AS try a/ 14 ISSN 0097-4463 ANNALS of CARNEGIE MUSEUM THE CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213 VOLUME 65 22 FEBRUARY 1996 NUMBER 1 CONTENTS ARTICLES Biology and immature stages of snail-killing flies belonging to the genus Tetanocera (Insecta: Diptera: Sciomyzidae). 1. Introduction and life his- tories of predators of shoreline snails B. A. Foote 1 New species of Rhyacophila from Virginia (Insecta: Trichoptera: Rhyacophilidae) Jan L. Sykora and Declan J. McCabe 1 3 New species of microcaddisflies from the eastern United States (Insecta: Trichoptera: Hydroptilidae) .... Steven C. Harris and Jan L. Sykora 17 A phylogenetic reassessment of the tadornine-anatine divergence (Aves: An- seriformes: Anatidae) Bradley C. Livezey 27 6 W6 Editors, ANNALS, BULLETIN and SPECIAL PUBLICATIONS: K. C. Beard B. C. Livezey D. R. Watters M. A. Schmidt, ELS, Assistant Editor Manuscripts, subscriptions, orders for individual numbers, and changes of address should be sent to: Office of Scientific Publications The Carnegie Museum of Natural History 4400 Forbes Avenue Pittsburgh, PA 15213-4080 Phone (412) 622-3287 Fax (412) 622-8837 E-mail: schmidtm@clp2.clpgh.org ANNALS OF CARNEGIE MUSEUM is published quarterly by The Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213-4080, by the authority of the Board of Trustees of Carnegie Institute. © 1996 Carnegie Institute. THE CARNEGIE MUSEUM OF NATURAL HISTORY @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 1, Pp. 1-12 22 February 1996 BIOLOGY AND IMMATURE STAGES OF SNAIL-KILLING FLIES BELONGING TO THE GENUS TETANOCERA (INSECTA: DIPTERA: SCIOMYZIDAE). L INTRODUCTION AND LIFE HISTORIES OF PREDATORS OF SHORELINE SNAILS B. A. Foote’ Research Associate, Section of Invertebrate Zoology Abstract An overview of the taxonomic composition, geographic distribution, and general biology of the family Sciomyzidae is presented. Literature dealing with the taxonomy and biology of the genus Tetanocera is summarized, and collecting, rearing, and preservation techniques are discussed. The natural histories, life cycles, and larval feeding habits of Tetanocera fuscinervis and T. silvatica, two species that prey on pulmonate snails occurring on wet shorelines, are presented. Introduction The acalyptrate Diptera family Sciomyzidae has a worldwide distribution and contains some 60 genera and 550 species (Knutson, 1987). The fauna of America north of Mexico consists of 21 genera and at least 175 species (Knutson et al., 1986). The biology of the family has been the focus of intensive study for over 40 years with the original impetus coming from the discovery of the snail-feeding habits of six species (Berg, 1953). Subsequently, numerous workers have de- scribed the immature stages and discussed the life histories, ecologies, and larval feeding habits of members of the family. Information is now available for over 200 species in the world, with particular attention to species occurring in the holarctic region. Larvae have been shown to prey on pulmonate aquatic snails, shoreline-inhabiting snails, terrestrial snails, snail eggs, slugs, and fingernail clams. Literature published before 1950 on the biology of Sciomyzidae has been discussed by Berg (1953) and Foote (1959). Berg (1961), Berg and Knutson (1978), and Knutson (1987) have summarized more recent findings. The genus Tetanocera has a holarctic distribution and contains some 40 species, with 30 species being recorded from the Nearctic region (Knutson, 1987). Keys and descriptions of the adults of species occurring in America north of Mexico are available in Steyskal (1959) and Orth and Fisher (1982). The present paper presents observations on the geographic distributions, habitat preferences, life histories, and larval feeding habitats of T. fuscinervis (Zetterstedt) and T. silvatica Meigen, two holarctic species having larvae that prey on pul- monate hygrophilous snails occurring in shoreline habitats. Subsequent papers will cover the biologies of species belonging to other trophic guilds, describe immature stages, and present a key to the third-instar larvae of the Nearctic species of Tetanocera. ' Department of Biological Sciences, Kent State University, Kent, Ohio 44240. Submitted 10 March 1995. 1 2 Annals of Carnegie Museum VOL. 65 Review of Literature The relatively few papers that have focused on the natural histories and larval feeding habits and/or described the immature stages of species of Tetanocera are discussed below. Dufour ( 1 849) apparently was the first worker to study the biology of a species of Tetanocera. He encountered a larva of T. ferruginea Fallen in mid-November, 1846, in floating mats of duckweed (Lemna sp.) and water- star wort (Callitriche sp.) in a small pond near Saint-Sever, France. This larva completed development in the laboratory and produced an adult on April 26, 1847. Dufour never observed larval feeding but did describe and illustrate the third-instar larva and puparium. Grtinberg (1910) described the habitat of the larvae of T. ferruginea in Europe as being among such aquatic plants as Lemna and Callitriche in slow-flowing water. He reported that pupae overwinter and that there are probably two or more generations a year. Brocher (1913) illustrated a larva of Tetanocera and stated that it resembled larvae of another sciomyzid genus, Sepedon, except that it was smaller and had much shorter lobes around the posterior spiracular disc. He reported that the larva left the water before forming a puparium and that the puparium did not float. Apparently, interspiracular processes (float hairs) were absent, but it is impossible to tell from his overly stylized drawing which species was involved. Lundbeck (1923) discovered floating puparia of 26 species of Sciomyzidae, including T. elata Fabricius, T. ferruginea, and T. silvatica, in marshes and ponds in Denmark during the spring months. He made no observations of the larvae, and his comment that ‘The larvae are phytophagous or feed on decaying vegetable matter, perhaps also being carnivorous on small objects ...” is unfounded. Lund- beck reared parasitoids belonging to a species of Ichneumonidae (subfamily Cryp- tinae) and wingless species of Proctotrupidae (Hymenoptera) from puparia of T. ferruginea. Johannsen (1935) described but did not illustrate the puparium of T. ferruginea from material collected amid vegetation along the margin of a shallow pond near Ithaca, New York. He noted the appearance of the anterior spiracles, and described the posterior spiracular disc and perianal pad (anal plate). He reported that the puparium of this species differed from those of species of Dictya in the appearance of the posterior spiracular disc and the length/width ratio of the perianal pad. Bhatia and Keilin (1937) described in considerable detail the larva of an un- determined species of Sciomyzidae that was obtained from a living land snail. Vertigo genesii Gredler, in England. They observed swallowing of air by the larva, a common sciomyzid habit, and noted that the larva imbedded itself in the prey snail until only the posterior spiracles were visible in the aperture of the shell. Their descriptions and drawings indicate that the larva was a member of the tribe Tetanocerini. The mouthhooks possessed four accessory teeth, the number usually found in larvae of Tetanocera, and the rows of “sensory papillae” found on the venter of each abdominal segment are recognizable in a few species of Tetano- cera. However, the presence of only five finger-like lobes (= papillae) on the anterior spiracles illustrated is a character not known for any Tetanocera larva. Berg (1953) made observations on T. ferruginea, T. fuscinervis (as unicolor Loew), and T. rotundicornis Loew. Larvae of the first species were free living and preyed on aquatic and shoreline snails. Newly hatched larvae of T. fuscinervis fed on pieces of a freshly killed Helisoma (Planorbidae), but died while still a 1996 Foote — Tetanocera Taxonomy and Biology 3 first instar. An adult of T. rotundicornis was reared from a larva that had infested a species of Oxyloma (Succineidae). He also presented evidence that overwinter- ing occurred as pupae in T. ferruginea and T. rotundicornis. In a later paper, Berg (1961) summarized the larval feeding habits and morphology of the immature stages of 84 species of Sciomyzidae, including 16 species of Tetanocera. Nielsen et al. (1954) discovered larvae of T. robusta Loew in wet areas in Iceland and suggested that the winter was passed as a pupa. They described and illustrated the third-iestar larva. Sods (1958) attempted to determine the number of generations produced each year for a variety of European species of Sciomyzidae by using data available on pinned specimens in museum collections. He reported that T. arrogans Meigen, T. hyalipennis Roser, and T. silvatica were bivoltine; whereas T. punctifrons Ron- dani and T. fuscinervis were univoltine. More recently, Berg et al. (1982) reported on seasonality, overwintering habits, and voltinism in north-temperate Sciomy- zidae. Species of Tetanocera were represented in three of the five distinct groups which they recognized. Berg and Neff (1959) discussed the use of larvae of certain predatory scio- myzids, including some species of Tetanocera, as biocontrol agents of aquatic snails serving as intermediate hosts of schistosome flukes that cause Schistoso- miasis in man. Foote (1961) discussed the life histories of 18 Nearctic species of Tetanocera and described and illustrated the eggs, larvae, and puparia of 12 species. Disney ( 1 964) reported that larvae of T. ferruginea were predators of pulmonate aquatic snails and described the larval habitat in England. He also recorded the ichneumonid wasp, Phygadeuon elegans Marshall, as a parasitoid of the pupa. Rozkosny (1965) described and illustrated the mature larva and puparium of T. ferruginea and the puparia of T. arrogans and T. elata. Later, Rozkosny (1967) discussed the biology and larval feeding habits of several European species of Sciomyzidae, including T. arrogans, T. elata, and T. ferruginea. He also described and illustrated the eggs, mature larvae, and puparia of these species. Although Foote (1963) briefly covered the slug-killing habits of two Nearctic species, T. plebeja Loew and T. valida Loew, the first detailed discussion of slug- killing habits in the genus was presented by Knutson et al. (1965). They presented information on the geographic distribution, habitat occurrence, life history, and larval feeding habits of T. elata, a Palearctic species. Other papers that discuss the biology of slug-killing species of Tetanocera are those of Trelka and Foote (1970) and Trelka and Berg (1977). Knutson (1963) did an extensive literature review of sciomyzid biology and taxonomy, gave detailed information on the natural history of 26 species of nine genera occurring in the Palearctic region, described the larvae and/or puparia of 18 species, and presented a key to the mature larvae and puparia of 19 species. He included information on seven species of European Tetanocera. Later, Knutson (1970) summarized the natural history of sciomyzid flies occurring in Sweden, including notes on 1 1 species of Tetanocera. Rozkosny (1966) summarized biological, ecological, distributional, and taxo- nomic information on the snail-killing flies of Czechoslovakia, including discus- sions of ten species of Tetanocera. Beaver (1972) discussed the life histories and larval feeding habits of eight species of Tetanocera occurring in England, including four species that also occur in North America. She reported that T. fuscinervis and T. silvatica had larvae that 4 Annals of Carnegie Museum VOL. 65 preyed on shoreline snails, whereas larvae of T. ferruginea and T. robusta attacked aquatic snails, Beaver (1973) presented oviposit! onal data for six European spe- cies, and later (1974a, 19741?) discussed intra-(four spp.) and interspecific (five spp.) competition in Tetanocera. Berg and Knutson (1978) reviewed the basic biology of sciomyzid flies, rec- ognizing eight trophic guilds in the family based on the feeding habits of the larvae. Gasc et al. (1984) described and illustrated the chorionic stucture of the egg of T. ferruginea, suggesting that the chorion served as a plastron in eggs that are deposited in habitats that are subject to immersion. As a result, embryonic de- velopment can continue even if eggs are completely immersed. Vala and Haab (1984) investigated the role of temperature and photoperiod in the development of T. ferruginea and the induction of pupal diapause. Manguin et al. (1985, 1988a, 19881?) and Manguin and Vala (1989) investigated the pred- atory behavior of T. ferruginea with respect to its foraging strategy, prey pref- erences, and the influence of snail biomass on numbers of prey consumed. Man- guin (1989) demonstrated sexual dimorphism in puparia and adults of T. ferru- ginea, noting that the sex of the adult could be predicted quite reliably by mea- suring the body length of the puparium. Vala and Gasc (1988) attempted to correlate the structure of the posterior spi- racular disc of sciomyzid larvae with their habitat preferences. They suggested that well-developed interspiracular processes (float hairs) and elongated peripheral lobes typified the aquatic predators, whereas these processes and lobes are greatly reduced in the more terrestrial species. Ferrar (1987a, 1987Z?) summarized knowledge of the biology and immature stages of the family Sciomyzidae, including several species of Tetanocera. Materials and Methods To avoid morphological descriptions and biological observations that may re- flect pecularities of local races, species of Tetanocera were reared from as many different geographic localities as possible. Bearings were initiated from material collected in Alaska, Colorado, Idaho, Montana, New York, Ohio, and Washington. Adults collected with a standard insect sweep net were placed in 8-dram shell vials and transported alive to the laboratory. Larvae of aquatic species in shallow- water habitats were found by submerging floating and emergent vegetation and allowing the larvae and puparia to float free. Many puparia were also found by searching through floating or stranded debris. Larvae of shoreline and terrestrial species were obtained by confining large numbers of field-collected snails and slugs in shallow plastic pans for at least ten days. The pans had a substrate of moist paper toweling and were covered by cheesecloth held in place by rubber bands. The pans were examined daily for dead and dying host individuals and for larvae and puparia. Snails that appeared to be infested were isolated in small petri dishes containing moist, shredded peat moss and examined daily for larvae. Bearings were maintained in large, well-lighted laboratories under room tem- perature conditions. Although relative humidities varied considerably in the lab- oratories, all rearings were held in small containers in which high humidities were maintained. Adults were confined in glass or plastic jars that were approximately 9.0 by 5.0 cm. The jars were covered with cheesecloth or fine screening and had a substrate of moist peat moss. Short lengths of cattail or short pieces of grass 1996 Foote — Tetanocera Taxonomy and Biology 5 served as resting sites for the adult flies. Small living snails or slugs were placed in each jar and a small pellet of a pasty mixture of honey and brewers’ yeast was appressed to the wall. In additional to the honey/yeast pellet, crushed snails were added to provide a protein source for the flies. Breeding jars were examined daily for eggs. These were removed with a fine camel’s hair brush and transferred to plastic petri dishes, with each dish receiving ten to 20 eggs. Eggs of terrestrial Tetanocera were placed on moist peat moss, and those of the more aquatic species were placed on short lengths of microscope slides floating in a small amount of water. Samples of eggs of each species were preserved in 10% formalin. When eggs of the aquatic species began to hatch, several small snails (under 5.0 mm) were added to the dishes. Dead snails were removed as they appeared and then replaced with living individuals. Cast exuviae were removed when pro- duced and preserved in 70% ethanol. Larvae of all three instars were killed in hot water and preserved in 70% ethanol. As puparia were formed, they were transferred to 8-dram shell vials containing a layer of moist peat moss and a cap of cheesecloth or fine screening. Most external structures of the larvae were examined without dissection, but the anterior spiracles were removed and mounted on microscope slides for detailed study. The anterior spiracles were macerated for 24 hr in a 10% solution of potassium hydroxide, rinsed in acid alcohol, dehydrated in a graded series of alcohol baths, and finally mounted on microscope slides in Canada Balsam or Permount. Cephalopharyngeal skeletons were recovered from cast exuviae, dis- sected out of preserved larvae, or removed from puparia. The skeletons were treated as described above and either mounted on microscope slides or studied directly in 70% ethanol by use of a dissecting microscope. All measurements were made with an American Optical Company ocular mi- crometer that had been calibrated with a stage micrometer. Drawings of eggs and larval structures were made by using an ocular grid of equal squares. Preliminary sketches were made on white paper that had been similarly gridded into equal- sized squares. Final drawings were made either on Strathmore drawing paper or on No. Wi Ross stipple board. Life History Studies Tetanocera fuscinervis (Zetterstedt), 1838; Insecta Lapponica Descripta: 737 Tetanocera fuscinervis is holarctic, occurring in Europe and Siberia as well as in North America. In the Nearctic region (Fig. 1), it is recorded from Newfound- land west to Alaska, and south to New York, Iowa, and Arizona (Knutson et al., 1986). Earlier American literature (e.g., Steyskal, 1959) referred to this species as T. unicolor Loew, 1847. In central New York, adults and puparia were found only in stands of emergent vegetation growing at the south end of the White Church Marsh located south of Ithaca. Adults were swept from a borrow pit bordering the abandoned railroad as well as along the south shore of the main marsh. Puparia were taken most abun- dantly floating amid the emergent sedges in the borrow pit. In northeastern Ohio, adults were taken only in an unshaded fen located in the Herrick Preserve in Portage County. In Colorado (three localities near Florissant in Teller County), Idaho (several localities in the northern half of the state), and Alaska (the Mata- nuska Valley and south of Anchorage), adults were collected in open or partially 6 Annals of Carnegie Museum VOL. 65 shaded marshes containing little water but supporting dense stands of emergent vegetation, especially species of Carex. This species was used in the first attempt to initiate a rearing of any sciomyzid by confining wild-caught flies for oviposition (Berg, 1953). Adults of T. fusci- 1996 Foote — Tetanocera Taxonomy and Biology 7 nervis {=T. unicolor) collected by Berg in an unshaded grass-sedge marsh five miles south of Anchorage, Alaska, on July 2, 1952, were confined in an experi- mental model of what has now evolved as a standard breeding jar. They mated soon after confinement and laid eggs on July 3 and 4, attaching them to damp sphagnum moss at the bottom of the container. Although motion inside the egg membranes was noticed on July 8, no hatching occurred on that day nor on the next. Some eggs had hatched by the morning of July 10; all, by noon of July 1 1. As there was then no knowledge that the minute first-instar larvae attack and kill living snails, they were offered pieces of freshly killed and chopped Helisoma subcrenatum (Carpenter). They fed sparingly and showed little evidence of de- velopment. Although the larvae survived well (up to 14 days), all died without molting into the second instar (Berg, 1953). Rearings were initiated by the present author from puparia taken at the White Church Marsh during March and April 1956 and from adults taken in June and early July 1958-60 in a small, partially shaded marsh located eight miles north of Sandpoint, Bonner County, Idaho (see Foote, 1961:148 for description of the Idaho site). Another rearing was initiated from adults collected on July 24, 1993, in an open grass-sedge marsh located one mile west of Florissant, Colorado. Many eggs were obtained from laboratory-reared and wild-caught females. On March 14, 1956, two females and a male that had emerged on March 13 were placed in a breeding jar. Mating was noticed first on March 19 and was seen frequently thereafter. Females began ovipositing on March 21, but laid only 16 eggs before dying on March 23. Another laboratory-reared female deposited 33 eggs between June 21 and July 1, 1958. Each of the females collected in nature in Idaho during June and early July produced between 70 and 204 eggs during periods ranging from eight to 13 days. Colorado-reared adults lived in the breed- ing jars from three to 21 days but did not oviposit. In the breeding jars, eggs were placed low on the glass jars, within two cen- timeters of the moss, on projecting sprigs of moss, and on shells of living and dead snails. One female deposited 95 eggs on the jar, 66 on projecting bits of moss, and 34 on shells of living Fossaria obrussa (Say). Those placed on moss were closely clustered in amorphous masses containing up to 30 eggs each. Eggs laid on the sides of breeding jars were more scattered and usually did not touch each other. Another female laid 108 eggs on the jar, three on shells, and none on moss. Under laboratory conditions, the incubation period was four to six days in the specimens collected in New York and Idaho but seven days in those from Alaska {n = 79: New York, 10; Idaho, 57; Alaska, 12). Larvae fed on small Gyraulus parvus (Say) and Helisoma spp. but developed more slowly than did other predaceous sciomyzid larvae reported in the literature. A larva that hatched on June 24 destroyed several small Helisoma, but did not molt into the second instar until July 1 . Seven other larvae remained in the second instar for nine to 1 1 days. Very few larvae reached the second instar, only four attained the third instar, and none formed a puparium. The causes of this heavy mortality rate are unknown. Knutson (1963) reared larvae of a European strain of T. fuscinervis (as T. unicolor) from hatching to pupariation, and did not mention a high mortality rate. However, only one adult emerged from his 19 puparia, and the puparia that were dissected contained mummified or decaying bodies. The 17 puparia found floating at the White Church Marsh from early March to April produced adults 11 to 16 days after being brought into the heated labo- ratory. Greatest emergence occurred in 11 to 12 days. 8 Annals of Carnegie Museum VOL, 65 In central New York, the earliest seasonal record for adults is June 10; the latest, September 2. Adults were collected occasionally throughout the summer months. Puparia were taken from early March to early April. In northern Idaho, the earliest record for adults is June 1; the latest, August 11. Because of the comparatively slow growth of the larvae, T. fuscinervis may have only one gen- eration per year. Overwintering evidently occurs in the pupal stage. Living puparia were collected on March 1 (more than three months before the earliest record for adults in this region) by breaking through the ice at the White Church Marsh. Two of eight puparia collected on March 12 produced ichneumonid wasps of an undetermined species on March 24. Tetanocera silvatica Meigen, 1830; Systematische Beschreibung der bekannten europaischen zweiflugeligen Insekten: 41. Tetanocera silvatica is a holarctic species that has a transcontinental distribution in Canada and the northern United States (Knutson et al, 1986), ranging from Alaska to Newfoundland and south to Minnesota, Colorado, and Arizona (Fig. 2). Adults of this species were most commonly encountered in permanent marshes or in marshy areas surrounding lakes. Other adults were taken in more temporary herbaceous wetlands, but none were encountered in wooded habitats. Near Flor- issant, Colorado, adults were taken on July 24, 1993, in association with T. fus- cinervis, Dictya montana Steyskal, Renocera johnsoni Cresson, Sepedon fuscT pennis Loew, and Limnia sandovalensis Fisher and Orth from a small (ca 1 acre), unshaded sedge marsh in which water levels had dropped noticeably as summer advanced. Adults of T. silvatica were especially common in somewhat drier, mar- ginal areas of the marsh. In Banff National Park on August 10, four adults that served as the basis of a laboratory rearing were swept from herbaceous vegetation growing in a floodplain marsh partially shaded by spruces. Small pools of open water one-half to one meter deep were scattered across the marsh. Three third- instar larvae of T. silvatica were found in shoreline debris around the shallow pools. Small aquatic snails of the genera Helisoma, Phy sella, and Gyraulus were abundant, and several individuals were crawling over the wet shoreline bordering the pools. Amphibious snails of the genera Catinella and Oxyloma were abundant on the shoreline vegetation. Another indication that the larvae are usually asso- ciated with shoreline habitats was my failure to recover larvae from floating veg- etation in western Alberta, central New York, northwestern Idaho, and central Colorado despite repeated attempts. Numerous matings in the breeding jars were noted at various times during the day between 10:00 a.m. and 9:30 p.m. The male’s position was similar to that observed in other species of the genus, except that his fore tarsi were placed close together on the middle of the female’s frons rather than lying along her eye orbits. The first 24 eggs were obtained on August 16 from the two females collected in nature on August 10. Seven eggs were affixed close together on the umbilical surface of a dead Helisoma, two were on the shell of a dead Lymnaea, six were placed on peat moss near a living Lymnaea, and the remaining nine eggs were scattered over the glass walls of the jar. A total of 71 eggs was deposited by these two females between August 16 and September 5. The incubation period lasted two to three days {n ^ 18), and newly hatched larvae were given small living Gyraulus and Lymnaea snails. As in T. fuscinervis, larval development in this species was somewhat prolonged, with the first stadium 1996 Foote — Tetanocera Taxonomy and Biology 9 lasting four to five days {n = 6). Another behavior that was somewhat unusual for species of Tetanocera was a tendency for the newly hatched larvae to feed gregariously within individual snails, with up to four larvae feeding together in a single Gyraulus. In contrast, second and third instars were more solitary in their 10 Annals of Carnegie Museum VOL. 65 feeding habits and did not occur together in any one snail. The second stadium was completed in five to six days {n = 3), but no larva managed to pupariate. Another rearing initiated by two females collected on July 24 in the marsh near Florissant, Colorado, was somewhat more successful in that adults emerged from two puparia obtained in the laboratory rearing. As in the earlier rearing, the newly hatched larvae commonly fed gregariously in such snails as small species of Gyraulus, Helisoma, Lymnaea, and Oxyloma, whereas older larvae fed singly. Between six and eight snails were consumed by two larvae during feeding periods of 22 and 27 days, respectively. The first stadium lasted four to six days {n = 3); the second, five days {n = 2); and the third, ten to 1 1 days {n = 2). The pupal period for a female was 17 days, whereas that of a male was 21 days. Mating among these reared adults began within three days after emergence, but no eggs were deposited. The earliest seasonal record for adults was obtained on May 20 (Minnesota); the latest, on August 17 (Alaska). Adults have been taken throughout the summer months in several localities, and it is probable that T. silvatica is at least bivoltine. The duration of the egg-to-egg period in the laboratory was 46 to 55 days, and there was no indication of diapause in any life stage. Discussion The habitat distribution of these two species of Tetanocera is quite similar in that both occur in open, largely unshaded herbaceous marshes with fluctuating water levels. More specifically, their larvae are associated with shorelines and can be found preying on stranded pulmonate aquatic and hygrophilic snails occurring in such situations. By frequenting shoreline habitats, the larvae probably escape competition from the more aquatic species of Tetanocera and other genera of Sciomyzidae. They probably do encounter competition from larvae belonging to other shoreline-dwelling genera of the family, such as species of Pherbellia, as well as from larvae of other species of Tetanocera. However, the specialization of the other species of shoreline Tetanocera on particular assemblages of gastro- pods probably mitigates interspecific competition within the genus. In contrast to such trophically specialized shoreline-dwelling species that prey on succineid snails {T. melanostigma Steyskal, T. oxia Steyskal, T. rotundicornis) or slugs {T. plebeja), larvae of T. fuscinervis and T. silvatica are rather generalized predators, frequently attacking stranded individuals of more aquatic species of Gastropoda. Competition for a limiting resource (stranded aquatic snails) may occur occa- sionally between T fuscinervis and T. silvatica. Adults of the two species com- monly occur in the same marsh at the same time, although the first species usually is more abundant. For example, the acre-sized marsh near Florissant, Colorado, supported a large population of T. fuscinervis, but a relatively small population of T. silvatica. Perhaps biotic (enemies) and abiotic (weather conditions) factors maintain populations of one or both species of Tetanocera below the carrying capacity of the marsh habitat such that there is only rarely a shortage of larval food. Acknowledgments Research was supported by grants from the National Science Foundation, National Institute of Allergy and Infectious Diseases, National Geographic Society, and the American Philsophical Society. Appreciation is expressed to G. C. Steyskal of Gainesville, Florida, for his expert help in determining species of Tetanocera. 1996 Foote — Tetanocera Taxonomy and Biology 11 This paper is dedicated to the memory of Dr. C. O. Berg, formerly of the Department of Entomology at Cornell University in Ithaca, New York. Literature Cited Beaver, O. 1972. Notes on the biology of some British sciomyzid flies (Diptera: Sciomyzidae). II. Tribe Tetanocerini. The Entomologist, 105:284-299. -. 1973. Egg laying studies on some British sciomyzid flies (Diptera: Sciomyzidae). Hydro- biologia, 43:1-12. — — — . 1974«. Laboratory studies on competition for food of the larvae of British sciomyzid flies (Diptera: Sciomyzidae. I. Intra-specific competition. Hydrobiologia, 44:443-462. . 1974^. Laboratory studies on competition for food of the larvae of some British sciomyzid flies (Diptera: Sciomyzidae). 11. Inter-specific competition. Hydrobiologia, 45:135-153. Berg, C. O. 1953. Sciomyzid larvae (Diptera) that feed on snails. Parasitology, 39:630-636. ■ 1961. Biology of snail-killing Sciomyzidae (Diptera) of North America and Europe. Ver- handlung der XI International Kongress ftir Entomologie, 1:197-202. Berg, C. O., and L. V. Knutson. 1978. Biology and systematics of the Sciomyzidae. Annual Review of Entomology, 23:239-258. Berg, C. O., and S. E. Neff. 1959. Preliminary tests of the ability of sciomyzid larvae (Diptera) to destroy snails of medical importance. Bulletin of the American Malacological Union, 25:11-13. Berg, C. O., B. A. Foote, L. Knutson, J. K. Barnes, S. L. Arnold, and K. Valley. 1982. Adaptive differences in phenology in sciomyzid flies. Memoir of the Entomological Society of Washington, 10:15-36. Bhatia, M. L., and D. Keilin. 1937. On a new case of parasitism of a snail (Vertigo genesii Gredl.) by a dipterous larva. Parasitology, 29:399^07. Brocher, F. 1913. L Aquarium de Chamber. Libraire Payot, Paris, 451 pp. Disney, R. H. L. 1964. A note on diet and habitats of the larva and an ichneumonid parasite of the pupa of Tetanocera ferruginea Fall. (Dipt., Sciomyzidae). Entomologist’s Monthly Magazine, 25: 88-90. Dufour, L. 1849. Histoire des metamorphoses du Tetanocera ferruginea. Annals of the Entomological Society of France, 7:67-79. Ferrar, P. 1987a. A Guide to the Breeding Habits and Immature Stages of Diptera Cyclorrhapha. Entomograph 8, Part 1: Text. E. J. Brill/Scandinavian Science Press, Copenhagen, 478 pp. — — — . 1987Z?. A Guide to the Breeding Habits and Immature Stages of Diptera Cyclorrhapha. Entomograph 8, Part 2: Figures. E. J. Brill/Scandinavian Science Press, Copenhagen, 429 pp. Foote, B. A. 1959. Biology and life history of the snail-killing flies belonging to the genus Sciomyza (Diptera: Sciomyzidae). Annals of the Entomological Society of America, 52:31-43. -- — — . 1961. Biology and immature stages of snail-killing flies belonging to the genus Tetanocera (Diptera: Sciomyzidae). Unpublished Ph.D. dissert., Cornell University, Ithaca, New York, 190 pp. Order No. 62-105, University Microfilms, Ann Arbor, Michigan (Dissertation Abstracts, 22: 3302-3303). — -. 1963. Biology of slug-killing Tetanocera (Diptera: Sciomyzidae). Proceedings of the North- central Branch of the Entomological Society of America, 18:97. Gasc, C., j. Vala, and j. Reidenbach. 1984. Etude comparative au microscope electronique a ba- layage structures chorioniques d’oeufs de cinq especes de Sciomyzidae a larves terrestres et aquatiques (Diptera). Annals de la Societe Entomologie de France, 20:163-170. GrUnberg, K. 1910. Diptera, Zweifliiger. Pp. 1-312, in Die Siisswasserfauna Deutschlands. Heft 2A (A. Brauer, ed.), Jena. Johannsen, O. a. 1935. Aquatic Diptera IT Orthorrhapha-Brachycera and Cyclorrapha. Memoir of the Cornell University Agricultural Experiment Station, 177:1-62. Knutson, L. V. 1963. Biology and immature stages of snail-killing flies of Europe (Diptera: Scio- myzidae). Unpublished Ph.D. dissert., Cornell University, Ithaca, New York, 390 pp. Order No. Mic 63-4822, University Microfilms, Ann Arbor, Michigan (Dissertation Abstracts, 24:445^46). — — — . 1970. Biology of snail-killing flies in Sweden (Dipt., Sciomyzidae). Entomologica Scandi- navica, 1:307-314. — . 1987. 84. Sciomyzidae. Pp. 927-940, in Manual of Nearctic Diptera, Volume 2 (J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, coords.). Research Branch, Agriculture Canada Monograph 28, Ottawa, 675-1332 pp. Knutson, L. V, J. W. Stephenson, and C. O. Berg. 1965. Biology of a slug-killing fly, Tetanocera data (Diptera: Sciomyzidae). Proceedings of the Malacological Society of London, 36:213-220. 12 Annals of Carnegie Museum VOL. 65 Knutson, L. V., R. E. Orth, T. W. Fisher, and W. L. Murphy. 1986. Catalog of Sciomyzidae (Diptera) of America north of Mexico. Entomography, 4:1-53. Lundbeck, W. 1923. Some remarks on the biology of Sciomyzidae, together with the description of a new species of Ctenulus from Denmark (Dipt.). Viddenskabelige Meddelelser fra Dansk Na- turhistoriske Forening I Kpbenhaven, 76:101-109. Manguin, S. 1989. Sexual dimorphism in size of adults and puparia of Tetanocera ferruginea Fallen (Diptera: Sciomyzidae). Proceedings of the Entomological Society of Washington, 91:523-528. Manguin, S., and J. Vala. 1989. Prey consumption by larvae of Tetanocera ferruginea (Diptera: Sciomyzidae) in relation to number of snail prey available. Annals of the Entomological Society of America, 82:588-592. Manguin, S., J. C. Vala, and J. M. Reidenbach. 1985. Predation de mollusques dulqaquicoles par les larves malacophages de Tetanocera ferruginea Fallen, 1820 (Diptera, Sciomyzidae). Canadian Journal of Zoology, 64:2832-2836. . 1988a. Action predatrice des larves de Tetanocera ferruginea (Diptera: Sciomyzidae) dans des systemes a plusieurs especies de mollusques-proies. Acta Oecologica/Oecologia Applicata, 9: 249-259. -. 1988£>. Determination des preferences alimentaires des larves de Tetanocera ferruginea (Dip- tera: Sciomyzidae), predateur de mollusques dulgaquicoles. Acta Oecologica/Oecologica Appli- cata, 9:353-370. Nielsen, P. O., O. Ringdahl, and S. L. Tuxen. 1954. The Zoology of Iceland. Volume III, Part 48a: Diptera I. Ejnar Munksgaard, Copenhagen, 189 pp. Orth, R. E., and T. W. Fisher. 1982. A new species of Tetanocera Dumeril from Colorado (Diptera: Sciomyzidae). Proceedings of the Entomological Society of Washington, 84:685-689. Rozkosny, R. 1965. Neue Metamorphosestadien mancher Tetanocera-oxicn (Sciomyzidae, Diptera). Zoologische Listy, 14:367-371. . 1966. Ceskoslovenske druhy malakofagni celedi Sciomyzidae (Diptera) (The Czechoslovak- ian species of the malacophagous family Sciomyzidae). Folia Facultatis Scientiarum Naturalium Universitatis Purkynianae Brunensis Biologia, 7:1-111. . 1967. Zur Morphologie und Biologic der Metamorphosestadien mitteleuropaischer Sciomy- ziden (Diptera). Acta Scientiarum Naturalium Academiae Scientiarum Bohemoslovacae Brno I, 4:117-160. Soos, A. 1958. 1st das Insektenmaterial der Museen fiir ethologische und okologische Untersuchungen verwandbar? Angaben iiber die Flugzeit und die Generationszahl der Sciomyziden (Diptera). Acta Entomologische Museum National Prague, 32:101-150. Steyskal, G. C. 1959. The American species of the genus Tetanocera Dumeril (Diptera). Papers of the Michigan Academy of Science, Arts, and Letters, 44:59-91. Trelka, D. G., and C. O. Berg. 1977. Behavioral studies of the slug-killing larvae of two species of Tetanocera. Proceedings of the Entomological Society of Washington, 79:475^86. Trelka, D. G., and B. A. Foote. 1970. Biology of slug-killing Tetanocera. Annals of the Entomo- logical Society of America, 63:877-895. Vala, J., and C. Gasc. 1988. Ecological adaptations and morphological variation in the posterior disc of larvae of Sciomyzidae (Diptera), Canadian Journal of Zoology, 68:517-521. Vala, J., and C. Haab. 1984. Etude experimentale du developement larvaire de Tetanocera ferru- ginea Fallen, 1820. Influences de la temperature et de la photoperiode, diapause pupale, biomasse alimentaire (Diptera: Sciomyzidae). Bulletin et Annales de la Societe Royale Entomologie de Belgique, 120:165-178. ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 1, Pp. 13-16 22 February 1996 NEW SPECIES OF RHYACOPHILA FROM VIRGINIA (INSECTA: TRICHOPTERA: RHYACOPHILIDAE) Jan L. Sykora Research Associate, Section of Invertebrate Zoology Declan J. McCabe’ Abstract A new species of rhyacophilid caddisfly from Virginia, Rhyacophila tricornuta is described and figured. This new species is related to R. nigrita, but differs from all other known species of R. nigrita subgroup mainly by the shape of the dorsoapical section of Segment IX bearing three short and rounded lobes. Introduction The genus Rhyacophila is widely distributed throughout the holarctic region. The Nearctic region supports rich and diverse populations of Rhyacophila that are included in several indigenous groups and complexes. One-hundred-twenty- six species were reported from North America including Greenland and Mexico (Morse, 1993). The R. invaria group includes 14 species all restricted to the eastern United States. Larvae in this group inhabit small, cold rivers and streams in the Appalachian region. According to Schmid (1970) within this group two subgroups could be easily distinguished, the R. invaria subgroup and the R. nigrita subgroup. Until now the R. nigrita subgroup included six closely related species that could be identified based on male genitalia characterized by transverse and high-crested abdominal Segment X, absence of horizontally produced lateral winglets on anal sclerites, and notched dorsoapical margin of Segment IX. In collections from Smyth County, Virginia, a new species of Rhyacophila in the R. nigrita subgroup has been recently discovered, and is described in this paper. Systematic Entomology Rhyacophila tricornuta, new species (Fig. 1-3) Diagnosis. — This species is characterized by its male genitalia. The unique features separating this species from other American Rhyacophila are three dor- soapical, rounded lobes on Segment IX and a dark, rounded process extended into an “undulated” apicolateral section located at the base of the anal sclerites. Description. — Male: length 12 mm. General appearance of specimen in alcohol brown. Head, thorax, and antennae black; legs yellowish; abdomen brown with darker terminal Segments VII, VIII, and IX. Ventromesal projection on Segment VII very short and smaller than in R. nigrita, corresponding in size to ventral ’ Department of Biological Sciences, University of Pittsburgh, Pittsburgh, Pennsylvania 15260; current address: Department of Biology, University of Vermont, Burlington, Vermont 05405. Submitted 19 April 1995. 13 14 Annals of Carnegie Museum VOL. 65 iit 1996 Sykora and McCabe — New Rhyacophila from Virginia 15 ‘‘dent” found on Segment VI in the latter species. Wing venation and general structure typical for genus. Male genitalia: Segment IX annular, narrow ventrally, wide dorsal section with double emargination forming lateral notch producing pair of dorsolateral lobes and very short, rounded dorsoapical process not reaching base of Segment X. Segment X with emarginated dorsoapical section resulting in V-shaped, triangular notch; from lateral view forming high-crested, rounded lobes covered with short bristles. Base of Segment X narrow with heavily sclerotized small, dark lateral lobes extending dorsoapically into an “undulated,” dark brown sclerite. Anal sclerites similar to those of R. nigrita with rounded apices from lateral and dorsal views. Inferior appendages each with basal segment as broad as long, apical seg- ment bilobed; tapered ventral lobe bearing narrow band of conspicuous dark spic- ules on dorsoapical margin; the ventral lobe 2.5 times as long as apically rounded dorsal lobe. Phallic apparatus large, mostly membranous, and similar to that of R. nigrita; the spiniform parameres straight and long; aedeagus sinuous, sclerotized and up- turned at apex. Female: unknown. Type Specimen. — Holotype, male, Virginia: Smyth Co., Mount Rodgers Na- tional Recreation Area, Hurricane Campground next to Hurricane Creek, May 22, 1994, Declan J. McCabe. The holotype is deposited at the Carnegie Museum of Natural History. Etymology. — Latin, meaning “three horns.” Discussion Rhyacophila tricornuta is a member of the R. invaria group and closely related to species included in the R. nigrita subgroup. This is based on similarity of several morphological features but especially on the shape of Segment X with transverse high crest and inferior appendages similar to those in R. nigrita and R. acutiloba (Schmid, 1970; Morse and Ross, 1971). However, the characteristic which separates this new species from all members of the R. nigrita group is the shape of the dorsoapical margin of Segment IX which in the R. invaria subgroup is extended into a single, solid or more-or-less forked dorsomesal lobe. In six species of the R. nigrita subgroup the dorsoapical margin of Segment IX is notched posteriorly in the center with no distinct lobes or projections. It is also of interest that the ventromesal projection on Segment VII is extremely short in R. tricornuta (30 fim). In males of R. nigrita the same process is usually over 60 jjim in length while the rudimentary ventromesal projection on Segment VI is approximately 20 jxm long. This process is missing on Segment VI of R. tricor- nuta. These morphological characteristics, especially the three-lobed Segment IX, set R. tricornuta apart from any other known species in the R. invaria group. How- ever, in order to define the placement of this species within R. nigrita subgroup, additional information on female morphology and possibly on immature stages is needed. Fig. 1-3. — Male genitalia of Rhyacophila tricornuta, n. sp. 1, left lateral view; 2, dorsal view; 3, phallic apparatus, ventral view. 16 Annals of Carnegie Museum VOL. 65 A single specimen of R. tricornuta was captured in a black-light trap near the confluence of two second-order streams — an unnamed tributary and Hurricane Creek. Hurricane Creek flows east in the valley formed between Hurricane Moun- tain and Bear Ridge while the unnamed creek flows west in the valley formed between Hurricane Mountain and Bobbys Ridge. The two combine at 36°42'45"N 81°42'45"W, and the resulting third order stream is called Comers Creek which flows north between Bobbys Ridge and Bear Ridge. The confluence is at elevation 852 m (2795 ft) in the Iron Mountains. The trap was operated continuously from shortly before nightfall until midnight. Additional Trichoptera species collected in the light trap included Goera fuscula Banks, Glossosoma nigrior Banks, and Dolophilodes distinctus (Walker). The following caddisfly larvae were collected by hand from the creeks in the sampling area; Rhyacophila Carolina Banks, R. fuscula Banks, Agapetus sp., Neophylax sp., Dolophilodes distinctus (Walker), and Pycnopsyche luculenta Betten. The streams in the area also supported a di- verse population of stoneflies, mayflies, and blepharicerid flies including species of Acroneuria, Peltoperla, Isoperla, Ephemerella, Epeorus, and Blepharicera. Based on this information the larvae of this new species inhabit second-order streams or seepage areas at higher elevations of the Appalachians in Virginia. Acknowledgments We are grateful to Dr. John S. Weaver III for reviewing the figures. Literature Cited Morse, J. C. 1993. A checklist of the Trichoptera of North America including Greenland and Mexico. Transactions of the American Entomological Society, 119:4-93. Morse, J. C., and H. H. Ross. 1971. Two new species of Rhyacophila from eastern North America. Journal of the Kansas Entomological Society, 44:403-405. Schmid, E 1970. Le genre Rhyacophila et la famille des Rhyacophilidae (Trichoptera). Memoires de la Societe Entomologique du Canada, 60:1-230. ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 1, Pp. 17-25 22 February 1996 NEW SPECIES OF MICROCADDISFLIES FROM THE EASTERN UNITED STATES (INSECTA: TRICHOPTERA: HYDROPTILIDAE) Steven C. Harris' Research Associate, Section of Invertebrate Zoology Jan L. Sykora Research Associate, Section of Invertebrate Zoology Abstract Three new species in the genus Hydroptila and one in the genus Mayatrichia from the eastern United States are described and illustrated: Hydroptila koryaki and H. fowlesi from West Virginia, H. homochitta from Mississippi, and M. tuscaloosa from Alabama. Introduction Although the caddisfly fauna of the eastern United States has been fairly well studied, ongoing collecting continues to reveal undescribed species. This paper describes four new species of microcaddisflies from West Virginia, Mississippi, and Alabama, three in the genus Hydroptila and one in the genus Mayatrichia. The Hydroptilidae, or microcaddisflies, include the smallest species of Trichop- tera, ranging in size from 1.2 to 6 mm. They are also the most speciose of all North American families with over 300 species (Morse, 1993), nearly one-third of which are in the genus Hydroptila. The genus Mayatrichia is much less species- rich with only five species known from North America (Morse, 1993). Terminology used in the descriptions follows that of Marshall (1979). Specimen length was measured from the tip of the head to the end of the wings and is given as a range when more than one specimen was available. Type material will be deposited at the Carnegie Museum of Natural History (CMNH); the National Museum of Natural History, Smithsonian Institution (NMNH); the Illinois Natural History Survey (INHS); the Florida State Collection of Arthropods (FSCA); Mis- sissippi State University, Department of Entomology (MSU); and the collections of the authors (SCH and JLS). Systematic Entomology Hydroptila koryaki, new species (Fig. 1-^) Diagnosis. — This species, a member of the H. waubesiana group, superficially resembles H. ouachita Holzenthal and Kelley and H. cheaha Harris in the spinose eighth segment and forked tenth tergum. Members of the H. waubesiana group, in general, are characterized by a short ventromesal process on Segment VII and a highly modified tenth tergite which may bear sclerotized hooks, elongate pro- cesses, curved rods, or sclerotized strips. The new species is easily recognized by ’ Department of Biology, Clarion University, Clarion, Pennsylvania 16214. Submitted 28 April 1995. 17 18 Annals of Carnegie Museum VOL. 65 Fig. 1^. — Hydroptila koryaki n. sp., male genitalia. 1, left lateral view; 2, ventral view; 3, dorsal view; 4, phallus, ventral view. the thin, curved inferior appendages which are fused mesally and the elongate tenth tergum which is hooked in lateral aspect. Description. — Male: length 2.6 mm. Antennae 26-segmented. Color brown in alcohol. Sternite of abdominal Segment VII with short apicomesal process. Seg- ment VIII narrowing posteriorly in lateral view, bearing numerous heavy setae at the apical margin; slightly emarginate in ventral view; deeply incised dorsally 1996 Harris and Sykora — New Microcaddisflies 19 with pair of small setose lobes, heavy spines laterally. Segment IX completely retracted within Segment VIII, generally quadrate in shape, incised slightly both anteriorly and posteriorly. Segment X elongate, extending nearly to posterior Seg^ ment VII; in lateral view narrowing apically to sclerotized, downturned hook, with dorsal hump subapically; in dorsal view narrow basally, divided apically into pair of thin, sinuate processes. Inferior appendages thin and elongate in lateral aspect, curving dorsad; in ventral view fused mesally, narrow basally, divided into two diverging arms distally each with sclerotized point on outer apical mar- gin, pair of stout setae subapically; in dorsal view basally with narrow sclerotized bands laterally which join anteriorly. Phallus long and narrow, paramere encircling shaft before midlength; ejaculatory duct protruding at apex. Female: unknown. Type Specimens. — Holotype, male (CMNH): West Virginia, Lewis County, Right Fork of the West Fork River, Walkers ville, 17 July 1993, light trap, J. L. Sykora. Paratypes: same, 13 July 1992, Shawn Campbell, 3 males (NMNH, SCH, JLS). Etymology.- — -Named for Michael Koryak from the U. S. Army Corps of En- gineers who selected the site where this species was collected. Remarks. — Under the dissecting microscope (SOX), Segment VIII in lateral view appears to narrow distally much as in H. waubesiana Betten, but this is an illusion created by the lateral curving of the segment. The species was collected adjacent to a small, slow-flowing stream composed mostly of pools with very few riffles. The stream is affected by human activity as it flows through a rural community and by farming activites, as evidenced by open grazing, erosion, and possibly animal wastes. Other species of Hydroptila collected at this site included H. amoena Ross, H. grandiosa Ross, H. hamata Morton, and H. fowlesi n. sp. Hydroptila fowlesi new species (Fig. 5-8) Diagnosis.^ — ^This species, a member of the H. tineoides group of Marshall (1979), is most similar to H. sandersoni Mathis and Bowles and other members of the H. amoena complex. Members of this complex have in common a long apicomesal process from the seventh sternum, beak-like inferior appendages, and a phallus divided into two apical processes. The new species is distinguished by the triangular shape of the tenth tergum and by the structure of the phallus, which is divided into apical processes unequal in length. Description.— M.^\c: length 2.8 mm. Antennae 27-segmented. Color brown in alcohol. Venter of abdominal Segment VII with long apicomesal process. Segment VIII annular, with venter longer than dorsum. Segment IX deeply excised ante- riorly in dorsal and ventral views; laterally with thin posterior lobe bearing nu- merous setae along posterodorsal and ventral margins, apodeme long and narrow extending to middle of Segment VII; square in dorsal view, incised at margins; in ventral view shallowly incised on posterior margin, bearing numerous setae laterally and submesally; internally with bilobed process bearing pair of setae posteriorly. Tergite X triangular, wide basally tapering to rounded apex, anteriorly fused to Segment IX; in lateral view sharply upturned basally. Inferior appendages beak-like, each with dorsolateral thumb-like projection bearing three setae on posterior margin, ventral projection tapering ventrad to sharp apex; fused along meson with wing-like lateral extensions in ventral view. Phallus with basal and 20 Annals of Carnegie Museum VOL. 65 Fig. 5-8. — Hydroptila fowlesi n. sp., male genitalia. 5, left lateral view; 6, ventral view; 7, dorsal view; 8, phallus, ventral view. 1996 Harris and Sykora — New Microcaddisflies 21 apical portions equal in length, with short paramere making nearly one revolution at midlength, apical portion divided into two thin processes; process bearing pro- truding ejaculatory duct straight and twice as long as sheath-like second process. Female: unknown. Type Specimen. — Holotype, male (CMNH): West Virginia, Lewis County, Right Fork of the West Fork River, Walkersville, 13 July 1992, light trap, Shawn Campbell. Etymology. — Named for J. M. Fowles from the U. S. Army Corps of Engineers who organized the light trap sampling program. Remarks. — That several species having a phallus divided into two straight api- cal processes would key to H. amoena was first recognized by Harris (1985) with the description of two closely related species. Two additional species were de- scribed by Mathis and Bowles (1990), who also presented a key to the five species in the complex and discussed characteristics separating the species. With the ad- dition of another species, the key of Mathis and Bowles is appended as follows: Amended Key to H. amoena complex 1. Aedeagus short, less than one-third as long as abdomen; phallic process lacking ejaculatory duct with apex flattened and pointed (Mathis and Bowles, 1990:fig. 4) H. artesa la. Aedeagus longer, greater than one-third as long as abdomen; apical process not as above (Fig. 8) ........................ . .................. ...... 2 2. Apex of Tergite X entire (Fig. 1) ... 2a Apex of Tergite X excised (Harris, 1985:fig. 8E) 3 2a. Tergite X narrow, nearly parallel-sided basally in dorsal view (Mathis and Bowles, 1990;fig. 9); Segment X directed caudad in lateral view (Mathis and Bowles, 1990:fig. 10); phallus with distal portion at least twice as long as base, apical processes equal in length (Mathis and Bowles, 1990:fig. 11) . . H. sandersoni 2b. Tergite X wide basally, tapering distally in dorsal view (Fig. 7); Segment X upturned in lateral view (Fig. 5); phallus with distal portion equal in length to base, apical processes unequal in length (Fig. 8) .H. fowlesi Hydroptila homochitta, new species (Fig. 9-12) Diagnosis. — This species, a member of the H. waubesiana group of Marshall (1979), is distinguished from the similar H. cretosa Harris and H. lonchera Blickle and Morse by the long, thin processes from the venter of Segment X. Such in- termediate appendages are found in several southeastern members of the H. wau- besiana group, including H. oakmulgeensis Harris, H. tridentata Holzenthal and Kelly, and H. poirrieri Holzenthal and Kelley, but in these species the structures tend to be sickle-shaped rather than long and narrow as in H. homochitta. The club-shaped, serrate apex of the tenth tergum is also diagnostic for the new spe- cies. Description. — Male: length 2. 3-2. 6 mm. Antennae 25-segmented. Color brown in alcohol. Venter of Segment VII with short, posteromesal process. Segment VIII rounded posteriorly in lateral view; in dorsal view deeply emargioate posteriorly; slightly emarginate on posterior margin in ventral aspect. Segment IX retracted within Segment VIII, rounded anteriorly in lateral view; in dorsal and ventral view anterior margin deeply incised, truncate posteriorly, and fused with Segment X. Tergum X rectangular in dorsal view, cleft distally with sclerotized spinal process at apex; in lateral view wide basally, narrowing at midlength, then wid- ening to rounded apex with numerous sclerotized serrations on posterodorsal mar- 22 Annals of Carnegie Museum VOL. 65 Fig. 9-12. — Hydroptila homochitta n. sp., male genitalia; 9, left lateral view; 10, ventral view; 11, dorsal view; 12, phallus, ventral view. 1996 Harris and Sykora — New Microcaddisflies 23 gin, ventral margin produced into thin, elongate sclerotized processes (interme- diate appendages) extending beyond Tergum X and curving ventrad. Inferior ap- pendages lightly sclerotized; in lateral view nearly parallel-sided, dorsal margin straight, ventral margin irregular and bearing numerous setae, narrowing apically to rounded tip; in ventral view wide basally and gradually tapering distally, scler- otized points apically and subapically on outer margin, series of stout setae di- rected laterad on ventral surface. Phallus long and narrow, widening basally and subapically, ejaculatory duct protruding at apex, thin paramere encircling shaft at midlength. Female: unknown Type Specimens. — Holotype, male (CMNH): Mississippi, Franklin County, Por- ter Creek, (T5N, R4E, S8NW), 29 June 1992, T. Schiefer and R. Fontenot. Para- types, same, 16 males (NMNH, INHS, CMNH, MSU, SCH, JLS); same, but 27 July 1992, 7 males (NMNH, INHS, FSCA); Franklin County, Tributary to McGehee Creek, (TGN, R4E, S26SW), 1 June 1992, T. Schiefer and R. Fontenot, 1 male, (NMNH). Etymology.- — Named for the Homochitto National Forest, collection locality for the species. Remarks. — The streams of the Homochitto National Forest where the new spe- cies was collected are all small, clear, and sandy-bottomed, flowing through rel- atively undisturbed mixed hardwood forests. Mayatrichia Tuscaloosa, new species (Fig. 13-16) Diagnosis. — Although similar to M. ponta Ross in the structure of the inferior appendages with three heavy setae at the apex, M. tuscaloosa differs in the brac- teoles being entire posteroventrally; a character which is shared with M. illobia Harris and Holzenthal and M. ayama Mosely. The new species is separated from the widespread M. ayama and M. illobia, which is only known from Costa Rica, on the basis of the rectangular inferior appendages. Description. — Male: length 2. 3-2.4 mm, antennae 18-segmented. Color brown in alcohol. Sternum VI with short posteromesal process. Segment VIII annular. Segment IX tapering anteriorly in lateral view; posterolateral processes (bracte- oles) narrow; ventral margin straight, entire; dorsal margin rounded; in dorsal view, Tergum IX deeply emarginate posteriorly, line of setae at margin, anteriorly with shallow, rounded incision; in ventral view bracteoles elongate and narrow, curving inward distally. Tergum X membranous, rounded apically; in lateral view triangular, indistinct. Inferior appendages in lateral aspect wide basally, narrowing distally, each bearing elongate setae; in ventral view nearly parallel-sided, each with three teeth bearing stout, elongate setae. Subgenital plate with posteromesal process directed ventrad; in ventral view truncate, slightly emarginate mesally, bearing pair of small setae on posteromesal margin. Phallus long and narrow, apex with small, dorsal projection. Female: unknown. Type Specimens. — Holotype, male (CMNH): Alabama, Tuscaloosa County, Big Sandy Creek at 4.5 mi south of Coaling on an unnumbered county road, 20 March 1991, S. Harris and T. Shepard. Paratype: same, 1 male (NMNH). Etymology. — Named for Tuscalosa County, Alabama. Remarks. — The species was found only in a single early spring collection. A 24 Annals of Carnegie Museum VOL. 65 Fig. 13-16, — Mayatrichia tuscaloosa n. sp., male genitalia. 13, left lateral view; 14, ventral view; 15, dorsal view; 16, phallus, lateral view. 1996 Harris and Sykora — New Microcaddisflies 25 collection made the following night and monthly collections through October at the same site failed to collect additional specimens. Collections made in March and April of the following year were also unsuccessful. The collection locality was at the headwaters of Big Sandy Creek in a transition zone where the stream emerged from a springfed limestone region out onto the coastal plain. Acknowledgments Mississippi specimens were obtained from Richard L. Brown of Mississippi State University and were collected as part of NSF grant BSR-9200856. West Virginia specimens were collected as part of contract DACW 59-92-M-0422 to JLS by the U. S. Army Corps of Engineers. The help of Tom Shepard of the Geological Survey of Alabama in the collection of the Alabama specimens is gratefully acknowledged. Literature Cited Harris, S. C. 1985. New microcaddisflies (Trichoptera: Hydroptilidae) from Alabama. Proceedings of the Entomological Society of Washington, 87:606-621. Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bulletin of the British Museum of Natural History (Entomology), 39:135-239. Mathis, M. L., and D. E. Bowles. 1990. Three new species of microcaddisflies (Trichoptera: Hy- droptilidae) from the Ozark Mountains, U.S.A. Proceedings of the Entomological Society of Washington, 92:86-92. Morse, J. C. 1993. A checklist of the Trichoptera of North America, including Greenland and Mexico. Transactions of the American Entomological Society, 119:47-93. ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 1, Pp. 27-88 22 February 1996 A PHYLOGENETIC REASSESSMENT OF THE TADORNINE-ANATINE DIVERGENCE (AVES: ANSERIFORMES: ANATIDAE) Bradley C. Livezey Associate Curator, Section of Birds Abstract Phylogenetic analysis of eight problematic genera of waterfowl {Stictonetta Plectropterus, Sarki- diornis, Hymenolaimus, Tachyeres, Merganetta, Malacorhynchus, and Salvadorina) and representa- tives of other tadornine genera and anatine tribes was undertaken using 114 morphological characters (39 skeletal, four tracheal, 1 1 natal, and 60 of the definitive integument) and a variety of weighting schemes, in an attempt to clarify relationships in the most poorly understood segment of the phylogeny of Anseriformes (Livezey, 1986<3). Under equal weighting of characters, 40 equally parsimonious trees were found that shared the following findings: (1) a sister-group relationship of Stictonetta to a clade comprising the Tadorninae and Anatinae; (2) a sister-relationship between Plectropterus and Sarki- diomis; (3) monophyly of the typical shelducks and sheldgeese (Tadorna, Cyanochen, Alopochen, Neochen, and Chloephaga); (4) monophyly of Hymenolaimus, Tachyeres, and Merganetta-, and (5) a sister-relationship between Salvadorina and Malacorhynchus. An analysis wherein skeletal characters were given twice the weight of other character groups resulted in four shortest trees which, in addition to the findings under equal weighting, indicated that: (1) Plectropterus + Sarkidiornis represent the sister-group of the typical shelducks and sheldgeese; (2) a sister-relationship between the clade com- prising Hymenolaimus, Tachyeres, and Merganetta and the preceding clade, collectively constituting the “tadornine” clade; and (3) a sister-relationship between Malacorhynchus P Salvadorina and an “anatine” clade comprising the remaining tribes and subtribes (Cairineae, Nettapodeae, Anateae, Ay- thyini, Mergini, and Oxyurini). An analysis in which skeletal, tracheal, natal, and definitive characters were assigned weights of four, three, two, and one, respectively, produced identical results to the preceding weighting scheme. Successive weighting by rescaled consistency indices produced two shortest trees that resembled those found under the two preceding protocols except: (1) the Mergini were placed as the sister-group of the “tadornine” clade defined above; and (2) the Cairineae (Cairina, Pteronetta, and Aix) were placed as the sister-group to Plectropterus + Sarkidiornis. Based on the comparative strengths of hypothesized homologies of skeletal characters and assessments of topolog- ical support (including bootstrapping, decay indices, and near-shortest trees) for all weighting schemes, a conservative classification was based on the a priori weighting schemes favoring skeletal characters. Maps of selected ecomorphological attributes on this phylogenetic tree revealed several distinct evo- lutionary trends, but incompletely resolved relationships among these diverse genera and within su- prageneric taxa limited inferences. Additional promising areas for anatomically based systematic study are identified, and the need for continued cladistic study of this poorly resolved but topologically critical segment of the phylogeny of Anseriformes is emphasized. Introduction Greater diversity of opinion has attended the classification of the following eight southern-hemisphere genera of waterfowl than in any assemblage of similar size in the Anseriformes: Stictonetta, Plectropterus, Sarkidiornis, Hymenolaimus, Tachyeres, Merganetta, Malacorhynchus, and Salvadorina (Table 1). Early tax- onomists typically included these genera within larger, more-familiar groups of waterfowl on the basis of superficial comparisons and very limited material (Eyton, 1838; Gray, 1841, 1871; Bonaparte, 1856; Sclater, 1880; Salvador!, 1895; Phillips 1922, 1923, 1925, 1926). In recent decades, taxonomic classifications of the Freckled Duck {Stictonetta Submitted 16 June 1995. 27 28 Annals of Carnegie Museum VOL. 65 Table 1. — A century of taxonomic classifications of eight taxonomically problematic genera of Anatidae. Genus Salvadori (1895) Phillips (1922-1925) Peters (1931) Stictonetta Anatinae, between Mar- maronetta and Hetero- netta. Anatinae"*, between Anas and Heteronetta. Anatinae, between Casar- ca and Tadorna. Plectropterus Plectropterinae, before Cairina and Sarkidior- nis. Plectropterinae, before Cairina and Sarkidior- nis. Plectropterinae, with An- seranas. Sarkidiornis Plectropterinae, between taxa currently merged into Cairina. Plectropterinae, between taxa currently merged into Cairina. Anatinae, between Neochen and Cairina. Hymenolaimus Merganettinae, between Salvadorina and Mer- ganetta. Merganettinae, between Salvadorina and Mer- ganetta. Nyrocinae% between Po- lysticta and Salvadori- na. Tachyeres Fuligulinae"', between Ay- thya and Bucephala. Fuligulinae"*, between Ay- thya and Bucephala. Nyrocinae, between Ay- thya and Bucephala. Merganetta Merganettinae, with Sal- vadorina and Hymeno- laimus. Merganettinae, with Sal- vadorina and Hymeno- laimus. Merganettinae (monotyp- ic), after “Merginae” (mergansers). Malacorhynchus Anatinae^, between Anas (part) and Marmaro- netta. Anatinae"*, between Anas (part) and Aix. Anatinae, between Anas (part) and Rhodonessa. Salvadorina Merganettinae, with Hy- menolaimus and Mer- ganetta. Merganettinae, with Hy- menolaimus and Mer- ganetta. Nyrocinae, with Polystic- ta and Hymenolaimus. ^ Generic taxa used by author(s) are replaced by currently accepted synonyms. Johnsgard (1979) listed no tribes within subfamilies. Salvadorina merged with Anas by author. ^ Skeletal specimens of Salvadorina not examined by Woolfenden (1961). naevosa) have included placements as: an aberrant dabbling duck (Delacour, 1938; Delacour and Mayr, 1945) or shelduck (Boetticher, 1942, 1952); the sole member of a tribe or higher taxon closely related to the true geese (Verheyen, 1953, 1955; Frith, 1964<3, 1964^?, 1965, 1967; Johnsgard, 1978); a very primitive sister-group of the majority of modern ducks, geese, and swans (Frith, 1967; Olson and Fed- uccia, 1980; Madsen et aL, 1988; Sibley and Ahlquist, 1990; Sibley and Monroe, 1990); or as both very primitive and a close relative of the stiff-tailed ducks (Fullagar et ah, 1990; Marchant and Higgins, 1990). Also, assignments of single species to monogeneric higher taxa has led to the phylogenetically misleading conclusion that each has, like Stictonetta, “no near living relatives” (e.g., Johns- gard, 1978:xx) or “is not closely related to any other tribe” (e.g., Edkins and Hansen, 1972:105). However, under an assumption of the monophyly of the biota of the earth, any species is the sister-group (i.e., closest relative) of some other group of organisms. At present, there is a consensus among most systematists that Stictonetta should be excluded from the true ducks (Anatinae), although other details of its relationships within the Anseriformes remain in dispute (Wolters, 1976; Johnsgard, 1979; Livezey, 1986^?; Madsen et ah, 1988; Sibley and Monroe, 1990). The Spur-winged Goose (Plectropterus gambensis) and Comb Duck (Sarki- diornis melanotus), traditionally considered to be members of an assemblage termed the “perching ducks,” or formally the Subfamily Cairininae or Tribe Cair- inini (e.g., Delacour, 1938, 1959, 1964; Delacour and Mayr, 1945; Boetticher, 1996 LiVEZEY T ADORNINE-An ATINE DIVERGENCE 29 Table 1. — Extended. Delacour and Mayr (1945) Boetticher (1952) Delacour (1954—1959) Anatinae, Anatini, as “aber- rant.” Anatinae, Tadornini, in mono- typic “section.” Anatinae, Anatini, after Hymen- olaimus, and Merganetta. Anatinae, Cairinini, after Sarki- diornis and Cairina. Anatinae, Cairinini, before Cairina and Sarkidiornis. Anatinae, Cairinini, after Sarki- diornis and Cairina. Anatinae, Cairinini, between Nettapus and Cairina. Anatinae, Cairinini, between Cairina and Nettapus. Anatinae, Cairinini, between Nettapus and Cairina. Anatinae, Anatini, as “aber- rant.” Anatinae, anatini, after Mala- corhynchus. Anatinae, Anatini, between Ma- lacorhynchus and Merganetta Anatinae, Tadornini, as “aber- rant.” Anatinae, Merganettini (mono- typic), after Oxyurini. Anatinae, Tachyerini, monoge- neric. Anatinae, Oxyurini, after Heter- onetta. Anatinae, Tadornini, after Lo- phonetta. Anatinae, Anatini, between Hy- menolaimus and Stictonetta. Anatinae, Anatini, as “aber- rant,” Anatinae, Anatini, merged with Anas. Anatinae, Anatini, before Hy- menolaimus. Anatinae, Anatini, between taxa currently merged into Anas. Anatinae, Anatini, between Rhodonessa and Hymenolai- mus. Anatinae, Anatini, merged with Anas. 1952; Johnsgard, 1965<2, 1978), were reassigned to a monotypic subfamily (Plec- tropterinae) and the shelducks (Tadorninae), respectively, by Livezey (1986a). Similarly, steamer-ducks (Tachyeres spp.; Livezey and Humphrey, 1992) and the Torrent Duck (Merganetta armata), for which persistent doubts concerning phy- logenetic relationships were finessed by some through allocations to monotypic tribes (Delacour and Mayr, 1945; Verheyen, 1953; Johnsgard, 1965a, 1966, 1978), also were included in the poorly resolved Tadorninae by Livezey (1986a). The Blue Duck (Hymenolaimus malacorhynchos) and the Pink-eared Duck (Malacorhynchus membranaceous), traditionally considered to be aberrant mem- bers of the dabbling ducks (Delacour, 1938; Delacour and Mayr, 1945; Boetticher, 1952; Delacour, 1956; Johnsgard, 1965a, 1965Z?, 1978, 1979; Kear and Steel, 1971; Sibley and Monroe, 1990), were included as well among the tadornines by Livezey (1986a). However, Olson and Feduccia (1980:22) suggested that Mala- corhynchus, together with Stictonetta and perhaps Merganetta, were '‘primitive relicts of an early anatid radiation and may be only distantly related to the re- mainder of the Anatidae.” Salvadori’s Duck (Salvadorina waigiuensis), for which skeletal and tracheal specimens were not available in the analysis performed by Livezey (1986a), also has been classified in recent decades as an unusual member of the dabbling ducks, and often merged with the genus Anas (Mayr, 1931; De- lacour and Mayr, 1945; Delacour, 1956; Johnsgard, 1965a, 1978, 1979). Subse- quent behavioral comparisons (Kear, 1975) and limited osteological data (MIL kovsky, 1989), however, indicate that the species may be more closely related to other waterfowl variably specialized for lotic habitats {Merganetta and Hymeno- laimus) or to the unique Malacorhynchus. Based in large part on these data, Livezey (1991) tentatively included the genus among the shelducks and allies 30 Annals of Carnegie Museum VOL. 65 Table 1. — Continued. Genus Woolfenden (1961) Johnsgard (1965“) Wolters (1976) Stictonetta Anserinae, Dendrocygnini (not examined). Anserinae, Stictonettini (monotypic), after An- serini. Stictonettinae (monotyp- ic), between Dendro- cygninae and Anseri- nae. Plectropterinae (monotyp- ic), between Tadorni- nae and Anatinae. Plectropterus Anatinae, Tadornini, be- fore Chloephaga and Cyanochen. Anatinae, Cairinini, be- fore Cairina and Sarki- diornis. Sarkidiornis Anatinae, Anatini, before Pteronetta and Cairina. Anatinae, Cairinini, be- tween Cairina and Pteronetta. Anatinae, before Cairina and Pteronetta. Hymenolaimus Anatinae, Anatini, be- tween Malacorhynchus and Tachyeres. Anatinae, Anatini, before Merganetta and Anas. Anatinae, between Mala- corhynchus and Tachy- eres. Tachyeres Anatinae, Anatini, after Malacorhynchus and Hymenolaimus. Anatinae, Tachyerini, be- tween Tadornini and Cairinini. Anatinae, between Hy- menolaimus and Mar- maronetta. Merganetta Anatinae, Merganettini, monotypic. Anatinae, Anatini, be- tween Hymenolaimus and Anas. Anatinae, between Lo- phonetta and Salvador- ina. Malacorhynchus Anatinae, Anatini, be- tween Salvadorina and Hymenolaimus. Anatinae, Anatini, be- tween Anas and Mar- maronetta. Anatinae, between Salva- dorina and Hymenolai- mus. Salvadorina Anatinae‘*, Anatini, be- tween Anas and Mala- corhynchus. Anatinae, Anatini, as monotypic subgenus of Anas. Anatinae, between Mer- ganetta and Malaco- rhynchus. (Tadorninae). Thirteen representative classifications of these eight genera spanning the last century are summarized in Table 1. Woolfenden (1961) presented the first study of the comparative osteology of waterfowl from the perspective of (phenetic) assessments of relationships. Livezey (19866) presented a species-level phylogenetic analysis of Tachyeres based on morphological comparisons, one which compared favorably with an assessment of genetic distances in the group (Corbin et ah, 1988) and biochemical patterns in secretions from the uropygial glands (Livezey et ah, 1986). Other contributions to the knowledge of the anatomy of these genera include, by genus: Stictonetta (Campbell, 1899; Frith, 19646; Olson and Feduccia, 1980), Plectropterus (Rand, 1954; Moller, 1969), Sarkidiornis (Garrod, 1875), Hymenolaimus (Kear and Bur- ton, 1971), Tachyeres (Livezey and Humphrey, 1986, 1992), Merganetta (Nie- thammer, 1952; Weller, 1968; Raikow, 1970), Malacorhynchus (Frith, 1955, 1967), and Salvadorina (Rothschild, 1895; Mayr, 1931; Kear, 1975; Mlikovsky, 1989). Tyler (1964) included several of these genera in a comparison of the mi- croanatomy of anseriform egg shells. Fossils assignable to any of these eight genera are limited to Pleistocene material of Sarkidiornis and subfossil material of Malacorhynchus (Brodkorb, 1964; Howard, 1964a; Olson, 1977, 1985; Worthy, 1995). 1996 Livezey — Tadornine-Anatine Divergence 31 Table 1. — Extended. Livezey (1986") Sibley and Monroe (1990) Johnsgard (1978) Johnsgard (1979)*’ Anserinae, Stictonettini (monotypic), after Cereopsini (monotyp- ic). Anatinae, Cairinini, be- fore Cairina and Sar- kidiornis. Anatinae, Cairinini, be- tween Cairina and Pteronetta. Anatinae, Anatini, be- fore Salvadorina^. Anatinae, Tachyerini, between Tadornini and Cairinini. Anatinae, Merganettini, between Cairinini and Anatini. Anatinae, Anatini, be- tween Anas and Mar- maronetta. Anatinae, Anatini, after Hymenolaimus as first member of Anas. Anserinae, between Cereopsis and Tador- nieae. Anatinae, before Cairi- na and Pteronetta. Anatinae, between Pteronetta and Net- tapus. Anatinae, between Merganetta and Sal- vadorina'^. Tadorninae, after Ta- dorna as last includ- ed genus. Anatinae, between Amazonetta and Hy- menolaimus. Anatinae, between Anas and Marmaro- netta. Anatinae, after Hymen- olaimus as first member of Anas. Stictonettinae (mono- typic), between An- serinae and Plectrop- terinae. Plectropterinae (mono- typic), between Stic- tonettinae and Tador- ninae. Tadorninae, Sarkidior- nithini (taxon amended; monotyp- ic). Tadorninae, Tadornini, in subtribe with Merganetta and Tachyeres. Tadorninae, Tadornini, in subtribe with Hy- menolaimus and Merganetta. Tadorninae, Tadornini, in subtribe with Hy- menolaimus and Tachyeres. Tadorninae, Tadornini, as subtribe between Tadorna and sheld- geese. Anatinae‘*, Anatini, pro visionally included in Anas. Stictonettinae (mono- typic), between Ox- yurinae and Cygni- nae. Anatinae, Anserini, be- tween tachyeres and Cairina. Anatinae, Anserini, be- tween Pteronetta and Nettapus. Anatinae, Anatini, be- tween Merganetta and Salvadorina Anatinae, Anserini, be- tween Tadorna and Plectropterus. Anatinae, Anatini, be- tween Amazonetta and Hymenolaimus. Anatinae, Anatini, be- tween Anas and Marmaronetta. Anatinae, Anatini, be- tween Hymenolaimus and Anas. Phylogenetic analyses using molecular characters are considered by some to hold potential for especially reliable inferences of phylogeny, but an assumption of inherent superiority lacks empirical support (Hillis, 1987; Sanderson and Don- oghue, 1989; Moritz and Hillis, 1990). Moreover, comparisons of studies based on molecular and morphological data, even where fundamentally phenetic meth- odologies were applied, share broad areas of agreement (e.g., Bledsoe and Rai- kow, 1990; Omland, 1994; see also Swofford, 1991). Unfortunately, most of the larger studies of Anseriformes employing molecular techniques included none of the eight genera considered here (Kessler and Avise, 1984; Patton and Avise, 1985; Scherer and Sontag, 1986). Numachi et al. (1983) included but a single member of this group of genera, Sarkidiornis, precluding any tests of relationships among members. Bottjer (1983) included Plectropterus and Sarkidiornis in his immunological comparisons; one especially distinctive finding was the low sim- ilarity between Plectropterus and the “perching ducks” Cairina, Chenonetta, and Aix. The electrophoretic comparisons of egg-white proteins by Sibley and Ahl- quist (1972:fig. 10-12) included Plectropterus, Sarkidiornis, Hymenolaimus, and Malacorhynchus, but not even a phenetic analysis of the intriguing patterns was presented. Other molecular analyses including two or more of these taxa also were phenetic in methodology, and therefore any inferred associations represent 32 Annals of Carnegie Museum VOL. 65 assessments of overall similarity rather than direct indications of phylogenetic relationship (Jacob and Glasser, 1975; Brush, 1976; Jacob, 1982; Jacob and Hoer- schelmann, 1993). Of the latter, the comparisons of feather proteins by Brush (1976) included six of the eight genera of interest here, but the resultant banding patterns proved difficult to interpret phylogenetically and rendered tribal assign- ments problematic for Stictonetta, Hymenolaimus, Merganetta, and Malacorhyn- chus. The analyses by Sibley and Ahlquist (1990) using DNA hybridization, and the related classification (Sibley et ah, 1988; Sibley and Monroe, 1990), also lacked any samples from these eight genera. The placement of Stictonetta by Madsen et al. (1988) in an analysis of 13 species of Anseriforrnes using DNA hybridization must be viewed with caution in light of criticisms of this general methodology (Cracraft, 1987; Houde, 1987; Sarich et ah, 1989; Springer and Krajewski, 1989), poor representation of taxa, and unfavorable metric properties evident in the com- parisons. The primarily osteological study by Livezey (1986<3) remains the only work in which even a majority of these problematic “tadornine” genera were analyzed. However, support for inferences in the topological vicinity of the divergence of the “shelducks” (Tadorninae) and the “surface-feeding ducks” (Anatinae) in the preliminary analysis by Livezey (1986<2), should be considered tentative. This assessment stems from several concerns: (1) the polychotomous topology depicted for the genera included in the tadornines (Livezey, 1986<3:fig. 1, 3); (2) the limited numbers of characters supporting groupings within this segment of the tree, e.g., those uniting Cyanochen with Hymenolaimus, Merganetta, and Tachyeres; (3) the impossibility of an assessment of support (e.g., bootstrapping of characters or consideration of suboptimal topologies) or the treatment of polymorphic character states at the time of the analysis; (4) three characters used by Livezey (1986a) to support clades in the tadornines — characters 15, 33, and 61^ — were judged unre- liable for these taxa based upon further study (Livezey, unpublished data); (5) several of the characters of the pelvic limb (including characters 56 and 69) show significant homoplasy evidently associated with diving habit (Livezey, 1986a; Faith, 1989), therefore groups united in part by these characters (e.g., Hymeno- laimus, Tachyeres, and Merganetta) merit confirmation; and (6) the profound limitations of software and computing platform increased the likelihood of sub- optimal solution sets (e.g., retention of more than 50 trees in memory for this data set typically caused a “crash” of the analysis). Livezey (1986a) indicated several areas of poor resolution and marginal support in the resultant phylogenetic hypothesis, and subsequent study discovered alter- native solutions to at least the basal segment of the tree (Livezey, 1989). In addition, reanalysis of the original data matrix compiled by Livezey (1986a) using improved versions of analytical software discovered multiple, slightly shorter al- ternative trees including several distinct topologies (J. Harshman, personal com- munication), the majority of which were the result of poorly resolved relationships in the vicinity of the divergence of “tadornine” and “anatine” genera. With respect to the eight genera of concern here, these alternative trees shared the following elements: (1) paraphyly of Stictonetta and Plectropterus to “tadornine” and “anatine” genera (as inferred by Livezey, 1986a); and (2) placement of Ma- lacorhynchus. Blue-winged Sheldgoose {Cyanochen cyanopterus), true shelducks (Tadorna) and other sheldgeese (Alopochen, Neochen, and Chloephaga), Sarki- 1996 Livezey — Tadornine-Anatine Divergence 33 diornis, and the clade comprising Hymenolaimus, Merganetta, and Tachyeres as paraphyletic to the genera included in the Anatinae by Livezey (1986^). Contradictory assessments of relationships in this group of genera “transition- al” between the anseriform subfamilies Tadorninae and Anatinae, as well as short- comings in analyses to date, justify a renewed effort to recover phylogenetic structure in this important group of waterfowl. Because of the phylogenetically central position of these genera, i.e., their topological proximity to the divergences of the tribes of the Anatinae (Livezey, 1986r?, 1991), a study of this kind provides an opportunity to assess available morphological evidence concerning the rela- tionships among the anatine tribes as well. This paper, one of a series of finer- scale analyses of the major subparts of the phytogeny of waterfowl (Livezey, 1991, 1995a, 1995 A 1995c, in press a, in press A in press c), presents a phylo- genetic analysis of the eight aforementioned genera and representatives of related tribes, using an enlarged set of morphological characters and improved analytical algorithms. This analysis is followed by ancillary considerations of morphological evolution, phylogenetic patterns of behavior and ecology, implications for histor- ical biogeography, and promising areas for future study for this most challenging aspect of the phytogeny of Anseriformes. Materials and Methods Taxonomy Composition of the Ingroup. — Determination of the broad limits of the com- paratively derived members of the Anseriformes, based on the cladistic analyses by Livezey (1986a, 1995a, in press c), conservatively included representatives of all groups exclusive of the screamers (Anhimidae), Magpie Goose (Anseranatidae: Anseranas semipalmata), whistling ducks and allies (Dendrocygninae), and true geese and swans (Anserinae). Comparisons included representatives of all taxa included in the eight genera of primary interest — Stictonetta, Plectropterus, Sar~ kidiornis, Hymenolaimus, Tachyeres, Merganetta, Malacorhynchus, and Salva- dorina. Representatives of other “tadornine” genera (Cyanochen, Alopochen, Neochen, Chloephaga, and Tadorna), also were included, which were inferred to constitute a monophyletic group in this series of phylogenetic analyses (Livezey, in press a). The Cape Barren Goose (Cereopsis novaehollandiae), considered by some to be an aberrant sheldgoose or an evolutionary “link” between the true geese and the sheldgeese (Delacour and Mayr, 1945; Delacour, 1954; Johnsgard, 1961a, 1978; Woolfenden, 1961; Frith, 1967; Rear and Murton, 1973), is excluded from this study on the basis of its inclusion among the true geese and swans (Livezey, 1986a, 1989, in press c). Also included are “basal” representatives (see below for methods of charac- terization) of the major clades of Anatinae: three subtribes of dabbling ducks or Anatini (Cairineae, Nettapodeae, and Anateae; Livezey, 1991), as well as the pochards (Aythyini; Livezey, in press b), seaducks (Mergini; Livezey, 1995^), and stiff-tailed ducks (Oxyurini; Livezey, 1995c). Inclusion of the separate sub- tribes of Anatini defined by Livezey (1991) in the analysis was necessitated by variation among the subgroups in characters determined to be informative as well among “tadornine” genera. Under the taxonomy proposed by Livezey (1991), the three subtribes of Anatini comprise the following genera: Cairineae {Cairina, Pteronetta, and Aix), Nettapodeae (Chenonetta and Nettapus), and Anateae (Ama- zonetta, Callonetta, Lophonetta, Speculanas, Mareca, and Anas). 34 Annals of Carnegie Museum VOL. 65 Composition of Included Genera. — -Five of the eight genera of primary interest in this analysis are considered to comprise single species by most taxonomists: Stictonetta, Plectropterus, Sarkidiornis, Hymenolaimus, and Merganetta (e.g., De- lacour and Mayr, 1945; Delacour, 1954, 1956, 1959; Johnsgard, 1961a, 1965a, \965b, 1978, 1979; Wolters, 1976). Segregation of Salvadori’s Duck in the genus Salvadorina (Woolfenden, 1961; Kear, 1975; Wolters, 1976; Mlikovsky, 1989; Sibley and Monroe, 1990; Livezey, 1991), instead of inclusion of the species within Anas (e.g., Delacour and Mayr, 1945; Boetticher, 1952; Delacour, 1956; Johnsgard, 1965a, 1978), delimits another monotypic genus in this group. Two of these purportedly monotypic genera^ — Sarkidiornis and Merganetta — include two or more qualitatively distinct taxa that might be accorded species rank. For example, in light of plumage differences and geographical isolation (Madge and Burn, 1988), partitioning of Sarkidiornis into a nominate African species S. melanotus and a Neotropical species S. sylvicola is consistent with both the emphasis of the “minimal diagnosable unit” under the phylogenetic species concept, and the requirements of distinct historical “identities” and independent evolutionary “tendencies” and “fate” under the evolutionary species concept (Wiley, 1978; Cracraft, 1983, 1988; McKitrick and Zink, 1988; Frost and Hillis, 1990). In practice, application of the phylogenetic species concept can approxi- mate the lineages conceptualized under the evolutionary species concept (Frost and Kluge, 1994). Numerous synapomorphies demonstrate the monophyly of Sar~ kidiornis (see beyond), however, and the included species were identical for the characters coded here; therefore the species-level taxonomy of the genus is not germane to this analysis. Similarly, substantial support for monophyly of Mer- ganetta (see beyond) and identical states of analyzed characters render inconse- quential any species-level interpretation one might impose on this poorly under- stood complex of three to six recognized forms for purposes of this analysis (Conover, 1943; Delacour, 1956; Johnsgard, 1978; Madge and Bum, 1988). The steamer-ducks {Tachyeres) are treated as a single generic taxon for this analysis; the genus comprises four confusingly similar species (Livezey and Humphrey, 1992), is demonstrably monophyletic (Livezey, 1986a, 1986Z?, prese'^nt study), members of the genus were identical for included characters, and the phytogeny of included taxa has been examined elsewhere (Livezey, 1986^?; Corbin et al., 1988). The Spur-winged Goose {Plectropterus gambensis) currently is considered to comprise two poorly differentiated subspecies (northern, comparatively pale gam- bensis and southern, comparatively dark niger) having a broad area of intergra- dation; the two forms are considered as a single analytical taxon here (Delacour, 1959; Johnsgard, 1978; Brown et al., 1982; Madge and Burn, 1988). No diag- nosable populations have been detected within the other four critical genera — Stictonetta, Hymenolaimus, Malacorhynchus, and Salvadorina (Delacour, 1956, 1959; Johnsgard, 1978, 1979). Specimens and Related Data Specimens Examined. — Study skins of adult birds were compared for deter- mination of plumage pattern and general structure of the externum in the collec- tions of several major museums, most importantly the National Museum of Nat- ural History and the American Museum of Natural History. Colors of the irides, bill, tarsi, and feet of adults also were characterized using study skins, but colors 1996 Livezey — Tadornine-Anatine Divergence 35 of unfeathered integument are not always reliably coded using dried specimens; consequently, colors of soft parts were confirmed using published descriptions (see below), notes on specimen labels, photographs of living individuals, and direct observation of living birds in avicultural collections or in the wild. Plumage patterns of downy young, specimens of which are comparatively rare in museum collections, were ascertained by comparisons of study skins (including an impor- tant collection at the Wildfowl and Wetlands Trust) and fluid-preserved specimens. Prepared skeletal specimens of adults were used for osteological comparisons. Numbers of prepared skeletons available varied significantly among taxa. Large series of skeletons (100 or more) were available for several groups (Tachyeres, Anas), whereas fewer than five were available for several others {Hymenolaimus, Merganetta); at least five skeletons were studied for other taxa, with the exception of Salvadorina (for which no complete skeletal specimen exists). Tracheal char- acters were determined using ossified elements in skeletal specimens and complete tracheae removed from fluid-preserved specimens (some previously mounted and dried). Published Data. — Published descriptions and illustrations of adult and subadult plumages supplemented data collected from direct study (Conover, 1943; Dela- cour, 1954, 1956, 1959, 1964; Johnsgard, 1965a, 1978; Weller, 1968; Todd, 1979; Humphrey and Livezey, 1982; Madge and Burn, 1988; Marchant and Higgins, 1990; Hoyo et al., 1992). Preliminary study of variation in natal plumages was accomplished using the illustrations and descriptions of downy young provided by Delacour and Mayr (1945, 1946), Delacour (1954, 1956, 1959, 1964), Frith (1955, 1964^7), Pengelly and Kear (1970), Kear (1972, 1975), and Humphrey and Livezey (1985). Variation and alternative interpretations of skeletal characters, most of which were based on my earlier analyses (Livezey, 1986a, 1989, 1991), were made using the works by Shufeldt (1909), Verheyen (1953), Rand (1954), Woolfenden (1961), Humphrey and Clark (1964), Cracraft (1968), Weller (1968), Moller (1969), and Raikow (1970). The few skeletal characters determinable for Salvadorina were coded using the descriptions of a few elements by Mayr (1931) and Mlikovsky (1989). Additional information on the tracheal anatomy of water- fowl was taken from Yarrell (1827), Garrod (1875), Campbell (1899), Mayr (1931), Niethammer (1952), Johnsgard (1961^, 1971), and Humphrey and Clark (1964). Definition of Characters Theory and Objectives. — Characters used in this phylogenetic analysis are dis- tinct morphological features, hypothesized to be homologous (de Pinna, 1991; Lipscomb, 1992; Hall, 1994), that comprise two or more states within the ingroup or differ between the ingroup and outgroups. Of the states defined, one is hy- pothesized as the primitive or plesiomorphous state based on outgroup compari- sons (see below), and the other(s) as the derived or apomorphic state(s). Definition of characters and included states are essential to any cladistic analysis, but rec- ognition of homology and description of the implicit distinctions between states remain the most challenging and influential aspect of phylogenetic inference (Pogue and Mickevich, 1990). Given that shared derived characters (synapomorphies) constitute the only em- pirical support for the delimitation of hierarchically nested sets of closest relatives (clades), the primary objective of character analysis is the discovery of such 36 Annals of Carnegie Museum VOL. 65 changes in states (transformation series) germane to the group of interest. Auta- pomorphies, character changes that are unique to terminal taxa, cannot contribute to the resolution of such groupings of included taxa (Wiley, 1981), although such differences traditionally have been accorded considerable weight in previous at- tempts at classification (e.g., Delacour, 1954; Johnsgard, 1961<3, 1965n, 1978). However, autapomorphies of the key eight genera of interest were included in this analysis for several reasons: (1) inclusion of autapomorphic divergence in cladis- tically based phylograms can provide insights into relative evolutionary rates, (2) autapomorphic characters of polytypic genera also are synapomorphic for included members, and (3) autapomorphic divergence can be depicted in phylogenetic trees independently of the character changes supporting topological structure and as- sociated summary statistics. Characters Defined. — A total of 1 14 morphological characters were defined for this analysis: 39 characters of the skeleton, four of the trachea and syrinx, 1 1 of the natal plumage, and 60 of the definitive plumage and soft parts (Appendix 1). Most skeletal characters were adapted from those described by Livezey (1986(2), with lesser numbers from Livezey (1989, 1991); for this analysis, states of these characters were restricted to those represented by the ingroup. This more detailed consideration of tracheosyringeal anatomy obviated the ad hoc weighting of the single (composite) syringeal transformation series defined in the analysis by Liv- ezey (1986(2). Behavioral patterns, especially courtship displays, have received considerable study in waterfowl (Delacour and Mayr, 1945; Delacour, 1954, 1956, 1959; Johns- gard, 1960a, 1961(2, 1962, 1965a, 1978). Such ethological data have provided insights into phylogenetic relationships of waterfowl (e.g., Johnsgard, 1961a) and have been shown to be comparable in consistency to morphological and molecular characters (de Queiroz and Wimberger, 1993). Formal inclusion of such data in the phylogenetic analysis, however, was precluded by numerous missing data, problems of coding behavioral characters termed “rare” or “uncommon” in a discrete scheme, and questions of homology and polarity not addressable from published descriptions. Each character was coded for the 19 members of the ingroup and a hypothetical “ancestor” (see below), producing a data matrix of dimension 20 X 114 (Ap- pendix 2). Missing data were as follows: 28 entries for skeletal characters of Salvadorina were coded as “missing”; two natal characters for the “hypothetical ancestor” were coded as “missing,” indicating an unknown polarity; and one entry for character 44, three entries for character 46, single entries for characters 47 and 48, and two entries for character 77 were treated as “missing” because of problematic comparability of states. In total, there were 38 matrix entries coded as “missing” (1.7% of matrix). The impact of missing data on phylogenetic in- ference has been considered by Nixon and Davis (1991) and Platnick et al. (1991). Fourteen characters were “multistate” (i.e., included two or more derived states), of which two (characters 29 and 78) were ordered, i.e., a particular or- dination of derived states was considered more probable on structural or devel- opmental grounds (Wilkinson, 1992). Analytical implications of multistate char- acters and ordering attract continuing controversy and empirical study (Hauser and Presch, 1991; Mickevich and Lipscomb, 1991; Lipscomb, 1992; Barriel and Tassy, 1993; Slowinski, 1993). Consequently, an assessment of the analytical im- pact of these two orderings was made by comparison with analyses in which all characters were treated as unordered. 1996 Li VEZE Y T ADORNINE- An ATINE DIVERGENCE 37 Characterization of Generic Ve^ ors. — Coding of states for characters found to be uniform among members of genera was straightforward. Most character states for the “typical” tadornines — Cyanochen, Alopochen, Neochen, Chloephaga, and Tadorna—wQVQ based largely on a separate species-level analysis (Livezey, in press a). Several characters for the polyspecific genera in this group {Chloephaga, Tadorna) varied among species. Similarly, character states for the “composite” vectors for the seven major subgroups of Anatinae, where invariant within groups, were made on the basis of the present study or taken from Livezey (1991). Where determinable on the basis of distribution of states within the species-level analyses of these genera (Livezey, 1991, 1995b, 1995c, in press b), the “basal” state (therefore presumptive locally ancestral state) was used to characterize the genus in this analysis. Where variation and distribution of states within a genus pre- cluded such a determination, the state was coded as polymorphic (character 4 for Cairineae, character 8 for the genera of Tadornini, character 82 for Anateae, and character 91 for Nettapodeae). Determination of Basal Polarities Outgroup Comparisons. — Basal polarities of characters, i.e., the states of char- acters judged to be ancestral for the ingroup under analysis, were inferred by reference to outgroups (Watrous and Wheeler, 1981; Maddison et ah, 1984), as opposed to ontogenetic criteria (de Queiroz, 1985; Mabee, 1989; Wheeler, 1990; Williams et ah, 1990). Based on the analysis by Livezey (1986<3), the most in- formative outgroups for study were the taxa of Anseriformes paraphyletic to the clade comprising Stictonetta, Plectropterus, Tadorninae, and Anatinae. Characters were compared against representative members of each of the subfamilies and tribes, with primary exemplars given in parentheses: Anseranatinae (Anseranas semipalmata), Dendrocygninae (Dendrocygna autumnalis and Thalassornis leu- conotus), and Anserinae (Cereopsis novaehollandiae, Branta canadensis, Anser anser, Coscoroba coscoroba, and Cygnus olor). Character states that were found in successive outgroups were considered plesiomorphous and used to define the states presumed to be ancestral for the ingroup. Hypothetical Ancestor. — A hypothetical ancestor, the set of character states judged to be primitive based on outgroup comparisons, was used to root the phylogenetic tree(s). This method provided a single, simple root for the ingroup without explicitly including representative outgroup taxa in this study, thereby avoiding ancillary analytical digressions concerning relationships among out- groups, considered elsewhere (Livezey, 1986<2, 1989, 1995a, in press c; Livezey and Martin, 1988). Monophyly of the ingroup under study here was demonstrated by Livezey (19860), and is further substantiated in the present analysis if one or more unambiguous synapomorphies for the taxa exclusive of the hypothetical ancestor are discovered; a less stringent criterion is a basal branch with positive length. Derivation of Trees Global Parsimony and Search Algorithms. — Phylogenetic reconstruction was based on the cladistic principle of grouping taxa on the basis of shared derived characters (synapomorphies), and the set of trees that accomplished this with a minimum number of hypothesized character transformations in the entire tree was sought (i.e., in accordance with the criterion of global parsimony). This approach 38 Annals of Carnegie Museum VOL. 65 was first formulated by Hennig (1966) and has been reviewed by Wiley (1981) and Wiley et al. (1991). Exhaustive searches of all possible trees were prohibited by the size of the data matrix; use of the branch-and-bound algorithm, a method guaranteed to discover all most-parsimonious trees (Hendy and Penny, 1982), also was not practical for a problem of this moderately large size (Swofford, 1993). Consequently, topological searches were undertaken using global branch swap- ping (a heuristic technique), the retention of ten trees at each step, using options for both simple and random order of entry of taxa (ten replications), and the retention of multiple equally parsimonious trees throughout the search. Also, three alternative branch-swapping methods (nearest-neighbor interchanges, tree-bisec- tion-reconnection, and subtree-pruning-regrafting) were used to avoid local optima or “islands'’ of suboptimal topologies (Maddison, 1991; Page, 1993). Several summary statistics were used to quantify the evidence supporting phy- logenetic trees (Kluge and Farris, 1969; Farris, 1989); total tree length; consis- tency index, both including (Cl) and excluding uninformative characters (Cl*); homoplasy index, both including (HI) and excluding uninformative characters (HI*); retention index (RI); and rescaled consistency index (RC). All phylogenetic analyses were performed using the software PAUP version 3.1 (Swofford, 1993) and MacClade version 3.0 (Maddison and Maddison, 1992) on a Macintosh Quad- ra 800. Weighting of Characters. — ^In addition to the equal (unit) weighting of char- acters employed in the baseline analyses, several alternative weighting schemes were used. Use of differential weighting in phylogenetic analysis remains contro- versial (Neff, 1986; Wheeler, 1986; Chippindale and Weins, 1994), but was in- cluded here to explore the impact of preconceived and post facto inferences con- cerning relative reliability of different character types. Two a priori differential weighting schemes were used: (1) ""two-group'" scheme — skeletal characters, con- sidered here to be evolutionarily more conservative than more-superficial mor- phological features based on studies by Livezey (1986(2, 1991, 1995(2, 1995 A 1995c, in press a, in press b, in press c), were given a weight of two whereas other character groups were given a weight of one; and (2) ""four-group" scheme — in which skeletal, tracheal, natal integumentary, and definitive integu- mentary characters were given weights of four, three, two, and one, respectively. Weighting of skeletal characters over those of the integument in this analysis was prompted further by the marginal confidence held in the homologies and the comparatively greater difficulties of variation and sexual dimorphism in several plumage characters included here (Appendix 1). Finally, I employed an a poster- iori weighting method — successive weighting (Farris, 1969; Carpenter, 1988) — in which characters are iteratively re weighted (base weight of 2) on the basis of rescaled consistency indices of characters. Consensus Trees. — Where multiple, equally parsimonious trees were discov- ered using the same set of analytical options, two consensus methods were em- ployed to summarize graphically the topological regions given support by the set of shortest trees: strict consensus trees (showing only those nodes shared by all equally parsimonious trees; Sokal and Rohlf, 1981) and majority-rule consensus trees (Margush and McMorris, 1981). The utility of consensus trees does not enjoy unanimous recognition (e.g., Barrett et al., 1991), and it has been shown that such summary trees are seldom interpretable in the same ways as the actual trees from which they are compiled (Swofford, 1991; Wiley et al., 1991). Accordingly, con- 1996 Livezey — T adornine-Anatine Divergence 39 sensus methods were used simply to summarize areas of congruence among equal- ly parsimonious trees. Assessment of Support Bootstrapping of Characters. — Standard summary statistics (e.g., consistency index) are flawed measures of empirical support for phylogenetic trees, because most are conditional in part on the number of taxa analyzed (Brooks et aL, 1986; Archie, 1989fl, 19891?, 1990; Sanderson and Donoghue, 1989; Faith and Cranston, 1991; Meier et aL, 1991). Consequently, stability of nodes in minimal-length trees in this analysis was assessed using bootstrapping, although the limitations of this method in phylogenetic contexts remain controversial (Carpenter, 1992; Davis, 1993; Hillis and Bull, 1993; Penny et aL, 1994). Applications involved generating 100 bootstrapped replicates using the same heuristic methods employed in the primary analyses, and summarizing those branches robust to this procedure through majority-rule consensus trees. The nature of the characters analyzed vi- olates the assumption of statistical independence required for the bootstrapped percentages to be viewed as statistically rigorous levels of confidence (Felsenstein, 1985a; Sanderson, 1989), therefore the procedure was employed here as one of several quantitative indices to empirical support for branches (Bremer, 1994). Phylogenetic ''Signal’' and Suboptimal Trees. — One measure of the phyloge- netic “signal” of a data set is the skewness statistic for the distribution of resultant tree lengths (Hillis, 1991; Huelsenbeck, 1991; Hillis and Huelsenbeck, 1991; Kal- lersjo et aL, 1992). Exhaustive generation of tree lengths was not feasible for a data set of this size, therefore the skewness statistics (gj) for random samples of 1000 trees (excluding the hypothetical ancestor) under each of the three a priori weighting schemes were calculated as an index to “phylogenetic signal” (Archie and Felsenstein, 1993). Values of gj, however, provide poorly focused indices to phylogenetic informativeness, distributional characteristics of g, remain poorly known, and values can be subject to significant bias (Kallersjo et aL, 1992). An alternative measure of support proposed by Bremer (1988), later dubbed the “decay index” by Donoghue et al. (1992), is the relative stability of nodes with respect to increased homoplasy (sensu lato, including both convergence and reversals). Specifically, the “decay index” for a given node is the minimal number of additional steps in tree length at which topologies not supporting the node are discovered. Estimates of the “decay index” were made for selected clades found to be robust under bootstrapping and various weighting regimes using exclusion- ary constraints in heuristic searches in PAUR Topologically Constrained Analyses. — The impact of particular topological constraints on the lengths of the shortest trees permissible for this matrix (i.e., the increases in total length) were estimated by performing searches constrained to preserve the specific phylogenetic relationship in question (while allowing to- pological modifications in other parts of the tree to accommodate this constraint). These comparisons were especially useful in quantifying the sacrifices in parsi- mony required to accommodate previously proposed phylogenetic relationships or traditional classifications in terms of the present data set. Phylogenetic Classification A Linnean classification, which incorporated inferred phylogenetic information while minimizing disruption of existing taxonomy, was prepared using the meth- 40 Annals of Carnegie Museum VOL. 65 ods described by Wiley (1979, 1981) and Wiley et al. (1991). These methods include the use of the “sequencing convention” to avoid daunting numbers of taxonomic ranks, annotation of taxa in unresolved polytomies as sedis mutabilis, and identification of possibly paraphyletic or polyphyletic taxa through annotation as incertae sedis. Taxa applied to higher taxonomic categories follow standard requirements of availability and seniority (International Commission of Zoological Nomenclature, 1985), based on published synonymies and classifications (Phillips, 1922, 1923, 1925, 1926; Boetticher, 1942, 1952; Wolters, 1976). Comparative Analyses General Objectives. — Phylogenetic hypotheses provide an opportunity to ex- amine the history of evolutionary change in selected life-historical parameters (Eldredge and Cracraft, 1980; Felsenstein, 1985^; Brooks and McLennan, 1991; Harvey and Pagel, 1991). In this study, a number of ecomorphological attributes not amenable to discrete coding and determination of homology were compiled and mapped a posteriori onto phylogenetic trees (Maddison, 1989; Mickevich and Weller, 1990) using MacClade 3.01 (Maddison and Maddison, 1992). Similar phylogenetic mapping was employed in analyses of other anseriform groups (Liv- ezey, 1995<2, \995b, 1995c, in press a, in press b, in press c). Detailed comparisons of the effects of different optimizations on the traces of attributes — e.g., ACCTRAN (length-neutral favoring of reversals over parallel- isms) and DELTRAN (length-neutral favoring of parallelisms over reversals) — were not performed because no justification was evident for favoring one method over another. Moreover, two alternative character optimizations — Dollo optimi- zation, in which it is required that each derived state is uniquely derived (Farris, 1977; Maddison et al., 1984), and the related option of irreversibility of character change (Sanderson, 1993) — were considered inappropriate for this analysis. How- ever, regions of the trees in which ambiguity of traces occurred are shown as “equivocal” (Maddison and Maddison, 1992). Quantitative methods for analysis of characters with respect to a phytogeny (e.g., Coddington, 1988, 1994; Pagel and Harvey, 1988; Grafen, 1989, 1992; Pagel, 1994), including the preliminary methods for tracing continuous characters permitted by MacClade, were not used primarily because of the inclusion of a number of higher taxa (e.g., Cairineae, Anateae) in the analyses that contain species exhibiting a diversity of states for the parameters of interest as well as poor resolution in parts of the phytogeny (see beyond). Attributes mapped include body mass (unweighted mean of sexes), mean clutch size, mean egg mass, relative clutch mass (mean mass of clutch divided by mean female body mass), preferred nest site (terrestrial or cavity), perching habit, rel- ative frequency of diving, and period of activity (diurnal, crepuscular, or noctur- nal). Data for these mapped attributes were taken from unpublished records as- sociated with specimens and from the literature (Niethammer, 1953; Delacour, 1954, 1956, 1959; Scott, 1954; Hallstrom, 1956; Hobbs, 1957; Warham, 1959; Johnsgard, \969a, 1961<3, 1962, \965a, \965b, 1978; Schon wetter, 1961; Johnson, 1963; Weller, 1964a, \96Ab, 1964c, 1976; Frith, 1967; Rear, 1970, 1975; Moffett, 1970; Siegfried, 1979; Clark, 1980; Wilson and Wilson, 1980; Brown et al., 1982; Humphrey and Livezey, 1982, 1985; Livezey and Humphrey, 1984a, 1984Z?; Ea- die et al., 1988; Livezey, 1988; Madge and Burn, 1988; Rohwer, 1988; Marchant and Higgins, 1990; Triggs et al., 1991, 1992; Williams, 1991; McNeil et al., 1992; Dunning, 1993). 1996 Livezey — Tadornine-Anatine Divergence 41 Results Shortest Trees Characters Weighted Equally. — Forty most-parsimonious trees were found us- ing equal weighting of characters, each having length 215, consistency index (Cl) of 0.670 (0.543 excluding uninformative characters. Cl*), homoplasy index (HI) of 0.367 (0.479 excluding uniformative characters, HI*), retention index (RI) of 0.641, and rescaled consistency index (RC) of 0.430. The skewness statistic (gj) for the unweighted matrix, based on 1000 randomly generated trees, was —0.612; this figure is substantially more negative than that expected for random data. A strict consensus tree of the 40 topologies revealed only broad inferences shared by the “simplest” interpretations of the equally weighted characters (Fig. 1): (1) monophyly of the taxa analyzed exclusive of Stictonetta; (2) a sister re- lationship between Plectropterus and Sarkidiornis; (3) monophyly of the typical shelducks and sheldgeese (Tadorna, Cyanochen, Alopochen, Neochen, and Chloe- phaga), with intergeneric relationships as inferred by Livezey (1996«); (4) mono- phyly of Hymenolaimus, Tachyeres, and Merganetta, wherein the last two genera are sister-groups; and (5) a sister relationship between Salvadorina and Malacor- hynchus. The solution set was unchanged if all characters were analyzed as unor- dered. A majority of the shortest trees (Fig. 2) indicated the following additional phylogenetic structure: (1) a sister relationship between Plectropterus + Sarki- diornis and the clade comprising the typical shelducks and sheldgeese (39 of 40 trees); (2) a sister relationship between the preceding two clades and that com- prising Hymenolaimus, Tachyeres, and Merganetta (38 of 40 trees); and (3) mono- phyly of the six taxa of “typical” anatines (Cairineae, Nettapodeae, Anateae, Aythyini, Mergini, and Oxyurini), within which the three subtribes of Anatini defined by Livezey (1991) are monophyletic and the two subtribes of smaller “perching ducks” are sister-groups. Comparative Consistencies. — Mean consistency indices for informative char- acters under equal weighting were: skeletal, 0.63 {n = 27; range, 0.33-1.00); tracheal, 0.75 {n = 2; range, 0.50-1.00); natal, 0.62 {n = 10; range, 0.20-1.00); and definitive integument, 0.64 {n = 27; range, 0.25-1.00). Consistency indices varied among the 40 trees, as a result of differences of single steps, in eight skeletal characters (characters 5, 10, 11, 21, 24, 25, 30, 35), one natal character (46), and ten characters of the definitive integument (55, 56, 91, 93, 95-100). ''Two-Group’’ Weighting.— K heuristic search employing the “two-group” a priori weighting scheme, within which skeletal characters were given a weight of two and all others retained weights of one, discovered four shortest trees shar- ing the following summary statistics: length = 290, Cl ^ 0.676 (Cl* = 0.554), HI = 0.372 (HI* = 0.474), RI = 0.674, and RC = 0.455. The skewness statistic (gi) for the “two-group” matrix, based on 1000 randomly generated trees, was —0.584, The four topologies differed only in the relationships inferred among the six taxa of “typical” anatines. A strict consensus tree of these four shortest to- pologies (Fig. 3) confirmed the phylogenetic structure shared by a majority of the trees found under equal weighting (Fig. 2), with the following additional reso- lutions: (1) the clade comprising Salvadorina and Malacorhynchus is inferred to be the sister-group of the six taxa of “typical” Anatinae; and (2) within the latter, the Mergini and Oxyurini are inferred to be sister tribes. A majority-rule consen- sus tree of the topologies under “two-group” weighting revealed that three of the 42 Annals of Carnegie Museum VOL. 65 Stictonetta Plectropterus Sarkidiornis Cyanochen Alopochen Neochen Chloephaga Tadorna Hymenolaimus Tachyeres Merganetta Salvadorina Malacorhynchus Cairineae Nettapodeae Anateae Aythyini Mergin'! Oxyurini Ancestor Fig. 1. — Strict consensus tree of 40 equally parsimonious trees for 19 taxa of Anatidae under equal weighting of characters. 