.3 mm; wide, width ap- proximately 55% length without spine; lateral mar- gin moderately curved; height approximately 45% width. Genal field gently convex, width approxi- mately 80% librigenal width. Border width ap- proximately 20% librigenal width; gently convex; lateral border furrow shallow, uniform depth; pos- terior border furrow absent. Genal spine unknown. Hypostoma, thorax, and pygidium unknown. Exoskeleton thin; fine granules on external sur- face of cranidium, genal caeca on anterior area of fixigena, preglabellar field (not on median swelling), and librigenae on external and internal surfaces. REMARKS. Chanciaopsis heteromorphos spans a morphological range typical of two genera. Spec- imens are grouped into two morphological types, one with a prominent median swelling and one without a prominent median swelling. Flowever, the strong or very weak development of the median swelling appears to be the only consistent difference between the two groups. There is no other break between the two morphologies and the two types represent extremes in a continuum. Variable fea- tures include the sagittal width of the posterior area of fixigena, depth of lateral glabellar furrows and axial furrows, tapering of glabella, width of anterior margin, and degree of downsloping of the anterior area of the fixigena and preglabellar field. Typically, the specimens with a median swelling have a steeper downsloping of the anterior area of fixigena and preglabellar lobe, wider anterior margin, and di- vergent anterior facial sutures, but some specimens without a median swelling also show these char- acters. Both morphologies of the species commonly occur together at the same locality. The specimens with a median swelling are unlike other Chanciaopsis species. Specimens without a median swelling are similar to C. odarayensis but differ in having larger palpebral lobes, concave an- terior border, and moderately posterolaterally di- rected ocular ridges. ETYMOLOGY. Latinized, named after the Swa- sey Limestone. HOLOTYPE. Cranidium (LACMIP 11953) from the Trailer Limestone, Dugway Range, Utah (LAC- MIP locality 15544). PARATYPES. LACMIP 11949 from LACMIP locality 15517; LACMIP 11950 from LACMIP lo- cality 15541; LACMIP 11951, 11952 from LAC- MIP locality 15543; LACMIP 11954 from LAC- MIP locality 15547; LACMIP 11955-11957 from LACMIP locality 15548. OTHER MATERIAL. LACMIP locality 15496, 3 cranidia; LACMIP locality 15500, 13 cranidia; LACMIP locality 15517, 8 cranidia; LACMIP locality 15541, 1 cra- nidium; LACMIP locality 15542, 1 cranidium; LACMIP locality 15543, 15 cranidia; LACMIP locality 15544, 2 cranidia; LACMIP locality 15545, 13 cranidia; LACMIP locality 15547, 2 cranidia; LACMIP locality 15548, 12 cranidia; LACMIP locality 15565, 1 cranidium; LACMIP locality 15568, 5 cranidia; LACMIP locality 15569, 6 cra- nidia; LACMIP locality 15570, 6 cranidia; LACMIP lo- cality 15571, 1? cranidium; LACMIP locality 15572, 1 cranidium. 106 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites Figure 68. Deltina limba, new genus and species, LACMIP locality 15531. 1, Stereo pair of holotype cranidium (LACMIP 11965), slightly exfoliated, xl3.0. 2, Paratype cranidium (LACMIP 11964), exfoliated, xl3.4. OCCURRENCE. Altiocculus Subzone, Ehmani- ella Biozone. Swasey Limestone, Drum Mountains and House Range, Utah. Trailer Limestone, Dug- way Range, Utah. Pole Canyon Limestone, Mem- ber E, Snake Range, Nevada. Genus Deltina , new genus TYPE SPECIES. Deltina limba, new species. DIAGNOSIS. Small Ptychopariida with subrec- tangular cranidium with a relatively large glabella elevated well above fixigena; moderately convex transversely; parallel to slightly convergent, mod- erate depth axial furrows; shallow to moderately shallow lateral glabellar furrows; rounded frontal lobe; downsloping, moderately narrow fixigenae; narrow or absent preglabellar field; convex, mod- erately to moderately strong curved, transversely wide, sagittally narrow, slightly tapering anterior border; moderate to short length palpebral lobes; moderate length posterior area of fixigenae. REMARKS. Deltina is most similar to Spencella in its elevated, relatively large, subquadrate glabella. Deltina differs from Spencella in having a greater transverse width; a uniform width anterior border; rounded frontal lobe; and longer posterior area of fixigenae. Deltina differs from other Middle Cam- brian ptychopariids in its relatively large, elevated glabella and in the character of the anterior border. The pygidium, thoracic segments, hypostoma, and librigenae are not known for Deltina. Deltina occurs in the Ehmaniella Biozone. ETYMOLOGY. Latinized, named for Delta, Utah, which is located near the type localities for both known species. Deltina limba , new species Figure 68 DIAGNOSIS. Deltina with cranidium with rel- atively wide preglabellar field (30-50% frontal area), moderate length palpebral lobes (30% glabellar length), anterior border curved moderately strong, granular ornamentation. DESCRIPTION. Cranidium small, length 2.8- 3.4 mm; rectangular, length 65-70% width; mod- erate convexity (sag. and trans.), height 30-35% width; anterior margin curved moderately strong, anterior margin width 65-80% cranidial width; pos- terior margin, excluding occipital ring, slightly backswept. Facial sutures parallel to slightly diver- gent from ends of palpebral lobes to anterior bor- der; moderately convergent across anterior border; strongly divergent posterior of palpebral lobes. Gla- bella elongate, length 75-80% and width 55-65% cranidial length; glabella width approximately 40% cranidial width; strongly to moderately tapered, width at anterior end 70-80% glabellar width; mod- erate convexity (sag. and trans.); frontal lobe mod- erately rounded, 3 lateral glabellar lobes. Axial fur- rows moderate depth, deeper posteriorly, slightly concave; preglabellar furrow moderate depth, moderately curved; lateral glabellar furrows shal- low, FI furrow bifurcated, directed posteriorly, F2 Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites ■ 107 furrow directed slightly posteriorly, F3 furrow di- rected slightly posteriorly. Occipital ring sagittal length 20-25% glabellar length, not elevated above glabella, moderately convex; weak occipital node; furrow straight, deeper laterally, shallowing medi- ally, moderately deep; posterior margin moderately curved, flattened medially. Frontal area length 20- 25% cranidial length, equally to subequally divided. Preglabellar field concave, downsloping, length 30- 50% frontal area length. Anterior border strongly convex, level, slightly tapering laterally, evenly curved, sagittal length 50-70% frontal area length. Anterior border furrow evenly curved, broad, mod- erate depth, shallower or as deep as axial furrows. Fixigena width, exclusive of palpebral lobe, 25- 45% glabellar width, low convexity, slightly down- sloping; anterior area of fixigena moderately down- sloping. Palpebral lobe wide, width approximately 40% lobe length; moderate length, length approx- imately 30% glabellar length; anterior margin lo- cated adjacent to 25-30% glabellar length from anterior margin of frontal lobe; furrow shallow depth. Ocular ridge moderately weak, directed moderately posterolaterally from glabella at 65-70° to axial trend. Posterior area of fixigena exsagittal width 30-40% glabellar length; length 60-65% gla- bella length; bluntly terminated. Posterior border strongly convex, slightly wider distally; border fur- row narrow, deep, shallowing laterally, curved slightly posteriorly. Librigena, hypostoma, thorax, and pygidium un- known. Exoskeleton thin; fine granules with scattered coarse granules on external surface and internal mold. REMARKS. Deltina limba differs from D. an - gustigena, new species, in having granular orna- mentation, a sagittally wider preglabellar field, and strongly curved and transversely wide anterior bor- der. ETYMOLOGY. Latin, limbus, named for the strongly convex anterior border. HOLOTYPE. Cranidium (LACMIP 11965) from the Trailer Limestone, Dugway Range, Utah (LAC- MIP locality 15531). PARATYPES. LACMIP 11962, 11963 from LACMIP locality 15526; LACMIP 11964 from LACMIP locality 15531. OTHER MATERIAL. LACMIP locality 15526, 1 cra- nidium; LACMIP locality 15529, 2 cranidia; LACMIP locality 15530, fragments; LACMIP locality 15491, 1 cra- nidium. OCCURRENCE. Altiocculus Subzone, Ehmani- ella Biozone. Trailer Limestone, Dugway Range, Utah. Swasey Limestone, Drum Mountains, Utah. Deltina angustigena, new species Figure 69 DIAGNOSIS. Deltina with cranidium with very narrow to narrow preglabellar field, moderate length to long palpebral lobes (30-45% glabellar length), moderately curved anterior border, smooth exo- skeleton. DESCRIPTION. Cranidium small, length 2.5- 4.7 mm; subrectangular, length 65-70% width; moderate convexity (sag. and trans.), height ap- proximately 40% width; anterior margin moder- ately curved, anterior margin width approximately 65% cranidial width; posterior margin, excluding occipital ring, slightly backswept. Facial sutures parallel to slightly divergent from ends of palpebral lobes to anterior border; moderately convergent across anterior border; strongly divergent posterior of palpebral lobes. Glabella elongate, length 80- 85% and width 50-65% cranidial length; glabella with 35-45% cranidial width; strongly to slightly tapered, width at anterior end 70-90% glabellar width; moderate convexity (sag. and trans.); frontal lobe moderately rounded, 3 lateral glabellar lobes. Axial furrows moderate depth, deeper posteriorly, slightly concave; preglabellar furrow shallow, mod- erately curved; lateral glabellar furrows shallow, FI furrow bifurcated, directed slightly posterior, F2 and F3 furrows directed slightly anterior. Occipital ring sagittal length 20-25% glabellar length, not elevated above glabella, slightly convex; weak oc- cipital node; furrow straight, deeper laterally, shal- lowing medially, moderate depth; posterior margin moderately curved, flatten medially. Frontal area length approximately 15% cranidial length, sub- equally divided. Preglabellar field very narrow to narrow, slightly concave, downsloping, length 5- 35% frontal area length. Anterior border strongly convex, level, slightly tapering laterally, evenly curved, sagittal length 65-95% frontal area length. Anterior border furrow evenly curved, broad, mod- erate depth, shallower or as deep as axial furrows. Fixigena width, exclusive of palpebral lobe, 30- 50% glabellar width, low convexity, slightly down- sloping; anterior area of fixigena moderately down- sloping. Palpebral lobe narrow, width 25-30% lobe length; moderate length to long, length 30-45% glabellar length; anterior margin located adjacent to 30-35% glabellar length from anterior margin of frontal lobe; furrow moderate depth. Ocular ridge moderate to moderately weak, directed very strongly posterolaterally from glabella at 50-55° to axial trend. Posterior area of fixigena exsagittal width approximately 25% glabellar length; length 45-50% glabella length; bluntly terminated. Posterior bor- der strongly convex, slightly wider distally; border furrow narrow, deep, shallowing laterally, curved slightly posteriorly. Librigena, hypostoma, thorax, and pygidium un- known. Exoskeleton thin; smooth on external and inter- nal surfaces. REMARKS. Deltina angustigena differs from D. limba in having a smooth exoskeleton, a sagittally narrower preglabellar field, narrower fixigena, and less strongly curved and narrower anterior border. Deltina angustigena differs from Spencella?. buttsi 108 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites Figure 69. Deltina angustigena, new genus and species. RC locality 374. 1, Stereo pair of holotype cranidium (KUMIP 204728), mostly exfoliated, x7.5. 2, Partial paratype cranidium (KUMIP 204729), exfoliated, xl3.1. 3, Stereo pair of paratype cranidium (KUMIP 204727), exfoliated, x9.0. (Resser, 1938b), which with it occurs, in having narrower fixigenae, rounded frontal lobe, and a non-swollen, non-tapered anterior border. ETYMOLOGY. Latin, angustus = narrow, gena = cheek, for its relatively narrow fixigenae. HOLOTYPE. Cranidium (KUMIP 204728) from the Dome Limestone, House Range, Utah (RC lo- cality 374). PARATYPES. KUMIP 204726 from RC locality 211; KUMIP 204727, 204729 from RC locality 374. OTHER MATERIAL. RC locality 211, 13 cranidia; RC locality 374, 17 cranidia; LACMIP locality 15462, 4 cranidia; LACMIP locality 15503, 8 cranidia. OCCURRENCE. Proehmaniella Subzone, Eh~ maniella Biozone. Dome Limestone, House Range and Drum Mountain, Utah. Genus Elrathina Resser, 1937 TYPE SPECIES. Conocephalites cordilleras Rominger, 1887. DIAGNOSIS. Small, micropygous Ptychopariida with subrectangular cranidium with convergent fa- cial sutures anterior of the palpebral lobes, glabella elevated above fixigena; parallel to slightly conver- gent, moderate depth axial furrows; shallow to ef- faced lateral glabellar furrows; strongly rounded frontal lobe; slightly downsloping, moderately wide fixigenae; narrow preglabellar field; convex, slightly to moderately curved, sagittally narrow, tapering anterior border; short palpebral lobes anterior of glabellar midlength. Free checks lack significant de- velopment of genal spines. Triangular pygidium. REMARKS. The diagnosis is a combination from Deiss (1939) and Palmer (in Palmer and Halley, 1979). Elrathina wheeler a, new species Figure 70 Elrathina, n. sp. Sundberg, 1991:388, fig. 2.6. DIAGNOSIS. Elrathina with cranidium with transversely wide anterior border, transversely wide fixigenae (70-80% glabellar width), moderate size recurved occipital spine. DESCRIPTION. Cranidium small to moderate size, length 3. 1-6.2 mm; subtrapezoidal, length 60- 70% width; moderately low convexity (sag. and trans.), height 25-40% width; anterior margin mod- erately curved, anterior margin width 60-70% cra- nidial width; posterior margin, excluding occipital ring, straight. Facial sutures moderately convergent from ends of palpebral lobes to middle of anterior border; strongly convergent across anterior half of border; moderately divergent posterior of palpebral lobes. Glabella moderately elongate, length 70-75% and width 45-55% cranidial length; glabellar width 30-35% cranidial width; slightly tapered, width at anterior end 85-90% glabellar width; moderate to moderately low convexity (sag. and trans.); frontal lobe bluntly rounded, 3 lateral glabellar lobes. Ax- ial furrows moderate depth, slightly concave; pre- glabellar furrow moderate depth, slightly curved; lateral glabellar furrows very shallow, FI furrow bifurcated, directed posteriorly, F2 and F3 furrows Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobitesl 109 Figure 70. Elrathina wheeler a, new species. 1, Stereo pair of holotype cranidium (LACMIP 11968), LACMIP locality 15519, partially exfoliated, x 6.8. 2, Stereo pair of paratype cranidium (LACMIP 11966), LACMIP locality 15518, x 7.9. 3, Larger paratype cranidium (LACMIP 11970) showing occipital spine, LACMIP locality 15519, partially exfoliated, x5.3. 4, Larger paratype free cheek (LACMIP 11972), LACMIP locality 15519, exfoliated, x4.6. 5, Smaller paratype free cheek (LACMIP 11973), LACMIP locality 15519, exfoliated, xll.6. directed laterally. Occipital ring sagittal length 20- 25% glabellar length, slightly convex; moderate length, recurved occipital spine; furrow straight, deep pits laterally, shallow medial; posterior margin slightly curved to flattened. Frontal area length 25- 30% cranidial length, subequally divided. Pregla- bellar field flat to slightly concave, slightly down- sloping, length 60-70% frontal area length. Ante- rior border strongly convex, upsloping, tapering laterally, evenly curved, sagittal length 30-40% frontal area length. Anterior border furrow evenly curved, moderate depth, shallowing medially and laterally, shallower than axial furrows. Fixigena width, exclusive of palpebral lobe, 70-80% gla- bellar width, low convexity, very slightly down- sloping. Palpebral lobe narrow to moderate width, width 25-35% lobe length; short, length 20-25% glabellar length; anterior margin located adjacent to 10-20% glabellar length from anterior margin of frontal lobe; furrow moderate depth. Ocular ridge moderately weak, larger specimens have dou- ble ridges, directed slightly posterolaterally from glabella at 80-85° to axial trend. Posterior area of fixigena exsagittal width 50-75% glabellar length; length 70-75% glabella length; sharply terminated. Posterior border moderately convex, distally wider; border furrow narrow, distally wider, deep to mod- erate depth, straight. Librigena small to moderate size, length 4.6 mm; narrow, width approximately 35% length without spine; lateral margin gently curved; moderate con- vexity, height approximately 40% width. Genal field level to moderately convex, width approximately 55% librigenal width. Border width approximately 35% librigenal width; slightly convex to flat; level to downsloping; lateral border furrow very shallow; posterior border furrow absent to very short, very shallow. Genal spine very short, approximately 5% librigenal length. Hypostoma, thorax, and pygidium unknown. 110 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites Figure 71. Spencella ? buttsi (Resser). 1, Stereo pair of cranidium (KUMIP hypotype 204757), locality RC 211, partially exfoliated, xl4.4. 2, Stereo pair of partial cranidium (KUMIP hypotype 204756), RC locality 211, exfoliated, xl2.7. Exoskeleton very thin; genal caeca on anterior area of fixigena and preglabellar field of cranidia and genal area of larger librigenae, some glabella with fine granules, otherwise smooth on external and internal surfaces. REMARKS. Elrathina wheelera is most similar to E. erecta Deiss, 1939, in its parallel-sided glabella but differs in having an occipital spine, wider fixi- genae, and a transversely wider, sightly curved an- terior border. Elrathina wheelera differs from other species of Elrathina in its wider anterior margin and fixigena. Elrathina wheelera occurs in the uppermost 0.5 m of the Swasey Limestone and overlying 20 cm of black, agnostoid-rich mudstones, which are here considered the Wheeler Shale. ETYMOLOGY. Latinized, named after the Wheeler Shale. HOLOTYPE. Cranidium (LACMIP 11968) from the Wheeler Shale, House Range, Utah (LACMIP locality 15519). PARATYPES. LACMIP 11966, 11967 from LACMIP locality 15518; LACMIP 11959, 11969- 11971, 11973 from LACMIP locality 15519. OTHER MATERIAL. LACMIP locality 15517, 1 cra- nidium; LACMIP locality 15518, 11 cranidia; LACMIP locality 15519, 27 cranidia; LACMIP locality 15563, 3 cranidia; LACMIP locality 15572, 3 cranidia. OCCURRENCE. Altiocculus Subzone, Ehmani- ella Biozone and lowermost Ptychagnostus gibbus Biozone. Swasey Limestone and Wheeler Shale, House Range, Utah. Member E, Pole Canyon Lime- stone, and Lincoln Peak Shale, Snake Range, Ne- vada. Genus Spencella Rasetti, 1963 TYPE SPECIES. Spencella montanensis Rasetti, 1963. DIAGNOSIS. Small, micropygous Ptychopariida with subrectangular cranidium with glabella ele- vated well above fixigena; slightly convergent, mod- erate depth axial furrows; shallow to effaced lateral glabellar furrows; bluntly rounded from lateral lobe; downsloping, moderately narrow to fixigenae; nar- row or absent preglabellar field; convex, moder- ately curved, sagittally narrow, strongly tapering, medially swollen anterior border; short length pal- pebral lobes; short (trans.) posterior area of fixi- genae. Librigenae narrow, rounded at genal angle. Pygidium small, alate, elevated axis, 1-2 shallow pleural furrow pairs, poorly defined border. REMARKS. The diagnosis is based on the de- scription of Rasetti (1963). The species reported here are questionably assigned to Spencella because there are no associated librigenae and pygidia, both of which are necessary to separate Spencella from Spencia Resser, 1939a (Rasetti, 1963:591). Spencella ? buttsi (Resser, 1938a) Ligure 71 Solenopleurella buttsi Resser, 1938a:99, pi. 6, figs. 24, 25; Shimer and Shrock, 1944:pl. 255, fig. 22. Spencella buttsi (Resser), Rasetti, 1963:591. Contributions in Science, Number 446 Sundberg: Middle Cambrian TrilobitesB 111 DIAGNOSIS. Spencella ? with cranidium with bluntly rounded glabella; convex axial furrows; moderately dorsally arched anterior border, 65- 70% of frontal area, medial swelling absent or poor- ly developed; moderately high glabellar convexity (sag. and trans.); and exoskeleton smooth. DESCRIPTION. Cranidium small, length 2.3- 3.1 mm; rectangular, length 60-80% width; mod- erate convexity (sag. and trans.), height 30-40% width; anterior margin moderately curved, anterior margin width 50-80% cranidial width; posterior margin, excluding occipital ring, straight. Facial su- tures slightly convergent from ends of palpebral lobes to middle of anterior border; strongly con- vergent across anterior half of anterior border; moderately divergent posterior of palpebral lobes. Glabella elongate, length 75-80% and width 60- 75% cranidial length; glabella width 40-50% cra- nidial width; moderately to slightly tapered, width at anterior end 75-90% glabellar width; high con- vexity (sag. and trans.); frontal lobe flattened to bluntly rounded, 3 lateral glabellar lobes. Axial fur- rows moderate depth, slightly concave; preglabellar furrow moderate depth, straight to slightly curved; lateral glabellar furrows shallow to moderate depth, FI furrow directed posteriorly, F2 and F3 furrows directed slightly posteriorly. Occipital ring sagittal length 20-25% glabellar length, slightly lower than glabella, slightly convex; occipital node absent(?); furrow straight, deep pits laterally, shallowing me- dial, moderate depth; posterior margin curved. Frontal area length 20-25% cranidial length, sub- equally divided. Preglabellar field flat to slightly concave, slightly downsloping, length 25-40% frontal area length. Anterior border strongly con- vex, slightly upsloping, slightly to not swollen me- dially, tapering laterally, evenly curved, sagittal length 60-75% frontal area length. Anterior border furrow evenly curved, moderate depth shallowing medially, shallower than axial furrows. Fixigena width, exclusive of palpebral lobe, 40-50% gla- bellar