1996 Li VEZE Y — T adornine-An ATiNE Divergence 43 Stictonetta Plectropterus Sarkidiornis Cyanochen Alopochen Neochen Chloephaga Tadorna Hymenolaimus Tachyeres Merganetta Salvadorina Malacorhynchus Cairineae Nettapodeae Anateae Aythyini Mergin'! Oxyurini Ancestor Fig. 2. — Majority-mle consensus tree of 40 equally parsimonious trees for 19 taxa of Anatidae under equal weighting of characters. Percentages of these 40 trees in which each branch was conserved are shown. 44 Annals of Carnegie Museum VOL. 65 Stictonetta Plectropterus Sarkidiornis Cyanochen Alopochen Neochen Chloephaga Tadorna Hymenolaimus Tachyeres Merganetta Salvadorina Malacorhynchus Cairineae Nettapodeae Anateae Aythyini Mergin'! Oxyurini Ancestor Fig. 3. — Strict consensus tree of four equally parsimonious trees for 19 taxa of Anatidae, wherein skeletal characters were given a weight of two and all other characters a weight of one (“two-group” weighting scheme). 1996 Livezey — Tadornine-Anatine Divergence 45 four shortest trees confirmed the sister relationship between the Anateae and the two, generically restricted subtribes of “perching ducks,” Cairineae and Netta- podeae (Fig. 4). This solution set is topologically identical to that based on an analysis in which all characters were treated as unordered. Numbers of unambiguous character changes supporting each branch in the first of the four most-parsimonious trees varied from the single synapomorphy uniting Salvadorina and Malacorhynchus with the “typical” anatines, to the seven uniting Hymenolaimus, Tachyeres, and Merganetta (Fig. 5); identities of supporting char- acters are shown in Figure 6. As in all the weighting schemes for these taxa, the most prominent component of differentiation among taxa was autapomorphy, which does not contribute to resolution of phylogenetic relationships. The topo- logical variants within the Anatinae differed primarily in the transformations in- ferred for characters 46, 56, and 99 (Appendices 1, 2). “FoMr-GroM/?” Weighting. — Under the “four-group” weighting scheme, wherein weights of 4, 3, 2, and 1 were applied to characters of the skeleton, trachea, natal integument, and definitive integument, respectively, a single most- parsimonious tree was found. The skewness statistic (gi) for the “four-group” matrix, based on 1000 randomly generated trees, was -0.588. The topology of this tree was identical to the first of the four shortest trees recovered under the “two-group” weighting scheme (Fig. 5, 6), and had the following summary sta- tistics: total length = 476, Cl = 0.676 (Cl* = 0.559), HI = 0.378 (HI* = 0.471), RI = 0.691, and RC = 0.468. Analysis using the same weighting scheme but in which all characters were treated as unordered resulted in a single tree that dif- fered in only two topological features: (1) Plectropterus was placed as the sister- group to all included taxa but Stictonetta (as in Livezey, 1986<3); and (2) the Cairineae was placed as the sister-group to other Anatinae, including Salvadorina + Malacorhynchus. Successive Weighting. — Using the equally weighted data matrix as a starting point (admitting 40 shortest trees), two successive reweightings of the characters (base weight 2, using RC) recovered, respectively, eight and two trees of equal length; these two trees were resistant to additional reweighting cycles. Both trees had the following summary statistics: length = 230; Cl = 0.857 (Cl* — 0.703); HI == 0.178 (HI* = 0.315); RI = 0.828; and RC = 0.709. The skewness statistic (^i) for 1000 randomly generated trees based on the successively weighted matrix was “0.772. A strict consensus tree of these two solutions differed from those for the short- est trees derived using a priori weighting schemes in two important ways (Fig. 7): (1) placement of the Mergini as the sister-group of all taxa exclusive of Stic- tonetta and the clade comprising Salvadorina, Malacorhynchus, Nettapodeae, An- ateae, Aythyini, and Oxyurini (a placement made on first iteration); and (2) place- ment of the Cairineae as the sister-group of the clade comprising Plectropterus and Sarkidiornis (placement made on the second iteration). The interrelated effects of reweighting on the distributions of character changes in the trees are difficult to interpret, but examination of supporting characters indicates that comparatively few characters underlie these two divergent inferences. Topologically identical series of solutions were found using successive weighting of unordered characters. The position of the Mergini reflects, at least in part, shared changes in char- acters 95 (lobation of hallux), 98 (ground color of secondary remiges), and 99 (color of wing linings). The position of the Cairineae in the successive weighting analysis evidently stems from a number of characters that members of the subtribe 46 Annals of Carnegie Museum VOL. 65 Stictonetta Plectropterus Sarkidiornis Cyanochen Alopochen Neochen Chloephaga Tadorna Hymenolaimus Tachyeres Merganetta Salvadorina Malacorhynchus Cairineae Nettapodeae Anateae Aythyini Mergin'! Oxyurini Ancestor Fig. 4. — Majority-rule consensus tree of four equally parsimonious trees for 19 taxa of Anatidae, wherein skeletal characters were given a weight of two and all other characters a weight of one (“two- group” weighting scheme). Percentages of these four trees in which each branch was conserved are shown. 1996 Li VEZE Y — T adornine-An ATiNE Divergence 47 Fig. 5. — One of four equally parsimonious trees for 19 taxa of Anatidae under the “two-group” weighting scheme, showing numbers of unambiguous characters supporting each branch. shared with Plectropterus, Sarkidiornis, or (less commonly) both genera: char- acters 5 (processus supraorbitalis), 10 (spina externa sterni), 64 (unfeathered facial region), 93 (coloration of tertials), 96 (black dorsal color), and 99 (color of wing linings). Of the characters primarily involved in these two inferences, one posed difficulties in codings for several taxa (character 99) and only three achieved a Cl of 1.00 (characters 5, 10, and 96). In addition, most of the foregoing characters suffered declines in consistency with the inclusion of more genera of waterfowl or species-level codings (Livezey, 1991, 1995^, in press a) of magnitudes in excess of those typically associated with the inclusion of additional taxa (San- derson and Donoghue, 1989). Support for Branches Bootstrapping of Characters. — Under equal weighting of characters, a major- ity-rule consensus tree of 100 bootstrapped replicates conserved only the nine clades shared by all 40 shortest trees (Fig. 1); support for these branches ranged from 56% for the four genera of sheldgeese to 99% for the ingroup exclusive of Stictonetta (Fig. 8). Identical groups were preserved in a majority-rule consensus tree of 100 bootstrapped replicates under the ‘"two-group” weighting scheme, with only minor differences in percentages (Fig. 9). A majority-rule consensus tree of 100 bootstrapped replicates under the “four-group” weighting scheme retained the same groupings with the exception of that uniting Salvadorina with Malacorhynchus (Fig. 10). Bootstrapping after two iterations of successive 48 Annals of Carnegie Museum VOL. 65 1996 LiVEZEY T ADORNINE-An ATINE DIVERGENCE 49 weighting produced a majority-rule consensus tree topologically identical to those derived from bootstrapping under equal weighting or “two-group” weighting (Fig. 8, 9), with the exception of the placement of the Cairineae as sister-group of Plectropterus and Sarkidiornis, a grouping retained in only 59% of the repli- cates (Fig. 11). Near-shortest Trees. — ^Under equal weighting, numbers of equally parsimoni- ous trees increased precipitously with admission of suboptimal solutions; retention of trees having only two additional steps resulted in the discovery of over 3500 trees (Table 2). Subjecting the matrix to “two-group” weighting lowered the numbers of topologies of near-minimal lengths considerably, and “four-group” weighting further narrowed the solution sets at near-minimal lengths (Table 2). Tallies of near-minimal tree lengths under successive weighting resembled those for the “four-group” weighting scheme (Table 2). Bremer or ''Decay'' Indices. — Stability of several clades common to most weighting schemes or consensus techniques with respect to additional homoplasy (i.e., suboptimal tree lengths) was measured using Bremer or “decay” indices. Monophyly of the ingroup exclusive of Stictonetta was resistant to increased homoplasy under all weighting schemes, e.g., requiring an additional six steps under equal weighting to disrupt the clade (Table 3). The clade comprising Hy- menolaimus, Tachyeres, and Merganetta showed moderately high resistance to increased homoplasy, an integrity that was magnified under regimes that weighted skeletal characters highly (Table 3). Three other recurrent clades — typical shel- ducks and sheldgeese, Plectropterus + Sarkidiornis, and Malacorhynchus + Sal- vadorina — required only one to three additional steps before alternative arrange- ments were admissible under most weighting regimes (Table 3). Ecomorphological Evolution Morphological Attributes. — For purposes of a posteriori mapping, the first of the four shortest trees derived using “two-group” weighting of characters was used (Fig. 5, 6), a tree sharing most of the well-supported elements common to trees based on other analyses (Fig. 1-4, 7-11). In spite of the generalizations necessary for coding parameters for genera and higher taxa in the ingroup, broad evolutionary patterns in body mass, sexual size dimorphism, and sexual dichro- matism were evident. Body mass appears to have undergone a substantial decrease in the common ancestor of the ingroup, although the ancestral state of this attrib- ute is questionable; significant, subsequent increases in body mass characterize three clades: Sarkidiornis + Plectropterus, the sheldgeese, and Tachyeres (Fig. 12A). Uncertainty in the states of associated stems makes the homology of the large body masses of the first two clades indeterminate. Patterns of sexual size dimorphism within the ingroup shows a similar pattern to that evident in body mass, although increases in the former include Tadorna (indicating homology in the increase in a clade comprising seven genera), Mer- ganetta (interpretable as homologous to that in Tachyeres), and the Cairineae (Fig. Fig. 6. — Detailed phylogenetic tree, one of four equally parsimonious trees for 19 taxa of Anatidae under the “two-group” weighting scheme, showing the characters unambiguously supporting each branch. Character numbers correspond to those used in the character descriptions (Appendix 1) and data matrix (Appendix 2). 50 Annals of Carnegie Museum VOL. 65 Fig. 7. — Strict consensus tree of the two shortest phylogenetic trees for 19 taxa of Anatidae converged upon after three successive reweightings of characters based on rescaled consistency indices, using equal initial weights and a base for reweighting of two. 12B). The moderately high dimorphism of the Cairineae is more parsimoniously interpreted under the alternative position of this group, as sister-group to Sarki- diornis and Piectropterus, supported by successive weighting (Fig. 7, 11). Sexual dichromatism manifested evolutionary patterns completely distinct from those for sexual size dimorphism (Fig. 12C). Comparatively minor, quantitative dichromatism appears to have evolved independently in Sarkidiornis, Tachyeres, and basal Anateae; this intermediate state evidently gave rise to pronounced, qual- itative dichromatism in a number of more-derived members of the latter subtribe (Livezey, 1991). Distinct, qualitative sexual dichromatism appears to have evolved in four other lineages independently: Chloephaga, Tadorna, Merganetta, and the remaining taxa of “typical” Anatinae (Fig. 12C). Parameters of Reproduction. — Polygyny without establishment of significant pair-bonds, inferred to be derived in this group of Anseriformes, is shared by the clade comprising Piectropterus and Sarkidiornis and the Cairineae (Fig. 12D). Among Anseriformes, parental investment by males is inversely correlated with sexual size dimorphism (Sigurjonsdottir, 1981), an attribute that was especially high in these three taxa (Fig. 12B; Livezey and Humphrey, 1984a). The associ- ation between paternal investment and sexual dichromatism in waterfowl is less clear (Johnsgard, 1962; Scott and Clutton-Brock, 1989). Sexual dichromatism in Anseriformes has traditionally been interpreted in terms of isolating mechanisms among closely related species (e.g., Sibley, 1957; Johnsgard, 1960A 1963), a perspective of questionable utility among these diverse taxa showing little or no 1996 Li VEZEY — T adornine-An ATiNE Divergence 51 85 99 56 79 60 71 86 65 62 Stictonetta Plectropterus Sarkidiornis Cyanochen Alopochen Neochen Chloephaga Tadorna Hymenolaimus Tachyeres Merganetta Salvadorina Malacorhynchus Cairineae Nettapodeae Anateae Aythyini Mergin'! Oxyurini Ancestor Fig. 8. — Majority-rule consensus trees for 19 taxa of Anatidae of 100 bootstrapped replicates under equal weighting of characters. 52 Annals of Carnegie Museum VOL. 65 62 99 74 78 70 81 87 64 58 Stictonetta Plectropterus Sarkidiornis Cyanochen Alopochen Neochen Chloephaga Tadorna Hymenolaimus Tachyeres Merganetta Salvadorina Malacorhynchus Cairineae Nettapodeae Anateae Aythyini Mergini Oxyurini Ancestor Fig. 9. — Majority-rule consensus trees for 19 taxa of Anatidae of 100 bootstrapped replicates under “two-group” weighting. 1996 Livezey — Tadornine-Anatine Divergence 53 52 98 85 84 72 86 87 65 Stictonetta Plectropterus Sarkidiornis Cyanochen Alopochen Neochen Chloephaga Tadorna Hymenolaimus Tachyeres Merganetta Salvadorina Malacorhynchus Cairineae Nettapodeae Anateae Aythyini Mergini Oxyurini Ancestor Fig. 10. — Majority-rule consensus trees for 19 taxa of Anatidae of 100 bootstrapped replicates under “four- group” weighting. 54 Annals of Carnegie Museum VOL. 65 93 59 100 84 72 53 87 100 76 96 Stictonetta Plectropterus Sarkidiornis Cairineae Cyanochen Alopochen Neochen Chloephaga Tadorna Hymenolaimus Tachyeres Merganetta Salvadorina Malacorhynchus Nettapodeae Anateae Aythyini Mergini Oxyurini Ancestor Fig. 11. — Majority-rule consensus trees for 19 taxa of Anatidae of 100 bootstrapped replicates for matrix subjected to successive weighting (see text for details). 1996 Livezey — T adornine-Anatine Divergence 55 Table 2. — Numbers of trees of minimal and near-minimal lengths under three a priori and one suc- cessive weighting schemes for 19 generic, subtribal, and tribal taxa of Anatidae. Weighting scheme Minimal Numbers of trees of length (m) m m + 1 m + 2 m + 3 m + 4 Unit (equal) 215 40 488 3503 — — “Two-group” 290 4 34 148 657 — “Four-group” 476 1 9 28 114 425 Successive 230 2 8 36 188 673 tendency toward hybridization (Scherer and Hilsberg, 1982) and in light of con= ceptual difficulties with this emphasis (Livezey, 1991, 1995^). Preferred nest site appears to have undergone two independent shifts to cavities, in Tadorna and (within the context of this particular tree, homologously) in the Cairineae and Nettapodeae (Fig. 12E). Modal clutch size varied greatly in this group (Fig. 12F), and states of terminal taxa rendered hypotheses of states inde- terminate for most stems and characterization of the Anateae, Mergini, Oxyurini, and Aythyini problematic. Suggested evolutionary changes in this parameter are (Fig. 12F): (1) an increase in the ancestor of the ingroup; (2) several increases within the shelducks and sheldgeese; (3) decreases, possibly homologous, in Hy- menolaimus and Merganetta; and (4) increases, interpretable as homologous with- in the context of this particular tree, in the Cairineae and Nettapodeae. Mean egg masses (g) for the eight key genera of the ingroup were: Stictonetta (66), Plectropterus (131), Sarkidiornis (65), Hymenolaimus (73), Merganetta (62), Tachyeres (115--167), Malacorhynchus (31), and Salvadorina (57). Egg mass is generally correlated with body mass in the ingroup, making a mapping of this parameter largely redundant with that presented for body mass (Fig. 12A). Explicit mapping of relative clutch mass also proved intractable for these taxa because of variation in the three parameters composing this ratio (female body mass, egg mass, clutch size), some of which are poorly known for some genera. In addition, generalizing this ratio for the tribal and generic vectors also proved problematic; the range of values within several subtribal and tribal taxa encompassed a greater range than all eight of the genera of primary interest combined. Estimates for relative clutch masses (percentages of female body mass, largely based on Roh- wer, 1988) for the key ingroup genera were as follows: Stictonetta (55), Plec- Table 3. — Bremer or “decay” indices for selected clades within 19 generic, subtribal, and tribal taxa of Anatidae under three a priori and one successive weighting schemes. Percentage of minimal total tree length represented by each index is given in parentheses to facilitate comparisons across weight- ing schemes. Clade Weighting scheme Equal Two-group Four-group Successive Tadorninae + Anatinae® 6 (2.8) 7 (2.4) 8(1.7) 6 (2.6) Tadornini*’ 1 (0.5) 2 (0.7) 3 (0.6) 1 (0.4) Plectropterus + Sarkidiornis 3(1.4) 5(1.7) 5(1.1) 5 (2.2) Hymenolaimus + Tachyeres + Merganetta 3 (1.4) 4(1.4) 20 (4.2) 9 (3.9) Malacorhynchus + Salvadorina 2 (0.9) 4(1.4) 1 (0.2) 2 (0.9) ® Includes all taxa analyzed herein except hypothetical ancestor and Stictonetta. Includes Cyanochen, Alopochen, Neochen, Chloephaga, and Tadorna. 56 Annals of Carnegie Museum VOL. 65 ri # . □ 0 u Body Mass (g) ordered I I < 500 I i 500-1500 I ' S 1500-2500 ■1 > 2500 uncertain ^Sequivocal . 1.40 :#H uncertain equivocal Fig, 12. — Mapping of eight ecomorphological attributes on the first of four minimal-length trees recov- ered under “two-group” weighting of characters (Appendices 1, 2): A — body mass, B — sexual size dimorphism, C — sexual dichromatism, D — mating system, E — typical nest site, F — clutch size, G — paternal attendance of broods, and H — diving habit. The tree shown is topologically identical to the single shortest tree found under the “four-group” weighting scheme, in which a soft polytomy involving the taxa Aythyini, Oxyurini, Mergini, Cairineae, Nettapodeae, and Anateae is depicted as resolved. 1996 Livezey — Tadornine-Anatine Divergence 57 □ Dichromatism unordered absent I J rudimentary I^H pronounced H’M uncertain ^Sequivocal ■ ■ u □ □ j-J 1 □ □ □ □ ■ □ □ □ on Uirpij L — 1 D — Mating System unordered □ monogamy ■I polygyny ^Huncertain Fig. 12. — Continued. tropterus (41), Sarkidiornis (31), Hymenolaimus (49), Merganetta (62), Tachyeres (30-35), Malacorhynchus (63), and Salvadorina (37). In addition to the three taxa in which polygyny is typical {Plectropterus, Sar- kidiornis, and Cairineae), a derived loss of paternal attendence of broods is char- acteristic of the Aythyini, Mergini, and Oxyurini; variation within the Anateae in 58 Annals of Carnegie Museum VOL. 65 E— Nest Site unordered terrestrial HH cavity uncertain ..r □ n □ □ E3 □ 00m xj T fez: ';'r F- ■ Clutch Size ordered CZ] 3-5 LZ6-8 ■I 9-11 ^Suncertain ^Sequivocal Fig. 12. — Continued. this attribute necessitated coding the taxon as polymorphic (Fig. 12G). Within the phylogenetic hypothesis used as a basis for mapping, the simplest evolutionary pattern suggested is a loss of paternal attendance in the ancestor of the “typical” Anatinae, with subsequent, independent reversals in the Nettapodeae and some Anateae. 1996 Li VEZE Y — T adornine-An ATiNE Divergence 59 G— Paternal Attendance unordered I I typical atypical ^Suncertain Ecological Trends. — Most ecological attributes successfully mapped in other anseriform groups (Livezey, \995b, 1995c, in press a, in press b, in press c) remain too poorly known for the key genera under study here, and furthermore were variable within the higher taxa included in this analysis. These inadequately documented attributes include: diel pattern, in which most species are probably 60 Annals of Carnegie Museum VOL. 65 crepuscular, but some like marine Tachyeres are influenced by tides (Livezey, 1988; McNeil et al., 1992); diet, in which most species probably feed on both plants and invertebrates, rarely vertebrates; aquatic habitat, in which all species are primarily freshwater except most Tachyeres, some Tadorna, some Chloepha- ga, and most Mergini; nest parasitism, which is probably infrequent in most in- group taxa as in most Anseriformes (Rohwer and Freeman, 1989); formation of creches (Eadie et al., 1988); parental carrying of young (Johnsgard and Kear, 1968) through the air (reported in Sarkidiornis, Tadorna, Alopochen, Chloephaga, and some members of the Cairineae, Anateae, Mergini, and Oxyurini) or on water (reported in Salvadorina, and some members of Chloephaga, the Mergini and Oxyurini); and migratory habit, wherein most species show variable seasonal movements, with the notable exceptions being sedentary Hymenolaimus and Mer- ganetta, and flightless species of Tachyeres. Diving habit is characteristic of four taxa in this analysis, evidently representing either: ( 1 ) three or four convergent acquisitions, the interpretation most consistent with most described evolutionary scenarios (Delacour and Mayr, 1945; Johnsgard, 1978; Faith, 1989); or (2) a single derivation in the ancestor of the Tadorninae and Anatinae followed by three independent losses (Fig. 12H). Possible homo- plasy in several morphological characters related to diving habit or its loss (e.g., characters 38, 41), both having locally high consistencies but showing global homoplasy (Livezey, 1986<2; Faith, 1989), might compromise their utility for phy- logenetic inference (especially with respect to Hymenolaimus, Merganetta, and Tachyeres). Several of these diving-related characters are coded for this analysis as lacking for tribes in which a majority of the members possess the trait (Ay- thyini, Mergini, Oxyurini), because the absence of the character in basal members of each tribe indicates the character was acquired independently in more-derived members of these groups (Livezey, 1995 A 1995c, in press b). Discussion Phylogenetic Inferences Points of Concern. — With the exception of the support for the monophyly of the ingroup exclusive of Stictonetta (i.e., Tadorninae + Anatinae), most members of the ingroup are characterized more by unique divergence than nested, unifying characters. For example, Merganetta and Tachyeres are inferred to be sister-genera on the basis of two unambiguous synapomorphies, whereas the genera are them- selves supported by 15 and 13 autapomorphies, respectively (Fig. 6). In part because of concerns regarding possible homoplasy stemming from convergent life histories, I remain somewhat suspicious with respect to the clade comprising Hymenolaimus, Merganetta, and Tachyeres, in part because of the possible effects of “long-branch attraction” (see below). However, there are seven unambiguous characters that support this clade (Fig. 6), the grouping is comparatively robust with respect to weighting schemes (Table 3; Fig. 1-11), and there is a significant sacrifice of parsimony (Table 4) required to accommodate the positions of these genera proposed by Johnsgard (1965a, 1978, 1979). The position of the Cairineae remains a challenging problem; this group com- prises three genera and five species traditionally included among the “perching ducks” (e.g., Delacour and Mayr, 1945; Delacour, 1959; Johnsgard, 1961a, 1965a, 1978, 1979), but was inferred by Livezey (1991) to be the sister-group of a clade comprising the Nettapodeae and Anateae. In this study, most weighting schemes 1996 Li VEZEY — T adornine-An ATiNE Divergence 61 Table 4. — Minimal numbers of additional steps required to preserve previously hypothesized taxo- nomic grouping among 19 generic, subtribal, and tribal taxa of Anatidae, under three a priori and one successive weighting schemes. Percentage of minimal total tree length represented by each index is given in parentheses to facilitate comparisons across weighting schemes. Weighting scheme Taxonomic group Equal Two-group Four-group Successive Tribe CairininF 2 (0.9) 6(2.1) 12(2.5) 3(1.3) Subfamily Anatinae^’ Subfamily Anatinae^ exclusive 10(4.7) 18 (6.2) 31 (6.5) 18 (7.8) of Hymenolaimus Tree based on Johnsgard 2 (0.9) 6(2.1) 12 (2.5) 5 (2.2) (1961a, 1978)'^ Tree depicted by Livezey 36(16.7) 62 (21.4) 117 (24.6) 58 (25.2) (1986a:fig. 1)“ 16 (7.4) 19 (6.6) 29 (6.1) 43 (18.7) Stictonetta + Oxyurini® 7 (3.3) 16(5.5) 31 (6.5) 9 (3.9) ^ Of the taxa defined herein, includes Plectropterus, Sarkidiornis, Cairineae, and Nettapodeae; Ama- zonetta and Callonetta, included in this tribe by Johnsgard (1978), are here included as basal members of the Anateae (Livezey, 1991). Includes Plectropterus, Sarkidiornis, Hymenolaimus, Merganetta, Malacorhynchus, Salvadorina, Cairineae, Nettapodeae, Anateae, Aythyini, Mergini, and Oxyurini (Johnsgard, 1965<2, 1978, 1979). ‘^Conservative reconstruction of phylogenetic diagrams depicted by Johnsgard (1961a, 1978) as per- mitted using the present taxonomic units (Fig. 13). Constraint follows Livezey (1986a:fig. 1), except Anatinae simplified to polytomy involving Salva- dorina (missing taxon), Cairineae, Nettapodeae, Anateae, Aythyini, and Mergini + Oxyurini. ® Grouping followed by Marchant and Higgins (1990). either: (1) failed to establish the higher-order relationships of this taxon (Fig. 1, 8-10); or (2) placed the Cairineae as the sister-group to the Nettapodeae {contra Livezey, 1991) with or without indicating a sister-relationship of the latter to the Anateae (Fig. 2-6). Under “four-group” weighting, the alternative treatment of two ordered characters as unordered alone resulted in the placement of the Cair- ineae as the sister-group of other Anatinae, along with an associated placement of Plectropterus as sister-group of Tadorninae + Anatinae. Under successive weighting, the Cairineae were inferred to be the sister-group of the “tadornine” clade comprising Plectropterus and Sarkidiornis (Fig. 7, 11); this was the only significant discrepancy between the findings of a priori and successive weighting schemes that were robust to bootstrapping. Pending further study, I recommend tentatively retaining the Cairineae, Nettapodeae, and Anateae as subtribes of the Anatini (as in Livezey, 1991), while noting the likely monophyly of the first two subtribes {contra Livezey, 1991) and the less well-supported alternative placement of the Cairineae (apart from the Nettapodeae) with Plectropterus and Sarkidiornis {contra Livezey, 1986<2, 1991). One analytical artifact of parsimony analysis is “long-branch attraction” or the tendency for branches showing many character changes to be joined under certain analytical circumstances (Felsenstein, 1978; Huelsenbeck and Hillis, 1993; Penny et al., 1994; Hillis, 1995; Huelsenbeck, 1995). Three couplets of taxa united in this analysis might be vulnerable to this artifact (Fig. 6): (1) Plectropterus and Sarkidiornis, (2) Merganetta and Tachyeres (perhaps also including Hymenolai- mus), and (3) Malacorhynchus and Salvadorina. The extent to which these group- ings were fostered by “long-branch attraction” cannot be tested at present, but the possibility should be considered in any assessment of these groupings of problematic genera. 62 Annals of Carnegie Museum VOL. 65 Apparent intertribal relationships within the Anatinae indicated in this analysis are among the weakest aspects of the study. None of the weighting schemes provided well-supported resolution of relationships among the Cairineae, Netta- podeae, Anateae, Aythyini, Mergini, and Oxyurini (Fig. 8-11). A sister-group relationship between the Mergini and Oxyurini was indicated in a number of the shortest topologies recovered (Fig. 3-6), an inference based on a single apomor- phy related to the diving-related modification of the pelvic girdle and very pos- sibly convergent (Appendix 1). The agreement between this inference and that indicated in the single tree depicted by Livezey (1986a) provides no independent confirmation because both studies shared most osteological characters, and fur- thermore supplementary analysis of the matrix used by Livezey (1986a) revealed that other arrangements were equally parsimonious (J. Harshman, personal com- munication). Comparison with Previous Studies. — Congruence among independent studies is an important component of the assessment of phylogenetic reconstruction (Mickevich and Johnson, 1976; Mickevich, 1978; Hillis, 1987, 1995; Miyamoto and Fitch, 1995). Unfortunately, little phylogenetic work has been performed for the taxa of concern here, a deficiency particularly marked for molecular data and which significantly limits the comparisons that are possible. Support in this anal- ysis for a clade comprising Plectropterus and Sarkidiornis as the sister-group of the four genera of “typical” shelducks and sheldgeese (Fig. 2-6) represents a significant departure from the position inferred for Plectropterus by Livezey (1986a). This revised inference, however, is consistent with the close relationship between these genera proposed by most recent workers (Delacour and Mayr, 1945; Delacour, 1959; Johnsgard, 1961a, 1965a, 1978, 1979). However, available evi- dence confirms the polyphyletic nature of the “Tribe Cairinini” as generally con- stituted (comprising nine genera and 13 species; e.g., Johnsgard, 1965a, 1978). The close relationships inferred among the genera Hymenolaimus, Tachyeres, and Merganetta (Fig. 1-11) are consistent with several other anatomical and be- havioral characteristics. All three are extremely territorial, even by tadornine stan- dards, and this pugnacity is shown by both sexes (Wright, 1965; Kear and Steel, 1971; Kear, 1972; Weller, 1976; Eldridge, 1985, 1986a, 1986^7; Livezey and Hum- phrey, 1985; Veltman and Williams, 1990). Also, in all three genera, territorial combat often involves blows using the variably developed metacarpal wing-knobs (Kear and Steel, 1971; Weller, 1968; Livezey and Humphrey, 1985; Eldridge, 1986^). In part because of inadequately defined states and a paucity of compar- ative data, no consistent indications of phylogenetic relationships are indicated by courtship displays and other ritualized behaviors in these three genera (Moynihan, 1958; Johnsgard, 1961a, 1965a, 1966; Eldridge, 1979, 1985, 1986a). The inferred positions of Salvadorina and Malacorhynchus, found herein to be sister-genera and together composing the sister-group of “typical” Anatinae under the two a priori weighting schemes (Fig. 1-11), represent significant departures from that inferred for Malacorhynchus by Livezey (1986a) or those assumed for Salvadorina by Livezey (1986a, 1991). The present hypothesis seems intuitively consistent with the “subfamilial intermediacy” implied by the placements of these genera as tadornine or as basal, “aberrant” anatines (Delacour and Mayr, 1945; Delacour, 1959; Johnsgard, 1965a, \965b, 1978) and is consistent with skeletal similarities noted by Mlikovsky (1989). This analysis indicates that the characters shared between Salvadorina and the genera Hymenolaimus and Merganetta cited by Kear (1975) are either plesiomorphous or homoplasious; however, the rec- 1996 Livezey — T adornine-Anatine Divergence 63 Fig. 13. — Phylogenetic tree summarizing the relationships among 19 taxa of Anatidae as depicted or described by Johnsgard (1961a, 1978), the basis for the analytical constraint used to estimate additional steps required to accommodate this hypothesis using the characters defined herein. Regions for which contradictory or inadequately detailed assessments were given are shown as polytomies. ommendation by Kear (1975) that Salvadorina be recognized as a genus distinct from Anas is supported. The territorial behavior of Salvadorina (Hallstrom, 1956; Kear, 1975) — shared by the two aforementioned genera, most Tadorninae, and some basal Anatini — also appears to be locally plesiomorphous. Although Kear (1975) reported that Salvadorina uses wing knobs in combat, the presence of “true” calcar alae (character 30) in this genus is unlikely based on examination of skin specimens (Appendices 1, 2). Searches under topological constraints (Table 4) were employed to provide quantitative assessments of the “penalties” (minimal increases in total tree length) of preserving groupings suggested by other classifications and hypotheses. To preserve as monophyletic the traditionally constituted “perching ducks” (assum- ing no specific topology within the group) requires, depending on weighting scheme, an additional 2-12 steps (1-3% increase in total length; Table 4). Con- straining the analyses to preserve as monophyletic the Anatinae as traditionally constituted (e.g., Johnsgard, 1965^3, 1978, 1979), without assuming any constraints regarding topology within the subfamily, requires an additional 10-31 steps or 5- 8% increase in tree length (Table 4). Exclusion of Hymenolaimus from this group revealed that a majority of the additional steps were attributable to the classifi- cation of this genus alone (Table 4). Analyses constrained to preserve all of the intergeneric relationships depicted by Johnsgard (1961a, 1978), graphically sum- marized in Figure 13, indicated that 36-117 additional steps were required using the present data matrix, representing an increase in total tree length of 17-25% (Table 4). A similar constrained analysis determined that the tree depicted by Livezey (1986a), to the level of detail permitted by taxa analyzed in both studies. 64 Annals of Carnegie Museum VOL. 65 would require 16^3 extra steps (6.1-18.7% increase in total length), depending on weighting scheme (Table 4). A sister-relationship between Stictonetta and the Oxyurini (cf. Fullagar et al., 1990; Marchant and Higgins, 1990), without assuming any specific placement of the composite group within the tree, requires a minimum of 7-3 1 additional steps, a 3-7% increase in total tree length (Table 4). Movement of Stictonetta with the Oxyurini (cf. Marchant and Higgins, 1990) to the position proposed for the latter by Sibley and Monroe (1990) — as sister-group of all Anseriformes exclusive of Anhimidae, Anseranas, Dendrocygna, and Thalassornis — requires several steps in addition to those required simply to link the Oxyurini with Stictonetta (Livezey, in press c). Proposed Classification of Modern Members Subfamilial Taxa. — Based on the phylogenetic inferences using the weighting schemes considered to favor the most reliable characters (conservatively sum- marized in Fig. 3, preferred variant shown in Fig. 4), I recognize three subfamilies within the present group of 19 taxa (Appendix 3): Stictonettinae (monotypic, Stictonetta naevosa), Tadorninae (three modern tribes, ten modern genera), and Anatinae (five modern tribes, one new, and one incertae sedis). This subfamilial taxonomy differs from that proposed by Livezey ( 1 986a) in the inclusion of Plec- tropterus, formerly a monotypic subfamily Plectropterinae Gray, 1840, as a tribe (with Sarkidiornis) in the Tadorninae (Appendix 3). Tribes of Tadorninae. — The Tadorninae here reconstructed (Fig. 3-6), comprise three tribes (Appendix 3): Tribe Merganettini, comprising Hymenolaimus, Tach- yeres, and Merganetta\ Tribe Plectropterini, comprising Plectropterus and Sarki- diornis', and Tribe Tadornini, comprising the subtribes Chloephageae (Cyanochen, Alopochen, Neochen, Chloephaga) and Tadorneae {Tadorna). Differences from the generic classification given by Livezey (1986a) include: (1) inclusion of Plec- tropterus (formerly a separate subfamily) with Sarkidiornis (formerly a separate tribe) as sister-groups in a single tribe of the Tadorninae; (2) monophyly of the Tadornini, in which the subtribe Tadorneae (comprising Tadorna) is the sister- group of the sheldgeese (formerly subtribes Cyanocheneae and Chloephageae; Livezey, in press a); and (3) exclusion of Malacorhynchus from the subfamily. Furthermore, contrary to the provisional allocation by Livezey (1991), Salvador- ina is not included in the Tadorninae. Tribes of Anatinae. — Five modern tribes are recognized in this subfamily (Ap- pendix 3), in which the order of all but the first is unresolved, hence the subfamily is annotated sedis mutabilis (Fig. 3, 4): Tribe Malacorhynchini, new taxon, com- prising Malacorhynchus and Salvadorina', Tribe Anatini, incertae sedis, tentative- ly comprising three subtribes Cairineae, Nettapodeae, and Anateae (Livezey, 1991); Tribe Aythini (Livezey, in press Z?); Tribe Mergini (Livezey, 1995^?); and Tribe Oxyurini (Livezey, 1995c). Generic composition of the Aythyini, Mergini, and Oxyurini remains unchanged from that proposed by Livezey (1986a), and the provisional generic composition of the Anatini advocated here follows that pro- posed by Livezey (1991). Provisional retention of the three subtribes of the An- atini recognized by Livezey (1991) is recommended, in part, to avoid the con- fusion that would stem from applying the tribal taxon “Cairinini,” traditionally including nine genera and 13 species, to a restricted group comprising only the Cairineae (three genera, five species) and Nettapodeae (two genera, four species), and excluding Plectropterus, Sarkidiornis, Amazonetta, and Callonetta. 1996 LIVEZEY T ADORNINE-An ATINE DIVERGENCE 65 Bio geo graphical Patterns Biogeographical patterns in this region of the phylogeny of Anseriformes are complicated by the broad but different distributional patterns shown by several taxa (e.g., Anateae, Aythyini, and Oxyurini). Furthermore, comparisons with the distributional limits of tribes given by Weller {\96Ad) are problematic because of important differences in the composition of tribes (especially Cairinini, Anatini) defined in this study and those used by Weller {\96Ad). In spite of these diffi- culties, the frequency of predominantly southern-hemisphere genera in these taxa {Stictonetta, Plectropterus, Sarkidiornis, most Tadornini, Hymenolaimus, Tachy- eres, Merganetta, Salvadorina, Malacorhynchus), as well as the southern distri- butions of basal members of the Anatini (Livezey, 1991) and Oxyurini (Livezey, 1995c), corroborate the hypothesis of southern origins for the Stictonettinae, Ta- dorninae, and Anatinae. This inference is consistent with a larger hypothesis of a southern-hemispheric origin for the Anseriformes (Livezey, \9^6a, 1989). The sister-relationship inferred for Merganetta and Tachyeres (Fig. 1-11) is at least consistent with the Neotropical distributions of the genera (Delacour, 1954, 1959). Similarly, the distributional limits of another counterintuitive pair of genera inferred to be sister-groups, Salvadorina and Malacorhynchus, is geographically “parsimonious” given the Australasian distributions of the two genera (Delacour, 1959). Several fossil taxa assignable to this region of the anseriform phylogeny also indicate the predominantly southern-hemisphere distributions of the member taxa. The peculiar genus Euryanas finschi from New Zealand (Van Beneden, 1875, 1876; Oliver, 1930, 1945) was most parsimoniously placed as the sister-group of the present ingroup exclusive of Stictonetta in an earlier, more-limited analysis (Livezey, 1989). Another subfossil endemic of New Zealand, Pachyanas chath- amica (Oliver, 1955), also appears to belong to the Tadornini (unpublished study of material recently collected by P. Millener). Other fossil taxa assignable to the Tadornini include the Madagascan genera Centrornis (Andrews, 1897) and Chen- alopex (Andrews, 1897; Brodkorb, 1964), and the Nearctic genera. Anabernicula (Ross, 1935; Howard, 1946, 1964<2, 1964Z?; Short, 1970) and Brantadorna (How- ard, 1963). Directions for Future Study A number of aspects of anseriform phylogeny remain poorly understood, and a majority of these involve the relationships among the taxa analyzed in the present analysis (Fig. 1-11). This state of affairs is ironic, given that the original motivation for the earlier analysis (Livezey, 1986r?), which formed the basis for this and other species-level studies, was the placement of the genus Tachyeres among the Tadorninae and Anatinae. This situation is particularly vexing in that these poorly resolved segments of the phylogeny of waterfowl include those among most tribes of the Anatinae, the largest subfamily in the order and one of particular importance for comparative studies (Sigurjonsdottir, 1981; Faith, 1989; Scott and Clutton-Brock, 1989). Poor resolution, combined with a great diversity of physical and ecological characters among genera in the vicinity of the “tador- nine-anatine divergence,” rendered many ecomorphological parameters all but noncomparable in this study (Fig. 12 and associated text). In order to resolve the phylogenetic relationships that remain resistant to anal- ysis, characters (morphological, molecular, or behavioral) of intermediate evolu- 66 Annals of Carnegie Museum VOL. 65 tionary conservatism are needed. Characters of greatest utility would be those which are less conservative than those uniting the Tadorninae and Anatinae, but more conservative than those accumulating as autapomorphies in the present anal- ysis (some of which are useful, however, as synapomophies for polytypic genera). Morphological systems deserving of particular attention in this regard are acces- sible only using anatomical (fluid-preserved) specimens, however, and therefore will not be useful for the placement of fossil taxa in this region of the anseriform phytogeny. These systems include the anatomical details of tracheal and syringeal myology and osteology, cranial and pelvic myology, pteryolography, and bucco- lingual integument. Acknowledgments This research was supported by National Science Foundation grants BSR-85 15523, BSR-9129545, and BSR-9396249, as well as grants from the National Museum of Natural History and the American Museum of Natural History. The hospitality of G. Mack and R. L. Zusi is much appreciated. In addition to access to specimens in the collections of Carnegie Museum of Natural History, I am grateful for loans and/or access to collections provided by the curatorial personnel of the following institutions: Division of Birds, National Museum of Natural History, Washington, D.C. (USNM); Department of Ornithology, American Museum of Natural History, New York, New York (AMNH); Division of Birds, Museum of Zoology, University of Michigan, Ann Arbor, Michigan (UMMZ); Division of Birds, Field Museum of Natural History, Chicago, Illinois (FMNH); Bird Group, Natural History Museum, Tring, Hertfordshire, United Kingdom (BMNH); Wildfowl and Wetlands Trust, Slim- bridge, Gloucester, United Kingdom (WWT); Department of Ornithology, Royal Ontario Museum, Toronto, Ontario, Canada (ROM); Department of Ornithology, Australian Museum, Sydney, Australia (AUM); and Division of Ornithology, Natural History Museum, University of Kansas, Lawrence, Kansas (KUMNH). I thank M. 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Transactions of the Linnean Society of London, 15:378-391. Appendix 1 Character Descriptions Characters are numbered continuously through skeletal, tracheal, natal, and de- finitive subgroups. Characters for which polarity remains undetermined are indi- cated by asterisks. Consistency indices (Cl) listed are those associated with trees derived under equal weighting of characters; ranges of indices are given for those characters in which CIs varied among equally parsimonious trees. 1996 LiVEZEY T ADORNINE-An ATINE DIVERGENCE 75 Fig. 14. — Maxillae of selected genera of Anatidae, laterodorsal views: A — Hymenolaimus malacor- hynchos (KUMNH 55871), B — Malacorhynchus membranaceus (UMMZ 204438), C — Merganetta {a.) armata (UMMZ 153796). Skeleton. — Those adapted from Livezey (1986a) are revised to include only rel- evant states and to assume local polarity. Synapomorphies for genera within sub- tribes of Anatini from Livezey (1991), as well as tribal synapomorphies for Ay- thyini (Livezey, in press b), Mergini (Livezey, 1995a), and Oxyurini (Livezey, 1995Z?), are not included. Characters 15, 33, and 61 from Livezey (1986a), and character 156 from Livezey (1991) are excluded as unreliable for these groups. States for Salvadorina are unknown except as determined from skin specimens, Mayr (1931), and Mlikovsky (1989). Nomenclature follows Baumel and Witmer (1993). 1. Maxilla, typical shape: (a) not “subconical”; (b) “subconical,” with es- sentially straight culmen (Alopochen intermediate, variable, sexually dimorphic). Note — see Fig. 14. Cl = 0.50. 2. Maxilla and (to lesser degree) mandibula, distinct recurvature: (a) absent, (b) present {Stictonetta extreme, T. tadorna most conspicuous of genus). Cl = 0.50. 3. Maxilla and mandibula, distally spatulate conformation: (a) absent, (b) present {Malacorhynchus; convergent state in subgenus Spatula of Anas). (Liv- ezey 1991: character 157.) Note — see Fig. 15. Cl = 1.00. 4. Cranium, os lacrimale, processus supraorbitalis and os frontale, margo la- teralis, prominent enclosed foramen (not to be confused with minute, largely oc- cluded foramina neurovascularia): (a) absent, (b) present {Alopochen variable, obscured in adult males), (a/b) polymorphic (derived state in Cairino). Cl = 1.00. 5. Cranium, os lacrimale, processus supraorbitalis: (a) essentially lacking or small, straight, and coplanar with dorsal surface of skull; (b) comparatively large, flat, often triangular in dorsal aspect, appressed to laterodorsal margin of orbit; (c) large, thick, rugose, dorsolaterally prominent; (d) long, slender, dorsally prom- inent. (Livezey 1986a.character 11.) Note — state “b” affected in appearance by 76 Annals of Carnegie Museum VOL. 65 Fig. 15. — Mandibulae of selected genera of Anatidae, ventral views: A — Hymenolaimus malachor- hynchos (KUMNH 55871), B — Malacorhynchus membranaceus (UMMZ 204434). variably developed sulcus glandulae nasalis in some genera; this and other aspects of os lacrimale in Stictonetta confounded by unique synostosis of processus or- bitalis with dorsally ossified Membrana jugolacrimale. Cl = 0.75-1.00. 6. Cranium, ossa frontales, pneumatic dorsal convexity (males): (a) absent; (b) present, rounded, confined to region immediately caudal to zona flexoria cran- iofacialis; (c) present, variably produced into two-parted prominence, the more- caudal one bilaterally compressed and interorbital in position. Note — revised state for Cyanochen based on examination of skins, few skeletons available inadequate for diagnosis of this variably developed, sexually dimorphic character; see Moller (1969) and Livezey (in pressa, in pressc). (Livezey, 1986a.*character 16, revised, corrected.) Cl = 1.00. 7. Cranium, basis cranii, lamina parasphenoidalis: (a) rounded; (b) distinctly convex ventrally, with prominent medial ridge (Anateae). (Livezey, 1991:char- acter 155.) Cl = 1.00. 8. Vertebrae cervicales, modal number: (a) 17; (a/b) polymorphic, 16-17; (b) 16. (Livezey, 1986a.-character 21, revised.) Cl = 0.86. 9. Sternum, corpus sterni, foramen pneumaticum: (a) present, (b) absent. (Liv- ezey, 1986a.character 78.) Cl = 0.50. 10. Sternum, rostrum sterni, spina externa: (a) lateromedially compressed flange (ancestor); (b) elongate, peg-like spine; (c) small or obsolete. (Livezey, 1 986<2.‘character 79, revised significantly.) Cl = 0.50-0.67. 1 1 . Sternum, rostrum sterni, spina interna: (a) absent; (b) present, small. (Liv- ezey, 1986a.character 82 [incorrect for Chenonetta]; Livezey, 1991 : character 151.) Cl = 0.33-1.00. 1996 Livezey — Tadornine-Anatine Divergence 77 12. Sternum, carina stemi, margo ventralis, profile in lateral view: (a) distinctly convex throughout length, (b) essentially straight for caudal half. (Livezey, 1986a: character 80.) Cl == 1.00. 13. Sternum, trabecula lateralis: (a) essentially equal in caudal extent to caudal terminus of linea intermusculus, (b) extend well caudad to caudal terminus of linea intermusculus. (Livezey, 1986a:character 81, revised.) Cl — 1.00. 14. Furcula, extremitas omalis claviculae, processus acrocoracoideus: (a) pres- ent, although of variable conspicuousness; (b) obsolete. (Livezey, 1986a:character 101.) Cl = 0.50. 15. Furcula, extremitas stemalis claviculae, apophysis furculae: (a) present, (b) obsolete. Note— -see Woolfenden (1961), Mlikovsky (1989). Cl = 0.33. 16. Scapula, extremitas cranialis scapulae, tuberculum coracoideum: (a) copla- nar with corpus scapulae, margo ventralis; (b) protrusive, distinctly ventral to corpus scapulae, margo ventralis. (Livezey, 1 986a;character 107.) Cl = 1.00. 17. Coracoideum, extremitas omalis coracodei, processus acrocoracoideus, subtending pneumatic depression: (a) absent, (b) present. (Livezey, 1986a;char- acter 95, Livezey, 1991: character 154.) Cl = 1.00. 18. Coracoideum, corpus coracoidei, facies ventralis, impressio m. supracora- coideus: (a) absent, (b) present. (Livezey, 1986a;character 96.) Cl = 1.00. 19. Coracoideum, extremitas sternalis coracoidei, facies articularis sternalis, crista ventralis: (a) ventrally prominent, forming buttress; (b) inconspicuous. (Liv- ezey, 1986a:character 100.) Cl = 0.50. 20. Humerus, extremitas proximalis, facies caudalis immediately distal to ca- put: (a) variably convex; (b) with transverse trench-like depression, distinctly concave. (Livezey, 1986a;character 24.) Cl = 1.00. 21. Humerus, extremitas proximalis, crista deltopectoralis: (a) margo dorso- cranialis prominent, rounded, facies caudalis concave; (b) margo dorsocranialis angular, facies caudalis convex. (Livezey, 1986a:character 25.) Cl = 0.25-0.50. 22. Humerus, extremitas proximalis humeri, tuberculum ventrale: (a) compar- atively proximal, exposing fossa pneumotricipitalis in caudal view; (b) compar- atively distal, largely or completely obscuring fossa pneumotricipitalis in caudal view. (Livezey, 1986a;character 27.) Cl = 0.50. 23. Humerus, extremitas proximalis humeri, fossa pneumotriciptalis, foramen pneumaticum: (a) present, (b) absent. (Livezey, 1986a;character 28.) Cl = 1.00. 24. Humerus, extremitas proximalis humeri, tuberculum dorsale: (a) prominent, buttressed; (b) not prominent, buttressed. (Livezey, 1986a:character 32.) Cl = 0.25-0.33. 25. Humerus, corpus humeri, margo caudalis (“capital shaft ridge”): (a) prom- inent, proximally directed toward caput humeri; (b) prominent, proximally di- rected toward tuberculum dorsale; (c) obsolete. (Livezey, 19 86a: character 22, or- dering not assumed.) Cl = 0.40-0.67. 26. Humerus, extremitas distalis humeri, tuberculum supracondylare ventrale: (a) not elevated, essentially coplanar with corpus humeri; (b) elevated, angled distally; (c) elevated, angled medially. (Livezey, 1986a:character 26.) Cl = 0.50. 27. Os carpi radiale: (a) not modified into spur, (b) modified into spur. (Liv- ezey, 1986a:character 49.) Note — see Rand (1954). Cl = 1.00. 28. Carpometacarpus, extremitas proximalis carpometacarpi, trochlea carpalis, labrum dorsalis, distal terminus, prominent swelling: (a) present, (b) absent. (Liv- ezey, 1986a:character 37.) Cl = 1.00. 29. Carpometacarpus, extremitas proximalis carpometacarpi, processus flexo- 78 Annals of Carnegie Museum VOL. 65 rius, orientation relative to shaft (ordered): (a) perpendicular to or proximally directed, (b) slightly distally directed, (c) distinctly distally directed. (Livezey, 1 986<3;character 41, revised.) Cl = 1.00. 30. Carpometacarpus, extremitas proximalis carpometacarpi, os metacarpale al- ulare, processus extensorius, enlargement into “spur” (calcar alae): (a) absent; (b) present, especially prominent in adult males. (Livezey, 1986a:character 42.) Cl = 0.33-0.50. 31. Carpometacarpus, extremitas proximalis carpometacarpi, fovea carpalis caudalis: (a) present but comparatively shallow; (b) present, comparatively deep, distinctly ovate. (Livezey, 1986fl.character 46.) Cl = LOO. 32. Carpometacarpus, extremitas proximalis carpometacarpi, trochlea carpalis, labrum ventralis: (a) of uniform depth, (b) distinctly thickened proximally. (Liv- ezey, 1986a.-character 47.) Cl = 0.50. 33. Carpometacarpus, corpus carpometacarpi, os metacarpale majus, facies dor- salis, impressio m. extensor metacarpi ulnaris, position relative to synostosis me- tacarpalis proximalis: (a) opposite, at least partly; (b) completely proximal. (Liv- ezey, 1986<2.-character 43, revised.) Cl = 0.50. 34. Illium, ala preacetabularis ilii, facies dorsalis, fossa iliaca dorsalis: (a) pre- senting smooth, only moderately concave surface; (b) containing deep, irregularly shaped depression. (Livezey, 1986a.character 113.) Cl = 1.00. 35. Pubis, scapus pubis: (a) concave dorsally, (b) convex dorsally. (Livezey, 1986a.character 115.) Cl - 0.50-1.00. 36. Femur, corpus femoris, craniocaudal curvature (lateral perspective): (a) ab- sent or slight, (b) moderate. (Livezey, 1 986a.-character 55.) Cl = 0.50. 37. Femur, corpus femoris, facies caudalis, linea intermuscularis caudalis: (a) present but not prominent; (b) prominent, produced into overhanging ridge prox- imally. (Livezey, 1986«.character 60.) Cl — 1.00. 38. Femur, extremitas distalis femoris, fossa poplitea: (a) shallow, (b) deep. (Livezey, 1986a.character 56.) Cl = 1.00. 39. Tarsometatarsus, corpus tarsometatarsi, facies dorsalis, margo medialis, dorsal prominence relative to margo lateralis: (a) essentially equal, (b) distinctly less prominent. (Livezey, 19 86a. character 75.) Cl = LOO. Trachea (Fig. 16). — Pertain to males. Nomenclature follows King (1989, 1993). (Livezey, 1986a.character 7 incorrect for Stictonetta, excluded.) 40. Asymmetrical bulla syringealis: (a) absent, (b) present (Malacorhynchus vestigial). (Livezey, 1986a.'character 6, revised.) Cl = 0.50. 41. Syrinx, narrow constriction between tympanum and bulla: (a) absent, (b) present. Note — see Garrod (1875). Cl = 1.00. 42. Bulla syringealis (if present), fenestrae: (a) absent; (b) present, but rela- tively prominent, widely distributed; (c) present, small, few, confined to craniola- teral margin. Cl = 1.00. 43. Bilobate bulbus trachealis: (a) absent; (b) present, males. Note — see Camp- bell (1899). Cl = 1.00. Natal Integument (Fig. 17). 44. Ground color of venter*: (a) yellow, (b) dusky, (c) white, (x) basal state not determinable (Mergini). Note — polarity difficult, see Livezey, 1991: character 121. Cl = 0.50. 1996 Li VEZEY T ADORNINE- An ATINE DIVERGENCE 79 Fig. 16. — Bullae syringeales of selected genera of Anatidae, males, ventral views: A — Plectropterus gambensis (USNM 431245), B — Sarkidiornis melanotus (AMNH 15667), C — Hymenolaimus mala= corhynchos (BMNH 1959- 14-1), D — Tachyeres patachonicus (KUMNH 77972), E — Merganetta {ar- mata) colombiana (ROM 93343), F — Malacorhynchus membranaceus (UMMZ 204438), G — Salva- dorina waigiuensis (WWT 1752). 45. Dark, vertical supraorbital stripe: (a) absent, (b) present. Cl = 0.50. 46. Dark auricular spot: (a) absent, (b) present, (x) noncomparable. Cl = 0.20~ 0.250. 47. Pale scapular spots*: (a) absent, (b) present, (x) unknown or basal state not determinable. Cl = 0.33. 48. Pale rump spots*: (a) absent, (b) present {Hymenolaimus rufous), (x) un- known or basal state not determinable. Cl = 1.00. 49. Dorsal scapular and rump spots (if present): (a) separate, (b) confluent. Cl = 0.33. 50. Pale, medial mantle stripe: (a) absent, (b) present. Cl = 1.00. 51. Foot color: (a) gray(ish), (b) bright yellow or orange. Cl ~ 1.00. 52. Dusky, oblique facial stripes: (a) absent, (b) present. Note — intraspecifi- cally variable. Cl = 1.00. 53. Sharp, dark preorbital (loral) stripe: (a) present, (b) absent (Cairineae ex- cluding Pteronetta). (Livezey, 1991: character 107.) Cl = 0.33. 54. Broad, pale supraorbital stripe, continuing caudoventrally around dark or- bital patch along sides of neck: (a) absent, (b) present. Cl = 1.00. Definitive Integument.— Chsucacter 101 of Livezey (1991) was deemed not reliably characterized at this scale and excluded. Redundant synapomorphies for anatine tribes (see other works), as well as characters defining clades within genera of Tadornini (Livezey, in press a) or clades within Tachyeres (Humphrey and Liv- ezey, 1985; Corbin et al., 1986; Livezey, 1986a; Livezey and Humphrey, 1992), also were excluded. 55. Rectrices: (a) not as follows (including narrow elongate tails of Merga- netta, Salvadorina); (b) long and broad, typically extending beyond feet in pre- pared skin specimens. (Livezey, 1991 : character 1.) Cl = 0.50-1.00. 80 Annals of Carnegie Museum VOL, 65 1996 LiVEZEY T ADORNINE-An ATINE DIVERGENCE 81 56. Dark, variably extensive postorbital stripe (juvenal and definitive basic plumages of both sexes, as well as definitive alternate of females in some): (a) absent, (b) present (possibly excluding micro-teal, Heteronetta). (Livezey, 1991: character 128.) Cl = 0.50-1.00. 57. Black, lateromedially compressed comb-like crista carnosa on culmen: (a) absent, (b) present. Cl = 1.00. 58. Sharp, black speckling of white face: (a) absent (but see Cairina scutu- lata), (b) present. Cl = 1.00. 59. Narrow, black shoulder marks at cranial margin of darkened sides: (a) absent, (b) present (obscured in S. sylvicola). Cl = 1.00. 60. Lesser dorsal wing coverts, contrasting pale (typically white) color: (a) absent {Plectropterus nonhomologous, see character d8), (b) present. Cl = 0.33. 61. Lesser dorsal wing coverts, contrasting pale color (if present): (a) white, (b) blue. Cl = 1.00. 62. Cranialmost lesser dorsal wing coverts, contrastingly white producing nar- row white leading edge of wing: (a) absent, (b) present. Cl = LOO. 63. Jagged, black coloration on sides: (a) absent, (b) present. Cl = 1.00. 64. Bare, variably colored facial region (from bill to eye to auricular region, including ramal and interramal areas of mandibles) in males: (a) absent, (b) pres- ent. Cl 0.50. 65. Single, bare, fleshy-red patches on sides of neck of adult males: (a) absent, (b) present. Cl = 1.00. 66. Crista nuchalis pennarum of black-tipped, white feathers from corona to base of neck: (a) absent, (b) present. Cl = 1.00. 67. Sides of head and neck, variable suffusion with yellowish buff: (a) absent, (b) present. Cl = 1.00. 68. Undertail coverts, variable suffusion with yellowish buff: (a) absent, (b) present. Cl = 1.00. 69. Well-defined single black lines from sides of rump extending ventrally and demarcating the belly from the undertail coverts: (a) absent, (b) present. Cl = 1.00. 70. Well-developed tomial flaps of maxilla, opposed by lateral extensions of mandibular: (a) absent, (b) present (Malacorhynchus extreme). Cl = 0.50. 71. Maxilla and mandibula, narrow, submergine conformation: (a) absent, (b) present. Cl = 1.00. 72. Mandibular rami, lateral compression defining long, very narrow inter- ramal region: (a) absent, (b) present. Cl = 1.00. 73. Black-and-white scalloping on sides and flanks: (a) absent, (b) present (terminal white band narrower in Salvadorina) . Cl = 1.00. 74. Sharply defined gray-and-chestnut breast and sides, contrasting sharply with white venter: (a) absent, (b) present. Cl = 1.00. Fig. 17. — Facial patterns of natal plumages of the Freckled Duck and selected genera of Anatidae, lateral views: A — Stictonetta naevosa (AUM 0.60097), B — Plectropterus gambensis (ROM 91832), C — Sarkidiornis melanotus (AMNH 731312), D — Hymenolaimus malacorhynchos (WWT 2359), E — Tachyeres patachonicus (USNM 48560), F — Merganetta (armata) turneri (FMNH 208125), G — Ma- lacorhynchus membranaceus (WWT 3572), H — Salvadorina waigiuensis (based on Kear, 1975:fig 2; see also Delacour, 1956:pl. XXIII). 82 Annals of Carnegie Museum VOL. 65 15. Narrow white barring of mantle: (a) absent, (b) present (comparatively extensive, including entire dorsum in Salvadorina) . Cl = 1.00. 76. Narrow white terminal white tips of secondary remiges: (a) absent, (b) present. Cl = 1.00. 77. Sharply defined black scalloping on otherwise white wing linings, ex- tending to axillars: (a) absent, (b) present (marks on axillars of Salvadorina tend- ing to spotting). Cl = 1.00. 78. Modal number of pairs of rec trices (ordered): (a) five, (b) six, (c) seven, (d) nine. Cl = 0.75. 79. Rectrices: (a) not elongate, narrow; (b) elongate, narrow. Note — see char- acter 55. Cl = 0.50. 80. Black-and-white barring of rump: (a) absent, (b) present. Cl = 0.50. 8 1 . Leg color: (a) black or gray, (b) yellow, (c) red. Cl = 0.50. 82. Bill color (breeding males): (a) gray; (b) pinkish white; (c) orange or red, including pink and coral; (d) yellow; (a/c) polymorphic (Anateae). Cl = 0.67. 83. Wing linings, fine vermiculations: (a) absent, (b) present. Cl = 1.00. 84. Major secondary coverts tipped with narrow white(ish) bars, producing cranial border to speculum (if any): (a) absent (Malacorhynchus, caudal only; T. radjah, broad, not homologous), (b) present (Anateae, excluding microteal). Cl = 0.33. 85. Rufous underparts (females): (a) absent, (b) present. Cl - 1.00. 86. Bifurcating, black, postorbital stripe: (a) absent, (b) present. Cl = 1.00. 87. Tarsus, dorsal surface: (a) reticulate, (b) scutellate. Cl = 1.00. 88. Contrastingly white venter: (a) absent, (b) present. Cl = 1.00. 89. Wing spurs, prominent claws: (a) absent, (b) present. Cl = 1.00. 90. Uniformly gray dorsal wing coverts: (a) absent, (b) present. Cl = 1.00. 91. Rump, ground color: (a) brown(ish); (b) black; (c) uniformly gray; (d) white; (a/b) polymorphic, black or brown. Cl = 0.57-0.67. 92. Dorsal chestnut wash: (a) absent, (b) present. Cl = 0.33. 93. Uniformly black tertials (in some extending to scapulars or entire dor- sum): (a) absent, (b) present. Cl = 0.25-0.33. 94. Demarcation of color between neck and breast: (a) absent, (b) present. Note — see Livezey (in press a). Cl = 0.50. 95. Lobation of hallux, and enlargement of subungual lobes in other digits: (a) absent (including basal Aythyini, Oxyurini), (b) present. Cl = 0.33-0.50. 96. Dorsum entirely glossy black: (a) absent, (b) present. Cl = 0.50-1.00. 97. Dorsum, iridescent greenish (to purplish) color on dark dorsum of wings: (a) absent; (b) present, including all of dorsal surface of wing except distal pri- maries and (in some) leading edge; (c) present, limited to secondary coverts (Cair- ineae in part, Nettapodeae variable); (d) present, limited to secondary remiges (Chloephageae variable). Cl = 0.75-1.00. 98. Secondary remiges, ground color: (a) brown(ish), (b) iridescent greenish black (including tribes in which border produces speculum), (c) largely white, (d) gray. Cl = 0.43-0.60. 99. Wing linings: (a) grayish brown, in some mottled with whitish {Mala- corhynchus white with brown bars); (b) “pure” white; (c) black; (d) gray. Cl = 0.43-0.60. 100. Pedal unguis: (a) not strongly developed, curved; (b) strongly developed, curved, subraptorial. Cl — 0.50-1.00. 101. Unguis of pedal digit III: (a) not qualitatively distinct from other digits; 1996 LivEZEY — Tadornine-Anatine Divergence 83 (b) distinctly broadened with marked concavity on plantar surface, medially dis- placed with enlarged lobate cutaneous pad enclosed in plantar cavity. Cl = 1 .00. 102. Interdigital webs of pedal digits, deep incisura: (a) absent, (b) present. Cl = 1.00. 103. Bill lamellae, extreme ventral elongation and increase in number: (a) ab- sent, (b) present. Cl = 1.00. 104. Bill lamellae, confinement to proximal half of maxilla: (a) absent, (b) present. Cl = 1.00. 105. Dark crown and nape band: (a) absent, (b) present. Note — possibly related to brown head of Salvadorina. Cl = 1.00. 106. White terminal tail band formed by narrow pale tips of rectrices: (a) absent, (b) present. Cl = LOO. 107. Dark periorbital patch: (a) absent, (b) present. Cl = 1.00. 108. Small pink postorbital spot: (a) absent, (b) present. Cl = 1.00. 109. Contrastingly buffy undertail coverts: (a) absent, (b) present. Cl = 1.00. 110. Head and neck contrastingly dark grayish black: (a) absent, (b) present. Cl = 1.00. 111. Uniformly blue-gray ground color of body plumage: (a) absent, (b) pres- ent. Cl = 1.00. 112. Reddish brown mottling of breast, mantle, and wing linings: (a) absent, (b) present. Cl = 1.00. 113. Scapulars, vexilla externum with contrasting, narrow, black margins: (a) absent, (b) present. Cl = 1.00. 114. Venter boldly striped (adult males): (a) absent, (b) present. Cl = 1.00. Attributes For Mapping (primitive states in boldface) A. Body mass (g, ordered): (a) <500, (b) 500-1500, (c) 1500-2500, (d) >2500. B. Sexual size dimorphism (ratio, ordered): (a) 1.05-1.20, (b) 1.21-1.40, (c) >1.40. C. Sexual dichromatism: (a) absent (others); (b) rudimentary, confined to quantitative differences in coloration of plumage and rhamphotheca (Anateae in- cluded here on the basis of states in basal members); (c) pronounced, involving distinct qualitative differences in pattern; (b/c) two states of almost equal fre- quency among members (Cairina and Pteronetta, b; Aix, c). D. Mating system: (a) monogamy, (b) serial polygyny. E. Typical nest site: (a) terrestrial (including over water), (b) cavity. F. Clutch size (median, ordered): (a) 3-5, (b) 6-8, (c) 9-11. G. Paternal attendance of broods: (a) typical (including poorly known Netta- pus), (b) atypical. H. Diving habit: (a) not developed, (b) developed (including only moderately specialized Salvadorina). 84 Annals of Carnegie Museum VOL. 65 Appendix 2 Data Matrix Matrix of 1 14 morphological characters (numbered, described in Appendix 1) used in the phylo- genetic analysis of Stictonetta, genera of “tadornines,” clades of “anatines,” and a hypothetical an- cestor, followed by eight attributes mapped a posteriori (lettered A-H). Skeletal characters are labelled “si” to “s39,” tracheal characters “tl” to “t4,” natal characters “nl” to “nil,” and those of definitive integument “dl” to “d60.” States are coded as lower-case letters, and questions marks (coded as “x” in Appendix 1) signify undetermined states. Character 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Taxon si s2 s3 s4 s5 s6 s7 s8 s9 slO sll sl2 sl3 sl4 sl5 1 Ancestor a a a a a a a a a a a a a a a 2 Stictonetta a b a a a a a a a b a a a a a 3 Plectroptenis a a a a b c a a a c a a a b a 4 Sarkidiornis a a a a b a a b a c a a a a a 5 Cyanochen b a a b b b a a/b a c a a a b a 6 Alopochen b a a b b b a a/b a c a a a b b 1 Neochen b a a b b b a a/b a c a a b b b 8 Chloephaga b a a b b b a a/b a c a a b b b 9 Tadorna a b a a b a a a/b a c a a a b a 10 Hymenolaimus a a a a a a a b a c a a a a b 1 1 Tachyeres a a a a c a a b b c a b a a b 12 Merganetta a a a a a a a b a c a b a a b 13 Salvadorina a a a a 7 a a 7 a b 7 7 a a b 14 Malacorhynchus a a b a a a a b a b b a a a a 15 Cairineae a a a a/b b a a b a c a a a a a 16 Nettapodeae b a a a a a a b a b b a a a a 17 Anateae a a a a a a b b a b b a a a a 18 Aythyini a a a a a a a b a b b a a a a 19 Mergini a a a a d a a b b b a a a a a 20 Oxyurini a a a a a a a b a b b a a a a Character 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 Taxon sl6 sl7 sl8 sl9 s20 s21 s22 s23 s24 s25 s26 s27 s28 s29 s30 1 Ancestor a a a a a a a a a a a a a a a 2 Stictonetta a a b a a a a a a b a a a a a 3 Plectropterus a a a a a a a a a b a b a b a 4 Sarkidiornis a a a b a a b a a b b a a c b 5 Cyanochen a a a b a a b a a c b a a a b 6 Alopochen a a a b a a b a a b b a a a b 1 Neochen a a a b a a b a a b b a a a b 8 Chloephaga a a a b a a b a a b b a a a b 9 Tadorna a a a b a a b a a b b a a a b 10 Hymenolaimus a a a b b b b a b b b a b a b 1 1 Tachyeres b a a b a a b a b b c a b a b 12 Merganetta a a a b a a b a a b b a b a b 13 Salvadorina 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 14 Malacorhynchus a a a b a a b b a b b a a a a 15 Cairineae a b a b a b b a b c b a a a a 16 Nettapodeae a a a b a b b a b c b a a a a 17 Anateae a a a b a b b a b c b a a a a 18 Aythyini a a a b a b b a b c b a a a a 19 Mergini a a a b a b b a b c c a a a a 20 Oxyurini a a a b a b b a b c b a a a a 1996 Livezey — Tadornine-Anatine Divergence 85 Appendix 2 — Continued. Character 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 Taxon s31 s32 s33 s34 s35 s36 s37 s38 s39 tl t2 t3 t4 nl n2 1 Ancestor a a a a a a a a a a a a a a a 2 Stictonetta a a a a a a a a a a a a b b a 3 Plectropterus a a b a a a a a a b b c a a a 4 Sarkidiornis a a a a a a a a a b b a a a a 5 Cyanochen a b a a a a a a a b a a a c a 6 Alopochen a a a a a a a a a b a a a c a 1 Neochen a a a a a a a a a b a a a c a 8 Chloephaga a a a a a a a a a b a a a c a 9 Tadorna a a a a a a a a a b a a a c a 10 Hymenolaimus a b a a a a a b a b a a a c b 1 1 Tachyeres b b b b a a b b b b a a a c b 12 Merganetta a b a a a b a b a b a a a c b 13 Salvadorina 7 7 7 7 7 7 7 7 7 b a a a c a 14 Malacorhynchus a a a a a b a a a b a a a c a 15 Cairineae a a a a a a a a a b a a a c a 16 Nettapodeae a a a a a a a a a b a a a c a 17 Anateae a a a a a a a a a b a a a c a 18 Aythyini a a a a a a a a a b a b a c a 19 Mergini a a a a b a a a a b a a a 7 a 20 Oxyurini a a a a b a a a a a a a a a b Character 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Taxon n3 n4 n5 n6 n7 n8 n9 nlO nil dl d2 d3 d4 d5 d6 1 Ancestor a 7 7 a a a a a a a a a a a a 2 Stictonetta ? a a a a a a a a a a a a a a 3 Plectropterus a b b a a a b a a a a a a a a 4 Sarkidiornis a b b a a a b a a a a b b b a 5 Cyanochen b b b a a a a a b a a a a a b 6 Alopochen b b b a a a a a b a a a a a b 1 Neochen b b b b a a a a b a a a a a a 8 Chloephaga b b b b a a a a b a a a a a b 9 Tadorna b b b a a a a a a a a a a a b 10 Hymenolaimus a a b a a a a a a a a a a a a 1 1 Tachyeres a b b a a b a a a a a a a a a 12 Merganetta b b b b b b a a a a a a a a a 13 Salvadorina b b b a a a a b a a a a a a a 14 Malacorhynchus 7 a b a a a a a a a a a a a a 15 Cairineae a b b a a a a b a b b a a a b 16 Nettapodeae a b b a a a a a a b b a a a a 17 Anateae b b b a a a a a a a b a a a a 18 Aythyini a b b a a a a b a a a a a a a 19 Mergini 7 7 7 a a a a a a a a a a a a 20 Oxyurini b b b b a a a a a a b a a a a 86 Annals of Carnegie Museum VOL. 65 Appendix 2 — Continued. Character 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 Taxon d7 d8 d9 dlO dll dl2 dl3 dl4 dl5 dl6 dl7 dl8 dl9 d20 d21 1 Ancestor a a a a a a a a a a a a a a a 2 Stictonetta a a a a a a a a a a a b a a a 3 Plectropterus a b b b b a a a a a a a a a a 4 Sarkidiornis a a a a a b b b b a a a a a a 5 Cyanochen b a a a a a a a a a a a a a a 6 Alopochen a a a a a a a a a a a a a a a 1 Neochen a a a a a a a a a a a a a a a 8 Chloephaga a a a a a a a a a a a a a a a 9 Tadorna a a a a a a a a a a a a a a a 10 Hymenolaimus a a a a a a a a a b a a a a a 1 1 Tachyeres a a a a a a a a a a a a a b a 12 Merganetta a a a a a a a a a a b a a a a 13 Salvadorina a a a a a a a a a a a a b a b 14 Malacorhynchus a a a a a a a a a b a a b a b 15 Cairineae a a a b a a a a a a a a a a a 16 Nettapodeae a a a a a a a a a a a a a a a 17 Anateae a a a a a a a a a a a a a a a 18 Aythyini a a a a a a a a a a a a a a a 19 Mergini a a a a a a a a a a a a a a a 20 Oxyurini a a a a a a a a a a a a a a a Character 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 Taxon d22 d23 d24 d25 d26 d27 d28 d29 d30 d31 d32 d33 d34 d35 d36 1 Ancestor a a C a a a a a a a a a a a a 2 Stictonetta a a C a a a a a a a a a a a a 3 Plectropterus a a C a a a c a a a a b a a a 4 Sarkidiornis a a c a a a a a a a a b a a a 5 Cyanochen a a c a a a a a a a a b a a a 6 Alopochen a a c a a c c a a a a b a a a 1 Neochen a a c a a c a a a a a b a a a 8 Chloephaga ? a c a a c a a a a a b a a a 9 Tadorna a a c a a c a a a a a b a a a 10 Hymenolaimus a a c a a a b a a a a b a a b 1 1 Tachyeres a a d a a a c a a a a b b a b 12 Merganetta a a a b b c c b b b b b a b b 13 Salvadorina b b c b b b d a b a a b a a a 14 Malacorhynchus b b b a b a a a a a a b a a a 15 Cairineae a a c a a a a a a a a b a a a 16 Nettapodeae a a c a a a a a a a a b a a a 17 Anateae a a c a a a a/c a b a a b a a a 18 Aythyini a a c a a a a a a a a b a a a 19 Mergini a a c a a a a a a a a b a a a 20 Oxyurini a a c a a a a a a a a b a a a 1996 LivEZEY — Tadornine-Anatine Divergence 87 Appendix 2 — Continued. Character 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 Taxon d37 d38 d39 d40 d41 d42 d43 d44 d45 d46 d47 d48 d49 d50 d5l d52 1 Ancestor a a a a a a a a a a a a a a a a 2 Stictonetta a a a a a a a a a a a a a a a a 3 Plectropterus b a b a a b b b b b a b a a a a 4 Sarkidiornis b a b a a b b b c b a a a a a a 5 Cyanochen c a a a a a d b b a a a a a a a 6 Aiopochen b b b b a a d b b a a a a a a a 1 Neochen b b b b a a d c c a a a a a a a 8 Chloephaga d a a b a a d c b a a a a a a a 9 Tadorna b b b b a a d b b a a a a a a a 10 Hymenolaimus c a a a b a a d d a b a a b a a 1 1 Tachyeres c a a a b a a c d a a a a a a a 12 Merganetta c a a a b a a b d a a a a a a a 13 Salvadorina a a a a b a a b a a a a a a a a 14 Malacorhynchus d a a a a a a a a a a a b a b b 15 Cairineae b a b a a b c b c b a a a a a a 16 Nettapodeae a/b a a a a a c b c a a a a a a a 17 Anateae a a a a a a a b c a a a a a a a 18 Aythyini a a a a a a a a a a a a a a a a 19 Mergini a a a a b a a b b a a a a a a a 20 Oxyurini a a a a a a a a a a a a a a a a Character 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 Taxon d53 d54 d55 d56 d57 d58 d59 d60 A B C D E F G H 1 Ancestor a a a a a a a a c a a a a a a a 2 Stictonetta a a a a a a a a b a a a a b a a 3 Plectropterus a a a a a a a a d c a b a/b b b a 4 Sarkidiornis a a a a a a a a c b b b a/b c b a 5 Cyanochen a a a a a a a a c c a a a b a a 6 Aiopochen a a a a a a a a c b a a a c a a 1 Neochen a a a a a a a a b b a a a c a a 8 Chloephaga a a a a a a a a c a/b c a a b a a 9 Tadorna a a a a a a a a b b c a b c a a 10 Hymenolaimus a a a a b b b a b a a a a a a b 1 1 Tachyeres a a a a a a a a d b b a a b a b 12 Merganetta a a a a a a a b a b c a a/b a a b 13 Salvadorina a a a b a a a a a a a a a a a b 14 Malacorhynchus b b b a a a a a a a a a a/b b a a 15 Cairineae a a a a a a a a b/c b/c b/c b b c b a 16 Nettapodeae a a a a a a a a a/b a c a b c a a 17 Anateae a a a a a a a a a/b a b a a b/c a/b a 18 Aythyini a a a a a a a a a/b a c a a b/c b b 19 Mergini a a a a a a a a a/b a c a a/b a/b b b 20 Oxyurini a a a a a a a a a/b a c a/b a a/b b b 88 Annals of Carnegie Museum VOL. 65 Appendix 3 Classification Formation and hyphenation of English group names follow conventions of Parkes (1978); capitalization of those for Sarkidiornis, Tacky eres, and Merganetta indicate currently recognized or tentatively proposed polytypy. Order Anseriformes (Wagler, 1831). Suborder Anseres Wagler, 1831. Family Anatidae Vigors, 1825. Subfamily Stictonettinae Welters, 1976. Genus Stictonetta Reichenbach, 1853. — Freckled Duck. Subfamily Tadorninae Reichenbach, “1850.” — Shelducks and allies. Tribe Merganettini Delacour and Mayr, 1945. Genus Hymenolaimus Gray, 1843. — Blue Duck. Genus Tachyeres Owen, 1875. — Steamer-ducks (see Livezey and Humphrey, 1992). Genus Merganetta Gould, 1842. — Torrent-ducks. Tribe Plectropterini, new taxon. Genus Plectropterus Stephens, 1824. — Spur-winged Goose. Genus Sarkidiornis Eyton, 1838. — Comb-ducks. Tribe Tadornini Delacour and Mayr, 1945 (see Livezey, in press a). Subtribe Chloephageae Boetticher, 1942. Genus Cyanochen Bonaparte, 1856. Genus Alopochen Stejneger, 1885. Genus Neochen Oberholser, 1918. Genus Chloephaga Eyton, 1838. Subtribe Tadorneae Boetticher, 1942. Genus Tadorna Oken, 1817. Subfamily Anatinae Swainson, 1837, sedis mutabilis. — Dabbling ducks and allies. Tribe Malacorhynchini, new taxon. Genus Malacorhynchus Swainson, 1831. — Pink-eared Duck. Genus Salvadorina Rothschild and Hartert, 1894. — Salvadori’s Duck. Tribe Anatini Delacour and Mayr, 1945; incertae sedis (see Livezey, 1991). Subtribe Anateae Boetticher, 1942. Subtribe Cairineae Boetticher, 1942. Subtribe Nettapodeae Livezey, 1991. Tribe Aythyini Delacour and Mayr, 1945 (see Livezey, in press c). Tribe Mergini Delacour and Mayr, 1945 (see Livezey, 1995Z?). Tribe Oxyurini Delacour and Mayr, 1945 (see Livezey, 1995c). INSTRUCTIONS FOR AUTHORS ANNALS f F CARNEGIE MUSEUM consist of con- tributions toll rlie earth sciences, life sciences and anthro- pology, in ?0! py 46 picas format (127 by 195 mm or 5 by 7% inche.};^’. Submit all manuscripts to the Office of Scientific Publications. 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D., W. Granger, and G. G. Simpson. 1928. Paleocene multituberculates from Mongolia. American Museum of Natural History Novitates, no. 331:1-4. 4) Chapter in an edited volume: Rausch, R. L. 1963. A review of the distribution of Holarctic mammals. Pp. 29^3, in Pa- cific Basin Biography (J. L. Gressitt, ed.). Bishop Museum Press, Honolulu, Ha- waii. 5) Unpublished dissertation: Smith, J. P. 1976. Review of Eocene mammals. Unpublished Ph.D. dissert.. University of California, Berkeley. 6) Book: White, M. J. D. 1961. The Chromosomes. Me- thuen and Co., Ltd., London. 7) Journal articles with usual volume and issue number: Anderson, W. I. 1969. Lower Mississippian con- odonts from northern Iowa. Journal of Paleontology, 43(4):9 16-928. Illustrations. All illustrations will be called figures, and are to be numbered in Arabic numerals. Three sets of illustrations are required, one (original artwork) for reproduction, two for reviewers. 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ISSN 0097-4463 Sdy "Xnnals 0/ CARNEGIE MUSEUM THE CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213 VOLUME 65 16 MAY 1996 NUMBER 2 CONTENTS ARTICLES Observations on the mammals of Tucam^ Province, Argentina. 1. Systematics, distribution, and ecology of the Didelphimorphia, Xenarthra, Chiroptera, Primates, Carnivora, Perissodactyla, Artiodactyla, and Lagomorpha Michael A. Mares, Ruben M. Barquez, Janet K. Braun, and Ricardo A. Ojeda 89 Biology and immature stages of snail-killing flies belonging to the genus Tetanocera (Insecta: Diptera: Sciomyzidae). 11. Life histories of pred- ators of snails of the family Succineidae B. A. Foote 153 A new genus of beaver (Mammalia: Castoridae: Rodentia) from the Arika- reean (Oligocene) of Montana and its bearing on castorid phytogeny William W. Korth 167 REVIEW Australian Beetles (J. F. Lawrence and E. B. Britton) .. Darren A. Pollock 181 Editors, ANNALS, BULLETIN and SPECIAL PUBLICATIONS: J. L. Carter B. C. Livezey D. R. Watters M. A. Schmidt, ELS, Assistant Editor Manuscripts, subscriptions, orders for individual numbers, and changes of address should be sent to: Office of Scientific Publications Carnegie Museum of Natural History 4400 Forbes Avenue Pittsburgh, PA 15213-4080 Phone (412) 622-3287 Fax (412) 622-8837 E-mail: schmidtm@clp2.clpgh.org ANNALS OF CARNEGIE MUSEUM is published quarterly by The Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213-4080, by the authority of the Board of Trustees of Carnegie Institute. © 1996 Carnegie Institute. THE CARNEGIE MUSEUM OF NATURAL HISTORY @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 2, Pp. 89-152 16 May 1996 OBSERVATIONS ON THE MAMMALS OE TUCUMAn PROVINCE, ARGENTINA. L SYSTEMATICS, DISTRIBUTION, AND ECOLOGY OF THE DIDELPHIMORPHIA, XENARTHRA, CHIROPTERA, PRIMATES, CARNIVORA, PERIS SOD ACT YL A, ARTIODACTYLA, AND LAGOMORPHA Michael A. Mares*’ ^ Research Associate, Section of Mammals Ruben M. BArquez^ Visiting Museum Specialist, Section of Mammals Janet K. Braun* Ricardo A. Ojeda"* Visiting Museum Specialist, Section of Mammals Abstract The systematics, distribution, and ecology of the Didelphimorphia, Xenarthra, Chiroptera, Primates, Carnivora, Perissodactyla, Artiodactyla, and Lagomorpha of Tucuman Province, Argentina, are sum- marized. Sixty-eight species belonging to 16 families are considered; two species are of probable occurrence (Tayassu pecari and Myrmecophaga tridactyla), and another species occurred in the prov- ince within the memories of locals living in the area (Cebus apella). A checklist also contains infor- mation on occurrence and status. Species accounts provide data on taxonomy, specimens examined and additional records, standard external and cranial morphological measurements, reproduction, molt- ing, habitat preferences, general natural history, and other information. Geography, climate, and veg- etation of the province are summarized. Mammal extirpations, potential threats to species, and mam- malian conservation are discussed. Resumen La sistematica, distribucion, y ecologia de los Didelphimorphia, Xenarthra, Chiroptera, Primates, Carnivora, Perissodactyla, Artiodactyla, y Lagomorpha de la provincia de Tucuman son resumidas. Sesenta y ocho especies perteneciendo a 16 familias son consideradas; tambien se incluye dos especies que probablemente ocurren en la provincia {Tayassu pecari y Myrmecophaga tridactyla) y otra que ocurria alK segun los relatos de habitantes de la zona. Una lista de especies conteniendo informacion sobre ocurriencia y estatus esta incluida. La historia natural de las especies incluye informacion sobre taxonomia, reproduccion, medidas, muda, y habitat. Tambien se comenta sobre la geografia, clima, y vegetacion de la provincia. Aspectos importantes de la conservacion de varias especies son mencion- adas. Introduction The small province of Tucuman is located in northwestern Argentina (Fig. 1). Its small size notwithstanding (22,524 km^, about the size of New Hampshire), ’ Oklahoma Museum of Natural History, University of Oklahoma, Norman, Oklahoma 73019. ^ Instituto Miguel Lillo, Universidad Nacional de Tucuman, 4400 Tucuman, Argentina. ^ Department of Zoology, University of Oklahoma, Norman, Oklahoma 73019. ^ Instituto Argentine de Investigaciones de las Zonas Aridas (lADIZA), C. C. 507, 5500 Mendoza, Argentina. Submitted 8 February 1995. 89 90 Annals of Carnegie Museum VOL. 65 Fig. 1 . — Map showing the location of Tucuman Province in Argentina and the major mountain chains and rivers. Tucuman is both exceptionally rich in the diversity of habitats that it supports, as well as in its vertebrate fauna. Indeed, with more than 100 species of mammals known to occur in the province, and many yet to be found, its richness of mammal species would suggest an area of much greater geographic extent. For example, the state of New Mexico, a large, topographically diverse area, supports 149 species of native mammals (Findley, 1987), a number that is somewhat higher than the number of species that will ultimately be found to occur in Tucuman. Yet New Mexico is 14 times larger than Tucuman. Clearly, Tucuman’s rich floristic diversity, great topographic relief, varied pat- terns of precipitation, and location at a conjunction of major macrohabitats (Ojeda and Mares, 1989) just slightly south of the Tropic of Capricorn, provide a mixture 1996 Mares et al. — TucumAn Mammals 91 of factors that has led to the province’s high richness of mammal species. More= over, because one of Argentina’s oldest cities is in the province (San Miguel de Tucuman — founded in 1565 and the city where Argentine independence from Spain was established in 1816), and because the city is an important agricultural and educational center, especially in the biological sciences, there have been many investigations of the province’s biota. These have included extensive studies on the flora of the province (e.g., Meyer, 1963; Digilio and Legname, 1966; Morello and Adamoli, 1968), as well as a number of reports on Tucuman’s mammals (e.g., Olrog, 1958, 1959, 1976; Olrog and Lucero, 1981; Lucero, 1983; Barquez et ah, 1991). Nevertheless, our recent field work surveying the mammals of the province continues to discover species new to the province and taxa (species and genera) new to science, indicating that the province cannot yet be considered to be well studied from the viewpoint of its mammals. Although a popular guide to the mammals of Tucuman Province was recently published (B^quez et al., 1991), neither it, nor the earlier works of various au- thors, provided detailed information on the distribution, systematics, or other tech- nical aspects of the biology of the province’s mammals. In this report, we provide data on the nonrodent fauna of Tucuman. (We will deal with the province’s com- plex rodent fauna in a subsequent paper.) We also review the physical setting of the province, its floral richness, its climate, and its history, for all of these have influenced the number of species of mammals that today occurs in the province. Finally, we place our observations in a conservation perspective, so that the status and importance of Tucuman’s mammals can be better appreciated and understood. Geography Tucuman is the smallest of Argentina’s provinces. Within its borders are a diversity of habitats that extend from the Chacoan lowlands in the east to the complex vegetation of the high mountains in the west. Tucuman contains a great deal of topographic diversity, ranging from low hills and small mountains in the east and northeast, to Andean massifs in the west. There are three major oro- graphic systems in Tucuman (Fig. 1). In the western parts of the province, Las Cumbres Calchaquies and Las Sierras del Aconquija extend from the northern to the southern borders. The Cumbres Calchaquies, the southernmost extension of the eastern Andean Cordillera, reach elevations of approximately 4500 m; in ad- dition to high, snow-covered peaks, the high mountains form many valleys and lakes. Numerous valleys having economic importance as ranchlands, farmlands, and tourist centers are found in the western highlands. One montane valley that is especially important is the Valle de Tafi, which is a rich bunchgrass prairie that crosses the Abra de Infiemillo at 3042 m and extends northward to the arid Valle de Santa Marfa. The depression formed by these two valleys separates the Cum- bres Calchaquies from the Sierras del Aconquija. The latter are composed of many mountains, among them Cerro Munoz, the Cerros de las Animas, the Nevados del Aconquija, and the Sierras de Santa Ana. Near the center of the province, just to the west of the capital city of San Miguel de Tucuman, is a series of low mountain formations known as the Serranfas de San Javier, mountains covered with a verdant subtropical forest. The Sierras de Burruyacu are a sub-Andean chain found in northeastern Tu- cuman. The easternmost mountains of the province include the Sierra de Medina, Cumbre del Nogalito, Sierra de la Ramada, and Sierra del Campo. These ranges 92 Annals of Carnegie Museum VOL. 65 Fig. 2. — Three east-west topographic profiles through the province of Tucuman showing major geo- graphic features taken along: A, latitude 26°25'; B, latitude 26°50'; and C, latitude 27°20'. Height is in meters above sea level. Population centers are designated by black rectangles. are generally considered the extreme southern limit of the sub- Andean mountains, which are most extensive in Salta and Jujuy provinces to the north (Anonymous, 1981). West of the Sierras del Aconquija and forming the extreme northwestern bound- ary of the province is a fourth mountain system, composed of the eastern faces of the Sierras de Quilmes that border the Valle de Santa Maria on the west, with the Cumbres Calchaquies bordering the valley on the east. The southern edges of these ranges are located in adjacent Catamarca Province. Indeed, the Cerro del Bolson in southern Tucuman along the Catamarca border is the highest point in the province at 5550 m. The great elevational variation in the province is a major factor contributing to its wide array of habitats. An indication of the topographic diversity of the prov- ince is given in Figure 2, which illustrates three topographic profiles across Tu- cuman at approximately 26°30', 26°52', and 27°16’ south latitude. The low, flat plain in the eastern part of the province corresponds to the “great Tucuman plain”; the gradual decline in elevation of the mountain ranges as one proceeds southward and eastward is also depicted. Figure 1 shows the major mountain chains and rivers of Tucuman. Climate Climatically, the province is at the limits of both the subtropical and the tem- perate regions. Thus, the area does not experience four distinct seasons, but rather has two well-defined ones. Summer months are hot and humid and winter months are cool and dry. Additionally, its pronounced elevational relief means that major 1996 Mares et al. — Tucuman Mammals 93 climatic zones will appear at different elevations in the mountains. Three basic climate types are distinguishable within Tucuman: tropical with a dry season in the northern and eastern portions, tropical montane in the central low mountains, and subtropical arid in the high mountains and western lowland valleys. Temperate climates characterize the higher mountainous regions of the prov- ince, but most of the province is influenced by the tropical montane climate. This is due to the north-south orientation of the mountain ranges that interrupt the westward movement of winds carrying moisture from the Atlantic Ocean. The humidity originating from the southern Atlantic air mass is retained by the trans- verse mountains, where orographic rain falls primarily on the eastern slopes. This produces a thick accumulation of clouds, generally between 2000-2500 m ele- vation in the mountains of central and west-central Tucuman, which may produce large amounts of rain. As might be expected, this rain results in rich floristic diversity along the eastern slopes and at mid elevations on the mountains. As the air mass continues to move westward, it becomes progressively more arid, until the drier climates of the western portions of the province develop. The subtropical climatic region, which has a distinct dry season, is restricted to a narrow band located in the eastern part of the province and a second narrow band in the arid mountains and lowland valleys that characterize western Tucuman. Most rivers in Tucuman descend from the eastern slopes of the mountains and unite in the valleys and ravines. The hydrology of the province is defined by the Rio Sail River basin (north-central Tucuman), the Rio Santa Maria (northwestern Tucuman), the Rio Tajamar or Cajon (northeastern Tucuman), and several mon- tane rivers in the south (Fig. 1). Rainfall in the province is greatest along the central mountain chains, reaching 3000 mm annually in the forests on the east-facing montane slopes just west of the capital city of San Miguel in the Serranias de San Javier. Both in far eastern Tucuman and in the far western parts of the province rainfall decreases markedly, with dry thorn scrub resulting in the east and desert scrublands in the west. Vegetation Tucuman supports a complex variety of major phytogeographic areas due to the province’s diverse topography, climate, and hydrology. The general pattern of macrohabitats is shown in Figure 3. For a more detailed description of the plants comprising these various vegetational communities, the reader should consult Ca- brera (1976), whose findings are summarized here. Three Neotropical dominions (each containing a number of major macrohabi- tats) are found in Tucuman. The Amazonian Dominion is represented by the Yungas Phytogeographic Province, the Chaco Dominion is represented by the Chaco, Prepuna, and Monte Phytogeographic provinces, and the Andean-Pata- gonian Dominion is represented by the High Andean and Puna Phytogeographic provinces. We describe these briefly. Yungas Phyto geo graphic Province. — The eastern plain of Tucuman originally supported a transitional forest with dominants that included the trees tipa and pacara (Tipuana tipu and Enterolobium contortisiliquum). According to Meyer and Weyrauch (1966), the plain also supported native cedars, i.e., cedro iCedrela lilloi), lapachos (Tabebuia avellanedae), and cebiles (Piptadenea macrocarpa). Since this forest primarily occurred in areas that have been subjected to extensive human habitation over more than four centuries, it has largely disappeared, being 94 Annals of Carnegie Museum VOL. 65 Fig. 3. — Some of the principle macrohabitats of Tucuman Province (after Cabrera, 1976; Vervoorst, 1982). replaced by orchards and plantations of sugar cane and tobacco (Cabrera, 1976; see also Roig, 1991). The eastern montane slopes of the central mountains support humid montane forest (Vervoorst, 1982), which is frequently divided into three forest types on the basis of elevation. The basal forest (or laurel forest) is found between 400™ 800 m elevation and includes laurel {Phoebe porphyria), palo amarillo {Termin- alia triflora), tarco (Jacaranda mimosifolia), tabaquillo {Solarium verbascifolium), tipa {Tipuana tipu), lapacho {Tabebuia avellanedae), nogal {Juglans australis), cedro {Cedrela lilloi), and San Antonio {Rapanea laetevirens). A forest of mir= taceous trees lies between 800-1400 m. This forest is dominated by horco molle {Blepharocalyx gigantea), and guil {Eugenia pseudomato), as well as several dom^ inant trees in other families, roblina {Ilex argentina), Tucuman laurel {Phoebe porphyria), nogal {Juglans australis), cedro {Cedrela lilloi), and some elements from the lower-elevation forest. An alder forest {Alnus jorullensis) is found at the 1996 Mares et al. — Tucuman Mammals 95 Fig. 4. — The montane subtropical forest of central Tucuman. (Photograph by M. A. Mares.) highest elevations (from 1400-2100 m) and occasionally intermixes with pine {Podocarpus parlatorei) forests (Meyer, 1963). At the higher elevations, in areas where soil conditions or microclimate lead to drier conditions, there is an increase in grasses, especially bunch grasses such as Festuca, Stipa, Chloris, and Calamagrostis. Quehoales “pines” (Polylepis aus- tralis) occur in ravines lying between 1900-2300 m, although in some regions they are found up to 3000 m elevation. The two lower levels are characterized by largely Neotropical plant species, but the upper elevation forest exhibits a number of Holarctic elements, including Alnus, Juglans, Anemone, Ranunculus, Thalictrum, and Geranium. These species extended their distributions southward along the Andean Cordillera during earlier geologic periods. The montane forest, excluding the monospecific stands of alders and conifers, is normally a dense forest in which various strata are differentiated according to light requirements. Trees of the upper stratum reach approximately 30 m in height and include such species as laurel {Phoebe porphyria), horco molle (Blepharo- calyx gigantea), cedars (Cedrela), horco cebil {Piptadenia excelsa), and nogal {Juglans australis). The second stratum is formed by trees that do not exceed 20 m in height — cochucho {Fagara coco), San Antonio {Rapanea laetevirens), ramo {Cupania vernalis), chalchal {Allophyllus edulis), and palo luz {Prunus tucuma- nensis). The shrub stratum is characterized by ramo tucumano {Boehmeria cau- data), hediondilla {Solanum lorentzi), Celtis triflora. Piper tucumanus, Urera bac- cifera, Miconia ioneura, Cestrum lorentzianum, and Psychotria carthagenensis. The lowest stratum is dominated by helecho {Pteris deflexa) and the herbaceous stratum by Oplismenus hirtellus, Panicum cf. demissum, and Axonopus compres- sus. Throughout this forest, vines, lianas, and epiphytes are abundant, especially on the laurels and tipas; mosses, lichens, and ferns are abundant on the ground. Forest canopy vegetation is shown in Figure 4. Chaco Phyto geo graphic Province. — The Chaco Phytogeographic Province is 96 Annals of Carnegie Museum VOL. 65 Fig. 5. — The montane Chacoan forest of northern Tucuman Province at Las Juntas, 22 km W Cho- romoro. (Photograph by J. K. Braun.) found on the large extension of the flat plain of eastern Tucuman where the Chaco proper reaches its western limit. The Chaco, a xerophytic forest with floristic affinities to the monte desert and the high elevation prepuna, is an extensive thorn scrub characterized by Schinopsis lorentzii and numerous thorny trees, such as species in the genera Cercidiuniy Acacia, Prosopis, and Geojfroea. Chaco serrano, or montane Chaco, characterizes the lower slopes of the drier eastern mountain ranges and the western slopes of the mountain ranges located to the east and northeast of the Sierras de San Javier. In this area, the Chaco is characterized by quebracho montano (Schinopsis haenkeana). In general, the Chaco has been heavily exploited throughout the province, resulting in invasions of vinal (Pro- sopis ruscifolia) from the northeast and jarilla (Larrea spp.) from the south. The Chaco of Tucuman, with annual precipitation ranging from 500-800 mm, is much drier than the eastern Chaco of north-central and eastern Argentina (Fig. 5). A major Chacoan plant community in Tucuman is the quebrachal community, composed of trees of quebracho Colorado (Schinopsis lorentzii) and quebracho bianco (Aspidosperma quebracho-bianco). Other important trees include itm (Pro- sopis kuntzei), algarrobo negro (Prosopis nigricans), chanar (Geojfroea decorti- cans), mistol (Zizyphus mistol), quayacan (Caesalpinia paraguariensis), and som- bra de toro (Jodina rombifolia). Cardones (Trichocereus and Cereus), quimil (Opuntia quimilo), and other cacti and epiphytic bromeliads are also present. Monte Phytogeographic Province. — In Tucuman, the monte is limited to areas west of the Cumbres de Calchaqufes. Throughout this region rain is scarce (never more than 200 mm annually) and falls primarily between October and March (summer), peaking in January. Average annual rainfall (based on 47 years of records) at Santa Maria, Catamarca Province, immediately south of the Tucuman border in the monte desert, is 181.7 mm (Morello, 1958). In Tucuman Province proper, average annual rainfall does not exceed 160 mm at Colalao del Valle and 1996 Mares et al. — Tucuman Mammals 97 Fig. 6. ““The monte desert of northwestern Tucuman Province at the Quilmes mins. (Photograph by J. K. Braun.) Amaicha del Valle (based on eight years of data). The monte is characterized by jarilla, or creosotebush {Larrea divaricata and Larrea cuneifolid), and various trees and shmbs {Prosopis, Tricomaria, Mimosa, Bulnesia, Cassia) (Fig. 6). In^ termontane forests formed by Prosopis flexuosa, Geojfroea decorticans, Acacia visco, and Atamisquea emarginata extend along temporary rivers or along the borders of the dry western valleys (Vervoorst, 1982). Colum.nar cacti {Trichcer- eus) are also common. Prepuna Phyto geo graphic Province. — In the provinces of Tucuman, Catamar- ca, and La Rioja, this phytogeographic province is generally found at elevations between 200'0=-3000 m and is located either between the Chaco and Puna Phy- togeographic provinces, or between the Monte and Puna Phytogeographic prov- inces (Cabrera, 1976). In general, the presence or absence of the prepuna depends upon the altitude, arrangement, and oriicntation of the mountain ranges. The cli- mate is dry and rainfall occurs in the summer. It is strongly related to the Monte Province, but differs fundamentally by the absence of Larrea and by the abun- dance of many species of columnar cacti (genus Trichocereus) and terrestrial, ring- or cushion-shaped ground bromeliads {Dyckia, Deuterocohnia, AbromeT tielia). Prepuna plant communities (Fig. 7) are able to grow on rocky mountain slopes with both eastern and western exposures (Vervoorst, 1982). Puna and High Andean Phytogeographic Provinces.— In Tucuman, the Andean Patagonian Domain is represented by the Puna and High Andean Phytogeographic provinces (Fig. 7). The latter is found in high mountain regions from the peaks down to the tree line. In Tucuman it occurs on the Aconquija range and the Nevados del Cajon. Normally, the soils in the high Andean Phytogeographic Province are rocky, stony, or sandy. Bogs may occur in moist areas. The montane climate is cold and dry, and the winds strong. There are only two endemic families of plants in this province, but there are a number of endemic genera. The dominant 98 Annals of Carnegie Museum VOL. 65 Fig. 7. — The prepuna northwest of El Infiernillo above Tafi del Valle. Puna vegetation can be seen in the background on the higher slopes. (Photograph by J. K. Braun.) vegetation is grassland steppes or steppes of low-growing plants having large roots (e.g., Adesmia, Baccharis, Oxalis, Senecio). The puna occurs on high mountains and plateaus, typically between 3400-4500 m. The soils are poor in organic material and are frequently rocky or sandy. The dominant vegetation is the shrub steppe, although herbaceous steppes and bog areas are not uncommon. The puna is related to the Patagonian region and many dominant plant genera are shared between them, including Junellia, Fabiana, Chuquiraga, Nardophyllum, Adesmia, and Mulinum (Cabrera, 1976). Methods General — Specimens were collected using standard Museum Special snap traps, Victor rat traps, Havahart traps (48 X 15 X 15 cm), gopher traps, leg-hold traps, Sherman live traps (7.6 X 7.6 X 25.4 cm and 12.7 X 12.7 X 38.1 cm), home-made live traps, mist nets, and .22 caliber rifles. All specimens were pre- pared as skins plus skulls or skeletons, or preserved in 10% formalin and alcohol. Beginning in 1990, tissues were collected and frozen in liquid nitrogen and rep- resentative specimens were karyotyped. Data collected included standard external measurements, reproductive condition, and molting pattern. Ecological or natural history data were noted when specimens were collected ancillary to ecological or physiological research. Specimens collected by us over the past quarter century are deposited in the following museums: Carnegie Museum of Natural History, Pittsburgh; Coleccion Miguel Lillo, Instituto Miguel Lillo, Tucuman; lADIZA, Coleccion de Mamiferos, Mendoza; Oklahoma Museum of Natural History, Uni- versity of Oklahoma, Norman; Texas Cooperative Wildlife Collection, Texas A & M University, College Station (frozen tissues and chromosomal preparations). Taxonomy. — We have chosen to follow, with four exceptions, the most recent 1996 Mares et al, — Tucuman Mammals 99 Fig. 8. — Collecting localities for mammals listed in the text. Numbers correspond to those given in the Gazetteer. edition of “Mammal Species of the World: A Taxonomic and Geographic Ref- erence” (Wilson and Reeder, 1993) for the taxonomic arrangement of families, subfamilies, genera, and species. Sturnim oporaphilum is used instead of 5*. bo- gotensis. Based on studies using allozyme and mitochondrial DNA data (Baker et aL, 1988; Morales and Bickham, 1995), we recognize Lasiurus blossevillU for the South American red bats. Eumops patagonicus is recognized as a distinct species; this species is included in Eumops bonariensis in Wilson and Reeder (1993). We follow Garcia-Perea (1994) in the recognition of Lynchailurus as a distinct genus containing three species; L. pajeros is found in Tucuman Province. Collecting Localities.^ — The principal localities for specimens cited in the text in Tucuman Province are shown in Figure 8. The Gazetteer lists the localities and 100 Annals of Carnegie Museum VOL. 65 coordinates of south latitude and west longitude in degrees and minutes for all specimens cited in the text that could be located. Localities are listed alphabetic cally and numerically; departments and numbers corresponding to numbered lo- calities in Figure 8 are given in parentheses. Coordinates were obtained from Gazetteer No. 103 (Argentina) of the United States Board of Geographic Names prepared by the Office of Geography, Department of Interior, Washington, D. C. (Anonymous, 1968), used in conjunction with various maps. Localities have been kept as they appeared on the original labels, except for obvious misspellings. Maps of Tucuman published by the Automovil Club Argentine (1974, 1977, 1982, 1986), the series of 1:500,000 provincial maps published the EJercito Argentino, lestituto Geografico Militar in 1966 (number 2966) and 1972 (number 2766), and the 1:250,000 maps of the province published by the EJercito Argentino, Institute GeogrMco Militar in 1986 (number 2766cIV), 1987 (number 2766^111), and 1988 (number 2766-1) were used to plot localities. Aeronautical maps TPC Q-27A (1: 500,000; 1988) and Q-27 (1:1,000,000; 1973), prepared and published by the Defense Mapping Agency Aerospace Center, were also used to plot localities. Checklist of the Mammals of TucumAn Province A list of the orders, families, genera, and species of mammals known or ex- pected to occur in Tucuman Province, Argentina, is given in Table 1. The des- ignations of relative abundance are largely subjective and are based on our long experience with the species of the province. We present them only to guide bi- ologists as to which species might be of special concern for conservation and which might be relatively easy to study in the province. In some cases, we have actual mark-recapture data, or collecting data, to estimate commonness or rarity. In other cases, we have formed impressions as to whether or not a species is common or rare based on experiences in capturing or observing the species. Species Accounts In the following sections we present the individual species accounts for those mammal species known to occur in Tucuman Province. Accounts are arranged according to order, family, and subfamily (where appropriate). Genera and species are arranged alphabetically within families or subfamilies. For each scientific name, the authority and literature citation for the first recognized description are given. In the measurements presented for each species, mean and range for adult males and females are listed separately. For two or fewer specimens, only the measurements are listed. Measurements are given in millimeters. Specimens from the following museum collections (acronyms given in paren- theses) were examined: The Natural History Museum, London, England (BMNH); Carnegie Museum of Natural History, Pittsburgh, Pennsylvania (CM); Coleccion Lillo, Institute Miguel Lillo, Tucuman, Argentina (CML); Coleccion de Mamf- feros, Centro Regional de Investigaciones Cientificas y Tecnicas— ™CRICYT, Men- doza, Argentina (lADIZA); Facultad Ciencias Exactas y Naturales, Mamiferos, Universidad Nacional de Buenos Aires, Buenos Aires, Argentina (FCM); Museo Argentino de Ciencias Naturales “Bernardino Rivadavia,” Buenos Aires, Argen- tina (MACN); Oklahoma Museum of Natural History, University of Oklahoma, Norman, Oklahoma (OMNH); personal collection of Ruben Barquez, Tucuman, Argentina (RMB); The Museum, Texas Tech University, Lubbock, Texas (TTU). 1996 Mares et al. — TucumAn Mammals 101 Order Didelphimorphia Family Didelphidae Subfamily Didelphinae Didelphis albiventris Lund Didelphis albiventris Lund, Konigelige Danske Videnskabernes Selskabs Afhandlinger, Kjobenhavn, p. 20, 1840. Specimens Examined. — (26) Aconquija, Concepcion, 5 (BMNH); Cerro del Campo, 800 m, 1 (BMNH); El Cadillal Dike, 25 km NW San Miguel de Tucuman, 2 (CM); El Cajon, 1 (CML); 4 km W of junction 338 and road to Horco Molle, on road to San Javier, 2,750 ft., 1 (OMNH); Biological Reserve at Horco Molle, 2,400 ft., 3 (1 CML, 1 lADIZA, 1 OMNH); Los Romanos, 1 (CML); Mala Mala, 1 (BMNH); Escuela Normal, Monteros, 1 (MACN); Piedra Tendida, 12 km WNW Burruyacu along Rfo Cajon, 2,500 ft., 2(1 lADIZA, 1 OMNH); San Fernando, 1 (CML); San Pedro de Colalao, 3 (CML); at km marker 42, on highway 364, south of San Pedro de Colalao, 4,700 ft., 3(1 CML, 1 lADIZA, 1 OMNH); Tucuman, no specific locality, 1 (BMNH). Additional Records. — Aguas Chiquitas (Cajal, 1976); Pozo Hondo (R. M. Barquez, personal obser- vation); Dique Escaba (J. K. Braun, personal observation); Santa Rosa de Leales (Massoia and Fornes, 1965); Taff del Valle (J. K. Braun, personal observation). Measurements. — External measurements (eight males, ten females, unless noted otherwise): total length, 518.5 (292.0=723.0), 588.2 (370.0=770.0); tail, 237.4 (84.0-337.0), 294.6 (175.0-390.0); hindfoot, 45.0 (32.0-54.0), 46.0 (30.0-60.0); ear, 49.7 (40.0-55.0), 50.2 (39.0-56.0); weight (four males, four females), 331.0 (210.0-508.0), 354.8 (144.0-910.0). Cranial measurements (two males, four fe- males, unless noted otherwise): greatest length of skull, 77.5, 94.0, 77.4 (52.0- 93.9); condylobasal length, 76.0, 91.0, 75.1 (45.7-91.8); least interorbital length, 10.0, 10.7, 10.2 (9.8-10.6); zygomatic breadth, 38.7, 53.6, 40.8 (28.0-48.3); breadth of braincase (two males, three females), 21.0, 27.4, 24.6 (23.7-25.8); length of maxillary toothrow, 21.0, 36.1, 27.7 (18.9-35.6); palatal length (one male, four females), 55.1, 45.0 (30.2-54.2); length of mandibular toothrow (one male, two females), 37.5, 32.3, 37.9; greatest length of mandible (one male, two females), 76.6, 60.0, 73.6. Reproduction.—See Table 2. Molt. — A subadult female and male were molting in January. Habitat. — This species is widely distributed throughout Argentina from north of 41°S latitude in Rfo Negro Province northward. Specimens were collected in transitional forest, areas bordering transitional forest, and in pine forest. Individ- uals were captured in Havahart traps, Sherman live traps, and in Victor rat traps. Both individuals from Piedra Tendida were captured in Havahart traps baited with either lunchmeat or sardines (M. L. Campbell, personal observation). One juvenile was caught in a trap set in a tree. Lucero (1983) noted that this species inhabits a variety of habitats including cities and urban areas. It is found at altitudes up to 2500 m (Lucero, 1983). The species is seldom found in arid or semiarid areas, preferring mesic agricultural zones, forests, and other moist habitats. The opossum from Taff del Valle was seen crossing the road (Highway 307) north of Taff del Valle at night (J. K. Braun, personal observation). Although we did not observe any individuals, locals at Dique Escaba indicated that D. albiventris was common in the area (J. K. Braun, personal observation). Remarks.— is some question regarding the specimens in the Natural His- tory Museum, London, collected by Shipton from Concepcion, Aconquija. The location on the skin tag is listed as Concepcion, but the locations on the skull tags have been changed to Aconquija. Cajal (1976) reported on the population ecology of this species in Tucuman Province. 102 Annals of Carnegie Museum VOL. 65 Table 1. — Checklist of the mammals ofTucumdn Province. The occurrence of each species is indicated as being: (*) collected, seen, or museum specimens examined by us; (1) reported in the literature; (p) of probable occurrence due to the presence of the species in adjacent provinces in similar habitats. In addition, we note whether, in the province of Tucumdn, the species is common (c), uncommon (u), rare (r), in danger of extirpation (e), or probably extirpated from the province (x). Taxon Occurrence Status Order Didelphimorphia Family Didelphidae Subfamily Didelphinae Didelphis albiventris *, 1 C Lutreolina crassicaudata * u Monodelphis dimidiata * r Thylamys elegans 1 c Thylamys pallidior *, 1 u Order Xenarthra Family Dasypodidae Subfamily Dasypodinae Cabassous chacoensis r Chaetophractus vellerosus *, 1 c Euphractus sexcinctus * u Priodontes maximus P r?, X? Tolypeutes matacus 1 e Family Myrmecophagidae Myrmecophaga tridactyla X? Tamandua tetradactyla * u Order Chiroptera Family Phyllostomidae Subfamily Phyllostominae Chrotopterus auritus * r Subfamily Stenodermatinae Artibeus planirostris * c Sturnira erythromos * u Sturnira lilium * c Sturnira oporaphilum * r Subfamily Desmodontinae Desmodus rotundus * c Family Vespertilionidae Subfamily Vespertilioninae Eptesicus diminutus * u Eptesicus furinalis * u Histiotus macrotus * u Lasiurus blossevillii 1 c Lasiurus cinereus * c Lasiurus ega 1 c Myotis albescens * c Myotis keaysi * c Myotis levis * c Myotis nigricans *, 1 c Family Molossidae Eumops bonariensis * c Eumops dabbenei * u Eumops glaucinus * c Eumops patagonicus * c Eumops perotis *, 1 c Molossops temminckii *, 1 c Molossus molossus * c Nyctinomops macrotis * c Promops nasutus =1= c Tadarida brasiliensis *, 1 c 1996 Mares et al. — TucumAn Mammals 103 Table 1. — Continued. Taxon Occurrence Status Order Primates Family Cebidae Subfamily Cebinae Cebus apella X? Order Carnivora Family Canidae Cerdocyon thous c Pseudalopex culpaeus u Pseudalopex griseus u Pseudalopex gymnocercus * c Family Felidae Subfamily Felinae Herpailurus yaguarondi * u Leopardus pardalis *, 1 u Lynchailurus pajeros * r Oncifelis geojfroyi 1 c Oreailurus jacobita I r Puma concolor 1 c Subfamily Paotherieae Panthera onca X Family Mustelidae Subfamily Lutrinae Lontra longicaudis sft e Subfamily Mephitinae Conepatus chinga c Subfamily Mustelinae Eira barbara * u Galictis cuja * c Lyncodon patagonicus 1 u Family Procyonidae Subfamily Procyoninae Nasua nasua u Procyon cancrivorus * u Order Peris sodacty la Family Tapiridae Tapirus terrestris *, 1 p?, X? Order Artiodactyla Family Tayassuidae Catagonus wagneri sfs r Pecari tajacu * c Tayassu pecari 1 p ? Family Camelidae Lama guanicoe 1 u Vicugna vicugna 1 X Family Cervidae Subfamily Odocoileieae Hippocamelus antisensis * u Mazama americana * u Mazama gouazoupira c Ozotoceros bezoarticus 1 X Order Lagomorpha Family Leporidae Sylvilagus brasiliensis u VOL. 65 104 Annals of Carnegie Museum ^ § •S § s ^ « s Q "S Oo 2 1 ll a 2 s g " I ^ h " a ^ s ^ 0 . -a h 1 - ^:§ ^ I ^ -s ao S § ;r 0 a S ^ § “f^ 1 P (N X X CM 00 00 < X s < < m ^ < < < < C^ 00 00 oo o J a“• < < < < 04 ^ H < C/!l H < < »r) CO < < < < < ^ 04 *— I cn U > 04 X s •a I .a |l s s i:i •S I S, Q s s S o I ki e S ^ s ^ pq It S § a 6^ |S I ^ ttj S' a s ? I § brasiliensis (3 X 2) 114 Annals of Carnegie Museum VOL. 65 in transitional forest, primarily in nets near sources of water. Barquez and Ojeda (1992) noted that all Chacoan localities in the province are near Yungas forest. Remarks. — Because individuals of this species have been confused with S. HI- ium and S. oporaphilum, little is known (with confidence) about the biology of this species (but see Barquez, 1987). Specimens have been captured in January (n = 7), May (n = 1), June (n = 1), July {n = 4), September (n = 10), October {n = 6), and November (n = 1). At Horco Molle, this was the only species captured; individuals of Tadarida brasiliensis were captured several days earlier (Mares et ah, 1995). Other species captured at Las Juntas include Artibeus plan- irostris, Histiotus macrotus, Lasiurus blossevUlii, and Tadarida brasiliensis. My- Otis keaysi and Myotis nigricans also were captured at Piedra Tendida, and at San Pedro de Colalao Myotis levis also was captured. Sturnira lilium (E. Geoffroy) Phyllostoma lilium E. Geoffroy, Annales Museum d’Histoire Naturelle, Paris, 15:181, 1810. Specimens Examined. — (57) Agua Colorada, 1 1 (9 MACN, 2 TTU); Casa de Piedra, Rio Los Sosa, 4 (CML); Dique San Ignacio, 1 (CML); El Cadillal, 1 (OMNH); Horco Molle, 15 km W San Miguel de Tucuman, 17 (10 CM, 7 OMNH); Ingenio San Pablo, 1 (MACN); La Cocha, Dique San Ignacio, 1 (OMNH); Los Sarmientos, 1 (CML); Playa Larga, 8 (CML); Rio Los Sosa, Ruta 307, km 19.7, 700 m, 1 (OMNH); Ruta 307, km 19.7, camino a Tafi del Valle, 750 m, 5 (OMNH); Ruta 307, km 23.9, 850 m, 1 (OMNH); Ruta 307, km 24.9, camino a Tafi del Valle, 850 m, 2 (OMNH); San Miguel de Tucuman, 2 (BMNH); San Pedro de Colalao, 1 (TTU). Measurements. — External measurements (eight males, 13 females, unless noted otherwise): total length, 60.5 (57.0-64.5), 61.1 (51.0-70.0); hindfoot, 12.3 (9.2- 14.1), 12.2 (9.3-14.8); ear, 17.6 (16.0-19.5), 17.2 (13.0-19.5); forearm (nine males, 13 females), 44.1 (41.7-46.0), 43.1 (38.7-45.9); weight (eight males, 11 females), 22.8 (20.0-28.0), 21.7 (17.0-25.0). Cranial measurements (eight males, 13 females, unless noted otherwise): greatest length of skull (eight males, 12 females), 23.1 (22.1-23.9), 22.2 (20.1-23.3); condylobasal length (eight males, 12 females), 21.1 (20.3-21.8), 20.2 (18.4-21.0); least interorbital length (nine males, 13 females), 6.1 (5. 8-6. 5), 6.0 (5. 5-6. 8); zygomatic breadth (seven males, nine females), 13.7 (13.0-14.0), 13.3 (12.1-14.3); breadth of braincase, 10.8 (10.2-11.3), 10.6 (9.5-11.2); length of maxillary toothrow (nine males, 11 fe- males), 6.8 (6. 3-7.4), 6.6 (5. 5-7. 3); palatal length (seven males, 13 females), 9.7 (9.4-10.0), 9.2 (8.4-10.0); length of mandibular toothrow, 6.5 (6. 2-6. 9), 6.3 (5.0- 6.7); greatest length of mandible (eight males, 12 females), 14.6 (13.8-15.5), 14.0 (13.0-15.0). Reproduction. — See Table 3. Barquez (1988) reported on the reproductive bi- ology of bats sampled between 12-16 October at Ruta 307, km 23.9. Eor females sampled, 41% were pregnant, 6% were lactating, 41% had open (perforate) va- ginas but without palpable fetuses, and 12% had closed vaginas. Males with abdominal testes comprised 58% of the sample; 42% had scrotal testes. During a later survey (28-30 October), all males (100%) had scrotal testes, 50% of the females had small palpable fetuses, and 50% were near parturition. In mid-Jan- uary, young were flying and there was a decrease in reproductive activity. Samples at this time showed 4% of males with scrotal testes, 43% were adults with ab- dominal testes, and 53% were juveniles with abdominal testes. Adult females were lactating, but 77% of all females were juveniles. Individuals captured at a nearby location in May were not reproductivly active and population densities were lower. Habitat. — In Argentina, S. lilium is distributed in the northwestern provinces 1996 Mares et al. — -Tucuman Mammals 115 of Jujuy, Salta, Tucumae, and Catamarca. It is also found in the northeastern provinces of Chaco, Formosa, Misiones, Entre Rios, and Santa Fe (Barquez, 1987). Lucero (1983) noted that S. lilium is locally common in humid and tram sitional forests. It apparently prefers the low vegetation forest stratum (Barquez, 1987). Few specimens document the presence of this species in the Chaco of Tucuman. Two localities, Agua Colorada and El Cadillal, are gallery forests. Bar- quez and Ojeda (1992) suggested that these records reflect a shift from forests to the Chaco during periods of limited availability of fruit. Remarks.— At Rfo Los Sosa, Rt. 307, km 23.9, this species was captured with Artibeus planirostris, Histiotus macrotus, Lasiurus cinereus, and Tadarida bras- iliensis. Sturnira lilium commonly eats the fruit of the Solanaceae, although at Horco Molle it has been found eating fruit of the introduced moras (mulberry). This species is not common in urban environments, although Barquez (1987) reported that pellets from Barn Owls {Tyto alba) found in buildings in the city of Tucuman contained skulls of young and subadult S. lilium. Owls might prey on S. lilium as they fly through the city or else are foraging in nearby forests. Sturnira oporaphilum (Tschudi) Phyllostoma oporaphilum Tschudi, Untersuchungen iiber die Fauoa Peraana. Therologie. Scheitlio ued Zollikofer, St. Gallen, Switzerland, p. 64, 1844. Specimen Examined.— {!) Casa de Piedra, Ruta 307, km 23.9, 850 m, 1 (OMNH). Additional Record.— A specimen identified as this species was recently captured and released at Piedra Tendida (P. Capllonch, personal communication). Measurements.— External measurements (one male): total length, 58.5; hind- foot, 9.5; ear, 18.5; forearm, 44.2; weight, 21.0. Cranial measurements (one male): greatest length of skull, 22.7; condylobasal length, 20.7; least interorbital length, 6.2; zygomatic breadth, 13.6; breadth of braincase, 10.9; palatal length, 9.0; length of mandibular toothrow, 6.5; greatest length of mandible, 13.5. Reproduction.— Sec Table 3. Habitat.- — -In Argentina, S. oporaphilum is distributed in the transitional forest habitats of Jujuy, Salta, and Tucuman. Remarks. — Barquez (1987) reviewed the biology of this species, which is little known in Argentina. In Wilson and Reeder (1993), this species is listed as S. bogotensis. In the localities where it has been captured, it is sympatric with S. lilium and S. erythromos. Subfamily Desmodontinae Desmodus rotundus (E. Geoffroy) Phyllostoma rotundus E. Geoffroy, Annaies Museum d’Histoire Naturelle, Paris, 15:181, 1810. Specimens Examined ~ — (13) El Cadillal, 1 (CML); Horco Molle, 15 km SW San Miguel de Tucu- man, 1 (CM); Monte Bello, 3 (CML); Piedra Buena, 1 (TTU); Playa Larga, 1 (CML); Rio Los Sosas, Ruta 307, km 23.9, 1 (OMNH); Vipos, Estancia San Pedro, 3 (CML). Measurements.— External measurements (three males, two females, unless not- ed otherwise): total length, 82.9 (80.0-88.0), 85.0, 91.0; hindfoot, 16.7 (16.0- 17.0), 16.0, 18.6; ear, 17.9 (17.0-19.6), 18.0, 20.7; forearm (four males, two females), 62.1 (60.0-64.0), 62.0, 65.2; weight (one male, one female), 43.0, 48,0. Cranial measurements (four males, two females, unless noted otherwise): greatest length of skull, 24.7 (24.2-25.4), 24.7, 25.6; condylobasal length, 22.4 (22.2- 22.6), 22.5, 22.9; least interorbital length, 5.6 (5. 2-6.0), 5.7, 5.4; zygomatic 116 Annals of Carnegie Museum VOL. 65 breadth, 123 (11.9-12.7), 12.5, 11.6; breadth ofbraiecase, 12.1 (11.1-12.6), 12.8, 12.8; length of maxillary toothrow (one female), 3.5; palatal length (three males, one female), 9.0 (8. 9-9. 2), 8.8; greatest length of mandible (one male, one fe- male), 14.2, 16.1. Reproduction.-Sce Table 3, Habitat.— Vampim bats are widely distributed in Argentina as far south as approximately 32°S latitude. In Tucuman, these bats are found in savannas, grass- lands, and open rural areas up to 2000 m (Oirog and Lucero, 1981; Lucero, 1983). ^^marij'.-^Although groups composed of up to 5000 individuals have been reported in other parts of the range, in Tucuman vampire bats form much smaller aggregations. They commonly are found in wells in the countryside. B^quez (1987) reviewed the biology of this species in Argentina. Family Vespertilionidae Subfamily Vespertilioninae Eptesicus diminutus Osgood Eptesicus diminutus Osgood, Field Museum of Natural History, Zoological Series, 10:197, 1915. Specimens Examined. — (4) Aguas Chiquitas, Sierras de Medina, 800 m, 3 (CM); El Cadillal, Es- tacion de Piscicultura, 1 (CM). Measurements. — External measurements (one male, two females): total length, 86.0, 89.0, 91.0; tail, 32.0, 35.0, 32.0; hindfoot, 6.2, 6.8, 7.0; ear, 13.3, 13.5, 13.6; forearm, 32.3, 32.7, 34.0; weight, 4.9, 5.7, 6.3. Cranial measurements (one male, two females, unless noted otherwise): greatest length of skull, 13.1, 13.6, 13.3; condylobasal length, 12.4, 12.6, 12.9; least interorbital length, 3.6, 3.8, 3.8; zy- gomatic breadth, 8.6, 8.9, 8.9; breadth of braincase, 6.5, 6.7, 7.1; length of max- illary toothrow, 4.7, 5.0, 4.9; palatal length, 4.7, 5.3, 5.2; length of mandibular toothrow, 4.2, 4.5, 4.6; greatest length of mandible (two females), 9.9, 9.5. Reproduction. — See Table 3. Habitat. — Eptesicus diminutus is widely distributed in Argentina, although mu- seum records are scarce. In addition to the localities given above for Tucuman, this species is known from one locality each in the provinces of La Pampa, Buenos Aires, Santa Fe, Corrientes, Misiones, Jujuy, and Salta (Barquez, 1987; Barquez and Lougheed, 1990). Remarks. — The taxonomic history of this taxon was discussed by Barquez (1987). Specimens were captured in December (n = 1) and January (n = 3). Little is known of the biology of this species. Eptesicus furinalis (d’Orbigny) Vespertilio furinalis d’Orbigny, Voyage dans I’Amerique Meridionale (le Bresil, la Republique orien- tale de Uruguay, la Republique Argentine, la Patagonie, la Republique du Chile, la Republique de Bolivia, la Republique du Perou) execute pendant das annees 1826, 1827, 1828, 1829, 1830, 1831, 1832, et 1833. Pitois-Levrault, et cie, Paris, Strasbourg, 4:13, 1847. Specimens Examined. — (13) Aguas Chiquitas, Sierras de Medina, 800 m, 4 (CM); Concepcion, 3 (BMNH); El Cadillal, 1 (CML); Las Mesadas, 1 (CML); Las Talas, 1 (CML); Las Talas, 4 km al N de Bella Vista, 1 (OMNH); San Miguel de Tucuman, 2 (1 BMNH, 1 CML). Measurements. — External measurements (one male, four females): total length, 102.0, 104.2 (99.5-111.0); tail, 40.0, 41.6 (37.0-45.0); hindfoot, 7.7, 8.6 (7.1- 10.1); ear, 15.0, 15.6 (14.7-17.3); forearm, 38.8, 39.7 (37.6-41.2); weight, 8.5, 10.2 (7.4-13.0). Cranial measurements (one male, four females): greatest length 1996 Mares et al. — Tucuman Mammals 117 of skull, 15.0, 15.5 (14.8-^16.0); condylobasal length, 14.5, 15.0 (143^153); least interorbital length, 3J, 3.8 (3.7=4. 0); zygomatic breadth, 10.4, 10.8 (9.8-113); breadth of braincase, 7.0, 7.7 (7. 2-8.0); length of maxillary toothrow, 5.6, 5.9 (5. 7=6.0); palatal length, 6.1, 6.5 (6 0=7.4); length of mandibular toothrow, 5.3, 5.8 (5. 5=6. 5); greatest length of mandible), 11.3, 11.9 (11.2=12.2). Habitat. — -Eptesicus furinalis is distributed from 36°S latitude northward, ex- cluding the provinces of San Luis and San Juan (Barquez, 1987; Barquez and Lougheed, 1990). Remarks.- — Eptesicus f findleyi occurs in the montane regions of northwestern Argentina (Williams, 1978). Myers and Wetzel (1983) synonymized this taxon with E. f. furinalis, an arrangement that was followed by Barquez (1987). Little is known about the biology of this species in Argentina. Myers (1977) and Myers and Wetzel (1983) reported on the biology of this species in Paraguay. Histiotus macrotus (Poeppig) Nycticeius macrotus Poeppig, Reise in Chile, Peru und auf dem Amazonenstrome wahrend der Jahre 1827-1832. Friedrich Fleischer, Leipzig, 1:451, 1835. Specimens Examined. — =(29) Aota Mapu, 1 (MACN); El Cadillal, 1 (CML); E! Naranjal, 2 (1 MACN, 1 TTU); El Naranjo, 8 (3 MACN, 5 TTU); El Nogalar, km 43, Ruta 307, 1700m, 4 (CML); Horco Molle, 15 km W San Miguel de Tucuman, 1 (CM); Las Juntas, 22 km W Choromoro on Hwy 312, 3,500 ft., 1 (OMNH); Norco, 2500 m, 2 (BMNH); San Miguel de Tucuman, 4 (2 BMNH, 1 CM, 1 CML); San Pedro de Colalao, 4 (CML); Tafi, 2000 m, 1 (BMNH). Measurements.— ^E-Ki&Tn^l measurements (six males, two females, unless noted otherwise): total length (two males, two females), 107.0, 113.0, 110.0, 109.0; tail (four males, two females), 55.0 (50.0-59.0), 46.0, 49.0; hindfoot (five males, two females), 9.2 (7.0-10.0), 9.8, 8.5; ear, 34.1 (27.0=38.0), 37.1, 34.5; forearm, 46.7 (44.0=48.9), 45.2, 47.7; weight (two males, one female), 9.0, 10.5, 11.0. Cranial measurements (four males, two females, unless noted otherwise): greatest length of skull, 17.5 (16.7=18.1), 17.7, 16.5; condylobasal length, 16.3 (15.7=17.0), 17.0, 17.1; least interorbital length (six males, two females), 4.1 (3. 8-4.6), 4.0, 4.2; zygomatic breadth, 10.6 (10.1=11.2), 10.1, 10.6; breadth of braincase, 8.4 (8.3- 8.4) , 7.7, 8.0; length of maxillary toothrow (five males, two females), 6.0 (5.0- 6.4) , 6.0, 6.0; palatal length (four males, one female), 7.3 (6. 8-7. 9), 7.2; length of mandibular toothrow, 6.2 (5. 6-6. 7), 5.5, 6.4; greatest length of mandible, 12.2 (11.8=12.5), 11.8, 12.1. Reproduction.— EmquGz (1987) reported that specimens captured in May and June were not reproductively active. Habitat. — Histiotus macrotus is found in the forests of the northwestern part of the province and ranges into the arid Monte Phytogeographic Province (Bar- quez, 1987). This species is found from Jujuy and Salta south to Rio Negro (Barquez, 1987; Barquez and Lougheed, 1990). In transitional forest at Las Juntas, a single individual of this species was captured in a net placed over the Rio Choromoro (Mares et aL, 1995). Remarks. — At Las Juntas, this species was captured with Artibeus planirostris, Sturnira erythromos, Lasiurus blossevillii, and Tadarida brasiliensis. Adults were collected in February {n = 5), March {n = 1), July {n = 1), September {n = 1), and November (w = 1). At San Pedro de Colalao, individuals were captured inside a dwelling and were flying over the city (Barquez, 1987). Little is known about the biology of this species in Argentina. 118 Annals of Carnegie Museum VOL. 65 Lasiurus blossevillii (Lesson and Garnot) Vespertilio blossevillii Lesson and Garnot, Bulletin des Sciences Naturelles et de Geologic, 8:95, 1826. Specimens Examined. — (23) Acheral, 1 (CML); Agua Rosada, 1 (CML); Concepcion, 3 (BMNH); Horco Molie, 1 (TTU); Ingeeio Amalia, 1 (CML); Ingenio Santa Lucia, 1 (CML); Instituto Lillo, 1 (CML); Las Juntas, 22 km W Choromoro on Hwy 312, 3,500 ft., 1 (OMNH); Los Vazquez, 445 m, 3 (BMNH); San Miguel de Tucuman, 6 (3 BMNH, 3 CML); San Pedro de Colaiao, 2 (CML); Timbo Nuevo, 1 (CML); Tucuman, 1500 m (no specific locality), 1 (BMNH). Additional Record. — El Colmenar (Romana and Abalos, 1950). Measurements.— measurements (five males, three females, unless not- ed otherwise): total length, 97.4 (92. 5-104. 0), 103.3 (100.0-105.0); tail, 48.5 (45.0-53.0), 49.7 (47.0-52.0); hindfoot, 8.2 (6.5-11.0), 7.8 (7.5-8.0); ear, 9.4 (4.0-13.0), 9.3 (6.0-11.0); forearm (eight males, three females), 38.7 (36.9-40.0), 40.2 (39.0-40.9); weight (one male), 9.0. Cranial measurements (seven males, three females, unless noted otherwise): greatest length of skull, 11.8 (11.2-12.8), 11.8 (11.5-12.1); condylobasal length, 10.5 (9.5-11.3), 11.7 (11.5-11.8); least interorbital length, 4.3 (4. 2-4. 5), 4.4 (4. 3-4. 5); zygomatic breadth (five males), 8.5 (8. 2-8. 9), 9.3 (9. 2-9.4); breadth of braincase (six males, three females), 7.0 (6. 8-7. 2), 7.5 (7.4-7. 7); length of maxillary toothrow (five males, three females), 3.8 (3.7-4.0), 4.2 (4. 1-4.3); palatal length (six males), 3.7 (2.8-4.2); length of mandibular toothrow (four males, three females), 4.3 (4. 2-4.4), 4.8 (4.8-4.9); greatest length of mandible (four males, three females), 8.3 (8. 0-8. 7), 9.2 (9.1- 9.3). Habitat. — Lasiurus blossevillii is distributed throughout northern Argentina, It has not been reported, however, from the provinces of Santa Fe and Santiago del Estero (B^quez, 1987). It appears to be rare in the Chaco. In Tucuman, this bat has been captured in many different habitats. Open areas appear to be preferred, although individuals have been captured flying over rivers in forests. At Las Juntas, the habitat was montane transitional forest. The single red bat collected at this site was taken in a net placed over the Rio Choromoro (Mares et aL, 1995). Remarks.- — At Las Juntas, this species was captured with Artibeus planirostris, Sturnira erythromos, Histiotus macrotus, and Tadarida brasiliensis. Wilson and Reeder (1993) listed this species under L. borealis. We have chosen to follow Baker et al. (1988) and Morales and Bickham (1995) in recognizing the distincL ness of red bats in the western and southern portions of their range. Lasiurus cinereus (Beauvois) Vespertilio cinereus Beauvois, A scientific catalogue of Peale’s Museum. S. H. Smith, Philadelphia, p. 18, 1796. Specimens Examined. — (7) Las Mesadas, 1 (CML); Los Vazquez, 1 (BMNH); Marcos Paz, 1 (CML); Playa Larga, 1 (OMNH); San Miguel de Tucuman, 2 (1 BMNH, 1 CML); Tafi Viejo, 1 (CML). Measurements, — External measurements (one male, two sex unknown): fore- arm, 52.6, 49.0, 52.7. Cranial measurements (one male, one sex unknown, unless noted otherwise): greatest length of skull, 15.7, 15.0; condylobasal length, 15.6, 14.9; least interorbital length, 5.4, 5.0; zygomatic breadth (one sex unknown), 11.6; breadth of braincase, 8.7, 8.7; palatal length (one sex unknown), 4.7. Habitat. — In Argentina, L. cinereus is distributed in most provinces from 40°S latitude northward, although no records are known from the provinces of San Juan, Chaco, Formosa, and Misiones (B^quez, 1987). In Tucuman, L. cinereus roosts in trees in gardens and along the streets. 1996 Mares et al. — TucumAn Mammals 119 Remarks. — Lasiurus cinereus is solitary. There is some evidence from North America that indicates that L. cinereus may migrate, although there are no data to support this in Tucuman Province or in Argentina. Lasiurus ega (Gervais) Nycticeus ega Gervais, in F. Compte de Castelnau, ExpMition dans les parties centrales de I’Amerique du Sud de Rio de Janeiro a Lima, et de Lima au Para; executee par ordre du gouvernement frangais pendant les annees 1843 a 1847, sous la direction de Grancis de Castelnau. P. Bertrand, Paris, (Sec. 7), Vol. 1, Pt. 2 (Mammiferes):73, 1856. Specimens Examined. — (9) Acheral, 1 (CML); Ciudad Capital, 1 (OMNH); Los Vazquez, 445 m, 2 (BMNH); San Miguel de Tucuman, 450 m, 4 (1 BMNH, 3 CML); Tafi Viejo, 1 (CML). Additional Record. — Munecas (Romana and Abalos, 1950). Measurements. — External measurements (two males, three females, unless not- ed otherwise): total length, 120.0, 106.0, 119.2 (115.0-124.0); tail, 52.0, 46.0, 55.8 (50.0-60.5); hindfoot, 9.0, 6.5, 8.3 (5.0-10.0); ear, 20.0, 15.0, 15.3 (14.0- 16.0) ; forearm (two males, four females), 47.0, 42.7, 46.8 (42.0-49.1); weight (one male), 13.0. Cranial measurements (one male, two females, unless noted otherwise): greatest length of skull, 16.4, 16.5, 17.0; condylobasal length, 15.4, 14.9, 16.0; least interorbital length, 4.5, 4.5, 4.5; zygomatic breadth (one male, one female), 11.2, 11.4; breadth of braincase, 8.2, 8.2, 8.7; length of maxillary toothrow (one male), 5.8; palatal length (one male), 5.5; length of mandibular toothrow (one male), 5.5; greatest length of mandible (one male), 12.1. Reproduction. — See Table 3. Habitat. — Lasiurus ega is known from northwestern, central, and eastern Ar- gentina, excluding the provinces of Chaco, Santiago del Estero, and Entre Rios, and northern Santa Fe (Barquez, 1987). In Tucuman, this bat is common in the humid and transitional forests. Lasiurus ega often uses plants, especially palms, in city gardens as refuges; they have also been reported to roost in the roofs of dwellings. Remarks. — Little is known about the biology of this species in Argentina, My- ers (1977) discussed the reproductive biology of this species in Paraguay. Geo- graphic variation in L. ega in the Chaco of Paraguay and Bolivia was discussed by Myers and Wetzel (1983), It is thought to be migratory. Myotis albescens (E. Geoffroy) Vespertilio albescens E. Geoffroy, Annales Museum d’Histoire Naturelle, Paris, 8:204, 1806. Specimens Examined. — (15) Aguas Chiquitas, Sierras de Medina, 800 m, 1 (CM); Concepcion, 1 (MACN); El Cadillal, Usina, 3 (CML); La Cocha, Dique San Ignacio, 1 (OMNH); Las Talas, 1 (CML); Las Talas, 4 km al N de Bella Vista, 1 (OMNH); Monteagudo, 4 (2 FCM, 1 MACN, 1 TTU); 3 km E Ticucho, cola de Cadillal Dique, 1 (OMNH); Ticucho, cola de embalse El Cadillal, 1 (OMNH); Tranquitas, 1 (MACN). Measurements. — External measurements (two males, four females, unless noted otherwise): total length, 83.0, 90.0, 88.6 (84.0-94.0); tail, 25.0, 31.0, 34.6 (31.0- 40.0) ; hindfoot, 7.5, 8.0, 7.4 (5.0-8.9); ear, 15.5, 14.0, 14.6 (14,0-15.7); forearm, 35.2, 34.1, 36.6 (35.9-37.3); weight (two males, three females), 7.0, 6.4, 7.8 (7.5- 8.0) . Cranial measurements (two males, three females, unless noted otherwise): greatest length of skull, 14.4, 13.6, 14.6 (14.2-15.0); condylobasal length, 13.4, 13.0, 13.7 (13.4-14.0); least interorbital length, 4.4, 4.0, 4.1 (4. 1-4.1); zygomatic breadth (two males, two females), 8.6, 8.6, 8.6, 8.7; breadth of braincase, 7.4, 6.9, 7.2 (7. 1-7.3); length of maxillary toothrow, 5.3, 5.2, 5.3 (5. 2-5. 5); palatal 120 Annals of Carnegie Museum VOL. 65 length (two males, two females), 5.5, 5.4, 5.2, 5.8; length of mandibular toothrow (one male, two females), 4.9, 5.2, 5.3; greatest length of mandible, 9.2, 10.2, 9.5 (9.3-9.6). Reproduction. — See Table 3. Habitat.— My Otis albescens is known from the provinces of the northwest and the northeast to south of Buenos Aires Province (Olrog and Lucero, 1981; Bar= quez, 1987). In Tucuman, this species is found in grasslands and savannas (Lu- cero, 1983), in forests, along rivers, and in open Chacoan thorn scrub (Barquez, 1987). Remarks. — At Las Talas, individuals of M. albescens, Eptesicus furinalis, and Tadarida brasiliensis were found in the roof of a rural dwelling (Barquez, 1988). Myotis keaysi J. A. Allen Myotis ruber keaysi J. A. Allen, Bulletin of the American Museum of Natural History, 33:383, 1914. Specimens Examined. — (7) Agua Colorada, 3 (MACN); El Nogalar, km 43, Ruta 307, 1700 m, 1 (CML); Piedra Tendida, 12 km WNW Burruyacu, along Rio Cajon, 2,500 ft., 1 (OMNH); Tranquitas, 2 (1 MACN, 1 TTU). Measurements. — External measurements (one male): total length, 90.0; tail, 37.0; hindfoot, 8.0; ear, 14.0; forearm, 40.0; weight, 6.0. Cranial measurements (one male): greatest length of skull, 13.9; condylobasal length, 13.3; least inter- orbital length, 3.7; zygomatic breadth, 8.8; breadth of braincase, 7.1; length of maxillary toothrow, 5.1; palatal length, 5.8; length of mandibular toothrow, 5.2; greatest length of mandible, 9.2. Habitat. — In Argentina, this species is known only from Salta and Tucuman (Barquez, 1987). In other parts of its range, M. keaysi is commonly found at localities above 2000 m (Barquez and Ojeda, 1992). In Tucuman, however, it has been captured at much lower elevations in transitional forest and montane forests. These localities are situated primarily in the northeastern part of the province, near Chacoan vegetation, and isolated from the principal mountain chains to the west (Barquez, 1987; Barquez and Ojeda, 1992). Remarks. — This species, in addition to Sturnira erythromos and Myotis nigri- cans, was captured in nets placed along a river at Piedra Tendida (Mares et al., 1995). An adult male was captured in June at Piedra Tendida. Bedford and Ei- senberg (1992) do not show this bat as occurring in the southern cone. Myotis levis (1. Geoffroy) Vespertilio levis I. Geoffroy, Annales des Sciences Naturelles, Zoologie, ser. 1, 3:444-445, 1824. Specimens Examined. — (55) Agua Rosada, 2 (CML); Aguas Chiquitas, 25 km NE San Miguel de Tucuman, 800 m, 5 (CM); Amaicha del Valle, 2000 m, 1 (OMNH); Colalao del Valle, 1 (BMNH); Concepcion, 4 (3 BMNH, 1 MACN); Dique San Ignacio, 1 (CML); El Cadillal, 1 (CML); Horco Molle, 1 (CM); La Cienaga, 2500 m, 1 (BMNH); La Higuera, 1 (CML); Las Pavas, Aconquija, 1 (MACN); Las Talas, 3 (CML); Leocadio Paz, 1 (MACN); Norco, Vipos, 2500 m, 2 (BMNH); San Miguel de Tucuman 7 (5 BMNH, 2 MACN); San Pedro de Colalao, 1 (CML); at km marker 42, on highway 364, south of San Pedro de Colalao, 4,700 ft., 13 (3 CML, 3 lADIZA, 7 OMNH); Tacanas 1 (CML); Tafi del Valle, 1 (CML); Tucuman, no specific locality, 2 (BMNH); Vipos, 5 (CML). Measurements. — External measurements (five males, 16 females, unless noted otherwise): total length, 86.6 (75.0^91.0), 89.3 (77.0-100.0); tail, 39.0 (36.0- 40.0), 40.2 (35.5-42.0); hindfoot (five males, 15 females), 7.8 (5. 0-9.0), 8.7 (7.3- 9.3); ear, 14.3 (14.0-15.5), 14.0 (13.0-16.0); forearm (four males, 15 females), 38.0 (36.8-40.0), 38.0 (36.7-40.0); weight (four males, 14 females), 5.2 (4.3- 1996 Mares et al. — Tucuman Mammals 121 5.5), 6.1 (4.4~8.0). Cranial measurements (two males, seven females, unless noted otherwise): greatest length of skull, 14.0, 14.4, 14.3 (14.1-14.8); condylobasal length, 13.3, 13.5, 13.5 (13.0-14.0); least ieterorbital length, 3.5, 3.6, 3.5 (3.4- 3.7); zygomatic breadth (two males, six females), 8.6, 8.2, 8.6 (8. 2-9.0); breadth of braiecase, 7.1, 6.8, 6.9 (6.6-7.2); length of maxillary toothrow (two males, six females), 5.4, 5.6, 5.6 (5.4-5. 8); palatal length (two males, six females), 6.4, 7.0, 6,6 (5.9-7.4); length of mandibular toothrow (two males, two females), — , 5.7, 5.8, 6.0; greatest length of mandible (two males, six females), 10.1, 10.2, 10.6 (10.4-10.8). Reproduction.ScQ Table 3. Barquez (1987) reported capturing lactating fe- males from the end of December to the end of January in transitional forest at Aguas Chiquitas, Molt.^ — -Three adults, captured in January at San Pedro de Colalao, were molting on the dorsum. Habitat.— This species is widely distributed throughout northern, central, and eastern Argentina as far south as Neuquen Province (Barquez, 1987). Although preferring open, natural habitats, it is found also in urban areas; elevation ranges up to 3500 m (B^quez, 1987). Many of the specimens were captured in transi- tional forest. Individuals from San Pedro de Colalao were collected in a rural dwelling; the surrounding habitat was transitional forest with a lot of secondary growth. Remarks.— My Otis levis is one of the most common vespertilionid bats in the province, although little is known about its biology. This species is easily confused with other Myotis. At San Pedro de Colalao this species was captured with Stur- nira erythromos. Myotis nigricans (Schinz) Vespertilio nigricans Schinz, Das Thierreich, 1:179, 1821. Specimens Examined.- — (8) Acheral, 1 (CML); Agua Colorada, 1 (TTU); Cerro del Campo, 1 (CML); El Cadillal, 25 km N San Miguel de Tucuman, 1 (CM); El Naranjo, 1 (TTU); Piedra Teedida, 12 km WNW Burrayacu, along Rfo Cajon, 2,500 ft., 1 (OMNH); Taff Viejo, 2 (CML). Additional Records. — Monte Bello, Rio Chico (Romana and Abalos, 1950). Measurements.— External measurements (two males): total length, 78.0, 78.0; tail, 38.0, 31.0; hindfoot, 6.9, 7.0; ear, 15.7, 14.0; forearm, 33.0, 33.0; weight, 4.5, 3.5. Cranial measurements (one male): greatest length of skull, 13.1; con- dylobasal length, 12.8; least interorbital length, 3.4; zygomatic breadth, 8.3; breadth of braiecase, 6.4; length of maxillary toothrow, 5.1; palatal length, 5.7; length of mandibular toothrow, 5.3; greatest length of mandible, 9.7. Reproduction.- — See Table 3. Little is known about the reproductive biology of this species in Argentina. Moll— The individual from Piedra Tendida was not molting. Habitat.— In Argentina, this species is restricted to the northern provinces, Ju- juy, Salta, Tucuman, Santiago del Estero, Chaco, Formosa, Corrientes, and Mi- siones (Barquez, 1987). Based on capture records, M. nigricans prefers humid and transitional forests. B^quez (1987) noted that this species has been captured in a variety of habitats, from Yungas forest to urban areas, in cities and in rural areas, under roofs and under tree bark. Remarks.- — Other bats collected at Piedra Tendida were Sturnira erythromos and Myotis keaysi (Mares et aL, 1995). Individuals have been captured in February (n = 1), March (n = 1), and July (n = 1). 122 Annals of Carnegie Museum VOL. 65 Family Molossidae Eumops bonariensis (Peters) Promops bonariensis Peters, Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin, 1874:232, 1874. Specimens Examined. — (17) Aguas Chiquitas, 3 (CML); Aguas Chiquitas, Sierras de Medina, 800 m, 4 (CM); Reserva Provincial “Aguas Chiquitas” El Cadillal, 3 (CM); Las Talas, 4 km al N de Bella Vista, 3 (OMNH); 29 km NE San Miguel de Tucuman, 1 (CM); 28 km NE San Miguel de Tucuman, 3 (CM). Measurements. — External measurements (three males, eight females): total length, 121.0 (117.0-128.0), 115.1 (105.0-129.0); tail, 40.5 (38.0-43.0), 37.8 (33.0-41.0); hindfoot, 9.6 (8.3-10.3), 8.4 (7.0-10.2); ear, 22.0 (21.4-23.0), 22.7 (21.9-24.0); forearm, 47.9 (47.3-48.7), 48.0 (44.8-49.0); weight, 18.2 (17.5- 19.0), 17.4 (15.5-20.0). Cranial measurements (three males, eight females, unless noted otherwise): greatest length of skull, 19.3 (19.1-19.5), 18.9 (18.3-19.2); condylobasal length, 18.3 (18.1-18.6), 18.0 (17.2-18.4); least interorbital length, 4.3 (4.2-4.3), 4.2 (4.0-4.3); zygomatic breadth, 11.8 (11.5-12.0), 11.5 (1 Fi- ll.7); breadth of braincase, 9.6 (9. 5-9. 7), 9.3 (8. 8-9. 7); length of maxillary tooth- row, 7.5 (7. 3-7. 6), 7.3 (7. 1-7.5); palatal length (three males, seven females), 7.3 (7. 1-7.4), 7.6 (7. 2-7. 9); length of mandibular toothrow, 6.7 (6. 6-6. 9), 6.7 (6.5- 7.3); greatest length of mandible, 14.0 (13.8-14.1), 13.8 (13.6-14.0). Reproduction. — See Table 3. Barquez (1987) reported that individuals captured at Las Talas at the end of March and the end of April were not reproductively active. Habitat. — In Argentina, E. bonariensis has been recorded from the provinces of Buenos Aires, Santa Fe, Entre Rios, Cordoba, and Tucuman (Barquez, 1987). In Tucuman, this species prefers dense transitional forests, but may also be found in rural and open areas, even in roofs and tree holes. At Aguas Chiquitas, a locality surrounded by Chaco serrano, individuals were collected in nets placed across areas of calm water in the forest. Remarks. — Barquez (1987) discussed the relationship of E. bonariensis and E. patagonicus, revalidating the distinction of E. patagonicus. Both species are found in sympatry in Tucuman Province. At Las Talas E. bonariensis were collected with E. patagonicus (reproductively active), E. furinalis, M. albescens, M. levis dinellii, and T. brasiliensis (Barquez, 1987). The biology of this species in Ar- gentina is poorly known. Eumops dabbenei Thomas Eumops dabbenei Thomas, Annals and Magazine of Natural History, ser. 8, 13:481, 1914. Specimens Examined. — (2) San Miguel de Tucuman, 2 (CML). Reproduction. — A juvenile (with cartilaginous phalanges and deciduous upper incisors) was captured in a house in mid-December in the city of Tucuman (Bar- quez and Lougheed, 1990; Barquez and Ojeda, 1992). Habitat. — In Argentina, E. dabbenei is known only from the provinces of Cha- co, Santa Fe, and Tucuman (Barquez, 1987; Barquez and Lougheed, 1990; Bar- quez and Ojeda, 1992). Remarks. — Little is known about the biology of this species in Argentina. Eumops glaucinus (Wagner) Dysopes glaucinus Wagner, Archiv fiir Naturgeschichte, 9 (1):368, 1843. Specimens Examined. — (2) San Miguel de Tucuman, 2 (CML). 1996 Mares et al. — Tucuman Mammals 123 Habitat.— In Argentina, E. glaucinus is known only from the provinces of Jujuy, Salta, and Tucuman (Barquez, 1987; Barquez and Lougheed, 1990). The inclusion of the provinces of Catamarca and La Rioja by Olrog and Lucero (1981) is not documented by specimens. In the city of Tucuman it roosts in trees and in cracks in buildings. Eumops patagonicus Thomas Eumops patagonicus Thomas, Annals and Magazine of Natural History, sen 9, 13:234, 1924. Specimens Examined.— {IT} Burrayacu, 1 (MACN); Gobernador Piedrabuena, 20 (17 MACN, 3 TTU); Las Taias, 4 km al N de Bella Vista, 1 (OMNH). Measurements.— ExtGin^l measurements (one male): total length, 92.4; tail, 35.8; hindfoot, 7.4; ear, 19.7; forearm, 45.4; weight, 15.0. Cranial measurements (one male): greatest length of skull, 18.4; condylobasal length, 16,5; least inter- orbital length, 4.0; zygomatic breadth, 10.4; breadth of braincase, 9.0; length of maxillary toothrow, 6.6; palatal length, 6.6; greatest length of mandible, 13.1. Reproduction.—SGC Table 3. Habitat.— In Argentina, this species is known from an isolated record in Chubut (type locality), Buenos Aires, Santa Fe, and all of the northern provinces except Jujuy and Catamarca. In Tucuman, E. patagonicus is found in open suburban and urban areas, especially in the Chaco. Remarks.— BmquQx (1987) discussed the relationship of this species with E. bonariensis and synonymized E. b. beckeri with E. patagonicus. Eumops perotis (Schinz) Molossus perotis Schinz, in Cuvier, Das Thierreich, 1:870, 1821. Specimens Examined.— {AO) Concepcion, 27 (7 BMNH, 20 MACN); Las Talitas, 1 (CML); Plaza Independencia, San Miguel de Tucuman, 1 (CM); San Miguel de Tucuman, 10 (2 BMNH, 8 CML); San Pedro de Colalao, 1 (CML). Additional Record.- — -Caspichango (Massoia, 1976). Measurements.— External measurements (three males, seven females, unless noted otherwise): total length, 172.7 (170.0^178.0), 167.5 (155.0^180.0); tail, 60.3 (60.0-61.0), 61.8 (55.0-70.0); hindfoot (four males, six females), 16.1 (15.0-18.4), 15.0 (14.0-16.0); ear, 42.3 (35.0-49.0), 34.7 (33.0-35.0); forearm (six males, five females), 78.2 (75.8-82.5), 78.3 (77.3-80.0); weight (two males), 52.0, 65.0. Cranial measurements (four males, one female, unless noted other- wise): greatest length of skull, 31.0 (30.2-31.7), 29.3; condylobasal length, 30.4 (29.0-30.8), 27.2; least interorbital length (five males, three females), 5.3 (5.1- 5.6), 5.8 (5.6-5.9); zygomatic breadth, 18.4 (17.5-18.8), 19.5; breadth of brain- case, 13.4 (12.6-14.9), 12.6; length of maxillary toothrow (three males, four fe- males), 12.4 (11.9-12.7), 12.9 (12.5-13,2); palatal length (four males, three fe- males), 12.7 (9.0-14.2), 13.9 (13,2-15.0); length of mandibular toothrow (three males, four females), 13.4 (12.2-14.1), 13.8 (13.3-14.0); greatest length of man- dible (three males, four females), 24.4 (24.1-24.7), 23.3 (23.0-23.6). Reproduction.— Table 3. Barquez (1987) reported catching a juvenile in the city of Tucuman in mid-December that had deciduous dentition and cartilag- inous phalanges. Habitat. — -This species ranges throughout all of northern Argentina, except the Chaco; it is distributed southward to central Cordoba (Barquez, 1987). In the city 124 Annals of Carnegie Museum VOL. 65 of Tucuman these bats live in palm trees and under roofs. In natural areas, they have been known to roost in rock crevices. Molossops temminckii (Burmeister) Dysopes temminckii Burmeister, Systematische iibersicht der Thiere Brasiliens. Georg Reimer, Berlin, p. 72, 1854. Specimens Examined. — (9) Cerro del Campo, 2 (CML); El Cadillal, 1 (CML); La Higuera, 1 (CML); Las Mesadas, 3 (CML); Puesto Portadera, 2(1 MACN, 1 TTU). Additional Record. — San Miguel de Tucuman (Fornes and Massoia, 1968). Measurements. — External measurements: forearm (two males, four females), 31.4, 31.7, 31.1 (30.0-31.5). Cranial measurements (two males, two females, unless noted otherwise): greatest length of skull (one male, one female), 13.8, 13.9; condylobasal length (one female), 14.8; least interorbital length (two males, three females), 3.7, 3.6, 3.7 (3. 7-3. 7); breadth of braincase, 7.0, 7.0, 7.0, 7.2; length of maxillary toothrow (one male, two females), 5.3, 4.9, 5.2; palatal length (one female), 6.1; length of mandibular toothrow (one male, two females), 5.5, 5.6, 5.8. Reproduction. — See Table 3. A male with scrotal testes was collected at El Cadillal in mid-October (B^quez, 1987). Habitat. — In northwestern Argentina (Jujuy, Salta, Santiago del Estero, and Tucuman), the preferred habitat is Chaco. This species is also known from the Chaco of Formosa, Santa Fe, Corrientes, and Chaco provinces; isolated records have also been reported from Buenos Aires and Chubut provinces (Barquez, 1987). Remarks. — Myers and Wetzel (1983) reported dramatic geographic variation in this species in the Chaco of Paraguay, Bolivia, Brazil, and northern Argentina. Individuals are small and light in color in the northwestern Chaco, and large and dark in the eastern Chaco. Specimens from the base of the Andes are as large as, or larger than, those from the eastern Chaco. It appears that specimens from Tucuman Province follow this trend. Molossus molossus (Pallas) Vespertilio molossus Pallas, Miscellanea Zoologica. Hagae Comitum, pp. 49-50, 1766. Specimens Examined. — (7) Ciudad Tucuman, casa centrica, 1 (OMNH); El Manantial, 1 (CML); San Miguel de Tucuman, 3 (CML); San Pedro de Colalao, 1 (CML); Trancas, 1 (CML). Measurements. — External measurements (one female): total length, 98.0; tail, 40.0; hindfoot, 12.0; ear, 7.0; forearm, 38.0; weight, 10.0. Reproduction. — See Table 3. Habitat. — In Argentina, M. molossus is found in the northwestern provinces of Jujuy, Tucuman, Salta, and Santiago del Estero. In eastern Argentina, it ranges from Formosa, Corrientes, Chaco, and Misiones south to Entre Rios and northern Buenos Aires (Barquez, 1987). In Tucuman, this species roosts in tree holes and crevices, roofs, and tunnels. It is common in both urban and suburban areas. Remarks. — This bat may form large colonial groups. Bedford and Eisenberg (1992) do not show this species as occurring in Tucuman Province. Nyctinomops macrotis (Gray) Nyctinomus macrotis Gray, Annals of Natural History, 4:5, 1840. Specimens Examined. — (6) San Miguel de Tucuman, 6 (CML). 1996 Mares et al.— -TucumAn Mammals 125 Reproduction.— See Table 3, Habitat. — In Argentina, N. macrotis has been reported only from the provinces of Jujuy, Salta, Tucumae, Catamarca, and La Rioja (Barquez, 1987). ^^M£irA^.-=-This species is common in the city of Tucuman where it uses build- ings for shelter (B^quez, 1987; Barquez and Ojeda, 1992). Little is known about the biology of this species in Argentina. Promops nasutus (Spix) Molossus nasutus Spix, Simiaram et Vespertilionum Brasiliensium species novae. Monachii, p. 58, 1823. Specimens Examined. — -(3) Rfo Loro, 2 (OMNH); San Miguel de Tucuman, 1 (BMNH). Reproduction.- — See Table 3. HabitaP — Promops nasutus is distributed in the Argentine provinces of Salta, Jujuy, Tucuman, Catamarca, Santiago del Estero, and Misiones (Barquez, 1987). Remarks.— The individuals from Rfo Loro were captured as they left a roost located in a crevice in a river bank (Barquez, 1987). Tadarida brasiliensis (L Geoffroy) Nyctinomus brasiliensis L Geoffroy, Aonales des Sciences Naturelles, Zoologie, 1:343, 1824. Specimens Examined. — (113) Burruyacu, 6 (5 MACN, 1 TTU); Cerro San Javier, 1 (CML); Con- cepcion, 10 (5 BMNH, 5 MACN); Dique San Ignacio, 20 (CML); El Cadillal, 1 (CML); Estancia El Cavao, 1 (CML); Estancia San Pedro, Vipos, 2 (CML); Gobernador Piedrabuena, 5 (2 MACN, 3 TTU); Horco Molle, 12 (CM); Biological Reserve at Horco Molle, at residencia, 3 (1 lADIZA, 2 OMNH); Iglesia, 3 (TTU); Instituto Lillo, San Miguel de Tucuman, 1 (CML); Las Juntas, 22 km W Choromoro on Hwy 312, 3,500 ft., 5 (3 lADIZA, 2 OMNH); Las Mesadas, 1 (CML); Las Pavas, 5 (MACN); Las Talas, 2 (CML); Las Talas, 4 km al N de Bella Vista, 2 (OMNH); Marcos Paz, 1 (CML); Playa Larga, 4 (CML); San Miguel de Tucuman, 14 (3 BMNH, 4 USNM, 7 CML); San Pedro de Colalao, 2 (CML); Ruta 307, km 19.7, camieo a Taff del Valle, 750 m, 3 (OMNH); Tacanas, 2 (CML); Tapia, 1 (CML); Timbo Nuevo, 4 (1 MACN, 3 TTU); Vipos, 2 (CML). Additional Records. — Ingenio Lules (Romana and Abalos, 1950); Monteagudo (Fornes and Massoia, 1967). Measurements.- — ^External measurements (nine males, eight females, unless not- ed otherwise): total length, 96.8 (91.0^102.0), 98.6 (87.7-105.0); tail, 35.9 (33.0- 40.0) , 37.2 (30.0-41.0); hindfoot, 7.9 (7.2-9.0), 8.6 (7.2-10.0); ear, 17.2 (13.0- 21.5), 18.5 (12.0-21.0); forearm (seven males, ten females), 43.3 (4L3-45.0), 43.8 (42,0-45,0); weight (six males, six females), 12.2 (10.0-14.0), 11.5 (10.0- 12.0) . Cranial measurements (ten males, five females, unless noted otherwise): greatest length of skull, 16.7 (15.4-17.2), 16.5 (16.1-16.9); condylobasal length, 15.7 (14.5-16.2), 15.5 (15.5-15.7); least interorbital length (ten males, six fe- males), 4.0 (3. 8-4. 2), 4.0 (3. 9-4.0); zygomatic breadth (seven males, four fe- males), 9.9 (9.3-10.4), 9.6 (9. 2-9. 9); breadth of braincase (ten males, six females), 8.4 (7. 9-8. 6), 8,3 (8. 0-8. 6); length of maxillary toothrow (ten males, six females), 5.7 (4.7-6.4), 6.0 (4. 8-6. 7); palatal length (seven males, five females), 6.8 (6.1- 7.1), 6.2 (3. 6-7. 5); length of mandibular toothrow (six males, five females), 6.2 (5. 9-6. 6), 7.6 (6.2-12.0); greatest length of mandible (five males, four females), 12.1 (11.7-12.6), 11.9 (11.7-12.1). Reproduction.See Table 3. Molt. — An adult male captured at Horco Molle in January was molting on both the dorsum and venter. Habitat. — -This is one of the most widely distributed species in Argentina, 126 Annals of Carnegie Museum VOL. 65 found in all provinces north of 43°S latitude, except La Rioja, Chaco, and Cor- rientes (B^quez, 1987). It is most common in transitional forest, Chaco, and montane forest. The specimens collected at Horco Molle were roosting behind a rain gutter. Remarks.-— In the province, this species is colonial and forms large groups that roost in caves, holes, crevices, tunnels, and roofs. In the city of Tucuman, T. brasiliensis is one of the most common species and is found roosting in building crevices and roofs. At Las Juntas, Artibeus planirostris, Sturnira erythromos, His- tiotus macrotus, and Lasiurus blossevillii also were captured. Sturnira erythromos also was captured at Horco Molle. Order Primates Family Cebidae Subfamily Cebinae Cebus apella (Linnaeus) Simia apella Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth edition. Laurentii Salvii, Stock- holm, 1:28, 1758. Specimens Examined. — (0) None. Additional Record. — Dique Escaba (M. L. Campbell, personal observation). Remarks. — Local inhabitants in the area surrounding the Embalse Escaba in southwestern Tucuman indicated that monkeys had been present in the area as late as the early 1980s (M. L. Campbell and J. K. Braun, personal observation), but we have been unable to confirm this. Bedford and Eisenberg (1992) do not show this species as occurring in the province. Order Carnivora Family Canidae Cerdocyon thous (Linnaeus) Cams thous Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Volume 1. Regnum Animale. 12th edi- tion. Laurentii Salvii, Stockholm, 1:60, 1766. Specimens Examined. — (4) Monteros, 1 (MACN); Taficillo, 1 (CML); Tapia, 500 m, 1 (BMNH); Vipos, 1 (BMNH). Measurements. — External measurements (one female): total length, 932.0; tail, 322.0; hindfoot, 140.0; ear, 67.0. Cranial measurements (one female): greatest length of skull, 127.6; condylobasal length, 133.0; least interorbital length, 26.2; zygomatic breadth, 74.4; breadth of braincase, 46.1; length of maxillary toothrow, 59.0; palatal length, 59.3; length of mandibular toothrow, 62.9; greatest length of mandible, 106,4. Habitat.- — Cerdocyon thous is distributed in northern Argentina east of the An- des, including the provinces of Salta, Jujuy, Tucuman, Formosa, Misiones, Cor- rientes, Santa Fe, Chaco, and Santiago del Estero (Olrog and Lucero, 1981). Lucero (1983) and Olrog and Lucero (1981) noted that this species prefers tran- sitional and humid forests, and mountains, where it occurs up to 2500 m. Remarks. — One individual each was captured in May, July, and October. The biology of this species was summarized by Berta (1982). Ginsberg and Macdonald (1990) noted that it is considered out of danger by the Argentine Wildlife Board 1996 Mares et al. — Tucuman Mammals 127 (Direccion Nacional de Fauna Silvestre). Nothing is known of the biology of this species in the province. Pseudalopex culpaeus (Molina) Cams culpaeus Molina, Saggio sulla storia eaturale del Chili. S. Tommaso d’ Aquino, Bologna, p. 293, 1782. Specimens Examined. — (4) Acoequija, 1 (BMNH); Agua Rosada, 1 (CML); El Duraznito, 1 (CML); San Javier, 1 (CML). Measurements.— measurements (one female): hindfoot, 130.0; ear, 90.0, Cranial measurements (one female, two sex unknown): greatest length of skull, 130.6, 129.8, 129.2; condylobasal length, 129.4, 124.3, 129.0; least inter- orbital length, 23.1, 23.8, 24.0; zygomatic breadth, 67.9, 66.5, 67.5; breadth of braincase, 44.7, 44.5, 45.3; length of maxillary toothrow, 59.5, 58.2, 58.2; length of mandibular toothrow, 66.2, 64.9, 65.8; greatest length of mandible, 95.3,— — . Habitat.— In. Argentina, the distribution of P. culpaeus extends as a narrow band from the northwest, widening southward to include all of the southern prov- inces south of Buenos Aires Province (Olrog and Lucero, 1981). This species prefers arid areas and steppes in the mountains at altitudes up to 4500 m (Olrog and Lucero, 1981; Lucero, 1983). Remarks.— Pseudalopex culpaeus has an Appendix II ranking in Argentina (Fuller and Swift, 1984). The biology of this species was summarized by Ginsberg and Macdonald (1990), who also noted that it has endangered status in Argentina. The effects of hunting on populations of this species in Neuquen Province was examined by Novaro (1995). Pseudalopex griseus (Gray) Vulpes griseus Gray, Magazine of Natural History (Charlesworth’s), 1:578, 1837. Specimens Examined. — (4) Cerro del Campo, 800 m, 2 (BMNH); Tapia, 500 m, 1 (BMNH); Vipos, 500 m, 1 (BMNH). Measurements. — External measurements (two males, two females): total length, 887.0, 829.0, 884.0, 887.0; tail, 347.0, 328.0, 322.0, 317.0; hindfoot, 131.0, 125.0, 120.0, 124.0; ear, 81.0, 70.0, 80.5, 82.5. Cranial measurements (two males): great- est length of skull, 124.4, 121.7; condylobasal length, 128.8, 125.4; least inter- orbital length, 24.8, 23.4; zygomatic breadth, 70.8, 70.7; breadth of braincase, 45.6, 45.9; length of maxillary toothrow, 59.3, 59.3; palatal length, 56.4, 56.8; length of mandibular toothrow, 66.0, 66.0; greatest length of mandible, 97,6, 97.6. Habitat.— In Argentina, the geographic distribution of this species is similar to that of P. culpaeus, although it is not found in Jujuy and eastern Salta (Olrog and Lucero, 1981). Moreover, P. culpaeus is a fox of the high mountains, whereas P. griseus is primarily a lowland animal. This fox frequents steppes and arid shrubby regions in western, southern, and northeastern Tucuman Province (Lucero, 1983). Remarks.— Two specimens were captured in August and two were captured in October. The Cerro del Campo specimens were listed as P. gracilis by Thomas (1926). In Argentina, this species is considered vulnerable (International Union for the Conservation of Nature, 1990, 1994). The biology of this species was summarized by Ginsberg and Macdonald (1990), who noted its endangered status. Hunting of this species has been banned in Tucuman Province, where little is known about its biology. 128 Annals of Carnegie Museum VOL. 65 Pseudalopex gymnocercus (Fischer) Procyon gymnocercus Fischer, Zoognosia, 3:xi, 178, 1814. Specimens Examined. — (11) Agua Rosada, 1 (CML); Burruyacu, 1 (MACN); Chicligasta, 1 (CML); El Simbolar, 1 (CML); El Timbo, 4 (CML); Monteros, 1 (MACN); Nueva Espana, 1 (CML); Taff Viejo, 1 (CML). Measurements. — External measurements (one male, one female): total length, 1010.0, 835.0; tail, 330.0, 305.0; hindfoot, 110.0, 120.0; ear, 80.5, 75.0. Cranial measurements (one male, three females, one sex unknown, unless noted other- wise): greatest length of skull (one male, one female, one sex unknown), 129.3, 123.6, 123.5; condylobasal length, 125.4, 123.5 (120.0-127.7), 117.5; least inter- orbital length (one male, two females, one sex unknown), 20.5, 22.5, 20.9, 23.1; zygomatic breadth (one male, three females), 63.6, 67.5 (64.0-71.4); breadth of braincase, 45.9, 45.2 (43.9-45.9), 43.1; length of maxillary toothrow (one male, one female, one sex unknown), 56.8, 55.0, 50.9; palatal length (three females, one sex unknown), 63.5 (61.5-64.6), 60.9; length of mandibular toothrow (one male, one sex unknown), 64.6, 60.0; greatest length of mandible (one male, one sex unknown), 93.2, 89.5. Habitat. — According to Olrog and Lucero (1981), P. gymnocercus inhabits all of northern and central Argentina except Misiones Province. The southern distri- butional limits are northern Rio Negro and Neuquen provinces. This fox is found in savannas, mountains, and forests up to 2000 m in elevation (Lucero, 1983). Remarks. — Specimens have been collected in nearly every month: February {n = 1), March {n = 1), April {n — 1), May {n = 2), June {n = 1), July {n = 2), September {n = 1), and November {n = 1). Ginsberg and Macdonald (1990) summarized the biology of this species. Crespo (1971) studied the ecology of P. gymnocercus in La Pampa Province. Family Felidae Subfamily Felinae Herpailurus yaguarondi (Lacepede) Felis yaguarondi Lacepede, in Azara, Voyage dans I’Amerique Meridionale. Imprimieur-Libraire, Par- is, Atlas, 1809, pi. 10, 1809. Specimens Examined. — (3) Concepcion, 1 (CML); El Bracho, 1 (CML); Villa Nougues, 1 (CML). Additional Record. — Dique Escaba (M. L. Campbell, personal observation). Measurements. — Cranial measurements (one male): greatest length of skull, 96.5; condylobasal length, 91.9; least interorbital length, 17.3; zygomatic breadth, 62.0; breadth of braincase, 42.1; length of mandibular toothrow, 20.7; greatest length of mandible, 58.1. Habitat. — This species is distributed throughout northern and central Argentina south to northern Neuquen and Rio Negro (Olrog and Lucero, 1981). Herpailurus yaguarondi prefers savannas, mountains, and forests to the southern limits of the province (Olrog and Lucero, 1981; Lucero, 1983). Remarks. — The record of this species at Dique Escaba is based on information obtained from local residents who indicated that a darkly colored cat was present in the area (M. L. Campbell, personal observation). The status of this cat in Argentina is considered indeterminate (i.e., it is known to be either endangered, vulnerable, or rare, but there is too tittle information to denote a category). In Argentina, this species has an Appendix II ranking (Fuller and Swift, 1984). 1996 Mares et al. — TucumAn Mammals 129 Leopardus pardalis (Linnaeus) Felis pardalis Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth edition. Laurentii Salvii, Stock- holm, 1:42, 1758. Specimens Examined. — (2) Cerro del Campo, 1 (CML); La Fronterita, 1 (MACN). Additional Records. — Burruyacu; La Cocha (Olrog, 1976). Measurements. — Cranial measurements (one male): greatest length of skull, 101.1; condylobasal length, 94.8; least interorbital length, 18.6; zygomatic breadth, 65.8; breadth of braincase, 51.3; greatest length of mandible, 64.6. Habitat.— Ocelots are found only in the northernmost Argentine provinces and Tucuman. Olrog and Lucero (1981) and Lucero (1983) noted that this species is found in transitional forests, mountains, and savannas to southern Tucuman Prov- ince. Remarks. — ^Cabrera (1961:177) listed a specimen of Leopardus wiedii from Burruyacu, Tucuman. Leopardus wiedii does not exist in the province. The spec- imen described is a young individual corresponding to the Leopardus pardalis from Cerro de Campo listed above (Olrog, 1976). This cat was listed as vulnerable in Argentina in 1990 (International Union for the Conservation of Nature, 1990), however it now is unlisted (International Union for the Conservation of Nature, 1994). In Argentina, this subspecies is listed in the Appendix I category and is protected (Fuller and Swift, 1984). This species is apparently still relatively com- mon in the province, although it is not easily observed. The specimen from Cerro del Campo was collected in May. Specimens from Burruyacu were collected in 1971 and 1972, and from La Cocha in 1973 (Olrog, 1976). Lynchailurus pajeros (Desmarest) Felis pajeros Desmarest, Chat, Felis, Linn., Briss. Schreb., Cuv., in Nouveau dictionnaire d’Histoire Naturelle, appliquee aux art, a Fagriculture, a Feconomie rurale et domestique, a la medecine, etc. Par une societe de naturalistes et d’agriculteurs (nouvelle edition), 6:114, 1816. Specimen Examined. — (1) Concepcion, 1 (MACN). Additional Records. — San Pedro de Colalao (M. A. Mares, personal observation); La Florida Re- serva Provincial; Santa Ana Reserva Provincial (Anonymous, in press). Habitat.— species is distributed throughout most of Argentina, although its presence in the Chaco is questionable (Olrog and Lucero, 1981; Garcia- Perea, 1994). Lynchailurus pajeros is found in mountains, grasslands, and savannas in the western part of the province (Lucero, 1983). Remarks. — Lucero (1983) mentioned that no recent data indicate its presence in the province, but Mares photographed a specimen taken above San Pedro de Colalao in 1976. In Argentina, this species has an Appendix II ranking (Fuller and Swift, 1984). The status of L. pajeros is given as indeterminate by the Inter- national Union for the Conservation of Nature (1994). Records of this species are rare. The animal photographed by Mares (Mares and Ojeda, 1984:583) was taken in the high-elevation grasslands in the area near San Pedro de Colalao. Garcia- Perea (1994) reviewed the systematics and biogeography of the members of this genus. Oncifelis geojfroyi (d’Orbigny and Gervais) Felis geojfroyi d’Orbigny and Gervais, Nouveau bulletin des sciences par la Societe Philomatique, Paris, 1844:40, 1844. 130 Annals of Carnegie Museum VOL. 65 Specimens Examined. — (1 1) Atahona, 1 (CML); Concepcion, 1 (MACN); Departamento Burruyacu, 1 (CML); La Ramada, 1 (CML); Leales, 1 (CML); Quebrada de la Angostura, 1 (CML); Quebrada de los Matos, 1 (CML); San Pedro de Colalao, Junta de los Rios, 2 (CML); Trancas, 1 (CML); Vipos, 500 m, 1 (BMNH). Additional Records. — El Manantial (Burmeister, 1943); Piedra Tendida, 12 km WNW Burruyacu, along Rio Cajon, 2,500 ft. (M. L. Campbell, in litt.); Pozo Hondo (R. M. Barquez, personal obser- vation); San Pedro de Colalao (M. A. Mares, personal observation). Measurements. — External measurements (one young female): total length, 679.0; tail, 255.0; hindfoot, 94.0; ear, 44.0; weight, 81.2. Cranial measurements (one male, three females, unless noted otherwise): greatest length of skull, 91.6, 78.6 (65.7-88.6); condylobasal" length (one male, two females), 88.J, 61.2, 76.1; least interorbital length, 16.2, 14.3 (11.2-17.2); zygomatic breadth, 61.4, 53.9 (43.4-64.9); breadth of braincase, 44.6, 40.6 (39.1-43.0); length of maxillary toothrow (one male, two females), 22.1, 19.1, 21.1; length of mandibular toothrow (one male, two females), 21,5, 19.3, 19.2; greatest length of mandible, 58.5, 48.8 (39.2-57.8). Habitat. — Oncifelis geojfroyi is distributed throughout all of Argentina except Tierra del Fuego (Olrog and Lucero, 1981). In 1990, locals stated that this species of cat occurred in the transitional forest around Piedra Tendida (M. L. Campbell, personal observation). Remarks. — In Argentina, this species has an Appendix II ranking (Fuller and Swift, 1984). One specimen was captured in September and one in October. Xim- enez (1975) summarized the biology of this species. Oncifelis geoffroyi is com- mon in the province, especially in the Chaco and monte habitats; it is less common in forested habitats. Oreailurus jacobita (Cornalia) Felis jacobita Cornalia, Memorie Societa Italiana di Scienze Naturali e Museo Civico di Storia Na- turale di Milano, 1:3, 1865. Specimen Examined. — (1) Morro del Zarzo, 5000 m, 1 (MACN). Additional Records. — Laguna Huaca Huasi, Cumbres Calchaquies, approximately 4250 m; Campo de la Gallina, Serrania de Quilmes o El Cajon (Scrocchi and Halloy, 1986); Cumbres Calchaquies (Lucero, 1983). Measurements. — External measurements (one sex unknown): total length, 1120.0; tail, 480.0; hindfoot, 115.0. Habitat. — Oreailurus jacobita inhabits the high altitude arid and semiarid mountains found in the western part of the province. The Andean cat has a dis- tribution restricted to high elevations in Jujuy, Salta, Tucuman, and Catamarca provinces (Cabrera, 1961; Olrog and Lucero, 1981). Remarks. — The locality of the specimen examined was given as Sarzo, Acon- quija, Tucuman, by Cabrera (1961:205); the locality given above is that which appears on the specimen tag. In the Argentine Andes, O. jacobita was considered rare (International Union for the Conservation of Nature, 1990); currently its status is indeterminate (International Union for the Conservation of Nature, 1994). This species has an Appendix I listing in Argentina and is protected (Fuller and Swift, 1984). S. Halloy photographed this species at Laguna Huaca Huasi on a morning in 1980. Scrocchi and Halloy (1986) suggest that this species may be confused with Lynchailurus pajeros garleppi, which is similar in appearance, but which occurs along the eastern slopes of the Peruvian Andes. Little is known about the biology of O. jacobita. 1996 Mares et al. — TucumAn Mammals 131 Puma concolor (Linnaeus) Felis concolor Linnaeus, Mantissa Plantamm, 2:522, 1771. Specimens Examined. — (15) Burruyacu, 1 (CML); Cajon de la Ovejeria, Chorrillos, 5 (CML); Chi- cligasta, 1 (CML); Gobernador Garmendia, 1 (CML); Graneros, 1 (CML); Leales, 1 (CML); Los Puestos, 1 (CML); Rfo Nio, 1 (CML); San Miguel de Tucuman, 2 (CML); Trancas, 1 (CML). Additional Records. — Dique Escaba (M. L. Campbell, personal observation); El Manantial (Bur- meister, 1943); Laguna Huaca Huasi (N. Ovrusky, personal communication). Measurements. — Cranial measurements (one male, three sex unknown, two im- mature females): greatest length of skull, 156.0, 163.3 (154.0-181.0), 145.4, 121.9; condylobasal length, 148.0.^ 155.0 (143.0-174.0), 133.2, 108.3; least inter- orbital length, 29.6, 31.4 (28.2-35.0), 27.4, 21.9; zygomatic breadth, 109.3, 112.5 (102.4-126.6), 96.2, 79.3; breadth of braincase, 64.6, 66.9 (66.0-67.5), 64.9, 61.8; length of maxillary toothrow, 52.6, 54.5 (52.0-58.8), — , — ; palatal length, 69.5, 69.9 (64.9-76.8), — , — ; length of mandibular toothrow, 62.7, 60.6 (57.3- 66.3),™,—; greatest length of mandible, 113.0, 121.3 (110.0-135.0), 103.5, 81.1. Habitat. — Mountain lions are common in the Chacoan thorn scrub, the monte, and in the mountains of Tucuman Province. Remarks. — In Argentina, this species has an Appendix II ranking (Fuller and Swift, 1984). Currier (1983) summarized the biology of this species. Individuals have been collected in January {n = 1), February {n = 1), April {n = 1), June {n = 1), July {n = 5), September {n = 1), and November {n = 1). Specimens from Leales and Cajon de la Ovejeria are young individuals. Local residents of Dique Escaba indicated that this species occurred in the area (M. L. Campbell, personal observation). Pumas are as common in the Chaco as in the more moun- tainous regions of the province. Little is known about the biology of the species in Argentina. Cajal and Lopez (1987) found that in the San Guillermo Reserve, San Juan Province, guanacos and vicunas were an important component of the diet of pumas. Subfamily Pantherinae Panthera onca (Linnaeus) Felis onca Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, spe- cies cum characteribus, differentiis, synonymis, locis. Tenth edition. Laurentii Salvii, Stockholm, 1:42, 1758. Specimens Examined. — (0) None. Additional Record. — Tucuman, no specific locality given (De Moussy, 1864). Remarks. — This species is probably extinct in the province. Originally wide- spread in distribution, the range of this species has constricted due to hunting and deforestation (Roig, 1991). It still occurs in the neighboring provinces of Salta and Jujuy. Redford and Eisenberg (1992) did not extend the distribution of this species to Tucuman Province. Family Mustelidae Subfamily Lutrinae Lontra longicaudis (Olfers) Lutra longicaudis Olfers, in Eschwege, Journal von Brasilien, oder Vermischte nachrichten aus Bras- ilien, auf wissenschaftlichen reisen gesammett von W. C. von Echwege . . . Weimer, Im verlage des Gr. H. S. pr. Londes-industries-comptoirs, 15(2):233, 1818. Specimens Examined. — (1) Rio Colorado, Famailla, 1 (CML). 132 Annals of Carnegie Museum VOL. 65 Measurements. — External measurements (one female): total length, 1030.0; tail, 410.0; hindfoot, 60.6. Cranial measurements (one female): greatest length of skull, 106.7; condylobasal length, 106.2; least interorbital length, 20.8; zygomatic breadth, 67.9; breadth of braincase, 52.6; length of maxillary toothrow, 28.5; palatal length, 48.0; length of mandibular toothrow, 33.9; greatest length of man- dible, 68.4. Habitat. — This otter prefers lagunas, estuaries, and rivers (Olrog and Lucero, 1981; Lucero, 1983). Remarks. — Otters are rare in the province. Cabrera (1957) and Olrog and Lu- cero (1981) listed this species as L. platensis. Although found widely in central and northern Argentina, the distribution is becoming fragmented. The high value placed on the fur encourages illegal hunting. This species is considered in the Appendix I category by Argentina and is protected (Fuller and Swift, 1984). A conservation action plan and status report is available for the otter (Foster-Turley et al., 1990). Subfamily Mephitinae Conepatus chinga (Molina) Viverra chinga Molina, Saggio sulla storia naturale del Chili. S. Tommaso d’ Aquino, Bologna, p. 288, 1782. Specimens Examined. — (5) Estancia El Cavao, 1 (CML); La Angostura, 1 (CML); La Cuesta, 1 (CML); Quebrada de las Burras, 1 (CML); Rfo Las Tacanas, 1 (CML). Additional Record. — El Infiemillo (R. M. Barquez, personal observation); El Infiemillo, about 3500 m (R. A. Ojeda, personal observation). Measurements. — External measurements (one female): total length, 660.0; tail, 250.0; hindfoot, 60.0. Cranial measurements (three females, two sex unknown, unless noted otherwise): greatest length of skull, 64.0 (59.1-67.0), 62.8, 64.3; condylobasal length, 61.6 (56.1-64.5), 61.1, 63.5; least interorbital length, 17.1 (16.4-17.7), 16.8, 15.3; zygomatic breadth (one female, one sex unknown), 40.2, 37.6; breadth of braincase, 31.7 (31.0-32.1), 29.6, 27.0; length of maxillary tooth- row, 20.0 (19.0-20.5), 19.9, 19.1; palatal length (one female), 28.2; length of mandibular toothrow, 22.4 (19.5-23.9), 22.4, 21.9; greatest length of mandible, 40.6 (37.1-43.6), 40.1, 39.8. Habitat. — Conepatus chinga prefers savannas, mountains, and forests as well as the arid, shrubby areas of western Tucuman Province up to 3500 m (Cabrera, 1957; Lucero, 1983). Remarks. — The extreme variation in coloration and size and stripe morphology shown by Conepatus has led to the description of numerous species. A systematic revision is needed. The few records that exist for the province do not permit us to make a detailed assessment of the color variation. In most systematic works, C. rex, which was described by Thomas (1898), is included in chinga. The in- dividual observed at El Infiemillo was completely white dorsally and had a white tail, a description that corresponds to that of C. rex (Olrog and Lucero, 1981). The distribution of C. rex would include the mountainous areas of the province. Subfamily Mustelinae Eira barbara (Linnaeus) Mustela barbara Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth edition. Laurentii Salvii, Stock- holm, 1:46, 1758. 1996 Mares et al. — TucumAn Mammals 133 Specimens Examined. — (5) Chicligasta, 1 (CML); Cumbres de San Pablo, 1 (BMNH); Monteros, 1 (MACN); Tucuman, no specific locality, 1 (MACN); Yerba Buena, 1 (CML). Measurements. — External measurements (one male): total length, 1030.0; tail, 420.0; hindfoot, 78.0; ear, 35.0. Cranial measurements (one male): greatest length of skull, 118.0; condylobasal length, 104.0; least interorbital length, 25.6; zygo- matic breadth, 71.6; breadth of braincase, 46.5; length of maxillary toothrow, 30.4; palatal length, 59.3; length of mandibular toothrow, 40.3; greatest length of mandible, 77.2. Habitat. — Eiras are found in northern Argentina east of the Andes and as far south as southern Santiago del Estero Province (Olrog and Lucero, 1981). Lucero (1983) notes that this species is found in savannas, forests, and woodlands. Remarks. —-IndiNidudil specimens have been collected in July (n = 1) and Au- gust (n = 1). Eiras are quite common in the province, although records of them are scarce. An individual was seen in dense forest above 1800 m along Highway 65 in Catamarca Province near the Catamarca-Tucumm provincial border (M. A. Mares, personal observation). Galictis cuja (Molina) Mustela cuja Molina, Sagio sulla storia naturale del Chili, S. Tommaso d’ Aquino, Bologna, p. 291, 1782. Specimens Examined. — (22) Concepcion, 15 (1 CML, 2 MACN, 12 BMNH); El Infiernillo, 1 (CML); El Portezuelo, 1 (CML); El Timbo, 2 (CML); La Higuera, 1 (CML); La Reduccion, 1 (CML); Tafi Viejo, 1 (CML). Additional Record. — Cumbres Calchaquies (E. Dominguez, S. Halloy, and E. Teran, personal com- munication). Measurements.— ~~¥.yAQ.tndl measurements (five males, six females, unless noted otherwise): total length (four males, six females), 512.5 (470.0-550.0), 480.3 (360.0-560.0); tail, 144.0 (120.0-170.0), 139.3 (100.0-160.0); hindfoot, 57.1 (50.0-70.0), 50.0 (45.0-55.0); ear, 23.5 (20.5-25.0), 20.5 (16.0-29.0). Cranial measurements (six males, unless noted otherwise): greatest length of skull, 70.0 (58.8-74.4); condylobasal length, 69.9 (58.5-74.9); least interorbital length, 16.6 (14.6-17.9); zygomatic breadth, 42.3 (36.1-44.5); breadth of braincase, 34.3 (25.3-37.8); length of maxillary toothrow, 19.3 (11.8-22.0); palatal length (five males), 32.2 (27.4-34.7); length of mandibular toothrow, 23.9 (19.8-25.4); great- est length of mandible, 42.8 (36.5-45.6). Habitat. — Galictis is found in nearly all of Argentina except Santa Cruz Prov- ince and Tierra del Fuego (Olrog and Lucero, 1981). This mustelid prefers moun- tains, savannas, and arid scrubby regions; it is found up to 3500 m (Olrog and Lucero, 1981; Lucero, 1983). Remarks. — Individuals have been captured in May {n = 1), June {n = 2), July {n = 1), August {n = 2), September {n = 1), and October (n = 3). Lyncodon patagonicus (de Blainville) Mustela patagonica de Blainville, Osteographie, on description iconographique comparee du squelette et du systeme dentaire des mammiferes recents et fossiles . . . par H.-M. Ducrotay de Blainville, . . . precede d’une etude sur la vie et les travaux de M. de Blainville, par M. P. Nicard. J.-B. Bailliere et fils, 1839-1864, 4 volumes. Mammalia, partie 10 (Viverra):!, 1842. Specimen Examined. — (1) Banda del Rio Saif, 1 (CML). Additional Records. — Amaicha del Valle; Colalao del Valle (Olrog, 1976); El Timbo (Olrog, 1958). Measurements. — External measurements (one female): total length, 330.0; tail. 134 Annals of Carnegie Museum VOL. 65 190.0; hindfoot, 29.5; ear, 15.0. Cranial measurements (one female): greatest length of skull, 45.7; condylobasal length, 46.0; least interorbital length, 10.9; zygomatic breadth, 24.3; breadth of braincase, 21.3; length of mandibular tooth- row, 15.8; greatest length of mandible, 27.1. Habitat. — Lyncodon is widely distributed in Argentina from Tucuman Province in the north to Cordoba Province in the east and southward to Santa Cruz Prov- ince. This species is found in arid, shrubby terrain in the lowlands, and in the mountains up to 2000 m (Olrog and Lucero, 1981; Lucero, 1983). Remarks. — Olrog (1958) noted that this species was not rare in the province and that it was common in the Valle Calchaquies up to 2000 m. We have not been able to verify the existence of the specimens listed in the additional records. Olrog (1958, 1976) noted that these specimens from Amaicha del Valle and Col- alao del Valle were collected in November; those from El Timbo were collected in May. The specimen from Banda del Rio SaK was collected in June. Family Procyonidae Subfamily Procyoninae Nasua nasua (Linnaeus) Viverra nasua Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Volume 1. Regnum Animale. 12th edi- tion. Laurentii Salvii, Stockholm, 1:64, 1766. Specimens Examined. — (3) Cerro San Javier, 1 (CML); Tafi Viejo, 1 (CML); Trancas, 1 (CML). Measurements. — Cranial measurements (one subadult sex unknown): greatest length of skull, 104.0; least interorbital length, 23.2; zygomatic breadth, 53.9; breadth of braincase, 44.3; length of maxillary toothrow, 34.6; palatal length, 61.6; length of mandibular toothrow, 36.8; greatest length of mandible, 71.0. Remarks. — The specimen from Cerro San Javier is a young animal. This species is not commonly observed in the province, which constitutes its southern distri- butional limits. Two populations are found in Argentina. The first occurs in the provinces of Chaco, Corrientes, Formosa, Misiones, and Santa Fe. The second, found in northwestern Argentina, occurs in Jujuy, Salta, and Tucuman. The Gran Chaco appears to separate the two. Mares et al. (1981, 1989) comment on the biology of this species in the neighboring province of Salta. Redford and Eisen- berg (1992) did not extend the distribution of this species to Tucuman Province. Procyon cancrivorus (G. Cuvier) Ursus cancrivorus G. Cuvier, Tableau elementaire de I’histoire naturelle des Animaux, Paris, p. 113, 1798. Specimens Examined. — (5) Chicligasta, 1 (CML); Concepcion, 3 (1 BMNH, 1 MACN, 1 CML); Los Vazquez, 1 (BMNH). Additional Records. — Dique Escaba (M. L. Campbell, personal observation); 2 km below La He- ladera along Hwy 307, 3,500 ft. (J. K. Braun, personal observation; M. L. Campbell, personal obser- vation); Piedra Tendida (R. M. Barquez, personal observation). Measurements. — External measurements (one male, one female): total length, 950.0, 880.0; tail, 330.0, 330.0; hindfoot, 135.0, — ; ear, 60.0, — . Cranial mea- surements (one male, one female): greatest length of skull, 118.4, 115.0; condy- lobasal length, 125.5, 121.0; least interorbital length, 28.8, 26.9; zygomatic breadth, 92.2, 83.0; breadth of braincase, 61.2, 57.1; length of maxillary toothrow, 49.7, 48.7; palatal length, 62.5, 63.0; length of mandibular toothrow, 55.7, 54.3; greatest length of mandible, 96.9, 93.1. 1996 Mares et al. — TucumAn Mammals 135 Habitat. — Crab-eating raccoons are, in general, found in savannas, forests, and woodlands, usually along water courses (Lucero, 1983). Procyon cancrivorus is found in all of northern Argentina east of the Andes as far south as San Luis Province (Olrog and Lucero, 1981); Cabrera (1957) had earlier placed the south- ern distributional limits in northern Catamarca Province. The vegetation at La Heladera was Yungas forest dominated by laurel and Mirtaceae (M. L, Campbell, personal observation). The scat, found in the cavity of a very large tree, contained freshwater crabs and large land snails (J. K. Braun, personal observation; M. L. Campbell, personal observation). Remarks. — -Single specimens have been captured in September, October, and December. The record of this species at Dique Escaba is based on information obtained from local residents (M. L. Campbell, personal observation). Order Perissodactyla Family Tapiridae Tapirus terrestris (Linnaeus) Hippopotamus terrestris Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth edition. Laurentii Salvii, Stockholm, 1:74, 1759. Specimens Examined. — (3) Las Pavas, 3 (CML). Additional Record. — Departamento Famailla (Lucero, 1983). Remarks. — Lucero (1983) noted that the tapir has been extinct in the province since 1940, although an isolated population may still exist is some of the heavily forested areas. This species is listed as an Appendix II species in Argentina and is protected (Fuller and Swift, 1984). Historically, tapirs were found in the Yungas forest and gallery forests of the Chacoan thorn scrub, always near water. The distribution has contracted due to hunting pressure and habitat modification (Roig, 1991). Tapirs are found in the provinces of Misiones, Corrientes, Formosa, Chaco, Salta, and Jujuy (Olrog and Lucero, 1981). Order Artiodactyla Family Tayassuidae Catagonus wagneri (Rusconi) Platygonus carlesi wagneri, Rusconi, Anales del Museo Nacional de Historia Natural “Bernardino Rivadavia,” 36:231.1930. Specimens Examined.— -{Q) None. Additional Records. — No specimens of this species have been collected in the province but Olrog, Ojeda, and Barquez (personal observation) have seen signs of its presence in the northeastern part of the province near the specimen locality (in Santiago del Estero) cited by Olrog et al. (1976). Lucero (1983) also suggested that this peccary may occur in the province. This species was found by Mares in neighboring Santiago del Estero, just a few kilometers from the Tucuman border. Remarks. — The Chacoan peccary is listed as vulnerable in Argentina (Inter- national Union for the Conservation of Nature, 1990). It is endemic to the Gran Chaco of Argentina, Bolivia, and Paraguay. The biology of this species is sum- marized by Mayer and Wetzel (1986). Pecari tajacu (Linnaeus) Sus tajacu Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth edition. Laurentii Salvii, Stockholm, 1: 50, 1758. 136 Annals of Carnegie Museum VOL. 65 Specimens Examined. — (4) Cerro del Campo, 1800 m, 3 (CML); Departamento Chicligasta, 1 (CML). Additional Records. — Aguas Chiquitas (P. Capllonch, personal communication); Dique Escaba (M. L. Campbell, personal observation); El Nogalar, = Rio los Sosa, km 43.0, Ruta 307 (S. Ojeda, personal communication); Piedra Tendida, 12 km WNW Burruyacu, along Rio Cajon, 2,500 ft. (M. L. Camp- bell, personal observation). Measurements. — Cranial measurements (one female, one subadult sex un- known): greatest length of skull, 450.0, 203.0; condylobasal length, 215.0, 172.0; least interorbital length, 54.0, 44.5; zygomatic breadth, 117.8, 85.5; breadth of braincase, 53.9, 52.9; length of maxillary toothrow, — , 46.2; palatal length, 142.0, 115.0; length of mandibular toothrow, — , 53.0; greatest length of mandible, — , 131.0. Habitat. — The distribution of the collared peccary includes north-central Ar- gentina east of the Andes and the Yungas forest (Olrog and Lucero, 1981). The southernmost distributional limits are San Luis and northeastern Mendoza prov- inces. This species is found in savannas, woodlands, and forests of thick vege- tation (Olrog and Lucero, 1981; Lucero, 1983). Meyer and Weyrauch (1966) reported that this peccary occurred in subtropical humid forest. Locals interviewed at the Piedra Tendida locality indicated that this peccary is found in the area, but is heavily hunted; it has retreated into the surrounding hills (M. L. Campbell, personal communication). The record of this species is based on information ob- tained from locals at Dique Escaba who indicated that this species is present in the area (M. L. Campbell, personal communication). Remarks. — The peccary captured at Cerro del Campo is a young individual. Specimens were collected in April (n = 1) and September (n = 1). Although this species is common in the province, its populations have been reduced by hunting and habitat conversion. Tayassu pecari (Link) Sus pecari Link, Beitrage zur Naturgeschichte, 2:104, 1795. Specimens Examined. — (0) None. Remarks. — Information on the biology of this species was presented by Mayer and Wetzel (1987). This species tends to inhabit the forested regions of northern Argentina (Olrog and Lucero, 1981). Its presence in the province is unknown, although Meyer and Weyrauch (1966) reported it as a component of the fauna of the subtropical humid forest. Family Camelidae Lama guanicoe (Muller) Camelus guanicoe Muller, Mit einer ausfiihrlichen erklarung ausgefertiget. Des ritters Carl von Linne . . . Vollstandigen natursystems supplements und register-ban iiber aler sechs theile oder classen des thierreichs. G. N. Raspe, Nurnberg, p. 50, 1776. Specimen Examined. — (1) Nevados del Aconquija, Glacial del Cochuna, 4550 m, 1 (CML). Additional Records. — Sierras del Cajon; Cumbres Calchaqufes; Cumbres de Santa Barbara; Sierra del Aconquija (Cajal, 1983); Sierras del Aconquija (De Moussy, 1864); Cumbres Calchaquies, Alto de Munoz (E. Dominguez, S. Halloy, and E. Teran, personal communication). Measurements. — Cranial measurements (one sex unknown): greatest length of skull, 280.0; condylobasal length, 267.0; least interorbital length, 101.3; zygo- matic breadth, 123.7; breadth of braincase, 69.7; length of maxillary toothrow, 62.6; palatal length, 143.5. 1996 Mares et al, — Tucuman Mammals 137 Habitat. — -This species is distributed from southern Argentina (Santa Cruz Province), where they are relatively abundant, northward to Jujuy Province, In the northern parts of the distribution, guaeacos are much less abundant. In Tu- cuman, guanacos inhabit mountains up to 4000 m (Olrog and Lucero, 1981; Lu- cero, 1983). Remarks.- — -Guaeacos are listed as an Appendix II species in Argentina (Fuller and Swift, 1984). The biology of the guaeaco has been summarized in Cajal (1991) and Torres (1992). Guanacos are present in La Florida Natural Park, but there are no density estimates. Cajal (1983) estimated 1000 guaeacos were present in the Cumbres de Santa Barbara. Aconquija has been proposed as a protected area. This species was originally distributed throughout most of the country, in- cluding Chaco, Santa Fe, and Buenos Aires provinces. The range has constricted due to habitat modification and hunting pressure. Cajal (1983) and Cajal and Amaya (1985) reviewed diverse aspects of the biology and history of the guanaco. Vicugna vicugna (Molina) Camelius (sic) vicugna Molina, Saggio sulla storia eaturale del Chili. S. Tommaso d’Aquino, Bologna, 1782:313. Specimens Examined. — (0) None. Additional Records.— Si&[rd& del Aconquija, Departamento Tafi del Valle (De Moussy, 1864). Remarks.— are unable to verify the precision of the information given by De Moussy (1864). Vicunas are distributed in the high Andean regions from Jujuy to northern Mendoza (Olrog and Lucero, 1981), although Cajal and Puig (1992) noted that the distribution extended only to northern San Juan Province. Bedford and Eisenberg (1992) did not extend the distribution of this species to Tucuman Province. Family Cervidae Subfamily Odocoileinae Hippocamelus antisensis (d’Orbigey) Cervus antisensis d'Orbigny, Aonales Museum d’Histoire Naturelle, Paris, 3:91, 1834. Specimen Examined.— {!) Cumbres del Aconquija, 1 (CML). Additional Record. — Laguna Huaca Huasi (S, Halloy, personal communication). Measurements.- — ^Cranial measurements (one male): greatest length of skull, 255.0; least interorbital length, 73.5; breadth of braincase, 71.0; length of max- illary toothrow, 67.1; palatal length, 116.2; length of mandibular toothrow, 74.7; greatest length of mandible, 227.0. Habitat.— The huemul is found in the Andes of Salta, Jujuy, Tucuman, Cata- marca, and La Rioja (Olrog and Lucero, 1981). This species inhabits the rocky steppes between 3000-4000 m (Olrog and Lucero, 1981; Lucero, 1983). Remarks.— Thi^ deer is considered vulnerable in Argentina (International Union for the Conservation of Nature, 1990, 1994) and is ranked as an Appendix I species (Fuller and Swift, 1984) by Argentina and is protected. The biology was summarized by Moreno (1993). Huemuls were intensively hunted; this species is in danger of being extirpated from the province, although several populations still occur in the central highlands. 138 Annals of Carnegie Museum VOL. 65 Mazama americana (Erxleben) Moschus americanus Erxleben, Systema regni animalis per classes, ordines, genera, species, varietates, cum synonymia et historia animalium. Classis. I. Mammalia. Weygandianis, Lipsiae, 1:324, 1777. Specimens Examined. — (0) None. Additional Records. — On two separate occasions Mares observed these deer standing on the road to Tafi del Valle at the approximate altitude of Piedras Coloradas; Dique Escaba (M. L. Campbell, personal communication); 0.5 km N of Las Higuerillas on Hwy 308, 3,000 ft. (M. L. Campbell, personal communication); Tucuman, no specific locality (Meyer and Weyrauch, 1966). Habitat. — The distribution of M. americana includes the forested regions of Jujuy, Salta, Tucuman, Formosa, Chaco, Santa Fe, Corrientes, and Misiones (Ol- rog and Lucero, 1981). In Tucuman, this species is found in transitional and humid forests; the southernmost limits of its distribution are along the border with Ca~ tamarca Province (Olrog and Lucero, 1981; Lucero, 1983). Remarks. — The biology of this species has been summarized by Moreno (1993). It is either rare or difficult to observe in its preferred habitat of dense, humid forest. Meyer and Weyrauch (1966) report this deer in the subtropical humid forest of Tucuman. At Dique Escaba, locals indicated that M. americana occurred in the area; four piles of scat were found at the Las Higuerillas locality (M. L, Campbell, personal communication). Mazama gouazoupira (G. Fischer) Cervus gouazoupira G. Fischer, Zoognosia, 3:465, 1814. Specimens Examined. — (4) Chicligasta, 3 (CML); Leales, 1 (CML). Additional Records. — Near the campground near Cochuna (R. A. Ojeda, personal observation); El Cadillal (personal observation); Piedra Tendida, 12 km WNW Burruyacu, along Rio Cajon, 2,500 ft. (M. L. Campbell, personal communication). Measurements. — Cranial measurements (two females): greatest length of skull, 159.5, 169.0; condylobasal length, 147.8, 164.0; least interorbital length, 39.8, 39.9; zygomatic breadth, 72.0, 74.0; breadth of braincase, 55.6, 54.1; length of maxillary toothrow, 40.6, 52.6; palatal length, 92.6, 106.5; length of mandibular toothrow, 45.4, 61.5; greatest length of mandible, 132.9, 143.0. Habitat. — Mazama gouazoupira is more widely distributed than M. americana, occurring from east of the Andes to the northeastern provinces, southward to San Luis Province (Olrog and Lucero, 1981). Lucero (1983) notes that this species is abundant in the open savannas, woodlands, thickets, and mountains up to 2000 m along the border with the province of Catamarca. This deer seems to prefer the Chacoan habitat found in the eastern parts of the province. Remarks. — Moreno (1993) summarized the biology of this species. Scat of this species was found at Piedra Tendida (transitional forest) and hunters indicated that it was a common species in the area (M. L. Campbell, personal communi- cation). Although there are few records and little is known of its biology, this species is more abundant than M. americana. Ozotoceros bezoarticus (Linnaeus) Cervus bezoarticus Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth edition. Laurentii Salvii, Stockholm, 1:67, 1758. Remarks. — An expedition report dated 1825-1826 stated that this species once occurred in large numbers in the region between Vipos and Trancas (Roig, 1991). 1996 Mares et al. — Tucuman Mammals 139 Jackson (1987) and Moreno (1993) have summarized the biology of this species. It is no longer found in the province. Redford and Eisenberg (1992) did not extend the distribution of this species to Tucuman Province. Order Lagomorpha Family Leporidae Sylvilagus brasiliensis (Linnaeus) Lepus brasiliensis Linnaeus, Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth edition. Laurentii Salvii, Stock- holm, 1:58, 1758. Specimens Examined. — (8) Campo de las Chunas, San Pedro de Colalao, 1 (CML); Cerro del Campo, 800 m, 1 (BMNH); El Portezuelo, San Pedro de Colalao, 2 (CML); El Taficillo, 3 (CML); Las Mesadas, 1 (CML). Additional Records. — El Cadillal (R. M. Barquez, personal observation); Playa Larga (R. M. Bar- quez, personal observation). Measurements. — External measurements (two males): total length, 390.0, 348.0; tail,“=, 19.0; hindfoot, 80.0, 74,0; ear, 75.0, 60.5. Cranial measurements (two males): greatest length of skull, 67.4, 70.8; condylobasal length, 60.5, 58.0; least interorbital length, 14.2, 17.2; zygomatic breadth, 32.7, 33.5; breadth of braincase, 25,3, 24.6; length of maxillary toothrow, 14.0, 12.6; palatal length, 27.8, 31.0; length of mandibular toothrow, 14.0, 12.7; greatest length of mandible, 46.8, 62.5. Habitat.-— Lucero (1983) gives the habitat as transitional forest, open wood- lands, and savannas, in mountains up to 2000 m. Remarks.— This herbivorous species is generally nocturnal and solitary. Discussion Our data provide a great deal of information on a rich fauna of mammals that has not been well studied to date. Certainly the many specimens will prove to be useful to systematists in the years to come, as will the tissues and chromosomes. Moreover, to date there has been little quantitative data on habitat selection, dis- tribution, and reproduction. Regarding the latter, there have been few studies to offer comparative data on patterns of reproduction for most groups of mammals in South America (Hamlett, 1992; Harder, 1992; Lacher, 1992). South American bats have also received little attention from the standpoint of their reproductive patterns; however, Myers (1977) and Myers and Wetzel (1983) provided a great deal of information on bats of the Chaco of Paraguay, located not far from Tuc- uman Province. Pearson and Pearson (1989) studied reproduction in three species of vespertilionid bats in southern Argentina, located much farther away from Tucuman Province than Paraguay, geographically speaking, although still within Argentina. In the Chaco, it was shown that reproductive activity for four species of ves- pertilionid bats began as early as the winter month of May, when spermatozoa were found in the reproductive tracts of females (Myers, 1977). Reproductively active Chacoan bats from a wide variety of taxa were found mainly from August through October (Myers and Wetzel, 1983). In southernmost Argentina, the bats that were studied were pregnant in the late spring and summer months of October through December (Pearson and Pearson, 1989). We did not do histological or other microscopic examinations of specimens, although fluid-preserved specimens are available for laboratory analyses. In examining the data for all the bats col- 140 Annals of Carnegie Museum VOL. 65 lected, we found that almost all gross reproductive activity (as determined by embryos, enlarged testes, and lactation) occurred during the late spring and sum- mer months of September through January. Only a few individuals of any species showed any evidence of reproductive activity during the winter (May-July) (e.g., Sturnira erythromos, Artibeus planirostris, Lasiurus ego), and none was pregnant or lac taring. These results are in accord with the studies conducted in Paraguay and in southern Argentina. Where winters are harsh in Tucuman (e.g., the high mountains), we would expect that bats will show the strict limitations on month of parturition described by Pearson and Pearson (1989). Where conditions are somewhat less stringent in winter (e.g., the Chaco or moist subtropical forest), we would expect bats to show occasionally some activity in colder months. The bats of Tucuman Province are clearly responding to the area in a manner that suggests that parturition is keyed to the warmer months of the year, a pattern that may obtain until truly tropical conditions are encountered, when much more com- plex patterns of reproduction are possible (Wilson, 1979). Only a larger data base will clarify these patterns over a broad geographic and climatologic scale, as well as under different conditions of allopatry, sympatry, and seasonality of food re- sources. Our work documents that 68 nonrodent species of mammals occur, or have recently occurred, in Tucuman Province. Additionally, we have suggested that Priodontes maximus, the giant armadillo, may still be found in the Chacoan scrub- lands of extreme northeastern Tucuman. The species is known to occur in the neighboring provinces of Salta and Santiago del Estero. Although there are likely several provinces in Argentina that support a similar number of mammal species as Tucuman, these provinces are much larger (e.g., Salta, which is seven times larger than Tucuman) and have a portion of their territory located within the tropics. Because of its small size, long history of extensive human habitation, economic and agricultural development, and limited size of macrohabitats, Tucuman can be considered to be a model system for studying the effects of human population density, economic development, and modern agricultural techniques on the natural environment. With its moist subtropical forest, rich montane woodlands, extensive lowland Chacoan thorn scrub, high-elevation and low-elevation deserts, and gla- cial watersheds, the province has experienced, and is experiencing, the challenges of rapid development, while attempting to control in some manner the effects of such development on the natural environment. Certainly the history of economic development in Tucuman has had a pronounced influence on the natural habitats of the province and on its mammal fauna. From the conservation standpoint, some of the most important species are non- rodents. Indeed, Argentina has only one threatened or endangered rodent (the chinchilla. Chinchilla lanigera), although some additional potential candidates for protection exist. Still, all other protected, threatened, and endangered mammals are nonrodents and nonbats. Although Tucuman has supported European human populations for almost four-and-a-half centuries, and native populations for mil- lennia, and even though it is presently Argentina’s most densely settled province, there have been relatively few extirpations of mammals. Six species have likely been extirpated from the province during historic times: the jaguar (P. onca), tapir (Tapirus), pampas deer (Ozotoceros), vicuna {Vicugna), black-capped capuchin monkey {Cebus), and giant anteater {Myrmecophaga). The first four were eliminated due to hunting, whereas the giant anteater and the 1996 Mares et al. — Tucuman Mammals 141 monkey may have occurred in low numbers in the dry Chaco (anteater) or low montane forests (monkey) and been eliminated due both to hunting pressures and habitat modification. Two other species, the three-banded armadillo (Tolypeutes) and the otter (Lontra) are in very low numbers in Tucuman, their populations reduced by hunting and habitat modification. Both were likely common in the past. There are four other species in Tucuman that are rare, and hunting pressures almost certainly have had an adverse affect on their numbers, but most of these were probably rare in any case (e.g., Cabassous, L. pajeros, O. jacobita, Cata~ gonus). Thus, when viewed as a fauna of more than 100 species, the mammals of Tucuman would seem to have been reduced by “only” six species. But this reduction in the number of species supported in the province has not occurred in a haphazard manner, with species disappearance occurring across taxonomic cat- egories. Rather it is the larger mammals that have been most strongly affected. We will deal with the rodents in a subsequent paper, but few rodents appear to have disappeared from the province due to human activities. Larger mammals are more susceptible to human depredation, serving as food for humans in some cases, competitors in others, and dangerous predators in at least one case. Among Tucuman’s 68 nonrodent species, fully 12 species, or 19% of the non- rodent fauna, have either been eliminated, or are in danger of being extirpated from the province. Among these are the largest-bodied species of mammals found in the province, as well as several important carnivores. If we also exclude bats from consideration, we find that of the 39 nonbat and nonrodent species that occurred in Tucuman Province within the last 200 years, 12 of 39, or 31%, have been extirpated or are at very low population levels, thus almost a third of the larger mammals are no longer found in the province. This is a significant loss of economically and ecologically important species, as well as a loss of species that were spectacular inhabitants of the rich habitats of Tucuman Province. We suggest that it is not too late to act to recover a significant proportion of the species that have been lost. There is little hope of bringing back a controversial species such as the jaguar, but vicunas could be reintroduced to the high mountains of the province and tapirs to the forests. Monkeys could also be brought back to the forest they inhabited only a relatively short time ago. Similarly, Ozotoceros would not appear to present special difficulties for reintroduction. By setting aside Chacoan thorn scrub reserves, the giant armadillo, giant anteater, Cabassous, and Tolypeutes could also be either protected or reintroduced. The recent history of the province involves failed efforts (fortunately) to intro- duce exotic species such as red deer {Cervus elephas) to the alder forests and montane grasslands. It would take much less effort, and be more logical, to re- cover the fauna of large mammals that has disappeared from Tucuman, thereby impoverishing the fauna of the province and the lives of the people who live there. The ecological situation of Tucuman, while serious, is far from hopeless. If provincial leaders can find the will to act, and if the citizenry can be educated to appreciate its own rich mammal fauna rather than the fauna of Europe, it is highly likely that the large-mammal fauna of Tucuman can be returned to a good approximation of the fauna that occurred in the province at the time of the col- onization of South America. In this way, Tucuman could serve not as an indicator of how important species disappear as a result of human activities, but rather how major faunal elements can be preserved in managed reserves and, with protection, in the natural habitats where they once roamed. 142 Annals of Carnegie Museum VOL. 65 Acknowledgments Many individuals have participated in field research. Most recently, the following have assisted in collecting and preparing specimens: S. Adams, M. L. Campbell, P. Capllonch, R. L. Humphrey, M. Guevarra, E. Pescetti, and S. Peurach. Funding for field research was provided by a grant to M. A. Mares from the National Science Foundation (BSR-8906665), among other sources. Additional support was provided by the Oklahoma Museum of Natural History and the Department of Zoology, University of Oklahoma. C. Kacmarcik produced the figures. We appreciate the many helpful suggestions of P. Myers, B. Patterson, and an anonymous reviewer, that resulted in a much improved manuscript. 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Annals and Magazine of Natural History, series 1, 7:277-283. . 1902. On mammals collected at Cruz del Eje, central Cordoba, by Mr. P. O. Simon. Annals and Magazine of Natural History, series 7, 9:237-245. . 1907. On Neotropical mammals of the genera Callicebus, Reithrodontomys, Ctenomys, Dasypus, and Marmosa. Annals and Magazine of Natural History, series 7, 20:161-168. . 1926. The Spedan Lewis South American exploration. III. On mammals collected by Sr. Budin in the province of Tucuman. Annals and Magazine of Natural History, series 9, 17:602- 609. Torres, H., ed. 1992. South American Camelids. An Action Plan for Their Conservation. Interna- tional Union for the Conservation of Nature, Gland, Switzerland, 58 pp. Vervoorst, F. 1982. Noroeste. Pp. 9-24, in Conservacion de la Vegetacion Natural en la Republica Argentina. Simposio XVIII Jornadas de Botanica, Fundacion Miguel Lillo, Tucuman, Argentina. Wetzel, R. M. 1980. A revision of the naked-tailed armadillos, genus Cabassous McMurtrie. Annals of Carnegie Museum, 49:323-357. . 1982. Systematics, distribution, ecology, and conservation of South American edentates. Pp. 345-376, in Mammalian Biology in South America (M. A. Mares and H. H. Genoways, eds.). Special Publications Series, Pymatuning Laboratory of Ecology, University of Pittsburgh, 6:1- 539. . 1985. Taxonomy and distribution of armadillos, Dasypodidae. Pp. 23-46, in The Evolution and Ecology of Armadillos, Sloths, and Vermilinguas (G. G. Montgomery, ed.). Smithsonian Institution Press, Washington, D.C., 451 pp. Williams, D. F. 1978. Taxonomic and karyologic comments on small brown bats, genus Eptesicus, from South America. Annals of Carnegie Museum, 47:361-383. Wilson, D. E. 1979. Reproductive patterns. Pp. 317-378, in Biology of Bats of the New World Family Phyllostomatidae. Part III (R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds.). Special Publications, The Museum, Texas Tech University, 16:1-441. Wilson, D. E,, and D. M. Reeder. 1993. Mammal Species of the World: A Taxonomic and Geo- graphic Reference. Smithsonian Institution Press, Washington, D.C., 1206 pp. XiMENEZ, A. 1975. Felis geoffroyi. Mammalian Species, 54:1-4. Yepes, j. 1929. Notas sobre la distribucion geografica del pichi ciego menor (Chlamyphorus trun~ cams) y pichi lloron {Chaetophractus vellerosus). Physis, 9:439-446. . 1935. Las especies Argentinas del genero Cabassous (Dasypodidae). Physis, 11:438-444. Gazetteer Localities are listed in both alphabetical and numerical order. Departments are given in parentheses. Numbers in parentheses correspond to numbered localities in Figure 8. Acheral (Monteros)— 27°07' 65°27' (109). Aconquija (Chicligasta) — 27°13' 66°08' (99). Aconquija, 3000 m (Chicligasta) — 27°13' 66°08' (99). Agua Colorada (Burruyacu) — 26°26' 64°53' (72). Agua Rosada (Trancas) — 26°22' 65°26' (15). Agua Rosada, San Pedro de Colalao (Trancas) — 26°22' 65°26' (15). Aguas Chiquitas (Burruyacu) — 26°37' 65°12' (35). Aguas Chiquitas, El Cadillal, 800m (Burruyacu) — 26°37' 65°12' (35). Aguas Chiquitas, Sierras de Medina, 800m (Burruyacu) — 26°37' 65° 12' (35). Aguas Chiquitas, 25 km NE San Miguel de Tucuman, 800 m (Burruyacu) — 26°37' 65°12' (35). Alderetes (Cruz Alta)— 26°49' 65°08' (63). Amaicha del Valle (Tafi del Valle)— 26°36' 65°55' (1). Amaicha del Valle, 2000 m (Tafi del Valle)— 26°36' 65°55' (1). Anta Mapu (Burruyacu) — 26°31' 64°43' (74). Atahona (Simoca)— 27°25' 65°17' (83). 146 Annals of Carnegie Museum VOL. 65 Banda del Rio Sail (Cruz Alta)— 26°50' 65° 10' (62). Biological Reserve at Horco Molle, 2,400 ft. (Yerba Buena) — 26°45' 65°2r (45). Biological Reserve at Horco Molle, near residencia, 2,400 ft. (Yerba Buena) — 26°45' 65°21' (45). Burruyacu (Burruyacu) — 26°30' 64°45' (73). Burruyacu, 1300 m (Burruyacu) — 26°30' 64°45' (73). Cafayate, 45 km S, along Hwy 40 (Tafi del Valle)— 26°20' 65°58' (4). Cajon de la Ovejeria, Chorrillos (Burruyacu) — 26°2r 64°55' (70). Campo de la Gallina, Serrania de Quilmes o El Cajon (Tafi del Valle) — (not located). Capital, San Miguel de Tucuman (Capital) — 26°50' 65°13' (50). Casa de Piedra, Rio Los Sosa (Monteros) — 27°03' 65°37' (106). Casa de Piedra, Ruta 307, km 23.9, 850m (Monteros) — 27°03' 65°37' (106). Caspichango (Monteros) — 27°04' 65°30' (112). Cerro de Raco, Trancas (Tafi Viejo) — 26°43' 65°30' (56). Cerro de Tafi Viejo, 2400 m (Tafi Viejo) — 26°42' 65°17' and 26°44' 65°16' (between 38 and 44). Cerro del Campo (Burruyacu) — 26°35' 64°57' (68). Cerro del Campo, 800 m (Burruyacu) — 26°35' 64°57' (68). Cerro del Campo, 1800 m (Burruyacu) — 26°35' 64°57' (68). Cerro San Javier (Yerba Buena) — 26°47' 65°24' (42). Chicligasta (Simoca) — 27°28' 65°07' (84). Ciudad Capital (Capital) — 26°50' 65°13' (50). Ciudad Tucuman, casa centrica (Capital) — 26°50' 65° 13' (50). Colalao del Valle (Tafi del Valle)— 26°22' 65°57' (3). Concepcion (Chicligasta) — 27°20' 65°35' (93). Cuesta del 25, Ruta 9, entrada al Cadillal (Tafi Viejo)— 26°37' 65°12' (34). Cumbres Calchaquies — (no specific locality) 26°27' 65°43' (5). Cumbres de San Pablo — (probably 52). Cumbres de Santa Barbara (Trancas) — 26°00'-26°15' 65°43' (11). Cumbres del Aconquija — (no specific locality) 27° 13 66°08 (98, 99) to 27°00' 65°55' (100). Departamento Burruyacu — (no specific locality) 26°30' 64°55'. Departamento Chicligasta — (no specific locality) 27°20' 65°35'. Departamento Famailla — (no specific locality) 26°55' 65°30'. Dique Escaba (J. B. Alberdi)— 27°40' 65°46' (94). Dique San Ignacio (La Cocha) — 27°44' 65°40' (89). El Bracho (Cruz Alta)— 26°59' 65°11' (61). El Cadillal (Tafi Viejo)— 26°37' 65°12' (34). El Cadillal Dike, 25 km NW San Miguel de Tucuman (Tafi Viejo)— 26°37' 65°12' (34). El Cadillal, Estacion de Piscicultura (Tafi Viejo) — 26°37' 65°12' (34). El Cadillal, Usina (Tafi Viejo)— 26°37' 65° 12' (34). El Cadillal, 25 km N San Miguel de Tucuman (Tafi Viejo) — 26°37' 65° 12' (34). El Cajon (Burruyacu) — 26°32' 64°52' (69). El Duraznito (Tafi Viejo)— 26°40' 65°11' (37). El Indio, Ruta 307 (Monteros)— 27°02' 65°40' (104). El Infiernillo (Tafi del Valle)— 26°44' 65°47' (119). El Manantial (Lules)— 26°5 1 ' 65°17' (53). El Naranjal (Monteros)— 27°02' 65°41' (105). El Naranjo (Burruyacu) — 26°40' 65°03' (65). El Nogalar, km 43, Ruta 307, 1700m (Monteros)— 27°01' 65°40' (111). El Nogalar, = Rio los Sosa, km 43.0, Ruta 307 (Monteros) — 27°01' 65°40' (111). El Portezuelo (Trancas)— 26°08' 65 °27' (21). El Simbolar (Trancas) — 26°14' 65°23' (19). El Timbo (Burruyacu) — 26°42' 65°08' (41). Embalse Escaba (J. B. Alberdi)— 27°40' 65°46' (94). Escuela Normal, Monteros (Monteros) — 27°10' 65°30' (102). Estancia El Cavao (Burruyacu) — 26°30' 64°45' (71). Estancia San Pedro, Vipos (Trancas) — 26°29' 65°22' (30). Garmendia, Departamento Burruyacu (Burruyacu) — 26°34' 64°33' (75). Gobernador Garmendia (Burruyacu) — 26°34' 64°33' (75). Gobernador Piedrabuena (Burruyacu) — 26°45' 64°40' (76). Graneros (Graneros) — 27°39' 65°27' (90). Horco Molle (Yerba Buena)— 26°45' 65°21' (45). Horco Molle, 15 km SW San Miguel de Tucuman (Yerba Buena) — 26°45' 65°21' (45). 1996 Mares et al. — TucumAn Mammals 147 Horco Molle, 15 km W San Miguel de Tucuman (Yerba Buena) — 26°45' 65°2r (45). Horco Molle, 25 km NW San Miguel de Tucuman (Yerba Buena) — 26°45' 65°2r (45). 4 km W of junction 338 and road to Horco Molle, on road to San Javier, 2,750 ft. (Yerba Buena) — 26°49' 65°2r (57). Iglesia (not located). Ingenio Amalia (Capital) — 26°52' 65°13' (51). Ingenio Lules (Lules) — 26°55' 65°20' (55). Ingenio San Pablo (Famailla)— 26°54' 65° 19' (54). Ingenio Santa Lucfa (Monteros) — 27°06' 65°3T (108). Instituto Lillo (Capital)— 26°49' 65°13' (50). Instituto Lillo, San Miguel de Tucuman (Capital)— 26°49' 65°13' (50). La Angostura (Taff del Valle)— 26°55 ' 65°41' (114). La Cienaga, 2500 m (Taff del Valle)— 26°46' 65°39' (117). La Cocha (La Cocha)— 27°47' 65°34' (87). La Cocha, Dique San Ignacio (La Cocha) — 27°44' 65°40' (89). La Cocha, 380m (La Cocha)— 27°47' 65°34' (87). La Cuesta (Trancas) — (not located). La Florida Reserva Provincial — (no specific locality). La Fronterita (Famailla)— 27°02' 65°28' (113). La Heladera, 2 km below, along Hwy 307, 3,500 ft. (Monteros) — 27°01' 65°40' (103). La Higuera (Trancas)— 26°23' 65°26' (27). La Quebradita, 12 km W, km 81 along Hwy 307, 9,500 ft. (Taff del Valle)— 26°44' 65°45' (116). La Ramada (Burruyacu) — 26°42' 64°57' (67). La Reduccion (Lules)— 27°57' 65°22' (59). Laguna Huaca Huasi (Taff Viejo) — 26°41' 65°44' (120). Laguna Huaca Huasi, Cumbres Calchaqufes, approximately 4250 m (Taff Viejo) — 26°41' 65°44' (120). Las Higuerillas, 0.5 km N on Hwy 308, 3,000 ft. (J. B. Alberdi) — 27°47' 65°50' (95). Las Higuerillas, 5 km N on Hwy 308, 2,900 ft. (J. B. Alberdi)— 27°44' 65°50' (95). Las Juntas, 22 km W Choromoro on Hwy 312, 3,500 ft. (Trancas) — 26°24' 65°31' (26). Las Mesadas (Trancas) — 26°27' 65°30' (14). Las Pavas (Chicligasta)— 27°15' 65°52' (97). Las Pavas, Aconquija (Chicligasta) — 27°15' 65°52' (97). Las Talas (Leales)— 27°00' 65°17' (60). Las Talas, 4 km al N de Bella Vista (Leales)— 27°(X)' 65°17' (60). Las Talitas (Monteros) — ^(not located). Leales (Leales)— 27° 12' 65°18' (78). Leocadio Paz (Trancas) — 26°09' 65° 18' (23). Los Puestos (Leales)— 27° 17' 65°00' (80). Los Romanos (Leales) — 27°24' 65°08' (82). Los Sarmientos (Rfo Chico) — 27°24' 65°41' (91). Los Vazquez (Capital) — 26°50' 65° 13' (50). Los Vazquez, 445m (Capital) — 26°50' 65°13' (50). Lules, 3 km W (Lules)— 26°56' 65°23' (58). Mala Mala (Lules)— 26°47' 65°33' (118). Marcos Paz (Yerba Buena) — 26°49' 65°17' (47). Monte Bello (Rfo Chico)— 27°22' 65°45' (92). Monte Bello (Trancas)— 26° 14' 65°33' (7). Monteagudo (Simoca) — 27°31' 65°17' (85). Monteros (Monteros)— 27° 10' 65°30' (102). Morro del Zarzo, 5000 m (Taff del Valle)— 27°00' 65°54' (100). Munecas (Capital)— 26°47' 65°15' (52). Near the campground near Cochuna (Concepcfon) — ^27°18' 65°55' (121). Nevados del Aconquija, Glacial del Cochuna, 4550 m (Chicligasta) — 27°13' 66°08' (98). Norco, 2500 m (Trancas)— 26°29' 65°22' (28). Norco, Vipos, 2500 m (Trancas)— 26°29' 65°22' (28). Nueva Espafia (Leales) — “27°16' 65°12' (81). Piedra Buena (Burruyacu) — 26°44' 64°40' (76). Piedra Tendida (Burrayacu) — 26°30' 64°52' (69), Piedra Tendida, 12 km WNW Burruyacu, along Rfo Cajon, 2,500 ft. (Burruyacu) — 26°30' 64°52' (69). Piedras Coloradas (Monteros) — 27°04' 65°40' (101). Playa Larga (Monteros)— 27°03 ' 65°40' (107). 148 Annals of Carnegie Museum VOL. 65 Plaza Independencia, San Miguel de Tucuman (Capital) — 26°50' 65°13' (50). Pozo Hondo (Graneros) — 27°49' 65°20' (86). Puesto Portadera (Burmyacu) — (not located). Quebrada de la Angostura (Trancas) — 26°12' 65°3r (8). Quebrada de las Burras (Trancas) — 26°19' 66°04' (6). Quebrada de los Matos (Trancas) — 26°20’ 65°22' (13). Quebrada de Lules, 11 km NW San Pablo (Lules)— 26°52' 65°25' (48). Quebrada del Toro (Trancas)— 26°1 8' 65°21' (20). Raco, Sierra San Javier (Taff Viejo) — 26°38' 65°26' (33). Rearte Norte (Trancas) — 26°06' 65°28' (25). Reserva Provincial “Aguas Chiquitas" El Cadillal (Burmyacu) — 26°37' 65°12' (35). Reserva Provincial El Cadillal “Aguas Chiquitas" (Burmyacu) — 26°37' 65°12' (35). Rfo Colorado (Famailla)— 27°09' 65°21' (110). Rio de Los Sosa, km 19.7, Ruta 307, camino a Taff del Valle, 700 m (Monteros) — 27°05' 65°40' (96). Rio de Los Sosa, km 19.7, Ruta 307, camino a Taff del Valle, 750 m (Monteros) — 27°05' 65°40' (96). Rfo de Los Sosa, km 23.9, Ruta 307, camino a Taff del Valle, 850 m (Monteros) — 27°04' 65°40' (106). Rfo de Los Sosa, km 24.9, Ruta 307, camino a Taff del Valle, 850 m (Monteros) — 27°04' 65°40' (106). Rfo Las Tacanas (Trancas) — 26° 12' 65°27' (10). Rfo Loro (Burmyacu) — 26°37' 65° 12' (34). Rfo Los Sosas, Ruta 307, km 23.9 (Monteros) — 27°04' 65°40' (106). Rfo Nfo (Burmyacu) — 26°26' 64°56' (66). Ruta 307, km 19.7, camino a Taff del Valle, 750 m (Monteros) — 27°05' 65°40' (96). Ruta 307, km 23.9, 850 m (Monteros)— 27°04' 65°40' (106). San Fernando (Trancas) — 26°22' 65°27' (17). San Javier (Yerba Buena)— 26°47' 65°23' (42). San Javier, Ciudad LFniversitaria (Taff Viejo) — 26°48' 65°23' (43). San Miguel de Tucuman (Capital) — 26°50' 65°13' (50). San Miguel de Tucuman, 450m (Capital) — 26°50' 65° 13' (50). San Miguel de Tucuman, 28 km NE (Burmyacu) — 26°37' 65°07' (64). San Miguel de Tucuman, 29 km NE (Burmyacu) — 26°37' 65°07' (64). San Miguel de Tucuman, 17 km NW (Taff Viejo)— 26°41 ' 65°18' (18). San Miguel de Tucuman, 25 km NW (Taff Viejo) — 26°38' 65°20' (22). San Pablo (Lules)— 26°52' 65° 19' (54). San Pablo, 11 km al NO (Lules)— 26°52' 65°25' (48). San Pedro de Colalao (Trancas) — 26°14' 65°29' (16). San Pedro de Colalao, Junta de los Rfos (Trancas) — 26° 14' 65°29' (16). San Pedro de Colalao, 1 1 km S (Trancas)— 26°20' 65°33' (12). San Pedro de Colalao, south of, at km marker 42, on highway 364, 4,700 ft. (Trancas) — 26°20' 65°33' (12). Santa Ana Reserva Provincial (Rfo Chico and J. B. Alberdi) — 27°30' 65°55' (88). Santa Rosa de Leales (Leales) — 27°09' 65°15' (77). Serranfa de Quilmes-Cajon (Taff del Valle) — 26°35' 66°10' (2). Sierra de Tucuman — (no specific locality). Sierra del Aconquija (Monteros and Taff del Valle) — 27°00' 65°55' (100). Sierras de Medina, Aguas Chiquitas, “El Cadillal" (Burmyacu) — 26°37' 65°12' (35). Sierras del Cajon (Taff del Valle)— 26°35' 66° 10' (2). Tacanas (Leales)— 27°08' 64°49' (79). Taco Yana (Trancas) — 26°11' 65°30' (9). Taff, 2000 m — (no specific locality). Taff del Valle (Taff del Valle)— 26°52' 65°41' (115). Taff Viejo (Taff Viejo)— 26°44' 65° 16' (44). Taficillo (Taff Viejo)— 26°42' 65°17' (38). Tapia (Trancas)— 26°36' 65°18' (32). Tapia, 500 m (Trancas) — 26°36' 65° 18' (32). Tapia de Tucuman, 600 m (Trancas) — 26°36' 65° 18' (32). Ticucho (Trancas)— 26°31' 65°15' (31). Ticucho, cola de embalse El Cadillal (Trancas) — 26°31' 65° 14' (39). Ticucho, entrando por cola del Dique el Cadillal (Trancas) — 26°31' 65°14' (39). Ticucho, 3 km E, cola de Cadillal Dique (Trancas) — 26°3r 65°14' (39). 1996 Mares et al. — Tucuman Mammals 149 Timbo Nuevo (Burruyacu) — 26°42' 65°07' (40). Trancas (Trancas) — 26°13' 65°17' (24). Tranquitas (Burruyacu) — 26°37' 65°02' (36). Tucuman — (no specific locality). Tucuman, 450 m (Capital)— 26°50' 65°13' (50). Tucuman, 1500m — (no specific locality). Villa Nougues (Lules)— 26°5 C 65°23^ (52). Vipos (Trancas)— 26°29' 65°22' (29). Vipos, Estancia San Pedro (Trancas)— 26°29' 65°22' (30). Vipos, 500 m (Trancas)— 26°29' 65°22' (29). Yerba Buena (Yerba Buena) — 26°49' 65°19' (46). 1. Amaicha del Valle (Tafi del Valle)— 26°36' 65°55'. Amaicha del Valle, 2000 m (Tafi del Valle)— 26°36' 65°55'. 2. Serranfa de Quilmes-Cajon (Tafi del Valle)— 26°35' 66° 10'. Sierras del Cajon (Tafi del Valle)— 26°35' 66° 10'. 3. Colalao del Valle (Tafi del Valle)— 26°22' 65°57'. 4. Cafayate, 45 km S, along Hwy 40 (Tafi del Valle)— 26°20' 65°58'. 5. Cumbres Calchaquies-(Trancas and Tafi del Valle) — 26°27' 65°43'. 6. Quebrada de las Burras (Trancas) — 26°19' 66°04'. 7. Monte Bello (Trancas)— 26° 14' 65°33'. 8. Quebrada de la Angostura (Trancas) — 26°12' 65°3r. 9. Taco Yana (Trancas)— 26°1 1 ' 65°30'. 10. Rfo Las Tacanas (Trancas) — 26° 12' 65°27'. 11. Cumbres de Santa Barbara (Trancas) — 26°00'-26°15' 65°43'. 12. San Pedro de Colalao, 11 km S (Trancas)— 26°20' 65°33'. San Pedro de Colalao, south of, at km marker 42, on highway 364, 4,700 ft. (Trancas) — 26°20'65°33'. 13. Quebrada de los Matos (Trancas) — 26°20' 65°22'. 14. Las Mesadas (Trancas) — 26°27' 65°30'. 15. Agua Rosada (Trancas) — 26°22' 65°26'. Agua Rosada, San Pedro de Colalao (Trancas) — 26°22' 65°26'. 16. San Pedro de Colalao (Trancas) — 26°14' 65°29'. San Pedro de Colalao, Junta de los Rios (Trancas) — 26°14' 65°29'. 17. San Fernando (Trancas) — 26°22' 65°27'. 18. San Miguel de Tucuman, 17 km NW (Tafi Viejo) — 26°41' 65°18'. 19. El Simbolar (Trancas) — 26° 14' 65°23'. 20. Quebrada del Toro (Trancas) — 26°18' 65°21'. 21. El Portezuelo (Trancas) — 26°08' 65°27'. 22. San Miguel de Tucuman, 25 km NW (Tafi Viejo) — 26°38' 20°12'. 23. Leocadio Paz (Trancas) — 26°09' 65°18'. 24. Trancas (Trancas) — 26°13' 65° 17'. 25. Rearte Norte (Trancas) — 26°06' 65°28'. 26. Las Juntas, 22 km W Choromoro on Hwy 312, 3,500 ft. (Trancas) — 26°24' 65°31'. 27. La Higuera (Trancas)— 26°23' 65°26'. 28. Norco, 2500 m (Trancas) — 26°29' 65°22'. Natnorco, Vipos, 2500 m (Trancas)— 26°29' 65°22'. 29. Vipos (Trancas)— 26°29' 65°22'. Vipos, 500 m (Trancas) — 26°29' 65°22'. 30. Estancia San Pedro, Vipos (Trancas) — 26°29' 65°22'. Vipos, Estancia San Pedro (Trancas) — 26°29' 65°22'. 31. Ticucho (Trancas)— 26°31' 65° 15'. 32. Tapia (Trancas) — 26°36' 65° 18'. Tapia, 500 m (Trancas)— 26°36' 65° 18'. Tapia de Tucuman, 600 m (Trancas) — 26°36' 65° 18'. 33. Raco, Sierra San Javier (Tafi Viejo) — 26°38' 65°26'. 34. Cuesta del 25, Ruta 9, entrada al Cadillal (Tafi Viejo) — 26°37' 65°12'. El Cadillal (Tafi Viejo)— 26°37' 65° 12'. El Cadillal Dike, 25 km NW San Miguel de Tucuman (Tafi Viejo) — 26°37' 65°12'. El Cadillal, Estacion de Piscicultura (Tafi Viejo) — 26°37' 65°12'. El Cadillal, 25 km N San Miguel de Tucuman (Tafi Viejo) — 26°37' 65°12'. 150 Annals of Carnegie Museum VOL. 65 El Cadillal, Usina (Tafi Viejo)— 26°37' 65° 12'. Rio Loro (Burruyacu) — 26°37' 65° 12'. 35. Aguas Chiquitas (Burruyacu) — 26°37' 65° 12'. Aguas Chiquitas, El Cadillal, 800m (Burruyacu) — 26°37' 65°12'. Aguas Chiquitas, Sierras de Medina, 800m (Burruyacu) — 26°37' 65° 12'. Aguas Chiquitas, 25 km NE San Miguel de Tucuman, 800 m (Burruyacu) — 26°37' 65°12'. Reserva Provincial El Cadillal “Aguas Chiquitas" (Burruyacu) — 26°37' 65°12'. Reserva Provincial “Aguas Chiquitas" El Cadillal (Burruyacu) — 26°37' 65°12'. Sierras de Medina, Aguas Chiquitas, “El Cadillal" (Burruyacu) — 26°37' 65°12'. 36. Tranquitas (Burruyacu) — 26°37' 65°02'. 37. El Duraznito (Tafi Viejo)— 26°40' 65°11'. 38. Talicillo (Tafi Viejo)— 26°42' 65°17'. 39. Ticucho, entrando por cola del Dique el Cadillal (Trancas) — 26°31' 65°14'. Ticucho, cola de embalse El Cadillal (Trancas) — 26°31' 65° 14'. Ticucho, 3 km E, cola de Cadillal Dique (Trancas) — 26°31' 65° 14'. 40. Timbo Nuevo (Burruyacu) — 26°42' 65°07'. 41. El Timbo (Burruyacu) — 26°42' 65°08'. 42. Cerro San Javier (Yerba Buena) — 26°47' 65°24'. San Javier (Yerba Buena) — 26°47' 65°23'. 43. San Javier, Ciudad Universitaria (Tafi Viejo) — 26°48' 65°23'. 44. Tafi Viejo (Tafi Viejo) — 26°44' 65° 16'. 45. Horco Molle (Yerba Buena)— 26°45' 65°21'. Horco Molle, 15 km W San Miguel de Tucuman (Yerba Buena) — 26°45' 65°21'. Horco Molle, 15 km SW San Miguel de Tucuman (Yerba Buena) — 26°45' 65°21'. Horco Molle, 25 km NW San Miguel de Tucuman (Yerba Buena) — 26°45' 65°21'. Biological Reserve at Horco Molle, 2,400 ft. (Yerba Buena) — 26°45' 65°21'. Biological Reserve at Horco Molle, near residencia, 2,400 ft. (Yerba Buena) — 26°45' 65°21'. 46. Yerba Buena (Yerba Buena)— 26°49' 65° 19'. 47. Marcos Paz (Yerba Buena)— 26°49' 65° 17'. 48. San Pablo, 1 1 km al NO (Lules)— 26°52' 65°25'. Quebrada de Lules, 11 km NW San Pablo (Lules) — 26°52' 65°25'. 49. Munecas (Capital)— 26°47' 65° 15'. 50. Los Vazquez (Capital)— 26°50' 65° 13'. Los Vazquez, 445m (Capital) — 26°50' 65° 13'. San Miguel de Tucuman (Capital) — 26°50' 65° 13'. Tucuman, 450 m (Capital) — 26°50' 65° 13'. San Miguel de Tucuman, 450m (Capital) — 26°50' 65° 13'. Capital, San Miguel de Tucuman (Capital) — 26°50' 65° 13'. Ciudad Capital (Capital)— 26°50' 65° 13'. Ciudad Tucuman, casa centrica (Capital) — 26°50' 65° 13'. Instituto Lillo (Capital) — 26°49' 65° 13'. Instituto Lillo, San Miguel de Tucuman (Capital) — 26°49' 65°13'. Plaza Independencia, San Miguel de Tucuman (Capital) — 26°50' 65° 13'. 51. Ingenio Amalia (Capital) — 26°52' 65°13'. 52. Villa Nougues (Lules) — 26°51' 65°23'. Cumbres de San Pablo — (probably 55). 53. El Manantial (Lules)— 26°5 1 ' 65° 17'. 54. Ingenio San Pablo (Famailla) — 26°54' 65° 19'. San Pablo (Lules)— 26°52' 65°19'. 55. Ingenio Lules (Lules)— 26°55' 65°20'. 56. Cerro de Raco, Trancas (Tafi Viejo) — 26°43' 65°30'. 57. 4 km W of junction 338 and road to Horco Molle, on road to San Javier, 2,750 ft. (Yerba Buena)— 26°49' 65°21'. 58. Lules, 3 km W (Lules)— 26°56' 65°23'. 59. La Reduccion (Lules)— 27°57' 65°22'. 60. Las Talas (Leales)— 27°00' 65° 17'. Las Talas, 4 km al N de Bella Vista (Leales) — 27°00' 65° 17'. 61. El Bracho (Cruz Alta)— 26°59' 65°11'. 62. Banda del Rio Sail (Cruz Alta) — 26°50' 65° 10'. 63. Alderetes (Cruz Alta)— 26°49' 65°08'. 1996 Mares et al. — Tucuman Mammals 151 64. San Miguel de Tucuman, 28 km NE (Burruyacu) — 26°37' 65°07'. San Miguel de Tucuman, 29 km NE (Burruyacu) — 26°37' 65°07'. 65. El Naranjo (Burruyacu) — 26°40' 65°03'. 66. Rio Nio (Burruyacu) — 26°26' 64°56'. 67. La Ramada (Burruyacu) — 26°42' 64°57'. 68. Cerro del Campo (Burruyacu) — 26°35' 64°57'. Cerro del Campo, 800 m (Burruyacu) — 26°35' 64°57'. Cerro del Campo, 1800 m (Burruyacu) — 26°35' 64°57'. 69. El Cajon (Burruyacu) — 26°32' 64°52'. Piedra Tendida (Burruyacu) — 26°30' 64°52'. Piedra Tendida, 12 km WNW Burruyacu, along Rio Cajon, 2,500 ft. (Burruyacu) — 26°30' 64°52'. 70. Cajon de la Ovejeria, Chorrillos (Burruyacu) — 26°2r 64°55'. 71. Estancia El Cavao (Burruyacu) — 26°30' 64°45'. 72. Agua Colorada (Burruyacu) — 26°26' 64°53'. 73. Burruyacu (Burruyacu) — 26°30' 64°45'. Burruyacu, 1300 m (Burruyacu) — 26°30' 64°45'. 74. Anta Mapu (Burruyacu) — 26°31' 64°43'. 75. Garmendia, Departamento Burruyacu (Burruyacu) — 26°34' 64°33'. Gobernador Garmendia (Burruyacu) — 26°34' 64°33'. 76. Gobernador Piedrabuena (Burruyacu) — 26°40' 64°40'. Piedra Buena (Burruyacu) — 26°44' 64°40'. 