width, low convexity, very slightly down- sloping. Palpebral lobe wide, width 40-45% lobe length; moderate length, length 30-40% glabellar length; anterior margin located adjacent to 15-25% glabellar length from anterior margin of frontal lobe; furrow moderate depth. Ocular ridge mod- erate to moderately weak, directed moderately pos- terolaterally from glabella at 70-75° to axial trend. Posterior area of fixigena exsagittal width 30-35% glabellar length; length 50-65% glabella length; bluntly terminated. Posterior border strongly to moderately convex, distally wider; border furrow narrow to moderately wide, deep to shallow, straight. Librigena, hypostoma, thorax, and pygidium un- known. Exoskeleton thin; smooth on external and inter- nal surfaces. REMARKS. Rasetti (1963) assigned Solenopleu- rella buttsi to Spencella, although neither the li- brigena nor the pygidium that are needed to place the species in the genus are known. For this reason, this species is questionably assigned to Spencella here. Spencella ? buttsi differs from other species of Spencella illustrated by Rasetti (1963) and Palmer (1968) in its narrow anterior border, absent or very slight medial inbending of anterior border furrow, and/or bluntly rounded glabella. Spencella ? buttsi from the Great Basin does not differ significantly from the Appalachian specimens figured by Resser (1938a). Spencella min or i (Resser, 1938a) may be conspecific with 5.? buttsi , but the illustrations of Resser are inadequate for the comparison of the two species. HYPOTYPES. KUMIP 204756, 204757 from RG locality 211. OTHER MATERIAL. RC locality 211, 12 cranidia; RC locality 374, 20 cranidia; LACMIP locality 15462, 4 cranidia; LACMIP locality 15503, 8 cranidia. OCCURRENCE. Proehmaniella Subzone, Eh - maniella Biozone. Dome Limestone, House Range and Drum Mountains, Utah. Spencella ? utahema, new species Figure 72 DIAGNOSIS. Spencella ? with cranidium with bluntly rounded to rounded frontal lobe; straight to convex axial furrows; moderately to strongly, dorsally arched anterior border, 60-95% of frontal area, medial swelling strong; moderately high gla- bellar convexity (sag. and trans.); and fine granular ornamentation. DESCRIPTION. Cranidium small, length 2.8- 3.5 mm; subtrapezoidal, length 65-95% width; moderate convexity (sag. and trans.), height 30- 45% width; anterior margin moderately curved, an- terior margin width 55-80% cranidial width; pos- terior margin, excluding occipital ring, straight. Fa- cial sutures slightly convergent to parallel from ends of palpebral lobes to middle of anterior border; strongly convergent across anterior half of anterior border; moderately divergent posterior of palpebral lobes. Glabella elongate, length 75-85% and width 55-65% cranidial length; glabella width 35-50% cranidial width; strongly tapered, width at anterior end 70-75% glabellar width; high convexity (sag. and trans.); frontal lobe flattened to bluntly round- ed, 3 lateral glabellar lobes. Axial furrows moderate depth to deep, slightly convex to straight; pregla- bellar furrow moderate depth, slightly curved; lat- eral glabellar furrows shallow to moderate depth, FI furrow bifurcated, directed posteriorly, F2 and F3 furrows directed slightly posteriorly to laterally. Occipital ring sagittal length 20-25% glabellar length, slightly lower than glabella, strongly con- vex; occipital node to short spine; furrow straight, deeper laterally, moderate depth; posterior margin moderately curved. Frontal area length 15-25% cranidial length, unequally divided. Preglabellar field 112 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites Figure 72. Spencella ? utahensa, new species. 1, 2, Stereo pair and anterior view of holotype cranidium (LACMIP 11975), LACMIP locality 15536, exfoliated, x 11.6. 3,4, Paratype cranidium (LACMIP 11978), LACMIP locality 15580, exfoliated, x!2.8. slightly convex, downsloping, length 5-40% frontal area length. Anterior border strongly convex, level, swollen medially, strongly tapering laterally, evenly curved, moderately to strongly arched dorsally, sag- ittal length 60-95% frontal area length. Anterior border furrow evenly curved, moderate depth to deep, uniform depth to shallowing medially, as deep as axial furrows. Fixigena width, exclusive of pal- pebral lobe, 40-50% glabellar width, weak to strong convexity, slightly downsloping. Palpebral lobe narrow to moderately wide, width 25-35% lobe length; moderate length, length approximately 30% glabellar length; anterior margin located adjacent to 20-30% glabellar length from anterior margin of frontal lobe; furrow shallow to moderate depth. Ocular ridge moderately weak to weak, directed moderately posterolaterally from glabella at 70-75° to axial trend. Posterior area of fixigena exsagittal width 30-40% glabellar length; length 40-55% gla- bella length; bluntly terminated. Posterior border strongly convex, wider slightly distally; border fur- row narrow to moderately wide, deep, wider lat- erally, straight. Librigena, hypostoma, thorax, and pygidium un- known. Exoskeleton thin; finely granulated on external surface and internal mold. Contributions in Science, Number 446 REMARKS. Spencella ? utahensa is most similar to S .? convexa Rasetti, 1963, in the moderately to strongly dorsally convex anterior border, but differs in having a more swollen anterior border and less rounded frontal lobe. Spencella ? utahensa differs from all other species of Spencella in having a mod- erately to strongly dorsally convex anterior border and granular ornamentation. ETYMOLOGY. Latinized, named after the State of Utah. HOLOTYPE. Cranidium (LACMIP 11975) from the Trailer Limestone, Dugway Range, Utah (LAC- MIP locality 15536). PARATYPES. LACMIP 11974 from LACMIP locality 15495; LACMIP 11976 from LACMIP lo- cality 15538; LACMIP 11977 from LACMIP lo- cality 15541; LACMIP 11978 from LACMIP lo- cality 15580. OTHER MATERIAL. LACMIP locality 15495, 6 cranidia; LACMIP locality 15496, 5 cranidia; LACMIP locality 15499, 1 cranidium; LACMIP locality 15515, 1 cranidium; LACMIP locality 15516, 1 cranidium; LAC- MIP locality 15517, 2 cranidia; LACMIP locality 15526, 3 cranidia; LACMIP locality 15528, 1 cranidium; LAC- MIP locality 15534, 1 cranidium; LACMIP locality 15536, 6 cranidia; LACMIP locality 15538, 2 cranidia; LACMIP locality 15540, 2 cranidia; LACMIP locality 15541, 15 Sundberg: Middle Cambrian TrilobitesH 113 Figure 73. Trymataspis cf. T. depressa Robison. Small cranidium (LACMIP hypotype 11979), LACMIP locality 15543, partially exfoliated, x 10.6. cranidia; LACMIP locality 15543, 2 cranidia; LACMIP locality 15547, 1 cranidium; LACMIP locality 15548, 1 cranidium; LACMIP locality 15580, 3 cranidia. OCCURRENCE. Altiocculus Subzone, Ehmani- ella Biozone. Swasey Limestone, Drum Mountains and House Range, Utah. Trailer Limestone, Dug- way Range, Utah. Pole Canyon Limestone, Mem- ber E, Patterson Pass. Genus Trymataspis Robison, 1964a Trymataspis cf. T. depressa Robison, 1964a Figure 73 REMARKS. A small cranidium that resembles Trymataspis depressa Robison, 1964a, was found in the upper portion of the lower member of the Trailer Limestone, Dugway Range. This specimen differs from T, depressa in having a shorter glabella and less prominent occipital spine. HYPOTYPE. LACMIP 11979 from LACMIP lo- cality 15543. OCCURRENCE. Altiocculus Subzone, Ehmani- ella Biozone. Trailer Limestone, Dugway Range, Utah. Genus Yuknessaspis Rasetti, 1951 TYPE SPECIES. Yuknessaspis paradoxa Raset- ti, 1951. EMENDED DIAGNOSIS. Multisegmented, rni- cropygous Ptychopariida with cranidium with mod- erate length glabella, lateral glabellar furrows shal- low to absent, frontal glabellar lobe truncated to slightly rounded, deep preglabellar furrow, anterior portion of axial furrows very deep, straight to con- cave axial furrows, ocular ridges typically strong, palpebral lobe moderate length, posterior area of fixigenae wader than occipital ring, posterior border furrow distally curved anteriorly, fixigenae mod- erately wade, convex preglabellar field and anterior area of fixigena, anterior border slightly to strongly convex. REMARKS. Rasetti (1951) based this genus on three specimens. The diagnosis is modified to reflect the new7 species Y. benningtonis. Yuknessaspis ben - ningtonis is a variable species with three subspecies. The subspecies vary considerably in the shape of the frontal area, glabella shape, axial furrows, and posterior area of fixigenae. Y uknessaspis benningtonis , new species Figures 74-76 Yuknessaspis, n. sp. Sundberg, 1991:388, fig. 2.10. DIAGNOSIS. Yuknessaspis with cranidium with medial swelling, downsloping preglabellar field and anterior area of fixigena. Figure 74. Yuknessaspis benningtonis benningtonis, new species and subspecies. I, Stereo pair of paratype cranidium (LACMIP 11980), LACMIP locality 15495, partially exfoliated, x 8.7. 2, 3, Dorsal and anterior view of holotype cranidium (LACMIP 11983), LACMIP locality 15543, partially exfoliated, x5.0. 114 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites Figure 75. Yuknessaspis benningtonis convexus, new species and subspecies. 1, Stereo pair of partial paratype cra- nidium (LACMIP 11985) with posterior area of fixigena preserved, LACMIP locality 15497, mostly exfoliated, x5.0. 2-4, Stereo pair, lateral, and anterior view of holotype cranidium (LACMIP 11984), LACMIP locality 15497, partially exfoliated, x 10.3. 5, Paratype free cheek (LACMIP 11987), LACMIP locality 15538, exfoliated, x5.1. 6, Stereo pair of partial paratype cranidium (LACMIP 11988), LACMIP locality 15538, partially exfoliated, x6.9. DESCRIPTION. Cranidium small to moderate size, length 2. 2-6.2 mm; subtrapezoidal, length 35- 60% width; strong sagittal and moderate transverse convexity, height 20-45% width; anterior margin slightly to moderately curved, anterior margin width 40-70% cranidial width; posterior margin, exclud- ing occipital ring, very slightly backswept to lateral margins curved anteriorly. Facial sutures parallel to moderately divergent from ends of palpebral lobes to halfway to anterior border; moderately conver- gent from halfway to anterior border to across an- terior border; very strongly divergent posterior of palpebral lobes. Glabella moderate length, length 65-70% and width 40-55% cranidial plan view length; glabella width 20-30% cranidial width; moderately to very strongly tapered, width at an- terior end 55-80% glabellar width; moderate con- vexity (sag. and trans.); frontal lobe bluntly rounded to truncated, 3 lateral glabellar lobes. Axial furrows very deep to moderately deep anterior of ocular ridge, shallower posteriorly, straight to slightly con- vex; preglabellar furrow moderate depth to deep, Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites ■ 115 Figure 76. Yuknessaspis benningtonis planus , new species and subspecies. 1, Stereo pair of holotype cranidium (LACMIP 11990), LACMIP locality 15548, exfoliated, x7.5. 2, Stereo pair of small paratype cranidium (LACMIP 11989), LACMIP locality 15495, slightly exfoliated, x 10.8. slightly curved; lateral glabellar furrows very shal- low, FI furrow bifurcated, directed posteriorly, F2 furrow directed slightly posteriorly, F3 furrow di- rected laterally. Occipital ring sagittal length 20- 35% glabellar length, not elevated above glabella, moderately convex; small occipital node; furrow curved anteriorly, moderate depth, deeper laterally; posterior margin moderately curved. Frontal area plan view length 25-30% cranidial plan view length, unequally divided. Preglabellar field slightly to moderately convex, moderately to very strongly downsloping, plan view length 80-100% frontal area plan view length; medial swelling, low sagit- tally elongate to high to low round. Anterior border slightly to strongly convex, level, slightly to mod- erately tapering laterally, evenly to unevenly curved, sagittal plan view length 0-20% frontal area plan view length. Anterior border furrow evenly to un- evenly curved, very shallow to moderate depth, slightly to greatly shallower than axial furrows. Fixigena width, exclusive of palpebral lobe, 65- 80% glabellar width, very low convexity, upsloping. Palpebral lobe narrow to moderate width, width 25-35% lobe length; moderate length, length 35- 40% glabellar length; anterior margin located ad- jacent to 5-25% glabellar length from anterior mar- gin of frontal lobe; furrow very shallow to shallow. Ocular ridge strong adjacent to glabella, moderately strong laterally, directed slightly posterolaterally from glabella at 75-85° to axial trend. Posterior area of fixigena exsagittal width 35-50% glabellar length; length 90-155% glabella length; projecting slightly posteriorly to bending strongly anteriorly; roundly terminated. Posterior border high, strongly convex, wider laterally, curving partially or com- pletely around border furrow; border furrow nar- row, deep, straight to curving posteriorly, distally curved anteriorly, wider laterally. Librigena moderate size, length 7.1 mm; mod- erately wide, width approximately 55% length with- out spine; lateral margin slightly, unevenly curved. Genal field moderately convex, width approxi- mately 60% librigena! width. Border width ap- proximately 20% librigenal width; gently convex; lateral border furrow shallow, shallowing anteri- orly; posterior border furrow absent. Genal spine broad based, wide, length unknown. Hypostoma, thorax, and pygidium unknown. Exoskeleton thin; coarse to fine genal caeca on preglabellar field, anterior area of fixigena, and ge- nal field of librigena, smooth to scattered moder- ately coarse granules on cranidia, fine pitting on lateral border and genal spine of librigena on ex- ternal surface and internal mold. REMARKS. Yuknessaspis benningtonis con- tains three new subspecies, Y. benningtonis ben- ningtonis, Y. benningtonis convexus, and Y. ben- ningtonis planus. The subspecies occur either together or within short stratigraphic intervals (less than 1 m), suggesting they are different morphol- ogies of the same species. In the Dugway Range, the three subspecies occur within a 0.6-m-thick se- quence of bioclastic limestones at the top of the lower unit of the Trailer Limestone. Yuknessaspis benningtonis convexus occurs in the lower 30 cm of section; Y. benningtonis benningtonis occurs in 116 ■ Contributions in Science, Number 446 Sandberg; Middle Cambrian Trilobites the next 10 cm of strata; and Y. benningtonis pla- nus occurs in the upper 15 cm of the section. In the Drum Mountains, Y. benningtonis bennington- is and Y. benningtonis planus occur together in the same bed with some specimens with intermediate morphologies, which suggests a close relationship of at least these two, very different looking sub- species. Yuknessaspis benningtonis differs from Y. para- doxa Rasetti in having a downsloping preglabellar field and anterior area of fixigena and a medial swelling. ETYMOLOGY. Latinized, named for the col- lector of the holotype, J Bret Bennington. HOLOTYPE. Cranidium (LACMIP 11983) from the Trailer Limestone, Dugway Range, Utah (LAC- MIP locality 15543). PARATYPES. See subspecies. OTHER MATERIAL. See subspecies. OCCURRENCE. Altiocculus Subzone, Ebmani- ella Biozone. See subspecies. Yuknessaspis benningtonis benningtonis, new subspecies Figure 74 Yuknessaspis, n. sp. Sundberg, 1991:388, fig. 2.10. DIAGNOSIS. Yuknessaspis benningtonis with cranidium with sagittally elongate, low, medial swelling; very strong to moderately strong down- sloping preglabellar field and anterior area of fixi- gena; slightly tapered glabella; posterior portion of axial furrows shallow; slightly convex anterior bor- der; transversely long (115-155% glabellar length), anteriorly bent posterior area of fixigenae. REMARKS. Yuknessaspis benningtonis ben- ningtonis differs from Y. benningtonis convexus in having a more tapered glabella, longer posterior area of fixigenae that are backswept, shallower axial furrows, slightly convex anterior border, less steep- ly downsloping preglabellar field, elongate and low medial swelling, and no genal caeca on the pregla- bellar field and anterior area of fixigena. Yuknes- saspis benningtonis benningtonis differs from Y. benningtonis planus in having a stronger down- sloping preglabellar field and anterior area of fixi- gena, converging facial sutures anterior of palpebral lobes, longer posterior area of fixigenae that are anteriorly bent, and slightly convex anterior border. Specimens found in the Drum Mountains show variation in the degree of downsloping of the fron- tal region indicating that the variation of the sub- species ranges into the morphological realm of Y. benningtonis planus. ETYMOLOGY. Latinized, named for my field partner, J Bret Bennington. HOLOTYPE. Cranidium (LACMIP 11983) from the Trailer Limestone, Dugway Range, Utah (LAC- MIP locality 15543). PARATYPES. LACMIP 11980-11982 from LACMIP locality 15495. Contributions in Science, Number 446 OTHER MATERIAL. LACMIP locality 15495, 1 cra- nidium; LACMIP locality 15580, 1 cranidium. OCCURRENCE. Altiocculus Subzone, Ehmani- ella Biozone. Swasey Limestone, Drum Mountains, Utah. Trailer Limestone, Dugway Range, Utah. Pole Canyon Limestone, Member E, Patterson Pass, Ne- vada. Yuknessaspis benningtonis convexus, new subspecies Figure 75 DIAGNOSIS. Yuknessaspis benningtonis with cranidium with rounded very weak to moderately strong medial swelling; very strong downsloping preglabellar field and anterior area of fixigena form- ing sharp flexure in front of glabella; slightly tapered glabella; posterior portion of axial furrows deep; moderately to strongly convex anterior border; transversely shorter (100-110% glabellar length) slightly backswept posterior area of fixigenae. REMARKS. Yuknessaspis benningtonis con- vexus differs from Y. benningtonis benningtonis as discussed above. Yuknessaspis benningtonis con- vexus differs from Y. benningtonis planus in having a strongly downsloping preglabellar field and an- terior area of fixigena, round medial swelling, con- vergent facial sutures anterior of palpebral lobes, moderately coarse genal caeca on frontal region, and deeper posterior portion of the axial furrows. The librigena described above belongs to this subspecies. ETYMOLOGY. Latin, named for the convex frontal region. HOLOTYPE. Cranidium (LACMIP 11984) from the Swasey Limestone, Drum Mountains, Utah (LACMIP locality 15497). PARATYPES. LACMIP 11985 from LACMIP locality 15497; LACMIP 11986-11988 from LAC- MIP locality 15538. OTHER MATERIAL. LACMIP locality 15498, 1 cra- nidium; LACMIP locality 15536, 1 cranidium; LACMIP locality 15538, 4 cranidia; LACMIP locality 15540, 2 cra- nidia; LACMIP locality 15541, 2 cranidia; LACMIP lo- cality 15568, 1 cranidium. OCCURRENCE. Altiocculus Subzone, Ehmani- ella Biozone. Swasey Limestone, Drum Mountains, Utah. Trailer Limestone, Dugway Range, Utah. Pole Canyon Limestone, Member E, Snake Range, Ne- vada. Yuknessaspis benningtonis planus, new subspecies Figure 76 DIAGNOSIS. Yuknessaspis benningtonis with cranidium with sagittally elongate medial swelling; moderately downsloping preglabellar field and an- terior area of fixigena, tapered glabella, posterior portion of axial furrows shallow; moderately con- vex anterior border; transversely shorter (approxi- Sundberg: Middle Cambrian TrilobitesB 117 Figure 77. Ptychopariida sp. unidet. 1, Stereo pair of cranidium (LACMIP hypotype 12009), LACMIP locality 15492, not exfoliated, x 9.8. 2, Cranidium (LACMIP hypotype 11961), LACMIP locality 15489, not exfoliated, x 11.4. 3, Stereo pair of cranidium (LACMIP hypotype 350), LACMIP locality 15492, partially exfoliated, x 8.4. mately 90% glabellar length), slightly backswept posterior area of fixigenae. REMARKS. Yuknessaspis benningtonis planus differs from Y. benningtonis benningtonis and Y. benningtonis concaua as discussed above. Yuknes- saspis benningtonis planus is similar to the Y. para- doxa but differs in having a rounded frontal lobe, shallower axial furrows, moderately convex ante- rior border, and weaker genal caeca in the frontal region. ETYMOLOGY. Latin, planus = level, for the relatively level pregiabellar field. HOLOTYPE. Cranidium (LACMIP 11990) from the Trailer Limestone, Dugway Range, Utah (LAC- MIP locality 15548). PARATYPE. LACMIP 11989 from LACMIP lo- cality 15495. OTHER MATERIAL. LACMIP locality 15545, 2 cranidia; LACMIP locality 15547, 1 cranidium. OCCURRENCE. Altiocculus Subzone, Ehmani- ella Biozone. Swasey Limestone, Drum Mountains, Utah. Trailer Limestone, Dugway Range, Utah. Genus Uncertain Ptychopariida sp. A Figure 77 DESCRIPTION. Cranidium small, length 2.4- 3.9 mm; subtrapezoid, length 70 — 80% width; moderately low convexity (sag. and trans.), height 25-35% width; anterior margin slightly to moder- ately curved, anterior margin width 65-80% cranid- ial width; posterior margin, excluding occipital ring, slightly backswept. Facial sutures parallel to slightly divergent from ends of palpebral lobes to half the distance of the anterior border; moderately to strongly convergent across anterior portion of an- terior border; strongly divergent posterior of pal- pebral lobes. Glabella elongate, length 75-80% and width 50-60% cranidia! length; glabella width 40- 45% cranidial width; very strongly to slightly ta- pered, width at anterior end 55-90% glabellar width; moderate convexity (sag. and trans.); frontal lobe moderately to bluntly rounded, 3(?) lateral glabellar lobes. Axial furrows moderate depth, deeper pos- teriorly, slightly concave to straight; pregiabellar furrow shallow to moderate depth, moderately to slightly curved; lateral glabellar furrows very shal- low or absent. Occipital ring sagittal length 25-30% glabellar length, elevated above glabella, moder- ately convex; very weak occipital node; anteriorly curved furrow, deeper laterally, shallowing medi- ally, moderately deep; posterior margin moderately curved, flattened medially. Frontal area length ap- proximately 25% cranidial length, subequally di- vided. Pregiabellar field slightly concave, down- sloping, length 40-45% frontal area length. Anterior border strongly convex to flat, level, slightly to strongly tapering laterally, evenly curved, sagittal length 55-60% frontal area length. Anterior border furrow evenly curved, narrow, moderately shallow to moderate depth, shallower medially, shallower than axial furrows. Fixigena. width, exclusive of palpebral lobe, 40-50% glabellar width, low con- vexity, level to slightly downsloping; anterior area of fixigena moderately downsloping. Palpebral lobe narrow to wide, width 20-35% lobe length; mod- erate length, length 30-35% glabellar length; an- terior margin located adjacent to 15-20% glabellar 118 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites length from anterior margin of frontal lobe; furrow shallow depth. Ocular ridge weak, directed mod- erately posterolaterally from glabella at 65-75° to axial trend. Posterior area of fixigena exsagittal width 35-40% glabellar length; length 50-60% glabella length; bluntly terminated. Posterior border strong- ly convex, wider slightly distally; border furrow narrow, deep, wider laterally, curved slightly pos- teriorly. Librigena, hypostoma, thorax, and pygidium un- known. Exoskeleton thin; smooth on external and inter- nal surfaces. REMARKS. The nomenclature of Ptychopariida sp. A is left open even though several specimens are known. Although the species is of the general ptychopariid morphology, it cannot be placed in any established genus. Specimens are variable in glabellar shape and anterior border convexity and tapering. These features are typically used to dif- ferentiate genera and species, but the mixing of characters so that no clear separation can be made suggests that these specimens belong to only one species. More specimens are needed from individ- ual localities to establish intrapopulation variation. Ptychoparaiida sp. A can be separated from Deltina in having a smaller glabella, shallower lateral gla- bellar furrows, and concave axial furrows. Ptycho- pariida sp. A can also be separated from Spencella in having longer posterior area of fixigenae and more tapering, less elevated, conical glabella. HYPOTYPES. LACMIP 11961 from LACMIP locality 15489; LACMIP 12009, 12010 from LAC- MIP locality 15492. OTHER MATERIAL. LACMIP locality 15489, 2 cranidia; LACMIP locality 15491, 1 cranidium; LACMIP locality 15493, 2 cranidia; LACMIP locality 15527, 1 cra- nidium; LACMIP locality 15532, 4 cranidia; LACMIP locality 15533, 3 cranidia; LACMIP locality 15534, 3 cra- nidia. OCCURRENCE. Altiocculus Subzone, Ehmani- ella Biozone. Swasey Limestone, Drum Mountains, Utah. Trailer Limestone, Dugway Range, Utah. ACKNOWLEDGMENTS This paper was part of a Ph.D. dissertation (Sundberg, 1990) completed at Virginia Polytechnic Institution and State University. I wish to thank the following people for their assistance in this project. Richard K. Bambach and J. John Sepkoski, Jr., reviewed the systematic portion of my dissertation. J Bret Bennington was my field partner in Utah and Nevada and translated Russian trilobite de- scriptions. Access to trilobite collections were provided by Richard A. Robison, University of Kansas; A1 Kamb, University of Kansas Museum of Invertebrate Paleontol- ogy; Michael E. Taylor, United States Geological Survey, Denver; and Fred Collier, United States National Muse- um. Funding for dissertation research was from my par- ents, Larry and Eileen Sundberg; my wife, Debbie Whit- ney-Sundberg; Society of Sigma Xi; Department of Geological Sciences, Virginia Polytechnic Institution and State University; and Appalachian Basin Industrial Asso- ciates. I also wish to thank Edward C. Wilson, who re- viewed the manuscript, and Richard Robison and Fred- erick R. Schram, who provided additional comments. Catherine Agra assisted in Greek and Latin construction of scientific names. LITERATURE CITED Beecher, C.E. 1897. Outline of a natural classification of the trilobites. American Journal of Science, series 4, 3:89-106, 181-207. Briggs, D.E.G., and D. Collins. 1988. A Middle Cam- brian chelicerate from Mount Stephen, British Co- lumbia. Palaeontology 31:779-798. Campbell, D.P. 1974. 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Ca- nadian Alpine Journal 1:232-248. . 1916. Cambrian geology and paleontology of Cambrian trilobites. Smithsonian Miscellaneous Collections 64:303-456. - — 1918. Appendages of trilobites. Smithsonian Miscellaneous Collections 67:115-216. . 1924. Cambrian and Ozarkian trilobites. Smith- sonian Miscellaneous Collections 75:53-60. . 1925. Cambrian and Ozarkian trilobites. Smith- sonian Miscellaneous Collections 75(3):1-146. Westergard, A.H. 1936. Paradoxides oelandicus beds of Oland. Sveriges Geologiska Under soekning, series C, 394:1-66. White, C. A. 1874. Preliminary report upon invertebrate fossils, with descriptions of new species. United States Geographical Exploration of the West 100th Merid- ian [Wheeler Survey], pamphlet, 27 pp. White, W.W., III. 1973. Paleontology and depositional environments of the Cambrian Wheeler Formation, Drum Mountains, west-central Utah. Unpublished M.S. thesis, University of Utah, Salt Lake City, 135 pp. Whitehouse, F.W. 1945. The Cambrian faunas of north- eastern Australia. Part 5. The trilobite genus Do- rypyge. Memoirs of the Queensland Museum 12(3): 117-123. Whittington, H.B. 1975. Trilobites with appendages from the Middle Cambrian, Burgess Shale, British Colum- bia. Fossils and Strata 4:97-136. . 1980. Exoskeleton, moult stage, appendage morphology and habits of the Middle Cambrian tri- lobite Olenoides serratus. Palaeontology 23:171- 204. . 1992. Trilobites. Suffolk, England: Boydell Press, 145 pp., 120 pis. Young, G.A., and R. Ludvigsen. 1989. Mid-Cambrian trilobites from the lowest part of the Cow Head Group, western Newfoundland. Geological Survey of Canada, Bulletin 392:1-49. Received 7 January 1993; accepted 4 August 1993. APPENDIX: LOCALITY REGISTER DRUM MOUNTAINS SECTIONS. Chi, Dol, Do2, Wl, Swl, Sw2, and Bl. LOCATION. Section 17, T. 15 S., R. 11 W., USGS topographic map Drum Mts. Well 7.5' Quadrangle, Utah (1971); Drum Mountains, Millard County, Utah (Fig. 78). DESCRIPTION. The measured sections in the Drum Mountains cover an incomplete interval of the Chisholm Formation (54.5 m, sections Chi and Dol) and complete intervals of the Dome Limestone (54.7 m, section Do2), Whirlwind Formation (42.5 m, section Wl), and Swasey Limestone (60.5 m, sections Swl and Sw2) (Figs. 79-81). A short interval from the uppermost Swasey and lower- most Wheeler Shale (1.55 m, section Bl) was also mea- sured and collected. The sections were measured in north- ward-trending canyons approximately 1.1 km (0.7 mi) north-northwest of the proposed stratotype for agnostid biozones in the Wheeler Shale (Rowell et al., 1982). FOSSIL LOCALITIES. LACMIP 15454. Chisholm Formation, section Dol, 47 m above base of section, float. LACMIP 15455. Chisholm Formation, section Do2, 2.0 m above base of section, collected adjacent to section from float. Figure 78. Location of stratigraphic sections in the Drum Mountains, Utah. From Drum Mts. Well (1971) and Lady Laird Peak (1988) 7.5' quadrangles. Contour intervals 20 and 40 ft. 122 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites LACMIP 15456. Chisholm Formation, section Do2, 2.35 m above base of section. LACMIP 15457. Chisholm Formation, section Do2, 2.37 m above base of section. LACMIP 15458. Chisholm Formation, section Do2, 2.39 m above base of section. LACMIP 15459. Chisholm Formation, section Do2, 4.0- 4.5 m above base of section, float. LACMIP 15460. Chisholm Formation, section Do2, 5.0 m above base of section, float. LACMIP 15461. Chisholm Formation, section Do2, 6.0 m above base of section. LACMIP 15462. Dome Limestone, section Do2, 39.5 m above base of section. LACMIP 15463. Dome Limestone, section Wl, 2.0 m above base of section. LACMIP 15464. Dome Limestone, section Wl, 3.2 m above base of section. LACMIP 15465. Whirlwind Formation, section Wl, 20.6 m above base of section. LACMIP 15466. Whirlwind Formation, section Wl, 21.2 m above base of section. LACMIP 15467. Whirlwind Formation, section Wl, 28.9 m above base of section. LACMIP 15468. Whirlwind Formation, section Wl, 29.4 m above base of section. LACMIP 15469. Whirlwind Formation, section Wl, 29.9 m above base of section. LACMIP 15470. Whirlwind Formation, section Wl, 30.9 m above base of section. LACMIP 15471. Whirlwind Formation, section Wl, 31.5 m above base of section. LACMIP 15472. Whirlwind Formation, section Wl, 34.9 m above base of section. LACMIP 15473. Whirlwind Formation, section Wl, 35.8 m above base of section. LACMIP 15474. Whirlwind Formation, section Wl, 37.6 m above base of section. LACMIP 15475. Whirlwind Formation, section Wl, 42.0 m above base of section, float. LACMIP 15476. Swasey Limestone, section Swl, 0.4 m above base of section. LACMIP 15477. Swasey Limestone, section Swl, 0.7 m above base of section. LACMIP 15478. Swasey Limestone, section Swl, 1.2 m above base of section. LACMIP 15479. Swasey Limestone, section Swl, 1.6 m above base of section. LACMIP 15480. Swasey Limestone, section Swl, 4.5 m above base of section. LACMIP 15481. Swasey Limestone, section Swl, 5.0 m above base of section. LACMIP 15482. Swasey Limestone, section Swl, 6.3 m above base of section. LACMIP 15483. Swasey Limestone, section Swl, 9.1 m above base of section. LACMIP 15484. Swasey Limestone, section Swl, 16.8 m above base of section. LACMIP 15485. Swasey Limestone, section Swl, 18.2 m above base of section. LACMIP 15486. Swasey Limestone, section Swl, 19.7 m above base of section. LACMIP 15489. Swasey Limestone, section Sw2, 19.7 m above base of section. LACMIP 15490. Swasey Limestone, section Sw2, 20.5 m above base of section. LACMIP 15491. Swasey Limestone, section Sw2, 21.5 m above base of section. o o o © © £? ~W] - _Q a, g C/5 ’5b © _2 oo o 3 00 »- c &0 e°a ZO CA e ■8 •e ON hv bb £ so P3 m £ Xfl n ° ON r? i-i o LQ in id in m 31BJ3UJ0|8U03 — jOAPig <— auoisujpjft OUOJS^DPd 3UO)S33f DBM Duoispnui D|IM|N CO O to to ^ ^ 3U0]S3Uin ^SSBMQ Q © ’ ^ .o in in in in 3S in in m m 't in TM' f i ( llili i S \fh ( vH\ T J lift Ik IH-Ol H- UOlJBUIJOj pUlMjJiq^ 128 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites Figure 84. Detailed sections W2, Sw3, and B3 from the House Range showing LACMIP localities. (See Fig. 79 for lithologic symbols.) Figure 85. Location of stratigraphic sections in the Dugway Range, Utah. From Dugway Range NE 7.5' Quadrangle (1953). Contour interval 40 ft. LACMIP 15515. Uppermost Swasey Limestone, section B3, 3.14 m above base of section. LACMIP 15516. Uppermost Swasey Limestone, section B3, 3.27 m above base of section. LACMIP 15517. Uppermost Swasey Limestone, section B3, 3.44 m above base of section. LACMIP 15518. Uppermost Swasey Limestone, section B3, 3.51 m above base of section. LACMIP 15519. Uppermost Wheeler Shale, section B3, 3.60 m above base of section. RC 107. Dome Limestone, collected approximately 23 m above the base of the formation. North side of Marjum Canyon, approximately 1.6 km east of canyon mouth, House Range, Millard Co., Utah. Collected by R. Ro- bison, 1967. RC 211. Dome Limestone, collected from a large block at the base of the Dome Limestone cliff on a northeast- trending ridge approximately 1.7 km from the east end of Marjum Canyon. The fossils are from approximately 5 m below the top of the block. (Redescribed from Oldroyd [1973] and KUMIP locality cards.) Collected by John D. Oldroyd. RC 374. Dome Limestone approximately 23 m above base of formation(?). North side of isolated pinnacle adja- cent to massive cliff on north side of Marjum Cyn. NWV4, NEV4, SWV4, section 15, T. 18 S., R. 14 W., Notch Peak (same as 107 and 211). Collectors Robison and Oldroyd, 1975. RC 375. Dome Limestone at base of slope southeast of KUMIP locality 374; talus from unknown unit. Col- lector Oldroyd, 1975. DUGWAY RANGE SECTIONS. Ssl(c), Trl, and B5. LOCATION. Section 3 (projected), T. 11 S., R. 11 W., USGS topographic map Dugway Range 7.5' Quadrangle, Utah (1953); Dugway Range, Juab County, Utah (Fig. 85). DESCRIPTION. The measured sections in the Dugway Range cover incomplete intervals of the Shadscale For- mation (32 m, section Ssl[c]) and Trailer Limestone (37.5 m, sections Trl and B5) (Fig. 86). Section B5 consists of the upper 12.5 m of section Trl. The sections are located on low-lying hills on the southeast side of the Dugway Range approximately 3.7 km (2.3 mi) west-northwest of the Dugway Topaz Well and 1.1 mi north of Shadscale Canyon. FOSSIL LOCALITIES. LACMIP 15520. Shadscale Formation, section Ssl(c), 4.6 m above base of section. LACMIP 15521. Shadscale Formation, section Ssl(c), 6.7 m above base of section, float. LACMIP 15522. Shadscale Formation, section Ssl(c), 11.3 m above base of section, float. LACMIP 15523. Shadscale Formation, section Ssl(c), 13.8 m above base of section, float. LACMIP 15525. Shadscale Formation, section Ssl(c), 23.5 m above base of section, float. LACMIP 15526. Lower Trailer Limestone, sections B5 and Trl, 25.80 m above base of sections. LACMIP 15527. Lower Trailer Limestone, sections B5 and Trl, 25.95 m above base of sections. LACMIP 15528. Lower Trailer Limestone, sections B5 and Trl, 26.75 m above base of sections. LACMIP 15529. Lower Trailer Limestone, sections B5 and Trl, 27.05 m above base of sections. LACMIP 15530. Lower Trailer Limestone, sections B5 and Trl, 27.15 m above base of sections. LACMIP 15531. Lower Trailer Limestone, sections B5 and Trl, 27.37 m above base of sections. Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites ■ 129 Dugway Range Ssl(c) PQ in m m lo in in in 3!CJ3UI0|SU03 ” |3abj8 OONLO^fONH OOCC^VO Penmen in m m in in m in mininm inininminmin mininm i-H t— ( r-H t— ^ ' r-* 1 HHHr' W A \ I / ]/// W ' c _o o H) on c G .S "C o DC .£9 >IUL ,(3)ISS jaddn J3MOI auojsauin uoncunoj aieospuiis — • -i snjnojojijv | tjjaiuvwy^ l L vjpi^WJ]3 | vjjsluvwysojj vunajdossojQ ui 130 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites Figure 86. Generalized stratigraphic section of the Ehmaniella Biozone in the Dugway Range, Utah, and detailed sections Ssl(c), Trl, and B5 showing LACMIP localities. (See Fig. 79 for lithologic symbols.) Figure 87. Location of stratigraphic sections in the Snake Range, Nevada. From Minerva Canyon 7.5' Quadrangle (1987). Contour interval 40 ft. LACMIP 15532. Lower Trailer Limestone, sections B5 and Trl, 29.95 m above base of sections. LACMIP 15533. Lower Trailer Limestone, sections B5 and Trl, 30.10 m above base of sections. LACMIP 15534. Lower Trailer Limestone, sections B5 and Trl, 31.40 m above base of sections. LACMIP 15535. Lower Trailer Limestone, sections B5 and Trl, 31.50 m above base of sections. LACMIP 15536. Lower Trailer Limestone, sections B5 and Trl, 36.40 m above base of sections. LACMIP 15537. Lower Trailer Limestone, sections B5 and Trl, 36.50 m above base of sections. LACMIP 15538. Lower Trailer Limestone, sections B5 and Trl, 36.54 m above base of sections. LACMIP 15540. Lower Trailer Limestone, sections B5 and Trl, 36.56 m above base of sections. LACMIP 15541. Lower Trailer Limestone, sections B5 and Trl, 36.61 m above base of sections. LACMIP 15542. Lower Trailer Limestone, sections B5 and Trl, 36.75 m above base of sections. LACMIP 15543. Lower Trailer Limestone, sections B5 and Trl, 36.80 m above base of sections. LACMIP 15544. Lower Trailer Limestone, sections B5 and Trl, 36.94 m above base of sections. LACMIP 15545. Lower Trailer Limestone, sections B5 and Trl, 36.97 m above base of sections. LACMIP 15547. Lower Trailer Limestone, sections B5 and Trl, 37.03 m above base of sections. LACMIP 15548. Lower Trailer Limestone, sections B5 and Trl, 37.10 m above base of sections. SNAKE RANGE SECTIONS. Peel, Peel, Pce3, and B6 LOCATIONS. Peel — section 4 (projected), T. 11 N., R. 68 E., LfSGS topographic map Minerva Canyon 7.5' Quadrangle, Nevada (1987); Snake Range, White Pine County, Nevada (Fig. 87). Peel — sections 4 and 9 (pro- jected) T. 1 1 N., R. 68 E., USGS topographic map Minerva Canyon 7.5' Quadrangle, Nevada (1987); Snake Range, White Pine County, Nevada (Fig. 87). Pce3 and B6 — section 3 (projected), T. 1 1 N., R. 68 E., USGS topographic map Minerva Canyon 7.5' Quadrangle, Nevada (1987); Snake Range, White Pine County, Nevada (Fig. 87). DESCRIPTION. The measured sections in the Snake Range cover an incomplete interval of Member C of the Pole Canyon Limestone (14 m, section Peel) and a com- plete interval of Member E of the Pole Canyon Limestone (89.5 m, section Peel) (Figs. 88 and 89). Only the low- ermost portion of Member D of the Pole Canyon Lime- stone (6 m, section Peel) was measured. A short interval from the uppermost Member E of the Pole Canyon Lime- stone and lowermost Lincoln Park Formation (4.0 m, section B6) was also measured and collected. The section for Member C of the Pole Canyon Limestone (Peel) was measured on the north side of Swallow Canyon, east 0.75 km (0.47 mi) from the mouth of the canyon. The sections for Member E of the Pole Canyon Limestone (Peel, Pce3, and B6) were measured on the south side of Swallow Canyon, east approximately 0.95 km (0.6 mi, section Peel) and 2.0 km (1.25 m, sections Pce3 and B6) from the mouth of the canyon. FOSSIL LOCALITIES. LACMIP 15549. Pole Canyon Limestone, Member C, section Peel, float. LACMIP 15550. Pole Canyon Limestone, Member C, section Peel, 8.5 m above base of section. LACMIP 15551. Pole Canyon Limestone, Member C, section Peel, 8.5 m above base of section, float. LACMIP 15552. Pole Canyon Limestone, Member C, section Peel, 8.8 m above base of section. Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites ■ 131 0> u 0H .. . m m mm m in to mm in in in mm in in qr\ no ON do OJD a m & CJ & es ti m ntsom m m in m imn inm imniflr* d .2 o » C/3 e c N •c o ffi .90 • £*>d ,|33d 1 133d QJ 2 2. 3 tO Figure 89. Detailed sections Pce3 and B6 from the Snake Range showing LACMIP localities. (See Fig. 79 for lithologic symbols.) LACMIP 15553. Pole Canyon Limestone, Member C, section Peel, 9.0 m above base of section. LACMIP 15554. Pole Canyon Limestone, Member E, section Peel, float near base of section. LACMIP 15555. Pole Canyon Limestone, Member E, section Peel, 12.2-17.0 m above base of section, float. LACMIP 15556. Pole Canyon Limestone, Member E, section Peel, 14.5 m above base of section. LACMIP 15557. Pole Canyon Limestone, Member E, section Peel, 15.7 m above base of section, float. LACMIP 15558. Pole Canyon Limestone, Member E, section Peel, 16.3 m above base of section, float. LACMIP 15559. Pole Canyon Limestone, Member E, section Peel, 17.2 m above base of section, float. LACMIP 15560. Pole Canyon Limestone, Member E, section Peel, 60.7 m above base of section. LACMIP 15561. Pole Canyon Limestone, Member E, section Peel, 68.5 m above base of section. LACMIP 15562. Pole Canyon Limestone, Member E, section Peel, 69.5 m above base of section. LACMIP 15563. Lincoln Peak Formation, section B6, first meter of formation (2. 9-4.0 m above base of section), float. LACMIP 15564. Pole Canyon Limestone, uppermost Member E, section B6, 2.19 m above base of section. LACMIP 15565. Pole Canyon Limestone, uppermost Member E, section B6, 2.25 m above base of section, collected along strike. LACMIP 15566. Pole Canyon Limestone, uppermost Member E, section B6, 2.25 m above base of section. LACMIP 15567. Pole Canyon Limestone, uppermost Member E, section B6, 2.43 m above base of section. LACMIP 15568. Pole Canyon Limestone, uppermost Member E, section B6, 2.47 m above base of section. LACMIP 15569. Pole Canyon Limestone, uppermost Member E, section B6, 2.60 to 2.80 m above base of section. LACMIP 15570. Pole Canyon Limestone, uppermost Member E, section B6, 2.55 m above base of section. LACMIP 15571. Pole Canyon Limestone, uppermost Member E, section B6, 2.85 m above base of section. LACMIP 15572. Pole Canyon Limestone, uppermost Member E, section B6, 2.93 m above base of section. USGS 1982-CO. Pole Canyon Limestone, Minerva Dis- trict. PATTERSON PASS (SOUTHERN SCHELL CREEK RANGE) SECTIONS. Pce4 and B7. LOCATION. Section 36 (projected), T. 9 N., R. 64 W., USGS topographic map Milk Ranch Spring 7.5' Quad- rangle, Nevada (1973); Southern Schell Creek Range, Lin- coln County, Nevada (Fig. 90). DESCRIPTION. The measured sections in the south- ern Schell Creek Range cover Member E (148 m, section Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites ■ 133 Figure 90. Location of stratigraphic sections in the Patterson Pass area, southern Schell Creek Range, Nevada. From Single Pass SE (1969) and Milk Ranch Spring (1973) 7.5' quadrangles. Contour interval 40 ft. Pce4) of the Pole Canyon Limestone and a short interval from the uppermost Member E of the Pole Canyon Lime- stone and lowermost Patterson Pass Shale (2.5 m, section B7) (Fig. 91). The sections for Member E of the Pole Canyon Limestone (Pce4 and B7) were measured on the north side of canyon approximately 1.4 km (0.9 mi) west of Patterson Pass. FOSSIL LOCALITIES. LACMIP 15573. Pole Canyon Limestone, Member E, section Pce4, 7.7 m above base of section. LACMIP 15574. Pole Canyon Limestone, Member E, section Pce4, 8.2 m above base of section. LACMIP 15575. Pole Canyon Limestone, Member E, section Pce4, 8.8 m above base of section. LACMIP 15576. Pole Canyon Limestone, Member E, section Pce4, 13.5 m above base of section. LACMIP 15577. Pole Canyon Limestone, Member E, section Pce4, 17.4 m above base of section. LACMIP 15578. Pole Canyon Limestone, Member E, section Pce4, 17.5 m above base of section. LACMIP 15579. Pole Canyon Limestone, Member E, section Pce4, 20.4-22.5 m above base of section, float. LACMIP 15580. Pole Canyon Limestone, uppermost Member E, section B7, 0.45 m above base of section. LACMIP 15581. Patterson Pass Shale, uppermost Mem- ber E, section B7, 0.53 m above base of section. NORTHERN EGAN RANGE SECTIONS. Ell and B8. LOCATION. T. 22 N., R. 63 E., USGS topographic map Borchert Springs, Nevada 7.5' Quadrangle, Nevada (1982); northern Egan Range, White Pine County, Nevada (Fig. 92). DESCRIPTION. The measured section in the northern Egan Range covers an incomplete interval of the Eldorado Formation (240 m, section Ell) and Secret Canyon For- mation (5 m, section Ell) (Fig. 93). A short interval from the shale unit at the base of the measured section (1.5 m, section B8; not the base of the Eldorado Formation) was also measured and collected. The lower portion of the section is located on an east-west canyon side and the upper portion is located on the adjacent ridge to the north, on the east face of the northern Egan Range, approxi- mately 2.0 km (1.25 mi) due west of a location on road at approximately 1 6.6 km (10.4 mi) south of Cherry Creek. The locations of the sections on the locality map are approximate; they were originally plotted on the Ely, Ne- vada; Utah, USGS 2° Quadrangle (1971). FOSSIL LOCALITIES. LACMIP 15582. Eldorado Formation, section Ell, float. LACMIP 15583. Eldorado Formation, section Ell, 2.0 m above base of section, found along strike, float. LACMIP 15584. Eldorado Formation, section Ell, 4.0- 7.0 m above base of section, float. LACMIP 15585. Eldorado Formation, section Ell, 5.3 m above base of section, found 3 m along strike. LACMIP 15586. Eldorado Formation, section Ell, 5.7 m above base of section, found 3 m along strike. LACMIP 15587. Eldorado Formation, section Ell, 5.7 m above base of section. LACMIP 15588. Eldorado Formation, section Ell, 6.5 m above base of section, found 4 m along strike. 