77. Santa Rosa de Leales (Leales) — 27°09' 65° 15'. 78. Leales (Leales)— 27° 12' 65°18'. 79. Tacanas (Leales) — 27°08' 64°49'. 80. Los Puestos (Leales)— 27° 17' 65°00'. 81. Nueva Espana (Leales)— 27° 16' 65° 12'. 82. Los Romanos (Leales) — 27°24' 65°08'. 83. Atahona (Simoca)— 27°25' 65° 17'. 84. Chicligasta (Simoca)— 27°28' 65°07'. 85. Monteagudo (Simoca) — 27°31' 65°17'. 86. Pozo Hondo (Graneros) — 27°49' 65°20'. 87. La Cocha (La Cocha)— 27°47' 65°34'. La Cocha, 380m (La Cocha)— 27°47' 65°34'. 88. Santa Ana Reserva Provincial (Rio Chico and J. B. Alberdi) — 27°30' 65°55'. 89. Dique San Ignacio (La Cocha) — 27°44' 65°40'. La Cocha, Dique San Ignacio (La Cocha) — 27°44' 65°40'. 90. Graneros (Graneros) — 27°39' 65°27'. 91. Los Sarmientos (Rfo Chico) — 27°24' 65°41'. 92. Monte Bello (Rio Chico)— 27°22' 65°45'. 93. Concepcion (Chicligasta) — 27°20' 65°35'. 94. Dique Escaba (J. B. Alberdi)— 27°40' 65°46'. Embalse Escaba (J. B. Alberdi) — 27°40' 65°46'. 95. Las Higuerillas, 0.5 km N on Hwy 308, 3,000 ft. (J. B. Alberdi)— 27°47' 65°50'. Las Higuerillas, 5 km N on Hwy 308, 2,900 ft. (J. B. Alberdi)— 27°44' 65°50'. 96. Rfo de Los Sosa, km 19.7, Ruta 307, camino a Taff del Valle, 700 m (Monteros) — 27°05' 65°40'. Rfo de Los Sosa, km 19.7, Ruta 307, camino a Taff del Valle, 750 m (Monteros) — 27°05' 65°40'. Ruta 307, km 19.7, camino a Taff del Valle, 750 m (Monteros) — 27°05' 65°40'. 97. Las Pavas (Chicligasta)— 27°15' 65°52'. Las Pavas, Aconquija (Chicligasta) — 27°15' 65°52'. 98. Nevados del Aconquija, Glacial del Cochuna, 4550 m (Chicligasta) — 27°13' 66°08'. 99. Aconquija (Chicligasta) — 27°13' 66°08'. Aconquija, 3000 m (Chicligasta) — 27°13' 66°08'. 100. Morro del Zarzo, 5000 m (Taff del Valle)— 27°00' 65°54'. Sierra del Aconquija (Monteros and Taff del Valle) — 27°00' 65°55'. 101. Piedras Coloradas (Monteros) — -27°04' 65°40'. 102. Monteros (Monteros) — ^27°10' 65°30'. Escuela Normal, Monteros (Monteros) — 27° 10' 65°30'. 103. La Heladera, 2 km below, along Hwy 307, 3,500 ft. (Monteros) — 27°01' 65°40'. 104. El Indio, Ruta 307 (Monteros)— 27°02' 65°40'. 105. El Naranjal (Monteros)— 27°02' 65°41'. 106. Casa de Piedra, Rfo Los Sosa (Monteros) — 27°03' 65°37'. 152 Annals of Carnegie Museum VOL. 65 Casa de Piedra, Ruta 307, km 23.9, 850m (Monteros) — 2T03' 65°37'. Rio Los Sosas, Ruta 307, km 23.9 (Monteros) — 27°04' 65°40'. Rio de Los Sosa, km 23.9, Ruta 307, camino a Tafi del Valle, 850 m (Monteros) — 27°04' 65°40'. Rio de Los Sosa, km 24.9, Ruta 307, camino a Tafi del Valle, 850 m (Monteros) — 27°04' 65°40'. Ruta 307, km 23.9, 850 m (Monteros)— 27°04' 65°40'. 107. Play a Larga (Monteros) — 27°03' 65°40'. 108. Ingenio Santa Lucia (Monteros) — 27°06' 65°31'. 109. Acheral (Monteros) — 2T01' 65°27'. 110. Rio Colorado (Famailla)— 27°09' 65°2r. 111. El Nogalar, km 43, Ruta 307, 1700m (Monteros) — 27°01' 65°40'. El Nogalar, = Rio los Sosa, km 43.0, Ruta 307 (Monteros) — 27°01' 65°40'. 112. Caspichango (Monteros) — 27°04' 65°30'. 113. La Fronterita (Famailla) — 27°02' 65°28'. 114. La Angostura (Tafi del Valle)— 26°55' 65°41'. 115. Tafi del Valle (Tafi del Valle)— 26°52' 65°4r. 116. La Quebradita, 12 km W, km 81 along Hwy 307, 9,500 ft. (Tafi del Valle) — 26°44' 65°45'. 117. La Cienaga, 2500 m (Tafi del Valle)— 26°46' 65°39'. 118. Mala Mala (Lules)— 26°47' 65°33'. 119. El Infiernillo (Tafi del Valle)— 26°44' 65°47'. 120. Laguna Huaca Huasi (Tafi Viejo) — 26°41' 65°44'. Laguna Huaca Huasi, Cumbres Calchaquies, approximately 4250 m (Tafi Viejo) — 26°41' 65°44'. 121. Near the campground near Cochuna (Concepcion) — 27°18' 65°55' (135). Campo de la Gallina, Serrania de Quilmes o El Cajon (Tafi del Valle) — (not located). Cerro de Tafi Viejo, 2400 m (Tafi Viejo)— 26°42' 65°17' and 26°44' 65°16 (between 38 and 44). Cumbres del Aconquija — (no specific locality) 27°13' 66°08' (98, 99) to 27°00' 65°55' (100). Departamento Burruyacu — (no specific locality) 26°30' 64°55'. Departamento Chicligasta — (no specific locality) 27°20' 65°35'. Departamento Famailla — (no specific locality) 26°55' 65°30'. Iglesia — (not located). La Cuesta (Trancas) — (not located). La Florida Reserva Provincia — (no specific locality). Las Talitas (Monteros) — (not located). Puesto Portadera (Burruyacu) — (not located). Sierra de Tucuman — (no specific locality). Tafi, 2000 m — (no specific locality). Tucuman — (no specific locality). Tucuman, 1500 m — (no specific locality). ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 2, Pp. 153-166 16 May 1996 BIOLOGY AND IMMATURE STAGES OF SNAIL-KILLING FLIES BELONGING TO THE GENUS TETANOCERA (INSECTA: DIPTERA: SCIOMYZIDAE). IL LIFE HISTORIES OF PREDATORS OF SNAILS OF THE FAMILY SUCCINEIDAE B. A. Foote^ Research Associate, Section of Invertebrate Zoology Abstract Information on the life histories and larval feeding habits of four Nearctic species of Tetanocera that prey on snails belonging to the pulmonate snail family Succineidae are presented. The four species are quite prey-specific, as larvae of Tetanocera melanostigma Steyskal prey on species of Succinea, those of T. oxia Steyskal and T. spirifera Melander attack snails of the genus Catinella, and larvae of T. rotundicornis Loew are associated with the genus Oxyloma. All four species are multivoltine and occur in marshy or swampy habitats containing large populations of succineid snails. Introduction This is the second of a series of papers covering the life histories and larval feeding habits of the North American species of Tetanocera. The first paper (Foote, 1996) reviewed literature, gave information on rearing techniques, and discussed the biology of two species whose larvae attack shoreline-inhabiting pulmonate snails. In this paper I present information on the life histories of four species of Te- tanocera, T. melanostigma Steyskal, T. oxia Steyskal, T. rotundicornis Loew, and T. spirifera Melander, whose larvae prey on species of succineid snails. The larval feeding habits are described, and information on trophic and spatial resource par- titioning in this guild of snail-killing Tetanocera is presented. Species of the three North American genera of pulmonate snails belonging to the family Succineidae are commonly called “amphibious” because of their oc- currence in wetland habitats, even though they are not able to survive prolonged submersion. They are frequently encountered in shoreline habitats, although Suc- cinea ovalis Say is more common in swampy and mesic forests. Species of Ox- yloma seemingly prefer open herbaceous marshes, whereas Catinella species more commonly occur in shaded habitats beneath shrubs and along the wooded banks of streams. Life Histories Tetanocera melanostigma Steyskal Steyskal, 1959. Papers of the Michigan Academy of Science, Arts, and Letters, 44:60. Tetanocera melanostigma is restricted to the Nearctic region (Knutson et al., 1986), where it is known to range from New Hampshire to Ontario and south to New York and Ohio (Fig. 1). This species was taken in central New York and northeastern Ohio in moist to ' Department of Biological Sciences, Kent State University, Kent, Ohio 44242. Submitted 28 April 1995. 153 154 Annals of Carnegie Museum VOL, 65 wet floodplain forests and around shaded hillside seepages at the foot of forested slopes. Two sites were especially productive of flies. The first site was the flood- plain bordering the outlet stream of Cayuta Lake in Schuyler County, New York, where a swamp forest is broken into a mosaic of habitats. The forest canopy. 1996 Foote — Tetanocera Taxonomy and Biology II 155 composed mostly of American elm (Ulmus americana L.), is irregularly broken and allows considerable illumination to reach the forest floor. As a result there is a luxuriant, if discontinuous, herbaceous layer. Colonies of cattail {Typha latifolia L.) are found in the wetter, more open sites, and there is a thick layer of decaying leaf litter in the more forested areas. The soil is permanently moist, and during the spring months the whole area commonly is covered by 3 cm or more of water. When the area was visited during early September, Succinea snails, the preferred food of the larvae, were most abundant on the leaves of cattail, although adults of T. melanostigma were more widely distributed over the floodplain. Unfortu- nately, no larvae were found in the snails collected at this site. The second locality visited was a seepage located at the foot of a forested, northwest-facing slope lying along the Ringwood Branch of Cascadilla Creek about 1 1 km east of Ithaca, New York. The area is well shaded by deciduous trees and covered by a luxuriant layer of herbs, mostly pale touch-me-not {Impatiens pallida Nuttall) and joe-pye- weed {Eupatorium maculatum L.). The highly organic soil is always wet and may be submerged during the spring and early summer months. Succinea snails were abundant on the leaves of the joe-pye-weed, less common in the stands of Im- patiens, and quite rare on a narrow-leaved species of Carex. All snails were inactive and attached to the leaves by a mucous secretion when collected on July 14 and 30. Four (11%) of the 35 Succinea collected on July 14 contained one or more larvae of T. melanostigma, but a larva was found in only one (4%) of 23 snails taken on July 30. Throughout the summer, many viable eggs were obtained from both feral and laboratory-reared females. Most eggs were placed on peat moss in the jars, but a few were scattered over the glass walls. Although eggs were not found in nature, it is probable that they are deposited on vegetation near the snail prey. The in- cubation period lasted five to seven days under laboratory conditions {n = 35). Observations were made on the activities of 20 first instars that hatched during the summer months. Groups of five to eight larvae were placed in each of several large Stender dishes containing moist sand and one or two living S. ovalis. The larvae moved slowly over the sand, stopping frequently and lifting the anterior parts of their bodies off the substrate and waving them to and fro. When they brushed against a Succinea, larvae became highly excited and attempted to crawl up onto the shell. Many then crawled onto the lateral surfaces of the expanded foot and attempted to insert themselves between the foot and the collar forming the edge of the mantle. However, only 14 (70%) of the 20 larvae actually managed to invade seven snails. Other larvae either failed to contact their potential prey or became entangled in the mucous secretion produced by the snails. These latter larvae invariably succumbed and were later found imbedded in small pellets of mucus along the sides and bottoms of the rearing dishes. Examination of three infested snails three or four days after larval attack revealed that each had one to three larvae embedded between the foot and mantle collar. One had three larvae along one side of the foot, and the other two each had a larva present on one side of the foot. No larvae had invaded the breathing orifice of the snails, although three were lying on either side of the opening. The snail containing three larvae remained alive for five days, apparently succumbing when the larvae molted to the second instar. This snail decayed rapidly after its death, but the larvae con- tinued to feed for another day on the decomposing flesh. The following day, all three larvae had left the shell and were crawling about on the moist sand. A living, uninfested Succinea was added to the dish, and by the next day, two of 156 Annals of Carnegie Museum VOL. 65 the larvae were embedded between the foot and the mantle collar. The third larva had died. The two living larvae continued to feed on the snail and molted to the third instar, but only one larva managed to complete development and form a puparium. The second larva was found dead on the moist sand. In all, only two puparia were obtained from the 14 larvae that initially had infested the Succinea snails. Both puparia were formed in the moist sand. The four Succinea containing five larvae collected on July 14 were placed in individual Stender dishes containing moist, coarse sand. One snail was alive, but very feeble, and retracted into its shell when collected from a sedge leaf. Its breathing orifice opened and closed spasmodically, and waves of faint tremors passed over its foot. Two second-instar larvae were imbedded between its foot and mantle collar. Each larva had most of its body buried in the snail, and only the posterior spiracles were visible at the surface of the prey. On the morning of July 16, the snail was dead, but both larvae were still in place laterad of the foot. Two second-instar cephalopharyngeal skeletons and one posterior spiracular disc were recovered from the decaying foot of the snail, indicating that both sciomyzid larvae had molted to the third instar. By July 17 the snail had been reduced to a few shreds of tissue, and only one larva was still within the shell. The second larva was lying quietly on the sand. Another Succinea was placed in the Stender dish, and by the following day it was invaded by the larva that last left the original snail. A third Succinea was added to the dish on July 18. On July 19 both larvae had vacated the snails and lay on the moist sand. The Succinea added the previous day was alive, although inactive, and unattacked. On July 20, one larva burrowed into the sand, while the other invaded the Succinea. The snail soon showed dis- tress by producing frothy bubbles around its breathing orifice and a dark yellow mucous secretion on its foot. The snail was quickly killed and largely consumed, and by July 21 both larvae had formed puparia below the surface of the sand. Another snail was not observed to contain a larva until July 17 when its re- tracted condition, waves of faint tremors passing over its foot, and an abundance of frothy bubbles around the breathing orifice indicated that it was in distress. At first, no larva could be seen, but within a minute, a small second instar thrust its posterior spiracles to the surface of the flesh laterad of the foot. The snail finally died on July 23, nine days after being collected, presumably harboring a devel- oping larva all that time. By the ninth day the larva had moved to a position outside the mantle against the parietal wall of the shell. By the following day the snail was badly decomposed and partly covered by fungal hyphae, but the larva, now a third instar, was still within the host’s shell, although not in a feeding position, as its head was directed away from the decaying flesh. On July 26 the larva left the snail and partly buried itself in the moist sand, where it underwent pupariation during the night of July 27. None of the five larvae found in the four field-collected Succinea attacked a second snail, except for the two in the first described snail whose food supply had been restricted by intraspecific competition. All larvae killed their prey and all partly or completely buried themselves in the moist sand before undergoing pupariation. The various rearings indicated that larvae of T. melanostigma tend toward parasitoidism, as they remain within the food snail for extended periods of time before killing it and probably are capable of completing development within one large snail not containing other larvae. However, they may leave de- caying and largely consumed snails to attack another living prey. 1996 Foote — Tetanocera Taxonomy and Biology II 157 Under laboratory conditions the first larval stadium lasted six to eight days; the second, three to five days; and the third, from five to 1 1 days (n = 10). Usually a few hours intervened between the time a larva left its snail food and pupariated. Some larvae pupariated within four hours after leaving their hosts, but others waited nearly three days before forming puparia. All puparia were oriented parallel to the bottom of the dish or at a slight angle from the horizontal. Under laboratory conditions the pupal stage lasted 14 to 17 days (n = 8). The period between emergence of a female and her first oviposition lasted from two to five days. Two females that emerged on August 1 1 began laying eggs on August 13, although neither fly had mated. All their eggs failed to hatch. A wild- caught female produced a total of 213 eggs over a period of 46 days (May 31 to July 15). Her daily egg production was quite variable, ranging from three to 22. Evidently, T. melanostigma larvae can also develop on other species of sue- cineid snails. Of 100 living Oxyloma ejfusa (Pfeiffer) collected at the Inlet Valley marsh on August 13, 1956, two contained a single third instar of T. melanostigma. The larvae pupariated in moist sand on August 30 and 31, and adults emerged on September 13 and 14. In central New York, the earliest seasonal record for adults was May 25, 1957; the latest date was September 5, 1956. Adults were taken in each of the summer months. In northeastern Ohio, adults were collected between early June and late August. Because the life cycle requires about 45 days and females taken at various times during the summer laid viable eggs, T. melanostigma probably has at least two annual generations. Winter is probably passed as a pupa, although no over- wintering puparia were found. Tetanocera oxia Steyskal, 1959 Steyskal, 1959. Papers of the Michigan Academy of Science, Arts and Letters, 44:80. Tetanocera oxia is known only from the Nearctic region (Knutson et al., 1986), where it ranges from Alberta to Newfoundland and south to Colorado and New York (Fig. 2). Adults were collected near Kent in northeastern Ohio, in hydrophilic vegetation in an alder (Alnus sp.) grove separating an open marsh and an adjacent lowland forest. Tetanocera rotundicornis was more common in the more open areas of the marsh. At Butternut, Minnesota, T. oxia was found in a completely unshaded, grass-sedge meadow (L. V. Knutson, personal communication). In Montana, adults were collected from a mixture of herbaceous and woody vegetation, mostly shrubby willows {Salix spp.), bordering an open marsh. Other species of Scio- myzidae commonly collected with T. oxia included Atrichomelina pubera (Loew), Pherbellia griseola (Falle’n), P. nana (Falle’n), P. vitalis (Cresson), Antichaeta borealis Foote, Renocera brevis (Cresson), Tetanocera annae Steyskal, T. fusci- nervis (Zetterstedt), T. plebeja Loew, and T. rotundicornis. Gastropods that were particularly abundant in habitats supporting good populations of T. oxia were species of Catinella and Oxyloma and slugs of the genus Deroceras. Aquatic snails were also common and became stranded as water levels dropped during the summer months. Rearings were initiated from two females and three males collected June 1-4 around the margins of a sedge marsh near Kent, Ohio. Adults collected on June 1 mated on June 2, and a female began ovipositing on June 15. Only one male and one female survived more than 30 days in the 158 Annals of Carnegie Museum VOL. 65 I m IC<0 lAfi laboratory. The male lived until September 5, and the female lived until Septem- ber 15. This female deposited 99 eggs between June 15 and September 2. Most eggs were scattered over the peat moss substrate in the breeding jar, but 22 were attached to the cheesecloth covers and eight were affixed to the glass walls. Seven 1996 Foote — Tetanocera Taxonomy and Biology II 159 eggs were placed along the edges of a blade of grass. In nature, eggs probably are scattered on low-growing herbaceous vegetation or possibly scattered on the moist surface litter. The incubation period lasted five to seven days, and nearly all eggs were viable. Because T. oxia is closely related to T. rotundicornis, a predator of Oxyloma, and because snails of the family Succineidae were abundant in the adult habitat, the newly hatched larvae were exposed to individuals of the three genera of this snail family as well as to species of more aquatic and terrestrial snails. The larvae of T. oxia ignored species of Gyraulus, Aplexa, Physa, Helisoma, Planorbula, Lymnaea, and polygryid land snails, but did respond positively to the three suc- cineid genera. Newly hatched larvae were quite active and constantly wandered about the rearing dish. They showed a distinct tendency to crawl up the sides of the dish, and many desiccated on the lid. Although they crawled over all species of gas- tropods introduced to the dish, they were particularly attracted to C. avara (Say). In contrast, they displayed little or no interest in the other two succineid genera, Oxyloma and Succinea. Larvae first moved up onto the shell of Catinella and wandered about for a few minutes until they made contact with the fleshy mantle exposed in the aperture of the shell. Larvae then moved onto the mantle and attempted to implant themselves between the collar of the mantle and the side of the foot like newly-hatched larvae of T. rotundicornis. Up to four larvae were found in any one snail, but usually only one or two larvae were found in place. Most larvae came to rest near the breathing pore of the snail but several embedded themselves at the anterior end of the foot. Larvae remained in place for at least three days while they molted into the second instar, although only 12 managed to reach this stage. Only three molted into the third instar. One of these eventually formed a puparium. Many larvae apparently were rubbed off the surface of the snail’s flesh as it moved across the substrate. Other larvae became trapped in accumulations of mucus. Infested snails remained alive for four to ten days and most remained active. However, a few snails affixed themselves to the wall or lid of the rearing dish and remained inactive while being consumed by the fly larvae. Snails infested by two or more larvae generally died two to four days earlier than individuals with only one larva. Most snails died shortly after the larvae molted into the third instar. Larvae generally remained between the foot and mantle collar until they approached the final instar. Late in the second stadium or early in the third, larvae typically moved to a position between the mantle and shell. Larvae abandoned their snails and became free-roaming predators shortly after the prey expired. The single larva that eventually formed a puparium remained within its original snail host until shortly after its second molt, but it killed an additional five snails in the third stadium. Although newly hatched larvae refused to attack Oxyloma and Succinea, the third instars were more catholic in their diet and attacked snails belonging to all three genera of Succineidae. When an older larva contacted a snail, it quickly moved up onto the mantle and implanted itself be- tween the mantle and the shell. The snail was killed within a few hours, and the larva rarely remained with the snail for more than ten hours. During the one- to three-day intervals between feedings, larvae remained inactive in the peat moss. The first stadium lasted two to four days (w = 4); the second, two to six days {n ~ 4); and the third, about 17 days {n — 1). The larva that matured did not feed for four days prior to pupariating and remained buried in the peat moss. The pupal period was 20 days. 160 Annals of Carnegie Museum VOL. 65 With an incubation period of five to seven days, a larval period of 21 to 27 days, and a pupal period of 20 days, the total developmental period was 46 to 54 days. No evidence of a diapause stage was obtained. At least two generations a year are produced at the latitude of northeastern Ohio. The discovery of two floating puparia in a marsh near Ithaca, New York, on April 22 suggested that overwintering occurs as a pupa. Adults emerged from these overwintering puparia on April 29, seven days after the puparia were placed under room temperatures. The earliest (May 6) and latest (August 13) collection dates for adults were both obtained near Butternut, Minnesota. In central New York, the earliest seasonal record for adults was June 10; the latest date was September 4. In northeastern Ohio, adults were collected between mid-May and mid-July, but were most com- mon in mid-June. As a result of severe droughts in Ohio during the summers of 1987 and 1991, succineid populations were greatly reduced. As would be ex- pected, populations of succineid predators have suffered, and no adults of T. oxia have been collected by me despite repeated effort in northeastern Ohio since 1992. The larva is very light gray, similar to that of T. rotundicornis. The body wall appears velvety when dry, far more transparent when wet, with heart, intestine, tracheae, and Malpighian tubules very evident. Larvae of T. oxia lack the con- spicuous body tubercles and float hairs that characterize aquatic larvae in this genus, and the lobes of the posterior spiracular dies are reduced. Available evidence indicates that the larval feeding habits of T. oxia are very similar to those of T. rotundicornis, T. melanostigma, and T. spirifera. As with the larvae of these other species, those of T. oxia feed as parasitoids on Succi- neidae, living within the snails for prolonged periods before killing them. Like all other tetanocerine parasitoids (Berg, 1961; Berg and Knutson, 1978), these larvae evidently leave the host snail shells to pupariate in or on the soil or in ground litter. Tetanocera rotundicornis Loew 1861. Berlin Entomologische Zeitschrift, 5:344. Restricted to the Nearctic region (Knutson et al., 1986), T. rotundicornis is recorded from Ontario west to Alaska, south to North Carolina, Tennessee, and Oregon (Fig. 3). In northeastern Ohio, central New York, and Idaho, adults were found most commonly in open vernal and permanent marshes containing large populations of Oxyloma snails. However, they do not occur in all habitats containing large Oxyloma populations. They tended to be more common in unshaded sites. Adults were not taken in wooded swamps, around shaded hillside seepages, or along sluggish woodland streams, although Oxyloma snails frequently were abundant in such habitats. Rearings were initiated from larvae, puparia, and adults taken at the Inlet Val- ley, Floral Avenue, and White Church marshes in Tompkins County, New York, and from a large marsh located 9.6 km east of Kent, Portage County, Ohio. Eggs were deposited repeatedly by laboratory-reared females and by wild-type adults collected in June, July, and August. Over two-thirds of the eggs were placed on projecting sprigs of peat moss used as a substrate in the breeding jars, but several were placed on short lengths of Typha and on the glass walls. Less than ten were laid on the shells of living Oxyloma and other snails. Eggs were mostly scattered over the substrate, and no clusters of more than four eggs were observed. 1996 Foote — Tetanocera Taxonomy and Biology II 161 ira ten Mr» Under laboratory conditions, hatching occurred in seven to ten days (n = 28, Inlet Valley marsh). The newly hatched larvae were quite active and moved rapidly, if erratically, over the substrate. They stopped frequently and waved the anterior fifth of the 162 Annals of Carnegie Museum VOL. 65 body back and forth in the air. Once contact was made with the flesh of an Oxyloma snail, the larva attempted to move up onto the expanded foot, where it crawled about slowly. After these exploratory movements, larvae usually attempt- ed to insert themselves between the foot and the collar of the mantle, but a few came to rest outside of the mantle against the shell. Many larvae became trapped and died in the mucus of the snail, and others were rubbed off the foot as snails glided over the rough substrate. As many as five larvae were found embedded up to their posterior spiracular discs in one snail, but usually only one or two larvae were found in each snail. Larvae infested only succineid snails belonging to the genus Oxyloma, and attempts to rear them on snails of other families were un- successful. In central New York and northeastern Ohio, the snail species utilized as larval food is O. ejfusa, but elsewhere in North America other species are used. In Alaska, Berg (1953) found larvae attacking O. decampi gouldi Pilsbry. On July 2, 20 larvae that hatched from eggs obtained on June 25 were placed in four rearing dishes, each containing two living O. ejfusa. Only one snail was infested by July 10, and it contained two second instars. The remaining 18 larvae were found dead in balls of mucus or desiccated on the sides and tops of the rearing dishes. By July 12 the moribund infested snail had secreted an epiphragm across the shell aperture and had attached itself to the wall of the dish. Its breath- ing orifice opened and closed sporadically and faint tremors passed over its re- tracted foot. One larva was located laterad of the foot, but the second larva had temporarily disappeared. On July 16 both larvae were in the third instar. One was lying on the moist sphagnum and when placed on a living snail did not attack and quickly returned to the moss, where it died the next day. The other larva had also left the original snail and had embedded itself up to its spiracular disc in a second Oxyloma. This snail was inactive, retracted, and attached to the wall of the dish. By July 18, the larva had left the second snail and was lying on the glass lid of the dish with its gut filled with food. Other living Oxyloma were added, and the larva consumed two more individuals before forming a puparium in the peat moss on July 24. It had remained a larva for 22 days (July 2 to 24). On July 5, 11 larvae were obtained from eggs laid on June 28. The larvae behaved similarly to those of the first rearing, and the one larva that completed development consumed three snails before pupariating on July 24. No other larvae reached the third instar. Perhaps they were unable to develop in the rearing dishes as the dead snails began to decay. Other rearings initiated from eggs were un- successful, for although many larvae began feeding on the Oxyloma snails, all died while still within the decaying bodies of the snails. Oxyloma collected in nature frequently contained larvae of T. rotundicornis. Collections of 100 snails made weekly at the Inlet Valley marsh between May and September produced 12 larvae. The first larva was found on June 13; the last, on September 26. Infestations remained at low levels (zero to three infested snails per week) throughout the summer. Although a few snails contained two larvae, only one larva was able to complete development on the original snail. The other larva either died or attacked a second Oxyloma, My rearing records indicate that T. rotundicornis larvae are predaceous but show parasitoid tendencies. They developed only on succineid snails, fed on a snail for several days before killing it, and killed only a few individuals during larval life. The larvae lack float hairs, are light colored, and have greatly reduced lobes around the posterior spriacular disc, all characters found in larvae adapted to feeding on terrestrial snails. 1996 Foote — Tetanocera Taxonomy and Biology II 163 Puparia obtained from laboratory-reared larvae always were formed away from their prey, usually on or slightly buried in the shredded peat moss. The 65 puparia collected in nature between February and May were also found apart from shells; nearly all were floating at the water surface among emergent vegetation. Because the puparia lack float hairs and do not have the posterior end upturned, they are not well adapted for floating compared to the truly aquatic species (Berg and Knutson, 1978). They probably are formed on moist soil during the late summer and are carried into open water when the marshes refill with water during the following spring. The five puparia obtained in the laboratory rearings produced adults in 12 to 15 days. Of the 65 puparia collected in nature, 38 (59%) produced adults in 12 to 18 days when brought into the heated laboratory. Reared females were seen to copulate two days after emerging, and copulation was repeated frequently during succeeding days. Mating lasted from less than ten minutes to more than two hours. Fertilized females began ovipositing within five days after emerging. An unfertilized virgin female laid 26 eggs in 18 days. Re- cords were not obtained of the number of eggs laid by laboratory-reared females, but two individuals field-collected on June 20 subsequently deposited 108 and 248 eggs each in 32 days (June 25 to July 27). Adults lived in the breeding jars up to 38 days. In central New York, the earliest seasonal record for adults was obtained on May 23; the latest record was on September 4. Adults commonly were taken throughout the summer. Probably there are at least two generations per year, as the life cycle can be completed in 56 days. No cessation of development other than that due to cold weather was observed. A puparium formed on October 5 by a larva collected on September 26 at the Inlet Valley marsh produced an adult in 14 days. That overwintering takes place in the pupal stage is indicated by the discovery of puparia containing viable pupae as early as February 9 at the Inlet Valley marsh in New York. Puparia were collected commonly during March, April, and early May. Seventeen puparia collected in the field during March and April produced ich- neumonid wasps. Six (29%) of 21 puparia collected between March 27 and April 2 at the Floral Avenue marsh gave rise to ichneumonids, and records acquired over several years indicate that the rate of parasitization varies from three to 40%. Apparently at least two species of Ichneumonidae are involved. Tetanocera spirifera Melander 1920. Annals of the Entomological Society of America, 13:330. Tetanocera spirifera is restricted to the Nearctic region, (Knutson et al., 1986), where it ranges from Manitoba west to Alaska, and south to Colorado and Idaho (Fig. 4). Adults were taken in open freshwater marshes in which water levels fluctuated greatly, although specimens were not taken in marshy habitats which became dry in summer. In southwestern Montana, adults were taken by sweeping sedges and grasses along a small, mostly unshaded stream. In Colorado, Alberta, and Alaska, adults were taken in open marshes dominated by sedges (Carex spp.). Succineid snails were abundant in all habitats examined. Rearings were initiated on June 28, 1968, from adults collected near East Creek Campground in Beaverhead National Forest in southwestern Montana. Mating was observed in the breeding jars on June 30, and the first eggs were deposited 164 Annals of Carnegie Museum VOL. 65 on July 1. Two females deposited a total of 106 eggs in 25 days either onto projecting sprigs of peat moss or affixed to the glass walls of the breeding jars. Five eggs were placed on the shells of dead Oxyloma. No clustering was noted, and eggs were scattered widely over the substrate. 1996 Foote — Tetanocera Taxonomy and Biology II 165 The incubation period lasted five to six days (n = 25) under laboratory con- ditions. Because the newly hatched larvae lacked interspiracular processes (float hairs) and had reduced lobing around the posterior spiracular disk, it was assumed that they were more terrestrial and probably fed on shoreline or land gastropods. Small individuals of genera of fingernail clams (Sphaerium), aquatic snails {Aplexa, Helisoma, Lymnaea, Phy sella), succineid snails {Oxyloma, Catinella, Succinea), land snails {Discus, Gyraulus, Zonitoides), and slugs {Deroceras) were added to the rearing dishes. After 24 hours of exposure to these molluscs, no attacks on any of the genera of fingernail clams, aquatic or land snails, or slugs had occurred. However, larvae did display interest in the genera of Succineidae and were particularly attracted to a species of Catinella. After another 24 hours, first instars were found in several specimens of that genus, but no snails of the genera Oxyloma and Succinea were invaded. Most of the larvae were embedded up to their posterior spiracles in the slit that runs between the breathing pore and foot. A few larvae were embedded in the side of the foot in front of the breathing pore, and one was embedded in the anterior end of the foot. Up to three larvae were found in each snail, although most snails were invaded by only one larva. Larvae remained in place for at least two days while the snail continued to move about the rearing dishes and even feed. However, by the third day, most of the snails had retracted into their shells and several had affixed their shells to the sides of rearing dishes. Snail death usually occurred on the third day, although one snail remained alive for four days. The larvae continued to feed for another day or so after death as scavengers on the decaying snail flesh. They then aban- doned the original snail host and attacked a second individual. Placement of the older larva differed from that of the newly hatched first instar in that the second instar placed itself on the flesh of the snail between the mantle and the shell. Death of the second infested snail occurred within one day, and larvae commonly then attacked a third or even a fourth individual. Older larvae were somewhat more catholic in their choice of prey in that a few individuals consumed snails belonging to the genus Oxyloma, however, none attacked species of nonsuccineid snails, slugs, or fingernail clams. After abandoning the final host, larvae usually buried themselves in the peat moss lining the rearing dish and remained inactive for 18 to 24 hours. The one larva that survived formed its puparium in the peat moss. The first larval stadium lasted one to two days {n = 8); the second, two to four days {n = 4); and the third, seven days {n = 1). No adult emerged from the single puparium that was formed, so the length of the pupal period remains undeter- mined. The earliest seasonal record for adults was June 10 (Pitkin, Colorado); the latest record was August 11 (Banff, Alberta). Adults have been captured during all of the summer months. No information was obtained on overwintering habits or on voltinism. Discussion How are resources, including space, food, and time, partitioned among these four species of Tetanocera? Obviously, there are differences in geographic dis- tribution of the four species. Tetanocera melanostigma, T. oxia, and T rotundo- cornis share similar geographic ranges, although the last species is somewhat more broadly distributed. In contrast, T. spirifera is strictly western and seemingly 166 Annals of Carnegie Museum VOL. 65 overlaps only T. rotundicornis in geographic occurrence. Also, the resource di- mension involving habitat is partitioned. Tetanocera melanostigma was found most commonly in wet to mesic woodlands, whereas the other three species were found in more open marshes. Food is definitely partitioned among the four species. Newly hatched larvae of T. melanostigma attack Succinea, those of T. rotundicornis prey on Oxyloma, and larvae of T. oxia and T. spirifera apparently are restricted to Catinella. Interest- ingly, the last two species that utilize a common prey occurring in open marshes have somewhat different geographic distributions (Fig. 2, 4), thus they rarely occur together. All four species seemingly have very similar phenologies and thus do not avoid competing temporally. The biology of the four Nearctic species is very similar to that reported for T. arrogans Meigen in Europe. Knutson (1963) reported that the newly hatched larvae of that species are largely restricted to Oxyloma and Succinea snails, al- though older larvae successfully fed on two other genera of land snails. An unexplored question deals with the possible competition for succineid snails that may occur, as at least 12 sciomyzid species {Sciomyza aristalis [Coquillett], S. dryomyzina Zetterstedt, S. simplex Falle’n, Pherbellia punctata maculata [Cres- son], Pteromicra anopla Steyskal, Antichaeta borealis Foote, A. testacea Melan- der, Hoplodictya spinicornis [Loew], and the four species of Tetanocera) and two species of Calliphoridae (Melanomyia obscura (Townsend) and M. ordinaria [West]) are known to prey on snails of that family. Acknowledgments Research was supported by grants from the National Science Foundation, National Institute of Allergy and Infectious Diseases, and the National Geographic Society. Appreciation is expressed to G. C. Steyskal of Gainesville, Florida, for his expert help in determining species of Tetanocera. This paper is dedicated to the fond memory of Dr. C. O. Berg, Department of Entomology at Cornell University, who inspired students around the world to pay attention to the details of fly biology and immature stages. Literature Cited Berg, C. O. 1953. Sciomyzid larva (Diptera) that feed on snails. Parasitology, 39:630-636. . 1961. Biology of snail-killing Sciomyzidae (Diptera) of North America and Europe. Ver- handlung der XI International Kongress de Entomologie, 1:197-202. Berg, C. O., and L. V. Knutson. 1978. Biology and systematics of the Sciomyzidae. Annual Review of Entomology, 23:239-258. Foote, B. A. 1996. Biology and immature stages of snail-killing flies belonging to the genus Tetan- ocera (Insecta: Diptera: Sciomyzidae). I. Introduction and life histories of predators of shoreline snails. Annals of Carnegie Museum, 65:1-12. Knutson, L. V. 1963. Biology and immature stages of snail-killing flies of Europe (Diptera: Scio- myzidae). Unpublished Ph.D. dissert., Cornell University, Ithaca, New York, 390 pp. Order No. Mic 63-4822, University Microfilms, Ann Arbor, Michigan (Dissertation Abstracts, 24:445-446). Knutson, L. V., R. E. Orth, and W. L. Murphy. 1986. Catalog of Sciomyzidae (Diptera) of America north of Mexico. Entomography, 4:1-53. ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 2, Pp. 167-179 16 May 1996 A NEW GENUS OF BEAVER (MAMMALIA: CASTORIDAE: RODENTIA) FROM THE ARIKAREEAN (OLIGOCENE) OF MONTANA AND ITS BEARING ON CASTORID PHYLOGENY William W. Korth^ Research Associate, Section of Vertebrate Paleontology Abstract The holotypes of two Arikareean castorids from Montana, Steneofiber complexus Douglass and S. hesperus Douglass were reexamined in light of recent synonymies of these species with other European and Asian species. The two Montana species are synonyms but are not synonymous with any previ- ously described species of castorid from either Europe or Asia. A new genus, Neatocastor, is named for the Montana beaver, N. hesperus. The new genus is most closely related to the otherwise Chadronian to Whitneyan-aged castorid Agnotocastor Stirton and the Hemingfordian to Barstovian Ambly castor Matthew. A combination of retained primitive characters and derived characters of the skull and dentitions of these genera unite them systematically. This group of genera represents an early offshoot of the Castoridae that is not ancestral to any other group of castorids. Introduction Douglass (1901) described two beavers from the Miocene of Montana, Steneo- fiber hesperus and S. complexus, but was uncertain of their age of occurrence. Matthew (1902:301) listed these species as “White River” in age (= Oligocene). Stirton (1935) transferred both of these species to his new genus Monosaulax and placed them questionably in the late Miocene (= Barstovian). Most recently, Xu (1994) synonymized these two species (along with four other Eurasian species) under Steneofiber hesperus and recognized them as occurring in the late Arika- reean. A detailed examination of the holotypes of S. hesperus and S. complexus has allowed for the recognition of a new genus of castorid and a detailed discus- sion of the relationships of these species. Dental nomenclature used follows that of Stirton (1935). Abbreviations for institutions are: AMNH, American Museum of Natural History; CM, Carnegie Museum of Natural History; KU, University of Kansas, Museum of Natural His- tory; USNM, National Museum of Natural History, Smithsonian Institution; YPM-PU, Princeton Collection, Yale-Peabody Museum. Systematic Paleontology Order Rodentia Bowdich, 1821 Superfamily Castoroidea Gray, 1821 Family Castoridae Gray, 1821 Neatocastor, new genus Type and Only Species. — Steneofiber hesperus Douglass, 1901. Range. — Arikareean (late Oligocene) of Montana. ’ Rochester Institute of Vertebrate Paleontology, 928 Whalen Road, Penfield, New York 14526. Submitted 28 September 1995. 167 168 Annals of Carnegie Museum VOL. 65 Diagnosis. — Castorids with elongate rostra; posterior palatine foramina within palatine bone; dorsally, nasals extend more posteriorly than premaxillaries; com- plex occlusal pattern cheek teeth (multiple fossettes preserved on worn teeth, irregularities of enamel preserved on lophs); cheek teeth subhypsodont and rooted; dP^ present; anterior face of upper and lower incisors broadly convex anteriorly; upper cheek tooth rows parallel, widely spaced; premolars subequal in size to molars in early stages of wear. Etymology. — Greek, neatos, last or utmost; Latin, castor, beaver. Discussion. — The cheek tooth patterns of Neatocastor are more complex than in most other castorids which have a simplified series of reentrant valleys. The rostrum of Neatocastor is more elongated than in other castorids, similar to the elongation seen in eutypomyids. The skull and dentition of Neatocastor most closely resemble those of the Chadronian to Whitneyan castorid Agnotocastor. The latter, however, possesses a P^, unlike any other castorid. The skull of Nea- tocastor is not flattened and broad as in palaeocastorines. Wahlert (1977) noted two primitive features of the skull that separated Agnotocastor from all other castorids: the stapedial foramen was retained in the bulla and the posterior palatine foramina were entirely within the palatine bone. In all other castorids, the stapedial foramen is lost and the posterior palatine passes through the palatine-maxillary suture more anteriorly than in Agnotocastor. The bulla of Neatocastor is not preserved but it is evident that the posterior palatine foramen is entirely within the palatine bone as in Agnotocastor. The posterior extent of the nasals dorsally on the skull of Neatocastor (more posterior than premaxillaries) is unlike any castorines, palaeocastorines, or castoroidines in which the nasals are level with or shorter than the premaxillaries. The incisors of palaeocastorines are anteriorly flattened unlike those of Neatocastor and all other castorids. The upper cheek tooth rows are parallel and widely spaced in Neatocastor unlike those of other castorids except Agnotocastor. The premolars in castorines, castoroidines, and most palaeocastorines are the largest of the cheek teeth. The premolars in Nea- tocastor are subequal in size to the molars in early stages of wear. Because the crowns of the molars taper toward their bases, in latest stages of wear the molars become shorter (anteroposteriorly) than the premolars in Neatocastor and all other castorids. Neatocastor differs from species of Agnotocastor in being larger with higher- crowned cheek teeth. Cranially, there is little difference between these two genera. Both are characterized by elongate rostra, premolars subequal in size to the molars (not enlarged), and having the maxillary tooth rows parallel rather than divergent. The complexity of the occlusal pattern of the cheek teeth of Neatocastor is quite similar to that of Agnotocastor: the upper molars retain the elongate J-shaped mesoflexus with multiple fossettes, and the lower cheek teeth preserve the irreg- ularities of the enamel along the borders of the various fossettids even in worn specimens. Agnotocastor retains a (primitive for castorids) whereas in Neato- castor a dP^ is present in juvenile individuals but is not replaced by the permanent tooth. Neatocastor hesperus (Douglass, 1901) (Fig. 1, 2; Table 1) Steneofiber complexus Douglass, 1901:249. Steneofiber hesperus Douglass, 1901:243. 1996 Korth — New Beaver from Montana 169 Monosaulax complexus (Douglass) Stirton, 1935:421. Monosaulax hesperus (Douglass) Stirton, 1935:422. "‘‘Monosaulax’’'’ complexus (Douglass) Wilson, 1960:72. "‘Monosaulax’’’’ hesperus (Douglass) Wilson, 1960:72. Steneofiber hesperus (= complexus) Douglass: Xu, 1994:85. Type Specimen. — CM 711, left mandible with complete dentition. Referred Specimen. — CM 712 (holotype of Steneofiber complexus), partial skull with incisors, dP'^- and unerupted on the right side, and associated mandibles with incisors and dP4-M3; right P4 preserved in crypt. Horizon and Locality. — Holotype from Blacktail Deer Creek, Beaverhead County, Montana, Re- ferred specimen from gray sandy layer that cuts through White River-age rocks on the west side of the Madison River nine or ten miles south of Three Forks, Gallatin County, Montana. Age. — CM 71 1 is from the Arikareean (late Oligocene) Black Tail Deer Creek fauna (see Hibbard and Keenmon, 1950). CM 712 was described from a horizon of unknown age by Douglass (1901). An age determination was not made by Douglass because of the lack of associated mammal fossils. Later, Wood (1933) described specimens of the Arikareean rhinocerotid Diceratherium armatum from the same beds but a different locality, establishing the age as Arikareean. Stirton (1935) mistakenly listed both S. hesperus and S. complexus as being questionably late Miocene (= Barstovian) in age. The age of both CM 712 and CM 711 is clearly Arikareean as noted by Xu (1994). Description. — The skull of N. hesperus (CM 712) is badly damaged and little more than the size and shape of the rostrum is preserved. The rostrum is long (upper diastema 22.7 mm in length), laterally narrow, and not very deep (dorsoventral measurement 1 1 .9 mm) compared to contempora- neous palaeocastorine beavers. The only recognizable foramina on the rostrum are the incisive foram- ina and the infraorbital foramen on the left side. The incisive foramina are long (3.9 mm) and narrow slits positioned in the center of the anterior part of the palate (diastema). The premaxillary-maxillary suture intersects the center line of the rostrum at the posterior end of the incisive foramina. The infraorbital foramen is low on the rostrum and dorsal to the posterior ends of the incisive foramina. Laterally, the infraorbital foramen is shielded by a flange of bone. Dorsally, the nasals and premaxillary bones extend posteriorly, level with the anterior margin of the orbits. The nasals extend about 1.8 mm farther posterior than the premaxillaries. A parasagittal crest arises from the posterior margin of the orbit and is directed posteriorly and slightly medially. Only the anterior part of the right parietal is preserved, so it cannot be determined whether there is a sagittal crest at the back of the skull. In Agnotocastor there is a single sagittal crest that runs the length of the cranium from a point even with the center of the orbits. The zygomatic plate is broken away on both sides of the skull and the medial orbital wall is badly crushed, eliminating any evidence of foramina or sutures. However, most of the anterior portion of the palate is preserved. The palatine-maxillary suture extends anteriorly to the level of the anterior margin of the first molars. The posterior palatine foramina are even with the center of M' within the palatine bone. The palatal surface is smooth with no indication of any grooves or furrows. The tooth rows are parallel. The ventral surface of the upper diastema is nearly flat with only a slight dorsal inclination. The upper incisors are broad with gently convex enamel faces. They appear somewhat procumbant, similar to the condition in Palaeocastor (Stirton, 1935:fig. 31). The cheek teeth are subhypsodont. P'* and the first two molars are subequal in size, M^ is slightly smaller. On the right side of the skull there is a small peg anterior to the tooth row which appears to be the root of dP^. In a maxillary specimen listed by Rasmussen (1977) as ""Monosaulax’' complexus there is no P^ associated with the adult dentition, and since dP"^ is still present in CM 712, it is most likely that the root belongs to a deciduous premolar rather than a permanent P^. DP"* is lower crowned and slightly smaller than the molars; its occlusal pattern is complex and reminiscent of the molars. There is a long paraflexus that extends lingually, and the sides of the paraflexus are irregular, not straight. A shallow hypoflexus extends from the lingual margin of the tooth anterobuccally and nearly meets the lingual end of the paraflexus. The anterior arm of the hypocone and posterior arm of the protocone fuse near the center of the tooth and extend to the buccal margin of the tooth as a thin loph (= ?mesoloph) which bisects the occlusal surface of the tooth. Anterior to the buccal end of this loph is a minute, circular fossette between the ?mesoloph and the paraflexus. Posterior to the ?mesoloph are three separate fossettes. The largest one is crescentic and forms most of the posterior border of the ?mesoloph. A smaller, ovate fossette is on the posterobuccal corner of the tooth. The last fossette is minute and is near the center line of the tooth along the posterior margin. P'* is preserved in CM 712 on the left side but still remains in the crypt. The surface consists of a 170 Annals of Carnegie Museum VOL. 65 J L I cm 1996 Korth — New Beaver from Montana 171 5 mm Fig. 2. — Dentitions of Neatocastor hesperus. A, C-G, CM 712. B, H, CM 711 (holotype). A, cross section of right Ij. B, cross section of left I,. C, right M’-M^. D, right P"* (in crypt). E, left dP4. F, left dP4-M3. G, partial right P4 (in crypt). H, left P4-M3. Fig. 1. — Skull and mandible of Neatocastor hesperus, CM 712 (holotype of ''Steneofiber complexus''). A, right lateral view of skull. B, ventral view of skull. C, dorsal view of skull. D, lateral view of left mandible. E, lateral view of right mandible. 172 Annals of Carnegie Museum VOL. 65 series of lophs that are covered with enamel that is crenulated and irregular. As in dP'^ there is a deep parafossette and shallow hypofossette that nearly meet. There is a central mesoflexus that extends lingually for nearly the entire width of the tooth. Between the mesoflexus and paraflexus is a smaller, ovate fossette along the buccal margin of the tooth. Posteriorly there is a wide (buccolingually) me- tafossette that runs the entire width of the tooth. The metafossette opens posteriorly through a shallow valley that may close in early stages of wear. All of the lophs that separate the flexi and fossettes of P^ are covered with irregular enamel that may produce any number of minute fossettes once the tooth is in occlusion and the lophs begin to wear. M' and are equal in size. As in F* the flexi and fossettes have irregular walls reflecting the enamel irregularities along these lophs. On M’, there is a long parafossette that may have opened buccally when unworn. The parafossette curves posteriorly at its lingual end, forming a “J” in shape. Similarly, the mesoflexus curves posteriorly at its lingual end, extending posteriorly to the posterior margin of the tooth. The hypoflexus is shallow and directed anterobuccally. Two other small fossettes are present along the buccal edge of the tooth. The first is the smallest and is situated between the paracone and the mesoflexus. The posterior one (metafossette) is larger and located in the postero- buccal corner of the tooth posterior to the metacone. The hypostria extends nearly to the base of the tooth on the lingual side. The mesostria is short, extending less than half the buccal height of the crown. All of the upper molars taper dorsally. In late wear this tapering is manifest in the anteropos- terior shortening of the molars. is nearly identical to M' except in a few features which may be due to a lesser amount of wear. On the hypoflexus is continuous with the paraflexus on the left side but not on the right. Also on the right, the mesoflexus is continuous with the metafossette at the lingual end. On the left side a posteriorly directed portion of the mesoflexus is closed anterior to the metafossette. A distinct meso- cone is present on both M^s. is the smallest of the cheek teeth and is in a very early stage of wear. The paraflexus and hypoflexus are confluent and the paraflexus has a shallow anterior opening at its buccal end. A distinct mesocone is present. There are four buccolingually elongate fossettes along the buccal margin of the tooth. The mandible is dorsoventrally deep, as is typical of castorids. The masseteric fossa ends anteriorly below the anterior margin of M,. The diastema is shallow, concave dorsally, and nearly as long as the length of the tooth row. The mental foramen, preserved only on CM 711, is at middepth of the mandible just anterior to the anterior margin of P4. The ascending ramus rises steeply dorsally, begin- ning at the level of M2, and a broad, shallow valley separates the tooth row from the medial border of the ascending ramus. The coronoid process is short and extends vertically, well dorsal to the occlusal surface of the cheek teeth. The articular condyle is directly posterior to the coronoid and just slightly ventral to it. The angle of the mandible is medially deflected, not in line with the ascending ramus. The lower incisor, like the upper incisor, is broad and gently convex anteriorly. As in the upper cheek teeth, the lophs of the lower cheek teeth are covered with enamel that has numerous irregular- ities. The dP4 is smaller than the molars and much lower crowned. In occlusal shape, it is narrow anteriorly and wider posteriorly. The hypoflexid is short and posteriorly directed. There is a small buccolingually elongated fossettid along the posterior edge of the tooth. The mesofossettid is split into two by a short loph (= ?mesolophid). On the right side, this fossettid is completely divided into two separate fossettids, whereas on the left side these two smaller fossettids are still joined at their lingual ends. Anteriorly, there is a crescentic parafossettid. The lower permanent premolar is preserved not only in the holotype (CM 711) but also in the crypt below dP4 on the right side of CM 712. The premolar is subequal in size to the anterior molars. Anteriorly there is a long parafossettid that extends the entire width of the tooth. It curves anteriorly at its buccal end and opens anteriorly through a shallow valley in the anterior wall of the tooth. The posterolingual end of the parafossettid appears to close off from the remainder of the fossettid in late stages of wear, isolating an additional fossettid. The mesoflexid is very shallow, and a small triangular fossettid (mesofossettid) is isolated near the center of the tooth. The hypoflexid is also shallow and split buccally by a distinct mesoconid. The metafossettid is buccolingually elongated and runs nearly the entire width of the tooth. Between the metafossettid and the mesofossettid are numerous minute fossettids that may disappear in later wear. Both the mesostriid and hypostriid extend to the base of the crown. The unworn molars of CM 712 preserve more complexity of the occlusal surface than do the more worn teeth of the holotype. The first two molars of the holotype consist of a deep mesoflexid and hypoflexid that extend at least to the centerline of the tooth, and two transversely elongate fossettids, parafossettid and metafossettid. Both the parafossettid and mesoflexid curve anteriorly at their buccal ends. Irregularities along the edges of the fossettids are remnants of the kind of enamel irregularities seen on the more unworn molars. The hypostriid extends to the base of the crown, whereas the 1996 Korth — New Beaver from Montana 173 mesostriid is very shallow on the lingual side and would be eliminated after only moderate wear. The last molar of the holotype is similar to the anterior molars but is smaller and slightly reduced poste- riorly. The less wore molars of CM 712 have the same basic pattern as those of the holotype but retain numerous minute lophules and reentrants in the fossettids. -Stilton (1935) referred both Steneofiber complexus and S. hes~ perus to the generally Barstovian-aged genus Monosaulax. Wilson (1960) noted that this was an incorrect allocation that was based only on the shape of the incisors but he did not allocate these species to any other genus, citing them only as '"Monosaulax.'' Xu (1994) listed S. complexus as a junior synonym of S. hesperus with no discussion of the comparative morphology of their respective type specimens. It does appear, however, that these two species are synonyms. The holotype of S. complexus is a young individual that retains the deciduous dentition. However, both the right permanent upper and lower premolars are pre= served in their crypts, which allows for direct comparison with the holotype of N. hesperus (CM 711). The lower premolars are essentially identical on CM 712 and CM 711, differing only in the greater complexity of the unerupted tooth in CM 712. Similarly, the lower molars of CM 712 are more complex than those of CM 711, in which the reduction in complexity is clearly due to its more advanced stage of wear. Douglass (1901), in describing both these species, cited four differences: 1) the lower incisor of S. complexus was less convex than that of S. hesperus, 2) the angle of the mandible was broadly rounded in S. complexus and acute in S. hesperus, 3) the buccal cusps of the lower cheek teeth were rounded externally in S. complexus and angular in S. hesperus, and 4) the lingual border of the lower cheek teeth of S. complexus was concave (not straight). These features do not appear to be consistent with the holotypes of these species. The anterior surface of the lower incisor is nearly identical in CM 711 and CM 712. The angle of the mandible in both specimens is broken, so the complete shape is not discernible. As for the shape of the lower cheek teeth, the differences cited by Douglass (1901) are clearly due to the greater degree of wear on CM 711. In crown height and complexity of the occlusal surface of the cheek teeth, and cross-sectional shape of the lower incisor, S. complexus is identical to S. hesperus. In size, the lower cheek teeth of the holotype of N. hesperus are only slightly larger than CM 712 (Table 1). The greatest difference between N. hesperus and species referred to Monosau- lax is the lack of complexity of the lophs of the cheek teeth and the relatively enlarged premolars of the latter. The lophs separating the fossettes (-ids) or flexi on the cheek teeth of Monosaulax species are simple, smooth, and generally straight. The lophs of the cheek teeth of N. complexus are complicated with minute ridges and crenulations. The premolars of Monosaulax are larger than any of the molars, whereas in Neatocastor the premolars are approximately equal in size to at least the first molars in early stages of wear. Xu (1994) included the European species Steneofiber jaegeri and S. depereti, as well as two Asian species, Monosaulax changpeinensis and M. tungurensis, as synonyms of S. hesperus. He cited three features that united these species and separated them from Eucastor (which he cited as a senior synonym of Monosau- lax): 1) jugal contacts the lacrimal bone and straight masseter ridge (masseteric fossa adjacent to the infraorbital foramen); 2) condyle, coronoid process, and angle of the mandible form straight line in posterior view (not aligned as in Eucastor), and 3) Eucastor has a cheek tooth pattern with fewer flexi. 174 Annals of Carnegie Museum VOL. 65 Table 1 . — Dental measurements of Neatocastor hespems. Abbreviations: a-p, anteroposterior length; tr, maximum transverse width; * indicates estimated dimensions of permanent premolars in crypt. CM 711 CM 712 Right Left I, a-p 3.68 3.62 3.49 tr 3.71 3.50 3.41 dP4 a-p 4.28 3.50 tr 2.99 3.12 P4 a-p 5.10 4.50* tr 4.40 3.30* M, a-p 3.80 3.22 3.47 tr 4.30 4.15 3.94 a-p 4.00 3.54 3.49 tr 4.05 3.82 4.04 M3 a-p 3.80 3.21 2.96 tr 3.60 3.30 2.96 V a-p 3.22 3.40 tr 3.69 3.71 dP^ a-p 3.40 tr 3.85 P4 a-p 3.80* tr 3.80* M' a-p 3.40 3.20 tr 4.10 4.00 M2 a-p 3.00 2.95 tr 4.00 3.90 M2 a-p 2.30 2.50 tr 3.50 3.65 The use of these characters is not consistent with the record of ''Steneofiber hesperus.’" The only known skull of N. Hesperus is CM 712, and it is damaged in such a way that these particular cranial features are not preserved at all. Sim- ilarly, of the three lower jaws of the American beaver, none has all three processes of the mandible preserved (condyle, coronoid, angle), so their alignment is im- possible to determine. The right mandible of CM 712 preserves a part of the angle and it is more medial than the remainder of the ascending ramus, at least hinting that the posterior mandibular processes are unlike those of Steneofiber. The skull of N. Hesperus also preserves the root of dP^ on the right side. In Steneofiber, as in all other castorids except Agnotocastor, this tooth is lost. As for the number of flexi on the cheek teeth, Xu (1994) may be correct in number, but his discussion and figures do not detail the great amount of complexity in the pattern of N. Hesperus teeth. The cheek teeth of Steneofiber, as well as Monosaulax and Eu~ castor, are a series of reentrant valleys separated by smooth, straight lophs. The lophs of the cheek teeth of Neatocastor are far more complicated with numerous lophules and tuberosities that are preserved even in worn teeth. Some cranial features that Xu (1994) did not consider also preclude Neatocastor from being a species of Steneofiber. The most obvious feature of the skull of N. Hesperus is the elongation of the rostrum, which it shares with Agnotocastor. In the type of N. Hesperus the upper diastema is 175% the length of the upper tooth row. In Agnotocastor praetereadens this ratio is 150%. In the skulls of Steneofiber figured by Stirton (1935:fig. 14-18) the ratio of the diastemal length to the tooth row ranges from 100 to 125%. The incisive foramen is also relatively larger in 1996 Korth — New Beaver from Montana 175 Neatocastor and Agnotocastor. The ratio of the length of the incisive foramen to the length of the upper diastema in Agnotocastor and Neatocastor is 0.39 and 0.33, respectively. In the figured skulls of Steneofiber this ratio is 0.23-0.25, much shorter. The more posterior extent of the nasals on the dorsal skull of Neatocastor is not present in skulls of Steneofiber where the premaxillary and nasal bones extend posteriorly to the same level as in other castorids. Wahlert (1977) separated Agnotocastor from all other castorids based on two primitive characters: presence of a stapedial foramen on the bulla (lost in other castorids), and posterior palatine foramina entirely within the palatine bone (along palatine-maxillary suture in other castorids). In a skull of S. castorinus (AMNH 11010) are two pairs of posterior palatine foramina, a large pair along the pala- tine-maxillary suture (typical of castorines), and a minute pair posterior to them in the palatine bone. The skull of N. hesperus preserves only the anterior portion of the palatine bone on the palate and the entire palatine-maxillary suture is distinguishable. The posterior palatine foramina are posterior to the suture, the condition in Agnotocastor and unlike Steneofiber and all other castorines. Clearly, N. hesperus is not a species of Steneofiber. The European species that Xu (1994) listed as synonyms of ''Steneofiber'' hesperus cannot be referred to Neatocastor. Not only do the cheek teeth of these species lack the complexity of N. hesperus, but they are markedly larger. Wilson (1960) noted that a species he referred to Anchitheriomys from the Hemingfordian of Colorado was similar in size to both Steneofiber depereti and S. jaegeri. This Colorado species is 30% larger than N. hesperus (Wilson, 1960:67). Stehlin and Schaub (1951:fig. 80) figured an upper molar of S. jaegeri that is nearly 40% larger than the upper molars of N. hesperus. Stirton (1935:415) listed dental mea- surements of "Steneofiber" depereti that are nearly twice as large as the dental measurements of N. hesperus (Table 1). Also, no species of Steneofiber is known to have a dP^ as in Neatocastor. These European species clearly are not synonyms of N. hesperus and should be retained in Steneofiber as originally designated. The Asian species listed as synonyms of "S." hesperus by Xu (1994), likewise, cannot be referred to this species. The early Miocene Monosaulax changpeiensis differs from the Montana species in a number of dental features. The P4 of M. changpeiensis is characterized by a deep mesoflexid that bends anteriorly, fusing with a parafossettid that is closed anteriorly (Li, 1962:fig. 1). In a worn P4 the flexid is closed lingually but the fossettid maintains its long and curved shape. The P4 of N. hesperus has an open anterior flexid, and there is a loph that separates the mesoflexid from the parafossettid. Another feature that separates the P4 of M. changpeiensis from N. complexus is the morphology of the metafossettid. In the unworn specimen of M. changpeiensis are two small fossettids, and in the worn specimen is a single, small fossettid that is obliquely oriented. The P4 of N. hesperus has only a single metafossettid that is long and buccolingually oriented. The lophs of the cheek teeth of N. hesperus preserve a number of minor irregu- larities not present in the teeth of M. changpeiensis. The mesostriids on the molars of M. changpeiensis also appear to extend lower on the crowns than those of N. hesperus (Li, 1962:pl. 1, fig. Ic). The later Miocene Monosaulax tungurensis is known from a mandible that retains dP4 along with the molars as does the type of "S." complexus (CM 712). The molars of M. tungurensis have lophs that are simpler (lack irregularities) than the molars of N. hesperus. The dP4 of M. tungurensis consists of two simple fossettids (para- and metafossettid) and a simple mesoflexid and hypoflexid (Li, 176 Annals of Carnegie Museum VOL. 65 1963:pL 1). The dP4S of N. hesperus (CM 712) are worn to the same level as that of the holotype of M. tungurensis but are far more complex. The parafossettid is elongated and bent at a right angle near its center. The mesoflexid is divided into two by what appears to be a mesolophid. The metafossettid is similar to that of the dP4 of M. tungurensis in shape and orientation, but preserves irregular sides, reflecting the generally more complex nature of the lophs of the tooth. There is a steep wall that extends to about the middle of the mandible anterior to P4 in M. tungurensis. The diastema of the mandibles of N. hesperus are shallow and gently convex upward. Clearly, these Asian species cannot be referred to N. hesperus as suggested by Xu (1994). Scott (1893) named Steneofiber montanus from the Arikareean of Montana based on a maxilla, some lower cheek teeth, and some associated postcranial bones (YPM-PU 10456). Matthew (1902) figured the teeth of S. montanus and noted that it had more complex teeth than Palaeocastor (= Steneofiber) nebras- kensis and was similar in size to S. hesperus. He did not synonymize S. hesperus and S. montanus because they represented individuals of very different ages {S. hesperus young, S. montanus old) but noted their great similarity. Later, Stirton (1935) included Scott’s species as lAgnotocastor montanus, but cited the holotype as being lost. Both Stirton and Matthew recognized the complexity of the occlusal pattern of the cheek teeth of S. montanus, a characteristic of Neatocastor hesperus. It is possible that S. montanus and N. hesperus are synonyms. The holotype of the former (YPM-PU 10456) does exist but the dental elements of the specimen have been lost, so no direct comparison can be made. In the future, if a larger sample of N. hesperus or S. montanus is discovered, and individuals of the same age as the holotype of the other species are available, then this comparison can be made. If these species do prove synonymous, then the species would have to be referred to Neatocastor montanus (Scott). Systematic Position of Neatocastor The most recent classification of the Castoridae listed three subfamilies: Cas- torinae, Palaeocastorinae, and Castoroidinae (Korth, 1994). The genus Agnoto- castor was excluded from any of these subfamilies because of its possession of primitive characters of the skull and dentition. Wahlert (1977) noted that the skull of Agnotocastor was more primitive than all other castorids. Wahlert cited no derived characters of Agnotocastor and left it as a sister group to all other castorids (Wahlert, 1977:fig. 3). Dentally, Agnotocastor retains a P^ which is lost in all other castorids, another primitive retention of the genus (Stirton, 1935). The char- acters that unite Agnotocastor and Neatocastor, along with the primitive features of the skull (position of the posterior palatine foramina, at least dP^ retained), are the elongation of the rostrum, larger incisive foramina, more posterior extent of the nasals, and the greater complexity of the occlusal surface of the cheek teeth. These specializations, as well as the primitively retained characters of the skull cited by Wahlert (1977), are also shared with the problematical genus Amblycastor and the Eutypomyidae. The Eutypomyidae have brachydont to submesodont cheek teeth (lower- crowned than either Agnotocastor or Neatocastor) with complex occlusal patterns, the last premolars subequal in size to the molars, the upper tooth rows parallel, and P^ or dP^ retained; these dental features are shared by Agnotocastor and Neatocastor. Eutypomyids have elongate rostra, nasals that extend more poster!- 1996 Korth — New Beaver from Montana 177 orly than the premaxillaries dorsally, and retain the stapedial foramen in the bulla (only definitely known in Agnotocastor among the former genera) and the pos- terior palatine foramen is entirely within the palatine bone, features shared by Agnotocastor and Neatocastor. The only feature used by Wahlert (1977) to sep- arate the Eutypomyidae from the Castoridae was the occlusal pattern of the cheek teeth. The cheek teeth of Agnotocastor and Neatocastor, while not being as com- plex as that of Eutypomys, are more complex than other castorids. Both Agno- tocastor and Neatocastor have higher-crowned cheek teeth than eutypomyids, and the crowns of the molars taper towards their roots, typical of all castorids. In later stages of wear, because of this tapering of the crowns, the molars are reduced in length (anteroposteriorly) making the premolars appear larger. This “narrowing” of the molars is not present in Eutypomys. The problematical Hemingfordian and Barstovian genus Amblycastor (possibly synonymous with Anchitheriomys) also shares the elongated rostrum, posterior extension of the nasals, and complex occlusal patterns of the cheek teeth with Agnotocastor, Neatocastor, and eutypomyids (Wilson, 1960; Voorhies, 1990). Korth (1994) referred Amblycastor and Anchitheriomys to the Eutypomyidae based on these characters, whereas previously these latter two genera had been considered castorids (Stirton, 1935; Simpson, 1945). Amblycastor has cheek teeth that are at least as high-crowned as those of Neatocastor. Voorhies (1990) dem- onstrated that there was evidence for the presence of a dP^ Anchitheriomys {= Amblycastor) fluminis. A nearly complete skull of this latter species from the Barstovian of Nebraska (USNM 298914, not yet described fully) has the posterior palatine foramina on this specimen entirely within the palatine bone as in Ag- notocastor, Neatocastor, and eutypomyids. The cheek teeth of Amblycastor and Anchitheriomys have several castorid characteristics not present in eutypomyids: 1) premolars are the largest of the cheek teeth; 2) crowns of the molars taper toward their bases (also present in Agnotocastor and Neatocastor)', and 3) the tooth rows diverge posteriorly, although not as strongly as in other castorids (for descriptions of the skull and dentition of Amblycastor and Anchitheriomys see Stirton, 1935; Wilson, 1960; Voorhies, 1990). The incisors of Amblycastor have been described as procumbent (extending anterior to the premaxillaries), similar to those of Neatocastor and Agnotocastor (Wood, 1937:pl XXVIII, fig. 1), and distinct from those of castorines such as Steneofiber and Castor (Stirton, 1935: fig. 15, 16; Voorhies, 1990). Agnotocastor, Neatocastor, and Amblycastor retain a primitive cranial feature lost in all other castorids: posterior palatine foramina entirely within palatine bone. Other features of the skull shared by these genera are the relative elongation of the rostrum, the nasals extending farther posterior than the premaxillaries dorsally, and procumbent incisors. These features, along with the complexity of the cheek teeth, unite these three genera. Within this group of genera, Neatocastor is clearly intermediate in dental characters. The cheek teeth are higher crowned than in Agnotocastor and lower crowned than in Amblycastor (and Anchitheriomys), and the degree of complexity of the occlusal surface of the cheek teeth of Neatocastor is less than in Agnotocastor and more complex than in Amblycastor. However, the Asian species of Anchitheriomys have occlusal patterns of the cheek teeth as complex as those of eutypomyids. If elongation of the rostrum and complex occlusal pattern of the cheek teeth are considered derived characters, then Agnotocastor, Neatocastor, and Ambly- castor represent a distinct clade among castoroids closely related to the Eutypo- 178 Annals of Carnegie Museum VOL. 65 Fig. 3. — Cladogram of proposed relationships. Explanation of nodes: 1, Castoroidea: sciuromorphy; uniserial enamel of incisors; interorbital foramen posterior to optic foramen; sphenopalatine foramen surrounded by maxillary; posterior maxillary foramen enclosed; dorsal palatine foramen in maxillary or along orbitosphenoid-maxillary suture (see Wahlert, 1977:fig. 3). 2, Castoridae: intermediate to large-size rodents; cheek teeth at least mesodont; castorid crown pattern of cheek teeth (= crowns of molars taper towards base; occlusal patterns of cheek teeth series of lophs and reentrant valleys [cusps indistinguishable], patterns become simpler with wear). 3, (unnamed taxon): elongate rostrum; occlusal pattern of cheek teeth complex; upper incisors procumbent; nasals extend further posteriorly than maxillaries dorsally. 4, Palaeocastorinae, Castorinae, Castoroidinae: and dP^ lost; stapedial foramen lost; posterior palatine foramen within palatine-maxillary suture; upper tooth rows strongly diverge posteriorly; premolars larger than molars. myidae. However, if these features are considered primitive for the Castoroidea or convergent with eutypomyids, then these genera represent a primitive group of castorids. The characteristic tapering of the molars that results in a narrowing with age is clearly a dental character that is shared by Agnotocastor, Neatocastor, Amblycastor, and all other castorids. Agnotocastor, Neatocastor, and Anchitheriomys represent a distinct subfamily of beavers. A name is being proposed formally elsewhere along with a complete description of the skull of Anchitheriomys (Korth and Emry, in press). The rela- tionships of this group of genera are presented in Fig. 3. Acknowledgments The specimens described above were loaned by permission of Dr. M. R. Dawson (CM). Assistance in locating the specimens, preparation, and helpful discussion of the locality data of the specimens 1996 Korth — New Beaver from Montana 179 was provided by A. Tabrum (CM). Drs. J. H. Wahlert (AMMH) and R. J. Emry (USNM) assisted with discussions of cranial anatomy of beavers. Camera and photographic equipment were provided by Dr. J. A. Massare of the State University of New York, College at Brockport. Literature Cited Douglass, E. 1901. Fossil Mammalia of the White River beds of Montana. Transactions of the American Philosophical Society, 20:237-279. Hibbard, C. W., and K. A. Keenmon. 1950. New evidence of the Lower Miocene age of the Blacktail Deer Creek Formation in Montana. Contributions to the Museum of Paleontology, University of Michigan, 8:193-204. Korth, W. W. 1994. The Tertiary Record of Rodents in North America. Plenum Press, New York, 318 pp. Korth, W. W., and R. J. Emry. In press. The skull of Anchitheriomys and a new subfamily of beavers (Castoridae, Rodentia). Journal of Paleontology. Li, C. K. 1962. A Tertiary beaver from Changpei, Hopei Province. Vertebrata PalAsiatica, 6:72-77. . 1963. A new species of Monosaulax from Tung Gur Miocene, Inner Mongolia. Vertebrata PalAsiatica, 7:240-243. Matthew, W. D. 1902. A horned rodent from the Colorado Miocene, with a revision of the mylagauli, beavers, and hares of the American Tertiary. Bulletin of the American Museum of Natural History, 16:291-310. Rasmussen, D. L. 1977. Geology and mammalian paleontology of the Oligocene-Miocene Cabbage Patch Formation, central-western Montana. Unpublished Ph.D. dissert.. University of Kansas, 775 pp. Scott, W. B. 1893. The mammals of the Deep River beds. American Naturalist, 27:680. Simpson, G. G. 1945. The principles of classificatoin and a classification of mammals. Bulletin of the American Museum of Natural History, 85:1—350. Stehlin, H. G., and S. Schaub. 1951. Die Trigonodontie der simplicidentaten Nager. Schweizerische Palaontologische Abhandlungen, 67:1-385. Stirton, R. a. 1935. A review of the Tertiary beavers. University of California Publications in the Geological Sciences, 23:391-458. Voorhies, M. R. 1990. Vertebrate paleontology of the proposed Norden Reservoir area. Brown, Cherry and Keya Paha counties, Nebraska. Technical Report 82-09, Division of Archeological Research, University of Nebraska-Lincoln, 731 pp. Wahlert, J. H. 1977. Cranial foramina and relationships of Eutypomys (Rodentia, Eutypomyidae). American Museum Novitates, 2626:1—8. Wilson, R. W. 1960. Early Miocene rodents and insectivores from northeastern Colorado. University of Kansas Paleontological Contributions, Vertebrata, 7:1-92. Wood, A. E. 1937. The mammalian fauna of the White River Oligocene. Part II. Rodentia. Trans- actions of the American Philosophical Society, 28:155-269. Wood, H. E., II. 1933. A fossil rhinoceros (Diceratherium armatum Marsh) from Gallatin County, Montana. Proceedings of the United States National Museum, 82:1-4. Xu, X. 1994. Evolution of Chinese Castoridae. Pp. 77-98, in Rodent and Lagomorph Families of Asian Origins and Diversification (Y. Tomida, C. Li, and T. Setoguchi, eds.). National Science Museum, Tokyo, Science Museum Monographs no. 8. - i- ■ BL - 's: '.-. W' .■'f , E . Ac vv ''u i • ;.V(a' * .' 'i si ^ f ^•: ' '-si!Sfh_ y: ' » jJ s.iilI*TI>l .'-•^.T .. -Jtrr ' ;w .fT^.v- . * I * f'i ' a Bp: ‘' V;.-. ^ •;■■' '■ n'fc. «,J»Si.,,..i>„-“' ■■-■■''■;'^(a»*v^ . ., • . ,.-' 1 r* •'. ‘ f‘J. '.nt3#l3 ^ . , . ■ V A- •^ • ' :/y ^ w-l *«pI/^037 1 :■.' ■ - f‘v v.if . :'SJ mL:,:.. r-r' - '» -= '• ^ ' ' , It '-'» ‘;’ ‘ »-'--fU^-4i|l ■ . ■ - ^ 5! ii.i r: **'**-’^' ♦iVi rf *!.-/ nr* ' . -.^ - ,, .,; •■\v |S?;v.i.jli^ a*«a ■: •- 1 '.. :««ip'.-i ^ JT- Rl «-e- ' - 'U . V' ‘*i ■ • 0f* |,f(,;'*WSi.’|F 1 la • . *li fi?! iT3 ■ - : ii ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 2, Pp. 181-182 16 May 1996 REVIEW AUSTRALIAN BEETLES. Lawrence, J. F., and E. B. Britton. 1994. Mel- bourne University Press, Carlton, x + 192 pp. ISBN 0-522-84519-3. $44.95. The much-anticipated second edition of Insects of Australia (CSIRO, 1991) replaced and modernized the first edition of 1970. There were many improvements in the new version, not the least of which was the chapter on beetles. The new Coleoptera classification presented was based on detailed study of both larval and adult characters, and was state-of-the-art. However, the second edition had been in the planning and production phase long before its publication in 1991, and therefore contained some information which was obsolete or otherwise faulty. This spinoff volume on Australian beetles, while very similar to the chapter on beetles in the new Insects of Australia, was published to correct some of these problems. The 35-page introduction to the book is much more than a simple preface, and is divided into several sections: fossil history, habitats of Australian beetles, col- lecting methods, anatomy of adults, immature stages, biology, and special bio- geographic features of Australian beetles. All are treated extensively, and the section on adult anatomy may serve as a reference for the correct use of terms for structures of adult beetles. The next major part of the book is devoted to the classification and keys to both adults and larvae of Australian beetles. The classification lists all recognized families of beetles, including those not found in Australia, with the number of Australian species (both described and known undescribed) for each. A total of 28,300 species is enumerated in 117 families, with 43 families unknown in Aus- tralia. A comparison between this classification and the one in Insects of Australia reveals a few differences of varying importance. For example, the Eucinetiformia has been changed to Scirtiformia, Rhizophagidae to Monotomidae, and Cephal- oidae to Stenotrachelidae. Perimylopidae, Boridae, and Pyrochroidae are added to the Australian fauna, while Chalcodryidae, Pedilidae, and Allocorynidae are re- moved. A brief explanation of the classification is given, with comparisons made to that of Crow son. The somewhat intimidating key to adults uses 255 couplets, with many families keying out in a number of places, e.g., Tenebrionidae in eight places, and Salpin- gidae in seven. The larval key is longer, with 266 couplets. Both keys are not strictly dichotomous; some “couplets” are actually triplets or quadruplets. This design is an artifact of using the program DELTA (Description Language for Taxonomy) to store the data on which the keys were based, and creates no prac- tical difficulties in using the keys. The book’s largest part is devoted to the actual treatments of Australian beetle families. Provided for each are diagnostic, abbreviated, descriptions of both larval and adult stages. The names of important structural features are abbreviated, which, although saving space, require some page flipping to and from the list of abbreviations, until they are memorized. After this diagnosis, there is a general discussion of structural peculiarities of the family, included Australian taxa and their classification within the family, a brief account of the natural history, and 181 182 Annals of Carnegie Museum VOL. 65 pertinent references. Within this section, the illustrative material is truly excellent. Each family has at least one representative adult, and for many, also the larva, illustrated. There are 16 plates including scanning electron micrographs of larval and adult structures, and some color paintings and photographs of adult beetles. Within some of the larger families (e.g., Carabidae, Staphylinidae, Scarabaeidae, Cerambycidae, and Chrysomelidae), separate keys are given for subfamilies or tribes. While this book was written to treat the Australian beetle fauna extensively, its value goes far beyond Australia. It is immediately obvious to the reader that an immense amount of time and many thousands of specimens have gone into the production of this book. The combination of the authoritative text, inclusion of both larval and adult stages, and the superb illustrations make the work a necessity for all serious students of beetle systematics, and the affordable price tag is a welcome change! DARREN A. POLLOCK, Rea Postdoctoral Fellow, Section of Invertebrate Zo- ology, r. / 'i v > M' ■%' / ■r % - ■t •I* . V- L INSTRUCTIONS FOR AUTHORS ANNALS OF CARNEGIE MUSEUM consist of con- tributions to the earth sciences, life sciences and anthro- pology, in 30 by 46 picas format (127 by 195 mm or 5 by 7% inches). Submit all manuscripts to the Office of Scientific Publications. Authors should give particular at- tention to scientific content, format, and general style for the ANNALS. Manuscripts that do not conform to the style of the ANNALS will be returned to the author immediately. Every manuscript will be peer reviewed by at least two outside persons. Authors will be asked to subsidize, if funds are available, any or all costs of publication (approximately $ 100/page printed). Manuscript style. — Articles should include the follow- ing items in this order: title page, abstract, text (with desired headings), acknowledgments, literature cited, ta- bles, figure captions, and copies of illustrations. All manuscripts must be typed double-spaced on standard 8% by 1 1 inch white bond paper, with at least one inch margins all around, and submitted in triplicate— -Sin orig- inal for the editors, and two review copies. All pages should be numbered, including tables, literature cited, and the list of figure captions. Only correspondence qual- ity or better dot matrix printouts will be accepted; draft mode is unacceptable. A fixed-space font (e.g.. Courier) is preferred. Title Page. The title, on a separate title page should be brief, include the animal or plant group involved, and appear two to three inches below the top margin. Include the authors name(s) and the affiliations of non-CMNH authors. In the case of multiple authorship indicate the address to which proofs should be sent. Abstracts. Abstracts should be short, double-spaced and substantive and included at the head of the first page of text. Text. Do not right justify text or break (hyphenate) a word at the end of a line. Footnotes and acknowledg- ments as footnotes are unacceptable. All text is double- spaced. Tables and figure legends. The list of figure legends and each table should be typed consecutively on indi- vidual pages separate from the text. Tables must be dou- ble-spaced throughout with no vertical lines. Abbreviations. Refer to the CBE Style Manual for cor- rect abbreviations. Mammalian dentition: use capital let- ters (I, C, P, M, D, for incisor, canine, premolar, molar, deciduous, respectively) and superscript/subscript num- bers (M^, P3) to designate upper and lower teeth. Measurements. Metric units should be used, if possi- ble. Literature Cited. All references in text must appear in the Literature Cited section. The data (author, date and page) in both citations must agree. Do not abbreviate the titles of periodicals or serials. The following style, double-spaced, should be used in Literature Cited: 1) Two authors in an institutional series: Matthew, W. D., and W. Granger. 1923. The fauna of the Houldjin Gravels. American Mu- seum of Natural History Novitates, no. 97: 1-6. 2) Same authors repeated — use three-em dash: . 1923. The fauna of the Ardyn Obo For- mation. American Museum of Natural History Novitates, no. 98:1-5. 3) Same authors plus a third author — repeat all authors: Matthew, W. D., W. Granger, and G. G. Simpson. 1928. Paleocene multituberculates from Mongolia. American Museum of Natural History Novitates, no. 331:1-4. 4) Chapter in an edited volume: Rausch, R. L. 1963. A review of the distribution of Holarctic mammals. Pp. 29-43, in Pa- cific Basin Biography (J. L. Gressitt, ed.). Bishop Museum Press, Honolulu, Ha- waii. 5) Unpublished dissertation: Smith, J. P. 1976. Review of Eocene mammals. Unpublished Ph.D. dissert.. University of California, Berkeley. 6) Book: White, M. J. D. 1961. The Chromosomes. Me- thuen and Co., Ltd., London. 7) Journal articles with usual volume and issue number: Anderson, W. I. 1969. Lower Mississippian con- odonts from northern Iowa. Journal of Paleontology, 43(4):9 16-928. Illustrations. All illustrations will be called figures, and are to be numbered in Arabic numerals. Three sets of illustrations are required, one (original artwork) for reproduction, two for reviewers. 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ISSN 0097-4463 AS ANNALS d/ CARNEGIE MUSEUM THE CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213 VOLUME 65 30 AUGUST 1996 NUMBER 3 CONTENTS ARTICLES The taxa of fossil Mollusca introduced by Hermann von Ihering Juan Jose Parodiz 183 Two new species of Xyris from Cuba Armando Urquiola Cruz and Robert Krai 297 A new plesiadapiform (Mammalia: Plesiadapiformes) from the early Eocene of the Bighorn Basin, Wyoming Kenneth D. Rose and Thomas M. Bown 305 i^b'RARltS* Editors, ANNALS, BULLETIN and SPECIAL PUBLICATIONS: J. L. Carter, Ph.D B. C. Livezey, Ph.D D. R. Watters, Ph.D M. A. Schmidt, ELS, Assistant Editor Manuscripts, subscriptions, orders for individual numbers, and changes of address should be sent to: Office of Scientific Publications Carnegie Museum of Natural History 4400 Forbes Avenue Pittsburgh, PA 15213-4080 Phone (412) 622-3287 Fax (412) 622-8837 E-mail: schmidtm@clp2.clpgh.org ANNALS OF CARNEGIE MUSEUM is published quarterly by The Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213-4080, by the authority of the Board of Trustees of Carnegie Institute. © 1996 Carnegie Institute. THE CARNEGIE MUSEUM OF NATURAL HISTORY 0 This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 3, Pp. 183-296 30 August 1996 THE TAXA OF FOSSIL MOLLUSCA INTRODUCED BY HERMANN VON IHERING Juan Jose Parodiz Curator Emeritus, Section of Invertebrate Zoology Abstract Between 1896 and 1921 Hermann von Ihering described or named 352 taxa of fossil molluscs, from genera to subspecies, and many “varieties.” Most of the taxa were from southern Argentina (Pata- gonia), but some were from Chile, Uruguay, and Brazil. When combined with other species described by Arnold E. Ortmann in 1902, these fossils constitute the basis of our knowledge of the Tertiary malacofauna of the region. Ihering’s collection was acquired in 1920 by the Museo Argentine de Ciencias Naturales, and during the 1940s the author organized and catalogued the collection of more than 900 lots. This paper deals with Ihering’s taxa and: 1) updates the taxonomic nomenclature for both genera and species; 2) indicates the correct type localities, many of which were uncertain at the time of description; 3) clarifies the stratigraphic position and geologic age of the deposits where the fossils were found; and 4) reconsiders some of Ihering’s conclusions about the origin and distribution of the fauna in relation to climatic changes in the region during the Tertiary. Ihering’s stratigraphy overestimated the age of deposits. There were no intercalations between the Cretaceous and Eocene; Paleocene species were believed to be Cretaceous; marine formations of the Miocene were dated as Eocene, and so on. He also subdivided formations in order to indicate that parts of them were older. The relative ages of strata were based on the proportion of extinct species versus surviving ones. When synonyms are recognized, and many infrasubspecific names eliminated as nontaxonomic variations, such estimation is untenable. Most of the mid-Tertiary species belong to genera that are represented in the Recent, and from zones warmer than the present climate in the Patagonian and Magellanian regions. Climatic changes were not local phenomena but the consequence of plate tectonics. Movements of plates not only changed their location, but also the orientation of the masses of land by clockwise rotation. The plates, formerly separated, united into a new continent, and rotation reversed to counterclockwise bringing them into their present position. Ihering considered them a single, unique mass, and named them Archhelenis (the northeastern part including Brasilia) and Archiplata (the southwestern part). Although land-bridge theories have been replaced by interpretations from plate tectonics, Ihering’s nomenclature for the land masses is still useful and valid. New combinations are proposed for 32 species, and lectotypes are designated for Mactra indistincta Ihering (1897), Neoinoceramus ameghinoi Ihering (1902), Nucula tricesima Ihering (1897), Phacoides rocana Ihering (1907), Panopea thomasi Ihering (1914), and Potamides patagonensis Ihering (1897). Aturia (Sphenaturia) brueggeni Ihering, 1921, is selected as the type species of Sphenaturia Ihering, 1921 (Nautilidae). Introduction Between 1896 and 1921 Hermann von Ihering introduced 352 taxa of fossil Mollusca from the early Tertiary to the Pleistocene of southern South America, including genera, subgenera, species, subspecies, and infrasubspecific taxa. I had the opportunity, in 1940, of organizing this material, verifying the data, and cat- aloguing the more than 900 lots that constitute Ihering’s collection. The present work attempts to update taxonomic nomenclature and synonymy, to clarify the stratigraphic position of strata where the fossils were found, and to indicate the correct type localities. Submitted 29 January 1990. 183 184 Annals of Carnegie Museum VOL. 65 Fig. 1. — Hermann von Ihering: born 9 October 1850, died 25 February 1930. Hermann von Ihering (Fig. 1) was born at Kiel, Germany, on October 9, 1850. He studied at Giessen, Berlin, and Gottingen, and was graduated in medicine and natural sciences. As a young naturalist he went to Italy and practiced at the Naples Zoological Station and at Messina to the south. When he returned to Germany he became an assistant to Professor C. Clauss in Gottingen, was “Privat Dozent” at Erlangen, and finally worked in Leipzig with Professor R. Leuckart. Hermann intended to make his home in the city of Henen (Geissen) by marrying a bourgeois girl, to which his father would not consent. At age 30 he was required to marry. 1996 Parodiz — Ihering Mollusca 185 instead, an aristocratic young lady, Anne Maria Clare von Bozel. He projected a trip to South America and his father demanded that, upon his return, he accept a promised professorship at the University of Warsaw in Poland. However, Ihering never took the position. He was already a well-known zoologist when, in 1880, he took permanent residence in Brazil, where he lived for 40 years. He was appointed “Naturalist” of the Museu Nacional at Rio de Janeiro, and later he assumed the responsibility of organizing a new museum in Sao Paulo, State of Rio Grande do Sul. The Museu Paulista was inaugurated under his direction. His bibliography of this period was considerable, in different branches of natural sciences, published in European journals, and also in his own Revista do Museu Paulista. In 1920 he was at the Museu Catharinense, in Florianopolis, Santa Ca- tharina; by that time he had more than 300 scientific publications to his credit. His greatest dedication was to the field of malacology, not only on the taxonomy of living and fossil Mollusca, but also in anatomical research, especially on the nervous system. As early as 1876 he had defined and created the class Amphi- neura, and recognized the affinities of the Aplacophora to the chitons. His nu- merous publications on molluscan paleontology, together with A. E. Ortmann’s (1902) in the Reports of the Princeton Expedition to Patagonia, laid the foundation of our knowledge of the Tertiary malacofauna of southern South America. Iher- ing’s zoogeographic and paleogeographic studies were epitomized in “Archhe- lenis und Archinotis” published in Leipzig in 1907. While residing in Sao Paulo, Ihering was in permanent correspondence with the Argentine paleontologist Florentino Ameghino, from whom he received the fossil materials that his brother, Carlos Ameghino, collected during 18 years of arduous travels in Patagonia. Carlos Ameghino also provided the geological in- formation on which Florentino Ameghino based the dating of his stratigraphic series which Ihering followed. Those views, however, went through considerable modification by the middle of this century. He had supported, as had other pa- leogeographers of his time (including Ortmann), the theories of land bridges ex- isting between Africa and South America prior to the Neogene (Ihering, 1900). Current tectonic concepts provide a better explanation for such paleogeographic relationships, but Ihering’s nomenclature, given for land masses at the beginning of the Tertiary, is still valid. A catalogue of the Ihering Collection (Ihering, 1914) was printed when von Ihering left the Museu Paulista in 1914 and took the collection with him (he considered the collection to be his own property). About 1919 the collection was purchased from Ihering by the Buenos Aires Museum. Each lot listed in the published 1914 catalogue was subsequently assigned a number from the Catalogue of Fossil Mollusca in the Museo Argentino de Ciencias Naturales. The numbers preceded by # in the following list are the previously unpublished numbers from that catalogue, which began with the Ihering Collection. A record of those num- bers exists as annotations by Parodiz in a copy of Ihering’s catalogue (1914) in the library of Carnegie Museum of Natural History (CMNH). All lots and type material were checked by Parodiz in 1940, and during the preparation of this paper in 1987 they were checked again. During his residence in Brazil, Ihering made frequent trips to the La Plata region, keeping close connection with his collaborators. His last visit to Buenos Aires was in 1919, and it was during that period that his collection of fossils was acquired for the National Museum in that city. When Ihering returned to Europe at the age of 70, he stayed again for a while 186 Annals of Carnegie Museum VOL. 65 at the Naples Zoological Station. Finally he settled in Budingen, and married his second wife Mela Buff, the same person his father had not permitted him to marry half a century before. He died on February 25, 1930. Although Ihering’s fossil collection was founded mainly on Ameghino’s spec- imens, he also received much material from other collectors, geologists, and ma- lacologists. Among those who contributed to the collection were A. E. Ortmann, M. Cossmann, R. A. Philippi, R. Hauthal, S. Roth, and his envoy to Patagonia in 1895, J. Bicego. The contents of this paper were presented to the Tenth International Congress of Malacology held during August 1989 in Tubingen, Germany. Methods For the taxa listed alphabetically in this paper, the original name is given first followed by references. Then the current combination or senior synonym is given after “ = ” if differing from the original name, followed in some cases by junior synonyms. The family name on the right side of the citation corresponds to place- ment of the genus in the current combination. Holotypes are indicated only when the description was based on an original lot consisting of a single specimen. All other type materials are indicated as syntypes, since in many cases the specimen marked “type” in the collection does not coincide with the one figured in the original description, or more than one specimen is marked “type,” or the type lot indicated by Ihering in the collection is not from the published type locality. There are several names where no specimen was figured and no specimen is marked as being a type. Finally, in some cases Ihering marked as “type” a better specimen received after the date of description. In Ihering (1907a), species names are separated by commas from a worker’s name and associated literature citation. It is therefore not possible to determine if that person and citation are the original author and description of the preceding name, or only a subsequent citation referring to that name. It was necessary to consult those citations directly in order to determine this difference. The stratigraphical notation of Ihering is given in parentheses on the same line of the reference (usually the last where the author corrected his own earlier as- signments), and in the author’s original spelling (French, Spanish, or Portuguese). The relative geological age according to current knowledge is given on the line Stratigraphy. The catalogue numbers correspond with few exceptions to those given to the collection in the Museo Argentino de Ciencias Naturales. There are a number of species, subspecies, or “varieties” that Ihering (1914) listed or described which were not found in the collection; in those cases, their absence is clearly stated. The date of publication for names proposed by A. D. d’Orbigny follow Parodiz (1957). Localities Alsina, Valle. Rio Negro Province, with strata of the Roca and Salamanca For- mations (Paleocene), indicated by Ihering for species he described as “upper cretaceous” or “etage guaranieen.” See also Puente Alsina. de San Julian. Narrow harbor at San Julian, Santa Cruz. Bajada de Parana. Cliff below the city of Parana, Entre Rios Province. Type section for some species from the Parana Eormation. 1996 Parodiz— -Ihering Mollusca 187 Bajo de la Pava, Lowland in the vicinity of Puerto Deseado, with “Sierra Laziar strata” (Laziarense), Araucanean Formation, Pliocene. Bajo de las Flechas. Lowland near the lower section of the Rio Deseado, southeast of Cerro Moro, near the coast of Santa Cruz. Bajo de San Julian. See San Julian. Boqueron. Punta Boqueron. On Bahia Inutil, Magellan Strait, Tierra del Fuego. Buentiempo. Cabo Buen Tiempo. Cape Fairweather, southern Santa Cruz; locality first explored by Hatcher (1897). Perforations made by Yacimientos Petroli- feros Fiscales (Argentina Oil Bureau) 15 km W of this locality, found con- tinental deposits at 93 m (Santa Cruz Formation), and Tertiary marine de- posits at 665 m. See Feruglio, 1949-1950:279. Camarones, Bahia. Bay immediately north of the Golfo San Jorge. Canada de los Artilleros. Long gorge running east from Cerro Cuadrado to the Rio Chico in Santa Cruz. Yegua Quemada and Las Cuevas are localities along it. Cahadon Blanco. South side of Colhue Huapi Lake, Chubut Province. Carmen Sylva. Tierra del Fuego. Species of this locality are mostly of the Ma- gellanian, and some of the Patagonian formations. Casamayor, Punta Casamayor. Point on southern part of Golfo San Jorge in Santa Cruz, north of Mazaredo. In geopaleontological bibliography sometimes spelled, Casa-Maior or Casamajor. Cerro Bagual or Cerro de los Baguales. North of the Rio Sehuen Valley west of Mata Amarilla, Santa Cruz. Thick deposits of the Salamanca Formation. Do not confuse with Sierra Baguales which is in south-central Santa Cruz. Chapadmalal. Creek and town south of Mar del Plata, Buenos Aires Province. Chico River, Rio Chico of Chubut Province. Tributary of Chubut River running northeast from Colhue Huapi Lake, eastern Chubut Province. Chico River, Rw Chico of Santa Cruz. Tributary of Rio Sehuen running southeast. Chocoi, Punta Chocoi. On Magellan Strait at Carelmapu, Chile. Colhue Huapi. Lake and locality in southeast Chubut Province, east of Lake Muster. Colloncura, Rio. Tributary of the Limay River in southern Neuquen Province. Corral Foyel. Locality on the Andean zone of Nahuel Huapi, along the Foyel River and Foyel Lake. First explored by the geologist Santiago Roth in 1897- 98, and later by R. Hauthal (1898). Deseado, Puerto Deseado, Port Desire. South of Golfo San Jorge, at the mouth (or ria) of Deseado River in Santa Cruz. Deseado Island. An early Tertiary island in the San Jorge Sea, practically sepa- rating northern and southern parts of that sea during the Paleocene. Fairweather, Cape. See Buentiempo. Foyel, Rio, Lago. See Corral Foyel. Foz de Santa Cruz. In Ihering (1914); mouth (or ria) of Santa Cruz River. General Roca. Roca in the geopaleontological literature. Town in the upper Rio Negro Valley, east of the junction with Rio Limay, which was earlier known as “Fresno Menuco,” Rio Negro. Homo Viejo [sic]. Error for Homo Viejo, near Roca, Rio Negro. Jahuel or Jaguel. Below the east slope of Sierra Auca Mahuida and Bajo del Palo, south of Rio Colorado, Neuquen Province; type section for many spe- cies from Paleocene fresh- water deposits. Jack Harvey. South of Monte Observacion and south of Coyle, Santa Cruz. Jegua Quemada. See Yegua Quemada. 188 Annals of Carnegie Museum VOL. 65 La Cueva or Las Cuevas. Six mi S of the mouth of Rio Santa Cruz and north of Rio Coyle, near Yegua Quemada, and east of Monte Leon. Laziar, Sierra Laziar. 50 km NW of Puerto Deseado, Santa Cruz; type section for species from deposits of the Araucanean Formation, Pliocene. Lele or Leleque. Creek in northwest Chubut Province, at the base of Sierra Le~ leque, halfway between norquinco and Esquel. Lujan. Old city on the Lujan River, 100 mi N of Buenos Aires. Pleistocene de- posits. Maiten, Maiten Valle. Rio Negro Province. Malaspina. 25 km W of Bahia Bustamante, northern part of Golfo San Jorge, Chubut Province. Mata Amarilla. On the north slope of Cerro del Castillo and 20 km E of Park- Aik, Santa Cruz. Monte Cazador (Capas with LahilUa luisa). A number of marine species were described by Wilckens from this locality that were confused with the strata of Salamanca (Paleocene); these are Upper Cretaceous. Monte Espejo. Marine terrace located near San Julian, Santa Cruz, north of Rio Seco and 50 km S of Bahia Laura. Monte Leon. South of the mouth of Rio Santa Cruz, 5 mi inland from Punta Leon, Santa Cruz. Monte Observacion. Hill between mouth of Rio Santa Cruz and Puerto Coig. Owen Point. Collecting locality on coast of Santa Cruz, named by Hatcher. Pan de Azucar. Promontories north of the lower Rio Chico Valley, near Gaiman, with Paleocene to Miocene strata, Chubut. Park~Aik or Pari-Aike. Name of an estancia (ranch) that was located 20 km W of Mata Amarilla, along the Rio Sehuen, Santa Cruz. Piedra Clavada. A “torreon” or tower produced by eolic erosion of sandstones from the Sehuenan beds, in the Rio Sehuen Valley; west of Kohuel Kaike on the railroad between Puerto Deseado and Colonia Las Heras, Santa Cruz. Puente Alsina. South of Buenos Aires city. Perforation found Miocene strata of the Parana Formation at a depth of 80 m. Puerto Militar: now Puerto Belgrano. Naval base in southwest Buenos Aires Province at Bahia Blanca. The vicinity is the type section for species of the marine Belgrano Formation, early Pleistocene. Pueyrreddn, Lago. Lake in western Santa Cruz, on the border between Argentina and Chile. Punta Atlas. Point midway between Rawson and Camarones, Chubut. Punta Nodales. On the Golfo San Jorge, 20 km N of Comodoro Rivadavia, Chu- but Province. Punta Nueva, Nava, Nova. Point between Bahia Mazaredo and Bahia Bustamante, Golfo San Jorge, Santa Cruz. Rada Tilly. Harbor near Comodoro Rivadavia, Golfo San Jorge. Salamanca, Pico. Peak on the Golfo San Jorge, near Bahia Solano, Santa Cruz, between Puerto Viser and Punta Matalinares. Marine Paleocene strata. San Julian, Bajo de San Julian. Southern Santa Cruz lowlands. Typical strata of the Lower Patagonian Formation (Juliense). Santa Cruz. Santa Cruz Province. Port at the mouth of the Santa Cruz River. Ihering’s references to Santa Cruz for fossils of different strata are not always clear; it may be the locality of that name or some other place in Santa Cruz Province (a Territory at the time). 1996 Parodiz — Ihering Mollusca 189 Santa Maria da Bocca do Monte, Rio Grande do Sul, Brazil. Sehuen Aik. Near Mata Amarilla, on the Valley of Rfo Sehuen, Santa Cruz. Some- times referred to by Ihering as Sehuen-Ark. Sehuen River (also called Chalia). Runs from the vicinity of Lago Viedma in Santa Cruz to become a tributary of the Rfo Chico. The name Chalia was applied for its vicinity to an estancia (ranch) of that name near Mata Amar- illa. Shell Gap. A locality mentioned by Ortmann (1902) and named by Hatcher, on the upper section of Rio Chico in Santa Cruz, near El Portezuelo. Tres Puntas. Cape south of Golfo San Jorge, near Deseado, Santa Cruz. Valcheta, Travesia. Area along the Gualicho River in western Rfo Negro. Villarino, Punta Villarino. On Peninsula Valdez, Chubut. Yegua Quemada. North of, and very close to. La Cueva, a few miles south of the mouth of the Santa Cruz River, east of Monte Leon. Stratigraphic Units Upper Cretaceous Senonian—Chubutian Strata with Lahillia luisa (Pehuenche-Pehuenchense),—The complex of Upper Cretaceous deposits, including the Lahillia luisa beds, were known to early au- thors as Chubutiano or Chubutense. According to E. Feruglio (1949=1950) that complex of sediments in southern Patagonia reaches a thickness of 2300 m. Ge- ologists recognize that a considerable hiatus exists between the Senonian and the marine deposits of the Paleocene. The fauna of invertebrates at the end of the Cretaceous showed more global geographic relationships with a complete change in the deposits of the Eocene, with no species affinities. In Ihering 's time, the concept of the Paleocene was not yet developed, and the Cretaceous-Tertiary boundary was not established. Marine strata now known as Paleocene were in- cluded in the Cretaceous. Even early in this century, authors such as Wichmann (1927) and Windhausen (1918; 1931, in part), parallelled Roca and Salamanca formations with the strata containing Lahillia luisa. Until near the end of the Cretaceous the conditions in the Patagonian region were of continental character. Then epirogenical movements began and continued into the early Tertiary, especially in the area now corresponding to the Golfo San Jorge, which was transformed into a great basin (Feruglio, 1949-1950:17). Tertiary Paleocene Guaranitic.—lhmng, Ameghino, and other early writers on Patagonian stratig- raphy used the name “formacion guaranitica” for a complex of strata correspond- ing globally to the Paleocene-Eocene, derived from d’Orbigny’s (1842) “Tertiarie Guaranien,” “Formacion Guaranitica” of Coring (1881-1882), and “Guaranitic Beds” of Hatcher (1897). Other authors included it in the “Pehuenche” (Upper Cretaceous). These sediments contain Ostrea guaranitica Ihering (= Exogyra guaranitica [Ihering]), Corbula sehuena Ihering (= Eriphyla sehuena [Ihering]), and Potamides patagonensis Ihering; it is recognized as a brackish-water unit of the Paleocene of the Sehuen Valley of Santa Cruz. See Sehuen Formation. Roca Formation (Rocanense). — -The marine deposits of Ihering’s (1903) Ro- caneen, or Rocanense of other authors, were named after the type locality, the 190 Annals of Carnegie Museum VOL. 65 village of General Roca on the left bank of the Rio Negro, Rio Negro Province. The Roca beds represent the earlier Tertiary ingression of the San Jorge Sea in its northern section. Ihering maintained that the marine Roca was of Upper Cre- taceous age. Windhausen (1916) was the first to report that the “Roca horizon corresponds probably to the lowest part of the Tertiary.” While the malacofauna of Roca (and Salamanca) appears as a precursor of the subsequent Tertiary marine faunas, and differs from that of the Upper Cretaceous, it contains some charac- teristic old elements, such as Odontogryphaea rostrigera (Ihering), Odontogry- phaea rocana (Ihering) (= Odontogryphaea rothi [Bohm, 1903]), and Ampullina {Pseudotylo stoma) romeroi (Ihering). Rw Chico Formation. — This continental formation, named and defined by G. G. Simpson (1933), contains the oldest Patagonian mammal fauna. Land molluscs are very rare (Thaumastus patagonicus Parodiz), but the consideration of the strata is important for correlations with marine and fresh-water strata of the Paleocene. It overlies, and is in part coeval with, the marine Salamanca, but the beginning of the Rio Chico sedimentation was most probably Danian; its recognized base is laterally equivalent to the Sehuen beds (Montian) and developed considerably during the Thanetian. Jahuel Formation. — Windhausen (1918) designated this formation as “Capas de Jahuel” strata of that locality, near the eastern slope of Sierra Mahuida, south of the Colorado River in Neuquen. The greater development is in Rio Negro, near Roca (Windhausen also called it “lower Roca”). It corresponds to the “Sen- onian lacustre” of Wichmann (1927) and in part to the Jahuelense of Feruglio (1949-1950). By mechanics of deposition it fills the eroded depressions of Roca, and sometimes is parallel to Rfo Chico. The deposits are fresh- water, and correl- ative to the southern brackish-water Sehuenan, The fresh-water fossils of Jahuel are abundant. The species belong to genera that appear for the first time in Patagonia, some still living such as Diplodon and Cyanocyclas (= Neocorbicula), or extinct such as Paleoanculosa. They indicate a migration from the Northern Hemisphere that was correlated with that of the early mammals of Patagonia. Salamanca Formation. — This is the southern facies of the San Jorge Sea, as Roca was of the north. It is frequently referred to in the literature as Salam- anque[i]ano or Salamanqueen. Although both Salamanca and Roca began in the Danian, Salamanca lasted longer into the Montian with greater development. The type section is at Salamanca Peak near the coast of the Golfo San Jorge, Santa Cruz. Feruglio (1949-1950) found the strata underlying the Rio Chico Formation, with brackish-water intercalations of the Sehuenan in situations that represent deltaic environments, and with thin layers containing fossil plants indicating tran- sition to the continental conditions that prevailed during the Eocene. Although there is a notable hiatus between the fauna of the Paleocene and the beginning of the Patagonian Formation in the Lower Miocene, there are a number of related species. Stenzel (1945) still considered Roca and Salamanca as a single mid-upper Paleocene unit. Lithologically the facies are distinguishable: Salamanca sediments are clay and sandstone, and those of the Roca are mostly limestone. Feruglio (1949-1950:307) identified 67 forms of marine invertebrates which are exclusive to the Salamanca. Ihering described 14. species that occur in the Salamanca Formation, among which Gryphaea pyrotheriorum and Odontogry- phaea rostrigera also occur in the Roca Formation. 