134 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites r- [3AEiS SUOJSUSSiS C3 C/3 Pu M ^s OJ Contributions in Science, Number 446 Sundberg: Middle Cambrian TrilobitesH 135 Figure 91. Generalized stratigraphic section of the Rbmanieila Biozone in the Patterson Pass area, Nevada, and detailed sections Pce4 and B7 showing LACMIP localitie 79 for lithologic symbols.) Figure 92. Location of stratigraphic sections in the northern Egan Range, Nevada. From Borchert Springs (1982) and Monte Neva Hot Springs (1982) 7.5' quadrangles. Contour interval 10 m. LACMIP 15589. Eldorado Formation, section Ell, 8.3 m above base of section, found 4 m along strike. LACMIP 15590. Eldorado Formation, section Ell, 115.0 m above base of section. LACMIP 15591. Eldorado Formation, section Ell, 148.5 m above base of section. LACMIP 15592. Eldorado Formation, section Ell, 148.7 m above base of section. LACMIP 15593. Eldorado Formation, section Ell, 149.6 m above base of section. LACMIP 15594. Eldorado Formation, section Ell, 190.0 m above base of section. LACMIP 15595. Eldorado Formation, section Ell, 190.2 m above base of section. LACMIP 15596. Eldorado Formation, section Ell, 191,0 m above base of section. LACMIP 15597. Eldorado Formation, section Ell, 191.5 m above base of section. LACMIP 15598. Eldorado Formation, section Ell, 192.5 m above base of section. LACMIP 15599. Eldorado Formation, section Ell, 206.0 m above base of section. LACMIP 15600. Eldorado Formation, section Ell, 228.5 m above base of section. LACMIP 15601. Eldorado Formation, section Ell, 231.0 m above base of section. LACMIP 15602. Eldorado Formation, section Ell, 237.0 m above base of section. LACMIP 15603. Eldorado Formation, section Ell, 238.5 m above base of section. LACMIP 15604. Eldorado Formation, section Ell, 238.5 m above base of section, found approximately 10 m along strike. LACMIP 15605. Eldorado Formation, section Ell, 238.7 m above base of section, found approximately 10 m along strike. LACMIP 15606. Eldorado Formation, section Ell, 238.8 m above base of section. LACMIP 15607. Eldorado Formation, section B8, 0.00 m above base of section. LACMIP 15608. Eldorado Formation, section B8, 0.30 m above base of section. LACMIP 15609. Eldorado Formation, section B8, 0.45 m above base of section. LACMIP 15610. Eldorado Formation, section B8, 0.50 m above base of section. LACMIP 15611. Eldorado Formation, section B8, 0.60 m above base of section. LACMIP 15612. Eldorado Formation, section B8, 0.80 m above base of section. LACMIP 15613. Eldorado Formation, section B8, 0.90 m above base of section. 136 ■ Contributions in Science, Number 446 Sundberg: Middle Cambrian Trilobites O O O ~ \Ov Q NO O in minio v T O CN “ o o nO in o o \Q in & in in rsfs f\r*^ IV 3 fw. sms UOXUB^ 13J33S oo in no in t* ON ON CJN 0N ON in in m m in in in mm in "T” o m ni 'W, (\rv ^ ^ 1 ® ® £><§) C/5 T3 6£ C ce 0S5 ■c o X *-» 3IBMS g UOXUBQ J3JD3S eflHHeHHbix ^aHHbix no MeTpunecKHM h HeMeTpnHecKHM npH3HaKaM corjiacyioTCH c rnnoTe3on, b cootbctctbuh c kotopoh HeMeTpHHecKne npH3HaKH ne HeinpajibHbi. nocKonbKy HeMeTpunecKHe npH3HaKH oSjianaiOT MeHbineH pa3peinaioineH cnocoSnocTbio h nocKOjibKy npn nojiyneHHH BOcnponsBOAHMbix on,eHOK 3thx npH3HaKOB B03HHKaioT Tpy^HOCTH, peKOMeH/jyeTCH hx Hcnojib30BaTb npn HCCjieAOBaHHH nonyjumHH nejib(])HHOB TOJibKo b coneTaHHH c MeTpHnecKHMH npH3HaKaMH, y^ejiBH oco6oe BHHMaHHe coSjnoneHHio ycjiOBHH, o6ecneHHBaiomHX MaKCHMajibHyio conocTaBHMOCTb naHHbix. 1. Southwest Fisheries Science Center, P.O. Box 271, La Jolla, California 92038. 2. Institute of Developmental Biology, Vavilov Street 26, Moscow 117334, Russia. Contributions in Science, Number 447, pp. 1-15 Natural History Museum of Los Angeles County, 1994 ISSN 0459-8113 INTRODUCTION Non-metric characters (features coded by state, e.g., “present” or “absent”) are potentially more useful than metric characters for discriminating popula- tions of mammals because (1) a large number of relatively independent characters can more easily be defined and (2) the characters are supposedly more “neutral” in relation to adaptation and there- fore good indicators of degree of gene flow be- tween populations (Sjovold, 1973; Hartman, 1980). An additional attraction of non-metric characters for the cetologist is that they are less affected by preparation methods and specimen handling; for example, dolphin skulls in museums very frequently have broken rostra or dried tissue remaining on the rostrum tips, broken pterygoids, etc., and this often leads to missing values for several measurements. Non-metric characters have been used in popu- lation analysis of several marine mammals: gray seals, Halichoerus grypus (Berry, 1969); ribbon seals, Phoca fasciata, and spotted seals, P. largha (Fe- doseev, 1984); pantropical spotted dolphins, Ste- nella attenuata, and spinner dolphins, S. longiros- tris (Perrin et al., 1982; Yablokov et al., 1983); and harbor propoises, Phocoena phocoena (Kinze, 1985, 1990), but the quality of their analytic power rel- ative to that of metric characters has not been crit- ically examined. The purpose of the present study was to compare the power of comparable suites of metric and non-metric characters to discriminate among manifestly isolated populations of dol- phins— in this case, three geographically isolated populations of the short-beaked common dolphin, Delphinus delphis — with the intention of possibly using only one or the other of the two kinds of characters in future analyses of population structure for this and other dolphin species. MATERIALS AND METHODS THE SAMPLES A total of 181 skulls of sexually mature adults was in- cluded in the analyses (Appendix 1). They came from the eastern tropical Pacific (73), the western and mid-North Atlantic (71), and the Black Sea (37). Distal fusion of the premaxillae and maxillae was used as an approximate criterion of sexual maturity (Dailey and Perrin, 1973). The eastern Pacific skulls (in the U.S. National Museum of Natural History [USNM], Washington, D.C.) were of dol- phins killed incidentally during commercial tuna-seining operations. The dolphins were captured in nine purse- seine hauls by several vessels over a period of three weeks in the same general area in oceanic waters off Central America. The common dolphins in this area are thought to belong to a single breeding population (Perrin et al., 1985). The North Atlantic specimens (also in the USNM) came from standings and scientific captures and from in- cidental kills in commercial trawls and purse seines along the east coast of North America from Nova Scotia to Florida (67) and the mid-North Atlantic; it is not known how many breeding populations are represented in this sample. The Black Sea material (in the Zoological Mu- seum of Moscow State University) came from the former Soviet commercial dolphin fishery; the population there has been recognized as a subspecies, Delphinus delphis ponticus (Tomilin, 1957). METRIC CHARACTERS Twenty-seven skull measurements were taken as de- scribed in Perrin (1975) and are listed in Appendix 2. Tooth counts were treated as continuous variables, for a total of 31 metric variables. NON-METRIC CHARACTERS For the non-metric analyses, we began with the characters used in Perrin et al. (1982) and Yablokov et al. (1983). Analyses were performed before the final scorings to quantify variation among observers, variation with sex of the specimen, and correlations among the characters. Pre- liminary results of these analyses were reported in Perrin et al. (1988). Upon examination of Delphinus material at the Southwest Fisheries Science Center and a preliminary fist-round scoring of the eastern tropical Pacific series at the USNM by WFP and AVY (in 1987), we deleted some of the characters and added others, to arrive at a total of 62. A number of additional characters were considered but not included, because of very low variability or dif- ficulty of scoring. WFP and AVY then each scored the series independently twice, with redefinition, deletion, and addition of characters between the rounds, to generate two separate data sets (WFP and AVY). Logistics pre- vented simultaneous “blind” scoring of all specimens at the same location and time. Methods Used for Correlation Analyses It is to be expected that at least some of the non-metric variables that were scored would be highly correlated with others. In such cases, it may be possible to drop one or more characters without losing significant information. When using metric variables, the Pearson product-mo- ment correlation coefficient is typically used to describe such inter-relationship between variables. We used a non- metric analog to the Pearson statistic which calculates a coefficient of correlation based on a chi-square approach (Urbach, 1964). The coefficient, R, can have values ranging from 0 (no correlation) to 1.0 (perfect correlation). It is calculated as _ chi-square maximum possible chi-square NV(*. - W„ - 1)’ where N = number of specimens, ka = number of classes of character A, and kb ~ number of classes of character B. The significance of the resulting correlation was de- termined by applying a chi-square contingency test. Cor- relation coefficients were therefore not calculated for some characters that had only two states with observations (a common state and a very rare state). For purposes of the analyses, the states of characters with more than three states (i.e., numbers of foramina or fenestrations) were aggregated into three pooled “states,” e.g., 1-3, 4-6, and 7-9 for one character. The same approach was used for analysis of correlations with sex of the specimen. Comparison of Observers Agreement between WFP and AVY in the first round of scoring ranged between 100% and 55-60% for the initial 2 ■ Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins 62 characters (Fig. 1A). After detailed discussion of the individual characters, 17 were dropped from the series because of low agreement, low variability, or problems of character-state definition. Seven new characters were devised and added to the series, for a new total of 52. Agreement in the second round of scoring was better (Fig. IB), but there was still very significant disagreement (more than 10%) for about one-third of the characters. Correlations among the Characters Five hundred fifty-four correlation analyses were possible (the number of observations for each state of the char- acters was five or greater) for the WFP data set and 583 for the AVY set; details are given in Perrin et al. (1988). The total numbers of significant correlations (p < 0.05) between variables were 44 (WFP) and 46 (AVY). The expected number of significant comparisons given 550- 600 pairwise tests is approximately 28-30 by chance alone. Flowever, 19 of the significant correlations were common to both sets of data. Six (WFP) and seven (AVY) of the significant correlations occurred with paired left/right characters for the same skull feature; of these five were present in both data sets. We concluded that correlations between the characters are significantly more common than would be expected by chance alone and that an appreciable amount of the redundancy in the variables could be eliminated by using only one of a left/right pair of characters. Correlations with Sex of the Specimen For the WFP data set, 34 analyses of correlation with sex were possible; for the AVY set, 36 were possible. Four characters were significantly correlated with sex in the WFP set and five in the AVY set (p < 0.05). Only one or two characters could be expected to be significantly cor- related with sex by chance alone. Two characters were correlated with sex in both data sets; these were consid- ered to be good candidates for characters to be avoided in comparisons of samples with different or unknown sex compositions. Further Reduction of the Character Set Based on the results of the comparisons of observers, 18 of the 52 characters were dropped from the set for com- parison of the geographical samples. These were the char- acters for which observer agreement was less than 90% in the second comparison round (Fig. IB). One of the characters deleted because of low repeatability was also significantly correlated with sex in both of the data sets. Because many of the Black Sea specimens were of un- known sex, the other character correlated with sex was also deleted. Of the five character pairs with significant within-pair correlation in both character sets, either one or both members of the pair were deleted for three pairs based on low observer agreement as described above. For the other two such character pairs, only the left-side mem- ber was retained. After all these deletions, 31 characters remained in the set for the geographical analyses (not 26, as stated in Perrin et al., 1988). A further modification was to assign three possible states for all characters that were not counts of foramina, with the third state being “intermediate,” “at the level of . . . ,” or “uncertain,” as appropriate, to allow more homogeneous analysis and interpretation of the data. Upon examination of the Black Sea series, 5 of the 31 characters proved to be unusable because of damage to the specimens (see Discussion). The remaining 26 char- 100 90 80 70 60 50 40 AGREEMENT (percent) Figure 1. Histograms of percentage of agreement be- tween WFP and AVY in (A) first round of scoring and (B) second round of scoring 73 skulls of Delphinus delphis for non-metric characters. In A, shaded portions represent characters deleted for the second round. In B, numbers represent characters as listed in Appendix 1 of Perrin et al. (1988); circled characters were added as new characters for the second round. acters are described in Appendix 3. WFP and AVY mea- sured and scored the Black Sea series in Moscow on 4- 5 November 1988 and the two USNM series in Wash- ington, D.C., from 31 March to 5 April 1989. For the non-metric characters, each scored all the skulls sepa- rately. The two sets of scores were compared, and dif- ferences were resolved through discussion and re-exam- ination of the skulls, to arrive at a single agreed data set for each of the three geographical samples. ANALYTICAL METHODS USED IN THE GEOGRAPHICAL COMPARISONS The statistical analyses of both the metric and the non- metric data included two approaches. First, multivariate statistical tests were used to determine whether the mea- sured traits were significantly different among the three geographical samples. Second, discriminant function anal- ysis was used to determine how well collection location could be predicted from observed characters using a dis- criminant function which maximizes the difference among area samples. Metric Analyses A multivariate analysis of variance (MANOVA, BMDP4V, Dixon et al., 1990) was used to test whether the areas Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins ■ 3 Table 1. Statistics for skull measurements (mm) of geographical series of Delphinus delphis. Sample sizes in parentheses. See Appendix 2 for explanation of acronyms. Black Sea Eastern tropical Pacific Western North Atlantic Measurement Range Mean SD Range Mean SD Range Mean SD 1. CBLN 374-420 (36) 394.5 11.09 397-460 (73) 425.8 14.41 409-478 (68) 440.9 15.75 2. ROST 218-258 (36) 237.4 9.38 233-286 (72) 256.5 11.33 240-299 (69) 270.8 11.91 3. ROSW 75-87 (36) 80.9 2.87 82-103 (73) 92.3 4.37 81-102 (69) 89.9 4.54 4. ROSQ 49-59 (36) 53.8 2.57 51-68 (73) 60.5 3.43 53-70 (69) 61.9 3.53 5. ROSH 42-52 (36) 46.1 2.70 45-58 (73) 51.4 2.96 44-62 (69) 54.8 3.59 6. ROS3 29-42 (36) 33.9 2.66 33-44 (73) 38.3 2.40 32-47 (68) 40.0 3.79 7. PMXH 19-25 (36) 21.8 1.57 21-30 (73) 25.3 1.85 20-31 (69) 25.0 1.92 8. TIPN 264-305 (36) 283.2 9.64 281-339 (73) 307.8 13.11 279-345 (69) 317.0 14.30 9. TIPP 257-296 (23) 273.9 10.40 273-332 (72) 301.3 12.34 279-343 (67) 312.4 13.53 10. PREO 142-162 (37) 152.3 4.85 152-186 (73) 167.3 6.24 157-185 (69) 172.1 6.56 11. POST 164-182 (37) 172.7 4.67 173-203 (72) 187.2 6.45 177-204 (69) 191.5 6.49 12. ZYGO 161-179 (37) 170.6 4.44 171-200 (73) 184.3 6.41 173-202 (70) 189.1 6.71 13. EXTN 40-52 (37) 45.2 2.25 43-53 (73) 47.5 1.96 43-54 (70) 48.3 2.57 14. PMXM 62-75 (37) 67.8 2.64 68-82 (73) 74.2 2.74 64-84 (69) 75.2 3.75 15. PARI 128-150 (37) 140.1 4.86 139-165 (73) 152.1 5.09 143-165 (70) 153.7 5.09 16. BRNH 92-109 (37) 101.1 3.47 97-113(73) 104.6 3.67 104-124 (70) 113.0 3.83 17. BRNL 100-114(37) 107.4 3.58 105-124 (72) 115.1 3.67 110-132 (69) 119.9 4.24 18. FOSL 62-73 (37) 67.0 2.71 58-76 73) 65.3 3.96 61-84 (70) 71.0 4.41 19. FOSH 45-57 (37) 50.8 3.02 38-60 (73) 48.3 3.94 48-64 (70) 55.2 3.46 20. ORB! 43-54 (37) 48.0 2.24 47-59 (73) 51.7 2.16 46-58 (69) 50.9 2.24 21. ANTO 34-45 (37) 38.8 2.61 38-53 (73) 43.9 2.70 39-54 (69) 47.2 3.22 22. INTN 46-54 (37) 49.5 2.45 47-59 (73) 52.6 2.71 45-65 (70) 56.0 3.69 23. UPRO 179-216(34) 196.8 8.48 192-248 (73) 218.4 10.52 200-259 (69) 228.4 11.10 24. UPTL 41-48 (31) 44.7 1.97 43-54 (71) 49.0 2.47 42-53 (64) 48.3 2.26 25. UPTR 41-51 (33) 45.1 2.30 43-54 (72) 49.1 2.19 44-53 (64) 48.3 2.26 26. LOTL 43-51 (32) 46.3 2.02 43-54 (70) 47.9 2.50 44-52 (64) 47.1 2.11 27. LOTR 43-50 (32) 46.3 2.04 42-53 (71) 47.9 2.57 43-53 (64) 47.1 2.20 28. LORO 180-210(33) 196.9 7.89 190-230 (72) 214.3 8.91 195-255 (67) 225.1 11.71 29. RAML 307-352 (33) 334.3 10.47 334-391 (72) 360.5 12.63 350-411 (67) 377.4 14.08 30. RAMH 59-67 (33) 61.6 1.92 59-70 (72) 64.2 2.26 62-75 (68) 68.5 2.88 31. TOOW 2. 6-3.4 (29) 3.13 0.156 2.6-3.4 (64) 2.98 0.173 3.0-3. 9 (47) 3.40 0.215 differed in the metric characters. Stepwise discriminant analysis (BMDP7M, Dixon et al., 1990) was used to iden- tify metric characters contributing to the discrimination of source area. In this analysis, variables are entered one at a time, based on their ability to improve on the sepa- ration of the sample means (estimated as a linear com- bination of variables). Variables are added until the im- provement of separation based on an F-ratio test falls below an arbitrary value (in this case, F < 4.0). (Note: We view this approach as empirical and do not ascribe statistical significance to variables that it identifies.) Jack- knife methods are used to estimate the number of cases (skulls) that are correctly identified; i.e., each specimen is classified based on a discriminant function that is com- puted excluding that specimen. Three specimens (two from the North Atlantic and one from the Black Sea) were deleted from the metric analyses because of a large number of missing values, leaving a total sample of 178. Missing values for the re- maining specimens were estimated using linear regression of the variable in question on the other variables (Dixon et al., 1990). Non-Metric Analyses Non-metric data violate many of the assumptions of stan- dard multivariate tests designed for metric data. There- fore, we used an index of similarity F which tests pairwise group (in this case, area) differences in multivariate non- metric characters (Zhivotovskiy, 1982): where r, = + Vprf2 + • • • + Vm~; T„ = index of similarity for n characters; r, = index of similarity for the zth character; p,, p2, . . ., pm = propor- tion of states 1,2,..., m, in one sample; qx, q2 , . . ., qm = proportion in other sample; and m, = number of states for character i. The statistical significance of the above similarity index 4 ■ Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins Table 2. Statistics for stepwise discriminant analysis of skulls of Delphinus delphis from three areas based on metric characters.3 See Appendix 2 for explanation of acronyms. Character F-value to enter Order of entry Coefficients I Ii 16. BRNH 154.69 1 0.47163 0.02491 3. ROSW 82.90 2 0.56276 -0.26132 31. TOOW 29.13 3 -0.41130 0.20190 17. BRNL 21.80 4 -0.20619 -0.36730 15. PARI 14.39 5 0.55171 -0.47189 19. FOSH 28.90 6 -0.51689 0.39200 2. ROST 11.30 7 -2.08716 0.04097 8. TIPN 14.22 8 1.65711 0.59925 10. PREO 8.61 9 -0.28245 -0.48738 23. UPRO 7.47 10 0.36557 -0.80257 22. INTN 6.61 11 -0.28309 0.29144 a. For discriminant function; standardized by pooled within-group variances. Table 4. Jackknife classifications for stepwise discrim- inant functions based on (a) 11 metric characters, (b) 8 non-metric characters, and (c) a combination of 7 metric and 2 non-metric characters. ETP = eastern tropical Pa- cific; WNA = western North Atlantic. True group Predicted group Black Sea WNA ETP Total Percent- age correct (a) Black Sea 36 0 0 36 100.0 WNA 0 67 2 69 97.1 ETP 1 1 71 73 97.3 Total 37 68 73 178 97.8 (b) Black Sea 28 0 9 37 75.7 WNA 4 61 4 69 88.4 ETP 9 5 57 71 80.3 Total 41 66 70 177 82.5 (c) Black Sea 36 0 0 36 100.0 WNA 3 62 2 67 92.5 ETP 3 2 66 71 93.0 Total 42 64 68 174 94.3 was tested using the method of Zhivotovskiy (1982). Giv- en the following expression, X = BN,N2 / _ y m+njv ti r, 5 where N, = number of individuals from group i and Z0 = number of cases with zero observed frequency in one group but not in the other, it can be shown that X is distributed approximately as x2 with degrees of freedom equal to 2”=i (m< — 1) (Yablokov et al., 1983). This was tested using standard tables. We also developed a simple bootstrap approach to test whether the non-metric traits differed among all areas simultaneously (analogous to an ANOVA test). The boot- strap test statistic is based on the mean number of dif- ferences in character states for all pairwise specimen com- parisons. The statistic used is the ratio, D, of the mean number of differences between groups to the mean within groups. Based on an assumption of no relationship be- tween character states and area, the expected value of this ratio is 1.0. The expected distribution of the ratio given this null hypothesis was computed by Monte Carlo sampling. Group (area) membership was randomly as- signed to each specimen (sampling without replacement and maintaining the observed sample sizes) and D was calculated. This bootstrap sampling procedure was re- peated 1000 times and the probability distribution of D estimated. Statistical inference was based on a comparison Table 3. Classification functions3 for stepwise discriminant analysis of skulls of Delphinus delphis from three areas based on metric characters. See Appendix 2 for explanation of acronyms. ETP = eastern tropical Pacific; WNA = western North Atlantic. Character Black Sea ETP WNA 2. ROST 1.62843 1.23656 2.17601 3. ROSW 0.01864 0.56331 -0.13070 8. TIPN -0.35333 -0.29916 -0.92628 10. PREO 1.34214 1.59824 1.80625 15. PARI 2.07914 2.70658 2.13252 16. BRNH 3.26663 3.03982 3.66798 17. BRNL 4.32538 4.63624 4.87718 19. FOSH 0.60288 -0.16514 0.58942 22. INTN -0.08477 -1.68253 -1.19496 23. UPRO -0.71854 -0.30765 -0.50764 31. TOOW 54.83513 45.05722 57.18664 Constant -793.27875 -890.32617 -983.51996 a. Used with original measurements. Add products of measurement and corresponding function values to constant; classify to area depending on which results in highest value for classification function. Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins ■ 5 B- Black Sea W- Western North Atlantic E-Eastern Tropical Pacific •- Group Mean FIRST CANONICAL VARIABLE Figure 2. Results of discriminant analysis of geographical samples of skulls of Delphinus delphis based on metric characters. of the observed value of D to its expected distribution given the hypothesis of no difference between areas. A transformation of the non-metric data was required before they could be analyzed with standard discriminant analysis. Of the 26 non-metric characters, 21 had three observed states (coded 0, 1, or 2) and 5 characters had two observed states (coded 0 or 1). Because these coded values are arbitrary, they could not be used directly in a linear model such as discriminant analysis. The non-metric variables were therefore converted to binary variables. A character that can assume only one of two possible states was represented by one binary variable. A character that could assume one of three possible states was coded as two binary variables, with values 0/0, 0/1, and 1 /0. The original 26 non-metric characters were thus coverted into 47 binary variables. A stepwise discriminant analysis (BMDP7M, Dixon et al., 1990) was used to identify which binary (or dummy) variables added to the discrimination of area. (It is important to note that when a character is coded as two binary variables, only one of these might contribute to the discrimination.) As with the metric data, the F-ratio for inclusion was set at 4.0 and classification of specimens to area was evaluated with jackknifed clas- sification. Four specimens (two from the eastern Pacific and two from the Atlantic) were eliminated from the non-metric analyses because of missing values, yielding a total sample size of 177. Combined Metric and Non-Metric Analyses We also applied discriminant function analysis to a com- bination of metric and non-metric characters. Only those metric characters or non-metric binary variables which we found to contribute significantly to either the metric or the non-metric discriminant functions were included. Again, we used a stepwise approach to determine which of the combined set of characters contributed most to discrimination of the samples. For this analysis we deleted the three specimens deleted for the metric analysis and the four deleted for the non-metric analysis, leaving a sample of 174. RESULTS STATISTICAL TESTS OF DIFFERENCES AMONG THE SAMPLES The samples from the three areas differed sig- nificantly in both metric and non-metric characters. Descriptive statistics for the metric characters are given by area in Table 1. The differences among the groups for the metric characters were large (MANOVA, Wilks’ lambda likelihood ratio statis- tic = 0.0288, p < 0.0001). For the non-metric char- acters, the similarity indices were 0.95 (Black Sea vs. western North Atlantic), 0.97 (western North Atlantic vs. eastern tropical Pacific), and 0.97 (Black Sea vs. eastern tropical Pacific); all these pairwise differences are very significant (chi-square test, p < 0.001). For the bootstrap test, the mean number of within-area pairwise differences was 5.51 characters (i.e., two specimens chosen randomly from one area were different in an average of 5.51 out of the 26 non-metric characters). The mean number of be- tween-area pairwise differences was 6.31. The ratio of within- to between-area differences thus was 6 ■ Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins Figure 3. Distribution of posterior probability of cor- rect classification for specimens correctly classified (hatched) and incorrectly classified (shaded), for metric and non-metric discriminant analysis of geographical sam- ples of Delphinus delphis. 0.874. A value this small was never observed in 1000 bootstrap samples (hence, p < 0.001). DISCRIMINANT ANALYSES The metric characters had higher discriminating power than the non-metric characters, but both were efficient. In the metric analyses, 1 1 characters added to the discrimination of sample origin (Ta- bles 2 and 3). The resulting first and second ca- nonical variables show excellent separation of sam- ples by area of origin (Fig. 2). The jackknife method gives an estimated 97.8% correct classification based solely on these 11 characters (Table 4, Fig. 3). Using non-metric characters, only eight binary variables added to the discrimination of sample or- igin. These eight corresponded to eight different non-metric characters (Table 5). The first two ca- nonical variables showed good separation of the western North Atlantic sample from the other two but considerable overlap between the Black Sea and Table 5. Statistics for stepwise discriminant analysis of skulls of Delphinus delphis from three areas based on non-metric characters (numbers in parentheses refer to character numbers in Appendix 3), including standard- ized (by pooled within variances) coefficients for canon- ical variables. Step Binary vari- able F-value Coefficients to enter I II 1 9(4) 220.24 -0.31316 0.02323 2 29(15) 16.72 0.97293 -0.26376 3 31 (16) 8.90 -0.06172 0.19337 4 10(5) 6.72 0.10358 -0.14793 5 25(13) 5.07 0.02593 0.18923 6 48 (26) 4.72 0.02346 0.37229 7 18 (9) 4.65 0.00641 0.24421 8 23 (12) 4.32 0.00000 0.00000 Constant: 1.55151 -1.09187 eastern tropical Pacific samples (Fig. 4). This was also reflected in the jackknife classification matrix (Table 4), in which 18 of 31 mis-classifications oc- curred between these two areas. The average cor- rect classification was 82.5%. The dispersion of pos- terior probabilities (Fig. 3) was also very much greater than for the metric analysis. When the 1 1 metric characters from the metric analysis and the 8 non-metric characters were com- bined in a third stepwise discriminant analysis, 7 metric and 2 non-metric characters contributed to discrimination of sample origin (Table 6). The jack- knife estimate of correct classification was 94.3% (Table 4). In the first canonical plane, the geographic areas are very distinct (Fig. 5). DISCUSSION RELATIVE POWER OF THE TWO TYPES OF CHARACTERS While both the metric and non-metric characters were effective in discrimination, the two types of characters had different resolving power for differ- ent pairs of the three geographical samples. In the combined analysis, a non-metric character was the first variable entered and was very powerful (as in- dicated by the initial F-value to enter; Table 6) in separating the western North Atlantic sample from the combined Black Sea and eastern tropical Pacific samples, and the additional non-metric variable that entered the analysis also contributed (almost ex- clusively) to this separation. The seven metric char- acters that entered the analysis mainly contributed to separation of the Black Sea and eastern tropical Pacific samples. If the non-metric characters are indeed neutral, this may indicate that the western North Atlantic population is relatively the most isolated, with the least genetic exchange with the Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins ■ 7 5 B= Black Sea, W= Western North Atlantic, E= Eastern Tropical Pacific _Q) JD co co > ”c0 o 'c o c co O -O c o o CD CO 4 3 2 1 0 -1 -2 w w w wO^w W w w WWW w w w'W .Ayv EW Wwvv M/Ww B w BB B B B B B B B B B c pB Wn R B E r E B F ° ® W® W ^ " 1%-'B Ee%%B p -B E E E W ►' E E| E.E W_ / w E E = B E JE ^ /e Efcf E E? K E E -3 +- -4 -2 0 First Canonical Variable Figure 4. Results of discriminant analysis of geographical samples of skulls of Delphinus delphis based on all 26 non- metric characters. other populations. The converse is that genetic ex- change is or has been greatest between the Black Sea and the eastern tropical Pacific. While possible, this seems unlikely. An alternative explanation is that the non-metric characters are not really neutral Table 6. Statistics for stepwise discriminant analysis of skulls of Delphinus delphis from three areas based on combined metric variables and non-metric characters, including standardized (by pooled within variances) co- efficients for canonical variables. Non-metric characters listed in Appendix 3; metric acronyms in Appendix 2. F- value _ Coefficients Step Character to enter I II 1 Non-metric 4 233.41 -0.73182 -0.21326 2 ROST 95.55 0.30877 -0.04114 3 BRNH 31.19 0.22978 -0.63139 4 PARI 19.77 0.43258 -0.75298 5 FOSH 31.99 -0.40402 0.05757 6 INTN 15.14 -0.25809 -0.29547 7 TOOW 10.58 -0.34667 0.56602 8 BRNL 9.10 -0.42534 0.22506 9 Non-metric 5 6.57 -0.23313 0.29519 Constant: 19.54346 26.24044 or are somehow linked chromosomally to metric characters directly subject to selection pressure (i.e., the results for both character sets reflect primarily differential selection pressures rather than genetic isolation) and that the closer morphological dis- tance between the Black Sea and eastern tropical Pacific samples in both character suites represents adaptive convergence. If this is the case, then the metric characters must be considered to be rela- tively more useful, as their use resulted in fewer errors of classification. The differential power to resolve various pairs of geographical samples could also relate to the fact that the western North Atlantic sample was drawn from a much larger area and was more heteroge- neous than the other two samples, in both metric (Fig. 6) and non-metric (Fig. 4) characters. These questions could usefully be addressed by a comparison of metric and non-metric differences between two parapatric populations that are dif- ferentiated morphologically but between which some gene flow occurs (as evidenced by intergra- dation or hybridization). Ideally, this would be cou- pled with a parallel analysis of DNA (e.g., sequenc- ing or restriction-enzyme analysis of highly variable regions of the mitochondrial DNA molecule) to estimate relative gene flow between the popula- 8 ■ Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins B - Black Sea W - Western North Atlantic E - Eastern Tropical Pacific • - Group Mean FIRST CANONICAL VARIABLE Figure 5. Results of discriminant analysis of geographical samples of skulls of Delphinus delphis based on combined metric and non-metric characters. tions. The question of the relative coherence of the Atlantic sample should be addressed by analyses of samples from more limited areas as they become available. REPEATABILITY The two scorers had considerable difficulty in main- taining repeatability in scoring the non-metric char- acters. Despite the character states ostensibly being objectively defined, there proved to be some diffi- culty and subjectivity in consistently discriminating between them for some skulls, because of subtle variation among individual skulls in size, shape, and placement of the features, and the scorers found that there was a constant need to cross-check and consult with each other to prevent character drift over a period of even a few days. We believe that there is a definite possibility that significant drift may have occurred between the Moscow and Washington, D.C., sessions. To address this prob- lem, ideally all specimens to be included in a non- metric study should be physically pooled and scored in random order in a short period of time in one location. The comparability of data collected by different investigators or by one investigator at widely separated times must be questioned. Of course, poor comparability of data collected by different workers is not a problem only of non- metric characters; even meristic characters can show Figure 6. Coefficients of variation (CV) for 31 cranial characters for three geographical samples of Delphinus delphis. The characters on the horizontal axis are in order of decreasing CV; the order differs among the three series. The areas under the curves are proportional to the total relative variation for the characters. Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins ■ 9 such variation (Perrin et al., 1987). Of special con- cern here, however, is possible drift for single work- ers over a relatively short period of time. Because of their apparent lesser resolving power and because of the difficulties concerning repeat- ability, we recommend that non-metric characters be used in dolphin population studies only in com- bination with metric characters. If non-metric char- acters are used, special attention should be paid to controlling conditions to maximize comparability of data sets. ACKNOWLEDGMENTS The research was carried out as part of the Marine Mam- mal Project under the U.S.-U.S.S.R. Environmental Agree- ment of 1972; our thanks go to V.A. Zemskiy, R.V. Mil- ler, N.A. Popov, S.G. Kohl, W. Silverman, and others who have made the Marine Mammal Project work. The specimens were collected by scores of field biologists and technicians aboard commercial fishing vessels, at landing sites, and from beaches in the case of strandings; this type of research would be impossible without the efforts of such people and cooperation of members of the fishing community and the general public, and we owe them very much. J.G. Mead kindly allowed us to work with the specimens in the U.S. National Museum, and C.W. Potter helped us extract the collection data for them. O.L. Ros- solino allowed access to the specimens in the Zoological Museum of Moscow State University, and I.Ya. Pavlinov assisted us there. G.J. Walker took part in initial discus- sions of the proposed work. J.V. Kashiwada and J.W. Gilpatrick, Jr., assisted with entry and editing of the data. L.C. Prescott and C. Ratcliffe assisted in preparation of the manuscript; R.M. Allen prepared the figures. D.P. DeMaster and I. Barrett read the manuscript and advised us on how it could be improved. We thank all of these people and others who helped us. LITERATURE CITED Berry, R.J. 1969. Non-metric skull variation in two Scot- tish colonies of the Gray seal. Journal of Zoology ( London ) 157:11-18. Dailey, M.D., and W.F. Perrin. 1973. Helminth parasites of porpoises of the genus Stenella in the eastern tropical Pacific, with descriptions of two new spe- cies: Mastigonema stenellae gen. et sp. n. (Nema- toda: Spiruroidea) and Zalophotrema pacificum sp. n. (Trematoda: Digenea). Fishery Bulletin, U.S. 71: 455-471. Dixon, W.J., M.B. Brown, L. Engelman, and R.I. Jennrich (eds.). 1990. BMDP statistical software manual. Two vol. Berkeley, Calif.: Univ. Calif. Press, 1386 PP- Fedoseev, G.A. 1984. Use of nonmetrical characters of skulls of Bering Sea seals in a study of the phenotypic structure of their populations. In Soviet- American cooperative research on marine mammals. Volume l — Pinnipeds, ed. F.H. Fay and G.A. Fedoseev. NOAA Technical Report NMFS 12, 99 pp. Hartman, S.E. 1980. Geographic variation analysis of Dipodomys ordii using nonmetric cranial traits. Journal of Mammalogy 61:436-448. Kinze, C.C. 1985. Intraspecific variation in Baltic and North Sea harbour porpoises ( Phocoena phocoena (L., 1758)). V idenskahelige Meddelelser fra Dansk Naturhistorisk Forening 146:63-74. Kinze, C.C. 1990. Non-metric analysis of harbour por- poises (Phocoena phocoena) from the North and Baltic Seas: Implications for stock identity. Inter- national Whaling Commission meeting document SC/42/SM35, 23 pp. (available from IWC). Perrin, W.F. 1975. Variation of spotted and spinner porpoise (genus Stenella) in the eastern tropical Pa- cific and Hawaii. Bulletin of the Scripps Institution of Oceanography 21:1-206. Perrin, W.F., E.D. Mitchell, J.G. Mead, D.K. Caldwell, M.C. Caldwell, P.J.H. van Bree, and W.H. Dawbin. 1987. Revision of the spotted dolphins, Stenella spp. Marine Mammal Science 3:99-170. Perrin, W.F., M.D. Scott, G.J. Walker, and V.L. Cass. 1985. Review of geographical stocks of tropical dolphins ( Stenella spp. and Delphinus delphis) in the eastern Pacific. NOAA Technical Report NMFS 28, 28 pp. Perrin, W.F., A.V. Yablokov, and V.L. Cass. 1982. Pre- liminary report on the use of non-metrical skull char- acters to discriminate populations of pelagic dol- phins. Southwest Fisheries Center Administrative Report LJ-82-14, 32 pp. Perrin, W.F., A.V. Yablokov, J. Barlow, and M.V. Mina. 1988. The use of non-metrical skull characters for analyzing geographical variation in the common dol- phin, Delphinus delphis. Part I: Selection of char- acters. Southwest Fisheries Center Administrative Report LJ-88-20, 24 pp. Sjovold, T. 1973. The occurrence of minor non-metrical variants in the skeleton and their quantitative treat- ment for population comparisons. Homo 24:204- 233. Tomilin, A.G. 1957. Kitoobraznye [Cetaceans], vol. 9. In Zveri SSSR i prilezhanshchikh stran. Zveri vos- tochnoi Evropy i severnoi Azii [Mammals of the U.S.S.R. and adjacent countries. Mammals of east- ern Europe and northern Asia], ed. V.G. Heptner. Moscow: Akad. Nauk SSSR, 756 pp. (Transl. by O. Ronen, Cetaceans. In Mammals of the U.S.S.R. and adjacent countries. Mammals of eastern Europe and adjacent countries [s/c]. Israel Program for Scientific Translations, Jerusalem, 1967, 717 pp.) Urbach, V.Y. 1964. Biometric methods [in Russian], Moscow: Nauka, 416 pp. Yablokov, A.V., W.F. Perrin, and M.V. Mina. 1983. Otsenka feneticheskikh otnosheniy gruppirovok del’finov na osnove analiza nemetricheskikh variatsiy cherepa [Evaluation of phenetic relations among groups of dolphins using analysis of non-metrical cranial varation]. Zoologicheskiy Zhurnal 62:1887- 1896. Zhivotovskiy, L.A. 1982. Pokazateli populyatsponnoy izmenchivosti po polimorfnym priznakam [Indices of population variation in polymorphic characters]. In Fenetika populyatsiy, ed. A.V. Yablokov, 38-44. Moscow: Nauka. Received 2 March 1994; accepted 23 March 1994. APPENDIX 1 Specimens used in analyses of geographical samples of Delphinus delphis are listed below. Black Sea specimens are in the collection of the Zoological Museum, Moscow State University, Moscow (MSUZM); eastern tropical Pa- cific and western North Atlantic specimens are in the U.S. 10 ■ Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins National Museum of Natural History, Washington, D.C. (USNM). BLACK SEA Batumi, 4 (MSUZM S-5411, S-80352-53, 62); Yalta, 13 (MSUZM S-50804-05, S-80335-36, S-80339-40, S-80363, S-80372-73, S-80376, S-80379, S-80534); Novorossiysk, 14 (MSUZM S-80334, 345, 349, 355, 359, 367, 369, 374- 75, 378, 383, 385); unknown localities on the Soviet coast of the Black Sea, 6 (MSUZM S-80342, 347, 360, 377, 387, 389). EASTERN TROPICAL PACIFIC 9°N, 84°10'W, 14 (USNM 487766-67, 69-70, 73-82); 8°25'N, 84°W, 19 (USNM 487807, 09-12, 15, 17-26, 28- 30); 9°18'N, 84°46'W, 1 (USNM 487480); 8°08'N, 87°54'W, 5 (USNM 5000060-61, 65-66); 7°32'N, 83°54'W, 18 (USNM 5000069-80, 82-85, 87-88); 7°25'N, 84°02'W, 1 (USNM 500092); 8°14'N, 83°15'W, 1 (USNM 500116); 7°45'N, 90°50'W, 12 (USNM 500315-16, 20-23, 34-39); 8°40'N, 84°32'W, 2 (USNM 500340-41). WESTERN NORTH ATLANTIC Nova Scotia, 2 (USNM 300191-92); 43°32'N, 64°25'W, 1 (USNM 267968); 42°58'N, 63°43'W, 1 (USNM 267969); 40°11'N, 73°33'W, 2 (USNM 571202-03); 39°39'N, 71°54'W, 1 (USNM 20873); 39°20'N, 72°44'W, 7 (USNM 550808-15); 39°N, 73°35'W, 1 (USNM 550781); 38°48'N, 73°05'W, 1 (USNM 550779); 38°54'N, 73°01'W, 1 (USNM 550870); 38°56'N, 72°53'W, 1 (USNM 550750); 38°38'N, 73° 14'W, 2 (USNM 550774-75); 38°38'N, 73°14'W, 1 (USNM 550775); 38°35'N, 73°17'W, 1 (USNM 550550); 38°14'N, 73°44'W, 1 (USNM 550862); 38°1 1'N, 73°46'W, 1 (USNM 550780); 37°57'N, 74°39'W, 1 (USNM 550927); 37°47'N, 74°52'W, 1 (USNM 550776); 37°44'N, 74°44'W, 1 (USNM 550987); 37°27'N, 74°28'W, 1 (USNM 504654); 37°18'N, 75°01'W, 1 (USNM 550885); Massachusetts, 19 (USNM 21525-5, 5500041, 192, 201, 205-07, 211, 450, 470, 4756, 478, 841, 846-47, 849, 571232); New York, 5 (USNM 7063, 504262-63, 504421, 550755); New Jer- sey, 1 (USNM 504407); North Carolina, 8 (USNM 4115, 21917-8, 305126, 504219, 504406, 504877-78); Virginia, 7 (USNM 22528, 22560, 395923, 504323, 504958, 550065, 550819); Florida, 1 (USNM 267478); 46°55'N, 27°51'W, 1 (USNM 504631). APPENDIX 2 Metric skull characters used in the analysis of samples of Delphinus delphis from three areas, with acronyms used in tables, are listed below. Measurements are illustrated in Perrin (1975). Tooth width (no. 31) was measured in millimeters and tenths; all others in millimeters. Character Acronym 1. Condylobasal length CBLN 2. Length of rostrum ROST 3. Rostrum width at base ROSW 4. Rostrum width at quarter-length ROSQ 5. Rostrum width at half-length ROSH 6. Rostrum width at three-quarters-length ROS3 7. Width of premaxillaries at half-length PMXH 8. Tip of rostrum to external nares TIPN 9. Tip of rostrum to internal nares TIPP Character Acronym 10. Preorbital width PREO 11. Postorbital width POST 12. Zygomatic width ZYGO 13. Width of external nares EXTN 14. Maximum width across premaxillaries PMXM 15. Parietal width PARI 16. Braincase height BRNH 17. Braincase length BRNL 18. Length of temporal fossa FOSL 19. Height of temporal fossa FOSH 20. Length of orbit ORBI 21. Length of antorbital process ANTO 22. Width of internal nares INTN 23. Length of upper toothrow UPRO 24. Number of upper left teeth UPTL 25. Number of upper right teeth UPTR 26. Number of lower left teeth LOTL 27. Number of lower right teeth LOTR 28. Length of lower toothrow LORO 29. Length of ramus RAML 30. Height of ramus RAMH 31. Width of tooth at alveolus TOOW APPENDIX 3 Non-metric cranial characters used in the analysis of geo- graphical samples of Delphinus delphis are as follows. IN DORSAL VIEW 1. Position of anterior edge of mesial infraorbital fo- ramen (a in App. Fig. 1) relative to the level of the anteriormost projection of the antorbital process, on left side. Note: The level of the antorbital process must be determined accurately; this is best done with a flexible ruler or other straight-edge. Codes: 1 = ahead, 2 = behind, 3 = at level, or uncertain. 