1996 Parodiz — Ihering Mollusca 191 Sehuen Formation (Piso Sehuense). — The 'Tiso Sehuense” or Sehueneen was first reported by Ameghino as coeval with Roca, later placed above it, and finally (Ameghino, 1906) below Roca-Salamanca. The sequence given by Ihering (1907i3) was (still considering it Cretaceous): “Pyrotherienne” (now Deseado For- mation, base of the Oligocene); disconformity; “Sehueneen” (now Paleocene, Monti an-Thaneti an) . The sediments were localized along the Sehuen, also called Chalia River, Santa Cruz. It represents the lateral brackish water of the terrestrial Rio Chico Forma- tion, equivalent of what Jahuel is to Roca in the north. It corresponds to the phase of regression of the San Jorge Sea, and the brackish-water Potamides and Eriphyla are found mixed with Ostrea and Struthiolaria. Boca-Levu. —These early marine deposits of the Tertiary of Chile are placed between Paleocene and Eocene (Feruglio, 1949-1950:230, 231). Eocene Casamayor Eormation. — This is the “capas with Notostylops'' or Notostylo- pense in the nomenclature of Ihering and Ameghino. Defined by Simpson (1933), it overlies the Rio Chico Formation and at its beginning was lateral to the re- gression of the San Jorge Sea. Its age is Lower Eocene, and together with the Muster Formation above, was called a unit by most geologists, the “Sarmiento group,” or “tuffs with mammals.” Molluscs are few but characteristic: Stro~ phocheilus chubutensis Ihering, Strophocheilus hauthali Ihering (= Megalobuli- mus hauthali [Ihering]), Paleobulimulus eocenicus Parodiz, and Chilina stenos- tylops Parodiz, all found in the upper section of the complex (Parodiz, 1949fl). Oligocene Magellanian Formation. — -This consists of marine deposits in the southern part of Santa Cruz Province, designated by Hatcher (1897, 1900) as the Magellanian beds, Lower Oligocene. Ihering placed it between the Santa Cruz and the Pata- gonian formations, but actually it is underlain by both. It represents the beginning of the great marine ingression that continued in the north in the Lower Miocene (the Patagonian Formation). According to Feruglio (1949-1950), who conducted extensive research, the age of the Magellanian must be between the middle and the end of the Oligocene. The separation of the Magellanian and Patagonian is geographical rather than stratigraphical, even if the Patagonian is younger as a whole. In Tierra del Fuego, the Strata of Boqueron at Bahia Inutil are, at their base, equivalent with the Magellanian, but also part of them belong to the Pata- gonian Formation. A number of forms described by Ihering for the Magellanian {Glycymeris cuevensis, Turritella ambulacrum sylva, Venericardia sylva, and Comine lla fueguensis) were found also in the Patagonian Formation and the Na- vidad beds of Chile. The greater age assigned by Flatcher for the Magellanian (Lower Oligocene) is probably correct. Queguay Formation. — This is the “Calizas de Queguay” (Queguay Lime- stone), the oldest Tertiary deposits known from Uruguay, lying in disconformity over the Asencio sandstones (Upper Cretaceous) and below the Fray Bentos clay (Miocene). Chronologically they are coeval with the Magellanian, and correspond to the lacustrine-eolic system with Strophocheilus, Paleoanculosa, and ''Plan- orbis'" waltheri. These strata are found mainly on the Queguay River Valley, north of Paysandu. 192 Annals of Carnegie Museum VOL. 65 Miocene Santa Cruz Formation. — This vast terrestrial sedimentation began to develop north and lateral to the Upper Magellanian, and continued coeval to the Patago- nian Formation, finally overlying it. Ihering {1^91 a) referred the marine fauna he described to the Santacruzense, and in 1907 indicated its relative position to be above, but mostly coeval, with the Patagonian that he considered to be at the Eocene-Oligocene boundary (Ihering, 1901 a). Feruglio (1949-1950) found it fill- ing also the eroded surfaces of the Patagonian, beginning with the Lower Miocene and progressing upwards thereafter. Its extension into the middle Miocene forms a hiatus between the marine faunas of the Patagonian and the Rio Negro forma- tions. In contrast with the molluscan contents of the Paleocene and Eocene continental formations, no land or fresh-water shells are known from the Santa Cruz For- mation. The absence is impressive compared with the abundant fluvio-terrestrial malacofauna in Peruvian, Ecuadorian, and Colombian deposits of the same age (Pilsbry and Brown, 1917; Bristow and Parodiz, 1982). Lower Miocene Marine Deposits. — Three large marine ingressions, the Oli- gocene Magellanian, the transition from Upper Oligocene to the early Miocene Patagonian, and the Navidad beds of Chile, covered a vast area. Between the northeastern part of the Patagonian Sea on the coast of Chubut, and the Navidad beds in the northwest, there was a triangular, uncovered peninsula of large di- mensions. To the south, the sea covered all the territory down to Tierra del Fuego, intercalated with some areas of terrestrial sedimentation (Santa Cruz Formation). Patagonian Formation. — D’Orbigny (1842) gave the name of “Tertiaire Pa- tagonien” to all the deposits of marine sedimentation from the north at Parana, to the south in Patagonia. Above it he placed the “Argile Pampienne” (Pliocene- Pleistocene), and below the “Tertiaire Guaranien.” The classification of the sed- imentary complex used by Ihering (1897Z?) when he described a large number of fossil Mollusca was the same dichotomy that Ameghino had made the same year: Patagonian and Superpatagonian formations. He also correlated the Patagonian with the terrestrial Santa Cruz Formation, placing both in the Eocene. Thus, most of the species were indicated as being from the “Form. Santacruzense” and, as it is explained in the appendix (Ihering, I897Z?:317-3I9), distinguished from the Patagonian Formation which he considered to be relatively older. Upper Eocene. In his larger work of 1907(<3) he named all these deposits in a global sense as “Pan-Patagonian” in which he established some chronological levels: above was the Superpatagonian made coeval with the Santa Cruz Formation, and better rep- resented at localities such as La Cueva and Yegua Quemada; the middle was occupied by the Monte Leon strata (Leonense) and the San Julian (Juliense); at the base were the strata of Bahia Camarones (Camaronense) or “Patagoneen in- fmeur.” Such divisions were based only on the relative relationship between the paleontological contents and living species. Many of the taxa named by Ihering were subspecific or infrasubspecific. The clarification of the taxonomy, as well as the recognition of many synonyms, makes his stratigraphic classification untenable, apart from the fact that the Eocene in Patagonia was predominantly a continental sedimentation. Ortmann (1902), study- ing the fossils from Hatcher’s expedition to Patagonia, concluded that the Pata- gonian Eormation was indivisable and entirely Miocene. Finally, the paleontolo- gist George G. Simpson (1932, 1933, 1935, 1940, 1941), and the geologist Egidio 1996 Parodiz — Ihering Mollusca 193 Femglio (1933, 1935, 1949, 1950) maintained that the Patagonian Formation can- not be older than Lower Miocene. But very probably the sedimentation, at its base, began as early as the Upper Oligocene, and as such these strata were in- dicated by modern authors (Del Rio and Morra, 1984; Erdmann and Morra, 1985; Morra, 1985) as the San Julian Formation (Ihering’s Juliense), and sometimes even older in age. Since the older marine Oligocene is represented by the Ma- gellanian Formation, the base of the Patagonian cannot be older than Upper Oli- gocene. However, the paleontological contents of the Patagonian {sensu lato) in their majority, even those included in the lower level, show features that are characteristically Lower Miocene. In the stratigraphic indications of our list of species, the age is given as such for the Patagonian Formation, although in many cases they belong to the Juliense (such as those materials from the Canada de los Artilleros). Actually the fauna of the earlier Patagonian Formation is transitional between Upper Oligocene and Lower Miocene. The Navidad beds of Chile are synchronic with the Patagonian Formation. The Patagonian Formation contains the richest of the Tertiary marine malaco- faunas of South America. Feruglio (1949-1950) estimated 325 recognizable spe- cies: 153 bivalves, 168 gastropods, four scaphopods, and one cephalopod. Ihering described many species from the Patagonian Formation that belong to genera that today represent elements of warmer climate in the northern zone. Among these are Abra, Lucina, Sanguinolaria, Aporrhais, Tibia, Laevicardium, Macoma, and Glycymeris. The absence of these and many other genera in the malacofauna of Recent Patagonian and Magellanian regions reflects displacement by plate tecton- ics rather than “local” changes in temperature. Collon—Cura Beds (Colloncurense). — These are terrestrial and in part fresh- water sediments in the western part of the Chubut Province. Kraglievich (1930) named it “Formacion Friaseana” (from Lake Frias) and considered the strata as Oligocene. Simpson (1940) and Feruglio (1949-1950) placed the Collon-Cura in the Middle or Upper Miocene, above the Santa Cruz and Patagonian formations. Molluscs are rare: Diplodon rothi Ihering. Fray Bentos Formation (Fray Bentos Clay). — This consists of strata in Uru- guay, on top of the Queguay limestone (Oligocene) and below (hiatus) the Ca- macho Formation (Pliocene). Parana Formation (Paranense of Authors; Also Known as Mesopotamian). — This consists of rich marine sediments in northeastern Argentina. The type section is located in the vicinity of Parana city, Entre Rios Province, and also found in deep perforations in Buenos Aires and Uruguay. It contains many molluscan gen- era that denote tropical or semitropical conditions. Authors differed about its age which was indicated as old as Oligocene and as young as Pliocene, but the actual position is in the Upper Miocene. It contains a larger number of extinct species than the next formation, the Araucanean, which is Pliocene. In Uruguay, the Parana Formation is particularly represented in the Department of Colonia, where it is known as the Camacho Formation (sandstone), correlated by Caorsi and Goni (1958:59) to the “Transgresion marina entreriana” but given the age of Pliocene. Also, fossils found at Punta Gorda on the Uruguay River are the same as those from perforations at Chuy on the Atlantic coast which are Miocene in age. Ihering used the name “Formacion Entreriana” as did several other authors. However, the name “Entreriense” or “Entreriano of Patagonia” was also applied to its chronological equivalent in Rio Negro, the “Rionegrense” or Rio Negro 194 Annals of Carnegie Museum VOL. 65 Formation. More difficult to define is the Araucano-Entreriano complex under which Kraglievich (1930), Simpson (1940), and others united the ‘Taranense,” the '‘Rionegrense,” and the Araucanean formations. Rio Negro Formation (Rionegrense). — The “Rionegrense” was equal to what some authors called the ''Entreriense” of northern Patagonia. The type section is Puerto Piramides in Golfo Nuevo, Chubut. There are similarities between the faunas of the Rio Negro and Parana formations. It is relatively younger than Parana, and corresponds to a later, southern ingressive phase of the Parana Sea, which marks the boundary between the Miocene and Pliocene, although distin- guishable from the Pliocene Araucanean. It has in common with the Parana For- mation species of Dosinia and Lucina, but the fauna is generally poorer with some characteristic species such as Amusium cossmanni Ihering. Of the 52 species known from the Rio Negro beds, 23 are exclusive and 26 were known in the Lower Miocene, showing faunistic continuity with chronological changes along the mid-Tertiary Patagonian strand. Pliocene Fairweather Beds (Fairweatheriense or Buentiempense).— This stratigraphic unit was reported by Hatcher (1897) for the Miocene of southern Santa Craz, According to Feruglio (1949-1950) it belongs to the Upper Pliocene. It appears coeval with northern deposits at Chapadmalal as an upper section of the Arau- canean complex. Araucanean Formation. — -The “Araucano” is a complex of sediments first ob- served by Charles Darwin in December 1833 and January 1834. Doring (1881- 1882) named it “Tehuelche” of which the larger part belonged to the Pleistocene. It consists of continental sediments and marine terraces of extensive development in the provinces of Chubut and Santa Cruz, particularly at Sierra Laziar (see Feruglio, 1933). Sierra Laziar Strata (Laziarense). — Most of the fossil Mollusca named by Iher- ing from the “Araucano” are from Sierra Laziar. Although it contains a good number of extinct species, it shows the transition from the ancient to the Recent fauna; some species or subspecies named by Ihering from this strata are synonyms of Recent ones: Pupilia aperta tehuelcha Ihering = Fissurellidea megatrema d’Orbigny; Neomphalius princeps Ihering = Tegula patagonica (d’Orbigny); Tro- phon necocheanus Ihering = Trophon lamellosus (Gmelin); Trophon inornatus gradatus Ihering = Trophon varians (d’Orbigny); Trophon varians montenus Ihering = Trophon varians (d’Orbigny); Chione vindex Ihering = Protothaca antiqua (King); Marcia scutata tehuelcha Ihering = Samarangia exalbida (Chem- nitz). Quaternary Pleistocene Belgrano Formation (Belgranense).— This is the first marine ingression of the Pleistocene. It is also known as the “Pampeano marine” of other authors. The sedimentation in the north was coeval with the terrestrial Buenos Aires Formation; the intercalations reveal the oscillation of the Atlantic strand. Geographically it is divided into two sections: Belgrano North, with greater development in Buenos Aires Province with almost all of the species still living; Belgrano South, in southern Santa Cruz, which contains the same species found in the Belgrano 1996 Parodiz — Ihering Mollusca 195 North. The southern section was also called “Deseadense,” but that term was used by Kraglievich (1930) and other authors for a different stratigraphic unit of terrestrial deposits. Buenos Aires Formation (Bonaerense). — This formation consists of very ex- tensive eolic sedimentation, synchronic and alternating with the Belgrano For- mation on the coast. It is very poor in land shells: Peronaeus {Lissoacme) amegh- inoi (Ihering), Strophocheilus charruanus (Frenguelli). In Uruguay, the same co- eval deposits are known as “Loess of Arazati.” Querandi Formation (Querandino). — This is the latest of the marine ingres- sions at the end of the Pleistocene. It was of very short duration but its deposits are found farther inland than Belgrano, up the Parana River, and of great devel- opment in Uruguay where the Belgrano beds are absent. The Querandi Formation coincided with the tectonic movement that produced the great fault of the Parana, and subsequent marine regression formed the La Plata estuary. It contains species that now live north of that estuary, such as Diplodon charruanus lujanensis Iher- ing (= Diplodon delodontus [Lamarck]) in some fresh- water intercalations. In Uruguay it is also known, in part, as the “Clays of Vizcaino.” Tertiary Climate and Paleogeographic Considerations Ihering’s important contribution to the geopaleontological history of South America was the introduction of the concept of different land masses that he named Archhelenis, Archiplata, and Archinotis (Ihering, 1907^). These ideas were elaborated two decades earlier than Wegener’s 1916 presentation of his continental drift theory (Wegener, 1924). Of Archhelenis, which in Ihering’s paleogeography united Africa with South America, he said (Ihering, 1907<3:483), “Le continent qui unissait le Bresil a I’Afrique, commencait a disparaitre pendant la formation cretacee.” That indicates that his idea of Archhelenis was not just of a bridge but of a continent. That same year (Ihering, \901b) he epitomized and developed his ideas in the book “Ar- chhelenis und Archinotis”; in it he illustrated Brasilia and Archiplata united to Africa by Archhelenis, even projecting it into the Indian Ocean in a mass of land of tremendous proportions. Actually Brasilia was the same as Archhelenis, or part of it, and its tectonic plate had been separated from Africa long before the Cretaceous. It was a large mass of land between the northern sea, Tethys, and the southern. Nereis. Ancient faunistic elements of Gondwanian origin developed into new groups by isolation in Archhelenis. Families which later became the Mycetopodidae and the Stro- phocheilidae in South America are good examples. West of Archhelenis was Archiplata, separated by a long, narrow sea. Up to the early Tertiary Archiplata was connected to Archinotis (Antarctica), and the land, with Patagonia at the center, extended as far north as Peru. During the Paleocene there suddenly appeared a large fauna of mammals, fresh-water mol- luscs, and elements of other zoological groups that were closely related to those of the Cretaceous of North America. The migration must have taken place through some connection between northern and southern regions of the Western Hemi- sphere. The connection was interrupted during the entire Tertiary and the southern faunas developed in isolation. There is also evidence of identical, or very closely related, species discovered in mid-Tertiary sediments of Antarctica and Patagonia. These are littoral molluscs that indicate a common continental shelf existing be- 196 Annals of Carnegie Museum VOL. 65 tween the two regions and a similarity in climatic conditions. Because of the subtropical, sometimes tropical, character of the Tertiary marine malacofauna, Ihering assumed that important climate changes had taken place and that the composition of the faunas was affected sometimes by barriers, sometimes by migrations. A. E. Ortmann supported the Archhelenis-Archiplata proposition, although he strongly disagreed with Ihering about the age of the Patagonian deposits. After Ortmann’s (1902) and Ihering’s (1907a, 19071?) paleontological work, when ge- ologists began to combine their ideas with new concepts of plate tectonics, still they interpreted, as Wegener did in his continental drift theory, the displacement of the South American continent westward as if it had been a solid and unified mass, instead of being formed of different plates originally separated, that in- cluded also Archiguiana. The mechanics of plate tectonics displaced Archiplata in a clockwise direction toward a west-northwest position relative to that which the continent occupies today. It remained connected with Antarctica which was also in lower latitudes. In such position, the east coast of Archiplata was facing more to the south than at present. The gyration must have advanced the tip of the continent to latitude 30-35°S. After the middle Tertiary, the movement of the “pendulum” turned counterclockwise until reaching the Recent position with its tip at latitude 56°S. These oscillating movements explain the climate changes that affected the marine as well as the fresh-water faunas, and their distribution. The author who first proposed the rotation theory was H. Modell (1950). After many years of studying the taxonomy and geographic distribution of the super- families Unionacea and Mutelacea in relation to climate, Modell arrived at a logical conclusion. Modell’s schematic illustration (1950:143) seems somewhat exaggerated, but it is supported by many biogeographic facts related to the changes in climate that occurred between the Tertiary and Recent in South Amer- ica. The unification of Archhelenis- Archiplata began as early as the first phase of the Andean orogeny on the Andean syncline, but a communication between the Pacific and the new Atlantic Ocean still existed in the Miocene, and probably until the Pliocene. The same gyration that modified the position of the South American continent separated Antarctica, its plate drifting south. Abbreviations ACM AJS AMNBA ARMGB ASCA BGSA BSGF BIGU CMNH CZMHN Annals of Carnegie Museum, Pittsburgh. American Journal of Science. Anales del Museo Nacional de Buenos Aires. Anali del Reale Museu Geologico di Bologna. Anales de la Sociedad Cientifica Argentina. Bulletin of the Geological Society of America. Bulletin de la Societe Geologique de France. Boletm del Instituto Geologico del Uruguay, Montevideo. Carnegie Museum of Natural History, Pittsburgh. Comunicaciones Zoologicas del Museo de Historia Natural de Mon- tevideo. International Commission on Zoological Nomenclature. Museo Argentine de Ciencias Maturates, Buenos Aires. ICZN MACN 1996 Parodiz — Ihering Mollusca 197 NDMG Nachrichtsblatter Deutsche Malakozoologische Gesellschaft, Frank- furt. NJMGP Neues Jahrbuch fur Mineralogie, Geologic, und Palaoetologie, Stuttgart. NPRMP Notas Preliminares da Revista do Museu Paulista, Sao Paulo. PAPS Proceedings of the American Philosophical Society. RMLP Revista del Museo de La Plata, Argentina, RMP Revista do Museu Paulista, Sao Paulo. List of New Combinations abavus Ihering, 1907. Neomphalius = Tegula abava (Ihering), new combination ameghinoi Ihering, 1907. Cucullaria. = Barbatia ameghinoi (Ihering), new combination ameghinoi Ihering, 1896. Voluta = Adelomelon ameghinoi (Ihering), new combination americanus Ihering, 1907. Neomphalius = Tegula americana (Ihering), new combination burmeisteri Ihering, 1907. Cymbiola = Adelomelon burmeisteri (Ihering), new combination camaronesia Ihering, 1907. Area = Anadara camaronesia (Ihering), new combination cannada Ihering, 1907. Cymbiola becki “var.” = Adelomelon cannada (Ihering), new combination, new status chicoana Ihering, 1907. Hadriania jorgensis “var.” MURICIDAE = Trophon chicoanus (Ihering), new combination, new status chubutiana Ihering, 1907. Scalaria EPITONIIDAE = Epitonium chubutianum (Ihering), new combination contorta Borchert, 1901. Modiola HYRIIDAE = Diplodon contortus (Borchert), new combination dalli Ihering, 1907. Lyria VOLUTIDAE = Enaeta dalli (Ihering), new combination dautzembergi Ihering, 1897. Triton (Argobuccinum) RANELLIDAE: RANELLINAE = Fusitriton dautzembergi (Ihering), new combination elegans Ortmaee, 1900. Urosalpinx MURICIDAE: OCENEBRINAE = Ocenebra elegans (Ortmann), new combination jorgensis Ihering, 1907. Hadriania MURICIDAE: OCENEBRINAE = Crassilabrum Jorgensis (Ihering), new combination Juliana Ihering, 1907. Area ARCIDAE = Anadara Juliana (Ihering), new combination major Ihering, 1899. Cancellaria gracilis “var.” CANCELL ARIIDAE = Naronia major (Ihering), new combination, new status monoceros Ihering, 1907. Trophon MURICIDAE = Thais monoceros (Ihering), new combination TROCHIDAE ARCIDAE VOLUTIDAE TROCHIDAE VOLUTIDAE ARCIDAE VOLUTIDAE 198 Annals of Carnegie Museum VOL. 65 multicoronatus Ihering, 1907. Vermetus = Dendropoma multicoronatus (Ihering), new combination patagonica Ihering, 1897. Bulla = Retusa patagonica (Ihering), new combination paucispina Ihering, 1897. Pholas = Netastoma paucispina (Ihering), new combination pyrotheriana Ihering, 1907. Polynices = Lunatia pyrotheriana (Ihering), new combination quemadensis Cossmann, 1899. Cirsotrema = Opalia quemadensis (Cossmann), new combination VERMETIDAE 7RETUSIDAE PHOLADIDAE NATICIDAE EPITONIIDAE rada Ihering, 1907. Lotorium (Lampusia) RANELLIDAE: CYMATIINAE = Semitriton rada (Ihering), new combination rocana Ihering, 1907. Phacoides LUCINIDAE = Lucina rocana (Ihering), new combination rocanum Ihering, 1907. Cardium (Hemicardium) = Trachycardium rocanum (Ihering), new combination CARDIIDAE salobris Ihering, 1907. Cyrena CORBICULIDAE = Cyanocyclas salobris (Ihering), new combination santacruzense Ihering, 1914. Cardium CARDIIDAE = Laevicardium santacruzense (Ihering), new combination santacruzensis Ihering, 1907. Chenopus APORRHAIDAE = Drepanocheilus santacruzensis (Ihering), new combination schucherti Ihering, 1914. Gibbula ARCHITECTONICIDAE = Architectonica schucherti (Ihering), new combination stromeri Ihering, 1914. Fusus RANELLIDAE: RANELLINAE = Fusitriton stromeri (Ihering), new combination tridentata Ihering, 1897. Cuculearia [sic] ARCIDAE = Barbatia tridentata (Ihering), new combination usurpator Ihering, 1907. Lotorium {Lampusia) RANELLIDAE: CYMATIINAE = Cymatium usurpator (Ihering), new combination. List of Taxa abavuSf 1907. Neomphalius TROCHIDAE 1907a. AMNBA, 14:134, pi. 4, fig. 9a-b. 1914. NPRMP, 1(3):76 (Patagonico Inferior). = Tegula abava (Ihering), new combination. Type Locality. — Southeast of Punta Nova, Golfo San Jorge, Santa Cruz. Syntypes. — #591, five specimens, coll. Ameghino, 1900. Stratigraphy. — Lower section of the Patagonian Formation (Lower Miocene). Remarks. — The shell is higher than in the living T. patagonica (d’Orbigny) and therefore more similar to Calliostoma. aequistriata^ 1907. Gibbula cuevensis “van” ARCHITECTONICIDAE 1907a. AMNBA, 14:132. Not figured; diagnosis only. 1914. NPRMP, 1(3):73 (Patagonico Superior). 1996 Parodiz — Ihering Mollusca 199 = Valdesia dalli (Ihering). Type Locality.— -Cdibo Tres Puntas, Golfo San Jorge, Santa Cruz. Syntypes.- — #573, two specimens, coll. Ameghino, 1900. Other #574, southeast of Punta Nova, one specimen, coll. Ameghino, 1900; #575, Punta Casamayor, two specimens, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). ameghinoi, 1908. Bulimulus ORTHALICIDAE: BULIMULINAE 1908. AMNBA, 17:429 (not figured). 1944. Parodiz, CZMHN, l(17):l-3, pi. 1, fig. 1-6. = Peronaeus {Lissoacme) ameghinoi (Ihering). Type Loca/ify.—Chapadmalal, Buenos Aires Province. Type.— Not in the collection. Stratigraphy.— Buenos Aires Formation (Pleistocene). Remarks. — Bulimulus sporadicus pampa Ihering is also a synonym. ameghinoif 1897. Cancellaria CANCELLARIIDAE 1897^. RMP, 2:310, pi. 3, fig. 2, and pi. 4, fig. 15. 1907a. AMNBA, 14:215. Admete ameghinoi. 1914. NPRMP, 1(3): 112. = Admete ameghinoi (Ihering). Type Locality. — Yegua Quemada, Santa Cruz. Syntypes.— #900, four specimens, coll. Ameghino, 1893. Stratigraphy.— Fatagonia Formation (Lower Miocene). Remarks. — This species resembles the living Admete magellanica Strebel. ameghinoi, 1907. Cardium CARDIIDAE 1907a. AMNBA, 14:292, pi. 11, fig. 72a-b. 1914. NPRMP, 1(3):47 (Patagonico Inferior). Type Locality.— Santa Cruz. Holotype. — #391, one valve, coll. J. Bicego, 1897 (by monotypy). Other Lot. — #392, Punta Nova, one specimen, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). ameghinoi, 1907. Cucullaria ARCIDAE 1907a. AMNBA, 14:233, text fig. 8. 1914. NPRMP, 1(3): 16 (Patagonico Inferior). = Barbatia {Cucullaria) ameghinoi (Ihering), new combination. Type Locality.— Fan de Azucar at San Julian, Santa Cruz. Holotype. — #99, one valve, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). ameghinoi, 1907. Lutraria MESODESMATIDAE 1907a. AMNBA, 14:26, pi. 3, fig. 17a-b. Type Locality. — Roca, Rio Negro. Type. — Not in the collection. Stratigraphy. — Roca Formation (Paleocene). 200 Annals of Carnegie Museum VOL. 65 Remarks, —ThGic is an unnumbered specimen in the collection, but not listed in the 1914 catalogue. The illustration is unlike Lutraria and resembles a meso- desmatid. ameghinoi, 1897. Modiola MYTILIDAE 18971?. RMP, 2:233, pL 6, fig. 43. 1902. Ortmann, p. 21, pi. 25, fig. 2. Modiolus. 1907a. AMNBA, 14:275. Modiolus ameghinoL 1914, NPRMP, 1(3):38 (Patagonico Inferior and Superpatagonico). = Modiolus ameghinoi (Iherieg). Syntypes. — #316, two valves, coll. Ameghino, 1893. Other Lots,— 4^311 , southeast of Punta Nova; #318, Santa Cruz, coll. Amegh- ino, 1900. Stratigraphy. — -Patagonian Formation (Lower Miocene). Remarks.— Thi^ species was reported by Wilckens (1911) from the Antarctic at Seymour Island. This species resembles Mytilus pseudochorus Doello-Jurado, 1922. ameghinoi, 1902. Neoinoceramus MYTILIDAE: MYTILINAE 1902L PAPS, 41(6): 134, pi. 19, fig. 3-4. 1907a. AMNBA, 14:243, fig. 10. 1914, NPRMP, 1(3):20 (Patagonico Inferior). Type Locality. — -Southeast of Punta Nueva. Lectotype. — #136, an intact minor valve, corresponding to the figures in the original description, here designated. Stratigraphy. — -Patagonian Formation (Lower Miocene). Remarks. — Lot #136 in the Ihering Collection consists of two specimens from Golfo San Jorge, southeast of Punta Nueva. Only one specimen is listed in the catalogue (Ihering, 1914:20) and is a fragment of the paralectotype. It is a single complete large valve, 20 cm long, coll. Ameghino, 1900. Although listed in the catalogue it does not correspond to fig. 3--4 in the original description (Ihering, 19021?). The other specimen in Lot #136 is the lectotype designated above, and is not listed in the 1914 catalogue. ameghinoi, 1907. Myochlamys centralis “var.” PECTINIDAE 1897L RMP, 2:229, pi. 8, fig. 49. Pecten centralis. 1902. Ortmann, 2:114, pi, 21-22, fig. la-c {Pecten proximus); 2:116, pL 23, fig. la-b {Pecten cen- tralis). 1907a. AMNBA, 14:253 (Patagonico). Myochlamys centralis. = Chlamys centralis Sowerby, 1846. Type Locality.- — -Santa Cruz. Holotype.—dt24l, specimen received from. Ortmann as P. proximus (by mono- typy). Other Lot — #240, Yegua Quemada, six valves, coll. Ameghino, 1900. Stratigraphy.— Fatagonimi Formation (Lower Miocene). ameghinoi, 1902. Ostrea 1902a. AMNBA, 4:114, fig. 4-7. 1903. AMNBA, 9:213. OSTREIDAE 1996 Parodiz — Ihering Mollusca 201 1907a. AMNBA, 14:44 (Salamanqueen). 1914. NPRMP, 1(3):23 (Formacion Guaranitica with Gryphaea pyrothe riorum). Type Locality. — ^West of Rio Chico, Chubut, Syntypes.— 4^165, one complete specimen and 16 valves. Other Lot.— #166, Puerto Malaspina, three valves; no indication of collector. Stratigraphy.—SalsLmsinca. Formation (Paleocene). Remarks.Sce Ostrea rocana Ihering. ameghinoi, 1897. Struthiolaria STRUTHIOLARIIDAE 1897^. RMP, 2:289, text fig. 14. 1900. Ortmann, p. 380. Struthiolaria chilensis (not S. chilensis Philippi, 1887). 1904. Cossmann, p. 104, pi. 8, fig. 3-4. Struthiolaria omata (not S. omata Sowerby, 1846). 1907a. AMNBA, 14:172. 1914. NPRMP, 1(3):95 (Superpatagonico). Type Locality. — Originally ‘‘La Cueva and Santa Cruz.” Syntypes.— #763, Yegua Quemada, designated by Ihering (1914) as “Type lot,” 12 specimens, coll. Ameghino. Other Lots. — #764, Canada de los Artilleros, coll. Ameghino, 1900; #765, Santa Cruz; #766, Santa Cruz. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.— Two lots were received from Ortmann: #765 as S. chilensis Philippi, and #766 as S. densestriata Ihering. ameghinoi, 1903. Turritella TURRITELLIDAE 1903. AMNBA, 9:213, pi. 2, fig. 14. 1907a. AMNBA, 14:48. 1914. NPRMP, 1(3): 89 (Salamanqueano). Type Locality.— -Rio Chico, Chubut. Holotype. — #717, coll. Ameghino, 1900 (by monotypy). Stratigraphy.— Salamanca Formation (Paleocene). ameghinoi, 1896. Voluta VOLUTIDAE 1896. NDMG, 7:97. \^91b. RMP, 2:302, text fig. 17. 1899Z?. Cossmann, p. 125, fig. 18. Cymbiola. 1902. Ortmann, p. 233. 1907a. AMNBA, 14:209. Cymbiola. 1914. NPRMP, 1(3): 109 (Superpatagonico). = Adelomelon (Pachycymbiola) ameghinoi (Ihering), new combination. Type Locality. — La Cueva, Santa Cruz. Holotype.— #SS3a, coll. Ameghino (by monotypy). Other Lors.— #884, La Cueva, a very large specimen, coll. Ameghino; #885, Rada Tilly, San Jorge, one specimen, coll. Ameghino; #886, Golfo San Jorge, one specimen, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.— Specimen #295 differs from typical ameghinoi, being more globose and without tuberculae. This species is ancestral to A. (P.) feruglioi (Doello- Jurado, 1931) from the Pliocene and the living A. (P.) brasiliana (Solander, 1786). 202 Annals of Carnegie Museum VOL. 65 Ameghinomya, 1907. VENERIDAE 1907a. AMNBA, 1907:71, 306. New subgenus. = Protothaca Dali, 1902. Type Species.— Chione argentina (Ihering, 1897Z?). Stratigraphy. — Patagonian Formation (Lower Miocene) and Parana Formation (Upper Miocene). Remarks. — Close to Chione and according to Ihering represented only in the Patagonian, but poorly defined. Although Ihering mentioned Protothaca in rela- tion with other forms (see Chione cossmanni), he did not compare Ameghinomya with Protothaca. See Venus volckmanni argentina. ameghinorum, 1907. Venericardia 7ARCIDAE 1907a. AMNBA, 14:23, pi. 3, fig. 13a-b. 1914. NPRMP, 1(3):42 (Roca). = INoetia ameghinorum (Ihering). Type Locality. — Roca, Rio Negro. Holotype. — #340, an artificial cast received from Florentino Ameghino in 1904 which corresponds to the figured specimen (by monotypy). Stratigraphy. — Roca Formation (Paleocene). Remarks. — This species has high umbos anteriorly produced and probably be- longs to Noetia\ unlikely a Venericardia. americanus^ 1907. Neomphalius TROCHIDAE 1907a. AMNBA, 14:134, pi. 4, fig. 7a-b. 1914. NPRMP, 1(3):76 (Superpatagonico). = Tegula (Promartynia) americana (Ihering), new combination. Type Locality. — Canada de los Artilleros, Santa Cruz. Holotype. — #592, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — The shell is high, Calliostoma-Xike but with an umbilicum “circon- scrit en dehors par una crete spirale,” as in the subgenus Promartynia Dali, 1909. archipatagonica, 1907. Urosalpinx MURICIDAE 1897Z?. RMP, 2:297. Trophon leucostomoides (not Triton leucostomoides Sowerby, 1846). 1907a. AMNBA, 14:189, pL 6, fig. 35. 1914. NPRMP, 1(3):98 (Superpatagonico). = Ocenebra (Ocinebrina) elegans (Ortmann), new combination. Type Locality. — La Cueva, Santa Cruz. Holotype. — #795, coll. Ameghino, 1900 (by monotypy). Other Lot. — #796, Yegua Quemada, one specimen, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This is a more elongate species than the Recent Urosalpinx rushi (Pilsbry). archiplata “Ihering,” 1931. Bulimus STROPHOCHEILIDAE 1931. Walther, p. 31, given as “fi. archiplata Ihering” in litteris. Nomen nudum. = Strophocheilus charruanus (Frenguelli, 1930). 1996 Parodiz — Ihering Mollusca 203 Remarks. — See Klappenbach and Olazarri (1966:234) and Parodiz (1969:172, pi. 18, fig. 2). arenmia, 1907. Bulla BULLIDAE 1907<3. AMNBA, 14:339. New name for Bulla remondi in Ortmann (1899:431) and Ortmann (1902: 245, pi. 37, fig. 7a-=-b), not Bulla remondi Philippi, 1887. Type Locality. — -The specimen figured by Ortmann (1902) is from Punta Are- nas, Chile. Stratigraphy. — Magellanian Formation (Lower Oligocene). arenophila, 1907. Macrocallista VENERIDAE 1907«. AMNBA, 14:349. New name for Meretrix iheringi in Ortmann (1902:142, pi. 28, fig. 5a-b), not Meretrix iheringi Cossmann, 1898. = lAmiantis arenophila (Ihering). Type Locality.— In Ortmann (1902): Punta Arenas, Chile. Stratigraphy. — Magellanian Formation (Lower Oligocene). argemtinaf 1897. Turritella TURRITELLIDAE 1897/?. RMP, 2:286. Not figured (Form. Patagonica). 1907a. AMNBA, 14:162. Turritella ambulacrum argentina. 1914. NPRMP, 1(3):88. Turritella ambulacrum argentina. = Turritella ambulacrum Sowerby, 1846. The different “varieties” of T. ambu- lacrum— T. a. steinmanni, T. a. pyramidesia, and T. a. argentina— dim all the same species. Type Locality. — Rfo Santa Cruz, Santa Cruz. Syntypes. — Not in the collection. Stratigraphy.— Pdiidigonmn Formation (Lower Miocene). Remarks. — The type lot with 17 specimens as reported by Ihering (1914) is not in the collection. The other lot, #709, is a conglomerate. argentina, 1897. Venus volckmanni “van” VENERIDAE 1897/?. RMP, 2:252, pL 7, fig. 45. 1904. RMLP, 11:231. Venus argentina. 1907a. AMNBA, 14:291, 385. Chione (Ameghinomya) argentina. 1914. NPRMP, 1(3):55 (Superpatagonico). — Protothaca argentina (Ihering). Type Locality. — Santa Cruz. Syntypes. — Not in the collection as listed in Ihering (1914). Other Lots.— ^460, Rada Tilly, two specimens, coll. C. Ameghino, 1900; #461, Trelew, Chubut, a fragment, coll. S. Roth (from “formagao entreriana”); #462, Sierra de los Baguales, three casts, coll. Hauthal, 1897; no number, Santa Cruz, one specimen, coll. J. Bicego, 1897. Stratigraphy. — -Patagonian Formation (Lower Miocene) and Parana Formation (Upper Miocene). Remarks. — Venus paranensis Borchert, 1901:38, pi. 3, fig. 16-18, from Parana, Entre Rios, is a synonym. E. Feruglio also found it in the Patagonian of Cerro Otto. 204 Annals of Carnegie Museum VOL. 65 argentinus, 1907. Actaeon ACTAEONIDAE 1907«. AMNBA, 14:117, pL 4, fig. 2. 1914, NPRMP, 1(3):68 (Superpatagonico). Type Locfllilj. —“Canada de Los Artilleros, Santa Craz. Syntypes.—^545, two specimens, coll. Ameghino, 1900. Stratigraphy.— Formation (Lower Miocene). Austmcomimella, 1907. BUCCINIDAE: PHOTINAE 1907fl. AMNBA, 14:190, 344. New subgenus. Type Species.— Cominella fueguensis Ihering. Stratigraphy, — Patagonian Formation (Lower Miocene). “Subgenus, “intermediaire entre Cominella s.s. et PtychosalpinxS' Cominella sensu stricto and other subgenera are from New Zealand; PtychosaT pinx is known only from the Miocene of North America. bagualesia, 1907. Panopaea [sic] HIATELLIDAE 1907£i. AMNBA, 14:325, pi. 13, fig. 86a-b. 1914. NPRMP, 1(3):63. = Panopea bagualesia Ihering. Type Locality.— Sictxdi de los Baguales, Santa Cruz [given by Ihering (1914) as Chubut in error]. Holotype.^ — #511, one cast with part of the shell, coll. Hauthal, 1897 (by mono^ typy). Stratigraphy.— V^idigonmn Formation (Lower Miocene). Feruglio (1949-1950) reported the species from Rfo Turbio. Barnea sp. PHOLADIDAE 1914. NPRMP, 1(3):65. = ICyrtopleura lanceolata (d'Orbigny, 1846). Specimens.— tt522, one cast, from la Paz, Entre Rios, coll. S. Roth. Stratigraphy.— PdLmn^ Formation (Upper Miocene), Similar to Barnea omata Borchert, 1901:45, pL 4, fig. 3. bellicosum, 1907. Callio stoma TROCHIDAE 1907a. AMNBA, 14:439, pi. 17, fig. 117. 1914. NPRMP, 1(3):77 (“Post-Pampeano”). = Calliostoma militaris Ihering. Type Locality.— Pucilo Militar (now Puerto Belgrano), Prov. Buenos Aires. Syntypes. — 4t591, three specimens, coll. Ameghino, 1905. Stratigraphy.— Belgrano Formation (Pleistocene). Recent: Rio de Janeiro to Prov. de Buenos Aires. Remarks.— Clench and Turner (1960:23) listed this species as a synonym of Calliostoma jucundum (Gould), but it is actually C. militaris; the descriptions of the two species are almost identical. bicegoiy 1899. Lotorium 1899a. NJMGP, 2:29, pi. 1, fig. 8. 1907a. AMNBA, 14:177 (Patagonico). RANELLIDAE: CYMATIINAE 1996 Parodiz— -Ihering Mollusca 205 1914. NPRMP, 1(3): 97. Lotorium (Sassia) bicegoL = Cymatium bicegoi (Ihering). Type Locality. —SsmtSL Cruz. Syntypes.— ^790, two specimens, coll. J. Bicego. Stratigraphy.— Patagonian Formation (Lower Miocene). Remarks.— Of this and other species of “Lotorium/' Ihering (1907fl:174) said that he did not know the correct stratigraphic position; however, he mentioned Upper Eocene (unlikely) and Superpatagonian. boehmi, 1907. Venericardia iheringi “var.” CARDITIDAE 1907a. AMNBA, 14:23. Not figured. = Venericardia iheringi Bohm, 1903. Type Locality.— R.oca, Rio Negro. Not in the collection under this name. Stratigraphy.— Koca Formation (Paleocene). Remarks two specimens “plus ventra, moins hault” were included in 1914 in the lot of Venericardia iheringi (#355); Ihering added: “PeuLetre c’est une variation individuelle.” borchertiy 1907. Tritonidea BUCCINIDAE: PISANIINAE 1907a. AMNBA, 14:369, New name for Turbinella subcraticulata d’Orbigny in Borchert (1901:50, pi. 5, fig. 1-2), not T. subcraticulata d’Orbigny. Type Locality.— Parana, Entre Rios (Borchert). Stratigraphy.— Parana Formation (Upper Miocene). Remarks.— The genus Tritonidea Swainson, 1840, for this species was sug- gested to Ihering by W. H. Dali. bruggeni [sic], 1921. Aturia (Sphenaturia) NAUTILIDAE 1921. Physis, Buenos Aires, 5:76. Not figured. = Aturia (Sphenaturia) brueggeni Ihering. Type Locality.— Punta Boqueron, Bahia Inutil, Tierra del Fuego. Status unknown. Stratigraphy.— “Pstrato^ de Boqueron” (below Patagonian Formation). Referring to the holotypes of this species and A. (5.) felschi, Ihering (1921) stated, “Los ejemplares tipicos quedan depositados en este Museo [Museo Argentieo de Ciencias Naturaies, Buenos Aires].” This species is the type of the subgenus Sphenaturia Ihering. Femglio (1949-1950) places it below the Pata- gonian. bullia, 1907. Melania PLEUROCERIDAE 1907a. AMNBA, 14:464, pL 18, fig. 127. 1914. NPRMP, 1(3):93 (Saiamanqueano). = Paleoanculosa bullia (Ihering), see Parodiz, 1969:125, pi. 12, fig. 7-9; pL 14, fig. 2; pL 10, fig. 2-4, 6. Type Locality.— Rio Chico, Chubut. Holotype.—#153, coll. Ameghino (by monotypy); deposited in CMNH. Stratigraphy.— Colhue Huapi Formation (Colhuehuapian; Thanetian-Montian, ? Paleocene). 206 Annals of Carnegie Museum VOL. 65 Remarks.— Melania pehuenchensis Doello=Jurado (1927:415; see Wichmann, 1927:fig. 77) and M. ameghiniana Doello-Jurado (1927:413; see Wichmann, pi. 10, fig. 61-71, and pL 11, fig. 74) are synonyms of P. bullia (Ihering). The holotype was received by Parodiz (1969:pl. 14, fig. 2) in exchange from Dr. Max Biraben, at that time Director of the Museo Argentine de Ciencias Naturales. burckhardti, 1903. Cardita CARDITIDAE 1903. AMNBA, 9:205. New name for Cardita morganiana in Burckhardt (1902:216), not Cardita morganiana Rathbun, 1875. Type Locality. — Malargue, near Roca, Rio Negro. Type. — No material described by Burckhardt is in the collection. Stratigraphy. — Roca Formation (Paleocene). Remarks. — Specimens from Roca were misidentified by Burckhardt (1902) as Cardita morganiana Rathbun, but that species is from the Maria Farinha For- mation in Brazil. burckhardtiy 1907. Dosinia VENERIDAE 1907a. AMNBA, 14:26. New name for Dosinia brasiliensis Burckhardt, 1902:216, pi. 4, fig. 8-12, not D. brasiliensis White, 1887. 1914. NPRMP, 1(3):50 (Cretacee Superieur). Type Locality. — Roca, Rio Negro. Type. — No material described by Burckhardt is in the collection. Stratigraphy. — Roca Formation (Paleocene). burckhardti, 1903. Turritella TURRITELLIDAE 1903. AMNBA, 9:208, pi. 1, fig. 9. 1907a. AMNBA, 14:38. 1914. NPRMP, 1(3):90. Turritella doeringi Bohm (Cretacee Superieur). = Turritella doeringi Bohm, 1903. Type Locality. — Roca, Rio Negro. Syntypes. — #729, three casts, coll. Santiago Roth. Stratigraphy. — Roca Formation (Paleocene). burmeisteri, 1907. Cymbiola VOLUTIDAE 1897/?. RMP, 2:303. Valuta dorbignyana Philippi. 1899a. NJMGP, 2:33. Voluta dorbignyana Philippi. 1902. Ortmann, p. 230, pi. 36, fig. 1 (V. d’orbignyana), and p. 232, fig. 37a-b (V. domeykoana, not domeykoana Philippi). 1907a. AMNBA, 14:210. Cymbiola burmeisteri Ihering. 1914. NPRMP, 1(3): 109-1 10. Cymbiola {Cymbiola) burmeisteri Ihering and Cymbiola {Miomelon) orbignyana [sic] Philippi (Patagonico). = Adelomelon burmeisteri (Ihering), new combination. Type Locality.— Cruz. Syntypes.^ — #880, two specimens plus four casts, Bicego Collection; #881, Si- erra de los Baguales, one complete specimen, coll. Hauthal. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Ortmann’s figures (1902) of V. dorbignyana and V. domeykoana are similar, and those on pi. 36, fig. lb, d, e, are juveniles. Figures la-lb in Ortmann (1902) are Adelomelon {P achy cymbiola) pilsbryi (Ihering). See Voluta pilsbryi Ihering. 1996 Parodiz-=-Ihering Mollusca 207 burmeisteri^ 1907. Dosinia VENERIDAE 1907a. AMNBA, 14:301, text fig. Oa-b. 1914. NPRMP, 1(3):50 (Patagonico Inferior). = Dosinia (Dosinella) burmeisteri (Ihering). Type Locality.— Bdijo de Sae Julian, Santa Craz. Holotype.—4A\6, one valve, colL Ameghino, 1900 (by monotypy). Stratigraphy.— Vdiidigonmn Formation (Lower Miocene). Remarks.- — According to Dali (in consultation with Ihering), this species be- longs to the subgenus Dosinella. burmeisteri^ 1907. Psammobia PSAMMOBIIDAE 1907a. AMNBA, 14:312, pL 11, fig. 81a-b. 1914. NPRMP, 1(3):57 (Patagonico Inferior). = Gari burmeisteri (Ihering). Type Locality.— CMtiMonQS, Chubut. Syntypes.—#411 , only three valves in the collection from the six reported by Ihering (1914). Stratigraphy.— PdilAgomm. Formation (Lower Miocene). Remarks.- — E. Feruglio (1949-1950) reported the species from Rio Turbio. burmeisteri^ 1907. Tellina TELLINIDAE 1907a. AMNBA, 14:27, pL 3, fig. 16. Type Locality. — -Roca, Rio Negro. Type.— Not in the collection. Stratigraphy.— Koc^ Formation (Paleocene). Remarks.— Ihoiing {1901a) states, “Seulement un moule interne.” The figured internal cast appears to be of the subgenus Strigilla Turton, 1832, and close to S. carnaria, a Recent species ranging from Florida to Bahia Blanca, Argentina. Iher- ing did not compare this species with living ones because he thought it was from the Cretaceous. cahuillensis, 1907. Turritella TURRITELLIDAE 1907a. AMNBA, 14:342. New name for Turritella affinis in Hupe, 1854:155 (pi. 2, fig. 7), preoccupied by Turritella affinis Muller, 1851, stated by Ihering to be a Cretaceous species. 1914. NPRMP, 1(3):90 (“Terciario de Chili”). Type Locality. — Curaumulli, Valparaiso, Chile. Sy ntypes. SI 27 , received from the Valparaiso Museum, three specimens, are possible syntypes. Stratigraphy. — Tertiary. Remarks.— 1 have not been able to locate a reference for Muller, 1851, as given in Ihering (1907a: 342). It is possible that Ihering never saw original descriptions for T. affinis Muller or T. affinis Hupe, and his use of the latter name was based on a questionable identification in Philippi (1887:71). In any event, the name cahuillensis reflects the type locality of that name, given by Hupe (1854:155) as Inseln Chiloe, Cahuil, Colchagua. Ihering (1897^:289) indicated that Moericke and Steinmann (1896:555) identified a specimen as T. affinis Philippi, and without explanation stated his belief that the species was probably the same as T. affinis Hupe. This conclusion appears consistent with later placement of "'Turritella af- 208 Annals of Carnegie Museum VOL. 65 finis R. A. Philippi {nec Hupe)” as a synonym of T, cahuillensis (Ihering, 1914: 90). Without study of original descriptions and type materials, it is not possible to determine the correct synonymy for specimens identified by various authors as T. ajfinis. callophyla, 1903. Exogyra OSTREIDAE 1903. AMNBA, 9:199. Not figured. 1907a. AMNBA, 14:10, 144. 1914. NPRMP, 1(3):20 (Cretaceo Superior, Roca). Type Locality. — Roca, Rio Negro. Holotype. — #138, one specimen (by monotypy). Other Lots. — #139, Roca, two complete specimens and five valves, coll. Ameghino, 1907; #140, Rio Negro, one specimen, coll. Ameghino, “Cretaceo horizonte mais inferior.” Stratigraphy. — Roca and Salamanca formations. Remarks. — Feruglio found the species in the Salamancan beds north of Colhue Huapi in the lower valley of the Rio Negro in Chubut (Paleocene). Burckhardt (1902:215) referred to it as Exogyra lateralis Nilsson; the original lateralis of Nilsson is not an Exogyra, but a Gryphaea. camaronesia, 1907. Area ARCIDAE 1907a. AMNBA, 14:238, text fig. 9. 1914. NPRMP, 1(3): 18 (Patagonico Inferior). = Anadara carnaronesia (Ihering), new combination* Type Locality. — Camarones, Chubut. Syntypes. — #104, 34 valves, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — The specimen figured in Ihering (1907a) is a juvenile, similar to Anadara brasiliana (Lamarck). carnaronesia^ 1907. Crenella divaricata MYTILIDAE 1907a. AMNBA, 14:277, text fig. lla-b. 1914. NPRMP, 1(3):39 (Patagonico Inferior). = Crenella carnaronesia (Ihering). Type Loca/ily.— Camarones, Chubut. Syntypes. — #323, 12 valves, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.— It differs from Crenella divaricata (d’Orbigny), a Recent species ranging from North Carolina to Argentina, by being narrower and more squamose. This species resembles Crenella duplisensis Dali of the Tertiary of Florida. carnaronesia^ 1907. Diplodonta villardeboaena [sic] “var.”' UNGULINIDAE 1907a. AMNBA, 14:290. Not figured. 1914. NPRMP, 1(3):47. = Diplodonta vilardeboaena (d’Orbigny, 1846). Type Locality. — Camarones, Chubut. Syntypes.— #390, three valves, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). 1996 Parodiz — Ihering Mollusca 209 Remarks. — Taras Risso, nomen dubium (= Diplodonta). camaromesia, 1907, Glycimeris [sic] GLYCYMERIDIDAE 1907fl. AMNBA, 14:240, pi. 7, fig. 52a-b. 1914. NPRMP, 1(3): 18 (Patagonico Inferior). = Glycymeris camaronesia Iherieg, Type Locality. — Camarones, Chubut. Syntypes. four large valves (five smaller valves mentioned in Ihering [1914] missing from the collection), coll. Ameghino, 1900. “Patagonian Formation (Lower Miocene). camaronesia, 1907. Leda NUCULANIDAE 1907a. AMNBA, 14:230, pL 7, fig. 50a™b. 1914. NPRMP, 1(3): 14 (Patagonico Inferior). = Nuculana (Sacceila) camaronesia (Ihering). Type Locfllilj. “Camarones, Chubot. Syntypes.—#H0, one complete specimen (Ihering listed four) plus 22 isolated valves, coll. Ameghino, 1900; #81, south of Colhue Haupi, coll. Ameghino, 1900. ftrarigrap/iy. “Patagonian Formation (Lower Miocene). Remarks.— Leda Schumacher, 1817 (= Nuculana Link, 1807). camaronesia, 1907. Tellina patagonica “var.” TELLINIDAE 1907a. AMNBA, 14:315. Not figured. 1914. NPRMP, 1(3):58 (Patagonico Inferior). = Tellina patagonica Ihering. Type Loca/fry. “Camarones, Chubut. Syntypes.-^A^l , two valves; coll. Ameghino, 1900. Stratigraphy.— Formation (Lower Miocene). Remarks. — The syntype valves differ from each other and represent variation in T. patagonica. camaronesia, 1907. Tornatina SCAPHANDRIDAE [CYLICHNIDAE] 1907a. AMNBA, 14:119, pL 4, fig. 3. 1914. NPRMP, 1(3): 68 (Patagonico Inferior). Type Locality.— Chubut. Syntypes. — #547, ten specimens, coll. Ameghino, 1900. “Patagonian Formation (Lower Miocene). camaronesia, 1907, Turritella breantiana “var.” TURRITELLIDAE 1907a. AMNBA, 14:164, pi. 4, fig. 23. 1914. NPRMP, 1(3):89 (Patagonico Inferior). = Turritella breantiana d’Orbigny, 1847. Type Lom/ily. “Camarones, Chubut. Syntypes.— #119, 15 specimens, coll. Ameghino, 1900. Stratigraphy.— Vdii^gonim. Formation (Lower Miocene). .^^wari^.“This form was among those reported by Ihering (1899a) as T. breantiana indecussata. Although both are of the same strata, T. camaronesia is 210 Annals of Carnegie Museum VOL. 65 found only north of the San Jorge Gulf, while indecussata is more abundant south of it. camaronesia, 1907. Venericardia CARDITIDAE 1902. Ortmann, p. 128, fig. Sa-c. Cardita patagonica (not C. patagonica Sowerby, 1846). 1907a. AMNBA, 14:28, pi. 10, fig. 69a-d (Patagonico Inferior). Type Loca/ity.— Camarones, Chubut. Syntypes.—^?>A9, 100 valves, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). camaronesium, 1907. Calliostoma TROCHIDAE 1907a. AMNBA, 14:137, pi. 4, fig. 11. (Only one figure is on plate 4; text says “a-b.”) 1914. NPRMP, 1(3):77 (Patagonico Inferior). Type Locality. — Camarones, Chubut. Holotype. — #598, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Ihering compared this species with Calliostoma santacruzense Ort- mann, but it is more similar to C. cossmanni Ortmann. cannada, 1907. Cymbiola becki “van” VOLUTIDAE 1907a. AMNBA, 14:211, pi. 6, fig. 42a-b. 1914. NPRMP, 1(3): 109 (Superpatagonico). = Adelomelon (Adelomelon) cannada (Ihering), new combination; new status. Type Locality.— Cdifididdi de los Artilleros, Santa Cruz. Holotype. — #879, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species is ancestral to the living A. (A.) becki (Broderip) and A. (A.) barattini Klappenbach and Ureta, 1966. cannada„ 1907. Venericardia CARDITIDAE 1907a. AMNBA, 14:287, pi. 10, fig. 71a-b. 1914. NPRMP, 1(3):42 (Superpatagonico). Type Locality.— CdiVididdi de los Artilleros, Santa Cruz. Syntypes.^3AA, 38 valves, coll. Ameghino, 1900; #345, Pan de Azucar, one valve, coll. Ameghino, 1900; #346, Bajo San Julian, two valves, coll. Ameghino, 1900; #347, La Cueva, seven valves, coll. Ameghino, 1900; #348, Yegua Que- mada, 15 valves, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). cannada, 1907. Volvulella RETUSIDAE 1907a. AMNBA, 14:119, pi. 4, fig. 4. 1914. NPRMP, 1(3):69 (Superpatagonico). Type Locality.— Cdifididdi de los Artilleros, Santa Cruz. Syntypes. — #549, two specimens, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species resembles the Recent species V. persimilis (Morch) which ranges from North Carolina to Uruguay. 1996 Parodiz — Ihering Mollusca 211 capa^ 1907. Ostrea hemisphaerica [sic] ''van” OSTREIDAE 1907fl. AMNBA, 14:12. Not figured. 1914. NPRMP, 1(3):22. Ostrea neuquena capa (Rocaeense). = Ostrea neuquena Iherieg. Type Locality.^ — -Roca, Rio Negro. Syntypes.-^^151 , five valves, plus three complete. Stratigraphy.— Formation (Paleocene). Remarks.— Them are no Ostrea hemispherica d'Orbigny in the collection. See Ostrea hemisphaerica [sic] "van” paca Ihering. camloameghinoiy 1902. Aturia NAUTILIDAE 1902fe. PAPS, 41(69):134, pi. 19, fig. i=-2. 1907a. AMNBA, 14:116; '"caroli-ameghinoi.” 1914. NPRMP, 1(3):68 (Patagonico Medio). Type Locality.- — ^Cabo Tres Puntas, Santa Cruz. Holotype.- — #544, coll. Ameghino, 1900 (by monotypy). Stratigraphy.— Vdiidigonimi Formation (Lower Miocene); also in the “Estratos de Boqueron,” Tierra del Fuego. carolodarwimiy 1907. Polynices NATICIDAE 1907a. AMNBA, 14:89. Nomen nudum; see Natica tumbeziana Ihering. cma^ 1907. CMone VENERIDAE 1907fl. AMNBA, 14:308, pi. 12, fig. 78. 1914. NPRMP, 1(3):55 (“Casa-Maior,” Patagonico Inferior). = Protothaca antiqua (King, 1832). Type Locality— Tmii^ Casamayor, Santa Cruz. Holotype.— ^456, coll. Ameghino, 1900 (by monotypy). Other Lot.-^#451 j Travesia Valcheta, coll. C. Burmeister, one fragment. Stratigraphy.— P^ildigomwi Formation (Lower Miocene). Remarks.— Thi^ is the oldest geological record of the Recent P. antiqua. chalcedomica^ 1903. Cytherea VENERIDAE 1903. AMNBA, 9:216, pL 2, fig. 18a-b. 1907a. AMNBA, 14:48. Meretix chalcedonica. 1914. NPRMP, 1(3):51 (Salamanqueen). [“Posigao systematica dubiosa”]. Type Locality.— Colhue Huapi, Chebut. Syntypes.— #422, three valves, crystalized into chalcedony. Slrarigrap/iy.-^-Salamanca Formation (Paleocene). ^^wari^.-^-This species is neither a Cytherea nor a Meretrix; it most likely belongs in Pitar. cMcoama, 1907. Hadriania jorgensis "van” MURICIDAE 1907a. AMNBA, 14:186, pL 6, fig. 33a--b. 1914. NPRMP, 1(3):99 (Patagonico Inferior). = ITrophon chicoanus (Ihering), new combination. Type Locality. — -Rio Chico, Chubut. 212 Annals of Carnegie Museum VOL. 65 Holotype. — #806, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Differs from Nucella jorgensis (Hadriania in Ihering, 1907a) and does not belong in that genus. chicoanus, 1907. Vermetus VERMETIDAE 1907«. AMNBA, 14:167. New name for Vermetus cf. intortus in Ortmann (1902:198, pi. 32, fig. 1), not Serpula intorta Lamarck, 1818. = IPetaloconchus chicoanus (Ihering). Type Locality. — Rio Chico, Chubut (Ortmann, 1902). Also from Lake Pueyr- redon. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Vermetus intortus (Lamarck) is a European species currently placed in Petaloconchus H. C. Lea, 1843. chubutensis, 1903. Rostellaria 7APORRHAIDAE 1903. AMNBA, 9:220, pi. 2, fig. 17. 1907a. AMNBA, 14:29. Aporrhais (Rocaneen). 1914. NPRMP, 1(3):94 (Salamanqueen). Type Locality. — Rio Chico, Chubut. Syntypes. — #756, two specimens, coll. Ameghino, 1900. Stratigraphy. — Salamanca Formation (Paleocene). Remarks. — Ihering transferred this species to the Aporrhais but it is more likely a species of Struthiolaria. chubutensis, 1904. Strophocheilus STROPHOCHEILIDAE 1904. RMLP, 2:241, pi. 2, fig. 13-14 (“Notostylops” beds). 1907a. AMNBA, 14:463. 1914. NPRMP, 1(3):70 (Cretaceo Superior, “Notostylopeano”). 1969. Parodiz, ACM, 40:170. Type Locality. — Canadon Blanco, south side of Lake Colhue Huapi, Chubut. Lectotype. — #560, cast, coll. Ameghino (reported in error as Hauthal by Ihering (1914). Stratigraphy. — Casamayor Formation (Lower Eocene). Remarks. — The lectotype was designated by Parodiz (1949/7:181, fig. la) as the specimen figured in Ihering (1904, pi. 2, fig. 13). Parodiz (1949/?) also clarified the status of 15 paralectotypes. As for 5. hauthali, the collector of the type lot was C. Ameghino, not Hauthal as reported by Ihering (1914). Three of the para- lectotypes from Rio Chico, Chubut (#560), became the holotype and paratypes for Strophocheilus avus Parodiz (1949/?: 181), a name later synonymized with S. {Megalobulimus) hauthali (Ihering) by Parodiz (1969). chubutiana, 1907. Scalar ia EPITONIIDAE 1907a. AMNBA, 14:364, pi. 14, fig. 99. = Epitonium chubutianum (Ihering), new combination. Type Locality. — Puerto Piramides, Chubut. Type. — Not in the collection. Stratigraphy. — Rio Negro Formation (Upper Miocene). 1996 Parodiz — Ihering Mollusca 213 Remarks. — Ihering (1907^:465) reported this species in a perforation from Pu- ente Alsina at 73 m. clarae, 1907. Ostrea OSTREIDAE 1907«. AMNBA, 14:10, pi. 1, fig. 6a, pi. 2, fig. 6a-c. 1914. NPRMP, 1(3):22 (Roca). Type Locality. — Roca, Rio Negro. Syntypes.- — #158, six valves, coll. Ameghino, 1904. Stratigraphy. — Roca Formation (Paleocene). Remarks. — Another lot is in the collection from Park- Aik (1907) or “Par-aiqu” (1914, one valve, “formagao guaranitica, classificagao incerta”)- clessini, 1907. Turbonilla PYRAMIDELLIDAE: TURBONILLINAE 1907«. AMNBA, 14:145. New name for Turbonilla iheringi Clessin (1900:174, pi. 36, fig. 7), not T. iheringi Cossmann (1899fif:235, pi. 10, fig. 12), and not T. iheringi Clessin (1900:168-169, pi. 35, fig. 5). Type Locality. — “Siidbrasilien, Igt. v. Ihering, Mus. BeroL,” as given in Clessin (1900:174). Stratigraphy.— V^itdigonm Formation (Lower Miocene). Remarks. — Clessin (1900) separately described two specimens under the name Turbonilla iheringi without mention of their primary junior homonymy with each other and with the senior valid name T. iheringi Cossmann, 1899. Ihering (1907<3) observed that Clessin’s material consisted of two species, the first of which he let stand as T. iheringi Clessin (1900:168) despite its junior status to T. iheringi Cossmann. The other species {T. iheringi Clessin, 1900:174) was named Turbon- illa clessini by Ihering. He noted that Cossmann had already used T. iheringi, and without regard for priority of the available names he offered Turbonilla eopa- tagonica as an unnecessary replacement name for the valid senior name, T. iher- ingi Cossmann. Turbonilla iheringi Clessin (1900:168) remains a junior homonym of Cossmann’s name, and requires a replacement name. colhuapensis, 1903. Diplodon HYRIIDAE 1903. AMNBA, 9:217, fig. 2. 1907a. AMNBA, 14:466 (“. . . depots qui sont probablement tertiares.”) 1914. NPRMP, 1(3):36 (Cretaceo). 1969. Parodiz, ACM, 40:53-54. Type Locality. — Colhue Huapi, Chubut. Lectotype. — #295, coll. Ameghino; in CMNH. Stratigraphy. — Sehuen Formation (Paleocene). Remarks.— The lectotype was designated by Parodiz, 1969:54, pi. 1, fig. 1, 3- 4 (fig. 2 is a paratype) based on material received in exchange from Dr. Max Biraben, at that time Director of the Museo Argentino de Ciencias Naturales. This species will be included in a new genus (Parodiz, in preparation). Cominella sp. BUCCINIDAE: PHOTINAE 1914. NPRMP, 1(3): 105 (Patagonico). Locality. — Punta Atlas, Chubut. Specimens. — #859, coll. Ameghino. 214 Annals of Carnegie Museum VOL. 65 Stratigraphy. — Patagonian Formation (Lower Miocene). concorSy 1902. Gryphaea GRYPHAEIDAE 1902a. AMNBA, 4:11, 114, fig. 1-3 (Formation Guaranitique). 1903. AMNBA, 9:210. 1904. RMLP, 11:210. 1907a. AMNBA, 14:8, 43 (Salamanqueen). 1914. NPRMP, 1(3):21 (“etage guaranieen”). Type Locality. — Valle Alsina, Chubut. Syntypes. — #144, five valves, coll. G. Burmeister, 1887. Other Lot. — #145, Pico Salamanca, two valves. Stratigraphy. — Salamanca Formation (Paleocene). confiniSy 1897. Marginella MARGINELLIDAE 1897Z?. RMP, 2:307, pi. 3, fig. 8 (Formagao Santacruzense). 1907a. AMNBA, 14:200. Neoimbricaria (Superpatagonico). = Neoimbricaria quemadensis (Ihering). Type Locality. — La Cueva, Santa Cruz. Holotype. — #866, coll. Ameghino (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Marginella gracilior Ihering is another synonym of N. quemadensis (Ihering). consimilis, 1897. Natica NATICIDAE 1897Z?. RMP, 2:283, text fig. 12 (Formagao Santacruzense). 1899a. NJMGP, 2:28. Natica famula Philippi. 1907a. AMNBA, 14:156. Polynices magellanica “van” 1914. NPRMP, 1(3):85. Polynices magellanica {consimilis) (Superpatagonico). = Lunatia magellanica Hombron and Jacquinot, 1834. Type Locality. — La Cueva, Santa Cruz. Syntypes. — #672, two specimens, coll. Ameghino; #673, San Jorge, ten speci- mens, coll. J. Bicego; #674, Golfo San Jorge, two specimens, coll. Ameghino; #675, Deseado (Formacion Araucana), one specimen, coll. Ameghino, 1900. Stratigraphy, — Patagonian Formation (Lower Miocene) and also Araucanean Formation (Pliocene). Remarks. — Natica famula Philippi referred to by Ihering (1899(3) is a species from Chile, different from consimilis, although it is also found in Santa Cruz (#668~#670 in the Ihering Collection). It is related to a longer species, Lunatia pyrotheriorum Ihering from the Patagonian. Conus sp. CONIDAE 1907a. AMNBA, 14:448. = Conus clenchi Martins, 1943. Recent. Locality. — Reported from “Pampien de Puerto Militar aff. C. portoricanus Hwass.’' The type locality of Conus clenchi Martins is Puerto Deseado, Santa Cruz. Stratigraphy. — Belgrano Formation (Lower Pleistocene). Remarks. — Conus carcellesi Martins, 1945, is a synonym of C. clenchi ac- cording to Rios (1975:124). 1996 Parodiz — Ihering Mollusca 215 Corbiculella, 1907. 1907a. AMNBA, 14:462, 469. New subgenus of Corbicula Megerle. 1969. Parodiz, ACM, 40:98. Subgenus of Neocorbicula Fischer. CORBICULIDAE Type Species. — Corbicula tenuis Ihering. Stratigraphy. — Parana Formation (Upper Miocene). cossmanni, 1907. Amussium [sic] PROPEAMUSSIIDAE 1907a. AMNBA, 14:260, pi. 9, fig. 59a-b. 1914. NPRMP, 1(3):35 (Patagonico). = Amusium cossmanni Ihering. Type Locality. — Punta Atlas, Chubut. Holotype. — #288, one valve, coll. R. Valentin, 1897 (by monotypy). Stratigraphy. — Parana Formation (Entre Rios) and Rio Negro Formation (Ri- onegrense) in Chubut (Upper Miocene). Remarks. — Ihering (1907a) remarked that this species was found together with Ostrea hatcheri Ortmann. Amusium cossmanni resembles Amusium darwinianum (d’Orbigny), which is also from the Parana Formation. cossmanni^ 1907. Chione VENERIDAE 1907a. AMNBA, 14:350. New name for Venus chiloensis in Ortmann (1900:378) and Ortmann (1902: 137, pi. 27, fig. 10), not Venus chiloensis Philippi, 1887. = IProtothaca cossmanni (Ihering). Type Locality. — Punta Arenas, Chile (Ortmann). Stratigraphy. — -?Lower Miocene. Remarks. —WiQnng (1907a:350) remarked that chiloensis belongs to Protothaca while cossmanni has the sculpture of Chione, but that is not a generic definition. cossmanni, 1907. Cominella BUCCINIDAE: PHOTINAE 1902. Ortmann, p. 209, pi. 39, fig. 18. Buccinum obesum var. minor (Philippi). 1907a. AMNBA, 14:191 (Santacruzense). 1914. NPRMP, 1(3): 105. = Cominella minor (Philippi, 1887). Type Locality. — Santa Cruz. Holotype. — #857, received from Ortmann (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Feruglio (1949-1950) reported the species for Rio Turbio. cossmanni, 1907. Lima LIMIDAE 1907a. AMNBA, 14:264, pi. 9, fig. 62. 1914. NPRMP, 1(3):29. Lima (Limatula) (Patagonico Inferior). Type Locality. — Punta Nova (Nueva), Golfo San Jorge. Syntypes. — #225, three valves, coll. Ameghino; #226, Casamayor, one valve, coll. Ameghino, 1900. Other Lot. — #227, “. . . valva isolada que foi examinada pelo snr. Cossmann.” (Ihering, 1914). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — The three species of Lima described by Ihering are all from the 216 Annals of Carnegie Museum VOL. 65 same type locality and same stratigraphic position, but they are easily distin- guished. cossmanni^ 1907. Proscaphella VOLUTIDAE 1907a. AMNBA, 14:206, pL 6, fig. 41a~b. 1914. NPRMP, 1(3): 110, Cymbiola (Miomelon) (Patagonico). = Adelomelon {P achy cymbiola) cossmanni (Ihering). Type Locality. — Santa Cruz. Syntypes. — #888, two specimens, coll. J. Bicego. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Ihering (1907a): . . intermediaire entre P. gracilis et quemaden- sis. “ cossmanni, 1903. Rostellaria APORRHAIDAE 1903. AMNBA, 9:219, pi. 2, fig. 15. 1907a. AMNBA, 14:49. Aporrhais (Salamanqueen). 1914. NPRMP, 1(3):94 (Salamanqueano). = Aporrhais cossmanni (Ihering). Type Locality. — Rio Chico, Chubut. Syntypes. — #757, two specimens (Ihering [1907a]: “L’ unique exemplaire”), coll. Ameghino, 1900. Stratigraphy. — Salamanca Formation (Paleocene). Remarks. — The syntypes have the siphonal canal and peristoma missing but the body whorl is of Aporrhais. cossmanni, 1907. Scalaria EPITONIIDAE 1907a. AMNBA, 14:141, pi. 4, fig. 12. Primary junior homonym of Scalaria (Nodiscala) cossmanni de Boury, 1889. 1914. NPRMP, 1(3):80. Cirsotrema quemadense [sic] Cossmann (Superpatagonico). = Opalia {Nodiscala) quemadensis (Cossmann), new combination. Type Locality. — Yegua Quemada, Santa Cruz. Holotype. —^616, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.— Opalia {Nodiscala) eolis Clench and Turner, 1950, is a closely re- lated species. cossmanni, 1907. Tenagodus SILIQUARIIDAE 1907a. AMNBA, 14:166, pi. 5, fig. 25. 1914. NPRMP, 1(3):91 (Superpatagonico). = Siliquaria cossmanni (Ihering). Type Locality. — Canada de los Artilleros, Santa Cruz. Holotype. — #743, a small fragment of tube, coll. Ameghino, 1900 (by mono- typy)- Stratigraphy. — Patagonian Formation (Lower Miocene). crucis, 1907. Solen 1897^. RMP, 2:263. Solen elytron Philippi. 1907a. AMNBA, 14:318, pi. 12, fig. 83. SOLENIDAE 1996 Parodiz— Ihering Mollusca 217 1914. NPRMP, 1(3):59 (Superpatagoeico). Type Locality.— Ycgua. Quemada, Santa Cmz. Holotype.—4tA9l, colL Ameghino (by monotypy). Other Lot.— #492, Rada Tilly, Golfo San Jorge, one specimen, coll. Ameghino. Stratigraphy.— V^l^gonim. Formation (Lower Miocene). .^ewflrfo.=™This species resembles Solen scalprum Broderip. cruzemsis, 1907. Genotia TURRIDAE 1907a. AMNBA, 14:222. New oame for Genotia cuevemis in Ortmane (1902:240, pi. 37, fig. 2), not Genota cuevemis Ihering, 1897 (Superpatagonico). = Genota cuevensis (Ihering). Type Locality.— Mouth of the Santa Cruz River (Ortmann). Stratigraphy.— V^ldigommi Formation (Lower Miocene). -This name was unnecessarily proposed for large forms correctly identified by Ortmann (1902) as Genota cuevensis Ihering. Genotia Fischer, 1883 (= Genota H. and A. Adams, 1853). cruzialis^ 1907. Phacoides promaucana “var.” LUCINIDAE 1907fl. AMNBA, 14:288, pL 13, fig. 90a-=b. 1914. NPRMP, 1(3):45 (Superpatagonico). = Lucina cruzialis (Ihering). Type Locality. Yegua Quemada, Santa Cruz. Syntypes.—#314, coll. Ameghino. Other Lots.- — #375, Santa Cruz, three specimens, coll. J. Bicego, 1897; #376, Jack Harvey, one specimen; #377, Pan de Azucar, two specimens, coll. Ameghino, 1900; #378, Casamayor, 11 specimens, coll. Ameghino, 1900; #379, Cabo Tres Puntas, one valve, coll. Ameghino, 1900; #380, Punta Nodales, three specimens, coll. Ameghino, 1900; “Region de Sur,” Golfo San Jorge, two casts, coll. Amegh- ino, 1900. Stratigraphy.— P^ldk-goniMi Formation (Lower Miocene). cuevemsiSy 1897. Genota TURRIDAE 1897^. RMP, 2:313, pL 3, fig. 10. 1902. Ortmann, p. 240, pL 37, fig. 2. Genota cuevemis. 1907fl. AMNBA, 14:222. Genotia. 1914. NPRMP, 1(3):113 (Superpatagonico). Type Locality.- — La Cueva, Santa Cruz. Holotype.— #911, coll. Ameghino (by monotypy). Other Lots.— #912, Cabo Tres Puntas, one specimen, colL Ameghino; #913, Golfo San Jorge, coll. Ameghino. Stratigraphy.— P^dlB-gonim. Formation (Lower Miocene). Remarks. — -See note on G. cruzensis for Genota vs. Genotia. cuevensis, 1897. Gibbula fracta “var.” ARCHITECTONICIDAE 1897^. RMP, 2:274, pL 3, fig. 2. 1907a. AMNBA, 14:131. Gibbula cuevensis (Patagonico Superior), 1914. NPRMP, 1(3):72 (Superpatagoeico). = Valdesia dalli (Ihering). 218 Annals of Carnegie Museum VOL. 65 Type Locality.— To. Cueva, Santa Cruz. Holotype.—4t51\, coll. Ameghino (by monotypy). Other Lot. — #572, Canada de los Artilleros, six specimens, coll, Ameghino. Stratigraphy. ^ — -Patagonian Formation (Lower Miocene). cuevensiSy 1897. Pectunculus pulvinatus “var.” GLYCYMERIDIDAE 1897L RMP, 2:238, pi. 7, fig. 46 and pi. 8, fig. 50 (Form, santacrazeesis). 1907fl. AMNBA, 14:240, 345, 373. Glycimeris cuevensis [sic], 1914. NPRMP, 1(3): 19 (Superpatagonico and Magellanico). = Glycymeris cuevensis (Ihering). Type Locality. — La Cueva, Santa Cruz. Syntypes.— #122, two valves, coll. Ameghino. Other Lots. — #123, Yegua Quemada, six valves; #124, Santa Cruz, one valve, coll. J. Bicego; #125, Manantial Salado, one valve, coll. Ameghino, 1900; #126, Canada de los Artilleros, two large valves and 30 smaller valves, coll. Ameghino, 1900; #127, Puerto Piramides, Chubut, one valve, coll. Ameghino, 1903 (label says “from Dr. E Lahille”); #128, “Fin de Barrancas” [?], three valves from Dr. Lahille; #129, Punta Arenas (Magellanico), one specimen received from Ortmann under the name Pectunculus ibari Philippi. Stratigraphy. — Magellanian (Lower Oligocene) and Patagonian Formation (Lower Miocene). Remarks. — E. Feruglio (1949-1950) found the species at Rio Turbio. Glycy- meris pulvinatus Lamarck from the Tertiary of Europe has no connection with G. cuevensis. cuevemiSy 1897. Turbonilla PYRAMIDELLIDAE 1897L RMP, 2:276, text fig. 11 (Formagao Santacmzense). 1907«. AMNBA, 14:144. 1914. NPRMP, 1(3):81 (Superpatagonico). Type Locality. — La Cueva, Santa Cruz. Holotype.— #635, coll. Ameghino (by monotypy). Stratigraphy. — -Patagonian Formation (Lower Miocene). Remarks. — See note under Turbonilla observationis (Ihering). Cymbiola (Pachycymbiola) sp, VOLUTIDAE 1914. NPRMP, 1(3):110 (Forma^ao Entreriana ou Araucana). = Adelomelon {Pachycymbiola) feruglioi (Doello-Jurado, 1931). Locality. — The undetermined species of Ihering is from Puerto Piramides, Chu= but; the type of C. feruglioi Doello^Jurado is from Sierra Laziar, Santa Cruz. Specimens.- — #887, one incomplete specimen, coll. Hauthal. Stratigraphy. — Sierra Laziar (Laziarense), Araucanean Formation (Pliocene). Remarks. — This species is intermediate between Adelomelon {Pachycymbiola) ameghinoi (Ihering) and A. {P.) brasiliana (Solander). dalliy 1907. Calliostoma 1901 a. AMNBA, 14:437, pi. 17, fig. 114a-c. (Deseado Formation). 1914. NPRMP, 1(3):77 (Pampeano). = Calliostoma militaris Ihering. TROCHIDAE 1996 Parodiz — Ihering Mollusca 219 Type Locality. — Puerto Militar, Prov. Buenos Aires. Syntypes.— ^602, three specimens, coll. Ameghino, 1905. Stratigraphy. — Belgrano Formation (Lower Pleistocene). Remarks. — CalUostoma dalli Ihering (1907a), C, amazonicum Finlay (1930), and C. quequensis Carcelles (1944) are all synonyms of C. militaris. See C. bellicosum Ihering. dalli, 1907. Crassinella CRASS ATELLIDAE 1907a. AMNBA, 14:280, text fig. 12a-b. 1914. NPRMP, 1(3):41 (Patagonico Inferior). Type Loca/iTy.— Camarones, Chubut. Syntypes.—^3?>A, 100 valves (the one figured is a right valve, 4.5 X 4 mm), coll. Ameghino, 1900. Stratigraphy. — -Patagonian Eormation (Lower Miocene). Remarks.— See C. acutus Dali from the Pliocene of Florida. dalli, 1897. Cucullaea CUCULLAEIDAE 18971?. RMP, 2:241, pi. 7, fig. 47, and pi. 8, fig. 54 (Formagao Santacmzense). 1907a. AMNBA, 14:232. Cucullaea alta Sowerby. 1914. NPRMP, 1(3): 14. Cucullaea alta Sowerby (Patagonico). = Cucullaea alta Sowerby, 1846. Type Locality.— Jsick. Harvey, Santa Cruz. Type.— Hoi in the collection. Other Lots. — #84, one complete specimen, coll. Ameghino, 1900; #85, two valves; #86, Santa Cruz, two casts, coll. J. Bicego; #87, Camarones, two fragments attached to Ostrea, coll. Ameghino, 1900; #88, Canada de los Artilleros, one valve, coll. Ameghino, 1900; #89, Punta Casamayor, one valve, coll. Ameghino, 1900; #90, Cabo Tres Puntas, coll. Ameghino, 1905; #91, Cabo Tres Puntas, coll. Ameghino, 1900; #92, Punta Nueva, one complete specimen, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). dalli, 1897. Gibbula ARCHITECTONICIDAE 1897/?. RMP, 2:273, pi. 3, fig. 1, and pi. 4, fig. 13 (Forma9ao Santacmzense). 1902. Ortmann, p. 171, pi. 31, fig. 9a-b. 1907a. AMNBA, 14:131. 1914. NPRMP, 1(3):73 (Superpatagonico). = Valdesia dalli (Ihering). Type Locality. — -Jack Harvey, Santa Cruz. Syntypes. — -#576, four specimens, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.- — This is the large form of G. cuevensis (= dalli); see also Gibbula cuevensis and Gibbula cuevensis fracta Ihering. See Del Rio (1985) for the de- scription of the genus Valdesia. dalli, 1907. Lithodomus patagonica [sic] MYTILIDAE: LITHOPHAGINAE 1907a. AMNBA, 14:275. 1914. NPRMP, 1(3):39 (Patagonico Medio). = Lithophaga patagonica (d’Orbigny, 1846). 220 Annals of Carnegie Museum VOL. 65 Type Locality. — Rada Tilly, Golfo San Jorge. Holotype. — #320, by original designation, “exemplaire typique 32 mm,” two paratypes, coll. Ameghino, 1901, found perforating Ostrea hatcheri. Stratigraphy.-— Faiagonisin Formation (Lower Miocene). Remarks. — This species resembles Lithodomus platensis Philippi (1893) from the Parana Formation. dalli, 1907. Lyria VOLUTIDAE 1907a. AMNBA, 14:204, pL 14, fig. 9. 1914. NPRMP, 1(3): 108 (Patagonico Medio). = Enaeta dalli (Ihering), new combination. Type Locality. — Cabo Tres Puntas, Santa Cruz. Holotype. — #874 (a very small specimen), coll. Ameghino, 1905 (by mono- typy)- Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — On consultation with H. W. Dali, Ihering placed this species in Lyria. There is another lot, #875, assumed to be a different species by Ihering, but too incomplete for description. dalli, 1907. Phacoides LUCINIDAE 1907a. AMNBA, 14:284. Not figured. 1914. NPRMP, 1(3):46 (Superpatagonico). — Lucina dalli (Ihering). Type Locality. — Canada de los Artilleros, Santa Cruz. Syntypes. — #386, six valves, coll. Ameghino, 1900. Stratigraphy. — Patagonian Eormation (Lower Miocene). Remarks. — Phacoides Blainville, 1825 (= Lucina Bruguiere, 1797). dalli, 1907. Venericardia CARDITIDAE 1907a. AMNBA, 14:413, pi. 16, fig. 108a-b. 1914. NPRMP, 1(3):43 (Formagao Araucana). Type Locality. — Sierra Laziar, Santa Cruz. Syntypes. — #353, three valves, coll. Ameghino, 1900. Stratigraphy.— ArancanQan Formation (Lower Pliocene). dautzembergi, 1897. Triton (Argobuccinum) RANELLIDAE: RANELLINAE 18971?. RMP, 2:293, text fig. 16 (Formagao Santacruzense). 1907a. AMNBA, 14:176. Lotorium. 1914. NPRMP, 1(3):97. Lotorium {Lampusia) (Superpatagonico). = Fusitriton dautzembergi (Ihering), new combination. Type Locality. — Yegua Quemada, Santa Cruz. Holotype. — #791, the specimen is small, juvenile (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). densestriata, 1897. Struthiolaria ornata “var.” STRUTHIOLARIIDAE 1897^. RMP, 2:292, text fig. 15. 1907a. AMNBA, 14:171. 1914. NPRMP, 1(3):95 (Patagonico Medio-Inferior). 1996 Parodiz — Ihering Mollusca 221 = Struthioiaria ornata Sowerby (1846). Type Locality.— Not reported in the original description, but the specimens of the “van” in the '‘type lot” were labelled as being from Santa Craz. Syntypes.— 4769, seven specimens, colL Ameghino. Other Lots.— ^716, Pan de Azucar, colL Ameghino; #771, Manaetial Salado, three specimens, colL Ameghino; #772, Cabo Tres Puntas, six specimens, colL Ameghino; #773, Sierra de los Baguales, eight specimens, colL Ameghino; #774, Corral Foyel, three specimens, coll. Hauthal; #775, Golfo San Jorge, five speci- mens, coll. Ameghino. Stratigraphy.— FdLl^gommi Formation (Lower Miocene). Remarks.— Of Struthioiaria ornata Sowerby (1846:260, pL 4, fig. 62) there are also five lots in the Ihering Collection: #777, Santa Cruz, 1 1 specimens, coll. J. Bicego; #778, ten specimens, coll. Hatcher, received from Ortmann; #779, “Foz do Rio Santa Craz,” one specimen received from Ortmann; #780, La Cueva in conglomerate; #781, Canada de los Artilleros, one specimen, coll. Ameghino. From Park- Aik, Rio Sehuen there is one cast, #782, collected by C. Ameghino, and labelled as ""Struthioiaria sp.” which is probably S. ornata (see Struthioiaria prisca). derbyi, 1907. Diplodon HYRIIDAE 1907fl. AMNBA, 14:466, pL 18, fig. 128 (Cretaceous). 1914. NPRMP, 1(3):36 (Terciario). 1969. Parodiz, ACM, 40:82, pi. 8, fig. 3, Fossula derbyi (Ihering). = Fossula derbyi (Ihering). Type Locality.— Marfa da Bocca do Monte, Rio Grande do Sul, Brasil. Holotype.— 4297 , upper portion of a valve, coll. Orville A. Derby (by mono- typy)- deseademse^ 1914. Calliostoma 7TURBINIDAE 1914. NPRMP, 1(3):77, pL 3, fig. 12 (Foraia^ao Araucana). Type Locality.— Deseado, Santa Cruz. Holotype.—4603, coll. Ameghino, 1900 (by monotypy). Stratigraphy.— Armcmman Formation (Pliocene) Remarks.— This species is not a Calliostoma, but rather more like Turbo. The holotype is a broken specimen but in the figure it shows complete ornamentation. deseadensiSf 1907. Myochlamys PECTINIDAE 1907a. AMNBA, 14:410, pL 15, fig. 102a-^b. 1914. NPRMP, 1(3):30 (Formayao Araucana). = Chlamys deseadensis (Ihering). Type Locality.— Laziar, Syntypes.— #242, 12 valves. Stratigraphy.— (Taziaiense) Araucaneae Formation (Lower Pliocene). deuterolivella^ 1907. Volvaria MARGINELLIDAE 1907a. AMNBA, 14:197. New name for Marginella oliviformis in Ortmann (1900:376), not Margi- nella oliviformis Tuomey and Holmes, 1857. 1914. NPRMP, 1(3): 106. Marginella {Volvaria). deuteroliveila (Patagonico), 222 Annals of Carnegie Museum VOL, 65 = IMarginella {sensu lato) deuterolivella (Ihering). Type Loua/ily.— Mouth of Santa Cmz River (Ortmann). Stratigraphy.— Vdiidigomdin Formation (Lower Miocene). Specimens of this species were misidentified as Marginella olivella Reeve, 1865, by Ortmann (1902:225, pL 35, fig. 3a, b). This species is more elongated than any living species of Marginella from the southwestern Atlantic, and is Columbella~\ik.e, in shape. There is one specimen (#863, Golfo San Jorge, colL Ameghino) in the Ihering Collection. duMosa, 1907. Gibbula ARCHITECTONICIDAE 1907a. AMNBA, 14:130, pL 4, fig. 91a-b. 1914. NPRMP, 1(3):73 (Patagonico Medio). = Valdesia dalli (Ihering). Type Locality. — Golfo San Jorge. Holotype. — #577, (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species was based on a distorted specimen of ""Gibbula'' dalli dynmtes, 1907. Nucula NUCULIDAE 1907a. AMNBA, 14:3, pL 1, fig. 2a-b (Rocaneen). 1914. NPRMP, 1(3): 12 (Roca, Cretaceo Superior, Roca). Type Locality. — General Roca, Rfo Negro. Syntypes. — #64, two casts, received from Ameghino in 1905. Stratigraphy. — Roca Formation (Paleocene). elongatUy 1914. Panopaea [sic] nucleus HIATELLIDAE 1914. NPRMP, 1(3):64. Not figured (Cretaceo Superior, Roca). = Panopea nucleus (Ihering). Type Locality.— Coitdil FoyeL Holotype.--^5\l , one cast, coll. Hauthal (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.— Th&tQ is another specimen in the collection (MACN #2480), Foyel River. Ihering’s brief diagnosis indicates a form higher than nucleus with the margin slightly convex (Ihering: , me parece uma variedade”). Entacanthus, 1907. MURICIDAE 1907a. AMNBA, 14:14:183. New subgenus of Trophon Montfort. = Thais (Stramonita) Schumacher, 1817. Type Species. — Trophon monoceros Ihering. Stratigraphy.— PdilAgonmn Formation (Lower Miocene). entreriana, 1907. Anomalocardia VENERIDAE 1907a. AMNBA, 14:386. 1914. NPRMP, 1(3):57 (Forma^ao Eotreriana). Type Locality. — Parana, Entre Rios. Syntypes.— ^476, five valves, Bravard coll. Remarks. — The syntypes were identified by Borchert (1901:39, pL 3, fig. 12- 1996 Parodiz — Ihering Mollusca 223 21) as Venus brasiliana (Gmelin). The specimens were sent to W. H. Dali who confirmed it to be a different species but still very similar to Anomalocardia brasiliana (Gmelin). entreriana, 1907. Dosinia VENERIDAE 1901. Borchert, p. 36. Dosinia patagonica, not Dosinia patagonica Philippi, 1893. 1907a. AMNBA, 14:383, Not figured. = Dosinia meridionalis Ihering. Type Locfif/fry.— -Parana, Entre Rios (Borchert). Type. — Not in the collection. Stratigraphy. — Parana Formation (Upper Miocene); also in the Patagonian For- mation. Remarks.— -Ihmng (1907a): “J’ai examine la coquille decrite par Borchert et verifie quelle n’est pas identique avec Dosinia patagonica."" entreriana, 1907. Leda NUCULANIDAE 1901. Borchert, p. 32, Leda patagonica, not Leda patagonica d’Orbigny, 1846. 1907a. AMNBA, 14:371. Not figured. = Nuculana entreriana (Ihering). Type Locality. — Parana, Entre Rios (Borchert). Type. — Not in the collection. Stratigraphy. — Parana Formation (Upper Miocene). Remarks.- — Other ""Leda'' in the collection are only from Patagonia, not from Parana. entrerianuSy 1907. Tagelus gibbus PSAMMOBIIDAE: SOLECURTINAE 1901. Borchert, p. 21, pi. 3, fig. 26. Solecurtus platensis d’Orbigny. 1907a.— AMNBA, 14:387. Not figured. = Tagelus plebeius (Solander, 1823). Recent. Type Locality. — ^Parana, Entre Rios (Borchert). Stratigraphy. — Parana Formation (Upper Miocene). Remarks.- — Solecurtus platensis d’Orbigny, Tagelus gibbus of authors, and T. plebeius (Solander) are all synonyms for the same species which is widely dis^ tributed in the western Atlantic. eopatagonica, 1907. Turbonilla PYRAMIDELLIDAE: TURBONILLINAE 1907a. AMNBA, 14:144. New name for Turbonilla iheringi in Cossmann (1899a:235, pi. 10, fig. 12), not T. iheringi (Clessin, 1902). 1914. NPRMP, 1(3):82 (Superpatagonico). = Turbonilla iheringi Cossmann, 1899. Type Locality.— YQgxxdL Quemada, Santa Cruz. Stratigraphy. — Patagonia Formation (Lower Miocene). Remarks. — This name is an unnecessary replacement name for T. iheringi Cossmann. The only specimen in the collection is #636, collected by Ameghino in 1900, and therefore not the type of Cossmann’s ""T. iheringi" described in 1899. The four species of Turbonilla named by Ihering, T. clessini, T. cuevensis, T. observationis, and T. eopatagonica (= T. iheringi Cossmann) are all similar 224 Annals of Carnegie Museum VOL. 65 to the Recent Turbonilla {Pyrgiscus) uruguayensis Pilsbry. See Remarks section under T. clessinL espejoana “Ihering,” 1902. Chlamys PECTINIDAE 1902. Ameghino, ASCA, 54:224. Nomen nudum. = Pecten praenuncius Ihering, 1897k eupatagonica, 1907. Myochlamys PECTINIDAE 1907a. AMNBA, 14:259, pi. 9, fig. 58. 1914. NPRMP, 1(3):30 (Superpatagonico). = Chlamys eupatagonicus (Ihering). Type Locality. — Canada de los Artilleros, Santa Cruz. Holotype. — #243, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). eupyga, 1907. Chione VENERIDAE 1907a. AMNBA, 14:310, pi. 12, fig. 80. 1914. NPRMP, 1(3):55 (Patagonico Superior). Type Locality. — San Julian, Santa Cruz. Holotype. — #452, coll. Ameghino, 1900 (by monotypy). Other Lot. — #453, Rada Tilly, Santa Cruz, coll. S. Roth, reported from “Roca.” Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — The specimen described and figured by Ihering is a fragment of a valve in poor condition for correct identification; it may belong in Chione rneri- dionalis Sowerby. faira^ 1907. Ostrea OSTREIDAE 1907a. AMNBA, 14:406. 1914. NPRMP, 1(3):25 (Formagao Araucana). Type Locality. — Cabo Fairweather (Buentiempo) Santa Cruz. Syntypes.—^\96, two large specimens received from Ortmann. Stratigraphy. — Araucanean Formation (Pliocene). Remarks. — Ostrea ingens in Ortmann (1902:99, pi. 18, fig. la-b) is in part this species. fairuSy 1907. Trophon inornatus MURICIDAE 1907a. AMNBA, 14:402. 1914. NPRMP, 1(3): 100 (Formagao Araucana). = Trophon {Austrotrophon) lamellosus (Gmelin, 1791). Type Locality. — Cabo Fairweather (Buentiempo), Santa Cruz. Syntypes. — #810, two specimens, coll. Hatcher, received from Ortmann. Stratigraphy.- — Araucanean Formation (Pliocene). felschiy 1921. Aturia (Sphenaturia) NAUTILIDAE 1921. Physis, Buenos Aires, 5:76. Type Locality. — Punta Chocoi, Carelmapu, Chile. 1996 Parodiz — Ihering Mollusca 225 Holotype. — Collected by J. Flesch; status unknown. Stratigraphy. — “Estratos de Boqueron” (below Patagonian Formation). Remarks. — Referring to the holotypes of this species and A. {S.) brueggeni, Ihering (1921) stated, “Los ejemplares tipicos quedan depositados en este Museo [Museo Argentino de Ciencias Naturales, Buenos Aires].” fiUpponei [sic], 1907. Lotorium RANELLIDAE: CYMATIINAE 1907a. AMNBA, 14:443, pL 17 (text states in error “XVIII”) fig. 22a-b. = Cymatium (Cabestana) felipponei (Ihering), emendation. Type Locality. —Maldonaido, Uruguay. Type. — Not in the collection. Stratigraphy. — Querandi Formation (Pleistocene). Remarks. — Ihering mentioned another specimen (1907<7) from Punta Carretas, Montevideo, also received from E Felippone of Uruguay. Fimbria sp. LUCINIDAE 18971?. RMP, 2:219. Specimens. — ^Not in the collection. Remarks.- — A single valve from Yegua Quemada, Santa Cruz, which Ihering said was CorbisAdko^, but different from ''Fimbria^" patagonica (Philippi). Corbis patagonica Philippi is of questionable taxonomic position. Fimbria M. von Muhl- feld, 1811; not Bohadsch, 1761 (= Corbis Cuvier, 1817). fissocostalis, 1899. Pecten PECTINIDAE 1899a. NJMGP, 2:11, pi. 1, fig. 1. 1907a. AMNBA, 14:255. Myochlamys. 1914. NPRMP, 1(3):31 (Patagonico). = Chlamys (Zygochlamys) fissocostalis (Ihering); see Morra (1985). Type Locality. — Santa Cruz. Syntypes.—^2AA, two complete specimens and four valves, coll. J. Bicego, 1897. Other Lot. — #245, fragment of a large valve, coll. Bicego; probably same lo- cality. Stratigraphy.-— Palagonmn Eormation (Lower Miocene). foyela, 1914. Myochlamys PECTINIDAE 1914. NPRMP, 1(3):33, pL 1, fig. 3 (Patagonico). = Chlamys foyela (Ihering). Type Locality.— Corral Foyel, Neuquen. Syntypes. — #273, three valves and pieces in conglomerate, coll. Hauthal. Stratigraphy.— Patagonian Formation (Lower Miocene). fracta, 1897. Gibbula ARCHITECTONICIDAE 1897Z?. RMP, 2:273, pi. 3, fig. 2 (Forma9ao Santacruzense). 1907a. AMNBA, 14:131, “et Van cruzis"’ not described. 1914. NPRMP, 1(3):73 (Superpatagonico); the specimen figured in 1897 is identified here as cuevensis. = IValdesia fracta (Ihering). 226 Annals of Carnegie Museum VOL. 65 Type Locality. — Yegua Quemada, Santa Cruz. Holotype. — #578, one specimen in fragments (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Another specimen from the same locality is reported as Gibbula diametralis Cossmann. The differentiation of ‘‘G.” fracta cuevensis is very dif- ficult; all specimens from the same area of Golfo San Jorge are probably indi- vidual variations. fraus, 1907. Diplodon HYRIIDAE 1907a. AMNBA, 14:467, pi. 18, fig. 129. Unnecessary replacement name for Modio la contorta Borch- ert, 1901:27, pi. 2, fig. 3-4, not Unio contortus Heude. 1969. Parodiz, ACM, 40:74, pL 7, fig. 5. = Diplodon contortus (Borchert, 1901), new combination. Type Locality. — Parana, Entre Rios (Borchert), Holotype. — Examined by Parodiz in 1971 in the Senckenberg Museum, Erank- furt. Stratigraphy. — Parana Eormation (Upper Miocene). Remarks. — The name contorta is not preoccupied in Diplodon', therefore, Di- plodon fraus Ihering, based on the type of Modiola contorta Borchert, is an unnecessary replacement name in that genus. This species belongs to the Diplodon parallelipipedon group. freti, 1907. Venericardia sulcolunularis CARDITIDAE 1907a. AMNBA, 14:347. Not figured. Type Locality. — Carmen Sylva, Tierra del Fuego. Type. — Not in the collection. Stratigraphy. — Magellanian Formation (Lower Oligocene). fuegintty 1907. Struthiolaria chilensis STRUTHIOLARIIDAE 1907a. AMNBA, 14:343. Not figured. = Struthiolaria chilensis Philippi, 1887. Type Locality. — Carmen Sylva, Tierra del Fuego. Stratigraphy. — Magellanian Formation (Lower Oligocene). Remarks. — There are no specimens in the collection under the name fuegina, but there is one specimen of S. chilensis, labelled by Ihering as “cotypo,” from Matanzas, Tertiary of Chile, received from R. A. Philippi. Struthiolaria chilensis is related to S. nordenskjoldi Wilckens, 1911, from the Tertiary of Seymour Island, Antarctica. fueguensisy 1907. Cominella BUCCINIDAE: PHOTINAE 1907a. AMNBA, 14:343, pi. 14, fig. 97a-b. 1908. Steinmann and Wilckens, 4(6):60, pi. 7, fig. 3-5. Cominella obesa "'var.’’' Jueguina. 1914. NPRMP, 1(3): 105 (Superpatagonico). = Cominella (Austrocominella) fueguensis Ihering. Type Locality. — Barrancas de Carmen Sylva, Tierra del Fuego. Syntypes. — #858, two specimens, coll. Backhausen. Stratigraphy. — Patagonian Formation (Lower Miocene). 1996 Parodiz — Ihering Mollusca 227 Remarks,— This species is the type of the subgenus Austrocominella Ihering, 1907. It was also found by Femglio at Rio Turbio. gibber, 1907. Tellina TELLINIDAE 1907a. AMNBA, 14:456, pL 18, fig. i26a-b. 1914. NPRMP, 1(3):59 (Pampeano and post-Pampeano). Type Locality,— Pimilo Militar (Puerto Belgrano), Buenos Aires Province. four valves from what Ihering indicates as Pampeano, and one valve posPPampeano, coll. Ameghino. Stratigraphy.— Bclgmiio Formation (Pleistocene) and Recent. Remarks.- — The Recent distribution of this species is from Rio de Janeiro to Chubut. golfina, 1907. Gibbula iheringi “van” BUCCINIDAE: PHOTINAE 1907a. AMNBA, 14:133. Not figured. 1914. NPRMP, 1(3):74 (Patagonico). = Gibbula iheringi Cossmann, 1899. Type Locality.— Cdho Tres Puntas, Santa Cruz. Holotype.- — #582, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — -Patagonian Formation (Lower Miocene). Ihering commented “Un des exempiaires que je possede” probably referring to G. iheringi Cossmann from Yegua Quemada, of which the “var.” goifina is a synonym. golfonia, 1914. Comineila BUCCINIDAE: PHOTINAE 1914, NPRMP, 1(3): 105, pi. 3, fig. 5 (Patagooico Medio). = Comineila {Austrocominella) golfonia Ihering. Type Locality.— Cdiho Tres Puntas, Santa Cruz. Syntypes. — #860, seven specimens, coll. Ameghino. Stratigraphy.— TAi^Lgommi Formation (Lower Miocene). Remarks.— Comineila iheringi Feruglio, 1935, from the Salamanca Formation has affinities with this species. gracilior, 1897. Marginella MARGINELLIDAE 1897fe. RMP, 2:308, text fig. 18 (Forma^ao Santacmzense). 1907a. AMNBA, 14:200. Neoimbricaria. 1914. NPRMP, 1(3): 107 (Superpatagoeico). = Neoimbricaria quemadensis (Ihering). Type Locality.— Quemada, Santa Cruz. Holotype.— ^^61 , coll. Ameghino, 1900 (by monotypy). Patagonian Formation (Lower Miocene). Remarks.—See Marginella confinis Ihering and Marginella quemadensis Iher^ ing. gracilior, 1896. Voluta VOLUTIDAE 1896. NDMG, 7:96. New name for Voluta gracilis in Philippi (1887), not Voluta gracilis Swainson, 1842. 18971?. RMP, 2:305. Voluta pMlippianus Dali. 228 Annals of Carnegie Museum VOL. 65 1907a, AMNBA, 14:206. Proscaphella gracilior (Patagonico). 1914. NPRMP, 1(3): 110. Cymbiola (Miomelon) gracilior. = Miomelon gracilior (Ihering). Type Locality. —Sa.nta. Craz. Syntypes.-—4t^^9, Santa Cmz, one specimen and one cast, coll. J. Bicego, 1897. Stratigraphy.— Patagonim Formation (Lower Miocene). gracilis f 1897. Cancellaria CANCELLARIIDAE 1897L RMP, 2:310, pi. 3, fig. 11 (Forma^ao Santacrazense). 1907a. AMNBA, 14:213. 1914. NPRMP, 1(3):111 (Superpatagonico). Type Locality.— La Cueva, Santa Craz. Holotype.—4tH96, coll. Bicego, 1897 (by moeotypy). Other Lot— ^^91 , Canada de los Artilleros, two specimens, colL Ameghino. Stratigraphy.— Palagonian Formation (Lower Miocene). Remarks.— Cancellaria gracilis observatoria Ihering is a synonym. gradatUy 1907. Trophon inornatus “var.” MURICIDAE 1907a. AMNBA, 14:402. Nomen nudum. Remarks.— Th^ name gradata described on the same page was given to one of the two specimens of T. inornatus fairus. See also Trophon varians gradata Iher- ing, 1897. gradata, 1897. Trophon varians “var.” MURICIDAE 1897 A RMP, 2:297, 323 (Pan-Patagonico). 1907a. AMNBA, 14:403. 1914. NPRMP, 1(3): 101. T. varians “gradatus” (Formagao Araucana). = Trophon varians (d’Orbigny, 1841). Type Locality.— "'LnliQ S. Jorge e Deseado,” Santa Craz. Holotype. — #833, coll. Ameghino (by monotypy). Other Lots (as Trophon varians). -=#829, San Juan [sic] must be San Julian, four specimens, coll. Ameghino; #830, Sierra Laziar, ten specimens, coll. Amegh- ino; #831, Monte Espejo, five specimens, coll. Ameghino; #832, Golfo San Jorge, “Pampeano,” five specimens, coll. Ameghino. S?rarigra|?/iy.=Araucanean Formation (Pliocene) and Belgrano Formation (Pleistocene); Recent, common. guaranitica, 1899. Ostrea OSTREIDAE 18991?. ASCA, 47:63-64 (Piso Sehueese, Formacion Guaranitica), 1902a. AMNBA, 7:116. 1903. AMNBA, 9:213. 1907a. AMNBA, 14:50 (Sehuenfen). 1914. NPRMP, 1(3):2 (Guaraniee). = Exogyra guaranitica (Ihering). Type Locality.— Park- AiL (Sehuen-Aik) Sehuen Valley, Santa Craz. Syntypes.—OnG lot of nine specimens is in the collection with one specimen, #149, marked as “type.” Stratigraphy.— SQfm&nan Formation (Paleocene). 1996 Parodiz — Ihering Mollusca 229 Remarks.— An incomplete specimen in the Ihering Collection, #150 from San Juan Creek, Santa Ana, Misiones, was identified as guaranitica but probably does not belong in the same Patagonian species. Ihering (1907a:52) stated that “Les huitres de San Juan. . .ne peuvent non plus servir pour reconnaitre Page geolo= gique, de depots respectifs [Upper Cretaceous or Tertiary].” The name ''guar- anitica” was taken from an ethnic root of the north, and is confusing for a Pat- agonian species. See Stratigraphic Units. guassus [sic], 1907. Psammobia PSAMMOBIIDAE 1907a. AMNBA, 14:312, pi. 14, fig. 95 (Patagonico). 1914. NPRMP, 1(3):57. Psammobia guassu (Patagonico Inferior). = Gari guassu Ihering (guassus is a typographical error). Type Locality.— Bajo de San Julian, Santa Cruz. Holotype.—#Al%, one large cast, coll. Ameghino (by monotypy). Stratigraphy.- — Patagonian Formation (Lower Miocene). Remarks.— lhQ,nn% (1914) explained that the word “guassu” in Tupi language means large and can be used, Latinized, as an adjective; but Tupi are Indians of southern Brazil, and why the word was used for a Patagonian species is difficult to understand. hauthali, 1907. Mytilus chorus “var.” MYTILIDAE imib. RMP, 2:232, pL 9, fig. 55. Mytilus aff. chorus. 1902. Ortmann, p. 120, pi. 25, fig. la-~b. 1907a. AMNBA, 14:273. Mytilus chorus “var.” hauthali. = Chloromya hauthali (Ihering). Type Locality.- — Mayten, Rio Chubut. Syntypes. — -#304, three specimens in a conglomerate, coll. Hauthal. Stratigraphy.— BdLi2igonmn Formation (Lower Miocene). Remarks. — This species has affinities with Mytilus pseudochorus Doello-Jura- do, 1922. hauthali, 1904. Strophocheilus MEGALOBULIMIDAE 1904. RMLP, 11:14, pi. 2, fig. 12. 1907a. AMNBA, 14:463 (Notostylopeen). 1914. NPRMP, 1(3):70 (Cretaceo Superior). 19496. Parodiz, Physis, 20:183. Megalobulimus hauthali (Ihering). 1969. Parodiz, ACM, 40:175 = Megalobulimus hauthali (Ihering). Type Locality.— CdindAon Blanco, between the rivers Senguer and Chubut. Type Material.- — Lectotype, #557, coll. Ameghino (not Hauthal); three paralec- totypes, same data as lectotype. Stratigraphy.— Cn^^mdiy or Formation (Lower Eocene); also Musters Formation (for Colhue Huapi, Astraponotense of authors; Upper Eocene). Remarks.— Tho lectotype was subsequently designated by Parodiz (1969:176). As in the case of Strophocheilus chubutensis, the collector of the type lot was C. Ameghino, not Hauthal as reported by Ihering (1914); the original label is in the handwriting of C. Ameghino, and indicates “capas de Astraponotus,'' not Notos- tylops. Strophocheilus avus Parodiz (1949^) was synonymized with S. (Megalob- ulimus) hauthali (Ihering) by Parodiz (1969). 230 Annals of Carnegie Museum VOL. 65 hauthali, 1907. Turritella TURRITELLIDAE 1907a. AMNBA, 14:163, pi. 5, fig. 22. 1914. NPRMP, 1(3):90 (Patagonico Inferior). Type Loca/ity.— -Camarones, Chubut. Syntypes. — #731, six specimens, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species has affinities with Turritella malaspina Ihering of the Salamanca Formation. huttoniy 1907. Cardium CARDIIDAE 1907a. AMNBA, 14:291. Holotype. — A single example sent to Ihering by F. W. Hutton; not in the col- lection. Type Loc<3//fy. —Waipara, New Zealand. Stratigraphy. — Miocene. Remarks. — Briefly diagnosed in the discussion of Cardium philippi Ihering. Ihering {\901a) proposed the new name C. huttoni for a specimen from the Mio- cene of Waipara, New Zealand, sent by F. W. Hutton and misidentified as Cardium multiradiatum Sowerby, 1833 (not Philippi, 1887). The type may have been re- turned to Hutton, but this is not certain. Also see Cardium philippii Ihering. improvisa, 1914. Columbella (Anachis) COLUMBELLIDAE 1914. NPRMP, 1(3): 106. Not figured (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Syntypes. — #861, seven specimens, coll. Ameghino. Stratigraphy. — ^Patagonian Formation (Lower Miocene). indecussata, 1899. Turritella breantiana “van” TURRITELLIDAE 1897L RMP, 2:288. Turritella breantiana d’Orbigny. 1899a. NJMGP, 2:26. Turritella b. “van” indecussata. 1907a. AMNBA, 14:164, pi. 5, fig. 23, T. breantiana. 1914. NPRMP, 1(3):89 refers to fig. 23 of 1907a as T. indecussata. = Turritella breantiana d’Orbigny, 1847. Type Locality. — Santa Cruz. Holotype. — #4229, coll. J. Bicego, 1895 (by monotypy). Other Lots. — #721, Manantial Salado, one specimen, coll. Ameghino; #722, Bajo de las Flechas, Santa Cruz, one specimen, coll. Ameghino; #723, Punta Casamayor, one specimen, coll. Ameghino; #724, Golfo San Jorge, one specimen, coll. Ameghino; #725, Cabo Tres Puntas, two specimens, coll. Ameghino; #726, Canada de los Artilleros, four specimens, coll. Ameghino. Stratigraphy. — -Patagonian Formation (Lower Miocene). Remarks. — There are no lots labelled as T. breantiana in the collection to compare with “van” indecussata. Turritella iheringi Cossmann, 1898, and T. tricincta Ihering, 1897 (not T. tricincta Hutton, 1873), are synonyms of T. brean- tiana d’Orbigny, 1847. indistinctUy 1897. Mactra 1897A RMP, 2:262, text fig. 3 (Formagao Santacruzense). 1907a. AMNBA, 14:320. MACTRIDAE 1996 Parodiz — Ihering Mollusca 231 1914. NPRMP, 1(3):60 (Superpatagonico). = Mactra patagonica d’Orbigny, 1846. Type Locality. — Yegua Quemada, Santa Cruz. Lectotype. — #497, the only specimen that remains in the collection from the three valves reported by Ihering (1914), coll. Ameghino, 1905, here designated. Stratigraphy. — Patagonian Formation (Lower Miocene). jeguaensis, 1897. Tellina TELLINIDAE 1897^. RMP, 2:260, pi. 5, fig. 33 (Forma^ao Santacmzense). 1907«. AMNBA, 14:315. ?“var.” of '"striata'' Turton. 1914. NPRMP, 1(3):58 (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Syntypes. — #484, three specimens, coll. Ameghino, 1895. Other Lor.— #485, Canada de los Artilleros, one valve, coll. Ameghino. S'rrar/grap/iy.— Patagonian Formation (Lower Miocene). jorgea, 1914. Mactra MACTRIDAE 1914. NPRMP, 1(3):60, pi. 2, fig. 6a-b (Patagonico Inferior). Type Locality. — Southeast of Punta Nova (Nueva), Santa Cruz. Syntypes. — #496, two specimens, crystalized into chalcedony, coll. Ameghino, 1905. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — ^The hinge area of the specimens is missing, for which reason Iher- ing stated “duvidosa a sua posigao systematica.” The valves are shorter but with the apex higher than in Mactra patagonica d’Orbigny. jorgensis, 1907. Bulla patagonica “van” 7RETUSIDAE 1907a. AMNBA, 14:121. Not figured. 1914. NPRMP, 1(3):69 (Supra e Patagonico Medio). = IRetusa patagonica (Ihering), new combination. Locality. — Punta Casamayor, Santa Cruz. Holotype. — #554, coll. Ameghino, 1900 (by monotypy). Other Lot. — #553, one specimen, Golfo San Jorge, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). jorgensis^ 1907. Cardium pisum “van” CARDIIDAE 1907a. AMNBA, 14:294. Not figured. 1914. NPRMP, 1(3):48 (Patagonico Medio). = Cardium jorgensis Ihering, 1907, new status. Type Locality. — Cabo Tres Puntas, Santa Cruz. Holotype.- — #405, colL Ameghino, 1905 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). jorgensis, 1907. Hadriania MURICIDAE: OCENEBRINAE 1907a. AMNBA, 14:185, pi. 6, fig. 32a-b. 1914. NPRMP, 1(3):99 (Patagonico Medio a Inferior). = Crassilabrum jorgensis (Ihering), new combination. 232 Annals of Carnegie Museum VOL. 65 Type Locality.— ¥unidL Casamayor, Santa Craz. Holotype.- — #804, coll. Ameghieo, 1900 (by moeotypy). Other Lot.— two specimens, Rio Chico, Chubut, coll. Ameghino, 1900. Stratigraphy.— FaiRgonian Formation (Lower Miocene). Remarks.— The valid generic name is Crassilabrum Jousseaume, 1880. The replacement name Antimurex Cossmane, 1903, on grounds of preoccupation, is not valid because the presumed senior homonym Crassilabrum Megerle in Scud- der (1882a: 88 and 1882l?:81) is a nomen nudum. jorgensis, 1907. Pecten PECTINIDAE 1907a. AMNBA, 14:258, pi. 9, fig. 57a, d. 1914. NPRMP, 1(3):34. Myochlamys (Patagonico Medio). = Chlamys (Zygochlamys) jorgensis (Ihering); see Morra (1985). Type Locality.— Punta Casamayor, Santa Cmz, Syntypes. ■—^274, one complete specimen plus one valve, coll. Ameghino. Other Lots.— ^215, same locality, one valve, coll. Ameghino, 1900; #276, Pun- ta Nodales, three valves, coll. Ameghino, 1900; #277, Punta Nova (Nueva), one intact specimen and six other valves, coll. Ameghino, 1900; #278, Fondo del Bajo de San Julian, Santa Cruz, one valve, coll. Ameghino, 1900; #279, Rio Chico, Chubut, one valve, coll. Ameghino, 1900. Patagonian Formation (Lower Miocene). Remarks.— Found also at Cerro Otto by E. Femglio. jorgensis f 1907. Pleurotoma TURRIDAE 1907a. AMNBA, 14:220, pL 7, fig. 47. 1914. NPRMP, 1(3):113 (Patagonico Inferior). Type Locality.— Caho Tres Puntas, Santa Cmz. Holotype.— 4t906, incomplete specimen, coll. Ameghino, 1900 (by moeotypy). Stratigraphy.— Fatagonian Formation (Lower Miocene). Remarks.— A more elongated species than Pleurotoma santacruzensis Ihering, otherwise very similar (both species based on a single specimen). jorgensisy 1907. Polynices NATICIDAE 1907a. AMNBA, 14:158, pL 5„ fig. 19. 1914. NPRMP, 1(3):84 (Patagonico Medio). = Polynices famuia (Philippi). Type Locality.— Cabo Tres -Puntas, Santa Cmz. Holotype.— #611, coll. Ameghino, 1900 (by moeotypy). Strarigrap/iy.— Patagonian Formation (Lower Miocene).. Ihering (1907a: 158) states, “Cette espece parait voisine de la P. famuia Phil., mais . . , foverture plus grand. ...” In the unique specimen the aperture is broken, but all other visible characters are equal to famuia, Juliana^ 1907. Area ARCIDAE 1907fl. AMNBA, 14:238, pL 7, fig. 5a--b. 1914. NPRMP, 1(3):18 (Patagonico Inferior). = Anadara Juliana (Ihering), new combination. Type Locality.— Fan de Azucar, Santa Cmz (Forma^ao Patagonica Inferior). 1996 Parodiz — Ihering Mollusca 233 Holotype.—^\\5, single valve, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Juliana, 1907. Bullinella SCAPHANDRIDAE [CYLICHNIDAE] 1907a. AMNBA, 14:121, pi. 4, fig. 6. 1914. NPRMP, 1(3):69 (Superpatagonico). = Cylichna Juliana (Ihering). Type Locality. — Canada de los Artilleros, San Julian, Santa Cruz. Syntypes. — -#555, nine specimens, coll. Ameghino, 1900. Stratigraphy.— Vdit^gonmn Eormation (Lower Miocene). Remarks.— Bullinella Newton, 1891 (= Cylichna Loven, 1846). The peristoma in this species is very thick and marginated. Juliana, 1907. Calyptraea pileus “var.” CREPIDULIDAE 1907a. AMNBA, 14:148. Not figured. 1914. NPRMP, 1(3):83 (Patagonico Inferior). = Calyptraea pileus Lamarck, 1822. Type Locality. — Fondo del Bajo de San Julian, Santa Cruz. Holo type. —#654, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — ^Patagonian Formation (Lower Miocene). Remarks. — The description of the ‘"van” was not comparative and insufficient for recognition of a distinct form, but the type specimen corresponds to Calyp- traea pileus Lamarck ranging from the middle Tertiary to Recent. Do not confuse with Calyptraea pileolus d’Orbigny (= Trochita americana Gray). Juliana, 1907. Cerithiopsis CERITHIOPSIDAE 1907a. AMNBA, 14:169, text fig. 7. 1914. NPRMP, 1(3):92 (Superpatagonico). Type Locality. — Canada de los Artilleros, Santa Cruz. Holotype. — #750, coll. Ameghino, 1900 (by monotypy). Stratigraphy.— VdiidigoniMi Formation (Lower Miocene). Remarks.— In its large number of whorls and deep sutures it resembles the Recent Cerithiopsis exilis (C. B. Adams, 1850) which ranges from the Antilles to Brazil, but the whorls are less convex. Juliana, 1907. Corbula pulchella CORBULIDAE 1907a. AMNBA, 14:322, pi. 12, fig. Sla-c. 1914. NPRMP, 1(3):62. Corbula pulchella hatcheri Ortmann (Superpatagonico, Patagonico Inferior). = Corbula hatcheri Ortmann, 1900. Type Locality.- — Punta Nova (Nueva) Santa Cruz. Syntypes.— #504, four specimens, coll. Ameghino, 1900. Other Lots.— #503, Mouth of Santa Cruz River, one specimen of C. hatcheri received from Ortmann; #505, south of Colhue Huapi, one specimen plus one valve, coll. Ameghino, 1900; #506, Golfo San Jorge, five specimens, coll. Amegh- ino, 1900; #507, Camarones, 60 specimens, coll. Ameghino, 1900; #508, Canada de los Artilleros, 15 specimens, coll. Ameghino, 1900. Stratigraphy.— ^nlngoninn Formation (Lower Miocene). 234 Annals of Carnegie Museum VOL. 65 Remarks. — This species has similarities with the living Corbula lyoni Pilsbry; Recent, Buenos Aires and Uruguay. Juliana^ 1907. Dentalium sulcosum “van” DENTALIIDAE 1907a. AMNBA, 14:224. 1914. NPRMP, 1(3):67. Dentalium sulcosum julianum (emend.) (Patagonico Superior). Type Locality. — Canada de los Artilleros, Santa Cruz. Syntypes. — #541, ten specimens, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — See Dentalium sulcosum camaronesis. Juliana^ 1907. Myochlamys PECTINIDAE 1902. Ameghino, ASCA, 54:224 (in litteris as Chlamys). 1907a. AMNBA, 14:253, pi. 8, fig. 56a-b. Myochlamys. 1914. NPRMP, 1(3):34 (Patagonico Inferior, Medio). = Chlamys Juliana (Ihering). Type Locality. — Southeast of Punta Nova (Nueva), Santa Cruz. Holotype. — #280, coll. Ameghino, 1900 (by monotypy). Other Lots. — -#281, Pan de Azucar, one valve, coll. Ameghino, 1900; #282, Punta Casamayor, one valve, coll. Ameghino, 1900; #283, north of Rio Seco and San Julian, Santa Cruz, one valve, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Juliana, 1907. Neoimbricaria quemadensis “van” MARGINELLIDAE 1907a. AMNBA, 14:199. Not figured. 1914. NPRMP, 1(3): 108 (Superpatagonico). = Neoimbricaria quemadensis (Ihering). Type Locality. — San Julian, Santa Cruz. Syntypes. — #871, 15 specimens, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This form was diagnosed within the discussion of N. quemadensis as having an obsolete spiral sulcus, but this is a character extremely variable in quemadensis and other species. A synonym of this species is N. gracilior (Iher^ ing). Juliana, 1907. Urosalpinx elegans “van” MURICIDAE: OCENEBRINAE 1907a. AMNBA, 14:187, pi. 6, fig. 34a-b. 1914. NPRMP, 1(3):98. Urosalpinx elegans Ortmann (Superpatagonico). = Ocenebra (Ocinebrina) elegans (Ortmann), new combination. Type Locality. — Canada de los Artilleros, Santa Cruz. Syntypes. — #799, ten specimens, coll. Ameghino, 1900. Other Lot. — #798, Yegua Quemada, Santa Cruz, two specimens, C. Ameghino. Stratigraphy.— ^dLidigonmn Formation (Lower Miocene). Remarks. — The species U. elegans Ortmann does not belong in Urosalpinx but rather with Ocinebrina Jousseaume, 1880. What Ihering describes as “van” Ju- liana are juveniles and not distinct from elegans. 1996 Parodiz- — Ihering Mollusca 235 Julianum, 1907. Pitar VENERIDAE 1907£i. AMNBA, 14:302, pL 11, fig. 74a-b. 1914. NPRMP, 1(3):51 (Patagonico Inferior). Type Locality.— Bajo de San Julian (in the original “Baie de San Julian”) Santa Cruz. Syntypes.- — #424, three fragments of valves, coil. Ameghino, 1900. Stmtigmphy.—PatSLgonmn Formation (Lower Miocene). Remarks.- — The incomplete valves do not show well the configuration of the shells, but from the umbo and cardinal area they are clearly a species of Pitar. E. Feruglio (1949-1950) mentions the species for Rio Turbo. julkiFiMS, 1907. Vermetus ANNELIDA: POLYCHAETA 1907a. AMNBA, 14:166, pL 5, fig. 6. 1914. NPRMP, 1(3):92 (Patagonico Inferior). Type Locality.— yimimlml Salado, San Julian. Syntypes.—WJ 45, a conglomerate with several individuals. Other Lots.— ^146, Pan de Azucar, Santa Cruz, numerous in rock, coll. Amegh- ino, 1900; #747, Canada de los Artilleros, three specimens, coll. Ameghino, 1900. Stratigraphy.— Pdil^gomMi Formation (Lower Miocene). -These are not molluscs but probably tubes of polychaetes (Anne- lida). juliensiSy 1907. Pododesmus ANOMIIDAE 1907fl. AMNBA, 14:267, pL 10, fig. 65a^b. 1914. NPRMP, 1(3):27 (Patagonico Inferior). Type Locality.— Pmi de Azucar, Santa Cruz. Syntypes.—42ll , Manantial Salado, six valves, coll. Ameghino, 1900; #219, Cabo Tres Puntas, Santa Cruz, one valve, coll. Ameghino, 1905. Stratigraphy.— Pdiidigomwi Formation (Lower Miocene). kokemi 1899. Crassatella CRASSATELLIDAE 1899fl. NJMGP, 2:17, pL 2, fig. 2. 1907«. AMNBA, 14:279. Crassatellites. 1914. NPRMP, 1(3):40 (Patagonico Inferior). Type Locality. — -Santa Cruz. Syntypes.— #325, five valves, coll. J. Bicego. Other Lots.— #326, Cabo Tres Puntas, three valves, coll. Ameghino, 1900; #327, Punta Casamayor, two specimens, coll. Ameghino, 1900. Stratigraphy.- — ^Patagonian Formation (Lower Miocene). lacustris, 1907. Hadriania MURICIDAE: OCENEBRINAE 1907a. AMNBA, 14:185. New name for material misidentified as Urosalpinx pyriformis (Ihering) in Ortmaen (1902:220, pi. 34, fig. 11), not Trophon pyriformis Ihering, 1897L 1914. NPRMP, 1(3):99. Hadriania hatcheri Ortmaen (Patagonico Medio). = Crassilabrum hatcheri (Ortmann, 1900). Other Lots.— #^02, Punta Casamayor, one specimen, C. Ameghino, 1900.; #803, Punta Nodales, one specimen, coll. Ameghino, 1900. Stratigraphy.— P^ldigonmn Formation (Lower Miocene). 236 Annals of Carnegie Museum VOL. 65 lahillei, 1907. Calliostoma TROCHIDAE 1907fl. AMNBA, 14:440, pi. 17, fig. 118 (“Pampeano”). = Calliostoma jucundum (Gould, 1849). Type Locality. — Punta Villarino, Chubut. Type.— 'Hoi in the collection. Remarks. — Included by Ihering among the species from the 'Tampeano” (Pleistocene), this is not a fossil. lahillei, 1907. Pitar VENERIDAE 1907a. AMNBA, 14:303, pi. 11, fig. 75a-b. Not fully described but reported as a new, Recent species in the discussion of Pitar julianum. Remarks. — This form, being Recent, is not in the collection but the figures show a higher shell than the living Pitar rostrata (Koch). As the latter species is extremely variable, P. lahillei may be only an individual variation. See Pitar lahillei ortmanni. laziarina, 1907. Amiantis VENERIDAE 1903. Ameghino, ASCA, 54:190. Cytherea laziarina Ihering, in litteris. 1907a. AMNBA, 14:414, pi. 16, fig. 110. Amiantis. 1914. NPRMP, 1(3):54 (Tehuelche Medio). Type Locality. — Sierra Laziar, Santa Cruz. Syntypes. — #431, ''Chione'' laziarina on label, seven valves, coll. Ameghino, 1900. Stratigraphy. — Araucanean Formation (Lower Pliocene). laziarina, 1907. Myochlamys PECTINIDAE 18971?. RMP, 2:323. Pecten aff. centralis Sowerby. 1902. Ameghino, ASCA, 54:190. Pseudoamusium laziarium, nomen nudum. 1907a. AMNBA, 14:409, pi. 16, fig. 104a-b. 1914. NPRMP, 1(3):35 (Forma^ao Araucana). = Chlamys laziarina (Ihering). Type Locality. — Sierra Laziar, Santa Cruz. Syntypes. — #284, four valves, coll. Ameghino, 1900; #285, Bajo de la Pava, Santa Cruz, one valve, coll. Ameghino, 1897. Stratigraphy. — Araucanean Formation (Lower Pliocene). laziarium, 1907. Calliostoma TROCHIDAE 1907a. AMNBA, 14:399, pi. 16, fig. 105a-b. 1914. NPRMP, 1(3):78 (Formagao Araucana). Type Locality: — -Sierra Laziar, Santa Cruz. Holotype. — #606, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Araucanean Formation (Lower Pliocene). levuana, 1907. Calyptraea CREPIDULIDAE 1907a. AMNBA, 14:523. New name for Trochita araucana in Philippi (1887:87), not Trochita ar- aucana Lesson, 1831. 1914. NPRMP, 1(3): 84. Calyptraea pueyrredona (Patagonico). = Calyptraea pueyrredona Ihering. 1996 Parodiz — Ihering Mollusca 237 Type Locality. — Lago Pueyrredon, Santa Cruz. Syntypes. — #666, two specimens, Lagoa Pueyrredon, coll. Hatcher, received from Ortmann as Galerus araucanus, labelled as “cotypes.” Stratigraphy.— -Patagonian Formation (Lower Miocene). Remarks. — -Ihering mistakenly proposed two new names, C. levuana and C. pueyrredona, for the species misidentified by Philippi (1887) as Trochita arau- cana Lesson, The name C. levuana Ihering is therefore a junior objective synonym and Ihering did not list it in his 1914 catalogue, but placed the corresponding lot under C. pueyrredona. There are five other lots of Calyptraea sp. similar to lot #666 in the Ihering Collection without indication of locality. longior, 1897. Crassatella CRASSATELLIDAE 1897Z?. RMP, 2:247, pi. 5, fig. 34, pi. 6, fig. 37 (Forma^ao Santacruzense). 1902. Ortmann, p. 125, pi. 27, fig. 2. Cras satellites. 1907fl. AMNBA, 14:272. CrassatelUtes. 1914. NPRMP, 1(3):40 (Superpatagonico). Type Locality.— Y&gna Quemada, Santa Cruz. Holotype. — #328, one valve, coll. Ameghino, 1900 (by monotypy). Stratigraphy.— Patagonian Formation (Lower Miocene). Remarks.— Othoi lot reported by Ihering (1914) from Punta Casamayor is miss- ing from the collection. According to Femglio (1949-1950), C. longior has af- finities to Crassatella chubutiensis Femglio. CrassatelUtes Kmeger, 1823, is an unavailable name (= Crassatella Lamarck, 1799). luciferus, 1904. Helcioniscus PATELLIDAE: NACELLINAE 1904. RMLP, 11:235, fig, 7-8. 1907a. AMNBA, 14:123 (araucaniense). 1914. NPRMP, 1(3):71 (Araucano on Pampeana). = Cellana lucifera (Ihering). Type Locality.— PSo Deseado, Santa Cruz, Syntypes.— 41563, two specimens, coll. S. Roth. Stratigraphy.— ''Deseado beds” coeval and equivalent to the Belgrano For- mation in the north (Lower Pleistocene). Remarks. — -Helcioniscus Dali, 1871 Cellana H. Adams, 1869). lujanensis, 1907. Diplodon charruanus HYRIIDAE 1907a. AMNBA, 14:468. Not figured. 1914. NPRMP, 1(3):36 (“Pampeano superior”)- = Diplodon delodontus (Lamarck, 1819). Type Locality.