2. Position of posterior edge of lateral infraorbital fo- ramen (first, if more than one on each side) (b in App. Fig. 1) relative to posterior edge of mesial infraorbital foramen (a in App. Fig. 1), on left side. Codes: 1 = ahead, 2 = behind, 3 = at the level, or uncertain. 3. Number of vertical perforations (visible in vertical view) of facial area of maxillary by foramina, on right side. Codes: 1 - 1, 2 = 2, . . ., 9 = 0. 4. Dorsal projection of mesethmoid between premax- illaries near base of rostrum (c in App. Fig. 1). Codes: 1 = below surface of premaxillaries, 2 = above surface, 3 = at surface, or uncertain. 5. Contact or near-contact ( < 1 - mm gap) between pre- maxillaries (or exposed maxillaries) in prenarial tri- angle (c in App. Fig. 1). Caution: The skull must be rotated to allow detection of overlapping bones. Codes: 1 = present, 2 = absent, 3 = uncertain. 6. Contact between premaxillary and nasal (no inter- vening bones), on left side (e in App. Fig. 1). Codes: 1 = present, 2 = absent, 3 = uncertain. Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins ■ 1 1 Appendix Figure 1. Dorsal view of dolphin skull, showing features mentioned in descriptions of non-metric characters. Codes: 1 = beyond (posterior to) anterior edge of left nasal, 2 = not beyond (anterior to) anterior edge of left nasal, 3 = at the level of anterior edge of left nasal or uncertain. 7. Same on right side. 8. Relative extension of ascending process of left pre- maxillary. Note: Do not confuse with anterior ex- tension of nasal between premaxillary and maxillary. 1 2 ■ Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins Appendix Figure 2. Occipital view of dolphin skull, showing features mentioned in descriptions of non-metric characters. IN OCCIPITAL VIEW (Resting Right-Side-Up on Table) 9. Shape of foramen magnus (App. Fig. 2). Codes: 1 = piriform, 2 = smooth oval/circle, 3 = uncertain. 10. Number of fenestrations in occipital (near foramen magnum) and exoccipital (a in App. Fig. 2), on left side. Codes: 1 = 1, 2 = 2, 3 = 3, . . 9 = 0. 11. Same on right side. 12. Groove between occipital condyles, in occipital pro- file (b in App. Fig. 2). Codes: 1 = clearly present, 2 = clearly absent, 3 = absent, or uncertain. IN VENTRAL VIEW 13. Small foramen between occipital condyles in bas- occipital (a in App. Fig. 3). Codes: 1 = present, 2 = absent, 3 = uncertain. 14. Mesial fenestrations in basoccipital (b in App. Fig. 3). Codes: 1 = present and clearly natural, 2 = absent, or clearly postmortem damage, 3 = present, but possibly postmortem damage (= uncertain). 15. Position of posterior end of vomer3 (c in App. Fig. 3) relative to line connecting mesial ends of sutures between alisphenoids and exoccipitals (d in App. Fig. 3). Codes: 1 = behind (ignoring small irregularities), 2 = in line or ahead, 3 = uncertain. 16. Lateral fenestration in vomer3 (e in App. Fig. 3), on left side. Codes: 1 = present, 2 — absent, 3 = uncertain. 17. Same on right side. 18. Mesial fenestration in vomer3 (f in App. Fig. 3). Codes: 1 = present, 2 = absent, 3 = uncertain. 19. One or more fossae between the mesial and posterior lacerate foramina (g in App. Fig. 3), on left side. Codes: 1 = present, 2 — absent, 3 = uncertain. 20. Position of interior opening of supraorbital foramen (h in App. Fig. 3), on left side. Codes: 1 = behind (under) frontal trabeculum (hid- a. Erroneously called basisphenoid in Yablokov et al. (1983) and Perrin et al. (1982, 1988). Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins ■ 13 Appendix Figure 3. V entral view of dolphin skull, showing features mentioned in descriptions of non-metric characters. 14 ■ Contributions in Science, Number 447 Perrin et al.: Comparing Cranial Characters in Dolphins den in perpendicular view), 2 — anterior to frontal trabeculum (visible), 3 = at trabeculum, or uncer- tain. 21. Position of anterior edge of large ventral foramen in maxillary (i in App. Fig. 3) relative to posterior edge of base of jugal (j in App. Fig. 3), on left side. Codes: 1 = anterior to edge of jugal, 2 = posterior to edge, 3 = at edge, or uncertain. 22. Same on right side. 23. Foramen-like fossa in internal roof of temporal fossa (k in App. Fig. 3; shown on right side), on left side. Codes: 1 = present, 2 = absent, 3 = uncertain. 24. Same on right side. 25. Configuration of optical hiatus (1 in App. Fig. 3), on left side. Codes: 1 = perforated in vertical view (>l-mm opening), 2 = not perforated, 3 = uncertain. 26. Same on right side. Contributions in Science, Number 447 Perrin et at.: Comparing Cranial Characters in Dolphins ■ 15 Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Number 448 6 October 1994 Contributions in Science Diadasia, Subgenus Dasiapis, in North America (Hymenoptera: Anthophoridae) Roy R. Snelling W5S Natural History Museum of Los Angeles County Serial Publications of THE Natural History Museum of Los Angeles County Scientific Publications Committee James L. Powell, Museum President Daniel M. Cohen, Committee Chairman Brian V. Brown Kenneth E. Campbell Kirk Fitzhugh Edward C. Wilson Robin A. Simpson, Managing Editor The scientific publications of the Natural History Mu- seum of Los Angeles County have been issued at irregular intervals in three major series; the issues in each series are numbered individually, and numbers run consecutively, regardless of the subject matter. • Contributions in Science, a miscellaneous series of tech- nical papers describing original research in the life and earth sciences. • Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth sci- ences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now published by the Museum in Contributions in Science. • Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through the Museum Book Shop. A catalog is available on request. The Museum also publishes Technical Reports, a mis- cellaneous series containing information relative to schol- arly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum’s Scientific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Tech- nical Reports may be obtained from the relevant Section of the Museum. Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Printed at Allen Press, Inc., Lawrence, Kansas ISSN 0459-8113 D i ad asia. Subgenus Dasiapis, in North America (Hymenoptera: Anthophoridae) Roy R. Snelling1 ABSTRACT. The status of the six species-group names applied to Diadasia, subgenus Dasiapis, in North America is reviewed and three species are recognized. D. olivacea (Cresson) is known only from Mexico. D. ochracea (Cockerell) (=D. blaisdelli Cockerell) occurs over much of the western United States and adjacent Mexico. D. tropicalis (Cockerell) ( =Melissodes spilognathus Cockerell and M. tenuicincta Cockerell, both N. SYN.) is common and widespread in Mexico but occurs in the United States (Arizona, Texas) and ranges south to Honduras and El Salvador; specimens tentatively assigned to D. tropicalis have been seen from Venezuela. The three species are separated by means of a key. Character state variation is briefly discussed and distribution records are cited. Male terminalia are illustrated for the three species, and the known distributions within Mexico and Central America are illustrated in a map. INTRODUCTION Dasiapis Cockerell, 1903a (type species: D. ochra- cea Cockerell, 1903a; monobasic) is a small sub- genus of Diadasia Patton, 1879, with representa- tives in both North and South America. To date, six species-group names have been proposed for the North American forms, but there has never been any attempt to clarify the relationships of the several names. While some synonymy has been pro- posed, it has been based on supposition, not upon examination of available material, including the rel- evant types. In an effort to clarify the status of the North American Dasiapis species, I have seen all the types and evaluated them in terms of observable character state variation in large numbers of mu- seum specimens. DISCUSSION The species of Diadasia in America north of Mex- ico were first reviewed by Timberlake (1941), but none of the species were treated in detail. The status of the subgenus Dasiapis was discussed by Mich- ener (1954), and its most important characteristics were set forth in a table to compare them with those of other subgenera of Diadasia. Dasiapis may be distinguished from other subgenera of Dia- dasia by the following combination of character- istics: clypeus, or at least base of mandible, yellow- marked; inner mesotarsal and both metatarsal claws of male rounded, the protarsal and outer mesotarsal claws acute; female protarsal claws acute, meso- and metatarsal claws rounded; male middle flagellar segments longer than broad; pterostigma moderate in size. 1. Entomology Section, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los An- geles, California 90007. Adlakha (1969) revised the Diadasia in America north of Mexico. All the species were described in detail, male terminalia were illustrated, geograph- ical distributions were cited, and known biologies were described. Adlakha’s key to the species was a slight modification of the earlier one by Timberlake (1941). Adlakha’s figures of the male terminalia se- riously misrepresent the distribution of pilosity. The one United States species of subgenus Dasiapis (D. ochracea) was stated to be a synonym of the Mex- ican D. olivacea, but without supporting evidence. Most recently, Hurd (in Krombein et al., 1979) listed D. ochracea as a presumably valid species but noted that the above synonymy had been proposed. In addition to the above names, four others apply to North American species of Dasiapis: D. tropi- calis Cockerell, 1918, from Mexico; D. blaisdelli Cockerell, 1924, from California; both Melissodes tenuicincta and M. spilognathus, described by Cockerell (1949), from Honduras were correctly assigned to Diadasia by LaBerge (1956). Of these, D. blaisdelli has been generally recognized as a syn- onym of D. ochracea (e.g., Michener in Muesebeck et al., 1951; Adlakha, 1969). After examining the available material I have concluded that there are three species of Dasiapis in North America. These may be separated by the following key. KEY TO NORTH AMERICAN DASIAPIS la. Male; scopa absent from metatibia and anten- na 13-segmented 2 b. Female; scopa present on metatibia and anten- na 12-segmented 4 2a. Head width 3. 5-3. 8 mm; inner eye margins more strongly convergent below, upper inter- orbital distance about 1.27 times lower; upper frons sharply roughened and slightly shining between well-defined punctures that extend to Contributions in Science, Number 448, pp. 1-8 Natural History Museum of Los Angeles County, 1994 margin of anterior ocellus olivacea (Cresson) b. Head width 2. 7-3. 3 mm; inner eye margins more weakly convergent below, upper inter- orbital distance about 1.2 times lower; upper frons shiny, weakly tessellate or smooth be- tween weak punctures that do not usually ex- tend to anterior ocellus 3 3a. Median one-half or more of disc of meso- scutum either impunctate or with sparse punc- tures that are little greater in diameter than hairs arising from them and are conspicuously finer than punctures near parapside; femora black ochracea (Cockerell) b. Center of mesoscutum closely punctate (except occasionally along midline), punctures about equal to those near parapside; femora reddish tropicalis (Cockerell) 4a. Head width 3. 8-4.1 mm; upper frons sharply roughened and slightly shiny between deep punctures, with fine, sharply defined punctures extending to margin of anterior ocellus; clypeal disc, seen from below, depressed or flattened across middle one-third or more, punctures of disc separated by 0.66 times puncture diameter or less olivacea (Cresson) b. Head width 2. 8-3. 6 mm; upper frons moder- ately shiny between obscure punctures, none of which extend to margin of anterior ocellus; clypeal disc, seen from below, regularly convex (slightly depressed along midline in some spec- imens), punctation various, but usually most punctures separated by more than puncture di- ameter 5 5a. Mesoscutum dull, center of disc without ob- vious punctures and posterolateral area with punctures (when present) largely effaced by dense tessellation; upper frons (within 1 .5 times OD from anterior ocellus) shiny with at most scattered fine, obscure punctures and often with no punctures; femora and tibiae dark brownish ochracea (Cockerell) b. Center of mesoscutum dull to slightly shiny and densely punctate, punctures not apprecia- bly finer than those near parapside; postero- lateral area of mesoscutum subcontiguously punctate; upper frons often with conspicuous punctures that are greater in diameter than hairs arising from them; femora and tibiae usually reddish but may be light red-brown tropicalis (Cockerell) Diadasia ( Dasiapis ) ochracea (Cockerell) Figures 3, 8-9, 11 Dasiapis ochracea Cockerell, 1903a:450; 6. Cock- erell, 1906:74, 100. Melissodes olivacea: Cockerell, 1903b:77. MIS- IDENTIFICATION. Diadasia hlaisdelli Cockerell, 1924:54; 9. Timber- lake, 1941:11. Diadasia ochracea: Timberlake, 1941:2, 6; 9 6. Michener, in Muesebeck, Krombein, and Townes, 1951: 1220. Hurd, in Krombein, Hurd, Smith, and Burks, 1979:2120. Diadasia olivacea: Linsley and MacSwain, 1957: 215-217 (biology). Adlakha, 1969:55, 60, 126- 131. Eickwort, Eickwort, and Linsley, 1977:1- 17 (biology). Rozen, Eickwort, and Eickwort, 1978:1-24 (parasite). MISIDENTIFICATION. The taxonomic status of D. ochracea has been confused since 1957, when Linsley and MacSwain quoted the statement made to them by P.H. Tim- berlake that he believed D. ochracea to be a syn- onym of D. olivacea, even though he had not seen the type of either name. Adlakha (1969) was un- equivocal about the matter: he treated the two as synonymous, but, again, there is no indication that he studied the types of either. I have examined all of the relevant North American types and con- cluded that D. olivacea and D. ochracea are distinct species, as is D. tropicalis from Mexico, and that D. hlaisdelli from California is a synonym of D. ochracea, as first suggested by Timberlake (1941). Michener (in Muesebeck et ah, 1951) included D. hlaisdelli as a synonym of D. ochracea. The several biological studies cited above all re- fer to this species, not to D. olivacea. Although the bulk of the floral visitations of D. ochracea involve species of Sphaeralcea (Malvaceae), there are rec- ords from other malvaceous genera also: Gossyp- ium, Malacothamnus, Malva, and Sida. Other plants visited, presumably for nectar, are the fol- lowing: ASTERACEAE: Baccharis glutinosa, Cir - sium sp., Gaillardia pulchella, Helenium hoopesii, Helianthus parviflora, Heterotheca subaxillaris, Hymenothrix sp., Verbesina encelioides, and Vi- guiera sp.; BORAGINACEAE: Heliotropium cu- rassavicum; FABACEAE: Medicago sativa and Phaseolus wrightii; LILIACEAE: Calochortus sp.; PAPAVERACEAE: Argemone sp.; POLYGONA- CEAE: Eriogonum sp. and Antigonon leptopus; PORTULACEAE: Portulaca sp.; ZYGOPHYL- LACEAE: Kallstroemia grandiflora; and CACTA- CEAE: Opuntia sp. Most of these records are cited in Adlakha (1969). The range of D. ochracea in the United States is dealt with in detail by Adlakha (1969); in summary, this species ranges from Texas west to California, thence north to the State of Washington; it occurs in southern Utah and Colorado. Although there are no records for western Kansas and Oklahoma, D. ochracea may occur in suitable habitats in both states. Since Adlakha did not treat the Mexican distribution in detail, the following records are pub- lished to supplement the United States records (see Fig. 11). MEXICO, Baja California: 1 9, Santo Tomas, 10 May 1938 (C.E. Norland; LACM); 1 9, 2 km NE Polvora, 13 July 1953 (R.R. Snelling; LACM), on Gossypium; 19 ,6 km SE Paredones, 2 July 1953 (R.R. Snelling; LACM), on Gossypium; 1 <3, 12 km SW Mexicali, 14 July 1953 (R.R. 2 ■ Contributions in Science, Number 448 Snelling: Diadasia, Subgenus Dasiapis Figures 1-3. Diadasia ( Dasiapis ) spp., male genital capsule, ventral view: 1, D. tropicalis; 2, D. olivacea; 3, D. ochracea. Scale line = 0.50 mm. Snelling; LACM), on Sida hederacea. Chihuahua: 12 99, 3 <3<3, Catarinas, 5800 ft., 25-26 July 1947 (Cazier, Gertsch, Michener, Schrammel; UCD); 1 9, 2 68, 42 mi SW Ca- margo, 4900 ft., 15 July 1947 (C.D. Michener; UCD); 1 <3, 25 mi SW Camargo, 14 July 1947 (C.D. Michener; UCD); 3 88, 20 mi SW Camargo, 13 July 1947 (C.D. Mich- ener; UCD), on Cevallia sinuata; 1 9, Agua Caliente, Santa Barbara Dist., 14 July 1947 (C.D. Michener; UCD); 2 99, 1 8, 16 mi SE Chihuahua, 11 July 1947 (Gertsch, Michener, Schrammel; UCD); 1 9, 65 mi N Chihuahua, 22 Aug. 1964 (R.C. Dickson; UCD); 1 9, 30 mi NW Chihuahua, 4900 ft., 27 July 1953 (Univ. Kans. Mex. Exped.; UKAN); 1 <3, 239 km S Ciudad Juarez, 1 July 1947 (C.D. Michener; UCD), on Sphaeralcea; 1 9, 5 mi N Escalon, 20 Sept. 1970 (G.E. and R.M. Bohart; BBSL); 1 6, 9 mi S Hidalgo de Parral, 22 July 1967 (R.C. Gardner and C.R. Kovacic; UCD); 7 <3<3, 29 mi E Hidalgo de Parral, 5000 ft., 21 July 1956 (H.A. Scullen; UCD); 1 9, 33 mi N, 38 mi E Hidalgo de Parral, 5000 ft., 21 June 1956 (H.A. Scullen; UCD); 1 8, 10 mi W Jimenez, 11 Sept. 1950 (R.F. Smith; UCD); 4 86, 10 mi N Jimenez, 21 Sept. 1970 (G.E. and R.M. Bohart; BBSL); 1 <3, 17 mi W Jimenez, 4600 ft., 26 July 1953 (Univ. Kans. Mex. Exped.; UKAN), on Grindelia sp.; 1 6, 21 mi N Jimenez, 9 Sept. 1973 (W.J. Hanson and B.A. Haws; BBSL); 1 9, La Cruz, 13 July 1947 (C.D. Michener; UCD), on Cevallia sp.; 1 9, 2 <3<3, Delicias, 13 July 1947 (C.D. Michener; UCD); 2 <3<3, 10 mi S Delicias, 13 July 1947 (C.D. Michener; UCD); 1 9, Salaices, 5200 ft., 21 Sept. Contributions in Science, Number 448 Snelling: Diadasia, Subgenus Dasiapis ■ 3 Figures 4-10. Diadasia ( Dasiapis ) spp.: 4-5, D. olivacea, male metasomal sternum 8, ventral and lateral views, respectively; 6-7, D. tropicalis, same; 8-9, D. ochracea, same; 10, D. olivacea, male sternum 7, ventral view. Scale line = 0.50 mm. 1947 (G.M. Bradt; UCD); 1 6, Samalayuca, 6 Aug. 1950 (R.F. Smith; UCD); 6 66, 15 km S Samalayuca, 1 Aug. 1974 (E.M. and J.L. Fisher; LACM); 1 9, 3 66, Santa Barbara, 6300 ft., 17-18 July 1947 (Cazier, Michener, Schrammel; UCD); 1 6, Villa Ahumada, 3700 ft., 28 July 1953 (Univ. Kans. Mex. Exped.; UKAN). Coahuila: 1 9, Guadalupe, 23 Aug. 1947 (C.D. Michener; UCD); 1 9, Paila, 3900 ft., 21 July 1947 (C.D. Michener; UCD); 2 66, 23 mi N Saltillo, 11 Aug. 1959 (L.A. Stange and A.S. Menke; UCD); 1 6, San Pedro de Colonias, 3700 ft., 20 Aug. 1947 (C.D. Michener; UCD). Durango: 1 9, Bermejillo, 1 Nov. 1965 (G.E. and A.S. Bohart; BBSL), on Sphaeralcea sp.; 1 9, same except 5 Oct. 1966 (BBSL); 1 6, 20 mi N Bermejillo, 4 Oct. 1966 (G.E. and A.S. Bohart; BBSL), on Sphaeralcea sp.; 1 9, 1 6, Ceballos, 5 Oct. 1966 (G.E. and A.S. Bohart; BBSL); 1 9, Cuencame, 31 Oct. 1966 (G.E. and A.S. Bohart; BBSL), on Sphaeralcea sp.; 1 9, Durango, 6300 ft., 17 June 1956 (H.A. Scullen; UCD); 3 5<5, 14 mi NE Durango, 6200 4 ■ Contributions in Science, Number 448 Snelling: Diadasia , Subgenus Dasiapis Contributions in Science, Number 448 Sneliing: Diadasia, Subgenus Dasiapis ■ 5 Figure 11. Known distribution in Mexico and Central America of D. ochracea (solid circles), D. olivacea (star), and D. tropicalis (open circles). ft., 17 June 1956 (H.A. Scullen; UCD); 9 <3<3, 156 mi N Durango, 6400 ft., 20 June 1956 (H.A. Scullen; UCD); 2 99, El Tascate, 6400 ft., 28 July 1947 (C.D. Michener; UCD); 4 99, La Loma, 4100 ft., 20 Aug. 1947 (C.D. Mich- ener; UCD); 1 6, 34 mi NW La Zarca, 5800 ft., 25 July 1953 (Univ. Kans. Mex. Exped.; UKAN); 1 9, Pedricena, 18 Oct. 1968 (G.E. Bohart; BBSL), on Sphaeralcea sp.; 1 9, Pedricena, 4500 ft., 19 Aug. 1947 (C.D. Michener; UCD); 1 9, 20 mi S Rodeo, 21 Aug. 1960 (D.C. Rentz; LACM); 1 9, 1 <3, San Juan del Rio, 5200 ft., 30 July 1947 (C.D. Michener; UCD); 8 99, 1 6, Yerbanio, 6700 ft., Cuencame Dist., 19 Aug. 1947 (Schrammel; UCD). Hi- dalgo: 2 99, Actopan, 6500 ft., 27 Aug. 1962 (Ordway and Marston; UKAN), on Malva sp.; 3 99, 7 66, Ixmiquilpan, 5300 ft., 25 June 1953 (Univ. Kans. Mex. Exped.; UKAN), on Sphaeralcea sp.; 1 6, same except 29 July 1954, no floral data (UKAN); 1 6, Tezontepec, 6600 ft., 21 mi SW Actopan, 27 Aug. 1962 (Ordway and Marston; UKAN); 10 66, Zimapan, 8 July 1968 (M.S. Wasbauer and J.E. Slansky; UCD). Jalisco: 4 99, 2 66, 6 mi NE Jalastitlan, 6000 ft., 19 July 1954 (Univ. Kans. Mex. Exped.; UKAN). Oaxaca: 1 <3, Nejapa, 2500-3000 ft., 77 mi W Tehuan- tepec, 22 Aug. 1963 (H.V. Weems; FSCA). San Luis Po- tost: 3 66, 13 mi NW Ciudad del Maiz, 3700 ft., 22 Aug. 1954 (Univ. Kans. Mex. Exped.; UKAN). Zacatecas: 1 <3, Rio Grande, 18 Oct. 1968 (G.E. Bohart; BBSL); 1 6, 34 mi S Zacatecas, 6900 ft., 15 June 1956 (H.A. Scullen; UCD). Diadasia ( Dasiapis ) olivacea (Cresson) Figures 2, 4-5, 10, 11 Melissodes olivacea Cresson, 1878:216; 2 <3. Cres- son, 1916:126; 2 (lectotype). Eucera olivacea: Dalla Torre, 1896:242. Dasiapis olivacea: Cockerell, 1906:100. Lutz and Cockerell, 1920:583. Cresson (1916) selected as lectotype a female from the original series of both sexes; the statement by Hurd (in Krombein et al., 1979) that the type is a male is incorrect. The type series is from an un- specified Mexican locality, collected by Sumichrast and possibly from the vicinity of Cordoba or Ori- zaba, (Cresson, 1868). Other than the type, I have seen a series of 10 females and 6 males from 9 km W Tizapan el Alto, 1585 m elev., Jalisco, Mexico, 12 Sept. 1976 (C.D. George and R.R. Snelling; LACM) (see Fig. 11). Both sexes were at flowers of an unidentified malvaceous plant. Females were also at a small nesting site in an area of sparse, tall grass. A few females of the cleptoparasitic anthophorid, Protepeolus integer Linsley, were at the nesting site. Although both sexes are very similar to the better known D. ochracea, they are conspicuously larger. The pubescence of the head and body is a darker, more brownish yellow than in D. ochracea, though older specimens may prove to be nearly as pale as that species. In addition to the other characteristics cited in the key, it may be noted that the genal and gular areas of D. olivacea are distinctly tessellate and only slightly shiny, rather than smooth and shiny as in D. ochracea. Diadasia ( Dasiapis ) tropicalis (Cockerell) Figures 1, 6-7, 11 Dasiapis tropicalis Cockerell, 1918:27; 9. Dasiapis olivacea: Cockerell, 1949:462; 9. MIS- IDENTIFICATION. Melissodes tenuicincta Cockerell, 1949:464; 2. NEW SYNONYMY. Melissodes spilognathus Cockerell, 1949:468; 2. NEW SYNONYMY. Diadasia spilognathus: LaBerge, 1956:1179. Diadasia tenuicincta: LaBerge, 1956:1179. I have examined the types of all of the above nominate species and believe them to be conspe- cific. The type of D. tropicalis is from Pueblo Viejo, Veracruz, Mexico, and those of M. spilognathus and M. tenuicincta are from Zamorano, Honduras; all are in the USNM. This species is very similar to D. ochracea, and the separation of the two is not always an easy matter. It may well be that further study will show that D. tropicalis is nothing more than a southern variant of D. ochracea. Both species are present in Mexico, but there appears to be little sympatry (see Fig. 1 1). In general, D. ochracea is a more northern species and is largely confined to the central highlands in Mexico. Al- though D. tropicalis ranges sufficiently far north to barely enter the United States in Texas and Arizona, the main distribution lies in Mexico, where it is principally a lowland species. There are few spec- imens available from south of Mexico, but the dis- tribution of D. tropicalis clearly extends at least to El Salvador. Specimens from several localities in Venezuela may also belong to D. tropicalis. Both sexes are available, and they closely resemble the Central American material of D. tropicalis in most features. They do differ, however, in that the punc- tation of the head and mesosoma tends to be weak and shallow, much as in D. ochracea, but in density more like D. tropicalis. For the present they are assigned to D. tropicalis but may represent another species. The correct placement of these specimens must await the acquisition of additional material and a better understanding of the South American species placed in Dasiapis. The most obvious difference between the two species is, unfortunately, one of color: D. ochracea is a dark-legged species, with the femora and tibiae dark brownish, while in D. tropicalis the legs, be- yond the coxae, are consistently reddish to yellow- ish red. This is a very consistent difference. In itself, this would hardly be justification for recognizing D. tropicalis as a species apart from D. ochracea, but this color difference does correlate with the punctation of the mesoscutum. Both species have a dull, densely tessellate meso- scutum. In the case of D. ochracea, there are scat- tered minute (about 0.01 mm diam.) punctures over the middle of the mesoscutal disc that are hardly 6 ■ Contributions in Science, Number 448 Snelling: Diadasia, Subgenus Dasiapis greater in diameter than the hairs arising from them. Such punctures are conspicuously finer than those adjacent to the parapsidal line. Although the pos- terolateral area of the mesoscutum of D. ochracea is usually without punctures, or with only very mi- nute ones, some specimens do possess larger punc- tures that are very shallow and difficult to discern. The mesoscutal punctation of D. tropicalis is quite different. In most specimens, the entire disc is closely punctate with well-defined punctures that are about 0.025 mm in diameter. Some specimens may have the punctures of the central part of the disc largely effaced, but these are still coarser than those of D. ochracea. There are also slight differences in the punctation of the upper frons, but so much variation exists that this is an unreliable, only weakly corroborative, character. Usually, the frons immediately below the ocelli is smooth and shiny in D. ochracea, with only scattered minute punctures. In D. tropicalis this area is often sharply tessellate and with sparse, well-defined coarser punctures in addition to mi- nute punctures. Nothing is known of the biology of D. tropicalis other than that provided by the floral data associ- ated with some of the specimens examined. Most of these are for the family Malvaceae, the apparent usual pollen source for species of Dasiapis. I have examined the following material of D. tropicalis (see Fig. 11). EL SALVADOR: 3 99, 5 mi N Quezaltepec, 23 Aug. 1961 (M.E. Irwin; UCD). GUATEMALA: 3 92, 9 56, E! Progreso, 99 km NE Guatemala City, 21 Sept. 1975 (J.L. Neff; CTMI), on Sida “7084” (2 22, 2 66) and Tribulus sp. (1 2, 7 55). HONDURAS: 1 2, Zamorano, Dec. (Vidales; type of Melissodes tenuicincta ; USNM No. 58546); 1 2, Zamo- rano, 1 Dec. (A. Pelen; type of M. spilognathus; USNM No. 58570). MEXICO, Baja California Sur: 1 2, 48 km S Loreto, 425 m, 14 Sept. 1983 (R.R. Snelling; LACM). Chiapas: 1 5, Ocozocuautla, 14 Sept. 1974 (W. Hanson and G. Bo- hart; BBSL); 6 22, 3 55, Tuxtla Gutierrez, Oct. 1954 (N.L.H. Kraus; UKAN); 1 5, 9 mi N Villa Flores, 12 Aug. 1963 (F.D. Parker and L.A. Stange; UCD). Chihuahua: 1 5, Ciudad juarez, 3700 ft., 28 July 1953 (Univ. Kans. Mex. Exped .; UKAN), on Sphaeralcea. Nayarit: 1 5, A capo - neta, 29 Sept. 1966 (G.E. and A.S. Bohart; BBSL); 1 2, Acaponeta, 12 Oct. 1968 (G.E. Bohart; BBSL), on “orange Cosmos ”; 1 2, Ruiz, 29 Sept. 1966 (G.E. and A.S. Bohart; BBSL). Nuevo Leon: 1 2, 37 mi NE China, 13 July 1954 (Univ. Kans. Mex. Rxped.; UKAN). Oaxaca: 1 5, 100 mi S Acayucan (Yucatan), in Oaxaca, 600 ft., 18 Aug. 1963 (Scullen and Bolinger; UCD); 1 2, 19 mi SE El Camaron, 2800 ft., 19 Aug. 1963 (Scullen and Bolinger; UCD); 2 55, 18 mi N La Ventosa, 21 Aug. 1966 (J.B. Karren; UKAN), on Waltheria americana; 1 5, 23 mi S Marias Romero, 14 Aug. 1963 (F.D. Parker and L.A. Stange; UCD); 5 22, 1 5, 4 mi N Pochutla, 150 m, 11 Oct. 1975 (J.L. Neff; CTMI), on Sida “7117”; 1 5, 8 mi W Tapanatepec, 400 ft., 10 July 1953 (Univ. Kans. Mex. F.xped.; UKAN); 1 2, 2 55, 12 mi W Tehuantepec, 11 Sept. 1974 (G.E. Bohart and W.J. Hanson; BBSL); 1 6, 35 W Tehuantepec, 16 Sept. 1974 (W. Hanson and G. Bohart; BBSL); 2 56, 4 mi N To tol apart, 1849 m, 15 Sept. 1975 (J.L. Neff; CTMI), on Sida “7063”; 1 2, 4 mi W Zanatepec, 200 m, 16 Sept. 1975 (J.L. Neff; CTMI); 1 2, same locality and collector, 7 Oct. 1975, on Sida (CTMI). San Luis Potosu 1 5, El Salto, 1600 ft., 24 Aug. 1954 (Univ. Kans. Mex. Exped.; UKAN); 1 2, 5 55, El Salto, 1250 ft., 4 Sept. 1962 (Ordway and Marston; UKAN); 3 22, 4 55, El Salto, 20-21 July 1962 (Univ. Kans. Mex. Exped.; UKAN). Sinaloa: 1 6, Culiacan, 10 Oct. 1968 (G.E. Bohart; BBSL), on Euphor- bia sp.; 1 2, 2 66, Guamuchil, 27 Oct. 1966 (G.E. and A.S. Bohart; BBSL, UCD); 1 2, 4 56, Escuinapa, Sept. 1966 (G.E. and A.S. Bohart; BBSL), 1 5, on Convolvulus sp.; 1 2, 2 66, near Guasave, 27 Oct. 1965 (G.E. and A.S. Bohart; BBSL), on Kallstroemia sp.; 1 2, 5 65, Elota, 10 Oct. 1968 (G.E. Bohart; BBSL); 3 22, 2 55, 16 km N Los Mochis, 152 m, 30 Sept. 1976 (C.D. George and R.R. Snelling; LACM), on Anligonon leptopus; 5 22, 15 55, 20 mi N Los Mochis, 9 Oct. 1968 (G.E. Bohart; BBSL), on “Malva- ceae”; 3 55, 31 km N Mazatlan, 76 m, 29 Sept. 1976 (C.D. George and R.R. Snelling; LACM); 1 6, Mazatlan, 1 1 Oct. 1968 (G.E. Bohart; BBSL); 1 6, Rosarito, 29 Sept. 1966 (G.E. and A.S. Bohart; BBSL) on “tall yell, comp.” Sonora: 1 6, Alamos, 27 Sept. 1966 (G.E. and A.S. Bohart; BBSL), “near Malva ”; 7 22, 6 65, Alamos, 8 Oct. 1968 (G.E. Bohart; BBSL), on Malvaceae; 2 22, 13 55, Alamos, 7 Sept. 1970 (G.E. and R.M. Bohart; BBSL); 3 22, 1 6, Alamos, 4 Sept. 1970 (W.J. Hanson and T.L. Whitworth; BBSL); 5 22, SE of Alamos, 5 Sept. 1970 (W.J. Hanson and T.L. Whitworth; BBSL); 1 5, Belen, 28 Sept. 1966 (G.E. and A.S. Bohart; BBSL), on Convolvulus ; 1 2, 10 mi S Guay- mas, 8 Oct. 1968 (G.E. Bohart; BBSL), on Antigonon [leptopus]; 2 22, 25 mi N Guaymas, 26 Sept. 1966 (G.E. and A.S. Bohart; BBSL); 1 6, Masiaca, 9 Oct. 1968 (G.E. Bohart; BBSL), on Kallstroemia; 2 22, 1 5, Obregon, 26 Sept. 1966 (G.E. and A.S. Bohart; BBSL). Tamaulipas: 1 5, 6.2 mi S San Fernando, 8 June 1966 (Univ. Kans. Mex. Exped.; UKAN); 1 5, Tampico, 26 Mar. 1951 (W.P. Ste- phen; UKAN); 1 6, 2.6 mi N Nuevo Morelos, 900 ft., 4 Sept. 1962 (Ordway and Marston; UKAN); 1 6, 8 mi S Jimenez, 15 June 1953 (Univ. Kans. Mex. Exped.; UKAN), on Parthenium hysterophorus; 2 22, 1 6, Paditta, 15 June 1953 (Univ. Kans. Mex. Exped.; UKAN), on Cevallia sin- uata; 1 2, 1 5, 6.6 mi W Antigua Morelos, 400 ft., 3 Sept. 1962 (Ordway and Naumann; UKAN). Veracruz: 2 66, 5 mi NE Tenajas, 18 Aug. 1963 (F.D. Parker, L.A. Stange; UCD); 1 2, Pueblo Viejo, 8 Dec. 1909 (F.C. Bishopp; type of Dasiapis tropicalis; USNM No. 22925). Zacatecas: 1 6, 10 mi S Jalpa, 17 Sept. 1970 (G.E. and R.M. Bohart; BBSL). UNITED STATES, Arizona: 1 2, Yaqui Canyon area, 5700 ft., Huachuca Mrs., Cochise Co., 29 Aug. 1972 (R.R. Snelling; LACM); 1 2, Huachuca Mts., Cochise Co., 30 Aug. 1953 (G.D. Butler; LACM), on Sida procumbens. Texas: 1 6, Progreso, Hidalgo Co., 12 Apr. 1950 (Mich- eners, Rozens, Beamers, Stephen; UKAN), on Verbena sp. VENEZUELA: 1 2, 2 55, Falcon, near Marpare, 19 Dec. 1976 (J.L. Neff; CTMI), on Tribulus. 1 2, 6 66, Escuela Naval, Parroquia Catia la Mar, Distrito Federal, various dates between 30 June and 31 Oct. (N. Ramirez, LACM), on Bastardia viscosa (2 65), undet. Malvaceae (1 2, 1 5), Lantana (1 6), Heliotropium (1 6), and undet. Zygophyllaceae (1 6); 1 2, Estacion Biologia de Sos Llanos, Calabozo [8°56'N, 67°25'W], Guarico, 2 Oct. 1982 (N. Ramirez; LACM), on undet. Malvaceae. ACKNOWLEDGMENTS Although this study was based primarily on specimens in the collections of the Natural History Museum of Los Angeles County (LACM), I have also utilized material Contributions in Science, Number 448 Snelling: Diadasia, Subgenus Dasiapis ■ 7 from the Bee Biology and Systematics Laboratory, Logan, Utah (BBSL), the Central Texas Melittological Institute, Austin, Texas (CTMI), the Florida State Collection of Arthropods, Gainesville, Fla (FSCA), the Snow Entomo- logical Museum of the University of Kansas, Lawrence (UKAN), and the University of California at Davis (UCD). To Terry Griswold, Jack Neff, Lionel Stange, Rob Brooks, and Lynn Kimsey, respectively, of these institutions, my thanks for making material available. Special thanks are extended to Don Azuma, Academy of Natural Sciences of Philadelphia, and Ron McGinley, National Museum of Natural History, for loaning the types essential to this study. LITERATURE CITED Adlakha, R.L. 1969. A systematic revision of the bee genus Diadasia Patton in America north of Mexico (Hymenoptera: Anthophoridae). Ph.D. thesis, Uni- versity of California, Davis, 188 pp. Cockerell, T.D.A. 1903a. North American bees, and a new homopteran. Annals and Magazine of Natural History (series 7) 12:442-455. . 1903b. A catalogue of the bees of California. Psyche 10:74-78. . 1906. The North American bees of the family Anthophoridae. T ransactions of the American En- tomological Society 32:63-116. . 1918. Neotropical bees, principally collected by Professor Bruner in Argentina. Transactions of the American Entomological Society 44:25-38. . 1924. Anthophorid bees in the collection of the California Academy of Sciences. Pan-Pacific En- tomologist 1:49-56. . 1949. Bees from Central America, principally Honduras. Proceedings of the United States Na- tional Museum 98:429-490. Cresson, E.T. 1868. Catalogue of a collection of Hy- menoptera made by Prof. E. Sumichrast near Cor- dova, Mexico. Transactions of the American En- tomological Society 2:1-38. . 1878. Descriptions of new species of North American bees. Proceedings of the Academy of Nat- ural Sciences of Philadelphia 30:181-221. . 1916. The Cresson types of Hymenoptera. Memoirs of the American Entomological Society 1: 1-141. Dalla Torre, C.G. de. 1896. Catalogus Hymenoptero- rum. X. Apidae (Anthophila). Leipzig, 648 pp. Eickwort, G.C., K.R. Eickwort, and E.G. Linsley. 1977. Observations on nest aggregations of the bees Dia- dasia olivacea and D. diminuta (Hymenoptera: An- thophoridae). Journal of the Kansas Entomological Society 50:1-17. Krombein, K.V., P.D. Hurd, Jr., D.R. Smith, and B.D. Burks. 1979. Catalog of Hymenoptera in America north of Mexico. Washington, D.C.: Smithsonian Institution Press, 2:1199-2209. LaBerge, W.E. 1956. A revision of the bees of the genus Melissodes in North and Central America. Part I (Hymenoptera, Apidae). University of Kansas Sci- ence Bulletin 37:911-1194. Linsley, E.G., and J.W. MacSwain. 1957. The nesting habits, flower relationships, and parasites of some North American species of Diadasia (Hymenoptera: Anthophoridae). Wasmann Journal of Biology 15: 199-235. Lutz, F.E., and T.D.A. Cockerell. 1920. Notes on the distributions and bibliography of North American bees of the families Apidae, Meliponidae, Bombi- dae, Euglossidae, and Anthophoridae. Bulletin of the American Museum of Natural History 42:491-641. Michener, C.D. 1954. Bees of Panama. Bulletin of the American Museum of Natural History 104:1-176. Muesebeck, C.F.W., K.V. Krombein, and H.K. Townes. 1951. Hymenoptera of America north of Mexico. Synoptic catalog. Washington, D.C.: United States Department of Agriculture, Agricultural Monograph 2, 1420 pp. Patton, W.H. 1879. Generic arrangement of the bees allied to Melissodes and Anthophora. Bulletin of the United States Geological and Geographical Survey 5:471-479. Rozen, J.G., K.R. Eickwort, and G.C. Eickwort. 1978. The bionomics and immature stages of the clepto- parasitic bee genus Protepeolus (Anthophoridae, Nomadinae). American Museum Novitates 2640:1- 24. Timberlake, D.H. 1941. Synoptic table of North Amer- ican species of Diadasia (Hymenoptera, Apoidea). Bulletin of the Brooklyn Entomological Society 36: 2-11. Accepted 4 May 1994. 8 ■ Contributions in Science, Number 448 Snelling: Diadasia, Subgenus Dasiapis Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Number 449 6 October 1994 *2/ (■ Contributions in Science Revision and New Species of the Apocephalus ( Mesophora ) TRUNCATICERCUS- INFRAGROUP (Diptera: Phoridae) Brian V. Brown Natural History Museum of Los Angeles County Serial Publications of THE Natural History Museum of Los Angeles County Scientific Publications Committee James L. Powell, Museum President Daniel M. Cohen, Committee Chairman Brian V. Brown Kenneth E. Campbell Kirk Fitzhugh Edward C. Wilson Robin A. Simpson, Managing Editor The scientific publications of the Natural History Mu- seum of Los Angeles County have been issued at irregular intervals in three major series; the issues in each series are numbered individually, and numbers run consecutively, regardless of the subject matter. # Contributions in Science, a miscellaneous series of tech- nical papers describing original research in the life and earth sciences. 9 Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth sci- ences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now published by the Museum in Contributions in Science. • Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through the Museum Book Shop. A catalog is available on request. The Museum also publishes Technical Reports, a mis- cellaneous series containing information relative to schol- arly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum’s Scientific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Tech- nical Reports may be obtained from the relevant Section of the Museum. Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, California 90007 Printed at Allen Press, Inc., Lawrence, Kansas ISSN 0459-8113 Revision and New Species of the Apocephalus ( Mesophora ) TRUNCATICERCUS -INFRAGROUP (Diptera: Phoridae) Brian V. Brown1 ABSTRACT. The putatively monophyletic Apocephalus ( Mesophora ) truncaticercus- infragroup is diag- nosed with hypothesized synapomorphic character states. Three new species are described: A. grandiflavus from the Dominican Republic, and two from Costa Rica, A. nitifrons and A. satanus. The previously unknown females of A. brevicercus and A. truncaticercus are recognized, and both species are redescribed. A key for the identification of adults of this group is presented. Species are extremely similar, and little can be said about their phylogenetic relationships at this time, except that it is possible that A. brevicercus and A. grandiflavus are exclusive sister species. INTRODUCTION In my revision of the parasitic genus Apoceph- alus, subgenus Mesophora (Brown, 1993), I de- scribed two new, closely related species: A. trun- caticercus and A. brevicercus. Both were known only from male specimens collected at a single lo- cality in Costa Rica. Since that time, a number of new specimens have been collected, including the apparent females of A. truncaticercus and A. brevi- cercus, as well as three new species from Costa Rica and the Dominican Republic. METHODS AND MATERIALS Treatment of specimens and terms used are the same as in my previous publications (e.g., Brown, 1992). In the descriptions, the following ratios are used: frontal ratio = height of frons divided by width of frons slightly below the anterior ocellus; costal ratio = distance from basicosta to costal apex, divided by distance from basicosta to wing apex; and costal sector ratio = distance, along the costa, from humeral crossvein to apex of vein Rt (sector l):Rj to R2+3 (sector 2) : R2+3 to R4+5 (sector 3) (all sector mea- surements are standardized by dividing by the length of sector 3). Unless fewer specimens were available, at least 10 individuals were measured. Latitude and longitude coordinates are quoted as on data labels, either as degrees and minutes (e.g., 35°30'N) or as decimal degrees (e.g., 35.5°N). Depositories of the material examined are as follows: CMNH Section of Invertebrate Zoology, Carnegie Mu- seum of Natural History, 4400 Forbes Avenue, Pittsburgh, PA 15213-4080, USA (C.W. Young) INBIO Instituto Nacional de Biodiversidad, A.P. 22- 1. Entomology Section, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los An- geles, California 90007. Contributions in Science, Number 449, pp. 1-7 Natural History Museum of Los Angeles County, 1994 3100, Santo Domingo, Heredia, Costa Rica (M. Zumbado) LACM Entomology Section, Natural History Museum of Los Angeles County, 900 Exposition Bou- levard, Los Angeles, CA 90007, USA (B.V. Brown) MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, USA (on indefinite loan to the author) MICR Museo de Insectos, Universidad de Costa Rica, San Pedro, San Jose, Costa Rica (P.S. Hanson) USNM United States National Museum, Smithsonian Institution, Washington, DC 20560, USA (on indefinite loan to the author) SYSTEMATICS Apocephalus Coquillett, 1901, subgenus Mesophora Borgmeier, 1937 EMENDED DIAGNOSIS. The genus Apoceph- alus was diagnosed previously (Brown, 1993, p. 218), but because of the atypical structure of females of the A. truncaticercus- infragroup, the following change, shown in italic, should be made: . . . an- terior margin [of ovipositor] usually with dark pe- riphery in dorsal view; species with a uniformly darker ovipositor lack middle and lower fronto- orbital seta. The diagnosis of subgenus Mesophora given pre- viously (Brown, 1993, p. 218) is still adequate, al- though females of the A. truncaticercus- infragroup must be added to A. antennatus Malloch as species lacking dark abdominal glands. Apocephalus truncaticercus-'miragroup Brown, 1993 DIAGNOSIS. Male with apex of flagellomere 1 markedly narrowed at base (Fig. 1). Female flagel- Figures 1-6. Apocephalus ( Mesophora ) truncaticercus- group species (scale bar = 0.1 mm; Figs. 1-4 to same scale). 1-3. A. brevicercus Brown. 1, Male frons and flagellomere 1, frontal; 2, female head, lateral; 3, male head, lateral. 4. A. nitifrons new species, male head, lateral. 5, 6. A. brevicercus Brown. 5, Female ovipositor, dorsal; 6, female foretarsomeres 3-5. Abbreviations: 5 = tarsomere 5; A = anterior projection of ovipositor; F = flagellomere 1; P = palpus. lomere 1 pyriform (Fig. 2). Female lacking lower and middle fronto-orbital seta; lacking darkly scler- otized abdominal glands; ovipositor dark, evenly sclerotized, without darker anterior margin; tar- somere 5 and claw enlarged, especially on foreleg (Fig. 6). NOTES ON DIAGNOSIS AND RECOGNI- TION. As a group, males of these species are rec- ognized by the elongate, narrowed apex of flagel- lomere 1, but species recognition within the group is difficult. All are extremely similar, with only slight structural differences. Within the Phoridae, species are defined narrowly at present, and research has shown that there are groups of extremely similar cryptic forms within some genera (Disney, 1989). These forms generally are recognized as separate species. Therefore, with these guidelines in mind, three new species are described below. Collection of further specimens from more sites will resolve the species limits in this complex. Females of this group are highly atypical. They lack the dark abdominal glands found in most other species of subgenus Mesophora, and the ovipositor is dark and uniformly sclerotized, unlike that of most other Apocephalus. Females can be recog- nized relatively easily by the lack of both middle 2 ■ Contributions in Science, Number 449 Brown: Revision of Apocephalus and lower fronto-orbital setae, both of which are otherwise absent in only A. limai Prado and Apo- cephalus unnamed species A (of Brown, 1993). Both of these other species have ovipositors with dark anterior margins and smaller apical tarsomeres. Furthermore, A. truncaticercus- infragroup fe- males are structurally divergent from males. This is especially striking in A. truncaticercus and A. brevi- cercus, species that have mostly brown-colored males and yellow-colored females. One of the few characters that link the two sexes is the presence of enlarged tarsal claws, which are especially pro- nounced in females. PHYLOGENETIC RELATIONSHIPS. These species are part of the A. wheeleri- subgroup (Brown, 1993), diagnosed by the presence of a rounded dor- sal apex of the female ovipositor. Monophyly of the A. truncaticercus- infragroup is supported by the following hypothesized synap- omorphic character states: 1. Flagellomere 1 of males narrowed abruptly on apical two-thirds (Fig. 1). 