— L\x]an, Buenos Aires Province (“Post-Pampeano”). Syntypes.— One specimen marked “type” exists at the Museum Senckenberg, Frankfurt; in the Ihering Collection there are two other valves (#294), both also marked as “type.” Neither of these are charruanus. Stratigraphy. — Upper Buenos Aires Formation (Pleistocene). Remarks. — See Parodiz (1973). I have observed fossil Diplodon specimens, from Lujan and other places, in strata indicated as “Lujanense” (Upper Pleisto- cene) and none of them are similar to D. charruanus (d’Orbigny), but belong to various species including D. delodontus (Lamarck), D. piceus (Lea), and D. rhu- 238 Annals of Carnegie Museum VOL. 65 acoicus (d’Orbigny). See Parodiz, 1968:7. Unio charruanus “van” lujanensis Doring (1884:328) is a nomen nudum. madryna, 1907. Ostrea OSTREIDAE 1907a. AMNBA, 14:407, pi. 15, fig. lOla-d. 1914. NPRMP, 1(3):25 (Formagao Araucana). Type Locality. — Puerto Madryn, Bahia Nueva, Chubut. Syntypes.— 41191 , seven isolated valves, coll. Ameghino, 1900. Stratigraphy. — Araucanean Formation (Pliocene). Remarks. — Ostrea madryna is intermediate between O. hatcheri Ortmann and O. orbignyi Ihering. magellanica, 1899. Pinna semicostata “van” PINNIDAE 1899fl. NJMGP, 2:213. Pinna. 1907a. AMNBA, 14:242. Atrina magellanica. 1914. NPRMP, 1(3): 19 (Patagonico Medio). = Atrina magellanica (Ihering). Type Locality. — Santa Cruz. Syntypes. — #131, ten valves (fragments); #132, Cabo Tres Puntas, two incom- plete specimens, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene); also Magellanian For- mation (Lower Oligocene) (Feruglio, 1949-1950, at Epuyen and Arroyo Nireco). magellanica^ 1907. Ringicula RINGICULIDAE 1907a. AMNBA, 14:120, pi. 4, fig. 5. 1914. NPRMP, 1(3):69 (Patagonico Inferior). Type Locality. — Camarones, Chubut. Syntypes. — #550, two incomplete specimens, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species is similar to the Recent R. semistriata (d’Orbigny), but with fewer whorls and larger shell. major, 1899. Cancellaria gracilis “van” CANCELLARIIDAE 1899a. NJMGP, 2:35, pi. 2, fig. 10. 1907a. AMNBA, 14:215, pi. 7, fig. 43. Sveltia major. 1914. NPRMP, 1(3):111 (Superpatagonico). = Naronia {Sveltia) major (Ihering), new combination. Type Locality. — Santa Cruz. Holotype.—^H9H, in clayish rock containing specimens of Struthiolaria amegh- inoi, coll. J. Bicego, 1895 (by monotypy). Stratigraphy.— Fatagonian Formation (Lower Miocene). malaspina, 1903. Turritella TURRITELLIDAE 1903. AMNBA, 9:217, pi. 2, fig. 13. 1907a. AMNBA, 14:48. 1914. NPRMP, 1(3):91 (Salamanqueano). Type Locality. — Malaspina, Chubut. Syntypes. — #735, three specimens, coll. Ameghino, 1900. 1996 Parodiz — Ihering Mollusca 239 Stratigraphy, — Salamanca Formation (Paleocene). Remarks. — -Feruglio (1949-1950) mentioned the species from Canadon Perdi- do, north of Comodoro Rivadavia. It has affinities with T. australis Ihering of the Patagonian Formation. Marginella sp. MARGINELLIDAE 1914. NPRMP, 1(3); 106 (Forma§ao Entreriana). Locality.- — La Paz, Entre Rios. Specimens.— ^^62, cast of an unidentifiable species, coll. S. Roth. Stratigraphy. — Parana Formation (Upper Miocene). Marginella {Volvaria) sp. MARGINELLIDAE 1914. NPRMP, 1(3): 107. Locality. — Cabo Tres Puntas, Santa Cruz. Specimen.— ^^64, one specimen, coll. Ameghino. Stratigraphy.— Pdiidigonmn Formation (Lower Miocene). Remarks. — Volvaria Jousseaume, 1875 (= Volvarina Hinds, 1844). matanzasense^ 1907. Dentalium 7DENTALIIDAE 19076E. AMNBA, 14:224. New name for Dentalium sulcosum in Philippi, 1887, not D. sulcosum Sowerby, 1846. 1914. NPRMP, 1(3):66 (“Terciario antiguo”). Type Locality. — Matanzas, Chile. Type. — #531, one specimen received from R. A. Philippi. Stratigraphy. — Probably Navidad Formation (Miocene). matthewi, 1914. Siphonalia FASCIOLARIIDAE: FUSININAE 1914. NPRMP, 1(3): 102, pi. 3, fig. 14 (Patagonico). = Fusinus subrectum (Ihering). Type Locality. — Golfo San Jorge, Santa Cruz. Holotype. — “#845a, one cast, distorted by pressure, with fragments of the shell, coll. Ameghino (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species and S. nodocincta Ihering are both synonyms of Fu- sinus subrectum (Ihering). mendozana, 1907. Exogyra ostracina OSTREIDAE 1907fl. AMNBA, 14:37, text fig. 1 (etage Rocaneen). = Exogyra mendozana Ihering. Type Locality. — “Territorios Nacionales al Sud de Mendoza”; = present Neu- quen Province, or Rio Negro. Holotype. — #4455 (by monotypy). Stratigraphy. — Roca Formation (Paleocene). Remarks. — Label data on the type: “from FI. Ameghino collection found 15 April [1882].” Not listed in catalogue (Ihering, 1914). The specimen was found together with Gryphaea burckhardti Bohm which is known from the Roca For- mation. 240 Annals of Carnegie Museum VOL. 65 meridionalis, 1897. Dosinia VENERIDAE 1897fe. RMP, 2:256, pi. 6, fig. 41 (Forma§ao Santacrazense). 1907^2. AMNBA, 14:300. 1914. NPRMP, 1(3):51 (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Syntypes. — #419, five valves, coll. Ameghino; #420, Canada de los Artilleros, one small valve, coll. Ameghino; #421, Parana, three casts and fragments of valves, coll. J. Bicego, 1895. Stratigraphy.— Patagonian Formation (Lower Miocene); Parana Formation (Upper Miocene); Araucanean Formation (Pliocene); Recent. militarise 1907. Calliostoma TROCHIDAE 1907«. AMNBA, 14:438, pi. 17, fig. 116 (Pampeano). Type Locality.— Pn&xto Militar (now Puerto Belgrano), Prov. Buenos Aires). Syntypes.— lyoscnbod from two specimens; neither are in the Ihering Collection or his 1914 catalogue under this name; the lot used in the description of C. militaris is under C. dalli in the Ihering Collection, labelled with an incorrect locality (Deseado). Priority of C. militaris over C. dalli follows Clench and Turner (1960:75). Stratigraphy.— Polgiano Formation (Pleistocene). militaris, 1907. Tornatina SCAPHANDRIDAE [CYLICHNIDAE] 1907«. AMNBA, 14:435, pi. 17, fig. 133 (Pampeano). = Tornatina candei (d’Orbigny, 1841). See Carcelles and Parodiz, 1938. Type Locality. — Puerto Militar (now Puerto Belgrano), Prov. Buenos Aires. Syntypes.— Two specimens, coll. Ameghino. Stratigraphy.— Potgxano Formation (Pleistocene); Recent. monoceros, 1907. Trophon MURICIDAE 1907fl. AMNBA, 14:183, pi. 14, fig. 92. 1914. NPRMP, 1(3):98. Entacanthus (Patagonico). = Thais (Stramonita) monoceros (Ihering), new combination. Type Locality. — Santa Cruz. Syntypes. — #801, two specimens, coll. Ameghino, 1900. Stratigraphy.— Patagonian Formation (Lower Miocene). montenus, 1907. Trophon varians MURICIDAE 1907a. AMNBA, 14:404. Not figured. 1914. NPRMP, 1(3): 101 (Formagao Araucana). = Trophon varians (d’Orbigny, 1841). Type Locality. — -Monte Espejo, Santa Cruz. Syntypes. — #834, three specimens. Stratigraphy.— Arancanoan Formation (Pliocene). Remarks. — In the original description two specimens were cited, but only one specimen was noted in the 1914 catalogue. 1996 Parodiz — Ihering Mollusca 241 multicoronatus, 1907. Vermetus VERMETIDAE 1907«. AMNBA, 14:167, pi. 5, fig. 27. 1914. NPRMP, 1(3):92 (Patagonico Inferior). = Dendropoma multicoronatus (Ihering), new combination. Type Locality.— Hoxih of Rio Seco (this may be the same as Monte Espejo). Syntypes. — #748, three specimens (Rio Seco and San Julian mixed). Stratigraphy.— Patagonmn Formation (Lower Miocene). Remarks. — The fragments of the tubes seem to belong in the genus Dendro- poma Morch, 1861 {nomina conservanda over Spiroglyphus Daudin, 1800; Si- phonium Gray, 1847. The only species known in the southwestern Atlantic is D. irregularis (d'Orbigny). multicostata^ 1897. Cucullaea CUCULLAEIDAE 1897^. RMP, 2:240, pi. 4, 20 and pi. 5, fig. 29. 1899a. NJMGP, 2:13. 1904. RMLP, 11:9, pi. 1, fig. 2-4. 1907a. AMNBA, 14:232. 1914. NPRMP, 1(3): 15 (Patagonico Inferior and Superpatagonico). Type Locality. — ^La Cueva, Santa Cruz. Holotype.— ^93, coll. Ameghino (by monotypy). Other Lots.- — #94, Santa Cruz, three valves, coll. Bicego, 1897; #95, Canada de los Artilleros, one valve, coll. Ameghino, 1900; #96, Punta Casamayor, six complete specimens and five valves, coll. Ameghino, 1900; #97, Sierra de los Baguales, four specimens, coll. Hauthal. Stratigraphy. — Patagonian Formation (Lower Miocene). Myochlamys, 1907. PECTINIDAE 1907a. AMNBA, 14:106, 251. New genus. GenuS“group name proposed to replace Chlamys Roding, 1798, which was thought to be invalid. The type species of Chlamys Roding is Pecten islandicus Muller, 1776, by subsequent designation of Hermannsen, 1846, and that species would also be the type species of Myochlamys Ihering. Since names from the Bolten Collection are now accepted by the ICZN as authored by Roding (1798), Myochlamys Ihering is an unnecessary replacement name and junior ob- jective synonym of Chlamys Roding. In any event, Myochlamys Ihering, 1907, is preoccupied by Myochlamys Fairmaire, 1876 (Insecta: Coleoptera). necocheanuSy 1907. Trophon MURICIDAE 1907a. AMNBA, 14:404, pi. 14, fig. 106 (Forma^ao Araucano). 1914. NPRMP, 1(3): 100. Trophon laciniatus necocheanus. = Trophon (Austrotrophon) lamellosus (Gmelin, 1791). Type Loca/iYy. —Necochea, Prov. of Buenos Aires. Type.— Hoi in the collection. Stratigraphy. — Araucanean Formation (Pliocene); Recent. Remarks. — Lot #823 contains two specimens from Sierra Laziar and was la- belled by Ihering as “Type,” but this does not agree with the type locality pub- lished in the original description. The two specimens are juveniles of T. lamellosus (Gmelin). Feruglio (1949-1950:268) reported the affinities of T. necocheanus Ihering with T. disparides Wilckens (1911) from Seymour Island in the Antarctic. 242 Annals of Carnegie Museum VOL. 65 negroina, 1907. Chlamys patagonensis “van” PECTINIDAE \901a. AMNBA, 14:22, pi. 3, fig. 11 (etage Rocaneen). = Chlamys patagonensis (d’Orbigny, 1842). Type Locality. — -Rio Negro, kilometers 1216 and 1217. Type. — Not in the collection and not listed in the 1914 catalogue. Stratigraphy. — ?Roca Formation. Remarks. — Ihering (1907<2) said he could not “Juger definitivement de la re- lation de cette forme avec Ch. patagonensis de la formation patagonienne.” neuquena, 1907. Ostrea OSTREIDAE 1899A ASCA, 47:64. Ostrea hemispherica d’Orbigny. 1907a. AMNBA, 14:45, text fig. 3a-c. 1914. NPRMP, 1(3):22 (Cretaceo Superior). Type Locality.— Rocdi, Rio Negro (not Neuquen as the species-group name suggests). Syntypes. — Those specimens figured in Ihering (1907a:text fig. 3a-c) including one marked #151 . Other Lots. — #152, San Juan, Puerto Santa Ana, Misiones, northeast Argentina, fragment reported from “Formacion Guaranitica” of uncertain identification; #153, Roca, Rio Negro; #154, Malaspina, Santa Cruz, five valves; #155, “Cre- taceo de Rio Negro Kilometro 1216,” two valves, coll. Ameghino, 1904. Stratigraphy. — Roca and Salamanca formations (Paleocene). Remarks. — E. Feruglio also found the species in the “Salamanqueano” of the valley of the Rio Chico. The valves from Misiones are fragments of O. guaran- itica Ihering; those from kilometer 1216 on the Rio Negro were probably from or near Roca. Ostrea hemispherica capa Ihering and O. h. paca Ihering are syn- onyms of this name. nodocincta, 1907. Siphonalia FASCIOLARIIDAE: FUSININAE 1897A RMP, 2:3299. Siphonalia aff. nodosa Martens. 1907a. AMNBA, 14:193, pL 6, fig. 37. Siphonalia nodocincta. 1914. NPRMP, 1(3): 102 (Patagonico Medio). = Fusinus subrectus (Ihering). Type Locality. — Cabo Tres Puntas, Santa Cruz. Holotype. — #841, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — The type specimen has only the upper part of the aperture preserved and the siphonal canal is missing. On the Argentine coast (Recent) the genus is represented by Fusinus frenguelli (Carcelles, 1953). The species most similar to F. subrectus is F. brasiliensis (Grabau, 1903). nodosoplicatus, 1897. Pecten PECTINIDAE 1897^7. RMP, 2:227, pi. 5, fig. 36 (Formagao Santacruzense). 1907a. AMNBA, 14:256. Myochlamys. 1914. NPRMP, 1(3):31 (Patagonico Medio-Superior). = Chlamys (Zygochlamys) nodosoplicatus (Ihering); see Morra (1985). Type Locality. — La Cueva, Santa Cruz. Syntypes. — #251, four valves, coll. Ameghino, 1900. 1996 Parodiz — Ihering Mollusca 243 Other Lots. — #252, Yegua Quemada, Santa Cruz, five valves, coll. Ameghino, 1900; #253, Monte Espejo, Santa Cruz, one valve, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. —Tht name was first given in Ihering (1897i?:227) as Pecten nodo- sopiicatus [sic], an incorrect original spelling for P. nodosoplicatus. nucleus, 1899. Glycimeris [sic] HIATELLIDAE 1899a. NJMGP, 2:23, pL 1, fig. 7. 1907a. AMNBA, 14:326, pi. 12, fig. 85a-b. Panopaea [sic]. 1914. NPRMP, 1(3):62 (Patagonico). = Panopea nucleus (Ihering). Type Locality. — Cabo Tres Puntas, Santa Cruz. Holotype. — #515, coll. Ameghino (by monotypy). Other Lots. — #513, Santa Cruz, one specimen, coll. J. Bicego, 1895, reported by Ihering (1914) as type but is not from type locality; #514, Corral Foyel, one cast, coll. Hauthal; #516, Sierra de los Baguales, one cast, coll. Hauthal; #4464, San Julian, coll. Ameghino; #2480, Rio Foyel, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species is related to Panopea nucleoides Wilckens (1911) from the Tertiary of Antarctica. observationis, 1907. Turbonilla PYRAMIDELLIDAE 1907a. AMNBA, 14:144. New name for one of the specimens of Turbonilla cuevensis misidentified by Ortmann (1902:174, pi. 33, fig. 8a-b), not Turbonilla cuevensis Ihering, 18971?. Type Locality. — Monte Observacion, Santa Cruz. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — A questionable species; Ihering (1907^:144) states, “Espece inter- mediaire entre T. cuevensis and T. Iheringi. observatoria, 1907. Cancellaria gracilis “van” CANCELLARIIDAE 1907a. AMNBA, 14:214. New name for specimens of C. gracilis as figured by Ortmann (1902:235, pi. 36, fig. 3a-b). = Cancellaria gracilis (Ihering). Type Locality. — Monte Observacion, Santa Cruz. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — No description; in the discussion of Cancellaria gracilis Ihering as “differente par les caracteres de Poverture”; these characters are insufficient to identify the form. observatoria, 1907. Nucula NUCULIDAE 1907a. AMNBA, 14:227, pi. 7, fig. 49a-b. 1914. NPRMP, 1(3): 13 (Patagonico Superior). Type Locality. — Monte Observacion, Santa Cruz. Holotype. — #70, one specimen within a lot of Nucula semiornata d’Orbigny received from Ortmann (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Ihering (1914:13) gives the type locality as Camarones from spec- imens collected by C. Ameghino in 1900, but the original description (Ihering, 244 Annals of Carnegie Museum VOL, 65 190742:227) states the type locality was Monte Observacion from which the spe= cies derives its name. Nucula semiornata d’Orbigny, 1846, of which N. reticularis Ortmann, 1902, is a synonym, is also found at both localities. octocostatum, 1897. Dentalium DENTALIIDAE 18971?. RMP, 2:266, pL 4, fig. 16 (Forma^ao Santacruzense). 1907a. AMNBA, 14:225. Dentalium octocostellatum Pilsbry and Sharp. 1914. NPRMP, 1(3):66 (Patagonico Inferior-Superior). = Dentalium octocostellatum Pilsbry and Sharp, 1898. Type Locality, — Yegua Quemada, Santa Cruz. Syntypes. — #532, nine specimens, coll. Ameghino, 1894. Other Lots. — #533, 24 specimens, Camarones, coll. Ameghino, 1900; #534, Canada de los Artilleros, 100 specimens, coll. Ameghino, 1900. Stratigraphy. — Patagonian Eormation (Lower Miocene). Remarks. — Dentalium octocostellatum Pilsbry and Sharp (1898:211) replaced D. octocostatum Ihering which was preoccupied by Dentalium octocostatum Fraas, 1867. Odontogryphaea^ 1903. GRYPHAEIDAE 1903. AMNBA, 9:194. New subgenus of Gryphaea Lamarck. 1907a. AMNBA, 14:43, 44. 1945. Stenzel, BGSA, 56(12, pt. 2): 1202. Type Species. — Gryphaea rostrigera Ihering, by subsequent designation of Stenzel, 1945. Stratigraphy. — Salamanca Formation (Paleocene). Remarks. — Ihering made Odonto gryphaea a subgenus of Gryphaea Lamarck for the early Tertiary species with crenulated margins. Stenzel considered it a genus, with Gryphaea rostrigera Ihering as the type species. According to Stenzel Odonto gryphaea is convergent with Gryphaea but not closely related. Outside Patagonia Odonto gryphaea is known from the Lower Eocene of the southeastern United States (O. thirsae [Gabb, 1862]; see Palmer and Brann, 1965:225) and from the Paleocene of France and India (Stenzel, 1945). orbignyi, 1897. Ostrea OSTREIDAE 1897# RMP, 2:222, pi. 9, fig. 52 (Formagao Santacruzense). 1907a. AMNBA, 14:17, pi. 2, fig. 9 (spelled d’orbignyi). 1914. NPRMP, 1(3):24 (Patagonico and Superpatagonico). Type Locality, — La Cueva, Santa Cruz. Syntypes. — #186, six valves, coll. Ameghino, 1904. Other Lots. — #187, Santa Cruz, two valves, coll. Ameghino; #188, Yegua Que- mada, one valve, coll. Ameghino; #189, Sierra de los Baguales, five valves, coll. Hauthal; #190, Deseado, one valve, coll. S. Roth; #191, Deseado, three valves, coll. S. Roth. Stratigraphy. — Patagonian Formation (Lower Miocene). ortmanni, 1907. Leda NUCULANIDAE 1907a. AMNBA, 14:229. New name for Leda oxyrhyncha in Ortmann (1902:83, pL 23, fig. 2a-b), not Leda oxyrhyncha Philippi, 1887. = Nuculana ortmanni (Ihering). 1996 Parodiz — Ihering Mollusca 245 Type Locality.— Cruz and Rio Negro (Ortmann). Stratigraphy.- — Patagonian Formation (Lower Miocene). Remarks. — See Leda camaronesia for note on Nuculana. ortmanni, 1899. Lucina LUCINIDAE 1899a. NJMGP, 2:18, pi. 2, fig. 3. 1902. Ortmann, p. 131, pi. 27, fig. 5. 1907a. AMNBA, 14:289, Phacoides. 1914. NPRMP, 1(3):46 (Patagonico Inferior-Medio). = Lucina ortmanni (Ihering). Type Locality.— C^o Tres Puntas, Santa Cruz. Syntypes. — #384, five valves, coll. Ameghino. Other Lots. — #382, Santa Cruz, five specimens (labelled “typo”); #383, Bajo de San Julian, one specimen, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). ortmanni, 1907. Marcia VENERIDAE 1907a. AMNBA, 14:304. New name for Venus navidadis in Ortmann (1902:141, pi. 27, fig. 12), not Venus navidadis Philippi, 1887. 1914, NPRMP, 1(3):52. Marcia navidadis Philippi. = Samarangia navidadis (Philippi). Type Loca/jiy.— Talcahuano, Chile (Philippi, 1887). #433, one specimen, received from the Valparaiso Museum. Stratigraphy.-— Te,^X\^d:y Inferior of Chile (Philippi, 1887). ortmanni, 1907. Pitar lahillei VENERIDAE 1907a. AMNBA, 14:453, pi. 18, fig. 124a-b. 1914. NPRMP, 1(3):51 (Pampeano). = Pitar rostrata (Koch, 1844). Type Locality.— 12 de Cabo Tres Puntas, Santa Cruz (probably Sierra La- ziar). Syntypes. — #425, nine valves, coll. Ameghino, 1900. Other Lot. — #426, one valve, coll. Ameghino, 1900. Stratigraphy. — Araucanean Formation (Pliocene). Remarks. — Femglio (1949-1950, vol. 3:81) reports the species from the Plio- cene of San Julian. It is common in the Recent. See Pitar lahillei Ihering. ortmanni, 1907. Polynices NATICIDAE 1907a. AMNBA, 14:157, pi. 5, fig. 18. New name for Natica ovoidea in Ortmann (1902:380), not Natica ovoidea Philippi, 1887. 1914. NPRMP, 1(3):85 (Patagonico— Superpatagonico). = Polynices famula (Philippi). Type Locality. — -Santa Cruz (specimens figured by both Ortmann and Ihering). Other Lots.— ^616, Yegua Quemada, coll. Ameghino, 1900; #677, two speci- mens, coll. J. Bicego, 1897. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.— Polynices jorgensis Ihering is also a synonym. 246 Annals of Carnegie Museum VOL. 65 ortmanni, 1907. Tellina PSAMMOBIIDAE 1907fl. AMNBA, 14:313, 315. New name for Tellina tehuelcha in Ortmann (1902:147, pi. 29, fig. 3), not Tellina tehuelcha Ihering, 1899. 1914. NPRMP, 1(3):58. Sanguinolaria tehuelcha (Ihering). = Sanguinolaria tehuelcha (Ihering). Type Locality.—Shell Gap at Rio Chico, Santa Cruz (Ortmann). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — There is no material in the Ihering Collection under this name, and only the type lot is curated under the corresponding Tellina tehuelcha Ihering. ortmanni, 1907. Urosalpinx MURICIDAE 1907a. AMNBA, 14:188. New name for material misidentified as Urosalpinx leucostomoides (Sow- erby) in Cossmann (1899a:pl. 10, fig. 7), not Triton leucostomoides Sowerby (1846:383, fig. 60). 1914. NPRMP, 1(3):98 (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Syntypes. — #800, four specimens, coll, Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species resembles Urosalpinx archipatagonica Ihering from the same locality. It is synonymous with Urosalpinx ''pleurotomoides'’ Cossmann (1903:50), lapsus calami and misidentification of U. leucostomoides (Sowerby). osborni, 1914. Gibbula 7TROCHIDAE 1914. NPRMP, 1(3):75, pi. 3, fig. 11 (Patagonico Inferior). Type Locality.— Pan de Azucar, Santa Cruz. Holotype. — #590, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — It may not be a Gibbula', it is more TwrZ^o-like; probably a new genus. paca, 1907. Ostrea hemisphaerica [sic] “van” OSTREIDAE 1907a. AMNBA, 14:12. Not figured (Roca). 1914. NPRMP, 1(3):22. Ostrea neuquena paca (Roca, Cretaceo Superior). = Ostrea neuquena Ihering. Type Locality.— Poca, Rio Negro. Syntypes. — #156, three valves, coll. A. Romero, 1903. Stratigraphy. — ^Roca Formation (Paleocene). pachychila, 1907. Bulla BULLIDAE 1907a. AMNBA, 14:363. New name for Bulla aff. elegans in Borchert (1901:47, pi. 4, fig. 7), not Bulla elegans Gray, 1825 (= Haminea guildingi [Sowerby]). Type Locality. — Parana, Entre Rios. Stratigraphy. — Parana Formation (Upper Miocene). Remarks. — Ihering (1907^:363) states, “La determination de Borchert est con- sequent inexacte, non seulement pour Lespece, mais aussi pour le genre au sous- genre. ...” palaeopatagonica, 1903. Cardita 1903. AMNBA, 9:215, fig. 12. 1907a. AMNBA, 14:24, 47. Venericardia (Rocaneen et Patagoneen). CARDITIDAE 1996 Parodiz — Ihering Mollusca 247 Type Locality.- — Rfo Chico, Chubut. Hoiotype.-~^360, coll. Ameghino, 1900 (by monotypy). Stratigraphy.— PaXSLgonmn Formation (Lower Miocene). "Ihering (1904) referred to another specimen, a cast from Roca, which is not listed in the 1914 catalogue and was not found in the collection. The species has affinities with Venericardia inequalis Philippi, also from the Pata- gonian Formation. It is similar to ""Cardita"' iheringi Bohm, 1903:72 (= Cardita burckhardti Ihering; see Burckhardt, 1902:216, fig. 1-7). pampUy 1914. BuUmulus sporadicus ORTHALICIDAE: BULIMULINAE 1914. NPRMP, 1(3):70~7L Not figured. 1944. Parodiz, CZMHN, 1(17): 1-2, fig, 2-3. Peronaeus (Lissoacme) ameghinoi (Ihering). = Peronaeus {Lissoacme) ameghinoi (Ihering). See BuUmulus ameghinoi Ihering. Type Locfllily.— “Buenos Aires.” Syntypes.— 41=562, coll. Ameghino, 1910. Stratigraphy.— Buenos Aires Formation (Pleistocene). Remarks.- — The type locality of P. (L.) ameghinoi is Chapadmalal, Buenos Ai- res Province; the specimens of B. pampa are probably from Lujan where Amegh- ino made extensive collections. The species ameghinoi occurs in the region of San Matias Gulf as the subspecies P. (L.) ameghinoi madrynensis Parodiz, 1944. pampa^ 1914. Siphonaria lessoni SIPHONARIIDAE 1914. NPRMP, 1(3):69. Not figured (Pampeano Inferior). = Siphonaria lessoni (Blainville, 1826). Type Locality.— "'Buenos Aires” probably near or at Chapadmalal. Syntypes.— #556, two specimens, coll. Ameghino, 1910. Sfrafigraj?/!};.— -B'Clgrano Formation (Lower Pleistocene); Recent. pampeama^ 1907. Bullia gradata NASSARIIDAE 1907fi. AMNBA, 14:445. Not figured. 1914. NPRMP, 1(3):105 (Pampeano). 1933. Feraglio, ARMGB, 8 bis:236, 239. 1939. Parodiz, Physis, 17:754, fig. 3-4. Buccinanops. = Buccinanops gradatum (Deshayes, 1844). Type Locality.— Buetlo Militar, Prov. Buenos Aires. Syntypes.—#H55, 19 specimens, coll. Ameghino, 1910. Stratigraphy.— Belgmno Formation (Lower Pleistocene); also Recent and com- mon from Uruguay to northern Patagonia. pampeano, 1907. Chione VENERIDAE 1907a. AMNBA, 14:454, pL 18, fig. 125a-b. 1914. NPRMP, 1(3);56. Type Locality.San Julian, Santa Cruz. Syntypes. — #468, 18 valves, coll. Ameghino, 1900. Stratigraphy.— Deseado beds coeval with the Belgrano Formation (Pleisto- cene), Remarks.— This is most likely a synonym of Protothaca antiqua (King), the specimens in the type lot being slightly more elongated. 248 Annals of Carnegie Museum VOL. 65 panensiSy 1914. Area 7ARCIDAE 1914. NPRMP, 1(3): 16-17, pi. 2, fig. 1 (Formagao Patagonica). Type Locality.-— Pan de Azucar, Santa Cruz. Holotype. — #109, a cast, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — The unique specimen is an incomplete cast of questionable specific determination, but not likely an Area. panis, 1907. Panopaea [sic] HIATELLIDAE 1907«. AMNBA, 14:327, pi. 13, fig. 87. 1914. NPRMP, 1(3):64, pi. 2, fig. 9a-b (Patagonico Inferior). = Panopea panis Ihering. Type Locality. — Pan de Azucar, Santa Cruz. Holotype. — #518, corresponding to the specimen figured in Ihering (1914), coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Ihering (1914:64) wrote that the species was not figured until 1914, but he did figure a specimen in 1907 without mentioning it in the text. The two figures {1901 a and 1914) appear slightly different, but the former is a photograph and the latter is a drawing. The original description is based on “P unique ex- emplaire” interpreted here as the specimen figured in the 1914 catalogue. Panope and Panopaea of authors are misspellings of Panopea. patagonensis, 1907. Pododesmus ANOMIIDAE 1907«. AMNBA, 14:266, pi. 9, fig. 63a-b. 1914. NPRMP, 1(3):27 (Patagonico Inferior). Type Locality. — Pan de Azucar, Santa Cruz. Syntypes. — #213, six valves. Other Lots. — #214, Pan de Azucar, Santa Cruz, four valves, coll. Ameghino, 1900; #215, Rio Chico, Chubut, one valve, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). patagonensis, 1897. Potamides POTAMIDIDAE 1897^. RMP, 2:315, text fig. 20 (first of two figures numbered 20; the other figure numbered 20 on page 316 is Ostrea pyrotheriorum.) 1899ZJ. ASCA, 47:67. Melania sp. 1907a. AMNBA, 14:52. 1914. NPRMP, 1(3):92 (Salamanqueano). 1969. Parodiz, ACM, 40:152. Type Locality. — “Lehuen-ark” error for Sehuen Aik (e). Valley of Sehuen Riv- er. Lectotype. — #749, the specimen figured by Ihering (1897^:text fig. 20) is here designated the lectotype. The type lot consisted of nine syntypes. Stratigraphy. — Sehuenan Formation (Paleocene). The name P. patagonensis was mentioned frequently in geological literature by Ameghino, Windhausen, Frenguelli, Feruglio, and others, and some of the references may correspond to Potamides chaliana Parodiz, 1969. 1996 Parodiz — Ihering Mollusca 249 patagonemsiSf 1903. Rostellaria STROMBIDAE 1903. AMNBA, 9:209. Not figured. 1904. RMLP 11:239, pi. 2, fig. 11. 1907£I. AMNBA, 14:29. Aporrhais (Rocan&n), 1914. NPRMP, 1(3):94 (Salamanqueaeo). = Tibia patagonensis (Ihering). Type Locality.— Rio Negro. Syntypes.— #759, two casts, specimens collected by C. Ameghino and S. Roth in the same lot. Stratigraphy.— Roca Formation (Paleocene). Remarks.— Aporrhais rocai Bohm, 1903:72, is synonymous but unavailable {nomen nudum). Both Rostellaria patagonensis and R. rocai were from the same locality and strata, and known only by internal casts; the variations in both species show similarities. Rostellaria Lamarck, 1799 (= Tibia Roding, 1798). patagonica, 1907. Abra SEMELIDAE 1907a. AMNBA, 14:316, pi. 2, fig. 82a-b. 1914. NPRMP, 1(3):59 (Patagonico Inferior). = 7 Abra Uruguay ensis (Pilsbry, 1879). Recent. Type Locality.— Chubut. Syntypes.— #4^9, 25 valves, colL Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). patagonica, 1897. Area ARCIDAE 1897^. RMP, 2:235, pi. 4, fig. 25, and pL 5, fig. 30. 1902. Ortmann, p. 93, pL 25, fig. 3a-b. 1907a. AMNBA, 14:236. Area umbonata Lamarck. 1914. NPRMP, 1(3): 18 (Superpatagonico). = Area umbonata Lamarck. Type Locality.— Yegua Quemada, Santa Cruz. Type.—ThG type lot containing four valves reported by Ihering (1914) is not in the collection. Other Lots.- — #118, Cabo Tres Puntas, colL Ameghino, 1900; #119, Canada de los Artilleros, 25 valves, coll. Ameghino, 1900; #120, San Julian, one specimen, coll. Ameghino, 1900. Stratigraphy.— Fatagonmn Formation (Lower Miocene). patagonica, 1897. Bulla 7RETUSIDAE 1897^. RMP, 2:271, text fig. 8 (Fomia^ao Santacrazeese), 1902. Ortmann, p. 246, pL 37, fig. 8a, c, 1907a. AMNBA, 14:121. 1914. NPRMP, 1(3):69 (Superpatagonico). = IRetusa patagonica (Ihering), new combination. Type Locality.— Yegua Quemada, Santa Craz. Syntypes.-— #551, 25 specimens, coll. Ameghino. Other Lot.— #552, Canada de los Artilleros, 12 specimens, coll. Ameghino, Patagonian Formation (Lower Miocene). 250 Annals of Carnegie Museum VOL. 65 patagonicttf 1907. Lahillia angulata CARDIIDAE 1897Z?. RMP, 2:257, text fig. 2 on p. 258. Amathusia angulata Philippi. \901a. AMNBA, 14:294. Lahillia angulata patagonica Ihering. 1914. NPRMP, 1(3):50 (Patagonico Inferior-Super). = Lahillia angulata (Philippi, 1887). Type Locality. — Sierra de los Baguales, Santa Cruz. Holotype. — #413, cast with portion of the shell, coll. Hauthal (by monotypy). Other Lots. — #412, Santa Cruz, one cast, coll. J. Bicego, 1897; #414, Yegua Quemada, one valve, large, 21 cm, but broken. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.-— In the collection there is also a cast in plaster of Paris, #4462, from the specimen (?type) of Amathusia angulata sent to Ihering by Philippi from Navi- dad, Chile. Cossmann (1899c: 134) listed the species as Iheringia angulata (Philippi). patagonica^ 1907. Lima LIMIDAE 1907a. AMNBA, 14:262, pi. 9, fig. 60. 1914. NPRMP, 1(3):29 (Patagonico Medio-Inferior). Type Locality. — Punta Nueva, Santa Cruz. Syntypes. — #230, nine valves, coll. Ameghino, 1900; #231, Golfo San Jorge, one valve, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). patagonica^ 1897. Tellina TELLINIDAE 1897/?. RMP, 2:261, pi. 5, fig. 261 (Forma^ao Santacruzense). 1907a. AMNBA, 14:315. 1914. NPRMP, 1(3):59 (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Syntypes.— three valves, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — See Tellina patagonica camaronesia Ihering. patagonica, 1897. Voluta MARGINELLIDAE 1897/?. RMP, 2:306, pi. 3, fig. 6 (Formagao Santacruzense). 1907a. AMNBA, 14:200. Neoimbricaria. 1914. NPRMP, 1(3): 107. = Neoimbricaria patagonica (Ihering), as figured in Wenz, 1938:1297, fig. 3701. Type Locality. — Yegua Quemada, Santa Cruz. Syntypes. — #868, five specimens, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). COSTELLARIIDAE [VEXILLIDAE] patagonica, 1907. Vulpecula 1907a. AMNBA, 14:199, pi. 7, fig. 48a-b. 1914. NPRMP, 1(3): 107 (Patagonico Medio). = Vexillum patagonicus (Ihering). Type Locality. — Golfo San Jorge, Santa Cruz. Holotype. — #865, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Vulpecula Blainville, 1824 (= Vexillum Roding, 1798). 1996 Parodiz — Ihering Mollusca 251 patagonicunty 1907, Cardium CARDIIDAE 1907fl, AMNBA, 14:293, pL 11, fig. 73. 1914. NPRMP, 1(3):48 (Patagonico). Type Locality.— Cruz, Holotype.— #396, colL J, Bicego, 1897 (by monotypy). Other Lot— #391 , Cabo Tres Puetas, three valves, coll. Ameghino, 1900. Stratigraphy.— ¥diidigommi Formation (Lower Miocene). patagomicuSy 1907. Actaeon ACTAEONIDAE 1907fl. AMNBA, 14:117, pL 4, fig. 1. 1914, NPRMP, 1(3):68 (Superpatagonico). Type Locality.- — Canada de los Artilleros, Santa Cruz. Syntypes.— #546, two specimens, coll. Ameghino, 1900. -Patagonian Formation (Lower Miocene). Remarks.— A very distinct, more elongated species than Actaeon argentinus Ihering from the same locality and strata, and perhaps not congeneric. patagomicuSy 1907. Cras satellites CRASS ATELLIDAE 1907fl. AMNBA, 14:279. 1914. NPRMP, 1(3):40 (Patagooico Inferior). = Crassatella kokeni Ihering. Type Locality.— CsL-maiones, Chubut. Syntypes.— #332, two valves, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.- — -Ihering named C. patagonicus as being different only in size from C. kokeni as figured by Ortmann (1902:123, pL 38, fig. 10). He gave neither description nor illustrations of C. patagonicus. Cras satellites Krueger, 1823 (= Crassatella Lamarck, 1799). patagonicus, 1904. Helcioniscus luciferus PATELLIDAE: NACELLINAE 1904. RMLP, 11:285. Not figured. 1907«. AMNBA, 14:436. 1914. NPRMP, 1(3):436 (Forma^ao Pampeana). = Cellana lucifera (Ihering). Type Locality.— Deseado, Santa Cruz. Syntypes.-— #564, two specimens, coll. Ameghino, 1900. Stratigraphy. — -“Deseadense,” Pleistocene marine of Deseade (= Belgrano Formation). Remarks.- — -See Helcioniscus luciferus Ihering. paucicostata, 1907. Scalaria rugulosa “van’’ EPITONIIDAE 1907a. AMNBA, 14:140. Not figured. 1914. NPRMP, 1(3):81 (Patagonico Inferior). = Opalia (Nodoscala) rugulosa (Sowerby, 1846). Type Locality.— Mananiml Salado, Santa Cruz. Holotype.— #631 , coll. Ameghino, 1900 (by monotypy). Stratigraphy.— Vaiagonmn Formation (Lower Miocene). 252 Annals of Carnegie Museum VOL. 65 Remarks. — Scalaria'' rugulosa Sowerby, 1846, is a species of wide distri- bution from New Zealand to Chile and is also abundant in the Tertiary of Pata- gonia in Chubut and Santa Cruz. pauciradiata^ 1899. Cardium philippii CARDIIDAE 1899«. NJMGP, 2:15. 1907a. AMNBA, 14:104, 291. Cardium philippii pauciradiatum. 1914. NPRMP, 1(3):48 (Patagonico). = Cardium philippii Ihering. Type Locality. — Santa Cruz. Syntypes. — #402, three valves, coll. Bicego, 1897. Stratigraphy. — In Ihering’s table (1907a: 104) the provenance of pauciradiata is uncertainly indicated. Probably Patagonian Formation (Lower Miocene). Remarks. — There is one additional specimen in the Ihering Collection, #2481, from Rio Foyel, collector unknown. paucispina, 1897. Pholas PHOLADIDAF 1897i?. RMP, 2:265, text fig. 5 (Forma^ao Santacruzense). 1907a. AMNBA, 14:329. Barnea paucispina (Ihering). 1914. NPRMP, 1(3):65 (Superpatagonico). = INetastoma paucispina (Ihering), new combination. Type Locality. — Yegua Quemada, Santa Cruz. Syntype. — #521, only one of the three valves mentioned by Ihering (1914); coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — The specimen figured in Ihering (1897^) is a restoration. Nettas- tomella Carpenter, 1865, is an unnecessary replacement name for Netastoma Car- penter, 1864. paucisquamatus, 1907. Pododesmus ANOMIIDAE 1907a. AMNBA, 14:268, pi. 10, fig. lOa-b. 1914. NPRMP, 1(3):27 (Patagonico Inferior). Type Locality. — Southeast of Punta Nueva, Santa Cruz. Holotype.—^220, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). pazensiSf 1914. Pododesmus 7ANOMIIDAE 1914. NPRMP, 1(3):28, pi. 2, fig. 2 (Forma§ao Entreriana). Type Locality. — La Paz, Entre Rios. Holotype. — #223, coll. S. Roth (by monotypy). Stratigraphy. — Parana Formation (Upper Miocene). Remarks. — Ihering (1914:28) states, “. . . esta descrip^ao ha de ser considerada como provisoria.” The generic position of this species is uncertain, but it is not a species of Pododesmus. percrassttf 1897. Ostrea OSTREIDAE 1897^. RMP, 2:221, text fig, 1 (and pi. 9, fig. 53 as Ostrea patagonica of authors; the true O. patagonica d’Orbigny in the plate is fig. 52). 1996 Parodiz— =Ihering Mollusca 253 1902, Hauthal, RMLP, 10:46. Ostrea hatcheri Ortmann. 1902. Ortmann, p. 99. Ostrea ingens in part. 1907£I. AMNBA, 14:18. Ostrea hatcheri Ortmann. 1914. NPRMP, 1(3):23, Ostrea hatcheri (Patagonico Superior). = Ostrea hatcheri Ortmaen, 1897. Type Locality.— Cruz. Syntypes.—#113, five of the six valves in original type lot are present in the collection. Other The name is preoccupied by Ostrea percrassa Conrad, 1840, a Miocene species from North America. There are other lots in the collection, under Ostrea hatcheri #174-183, from different localities including Puerto Madrye, Camarones, Pan de Azucar, north of Rio Seco, Rfo Chico of Chubut, Punta No- dales, southeast of Punta Nueva, and Deseado, collected by S. Roth, J. Bicego, and C. Ameghieo. There are a total of 90 specimens in these lots. Stratigraphy.— Vdilstgonim. Formation (Lower Miocene). perplama, 1897. Teliina PSAMMOBIIDAE 18971?. RMP, 2:259, pL 6, fig. 39 (Forma^ao Santacrazense). 1907fl. AMNBA, 14:313. Sanguinolaria. 1914. NPRMP, 1(3):58 (Superpatagonico). = IPsammotella perplana (Ihering). Type Lom/ily.— La Cueva, Santa Cruz. Syntype.—4A%1^ only one specimen remaining of the three listed by Ihering (1914), coll. Ameghino, 1900. Stratigraphy.— PdildigomMi Formation (Lower Miocene). PhacoMes sp. LUCINIDAE: MILTHINAE 1914. NPRMP, 1(3):46. = Miltha iheringiarta Doello-Jurado (1919:558). Locality.— Paz, Entre Rios. Specimens.— One specimen from Puerto Madryn, Chubut, coll. J. Bicego, 1897; there are three other casts in the collection, #388 from La Paz, Entre Rio, coll. S. Roth. Stratigraphy.— The species is known from, both Parana Formation of Entre Rios and Rio Negro Formation of Chubut (Upper Miocene). pMMppii 1897. Cardium CARDIIDAE 18971?. RMP, 2:249, pi. 6, fig. 40 (Forma^ao Santacrazense). 1907a. AMNBA, 14:291. 1914. NPRMP, 1(3):48 (Superpatagonico). Type Locality.— Yegua Quemada, Santa Cruz. Holotype.—#39H, one valve, C. Ameghino (by monotypy). Other Lots.— #399, Yegua Quemada, Santa Craz, six valves, coll. Ameghino; #400, Santa Cruz, three casts, coll. J. Bicego, 1897; #401, Canada de los ArtiL leros, one valve, coll. Ameghino, 1900. Stratigraphy.— PaiagomMi Formation (Lower Miocene). This is Cardium multiradiatum (in part) in Philippi (1887:171, pL 38, fig. 9), not Cardium multiradiatum Sowerby, 1833. See also Cardium huttoni Ihering. 254 Annals of Carnegie Museum VOL. 65 philippiif 1907. Gibbula 7RAPHISTOMATIDAE 1907fl. AMNBA, 14:132, pi. 4, fig. 6a-c. 1914. NPRMP, 1(3):74 (Patagonico Medio). Type Locality. — Cabo Tres Puntas, Santa Cruz. Syntypes. — #585, four specimens, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.- — This species belongs to the Pleurotomariacea, and to a genus similar to Trophidostropha Longstoff from the European Carboniferous, but probably undescribed. E. Feruglio collected it at Epuyen. piconuSy 1907. Pecten PECTINIDAE 1907a. AMNBA, 14:46, text fig. 4. 1914. NPRMP, 1(3):36 (Salamanqueano). Type Locality. — Pico Salamanca, Santa Cruz. Holotype. — #292, single valve, coll. Ameghino, 1909 (by monotypy). Stratigraphy. — Salamanca Formation (Paleocene). Remarks. — This species may belong in Pecten sensu lato; its taxonomic posi- tion is questionable. pilsbryi, 1899. Voluta VOLUTIDAE 1899a. NJMGP, 2:34, pi. 12, fig. 9. 1902. Ortmann, pi. 37, fig. la-b. 1907a. AMNBA, 14:210. Cymbioia. 1914. NPRMP, 1(3): 109 (Superpatagonico). = Adelomelon {P achy cymbioia) pilsbryi (Ihering). Type Locality. — Santa Cruz. Syntypes, — #883, one specimen plus two casts, coll. J. Bicego, 1897. Stratigraphy. — Patagonian Formation (Lower Miocene). pisunty 1907. Lima LIMIDAE 1907a. AMNBA, 14:263, pi. 9, fig. 61a-c. Limatula. Type Locality. — Southeast of Punta Nueva, Santa Cruz. Syntypes. — #228, 16 valves, coll. Ameghino, 1900; #229, Punta Casamayor, 12 valves, coll. Ameghino, 1900. Stratigraphy. — Patagonian Formation (Lower Miocene). platUy 1907. Cardita CARDITIDAE 1907a. AMNBA, 14:451, pi. 18, fig. 23a-b. = Carditamera plata (Ihering). Type Locality. — Monte Hermoso, Prov. de Buenos Aires. Type. — Not in the collection. Stratigraphy.- — Belgrano Formation (Lower Pleistocene) and Querandinan (Up- per Pleistocene); Recent, plicifertty 1897. Marginella 18971?. RMP, 2:308, text fig. 19. 1902. Ortmann, p. 225, fig. 2 (with supplementary description). MARGINELLIDAE 1996 Parodiz—Ihering Mollusca 255 1907fl. AMNBA, 14:200. Neoimbricaria. 1914. NPRMP, 1(3): 107. Neoimbricaria (Superpatagoeico). = 1 Neoimbricaria plicifera (Ihering). Type Locality,- — Yegua Quemada, Santa Craz. Holotype,- — #869, coll. Ameghino, 1900 (by monotypy). Stratigraphy.^Patagonmn Formation (Lower Miocene). portenittf 1907. Myochlamys patagonensis PECTINIDAE 1907fl. AMNBA, 14:377. Not figured. = Chiamys patagonensis portenia (Ihering). Type Localily.'— Puente Alsina, Buenos Aires, from a perforation 80 m deep. rype.=Not in the collection. Stratigraphy.— Formation (Upper Miocene). Remarks.— In the original description Ihering stated that he received several specimens from Ameghino. praenumciuSj 1897. Pecten PECTINIDAE 18971?. RMP, 2:230. Not figured. 1902. Ameghino, ASCA, 54:224. Chiamys espejoana Ihering, in litteris. 1907£I. AMNBA, 14:251, pi. 7, fig, 54. Myochlamys. 1914. NPRMP, 1(3):33 (Patagonico Inferior-Superpatagonico). = Chiamys praenuncius (Ihering). Type Locality.— Punl^ Nova (Nueva), Santa Cruz. Syntypes.—^26H, a complete specimen, eight valves. Other Lots.- — #269, Monte Espejo, four valves, coll. Ameghino; #270, Yegua Quemada, Santa Cruz, two valves, coll. Ameghino. Patagonian Formation (Lower Miocene). princepSf 1907. Neomphaiius TROCHIDAE 1907a. AMNBA, 14:135, pi. 4, fig. 8a~b. 1914. NPRMP, 1(3):76 (Patagonico Medio). = Tegula patagonica (d’Orbigny, 1840). Type Locality.— Punta Casamayor, Santa Cruz. Hoiotype.— #594, coll. Ameghino, 1900 (by monotypy). Stratigraphy.— Patagonian Formation (Lower Miocene). Ihering did not compare this species with T. patagonica which is known from the Araucaneaii Formation (Pliocene), and from Rio de Janeiro to northern Patagonia (Recent). pnsca, 1907. Modiolarca 7GAIMARDIIDAE 1907a. AMNBA, 14:276, pL 14, fig. 94. 1914. NPRMP, 1(3):39 (Patagonico Medio). = IGaimardia prisca (Ihering). Type Localily. -^Southeast of Punta Nueva, Santa Cruz. Holotype.— #322, coil. Ameghino, 1900 (by monotypy). -Patagonian Formation (Lower Miocene). Remarks.— The holotype is a very poor specimen for correct generic identifi- cation. 256 Annals of Carnegie Museum VOL. 65 prisca, 1907. Olivancillaria OLIVIDAE 1907«. AMNBA, 14:370, text fig. 15. 1914. NPRMP, 1(3): 108 (Forma^ao Entreriana). Type Locality.^ — Parana, Entre Rios. Holotype. — #873, one cast, coll. S. Roth (by monotypy). Stratigraphy. — Parana Eormation (Upper Miocene). Remarks. — This is Oliva reticularis brasiliana in Borchert (1901:47, pi. 4, fig. 8), not O. reticularis Lamarck, 1811, or O. brasiliana Lamarck, 1811. Similar to the Recent species Olivancillaria elongata (Eischer, 1807) from Brazil and Uru- guay. prisca, 1907. Struthiolaria STRUTHIOLARIIDAE 1907a. AMNBA, 14:28, pi. 3, fig. 18 (Cretacee Superieur). Type Locality. — Roca, Rio Negro. Type. — Not in the collection under this name. Stratigraphy. — Roca Formation (Paleocene). Remarks.- — The type material was stated to consist of “un moule externe mal conserve.” One specimen in the collection, #782 from Park- Aik, is not well pre- served and was listed by Ihering (1914:96); it may be this species. See Struthio- laria ornata “van” densestriata. Proscaphella, 1 907 . VOLUTID AE 1907a. AMNBA, 14:205. New genus. Type Species. — Valuta gracilior Ihering {= V. gracilis Philippi, 1887). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — = Miomelon Dali, 1907 (type species: Volutilithes philippianus Dali; = V. gracilis Philippi, 1887). pruniformiSy 1907. Marginella MARGINELLIDAE 1907a. AMNBA, 14:370. New name for Marginella aff. prunum in Borchert (1901:48, pi. 6, fig. 11), not Marginella prunum Swainson, 1822. Type Locality. — Parana, Entre Rios (Borchert). Stratigraphy. — Parana Formation (Upper Miocene). pseudopatagonicuy 1899. Cardita CARDITIDAE 1899a. NJMGP, 2:16. 1907a. AMNBA, 14:283. Venericardia patagonica (Sowerby). 1914. NPRMP, 1(3):44. Venericardia pseudopatagonica (Superpatagonico). = Venericardia patagonica (Sowerby). Type Locality. — La Cueva, Santa Cruz (in Ihering [1914] changed to Yegua Quemada). Syntypes.— ^365, two specimens, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species was based on juveniles of V. patagonica. pseudopatagonica, 1907. Corbula CORBULIDAE 1907a. AMNBA, 14:388. Not figured; new name for Corbula patagonica in Borchert (1901:44, pi. 4, fig. 31-32), not Corbula patagonica d’Orbigny, 1846. 1996 Parodiz^-Ihering Mollusca 257 Type Locfllilj. “-“Parana, Entre Rios (Borchert). Stratigraphy. — Parana Formation (Upper Miocene). Pseudotylostoma^ 1903, 1903. AMNBA, 9:207. New geous. NATICIDAE Type Species.— Pseudotylostoma romeroi Iheriiig. Stratigraphy.— Roca Formation (Paleocene). Remarks. See Pseudotylostoma romeroi. pueyrredona^ 1907. Calyptraea CREPIDULIDAE 1907a. AMNBA, 14:149. New name for Trochita araucana in Philippi (1887:87, pi. 11, fig. 1), not Trochita araucana Lesson, 1831. 1914. NPRMP, 1(3):84 (Patagoeico). Type Locfliily. -—Northwest Santa Cruz at Lake Pueyrredon. Syntypes.—4t666, two specimens, coll. Hatcher, received from Ortmann. Stratigraphy. — -Patagonian and Navidad formations (Feruglio, 1949-1950) (Lower Miocene). Remarks.- — This species was misidentified as Galerus araucanus by Ortmann (1902:181, pL 32, fig. 7a-b). Trochita araucana Lesson is a synonym of Calyp- traea radians Lamarck, In the same publication (1907£i:253) Ihering gave another new name, Calyptraea levuana, for the species misidentified as Trochita araucana by Philippi (1887). pueyrredonOf 1907. Leda NUCULANIDAE 1907a. AMNBA, 14:229. New name for Leda errazurizi ie Ortmann (1902:84, pL 26, fig. 3a--b), not Nucula errazurizi Philippi (1887:189, pL 41, fig. 11). 1914. NPRMP, 1(3): 14 (Formagao Patagonica). = Nucuiana pueyrredona (Ihering). Type Locality.— hago Pueyrredon, Santa Craz (Ortmann). Syntypes.—#H3, two valves received from Ortmann and reported as “cotypos” are in collection. Stratigraphy. — -Patagonian and Navidad formations (Lower Miocene). pueyrredona, 1907. Venericardia CARDITIDAE 1907a. AMNBA, 14:286. New name for Cardita volckmanni in Ortmann (1902:126, pi. 26, fig. 6), not Cardita volckmanni Philippi, 1887 (“Terciario antiguo”)- 1914. NPRMP, 1(3):45. Venericardia volckmanni (Philippi). = Venericardia volckmanni (Philippi). Type Locality.— Lago Pueyrredon, Santa Cruz, as Ihering referred to specimens listed in Ortmann (1902). The type locality for Cardita volckmanni Philippi is Tubul, Chile. Stratigraphy.— I'Navidad Formation (?Miocene). Remarks.— valve received from Philippi (Lot #372) is in the Ihering Col- lection under volckmanni, not under pueyrredona. puntana, 1907. Pecten patagonensis 18971?. RMP, 2:226. Pecten patagonensis d’Orbigny. 1907a. AMNBA, 14:257. Myochlamys patagonensis puntana. PECTINIDAE 258 Annals of Carnegie Museum VOL. 65 1914. NPRMP, 1(3):32 (Formagao Entreriana Patagonico). = Chlamys patagonensis (d’Orbigny, 1842). Type Locality. — Golfo San Jorge, Punta Nova (Nueva), Santa Cruz. Syntypes. — #267, 12 valves, coll. Ameghino, 1900. Stratigraphy.— P^idigonmn Formation (Lower Miocene). Remarks. — Curated in the Ihering Collection under Myochlamys patagonensis puntana. puntarenasensisy 1907. Polynices NATICIDAE 1907a. AMNBA, 14:341. New name for Natica chiloensis in Ortmann (1899:431) and Ortmann (1902, pi. 186, fig. la-b), not Natica chiloensis Philippi, 1887. 1914. NPRMP, 1(3):85 (Magellanico Inferior). Type Locality. — Punta Arenas, Chile (Ortmann). Syntypes. — #678, three specimens received from Ortmann and reported as “co- types'’ are in the collection. Stratigraphy. — Magellanian Formation (Lower Oligocene). puntasium, 1907. Calliostoma TROCHIDAE 1907a. AMNBA, 14:136, pi. 4, fig. lOa-c. 1914. NPRMP, 1(3):79 (Patagonico Medio). Type Locality. — Cabo Tres Puntas, Santa Cruz. Types. — Not in the collection. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Ihering compared C. puntasium to Calliostoma rioense Dali (= C. jucundum [Gould]), but considering the nature of the columella in the illustration it appears to belong in the subgenus Kombologion Clench and Turner, 1960. Two specimens were listed in the 1914 catalogue. pyramidesitty 1907. Turritella ambulacrum TURRITELLIDAE 1907a. AMNBA, 14:367. Not figured. 1914. NPRMP, 1(3):88 (Formagao Entreriana). = Turritella ambulacrum Sowerby, 1846. Type Locality. — Puerto Piramides, Chubut. Holotype. — #710, coll. Ameghino, 1903 (by monotypy). Other Lot. — #711, Trelew, Chubut, three specimens, coll. S. Roth. Stratigraphy. — Rio Negro Formation (Upper Miocene) and also Camacho For- mation in Uruguay which is coeval with the Parana and Rio Negro formations. Remarks. — Although this material is both allopatric and allochronic with nom- inotypical T. ambulacrum, it is not different and the name is placed in synonymy. Turritella ambulacrum Sowerby of southern range in Santa Cruz is represented in the collection by ten lots from different localities. See also T. steinmanni Iher- ing, T. argentina Ihering, and T. ambulacrum sylva Ihering. pyramidesiunty 1907. Pecten oblongus PECTINIDAE 1907a. AMNBA, 14:375. Not figured. 1914. NPRMP, 1(3):36. Pecten oblongus Bravard (Formagao Entreriana). = Chlamys oblongus pyramidesium (Ihering). Type Locality. — Puerto Piramides, Chubut. 1996 Parodiz — Ihering Mollusca 259 Types.— Not in the collection. Other Lot— #291, Pecten oblongus Bravard, three specimens, coll. S. Roth, 1903. Stratigraphy.— PdimnA and Rfo Negro formations (Upper Miocene). Ihering (1907) reports specimens from this locality collected by Ameghino in 1903, but none are in the collection under this name; only ""Pecten"" oblongus from Entre Rios. pyriformiSy 1897. Trophon MURICIDAE 1897L RMP, 2:295, pL 3, fig. 5 (Forma^ao Saetacrazense). 1907£i. AMNBA, 14:183, pi. 5, fig. 31. 1914. NPRMP, 1(3): 101 (Superpatagonico), Type Locality.— Ycgu^ Quemada, Santa Cruz. Syntypes.- — #'828, one specimen and one fragment (juveniles), colL Ameghino. Stratigraphy.— P^l^gonimi Formation (Lower Miocene). Remarks.- — These may belong in Trophon lamellosus (Gmelin). pymthermma, 1907. Polynices NATICIDAE 1907a. AMNBA, 14:154, pL 5, fig. 17. 1914. NPRMP, 1(3):85, pL 3, fig. 13 (Patagonico Inferior). = Lunatia pyrotheriana (Ihering), new combination. Type Locality. — Camarones, Chubut. Syntypes.—4t679, six large specimens and nine smaller, coll. Ameghino. Patagonian Formation (Lower Miocene), also known from the Magellanian Formation (Lower Oligocene) of Rio Turbio. pyrotheriorum, 1897. Ostrea GRYPHAEIDAE 1897L RMP, 2:315, text fig. 20 on p. 316 [there are two figures numbered 20, the first on page 315 is Potamides patagonensis]. 1902a. AMNBA, 4:116. Gryphaea (Salamaequeano). 1903. Tournouer, BSGF, 3:473. 1907a. AMNBA, 14:44. Gryphaea. 1914. NPRMP, 1(3):21 (Cretaceo Superior “salamanqu&n”). = Gryphaea pyrotheriorum (Ihering). Type Locality.- — ^Mamelones de Pinedo (also known as Tetas de Pinedo), Santa Cruz. Syntypes.—4tlA6, one complete specimen and one valve. Stratigraphy.— and Salamanca formations (Paleocene). Remarks.^ — ^The species was also mentioned by Tournouer (1903) from Colhue Huapi, and by Feraglio (1 949-1 950) from Punta Peligros and Puerto Viser. quadrisulcata, 1897. Perna ISOGNOMONIDAE 1897L RMP, 2:231, pi. 9, fig, 54 (Superior formationis patagonicae). 1902. Ortmane, p. 97, pi. 24, fig. 2a~b (Lake Pueyrredon). 1907a. AMNBA, 14:243. 1914. NPRMP, 1(3):20 (Superpatagonico). Type Locality.— La Cueva, Santa Cmz. Holotype.- — #134, single valve, coll. Ameghino (by monotypy). Other Lol— #135, Cabo Tres Puntas, two valves. 260 Annals of Carnegie Museum VOL. 65 Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Lot #135 was reported by Ihering (1914) as “typo"’ but it was collected by C. Ameghino in 1900, after the original description and from a different locality. This species is only tentatively placed in Perna. quemadensis, 1907. Fusus FASCIOLARIIDAE: FUSININAE 1907a. AMNBA, 1907 14:195, pi. 6, fig. 39. 1914. NPRMP, 1(3): 103 (Superpatagonico). = IFusinus quemadensis (Ihering). Type Locality. — Yegua Quemada, Santa Cruz. Holotype. — #850, coll. Ameghino (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Fusus of authors (= Fusinus Rafinesque, 1815); the canal in this species is very short. This species resembles Fusinus pilsbryi (Ortmann) which was originally described in the genus Fusus (Ortmann, 1900:375). quemadensis^ 1897. Glycimeris [sic] HIATELLIDAE 1897Z?. RMP, 2:264, text fig. 4 on p. 265 (Formagao Santacruzense). 1907a. AMNBA, 14:328. Panopaea [sic]. 1914. NPRMP, 1(3):63 (Superpatagonico). = Panopea quemadensis (Ihering). Type Locality. — Yegua Quemada, Santa Cruz. Holotype. — #519, a restored valve, coll. Ameghino (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This is a species close to the Recent Panopea abbreviata Valenci- ennes, 1839. quemadensis^ 1897. Marginella MARGINELLIDAE 1897^. RMP, 2:307, pi. 3, fig. 9, and pi. 4, fig. 14. 1907a. AMNBA, 14:199, pi. 6, fig. 40a-b. Neoimbricaria. 1914. NPRMP, 1(3): 107. Neoimbricaria '' guemadesia" (Patagonico). = Neoimbricaria quemadensis (Ihering). Type Locality. — Yegua Quemada, Santa Cruz. Holotype. — #870, coll. Ameghino (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). quemadensis, 1897. Pecten PECTINIDAE 1897# RMP, 2:228, pi. 6, fig. 38 (Formagao Santacruzense). 1907a. AMNBA, 14:256. Myochlamys. 1914. NPRMP, 1(3):33 (Superpatagonico). = Chlamys (Zygochlamys) quemadensis (Ihering). Type Locality. — Yegua Quemada, Santa Cruz. Syntypes. — #271, five valves, coll. Ameghino; #272, Las Cuevas, one specimen, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Lot #272 was reported as “typo” in Ihering (1914), but it was collected after the original description and from a different locality. See Morra (1985). 1996 Parodiz — -Ihering Mollusca 261 quemadensis^ 1897. Terebra costellata 'Var.” TEREBRIDAE 18971?. RMP, 2:311 (Formagao Santacrazense). 1902, Ortmane, p. 236, pi. 36, fig. 5. Terebra costellata Sowerby. 1907fl. AMNBA, 14:216. Terebra quemadensis. 1914. NPRMP, l(3):li2 (Seperpatagonico). = Terebra quemadensis Iherieg. Type Locality.- — -Yegua Quemada, Santa Cmz. Holotype.—4t9Ql, coll. Ameghino (by monotypy). 5'rrarigra|?/iy. -"-Patagonian Formation (Lower Miocene). Remarks.— ThQ type locality of Terebra costellata Sowerby is Navidad, Chile; T. quemadensis is a different species. See T. costellata santacruzensis Ihering. quemadensis^ 1896. Voluta VOLUTIDAE 1896, NDMG, 7:97. 18971?. RMP, 2:304, pi. 3, fig. 7 (Forma^ao Santacrazense). 1899fl. NJMGP, 2:34. 1907a. AMNBA, 14:207, Proscaphella. 1914. NPRMP, 1(3): 111. Cymbiola (Miomelon) (Superpatagonico). = Miomelon quemadensis (Ihering). Type Locality.— Ycgu^ Quemada, Santa Craz. liolotype.- — #892, coll. Ameghino (by monotypy). Other Lot.— #^93, Maeantial Salado, Santa Cruz, two incomplete specimens, reported as Cymbiola sp. aff, to quemadensis, coll. Ameghino. Patagonian Formation (Lower Miocene). Remarks.- — ^Iherieg (18971?) stated that this species may be synonymous with V. phiiippiana Dali, which is a synonym of Adelomelon graciiior (Ihering, 1896). querandina, 1907. Turbonilla PYRAMIDELLIDAE: TURBONILLINAE 1907a. AMNBA, 14:442, pL 17, fig. 121 (Pampeano Superior). = Turbonilla (Pyrgiscus) uruguayensis Pilsbry, 1897. Type Locality.— Bshm Blanca, formation pampienne superieur. Type.— Hoi in the collection. Stratigraphy.— Qncmndi Formation (Late Pleistocene). Recent in Uruguay and Buenos Aires Province. rada, 1907. Lotorium (Lampusia) RANELLIDAE: CYMATIINAE 1907a. AMNBA, 14:175, pi. 5, fig. 29a-"b (text states fig. 26 io error). 1914. NPRMP, 1(3):97 (Superpatagonico). = Semitriton rada (Ihering), new combination. Type Locality.— Rada Tilly, Santa Cruz. Syntypes.— #793, three specimens, coll. Ameghino, 1900. S'frarigrap/i};.— Patagonian Formation (Lower Miocene). Remarks.— This species resembles Semitriton dennanti (Tale) from the Eocene of Australia but has a shorter spire and stronger nodulous cords. rada, 1914. Marcia VENERIDAE: SAMARANGIINAE 1914. NPRMP, 1(3):53, pi. 1, fig. 5a-b (Superpatagonico). = ISamarangia exalbida (Chemnitz, 1795). 262 Annals of Carnegie Museum VOL. 65 Type Locality. — Rada Tilly, Santa Cruz. Holotype.—^43S, juvenile, coll. Ameghino (by monotypy). Other Lots.‘ — Unnumbered, Sierra de los Baguales, seven casts, coll. Hauthal. Stratigraphy. — ^Patagonian Formation (Lower Miocene); Recent. radana, 1907. Myochlamys geminata “van” PECTINIDAE 1907a. AMNBA, 14:255. Not figured. 1914. NPRMP, 1(3):35 (Patagonico Inferior). = Chlamys geminata (Sowerby, 1846). Type Locality. — Rada Tilly, Santa Cruz. Sy nty pe. --^2H7 , only one specimen reported as “cotypo” in the collection, C. Ameghino, 1900. Stratigraphy. — Patagonian Eormation (Lower Miocene). Remarks. — There was another specimen mentioned in the description from Piedra Clavada but it was not in the collection. Although there are no specimens of Chlamys geminata Sowerby in the collection, Ihering (1907a) mentioned the species had been collected at several localities: Bajo de San Julian, Manantial Salado, Pan de Azucar, Monte Espejo, Punta Nodales, Punta Nueva, and Cama- rones. Eeruglio (1949-1950) found the species geminata at all levels of the Pat- agonian, but listed the form radana with doubt. rionegrensis, 1903. Modiola MYTILIDAE 1903. AMNBA, 9:205. Not figured. 1904. RMLP, 11:233, pi. 1, fig. 5. 1907a. AMNBA, 14:38. Modiolus. 1914. NPRMP, 1(3):39 (Cretaceo Superior). Type Locality. — Roca, Rio Negro. Syntypes. — #319, two specimens, coll. S. Roth. Stratigraphy. — Roca Formation (Paleocene). Remarks. — -This is probably a Brachidontes Swainson. rionegrensiSy 1903. Ostrea OSTREIDAE 1903. AMNBA, 9:202, pi. 1, fig. 5a-b (text shows 5-6). 1907a. AMNBA, 14:14, pi. 2, fig. 7; see also p. 46 (Rocaneen). 1914. NPRMP, 1(3):22 (Cretaceo Superior). Type Locality. — Roca, Rio Negro. Syntypes. — #160, four valves, coll. S. Roth. Other Lots. — #161, Roca, two valves, coll. Ameghino, 1904; #162, Homo Viejo [sic] for Homo Viejo near Roca, ten valves, coll. Ameghino, 1904. Stratigraphy. — Roca Formation (Paleocene). Remarks. — The specimen figured in Ihering (1907a) differs from the original of Ihering (1903). In the Danian of Cerro Dorotea, in the Magellanian region, the species is characteristic (Feruglio, 1949-1950). rioplatensis^ 1907. Columbella COLUMBELLIDAE 1907a. AMNBA, 14:369. New name for Columbella acuta in Borchert (1901:50, pi. 5, fig. 3.), not Anachis acuta Stearns, 1873. = 1 Anachis moleculina (Duclos, 1840). 1996 Parodiz — Ihering Mollusca 263 Type Locality,— Rios (Borchert). Stratigraphy.— WmmiA Formation (Upper Miocene). Remarks.— On consultation with W. H. Dali, Ihering confirmed that C acuta Stearns was a synonym of C albella Dali and different from C rioplatensis. The name rioplatensis given by Ihering might refer to deep perforations in the zone of La Plata River (?Puente Alsina) 230 ft deep. rocama^ 1903. Cucullaea CUCULLAEIDAE 1903. AMNBA, 9:204, pL 1, fig. 7. 1907. NPRMP, 14:35 (Rocanfen). Type Locality.— Koca, Rio Negro. Type. — =Not in the collection. Stratigraphy.— Roca Formation (Paleocene). Remarks.— In the work of 1914, no Cucullaea were listed from the Paleocene of Roca as indicated in the original description. mcmm, 1903. Gryphaea GRYPHAEIDAE 1903. AMNBA, 9:198. Not figured. 1907«. AMNBA, 14:7, pi. 1, fig. 5a™b (Rocaneen), and p. 36. 1914. NPRMP, 1(3):20. Gryphaea rothi Bohm. = lOdonto gryphaea rothi (Bohm, 1903). Type Locality.- — Roca, Rio Negro. Stratigraphy.— Roca Formation (Paleocene). Remarks. — ^The description of G. rothi Bohm appeared a few weeks before G. rocana Ihering. Gryphaea rocana was based upon specimens figured by Burck^ hardt (1902:214, pL 3, fig. 1-3) as Gryphaea aff. pitcheri, not G. pitcheri Morton, 1834. rocana^ 1907. Phacoides LUCINIDAE 1907£I. AMNBA, 14:25, pi. 3, fig. 5 (Rocaneen). 1914. NPRMP, 1(3):46. = Lucina (Lucinisca) rocana (Ihering), new combination. Type Locality.— Roca, Rfo Negro. Lectotype.—#3H5, the only specimen in the collection of the three listed in the original description, coll. Ameghieo, 1904, here designated the lectotype. Stratigraphy. — -Roca Formation (Paleocene). Remarks.— Phacoides Blainville, 1825 (= Lucina Bruguiere, 1797). rocana, 1903. Ostrea OSTREIDAE 1902. Burckhardt, p. 214, pi. 2, fig. 11“-19. Ostrea aff. bomilcaris Coquand. 1903. AMNBA, 9:203. Ostrea rocana. Not figured. 1907a. AMNBA, 14:13. O. ameghinoi rocana. 1914. NPRMP, 1(3):23. O. ameghinoi rocana (Rocaneen). Type Locality.— Roca, Rio Negro. Holotype.—^l61 , colL Ameghino, 1901 (by monotypy). Other Lots.- — #168, Roca, eight valves, coll. S. Roth; #169, Neuquen, one well- preserved specimen; #170, Roca, one complete specimen, coll. Ameghino, 1904; #171, same five valves; #172, km 1216-1218, Rfo Negro, one valve. 264 Annals of Carnegie Museum VOL. 65 Stratigraphy.— Roca Formation (Paleocene). mcamumf 1907. Cardium (Hemicardium) CARDIIDAE 1907a. AMNBA, 14:25, pL 3, fig. 14a--b (Rocan&n). = Trachycardium rocanum (Ihering), new combination. Type Locality.— Rocsl, Rio Negro. Ty/je.— Not in the collection; the figured specimen is an internal cast. Stratigraphy.- — Roca Formation (Paleocene). mchai, 1907. Tylostoma NATICIDAE 1907a. AMNBA, 14:40, text fig. 2 (“Cretaceous”). = AmpulUna (Pseudotylostoma) rochai (Ihering). Type Locality.— Rio Grande do Norte, Brasil. Type.”— Not in the collection; the single described specimen was collected by Francisco da Rocha. Stratigraphy.— Rmhdhly Paleocene. mmeroif 1903. Nautilus NAUTILIDAE 1903. AMNBA, 9:191, pL 1, fig. 1 (Cretacee Superieur). 1907a. AMNBA, 14:41 (Rocaneen). 1914. NPRMP, 1(3):68 (Cretaceo Superior de Roca). Type Locality.- — Roca, Rfo Negro. Syntype.—4t5A3, labelled “cotypo,” coll. Ameghino, 1901. Stratigraphy.— Roc^ Formation (Paleocene). romemiy 1903. Pseudotylostoma NATICIDAE 1903. AMNBA, 9:207, pi. 1, fig. 8. 1907a. AMNBA, 14:39. 1914. NPRMP, 1(3):87. = AmpulUna (Pseudotylostoma) romeroi (Ihering). See Wenz, 1938:1028. Type Locality.— Rocdi, Rfo Negro. Syntype.— #693, one cast labelled “cotypo,” coll. Ameghino, 1904[!]. Stratigraphy.— Roca Formation (Paleocene). rfo. —Although the label for the syntype states the specimen was from C. Ameghino, 1904, the catalogue (Ihering, 19 1 A) names Colonel A. Romero as the collector. mstrigerUy 1902. Gryphaea concors “var.” GRYPHAEIDAE 1902a. AMNBA, 4:113. Not figured. 1903. AMNBA, 9:211, pi. 2, fig. 10. 1907a. AMNBA, 14:43. Gryphaea rostrigera (Salamanqueano). 1945. Stenzel, BGSA, 56(12, pt, 2): 1202. Odontogryphaea. = Odontogryphaea rostrigera (Ihering). Type Locality.— of Rio Chico, Chubut. Syntypes.—#IA1 , two complete specimens and six valves. Other Lots.— ^14^, Puerto Malaspiea, Chubut, one complete specimen and ten valves. 1996 Parodiz — Ihering Mollusca 265 Strati graphy.SdlMndLnc2L Formation (Paleocene). Remarks. — -This species is the type species of the genus Odontogryphaea Iher- ing. rothif 1904. Diplodon HYRIIDAE 1904. RMLP, 11:232, pi. 1, fig. 6. 1907«. AMNBA, 14:466. 1914. NPRMP, 1(3):37 (“Terciario antiguo”). 1969. Parodiz, ACM, 40:69, pi. 7, fig. 4. Type Locality. — Arroyo Lele, N. W. Chubut. Syntypes. — #299, not in collection; the lectotype is deposited at CMNH (see Remarks). Stratigraphy. — Colloncura beds (Simpson, 1933; Femglio, 1949-1950), = Fri- asean Formation (Kraglievich, 1930). Upper Miocene. Remarks.— T\i& lectotype was designated by Parodiz (1969) from a specimen in Lot #299 in the Ihering Collection received in exchange from Dr. Max Biraben, at that time Director of the Museo Argentino de Ciencias Maturates. The species is also found at Arroyo Lepa given as “type,” coll. Hauthal. rothi^ 1903. Rostellaria STROMBIDAE 1903. AMNBA, 9:204. 1904. RMLP, 11:12, pi. 2, fig. 9-10. 1907^1. AMNBA, 14:28. 1914. NPRMP, 1(3):94 (Salamanqueano). = Tibia rothi (Ihering). Type Locality. — Roca, Rio Negro. Syntypes. — #760, nine casts, coll. S. Roth; #761, a “Variety,” three casts, coll. Ameghino-S. Roth. Stratigraphy. — Roca Formation (Paleocene). Remarks.- — See note under Rostellaria patagonensis. There are two lots in the collection under Rostellaria “sp.” (undetermined species), #762 consisting of two casts from Valle Alsina, Chubut, coll. Ameghino, and #4233, a single cast from Roca, coll. S. Roth. Salamanca, 1903. Chlamys PECTINIDAE 1903. AMNBA, 9:214, pi. 1, fig. 11. 1907(3. AMNBA, 14:47. Myochlamys (Salamanqueano). Type Locality. — Pico Salamanca, Santa Cruz. Syntypes. — #286, one complete specimen, seven valves, coll. Ameghino, 1900. Stratigraphy.— -Sdildirndincdi Eormation (Paleocene). Remarks.— This, species appears to be ancestral to Chlamys patagonicus (d’Orbigny). Femglio (1949-1950) reports it from the Valley of the Rio Chico. salobHs, 1907. Cyrena CORBICULIDAE 1907a. AMNBA, 14:413, pi. 16, fig. 109 (Formagao Araucana). 1914. NPRMP, 1(3):41. 1969. Parodiz, ACM, 40:95, fig. 14-15. Neocorbicula. = Cyanocyclas salobris (Ihering), new combination. Type Locality. — Sierra Laziar, Santa Cmz. 266 Annals of Carnegie Museum VOL. 65 Type Material. — #336, lectotype and one paralectotype, coll. Ameghino; sub- sequent designation by Parodiz, 1969. Stratigraphy. — Araucanean Formation (Lower Pliocene). Remarks. — Cyanocyclas Blainville, 1818, and Neocorbicula Fischer, 1887, are subjective synonyms. This synonymy was not recognized correctly in Parodiz and Hennings (1965). I prefer to treat Cyanocyclas as a genus distinct from Corbicula because species of the former have a pallial sinus, longer siphons, and are re- stricted to the Neotropical region. Living species of Cyanocyclas are viviparous. santacruzensCf 1914. Cardium CARDIIDAE 1914. NPRMP, 1(3);49, pi. 1, fig. 4a-b. = Laevicardium santacruzense (Ihering), new combination. Type Locality. — Yegua Quemada, Santa Cruz. Holotype. — #411, a broken valve, coll. Ameghino (by monotypy). Stratigraphy.- — Patagonia Formation (Lower Miocene). santacruzensis, 1907. Chenopus APORRHAIDAE 1907a. AMNBA, 14:170. New name for Aporrhais araucana in Ortmann (1902:200, pi. 33, fig. 9), not Aporrhais araucana Philippi, 1887, from the Tertiary of Chile. = Drepanocheilus santacruzensis (Ihering), new combination. Type Locality. — Santa Cruz (Ortmann). Stratigraphy.^ — Patagonian Formation (Lower Miocene). Remarks.— Thi^ species differs from D. araucanus (Philippi) by having axial instead of spiral sculpture. santacruzensis, 1907. Pleurotoma TURRIDAE 1907a. AMNBA, 14:219, pi. 7, fig. 46. 1914. NPRMP, 1(3):113 (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Holotype. — #907, incomplete specimen, coll. J. Bicego, 1897 (by monotypy). Other Lot. — #908, Canada de los Artilleros, one specimen, coll. Ameghino. Stratigraphy.— Formation (Lower Miocene). santacruzensis, 1897. Polynices NATICIDAE 1897Z?. RMP, 2:280. Natica solida Sowerby, 1846 (not N. solida Blainville, 1825). 1907a. AMNBA, 14:152, pi. 5, fig. 5. Polynices santacruzensis. 1914. NPRMP, 1(3):86 (Superpatagonico). Type Locality.— CsiheidB. de los Artilleros, Santa Cruz. Syntypes. — #680, seven specimens. Other Lots. — #681, Yegua Quemada, coll. Ameghino; #682, La Cueva, 12 spec- imens, coll. Ameghino. Stratigraphy.— P^ldigonmn Formation (Lower Miocene). Remarks. — The species has affinities with Polynices lacteus (Guilding, 1830) which is Recent and ranges from North Carolina to Brazil. See note on Natica tumbeziana Ihering. santacruzensis, 1899. Tellina 1899a. NJMGP, 2:22, pi. 2, fig. 5. 1907a. AMNBA, 14:314. Macoma. TELLINIDAE: MACOMINAE 1996 Parodiz — Ihering Mollusca 267 1914. NPRMP, 1(3):58. Macoma (Patagonico). = Macoma santacruzensis (Ihering). Type Locality. Cruz. Syntypes. — #483, four casts, coll. J. Bicego, 1897. Stratigraphy.— Formation (Lower Miocene). santacruzensis, 1897. Terebra costellata “van” TEREBRIDAE 1897^. RMP, 2:311. Terebra costellata Sow. ?“var.” 1907<3. AMNBA, 14:216. Terebra quemadensis santacruzensis. 1914. NPRMP, 1(3): 112. Terebra quemadensis santacruzensis (Superpatagonico). = Terebra santacruzensis Ihering. Type Locality.— Ldi Cueva, Santa Cruz. Holotype. — #902, coll. Ameghino, 1895 (by monotypy). Other Lot.— #903, Canada de los Artilleros, three specimens. Stratigraphy. — Patagonian Formation (Lower Miocene). santacruzensis, 1897. Trophon laciniatus MURICIDAE 1897^. RMP, 2:294, pi. 3, fig. 4. 1907a. AMNBA, 14:182. 1904. RMLP, 11:230. Trophon laciniatus. 1914. NPRMP, 1(3): 100 (Patagonico Inferior-Medio). = Trophon (Austrotrophon) lamellosus (Gmelin, 1791). Type Locality. — La Cueva, Santa Cruz. Holotype.— #%l\, coll. Ameghino, 1895 (by monotypy). Other Lots. — #812, Yegua Quemada, two specimens, coll. Ameghino; #814, Manantial Salado, four specimens, coll. Ameghino; #815, Pan de Azucar, four specimens, coll. Ameghino; #816, Owen Point, four specimens, received from Ortmann; #817, Canada de los Artilleros, three specimens, coll. Ameghino; #818, Rada Tilly, one specimen, coll. Ameghino; #819, Punta Nodales, one specimen, coll. Ameghino; #820, Punta Casamayor, one specimen, coll. Ameghino; #822, Corral Foyel, one cast, coll. Hauthal. Stratigraphy.— Pdiidigonmn Formation (Lower Miocene). saxicava, 1907. Barnea PHOLADIDAE 1907a. AMNBA, 14:325, pi. 14, fig. 96. 1914. NPRMP, 1(3):63 (Patagonico Inferior). Type Locality. — Bahia Camarones, Chubut. Type. — Not in the collection. Stratigraphy.— PditZigonmn Formation (Lower Miocene). Remarks. — This species was based on an unique specimen listed in Ihering (1914), and because of the unsatisfactory illustration the species is of questionable status. schucherti, 1914. Gibbula ARCHITECTONICIDAE 1914. NPRMP, 1(3):74, pi. 3, fig. lOa-b (Patagonico Inferior). = Architectonica schucherti (Ihering), new combination. Type Locality. — Manantial Salado, Santa Cruz. Holotype. — #856, coll. Ameghino, 1900 (by monotypy). 268 Annals of Carnegie Museum VOL. 65 Other Lots. — #587, Cabo Ties Puntas, three specimens, coll. Ameghino, 1900; #588, Rada Tilly, one cast, coll. Ameghino, 1900; #589, Sierra de los Baguales, two casts, coll. HauthaL Stratigraphy.— Formation (Lower Miocene). scutata, 1907. Marcia VENERIDAE: SAMARANGIINAE 1907a. AMNBA, 14:303, pL 11, fig. 76. 1914. NPRMP, 1(3):52 (Patagonico). == Samarangia scutata (Ihering). Type Locality.— Funta Casamayor, Santa Cruz. Holotype.—WJ31 , coll. Ameghino (by monotypy). Other Lot.— #435, Santa Cruz, eight casts, colL Bicego, 1897. Stratigraphy.— Fdildigoniwi Formation (Lower Miocene). .^emarfo.“-“This species is close to the Recent Samarangia exalbida (Chemnitz, 1795). See also '"Marcia"' scutata tehuelcha Ihering. sehuemUf 1907. Corbula 7CORBICULIDAE 1907a. AMNBA, 14:51, text fig. 5 (Formation “Guaranitica”)- 1914. NPRMP, 1(3):62 (Cretaceo Superior). 1949-1950. Feruglio 1:258, 273, 274. Eriphyla. 1969. Parodiz, ACM, 40:101. Eriphyia (Paleocene). = Eriphyia sehuena (Ihering). Type Locality.— MditdL Amarilla on the Sehuen River (also called Chalia), Santa Cruz (delimited by Parodiz, 1969). Lectotype.— 41509, coil. Ihering, deposited at CMNH (see Remarks). Stratigraphy.— Sthucndin beds (Paleocene). Remarks.— A lectotype was designated by Parodiz (1969) from Lot #509 based on specimens from the Ihering Collection received in exchange from Dr. Max Biraben, at that time Director of the Museo Argentino de Ciencias Naturales. Eriphyia Gabb, 1864, is known from the Mesozoic of the Archiplata of Bolivia and Ecuador, and also from the Tertiary in the Puca Formation (Paleocene). sierranttf 1907. Panopaea [sic] HIATELLIDAE 1902. Ortmann, p. 154, pi. 30, fig. 2. Panopaea [sic] quemadensis Ihering. 1907a. AMNBA, 14:328, pL 13, fig. 38. 1914. NPRMP, 1(3):65 (Patagonico). = Panopeal quemadensis (Ihering). Type Locality.— de los Baguales, Santa Cruz. Holotype.— 4520, one cast, coll. Hauthal (by monotypy). Stratigraphy.— Fa.ta.gonmn Formation (Lower Miocene). Remarks.— The type cast is only an upper portion not showing the characteristic surface that Ihering mentioned in the original description. Sphenaturia^ 1921. NAUTILIDAE 1921. Physis, 5:76. New subgenus of Aturia Bronn. Type Species.- — Aturia (Sphenaturia) brueggeni Ihering, here designated. Stratigraphy.— de Boqueroe” (below Patagonian Formation). Described as a subgenus of Aturia Bronn, 1838, without original selection of a type species. 1996 Parodiz— Ihering Mollusca 269 splendida, 1897. Cytherea VENERIDAE 1897^. RMP, 2:255, pi. 6, fig, 42 (Formagao Santacrazense). 1907a. AMNBA, 14:302. 1914. NPRMP, 1(3):51. Macrocallista iheringi Cossmann (Superpatagonico). = Macrocallista iheringi (Cossmann, 1895) (= Amiantis Carpenter, originally Meretrix Lamarck) . Type Locality. Quemada, Santa Cruz. Syntypes.--~^4-23, 11 valves, one separated with a label: “C. explendida [sic] = C. iheringi Cossmann.” Strarigrap/zy.— Patagonian Formation (Lower Miocene). steinmanni, 1897. Turritella TURRITELLIDAE 1897^. RMP, 2:289. Not figured. 1907a. AMNBA, 14:162. Turritella ambulacrum steinmanni. 1914. NPRMP, 1(3):88. Turritella ambulacrum steinmanni (Patagonico and Superpatagonico). = Turritella ambulacrum Sowerby, 1846. Type Locality.- — Santa Cruz. Holotype. — #712, coll. Ameghino (by monotypy). Other Lots. — #713, Casamayor, three specimens, coll. Ameghino; #714, Mam antial Salado, five specimens, coll. Ameghino; #715, Camarones, 40 specimens, coll. Ameghino. Strafigrap/iy.— Patagonian Formation (Lower Miocene). Remarks. — Turritella ambulacrum argentina Ihering is also a synonym. Ihering (1897^:332) referred to T. cf. steinmanni from the Parana Formation, which in Ihering (I907<2) was included in the synonymy of T. americana Bravard, a com- mon species of the Upper Miocene. striatissima, 1903. Rostellaria STRUTHIOLARIIDAE 1903. AMNBA, 9:219, pi. 2, fig. 16. 1907a. AMNBA, 14:49. Aporrhais striatissima. 1914. NPRMP, 1(3):94 (Salamanqueano). = Struthiolaria striatissima (Ihering). Type Locality.-— Rio Chico, Chubut. Syntypes. — #768, two specimens, coll. Ameghino. Stratigraphy. — Salamanca Formation (Paleocene). Remarks. — Other Rostellaria in the Ihering Collection correspond by priority to Tibia, but striatissima has the characteristics of a Struthiolaria. striatolamellata, 1897. Venus VENERIDAE 1897/?. RMP, 2:253, pi. 7, fig. 44 (Formagao Santacruzense). 1902. Ortmann, p. 140, pi. 28, fig. 4. Venus darwini Ortmann. 1907a. AMNBA, 14:305. Marcia. 1914. NPRMP, 1(3):53. Marcia (Superpatagonico). = Samarangial exalbida (Chemnitz, 1795). Type Locality. Quemada, Santa Cruz. Syntypes. — #437, four valves, coll. Ameghino. Stratigraphy. — ^Patagonian Formation (Lower Miocene); also Recent. Remarks. — This species resembles Venus navidaensis Philippi from the Tertiary 270 Annals of Carnegie Museum VOL. 65 of Chile. '"Marcia'' pupilla Feruglio, 1935, from the Salamanca Formation is also very similar to striatolamellata. stromeriy 1914. Fusus RANELLIDAE: RANELLINAE 1914. NPRMP, 1(3):104. Not figured (Patagonico Medio). = Fusitriton stromeri (Ihering), new combination. Type Locality. — Cabo Tres Puntas, Santa Cruz. Syntypes.^ — #851, two specimens, coll. Ameghino; #852, southeast of Punta Nueva, one specimen, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.— Ihering (1907^?: 194) assigned the specimens from Cabo Tres Puntas to Fusus archimedis Ortmann (1900:374). Struthiolaria sp. STRUTHIOLARIIDAE 1914. NPRMP, 1(3):96. Locality. — Pio [Piso] Sehuen (Cretaceo superior). Specimens. — #782, from Park-Aik, Santa Cruz, coll. Ameghino, Stratigraphy. — Sehuenan Formation (Paleocene). Remarks. — The “piso” Sehuen (Sehuenan) is a fresh-brackish-water strati- graphic unit of the Paleocene. The specimen, not well preserved, must be from debris of the Salamanca; it was not reported by Ihering (1907a), and probably is S. prisca Ihering. subrecta, 1899. Siphonalia dilatata FASCIOLARIIDAE: FUSININAE 1899a. NJMGP, 2:30. 1902. Ortmann, p. 211, pi. 34, fig. 4. Siphonalia domeykoana (Philippi). 1907a. AMNBA, 14:192. Siphonalia subrecta. 1914. NPRMP, 1(3): 102 (Patagonico Medio-Super). = Fusinus subrectum (Ihering). Type Locality. — Santa Cruz, Syntypes. — #842, five specimens, coll. J. Bicego, 1895. Other Lots. — #843, Santa Cruz, one specimen received from Ortmann. #844, Cabo Tres Puntas, five specimens, coll. Ameghino; #845, Canada de los Artilleros, six specimens, coll. Ameghino. Stratigraphy.— P^tdLgonmn Formation (Lower Miocene). Remarks. — Siphonalia nodocincta Ihering and S. matthewi Ihering are syn- onyms of this name. subtenuis, 1897. Natica NATICIDAE 1897Z?. RMP, 2:284, text fig. 13 (Forma^ao Santacruzense). 1907a. AMNBA, 14:157. Polynices. 1914. NPRMP, 1(3):86 (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Holotype. -#692, coll. Ameghino, 1895 (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species has affinities with P. ortmanni Ihering. It was reported by Wilckens (1911) from the Tertiary of Seymour Island, Antarctica, and may 1996 Parodiz — Ihering Mollusca 271 belong in Magnatica Marwick (1924:545), a genus described from the Miocene of New Zealand. subtrigona^ 1907. Glycimeris [sic] GLYCYMERIDIDAE 1907a. AMNBA, 14:241, pi. 7, fig. 53a-b (in text as fig. 23). 1914. NPRMP, 1(3): 19 (Patagonico Inferior). = Glycymeris subtrigona Ihering. Type Locality.— CdLmdiXon&s, Chubut. Syntypes.—^130, 32 valves, coll. Ameghino, 1900. Stratigraphy. — -Patagonian Formation (Lower Miocene). Remarks. — Glycymeris Da Costa, 1778, was considered a nomina conservanda by the ICZN (Opinion 1414). suburbana^ 1907. Crassatellites CRASSATELLIDAE 1907a. AMNBA, 14:381, pi. 14, fig. lOOa-b. = Crassatella suburbana (Ihering). Type Locality. — Puente Alsina, a suburb south of Buenos Aires city, from a perforation, 230 ft. Type.- — Not in the collection. Stratigraphy. — Parana Formation (Upper Miocene). Remarks. — The only valve was described in great detail. There was no indi- cation of collector (probably Ameghino). subventricosa, 1897. Eulima EULIMIDAE 1897/?. RMP, 2:274, text fig. 9. 1907a. AMNBA, 14:142. 1914. NPRMP, 1(3):81 (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Syntypes. — #632, two specimens, coll. Ameghino. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — This species is a large (20 mm in length) lEulima, a rarity among the Patagonian gastropods, fossil or Recent. sulcolunulariSy 1907. Venericardia CARDITIDAE 1907a. AMNBA, 14:285, pi. 10, fig. 70a-b. 1914. NPRMP, 1(3):45 (Patagonico Medio). Type Locality. — Bajo de las Flechas, Santa Cruz. Holotype.—^361 , coll. Ameghino, 1900 (by monotypy). Other Lots. — #366, Golfo San Jorge, one complete specimen and four valves, coll. Ameghino, 1900; #368, Golfo San Jorge, three valves; #369, Pan de Azucar, two valves. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — In the 1914 catalogue a lot from Golfo San Jorge was listed as “Typo.” This species resembles Venericardia cossmanni Ihering. suturalis, 1897. Odostomia 1897/?. RMP, 2:275, text fig. 19 (Formagao Santacruzense). 1902. Ortmann, p. 173, pi. 33, fig. 7a-b. 7PYRAMIDELLIDAE 272 Annals of Carnegie Museum VOL. 65 1907a. AMNBA, 14:143, pi. 4, fig. 13. 1914. NPRMP, 1(3): 81 (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Holotype.— #633, one specimen, coll. Ameghino (by monotypy). Stratigraphy.— -FatagonisLU Formation (Lower Miocene). Remarks.— The type specimen is the one figured in Ihering (1907a). The orig- inal figure of 1897^ is much different and must be an error. The acute spire and long “goniostome-like” aperture are unlike any Recent species. sylva^ 1907. Turritella ambulacrum TURRITELLIDAE 1907a. AMNBA, 14:342. Not figured. 1914. NPRMP, 1(3):89 (Magellanico). = Turritella ambulacrum Sowerby, 1846. Type Locality. — Carmen Sylva, Chile. Syntypes.—#1\6, two specimens, received from C. Backhausen. Stratigraphy. — ^Magellanian Formation (Lower Oligocene). Remarks. — Only two specimens remain of the five mentioned in the original description. See Turritella ambulacrum argentina Ihering, T a. steinmanni Iher- ing, and T. a. pyr amide sia Ihering. sylva, 1907. Venericardia CARDITIDAE 1907a. AMNBA, 14:346, pi. 14, fig. 98a-b (Magellanico). Type Locality. — Carmen Sylva, Chile. Type. — Not in the collection. Stratigraphy. — Magellanian Formation (Lower Oligocene). tehuelcha, 1907. Cucullaea CUCULLAEIDAE 1907a. AMNBA, 14:5, pi. 1, fig. 3a-b (Rocaneen). Type Locality. — ^Roca, Rio Negro. Type. — Not in the collection. Stratigraphy. — ^Roca Formation (Paleocene). Remarks.— This species was based on a cast and the generic assignment is uncertain. The locality indicates the Roca Formation, but there are no Cucullaea from the Paleocene in the Ihering Collection. tehuelcha, 1914. Marcia scutata VENERIDAE: SAMARANGIINAE 1914. NPRMP, 1(3):53. Not figured; title of original description states ""subsp. but description says “variedade” (Formagao Araucana). = ISamarangia exalbida (Chemnitz, 1795). Type Loca//ry.— Between San Jorge and Deseado, Santa Cruz. Holotype.- — #436, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Araucanean Formation (Pliocene); Recent. tehuelcha, 1907. Psammobia PSAMMOBIIDAE 1907a. AMNBA, 14:417, pi. 17, fig. 112a-b. 1914. NPRMP, 1(3):58 (Formagao Araucana). = Gari tehuelcha (Ihering). 1996 Parodiz — Ihering Mollusca 273 Type Locality. — Sierra Laziar, Santa Cruz. Syntypes.—TtA^O, three valves, coll. Ameghino, 1900. Stratigraphy. — Araucanean Formation (Lower Pliocene). Remarks.-— Psammobia Lamarck, 1817, is a synonym of Gari Schumacher, 1817. tehuelchtty 1907. Pupilia aperta FISSURELLIDAE 1907£i. AMNBA, 14:399. Not figured. 1914, NPRMP, 1(3):72 (Formagao Araucana). = Fissurellidea megatrema d’Orbigny, 1841. Type Locality.— Si&rxdL Laziar, Santa Cruz. Holotype.— #566, coll. Ameghino, 1900 (by monotypy). Stratigraphy. — Araucanean Eormation (Lower Pliocene); also Recent. Remarks. — The species described by Sowerby (1892) as Pupilia aperta was from South Africa. tehuelcha, 1899. Tellina PSAMMOBIIDAE 1899a. NJMGP, 2:21, pi. 2, fig. 4. 1907a. AMNBA, 14:313. Sanguinolaria. 1914. NPRMP, 1(3):58. Sanguinolaria (Patagonico). = Sanguinolaria tehuelcha (Ihering). Type Locality. — Santa Cruz. Holotype. — #482, cast with pieces of the original valve, coll. Ameghino, 1900 (by monotypy). Stratigraphy, — Patagonian Formation (Lower Miocene). tehuelchana^ 1907. Venericardia CARDITIDAE 1902. Ameghino, ASCA, 54:190. Cardita tehuelchana Ihering, in litteris. 1907a. AMNBA, 14:412, pi. 16, fig. 107a-b. Venericardia. 1914. NPRMP, 1(3):45 (Formagao Araucana). Type Locality. — Sierra Laziar, Santa Cruz. Syntypes.— #37 1 , 16 valves, coll. Ameghino. Stratigraphy. — Araucanean Formation (Lower Pliocene). tehuelchuntf 1907. Calliostoma TROCHIDAE 1907a. AMNBA, 14:437, pi. 17, fig. 115a-b. 1914. NPRMP, 1(3):74 (Pampeano). = Calliostoma consimilis Smith, 1881. Recent. Type Locality. — Puerto Deseado, Santa Cruz. Holotype.— #615, coll. Ameghino, 1907 (by monotypy). Stratigraphy. — Deseado beds, corresponds to the Belgrano Formation of the north (Pleistocene). Remarks. — The original description indicates a unique specimen, but Ihering (1914) list two specimens in error. tenuis, 1907. Corbicula 1907a. AMNBA, 14:468, pi. 18, fig. 13c. 1914. NPRMP, 1(3):41 (Formagao Entreriana). CORBICULIDAE 274 Annals of Carnegie Museum VOL. 65 1920. Frenguelli, 24:108. 1969. Parodiz, ACM, 40:98, pi. 10, fig. 12, 13. Neocorbicula tenuis (Ihering). Type Locality. — Parana, Entre Rios, Syntypes. — #337, four valves, labelled as “cotypos.” Other Lot. — #338, La Paz, Entre Rios, conglomerate with numerous impres- sions of valves, coll. S. Roth. Stratigraphy.— VdiVdinA Formation (Upper Miocene). Remarks. — Frenguelli (1920) said that he found “thousands” of badly pre- served shells in the deposits at Parana; these were probably Cyanocyclas limosa (Maton), as Corbicula tenuis is very rare in collections. Ihering said that the specimens were found in a mass of clay from the Bravard collection that also contained Diplodon fraus Ihering. Corbicula tenuis Ihering is the type species of the subgenus Corbiculella Ihering, 1907. thomasi, 1914. Panopaea [sic] HIATELLIDAE 1914. NPRMP, 1(3):63, pi. 2, fig. 8a-b (Cretaceo Superior). = Panopea thomasi Ihering. Type Locality. — Rio Chico, Chubut. Type Material — #474, two casts, coll. Ameghino. The figured specimen, 27 mm long, is here designated the lectotype. Stratigraphy. — Salamanca Formation (Paleocene). trespunta^ 1907. Cerithiopsis CERITHIOPSIDAE 1907«. AMNBA, 14:168, text fig. 6. 1908. Cossmann, p. 104. Colina trespunta (Ihering). 1914. NPRMP, 1(3):93 (Patagonico Medio). Type Locality. — Cabo Tres Puntas, Santa Cruz. Holotype. — #751, coll. Ameghino (by monotypy). Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks. — Cerithiopsis emersoni (C. B. Adams, 1839), Recent, is a very sim- ilar species occurring from Massachusetts to Brazil. tricesima, 1897. Nucula NUCULIDAE 1897^. RMP, 2:243, pi. 4, fig. 21, and pi. 5, fig. 27. 1907a. AMNBA, 14:226 (Superpatagonico). 1914. NPRMP, 1(3): 13 (Superpatagonico). Type Locality. — La Cueva, Santa Cruz. Lectotype. — #75, coll. Ameghino, 1895. The original description reports three specimens, broken; only one remains in the collection, and is here designated the lectotype. Stratigraphy. — Patagonian Formation (Lower Miocene). tricinctUf 1897. Turritella TURRITELLIDAE 1897/?. RMP, 2:pl. 3, fig. 3 (Formagao Santacruzense). 1902. Ortmann, p. 135, pi. 31, fig. 14a-b. Turritella breantiana d’Orbigny. 1907a. AMNBA, 14:165, pi. 5, fig. 24. Turritella iheringi Cossmann. 1914. NPRMP, 1(3):90. Turritella iheringi Cossmann (Superpatagonico). = ^Turritella breantiana d’Orbigny, 1847. 1996 Parodiz— Ihering Mollusca 275 Type Locality.— YQg\x2i Quemada, Santa Cruz. Syntypes.—#132, ten specimens, coll. Ameghino. Other Lots.--^733, La Cueva, one specimen, coll. Ameghino; #734, Canada de los Artilleros, 18 specimens, coll, Ameghino. Stratigraphy. — ^Patagonian Formation (Lower Miocene). Remarks. — There are no specimens in the collection under Turritella tricincta and all material of that species are labelled T. iheringi as listed in the catalogue (Ihering, 1914:90-91). tridentata^ 1897. Cuculearia [sic] ARCIDAE 1897Z?. RMP, 2:237, pi. 4, fig. 22, and pi. 5, fig. 28. 1907a. AMNBA, 14:233. Cucullaria. 1914. NPRMP, 1(3): 15. = Barbatia tridentata (Ihering), new combination. Type Locality. — Yegua Quemada, Santa Cruz. Syntypes. — #98, two broken valves, coll. Ameghino, 1895. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.— ThQ original description reports “an unique specimen.” ''Cucu- learia'' was an error for Cucullaria. tumbeziana, 1907. Natica NATICIDAE 1907a. AMNBA, 14:153. New name for Natica darwini in Philippi (1887:80, pi. 9, fig. 28), not Natica darwini Hutton, 1886 (the latter in turn a new name for Natica solida Sowerby [1846]), not Natica solida Blainville (1825). Type Locality. — “Eormation Cretacee, Chile” (Philippi, 1887). Stratigraphy. — Probably the Navidad Eormation (Miocene). Remarks. — In the collection there is a single cast from Lebu, Chile, received from R. A. Philippi as "Polynices" darwini Hutton, but no specimens labelled as N. tumbeziana. Ihering (1907a: 89) said that "Polynices Carolodarwini Ih. [’est substitue dans la Nouvelle-Zelande] par P. darwini Hutton.” I have not found any previous reference to Polynices carolodarwini. unifascialiSf 1897. Pleurotoma discors “van” TURRIDAE 1897^. RMP, 2:312. Discussed under Pleurotoma discors Sowerby. 1902. Ortmann, p. 239, pi. 36, fig. 7a-b. Pleurotoma discors “Var.” 1907a. AMNBA, 14:219, pi. 7, fig. 45. Pleurotoma unifascialis. 1914. NPRMP, 1(3):113 (Superpatagonico). Type Locality. — Yegua Quemada, Santa Cruz. Holotype. — #909, coll. Ameghino (by monotypy). Stratigraphy. — Patagonian Eormation (Lower Miocene). Remarks. — This species probably belongs in Bathystoma. Pleurotoma Link, 1807 (= Turris Roding, 1798). usurpator^ 1907. Lotorium (Lampusia) RANELLIDAE: CYMATIINAE 1907a. AMNBA, 14:175, pi. 5, fig. 28a-b. 1914. NPRMP, 1(3):98 (Superpatagonico). = Cymatium usurpator (Ihering), new combination. Type Locality.— KziddL Tilly, Santa Cruz. 276 Annals of Carnegie Museum VOL. 65 Syntypes. — #794, two specimens, coll. Ameghino, 1900. Stratigraphy.— ~V2LldLgonmn Formation (Lower Miocene). Remarks.- — Cymatium usurpator (Ihering) and C. rada (Ihering) are both from the same locality and strata, but they can be distinguished by their shape and sculpture. valchetanuSy 1907. Pododesmus ANOMIIDAE 1907a. AMNBA, 14:267, pi. 10, fig. 64a-b. 1914. NPRMP, 1(3):27 (Patagonico). Type Locality. — “Travesia” Valcheta, Gualicho River, Rio Negro. Syntypes.- — #216, seven upper valves (12 were listed in the original descrip- tion), coll. C. Burmeister, 1887. Stratigraphy. — Patagonian Formation (Lower Miocene). vindex, 1907. Chione VENERIDAE 1907fl. AMNBA, 14:416, pi. 16, fig. lla-b. 1914. NPRMP, 1(3):55 (Formacion Araucana). = Protothaca antiqua (King). Type Locality. — Sierra Laziar, Santa Cruz. Holotype. — #454, one broken valve, coll. Ameghino (by monotypy). Stratigraphy. — Araucanean Eormation (Lower Pliocene); Recent. waltheri “Ihering,” 1931. Planorbis PLANORBIDAE 1931. Walther, BIGU, 13:192. Name credited to Ihering in litteris. 1958. Caorsi and Goni, BIGU, 37:58. 1969. Parodiz, ACM, 40:165, pi. 18, fig. 1. Taphius waited (emended as walthen). = Taphius waltheri Parodiz (= Biomphalaria waltheri [Parodiz]). Remarks. — Describing Taphius walteri [sic] (= waltheri, Parodiz, 1969), I used the name which was a nomen nudum in Walther (1931) because it was frequently mentioned as such in the Uruguayan geological literature. Taphius Adams and Adams, 1855, is the most abundant genus of South American planorbids; the name was arbitrarily synonymized by the ICZN under the African Biomphalaria Preston, 1910, on the grounds that the two genera are anatomically the same, but disregarding the 50-year rule for conservation of names. The type locality for Taphius walteri [sic] Parodiz, 1969, is Palmitas, Soriano, southeastern Uruguay, from the Fray Bentos Formation (Miocene). The holotype in CMNH is a specimen collected by A. Berro in 1927. whitei, 1903. Pseudotylo stoma NATICIDAE 1903. AMNBA, 9:307. New name for Tylostoma torrubiae in White, 1887, not T. torrubiae Sharpe, 1849. = Ampullina {Pseudotylo stoma) whitei (Ihering). Type Locality. — Sergipe, Brazil. Stratigraphy. — ? “Upper Cretaceous. ’ ’ wilckensi, 1907. Ostrea 1907a. AMNBA, 14:13, pi. 2, fig. lOa-b, and pi. 3, fig. 10c. 1914. NPRMP, 1(3):23 (“cretaceo superior horizonte medio”). OSTREIDAE 1996 Parodiz — Ihering Mollusca 277 Syntypes.— ^163, two valves. Stratigraphy.~~R.ocd^ Formation (Paleocene). Zygochlamys, 1907. PECTINIDAE 1907«. AMNBA, 14:250. New subgenus of Chlamys Roding. Type Species. — Chlamys geminata Sowerby. Stratigraphy. — Patagonian Formation (Lower Miocene). Remarks.— This subgenus includes C. fissocostatus Ihering, C. nodosoplicatus Ihering, C. quemadertsis Ihering, and C. jorgensis Ihering. For a revision of Zyg- ochlamys, see Morra (1985). Synonymic List of Fossil Taxa Described by Ihering All names proposed by Ihering are organized below in a synonymic list fol- lowing the classification of Vaught (1989). The placement of some fossil taxa follows Wenz (1938). Names of the species group and of the genus group are alphabetically arranged under higher categories (family level and above). Original combinations of names proposed by Ihering are in boldface; all synonyms are indented below the name considered valid. In some cases, combinations are un- known or tentative, and this uncertainty is indicated by placing the genus-group name within quotation marks (e.g., ''Gibbula""). PHYLUM ANNELIDA Class Polychaeta ^^Vermetus"^’ julianus Ihering, 1907 PHYLUM MOLLUSCA Class Gastropoda Subclass Prosobranchia Order Archaeogastropoda Superfamily Pleurotomarioidea Family Raphistomatidae “Gibbula^^ philippii Ihering, 1907 Superfamily Fissurelloidea Family Fissurellidae Fissurellidea megatrema d’Orbigny, 1841 Pupilia aperta tehuelcha Ihering, 1907 Superfamily Patelloidea Family Fissurellidae Cellana lucifera (Ihering, 1904) Helcioniscus luciferus Ihering, 1904 Helcioniscus luciferus patagonicus Ihering, 1904 Superfamily Trochoidea Family Trochidae Calliostoma camaronesium Ihering, 1907 Calliostoma consimilis Smith, 1881 Calliostoma tehuelchum Ihering, 1907 Calliostoma militaris Ihering, 1907 Calliostoma amazonicum Finlay, 1930 278 Annals of Carnegie Museum VOL. 65 Calliostoma bellicosum Ihering, 1907 Calliostoma dalii Ihering, 1907 Calliostoma quequensis Carcelles, 1944 Calliostoma jucundum (Gould, 1849) Calliostoma lahillei Ihering, 1907 Calliostoma laziarium Ihering, 1907 Calliostoma puntasium Ihering, 1907 ^^Gibbula^’ osborni Ihering, 1914 Tegula patagonica (d’Orbigny, 1840) Neomphalius princeps Ihering, 1907 Tegula abava (Ihering, 1907), new combination Neomphalius abavus Ihering, 1907 Tegula (Promartynia) americana (Ihering, 1907), new combina- tion Neomphalius americanus Ihering, 1907 Family Turbinidae ‘‘Calliostoma^^ deseadense Ihering, 1914 Order Mesogastropoda Superfamily Cerithioidea Family Potamididae Potamides patagonensis Ihering, 1897 Family Pleuroceridae Paleoanculosa bullia (Ihering, 1907) Melania bullia Ihering, 1907 Melania pehuenchensis Doello-Jurado, 1927 Melania ameghiniana Doello-Jurado, 1927 Family Turritellidae Turritella ambulacrum Sowerby, 1846 Turritella ambulacrum sylva Ihering, 1907 Turritella ambulacrum pyramidesia Ihering, 1907 Turritella argentina Ihering, 1897 Turritella steinmanni Ihering, 1897 Turritella ameghinoi Ihering, 1903 Turritella breantiana d’Orbigny, 1847 Turritella breantiana ‘‘var.’^ camaronesia Ihering, 1907 Turritella breantiana “var.” indecussata Ihering, 1899 Turritella iheringi Cossmann, 1898 Turritella tricincta Ihering, 1897 Turritella doeringi Bohm, 1903 Turritella burckhardti Ihering, 1903 Turritella cahuillensis Ihering, 1907 Turritella hauthali Ihering, 1907 Turritella malaspina Ihering, 1903 Family Siliquariidae Siliquaria cossmanni (Ihering, 1907) Tenagodus cossmanni Ihering, 1907 1996 Parodiz- — Ihering Mollusca 279 Family Vermetidae Dendropoma multicoronatus (Ihering, 1907), new combination Vermetus multicoronatus Ihering, 1907 Petaloconchus cMcoanus (Ihering, 1907) Vermetus cMcoanus Ihering, 1907 Superfamily Stromboidea Family Stmthiolariidae StrutMolaria amegMnoi Ihering, 1897 Struthiolaria chilemis Philippi, 1887 StrutMolaria cMlensis fuegina Ihering, 1907 Struthiolaria ornata Sowerby (1846) StrutMolaria ornata “var.” densestriata Ihering, 1897 StrutMolaria prisca Ihering, 1907 StrutMolaria sp., Ihering, 1914 Struthiolaria striatissima (Ihering, 1903) Rostellaria striatissima Ihering, 1903 Family Aporrhaidae Rostellaria chubutensis Ihering, 1903 Aporrhais cossmanni (Ihering, 1903) Rostellaria cossmanni Ihering, 1903 Drepariocheilus santacruzensis (Ihering, 1907), new combination Chenopus santacruzensis Ihering, 1907 Family Strombidae Tibia patagonensis (Ihering, 1903) Rostellaria patagonensis Ihering, 1903 Aporrhais rocai Bohm, 1903, nomen nudum Tibia rothi (Ihering, 1903) Rostellaria rotM Ihering, 1903 Superfamily Crepiduloidea Family Crepidulidae Calyptraea pileus Lamarck, 1822 Calyptraea pileus “var.” jmIiii#1£i Ihering, 1907 Calyptraea pueyrredona Ihering, 1907 Calyptraea levuana Ihering, 1907 [junior objective synonym] Superfamily Naticoidea Family Naticidae Ampullina {Pseudotylostoma) Ihering, 1903 Pseudotylostoma Ihering, 1903 Ampullina {Pseudotylostoma) rochai (Ihering, 1907) Tylostoma rochai Ihering, 1907 Ampullina {Pseudotylostoma) romeroi (Ihering, 1903) Pseudotylostoma romeroi Ihering, 1903 Ampullina {Pseudotylostoma) whitei (Ihering, 1903) Pseudotylostoma whitei Ihering, 1903 Lunatia mageilanica Hombron and Jacquinot, 1834 Natica consimilis Ihering, 1897 Lunatia pyrotheriana (Ihering, 1907), new combination Polynices pyrotheriana Ihering, 1907 280 Annals of Carnegie Museum VOL. 65 Polynices carolodarwini Ihering, 1907 [nomen nudum] Polynices famula (Philippi, 1887) Polynices Jorgensis Ihering, 1907 Polynices ortmanni Ihering, 1907 Polynices puntarenasensis Ihering, 1907 Polynices santacruzensis Ihering, 1897 Natica subtenuis Ihering, 1897 Natica tumbeziana Ihering, 1907 Superfamily Tonnoidea Family Ranellidae Cymatium bicegoi (Ihering, 1899) Lotorium bicegoi Ihering, 1899 Cymatium (Cabestana) felipponei (Ihering, 1907) [emendation] Lotorium filipponei [sic] Ihering, 1907 Cymatium usurpator (Ihering, 1907), new combination Lotorium {Lampusia) usurpator Ihering, 1907 Fusitriton dautzembergi (Ihering, 1897), new combination Triton {Argobuccinum) dautzembergi Ihering, 1897 Fusitriton stromeri (Ihering, 1914), new combination Fusus stromeri Ihering, 1914 Semitriton rada (Ihering, 1907), new combination Lotorium {Lampusia) rada Ihering, 1907 Superfamily Cerithiopsoidea Family Cerithiopsidae Cerithiopsis Juliana Ihering, 1907 Cerithiopsis trespunta Ihering, 1907 Superfamily Epitonioidea Family Epitoniidae Epitonium chubutianum (Ihering, 1907), new combination Scalaria chubutiana Ihering, 1907 Opalia (Nodiscala) quemadensis (Cossmann), new combination Scalaria cossmanni Ihering, 1907 Opalia {Nodoscala) rugulosa (Sowerby, 1846) Scalaria rugulosa paucicostata Ihering, 1907 Superfamily Eulimoidea Family Eulimidae Eulima subventricosa Ihering, 1897 Order Neogastropoda Superfamily Muricoidea Family Muricidae Crassilabrum jorgensis (Ihering, 1907), new combination Hadriania jorgensis Ihering, 1907 Crassilabrum hatcheri (Ortmann, 1900) Hadriania lacustris Ihering, 1907 Ocenebra {Ocinebrina) elegans (Ortmann, 1900), new combina- tion Urosalpinx archipatagonica Ihering, 1907 Urosalpinx elegans Juliana Ihering, 1907 1996 Parodiz — Ihering Mollusca 281 Thais (Stramonita) Schumacher, 1817 Trophon (Entacanthus) Ihering, 1907 Thais {Stramonita) monoceros (Ihering, 1907), new combination Trophon monoceros Ihering, 1907 Trophon chicoanus (Ihering, 1907), new combination Hadriania jorgensis ^‘var.” chicoana Ihering, 1907 Trophon inornatus ‘^var.” gradata Ihering, 1907 [nomen nudum} Trophon lamellosus (Gmelin, 1791) Trophon inornatus fairus Ihering, 1907 Trophon laciniatus santacruzensis Ihering, 1897 Trophon necocheanus Ihering, 1907 Trophon pyriformis Ihering, 1897 Trophon varians (d’Orbigny, 1841) Trophon varians ‘‘var.” gradata Ihering, 1897 Trophon varians montenus Ihering, 1907 Urosalpinx ortmanni Ihering, 1907 Urosalpinx ''pleurotomoides'’ Cossmann, 1903 Family Buccinidae Cominella (Austrocominella) Ihering, 1907 Cominella fueguensis Ihering, 1907 Cominella minor (Philippi, 1887) Cominella cossmanni Ihering, 1907 Cominella golfonia Ihering, 1914 Cominella sp., Ihering, 1914 [unidentifiable] Gibbula iheringi Cossmann, 1899 Gibbula iheringi “var.” golfina Ihering, 1907 Tritonidea borcherti Ihering, 1907 Family Columbellidae Columbella (Anachis) improvisa Ihering, 1914 Anachis moleculina (Duclos, 1840) Columbella rioplatensis Ihering, 1907 Family Nassariidae Buccinanops gradatum (Deshayes, 1844) Bullia gradata pampeana Ihering, 1907 Family Fasciolariidae Fusinus quemadensis (Ihering, 1907) Fusus quemadensis Ihering, 1907 Fusinus subrectum (Ihering, 1899) Siphonalia dilatata subrecta Ihering, 1899 Siphonalia nodocincta Ihering, 1907 Siphonalia matthewi Ihering, 1914 Family Volutidae Adelomelon ameghinoi (Ihering, 1896), new combination Voluta ameghinoi Ihering, 1896 Adelomelon burmeisteri (Ihering, 1907), new combination Cymbiola burmeisteri Ihering, 1907 Adelomelon cannada (Ihering, 1907), new combination 282 Annals of Carnegie Museum VOL. 65 Cymbiola becki ^‘var.” cannada Ihering, 1907 Adelomelon cossmanni (Ihering, 1907) Proscaphella cossmanni Ihering, 1907 Adelomelon ferugUoi (Doello-Jurado, 1931) Cymbiola (Pachy cymbiola) sp., Ihering, 1914 Adelomelon (Pachycymbiola) pilsbryi (Ihering, 1899) Voluta pilsbryi Ihering, 1899 Enaeta dalli (Ihering, 1907), new combination Lyria dalli Ihering, 1907 Miomelon Dali, 1907 Proscaphella Ihering, 1907 Miomelon gracilior (Ihering, 1896) Voluta gracilior Ihering, 1896 Miomelon quemadensis (Ihering, 1896) Voluta quemadensis Ihering, 1896 Family Olividae Olivancillaria prisca Ihering, 1907 Family Marginellidae Marginella {sensu lato) deuterolivella (Ihering, 1907) Volvaria deuterolivella Ihering, 1907 Neoimbricaria quemadensis (Ihering, 1897) Marginella confinis Ihering, 1897 Marginella gracilior Ihering, 1897 Marginella quemadensis Ihering, 1897 Neoimbricaria quemadensis “var.” jM/iawa Ihering, 1907 Marginella pruniformis Ihering, 1907 Marginella sp., Ihering, 1914 [unidentifiable] Marginella {Volvaria) sp., Ihering, 1914 [unidentifiable] Neoimbricaria patagonica (Ihering, 1897 Voluta patagonica Ihering, 1897 Neoimbricaria plicifera (Ihering, 1897) Marginella plicifera Ihering, 1897 Family Costellariidae [Vexillidae] Vexillum patagonicus (Ihering, 1907) Vulpecula patagonica Ihering, 1907 Superfamily Cancellarioidea Family Cancellariidae Admete ameghinoi (Ihering, 1897) Cancellaria ameghinoi Ihering, 1897 Cancellaria gracilis Ihering, 1897 Cancellaria gracilis “var.” observatoria Ihering, 1907 Naronia {Sveltia) major (Ihering, 1899), new combination, new status Cancellaria gracilis “var.’’ major Ihering, 1899 Superfamily Conoidea Family Conidae Conus clenchi Martins, 1943 1996 Parodiz — Ihering Mollusca 283 Conus sp.5 Ihering, 1907 Conus carcellesi Martins, 1945 Family Turridae Genota cuevemis Ihering, 1897 Genotia cruzensis Ihering, 1907 Pleurotoma discors unifascialis Ihering, 1897 Pleurotoma jorgensis Ihering, 1907 Pleurotoma santacruzensis Ihering, 1907 Family Terebridae Terebra quemadensis Ihering, 1897 Terebra costellata ‘‘var.” quemadensis Ihering, 1897 Terebra santacruzensis Ihering, 1897 Terebra costellata ‘‘var.” santacruzensis Ihering, 1897 Subclass Heterobranchia Superfamily Architectonicoidea Family Architectonicidae Architectonica schucherti (Ihering, 1914), new combination Gibbula schucherti Ihering, 1914 Valdesia dalli (Ihering, 1897) Gibbula cuevensis ‘‘var.” aequistriata Ihering, 1907 Gibbula dalli Ihering, 1897 Gibbula dubiosa Ihering, 1907 Gibbula fracta ^‘var.” cuevensis Ihering, 1897 Valdesia fr acta (Ihering, 1897) Gibbula fracta Ihering, 1897 Superfamily Pyramidelloidea Family Pyramidellidae Odostomia suturalis Ihering, 1897 Turbonilla clessini Ihering, 1907 Turbonilla iheringi Clessin, 1900:174 [primary junior hom- onym] Turbonilla cuevensis Ihering, 1897 Turbonilla iheringi Cossmann, 1899 Turbonilla eopatagonica Ihering, 1907 [unnecessary replace- ment name] Turbonilla iheringi Clessin, 1900:168 [primary junior homonym without replacement name] Turbonilla observationis Ihering, 1907 Turbonilla uruguayensis Pilsbry, 1897 Turbonilla querandina Ihering, 1907 Subclass Opisthobranchia Order Cephalaspidea Superfamily Philinoidea Family Actaeonidae Actaeon argentinus Ihering, 1907 Actaeon patagonicus Ihering, 1907 284 Annals of Carnegie Museum VOL. 65 Family Ringiculidae Ringicula magellanica Iherieg, 1907 Family Scaphandridae Cylichna Juliana (Ihering, 1907) Bullimella Juliana Ihering, 1907 Tornatina camaronesia Ihering, 1907 Tornatina candei d’Orbigny, 1841 Tornatina militaris Ihering, 1907 Family Bullidae Bulla arenmia Ihering, 1907 Bulla pachycMla Ihering, 1907 Family Retusidae Retusa patagonica (Ihering, 1897), new combination Bulla patagonica Ihering, 1897 Bulla patagonica jorgensis Ihering, 1907 Volvulella cannada Ihering, 1907 Subclass Pulmonata Order Basommatophora Superfamily Siphonarioidea Family Siphonariidae Siphonaria lessoni (Blainville, 1826) Siphonaria lessoni pampa Ihering, 1914 Superfamily Planorboidea Family Planorbidae Biomphalaria waltheri (Parodiz, 1969) Planorbis waltheri “Ihering,” ex Walther, 1931 Taphius walteri [sic] Parodiz, 1969 [incorrect original spelling] Taphius waltheri Parodiz, 1969 [emendation] Order Stylommatophora Suborder Sigmurethra Superfamily Orthalicoidea Family Orthalicidae Peronaeus (Lissoacme) ameghinoi (Ihering, 1908) Bulimulm ameghinoi Ihering, 1908 Bulimulus sporadicus pampa Ihering, 1914 Peronaeus {Lissoacme) ameghinoi madrynensis Parodiz, 1944 Superfamily Strophocheiloidea Family Strophocheilidae Strophocheilus charruanus (Frenguelli, 1930) Bulimus archiplata “Ihering” ex Walther, 1931 {nomen nu- dum} Strophocheilus chubutensis Ihering, 1904 Family Megalobulimidae Megalobulimus hauthali (Ihering, 1904) Strophocheilus hauthali Ihering, 1904 Strophocheilus avus Parodiz, 1949 1996 Parodiz — ^Ihering Mollusca 285 Class Cephalopoda Subclass Nautiloidea Order Nautilida Family Nautilidae Aturia camloameghimoi Ihering, 1902 Aturia (Sphenaturia) Ihering, 1921 Aturia {Sphenaturia) brueggeni Ihering, 1921 Aturia (Sphenaturia) briiggeni [sic] Ihering, 1921 Aturia (Sphenaturia) felscM Ihering, 1921 Nautilus romeroi Ihering, 1903 Class Bivalvia Subclass Protobranchia Order Nuculoida Superfamily Nuculoidea Family Nuculidae Nucula dynastes Ihering, 1907 Nucula observatoria Ihering, 1907 Nucula tricesima Ihering, 1897 Superfamily Nuculanoidea Family Nuculanidae Nuculana camaronesia (Ihering, 1907) Leda camaronesia Ihering, 1907 Nuculana entreriana (Ihering, 1907) Leda entreriana Ihering, 1907 Nuculana ortmanni (Ihering, 1907) Leda ortmanni Ihering, 1907 Nuculana pueyrredona (Ihering, 1907) Leda pueyrredona Ihering, 1907 Subclass Pteriomorphia Order Arcoida Superfamily Arcoidea Family Arcidae Anadara camaronesia (Ihering, 1907), new combination Area- camaronesia Ihering, 1907 Anadara Juliana (Ihering, 1907), new combination Area Juliana Ihering, 1907 Area panensis Ihering, 1914 Area umbonata Lamarck, 1819 Area patagonica Ihering, 1897 Barbatia (Cucullaria) ameghinoi (Ihering, 1907), new combina- tion Cucullaria ameghinoi Ihering, 1907 Barbatia tridentata (Ihering, 1897), new combination Cuculearia [sic] tridentata Ihering, 1897 Noetia ameghinorum (Ihering, 1907) Venericardia ameghinorum Ihering, 1907 Family Cucullaeidae Cucullaea alta Sowerby, 1846 286 Annals of Carnegie Museum VOL. 65 Cucullaea dalli Ihering, 1897 Cucullaea multicostata Ihering, 1897 Cucullaea rocana Ihering, 1903 Cucullaea tehuelcha Ihering, 1907 Superfamily Limopsoidea Family Glycymerididae Glycymeris camaronesia Ihering, 1907 Glycimeris [sic] camaronesia Ihering, 1907 Glycymeris cuevensis (Ihering, 1897) Pectunculus pulvinatus “var.” cuevensis Ihering, 1897 Glycymeris subtrigona Ihering, 1907 Glycimeris [sic] subtrigona Ihering, 1907 Order Mytiloida Superfamily Mytiloidea Family Mytilidae Crenella camaronesia (Ihering, 1907) Crenella divaricata camaronesia Ihering, 1907 Chloromya hauthali (Ihering, 1907) Mytilus chorus “var.” hauthali Ihering, 1907 Lithophaga patagonica (d’Orbigny, 1846) Lithodomus patagonica [sic] dalli Ihering, 1907 ^^Modiola'' rionegrensis Ihering, 1903 Modiolus ameghinoi (Ihering, 1897) Modiola ameghinoi Ihering, 1897 Neoinoceramus ameghinoi Ihering, 1902 Order Pterioida Suborder Pteriina Superfamily Pterioidea Family Isognomonidae Perna quadrisulcata Ihering, 1897 Suborder Pinnina Superfamily Pinnoidea Family Pinnidae Atrina magellanica (Ihering, 1899) Pinna semicostata ^^var.” magellanica Ihering, 1899 Order Limoida Superfamily Limoidea Family Limidae Lima cossmanni Ihering, 1907 Lima patagonica Ihering, 1907 Lima pisum Ihering, 1907 Order Ostreoida Suborder Ostreina Superfamily Ostreoidea Family Ostreidae Exogyra callophyla Ihering, 1903 Exogyra guaranitica (Ihering, 1899) 1996 Parodiz — Ihering Mollusca 287 Ostrea guaranitica Ihering, 1899 Exogyra mendozana Ihering, 1907 Exogyra ostracina mendozana Ihering, 1907 Ostrea ameghinoi Ihering, 1902 Ostrea clarae Ihering, 1907 Ostrea faira Ihering, 1907 Ostrea hatcheri Ortmann, 1897 Ostrea percrassa Ihering, 1897 Ostrea madryna Ihering, 1907 Ostrea neuquena Ihering, 1907 Ostrea hemisphaerica [sic] “var.” capa Ihering, 1907 Ostrea hemisphaerica [sic] “var.” paca Ihering, 1907 Ostrea orbignyi Ihering, 1897 Ostrea rionegrensis Ihering, 1903 Ostrea rocana Ihering, 1903 Ostrea wilckensi Ihering, 1907 Family Gryphaeidae Gryphaea concors Ihering, 1902 Gryphaea pyrotheriorum (Ihering, 1897) Ostrea pyrotheriorum Ihering, 1897 Odonto gryphaea Ihering, 1903 Gryphaea (Odontogryphaea) Ihering, 1903 Odonto gryphaea rostrigera (Ihering, 1902) Gryphaea concors ^^var.” rostrigera Ihering, 1902 Odontogryphaea rothi (Bohm, 1903) Gryphaea rocana Ihering, 1903 Suborder Pectinina Superfamily Pectinoidea Family Pectinidae Chlamys Roding, 1798 Myochlamys Ihering, 1907 [unnecessary replacement name; preoccupied] Chlamys {Zygochlamys) Ihering, 1907 Chlamys centralis Sowerby, 1846 Myochlamys centralis “var.” ameghinoi Ihering, 1907 Chlamys deseadensis (Ihering, 1907) Myochlamys deseadensis Ihering, 1907 Chlamys eupatagonicus (Ihering, 1907) Myochlamys eupatagonica Ihering, 1907 Chlamys fissocostalis (Ihering, 1899) Pecten fissocostalis Ihering, 1899 Chlamys foyela (Ihering, 1914) Myochlamys foyela Ihering, 1914 Chlamys geminata (Sowerby, 1846) Myochlamys geminata “var/’ radana Ihering, 1907 Chlamys jorgensis (Ihering, 1907) Pecten jorgensis Ihering, 1907 Chlamys Juliana (Ihering, 1907) Myochlamys Juliana Ihering, 1907 288 Annals of Carnegie Museum VOL. 65 Chlamys laziarina (Ihering, 1907) MyocMamys laziarina Ihering, 1907 Chlamys nodosopUcatus (Ihering, 1897) Pecten nodosopUcatus [sic] Ihering^ 1897 [incorrect original spelling] Pecten nodosopUcatus Ihering, 1897 Chlamys patagonensis (d’Orbigny, 1842) Pecten patagonensis puntana Ihering, 1907 Chlamys patagonensis ^‘var.” negroina Ihering, 1907 Chlamys patagonensis portenia (Ihering, 1907) MyocMamys patagonensis portenia Ihering, 1907 Chlamys praenuncius (Ihering, 1897) Chlamys espejoana ^ ^Ihering” ex Ameghino, 1902 [nomen nu- dum] Pecten praenuncius Ihering, 1897 Chlamys oblongus pyramidesium (Ihering, 1907) Pecten oblongus pyramidesium Ihering, 1907 Chlamys quemadensis (Ihering, 1897) Pecten quemadensis Ihering, 1897 Chlamys salamanca Ihering, 1903 Pecten piconus Ihering, 1907 Family Propeamussiidae Amusium cossmanni Ihering, 1907 Amussium [sic] cossmanni Ihering, 1907 Superfamily Anomioidea Family Anomiidae Pododesmus Juliensis Ihering, 1907 Pododesmus patagonensis Ihering, 1907 Pododesmus paucisquamatus Ihering, 1907 Pododesmus pazensis Ihering, 1914 Pododesmus valchetanus Ihering, 1907 Subclass Paleoheterodonta Order Unionoida Superfamily Unionoidea Family Hyriidae Diplodon colhuapensis Ihering, 1903 Fossula derbyi (Ihering, 1907) Diplodon derbyi Ihering, 1907 Diplodon contortus (Borchert, 1901), new combination Diplodon fraus Ihering, 1907 [unnecessary replacement name] Diplodon delodontus (Lamarck, 1819) Diplodon charruanus lujanensis Ihering, 1907 Diplodon rothi Ihering, 1904 Subclass Heterodonta Order Veneroida Superfamily Lucinoidea Family Lucinidae Fimbria sp., Ihering, 1897 [unidentifiable] 1996 Parodiz — Ihering Mollusca 289 Lucina cruzialis (Ihering, 1907) Phacoides promaucana “var.” cruzialis Ihering, 1907 Lucina dalli (Ihering, 1907) Phacoides dalli Ihering, 1907 Lucina ortmanni (Ihering, 1899) Lucina ortmanni Ihering, 1899 Lucina rocana (Ihering, 1907), new combination Phacoides rocana Ihering, 1907 Miltha iheringiana Doello-Jurado (1919) Phacoides sp., Ihering, 1914 Family Ungulinidae Diplodonta vilardeboaena (d’Orbigny, 1846) Diplodonta villardeboaena [sic] “var.” camaronesia Ihering, 1907 Superfamily Carditoidea Family Carditidae Cardita burckhardti Ihering, 1903 Cardita palaeopatagonica Ihering, 1903 Carditamera plata (Ihering, 1907) Cardita plata Ihering, 1907 Venericardia camaronesia Ihering, 1907 Venericardia cannada Ihering, 1907 Venericardia dalli Ihering, 1907 Venericardia iheringi Bohm, 1903 Venericardia iheringi “var.^’ boehmi Ihering, 1907 Venericardia patagonica (Sowerby, 1846) Cardita pseudopatagonica Ihering, 1899 Venericardia sulcolunularis Ihering, 1907 Venericardia sulcolunularis freti Ihering, 1907 Venericardia sylva Ihering, 1907 Venericardia tehuelchana Ihering, 1907 Venericardia volckmanni (Philippi, 1887) Venericardia pueyrredona Ihering, 1907 Superfamily Crassatelloidea Family Crassatellidae Crassinella dalli Ihering, 1907 Crassatella kokeni Ihering, 1899 Crassatellites patagonicus Ihering, 1907 Crassatella longior Ihering, 1897 Crassatella suburbana (Ihering, 1907) Crassatellites suburbana Ihering, 1907 Superfamily Cardioidea Family Cardiidae Cardium ameghinoi Ihering, 1907 Cardium huttoni Ihering, 1907 Cardium patagonicum Ihering, 1907 Cardium philippii Ihering, 1897 Cardium philippii pauciradiata Ihering, 1899 290 Annals of Carnegie Museum VOL. 65 Cardium jorgensis Ihering, 1907, new status Cardium pisum jorgensis Ihering, 1907 Laevicardium santacruzense (Ihering, 1914), new combination Cardium santacruzense Ihering, 1914 Lahiilia angulata (Philippi, 1887) Lahillia angulata patagonica Ihering, 1907 Tr achy cardium rocanum (Ihering, 1907), new combination Cardium (Hemicardium) rocanum Ihering, 1907 Superfamily Mactroidea Family Mactridae Mactra patagonica d’Orbigny, 1846 Mactra indistincta Ihering, 1897 Mactra jorgea Ihering, 1914 Family Mesodesmatidae Lutraria ameghinoi Ihering, 1907 Superfamily Solenoidea Family Solenidae Solen crucis Ihering, 1907 Superfamily Tellinoidea Family Tellinidae Macoma santacruzensis (Ihering, 1899) Tellina santacruzensis Ihering, 1899 Tellina burmeisteri Ihering, 1907 Tellina gibber Ihering, 1907 Tellina Jeguaensis Ihering, 1897 Tellina patagonica Ihering, 1897 Tellina patagonica ^^var.” camaronesia Ihering, 1907 Family Semelidae Abra uruguayensis (Pilsbry, 1879) Abra patagonica Ihering, 1907 Family Psammobiidae Gari burmeisteri (Ihering, 1907) Psarnrnobia burmeisteri Ihering, 1907 Gari guassu (Ihering, 1907) Psammobia guassus [sic] Ihering, 1907 Gari tehuelcha (Ihering, 1907) Psammobia tehuelcha Ihering, 1907 Psammotella perplana (Ihering, 1897) Tellina perplana Ihering, 1897 Sanguinolaria tehuelcha (Ihering, 1899) Tellina ortmanni Ihering, 1907 Tellina tehuelcha Ihering, 1899 Tagelus plebeius (Solander, 1823) Tagelus gibbus entrerianus Ihering, 1907 Solecurtus platensis d’Orbigny, 1846 1996 Parodiz — Ihering Mollusca 291 Superfamily Gaimardioidea Family Gaimardiidae Gaimardia prisca (Ihering, 1907) Modiolarca prisca Ihering, 1907 Superfamily Corbiculoidea Family Corbiculidae Eriphyla sehuena (Ihering, 1907) Corbula sehuena Ihering, 1907 Corbicula (Corbiculella) Ihering, 1907 Corbicula {Corbiculella) tenuis Ihering, 1907 Corbicula tenuis Ihering, 1907 Cyanocyclas salobris (Ihering, 1907) Cyrena salobris Ihering, 1907 Superfamily Veneroidea Family Veeeridae Amiantis arenophila (Ihering, 1907) Macrocallista arenophila Ihering, 1907 Amiantis laziarina Ihering, 1907 Anomalocardia entreriana Ihering, 1907 Chitme eupyga Ihering, 1907 Chione pampeana Ihering, 1907 ^^Cytherea^^ chalcedonica Ihering, 1903 Dosinia burckhardti Ihering, 1907 Dosinia (Dosinella) burmeisteri (Ihering, 1907) Dosinia burmeisteri Ihering, 1907 Dosinia meridionalis Ihering, 1897 Dosinia entreriana Ihering, 1907 Macrocallista iheringi (Cossmann, 1895) Cytherea spiendida Ihering, 1897 Pitar julianum Ihering, 1907 Pitar lahillei Ihering, 1907 Pitar rostrata (Koch, 1844) Pitar lahillei ortmanni Ihering, 1907 Protothaca Dali, 1902 Chione (Ameghinomya} Ihering, 1907 Protothaca antiqua (King, 1832) Chione casa Ihering, 1907 Chione vindex Ihering, 1907 Protothaca argentina (Ihering, 1897) Venus volckmanni ^‘¥ar.” argentina Ihering, 1897 Venus paranensis Borchert, 1901 Protothaca cossmanni (Ihering, 1907) Chione cossmanni Ihering, 1907 Samarangia navidadis (Philippi, 1887) Marcia ortmanni Ihering, 1907 Samarangia exalbida (Chemnitz, 1795) Venus strialolamellata Ihering, 1897 Marcia rada Ihering, 1914 Marcia scutata tehuelcha Ihering, 1914 292 Annals of Carnegie Museum VOL. 65 Samarangia scutata (Ihering, 1907) Marcia scutata Ihering, 1907 Order Myoida Suborder Myina Superfamily Myoidea Family Corbulidae Corbula hatcheri Ortmann, 1900 Corbula pulchella Juliana Ihering, 1907 Corbula pseudopatagonica Ihering, 1907 Superfamily Hiatelloidea Family Hiatellidae Panopea bagualesia Ihering, 1907 Panopaea [sic] bagualesia Ihering, 1907 Panopea nucleus (Ihering, 1899) Glycimeris [sic] nucleus Ihering, 1899 Panopaea [sic] nucleus elongata Ihering, 1914 Panopea panis Ihering, 1907 Panopaea [sic] panis Ihering, 1907 Panopea quemadensis (Ihering, 1897) Glycimeris [sic] quemadensis Ihering, 1897 Panopaea [sic] sierrana Ihering, 1907 Panopea thomasi Ihering, 1914 Panopaea [sic] thomasi Ihering, 1914 Superfamily Pholadoidea Family Pholadidae Cyrtopleura lanceolata (d’Orbigny, 1846) Barnea sp., Ihering, 1914 Barnea saxicava Ihering, 1907 Netastoma paucispina (Ihering, 1897), new combination Pholas paucispina Ihering, 1897 Class Scaphopoda Order Dentaliida Family Dentaliidae Dentalium sulcosum Juliana Ihering, 1907 Dentalium matanzasense Ihering, 1907 Dentalium octocostellatum Pilsbry and Sharp, 1898 Dentalium octocostatum Ihering, 1897 [preoccupied name] Acknowledgments I acknowledge with gratitude the use of facilities at the Museo Argentine de Ciencias Naturales in Buenos Aires, and the assistance of the late Directors, Dr. Max Biraben and Dr. Jose Marfa Gallardo, and the Curator of Invertebrate Paleontology, Mrs. Nora Cabaleri. I am indebted to Dr. John E. Rawlins, Section of Invertebrate Zoology, CMNH, Pittsburgh, Pennsylvania, for his valuable sugges- tions and revisions of various drafts of this paper, and Mary Ann Daman of the same Section for technical assistance with the preparation of the manuscript. Bernadette G. Gallery, Kenneth J. Boss, and Richard I. Johnson provided information on references not available at CMNH. I also wish to pay tribute to the memory of Dr. Egidio Feruglio, who, during many decades of arduous exploration, contributed so much to our knowledge of Patagonian geology. 