2. Right surstylus of male with number of setae reduced to three or fewer (Figs. 7, 9, 11). The holotype of A. truncaticercus is an unusual ex- ception, with more setae present. 3. Apex of male cercus truncate, slightly enlarged apically (Brown, 1993: figs. 8, 9, 29, 30). 4. Female tarsomere 5 and claw enlarged, especial- ly on foreleg (Fig. 6). 5. Middle fronto-orbital seta absent on female. 6. Female ovipositor darkly sclerotized through- out. 7. Female ovipositor with long, narrow anterior process (Fig. 5). Within this group, I cannot reconstruct the phy- logenetic relationships at this time. Species are ex- tremely similar, with the characters separating them of little apparent phylogenetic value. In my search for synapomorphic character states to group spe- cies, I considered the following: 1. Palpus enlarged, inflated, with setulae reduced (Fig. 3). Most outgroup taxa have a relatively small palpus with well-developed setulae. This is the most promising character state, and represents a potential synapomorphy of A. gran- diflavus and A. brevicercus. It also occurs in a few other species of subgenus Mesophora, out- side of the A. truncaticercus- infragroup. 2. Body color mostly dark brown. Outgroup taxa, other species of Apocephalus, generally are yellow-colored. The phylogenetic value of this character state is dubious. Specimens of A. brevicercus and A. truncaticercus can be either brown or yellow, and some other, unrelated, species of subgenus Mesophora are brown to a varying extent. Brown body color as a synapomorphy would group A. brevicercus (most specimens), A. nitifrons, A. satanus, and A. truncaticercus, conflicting with the other proposed synapomorphy, above. Analysis of these scanty data seems futile. When females are known for all species, some further character states likely will become apparent. LIFE HISTORY. The hosts of these flies are un- known, but their closest relatives are parasitoids of beetles of the family Lampyridae (Brown, 1994; Lloyd, 1973). These other species are multiple par- asitoids of their hosts, and the abdomen of one specimen of A. grandiflavus contained 22 mature eggs. KEY TO ADULT MALES 1 Setae of cercus and proctiger of normal size, longer and thicker than short setae of epan- drium (Brown, 1993: figs. 9, 30); hind femur of uniform color 2 Setae of cercus and proctiger markedly re- duced, subequal to short setae of epandrium (Brown, 1993: figs. 8, 29); hind femur with abrupt, distinctive darkening at apex Apocephalus brevicercus Brown 2(1) Palpus of male small with setae normal, thick, moderately long, pointed (Fig. 4); right side of epandrium, in addition to ventral setae, with 1-3 lateral setae (Figs. 9, 11) 3 Palpus of male enlarged, with setae reduced, thin, short, stubby (Fig. 3); right side of epan- drium, in addition to ventral setae, with nu- merous lateral setae (Fig. 7) Apocephalus grandiflavus new species 3(2) Frons glossy, dark brown Apocephalus nitifrons new species Frons opaque, dark brown to yellow .... 4 4(3) Flagellomere 1 dark brown throughout; hind femur dark brown; palpus brown; epandrium with anterior part short, bulging anteriorly (Figs. 11, 12) Apocephalus satanus new species Flagellomere 1 yellowish at base, apically darker; hind femur yellowish brown; palpus yellow; epandrium with anterior part short, straight (as in Figs. 9, 10) Apocephalus truncaticercus Brown KEY TO ADULT FEMALES (Note: Female of A. nitifrons unknown; females of A. grandiflavus and A. truncaticercus cannot be separated at this time.) 1 Tip of hind femur dark brown to black, con- trasting sharply with yellowish color of rest of femur; frons glossy yellow Apocephalus brevicercus Brown Tip of hind femur yellowish brown or dark brown, color little differentiated from rest of femur; frons opaque, yellow or brown 2 Contributions in Science, Number 449 Brown: Revision of Apocephalus H 3 2 Body color, including halter and palpus, dark brown Apocephalus satanus new species Body color yellow Apocephalus grandiflavus new species Apocephalus truncaticercus Brown Apocephalus brevicercus Brown, 1993 Figures 1-3, 5, 6 SPECIES RECOGNITION AND GEOGRAPH- ICAL DISTRIBUTION. Males are recognized eas- ily by the reduced setae of the cercus. Females can be separated from those of other species by the glossy yellow frons and the darkened apex of the hind femur. Besides several localities in Costa Rica, this spe- cies occurs also in the Dominican Republic. Unlike specimens from Costa Rica, which have the frons brown, some specimens from the Dominican Re- public have the frons yellow. Furthermore, some newly collected specimens from Costa Rica and the Dominican Republic have the halter yellow, rather than brown as in the original description. The females are strikingly divergent from the males: females are yellow, whereas males are mostly dark brown in color. The original association of the sexes was based on co-occurrence of male and female specimens in Malaise trap samples from San Gerardo, Costa Rica, where apparently A. brevi- cercus is the only Mesophora species present. DESCRIPTION. Body length 1.6-2. 3 mm. Frons yellow or brown; opaque in male, glossy yellow in female. Mean frontal ratio 0.79. Color of flagel- lomere 1 yellowish at base, apically darker in male; flagellomere 1 of female yellow. Flagellomere 1 of female pyriform (Fig. 2). Palpus yellow. Palpus of male enlarged, with setae reduced, thin, short, stub- by (Fig. 3). Palpus of female small to enlarged, with setae normal, thick, moderately long, pointed (Fig. 2). Dorsum of thorax yellow to dark brown. Pleural regions yellow to dark brown. Legs yellowish brown. Hind femur yellowish brown, with abrupt, distinc- tive darkening at apex. Mean costal ratio 0.56, range 0.54-0.59. Mean costal sector ratio 3.96:3.45:1, range 3.08-5.50:2.71-4.50:1. Wing vein R2+3 well developed. Halter yellow or brown. Ventral setae of female segment 6 absent. Venter of male segment 6 bare. In cleared specimens, female dorsal abdom- inal glands invisible. Female Terminalia. Female intersegment 6-7 without distinct sclerite. Ovipositor (Fig. 5) trian- gular; dorsally broad; anterodorsal portion nar- rowed, elongate; anteroventral margin smooth, without notch; posterodorsal apex of ovipositor rounded; in lateral view, posteroventral apex of ovipositor straight; posteroventral apex of ovipos- itor rounded. Stylet with lateral barbs. Dorsal scler- ite consisting of two small processes. Male Terminalia. Epandrium globular, with an- terior part short, straight, with setae near cercus not markedly larger than other epandrial setae. Left side of epandrium with thick, ventral setae. Right side of epandrium with faint, unsclerotized dorsolateral ridge; in addition to ventral setae, with one to three lateral setae. Right surstylus short, rounded, lacking ventral carina. Number of setae on right surstylus two to three. Cercus yellow. Setae of cercus and proctiger markedly reduced, subequal to short setae of epandrium. NEW MATERIAL EXAMINED. COSTA RICA: San Jose: 20 km S. Empalme, 9.63°N, 83.85°W, 19, ix.1988, P. Hanson, Malaise trap, 2800 m (LACM), 6 km N San Gerardo, 9.95°N, 84.05°W, 29, ix.1992, 39, x.1992, IS, 89, xi.1992, 39, xii.1992, IS, 39, i.l 993, P. Hanson, Ma- laise trap, 2800 m (INBIO, LACM, MCZ, MICR, USNM), 26 km N San Isidro, 9.5°N, 83.72°W, IS, ii-v.1992, P. Hanson, Malaise trap, 2100 m (LACM); Zurqui de Mo- ravia, 10.05°N, 84.02°W, 19, x-xii.1990, 19, iii.1991, 1<3, iv.1991, IS, vi.1991, 29, xii.1991-ii.1992, 19, iv-v.1993, IS, l-15.vi.1993, P. Hanson, Malaise trap, 1600 m (LACM, MCZ, MICR). DOMINICAN REPUBLIC: Independen- ce: 5 km NNW Angel Feliz, 18.68°N, 71.78°W, IS, 13- 15.X.1991, IS, 21-22.vii.1992, J. Rawlins et al, cloud forest, 1780 m (CMNH, LACM); Pedernales: La[s] Abeja[s], 38 km NNW Cabo Rojo, 18.15°N, 71.63°W, IS, 19, 15.vii.1987, J. Rawlins & R. Davidson, 1250 m (CMNH, LACM), Upper Las Abejas, 38 km NNW Cabo Rojo, 18.15°N, 71.63°W, 29, 22.vii.1990, L. Masner, sweeping, mesic deciduous forest, 1350 m (CMNH). Apocephalus grandiflavus new species Figures 7, 8 SPECIES RECOGNITION. Males can be rec- ognized by the enlarged, yellow palpi and the great- er number of setae on the right side of the epan- drium. Females can be separated from those of A. brevicercus (the only other species of this group known from the Dominican Republic) by the hind femur, which is without dark color at base. DESCRIPTION. Body length 1.4-2 mm. Frons yellow, opaque. Mean frontal ratio 0.78. Color of flagellomere 1 yellowish at base, apically darker in male, female with flagellomere 1 yellow. Flagello- mere 1 of female pyriform. Palpus yellow. Palpus of male enlarged, with setae reduced, thin, short, stubby. Palpus of female small to elongate, with setae normal, thick, moderately long, pointed. Dor- sum of thorax yellow. Pleural regions yellow. Legs yellow. Hind femur yellowish brown; evenly col- ored throughout. Mean costal ratio 0.55, range 0.53- 0.58. Mean costal sector ratio 4.99:4.13:1, range 4.25-6.33:3.67-5.00:1. Wing vein R2+3 well devel- oped. Halter yellow. Ventral setae of female seg- ment 6 absent. Venter of male segment 6 bare. In cleared specimens, female dorsal abdominal glands invisible. Female Terminalia. Female intersegment 6-7 without distinct sclerite. Ovipositor triangular; dor- sally broad; anterodorsal portion narrowed, elon- gate; anteroventral margin smooth, without notch; posterodorsal apex of ovipositor rounded; in lateral view, posteroventral apex of ovipositor straight; posteroventral apex of ovipositor rounded. Stylet with lateral barbs. Dorsal sclerite consisting of two small processes. 4 ■ Contributions in Science, Number 449 Brown: Revision of Apocephalus Figures 7-12. Apocepbalus ( Mesophora ) truncaticercus- group species, male terminalia, cercus and proctiger truncated, right side and left side (scale bar = 0.1 mm; all figures to same scale). 7, 8. A. grandiflavus new species. 9, 10. A. nitifrons new species. 11, 12. A. satanus new species. Abbreviations: B = bulge of epandrium; R = right surstylus. Contributions in Science, Number 449 Brown: Revision of Apocepbalus ■ 5 Male Terminalia. Epandrium globular, with an- terior part elongate (Figs. 7, 8), with setae near cercus not markedly larger than other epandrial setae. Left side of epandrium with thick, ventral setae. Right side of epandrium with faint, unscler- otized dorsolateral ridge; in addition to ventral se- tae, with numerous lateral setae. Right surstylus short, rounded, with ventral carina short, extended partially across surstylus. Number of setae on right surstylus two. Cercus yellow. Setae of cercus and proctiger of normal size, longer and thicker than short setae of epandrium. GEOGRAPHICAL DISTRIBUTION. Known only from some middle-elevation sites in the Do- minican Republic. DERIVATION OF SPECIFIC EPITHET. The name of this species is based on Latin words mean- ing “large” and “yellow,” referring to the palpus of male specimens. HOLOTYPE. 5, DOMINICAN REPUBLIC: Pedernales: 3.3 km NE Los Arroyos, 18°15'N, 71°45'W, 16-18.vii.1990, L. Masner et al., wet montane forest, sweep samples, 1450 m (CMNH). PARATYPES. DOMINICAN REPUBLIC: Azua: 7 km WNW Barrero, 18.35°N, 70.97°W, 1 <3, 25- 26.vii.1992, C. Young et al., cloud forest adjacent to disturbed forest, 860 m (CMNH); Pedernales: La[s] Abeja[s], 38 km NNW Cabo Rojo, 18°09'N, 71°38'W, 186, 302, 15.vii.1987, J. Rawlins & R. Davidson, 1250 m (CMNH, LACM, MCZ), Upper Las Abejas, 12, 22.vii.1990, L. Masner, sweeping, mesic deciduous forest, 1350 m (CMNH), 186, 62, same data as holotype (CMNH, LACM, USNM), 5 km NE Los Arroyos, 18°15'N, 71°45'W, 12, 17- 18.vii.1990, 16, 28.vii.l990,C.W. Younger/., 1680 m (CMNH). Apocephalus nitifrons new species Figures 2, 9, 10 SPECIES RECOGNITION. Males can be rec- ognized by the shiny, dark brown frons, brownish palpus, and brown halter. Females are unknown. DESCRIPTION. (Male only, female unknown.) Body length 1 .3-1 .6 mm. Frons brown, glossy. Mean frontal ration 0.73. Color of flagellomere 1 yellow- ish at base, apically darker. Palpus brown, small, with setae normal, thick, moderately long, pointed (Fig. 2). Dorsum of thorax dark brown. Pleural regions dark brown. Legs yellowish brown. Hind femur yellowish brown, evenly colored through- out. Mean costal ratio 0.56, range 0.53-0.58. Mean costal sector ratio 3.98:3.48:1, range 3.78-4.50:3.33- 3.75:1. Wing vein R2+3 well developed. Halter brown. Venter of male segment 6 bare. Male Terminalia. Epandrium globular, with an- terior part short, straight (Figs. 9, 10), with setae near cercus not markedly larger than other epan- drial setae. Left side of epandrium with thick, ven- tral setae. Right side of epandrium with faint, un- sclerotized dorsolateral ridge; in addition to ventral setae, with one to three lateral setae. Right surstylus short, rounded, with ventral carina short, extended partially across surstylus. Number of setae on right surstylus one. Cercus brown. Setae of cercus and proctiger of normal size, longer and thicker than short setae of epandrium. GEOGRAPHICAL DISTRIBUTION. Known only from two sites in Costa Rica. DERIVATION OF SPECIFIC EPITHET. The name of this species is derived from Latin words meaning “shiny frons.” HOLOTYPE. 6, COSTA RICA: San Jose: Zurqui de Moravia, 10.05°N, 84.02°W, vii.1992, P. Han- son, Malaise trap, 1600 m (LACM). PARATYPES. COSTA RICA: Puntarenas: Las Alturas, 8.95°N, 82.83°W, 36, i.l 992, P. Hanson, Malaise trap (LACM, MICR); San Jose: Zurqui de Moravia, 10.05°N, 84.02°W, 16, x-xii.1990, 16, xii- ii.l 992, 16, vii.1992, P. Hanson, Malaise trap, 1600 m (LACM). Apocephalus satanus new species Figures 11, 12 SPECIES RECOGNITION. Males and females can be recognized by the overall dark brown color. DESCRIPTION. Body length 1 .4-1.8 mm. Frons brown, opaque. Mean frontal ratio 0.84. Color of flagellomere 1 brown. Flagellomere 1 of female pyriform. Palpus brown. Palpus of male small, with setae normal, thick, moderately long, pointed. Pal- pus of female small, with setae normal, thick, mod- erately long, pointed. Dorsum of thorax dark brown. Pleural regions dark brown. Legs dark brown. Hind femur dark brown, evenly colored throughout. Mean costal ratio 0.57, range 0.56-0.59. Mean cos- tal sector ratio 3.61:2.53:1, range 3.17-4.20:2.17- 2.80:1. Wing vein R2+3 well developed. Halter brown. Ventral setae of female segment 6 absent. Venter of male segment 6 bare. In cleared speci- mens, female dorsal abdominal glands invisible. Female Terminalia. Female intersegment 6-7 without distinct sclerite. Ovipositor triangular; dor- sally broad; anterodorsal portion narrowed, elon- gate; anteroventral margin smooth, without notch; posterodorsal apex of ovipositor rounded; in lateral view, posteroventral apex of ovipositor straight; posteroventral apex of ovipositor rounded. Stylet with lateral barbs. Dorsal sclerite consisting of two small processes. Male Terminalia. Epandrium globular, with an- terior part short, bulging anteriorly (Figs. 11, 12), with setae near cercus not markedly larger than other epandrial setae. Left side of epandrium with thick, ventral setae. Right side of epandrium with faint, unsclerotized dorsolateral ridge; in addition to ventral setae, with one to three lateral setae. Right surstylus short, rounded, with ventral carina short, extended partially across surstylus. Number of setae on right surstylus three. Cercus brown. Setae of cercus and proctiger of normal size, longer and thicker than short setae of epandrium. GEOGRAPHICAL DISTRIBUTION. Known from three middle-elevation sites in Costa Rica. DERIVATION OF SPECIFIC EPITHET. The 6 ■ Contributions in Science, Number 449 Brown: Revision of Apocephalus species is named after Satan, an indirect reference to darkness, and refers to the dark color of these flies. HOLOTYPE. 3, COSTA RICA: San Jose: Zurqui de Moravia, 10.05°N, 84.02°W, iv-v.1993, P. Han- son, Malaise trap, 1600 m (LACM). PARATYPES. COSTA RICA: Puntarenas: Mon- teverde, 10.1°N, 83.43°W, 13, 12-15.vi.1974, J.P. Donahue, Malaise trap (LACM); San Jose: 14 km N San Isidro, 9.37°N, 83.7°W, 13, 20-23.vi.1974, J.P. Donahue, Malaise trap (LACM), Zurqui de Moravia, 10.05°N, 84.02°W, 19, v.1992, P. Hanson, Malaise trap, 1600 m (LACM). Apocephalus truncaticercus Brown, 1993 NOTES ABOUT NEW MATERIAL. Addition- al specimens of this species have been collected. Unlike the holotype, these new specimens have only one or two setulae on the right surstylus. Specimens from La Cangreja have a yellower frons and halter, unlike the mostly dark brown specimens from Zur- qui. Based on this color difference, I considered separating the La Cangreja specimens as a new spe- cies. Because no other differences were found, and because other species vary markedly in color, I abandoned this separation. DESCRIPTION. Body length 1.3- 1.5 mm. Frons yellow to brown, opaque. Mean frontal ratio 0.7. Color of flagellomere 1 yellowish at base, apically darker in male, female with flagellomere 1 yellow, pyriform. Palpus yellow. Palpus of male small, with setae normal, thick, moderately long, pointed. Pal- pus of female small, with setae normal, thick, mod- erately long, pointed. Dorsum of thorax yellow. Pleural regions yellow. Legs yellow to yellowish brown. Hind femur yellowish brown; evenly col- ored throughout. Mean costal ratio 0.56, range 0.54- 0.58. Mean costal sector ratio 4.36:2.98:1, range 3.33-6.33:2.50-3.67:1. Wing vein Rs slender, sub- equal to or thinner than costa. Wing vein R2+3 well developed. Halter yellow, or brown. Ventral setae of female segment 6 absent. Venter of male segment 6 bare. In cleared specimens, female dorsal abdom- inal glands invisible. Female Terminalia. Female intersegment 6-7 without distinct sclerite. Ovipositor triangular; dor- sally broad; anterodorsal portion narrowed, elon- gate; anteroventral margin smooth, without notch; posterodorsal apex of ovipositor rounded; in lateral view, posteroventral apex of ovipositor straight; posteroventral apex of ovipositor rounded. Stylet with lateral barbs. Dorsal sclerite broadly truncate. Male Terminalia. Epandrium globular, with an- terior part short, straight, with setae near cercus not markedly larger than other epandrial setae. Left side of epandrium with thick, ventral setae. Right side of epandrium with faint, unsclerotized dorsolateral ridge; in addition to ventral setae, with one to three lateral setae. Right surstylus short, rounded, with ventral carina short, extended partially across sur- stylus. Number of setae on right surstylus one to many. Cercus yellow. Setae of cercus and proctiger of normal size, longer and thicker than short setae of epandrium. NEW MATERIAL EXAMINED. COSTA RICA: Car- tago: La Cangreja, 9.8°N, 83.97°W, 73, vii.1991, 23, viii- ix.1991, 33, xi.1991, 33, iii— v.l 992, 63, 19, vi-vii.1992, P. Hanson, Malaise trap, 1950 m (CMNH, INBIO, LACM, MCZ, MICR); San Jose: Zurqui de Moravia, 10.05°N, 84.02°W, 23, vii.1992, P. Hanson, Malaise trap, 1600 m (LACM). Other A. fruncaficercus-Infragroup Specimens A single female of an as yet undescribed species (CMNH) was collected in Ecuadar. There probably are many such undescribed species in the Andes of South America. The material described in this paper raises the number of species of subgenus Mesophora to 31, in a group that originally contained only 6 species. I expect that further new species will be collected when other middle-elevation forests in South and Central America are surveyed. ACKNOWLEDGMENTS I am grateful to Julian Donahue, Paul Hanson, and Chen Young for allowing me to examine the material upon which this revision is based. I thank Brian Harris, Betty Oefibaugh, and Vicky Brown for technical assis- tance. Senor Jorge Arturo Lizano owns the property at Zurqui de Moravia; all entomologists are grateful to him for allowing us to collect insects on his land. This research was supported by the Natural History Museum of Los Angeles County Weiler Biodiversity Research Fund. LITERATURE CITED Borgmeier, T. 1937. Um nova especie de Apocephalus (Dipt. Phoridae), endoparasita de Chauliognathus fallax Germ. (Col. Cantharidae). Revista de Ento- mologia, Rio de Janeiro 7:207-216. Brown, B.V. 1992. Generic revision of Phoridae of the Nearctic Region and phylogenetic classification of Phoridae, Sciadoceridae and Ironomyiidae (Diptera: Phoridea). Memoirs of the Entomological Society of Canada 164:1-144. Brown, B.V. 1993. Taxonomy and preliminary phylog- eny of the parasitic genus Apocephalus , subgenus Mesophora (Diptera: Phoridae). Systematic Ento- mology 18:191-230. Brown, B.V. 1994. Life history parameters and new host records of phorid parasites of fireflies. Coleopterists ’ Bulletin 48:145-147. Coquillett, D.W. 1901. Apocephalus Coquillett, nov. gen. Proceedings of the Entomological Society of Washington 4:501. Disney, R.H.L. 1989. Scuttle flies — Diptera, Phoridae, genus Megaselia. Handbooks for the Identification of British Insects 10:1-155. Lloyd, J.E. 1973. Firefly parasites and predators. Cole- opterists ' Bulletin 27:91-106. Submitted 31 March 1994; accepted 25 May 1994. 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