1996 Parodiz — ^Ihering Mollusca 293 Literature Cited Ameghino, F. 1902. L’Age des formations sedimentaires de Patagonie. Aeales de la Sociedad Cien- tifica Argentina, 54:161-180, 220-249, 283-242, — - — 1906. Les formations sedimentaires du Cretacee Superieur et du Tertiaire de Patagonia. Aeales del Museo Nacional de Buenos Aires, 8:1-568. Bohm, J. 1903. Fossilien von General Roca am Rio Negro [without title under “Protokole”; title in index]. Zeitschrift der Deutsche Geologische Gesellschaft, 54:71-72. Borchert, a. 1901. Die Molluskeefauea das Alter der Parana-Stuffe. Neues Jahrbuch fur Mineral- ogie. Geologic, und Palaontologie (Stuttgart), 14:1-78. Bmstow, a., and J. J. Parodiz. 1982. The stratigraphic paleontology of the Tertiary eon-marine sediments of Ecuador. Bulletin of Carnegie Museum, 19:1-53. Burckhardt, C. 1902. Le gisement supracretacique de Roca. Revista del Museo de La Plata, Zool- ogia, 10:207-223, 4 pL Caorsi, j. H., and Q. j. Goni. 1958. Geologfa Uraguaya. Boletin del Institute Geologico del Uruguay, 33:1-73. Carcelles, a. 1944. Catalogo de los Moluscos marines de Puerto Quequen. Revista del Museo de La Plata, Zoologia, 3(241):233-310. Carcelles, A., and J. J. Parodiz. 1938. Moluscos del cooteeido estomacal d& Astropecten cinguiatus. Physis (Buenos Aires), 12:255-266. Clench, W. J., and R. D. Turner. 1950. The genera Sthenorytis, Cirsotrema, Acirsa, Opalia, and Amaea in the western Atlantic. Johnsonia, 2(29)221-246. — — — . 1960. The genus Calliostoma in the western Atlantic. Johnsonia, 4(40): 1-80. Clessin, S. 1900. Eulimidae. Parts 456-460, pp. 121-200, in Systematisches Conchylien-Cabinet von Martini und Chemnitz (edition of H. C. Kuester), Band 1, Abteilung 28. CossMANN, M. 1899a. Description de quelques coquilles de la formation Saetacrazienne en Patagonie. Journal de Conchyliologie, 47(3):223-242. — . 1899A Essais de Paleoconchologie comparee. Livre 3. Paris, 201 pp., 8 pi. — — — . 1899c. Rectifications de nomenclature. Revue Critique de Paleozoologie (Paris), 3:133-139, 176-178. — — — . 1903. Essais de Paleoconchologie comparee. Livre 5. Paris, 215 pp., 9 pi. — — . 1904. Essais de Paleoconchologie comparee. Livre 6. Paris, 151 pp. . 1908. Rectifications de nomenclature. Revue Critique de Paleozoologie (Paris), 12:64-268. Del Rio, C. J. 1985. Primera mencion de la subfamilia Architectonicinae en el Terciario de la Pa- tagonia (Republica Argentina). Ameghiniana (Buenos Aires), 22(3-4):263-268. Del Rio, C., and G. A, Morra. 1984. Representantes de la subfamilia Pseudomelaxinae en el Ter- ciario de Patagonia. Ameghiniana (Buenos Aires), 22(1-2): 11 1-1 15. Doello-Jurado, M. 1919. Uee nouvelle espece de “Miltha” du tertiaire du I’Argentine. Physis (Buenos Aires), 4:558-562. ■ 1922. Un nuevo Mytilus fosil del Terciario de Patagonia. Anales de la Sociedad Cientifica Argentina, 94:86-90. 1927. Noticia preliminar sobre los moluscos fosiles de agua dulce mencionados por Wich- mann. Boletm de la Academia Nacional de Ciencias (Cordoba, Argentina), 30:407-416. . 1931. Una nueva especie de Voluta fosii de Patagonia. Anales del Museo Nacional de Buenos Aires, 36:515-519. Doring, A. 1881-1882. Informe Oficial de la Comision Cientifica Agregada al Estado Mayor General de la Expedicion al Rfo Negro. Buenos Aires, 3:299-530. . 1884. Estudios Hidrognosticos. Boletm de la Academia Nacional de Ciencias (Cordoba, Argentina), 6:328. Erdmann, S,, and G. A. Morra. 1985. Nuevos moluscos de la Formacion San Julian. Ameghiniana (Buenos Aires), 22(3— 4): 289-295. Feruglio, E. 1933. I terrazi marini della Patagonia. Anali del Reale Museu Gelogico di Bologna, 8(bis):236. — — — . 1935. Relacioees estratigraficas y faunisticas entre los estratos cretaceos y terciarios in ia region del Lago Argentino y ee la del Golfo de San Jorge. Boletm Informativo de Petroleos (Buenos Aires), nos. 120 and 130. — — . 1949-1950. Descripcion Geologica de la Patagonia. Publication of the Yacimientos Petro- liferos Fiscales (Buenos Aires), 3 volumes, 1112 pages. Frenguelli, j. 1920. Contribucion al conocimiento de la geologfa de Entre Rios. Boletm de la Academia Nacional de Ciencias (Cordoba, Argentina), 24:55-256. 294 Annals of Carnegie Museum VOL. 65 Hatcher, J. B. 1897. On the geology of southern Patagonia. 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Os Moluscos dos terrenos Terciarios da Patagonia. Revista do Museu Paulista (Sao Paulo), 2:217-382. . 1899a. Die Conchylien der Patagonisches Formation. Neues Jahrbuch fiir Mineralogie, Geo- logie, und Palaontologie, 2:1-46. — . 1899Z?. Descripcion de la Ostrea guaranitica. Anales de la Sociedad Cientffica Argentina, 47:63-64. — . 1900. The History of the Neotropical Region. Science, 1900:857-864. . 1902a. Historia de las Ostras Argentinas. Anales del Museo Nacional de Buenos Aires, 4: 109-123. . 1902^7. On the molluscan fauna of the Patagonian Tertiary. Proceedings of the American Philosophical Society, 41:132-137. — — . 1903. Les mollusques des terrains cretaciques superieurs de F Argentina orientale. Anales del Museo Nacional de Buenos Aires, 9:193-229. — . 1904. Nuevas observaciones sobre moluscos cretaceos y terciarios de Patagonia. Revista del Museo de La Plata, Zoologfa, 11:227-242. 1907a. Les mollusques fossiles du Tertiaire et du Cretacee Superieur de FArgentine. Anales del Museo Nacional de Buenos Aires, 14:1-611. . 1907^7. Archhelenis und Archinotis. W. Engelmann, Leipzig. 350 pp. . 1908. Mollusques du pampeen de Mar del Plata et Chapadmalal. Anales del Museo Nacional de Buenos Aires, 17:429. . 1914. Catalogo de moluscos Cretaceos e Terciarios da Argentina da collegao do author [with appendix in German]. Notas Preliminares da Revista do Museu Paulista (Sao Paulo), 1(3): 1-148. — — — . 1921. Nautilidos del Terciario de Chile y Argentina. Physis (Buenos Aires), 5:76. Klappenbach, M., and O. Olazarri. 1966. Notas sobre Strophocheilidae: Strophocheilus charruanus (Frenguelli). Revista de la Facultad de Humanidades y Ciencia (Montevideo), 22:233-238. Kraglievich, L. 1930. La Formacion Friaseana. Physis (Buenos Aires), 10:127-161. Martins, E. A. 1943. Uma nova especie de Molusco do genero Conus do Brasil (C. clenchi). Boletim do Museu Nacional do Rio de Janeiro, 128:1-8. . 1945. Conus carcellesi, un nuevo gastropodo marino de la Rep. Argentina. 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ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 3, Pp. 297-303 ' 30 August 1996 TWO NEW SPECIES OE XYRIS FROM CUBA Armando Urquiola Cruz^ Robert KraU Research Associate, Section of Botany Abstract Two new species of Xyris (Xyridaceae) are described and illustrated, both endemic to western Pinar del Rio Province, Cuba. Differences between these and two closely related species, X. navicularis Griseb. and X. bicarinata Griseb., are discussed. A key to the four species is included. Resumen Se describen e ilustran dos especies nuevas de Xyris (Xyridaceae) eedemicas del occidente de Pinar del Rio Provincia, Cuba. Se discuten las diferencias entre estas y la cercana X. navicularis, X. bicar- inata, ademas se dan las claves para las mismas. Introduction During an examination of material of Xyridaceae for the Flora of the Greater Antilles, the senior author had difficulty with some morphologic extremes of what previously had been identified as Xyris navicularis, particularly with samples got- ten by him and others from far western Pinar del Rio Province. Further fieldwork by him in western Pinar del Rio has served not only to confirm extreme variation, but also to find populations of two related and yet distinctly different species. These two novelties, now sufficiently vouchered, are described, figured, and dis- cussed below. Since there are also some character states in this complex of species that are shared by yet another Cuban endemic, X. bicarinata Griseb., we include and compare that species also. Institutional abbreviations in text follow Holmgren et al., 1990. Description of New Species Xyris bissei Urquiola & Krai, species novum (Fig. 1) Herba perennis, densicaespitosa; radices fibrosa. Caules breves. Folia princi- palia anguste linearia, 8-15 cm longa, erecta vel anguste flabellate expansa, va- ginis scaporum longiora. Laminae foliorum compressae, tortae, 2-3 mm latae, vaginis 3-4-plo longiores, longitudine subtiliter multinervosae, brunneolo-ferru- gineae vel testaceae; apices gradatim, turn abrupte incur vato-acuti, leviter incras- sati; margines cartilagineo-incrassati, grosse papillosi vel scabrelli; vaginae inte- grae, carinatae, carinibus ad apicem leviter papillosis, lateribus valde longitudinae multicostatae, basi brunneae, glabrae, valde pluricostatae, marginibus in laminas gradatim convergentibus, ad apicem ligulam erectam vel excurvatam acutam 1.5- ’ Departamento de Biologia, Instituto Superior Pedagogico de Pinar del Rio, Cuba. ^ Department of Biology, Vanderbilt University, Nashville, Tennessee. Submitted 19 June 1995. 297 298 Annals of Carnegie Museum VOL. 65 2 mm longam fascientes, infime gradatim dilatatae. Vaginae scaporum laxae, valde convolutae sed apertae, laminis brevibus. Scapi erecti, subfiliformes, torti et flexuosi, vulgo bicostati, distaliter parum compressi aut ovales, 0.7-1 mm lati, costis duas angustata scabropapillatas fascientes. Spicae obovoideae, late ovoideae ubi maturae, 8-13 mm longae, pluriflorae, breviter attenuatae; bracteae laxe spir- aliter imbricatae, tenues, pallide brunneolae, margine late scariosae, involuto-lac- eratescens; bracteae steriles ca. 4, mox valde excurvatae vel recur vatae, oblongo- ellipticae vel anguste ovatae, 5-5.5 (-6) mm, areis dorsalibus linearibus bracteam aequantibus, viridis, unicostalis; bracteae fertiles late ovatae aut ellipticae, 5.5- 6.5 mm longae, ecarinatae, leviter convexae, ad apicem obtusae et areis dorsalibus brevibus, ovatis, indistinctis, dilutis, ad apicem leviter carinatis. Sepala lateralia libera, subequilateralia, valde curvata, elliptica, ca. 5 mm longa, acuta, ala carinali firma, prominente crispato-ciliolata. Laminae petalorum late rhombeo-orbicula- tae, luteolae, margine denticulatae. Staminodia bibrachiata, brachiis longipeni^ cillata. Antherae oblongae, ca. 2 mm longae, profunde bifidae et auriculatae; filiis brevis, 0.5-0. 6 mm longis. Capsula ellipsoideae vel anguste obovoideae, 3 mm longae, apiculatae, pallide cinereo-brunneolae; placenta marginalis. Semina nu- merosa, ellipsoidea, 0.5 mm longa, pallide brunneola, translucida, longitudine vab de multicostata. Slender but stiff and densely cespitose, hard-based perennial 20-60 cm high. Roots coarse, fibrous. Stems short. Principal foliage leaves ascending or in narrow fans, longer than the scape sheaths, 8-15 cm long; blades linear, flat, 2-3 mm wide, narrowed gradually to an incurved-acute, slightly thickened tip, the edges cartilaginous-thickened, scabridulous-papillate, the surface smooth, finely nerved, greenish-brown; sheaths entire, keeled, ^-^4 as long as blades, with edges forming an erect or slightly excurved scarious ligule 1.5-2 mm long at narrowed base of blade, and gradually dilating downward, the base smooth, deep brown, strongly few-costate, the keel costa papillate-scabridulous. Scape sheaths loosely tubular, strongly convolute but open, short-bladed. Scapes wiry, twisted and flexuous, mostly two-costate, distally slightly compressed or oval, 0.7-1 mm wide, the costae forming two strong, scabro-papillate edges. Spikes ovoid, maturing broadly ovoid, 8-13 mm long, of several spirally and loosely imbricate, thin, light-brown bracts; sterile bracts mostly four, elliptic to narrowly ovate, 5-5.5 (-6) mm long, broadly acute, soon strongly excurved or recurved, lacerate, with pale or pale green, narrowly elliptic, elongate dorsal areas, carinate; fertile bracts broadly ovate or elliptic, 5. 5-6. 5 mm long, ecarinate, slightly convex with indistinct, ovate, paler, apically low-carinate dorsal areas, the apex obtuse or slightly emarginate, the thin borders strongly involute, becoming strongly lacerate. Lateral sepals free, subequilateral, strongly curvate, elliptic, ca. 5 mm long, acute, the firm but broad keel prominently crisped-ciliate. Petal blades rhombic-orbicular, 5 mm long, yel- low, low-toothed. Staminodia bibrachiate, the branches long-penicillate. Anthers oblong, ca. 2 mm long, deeply bifid and auriculate, on short filaments 0.5-0. 6 mm long. Capsule ellipsoid or narrowly obovoid, 3 mm long, apiculate, pale gray- Fig. 1. — Xyris bissei (Urquiola et al. 7876). a. Habit sketch; b. leaf tip; c. leaf midblade; d. leaf blade- sheath junction; e. leaf base; f. spike; g. lowest spike bract (left), -fertile bract (right); h. lateral sepal; i. petal blade and stamen; j. staminode and enlarged apical sector of beard hair (below); k. stylar apex; 1. capsule; m. seed. 299 1996 Cruz and Kral — New Cuban Xyris Icm 300 Annals of Carnegie Museum VOL. 65 brown, the placentation parietal. Seeds ellipsoid, 0.5 mm long, pale amber, trans- lucent, with ca. ten strong, horizontal, parallel ribs per side. Type. — CUBA. Pinar del Rio: Pinares sobre arenas blancas, proximo a la Laguna de Los Carneros, Santa Teresa, Guane, m.s.m. 5, 12 Apr. 1993, A. Ur- quiola, A. Urquiola f, E. Lezcano, R. Novo 7876 (holotype: HPPR; isotypes: GH, HAJB, HAG, NY, VDB). Etymology. — This species is named in honor of the late Johannes Bisse, an exceptional worker on the Cuban flora and teacher of many present-day Cuban botanists. Xyris mantuensis Urquiola & Krai, species novum (Fig. 2) Herba perennis, gracilis, densicaespitosa; radices fibrosa. Caules breves. Folia principalia anguste linearia, 15-20 (-25) cm longa, erecta vel anguste flabellate expansa, vaginis scaporum longiora. Laminae foliorum compressae, tortae vel rectae, 1-2 (-3) mm latae, vaginis 4-5-plo longiores, integrae, longitudine sub- tiliter multinervosae, olivaceae; apices gradatim decrescentes, anguste acuti, vulgo prominente scabro-papillatae; margines tenues, a basin dorsaliter papillosi vel cil- iatae; vaginae integrae, carinatae, carinibus ad apicem leviter scabridis, lateribus valde longitudine multicostatae, basi atroferrugineae aut brunneolae, nitidae, mar- ginibus in laminas gradatim convergentibus, ad apicem ligulam erectam vel ex- curvatam acutam 1.5-2 mm longam fascientes, infime gradatim, turn abrupte di- latatae. Vaginae scaporum laxae, conduplicatae, a medio ad apicem apertae, lam- inis breves. Scapi erecti, subfiliformes, torti et flexuosi, a basin teres, multicostali, apicem versus vulgo bicostati, ca. 1 mm lati, plerumque costis duas alas valde papillosas aut ciliatas fascientes. Spicae anguste ellipsoideo-cylindricae vel lan- ceoloideae, L2-L6 cm longae, pluriflorae; bracteae steriles mox 4, pari infimo oblongo, valde carinato, carinis scabridis, obtuso, 3-3.5 mm longo, paribus intimis fertilibus leviter breviores, in fertiles gradatim transientes; bracteae fertiles obov- atae vel late ellipticae, 5.5-6 mm longae, integrae turn laceratae, convexae, areis dorsalibus late ovatis, brevis, indistinctis, subapice carinatis. Sepala lateralia lib- era, subequilateralia, valde aut leviter curvata, 3.5-5 mm longa, acuta; ala carinali a medio ad apicem ciliolata. Laminae petalorum obovatae, ca. 5 mm longae, late rotundatae, serratae. Staminodia bibrachiata, brachiis longipenicillatis. An- therae oblongae, ca. 2 mm longae, profunde retusae et auriculatae; fila lata, ca. 7 mm longa. Capsula ellipsoidea, 4 mm longa, placentatio parietalis. Semina ellipsoidea, apiculata, 0.5 mm longa, translucida, longitudine valde multicostata. Herb perennial, slender, densely cespitose, hard-based, 30-45 cm high. Roots coarsely fibrous. Stems short. Principal foliage leaves ascending in narrow fans, 15-20 (-25) cm long; blades flattened, twisted or straight, narrowly linear, 1-2.5 (-3) mm wide, tapering gradually to slightly thickened, incurved, narrowly acute tip, the margin slightly if at all thickened, papillate to ciliate at least toward base, Fig. 2. — Xyris mantuensis {Urquiola et al. 7880). a. Habit sketch; b. leaf apex; c. leaf midblade; d. leaf blade-sheath junction; e. leaf base; f. spike; g. extreme example of midscape costa; h. lower spike bract (left), -fertile bract (right); i. lateral sepal; j. petal blade, stamen; k. staminode and enlarged apical sector of beard hair (below right); 1. capsule valve, adaxial side; m. capsule, valves pulled apart; n. seed. 1996 Cruz and Kral — New Cuban Xyris 301 OS mm 302 Annals of Carnegie Museum VOL. 65 surface smooth, greenish-maroon with several low nerves, blade base narrowed at junction with sheath; sheaths entire, multicostate, keeled distally, keel costa strongly scabro-ciliate, at blade junction producing an erect to excurved ligule 1.5-2 mm, gradually, then abruptly dilating downsheath to a broad base, distally pale yellow-green, proximally deep red-brown or brownish, lustrous. Scape sheath conduplicate but loose, open most of its length, and with a short, linear blade similar to foliage leaf. Scapes subfiliform, twisted and flexuous, proximally terete, several-costate, costae narrow but strong, distally with fewer, often two, costae, ca. 1 mm wide, the strong costae forming edges, these commonly intermittently papillate to strongly tuberculate-scabrid, or retrorsely so. Spikes narrowly ellip- soid-cylindric or lanceoloid, 1.2-1. 6 cm long, of many subdecussately arranged, loosely imbricate, ascending, tan to brown bracts with indistinct dorsal areas; sterile bracts usually four, the lower pair oblong, 3-3.5 mm long, blunt, carinate, the keel raised and scabrid, the upper pair slightly longer, broader; fertile bracts broadly elliptic or narrowly obovate, convex, 5.5-6 mm long, entire aging lac- erate, the paler dorsal area with a low carina subapically. Lateral sepals free, subequilateral, strongly or slightly curved, 3.5-5 mm long, the broadly rounded apex coarsely toothed. Petal blades obovate, ca. 5 mm long, broadly rounded, serrate. Staminodia bibrachiate, branches long-penicillate-hairy. Anthers oblong, ca. 2 mm long, deeply bifid and auriculate, the connective broad, on short flattened filaments ca. 0.7 mm long. Capsule ellipsoid, 4 mm long, placentation three- parietal. Seeds broadly ellipsoid, 0.5 mm long, amber, with ca. ten strong, mostly parallel ribs per side. Type. — CUBA. Pinar del Rio: Pinares sobre arenas blancas de Playa La Cana, Mantua, m.s.m. 5, 1 Jan. 1994, A. Urquiola & A. Urquiola f 7880 (holotype HPPR; isotypes GH, HAC, HAJB, NY, VDB). Etymology. — This species is named for the type locality. Discussion There are two other species of Xyris in the Antilles with which the two new species could be confused and which can co-occur with them in the white sand savanna or pineland ecosystems of Pinar del Rio, namely X. navicularis Grisebach (as mentioned), the nearest morphologically to X. mantuensis, and X. bicarinata Grisebach, which resembles X. bissei in spike and somewhat in leaf. These four species are compared by means of the following key: la. Mature (seeding) spikes broadly ovoid or obovoid, with at least the lower bracts strongly excurved or recurved. 2a. Leaf blades narrowly linear or filiform, 1-2 mm wide, the edges thickened, often with a median furrow, smooth or nearly so, and not in color contrast with the rest of the blade, medially with 1-3 strongly raised nerves, the others lower or indistinct X. bicarinata 2b. Leaf blades linear, 2-5 mm wide, the edges a narrow or strong, cartilaginous, smooth to papillate band in pale contrast with the rest of the blade, the faces with more and less strongly raised nerves. 3a. Bracts aging lacerate and also strongly revolute-inrolled; blades of principal leaves mostly 2-3 mm wide, their margins moderately to strongly papillate or rugulose- papillate; scape costae mostly papillate or scabridulous. ... .X. bissei 3b. Bracts aging lacerate but scarcely if at all revolute; blades of principal leaves 2-5 mm wide, their margins mostly smooth, rarely sparsely papillate; scape costae smooth to sparsely papillate .X. navicularis lb. Mature (seeding) spikes narrowly ovoid to lanceoloid or ellipsoid-cylindric, the bracts (spreading-) ascending to erect. 4a. Spikes lance-cylindric, narrowly ellipsoid, or cylindric, 0.4-0.7 cm thick, mostly widest 1996 Cruz and Kral — New Cuban Xyris 303 at or near middle; leaf sheath apex with ligule 1-1.5 mm long; margins of leaf blades remotely papillate to papillate or even scabrociliate; the most prominent scape costae papillate to intermittently scabrociliate, often retrorsely so . .X. mantuensis 4b. Spikes ovoid to lance-ovoid or broadly cylindric, 0.7- 1.0 cm thick; leaf sheath apex with ligule 0.2-0. 5 mm long or ligule lacking; margins of leaf blades smooth, rarely sparsely papillate; scape costae smooth or sparsely or distantly papillate. .X navicularis Of the four species treated in the key above, only X navicularis is found outside Cuba, appearing in savanna or pine savanna in Belize, Honduras, Nicaragua, and (rarely) in Venezuela. The rest are confined to white-sand savannas in western Cuba. There all four occur, but the two novelties are thus far known only from a few populations within a very narrow area of far western Pinar del Rio. Literature Cited Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index Herbariorum. Part I: The Herbaria of the World. Eighth Edition. Regnum Vegetabile, 120:1-693. ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 3, Pp. 305-321 30 August 1996 A NEW PLESIADAPIFORM (MAMMALIA: PLESIADAPIFORMES) FROM THE EARLY EOCENE OF THE BIGHORN BASIN, WYOMING Kenneth D. Rose^ Research Associate, Section of Vertebrate Paleontology Thomas M. Bown^ Research Associate, Section of Vertebrate Paleontology Abstract Dentaries of a very small, specialized plesiadapiform from the lower Eocene Willwood Formation are the basis for a new genus and species, Picromomys petersonorurn. It was the smallest known plesiadapiform, weighing only about 10 g. Like some other plesiadapiforms, the new taxon has a strongly procumbent, hypertrophied medial incisor, followed by very reduced dentition between the incisor and P4. Unlike all other plesiadapiforms, it has a unique semimolariform P4 reminiscent of Mj in picrodontids, as well as lower molars bearing an accessory trigonid cusp anterobuccal to the pro- toconid. The closest known relative of Picromomys appears to be Bridgerian Alveojunctus, and we group these two genera here in the new family Picromomyidae. Picromomyids share derived similar- ities with several plesiadapiform families but are probably most closely related to either Micromo- myidae or uintasoricine Microsyopidae. Introduction Field work in early Wasatchian strata of the Willwood Formation of the Bighorn Basin in 1994 yielded dentaries of a highly distinctive new plesiadapiform. The new form is unusual in several respects, including a highly modified lower fourth premolar, presence of an accessory trigonid cusp on Mi_2, and its extremely small size. It also shares numerous derived traits with various plesiadapiforms assigned to four different families, but there is no compelling evidence to support a special relationship to any one particular family. Hence homoplasy must account for many of the similarities. The only strongly supported relationship is to the poorly known Bridgerian species Alveojunctus minutus, and even this alliance is based on very limited evidence. The latter has been considered to be a uintasoricine microsyopid (Bown, 1982; Gunnell, 1989), but its apparent similarities to microsyopids could well be convergent, for it also shares potential synapomorphies with other fami- lies. The new taxon and Alveojunctus are here placed in a new family of plesia- dapiforms because of their unusual specializations and the lack of an unequivocal relationship to any known family of plesiadapiforms. Recognition of new species of fossil mammals from the lower Eocene Will- wood Formation is not unusual, despite more than a century of field work in these strata. Discovery of new genera or families, however, is considerably rarer. It ^ Department of Cell Biology and Anatomy, The Johns Hopkins University School of Medicine, Bal- timore, Maryland 21205. ^ Adjoint Associate Professor, Department of Geological Sciences, University of Colorado, Boulder, Colorado 80309. Submitted 30 October 1995. 305 306 Annals of Carnegie Museum VOL. 65 serves as a reminder that we still have much to learn about mammalian diversity during the Wasatchian — even in well-studied sequences. Abbreviations used in text are: AMNH, American Museum of Natural History, New York, New York; USGS, US Geological Survey, Denver, Colorado; USNM, National Museum of Natural History, Smithsonian Institution, Washington, D.C.; YPM-PU, Peabody Museum, Yale University, New Haven, Connecticut. Systematic Paleontology Order Plesiadapiformes Simons and Tattersall, in Simons, 1972 Picromomyidae, new family Type Genus. — Picromomys, new genus. Included Genera. — Type genus and Alveojunctus. Distribution. — Wasatchian-Bridgerian (early-middle Eocene) of Wyoming. Diagnosis. — Diminutive plesiadapiforms having the following combination of features: very low-crowned lower molars with shallow hypoflexids, low and an- teriorly canted trigonids, and hypoconulids shifted towards entoconids but sepa- rated by a distinct entoconid notch; P4 larger than M, and with open, anteriorly canted trigonid and large, squared talonid with fully buccal cristid obliqua and no hypoflexid; anterobuccal cingulids poorly developed or absent. Discussion. — Although Picromomys and Alveojunctus are extremely rare and their known fossils very fragmentary, they share several unusual features that suggest close relationship. Moreover, they are dentally so distinctive that they cannot reasonably be assigned to any named family. No other plesiadapiforms have modified P4 in the same way. Nonetheless, there can be little doubt that both are plesiadapiforms (see Discussion following description of the new species), and there are derived resemblances to members of four different families: Micro- momyidae, Microsyopidae, Picrodontidae, and Paromomyidae (see below). In no case are the similarities very compelling, however, and the likelihood that most of them result from convergence is equally plausible. It is possible that more complete material will strengthen the alliance of Picro- momyidae with one of those families — most probably Micromomyidae or Mi- crosyopidae, to judge from present evidence — and eventually warrant reclassifi- cation (e.g., as a subfamily of Micromomyidae or a tribe of Uintasoricinae). How- ever, family level separation seems justified by the kind and extent of dental apomorphies present in picromomyids. Many plesiadapiforms exhibit modifica- tions of antemolar teeth while retaining rather conservative molars. In particular, a specialized, usually hypertrophied P4 characterizes several plesiadapiform fam- ilies (viz. Carpolestidae, Paromomyidae, Micromomyidae), each of which has modified this tooth in a different manner. The defining specializations of picro- momyids are comparable to any of these. Bridgerian Alveojunctus is known from only a small number of isolated teeth (Fig. 1) representing two or three loci, P4, M2(?), and possibly M3. The single molar (the holotype) known from the Aycross Formation was identified as Mj by Bown (1982), but in comparison with the new species appears more likely to be M2. As noted by Bown, an M3 from the early Bridgerian Cathedral Bluffs Tongue (Wasatch Formation, Green River Basin), referred to cf. Niptomomys sp. by West and Dawson (1973), may also belong to Alveojunctus. These two molars are distinctive and differ from those of the new species in uniquely having the trigonid basin conjoined with the talonid basin through a deep trigonid notch. P4 of AT 1996 Rose and Bown — New plesiadapiform from Wyoming Eocene 307 Fig. L- — SEM stereopairs of picromomyids. A, Picromomys petersonorum, n. gen. and sp., holotype (USNM 487900), right dentary with base of I, and P4-M3 in occlusal view. B-E, Aiveojunctus minutus. B, unworn left P4 (USNM 250589); C, worn right P4 (USNM 251446); D, right holotype (USGS 2005); B-D in occlusal view, E, left P4 (USNM 250589) in buccal view. veojunctus differs from that of the new species in having a talonid with a clear basin and a particularly prominent entoconid, which is often swollen into the basin. Picromomys, new genus Type and Only Known Species.— Picromomys petersonorum, new species. Diagnosis.- — Very small plesiadapiform (Mj length = LI mm, just over half the linear dimensions of Aiveojunctus) with enlarged, nearly horizontal medial incisor and only two small teeth between Ij and P4. Differs from Aiveojunctus 308 Annals of Carnegie Museum VOL. 65 and all other plesiadapiforms in having a prominent cusp-like expansion on M,_2 anterobuccal to the protoconid. P4 trigonid with poorly defined blunt cusps. P4 talonid about same length as trigonid and wider than trigonid, as in Alveo- junctus\ and flat (without basin), sloping posterolingually, and lacking distinct hypoconulid and entoconid, all in contrast to Alveojunctus. Distribution. — Wasatchian (early Eocene) of Wyoming. Etymology. — Greek pikros, bitter, in allusion to the superficial resemblance of P4 to the first lower molar of Paleocene Picrodus, + Omomys, middle Eocene primate and suffix commonly used for small primitive primates and plesiadapiforms. Picromomys petersonorum, new species (Fig. 1, 2) Holotype.- — USNM 487900, right dentary with P4-=M3, base of I|, and two al- veoli between incisor and P4; left dentary with base of Ij, incomplete P4, and all other alveoli. Only known specimen; collected by T. M. Bown in 1994. The two dentaries are assumed to represent the same individual because they are the same size, are similarly worn, and were found together in situ during quarrying, within about 1 cm of each other. Locality and Horizon. — A stage 1 paleosol in meander-belt mudstones at USGS locality D-2035 in the Dorsey Creek badlands, SE Va Sec. 12, T.50 N., R.95 W., Big Horn County, Wyoming; 390 m-level of the Willwood Formation, early Was- atchian. See Bown and Kraus (1987) and Bown et al. (1994) for further infor- mation on Willwood sediments and stratigraphy. Diagnosis. — As for genus. Etymology. — For Ted and LaDean Peterson of Worland, Wyoming, in recognition of years of friend- ship and generous assistance to our expedition. Description. — The dentary appears relatively deep because of the very low-crowned molars, but it is actually comparatively shallower than in some paromomyids. Two mental foramina are present, the larger one positioned below the front of P4 and the other under the trigonid of Mj. The cheek tooth row is not perfectly straight in superior view, but forms a gentle arc directed slightly mediad at the posterior end, a common configuration in plesiadapiforms. The lower dentition consists of a hypertrophied, procumbent anterior tooth, assumed to be Ij, and six postcanine teeth. The base of I,, present in both dentaries, is oriented essentially horizontally; the crowns of both are missing. The limited portion preserved, however, provides significant information bearing on the affinities of Picromomys. The tooth is laterally compressed, with an elliptical cross section that is almost flat medially and convex laterally. A faint horizontal ridge is present on the anterolateral aspect near the broken edge of I,, whereas the dorsal aspect is smoothly rounded. The preserved part of the incisor approximates that of paromomyids and micromomyids more closely than that of other plesiadapiforms, such as carpolestids or plesiadapids. Moreover, it is distinctly different from the diagnostic lanceolate lower incisor of microsyopids (including Navajovius, Arctodontomys, Microsyops, Niptomomys, and Uintasorex). In the latter group, the incisor has an abrupt dorsal ex- pansion just distal to its base, and the crown is rotated medially. As a result, the homologue of the lateral crest in other plesiadapiform incisors is reoriented to form a sharp dorsal margin in microsy- opids, and what was the “dorsal” surface faces medially. This sharp dorsal margin is absent in Picromomys. Only two alveoli are present between I, and P4, making Picromomys one of the most derived plesiadapiforms in terms of antemolar reduction. The dimensions and orientation of the two alveoli make it probable that they held two single-rooted teeth rather than a single, large double-rooted premolar. The first alveolus is smaller and slightly lower than the one behind, and is separated from Ij by a short diastema. The inclined position of this alveolus suggests that it housed a somewhat procumbent tooth whose crown projected mesially over the front of its root, similar to the condition in apatemyids, carpolestids, and the micromomyid Tinimomys, as well as in some soricids, but not so closely appressed to I, as in uintasoricines. The homologies of this tooth are equivocal in Picromomys and several of the other taxa; here it is tentatively identified as P2, but it could just as well be I2 or 1996 Rose and Bown — New plesiadapiform from Wyoming Eocene 309 Fig. 2. — Picromomys petersonorum, n. gen. and sp., holotype right dentary (USNM 487900) in lateral, occlusal, and medial views. Scale = 1 mm. the canine. The second alveolus is elliptical and somewhat larger, and probably held a tooth (P3) with a fused double root as in Uintasorex. P4 is the most distinctive tooth and, apart from the noted resemblance to Alveojunctus, is unique in form among plesiadapiforms. The trigonid is tall but anteriorly canted, with a compact and wedge- shaped summit bearing blunt, ill-defined cusps or, more precisely, crests. The crests form a trefoil or Y-shaped pattern, a short anterior arm representing the paracristid, and a longer posterolingually directed arm joining an indistinct metaconid, which is indicated by a subtle swelling on the back of the trigonid. Near the midpoint (site of the protoconid?) a lower, blunt crest projects posterobuccally and merges into a sharper crest that descends the posterolateral wall of the trigonid. There it meets 310 Annals of Carnegie Museum VOL. 65 the talonid just lateral to the cristid obliqua on the buccal tooth margin, producing a faint hint of a hypoflexid. The unusual appearance of the trigonid probably results from a combination of wear and weakly defined cusps, and has been interpreted in part by comparison with a series of unworn and worn P4S of Alveojunctus (see Fig. 1), its closest morphological counterpart. The talonid of P4 is about the same length as the trigonid and is slightly broader than long. The cristid obliqua is high and fully buccal and the entocristid is much lower. These crests meet the straight postcristid at right angles, and all three crests are situated on the periphery of the talonid, barely elevated above the enclosed surface. The hypoconid, a rounded conical swelling at the posterolateral corner of the tooth, is the highest point on the talonid and the only distinct cusp. Nonetheless, it is hardly any higher than the cristid obliqua. From the hypoconid, the postcristid slopes medially to a low point at the lingual margin, where it meets the entocristid. This is normally the location of the entoconid in plesiadapiforms and other mammals that have well-developed P4 talonids, but in Picro- momys it is the lowest part of the talonid. From there, the entocristid ascends gradually to its highest point at the posterolingual border of the trigonid; hence, there is no talonid notch. Neither hypoconulid nor entoconid cusps are discernible, although a small bump of enamel near the hypoconid may be the remnant of a hypoconulid. Rather than having a basin, the talonid is flat. It is distinctly lower lingually than buccally, and bears an irregular fold of enamel that crosses diagonally from just inside the hypoconid to the anterolingual corner. P4 lacks cingulids. The lower molars are very low crowned, with trigonids that are sharply canted anteriorly and appear anteroposteriorly constricted in lateral view. Although of roughly similar dimensions, the molars in- crease in length in the sequence M2 < Mj < M3. The standard trigonid cusps are bunodont, the metaconid tallest and the paraconid low and reduced, closely resembling the arrangement in Tinimo- mys. The trigonids are transversely narrow and become progressively more compressed anteroposte- riorly from M, to M3. The most distinctive feature of the molars is an autapomorphous cusp-like swelling, with a well-developed, beveled wear facet, situated just anterobuccal to the protoconid. It is large and columnar on M,, somewhat smaller on M2, and barely visible on M3. Apatemyids also have an accessory trigonid cusp anterior to the protoconid, but the resemblance to Picromomys is quite superficial. The molar talonids are increasingly broader and have increasingly deeper basins from M, to M3. The talonid of M, is slightly longer than wide, whereas that of M2 is wider than long. The talonid of M3 is slightly broader still, but the third lobe is only moderately expanded, unlike the enlarged third lobe of paromomyids and plesiadapids. The cristid obliqua joins the trigonid near the protoconid on M, and is progressively more buccal on M2_3, resulting in very shallow hypoflexids. A faint mesoconid is present on M,. The hypoconulid and entoconid are especially prominent on M,_2. The hypoconulid is lenticular or crescentic, not round or conical like the hypoconid on M2, and is distinctly shifted toward the entoconid but separated from the latter by a distinct, shallow notch. These conditions are closely approximated in uintasoricine microsyopids {Uintasorex more than Niptomomys) and in mi- cromomyids (especially Tinimomys). Unlike P4, the molars have a well-defined talonid notch between the trigonid and entocristid. The molar ectocingulids are poorly developed, being best expressed at the hypoflexids but virtually absent anteriorly. The talonid of M3 has weakly crenulated enamel. Measurements (mm) of the Holotype. — P4 length = 1.30, breadth = 0.70; Mj length = 1.10, breadth = 0.75; M2 length = 1.00, breadth = 0.80; M3 length = 1.20, breadth = 0.75; depth of dentary below Mj = 2.70. Discussion. — With a P4-M3 length of 4.5 mm, the holotype belonged to an animal much smaller than any known primate, and even smaller than the dimin- utive micromomyid plesiadapiforms Tinimomys and Micromomys and the mi- crosyopid Berruvius, up to now the smallest known plesiadapiforms. It weighed only 9.0-10.6 g, as estimated from regressions of Mj area on body mass in all extant primates, or in prosimians, respectively (Conroy, 1987). A somewhat higher estimate, 26 g, is obtained using the regression of Gingerich et al. (1982) but, as they note, this equation overestimates the weight of insectivorous primates. Whichever regression is used, Picromomys apparently was no more than one-half to two-thirds the size of Tinimomys (Rose et al., 1993). Picromomys resembles plesiadapiforms in several presumably derived dental traits, including presence of a hypertrophied, procumbent incisor together with low-crowned molars having reduced paraconids and broad, basined talonids. Also 1996 Rose and Bown — -New plesiadapiform from Wyoming Eocene 311 like many plesiadapiforms, it is further characterized by reduction in number and size of antemolar teeth between the enlarged incisor and P4, as well as enlargement and specialization of P4, which has a trigonid taller than any other tooth. Although some omomyids can be similarly characterized, the combination of features in Picromomys conforms much more closely with that in plesiadapiforms. Thus as^ signment of Picromomys to Plesiadapiformes seems beyond doubt. The details of its dental morphology, however, indicate that Picromomys belongs to a clade divergent from other known plesiadapiforms. Based on the very low-crowned molars and unusual structure of P4, the only other known member of this clade is Alveojunctus. From comparisons summarized below, it will be apparent that the precise re- lationships and phylogenetic position of Picromomys and the Picromomyidae are uncertain. Picromomys shares numerous apparently derived dental characters, in- dicating potential relationship, with four known plesiadapiform clades: Paromo- myidae, Picrodontidae, Micromomyidae, and uintasoriciee Microsyopidae. As mentioned above, the molar trigonid arrangement in Picromomys is also vaguely reminiscent of that in Apatemyidae, but in the latter the accessory trigonid cusp is low and well anterior to the protoconid, forming a quadrilateral with the other trigonid cusps. There are no other compelling resemblances to apatemyids. Phylogenetic Position of Picromomys Cladistic analysis (using Heenig86; Farris, 1988) of 29 characters of the lower dentition in 12 plesiadapiform genera (see Appendices 1 and 2) was undertaken in an effort to resolve the relationships of Picromomys. The analysis yielded two equally most parsimonious trees, each with considerable homoplasy (tree length = 72, consistency index = 0.48, retention index = 0.53), one grouping picro- momyids in a clade with micromomyids, and the other grouping them succes- sively with Niptornomys, Picrodus, and Ignacius (Fig. 5). When the lanceolate incisor of microsyopids (widely considered to be an important synapomorphy of the family) was weighted 2, a single shortest tree (B) of 73 steps resulted, tree A having 74 steps under this weighting. The two contrasting trees, and concomitant uncertain position of Picromomys, result in part from the very limited anatomical evidence and suggest that lower dentitions alone-=-at least the dental characters used in this analysis— are insuf- ficient to establish interrelationships among plesiadapiforms with confidence. In this regard it is noteworthy that neither of the two shortest trees corroborates the widely accepted holophyly of Paromomyidae, sensu stricto {Paromomys and Ig- nacius in Fig. 5). Another important factor contributing to the uncertainty is the admitted ambiguity in coding several characters, usually resulting from either questionable homology (e.g., of antemolar teeth) or the inherent difficulty in cod- ing intermediate states of continous characters. Specific ambiguities are noted in Appendix 1. The variable nature of some of the characters makes it difficult to code them consistently, and further complicates character assessment. Nonethe- less, an attempt to resolve relationships on this basis is justified because the lower dentition has been the basis for much of the systematics of plesiadapiforms, and is the only evidence available for Picromomys; moreover, it is adequate to ascer- tain the uniqueness of Picromomys. In both trees, Picromomys and Alveojunctus compose a monophyletic group supported by the following synapomorphies (one or both trees, indicated by A, 312 Annals of Carnegie Museum VOL. 65 B, or AB): P4 larger than Mj (lOA) and with metaconid present (27B) and talonid as long as trigonid (llAB), a buccally oriented cristid obliqua on P4 (14AB), shallow hypoflexids (16B), low, mesially canted trigonids (17B), extreme molar brachydonty (18B), weak or absent anterobuccal cingulids (19B), moderately ex- pressed molar paraconids (21A-reversal), and molar hypoconulids close to ento- conids (28 A). Only character 11(1), however, is unique to picromomyids. This combination of common traits results in a closer correspondence between these two genera than either one has with any other plesiadapiform. Even so, both Picromomys and Alveojunctus are very incompletely known, and their proposed alliance remains to be tested as the record improves. Comparisons with Other Plesiadapiformes Paromomyidae. — Like Picromomys, both Paromomys and Ignacius have rela- tively low-crowned molars (character 18) and broad talonids on P4 (13). Ignacius further resembles Picromomys in having a strongly procumbent medial incisor (2), reduced anterior dentition between Ij and P4, a buccally oriented cristid ob- liqua on P4 (14), shallow hypoflexids (16), weak molar ectocingulids (19), and low and mesially canted molar trigonids (17) (Fig. 3B, 4B). Several of these traits are also shared with Picrodus. Both Picromomys and Ignacius also have relatively deep dentaries, but this apparent resemblance may be exaggerated because both forms are brachydont. As already noted, holophyly of the Paromomyidae sensu stricto is not supported by either of the two shortest trees in the parsimony analysis, but this almost certainly reflects inadequacy of the characters used here rather than lack of close relationship. In fact, two important traits, a broad, basined talonid on P4 (13) and an expanded third lobe on M3 (25), would support that grouping. All known paromomyids except Paromomys are even more derived in antemolar tooth loss than Picromomys, and all (including Paromomys) tend to emphasize crests over cusps, in contrast to picromomyids. This disparity, presumably related to a dif- ference in diet, suggests that picromomyids and paromomyids are not very closely related. Picrodontidae. — Picromomys shares several derived traits with Torrejonian Pi- crodus, including very low-crowned molars (character 18) with shallow hypo- flexids (16), weak ectocingulids (19), transversely compressed trigonids (20) that are low and mesially canted (17), and variably crenulated enamel (26) (Fig. 3H, 4H). In both taxa P3 is one-rooted (9). None of these characters is uniquely shared by these two genera, however. Also intriguing is the resemblance between the unusual enlarged cheek teeth of these two genera; but the teeth are believed to be nonhomologous (P4 in Picromomys, Mj in Picrodus; Szalay, 1968), and close inspection shows the similarity to be far from precise. Consequently, close rela- tionship between picromomyids and picrodontids is unlikely, and closer probable relationships are suggested by the cladistic analysis. Uintasoricine Microsyopidae. — Tree A indicates the Wasatchian uintasoricine Niptomomys as the sister taxon of picromomyids, based on the shared presence of a strongly procumbent incisor (2), P4 with a paracristid (12[l]-reversal), a metaconid (27), and a broad but unbasined talonid (13), prominent molar hypo- conulids (23-reversal), and an entoconid notch (29). These characters apply equal- ly to Bridgerian Uintasorex (see Szalay, 1969), which further resembles Picro- momys (but not Alveojunctus) in having a low entoconid region on P4. Uintasor- i996 Rose and BowN"=New plesiadapiform from Wyoming Eocene 313 icines also resemble picromomyids in having very brachydont molars with very broad talonid basins. There are other derived similarities as well which are also present in successive outgroups. However, the fundamental differences in the form of P4, the molar trigonids (transversely broad and mesiodistally compressed, with more reduced paraconids in uintasoricines), and especially Ij, together with the improbable required reversal (regeneration of a P4 paraconid following its loss), suggest that the observed resemblances between uintasoricines and picromomyids could well be convergent, Micromomyidae.—-MicmmQmyid^ are derived, like picromomyids, in having P4 larger than Mj (10), transversely compressed molar trigonids (20, Picromomys only), and molar hypoconulids shifted toward the entoconid (28) (see Fig. 3C, D; 4C, D; and 5B, node 3). Additional derived traits found in picromomyids and Tinimomys (Fig. 5B, node 4), include a strongly procumbent Ij (2), loss of several antemolar teeth (4, 6, 8), a distinct entoconid notch on lower molars (29), and a broad, basined P4 talonid with an enlarged entoconid (13[2], 15 [2], Alveojunctus only). Tinimomys and picromomyids also have similar bunodont trigonid cusps, but this resemblance is so difficult to characterize precisely that we have not included it in the cladistic analysis. Overall, these resemblances are weakly sug- gestive of relationship between picromomyids and micromomyids. Characters for Micromomys used in this analysis were based principally on Tiffanian M. fremdi (Fox, 1984), the most primitive known micromomyid. In our opinion, micromomyids have been oversplit generically, and we recognize only the genera Micromomys and Tinimomys in this report. The features originally cited to distinguish Chaiicomomys from Micromomys (Beard and Houde, 1989) and Myrmekomomys from Tinimomys (Robinson, 1994) do not justify their generic separation. Chaiicomomys closely resembles Micromomys in its taller P4 with much smaller talonid basin, more acute (less bunodont) molar cusps, and several other traits cited by Beard and Houde to differentiate Chaiicomomys from TinT momys. Hence there seems little reason to separate it generically from Micro- momys. At the same time, '"Chaiicomomys” antelucanus strengthens the proba- bility of a close relationship between Micromomys and Tinimomys. The putative generic distinctions of Myrmekomomys compared to Tinimomys (“relatively great- er molar trigonid relief and taller talonid. . . . hypoconulid of Mi_2 more devel- oped;” Robinson, 1994:86) appear to vary intraspecifically in the Will wood sam- ple of r. graybulliensis. The only character that differs from known samples is the more constricted hypoconulid lobe of M3 in the holotype of Myrmekomomys loomisi, a feature that could be an intraspecific variant and is unlikely to warrant generic recognition. In the present study, therefore, we consider Micromomys to include Chaiicomomys, and Tinimomys to include Myrmekomomys. Discussion and Conclusions Picromomys is a highly distinctive new plesiadapiform, demonstrably allied closely with only one other known taxon, Alveojunctus. These two genera are allocated to a new family, Picromomyidae, to recognize their highly distinctive dental anatomy and their ambiguous phylogenetic position among plesiadapi- forms. Picromomyids possess a number of derived characters suggesting affinity with uintasoricine microsyopids, but the combination of derived resemblances supporting a relationship between picromomyids and micromomyids, especially Tinimomys, seems a little less susceptible to convergence and requires fewer 314 Annals of Carnegie Museum VOL. 65 Fig. 3. — Plesiadapiform lower right dentitions in lateral view. A, Picromomys petersonorum, n. gen. and sp. (USNM 487900). B, Ignacius fremontensis (YPM-PU 14790, reversed; Paromomyidae). C, Tinimomys graybulliensis (after Rose et al., 1993; Micromomyidae). D, Micromomys antelucanus (after Beard and Houde, 1989; Micromomyidae). E, Niptomomys doreenae (after Rose et al., 1993; incisor restored from N. thelmae; Microsyopidae). F, Navajovius kohlhaasae (AMNH 17390, holotype, reversed; Microsyopidae). G, Palenochtha minor (AMNH 35451, anterior after Gunnell, 1989; Palae- chthonidae?). H, Picrodus silberlingi (AMNH 35456, incisor after Szalay and Delson, 1979). Scales = 1 mm. 1996 Rose and Bown — New plesiadapiform from Wyoming Eocene 315 E F Fig. 3. — Continued. improbable reversals or autapomorphies than other arrangements. Nonetheless, the high degree of homoplasy, including multiple convergent autapomorphies, re- quired in both of the two shortest trees does not lend particular confidence to either one, and additional anatomical evidence is probably needed to resolve these relationships better. When first described, Micromomys and Tinimomys were assigned to the Paro- momyidae, tribe Micromomyini (Szalay, 1973, 1974). Bown and Rose (1976; Rose and Bown, 1982) used a more restricted concept of Paromomyidae, and transferred Micromomys, Tinimomys, and several other taxa formerly included in Paromomyidae to the Microsyopidae, an assignment followed by Fox (1984) and Gunnell (1989). More recent studies have suggested that Micromomyini diverged 316 Annals of Carnegie Museum VOL. 65 Fig. 4. — Plesiadapiform lower dentitions in occlusal view; same specimens as in Fig. 3. A, Picromomys petersonorum, n. gen. and sp.; B, Ignacius fremontensis; C, Tinimomys graybulUensis; D, Micromomys antelucanus; E, Niptomomys doreenac, F, Navajovius kohlhaasae; G, Palenochtha minor, H, Picrodus silberlingi. Scales = 1 mm. 1996 Rose and Bown — New plesiadapiform from Wyoming Eocene 317 >3 >4 >5 ► 6 Tree A 4T Purgatorius Palaechthon Tinimomys Micromomys Palenochtha Paromomys Navajovius Ignacius Picrodus Niptomomys Picromomys Alveojunctus Tree B {e: Purgatorius Palaechthon Alveojunctus Picromomys Tinimomys Micromomys Palenochtha Paromomys Niptomomys Navajovius Ignacius Picrodus Fig. 5. — Possible relationships among picromomyids and some other plesiadapiforms: the two shortest trees based on lower dental characters given in Appendices 1 and 2, resulting from an exhaustive search using the implicit enumeration algorithm of Hennig86, which finds all trees of minimal length (Farris, 1988). Tree length = 72, consistency index = 0.48, retention index = 0.53. Ordering the characters did not change the tree topologies. Character optimization was performed using CL ADOS (Nixon, 1992). Nodes in tree A are supported by the following synapomorphies (change is from 0 to 1 for binary characters; state indicated for multistate characters): node 1 — 5, 7, 8(1); node 2 — 12(1); node 3 (Mi- cromomyidae) — 4, 10, 20, 24, 28; node 4—23; node 5 — 18(1), 21, 22; node 6 — 4, 12(2), 19; node 7—6(1), 16, 17; node 8—9, 18(2); node 9—2, 12( 1-reversal), 13(1), 23 (0-reversal), 27, 29; node 10 (Picromomyidae) — 10, 11, 14, 21 (0-reversal), 28. Autapomorphies required in this tree are: Palae- chthon— 21, 27; Tinimomys — 2, 6(2), 13(2), 15(2), 18(1), 25, 28, 29; Micromomys — 8(0“reversal); Palenochtha — 2; Paromomys — 8(0-reversal), 13(2), 25, 26; Navajovius — 3, 23(0-reversal), 27; Ig- nacius— 2, 6(2), 8(2), 13(2), 14, 21 (0-reversal), 25; Picrodus — 8(2), 20, 26; Niptomomys — 3, 4(0- reversal), 16 (0-reversal); Picromomys — 6(2), 15(1), 20, 22(0-reversal), 26; Alveojunctus — 12(0-re- versal), 13(2), 15(2). Nodes in tree B are supported by the following synapomorphies: nodes 1 and 2 (same as tree A); node 3—4, 10, 20, 28; node 4 — 2, 6(2), 13(2), 15(2), 18(1), 29; node 5 (Picromomyidae) — 11, 14, 16, 17, 18(2), 19, 27; node 6—23; node 7—18(1), 21, 22; node 8^, 12(2), 19; node 9 (Microsy- opidae) — 3(2), 23(0-reversal), 27; node 10 — 8(2), 16, 17. Taxa whose required autapomorphies differ on tree B: Alveojunctus — 12(0-reversal), 20 (0~reversal); Picromomys — 9, 13(1), 15( 1-reversal?), 26; Tinimomys — 24, 25; Micromomys — 8(0-reversal), 24; Niptomomys — 2, 4(0~reversal), 6, 9, 12( 1-re- versal), 13, 17, 18(2), 29; Navajovius — none; Ignacius — 2, 6(2), 13(2), 14, 21 (0-reversal), 25; Pi- crodus-—9, 18(2), 20, 26. from other plesiadapiforms early in their history (Fox, 1984; Beard and Houde, 1989) and belong to a clade, the Micromomyidae, separate from other families (Beard, 1993; Rose et ah, 1993). This conclusion is corroborated by the cladistic analysis presented here, but formal grouping of Micromomyidae and Picromo- myidae in a higher taxon Micromomyiformes Beard, 1993, is premature from the evidence currently available (see also MacPhee et ah, 1995). Both picromomyid genera were very small plesiadapiforms, and Picromomys was especially diminutive (—10 g), approaching the minimum known size for mammals. Plesiadapiforms are dentally reminiscent of various extant diprotodon- tian marsupials such as petaurids, burramyids, and phalangerids (Cartmill, 1974). In this respect Picromomys and Tinimomys are particularly convergent on the tiny diprotodont Acrobates. The latter also has a hypertrophied and procumbent medial 318 Annals of Carnegie Museum VOL. 65 incisor, behind which are two peg-like teeth, followed by two premolariform teeth with tall trigonids and low-crowned molars with broad talonid basins. Ac rotates is one of the smallest known marsupials, weighing only 12-14 g; it feeds on insects, larvae, and nectar (Nowak, 1991)=^ — a diet constrained by its dental mor- phology and small body size. The same constraints suggest that the diet of Pi~ cromomys was probably very similar. Acknowledgments We thank Drs. K. C. Beard, R. J. Emry, and R Robinson for providing access to specimens and casts used in this study, and Dr. Robinson for supplying locality data that led to the discovery of the quarry at locality D-2035. Vivian Noble and Jay Mussell assisted with SEM photography, and Mary Silcox provided invaluable help with the cladistic analysis. The drawings were expertly prepared by Elaine Kasmer. For their insightful commentaries on the manuscript, which resulted in numerous improvements, we thank Drs. K. C. Beard and G. F. Gunnell, Mary Silcox, and an anonymous reviewer. Fossils of Picromomys were collected under BLM permit 183-WY-PA94. This research was supported in part by NSF grant IBN-94 19776 to KDR. Literature Cited Beard, K. C. 1993. Phylogenetic systematics of the Primatomorpha, with special reference to Der- moptera. Pp. 129-150, in Mammal Phylogeny, Volume 2, Placentals (F. S. Szalay, M. J. Novacek, and M. C. McKenna, eds.). Springer- Verlag, New York. Beard, K. C., and P. Houde. 1989. An unusual assemblage of diminutive plesiadapiforms (Mammalia, ?Primates) from the early Eocene of the Clark’s Fork Basin, Wyoming. Journal of Vertebrate Paleontology, 9:388-399. Bown, T. M. 1982. Geology, paleontology, and correlation of Eocene volcaniclastic rocks, southeast Absaroka range. Hot Springs County, Wyoming. US Geological Survey Professional Paper, 1201- A:l-75. Bown, T. M., and M. J. Kraus. 1987. Integration of channel and floodplain suites, I. Developmental sequence and lateral relations of alluvial paleosols. Journal of Sedimentary Petrology, 57:587- 601. Bown, T. M., and K. D. Rose. 1976. New early Tertiary primates and a reappraisal of some Plesia- dapiformes. Folia Primatologica, 26:109-138. Bown, T M., K. D. Rose, E. L. Simons, and S. L. Wing. 1994. Distribution and stratigraphic correlation of upper Paleocene and lower Eocene fossil mammal and plant localities of the Fort Union, Willwood, and Tatman formations, southern Bighorn Basin, Wyoming. US Geological Survey Professional Paper, 1540:1-103. Cartmill, M. 1974. Rethinking primate origins. Science, 184:436-443. Conroy, G. C. 1987. Problems of body- weight estimation in fossil primates. International Journal of Primatology, 8:115-137. Farris, J. S. 1988. Hennig86 Version 1.5. Program and documentation. Port Jefferson Station, New York (privately produced). Fox, R. C. 1984. The dentition and relationships of the Paleocene primate Micromomys Szalay, with description of a new species. Canadian Journal of Earth Sciences, 21:1262-1267. Gingerich, P. D., B. H. Smith, and K. Rosenberg. 1982. Allometric scaling in the dentition of primates and prediction of body weight from tooth size in fossils. American Journal of Physical Anthropology, 58:81-100. Gunnell, G. F. 1989. Evolutionary history of Microsyopoidea (Mammalia, ?Primates) and the rela- tionship between Plesiadapiformes and Primates. University of Michigan Papers on Paleontology, 27:1-157. MacPhee, R. D. E., K. C. Beard, C. Flemming, and P. Houde. 1995. Petrosal morphology of Tini- momys graybulliensis is plesiadapoid, not microsyopoid. Journal of Vertebrate Paleontology, 15, supplement to no. 3:42 A. Nixon, K. C. 1992. CLADOS version 1.2, IBM PC-compatible character analysis program (privately produced). Nowak, R. M. 1991. Walker’s Mammals of the World. Fifth Edition, Volume 1. Johns Hopkins University Press, Baltimore, Maryland. 642 pp. Robinson, P. 1994. Myrmekomomys, a new genus of micromomyine (Mammalia, ?Microsyopidae) 1996 Rose and Bown — -New plesiadapiform from Wyoming Eocene 319 from the lower Eocene rocks of the Powder River Basie, Wyoming. Contributions to Geology (University of Wyoming), 30:85-90. Rose, K. D., and T. M. Bown. 1982. New plesiadapiform primates from the Eocene of Wyoming and Montana. Journal of Vertebrate Paleontology, 2:63-69. Rose, K. D., K. C. Beard, and P. Houde. 1993. Exceptional new dentitions of the diminutive pie- siadapiforms Tinimomys and Niptomomys (Mammalia), with comments on the upper incisors of Plesiadapiformes. Annals of Carnegie Museum, 62:351-361. SzALAY, E S. 1968. The Picrodontidae, a family of early primates. American Museum Novitates, 2329:1-55. — — . 1969. Uintasoricieae, a new subfamily of early Tertiary mammals (?Primates). American Museum Novitates, 2363:1-36. . 1973. Nev/ Paleocene primates and a diagnosis of the new suborder Paromomyiformes. Folia Primatologica, 19:73-87. . 1974. A new species and genus of early Eocene primate from North America. Folia Pri- matologica, 22:243-250. SzALAY, F. S., AND E. Delson, 1979. Evolutionary History of the Primates. Academic Press, New York, 580 pp. West, R. M„ and M. R. Dawson. 1973. Fossil mammals from the upper part of the Cathedral Bluffs tongue of the Wasatch Formation (early Bridgerian), northern Green River Basin, Wyoming. Contributions to Geology (University of Wyoming), 12:33-41. Appendix 1 Description of Characters Used in Phylogenetic Analysis Characters were polarized with respect to Purgatorius as the outgroup, using 0 as the primitive state and 1 or 2 for derived states. Characters 6, 8, 12, and 18 were treated as ordered; all others are ueordered. Intraspecifically variable traits were coded according to the predominant character state. Autapomorphic char- acters known to be present in only one of the genera analyzed are excluded from the analysis. 1. Ij small to moderate (0); hypertrophied (1). Although incisors have not been described for Purgatorius, alveoli suggest that its medial incisor was relatively less enlarged than in other plesiadapiforms. 2. Ij somewhat inclined (0); strongly procumbent to horizontal (1). 3. Ij laterally compressed (1); laterally compressed and lanceolate (2). The primitive state is unknown; both states observed could be derived, hence no taxa are scored as 0. 4. I2 present (0); absent (1). Scoring of this character is obviously dependent on the homologies of the anterior dentition, which are equivocal in most taxa used here. See Appendix 2 for dental formulae used in this analysis. 5. I3 present (0); absent (1). This character is subject to the same ambiguity as character 4, but relative tooth sizes support the derived state in most plesiadapi- forms. 6. Canine moderately large: alveolus usually >, or crown higher, than I2 and Pi or 2 (0); noticeably reduced, smaller than adjacent teeth (1); absent (2). Same ambiguity as character 4. In addition, relative size is often difficult to judge due to poor preservation. 7. Pj present (0); absent (1). Equivocal in Palenochtha because of conflicting interpretations of the dental formula. 8. P2 two-rooted (0); one-rooted (1); absent (2). Same ambiguity as character 4. 9. P3 two-rooted (0); one-rooted (1). 10. P4 about the size of Mj or smaller (0); larger than Mj (1). 11. P4 talonid shorter than trigonid (0); about as long as trigonid (1). 12. P4 trigonid with distinct paraconid (0); with paracristid with or without 320 Annals of Carnegie Museum VOL. 65 vestigial paraconid (1); with protoconid only, no trace of paraconid or paracristid (2). Ambiguous in Picromomys, owing to wear and/or morphology. 13. P4 talonid narrow (0); broad but without basin (1); broad and with basin (2). 14. P4 cristid obliqua joins trigonid near its midline axis (0); near buccal margin of trigonid (1). Although coded “0,” Paromomys displays a somewhat interme- diate condition. 15. P4 talonid with hypoconid and weak entoconid (0); entoconid absent (1); with enlarged entoconid (2). 16. P4 and molar hypoflexids distinct (0); shallow to absent (1). Characters 13, 14, and 16 are often but not always correlated. 17. P4 and molar trigonids relatively tall and erect (0); lower and strongly anteriorly inclined (1). 18. Molars relatively high crowned (0); of moderate height (1); very low crowned (2). Scoring was based on mean index of M2 trigonid height/length, with three arbitrary divisions: > 0.79 (0), 0.60-0.71 (1), ^ 0.53 (2). 19. Anterobuccal cingulids well developed (0); weak or absent (1). 20. Molar trigonids transversely broad (0); transversely compressed (1). Inter- mediate condition in Palenochtha. 21. Molar paraconids moderately reduced (0); greatly reduced or absent (1). The utility of this character, except for extreme manifestations, is dubious because of its intraspecifically variable and continuous nature. 22. M, mesoconid weak (0); absent (1). Intraspecifically variable in Tinimomys and perhaps others. 23. Mi_2 hypoconulids prominent (0); small or indistinct (1). Ambiguous state in Micromomys due to intermediate expression. 24. Molar talonid cusps and crests peripheral (0); set in from margin as a result of basal inflation (1). 25. M3 third lobe not expanded (0); expanded (1). 26. Enamel smooth (0); crenulated to a variable degree (1). 27. P4 trigonid without metaconid (0); with metaconid (1). 28. Mi_2 hypoconulids central in position (0); distinctly closer to entoconid (1). 29. M,_2 without entoconid notch (0); with distinct notch between hypoconulid and entoconid (1). Variable in Niptomomys examined. 1996 Rose and Bown—New plesiadapiform from Wyoming Eocene 321 Appendix 2 Character Matrix Used for Phylogenetic Analysis Characters deemed too ambiguous to score are indicated by ‘‘a” and were scored as missing data (“?”) for the analysis. At right is the lower dental formula as- sumed in this analysis. Where tooth number varies within a genus, the most primitive known formula is given. Purgatorius 0??00 00000 00000 00000 00000 0000 3-1-4-3 Palaechthon ???01 01100 00000 00000 10000 0100 2-1-3-3 Palenochtha 11101 01100 01000 0000a 00100 0000 2-1-3-3 Picromomys mil 21111 mil 11211 00000 1111 l-0~3-”3 Aiveojunctus 79997 ????! 10212 11210 0?00? ?m ? Paromomys 10?01 01000 01200 00100 11101 1000 2"l-3~3 Ignacius mil 21200 02210 11110 01101 0000 1-0-2-3 Navajovius 10211 01100 02000 00110 11000 0100 l~l-3-3 Micromomys 10111 01001 01000 00001 OOalO 0010 l~-l-3~3 Tinimomys mil 21101 01202 00101 00011 0011 l-0~3-3 Niptomomys 11211 11110 01100 01210 11000 0101 l”l=3-3 Picrodus 10??1 ?1210 02000 11211 11100 1000 ?-?-2“3 INSTRUCTIONS FOR AUTHORS ANNALS OF CARNEGIE MUSEUM consist of con- tributions to the earth sciences, life sciences and anthro- pology, in 30 by 46 picas format (127 by 195 mm or 5 by 7% inches). Submit all manuscripts to the Office of Scientific Publications. 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' /I f7>< a;h- ISSN 0097-4463 ANNALS of CARNEGIE MUSEUM THE CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213 VOLUME 65 27 NOVEMBER 1996 NUMBER 4 CONTENTS ARTICLES Archaeology of Trants, Montserrat. Part 3. Chronological and settlement data James B. Petersen 323 Pipunculidae (Insecta: Diptera) of the Dominican Republic: New records and description of new species J. A. Rafael 363 New genera and new species of Colpurini (Insecta: Heteroptera: Coreidae) fropi Philippines, Borneo, and Malaya.... Harry Brailovsky 383 On the genus Acantholybas Breddin (Insecta: Heteroptera: Coreidae: Col- purini), with description of one new species from Java................. Harry Brailovsky 393 Index to Volume 65 401 Editors, ANNALS, BULLETIN and SPECIAL PUBLICATIONS: J. L. Carter, Ph.D B. C. Livezey, Ph.D D. R. Watters, Ph.D M. A. 'Schmidt, ELS, Assistant Editor Manuscripts, subscriptions, orders for individual numbers, and changes of address should be sent to; Office of Scientific Publications Carnegie Museum of Natural History 4400 Forbes Avenue Pittsburgh, PA 15213-4080 Phone (412) 622-3287 Fax (412) 622-8837 E-mail: scipubs@clpgh.org ANNALS OF CARNEGIE MUSEUM is published quarterly by The Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213-4080, by the authority of the Board of Trustees of Carnegie Institute. © 1996 Carnegie Institute. THE CARNEGIE MUSEUM OF NATURAL HISTORY @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 4, Pp. 323-361 27 November 1996 ARCHAEOLOGY OF TRANTS, MONTSERRAT PART 3. CHRONOLOGICAL AND SETTLEMENT DATA James B. Petersen* Research Associate, Division of Anthropology Abstract Ongoing analyses of the prehistoric Amerindian Trants site in Montserrat include chronological and settlement data obtained from several episodes of field work. Artifact distributions suggest that Trants is larger than most other Saladoid period sites in the West Indies, The radiocarbon dates, artifacts, and stratigraphic associations further suggest that Trants was occupied earlier than many other Saladoid sites and over a long span, ca. 500 b.c.-a.d. 410, or later. Phosphate fractionation and other sediment data confirm the intensity and duration of occupation, and with artifact distributions discussed else- where suggest that the prehistoric settlement had a stable circular ring and plaza configuration. The implications of these data for research on the beginning and development of the Saladoid period in the West Indies and Lowland South America are discussed. Introduction Ongoing analyses of the prehistoric Trants site, located on the island of Mont- serrat in the northern Lesser Antilles, suggest that it is unusual in many respects. Dated as early as ca. 500 b.c., the Trants site documents regional settlement in the broad West Indies by ceramic-making populations much earlier than previ- ously suspected, along with a few other recently studied sites. Moreover, emerging settlement pattern data from Trants document that the earliest groups known as Saladoid, with linkages to the Orinoco River drainage in Venezuela, may have been larger than commonly reconstructed, perhaps including hundreds of residents at a site like Trants. In fact, the size and character of the Amerindian deposits at Trants suggest it was a large, apparently circular village centered on a plaza from the early Saladoid period onward. This article uses chronological and settlement data to help establish the size and configuration of the Trants site, as well as its antiquity and duration. After a brief discussion of relevant field work, radiocarbon dating, and ceramic artifacts, particular emphasis is given to the sediment analyses. Phosphate fractionation and other sediment data corroborate the very intensive and long-term use of the site during the early and later Saladoid period, ca. 500 b.c.-a.d. 410 or later, as es- tablished on the basis of radiocarbon dates and diagnostic ceramics. Although still evolving, these and other data suggest that the circular, ring-shaped Trants site is analogous to some Amerindian settlements in Lowland South America and the insular Caribbean known through the ethnographic record. These findings suggest that past perceptions about the antiquity and scale of Saladoid period develop- ments in the Caribbean and Lowland South America may need to be reevaluated on the basis of research at Trants. ’ Archaeology Research Center, University of Maine at Farmington, Farmington, Maine 04938. Submitted 1 November 1993. 323 324 Annals of Carnegie Museum VOL. 65 ! Fig. 1. — The Trants site on the eastern coast of Montserrat in the northern Lesser Antilles. Trants Site Research The Trants site (MS-Gl) is located on the eastern or windward shore of the small volcanic island of Montserrat in the northern, Leeward Islands portion of the Lesser Antilles in the insular Caribbean (Fig. 1). Montserrat lies about 670 km to the north of mainland South America and 300 km to the east of the Virgin Islands and the more distant Greater Antilles. Thus, Trants is situated in the east- ern portion of the insular Caribbean. The island is relatively high for its small size, extending to about 923 m above sea level and covering an area of about 98 km^ (Watters, 1980). Level areas are rare on the steep volcanic slopes of Mont- serrat and Trants occupies a portion of the only large level area on the windward coast, along with a historic estate of the same name. This landform apparently consists of ancient volcanic deposits, as described further below (Fig. 2, 3). The site lies about 9.1=18.2 m above mean sea level at present. Long before Amerindians of the Saladoid period came to Trants, the island was bom as the product of volcanic activity in the Pliocene=Pleistocene epoch (MacGregor, 1939; Martin-Kaye, 1969; Rea, 1974; Wadge and Isaacs, 1988; Mau- ry et al., 1990). The site proper and the broader landforms on both sides of Farm River are related to early pyroclastic flows from the youngest of five volcanic centers, the Soufriere Hills. Dated to ca. 22,000-14,000 b.c. or earlier, these flow deposits were likely laid down quickly and dramatically in multiple events, wheth- er they represent mud flows or a Pelean type nuee ardente (“glowing cloud”)- Although most of the island’s volcanoes are inactive, the Soufriere Hills remain quite active (MacGregor, 1939; Rea, 1974; Wadge and Isaacs, 1988), as seen most recently in 1995-1996. The Trants archaeological site was first brought to the attention of archaeolo- gists in a publication by Mark Harrington (1924). Harrington published prelimi- nary observations about the private Howes artifact collection from Trants which had been acquired by the Museum of the American Indian (MAI), Heye Foun- 1996 Petersen — Trants Archaeology, 3 325 KEY: SYSTEMATICALLY S UR FAC E - C OL L E C T E D FIELDS WITH SELECTED TEST PITS Fig. 2.-— Overview map of the Trants site showing systematic collection strategies and field designa- tions during the 1990 field work. lOm WIDE COLLECTION CORRIDORS WITH TEST PITS EVERY 50m dation (see Watters and Scaglion, 1994). In particular, the rather substantial Howes shell and stone bead collection led Harrington to conclude that the Trants site was a “West Indian Gem Center,” beginning long-term reference to the site in this light. Unfortunately, Harrington did not visit Montserrat and so he never published full details about the site. In 1978-1979, David Watters was the first archaeologist to conduct systematic field work at Trants as part of his dissertation research (see Watters, 1980, 1994). Over ten years later, Watters and Petersen codirected more extensive testing of the Trants site in 1990 as a joint undertaking of the Carnegie Museum of Natural History (CM) and the University of Maine at Farmington (UMF) (Petersen and 326 Annals of Carnegie Museum VOL. 65 Fig. 3. — General view of the Trants site and existing airport on the eastern coast of Montserrat, facing southeast. Watters, 1991; Watters, 1994). The 1990 field work was done at the request of the Montserrat National Trust given the likelihood of site disturbance by the pro- posed expansion of Blackburne Airport, Montserrat’s only airport (see Fig. 2, 3). A supplemental field inspection was undertaken in 1992 after backhoe “trial bar- row” (or borrow) pits were excavated at the site by airport planners. Various individuals have undertaken analysis of the 1978-1979, 1990, 1992, and MAI (now Smithsonian Institution) collections (see Steadman et ah, 1984; Bartone and Crock, 1991; Crock, 1993; Newsom, 1993; Reitz, 1994; Watters and Scaglion, 1994; Petersen and Watters, 1995; Reitz and Dukes, 1995). Although further field work at the site was undertaken in 1995 by CM and UMF (Petersen et ah, 1995; Watters, 1995), this paper primarily addresses results of the 1978-1979 and 1990 research at the Trants site, although some of the 1995 results are cited, as appro- priate. Details about the 1978-1979 and 1990 Trants field work have been presented elsewhere (see Watters, 1980, 1994; Petersen and Watters, 1991, 1995; Watters and Petersen, 1995) and need not be repeated here, with the exception of a few specifics. Limited surface collection and a single 2.0 m X 2.0 m excavation unit were used to test the site in 1978-1979. Extensive surface collection, 29 0.5 m X 0.5 m test pits and three 1.0 m X 1.0 m test units were used to further sample Trants in 1990 (see Fig. 2). The 1978-1979 and 1990 excavations covered a combined area of only 14.25 m^, but they have provided important artifact and sediment samples from the site. Sediments at the Trants site are quite coarse overall, representing slightly weath- ered sandy deposits from pyroclastic and/or mud flows related to eruption of the Soufriere Hills on the basis of geological research (MacGregor, 1939; Rea, 1974). 1996 Petersen — Trants Archaeology, 3 327 Alternatively, the deposits at Trants have been designated by soil scientists as “protosols,” relatively young developing soils in an “outwash” “delta” of “al- luvium,” or “colluvium,” presumably related to the Farm and Paradise rivers (Lang, 1967:10, 13, 18, fig. II) to the south of the site (see Fig. 2). This latter attribution seems to be contradicted by the local topography, however, which shows the lowermost portion of Farm River Ghaut (near Trants village) deeply crosscutting a long, gently sloping landform extending from the base of the steep Soufriere Hills (near the modern villages of Bethel and Bramble) northeastward to the sea. The Trants stratigraphy is quite variable across the areas sampled, including a combination of relatively fine and very coarse-grained sediments. In the archae- ological excavations, the natural, noncultural strata range horizontally from sandy, rock-free zones to dense, rocky strata, as described below in the discussion of the sediment analyses (Fig. 4”9). This extreme horizontal variability is directly at- tributable to lateral flow variations in the volcanic deposits across the large site area. Within a defined “core” area of the site, both natural and culturally-modified strata were discerned in 1990 and 1992, as well as during the recent 1995 field work. The topography in the local area has been derived from a combination of the original volcanic deposits, later natural processes such as weathering and possible erosion, modification by the prehistoric Amerindian (and historic Euro- pean and African) occupants, and one or more possible natural catastrophes during the time of the Amerindian occupation (Petersen et al., 1995). The extensive surface collection at Trants in 1990 was conducted in two cross- cutting corridors and a larger area elsewhere at this site (see Fig. 2; Watters, 1994; Watters and Petersen, 1995). The 1990 field work established an overall site size of nearly 600,000 m^ on the basis of the full extent of observed and suspected prehistoric artifact distributions. However, the unequivocal “core” area of about 50,000 m^, or about 5.0 hectares, was defined from the 1990 and 1992 field work; this has been revised to an area of about 62,000 m^, or 6.2 hectares, on the basis of the 1995 research. Historic disturbance of the site in the “core” area has largely been confined to shallow hand and limited plow cultivation of sugar cane and cotton. The now abandoned Trants Estate and several existing roads to the estate buildings and Blackburne Airport also have caused localized disturbance to the prehistoric deposits. Thus, the prehistoric site is rather large and relatively little disturbed such that its original configuration can be still determined. Radiocarbon Dating A total of eight radiocarbon dates were first obtained for the Trants site, two from the 1978-1979 testing and six from the 1990 testing (Table 1). These ab- solute dates establish that the site was minimally occupied from ca. 480 b.c. to A.D. 330; an additional 11 radiocarbon dates have recently been obtained from the 1995 research, extending the sequence to ca. a.d. 410 or later (Petersen et al., 1995). Along with another date of ca. 440 b.c. from the Radio Antilles site in Montserrat, two of the Trants dates are among the very oldest reliable published dates known for the Saladoid period anywhere in the West Indies. In fact, only single published dates from the Fond Brule site in Martinique and the Hope Estate site in St. Martin are clearly older than the oldest Trants (and Radio Antilles) dates from Montserrat (Mattioni, 1982; Rouse, 1989; Haviser, 1991). In concert with recent research at Hope Estate and scattered evidence 328 Annals of Carnegie Museum VOL. 65 Fig. 4. — Detail map of the Trants site showing site “core” area in fields 8-11, 1990 test units and other selected excavations, and 1992 “barrow” pits. Note site grid designations along margins. 1996 Petersen-^Trants Archaeology, 3 329 WEST WALL K E Y ■ [aI black (iOYR 2/!) SANDY LOAM - LOAMY SAND [ej] BLACK (7.5 YR 2/0) SANDY LOAM [ii] VERY DARK GRAYiSH BROWN ^ (10 YR 3/2) LOAMY SAND Pel DARK YELLOWISH BROWN (10 YR 4/4) SAND POTTERY SCALE IN METERS Fig. 5.— West wall profile of 1990 test unit N396 E571 in field 8 at the Traots site. Note in situ prehistoric ceramic sherds and location of sediment column 2; lower portion of field stratum B2 was radiocarbon dated to 480 B.c. ± 80. 330 Annals of Carnegie Museum VOL. 65 WEST WALL NORTH WALL N420.5 E644 N42I.5 E644 N42I.5 E645 0.00 M 0.G.S. Ha] brown / DARK BROWN (10 YR 4/3) ' ' LOAMY SAND [¥] YELLOWISH BROWN to DARK YEL- ' ' LOWISH (10 YR 5/4-4/4) LOAMY SAND with PEBBLES andCOBBLES [cTl GRAYISH BROWN (10 YR 5/2) LOAMY SAND BROWN / DARK BROWN (10 YR 4/3) ' ' LOAMY SAND-SANDY LOAM [dI YELLOWISH BROWN to DARK YEL- ' LOWISH BROWN (10 YR 5/4-4/4) LOAMY SAND with PEBBLES and ROCKS IrsI BLACK (10 YR 2/1) SANDY LOAM POTTERY o SCALE IN METERS • 0.10 0.20 • 0.30 L40 Fig. 6. — Composite west and north wall profiles of 1990 test unit N421-N422 E645 in field 10 at the Trants site. Note in situ ceramic sherds, rocks, feature 3, and location of sediment column 3; field stratum Cl was radiocarbon dated to a.d. 210 ± 90, and field stratum C2 was radiocarbon dated to 10 B.c. ± 90. 1996 Petersen — Trants Archaeology, 3 331 Fig. 7.— General view of west wall of 1990 test unit N421-N422 E645 in field 10 at the Trants site. Note tagged field strata and feature 3. elsewhere in the West Indies (e.g., Rouse, 1989), the Trants radiocarbon dates suggest that various estimates of the timing of Saladoid period migration from the mainland into the insular Caribbean have been too conservative, generally thought to have occurred at about a.d. 1 or later (e.g., Willey, 1971; Rouse, 1976; Meggers and Evans, 1978; Rouse and Allaire, 1978; Goodwin, 1979:78; cf. Rouse, 1989, 1992). Acceptance of the beginnings of Saladoid settlement as early as ca. 500 b.c. in the West Indies means that Saladoid populations may have been present for only a relatively brief period in the Orinoco River drainage prior to their colonization of the West Indies. This is especially the case if one accepts the “short” chro- nology for the Saladoid period in Venezuela; that is, beginning at ca. 1000 b.c., rather than 2000 b.c., as has been typically suggested (e.g.. Rouse and Allaire, 1978:441-442; Roosevelt, 1980:193-196; Rouse, 1992:75; cf. Meggers and Evans, 1978; Sanoja and Vargas, 1983). The “short” chronology for the Saladoid period and new dates from the insular Caribbean may support the hypothesis that Saladoid migration commenced in Venezuela due to climatic changes which brought increased aridity (Meggers, 1987), but this is by no means certain. Any number of factors may be responsible for Saladoid dispersal in the insular Caribbean and Lowland South America alike, and better resolution is needed for its dating and relationships all across the broad region. 332 Annals of Carnegie Museum VOL. 65 Fig. 8. — West wall profile of 1990 test unit N596 E571 in field 9 at the Trants site. Note in situ ceramic sherds, rocks, and location of sediment column 1; upper portion of field stratum B was radiocarbon dated to a.d. 60 ± 70, and the upper stratum C was radiocarbon dated to 440 b.c. ± 90. 1996 Petersen — -Trants Archaeology, 3 333 Fig. 9. — General view of west wall of 1990 test unit N596 E571 in field 9 at the Trants site. Note tagged field strata. Trants Amerindian Ceramics Diagnostic prehistoric artifacts from the Trants site, primarily Amerindian ce- ramics, are found in the collections curated by the MAI, from the various episodes of systematic field work, and at the Montserrat National Trust Museum. The goal of the discussion here is to further establish the early and later Saladoid period antiquity of the site, rather than to present exhaustive details about the prehistoric ceramics and other artifacts diagnostic of the Amerindian occupants. Although some of the lithic artifacts are temporally diagnostic as well, especially the lithic beads (Bartone and Crock, 1991; Watters and Scaglion, 1994), the vast majority of the diagnostic artifacts known from Trants are hand-built, open-fired Amerin- Table 1. — Radiocarbon dates for the Trants site. Radiocarbon years b.f Years b.c./a.d. Sample no. Context Depth (in cm b.s.) 2430 ± 80 480 B.C. Beta~44828 N396E571 50-70C 2390 ± 90 440 B.C. Beta-41682 N596E571 70--80C 2140 ± 80 190 B.C. Beta- 18489 1979 unit 40-50 1960 ± 90 10 B.C. Beta-41680 N422E645 70-80C 1890 ± 70 A.D. 60 Beta-41678 N596E571 30-40B 1750 ± 80 A.D. 200 Beta-41679 TP N407E634 40-60B 1740 ± 90 A.D. 210 Beta-41681 N422E645 61C 1620 ± 90 A.D. 330 Beta- 18582 1979 unit 40-50 334 Annals of Carnegie Museum VOL. 65 dian ceramics. More complete details about these ceramics, including stratigraphic and radiocarbon associations, among other issues, are presented elsewhere (Pe- tersen and Watters, 1995), and a complete, exhaustive taxonomy will be presented in the future. Cross dating of the Amerindian ceramics corroborates the available radiocarbon dates for Trants. Such local and regional comparisons generally confirm the place- ment of most, if not all, ceramics to Saladoid period, that is, ca. 500 b.c. to a.d. 600. The Trants ceramics are best related to Cedrosan Saladoid ceramics, as pre- viously defined by Rouse (1992). Alternatively, these ceramics can be assigned to the Early Ceramic period or age (e.g., Haviser, 1991) and some are even at- tributable to the Huecan Saladoid, or Huecoid “culture” (Chanlatte Baik and Narganes Storde, 1990; Rodriguez, \99\a\ Rouse, 1992). This discussion avoids such differentiation, however, given the close technological relationship of all ceramics at Trants (Petersen and Watters, 1995). The available Trants ceramic samples suggest that the entire sequence for the Saladoid period in the Lesser Antilles is represented at Trants. Since most of the systematically recovered ceramics are not decorated (depending on whether solely slipped specimens are included or not, about 1 1=14% of the sample of 2752 sherds excavated from the three 1990 test units are decorated, for example) and are not rim sherds (Petersen and Watters, 1991, 1995), it is possible that some are, in fact, attributable to the post-Saladoid period. However, there is very little, if any, obvious evidence of post-Saladoid occupation in any of the Trants collections; reliable diagnostic artifacts of the post-Saladoid period or Late Ceramic age are not present. The Trants Amerindian ceramics represent a diverse set of vessel forms which closely match those known from elsewhere in the region (e.g.. Rouse, 1974, 1992; Petitjean Roget, 1978; Mattioni, 1982; Chanlatte Baik and Narganes Storde, 1990; Rodriguez, 1991a; Versteeg and Schinkel, 1992). These include, but are not lim- ited to, effigy and noneffigy bowls of various sizes and forms (Fig. 10), cylindrical pots or jars of various sizes (Fig. 11), “incense burners,” and griddles, as well as rare figurines and other forms. Both molded ceramic spindle whorls (in contrast to later post-Saladoid forms which were recycled from broken sherds), and conical and cylindrical ceramic body stamps are also known from the site. Although most of the Trants ceramics are undecorated, some exhibit classic Saladoid types of decoration such as zone-incised crosshatched (ZIC) and white- on-red (WOR) motifs (Fig. 12-15). These two types are quite distinctive, each of which has its own vessel forms as well as distinctive decoration. Both ZIC and WOR are widespread across the site and are found together in intact subsurface deposits. Notably, the Huecan and Cedrosan Saladoid (sensu Rouse, 1992) ceram- ics co-occur in midden deposits at Trants, supporting the close technological sim- ilarity noted above. Thus, it is possible that these are not as different as has been suggested elsewhere (e.g., Chanlatte Baik and Narganes Storde, 1990; Chanlatte Baik, 1991; Haviser, 1991; Rouse, 1992), at least not at the Trants site. Contextual analysis of the Trants ceramic sample from the 1995 excavations should help resolve this matter, as should ongoing work at the Hope Estate site in St. Martin (Haviser, 1991; J. Haviser, C. Hoffman, and M. Hoogland, personal communi- cations, 1993), among other sites. Regardless of this taxonomic issue, evidence of vessel morphology, decoration per se, and rim adomos (Fig. 16) clearly establish the widespread connections between the Trants ceramics and others known from the Lesser Antilles, the Virgin 1996 Petersen — Trants Archaeology, 3 335 Fig. 10. — Saladoid Amerindian ceramic turtle effigy bowl rim sherd from the Trants site. Note zone- incised crosshatched and node decoration, and turtle feet. 336 Annals of Carnegie Museum VOL. 65 Fig. 11. — Saladoid Amerindian ceramic bowl rim sherd from the Trants site. Note node decoration and strap handle. 1996 Petersen — Trants Archaeology, 3 337 Fig, 12. — ^Saladoid Amerindian articulating ceramic rim and neck sherds from the Trants site. Note white-on-red decoration. 338 Annals of Carnegie Museum VOL. 65 Fig. 13. — Saladoid Amerindian ceramic rim, neck, and body sherds from the Trants site. Note white- on-red decoration (all except middle right specimen) and white-and-black-on-red decoration (middle right specimen). 1996 Petersen — Trants Archaeology, 3 339 Fig. 14.— Saladoid Amerindian ceramic rim sherds from the Trants site. Note zone-incised cross- hatched decoration on rim flanges. Islands, and Puerto Rico in the West Indies, as well as the South American main- land (Petersen and Watters, 1995). Montserratian Saladoid potters shared very similar ceramic styles with other potters over a very large area, more than 2500 km in extent, including both their mainland and insular distribution. This social interaction is further corroborated by the lithic raw materials and some of the forms represented among the copious beads known from Trants, as discussed elsewhere (e.g., Bartone and Crock, 1991; Crock, 1993; Watters and Scaglion, 1994). These likewise show connections to other Antillean islands and mainland South America. 340 Annals of Carnegie Museum VOL. 65 Fig. 15. — Saladoid Amerindian ceramic rim sherd from the Trants site. Note zone-incised and node punctate decoration. Trants Sediment Data Sediment analyses can be used to assess the natural and cultural processes which are responsible for the modem configuration of a given archaeological setting (Hassan, 1978). Combining analysis of sediment grain size, for example, with various chemical, pH, and organic matter determinations, among others, may allow evaluation of the natural processes of sediment origin, transport, deposition, and weathering, along with human alteration thereafter through enrichment from refuse disposal and other mechanisms. Although such analyses have been infre- quently conducted in the context of Caribbean and Lowland South American archaeology, they provide an important means of studying site geomorphology and settlement stmcture where they have been undertaken (e.g.. Smith, 1980; Eden et al., 1984; Eidt, 1984; Robinson, 1985). Sediment samples were obtained from all of the subsurface test pits and test units excavated at Trants in 1990 as part of the overall site investigation. Incre- mental 10 cm X 10 cm sediment samples, maximally 10 cm deep within natural 1996 Petersen — ^Trants Archaeology, 3 341 Fig. 16.— Saladoid Amerindian ceramic adomos from the Trants site. Note white-on-red, black-on red, and white-and-black-on-red decoration. 342 Annals of Carnegie Museum vol. 65 Fig. 17. — Collection of sediment column 3 in 1990 test unit N421-N422 E645 at the Trants site. Note rocks removed largely from stratum B. Stratigraphy, were collected from the three 1990 test units, with care taken not to mix natural strata (Fig. 17). Smaller sediment grab samples were removed from the test pits to enable analysis of sediment variability across the extent of the site. Grain size characterization, pH, organic matter, phosphate fractionation, and other chemical analyses were conducted using the professional consulting services of the University of Wisconsin-Milwaukee Sediment Laboratory and the methods of the late Robert Eidt (e.g., 1984, 1985). Of these analyses, the phosphate frac- tionation data were suspected to provide the most objective assessment of past land use at and near the Trants site given the long-term stability of phosphates in archaeological sediments (e.g., Sjoberg, 1976; Eidt, 1985:180-183; Lillios, 1992). Phosphate fractionation values are typically reported as a sum of three inorganic fractions, I-III, representing different combinations of aluminum, iron, and cal- cium; the ratio of fractions II/I can even be used as a relative dating technique, with higher ratios denoting greater antiquity (Eidt, 1984). The three complete sediment columns from the 1990 test units at Trants were completely processed for grain size, pH, organic matter, and chemical analyses; a total of 34 samples were thus analyzed (Table 2, 3). Sediment column numbers 1 and 3 from units N596 E571 and N421-N422 E645, respectively, sampled artifact-rich stratified midden deposits, while column 2 from unit N396 E571 sampled a very notable “anthrosol” composed of dark, organic-rich soil with somewhat fewer obvious subsistence remains (see Fig. 5-9). This anthrosol in the area of unit N396 E571 is much like terra preta, or “black earth,” as known from archaeological sites in the Lowlands of South America (e.g.. Smith, 1980; Eden et al., 1984). Phosphate fractionation analysis was undertaken for only selected increment 1996 Petersen — Trants Archaeology, 3 343 Table 2. — Summary of sediment grain size data for the Trants site. Provenience PN no. % Sand Grain size data % Silt % Clay N596 E571, column sample 1 0-1 OA 2143 73.0 22.0 5.0 10-20A 2144 73.0 22.0 5.0 20-24A 2145 76.0 18.0 6.0 24-30B 2146 74.0 20.0 6.0 30-40B 2147 74.0 21.0 5.0 40-50B 2148 76.0 20.0 4.0 50-60B 2149 76.0 20.0 4.0 60-65B 2150 80.0 16.7 3.3 65-70C 2151 84.0 14.0 2.0 70-80C 2152 85.0 12.0 3.0 80-90D 2153 87.0 10.0 3.0 N396 E571, column sample 2 O-lOA 2160 76.0 19.0 5.0 10-20 A 2161 74.0 21.0 5.0 20-30B 2162 63.0 28.0 9.0 30-40B 2163 61.0 29.0 10.0 40-50B 2164 69.0 22.0 9.0 50-60B 2165 82.0 15.0 3.0 60-70C 2166 87.0 11.0 2.0 70-80C 2167 89.0 11.0 0.0 80-90C 2168 92.0 8.0 0.0 90-100C 2169 94.0 6.0 0.0 100-1 IOC 2170 95.0 5.0 0.0 N422 E645, column sample 3 0-1 OA 2180 76.0 19.0 5.0 10-20A 2181 76.0 19.0 5.0 20-30B 2182 82.0 15.0 3.0 30-40B 2183 84.0 13.0 3.0 40-50B 2184 84.0 13.0 3.0 50-58C 2185 74.0 23.0 3.0 58-60C 2186 75.0 18.0 7.0 60-70C 2187 78.0 16.0 6.0 70-80D 2188 80.0 15.0 5.0 80-90D 2189 84.0 13.0 3.0 90-100D 2190 85.0 13.0 2.0 100-105D 2191 85.0 13.0 2.0 samples from the same three Trants sediment columns and several of the 1990 test pits (Table 4). The test pit samples were selected to cover somewhat the extent of the two sampling transects, east-west and north-south, across the site area, as well as a few essentially “sterile” site margin or off=site areas to enable broad comparison. Watters (1994:table 1) provides a complete list of all excava- tions before the 1995 field work. Eleven phosphate fractionation samples have been analyzed to date, in all cases except two clearly from sub-cultivation zone contexts. Of all the results of the sediment analyses, perhaps most striking are those related to grain size (Fig. 18-20) and phosphate fractionation, the latter described separately below. The three full vertical sediment columns, taken in areas of obvious artifact concentrations, reveal that the macroscopically obvious anthrosol, field strata B1 and B2, in column 2 from test unit N396 E571 (see Fig. 5) has the highest percentages of fine-sized silt and clay relative to other column 2 samples and those from the other two sediment columns (see Table 2, Fig. 18). The deepest portion of the anthrosol in unit N396 E571, stratum B2, has been 344 Annals of Carnegie Museum VOL. 65 Table 3.— Summary of sediment pH, organic matter, and chemical data for the Trants site. Chemical and other data Sol. Salts OM K Ca Mg mho X p Provenience PN no. pH % ppm ppm ppm 10-3 ppm N596 E571, column 0-1 OA 2143 7.0 3.1 147 1476 219 0 5 sample 1 10-20A 2144 7.2 3.1 103 1350 152 0 2 20-24A 2145 7.7 2.8 73 2154 162 0 7 24-30B 2146 7.8 2.8 55 3144 158 0 6 30-40B 2147 7.8 2.9 38 3684 137 0 <0.5 40-50B 2148 7.9 3.0 1380 4182 128 0 <0.5 50-60B 2149 7.9 2.1 1114 3732 149 0 <0.5 60-65B 2150 8.1 1.6 1200 636 125 0 <0.5 65-70C 2151 8.1 2.1 205 636 195 0 <0.5 70-80C 2152 8.2 1.9 141 3246 189 0 0.5 80-90D 2153 8.2 1.7 117 3678 293 0 13 N396 E571, column 0-1 OA 2160 7.8 3.0 298 1512 257 0 6 sample 2 10-20A 2161 7.8 3.3 242 1668 163 0 9 20-30B 2162 7.8 4.6 161 2496 241 0 5 30-40B 2163 7.8 4.2 128 2346 248 0 4 40-50B 2164 7.9 3.1 88 1980 222 0 4 50-60B 2165 7.9 2.0 69 1686 218 0 3 60-70C 2166 8.0 1.8 61 1140 179 0 1 70-80C 2167 7.9 1.9 60 906 167 0 <0.5 80-90C 2168 8.1 1.5 48 642 128 0 1 90-100C 2169 8.0 1.4 67 744 169 0 <0.5 100-1 IOC 2170 8.1 1.7 47 1320 240 0 <0.5 N422 E645, column 0-1 OA 2180 7.9 3.8 70 2280 326 0 5 sample 3 10-20A 2181 8.0 2.5 94 2490 137 0 2 20-30B 2182 8.1 2.0 70 1590 121 0 1 30-40B 2183 8.3 1.7 104 1452 108 0 2 40-50B 2184 8.4 1.4 32 1344 67 0 <0.5 50-5 8C 2185 8.2 2.5 44 3666 126 0 <0.5 58-60C 2186 8.3 1.8 53 3912 142 0 <0.5 60-70C 2187 8.4 2.2 1230 3384 137 0 <0.5 70-80D 2188 8.5 1.9 1238 3234 165 0 <0.5 80-90D 2189 8.5 1.7 67 2310 145 0 <0.5 90-100D 2190 8.5 1.6 56 2487 190 0 <0.5 100-105D 2191 8.5 1.6 67 2526 205 0 7 Table 4. — Summary of sediment phosphate fractionation data for the Trants site. Provenience PN no. Fraction la (ppm) Fraction Ib (ppm) Fraction II (ppm) Fraction III (ppm) Total phosphate (ppm) Ratio II/I N596 E571, 24-30B 2146 17.0 283.0 1,264.0 1,192.0 2,756.0 4.21 N422 E645, 20-30B 2182 14.0 95.0 379.0 251.0 739.0 3.49 N422 E645, 50-5 8C 2185 12.0 292.0 994.0 1,390.0 2,688.0 3.39 N396 E571, O-lOA 2160 52.0 61.0 345.0 381.0 839.0 3.04 N396 E571, 20-30B 2162 37.0 45.0 245.0 323.0 650.0 2.99 T3 PI, 25-30 cm 1672 7.5 1.9 4.5 120.5 134.4 0.47 TP N421 E245, 20 cm 1897 7.3 3.5 13.5 53.9 78.2 1.25 TP N421 E495, 40 cm 1838 46.4 25.7 112.6 185.2 369.9 1.56 TP N421 E745, 30 cm 1879 17.0 7.7 22.5 82.8 130.0 0.91 TP N495 E571, 40 cm 1887 22.1 23.6 100.8 235.5 382.0 2.20 TP N521 E646, 35 cm 2009 9.2 302.4 1,954.5 978.4 3,244.5 6.27 1996 Petersen — Trants Archaeology, 3 345 COLUMN SAMPLE 1 N576 E57i 0.90 100% OF SAMPLE Fig. 18. — Vertical distribution of sediment grain sizes plotted against field strata at the Trants site. 346 Annals of Carnegie Museum VOL. 65 (PPM) 0 20 40 60 80 100 200 400 600 800 1000 2000 3000 4000 KEY: • PHOSPHORUS — — POTASSIUM --MAGNESIUM —CALCIUM Fig. 19. — Vertical distribution of sediment chemical data plotted against field strata at the Trants site. 1996 Petersen — Trants Archaeology, 3 347 COLUMN SAMPLE N576 E5?l COLUMN SAMPLE 2 N396 E57I COLUMN SAMPLE 3 N422 E645 PERCENT ORGANIC MATTER 0 8% PERCENT ORGANIC MATTER PERCENT ORGANIC MATTER Fig. 20.-=Vertical distribution of pH and organic matter data plotted against field strata at the Trants site. 348 Annals of Carnegie Museum VOL. 65 dated to ca. 480 b.c. (see Table 1). The anthrosol is very dark colored and certainly represents sediment alteration caused by the Amerindian occupants. The fine-sized silt and clay fractions increase significantly in the anthrosol over the original sandier sediments, as noted in the difference between samples from a depth of ca. 20-60 cm and those both above and below them (see Fig. 18). This anthrosol area lies at the southern end of the site. The deeper samples in Trants sediment column 2, originating below the an- throsol, or below about 60 cm, represent the original, unaltered site sediments and these are only about 5-13% silt and clay. In contrast, the preserved anthrosol proper, or sediments ca. 20-60 cm deep, includes 18-39% silt and clay, an in- crease of over 700% at most in terms of the percentages of the nonsand fractions (see Table 2). Even the upper 20 cm of sediments in column 2 are generally coarser than the preserved anthrosol, including about 24-26% silt and clay. Nat- ural weathering and soil horizon formation before the Amerindian occupation and the Amerindians themselves all seemingly increased the fine sediments in sedi- ment column 2, but historic cultivation and perhaps other mechanisms have slight- ly coarsened the upper portion relative to the intact anthrosol. The other two Trants sediment columns, both of which sampled artifact-rich middens in test units N422 E645 and N596 E571 (nos. 3 and 1, respectively), also show finer-sized sediments in correlation with the prehistoric middens. How- ever, other factors, natural and/or cultural, have affected their specific configura- tions to some degree (see Table 2, Fig. 18). This is most clearly the case for column 3 in unit N422 E645 which crosscuts a buried midden, field strata Cl and C2, and an upper, historically plow-disturbed midden, field stratum A (see Fig. 6, 7). These include 22-26% and 24% silt and clay for the buried midden and the upper, disturbed midden, respectively. Strata Cl and C2 in unit N422 E645 have been dated to ca. a.d. 210 and 10 b.c, respectively (see Table 1). Sediment column 3 originated towards the eastern end of the site and it is some- what close to the original shoreline. Stratum B overlies strata Cl and C2 in column 3 and it is a largely noncultural deposit of some sort, including a series of large, rounded cobbles, perhaps the result of a storm-surge of water due to a natural catastrophe during the time of the Saladoid occupation (see Fig. 6, 7, 17). It is also possible the strata B deposits resulted from volcanic activity during the Saladoid occupation, but their clustered distribution and proximity to the old shoreline in field 10 (and in field 9, described below) suggests that they are more likely the product of one or more storm-surge events (Petersen et al., 1995). Underlying stratum D presumably represents the original, unaltered site sedi- ments in column 3. The coarsest sediments in column 3 lie between the two middens and beneath the buried midden, including 16-18% silt and clay for the noncultural deposit in stratum B and 15-20% silt and clay for stratum D. Trants sediment column 1 from unit N596 E571 was taken from an area of very dense prehistoric artifacts at the north end of the site, one of the densest midden areas discovered during the 1990 (and more recent) field work (see Fig. 8, 9). In this case, the fractions of nonsand and sand constituents in the midden sediments of column 1 remain more or less constant from the ground surface downwards to a depth of ca. 65 cm, including 20-27% silt and clay in strata A and B, the historic cultivation zone and the intact prehistoric midden, respectively. Although not isolated as a separate stratum, a series of anomalous rock deposits are present in a portion of stratum B in unit N596 E571 as sampled by column 1996 Petersen- — Trants Archaeology, 3 349 1, sometimes including rocks weighing 75 kg or more lying directly on top of artifacts (see Fig. 8, 9). Stratum B in unit N596 E571 has been dated ca. a.d. 60 (see Table 1). Coarser sediments lie deeper still in stratum C of -column 1, con- taining a few prehistoric artifacts and dated 440 b.c. (see Table 1), as well as in noncultural stratum D. These include a range of 13-16% silt -and -clay for the two combined strata. It is difficult to interpret column 1 other than to note some degree of sediment grain size alteration in the upper portion of the column due to the Amerindian occupation, with slight fining of the midden sediments relative to the deeper sediments. The dense prehistoric occupational debris in this area seemingly re- flects intensive activities which may have homogenized the cultural deposits to some degree. Alternatively, the presumed natural disturbance in stratum B reflect- ed by the large rocks, somewhat like stratum B in unit N422 E645, may have coarsened the cultural deposits in part, or perhaps some combination of these and/ or other factors pertains. The rocky deposits may be attributable to one or more storm-surge events during the Saladoid occupation given their proximity to the old shoreline in field 9, much like those in field 10, as described above (Petersen et aL, 1995). The chemical analyses also help to delineate the relative degree of sediment alteration in the sampled portions of the Trants site (Fig. 19; see Table 3). Again, this alteration is due to a combination of cultural and/or natural factors. Available (or exchangeable) calcium (Ca) values peak in all three columns in obvious cor- relation with the Amerindian aethrosols and middens. In the case of column 1 from unit N596 E571, the highest calcium values of 3144-4182 ppm correlate with the preserved Saladoid midden in stratum B, but the deepest and clearly noncultural samples in lower stratum C and stratum D are likewise high at 3246- 3678 ppm. The cultivation-disturbed Amerindian midden in stratum A and the deposits in upper stratum C have lower combined calcium values of 636-2154 ppm. It is likely that the two calcium peaks in column 1 represent cultural en- richment to some degree in stratum B, but in lower stratum C and stratum D the high calcium concentration is perhaps due to natural, presumably geological fac- tors. Percolation downward is negated as an explanation by the low calcium values at the top of stratum C. Calcium data from the other two sediment columns are somewhat different than column 1 in that they each show single peaks, in both cases quite likely related to cultural enrichment. However, in column 2 from unit N396 E571 there is a tendency towards a second peak evident at the very bottom, perhaps reflecting a certain degree of calcium concentration under natural conditions (see Fig. 19). The upper, more obvious calcium, peak correlates with the top of the intact stratum B anthrosol in column 2, wit!) a peak value of 2496 ppm. Other calcium values for the intact anthrosol are 1686-2346 ppm. The cultivation zone, or stratum A, portion of the column has calcium values of 1512-1668 ppm and underlying, sterile stratum C has calcium values of 642-1 140 ppm and peaks at 1320 ppm at the very bottom of column 2. Calcium values in sediment column 3 provide the clearest case of calcium enrichment related to cultural factors. A single strong peak for calcium at 3912 ppm is correlated with the buried midden, or field stratum C2, with other high calcium values of 3384-3666 ppm for strata Cl and C2 as well (see Fig. 19). This buried midden was relatively rich in land crab subsistence -remains and it is likely that these shells contributed to this peak. The cultivation-disturbed upper 350 Annals of Carnegie Museum VOL. 65 midden has calcium values of 2280”-2490 ppm, while largely noncultural and rocky stratum B has calcium values of only 1344-1590 ppm, the lowest for the column; stratum B consists of the possible storm-surge deposits noted above. Once again there is a tentative trend towards calcium concentration at the bottom of column 3 in stratum D, where calcium values of 2310-3234 ppm may reflect percolation downward from the overlying midden and/or some natural concentra- tion. Available potassium (K) peaks similarly to calcium, but in this case only within the middens in columns 1 and 3 from units N596 E571 and N422 E571, respec- tively, and not within the anthrosol of column 2 from unit N396 E571 (see Table 3, Fig. 19). In column 1, a strong potassium peak of 1380 ppm occurs in the middle of the intact stratum B midden paralleling the calcium peak; other potas- sium values in stratum B range from 38-1200 ppm. The disturbed midden in overlying stratum A has potassium values of 73-147 ppm, while underlying strata C and D have combined potassium values of 117-205 ppm. The potassium con- centration in stratum B is thus likely due to cultural factors, but it has seemingly percolated downward to some degree towards the bottom of the midden. Sediment column 2, containing the anthrosol, has its highest potassium values of 242-298 ppm at the surface in the disturbed midden of stratum A, with values that drop from there steadily downward into strata B1 and B2, or the anthrosol, and underlying stratum C; potassium values for these strata combined range from 47-161 ppm. This simple distribution of potassium in column 2 is probably due to Amerindian and/or modern enrichment with relatively little percolation down- ward. In sediment column 3, potassium values exhibit a single clear peak at 1230- 1238 ppm at the bottom of stratum C2, the buried midden, and the top of under- lying stratum D. Much lower potassium values of 70-94 ppm, 32-104 ppm, and 44-53 ppm characterize overlying stratum A, the disturbed upper midden, the largely noncultural rocky deposit in stratum B, and upper stratum C (Cl and the top of C2) respectively, while the deeper deposits of stratum D have potassium values of 56-67 ppm. This distribution suggests that it is the rich cultural deposits of the buried midden which produced the notable potassium peak in stratum C2, with its migration thereafter downward to its interface with deeper stratum D, again directly paralleling the calcium peak in the same column. Available magnesium (Mg) values seem to vary little across the different field strata in all three sediment columns, although these values are somewhat variable through the strata in each case (see Fig. 19, Table 3). In midden-rich column 3, a peak magnesium value of 326 ppm occurs at the surface in the disturbed upper midden of stratum A and drops steadily to a low of 67 ppm at the interface of largely noncultural rocky stratum B and the underlying midden, strata Cl and C2, and then climbs slightly to 205 ppm in the deepest part of the column in stratum D. Likewise, magnesium peaks near the top and the bottom of columns 1 and 2, with a surface value of 219 ppm and a basal value of 293 ppm for column 1, and a surface value of 257 ppm and a basal value of 240 ppm for column 2. Finally, of the other chemical values, available phosphorus (P) is generally very low in all the samples, except basal peaks occur in columns 1 and 3, while soluble salts are nonexistent. Of all three sediment columns, the highest organic matter (OM) values, 4.2- 4.6%, are strongly correlated with intact stratum Bl, the upper anthrosol, beneath the stratum A historic cultivation zone in column 2 from unit N396 E571 (Fig. 1996 Petersen— Tr ANTS Archaeology, 3 351 20; see Table 3). This dark-colored, organic-rich deposit had a notable calcium peak at the same depth, suggesting obvious organic and chemical alteration related to the Amerindian anthrosol (see Fig. 5). Other organic matter values for column 2 include 1.4-3. 3% from above and below stratum Bl. In the two other sediment columns, the highest organic matter values are cor- related with the historic cultivation zone at the surface (see Table 3, Fig. 20). These values, generally 2. 5-3. 8%, are what might be expected from the intentional (or unintentional) historic incorporation of modern vegetal crop residue and other potential organic fertilizers into the site sediments. One secondary organic matter peak of 2.5% occurs at the top of the buried midden in column 3. Most notably, even with such historic enrichment, none of the organic matter values for columns 1 and 3 match or exceed the anthrosol values for column 2. In comparison, an earlier soils study in Montserrat recorded organic matter values of 1.01-1.72% at five localities related to the ''azonal” soils around the Trants Estate, with a mean of 1.49% (Hardy et aL, 1949). Of 29 terra preta sediment samples from the Brazilian Amazon, organic matter values range from 0.45% to 4.70%, with most values less than 2.0% (Smith, 1980:table 1), further reflecting the high relative value of some of the Trants samples. Uniformly high pPI values, 7. 0-8. 5, characterize all three Trants sediment coF umns and in all cases the values increase downward from the surface (see Table 3, Fig. 20). Historic cultivation may have lowered the pH in the uppermost stratum in each case. Curiously, there is a small decrease in pH from largely noncultural rocky stratum B downward to the buried stratum Cl midden in column 3, perhaps again reflecting cultural alteration of some sort. In any case, all of these pH values are neutral to basic and they help account for the superb preservation of faunal remains at the Trants site, at least within the midden areas (Steadman et aL, 1984; Reitz, 1994; Reitz and Dukes, 1995). The Trants pH values are likely high because of a combination of cultural modification due to Amerindian refuse disposal and the natural mildly acidic to basic conditions of the volcanic sediments in Montserrat. Refuse disposal made the sediments even less acidic, building on the favorable conditions naturally inherent in the sediments. In fact, previous soil analyses on the island have es- tablished naturally occurring average pH values of about 6. 0-6.3 to 7.1 island- wide, with an overall average of 6.5 (Hardy et aL, 1949; Lang, 1967). Such relatively nonacidic conditions have been caused by the generally basic character of the bedrock types (Lang, 1967; Rea, 1974). Earlier soil studies have specifically recorded a pH range of 6.6 to 7.2 for five determinations around the Trants Estate, with an average pH value of 6.9 (Hardy et aL, 1949). The results of the phosphate fractionation analyses are the most interesting of all the available sediment data (Fig. 21; see Table 4). The 11 values derived from nine different excavations, including test units and test pits across the site and the site margins, generally confirm the circular ring configuration of the prehistoric artifact distributions (see Fig. 4). Although only a limited number of samples have been analyzed thus far for phosphate fractionation, they document a single peak in intensity along the east-west sampling transect and two peaks along the north- south transect, directly paralleling the Amerindian artifact distributions (Watters, 1994: fig. 8-10; Watters and Petersen, 1995: fig. 4-7). Moreover, the phosphate fractionation data establish a dramatic variation across the site and nearby areas, with low values of 78 ppm and 130 ppm near or beyond the site limits in fields 13 and 10 on the west and east, respectively (see Table 4, 352 Annals of Carnegie Museum VOL. 65 T 1 I ! 1 I T 1 I N 600 I N 400 M 300 P = I30 N 42! E 745 P = 2,756 N 596 E 571 N 52 I E 646 N 42 I E 645 P=382 •H N 495 E57I P-839„., N 396 E 57 I P = 370 KEY: P PHOSPHATE VALUE METERS 0 100 200 300 FEET 1 1 1 J I 1 ■ 1 P = 78 N 42 I E 245 4 , 1 L E 700™ E 600 "■ E 500 — E 400 — E 300- Fig. 21. — Horizontal distribution of phosphate fractionation data plotted on the metric grid at the Trants site. 1996 Petersen — Trants Archaeology, 3 353 Fig. 4, 21). Another low value of 134 ppm is documented for field 2 to the south of the ‘‘core” site area towards Farm River Ghaut. Intermediate values of 370, 382, 650, 739, and 839 ppm characterize the site “core” area (see Fig. 4, 21). Very high phosphate fractionation values of 2756 ppm in sediment column 1 in field 9, 2688 ppm in sediment column 3, and 3245 ppm in a test pit in field 10 document intensive soil alteration by the Amerindian occupants, clearly marking the ring configuration on three sides. Eidt (1984:43) has characterized phosphate values of 10-200 ppm as representing “a range of economic activities such as those from ranching to hack farming,” while values above 200 ppm are indicative of “more intensive activities such as dwelling areas, intensive gardening, manufac- turing, etc.” He goes on to say “extremely high” values “above 2,000 ppm, for example” are representative of “burials, garbage pits, slaughter areas, urbanized zones, etc.” Elsewhere, Eidt (1984:95) notes that such high values are indicative of “intensive residential activities of the type found in Middle East tells or in urbanized zones.” In direct comparison with the values reported here, given use of the same techniques by the same laboratory, the highest of the Trants values also exceed those recently reported for control and archaeological sediments from Bronze Age and Medieval deposits in Portugal (Lillios, 1992:table 1, 2). Another important aspect of the phosphate fractionation data is relative age estimation through comparison of two of the three constituent fractions. Although not very useful at this stage in differentiating the age of the prehistoric Saladoid period deposits from one another, the ratio of phosphate fractions II/I is reflective of gross time differences across the areas sampled. The highest Trants ratio values, 2.99-6.27, are all attributable to the obvious prehistoric midden and anthrosol deposits, with a central plaza ratio of 2.20 (see Table 4). In contrast, the ratios for the samples outside of the “core” area are as low as 0.47 for test pit T3 PI beyond the “core” in field 2 to as high as 1.56 for test pit N421 E495 near (but outside) the ring midden. These latter ratios may be reflective of modem phos- phate enrichment (and relatively low phosphate values) produced primarily through historic farming, while the higher ratios clearly reflect the greater age (and much higher phosphate values) of the prehistoric Saladoid deposits. Thus, the phosphate fractionation data confirm the general site configuration first noted on the basis of Amerindian artifacts. They also provide an absolute means of establishing the intensity of occupation across the site area. It seemingly represents a circular or elliptical midden, approximately 250 m X 200 m in size on the basis of the 1995 field work, with a more or less open central plaza, or “common ground.” Unidentified house stmctures likely lie between the central area and the midden, perhaps like those known from the Golden Rock site in St. Eustatius (Versteeg and Schinkel, 1992). Tentative evidence of a stmcture may have been identified during the 1995 field work at Trants (Petersen et ah, 1995). The chronological data, including matching early radiocarbon dates and ceramic styles, establish that at least the northern and southern portions of the Trants site were occupied contemporaneously. Moreover, the radiocarbon dates and the dense cultural deposits establish a relatively long span of occupation at Trants, at least 800-900 years in duration and possibly longer. It is still unclear if the site rep- resents a continuous Saladoid occupation or various episodes of occupation, but if it was not continuously occupied, one would expect at least partial obliteration of the ring midden from one occupation to another, unless the later occupants took great care to align their settlements with those that came before. Even with continuous occupation, it might be expected that the ring pattern would have been 354 Annals of Carnegie Museum VOL. 65 effaced by reconfiguration of the settlement over time, with house replacement, for example, disturbing the circular settlement and open plaza plan. The long time span of occupation and the strong likelihood that the Amerindian occupants modified the site topography (Petersen and Watters, 1991; Petersen et al., 1995) make the clarity of the preserved ring configuration at Trants all the more remarkable, regardless of whether there was one or more occupations there during the Saladoid period. The Amerindian occupants must have had a very strong cultural imperative which led them to preserve their circular settlement and plaza plan over this long period. Moreover, it seems likely that this imperative was related to maintenance of the open plaza, rather than to the precise positioning of houses and other structures around it. Research Implications of Archaeological and Ethnographic Analogues Habitation sites in the South American Lowlands can be both large and rep- resentative of long-term occupation on the basis of archaeological and ethnograph- ic examples (e.g., Myers, 1973; Smith, 1980; Posey, 1987; Roosevelt, 1991). Among the four major types of ethnographic settlements in the South American Lowlands, as recognized in a typology developed by Myers (1973:243--246), the circular village and plaza configuration with multiple house structures is partic- ularly germane to Trants; the other three types in this typology include single- family and multiple-family single-structure settlements, and multiple-structure set- tlements arranged in a linear or irregular fashion. Alternatively, there are three forms of lowland settlements in another similar classification developed by Roe (1987): the “Linear Riverine” type, the “Oval Upper Tributaries” type, and the “Round Interfluvial” type, correlating with the three gross environmental cate- gories referred to in their designations. The archaeological implications of these ethnographic settlement plans have been hotly debated among regional researchers in the Lowlands in terms of set- tlement size, form, and duration, as well as resident population. Not all scholars are willing to recognize extensive, long-term occupations at some lowland sites, choosing instead to see them as aggregate deposits resulting from repeated small- scale occupations (e.g., Meggers and Evans, 1957, 1978; Evans and Meggers, 1960). In general, archaeological sites in the South American Lowlands are very poorly known in terms of their sizes and spatial configurations. Most lowland sites have been recognized as being smaller than 0. 5-1.0 hectare in extent, al- though larger sites of up to 8.0-9.0 hectares are known (Myers, 1973:tables 2- 4, 7-10; Roosevelt, 1980:tables 16-18). At least a few sites in the Central Amazon region are even larger than 10 hectares in size, and at least one is apparently as large as 80 hectares (Myers, 1973:table 7). The ethnographic circular village and plaza configuration, or “Round Interflu- vial” type, seems more characteristic of certain environments in any case. The savannah grassland environments of central and eastern Brazil are pertinent, for example, where various Ge-speakieg and closely related groups, different cultures in the Upper Xingu region, and the Tupinamba along the Atlantic coast, among others, have had circular plaza-oriented settlements. Seemingly less common in densely forested areas, circular settlements with multiple separate houses are typ- ically divided into secular domestic and public political/ceremonial zones, cor- responding to outer house ring and plaza spaces, respectively; middens and house gardens lie behind the houses on the outer margin of the house ring in most cases (Myers, 1973; Maybury-Lewis, 1979; Novaes, 1983; Costa and Malhano, 1987). 1996 Petersen — Trants Archaeology, 3 355 This teotative environmental linkage may be biased by the incomplete nature of the ethnographic record, however, and the poorly known prehistoric cultures of the Lowlands may have used this configuration over a much broader area. Certainly, circular single-family and circular multiple-family single structures, with open plazas in some of the latter cases, are known in forested areas (Myers, 1973). Moreover, Roe (1987) has suggested that all of the various lowland settle- ment configurations, including the circular village and plaza plan, are linked by a common cosmological model of concentric space related to a basic sexual du- ality or opposition among the Amerindians. In the West Indian archipelago, the ethnographic record is even more poorly known or nonexistent for settlement size, form, and duration across much of the region (Robinson, 1985:G1-G2; Curet, 1992; Keegan, 1992:167). Rouse (1992: 9) notes that Classic Taino houses in Hispaniola known through ethnohistory “were irregularly arranged around a central plaza” and the permanent villages “contained an average of one thousand to two thousand people and ranged in size from a single building to twenty to fifty houses, all made of wood and thatch.” To the east in the Lesser Antilles, Rouse (1992:17-18) suggests that the very poorly known villages of the Eastern Taino were probably like those of the Western Taino; that is, they may have “contained only an estimated 120-225 people, divided among twelve to fifteen houses.” The West Indian archaeological record is not much better for such concerns given the general paucity of broad-scale sampling and large block excavations (but see a few recent exceptions, e.g., Mattioni, 1982; Drewitt, 1991; Rodriguez, 1991^; Siegel, 1991; Curet, 1992; Versteeg and Schinkel, 1992). However, where horizontal excavations have been extensive and/or sampling has been designed to address this issue, the majority of known Lesser Antillean sites are a few hundred to a few thousand square meters in size, with nearly all sites less than 10,000 m^ or 1.0 hectare in extent (e.g.. Rouse, 1976; Watters, 1980; Wilson, 1989; Haviser, 1991; Watters and Petersen, 1992). A few notable Saladoid period sites larger than 1 .0 hectare can be specified for the Lesser Antilles, such as Pearls in Grenada, Indian Creek in Antigua, and Golden Rock in St. Eustatius (Rouse, 1974; Cody, 1991; Versteeg and Schinkel, 1992). Likewise, other Saladoid sites in Puerto Rico (e.g., Siegel 1989, 1991, 1992; Rodriguez, 1991a) are also large, but none of these are clearly larger than Trants. In fact, Trants is larger than most post-Saladoid sites in the region, al- though a few sites may approach or exceed its size on the basis of available information (e.g., Wilson, 1989; Rodriguez, 1992). Thus, several factors about the Trants site at first seem anomalous or unique in the local record, especially for the earliest settlement by ceramic -making Sa- ladoid groups. These include multiple early dates for Trants, its large size, up- wards of 5-6 hectares or more for the “core” area and upwards of 50-60 hectares for the full scatter of artifacts, its configuration as an apparently stable, circular or oval midden (and corresponding unexposed structures) around an open plaza, and the phosphate fractionation and other sediment data which suggest both in- tensive and long-term occupation. At this point, it is obviously premature to put much stock in precise population estimates for Trants, but it seems quite likely that as many as 200-300 individuals may have occupied the site at any one time. Considerably more or perhaps fewer individuals may have been resident at Trants, depending on the size and number of the houses. Although quite large for typical reconstructions of Saladoid groups. 356 Annals of Carnegie Museum VOL. 65 an estimate of 200-300 people is reasonable on the basis of ethnographic infor- mation drawn from the South American Lowlands where circular settlements of similar size are known, as discussed above, along with rare examples of still larger circular sites (e.g., Myers, 1973; Novaes, 1983; Costa and Malhano, 1987; Posey, 1987; Frechione, 1990). In ethnographic examples, the midden almost always lies outside of the circular village consisting of multiple houses, with the central plaza left largely empty. A ceremonial structure, often a men’s house, may occupy the very center of the plaza (e.g., J. Crocker, 1985; Costa and Malhano, 1987; Roe, 1987; W. Crocker, 1990; Heckenberger and Petersen, 1995). A more complicated picture may pertain to West Indian archaeological sites and other, eoncircular settlement forms were likely used too. Nonetheless, circular settlement configurations are locally known or posited for Saladoid period sites in Antigua, St. Eustatius, and Puerto Rico (e.g.. Rouse, 1974; Siegel, 1989, 1991; Rodriguez, 1991a, 1992; Versteeg and Schinkel, 1992). The stable configuration and character of the Trants deposits, including both anthrosols and middens, and the phosphate fractionation data suggest that the site was occupied more or less continuously, as do the chronological data including radiocarbon dates and diagnostic artifacts. Quite obviously, if one assumes such an occupation at the Trants site, no analogues can be posited for the local eth- nographic record, but comparably long or longer occupations are known at other archaeological sites in the insular Caribbean (e.g.. Rouse, 1976; Siegel, 1991; Rodriguez, 1992). Additional excavation, artifact analyses, sediment analyses, and radiocarbon dating will be necessary to verify the inferences about Trants offered here. Summary and Conclusions Several conclusions are based on the available data from the Trants site and attendant interpretations. The first is that recent research at Trants, along with a few other Saladoid period sites in the region, requires reconsideration of the tim- ing and scale of Saladoid settlement of the West Indies. Saladoid groups seem to have come to the Caribbean islands considerably earlier than previously suspect- ed; that is, by ca. 500 b.c. or so, on the basis of evidence from Martinique, Montserrat, and St. Martin. Secondly, Saladoid groups established variably-sized settlements, with some such as Trants being quite large early on and presumably occupied by correspond- ingly large groups of perhaps as many as 200-300 people or more. It is possible that different sizes among the Saladoid sites reflect early establishment of settle- ment hierarchies in the region, but this matter awaits detailed consideration else- where. In any case, not only did the Saladoid settlers occasionally establish large sites, but some were occupied for long periods, over many centuries in various cases. Intensive occupation is documented by dense cultural deposits at different sites, along with newly obtained results of sediment analyses as reported here for Trants. The Trants data suggest a stable settlement configuration, even if the site was not continuously occupied. The remarkable clarity of the ring midden and interior plaza even after 800-900 years or more of Saladoid occupation at Trants suggests that a very strong cultural imperative was operative; the central plaza endured more or less intact in spite of undoubted house reconstruction and other potentially disruptive activities at Trants. 1996 Petersen — Trants Archaeology, 3 357 Thirdly, recently obtained settlement data from sites such as Trants, Golden Rock, and Maisabel, among others, tentatively establish that Saladoid groups brought more than their Lowland South American material culture and aspects of their subsistence practices with them to the West Indies. They also apparently brought traditional settlement patterns and a corresponding cosmology (Hecken- berger and Petersen, 1995). Saladoid settlements almost certainly reflect clear conceptions of secular and public political/ceremonial space, as previously posited for the later Saladoid period Golden Rock and Maisabel sites (Siegel, 1989, 1992; Versteeg and Schinkel, 1992; see also Roe, 1987). Trants suggests that this pattern was present among the earliest Saladoid settlers too. In summary, a full understanding of Saladoid period sites including site struc- ture and related settlement data, among other topics, will only be possible through broad-based research. Refinement of past notions of social structure and com- plexity in West Indian prehistory should be possible very soon. Evidence of re- peated and continuous widespread social interaction, as expressed in common diagnostic artifacts over time, is obvious for the Saladoid period, with linkages from Puerto Rico southward through the Lesser Antilles to mainland South Amer- ica. Likewise, the available evidence suggests very extensive trade over much the span of the West Indian archipelago and a portion of the South American Low- lands during the Saladoid period (e.g., Boomert, 1987; Cody, 1991; Rodriguez, 1991^?; Watters and Scaglion, 1994). This evidence supports the inference that Saladoid social complexity, along with its antiquity, may have been underesti- mated in the past. In fact, much of the social complexity that was to become characteristic of ethnohistoric cultures in the West Indies may well have been brought there by the first Saladoid settlers. Acknowledgments The author extends thanks first and foremost to David Watters who provided an introduction to West Indian prehistory as free of bias as humanly possible. David’s approach can be simply stated: “Here’s all of the literature you want; we don’t know enough to say much about many issues; draw your own conclusions!” I will be always thankful for this wise and appropriate introduction to a very complicated and still understudied field of inquiry. David also introduced me to the Trants site and that’s another reason to give him thanks. The UMF Archaeology Research Center and the Carnegie Museum of Natural History jointly funded recent (pre-1995) field work at the Trants site, visits before and after that time, as well as analyses of previous and recent site collections. At UMF, Belinda Cox prepared the drafted graphics. Bill Crandall and Shirley Thompson helped with manuscript preparation, and Laurie Kidd helped with sediment sample preparation, Fred Dearnley of the UMF Media Center kindly printed the photographs from the author’s negatives and he also printed copies of the drafted graphics. The 1990 field work was con- ducted by Robert Bartone, Ellen Cowie, John Crock, Elizabeth Subin, David and Cathy Watters, and the author. Bob Bartone made a particularly significant contribution in serving as the project coordi- nator for UMF. All the sediment samples were analyzed at the University of Wisconsin-Milwaukee (UW-M) Soils and Physical Geography Laboratory by Bob Brinkman and his staff. As with all their work, the services of the UW-M lab are greatly appreciated and any success gained from this aspect of the overall research is largely due to their expertise. Irving Rouse and two anonymous reviewers, Mary Ann Schmidt, and David Watters all provided useful help and advice during the preparation of this article. An earlier version of this paper was presented in the symposium, “Early Saladoid Period Settlement of the Lesser Antilles: Implications from the Trants Site, Montserrat,” at the 58th annual meeting of the Society for American Archaeology held in St. Louis in 1993. 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Prentice- Hall, Inc., Englewood Cliffs, New Jersey. Wilson, S. M. 1989. The prehistoric settlement pattern of Nevis, West Indies. Journal of Field Archaeology, 16:427-450. ' c V.',. K TV " ■•VT} / t ii ' f •■■ '- • ■ ■» ,r '- » ' ^ - jA^lr **4 ^.i »* V' ' 'm f%,» V ^ ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 4, Pp.. 363-381 27 November 1996 PIPUNCULIDAE (INSECTA: DIPTERA) OF THE DOMINICAN REPUBEIC: NEW RECORDS AND DESCRIPTION OF NEW SPECIES J. A. Rafael^ Abstract The Pipunculidae of the Dominican Republic are reviewed. Keys to genera, subgenera* and species of West Indies Pipunculidae are presented with notes on geographical distribution. Seventeen species in seven genera of Pipunculidae are reported from the Dominican Republic. Eight new species are described and illustrated: Cephalosphaera insuiaris, Cephalops pedemalemis, Eudorylas curvicau- datus, E. dominicanemis, Metadorylas antillemis, M. comutus, M. pilosus, and M. youngi. All Neo- tropical species of the Cephalops iatifrons (Hardy) group are transferred to the genus Micmcephalops De Meyer as new combinations: M. inermus (Hardy), M. Iatifrons (Hardy), M, Yavilateralis (Hardy), M. tramversalis (Rafael), and M. williamsi (Hardy). Eight species are recorded from the Dominican Republic for the first time: Basileunculus aliceae Rafael, Cephalops varius stygius (Hardy), Metador- ylas cressoni Johnson, M. dominicemis Scarbrough and Knutson, Microcephalops Iatifrons (Hardy), M. williamsi (Hardy), Tomosvaryella mexicanensis Ale-Rocha and Rafael, and T. tuberculata Hardy. The previously unknown male of Cephalosphaera mocaensis Hardy and Microcephalops Iatifrons (Hardy) are described for the first time. Introduction Pipunculidae is a poorly represented family in the Dominican Republic fauna. The only previously recorded species is Cephalosphaera mocaensis Hardy, based on a single female specimen (Rafael, 1992). The opportunity to study recently collected material in the collections of the Carnegie Museum of Natural History (CMNH) revealed eight undescribed species, and new distribution records for eight species for that country. It is hoped that this contribution will stimulate interest in collecting and studying this fascinating, though poorly collected, family from the Dominican Republic and elsewhere. Most of the available studied specimens were collected in July, with a few from October and December. Those collected late in the season represented taxa dif- ferent from those collected in July. It is inevitable that additional species and records will be found when more collections are made in other seasons. The material studied was taken from intercept traps, light traps, and sweep samples. Genitalia were studied by removing the abdomen, after which it was placed in a microvial with glycerin and pinned along with the specimen. Taxo- nomic keys including geographical distribution records for the New World were compiled for all the species studied from the West Indies. Primary types are deposited in the collections of CMNH, and representative paratypes and other specimens have been deposited at the Instituto Nacioeal de Pesquisas da Ama- zonia, Manaus, Brazil (INPA). The generic classifications adopted here follows Rafael and De Meyer (1992). Since the pipunculid fauna of the Dominican Republic and of other islands in the West Indies is still largely unknown, it is premature to conduct a zoogeographic ‘ Institute Nacional de Pesquisas da Amazonia (INPA), Caixa Postal 478, 69011-970, Manaus, Ama- zonas, Brazil. 2 Fellow of the Conselho Nacional de Desenvolvimento Cientifico e TecnoMgico. Submitted 24 March 1995. 363 364 Annals of Carnegie Museum VOL. 65 and phylogenetic study at the present time. The following is primarily a taxonomic study and should provide a foundation for future systematic research. Key to Genera and Subgenera of Pipunculidae from the West Indies (Genera not recorded, but of probable occurrence in the West Indies, are also included) 1 Ocellar bristles present; head hemispherical; postcranium narrow; veins M and dm^-cu absent ................................................. Chalarus Walker 1 ' Ocellar bristles absent; head nearly spherical; postcranium swollen; wing venation com- plete ............................................................ 2 2(1') 2' 3(2) 3' 4(3) A' 5(3') 5' 6(5) 6' 7(5') 7' 8(7) 8' 9(2') 9' 10(9) 10' 11(9') 11' 12(11) 12' Proepisternum with fan-like tuft of hairs . . Proepisternum without fan-like tuft of hairs Scutum with conspicuous pilosity anteriorly, dorsocentral hairs not differentiated ...... 4 Scutum without pilosity between rows of dorsocentral hairs ..................... 5 . . Pipunculus Latreille .Parapipunculus Rafael Vein M2 present .................................. Vein M2 present ................................. Cephalosphaera Enderlein 6 Vein M2 absent .................................................... 7 Male abdominal syntergosternite VIII with membranous area reaching epandrium ..... ......................................... C. (Neocephalosphaera) Enderlein Male abdominal syntergosternite VIII with membranous area not reaching epandrium ............................................ C (Cephaiosphera) De Meyer Face equal or subequal in width to lower portion of frons; frons not inflated; flagellum distinctly larger than pedicel ............................... Cephalops Fallen 8 Face narrower than lower portion of frons; frons inflated; flagellum small, slightly larger than pedicel .................................... . Microcephalops De Meyer Male abdominal syntergosternite VIII with membranous area reaching epandrium; ab- domen broad and shortened ....................... C (Semicephalops) De Meyer Male abdominal syntergosternite VIII with membranous area apical; abdomen elongate .................................................. C. {Cephalops) Fallen Pterostigma absent; third section of costa shorter or equal in length to fourth section of costa .......................................................... 10 Pterostigma present; third section of costa longer than fourth section of costa ....... 11 Crossvein r-m located near base of cell dm; abdomen clavate ...... Dorylomorpha Aczel Crossvein r-m located near middle of cell dm; abdomen cylindrical . . Tomosvaryella Aczel Tergite VI visible dorsally in males ..................................... 12 Tergite VI not visible dorsally in males .................................. 13 Apex of flagellum obtuse below; ventral ctenidia present on mesofemur only; tergites with oblique gray pruinose spot posterolaterally ................ .Elmohardyia Rafael Apex of flagellum acuminate to subfiliform; ventral ctenidia present on profemur and mesofemur; tergites without gray pruinose spots ................. Metadorylas Rafael 13(11') Pedicel with many long hairs; abdomen with conspicuous hairs laterally ........... 14 13' 14(13) 14' Pedicel with a few short hairs; abdomen with small inconspicuous hairs . . Eudorylas Aczel Distal section of vein M, straight; aedeagus trifid ................. Allomethus Hardy Distal section of vein Mj curved; aedeagus simple and heavily sclerotized ......... ................................................... BasUeunculus Rafael System actic Accounts Genus Chalarus Walker This genus is cosmopolitan in distribution but has not been recorded from the Dominican Republic. Two species of Chalarus have previously been recorded 1996 Rafael — Pipunculidae 365 from Trinidad (Rafael, 19900). A third species, C. spurius (Fallen) was recorded from Dominica by Scarbrough and Knutson (1989) based on a female specimen. Further studies (Rafael, 1988o, 1990o; Jervis, 1992) indicated that C. spurius represents a species complex of six species, and is restricted to Europe in distri- bution. Records from the New World and other regions (Afrotropical, Palaearctic, Oriental, and Australian) are questionable and therefore this species was not in- cluded in the following key. Key to Species of Chalarus from the West Indies 1. Cell r, closed (Trinidad, Brazil) C. connexus Rafael 1' Cell r, open (Trinidad, Brazil) C. amazonensis Rafael Genus Cephalosphaera Enderlein Cephalosphaera is cosmopolitan in distribution. Recently it was divided by De Meyer (1994) into two subgenera: Cephalosphaera sensu stricto Enderlein and Neocephalosphaera De Meyer. Both are widespread from Canada to Chile in the New World. Key to Species of Cephalosphaera from the West Indies 1 Femora yellow to chestnut, sometimes with brown discoloration (Jamaica, Dominican Repub- lic) C. insularis, new species 1 ' Femora dark brown to black, with bases and apices yellow 2 2(T) Flagellum yellow, pedicel brown; syntergosternite VIII not divided by membranous area; aedeagus short (Dominican Republic) ........................ C. mocaensis (Hardy) 2' Flagellum and pedicel brown to black; syntergosternite VIII divided by membranous area on the dorsum and the venter; aedeagus long, spiraled (Jamaica) . . . C. jamaicensis (Johnson) Cephalosphaera {Cephalosphaera) mocaensis (Hardy) (Fig. 1-6) Dorilas {Cephalosphaera) mocaensis Hardy, 1948Z?:127, fig. 4a, b. Description. — Male: frons and face silvery gray pruinose. Antennae with scape and pedicel dark brown to black; flagellum yellow, long and acute (Fig. 1). Thorax dark brown to black, gray-brown pruinose on the dorsum and gray pruinose on sides. Postpronotal lobe brownish yellow on margins. Propleural fan well developed. Legs with coxae black, trochanters yellow, femora black with base and apex yellow, tibiae and tarsi yellow, except metatarsomeres brown. Wings with third costal section twice as long as fourth and equal in length to fifth. Crossvein r-m located between basal third and basal two-fifths of cell dm. Halteres yellow. Abdomen dark brown to black, brown pruinose, velvety black on bases of tergites III-V and gray pruinose in dorsal view on tergite I and posterolaterally on tergites II-V; gray pruinosity larger on tergite V. Terminalia with apical membranous area on synter- gosternite VIII. Surstyli symmetrical (Fig. 2). Parameres and aedeagus as in Fig. 3, 4. Specimens Examined. — DOMINICAN REPUBLIC: Pedernales, 26 km N Cabo Rojo, 18-06N 71- 38W, 730 m, 13-25 July 1990, L. Masner, J. Rawlins, C. Young, deciduous forest, intercept trap (two males, two females, one female without vein M2); 13-20 July 1990 (one male, one female, INPA); Pedernales, 7 km NE Los Arroyos, 1870 m, 18-16N 71-44W, 15 July 1990, L. Masner, J. Rawlins, C. Young, intercept trap (four females, two females without vein M2); Pedernales, 23.5 km N Cabo Rojo, 18-06N 71-38W, 540 m, 19-25 July 1990, L. Masner, J. Rawlins, C. Young, deciduous forest, intercept trap (one male); 19-21 July 1990 (two males, two females); Pedernales, 26 km N Cabo Rojo, 18-06N 71-38W, 730 m, 26-27 September 1991, R. Davidson, C. Young, S. Thompson, J. Rawlins, wet deciduous forest (one female, INPA, without vein M2); Pedernales, 5 km NE Los Ar- royos, 18-15N 71-45W, 1680 m, 17-18 July 1990, C. Young, J. Rawlins, S. Thompson (one male, INPA); Pedernales, La Abeja, 38 km NNW Cabo Rojo, 18-09N 71-38W, 1250 m, 15 July 1987, J. Rawlins, R. Davidson (one male, INPA); Independencia, 32 km NW La Descubierta, Sierra de Neiba, 1850 m, 1-5 Dec. 1991, L. Masner, S. Peck, mountain forest, Malaise trap (one male, INPA); Bara- 366 Annals of Carnegie Museum VOL. 65 Fig. 1-23. — Pipunculidae from the Dominican Republic. 1-6. Cephalosphaera mocaensis, male: 1, antenna; 2, surstyli; 3, parameres and aedeagus; 4, parameres, ventral view; 5-6, parameres (variation). 7-11. Cephalosphaera insularis, 1, surstyli; 8, inner surstylus, lateral view; 9, parameres and aedeagus; 10, ejaculatory apodeme; 11, ovipositor. 12-17. Cephalops pedemalensis, male: 12, antenna; 13, 1996 Rafael— =PiPUNCULiDAE 367 hoea, 5 km SE Polo, slopes of Loma La Torre, 18“03N 71“16W, 16 July 1992, disturbed forest with coffee, C. Young, R. Davidson, S. Thompson, J. Rawlins (one male). Remarks.— Cephalosphaera mocaensis is the only pipunculid species previ- ously recorded from the Dominican Republic (Hardy, 1966), and the present study shows that it is a frequently collected species. The original description was based on one female specimen, and the following is the first description of the male. This species was previously classified in the subgenus Neocephalosphaera by De Meyer (1994). However, based on the male characters pointed out by De Meyer (1994), this species belongs to the subgenus Cephalosphaera. Cephalosphaera mocaensis is most closely related to C incomitata (Hardy) from Argentina on the basis of characteristics of the male terminalia. It differs from the latter species by the yellow flagellum and by the postpronotal lobe (humeri) being concolorous with scutum. Further study with more specimens from the northern part of South America will likely reveal that C incomitata is a junior synonym of C mocaensis. The male specimens collected from the Dominican Republic show variations in the parameres (Fig. 5, 6). Variation also occurs in wing venation. Vein M2 is absent from the left wing of one male and from both wings of three females. These three female specimens key to the genus Cephalops, but they match the description of Cephalosphaera mocaensis in all respects, except the absence of vein M2. Cephalosphaera {Cephalosphaera) insularis Rafael, new species (Fig. 7^11) DwgiioyA,— This species differs from all other Cephalosphaera species by the following combination of characters: antennae yellow, scutellum with yellow mar- gin, tergites III-V with narrow velvety black bands across the bases, surstyli symmetrical, forcipate, and parameres with a distinct projection at apex. Description.~—M.^e: body length 4.0 mm; wings 4.7 mm. Antennae yellow; flagellum acutum. Eyes joined on the frons for a distance slightly greater than the length of frontal triangle. Front and face silvery gray prainose. Thorax dark brown to black, brown prainose dorsally, gray prainose laterally. Postpronotal lobes yellow, Scutellum yellow at margin, with long bristles. Propleural fan with long, pale bristles. Legs yellow, except coxae brown, metafemor brown on distal half, and all fifth tarsomeres light brown. All femora with ventral ctenidia. Wings slightly iefuscated; pterostigma brown. Third costal section about as long as fourth. Crossvein r-m located near basal one-third of cell dm. Section between cell dm and vein M2 longer than vein dm-cu. Halteres light yellow. Abdomen dark brown to black, brown prainose, with gray praieosity on tergite I and laterally on tergites II-V. When illu- minated at a certain angle tergites III-V with nairow interrapted velvety black bands across the bases. Tergite I with 6-8 bristles laterally. Terminalia with syntergosternite VIII about equal length as tergite V and with distal membranous area. ^ Surstyli (Fig. 7, 8) symmetrical, forcipate. Parameres (Fig. 9) with a distinct projection at apex. Ejaculatory apoderoe as in Fig, 10. Female: body length 3. 6-4.0 mm; wings 4. 2-4.5 mm. Frons and face silvery gray prainose, except froos black near ocellar triangle. Antennae yellow, long, and acute. All coxae brownish yellow. Abdomen brown prainose, except tergites I and II gray prainose, the latter with narrow band brown prainose across posterior margin; tergites III-VI with narrow velvety black band across the bases and gray prainose 00 the sides. Other aspects as in male. Terminalia (Fig. 11) with syntergosternite VII + VIII concolorous with abdomen, epandrium and surstyli; 14, inner surstyius, lateral view; 15, parameres and aedeagus; 16, parameres, ventral view; 17, ejaculatory apodeme, 18-23. Microcephaiops latifrons, male: 18, antenna; 19, ab- domen; 20, terminalia; 21, parameres; 22, aedeagus; 23 ejaculatory apodeme. All figures to same scale as Fig. 1 (== 0,1 mm), except Fig. 11 and 19 (scale = 0.5 mm). 368 Annals of Carnegie Museum VOL. 65 rather cylindrical dorsally, flattened ventrally; ovipositor bright yellow, curved outward and longer than syntergosternite VII + VIII. Type Material— }^o\oiypQ: male (CMNH). Verbatim text of three pin labels: “DOMINICAN REPUBLIC: Pedernales, 26 km N Cabo Rojo, 18^06N 71~38W, 730 m” / “13-25 July 1990, L. Masner, J. Rawlins, C. Young, wet deciduous forest, intercept trap” / “HOLOTYPE Cephalosphaera insularis Rafael [red pa- per]”. Paratype: eight males seven females, topotypic, (two males, three females, INPA); JAMAICA: Portland, just east of Hardwar Gap, 22 Aug. 1980, 1100 m, A. Norrbom (one male). Etymology. — The specific name refers to the island locality where the type series was collected (from Latin insularis = of an island). Remarks. — Cephalosphaera insularis is closely related to C. panamaensis (Har- dy). It differs by the smaller size, yellow antennae, and scutellum yellow at mar- gin. In the terminalia, it differs by the shape of the parameres (which possess a distinct projection at apex in insularis and it is bifid in panamaensis) and by the aedeagus (which is short in insularis and elongated and with a bifid dorsal pro- jection in panamaensis). Cephalosphaera spp. Remarks. — Four female specimens cannot be identified to species and were not associated with any males. Two females have ovipositors curved outward and longer than those of C. panamaensis^ the other two female specimens have straight ovipositors. It is clear that more Cephalosphaera species remain to be described from the Dominican Republic. Specimens Examined. — DOMINICAN REPUBLIC: Pedernales, 23.5 km N Cabo Rojo, 18-06N 71- 38 W, 540 m, 19-25 July 1990, L. Masner, J. Rawlins, C. Young, deciduous forest, intercept trap (three females); Pedernales, La Abeja, 38 km NNW Cabo Rojo, 18-09N 71-38W, 1250 m, 15 July 1987, J. Rawlins, R. Davidson (one female). Genus Cephalops Fallen Cephalops is a cosmopolitan genus and has a wide distribution in the New World, ranging from Canada to Chile. Recently the genus Cephalops was sepa- rated by De Meyer (1994) into four subgenera: Cephalops sensu stricto Fallen, Semicephalops De Meyer, Parabeckerias De Meyer, and Beckerias Aczel. The last two subgenera are not recorded in the New World. Key to Species of Cephalops from the West Indies 1 Flagellum dark brown to black; crossvein r-m located at basal third of cell dm; third section of costa twice as long as fourth; surstyli equal to or longer than epandrium (Dominican Re- public) . C. pedemalensis, new species 1 ' Flagellum yellow; crossvein r-m located near middle of cell dm; third section of costa subequal to fourth; surstyli shorter than epandrium (Jamaica, Dominican Republic) .............. C. varius stygius (Hardy) Cephalops {Cephalops) pedemalensis Rafael, new species (Fig. 12-17) Diagnosis. — This species differs from other described species of Cephalops by the predominantly brown to black antennae and legs, short acute flagellum, and 1996 Rafael — Pipunculidae 369 crossvein r~m at basal third of cell dm. Surstyli are forcipate, subsymmetrical, both curved inward, with the outer one slightly shorter than inner one. Description. — ^(Male only, female unknown.) Body length 3.1 mm; wings 3.5 mm. Frons brown pruinose. Face gray pruinose. Antennae dark brown; flagellum acute (Fig. 12), light brown at margin. Thorax dark brown to black, brown pruinose dorsally, gray pruinose laterally. Propleural fan with five white bristles. Legs predominantly black with trochanters, apex of femora, base and apex of tibiae, and basal tarsomeres yellow. All femora with ventral ctenidia. Wings slightly infuscated; pterostigma brown. Third costal section two times longer than fourth and as long as fifth section. Halteres white, slightly yellow at apex. Abdomen dark brown to black, brown pruinose, with gray pruinosity poster- olaterally; when illuminated at a certain angle, tergites II--V show velvety black bands across bases. Terminalia: syntergosternite VIII with apical membranous area. Surstyli subsymmetrical, curved in- ward (Fig. 13, 14). Parameres and aedeagus as in Fig. 15, 16. Ejaculatory apodeme as in Fig. 17. Type Material— \io\oXypc: male (CMNH). Verbatim text of three pin labels: “DOMINICAN REPUBLIC: Pedemales. 23.5 km N Cabo Rojo, 18-06N 71- 38W” / “26-27 September 1991 J. Rawlins, R. Davidson, C. Young, S. Thomp- son, Wet deciduous forest” / “HOLOTYPE Cephalops pedernalensis Rafael [red paper].” Paratype: one male, Pedemales, La Abeja, 38 km NNW Cabo Rojo, 18- 09N 71~38W, 1250m, 15 July 1987, J. Rawlins, R. Davidson (INPA). Etymology. — The species name refers to the locality Pedemales where the type series was collected. Remarks. — This species is being treated in the subgenus C. {Cephalops) Fallen by the presence of one or more erected spines on the metatibiae median anteriorly and with membranous area not reaching the epandrium. Following the key of Rafael (1991), specimens of this species will run to couplet 16 for C. pauculus and C penepauculus, described from Brazil and Argentina respectively. The new species clearly shows characteristics of Cephalosphaera species in the terminalia and it could well belong to this genus instead of Cephalops. Both of these genera are morphologically very closely related sister groups within the same monophy- letic lineage (Rafael and De Meyer, 1992), and the character vein M2 has been considered as the only character to separate the two genera. Since this character is invalidated to date, pedernalensis is being treated in the genus Cephalops. This should be further studied to clarify the status of these two genera. Cephalops (Semicephalops) varius stygius (Hardy) Dorilas (Dorilas) stygius Hardy, 1948a:5, fig. 4a--c. Remarks. — Cephalops varius stygius possesses the apomorphic characteristic, the membranous area reaching the epandrium, stated by De Meyer (1994) for the subgenus Semicephalops De Meyer. Cephalops stygius was treated as a subspecies of varius with the observation that it might be only a synonym of that species (Rafael, 1991). This is the first record subsequent to its original description from Jamaica. Specimens Examined. — DOMINICAN REPUBLIC: Pedemales, 37 km N Cabo Rojo, 18-09N 71- 35W, 1500 m, 11 July 1987, R. Davidson, J. Rawlins (one female); Pedemales, La Abeja, 38 km NNW Cabo Rojo, 18-09N 71-38W, 1250 m, 15 July 1987, J. Rawlins, R. Davidson (one female, INPA); Pedemales, 9.7 km NE Los Arroyos, 18-16N 71-44W, 2070 m, 15-16 July 1990, J. Rawlins, C. Young, L. Masner (one male); Pedemales, 23.5 km N Cabo Rojo, 18-06N 71-38W, 510 m, 19- 25 July 1990, L. Masner, J. Rawlins, C. Young, deciduous forest, intercept trap (one female, INPA); Pedemales, 1 km S Los Arroyos, 1125 m, 18-14N 71-45W, 18 Oct. 1991, R. Davidson, C. Young, S. Thompson, J. Rawlins, second growth forest (one female). 370 Annals of Carnegie Museum VOL. 65 Cephalops sp. Remarks.— ThttQ female specimens with antennae and legs entirely yellow and abdomen partially yellow cannot be identified to species. No male association can be established with these females. Specimens Examined. — DOMINICAN REPUBLIC: Pedemales, 26 km N Cabo Rojo, 730 m, 18- 06N 71-38W, 26-27 Sept. 1991, R. Davidson, C. Young, S. Thompson, J. Rawlins, wet deciduous forest (one female, INPA); Hato Mayor, Parque Los Haitises, 3 km W Cueva de Arena, 19-04N 69- 29W, 20 m, 7-9 July 1992, R. Davidson, J. Rawlins, S. Thompson, C. Young, mesic lowland forest (two females). Genus Microcephalops De Meyer This genus was established recently by De Meyer (1989) to encompass four Nearctic and one Palaearctic species of small size, face distinctly narrower than the lower portion of the frons, and the flagellum only slightly larger than pedicel. This genus also contains the following Neotropical species treated in the Ce- phalops latifrons group by Rafael (1991). Microcephalops inermus (Hardy), new combination = Dorilas (Eudorylas) inermus Hardy, 1954:26 Microcephalops latifrons (Hardy), new combination = Dorilas {Dorilas) latifrons Hardy, 1948^:1 Microcephalops ravilateralis (Hardy), new combination = Pipunculus (Pipunculus) ravilateralis Hardy, 1965<3:234 Microcephalops transversalis (Rafael), new combination = Pipunculus {Pipunculus) transversalis Hardy, 1965^:22 Microcephalops williamsi (Hardy), new combination = Dorilas {Dorilas) williamsi Hardy, 1954:54 Key to Species of Microcephalops from the West Indies 1 Tergites II-IV velvety dark brown to black, without bands of pruinosity; both surstyli curved inward at apex (Brazil, Colombia, Dominica, Dominican Republic) ....... M. williamsi (Hardy) 1' Tergites II-IV brown pruinose across posterior margin; outer surstylus slightly curved inward at apex (Dominican Republic, Jamaica) ... M. latifrons (Hardy) Microcephalops latifrons (Hardy), new combination (Fig. 18-23) Dorilas {Dorilas) latifrons Hardy, 1948a: 1, fig. la, b. Remarks. — This species is known from the female and has previously been recorded only from Jamaica. It was recharacterized by Rafael (1991). One male fitting the female description apparently belongs here, and is described below. Description. — Male: body length 3.2 mm; wings 3.7 mm. Frons black, brown pruinose, with a small shining black point medially. Face narrower than frons, slightly gray pruinose. Antennae (Fig. 18) with flagellum short, acute. Abdomen (Fig. 19) dark brown to black, velvety black pruinose across bases of tergites II-V; metallic brown pruinose on tergite I and across posterior margin of tergites II- IV; tergite V sub-shining black on posterior half. In other respects as in female description. Terminalia: syntergosternite VIII with apical membranous area. Surstyli asymmetrical (Fig. 20), outer surstyli curved inward distally. Parameres simple (Fig. 21). Aedeagus trifid (Fig. 22) with pair of membranous projections. Ejaculatory apodeme as in Fig. 23. Specimens Examined. — DOMINICAN REPUBLIC: Pedernales, 37 km N Cabo Rojo, 4 km E La Abeja, 18-lON 71-37W, 1440 m, 13-16 July 1987, R. L. Davidson, J. E. Rawlins (one male); Ped- 1996 Rafael — Pipunculidae 371 emales, 23.5 km N Cabo Rojo, 18-06N 71-38W, 540 m, 20 July 1990, C. W. Young, J. E. Rawlins, S. Thompson (one female). Microcephalops williamsi (Hardy), new combination Dorilas (Dorilas) williamsi Hardy, 1954:54, fig. 27a-c. species is widespread across southern Brazil. It is also recorded from Colombia (Rafael, 1991) and Dominica (Scarbrough and Knutson, 1989). Specimens Examined. — DOMINICAN REPUBLIC: Pedernales, La Abeja, 38 km NNW Cabo Rojo, 18-439N 71-38W, 1250 m, 15 July 1987, J. Rawlins, R. Davidson (one male); Barahoea, 9,2 km NW Paraiso, confluence of Rio Nizao and Rio Coltico, 18-03N 71~12W, 230 m, 9-10 Aug. 1990, J. Rawlins, S. Thompson (one male, INPA). Genus Basileunculus Rafael Basileunculus is a Neotropical genus previously recorded from Brazil, El Sab vador, Guyana, Panama, and now the West Indies. Basileunculus aliceae Rafael Basileunculus aliceae Rafael, 1987:630, fig. 11-13. Remarks. — This species was described by Rafael (1987) from the states of Rio de Janeiro and Parana in Brazil. The Dominican Republic specimens represent the first record of the species subsequent to its original description. This is also the only species of the genus Basileunculus from the material examined. The specimens present differ in the following ways from the original description: frons, face, and mesopleuron rather gray prainose; tibiae predominantly black with only base and apex yellow. Specimens Examined. — DOMINICAN REPUBLIC: Pedernales, 37 km N Cabo Rojo, 4 km E La Abeja, 18-lON 71-37W, 1440 m, 13-16 July 1987, R. Davidson, J. Rawlins (one male); Pedernales, 26 km N Cabo Rojo, 18-06N 71~-38W, 13-25 July 1990, L. Masner, J. Rawlins, C. Young, wet deciduous forest, intercept trap (two males, two females, INPA); Pedernales, 5 km NE Los Arroyos, 18-15N 71-45W, 1680 m, 28 July 1990, C. W. Young, J. E. Rawlins, S. Thompson (one female); Elias Pina, Sierra de Neiba at crest, 5.5 km NNW Angel Feliz, 1800 m, 18-41N 71-47W, 15 Oct. 1991, R. Davidson, C. Young, S. Thompson, J. Rawlins, cloud forest (two males); Independencia, 32 km NW La Descubierta, Sierra de Neiba, 1850 m, 01-05 Dec. 1991, L. Masner, S. Peck, mountain forest. Malaise trap (one male, one female). Genus Metadorylas Rafael Metadorylas is limited to the New World. It is best known in the Neotropical region (Rafael, 1990^), and revisionary studies are needed for the Nearctic region. Key to Species of Metadorylas from the West Indies 1 Abdomen predominantly yellow to rufous (Bahamas, Dominican Republic, Jamaica) .... M. cressoni (Johnson) 1' Abdomen dark brown to black 2 2(L) Legs predominantly yellow 3 2' At least femora predominantly dark brown to black 6 3(2) Antennae predominantly yellow to chestnut-brown 4 3' Antennae with scape and pedicel chestnut-brown to black, flagellum yellow . 5 4(3) Parameres with distinct hairs before the recurved processes located near middle; female syntergosternite globose (Dominican Republic) ................ M youngi, new species 372 Annals of Carnegie Museum VOL. 65 4' Parameres without hairs and with processes located at apex not recurved; female synter- gosternite VII + VIII with deep median groove (Dominican Republic, Dominica) ...... M. dominicensis Scarbrough and Knutson 5(3') Abdomen opaque brown to black, brown pruinose; male with a median tubercle on the inflated sternite VI; parameres with two apical recurved processes (Dominican Republic) ................................................ M. comutus, new species. 5' At least tergites III-V shining black, without pruinosity; male sternite VI less inflated and without median tubercle (Mexico, Costa Rica, Grenada, Trinidad, Colombia, Ecuador, Bra- zil, Argentina) M spinosus (Hardy) 6(2') Scape and pedicel brown to black, flagellum yellow; parameres without dorsal recurved processes, only with small subapical spines; outer gonopod not developed (Dominican Re- public) M. antillensis, new species 6' Antennae predominantly dark brown to black; parameres simple with outer gonopod well developed and densely haired (Dominican Republic) ............ M. pilosus, new species Metadorylas youngi Rafael, new species (Fig. 24-29) Diagnosis. — This species differs from all other described Metadorylas species by the following: yellow antennae, postpronotal lobes, halteres, legs, epandrium, and surstyli; profemora and mesofemora with ventral ctenidia; protibiae and me- sotibiae with posteroventral apical bristle; and parameres simple, long bristled with two subapical, recurved processes dorsally. Description. — Male: body length 3.1 mm; wings 3.7 mm. Antennae yellow, flagellum long, acu- minate (Fig. 24). Frons and face gray pruinose. Thorax dark brown to black, brown pruinose dorsally and gray pruinose laterally (mesopleuron and postnotum). Postpronotal lobe yellow. Legs yellow, except fifth tarsomeres brown. Profemora and mesofemora with short ventral ctenidia. Protibiae and mesotibiae with posteroventral bristle. Wings hyaline, except for brown pterostigma; crossvein r-m located near basal one-third of the cell dm; last section of vein M, almost straight. Halteres yellow. Abdomen dark brown to black, brown pruinose, except tergite I dorsally and laterally and tergites II- V posterolaterally with gray pruinosity. Terminalia with large membranous area apically on synter- gosternite VIII; surstyli (Fig. 25) asymmetrical; parameres and aedeagus as in Fig. 26, 27. Female: body length 3.1 mm; wings 3.6 mm. Frons silvery gray pruinose near antennae, becoming white pruinose dorsally and with a shiny black median ridge extended from ocellar triangle to area of larger eye facets. Posteroventral apical bristles of protibiae and mesotibiae longer than in male. Tergites II- V more dense gray pruinose posterolaterally than in male. Terminalia (Fig. 29) with ovipositor yellow, straight, shorter than syntergosternite VII + VIII. Type Material— WoXoiypQ: male (CMNH). Verbatim text of three pin labels: “DOMINICAN REPUBLIC: Pedemales. 26 km N Cabo Rojo. 18-06N 71-38W. 730 m” / “13-25 July 1990 L. Masner, J. Rawlins, C. Young, Wet deciduous forest; intercept trap” / “HOLOTYPE Metadorylas youngi Rafael [red paper].” Paratypes: Thirteen males and four females, topotypic, 15 July 1987, J. Rawlins, R. Davidson (three males, one female, INPA); 21 July 1990, L. Masner; 19-25 July 1990, L. Masner, J. Rawlins, C. Young. Etymology. — This species is named in honor of one of the collectors. Dr. Chen W. Young, Carnegie Museum of Natural History. Fig. 24-33. — Metadorylas from the Dominican Republic. 24-29. Metadorylas youngi, male: 24, an- tenna; 25, terminalia; 26, parameres and aedeagus; 27, parameres, ventral view; 28, surstyli, variation in a paratype; 29, female terminalia. 30-33. Metadorylas cornutus, male: 30, terminalia; 31, parameres and aedeagus; 32, parameres, ventral view; 33, ejaculatory apodeme. All figures to same scale as Fig. 24 (= 0.1 mm), except Fig. 29 (scale = 0.5 mm). 1996 Rafael — Pipunculidae 373 374 Annals of Carnegie Museum VOL. 65 Fig. 34-39. — Metadorylas from the Dominican Republic. 34-36. Metadorylas antillensis, male: 34, terminalia; 35, inner surstylus, lateral view; 36, parameres and aedeagus. 37-39. Metadorylas pilosus, male: 37, antenna; 38, terminalia; 39, parameres and aedeagus, showing outer gonopod. All figures to same scale as Fig. 34 (= 0.1 mm). Remarks. — Metadorylas youngi resembles in most respects the description of M. dominicensis Scarbrough and Knutson, recently described from Dominica. It can be easily separated from M. dominicensis by characters of the terminalia. One male with different surstyli (Fig. 28), one of the most variable characters in pi- punculids, belongs here because it has identical aedeagus and parameres. Metadorylas cornutus Rafael, new species (Fig. 30-33) Diagnosis. — This species differs from all other Metadorylas species by its ex- tremely large stemite VI inflated with a median tubercle, and by its complex parameres, short bristles with two apical recurved processes. Description. — (Male only, female unknown.) Male: Body length 3.8 mm; wings 4.4 mm. Antennae yellow, with scape and pedicel black; flagellum long, acuminate. Frons and face gray pruinose. Thorax dark brown to black, brown pruinose dorsally and gray pruinose laterally (mesopleuron and postno- tum). Postpronotal lobe yellow. Legs yellow, except fifth tarsomeres brown. Profemora and mesofe- mora with short ventral ctenidia. Protibiae and mesotibiae with posteroventral bristle. Wings hyaline, except for brown pterostigma; crossvein r-m located near basal one-third of the cell dm. Halteres yellow. Abdomen dark brown to black, brown pruinose, except tergite I dorsally and laterally and tergites II-V posterolaterally with gray pmieosity. Terminalia (Fig. 30) with sternite VI inflated with median tubercle; syntergosternite VIII with small membranous area; surstyli asymmetrical, the outer with an apical sinus. Parameres and aedeagus as in Fig. 31, 32. Parameres with two recurved processes distally. Ejaculatory apodeme as in Fig. 33. 1996 Rafael=-Pipijnculidae 375 Type Material— Holoiypc: male (CMNH). Verbatim text of three pie labels: ‘'DOMINICAN REPUBLIC: Pedemales. 33 km NE Los Arroyos. 18-15N 71- 45W 1450m” / "16-18 July 1990, L. Maseer, J. Rawlins, C. Young, Wet mourn tain forest, sweep samples” / "HOLOTYPE Metadorylas cornutus Rafael [red paper].” Eiy/wolo^y.—The specific name refers to the two recurved processes at the apex of the parameres in male terminalia (from Latin cornus = horn). Remarks.— Metadorylas cornutus is closely related to the new species M. youn- gi described above. The distinct black scape and pedicel will instantly separate the two species. The male genitalia also provide reliable specific characters. The male of M. cornutus has peculiar terminalia with stemite VI inflated with a median tubercle, the outer surstyli with an apical sinus, and the parameres with two re- curved apical processes. Metadorylas antillensis Rafael, new species (Fig. 34-36) This species differs from all other Metadorylas species by its black coxae, trochanters, and femora, by the extremely large bases of the surstyli, and by parameres being almost straight dorsally with small subapical spines. Description. — (Male only, female unknown.) Body length 3.0 mm; wings 3.6 mm. Antennal scape and pedicel brown, flagellum yellow. Froes with a narrow, dark median ridge. Thorax dark brown to black, brown prainose dorsally and gray prainose laterally on mesopleuron and postnotum. Postproe- otal lobe yellow. Legs with coxae, trochanters black; femora black with base and apex yellow; tibiae and tarsi yellow, except fifth tarsomeres brown. Profemora and mesofemora with short ventral ctenidia. Protibiae and mesotibiae with posteroventral bristle. Wings hyaline, except for brown pterostigma; cross vein r-m located near basal one-third of the cell dm; last section of vein Mj almost straight. Halteres yellow. Abdomen dark brown to black, brown prainose, except tergite I dorsally and laterally and tergites II-V posterolaterally with gray prainosity. Terminalia (Fig. 34) with large membranous area apically on syntergostemite VIII; surstyli asymmetrical, in lateral view as in Fig. 35; parameres and aedeagus as in Fig. 36. Parameres without dorsal processes but with small subapical spines. Type Material.— Holotype: male (CMNH). Verbatim text of three pin labels: "DOMINICAN REPUBLIC: Pedemales. 23.5 km N Cabo Rojo, 18-06N 71- 38W 540 m” / "19-25 July 1990, L. Masner, deciduous forest, intercept trap” / "HOLOTYPE Metadorylas antillensis Rafael [red paper].” Paratype: one male, topotypic, 19-21 July 1990 (INPA). Etymology. — The species name refers to the Antilles. R^iwariy.— The male surstyli of Metadorylas antillensis are superficially similar to M. spinosus (Hardy). It can be separated from that species by the key characters presented above and by the parameres which possess small dorsal subapical spines instead of the dorsal processes present in M. spinosus. Metadorylas cressoni (Johnson) Pipunculus cressoni Johnson, 1919:433. Remarks. — Three specimens (two in poor condition, sex undetermined) may be referred to M. cressoni. The male terminalia are identical to those figured of the holotype (Rafael, 1990^). This species has been recorded previously only from the Bahamas (New Providence) and Jamaica. Specimens Ljcaiwmgrf.— DOMINICAN REPUBLIC: Dajabon, Mariano Cestero, 13 Aug. 1980, 650 376 Annals of Carnegie Museum VOL. 65 A. Norrbom (one specimen); 14 Aug. 1980 (one male); Azua, Padre la Casa, 500 m, 18 Aug, 1980, A. Norrbom (one specimen). Metadorylas dominicensis Scarbrough and Knutson Metadorylas dominicensis Scarbrough and Knutson, 1989:531, fig. 11-15 Remarks.— One female with deep median groove on syntergostemite VII + VIII agrees with the description of M. dominicensis from Dominica. This is the first record of this species subsequent to its original description. Specimens ExammeJ.— DOMINICAN REPUBLIC: Pedemales, 1 km S Los Arroyos, 1125 m, 18- ■14N 71-45W, 18 Oct. 1991, R. Davidson, C. Young, S. Thompson, J. Rawlins, second growth forest (one female). Metadorylas pilosus Rafael, new species (Fig. 37-^39) Diagwoj'fA.— This species differs from all other described Metadorylas species by its well-developed and densely haired outer goeopod, and by the outer sursty- lus being larger than the inner surstylus and having an apical sinus. Description.— only, female unknown.) Body length 2,9 mm; wings 3.2 mm. Antennae (Fig. 37) brown to black, flagellum acuminate. Thorax dark brown to black, brown pruinose on dorsum, gray on the sides. Postpronotal lobes and halteres yellow. Femora predominantly black, profemur without ventral ctenidia; tibiae yellow with small posteroventral bristle on protibiae and mesotibiae. Wings hyaline with brown pterostigma; last section of vein Mj slightly curved; vein CuAj as long as vein dm-cu. Abdomen dark brown to black, brown pruinose, except tergites II-V laterally and pos- terolaterally with gray pminosity. Terminalia: syntergostemite VIII with apical membranous area (Fig. 38) ; outer surstylus with apical sinus and larger than inner surstylus. Parameres and aedeagus as in Fig. 39. Parameres simple; outer gonopod curved, long, thin, and densely haired. Type Materia/.— Holotype: male (CMNH). Verbatim text of three pie labels: ^‘DOMINICAN REPUBLIC: Pedemales. 23.5 km N Cabo Rojo. 18-06N 71- 38W 540 m*’ / ‘‘19-21 July 1990, L. Masner, J. Rawlins, C. Young, Deciduous forest, intercept trap” / “HOLOTYPE Metadorylas pilosus Rafael [red paper].” Paratype: one male, topotypic, 19-25 July 1990 (INPA). Etymology.— The specific name refers to the densely haired gonopod in male terminalia (from Latin pilus — hair). Remarks.— Metadorylas pilosus is the third known species with outer goeopod densely haired. The other two species are M. disgregus (Hardy) described from Argentina and M. gratiosus (Kertesz) described from Paraguay. Metadorylas pi- losus can be separated from both species by the following characters of the teu minalia: outer surstylus larger than inner one; parameres simple; outer gonopod thinner. Genus Eudorylas Aczel Eudorylas is a cosmopolitan genus, and it occurs from Canada to Chile in the New World. Key to Species of Eudorylas from the West Indies 1 Postpronotal lobes dark brown to black, concolorous with scutum .................. 2 1 ' Postpronotal lobes yellow, scutum dark brown to black .......................... 3 2(1) Legs predominantly yellow, except coxae and distal tarsomeres black and brown respec- 1996 R/iFAEL— PiPUNCULIDAE 377 lively; crossvein r-m placed at basal fourth of cell dm (Mexico, Costa Rica, St. Vincent) willistoni (Kertesz) 2’ Legs predominantly black; cross vein r=m placed near middle of cell dm (Dominican Re- public) curvicaudatus, new species 3(1') Combined length of third and fourth costal section longer than fifth; cross vein r-m placed near middle of cell dm; last section of vein M, slightly curved; membranous area of the male terminalia extended (Fig. 46) (Dominican Republic) . . . E. dominicanensis, new species 3" Combined length of third and fourth costal section shorter than fifth; crossveie r-m placed near basal third of ceil dm; last section of vein Mj straight; membranous area of the male terminalia not extended (Puerto Rico, Peru, Brazil) regalis (Curran) Eudoryias dominicanensis Rafael, new species (Fig. 40-43) Diagnosis,— Antcnnm dark brown to black; flagellum long and acuminate; legs predominantly black; mesopleuron gray praieose; crossveie r-m placed near mid- dle of cell dm; membranous area of the male terminalia extended. Description. — -(Male only, female unknown.) Body length 4.1 mm; wings 4.7 mm. Froes and face silvery gray pruinose. Antennae (Fig. 40) brown to black with flagellum long and acuminate. Thorax brown to black, brown pruinose dorsally, gray pruinose laterally, Postpronotal lobe yellow. Legs predominantly black with base and apex of the femora narrowly yellow, base and apex of the tibiae widely yellow and basal two to three tarsomeres yellow. All femora with short ventral ctenidia. Wings slightly infuscated; pterostigma brown; third costal section as long as fourth, both longer than fifth; crossvein r— m placed near middle of cell dm; last section of vein M, slightly curved. Abdomen dark brown to black, brown praieose dorsally, gray pruinose on tergite I and laterally on tergites II- V. Teiminalia with syetergostemite about equal in length to tergite V and with extended down-curved membranous area. Surstyli asymmetrical (Fig. 41). Epandrium narrow. Parameres and aedeagus as in Fig. 42. Ejaculatory apodeme as in Fig. 43. Type Material— Holotype: male (CMNH). Verbatim text of three pin labels: “DOMINICAN REPUBLIC: Pedemales. 7 km NE Los Arroyos. 1870 m, 18- 16N 71-44W” / “15 July 1990 L. Masoer, C. Young, J. Rawlins, intercept trap” / “HOLOTYPE Eudoryias dominicanensis Rafael [red paper].” Etymology. ^ — -The species names refers to the country Dominican Republic. Remarks,— Eudoryias dominicanensis closely resembles E, curvicaudatus in the appearance of the male terminalia. It can be differentiated by the longer fla- gellum and by the less symmetrical surstyli in E. dominicanensis. Eudoryias curvicaudatus Rafael, new species (Fig. 44-48) Antennae dark brown to black; flagellum short and acute; legs black; mesopleuron brown pruinose; cross vein r-m placed near middle of cell dm; membranous area of the male terminalia extended and down-curved distally. Description.— iM.3L\Q only, female unknown.) Body length 3.2 mm; wings 3.7 mm. Frons brown pruinose; face silvery gray pruinose. Antennae (Fig. 44) brown to black with flagellum short, acute, and yellowish under strong illumination. Thorax brown to black, dusted with brown praioosity. Legs black, except apex of the femora and base of the tibiae narrowly yellow. All femora with short inconspicuous ventral ctenidia. Wings slightly infuscated; pterostigma brown; third costal section slightly longer than fourth; third and fourth sections combined slightly longer than the fifth section; crossvein r-m placed near middle of cell dm; last section of vein Mj curved. Abdomen (Fig. 45) dark brown to black, brown pruinose dorsally, light brown pruinose on tergite I and gray pruinose laterally on tergites II- V. Terminalia with syntergostemite about two-thirds length of the tergite V, with ex- tended down-curved membranous area. Surstyli subsymmetrical (Fig. 46). Epandrium narrow. Para- meres and aedeagus as in Fig. 47, Ejaculatory apodeme as in Fig. 48. 378 Annals of Carnegie Museum VOL. 65 Fig. 40-48.- — Eudorylas from the Dominican Republic. 40-43. Eudorylas dominicanensis, male: 40, antenna; 41, epandrium and surstyli; 42, parameres and aedeagus; 43, ejaculatory apodeme. 44-48. Eudorylas curvicaudatus, male: 44, antenna; 45, abdomen; 46, terminalia; 47, parameres and aedeagus; 48, ejaculatory apodeme. Ail figures to same scale as Fig. 40 (= 0.1 mm), except Fig. 45 (scale = 0.5 mm). Type Material— Rololype: male (CMNH). Verbatim text of three pin labels: “DOMINICAN REPUBLIC: Hato Mayor. Parque Los Haitises, 3 km W Cueva de Arena, 19~04N 69-29W” / “20 m, 1^9 July 1992 R. Davidson, J. Rawlins, S. Thompson, C. Young, mesic lowland forest” / “HOLOTYPE Eudorylas curv- icaudatus Rafael [red paper].” Etymology.— The specific name refers to the extended down-curved membranous area in male terminalia (from Latin curvus = curve, caudatus = having tail). Remarks.— Eudorylas curvicaudatus differs from other Neotropical species by the terminalia with an extended, down-curved membranous area and by the sub- symmetrical surstyli. 1996 Rafael — Pipunculidae 379 Eudorylas spp. Remarks.— Eom female specimens representing three different species cannot be identified and were not associated with males. Specimens Examined. — DOMINICAN REPUBLIC: Peravia, Arroyo Canas, 650 m, 08 Aug. 1980, A. Norrbom (one female); Pedernales, 37 km N Cabo Rojo, 4 km E La Abeja, 18-lON 71-37W. 1440 m, 13“15 July 1987, R. Davidson, J. Rawlins (one female); Pedernales, 23.5 km N Cabo Rojo, 18- 06N 71-38W, 540 m, 19-25 July 1990, L. Masner, J. Rawlins, C. Young, deciduous forest, intercept trap (one female); Pedernales, La Abeja, 38 km NNW Cabo Rojo, 18-09N 71-38W, 1250 m, 15 July 1987, I. Rawlins, R. Davidson (one female) Genus Tomosvaryella Aczel Tomosvaryella is a cosmopolitan genus and it has been recorded from Canada to Chile. Seven species have been reported from the West Indies, and six species were observed in this study. The excluded species is T. polita (Williston) de- scribed from St. Vincent. The original description was very brief and generalized and it fits all the described species from the West Indies. Since the type has been lost, no comparison can be conducted to verify the status of this species, therefore it has been excluded from the present study. Key to Species of Tomosvaryella from the West Indies 1 Syntergosternite VIII symmetrical, subhemispherical; metatrochanters with short square- topped process densely white pubescent (USA, Mexico, Belize, Nicaragua, Costa Rica, Panama, Bahamas, Cuba, Puerto Rico, Jamaica, Colombia, Peru, Brazil, Chile) ........ T. subvirescens (Loew) 1 ' Syntergosternite VIII asymmetrical in dorsal view, compressed to right; metatrochanters not as above ........................................................... 2 2(1') Metatrochanters with two spiniform ventral processes (USA, Mexico, Bahamas, Brazil) . . T. bidens (Cresson) 2' Metatrochanters with only one ventral process 3 3(2') Metatrochanters with long process, about as long as width of trochanters; surstyli with apex wider than base; parameres without dorsal spine (Mexico, Costa Rica, Cuba, Haiti, Domin- ican Republic, “Hispaniola,” Jamaica, Dominica, St. Lucia) . . .T. tuberculata Hardy 3' Metatrochanters with short obtuse process or small inconspicuous ridge 4 4(3') Metafemur with sparse long and slender bristles on posterior face; surstyli with apex as wide as base; parameres with one dorsal spine and with one branch of the aedeagus serrate (Mexico, Dominican Republic) T. mexicanensis Ale-Rocha and Rafael 4' Metafemur with short bristles 5 5(4') Surstyli rather simple with bristles on dorsal protuberance, more distinct in lateral view (Mexico, Cuba, Dominica, Colombia) T. scopulata Hardy 5' Surstyli forcipate, distinctly larger at base, without bristles on dorsal protuberance (Puerto Rico, Dominica) T. spangleri Scarbrough and Knutson Tomosvaryella tuberculata Hardy Tomosvaryella tuberculata Hardy, 1948a: 11, fig. 7a-d. Remarks. — ^This species is widespread across Central America and the West Indies, being recorded in Mexico, Costa Rica, Cuba, Dominica, Bahamas, Haiti, Jamaica (Scarbrough and Knutson, 1989; Ale-Rocha and Rafael, 1995) and now Dominican Republic and St. Lucia. Specimens Examined. — DOMINICAN REPUBLIC: Azua, Padre la Casas, 500 m, 18 Aug. 1980, A. Norrbom (one male); Pedernales, 37 km N Cabo Rojo, 18-09N 71-35W, 1500 m, 11 July 1987, 380 Annals of Carnegie Museum VOL, 65 R. Davidson, J. Rawlins (one female); Pedernales, 23.5 km N Cabo Rojo, 18~06N 71-38W, 540 m, 19-25 July 1990, L. Masner, J. Rawlins, C. Young, deciduous forest, intercept trap (one male, INPA). ST. LUCIA: Anse La Raye, Anse Galet, 1 km SSW Anse La Raye, 13-56N 6 1-03 W, 50 m, 21-30 June 1991, J. E. Rawlins, S. A. Thompson (one male). Tomosvaryella mexicanensis Ale-Rocha and Rafael Tomosvaryella mexicanensis Ale-Rocha and Rafael, 1995:416, fig. 29-35. Remarks. — This is the first record of this species subsequent to its original description from Mexico. Specimens Examined. — DOMINICAN REPUBLIC: Azua, Padre la Casas, 500 m, 18 Aug. 1980, A. Norrbom (two males, CMNH; two males, INPA). Genus Elmohardyia Rafael Elmohardyia is a New World genus, better known in the Neotropical region through recent studies (Rafael, 1988Z?). This genus has not been recorded from the Dominican Republic, but two species have been previously recorded from Jamaica and Trinidad (Rafael, 1988^). Key to Species of Elmohardyia from the West Indies 1 Tergite II predominantly gray pruinose; tergites III-V with gray pruinose spots visible in dorsal view; inner surstylus about four times longer than outer one (Jamaica, Costa Rica, Guyana) ... E. gowdeyi (Curran) 1' Tergite II brown pruinose; only tergite V with gray pruinose spot visible on dorsal view; inner surstylus slightly longer than outer one (Trinidad, Brazil) .......... .E. trinidadensis (Hardy) Acknowledgments I thank Dr. Chen W. Young, Section of Invertebrate Zoology, Carnegie Museum of Natural History, for the loan of the specimens and for reviewing an early draft of the manuscript. I also thank two anonymous reviewers for their comments. Literature Cited Ale-Racha, R., and j. a. Rafael. 1995. Tomosvaryella Aczel, do Mexico, America Central e An- tilhas (Diptera: Pipunculidae). Revista Brasileira de Zoologia, 12:405-430. De Meyer, M. 1989. Systematics of the Nearctic species of the genus Cephalops Fallen (Diptera, Pipunculidae). Bulletin de ITnstitut Royal des Sciences Naturelles de Belgique, Entomologie, 59: 99-130. — - — — . 1994. Phylogenetic relationships within the Cephalopsini (Diptera, Pipunculidae). Bulletin et Annales de la Societe Royale Beige d’ Entomologie, 130:7-18. Hardy, D. E. 1948a. Neotropical Dorilaidae (Pipunculidae) studies; Part 1 (Diptera). Psyche, 55:1- 15. — . 1948Z?. New and little known Neotropical Dorilaidae (Pipunculidae-Diptera). Journal of the Kansas Entomological Society, 21:124-133. . 1954. Neotropical Dorilaidae studies, part III. Brazilian species and key to the known species of Dorilas sens. lat. Boletim do Museu Nacional Zoologia, 123:1-60. --- — — . 1965a. The Pipunculidae of Argentina. Acta Zoologica Lilloana, 19:187-241. — — — . \965b. Neotropical Pipunculidae (Diptera) studies, part IV. Further studies of Brazilian spe- cies. Arquivos de Zoologia 4:1-68. — — — . 1966. Family Pipunculidae (Dorilaidae). 45:1-15, in A Catalogue of the Diptera of the Americas South of the United States (N. Papavero, ed.). Museu de Zoologia, Universidade de Sao Paulo, Sao Paulo, Brazil. Jervis, M. A. 1992. A taxonomic revision of the pipunculid fly genus Chalarus Walker, with particular reference to the European fauna. Zoological Journal of the Linnean Society (London), 105:243- 352. 1996 Rafael- — Pipunculidae 381 Johnson, C. W. 1919. A revised list of the Diptera of Jamaica. Bulletin of the American Museum of Natural History, 41:421-449. Rafael, J. A. 1987. Basileunculus, um genero novo de Pipunculidae (Diptera) da regiao neotropical, com chave para as especies. Acta Amazonica, 16/17:627-634. — 1988«. Pipunculidae (Diptera) neotropicais da regiao Neotropical. I. Redescriyao de Chalarus chiiensis Collin, comb. n. e descrigao de duas especies novas da Amazonia. Revista Braxileira de Zoologia, 5:1-9. ■ 1988Z?. Pipunculidae (Diptera) neotropicais do genero Elmohardyia. Acta Amazonica 18: 223-264. 1990a. Revisao das especies neotropicais do genero Chalarus Walker, 1834 (Diptera, Pipun- culidae). Iheringia, Serie Zoologia, 70:45-53. 1990/?. As especies neotropicais de Metadorylas Rafael (Diptera: Pipunculidae). Boletim do Museu Paraense Emilio Goeldi, Serie Zoologia, 6:127-164. — — — . 1991. Revisao das especies neotropicais do genero Cephalops Fallen (Diptera: Pipunculidae). Acta Amazonica, 20:353-390. 1992. A review of the Neotropical species of big-headed flies, genus Cephalosphaera (Dip- tera, Pipunculidae). Pp 631-646, in Insects of Panama and Mesoamerica: Selected Studies (D. Quintero and A. Aiello, eds.). Oxford University Press, Oxford, United Kingdom. Rafael, J. A., and M. De Meyer. 1992. Generic classification of the family Pipunculidae (Diptera): A cladistic analysis. Journal of Natural History, 26:637-658. Scarbrough, A. G., and L. V. Knutson. 1989. Asilidae, Bombyliidae, Conopidae, and Pipunculidae (Diptera) of Dominica, West Indies. Florida Entomologist, 72:519-537. II..- fl. . a:-. . f ^JHPI ■ . ■!5»ii - - .- ■■ i-- • % ■ M- - ^ ■<* ^ > ■!«««»{< ■. *; 'I ti^< - .4 j * ft ^ ^ f 1 1 <0 './ I >3bJUi . ^ - a;, v ^ . (■* .«!,! ■ ■ 7r-> *tj i f> ^ ,‘ ■? i , It -r ;■■!.' ^ • r.*Mr . ■•'■V 5.#' j- ' ! - f ^ ^...:U i. ^ " -;'l^''&i^-'”' >- f' ■ ■ - ^ -. „ _;■ '• jet:* . -4-^ ^ V . V ■■■ ' ^ : .^7}_y‘hJi '. ■■ J I ..'.ii^.» .- i. . V‘ uv. ’’'i - . ,i '* ‘'•V’^'Si J ' ;' .■ i^^ii'lf' ' ‘ '4 . .' ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 4, Pp. 383-391 27 November 1996 NEW GENERA AND NEW SPECIES OF COLPURINI (INSECTA: HETEROPTERA: COREIDAE) FROM PHILIPPINES, BORNEO, AND MALAYA Harry Brailovsky^ Research Associate, Section of Invertebrate Zoology Abstract Three new genera of Colpurini (Coreidae) are described, each containing a single undescribed species: Kinabaluhygia sabah from Sabah (Borneo), Eludohygia parvioculata from Malaya, and Bau- mannhygia insulata from Mindanao (Philippines), Adult dorsal habitus for each are illustrated, and drawings of the male genital capsule are provided. Introduction The tribe Colpurini is represented in the South Pacific Islands by a number of species that show various bizarre morphological specialization. The most striking features of these genera are the remarkable reduction of the hemelytra, the de^ velopment of the head including the eyes, and the high degree of endemicity. The three genera described at this time complement a series of papers prepared by the author to clarify the systematics of the tribe (Brailovsky 1990, 1993a, 1993 A 1993c; Brailovsky et aL, 1992; Brailovsky and Ortega Leon, 1994). The following abbreviations are used for the institutions cited in this paper: BMNH, The Natural History Museum, London, United Kingdom; FMNH, Field Museum of Natural History, Chicago, Illinois; QMBA, Queensland Museum, Brisbane, Australia; UNAM, Coleccion Entomologica del Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico D. E, Mexico. Systematic Entomology Kinabaluhygia, new genus (Fig. 1-4) Diagnosis.— Kinabaluhygia, n. gen., is the only genus within the tribe Colpurini that exhibits a conspicuous globosity on the side of the head in front of and close to the eyes, with the apex truncated. This new genus closely resembles Wygohygia Brailovsky (1993) in having the eyes small and protruding on short stalks, head quadrate, ocelli present, buccula with short anterior projection, postocular tubercle protuberant, tylus unarmed, scu^ tellum flattened, and plica and fissura on abdominal sternite VII of the female. It differs from other Colpurini in a number of important characteristics: sides of head in front of and close to eyes conspicuously globose, raised above eyes and apically truncated; hemelytra brachypterous, with hemelytral membrane re- duced; posterior lobe of metathoracic peritreme small and globose; callar region uniformly and slightly convex; posterior pronotal lobe near middle third flat; and anteneiferous tubercle unarmed. In Wygohygia, the sides of head in front of eyes ‘ Institute de Biologia, UNAM, Depto. de Zoologia, Apdo, Postal 70153, Mexico 04510 D. F. Mexico. Submitted 31 March 1995. 383 384 Annals of Carnegie Museum VOL. 65 are laterally globose, never raised above eyes, and apically projected on a long and robust lobe; the hemelytra are submacropterous with the hemelytral mem- brane well developed with few veins furcate; posterior lobe of the metathoracic peritreme small and sharp, callar region transversely convex and separated along midline by slightly longitudinal depression; posterior pronotal lobe near middle third with two lateral and irregular convexities separated by longitudinal depres- sion; antenniferous tubercle armed with a long robust spine. Fig. 2-6. — Colpurini. Fig. 2-4, Kinabaluhygia sabah Brailovsky; 2, female genital plates, caudal view; 3, female genital plates, lateral view; 4, male genital capsule, caudal view; 5, Baumannhygia insulata Brailovsky, male genital capsule, caudal view; 6, Eludohygia parvioculata Brailovsky, male genital capsule, caudal view. Brailovsky— New Colpurmi 386 Annals of Carnegie Museum VOL, 65 Wygohygia, known only from New Guinea, is much more robust and longer than 12 mm (female; male unknown) and Kinabaluhygia, recorded from Borneo, is shorter, less than 10.50 mm. Description. — Head: quadrate, wider than long, sunken between lateral lobes; tylus unarmed, api- cally globose, extending anteriorly to the jugae and more raised in lateral view; jugae unarmed; sides of head in front of and close to eyes globose, elevated, conspicuously convex, and raised above eyes; antennal segment I robust, thickest, slightly curved outward, and shorter than head; segments 11 and III cylindrical and slender; segment IV fusiform; segment II longest; segment IV the shorter; segment I longer than III (male) or shorter than III (female); ocelli reduced, not elevated, and difficult to see; preocellar pit weakly deep; eyes small, pimple-like, protruding on short stalks; postocular tubercles protuberant; vertex and frons sunken and flattened; buccula rounded, short, elevated, not projecting beyond antenniferous tubercle, with a short anterior projection; rostrum long, reaching posterior third of abdominal sternite V (male) or anterior third of VI (female); rostral segment I reaching posterior gular region; mandibular plate absent. Thorax: pronotum wider than long, trapeziform, nondeclivent; collar wide; frontal angles projecting forward as rounded thickened lobes; humeral angles rounded, not exposed and slightly elevated; anterolateral borders weak and obliquely straight; posterolateral and posterior borders straight; callar region slightly convex; posterior margin with two lateral convexities. Anterior lobe of metathoracic peritreme reniform, posterior lobe small, subglobose. Legs: femora unarmed; tibiae sulcate. Scutellum: longer than wide (female) or as long as wide (male); triangular, with subacute apex; disc flat. Hemeiytra: brachypterous, reaching median third of abdominal segment IV; clavus and corium fused into a coriaceous pad, and the wings meeting each other along the midline; hemelytral membrane reduced. Abdomen: connexival segments strongly elevated, higher than abdominal segments; angle of connexival complete; abdominal stemites with the medial furrow extending to posterior border of stemite V. Integument: body surface mostly shining, with short decumbent, silvery bristle-like setae, intermixed with long, erect setae located on the abdominal sterna. Head, pronotum, scutellum, hemeiytra, thorax, abdomen, and exposed parts of genital segments punctate. Head with frons and vertex and connexival segments practically smooth; antennal segments and legs covered by long and short setae. Male genitalia: genital capsule: postero ventral border with lateral diverging spine-like projections, medial process bilobed, and with submedian truncated lobes (Fig. 4). Female genitalia: abdominal sternite VII with plica and fissura; plica triangular reaching median third of sternite VII; gonocoxae I squarish, large; paratergite VIII short, subtriangular, with spiracle visible; paratergite IX square, longer than paratergite VIII (Fig. 2, 3). Distribution.--=Known only from Borneo. Etymology. — Named for its occurrence on Mt. Kinabalu (Borneo). Gender feminine. Type Species.- — -Kinabaluhygia sabah, n. sp. Kinabaluhygia sabah, new species (Fig. 1--4) Description. — -Dorsal coloration, male: head, pronotum, and scutellum bright black with pale yellow antenniferous tubercle and apex of scutellum; antennal segment I black, segments II and III dark red with basal and apical joint yellow, and IV yellow-orange with basal joint dark red; clavus, corium, connexival segments, and abdominal segments bright orange-chestnut tinged with red, except the posterior margin of the connexival II to VII ocher; hemelytral membrane dirty chestnut-orange. Ventral coloration, male: head and thorax bright black tinged with red on acetabulae; rostral segments I to IV, and anterior and posterior lobes of metathoracic peritreme bright pale orange with chestnut reflections; coxae dark chestnut-orange; trochanters yellow; femora bright dark red with basal joint yellow; tibiae bright dark red with one or two pale yellow-orange rings; tarsi bright yellow-orange; abdominal sternites and pleural margins bright brownish red except the posterior angle of pleural margins III to VII ocher; genital capsule black with bright brownish red reflections. Color, female: similar to male. Legs: tarsal segments I and III dark red with chestnut-orange re- flections; segment II mostly bright yellow-orange. Genital plates: gonocoxae I black with red reflec- tions except the external upper angle creamy yellow; paratergite VIII and IX black with red reflections. 1996 Brailovsky — New Colpurini 387 Measurements. — Male, female (mm). Head length 1.72, 1.72; width across eyes 1.88, 1.88; inter- ocular space 1.32, 1.32; interocellar space 0.48, 0.48; preocular distance 1.30, 1.34; length antennal segments: I, 1.52, 1.41; II, 2.16, 2.20; III, 1.48, 1.44; IV, 1.32, 1.30. Pronotum: total length 1.72, 1.92; width across frontal angles 1.68, 1.64; width across humeral angles 2.64, 2.60. Scutellar length 1.04, 1.02; width 1.04, 1.00. Total body length 10.10, 10.40. Type Specimens. — Holotype, male (QMBA): BORNEO, SABAH, Park Head- quarters, Mt. Kinabalu, 5000 ft, 24-26 December 1975, G. B. and S. R. Monteith. Paratype, one female (UNAM): same data as holotype. Etymology. — Named after the type locality. Eludohygia, new genus (Fig. 6, 7) Diagnosis.— Eludohygia, n. gen., is a medium-sized striking insect easily dis- tinguished from all other Colpurini by the pronounced dorsal convexity of the head, the minute eyes occupying only a small portion of the head surface (Fig. 7), tylus conspicuously enlarged, postocular tubercle weakly developed, buccula entirely without external teeth and femora armed with two rows of spines. Description. — Male. Head; pentagonal, longer than wide across eyes, remarkably convex dorsally; tylus conspicuously enlarged, unarmed, apically globose, extending anteriorly to the jugae, and more raised in lateral view; jugae unarmed, thickened and extended to middle third of tylus; antenniferous tubercle unarmed; sides of head in front of eyes unarmed and straight; antennal segment I robust, thickened, slightly curved outward and shorter than head; segments II to IV mutilated; ocelli absent; preocellar pit deep; eyes minute, occupying only a small portion of the head surface; postocular tubercles developed but not protuberant; buccula rounded, short, elevated, not projecting beyond an- tenniferous tubercles, without teeth; rostrum long, reaching posterior third of abdominal sternite VI; mandibular plate absent. Thorax: pronotum trapeziform, wider than long, nondeclivent; collar wide; frontal angles produced forward as rounded and globosus lobes; humeral angles rounded, not exposed; anterolateral borders obliquely straight, emarginated, and raised above pronotal disc; posterolateral border straight and posterior border slightly concave; pronotal disc with anterior half slightly concave and posterior half flat. Anterior lobe of metathoracic peritreme reniform, posterior lobe sharp, small. Legs: femora armed with two rows of robust spines along ventral surface; tibiae with sulcus difficult to see. Scutellum: wider than long, triangular, with sharp apex; disc flat. Hemelytra: brachypterous, reaching posterior third of abdominal segment III; clavus and corium fused into a coriaceous pad and the wings meeting each other along the midline; hemelytral membrane reduced. Abdomen: connexival segments strongly elevated, with posterior angle complete, not produced into spines; abdominal sternites with the medial furrow extending to posterior third of sternite V. Integument: body surface mostly dull, with head, anterior third of pronotum, thorax, and abdomen shining. Head, thorax, abdomen, and genital capsule strongly punctate; pronotum, scutellum, hemel- ytra, dorsal abdominal segments, and connexival with scattered punctures. Body with short decumbent silvery bristle-like setae, intermixed with long erect setae located on the abdominal sterna and legs. Male genitalia: genital capsule: posteroventral border complete, laterally enclosed by two medium- sized lobes (Fig. 6). Female. Unknown. Distribution. — Known only from Malaya. Etymology. — From the latin eludo = escape, avoid, or elude. Gender feminine. Type Species. — Eludohygia parvioculata, n. sp. Eludohygia parvioculata, new species (Fig. 6, 7) Description. — Male. Dorsal coloration; head bright black; antennal segment I pale chestnut-orange; pronotum with anterior half shining black and posterior half dull black with dark red reflections; 388 Annals of Carnegie Museum VOL. 65 anterolateral margin chestnut-orange; scutellum, hemelytra, connexival segments and dorsal abdominal segments dark red with posterior border of connexival ocher-yellow; hemelytral membrane dark chest- nut-orange. Ventral coloration: head bright black with dark red reflections; buccula (except basal third dark red) and rostral segments I to IV pale chestnut-orange-yellow; thorax and abdominal sternite bright orange-red with following areas pale chestnut-yellow-orange: anterior and posterior lobe of metathoracic peritreme, posterior margin of metapleura, acetabulae, posterior border of pleural seg- ments III to VII, and legs. Measurements. — Male (mm). Head length 1.84; width across eyes 1.68; interocular space 1.20; preocular distance 1.36; length antennal segment: I, 1.28 (segments II-IV mutilated). Pronotum: total length 1.60, width across frontal angles 1.76; width across humeral angles 2.76. Scutellar length LOO; width 1.32. Total body length 10.00. Type Specimen.— VLoloiypQ, male (BMNH): MALAYA, BUKIT PALAS, Cam- eron Highlands, 5400 ft, 5 May 1939, N. C. E. Miller. Etymology. — Referring to the appearance of the eyes in lateral view. Baumannhygia, new genus (Fig. 5, 8) Diagnosis. — The reduction of wings, the membrane absent, the prominent post- ocular tubercle, the ocelli absent, the unarmed legs, and tibial sulcus difficult to see might suggest a relationship with Grosshygioides Brailovsky (1993). In Bau- mannhygia, n. gen., known only from Philippine Republic, the antenniferous tu- bercles are armed, the head slightly convex dorsally, mandibular plate unarmed, scutellar disc and hemelytra weakly punctate, hemelytra micropterous, and the pronotal disc flat. In Grosshygioides, described from Australia, the antenniferous tubercles are unarmed, the dorsal head flat, mandibular plate armed, scutellar disc and hemelytra strongly punctate, hemelytra staphylinoid, and the pronotal disc never flat. Grosshygia Brailovsky (1993) like Baumannhygia, has the antenniferous tu- bercle armed, the mandibular plate unarmed, the eyes small and globular, post- ocular tubercle protuberant, pronotal disc nondeclivent, legs unarmed, tibiae with sulcus difficult to see, hemelytra micropterous, and the membrane absent. The two genera can be separated by the following combination of characters. In Bau- mannhygia, the head in dorsal view is slightly convex, buccula with small and blunt anterior projection, and callar region, including the pronotal and scutellar disc, flat. In Grosshygia, known from Australia, the head in dorsal view is con- spicuously convex and raised above eyes, buccula rounded without teeth, callar region nodulose, and scutellar disc raised and convexly rounded. Description. — Male. Head: quadrangular, wider than long, and dorsally slightly convex; tylus un- armed, apically globose, extending anteriorly to the jugae, and more raised in lateral view; jugae unarmed, thickened, and shorter than tylus; antenniferous tubercle armed with long robust spine; side of head in front of eyes unarmed and apically divergent; antennal segment I robust, thickest, slightly curved outward and shorter than head; segments II and III cylindrical and slender; segment IV fusi- form; segment II longest; segment I longer than III; segment III subequal to IV; ocelli absent; preocellar pit deep; eyes small, globular; postocular tubercle protuberant; buccula rounded, short, not projecting beyond antenniferous tubercle, with small and blunt anterior projection; rostrum long, reaching pos- terior third of abdominal sternite V; mandibular plate absent. Thorax: pronotum trapeziform, wider than long, nondeclivent; collar wide; frontal angles produced forward as rounded and globosus lobes; humeral angles rounded, not expanded; anterolateral borders obliquely straight and raised above pronotal disc; posterolateral borders straight, and posterior border slightly concave; pronotal disc flat. Anterior lobe of metathoracic peritreme reniform, posterior lobe sharp, small. Legs: femora unarmed; tibiae with a vague longitudinal sulcus. Scutellum: wider than long, triangular, with sharp apex; disc flat. 1996 Brailovsky — -New Colpurini 389 Fig. 1 .-—Eludohygia parvioculata Brailovsky, male. 390 Annals of Carnegie Museum VOL. 65 Hemelytra: micropterous reaching posterior third of abdominal segment II; wings reduced to small pads, separated from each other; clavus and corium fused; membrane absent. Abdomen: connexival segments strongly elevated, higher than abdominal segments; angle of con- nexival complete; abdominal sternites with the medial furrow extending to middle third of stemite V. Integument: mostly shining, with hemelytra, connexival segments, and dorsal abdominal segments dull. Head, thorax, abdomen, and exposed parts of genital segments strongly punctate; pronotum, scutellum, hemelytra, and connexival segments weakly punctate. Body with short decumbent silvery bristle-like hairs, intermixed with long erect setae located on the abdominal sternites, legs, and antennal segments. Male genitalia: genital capsule: postero ventral edge with two large and robust lobes, with deep V-shaped notch (Fig. 5). Female. Unknown. Distribution. — Known only from the Philippine Republic. Etymology. — I am please to name this new genus for Dr. Richard W. Baumann, distinguished American plecopterist. Gender feminine. Type Species.— Baumannhygia insulata, n. sp. Baumannhygia insulata, new species (Fig. 5, 8) Description. — Male. Dorsal coloration: head bright black; antennal segment I black, II and III dark red, and IV yellow-ocher with basal joint dark red; pronotum, scutellum, and hemelytra bright black 1996 Brailovsky- — New Colpurmi 391 with dark red reflections; coenexival segments dull black with dark red reflections, except the posterior margin, and dorsal abdominal segments dark red. Ventral coloration: bright black with following areas bright brownish red: acetabuiae, metapleura, abdominal stemite III to VII, and genital capsule; rostral segments I to IV bright orange-yellow; coxae bright red-brown; trochanters bright orange-yellow; femora dark red with basal joint bright orange-yellow; tibiae yellow with basal and apical joint dark red; tarsi pale chestnut-orange; anterior and posterior lobe of metathoracic peritreme dirty orange- yellow; pleural margin of abdominal stemite III to VII dark red with posterior border ocher-yellow. Measurements.— Mdilc (mm). Head length 1.78; width across eyes 1,96; interocular space 1.30; interocellar space 0.76; preocuiar distance 1.32; length antennal segments: I, 1.48; II, 204; III, 1.42; IV, 1.42. Pronotum: total length 1.48; width across frontal angle 1.80 mm; width across humeral angle 2.64. Scutellar length 0.92; width 1.08. Total body length 9.50. Type Specimens.— hlolotype, male (FMNH): PHILIPPINE REPUBLIC, DA- VAO PROVINCE, Mindanao (Calian), 31 December (without year), C. S. Clagg. Paratype, one male (UNAM): PHILIPPINE REPUBLIC, DAVAO PROVINCE, Mindanao (Calian), 11 July (without year), C. S. Clagg. Etymology. — This species is named for its island distribution. Acknowledgments I thank the following colleagues and institutions for the loan of specimens and other assistance relevant to this study: Gordon M. Nishida (Bernice P. Bishop Museum, Honolulu, Hawaii); Janet Margerison (The Natural History Museum, London, United Kingdom); P. P. Parrillo (Field Museum of Natural History, Chicago, Illinois); G. B. Monteith (Queensland Museum, Brisbane, Australia). Ernesto Barrera and Cristina Urbina from Institute de Biologia, UNAM, prepared the illustrations. Special thanks to the Consejo Nacioeal de Ciencia y Technologia (CONACyT) for financial assistance. Comments on the manuscript from the anonymous reviewers are greatly appreciated. Literature Cited Brailovsky, H. 1990. Three new species of Indo-Pacific Colpurini (Hemiptera: Heteroptera: Corei- dae). Pan-Pacific Entomologist, 66:292-=300. ■ — 1993g. New genera and new species of Colpurini (Heteroptera: Coreidae) from the Fiji Islands and New Guinea). Proceedings of the Entomological Society of Washington, 95:435-448. 19931?. New genera and new species of micropterous Colpurini from Buiii Islands and New Guinea (Heteroptera: Coreidae). Pan-Pacific Entomologist, 69:281-289. — — — , 1993c‘. A revision of the tribe Colpurini from Australia (Hemiptera-Heteroptera-Coreidae). Memoirs of the Queensland Museum, 34:35-60. Brailovsky, H., and G. Ortega Leon. 1994. Three new genera and six new species of Colpurini (Heteroptera: Coreidae) from New Guinea. Proceedings of the Entomological Society of Wash- ington, 95:500-515. Brailovsky, H., E. Barrera, and W. Lopez-Forment. 1992. Revision of the genus Tackycolpura Breddin (Hemiptera: Heteroptera: Coreidae: Colpurini). Pan-Pacific Entomologist, 68:122-132. V- ■- «. ^ -•„'•.■ "il^ ■i-."2:-^*^-.'ni il^ ^ ^ ^1. -'riv faiiin|i * ■!•, M? n»^-. t^ ..i . < •J^|'«'^ iS* ' ■ ,. -wit-, il^', ' <- ' •■ .’M i' rtji; • ■'■■'1‘’'f ■”3' ^M — nil-' .-’ - t. . .^ ’■ ■ • '"' "TVt ■'^^- f ■ ---^>> ' I ‘ i^ ' ' ^'' '• ■' ■M j,W^" ' * 1 i '■ ,. ' i ■t''‘•^r^^.^& .‘f^vJ ...:^ ' ^ V ^ ' ■.^' * } ' ■ '*<--' * * « I i '’ ■ ‘j'.. ' ' ■' ■ ■ ' / ^ 'i*-'*' ‘'‘^s-^ I, ja>>«!!f '’-ji I - -■’ «. * ' ' ■ „ ■'■'■ '■ ■■ "d^'^OLa:^ .. .|^'4 r ,. v n f * f., ■, . .n ■'- 4 : . * .JAV ■» -/ , i‘iri3u'^s- i-. ...#9>r'*i ’- » '’ih.v.7;tJ4'-4«W!P5^ **' .t.« '. ”- .’t/'^(MA--Jl...' v^k.-'ulpAnif- a, ;■ "1 il - . .>.- i.:.. J .. 'v*«H ; ‘^4’ J^ ' ■ ANNALS OF CARNEGIE MUSEUM VoL. 65, Number 4, Pp. 393=399 27 November 1996 ON THE GENUS ACANTHOLYBAS BREDDIN (INSECTA: HETEROPTERA: COREIDAE: COLPURINI), WITH DESCRIPTION OF ONE NEW SPECIES FROM JAVA Harry Brailovsky^ Research Associate, Section of Invertebrate Zoology Abstract Acantholybas steinbaueri a brachypterous new species, is described and illustrated from Java (Bali Island and Dammerman). The genus Acantholybas is redescribed, and a key to the four known species is provided. Introduction The genus Acantholybas was proposed by Breddin ( 1 899) to include the species A. longulm collected from Lombok Island and later recorded from Sumbawa (Blote, 1936). Subsequently Breddin (19001?) described the Acanthocolpura with one species, A. brunneus, from New South Wales, Australia, and in the same year (Breddin, 1900a) ^ynonymimd Acanthocolpura with Acantholybas and trans- ferred A. brunneus to Acantholybas. Bergroth (1909) described A. kirkaldyi from Tasmania (Semmens et aL, 1992). Acantholybas belongs to the group of Colpuriei with the abdominal stemite VII of the female complete, without plica or fissiira, and has not been previously reviewed or revised. There are no keys or modem descriptions for any of the species except A. brunneus (Woodward, 1951, 1953, 1961; Wise, 1958a, 19581?; Brailovsky, 1993). An examination of material representing Acantholybas spp. indicates that the critical differences between species are found in the development of the callar region, tylus, hemelytra, proportions of the antennal segments, and relative body size. This paper presents a summary of these comparisons. The following abbreviations identify the institutions where types are deposited: BPBM, Bernice P. Bishop Museum, Honolulu, Hawaii; UN AM, Coleccion En- tomologica del Instituto de Biologfa, Universidad Nacionai Autonoma de Mexico; ZIL, Zoological Institute, St. Petersburg, Russia. Systematic Entomology Acantholybas Breddin (Fig. 1--10) Acantholybas Breddin, 1899: 169=1 70. Diagnosis.— Acantholybas Breddin is distinguished by the globose, apically truncated or bifid tylus, the armed antenniferous tubercle, the armed buccula with an obvious spine near the middle third, the unarmed femora and the abdominal stemite VII of female without plica or fissura. Pachycolpuroides Brailovsky ' Institute de Biologfa, UNAM, Depto. de Zoologfa, Apdo. Postal 70153, Mexico 04510 D. F. Mexico. Submitted 31 March 1995. 393 394 Annals of Carnegie Museum VOL. 65 (1993), the most closely related genus, may be recognized by its smaller size, rounded buccula without teeth or spiny projection, and shorter rostrum not ex- tending beyond the anterior margin of the metastemum. Redescription. — Head: pentagonal, wider than long (across eyes) and dorsally slightly convex; tylus unarmed, apically globose or bifid, extending anteriorly to the jugae and more raised in lateral view; jugae unarmed, thickened, and shorter than tylus; antenniferous tubercle armed, lobes raised, diverging anteriorly, and apically subacute; side of head in front of eye short, unarmed, and obliquely straight; antennal segment I robust, thickest or more or less slender, slightly curved outward and shorter than head; segments II and III cylindrical and slender; segment IV fusiform; segment II longest; segment I shorter; segment IV longer or shorter than III (Fig. 2-4); ocelli tuberculate, well developed; preocellar pit deep; eyes large, hemispherical, sessile; postocular tubercle protuberant; buccula rounded, short, not projecting beyond antenniferous tubercle, with sharp spiny projection; rostrum reaching posterior third of abdominal sternite IV or anterior third of V; mandibular plate absent. Thorax: pronotum wider than long, nearly trapeziform, slightly bilobed, and nondeclivent; anterior lobe shorter than posterior lobe, both with lateral margins convexly rounded, not elevated, and slightly reflexed; collar wide; frontal angles produced forward as conical lobes; humeral angles rounded; posterolateral border straight and posterior border slightly concave; callar region transversely convex with or without median longitudinal depression. Anterior lobe of metathoracic peritreme reniform, posterior lobe sharp, small. Legs: femora unarmed; tibiae sulcate. Scutellum: longer than wide, wider than long, or equilateral; triangular, with the apex rounded or subacute and raised or not above disc; disc nearly flat. Hemelytra: macropterous forms with hemelytral membrane well developed, extending to the apex of abdomen or beyond. Brachypterous forms with clavus and corium distinctly separate; hemelytral membrane reduced, reaching the third abdominal segment; wings widely separated from each other, leaving the abdomen exposed mesally. Abdomen: connexival segments higher than abdominal segments, with posterior angle not produced into spines; abdominal sternites with medial furrow extending to anterior third of sternite V. Integument: body surface mostly dull, with head, thorax, and abdomen slightly shining. Pronotum, scutellum, clavus, corium, dorsal abdominal segments, ventral surface of head, thorax, abdominal sterna, and genital capsule strongly punctate. Dorsal surface of head with scattered punctures and connexival segments practically smooth. Body with short, decumbent, silvery bristle-like setae, inter- mixed with long erect setae located on the abdominal sterna, legs, and antennal segments. Male genitalia: genital capsule: posteroventral border broadly rounded, with a large internal emar- gination (Fig. 5). Parameres: body robust, with anterior lobe convex, and posterior lobe long (Fig. 6- 7, variation for the different view). Female genitalia: abdominal sternite VII complete, without plica or fissura. Genital plates: gonocoxa I oblique, with a convex and protruding external margin, slightly emarginate; paratergite VIII short, square, with visible spiracle; paratergite IX squarish, larger than paratergite VIII (Fig. 8-9). Distribution. — Acantholybas species have been collected from Lombok Island (Indonesia), Australia, Tasmania, New Zealand, and now recorded from Java. Type Species. — Acantholybas long ulus Breddin. Acantholybas steinbaueri, new species (Fig. 1-2, 5, 8-9) Description. — Color, male. Head: anterior lobe of the pronotum, scutellum, connexival segments, dorsal abdominal segments, thorax, and abdominal sterna bright to dark red-brown with dark orange reflections; apex of scutellum and anterior and posterior lobe of metathoracic peritreme yellowish white to pale yellow; antennal segments I-III dark orange-brown with reddish brown reflections; segment IV pale yellow with proximal end dark orange-brown; posterior lobe of pronotum, clavus, and corium dark to pale orange-yellow; rostral segments dark orange-brown with basal segment paler; superior area of the postocular tubercle, posterior edge of connexival segments IV-VI, and posterior Fig. 1. — Acantholybas steinbaueri Brailovsky, new genus, new species; female. 1996 Brailovsky— A cA#raoLraA5' Breddin 395 396 Annals of Carnegie Museum VOL. 65 2 3 1996 Brailovsky — Acantholybas Breddin 397 Fig. 2-9. — Antennae and genitalia of Acantholybas. 2, antennal segments of A. steinbaueri Brailovsky; 3, antennal segments of A, kirkaldyi Bergroth; 4, antennal segments of A. brunneus (Breddin); 5, male genital capsule of A. steinbaueri Brailovsky; 6-7, parameres of A. brunneus (Breddin); 8-9, female genital plates of A. steinbaueri Brailovsky; 8, caudal view; 9, lateral view. 398 Annals of Carnegie Museum VOL. 65 edge of pleural sterna IV-VI yellow; coxae dark reddish brown with orange reflections; trochanters dark orange-brown with dorsal face pale yellow; anterior and middle femora dark orange-brown; posterior femora dark orange-brown with a wide yellow ring located on the middle third; tibiae dark orange-brown; tarsi with dorsal surface dark orange-brown, and ventral surface pale yellow. Color, female. Similar to male. Structure. Head; tylus apically globose; antennal segment III shorter than IV. Pronotum: callar region transversely convex, with or without a vague, difficult-to-see, median longitudinal depression. Scutel- lum: wider than long (male) and longer than wide (female). Hemelytra: brachypterous, with the mem- brane reaching the third abdominal segment. Measurements. — Male, female (mm). Head length 1.32, 1.46; width across eyes 1.48, 1.60; inter- ocular space 0.90, 0.96; interocellar space 0.46, 0.51; preocular distance 0.86, 0.94; length antennal segments: I, 0.88, 1.04; II, 1.44, 1.56; III, 1.00, 1.08; IV, 1.08, 1.16. Pronotum: total length 1.52, 1.84; width across frontal angles 1.56, 1.48; width across humeral angles 2.40, 2.56. Scutellar length 1.04, 1.32; width 1.08, 1.24. Total body length 8.40, 9.90. Type Specimens. — Holotype, male (BPBM): JAVA, Bali L, Dadjan Danu, 29 March 1956, J. Winkler. Paratypes: one female (UN AM), data as for holotype; one male (ZIL), Dammerman, Idjen Kendeng, 1400 m, 15 June 1924 (no. 68-III). Etymology. — Named for Martin James Steinbauer in recognition of his recent rediscovery of Acan- tholybas kirkaldyi Bergroth. Key to the Known Species of Acantholybas 1 Brachypterous, with the wings widely separated from each other, leaving the abdomen exposed mesally; hemelytral membrane reduced, reaching the third abdominal segment (Fig. 1) (Java) A. steinbaueri, new species T Macropterous; hemelytral membrane well developed, reaching the apex of the abdomen (Fig. 10) 2 2 Antennal segment I longer than 1.00 mm; antennal segment II longer than 1.90 mm. (Lombok Island) A. longulus Breddin 2' Antennal segment I shorter than 0.90 mm; antennal segment II shorter than 1.60 mm. (Fig. 3, 4) ............................. ..... .............. 3 3 Antennal segment I robust; body robust and longer than 10.20 mm; scutellum wider than long or equilateral (Tasmania) A. kirkaldy Bergroth 3' Antennal segment I slender; body slender and shorter than 10.00 mm; scutellum longer than wide (Fig. 10) (Australia and New Zealand) .A. brunneus (Breddin) Acknowledgments I am grateful to the following individuals and institutions for the loan of specimens and other assistance: Gordon M. Nishida (BPBM); Tamas Vasarhelyi (Hungarian Natural History Museum); G. B. Monteith (Queensland Museum, Brisbane, Australia); Margaret Schneider (University of Queens- land, Brisbane, Australia); I. M. Kerzhner (ZIL). Ernesto Barrera, Albino Luna, and Cristina Urbina (UNAM) prepared the illustrations. Special thanks to the Consejo Nacional de Ciencia y Tecnologia (CONACyT) for financial assistance. Comments on the manuscript from the anonymous reviewers are greatly appreciated. Literature Cited Bergroth, E. 1909. Hemiptera Nova Orientalia. Annales de la Societe Entomologique de Belgique, 53:184-190. Blote, H. C. 1936. Catalogue of the Coreidae in the Rijksmuseum van Natuurlijke Historie. Part III. Coreinae, Second part. Zoologische Mededeelingen, 19:23-66. Brailovsky, H. 1993. A revision of the tribe Colpurini from Australia (Hemiptera-Heteroptera- Coreidae). Memoirs of the Queensland Museum, 34:35-60. Breddin, G. 1899. Hemiptera Insulae Lombok in Museo Hamburgensi asservata adiectis speciebus nonnullis, quas continet collectio auctoris. Jahrbuch der Hamburgischen Wissenschaftlichen An- stalten, 16:155-194. 1996 Brailovsky — Acantholybas Breddin 399 1900a. Materiae ad cognitionem subfamiliae Pachycephalini (Lybantini Olim) ex Hemip- teris-Heteropteris, Fam. Coreidae. Revue d’Entomologie, Caen, 19:194-217. 1900^. Hemiptera nonnulla regionis Australicae (figuris duodecim illustrata). Entomologische Nachrichten (Berlin), 26:33-46. Semmens, T. D., P. B. McQuillan, and G. Hayhurst. 1992. Catalogue of the Insects of Tasmania. Department of Primary Industry, Hobart, Tasmania. Wise, K. A. J. 1958a. Notes on insects pests, 1956-57. New Zealand Entomologist, 2:26-28. 1958^. Notes on insects pests, 1957-58. New Zealand Entomologist, 2:12-14. Woodward, T. E. 1951. The occurrence of Acantholybas brunneus Breddin in New Zealand (Het- eroptera: Coreidae). Transactions of the Royal Society of New Zealand, 79:206-209. 1953. Notes on Acantholybas brunneus Breddin (Heteroptera; Coreidae). New Zealand En- tomologist, 1:8-9. “ — — — . 1961. The Heteroptera of New Zealand. Part III. Coreidae, Berytidae, Tingidae, Cimicidae. Transactions of the Royal Society of New Zealand, 1:145-158. f VH jSXV. r r^i- -. f --. - ^ ii ,11. '.' .r j * , f ^ * <1 ' ■' ' )'♦ ’l» >^us * C4vfc . _^ . * - . 1 -~ X- ' ' ^7 >4 ’.'ir , ^r‘*’f‘^it 'V mniCJ ’ /. vt ’ * i,u^ ti ^ . . . A . , INDEX TO VOLUME 65 CONTENTS ARTICLES Biology and immature stages of snail-killing flies belonging to the genus Tetanocem (Insecta: Diptera: Sciomyzidae). I. Introduction and life histories of predators of shoreline snails V ....... B. A, Foote New species of Rhyacophila from Virginia (Insecta: Trichoptera: Rhyacop'Mlidae) ......................................... Jan L. Sykora and Decian J. McCabe New species of microcaddisflies from the eastern United States (Insecta: Trichoptera: Hydrop- tilidae) .................................. Steven C. Harris and Jan L. Sykora A phylogenetic reassessment of the tadomine-anatine divergence (Aves: Anseriformes: Anati- dae) ................................................ .Bradley C. Livezey Observations on the mammals of Tucuman Province, Argentina. L Systematics, distribution, and ecology of the Didelphimorphia, Xenarthra, Chiroptera, Primates, Carnivora, Perissodac- tyla, Artiodactyla, and Lagomorpha ........... Michael A. Mares, Ruben M. Barquez, Janet K. Braun, and Ricardo A. Ojeda Biology and immature stages of snail-killing flies belonging to the genus Tetanocem (Insecta: Diptera: Sciomyzidae). IL Life histories of predators of snails of 'the family Succineidae ........................................................ B. A. Foote A new genus of beaver (Mammalia: Castoridae: Rodentia) from the Arikareean (Oligocene) of Montana and its bearing on castorid phytogeny ................... William W. Korth The taxa of fossil Mollusca introduced by Hermann von Ihering Juan Jose Parodiz Two new species of Xyris from Cuba .......... Armando Urquioia Craz and 'Robert Krai A new piesiadapiform (Mammalia: Plesiadapiformes) from the early Eocene of the Bighorn Basin, Wyoming ......................... Kenneth D. Rose and Thomas M. Bown Archaeology of Trants, Montserrat. Part 3. Chronological and settlement data ..................................................... .James B. Petersen Pipunculidae (Insecta: Diptera) of the Dominican Republic: New records aiid descri'ptioo of new species ................................................ J. A. Rafael New genera and new species of Colpurini (Ins^ta: Heteroptera: Coreidae) from Philippines, Borneo, and Malaya ...................................... Harry Brailovsky On the genus Acantholybas Breddin (Insecta: Heteroptera: Coreidae: Colpurini), with descrip- tion of one new species from Java ........................... Harry Brailovsky 1 13 17 27 89 153 167 183 297 305 323 363 383 393 401 402 Annals of Carnegie Museum VOL. 65 REVIEW Australian Beetles (J. F. Lawrence and E. B. Britton) ............... .Darren A. Pollock 181 NEW TAXA NEW FAMILIES, GENERA, AND SPECIES Acantholybas steinbaueri, new species ....................................... 394 Baumannhygia, new genus ............................................... 388 Baumannhygia imulata, new species ........................................ 390 Cephalops (Cephalops) pedemalemis Rafael, new species ......................... 368 Cephalosphaera (Cephalosphaera) insularis Rafael, new species .................... 367 Eludohygia, new genus .................................................. 387 Eiudohygia parvioculata, new species ....................................... 387 Eudorylas curvicaudatus Rafael, new species .................................. 377 Eudorylas dominicanensis Rafael, new species ................................. 377 Hydroptila fowlesi, new species ............................................ 19 Hydroptiia homochitta, new species ......................................... 21 Hydroptila koryaki, new species ........................................... 17 Kinabaluhygia, new genus ............................................... 383 Kinabaluhygia sabah, new species .......................................... 386 Mayatrichia tuscaloosa, new species ........................................ 23 Metadorylas antillensis Rafael, new species ................................... 374 Metadorylas comutus Rafael, new species .................................... 374 Metadorylas pilosus Rafael, new species ..................................... 376 Metadorylas youngi Rafael, new species ...................................... 372 '\Neatocastor, new genus ................................................ 167 tPicromomyidae, new family ............................................. 306 ^Picromomys, new genus ................................................ 307 -\Picromomys petersonorum, new species ..................................... 308 Rhyacophila tricomuta, new species ........................................ 13 Xyris bissei Urquiola & Krai, species novum .................................. 297 Xyris mantuensis Urquiola & Krai, species novum .............................. 300 t Fossil taxa AUTHOR INDEX Barquez, Ruben M. ............................... Bown, Thomas M. ................................ Brailovsky, Harry ................................. Braun, Janet K. .................................. Cruz, Armando Urquiola ............................ 89 . . 305 383, 393 89 . . 297 1996 403 Foote, B. A. .................. 1, 153 Harris, Steven C. 17 Korth, William W. 167 Krai, Robert .................... ............................. 297 Livezey, Bradley C 27 Mares, Michael A. ..................................................... 89 McCabe, Declan J. 13 Ojeda, Ricardo A. ..................................................... 89 Parodiz, Juan Jose 183 Petersen, James B. 323 Pollock, Darren A. 181 Rafael, J. A. 363 Rose, Kenneth D 305 Sykora, Jan L. 13, 17 INSTRUCTIONS FOR AUTHORS ANNALS OF CARNEGIE MUSEUM consist of contributions to the earth sciences (including paleontol- ogy), life sciences, and anthropology, in 30 by 46 picas format (127 by 195 mm or 5 by 7% inches). Submit all manuscripts to the Office of Scientific Publications. 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